(Demetrio Boltovskoy) South Atlantic Zooplankton

South Atlantic
Zoo~lankton
Volume 1
Edited by
Demetrio Boltovskoy
Backhuys Publishers
South Atlantic
Zooplankton
Vol. 1
Pages 1 - 868
Edited by Demetrio Boltovskoy
[B
Backhuys Publishers, Leiden 1999
South Atlantic
Zooplankton
Vol. 2
Pages 869 - 1706
Edited by Demetrio Boltovskoy
lB
Backhuys Publishers, Leiden 1999
ISBN 90-5782-035-8
Volume 1: pages 1 - 868

Volume 2: pages 869 - 1706
© Backhuys Publishers, Leiden, The Netherlands 1999
AIl rights reserved. Nothing from this publication may be reproduced, stored in a computerized system or published in any form
or in any matter, including electronic, mechanical, reprographic or photographic, without prior written permission from the pub-
lishers, Backhuys Publishers, P.O. Box 321, 2300 AH Leiden, The Netherlands.
Printed in The Netherlands

Contents
Foreword vii
Authors ix
General biological features of the South Atlantic - Demetrio Boltovskoy, Mark Gibbons, Lawrence
Hutchings and Denis Binet 1
Foraminifera - Sylvia Kemle-von Mücke and Christoph Hemleben 43
Acantharia - Renate Bernstein, Stanley A. Kling and Demetrio Boltovskoy 75
Radiolaria Polycystina - Demetrio Boltovskoy 149
Radiolaria Phaeodaria - Stanley A. Kling and Demetrio Boltovskoy 213
Ciliophora - Wo/fgang Petz : 265
Tintinnoinea - Viviana A. Alder 321
Hydromedusae - Jean Bouillon 385
Siphonophorae - Phil Pugh 467
Cubomedusae and Scyphomedusae - Hermes W Mianzan and Paul Cornelius 513
Ctenophora - Hermes W Mianzan 561
Nemertina - Ray Gibson 575
Polychaeta - Erik Thuesen and María Ana Fernández Alamo 595
Heteropoda - Gotthard Richter and Roger R. Seapy 621
Pteropoda - Siebrecht van der Spoel and José R. Dadon 649
Cephalopoda - Kir N. Nesis 707
Ctenopoda and Onychopoda (=Cladocera) - Takashi Onbé 797
Ostracoda - Martin V Angel.. 815
Copepoda -Janet M. Bradford-Grieve, Elena L. Markhaseva, Carlos E.F. Rocha and Bernardo Abiahy 869
Mysidacea - Masaaki Murano 1099
Amphipoda - Georgyi Vinogradov : 1141
Euphausiacea - Mark J Gibbons, Vassily A. Spiridonov and Geraint A. Tarling 1241
Larval Decapoda (Brachyura) - Gerhard Pohle, Adilson Fransozo, María L. Negreiros-Fransozo and
Fernando L. Medina Mantelatto 1281
Chaetognatha - Jean-Paul Casanova 1353
Appendicularia - Graciela B. Esnal 1375
Pyrosomatida - Graciela B. Esnal 1401
Doliolida - Graciela B. Esnal and María C. Daponte 1409
Salpida - Graciela B. Esnal and María C. Daponte 1423
Fish larvae - Yasunobu Matsuura and María Pilar Olivar 1445
Bibliography 1497
Taxonomic index 1627
Foreword
Most biological oceanographers working on zoo- liographic references have been included. The most
planktonic organisms are familiar with the systematics relevant works dealing with the classification and dis-
and distribution of one group of animals, have a vague tribution of each taxon have been listed separately, in
understanding of a couple more, and usually know the "Suggested readings" sections provided at the end
very little about the rest. Our time is often barely suf- of each chapter.
ficient to keep abreast with the literature on our main
subject of interest, let alone dwelling into publications Species-specific distributional data for the South
centered on other groups. Furthermore, because of the Atlantic constitute the other major target ofthis work.
increasing degree of specialization, literature on these These, aside from furnishing an additional evidence
other groups is often either unavailable or ignored. for identification purposes, update our current know-
One of the main purposes of the present volume is to ledge on the biogeography of the zooplankton of this
fill this gap offering an updated, comprehensive com- vast area. In most cases distributional data were sum-
pendium where all marine zooplanktonic groups are marized in tables or maps in order to make them read- .
included. The book is chiefly aimed at surnmarizing ily accessible to the reader. Whenever possible, scat-
available data on the c1assification and geographic dis- tered records were interpreted in terms of the
tribution of 28 higher-Ievel taxa in the South Atlantic ecological settings derived and general ranges are
Ocean. General introductory information on morpho- suggested. In a few instances, however, when records
logy, reproduction, development, feeding, production are too scarce and spotty, only the isolated findings
rates, etc. is al so included, and the references provid- are given.
ed should be helpful in guiding the student toward the
relevant papers on each subject. It should be stressed that, for most chapters, the taxa
covered include not only those effectively reported to
In a few cases the chapters presented benefited from date from South Atlantic waters, but also the ones
recently published comprehensive reports for the whose known ranges suggest that their presence \TI
World Ocean and/or for restricted oceanic areas. In this ocean is very likely.
most others, however, overviews were painstakingly
put together on the basis of widely scattered informa- In short, the book provides a thoroughly documented
tion, including turn of the century expedition reports, review of what zooplankton species are present in the
a very substantial amount of papers of restricted dis- South Atlantic from the equator to 60oS, from the
tribution (cruise, internal institutional and grant coasts of South America to Africa, and what are their
reports, dissertations, etc.), unpublished data, and geographic and vertical ranges. In this respect, this is
much personal, critical input. Thus, because all chap- probably one of the first such attempts for a vast
ters were prepared by students of international stand- oceanic basin, one that is certainly in need of much
ing with ample experience in their respective fields, further research. General data on abundance and bio-
the information presented is by no means a mere com- mass, seasonality, sinking and relationships with the
pilation of previous literature, but represents highly sedimentary distribution of the corresponding fossi-
critical overviews. In some cases this resulted in lizable remains, horizontal and vertical diversity pat-
extensive taxonomic rearrangements, and in most terns, associations with currents and water masses,
changes with respect to previous data are suggested. ecophenotypic variations, etc. complement the di s-
tributional information provided.
As an identification tool, the information included in
this volume should suffice for general purpose speci- As much as possible I have tried to bring all chapters
fic, or at least generic or family-Ievel identifications. to a cornmon format. Differences, however, persist;
Detailed systematic studies will obviously require they reflect dissimilarities in the amount and com-
additional literature, for which reason abundant bib- plexity of information available for the various groups
Foreword V111
covered, as well as differences in style, approach and The assistance and permanent advice of Stanley Kling
interests of the 41 authors involved. throughout many months of hard work was a major
support and is very sincerely appreciated. Eleonora
Many friends and colleagues have helped to bring this Uliana and Michiko Matsubara patiently proofread
volume to fruition in one way or another. I would first thousands of manuscript pages and assembled hun-
like to thank all the authors for generously investing dreds of illustrations.
much of their valuable time in this endeavor, and espe-
cially for patiently bearing with my editorial demands. Editorial aspects of this work were partly financed by
Many coauthors, as well as other colleagues, were grants from the University ofBuenos Aires (UBA EX-
extremely helpful in providing comparative specimens 040), from the Consejo Nacional de Investigaciones
and published and unpublished data to others, raising Científicas y Técnicas, Argentina (CONICET PID-BID
the funds for covering travel expenses in order to joint- 0366), and from the Agencia Nacional de Promoción
Iy complete their sections, locating hard to get copy- Científica y Tecnológica, Argentina (PMT -PICT0395).
right clearances, allowing to use their unpublished or
copyrighted materials, etc. Among these, as editor I My contribution to this work is dedicated to my
am particularly grateful to Carina Lange, Janet father, Esteban Boltovskoy, whose guidance and sup-
Bradford-Grieve, Jean-Paul Casanova, Jordi Corbera, port over the years were a major influence in my life.
Paul Comelius, Mark Gibbons, Ray Gibson, Lena
Markhaseva, Yasunobu Matsuura, Hermes W.
Mianzan, Masaaki Murano, Makoto Omori, María Demetrio Boltovskoy
Pilar Olivar, Charrid Resgalla, Carlo~ E. F. Rocha and
Georgy Vinogradov.
Authors
Bernardo Abiahy Demetrio Boltovskoy

Universidade de Sao Paulo, Departamento de Ciencias Biológicas,
Departamento de Zoologia, Facultad de Ciencias Exactas y
Caixa Postal 11461, Naturales, Universidad de Buenos
CEP 05422 970 Sao Paulo, Brazil Aires, 1428 Buenos Aires;
(barroso@usp.br) Museo Argentino de Ciencias
Naturales "Bernardino Rivadavia";
Viviana A. Alder and Consejo Nacional de
Instituto Antártico Argentino, Investigaciones Científicas y
Consejo Nacional de Investigaciones Técnicas, Argentina
Científicas y Técnicas, and (demetrio@bg. fcen. uba.ar)
Departamento de Ciencias Biológicas,
Facultad de Ciencias Exactas y Jean Bouillon
Naturales, Universidad de Buenos Laboratoire de Zoologie,
Aires, 1428 Buenos Aires, Argentina Université Libre de Bruxelles,
(viviana@bg. fcen. uba.ar) Av. F.D. Roosevelt 50, Bruxelles,
B-I050, Belgium
Martín V. Angel Postal address: Rue Neuve 20,
Southampton Oceanography Centre, Oignies 5670, Belgium
. Empress Dock, (jbouillo@ulb.ac.be)
Southampton S014 3ZH, UK
(mva@soc.soton.ac.uk) Janet Bradford-Grieve
National Institute of Water and
Atmospheric Research, Box 14901,
Renate Bernstein Kilbirnie, Wellington, New Zealand
University of South Florida, (grieve@storm.greta.cri.llZ)
Department of Marine Science,
St. Petersburg Campus,
140 7th Ave. South, Jean-Paul Casanova
St. Petersburg, Florida 33701, USA Université de Provence,
(renate@seas. marine. usf. edu) Biologie Animale (Plancton),
case 18,3, Place Victor Hugo,
Denis Binet 13331 Marseille, Cedex 3, France
Antenne Orstom - Centre (bioplank@newsup.univ-rnrs.fr)
IFREMER, B.P. 21105-44311,
Nantes-Cedex, France Paul Cornelius
(binet@orstom.fr) Department of Zoology,
The Natural History Museum,
London SW7 5BD, UK
(pfsc@nhm.ac.uk)
".
x Authors
José R. Dadon Ray Gibson

Departamento de Ciencias Biológicas, School of Biological and Earth
Facultad de Ciencias Exactas y Sciences, Liverpool lohn Moores
Naturales, Universidad de Buenos University, Byrom Street,
Aires, 1428 Buenos Aires, and Liverpool L3 3AF, UK
Consejo Nacional de Investigaciones (r.gibson@livjm.ac.uk)
Científicas y Técnicas, Argentina
(dadon@bg. fcen. uba.ar) Christoph Hemleben I
Institut und Museum fur Geologie und
Cristina Daponte Palaontologie, Universitat Tübingen,
Departamento de Ciencias Biológicas, D-7400 Tübingen, Germany
Facultad de Ciencias Exactas y (christoph.hemleben@uni-tuebingen.de)
Naturales, Universidad de Buenos
Aires, 1428 Buenos Aires, Argentina
(daponte@bg. fcen.uba.ar) Lawrenee Hutehings
Sea Fisheries Research Institute,
Graciela B. Esnal Private Bag X2, Roggebaai 8012,
Departamento de Ciencias Biológicas, Republic of South Africa
Facultad de Ciencias Exactas y (lhutchin@sp.sfri.ac.za)
Naturales, Universidad de Buenos
Aires, 1428 Buenos Aires, and
Consejo Nacional de Investigaciones Sylvia Kemle-von Müeke
Científicas y Técnicas, Argentina Fachbereich Geowissenschaften,
(esnal@bg.fcen. uba.ar) Klagenfurter Strasse,
Universitat Bremen,
María Ana Fernández Álamo 28359 Bremen, Germany
Laboratorio de Invertebrados, (skemle@allgeo. uni -bremen.de)
Facultad de Ciencias, UNAM,
Apartado Postal 70-371, México, Stanley A. Kling
D.F. 04510, México 416 Shore View Lane, Leucadia,
J California 92024, USA
(mafa@hp.fciencias.unam.mx)
(skling@gs.ucsd.edu)
Adilson Fransozo
Departamento de Zoologia,
Instituto de Biociencias, UNESP -
Campus de Botucatu, CP 502, Lena Markhaseva
CEP 18618-000, Botucatu, SP, Brazil Zoological Institute,
(fransozo@surfnet. comobr) Russian Academy of Sciences,
Universitetskaya 1,
Mark Gibbons 199034 St. Petersburg, Russia
Zoology Department, University of (lena@markhaseva.zin.ras.spb.fU)
the Western Cape, Private Bag XI7,
Bellville 7535; Zoology Department, Yasunobu Matsuura
University of Cape Town, Instituto Oceanográfico,
Rondebosch 7700; Sea Fisheries Universidade de S1ioPaulo,
Research Institute, Prívate Bag X2, Cidade Universitária, Butantá,
Roggebaai 8012, 05508 S1ioPaulo, Brazil
Republic of South Africa (ymatsuur@usp.br)
(mark@botany.uwc.ac.za)
Authors Xl
Fernando L. Medina Mantelatto Takashi Onbé

Departamento de Biologia, Faculty of Applied Biological Science
FFCLRP-USP, Av. Bandeirantes 3900, Hiroshima University
Riberao Preto (SP), Brazil Higashi-Hiroshima 739
(flmantel@spider. usp. br) Japan
Postal address:
2-60-406 Saijo Nishi Honmachi,
Hermes W. Mianzan Higashi-Hiroshima 739, Japan
Instituto Nacional de Investigación (Fax: 81-824-22-8208)
y Desarrollo Pesquero (INIDEP),
Casilla de Correo 175, Wolfgang Petz
7600 Mar del Plata, and Consejo Institut fur Zoologie,
Nacional de Investigaciones Universitat Salzburg,
Científicas y Técnicas, Argentina Hellbrunner Strasse 34,
(hermes@mdp.edu.ar) A-5020 Salzburg, Austria
(wolfgang. petz@sbg.ac.at)
Masaaki Murano
Institute of Environmental Ecology, Gerhard Pohle
METOCEAN Co. Ud., Atlantic Reference Centre,
Riemon 1334-5, Ooigawa-cho, Huntsman Manne Science Centre,
Shida-gun, Shizuoka 421-02, Japan Sto Andrews, New Brunswick,
(murano@notes.metocean.co.jp) Canada EOG 2XO
(arc@sta.dfo.ca)
María L. Negreiros-Fransozo
Departame..1o de Zoologia, Phil Pugh
Instituto de Biociencias, UNESP Southampton Oceanography Centre,
Campus de Botucatu, CP 502, Empress Dock,
CEP 18618-000, Botucatu, SP, Brazil Southampton S014 3ZH, UK
(fransozo@surfnet. comobr) (prp@soc.soton.ac.uk)
Kir Nesis
P.P. Shirhnov Institute of Oceanology Gotthard Richter
of the Russian Academy of Sciences, Forschungsinstitut Senckenberg,
36 Nakhimov Avenue, Senckenberganlage 25,
117218 Moscow, Russia 6000 Frankfurt am Main 1,
(npar@fish.comcp .msk. su) Germany
(sbaszio@sng.uni- frankfurt.de)
María Pilar Olivar
Instituto de Ciencias del Mar, Carlos E.F. Rocha
CSIC, Paseo Joan de Borbó s/n, Universidade de Sao Paulo,
Barcelona 08039, Spain Departamento de Zoologia,
(polivar@icm.csic.es) Caixa Postal 11461, CEP 05422 970
Sao Paulo, Brazil
(cefrocha@usp.br)
••
Xli Authors
Roger Seapy Erik Thuesen

Department of Biological Science, Lab. n, The Evergreen State College,
California State University, Fullerton, Olympia, Washington 98505, USA
r California 92634, USA (thuesene@el wha. evergreen .edu)
(rseapy@fullerton.edu)
Vassily Spiridonov Siebrecht van der Spoel

Zoology Museurn of the University of Zoologisch Museum,
Moscow, Bolshaya Nikitnskaya, 6, Universiteit van Amsterdam,
Moscow 103009, Russia Postbus 94766,
(spiridonov@5 .zoomus.bio.msu.ru) 1090 GT Amsterdam,
The Netherlands
(spoel@bio. uva. nI)
Geraint Tarling
Scottish Marine Biological Association, Georgy Vinogradov
P.O. Box 3, Oban, Argyll, A.N. Severtzov Institute ofthe
Scotland PA34 4AD, UK Problems of Evolution,
(gant@wpo.nerc.ac.uk) Russian Academy of Sciences (RAS),
Lenin Ave. 33, Moscow 117071,
Russia
(egor@ecosys.sio.rssi.ru or
egor@invert.sevin.msk.ru)
General biological features of the South Atlantic
Demetrio Boltovskoy, Mark J. Gibbons, Lawrence Hutchings and Denis Binet
Introduction now been more or less well established for over 50

years, the lack of major modifications to the biologi-
The goal of this section is offering an overview of cal zonations defined half a century ago is not surpri-
some salient biological traits of the South Atlantic. The sing. However, due to the tight physical-biological
scheme outlined below reflects the known or assumed coupling, increasing knowledge about the physical
boundaries and gradients between discrete, structurally characteristics of the oceans paves the way for more
more or less homogeneous faunal domains, yet traits detailed biogeographic analyses. From this point of
other than specific composition (e.g., water masses and view, water temperature seems to strongly outweight
currents, primary production, biomass, seasonality, all other parameteres as far as life types are concerned
specific diversity) are relied upon heavily. Inventorial (but not for the distribution of abundance, biomass
analyses alone, often due to the paucity of data, fail to and productivity). As a result, biogeographic patterns
identify major ecologically and biogeographically generally follow the classical 9-belt system (paired
meaningful features. Furthermore, especially when polar, subpolar, transitional, and subtropical bands on
used alone and in a quantitative manner, faunal distri- both sides of a tropical or equatorial one). This 9-belt
butional data tend to break areas into too many regions, system is primarily derived from physical data, which
many of which have little or no ecological meaning raises the long-standing question as to whether the
(Fager and McGowan, 1963; Beklemishev, 1969; distribution of biogeographic domains reflects physi-
Dadon and Boltovskoy, 1982). cally defmed water masses. This question may seem
superfluous in the light of the wealth of reports where
Early biogeographic schemes of the South Atlantic the distribution of individual species and species
(and of the World Ocean) were almost exclusively assemblages are concluded to match water-mass pat-
based on qualitative data on the distribution of plant terns. However, because extensive interpolations are
and/or animal species. Hardly any quantitative infor- needed in order to establish large-scale zonations on
mation was then available, let alone data on primary the basis of the highly spotty biological information,
production, biomass, turnover rates, trophic relation- and since these interpolations are usually based on
ships, and the like. Surprisingly, however, comparison physical trends (chiefly temperature and salinity
of these early schemes with those accepted today shows fields), such comparisons are intrinsically biased.
that the size and shape of the major, world-wide bioge- Indeed, when objective data on individual species ran-
ographic provinces changed very little (Boltovskoy, ges are analyzed in detail, the water-mass vs. bioge-
1986, in press). Indeed, for the Southwestern Atlantic, ographic range coincidence is often vague
for example, the biogeographic zonations outlined as (Boltovskoy, 1986), especially in the pelagic realm
early as in the 1930's and 1940's (e.g., Hentschel, (see below). Nevertheless, because -as mentioned
1938; Bobrinskii et al., 1946) do not differ drastically above- it is the physical environment which primarily
from the "biogeochemical provinces" proposed recent- drives planktonic life in the oceans, water masses (and
ly on the basis of satellite imagery (Longhurst, 1995; their temperature gradients in particular) are intimate-
Longhurst et al., 1995; see Fig. 1, inset). ly linked with biogeographically distinct areas.
Several reasons converge to explain this situation. A major goal of biogeographic surveys is producing
Besides the fact that the early schemes proved good one or a few maps which reflect adequately the distri-
enough as to need only minor adjustments, it seems bution of life and life-related processes. Indeed, when
obvious that the physical environmental setting, as discussing biogeography in the pelagic realm of the
defined by water-mass properties and movements, is World Ocean, the first image that comes to mind is the
the main driving force behind the distribution of above-mentioned classical system of belt-like polar to
planktonic life in the seas. Because the general traits equatorial zones stretching across the globe. However,
of surface circulation and water-mass patterns have this picture changes radically if primary production, or
South Atlantic Zooplankton.

edited by D. Boltovskoy, pp. 1-42
© 1999 Backhuys Publishers, Leiden, The Netherlands
--.- ....•.-.,~--.----~
~-------..-.---~---.-.-, , ..-.---.~-'.-----•.... - ~.~.~-- -- .. " ..
, "'--.'"-..............
I
2 General biological features of the South Atlantic
Guinea
Tropical Oceanic Current
Neritic
Benguela
Current
Neritic
30°
Agulhas
Current
Neritic
~
0.... .... ~~~ - - - ----- 40°
~ >o('
00
••••..•••.•···~ooo
t 1)000
Sb . 0
u antarctic
.
cearuc
07>0000\ .
50°
Winter Summer
boundary boundary
Antarctic Oceanic
Guinea
Current
Coastal
South Subtropical •••• --- ••• -+

Convergence L.. -+
Subantarctic
Antarctic
•• Austral
L41~-=-- Polar --------------'
Biogeochemical provinces according to

Longhurst (1995)
Fig. 1. Sketch of major biogeographic domains in the South Atlantic. Western side is chiefly based on E. Boltovskoy (1970, 1981 a).
General biological features of the South Atlantic 3
Fig. 2. Areas with distinct biological characteristics, as defined in Table I. Gi: Guinea Current; Gu: Guiana Current; SSF: Southern Subpolar
Front; SST: Southern Tropical Front; see Table 1 for other abbreviations. From Greze, ed. (1984).
the abundance of life - rather than distinct species functional traits. Thus, much of the ocean surface is
assemblages - are considered (compare Fig. I with reportedly occupied by transitional areas where dissi-
Fig. 3 below). The distribution of endemics, of speci- milar ecosystems intermingle. However, while mixtu-
fic diversities, of particle size, of export production, re in the oceanic realm is definitely a major feature,
etc. will each yield a different map, all of which are our appreciation of its relative importance is largely
relevant to biogeography. Actually, one should proba- derived from the seasonal and interannual fluctuations
bly think of biogeography as a multilayered system of of the boundaries of "core communities". In other
maps, each layer representing a different property words, at any given point in time ecotones are proba-
and/or addressing a different aspect of the biota bly less important than our seasonally and multi-
(Boltovskoy, in press). However, such a multilayered annually integrated data in synoptic biogeographic
system is not amenable to summarizing in one or even schemes indicate. The rather large Transitional
a few simple and straightforward schemes that satis- Oceanic area illustrated in Fig. I, for example, is chie-
factorily represent a significant proportion of the traits fly based on multiseasonal and multiannual data as
of pelagic life. The divisions illustrated in Fig. I are, defmed by the latitudinally extreme locations where
therefore, a compromise solution to this maze of con- subantarctic plankters can be carried northwards with
flicting maps. For example, specific inventories of the the cold Malvinas (=Falkland) Current (northern
Tropical Oceanic domain may not differ greatly from limits), and subtropical ones southwards with warm
those of the Central Gyre (Fig. 1), yet they do have Brazil Current (southern limits) (cf. Fig. 8 below). The
quite dissimilar primary production values, zooplank- area actually dominated by mixed subantarctic-subtro-
tonic biomass, phytoplankton:zooplankton biomass pical assemblages during a restricted period is, howe-
relationships, etc. (cf. Table 1). ver, more limited (e.g., Boltovskoy et al., 1996).
As opposed to the terrestrial milieu, the dynamics of Table 1 (refer also to Fig. 2) summarizes some rele-
open-ocean systems enhances the importance of areas vant traits of the various biologically distinct areas
of gradients between neighboring communities, or that can be recognized in the South Atlantic. These
ecotones, rather than that of clearly circumscribed results are chiefly based on one of the most compre-
domains characterized by particular structural and hensive set of surveys carried out so far in the South
,-----CBel G\1a) ~
~ ~SE Br ~ CG
South B~i1 Malvinas WWind Drift Central Gyre
Benguela Equat. Trop. Subtrop. <=Falkland) Wpart E part North I Central [South
Mean annual temperature (0C) 16 26 22 10 3 22
Primary prod.
Greze (1984) 2469 175-344 115 120 146 296 273 123 I 94 i 201
Longhurst et al. (1995) 880 360-430 830 1300 330-370 210
(mg C m·2 day")
Koblentz-Mishke (1977) 250:. >500 150-250 <100-150 150-500 100-250 <100
Phytopl. abundance 6
(cells x 10 per I; 0-50 rn) (Greze, 1984) 3218 13 12 7 10 5 9 5 I 18 5
Phytopl. abundance (cells r'. 0-100 rn) (Semina, 1977) 10 3
_ >104
3
10 _10 4 2
10 _10 4
103_ >104 103_ >104 -. <10
2
!
Phytoplanktonic biomass (rng m·3; 0-50 rn) 2785 11 10 12 12 5 16 7 I 15 ! 11
5 1.8 3.4 1.4
Bioluminiscence (10. J.lW cm2), 0-100 m 25.2-74.7 3.2-11.8 2.8 2.8 8.0
Phytoplanktonic diversity (number of species) 110 264 155 154 80 46 92 176 I 233 I 78
Phytopanktonic diversity (a) 13 49 42 50 40 24 30 45 I 56 29
2
Zooplanktonic biomass (mg m· ) 70 69 32 23 30
Biomass of herbivores (% of total zooplankton) 47 27 15 22 8
Biomass of carnivores (% of total zooplankton) 0.5 25 15 6 8
General zoopl. diversity (comparative figures, 0-100 m) 3.1 3.91 4.39 3.34 2.52 4.45
Zoopl. abundance (ind m .3; 0-100 m; micro + mesozoopl.) 31816 6492 4565 6549 4858 8945
l
Zooplanktonic production (rng d· at 0-500 rn) 3740 1594 1200 846 580 1734
Number of copepod species recorded 176 215 280 161 300
Biomass Phytoplankton 67.4 5.0 4.7 2.2 8.0
relationships Zooplankton 2.8 34.6 16.8 5.7 21.9
(%) Bacterioplankton 29.8 60.4 78.5 92.1 70.1
Bacterioplanktonic biomass (mg m .3) 754 120 150 372 96
Table 1. Comparative values for various biological parameters in the major biogeographic domains of the South Atlantic. Where units are not clearly specified (in the original source) values
are reproduced for comparative purposes only. See Fig. 2 for location of areas. Data from Greze, ed. (1984), unless otherwise noted. The "South Equatorial" domain is roughly coincident with
the "Tropical Oceanic" domain of Fig. 1. Shaded cells denote highest values.
Chlorophyll a Primary production Zooplanktonic biomass

(mg m-3) (mg C m-2 d-t) (mg 1000 1-1 )
,..------,
CJ~
<0.5
->30
,..---,
~
<100
f'.,;J<~4ill _
i"·-~"~"'.
100-150 150-250
_
>250
CJ CJ
<25 25-50
!:S';l'~~;1__
50-100 100-200 >200
Fig. 3. Distribution of chlorophyll a (composite image based on data from 1978 through 1986 retrieved by the CZCS atellite) (left), primary
production (chiefly from Koblentz-Mishke, 1977) (middle), and zooplanktonic biomass (chiefly from Bogorov, 1974) (right).
Atlantic after the German "Meteor" expeditions of Over half of the South Atlantic is occupied by the
1925-1927, yet it should be noticed that some of them South Atlantic Central Gyre or South Atlantic Central
are based on few observations and therefore probably water mass, a tropical/subtropical area with extremely
do not represent adequately average values for the low nutrient loadings and, in consequence, with very
cprresponding area. For example, differences in pri- low phytoplanktonic and zooplanktonic populations.
/:nary production levels between the Benguela Current Primary poduction values range here around >0.1-0.2 g
and other areas seem too high when compared with C m-2 d-I (Koblentz-Mishke, 1977; Longhurst et al.,
other estimates. Nevertheless, because they constitute 1995; see Table 1), with phytoplankton concentrations
the results of a major interdisciplinary effort and for below 103 cells I-I (Semina, 1977). This area shows
some parameters (e.g., bioluminiscence, biomass pro- clearly both in chlorophyll a and in primary production
portions of the various trophic levels, zooplanktonic maps, as well as in those depicting the distribution of
production, bacterioplankton biomass) are the only zooplanktonic biomass (Fig. 3). In the vicinity of the
large-scale estimates available for the area, their use- equator biological richness is enhanced by equatorial
fulness for a general characterization of the South divergence and by seaward advection of nutrient- and
Atlantic is beyond doubt. biomass-rich Benguela upwelling waters (see below).
On the western side, the Amazon plume can also sea-
sonally contribute to the enhancement of plankton
General remarks on seasonality, biomass and diver- along the equator (see below). Specific compositions
sity in the pelagic domain of the South Atlantic between central and equatorial waters differ little,
although the richer equatorial ones tend to show lower
The quantitative distribution of pelagic life in the equitabilities (i.e., higher degrees of dominance of a
South Atlantic parallels those found in the other oce- few species over the rest of the assemblage). A few
ans: a large area of poor central waters bound to the organisms, however, seem to be especially adapted to
north and south by richer equatorial and subpolar these oligotrophic waters. Such is probably the case of
bands, respectively, with the biologically richest sec- the colonial polycystine Radiolaria (see "Radiolaria
tors circumscribed to the coastal regions, especially Polycystina" in this volume), which host symbiotic
along Africa (Fig. 3). algae in their protoplasm, presumably to compensate
for the scarcity of food in the medium. In some areas role in the biogeochemistry of the oceans. Regardless
the production of these symbiotic algae has been found of absolute values, more or less constant primary
to/exceed that of the free-living phytoplankton (e.g., in poduction levels throughout the year would permit a
the Gulf of Aden, according to Khmeleva, 1967). better coupling between production and consumption,
Mesozooplanktonic wet weight (0.2-20 mm size range) thus allowing for less organic matter to go unused in
in the Central Waters is below 25-50 mg m-3 (0-100 m; the trophogenic layer and sink toward the sea floor. In
cf. Bogorov, 1974; Rudjakov, pers. comm.; Fig. 3). contrast, sudden bursts in plant biomass, as those char-
acteristic of high-latitude and some coastal upwelling
The southern boundary of the Central Gyre is repre- systems (such as the Benguela system, see below),
sented by the West Wind Drift (WWD), a strong cir- have probably less chances to be used fully by the
cumglobal current carrying subantarctic and antarctic slower-reacting consumers, therefore exporting higher
waters from west to east. In the area of the study, the proportions of their matter to depth. These concepts,
gradient between the warmer waters north of the WWD discussed in detail in a paleoceanographic context by
and those to the south of it is probably the most con- Berger and Wefer (1990), are quite relevant to bio-
spicuous in biological terms. Indeed, it was the first geography insofar as they may help explain some of
(and sometimes the only) biogeographic limit proposed the differences between warm water and cold water
in early studies (e.g., Meisenheimer, 1905a), insofar as zooplanktonic asemblages. Thus, differences in spe-
planktonic species can be roughly divided into warm cies richness between tropics and poles are probably as
water and cold water organisms, the former associated much a result of extreme temperature conditions in the
with the tropics and subtropics, and the latter with the latter, as they are of the need to be able to adapt to long
subantarctic and the Antarctic (see Fig. 8 below). periods of starvation characteristic of polar regimes.
Besides temperature per se, several environmental The distribution of biomass in pelagic waters of the
parameteres are strongly modified south of 40-45°S, in South Atlantic seems to be also reflected by that of the
particular the concentration of nutrients, which gradu- numbers of species present, yet this relationship is not
ally increase from north to south, the growing impor- straightforward. Indeed, while for most groups diver-
tance of sunlight as a limiting factor for phytoplankton sity in the tropics is several times that at the pole, on
development, and the seasonality (varying temperature these large scales higher numbers of taxa are recorded
and insolation throughout the year). Typical P-P04 in the Transition Zone, rather than at the equator (Fig.
concentrations below 0.25 g at P 1-1 (0-50 m) in the 4). Thus, biologically poorest waters host the highest
Central Waters increase to >1.5 g at P 1-1 around 45- numbers of species. It should be noticed that the esti-
500S. Nitrates and silicates follow similar trends as mate reproduced in Fig. 4 is partially an artifact of
well. At 600S sunlight limits phytoplankton growth lumping the time-averaged record in a single graph.
between May and August, and from April to July at Insofar as it is the accumulated inventory resulting
700S (Voronina, 1977). The onset of spring autotroph- from sampling the same area throughout several sea-
ic gowth is determined by the increasing availability of sons and years, restricted time offsets would most
light and by the stratification (and consequent stabi- probably lower the figures indicated, especially for the
lization) of the upper water layer; the steep density- colder-water areas in general, and for the Transition in
gradient precludes phytoplankton cells from being particular. Nevertheless, high degrees of mixture of
drawn down away from the photic layer, thus favoring warm- and cold water organisms in the same sample
the buildup of plant biomass. In subantarctic waters are very common in the Transition, as is evident from
spring and summer stratification of this upper layer is the curve for the ostracods (Fig. 4), derived from data
due to heating. Further south, however, where winter collected during a single cruise. In the eastern South
to summer temperature gradients are minor, meltwater Atlantic further enhancements of specific diversity at
from the retreating ice-pack, which spreads along the transitional latitudes are also derived from the advec-
ice-edge as a large 20-50 m thick superficial lens, tion of Indian Ocean waters via eddies and rings asso-
plays a determining role (Smith and Nelson, 1986). ciated with the Agulhas Current (see below).
Seasonality is probably a major factor in the structur- From the ecological point of view this Transition is
ing of pelagic communities, and it also may playa key probably quite dissimilar from the domains that bound
Number of species
10 20 30 1 5 10 2 5 10 5 10 20 3010 20 30 40
00 r-----r ..•...•
-,----,--,
£
•..
J
o 200
tll c
o
.-
}
'~
Q)
<Jl
"'0 400 C
J
•.. ~
•.. ~
(1j
...J
600
, -V
Foram in ifera Chaetognatha Salpida Appendicularia Ostracoda
'--------'
Fig. 4. Approximate numbers of planktonic species that can be recorded throughout the year in pelagic waters of the South Atlantic. Estimate
by Boltovskoy (1982), based on several sources. Ostracod data are from a single cruise (Deevey, 1974).
it from north and south. These organisms are almost often the result of lack of information, than of the fact
exclusively expatriates from the neighboring systems, that the species in question does not actually live in the
since very few zooplankters seem to thrive here better South Atlantic. Sorting out these two influences on the
that in Subantarctic or Subtropical waters. Thus, they database is clearly impossible with any degree of con-
most probably are "terminal" assemblages (sensu fidence, yet comparisons between groups can yield
Beklemishev, 1969) of organisms circumstantially suggestive general trends useful for future work.
brought together, with little chances of surviving and
reproducing. In consequence, because they do not share One of the questions we attempted to address using
an evolutionary history, their interactions are loose, the information contained in this volume and previous
rather than based on mutual adaptation (Boltovskoy, data is how many of the species described for each
1986). zooplanktonic group occur in the South Atlantic. The
data reported for 24 zooplanktonic taxa (of a total of
30 covered in the volume, some were excluded from
Patterns of distribution of zoo planktonic species the analyses due chiefly to lack of adequate infonna-
tion) indicate that these percentages vary widely, from
Results presented in this volume offer a unique oppor- 100% (Cladocera) to as little as 10-15% (Nemertina,
tunity to assess in a comprehensive manner our present Ctenophora, Mysidacea) (Fig. 5).
understanding of some general aspects of specific rich-
ness in the area covered, as well as selected traits of the Some of the datapoints illustrated in Fig. 5 clearly
distribution patterns of the species present. Tentative deviate from the overall trend. For the Nemertines an
comparisons are drawn within and across major taxo- overall total of 97 marine planktonic species has been
nomic groups. Admittedly, interpretations are very described (R. Gibson, pers. comm.), only 11 of which
strongly influenced by both uneven intrinsic specific have been found in the South Atlantic. However, most
diversities, and by the very variable degrees of knowl- described nemertine species have only been recorded
edge of the various taxa concerned. Indeed, while po- once or twice, which clearly shows that available data
sitive records pose little problem, negative ones (i.e., are totally insufficient for drawing a world-wide dis-
the absence of a given species) are probably more tributional pattern for these scarce and delicate deep-
~----------------------------------,-100
2000
0:0
I::
.-({l'O
~ 1600
u..r::.
ClI III
a.~
__
III "0
ClI
•..
0'-
~
1200
ClI"O
...c-.::
E 0
:::J ~ 800
1::"0
-Cli
~...c
~'5 III 400
ClI
"0
0 o
Fig. S. Nwnbers of species known to date for each of the major marine zooplanktonic groups (shading), and proportions thereof recorded in
the South Atlantic (line).
dwelling animals. The Mysidacea are also atypical in some unrealistically low percentages (e.g., Acantharia,
that only 96 of the ca. 700 known species have been Cephalopoda, Radiolaria Phaeodaria, Copepoda). The
found in the South Atlantic. As opposed to the nemer- Radiolaria Polycystina, also with a seemingly relative-
tines, in this case the actual proportion of geographi- ly low coverage, constitute another exception. In this
cally restricted species may effectively be higher than case, the figure used as an estimate of the total number
for other groups. Most mysid species are closely asso- of Recent species described is a very rough approxi-
ciated with the sea floor, and therefore their ranges are mation; most of these taxa are very poorly defined, and
influenced by bottom topography, rather than by sur- in fact they are ignored altogether in most surveys (see
face water masses and currents. Dispersion by currents "Radiolaria Polycystina" in this volume). The species
is further hindered by their comparatively large size, covered in the corresponding chapter, which include
powerful swimming, and the lack of planktonic larval both actual and potential records, are therefore a selec-
stages. Thus, even though their coverage in the South tion of the most widespread and better defined poly-
Atlantic is poor, and future work will undoubtedly add cystines.
many species, the proportions of South Atlantic taxa
with respect to the overall world total will probably These considerations indicate that several of the data
increase further, rather than decrease (M. Murano, illustrated in Fig. 5 are governed by factors other than
pers. comm.). actual degrees of endemism within each group. The
overall specific richness within each major taxon, in
For most of the groups dealt with, species covered particular, seems to influence the numbers strongly.
include both those effectively recorded in the South The species-richest groups, such as the fish larvae
Atlantic, and the ones not yet recorded, but whose and the copepods, have proportionally the lowest
presence in this basin is very likely. In a few cases, numbers of South Atlantic species «30%); while for
however, general information available was too scarce the less diversified ones, such as the Cladocera,
or otherwise inadequate for this type of approach, and Pyrosomatida, Heteropoda, Salpida, Appendicularia
only actual records were considered. This resulted in and Euphausiacea, at least 60-70% of the species
COLDER WATER
~
~ .~~
ra
"'C
0
ra
"'C
ra
"'C
ra
~
~
"'C
0
c.
~
~
~
u
~
.;;;
•..0ra
..s:
ra
"'C
-~
C
~
..s: •..~
0
..s:
C. ~
:E
ra
c
.~
•..
~
~
.:::
-;
.~ .rara.:::
.e-
-
u t).O
0
c. 0
c. ra
0
~ ~ c. 0 0 <II
·e•..
C "t:I -e
--
..s: ~ u 0 ·c >- e
•.. ~ "'C ..s: .a.
"'C ra
..s: 0
•..
ra ~ ]! u 0
-
c. 0 0 QI C
C. .;;; C. c. c ra ..s: >- ra c. QI
ra ra
E ~ 0 >- ~ -; ~
QI
fIl ..s: c. "'0 "'0 c. ..s: QI
-« Q..
U ~ U tn L.U U 0 U V; c Q.. 0
1.1.. -e Q..
~
Equatorial(rropical
69%
(0-1005)
Central/Subtropical
(10-3505) 75%
Transitional/Temperate 58%
(35-4005)
Subantarctic 43%
(40-50°5)
Antarctic 29%
(>5005)
Total species
considered 147 101 456 55 97 29 49 20 102 37 82 10 164 39 44 163
100%
I-----l
Fig. 6. Numbers of species of 16 zooplanktonic groups occurring throughout the 5 main biogeographic areas of the South Atlantic, expressed
as percentages of the totals within each group considered (totals are indicated in the bottom row).
described were found to be present in the South the mixed subtropical-subantarctic area are probably
Atlantic. Groups which had locally-based (Brazil, underestimated by data illustrated in Fig. 6 (compare
Argentina, South Africa) specialists for a number of with Fig. 4), since they emphasize "core" areas of
years, such as the foraminifers, planktonic tunicates, occurrence, rather than expatriation ranges. As dis-
pteropods, euphausiids, chaetognaths, also seem to be cussed above, expatriation of both warm water and
better covered than those lacking local expertise. cold water organisms is extensive in the Transition
Zone (see Fig. 4). Diversity drops sharply in the sub-
In view of all these limitations, an accurate estimate of antarctic, and further decreases south of the Polar
the percentage of planktonic species present in the Front, where on average it is less than 50% that in the
World Ocean that inhabit the South Atlantic is clearly tropics and subtropics (Fig. 6).
impossible. Data for the groups which are probably
least biased, however, suggest that these proportions North-south decrease rates in species numbers, howev-
vary from 80-100% to below 50%. er, are uneven between groups. For the "warmest" taxa,
such as the Pteropoda and Copepoda, subantarctic and
In general terms, highest zooplanktonic specific rich- Antarctic representatives are only about 10-20% of the
ness occurs in the subtropical waters, roughly between totals present in the entire South Atlantic; on the other
10 and 35°S, followed by the equatorial belt (Fig. 6). extreme, for the Appendicularia and Phaeodaria
Numbers of species that can effectively be recorded in Antarctic waters host almost 50% of the species.
••....
Percentage of total species recorded in the South Atlantic

o )0 60%
L...L..ll-.-L1
0·10° 7%
-------- I
"'C
QI
10.20° 9%
c:
c: -/
~
c.
<II 20·30° 13%
QI
-- - - - - - - - - - - - - - - - - --I
-
"'C
::I
'';:; 30·40° 31%
.!!!
<II ---I
~9r/o
QI
•..0.0
QI
40·50°
QI
c
50·60° 9%
- ----------I
>600 13%
~ ~ ~ ~ ~ •..~ ~ ~ •..~ ~
-
QI
"'C ';: "'C .c: "'C
::! QI "'C ~
•..
0 ~ 0 ~ 0 0 1.1
~
QI
:-=c: o o
C.
.s:
"5 c.
0 c:
c.
QI
.c:
C. :§ 'r;;
1.1
~
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c.
- c.
1.1 0.0
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-
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c. :sc: ~
.c:
0 0 c: c ~
•..
-:;; o
E
4( QI
c.
C.
QI
QI
~
.c:
U QI
U
.c:
c. 0
o c:
o
.c:
::I ••• c.
C. U U 1.1.I
4( Vi
Fig, 7, Approximate latitudinal spans of the species of 12 zooplanktonic groups in the South Atlantic, expressed as percentages of total spe-
cies considered for each of the groups indicated.
Data available for some of the groups treated also ranges are asymmetric on the two sides of the ocean
allow a very rough estimate of the latitudinal span of (see Fig. 1), these results are clearly very rough. Never-
the zooplanktonic species covered. Fig. 7 shows that theless, we contend that they still do reveal a meaning-
few taxa are restricted to ranges below 10-20°, or ful trend (Fig. 8).
occupy areas in excess of 50°; instead, most zooplank-
ters occur over moderately large areas spanning By far highest numbers of distributional boundaries
around 40° in latitude. In terms of biogeographic divi- occur within the Transition Zone. This confirms the
sions, these estimates confirm some previous data widely accepted notion, already noticed in the first
(Boltovskoy, 1986) concluding that both widely cos- large scale biogeographic schemes proposed (e.g.,
mopolitan (i.e., tropical to polar), and strongly Meisenheimer, 1905a; see above), that the two major
restricted zooplanktonic species are rare (Fig. 7). In structural units in the area are the cold water and the
other words, most species are not restricted to anyone warm water assemblages. Quantification ofthis parti-
water mass; their ranges usually cover at least one cular limit as compared with other boundaries, howev-
entirely, and at least parts of one or two others. er, has seldom been attempted. Our results indicate that,
in these terms, the cold water vs. warm water boundary
Definition of faunistically discrete biogeographic units is about twice as important as the second strongest one,
is chiefly based on estimates of the distributional boun- the Polar Front, and about 3 times stronger than the one
daries of the species. In an attempt to validate the separating tropical from subtropical domains (Fig. 8).
scheme presented in Fig. 1, on the basis of the data These latter two also appear as significant breaks in the
available we calculated how many of the 687 taxa con- geographic ranges of the zooplankters considered. The
sidered appear or disappear at each degree latitude one centering on 17-200S represents the boundary
between 3 and 58°S. Since data used for this calcula- between the equatorial/tropical and the subtropical
tion are very approximate, and because most specific domains on the west, and the Angola/Benguela Frontal
Tropical-Subtropical Transition Subantarctic-Antarctic

boundary Zone boundary
..£1- ..£1-
30
.., ,--,
.A.
tJl
Q) -~ I
"...
U tJl
~.~
tJl
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E
20
..
0-
...••.. ,"
,
•.. Q) '"
Q)oo l.-... Iii'" ""IIIiiIr
\
.o c 10 r: V'
E~
••
:J ~
Z
o
00 100
- --I
200 300 400 500

Latitude South
Percent species in common
of those present in the two areas compared
Tropical Subtropical Transitional Subantarctic Antarctic
(0-2005) (21-3505) (36-4005) (41-5005) (51-6005)
Total species 580 578 383 285 185
Subtropical 84
Transitional 51 62
Subantarctic 31 38 62
Antarctic 16 21 35 60 I
Fig. 8. Top: numbers of species distribution limits for 687 selected zooplanktonic taxa in the South Atlantic as a function of latitude (curve
smoothed by a 5-point running mean). Bottom: compositional similarity between the biogeographic areas indicated as derived from the num-
bers of taxa shared. Based on pooled data for 687 taxa in 11 zooplanktonic groups.
Zone on the east (cf. Fig. 1, 27). In open-ocean waters allowing for the maintenance of permanent, exclusive-
this limit is probably less pronounced and occurs far- ly transitional populations are unknown (Reid et al.,
ther north, around 10 S (Fig. 1).
0
1978; Boltovskoy, 1986; see above). The numbers
illustrated in Fig. 8 (lower panel) support this notion
The bottom panel of Fig. 8 confirms the results of the insofar as the corresponding percentages of shared spe-
graph insofar as the numbers of species shared between cies are the second highest (62%). Interestingly, pene-
areas are generally inversely proportional to the num- tration of subantarctic organisms northwards seems
bers of specific distributional boundaries between much stronger than expatriation of subtropical organ-
them. Thus, 84% of the 629 tropical and/or subtropical isms southwards. Indeed, only 63% of the 609 species
species considered are present in both these areas, present in the subtropics are also recorded in the
whereas subantarctic and Antarctic waters share 60% Transition, whereas for the 285 subantarctic plankters
of the 294 taxa present in either. As opposed to the the figure is 90%. It is probable, however, that this dif-
above, the Transition Zone is actually an ecotone, a difference is at least partly due to submersion and north-
fuse boundary in itself, rather than a "community". ward expatriation of cold water organisms at subsur-
Almost all its inhabitants are expatriates from neigh- face and deep layers, rather than at the surface (see
boring areas, and recirculating physical mechanisms "Argentinian coastal and offshore waters" below).
The western South Atlantic and multiannually. Thus, the northern winter boun-
dary of the Trasition illustrated in Fig. 1 (western sec-
A comprehensive scheme of the major biogeographic tion) is defined by the northernmost winter records of
divisions in the Southwestern Atlantic was published typically subantarctic planktonic foraminifers
by E. Boltovskoy in 1970, which was extensively used throughout a 20 year period. Similarly, the southern
to define boundaries in the western part of Fig. 1. This winter boundary of the Transition reflects the south-
work was based on the analysis of the planktonic ernmost winter records of subtropical foraminifers in
Foraminifera from over 3500 samples (mostly super- the same time span. The Transition Zone was defined
ficial), collected over a ca. 20-year period. Despite as an area of mixing of subtropical and subantarctic
some drawbacks, including irregular temporal and faunas, as well as large areas occupied by tongues and
spatial coverage, paucity of information in the coastal patches of purely subtropical or purely subantarctic
and neritic areas (where planktonic foraminifers are organisms (see Fig. 14 below). However, in this time-
either very scarce or absent altogether), usage of only integrating image these features are necessarily
one zooplanktonic group (rather than several), this ommitted. Interestingly, the southern limit of this
scheme is the first contemporary large-scale attempt at Transition agrees well with subsequent information
defining a plankton-based zonation of this area. derived from satellite images, while the northern one
Details of these results were discussed in several sub- is noticeably displaced to the north (compare Fig. 1
sequent publications, and so far in general terms they herewith with Fig. 7 in Olson et aI., 1988). We con-
seem to hold not only for the foraminifers, but also for tend that submersion of cold water plankters, in par-
many other planktonic taxa. A major contribution of ticular the foraminifer Globoquadrina pachyderma,
this work was the recognition of the Transition Zone recorded in subsurface tows by E. Boltovskoy, is
(=Subtropical-Subantarctic Convergence Zone, or responsible for this mismatch (see below).
Brazil-Malvinas=Falkland Confluence), a large area
between approx. 30 and 46°S where subtropical Northern Brazil
waters of the Brazil Current collide, overlap and mix
with the subantarctic ones of the Malvinas (=Falkland) Along the Brazilian coast, the area toward the north-
Current. Previous and subsequent investigations con- west of Cabo San Roque (5 S) differs from that to the
0
tributed much information on details of the distribu- southwest of this site in that it hosts the mouth of the
tion of various planktonic organisms in this area. Amazon drainage basin. This river discharges about
Enrique Balech, in particular, on the basis of dinoflag- 190,000 m3 s-1 of fresh water into the sea, its plume
ellates and tintinnids, starting in the early 1940's made extending far into the open ocean where it can be
several major contributions to our understanding of traced up to 1000 km to the north and east of the river
biogeographic traits of the Transition Zone and of the mouth (Lentz, 1995; Limebumer et aI., 1995). The
coastal stretch between southern Brazil and Patagonia. nutrients thus exported are responsible for a permanent
However, because E. Boltovskoy's (1970, 1981a) is a diatom bloom between the 10 and 30 m isobaths (up to
time-integrating image, these details of seasonal and 30 mg Chi a m-3; Muller-Karger et al., 1988). During
interannual variations are not depicted in his map (Fig. February-May Amazon water flows into the Caribbean
1, western part). at ca. 90 ern sol in a broad (150-200 km) continuos
band of high pigment concentrations (>1.5 mg m-3; as
The domains defined by E. Boltovskoy (1970, 1981a; compared with background concentrations in oceanic
see Fig. 1, western part) include a Tropical zone, waters of 0.15 mg m-3). Between January and June this
stretching roughly from the equator to ca. 20oS, diffe- plume is carried northward, around a retroflection of
ring little from a Subtropical one (20 to 30-35°S, the Guiana Current, and then turns east, toward Africa,
depending on season and longitude); a vast offshore joining the North Equatorial Countercurrent (Muller-
Transition Zone (30-35°S to 46-48°S in pelagic Karger et al., 1988). This mechanism seems to be a
waters); a Subantarctic Zone; and south of the Polar major contributor to the low salinities observed across
Front the Antarctic Zone. Of these, the Transition the tropical Atlantic between 5-1O~, to enhanced zoo-
(and its boundaries with the neighboring subtropical planktonic biomass (Greze et aI., 1969), and to high
and subantarctic zones) is the most dynamic insofar and very variable rates of sedimentation of organic
as, as noted above, its limits vary widely seasonally aggregates in the northwestern Atlantic (Deuser et al.,
~~-----------------------------------------------------------------------------------------------------------------------------------------
1988). In view of its enormous discharge, the biologi- of 20 S the Brazil Current strengthens and moves
0
cal and biogeochemical significance of the Amazon somewhat farther offshore. Pires-Vanin et al. (1993)
plume (primary production, organic carbon burial, summarized some outstanding features of the physical
sediment accumulation) is such that it has been report- and biological oceanography around Ubatuba (approx.
ed to affect not only regional, but also global mass 24°S) (see also the chapter "Fish larvae" in this vol-
budgets (Nittrouer and DeMaster, 1996). Another ume). Seasonal changes are mainly driven by trans-
effect of this freshwater plume is the rather dramatic gressive movements of fresh South Atlantic Central
local drop in the diversity of most oceanic planktonic Waters onto the middle and inner shelf. In late spring
groups (e.g., chaetognaths, siphonophores and medu- and summer these nutrient-rich waters extend coast-
sae; Alvarifio, 1968). ward as a thick subsurface layer the top of which lies
within the photic zone. Nannoplanktonic poducers
Coastal circulation off north-eastern and central Brazil account for a major part of the chlorophyll a (around I
is chiefly defined by the southward-flowing oligo- mg m-3), while salps are the dominant primary consu-
trophic Brazil Current; because the shelf over this mers. In the winter South Atlantic Central Waters
entire area is narrow, typically neritic populations are retreat oceanward, thermic stratification disappears in
loosely delimited and pelagic flora and fauna invade the coastal region, and primary production is mainly
the coastal domain regularly. Along the northern coast fed by wind-driven mixing and sediment resuspension.
of Brazil, both phytoplanktonic biomass « 1000 cells Moderate faunal gradients occur vertically and perpen-
ml+, <0.2 g ChI a I-I; cf. Valentin, 1989), and primary dicularly to the coastline in association with the
production (ca. 300 g C m-2 y-l, cf. Longhurst, 1995) replacement of water masses of different origin (e.g.,
are low, with the exception of some restricted areas Bjornberg, 1981; Christoffersen, 1982). Overall pri-
(e.g., off Cabo Frio, at 23°S) where quasi-permanent mary production is low, generally comparable to that
wind-driven coastal upwelling conditions can enhance of the northern sector; in oceanic areas it ranges around
pimary production values and foster the development 0.45/lg C I-I h-I, while in coastal waters it is somewhat
of colder- and deeper-water planktonic assemblages at higher and more variable (1.08-15.52 pg C I-I h-1; cf.
the surface (Valentin, 1980, 1989). Metzler et al., 1997). As in most other oligotrophic
areas, regenerated production is dominant over new
Central Brazil production, with reduced forms of nitrogen accounting
for 74% (offshore) to 96% (inshore) of total nitrogen
Most of the biogeographic schemes proposed for this utilized by phytoplankton (Metzler et al., 1997).
area agree in the presence of a discontinuity around Isolated coastal and shelf-break upwelling cells, like
20-23°S, which is generally interpreted as a boundary the ones at 24°S and 2.6°S (shear-related vortices, pro-
between the tropical and the subtropical domains (Fig. bably enhanced by topographic discontinuities) may
9; see reviews in Dadon and Boltovskoy, 1982; host both higher production and unusually cold- and
Palacio, 1982; Bisbal, 1995). This break was already deep-water planktonic assemblages (Brandini et al.,
suggested almost 150 years ago on the basis of SST 1989; Brandini, 1990; Pires-Vanin et al., 1993); under
data (Dana, 1853; Fig. 9), and was subsequently con- these conditions the share of new production is pro-
firmed on the basis of floral and faunal data. It is espe- bably enhanced.
cially noticeable in benthic assemblages (benthic
foraminifers, hydrozoans, echinoderms, molluscs, Southern Brazil
crustaceans) and fishes, where a large poportion of the
north-Brazilian species is shared with the Caribbean At 25-28°S the axis of the Brazil Current moves even
(Vannucci, 1964; Semenov and Berman, 1977; farther offshore and the neritic area is seasonally influ-
Semenov, 1978). Because the Brazilian coast from 5 S 0
enced by Rio de la Plata and Lagoa dos Patos outflow
to ca. 30-35°S is dominated by the southbound Brazil waters which are carried northwards by the coastal,
Current, changes in the plankton at 20-23°S are minor Malvinas (=Falkland)-driven flow (Castello and
and gradual; nevertheless, modifications in the specif- Miiller, 1977; Hubold, 1980a, b; Brandini, 1990).
ic composition of various pelagic groups (e.g., plank- Chlorophyll a values are around 0.2-3.0 mg m-3, but
tonic foraminifers, cephalopods) have been reported isolated peaks of up to 25 mg m-3 can be recorded in
to occur at or slighly south of Cabo Frio (23°S). South association with the fertilizing effects of freshwater
.-"
(0
r-,
r-, 0'>
0'>
C
I1l
0
r-, E
•...
-.-
0'> (])
co
~ c-, ""0
N
lJ') 0' 0 c co
.-
("t')
..::£
.- .-
0'> ..0 I1l 0'>
lJ')
V'>
co 0'> > >
...s::::: .9 0
c 0
I1l
c
U
(])
X "0 (]) u
I1l I1l
0
C
co E
(])
I1l
I1l
0 co ~ w V1
a,
00 r------~....
~~
10or-----------------~~
20° /------------}--
Subtropical
30° /--------../---.,.,.
40° /---+------1
Subantarctic
50° ~-------I
Fig. 9. Biogeographic divisions of coastal South America according to various authors.
runoff (Hubold, 1980a, b) and colder waters of subant- smaller plankton of areas strongly influenced by the
arctic origin, especially during the late winter and Brazil Current, generally north of 25-26°S and off the
spring (Odebrecht and Garcia, 1997). Data on particu- slope, is dominated by nanoflagellates, coccolithophor-
late primary production rates off southern Brazil have ids, blue-greens and small «20 Jll11) naked ciliates;
been summarized by Odebrecht and Garcia (1997), while large naked and thecate heterotrophic dinoflagel-
reporting between 12.3 and 120.3 mg C m-2 h-I, with lates, diatoms, silicoflagellates, large «40-80 11m)
peaks in excess of 340 mg C m-2 h-1 (yearly average: naked ciliates and tintinnids are more abundant in areas
approximately 160 g C rrr-'). In general terms, plank- influenced by coastal and subantarctic waters (Brandini,
tonic production and biomass are highest inshore and 1988; Odebrecht and Djurfeldt, 1996; Odebrecht and
decrease oceanward, as one enters the oligotrophic Garcia, 1997). A roughly similar transition is also
waters of the Brazil Current (Vannucci and Almeida observed between winter communities (characterized
Prado, 1959; Vannucci, 1961; Hubold, 1980a, b; by large-sized producers and relatively short and highly
Brandini, 1990; Monro et al., 1997). However, occa- efficient food webs) and summer associations (domi-
sional upwellings of nutrient-rich South Atlantic nated by small producers and bacteria, which in turn
Central Water along the continental slope, especially give rise to more complex trophic relationships where
during the summer, may strongly enhance the produc- mucus-feeding mesozooplankters, like Thaliacea and
tivity of this offshore sector (Brandini, 1990). The Pteropoda, playa dominant role) (Castello et al., 1997).
550 500
300
I-uu ul >50% Tropical Water
• >50% Subantarctic Water
~ >50% Coastal Water
----._.-
Winter-Spr ing 1977

Chlorophyll a (August-N ovember)
(rng m") Tropical 0.02- 0.2
---
Subantarctic 0.2-2 .25--T-------
Coastal 0.12- 22.4
Zooplankton biomass
--------.------~
(displacement volume, Tropical 0.01-0.1 0.01-0.1
Water masses ml m -3, 1---'----. -t--
0.31-0.78
250 IJm - mesh net, Subantarctic 0.34-0.88 I
Autumn 1978 (10m) 0-200 rn) Coastal 0.04-0.81 0.31-1:.:..5=--_J
Fig. 10. Surface water masses and associated chlorophyll a concentrations and zooplankton biomass off southern Brazil, Uruguay and northern
Argentina at two times of the year. Based on data from Hubold (I980a, b).
Representative images of the distribution of water are circumscribed to the seaward sector, coastal ones
masses off the southern Brazilian, Uruguayan and stretch some 50-70 miles offshore, while the band
northern Argentinian coasts, and the associated phyto- separating these 2 assemblages is dominated by cold
and zooplanktonic abundances, are nicely illustrated water plankters (Sagitta tasmanica, Limacina retro-
by the results of Hubold (1980a, b), partially repro- versa) swept in by the Malvinas (=Falkland) Current
duced in Fig. 10. (Fig. 11). These subantarctic influences can be traced
as far north as 31-32°S, but during the summer the
Resgalla (1993) produced an interesting analysis of subtropical-subantarctic front retreats southwards
the zooplanktonic assemblages (Pteropoda, Cladocera beyond the Rio de la Plata estuary. However,
and Chaetognatha) off the Brazilian coast between 31 Malvinas (=Falkland)-related waters and the associat-
and 35°S (Fig. 11). During the summer, the area is ed benthic and planktonic biota are present year-round
dominated by neritic warm water fauna exclusively at depth along the Brazilian coast as far north as 20 S 0
(several Cladocera, Sagitta tenuis, Creseis virgula). In (E. Boltovskoy et aI., 1996 and references therein; see
winter, on the other hand, tropical-subtropical species below).
,-
I
Winter 1988
320
330
0-25 m 25-50 m
530
Sagitta tenuis
~ Podon polyphemoides
coastal~
Evadne nordmanni
Sagitta serratodentata
33 0
Tropical Sagitta minima
(Brazil C.) Sagitta enflata
Subtropical D Krohnitta subtilis

Sagitta decipiens
Subantarctic
(Malvinas C.)
D Sagitta tasmanica
Limacina retroversa
50-100 m
530
Fig. 11. Biological zonations of southern Brazilian coastal waters based on the distribution of zoo planktonic assemblages. From Resgalla (1993).
Rio de la Plata estuary beyond the area where most of the suspended silt and
clay sediments, primary production peaks noticeably,
At appoximately 35°S, the mouth of the Rio de la Plata exceeding conspicuously values typical of the neigh-
defines a major hydrographic and biogeographic boun- boring Malvinas (=Falkland) and Brazil waters. Both
dary. Fresh waters of the Parana-Uruguay drainage the estuary itself and the adjacent oceanic waters influ-
basin extend hundreds of miles into the sea thus form- enced by its plume are subject to ample temperature
ing an effective dispersal barrier for many planktonic and salinity fluctuations, partly due to seasonal chang-
species. The estuary of the Rio de la Plata has a total es, and also to a large extent as a result of prevailing
surface of about 30,000 km2 (Fig. 12A), with a mean wind, precipitation (and consequent runoff), and tide
feshwater runoff of 20,000 m3 s-l. In spite of the very conditions (Guerrero et al., 1997). Surface salinities
high nutrient loadings associated with the freshwater vary widely within the area of influence of estuarine
plume, primary production in the inner sections of the waters; yearly changes off Montevideo and Punta del
estuary is low due to the strong light-limiting condi- Este (Fig. 12A), for example, range from 13 to 35%0,
tions imposed by the high concentrations of suspended with spans up to 20%0 in a few days being common
inorganic particulates. Farther oceanward, however, (Ferrando, 1962; Milstein and Juanico, 1985). Usually
Surface salinity fields in the RIO de la Plata influence area

(February-March)
53°
OUTER ZONE
ATLANTIC
OCEAN
25
50
Typical salinity profile along the estuary (0/00)
55°
Fig. 12. Horizontal and vertical salinity fields and general zonation of the Rio de la Plata estuary (A and B, based on Boschi, 1988), and zoo-
geographic areas off its mouth, as defined by distinct assemblages of several crustacean zooplankters (copepods, amphipods, cladocerans,
euphausiids and ostracods collected from 54 stations in November, 1986) (based on Fernandez Araoz et al., 1991).
there also is a strong vertical salinity stratification, few other holo- and meroplankters have been identi-
with marine waters (saltier and denser) penetrating fied (Boschi, 1988; Milstein and Juanico, 1985),
deeper into the estuary along the bottom, while fresh including both species closely associated with subtrop-
waters extend farther oceanward on the surface (Fig. ical waters, and those which dwell best farther south,
12B). Planktonic populations in the estuary are almost in the Malvinas (=Falkland)-influenced subantarctic
exclusively represented by marine eurihaline species, ones. Off Punta del Este (Fig. 12A) mean yearly zoo-
whereas freshwater representatives are practically planktonic densities (180 um-mesh net) around 3,900
absent (Boschi, 1988). Among the former, several tin- indo m-2 have been recorded, with very conspicuous
tinnids (Souto, 1974), appendicularians (Esnal, 1973), qualitative and quantitaive variations in very short
ctenophores, medusae, cladocerans, copepods and a periods (Milstein and Juanico, 1985).
Fig. 13. SST satellite image of the Brazil-Malvinas (=Falkland) Confluence area on 3 November 1996; lighter shading denotes warmer waters
(NOAA, RSMAS).
Argentinian coastal and offshore waters images do indeed suggest a branching of the Brazil
Current north of the estuary of the Rio de la Plata,
Much of the literature dealing with the distribution of with a major arm deflecting to the southeast and a
benthic coastal/neritic flora and fauna concludes that minor one invading the shelf in a southwest direction
2 biogeographic provinces can be recognized along (Fig. 13). Alternatively, Balech (1986) suggested that
the coasts of southern Brazil to Tierra del Fuego: an this warmer water biota is the biological imprint of a
Argentine province, extending to 43°S, and a backflow of Malvinas (=Falkland) Current waters
Magellanic province between 43°S and the Drake which retroflect southward along the coast after hav-
Passage (e.g., Lopez Gappa and Lichtschein, 1988, ing been heated and partially mixed with coastal sub-
and references therein; Fig. 9). Planktonic groups do tropical waters at lower latitudes (the "Argentine
not seem to follow an identical pattern, yet a boundary Flux" sensu Severov, 1990). It should be pointed out,
at 43°S (or 47°S, see Fig. 1) is also often identified. however, that at least as far as spore- or cyst-forming
The neritic area located between Rio de la Plata and organisms (mainly phytoplankters) are concerned,
Peninsula Valdes (43°S), or even Cabo Dos Bahias local populations of warmer-water preferences could
(47°S), is of special interest insofar as, while general- be maintained in dormant stage on the sea-floor,
ly dominated by subantarctic species, it periodically blooming with the onset of adequate ambient condi-
hosts significant proportions of subtropical plankters tions in the water-column. This being the case, Brazil
(Balech, 1949, 1964a, 1976; E. Boltovskoy, 1970; water intrusions would not be required to explain their
Lange, 1985). Benthic assemblages in this area also presence in this area.
show clear relationships with those from Brazilian
shelf waters (e.g., benthic Foraminifera, cf. E. Mixing of subantarctic and subtropical waters off
Boltovskoy, 1964). These findings are interpreted as Southern Brazil down to ca. 45°S is nicely illustrated
the result of the (summer) intrusion of Brazil Current by results based on analyses of the provenance of
waters which slip between the main axis of the planktonic organisms recorded in the samples (i.e.,
Malvinas (=Falkland) Current and the coast and move hydrological indicators or tracers). Fig. 14 illustrates a
southward along the shelf. Summer SST satellite selection of these showing the wedge-like intrusion of
A c
D _ Subtropical
Mixed
I,TiIT:1 >50% Subtropical
[ZJ >50% Subantarctic
1,/ I Subantarctic waters
Fig. 14. Identification of various degrees of mixing of subtropical and subantarctic waters based on their planktonic contents along the Brazil-
Malvinas (=Falkland) front. A: based on dinoflagellates and tintinnids, averaged data for several years. B: same as A, for November-December
1962. C: based on planktonic Foraminifera, March. D, E: based on planktonic Foraminifera, Pteropoda and Chaetognatha, July (D) and January
(E). A: redrawn from Balech (1976); B: redrawn from Balech (1965); C: redrawn from E. Boltovskoy (1962); D, E: based on data from
Boltovskoy (1975).
the cold Malvinas (=Falkland) flow into the subtropi- in the spring and autumn, diatoms being the most
cal domain (Fig. 14A, B, D, E; see also Fig. 13), conspicuous producers (Carreto et al., 1995). The
extensions of Brazil Current waters to the east of this zooplankton hosts typically coastal species, (e.g.,
intrusion (Fig. 14A, B, C), as well as the coastal the copepods Paracalanus parvus, Calanoides
tongue with warm-water organisms reaching as far as carinatus, and the cladocerans Evadne nordmanni,
46-47°S (Fig. 14A; see also Fig. 13). These maps also Podon spp., Pen ilia avirostris; Fernandez Araoz et
stress the irregularity and extremely high temporal aI., 1991).
instability of the front in question (e.g., Olson et al.,
1988), and of the concomitant distribution patterns of (b) The Subantarctic shelf waters system is separated
the organisms concerned. from the coastal one by a coastal front, which
breaks down during the winter. A very strong sea-
Off Buenos Aires Province (ca. 37-39°S) a clear cross- sonal (summer) thermocline develops in this area
shore zonation is present (Fernandez Araoz et al., at depths of ca. 30 m. Phytoplankton growth has
1991; Carreto et aI., 1995; see Fig. 12C): two well defined maxima in spring and in autumn,
with a conspicuous gradient of higher values sea-
(a) In the inner or costal system (to ca. 30 km offshore) wards. This area is periodically influenced by Rio
wind- and tide-induced mixing yields a homogene- de la Plata outflow waters.
ous water-column year-round. Phytoplankton
growth is chiefly nitrogen-limited, but swift nutri- (c) The Malvinas (=Falkland) system is dominated
ent regeneration yields chlorophyll a values by the northeastward-flowing cool Malvinas
between 1 and 4 mg m-3. Seasonal variations in (=Falkland) waters and separated from the shelf
phytoplanktonic abundance are low, with maxima system by a well defined shelf-break front. Plants
t</·, I area sampled

• >1000 mm3/m3
Fig. 15. Zooplanktonic biomass (displacement volumes excluding fish eggs and larvae, 333 urn-mesh nets) off Argentina based on 18 cruises
carried out in 1978-1979 (see source for asumptions used for data conversions). Modified from Ciechomski and Sanchez (1983).
and animals are here represented by typically sub- makes them advantageous feeding grounds (C.
antarctic species (Fernandez Araoz et al., 1991), Campagna, pers. comm.).
although during the summer warmer-water forms
are common as well (Lange, 1985). In general terms, zooplanktonic abundances in this
area down to SSOSare highest during the spring and
The shelf-break front is a major oceanographic feature early summer, decreasing noticeably during the win-
along the southern Southamerican Atlantic coast ter (Ramirez, 1981b; Ciechomski and Sanchez, 1983;
between ca. 37 and SooS; active vertical water mixing Fig. IS). Typically cold water zooplankters (e.g., the
along this boundary is responsible for enhanced pri- pteropod Limacina retroversa), however, may be
mary production (3S0-4S0 g C m-2 v": Negri, 1993), most abundant over the shelf and slope at other times
and the concomitant concentrations of zooplankton of the year, like in the autumn (Dadon, 1990a).
(Ciechomski and Sanchez, 1983; Dadon, 1984a) (Fig.
3, IS, 16). Interestingly, radio-tracked elephant seals It should be noted that the large-scale biogeographic
from Peninsula Valdes (ca. 43°S) have been observed divisions described and illustrated for the Argentine
to spend many months at a time precisely within this Sea (Fig. I) reflect only marginally the strong longi-
front, where the high concentration of animal biomass tudinal component derived from the influence of the
Pteropoda and
Decapoda larvae Chaetognatha Surface isotherms
35°
40·
45·
50·
."
A C 55°
70" 65" 60· 55· 50· 70· 65· 60· 55· 50· 70· 65· 60" 55" 50·
Fig. 16. Biogeographic zonations for the Southwestern Atlantic neritic area based on early stages (protozoeae and zoeae) of Decapoda larvae col-
lected in May 1978 through February 1982 (A), and on Pteropoda Euthecosomata and Chaetognatha (Autumn 1978) (B). C: Surface temperature
fields in January-March (data collected between 1958 and 1970). A: modified from Bacardit (1986); B: modified from Dadon (1989); C: simpli-
fied from Brandhorst and Castello (1971).
Malvinas (=Falkland) Current. Indeed, at the same lat- ly higher depths. Waters of subantarctic ongm are
itude inshore waters are usually warmer and host present at depths of ca. 300 m off the coasts of Brazil
warmer-water organisms than those farther offshore, (Emilsson, 1961; Matsuura, 1986); and frequent
while the slope and adjacent oceanic area are domi- records of subantarctic and even Antarctic plankters in
nated by the cold, subantarctic Malvinas (=Falkland) upwelling cells as far north as 23°N (E. Boltovskoy,
Current flow, where only subantarctic plankters 1970; Valentin et al., 1987; Brandini, 1990) strongly
dwell. Thus, detailed studies often show that boundar- suggest that these organisms have been advected from
ies between dissimilar assemblages run roughly para- higher latitudes at subsurface depths. Typically sub-
lell to the coast, rather than perpendicularly to it (Fig. antarctic benthic organisms are recorded year round
16A, B), which is clearly matched by the north-south off Buenos Aires and Uruguay at depths of 150 to
orientation of the isotherms (Fig. 16C). Thus, north- >200 m. E. Boltovskoy et al. (1996), for example,
ward dispersion of cold water species is favored by found that, between 30 and 60oS, along 55°W, at any
these advective longshore processes, which are fur- given latitude the proportions of the cold water plank-
ther enhanced by the concomitant deformation of the tonic foraminifer Globoquadrina pachyderm a (with
associated temperature fields. respect to the entire foraminiferal assemblage) are
noticeably higher at 30-50 m than higher up in the
Additional complications to these geographic patterns water column. This submersion and equatorward
arise when the vertical component is brought into the advection of cold water zooplanktonic species is pro-
picture. While up to around 40 S the Malvinas
0
bably a world-wide phenomenon with important bio-
(=Falkland) Current occupies the entire water-column, geographic and, especially, paleoecologic implications
further north it submerges partially under the warmer (Boltovskoy, 1988, 1994; see chapter "Radiolaria
Brazil Current and pogresses northward at increasing- Polycystina" in this volume).
GUin~u
dome
c:::::::;.
.~f()
NECC ~ o'~\ Congo
~ ~'()'i 1- River
GC Gulf of Guinea \.o~
¢:::= :::::::::: ~> ,.-.-.-.---.--'~
C3~
SEC SEUC
~
~ %51~ Poi e Noire
~ SECC
Angola
¢:==
SEC
dorrle I
-,
~ AC
Sf"C Cape Frio
/ South Atlantic
Subtropical Gyre
BenC
AC: Angola Current

\
AgC: Agulhas Current
BenC: Benguela Current ~ ~gC
Brae: Brazil Current ~ \\c CUffent
CC: Congo Current south ~t\an
GC: Guinea Current
GCUC: Gabon-Congo Undercurrent
NECC: North Equatorial Countercurrent
SEC: South Equatorial Current
SECC: South Equatorial Count
SEUC: South Equatorial Undercurrent
Fig. 17. Sketch of the main hydrographic features and geographic reference points in the eastern South Atlantic mentioned throughout the text.
The tropical and subtropical eastern South Atlantic There is quite a bit of evidence to suggest that neritic
zooplankton communities in this area are distinct from
By comparison with the western South Atlantic, dedi- oceanic ones (Dessier, 1981; Barange et al., 1992;
cated biogeographic studies of the eastern region are Gibbons et al., 1995) although biogeographic affin-
far and few between and mostly of a patchy and ities amongst them are less clear. Certainly there are
taxa-specific nature. For example, Thiede (1975), three assemblages along the African coastline asso-
Robson (1983), Boltovskoy et al. (1995), have exam- ciated with the Gulf of Guinea (Bainbridge, 1972;
ined the distribution of some shell- and skeleton-pro- Binet, 1993), the Benguela Current and the Agulhas
ducing microplankton in the tropical and subtropical Current, respectively (Fig. 1), and within each there
east Atlantic; Gibbons et al. (1995) have looked at the are some permanent and seasonal boundaries to
biogeography of euphausiids around southern Africa, "sub-communities" (see below). The vicinity of the
and Pakhomov et al. (1994a) have studied commu- African continent affects the hydrology and the cli-
nities in the region between the Subtropical mate (circulation, coastal upwellings, river run-oft)
Convergence and the Polar Front. However, fully syn- and imparts a strong seasonal signal over the shelf. At
optic studies of multi-taxa communities are largely the coastal station off Pointe-Noire (4°47'S; see Fig.
absent and therefore discussions of biogeography have 17), the mean seasonal variation of surface tempera-
to be chiefly deductive. ture reaches 15°C,while 150 nautical miles offshore it
'"-- -------~----- ------------------

is only 7°C, and 3°C at 50 m (Dessier, 1981). Except exist over the Namibian shelf, where coastal upwell-
in the equatorial belt, the seasonal variability becomes ing is quite permanent. While these divisions are
attenuated offshore. Oceanic provinces seem less largely artificial because of considerable interaction
clearly associated with water mass type, although between them (especially south of200S) they conform
there are obvious latitudinal, thermal associations with to the biogeochemical provinces of the region pro-
the Polar Front and the Subtropical Convergence. posed by Longhurst (1995; see Fig. 1). Having said
that, a lack of congruence between the biogeography
Unlike the western South Atlantic, where the South and biogeochemistry has been observed for the coast-
American continent essentially extends from the al domains off Africa (Gibbons, 1997b) but future
equator to the pole and strongly limits the movement studies will undoubtedly change this.
of water from the Pacific Ocean Basin into the
Atlantic, the eastern boundary is open to the Indian Eastern tropical South Atlantic
Ocean south of Africa. Thus, superimposed upon the
north-south provincial make of the region are patterns South of the equator, the oceanic areas of the Gulf of
induced by contact with Indian Ocean water. Given Guinea can be divided into zones on the basis of
the general west-east flow of water at higher latitudes, hydrology and currents (Dufour and Stretta, 1973a;
the effects of this contact would be negligible were it Stretta, 1975; Voituriez and Herbland, 1977; Herbland
not for the fact that Indian Ocean water is actually and Voituriez, 1977; Gallardo et al., 1974; Voituriez
"fed" into the South Atlantic via the fast flowing and Herbland, 1982). At the equatorial divergence,
Agulhas Current. Although strictly speaking, the between 1DN and 3°S, there is a pronounced seasonal-
Agulhas Current retroflects to the east around 38°S, ity in the structure of the water column (Voituriez and
15°E (Hutchings and Boyd, 1992; see Fig. 17), its Herbland, 1977). During the warm period (October-
water enters the region via eddies (which move to the June) a thermal ridge develops due to the eastward
south) and rings (which move lazily across the entire flowing Equatorial Undercurrent (Fig. 18) under the
region from the SE to the NW), and it also gets caught westward flowing South Equatorial Current.
up in the northward flowing frontal jet system of the
Benguela ecosystem. Because subtropical Agulhas This ridge effectively separates a warm, shallow,
Current waters enter the South Atlantic in the transi- nutrient-poor upper layer from a cooler, nutrient-rich
tion zone between the Southern Ocean and the sub- deeper layer. Chlorophyll levels (up to >0.4 mg ChI a
tropical South Atlantic, the transition zone itself is m-3) and primary production (70 mg C m-2 h-1) are
exaggerated, and communities there are far more spe- high and are strongly stratified as a result of the inter-
cies rich than those at lower latitudes. The northward play between light and nutrient limitation, and peak in
extent of the transition zone in the eastern South the region of the thermocline. During the cool period
Atlantic is likely to be far greater than that on the west (July-September) the flow of the South Equatorial
owing to the strong northward flow in both the coast- Current is accelerated and the Equatorial Undercurrent
al and oceanic areas. Indeed, Agulhas Current species becomes much shallower (Fig. 18). Consequently,
have been recorded off the Gabonese-Congolese- high vertical shear develops between the two currents
Angolan shelf (roughly 0 to 15°S; Ducret, 1968; prob- which results in upwelling and the thermocline breaks
ably caught up in the shelf edge jet), as well as in the surface. During this period, chlorophyll (>0.8 mg
waters off northern Brazil (Ponomareva, 1990). m-3) and primary production (up to 100 mg C m-2 h+)
are high, especially at the surface, but they may be no
For the sake of convenience we have divided the higher than during the warm period as a result of
coastal eastern Atlantic into two areas around the increased turbulence (Voituriez and Herbland, 1977).
Angola-Benguela front (at 16-200S, also called the
Cape Frio front, see Fig. 1, 17). North of the Angola- Zooplankton biomass tends to be low at the core of
Benguela front (Fig. 26 below), along the Gabonese the divergence zone (0.2 ml m-3, settled volume) but
and Congolese coast, the physical and biological pro- increases with the latitude as populations and commu-
cesses are markedly seasonal, as they are too in the nities mature (0.56 ml m-3) (Dufour and Stretta,
region of the southern Benguela south of Luderitz (ca. 1973a). The biomass of zooplankton in this region
25°S). By contrast, more or less perennial conditions may be over 2.5 times greater than that of the sur-
Latitude
()o 50S 150S 50N ()o 1()OS 150S
50
100
150
Temperature (oC)
O-r~------~--------~------~--------~
-r~------~~------~------~--------~
50
.c
.•...
Q.
Q)
100
o
150
O-ri------,~-.--~~--------~------~
/
I
I
I
50 o If I I
", :;
..•.
__ " I
, ,
I
100 \ I
\ ..J
150
Chi a (mg m-3)
Fig.lS. Vertical transect across the equator to 15°S; temperature, nitrate and chlorophyll a from the surface to 150 m in January (left) and June-July
(right). From Voituriez and Herbland (1977).
-
rounding area, and only 1.5 times less than that Phytoplankton and zooplankton biomasses are strong-
observed in the Mauritanian upwelling region (Le ly correlated over much of the equatorial south
Borgne, 1977). The vertical distributions of chloro- Atlantic during the warm period, both horizontally
phyll and mesozooplankton (200-500 m) are similar (Fig. 19) and vertically (Fig. 20) (Dufour and Stretta,
(Le Borgne, 1977). The depth of the thermocline 1973a; Dandonneau, 1975; Le Borgne, 1977). This
deepens to the south of the equatorial divergence. agreement tends to break down during the cold period
in the enrichment areas, but improves to either side.
There is a surface movement of the upwelled water to Zooplankton communities comprise a greater number
5 S, especially during the cold season, which is
0
of higher trophic levels both during the warm (than
reflected by fairly high phytoplankton biomass and cold) period and at distances from the divergence and
production at the surface (Herb land and Voituriez, enrichment zones. Such communities are typical of
1977; Voituriez and Herbland, 1977). This may be oceanic waters which are generally species rich,
weakly augmented to 8 S during the warm period by
0
trophically diverse and said to be balanced. Those of
the advection of nutrient rich waters of Benguela cur- the divergence and enrichment zones are unusual, in
rent origin from the east (Dufour and Stretta, 1973a). the sense that they tend to be relatively poor in species
Zooplankton biomass decreases from north to south (mostly herbivores), and are probably unbalanced
(0.39 ml m-3 settled volume) across this area. with their environment (Dufour and Stretta, 1973a;
Dandonneau, 1975; Le Borgne, 1977).
Between 9-13°S, the surface water flow is weak and of
variable direction, although this is the theoretical area Although there are obvious differences in the zoo-
of the South Equatorial Countercurrent. The thermo- plankton communities associated with waters inside
cline may occur at depths in excess of 70 m. It is a and outside of the equatorial divergence, evidence to
region of low phytoplankton biomass (chlorophyll a support faunal regions associated with the clear zones
between 0.25 and 0.50 mg m-3) and production (21 mg of biomass and production is ambiguous. Boltovskoy
C m-2 h+), which peak in the region of the thermo- et al. (1996) failed to identify distinct polycystine
cline in the warm season and is scattered in the upper radiolarian assemblages with any of the zones, and
mixed layer in the cold season (Fig. 18). Production is noted a relative homogeneity of the assemblage across
largely based on regenerated nutrients. Zooplankton the region between 10<N and 17°S irrespective of
biomass is low (0.2 ml m-3, settled volume) (Dufour temperature, latitude, salinity or chlorophyll a. They
and Stretta, 1973a; Voituriez and Herbland, 1977). did observe distinct vertical stratification, however,
and as their study was conducted during the warm
Between 13-16°S, in the southern branch of the South period, it is possible that different assemblages might
Equatorial Current (or Benguela Drift), the surface have become clear during the cold season.
water flow is generally to the NW. Temperatures are
generally low and the water has salinity characteris- On approaching the African continental land mass,
tics of Benguela (and coastal upwelled) waters mixed the South Equatorial Countercurrent and South
with Subtropical Convergence waters. Thus, phyto- Equatorial Underrcurrent are deflected southwards.
plankton biomass (chlorophyll a: 0.3 to 0.8 mg m-3) This polewards deflection induces an uplift (or dom-
and primary production (around 50 mg C m-2 h+) are ing) of the isotherms and results in the formation of
rather high. They are distributed throughout the mixed the Angola Dome centred on 100S-9°E (Mazeika,
layer. Zooplankton biomass is low in the core of this 1968; Gallardo et al., 1974). Although the dome is a
region (0.2 ml m-3) but increases to the north and permanent structure at depth, it only has an influence
south (0.3 ml m-3) (Dufour and Stretta, 1973a). on primary production when the thermocline and
nutrient rich waters are uplifted to the euphotic zone.
Between 16 and 23°S, in the Subtropical Convergence This is seasonal and occurs when the trade winds
area the nutrient rich layer is too deep (100 m) to fuel decrease and facilitate the eastwards flow of surface
high productivity. Chlorophyll a varies between 0.2 waters (Voituriez and Herbland, 1982). Gallardo et al.
and 0.8 mg m-2, and primary production is less than (1974) showed that the thermocline rises to 10 m in
13 mg C m-2 h-l (Dufour and Stretta, 1973a). February-March, and although it does not induce any
Planktonic biomass (rng ash-free dry weight m-2)
o July A
• October
2000
1000
r-r B
I \ 40
I I
2500 f \
.
C1l
I t "
1/ ' ,
I 30 •,
.. >-N
I
" •, 0
....cO
... " '.
I
2000 /ft... / 0
o...E
'4\
I
'
20 0
0
J 0 200
1500 10
0
0
o E
o
....c-
0
• U
1000
40 60 80 100 120 140 S

Latitude
Fig. 19. Horizontal distribution of zooplankton biomass at 3°N-15°S (A). Relationships between chlorophyll a and zooplankton at 3°N-ISoS
(B). From Le Borgne (1977).
o 2 3 4 5 Prim. prod. (mg C m-3 h+)

! ! ! ! !
0, Chi a (mg m-3)

0, 10 No. copepods per 10 I
!
o
!
10 !
20
!
N03 - N (fl atg 1-1)
o __ ----------~~----------~------------_.
I
i.,
"
•I'
----.. .--
........ ..:..~:::-
50
:.- ~.......-.........
.I
-.
,i •I
.II ~
• .•.....•
Prim. prod.
Chi a
_. _ No. cope pods
I
100 ~ .-_ ..• NO) - N
Fig. 20. Vertical distribution of primary production, chlorophyll a, copepod abundance, and nitrates at 0 0
, SoW. From Le Borgne (1977).
_....,.,-------------------------------------
surface cooling or nutrient enrichment, production on According to Berrit (1966) the warm, low salinity
the side of the structure seems slightly higher than in «35%0) surface waters which spread south along the
the surrounding oceanic waters. This contrasts with coast (Fig. 21) to Pointe-Noire (4°42'S) during the
the situation north of the equator, where the Guinea warm season originate in the Gulf of Guinea where
dome (which arises due to the northwards deflection they would be formed in the eastern part of the gulf.
of the North Equatorial Countercurrent, which is itself Berritt (1966) called them "Guinean waters", but for
a stronger feature of the superficial tropical circula- Wauthy (1977) these waters result from a mixture of
tion than the South Equatorial Countercurrent) is tropical waters with the outflow of the Congo river.
associated with highly elevated phytoplankton bio- Indeed, a low salinity, swiftly flowing (0.5 to 2 knots),
mass and production (Voituriez and Herbland, 1982). northward directed current (the Congo Current) is
The effect of the Angola Dome on zooplankton popu- present in the upper 10 m (Fig. 21), between the
lations, processes and communities is unknown, mouth of the Congo River (6 S) and Cape Lopez
0
although secondary production probably balances (1°S) along the coast and beyond the edge ofthe shelf.
phytoplanktonic growth.
Water temperatures across the region are generally
high during the warm season. They decrease to the
Coastal area between 0 and 20 S (chiefly Guinea
0
south owing to the cold waters of the Benguela Current
Current) which are advected NW. Near the coast, north of Cape
Frio (about 18°S), a thermal front separates the warm
Physical processes and seasonality Angolan waters from the recently upwelled Benguela
water (Angola-Benguela Front, see below). North of
The main surface currents in the region include the this front, the water column is strongly stratified over
northward flowing Benguela Current, and the west- most of the area and displays the characteristic typical
ward or north-westward flowing South Equatorial tropical structure. The thermocline is pronounced and
Current (Fig. 17). These currents are influenced by its lower point (18°C) can be found at between 30-100
seasonal fluctuations in the St. Helena anticyclone m depth. The upper mixed layer is generally poor in
which also cause alternating periods of upwelling and nutrients and in plankton, but seasonal shoalings of the
stability at the coast. thermocline induce periodic enrichment processes.
The basic seasonal pattern is a pronounced warm peri- During the austral winter the St. Helena anticyclone is
od between October and May, followed by a cold sea- reinforced and the SE winds extend along the coast to
son from June to September. Cape Lopez (1OS). Because the winds are parallel to
the coast, extensive coastal upwelling of waters from
During the warm season, the anticyclone is the weak- the Equatorial Undercurrent is observed in the region
est and furthest south (Peterson and Stramma, 1991; from Cape Lopez to Cape Frio. The acceleration of
Gouriou, 1993). The SW monsoon winds penetrate the surface current and the shallowing of the
the African continent and give rise to significant pre- Equatorial Undercurrent reinforce the shear and
cipitation: the Zaire (or Congo) River has the second increase the vertical mixing which leads to the pro-
largest discharge after the Amazon. As its basin strad- duction of a fairly deep upper mixed layer. Along the
dles the meteorological equator, differences in the coast, the surface circulation is directed N from Cape
rainy seasons between northern and southern tributa- Frio to Cape Lopez (Fig. 21). Due to the continuity of
ries determine a four season regime at its mouth. upwelling north and south of Cape Frio, the thermal
Highest outflow is observed in December and a sec- gradient at the Angola-Benguela Front weakens.
ondary maximum takes place in May, while minimum
discharges are observed in July-August, with a second A four season regime can be observed in the coastal
minimum in March. Its considerable discharge, about areas from Cape Lopez to Cape Frio: the Great Warm
45,000 m3 s-1 on average, fluctuates between 23,000 Season (January-May), the Great Cold Season (June-
and 80,000 m3 s-1 (Eisma and van Bennekom, 1978). September), the Little Warm Season (October-
Its plume is directed to the W or NW and can be November) and the Little Cold Season (December).
detected 1000 km offshore (Piton, 1988). The minor upwelling season that takes place in
Distance offshore (nautical miles)

a
!
10 20 30 75 100 150 200
! ! ! ~~--~! ----~!----~--~!----~--~!
wsw
100 " A"""""""""""":'"\",,C,,"'::"'"'

. r-; >.~
~NNWflow
200 D SSEflow
Great Warm Season

---E 300
--
s:
+-'
a
Q.
a;
0
100
200
D
Great Cold Season
300
Fig. 21. Meridional transport off Pointe-Noire (4°50'S), from the coast to 200 miles offshore. Hatched: north-north-westwards, open: south-
south-eastwards. Isolines: velocities from 10 to 10 em s-l. From Guillerm (1975).
December is possibly a result of the secondary maxi- ficial layer of warm water with low salinity which is
mum observed at this time of year in the velocity of periodically pushed away by upliftment of the ther-
the South Equatorial Current. This Minor Cold mocline and upwelling.
Season, which is highly variable in intensity, is not
detectable offshore and decreases in importance in the The four seasons cycle
northern part of the area (near the seasonal front
at Cape Lopez). The Little Warm Season is less During the warm season, the water column is strati-
important in the south (near the Cape Frio front). fied and the thermocline is relatively deep (the 18°C
Offshore, the four seasons are replaced by a two isotherm is between 50 and 120 m off Pointe-Noire,
seasons cycle. Dessier, 1979). As a consequence, nutrient concentra-
tions (phosphate 0.3 atg 1-1), phytoplankton biomass
(0.5 mg m-3) and production (35 mg C m-2 h+) can be
Coastal area between Cape Lopez (1°S) and Cape low and are generally situated in the region of the
Frio (18°S) thermocline (Fig. 22).
The area, delimited to the north and the south by sea- The low phytoplankton biomass observed during the
sonal thermohaline fronts, is characterized by a super- warm period is reflected by low zooplankton biomass
Temperature at 17 m A 0.9 Phosphate at 5 m c

c:
...!.... 0.8
o
s.
v•...
::::l
.•...
r1S
•...
V
o,
E 16
~ J F M A M J J A SON 0
J F M A M J J A SON 0
Chlorophyll a B Primary production D

3 ~100
M
I 2
--'= 90
E ~ 80
00
E 70
~E U
00 60
E 50
~ 40
30~~~~~~~~~~~~~~
J F M A M J J A SON 0 J A SON 0
Month
7r---------------------------,
Zooplankton Month
~ 6 biomass
E
N
I
E 5
~
1:bO 4
'w 3
~
C2
c
Fig. 22. Coastal station off Pointe-Noire (4°50 S). Changes in 17 m temperature, 5 m phosphate and chlorophyll, primary production and zoo-
plankton dry weight, over a year. From Dessier (1981).
(dry weight: 1 g m-2, displacement volume <0.1 ml py the whole column. On average, nutrient concentra-
m-3), across the shelf (Fig. 22 and 23), although this tions are high (phosphate: 0.7 atg 1-1,Fig. 22). This in
may be elevated when the thermocline shoals or turn is reflected by a seasonal peak in coastal chloro-
(locally) in response to river runoff (Dessier, 1981). phyll a (2 mg m-3) and phytoplankton production (75
mg C m-2 h-I) at 5 m depth. Pigment diversity
At the end of May, the thermocline shoals and breaks (D4301D665) decreases from March to August indi-
the surface. Subsurface cold waters with high salinity cating that phytoplankton populations are the young-
(from the Equatorial Undercurrent) replace the tropi- est when the nutrient input is the largest (Dessier,
cal warm waters with low salinity. A first peak in 1981). Zooplankton dry weight (2.5 g m-2) and dis-
nutrients and phytoplankton occurs in June. After a placement volume (0.25 ml m-3) also tend to be high-
decrease in July, a second (and greater) maximum is er during the cold period across the shelf (Fig. 23). At
reached during August, when Benguela waters occu- the start of the cold season a marked increase in zoo-
______________________________________________ i.,:3.i1.':~~r~~ .•,~~;.d
.•
:ioI .':".t
Great Cold Season

August September
Zooplankton
volume
00 (ml 1000 m-3)
c=J <100
18~fJ~J100-200
GABON
•• 200-300
_ >300
CONGO
90 120 120
Fig. 23. Distribution of zooplankton displacement volumes across the shelf from Gabon to Cabinda. From Dessier (1981).
plankton biomass to the edge of the shelf is related to Zooplankton communities

a maximum in divergence and upwelling there (as
indicated by the high salinities), and to the develop- Dessier (1981) identified a number of distinct zoo-
ment of two large copepods species. plankton communities over the Gabonese-Congolese-
Angolan shelf from studies conducted over several
Cape Lopez thermohaline front years. These communities were generally associated
with either different temperatures or hydrological
At the beginning of the cold season, a very strong lat- regimes, and their presence in the region varied on a
itudinal front develops in the region of Cape Lopez seasonal basis. In essence, the region is characterized
(I°S) which separates the warm, less saline Guinean by two quite different communities associated with
waters from colder, high salinity waters to the south. either oceanic or neritic waters. The warm oceanic
This front joins the northern boundary of the equatori- group is persistently present north of Cape Lopez and,
al divergence (Wauthy, 1977) and during the cold sea- during the warm season, across the region beyond the
son confines thermophilic species, such as the copep- 100 m isobath. During summer it encroaches onto the
od Undinula vulgaris, to the Guinean waters north of shelf, south of Cape Lopez.
Cape Lopez. This front (around 24°C at the surface)
undulates and moves a little northwards or southwards The structure of frontal neritic communities is more
in June and July, before vanishing and letting Guinean complex and reflects their diverse origin and the sea-
waters spread southwards during the warm seasons. sonal variability of their environment. The composi-
When strong, this front is associated with a conver- tion of these communities varies with changes in
gence, and chlorophyll and zooplankton biomass hydrology and latitude, and seems to accurately
enhancements on its southern, cold side (Fig. 24) reflect warming and cooling events (Dessier, 1981).
(Dufour and Stretta, 1973b). For example, austral summer assemblages of the inner
shelf south of Cape Lopez may be supplemented by
2000'S 1030' 1000' 0030' 0000' 0030'N

Stations 20 ;1 22 ~3 24 ' 25 ~1'~~~ 28 I ~O
30 F
10~~~~2~1~2~2~~~~~~~~~~~~~2~4~'~;;;:~~::::::::~~
1--_-18
19
50
70
90
17- __ -
Temperature (OC)
10
_ 34.5---...:,34.0
30 -~---------
' _-- -----
50
70
,.. - ......
.•....
.....•.
- - - -- - - - - - - ---
90
--------
35.5-
10
"23.0 ~22.5
.c 30
+-'
0- 50
Q)
70
o 90
0.25
.75'-- -----~0.75
0.50--=.
----0.25
70
90
Chi a (mg m-3)
20 5_
10 __
____
--5-
70
90
Zooplankton (ind. m-3)
Fig. 24. Vertical transect across the thermohaline front (hatched) off Cape Lopez in June. From Dufour and Stretta (1973b).
those of Guinean waters (tropical waters with low During the cold season, the neritic communities can be
salinity), owing to the southwards movement of shelf divided up into those of the inner (depth <50 m) or
water then. These warm waters push the winter popu- outer shelf region. Upwelling mainly occurs at the
lations offshore and neritic communities are found in shelf break, and as the shelf is 25 nautical miles wide
the middle of the shelf or southwards. Highest bio- inner and outer communities can develop simultane-
masses are situated near the coast (Fig. 23), due in ously, aided by probable coastal divergence. Although
part to the regression of the outer shelf species and the both regions obviously share a number of species, the
development of inner shelf communities dominated numerical structure of the two neritic communities dif-
, by asexually reproducing cladocerans and thaliaceans. fers markedly. The winter communities may be aug-
mented either by offshore subsurface species which tion (8 mg C m-2 h+; Cadee, 1978). River phytoplank-
have been advected into the region by water of ton die rapidly on arrival at the sea and owing to both
Equatorial Undercurrent origin during June, or by high turbidity and the rapid change in salinity there is
coastal and upwelling species from the Benguela an inhibition of the growth of brackish water species.
Current in August due to the northwards movement of This results in a general decrease in chlorophyll bio-
the surface current then. The most thermophilic super- mass and production with increasing salinity to 20%0.
ficial species are driven offshore and north of the front Thereafter, an increase in phytoplankton biomass is
at Cape Lopez. Other less thermophilic species are observed at the edges of the plume to 30%0, after which
found in abundance just south of the front. On the outer nutrients become depleted (or imbalanced) and phyto-
shelf, they are mainly replaced by subsurface species plankton biomass and production become depressed
from off the shelf break, of which only Calanoides (Cadee, 1978). The arrival of Congo waters off
carinatus and Eucalanus monachus become abundant Pointe-Noire (4°50'S) is always marked by a drop in
owing to their particular maintenance strategies (Binet, phosphates, chlorophyll and production. Only silica
1979; Petit and Courties, 1976). A neritic winter com- enrichment persists (Dessier, 1981).
munity develops inshore and Paracalanus scotti and
Oithona nana overwhelm the plankton numbers. There are few studies of zooplankton in the vicinity of
the Congo River mouth. Dessier (1981) reported high
The influence of the Congo river planktonic (both phyto- and zooplankton) concentra-
tions between 4 and 60S during May, which may have
The generally positive effects of river run-off on phy- been related to the high runoff. He also reported high
toplankton biomass and production should be expect- densities of opportunistic zooplankton species (the
ed during the Great Warm Season, which is also the copepods Temora turbinata and Centropages chier-
period of maximum run-off. While some of the small- chiae, the cladoceran Penilia avirostris) over the shelf
er rivers (Ogoue, Kouilou) have a clear localized between 4 and 13°S in November and low numbers of
influence (Dessier, 1981), the beneficial effects of the these species in March. These too could be related to
Congo river plume on marine coastal plankton are not periods of high and low runoff, respectively. The
evident and contradictory observations are probably plankton content of these low salinity waters probably
related to sampling at different states of mixing depends on the time elapsed since the mixing between
between riverine and sea waters. The huge amount of continental and marine waters, and on the nutrients
water discharged overwhelm the region with approxi- remaining after the first blooms.
mately 45,000 m3 s-1 (Eisma and van Bennekom,
1978). Unlike the Amazon (see above), the Congo has The Cape Frio thermohaline front
a deep canyon at its mouth which canalizes its very
fast flow. The canyon's depth, the low tidal range and South of the area, in the vicinity of Cape Frio, the
high current velocities in the river lead to an extreme intensity of the southward flowing Angola Current is
case of stratified estuary in which the residence time seasonal and its convergent boundary with the
of the water in the mixed surface is very short. The Benguela Current fluctuates between 16-20 S (some-
0
influence of freshwater is thus confmed to a thin sur- times to 22°S) (Dias, 1983; Shannon and Agenbag,
face layer (10 m at the mouth and 30 m at 700 km) and 1987; Shannon et al., 1987; Fig 26). The more north-
the dilution of river waters in the sea takes place very erly position of the front occurs during the cold period
far from the mouth (Fig. 25). when upwelling over the Namibian shelf is strongest
and northward advection greatest. By contrast, it is fur-
Close to the river mouth, there is an increase in nitrate thest south during the warm period when southerly
and phosphate which Van Bennekom et al. (1978) winds are weakest and the South Atlantic Anticyclone
attribute to upwelling induced by estuarine circulation is furthest south. Thermal and salinity fronts between
(nitrate) and desorption from suspended matter (phos- these two systems do not coincide because the maxi-
phate). Silica concentrations decrease linearly with mum of salinity is not in the cold waters upwelled
salinity, which suggests weak utilization by diatoms. from the Benguela Current, but in the warm waters
The core of the plume is associated with surprisingly from the Equatorial Undercurrent, South Equatorial
low chlorophyll (1 mg m-3) and poor primary produc- Countercurrent and South Equatorial Undercurrent.
November 1976
So
..
70 Salinity (0/00)
/
\
(f.
• 200 m
26
So
.. . ....
.... II.., ~ :"
'~;3
,,~..~
70 Temperature
/: (...: "\::
(aC) •
. \
.\
So
x 0.2
j' 200m
\
i
90 100
• •
405
So •
60
•
70 Chi a (rng rrr-')
...•..•......
405
So
60
70
60 70 80 90 100 110 120E
Fig. 25. Zaire (or Congo) River plume: distribution of salinity, temperature, nitrate, surface chlorophyll a, and primary production in
November and April-May. From Cadee (1978).
...,&.,.•..•••••';iik.1!)G h••,,:;;;
------------------------------------------------------------------------------------------------------
Fig. 26. Changes in the SST in the region of the Angola-Benguela Front throughout the year. From Shannon et al. (1987).
There is a strong suggestion that two thermal fronts is observed north of 16°S throughout the year.
may exist in the region at times, one associated with Stratification is weaker to the south (especially during
Benguela upwelling and the other with a tongue of sur- the cold season) and the mixed layer can be fairly
face tropical and equatorial water: the Angola Tongue. deep. The front may be pushed offshore in northern
Namibia during the warm period by coastal upwell-
The Angola-Benguela Front (Fig. 26) is dynamic and ing. During the cold season upwelling occurs all along
can display quite rapid changes in latitudinal position the coast and the thermal gradient is weaker.
in response to changes in wind stress (Shannon et al.,
1987). The front is most clearly observed in the sur- Zooplankton communities to either side of the
face waters and strong, shallow thermal stratification Benguela-Angola Front are very different (Olivar,
General biological features of the .South Atlantic 35
1987;MacPherson, 1991;Barange et al., 1992). Those heat, salt and Indo-Pacific fauna into the South
to the north comprise many species with tropical affin- Atlantic. Occasionally, an Agulhas ring will move
ities which occur at low abundance, whilst those to northwards close to the shelf, entraining Indian Ocean
south comprise fewer species with coastal and cold water into the South Atlantic and also drawing neritic
water affinities, at high abundance. Southward move- water offshore into the subtropical gyre.
ment of Angola Current water can occur along the
shelf edge, favored by an abattment of the coastal Phytoplankton
Namibian upwelling. Such an unusual intrusion is
probably more frequent during the so called Phytoplankton biomass and productivity in the
"Benguela Nifios". It is marked by an inflow of gelat- Benguela region are enhanced by the wind-driven
inous carnivorous species, siphonophores and hydro- upwelling over the shelf. Brown et al. (1991) and
medusae into Namibian waters. The number of spe- Brown (1992) have described the distribution and pro-
cies in both groups decrease southward and the ductivity of phytoplankton on the west coast, while
siphonophores exhibit a weak inshore-offshore gra- Probyn et al. (1994) describe primary production and
dient (Pages and Gili, 1991). This illustrates the fact related processes on the Agulhas Bank. Highest chlo-
that fronts are boundaries between ecosystems of dif- rophyll a concentrations (5-10 mg m-3) occur close to
ferent trophic structure. The cold side is the lower the coast, associated with upwelling centres, with
level with high productivity and herbivores, whereas declining concentrations to less than 1 mg m-3 off-
the warm side is characterized by planktonic predators shore in the southern Benguela, but still greater than 2
such as cnidarians (north of Cape Frio), or even tunas mg m-3 beyond the 200 m isobath off Namibia, indi-
(north of Cape Lopez; Dufour and Stretta, 1973b). cating a wider productive zone in the northern
Benguela. Diatoms and dinoflagellates dominate the
biomass near upwelling centres, while small flagel-
Coastal area between 20 and 34°S (Benguela lates and other nanoplanktonic organisms dominate in
Current) aged upwelled waters. Clear successional changes
occur as upwelled waters stabilize, mix with sur-
Physical processes and seasonality rounding waters and mature (Fig. 27) (Mitchell-Innes
and Walker, 1991; Pitcher, 1988; Pitcher et al., 1991).
Wind-driven upwelling is the dominant forcing mech-
anism determining the physical characteristics of the Red tides are common throughout the Benguela
Benguela shelf region. Strong southerly winds blow Current and have caused numerous mortalities in shel-
permanently in the Luderitz area (25°S), which, com- tered bays when aggregations of toxic species develop
bined with the narrow shelf, make it probably the or when oxygen is depleted during the decay of the
most active upwelling centre in the world. Southerly bloom. In the southern Benguela there are distinct
winds become more seasonal to the north and south, seasonal patterns in the development of dinoflagellate
with peak winds in the austral winter in the northern populations, with dense aggregations occurring when-
Benguela, and in the austral summer in the southern ever physical conditions are favorable (Pitcher et aI.,
parts. The shelf is relatively wide and deep, except in 1995; Pitcher and Boyd, 1996). The affect of these
the extreme north and south and off Luderitz. A num- red-tides on zooplankton communities is unknown.
ber of active upwelling sites exist along the coast,
where upwelling is pulsed in response to changes in Primary production measurements are relatively rare in
coastal winds and the passage of coastally trapped the Benguela region, being limited to a small region
waves. The northern and southern boundaries are around the SW Cape where more than 10 measure-
characterized by intensive mixing with warm water of ments have been made within a limited area.
tropical origin, and shifts in the boundary conditions Elsewhere only 1-2 measurements have been docu-
can cause major perturbations to the productivity of mented in the literature, and are certainly inadequate to
the fish stocks (Shannon et al., 1992). The offshore describe seasonal, areal and interannual variations in
boundary is a complex series of jet currents, eddies primary productivity in the Benguela region. The
and frontal features such as filaments. Agulhas rings measurements that have been made indicate that the
break off from the retroflection area and transport southern Benguela is more productive (2.0 g C m-2d-1)
10
20
30
40
10
20
30
40
---E
-S
10
20
0-
~ 30
40
x: • ·\~__---.--.. f.~·~~· dinoflagellates

10 xl· •• .----_. & & 6 6
small • .•. .•. .• .• ""... small d
20 diatoms .•. ....
•......, flagellates
30
Coscinodiscus
gigas . "-, 6
... ................••
6
... ""''''
40 Dominant phytoplanktonic groups
30
40 Primary production (mg C m-3 h-l)
20 22 24 26 28 30 3 5 7 9 11 13 15
March April
Dates
Fig. 27. Changes in several parameters at a fixed location (32°33.2'S, 18°05.2'E) in the southern Benguela over two consecutive upwelling
events. From Mitchell-Innes and Walker (1991).
than the northern Benguela (1.2 g C m-2 d+), despite may yield different results. Seasonal changes in pro-
the higher mean biomass of phytoplankton in the north ductivity were reported by Brown et al. (1991) for a
(4.8 vs 2.2 mg chlorophyll a m-3, respectively). On the very limited area in the southern Benguela. Nitrogen
south coast, east of Cape Point, biomass is lower (1.5 uptake expe~nts (Probyn 1985, 1987, 1992) indi-
mg chlorophyll a m-3), and productivity is intermediate cate that mvch of the production on the continental
(1.8 g C m-2 d+), compared with the west coast. The shelf, downstream from the upwelling centres, is
high biomass, long turnover time and low productivity fuelled by recycled nutrients from the microbial food-
off Namibia is somewhat surprising, but as these data web, rather than by "new" nutrients advected from
are based on very few measurements future research elsewhere. The high organic content of the shelf sedi-
:;.I~/iiliiriliil·~_"'·'!!1,= --------.------ ...---

ments indicates considerable deposition of carbon. Part

of the organic matter is recycled into the upwelling
source water and part is sequestered in the sediments,
resulting in the Benguela system being both a source
and a sink for carbon dioxide (Hutchings et al., 1995).
Grazing control of the initial blooms in newly
upwelled water is relatively poor in the southern
20<>
Benguela, due in part to the poor spatial and temporal
matching between phytoplankton and mesozooplank-
ton in the strongly pulsed upwelling regime (Verheye,
1991; Hutchings et al., 1995).
Zooplankton Processes
Zooplankton studies in the Benguela Current have

largely focused on the regions of the most intense fish- Copepods
ing operations in the Southwestern Cape (33-35°S), (103 indo m-3)
and in the vicinity of Walvis Bay (23°S). Within these CJ <10

areas research has focused on the distribution and sea- ~ 10-30
sonal abundance of particular taxonomic groups (e.g., ~::::::::::I 30-100
euphausiids, copepods, cladocerans, tunicates, chae- ~ 100-300
tognaths), followed by more process-orientated and _>300
quantitative studies of life history strategies, biomass
fluctuations and productivity estimates of the domi-
nant species of copepods and euphausiids, which are
the most important components relevant to fisheries.
Fig. 28. Distribution of copepod biomass in the northern Benguela
in April 1986. From Olivar and Barange (1990).
Hutchings et al. (1991) described the biomass and pro-
duction of zooplankton in the Benguela region, based
on the limited measurements available. Highest bio-
mass occurrs downstream from major upwelling sites
Zooplanktonic
both off Namibia (23-25°S; Fig. 28) and off the SW dry mass
Cape coast (32-33°S; Fig. 29), on the inner to midshelf (g m-2)
region. The mean biomass of zooplankton was esti- D<0.5

mated to be quite similar in the northern and southern ~0.5-1.0
Benguela, approximately 0.8 to 1.0 g C m-2 in the ~ 1.0-1.5
330 ->1.5
upper 200 m. However, Timonin et al. (1992) conclud-
ed that moderate to high concentrations of deep-living
Calanoides carinatus live at depths of 300-700 m, and
total biomass in the offshore zone is therefore underes-
timated. In the extreme south and north, where warm
water boundaries occur, biomass is reduced. Few sea-
sonal studies are available; Andrews and Hutchings
(1980) show a distinct but erratic seasonal cycle in
zooplankton biomass in the southern Benguela at
34°S, with highest biomass occurring during the sum-
mer upwelling season (Fig. 30). Seasonality is reduced 170 180 200
slightly to the north (Pillar, 1987), while off Namibia
Kollmer (1963) showed little seasonal variation in Fig. 29. Distribution of mean monthly standing stock of zooplank-
zooplankton biomass in a limited area close to Walvis ton in the southern Benguela in 1977-1978. From Pillar (1987).
~---------------------.-----.---- -
5-.----------------------------------------------------,
N
~
E 4
1:
DO
3-month
.~ running
~ 3 mean
~
"eo
J1 2
erei
Q.
8
N 1
ONDJ FMAMJ J ASONDJ FMAMJ J AS ONDJ FM

1970 1971 1972 1973
Fig. 30. Seasonal variations in standing stock of zooplankton along a transect from approx. 34"S, 17°E to 33"27'S, I ?D15 'E across the shelf in
the southern Benguela. From Andrews and Hutchings (1980).
Bay. Elsewhere in the Benguela region zooplankton tion, phytoplankton community structure and cell size
surveys have been limited to irregular sampling on to be related to zooplankton grazing and production
a seasonal basis, but usually with comparisons of (Peterson et aI., 1990; Walker and Peterson, 1991).
"active upwelling" and "quiescent" conditions (e.g.,
Timonin et al., 1992). Zooplankton Communities
The adaptations of the dominant copepod Calanoides As with the coastal regions to the north of 20 S, there 0
carinatus and the euphausiid Euphausia lucens to the is a very pronounced frontal boundary between neritic
shelf circulation in the southern Benguela have been and oceanic communities that approximates the edge
studied by Verheye et al. (1991, 1992) and Pillar of the continental shelf (Barange et aI., 1992).
(1987), Pillar and Stuart (1988), and Pillar et al. Communities on the shelf across the region are char-
(1992). Similar studies of the dominant euphausiids acterized by low diversity and high abundance and
Nyctiphanes capensis and Euphausia hanseni were contain species which are typical of upwelling areas or
conducted off Namibia by Barange and Pillar (1992). cold-temperate shelf waters world-wide (Gibbons and
Laboratory rearing and shipboard measurements of Hutchings, 1996). These include Calanoides carina-
grazing, egg production and moulting rates have been tus, Euphausia lucens, Sagitta tasmanica, Muggiaea
made for a number of copepod species and Euphausia atlantica and Themisto gaudichaudii. Although the
lucens in the southern Benguela (Stuart and Pillar, frontal region is generally composed of a mixture of
1988, 1990; Borchers and Hutchings, 1986; Attwood neritic and oceanic species, it may contain species
and Peterson, 1989; Peterson and Painting, 1990). A which reach peak abundances here, such as Euphausia
30-day time series at a fixed location in St. Helena recurva (Pillar et al., 1991; Barange, 1990). The fron-
Bay in the southern Benguela (Bailey and Chapman, tal boundary may be intensified during the upwelling
1991) allowed the time scales relating to pulsed season (austral summer) in the south when Agulhas
upwelling to be linked through the phytoplankton suc- Bank and Agulhas Current water gets entrained in the
cessional sequences (Mitchell-Innes and Walker, northward flowing shelf-edge jet. Although perennial,
1991; Pitcher et al., 1991) to the responses and beha- . the front may also become blurred at times, especially
viour of the dominant copepods (Armstrong et al., in areas where the shelf is narrow (e.g., off capes
1991; Verheye, 1991). Transect studies across the Columbine and Point), and where as a consequence
southern Benguela shelf sampled water of differing frontal and oceanic water is advected shoreward dur-
ages after upwelling have allowed primary producing wind reversals. Under these circumstances, ocea-
~
!
I
::::. 4
nic species may be introduced into the coastal area. shore system (they have maintenance strategies which
While shelf communities south of Cape Columbine would favour this) but they could also be continullay
contain many of the characteristic neritic species of lost to entrainment in the South Atlantic gyre in the
the region, they are extremely labile, and changes in north and replenished from the south. But whatever the
their composition reflect the dynamic oceanography case, these species. which occur at low densities outside
of this region. Although species richness in this area the Benguela ecosystem, thrive within it.
can be extremely high (De Decker, 1984; Gibbons and
Hutchings, 1996), the communities are largely struc-
tured by physical processes and the region (to East Africa coastal
Hondeklip Bay, ca. 31°S) must be considered a transi-
tion zone. Just as the southern boundary to the Physical processes
Benguela ecosystem lies inside the physically defined
Benguela region (Gibbons et al., 1995), so too does The east coast of South Africa is bounded by the fast,
the northern boundary with the Angola Current southward flowing Agulhas Current which is com-
(Barange et al., 1992). Both are essentially described posed of warm, equatorial Indian Ocean water. The
by the extremes of advection of Agulhas Current and inner edge of the Agulhas Current tracks the edge of
Angola Current water respectively. Shelf communities the continental shelf, which is close inshore north of
to the north of Cape Columbine (to the Angola- East London, before moving offshore and westward
Benguela front) generally display a homogeneity in along the edge of the Agulhas Bank to retroflect in the
species composition, although cross shelf gradients region of 16 to 200E (Lutjeharms and van Ballegooyen,
that reflect mixing or community responses to the age- 1988). The Agulhas Bank is a roughly triangular exten-
ing of upwelled water are observed (Gibbons and sion of the continental shelf off southernmost South
Hutchings, 1996). There are, however, subtle differ- Africa. The western part of the region (West of Cape
ences in the numeric composition of zooplankton Agulhas) is considered to be part of the Benguela eco-
communities to either side of the perennial upwelling system (Shannon and Nelson, 1996) while the eastern
cell at Luderitz (Barange et al., 1992; Gibbons et al., part is directly affected by the Agulhas Current (Boyd
1995). It is clear that while the upwelling cell may not and Shillington, 1994) (Fig. 31). The structure of the
provide a permanent barrier to the longshore transport interior of the Bank is strongly influenced by wind
of holozooplankton from south to north (but see forcing and is deeply stratified in summer but well
Emmanuel et al., 1992 for intertidal communities) mixed in winter (Probyn et al., 1994). Easterly winds
there are differences in the seasonality. of upwelling in summer result in upwelling at the Capes and head-
(Shannon, 1985), production and biomass (Brown et lands along the coast and a cool water ridge may devel-
al., 1991; Pitcher et al., 1992), as well as differences op across the shelf from the eastern to the central
in the circulation of water across the shelf systems Agulhas Bank (Boyd and Shillington, 1994). This
(Barange and Pillar, 1992) that are reflected by the ridge serves to prevent the penetration of deep Agulhas
zooplankton communities. Current features onto the inner shelf. Conditions over
the outer region of the Agulhas Bank are forced by
Of particular interest here is the presence in the region oceanic processes and an upliftment of the isotherms
of species which also occur in the transitional, south may be induced by the Agulhas Current at the shelf
subtropical convergence (Fig. 1), such as Euphausia edge (east of Cape Agulhas) which can result in a per-
lucens and Sagitta tasmanica. These species are manently stratified water column there. Patterns of
thought of as characteristic zooplankters in the water flow at the edge of the Agulhas Bank are gene-
Benguela upwelling region and dominate their respec- rally W and SW to the east of Cape Agulhas and NW to
tive assemblages. It is thought that these species seed the west of Cape Agulhas. Water flow over the central
into the system from the south, presumably from South Bank is sluggish and of variable direction for much of
Atlantic Central Water, and undergoe population explo- the year but there is pronounced eastward flow inshore
sions in the productive waters of the South African west along the coast. The Agulhas Current penetrates onto
coast (Gibbons and Hutchings, 1996). The eventual fate the Agulhas Bank directly only in the region of the
of these populations is unknown. It is possible that Agulhas Bight (especially during summer), and this
these populations are self-perpetuating within the near- warm water may intensify the seasonal stratification of
Fig. 31. Map showing the different sectors of the Agulhas Bank. Broken lines represent the approximate depth of the upper mixed layer during
summer. Shading indicates the area where the most intense thermoclines are likely to occur over the bank. From Probyn et al. (1994).
the water column. It also serves to seed shelf commu- Phytoplankton

nities with Agulhas Current species. Filaments and
cyclonic eddies develop from instabilities along the Our understanding of phytoplankton communities and
northern boundary of the Agulhas Current while rings processes in the oligotrophic Agulhas Current is very
may form in the region of the retroflection. The rings limited. Rates of primary production are low (99 mg C
are deep (up to 1500 m), broad (up to 300 km diame- m-2 h+) by comparison with those on the Agulhas
ter) features which are generally situated well offshore Bank itself (Probyn et al., 1994) and biomass is con-
and whose only influence on coastal communities as centrated at the thermocline. The influence of shelf-
they move off into the south Atlantic is thought to be edge upwelling on production and biomass is of great-
surface entrainment (Duncombe Rae et al., 1992). est significance over the eastern Agulhas Bank where
Although it is possible that central water of Indian it results in a doming of the isotherms over the outer
Ocean origin (it is physically similar to Atlantic central shelf and leads to high subsurface chlorophyll maxima.
water) may get upwelled on to the west coast shelf du-
ring summer from the Agulhas rings, this is unknown. Chlorophyll biomass and production over the Agulhas
Filaments of Agulhas Current water are much shallow- Bank is strongly linked to the structure of the water col-
er (50 m) than rings and they may penetrate to Cape umn, the depth and intensity of the thermocline and to
Columbine in the southern Benguela ecosystem. They the forcing processes. Wind mixing in winter results in
occur just beyond the shelf edge and if entrained in the low production over the whole Agulhas Bank and chlo-
northward moving shelf-edge jet serve to intensify the rophyll biomass is distributed throughout the upper
frontal zone. They may be advected shoreward onto the mixed layer (generally greater than 70 m but between
Benguela shelf during wind reversals. 30-40 m in the east) (Fig. 32D). On the western
~Ii__~t;~_~. 1
Temperature (OC)
I
10
0
\
\
\
,,
\
20
,
-E~
...>=
Q..
40
,, Temp.
(])
0 60
80
100
A C D
0 2 4 6 8 10 0 2 4 6 8 10 0 2 4 6 8 10 0 2 4 6 8 10
Chlorophyll (rng m-3)

I I I I I I I I I I I I I
0 10 20 0 10 20 0 10 20 0 10 20
Nitrates (rnrnol rn-')
Fig. 32. Examples ofN03' chlorophyll and temperature profiles for sununer (A-C) and winter(D) on the Agulhas Bank. A: western Bank mid-
shelf; B: eastern Bank mid-shelf; C, D: western Bank inshore. From Probyn et al. (1994).
Agulhas Bank, the thermoclines may be very deep in ring summer) while those of the eastern area are char-
winter because of large amplitude internal waves and acterised by coastal communities dominated by
tidal stirring (Largier and Swart, 1987). As the water Calanus agulhensis (Verheye et al., 1994).
column begins to stabilise in spring surface blooms Communities over the Agulhas Bank are distinct from
develop which gradually deepen and decrease with those of the Agulhas Current (e.g. De Decker, 1984)
stratification during summer. Chlorophyll biomass is and tend to comprise typically coastal species such as
generally shallower and higher in the east than the west small calanoids and cladocerans. The communities in
(Probyn et al., 1994) because of the influence of the the Agulhas Current are typically tropical in structure
cold ridge and the strong, shallow thermocline induced and contain a preponderance of carnivores. The com-
by the shelf-edge upwelling there (Fig. 32A, B). mon herbivores are those which are capable of survi-
ving in oligotrophic waters and which feed primarily
Both phytoplankton biomass and production are high- on small cells. Their persistance outside these regions
er inshore (Fig. 32C) and downstream from the upwell- is thus unlikely given the altered structure of the food
ing foci (downwind of the prominent capes) than else- webs elsewhere. Having said that, the Agulhas
where on the Agulhas Bank and are lowest in the Current may seed the shelf with oceanic species such
central regions where water circulation is sluggish. The as Thalia democratica during summer and these can
phytoplankton communities influenced by coastal develop to dominate assemblages over the central
upwelling are dominated by large cells (as in the Bank where circulation is sluggish and waters oligo-
Benguela system) but these are thought to give way to trophic (e.g. Gibbons, 1997a).
smaller cells elsewhere. Those of the Agulhas Current
are likely to be dominated by small cells and are thus Zooplankton biomass is generally greater over the
unlikely to contribute much in either the way of bio- eastern Agulhas Bank (3.07 g dry wt m-2) than the
mass or production to the west coast (via filaments). western Agulhas Bank (1.21 g dry wt m-2), though
whether this is due to differences in primary produc-
Zooplankton processes and communities tion associated with the cold ridge or to differences in
fish predation is largely unknown. Fluctuations in
Zooplankton communities over the western Agulhas zooplankton biomass over the western Agulhas Bank
Bank have strong Benguela affinities (especially du- are strongly linked to fluctuations in phytoplankton
standing stock and coastal upwelling (Hutchings and reflect entrainment in the northward flowing shelf-
Nelson, 1985). Although zooplankton communities edge current. It would also suggest that conditions at
-over the Agulhas Bank may consume in excess of the front must be suitable for their continued survival,
50% of daily primary production, the food chain effi- if not reproduction. Certainly feeding conditions in this
ciency is thought to be low (3%) and much of the pri- region are thought to be good (Hutchings, 1992) and it
mary production may move through microhetero- does have a stable thermal structure (Barange, 1990)
trophs (Verheye et al., 1994). which may favour their persistence. Unlike conditions
on the Agulhas Bank, the survival of Agulhas Current
Our understanding of the zooplankton in the Agulhas expatriates in the nearshore waters of the west coast is
Current is confmed largely to taxonomy and we remain unlikely due to the variability of the nearshore environ-
ignorant of community processes. Zooplankton com- ment. Records of Agulhas Current species have been
munities in the current are much more diverse than made on the western side of the South Atlantic, espe-
those of the surrounding water masses and species cially in the tropical latitudes (Ponomareva, 1990). If
richness tends to decrease from east to west (Gibbons valid, these records can only suggest passage by
and Hutchings, 1996). Indicator species of the Agulhas Agulhas rings, and this has major implications for our
Current have been identified amongst euphausiids understanding of pelagic biogeography as a whole.
(Talbot, 1974), and chaetognaths (Stone, 1969;
Schleyer, 1977), as well as copepods (De Decker,
1984). These species have been detected in the near- Atkno~ledgements
shore waters of the Benguela system and at the front to
Hondeklip Bay up the west coast (Gibbons et al., Drs. Yasunobu Matsuura and Frederico Brandini' s
1995). Their presence there must largely reflect the critical comments on earlier versions of the manu-
influence of surface filaments. Further north along the script are appreciated. Drs. F. Shillington and B.
western seaboard of the African continent species Mitchell-Innes are gratefully acknowledged for their
attributed to the Agulhas Current have been observed comments on some of the physical and biological fea-
by Ducret (1968), Frontier (1968) and Godeaux and tures of the Agulhas Current. Several coauthors of the
Goffmet (1968). If these species really are of Agulhas present volume furnished data on their respective
Current origin then their presence so far north must groups of expertise.
Foraminifera
Sylvia Kemle-von Miicke and Christoph Hem/eben
Introduction 4 !lffi) and micropores (<1 !J.m) occur in non-spinose

species, whereas spinose species always exhibit nor-
Extant planktic Foraminifera occur in all pelagic envi- mal pores (4-7 !lffi) (Be, 1968). Pustules are present in
ronments of the World Ocean. Approximately 42 variable numbers on and within the rather smooth
morpho-species exist today, in contrast to several shell-wall of non-spinose species, or the wall surface
hundred fossil species. Of the 42 species, 35 have so far may be cancellate, as in some of the spinose species.
been recorded from the South Atlantic. Spinose species shed their spines during gametogene-
sis. Thus, shells settling on the sea-floor are seldom
Planktic Foraminifera have been recognized since the provided with spines, showing only spine holes.
early 18th century, one of the first good figures, Occasionally some remnants can be detected under the
including the cell, having been provided by Rhumbler SEM (Hemleben et al., 1989). The surface wall struc-
( 1911 ). They are used as stratigraphic markers to ture is either cancellate (honeycomb structure, see Fig.
determine ages of fossiliferous sediments, and since 51 below), or consists of irregularly distributed spine
Schott's (1935, 1936) key distributional studies, have bases. Microperforate species lack a cancellate texture
been used increasingly to trace recent and ancient cur- but may bear many small pustules. Microperforate spe-
rent systems (Vincent and Berger, 1981 ). cies have the smallest pore size, the highest pore con-
centration, but lowest porosity (Be, 1968).
Planktic Foraminifera form calcitic, multi-chambered
shells usually coiled either plani- or trochospirally, or In general, planktic Foraminifera are omnivorous.
the chambers may be hi- or triserially arranged (see Fig. However, spinose species may prefer a carnivorous
6.8 below). The wall is multi-layered and porous when diet, whereas the non-spinose species favor algal food.
seen from the outside. However, in life these pores are Animal prey observed within the cytoplasm of spinose
closed by organic material (Hemleben et al., 1989). In species includes ciliates, tintinnids, radiolarians,
all genera except one (Hastigerina), the primary wall is copepods (mostly calanoids), amphipods, pteropods,
bilamellar (Reiss, 1957), two calcitic layers separated heteropods, chaetognaths, and larvae of various kinds.
by a primary organic membrane. At each chamber for- All kinds of algae, mainly diatoms, but also prymne-
mation event a new organic and calcite layer is secret- siomonads (e.g., coccolithophorids and thin-walled
ed on top of the previous layer. Inside the test up to four green algae) are observed in the cytoplasm of non-spi-
calcite/organic layer couplets have been observed. nose species. Most spinose species host symbionts
Only subsequently organic layers are formed inside at (zooxanthellae; exceptions are Globigerina bulloides,
each chamber formation, creating the so-called inner Hastigerina pelagica and H. digitata, which are
organic layer (Hemleben et al., 1989). In most species devoid of symbionts), either dinoflagellates or chry-
a rather incomplete to thick calcite layer is deposited sophycophytes. This implies that spinose species live
prior to gametogenesis (gametogenic calcification), in the photic zone, although they may also survive in
which often covers spine holes (spines are resorbed deeper waters for a few days when expatriated from
during gametogenesis leaving holes or depressions in their original environment (e.g. , during storms, cf.
the wall) and their bases. In cases where species Schiebel et al., 1995). Some non-spinose species, like
migrate to deeper and/or cooler water, a secondary cal- Neogloboquadrina dutertrei, may host green algae
cite layer or crust is deposited on top of the normal facultatively , especially during spring blooms.
wall. The calcite crust has a coarse crystalline structure, Commensals and parasites are quite common in both
normally with euhedral or subhedral crystals. groups (Anderson eta/. , 1979; Hemleben et al. , 1989).
Following Murray ( 1897), one can distinguish between The reproductive cycle varies from species to species,
non-spinose and spinose species. Spines may be circu- although some similarities exist within various
lar or triangular in cross-section. Both normal pores (2- groups. Most of the spinose, symbiont-bearing spe-

edited by D. Boltovskoy, pp. 43- 73
44 Foraminifera
cies exhibit a lunar or semi-lunar cycle, reproducing 1976; Oberhansli et a/., 1992; Kemle-von Miicke,
at about either full moon or new moon. Their normal 1994; Ufkes et a!., 1998), as well as to characterize
life span ranges between 2 and 4 weeks (Spindler et ancient water masses over the last 100 million years.
a/., 1979; Hemleben eta/., 1989; Bijma eta/., l990a; They are used as paleoceanographic proxies (e.g. in
Erez et a/., 1991; Schiebel et a/., 1997). Some of the studies of stable isotopes of oxygen and carbon) to
non-spinose species follow the same reproductive determine temperature, salinity, oxygen content, and
scheme, while others, the deep-living non-spinose fertility in the changing ocean environment, thus con-
species, seem to follow a yearly cycle. All planktic tributing to understanding ancient ocean processes
Foraminifera perform ontogenetic vertical migrations. (see Hemleben eta/., 1989, for review).
The deep-living species, like Globorotalia truncatuli-
noides, reproduce in surface waters during spring and The geologic record of foraminifers dates back to the
then migrate downwards to layers as deep as 1000 m middle Jurassic. They became more abundant, and
or even deeper (Hemleben et al., 1985). Other spi- thus important for stratigraphy, since the mid-
nose, as well as non-spinose surface-dwelling species Cretaceous. During the first half of this century they
reproduce in the pycnocline or chlorophyll maximum were used principally for establishing zonal stratigra-
layer. Thus, each species seems to have its own typi- phy from the upper Valanginian to the Recent, finding
cal cycle and ontogenetically variable depth habitat. practical application mainly in oil exploration.
So far, only sexual reproduction (gamete release) has

been observed in laboratory cultures. However, a se- Methods
cond generation has never been successfully raised,
although zygotes and juveniles of up to four chambers Most surveys on the distributional patterns of planktic
have been observed. The ontogeny has been studied Foraminifera were based on sediment samples, rather
either using juveniles from plankton tows, or by back- than on plankton tows. Both methods are useful for
tracking the successive growth stages (chamber the study of distributional patterns (see chapter on
sequence). Most species show a distinct development "Radiolaria Polycystina"). However, comprehensive
described by Brummer et a/. (1986, 1987), who dis- biological studies must be based on plankton tows
tinguished five different stages: prolocular, juvenile, (Fig. 2, 3) and/or hand sampling by SCUBA methods
neanic, pre-adult, and terminal stage. This sequence are required.
usually involves a drastic change during the neanic
stage. The final chamber, built just prior to gameto- A third way of obtaining information on distribution-
genesis, may be morphologically different from the al patterns is by means of sediment traps. Time-series
previous ones: either larger or smaller, or equal in size sediment trap materials have several advantages over
to the previous chamber but different in other aspects. planktonic and/or sedimentary samples, e.g., larger
In the microperforate foraminifers a bulla (see below, sample-size, integration over preselected depth and
Fig. 5H) may be formed covering the aperture. time ranges.
The distribution pattern of living planktic Foraminifera None of these 3 approaches alone suffices to yield an
is determined chiefly by the climatic belts and their cur- adequate picture of the world-wide distribution pat-
rent systems. Thus, one can recognize antarctic, subant- terns of planktic Foraminifera, as all three have
arctic, transitional, subtropical, and tropical assembla- advantages as well as drawbacks (see "Radiolaria
ges (e.g., Be and Tolderlund, 1971; Hemleben eta/., Polycystina" in this volume). Information combined
1989). This pattern may be overprinted under special from various sources should therefore be used in order
hydrographic conditions, e.g., upwelling, shifts in to maximize the potential of these organisms for
boundary conditions, or seasonal events such as eleva- oceanographic and paleoceanographic surveys.
ted spring blooms or a sequence of heavy storms.
SCUBA sampling
Planktic Foraminifera are used to trace present-day Sampling planktic Foraminifera by SCUBA diving, in
ocean surface currents and their hydrographic proper- which specimens are captured by hand in solid contain-
ties (e.g., Boltovskoy, 1962, 1970; Cifelli and Benier, ers (Hemleben et a/., 1989), is very time consuming,
Foraminifera 45
but it is the best method for obtaining healthy speci- One has to be aware that plankton hauls are only spot
mens for culture experiments and cytological and ultra- checks in space and time, not accounting for the com-
structural studies in the transmission electronic micro- plete dynamics of species distributions. Thus, daily to
scope {TEM). However, SCUBA methods require weekly surface and subsurface plankton tows over
trained divers and well-equipped facilities to maintain extended time periods are needed to achieve a reason-
the specimens in laboratory cultures (Hemleben et a/., able picture of distribution patterns in the productive
1989), or to prepare specimens for TEM research. zone. Deeper (below ca. 80-100 m) plankton tows can
help evaluate processes in the flux zone where remin-
Plankton tows eralization of most juvenile specimens occurs. On the
Planktic Foraminifera have been collected in the past other hand, comparison of plankton tows and sedi-
with a wide variety of nets made with mesh of various ment samples can aid in revealing the effects of cur-
sized openings, towed through the water vertically, hor- rents and environmental conditions in the region.
izontally, or obliquely. Some have been equipped with
opening and closing devices to collect material from Sediment traps
descrete depths, sometimes fitted with flowmeters to Sediment traps are specially designed containers that
measure the amount of water represented by the sam- are deployed at various depths in the oceans for peri-
ples. Mesh sizes have varied between 10 and 300 11m in ods usually of several months to one or more years to
the last several decades. Hence, many data sets are dif- accumulate material settling from the water column
ficult to compare with one another, especially samples above (Honjo et a/., 1980; US GOFS , 1989). These
collected in the 60's and 70's compared to more recent samples describe quantitatively the seasonal or subsea-
decades. Today, most investigators use special opening sana) fallout usually below the productive layer plus
and closing nets, such as the Multinet (0.25 m2 open- organisms advected laterally from elsewhere (Siegel
ing) or the MOCNESS system (Wiebe eta/., 1976), and and Deuser, 1997), both modified by processes that
usually with mesh openings of either 64 or 100 11m. The affect settlement and preservation (Boltovskoy, 1994).
Multinet is usually towed at a speed of approximately Their shortcomings include restricted effectiveness
0.3-0.5 m s- 1. Samples are preserved with HgC1 2, for- when deployed at depths less than 500-700 m, the inte-
maldehyde, or methyl alcohol, and stored at 4°C on gration of the flux from several biologically dissimilar
board. In the laboratory samples are split with a rotary layers, distortion of the near-surface patterns due to
liquid splitter. Wet subsamples may be used without selective and non-selective destruction (dissolution,
further processing, or they may be combusted (< 150°C) grazing), and lateral advection of particles (see chapter
to remove the cytoplasm and other organic compo- "Radiolaria Polycystina").
nents. If living-to-dead ratios in the water-column
assemblage are sought, foraminiferal cytoplasm can be Sediment coring
stained using rose Bengal, Sudan black B, or eosin For paleoceanographic studies, the Multicorer is the
(Boltovskoy and Wright, 1976), in which case ashing is best gear for obtaining undisturbed sediment samples
not possible. However, after some practice "living" from the ocean floor. Usually, a Multicorer core co-
shells can be identified without staining because speci- vers the last 3 to 4 kyr. If longer cores are desired, a
mens containing cytoplasm are easy to distinguish from piston core or other coring devices can be used (see
empty ones by their color. Nevertheless, cytoplasmic chapter "Radiolaria Polycystina" in this volume).
remnants may persist in dead specimens for weeks and
months (Boltovskoy and Lena, 1970; Bernhard, 1988),
especially if they sink without contact with larger phy- Distribution
toplankton aggregates with dense bacterial populations.
Planktic Foraminifera occur in all oceans, although
Samples are thoroughly wet-sieved through a 30 11m they are scarce in shelf areas and closed marine basins
mesh sieve, oven-dried, and sieved again into suitable like the Red Sea. Densities depend greatly on food
grain size fractions (e.g., >250 !liD, 150-250 11m and availability and hydrographic characteristics, varying
<150 11m). Counting is usually performed on splits from less than 1 to over 100 specimens per m3 (e.g.,
containing approximately 300 specimens. A random Caron and Swanberg, 1990; see Fig. I, bottom panel),
check should be carried out in order to assure correct with vertical flux rates calculated between 0.1 and
aliquot size . 100 g CaC0 3 m-2 y-1.
46 Foraminifera
0
ooooo
WWS, tropical o~~~or: o oo o o o o o o o o o o ooo
C. ruber (white & pink) o~
C. trilobus, C. sacculifer i
C. menardii o
0
0
0
o a
"<P
Coastal
Gyre
upwelli~g
C. truncatulinoides (right) o"
c 0
c 0
aooo oooo o o oooooo o ooo o o ooo o a o oo o o oo o oo oooo o Do o

o o a ooooo o D
CWS, subtropical ocean
C. bulloides, N. pachyderma,
C. inflata, C. hirsuta,
C. truncatulinoides (left)
Specimens per m-3

50° 0.001-0.9
1-9
Foraminifera 47
Published planktic foraminiferal distributional pat- upper 50 m of the water column where temperature,
terns are based mainly on ocean floor sediment sam- food, and light (for symbiont photosynthesis) are
ples (Fig. 4). Species-specific geographic distribu- appropriate. Below this depth foraminiferal concentra-
tional data are moderately abundant in the literature, tions usually decrease exponentially. Thus, most of the
but scarce and fragmentary for most of the rare forms. shells originate from the upper 50 to 80 m, which
It is widely accepted that, in general terms, species should be kept in mind when using planktic foraminif-
occurrences are controlled chiefly by climatic belts eral abundances for paleoceanographic purposes.
(mostly temperature-dependent), by current systems,
and by food availability. All three boundary condi- Fig. 2 illustrates the occurrence and relative abundanc-
tions vary seasonally. Additionally, factors such as es of the most common foraminiferal species in the
reproductive behavior (e.g. semi-lunar, lunar, or year- Southwestern Atlantic (pooled data for 0-50 m). Notice
ly cycles), and local water properties (e.g., river dis- that, while the distribution of some species exhibits dis-
charge, such as the Zaire River, cf. Oberhiinsli et a/., tinct geographic patterns, others, such as Globigerinita
1992; Ufkes eta/., 1998) may also be of major impor- glutinata and Turborotalita quinqueloba (=Globi-
tance. Thus, the actual distribution of a species may gerina quinque/aha in Boltovskoy et a/., 1996), are
be rather patchy both in space and time. Despite these present at almost all locations, albeit in variable propor-
difficulties, planktic Foraminifera may be used to tions. For the eastern South Atlantic, species distribu-
trace surface water hydrographic conditions. tions in the plankton (0-50 m) are illustrated with a
series of maps based on the works ofWeferet a/. (1988,
Foraminiferal diversity decreases drastically from 1991 ), Oberhiinsli et al. ( 1992 ), and Kemle-von Miicke
approximately 15-20 species in a tropical assemblage, (1994) (Fig. 3). Fig. 4 offers a more detailed picture of
where 6-8 species dominate, to 5-6 species in the an- foraminiferal species-specific distributional patterns in
tarctic realm where only one species dominates (Fig. the temperate to tropical South Atlantic based on sedi-
2). Mixed assemblages occur where cold water currents mentary materials (from Pflaumann et al., 1996).
enter more temperate belts (Fig. 2), in upwelling areas,
or where river outflow creates a surface layer of low The following section offers comments on individual
salinity. Mixing of water masses either vertically or species distributional patterns and general ecological
horizontally usually leads to expatriation of species preferences (species are ordered alphabetically). A
with concomitant abundance drops, but it also may few taxa, for which data are too scarce, are omitted.
enhance a population if nutrient-rich water is intro-
duced. Candeina nitida: Tropical, rare.
Dentagloborotalia anfracta: Probably restricted to
Although at global scales the influence of temperature transitional-temperate to tropical waters, where it is
is dominant, its role decreases at more restricted moderately common. However, because of its small
scales, as most species tolerate rather wide tempera- adult size (120-200 Jlm) it may be routinely under-
ture ranges. At these scales foraminiferal distribution represented in standard collections, for which rea-
is strongly dependent on food availability. This may son its distribution pattern is poorly known.
partly explain observations of warm and cold water Globigerina bulloides (Fig. 2, 3C, 4A): Abundant in
species coexisting in the same water mass (e.g., subpolar and, to a much Jesser extent, polar water
Boltovskoy eta/., 1996; Fig. 2). masses. In tropical areas it favors nutrient-rich
upwelling waters, where it can reach high concen-
The species Globigerinoides ruber, Globigerinoides trations (e.g., near the coastal upwelling off
sacculifer, Globoturborotalita rubescens, Globigeri- Namibia and toward the eastern equatorial upwell-
nita glutinata and Turborotalita quinqueloba domi- ing region, where phosphate and nitrate concentra-
nate South Atlantic tropical to temperate assemblages, tions are higher than in other parts of the tropical
followed by Globorotalia menardii and Neoglobo- and subtropical South Atlantic, cf. 30 m water
quadrina dutertrei. Most individuals are found in the depth annual means in Levitus and Boyer, 1994).
Fig. l. Generalized planktic foraminiferal biogegraphic zones in the South Atlantic (top panel), and their abundance in the upper 10m of the
water column (bottom panel, from Be and Tolderlund, 1971).
Latitude south +>-
00
30 35 40 45 so 55
I I I I I I I I I I I I I I I I I I I I I I I I I _) Globorotalia menardii
-:J Globoturborotalita rubescens
_) Globigerinella siphonifera
..... =:/ Orbulina universa
(J)
...... ~ Hastigerina pelag ica
Ill ~ Neogloboquadrina dutertrei
3: - Globigerinoides ruber
E
..... Globigerinoides sacculifer
Ill ~ Globigerinoides conglobatus
$ - Globigerina fa lconensis
Transitional[ - Globorotalia inflata
Cosmopolitan - Globigerinita glutinata
lE3 I ~-
- Neogloboquadrina pachyderma (left)
-- Neogloboquadrina pachyderma (ri ght)

.....
(J)
......
Ill
•••Ill- Turborotalita quinqueloba
~
3: h IIIII- Globigerina bulloides
---c
0 _j Globigerinita uvula 3
u
~~ J - • I _j Globorotalia trun ca tulinoides
s·
l==:___ _ __;;:;::;:;:;::;:;;;;:;;;:::;;;::::;::;::;;;;;:;;;;;;;;;;~~=====;:;::;:;:;::;:;:;::;:;:;::;:::;;::====----=====---_j- - Globorotalia scitula
~
Pl
100%
50
0 '"'''"'_,~,- -,,-,,.
<X> R ~
Subantarctic
::> ~
Antarctic
Subtropical Warm Transitional
Transitional
1
-
~ ;~
r~ - y- , , 1 .
__,.,.
... 60°
. ....~
70° 60° 50° 40°

Foraminifera 49
Globigerina falconensis (Fig. 2): In the South stations with the last quarter (0-3 days before the
Atlantic, transitional to warm water. moon phase). This may agree well with the results
Globigerinella calida: Limited to warm waters. ofBijma eta/. (1990a), who document a semilunar
Globigerinella siphonifera (Fig. 2, 4B): Restricted to reproduction periodicity for Globigerinoides
temperate to warm water masses, conspicuous ruber. However, Van Leeuwen (I 989) also found
member of foraminiferal assemblages in the South higher percentages of Globigerinoides ruber at
Atlantic Central Gyre. 10°S, 10°E in the Angola Basin, and at the Equator
Globigerinita glutinata (Fig. 2, 3, 4C): The most on the sea floor, as well as lower frequencies off
widely distributed planktic foraminifer, occurring the south coast of Angola. It is therefore probable
from the equator to the poles. It seems to peak in that despite the influence of the reproduction cycle,
abundance in areas where nutrients are brought concentrations of Globigerinoides ruber in the
into the mixed layer. In the tropical South Atlantic water column still do reflect its preference for par-
it shows maximum abundances during April ticular oceanic areas. In the South Atlantic
through June. Its contribution to the total planktic Globigerinoides ruber pink form is generally more
foraminiferal fauna may be in the order of 8-18%. than 5 times less abundant than Globigerinoides
Globigerinita uvula (Fig. 2): Its distribution pattern ruber white form (cf. Fig. 4E and 4F).
seems to be obscured by its opportunistic behav- Globigerinoides sacculifer (Fig. 3B, 4G, H): Tropical
ior. Normally it is rare, but can be abundant at to subtropical. In the eastern South Atlantic it
restricted times and areas (e.g. , in cold waters of peaks above the Walvis Ridge (Fig. 3B, 4H), and
the southwestern South Atlantic, 30-60°S, cf. in the equatorial region. This distribution pattern
Boltovskoy et a!., 1996). In the eastern South may also be affected by its reproduction cycle.
Atlantic, Oberhansli et a!. ( 1992) found very few Maximum abundances occur I and 5 days after
individuals in the cold Benguela Current. full moon, which fits the reproductive cycle of a
Globigerinoides conglobatus (Fig. 2, 4D): Usually full lunar period observed by Bijma eta/. ( 1990a).
present in low numbers in warm to temperate Nevertheless, comparison of the abundance of
waters, but Benier (I 985) reported rather high con- Globigerinoides sacculifer with water tempera-
centrations from the eastern tropical Atlantic. Deep ture, salinity, and dissolved oxygen indicates
dwelling (at the base of the seasonal thermocline). enhanced numbers with increasing oxygen content
Globigerinoides ruber (Fig. 2, 3A, 4E, F): The most and lower salinity (Oberhansli et a!., 1992). This
common species in warm to temperate South suggests that the distribution pattern is not the
Atlantic waters. The temperature range of the result of reproductive behavior alone. In the
white form, however, is broader and extends into Walvis Ridge area, oxygen-rich subtropical sur-
temperate water masses. Thus, Globigerinoides face water eddies are mixed with lower salinity
ruber pink form may be considered a summer spe- water of the Benguela Current. The abundance
cies, whereas Globigerinoides ruber white form maximum at the equator corresponds roughly with
occurs year-round and dominates during the austral the oxygen-rich South Atlantic Central Water and
winter. In the eastern South Atlantic it is most the less saline surface water coming from the coast
abundant in the Angola Gyre, with significant con- of Equatorial West Africa (Levitus and Boyer,
centrations also in the equatorial upwelling area, 1994). This suggests that Globigerinoides sacculi-
but low numbers elsewhere (Fig. 3A). Its wide- fer is an indicator of oxygen-rich, low-salinity
spread distribution in the underlying sediment sug- waters, yet in other regions, like the Caribbean,
gests that the planktonic pattern is a result of repro- Globigerinoides sacculifer prefers higher sali-
ductive habits. Abundance maxima in the Angola nities, whereas Globigerinoides ruber occurs in
Basin coincide with the first quarter (2 days before low-salinity water masses. These contrasting
the moon phase), and at the Equator and northern observations suggest that, within certain limits,
Fig. 2. Latitudinal changes in the relative abundances of planktic Foraminifera (0-50 m, top panel) along a transect in the southwestern South
Atlantic (inset map at bottom right), in November 1993, and corresponding surface water temperatures (bottom panel). Pooled data (middle
panel) exclude Globorotalia in.flata and Globigerinita glutinata. Based on data from Boltovskoy et al. (1996).
50 Foraminifera
10°
oo
19(
IQO 17(
15(
13(
20°
11 C
90
1 10 J021Q3 1
70
'l1
30° 50
1 30
10
1000
55
10
: i ~[
1 1010'10' 10•
0
Neogloboquadrina
dutertrei menardii
0 1000 1----,.-- --,------1
0
10°
Fig. 3. Abundance of selected foraminiferal species (ind. m-3 of water filtered) in the surface layer (0-50 m) of the eastern South Atlantic. In
all cases values range between the maximum concentration indicated and 0 (shadings of decreasing darkness). For vertical profiles (ind. per
1000 m3 of water filtered), hatched bars indicate "live" specimens, white bars "dead" individuals. Data from Wefer et al. (1988, 1991), Kemle-
von Miicke (1994), and Oberhiinsli, et al. (1992).
salinity seems to play a minor role (Bijma et al., best showing this abrupt decline with depth; only
1990b). On the other hand, food availability may "dead" specimens are recorded in deeper net
explain these contrasting distributional patterns. hauls. Downward flux of "dead" shells blurs the
Field and laboratory observations indicate that downward decreasing trend somewhat (Kemle-
Globigerinoides sacculifer occupies more oligo- von Miicke, 1994).
trophic water masses and feeds mostly on cope- Globorotalia crassaformis (Fig. 3F, 41): Tropical to
pods, whereas Globigerinoides ruber prefers subtropical, normally not present south of 30°S
eutrophic environments. The depth distribution of (Van Leeuwen, 1989). Although only a minor con-
Globigerinoides sacculifer is typical of spinose, stituent of typical deep-living planktic foramini-
symbiont-bearing foraminiferal species (Fig. 3B): fers, it has an interesting habitat. It shows frequen-
highest concentrations in the upper 50 m, decreas- cy maxima at the eastern edge of the
ing sharply further down, with "living" specimens Angola-Benguela Front (17°10 S, 8°54 E), and at
Foraminifera 51
the Equator, both areas of high productivity in the water-column is not yet clear. The species
(Berger and Herguera, 1992). As previously might be transported by a southward-flowing sub-
reported by Jones (1967), its greatest abundance is surface current (Oberhansli eta/., 1992) because its
observed in the equatorial oxygen minimum zone preferred depth habitat here appears to be the lower
between 100 and 300 m (3F, 4I). Van Leeuwen photic zone or some 50 m deeper (e.g., 200-400 m).
(1989) noted a peak in sediments off the Zaire As previously observed by Jones (1967), the depth
River, where there is a pronounced oxygen mini- distribution of Globorotalia menardii indicates
mum layer at depths of 300-400 m (Van affinities with the Equatorial Undercurrent, which
Bennekom et a/., 1978). Optimum conditions for flows along the equator at 50-100m. Interestingly,
Globorotalia crassaformis include a shallow ther- a significant number of "living" Globorotalia
mocline, high productivity in surface waters, and a menardii were found at depths of 500-1000 m. This
distinct subsurface oxygen minimum where sink- suggests a wider depth habitat during ontogeny, as
ing organic matter is oxidized. Thus, enhanced is common for deep living globorotaliids. However,
numbers of Globorotalia crassaformis in the sedi- expatriation phenomena may be involved in this
ments may indicate this type of oceanographic set- pattern as well, as in many other species in, for
ting. In the western South Atlantic this species is example, the Red Sea, without showing the high
not found along the Brazilian coast between 8°N salinity Red Sea signals in e.g. ~)180. In the western
and 32°S (Benier, 1985), and has never been South Atlantic this species occurs in rather low
recorded farther south. numbers, seldom exceeding 3% of all foraminifers.
Globorotalia hirsuta (Fig. 4J): Scarce but widely dis- Globorotalia scitula (Fig. 2, 4M): Deep-living, rare in
tributed south of 20°S in subtropical to temperate transitional cooler waters and almost absent in
water masses. Deep- and long-living foraminifer low-latitude regions. Its pattern in the North
which usually reproduces in surface waters during Atlantic is quite different, as here it is common
austral mid-spring. along the temperate and subtropical belts.
Globorotalia injlata (Fig. 2, 4K): Often a dominant Globorotalia truncatulinoides (Fig. 2, 4N, 0): Its dis-
component in temperate waters (up to 33% of all tribution is similar to that of Globorotalia scitula,
foraminifers; see Boltovskoy et a/., 1996), but also but it is more common. The right coiling form
present in cold as well as warm areas, preferring the spreads farther equatorward than the left coiling
former. In the eastern South Atlantic it peaks one. Deep-living, reproduces in early austral
around 40°S. In the western South Atlantic and in spring in surface waters and migrates during the
the eastern North Atlantic it is abundant in spring. summer and fall to deeper layers. In the western
Its reproduction is closely related to the lunar cycle North Atlantic (Bermuda) it has been observed
(Schiebel and Hemleben, unpublished data). It living below 1000 m (Hemleben eta/., 1985).
feeds on phytoplankton available in the photic zone. Globorotalia tumida: Its distribution is similar to that
Globorotalia menardii (Fig. 2, 3E, 4L): In the south- of Globorotalia menardii, but it is less abundant.
western South Atlantic (30-60°S), this species is Globorotalia ungulata: Rare species with poorly
scarce, rarely exceeding 3% of the entire foramini- known ecological affinities; probably similar in its
feral assemblage (Boltovskoy eta/., 1996). In con- distribution to Globorotalia menardii.
trast, it is abundant in the eastern South Atlantic in Globoturborotalita rubescens (Fig. 2): Typically
the area of the Walvis Ridge, especially at 21 °43'S, warm water, occasionally occurring in fairly high
5°25'E, to the west of the Zaire River mouth, and at numbers (up to 3% of the spring assemblage in
the equator (Fig. 3E). Several authors consider tropical regions).
Globorotalia menardii a rather common member Hastigerina pelagica (Fig. 2): Tropical to subtropical,
of the fauna in the tropical ocean (Jones, 1967; Be, very rarely observed in sediments.
1977); thus, maximum abundances at the equator Neogloboquadrina dutertrei (Fig. 2, 3D, 4P): Frequent
are quite typical. Its high abundances above the in the eastern tropical to temperate South Atlantic,
Walvis Ridge (dominant in the cold Benguela less abundant in the subtropics. Fig. 4P shows that,
Current) are therefore inconsistent with its tropical in the eastern South Atlantic, highest concentra-
distribution pattern. Globorotalia menardii is very tions of N. dutertrei are recorded off the African
rare in sediments under the Benguela Current (Van coast. Previous studies indicate that this species
Leeuwen, 1989). The reason for its high abundance reaches peak concentrations where the thermocline
VI
2ao N
3 3 2
4
1ao 2.5 ltri/MtJJIJ
ao ~--
~ ----- - -
. ·E :
:> it!l.'
4
1ao ... ~· 4
2
ao _...... 2.5 , ~ 3 j 3
3ao. , -- 5 5 - 2
10 2 I 0 1 '"Ij
;~: 4
10
8
6
4 Globi erinita
C ~:.,"<i
....,.
)( )I 2
~1 °
D
Globigerinoides
;
2.
~
1ao 6 /1 / -
~· 8
ao
1ao
ll
2ao 6
8
6 6
3 oa ~J~l~i~~;ir~ : ,:
2 4
4
4a 6a
0
0 sao 4a 0 3a 0 2a 0
1ao ao 1ao 2a 0
6a 0 sao 4a 0
3a 0 2a 0
1ao ao 1ao 2a 0
2a 0
4
4
1ao 2
ao 4
4
1 ao
4
2a 0
3a 0
2
4a 0
2a 0
f=
.....
~
""!
s:>l
1ao
ao
::'I IV
7.5
1 ao 10
7.5
2a 0
5
7.5
2.5
. 2.5
3a 0
4a 0 -
6a 0 sao 4a 0 3a 0
2a 0
1a0 ao 1 ao 2a 0
6a 0 sao 4a0 3a 0
2a 0
1a0 ao 1ao 2a 0
Vl
w
Vl
~
20°
0.5
10° 0.5
oo1111
0.5
10°
0.5
zoo
30°
61
i;l
zoo §.
::l
2 4 ~
i;l
10°
oo
10°
zoo 4
30°
40° L-------------~~
60° 50° 40° 30° zoo 10° oo 10° zoo 60° 50° 40° 30° zoo 1QO oo 10° 20°
20° --~
3
1
0
0
. 2
lQ O .
oo
10°
20°
30°
~2
2
1
1 0 0
40° 2 "Tj
0
'"1
::r8
20°
0.5
, ..
··· -
~
'"1
!))
{<
10° 5 5 10
oo til. til.
lQ O
20°
30° u.:. - . w ..,Sf~~~\~-~. J·Z"·...>t.··

40°
IF ,~1:J
~t~:~. ~··
'" (3 5~ 'l! ~ 5
5
60° 50° 40° 30° 20° 10° oo 10° 20° 60° 50° 40° 30° 20° 10° oo 10° 20° V1
V1
2a 0
18 Vl
0\
4
1ao 2
1 1
ao Ill.
1ao
2a
30"r.
0
2
-- - 0.5
4a 0
-
1 ~
u1ill! l~iJh'r
- 0.5
· ···--------------------------------------
I
>-rj
0>-;
~
ss·
2a 0
4 ~
>-;
2 "l ~
1
1ao 2
ao Ill.
1
1 2
1ao
2a 0
3a 0
4a 0
6a 0 sao 4a 0
3a 0
2a 0 lQO ao lQO 2a 0
6a 0 sao 4a 0
3a 0
2a 0 1ao ao 1ao 2a 0
Foraminifera 57
is shallow and the chlorophyll maximum is located foraminiferal assemblage. Normally, however,
between 25 and 50 m (Fairbanks and Wiebe, 1980; very rare (<0.1 %), and therefore seldom recorded.
Fairbanks et al., 1982; Ravelo et al., 1990; Kemle- Sphaeroidinella dehiscens: Tropical to subtropical,
von Miicke, 1994). Both conditions are met in the very resistant to dissolution.
Guinea Basin, which may explain its high local Tenuitella iota: Prefers cool waters, but also occurs in
abundances. On the sea floor it exceeds 10% of the tropical ones. Always scarce.
entire foraminiferal thanatocoenosis, reaching 25% Tenuitella parkerae: Minor contributor to tropical to
in the Guinea Basin (Van Leeuwen, 1989). In the temperate assemblages.
North Atlantic this species occurs in tropical as Turborotalita clarkei: Known so far from temperate
well as subtropical regions, and its depth stratifica- to subtropical waters.
tion is tied to the physico-chemical and biological Turborotalita humilis: Temperate to subtropical,
properties of the upper water column (e.g., thermo- dwells in the photic zone.
cline depth, chlorophyll maximum). As shown Turborotalita quinqueloba (Fig. 2): In the South
experimentally, it forms a calcite crust when the Atlantic chiefly cold to temperate; especially abun-
water temperature drops below l5°C (Hemleben et dant during spring bloom conditions at low latitudes.
al., 1989).
Neogloboquadrina incompta (Fig. 4Q, as N. pachy-
derma right): As opposed to N. pachyderma left Taxonomy and identification
coiling, a typical subpolar to polar species, N.
incompta dwells in temperate to subtropical The classification of planktic Foraminifera used in this
waters. Fig. 4Q refers to a data set ofPflaumann et chapter was adopted from various sources (Loeblich
al. ( 1996), where these authors used a different and Tappan, 1988; Hemleben et al., 1989; Olsson et al.,
definition: N. pachyderma right coiling and inter- 1992). In general, the following major informal groups
grades. Thus, figure 4Q displays mostly N. can be identified: spinose species, non-spinose species,
incompta, but may also include forms allied to N. normally perforate species, and species without spines
dutertrei. but with micropores. The last group, which probably
Neogloboquadrina pachyderma (Fig. 2, 4R): N. derives from Cretaceous ancestors (Olsson et al.,
pachyderma left coiling is nearly restricted to cold 1992), has been called "microperforate". Micropores
water masses, and is very often the only planktic are not readily visible under the binocular microscope.
foraminifer in Antarctic and Arctic waters. In spinose species it is very difficult to see the spine
Antarctic specimens can withstand high salinities holes after a species has undergone gametogenesis,
(in excess of 80%o), and may live in small channels which is the case in more than 99% of sediment-
within the ice (Spindler and Dieckmann, 1986). derived specimens. However, spines are usually sup-
Orbulina universa (Fig. 2, 4S): In the South Atlantic ported by collar-like structures which after shedding
this species seems to be almost restricted to warm give the surface a rough "pustulose" appearance.
water masses and upwelling areas of the Benguela
front, with only a few specimens having been
observed in cooler waters in the western South Glossary
Atlantic (Boltovskoy, 1962). However, in the
North Atlantic it ranges from subpolar/temperate The following glossary defines some of the terms
regions to the tropics. most commonly used in descriptions of planktic
Orcadia riedeli: Cold water but tolerant to warmer Foraminifera.
waters, probably cosmopolitan.
Pulleniatina obliquiloculata (Fig. 4T): Probably Apertural lip (Fig. 5C): a shielding plate in the form
restricted to warm waters; in the tropics it can of a lateral outgrowth covering the aperture total-
account occasionally for up to 7% of the entire ly or partially.
Fig. 4. Distribution of planktic Foraminifera in sediments of tbe Soutb Atlantic (data set from Pflaumann et al., 1996). Contour plots represent
the relative abundance of the different species. For special remarks regarding Fig. 5G, H, N, 0, Q and R see species distribution and remarks.
58 Foraminifera
last
last chamber
chamber sacciform,
elongated
sutures B
(Giobigerinoides sacculifer)
left coiling
(spiral view) sutural
(Globorotalia inflata) A apertures
(secondary)
keel
rim
smooth
surface
apertural aperture
lip globose
chambers
(Candeina nitida)
D
(Globorotalia inflata)
(Globorotalia menardii)
extraumbilical
aperture
spiral~
f~ ;ld'.,"""l
equatorial
aperture,
slit-like
side
flat conical
angular,
conical
chambers
trochospiral coiling G
(lateral view)
(Globorotalia truncatulinoides)
bulla umbilicus
H J
honeycomb (hexagonal,
infralaminal apertural view cancellate) surface texture
apertures (Globorotalia scitula) (Giobigerinoides sacculifer)
Foraminifera 59
Apertural side (=ventral side) (Fig. 5B, C, D, F, H, 1): Inner organic lining: a thick, dense layer between the
side of the shell on which the aperture is located; inner test surface and the intratest cytoplasm of
only the last whorl is visible in this view. planktonic Foraminifera, especially in Globorotalia.
Aperture (Fig. 5C, D, E, F, G, H): opening (or multi- Juvenile stage: in foraminifers, the growth stage from
ple openings) in the chamber through which cyto- formation of the proloculus up to the beginning of
plasmic strands emerge. Shells may have one the neanic stage.
aperture or several; in the latter case there usually Keel (Fig. 5C): a peripheral crest along the outer mar-
is one main (larger) opening and several smaller gin of the chambers, usually thickened by addi-
ones (secondary or supplementary apertures). tional calcite deposition.
Aposymbiotic: barren of symbionts. Kumrnerform: term used to designate specimens in
Bilamellar wall: a primary wall composed of a pri- which the last chamber is equal to or smaller than
mary organic membrane separating two calcite the preceding one Berger ( 1969). The formation of
lamellae. The outer layer is deposited concurrent- kummer chambers may result from environmental
ly over the exterior of previously built chambers. stress or from reproductive traits (Hecht and
Bubble capsule: cytoplasmic alveoli surrounding the Savin, 1971 , 1972; Olsson, 1973).
test of Hastigerina pelagica and resembling a Left coiling (=levogyrous) (Fig. 5A): arrangement of
mass of soap bubbles. chambers in counter-clockwise direction with the
Bulla (Fig. 5H): thin calcitic cover over the aperture shell viewed from the spiral (or dorsal) side. When
leaving one or more small openings through which chambers are arranged in clockwise direction the
rhizopods emerge. shell is named right coiling (=dextrogyrous or
Carbonate compensation depth (=CCD): oceanic level dextral).
at which the solution rate of calcium carbonate Levogyrous: see left coiling.
equals the rate of supply from above. Lip: see aperturallip.
Calcite crust: secondary secretion of calcite over the Lunar periodicity (=lunar cycle): The synodic cycle of
exterior of the test, often resulting in a doubling of the moon phases, with a period of29.5 days.
the wall thickness in late growth stages. Micropores: small pores (< 1 Jlm) in the shell wall,
Cancellate (Fig. 5J): surface texture in which the characteristic of some non-spinose species.
pores are surrounded by hexagonal ridges. Monolamella: calcitic test wall deposited on top of an
CCD: see carbonate compensation depth. organic layer.
Chamber (Fig. 5A, B, E, G): each of the series of more Neanic stage: growth stage between the juvenile and
or less spherical structures or sections of a forami- the adult stages; a short transitional stage charac-
niferal shell; the initial chamber is the proloculus. terized by sharp changes in morphology.
Crust: see calcite crust. Non-spinose: devoid of spines on the surface of the
Dextral (=dextrogyrous, right coiling): see left coiling. test.
Dextrogyrous (=right coiling, dextral): see left coiling. Planispiral (Fig. 5F): coiling pattern in one plane.
Dorsal side: see spiral side. Pore: small canal penetrating the wall and closed by an
Euhedral crystals (or crystallites): crystals with regu- organic membrane, sometimes with micropores.
lar crystallographic shape, with well-defined crys- Primary organic membrane: organic layer between the
tal faces and interfacial angles. two initial calcite layers of rotaliid test walls; con-
Extraumbilical aperture (Fig. 5G): aperture located tinuous with the pore plate.
away from the umbilicus. Proloculus (Fig. 5A): initial chamber in Foraminifera.
Gametogenic calcification: facultative calcification Pustule (Fig. 6.18d, e): small protuberance of the wall,
prior to gametogenesis producing an additional with internal layers, serving as attachment site for
layer on top of the entire shell or parts of it. the rhizopodia.
Infralaminal (aperture) (Fig. 5H): Secondary aper- Right coiling (=dextrogyrous, dextral): see left coiling.
tures between the edges of the bulla and the under- Secondary aperture (Fig. 5E): additional aperture on
lying shell-wall. the spiral side of the shell.
Fig. 5. Characters used for identification of planktic Foraminifera.

60 Foraminifera
Septum: section of the shell wall separating two con- Superfamily Globorotaliacea Cushman,
tiguous chambers. 1927 (non-spinose and normal perforate)
Spines: thin calcite projections anchored to the wall; (genera Dentagloborotalia, Globorotalia,
they leave characteristic holes after spines are Neogloboquadrina, Pulleniatina
shed during gametogenesis. Superfamily Heterohelicea Cushman, 1927
Spinose: with spines on the shell surface; in (non-spinose and microperforate) (genera
Foraminifera this trait evolved during the earliest Candeina, Globigerinita, Tenuitella)
Cenozoic.
Spiral side (=dorsal side) (Fig. 5A): side of the shell Species so far recorded in the South Atlantic can be
from which all chambers are visible (if not over- grouped into 3 categories:
grown by additional calcite).
Streptospiral: irregular coiling pattern. Spinose species:
Subhedral crystals (or crystallites): sub-regular crys- Globigerina bulloides d'Orbigny, 1826
tals with poorly developed crystal faces and inter- Globigerina falconensis Blow, 1959
facial angles. Globigerinella calida (Parker, 1962)
Suture (Fig. 5A): depression of the shell-wall between Globigerinella siphonifera (d'Orbigny, 1839)
two adjacent chambers. Globigerinoides conglobatus (Brady, 1879)
Taphocoenosis (-thanatocoenosis ): assemblage of Globigerinoides ruber (d'Orbigny, 1839)
dead shells deposited on the ocean floor. Globigerinoides sacculifer (Brady, 1877)
Terminal stage: adult stage shortly before reproduction. Globoturborotalita rubescens (Hofker, 1956)
Test: hard shell of the Foraminifera. Globoturborotalita tenella (Parker, 1958)
Thanatocoenosis: assemblage of dead specimens of a Hastigerina digitata Rhumbler, 1911
population settling to the sea floor. Hastigerina pelagica (d 'Orbigny, I 839)
Trochospiral (Fig. 5G): spiral coiling pattern, resem- Orbulina universa d'Orbigny, 1839
bling conical snail shells. Orcadia riedeli (Rogl and Bolli, 1973)
Umbilicus (Fig. 5I): depression in the center of the Sphaeroidinella dehiscens (Parker and Jones, 1865)
apertural (or ventral) side. Turborotalita clarkei (Rogl and Bolli, 1973)
Ventral side: see apertural side. Turborotalita humilis (Brady, 1884)
Turborotalita quinqueloba (Natland, 1938)
Outline classification
The following classification system is herewith adopt- Non-spinose species:
ed: Dentagloborotalia anfracta (Parker, 1967)
Globorotalia crassaformis (Galloway and Wissler,
Kindom Protista Haeckel, 1886 1927)
Phylum Sacomastigophora Honigberg and Globorotalia hirsuta (d'Orbigny, 1839)
Balamuth, 1963 Globorotalia injlata (d'Orbigny, 1839)
Subphylum Sarcodina Schmarda, 1871 Globorotalia menardii (Jones and Brady, 1865)
Superclass Rhizopoda von Siebold, 1845 Globorotalia scitula (Brady, 1882)
Class Granuloreticulosea de Saedeleer, 1934 Globorotalia truncatulinoides (d'Orbigny, 1839)
Order Foraminifera d'Orbigny, 1826 Globorotalia tumida (Brady, 1877)
Superfamily Globigerinacea Carpenter, Globorotalia ungulata Bermudez, 1960
Parker and Jones, 1862 (bilamellar, spinose Neogloboquadrina dutertrei (d'Orbigny, 1839)
and normal perforate) (genera Globigerina, Neogloboquadrina incompta (Cifelli, 1961) (=right
Globigerinella, Globigerinoides, Globo- coiling N. pachyderma)
turborotalita, Orbulina, Sphaeroidinella, Neogloboquadrina pachyderma (left coiling)
Turbo rota/ita) (Ehrenberg, 1861)
Superfamily Hastigerinacea Bolli, Pulleniatina obliquiloculata (Parker and Jones, 1865)
Loeblich and Tappan, I 957 (monolamellar,
spinose and normal perforate) (genera Microperforate species:
Hastigerina, Orcadia) Candeina nitida d' Orbigny 1839
Foraminifera 61
Globigerinita glutinata (Egger, 1893) • Dentagloborotalia anfracta (Fig. 6.2) [=Globo-

Globigerinita uvula (Ehrenberg, 1861) rotalia anfracta]. Non-spinose species; adults
Tenuitella iota (Parker, 1962) from 120 to over 200 J..LII1 (Brummer and Kroon,
Tenuit ella parkerae (Bronnimann and Resig, 1971) 1988); thus, only large specimens normally caught
in standard plankton nets. Resembles Pulleniatina
In his 1981 review of the identification and distribution in streptospiral coiling during early ontogeny; and
of planktic foraminifers from the Southwestern also Globorotalia scitula, but is smaller, with
Atlantic, in addition to the above, E. Boltovskoy more rounded margin and higher aperture. Curved
( 1981 b) included the following species: G/obigerina sutures, rounded periphery and aperture extending
megastoma Earland, G/obigerinella adamsi (Banner from the .umbilicus to periphery are typical for this
and Blow), Globigerinoides elongatus (d'Orbigny), species. Small denticle-like pustules in front of
Globoquadrina conglomerata (Schwager), aperture are a unique feature not observed in any
Globorotalia cavernula Be, Globorotalia pumilio other planktic foraminiferal species. Ref.: Fleisher
Parker, and Globoquadrina hexagona Natland. (1974), Li (1987), Brummer (1988).
However, although E. Boltovskoy (1981b) included • Globigerina bulloides (Fig. 6.3). Large umbilical
them in his work in anticipation of future potential aperture without lip. Surface rather rough from
records in this area, most of these have still been only numerous spine collars. Aposymbiotic. Ref.:
recorded in the Indo-Pacific (some became extinct in Boltovskoy (1973), Febvre-Chevalier (1971),
the Atlantic during the Pleistocene, like Globoquadrina Malmgren and Kennett ( 1976, 1977, 1978),
cong/omerata, Globoquadrina hexagona), for which Kennett (1976), Kahn and Williams (1981),
reason they are not included here. Also two serial (hi- Caulet et al. (1992), Thiede and Jiinger (1992),
and tri-serial) planktic Foraminifera: Gallitellia vivans Puechmaille (1994 ), Kroon and Darling ( 1995),
and Streptochilus globulosus present elsewhere (Indian Ortiz et al. (1995), Rohling et a/. ( 1995), Gupta
Ocean, Pacific and North Atlantic) have not yet been and Mohan (1996), Spero and Lea (1996),
recorded from the South Atlantic. Their small size may Schiebel et al. (1997).
allow them to pass through meshes over 100 J..Lill. In any • Globigerina falconensis (Fig. 6.4). Spinose,
case, both species invariably occur in very low num- closely related to Globigerina bulloides, but with
bers (below 1% of the total standing crop). symbionts. Adults differ from Globigerina bul-
loides in smaller size, slightly elongated last
Detailed descriptions of these species can be found in chamber, smaller aperture, and distinct apertural
Loeblich and Tappan (1988), and in Saito et a/. lip. Ref.: Malmgren and Kennett (1977), Ortiz et
( 1981 ). E. Boltovskoy (1981 b) provides good sketch- a/. (1995).
es and a few SEM micrographs of most species that • Globigerinella calida (Fig. 6.5). Young individuals
occur in the area. difficult to distinguish from Globigerinella sipho-
nifera (similar spinosity and general morphology).
Species descriptions and remarks However, adults trochospiral and last 2-3 chambers
Table I presents the most stable characters for identi- become elongated. Ref.: Parker (1962).
fying the species covered in this review. Additional • Globigerinella siphonifera (Fig. 6.6) [=Globi-
comments on diagnoses are offered below. For each gerinella aequilatera/is]. Spinose, carnivorous
species, one or more references ("Ref.") provide during pre-adult and adult stages. Symbionts are
detailed descriptions, illustrations, and complementary chrysophytes and either aligned along spines or
distributional, ecological, paleoecological and biologi- evenly distributed within rhizopodial (pseudopo-
cal information. Common synonyms of some species dia}) network. Two known formae with slightly
are included in square brackets after the species name. different morphology involving more open or
closed (evolute or involute) coiling of last whorl.
• Candeina nitida (Fig. 6.1 ). Microperforate spe- Spines similar in both varieties: round and triangu-
cies with large proloculus (26 f.!m), smooth surface lar. Often placed in genus Hastigerina, which
and distinct sutural multi-apertures which are however, differs from Globigerinella siphonifera
unique among extant planktic species. Ref.: in wall structure, spinosity, cytoplasm, and
Brummer (1988), Sverdlove and Be (1985). absence of symbionts (see remarks for
62 Foraminifera
Hastigerina). Ref.: Beet al. (1973), Deuser et al. wide open in warm, saline environments, to slit-
( 1981 ), Caron et al. ( 1987b), Huber et al. ( 1997). like in salinities below 35%o. Typical for this spe-
• Globigerinita glutinata (Fig. 6. 7). Non-spinose, cies is the honeycomb (hexagonal) surface struc-
microperforate, trochospiral, with rather smooth ture, which is lacking in other spinose species (but
surface with irregularly distributed micropores. does occur in a non-spinose genus: Globo-
Small pustules also scattered over entire surface quadrina hexagona, from Indo-Pacific). Last
which, if well preserved, is whitish shiny. chamber sacciform, elongated. Ref.: Hecht ( 1974 ),
Aperture umbilical in adult specimens, often co- Be ( 1980), Be et al. (1981 ), Caron et al. (1981,
vered by a bulla. Ref.: Parker (1962), Kahn and 1987a), Duplessy et al. (1981 ), Brummer et al.
Williams (1981 ), Li (1987), Spindler et al. (1984), (1987), Hemleben et al. (1987), Bijma et al.
Brummer ( 1988), Ortiz et al. ( 1995). ( 1990a, 1990b, 1992, 1994 ), Spero and Lea
• Globigerinita uvula (Fig. 6.8) [=Globigerinita (1993). Remark: following widespread usage, we
bradyi]. Non-spinose, very small, microperforate, retain the present name for this species, rather than
readily recognizable by high trochospiral test, Globigerinoides quadrilobatus or Globigerinoides
somewhat like grape cluster. Surface structure trilobus. However, in the distributional data set,
resembles that of Globigerinita glutinata. Ref.: authors used the name Globigerinoides trilobus
Parker ( 1962), Li (1987), Saito et al. (1981 ), sacculifer for Globigerinoides sacculifer with a
Brummer (1988). sack-like terminal chamber, and Globigerinoides
• Globigerinoides conglobatus (Fig. 6.9). Spinose, trilobus trilobus for those devoid of this trait.
trochospiral, with 4 subspherical chambers in last • Globorotalia crassaformis (Fig. 6.13)
whorl, aperture umbilical. Usually 2-3 apertures [=Globorotalia punctulata]. Trochospiral, non-
on spiral side. Sutures rather deep. Adult speci- spinose, with 4-5 chambers in last whorl, rather
mens thick-walled because of additional calcite open umbilicus and usually calcite crust. Spiral
secreted when they migrate to deeper water. side flat, apertural side lower than in Globorotalia
• Globigerinoides ruber (Fig. 6.1 0, 6.11 ). truncatulinoides (more conical in latter). Aperture
Trochospiral, with 3 chambers in last whorl, aper- extraumbilical, no lip. Periphery rounded and only
ture umbilical over penultimate and antepenulti- slightly keeled on last and penultimate chamber in
mate chambers. On spiral side two smaller secon- adult specimens. Resembles Globorotalia injlata,
dary apertures visible. Two varieties (phenotypes): but with more angular outline. Ref.: Parker (1962),
white and pink forms (forma alba and forma rosa- Parker and Berger ( 1971 ), Lidz ( 1972), Rogl
cea), pink form only in the Atlantic. In the Indo- (1974), Zougary-Jaadi and Pujol (1987).
Pacific the pink variety became extinct during Globorotalia hirsuta (Fig. 6.14). Non-spinose,
Termination II (approximately 127 ky ago). White with 4 chambers in last whorl and scattered pus-
variety on average approximately 50 J..UTI smaller tules on older chambers. Usually rather smooth-
than pink. Reddish color from pigment mostly on walled; aperture extraumbilical - peripheral. Low
wall of inner whorl. Last chamber always whitish, trochospiral, bordered by distinct peripheral keel.
in contrast to Globoturborotalita rubescens whose Resembles Globorotalia scitula, but with fewer
entire test is reddish. Both varieties of chambers, contour more lobulate. Ref.: Glac;on et
Globigerinoides ruber have symbionts, indicating a al. (1973), Boltovskoy (1974a), Hemleben et al.
near-surface habitat. Ref.: Glac;on and Sigal ( 1969), ( 1985).
Yergnaud-Grazzini et al. (1973), Hecht (1974), • Globorotalia injlata (Fig. 6.15). Non-spinose, tro-
Kennett (1976), Brummer et al. (1987), Brummer chospiral, with 3-4 chambers in last whorl, spiral
and Kroon ( 1988), Gastrich and Bartha ( 1988), side flat, chambers on apertural side subspherical,
Hemleben et al. (1989), Oberhiinsli et al. ( 1992), aperture slit-like reaching from periphery towards
Kemle-von Miicke (1994), Kroon and Darling the umbilicus. Wall structure of different phases,
(1995), Ortiz et al. (1995), Wang et al. (1995). observed only in this species and in Pulleniatina
• Globigerinoides sacculifer (Fig. 6.12). Trocho- obliquiloculata. Growth of wall begins with bila-
spiral, with 3.5-4 chambers in last whorl (adults). mellar construction followed by secretion of addi-
Aperture bordered by small rim, may be slightly tional calcite layers on outer surface of test, includ-
extraumbilical. Height of aperture varies from ing formation of pustules over entire surface. When
Shell Chambers Aperture
Coiling
Coiling Pores Secondary
(A: Adult; (m: microperf.; Max.
direction Surface No. in Primary Additional apertures
P: planispiral; Spines Margin length Shape
Species S: streptospiral;
(R: right; L: texture n: normal
perforated) (mm)
last aperture structures
per chamber
whorl
left)
T: trochospiral)
): umbilical;
Candeina nitida T R very smooth m no rounded 0.75 globose 3 rimmed sutural
A: sutural
Dentagloborotalia
T low R&L smooth n no rounded 0.25 subquadrate 4.5-5 extraumbilical lip none
anfracta
medium to large,
Globigerina bulloides T low R&L n thin, circular rounded 0.8 globose 4 none none
rough umbilical
Globigerina large,
T low R&L rough n thin, circular rounded 0.45 globose 4 lip none
falconensis umbilical
circular to equatorial
Globigerinella calida T/P R&L rough n rounded 0.8 globose 4-6 none none
triangular extraumbilical
Globigerinella circular to
T, almost P R&L rough n rounded 0.9 globose 5-6 equatorial thin rim none
siphonifera triangular
Smooth to umbilical,
large lip,
Globigerinita glutinata Tlow R&L finely m no rounded 0.5 globose 4 often none
bulla
oustulose infralaminal
Smooth to umbilical,
lip may be
61
~
Globigerinita uvula Thigh R&L finely m no rounded 0.2 globose 4 sometimes none
present
oustulose infralaminal
Globigerinoides circular to Globose, last s·
~
Tlow R&L rough n rounded 1 3.5-4 umbilical rimmed 2
comdobatus subtriangular 3 compressed
medium
n circular rounded 0.8 globose 3 umbilical rimmed 2 ~
Globigerinoides ruber T low R&L
rough
Globigerinoides circular to globose, last
T low R&L honeycomb n rounded 1.3 3.5-4 umbilical rimmed 1
sacculifer subtriangular may be
Globorotalia smooth to angular,
T planoconvex L mostly n no keeled 0.65 4-5 extraumbilical lip none
crassaformis pustulose conical
T compressed, smooth to angular,
Globorotalia hirsuta R n no keeled 1 4-5 extraumbilical lip none
biconvex pustulose rhomboidal
Smooth,
T, spiral side flat, apert. area globose,
Globorotalia inflata L n no rounded 0.65 3.5 extraumbilical rimmed none
apert. side conical may be hemispherical
oustulose
elongate from
smooth to angular
Globorotalia menardii T low, flat L n no keeled 1.5 5-6 umbilicus to lip none
pustulose rhomboid
periphery
elongate from
T, compressed smooth to angular
Globorotalia scitula L& R n no rounded 0.65 5-6 umbilicus to lip none
biconvex pustulose rhomboid
periphery
Globorotalia T, spiral side flat, smooth to angular
L& R n no keeled 0.9 5-5.5 extraumbilical lip none
truncatulinoides apert. side conical pustulose conical
smooth to
angular
Globorotalia tumida Tlow L coarsely n no keeled 1.4 5-6 extraumbilical lip none
rhomboid 0'\
crvstalline ! (.;.)
Shell Chambers Aperture
~
Coiling
Coiling Pores
(A: Adult; (m: microperf.; Max. Secondary
P: planispiral; direction Surface
n: normal Spines Margin length Shape
No. in Primary Additional apertures
Species S: streptospiral;
(R: right; l: texture
perforated) (mm)
last aperture structures
per chamber
left) whorl
T: trochospiral)
angular
Globorotalia ungulata T L smooth n no keeled 1.2 5-6 extraumbilical lip none
rhomboid
Globoturborotalita
Tlow R&L rough n Thin, circular rounded 0.25 globose 4-5 umbilical lip n'one
rubescens
Globoturborotalita thin,
Tlow R&l rough n rounded 0.36 globose 4 umbilical rimmed 1
tenella trian~ular
equatorial to
Hastigerina digitata p smooth n triradiate rounded 5 globose 6 rimmed none
spiroumbilical
Hasti1:erina pela1:ica p smooth n triradiate rounded 1.2 hemispherical 6 equatorial thin lip none
Neogloboquadrina rough to umbilical
T Rmostly n no rounded 0.7 subglobose 4-6 umbilical none
dutertrei honeycomb teeth
Neogloboquadrina
Tlow R pustulose n no rounded 0.5 subglobose 4-5 extraumbilical lip none
incompta
Neogloboquadrina subglobose to
Tlow L pustulose n no rounded 0.5 4 extraumbilical lip none
~
pachyderma subquadrate
large multiple 0-1 (in
~
smooth to circular to
Orbulina universa j: T; A: spherical n rounded 1 spherical 1 11
pores 11 none juven iles
rough triangular
=apertures onlv) s·
~
triangular to
Orcadia riedeli T R&L smooth n rounded 0.2 globose 5 umbilical lip none
rough to
triradiate
hemispherical,
;
Pulleniatina umbilical to
5 R mostly smooth, n no rounded 0.8 later 3.5-4.5 rimmed none
obliquiloculata extraumbilical
shiny overlapping
Sphaeroidinella globose with extraumbilical, one
T ovate R&L smooth n circular rounded 1.3 nanges
4 rimmed
dehiscens slit-like (inconspicuo
smooth to
umbilica l to
Tenuite/la iota Tlow R&L finely m no rounded 0.22 globose 4-5 thin lip none
extraumbilica l
pustulose
smooth,
apert. area globose, umbilical to
Tenuitella parkerae Tlow R&L m no rounded 0.16 5 lip none
with fine last elongate extraumbilical
pustules
finely
Turborotalita clarkei Tlow R&L pustulose to n thin, circular rounded 0.15 angular 5 umbilical rimmed none
rough
finely extraumbilical,
Turborotalita humilis Tlow R&l pustulose to n thin, circular rounded 0.2 angular 6-8 sometimes rimmed none
rou~h infralaminal
extraumbilical,
Turborotalita smooth to globose,
Tlow R&L n thin, circular rounded 0.3 5 sometimes lip none
quinque/oba rough last expanded
-- ---
infralaminal
Foraminifera 65
individual migrates to deeper water (below the ther- a/. (1985), Pharr and Williams (1987), Lazarus et
mocline) calcite crust may form, subsequently al. (1995), Ravelo and Fairbanks (1995).
smoothed by thin calcite veneer, thus giving test • Globorotalia tumida (Fig. 6.19). Non-spinose,
shiny reflective appearance when viewed under low trochospiral, closely related to Globorotalia
binocular microscope. No keel. Ref.: Huang ( 1981 ), menardii, yet higher- and thicker-shelled than lat-
Ganssen (1983), Malmgren and Kennett (1983), ter. Last two chambers often twisted. Keel, lip and
Hemleben et a/. (1985), Ravelo and Fairbanks aperture prominent (as in Globorotalia menardii).
(1995), Rohling eta/. (1995). Both species may produce thick calcite crust. Ref. :
• Globorotalia menardii (Fig. 6.16) [=Globorotalia Malmgren et al. (1983), Ravelo and Fairbanks
cultrata] . Non-spinose, very low trochospiral, (1995).
large (> l mm), flat on both sides, with 5-6 cham- • Globorotalia ungulata (Fig. 6.20). Non-spinose,
bers in last whorl. With prominent keel (may similar to Globorotalia menardii, but with keel
be fimbriated), and broad lip over umbilical- band on crest of last chamber and fairly open
extraurnbilical aperture. Sutures straight on aper- umbilicus. Also resembles Globorotalia tumida,
tural side, curved on spiral side. Pustules thick on but walls thinner and smoother, apertural side
older chambers and in front of aperture, smaller more convex and angular. Ref.: Bermudez (1960).
towards periphery and on younger chambers. • Globoturborotalita rubescens (Fig. 6.21)
Calcite crust often visible, growing like pustules [=Globigerina rubescens ]. Spinose, with 4 globu-
from older to younger parts of test. Ref.: Scott lar chambers in last whorl, surface similar to that
(1973), Adelseck (1975), Hemleben et al. (1977), of Globigerinoides ruber. In Holocene samples
Bouvier-Sournagnac and Duplessy ( 1985), Healy- easily distinguished by reddish color throughout
Williams et a/. ( 1985), Schweitzer and Lohman entire test. Ref.: Parker ( 1976), Vincent (1976).
( 1991 ), Ravelo and Fairbanks (1995). • Globoturborotalita tenella (Fig. 6.22) [=Globi-
Globorotalia scitula (Fig. 6.17). Non-spinose, low gerinoides tenellus]. Very similar to Globotur-
trochospiral, with smooth surface, no keel and few borotalita rubescens, except for lack of red pigmen-
pustules on apertural side close to aperture. tation. Smaller than Globigerinoides ruber white
Umbilicus moderately deep, aperture stretching form, rather thick shelled. Surface structure more or
from umbilicus toward periphery, bordered by less cancellate and rather rough. Ref. : Parker (1958).
small lip. Resembles Globorotalia hirsuta, but • Hastigerina digitata (Fig. 6.23). Test transparent,
surface smoother, outline more rounded. Ref.: chambers claviform. Very similar to Hastigerina
Baumfalk eta/. (1987). pelagica, but with elongate chambers in last whorl
Globorotalia truncatulinoides (Fig. 6.18). Non- (see Hastigerina pelagica for additional remarks).
spinose, high trochospiral on the umbilical side, • Hastigerina pelagica (Fig. 6.24). Spinose, sym-
with slit-like aperture along base of last chamber biont-barren, with unique monolamellar wall. With
provided with broad lip. Spiral side flat, showing large triradiate spines with hooks on edges. Unique
total spire (if not encrusted by pustules). Pustule cytoplasmic bubble capsule surrounding test.
growth in clear gradient from older to younger Almost exclusively carnivorous. With well timed
chambers. Umbilicus fairly open and keeled lunar reproductive cycle, resorbing septa while
around periphery from neanic stage onward. In undergoing gametogenesis. Characteristically with
deeper water (>800 m) with rather thick calcite very low fossilization potential because of thin
crust. Last chamber wall, however, may be very walls (Spindler eta/. , 1979; Hemleben eta/., 1979;
thin from secretion in deeper water. Ref.: Be and Hemleben et a/., 1989). Ref.: Alldredge and Jones
Ericson (1963), Gla.;on and Sigal (1969), Rogl (1973 ), Hem! eben et a/. ( 1979), Spindler et a/.
(1974), Kennett (1976), Healy-Williams (1983), ( 1978, 1979).
Lohmann (1983, 1992), Lohmann and Malmgren • Neogloboquadrina dutertrei (Fig. 6.25)
(1983), Healy-Williams et al. (1985), Hemleben et [=Globoquadrina dutertrei]. Non-spinose, with
Table 1. Diagnostic characters of planktic foraminiferal species from the South Atlantic. Based partly on Be (1967) and E. Boltovskoy ( 198 1b),
with additions and emendations.
66 Foraminifera
4.5 to 7 subglobular chambers in last whorl. • Orcadia riedeli (Fig. 6.29) [=Hastigerinella riede-
Surface texture determined by regular pore pat- li]. Spinose, cosmopolitan but very rare. Small (less
tern. Last normal chamber usually broader than than 200 f.U11), trochospiral, with round as well as tri-
previous ones, positioned on top of first and penul- radiate spines. Wall thin, aperture opening towards
timate chamber of last whorl. Kummer chambers the umbilicus. Ref.: Boltovskoy and Watanabe
often present. Gametogenic calcification occurs (1981), Holmes (1984), Brummer et al. (1988).
regularly, calcite crust developed below 15°C. • Pulleniatina obliquiloculata (Fig. 6.30). Non-spi-
Ref.: Kennett (1976), Bouvier-Soumagnac and nose, wall structure similar to that of Globorotalia
Duplessy (1985), Srinivasan and Kennett (1976), injlata, although coiling is streptospiral, rather than
Thiede and Junger ( 1992), Kroon and Darling simply trochospiral. In deeper water with calcite
(1995), Ravelo and Fairbanks (1995), Ortiz et al. crust which becomes smooth by addition of veneer
(1995), Boltovskoy eta/. (1996). of very small crystals. Diet comprises more chry-
• Neogloboquadrina incompta (Fig. 6.26) sophytes than diatoms. Reproductive behavior sug-
[=Neogloboquadrina pachyderma, right coiling; gests month-long life span including gametogenic
Globoquadrina pachyderma, right coiling]. This calcification. Juveniles resemble Neoglobo-
species and Neogloboquadrina dutertrei seem to quadrina dutertrei. Ref.: Prell and Damuth (1978).
intergrade, although they are normally restricted • Sphaeroidinella dehiscens (Fig. 6.31 ). Spinose,
to different water masses. The non-spinose closely related to Globigerinoides saccu/ifer in
Neogloboquadrina incompta used here instead of adult morphology (Be and Hemleben, 1970), but
Neogloboquadrina pachyderma (right coiling) ontogenetic sequence (Huang, 1981) does not sup-
because of wall structure (pustules on surface, no port close linkage between these species. In pre-
ridge growth), and distribution pattern (preferen- adult stage Sphaeroidinella dehiscens distin-
tially in temperate to warm waters), thus different guished from G/obigerinoides sacculifer by very
from Neogloboquadrina pachyderma which usu- narrow aperture and calcite crust which accompa-
ally builds calcite crust. In the literature, nies shedding of spines prior to migration to deep-
Neogloboquadrina incompta is often described as er water. Thick, smooth, glassy calcite layer cov-
Neogloboquadrina pachyderma right coiling, as ers entire shell. Ref.: Bolli et al. (1985).
well as forms which intengrate into • Tenuitella iota (Fig. 6.32) [=Globigerinita iota].
Neogloboquadrina dutertrei. Ref.: Cifelli (1961 , Microperforate, non-spinose, almost planispiral
1973), Oberhiinsli eta/. (1992). during juvenile stage, aperture equatorial. Adults
• Neogloboquadrina pachyderma (Fig. 6.27) low trochospiral, aperture displaced towards aper-
[=Globoquadrina pachyderma, left coiling]. Non- tural side, small pustules scattered over test. Bulla
spinose, usually with 4 chambers in last whorl, may form in final stage. Resembles Tenuitella
general outline almost square. Aperture very nar- parkerae , but with fewer chambers, Jess com-
row, surface usually covered by heavy calcite pressed shell, with bulla, last chamber not lobu-
crust. Ref.: Bandy (1959), Bandy and Theyer late. Ref. : Cifelli and Smith (1970), Li (1987),
(1971), Malmgren and Kennett (1972), Vella Brummer ( 1988), Parker ( 1962).
(1974), Vilks (1975), Olsson (1976), Thompson • Tenuitella parkerae (Fig. 6.33) . Microperforate,
and Shackleton (1980), Healy-Williams et a/. ontogeny morphologically similar to those of
(1983), Reynolds and Thunell (1986), Donner and Tenuitella iota and Globigerinita glutinata.
Wefer (1994), Niirnberg (1995), Berberich (1996). However, adult stage without bulla, with more
• Orbulina universa (Fig. 6.28). Easy to distinguish elongate chambers, less globular than G/obigerinita
by last chamber: sphere encompassing weakly cal- glutinata. Ref.: Bronnimann and Resig (1971), Li
cified small trochospire which is mostly resorbed (1987), Brummer (1988), Saito et al. (1981).
during gametogenesis. Spines round and triangu- • Turborotalita clarkei (Fig. 6.34). Very small, spi-
lar. Adults feed mainly on calanoid copepods and nose. In pre-adult stage difficult to distinguish
other animal prey. With dinoflagellate symbionts. from Turborotalita quinqueloba. Also resembles
Ref.: Be et al. (1973), Vilks and Walker (1974), Turborotalita humilis, but with fewer chambers,
Weiner (1975), Caron et al. (1987b). lacking ventral lobulate expansion. Adult stage
Foraminifera 67
may be heavily encrusted in deeper water. Ref.: Be A.W.H., Tolderlund D.S. 1971. Distribution and
Brummer and Kroon (1988). ecology of living planktonic Foraminifera in sur-
• Turborotalita humilis (Fig. 6.35) [=Globigerina face waters of the Atlantic and Indian Oceans. In
humilis, Globigerina cristata. Spinose. Specimens "The micropalaeontology of oceans" (W.R.
from sediments spineless, with thick calcite crust Riedel, B.M. Funnell, eds.), Cambridge Univ.
(this feature also common to other species, such as Press., Cambridge, pp. 105-149. Although some-
Sphaeroidinella dehiscens or Turborotalita what outdated, still one of the best references for
clarkei). Spines mostly peripherical because of the geographic distribution and abundance of
very low trochospiral morphology. Symbionts are planktic foraminifers in the Atlantic Ocean.
chrysophytes, as in Globigerinella siphonifera. Boltovskoy E. 1981. Foraminifera. In "Atlas del
Ref.: Holmes (1984). Zooplancton del Athintico Sudoccidental y meto-
• Turborotalita quinqueloba (Fig. 6.36) [=Giobi- dos de trabajo con el zooplancton marino" (D.
gerina quinqueloba]. Small, spinose, low trocho- Boltovskoy, ed.), Public. Esp. lost. Nac. lnv.
spiral, aperture extraumbilical and with small to Desarrollo Pesq., Mar del Plata, pp. 317-352.
broad lip. Last chamber slightly elongate and cov- General review summarizing identification and
ering umbilicus. During late ontogenetic stages, geographic distribution of planktic Foraminifera
shortly before gametogenesis, calcite crust may be in Southwestern Atlantic waters.
added on top of normal bilamellar wall. In this Boltovskoy E., Wright R. 1976. Recent Foraminifera.
case spines are drastically reduced in number. Junk, The Hague, pp. 1-515. Textbook covering all
With dinoflagellate symbionts. Feeds mostly on aspects of the study offoraminifers , including the
algae and small zooplankton. Ref.: Asano et a/. benthic species.
(1968), Kroon et al. (1988). Hemleben C., Spindler M ., Anderson O.R. 1989.
Modern planktonic Foraminifera. Springer Verlag,
New York, pp. 1-363. Detailed review of the tax-
Acknowledgements onomy, biology, ecology, cytology and methods of
research ofplanktic foraminifers.
To Ingrid Breitinger, Horst Hiitterman and Dorothe Loeblich A.R., Tappan H. 1984. foraminiferal genera
Miihlen who were involved in this work at various and their classification. Van Nostrand Reinhold,
stages; to Barbara Donner who supplied SEM speci- New York, pp. 1-970. Guide to the general clas-
mens, and to Silvia Watanabe for useful suggestions sification of the Foraminifera.
and provision of several of the SEM photos. Parker F.L. 1962. Planktonic foraminiferal species in
Organization of sections in this chapter follows the Pacific sediments. Micropaleontology, 8:219-254.
editor's request in order to generally agree with those Useful guide for the identification ofmost planktic
adopted throughout the volume. Foraminifera, with excellent line-drawings.
Saito T., Thompson P.R. , Breger D. 1981. Systematic
Index of Recent and Pleistocene Planktonic
Suggested readings Foraminifera. Univ. Tokyo Press, Tokyo, pp. 1-
190. Nicely illustrated guide including all Recent
Be A.W.H. 1967. Foraminifera: families Globigerinidae planktic Foraminifera, with synonymies, diagnos-
and Globorotaliidae. Fiches d'Identif. Zoopl., Cons. es, and distributional remarks.
Perm. Int. Explor. Mer, 108, pp. 1-9. General iden- Vincent E., Berger W.H. 1981. Planktonic foraminife-
tification guide for Recent planktonic foraminifers. ra and their use in paleoceanography. In "The
Be A.W.H. 1977. An ecological, zoogeographic and Oceanic Lithosphere", The Sea (7) (C. Emiliani,
taxonomic review of Recent planktonic Foramini- ed), Wiley, New York, pp. 1025-1119. Comprehen-
fera. In "Oceanic Micropaleontology" (A.T.S. sive review ofpaleontological and paleoecological
Ramsay, ed.), Acad. Press, London, pp. 1-100. aspects offoraminiferal research.
Excellent review of the identification and distribu-
tion ofplanktic foraminifers from the World Ocean.
68 Foraminifera
Foraminifera 69
70 Foraminifera
Foraminifera 71
72 Foraminifera
Foraminifera 73
Fig. 6. Illustrations of the species. All scale bars are 50 f!m, unless otherwise noted.
Figure sources:
From E. Boltovskoy (l974a): 6.4a; from E. Boltovskoy (198lb): 6.8b; from Boltovskoy and Watanabe (1980): 634a; !rom Boltovskoy and
Watanabe (1986): 6.4b, 6.8c, 6.32b, 6.34b; from Saito eta/. (1981): 6.23; courtesy S. Watanabe: 6.8a. 6.32a. 6.34c. 6.35a. 6.35b. 6.35c; all oth-
ers original.
Acantharia
Renate Bernstein, Stanley A. Kling and Demetrio Boltovskoy
Introduction . copepodids and adults, pelagic molluscs, and other

sarcodine taxa (Caron and Swanberg, 1990).
Acantharia are delicate, free living, microphagic, Examination of cells under epifluorescent illumina-
exclusively marine microzooplanktonic protozoa. tion (blue excitation, red and orange emission) reveals
They are generally spherical, oblate or sometimes signs of microalgal prey including picoplankton sized
flattened, ranging in size from 0.05-5 mm in diameter. «2 urn) cyanobacteria (Michaels, pers. comm.).
Small bacteria can also be observed within food vac-
Acantharia share a feature with Radiolaria and the uoles with the appropriate staining techniques; how-
closely related group Heliozoa: the axopodia. These ever, it is unclear if they are actively consumed or
are long, radiating, unramified processes stiffened by accidentally incorporated with larger food. There are
patterned arrays of microtubules (Febvre-Chevalier no published quantitative estimates of grazing rates.
and Febvre, 1993). For this reason, all these taxa are
placed in the Class Actinopoda. Many species contain symbiotic algae generally
known as zooxanthellae. According to Febvre et al.
Distinguishing features are: (I) the characteristic (in press), in most families they are located in the
skeleton of strontium sulfate, based on a general plan endoplasm, although Schewiakoff (1926) reports that
of 10 diametral or 20 radial spicules arranged in a re- they are usually in the ectoplasm. In natural assem-
gular pattern called Muller's law (Muller, 1859; see blages of Acantharia, approximately half of the indi-
below and Fig. I); (2) division of the cell body into viduals have symbionts and the smallest and largest
two parts, a centrally located endoplasm (central cap- size classes have the lowest proportions of symbionts
sule), separated from the ectoplasm by the central cap- (Michaels 1988a, 1988b, 1991). Symbionts are found
sule membrane; the ectoplasm in many species further in all members of the order Arthracanthida (Febvre
subdivided by a distinctive outer layer termed the per- and Febvre-Chevalier, 1979; Febvre, 1990) at some
iplasmic or ectoplasmic cortex (Fig. 2); (3) the myo- part of the life cycle. Within a species, young tro-
nemes, bundles of contractile filaments resembling phonts and gamonts lack symbionts, and symbiont
umbrella ribs grouped around the spicules where they numbers appear to increase with size in the trophont
protrude from the peripheral cell body (Fig. 2, 6.19b). stage. Symbionts seem to be shed or consumed imme-
diately prior to gametogenesis. Acantharian sym-
Reproduction and the life cycle are incompletely bionts are of diverse biologic origins, including dino-
known. There is no evidence for asexual reproduction, flagellates and haptophytes, and some species have
although it was previously reported in one genus more than one symbiont type within the same host.
(Schewiakoff, 1926). Sexual reproduction takes place
in a gamont which maintains the aspect of the tro- Symbiont abundances vary within a host from a few
phont (vegetative cells), or in a cyst (gamontocyst; large dinoflagellates in some species, to many thou-
Fig. 6.127-6.158). Encystment involves a complete sands of smaller symbionts in others. Most Acantharia
reorganization of the mineral skeleton and a series of have symbionts at some stage in their life cycle,
mitotic and meiotic fissions. Thousands of biflagellate although in heterogeneous natural populations, general-
isogametes are formed in the gamonts and gamonto- ly about half of the individual Acantharia are aposym-
cysts, and shed within minutes. Further steps to the biotic (Michaels, 1991). These aposymbiotic indivi-
formation of the young aca~tharian remain unknown. duals are a mix of juveniles and near-reproductive
adults (which lose the symbionts). Symbiont reproduc-
Acantharia consume a wide variety of prey. Reports tion is likely under host control as the slow symbiont
from microscope observations of feeding vacuoles doubling times inferred by cell division (very few of
describe the remains of tintinnids and other types of the cells are seen in mitosis at anyone time) are much
ciliates, diatoms, dinoflagellates, copepod nauplii, lower than the rapid growth that could be inferred from

r----- ...
--.--
..--
76 Acantharia
c······nmh ...hh::.:-> polar (4 spicules)
'., tropical
'. (4 spicules) ;.,
I';~ tropical
0° -------. ~----.
equatorial
(2 spicules)
2700 ••• _= r/
'-•.~" ~900
.
spicules
"'<'\'~
tropical -, polar
spicules
225° 135°
<-"'-,_~_~_~ polar
18~ eq.uatorial
spicules
A SIDE VIEW POLAR VIEW
B
MODELS p: polar spicules p p

t: tropical spicules
e: equatorial spicules Acanthometra cylindrica
Fig.1. Spatial distribution of the 20 acantharian spicules according to Muller's law. A-D: original; E: from Haeckel (1887).
the carbon-specific rates of symbiont production (9.4 are most common, it is likely that nitrogen and phos-
ng C per ng symbiont C per day, Michaels, 1991). Most phorus are supplied to the association by the feeding
of the symbiont-fixed carbon is probably transferred to activity of the acantharian as dissolved nutrients are
the host, suggesting that symbiotic algae are an impor- present at very low concentrations. However, symbi-
tant part of the nutrition of Acantharia. These symbionts may play a role in retention of these nutrients
onts have extremely high saturation irradiances and can either by recycling through the host or by meeting the
continue to fix carbon even near the surface. This car- host energy demands and thus allowing a more effi-
I' bon can sometimes meet or exceed the inferred daily cient use of particulate nutrient material.
j metabolic requirements of the host for carbon
(Michaels, 1988a, 1988b, 1991; Caron et al., 1995). Acantharians also are thought to play an important
role in the budgets of strontium and other metallic ele-
Little is known about host-symbiont nitrogen dyna- ments in the oceans. Prior to the 1950's, dissolved
mics. In the oligotrophic settings in which Acantharia strontium (Sr) in seawater was considered a conserva-
Acantharia 77
_ axopod
~----------------==--=axoneme
filopods
myoneme
ectoplasmic
cortex
Eof> E'"
'" E
0..2-
ti '"
o u
OJ
, '
:- - - -~ - - - - - - - --
~------- nuclei
E digestive
of>
vacuoles
'"
0..
o
"c:
OJ
Fig. 2. Schematic cross section of an acantharian cell. From Reshetnjak (1981).
tive element, i.e. varying with salinity (e.g., Chow and ton induced X-ray ermssion analyses, Brass (1980)
Thompson, 1955). Using a flame photometric internal noticed a variety of trace elements - Ca, Mn, Ni, Zn,
standard technique, Mackenzie (1964) was the first to Pb, Sr, As and Br - in 10 acantharian skeletons. And
observe a significant increase in strontium to chlorinity although Arrhenius (1963) reported high concentra-
ratios (Sr/CI) at water depths of 500 to 800 meters, tions of barium (5400 ppm) in acantharian celestite
which he attributed to possible reaction with organic (SrS04), Brass (1980) found no barium in his study.
aggregates. More refined analytical methods (atomic
absorption and neutron activation techniques) estab- To date, low surface water Sr/CI ratios have been cor-
lished a trend in Sr/CI ratios in Pacific seawater sam- related with uptake of Sr by surface-dwelling acanthar-
ples, with significantly lower ratios in shallow waters ians. Conversely, comparatively high ratios (about 2-
from the eastern North Pacific than those from deeper 7% higher) in intermediate waters may be the result of
waters (Brass and Turekian, 1974). Subsequently, a rapid dissolution of sinking acantharian celestite debris
more sensitive technique using mass spectrometric iso- (Bernstein et al., 1987). To the extent that acantharians
tope dilution, demonstrated differences between affect seawater Sr/CI ratios, the trace metals reported in
Atlantic and Pacific Sr/CI ratios. In addition, Sr/CI their skeletons and cysts suggests that cycling of other
ratios in Pacific surface waters were shown to be oceanic elements may also be influenced. Molar ratios
approximately 2% lower than those in the deep Pacific. of Ba to Sr in acantharians have been determined by
Brass and Turekian (1974) suggested acantharians as a Arrhenius (1963) and Rieder et at. (1982) as 4 x 10-3.
possible mechanism for this depth trend. Based on pro- More recently, neutron activation analyses of Pacific
78 Acantharia
acantharian specimens yielded a (Bah/(Srh molar etal morphology almost exclusively for taxonomic dis-
ratio of (3.0±0.8) x 10-3 (Bernstein et al., 1992). These crimination. More recently, general descriptions of
molar ratios in acantharian celestite are approximately morphology and systematics, in some cases largely
10 times larger than the dissolved [Ba2+h/[Sr2+h compiled from Schewiakoff (1926), were produced by
molar ratios calculated for surface ocean waters Tregouboff (1953), Reshetnjak (1981), Cachon and
(Bruland, 1983). Consequently, sinking dead-acantha- Cachon (1985), and Febvre et al. (in press).
rian celestite may play a larger role in oceanic Ba Reshetnjak's (1981) monograph is particularly relevant
cycling than is the case for cycling of Sr. since it reviews all known acantharian species, includ-
ing distributional data, identification keys, and diagno-
To further test this hypothesis, individual acantharian ses. Present knowledge of the fine structure of these
organisms (skeletons and cysts) were analyzed for Sr protozoa, their life cycle, nuclear fission, endosymbio-
and Ba by inductively coupled plasma mass spectro- sis, and myoneme behavior are summarized in Febvre
metry (Bernstein et al., 1998). Specimens were (1990). A few modem ecological data are available,
obtained from four diverse areas of the world's including their abundance and distribution (Beers et al.,
oceans. Acantharian (Bah/(Srh molar ratios were 1975, 1982; Bishop et al., 1977, 1978, 1980; Massera
compared to dissolved Ba and Sr concentration ratios Bottazzi et aI., 1971; Massera Bottazzi and Andreoli,
from GEOSECS (Ostland et al., 1987) stations that 1974, 1982a, 1982b; Michaels, 1988a, 1988b, 1991;
corresponded most closely to the stations from which Michaels et al., 1995), their role in cycling carbon
the acantharians had been collected. Results showed through both photosynthesis by symbionts (Michaels,
that for 61 of the 64 specimens analyzed, (Bah/(Srh 1991; Caron et al., 1995), and their contributions to
molar ratios were significantly higher than those cal- sinking particles (Caron and Swanberg, 1990;
culated from the corresponding seawater samples. Michaels, 1991; Michaels et al., 1995).
Thus, acantharian particles exiting surface water
remove a larger fraction of Ba than Sr.
Methods
It is beyond the scope of this chapter to elaborate on
the thermodynamic principles supporting the data on The extreme solubility of the celestite skeletons of
enrichment of Ba in acantharian celestite relative to acantharians requires special collection and preserva-
BaiSr ratios in seawater (Bernstein et al., 1998). tion methods. Sampling is restricted to the water-co-
Nevertheless, it is important to point out that thermo- lunm, as they are not preserved in ocean bottom sedi-
dynamic principles and the reported inclusion of other ments. Solubility also prevents their retention in sea
trace metals in acantharian celestite (Michaels and water samples preserved by conventional methods:
Coale, unpublished data) suggest that acantharians addition of strontium to preserved samples is neces-
may also be important in the oceanic cycling of met- sary. With these provisos, various collecting tech-
als other than Sr and Ba. niques applicable to other delicate microplankton are
also appropriate for acantharians.
There is no direct evidence of the origin or geologic
history of the Acantharia. Since their skeleton dissolves Net samples can be collected by either simple, non-
rapidly in sea water after death, they do not fossilize. closing plankton nets (Massera Bottazzi and Andreoli
1974, 1982a, 1982b; Tan and Su, 1983), or more ela-
The Class Acantharia comprises around 50 genera and borate devices allowing multiple, discreet, vertically
:i
'I
150 species grouped into 18 families and 4 orders: stratified tows (Schreiber, 1960; Spindler and Beyer,
I Holacanthida, Symphyacanthida, Chaunacanthida and 1990). Nets have been employed in horizontal surface
Arthracanthida. The primary reference for the system- tows, as well as in oblique or vertical hauls from var-
I atics is the monograph by Schewiakoff (1926). This ying depths. Mesh sizes also vary, ranging from 25 to
I book contains a set of remarkable drawings and a clear >160 Jim, although sizes in the range of 160-350 urn
I and expansive description of acantharian taxonomy result in gross abundance underestimates. Estimates
based on accurate in vivo observations of 80 species. from nets are in the range of 1-500 indo m-3 (Massera
Schewiakoffs system takes the character of the cell Bottazzi et al., 1971; Massera Bottazzi and Andreoli,
body into account, whereas his predecessors used skel- 1974, 1982a, 1982b) compared to 1,000-50,000 m-3
I
I
I!
Acantharia 79
I based on pump or filtration methods (see below). This from Niskin bottles are similar to those from diver-col-
suggests that net sampling, especially with coarse lected samples; (2) plankton net samples (both 25 and
I meshes, is generally inadequate for these organisms 160 J.I.IIl mesh, and both horizontal and oblique tows)
as their small and fragile bodies get extruded through appear to capture far fewer specimens than do Niskin
I the gauze or destroyed beyond recognition in the sam- bottles; and (3) Niskin bottle specimens obtained from
ples (Michaels, 1988a, 1988b). the spout were more highly clumped than those
retrieved through the bottom valve. Possible explana-
Niskin bottles of 12-30 1 capacity at various intervals tions for acantharian underestimation by plankton nets
through the upper 160-200 meters of the water column include overestimation of the amount of water filtered,
have been employed successfully by Beers et al. (1975, damage to acantharian cells by the net, leading to dis-
1982), and Michaels et al. (1995). To minimize destruc- tortion and passage through the net, and net avoidance
tion of fragile acantharian cells during collection, by a buoyancy mechanism triggered by disturbance.
Graham et al. (1976) used a 1 liter polyethylene bag
attached to a spring-hinged sampling device controlled To concentrate acantharian cells, whole samples are
by a standard heavy-duty fishing rod and reel. The tech- often sieved through a series of screens separating the
nique was successful at depths from 1 to 300 m, and the material into discrete size classes (Beers et al. 1975,
number of acantharians collected was two orders of 1982; Bishop et al., 1977, 1978, 1980; Bernstein and
magnitude higher than those previously published. Betzer, 1991; Michaels et al., 1995), or allowed to set-
tle for 1-3 days and the supernatant fluid then gently
Pump sampling has been employed by several work- withdrawn (Michaels, 1991).
ers. Bishop et al. (1977, 1978, 1980) used large volume
in situ filtration systems to collect particulate matter Although Acantharia are often the most abundant
from 5-30 m3 of water from the upper 400 m. Water shelled sarcodines in the upper ocean (Caron and
was pumped at a rate of -6 m3 h-I through 53 J.I.IIl mesh Swanberg, 1990; Michaels, 1988a, 1988b, 1991;
screens, and then through acid-leached pre-combusted Michaels et al., 1995), their skeletons dissolve in tra-
glass filters to provide sample size fractions of 1 J.I.IIl, 1- ditional preservatives such as buffered formalin
53 J.I.IIl, and >53 J.I.IIl. Michaels et al. (1995) used a (Beers and Stewart, 1970), and thus tend to be missed
deck-mounted air-driven pump connected to a 200 m in assessments of microplankton abundance. Addition
long hose to collect water samples from various depths. of SrCl2 6H20 to fixed samples such that the final Sr
Canisters mounted close to the hose and near the pump concentration equals 80 mg 1-1has been widely adopt-
contained 300 and 20 J.I.IIl mesh Nitex socks to separate ed (Beers et al., 1975, 1982; Michaels, 1988b, 1991).
sample material into two size fractions. Higher Sr concentrations (160 to 240 mg 1-1) in for-
maldehyde-preserved samples, especially when buf-
Acantharian skeletons and cysts may be collected in fered with sodium borate, tend to produce a white pre-
sediment traps (Bernstein et al., 1987). Short deploy- cipitate and are therefore not recommended
ment times (on the order of24 hours), relatively shal- (Michaels, 1988b). Michaels (I 988b ) also concluded
low trapping depths, lack of preservatives that might that in samples with large numbers of acantharians,
enhance dissolution, and sample processing immedi- skeletal dissolution will raise the dissolved Sr concen-
ately after trap retrieval were instrumental in yielding trations and preclude further dissolution.
well-preserved organisms, although abundances are
still considered low (Bernstein and Betzer, 1991). Preservative formulations used in modem studies
Michaels (1991) used free-floating cylindrical traps include Bouin solution (Tan and So, 1983),2-3% for-
over periods of 54 days. Traps were charged with a malin buffered with borax or hexamethylenetetramine
salt gradient (NaCl, 50 g 1-1)to retain 10 ml I'! offor- (Massera Bottazzi et al., 1971), 5% formalin buffered
malin preservative and 72 mg 1-1 of Sr to maintain with hexamine and containing 72 mg 1-1Sr (Michaels,
acantharian integrity. 1991),2.5% gluteraldehyde with a sodium cacodylate
buffer and strontium chloride (80-160 mg l+)
Michaels (1988) compared the efficiency of various (Michaels, 1988b), 4% formaldehyde buffered with
acantharian collection and processing techniques to sodium borate, and strontium chloride (80-160 mg 1-1)
show that (1) acantharian density estimates obtained (Michaels, 1988b).
80 Acantharia
Filtration of samples onto Nuc1epore or Millipore fil- There are few time-series studies of Acantharia.
ters and subsequent drying is another cornmon tech- Schewiakoff (1926) described moderate seasonal
nique used to preserve skeletal and cystic material and changes in acantharian abundance and specific compo-
to facilitate examination using an electron microscope sition in the Mediterranean. During the winter some
(Bishop et al., 1977, 1978, 1980; Michaels, 1988b; species were observed to avoid the upper layer, sink-
Bernstein and Betzer, 1991), although it preserves the ing to depths of 50-200 m. In the South China Sea,
skeleton only. Reshetnjak (1981) recorded 23 species, 21 of which
were present during the surnmer only. At the VERTEX
Acantharians can be examined in wet samples under a time-series station in the northeast Pacific (33°N,
regular microscope or an inverted microscope. Larger 139°W) they tended to be more abundant in the sum-
specimens (> 100 Ilm) can also be counted under a dis- mer and fall, although their biomass did not show the
secting microscope. Best results are obtained with same pattem (Michaels, 1991). Integrated abundances
freshly collected, living specimens. Staining with were variable at the Bermuda Atlantic Time-series
boric carmine (Schreiber, 1960), or bengal rose (Beers Study site (32°N, 64°W) in the Sargasso Sea (Michaels
et al., 1975, 1982) may help visualize the soft parts. et al., 1995). Samples collected with plankton nets at a
Scanning electron microscopy (Bernstein et al., 1987, variety of locations throughout the Atlantic suggest
Bernstein and Betzer, 1991) is especially appropriate higher abundances in spring and lower abundances in
for high resolution of skeletal geometry. surnmer, although the data were not collected as an
explicit time-series (Massera Bottazzi and Andreoli,
1982a). In coastal areas, seasonal patterns are some-
Distribution times observed if oceanic advection displaces eutroph-
ic waters near the coast for a defined portion of the
Acantharia are cornmon in surface waters of tropical year. In the oligotrophic tropical and subtropical seas,
and subtropical oceans worldwide; in temperate and Acantharia are a regular, cornmon and conspicuous
polar seas they occur in much lower numbers. In very component of the microplankton year-round.
nearshore environments and coastal upwelling zones
they are usually scarcer. Reported acantharian abun- Because of difficulties associated with their collection,
dances vary widely, from <1 to 50 ind. 1-1 (Beers et al., preservation, and identification, data on acantharian
1975, 1982; Bishop et al., 1977, 1978, 1980; Michaels, geographic ranges are scarce. Available information,
1988a, 1988b, 1991; Michaels et al., 1995; surnmarized however, suggests that they are geographically (but not
in Caron and Swanberg, 1990). ecologically) cosmopolitan. With the exception of
Coleaspis obscura, which so far has been recorded
They are generally restricted to the illuminated upper only in high-Iatitude areas (Arctic and Antarctic), all
layer of the ocean, although some individual s have other species were found in tropical and/or subtropical
been collected in sediment traps thousands of meters waters (Reshetnjak, 1981; see Table 1). The number of
below the surface (Bernstein et al., 1987). On calm eurythermic species is comparatively low, probably
days larger Acantharia usually aggregate in the upper around 10-20 (Table 1), and most of their records in
few meters of the water colurnn, and may be concen- cold waters are associated with the influences ofwarm-
trated in Langmuir cells (windrows). This surface affin- water currents, like the GulfStream in the Atlantic, and
ity is likely related to the photosynthetic physiology of the Kuroshio in the Pacific (Reshetnjak, 1981).
their symbiotic algae. However, upon mechanical
(wind) or chemical (freshening of the surface layer by Despite low coverage of the ca. 150 known species
heavy rain) disturbance, their active buoyancy control (especially in the Indian and Pacific Oceans), 57 are
allows them to swiftly descend to lower strata (50-200 circumtropical and have been recorded in the 3 major
m), returning to the surface once adequate conditions oceans (Reshetnjak, 1981). Although the Atlantic (in
there have reestablished (Schewiakoff, 1926). During particular the Mediterranean) has been more inten-
periods of strong vertical mixing, their abundances sively sampled than the Indo-Pacific, few materials
become uniform with depth, but they tend to retum to come from the South Atlantic, which prec1udes draw-
the surface within a few days of restratification. ing conc1usions about their abundance and distribu-
tion in this basin. However, as noted above, judging
Acantharia 81
1---[
T
,
-
v
c:::
~
.-_.
"'C
~:.
QJ
i
I
'
i
V
t;:::
.-
V i
c:::
.~
"'C
V
'';;
V
~
< e <
=~--=l
-------"--- I
-- -----
----.--
Aconthaspis capil/ata
- •
•
-
Aconthaspis furcata
-- - ------
Aconthaspis hastata
• - ------1
Acanthochiasma fusiforme
• • • • • '-- --- •-----r i
Acanthochiasma hertwigi
•
- ----
•
-"
Acanthochiasma planum
'----" ---_._-- -- -------
Acanthochiasma quadrangulum
- --- I--- • - • • -- - • .. _-----,
• • • • • •
-"
Acanthochiasma rubescens
- ------
Acanthochiasma serrulatum
• • • • -_.-
Acanthocol/a cruciata
• • • • • •
•
---~
•
"-
Acanthocol/a solidissima
--
_~t::anth?_cr..rtha haeckeli
• • • • •-- •.....-.. --------
Acantholithium dicopum
• • • • ..
•
_---_.
Acantholithium stel/atum
• • • • • ----- -------
• • •
-
Acanthometra bulbosa
• • -------
Acanthometra cylindrica
r-------
Acanthometra dolichoscia
• •
•
• • • .---
---, I
I
Acanthometra Fusca
• •• ---- •
• • • • •
f------
• -I
,
Acanthometra
f---
pel/ucida
Acanthometra prismatica
• • ----------j
i
Acanthometra prismatica
f-- • • __ .1I
Acanthometra tetracopa
• • • •
--- • •
Acanthoplegma krohni
• • • • • • f--- •-------'
._---
Acanthospira spiralis
• • i
Acanthospira torta
•
Acanthostaurus bipennis
• • • --~
Acanthostaurus conacanthus
• • • • • i
Acanthostaurus henseni
-
Acanthostaurus purpurascens
•
• • • • • -----==l ----I
Amphiacon denticulatus
• • -- f--
I
Amphibelone anomala
• • • • •
Amphibelone cultellata
• • •
Amphibelone hydrotomica
• • • • • ------~I~
Amphibelone violina
• .-..
Amphilithium clavarium
-- -- • • • •• • ....~
• • • •
------
Amphilithium concretum
--------
Amphilonche diodon
• --- .. ------j
Amphilonche elongata
• • • • --1
I
i
Amphistaurus atlanticus
• • • • •
Amphistaurus
_--
.......
complanatus
• • • • • --- =J
---------------.--.---------
82 Acantharia
------ -_ .._-
I
U
'';:
'';: u c::: u
.~ I
(Ij
a.. a..
e
~
:E
U
.- ~ E
a.. Q.I
••• ~
•••
•••
U
I
.
e
""C
'; ~ ~
c::: -e
L -< Q..
---_.
~ ~ i____ ~~
lJ'\m~histaurus lanceolatus
Amphistaurus tetrapterus ._-
4---:--+--.-- •
1------+----+-
---- • ._-
•
I
• • • •
- -- ----- ---
Astrolithium bulbiferum
-_ _-------- .----
•
..
Astrolithium piriforme
-----
Astrolonche pectinata
1-----1 --- ..
-- • ·---1
• _.--- ..- ·--·----1'
Astrolonche pinnata •
. .!
--- ----+- ------- .----f-----.------
Astrolonche serrata
---_ ..
_---------- --!-----
•
• ----±--i
Coleaspis coronata
----- -_._._-+-----
•
•
----j-----j------t--- • --
·•
-----1
Coleaspis obscura •
• •
------- • +------.---- ---·--1
Coleaspis vaginata •
---_._---+-- ---+- .-----
----- J
Conacon foliaceus
--_.- -----
•
----+--- • • •
••
--- -- .. -----~
Coscinaspis isopora -----j-_._-
-----_.
Coscinaspis peripora •
-----.-- --.+-------1-------1-- ---
Cranisaspis parmipora •
-.---- -------+- --+--------
Derataspis polypora • , I
~~~ranophora_!!.uetschli --_-- _ .._.._._ 4-----e-------.- --t------+---------+--~-- _-~~_-:
Dic_t_yacanthatabu_'a_ta . • -+I • __ t--- __ •__ -j- -+-__ ._ _. __

. j
Dictyacantha tetragonopa •• ••
---j-----f-------+-----t----t--------j--- ---I ,
Dictyaspis furcata • ••• I
Dictyaspis solidissima ••••• I
~~~-:-~~:i-*-~~:
~_;!_E_f_':~_S-_-.••.= ....-_t---:---t---~~.:~~--~--~-=.~-----~+-- --.---~J
-.~~~::~~--:---.....,-f~-~--~.+-
[)jpl?.c_o_n_u_s
!'f!..~~p_h_y_'!_us___ . -+ --.--1-----t----------1-.- ..
-- ..--J
:--+--: : --=-=-
Diploconus saturnus •••••
~:~:~::~;:
f-~;-~-~t-~-·---
: :
!Dorataspis macropora---' ..-.--+---.----1--.--r.---.--+--.---t--
~:;::::~;::;;::;::: . .--;----.----+---.---r-----:---+---.--t--.----t----- ..... -~J

-------:
Gig~aCon abcisus
Gigartacon bitidus
Gigartacon co;:(Ji-fo-rm-i-s
--=~.
- -- -- -1---.- ••
--.~=--~~.===.==--:-----.--__+--.--t--.--_t_----~-------:
1-----
I'
-----+
--- .. ..--.--
• .
- -- ------- ------ -- - --j----- ._-- ----- ------ -----

~Gigartacon denticulatus • • • • • • •
!-Gigartaco~fragilis • --.- - -.----+----- • ---1--.----
I Gigartaconmuelleri ••••••• i
-r-_-_-
__
_ 1
f_H_-_~ii~~_
ma_tl_~m~nc_e'_'._at_um_
- -.- - --. ---.--t-----t-. --·---1
: :~~~~--:~-;m-£;eH:"
_ J-- -_:-_·-_1--.~
.•. _-_- -:_._-=E _=__ -_.
_--~--~=- ~;
Acantharia 83
u 0'u8
0';:
.~ u
1
s-
c E
tIl
10. 10. U
C
~ "'t:I !E ~ 10. QJ
J
U "'t:I
-;::; QJ
~ ~
-e ~ Q.. c ~
- - -~.-Cf=.---- ~===-::-:==*======~~=====-.;:::===~=-=
---_.----- __ ~
1---1-· :--.--
-- -
['::';;aconus c!!0tu~== __ -+-_
~Hexaconus echinatus - ---_ .. .
Hexaconus infundibulum
--t-.-----+----t--------+_---.-t----
Hexaconus nivalis
Hexaconus serratus
f------
1---- .....
--.
----
..
---
---------
Hexaconus trypanon
Hexalaspis heliodiscus
Hexalaspis sexalata
-.--
-...
----t--
---
...
.---t------
..
'--" -----+-
_
_
-----
_ _
...
- - -
-.----!---
_
-.---+------t--
_
_
- _
----j--
+----t--
...
_
--- ...
_
_
-
_
1-----+_--------+-----+
--'-- _. ---
---- -·-1
j
1-------- ....
---------. --j----t- ---j----j-- ---+-- --.....
--,
Hystrichaspis dorsata
Hystrichaspis rr-.'!_ti_c_a~~ --+_
Hystrichaspis serrata
.--. .--------t---
Icosaspis elegans
+------1-.------[------.
_
_
_
_
---+---- -----
_
_
~_
_
e__
___
_
_._.__ _ __ ._~
.. .1
~:~~: ;;~:ut~ia
Li!h_o_p~,!_~_e_n_es.tr_at_a
.. --_=-=_.
__-
__
~. __
. ---.--t--
+--------t--_-_-_=_~-_-_.-t
-
__
-._-
..-.
-
-·_1
- _
..._. J
~~~~~:~:::;~Se~;Zrc~-s----
--- .-.---j---
.~----+------~--- : .. -.-+---- -------.--.--------.-----.-~
.1----- -- ------1
Lonchostaurus rhombicus _ _ _ _ _
------- ----1- ..
----- ----. -j---- --+-
...
Lychnaspis giltschi - - - -_ - --_- ..-- .. -=J--.

Lychnaspis longissima _ _ _
I-- -- ---+------+
.. -+__ --j---_
-.---:~=--=--.+------~-~----.--+------- _._ -__'.'_'--,_~ :_==-- .-.-._:--.:._.~.~.-

..-._. ....
-.---
_~.~.-..~_--._-I'
i;§~ra-_~-i ~;-t:~-: · t.1
~.-~-~~-~-~-::~~:
:;:::: ----+- II!,
:T:_ :i ~__
~~~:~~t=1;-::-;-~:-b~--i~---- :__
~=~_~~~_u_~ I __
;~~:~i~-.~-::-:~-:-:-~-:-;-~-o~~;·~-_-~
: __ -.-....-... ~--+. _-_--~:--.-- _-~_i_-t_--_:_~-~:--·'--- ~-_-- -- ,
~hra_ct_o_pe_/~~~~_s~~aspis _. __~ ..._~_._ ~------~-t--~--- _ ...__
~~=f~::-:-5:-::-~~:---:-:-~-a-:~-r-:-a-:-t:-e-~/U5--
------ -------
--:-. .-::- ~-.
--.---l-------.J----..
m....!-t~-. . - _:----=1 -- -- ....
-- . I
PhJ"()_s_ta_u_r'!..s_si_c_u,!!~.__ _- . - - ._+-! __ - .1__ ...! --1
,Pleuraspis bipennis e _ _ _ I
:Pleuraspis c~~5tricta .-- .•----- ...-..- .....-- -----! I ---.-----1
rp-"iStaca-r;iha
I ._ _
dodecodon
_.. _ __ .. _
_-; - .----- i----
.--1-- ---.-- __ .. _._._ .. _
r--'- I" .-...
84 Acantharia
e
.~ -..:uu
"'C
c: •..
-e
--
Pristacantha mu/tidentata
•• •
Pristacantha octodon
Pristacantha polyodon
•• •
Pseudo/ithium bifidum
• •• • ••
Pseudo/ithium compressum
• •
Siphonaspis cristata
• •
Stauracantha orthostaura
• • • •
Stauracantha spinulose
•
Stauracon pallidus
•• •• • • •
Stauraspis stauracantha
•
1----~~-----------------+_------~-------_+------_4--------~------1_--------+_--------
Stauraspis xiphacantha
• • •
Tetra/onche arachnoides
• • •
Tetra/onche hastata
•
---------~~------+--------~------_+------~--------~-------4_------~
Tetra/onche quadrata
•
Tetra/onche rara •
Xiphacantha a/ata
Xiphacantha mu/tispina
••
• •
• • •
Xiphacantha quadridentata
Zygostaurus amphitectus
•
•
•
•
••
~~--------~-----------+--------~--------~------4_-------~------~--------~----------
•• ••
Zygostaurus cornutus
•
Zygostaurus sagitta/is
•
Table 1. Generalized distribution patterns of acantharian species (chiefly from Reshetnjak, 1981).
from the widespread geographic distribution of acan- Acanthin: term used by Haeckel to designate the
tharian species it is reasonable to assume that most of strontium sulfate of which acantharian skeletons
them could occur in the South Atlantic. Accordingly, are composed.
we include all the species so far described and cur- Accessory spines (Fig. 6.51, 6.61, 6.64, 6.72): spines
rently considered valid by Schewiakoff (1926) and/or other than the 20 radial spicules, usually smaller
Reshetnjak (1981) (see Table 1). than the latter and arising from the surface of the
latticed shell.
Apophyses: any protruding outgrowths of the skeletal
Taxonomy and identification meshwork. Apophyses on spicules may branch
repeatedly and remain separated from those of
Glossary neighboring spicules (Fig. 6.117b, c), or the
branches may coalesce forming elaborate latticed
The following glossary defines the terms most com- shells (Fig. 6.126a, b). Primary apophyses are
monly used for the description of Acantharia. those inserted directly into the spicule; secondary
ones rise from the primary, and so on.
Armor shell: the latticed shell of some Acantharia.
Acantharia 85
suture
(=sutural
furrow)
coronal
pore
spicule
aspinal
pore
-. sutural pores
Dictyaspis furcata
A B
aspinal
pores
suture
(=sutural
furrow)
-. coronal pores
Craniaspis parmipora
c D Icosaspis elegans
isolated polar plate
Fig. 3. Pore nomenclature. A: original; B-D: from Haeckel (1887).
Aspinal caveola: caveola in aspinal position (see aspi- Calymma (Fig. 2, 6.19): jelly-like, frothy extracapsu-
nal pores). lar cytoplasm (see central capsule).
Aspinal furrow: furrow in aspinal position (see aspi- Caveola: small depression or pit on shell surface.
nal pores). Central capsule (Fig. 2, 6.19): mucoid or chitinous sac
Aspinal pores (Fig. 3): large pores on the shell surface within the protoplasm, bounded by an organic,
located next to the site where the spicule pierces multi-perforated membrane (the central capsule
the shell. membrane) containing the intracapsular cyto-
Axoneme (Fig. 2): central, stiff shaft of the axopod plasm. Central capsules may be spherical, ellip-
consisting of patterned arrays of microtubules and soidal, elongate, polygonal, rhomboid, etc.
covered by a thin sheet of cytoplasm. Claviform: club-shaped.
Axopod (Fig. 2): long, radiating, unramified process Comet (Fig. 6.91, 6.92): sheath or sleeve around a
of the cell stiffened by a patterned array of micro- skeletal spicule encasing it partly or totally, often
tubules. with serrated edge (see mantle).
"-'
I
I
86 Acantharia
~f1
,'~,
;f
Astrolithidae Conaconidae
Stauracanthidae
Amphilithidae
.;~
Acanthocollidae
Stauracanthidae
Phyllostauridae
Acantharia 87
Coronal furrow: furrow in coronal position (see coro- microstrands. Myonemes are located peripherally
nal pores). in the cell, grouped as umbrella ribs around each
Coronal pores (Fig. 3A, C, D). Openings surrounding spicule. Myonemes can be flat, ribbon-shaped,
the aspinal pores. cylindrical, triangular, etc.
Cristae: Crests on shell surface. Parmal pores: Pores piercing the shield and bordered
Diametral spicule (Fig. 6.9, 6.11): term applied to only by united branches of apophyses.
acantharians in which all pairs of opposing radial Periplasmic cortex: see ectoplasmic cortex.
spicules are fused, or simply unseparated, at their Pores (Fig. 3): Perforations in the latticed skeleton.
basal ends; the 10 diametral spicules cross loosely Primary spicule: see spicule.
in the center of the shell. When opposing spicules Radial spicule: see diametral spicule.
are not fused in pairs but form a central star-shaped Reticulopod: pseudopodia that branch and anasto-
or spherical concretion, or join otherwise in the mose in a net-like (reticulate) pattern.
center, the term radial spicule (Fig. 4) is used. Ribs: elongate thickenings on the shell-wall, spicules,
Ectoplasm (Fig. 2): extracapsular cytoplasm. comets, etc. When strongly developed they are
Ectoplasmic (=periplasmic) cortex (Fig. 2): A well usually termed keels or wings.
developed, thick ectoplasmic layer at the outer Secondary apophyses: see apophyses.
edge of the ectoplasm, thus affording it a 2-layered Secondary spicule: see spicule.
appearance. Spicule (Fig. 4): each of the 10 diametral, or 20 radial,
Extrusomes (=kinetocysts): membrane-bound, sub- variously shaped solid rods that form the skeleton
pellicularly located extrusible bodies of various, of Acantharia. Larger spicules are called primary
probably non-homologous types. or main, as opposed to the secondary, which are
Fibrillar meshwork: a system of radial and annular smaller.
organic fibrillae which support and stretch the Spindle-shaped: rod-like, tapering toward both ends.
ectoplasmic cortex. Spines: see accessory spines.
Four-edged (Fig. 4, 6.37a): quadrangular in cross-sec- Sutural furrow (=suture) (Fig. 3C): a furrow at the
tion. suture or junction of plates in the shell wall.
Four-winged (Fig. 6.123b, c): 4-bladed in cross-section. Wing (Fig. 6.123c): Lath-like longitudinal plate
Keel: Sail-like longitudinal plate extending laterally extending laterally from spicule or bar (see keel,
from spicule or bar (see ribs, wing). ribs).
Kinetocysts: see extrusomes.
Lanceolate: Lance or sword-shaped.
Leaf-cross (Fig. 4): the proximal end of a radial spic- Morphology
ule, commonly expanded, with 4 narrow to broad
laminae or wings (thus cross-shaped in cross-sec- Characters of the skeleton
tion) in a structure fitted to corresponding struc-
tures of neighboring spicules at the center of the The acantharian skeleton, composed of strontium sul-
skeleton. Sometimes referring to the entire fused fate, consists of 10 diametral (or 20 radial), solid spi-
structure of all such formations; see leaf-star. cules which cross the cytoplasm. The distal ends of the
Leaf-star (Fig. 6.123c, 6.118): the expanded, star- spicules arise at precise points on the cell surface,
shaped structure consisting of the fused leaf-cross- which Miiller (1859) described in an analogy to the sur-
es; see leaf-cross. face of the earth. Two quartets of polar spicules emerge
Mantle: variously fonned covering or coat, common- at 60"N and 600S, 2 quartets of tropical spicules at 30"N
ly a comet-shaped structure. and 300S, and 1 quartet of equatorial spicules at 0°.
Myofriscs: see myonemes. Tropical spicules are turned 45 with respect to all oth-
Myonemes (=myofriscs) (Fig. 2, 6.19b): motor orga- ers, so that the polar and equatorial tips arise at longi-
nelles consisting of contractile bundles of nonactin tudes 0°, 90°, 180° and 2700, while the tropical tips
filaments (2-4 J.11TI), twisted by pairs in elementary arise at 45°,135°,225° and 315° (Fig. I). This distribu-
Fig. 4. Types ofacantharian spicules. Acanthocollidae: from Haeckel (1887); others from Reshetnjak (1981).
II --------------------------------------
I'
1,
88 Acantharia
I tion of spicules is a general rule in the group, and by the sutural pores (Fig. 3A). Alternatively, plate
although anomalous patterns have been recorded, they growth can stop short of closing the sphere (Fig.
;
1 invariably seem to represent malformations, immature 6.116c, 6. 116t). Smaller coronal pores are scattered
III' individuals, or methodological artifacts. The "acanthar-
ians" with 18 to 500 spicules described by Popofsky
between the aspinal and the sutural ones (Fig. 3A, D).
,j (1904) are now excluded from the group and consi- Acantharian shells may be either a delicate, densely
dered Radiolaria incertae sedis. The "Litholophus" perforated meshwork (e.g., Fig. 6. 125a), or robust and
Ii stage described by some authors is probably a method-

ological artifact (Reshetnjak, 1981).
thick-walled (Fig. 6.88b). The basic shell pattern may
be modified further in various groups by additional
11 structures. Thus, in Phractopeltidae there are 2 (rather
Variability in acantharian shapes arises chiefly from than 1) concentric spheres (Fig. 6.80b). In Lithopteri-
1
1 differences in the development of these spicules. In dae large wing-like meshwork apophyses develop on
over half of the species spicules are of equal length. In the distal ends ofthe 2 main spicules (Fig. 6.86, 6.87).
several families, however, 2 or 4 equatorial spicules In Diploconidae the ellipsoidal shell is provided with
are larger and/or different in shape from the others, two large opposite conical "sleeves" (comets) or man-
and either similar or different among themselves in tle (Fig. 6.81-6.85).
size and shape. Differences can also be found between
polar and tropical spicules, and can in turn lead to Although dimensions of the skeleton's elements are
marked changes in the overall body shape: spherical, an important diagnostic feature, size is subject to wide
ovoid, lenticular, etc. geographic variability. Nencini and Saglina (1968),
L for example, showed that the main spicules of
The shape of the spicules is highly variable (Fig. 4), Atlantic specimens of Amphibelone anomala (Fig.
I from simple cylindrical rods, to elaborate 4- or 6-blad- 6.16) are over twice as large (380 1JlIl) as those col-
ed structures with smooth or serrated (usually at the lected in the Mediterranean (170 urn). The overall
distal end) edges. Cylindrical spicules are probably the size, aspect, and complexity of the skeleton also
most primitive, and they also characterize the juvenile changes markedly with growth (Fig. 5).
stages of species with more elaborate adult spicules.
Cell body organization
In some species (e.g., Holacanthida, Acanthochiasma)
the spicules cross loosely in the center of the cell (Fig. The cell body consists of a central endoplasm and a
6.9a, 6.13b). In others, they fuse in the center into a peripheral ectoplasm. In most species the endoplasm,
sphere or a star-like mass which mayor may not be dis- which contains the nuclei, is colored brown, red, or
sociated in S04H2' In some forms (e.g., Acanthospira black by pigments. In 3 orders (Holacanthida,
amphicopa) the spicules are flattened in the center and Symphyacanthida, Chaunacanthida) the endoplasm is
twisted about each other (Fig. 6.4b, 6.6b, 6.7b). The dense in the cell center and becomes progressively
mode of junction of the spicules in the center of the clearer towards the periphery. Moreover, it is not dis-
shell is a most important diagnostic character. tinctly separated from the ectoplasm. In contrast, in the
fourth order (Arthracanthida), the endoplasm is easily
In over half of acantharian species the skeleton is rep- distinguished from the ectoplasm since it is limited by
n resented by the 10 spicules only, yet others develop a a thick capsular wall. The endoplasm includes nuclei
,
i'
complementary meshwork and/or plates forming a and most cell organelles such as large mitochondria,
" .
shell. These additional structures are derived from ribosomes, areas of rough endoplasmic reticulum, die-
anastomosing and merger of lateral apophyses of the tyosomes and various membrane-bound organelles. In
spicules. Thus, each spicule can generate a pentagonal some Symphyacanthida (Haliommatidium and
or hexagonal plate by lateral growth. This plate usual- Dicranophora), during the vegetative life the endo-
ly has 2 large pores right next to the spicule - the aspi- plasm contains a single large nucleus, probably poly-
nal pores (Fig. 3). The edges of the plate can come in ploid, while in all other Acantharia there are numerous
, i
. , contact with neighboring plates, in which case the small round or oblong nuclei. In many species, the tro-
suture (or sutural furrow) can remain visible or disap- phont contains hundreds to thousands of nuclei imme-
pear altogether. Sutural furrows are often interrupted diately prior to gametogenesis.
Acantharia 89
A B c
Fig. 5. Successive stages of growth of the skeleton of Pleuras pis costa/a. From Reshetnjak (1981), according to Schewiakoff (1926).
In the Holacanthida, Chaunacanthida and Symphya- Their lengths are in the range of about 5-90 urn. They
canthida, which encyst just before gametogenesis, the are either flat, triangular, or as long ribbons or short
endoplasm is progressively enriched in tiny oval cylinders. The myonemes are motor organelles
plates - the lithosomes. They are generated by the capable of active movement. Rapid and simultaneous
Golgi and are birefringent. They are transported contraction of all the myonemes produces inflation of
towards the cell surface and arranged to form the cyst the cortex and the volume ofthe ectoplasmic region is
wall (Fig. 6.128, 6.129, 6.130, 6.135). During this consequently increased. Such a coordinated activity
morphogenetic change the external networks and the of the myonemes may playa part in buoyancy regula-
myonemes are shed. tion (Febvre-Chevalier and Febvre, 1994).
Axopodia, slender processes radiating from the cell Outline classification

surface, are sensitive and can retract upon chemical or
physical stimulation, then re-grow progressively and Order Holacanthida Schewiakoff, 1926
slowly. They are stiffened by a central axis (axoneme) Family Acanthocollidae Febvre, Febvre and
consisting of patterned arrays of microtubules. Michaels, in press (genera Acanthocolla, Acantho-
cyrtha, Acanthospira)
The cytoplasm around each spicule is limited by the Family Acanthochiasmidae Haeckel, 1862,
cell membrane and contains 2-60 myonemes (depend- emend. Schewiakoff, 1926 (genus Acanthochiasma)
ing on the species). Myonemes are birefringent fibril- Family Acanthoplegmidae Schewiakoff, 1926,
lar bundles anchored by one end to the spicules and by emend. Reshetnjak, 1979 (genus Acanthoplegma)
the other end to the periplasmic cortex (Fig. 2, 6.19b). Order Symphyacanthida Schewiakoff, 1926
90 Acantharia
Family Amphilithidae Haeckel, 1882, emend. species for genera which comprise over 5 species. In
Schewiakoff, 1926 (genera Amphibelone, Amphi- addition, brief diagnoses for all the known species are
lithium) given. This section relies heavily of the works of
I Family Astrolithidae Haeckel, 1860, emend. Haeckel (1887), and especially Schewiakoff (1926)
i.
Schewiakoff, 1926 (genera Acantholithium, Astro- and Reshetnjak (1981).
lithium, Astrolonche, Heliolithium)
Family Pseudolithidae Schewiakoff, 1926 (genera Key to orders and families
Dicranophora, Haliommatidium, Pseudolithium) 1 Ten diametral spicules of equal or unequal length,
Order Chaunacanthida Schewiakoff, 1926 crossing or fitting together in cell center, dissoci-
Family Stauraconidae Schewiakoff, 1926 (gene- ated after treatment with sulfuric acid. Endoplasm
ra Amphiacon, Heteracon, Stauracon) with pigments and numerous nuclei. Ectoplasm
Family Conaconidae Schewiakoff, 1926 (genus clearer and less granulose than endoplasm, resem-
Conacon) bling a gelatinous sheet. Limit between endo-
Family Gigartaconidae Schewiakoff, 1926 plasm and ectoplasm generally poorly defmed.
(genus Gigartacon) Capsular wall near ectoplasmic cortex, poorly vis-
Order Arthracanthida Schewiakoff, 1926 ible in light microscopy. Cell surface irregular or
Suborder Sphaenacantha Schewiakoff, 1926 limited by thin periplasmic cortex. Cytoplasm
Family Acanthometridae Haeckel, 1862, emend. crossed by numerous thin, sensitive axonemes.
Schewiakoff, 1926 (genera Acanthometra, Amphi- Flat myonemes. Gametogenesis within a cyst:
lonche, Tetralonche) order Holacanthida (Fig. 6.1-6.15) .4
Family Dorataspidae Haecke1, 1862 (genera 1a 20 radial spicules 2
Aconthaspis, Coscinaspis, Craniaspis, Dictyaspis, 2 Bases of the 20 radial spicules indissociable in
Dorataspis, Hystrichaspis, Icosaspis, Lychnaspis, S04HZ' tightly fused into tiny sphere or star-like
Pleuraspis, Siphonaspis, Stauraspis) mass. Cell body round or oblong. Endoplasm 2-
Family Phractopeltldae Haeckel, 1862 (genus layered, inner one dense, containing nuclei,
Phractopelta) brown, yellow or reddish pigments, outer one
Family Diploconidae Haeckel, 1862 (genus with anastomosed cytoplasmic strands and axo-
Diploconus ) podial axonemes. Capsular wall visible in light
Family Lithopteridae Haeckel, 1887, emend. microscopy. Ectoplasm transparent. Periplasmic
Schewiakoff, 1926 (genus Lithoptera) cortex obvious in light microscope, consisting of
Family Hexalaspidae Haeckel, 1887 (genera elastic lattice with numerous pores through
Coleaspis, Hexaconus, Hexalaspis) which axopodia arise. Myonemes flat: order
Suborder Phyllacantha .Schewiakoff, 1926 Symphyacanthida (Fig. 6.16-6.33) 6
Family Phyllostauridae Schewiakoff, 1926 (gene- 2a Bases of the 20 radial spicules dissociable with
ra Acanthostaurus, Amphistaurus, Lonchostaurus, S04HZ 3
Phyllostaurus, Zygostaurus) 3 Bases of 20 radial spicules oblong, resembling
Family Stauracanthidae Haeckel, 1887 (genera grape seeds or bowling pins. Spicules cruciform
Phatnacantha, Pristacantha, Stauracantha, in cross section, generally serrated on two oppo-
Xiphacantha) site edges, smooth on other two. Spicules held at
Family Dictyacanthidae Schewiakoff, 1926 base in contractile matrix. Endoplasm black or
(genus Dictyacantha) reddish in cell center, clearer at periphery.
Capsular wall very conspicuous in some genera.
Identification Ectoplasm transparent. Cortex showing square
Because of the widespread occurrence of Acantharia pattern of elastic junctions. Myonemes ribbon-
and the scarcity of data for the South Atlantic, the sec- shaped, very long and mobile. order Chauna-
tion that follows includes all species so far described canthida (Fig. 6.34-6.41) 8
and. considered valid by Schewiakoff (1926) and/or 3a Bases of 20 radial spicules pyramidal with 4-6
Reshetnjak (1981). The identification-related infor- facets, with or without basal extensions juxta-
mation offered includes a general key for orders and posed or linked to form more or less cohesive
families, genus keys within each family, and keys to system. Endoplasm with numerous nuclei, pig-
Acantharia 91
ments, inclusions, and zooxanthellae. Thick cap- 9 Ectoplasm homogeneous, clear. Periplasmic cor-
sular wall. Ectoplasm limited by periplasmic tex without elastic junctions. Four-six myonemes
cortex. Myonemes cylindrical, generally numer- per spicule: family Conaconidae (Fig. 6.37)
ous. A few axopodia emerging between spicules: 9a Ectoplasm granular. Periplasmic cortex with
order Arthracanthida (Fig. 6.42-6.126). elastic junctions. Large size (spicules 0.5-2 mm
Order is divided into 2 suborders on basis of in length); 6-8 myonemes per spicule: family
shape and arrangement of spicules in their basal Gigartaconidae (Fig. 6.38-6.41)
region. Endoplasm always containing numerous 10 Proximal end of each spicule looking like a 5-6-
zooxanthellae. Contrary to three other orders of edged pyramid: suborder Sphaenacantha.
Acantharia, gametogenesis occurs in a gamont In this suborder the base of the spicule looks like
which keeps the aspect of the trophont. Whole a small pyramid without lateral extensions on
endoplasm converted into biflagellate gametes the ridges of the pyramid 11
which are shed across capsular wall at maturity lOa Proximal end of each spicule bearing 4-6 lateral
......................................................................... 10 extensions, resembling leaves or blades .
4 Long, equal, generally cylindrical spicules cross- Spicules assembled in cell center into star-like
ing in cell center. Gamontocyst round or oval, structure with bases delimiting 22 pyramidal
limited by small plates, showing remnants of spic- cavities. The pyramidal bases of the spicules
ules, forming small bundle in center of cyst 5 bear 4-6 lateral leaves or blades on each ridge:
4a Spicules equal or unequal, twisted at half length, suborder PhyUacantha 15
more or less intimately fitted or cemented with 11 Spicules of equal or unequal length, without
one another in cell center of adult vegetative apophyses. Thin capsular wall. Periplasmic cor-
forms: family Acanthocollidae (Fig. 6.1-6.8) tex with elastic junctions. Sixteen to forty myo-
5 Ectoplasmic rim irregular. Cortex and myonemes nemes per spicule: family Acanthometridae
very thin, poorly visible. Numerous granule- (Fig. 6.42-6.50)
studded axopodia: family Acanthochiasmidae lla Spicules bearing lateral hypophyses 12
(Fig. 6.9-6.14) 12 Spicules all of same length 13
5a Ectoplasm very narrow, limited by elastic peri- 12a Two to six spicules of different lengths with lat-
plasmic cortex. Capsular wall very thin, closely eral apophyses or armored plates 14
fitted in some areas to periplasmic cortex. Two 13 Spicules all of same shape, with single lattice of
flat myonemes per spicule, totally included under concave plates, joined with each other forming
periplasmic cortex. Ten-fifteen large spherical single intracytoplasmic shell. Endoplasm yellow,
granules scattered around each spicule just under red or green. Thin capsular wall. Periplasmic
periplasmic cortex: family Acanthoplegmidae cortex. Six to sixteen myonemes per spicule:
(Fig. 6.15) family Dorataspidae (Fig. 6.51-6.77)
6 Spicules of equal length and same shape 7 13a Spicules bearing 2-3 sets of plates and/or apo-
6a Two spicules longer and thicker than others: physes forming 2 or 3 concentric latticed shells.
family Amphilithidae (Fig. 6.16-6.21) Six to 8 myonemes per spicule: family
7 Spicules fitting with each other in spherical or Phractopeltidae (Fig. 6.78-6.80)
star-like structure. Capsular wall peripheral. 14 Two spicules longer and stronger than others,
Ectoplasmic cortex made of elastic threads: fam- bearing longitudinal cornets, and shell with two
ily Astrolithidae (Fig. 6.22-6.28) small oval perforations around each small spi-
7a Spicules very thin, long, all equal, smooth or cule. Acantharian resembling a butterfly bow:
bearing apophyses or latticed plates connected family Diploconidae (Fig. 6.81-6.85)
with each other to form perforate shell: family 14a Four spicules longer and thicker than others,
Pseudolithidae (Fig. 6.29-6.33) with flat triangular latticed plates on apical third
8 Twenty spicules of equal length and same shape, of length. Each plate consisting of 1-5 lateral
with conical proximal ends 9 apophyses cross-linked by 3-5 orthogonal bars
8a Spicules of unequal length. Ectoplasm with elas- delineating square lattice. Species flattened in
tic junctions, 8-12 myonemes per spicule: fami- plane of longest spicules. Endoplasm including
ly Stauraconidae (Fig. 6.34-6.36) numerous zooxanthellae. Six to eight short
T
92 Acantharia
I
cylindrical myonemes: family Litbopteridae 2a Spicules of approximately equal length and
(Fig. 6.86-6.87) same shape, or 1-2 larger and stouter, twisted j
14b Two to six spicules longer than others, all with and fitted in cell center. Endoplasm reddish. I
lateral thick armored plates or apophyses. Cell Five-six flat myonemes per spicule. Cyst bicon- !
body spherical or oblong. Numerous axopodia. ical, showing remnants of spicules after S04H2
Capsular wall overlaying armored shell, ecto- treatment: genus Acanthocyrtha (Fig. 6.4). One
plasm around shell covered with a periplasmic species: Acanthocyrtha haeckeli
cortex. Eight to 12 short myonemes: family 3 Two equatorial spicules much thicker than other
Hexalaspidae (Fig. 6.88-6.97) 8. Central part twisted, distal part cylindrical, flat-
15 Spicules generally cylindrical, without apophy- tened at both ends. Cell body flattened, square or
ses, all equal and of same shape or 2-4 spicules rectangular. Four myonemes. Two diametral spic-
longer and stronger than others. Endoplasm ules visible in cyst center: genus Acanthocolla
spherical, ellipsoidal or cushion-shaped, limited (Fig. 6.3). Two species: Acanthocolla cruciata,
by thick capsular wall. Sixteen to 80 cylindrical Acanthocolla solidissima
myonemes: family PbyUostauridae (Fig. 6.98-
6.115) • Acanthocolla cruciata (Haeckel, 1860) (Fig. 6.1).
15a Spicules bearing apophyses 16 Two main spicules longer and morphologically
16 Four apophyses looking like either simple or different from other 8. They may be equal, or
branched lateral wings or thorns, or bearing one (more rarely) differ in width, with blunt ends, or
or several secondary orthogonal apophyses with lanceolate, sharp distal ends. Sometimes with
which can form a latticed plate. Endoplasm longitudinal keel. Secondary spicules thinner and
spherical or polygonal, limited by thick capsular shorter than main. Length of main spicules: 0.16-
wall: family Stauracantbidae (Fig. 6.116-6.124) 0.18 mm, secondary spicules: 0.11 mm.
16a Latticed plates formed by slightly curved apo- • Acanthocolla solidissima (Popofsky, 1905) (Fig.
physes joining together into a latticed sphere. 6.2). All 10 spicules of nearly equal length, com-
Endoplasm spherical: family Dictyacanthidae pressed, double edged, sharp at both ends, fused at
(Fig. 6.125-6.126) center into central structure. Spicule length: 0.062
mm, width: 0.002 mm.
Order Holacanthida • Acanthocyrtha haeckeli Schewiakoff, 1926 (Fig.
6.5). Sixteen diametral spindle-shaped spicules; at
Family Acantbocollidae times, one or both main spicules longer, stouter, or
Family characterized by 10 cylindrical spicules, equal otherwise different from others; twisted and fused
or unequal, twisted at midpoint where they join and fit at center, forming central kernel. Cell body sphe-
more or less tightly, or cemented at center. Endoplasm rical or ovoid. Length of largest spicule: 0.3 mm,
includes brown pigments, some nuclei and organelles. diameter: 0.004-0.006 mm.
Ectoplasm clear, transparent,with anastomosed strands. • Acanthospira amphicopa (Haeckel, 1887) (Fig.
A thin, flexed capsular wall is overlain by a superficial 6.7). Sixteen skeletal spicules cylindrical in mid-
periplasmic cortex. A few thin axopodia emerge from dle, compressed at both ends. Both ends oar-
cell body. The myonemes (2-6 per spicule) are triangu- shaped, strongly compressed, 5-7 times as wide as
lar, very short. Gametogenesis within oblong cysts. in central portion. Spicule length: 0.12-0.16 mm.
• Acanthospira spiralis (Haeckel, 1887) (Fig. 6.8).
Key to genera of Acanthocollidae Spicules spindle-shaped, tapering from thicker
1 Spicules of equal length 2 central part towards two thin conical ends, rigid,
la Spicules of unequal length, similar shape, fused inelastic; their central part spirally convoluted in
at center 3 very peculiar cochlea-like manner. Ten spicules
2 Spicules simply twisted, crossing in cell center. propped one upon another by central screw.
Two flat myonemes. Cyst long and cylindrical: Central capsule dark, opaque. Spicule length: 0.2
genus Acanthospira (Fig. 6.6). Three species: to 0.3 mm, breadth of central spiral part: 0.01 mm.
Acanthospira amphicopa, Acanthospira spiralis, • Acanthospira torta (Popofsky, 1904).Spicules con-
Acanthospira torta stricted in middle, lance-shaped, distally flattened,
Acantharia 93
tips sharp. Central linkage as in Acanthospira spi- oblong cyst limited by very thin birefringent
ralis. Spicule length: 0.12-0.16 mm. plates. Remnants of spicules forming bundle in
cyst center. Myonemes extremely thin, appearing
Family Acantbocbiasmidae in EM as narrow flat bands.
Spicules long, straight, of equal length, simply • Acanthochiasma hertwigi Popofsky, 1904 (Fig.
crossed in cell center, generally cylindrical, smooth. 6.10). Two main diametral spicules 7-8 times
Endoplasm in 2 concentric zones. Ectoplasm vacuo- longer than other 8. Main spicules thinner in cen-
lar. Myonemes very thin. Gametogenesis in oblong or ter, gradually thickening distally, ending in 4-
spherical cysts. One genus: Acanthochiasma. Six spe- bladed arrow head-type structures. Length of main
cies: Acanthochiasma fusiforme, Acanthochiasma spicules: 0.34 mm, their thickness: 0.0048 mm;
hertwigi, Acanthochiasma planum, Acanthochiasma length of smaller spicules: 0.05 mm, thickness:
quadrangulum, Acanthochiasma rubescens, Acantho- 0.001 mm.
chiasma serrulatum. • Acanthochiasma planum Popofsky, 1904 (Fig.
6.11). Spicules smooth, thinner in center, distal
Key to species of Acanthochiasma parts thicker, with sharp tips. Length of spicules:
I All 10 diametral spicules of equal or almost 0.11 mm.
equal length 2 • Acanthochiasma quadrangulum Popofsky, 1904
la Two main (larger) diametral, and 8 smaller spic- (Fig. 6.12). Spicules quadrangular in cross section,
ules: Acanthochiasma hertwigi (Fig. 6.10) almost twice as thick at lance-like distal end as
2 Spicules smooth, cylindrical., 3 centrally. Spicule length: 0.09-0.10 mm, thickness
2a Spicules quadrate or ellipsoidal in cross section at distal end: 0.007-0.008 mm; diameter of cell
...........................................................................4 body: 0.15 mm .
3 Spicules long, flexible, almost cylindrical: • Acanthochiasma rubescens Haeckel, 1862 (Fig.
Acanthochiasma rubescens (Fig. 6.13) 6.13). Spicules cylindrical, thin, very flexible. Ecto-
3a Spicules spindle-shaped, 2-2.5 times shorter than and endoplasm barely distinguishable from one
in Acanthochiasma rubescens: Acanthochiasma another. Cell body in live specimens yellow-brown-
fusiforme (Fig. 6.9) ish with few zooxanthellae.: Pseudopodia all simi-
4 Spicules ellipsoidal in cross section 5 lar, branched and anastomosed reticulopods. Large
4a Spicules quadrate in cross section: Acantho- and common species. Spicule length: 0.8-1 mm.
chiasma quadrangulum (Fig. 6.12) • Acanthochiasma serrulatum Schewiakoff, 1926
5 Spicules smooth, thinner in center, expanding (Fig. 6.14). Spicules thin, slightly compressed,
toward distal ends, tips sharp, spicule length: margins faintly serrated. Spicule length: 0.55-0.60
0.11 mm: Acanthochiasma planum (Fig. 6.11) mm; cell body diameter: 0.05-0.06 mm.
5a Distal section of spicules with serrated edges;
spicule length: 0.55-0.60 mm: Acanthochiasma Family Acantboplegmidae
serrulatum (Fig. 6.14) Ten diametral spicules simply crossing in cell center,
long, equal, cylindrical, straight and very thin.
• Acanthochiasma fusiforme Haeckel, 1860 (Fig. Endoplasm with brown or yellowish inclusions and
6.9). Spicules spindle-shaped with tapering ends, numerous nuclei, clearer towards periphery. Capsular
0.3-0.6 mm long (almost 2.5 times shorter than in wall very thin, consisting of thin network. Ectoplasm
Acanthochiasma rubescens), stiff. Cell body limited by periplasmic cortex, consisting of hexagonal
bulky, with numerous sensitive axopodia radiating elastic lattice. Two large flat triangular myonemes
in all directions. Endoplasm consisting of central anchored on each spicule, completely covered by per-
black-reddish sphere surrounded by clear vesicu- iplasmic cortex. Large spherical granules scattered
lar sheet including most nuclei. Ectoplasm resem- around each spicule just under periplasmic cortex
bling transparent jelly striated by anastomosed (these spheres were confused with zooxanthellae by
cytoplasmic strands and axopodial axonemes. As Schewiakoff, 1926), so ectoplasm seems to bear small
in Acanthochiasma rubescens, periplasmic cortex umbrellas around the 20 spicules. As in other members
more visible when cell is completely inflated after of Holacanthida, gametogenesis takes place in an oval
long period of calm. Gametogenesis occurs in cyst after complete remodeling of cell. One genus
94 Acantharia
Acanthoplegma. Only one species: Acanthoplegma remaining spicules pin-like. Central capsule green,
. ! krohni . semitransparent. Length of anterior spicule: 0.1 to
• I
I
: 1
• Acanthoplegma krohni (Haeckel, 1860) (Fig. 0.2 mm, breadth: 0.02 to 0.03 mm; length ofposte-
6.15). Spicules thin, cylindrical, tips sharp, central rior spicule: 0.2 to 0.4 mm, breadth: 0.04 to 0.08
l: linkage loose. Cell body spherical. Each spicule

with 2 myonemes. Spicule length: 1.6 mm. •
mm; length of 18 smaller spicules: 0.05 to 0.15 mm.
Amphibelone hydrotomica (Haeckel, 1887) (Fig ..
6.19). Posterior main spicule usually slightly lon-
Order Symphyacanthida ger than anterior, both 4-bladed in cross section,
with sharp tips. All secondary spicules of similar
Family AmphiIithidae length, cylindrical, wide at base, tips sharp.
Skeleton of 20 radial spicules, of different size and Length of main spicules: 0.16-0.20 mm, width:
shape, two longer and thicker than other 18. All fused 0.012-0.014 mm; secondary, length: 0.1O-0.l4
in cell center. Cell body cylindrical or oval, endoplasm mm, width: 0.005 mm.
with numerous nuclei and zooxanthellae. Capsular wall • Amphibelone violin a (Haeckel, 1887) (Fig. 6.18).
thin, peripheral. Ectoplasm covered with periplasmic Main spicules quadrangular prismatic, almost vio-
cortex, attached to each spicule by 6-24 myonemes. lin-shaped, with four very broad, prominent, lamel-
lar wings, constricted in middle part, and broad-
Key to genera of Amphilithidae ened towards both ends; apex truncate pyramidal;
Two equatorial spicules cylindrical (referred to as base with large leaf-cross. Eighteen smaller spic-
anterior and posterior) much longer and thinner ules shorter (about three times shorter), at the base
than other 18. Anterior 2 times shorter than poste- quadrangular pyramidal, prismatic in distal part.
rior, other spicules 5-20 times shorter than the Central capsule spindle-shaped, opaque. Length of
posterior. Endoplasm with numerous zooxanthel- 18 secondary spicules: 0.04 to 0.08 mm.
lae: genus Amphilithium (Fig. 6.20, 6.21). Two • Amphilithium clavarlum (Haeckel, 1887) (Fig.
species: Amphilithium clavarium, Amphilithium 6.20). Main posterior spicule 0.5-1.5 times longer
concretum than anterior. Main spicules proximally 4-bladed
1a Two equatorial spicules 1.5-2 times longer than or 4-winged in cross section. Secondary spicules
other 18, both of about same length. Endoplasm 2.5 times shorter than main, thread-like, flexible.
cylindrical; with zooxanthellae and numerous Length of main spicules, anterior: 0.4 mm, poste-
nuclei. Six-eight short myonemes: genus rior: 0.7 mm; secondary spicules: 0.3 mm.
Amphibelone (Fig. 6.16-6.19). Four species: • Amphilithium concretum (Haeckel, 1887) (Fig.
Amphibelone anomala, Amphibelone cultellata, 6.21). Anterior main spicule usually 1.5 to 2.5
Amphibelone hydrotomica, Amphibelone violina times shorter than posterior, tips pointed.
Secondary spicules 5-8 and up to 20 times shorter
• Amphibelone anomala (Haeckel, 1860) (Fig. than main, cylindrical (very rarely main spicules
6.16). Two (more rarely 3 or 4) equatorial spicules as short as secondary ones). Cell body cylindrical.
. differing markedly from other 18, which are much Length of main spicules, anterior: 0.6-0.8 mm,
shorter. Main spicules of equal or almost equal posterior: 1.2-1.6 mm; secondary spicules: 0.08-
length, but shape usually differing: posterior 0.3mm.
thicker, with 3 wing-like extensions, anterior with
4. Main spicules with central cylindrical axis from Astrolithidae
which wing-like extensions protrude. Eighteen Skeleton of 20 radial spicules usually of equal size
secondary spicules thorn- or needle-like, thick- and same shape, linked at base into single homogene-
ened basally, tips sharp. Length of main spicules: ous either spherical or star-like mass. Endoplasm and
0.05-0.08 mm; secondary: 0.025-0.05 mm. ectoplasm separated by thin peripheral capsular wall.
• Amphibelone cultellata Haeckel, 1887 (Fig. 6.17). Periplasmic cortex as network of elastic threads
Main spicules of differing length and shape, cylin- between which narrow channels lead to internal
drical in distal part, four-flanged in proximal por- space. These splits often with electron dense material
tion with two broad flanges lying in equatorial which may correspond to defecation vesicles or
plane; posterior spicule two times as long and two degraded organelles. Six to 12 short flat, or 16-32
to four times as wide as anterior spicule; 18 shorter thread-like myonemes .
.3
~-----,---------------
Acantharia 95
Key to genera of Astrolithidae • Astrolithium piriforme (Popofsky, 1904) (Fig.

1 Spicules cylindrical, usually of equal length, 6.24). Strongly inflated basal section of spicules
sharp-pointed and intimately united at base, sharply separated from .distal, thread-like "part.
forming spherical central mass. Endoplasm Spicule length: 0.12-0.15 mm, width: .0.02 mm;
golden-yellow in center, granular at periphery: diameter of basal part: 0.04 mm.
genus Heliolithium (Fig. 6.28). One species: • Astrolonche pectinata (Müller, 1858). Spicules
Heliolithium aureum compressed, 4-edged, pyramidal at base, tips sim-
la Spicules rhombic or cylindrical only over part of ple or bifid. On proximal half, broader edges with
length 2 to 2 longitudinal rows of opposite apophyses with
2 Spicules rhombic: genus Acantholithium (Fig. 3-4 slender teeth in each row. Length of spicules:
6.22, 6.23). Two species: Acantholithium dico- 0.2 mm, width: 0.08 mm.
pum, Acantholithium stellatum • Astrolonche pinnata (Müller, 1858). Spicules in
2a Spicules cylindrical, bulbous at base: genus distal half compressed, linear, two-edged, with
Astrolithium (Fig. 6.24, 6.26). Two species: bifid apex; in proximal half three times as broad,
Astrolithium bulbiferum, Astrolithium piriforme lanceolate, pyramidal at base, without leaf-cross.
2b Spicules with 2-6 rows of sharp thorns at base. From two broader edges of proximal half, two lon-
Numerous axopodia: genus Astrolonche (Fig. gitudinal rows of opposite apophyses; four to six
6.25, 6.27). Three species: Astrolonche pectina- broad triangular teeth in each row. Length of spic-
ta, Astrolonche pinnata, Astrolonche serrata ules: 0.3 to 0.4 mm, breadth in outer half: 0.012
mm, in inner half: 0.03 to 0.04 mm.
• Acantholithium dicopum (Haeckel, 1860) (Fig. • Astrolonche serrata (Haeckel, 1860) (Fig. 6.27).
6.22). Spicules sharpened at both ends, quadran- Distal half of spicules thin,cylindrical,. or slightly
gular in cross section at center, gradually thicken- compressed, tapering toward simple or bifid apex;
ing towards periphery. All 10 spicules usually of proximal half3-6 times broader, 4-winged in cross
equal length, sometimes 1 longer. Ectoplasm section, with 6 longitudinal rows .of opposite
coarsely vacuolated. Nuc1ei small, numerous (up apophyses. Edges of broader wings giving rise to
to 120). Eight-twelve c1aviform myonemes next to 3-4 pairs of strong conical teeth., Edges of smaller
each spicule. Ectoplasmic cortex of calymma two- (sagittal) wings producing 3-4 pairs . of short,
layered. Cell body diameter: 0.1 mm; spicule broad, triangular teeth, which in turn bear slender
length: 0.16 mm. conical teeth on opposite sides. Spicule bases
• Acantholithium stellatum (Haeckel, 1887) (Fig. pyramidal, with small leaf-cross. Eight-twelve
6.23). Spicules 4-bladed, pyramidal, tapering myonemes per spicule. Central capsule large,
gradually ftom base toward distal end. Ectoplasm whitish, ectoplasmic cortex 2-layered. Length of
fmely granulated. Few (6-8) zooxanthellae. Each spicules: 0.2-0.4 mm, width of proximal section:
spicule surrounded by 8 pear- or egg-shaped myo- 0.016imm, distal: 0.002-0.004 mm.
nemes. Ectoplasmic cortex 2-layered. Cell body • Heliolithium aureum Schewiakoff.. 1926 (Fig.
diameter: 0.1 mm, spicule length: 0.14 mm. 6.28). Spicules strongly widened basally, tapering
• Astrolithium bulbiferum Haeckel, 1882 (Fig. distally, sharp. Some specimens with 2 equatorial
6.26). Spicules very thin, elastic, gradually . spicules much longer than other 18.In live speci-
expanding toward base, fused into homogeneous mens 'cupola-shaped extensions of calymma .can
mass at center. Soft parts spherical and amoeboid, be seen over spicules. Elastic fibrillae of calymma
colorless or brownish. Pseudopodia branching and form heptagons or hexagons around spicules, each
anastomosing; inc1uding numerous unbranched with 5-6 pores. Spicule length: 0.28 mm.
ones with axial threads. Central capsule mem-
brane lacking. Ectoplasmic cortex 2-layered; outer Fami1y Pseudolithidae
one thick, with strongly developed radial elastic Twenty spicules of equal length, same shape, very
filaments; 6-8 band-shaped myonemes. Spicule thin, with or without apophyses or plates. Endoplasm
length: 0.3-0.36 mm, width: 0.0007-0.001 mm; yellow-brownish in center, c1earer at periphery, with
diameter of basal part: 0.008 mm. single -Iarge nuc1eus and lithosomes. Ectoplasm very
c1ear, transparent, .with thin. cytoplasmic strands form-
96 Acantharia
ing wide-meshed pseudopodial network. Thin capsular plate pores of different sizes, always quadrangu-
walllirniting endoplasm near cell center. Periplasmic lar. Diameter of latticed sphere: 0.16-0.22 rnm;
cortex stretching between tips of spicules. Numerous spicule length: 0.6-0.88 rnm.
axopodia and 16 flat myonemes per spicule. • Haliommatidium tabulatum (Müller, 1856) (Fig.
6.30). Spicules sharp at both ends, proxirnally cylin-
Key to genera of Pseudolithidae drical, distally conical. At 1/3 length fmely perforate
1 Spicules smooth, cylindrical, or flattened, with rhomboid plates forro; plates fusing together tightly,
sharp edges linked into small sphere in cell cen- forrning thin-walled spherical or ellipsoidal shell.
ter. Cell body golden yellow or brown: genus Diameter of latticed shell: 0.20 by 0.22 mm; cell
Pseudolithium (Fig. 6.32, 6.33). Two species: body: 0.18 rnm; spicule length: 0.24-0.26 rnm.
Pseudolithium bifidum, Pseudolithium compres- • Pseudolithium bifidum (Haeckel, 1860) (Fig.
sum 6.33). Spicules cylindrical, pin-like, tips sharp,
la Spicules with 4 apophyses perpendicular to axis each with 16 myonemes. Ectoplasm much thicker
of spicules in proximal third of length 2 than endoplasm; ectoplasmic cortex two-layered,
2 Apophyses short and truncate: genus Dicrano- Diameter of cell body: 0.3 rnm; spicule length:
phora (Fig. 6.29). One rare species: Dicrano- 0.6-0.8 rnm; diameter of central spherule: 0.012-
phora buetschli 0.015 rnm.
2a Apophyses long, with orthogonal secondary and • Pseudolithium compressum (Haeckel, 1860) (Fig.
tertiary apophyses. Apophyses forrning slightly 6.32). Spicules distally cylindrical, proximally
.concave grids which join just before gameto gene- compressed and keeled, 4-edged (rhomboidal in
sis to forro cyst. Biflagellate gametes forro within cross section). Ectoplasm thinner than endoplasm.
gamontocyst, then shed through perforated shell: Diameter of cell body: 0.6-0.8 rnm; spicule length:
genus Haliommatidium (Fig. 6.30, 6.31). Three 0.6-0.8 rnm.
species: Haliommatidium cancellatum, Haliom-
matidium muelleri, Haliommatidium tabulatum Order Chaunacanthida
• Dicranophora buetschli. Schewiakoff, 1926 (Fig. Family Stauraconidae

6.29). All spicules of equal length, proximally 4- One, two, or four radial spicules longer than others.
edged, with 2 small opposed, forking apophyses, Cell center black or reddish. Peripheral endoplasm
distally cylindrical, sharp. Ectoplasrnic cortex of and ectoplasm c1ear. Ectoplasm with numerous elastic
calymma 2-layered. Shelllength: 1.6 rnm; spicule junctions. Myonemes very long, mobile ribbons.
length: 0.8 rnm; diameter of cell body: 0.2 rnm.
• Haliommatidium cancellatum Schewiakoff, Key to genera of Stauraconidae
,1926. Spicules stout, long, very strongly com- 1 One smooth, spatula-shaped spicule longer than
pressed but with only distal third plainly two- other 19. Bases of spicules resembling bowling
sided; opposing apophyses arising in proximal pins, inc1uded in very contractile central mass.
quarter .of spicule length and bearing secondary Cell center black. Endoplasm containing nuc1ei.
apophyses. These do not unite with those of adja- Ectoplasm c1ear, surrounded by-thin periplasmic
cent spicules to forro continuous lattice shell. cortex. Eight ribbon-like, very mobile myo-
Dimensions 'of latticed shell: 0.18-0.19 by 0.11 nemes. Cyst size 0.08-0.1 rnm; spicules 0.2,.0.4
mm; cell body, length: 0.17 rnm, width: 0.1 rnm; rnm: genus Heteracon (Fig. 6.35). One species: l
spicule length: 0.2-0.3 rnm. Heteracon biformis

• Haliommatidium muelleri Haeckel, 1860 (Fig. la Two spicules three times as long as others, all
6.31). Shell a spherical or egg-shaped lattice. serrate along 2 opposite sharp edges; other edges
Spicules very long, three to four times as long as smooth. Cell body brown or reddish. Cytoplasm
lattice shell; flattened in proximal third, quadran- granulose. Six-eight . myonemes: genus
gular, in distal portion two-edged; proxirnal por- Amphiacon (Fig. 6.34). One species: Amphiacon
tion of spicules produce rhombic lattice plates, denticulatus
which are fused through end branches with neigh- lb Four equatorial spicules longer than other 16.
boring lattice plates to forro lattice shell. Lattice Endoplasm granular, with nuc1ei, separated from
Acantharia 97
ectoplasm by obvious capsular wall. Ectoplasm Family Gigartaconidae

clear, limited by periplasmic corte x bearing rec- Spicules long, stiff, of equal size (1-2 mm long);
tangular lattice of e1astic junctions. Six to eight edges may be serrated. Bases of spicules resembling
ribbon-like, long and very dynamic myonemes grape seeds embedded in contractile matter.
per spicule: genus Stallracoll (Fig. 6.36). Qne Periplasmic cortex with e1astic junctions: Qne genus:
species: Stauraeon pa/lidus Gigartacoll (Amphiaeon and Heteracon are some-
times placed in this fami1y). Five species: Gigartaeon
• Amphiacon denticlIlatlls (Haeckel, 1860) (Fig. abcisus, Gigartaeon bifidus, Gigartaeon eordiformis,
6.34). Two main spicules 3 times longer and 2 Gigartaeon dentieulatus, Gigartaeon jragilis,
times thicker than other 18. Spicules 4-edged in Gigartaeon mue/leri.
cross section, 2 opposite edges fmely serrate, others • Gigartacoll abcislls (Popofsky, 1904) (Fig. 6.39).
smooth. Six to eight myonemes. Cytoplasm brown Radial spicules similar, 4-winged, smooth-edged,
or reddish, granulose. Cell body size: 0.24 by 0.16 very thin, widening rapidly from base, outside soft
mm; length ofmain spicules: 0.56 mm, secondary: parts equally wide to pointed distal ends. Basal
0.2mm. spicule ends heart- or grape seed-shaped, fitted
• Heteracon biformis (Popofsky, 1904) (Fig. 6.35). loosely in center. Cell body star-shaped, colorless;
Skeleton with 1 main and 19 secondary spicules. ectop1asm vacuo1ar. Two kinds of pseudopodia.
Main spicule 0.2-0.5 mm longer than others, Central capsule membrane lacking. Periplasmic
lance-shaped, 4-winged, edges smooth. Secondary cortex wide, structureless. Four to six very long
spicules equal in length, 4-winged, 2 opposed thread-like myonemes. Shell diameter: 0.9-1.0
edges smooth, other 2 serrated. Length of main mm; cell body: 0.48-0.5 mm.
spicule: 0.25-0.44 mm, secondary: 0.2-0.26 mm; Gigartacon bifidlls (Popofsky, 1904). All spicules
cell body: 0.08-0.13 mm. cy1indrical, pin-like, distally pointed. Basal spi-
• Stallracoll pallidlls (Claparede, 1855) (Fig. 6.36). cule ends grape seed-shaped, fitted 100sely in cen-
Spicules 4-winged, 4 main with fine serration, 3-4 ter. Six-eight Myonemes. Spicu1e 1ength: 0.2 mm,
times longer than 16 secondary, which are smooth; width 0.003 mm.
each with 12-16 long, thread-like myonemes. Gigartacoll cordiformis (Popofsky, 1904).
Diameter of cell body: 0.10-0.16 mm; length of Spicules conical, tapering gradually to point dis-
main spicules: 0.7 mm, width at base: 0.009 mm, tally; inner half rounded 4-edged, outer half circu-
widest part: 0.012 mm, at top: 0.004 mm; length of lar in cross section. Basal spicule ends heart- or
secondary spicu1es: 0.15-0.20 mm, width: 0.004 grape seed-shaped. Spicu1e length: 0.07-0.08 mm,
mm. width 0.003 mm.
Gigartacon denticlllatlls (Haeckel, 1887). AlI
Family Conaconidae spicules very long, 5-10 times longer than diame-
All spicules of equal length (0.1-0.2 mm long) with ter of cell body; sometimes 1-2 (less often 4) equa-
two opposite edges sharp and serrate; others smooth. torial spicu1es 2 times as long as others. Spicules
Bases of spicules conical, quadrangular in cross sec- 4-winged, at base 2 opposing edges serrated (ser-
tion, loosely linked in cell center, embedded in central rations small, distally directed), other 2 smooth.
mass of contractile materia!. Cell body spherical. Each spicule with 6-8 long, thread-like myo-
Endoplasm clear, some zooxanthellae. Tiny polygo- nemes. Shell diameter: 0.6-1 mm; cell body: 0.06
nal, birefringent plates foun just before encystment. mm; spicu1e length: 0.03 mm (main equatorial
Very thin capsular wall. Ectoplasm transparent. Qne spicules may reach 0.6 mm).
genus: COllacoll. Qne species: Conaeonjoliaeeus. Gigartacon fragilis (Haecke1, 1860) (Fig. 6.40).
Conacon joliacells (Haecke1, 1887) (Fig. 6.37). Spicules of equal 1ength, 4-winged, edges deeply
Spicules lance-shaped, tapering toward both ends, serrated (serrations small, distally directed, deep-
tips pointed, 4-edged, 2 opposing edges smoóth, est distally), 4-5 times longer than cell body,
other 2 serrated. Spicule length: 0.18-0.22 mm, tapering proximally and distally (broadest at mid-
maximum width: 0.006-0.008 mm; diameter of dIe), tips sharp (usually broken off in adults).
cell body: 0.08 mm. Basal ends with 4 pear-shaped thickenings.
Spicule length: to 1 mm.
98 Acantharia
• Gigartaeon muelleri (Haeckel, 1860) (Fig. 6.41). 3a Radial spicules claviform proximally, distal ends
Spicules 4-edged, pyramidal, 2 opposed edges ser- rounded. Central capsule milky white, endoplasm
rated proximally (serrations fme, densely arrayed, reddish: Acanthometra cylindrica (Fig. 6.43)
proximally directed), each with 6-8 thread-like 4 Spicules short, tips sharp: Acanthometra fusea
myonemes. Live specimens orange to gray-yellow 4a Spicules long, 2-5 times thicker at base than cen-
in color. Shell diameter: 0.5 mm; cell body: 0.09- trally 5
0.12 mm; spicule length: 0.2-0.24 mm, width: 5 Spicules cylindrical, slightly thinner centrally,
0.008 mm. bases about 2 times thicker; central capsule
opaque, unpigmented: Acanthometra bulbosa
Order Arthracanthida (Fig.6.42)
5a Spicules needle-like, bases conical, almost 5
Suborder Sphaenacantha times thicker; endoplasm with brownish pigment
Family Acanthometridae granules: Acanthometra dolichoscia (Fig. 6.44)
Skeleton consisting of 20 spicules of same .length and 6 Spicules 4-bladed, prismatic; central capsule very
same shape, or with 2-4 spicules longer than others. small, transparent: Acanthometra prismatica
Cell body polygonal or oblongo Endoplasm with zoo- 6a Spicules 4-bladed, prismatic; central capsule
xanthellae, limited by capsular wall. Ectoplasm clear, small, polygonal, with yellow, red or brownish
limited by periplasmic cortex. Cylindrical myonemes. pigment grains and purple-red oil globules:
Acanthometra tetraeopa (Fig. 6.46)
Key to genera of Acanthometridae
1 Thin spicules equal or subequal, all of same shape. The species Aeanthometra annulata was erected by
Cell body polygonal: genus Acanthometra (Fig. Popofsky in 1906 (as Zygaeantha annulata Popofsky,
6.42-6.46). Eight species: Aeanthometra annulata, 1906) from 2 poorly preserved specimens, based on
Aeanthometra bulbosa, Aeanthometra cylindriea, the presence of "rings" on the spicules. This species
Aeanthometra doliehoscia, Aeanthometra fusea, has not been .recorded since, and such unusual struc-
Aeanthometra pellucida, Aeanthometra prisma ti- tures on the spicules, absent in all other acantharians,
ea, Aeanthometra tetraeopa suggest that the spines in question do not belong to
la Two or four equatorial spicules longer and thick- Acantharia (sponge spicules?).
er than others 2 • Acanthometra bulbosa Haeckel, 1860 (Fig. 6.42).
2 Two equatorial spicules longer than others. Cell All spicules of equallength, very thin, cylindrical
body lengthened, elongate. Endoplasm with zoo- with sharp tips, about 2 times wider at base, fused
xanthellae, .limited by capsular wall. Ectoplasm in center forming large spherical body. Spicule
clear and limited by periplasmic cortex. Numerous length: 0.30-0.35 mm.
axopodia. 'Sixteen to 24 myonemes per spicule: • Acanthometra eylindriea (Haeckel, 1887) (Fig.
genus Amphilonche (Fig. 6.47, 6.48). Two spe- 6.43). All spicules of equal length, equal width
cies: Amphilonehe diodon, Amphilonehe elongata throughout, claviform proximally, bases widened
2a Four equatorial spicules longer than others: pyrarnidal, distal ends rounded. Central capsule
genus Tetralonehe (Fig. 6.49, 6.50). Four spe- milky white, endoplasm reddish. Spicule length:
cies: Tetralonche araehnoides, Tetralonche has- 0.4-0.8 mm.
tata, Tetralonehe quadrata, Tetralonehe rara • Acanthometra dolichoscia Haeckel, 1860 (Fig.
6.44). Spicules cylindrical, tapering distally. Basal l
Key to species of Acanthometra ends 2-3 times thicker than distal portion, ending
1 Spicules circular in cross section 2 in 5-6 edged pyrarnids fused into central body.
la Spicules quadrangular in cross section 6 Cell body polygonal, protoplasm whitish-gray
2 Spicules of equal diameter throughout 3 with abundant oil inclusions. Each spicule with 32
2a . Spicules tapering distally ..4 or more myonemes. Diameter of cell body: 0.2
3 .. Spicules thin, needle-like, flexible, with sharp mm; spicule length: 0.50-0.56 mm.
tips. Central capsule transparent, colorless, with • Acanthometra fusca Müller, 1856. Spicules cylin-
large vacuoles and red oil globules, 24-40 myo- drical, tapering distally, with sharp tips. Basal
nemes: Acanthometra pellucida (Fig. 6.45) ends pyramidal, fused into central body. Cell body
Acantharia 99
polygonal, with yellow, reddish to brownish pig- • Amphilonche elongata (Müller, 1856) (Fig. 6.48).
ment grains. Central capsule thin-walled. Nuc1ei Two main spicules of equal or almost equal
(ea. 400) usually distributed around periphery of length, 4-bladed or 4-winged in cross section near
endoplasm. Calymma single-layered. Each spicule base, distally cylindrical or spindle-shaped, tips
with 32 or more myonemes. Diameter of cell sharp, 3-7 times longer than secondary. Secondary
body: 0.23 mm; spicule length: 0.25 mm. spicules needle- or thorn-like. Length on main
• Acanthometra pellucida Mililer, 1856 (Fig. 6.45). spicules: 0.40-0.75 mm; secondary: 0.05-0.26
All 10 diametral spicules of equal size and same mm. Very variable species, Schewiakoff (1926)
shape, thin, cylindrical, flexible, with sharp tips. described 3 subspecies: Amphilonche elongata
Specimens with 1, 2, or 4 larger equatorial spicules belonoides, Amphilonche elongata amphipolaris
very rare. Spicules linked in cell center by edges and Amphilonche elongata heteracantha.
of 5- or 6-edged pyrarnidal bases. Cell body • Tetralonche arachnoides (Popofsky, 1904) (Fig.
polygonal, with yellow-green endoplasm, with 6.49). Four main spicules cylindrical, much longer
numerous zooxanthellae (usually in groups of2-3) and up to 3 times thicker than remaining 16, each
and nuc1ei. Endoplasm limited by thin, colorless, with 7 myonemes. Central capsule spherical, com-
transparent capsular wall. Ectoplasm c1ear, with pressed, very small and dark. Calymma poorly
delicate cytoplasmic strands. Periplasrnic cortex developed. Length ofinain spicules:'0.7-0.15 mm,
of20 polygonal pieces, fixed to each other by elas- width: 0.004 mm; length of secondary spicules:
tic junctions. Each spicule with 24-40 thread-like 0.33-0.48 mm, width: 0.002-0.003 mm; diameter
cylindrical myonenies (juveniles with 8-16). of the central capsule: 0.06-0.09 mm. '
Diameter of cell body: 0.24 mm; spicule length: • Tetralonche hastata (Haeckel, 1860). Four main,
0.43-0.56 mm. One of commonest and most abun- equatorial spicules cylindrical in the basal half,
dant acantharians world-wide. spear-shaped or lanceolate in distal half, with thin
• Acanthometra prismatica (Haeckel, 1887). edges, little longer, but five to eight times as broad
Spicules prismatic, smoothly 4-edged in cross sec- as sixteen smaller spicules, which are cylindrical, of
tion, very long and thin, linked in cell center by equal' breadth throughout whole length.rwith bifid
edges of small 5- or 6-edged pyrarnidal bases. Cell apex. Central capsule yellow, spherical, or lenticu-
body spherical, colorless, with abundant zooxan- lar. Length of the 4 major spicules: 0.066 mm,
thellae. Ectoplasmic cortex single-layered, breadth in rniddle: 0.005 mm, at the apex: 0.008
Spicules with 16 myonemes. Diameter of cell mm; length of 16 smaller spicules: 0.054 mm,
body: 0.07 mm; spicule length: 0.3 mm. breadth: 0.001 mm.
• Acanthometra tetracopa Müller, 1856 (Fig. 6.46). • Tetralonche quadrata (Haeckel,' 1887)~ Four
Spicules 4-edged in cross section, tapering distal- main, equatorial spicules slightly compres sed,
ly, linked in cell center by edges of small 5- or 6- almost cylindrical in basal half; broader, lanceo-
edged pyramidal bases. Cell body spherical or late in distal half, with simple apex. Eight tropical
polygonal, with numerous green zooxanthellae, spicules a little shorter, but on1y half as broad,
yellow, red, or brownish pigment grains, and pur- sword-shaped, two-edged, of equal breadth, with
ple-red oil globules. Each spicule with 16-32 myo- simple apex. Eight polar spicules very short, coni-
nemes. Ectoplasm single-layered. Diameter of cell cal, also with simple apex. Length of 4 equatorial
body: 0.17 mm; spicule length: 0.27-0.3 mm. spicules: 0.18 mm, of 8 tropical spicules: 0.14
• Amphilonche diodon Haeckel, 1887 (Fig. 6.47). mm, of 8 polar spicules: 0.04 mm.
Two main spicules compressed, lanceolate, two- • Tetralonche rara (popofsky, 1904) (Fig. 6.50).
edged, tapering from broader rniddle towards two Four main 4-winged spicules longer than 16 secon-
ends; apex simple, base a small pyramid, without dary conical. One pair of main slightly longer than
leaf-cross. Eighteen smaller spicules sword- other and faintly keeled. All spicule tips pointed,
shaped, only two thirds or one half as long, and bases with small pyrarnids fused into central sphe-
one third or one half as broad, each with two oppo- rule. Length oflarger main spicules: 0.07-0.1 mm,
site teeth in distal parto Central capsule ellipsoidal. width: 0.004-0.006 mm; length of smaller main
Length of 2 major spicules 0.13 mm; breadth 0.2 spicules: 0.05-0.06 mm, width: 0.003-0.005 mm;
mm; length ofthe 18 secondary spicules 0.1 mm. length of secondary spicules: 0.03-0.05 mm, width:
0.002-0.004 mm.
r'
I
100 Acantharia
Family Dorataspidae small sutural pores. Central capsule generally pink.

Twenty spicules of same shape and generally same Twelve-sixteen myonemes per spicule: genus
length, with 2-4 cross-shaped ramified processes or 5- Icosaspis (Fig. 6.62-6.64). Three species: Icosas-
6 angular porous plates linking to form latticed or pis elegans, Icosaspis ornata, Icosaspis serrulata
annored shell. Conical bases of spicules linked or 5 Shield plates generally linked along polygonal
closely associated, forming mass in cell center. Cell sutural line. External surface of plates smooth
body spherical, with yellow or red endoplasm, sur- ........................................................................... 6
rounded by capsular wall. Ectoplasm with numerous 5a Shields without sutural line. External surface of
zooxanthellae. Periplasmic cortex covering ectoplasm shell bearing caveolae and network of combs
and linked to spicules by 6-16 myonemes. and perpendicular spines 8
6 In each shield 2 aspinal pores surrounded by one
Key to genera of Dorataspidae cycle of sutural pores localized along suturalline.
1 Spicules with 2-4 cross-shaped apophyses linked Endoplasm colorless or red. 6-12 myonemes per
with neighbors forming latticed shell 2 spicule: genus Dorataspis (Fig. 6.56-6.59). Seven
la Spicules with 5-6 polygonal, porous, or armored species: Dorataspis choanopora, Dorataspis gla-
plates forming spherical oroblong shell 5 diata, Dorataspis loricata, Dorataspis macracan-
2 Cylindrical spicules bearing two apophyses on tha, Dorataspis macropora, Dorataspis micropo-
proximal third, either unramified or branched as ra, Dorataspis polypora
fork and linked with apophyses of neighboring 6a In each shield 2 aspinal pores surrounded by one
spicules to delineate large- meshed shell. At or several cycles of coronal pores 7
junction points, 1-2. harpoon-shaped spines 7 Coronal pores forming cycles between aspinal
pointing towards exterior. Endoplasm yellowish. and sutural pores: genus Coscinaspis (Fig. 6.52).
Eight cylindrical myonemes per spicule: genus Two species: Coscinaspis isopora, Coscinaspis
Pleuraspis (Fig. 6.73, 6.74, 6.76). Four species: peripora
Pleuraspis bipennis, Pleuraspis constricta, 7a No sutural pores, junctions between adjacent
Pleuraspis costata, Pleuraspis sarmentosa shields conspicuous. Numerous coronal pores
2a Spicules with four opposite processes, which surrounding aspinal pores: genus Craniaspis
may be ramified, forming latticed plate 3 (Fig. 6.53). One species: Craniaspis parmipora
3 Secondary or tertiary apophyses linked with 8 AlI shields identical.. 9
secondary or tertiary apophyses of neighboring 8a Six shields on 2 equatorial spicules and 4 polar
spicules forming small-meshed latticed shell. spicules bearing large aspinal fÚrrows, each with
Cylindrical spicules: genus Stauraspis (Fig. 6 aspinal pores. Other 14 shields with caveolae,
6.77). Three species: Stauraspis echinoides, each with 2 pores. Each aspinal furrow sur-
Stauraspis stauracantha, Stauraspis xiphacantha rounded by 12-16 smaller sutural furrows bear-
3a Apophyses linked partly with each other, partly ing pores: genus Siphonaspis (Fig. 6.75). One
. with apophyses of neighboring spicules forming species: Siphonaspis cristata
latticed shell 4 9 Each aspinal caveola surrounded by aspinal
4 Each spicule bearing perforated plates with four pores and by 10-12 sutural furrows lO
cross-shaped pores (aspinal pores). Plates linking 9a In addition to sutural furrows, also coronal fur-
with neighboring plates by processes delineating rows, all with pores; combs with accessory
8-12 large pores (sutural pores). Endoplasm yel- spines: genus Aconthaspis (Fig. 6.51). Three I
lowish, with 8-16 myonemes per spicule: genus species: Aconthaspis capillata, Aconthaspis fur-
Lychnaspis (Fig. 6.65-6.72). Nine species: cata, Aconthaspis hastata
Lychnaspis giltschi, Lychnaspis longissima, 10 Sutural furrows with alternating parts with blind
Lychnaspis maxima, Lychnaspis mintma, pores and parts without pores. Endoplasm dark-
Lychnaspis polyancistra, Lychnaspis serrata, red: genus Hystrichaspis (Fig. 6.60, 6.61). Three
Lychnaspis strelkovi, Lychnaspis. undulata, species: Hystrichaspis dorsata, Hystrichaspis
Lychnaspis wagenschieberi fruticata, Hystrichaspis serrata
4a Each plate bearing 8 aspinal surrounded by 1-2 lOa All sutural furrows lacking pores. Endoplasm
cycles of smaller "coronal pores" and 1 cycle of colorless spherical or ellipsoidal. Eight myo-
Acantharia 101
neme s per spicu1e: genus Dictyaspis (Fig. 6.54, elliptical aspinal pores, 4-12 small (2-4 times
6.55). Two species: Dictyaspis jurcata, Dicty- smaller than aspinal) circular coronal pores, sur-
aspis solidissima rounded by 5-6 bi-Iobed or irregular sutural pores
ofvariable size. Shell diameter: 0.3 rnm; length of
• Acollthaspis capillata Haeckel, 1887. Shell very main spicules: 0.12-0.15 rnm.
dark and thick walled, opaque, with very numer- • Cralliaspis parmipora Haeckel, 1887 (Fig. 6.53).
ous (300-400 or more?) deep funnel-shaped dim- Shell thin-walled, with smooth surface, without
pIes, each perforated by small circular pore (40 crests and dimples. No sutural pores, all twenty
aspinal, 200-300 coronal, and 50-100? sutural p1ates connected by perfect sinuous sutures. Each
pores). High crests between dimples with very p1ate with two elliptical aspinal pores, 2 to 5 times
numerous simple accessory spines, nearly half as broader than numerous, roundish irregularly scat-
long as radius, so shell appears covered with hairs. tered coronal pores (30 to 50 on each p1ate). Radial
Radial main spicules very long and thin, cylindri- spicules internal, not prolonged on outside of she11.
cal, longer than diameter of shell. Diameter of Diameter of shell: 0.16 rnm, aspinal pores: 0.01
shell: 0.2 rnm, ofpores: 0.012 rnm. rnm, coronal pores: 0.002 to 0.005 rnm.
• Acollthaspis furcata Haeckel, 1887. Shell thick- • Dictyaspis furcata (Haecke1, 1887) (Fig. 6.54).
walled, dimpled, with network of crests, of same Main spicules cy1indrical, strongly compressed,
construction as A. has tata, with 170-190 dimp1es sharp at both ends. Secondary spicules rarnified,
(20 aspina1, 100-110 coronal, and 50-60 sutural). radiating from prominent pore frames. Aspinal
All dimples and pores nearly of same size. Each pores 1arge, subcircular, surrounded by 9-12 pen-
node of crested network bears stout spine, forked tagonal depressions with one smaller, circular
at base, with two divergent straight branches. sutural pore each. Spicule 1ength: 0.15 rnm'
Radial main spicu1es two-edged, sword-like, near- diameter of lattice sphere: 0.16 rnm; aspina1 pores:
ly as long as diameter of she11.Diameter of shell: 0.006-0.01 rnm; sutural pores: 0.003-0.005 rnm.
0.13 rnm, ofpores: 0.008 rnm. • Dictyaspis solidissima (Haeckel, 1862) (Fig.
• Acollthaspis hastata Haecke1, 1887 (Fig. 6.51). 6.55). Spicules very strongly compressed, lamel-
Shell thick-walled, with numerous (170-190) dim- lar, both ends sharp, each with 6-8 short myo-
pIes separated by network of elevated prominent nemes. All pores in funnel-shaped depressions;
crests. In center of each plate a larger dimp1e with oval aspinal pores surrounded by 10-12 smaller
2 aspinal pores, surrounded by corona of lOor 12 sutural pores. Cell body within 1attice shell spheri-
smaller dimples, each with single pore (coronal cal, colorless, with many zooxanthellae. Ecto-
and sutural alternately). All pores circular or plasmic cortex single-layered. Spicule length:
roundish, nearly of same size. If shell quite regu- 0.084-0.11 rnm; diameter of lattice shell: 0.08-0.1
1arly developed, with 50-60 sutural pores and 100- rnm; of aspinal pores: 0.01 by 0.006 rnm, of sutu-
110 coronal pores (6 in each equatorial p1ate, 5 in ral pores: 0.004 by 0.003 rnm.
each of other plates). Knobs of joining crests con- • Dorataspis choallopora Tchang and Tan, 1964.
ical, each with simple short accessory spine. Spicules all of same length, compressed. Four
Radial main spicules compressed, at dista1 end equatorial and four of 8 polar spicules wider than
spear-shaped, with rhomboida1, more or less hook- 12 remaining spicules. Apophysal plates fused
like apophyses, about as long as radius. Diameter into ellipsoida1 shelllacking sutures. Plates of four
of shell: 0.18 rnm; of pores and bars: 0.01 rnm. equatorial and of four polar spicules are hexago-
• Coscillaspis isopora Haeckel, 1887. Skeletal spi- nal, other 12 pentagonal. Gn both sides of each
cu1es cylindrical inside, conica1 outside thick- spicule two aspinal pores and 5-6 quite larger
walled 1attice she11. Two reniform aspinal pores, sutural pores varying slightly in size. All aspinal
5-6 round coronal pores, and 5-6 ovoid sutural and sutural pores with thin membranous skeletal
pores. All pores of approximate1y same size. Shell collar. Surface of shell with many compressed
diameter: 0.2 rnm; pore and bar width: 0.012 rnm. accessory spines which may be shortly forked.
Coscillaspis peripora Haeckel, 1887 (Fig. 6.52). Body length tip to tip 0.22 rnm, diameter of lattice
Shell thin-walled. Spicu1es cylindrical, slightly shell: 0.11 rnm, spicule length: 0.05 rnm, spicule
compressed or lance-shaped. Two small round or width: 0.005-0.01 rnm.
11
I 102 Acantharia
• Dorataspis gladiata Haeckel, 1887 (Fig. 6.57). • Dorataspis polypora Haeckel, 1887. Shell with 54
Shell ellipsoidal or spherical. Spicules strongly sutures and 160-200 sutural meshes, with 8 hexago-
/ compressed, two-edged, narrowing gradually from nal and 12 pentagonal plates. Aspinal pores kid-
middle toward each end; basal ends with conical ney-shaped, surrounded by 15-20 very small,
structures which fuse into central spherule. round sutural pores (three to four at each suture).
Aspinal pores triangular or elongated, surrounded Radial spicules thick, in inner longer part cylindri-
by 10-12 sutural pores, latter 3-4 times smaller cal, in outer shorter part conical. Differs from all
than former. Shell surface with inconspicuous other species in multiplication of sutural pores.
sutures, rough, with furrows, pustules or tiny Diameter ofshell: 0.2 mrn, ofaspinal pores: 0.016
thorns. Diarneter of shell (without spicules): 0.08- mrn, of sutural pores: 0.004 mrn.
0.1 mrn; length ofmain spicules: 0.08-0.1 mrn. • Hystrichaspis dorsata Haeckel, 1887 (Fig. 6.60).
• Dorataspis loricata Haeckel, 1860 (Fig. 6.58). Shell spherical. Each spicule arising from a pentago-
Shell spherical. Spicules with elliptical cross sec- nal or hexagonal pit with large ovoid to circular
tion, flattened; pyrarnidal bases joining in middle aspinal pores, surrounded by smaller pits. Among
into star-shaped body. Size of aspinal pores latter are 5-6 larger, pentagonal, poreless ones, at
strongly variable, ellipsoidal or kidney-shaped, vertices of plates, and 5-6 smaller, quadrate, with
surrounded by 5-6 sutural pores 2 times smaller to one sutural pore each. All pits .conspicuously
as large as aspinal pores. In adult specimens all framed. Size of sutural pores variable from almost
pores funnel-shaped, outer edges of funnels den- as large as aspinal to half their size. In adult speci-
tate. Sites of junction of 3-4 plates give rise to mens sides and angles of frame crests give rise to
short spines. Diarneter of shell (without spicules): thin spines of variable length. Shell diarneter
0.14-0.16 mrn; length ofmain spicules: 0.21 mrn; (excluding spicules): 0.36-0.58 mrn; spicule length:
secondary: 0.01-0.016 mm. 0.18-0.58 mrn; length of spines: 0.08-0.04 mrn.
• Dorataspis macracantha Haeckel, 1887. Shell with • Hystrichaspisfruticata Haeckel, 1887 (Fig. 6.61).
52 sutures and 52 sutural meshes; with 4 hexagonal Spicules cylindrical, compressed. Accessory
and 16 pentagonal plates. Aspinal pores round and spines very numerous, branched. At spicule bases
small, surrounded by 5-6, twofold smaller sutural in larger pits oval aspinal pores surrounded by
pores (one at each suture). Radial spicules quad- smaller pits without pores or with one sutural pore
rangular, prismatic, section external to lattice shell as large as to half size of aspinal pores. Shell
2-3 times longer than section within shell. diameter (excluding spicules): 0.12-0.18 mrn;
Diameter of shell.. 0.12 mrn, of parmal pores: aspinal pores: 0.007-0.013 mrn; sutural pores:
0.008 mrn, of sutural pores: 0.004 mrn. 0.004-0.01 mrn.
• Dorataspis macropora Haeckel; 1887 (Fig. 6.59). • Hystrichaspis serrata Haeckel, 1887. Radial spi-
Shell thick-walled, sutures hardly visible, surface cules compressed, two-edged, with serrated edges,
pustulose or spiny. Spicules outside shell short, about as long as shell diarneter. Shell with numer-
proximally wide, then narrowing and strongly flat- ous funnel-shaped dimples, majority blind, some
tened. Aspinal pores very large. Sutural pores 8 perforated; 40 aspinal pores of sarne size as sutu-
times smaller than aspinal, funnel-shaped, 1-2 on ral pores. Crests between dimples covered with
each suture. Diarneter of shell (without spicules): denticulate accessory spines. Diarneter of shell:
0.12-0.13 mrn; length of main spicules: 0.06-0.07 0.21 mrn, pores: 0.012 mrn.
mrn. • Icosaspis elegans Haeckel, 1887 (Fig. 6.62).
• "Dorataspis micropora Haeckel, 1887 (Fig. 6.56). Spicules cylindrical, outside shell somewhat com-
Shell massive, ovoid or spherical. Spicules outside pressed, at apophyses twice as wide as inside.
shell long, sword-shaped, basal ends conical, Large-meshed latticed sphere with wide bars.
joined at center into spherical or ellipsoidal body. Aspinal pores large, elongated, surrounded by 2-3
Sutures well marked. Usually 1 sutural pore per cycles of smaller, oval coronal pores, surrounded in
suture, seldom 2 pores, 5-6 times smaller than turn by cycle oflarger sutural pores. Diarneter oflat-
aspinal. Diameter of shell (without spicules): 0.14- ticed sphere (without spicules): 0.3-0.4 mrn; length
0.16 mrn; length of spicules: 0.06-0.07 mrn. of spicules outside of latticed sphere: 0.3-0.4 mrn.
Acantharia 103
• Icosaspis ornata Haeckel, 1887 (Fig. 6.63). • Lychnaspis minima Haeckel, 1887. Parmal pores
Spicules thin, 4-edged, proximal parts with lat- circular, very small, only half as broad as thick
ticed plates which coalesce into sphere with thin bars, much smaller than irregular sutural meshes.
bars. Each spicule with 8 large 8 to 5-sided aspinal Accessory spines (100-110) half as long as radius,
pores, surrounded by 1-2 cyc1es of small, polygo- zigzag. Radial spicules thick, in inner part cylin-
nal coronal pores, surrounded in turn by 1 cyc1e of drical, in outer shorter part conical, of very varia-
sutural pores. Each spicule with 12-16 thread-like ble size. Diameter of shell: 0.05 rnm, coronal
myonemes. Diameter of latticed sphere: 0.24 rnm; pores: 0.002 rnm, sutural pores: 0.006 to 0.012
length of spicules: 0.2 rnm; diameter of cell body: rnm, bars: 0.004 rnm.
0.08 rnm. • Lychnaspis polyancistra (Haeckel, 1860) (Fig.
• Icosaspis serrulata (Haeckel, 1887) (Fig. 6.64). 6.70). Shell with small pores. Spicules inside shell
Parmal meshes 400-500; in center of each plate a cylindrical, outside shell conical, 2 times broader
cross of four very large pentagonal or roundish and longer than inside. Small aspinal pores oval,
aspinal pores, and around this a circ1e of 16 to 20 twice as broad as bars; sutural pores 2 times larger
much smaller irregular, polygonal, coronal pores; than aspinal. Accessory spines half shell radius,
latter about same size as sutural pores. On each zigzag. Diameter of latticed sphere: 0.2 rnm;
condyle one thin zigzag-shaped accessory spine, length of spicules: 0.09-0.1 rnm; length of acces-
nearly as long as radius. Twenty radial spicules sory spines: 0.02-0.03 rnm.
very long, quadrangular, prismatic; on inside thin- • Lychnaspis serrata Haeckel, 1887 (Fig. 6.71).
ner and smooth, on outside thickened, with four Parmal pores pentagonal, three to four times as
rows of recurved teeth, serrated. Diameter of the broad as thick bars, smaller than irregular sutural
shell: 0.18 rnm, aspinal pores: 0.02 rnm, other meshes. Accessory spines (200-300) scarcely one-
pores: 0.002 to 0.01 mm.ibars: 0.003 rnm. third as long as radius, zigzag, with strong denti-
• Lychnaspis giltschi Haeckel, 1887 (Fig. 6.65). c1es. Radial spicules very long, stout, quadrangu-
Spicules thin, cylindrical, pin-shaped, equally thick larly-prismatic (two edged according to Popofsky,
and long inside and outside she1l. Pores of latticed 1906b), with four serrated edges. Diameter of
sphere roundish, bars thin. Aspinal pores pentago- shell: 0.22 rnm, coronal pores: 0.015 rnm, sutural
nal, more rarely oval, 10-20 times larger than bars. pores: 0.012 to 0.03 rnm, bars: 0.004 rnm.
Accessory spines undulating, short (6 times short- • Lychnaspis strelkovi Reshetnjak, 1977 (Fig. 6.68).
er than shell radius). Diameter of sphere: 0.2-0.3 Shell with small pores. Spicules inside shell cylin-
rnm; spicule length (outside sphere): 0.10-0.15 drical, outside shell conical, as long outside as
rnm; length of accessory spines: 0.05 rnm. inside. Aspinal pores of different shape and dispo-
• Lychnaspis longissima Haeckel, 1887 (Fig. 6.66). sition: at each spicule base are 2 small and 2 large
Skeletal spicules inside lattice shell thin, outside pores. Around aspinal 1 cyc1e of 7-12 coronal
very thick and 8-12 times as long as portions pores. Shell diameter: 0.16 rnm; spicule length: 0.7
inside shell; lattice beams wide, round in cross- rnm; diameter oflarge aspinal pores: 0.01 rnm.
section. Aspinal pores small, elongate-ovoid, as • Lychnaspis undulata Haeckel, 1887 (Fig. 6.72).
wide as the lattice beams. Accessory needles long, Shell with broad bars. External part of spicules
four times as long as shell radius. Diameter of compressed, 2-edged, 2 times longer than internal.
shell: 0.08 rnm, coronal pores: 0.002 rnm, sutural Aspinal pores elongated to oval, 2-4 times larger
pores: 0.01 to 0.015 rnm, bars: 0.003 rnm; length than bars. Accessory spines half radius of sphere,
of radial spicules: 0.3 to 0.5 rnm. zigzag. Diameter of latticed sphere: 0.26-0.36
• Lychnaspis· maxima Haeckel, 1887 (Fig. 6.67). rnm; spicule length: 0.13-0.18 rnm; length of
Sphere with large pores, wide bars. Spicules 4- accessory spines: 0.02-0.04 rnm.
edged proximally, cylindrical distally, section out- • Lychnaspis wagenschieberi Haeckel, 1887 (Fig.
side sphere 3-5 times longer than within sphere. 6.69). Skeletal spicules long and thin, quadrangu-
Aspinal pores oval, 5-7 times larger than bars. lar-prismatic or compressed, smooth. Lattice
Accessory spines very long, 2 times shell radius, beams wide and flattened. Aspinal pores triangu-
zigzag. Spicule length: 0.31-0.50 rnm; accessory lar to pentagonal or roundish, three to four times
spines: 0.15-0.20 rnm; shell diameter: 0.6-0.1 rnm. wider than the lattice beams. Accessory needles
104 Acantharia
long, as long as shell radius, with long recurved • Siphonaspis cristata (Haeckel, 1887) (Fig. 6.75).
denticles, very zigzag. Diameter of shell: 0.16 Shell with numerous (100-200?) funnel-shaped
/ rnm, coronal pores: 0.014 rnm, sutural pores: dimples, each pierced at bottom by one of more
0.015 to 0.02 rnm, bars: 0.004 rnm. pores. Twenty larger dimples in center of plates
• Pleuraspis bipennis (Müller, 1858) (Fig. 6.73). pierced by radial spicules; among these 14 contain
Spicules thin, two-edged or quadrangular-prisma- each a couple of aspinal pores; 6 others much lar-
tic, same diameter from base to origin of apophy- ger, and contain each six larger pores; these six
seso Delicate lattice shell spherical, with very large plates are two opposite equatorial plates and four
subcircular to irregularly polygonal pores with polar plates, placed in same meridian plane
thin bars. Basal ends of spicules form a central ("hydrotomical plane"); in each of these six
mass. Sutures give rise to short, smooth accessory "hydrotomical dimples" two pores opposite to one
spines. Diameter of sphere: 0.11 rnm; spicule another on two edges of leaf-shaped spicule, four
length: 0.11 rnm. others opposite in pairs on both flat sides. Twenty
• Pleuraspis constricta (Haeckel, 1887) (Fig. 6.74). radial spicules leaf-like, compressed, of equal
Skeletal spicules strongly flattened, two-edged; length; shell spherical. Crests between dimples are
each spicule twice constricted outside the lattice dentated by series of small accessory spines.
shell, and tapering to a point. Strong apophyses Diameter ofshell: 0.15 rnm, ofaspinal pores: 0.01
twice forked, apophysal ends widening to form rnm, of sutural pores: 0.005 rnm.
knuckle-like joints uniting with those from adja- • Stauraspis echinoides (Haeckel, 1887). Skeletal
cent spicules to form fine-meshed and thick- spicules cylindrical inside the lattice shell, conical
beamed lattice shell. Diameter of shell: 0.11 rnm, outside, with tips sometimes wavy; apophyses
of large meshes: 0.04 to 0.05 rnm, of small mesh- forked twice to thrice, with 16-24 branches per
es: 0.01 rnm; breadth ofspicules: 0.01 rnm. spicule; these branches bearing accessory needles
• Pleuraspis costata (Müller, 1856) (Fig. 6.76). at all distal ends. Apophysál branches connecting
Spicules cylindrical or compressed, broadest with those of neighboring needles to form lattice
where apophyses arise, narrowing gradually shell. Maximum body length: -120 rnm, diameter
toward both ends. Sphere with large pores of irreg- of sphere: 0.14 rnm, spicule length: -0.60 rnm,
ular size and shape, and wide bars, forked once. spicule width: 0.002-0.009 rnm.
Basal ends of spicules forming central mass. • Stauraspis stauracantha (Haeckel, 1862) (Fig.
Accessory spines with barb-like hooks, length 1/3 6.77). Spicule elliptical in cross section, flattened
of spicules. Diameter of sphere: 0.09-0.10 rnm; to flattened quadrangular. Latticed sphere with
spicule length: 0.10-0.15 rnm. thin, bars, forked once or twice from spicules.
• Pleuraspis 'sarmentosa Tchang and Tan, 1964. Pores of two types: large (about 22), and some-
One principal pair of apophyses arising from proxi- what smaller, elongate (about 120). No accessory
mal part of each spicule and participating in for- spines.: 0.11-0.12 rnm; spicule length: 0.1 rnm.
mationof lattice shell; more distally along each • Stauraspis xiphacantha Haeckel, 1887. Radial
spicule, one or more additional pairs of opposing, spicules stout, cylindrical in inner half, conical in
simple or dendroid branches not uniting with one shorter outer half, each producing cross-like quartet
another. Proximal end of each spicule with two ofunbranched apophyses which connect with those
opposite principal apophyses. Each principal of neighboring spicules to form lattice shell with 22
apophysis with two opposite thin, blade-like mesh apertures. Meshes of shell six to eight times
branches uniting to form oval or ellipsoidal shell. as broad as bars. Diameter of shell: 0.12 rnm;
Branches of principal apophyses irregular, pores breadth of spicules and bars: 0.008 to 0.01 rnm.
on surface of shell also irregular. Spicules two-
edged, compressed, all of nearly equal length; in Family Phractopeltidae
early development two of these spicules longer Skeleton is composed of 20 radial spicules joined at
than others. Myonemes 8-35. Maximum body proximal ends, bearing 2 concentric latticed shells
length: 0.43 rnm, maximum spicule length: 0.17 each formed of 20 latticed plates. Distally, spicules
rnm, spicule width: 0.002-0.01 rnm. may bear branched apophyses, which never form third
shell. Endoplasm with pigments and zooxanthellae:
Acantharia 105
genus Phractopelta (Fig. 6.78-6.80). Five species: spicule with 2 large quadrate aspinal pores, and 2
Phractopelta cruciata, Phractopelta dorataspis, smaller oval ones, surrounded by 5-6 two-lobed,
Phractopelta histrix, Phractopelta stauropora, polygonal irregularly shaped sutural pores. Cell
Phractopelta tessaraspis. body spherical, dark cherry red, enclosed in outer
• Phractopelta cruciata (Haeckel, 1887) (Fig. 6.78). shell. Ectoplasmic cortex single-layered. Six-eight
Apophyses of long polar and tropical spicules short myonemes per spicule. Diameter of cell
crossed by transverse beam, which is again crossed body: 0.06-0.07 rnm; spicule length: 0.27 rnm;
by two perpendicular branches parallel to apophy- diameter of outer shell: 0.09-0.097 rnm, inner:
ses; branches with free ends, not coalescing. At 0.03-0.037 rnm; large aspinal pores: 0.016 by
base of each spicule (in outer shell) two large ellip- 0.014 rnm, small aspinal pores: 0.012 by 0.06-0.08
tical aspinal pores, larger than other irregular rnm, sutural pores: 0.012 by 0.008 rnm.
pores. Four equatorial spicules simple, without
apophyses, of same size as other sixteen. Diameter Family Diploconidae
of outer shell: 0.12 rnm, ofinner 0.05 rnm. Skeleton consisting of 20 radial spicules of different
• Phractopelta dorataspis Haeckel, 1887 (Fig. shape and different length, among which 2 equatorial
6.79). Spicules 2 times longer than external shell spicules are longer and stronger than other 18, each of
diameter, cylindrical, tapering to sharp tips, usual- 2 main spicules with cornet. Spicules ' pierce thick,
ly of equal size and shape; their basal ends provid- ellipsoidal, armored shell. Around each spicule on
ed with small pyramids which coalesce into sphe- upper part of shell are 2 oval aspinal pores surround-
rule in center of the shell. Outer shell spherical to ed by 5-6 sutural pores. Cornets striated longitudinal-
ellipsoidal, 2.5-3 times larger than internal spheri- ly and serrate at edge. Endoplasm with nuclei, lipid
cal shell. Two aspinal pores oval elongate or reni- inclusions, and zooxanthellae. Axopods pass across
form, very variable in shape and size. Coronal and sutural pores. Periplasmic cortex suspended from
sutural pores elongated, irregular. Diameter of apex ofsmall spicules by 6-12 myonemes: One genus:
external shell: 0.08-0.09 rnm, internal: 0.03-0.035 Diploconus (Fig. 6.81-6.85). five species: Diploconus
rnm; spicule length: 0.17 rnm. cyathiscus, Diploconus cylindrus, Diploconus fasces,
• Phractopelta histrix (Müller, 1856). Spicules elon- Diploconus hexaphyllus, Diploconus satumus.
gate-conical outside outer shell, as long or twice as • Diploconus cyathiscus Haeckel, 1887 (Fig. 6.81).
long as outer shell diameter. Pair of small knob-like Mande of double cone thin walled, not com-
or forked apophyses often on some or all spicules pressed; circular in cross section, .contour strongly
outside outer shell. Basal ends of spicules fused into convex, surface with six stronger and many small-
central mass. Shell spherical, thick-walled, pores er ribs, margin with numerous, straight and long,
circular, regular. Each spicule surrounded by 4 aspi- parallel denticles. Diameter of mouth half as long
nal pores, about 2-3 larger on outer shell than on as whole shell, one and a halftimes as long ~s equa-
inner one. Diameter of outer shell: 0.075 rnm, inner torial diameter. Two main spicules prismatic, one-
shell: 0.03 rnm; spicule length: 0.65 rnm. fourth longer than conical sheath or cornet. Sixteen
• Phractopelta stauropora (Haeckel, 1887). smaller spicules about half as long, thin, straight.
Apophyses of eight tropical spicules crossed by Two quadrangular-prismatic equatorial spicules
transverse beam, which is again crossed by two thick and short. Length of shell: 0.2 rnm; equatori-
perpendicular branches; by union of these branch- al breadth: 0.07 rnm, polar breadth: 0.1 rnm.
es each tropical spicule forms square shield with • Diploconus cylindrus Haeckel, 1887 (Fig. 6.82).
four crossed pores and 12 to 16 marginal spikes. Central shell small, pitted, without sutural fur-
On base of each spicule (in surface of outer shell) rows. Cornets very long, cylindro-conical, encas-
four crossed aspinal pores which correspond to ing up to 3/4 of main spicules, distal parts 1.5
Ithose of free shields; somewhat larger than other times wider than the proximal ones, with many
pores. Diameter of outer shell: 0.12 rnm, of inner: (> 15) longitudinal wavy ribs, 6 of which are more
0.06rnm. conspicuous than others. On distal edge of cornet
• Phractopelta tessaraspis Haeckel, 1887 (Fig. ribs extend into spines. Main spicules 6-edged,
6.80). Spicules of external shell compressed, two- their free ends pyramidal, tips sharp. Eighteen se-
edged, with 2 opposite branching apophyses. Each condary spicules of equal length, needle-like.
106 Acantharia
Length of main spicules: 0.16 mm; secondary: One genus: Lithoptera (Fig. 6.86, 6.87). Two species:
0.05 mm; diameter of central shell: 0.04 mm; Lithoptera fenestrata, Lithoptera muelleri.
length of cornets: 0.11 mm. • Lithoptera fenestrata Müller, 1858 (Fig.-6.86).
• Diploconus fasces Haeckel, 1860 (Fig. 6.83). Distal ends of 4 main (equatorial) spicules form-
Central shell ellipsoidal, small, thick-walled, ing large, triangular, latticed plates with quadran-
sharply set off from cornets. Cornets conical, thin- gular pores. Tropical spicules about 1/3 as long as
walled. Main spicules cylindro-conical, with lon- equatorial, polar 1/2 as long, very thin, with sharp
gitudinal keels. Secondary spicules 2 times short- tips, basal part pear-shaped, inflated. Distal ends
er than main, strongly compressed. Length of ofsecondary spicules with 1-2 pairs ofthin oppos-
main spicules: 0.1 mm; secondary: 0.05 mm; ing apophyses at right angles, which in tum also
diameter of central shell: 0.03-0.04 mm; length of bear apophysesat right angles; these branches
cornets: 0.05-0.06 mm. coalesce into a delicate meshwork. The number
• Diploconus hexaphyllus Haeckel, 1887 (Fig. and degree of development of apophyses varies
6.84). Mantle of double cone compressed, thick widely (may be absent altogether in young speci-
walled, with elliptical transverse section; its con- mens); this variability led Haeckel (1887) to estab-
tour straight; surface with six stronger and nume- lish several species, which were subsequently syn-
rous smaller straight ribs and six marginal dentic- onymized by Schewiakoff (1926) into L.
ulate incisions. Central shell nearly spherical. fenestrata. Length of main spicules: 0.24 mm;
Diameter of mouth one-third as long as whole width of latticed meshwork: 0.32 mm; length of
shell and somewhat smaller than equatorial diam- tropical spicules: 0.15 mm, polar: 0.12 mm.
eter. Larger equatorial spicules about twice as long • Lithoptera muelleri Haeckel, 1860 (Fig. 6.87).
as sheath, pyramidal, with emarginate point and Four strongly compressed main spicules with
six wings. Smaller equatorial spicules short and large, wing-like latticed apophyses forming solid,
strong, pyramidal. Sixteen other spicules thin, centrally somewhat concave, meshwork. Sixteen
straight, conical, Length of shell 0.3 mm; equato- secondary spicules short, forming delicate mesh
rial breadth 0.11 mm, polar breadth 0.1 mm. with their apophyses. Main spicules with 8 pairs of
• Diploconus saturnus Haeckel, 1879 (Fig. 6.85). myonemes; secondary with 5-6 pairs. Cell body
Central shell ellipsoidal, less often spherical, sur- quadrangular. Shell: 0.42 mm; cell body: 0.18
face smooth, sutural furrows conspicuous. mm; length of main spicules: 0.21 mm, tropical
Secondary spicules almost as long as pyramidal spicules: 0.12-0.13 mm, polar spicules: 0.09 mm.
main spicules, pin-like, curved. Cornets bell-
shaped, surrounding main spicules to about half Family Hexalaspidae
their length; c1early delimited from central shell; Among 20 spicules, 2-6 differ in shape, longer and
twice as broad at distal end as at the base; surface stronger than others. Spicules assembled into central
covered by many, (>.16) ribs which extend into mass resembling star, at half-length with thick
small thorns at edge of cornet. Length of main spic- armored plates with numerous spines at external sur-
ules: 0.08 mm; secondary: 0.05 mm; diameter of face and pores edged with serrate rings. Plates also
central shell: 0.04 mm; length of cornets: 0.03 mm. with sheets surrounding all spicules or on1y major
ones. Endoplasm in armored shell. Capsular wall out-
Fami1y Lithopteridae side of shell. Eight to 12 myonemes.
Family .characterized by four spicules longer and
thicker than others, with flat, triangular, latticed plates Key to genera of Hexalaspidae
on apical third of length. Each plate of 1-5 lateral 1 Two main spicules (pair of opposing equatorial
apophyses cross-linked by 3-5 orthogonal bars delin- spicules) longer, wider, and/or differently shaped
eating square-based lattice. Two species of this fami- from remaining 18 spicules; 18 spicules equal;
ly flattened in plane of largest spicules. Endoplasm sheaths or sleeves around all skeletal spicules
with numerous zooxanthellae and small nuc1ei, limit- encasing not over one-third of length of needle.
ed by thick fibrillar meshwork through which axopo- Armor shell ellipsoidal: genus Coleas pis (Fig.
dia pass. Traffic of tiny kinetocysts active along axo- 6.88, 6.89). Three species, Coleaspis coronata,
podia. Myonemes short, cylindrical, not very mobile. Coleaspis obscura,' Coleaspis vaginata
Acantharia 107
la Six main spicules (two opposing equatorial spi- edges of pits protruding, forming sheaths with ser-
cules and 4 polar spicules all in same plane, the rated crests. Bases of spicules also surrounded by
so-called "hydrotomical plane" ofHaeckel, 1887) sheaths, equally large or nearly so, extending to
distinctIy longer, thicker and with different shape 1/4-1/3 of the spicule length; sheaths around main
than other 14 more rudimentary spicules .2 spicules better developed than those around secon-
2 Skeletal spicules lacking sheaths, with accessory dary ones; sheath rims producing long pin-like
spines, armor shell lens or disc shaped covered accessory spines. Within sheath, two opposing
with jagged circles of accessory spines: genus ribs extend from narrower edges of spicules,
Hexalaspis (Fig. 6.96, 6.97). Two species: dividing aspinal pit into 2 parts, with 1 aspinal
Hexalaspis heliodiscus, Hexalaspis sexalata pore in each. Aspinal pits surrounded by 8-12
2a Sheaths around all spicules or only around six smaller sutural pits; all pits bordered with small,
main spicules; sheaths significantIy better devel- pin-like accessory spines. Diameter of shell
oped on latter. Armor shell ellipsoidal, rarely (excluding spicules): 0.12 by 0.11 rnm; length of
spherical: genus Hexaconus (Fig. 6.90-6.95). Six equatorial spicules: 0.11-0.13 rnm, polar: 0.11
species: Hexaconus ciliatus, Hexaconus echina- rnm, secondary: 0.09 rnm.
tus, Hexaconus infundibulum, Hexaconus nival- • Hexaconus echinatus Haeckel, 1887 (Fig. 6.91).
is, Hexaconus serratus, Hexaconus trypanon Six main spicules of unequal size; tWo equatorial
with sheaths twice as large as the four polar spi-
• Coleas pis coronata (Müller, 1858) (Fig. 6.88). cules. These six spicules six-edged, pyramidal, with
Shell ovoid, very thick-walled. All 20 spicules of basal half enveloped by very large, conical, sulcate
nearly equal form, 2 main slightIy longer and sheaths, twice as broad on dentate distal mouth as
thicker. Crests between dimples of shell-wall ser- on narrower base. Other fourteen spicules very thin,
rated; comets around spicules short, with regular- two-edged, half as long, with low sheaths. Diameter
ly spiny edges. Shell diameter: 0.38-0.4 rnm (0.17- of shell: 0.15 rnm; length of equatorial spicules:
0.2 rnm according to Haeckel, 1887); length of 0.13 rnm, ofpolar spicules: 0.08 rnm.
main spicules: 0.14-0.19 rnm, secondary: 0.12- • Hexaconus infundibulum (Haeckel, 1887) (Fig.
0.15 rnm. 6.92). Six main spicules of unequal size, two equa-
• Coleaspis. obscura Haeckel, 1887. Shell ovoid, torial about twice as long as four polar; all six spi-
very thick-walled, dark, opaque. Two main spicules of same form, quadrangular prismatic, point-
cules slightIy to 1 1/2 times longer than other 18. ed at distal pyramidal end. Sheaths conical, two to
Comets around spicules almost reaching spicule three times as broad at denticulate distal mouth as
ends, with several longitudinal keels, their edges at narrow base, sulcate; mande of cone concavely
with strong spines. Shell diameter: 0.08-0.1 rnm; vaulted. Large sheaths envelop two-thirds or three-
main spicules: 0.05-0.06 rnm. fourths of spicules. Diameter of shell: 0.1 to 0.13
• Coleaspis vaginata Haeckel, 1887 (Fig. 6.89). Shell rnm; length of equatorial spicules: 0.15 to 0.2 rnm,
ovoid, thick-walled. All twenty spicules of nearly of sheaths: 0.1 to 0.15 rnm; basal breadth oflatter:
equal size and form, nearly as long as diameter of 0.04 rnm, distal breadth: 0.12 rnm.
shell, conical; larger equatorial spicules cylindrical • Hexaconus nivalis (Haeckel, 1887) (Fig. 6.93). All
or conical, with three longitudinal keels and up to 6- six Main spicules of nearly equal size and equidis-
sided; about one-third longer than remaining 18 tant, thin, lanceolate, compressed, somewhat short-
conical or two-edged spicules. Crests between dim- er than diameter of hexagonal shell. Sheaths nearly
pIes of shell-surface serrated, forming truncate con- prismatic, conical in distal third, each with twelve
ical sheath with crested surface and serrated mouth prominent parallel crests, separated by twelve deep
around basal half of each spicule, with teeth pro- furrows; contracted distal opening or mouth denti-
longed into short accessory spines. Major axis of culate. Sheaths as long as radius of shell, envelop-
shell 0.12 rnm, secondary 0.1 rnm; length of spic- ing two-thirds of spicules. Sutural pits polygonal,
ules: 0.08 rnm, basal breadth 0.01 rnm. separated from one another by protruding crenulate
• Hexaconus ciliatus Haeckel, 1887 (Fig. 6.90). border. Diameter of shell: 0.15 rnm, thickness of
Shell thick-walled, ellipsoidal, surface covered wall: 0.08 rnm; length of spicules: 0.1 rnm, of
with circular aspinal pits with oval aspinal pores, sheath: 0.007 rnm; breadth oflatter: 0.04 rnm.
108 Acantharia
• Hexaconus serratus Haeckel, 1887 (Fig. 6.94). Suborder Phyllacantha

Shell thick-walled, ellipsoidal, surface covered Family Phyllostauridae _
/ with circular aspinal pits with kidney-shaped aspi- Skeleton of 20 spicules without apophyses, either of
nal pores, edges of pits protruding, forming same length and similar shape or different, 2'-4 longer
sheaths reaching 1/2 spicule length. Sheaths with and stronger than others. Base of each spicule with 4
longitudinal ribs ending in large spines at sheath large lateralleaf-shaped extensions (leaf-cross), cruci-
margino Sheath on secondary spicules less deve- form in cross section. Cell body surrounded by thick
loped. Size of shell (exc1uding spicules): 0.16 by capsular wall, spherical, oblong or quadrangular,
0.14 rnm; lengthofequatorial spicules: 0.15 rnm, looking like cushion, consisting of endoplasm with
polar: 0.13 rnm, secondary: 0.1 mm; length of various inc1usions and peripheral ectoplasm.
sheaths on main spicules: 0.02-0.04 rnm, on se- Axopodia arise from ectoplasmic cortex between
condary: 0.01 rnm. spicules. Ectoplasmic cortex of polygonal pieces con-
• Hexaconus trypanon (Haeckel, 1887) (Fig. 6.95). nected by elastic junctions.
Six main spicules of unequal size, two equatorial
about as long as diameter of shell and near1y twice Key to genera of Phyllostauridae
as large as four polar; all of same form, quadran- 1 Spicules of same length and same shape. One or
'.gular prismatic, with prominent edges and pointed two spicules may be longer than others: genus
distal ends. Sheaths six-sided prismatic, with Phyllostaurus (Fig. 6.109-6.113). Four species:
strong prominent edges and thinner parallel ribs Phyllostaurus cuspidatus, Phyllostaurus echi-
between them; distal mouth with six strong dentic- noides, Phyllostaurus quadrangulus, Phyllo-
ulated teeth. Sheaths envelop two-thirds of spi- staurus siculus
cules,as broad as radius of shell. Sutural- pits 1a Two to four spicules longer than others, of dif-
round, surrounded by somewhatknob-like acces- ferent shape 2
sory spines. Diameter of shell: 0.13 rnm; length of 2 Two spicules longer than others, of different
equatorial spicules: 0.12 rnm, of sheaths: 0.08 shape. Endoplasm oblongo 16-20 myonemes per
rnm; breadth oflatter: 0.06 rnm. spicule: genus :Amphistaurus (Fig. 6.103-
• Hexalaspis heliodiscus Haecke1, 1887 (Fig. 6.96). 6.106). Four species: Amphistaurus atlanticus,
Shell thick-walled, opaque, lenticular or discoidal, Amphistaurus complanatus, Amphistaurus lan-
with short spicules. Surface covered with subcir- ceolatus, Amphistaurus tetrapterus
cular to oval deep, funnel-shaped pits. Six main 2a Four spicules longer and stronger than 16 others,
spicules equal in shape and size, massive, usually which are identical or different .3
smooth, less often faintly edged or keeled, tips 3 Four main spicules identical, 16 others of differ-
blunt1y rounded, sometimes two equatorial spi- ent shape, though all similar, or polar spicules a
cules longer and thicker than other four main little shorter and flattened than tropical spicules.
(polar) spicules. Secondary spicules much shorter Endoplasm quadrangular, red, 16-30 myonemes:
and thinner than main (they brake off and dissolve genus Acanthostaurus (Fig. 6.98-6.102). Four
easi1y, therefore often lacking). Diameter of shell species: Acanthostaurus bipennis, Acantho-
(exc1uding spicules): 0.11 rnm; length of main staurus conacanthus, Acanthostaurus henseni,
equatorial.spicules: 0.1 rnm, polar: 0.08 rnm, sec- Acanthostaurus purpurascens
ondary: 0.06 rnm. 3a Main spicules different from one another .4
• Hexalaspis, sexalata Haeckel, 1887 (Fig. 6.97). 4 Two main opposite spicules identical two by
. Six main spicules of unequal size; two equatorial two. Endoplasm oblong to rhomboidal: genus
spicules six-sided, about as !long as shell-radius, Lonchostaurus (Fig. '6.107, 6.108). Two spe-
twice 'as long as four polar spicules, which are cies: Lonchostaurus bifurcus, Lonchostaurus
three-edged or pyramidal. Fourteen other spicules rhombicus
only half as long and one-fourth as broad as latter, 4a Rhomboidal acantharians. Only 2 main spicules
little prominent. Central shell spherical. Diameter along transversal axis are identical with one
of shell: 0.15 rnm; length of 2 equatorial spicules: another; other 2 spicules along longitudinal axis
0,08 rnm, of 4 polar spicules: 0.04 rnm, of 14 other are different. Tropical and polar spicules also
spicules: 0.02 rnm. ., different: genus Zygostaurus (Fig. 6.114,
Acantharia
6.115). Three species: Zygostaurus amphitectus, spicule. Central capsule light-colored. Length of
Zygostaurus cornutus, Zygostaurus sagittalis main spicules: 0.7-1.33 mm, width: 0.008 mm;
length of secondary: 0.2-0.35 mm, width: 0.002-
• Acanthostaurus bipennis Haeckel, 1887 (Fig. 0.003 mm.
6.98). Main spicules linear, sword-shaped, slight- • Amphistaurus complanatus (Haeckel, 1860) (Fig.
ly to strongly compressed, two- to four-edged; of 6.105). Two main spicules cylindrical to very
nearly equal breadth over whole length. Apex slightly compressed, linear, of equal breadth
bifid. Remaining spicules strongly compressed, throughout whole length; apex emarginate or
lanceolate. Base with smallleaf-cross. Four equa- bifid; base with large leaf-cross of double breadth.
torial spicules of same form as sixteen others, but Eighteen smaller spicules strongly flattened, two-
twice as long and as broad. About 20 myonemes. edged, needle-like along narrow side, one-fourth
Length of 4 major spicules: 0.4 to 0.5 mm, to one-half as large. Central capsule ellipsoidal,
breadth: 0.02 mm; length of 16 secondary: 0.2 to yellow, pellucid. Length of2 major spicules: 0.15
0.3 mm, breadth: 0.01 mm. to 0.25 mm, breadth: 0.004 to 0.006 mm; length of
• Acanthostaurus conacanthus Haeckel, 1887 18 secondary spicules: 0.05 to 0.1 mm.
(Fig. 6.100). Main spicules conical, tapering dis- • Amphistaurus lanceolatus (Haeckel, 1887) (Fig.
tally, ending in blunt tips. Four primary spicules 2 6.106). Two main spicules compressed, lanceolate,
times longer and 3 times thicker than sixteen se- two-edged, gradually tapering from broader mid-
condary; latter strongly compressed, with sharp dle towards two ends; apex simple; base a large
tips. Polar spicules sometimes shorter than tropi- pyramid, with broad leaf-cross. Eighteen smaller
cal. Cruciform leaf-cross especially well devel- spicules about half as long, pyramidal, with simple
oped on equatorial and tropical spicules, forming conical or bristle-shaped apex. Central capsule len-
large leaf-star almost filling central capsule. telliptical, pink, 'opaque, Length of 2 major spic-
Length of main spicules: 0.15-0.20 mm; secon- ules: 0.2 mm, of 18 secondary: 0.1 mm.
dary: 0.09-0.10. • Amphistaurus tetrapterus (Haeckel, 1860). Main
• Acanthostaurus henseni (Popofsky, 1905) (Fig. spicules 4-winged, of equal length. Secondary
6.101). Four main equatorial spicules broad, 4- spicules 1/2 to 2/3 as long as rnain, 4-winged. All
winged in cross section, pyramidal. Secondary spicules broaden near base, tapering distally, with
spicules of same length or slightly longer, com- 20 myonemes. Central capsule ellipsoidal, com-
pressed but not two-edged, tips blunt. Cell body pressed, opaque, yellow to dark brown. Length of
quadrangular. Length of main spicules: 0.09 mm; main spicules: 0.12-0.2 mm, width: 0.013-0.025
secondary: 0.07-0.1 mm. mm; .secondary, length: 0.06-0.15 mm, width:
• Acanthostaurus purpurascens (Haeckel, 1860) 0.08-0.016 mm; central capsule, major axis: 0.1-
(Fig. 6.102). Main spicules cylindrical, narrowing 0.15 mm, minor axis: 0.08-0.1 mm.
distally, slightly compressed, les s often keeled or • Lonchostaurus bifurcus Haeckel, 1887 (Fig.
4-bladed in cross section basally or distally, tips 6.107). Four equatorial spicules compressed, nearly
rounded. Sixteen secondary spicules 1.5 times rectangular, forked; one pair of opposing equatorial
shorter than main, more strongly compressed, spicules with two, other pair with four homs; two
two-edged; 8 polar spicules may be less com- opposite spicules of longitudinal axis about: one-
pressed and shorter than 8 tropical. Equatorial third longer and broader than those of transverse.
spicules with 4 leaf-crosses, tropical and polar Sixteen smaller spicu1es of same form, 'also forked,
with 2. Leaf-crosses on equatorial and tropical eight tropical half as large, forked twice; and eight
spicules 3 times broader than maximum spicule polar spicules scarcely one-fourth as large as two
diameter. Length of main spicules: 0.3-0.4 mm, longitudinal spicules, forked once. Length of equa-
secondary: 0.2-0.3. torial cross: 0.36 mm, breadth: 0.26 mm.
• Amphistaurus atlanticus (Popofsky, 1904) (Fig. • Lonchostaurus rhombicus (Haeckel, 1887) (Fig.
6.103). Main spicules ofequallength, 4-winged in 6.108). Two equatorial spicules largest, cylindri-
cross section (wings narrow), distally expanded, calor slightly swollen in the middle - spindle-
tips sharp. Eighteen secondary spicules shorter shaped, sometimes slightly compressed, 4-sided in
and thinner, tips pointed. Twenty myonemes per cross section; distal part . tapering, tips blunt,
110 Acantharia
rounded. Other 2 equatorial spicules shorter but slightly bent homs; other, directed toward back
wider, may bear 2 longitudinal keels in the distal (posterior), may be spindle-, lance- or sword-
part, in which case they are pyramidal, or may be shaped, and may also bear 2 diverging or parallel
sword-shaped. Sixteen secondary spicules shorter homs. Tropical spicules much shorter, ending in 2
than equatorial, tapering distally, compressed, small diverging homs. 'Polar spicules arborescent,
two-edged, tips sharp; may be all alike, or 8 polar reduced in size and number, may be rudimentary.
may be slightly shorter and thinner than 8 tropical. Central capsule rhomboidal or pillow-shaped.
Leaf-star very large, occupying almost entire cen- Length of equatorial spicules: 0.4-0.8 mm.
tral capsule. Length of larger equatorial spicules: • Zygostaurus cornutus Haeckel, 1887. Anterior
0.1 mm, smaller: 0.08 mm; secondary spicules: spicule little-different from two lateral spicules,
0.06 mm; breadth of leaf-cross: 0.06 by 0.04 mm. each with two divergent straight homs, shorter
• Phyllostaurus cuspidatus (Haeckel, 1862) (Fig. than basal parto Posterior spicule simple, triangu-
6.110). All spicules of equal size and shape, 4- lar, shorter than three former. Eight tropical spi-
winged, tips sharp, with small leaf-cross. Cell cules of equal size and form, doubly forked, each
body spherical, light yellow to light brown. with four thin, bristle-shaped, little divergent teeth.
Ectoplasmic cortex. single-layered. Each spicule Eight polar spicules short, simply forked. Length
with 40 myonemes. Diameter of cell body: 0.1- of equatorial cross: 0.5 mm, breadth: 0.4 mm.
0.16 mm; spicule length: 0.4-0.6 mm; breadth of • Zygostaurus sagittalis Haeckel, 1887 (Fig. 6.115).
leaf-cross: 0.012-0.08 mm. Anterior spicule very different from others, with
• Phyllostaurus echinoides (Claparede, 1855) (Fig. two equal, strongly divergent, bent homs. Lateral
6.111). AlI spicules of equallength, 4-edged, tips spicules little smaller, but with two very unequal
sharp. Cell body spherical. Central capsule homs (anterior shorter than posterior). Posterior
opaque. Spicule length: 0.3-0.5 mm; breadth of spicule simple, sword-like or triangular, two-
leaf-cross: 0.02-0.03 mm. edged, longer than anterior spicule. Eight tropical
• Phyllostaurus quadrangulus (Haeckel, 1887) spicules different, four anterior smaller, with long-
(Fig. 6.112). Spicules of equal shape and size, 4- . er homs; four posterior broader, with shorter homs;
winged, very wide in proximal part (inside central anterior hom of each tropical spicule longer than
capsule), tips sharp, leaf-cross very large. Cell posterior. Eight polar spicules rudimentary, simple,
body spherical; bright red. Diameter of cell body: very short. Length of equatorial cross: 0.6 mm,
0.09 mm; length of spicules: 0.18-0.36 mm; breadth: 0.4 mm.
breadth of leaf-cross: 0.024 mm.
• Phyllostaurus siculus (Müller, 1858) (Fig. 6.113). Family Stauracanthidae
Spicules cylindrical, equal in size and shape, taper- Skeleton of 20 spicules of equallength and generally
I ing distally, tips sharp; more rarely compressed, same shape, with 4 longitudinal wings so that spicule
flexible. Leaf-cross 2-3 times broader than spi- is cruciform in cross section. Wings bear simple or
,I cules. Sometimes spicules of equal width through- branched apophyses which may link with 4 or more
,I
out length, leaf-cross up to 4-6 times broader than apophyses of neighboring spicules forming latticed
spicules. In some specimens 2 opposing equatorial shell. Bases of spicules with 4 lateral blade-shaped
spicules 1.5 times longer than others, yet shape and extensions delineating deep cavities between spicules.
width similar to that of other 18 spicules. Endoplasm spherical to polygonal, surrounded by
Sometimes pyramidal bases of spicules very long, thick capsular wall. Periplasmic cortex thick, with
almost reaching central capsule, 4-edged within elastic junctions, linked to spicule by 40-60 myo-
central capsule; thin, needle-like outside. Spicule nemes resembling long cylindrical threads.
length: ; 0.3-0.4 mm, length of basal pyramids:
0.024-0.03 mm. Two subspecies: Phyllostaurus Key to genera of Stauracanthidae
siculus catervatus Schewiakoff, 1926, and Phyllo- 1 Spicules with apophyses, complete lattice plates
staurus siculus quadrifo/ius Schewiakoff, 1926. absent 2
• Zygostaurus amphitectus Haeckel, 1887 (Fig. la Spicules bearing rectangular-pored latticed
6.114). Two main vertical spicules of different plates orthogonal to direction of spicule. Each
shape: one directed forwards, with 2 diverging, plate consisting of series of orthogonal apophy-
! I
Acantharia 111
ses: genus Phatnacantha (Fig. 6.116). One spe- • Pristacantha octodon Haeckel, 1887 (Fig. 6.117).
cies: Phatnacantha icosaspis Each spicule bears 8 apophyses arranged in 2 ver-
2 Apophyses unbranched 3 ticils, longest apophyses are closer to base of spi-
2a Apophyses branched. Secondary and tertiary cule, getting shorter distally. Length of spicules:
apophyses at right angles, crossing but never 0.64-0.65 mm; length of longest apophyses:
fonning latticed plates: genus Stauracantha 0.012-0.028 rnm, shortest: 0.009-0.012 rnm;
(Fig. 6.122). Two species: Stauracantha orthos- length ofbasal pyramids: 0.028-0.038 rnm.
taura, Stauracantha spinu/osa • Pristacantha polyodon Haeckel, 1887 (Fig.
3 Each spicule bearing 4 crosswise apophyses 6.119). Spicules in distal half four-sided prismatic,
which can resemble thoms, stumps, or wings: thin, with pyramidal apex; in basal half much
genus Xiphacantha (Fig. 6.123, 6.124). Three broader, four-winged, with large leaf-cross at
species: Xiphacantha a/ata, Xiphacantha multi- base. From four wings ofbasal halfvariable num-
spina, Xiphacantha quadridentata ber of slender teeth or triangular apophyses, com-
3a On each spicule 2-6 groups of apophyses, never monly sixteen to twenty-four, often irregular (four
crosswise: genus Pristacantha (Fig. 6.117- to six from each wing). Length of spicules: 0.3 to
6.120). Four species: Pristacantha dodecodon, 0.4 rnm, breadth in distal part: 0.007 to 0.01 rnm,
Pristacantha mu/tidentata, Pristacantha octodon, in basal part: 0.02 to 0.03 rnm. .»:
Pristacantha po/yodon • Stauracantha orthostaura Haeckel, 1887 (Fig.

6.122). Spicules long or short; usually short, in
• Phatnacantha icosaspis Haeckel, 1887 (Fig. basal third (to point where apophyses are inserted)
6.116). In proximal l/4th spicules with quadran- of uniform thickness, prismatic, from there on
guIar latticed plates of thin, compres sed bars. At tapering, tips sharp. Less often specimens with
first bars form 4 large pores around each spicule, long, needle-like spicules, as well. as short and
subsequent development of bars forms 8 coronal long spicules. Apophyses very variable in shape.
pores around aspinal ones. Both aspinal and coro- Primary branches of apophyses cylindrical or
nal pores may be further divided by additional slightly compressed; .approximately from center
growth of bars within them. Growth of latticed arise short, opposing secondary branches, which
plates may be asynchronous on different spicules. may in turn give rise to perpendicularly inserted,
Leaf-star large, occupying almost entire central short tertiary branches of irregular distribution.
capsule. Spicule length: 0.32-0.40 rnm; latticed Sometimes primary branches give rise to several
plates: 0.09 by 0.09 rnm; leaf-cross: 0.04 rnm. (rather than 1) pairs ofsecondary ones. Secondary
• Pristacantha dodecodon Haeckel, 1887 (Fig. and tertiary branches may cross each other at right
6.118). Spicules four-winged, of lanceolate out- angles, but never coalesce (as in the genus
line, tapering from broader middle towards two Phatnacantha). Leaf-star occupying a small por-
ends. Apex pyramidal. Base with large leaf-star. tion of central capsule. Spicule length: 0.35-0.45
From four wings arise in basal part (between first rnm; primary branches: 0.05-0.08 rnm; secondary:
and second third of length) twelve triangular 0.016-0.03 rnm, tertiary: 0.008-0.01 rnm.
apophyses (three from each wing, arranged in • Stauracantha spinulosa (Haeckel, 1860). Spicules
crowns). Length of spicules: 0.3 rnm, breadth in stout, four-sided prismatic, of nearly equal breadth
middle: 0.03 rnm. throughout whole length, on distal apex truncated,
• Pristacantha multidentata Schewiakoff, 1926 two-edged. Base with large wing-cross. Four
(Fig. 6.120). Spicules of equal size and shape, 4- apophyses about in middle of each spicule, conical,
winged, tapering distally; edges in basal l/4th straight, spinulate, twice to four times as long as
smooth, distally with hooks or thoms, first one to basal breadth of spicule. Central capsule opaque,
three quartets needle-like; each with 40 thread-like yellow. Length of spicules: 0.27 rnm, breadth:
myonemes. Basal pyramids small. Cell body 0.014 rnm; length ofapophyses: 0.05 rnm.
spherical, central. capsule opaque, brown-red. • Xiphacantha alata (Müller, 1858) (Fig. 6.123).
Ectoplasmic cortex single-Iayered. Diameter of Spicules equal in size and shape, 4-winged, in
cell body: 0.2-0.3 rnm; spicule length: 0.8-1.3 basal fourth with four large crosswise lamellar
rnm; breadth of basal pyramids: 0.028-0.03 rnm. wings, wing edges with fme serrations; each with
50 thread-like myonemes. Basal leaf-star small.
[
112 Acantharia
Cell body spherical or polygonal, central capsule Spicule length: 0.2-0.36 rnm; diameter of lattice
thick, opaque, brown-yellow or brown-green, shell: 0.2-0.3 rnm; diarneter of leaf-cross: 0.032
Diarneter of cell body: 0.2 rnm; length of spicules: rnm; diarneter of cell body: 0.14 rnm.
0.64-0.80 rnm; width of wings: 0.024-0.068 rnm;
length ofbasalleaf-star: 0.015-0.037 rnm.
• Xiphacantha multispina (Müller, 1858) (Fig. Acknowledgments
6.121). Spicules equal in size and shape, proximal-
ly 4-winged, distally 8-winged, tips sharp, from This work is based partly on an earlier manuscript ver-
middle 8 short stump-like apophyses; each with 40 sion prepared by Anthony Michaels, which in turn was
thread-like myonemes. Basal leaf-star very large. derived from a surnmary written by C. Febvre, J.
Cell body polygonal, central capsule red-brown, Febvre and A. Michaels. We are indebted to A.
Diarneter of cell body: 0.12 rnm; length of spic- Michaels for putting this initial draft and other pertinent
ules: 0.25 rnm, width: 0.017 rnm; length of basal materials at our disposal, as well as to John Lee for pro-
leaf-star: 0.06 rnm. viding unpublished graphic material s for consultation
• Xiphacantha quadridentata (Milller, 1858) (Fig. purposes. The assistance of Georges Merinfeld for the
6.124). Spicules equal in size and shape, 4-winged, identification of several of the SEM images is greatly
tips sharp, from their basal third arise 4 short appreciated. Michiko Matsubara provided efficient
opposed apophyses; each with 40-60 myonemes. technical assistance. This work was partly financed by
Basal leaf-star very large. Cell body spherical, cen- grants from the University ofBuenos Aires (UBA EX-
tral capsule red-brown, Diarneter of cell body: 0.14 040), from the Consejo Nacional de Investigaciones
rnm; length of spicules: 0.45-0.8 rnm, width at base: Científicas y Técnicas, Argentina (CONICET PID-BID
0.009-0.016 rnm, at level of apophyses: 0.009-0.012 0366), and from the Agencia Nacional de Promoción
rnm; length ofbasalleaf-star: 0.024-0.028 rnm. Científica y Tecnológica, Argentina (PMT-PICT0395).
Farnily Dictyacanthidae
Skeleton of 20 spicules of sarne length and sarne shape, Suggested readings
with latticed plates which link with each other to form
latticed shell. Cell body spherical, surrounded by thick Febvre c., Febvre J., Michaels A. In press. Subphylum
capsular wal1, with several nucIei, pigments, and zoo- Acantharia Haeckel, 1881. In "An illustrated guide
xanthellae. Ectoplasm limited by thick periplasmic cor- to the Protozoa" Second Edition, (J. Lee, ed.).
tex with radial and concentric elastic junctions, sus- Overview of acantharian biology and classifica-
pended from spicules by 50 thread-like myonemes. One tion with examples ofrepresentative species.
genus: Dictyacantha (Fig. 6.125,6.126). Two species: Haeckel E. 1887. Report on Radiolaria collected by
Dictyacantha tabulata, Dictyacantha tetragonopa. H.M.S. Challenger during the years 1873-1876.
• Dictyacantha tabulata (Haeckel, 1887) (Fig. Rep. Sci. Res. Voyage H.M.S. Challenger 1873-
6.125). Spicules of equal length and shape, thin, 76, 18:1-1803. Detailed monograph covering
long, 4-edged, gradually tapering distally. Each acantharian morphology and classification witb
with 20 myonemes. Basal ends with small leaf- superb illustrations. Many of the species erected
[1 cross. Spherical fine-meshed lattice shell with in this work were subsequently synonymized or
quadrangular pores, thin bars. Cell body spherical, renamed by Schewiakoff(1926).
opaque, dark red. Central capsule membrane Reshetnjak V.V. 1981. Akantarii. Fauna SSSR, 123,.1
thick. Spicule length: 0.58 rnm; diarneter oflattice Akad. Nauk SSSR, Zoo1. Inst., Nauka, Leningrad,
shell: 0.28-0.32 rnm, of leaf-cross: 0.016 rnm; cell pp. 1-210. Excel!ent review of al! aspects of acan-
body: 0.19 rnm. tharian studies, with detailed treatment ofthe tax-
• Dictyacantha tetragonopa (Haeckel, 1860) (Fig. onomy down to the species level (in Russian).
6.126). All spicules alike, short, massive, 4- SchewiakoffW. 1926. Die Acantharia des Golfes von
winged, sword-shaped, bases with large leaf- Neape1. Fauna Flora Golfo Napoli, Monogr. 37:1-
cross, each with 50 thread-like myonemes. 755. A cornerstone monograph based on
Coarse-meshed lattice shell with large pores, thick Mediterranean materials, covering al! aspects of
bars. Cell body spherical, opaque, yellow-brown. the knowledge ofthe group.
I
11
Acantharia 113
HOLACANTHIDA
ACANTHOCOLLlDAE
Acanthocolla cruciata
6.1b
'J. 6.2
6.4a 6.4b
Acanthocolla solidissima Acanthocolla sp.
Acanthocyrtha sp.
Acanthospira sp. 6.6b

114 Acantharia
HOLACANTHIDA
ACANTHOCOLlI DAE
j.
¡
isolated
spicules 6.8
Acanthospira spiralis
Acanthospira amphicopa 6.7b
HOLACANTHIDA
ACANTHOCH IASMI DAE
Acanthochiasma
hertwigi
6.10
isolated
spicule
6.12
6.11
6.9b
Acanthochiasma
Acanthochiasma fusiforme Acanthochiasma planum
quadrangulum
Acantharia 115
HOLACANTH IDA
ACANTHOCHIASMIDAE
r
/
6.14a
6.13a 6.13b
Acanthochiasma rubescens
HOLACANTHIDA
ACANTHOPLEGMI DAE
!
I
Acanthochiasma serrulatum
Acanthoplegma krohni
116 Acantharia
SYMPHYACANTHIDA
, AMPHILlTHIDAE
6.16c
6.16b
6.16d
Amphibelone anomala
í ';\ "\ '-..:, \
'\"" ;~:
':'
': ':,
..
:r
"Amphibelone Amphibelone
6.17 cultellata Amphibelone violina?
. violina
Acantharia
SYMPHYACANTH IDA
AMPHILlTHIDAE
6.19d
6.19c
6.19a
Amphibelone hydrotomica
, \
., '
:~
..
detail "
• í
i,
6.20a 6.20b 6.21a :' 6.21c
}
Amphilithium Amphilithium
c1avarium concretum
118 Acantharia
SYMPHYACANTHIDA
ASTROLlTHIDAE
I
.1
Acantholithium dicopum
Acantholithium stellatum Astrolithium piriforme

6.25
" Astrolonche sp.
11
1.
~I Astrolithium
Astrolonche
6.26a serrata
I
bulbiferum
,1
I
.;/; r
Heliolithiuthf" 6.27b
6.28a
aureum
Acantharia 119
SYMPHYACANTHIDA
PSEU DOllTH IDAE
6.29b
6.29a Dicranophora buetschli

6.30
Haliommatidium
tabulatum
6.31b
Haliommatidium muelleri
Pseudolithium compressum Pseudolithium bifidum

120 Acantharia
CHAUNACANTHIDA
STAURACONIDAE
6.35a
Amphiacon
denticulatus
6.34b
6.35b
6.36a Heteracon biformis
Stauracon pallidus
I
l' .11
,
1,
"
.. _ __
.. _-_._._._-~-----~---------
...
Acantharia 121
CHAUNACANTHIDA
CONACONIDAE
Conacon foliaceus I
r
!
I
l'
6.37a 6.37b
CHAUNACANTHIDA
GIGARTACONIDAE
Gigartacon sp,
6.38
Gigartacon fragilis
Gigartacon abcisus
6.40a
isolated
spicule
6.40b 6.40c
1\ Gigartacon muelleri
122 Acantharia
ARTHRACANTHIDA
ACANTHOMETRIDAE
6.42ai 6.42b 6.43a isolated

spicule
Acanthometra bu/bosa
Acanthometra
cy/indrica
6.43b
6.44
Acanthometra
dolichoscia Acanthometra pe//ucida,
isolated \.
, spicule
1" -v
Amphilonche Acanthoietra
6.47
diodon tetrecope 6.46a 6.46b
Acantharia !( 123
ARTHRACANTHIDA
ACANTHOMETRI DAE
, ..
i.:·
Amphilonche
6.48a f \ 6.48b \ elongata 6.48c
n.
---------
-:
6.49 Tetralonche arachnoides 6.50 Tetralonche rara

124 Acantharia
lL
ARTHRACANTHIDA
DORATASPIDAE
Coscinaspis peripora Craniaspis parmipora

Aconthaspis hastata
Dictyaspis furcata
Dictyaspis so/idissima
6.55c
Dorataspis micropora
Acantharia 125
ARTHRACANTHIDA
DORATASPIDAE
"' , 6.57b
Dorsisspis gladiata Oorataspis gladiata?
Dorataspis
\ macropora
\
Dorataspis lorícata
\
6.61b
.' 'f' . l~ detail
Hystrichaspis fruticata
[ J
126 Acantharia
ARTHRACANTHIDA
DORATASPIDAE
isolated polar
plate
6.62a equatorial section
through the
central capsule
6.62c
Icosaspis elegans
isolated plate '6.63
Jcosaspis ornata
Icosaspis serrulata
Acantharia 127
ARTHRACANTHIDA
DORATASPIDAE
6.65b
Lychnaspis giltschi 6.65c
t 6.66
Lychnaspis
longissima
6.69
Lychnaspis strelkovi Lychnaspis wagenschieberi

128 Acantharia
ARTHRACANTHIDA
DORATASPIDAE
6.70a
Lychnaspis polyancistra
Lychnaspis serrata
Lychnaspis undulata 6.72b

Acantharia 129
ARTHRACANTHIDA
DORATASPIDAE
"Pleuraspis bipennis
6.74
Pleuraspis constricta
Siphonaspis cristata
6.77b
~U'~r\v'~
~~~~
isolated
spicule 6.77c
Stauraspis stauracantha
130 Acantharia
ARTHRACANTHIDA
PHRACTOPELTIDAE
6.78
Phractopelta cruciata 6.79a
Phractopelta
dorataspis
6.79b
6.80a
Phractopelta tessaraspis
Acantharia 131
ARTHRACANTHIDA
DIPlOCONIDAE
Diploconus
cyathiscus
Diploconus fasces
6.82
Diploconus
cylindrus
6.85a 6.85b
J"
Diploconus satu¡~us
132 Acantharia
4-lobed central
capsule shown
6.86a 6.86b
Lithoot
________ r: era fenestrata
6.87a
4-lobed central
capsule shown
L Iithoptere muelleri
6.87b
Acantharia 133
ARTHRACANTHIDA
HEXALASPIDAE
6.88b
Coleaspis vaginata
Hexaconus ciliatus
.'
"
6.90b
Hexaconus ciliatus?
Hexaconus echinatus
134 Acantharia
ARTHRACANTHIDA
HEXALASPIDAE
6.92
Hexaconus
infundibulum
Hexaconus nivalis 6.93
Hexaconus serratus
juvenile \, 6.94d
\
I
'1
I
¡
L
Acantharia 135
ARTHRACANTHIDA
HEXAlASPIDAE
Hexaconus trypanon
Hexa/aspis sexa/ata ,..

"".... 1", "
;."-
¡ .~ Hexalaspis heliodiscus
136 Acantharia
ARTHRACANTHIDA
PHYLLOSTAURIDAE
Acanthostaurus
bipennis
I
6.98b
" Acanthostaurus sp.I
6.100b
Aeenthosteurus conacanthus
\
6.100a
¡
I
6.101 Acanthostaurus
I Acanthostaurus henseni purpurascens 6.102
1I
!
Acantharia 137
ARTHRACANTHIDA
PHYLLOSTAURIDAE
\,
¡
¡
I
6.103b 6.104
6.103a
Amphistaurus sp.
6.105
Amphistaurus atlanticus
Amphistaurus complanatus
¡
\ .
\ 6.106
Amphistaurus lanceolatus Lonchostaurus bifurcus

138 Acantharia
ARTHRACANTHIDA
PHYLLOSTAURIDAE
r
I
\
\ \ 6.108c
Lonchostaurus rhombicus
\ "
\~
isolated
\
spicule
\,
Phyllostaurus sp. 6.109
Phyllostaurus cuspidatus
Acantharia 139
ARTHRACANTHIDA
PHYLLOSTAU RI DAE
.f:\
g :
J : 6.111a
i
¡
1 ;
6.111d I Phyllostaurus echinoides
isolated
spicules
6.111b 6.111 e
isolated
spicule
6.112a
6.112b
Phyllostaurus quadrangulus
140 Acantharia
ARTHRACANTHIDA
PHYLLOSTAURIDAE
6.113a
Phyllostaurus siculus ¡
./ 6.114a
6.113b
r--
. quadrangular central 6.114b

capsule shown
Zygostaurus amphitectus
v
Zygostaurus
sagittalis
Acantharia 141
ARTHRACANTHIDA
STAURACANTHIDAE
\
\'.
......
6.116b
6.116d
Phatnacantha icosespis
Phatnacantha icosaspis ?
6.118
6.117b detail of
6.117c central
Pristscenthe part and
6.117a Pristecenthe octodon dodecodon leaf-cross
142 Acantharia
ARTHRACANTHIDA
STAURACANTHIDAE
6.119
Pristacantha polyodon
/ 6.120
Pristacantha multidentata
Xiphacantha multispina 6.121

b!:i
6.122a
6.122b
Stauracantha orthostaura
Acantharia 143
ARTHRACANTHIDA
STAURACANTHIDAE .' , \.
6.123b
Xiphacantha a/ata
Xiphacantha quadridentata 6.124

144 Acantharia
ARTHRACANTHIDA
DICTYACANTHIDAE
\ ITr
1\ 1#
6.125a
Dictyacantha tabulata 6.125b
Ii
6.126b
isolated
Dictyacantha tetragonopa spicule
Acantharia 145
Acantharian cysts
probably arder Halacanth ida
prabably arder Chaunacanthida

¡oa
146 Acantharia
Acantharian cysts
probably order Chaunacanthida
probably family Pseudolithidae
probably family Stauraconidae
6.150a 6.150b 6.151a 6.151b
11
Acantharia 147
Acantharian cysts
family unknown
6.152a 6.152b
6.155 6.156 6.157 6.158
Fig.6. Illustrations of representative acantharian species. Notice that while most ofSchewiakoff's (1926) figures depict whole individuals, those
taken from other sources, as well as the SEM photos, show the skeletal elements only. In many cases species names are different from those given in
the originals, following assignments and synonyrnies proposed by Schewiakoff( 1926) and Reshetnjak (1981). Specific identity of acantharian cysts
is largely unknown; howevcr, because these organisms can be rather cornmon in plankton samples, SEM photos of a representative se1ection of
forms is provided. Figure sources: From Haeckel (1887): 6.7a, 6.7b, 6.16b, 6.17, 6.18a, 6.19a, 6.19b, 6.26a, 6.30, 6.31a, 6,43a, 6,47, 6,48b, 6.51,
6.52,6.53,6.54, 6.55c, 6.56, 6.57b, 6.60b, 6.60c, 6.60d, 6.61a, 6.61b, 6.62a, 6.62b, 6.62d, 6.62e, 6.64, 6.65a, 6.65b, 6.65c, 6.66, 6.67a, 6.70a, 6.70b,
6.72a, 6.74, 6.75, 6.76a, 6.77b, 6.77c, 6.77d, 6.77e, 6.78, 6.79b, 6.80a, 6.81, 6.84, 6.86b, 6.86c, 6.88b, 6.89a, 6.90a, 6.91, 6.92, 6.93, 6.94a, 6.95,
6.96b, 6.97, 6.106, 6.107, 6.108a, 6.114b, 6.115a, 6.116c, 6.117b, 6.118, 6.119, 6.122a, 6.123b, 6.123c; 6.125a; courtesy ofA. Michaels: 6.3, 6,4a,
6,4b, 6.6a, 6.6b, 6.25, 6.29b, 6.34a, 6.37a, 6.38, 6.104, 6.109, 6.126c; from Popofsky (1904): 6.2, 6.24, 6.39, 6,49, 6.50, 6.88a, 6.101, 6.103a; from
Popofsky (1905): 6.103b; from Popofsky (1906a): 6.10, 6.11, 6.12, 6.98a, 6.98b, 6.111a, 6.111b, 6.111c, 6.111d, 6.112a; from Popofsky (1906b):
6.69,6.71; from Reshetnjak (1981): 6.1a, 6.9a, 6.13b, 6.16c, 6.19c, 6.20b, 6.21c, 6.22, 6.27b, 6.28a, 6,40b, 6,42b, 6,48c, 6.55b, 6.58a, 6.59, 6.67b,
6.68, 6.73a, 6.85a, 6.87b, 6.94b, 6.94c, 6.100b, 6.108e, 6.113a, 6.114a, 6.116d, 6.117c; from Schewiakoff(1926): 6.1b, 6.5, 6.8, 6.9b, 6.13a, 6.14a,
6.14b, 6.15, 6.16d, 6.19d, 6.20a, 6.2Ia, 6.21b, 6.23, 6.26b, 6.27a, 6.28b, 6.29a, 6.31b, 6.32, 6.33, 6.34b, 6.35a, 6.35b, 6.36a, 6.36b, 6.37b, 6,40a,
6,40c, 6.41, 6,42a, 6.43b, 6,44, 6,45, 6,46a, 6,46b, 6,48a, 6.55a, 6.57a, 6.58b, 6.60a, 6.62c, 6.63, 6.67c, 6.72b, 6.73b, 6.76c, 6.77a, 6.79a, 6.80b,
6.82, 6.83a, 6.85b, 6.86a, 6.87a, 6.89b, 6.89c, 6.90b, 6.94d, 6.96c, 6.100a, 6.102, 6.105, 6.108c, 6.110, 6.112b, 6.113b, 6.116a, 6. 116b, 6.117a,
6.120,6.121, 6.122b, 6.123a, 6.124, 6.125b, 6.126b; original: 6.1c, 6.l6a, 6.18b, 6,48d, 6.57c, 6.76b, 6.83b, 6.85c, 6.86d, 6.89d, 6.90c, 6.96a, 6.99,
6.108b, 6.108d, 6.115b, 6.116e, 6.116f, 6.123d, 6.126a, 6.127 through6.158.
,
\
I
I•
Radiolaria Polycystina
Demetrio Boltovskoy
Introduction both ends with other elements - the bars (Fig. 2F), or
formations attached to the rest of the shell by one end
Polycystine radiolarians are exclusively marine, pela- only - the spines (Fig. 2A, B, G). All skeletal elements
gic, solitary or colonial protists provided with actino- are composed of amorphous silica (Si02 nH20).
pods. Polycystines comprise the Collodaria, a small There is a perplexing variety of shapes in which these
group lacking a skeleton (Fig. 16.1, 6-8, 11, 12), or bars and spines can be arranged in order to form the
provided only with scattered mono- or polyaxonic skeleton, from simple latticed spheres (Fig. 16.13) or
spicules (Fig. 16.3-5, 16.9,16.10); and the Spumellaria a few anastomosed spines (Fig. 16.119), to elaborate
and Nassellaria, most of which have a well developed constructions with several concentric spheres (Fig.
siliceous latticed or spongy skeleton (see "Outline 2B) or multilocular conical structures with protruding
classification ..." below). Solitary species (the greatest latticed or solid appendages known as wings, feet,
majority) range between 20-30 um to about 300 um, teeth, etc. (Fig. 3).
but colonies (some Collodaria and the spumellarian
farnily Collosphaeridae) may in exceptional cases be Very little is known about the reproduction of the
as long as 3 m (Swanberg, 1979). The siliceous skele- Radiolaria. In addition to vegetative reproduction
tons of the polycystines are a major contributor to the (Hollande and Enjumet, 1953), including binary fis-
sedimentary flux, their earliest records dating back to sion at various ontogeneting stages (Anderson and
the Cambrian. Paleozoic, Mesozoic and Cenozoic Swanberg, 1981; Anderson and Gupta, 1998), the pro-
sequences furnish detailed records for evolutionary, duction ofbiflagellated swarmers was observed, but it
stratigraphic and paleoecologic analyses. is not known if the swarmers are asexual dissemules
or motile gametes (Anderson, 1983a). Although no
A distinguishing feature of all radiolarians (polycys- direct estimates have been made so far, it is generally
tines and phaeodarians) is the central capsule membra- assumed that individual radiolarian life spans are
ne, a proteinaceous perforated membrane that divides around 2-4 weeks (Anderson 1983a; Caron and
the cytoplasm into two areas: the endoplasm or intra- Swanberg, 1990).
capsular cytoplasm, and the calymma or extracapsular
cytoplasm (Fig. lA). This central capsule is either Polycystines consume a wide variety of prey includ-
spherical (in many Spumellaria), or elongated and ing bacteria, algae, protists, copepods, appendicular-
pyriform (in most Nassellaria, Fig. lB). The intracap- ians, and other small zooplankton (Anderson 1983a,
sular cytoplasm contains reserve substances and major 1993; Caron and Swanberg, 1990). AIgal symbionts,
cytoplasmic organelles (nucleus or nuclei, mitochon- when present, secrete photosynthetic products that are
dria, and other organelles, except for the digestive vac- assimilated by the host as a nutritional source
uo1es), and is generally believed to be responsible for (Anderson 1983b).
the functions of reproduction, biochemical synthesis
and energy production. The calyrnma is the frothy or The first published descriptions of Radiolaria date
web-like extracapsular cytoplasm where the digestive back to the early XIX century. Between approximate-
vacuoles are located. AIgal symbionts, when present, ly 1850 and 1900, C. G. Ehrenberg, J. Müller, R.
are enclosed within vacuo les usually located in the Hertwig, A. Popofsky, and especially E. Haeckel
calyrnma. Colonial forms have a gelatinous sheath described thousands of new species and provided the
containing numerous central capsules interconnected first comprehensive classification systems (Riedel,
by a rhizopodial network (Fig. 16.6-9, 16.11, 16.12). 1967a). After a period of little activity, interest in the
Radiolaria was renewed around 1950, and somewhat
Polycystine skeletons are typically constructed of a later further fostered by the rich sedimentary mate-
network of structures which can be either connected at rials recovered by the Deep Sea Drilling Project.

150 Radiolaria Polycystina
Spumellaria Nassellaria
apical
horn
cephalis
abdomen
Colonial Polycystina (Spumellaria)
Solenosphaera collina Acrosphaera spinosa Collosphaera huxleyi

(=5. chierchiae)
Fig. 1. Main features of a polycystine cell (A, B), and general view of entire radiolarian colonies (C-E). A: slightly modified from Kling (1978)
and Boltovskoy (1981e); B: from Hollande and Enjumet (1960); C, D: by courtesy ofNeil Swanberg (from Swanberg, 1979); E: from Strelkov and
Reshetnjak (1971).
Because of their application to stratigraphy, polycys- plus names; however, only 100-150 of these are pri-
tine studies have traditionally been within the realm of marily concemed with radiolarían studies. Almost all
geologists/paleontologists, with biologically-oriented these workers are geologists focusing their interest on
publications representing less than 10% of the overall stratigraphic and paleoceanographic problems, espe-
total produced to date (A. Sanfilippo, pers. cornm., cially dealing with Paleozoic and Mesozoic deposits;
1997). The directory included in the 1994 issue of interest in Cenozoic faunas has been dwindling over
Radiolaria (Newsletter for the Intemational the last few years. Biologically-oriented research
Association of Radiolarian Paleontologists) lists 400- based on samples from the water-column has even
Radiolaria Polycystina 151
fewer specialists, and at present they are probably less few other microzooplanktonic taxa, pose serious
than 10-20 world-wide. Since 1834 approximately methodological difficulties. Indeed, they are too small
3500 works on polycystine radiolarians have been (around 20-30 to 300 um) to collect effectively with
published (over half of these on Cenozoic faunas, standard zooplanktonic nets (100 to 300 um in pore
about 35% on Mesozoic, and 15% on Paleozoic; A. size), yet too scarce in most areas to yield adequate
Sanfilippo, pers. cornrn., 1997). catches with water-bottles or low-powered pumps.
Thus, fme-meshed nets have to be employed, which
significantly complicates not only the concentration
Methods of the radiolarians (due to the concomitant retrieval of
other organisms, some of which, like the diatoms,
Provenance and collection of materials cannot be fractioned out later; see Swanberg and Eide,
1992), but also because net clogging jeopardizes sub-
Most of the surveys on extant polycystine radiolarians sequent estimations of the volume of water filtered
published to date are based on samples of their skele- (Tranter and Smith, 1968; Boltovskoy, 1981b). In
tons preserved in the surface sediments, rather than on order to avoid clogging by smaller particles thus
plankton samples. Sediment samples have some ensuring better estimates of the volume of water fil-
advantages over water-column material s, but also se- tered and larger sample-sizes, meshes ranging
veral important shortcomings (see "Sedimentary vs. between 60 and up to 100 um are traditionally used
water-column materials" below). for polycystine studies in the water-column. It should
be stressed, however, that both absolute quantitative
A variety of sediment coring and grabbing devices estimates of radiolarian abundance, and the propor-
have been used throughout the years for analyses of tions of at least some species and developmental sta-
the polycystines from the upper centimeters of the ges may be seriously biased in these collections:
sediments (Kennett, 1982). Gravity and Kasten corers Boltovskoy et al. (1993a) reported that in sediment
are among the simplest, permitting retrieval of up to a trap materials from the tropical Atlantic shells below
few meters of sediment at a time from practically any 40-60 !lID represent roughly 50% of the overall poly-
depth. Piston corers have been used widely due to cystine fauna.
their ability to recover long sedimentary sequences, up
to 20-30 m in length. However, all these devices tend Estimates of radiolarian abundances in the water-co-
to disturb the sediments, especially the uppermost lumn must be performed with flow-metered nets;
layer which is of particular importance for the analy- clogging of the meshes, .in particular of those with
sis of Recent assemblages. Box corers (rectangular, small pores, makes assessment of the volume of water
shallow, ea. 1 m coring boxes which ensure complete filtered based on distance towed and mouth diameter
closure of water-flow passages after sampling and extremely unreliable (Tranter and Smith, 1968;
before leaving the seabed, thus minimizing sample Boltovskoy, 1981a, b). Thus, whenever unflow-
washout during ascent) are preferable for retrieval of metered nets are employed, such as those derived
the top layer of the sediments. However, the fact that from Tucker's (1951) opening-closing mechanism
box core samples usually lack the thin uppermost phy- (e.g., the Multinet, based on Bé's, 1962, design; the
todetrital film characteristic of most sediments (Billett MOCNESS, Wiebe et al., 1976; the RMT 1+8, Baker
et al., 1983) suggests that the bow-wave of the device et al., 1973), it is strongly recornrnended that evalua-
is strong enough as to wash away any mobile particles tion of radiolarian concentrations be avoided (species
before hitting the bottom. Multicorers, an arrangement proportions, on the other hand, are in principIe unaf-
of several short coring tubes mounted on a rigid frame fected in these samples).
seem to overcome this problem successfully as they
have been shown to collect phytodetritus, as well as For assessment ofthe delicate colonial forms, as well
significantly higher numbers of macrobenthic speci- as for studies of feeding, growth, metabolism, etc. of
mens than box corers (Bett et al., 1994). live individuals, specimens are collected by divers
(e.g., Swanberg, 1979), or by means of very short and
Plankton samples for radiolarian studies are usually slow plankton tows, thus ensuring a better preserva-
collected with nets. However, this group, as well as a tion of the protists (Matsuoka, 1992).
r
second
primary
-"--spines
f (straight,
~ 3-blaaed)
second
(first cortical) u ¡ I
shell pores with
~ __ outernmost hexagonal frames
(second cortical)
shell
Sty/atractus cubical
spp, group medullary
shell
T ·~·····'·'·
,
\
!:
TYPES OF SPINES
lENTELlI PTICAl
SHEllS
-~ ~.,~; .. "";--,)
3-bladed,
serrated
edges
3-bladed,
club-like
~
Spongotrochus
gleclelis
O
branched
3-bladed, cylindrical
twisted girdles
~ lateral
polar "y
caps
medullary
shell
beams
mantle
"-. ..
transverse
Didymocyrtis Cubotho/us spp. group
tetratha/amus
Phorticium elevei saggital
fl'
SPONGY SHEllS
I
11 (i)
spongy, with
spongy shell, concentric
no structure chambered rings
central
Sty/och/amydium chamber
asteriscus arms with
Sty/odictya
multispina
patagium
Dictyocoryne profunda
chambered ~
rings
Amphirhopa/um ypsi/on
W
Sediment trap techniques have undergone major the hydrodynamic properties of partic1e accumulation
improvements in the last years, thus constituting a very in the traps, these devices are most effective when
useful tool for the collection of polycystine materials deployed at depths in excess of 500-700 m (US
(US GOFS, 1989; Lange and Boltovskoy, 1995). GOFS, 1989; Lange and Boltovskoy, 1995). As a
Simple sediment traps consist of a concentrating cone result, they integrate the flux from several biological-
or funnel which tapers into a collecting jar; the array, ly dissimilar layers (e.g., Kling, 1979; Kling and
which can have either one or several traps, is moored to Boltovskoy, 1995). Furthermore, sinking skeletons
the botlom or drifts with the current suspended from a intercepted at these depths may not adequately reflect
buoy at the surface. Time-series models are deployed at their standing stocks at the surface, nor their specific
different oceanic locations for periods up to ayear or composition. Boltovskoy and Alder (1992) conc1uded
more, and are provided with a mechanism which replac- that, in the Weddell Sea, over 90% of the polycystines
es the collecting cup at predetermined intervals thus that inhabit the upper 400 m are destroyed (probably
yielding a detailed record of the changes in the amount due to fragmentation by grazing) before reaching 400-
and type of flux throughout several seasons (Honjo and 900 m of depth. Subsurface advection of shells pro-
Doherty, 1988; Lange and Boltovskoy, 1995). duced at higher latitudes and integration oflow protist
abundances over large depth intervals may be respon-
Sediment-trap materials have some important advan- sible for the fact that, in the eastern equatorial
tages over planktonic collections. Sample-size is usu- Atlantic, polycystine assemblage compositions
ally much larger in sediment traps than in plankton recorded in plankton samples at 0-300 m are totally
nets, with fluxes as high as 200,000 shells m-2 day-l different from those recovered in traps at 800-2000 m
having been recorded in the equatorial Atlantic (Boltovskoy et al., 1996; Fig. 4).
(Boltovskoy et al., 1996; see al so Table 3 in
Boltovskoy et al., 1993a). Seasonal plankton collec- It should be borne in mind that the yields of sediment
tions are composed of a sequence of snapshots which trap samples are not amenable to direct comparisons
represent but an insignificant proportion of the total with those of plankton samples: while the former are
time elapsed between tows, and may therefore not an expression of the downward flux, which in turn is
only under- or overestimate mean protist abundances associated with productivity and preservation, quanti-
(e.g., Bé et al., 1985), but also yield "atypical" speci- tative plankton samples give information on standing
fic assemblages. Time-series sediment trap samples, stock only. Hence, compositional differences may not
on the other hand, integrate over preselected depth only reflect advection, destruction by grazing, etc.,
and time ranges, thus averaging the overlying plank- but also biological traits of the species considered.
ton over restricted periods which yield adequate Tlius, a scarce species with high reproduction, mortal-
chronological resolution to allow pinpointing the re- ity and output rates may be rare in the plankton but
lative importance of limited offsets of the yearly abundant in the underlying sediment trap (Kling and
cyc1e. Furthermore, since seasonal variations in total Boltovskoy, 1995; Boltovskoy et al., 1996).
mas s flux are usually closely coupled with primary
production in the upper mixed layer (Honjo et al., As with other zooplanktonic groups, analyses of
1982, 1988; Deuser et al., 1983, 1990; Wefer, 1989), radiolarian vertical distribution patlerns are usually
comparison of total flux vs. radiolarian numbers and performed with the aid of vertically stratified plankton
specific makeup can furnish first hand information on tows (e.g., Renz, 1976; Kling, 1979; Dworetzky and
indicators (and paleoindicators) of the biological pro- Morley, 1987; Kling and Boltovskoy, 1995; Abelmann
ductivity of the associated water masses. and Gowing, 1997). However, because their identifi-
cation is based on the siliceous skeleton which pre-
Sediment trap material s, however, also have some serves after the death of the cell, in order to discrimi-
shortcomings. Because of limitations associated with nate live vs. dead protists in the subsurface layers the
Fig.2. Characters used for the identification of spumellarian radiolarians. A, B, D, E, G, H, K: frorn Haeckel (1887); C, F, J, L, M, P, Q: orig-
inal; 1: from Riedel (1958); R: from Nigrini and Moore (1979); N: from Dreyer (1889); O: from Petrushevskaya (1967); S: from Hollande and
Enjumet (1960).
MAIN SPINES OF THE INTERNAL SKELETON TYPES OF CEPHALlS

apical apical frontal ring cephalis
- se~~~~iry
(~~nh~~
~~o:--r
. í. ¡7
~:~:n
L~ve~~~al
saggital
nng
spine ~~cePhalismb~~a~n~~~
v:~~:1 collar 71~\ ~1tD~2nng

sagittal
partially
submerged
into thorax
V""
left) ~
dorsal
spine~ I
'.
. .
~
spme
main lateral
"spines
4D
7t
dorsal
spine.
pore~-=
main lateral
A¡-==- ~~
.
.
Ci
e .t. axial spine
(=axobate)
(right, left)
bar
sprne
: median axial spine
(=axobate)
'", /'
basal nng
Acanthodesmia viniculata
cephalis cephalis
Antarctissa denticulata
completely
group?
submerged
into thorax
(i) C1athrocorys teuscheri

oCarpocanium spp. group?
TYPES OF CEPHALlS
Acrobotrys spp. group

MULTILOCULAR CEPHALlS
SHELL STRUCTURES
PORE
ARRANGEMENT
G~
peristome
TRANSVERSE
ROWS
CHECKERED or
HEXAGONAL
LONGITUDINAL
ROWS
Lamprocyclas maritalis
V-''"- __ velum
Antarctissa denticulata (closed rnouth)
Lipmanella dictyoceras
group?
Vl 18 60-=:------------=------, 100
(])
e 15
'';:¡ 40
ECo
Vl
>- 10
SO O~
U Oc
>- S
20 M~
O
Q.
oa..
l-
O O O
.•..ro 18 60~------------~ 100
.•..O 15
'+- 40
O
(])
10
SO
eo
.•..ro S
20
e
(])
...
U
(])
O
(j) (j) (j) (j) (j)
O
o, re "'O
re re re re
"O "O
~~
':i :~ 'E =..c"O.!!!u
(j)
:O-ºE
o >- o
eo e, e
.~ O
~ U
e "¡::j U
O
c... -<u O
U
V'l
Species Families Orders
Fig. 4. Comparison of the mean percentage abundances for several nwnerically dominant polycystine species, families, and the 2 orders,
recovered in 0-300 m plankton tows with those from time-series sediment traps deployed at 853 m, in the eastern equatorial Atlantic. From
Boltovskoy et al. (1996a).
cytoplasm is often stained with rose Bengal, Sudan that stained protoplasm was present in individuals
black B, or eosin (Petrushevskaya, 1971b; Swanberg up to 4 weeks after actual death of the cell. These lap-
and Bjerklund, 1986; Abelmann and Gowing, 1997). ses are significantly longer than the time it takes a
Although this technique can furnish some clues on the radiolarian shell to reach the sea-floor (Takahashi and
living depth ranges of the species, it does not provide Honjo, 1983).
unequivocal information because of uncertainties
associated with the speed of decomposition of the Unless special cytological studies are required (e.g.,
protists' cytoplasm. Boltovskoy and Lena (1970), for Petrushevskaya, 1986), plankton and sediment trap
example, concluded that specimens of several plank- samples can be preserved in 4-5% formaldehyde; the
tome Foraminifera still contained protoplasm in their addition of picric acid to the solution enhances the
shell 98 days after death. Bernhard (1988) compared preservation of the colonies, yet acidification should
estimates of the proportions of presumably live ben- be avoided if the calcareous plankton is to be saved
thic Foraminifera as indicated by rose Bengal and from dissolution (see "Foraminifera").
Sudan black B staining and by ATP assay, concluding
Fig.3. Characters used for the identification ofnassellarian radiolarians. A, N: from Petrushevskaya (1981); B: from Riedel (1958); C, D, E,
G, 1, J, K, M, 0, Q, R, W: from Petrushevskaya (1971a); F: from Haeckel (1887); H, S, T, U, V: Original; L: from Riedel (1957); P: from Paverd
(1995); X: from Petrushevskaya (1967).
Sample preparation and analysis Moore (1973) proposed a convenient method which
allows quantification of the number of radiolarian
The following section offers some general comments shells per unit weight of sediment. Before processing
on the preparation of whole samples for routine count- as described above, the sample is dried and weighed.
ing and identification procedures. It does not review This weighed sediment is then c1eaned and sieved,
the methods involved in special cytological and ultra- and all the resulting residue is poured into a large
structural studies (se e Anderson, 1983a, for a review (e.g., 5 1) beaker full of distilled water, on the bottom
ofthese topics), as well as those used for detailed tax- of which one or two cover gasses have been posi-
onomic work, which can involve thin-sectioning, tioned. The water with the sediment in the beaker is
etching and polishing, etc. (Riedel and Sanfilippo, then thoroughly stirred (avoiding rotational motion,
1977; Boltovskoy et al., 1983; Petrushevskaya, 1986). which will result in centrifugal fractionation) for
achieving a random distribution of the partic1es, and
Pelagic surface sediments are usually c1ean enough as the sediment is allowed to settle. With the aid of a
to require little treatment before preparation of the siphon all but 3-5 cm of water are removed, and the
slides. Elimination of the organic matter and disaggre- remainder is evaporated with an overhead infrared
gation of the materials is achieved boiling the sample lamp.When the surface of the cover glasses is dry
(5-10 g) for a few minutes in a beaker with water to they are removed from the beaker and mounted as
which hydrogen peroxyde (10%, 300 mI per liter) and described above. The slide thus prepared will contain
tetrasodium pyrophosphate (10 g per liter) have been a fraction of the radiolarian shells present in the orig-
added. Disaggregation, c1eaning and removal of c1ay inal sample, this fraction being equivalent to the pro-
coatings and infilling partic1es can be aided by treating portion that the surface of the cover glass makes of
the sample in a gentle ultrasonic bath. For further dis- that of the surface of the bottom of the beaker.
aggregation of heavily indurated sediments various
products, such as kerosene, paint thinner, or ammonia Preparation of plankton and sediment trap samples is
can be helpful (the sediment is dried, soaked in the sol- somewhat more labororious due to the large amounts
vent, and then immersed in water, upon which disag- of organic material they contain. When both absolute
gregation usually occurs rapidly). If ca1careous materi- radiolarian concentrations and specific inventories are
al is abundant it can be removed with a few drops of sought, it is recommended that counting be performed
hydrochloric acid (after eliminating the hydrogen per- separately from the identifications. Polycystines can
oxyde by wet-sieving). The resulting c1ean material is be counted (although not identified) in whole, unpro-
then sieved with abundant water in order to eliminate cessed samples in counting chambers under the
the reagents and smaller partic1es. The mesh size used inverted microscope (Hasle, 1978; Boltovskoy,
depends on the aims of the study; most surveys rou- 1981c; Villafañe and Reid, 1995). Subsequently,
tinely employ 40-60 um-meshes, yet these, as either the entire sample or a subsample can be treated
described above, miss many of the smaller species, as in order to eliminate all organic matter leaving the
well as most developing forms. If precise abundance c1ean siliceous skeletons that will be mounted as
estimates are sought, mesh openings around 15 to 20 described above for sedimentary materials. It should
um should be employed, although these will retain be bom in mind, however, that radiolarian cells are
large numbers of unidentifiable skeletal fragments, as often very difficult to recognize in preserved, unpro-
well as non-radiolarian material (especially diatoms), cessed plankton samples. The siliceous skeleton, usu-
which can make subsequent observation more labori- ally the most conspicuous distinguishing feature, is
ous. The c1ean residue in the sieve is pipetted onto obscured by the cytoplasm to such an extent that
glass microscope slides, dried, and soaked with a few radiolarians are easily confused with other planktonic
drops of xylene; before the xylene has evaporated the protists, fecal pellets, eggs, various organic aggre-
mounting medium is added and covered with a cover gates, debris, etc. Adding a few drops of hydrogen
glass. Canada Balsam is most often used for these peroxide and/or hydrochloric acid, which slowly
preparations, although it takes longer to harden than digest the organic matter, and comparing the dubious
some other synthetic materials, commercially known partic1es before and after treatment can greatly help to
as Norland, Pleurax, Hyrax or Depex. I pinpoint radiolarian cells (Alder, pers. comm.).
Several different methods have been used for elimi- Scanning electron Light
nating organic material from water-column samples, microscope microscope
including high- and low-temperature ashing, oxidiz-
ing with hydrogen peroxide and/or ultraviolet light,
etc. (see review in Boltovskoy et al., 1983). One of
the most widespread, however, is that proposed by
Simonsen (1974) for cleaning diatom frustules. The
plankton sample is rinsed with abundant fresh water
(wet-sieving), and placed in a beaker to which an
equal volume of saturated KMn04 is added; it is then
left for 24 hs. A volume of concentrated Hel equiva-
lent to that already contained in the beaker is subse-
quently added to the sample; the dark brown liquid is
gently heated until it becomes transparent or light yel-
low. Once the sample has cooled, it is sieved again
thoroughly with fresh water and rinsed with distilled
water. The residue is pipetted onto microscope glass
slides as described above.
Analysis ofthe specimens is best performed in mount-

ed slides, which by transparency permits observing the
internal structures (such as medullary shells, spiral
structures, etc.), and the wall-thickness. In addition,
slight varíations in the depth offield allow one to deter-
mine whether a shell of circular outline is a disc (in
which case most of the surface is in focus simultane-
ously), or a sphere (either the central part or the peri-
phery are in focus). Photographs taken in the light
microscope have the advantage of being readily com-
parable to mounted specimens. The scanning electron
microscope (SEM), on the other hand, is especially
suitable for analyzing the surface morphology, but only
in specimens with large openings in the outermost
shell, or in those partially broken, can internal struc- Fig. 5. Scanning electron microscope (SEM) and light-microsco-
tures be observed. SEM photographs produce very py photographs of the same radiolarian specimens. A, A':
appealing results, but their comparison with routine Crornyechinus antarctica; B, B': Larcopyle butschlii; C, C':
Lithelius nautiloides. Notice that while SEM pictures yield great
collections mounted in slides is tricky (see Fig. 5).
details of the surface of the shell-wall, they conceal all internal
Ideally, both techniques should complement each other structures, most of which are important for identification purposes.
(Boltovskoy et al., 1983, described a method which From Boltovskoy (l98le), and Boltovskoy et al. (1983).
allows performing light and SEM observations and
photographs ofthe same radiolarían specimens; Fig. 5).
bias in sample-based particle counts (see reviews in
Assessment of radiolarian species-specific absolute Venrick, 1978a, 1995; Frontier, 1981), and in the
and relative abundances are based on identifications appraisal of species proportions (Patterson· and
and counts. Since any given slide often contains thou- Fishbein, 1989; Buzas, 1990). Patterson and Fishbein
sands of polycystine shells, the researcher is forced to (1989) concluded that for species representing >50%
decide how many specimens should be. identified and of the overall taxocoenosis at least 50 specimens
counted in order to achieve an adequate estimate of should be counted in order to achieve reliable percent-
overall numbers and species proportions. Several age data, 300 counts for species which comprise
methods have been proposed for the assessment of approximately 10% of a sample, 500-1000 counts for
•
species that make up 5%, and counts of several thou- radiolarian skeletons (see Fig. 7 below). Plankton
sands for those that comprise 1%. Unfortunately, in samples yield a snapshot-type image of the composi-
the case of the polycystines these efforts are unrealis- tion of the assemblages, which does not necessarily
tic because in any given sample containing 100-150 adequately reflect long-term trends. The dai1y, season-
species on1y 1-3 are above 10%, and 70-90 occur at al and interannual variability involved is smoothed out
levels below 1% (see "Geographic and vertical distri- in the sedimentary record, which may be a welcome
bution" below; Fig. 7). In terms of the amount of trait when general pattems are sought, Further, sedi-
information attained, it is more profitable to analyze mentary materials are more readily available from the
more samples at a lower resolution, than to examine various repositories around the globe than p1ankton
fewer sites at these statistically more reliable levels. samples. In any case, p1ankton samples not collected
Thus, in practice proportions are estimated in bulk, for microplanktonic purposes may be use les s for
regardless of the individual species abundances, usu- radiolarian studies due to inadequate net mesh-size ..
ally scanning 300-600 specimens per sample. It is
cornmon practice to identify the first 300-600 indivi- On the other hand, interpretation of the geographic
duals on the slide, and then check the rest of the slide distribution of extant radiolarian assemblages on the
or slides for the given sample in order to account for basis of sediment samples presents several important
the rarer taxa. The relative abundances of the latter are drawbacks (Boltovskoy, 1988, 1994, 1995; Kling and
estimated approximately, and they are usually exclud- Boltovskoy, 1995; Fig. 6). On their way to the sea-
ed from subsequent general numerical analyses (e.g., floor and after settling, radiolarian remains are grazed
multivariate techniques, such as cluster and factor upon by various consumers thus breaking their skele-
analysis) because of the uncertainties associated with tons into unidentifiable fragments. Because more de-
their assumed absences. It should be stressed, howev- licate shells are destroyed more readily than the more
er, that the counting effort necessary for reliable esti- robust ones, specific makeups on the bottom and at
mates of the fractional abundance of the rare species middepths can differ significantly from the living
is inversely proportional to the equitability of the assemblage in the upper water-column (Boltovskoy et
assemblage. Thus, when the sample is strongly domi- al., 1993b, 1996). Selective dissolution ofwhole sili-
nated by a single or only a few taxa, such as in polar ceous skeletons en route to the sea-floor and after dep-
areas (see Fig. 10 below), chances of recording the osition, although often advocated as an important
rare polycystines in random sequential counts are low source of plankton vs. sediments dissimilarities (e.g.,
because the observer repeatedly hits the dominant Petrushevskaya, 1971b; Renz, 1976), is probably
species. On the contrary, as equitability increases so much less critical than fragmentation due to grazing
does the probability of logging a so far unrecorded (Boltovskoy and Alder, 1992; Morley et al., Ms).
species with every new specimen scanned. Bottom material s can be reworked after deposition (as
a result of which non-Recent deposits, sometimes
characteristic of quite dissimilar oceanographic set-
Geographic and vertical distribution tings, are brought up to the surface layer, or win-
I nowed by bottom currents (dislodging settled ske1e-
\, Sedimentary vs. water-column materials tons and carrying them thousands ofkilometers away;
Fig. 6). Sediments integrate the imprint of near-sur-
As opposed to other zooplanktonic groups, studies on face faunas (which are generally associated with sur-
1: the geographic distribution of extant polycystines have ficial temperature, salinity and primary production
I
been chiefly based on sedimentary - rather than on fields, as well as with currents and water masses),
planktonic - materia1s. As mentioned above, sediment with the meso- and bathypelagic species whose geo-
samples present some advantages, but also several graphic distribution is uncoupled with upper-water
important shortcomings. oceanography (Fig. 6). In general terms the sedimen-
tary distributions of cold-water species tend to show
Whereas polycystine abundances seldom exceed 5 conspicuous equatorward extensions as compared
cells per liter in the plankton (e.g., Caron and with their planktonic pattems. This distortion is most
Swanberg, 1990), one gram of (dry) surface sediments probably due to the fact that extended survival of the
can contain thousands to hundreds of thousands of expatriated cold water taxa is facilitated by submer-
1
o ••
Reproduction
~!
00000
000
la
o
Fig. 6. Schematic diagram of the mechanisms that can distort the sedimentary imprint of the planktonic pattern of fossilizable microplankton
in general, and ofpolycystine radiolarians in particular (see text for detailed explanation). From Boltovskoy (1995).
sion (Boltovskoy, 1988, 1994; Fig. 6); as a conse- between presumably neritic vs. oceanic radiolarian
quence, sediment-derived species-specific ranges may assemblages have been described occasionally in the
wrongly suggest an enhanced tolerance to gradients in literature (Kruglikova, 1984), and even used for paleo-
the ecological factors. environmental reconstructions (e.g., Palmer, 1986).
However, with the probable exception of specific
Despite the limitations outlined, because radiolarian diversities, which indeed seem lower in neritic assem-
distributional studies based on water-column mate- blages (Nishimura et al., 1997), and the fact that a few
rials are extremely scarce, the biogeographic data selected polycystines are probably less intolerant to
surnmarized in Table 1 had to draw heavily on reports near-shore conditions than the bulk, most other traits
that used surface sediments. (such as Spumellaria:Nassellaria proportions, percent-
ages of Spongodiscidae, percentages of "spiny
Geographic patterns Porodiscidae", percentages of small "Cyrtoidea"; cf.
Kruglikova, 1984) need further confmnation.
Polycystines are typically open-ocean organisms,
occurring throughout the World Ocean. However, dis- Polycystine densities are typically around 0.3-1 cells
tinct coastal associations, while uncornmon or absent per liter, but values exceeding 50 ind. 1-1 have been
altogether in areas with an extended shelf, such as the recorded in some productive areas (Caron and
Southwestem Atlantic (Boltovskoy, 1980), have been Swanberg, 1990). The quantitative distribution of poly-
described in various studies. For example, Norwegian cystines in surface sediments of the South Atlantic is
fjords host dense and diverse radiolarian assemblages, illustrated in Fig. 7. This pattem is probably an
which differ from those of the open Norwegian Sea approximate representation of their concentrations in
(Swanberg and Bjerklund, 1986, 1987, 1992). the water-column as well, and it also roughly reflects
Interestingly, two of these fjord species, Rhizoplegma the overall distribution of primary production (e.g.,
boreale and Phormacantha hystrix/Plectacantha oikis- Koblentz-Mishke and Vedemikov, 1977), and ofphy-
kos, have been found to strongly dominate (up to 47% toplanktonic (Semina, 1977) and zooplanktonic
of all polycystines) shallow, coastal sediments around (Bogorov et al., 1968) biomasses. Highest numbers of
Antarctica (Nishimura et al., 1997). General differences polycystines would thus be expected along the upwell-
00
Shells per gram of
dry surface sediment
UJ <1,000
200 [ZJ 1,000-10,000
D 10,000-50,000
lB 50,000-100,000
400
• >100,000
600400 200 00 200 400
Fig.7. Quantitative radiolarian distribution in the surface sedimentary layer ofthe South Atlantic. Redrawn from Goll and Bjerklund (1974).
ing areas off Africa (Abelmann and Gowing, 1997), cal Atlantic (201,064 shells m-2 day+; cf. Boltovskoy
where the highest radiolarian fluxes have been recor- et al., 1996).
ded to day (Boltovskoy et al., 1996), and in the equa-
torial current system. In the southem part of the ocean The numbers of species that inhabit the different cli-
high densities are probably associated with the subant- matic zones of the World Ocean are difficult to esti-
arctic belt and its northem extensions, the Malvinas mate because most authors restrict their scopes to
(=Falkland) and the Benguela Currents. In a transect some 20-40 more or less well-defined morphotypes,
between the Antarctic and approximately 30oS, 100E ignoring the rest of the species. The few surveys that
(offNamibia), Abelmann and Gowing (1997) recorded (presumably) did attempt to identify all the skeletons
highest polycystine densities at 100-300 m in Antarctic recorded indicate that these numbers oscillate around
waters, and at 0-150 m in subantarctic waters (up to 0.3 100-200 for the tropics and subtropics, dropping to
ind. 1-1; these values, however, may be somewhat some 50-60 at the poles (Fig. 8). This decrease, how-
underestimating, see Fig. 13 and Boltovskoy and ever, is often punctuated by an isolated peak in the
Alder, 1992) . In the Southwestem Atlantic (30-60 S, 0
transitional areas which usually host both cold water
along 55°W), surface (5-15 m) layers were found to and warrn water taxa, especially in the sediments (see
host 0.5 polycystines per liter on the average, with Fig. 9) (Boltovskoy, 1981d, 1982, 1986).
maximum concentrations of 3 shells per liter (Alder et
al., 1997). Lowest numbers are those present in Central Despite these rather high numbers, very few of the
Gyre and Tropical/Subtropical waters (see Fig. 7). species are abundant in any given sample. In terrns of
their relative contribution to the overall polycystine
Flux rates of radiolarian shells at depths between 50 assemblage, usually only 1-3 species exceed.l0%, and
and ea. 5000 m vary from 0-4 to over 100,000 ind. m-2 up to 5 represent over 5%; radiolarians whose average
day+ (Boltovskoy et al., 1993a), with highest numbers percentage abundances are below 1% of the fauna
having so far been recorded in the north-eastern tropi- usually comprise 70-90% of all the species recorded
194m
162d
• I
J13d
• ~ I
Fig.8.
~--~~
~----_.,~--~-~~ 59n
..
75d
I
Numbers ofpolycystine species reported in various surveys which presumably attempted to identify all members ofthis group in their
samples (some figures are approximate). a: Benson, 1966 (surface sediments from the Gulf of California); b: Renz, 1976 (0-100 m plankton and
surface sediments, selected stations); e: Boltovskoy and Riedel, 1980 (0-1500 m plankton); d: Boltovskoy, 1987 (surface sediments); e:
Boltovskoy and Riedel, 1987 (0-100 m plankton); f: Swanberg and Eide, 1992 (0-400 m plankton); g: Boltovskoy el al., 1993a (sediment trap
samples from 853 m); h: Boltovskoy el al., 1996 (sediment trap samples from 2195 m); i: Boltovskoy el al., 1996a (0-300 m plankton); j: Kling
and Boltovskoy, 1996a (0-2000 m plankton, selected samples); k: Paverd, 1995 (5 m plankton); 1: Welling, 1990 (1000-1500 m sediment traps
and surface sediments); m: Takahashi, 1981 (400-5500 m sediment traps); n: Abelmann, 1992 (300-2500 m sediment traps); o: Bjarklund, 1973,
and Swanberg and Bjerklund, 1987 (0-1000 m fjord plankton samples and surface sediments); p: Abelmann and Gowing, 1997 (0-1000 m ver-
tically stratified plankton samples from 7 stations).
(Fig. 10). Of the 164 polycystines inc1uded in this Southwestern Atlantic records up to that date, which
review, around 10 can attain average proportions in basically represented 7 reports (Haeckel, 1887; Hays,
excess of 10% in any given area, 12-15 morphotypes 1965; Nigrini, 1967; Goll and Bjerklund, 1974;
can reach 5-7%, and ea. 50-70 are normally around 1- Lozano and Hays, 1976; Morley, 1977; and
3% (Table 1). The remaining half of the polycystine Boltovskoy and Riedel, 1980), chiefly based on sedi-
species are present at levels below 1%. Highest dom- mentary materials. This objective compilation pro-
inances are associated with polar environrnents, duced a spotty picture with no discernible patterns. In
where a single species or species group can account the 15 years elapsed since that review several contri-
for 25-40% of the assemblage (e.g., Antarctissa spp. butions based on South Atlantic materials appeared,
in the Antarctic, cf. Boltovskoy, 1987; Amphimelissa but they mostly focused on downcore analyses (e.g.,
setosa in the Greenland Sea, cf. Swanberg and Eide, Pisias and Moore, 1978; Coco, 1982; Weaver, 1983;
1992; Phormacantha hystrix/Plectacantha oikiskos Bjerklund and Jansen, 1984; Grinstead, 1984; Charles
and Rhizoplegma boreale in coastal Antarctic sedi- and Morley, 1988; Alperin, 1987), or were restricted
ments, cf. Nishirnura et al., 1997; see Fig. 10). geographically to rather small areas (Robson, 1983;
Dworetzky and Morley, 1987; Boltovskoy et al.,
Species-specific distributional data for the South 1993a, b, 1995, 1996a, b; Abelmann and Gowing,
Atlantic are scarce and fragmentary. Boltovskoy 1997). Thus, in order to fumish a more comprehen-
(1981e) produced a detailed listing of all known sive insight into polycystine biogeography in the
Geographic distribution Numbers of species Specific diversity

of samples recorded (Shannon-Wiener)
50 100 140 4 5
0° ---::;.----*------------ --------------_.
'_-...
__ 1;(~
ffr
20° 1;(1;(
_--------- -. 1;( ------
~
400
..
--------------
- _/~/
,"* ',.....
~
_----_...-----
,
.s->:
1;(
---------------1;(--------------------------.
1;(
60°
---- Majar fronts
1400 160° 180° 160°
Fig. 9. Fluctuations in polycystine numbers of species and specific diversity in surficiaI bottom sediments along a transect from the equator to
the Antarctic in the south Pacific Based on data from Boltovskoy (1987).
South Atlantic, distributional species-specific data are it's highest abundance. Total ranges are, as a rule,
referred to the 7 distinct areas illustrated in Fig. 11). much larger, and usually cover several climatic zones.
These divisions take into account the distribution of Data presented in Fig. 12 are based on surface sedi-
general planktonic biogeographic provinces (e.g., E. mentary material s, and therefore most probably offer a
Boltovskoy, 1970; Koblentz-Mishke and Vedemikov, somewhat blurred image of actual distributions in the
1977; Boltovskoy, 1979, 1981d, 1982, 1986; Dadon plankton. This is particularly noticeable in the case of
and Boltovskoy, 1982; Longhurst, 1995), as well as the cold water pattems, where typically subantarctic
radiolarian-based biogeographic pattems (Goll and and Antarctic species are recorded as far north as 30oS.
Bjerklund, 1974; Morley, 1977; see Fig. 11, insets A As described above (see "Sedimentary vs. water-co-
and B). For some of the especially abundant and bet- lumn materials"), submersion with cold currents and
ter defmed taxa relative (percentage) contributions to subsurface or deep water northward displacement of
all polycystines can be predicted with reasonable these shells is presumably responsible for these biases.
II accuracy. For most others, however, only a very
rough indication oftheir numbers (abundant, present) The information used to compile Table 1 was not
can be offered for the time being. restricted to data from the South Atlantic Ocean, but
was extracted from many reports on various oceanic
It should be stressed that specific distribution ranges areas, putting special emphasis on water column-
are hardly ever actually restricted to any ofthe biogeo- based surveys (see "Sedimentary vs. water-column
graphic domains indicated in Fig, 11. Rather, as illus- materials" above). Although very subtle differences
trated in Fig, 12, these areas define the "core" ranges between oceanic basins probably do exist (Nigrini,
of the species, i.e., the sector where the taxon reaches 1967; Goll and Bjerklund, 1974), polycystine species
Percentage of total polycystines in sample
1:~""m •••.••
u._......... Equatorial (128 .pp.) I
30~-----------------------------------------.
Transitional (78 spp.)
20
10
o
40-r--------------------------,
Antarctic (52 spp.)
30
20
10
O
Polycystine species
Fig. 10. Changes in the species richness and the equitability of radiolarian asseroblages, as exeroplified by single sarople yields from an
Equatorial area (from Boltovskoy, 1987, surface sediroents), a Transitional area (froro data in Boltovskoy and Riedel, 1987,0-100 m plankton),
and froro the Antarctic (from data in Boltovskoy, 1987, surface sediroents).
are chiefly restricted in their distribution by climatic South Equatorial Current, tropical assemblages differ
and productivity fields, rather than by ocean basins, as little from the subtropical ones. Central Gyre fauna is
are most other pelagic planktonic organisms. Thus, also very similar to the Tropical and Subtropical one,
with very few exceptions, similar assemblages char- yet these oligotrophic waters, characterized by very
acterize the equatorial circumglobal belt, the subtro- low overall plankton abundances, host enhanced pro-
pical zones of the two hemispheres, and the polar portions of several colonial radiolarians.
waters (Petrushevskaya, 1971a). Geographic endem-
ics are rare, probably accounting for les s than 5% of Vertical profiles
all the species (one outstanding example is
Antarctissa spp., Fig. 16.104, which is absent in the Vertical profiles of total radiolarian abundance in
Arctic, but dominates both the plankton and the sedi~ tropical and subtropical waters indicate that the bulk
ments of the Antarctic zone). oftheir populations is usually located in the upper 50-
100 m (Petrushevskaya, 1971b; Renz, 1976;
It should be bom in mind that the degree of mixture Dworetzky and Morley, 1987; Kling, 1979; Kling and
between most ofthe areas shown in Fig. 11 is extreme- Boltovskoy, 1995; Abelmann and Gowing, 1997; see
ly large. For example, in the westem South Atlantic Fig. 13A-C, F, G). Quite often several discrete maxi-
the Transition Zone stretches up to almost 15 degrees ma are recorded, one at or near the surface, and a sec-
in latitude (ea. 34-35°S to 47-48°S; see "General bio- ond one between 50 and 100 m (Petrushevskaya,
logical features of the South Atlantic"); Subantarctic 1971b; Kling and Boltovskoy, 1995). In the Antarctic,
species are regularly found here in the same tows as however, peak abundances seem to be associated with'
the Subtropical representatives (E. Boltovskoy, 1970, the Warm Deep Water and occur deeper, at 200-400
1981a, b; E. Boltovskoy et al., 1996). Because the m (Petrushevskaya, 1967; Boltovskoy and Alder,
Brazil current is a southwest flowing branch of the 1992; Abelmann and Gowing, 1997; Fig. 13D, E).
Tropical/
Subtropical
" I
',; .":. '.'.';: .:.' 1;
!,.. r
, , : l' ; I l' !;.;. I : [ ,
2'-~t"_'Central Gyre='~' g'§! '~?

~ 1 (SübtropiÓtl)-=-; i;
., I i . i i .
¡ r:
'11 ti t 1
300
1" ,
Polar
Transitiona
I
I
i,
,i' ' ; , Suba'ntarctiC'
-l II i
Antarctic
:1 Fig. 11. Main biogeographic areas characterized by different radiolarian specific assemblages. Inset maps show the distribution of selected
radiolarian species in surface sediments indicative of major biogeographic pattems (A, according to Goll and Bjerklund, 1974); and Morley's
(1977) biogeographic divisions of the South Atlantic based on the polycystine contents of 57 surface sediment samples (B).
Many radiolarian species occupy discrete depth inter- masses which move vertically as well as horizontally.
vals of the water colurnn. Kling and Boltovskoy For example, in the eastern subtropical Pacific inshore
(1995), on the basis of a series of plankton tows in the and oceanic 0-25 m waters can host a typically warm-
upper 2000 m in the eastern sub tropical Pacific water assemblage associated with the Central Water
defined the following characteristic layers: (1) surface which is advected coastward by the Southern
(with maxima at O m, 25 m, O and 50 m, 50 m, or O California Eddy, while midway between these two
and 100 m), (2) subsurface (maximum at 100 m), (3) sites the same depths are inhabited by a conspicuous-
deep (maxima at 200 m, 200 and 300 m, or 300 m), ly different, colder-water assemblage associated with
and (4) species peaking below 300 m. Roughly simi- the cooler waters of the California Current (Kling and
lar zonations were established by other authors as well Boltovskoy, 1995). Many cold water radiolarians that
(e.g., Renz, 1976; Dworetzky and Morley, 1987; inhabit the upper layers at high latitudes submerge
Kling, 1979). Worldwide depth zonations, however, with their corresponding water masses and can be
cannot be defined in terms of fixed depths beca use the found at depth in mid- and low-latitude areas (Kling,
distribution of radiolarian species is related to water 1976; Boltovskoy, 1988; Steineck and Casey, 1990).
Tropical/
Subtropical
o o
o Heliodiscus Acrosphaera Acrosphaera Collosphaera
asteriscus L- murrayana
~ spinos« ~roup? tuberosa
Central Gyre
Transitional
Subantareti e/ Antarcti e
Antarctissa
denticulata
group?
o
Antarctic
.0-.
1.20.:
~
. >50 .
Fig. 12. Examples of specific polycystine distribution pattems (in surface sediments; compare with Fig. 11). From Morley (1977).
Cold
Warm water
water
•..>-
QJ
/ij /ij
u
.';::: Depth
/ij-d /ij.
e c: •..~
u
- -
lo) u u
.¡: ._ "'C .5.. o
•.. o
,Se.. -;c..
o •.. -.;::: .'8•.. (m)
~..c:
:::let)
C'".- :::lC'"o==
1.1.I ..c: 1.1.I_
/ij
-•.. -
c:
QJ
U
•..
o
..Q
:::l
V'l
.¡¡;
e
..:~
c:
~
..Q
:::l
V'l -e
~
c:
Acanthodesmia viniculata P P 5 P P O
Acanthosphaera actinota P P
Acanthosphaera dodecestvle P P P P
Acanthosohaera oinchuda P P P
Acrobotrys sep. group
Acrosoheere murravana 5 P P P P
Acrosoheer« soinose l!roup? P 10 5 P
Actinomma antarcticum ~roup? P 5 5 0-200
Actinomma ercedoohorum P
Actinomma leptodermum P P P
Actinomma sol P P P P P 0-100
Amphirhopalum vosilon P A A A P
Amohisovris reticulata A A A P P
Antarctissa denticulata ~roup? 5 10 0-200
Anthocvttidium oohirense P P 25-50
Anthocvrtidium zeneuebeticum A P P P P
Arechnocorvs circumtexta P P 25-100
Arachnosphaera myriacantha P
Artostrobus annulatus P P P A >100
Astrosoheere hexeeonelis P
Botrvocepheline armata P P
Botrvocvrtis scutum P 5 5 5
Botrvoovle dictvoceohalus P P
Botryostrobus eauilonsris P 5 >100
Botryostrobus aurituslaustralis p p p P 5 5 P 100
Buccinosoheere inveglnete P A P P
Callimitra carolotae P
Carpocanium spp. group? p p 5 P P O
Carposphaera acanthoohora P P P A P
Cenospheere spp, l!roup 5 5 5 5 P
Centrobotrys thermophila P
Centrocubus c1adostylus P P
Ceohslosovris c1athrobursa P P
Cladococcus cervicornis P P
Cladococcus megeceros p p
Clathrocanium coarctatum P P P A O
Clethrocotvs teuscheri A P P p P
Clathrocyclas cassiopeiae P
Collosphaera huxlevi P A P P
Collosoheere mectooore P A P P
Collosphaera tuberosa P A P P
Cornutella profunda Cosmopolitan >100
Corocslvotre cervus P P P P 5 P 0-> 100
J
Cold
Warm water
water
>-
•..
~
"'¡:¡j "'¡:¡j
u
0';:
Depth
"'¡:¡j-ci
"'¡:¡j0
e o u r::: •..
u
-
o¡: 0_ "'C 0c.. o u
•.• o ~ 0';: (m)
- -
,Se.. o •.• "'¡:¡j 0';:
•.. •..
o •..~
u
-
r:::
~.c ~e.. r:::
O¡¡;
..s::;¡ r::: ~
..s::;¡
::::1 0.0 ::::1 ::
c"o_ ~ ~ r:::
C"'o ::::1 ::::1
1.1.I.c 1.1.I- U (J') ~ (J') -e
Corocelvotre columba P P A P
Corocalyptra kruegeri P
Cromvechinus antarctica 5 >100
Cromvechinus icosacanthus P
Cromyomma circumtextum P
Cubotholus spp. group
Cycladophora davisiana P 5 5
Cyrtopera laguneula Cosmopolitan? >100
Dictvocephelus papillosus Cosmopolitan?
Dictvocorvne p,lofunda 5 5 P P P O
Oietyocoryne truncatum P P P P
Dictvoohimus greciiioes P P A 5 100-1000
Dictvoohimus hirundo Cosrnopol itan?
Dictvoohimus infabricatus P
Didvmocvrtis tetrathalamus 10 10 10 5 A O
Diovliss« bensoni P?
Druooetrectus irreguletis P
Euchitonia ele1!ans/furcatagroup? P 5 10 5 P O
Eucvrtidium aeuminatum P P P A P O
Eucvrtidium anomalum A P P A P
Eueyrtidium hexastiehum P P P P P P 25-100
Heliaster hexegonium P
Heliodiscus asteriseus P P A P P O
Heliosoma echinaster P
Helotholus histricosa P P A A 0-100
Hexaeontium aristarehi P P
Hexacontium armatum/hostile zroup P P 5 P P
Hexaeontium leevigetum P
Lamprocyclas hannai P?
l.emorocvcles maritalis zrouni P P P A A 50-100
Lamprocyclas nigriniee P P P P 100->100
Lamoromitra coronata P
Lempromitre danaes P
Lamoromitra auedricusois P 100-> 100
Lampromitra schultzei P
Lerconvle butschlii P P P P A A P >100
Larcospira quedrengule P A P A P O
Liomenell« bombus P
LipmanelJa dictyoceras P P P P 5 O
Litharachnium tentorium P 100
Lithelius minor P P P 5 P P P
Lithelius nautiloides P P 5
Lithooere bacea P A P P O
Warm water Cold w,
•..>-
CIJ
~ ~
v
0';;
•..
v
~-é 0_"'t:I o
e v e
-"'"' -"'
-..:vv
~o
o¡: "'0
05.. o Depth
•..
- -
o •..~•.. 0';;
.se.. ~e..
"'.,c
•..o °Vi
e
e (m)
e ..c ..c
"'•..
:lt)() :l~
C"'o_ CIJ :l :l e
1.I.I.,c C"'o
1.1.I_ U CJl 1- CJl <1::
-~~---
Lithostrobus hexagonalis P 0-100
Lophophaena butschlii P
-
Lophophaena hispida P
Lophospyr~entagona pentagona P
Neosemantis distephanus P -- O
N~phrospyris renilla P
Octodendron cubocentron P
Octopyle stenozona group? 5 10 10 5 5 O
Peromelissa phalacra _. P
Perypiramis circumtexta Cosmopo itan >100
Phormacantha hystrix P P P --
Phormospyris stabilis scaphipes P P P A A 25-100
Phormospyris stabilis stabilis P
Phormostichoartus corbula P P P P P A A 100->100
--
Phorticium elevei P 5 --
Plagoniidae gro~ lO 5 5 A A A 5
Plegmosphaera entodictyon P
Plegmospha.eraexodictyon P -1-- ----
Plegmo?Il.haera pacl!y'p!~gma -
P --
Pseudocubus obeliscus P P P 5
-
Pterocanium ele~ P --
Pterocanium praetextum P P 5 5 -
P O
Pterocanium trilobum _--_
.. ..
P P P P 5 O
Pterocorys herwigii P P P A P O
Pterocorys minythorax P P P P 10 O
Pterocorys zancleus P P P A P --
Pterocyrtidium dogieli --
P?
Pteroscenium pinnatum P
Pylolena armata group? P P P A P
Pylospyra octop~ P?
Saccospyris antarctica P 5
Saturnalis circularis P P P
Sethoconus anthocyrtis P-
Sethodiscus macococcus P P
Sethophormis aurelia P O
Sethophormis rotula P P P
Siphocampe arachnea P P A >100
Siphocampe lineata P P 5
Siphonosphaera martensi A A P P --
Siphonosphaera polysiphonia P A 5 P P O
_Solenosphaera chierchiae A A P P
Solenosphaera polysolenia A A P P --
Cold
Warm water
water
>-
...
~
-; -;
1.1
o,¡::
Depth
-;-é -;0
e 1.1 c: ...~
1.1
- -...
o_"t:j \J 0Q. o
1.1
o¡: o,¡:: (m)
-
•.• o
.se.. o •.• -; ... ...
o,¡::
-o 1.1
o¡¡; e
~.s::. ¡:¡¡e.. c: c: ~ ~
::1 0.0 ::1 ~ ~
..&:l
•..
~ ..&:l
e
C"'o- C"'o ::1 ::1
w.s::. w_ U <Jl t- <Jl -e
Spirocyrtis sca/aris P --
Sooneester tetras irrezuleris P P P A
Spongaster te tras te tras -
5 5 P P P O ---
Spongocore cy/indrica --
P ---
P P P P 100
Spongodictyon spongiosum P P P P P
Spongodiscf!§ resurgens _ 5 P P P 10 P 5
Spongo/iva ellipsoides ~~.-.~-- P
5J:2..ongop/egma antarcticum P
Soongoplegme rugoss P
Spongopyle se tosa P P P P P 5 5 >100
Spongosphaera streptaca[Jtha P P P 5 P lO? O
Spongotrochus glacialis 5 P 5 10 -- P P 5 0-100
Spongurus py/omaticus ------
P P --
P A 0-1000
~.!]gprus sp~group? --- 5
---
P P P P A 10 >100 ---
Stichopi/~um bicorne -•..... -.-- P •...
Stylatractf!~~roup --- P 5 5 P
Stylochlamrdium asteriscus P
---- P P 5 P --
Stylodictya aculeata p P 5 5
---
Stvlodictve multispine 5 5 5 10 10
Sty/osphaera me/pomene -- P
Styptosphaera spumacea P
Tetrap/ecta pinigera -- P
Thecosphaera inermis - -
P P P A A
Jheoco.!J1hium trachelium -
P P P P 5 P 0-100
Theopilium tricostatum P
--- 1-------- P P 5 P P P O
Th0/osf!Y..@~~!yJcornis -
Tholospvris anthophora P P P
Tholospyris ramosa -----------
P
Th%spyis tripodiscus P
Tribonosphaera centripetajjs P A P P
Triceraspyris antarctica P 5 10 ---
Tricolocempe cy/indrica --
P P P P
Trisu/cus triacanthus P P P P P
Zvuocircus oroductus P P 5 P P
Table 1. General distributional data for the polycystine species treated. Geographic distribution areas refer to those indicated in Fig. 11. Most
abundant species are denoted with bold lettering. Numbers in body of table indicate approximate pecentage contribution to the correponding
assemblages; 10: between 5 and slightly over 10%, exceptionally up to 20-30%; 5: up to 5%, usually around 1-3%; A: abundant; P: present. Depths
defined are those ofpeak abundance ofthe species as derived from various reports, mostly based on materials from the Pacific Ocean. These vertical
ranges are probably generally valid for middle and low latitudes, but not for the Subantarctic andAntarctic areas.
Percentage of Iive
Polycystines per liter
polycystines
o 0.2 0.4
O
O 0.5 1.5 2
_ •••• -"'------1
O 50 100
o O
~
500 500 500

I
-
S
..c
~
S 1000 ------------------------
1000 fr1000
..c B O
.¡...J
Q..
(J.)
O
1500 ------------------------ 1500
2000 -'----------'
H
Polycystines
per liter
O 5 O 5
O O
..c
.¡...J
Q..
(J.) 100 .... 100
~ O
-E
..c
.¡...J
150
Q.. 100
(J.)
O 200
200 200 200
300
400 .
Fig. 13. Vertical distribution patterns ofpolycystine radiolarians in various oceanic areas. In all cases total numbers ofshells recorded are illus-
trated, which most probably significantly overestimates in situ living populations, especially below 100-200 m; panel H shows assumed propor-
tions oflive polycystine cells at various depths in the water colurnn. A, B, H: from data in Kling and Boltovskoy (1995); e, D, E: from data in
Renz (1976); F, G: from data in Boltovskoy and Alder (1992).
Siphocampe arachnea (Fig. 16.167), for example, is a Changes in the proportions of presumably living poly-
dominant component of surface Pacific Arctic and cystine cells with depth have been assessed in a few
Subarctic plankton; in the central north Pacific it studies. Boltovskoy et al. (l993a), based on extensive
peaks at 100-300 m, and at 300-1000 m in the sub- sediment trap data, concluded that numbers of live
tropical eastem Pacific (Boltovskoy, 1994). specimens decrease drastically downwards (e.g.,
·.ooo·_o .0.0 ••• 00 •••• 0_0. ..0_0 •• _ •• _
aprox. 100% at O m, 50-60% at 100 m, 20-40% at 200 In spite of these shortcomings and the time elapsed,
m, 10-20% at 500 m, 5% at 1000 m; see Fig. 13H). advances in the development of a better classification
These results generally agreee with other studies (e.g., system have been very limited. Efforts to depart from
Petrushevskaya, 1971a; Kling and Boltovskoy, 1995). and improve upon the classification schemes inherit-
On the other hand, Abelmann and Gowing (1997) esti- ed from earlier workers have mainly followed two dif-
mated much higher proportions of living cells at com- ferent approaches: cytological techniques and evolu-
parable levels in the water column: over 90% at 100- tionary studies.
200 m, around 70% at 300-500 m. It should be noticed
that staining techniques, which are usually applied for Hollande and Enjumet (1960), Cachon and Cachon
these estimates, do not adequately differentiate (1972), Petrushevskaya el al. (1976), Petrushevskaya
between live and dead cells (see above "Provenance (1981) proposed revisions which use not only the
and collection of materials"), for which reason it is skeleton (as most other classifications), but also cyto-
probable that concentrations of living specimens plasmic features, in particular the "nucleoaxopodial
below 50-100 m are systematically overestimated in complex" (sensu Petrushevskaya, 1981). Although
such surveys (Boltovskoy el al., 1993a; see Fig. 13H). these schemes are probably sounder in biological
terms, their applicability to fossil and subfossil mate-
As with geographic pattems, data on the depths at rials lacking the protoplasm is problematic, which is
which the various species peak listed in Table 1 have one of the reasons for their very limited acceptance
been compiled from reports on different oceanic among radiolarian workers.
areas. It is anticipated that they are generally valid for
subtropical and tropical environments worldwide; at Analyses of evolutionary lineages in geological
higher latitudes, however, some deep species may sequences were somewhat more succesful than cyto-
occur closer to the surface, while in the Antarctic the logical techniques in defining characters applicable to
bulk of the asssemblages seems to occupy deeper classification. Based on evolutionary evidence, Riedel
layers (see above). and Sanfilippo (1986) produced an interesting critical
review of the most important skeletal traits used by
Haeckel. They concluded that some of them (e.g.,
Taxonomy number of segments, number of supplementary con-
centric spheres, number of feet, number of rays and of
Morphology and c1assification systems equatorial spines in discoidal Spumellaria, presence
and nature ofthoracic wings) have little or no suprag-
Based on his previous monograph of 1862, and espe- eneric value. In contrast, several others (especially
cially on the extensive collections of HMS Challenger, cephalic structure, but also pore arrangement, shell
Haeckel (1887) produced the first comprehensive terminations in Nassellaria, etc.), traditionally consi-
system of radiolarian classification, encompassing over dered as of minor value, are conservative through
3,000 species, 2,400 of which were new to science. time, reveal evolutionary lineages and, therefore, are
Although Haeckel's work is still a compulsory refer- relevant for higher-rank divisions.
ence guide for anyone attempting to deal with the iden-
tification of these organisms, it has for some time been Riedel (1967b, 1971), Petrushevskaya (1965, 1971a),
evident that it does not satisfactorily represent natural Goll (1968, 1969), Sanfilippo and Riedel (1970),
relationships. Indeed Haeckel's groupings are only Zhamoida and Kozlova (1971), Foreman (1973),
based on morphological similarities without the sup- Dumitrica (1988, 1989) based on skeletal features
port of continuity in the fossil record, rather than on alone worked out altemative classifications, either for
demonstrable evolutionary sequences. In addition, the the entire order or for selected polycystine groups. Of
rigidity of his diagnoses, based chiefiy on strict geo- these, Riedel' s (1967b, 1971) suprageneric system
metric considerations (Fig. 14), ignores the ample intra- has become the most widely accepted for extant and
specific variability of the polycystines. As a result, Cenozoic radiolarians, and is the one adopted here-
many of his described "species" are but slightly differ- with (with slight modifications; see also Kling, 1978;
ent morphotypes or even developmental stages of the Boltovskoy, 1981 e; Anderson el al., 1996). It should
same organism (se e Figs. 13 and 14). be stressed, however, that this system does not over-
Spumellaria
ealiad a r ia (skeleton absent or not latticed)
Sphaerellaria (latticed or spongy skeleton present)
Prunoidea (shell ellipsoidal or cylindrical with constrictions)
Discoidea (shell biconvex or flat disc)

Larcoidea (shell triaxon-ellipsoid)
Sphaeroidea (one to several concentric spheres)
outermost
sphere
spongy,
with or w/o
.•••
1-------- outermost sphere latticced --------,,--l.~ medullary
1 sphere 2 spheres 3 spheres 4 spheres 5 spheres shells
l.iosohaerida
No radial spines
Stylosphaerida
Two main radial spines
Staurosphaerida
Four main radial spines
Cubosphaerida
Six main radial spines
'"
""C
E
E
o
°00
c:
~
Vl
Fig. 14. Schematic representation ofthe characteres used by Haeckel (1887) for the classification ofthe Sphaeroidea (=Actinommidae).
come many of the above-mentioned problems, and is there is fairly good agreement, binornial nomenc1ature
therefore a compromise provisional c1assification. alone very often fails to pinpoint unequivocally a given
Several of the family-level definitions, especially in morphotype because different names are applied to the
the Spumellaria, are rather vague and generally used same species and, conversely, identical organisms are
as a lumping black box for the many forms with com- reported under different specific and even generic
plex morphologies and poorly understood relation- names (see Boltovskoy and Jankilevich, 1985).
ships (e.g., Litheliidae, Pyloniidae, Tholoniidae). Because a very substantial proportion of the original
species descriptions were published in old and often
Specific identification ofthe polycystines is a time-con- hard to get monographs, some authors find it faster and
suming and frustrating task. With the exception of the easier to create a "new species" for the unusual-looking
few abundant and widespread species on whose names skeleton in the slide, than to comb the dusty books in
Actinornrnidae
Pyloniidae Phorticium elevei
100 um
Plagoniidae Arachnocorallium spp.
Fig. 15. Successive stages of growth ofpolycystine skeletons. From Petrushevskaya (1962, 1967).
search of an adequate, already established name. The following illustrated glossary of most comrnonly
Ecologically, paleoecologically and stratigraphically- used terms for the description of polycystine skeletons
oriented studies often underestimate the importance of is chiefly based on the listing compiled by
a stable and consistent naming system; the lack of spe- Petrushevskaya (1981). Capital letters (in parenthe-
cies illustrations in these reports allows the wrong desi- ses) denote the group for which the term is used (N:
gnations to go undetected. This not only hinders build- Nassellaria; S: Spumellaria).
up of useful information, but al so significantly
degrades the overall quality of radiolarian-based data Abdomen (N) (Fig. 30, P): the third segment of nas-
for other applications. Recent literature has abundant sellarian multisegmented shells.
examples of this bias, which introduces even more Aboral, aboral pole (N): skeletal section located at the
chaos into the already anarchic situation inherited from opposite extreme of the mouth or aperture.
turn of the century works. Indeed, this may be a major Nassellarian growth starts with this part of the
reason for the waning use of radiolarians in stratigraph- skeleton.
ic and paleoecologic work. Adoral teeth: see teeth.
Annular strictures: see strictures. Axoneme (N, S): the central shaft ofparallel microtu-
Antecephalic chamber: see antecephalic lobe. bules of the axopodia (cytoplasmic structure).
Antecephalic lobe (N) (Fig. 3M, N): Section of the Axoplast (N, S) (Fig. lA): central nucleus of the axo-
cephalis at the base of which the dorsal spine is podial system (cytoplasmic structure).
located. Separated from the eucephalic lobe by the Bars (N, S) (Fig. 2F): siliceous anastomosed beams
apical spine. When this section is separated from that define the meshwork of polycystine shells,
the rest of the shell by a pored wall it is called the separated by pores.
antecephalic chamber. Basal feet: see feet.
Aperture: see mouth. Basal plate: see collar plate.
Apical cupola: see galea. Basal pores (N): see collar pores.
Apical horn (N) (Fig. 3P): external extension of the Basal ring (N) (Fig. 3C): the subcircular structure
apical spine. formed by the arches that separate the wall of the
Apical pore (N): wall-pore located at the base of the cephalis from that of the thorax. In reduced skele-
apical spine. tons, such as those of the Spyridae, these arches
Apical spine (N) (Fig. 3A, B): Internal cephalic spine make an actual ring, whereas in other families the
which branches off the median bar close to the ring is embedded into the shell-wall.
point of insertion of the dorsal spine. The apical Basal segment (N): the last segment in multisegmen-
spine can protrude outside of the cephalis, in ted N assellaria.
which case its external section is called apical Beams (S): siliceous rods joining contiguous shell
horno structures. In some families (e.g., Actinommidae,
Apical tube (N) (Fig. 3J): Tube-like projection on the Pyloniidae, Litheliidae) radial beams connect the
cephalis of the Artostrobiidae, homologous to the successive, concentric spheres (Fig. 2A) or con-
vertical spine. Also termed lateral tube or tubule. secutive whorls of the spiral (Fig. 2L).
Apophyses (N, S): any external or internal protruding Branched spines: see spines.
outgrowth of the skeletal meshwork. By-spines: see spines.
Appendages (N): any external or internal protruding Calymma (N, S) (Fig. lA): extracapsular cytoplasm
outgrowth of the skeletal meshwork. Riedel and (see central capsule).
Sanfilippo (1986) suggested that primary appen- Central capsule (N, S) (Fig. lA, B): a dense central mass
dages (those directly connected to or having ofcytoplasm (the endoplasm or intracapsular cytoto-
homologies with the internal spicule) are of high- plasm) with the nuclear material, as well as reserve
er phylogenetic significance than the secondary substances and major cytoplasmic organelles,
ones (not directly related to the internal spicule). bounded by an organic, perforated membrane which
Arches (N): anastomosed skeletal outgrowths of the separates it from the extracapsular protoplasm.
main spines, such as the one forming the upper Central chamber (S) (Fig. 2R): the central, spherical
section of the sagittal ring (Fig. 3A). structure of the skeleton of several Spongodiscidae.
Areolate (N, S): Referring to small, very regularly Cephalis (N): the first (uppermost) segment of the
repeated wall perforations or pores (see Radiolaria skeleton, which can be either undivided (Fig. 3H-
Phaeodaria, Fig. 1G). J), or divided into lobes or chambers (Fig. 3K-N).
Arms (S) (Fig. 2Q, R): elongate projections (usually The cephalis can be well differentiated from the
3) radiating from a central subcircular disc in the thorax (Fig. 3F, H-R), or it can be immersed into
shells of some Spongodiscidae. Arms can consist it partially (Fig. 3D) or totally (Fig. 3E).
of unstructured spongy meshwork (Fig. 2Q), or of Cervical apophyses (N): lateral outgrowths ofthe dor-
spongy meshwork with more or less clearly visible sal spine.
chambered rings (Fig. 2R). Cervical pores (N): paired pores in the base of the
Axial rod: see axial spine. cephalis between the primary lateral and the dor-
Axial spine (N) (Fig. 3A, B): spine projecting from sal spines.
the median bar and oriented toward the thorax and Cervical stricture (N) (Fig. 30): furrow or constric-
subsequent segments; can be simple or branched. tion which divides the cephalis from the thorax.
Also called axobate. Chamber (N): one of the several sections into which
Axobate: see axial spine. the cephalis of the N assellaria can be divided (see
antecephalic, lateral, eucephalic and postcefalic Funnel-shaped pores (N, S): pores the external open-
chambers). See also central chamber. ing ofwhich is conspicuously wider than the inter-
Chambered rings (S) (Fig. 20, R): Concentric or spiral nalone.
more or less visibly segmented rings around the cen- Galea (N): helmet-shaped portion of the shell of the
tral chamber in several Spongodiscidae (see arms). Nassellaria associated with the apical horn; its
Club-like spines: see spines. base is formed by the top of the cephalis. Also
Collar plate (N): the complex structure at the base of called apical cupola.
the cephalis formed by the spines and arches, all Gates (S) (Fig. 2L, M): large openings in the
lying on approximately the same planeo skeleton's meshwork, usually conspicuously larger
Collar pores (N) (Fig. 3C): pores in the collar plate. than the pores.
Collar stricture: see cervical stricture. Girdles (S) (Fig. 2L, M): circular or ellipsoidal skeletal
Columella (N): free portion of the apical spine (Fig. perforated plates arranged in 3 mutually perpendicu-
3A) in the cephalis of the Nassellaria. lar planes which form the skeleton of the Pyloniidae.
Cortical shell (S) (Fig. 2A, B): one of the outermost Internal spicule: see main spicule.
perforated spherical shells of the Spumellaria, Intracapsular cytoplasm: see central capsule.
located outside of the central capsule. Jugal pores (N): paired pores in the base of the cephalis
Cupolae (S) (Fig. 2C): dome-like protuberances ofthe between the primary lateral and the vertical spines.
skeleton of the Tholoniidae. Laterallobes (N) (Fig. 3K, L): lateral sections of the
Cylindrical spines: see spines. cephalis separated from the eucephalic portion by
Dorsal spine (N) (Fig. 3A, B): Internal cephalic spine cephalic arches.
projecting from the median bar at the opposite end Lateral spines (N) (Fig. 3A, B): Paired spines project-
of the vertical spine, next to the apical and secon- ing from the median bar. The main or primary lat-
dary lateral spines. The dorsal spine can protrude eral spines (right and left) are located in the vicinity
outside of the cephalis at the level of the cervical of the vertical spine, while the secondary ones
stricture. (right and left) are inserted close to the apical spine.
Equatorial plane (S): mainly applicable to lenticular, Lateral tube: see apical tube.
discoidal and biconvex spumellarian shells (Fig. Latticed meshwork (N, S) (Fig. 2D, E, J, L): siliceous
2H, 1); plane of maximum shell surface. meshwork ofbars separated by regular or irregular
Eucephalic lobe (N) (Fig. 3N): in species with a divid- pores of variable size, not spongy in appearance
ed cephalis, the section which hosts the median bar. (see spongy meshwork).
Extracapsular cytoplasm: see calymma. Lentelliptical shell (S) (Fig. 2H, 1): in the shape of a
Feet (N) (Fig. 3Q): External projections ofthe wall of biconvex disco
the thorax oriented down and sideways, usually in Lumbar stricture (N) (Fig. 30): constriction between
the number of3. The feet are often associated with the thorax and the abdomen.
the dorsal and primary lateral spines. Main lateral spines: see lateral spines.
First shell (S) (Fig. 2A, B): the innermost shell of the Main spicule: see main spines.
set of concentric spheres of an actinommid. Main spines (N) (Fig. 3A-C): basic skeletal elements
Sometimes also applied to members of other fam- of most Nassellaria, composed of the median bar
ilies with several successively larger shells totally from which the apical, vertical dorsal and lateral
or partially enclosing one another. Shells are num- spmes anse.
bered from the inside of the skeleton toward the Mantle (N, S): usually thin, delicate, lace-like mesh-
periphery; thus, the second shell is the one located work surrounding the main shell which appears in
next to- and outside of the first one, and so on. some fully grown polycystines.
Fourth shell: see first shell. Median bar (N) (Fig. 3A-C): the basic nassellarian
Frames (N, S) (Fig. 2D): ridges on the surface of the internal skeletal element which supports the api-
shell surrounding the pores. Frames around pores cal, vertical, dorsal and lateral spines. Its position
can be roundish or polygonal (pentagonal, hexa- defines the limit between cephalis and thorax.
gonal, etc.). Medullary shell (S) (Fig. 2A, B, J): one of the inner-
Fundamental spicule: see main spicule. most perforated spherical shells of the Spumellaria,
located inside the central capsule. The central Post-abdominal (N) (Fig. 30): refers to structures
smallest sphere «30 mm) is also called microsphere associated with the fourth and subsequent seg-
by some authors. ments of multisegmented Nassellaria (segments,
Meshwork (N, S): the combination of anastomosed pores, ribs, etc.).
rods and bars that form the external siliceous Postcephalic lobe (N) (Fig. 3N): one of the sections
skeleton of the polycystines. into which the cephalis can be divided, sometimes
Microsphere: see medullary shell. elongated into a tube.
Mitral ring (N): in the Spyridae, the skeletal ring Primary lateral spines: see lateral spines.
which lies in the plane parallel to that of the basal Primary spines: see spines.
ring and perpendicular to that of the sagittal ring. Pylome (S) (Fig. 2N): A larger opening on the exter-
Monolocular shell (N) (Fig. 16.111): Nassellarian nal shell of some Spumellaria, often extending
shell composed of a single segment. into a tube or surrounded by larger spines.
Monothalamous shell: see monolocular shell. Radial beams: see beams.
Mouth (N): A large basal opening in the last segment Ribs (N) (Fig. 3Q): elongated, usually unperforated
ofthe Nassellaria (also called aperture), which can thickenings in the shell-wall. Sometimes the ribs
be open (Fig. 30) or obliterated by a porous plate can extend into wings (Fig. 3R) or feet (Fig. 3Q).
or velum (Fig. 3X). Dorsal and primary lateral spines embedded in the
Multilocular cephalis (N) (Fig. 3K-N): cephalis pro- wall of the thorax can also form ribs.
vided with several chambers or lobes in addition Rosette-shaped (N, S): usually applicable to pores
to the eucephalic one. with a tri- or tetra-lobed outline (Fig. 2E).
Multilocular shell (N): see multisegmented shell. Sagittal plane (N): the plane defined by the position of
Multisegmented shell (N) (Fig. 30): Nassellarian shell the median bar (and also the apical, vertical, dor-
composed of several (as opposed to one) segments. sal and axial spines).
Neck (N): elongated section of shell joining cephalis Sagittal ring (N) (Fig. 3A): a skeletal ring formed by
and thorax. the median bar at the base, the vertical and apical
Nodal points: see nodes. spines at the front and back, respectively, and an
Nodes (N, S) (Fig. 2F): area of the shell where two or arch joining these two spines at the topo lt can be
more bars meet (=nodal points). either free or embedded into the skeletal meshwork.
Oral teeth: see teeth. Second shell: see first shell.
Patagium (S) (Fig. 2Q, R): a delicate, spongy mesh- Secondary lateral spines: see lateral spines.
work in some Spongodiscidae which differs from Secondary spines (=by-spines): see spines.
the main shell by its looser structure. Segment (N) (Fig. 30): one of the several joints or
Peristome (N) (Fig. 3P): differentiated rim around the sections which compose a Nassellarian skeleton,
mouth of the last segment, usually poreless, often separated by the adjoining ones by an internal cir-
bearing teeth or other structures. cular thickening of the shell-wall ancl/or by an
Phacoid shell (S): structure formed by two concentric external stricture.
spherical medullary shells enclosed in one lenticu- Septae (N): Well-developed internal divisions
lar cortical shell, characteristic of the families between the segments of nassellarian shells.
Coccodiscidae and Phacodiscidae. Sieve-plate: see porous sieve-plate.
Polar caps (S) (Fig. 2J): Additional skeletal growth in Spines (N, S): any internal or external rod or needle
the form of latticed cupolae at both ends of some with one free end. According to their relative size
coccodiscid shells. and number, spumellarian spines can be primary
Pore arrangement (N, S): spatial distribution of pores (larger, usually fewer in number; Fig. 2B) and sec-
of the shell-wall; regular distributions can be ondary (smaller, more numerous, sometimes bris-
checkered (Fig. 3U) or in rows, either transversal tle- or thorn-shaped; Fig. 2B). In cross-section
(Fig. 3S, T) or longitudinal (Fig. 3V, W). spines can be triangular (3-bladed) with smooth or
Pores (N, S) (Fig. 2D, E): perforations in the skeleton serrated edges, or cylindrical; straight, or twisted
of the polycystines. (Fig. 2G). The distal end of spines can be
Porous sieve-plate (S) (Fig. 2P): a thin, perforated unbranched or branched (Fig. 2G). Club-like
plate that covers the spongy surface of some spines have a swollen or thickened distal end
Spongodiscidae.
(Fig. 2G). Radial spines are those radiating cen- Subelass Acantharia Müller, 1858
trifugally from the center of the shell (Fig. 2B). Subelass Radiolaria Müller, 1858
Spongy meshwork (N, S) (Fig. 2N-S): sponge-like (as Superorder Phaeodaria Haeckel, 1879
opposed to latticed) siliceous meshwork of thin, Superorder Polycystina Ehrenberg, 1838,
irregular, more or les s densely packed bars, main- emend. Riedel, 1967
ly in some spumellarian families (Actinornmidae, arder Collodaria Haeckel, 1881
Spongodiscidae ). Family Thalassicollida Haeckel, 1862
Strictures (N) (Fig. 30): joints between contiguous Family Collozoida Haeckel, 1862
segments, often marked by conspicuous external Family Thalassosphaerida Haeckel, 1862
constrictions. Family Sphaerozoida Haeckel, 1862
Teeth (N) (Fig. 3P): protruding apophyses around the arder Spumellaria Ehrenberg, 1875
mouth, of variable length and thickness (=oral Family Collosphaeridae Müller, 1858,
teeth, adoral teeth, terminal teeth, sub terminal emend. Strelkov and Reshetnjak, 1971
teeth). Family Actinommidae Haeckel, 1862,
Terminal teeth: see teeth. emend. Sanfilippo and Riedel, 1980
Third shell: see first shell. Family Coccodiscidae Haeckel, 1862,
Thorax (N) (Fig. 30, P): second segment ofmultiseg- emend. Sanfilippo and Riedel, 1980
mented nassellarians. Family Phacodiscidae Haeckel, 1881
Tripod (N): section ofthe internal skeleton formed by Family Spongodiscidae Haeckel, 1862,
the dorsal and primary lateral spines (Fig. 3A, B). emend. Riedel, 1967
In some shells the tripod protrudes externally Family Litheliidae Haeckel, 1862
forming the feet (Fig. 3Q). Family Pyloniidae Haeckel, 1881
Tubereles (N, S): pustule-like lumps (condyles, Family Tholoniidae Haeckel, 1862
mamillae) on the surface of the siliceous shell. arder Nassellaria Ehrenberg, 1875
Unilocular shell: see monolocular shell. Family Spyridae (=Trissocyclidae,
Velum (N) (Fig. 3X): a porous plate obliterating the Acanthodesmiidae) Ehrenberg, 1847,
mouth. emend. Petrushevskaya, 1971
Ventral spine: same as axial spine. Family Plagoniidae Haeckel, 1881,
Vertical spine (N) (Fig. 3A, B): Main unpaired spine emend. Riedel, 1967
projecting from the median bar in the vicinity of Family Theoperidae Haeckel, 1881,
the primary lateral spines and the axial spine. emend. Riedel, 1967
Wings (N) (Fig. 3R): lateral apophyses of the thorax Family Carpocaniidae Haeckel, 1881,
usually connected to the shell-wall by siliceous emend. Riedel, 1967
meshwork, derived from external projections of Family Pterocorythidae Haeckel, 1881,
the dorsal and primary lateral spines. emend. Riedel, 1967
Family Artostrobiidae Riedel, 1967,
Outline classification emend. Foreman, 1973
Family Cannobotryidae Haeckel, 1881,
The following synopsis, based chiefly on the scheme emend. Riedel, 1967
proposed by Riedel (1967b, 1971), is restricted to
extant families only. As pointed out above, it is a pro- Order and family-level diagnoses, and identifica-
visional elassification very likely to change as more tion of the species
structural and evolutionary data are obtained. Taxa in
bold underlined characters are those treated in detail The overall total of living polycystine species is prob-
in this chapter. ably around 300-600, but this review ineludes only
176 taxa. Unfortunately, our present state of knowl-
Kingdom Protista Haeckel, 1886 edge is insufficient for a significantly greater cover-
Phylum Sarcodina Hertwig and Lesser, 1876 age. All polycystine families need detailed taxonomic
Class Actinopoda Calkins, 1909 work, but some are especially poorly known (e.g.,
Subelass Heliozoa Haeckel, 1886 Actinornmidae, Pyloniidae, Litheliidae, Tholoniidae,
----------------------------------------------------------------
Plagoniidae). However, the inexperienced student Order Collodaria (Fig. 16.1-12)

should not be discouraged by the fact that the available Solitary or colonial polycystines without a siliceous
literature fails to assist in identifying very significant skeleton, or provided with simple or branched spic-
proportions of the shells present in any given sample. ules scattered in the calymrna. According to Haeckel
As mentioned above, most radiolarian-based ecologi- (1887), this group comprises 4 families:
cal and paleoecological surveys restrict their scope to Family Thalassicollida: solitary cells, no skeletal ele-
some 20-40 polycystines, the rest being ignored alto- ments; genera Actissa (Fig. 16.1), Thalassocampe
gether. Although coverage of an ampler inventory is (Fig. 16.2), Thalassopila, Thalassicolla, and
c1early desirable, these restricted databases allow one Thalassophysa;
to draw very valuable environment-related conc1u- Genus Collozoida: colonial, no skeletal elements;
sions. Furthermore, in order to circumvent identifica- genus Collozoum (Fig. 16.6-8, 16.11);
tion-related problems, the use of family-level assign- Genus Thalassosphaerida: solitary, with siliceous
ments has been advocated for environmental and spicules scattered in the calymma; genera
biogeographic analyses (Kruglikova, 1981, 1987). Thalassosphaera, Thalassoxanthium (Fig. 16.3),
Physematium, Thalassoplancta, Lampoxanthium
The following section presents succinct information (Fig. 16.5);
for the identification of the taxa recorded in the South Genus Sphaerozoida: colonial, with siliceous spicules
Atlantic Ocean, as well as for several others whose scattered in the calymrna; genera Belonozoum,
presence in the area is very likely, but has not been Sphaerozoum (Fig. 16.10), Raphidozoum (Fig.
confirmed yet. 16.12).
When applicable, species names are followed [in Due to their fragility, members of this group preserve
square brackets] by some of the other most comrnon poorly in net plankton samples, and either do not pre-
names under which the form has been recently cited in serve at all or are represented on1y by their spicules in
the literature. Diagnostic information is provided as sedimentary materials. Partly because of these limita-
short remarks stressing the characters which help dif- tions, information on their c1assification and distribu-
ferentiate the taxon from closely related forms; they tion is extremely scarce, and no further details are
are not meant to replace detailed descriptions, but in given on this group in this chapter. Detailed reviews
most cases these remarks, in combination with the of the colonial radiolarians, including several
illustrations, should suffice for producing adequate Collodaria, were produced by Hollande and Enjumet
identifications ofthe radiolarians treated. Keys are not (1953), Strelkov and Reshetnjak (1971), and
adequate for this particular group due to their lack of Swanberg (1979). Most of these species have tropical
flexibility, and because they are more prone to lead to distribution ranges in the three major oceans. In the
misidentifications given the high number of rare poly- south Atlantic they are probably restricted to waters
cystine species not inc1uded in this review. In order to associated with the equatorial current system, the
facilitate identification of the taxa, in some cases Tropics/Subtropics, and the oligotrophic Central Gyre
illustrations have be en arranged into groups of species (Fig. 11).
which share some conspicuous characteristics, and for
ease of comparison similar morphotypes are located Order Spumellaria (Fig. 16.13-90)
near-by on the plates. Shell size is generally of minor Solitary or colonial radiolarians with a well-devel-
value for identification purposes, for which reason no oped shell of radial symrnetry or one derived from the
detailed morphometric information is given; the above. Variations in the type ofsymrnetry inc1ude spi-
dimensions specified are approximate and are fur- ral shells (e.g., Fig. 16.87), asymrnetric, discoidal or
nished with the sole purpose of a general scale refer- lenticular (biconvex) (Fig. 2H, 1; 16.60-65), triaxonic
ence. The designation "group" denotes categories of (Fig. 2Q, R), quadrangular (Fig. 16.66, 16.67), etc. In
unc1ear taxonomic affinities, where probably several many cases two axes of symrnetry can be c1early dif-
related taxa are lumped. For each ofthe species treat- ferentiated (Fig. 2J) but, as opposed to the Nassellaria,
ed one or more references ("Ref.") providing detailed the larger axis is homoaxonic. The central capsule
descriptions and illustrations are inc1uded. (organic) of these cells has many small pores.
Family Collosphaeridae (Fig. 16.13-24) (81-164 11m). Boltovskoy and Riedel (1980) re-
Colonial polycystines, each individual has a single, cognized 3 formae (A, B and C) of this species in
thin-walled, spherical or subspherical latticed shell. Southwestern Atlantic planktonic materials. Ref.:
The Collosphaeridae is the only group of colonial Strelkov and Reshetnjak (1971), Boltovskoy and
polycystines with complete latticed shells. Colonies Riedel (1980).
consist of a gelatinous mass (which obviously disap- • Buccinosphaera invaginata Haeckel (Fig. 16.17)
pears in the sedimentary record, as well as in many [=Collosphaera invaginata]. The smooth shell
net-plankton samples where it breaks down) in which produces several pored tubes directed toward the
hundreds to thousands of shells are irnmersed (Fig. center of the sphere. Rather small, irregular pores.
1C-E). The shape of the colony is not species-speci- Shell diameter: 100-130 11m. Ref.: Strelkov and
fic; it may be spherical, ellipsoidal, cylindrical, rib- Reshetnjak (1971), Nigrini (1971).
bon-shaped, etc., measuring up to several centimeters • Collosphaera hux/eyi Müller (Fig. lE, 16.13).
in length and a few millimeters in diameter. The sili- Shells with small to medium-sized pores scattered
ceous shells are always represented by a single perfo- about the surface only; no spines or tubes. Shell
rated sphere (internal spheres are never present), with diameter: 80-150 .11m. Ref.: Strelkov and
or without centrifugal (external) or centripetal (inter- Reshetnjak (1971): Boltovskoy and Riedel (1980).
nal) tubular projections andJor spines. Spines (when • Collosphaera macropora Popofsky (Fig. 16.15).
present) are conical (circular in cross-section). As No spines or tubes on shell surface; few very large
with most other polycystines, specific assignments are pores, sometimes angular. Shell diameter: 100-
based almost exclusively on the skeleton; however, 120 11m. Ref.: Strelkov and Reshetnjak (1971),
studies of entire colonies, which allow investigating Boltovskoy and Riedel (1980).
the intraspecific morphologic variability ofthe collos- • Collosphaera tuberosa Haeckel (Fig. 16.14). No
phaerids, indicate that quite dissirnilar shell morpho- spines or tubes on shell surface, but with conspicu-
types can coexist within the same colony, thus stress- ous lumps and depressions; many small, irregular-
ing the assumption that at least some of the specific ly shaped pores. Shell diameter: 50-300 11m.Ref.:
divisions based on the siliceous sphere alone are spu- Strelkov and Reshetnjak (1971), Boltovskoy and
rious (e.g., Kleijne, 1987; Petrushevskaya and Riedel (1980).
Swanberg, 1990). • Siphonosphaera martensi Brandt (Fig. 16.20).
• Acrosphaera murrayana (Haeckel) (Fig. 16.19) Each pore bears a short centrifugal tube, tube
[=Polysolenia murrayana]. Large pores, each sur- walls are imperforate. Shell diameter: 90-100 /lID.
rounded by a crown of short spines. Shell diame- Ref.: Strelkov and Reshetnjak (1971).
ter: 70-190 11m. Ref.: Strelkov and Reshetnjak • Siphonosphaera po/ysiphonia Haeckel (Fig.
(1971), Nigrini and Moore (1979). 16.23) [=?Siphonosphaera socialis, ?Siphono-
• Acrosphaera spinosa (Haeckel) group? (Fig. ID, sphaera tenera]. Tubes of variable length are -
16.18) [=Polysolenia spinosa, ?P. lappacea, ?P. present on some ofthe pores, tube walls are imper-
jlammabunda]. Irregular pores and many irregu- forate. Shell diameter: 80-120 11m. According to
larly arranged spines scattered about the surface, Strelkov and Reshetnjak (1971) S. socia lis Haeckel
some of the latter extending from the pore-rims. has usually smaller shells (50-80 11m).Ref.: Nigrini
Spine and pore patterns are variable. Shell diame- and Moore (1979), Boltovskoy and Riedel (1980).
ter: 60 to >200 11m.Various authors attempted to • So/enosphaera chierchiae Brandt (Fig. 1C,
group the different morphotypes within this taxon. 16.22) [=Otosphaera polymorpha, Solenosphaera
Nigrini and Moore (1979) recognized 4 closely collina]. The shell bears 3-4 (up to 6) short tubes
related species: Polysolenia aktios Nigrini, very with perforated walls whose distal end is tapered,
similar to Acrosphaera spinosa but restricted to ending in a slanted pore provided with 1-2 con-
the Subarctic (103-238 11m);Polysoleniajlamma- spicuous spines. Shell diameter: 65-100 11m.Ref.:
bunda (Haeckel), with somewhat larger pores and Strelkov and Reshetnjak (1971), Nigrini and
longer spines (81-136 11m);Polysolenia lappacea Moore (1979).
(Haeckel), spines curved and often interconnected • So/enosphaera po/yso/enia Strelkov and
(81-109 11m); and Polysolenia spinosa (Haeckel) Reshetnjak (Fig. 16.21). With many (rather than 3-
5, as in Solenosphaera zanguebarica) short tubes obvious that, based on this trait, Haeckel (as well as
with perforated walls, and smaller, more angular many other authors) assigned new names to growth
pores. Shell diameter: 90-280 um. Ref.: Strelkov stages still missing the outermost sphere(s) (see Figs.
and Reshetnjak (1971). 13 and 14). Furthermore, while growth of an actinom-
• Solenosphaera zanguebarica (Ehrenberg) group? mid as far as we know proceeds from the center
(Fig. 16.24) [=Solenosphaera polymorpha, toward the periphery (Fig. 15, upper panel), dissolu-
Otosphaera polymorpha, Disolenia zanguebarica, tion works in the opposite direction, innermost, more
?Disolenia quadrata]. General outline of shell delicate shells usually disappearing before the more
often subtriangular, the vertices extending into 3-4 robust cortical ones. Thus, material s from the sedi-
short tube-like protrusions with perforated walls ments offer yet another suite of "new species", this
and open ends. The rim of these tubes may bear time missing the medullary (rather than the cortical)
spines. Pores irregular in shape and size, but gener- shells.
ally roundish or subangular. A morphotype very • Acanthosphaera actinota (Haeckel) (Fig. 16.25)
similar to S. zanguebarica (and most probably con- [=Acanthosphaera tenuissima]. Single shell with
specific with it), usually cited as Disolenia quadra- large, regular polygonal meshes separated by very
ta (Ehrenberg) (Fig. 16.24a), differs by having thin bars. Nodal points bear short, bristle-shaped
somewhat larger tubes which are better differentiat- spines; no primary (conspicuously larger) radial
ed from the main body of the shell. Shell diameter: spines (broken-off on specimen photographed).
100-150 um. Ref.: Strelkov and Reshetnjak (1971). Shell diameter: 60-90 um. Ref.: Boltovskoy and
• Tribonosphaera centripetalis Haeckel (Fig. Riedel (1980).
16.16). Outer shell surface smooth, but with many • Acanthosphaera dodecastyla Mast (Fig. 16.26).
slender spines directed toward the center of the Single shell with large circular pores with very
sphere; pores irregular in size and shape. Shell conspicuous polygonal frames, many (10-20)
diameter: 100-120 um. Ref.: Strelkov and robust, 3-bladed spines. Shell diameter without
Reshetnjak (1971), Boltovskoy and Riedel (1980). spines: 50-80 um. Ref.: Popofsky (1913),
Boltovskoy and Riedel (1980).
Family Actinommidae (Fig. 16.25-59) • Acanthosphaera pinchuda Boltovskoy and
Solitary species with latticed or spongy spherical, Riedel (Fig. 16.28). Single shell with regular, sub-
subspherical, or ovoid shells (not lenticular); with or circular pores surrounded by conspicuous protru-
without medullary shells. Surface of shell is often ding frames which extend into thin, thread-like
covered with spines, but not tubes. All actinommids spines at the nodal points. No primary (conspicu-
posses either single or multiple, concentric spherical ously larger) radial spines. Shell diameter without
or ovoid shells. When several shells are present they spines: 90-120 um. Ref.: Boltovskoy and RiedeI
are connected to each other by radial beams which (1980).
pierce the cell. An enormous variety of forms was • Actinomma antarcticum (Haeckel) group? (Fig.
described in this family whose identification has tra- 16.48) [=Diploplegma banzare, Diploplegma
ditionally been based on Haeckel's (1887) system. aquatica]. Three concentric shells. Cortical shell
Haeckel based the classification of the actinommids composed of a very irregular, sometimes sponge-
(=suborder Sphaeroidea, exclusive of the Collo- like network of thick anastomosing bars. Second
sphaeridae) on the following characters (in decreasing shell large, irregularly shaped, spongy. First shell
order of importance; see Fig. 14): 1. Number of pri- small, circular, rarely visible. Nigrini (1967)
mary radial spines; 2. Number of concentric spheres; described Actinomma medianum, which differs
3. Position of concentric spheres (intra- or extracapsu- from A. antarcticum in that it has a simply latticed
lar), type and relative size of spines, presence of by- cortical shell and a more delicate medullary mesh-
spines, type of medullary shell, etc.). However, the work. Cortical shell diameter: 200-400 um. Ref.:
number of primary spines varies intraspecifically, Riedel (1958), Nigrini (1967).
whereas the number of main concentric spheres, • Actinomma arcadophorum Haeckel (Fig. 16.49).
which within some bounds might indeed be species- Medullary shells similar to Actinomma antarcticum;
specific (Riedel and Sanfilippo, 1986), can only be cortical shell is composed of an irregular, sponge-
used in the case of fully-grown individuals. It is quite like or lace-like network of very thin, delicate anas-
tomosing bars. Cortical shell diameter: 200-250 um, • Cenosphaera spp. group (Fig. 16.29) [=Ceno-
Ref.: Haeckel (1887), Nigrini and Moore (1979). sphaera elysia, C. compacta, C. hirsuta]. Single
•. Actinomma leptodermum (Jorgensen) (Fig. sphere, often thick-walled. Pores more or less re-
16.37) [=Echinomma leptodermum]. Three con- gular, circular, or irregular, of variable shape and
centric shells, pores on outermost shell relatively size, with or without polygonal frames, with or
large, subcircular, irregularly arranged; surface without thin, bristle-shaped by-spines or thoms.
covered by numerous short 3-bladed spines; no No primary (conspicuously larger) radial spines.
primary (conspicuously larger) radial spines. Highly variable group with many different mor-
Outermost shell diameter without spines: 60-120 photypes present chiefly in middle and high lati-
um. Ref.: Nigrini and Moore (1979). tudes (e.g., C. cristata Haeckel in Antarctic waters,
• Actinomma sol Cleve (Fig. 16.44) [=Theco- cf. Petrushevskaya, 1967; C. compacta Haeckel
sphaera radians]. Three concentric shells. and C. elysia Haeckel in subantarctic-transitional
Outermost with regular, circular pores with well areas, cf. Boltovskoy and Riedel, 1980). Shell
developed polygonal frames, usually bearing short diameter: 50-320 um. Ref.: Petrushevskaya
bristle-shaped spines on the nodes (broken off in (1967), Boltovskoy and Riedel (1980).
the specimen photographed); no primary (conspic- • Centrocubus cladostylus Haeckel (Fig. 16.58).
uously larger) radial spines (see remark for Medullary shell single, composed of bars that
Thecosphaera inermis). Outermost shell diameter: define a small cube (Fig. 2C); cortical shell a
70-100 um. Ref.: Hollande and Enjumet (1960), spongy meshwork arising irnmediately from the
Boltovskoy and Riedel (1980). medullary shell, supported by many large, 3-blad-
• Arachnosphaera myriacantha Haeckel (Fig. ed spines that protrude outside ofthe spongy mass.
16.45). Fully grown specimens with many (up to Cortical shell diameter without spines: 150-200
over 7; 4 in the specimen illustrated) concentric um. Ref.: Haeckel (1887).
spheres, the innermost with regular hexagonal • Cladococcus cervicornis Haeckel (Fig. 16.41).
meshes and cylindrical spines arising from the Single shell with very irregular sub-polygonal
nodes. At regular distances these spines produce pores; at many nodal points slightly curved, cylin-
branches which anastomose laterally forming sub- drical spines project radially dichotomizing
sequent spheres with a delicate, irregular, cobweb- repeatedly starting about 2/3 of the way from the
like network. Branching spines protrude from the cortical shell. Shell diameter without spines: ea.
surface of the last shell. Diameter of innermost 70 um. Ref.: Boltovskoy and Riedel (1980).
shell: ea. 100 um, Ref.: Haeckel (1862). • Cladococcus megaceros Hollande and Enjumet
• Astrosphaera hexagonalis Haeckel (Fig. 16.40). (Fig. 16.42). Similar to Cladococcus cervicornis,
Two shells and many (>6) primary radial spines. differs in that spines can be 3-bladed, thickening
Inner shell with regular polygonal meshes and thin toward their distal end; branches are short and
bars. Outer shell with very large, triangular mesh- dull, resembling the homs or a reindeer. Pores are
es formed by anastomosing of lateral branches more irregular in size and shape. Shell diameter
produced by the long, 3-bladed primary spines without spines: ea. 60 um. Ref.: Hollande and
arising from the first shell. Spines extend beyond Enjumet (1960), Boltovskoy and Riedel (1980).
the very open outer shell. Diameterof inner shell: • Cromyechinus antarctica (Dreyer) (Fig. 5A, A',
150 um. Ref.: Haeckel (1887). 14) [=Actinomma boreale, Cromyechinus boreal-
• Carposphaera acanthophora (Popofsky) (Fig. is]. Fully grown specimens with 4 concentric, lat-
16.39). One latticed cortical shell and one medul- ticed shells. Outermost thin-walled, with very
lary shell represented by an irregular formation of small pores. Third shell thicker, with large, irreg-
anastomosing centripetal bars arising from the ularly shaped pores. Surface covered with many
cortical shell (may be missing in poorly preserved short, stout, 3-bladed spines, at one of the poles
materials; not focused in specimen illustrated). spines may be denser and larger, forming a
Pores on outer shell irregular in shape and size, pylome. Major diameter of outermost shell: 100-
generally subcircular. Surface rough or thomy. 160 um. Ref.: Petrushevskaya (1967).
Diameter of cortical shell: 150-270 um. Ref.: • Cromyechinus icosacanthus Haeckel (Fig. 2B).
Benson (1966). Fully grown specimens with 4 latticed concentric
-í-----
shells and many (>6) stout, 3-bladed spines. delicate fourth shell. Outennost shell diameter
Outennost shell thin-walled, delicate, with very without spines: ea. 130 um. Ref.: Boltovskoy and
small, regular, circular pores. Third shell with Riedel (1980).
large, irregular, polygonal pores. Outennost shell • Hexacontium armatum/hostile Cleve group (Fig.
diameter without spines: ea. 160 um. Ref.: 16.32). [=Hexacontium armatum, Hexacontium
Haeckel (1887). hostile, THexacontium entacanthum]. Three con-
• Cromyomma circumtextum Haeckel (Fig. 16.47). centric latticed shells; surface of outennost usual-
Fully grown specimens with 4 latticed concentric ly thorny or spiny, pores medium-sized, regular or
shells and many 3-bladed spines of irregular size irregular in size and distribution, with or without
and distribution. Outennost shell very delicate, polygonal frames. Usually 6 main spines opposite
thin-walled, with thread-like bars and large, irreg- in pairs in three dimensive axes perpendicular to
ular, polygonal meshes; third shell with large, one another. Outennost shell diameter without
irregular pores and thick bars. Due to its delicacy, spines: 70-100 um. Poorly defined morphotype,
the fourth shell is very likely to be absent in sedi- probably includes several related species. Ref.:
mentary materials, in which case this species may Boltovskoy and Riedel (1980).
be identified as Hexa/onche aristarchi (see • Hexacontium laevigatum Haeckel (Fig. 16.31).
below). Outennost shell diame ter without spines: Similar to Hexacontium armatum/hostile, except
ea. 220 um. Ref.: Haeckel (1887). that the surface of the outennost shell is smooth,
• Druppatractus irregularis Popofsky (Fig. 16.27). and pores are smaller, circular, unframed of regu-
Two latticed shells, outermost thin-walled, lar size and distribution. Outennost shell diameter
smooth, with small regular, circular pores; medul- without spines: 70-120 urn. Ref.: Benson (1966),
lary shell pear-shaped, Two main polar spines of Nigrini and Moore (1979).
equal or different size (additional, usually smaller • Octodendron cubocentron Haeckel (Fig. 16.56).
spines may be present as well). Cortical shell Similar to Centrocubus cladosty/us, except that
diame ter without spines: ea. 80 um. Ref.: Benson spongy meshwork starts at some distance from the
(1966). cubical medullary shell. Outer shell diameter
• Heliaster hexagonium Hollande and Enjumet without spines: ea. 200 um. Ref.: Haeckel (1887).
(Fig. 16.46). Two latticed shells, medullary as in • Plegmosphaera entodictyon Haeckel (Fig. 16.54).
Carposphaera acanthophora (may be missing in Shell is a spongy mass with a central cavity; the
poorly preserved materials). Pores on cortical spongy meshwork is denser in the vicinity of the
shell very regular, polygonal, with thin bars; bris- central cavity than at the periphery of the shell.
tle-shaped spines on nodes. Outer shell diameter Shell diameter: ea. 200 um, Ref.: Hollande and
without spines: ea. 230 um. Ref.: Hollande and Enjumet (1960).
Enjumet (1960). • Plegmosphaera exodictyon Haeckel (Fig. 16.55).
• Heliosoma echinaster Haeckel (Fig. 16.43) Similar to P/egmosphaera entodictyon, except that
[=Tetrapeta/on e/egans]. Two concentric shells. the spongy meshwork is denser both in the vici-
Medullary shell composed of a loose network of nity of the central cavity and at the periphery of
thin anastomosing bars. Outer shell similar to that the shell, and looser midway. Shell diameter: ea.
of Heliaster hexagonium, except for the presence 400 urn. Ref.: Haeckel (1887).
of many (>6) long, 3-bladed primary spines. Outer • Plegmosphaera pachyplegma Haeckel (Fig.
shell diameter without spines: ea. 160 um. Ref.: 16.53). Similar to Plegmosphaera entodictyon,
Hollande and Enjumet (1960). except that the spongy meshwork is den ser at the
• Hexacontium aristarchi (Haeckel) (Fig. 16.33) periphery of the shell than in the vicinity of the
[=Hexa/onche aristarchi]. Similar to Hexacontium central cavity. Shell diameter: ea. 200 um, Ref.:
armatum/hosti/e, except that pores on outennost Hollande and Enjumet (1960).
shell are larger and more irregular in size and dis- • Saturnalis circularis Haeckel (Fig. 16.36). Two
tribution. May have 1 medullary shell (7). It is concentric latticed shells, the cortical with circular
probable that H aristarchi is a developmental to subcircular pores and rough surface. Two pri-
form of Cromyomma circumtextum, from which it mary radial polar spines (which extend as internal
differs by lacking the outennost, very slender and beams joining the 2 shells) joined distally by a cir-
,
'-
cular ring. Cortical shell diame ter without spines: • Styptosphaera spumacea Haeckel (Fig. 16.50).
70-80 urn. Ref.: Nigrini (1967). Shell is an irregular spongy mass without central
• Spongodictyon spongiosum (Müller) (Fig. 16.57) cavity. Shell diameter: ea. 450 um, Ref.:
[=Dictyosoma spongiosum]. Generally similar to Boltovskoy and Jankilevich (1985).
Spongoplegma rugosa, except that center hosts a • Thecosphaera inermis (Haeckel) (Fig. 16.38).
double medullary shell; second medullary shell Three concentric latticed shells; pores on outer-
usually incompletely developed. Outermost shell most relatively small, circular, regularly arranged,
diameter: 200 urn. Ref.: Müller (1858). without frames (in some specimens frames around
• Spongoplegma antarcticum Haeckel (Fig. 16.51). the pores can develop, in which case this species
Spongy mas s with a single irregular, sponge-like merges with Actinomma sol). Cortical shell usual-
medullary shell in the center. Outer shell diameter: ly smooth, barren of spines. Cortical shell diame-
ea. 150 urn. Ref.: Boltovskoy and Riedel (1980). ter: ea. 80 um. Ref.: Boltovskoy and Riedel (1980).
• Spongoplegma rugosa Hollande and Enjumet
(Fig. 16.52). Spongy mass with a single latticed Family Coccodiscidae (Fig. 2H, J)
medullary shell in the center (not focused in the Latticed discoidal or lenticular shell enclosing a single
illustration); spongy meshwork is looser toward or double medullary shell, and surrounded by an equa-
the periphery ofthe shell. Outer shell diameter: ea. torial zone of spongy or concentrically-chambered
300 um. Ref.: Hollande and Enjumet (1960). structures (Fig. 2H), or forms with an ellipsoidal corti-
• Spongosphaera streptacantha Haeckel (Fig. cal shell equatorially constricted enclosing a single or
16.59). Two concentric, spherical, latticed medul- double medullary shell (Fig. 2J). The formerly acti-
lary shells and one spongy cortical shell. With nommid subfamily Artiscinae was transferred to the
large, 3-bladed spines with serrated edges origi- Coccodiscidae by Sanfilippo and Riedel (1980) due to
nating in the second medullary shell and protrud- its phylogenetic affinities with extinct coccodiscids.
ing conspicuously outside ofthe spongy outermost • Didymocyrtis tetrathalamus (Haeckel) (Fig. 2J,
shell. Outer shell diameter without spines: ea. 300 16.77) [=Ommatartus tetrathalamus, Panartus
urn. Ref.: Hollande and Enjumet (1960). tetrathalamus]. Cortical shell ("twin shell") cylin-
• Stylatractus spp. group (Fig. 2A; 16.35). [= drical to ellipsoidal with a very conspicuous equa-
?Stylatractus, ?Axoprunum, ?Stylosphaera, torial constriction. Two medullary shells (outer-
?Xiphosphaera, ?Lithatractus, ?Xiphatractus]. most slightly compressed, innermost spherical)
Usually two latticed shells, outermost spherical to joined to the cortical by several radial beams con-
oval-shaped, often thick-walled with irregular fined to the equatorial plane (Fig. 16.77b). May
pores; medullary shell spherical. Two polar spines have polar caps (Fig. 2J) or spines, and supple-
of equal or different size, with or without secon- mental peripheric growth (mantle; Fig. 16.77a).
dary spines. Two-spined and usually two-shelled Height of cortical twin shell: 90-140 um, height of
actinommids comprise a large, sometimes abun- polar caps: 36-63 um. The subspecies D. t. coron-
dant, highly variable and very poorly studied atus is identical to D. t. tetrathalamus, except for
group. Outer shell diameter without spines: 100- the development of a corona of stout, unbranched,
150 um. Ref.: Nigrini and Moore (1979). 3-bladed spines on the distal ends ofthe twin shell
• Stylosphaera melpomene Haeckel (Fig. 16.34) and/or on the polar caps (Nigrini, 1970). Ref.:
[=Stylacontarium bispiculum]. Two or 3 (?) lat- Nigrini and Moore (1979).
ticed shells, outermost with characteristically sub- • Spongoliva e/lipsoides Popofsky (Fig. 16.76).
quadrangular outline, moderately thick-walled, Generally similar to D. tetrathalamus, except that
thorny, with irregularly shaped pores; medullary cortical shell is a much looser and irregular mesh-
shell spherical. Six stout beams join the two shells; work. Fully grown specimens usually have a well
usually two of these beams protrude outside as developed mantle. Height of cortical shell: 100-
short, 3-bladed polar spines, but the other two 250 um. Ref.: Benson (1966).
pairs may also extend slightly beyond the cortical
shell-wall. Polar spines of equal or different size. Family Phacodiscidae (Fig. 16.78, 16.79)
Outermost shell diameter without spines: 80-130 Lenticular, biconvex, latticed cortical shell, not sur-
urn. Ref.: Benson (1966). rounded by spongy or chambered structures, within
which a small, spherical single or double medullary unchambered arms radiating at equal or almost
shell is enclosed. The margin (but less commonly the equal angles from a central disco Length of arms
surfaces) of the cortical shell may bear radial spines. (from center of shell): 130-150 urn. Ref.: Nigrini
• Heliodiscus asteriscus Haeckel (Fig. 16.78). and Moore (1979).
Outer shell lenticular, with regularly arranged cir- • Dictyocoryne truncatum (Ehrenberg) (Fig.
cular pores and ea. 8-15 radial spines on the equa- 16.69). Very similar to D. profunda (and probably
torial plane; inner shell spherical. Diameter of cor- synonymous with it: intergrading specimens are
tical shell without spines: 120-200 um. Ref.: very common); differs in having much broader
Nigrini and Moore (1979). arms and usually a well developed patagium (Fig.
• Sethodiscus macrococcus Haeckel (Fig. 16.79). 2Q, R). Length of arms (from center of shell): ca.
Very similar to H asteriscus, but without radial 130 um. Ref.: Nigrini and Moore (1979).
spines. Diameter of cortical shell: ea. 150 urn. Ref.: • Euchitonia elegans/furcata (Ehrenberg) group?
Haeckel (1887), Boltovskoy and Riedel (1980). (Fig. 16.70) [=?Euchitonia furcata, Euchitonia
elegans]. Differs from D. profunda in that the 3
Family Spongodiscidae (Fig. 16.60-75) arms are more slender, longer, and their orienta-
Discoidal or cylindrical, spongy or finely chambered tion defines two larger angles and one smaller one.
skeleton, with or without surficial pore-plate, often E. elegans intergrades smoothly with Euchitonia
with radiating arms or marginal spines. The members furcata; end members differ in that distal tips of
of this family are characterized by possessing skele- arms taper in E. elegans (Fig. 16.70c), while in E.
tons which are partly or entirely spongy in appear- furcata their terminations are club-shaped and
ance. However, as opposed to the Actinommidae, more blunt (Fig. 16.70a, b). Length of arms (from
which can also have spongy skeletons, the center of shell): 150-300 um. Ref.: Nigrini and
Spongodiscidae are not spherical. Their overall shape Moore (1979).
can be lenticular (biconvex discs, Fig. 21), cylindrical • Spongaster tetras Ehrenberg irregularis Nigrini
(Fig. 16.74), quadrangular or subquadrangular in out- (Fig. 16.66). Generally similar to S. tetras tetras ,
line (Fig. 16.67), or Y-shaped (Fig. 2Q, R). With the except that shell is less regular, angles between
exception of the cylinders, all others are depressed or "arms" are uneven, and outline is an elongated
flattened (rather than circular in cross-section, Fig. rectange (rather than a square). Length of longer
21). Lenticular, quadrangular, and Y-shaped forms side: 140-260 um. Ref.: Nigrini and Moore (1979).
may be entirely composed of a spongy mass with no • Spongaster tetras tetras Ehrenberg (Fig. 16.67).
discemible structure (in which case the central part of Shell outline rectangular with rounded comers;
the skeleton is often thicker and/or denser, and there- denser spongy meshwork defines 4 equidistant
fore appears darker in the light microscope; Fig. 20, pear-shaped "arms" radiating from the center. Side
16.64), or may posses a small central chamber sur- ofrectangle: 150-300 um. Ref.: Nigrini and Moore
rounded by concentric or spiral, continuous or inter- (1979).
rupted bands (Fig. 2R). The surface of some forms • Spongocore cylindrica Haeckel (Fig. 16.74)
may be partly or totally covered with a very thin, po- [=Spongocore puella, Spongocore diplocylindrica].
rous sieve-plate, which in lenticular forms may extend Spongy cylinder with or without protruding spines
beyond the central spongy mass forming a delicate and mantle, often with two slight constrictions.
equatorial girdle around the periphery of the shell Totallength: 200-360 urn. Ref.: Benson (1966).
(Fig. 2P) (these morphotypes were formerly included • Spongodiscus resurgens Ehrenberg (Fig. 16.64).
in the family Porodiscidae). Spongy biconvex disc with no discemible struc-
• Amphirhopalum ypsilon Haeckel (Fig. 2R, 16.71, ture, without spines on edges or surfaces of disco
16.75). Shell with two opposite chambered arms, Diameter of disc: 100-400 um. Ref.: Boltovskoy
one of them may be bifurcated distally, in which and Riedel (1980).
case the shell outline becomes trigonal. Total • Spongopyle setosa Dreyer (Fig. 2N, 16.63)
length: 200-300 um. Ref.: Nigrini and Moore [=Spongopyle osculosa]. Similar to S. resurgens,
(1979). except for the presence of a tubular, spiny or
• Dictyocoryne profunda Ehrenberg (Fig. 2Q, notched pylome on the margino Diameter of disc:
16.68) [=Hymeniastrum euclidis]. Three spongy, 100-300 urn. Ref.: Riedel (1958).
ttn~
• Spongotrochus glacialis Popofsky (Fig. 21, 16.61). • Larcopyle butschlii Dreyer (Fig. 5B, B', 16.85).
Similar to S. resurgens, except for the presence of Ellipsoidal outer shell with irregular pores, surface
spines on the surfaces andlor on the margins ofthe often thorny or spiny, inner structure spiral. One
disco Can have an inconspicuous pylome. Diameter of the poles bears a pylome surrounded by larger
of disc (without spines): 100-500 um. Ref.: Riedel spines. Major diameter: 80-170 um. Ref.: Nigrini
(1958), Petrushevskaya (1967). and Moore (1979).
• Spongurus pylomaticus Riedel (Fig. 16.72). Shell • Larcospira quadrangula Haeckel (Fig. 16.86).
subcylindrical or elongate ellipsoidal, inner dense Shell consists of two open spirals arising from a
spongy meshwork surrounded by looser mesh- cornmon origino Breadth of shell: 120-250 um.
work, surface covered with bristle-like spines, Ref.: Nigrini and Moore (1979).
with a pylome at one ofthe poles. Length: 70-250 • Lithelius minor Jorgensen group? (Fig. 16.87).
um, Ref.: Riedel (1958), Petrushevskaya (1967). Tightly wound spiral with thorny or spiny surface,
• Spongurus spp. group? (Fig. 16.73). Irregular, outline circular or ovoid. Diameter: 80-150 um.
oval-elongate spongy mass, sometimes with con- Ref.: Nigrini and Moore (1979).
centric or spiral rings, with spines protruding at • Lithelius nautiloides Popofsky (Fig. 5C, C',
the poles. Length: ea. 100-150 urn. Ref.: 16.88). Small, spherical medullary shell surround-
Petrushevskaya (1967). ed by an involute spiral of 4-5 whorls which
• Stylochlamydium asteriscus Haeckel (Fig. 2P, increase in width outwards; surface spiny.
16.65) [=?Stylochlamydium venustum]. Biconvex Diameter: 100-230 um. Ref.: Petrushevskaya
spongy disc more or less clearly partitioned into (1967).
chambers by circular or spiral, continuous or bro- • Pylospira octopyle Haeckel (Fig. 16.90)
ken, and radial bands. Both surfaces covered by a [=?Phorticium pylonium]. A series of spirally
thin porous sieve plate which can extend beyond arranged chambers; outline ellipsoidal; surface
the central spongy mass. With or without protrud- spiny. Shell diameter: 80-130 um. Ref.: Nigrini
ing spines. Very similar morphotypes with an oval and Moore (1979).
outline and a pylome have been cited under the • Tholospira cervicornis Haeckel group (Fig.
name Stylochlamydium venustum Bailey (Fig. 16.89). Irregular meshwork with pores of variable
16.65b). Diameter of disc: 150-350 um. Ref.: size; surface spiny. Probably an artificial category
Boltovskoy and Vrba (1988). for lumping various juvenile, broken and poorly
• Stylodictya aculeata Jorgensen (Fig. 16.62). Flat known Litheliidae and Pyloniidae. Ref.:
disk not thickened in the center, with clearly Petrushevskaya (1967), Takahashi and Honjo
defined circular concentric rings, the innermost (1981).
rosette-shaped. Diameter of disc: ea. 150 um. Ref.:
Petrushevskaya (1967), Boltovskoy and Vrba Family Pyloniidae (Fig. 16.80-83)
(1988). The major part of the shell is composed of a series of
• Stylodictya multispina Haeckel (Fig. 20; 16.60) successively larger elliptical latticed girdles in three
[=Stylodictya validispina, Stylodictya tenuispina]. mutually perpendicular planes, with the major diame-
Similar to S. aculeata, except that all rings are cir- ter of each girdle being the minor diameter ofthe next
cular to subcircular (rather than rosette-shaped), larger one (Fig. 2L, M). The center is occupied by a
center may be thickened (darker), and usually with small ellipsoidal structure - the microsphere (see
marginal spines. Diameter of disc: 100-200 um, Dumitrica, 1989).
Ref.: Boltovskoy and Vrba (1988). • Dipylissa bensoni Dumitrica (Fig. 16.83). Shell
fomed by 3 systems of globular caps, in optical
Family Lithelüdae (Fig. 16.85-90) section appearing as oval concentric subhemi-
The lattice of the ellipsoidal, spherical or lenticular spheres interconnected by numerous thin radial
shell is totally or partially arranged along a bilaterally beams. Major diameter: 90-120 urn. Ref.:
syrnmetrical spiral. Although very abundant, due to Dumitrica (1988).
their complicated architecture the litheliids are poorly • Octopyle stenozona group? Haeckel (Fig. 2 L, M;
known, for which reason the morphotypes defined 16.80) [=Tetrapyle octacantha]. Shells with 2
may include several different forms. systems of latticed girdles (as described for the
diagnosis ofthe family, see above) and large gates syrnmetry of this group is characterized by the fact
(Fig. 2L, M). Major diameter: 100-250 um. Some that the two extremes of the major axis define two
authors (e.g., Benson, 1966; Nigrini and Moore, morphologieally different poles of the shell. One of
1979) recognize 2 species within this group (?): these, conventionally accepted as the top or anterior
Tetrapy/e octacantha Muller and Octopy/e steno- end, is where the cephalis is located. A widely recog-
zona Haeckel; while others (e.g., Welling, 1997) nized, albeit seldom utilized, feature of primary
consider o. stenozona as a rare tropical end mem- importance for the classifieation of the Nassellaria is
ber variant of T. octacantha (the wo froms were the internal skeleton. The internal skeleton consists of
found to covary in tropical plankton). According a complex set of spines and connecting bars enclosed
to Benson (1966), o. stenozona is distinguished in the cephalis (Fig. 3A-C), which allow comparison
by its very narrow second transverse and lateral of homologous structures in forms differing widely in
girdles, short saggital axis, generally quadrangular their external morphology. Unfortunately, analysis of
outline, and the presence of 2 heavy polar beams these features requires dedicated efforts at under-
which occupy the principal axis. Ref.: Nigrini and standing the complex spatial relationships involved.
Moore (1979). Furthermore, observation of this internal skeleton is
• Phorticium elevei (Jorgensen) (Fig. 2L, 16.82) only feasible with well preserved individual s oriented
[=Phorticium py/onium]. Generally resembling o. in the right position, which is seldom the case in spe-
stenozona, except that central part is a large struc- cimens mounted in permanent slides. In addition to
ture formed by a series of spirally arranged cham- the small scattered perforations typieal of the
bers. Major axis of shell: 150-200 um, Ref.: Spumellaria, the central capsule of the Nassellaria is
Petrushevskaya (1967). usually provided with a single larger pore.
• Pylolena armata Haeckel group? (Fig. 16.81)
[=?Hexapyle spp.]. A central pyloniid shell sur- Family Spyridae (=Trissoeyclidae) (Fig. 16.91-103)
rounded by 3 centrifugally radiating girdles whose The skeleton is represented by a well-developed D-
distal ends join forming a trigonal outline; this shaped sagittal ring (median bar and anastomosed ver-
structure may be surrounded by supplementary tical and apical spines), either free (Fig. 16.10 1) or
growth in the form of a spherical latticed test. embedded into the latticed cephalic wall, in which
Diameter of fully developed shell: ea. 150 um. case the eephalis is usually bilaterally lobed (Fig.
Ref.: Nigrini and Moore (1979). 16.93, 16.94, 16.100). Sometimes with thorax, abdo-
men always absent. The typical heteropolar nassellar-
Family Tholoniidae (Fig. 16.84) ian syrnmetry is often inconspicuous in the Spyridae.
Completely latticed shell, without larger openings, • Acanthodesmia viniculata (Muller) (Fig. 2C;
and with constrictions that define several (typically 6) 16.96) [=Giraffospyris angu/ata]. Shell composed
dome-shaped protuberances (Fig. 2K). of a D-shaped sagittal ring, a basal ring and a fron-
• Cubotholus spp. (Fig. 2K, 16.84) [=Th%nium tal ring (see Fig. 3C). Breadth of frontal ring: ea.
spp., Amphitholus spp., Cubotholus cf. orthoce- 140-180 urn. Ref.: Petrushevskaya (1971 a),
ras]. Cortical shell composed of 6 hemispherical Nigrini and Moore (1979).
cupo las, opposite in pairs on the poles of 3 mutu- • Amphispyris reticulata (Ehrenberg) (Fig. 16.95)
ally perpendicular axes. Variously shaped "central [=Liriospyris reticu/ata, Tlholospyris pro cera ].
chamber" with or without enclosed small medul- From the D-shaped sagittal ring 6 pairs of bars
lary shell. Diameter: ca. 150 um, Ref.: Haeckel arise whieh branch and anastomose forming the
(1887). latticed lateral walls of the shell. Breadth of shell:
ea. 230 um. Ref.: Nigrini and Moore (1979).
Order Nassellaria (Fig. 16.91-176) • Cephalospyris elathrobursa Haeckel (Fig. 16.102).
Solitary polycystines with a siliceous heteropolar Sub-ovoid shell with very delicate, thin wall and
shell, which can be represented by several fused spic- very small pores; main lateral spines extend as
ules only, by a D-shaped ring and associated spines, three-bladed or hollow and perforated feet. Shell
or by more elaborate, mono- or multilocular latticed height: ea. 200 um. Ref.: Petrushevskaya (1971a).
skeletons. With the exception of a few forms lacking • Lophospyris pentagona pentagona (Ehrenberg)
a well developed skeleton (Fig. 16.101, 16.119), the (Fig. 16.97) [=Lophospyris quadriforis, Lopho-
spyris pentagona]. Bars arising from sagittal, basal lady distributed circular pores and 3 massive, sim-
and frontal rings define large, very regular, polygo- ple or branched feet at the base; rudiments of tho-
nal pores. Skeletal bars and spines sharply 3-blad- racic lattice often present between feet. Shell
ed. Breadth of shell: ea 150 um. Goll (1976) erect- breadth: ca. 100 urn. Ref.: Petrushevskaya (1967).
ed the subspecies Lophospyris pentagona • Zygocircus productus (Hertwig) (Fig. 16.10 1).
hyperborea (Jorgensen) (not figured), which differs Pear-shaped or D-shaped, spiny, 3-bladed sagittal
from L. p. pentagona by the absence of tribladed ring. Major diameter: 90-140 um. Ref.:
lattice bars on the lateral surfaces of the shell. Ref.: Petrushevskaya (1971a).
Petrushevskaya (1971a), Goll (1976).
• Nephrospyris renilla Haeckel (Fig. 16.99) Family Plagoniidae (Fig. 16.104-124)
[=Nephrodictyum renilla]. The front and back of Skeletons restricted to a simple tri- or tetraxonic nas-
the sagittal ring produce branches that fork and sellarian spicule (Fig. 16.119), or a well developed
anastomose distally; the two sets of lattice plates system of main spines enclosed within a fully formed
thus formed are not interconnected laterally. cephalis (Fig. 3A, B). The degree of development of
Breadth ofshell: ea. 200 um. Ref.: Petrushevskaya the cephalis may vary from a few anastomosed bars
(1971a), Goll (1972). (Fig. 16.120, 16.124) to a well developed, latticed or
• Phormospyris stabilis scaphipes (Haeckel) (Fig. latticedlspongy chamber. Usually without postcephal-
16.93). Bi-lobulate, sagitally constricted, thin ic segments. In addition to several fairly well-defined
walled skeleton with 3 conspicuous feet protrud- species, the Plagoniidae comprise many probably
ing from basal ring. Breadth of shell: ea 80 um, related forms of obscure taxonomic status usually
Ref.: Goll (1976). cited under various generic names (see below). The
• Phormospyris stabilis stabilis (Goll) (Fig. classification of these forms needs detailed ad hoc
16.100). Cephalis thin-walled, bilobulate, separat- studies, for which reason many of them are provision-
ed by a conspicuous annular constriction from the ally lumped under the designation Plagoniidae group
conical thorax. Thorax open or closed. Both seg- in the present chapter.
ments with regular, circular pores. Breadth of • Antarctissa spp. group? (Fig. 3D, X; 16.104)
shell: ea 110 urn. Ref.: Goll (1976). [=?Antarctissa strelkovi, ?Antarctissa longa]. Shell
• Tholospyris anthophora (Haeckel) (Fig. 16.98). outline triangular to oval, cephalis partly sub-
Oval-shaped sagitally constricted skeleton with merged into thorax. Shell-wall thick. A. strelkovi
very heavy bars and circular pores. Shell breadth: (Fig. 3D) and A. longa differ from A. denticulata by
ea. 140 um. Ref.: Goll (1969, 1972). having thinner and spinier shell-walls. Shell height:
• Tholospyris ramosa (Haeckel) (Fig. 16.91) 100-160 um, Ref.: Petrushevskaya (1967).
[=Androspyris ramosa]. Shell pear-shaped, sagi- • Arachnocorys circumtexta Haeckel (Fig. 16.110).
tally constricted, with a well-developed galea and Spherical cephalis provided with numerous
3 feet whose distal ends can be spatulated or spines; those directed upwards are interconnected
forked. Shellheight: ea. 180 urn. Ref.: Takahashi by a spider web-like lattice of very thin bars; the
(1991). ones directed toward the base support an incipient
• Tholospyris spp. group (Fig. 16.103). Rings of thorax formed by a coarser lattice with irregular
variable size and form very cornmon in most pores, with several very large pores located in the
warm water materials; most of these are probably neck area. Overall shell height: ca. 140 um. Ref.:
juvenile representatives of various Spyridae. Petrushevskaya (1971a).
• Tholospyris tripodiscus Haeckel (Fig. 16.92). • Callimitra carolotae Haeckel (Fig. 16.123). The
Generally similar to T. ramosa, but with conspic- small, dome-shaped cephalis is provided with very
uously larger pores on both sides of the sagittal long apical (directed upwards), dorsal and main
ring and at the base of the galea; feet usually lateral spines (directed down and sideways) inter-
unbranched. Shell height: ea. 150 um, Ref.: connected by a delicate meshwork which forms 3
Petrushevskaya (1971a). basal plates and 3 lateral plates. Overall shell
• Triceraspyris antarctica (Haecker) (Fig. 16.94) height: ea. 200 um. Ref.: Haeckel (1887).
[=Triospyris antarctica, Phormospyris stabilis an- • Clathrocanium coarctatum Ehrenberg (Fig.
tarctica]. Heavy bilobulated cephalis with irregu- 16.114). Cephalis with a large, 3-bladed apical
---------------------------------~-----~ ------- -----

hom which may have lateral thread-like, anasto- and many long, thin spines. Thorax conical, its
mosing pojections. The dorsal and two main later- pores decreasing in size toward the base; some-
al spines, directed down and sideways, are joined times an incipient abdomen present. Overall shell
by narrow lattice plates which form a small thorax. height: ea. 150 urn. Ref.: Petrushevskaya (1971a).
Overall shell height: ea. 100 um, Ref.: • Neosemantis distephanus (Haeckel) (Fig.
Petrushevskaya (1971a). 16.120). Skeleton composed of an oval or pyri-
• Clathrocorys teuscheri Haeckel (Fig. 2G; form ring (fused main lateral spines) both poles of
16.112). Similar to C. coarctatum, except that the which are connected by a third bar (the apical
apical, dorsal and main lateral spines are joined by spine); from its base protrudes the dorsal spine.
a well-developed lattice. Overall shell height: ea. All skeletal elements very spiny. Major ring diam-
150-190 um. Ref.: Petrushevskaya (1971a). eter: 70-80 11m.Ref.: Petrushevskaya (1971a).
• Helotholus histricosa Jorgensen (Fig. 16.ll3). • Peromelissa phalacra (Haeckel) (Fig. 16.107)
Spiny, dome-shaped shell. Cephalis merging [=Psilomelissa phalacra, Lithomelissa monoce-
smoothly with thorax; thorax with large and irreg- ras]. Oval cephalis whose upper part is unperforat-
ular pores, without defined termination. Maximum ed or has very few, very small pores. The dorsal
shell width: ca. 100-120 um, Ref.: Petrushevskaya and main lateral spines emerge in the neck region
(1971a). as large, 3-bladed appendages. Overall shell
• Lampromitra coronata Haeckel (Fig. 16.115). height: ea. 100 um. Ref.: Petrushevskaya (197la).
Shell in the shape of a Chinese peasant hat. Thorax • Phormacantha hystrix (Jorgensen) (Fig. 16.111).
with subregular circular pores increasing in size Cephalis composed of a loose network of arches
slightly towards the base, with a well defined per- defining large, irregular pores. Thorax absent.
istome of a row of small pores and minute spines. Shell height: 60-70 11m. Ref.: Petrushevskaya
Shell diameter: ea. 200 um. Ref.: Petrushevskaya (1971a).
(1971a). • Plagonüdae group (Fig. 16.105). Includes many
• Lampromitra danaes (Haeckel) (Fig. 16.116) generally similar forms with a latticed cephalis and
[=Corocalyptra danaes]. Overall shell-shape sim- with or without a rudimentary thorax. The systema-
ilar to L. coronata. Pores on thorax regular, poly- tics of these sometimes extremely abundant forms
gonal, strongly increasing in size toward its base. is confused, and they are generally ignored in most
Rim of thorax represented by a very regular row of surveys. They include species cited under a variety
small, rectangular pores followed irnmediately by of generic names, such as Amphiplecta,
one of much larger pores. Shell diameter: ea. 180 Arachnocorallium, Arachnocorys, Ceratocyrtis,
um, Ref.: Haeckel (1887), as Clathrocyclas Dimelissa, Lophophaena, Lophophaenoma,
danaes. Micromelissa, Peromelissa, Psilomelissa, etc.
• Lampromitra quadricuspis Haeckel (Fig. • Pseudocubus obeliscus Haeckel (Fig. 16.122).
16.117). Generally similar to L. coronata, except Skeleton represented by the sharply 3-bladed
that cephalis is more elongate and pores on thorax edges of a 4-sided, truncated pyramid. Shell
are larger. Shell diameter: 120-350 um. Ref.: height: ea. 40 um. Ref.: Petrushevskaya (1971a).
Benson (1966). • Sethophormis aurelia Haeckel (Fig. 3F; 16.121).
• Lampromitra schultzei (Haeckel) (Fig. 16.118). Shell in the form of a Chinese peasant hat; cepha-
Similar to L. coronata, except that pores on thorax lis cupola-shaped; thorax with a very delicate,
are considerably larger and less regular. Peristome lace-like meshwork of irregular pores decreasing
with 2-3 rows of regularly aligned, small, subrec- in size toward the periphery and many radial sinu-
tangular pores. Shell diameter: ea. 100 um. Ref.: ous ribs. Shell diameter: 150-200 um. Ref.:
Boltovskoy and Riedel (1980). Petrushevskaya (1971a).
• Lophophaena butschlii (Haeckel) (Fig. 16.108). • Sethophormis rotula (Haeckel) (Fig. 16.124).
Elongated cephalis, the top of which is usually Central part of spider web-like skeleton is an hex-
unfinished, short conical thorax. Overall shell agonal ring whose vertices support 6 radiating
height: ea. 100 um. Ref.: Petrushevskaya (1971a). spines; 3 additional radial spines (dorsal and 2
• Lophophaena hispida (Ehrenberg) (Fig. 31; main lateral) merge in center of hexagon. All
16.109). Cephalis spherical, with very large pores spines produce rather regularly spaced anastomos-
i
""=--
ing lateral branches. Diameter of central hexagon: la-shaped thorax, provided with a large apical
ea. 40 um. Ref.: Petrushevskaya (1971a). horno Thorax with 3 small wings. Abdomen cylin-
• Tetraplecta pinigera Haeckel (Fig. 16.119). drical. Shell height: ea. 100-120 um. Ref.: Haeckel
Skeleton reduced to four equidistant 3-bladed (1887), as Pterocorys columba.
spines arising from a common central point; spines • Corocalyptra kruegeri Popofsky (Fig. 16.l36).
produce thin lateral braches which can anastomose Generally similar to C. columba; cephalis larger,
forming a delicate, irregular web. Length of each thorax and abdomen cylindrical. Shell height: ea.
spine: 25-30 um. Ref.: Haeckel (1887). 80 um. Ref.: Popofsky (1913).
• Trisulcus triacanthus Popofsky (Fig. 16.106). • Cycladophora davisiana (Ehrenberg) (Fig.
Small, sparsely perforated cephalis sitting on top 16.131) [=Theocalyptra davisiana, ?Artostrobus
of a conspicuously 3-lobulated thorax, lobes are jorgenseni]. Shell conical to campanulate.
most evident in the uppermost part, disappearing Cephalis subspherical, sparsely perforated, usually
gradually toward the base. Shell height: ea. 60 um, with 2 spines. Thorax conical to cylindrical, with
Ref.: Petrushevskaya (1971a). circular (proximally) to quadrate (distally) pores
increasing in size toward the base. Abdomen, when
Family Theoperidae (Fig. 16.125-151) present, flared out, wider than thorax, with quad-
Cephalis spherical or subspherical, relatively small, rate pores. Petrushevskaya (1967) described 3 sub-
often poreless or sparsely perforate. It usually bears an species of this form, of which C. davisiana
apical horno Internal spicule small and inconspicuous. (Ehrenberg) davisiana Petrushevskaya (Fig.
With one or more, sometimes up to over 10, usually 16.131 b) and C. davisiana (Ehr.) cornutoides
well-developed postcephalic segments. Generally, (Petrushevskaya) (Fig. 16.l31c) are often used in
cap- or helmet-shaped, or conical in overall outline. current literature (the last one is probably synony-
• Artostrobus annulatus (Bailey) (Fig. 16.125). mous with Artostrobus jorgenseni). Shell height:
Cephalis cup-shaped, poreless, with a thin apical ea. 100 um. Ref.: Riedel (1958), Petrushevskaya
horno Thorax cylindrical, with circular (proximal- (1967), Bjerklund and Ciesielski (1994).
ly) to subrectangular (distally) pores in transverse • Cyrtopera laguncula Haeckel (Fig. 30; 16.151)
rows, increasing in size toward the base. Shell [=Cyrtolagena laguncula, Stichopera pectinata).
height: up to 160 um. Ref.: Riedel (1958), Very typical multisegmented shell with conical
Petrushevskaya (1967). outline and clearly marked constrictions; last seg-
• Clathrocyclas cassiopeiae Haeckel (Fig. 16.126). ment may be open or closed. Shell height: 150-250
Small cephalis with a stout apical horn and many um, Ref.: Petrushevskaya (1971a).
smaller spines. Thorax large, campanulate, with • Dictyocephalus papillosus (Ehrenberg) (Fig.
irregular pores. Abdomen short, truncated, with 16.l35) [=Carpocanarium papillosum). Two-seg-
ragged, spiny termination. Shell height: 100-200 mented, thick-walled shell. Cephalis spherical.
um. Ref.: Haeckel (1887). Thorax oval, with circular, usually framed pores,
• Cornutella profunda Ehrenberg (Fig. 16.127). with 3 short wings, ending in a narrowed, poreless
Narrow, bilocular conical shell with very small peristome. Shell height: 70-90 um, Ref.:
subspherical pore1ess cephalis with or without api- Petrushevskaya (1967).
cal horno Pores on thorax circular, increasing in • Dictyophimus gracilipes Bailey (Fig. 16.142)
size toward the base. Shell height: 120-230 um, [=Pseudodictyophimus gracilipes, Dictyophimus
Ref.: Riedel (1958), Nigrini (1967). clevei). Cephalis partly submerged into thorax,
• Corocalyptra cervus (Ehrenberg) (Fig. 16.128). bears a large apical horno Thorax pyramidal or
Cephalis with a 1arge apical horn, often forked dis- conical, mouth open or closed, with 3 conspicuous
tally. Thorax large, campanulate, with regular, cir- legs (dorsal and lateral spines). Shell height (with-
cular to polygonal pores in transversal rows out feet): 55-90 um. Ref.: Petrushevskaya (1971a).
increasing in size distally. Abdomen restricted to a • Dictyophimus hirundo (Haeckel) (Fig. 16.140)
narrow brim with several rows of small pores. [=Pterocorys hirundo). Cephalis, globular, some-
Shell height: 100-200 um, Ref.: Benson (1966). times spiny, with a stout, 3-b1aded apical horno
• Corocalyptra columba (Haeckel) (Fig. 16.l32). Thorax truncate-conical to campanulate, spiny,
Subspherical cephalis partly submerged into cupo- with large circular pores and 3 ribs which extend
190 Radio1aria Polycystina
into mas sive, divergent feet. Highly variable spe- very small, spherical, hyaline. Thorax conical
cies (species group?). Shell height (without feet): proximally, flaring rapidly outward distally and, in
50-100 11m. Riedel (1958), Nigrini and Moore complete specimens, ending in a gently curved
(1979). brim. Diameter of fully-grown shells: up to 1 mm.
• Dictyophimus. infabricatus Nigrini (Fig. 3Q; Ref.: Benson (1966), Petrushevskaya (1971a).
16.143). Similar to D. hirundo, but shell is thinner • Lithopera bacca Ehrenberg (Fig. 16.129).
and pores are larger; cephalis usually wih 2 horns Spherical, often rough cephalis with an eccentri-
(apical and vertical spines), feet are smaller. Shell cally located apical horn partly submerged into an
height (without feet): 90-200 11m. Ref.: Nigrini oval thorax with closed mouth. Pores on thorax
and Moore (1979). regularly arranged. Shell height: 120-140 11m.
• Eucyrtidium acuminatum (Ehrenberg) (Fig. Ref.: Benson (1966).
16.13 7) [=? Eucyrtidium hexagonatum]. Small • Lithostrobus hexagonalis Haeckel (Fig. 16.141).
subspherical cephalis with apical horno Thorax Multisegmented, subconical shell with 5-9 joints
small, inflated, thick-walled. Abdomen and 4-5 with well marked strictures and angular shoulders.
post-abdominal segments thin-walled, with pores Pores very regular, subpolygonal, in transverse
arranged in longitudinal rows. Shell height: 120- rows. Shell height: 120-300 11m. Ref.: Benson
200 11m. Eucyrtidium hexagonatum Haeckel is (1966).
closely related to this species; according to Nigrini • Perypiramis circumtexta Haeckel (Fig. 16.130)
(1967) it can be distinguished from E. acuminatum [=?Plectopyramis dodecomma]. Cephalis very
by the sharp change in contour at the lumbar stric- small, ovate, hyaline. Thorax large, conical, with
ture. Ref.: Nigrini and Moore (1979). ea. 10 well defined longitudinal rows of subquad-
• Eucyrtidium anomalum (Haeckel) (Fig. 16.138). rate pores rapidly increasing in size toward the
Cephalis spherical, partly submerged into the large, base. Pores are usually not aligned transversely.
assymetric, conical, thin-walled thorax. Abdomen The form described as Plectopyramis dodecomma
and 2-3 postabdominal segments thin-walled, with Haeckel differs from P. circumtexta in that pores
pores in longitudinal rows. Maximum shell width: are aligned transversely as well as longitudally.
80-100 ITUll. Ref.: Petrushevskaya (1971a). Bathropyramis woodringi is also similar to the
• Eucyrtidium hexastichum (Haeckel) (Fig. above, but has thicker longitudinal bars and aligned
16.139). Cylindrical or conical shell with up to 9- horizontal (transverse) bars. Shell height: 100-300
10 segments, thin- walled, with pores arranged in um. Ref.: Riedel (1958), Nigrini and Moore (1979).
transversal rows. Maximum shell width: 65-80 • Pterocanium elegans (Haeckel) (Fig. 16.147).
11m.Ref.: Petrushevskaya (1971a). Cephalis relatively small, with two large horns
• Lipmanella bombus (Haeckel) (Fig. 16.133) (apical and vertical spines). Thorax a large, thin-
[=Dictyoceras cf. pyramidale]. Very characteristic walled, 3-sided pyramid with small, very regular
thorax where the dorsal and main lateral spines, subcircular pores and thin bars; edges of pyramid
which project down and sideways, are surmounted continue as 3 strong 3-bladed legs. Abdomen
by latticed keels; these keels define the 3 slightly cylindrical, may be rudimentary. Shell height
concave sides of the pyrarnidal thorax. Maximum (without horns): 180-260 11m. Ref.: Benson
shell width: 80-150 11m. Ref.: Benson (1966), (1966), as Pterocanium cf. elegans.
Petrushevskaya (1971a). • Pterocanium praetextum (Ehrenberg) group?
• Lipmanella dictyoceras (Haeckel) (Fig. 3R, (Fig. 16.146). Similar to P. trilobum, from which
16.134) [=Lipmanella virchowii, Dictyoceras vir- it sometimes is difficuIt to separate; differs by
chowii, Dictyoceras neglectum]. Cephalis large, having a thorax with more pronounced, angular
hemispherical, with a conspicuous apical horno shoulders, and often a better developed abdomen.
Thorax thin-walled, conical-inflated, with 3 wings Shell height (without horn and feet): 100-120 11m.
(dorsal and main lateral spines). Abdomen absent Ref.: Petrushevskaya (1971a).
or rudimentary. Shell height: 90-130 11m. Ref.: • Pterocanium trilobum (Haeckel) (Fig. 3H;
Petrushevskaya (1971a). 16.145). Cephalis relatively small, with a stout
• Litharachnium tentorium Haeckel (Fig. 16.148). apical horno Thorax an inflated tetrahedron with
Two-segmented, very characteristic shell. Cephalis regularly arranged circular pores, with 3 ribs
!
I
'1
extending into stout, 3-bladed, slightly curved apical horno Thorax campanulate, with circular,
feet. Abdomen, when present, rudimentary. Shell regular pores arranged hexagonally; peristome
height (without horn and feet): 100-120 um. Ref.: distinct, constricted, may bear terminal teeth.
Nigrini and Moore (1979). Abdomen absent. Maximum breadth of thorax:
• Pterocyrtidium dogieli Petrushevskaya (Fig. 90-140 um. Ref.: Nigrini and Moore (1979).
16.144) [=Sethoconus dogieli]. Cephalis herni- • Anthocyrtidium zanguebaricum (Ehrenberg)
spherical, almost poreless, thick-walled. Thorax (Fig. 16.153). Similar to A. ophirense, but apical
cylindrical, thorny, with very large, irregular, sub- horn is smaller, thorax less campanulate, narrow-
circular pores. Shell height: ca. 140 um, Ref.: er, and terminal teeth smaller or absent. Maximum
Petrushevskaya (1971a). breadth of thorax: 60-80 um. Ref.: Nigrini and
• Sethoconus anthocyrtis Haeckel (Fig. 16.150) Moore (1979).
[=Conarachnium polyacanthum, Lophocorys • Lamprocyclas maritalis group? Haeckel (Fig. 3P;
polyacantha]. Cephalis spherical, spiny. Thorax 16.158). Massive shell; cephalis oval, with a large
c1early differentiated from cephalis, very large, apical horno Thorax cupola-shaped, with regular,
conical, spiny, with large, regular, subpolygonal circular, framed pores. Abdomen separated by a
pores approximately in longitudinal rows; termi- conspicuous lumbar stricture, inflated, with larger,
nation ragged. Shell height: 200-300 urn. Ref.: regularly arranged, framed pores, usually with a
Haeckel (1887). well-defined poteless peristome with teeth. L. m.
• Theopilium tricostatum (Haeckel) (Fig. 16.149) maritalis Haeckel differs from L. m. Haeckel poly-
[=?Theocalyptra gegenbauri]. Chinese peasant pora Nigrini in having a less inflated abdomen.
hat-like shell with a small cephalis. Thorax wide- Maximum breadth of abdomen: 100-140 um. Ref.:
ly open, with small, regularly arranged pores, cir- Nigrini and Moore (1979).
cular proximally and becoming larger and more • Lamprocyrtis hannai Campbell and Clark (Fig.
polygonal distally, with 3 conspicuous symmetri- 16.154) [=Theoconus junionis, Lamprocyclas
cal ribs (dorsal and main lateral spines). junionis, ?Androcyclas gamphonycha]. Cephalis
Abdominal brim flat, with 5-8 rows of very regu- elongate, with a large 3-bladed apical horno
lar, quadrangular pores. Shell diameter: 130-300 Thorax campanulate, thick-walled, with subregu-
um. Ref.: Haeckel (1887), Benson (1966). lar, circular pores. Abdomen truncate-conical,
with large, subregular, circular pores and usually
Family Carpocaniidae (Fig. 16.170) with terminal and/or subterminal teeth. Maximum
The small, rudimentary cephalis is usually totally breadth ofabdomen: 90-150 urn. Ref.: Nigrini and
irnmersed in the large and well-developed thorax (Fig. Moore (1979).
3E). Abdomen absent or rudimentary. • Lamprocyrtis nigriniae (Caulet) (Fig. 16.157)
• Carpocanium spp. (Fig. 3E, 16.170) [=Carpo- [=Conarachnium nigriniae, Lamprocyrtis haysi].
canistrum spp.]. Shell outline subspherical to oval. Cephalis elongated, usually open proximally, with
Cephalis indistingishable from thorax, inc1uded a large 3-bladed horno Thorax campanulate, thin-
within its upper parto Peristome poreless, smooth walled, with large, subregular, circular pores
or provided with teeth. Height of shell: 80-130 increasing in size distally; peristome absent or
urn. Ref.: Nigrini and Moore (1979). weakly developed. No abdomen. Maximum
breadth of thorax:ca. 90 urn. Ref.: Nigrini and
Family Pterocorythidae (Fig. 16.152-162) Moore (1979).
Cephalis large, divided into 3 lobes by two lateral fur- • Pterocorys hertwigii (Haeckel) (Fig. 16.155)
rows directed obliquely and downward from the api- [=Theoconus hertwigii, Phormocyrtis fatuosa].
cal spine to the base of the cephalis. The upper Cephalis oval, horned. Thorax campanulate.
unpaired lobe is located above the two smaller paired Abdomen conical, with ragged termination.
ones (Fig. 3K, L); these basal paired lobes are not Thorax and abdomen thin-walled, with regular cir-
always conspicuous. Many pterocorythids are two or cular pores in longitudinal rows, with several con-
three-segmented, lacking postabdominal segments. tinuous or interrupted longitudinal poreless ribs.
• Anthocyrtidium ophirense (Ehrenberg) (Fig. Total shell height (exc1uding horn): 120-200 um.
16.152). Cephalis elongate with a large 3-bladed Ref.: Caulet and Nigrini (1988).
• Pterocorys minythorax (Nigrini) (Fig. 16.156) last ones, are arranged in clearly defined transverse
[=Theoconus minythorax]. Cephalis subspherical, rows (Fig. 3S). Usually elongated, multisegmented
homed. Thorax small, campanulate. Abdomen forms.
comparatively large, cylindrical, slightly flared, • Botryostrobus aquilonaris (Bailey) (Fig. 16.164)
with ragged termination. Pores on thorax and [=Lithocampe aquilonaris, Botryostrobus tumidu-
abdomen regular, circular. Total shell height lum]. Very thick-walled, spindle-shaped skeleton
(excluding hom): 120-200 um. Ref.: Caulet and with 6 poorly defmed joints, of which the fourth is
Nigrini (1988). the broadest. Poreless peristome usually present.
• Pterocorys zancleus (Muller) (Fig. 16.159) Shell height: 100-150 um. Ref.: Nigrini and
[=Theoconus zancleus, ?Pterocorys sabae, Moore (1979).
?Pterocorys campanula]. Cephalis subcircular, • Botryostrobus aurituslaustralis (Ehrenberg) (Fig.
homed. Thorax a truncated cone, with 3 small 3J; 16.168). [=Lithostrobus seriatus]. Multi-
wings. Abdomen cylindrical or conical, slightly segmented cylindrical shell of variable wall-thick-
flared. Total shell height (excluding hom): 100-230 ness. Cephalis subspherical, apical tube and apical
um. Ref.: Benson (1966), Petrushevskaya (1971a). spine usually visible. Thorax and post-thoracic seg-
• Pteroscenium pinnatum Haeckel (Fig. 16.162) ments inflated, separated by conspicuous strictures,
[=Verticillata hexacantha]. Campanulate one-seg- with 3-5 transverse rows of pores. Shell height:
mented shell with delicate lattice of small, circular 110-200 um. Ref.: Boltovskoy and Vrba (1989).
pores. Top of cephalis extends into a large, pro xi- • Phormostichoartus corbula (Harting) (Fig.
mally perforated, 3-bladed hom; base is prolonged 16.163) [=Lithocampe multiseriata, Siphocampe
into 3 stout, perforated, slightly bent feet. Total corbula]. Four-segmented shells. Cephalis and
shell height (including hom and feet): 220-280 thorax fused, thick-walled. Abdomen short, cylin-
um. Cephalis does not seem to be agree with the drical. Last segment usually 3-4 times longer than
family-level diagnosis; probably a plagoniid. Ref.: previous, both with circular pores arranged regu-
Haeckel (1887), Benson (1966). larly in transverse rows. Shell height: 130-170 um,
• Stichopilium bicorne Haeckel (Fig. 16.161). Ref.: Nigrini and Moore (1979).
Cephalis cap-shaped, with 2 stout, 3-bladed homs. • Siphocampe arachnea (Ehrenberg) (Fig. 16.167)
Thorax pyramidal (proximally) to cylindrical (dis- [=Lithomitra arachnea]. Cylindrical shell with 3-8
tally) , with 3 conspicuous wings. One or two joints, with 4 transverse rows of small, circular
cylindrical post-thoracic segments. Thorax and pores on the thorax and one on each subsequent
subsequent joints with small, circular, regularly segment. Surface covered with a typical net of
arranged pores. Although this species has tradi- irregular longitudinal and transverse ridges. Shell
tionally been ascribed to the Pterocorythidae, it's height: ea. 60-90 um. Ref.: Petrushevskaya (1967).
cephalis does not seem to be divided into lobes, • Siphocampe lineata (Ehrenberg) (Fig. 16.169)
and is therefore probably a theoperid. Ref.: [=Lithomitra lineata, Lithomitra nodosaria,
Haeckel (1887), Benson (1966). Siphocampe nodosaria]. Very similar to S. arach-
• Theocorythium trachelium (Ehrenberg) (Fig. nea, except that surface omamentation is restrict-
16.160) [=Calocyclas amicae, Lamprocyclas tra- ed to longitudinal, sinuous ridges. Shell height: ea.
che/ius, Lamprocyclas cranoides]. Cephalis spheri- 80-11 O um. Ref.: Petrushevskaya (1967).
cal, with a prominent, 3-bladed apical hom. Thorax • Spirocyrtis scalaris Haeckel group? (Fig. 16.166)
campanulate, inflated. Abdomen cylindrical, with a [=Spyrocyrtis scalaris/cornutella]. Thin-walled
slight medial constriction; peristome differentiated, shell composed of up to 8 joints with a typically
toothed. Pores on post-cephalic segments circular, angular outline increasing step-wise in width
regularly arranged. Total shell height (without toward the base. Pores on post-thoracic segments
hom): 150-200 um. Ref.: Petrushevskaya (1971a). quadrangular, arranged in very regular transverse
rows. Shell height: 100-200 um. Ref.: Nigrini
Family Artostrobiidae (Fig. 16.163-169) (1967), Petrushevskaya (1971a).
Spherical or subspherical cephalis, usually with an • Tricolocampe cylindrica Haeckel (Fig. 16.165)
apical tube directed obliquely upwards (Fig. 3J). The [=Siphocampium cy/indrica]. Cephalis and thorax
pores on all postcephalic segments, or at least on the fused, separated from rest of shell by a conspicu-
l
ous lumbar constnction. Abdomen and postab- one, and approximately equal in size. Thorax cylin-
dominal section cylindrical to spindle-shaped, drical, with a closed mouth in fully-grown speci-
without external constrictions, with several very mens. Shell thick-walled, surface rough.· Shell
regularly arranged transverse rows of small, circu- height: 110-160 um. Ref. : Petrushevskaya (1965).
lar pores. Shell height: ea. 90 um. Ref.: Benson
(1966), Pterushevskaya (1971a).
Acknowledgments
Family Cannobotryidae (Fig. 16.171-176)
Cephalis large, with several asymmetrical lobes Thanks are due to the many colleagues and publishers
(sometimes appearing as irregular bulges) (Fig. 3M). which kindly allowed using their published and
Mostly 2-segmented forms (cephalis and thorax), but unpublished illustrations for this work, and especially
sometimes with post-thoracic segments. to Neil Swanberg, Petrus van de Paverd, Amy
• Acrobotrys spp. (Fig. 3M, N; 16.171). Several Weinheimer and Cathy Nigrini for providing several
poorly defined Cannobotryidae of variable con- of the photographs included. Critical reviews by
struction (e.g., Acrobotrys sp. A, B, C in Stanley K1ing, Cathy Nigrini, Annika Sanfilippo and
Petrushevskaya, 1965; Acrobotrys cf. disolenia in Arny Weinheimer, as well as comments by Roger
Benson, 1966; Acrobotrys sp. A and B, Anderson, William Riedel and Luis O'Dogherty,
Cannobotryid sp. A in Boltovskoy and Riedel, were helpful for improving earlier vesions of the ma-
1987; etc.). nuscript. This work was partly financed by grants
• Botryocephalina armata Petrushevskaya (Fig. from the University of Buenos Aires (UBA EX-040),
16.173). Main part of cephalis represented by two and from the Consejo Nacional de Investigaciones
laterally fused hemispherical chambers of almost Científicas y Técnicas (CONlCET PID-BID 0366).
equal size (cephalic and antecephalic); with a long
apical spine. Thorax cylindrical, its distal section
can be poreless. Shell height: ea. 50 um. Ref.: Suggested readings
Petrushevskaya (1965).
• Botryocyrtis scutum (Harting) (Fig. 16.172) Anderson O.R. 1983. Radiolaria. Springer Verlag,
[=?Botryocyrtis caput-serpentis, ?Botryocyrtis New York, pp. 1-355. An excellent review o/
quinaria]. Large, multilobed cephalis. Thorax radiolarian biology and cytology.
very short, cylindrical. Abdomen longer. Anderson O.R., Nigrini C.A., Boltovskoy D.,
Sometimes one post-abdominal segment. Entire Swanberg N.R., Takahashi K. In press. Radiolaria.
shell, and especially its upper section, enclosed in In "An illustrated guide to the Protozoa" Second
a thick, spongy mantle. Shell height: 80-130 um. Edition, (1. Lee, ed.). A short review o/ po lycystin e
Ref.: Nigrini and Moore (1979). classification, with examples (identification keys
• Botryopyle dictyocephalus Haeckel (Fig. 16.174). andjigures) of the most widespread genera.
Cephalis chiefly composed of a large, ovoid, thin- Benson R.N. 1966. Recent Radiolaria from the Gulf
walled antecephalic chamber, and a much smaller, of California. Ph. D. Diss., Univ. Minnesota, pp.
spherical, thick-walled eucephalic chamber. 1-577. Contains some of the most detailed descrip-
Cervical constriction absent. Thorax cylindrical, tions and many photographs o/ 158 tropical and
with ragged termination or distally narrowing into subtropical polycystine species.
a short, poreless tube. Shell height: ea. 100 um. Boltovskoy D. 1981. Radiolaria. In "Atlas del
Ref.: Petrushevskaya (1965). Zooplancton del Atlántico Sudoccidental y méto-
• Centrobotrys thermophila Petrushevskaya (Fig. dos de trabajo con el zooplancton marino" (D.
16.175). Shell is a laterally compressed, very thin Boltovskoy, ed.), Publico Esp. Inst. Nac. Inv.
walled, pored cone, within which the spherical, Desarrollo Pesq., Mar del Plata, pp. 261-316.
thick-walled eucephalic chamber is enclosed. Shell General review o/ the systematics and distribution
height: ea. 100 urn. Ref.: Petrushevskaya (1965). o/ Recent polycystines from the Southwestern
• Saccospyris antarctica Haecker (Fig. 16.176). Atlantic Ocean. Inc/udes brief remarks on
Cephalis tri-lobulated, with the cephalic and antece- radiolarian biology, methods o/ research, histoy
phalic chambers much larger than the postcephalic o/ study, etc. (in Spanish).
COLLODARIA
Solitary species
, Thalassocampe
maxima
Thalassoxanthium
medusinum
Thalassoxanthium
cervicorne
(j nset: detai I
of spicule)
Colonial species
16.6a
16.9b
Collozoum inerme 16.7* Collozoum 16.8 entire colony
serprentinum Collozoum Rhaphidozoum
longiforme neapolitanum
1~;Ytf'
'<;¡'
\-h
¡ ,~., -', -.
16.10* '. ,/ .
Sphaerozoum, EJ,emmatum
single individual
COLLOSPHAERI DAE
Externa surface without spines or tubes External surface spiny
or thorny ,
16.13b
Collosphaera tuberosa 16.14b
16.17*
16.16b* Buccinosphaera invaginata

Acrosphaera
Tribonosphaera murrayana
centripetalis ----1
External surface with 3-4 to many tubes
Siphonosphaera
martensi
Solenosphaera
chierchiae
Solenos
! (
,------
196 Radiolaria Polycystina /7'.., /\
'.J
ACTI NOMMI DAE
Outermost shell latticed
16.25a 16.25b \\
Acanthosphaera actinota • 16.26b \
.f 16.27a
16.26a Acanthosphaera dodecastyla
16.28a* 16.28b*
Acanthosphaera pinchuda
Druppatractus
irregularis
16.31b
Hexacontium laevigatum 16.31 c* Cromyechinus
antarctica
16.32a 16.32b
Hexacontium armatum/hostile group 16.34a Hexacontium

aristarchi
1I
1
I
I~ 16.34b*
Stylosphaera
melpomene 16.36
Stylatractus spp. group
·._._------_._---~--------------
f
ACTI NOMMI DAE
16.37a 16.37b 16.38a

Thecos haera inermis
Actinomma leptodermum
16.39
Carposphaera
acanthophora
16.42* !{ '\
Cladococcus
megaceros
.~~~ ./ ~
-;~l'
• ~ l. \
16.43b '/"!. \ ;
Heliosome echinaster
!Jl
16.44a 16.44b
Actinomma 501
ACTI NOMMI DAE

. 16.48a Actinomtne
Aetínomma antaretíeum group? areadophorum
Outermost shell spongy
Styptosphaera
spumaeea Spongoplegma Plegmosphaera
rugosa paehyplegma
Plegmosphaera
exodíetyon
" 16.54
Plegmosphaera
entodíetyon
detailof "
central
Spongosphaera par! of
Spongodíetyon streptaea nth a shell
spongíosum
\
SPONGODISCI DAE
Circular outline
16.61 a
I 16.61 b
Spongotrochus
gracia lis
16.64
Spongopyle Spongodiscus
setosa resurgens
Quadrate outline
Spongaster tetras
irregularis
16.67 ~-:.< .~_ .

.••.. ,. . .•.• :>"
Spongaster tetras tetres
SPONGODISCIDAE COCCODISCI DAE

Elongated outline
16.73 \\
Spongocore
cylindrica
16.72a 16.72b Spongurus Amphirhopalum
16.77b
Spongurus pylomaticus spp. group? ypsilon cross
section
of Didymocyrtis
middle tetrathalamus
PHACODISCI DAE
PYLONIIDAE
Heliodiscus asteriscus 16.78b* ..

16.80a Octopyle stenozona group? 16.80b
detail of medullary shell,

cortical shell broken
16.79a Sethodiscus macrococcus 16.79b
16.81a 16.81 b
Pylolena armata
THOLONIIDAE
16.82b
Oipylissa
Cubotholus bensoni
spp. group 16.83b*
LITHELIIDAE
~
Larcopy/e butsch/ii 16.86a
Larcospira quadrangu/a
16.88a
16.87b
Lithe/ius
Lithe/ius minor
nauti/oides
group?
16.87c
16.90b*
16.90a
Th otosp ira Py/ospira octopy/e
cervicornis
202 Radiolaria Polycystina ...
l
)
1- !
/
SPYRIDAE
16.93a Phormospyris stabilis 16.93b

scaphipes
Tholospyris tripodiscus
16.94c
16.95a 16.95b
Amphispyris
teticule ta
Triceraspyris antarctica
,--------------------------------,
'----__ ----'-__ ---'---'----'- =-- __ --'----__ ---= ---' 16.1 00
16.99
Phormospyris
stabilis
stabilis
16.102
16.101a
Cephalospyris 16.103a
Zygocircus productus cfathrobursa Tholos
T Radiolaria Polycystina 203
PLAGONIIDAE
• ~í
16.10Sd 16.105e
Plagoniidae group
Trisulcus.triacanthus
16.108a
Arachnocorys
circumtexta
'"' 16.112a Clathrocorys teuscheri

Phormacantha hystrix r-----------------
16.113b
Clathrocanium coarctatum
Helotholus histricosa
204 Radiolaria Polycystina l}
/
PLAGONIIDAE
. Viewedir~m below, showing

Lemptomitre eoronata tetralobed central capsule
16.118b
Lampromitra
I sehultzei
II
16.122a
Tetrapleeta
pinigera Neosemantis distephanus
Pseudoeubus obeliseus
16.123d
16.123b Callimitra ea ro lotae
I
l·
II
THEOPERIDAE
16.125b
000°
00 O
OOOe¡
0000
0000
... -
-"")-
Artostrobus annulatus
Corocalyptra cervus
16.131c Lithopera
16.131b bacca Peripyramis circumtexta
:;";~r'::·":<.·;r-
~:::Il(~;;'.:.
\\
Cycladophora davisiana ¡[
16.136a
16.
135a
Lipmanella dictyoceras Oictyocephalus papillosus Corocalyptra kruegeri

THEOPERIDAE
16.139a
l'
Oictyophimus Pterocyrtidium dogieli

infabricatus
16.145b
16.147
Pterocanium praetextum Pterocanium

Pterocanium elegans
trilobum
j \-f )
THEOPERIDAE <:>
( !/
16.148b
Litharachnium tentorium
16.149c
Theopilium tricostatum
16.150b*
16.151a
~
Cyrtopera laguncula
¿
PTEROCORYTH I DAE
16. 16.154
153a
~
Lamprocyrtis
Anthocyrtidium zanguebaricum hannai
16.
157a
16.
158a
16.162c*
(
;
) '1 )
....1 "
ARTOSTROBIIDAE
Phormostichoartus corbu/a
I
16.16Sb
Tricolocsmpe
Spirocyrtis sca/aris 16.167a cy/indrica
Siphocampe arachnea
16.169a
Botryostrobus auritus/austra/is 16.168d

Siphocampe,
/ineata
CARPOCAN 11DAE
16.170a 16.170b 16.170c 16.170d 16.170e

Carpocanium spp. group?
CAN NOBOTRYI DAE
16.172d
Botryocyrtis scutum Botryocephal ina
armata
16.174b 16.174c
16.171e 16.171f
Acrobotrys spp.
16.175a
16.175b*
•••.. ..;".;.. .•..•• -'- ..r..:.;. ••.• 16.176a

Centrobotrys thermophila Saccospyris antarctica
Boltovskoy D. 1994. The sedimentary record of Arch. Mus. Nation. Hist. Natur., Paris, Ser. 7, 7:1-
pelagic biogeography. Progr. Oceanogr., 34: 135- 134. A major contribution toward a new classifi-
160. Review of the differences between microzoo- cation o/ spherical spumellarians based on cyto-
planktonic geographic patterns in the plankton logical traits (in French).
and in the surface sediments, and o/ the distorting Kling S.A. 1978. Radiolaria. In "Introduction to marine
mechanisms involved. rnicropaleontology" (B.U. Haq, A. Boersma, eds.),
Campbell A.S. 1954. Subclass Radiolaria. In EIsevier, New York, pp. 203-244. A concise and
"Treatise of invertebrate paleontology. Protista 3" very good review o/ all aspects o/ polycystine
(R.c. Moore, ed.), Geol. Soc.Amer. and Kansas knowledge, with emphasis on their applications to
Univ. Press, Lawrence, pp. DII-DI63. A sum- stratigraphy and paleoceanography.
mary o/ Haeckel 's (1887) system with minor revi- Lombari G., Boden G. 1985. Modem global radiolar-
sions and additions. ian distributions. Cushman Found. Foram. Res.,
Caron D.A., Swanberg N.R. 1990. The ecology of Spec. Pub. 16A, Washington, D.C. Charts o/ the
planktonic sarcodines. Aquatic Sci., CRC Press, relative (percentage) distribution o/ 70 polycys-
Boca Raton, 3:147-180. Useful summary o/ tine species in the surface sediments o/ the World
radiolarian biology; includes information on dis- Ocean, based on data o/ the CLIMAP project.
tribution and abundance, feeding, growth and Nigrini c.A., Moore T.C. 1979. A guide to Modem
reproduction and symbiosis o/ planktonic sarco- Radiolaria. Cushman Found. Foram. Res., Spec.
dines (Amoebae, Foraminifera, Radiolaria, Pub. 16, Washington, D.C., pp. SI-S142 + Nl-
Acantharia and Heliozoa). N106. A nicely illustrated catalogue o/ ea. 100
Haeckel E. 1887. Report on Radiolaria collected by Recent polycystine species, including descriptions
H.M.S. Challenger during the years 1873-1876. and remarks on their geographic distribution in
Rep. Sci. Res. Voyage H.M.S. Challenger 1873- surface sediments. This work summarizes current
76, 18:1-1803. Cornerstone work on radiolarian interpretation and names for the most common
classification, including the descriptions 0/2785 po/ycystine taxa.
new species with artistically superb illustrations. Petrushevskaya M.G. 1967. Radiolyarii otryadov
Hollande A., Enjumet M. 1960. Cytologie, évolution Spumellaria i Nassellaria Antarkticheskoi oblasti
et systématique des Sphaeroidés (Radiolaires). (po materialam Sovetskoi Antarkticheskoi Ekspe-
•••
Fig. 16. Illustrations of polyeystine speeies. Asterisks denote figures given with the original speeies deseription. Figure sourees: from Benson
(1966): 31b; from Bjerklund and Gol! (1986), with kind permission from Elsevier Seienee: 129; from Boltovskoy and Jankilevieh (1985), eour-
tesy ofGauthier-Vil!ar Editeur: 126a; from Boltovskoy and Riedel (1980): 28a, 28b, 82a, 110a; from Boltovskoy and Riedel (1987): 58a, 101a,
103a, 108a, lila, 161a, 170a, 172a, 173a, 176a; from Boltovskoy and Vrba (1988): 60a, 62a; frorn Boltovskoy (1981e): 13b, 16a, 29b, 37b,
79b; from Boltovskoy (1987): 56b; from Caulet (1971): 157b; frorn Cheng and Yeh (1989): 96a; from Durnitriea (1988): 83a, 83b; from Gol!
(1972): 97a, 98; from Gol! (1976): 93a, 93b, 94a, 100; from Gol! (1980): 157a; from Haeekel (1887): 1-5, 6a, 7,10, 16b, 17, 18e-e, 19b, 20,
31e, 34b, 40b, 47, 56a, 58b, 78b, 84b, 84e, 90b, 91, 92b, 112b, 115b, 116b, 117b, 119, 123d, 126b, 149b, 150b, 161b, 162b, 162e; from Hol!ande
and Enjurnet (1960): 42, 43b, 44b, 46b, 54, 55; from Kling (1977), with kind permission from Elsevier Seienee: 113b, 156a; from Kling (1978):
86a; from Matsuoka (1992), with kind permission from Elsevier Seienee: 69; from Matsuoka (1993): 166e; from Nigrini and Moore (1979):
19a, 22a; from Nigrini (1967), by eourtesy ofthe Seripps lnstitution ofOeeanography: 48a; from Nigrini (1968): 170e; from Nigrini (1971), by
eourtesy ofCambridge University Press: 71; from Paverd (1995): 14b, 15b, 18b, 57a, 90a, 136a, 141b, 147, 148b, 150a, 153b, 155b; from
Petrushevskaya (1965): 171b, 171f, 173b, 174e, 175b, 176e; from Petrushevskaya (1967): 2ge, 35b, 60e, 62e, 65e, 72b, 82b, 88b, 89, 94b, 94e,
104e, 131b, 13le, 135b, 163b, 164e, 165b, 168d, 167b, 169b; from Petrushevskaya (1971a): 95b, 96b, 97b, IOlb, 102, 103b, 103e, 106b, 107b,
108b, rose, 110b, ll1b, 113e, 114b, 120, 122b, 124b, 125b, 127b, 128b, 130b, 133b, 137e, 138b, 139b, 142b, 144b, 145e, 146b, 155e, 159b,
170d, 170e; from Petrushevskaya (1971e): 148a; from Petrushevskaya (1981): 172e, 172d; from Popofsky (1913): 25b, 26b, 41, 77a; from
Riedel (1958), by eourtesy ofthe Mawson Antaretie Col!eetion, University of Adelaide, Australia: 48e, 72a, 140b; from Sehaaf(1981): 123e;
from Strelkov and Reshetnjak (1971): 6b, 9, 21, 22b, 23b; Courtesy ofNeil Swanberg (from Swanberg, 1979): 6e, 8, 11, 12; from Takahashi
(1981): 40a; from Weinheimer (1994): 87b, 113a; Original: 13a, 14a, 15a, 18a, 23a, 24a, 24b, 25a, 26a, 27a, 27b, 29a, 30, 31a, 32a, 32b, 33a,
33b, 34a, 35a, 36, 37a, 38a, 38b, 39, 43a, 44a, 45, 46a, 48b, 49-53, 57b, 59a-e, 60b, 61a, 62b, 63, 64, 65a, 65b, 66-68, 70a-e, 73-76, 77b, 78a,
79a, 80a, 80b, 81a-e, 84a, 85a, 85b, 86b, 87a, 87e, 88a, 88e, 92a, 95a, 99, 104a, 104b, 105a-e, 106a, 107a, 109a, 109b, 112a, 114a, 115a, 116a,
116e, 117a, 118a, 118b, 121, 122a, 123a, 123b, 124a, 125a, 127a, 128a, 130a, 131a, 132a, 132b, 133a, 134, 135a, 136b, 137a, 137b, 138a, 139a,
140a, 141a, 142a, 143, l44a, 145a, 145b, 146a, 148e, 149a, 14ge, 151a, 152a, 152b, 153a, 154, 155a, 156b, 158a-e, 159a, 15ge, 160, 162a, 163a,
164a, 164b, 165a, 166a, 166b, 167a, 168a-e, 169a, 170b, 171a, 171e, 171d, 171e; 172b, 174a, 174b, 175a, 176b.
ditzii) [Radiolarians of the orders Spumellaria and Riedel W.R. 1971. Systematic c1assification of poly-
Nassellaria from the Antarctic region (on the basis cystine Radiolaria. In "The micropalaeontology of
of material s of the Soviet Antarctic Expeditions]. oceans" (W.R. Riedel, B.M. Funnell,eds.),
Issledovaniya Fauny Morei 9(12), 3:5-186. Cambridge Univ. Press., Cambridge, pp. 649-660.
Descriptions and illustrations of most Antarctic Outline of most widely used polycystine suprage-
radiolarians (in Russian, translated into English neric c/assification.
by WR. Riedel). Riedel W.R., Foreman H.P. 1995. Catalogue ofpoly-
Petrushevskaya M.G. 1971. Radiolyarii Nassellaria v cystine radiolaria. National Technical Information
planktone Mirovogo Okeana [Nassellarian Service, NTIS ID Number PB281000/LC. A cata-
radiolarians in the plankton of the World Ocean]. logue of the polycystine species described up to
Issledovaniya Fauny Morei 9(17):5-186. Very well 1930; icludes approx. 20,000 sheets in microfiche
illustrated review of practically all extant nassel- format containing species illustrations and
larian species; includes some data on ecology and descriptions (price: 1158 US dollars).
distribution in the World Ocean (in Russian, Sanfilippo A, Westbeg-Smith M.J., Riedel W.R. 1985.
translated into English by WR. Riedel). Cenozoic Radiolaria. In "Plankton stratigraphy"
Petrushevskaya M.G. 1981. Radiolyarii otryada (H.M. Bolli, J.B. Sunders, K. Perch-Nielsen, eds.),
Nassellaria Mirovogo Okeana [Radiolarians ofthe Cambridge Univ. Press, Cambridge, pp. 631-712. A
order Nassellaria from the World Ocean]. Nauka, detailed guide for Cenozoic stratigraphy based on
Leningrad, pp. 1-405. A catalogue and new c/as- polycystines. Inc/udes illustrations and descriptions
sification scheme of all extant and extinct nassel- of all diagnostic species, as well as ancillary data
larians (in Russian). on techniques and historie perspectives.
Petrushevskaya M.G. 1986. Radiolyarevyi analiz Strelkov AA, Reshetnjak V.V. 1971. Koloniyalnie
[Radiolarian analysis]. Nauka, Leningrad, pp. 1- radiolyarii Spumellaria mirovogo okeana [Colonial
200. A comprehensive guide covering all aspects spumellarian radiolarians from the World Ocean].
of the study of Recent and fossil Radiolaria (in Issledovaniya Fauny Morei 9(17):295-373.
Russian). Illustrated descriptions and distribution data for
most colonial polycystines of the World Ocean (in
Russian).
Radiolaria Phaeodaria
Stanley A. Kling and Demetrio Boltovskoy
Introduction structure, which extends into a reduced part, the tubu-

lar, straight to sinuous proboscis (Fig. lA, L). Electron
Phaeodarians are exclusively marine, planktonic organ- microscopy reveals that the striations are reflections of
isms that differ fundamentally from the Polycystina (the complex folding (see Cachon and Cachon, 1973; and
other superorder ofRadiolaria) by distinctive character- Anderson, 1983a for more detail). In addition to the
istics of the central capsular membrane, as well as by a astropyle, there are usually one or more accessory
mass of pigmented spherules, the phaeodium. The cen- pores termed parapylae. Typically there is a pair of
tral capsule is a unique part of radiolarian soft anatomy such pores (Fig. lA). A combination of astropyle and
that encloses an inner zone (intracapsulum or endo- paired parapylae is characteristic of many phaeodar-
plasm), separated by the capsular membrane from the ians, resulting in the name Tripylea which Hertwig
outer zone (extracapsulum or ectoplasm). Phaeodaria (1879) applied to the Phaeodaria. However, Haeckel
are generally larger forrns whose skeletal elements are (1887) wams that parapylae seem to be lacking
often hollow tubes rather than solid bars. The skeletons altogether in several families (Challengeridae,
are composed of opal, but with some organic content or Medusettidae, and Castanellidae), and in a few species
matrix that is still poorly understood. Almost all the spe- in other farnilies. Rarely, there may be only one, or
cies are solitary, except for the genera Nationaletta, several (in access of two), parapylae.
Tuscaridium and Tuscaretta, which have been observed
to form colonies of up to 16 individuals enclosed in a The phaeodium usually occupies the ectoplasm, but
common meshwork sphere (Haecker, 1908; Ling and may also extend into the endoplasm, and may rival the
Haddock, 1997) (Fig. 1Q, 6.153a). rest of the cytoplasm in volume. It consists of spherules
of various yellow, brown, or green hues, and occasion-
Phaeodarian skeletal geometry is quite varied and ally some red granules. Recent studies of phaeodia
complex, with a number of basic plans that are the (Gowing 1986, 1989, 1993b; Gowing and Bentham,
basis for recognizing families. One family produces 1994) show that it is an assemblage of food and waste
fairly simple latticed spheres, very similar to those in vacuo les containing a wide range of materials including:
many spumellarian polycystines (Fig. 6.57-6.76). In bacteria, prokaryotic and eukaryotic algae, algal cysts,
another group with similarly latticed walls the shells dinoflagellates, rnicroheterotrophs, large virus-like par-
are divided into two symmetrical, mirror-image ticles, loricae, coccoliths, diatom frustrules, tricho-
halves or valves, as in pelecypods (Fig. 6.116-6.121). cysts, cuticlar remnants and nematocysts of metazoans,
Other bivalved forms develop radially into elaborate- amorphous material, and siliceous skeletal fragments.
ly branched structures that eclipse the inner val ves It is presumed that these are representative of nutrition-
(Fig. IK, L). Several groups build an open polygonal al sources, although some of the debris may have
meshwork resembling the architectural structures entered the cells as remnants of secondary food sourc-
known as geodesic domes (Fig. 6.51a, 6.43, 6.143a, es such as fecal pellets or organic aggregates.
6.145). Species of one family construct skeletons by
accreting foreign material s, usually the siliceous Phaeodarians are heterotrophic feeders, and most spe-
skeletons of other protists, such as diatoms, polycys- cies so far investigated are generalists, with vacuo les
tines, and silicoflagellates (Fig. 6.134, 6.135). containing a wide variety of materials, as listed above.
An occasional specimen bearing vacuoles mainly with
The phaeodarian central capsule is double-walled, a single food type has been observed in a species
with a thicker outer wall placed very close to a thinner which in other cases contained various food types.
inner wall. One main opening in the wall is surround- And the concentrated resources in organic aggregates
ed by a characteristic phaeodarian anatomical feature, are considered a potentially important food source
the astropyle (Fig. lA, L). The main part is the oper- (Gowing and Bentham, 1994). Unlike many polycys-
culum, a flatly conical or cap-shaped, radially striated tines, phaeodarians apparently lack symbionts.

214 Radiolaria Phaeodaria
eetoplasm
(extracapsulurn)
nuclei
astropyle
(opereulum)
proboseis
Aulactinium actinastrum
PYRAMIDS (=PYRAMIDAL ELEVATIONS)
vertieilof
./ spines
radial spine ;~~ pyrarnidal
~ ortube
pyramidal
1 elevation
with axial
'P;~\
tube or pyrarnidal
elevation
without
axial rod
.. rod
Circoporidae
n\.
W ~ .rod
axial
eoronet of
4.._-.~;""'=~~=-~- ~ tangential
spine or tube terminal branehes
'\ (tangential mesh)
spinulate
Aulosphaeridae, Sagosphaeridae shaft
Radiolaria Phaeodaria 215
AREOLATE
STRUCTURE
(Challengeridae)
W
11\ ab?ral
splnes
Tuscaroridae
f)
Challengeridae
galea
o cross section
through
shelrwallthe
anchor
barbs
outer
lattice
mantle
Coelodend~idae
Aboral (posterior, distaD

Challengeridae Concharidae Medusettidae
Tuscarorid colony,
presumably Tuscaridium
cygneum (Murray)
Fig. 1. Main features of a phaeodarian ceIl (A), and characters used for the identification of phaeodarian radiolarians (B-P). A, F, G, K, L, 0,
P (left): from Haeckel (1887); B, C: from Borgert (1909b); D, E, 1, J: from Haecker (1908); G (photos): courtesy of M. Gowing; H, N: from
Borgert (1911); M: from Borgert (1906); P (right): courtesy ofRenate Bemstein; Q: courtesy of Steve Haddock.
Reproduction in the Phaeodaria is still poorly known. Phaeodarians have received little concentrated study
Flagellated swarmers have been observed and are gen- since the large expedition monographs of the late
erally presumed to represent a sexual stage, but their nineteenth and early twentieth centuries (see below).
exact function remains elusive. Apparent dimorphism Some recent studies of one or more families based on
in some species may be related to a sexual reproduc- sampling pattems approaching oceanic scale are:
tive stage (K1ing, 1971). Fission also occurs, often K1ing (1966, 1976), Tibbs (1969, 1976), and Tibbs
accompanied by large and numerous polyploid chro- and Tibbs (1986). Reshetnjak (1966) reviews collec-
mosomes that are visible even in the light microscope. tions from the northwestem Pacific and provides a
At least some species undergo an unusual mode of detailed account ofthe biology, morphology, system-
mitotic division (Anderson, 1983a; Grell, 1973). atics and world-wide distribution of the entire group.
The first phaeodarian species were described by Phaeodarian skeletons are rarely preserved in sediments.
Haeckel (1862), who placed two ofthree new genera in Either the peculiar siliceous shell material or the thinness
new families but did not otherwise separate them from of the skeletal e1ements, or both, result in such rapid dis-
sphaeroid polycystines. In 1878 Haeckel recognized solution rates that many species disappear from the
these species as a separate group, the Pansoleniae, water-column even before reaching the sea-floor
based on the tubular structure oftheir skeletal elements. (Takahashi, 1981). Nevertheless, a few species have
Hertwig (1879) placed Haeckel's genera in the been reported from sediments in some areas, and rarely
Tripylea in reference to the distinctive arrangement of in fonnations as old as Eocene (e.g., Dumitrica, 1964,
pores in the central capsule. Haeckel (1879) noted 1965,1973; Ling, 1991; Petrushevskaya andReshetnjak,
Hertwig's work ofthe same year, but observed that the 1980; Reshetnjak, 1971; Stadum and Ling, 1969).
set of three pores was not universal, renaming the Because of the lirnited fossil record, there is as yet no
group Phaeodaria after the typical anatomical structure. paleontological evidence of evolutionary relationships.
Geographic and vertical distribution 350

"O
~ 300
Phaeodaria inhabit all oceans at all water depths. Fig. (;
u
2, based on data from Reshetnjak (1966), illustrates the E: 250
Vl
total number of species recorded in all major oceanic .9! 200
basins, as well as the percentages of forms endemic to u
<lJ
each ofthem. Highest total numbers have been record- ~ 150
ed for the Atlantic and Pacific Oceans (343 and 276 ~ 100
<lJ
species, respectively), where also the highest degrees ..o
of endemism are observed. Indian Ocean inventories E 50
:J
are largely shared with the Pacifico As expected, the Z o

u u u
zO~Z¡::
CIl:
¡:: U
-1
CIl:
u:::
<C
<C
15 ~
Z
Z
¡::
polar seas host the lowest specific diversities.
~
<C It should <C
be noted, however, that to a large extent these esti-

mates reflect the different degrees of coverage of the
various areas. Relative differences between polar and
extrapolar domains, however, probably reflect actual
relationships more faithfully. Fig. 2. Numbers of phaeodarian species recorded in, and endemic
to, each major oceanic basin. Based on data from Reshetnjak (1966).
Unlike most other sarcodines, phaeodarians are typi-

cally deep-water animal s usually peaking in both Om~r----------------------------------
50~
abundance and diversity below 200 m. A detailed
depth zonation for the area of the Kurile-Kamchatka
trench was produced by Reshetnjak (1955,1966). Her
data (Fig. 3) indicate that only 2 (of the 103 species
recorded) inhabit the upper 50 m; approximately 30
more phaeodarians have restricted vertical ranges at
various depths; while over 50% of the taxa were 1000
2000 1-
retrieved from the broad depth interval of 50 to 2000-
8000 m. It should be borne in mind, though, that
3000
although uppermost records for a given species are
generally reliable, the lowermost ones often are not,
4000
since dead sinking shells can be counted together with
in situ living organisms (see Kling and Boltovskoy,
5000
1995). Interestingly, zoogeographic zonations based
on the degrees of endemism of phaeodarian assem-
6000
blages (Fig. 4) are more similar to zonations based on
bottom, deep-water organisms, than on those based on
planktonic groups. This circumstance reflects the 7000
deep-living habitat of a large fraction of the phaeodar-
ians described (Reshetnjak, 1966). 8000
These vertical patterns at a given locale, however, 50 species

11I"""-"""11
ttt-;;¡'o¡~,}_d_~l
may change significantly because of the dynamics of
deep ocean circulation, with species exhibiting quite stenobathyal
variable depth ranges over oceanic distances. For ••• eurybathyal
example, species tied to surface water masses in polar
or subpolar regions, where dense waters sink to form Fig. 3. Numbers of phaeodarian species recorded at different
the Intermediate Waters, are typically found at gradu- depth interva1s in the Kuri1e-Kamchatka trench area (northwestem
ally deeper horizons toward equatorial waters, Pacific). Based on data from Reshetnjak (1955, 1966).
TmpicalH-+-++++-~L
Padfic++++-t-+++++
Province ++++-H-+-H-
Deep Water Biogeographic Regions

~ ~ 1----::----
Atlantic Pacific North-Indian Antarctic
Fig. 4. Biogeographic zonation of the world ocean on the basis of deep-sea phaeodarians. Redrawn from Reshetnjak (1966).
where they appear at depths of hundreds or thousands or the Deep Water, to reappear in the opposite polar or
of meters. Thus, on regional scales, depth distribu- subpolar seas. In the Pacific Ocean, the Intennediate
tions of phaeodarians are not describable in tenns of Water circulates in anti-cyc1onic gyres that mimic the
fixed ranges. surface circulation (Reid, 1965), thus providing conti-
nuity for the water masses and their biological contents.
Vertical profiles in the North Pacific (Kling, 1966, Such bipolar patterns have been described for a number
1976) illustrate these kinds of relationships (Fig. 5). of zooplanktonic species in both the Atlantic (Pierrot-
Some species occupy the Subarctic Pacific Water Bults, 1974) and the Pacific Ocean (Alvariño, 1965),
Mass at depths as shallow as 25 meters, but sink to whereby the north and south polar or subpolar near-sur-
greater depths southward where a salinity minimum face populations are joined at depths of 800-1000 m
defines the Intennediate Water (Fig. 5C). Above the across the equator.
southern ranges of these species are others occupying
an unnamed salinity-maximum zone (Fig. 5B). And In addition to such physical factors, phaeodarian dis-
these in turn underlie surface-dwelling Central Water tributions may also be influenced by biological vari-
and equatorial species (Fig. 5A). ables. In the South Atlantic Meyer (1934) showed
close coupling between the absolute abundances of
Equatorward submergence may account for so-called Challengeridae and primary producers such as dia-
bipolar distributional patterns characteristic of many toms and peridinians, as well as between larger phae-
phaeodarian species. Thus, high-latitude species could odarians and total plankton and phosphate content.
pass under equatorial waters via the Intennediate Water Dworetzky and Morley (1987) showed depth zones
A Castanidium longispinum
100
200 1600
300
400~~~--~--~--~--~---L~~L--------------
B Castanea amphora
Om~------------------~------~-----------------
100
200
300
Castanidium variabile PííZ!1-49
om-r--¡n-~~~~
100J
Píí 150-499
500-4999
200
Individuals
300 per 1,000 m3
of water
filtered
Fig. 5. Horizontal and vertical distribution of 3 phaeodarian species with different water-rnass affinities in the north-eastern Pacifico Redrawn
from Kling (1976; with kind pennission from Elsevier Science).
for the upper 200 meters of the eastem equatorial ries: the Challenger Expedition (1873-1876), the
Atlantic related to various physical-chemical features Plankton-Expedition (1889), the Deutsche Tiefsee-
and to a chlorophyll maximum. Expedition (1898-1899), the Deutsche Südpolar-
Expedition (1901-1903), and the Deutsche Atlantische
Some authors (e.g., Haecker, 1908; Reshetnjak, 1966) Expedition (1925-1927). Stations were gene rally
suggested that deep-living species are generally larger widely spaced and relatively few closing net samples
in size than their shallower-dwelling relatives. were obtained. Nevertheless, many species were
Reshetnjak (1966), based on comparison of the size of reported from restricted geographic provinces, and in
9 Challengeridae, noticed that the shells of 3 species some cases relatively narrow depth ranges are sug-
which live at 50-400 m vary between 0.06 and 0.18 gested. Because of better sample coverage in near-
rnm, for 3 others characteristic of the 400-1500 m layer surface depths, ties to surface water masses and cur-
the size spans 0.18-0.30 rnm, while 3 more that inhabit rent systems are fairly well established by existing
the 1500-5000 stratum range from 0.30-0.65 rnm. South Atlantic data. It is also likely that a number of
species that appear at shallow depths in the Antarctic
It is difficult to specify precisely the distribution pat- and at greater depths at the equator are occupying the
tems of phaeodarian species in the South Atlantic Intermediate Water as has been shown in the North
because the data are sparsely distributed spatially and Pacific, and was suggested by earlier authors for the
temporally and have been gathered with various tech- South Atlantic (e.g. Meyer, 1934). However, continu-
niques and equipment. Most of the phaeodarian data ous data from great depths, that would be required to
come from reports of the large oceanographic expedi- substantiate such anticipated deep distributions, have
tions of the late nineteenth and early twentieth centu- not yet been gathered.
",
Warm water Cold water
0
•..
~
"i;
\,1
>- ti
•.. •..!U
'"
.:.t.
~ .......
_.-
- --
"i;-ci e . :ll Depth
.~ E0 •~ "C ~ U lo:' OU !U.t::
\,1
.s!U..c
Q,,!tI
o E t:..;
•..
"i; .-
.-
!ti
QI e ~ ti~
••• QI
(m) E
~
QI
~ Q,,6 ~
'"
eoo ::;¡ !U0 !U 'U
_ 'U c:.::
::::1 0.0,", ::::I::uc:r e
C".- ::;¡
E.2g~ ~ !U
•••
t:
QI
w..clJ U ~a:> ~~ ~~
Astracantha heteracantha P >1000? 15
Atlanticella bicornis P --_._-- ¡-._.
Atlanticella craspedota P P
Atlanticella olanctonica 0-3000 0-2000 15
Aulacantha cannulata P P P
800- 200- 50-
Aulacantha scolymantha P P 200-400 x P 7
1000 x? 400 x 1700 __ 'o
---
400-
Aulacantha scolymantha bathybia P P P P P P
1000
Aulastrum mirabile P P
Aulastrum monoceros P P
Aulastrum soinosum P 0-600 11
0-
Aulatractus fusiformis P P P P
1000
Auloceros arborescens birameus P P 18
250-
A. arborescens dichodendrum P P 18
500
A. arborescens irreguleris P 18
A. arborescens subeleeens P P P >500 18
A. arborescens trigeminus P 18
Aulographis bicurvata P P >400? 8
800-
Aulographis furcula P P 18
1000
400-
Aulographis pandora P 10
80Q
Auloereohis stellata P P [>400] 9
Auloereohis tetrancistra P P P 1>500] 10
Auloereohis tetrestvle ? ? Deep? 18
Auloereohis triengulum ? ? 18
Auloersohonium antarcticum 1>500] 11
Auloereohonium anthoides [>500] 14
Auloereohonium bicorne P P P >500 m 18
Aulozreohonium candelabrum ? ? -- 18
Aulogrephonium mediterraneum ? ? 18
Auloereohonium oulvinatum ? ?
Aulokleptes flosculus P P P 10
Aulokleotes ramosus P P P P 1>5001 9
A. ramosus varo antarctica ? P 2
A. ramosus varo longisoine ? ? 1
288-0,
Auloscena pyramidalis ? ? ? ?
272-0
Auloscena robustissima P 18
Auloscena verticillus hamata [>5001 11
Auloscena verticillus laevis P P 15
Auloscena verticillus tvoice ? ?
Aulospathis oinus P P P [> 1000] 15
-
~s::::
--.=~ Oepth
.
OU
.- '"
.-
!IJ
Q)
::J
(m)
s:::: 00
"' c:
¡A~/~{2~Jll;--;:;~i-~b¡i~~b·i¡w~ª __ .__
!\. variªl:JiJis aulQd..ffl..cJ!..oicf.es____ p .___
p
_ .
_ __ ---Ll-m
L__
_._?
?
[> 1OOOJ-
P __ ;-
__~Il
1.8_
[~~ggr
I
_6LJJos{2EJbJsvariaQilisdiQSio!l. . -- -. --.-.-- _ --- - E_e- E.__ .-+--.- 1L
A ulospathis _va~b~/is~~~~~ ~__ P .._ .__~ ~ __ e------ __1000 x? 9

Aulº82haera_bist~nar~a qvulu!!L P u. . . .. 1---. 1---
A. bistemaiÍA5..fe!....E;!1_tri9.!l.ªlis ._____ ..__.__ ._. . ._. [>SOOJ .___ ....2....

Aulosphae{a ele&YJ.tissim~ _ . . ..__ . 200-400 x? ..l2..
Au(oJRh..ª-eia filigera __.______.E.. .---f--.-. u. • _
I
Aulosphaera labradoriensis 0-400 <SOO?
IAuJp~phaera robu~n P P ..l.?O-(SOO?) __
= --.-
1-_
AulosplJ.aera triodolJ....__
Aulosphaera trispaJhis __ .
Aulosphaf:!!,a~rticillata.L~nt.
u. __
r.
•
E_.
nn
-+__
? ~-- ? ?__
•
=-=-_~=.--=-.-----" ~ 300-500
_12~0_
-2_00 x_ -_--_-
~=18
18
---
Aulosphae.ia_ verticjlLéi1aL~glJ-ª1-f. P..u. ._ _
BOJg~®Jla cª-Ll2ata____ __ _ _ 1- º::.4-ºº--- __ _ ___ __ .
:::::~~_:;:r::~:C~_
Ca.nn.~pb-ªera~!Jlª-!.c:JLc..ª
. 0~~Q9
n_u. _
0_~2_Q9_~___
_1--_O~.2_~ • ~ .. II • __ ._._____
2~~~¿oL---;
Castanariuf!1 antarsJicuf!l ._ _ . . 300-700 1~_ 1..
CastalJ!1rium favosuf12
(asta!l.9rium hooke!i_J.'
Castanea ª!IJ.J2ho{a..
._.. P
__
E_._. P L
~~~~--~=~_
._P.
_1-_
?
? u.
. -JZ.
_ 18
.-.----------=.= -~ J-=__.=::-= --.l
Ca~tanea gjpjJ_osa_ P!,_ _ _ _ __ __ _ JJL.

Castanea henseni P 16
Castªnella borgerti 1-_._ .. _ _ .____ _ _ .L_I--_____ _+-_
~astanella maxima -------------p----e. _ .._E.._ ___ L 1- .__ __ __ ..
-----l___
Castanella thomp~oni. f_+ __

P__ I-- I-__ ---f-- 16
Castanidium h.ª.e_cJ5~L __ . . ... __ f--t-------- -----·----1----+--·-------
CastanidiuJ!J.JongjJ.pjD_u.!Jl
__ ._. ._y_.I-_p____ _
---.- ..---t------+---
0-400 _ -JL. +___
Castanidium!!J.º.~e1.f!.xL . --E __..~-- .E._.. ------I-----r- 0-400 __ 1--. __
Castanidium spinosum __ . . ..._1-_ -- .....1--- _ ----i_-'0"----'4cO".~0_'__.
1---. 1---
_C_asJ_an_I_·di.t{m_vª!l._ho_eff~!!L_ _ . .1-____ . ... + 1--- --+ ,0"-'-400 14
~~f~~ElE!;~=~
Castanidium variabile P P P P 50-100,
~.~.~·-i~:=r~o-rt-~-=~~~-~~:,===(OO-6~-=1 -----L- I----.
-1-_ -
• • •. ••• __ ~._. u ••• •• ._._. • •
Ca~ªnoP?Ls_fragilis------ _.-- --1-- -- - --- __ ~-0..::2.S0 .--L----J 1..

I~..._-
._~?~=,~~_ - ~~...:~~
(1J_~R.eria naresi _.__ _ _ ~_ p_. ----I--L----P--- _'__0-15 c-li-.
;~:::;::::2~;::;:~_
=~:~~~!T:C:OO
u
•..>-
CII
~e .
u
~•..
~-c 0'8
•.. '"
eu ~ o~ . o :ll Depth
o;:
.s c. '"
o;:"t:I ~ U
o e s;
~ c.8 ~
~•..
•..e •.. -
Ou
0-
0-
'"
<1>
•...
tU~ -
e ~
u
0'8~
••• <1>
(m)
tU
E
CII
tU..c <1> eco
'" :J tU° ~:g c.::
:l
C'"o_"".=
:J
l.I.I..clJ
:l ~ U
~..Eg~
C"
CII
U
tU e
.:~ ~~ ~~
150- 900-
Chal/engeron diodon 50-1000 600-800 4
1050 1100
Chelleneeron radians
200- 50-
Challengeron wil/emoesii
400 x? 1000
Circooorus oxvaeanthus 0-400
Circooorus sexfuscinus ? ?
Circosoethis sexfurea ? ? P P ,
Coeleehinus weoiticornis 0-200 14
Coelodeeas ambulaerum P
Coelodeeas ovzmaea P P P <1000
Coelodendrum flabellatum ? ? ?
Coelodendrum fureatissimum P P 600 X
Coelodendrum laooaeeum ? ? ? DeeD
50- 0?-400, SO- SO- 50-
Coelodendrum ramosissimum P 18 I
400 X? -900? 400 400 400 I

I
Coelodendrum soinosissimum ? ? ? I
Coelodieeras spinosum ? ? ? I
Coeloereohis antaretiea P Deeo 11 i
Coeloereohis reeine P P P 18 !
I
60-
Conehellium eapsula
200 X I
60-
Conehidium eaudatum
200 x
I
r
Conehidium terebratula ? ? ? I
Conchophecus diatomeus ? ? ?
400-
Conehopsis eompressa
1000?
iI
Conchoosis orbieularis ? P
Cornueella maya P P
Euohvsetta dubia [>500] 14
50- 300- 900- 600- 700-
Euphysetta elegans 4
1000 1100 1100 800 x 900 x
Euohvsette lueani P P 0-800
Euohvsetta ousilla 0-400
400-
Haeekeliana darwiniana
1000?
Haeekeliana irrezuleris P P P >100 10
Haeekeliana ooreellana P P P 10
I
Haloeella gemms 0-300 0-300 I
Haloeella mama P P
Lirella bul/ata 0-400?
Lirella melo 0-1000?
!
I
Loboeella ptoteus P !
Medusetta ansata 0-400
Medusetta inflata ? ? ?
u •..
Q,I
-¡¡j
v el)
~•..
-¡¡j-d -¡¡j ,~ ,
>- e o ''8
•..
.s~..cEc.u'"
'¡:
~ ~o!:
r::r,- "
w..cLJ
<li
'¡: -o ~ u
o E s;
~ c.6 ~
~3:uo-
~.2g~
\,J
-¡¡j
-
U
•..
e
Q,I
--
,-
,-
~ <:
..:~
Ou
el)
eco
<li
'"
"
~.;::
-e 0-
~o
~~
~
val
''8~
••• <li
~"
-"
~~
Depth
(m)
~
~
E
Q,I
Medusetta rara ? ? ? i Deep

Nationa/etta Ireeilis ? ? I P
Phaeocol/a embieue 0-200
0-100 x
Phaeodina antarctica
-<600
I
Phaeodina triov/ea P I
II
Phervneel!e eestrule ? ? ? Deep
I
150-
P/anktonetta at/antica P P P P
01100
1000
Porospathis holostome >400 10
-4000
Protocvstis acornis <200 <200 15
400-
Protocystis balfouri p P ?
600 x
200- 50- 0-
Protocystis bicornuta
400 x 100 y 1100
Prptocy'stis_ fJic;uspis -_. -- - ----- o o - - - _.~ _. - ----- - -
0-200 _o _ _o o _ o -
50- O-50?, 200- 850- 700-
Protocystis harstoni 3
800 300-1100 1100 1050 900 x
Protocvstis heterocentrota 0-200
Protocvstis macleari ? ? ? Deeo
200-500 x,
Protocystis micrope/ecus 14
450-600 x
Protocystis murrayi ? ?
--
? - - D_eep
Protocvstis slozeetti ? ? ? Deep
200- 100- 100-
Protocystis swirei 5
600 200 x 200 x
800- 200- 50-
Protocystis thomsoni 5
1000 1100 1050
200-
Protocystis triangu/aris
500 x
800- 200- SO- SO-
Protocystis tridens 5
1000 400 800 100 x
400-
Protocystis tritonis ? ? 6
1000?
Protocvstis varians P P Mid
Protocvstis xiphodon P P P P 0-1000
Sagenoarium antarcticum P P 14
Seeenoerium chuni p p ? 18
Segenoerium dicranon P P 18
Segenoscene itmineerlene P [>3001 11
Segenoscene tetracantha P
Seeoscene castra P I P 11
Sezoscene oreetorium P 8
Tuscarantha braueri P 0-300 18
Tuscarantha hvdra P
Depth
(m)
Table 1. General distributional data for the phaeodarian species reported from the South Atlantic Ocean. Data are entered according to major bio-
geographic provinces indicated in Fig. 11 of the section on Polycystine Radiolaria. Where depth data suggest a reasonably restricted range for a
given province, the depths are entered under the appropriate heading. Generalized depth ranges are entered in the column headed "Depth". When
data are sufficient to suggest a concentration of individuals at a particular depth or depth-range, the depths are noted as maxima (abbreviated "x").
Remarks:
1: Warm water form; 2: polar cold water form; 3: deeper to South?; 4: deeper to South; 5: deeper to North?; 6: deep; 7:cosmopolitan; 8: bipolar?;
9: bipolar submergent?; 10: bipolar submergent; 11: bipolar; 12: Antarctic endemic? Bipolar?; 13: Antarctic endemic?; 14: Antarctic endemic;
15: Antarctic, north-submergent?; 16: also central North Pacific 200-300 max.; 17: also central North Pacific 100 max.; 18: also central North
Pacific?; 19: also Mediterranean, north-submergent?; 20: also Indian Ocean.
Table 1 summarizes the distribution of phaeodarian used by subsequent students of the group, and some
species reported from the South Atlantic Ocean. Data new families have been added. The morphology of
are entered according to major biogeographic provine- each family is so distinctive that there has been essen-
es reported in the literature. In addition to expedition tially no controversy as to their taxonomic identity.
reports and other sources cited above, data were com- Haeckel also proposed a c1assification of families into
piled from the following publications: Dworetzky and orders, which is presented here, although it is perhaps
Morley (1987), Gowing (1986, 1993a), Morley and in need of revision. It is based on skeletal morpholo-
Stepien (1985), Nóthig and Gowing (1991), Peters gy, and further analysis of the cytological relation-
(1929), Takahashi and Honjo (1981), Tibbs (1969, ships might provide a basis for a more natural order-
1976), and Tibbs and Tibbs (1986). Where depth data . level classification. The generic c1assification
suggest a reasonably restricted range for a given prov- proposed by Haeckel (1887) has been followed gener-
ince, the depths are entered under the appropriate ally in subsequent works, but occasional revisions have
heading. Generalized depth ranges are entered in the been suggested, and inconsistent usage (particularly
colurnn headed "Depth". When data are sufficient to among the challengerids) persists into modem times.
suggest a concentration of individuals at a particular
depth or depth-range, the depths are noted as maxima Remarkable intraspecific variations have been
(abbreviated "x"). described for this group. Reshetnjak (1966) noticed
that, for example, in Aulosphaera bisternaria and
Sagenoarium chuni the shape of the meshwork can be
Taxonomy either spherical, oval, spindle-shaped or pyriform.
Spherical and pear-shaped Castanidium variabile occur
Morphology and classification system regularly. Tuscarantha braueri 's pyramidal shell can
be either 3 or 4-sided. Tuscaretta tubulosa can be pear-
The c1assification of phaeodarian families presented shaped, egg-shaped or ovoid. Intraspecific variability is
here was put in place by Haeckel (1879). It has been not circumscribed to shape, but also affects other mor-
phologic traits. In Tuscaretta tubulosa, for example, the Galea (Coelodendridae): Dome-shaped to irregular
number of oral radial spines can vary from 2 to 5. In sack -like extension of valve of inner shell, usually
some cases this variability is probably associated with with triangular base (Fig. lL).
ecological settings: in some Tuscaroridae, for example, Girdle fissure (Coelodendridae): space separating the
the number of radial spines increases with decreasing two valves (Fig. lL).
water-temperature (Reshetnjak, 1966). Intracapsulum: protoplasm inside the central capsule
(Fig. lA).
Leg (Medusettidae): major appendage or spine (Fig.
Glossary 1M).
Mouth: a large, dominant pore or opening in the shell
Aboral: Not oral, located away from the mouth (Fig. (Fig. lB, M).
lN-P). Nasal style (=main style, odd style) (Coelodendridae): a
Alveoli: vacuole-like, fluid- or gas-filled inc1usions in single dominant style (see tergal styles) (Fig. lK).
the ectoplasm, usually in a distinct frothy (in Nasal tube (Coelodendridae): see rhinocanna (Fig. lL).
spherical forms concentric) layer (Fig. lA). Operculum: the main, cap-shaped to conical portion
Anchor barbs: barbs shaped like anchors (Fig. 1K). ofthe astropyle (Fig. lA, L).
Aperture: usually odd, major hole or opening in the Oral: pertaining to, or in the vicinity of, the mouth.
shell-wall (see mouth). Oral spines (=teeth, cf. Haeckel): spines surrounding
Apical hom: spine at the apex of the shell (Fig. 1M). the mouth (Fig. 1M).
Areolate: referring to complex, more or les s tubular, Parapylae: paired pores, smaller than, and structurally
regularly repeated wall perforations or pores (Fig. different from, the astropyle (Fig. lA).
1G). Peristome: differentiated area surrounding the mouth
Astral septum (septa): partition separating tubes where (Fig. 11, N, O).
they are conjoined. Phaeodium: mass of pigmented spherules in the cyto-
Astropyle: a conical or cap-shaped, radially striated, plasm (Fig. lA, L).
structure surrounding the main aperture in the cen- Primary shell (Medusettidae): main shell formed of
tral capsule, with a tubular extension, the probos- central elements.
eis (Fig. lA, L). Proboscis: see astropyle.
Basal pores (Tuscaroridae, Circoporidae): pores at the Pyramid (=pyramidal elevation) (Aulosphaeridae,
base of the spines (Fig. lB). Sagosphaeridae): structure consisting of spines
Central capsule: a sac enclosing the intracapsular pro- arranged in the shape of a pyramid (Fig. ID, E).
toplasm and the nuc1eus (Fig. lA, L.) Pyramidal elevation: see pyramid.
Coronet (=corona): crown-shaped arrangement of Pyramidal tubes: tubular spines arranged in a pyrami-
thoms at the tip of a spine or style (Fig. lF), of dal structure (Fig. ID, E).
pores at base of main-spines (Circoporidae, Fig. Radial spines: spines oriented as the radius ofthe shell
1C), or of other elements. (Fig. lA, B).
Cupo la: a dome-shaped structure. Radial tubes: tubular (hollow) spines oriented as the
Ectoplasm: protoplasm outside of the central capsule radius of the shell (Fig. 1D).
(Fig. lA). Rhinocanna (Coelodendridae): tubular lateral exten-
Endoplasm: protoplasm within the central capsule sion of galea along the central capsule (Fig. lL).
(Fig. lA). Spathilla(ae): term applied to more or less flattened,
Extracapsulum: part of the cell surrounding the cen- usually spine-bearing, knobs at the tips of spines;
tral capsule (Fig. lA). could also mean flattened, blade-like spines or
Feet: Radial spines or appendages arising from mar- thoms (Fig. lF).
gins of terminal opening (Fig. 11, J). Spinulate: beset with tiny spines (spinules) (Fig. lF).
Fenestrae: more or less large opening on the shell wall Style (Coelodendridae): prolonged tubular main spine
(Fig. 1H). (or tube) extending outward from valve (Fig. lK).
Frenulum (Coelodendridae): brace- or buttress-like Tangential meshes (Aulosphaeridae, Sagosphaeridae):
bar, tube, or delicate latticed bridge connecting meshes making up the main spherical shell (Fig.
galea to rhinocanna (Fig. lL). ID, E).
'P
Tangential tubes (Aulosphaeridae, Sagosphaeridae): capsule either strewn or regularly ordered. The skele-
tubes fonning the tangential meshes (Fig. ID, E). tal elements (if present) in these families are loosely
Teeth (=oral spines): spines surrounding the mouth associated in the cytoplasm, not united into a fixed
(Fig. 1H), or (Concharidae) tiny spines along structure. The taxonomic relationships among these
hingeline (Fig. 1P). families needs to be re-examined.
Tergal style (=lateral style, paired style) (Coeloden-
dridae): one of two similar, paired styles, usually Family Astracantbidae (Fig. 6.1, 6.2)
smaller than the nasal style (Fig. 1K). Skeleton of radiating hollow tubes which are proxi-
Valve: symmetrical half shell (Fig. 1L). mally fused. Rarely found.
Verticil: a set (e.g. of spines) arranged in a circle or
whorl around an axis (=whorl) (Fig. 1F). Family Aulacantbidae (Fig. 6.10-39)
Skeleton of numerous hollow spines radiating free
(unattached) from the central capsule. Usually with
Outline classification, order and family-level peripheral veil of fine tangential needles. Because the
diagnoses spines are unattached, they often appear isolated or in
small groups in samples.
The higher classification of families presented here fol-
lows that used in the chapter on Radiolaria Polycystina. Family Pbaeodinidae (Fig. 6.134-137)
I Phaeodarians lacking a self-secreted skeleton. In
I Kingdom Protista Haeckel, 1886
Phylum Sarcodina Hertwig and Lesser, 1876
some fonns, foreign siliceous particles (usually sili-
ceous skeletons of other organisms such as diatoms,
Class Actinopoda Calkins, 1909 polycystine radiolarians, silicoflagellates, or sponge
Il' Subclass Heliozoa Haeckel, 1886 spicules) may be irregularly distributed in the extra-
Subclass Acantbaria Müller, 1858 capsular cytoplasm. The foreign skeletons may
Subclass Radiolaria Müller, 1858 belong to a single species or be a mixed assemblage.
Superorder Pbaeodaria Haeckel, 1879 Fonns harboring foreign siliceous bodies have also
Order Pbaeocystina Haeckel, 1879 been placed in the family Caementellidae Borgert
Family Astracantbidae Haecker, 1908 (1909a), now placed in synonymy with the
Family Aulacantbidae Haeckel, 1862 Phaeodinidae by Cachon-Enjumet (1961). Specimens
Family Pbaeodinidae Haeckel, 1879 of this family may resemble fecal pellets or organic
Family Pbaryngellidae Haeckel, 1887 aggregates (Gowing and Coale, 1989). Hertwig
Order Pbaeospbaeria Haeckel, 1879 (1879) and Haeckel (1887) described a number of
Family Aulospbaeridae Haeckel, 1862 phaeodarian species bearing.essentially monospecific
Farnily Cannospbaeridae Haeckel, 1879 silicoflagellate assemblages in the genera Cannopilus,
Farnily Sagospbaeridae Haeckel, 1887 Distephanus, Dictyocha, and Mesocena, which were
Order Pbaeogromia Haeckel, 1879 originally erected for individual silicoflagellate tests.
Family Atlanticellidae Borgert, 1905 Haeckel (1879) described the family Cannorhaphidae
Family Castanellidae Haeckel, 1879 to accornmodate these species. Borgert (1890) recog-
Family Cballengeridae Murray, 1876 nized that many of these fonns were a separate group
Family Circoporidae Haeckel, 1887 of flagellates and separated them from the Radiolaria.
Family Lirellidae Ehrenberg, 1872 Some genera and species not tied to monospecific
Family Medusettidae Haeckel,1887 assemblages of silicoflagellates remain in the family
Family Porospatbidae Borgert, 1901 Cannorhaphidae (see e.g. Borgert, 1909a), but many
Family Tuscaroridae Haeckel, 1887 ofthem have never been illustrated, and the status ofthe
Order Pbaeoconcbia Haeckel, 1879 family remains in some doubt. Most or all of the
Farnily Coelodendridae Haeckel, 1862 remaining cannorhaphid species would now be placed
Family Concbaridae Haeckel, 1879 in the family Phaeodinidae.
Order Pbaeocystina (Fig. 6.1-6.2, 6.10-39, 6.134-137) Order Pbaeospbaeria (Fig. 6.40-56, 6.140-146)
Siliceous skeleton lacking altogether, or consisting of Siliceous skeleton of numerous hollow or solid tan-
hollow tubes or foreign objects outside the central gential tubes organized in a regular fashion into a
1
large, usually spherical or polyhedral, openly latticed circular to subcircular pores, numerous by-spines, and
shell. The outer lattice work is a regular arrangement sometimes larger main-spines, and with mouth, usual-
of polygons, most1y triangles, resembling architectu- ly circular, often surrounded by ring of teeth.
ralstructures known as geodesic domes.
Family Challengeridae (Fig. 6.77-98)
Family Aulosphaeridae (Fig. 6.40-55) Phaeodaria with a monaxial heteropolar, ovate to len-
Skeleton a spherical to ellipsoidal, 100se1y structured ticular shell, with reticulate surface sculpture of regu-
polygonal network of hollow tangential tubes. Tubes lar hexagonal indentations resembling the areolate
separated by astral septa where they meet at nodal structure of diatoms, with a mouth usually with teeth.
points. Radial tubes or pyramidal elevations common-
ly arise from nodal points. The structures resemble Family Pharyngellidae (Fig. 6.138)
architectural shapes known as geodesic domes. Shell outwardly similar to those of Challengeridae, but
with a tube-like structure (pharynx) directed inward
Family Cannosphaeridae (Fig. 6.56) from the mouth. Originally described by Haeckel
Two concentric spherical or subspherical shells con- (1887) as a subfamily of the Family Challengeridae.
nected by numerous thin radial bars. Inner shell sim-
ple, spherical or ovate, solid or latticed, with broad Family Circoporidae (Fig. 6.99-104)
mouth. Outer shell spherical or polyhedral, a 100se1y Phaeodaria with spherical to polyhedral shell, with
constructed polygonal network of hollow tangential porcellanous wall structure, with stellate ring of pores
tubes separated by astral stellate septa at nodal points. at bases of hollow radial spines. Mouth usually with
Radial tubes extend from nodes, and are not exten- teeth. Shell surface tabulate, panelled, or dimpled.
sions of the thin radial bars connecting inner and outer
shells. Rarely found. Family Lirellidae (Fig. 6.122-124)
Minute shell, ovate to ellipsoidal. Mouth with bent,
Family Sagosphaeridae (Fig. 6.140-146) slanted, or straight peristome, oral aperture terminal
Phaeodaria with large spherical (or sometimes polyhe- or subterminal. Surface sculptured with regular longi-
dral or pyriform), very delicate shell, of solid, very thin tudinal striae. Sometimes with apical spine at aboral
and long bars. Nodal points of the web-like network pole, or ring connecting aboral pole with lower lip of
fused, without astral septa. Mesh openings large, trian- oral aperture.
gular. Shell surface usually with radial spines and often
with pyramidal elevations. The structures resemble Family Medusettidae (Fig. 6.125-133)
architectural shapes known as geodesic domes. Phaeodaria with simple, ovate, hemispherical, or cap-
shaped shell of areolate structure (resembling diatom
Order Phaeogromia (Fig. 6.3-9,6.57-104,6.122-133, areolae), with hollow articulate feet surrounding a
6.138,6.139,6.147-158) widely open mouth. Distinctive tubular feet (or legs)
Siliceous skeleton lacking, or an individual lattice are subdivided by numerous, equally spaced trans-
shell, spherical, ovoid, or variously shaped. With a verse septa.
large mouth. Often with hollow spines with basal pore
fields. It is questionable whether all the families Family Porospathidae (Fig. 6.139)
placed in this order are closely related. Some should Shell surface panelled, tabulate, or covered by trizonal
probably be separated into one or more new orders. meshwork. With radial spines on all sides. With peri-
stomal tube.
Family Atlanticellidae (Fig. 6.3-9)
Skeleton may be lacking. When present, shell ring- Family Tuscaroridae (Fig. 6.147-158)
shaped, with 4 paired, hollow, articulated, vertical, Shell ovate, spindle-shaped, to nearly spherical, of
descending branches, appended to bladder- (or bub- solid porcellanous structure. A few pores at bases of
ble-) like central capsule. hollow radiallegs, placed syrnmetrically around main
axis and mouth. Surface smooth or spiny, not panelled
Family Castanellidae (Fig. 6.57-76) or tabulate. -
Spherical to subspherical regular lattice shells, with
J------------------------------------------------------~----~--~~--------- 52
Order Phaeoconchia (Fig. 6.105-121) Family Atlanticellidae

Siliceous skeleton bivalved, of two mirror-image 1 Central capsule bubble-shaped .2
halves, simple or with elaborate structures of branch- 1a Central capsule large, tomato-shaped: Atlanticella
ing radial spines. Although both families placed in 2 No skeleton 3
this order are basically bivalvar, the valves of the 2a Skeleton an irregular meshwork of siliceous nee-
Coelodendridae are spatially insignificant and structu- dles: Halocella
rally unlike the regularly latticed valves of the 3 Central capsule with variable number of broad
Concharidae. It seems likely that they should be lobes: Lobocella
placed in separate orders. 3a Central capsule with long hom- or arm-shaped
extensions from disk-shaped central mass:
Family Coelodendridae (Fig. 6.105-115) Cornucella
Phaeodaria with bivalved lattice shell, with conical
cupola or pyramidal galea at apical pole of both • Atlanticella bicornis Haecker (Fig. 6.3). Peristome
valves, therefore at opposite poles of the sagittal axis, coarsely areolate, with 4 perpendicular radial
each valve with 3 or more hollow radial tubes sym- spines. One pair of spines directed toward bladder-
metrically disposed, sometimes with branches form- (or bubble-) like central capsule, bifurcating at the
ing an outer bivalved mantle. The elaborate outer base. The other pair very long, directed toward the
mantle of tube branches is usually the dominant fea- sack-like central shell. Chambers in spines aligned
ture of the skeleton. Some authors have placed species in two rows. Central shell pyriform, thin-walled,
possessing a frenulum and rhinocanna into the family smooth-surfaced. Height of central shell: 0.36-0.42
Coelographididae Haeckel 1887, reserving the family rnm. Ref.: Reshetnjak (1966), Haecker (1908).
Coelodendridae for species lacking those structures. • Atlanticella craspedota Borgert (Fig. 6.4). Shell
with spindle-shaped central area, open at smaller
Family Concharidae (Fig. 6.116-121) (proximal) end, with thickened rim, distally point-
Spherical to ellipsoidal shell divided into two indepen- ed, with 4 divergent ascending primary branches,
dent, hernispherical to boat-shaped, equal or unequal forming small basket at indented base of tomato-
valves similar to the two valves of clam shells. shaped central capsule, each with single, simple,
divergent, descending secondary branch, about
twice as long as the central part of the shell that
Identification of species they surround. Length of central area: 0.32-0.40
rnm. Ref.: Schróder (19l3).
The species listed are nearly all of the ones that have • Atlanticella planctonica Borgert (Fig. 6.5). Shell
been reported from the South Atlantic, including the with spindle- or cudgel-shaped central area, open
late nineteenth to early twentieth century expedition at smaller end, with thickened rim, with 4 diver-
reports and subsequent literature. A few species that gent prirnary branches, forrning basket around
are extremely rare, or for which no illustration is tomato-shaped central capsule (not illustrated),
available, are omitted. Species are listed by families, each primary branch with two simple secondary
and families are listed alphabetically. Brief diagnostic branches, one medial and one distal, from opposite
keys to genera are included where they provide infor- side of primary branch, about twice as long as cen-
mation essential to identifying the species. tral shell area. Length of central area: 0.32-0.40
rnm. Ref.: Schróder (19l3).
Family Astracanthidae • Cornucella maya Borgert (Fig. 6.6). Central cap-
• Astracantha heteracantha (Haecker) (Fig. 6.1). sule bubble-shaped, with 3-7 hom- or arm-shaped
Skeleton of 30-40 hollow spines radiating from extensions. Central capsule wall double, with
the central focal point. Midway along each spine, large operculum. Protoplasm forms disc-shaped
numerous short, irregularly spaced, outwardly main mass, in which the nucleus lies, and from
curved branches bear terminal thoms proximally, which several plasma cords extend. Diameter: 0.5-
becoming terminally smooth-pointed and some- 1.0 rnm. Ref.: Schroder (19l3).
what thicker distally. Length of radial tubes: 1.8 • Halocella gemma Borgert (Fig. 6.7). Central cap-
rnm. Ref.: Reshetnjak (1966), Haecker (1908). sule bubble-shaped with a tube-shaped astropyle in
l the center of an operculum, with a wall seemingly
Radiolaria Phaeodaria
• Aulacantha cannulata Haeckel (Fig. 6.10). Radial

229
I of two layers. Intracapsular protoplasm mainly in tubes cylindrical in the proximal half, prismatic in
front of the operculum where it forms a disc, from the distal half, with 3 to 6, usually 4, prominent,
which a cord stretches to the opposite pole of the thick, parallel edges, which are dentate towards the
central capsule. Here it unravels "fountain-like" thicker distal end; their teeth short, conical, direct-
and forms a net under the central capsule wall with ed outwards, scarcely one fourth as long as the
its branches. Inconspicuous skeleton an irregular thickness of the tube. The edges are separated in
meshwork of silica needles, funnel shaped, at the the distal third by more or less deep furrows, like a
oral pole of the central capsule. Diameter: 0.30- channelled column. Length of tubes: 1.2-2.5 rnm.
0.50 rnm. Ref.: Schróder (1913). Ref.: Haeckel (1887).
• Halocella magna Schroder (Fig. 6.8). Central • Aulacantha scolymantha scolymantha Haeckel
capsule bubble-shaped, spherical, with well-devel- (Fig. 6.12). Radial tubes cylindrical, straight, of
oped operculum. Central capsule wall of two nearly equal breadth throughout their entire
membranes, showing afine pleated folding. In- length, somewhat inflated in the proximal basal
conspicuous skeleton an irregular meshwork of end, smooth throughout the greater part of their
silica needles, funnel shaped, at the oral pole of the length, but in the distal part dentate, with numer-
central capsule. Differs from H gemma mainly in ous (10 to 40) small pointed teeth, which are
its larger size and more conspicuous operculum. curved outwards, and shorter than the thickness of
Also similar to Cornucella maya, with which it the tube. A. scolymantha is the cornmonest aula-
may be conspecific. Diameter: about 1 rnm. Ref.: canthid, and perhaps the cornmonest phaeodarian,
Schróder (1913). species. Length of tubes: 0.5-1.0 mm, Ref.:
• Lobocella proteus Borgert (Fig. 6.9). Central cap- Haecker (1908), Reshetnjak (1966).
sule bubble-shaped, with a variable number of • Aulacantha scolymantha bathybia Haecker (Fig.
protuberances, with astropyle and significant 6.11). Differs from the nominate subspecies by
operculum. Soft parts and nuc1eus as in Halocella. being larger, with correspondingly somewhat long-
Skeleton unknown. Diameter: about 1 rnm. Ref.: er and thicker spines. Length oftubes: 0.7-1.5 rnm.
Schróder (1913). Ref.: Haecker (1908), Reshetnjak (1966).
• Auloceros arborescens birameus Irnmerman (Fig.
Family Aulacanthidae 6.13). Cylindrical radial tubes with 2 (seldom 3)
1 Distal end of radial spines with lateral branches .2 long terminal branches inserted at almost 90
la Distal end ofradial spines smooth or spiny, no lat- degrees, bifurcating twice. Spathillae usually
eral branches: Aulacantha absent or poorly developed. Body spherical, with
2 Radial spines with terminal branches only, no lat- 1 central capsule. Diameter: 1.8-3.0 rnm. Ref.:
eral branches 3 Reshetnjak (1966), Haecker (1908).
2a Radial spines with terminal and lateral branches: • Auloceros arborescens dichodendrum Immerman
Aulospathis (Fig. 6.14). Radial tubes nearly cylindrical, taper-
3 Terminal branches usually undivided (may be ing distally where wall thickness increases, with
bifurcated only) 4 corona of 2-3 terminal branches, which branch
3a Terminal branches of radial spines further divided again 1-3 times at acute angles, with terminal spa-
.............................................................................. 5 thillae. Length of tubes: 0.75-1.90 rnm. Diameter:
4 Terminal branches without lateral teeth: Aulogra- 3.0-4.5 rnm. Ref.: Tibbs (1976).
phis • Auloceros arborescens irregularis Haecker (Fig.
4a Terminal branches with lateral teeth: Aulograph- 6.15). Radial tubes nearly cylindrical, tapering
onium distally where wall thickness increases. Corona of
5 Radial spines with foreign objects (diatom frus- 3 short, strongly divergent terminal branches,
tules) inc1uded within: Aulokleptes which branch 3 times into secondary branches,
5a Radial spines without foreign objects (diatom with spathillae. Diameter: 2.0-3.5 rnm. Ref.: Tibbs
frustules) inc1uded within: Auloceros (1976).
· ¡:
• Auloceros arborescens subelegans Haecker (Fig. tubes: 0.5-1.5 mm. Ref.: Reshetnjak (1966), Tibbs
6.16). Radial tubes nearly cylindrical, tapering (1976).
distally where wall thickness increases, with coro- • Aulographis stellata Haeckel (Fig. 6.25). Radial
na of2-4 terminal branches,which branch again 1- tubes 60-70, cylindrical to club-shaped, in the lat-
2 times at obtuse angles, with small terminal spa- ter case gradually thickened towards the distal end.
thillae. Diameter: 2.0-3.0 mm. Ref.: Tibbs (1976). Terminal branches stellate, numerous (5 to 10 or
• Auloceros arborescens trigeminus Haeckel (Fig. more), diverging in all directions, smooth, straight
6.17). Radial tubes club-shaped, gradually thick- or slightly curved, slenderly conical, 3 to 5 times as
ened toward the inflated distal end. Terminal long as the distal breadth of the tube. Branches
branches short, scarcely longer than tube width, 2 very variable in size, sometimes very small. Length
constantly opposite, each with 3 equal, conical, of tubes: 0.4-0.8 mm. Ref.: Haecker (1908),
short secondary branches. No terminal spathillae. Reshetnjak (1966).
Length oftubes: 0.6-0.8 mm. Ref.: Haeckel (1887), • Aulographis tetrancistra Haeckel (Fig. 6.26).
Reshetnjak (1966). Radial tubes slender, cylindrical to slightly tapered
• Aulographis bicurvata Haecker (Fig. 6.21). Cylin- distally, straight. Terminal branches 4 to 6 (usually
drical radial tubes with 3-4 simple terminal branch- 4) 8 to 10 times as long as the tube is broad, slight-
es inserted at almost 90 degrees, distal ends curving ly curved and widely divergent, with a terminal
backwards, ending in spathillae. Body spherical to spathilla of 4 crossed recurved teeth. Length of
ovoid, with 1-8 central capsules. Diameter: 1.5-3.0 tubes: 0.5-0.7 mm. Ref.: Reshetnjak (1966), Tibbs
mm. Ref.: Haecker (1908), Reshetnjak (1966). (1976).
• Aulographis furcula Haeckel (Fig. 6.22). Radial • Aulographis tetrastyla Haecker (Fig. 27). Almost
tubes cylindrical, slender, slightly curved. Terminal cylindrical radial tubes, with 3-4 long terminal
branches usually 2, sometimes on single tubes 3, 8 branches at 120 degrees; terminal branches ending
to 10 times as long as the tube is broad, strongly in 2 small spines. Body kidney-shaped, with 2
curved, ascending vertically from a horizontal base, central capsules. Diameter: 2.5-3.5 mm. Ref.:
with a terminal spathilla of 4 crossed teeth at the Haecker (1908), Reshetnjak (1966).
distal end. Diameter: 1.2-2.0 mm. Ref.: Reshetnjak • Aulographis triangulum Haeckel (Fig. 6.28).
(1966), Tibbs (1976). Radial tubes slender, cylindrical, straight. Terminal
• Aulographis mohri Tibbs (Fig. 6.23). Skeleton of branches constantly 3, slender, slightly curved,
well-developed lenticular veil of tangential nee- nearly tangentially divergent, 8 to 10 times as long
dles and of several hundred radial tubes. Tubes as tube width, with a terminal spathilla of 4
small, slender, thin-walled, slightly tapered distal- recurved teeth in opposite pairs. Angles between
ly, each with distal crown of 2 to 4 (generally 3) branches equal. Length of tubes: 1.2 mm. Ref.:
terminal branches, curved outward and upward. Haeckel (1887), Reshetnjak (1966).
Distal end of each terminal branch with compara- • Aulographonium antarcticum Haecker (Fig. 6.29).
tively large, strongly silicified, cushion-like spa- Cylindrical radial tubes with 11-15 terminal branch-
thilla with 4 to 8 (generally 6 to 8) strong recurved es, central 1-3 branches short, straight, with tiny lat-
teeth. Some spathillae with short, stout apical pro- eral spinules, remaining peripheral branches longer,
cess. Length: 1.2-1.4 mm. Width: 0.2-0.3 mm. curved, surface almost totally smooth. Observed
Ref.: Tibbs (1976). only as isolated spines. No dimensions given. Ref.:
• Aulographis pandora Haeckel (Fig. 6.24). Radial Haecker (1908), Reshetnjak (1966).
tubes cylindrical to somewhat tapered distally, slen- • Aulographonium anthoides Haecker (Fig. 6.30).
der, straight or slightly curved. Terminal branches Radial tubes straight, cylindrical, somewhat tapered
of the tubes very variable in size and number (2 to distally, with crown of 4-17 terminal branches in
6, usually 3 to 4), without lateral teeth or terminal one or two whorls. Whorls sometimes around 1-3
spathillae, slender, conical, slightly curved, directed central branches, or central branches may be absent.
outwards, 2 to 4 times (rarely 5 to 8 times) as long Terminal branches sometimes dentate. Tube length:
as the breadth of the tubes. The number and size of 1.2-1.65 mm. Ref.: Tibbs (1976).
the simple terminal branches is extraordinarily var- • Aulographonium bicorne Haecker (Fig. 6.31).
iable, even within the same specimen. Length of Cylindrical radial tubes with 6-7 stout, almost
straight terminal branches, two branches central, es. Branches simple or branched, or covered with
others peripheral. All branches with irregularly numerous irregular, stout, solid thorns. Foreign
spiny surface and terminal spathillae. Diameter: siliceous detrital bodies (usual1y diatom frustules)
3.0-3.5 rnrn. Ref.: Haecker (1908), Reshetnjak arranged axially within radial tubes (a generic
(1966). character) (Fig. 6.20a). Veil of tangential needles
• Aulographonium candelabrum (Haecke1) (Fig. replaced largely by loose network of siliceous
6.32). Radial tubes c1ub-shaped, straight, thick- detritus. As many as 4 varieties or subspecies,
ened distally and constricted beyond the ovate, possibly distinct warm- and cold-water forms are
inflated, terminal knob. Knob similar to a candela- discussed by some authors. A. r. varo longispina
brurn with a corona of 6 to 9 strongly curved ter- Irnrnermann (a simple form with 2, 3, or more
minal branches, which are 8 to 10 times as long as smooth terminal teeth, not necessarily from a
the tube width, with scattered lateral dentic1es, and common origin), and A. r. varo antarctica
with a spathilla of 5 to 7 radial teeth. Length of Haecker (with robust spines and short branches),
tubes: 1.6-2.4 mm. Ref.: Haeckel (1887), both reported from the South Atlantic, are illus-
Reshetnjak (1966). trated. Length of tubes: 1.2-1.8 rnrn. Ref.: Tibbs
• Aulographonium mediterraneum (Borgert) (Fig. (1976).
6.33). Radial tubes almost cylindrica1 with 3-6 • Aulospathis pinus Haecker (Fig. 6.35). Radial
(usually 4) long terminal branches, one branch tubes cylindrical, tapering proximally and distally,
central, stouter, longer and straighter than others, sometimes somewhat curved and undulant in dis-
remaining peripheral branches with curved distal tal region of 3-12 subtenninal lateral branches.
ends. AlI branches smooth or covered with small Lateral branches usually in whorl, size decreasing
spinules, ending in spathillae. Diameter: 1.8-2.0 distally. Usually with 3 terminal branches. Thin
rnrn. Ref.: Reshetnjak (1966), Tibbs (1976). peripheral veil of tangential needles. Diameter:
• Aulographonium pulvinatum (Haeckel) (Fig. 2.2-3.5 rnrn. Ref.: Tibbs (1976).
6.34). Radial tubes club-shaped to cylindrical, • Aulospathis variabilis aulodendroides Haecker
straight, gradually thickened toward the distal end, (Fig. 6.36). Radial tubes more or less spindle-
'which bears a broad, circular, convex bulge. shaped, tapering toward both extremes, distal end
Margin of bulge bears 2 alternating verticils of with 4-9 terminal branches, without inflated termi-
radially divergent, straight terminal branches, nal knob. Proximal branches tend to be numerous,
which are 2 to 3 times as long as the tube is wide. not in whorl, but in a large stretch along the shaft.
Eachbranch with 2 opposite lateral rows of Length oftubes: to 2.5 rnrn. Ref.: Haecker (1908).
numerous small dentic1es, and bearing a terminal • Aulospathis variabilis bifurca Haeckel (Fig. 6.37).
spathilla with 6 to 8 recurved radial teeth. Length Radial tubes inflated ovate to spindle-shaped,
of tubes: 2.0-2.4 rnrn. Ref.: Haecker (1908), becoming very thin proximally, with inflated termi-
Reshetnjak (1966). nal knob with 2 (often a few with 3 or 4) divergent,
• Aulokleptes flosculus (Haeckel) (Fig. 6.20). Radial inwardly curved, hollow, terminal branches. Whorl
tubes c1ub-shaped, straight, tapered proximally, of 4 (often 5 or 6, more rarely 2 or 3) irregularly
each with 9-12 stout conical terminal branches crossed lateral branches, at about two thirds dis-
which are studded with thorns. Foreign siliceous tance from proximal end, with spathillae of 4-6
detrital bodies aligned axial1y within radial tubes (a curved teeth, sometimes also with a central tooth.
generic character). As many as 5 varieties or sub- Length of tubes: 1.5-2.5 rnrn. Ref.: Irnrnerman
species, possibly distinct warm- and cold-water (1904), Reshetnjak (1966).
forms are discussed by some authors. Note frustule • Aulospathis variabilis diodon Haeckel (Fig. 6.38).
of the diatom genus Rhizosolenia serving as a Radial tubes without inflated terminal knob,
basis for deposition of the spine shaft (a generic forked at the tapering distal end, with 2 divergent
character) in Fig. 6.20a. Diameter: 3.0-5.5 rnrn. curved branches of equal size. Proximal whorl
Ref.: Reshetnjak (1966), Tibbs (1976). cruciate, with 4 rather equal, crossed, straight, lat-
• Aulokleptes ramosus Immermann (Fig. 6.18, eral branches (sometimes 3 or 5 instead of 4 in sin-
6.19). Radial tubes c1ub-shaped, straight, pro xi- gle tubes). Length of tubes: 2.0-2.5 rnrn. Ref.:
mal1y tapered, with crown of 2-5 terminal branch- Haeckel (1887).
1[;.
• Aulospathis variabilis triodon Haeckel (Fig. poleo Radial tubes about as long as tangential bars.
6.39). Radial tubes somewhat spindle-shaped, Distal ends of tubes with whorl of 3-6 branches.
without inflated terminal knob, with 3 divergent, Terminal branches of distal whorl somewhat short-
slightly curved, terminal branches. Proximal er than those of inner whorl. Branches forked, with
whorl with variable number (usually 3, often 2 or spiny terminal pads. Proximal radial tubes some-
4, rarely more) of curved lateral branches. Tube times with one or a few lateral thoms. Length: 3-10
walls very thin. Length oftubes: 1.8-3.0 rnrn. Ref.: rnrn. Ref.: Haeckel (1887), Reshetnjak (1966).
Haeckel (1887), Reshetnjak (1966). • Auloscena pyramidalis Haeckel (Fig. 6.44).
Spheroidal lattice shell with triangular mesh open-
Family Aulospbaeridae ings. Radial tubes about as long as tangential bars,
1 Without pyramidal elevations 2 with crown of 3-6 divergent, straight or slightly
la With pyramidal elevations 3 curved, pointed terminal branches. Remainder of
2 With polygonal (heptagonal or hexagonal) mesh- each tube smooth or with a few lateral thoms.
es: Aulastrum Diameter: 2.5-4.0 rnrn. Ref.: Haecker (1908),
2a With triangular (more seldom quadrangular) mesh- Reshetnjak (1966).
es, spindle shaped: Aulatractus • Auloscena robustissima Haecker (Fig. 6.45).
2b With triangular (more seldom quadrangular) mesh- Cylindrical radial tubes large, massive, thick-
es, spherical to pyriform: Aulosphaera walled, with scattered spines on the surface, ending
3 With triangular (more seldom quadrangular) mesh- in 3-8 short, pointed terminal branches. Diameter:
es, spherical forms: Auloscena 3.5-4.0 rnrn. Ref.: Haecker (1908), Reshetnjak
(1966).
• Aulastrum mirabile Borgert (Fig. 6.40). Radial • Auloscena verticillus Haeckel (Fig. 6.46, 6.47).
spines about 3 times as long as the smooth tangen- Spheroidal to ovoidal lattice shell with triangular
tial bars, distally with about 10 whorls of usually mesh openings. Radial tubes 1-3 times as long as
8 branches with simple small buttons or small pyramidal bars, with crown of 12-30 recurved,
crowns of short prongs, proximal half with irregu- pointed, terminal branches, partially fused at base.
1ar1y arranged, pointed thoms. Diameter: 1.8-2.8 Distal subcoronal region ofhook-like lateral thoms,
rnrn. Ref.: Borgert (1904), Haecker (1908). sometimes in irregular verticils. Sometimes a few
• Aulastrum monoceros Haeckel (Fig. 6.41). Radial single, curved thoms on proximal part of tubes.
tubes simple, smooth, straight, cylindro-conical, Diameter: 1.85-3.75 rnrn. Besides the nominate sub-
about as long as the smooth tangential tubes or species (named A. v. typica by Haeckel, Fig. 6.46),
somewhat longer. Mesh openings irregularly two subspecies have been reported from the South
polygonal, usually pentagonal or hexagonal. AtIantic: A. v. hamata Borgert (12-25 terminal
Diameter: 3.0-4.0 rnrn. Ref.: Haeckel (1887). branches, 6-8 subcoronallateral thoms; Fig. 6.47);
• Aulastrum spinosum Borgert (Fig. 6.42). A. v. laevis Haecker (12-16 terminal branches, 3-8
Spheroidal lattice shell with subregular, generally subcoronallateral thoms). Diameter: 1.85-3.75 rnrn.
pentagonal or hexagonal mesh openings. Radial Ref.: Haecker (1908), Reshetnjak (1966).
tubes 2 to 5 times length of tangential bars. • Aulosphaera bisternaria Haeckel (Fig. 6.49).
Proximal half to two thirds of radial tubes with sin- Spherical, ovoidal, pear- or spindle-shaped shell,
gle or paired, irregularly distributed, curved, lateral with triangular mesh openings of various sizes.
thoms. Distal thoms usually in 1 or 2 loose whorls, Radial tubes about as long as tangential bars,
3 teeth in terminal position. Sometimes one or a few sometimes longer on one pole, proximal portions
thin, straight, centrifugal needles on tangential bars. with one or a few single lateral thoms, distal por-
Diameter: 1.4-1.8 mm. Ref.: Haecker (1908), tions with 2 whorls of 2-6 branches, sometimes
Reshetnjak (1966). distally forked, with spathillae. Terminal whorl
• Aulatractus fusiformis Haeckel (Fig. 6.43). with shorter branches, inner whorl branches about
Lattice shell generally spindle-shaped, with both twice as long, branches ofthe two whorls altemate
poles rounded at ends or with one end more point- positions. Diameter: 1.8-5.5 rnrn. Two subspecies
ed than the other, with triangular mesh openings. of this species have be en reported from the South
Tangential bars considerably stouter on rounded Atlantic: A. b. ovulum Borgert (whorls widely sep-
arated, 2-4 branches per whor1; Fig. 6.49b) is con- gential bars, distally with 3 whorls of 4 to 6 equidis-
sidered a warm-water species, and A. b. septen- tant, nearly perpendicular lateral branches.
trionalis Borgert (whorls narrowly spaced, 3-6 Diameter: as large as 3.0 by 3.5 mm. Ref.: Haecker
branches per whorl; Fig. 6.49c) is normally report- (1908), Reshetnjak (1966).
ed from the Antarctic. Diameter: 1.8-5.5 mm. Ref.: • Aulosphaera verticillata Haeckel (Fig. 6.55).
Haecker (1908), Reshetnjak (1966). Shell spheroidal with regular triangular mesh
• Aulosphaera elegantissima Haeckel (Fig. 6.50). openings. Radial tubes slender, straight, distally
Shell spheroidal with fair1y regular triangular tapered, about as long as tangential bars, each with
mesh openings. Radial tubes about 2 times as long 6-12 whorls of 3 or 4 curved simple lateral spines
as tangential bars, distal third very thin and with 8 without spathillae. Haecker (1908) differentiated a
to 12 whorls of 4 lateral branches, forked near the larger Antarctic form with few whorls of lateral
base to appear' as 8 branches. Diameter: 1.5-2.0 spines (Fig. 6.55b) that has been reported also by
mm in equatorial samples; 2.5-3.0 mm in Peters (1929) and Tibbs (1969) from Antarctic
Antarctic samples. Ref.: Haecker (1908). localities. Diameter: 1.5-2.5 mm. Ref.: Haecker
• Aulosphaera filigera Haecker (Fig. 6.48). Lattice (1908).
shell ellipsoidal, with equilateral triangular mesh
openings. Three spathillae-bearing filaments on Family Cannosphaeridae
the bars. Radial spines as long as bars, with later- • Cannosphaera antarctica Haeckel (Fig. 6.56).
al thoms and 3-4 terminal branches with buttons. Inner shell mammillate, with 15 to 20 radial rods
Length: 4.0-4.5 mm. Ref.: Schróder (1913). with numerous groups of 4 delicate, terminally
• Aulosphaera labradoriensis Borgert (Fig. 6.51). dentate lateral spines. Outer shell with irregular
Spheroidallattice shell with triangular mesh open- pentagonal or hexagonal mesh openings, with
ings. Radial tubes about as long as tangential bars, numerous scattered forks of 2 or 3 delicate, diver-
2-5 whorls of 3-5 curved lateral branches on distal gent, terminally dentate spines, and with 20 to 30
half of tubes, terminal whorl of 2-4 (usually 3) larger simple radial spines, about half as long as the
short, curved branches, paired lateral branches internal rods, distally with verticil of 3 to 5 slightly
sometimes on proximal half of tubes, lateral curved, simple, terminal branches. Diameter: 1.0-
branches pointed or rounded. Diameter: 1.2-2.7 2.2 mm. Ref.: Haeckel (1887).
mm. Ref.: Haecker (1908), Reshetnjak (1966).
• Aulosphaera robusta Haecker (Fig. 6.52). Shell Family Castanellidae
ovate, with triangular mesh openings. Stout radial 1 Without main spines .2
tubes with brownish thick walls, about as long as la With main spines 3
tangential bars, distally with 2 whorls of 2 to 5 l c With or without main spines 5
(gene rally 3) branches with terminal spathillae. 2 Mouth smooth: Castanarium
Branches in inner whorl slightly longer, very 2a Mouth with teeth: Castanella
slightly curved; in outer whorl shorter, generally 3 With simple main spines .4
arranged in a crown. Diameter: one whole speci- 3a With branched main spines: Castanopsis
men reported as 3.0 by 3.5 mm. Ref.: Haecker 4 Mouth smooth: Castanidium
(1908). 4a Mouth with teeth: Castanissa
• Aulosphaera triodon Haeckel (Fig. 6.53). 5 Mouth with thickened rim or low protuberances:
Spheroidal to ellipsoidal shell with triangular Castanea
(sometimes square) mesh openings. Radial tubes
generally smooth, about as long as tangential bars, • Castanarium antarcticum Haecker (Fig. 6.57)
with 2-4 (rarely 3, seldom 4) straight or slightly Thick-walled, yellowish shell with simple, circular
curved terminal branches, third to fifth as long as pores of irregular size, about 2-3 times bar-width.
radial tubes. Diameter: 1.2-4.0 mm. Ref.: Haeckel Mouth small, with smooth rim. By-spines about 2
(1887), Reshetnjak (1966). times as long as the average pore-diameter.
• Aulosphaera trispathis Haecker (Fig. 6.54). Shell Diameter: 0.55-0.85 mm. Ref.: Haecker (1908),
spheroidal to ellipsoidal with regular triangular Reshetnjak (1966).
mesh openings. Radial tubes about as long as tan-
• Castanariumfavosum Haecker (Fig. 6.58). Thick- slender by-spines with tiny bulb-like expansions
walled spherical shell, with circular to subcircular at the base. Diameter: 0.8 rnm. Ref.: Schróder
pores, hexagonal pore frames, thick bars, smooth (1913).
mouth, numerous thick, short, by-spines as long as • Castanella maxima Schmidt (Fig. 6.64). Thick-
pore diameter. Diameter: 0.50-0.75 rnm. Ref.: walled spherical shell, with nearly circular pores,
Haecker (1908), Reshetnjak (1966). slightly wider than bars, thick bars. Small mouth
• Castanarium hookeri Haeckel (Fig. 6.59). Very (about 3 times pore width), with 4 to 5 dagger-
thick-walled spherical, yellowish shell, with small shaped teeth, about 4 to 5 times as long as pore
(most about 2 times bar width) circular pores. width. Numerous by-spines, about 3 times as long
Hexagonal pore frames result in funnel-shaped as pore width. Diameter: 0.9-1.0 rnm. Ref.:
pores. Small (about 2 times pore diameter), circu- Schróder (1913).
lar mouth, surrounded by a ring of ordinary spines. • Castanella thomsoni Haeckel (Fig. 6.65, atypical
Spines of variable size (to about 2 times pore specimen with 2 pylomes). Large, moderately
diameter), but not differentiated into main- and thick-walled spherical shell, with circular to sub-
by-spines. Diameter: 0.55-1.0 rnm. Ref.: Haecker circular (rarely polygonal), small pores, 2-3 times
(1908). thickness ofbars, sometimes incompletely framed,
• Castanea amphora Haecker (Fig. 6.61). Large, with very numerous, long (about 2 times pore
thick-walled spherical (rarely ellipsoidal) shell, diameter) by-spines. Mouth small, circular, with
with small (about 2-3 times bar width) circular or 5-7 (rarely 3-10) uniform, robust, conical, long
sub circular to subpolygonal pores, often partially (twice length of by-spines), convergent teeth.
framed, wide bars. Circular mouth, with thick rim, Three subspecies were distinguished by Haecker
elevated as a collar- or crater-like structure. Short (1908) and one by Tibbs (1969), and two forms
(2/3 shell diameter) main-spines. Numerous short were recognized by K1ing (1966), based mainly on
(2 times pore diameter) by-spines. Diameter: 0.9- size and wall thickness. Polar forms are generally
1.3 rnm. Ref.: Haecker (1908), Reshetnjak (1966). larger. Diameter: 0.3-1.1 mm. Ref.: Haecker
• Castanea globosa Haecker (Fig. 6.60). Large, (1908), Reshetnjak (1966).
thick-walled spherical shell, with small (about 2 • Castanidium haeckeri Schróder (Fig. 6.71).
times bar width) subpolygonal (often trigonal) to Spherical to nearly ellipsoidal shell, with circular
subcircular pores, often partially framed, wide bars. to oval pores (about 2 times bar width). Small cir-
Very small, oval to irregular mouth, with thick rim, cular mouth with thick rim and 1-2 main-spines:
usually with one main-spine, sometimes with low Main-spines thick, about half shell radius, curved
elevations. Short, thick, main-spines, lacking in or flame-shaped. Numerous by-spines, about 2
some specimens. Numerous short (1-2 times pore times pore width. Diameter: 1.10-1.25 rnm. Ref.:
diameter) by-spines. Diameter: 0.6-1.25 rnm. Ref.: Schróder (1913).
Haecker (1908), Reshetnjak (1966). • Castanidium longispinum Haecker (Fig. 6.72).
• Castanea henseni Borgert (Fig. 6.62). Large, Thin-walled shell basically spherical, with eleva-
thick-walled spherical shell, with small (1-3 times tions at base of main-spines suggesting polyhedral
bar width) subcircular (elliptical) pores, wide bars. shape, with small (4-6 times bar width) circular to
Small mouth with thick rim, with blunt protuber- sub circular or polygonal pores. Bars often rectan-
ances projecting radially and tangentially into the gular in cross section, giving pores cylindrical
mouth opening. Short (half shell radius) main shape, especially in smaller specimens. Mouth
spines. Numerous short (2-3 times pore diameter), variable in size, usually with thin but distinct rim
thick by-spines. Diameter: 0.65-0.9 rnm. Ref.: and 1-3 main-spines. Main-spines long (longer
Haecker (1908), Reshetnjak (1966). than diameter), elevated and fenestrated at base.
• Castanella borgerti Schroder (Fig. 6.63). Thick- Diameter: 0.25-0.65 rnm. Ref.: Haecker (1908),
walled, yellowish spherical shell, with circular to Reshetnjak (1966).
oval pores, slightly wider than bars. Small mouth • Castanidium moseleyi Haeckel (Fig. 6.73). Very
(about 2 times pore width), with thin teeth, with large, thin-walled shell, with conical protuberanc-
tiny perpendicular branches at about mid-length, es at main-spine bases suggesting polyhedral
suggesting incomplete walls of basal pores. Very shape, with variable (2-8 time bar width) circular
to subpolygonal pores, narrow bars. Wide mouth • Castanissa schmidti Schróder (Fig. 6.69). Shell
with 1-3 main-spines, Main-spines slender, thin-walled, balloon-shaped, somewhat tapered
straight , and smooth, to thick, wavy and sculp- toward the mouth, with small irregularly circular
tured, about as long as shell diameter. Sparse by- to oval pores, about 3 times bar width, weakly
spines, 1-4 times pore diameter. Diameter: 0.725- framed. Mouth small, about 3 times pore width,
1.1 rnm. Ref.: Haecker (1908). with 7 strong spines, fused at their based into a
• Castanidium spinosum Schróder (Fig. 6.74). ring. Diameter: 0.75 rnm. Ref.: Schroder (1913).
Spherical, thick-walled, yellowish shell with sub- • Castanissa valdiviae Haecker (Fig. 6.68). Thick-
circular, irregular pores, almost twice as large as walled spherical shell, with small circular pores
bar width, with inconspicuous funnel-shaped 5 to about 1-2 times bar width. Mouth relatively small,
6-sided frames. Mouth relatively large, with 5-6 with a ring ofteeth and 1-3 main-spines, ofwhich
main-spines. Main-spines as long as shell diame- one is sculptured in a net-like pattem. Thick main-
ter. By-spines about twice as large as pore diame- spines, length about half the shell radius.
ter. Diameter: 0.53-1.2 mm. Ref.: Schróder Numerous by-spines, about 3 times pore width.
(1913), Reshetnjak (1966). Diameter: 0.8-1.0 rnm. Ref.: Schróder (1913).
• Castanidium vanhoeffeni Schróder (Fig. 6.75). • Castanopsis fragilis Haecker (Fig. 6.70). Thin-
Very thick spherical shell, with large circular to walled shell, weakly polyhedral, with subcircu1ar
oval pores, 1-2 times bar width, framed. Circular to oval pores, about 3-6 times bar width, wide
mouth (about 3 times pore width) with 2 short bars. Wide mouth, with a few rnain-spines. Main-
conical main-spines. Main-spines fairly sparse, spines on pyramids of 3-4 bars, simple, forked or
relatively short (about 4 times pore width), some weakly branched at the tips. By-spines about as
slightly curved. By-spines long (about 3 times long as pore width. Diameter: 0.7-0.8 rnm. Ref.:
pore width) and slender. Diameter: 0.7 rnm. Ref.: Schróder (1913).
Schróder (1913).
• Castanidium variabile Borgert (Fig. 6.76). Small, Family Challengeridae
moderately thick-walled spherical shell, with cir- Various authors have revised individual challengerid
cular to sub circular or subpolygonal large pores, genera and/or transferred species from their original
variable in size, 4-5 times width of bars, rarely genera. However, these piecemeal efforts have some-
weakly framed. Mouth average to fairly large, times resulted in conflicting generic criteria, and a
with 1-3 main-spines. Diameter: 0.7-1.0 rnm. Ref.: comprehensive natural c1assification of the entire
Haecker (1908), Reshetnjak (1966). family is still needed. We attempt in the generic diag-
• Castanissa nationalis Schmidt (Fig. 6.66). noses below to use non-overlapping criteria that are
Spherical shell, with circular funnel-shaped pores, consistent with both the original definitions and the
about as wide as bar width, framed. Mouth small, observed characters of the species discussed. It is pro-
circular, with 4-6 stout conical teeth, about same visional for the purposes of this review, and should
length as by-spines, with one flarne-shaped main- not be interpreted as proposing emendation of any of
spine, longer and thicker than other main spines, the genera treated.
often sculptured. Main-spines fairly straight, slen-
der, length about two fifths shell diameter. By- 1 Shell without large fenestrae near peristome 2
spines about two and one half times pore width. la Shell with 2 large fenestrae near peristome, 2 oral
Diameter: 0.33-0.53 rnm. Ref.: Schróder (1913). teeth on peristome, 1 or more marginal spines:
• Castanissa proteus Schmidt (Fig. 6.67). Fairly Challengeranium
thick-walled spherical shell, with irregularly cir- 2 Margin of shell smooth or with 1 or 2 isolated
cular pores, weakly framed. Mouth fairly small, teeth 3
about 3 times pore width, with teeth about as long 2a Margin of shell with row of numerous marginal
as by-spines, and 1-2 main-spines. Main-spines spines, with oral teeth: Challengeron
relatively thin, shorter than shell radius. Very 3 With 1 oral tooth, no marginal spines: Challengeria
numerous by-spines, about 3 times pore width. 3a With 2 or more oral teeth, margin smooth or with
Diameter: 0.7-1.0 rnm. Ref.: Schróder (1913). 1 or 2 isolated teeth: Protocystis
,
1,
• Challengeria naresi Murray (Fig. 6.78). Lenticular shaped or half conical, longer than shell radius,
shell, circular or ovate in lateral view, with thick with 2 pairs of conical, curved, divergent teeth.
wall. Peristome with simple tooth, about as long as Terminal teeth are the halves of a forked dorsal
radius, with 2 lateral basal ridges. Length: 0.5-0.65 tooth, diverging laterally. Lower teeth nearly hor-
rnm. Ref.: Haecker (1908), Reshetnjak (1966). izontal and directed ventrally. Diameter: 0.13-0.15
• Challengeria tritonis Haeckel (Fig. 6.79). Shell rnm. Ref.: Haeckel (1887), Takahashi and Honjo
ovate, slight1y compressed. Peristome with single (1981).
simple tooth, weakly S-shaped, more or less • Protocystis acornis Haecker (Fig. 6.84). Shell
curved, about third as long as shell. Diameter: ovate, thick-walled, slightly compressed laterally.
0.15-0.2 rnm. Ref.: Haeckel (1887). Peristome oblique, collar-like, with process
• Challengeria xiphodon Haeckel (Fig. 6.80). Shell extending into 3 teeth, of which middle one is
spheroidal, sometimes slightly compressed later- largest. Length: 0.07-0.08 rnm. Ref.: Haecker
ally, in lateral view circular to slightly oval. (1908).
Peristome with single, simple vertical tooth. Tooth • Protocystis balfouri (Murray) (Fig. 6.85). Shell
straight, triangular, trowel-like, length about shell lenticular, strongly compressed laterally, subcircu-
diameter. Diameter: 0.09-0.13 rnm. Ref.: Haecker lar in lateral view, aboral margin with 2 short
(1908). divergent spines, one dorsal, one ventral.
• Challengeranium diodon (Haeckel) (Fig. 6.77). Peristome vertical, slender, semi-tubular, about as
Shell ovoid, slight1y compressed, with one large long as shell radius, distal half forked, extending
conical spine and sometimes a few smaller spines into 2 sub-parallel teeth. Diameter: 0.08-0.2 rnm.
on aboral pole, half as long as radius. Peristome Ref.: Borgert (1911).
short and wide, collar-shaped, about twice as wide • Protocystis bicornuta Schróder (Fig. 6.86). Shell
as long, half as long as radius, obliquely inclined thick, spheroidal, sometimes slightly compressed
over mouth, with 2 divergent straight teeth, conical laterally, with 2 strongly divergent spines on abo-
and longer than shell radius. Base of each tooth ral margino Peristome somewhat oblique, extend-
with large ovate pore in wall of peristome. ing into 2 short lateral teeth and 1 medial tooth.
Diameter: 0.08-0.10 rnm. Ref.: Haeckel (1887). Length (including process): 0.1 mm. Ref.:
• Challengeron channeri Murray (Fig. 6.82). Shell Schróder (1913).
subcircular, lenticular, strongly compressed, with • Protocystis bicuspis Schróder (Fig. 6.87). Shell
14 to 18 slender, straight, rarely slightly curved, ovate, with 2 thick, parallel, median spines at the
radial spines on margino Spines straight, equidis- aboral poleo Peristome straight, collar-like, per-
tant, cylindrical, about as long as shell radius. pendicular to main axis of shell, with process
Peristome short, with 2 parallel, vertical, lateral, extending into 3 teeth, of which the middle one is
straight teeth, less than half length of spine. longest. Length: 0.16 rnm. Ref.: Schróder (1913).
Diameter: 0.14-0.17 rnm. Ref.: Haeckel (1887). • Protocystis harstoni (Murray) (Fig. 6.88). Shell of
• Challengeron radians Borgert (Fig. 6.81). Shell variable shape, compressed laterally; margin sub-
lens-shaped, compressed, nearly circular in side circular to somewhat triangular in lateral view.
view, with 18-26 equally spaced, radial teeth on Peristome oblique to vertical, with process extend-
the margin, about equal in length. Marginal spines ing into 2 parallel to sub-parallel teeth. Length:
conical, straight, scarcely third to at most half as 0.125-0.133 rnm. Ref.: Schróder (1913).
long as shell radius. Peristome short, channel- • Protocystis heterocentrota Peters (Fig. 6.89).
shaped, with 2 lateral, parallel, pointed teeth, ver- Shell spheroidal to ovoid, slight1y compres sed lat-
tical or slightly inclined ventrally. Peristome with erally, with 2 divergent spines on aboral margin,
teeth usually somewhat longer than the marginal one large dorsal, one smaller ventral. Distal end of
spines. Diameter: 0.13-0.15 rnm. Ref.: Borgert peristome process expanded into 3 small very
(1904). weakly developed teeth. Length (including spines
• Challengeron willemoesii Haeckel (Fig. 6.83). and process): 0.12-0.14 rnm. Ref.: Peters (1929).
Shell broadly ovate to nearly circular in lateral • Protocystis macleari (Murray) (Fig. 6.90). Shell
view, lenticular, with a variable number of margi- subcircular to subtriangular, lenticular, with sharp
nal spines of variable shape. Peristome helmet- marginal edge. Peristome semi-tubular, oblique
half cylinder, with 3 divergent, short, triangular flange-like teeth. Length (including process):
teeth. Dorsal odd tooth nearly vertical, paired lat- 0.15-0.18 rnm. Ref.: Peters (1929).
eral teeth sub-horizontal. Peristome about third to • Protocystis tridens Haeckel (Fig. 6.97). Shell
fourth as long as shell diameter. Diameter: 0.25- ovoid, slightly compressed laterally, thin-walled.
0.35 rnm. Ref.: Haeckel (1887). Peristome oblique and collar-like, extending into 3
• Protocystis micropelecus Haecker (Fig. 6.91). teeth, with middle tooth largest. Length: 0.07-0.15
Shell stout, thick-walled, shape complex, but rnm. Ref.: Haecker (1908), Reshetnjak (1966).
broadly hatchet-shaped in lateral view. Aboral • Protocystis varians Borgert (Fig. 6.98). Shell
surface flattened. Peristome process long, extend- compressed lenticular, elliptical, ovate, or nearly
ing into 2 curved, sub-parallel teeth. Length circular in side view, or nearly triangular with con-
(including process): 0.27-0.32 rnm. Ref.: Haecker vex sides. Mouth with a straight or slightly curved
(1908). process obliquely tilted over the opening, third to
Protocystis murrayi (Haeckel) (Fig. 6.92). Shell half the shell diameter, terrninating in three teeth,
spherical to subspherical. Peristome trapezoidal, one simple dorsal and two parallel and tilted
curved, collar-like lamella, width three times obliquely over the mouth. Middle tooth sometimes
length and equal to diameter of shell, its margin larger and stouter than the two paired ones, some-
with 6 triangular teeth, 2 convergent ventral, 2 times only in forrn of small protuberance, some-
extemal horizontal lateral, 2 ascending dorsal. times lacking altogether, with all interrnediate
Diameter: 0.15 mm. Ref.: Haeckel (1887). transitions. Length: 0.155-0.175 mm. Ref.:
• Protocystis sloggetti (Murray) (Fig. 6.93). Shell Borgert (1904).
triangular, strongly compressed, with three round-
ed comers, or nearly semi-circular. Aboral margin Family Circoporidae
straight, horizontal, longer than convex dorsal and I Shell with panelled structure and polygonal plates
ventral margins. Peristome with 2 parallel, trian- .............................................................................. 2
gular, pointed teeth, obliquely ascending, lamellar, la Shell spherical, with dimpled structure, with vari-
with inflated ovate base, about as long as shell able number of spines: Haeckeliana
radius. Diameter: 0.2-0.4 mm. Ref.: Haeckel 2 Shell octahedral (or spherical), with 6 radial
(1887). spines: l7ircoporus
• Protocystis swirei (Murray) (Fig. 6.94). Shell 2a Shell tetradecahedral, with 9 radial spines: Orco-
spheroidal to ovate, with single erect spine at abo- spathis
ral poleo Oblique peristome, with process extend-
ing into three teeth, of which the middle one is Orcoporus oxyacanthus Borgert (Fig. 6.99).
longest. Length: 0.095-0.37 rnm. Ref.: Haecker Shell nearly spherical, with 6 (seldom 7) strong
(1908). radial spines equal to or somewhat longer than
• Protocystis thomsoni (Murray) (Fig. 6.95). Shell shell diameter, with whorl of 3 (sometimes 4)
lentellipsoidal to lenticular, circular to ovate in lat- short, thick, slightly curved by-spines at the mid-
eral view, rather strongly compressed. Peristome die or nearer the base. Bases of spines with ring of
semi-tubular, nearly straight, extending obliquely 4 (seldom 3 or 5) elongate pores and a whorl of
and ventrally over oral opening, extending distally usually 8 long, thin by-spines, slightly curved at
into 2 parallel lateral teeth, sometimes with third the distal ends of the radial spines. Shell wall with
medial tooth. Length: 0.26-0.4 rnm. Ref.: Haecker alveolar structure. Mouth large, triangular or more
(1908), Reshetnjak (1966). rounded, with 3 narrow spines, with 1 or 2 basal
• Protocystis triangularis Peters (Fig. 6.96). Shell by-spines. Diameter: 0.14-0.2 rnm. Ref.: Borgert
stout, thick-walled, nearly triangular in lateral (1909b).
view. Dorsal wall broadly rounded, convex; v.en- • Grcoporus sexfuscinus Haeckel (Fig. 6.100).
tral wall narrow, slightly convexo Aboral wall flat- Shell subregular, octahedral, with 8 triangular,
tened to slightly concave. Ventral comer may con cave, somewhat unequal faces, covered with
expand into rounded projection. Peristome process extremely delicate network of small square dim-
long and twisted, originating from one side of dor- pies, furrowed by radial crests arising from bases
sal mid-line, and extending into 2 asyrnmetrical, of spines. Six radial spines about as long as shell
TI
diameter, four-sided, prismatic, with rounded sions, though some individual s may be somewhat
edges, with a verticil (sometimes two) of 4 transitional to that species. Future research may
crossed, curved bristles in basal half, distal apex indicate that the two species should be combined.
inflated, with trident or fuscina. Bases of spines Diameter: 0.36-0.45 rnm. Ref.: Haecker (1908),
with 4 crossed elliptical pores. Mouth cruciform, Reshetnjak (1966).
with 4 small teeth. Diameter: 0.20-0.25 rnm. Ref.:
Haeckel (1887). Family Coelodendridae
• Circospathis sexfurca (Haeckel) (Fig. 6.101). 1 Without frenulum and rhinocanna, no outer lattice
Spherical shell composed of irregular polygonal mantle 2
plates, with 6 radial spines, shorter than shell la With frenulum and rhinocanna, branches of tubes
diameter, covered with thin curved bristles. Radial anastomose to form outer bivalved lattice mantle.
spines with ring of 12-16 triangular pores, divided With simple galea .4
distally into 2 or 3 equal branches. Mouth with 3- 2 With nasal tubes 3
4 convergent teeth. Diameter: 0.55-0.75 rnm. Ref.: 2a Without nasal tubes. All tubes dichotomously
Haeckel (1887). branched: Coelechinus
• Haeckeliana darwiniana Haeckel (Fig. 6.102). 3 Tubes forked or dichotomously branched: Coelo-
Shell with 26-32 coronets, most with 5 (rarely 4 or dendrum
6) pores. Coronets smaller than their intervals and 3a Tubes developed as styles: Coelodiceras
than length ofby-spines. Dimples subcircular, half 4 With 6 styles, 1 odd, 2 paired on each valve: Coelo-
pore width. Main-spines, in center of coronets, graphis
longer than radius. Diameter: 0.3-0.42 rnm. Ref.: 4a With 10 styles, 1 odd, 4 paired on each valve:
Haeckel (1887). Coelodecas
• Haeckeliana irregularis Haecker (= H. labrador-
iana Borgert) (Fig. 6.103). Shell small, spherical, • Coelechinus wapiticornis Haecker (Fig. 6.105).
with thick wall. Pores circular to subpolygonal, Spherical skeleton. Galea with triangular oral and
uniform in size and shape, in syrnmetrical groups aboral surfaces. Aboral surface bridged to inner
(coronets) of 4-7 around bases of main spines, 1-2 valve. Rhinocanna near oral margin, bound to
times bar width, bars narrow to wide. Mouth cir- galea by two frenula. Two prirnary lateral tubes
1: cular to subcircular or elliptical, smooth except for and aboral tube each branched 5 to 7 times.
normally distributed by-spines. Main-spines fairly Primary tubes with 3 branches at first branching,
thick, straight, short, about as long as shell radius. other branches dichotomous, distal branches typi-
By-spines at intersections of surface ridges, length cally with very small, irregular teeth. Diameter:
1-3 times pore diameter, uniformo Surface orna- 2.2-2.8 rnm. Ref.: Haecker (1908).
mentation of low flat areas, very irregular and • Coelodecas ambulacrum Haecker (Fig. 6.107).
angular, separated by network of low, rounded Valves typically with ten styles, extending beyond
ridges. Diameter: 0.35-0.55 rnm. Ref.: Haecker externallattice, distal ends tapering and branching
(1908), Reshetnjak (1966). dichotomously 5 times producing 32 strongly
• Haeckeliana porcellana Murray (Fig. 6.104). divergent terminal branches in a low, flat tuft.
Shell small, spherical, with thick wall. Pores circu- Terminal branches slightly curved, with a few
1: lar to subpolygonal, very uniform in size, in sym- recurved hooks, with spathilla with 5-6 small teeth
I
metrical groups (coronets) around bases of by- at distal end of each branch. Length: 1.5-1.8 rnm.
I spines, 1-3 times bar width, bars narrow to wide. Ref.: Haecker (1908).
Mouth circular to subcircular or oval, of medium • Coelodecas pygmaea Haecker (Fig. 6.106).
size, smooth except for normally distributed by- Externallattice ovate in lateral view. Valves typi-
spines. Main-spines thick, straight, short, 1/3 to cally with ten styles, extending beyond external
3/3 shell diameter. By-spines 1-3 times pore diam- lattice, distal ends tapering and branching dichoto-
eter, uniformo Surface ornamentation of shallow mously 3 (rarely 4) times, producing 8 or 16 termi-
dish-shaped depressions, circular to subcircular, nal branches in a conical tuft. Terminal branches
separated by low rounded or sharp ridges. Differs long, slender, gene rally regularly undulant, with
from H. irregularis by less angular surface depres- recurved hooks. Spathilla with 3-5 stout, divergent
teeth at distal end of each terminal branch. Length: with frenulum. Nasal tubes branch at obtuse
1.3-1.5 rnm. Ref.: Haecker (1908). angles, with anchor threads. Anchor threads
• Coelodendrum flabellatum Haeckel (Fig. 6.108). bidentate, with terminal ends arranged like picks
Terminal branches fan-shaped, irregularly rami- with few short, opposing, sub terminal teeth.
fied, each of last two fork-branches divided into 4 Terminal branches of all tubes with strong, irregu-
or 5 diverging straight branches of different lar thorns. Length of nasal tubes: 2.0-2.2 rnm.
lengths. Last 8 or 10 ramules usually in meridion- Ref.: Haecker (1908).
al plane, their terminal knobs cap-shaped, with a • Coelographis antarctica Haecker (Fig. 6.115).
corona of recurved teeth. Diameter: 2.0-2.5 rnm. External lattice nearly pentagonal, shaped like
Ref.: Haeckel (1887). arrowhead in lateral view. Nasal style with 5 pairs
• Coelodendrum furcatissimum Haeckel (Fig. of larger branches and a few smaller, distal,
6.112). Spherical shell with hemispherical valves. unpaired branches, all in plane parallel to median
Galea arched and aborally concave. Nasal opening suture between shell halves. All paired and
with cIeft. Four primary radial tubes bifurcate 6 to unpaired branches further branched, distal ends
9 times at acute angles, branches smooth and extending to externallattice. Nasal style and 2 sec-
straight or slightly curved, very small knobs at end ondary lateral styles extend slightly beyond exter-
of each terminal branch, each with 3-4 short nal lattice. Protruding nasal style with terminal
recurved teeth. Diameter: 1.8-2.5 mm. Ref.: crown of 3 or 4 twice-dichotomous short, smooth,
Haecker (1908). terminal branches. Two protruding secondary lat-
• Coelodendrum lappaceum Haeckel (Fig. 6.109). eral styles with terminal crowns, generally with
Terminal branches regularly forked, with 2 equal, only two twice-dichotomous, short, smooth, ter-
straight or slightly curved fork-branches, diverg- minal branches. Length: 2.0-3.6 rnm. Ref.:
ing at acute angles and covered with small Haecker (1908), Reshetnjak (1966).
recurved hooks; their terminal knobs large, coni- • Coelographis regina Haeckel (Fig. 6.114). Shell
cal, with prominent apex and basal corona of 6 to mantle twice as long as wide, its frontal perimeter
8 recurved teeth. Diameter: 1.5-1.8 rnm. Ref.: an isosceles triangle, with triangular excision at
Haeckel (1887). base, its sagittal perimeter slenderly ovate. Nasal
• Coelodendrum ramosissimum Haeckel (Fig. style twice as long as paired tergal styles, former
6.110). Spherical shell with hemispherical valves. with 10-12 (10-30, Reshetnjak, 1966), latter with
Galea arched, with large nasal opening. Four pri- 5-6 alternate-cruciate pairs of lateral branches.
mary tubes bifurcate at right angles 4 to 6 times. Terminal coronets (on free distal ends of styles)
Branches smooth and straight, or slightly curved. palmate, with 10-12 (4-12, Reshetnjak, 1966)
Ends of terminal branches sometimes with very spinulate, irregular, finger-shaped branches.
small inflated disc with 4-7 (usually 4) short Length: 5.5 rnm. Ref.: Haeckel (1887), Reshetnjak
recurved teeth. Diameter: 0.55-1.8 rnm. Ref.: (1966).
Haecker (1908), Reshetnjak (1966).
• Coelodendrum spinosissimum Haeckel (Fig. Family Concharidae
6.111). Terminal branches regularly forked, with 1 Valves without sagittal keel, nearly hemispherical
two equal, smooth, straight, fork-branches, diverg- or slightly compressed 2
ing at right angles, their terminal knobs echinoid- la Valves compressed, watch-glass-shaped. Aboral
al, subspherical or club-shaped, with numerous hinge without horns, with a pair of low lips:
short radial thorns. Diameter: 2.0-2.2 rnm. Ref.: Conchophacus
Haeckel (1887). lb Valves laterally compressed, boat-shaped. Aboral
• Coelodiceras spinosum Haecker (Fig. 6.113). hinge without horns: Conchopsis
Shell late rally compressed, butterfly-shaped. 2 Aboral hinge without horns: Conche/lium
Galea large, convex, inflated, broadly triangular in 2a Aboral hinge with two horns, no apical horn:
oral and aboral views. Valves small, covered by Conchidium
aboral surface of galea. Rhinocanna long, thin,
tubular, very near oral margin of valve. Irregular • Conche/lium capsula Borgert (Fig. 6.116). Shell
margin around nasal opening of rhinocanna, latter usually nearly spherical, valves almost the same
size, about hemispherical or tapering slightly abo- 4 to 6 times as long as bars, in regular transverse
rally, rims sometimes slightly indented. Rims of rows. Length: 0.22 mm. Ref.: Haeckel (1887).
both valves with strong, conical teeth of about the • Conchopsis compressa Haeckel (Fig. 6.119).
same length, about 10-15 on each side of each Shelllenticular, strongly compressed, ratio of lon-
va1ve. Aboral rim widened into a lip- or nose- gitudinal to sagittal to lateral diameters = 10:9:3,
shaped structure, rounded or sometimes pointed. sagittal perimeter elliptical, cinctural and frontal
Pores circular, regularly arranged, about 2-3 times perimeters spindle-shaped. Borders of two boat-
bar width. Bars smooth or weakly framed. Length: shaped valves smooth in oral and aboral thirds,
0.022-0.024 mm. Ref.: Borgert (1907). dentate in middle fourth, about 40-44 very slender
• Conchidium caudatum (Haeckel) (Fig. 6.117). equal teeth on one lateral edge of each valve. In
Shelllenticular, strongly compressed, ratio of lon- half frontal perimeter 64-68 pores, in half sagittal
gitudinal to sagittal to lateral diameters = 6:5:4. perimeter 70-80 pores, in half equator 44-48
Free margins of two boat-shaped valves, dentate pores. Ventral and dorsal pores linear, 3 to 4 times
nearly in whole perimeter, in one lateral border of as long as circular, lateral pores. Length: 0.6-0.8
each valve 11 or 12 very large conical teeth, larg- mm. Ref.: Haeckel (1887).
est near half as high as valve. On aboral hinge 2 • Conchopsis orbicularis Haeckel (Fig. 6.121). Shell
divergent, very large horns, four-sided pyramidal, subcircular in outline, lenticular, strongly com-
acute, straight, with a few basal pores. Dorsal horn pressed on both sides. Sagittal perimeter nearly cir-
(of smaller val ve) somewhat shorter than ventral. cular, frontal and cinctural perimeters spindle-
Apical distance of both horns somewhat greater shaped. Borders of two boat-shaped valves smooth
than their length, about half length of shell. In half in 0.4 of oral part, and in 0.1 of aboral part, remain-
frontal perimeter (along one border of each valve) ing 0.5 middle part strongly dentate, with about 25
32-34 pores, in half sagittal perimeter 34-38, in straight, slender teeth on each side of one valve,
half equator 30-33. Pores smaller near girdle-fis- increasing in size from aboral towards oral poleo In
sure, irregularly quadrangular, in longitudinal the half lateral perimeter of the shell (along the
rows, separated by meridional crests, converging right and left borders of each valve) 60-65 pores, in
towards both poles of main axis. Length (without the half sagittal perimeter (along the keel of each
horns): 0.24-0.26 mm. Ref.: Haeckel (1887). valve) 80-85 pores. in half equator 60-65 pores.
• Conchidium terebratula Haeckel (Fig. 6.118). Diameter: 0.53-0.75 mm. Ref.: Haeckel (1887).
Shell subspherical, smooth, both valves nearly
equal in size and form, hemispherical. Three dimen- Family Lirellidae
sional axes almost equal. Margins of valves dentate Chamber subdivided by internal diaphragm: Borger-
in nearly whole periphery, on each side of one valve tella
11 or 12 strong conical teeth, of equal size. Aboral Chamber not subdivided by internal diaphragm:
hinge with 2 short, stout four-sided pyramidal horns Lirella
of equal length. Mouth with two equal short lips.
Pores subregular, circular, 3-4 times bar width, • Borgertella caudata (Wallich) (Fig. 6.122). Ovate
twice as larger in dorsal valve as ventral. Length: main chamber, with hollow spine at aboral pole,
0.24-0.28 mm. Ref.: Haeckel (1887). separated by an internal diaphragm and partially
• Conchophacus diatomeum (Haeckel) (Fig. by an external constriction from secondary cham-
6.120). Shell nearly spherical, slightly lenticular, ber, nearly as large as main chamber. Secondary
somewhat compressed dorso-ventrally, sagittal chamber connected to main chamber by eccentric
diameter therefore somewhat shorter than other tube through diaphragm. Secondary chamber
two. Borders of two hemispherical val ves nearly broadly curved to open at about right angle to
circular, quite smooth, about as wide as largest main axis, lower lip of peristome may be extend-
pores. In frontal perimeter of shell 44-50 pores, in ed as spine, or connect through a broadly arched
half sagittal perimeter 20-24, in half equator 30- filamentous bar (probably normal, but seldom pre-
33. Pores different in form and size, marginal served) to aboral spine. Surface with longitudinal
pores nearly small, circular; dorsal and ventral striae. Length: 0.037-0.048 mm. Ref.: Borgert
pores oblongish-hexagonal, twice as long as wide, (1904), Dumitrica (1973).
• Lirella bullata (Stadum and Ling) (Fig. 6.123). horizontal), remainmg ones shorter thoms
Shell small, ovoidal or ellipsoidal, with about 25- between the larger ones. Length: 0.110-0.116 mm.
35 longitudinal striae. Smooth open bubble pro- Ref.: Borgert (1906).
jects from mouth, with short apical spine at aboral • Euphysetta pusilla eleve (Fig. 6.128). Shell egg-
poleo Length: 0.48-0.58 mm. Ref.: Stadum and shaped, without apical hom, structure double.
Ling (1969), Takahashi and Honjo (1981). Longitudinallines 11 in 0.01 mm, very small are-
• Lirella melo (eleve) (Fig. 6.124) Ellipsoidal shell, olae, quincuncially arranged, 19 in 0.01 mm.
with oral aperture at end of short, stout, collar-like Small cylindro-conical legs, odd leg half as long
peristome. Peristome perpendicular to main axis. as shell, not branched. Length: 0.06 mm. Ref.:
Usually with short aboral spine. Surface with 25- eleve (1900), Borgert (1906).
40 longitudinal striae. Length: 0.07-0.14 mm. • Medusetta ansata Borgert (Fig. 6.129). Shell
Ref.: Haecker (1908). ovoid, weakly laterally compressed, surface with a
number (usually 14-16) ofmore or less prominent,
Family Medusettidae equally spaced, longitudinal, meridional ribs, usu-
With four equal feet: Medusetta ally restricted to the widest part of the shell, lack-
With 1 large and 3 small feet: Euphysetta ing on the aboral dome and the oral area. Four
With 8-10 feet: Planktonetta strong, divergent, distally pointed legs, connected
With 10-13 feet: Nationaletta in pairs by narrow threads into two lateral arches.
Unbound pair of legs on one side closer together
• Euphysetta dubia Schróder (Fig. 6.126). Shell and longer than the opposite pair. Outer side of
ellipsoidal, with wall of thick outer and thin inner legs branched, with longest, thickest branches usu-
layer, outer layer with faint longitudinal striae and ally toward the mouth. Base of each leg with thin
less distinct horizontal striae. Oral opening wide, bristle directed obliquely outwards, and often 3-5
with 3 small solid legs and one chambered leg, narrow, short spines on rim of the mouth within
smaller legs with an accessory thom, large leg the arches. Takahashi and Honjo (1981) distin-
about as long as shell, with distal thom-like acces- guished an Atlantic form with ovoid shell and feet
sory branch. Stout spine, often short, at aboral extending obliquely and bent abruptly at the
pole, usually erect, sometimes tilted ventrally. branching joint. Length: 0.065-0.075 mm. Ref.:
Length: 0.06-0.08 mm. Ref.: Schróder (1913). Borgert (1906), Takahashi and Honjo (1981).
• Euphysetta elegans Borgert (Fig. 6.125). Shell • Medusetta inflata Borgert (Fig. 6.130). Shell with
spheroidal to ovoidal, with stout spine at aboral dissimilar curving surfaces, flattened on one side,
pole, usually erect, sometimes sloping ventrally. in side view nearly circular or subcircular, smooth.
Surface with numerous horizontal and vertical On aboral pole 4 chambered, weakly divergent,
ridges, dividing surface into rather regular ovallegs, shorter than (often scarcely half as long
squares. Oral rim with 3 small legs and one large as) shell diameter, distally straight or bent slightly
chambered lego Large leg with 2 short, proximal, inward, with longer or shorter lateral branch about
lateral thoms, and distal auxiliary branch. Small in the middle or toward distal end. Terminal and
legs with 2 short distal thoms and 2 long, slender, lateral branches longer on one side off the shell.
proximal spines. Length: 0.06-0.95 mm. Ref.: Sometimes 1 or 2 short thoms on rim of mouth.
Haecker (1908), Reshetnjak (1966). Length: 0.065-0.085 mm. Ref.: Borgert (1906).
• Euphysetta lucani Borgert (Fig. 6.127). Shell • Medusetta rara (Borgert) (Fig. 6.131). Shell asym-
nearly circular or ovate, with dissimilar surface metrically flattened, nearly circular or subcircular
curvature, smooth. With one stronger, chambered, in side view or somewhat wider than long, smooth,
bent leg, nearly as long as shell diameter, distally without spines on the aboral poleo Oral main leg
tapered, with outwardly directed point, with 2 thin, simply pointed, chambered, somewhat longer than
pointed lateral spines on the convex side, the one shell length, more or les s strongly bent, with long
nearest the shell significantly longer. Mouth also lateral branch on convex side near mouth. Three
with a number of narrower spines, usually 3 larg- short oral spines, pointed, divergent, tooth-shaped,
er and often longer, wavy, outwardly bent (nearly developed from mouth rim. Sometirnes the oral
s
spines also bear a narrow lateral branch projecting assemblage of various diatoms, silicoflagellates,
obliquely or nearly horizontally. Length: 0.085- polycystine radiolarians, or sponge spicules.
0.096 rnm. Ref.: Borgert (1906). Diameter: 0.9-0.3 rnm. Ref.: Borgert (1909a).
• Nationaletta fragilis (Borgert) (Fig. 6.132). Shell • Phaeodina tripylea Haeckel (Fig. 6.137). Central
dome-shaped, its outer edge extended into 10-13 capsule spherical to somewhat flattened in the
slender areolate legs, with numerous perpendicular direction of the main axis. Astropyle domed,
spines which usually extend into tufts of anchor scarcely a third as wide as the central capsule
barbs, with straight or curved arms, branching dis- diameter. Parapylae of characteristic form and
tally into terminal spines. Oral opening surrounded location. (Figured specimen in process of fission
by ring of pores. Diameter: 0.4-0.6 rnm. Ref.: according to Haeckel, 1887). Diameter: 0.15-0.25
Haecker (1908). rnm. Ref.: Borgert (1909a).
• Planktonetta atlantica (Borgert) (Fig. 6.133)
Shell delicate, bladder-like, generally ovoid, or Family Pharyogellidae
pear-shaped to spherical. Surface with small, fine • Pharyngella gastrula Haeckel (Fig. 6.138). Shell
thoms, which sometimes have small basal discs. ovoid. Peristome with two parallel, vertical, straight
Central capsule lines inner shell. Spherical float to slightly curved, pointed teeth, sometimes
about same size as shell attached to one side. Wide obliquely tilted over mouth, half as long as shell
oval opening with stout collar-like structure. Four radius. Pharynx of same length, cylindrical or fun-
pairs of distally ramified (2-4 irregularly forked nel-shaped, outer aperture scarcely wider than
terminal branches) oral legs at periphery, sur- inner. Length: 0.30-0.36 rnm. Ref.: Haeckel (1887).
rounding oral opening, 1-3 times as long as shell,
with short, perpendicular lateral spines along Family Porospathididae
length, extending into tufts of anchor barbs. Shell surface panelled, tabulate, or covered by trizonal
Fibrous diaphragm of central capsule between meshwork, with radial spines, with peristomal tube:
mouth and shell body. Skeletal meshwork across Porospathis
oral opening, connecting float to shell and sup-
porting phaeodium. Length: 1.2-1.7 rnm. Ref.: • Porospathis holostoma Cleve (Fig. 6.139).
Haecker (1908), Reshetnjak (1966). Spheroidal shell with papillae tapering into thoms,
connected by regular triangular ridges, with oral
Family Phaeodioidae opening at flared distal end oflong cylindrical per-
• Phaeodina ambigua (Borgert) (Fig. 6.134). istome. Radial spines (13-30) over all of shell sur-
Rounded body, more or less significantly flat- face, but more abundant in oral than in aboral
tened, with two central capsules. lo the extracap- hemisphere, length 3-4 times shell diameter.
sulum outside the phaeodium is a covering, vari- Diameter: 0.05-0.16 mm. Ref.: Hülsemann
able in density, of foreign siliceous objects. (1963), Reshetnjak (1966).
Diameter: 0.19-0.3 rnm. Ref.: Borgert (1909a).
• Phaeodina antarctica (Schróder) (Fig. 6.135). Family Sagosphaeridae
Cell presumed to be originally spherical, but is 1 With simple latticed mesh (tops of pyramids not
later somewhat deformed, depending on materials joined by tangential bars) 2
incorporated into the skeleton. Central capsule la With double latticed mesh, with tops ofpyramids
ellipsoidal to spheroidal. Skeleton of encrusted joined by tangential bars: Sagenoarium
detrital siliceous material. Individuals in 2 Pyramids with axial rod (Fig. lE): Sagenoscena
Schróder' s illustrations shown with a variable 2a Pyramids without axial rod (Fig. ID): Sagoscena
mixture of skeletons from diatoms, silicoflagel-
lates, and polycystine radiolarians. Diameter: • Sagenoarium antarcticum Haecker (Fig. 6.140).
0.06-0.09 rnm. Ref.: Schróder (1913). Shell ovate, with pyramidal elevations of several
• Phaeodina loricata (Borgert) (Fig. 6.136). Surface long delicate spines, terminating in whorls of 3 or
of the cell covered with foreign siliceous objects, 4 small, thin branches with reniform terminal
ranging from the tests primarily of a single species pads. Diameter: one whole specimen reported as
of silicoflagellate to a (seemingly random) mixed 2.5 by 3.5 rnm. Ref.: Haecker (1908).
• Sagenoarium chuni Borgert (Fig. 6.141). Lattice Family Tuscaroridae

shell ovoidal, pear-, heart-, or spindle-shaped. Aboral spines inserted in the equatorial or aboral
Pyramids with 2-5 (usually 3) apical spines, with areas, peristome annular: Tuscarora
.2-3 distal whorls of 3-4 lateral branches. Branches Aboral spines inserted in the apical or aboral areas:
of terminal whorl with forked or tripartite spathil- Tuscarantha
lae; branches of inner whorl with simple spathil- Aboral spines arising from oral hemisphere of shell,
lae. Length: 3.0-6.8 mm. Ref.: Haecker (1908), in close vicinity of oral spines: Tuscaretta
Reshetnjak (1966). Aboral spines present at the aboral pole only:
• Sagenoarium dicranon Haecker (Fig. 6.142). Tuscarilla
Lattice shell spherical, ovoid, or balloon-shaped.
In balloon-shaped individuals, pyramids and their • Tuscarantha braueri Haecker (Fig. 6.147). Shell
apical spines longer on the blunt than the pointed triangular-pyramidal with rounded corners, or sel-
end. Apical spines thick, unbranched, with a weak- dom quadrangular-pyramidal, or triangular shield-
ly thorny, inflated termination. Length: 4.0-5.5 shaped. Peristome in shape of a short, wide tube,
mm. Ref.: Schróder (1913), Reshetnjak (1966). somewhat widened at the opening, with typically
• Sagenoscena irmingeriana Borgert (Fig. 6.143). developed basket-shaped peristomal termination.
Spheroidal to pear-shaped lattice shell. Pyramids Three long, widely divergent oral spines. Three
usually with 5-6 sides and stout axial rod, with one (seldom 2 or 4) very long, massive, divergent abo-
or a few (2-5, Reshetnjak, 1966) radial spines at ral spines. Length (including peristome): 1.4-2.3
apex, with crown of 10-20 slightly curved terminal mm. Ref.: Schróder (1913), Reshetnjak (1966).
branches, sometimes slightly fused at bases. Axial • Tuscarantha hydra Haecker (Fig. 6.148). Shell
rods sometimes with short lateral branches at apex spindle-shaped, thick-walled, with apical spine.
of pyramids, sometimes forming secondary spher- Peristome in shape of short, wide tube, widened at
ical lattice, rarely with poorly developed crowns the opening, with 5 well-developed oral spines,
of few short terminal branches. Length: to 3.5 or with several large basal pores, bent horizontally.
4.5 mm. Ref.: Haecker (1908), Reshetnjak (1966). Length: 2.8 mm. Ref.: Schróder (1913).
• Sagenoscena tetracantha Haecker (Fig. 6.144). • Tuscarantha luciae Haecker (Fig. 6.149) Shell bal-
Shell pear-shaped, with 4- to 6-sided pyramidal loon-shaped, with strongly arched aboral and grad-
elevations. Stout axial rods, extend radially as ually tapered oral halves. Aboral surface drawn out
spines with 3-6 (generally 4) short, stout, some- in a gradual curve into the very strong basal cone of
what curved, diverging terminal branches. the apical spine. Peristome in shape of short, wide
Pyramidal bars may also extend as lateral branch- tube, widened at the opening, with well-developed,
es at the apex. Diameter: one whole specimen basket-shaped peristomal termination. Three very
reported as 4.0 by 5.5 mm. Ref.rHaecker (1908). long (to twice shelllength) oral spines. Single api-
• Sagoscena castra Haeckel (Fig. 6.145). Shell cal spine, twice shelllength. Length: 2.8-3.0 mm.
spheroidal with regular pyramidal elevations of Ref.: Schróder (1913), Reshetnjak (1966).
solid rods that fuse in a thick node, and continue • Tuscaretta aeronauta Haecker (Fig. 6.150). Shell
as short, simple terminal spines, each with a spin- ovate, thin-walled. Peristome short, tubular, some-
ulate terminal knob, rarely as a single radial termi- what constricted in middle, with 3-4 oral spines on
nal spine. Diameter: 1.7-2.3 mm. Ref.: Haeckel terminal rim. Oral spines slightly divergent, weak-
(1887). ly dentate, lacking basal pores. 4-6 aboral spines
• Sagoscena praetorium Haeckel (Fig. 6.146). originate in oral hemisphere of shell. Height: 1.2-
Lattice shell spheroidal to ellipsoidal, with 3- to 6- 1.6 mm. Ref.: Haecker (1908).
sided pyramids, with 3-5 terminal spines. Terminal • Tuscaretta globosa Borgert (Fig. 6.151, 6.153).
spines with distal whorl of terminal branches, usu- Spheroidal or ellipsoidal shell, with short tubular
ally with 4 branches, each with terminal spathilla. peristome on oral pole, with 3 slender, equally
One or a few single or paired lateral branches spaced, slightly divergent oral spines on peristo-
irregularly along remainder of terminal spines. mal rim. One or two large triangular pores in
Length: 2.6-3.1 mm. Ref.: Haeckel (1887). broad base of each oral spine. Four to six equally
spaced, long, aboral spines, as ring around peri-
stome, curved toward aboral hemisphere, with 3-4 divergent aboral spines. Length: 1.2-1.6 mID. Ref.:
elliptical pores surrounding base. Oral and aboral Schróder (1913), Reshetnjak (1966).
spines with numerous curved thoms throughout • Tuscarilla nationalis Borgert (Fig. 6.i56). Shell
length. Length: 1.15-1.46 mID. Subspecies. Two triangular, pyramidal, extended into a tube at the
subspecies described by Haecker (1908) have oral end, widened into a funnel-shape, its rim with
been reported from the South Atlantic: T g. atlan- 6 (seldom 4-5) long, straight or weakly curved,
tica (larger, thick-walled, spheroidal, Fig. 6.151), evenly spaced oral spines, 3 times as long a shell
and T g. chuni (smaller, cap-shaped, with com- length or longer, with 1-3 rounded oblong basal
pressed oral-aboral axis, Fig. 6.153a, 6.153b). pores. Aboral comers with 3 straight, short,
Latter possibly Antarctic endemic. Ref.: Haecker strongly divergent aboral spines, as long or longer
(1908), Reshetnjak (1966). than shelllength, with ring of 3-5 rounded oblong
• Tuscaretta tubu/osa (Murray) (Fig. 6.152). Shell pores at their conically widened bases. Oral and
pear-, egg- or cupola-shaped. Peristome laterally aboral spines simply pointed, with numerous dis-
compressed, with slit-shaped opening, with two tally curved thoms. Length: 1.1-1.5 mID. Ref.:
slightly divergent oral spines, running parallel, Haecker (1908), Reshetnjak (1966).
converging or crossing. Two to five (usually 3 or • Tuscarilla similis Schroder (Fig. 6.157). Shell tri-
4) aboral spines bending sharply toward the aboral angular, shield-shaped. Peristome long, relatively
end. All spines with 4-5 basal pores, and curved narrow, extended into 3 divergent oral spines, with
thoms along the length. Length: 1.2-1.6 mID. Ref.: 2 rounded oblong basal pores. Comers of aboral
Schróder (1913), Reshetnjak (1966). side with 2 aboral spines, fairly steeply inclined
• Tuscarilla ampulla Haecker (Fig. 6.154). Sphe- upwards, with 2-3 basal pores. Length (including
roidal to ovoidal shell, with tubular peristome, peristome): 1.75 mID. Ref.: Schróder (1913).
with 3 or 4 aboral spines arising from near aboral • Tuscarora bisternaria Murray (Fig. 6.158). Shell
pole, with 3 or 4 straight or slight1y curved oral subspherical, pear-shaped, seldom with flattened
spines originating from lip of peristome. Pores at aboral surface. Peristome thick, ring-shaped, with 3
base of oral and aboral spines. Length (including straight, divergent oral spines, with 8-10 large basal
peristome): 0.7-1.5 mID. Ref.: Haecker (1908), pores, with stout, partly branched thoms at their
Reshetnjak (1966). bases, partly fused with the peristome. Three (sel-
l' • Tuscarilla campanella Haecker (Fig. 6.155). dom four) straight, weakly spiny, aboral spines,
Shell bell-shaped with nearly flat aboral surface. somewhat bent only at the base, usually in an equa-
Peristome very short and wide, cup-shaped, with torial position, altemating with oral spines. Length:
4-6 almost horizontal oral spines, with 3-4 basal 1.5-2.0 mID. Ref.: Schróder (1913), Reshetnjak
pores. Rim of aboral surface with 5-7 strongly (1966).
ASTRACANTH I DAE ATLANTICELLI DAE

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1: skeleton of
At/anticella bicornis At/anticella
¡i Astracantha umbellifera craspedota
i.;.!,.
~:. 4
Astrecenthe
heteracantha
6.1 fi
6.5
At/anticella p/anctonica
Cornucella maya
Ha/ocella gemma Ha/ocella magna
Lobocella proteus
AULACANTHIDAE
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6.10
6.11 A. s. barhybla A. s. scolymanrha
ti,!
Aulacantha Aulacantha scolymantha 6.12b ~.;; 6.12c
cannulata
~~1t.jIj}~
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6.15 ! ..
ti'
Auloceros arborescens birameus Auloceros arborescens
irregularis
Auloceros arborescens 6.18a

subelegans
!
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t 1
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Diatom frustule I! :
'1 i ramosus
(Rhizosolenia sp.) ¡!t ;
varo longispina
Auloceros within a radial 6.19
n ;
arborescens spine
6.20b
trigeminus
Aulokleptes flosculus
·r~---------------------------------
AULACANTHIDAE
6.23d
6.22
6.23b
Aulographis furcula Aulographis mohri 6.23c
Aulographis bicurvata
6.27
, Au I ographis
¡i'
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6.26 i
il
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I
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6.25bll ..: Aulographis

tetrencistrs Aulographis
Aulographis pandora Aulographis stellata triangulum
6.29.\
't Aulographonium
!; antarcticum
6.30 Aulogrephonium
Aulographonium bicorne
anthoides
~.
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;.~;
Aulographonium
6.331: Aulo¡¡raphonium
j mediterraneum Aulographonium
candelabrum pulvinatum
AULACANTHIDAE
,
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6.35 ;:
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6.37a
6.38 f I
pmus
6.36
I 6.37b ¡
11
A ulospathis
\'
11 '
variabilis
A ulospathis Aulospathis variabilis bifurca diodon
variabilis 6.39
aulodendroides
Aulospathis
variabilis
II
triodon
AULOSPHAERIDAE
, ! 6.40b
: v I ". t
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6.40a {'
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Aulastrum mirabile 6.41 a
6.41 b "----L__ --'
Aulastrum A ulatractus
monoceros fusiformis
6.43a
Aulastrum spinosum
T
i Radiolaria Phaeodaria 249
. ~-\)
AU LOSPHAERI DAE -~/
I
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6.44
6.46, 6.48
i
r
A. v. typíca oO, __ , ••_._~
A. v. hamata
Auloscena Auloscena verticillus Aulosphaera
robustissima fi/igera
6.49a
t
6.51 b
\;
" 6.51 a
Aulosphaera labradoriensis
:_s l' ., lt::=-!-"<-'
6.50a ::. 1,·-:;;0'

Aulosphaera
elegantissima Aulosphaera
robusta
6.54
!~
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Aulosphaera ,
--l'
trispathis "i
... '~ 6.55b
O¡l
6.55a ij
~ Antarctic
{i
form
:i 1
A. b. .septentrionelts i
1,[
I . .
:'1
'1 Equatorial ..~
Aulosphaera 6.53c, ,
form '
bisternaria 6.53a / .~.
Aulosphaera triodon Aulosphaera verticillata
250 Radiolaria Phaeodaria /,~
, ~\
1 I
-~./
CANNOSPHAERIDAE /
Cannosphaera antarctica
detailof
central shell radial
spine
CASTANELLIDAE <
~!
6.57' / ji !
Castanarium antarcticum
Castanarium hookeri
Castanarium favosum
I
1
1
I
6.60b
Castanea amphora
1,
Castanea globosa Castanea henseni
,1,
11
I
, !
/'-
! -J.-)
. T /
CASTANELLIDAE /
Castanella borgerti
Castanella maxima
Castanella thomsoni
6.66b
Castanissa nationalis
I
6.67 ¡ Castanissa valdiviae
Castanissa proteus
Castanissa schmidti Castanopsis fragilis

\
CASTANELLIDAE
Castanidium haeckeri
Castanidium longispinum
Castanidium moseleyi
Castanidium variabile
T
CHALLENGERI DAE
6.77a
Challengeranium diodon
6.78a 6.78b 6.79a 6.79b

Challengeria naresi
Challengeria tritonis
Challengeron radians 6.81 b

6.82 Challengeron channeri
Radiolaria Phaeodaria -------
254
/ ~))
r
CHALLENGERI DAE
6.83a 6.83b Chal/engeron willemoesii
6.85a 6.85b
Protocystis Protocystis balfouri 6.86 ~ 6.87
acornis Protocystis Protocystis
bicornuta bicuspis
6.90a
Protocystis Protocystis micropelecus

heterocentrota Protocystis macleari
CHALLENGERI DAE /! f\ \
I
,/
./
6.92b 6.93
Protocystis sloggettii
Protocystis murrayi
6.94a
Protocystis swirei
Protocystis triangularis Protocystis tridens Protocystis varians

CIRCOPORIDAE
Circoporus
6.100 sexfuscinus
6.99a 6.99b
Circoporus
oxya can thus
6.102b
Haeckeliana darwiniana
Circospathis sexfurca
!
/
Haeckeliana
porcellana
Haeckeliana
6.103 irregularis
--------
l
COELODEN DRI DAE r--:- ---,(i ~
Coelodecas pygmaea
Coelodecas
ambulacrum
\l.~'f
~:V
Coelodendrum flabellatum
II 6.109b
Coelodendrum
lappaceum Coelodendrum 6.111 ~.•
ramosissimum e
Coelodendrum
spinosissimum
one shell valve
6.113b
6.113a
~
Coelodiceres
spinosum Coelographis regina Coelographis antarctica
-- ...•...
/¡I r )
I IV¡ /
~/
CONCHARIDAE
6.116b
6.116a Conchellium capsula Conchidium terebratula
;.' .
1,'·
\
\,
-, 6.121
6.119 6.120~ -,
Conchidium '-- Conchopsis
caudatum Conchopsis compressa Conchophacus diatomeum orbicularis
LIRELLIDAE
6.123a
6.124a
6.124b
612~
Lirel/a Lirel/a
6.123b bul/ata melo
Borgertel/a caudata ~-"""""~--
..,.."
,!
MEDUSETTIDAE
Euphysetta e/egans
6.129b
Medusetta Medusetta ¡
ansata rara
Medusetta inflata
6.132
Nationaletta
fragilis .> float 6.133b
Planktonetta atlantica
í
. /\\(:))
PHAEODINIDAE \ '
,,--/ I
Phaeodina antarctica
Phaeodina loricata
PHARYNGELLIDAE
Phaeodina tripylea
POROSPATHIDAE
Porospathis holostome . 1
6.139c j
-
SAGOSPHAERI DAE ll)

/
Sagenoarium
chuni
Sagenoarium
antarcticum
Sagenoarium dicranon
~L/./
'1'
Sagenoscena
te tracantha
¡}
=»
Sagoscen!
Sagoscena
castra
TUSCARORIDAE
Tuscaretta Tuscaretta giobose

aeronauta etlentlce
Tuscarantha
braueri
Tuscarantha
hydra
I
J 1
I
1
I
Tuscarantha
/uciae
6.152c
Tuscaretta tubu/osa
Tuscaretta g/obosa chuni

Tuscarilla campanella
Tuscarilla
ampulla
Tuscarilla
Tuscarilla nationalis similis Tuscarora bisternaria
Acknowledgements radiolarians, including classifications of al!

groups, descriptions and illustrations of thou-
Thanks are due to colleagues and editorial offices for sands of species.
pennission to use their illustrations, and especially to Haecker V. 1908. Tiefsee-Radiolarien. Wiss. Erg.
Renate Bemstein, Marcia Gowing and Steve Haddock Deutsch. Tiefsee-Exp, Dampfer "Valdivia" 1898-
for providing several of the photographs inc1uded. 1899,14, T. 1:1-476; T. 2:477-706. A comprehen-
This work was partly fmanced by grants from the sive expedition report covering al! previous work,
University ofBuenos Aires (UBA EX-040), and from new species descriptions, and new distributional
the Consejo Nacional de Investigaciones Científicas y data.
Técnicas (CONICET PID-BID 0366). Kling S.A. 1976. Relation of radiolarian distribution
to subsurface hydrography in the North Pacifico
Deep-Sea Res., 23: 1043-1058. Relates subsurface
Suggested readings distributions of 'phaeodarians to water masses and
current systems.
Anderson O.R. 1983. Radiolaria. SpringerVerlag, New Meyer K. 1934. Die geographische Verbreitung der
York, pp. 1-355. A modern review of radiolarian tripyleen Radiolarien des Südatlantischen Ozeans.
biology, concerned mainly with polycystines, but Wiss. Erg. Deutsch. Atlant. Exp. "Meteor" 1925-
with considerable important information on phaeo- 1927, 12(2):122-198. Comprehensive horizontal
darians. and vertical distributional data for most phaeo-
Cachon-Enjumet M. 1961. Contribution a l'étude des darian families in a number of north-south and
Radiolaires Phaeodariés. Arch. Zoo1. Exp. Gén., east-west projiles in the South Atlantic.
100: 152-237. A detailed study of the biology Reshetnjak V.V. 1966. Glubokovodnie radilaoyarii
(including cytology) and systematics of some Phaeodaria severo-zapadnoi chasti Tichogo Okeana.
phaeodarian species. Fauna SSSR, Akad. Nauk SSSR, Zoo1. Inst, Nauka,
Gowing M.M., Bentham W.N. 1994. Feeding ecology Moskva: pp. 1-208. Although based on materials
of the phaeodarian radiolarians at the VERTEX from northwest Pacific, this comprehensive mono-
North Pacific time series site. J. Plankton Res., graph discusses taxonomic, biological, and dis-
16:707-720. This and previous studies by Gowing tributional data from al! previous litera ture.
and co-authors reveal, through detailed light and Schróder O. 1913. Die Tripyleen Radiolarien
electro n microscopy, the nature of the phaeodium (Phaeodarien) der Deutschen Südpolar Expedition
and general feeding ecology of a number of 'phae- 1901-1903. Deutsche Südpolar-Exp. 1901-1903,
odarian species. 14(Zoo1. 6):115-215. One ofthe latest expedition
Haeckel E. 1887. Report on Radiolaria collected by reports. Includes descriptions of southern ocean
H.M.S. Challenger during the years 1873-1876. species of al! major families, and summarizes pre-
Rep. Sci. Res. Voyage H.M.S. Challenger 1873- vious distributional data from al! oceans.
76, 18:1-1803. The basic classical work on
••• Fig. 6. Illustrations of phaeodarian species.

Figure sourees: eourtesy ofRenate Bemstein: 78b, 79b, 80a, 81b, 83b, 92b, 95e, d, 101b, 104b, 116e, 125e, 127a, 128b, 138e, 139b; from Borgert
(1904): 117, 122b, 138b; from Borgert (1906): 125a, 127b, 129, 131; from Borgert (1907): 116b; from Borgert (1909): 99a, 100, 103, 134, 136;
from Borgert (1910): 13ge; from Borgert (1911): 77e, 81a, 83e, 85a; from Borgert (1913): 7,9; from Dumitriea (1973): 122e; from Dworetzky and
Morley (1987), with kind permission from Elsevier Seienee: 66b, 116a; eourtesy ofM. Gowing: 77a, 91, 94b, 96, 97, 80e, 105a, 122a, 123a, 124a,
125b, 128a, 135; from Haeekel (1887): 10, 17,22,26,28,32,34,37,38,39,43,46, 49a, 53e, 56, 73, 83a, 92a, 79a, 101a, 102, 108, 111, 112, 114,
118, 119, 120, 121, 137, 145, 146, 158; from Haeeker (1908): 1,2,3,6, 12a, 13, 14, 15, 16, 18, 20b, 21b, 24b, 25b, 27, 29, 30, 31, 33, 35, 36, 40,
41,42,44,47,48, 49b, 4ge, 50, 51, 52, 53a, 53b, 55, 58, 59, 60b, 61, 68, 70, 72b, 82, 83a, 84, 85b, 90a, 93, 98, 99b, 105b, 106b, 107, 109, 113a,
113b, 115b, 130, 132, 133, 138a, 140, 141a, 141e, 142, 143, 144, 148, 149, 150, 151, 152e, 153, 155, 156; from Immermann (1904): 12b, 12e, 19,
20a; from Morley and Stepien (1985): 86, 88, 94a; from Murray (1885): 90b, 95b; from Peters (1929): 89a, b; from Reshetnjak (1966): 11, 21a,
24a, 25a, 45, 54, 60a, 62, 65, 72a, 74a, 76, 78a, 95a, 104a, 110, 115a, 141b, 147, 152a, 152b, 154; from Sehmidt (1908): 64, 66a, 67; from Schróder
(1913): 4, 5, 8, 57, 63, 69, 71, 74b, 75, 87, 106a, 126, 157; from Stadum and Ling (1969): 80b, 123b, 124b, 139a; from Tibbs (1976): 23.
Tibbs J.F. 1976. The Aulacanthidae (Radiolaria: Tibbs J.F., Tibbs S. D. 1986. Further studies on the
Phaeodaria)ofthe Antarctic seas. Biol. Ant. Seas 5, Phaeodaria (Protozoa: Radiolaria) ofthe Antarctic
Ant. Res. Ser. 2, Amer. Geophys. Union, pp. 21-65. seas. Biol. Ant. Seas 16, Ant. Res. Ser. 4, Amer.
Species deseriptions and new distributional data lor Geophys. Union, pp. 167-202. New and old dis-
this lamily in the Paeifie sector 01 the Antaretie. tributional data on most lamilies 01phaeodarians
. in the Paeific and Atlantie seetors 01 the Antaretic
are diseussed.
Ciliophora
Wolfgang Pefz
Introduction and deriving nutritive benefits from them (Laval-Peuto

and Rassoulzadegan, 1988; Stoecker et al., 1989a;
This chapter concems the aloricate or "naked" ciliates Sanders, 1991). At least one species, Myriollecfa rubra,
(for the purpose ofthis chapter, non-tintinnid ciliates), is apparently exclusively autotrophic depending on a
rather fragile organisms occurring virtually every- cryptophycean endosymbiont (Taylor el al., 1971;
where: in salt and fresh water (pelagic, benthic, inter- Lindholm, 1985; Crawford, 1989).
stitial), soil, tree bark, snow, ice brine, sewage treat-
ment plants, anoxic environments, etc. Several Reproduction of ciliates is typically by asexual binary
species are ecto- or endocornmensals but only a few fission. Many ciliates divide homeotropically (homot-
are true parasites. Some 8000 extinct and extant spe- hetogenically), i.e. the fission plane is transverse to
cies are currently known, but many more still await the main body axis, but oligotrichs divide enantiotro-
discovery (Lynn and Corliss, 1991). pically, i.e. both daughters (pro ter, opisthe) are con-
nected at their posterior ends (Fauré-Fremiet, 1953;
The Ciliophora, although a highly differentiated and Petz and Foissner, 1992). Sessile and sedentary spe-
diverse assemblage, are considered a monophyletic cies (e.g., chonotrichs, suctorians, peritrichs, follicu-
taxon. With the exception ofthe small primitive groups linids) often reproduce by budding, a kind of unequal
primociliatids (without nuclear dualism) and karyore- binary fission. The smaller bud (larva) looks very dif-
lictids (with a diploid, nondividing macronucleus), ferent from the paren tal cell and is usually a dispersal
ciliates are characterized by having two kinds ofnuclei: form (Corliss, 1979; Small and Lynn, 1985). Some
a micro- and a macronucleus. They also have few to species (e.g., colpodids, the freshwater fish parasite
many cilia or compound ciliary organelles (at least in Ichthyophthirius) divide into more than two offspring
some stages oftheir life cycle), a complex infraciliatu- (palintomy, polytomy), which usually takes place in a
re composed of single, paired, or multiple basal bodies reproductive cyst. Multiple budding occurs in some
with closely associated microtubules and microfibrils. suctorians and chonotrichs (Jankowski, 1973; Lynn
And they conjugate, which is a sexual phenomenon. and Corliss, 1991). During conjugation two speci-
Some ciliates are mouthless (e.g., astomes and mens of presumably differing mating types fuse par-
Myrionecta rubra) or polystomic (suctorians), but the tially, their nuclei also merging. After separation of
vast majority have a single feeding apparatus. Most the organisms, both exconjugants divide.
ciliates possess pellicular alveoli (unit membrane-
bound sacs beneath the plasma membrane) and tubular Ciliates have been investigated ever since Antoni van
mitochondrial cristae (Lynn and Corliss, 1991). Leeuwenhoek first discovered free-living forms with
his microscope (actually a hand lens) in 1674. Many
Most ciliates are naked, but several groups and species new species were described about 100 years later by
form a more or less robust lorica (e.g., tintinnids, this O. F. Müller, and in the early 19th century e.g., by M.
volume). Aloricate ciliates comprise several size clas- Bory de StoVincent, C. G. Ehrenberg and F. Dujardin.
ses ranging from about 10 ¡.un to 4.5 rnm, but stalked In the late second half of that century, O. Bütschli
forms and colonies may be come distinctly larger (1887 -1889) established the first comprehensive cilia-
(Corliss, 1979). They are trophically diverse, most te classification. In the early 20th century, A. Kahl
being heterotrophs (bacterivores, herbivores, camivo- produced a comprehensive guide to genera and spe-
res, detritivores) consuming a wide variety of prey cies of almost every major ciliate group, a work still
(e.g., bacteria, cyanobacteria, coccolithophorids, algae, highly valued today. Nowadays, interest in the often
fungi, flagellates, ciliates, small metazoans, organic neglected ciliates is increasing again since it is beco-
particles, and dissolved organic matter; Corliss, 1979). ming clear that they play an important role in the eco-
Some, especially planktonic oligotrichs, are mixot- system. They can be valuable bioindicators used, for
rophs, retaining functional chloroplasts from their prey instance, in assessing the quality of running waters.
Soufh Atlantic ZooplanÁ:ton.

edited by D. Boltovskoy. pp. 265-319
266 Ciliophora
Area Region System Season Abundance Biomass Reference
Southern Oc. Antarctic Open Oc. Sp 1000-5000 I Klaas (1997)

Southern Oc. 0.02-4.8 Garrison and Buck (1989)
Antarctic Open Oc. Sp, A__ 0.4-1900 +
Southern Oc. Antarcti~en Oc. Su 1000-9000 Buck and Garrison (1983)
Southern Oc. Antarctic Open Oc. Su 220-760 Froneman and Perissinotto (1996a)
Southern Oc. Antarctic Open Oc. Su 100-7000 t "-Mamaeva (1989)
Southern Oc. Antarctic Open Oc. W 0-4660 0-0.07 Gowing and Garrison (1992)
-+-
Southern Oc. Antarctic Shelf W 220-1900
---+-Garrison et al. (1993)
Southern Oc. Antarctic Bay Yearly 200-2300 t 0.5-13 Leakey et al. (1994b)
I
Southern Oc. Klaas (1997)
Subantarctic Open Oc. + Sp 1000-10 000
Southern Oc. Froneman and Perissinotto (1996a)
Southern Oc. Subantarctic ~en OCi- Su 260-650
Froneman and Perissinotto (1996b)
Pacific Oc. 0.3-11 Strom et al. (1993)
Subarcti~en Oc. Sp, S_u~_500-28 000 t
Pacific Oc. Tumantseva (1992)
Subarcti~en Oc. Su 1460-15
20-150 000
i
Subantarctic Open 0rcw
Pacific Oc. Subarctic W 400-6120 t 0.3-6 Martin and Montagnes (1993)
Atlantic Oc. Subarctic Coastal
Estuary Yearly ca. 110-55 000 Sanders (1995)
Atlantic Oc. Transition Open Oc. Sp 800-4000 Stoecker et al. (1994b)
Atlantic Oc. Transition Open Oc. Su 60-1340 Froneman and Perissinotto (1996a)
+-
Pacific Oc. Transition Open Oc. Su 90-3230 t 0.4-14 Sime-Ngando et al. (1992)
Atlantic Oc. Transition Slope Su 1900-2800 t
~cker et al. (1989b)
Baltic Sea Transition Shelf Sp 4600-42 200 t Mamaeva (1988)
Irish Sea Transition Shelf Sp, Su 21 00- 14 000 t about 1-65 Edwards and Burkill (1995)
Atlantic Oc. Transition Shelf Su 600-13000 t
~Stoecker et al. (1989b)
Pacific Oc. Transition Coastal Sp 500-45000 t 0.9-9 Beers et al. (1980)
Atlantic Oc. Transition Coastal Yearly 20-6400 Capriulo and Carpenter (1983)
Atlantic Oc. Transition Coastal Yearly 350-6000 t 0.06-5 Montagnes et al. (1988a)
Atlantic Oc. Transition Coastal Yearly 300-22 000 t Stoecker et al. (1987)
Atlantic Oc. Transition Estuary Sp 0-225000 Dale and Burkill (1982)
Atlantic Oc. Transition Estuary Yearly 0-45000 Sanders (1987)
Atlantic Oc. Transition Estuary Yearly 700-16000 t 1.4-219 Leakey et al. (1992)
Atlantic Oc. Transition Estuary Yearly Dolan and Coats (1990)
1800-17 200 t + about 4-12
Baltic Sea -L Transition . Estuary Yearly Smetacek (1981)
0-92 000 t t <1-56
Dale and Dahl (1987a)
Sp --+-
North Sea
Atlantic Oc.
Atlantic Oc.
Transition----:=
Trans./subtrop.
Trans./subtrop.
Bay
Open Oc.
Open Oc. I
Sp
Su
1300-1 360 000
0-4000 t
<30 to > 1000 t
¡ Mamaeva
Mamaeva
(1984a)
(1982a)
Sp,Su 100-5000 t 0.1-2.3 Lessard and Murrell (1996)

Atlantic Oc. Subtrop~pen Oc+-
Su 330-920 0.1-0.3 Beers et al. (1975)
Pacific Oc. ~ubtropic Open Oc.
Adriatic Sea Subtropic Coastal Su 160-1960 t Revelante and Gilmartin (1990)
Adriatic Sea Subtropic Coastal Yearly 20-39300 0.02-37 +-Revelante and Gilmartin (1983)
Indian Oc. Tropic Open Oc. Su, A 5-610 t Mamaeva (1982b)
Indian Oc. Tropic Open Oc. A 30-820 t

0.1-1.2 ~key et al. (1996)
Pacific Oc. Tropic Open Oc. W 120-720 Beers and Stewart (1971)
Atlantic Oc. Tropic Open Oc. Su 0-7200 (mean 534)

J'Aargalef (1973)
Atlantic Oc. Tropic Upwelling Su 0-3500 (mean 978) Margalef (1973)
Caribbean Sea Tropic Coastal Yearly 970-3930 0.3-2.1 Lynn et al. (1991)
Ciliophora 267
COPEPOD
/ NAUPlIUS
Myrionecta PHOTOSYNTHETIC
rubra DIATOM MICROFlAGEllATE
DINOFlAGEllATE -........ \
BACTERIOPlANKTON
Fig.1. Trophic pathways involving marine ciliates. From Pierce and Tumer (1992).
Ciliates are al so used as ideal model organisms in cell and fish larvae. Thus, they are an important link
biology and are popular study objects in molecular between lower trophic levels and the larger zooplank-
biology and biochemistry. ton consumers (Fig. 1; e.g., Pomeroy, 1974; Taylor,
1982; Azam et al., 1983; Porter et al., 1985; Jonsson,
Because of their fragility and lack of a pennanent ske- 1986; Sherr et al., 1986; Albright et al., 1987;
leton, naked ciliates are under-represented in marine Fenchel, 1987, 1988; Sherr and Sherr, 1987, 1988;
plankton samples and were thus largely neglected in Rassoulzadegan et al., 1988; Stoecker et al., 1989b;
the pasto However, they make up the bulk ofthe cilia- Bernard and Rassoulzadegan, 1990; Lynn and
te cornmunity in the pelagic realm and can be up to 50 Montagnes, 1991; Pierce and Tumer, 1992; Lonsdale
times more abundant than the tintinnids (e.g., et al., 1996). But as species identification is someti-
Smetacek, 1981; Tumantseva and Kopylov. 1985; mes difficult and time-consuming (e.g., because of
Dale, 1987; Pierce and Tumer, 1992) reaching densi- silver impregnation), ecological investigations were
ties as high as 1.36 x 106 ind.!1 (Dale and Dahl, 1987a; often restricted to size-classes or even communities.
Table 1). Recent investigations show that aloricate Species composition and distribution are still not
ciliates occur in high numbers and consume signifi- well-researched.
cant quantities of autotrophic and heterotrophic
microbial production. They are eaten in tum by other
protozoans, many metazoans (especially copepods),
..
Table 1. Range of abundance (ind. 1-1; data including tintinnids are denoted with "1") and biomass (mg e m-3) of planklonic ciliales in various
areas.
268 Ciliophora
Methods Plankton nets and screens are unsuitable for quantita-

tive sampling because a large proportion of the indi-
Provenance and collection of materials viduals is lost (Gifford, 1985). In nets of 20 and 44 Ilm
mesh size, for instance, losses of aloricate ciliates
For enumeration of aloricate ciliates only whole-water ranged from 74-98% (Brownlee and Jacobs, 1987).
techniques are recornmended (Gifford, 1985; Finlay These simple tools may thus only be utilized for a
and Guhl, 1992; Pierce and Tumer, 1992). Usually, qualitative taxonomic survey. For this, a mesh size of
several hundred millilitres to some litres of seawater 10 Ilm or less is necessary to catch the many smaller
are collected by various types of water bottles aloricate species. As ciliates are often slender and
(Venrick, 1978b). Whereas sma11 capacity bottles are very flexible, even 50 Ilm long specimens can pass
adequate for biomass-rich inshore waters, larger bot- through the pores of a 10 Ilm screen (Foissner et al.,
tles are necessary for the oligotrophic open ocean. 1991). So nets have to be towed very gently. Sediment
However, the latter usua11y lack mixing devices, so traps are equa11y unsuitable beca use they collect the
grave errors in abundance estimates may result from ciliate population incompletely (Antia, 1991), and
under-representation of particles settling rapidly in species may multiply or disappear in the containers.
subsamples (Gardner, 1977). Containers provided with
windows, like Van Dorn bottles, pose the problem that Sample preparation and analysis
light-sensitive species may move to or away from
them. which can bias subsampling. Standard Niskin The following are only some general comments for
bottles may be toxic to protozoa, therefore ultraclean the determination and enumeration of ciliates. For
techniques or GoFIo bottles should be used instead accurate species identification, in vivo observations
(Fitzwater et al., 1982; Chavez and Barber, 1987). are necessary, preferably with interference contrast.
Important characters, e.g., movement, cortical gra-
For elucidation ofthe vertical or horizontal micro-dis- nules and their colour, presence and shape of extru-
tribution, special water samplers take several samples somes, are often not preserved or do not stain in fixed
simultaneously at 10-20 cm intervals (Blaker, 1979; material. However, an experienced researcher should
Owen, 1981). They collect only rather small amounts be able to identify many species in live samples, at
of water, but if used in a chlorophyll-maximum or least after some preliminary work in the area, which
density discontinuity, the number of organisms may considerably shortens sample processing. Sometimes,
be adequate for meaningful data (Paranjape, 1991). particularly for taxonomic investigations, determina-
Macro-aggregates and associated ciliates are best col- tions have to be verified by sil ver impregnation.
lected by scuba divers or submersibles (Silver et al., Several techniques are available for this, of which the
1978; Caron et al., 1982). protargol method is the most universal. These proce-
dures and some modifications as well as preparation
Pump samplers have to be chosen according to mode for scanning electron microscopy are described in
of operation and study requirements (Beers, 1978; detail by Foissner (1991) and in the "Protocols in
Miller and Judkins, 1981). They can be used either at Protozoology" (Lee and Soldo, 1992).
discrete depths or for sampling over vertical, oblique,
or horizontal paths. Additionally, they can be fitted Traditiona11y, however, a variety of fixatives is utilized
with continuous-flow sensors for measuring physical in different concentrations in protozooplankton studies:
and chemical parameters (Paranjape, 1991). Acid Lugol's iodine: 10 g iodine and 20 g potassium .
Mechanisms also have been developed for keeping iodide dissolved in 200 mI disti11ed water, then add 20
the pump-intake at constant depth even when the ship mI glacial acetic acid; used in 0.4-10% final concen-
is ro11ing (Paranjape, 1991). While pumps are capable tration, precludes most silver impregnation methods
of co11ecting a11 size-classes of ciliates, ce11s may and epifluorescence microscopy;
become damaged or lost during concentration and fi- Bouin: 150 mi saturated aqueous picric acid, 50 mI
xation (Sorokin, 1977, 1981; Dale and Burki11, 1982). formaldehyde, 10 mI glacial acetic acid; prepare
Pumped samples thus underestimate the ciliate popu- immediately before use;
lation. However, compared to bottles or nets, they Modified Bouin: buffered concentrated formalde-
may yield much better spatial resolution. hyde saturated with picric acid, add glacial acetic acid
Ciliophora I 269
immediately before use so that the final concentration and may cause losses of over 75% (Gifford, 1985;
of acetic acid is 1% (v/v) after addition ofthe sample; Laval-Peuto and Rassoulzadegan, 1988).
use a fixative:sample ratio of 1:19 for brackish and of
1:10 for open ocean water (Lee el al., 1985); AII fixatives change the volumes of ciliates; i.e. they
Glutaraldehyde: 1-6% final concentration; addition usually shrink. Thus, the biomass derived from pre-
of 2% tannic acid reduces cell shrinkage (Choi and served samples is considerably lower than the stand-
Stoecker, 1989); ing stock ifused without correction. Pierce and Turner
Formaldehyde: 0.4-2% final concentration, buffered (1992) concluded that the error caused by cell shrink-
with hexamethylenetetramine; age and loss from fixation and handling may under-
Mercuric chloride: 380 mI saturated aqueous mercur- estimate the true aloricate ciliate biomass by at least an
ic chloride (60 g mercuric chloride dissolved in 1 1 of order of magnitude. Compared with the live vo1ume,
boiling distilled water), 100 mI formaldehyde, 30 mI formaldehyde shrinks cells the 1east, viz. 10-20% on
glacial acetic acid; used in 2.5% final concentration. average. Lugol's solution and glutaraldehyde decrease
cell volumes by 13-45% (Choi and Stoecker, 1989;
Although widely used, fonnaldehyde is particularly Ohman and Snyder, 1991; Leakey el al., 1994a). But
inappropriate for preservation of marine ciliates mercuric-chloride-fixed cells are about 20% larger
because most specimens burst (up to 72% los s relative than those preserved in 2% acid Lugol (Sime-Ngando
to Lugol), and many are strongly distorted (Pace and el al., 1992). Similarly, 5% Bouin usually shrinks cells
Orcutt, 1981; Reve1ante and Gilmartin, 1983; Leakey less than 10% Lugol or the quantitative protargol stain
el al., 1994a; Stoecker el al., 1994a). Fewest losses (Jerome el al., 1993; Stoecker el al., 1994a).
occur with Lugol's iodine, higher concentrations, e.g., Consequently, the ratio of cell carbon content to vo-
10% or 20%, usually yielding more ciliates (Stoecker lume depends on the fixative used and its concentra-
el al., 1994a). This fixative is followed by Bouin (up tion. Putt and Stoecker (1989) experimentally deter-
to 46% loss relative to Lugol), and glutaraldehyde (up mined carbon contents in preserved ciliates at 0.19 pg
to 66% loss; Revelante and Gilmartin, 1983; Leakey C ¡.rm-3 in 2% Lugol, and 0.14 pg C ¡.rm-3 in 2% for-
el al., 1994a; Stoecker el al., 1994a). Particular1y 1ess maldehyde, whereas an average of 0.11 pg ¡.¡m-3is
destructive is mercuric chloride but it is costly and suggested for living specimens (Turley el al., 1986).
highly toxic (Pace and Orcutt, 1981; Laybourn-Parry,
1992). In contrast to seawater, the most destructive Particular attention should be paid to the method of
fixative in freshwater is Lugol's solution, causing enumeration, since a formally adequate sampling pro-
58% 10ss after nine months, followed by glutara1de- cedure cannot replace an inappropriate counting tech-
hyde (29%), fonnaldehyde (24%), and mercuric chlo- nique. The most accurate method is live counting of
ride (15%) (Sime-Ngando and Groliere, 1991). unconcentrated water samples as demonstrated by
comparative studies (Sorokin, 1977, 1981; Dale and
From the above mentioned studies it may be conclu- Burkill, 1982; Sime-Ngando el al., 1990). The last
ded that the effects of fixatives vary with concentra- study also found that live counting of filtered samp1es
tion, salinity of sample, ciliate species, perhaps the provided good results (but see above). For enumera-
organism's nutritive state and apparently the duration tion, the water samp1e (0.4-30 mi) is put in a Dolfuss
of samp1e storage. A further problem with Lugol is or other shallow counting chamber or a Bogorov
that it stains ciliates and detritus the same colour, counting tray. The entire sample or a fraction is exam-
which may cause significant counting errors in the ined using a stereo microscope at 25-60x magnifica-
presence of abundant particulate organic matter (Pace tion and bright or dark field illumination (Dale and
and Orcutt, 1981). Burkill, 1982; Sorokin, 1982; Sime-Ngando el al.,
1990). Shallow counting chambers with a lid are ge-
There are several methods for concentration. But like nerally well suited for shipboard use, but the Bogorov
fixatives they can severely reduce aloricate ciliate num- tray may be used only on firm ground. There, and if
bers (Sorokin, 1981). Filtration of samples, which is conditions are fairly stable on a big ship, the simplest
widely used, may decrease the abundance of aloricate counting method is to transfer the sample dropwise
ciliates by up to 95% (Sorokin, 1977; Gifford, 1985). onto a c1ean slide and inspect it under a compound
Reverse filtration can be only slightly less destructive, microscope at 40x magnification.
270 Ciliophora
Live counting is a quick, well tested, and cheap me- dances are about 13% lower than with direct counting
thod for the enumeration of ciliates (Dale and BurkilL (Montagnes and Lynn, 1987).
1982; Sorokin, 1982; Foissner, 1983; Lüftenegger el al.,
1988; Sime-Ngando el al., 1990). Counting of 1 or 5 mi Especially small ciliates are often counted using epi-
samples takes 15-120 minutes (Dale and Burkill, 1982; fluorescence microscopy (Sherr and Sherr, 1983).
Sime-Ngando el al., 1990). The volume ofwater inves- Aliquots of preserved samples are filtered onto black
tigated depends on the concentration of ciliates, i.e. a polycarbonate filters and stained with DAPI, profla-
smaller volume (about 1 mi) is sufficient in near-shore vin, fluorescein isothiocyanate (FITC), or acridine
rich waters, but if specimens are scarce «5 ind./ml), as orange. This permits distinction between autotrophic
in the oligotrophic open ocean, a larger volume (about and heterotrophic cells as specimens containing plant
5-15 mi) has to be counted. An advantage oflive count- pigments fluoresce either red or orange depending on
ing is that it allows species identification. This requires the type of pigment present (Stoecker el al. 1989b).
some experience, but in contrast to preserved samples But species identification is usually impossible. The
important morphological characters (shape, size, use of electronic counters like the Coulter Counter
colour, extrusomes, movement) are not lost or distorted, requires considerable experience and thus often yields
which makes correct determination much easier. unrealistic figures (Sheldon, 1978).
However, a crucial drawback of live counting is that it
has to take place irnmediately after sampling, i.e. at least
on the same day. This limits considerably the number of Geographic and vertical distribution
samples that can be taken and processed.
Not much is known about the biogeography of marine
More cornmon than live counting of planktonic ciliates aloricate ciliates. Since many specimens become lost
is enumeration of fixed and concentrated material using or severely distorted in traditionally fixed plankton
an inverted microscope following the procedure of samples (see above), they usually cannot be identified.
Utermohl (1958). However, this method is less satis- The Iimited data available show that the ciliate com-
factory beca use almost all aloricate ciliates may have munity is often dominated numerically by oligotrichs
been lost during prior preservation and concentration of the genus Slrombidium (e.g., Beers el al., 1971;
(see above). Furthermore, as specimens are frequently Mamaeva, 1984a,b, 1986; Tumantseva and Kopylov,
shrunk, distorted, darkly stained, or masked by detritus 1985). The distribution of the species recorded in the
in the settled samples, identification to genus or species South Atlantic Ocean is given in Table 2.
level is usually impossible (Sorokin, 1977; Pace and
Orcutt, 1981; Leakey el al., 1994a). Geographic patterns
An altemative to counting fixed samples is the quan- As with phytoplankton and bacterioplankton, den-
titative protargol stain of Montagnes and Lynn sities of aloricate ciliates decrease from coastal
(1987), which also allows preliminary species iden- regions towards the open ocean (Table 1). Peak num-
tifications. The organisms are fixed with Bouin and bers and biomass generally occur in bays and estuar-
concentrated on a Millipore cellulose filter (0.8-3.0 ies. But high densities are also found in nutrient-rich
!lm pore size; suction <100 rnm Hg). The material on upwelling areas and divergences in the open ocean,
the fi1ter is then embedded in agar, stained and count- whereas zones of convergence yield particularly low
ed. By this, a permanent record of the cornmunity is values (Mamaeva, 1982b, 1986; Porter el al., 1985;
obtained. However, as in fixed samples, characters Layboum-Parry, 1992). This suggests that ciliate dis-
present solely in living individual s are lost and defi- tributions depend more on factors like food supply
nitive species determinations are often impossible. than on geographic latitude or ocean basins. This is
Another disadvantage is that the procedure is costly, corroborated by positive correlations between ciliates
time consuming, and not suitable for routine treatrnent and chlorophyll concentration and, less frequently,
of many samples. However, the modification by bacterioplankton biomass (for review see Lynn and
Skibbe (1994) is distinctly quicker (4 hours/prepara- Montagnes, 1991). It must be noted, however, that
tion) and works also with Lugol-fixed cells. The effi- differences in numbers between studies may also be
ciency is similar to that of settling chambers but abun- caused by differing methods (see above).
Ciliophora
Tropical/Subtropical Transitional Subantarctic Antarctic

Species Reference
C c S e N e ~ r c ~ B eN 0 N O N 0
-
r--- -7 7 - 7 - -- 8
Acineta tuberosa
Actinotricha saltans - - - + - 1 - - I - , -
- - - + - - ,
Aegyriana oliva - I -
Cohnilembus verminus
Condylostoma patens
Corynophrya campanula 1 - - - - - 1 - - 1 - - 1 - 1 - + * 3
- - - -
PAP
- + - - I -
Didinium gargantua
Didrnium sp. 4 ~ 7 --
Diophrys appendiculata
Dysteria brasiliensis
Dysteria compressa
Dysteria monos tyla
-
-
-
-
-
-
-
-
-
+
+
+
-
-
-
-
-
-
' 1.2
Enchelyodon laevis
P- - - - - - + - , - - I
Euplotes charon
- - -
-
-
-
-
- + - -
Euplotes harpa
- - -- I - -
- -I + - - - I - 1,2
Euplotes vannus - - - - - - + I 8
Euplotes sp. - + - - I + i 3
Gruberia lanceolata - - - +
Holophrya pelagica + - 1 - +I
I
Holosticha diademata -- -- - L
Hypotrichida gen. sp. -
-- - - -
Kentrophyllum setigerum
- -
- - - +
Laboea strobila -P
- - - -
Laboea sp. - - - - - - - 1 -
I
Leegaardiella spp. - - - - - -
Licnophora auerbachi - - - + - - 1,2
Litonotus cygnus I- - - I + - - - 1 - - I 1,2
- - - + --
I
Litonotus fasciola 1 - - - - - - - 1 f2
Lohmanniella spp. - - - I - - 1 _ _ + - ~ - 10
- 1I -
Mesodinium acarus -
- --- -- -- 1,2 -
Mesodinium pulex 1,2

11-1 - +, - -
- - - -
- -
+
-
~ - 1
;I -- 4
272 Ciliophora
~ Species
Tropical/Subtropical
G c S e N e B r l ~ ~ B eN
Transitional
0
Subantarctic
N , 0
Antarctic
N 0
Reference
- + - -
I Pseudocohnilembus pusillus
Stentor polymorphus
I Strobilidium typicum
- -
Strobilidium sp.
-- - -- --
Strombidium conicoides
~trombidiurnconicum
Strombidium spp.
fiarina fusus
Vorticella ocean1
I
Zoothamnium elegans
Be: Benguela Current; Br: Brazil Current; Cg: Central Gyre; Gc: Guinea Current;
N: neritic; Ne: North Equatorial Current; 0: oceanic; Se: South Equatorial Current.
*: exact location uncertain
Table 2. Records of planktonic aloricate ciliates in the South Atlantic Ocean (+: found, -: not found). References: 1: Faria et al. (1922); 2: Pinto
(1925); 3: Schroder (1906); 4: Leakey et al. (1994b); 5: Hada (1970); 6: Mamaeva (1989); 7: Klaas (1997); 8: Petz, Song and Wilbert (unpubl.);'
9: Lohmann (1920); 10: Fonda Umani and Monti (1991); 11: Horstman (1981); 12: Petz et ul. (1995a); 13: Faria and da Cunha (1917).
Aloricate ciliates are often distinctly (up to 50 times) cate ciliates is rather similar in nearshore and open
more abundant than the loricate tintinnids (e.g., Beers ocean areas. Oligotrichs like Strombidiunz, or occasion-
et al., 1980; Smetacek, 1981; Mamaeva, 1982a; Dolan ally Lohmanniella or Tontonia, usually dominate (e.g.,
and Marrast, 1995). It appears that the percentage of Beers et al., 1975; Mamaeva, 1983, 1984a,b, 1986;
aloricates is slightly lower in neritic waters, where tin- Revelante and Gilmartin, 1983; Tumantseva and
tinnids even dominate occasionally, and increases Kopylov, 1985; Stoecker et al., 1994b; Dolan and
towards the open ocean (Margalef, 1973; Revelante, Marrast. 1995). For instance, in the Atlantic sector of
1981; Capriulo and Carpenter. 1983; Paranjape et al.. the Southern Ocean the oligotrich genera Stronzbidium,
1985; Leakey et al.. 1994b; Sanders. 1995). At least at Laboea and Strobilidizrm make up 83-99% of total cili-
the generic level, the community composition of alori- ate carbon (Klaas. 1997). However. species-specific
Ciliophora 273
uwOwee ~
e~ ~ ~0.2
U0.8
§w ~~uOu~-
~
- ~
U ~O
ro
ro w 0.2
~
ro
OA
0.8
0.6
E 1~
0.4 0.0
0.6 1.0
ro
ro
O~ ~
C)
D>20flm
D 10-20flm
1<10flm
8 14 20 27 5 9 15 21 5 11 19 25 17 20 25
May June August September
Fig.2. Size distribution of integrated ciliate abundance (A) and biomass (8) for 16 vertical profiles (0-80 m) in the subarctic Pacific (cells
sorted by width). From Strom el al. (1993).
distributional data are very scarce. These would be in 10w productivity zones, and decreases with depth
especially interesting since Borror (1980) doubted that (Fig. 3; Beers and Stewart, 1969; Margalef, 1973;
individual marine species have a cosmopolitan distri- Revelante and Gilmartin, 1983, 1990; Mamaeva,
bution. 1984b, 1986, 1990; Paranjape el al., 1985; Edwards
and Burkill, 1995; Froneman and Perissinotto,
Data on dominance of single species are rare since 1996a). There may be one or sometimes two discrete
only genera or size classes are often differentiated. abundance maxima. The first occurs at or near the sur-
From the very few studies available it appears that, as face (0-25 m) and the second at the thermocline if it is
in other zooplankton groups, rather few aloricate spe- marked (Beers and Stewart, 1969; Margalef, 1973;
cies are abundant in any given sample. For instance, Mamaeva, 1986, 1990; Sime-Ngando el al., 1992).
in the open Baltic Sea 1-3 species contributed > 10%
to total ciliate abundance, 0-3 taxa between 5-10% The vertical distribution of ciliate abundance ancl/or
and 50-80% of the species contributed less (Leppanen biomass frequently reflects that of the phytoplankton,
and Bruun, 1986). Small ciliates «20-30 Ilm) often and less distinctly that of bacterial biomass, indicating
dominate numerically, whereas larger species contri- a predator/prey relationship (Fig. 4; Beers el al., 1980;
bute more to biomass (Fig. 2; Beers el al., 1980; Strom el al., 1993; Dolan and Marrasé, 1995; Leakey el
Revelante el al., 1985; Sherr el al., 1986; Sushin el al., 1996). Mixotrophic species and the phototrophic
al., 1986; Revelante and Gilmartin, 1987; Lynn el al., Myrionecla rubra occur predominantly in the upper
1991; Strom el al., 1993; Lessard and Murrell, 1996; layer but are capable of diurnal vertical migrations
Tamigneaux el al., 1997). (Dale, 1987; Stoecker el al., 1989b; Dolan and
Marrasé, 1995). The latter species, for instance, usual-
Vertical profiles Iy occurs at the surface layer during most ofthe day and
migrates down to 30-40 m at night (Crawford, 1989).
The majority of ciliates is usually found in the upper Dolan and Marrase (1995) even found a M. '-lIbra peak
40 m in highly productive areas and between 50-75. m at 80 m depth. At the surface and during red-tide
274o~9 oOOl/")
N "'" qoq 10 N
o22
l/")OO '"CD
oN o
O 60
Ciliophora100
20
..r: Q. 100
Q)
o
140
Fig.3. Vertical distribution of ciliate biomass (mg m-3) at several stations in the Antarctic zone of the Atlantic Ocean (A), and in the central
part ofthe Indian Ocean (B). From Mamaeva (1990).
20o40
N
~o80
co
60
100
40
80 ~
160
40
10
20
120
30 oo oqbiomass
'"Chlorophyll
l/")
l/") OprofiJes ofStation 4C 1-1) biomass86and chlorophyll
Station
Station
owestem
------
---+-
O(í
...s::
Indian
Depth
80
120
Q)
euphotic
Vertical
Ciliate
Bacterial
Ocean.
of zone From
biomass
a (xl ciliate
0-2,C fJg
(fJg and
1-1)
Leakey el al. (]996).
bacteria] a concentration at 3 stations of contrasting productivity in the north-
o
E
200
40
160
Ciliophora 275
blooms, when it reaches up to 254 x 106 ind.!l, M. rubra est classifications may be regarded only as transient
may be the numerically dominant species (Fenchel, and are likely to change again shortly.
1968; Leppiinen and Bruun, 1986). Too few data are
available to define depth zonations for other species. The following illustrated glossary of terms widely
used for describing ciliates is based mainly on compi-
lations by Corliss (1979), Corliss and Lom (1985),
Taxonomy and identification Foissner el al. (1995), and Foissner and Dragesco
(1996a). Capital letters (in parentheses) denote the
Classification systems group for which the term is used (C: cyrtophorids; G:
gyrnnostomates; H: hypotrichids; K: karyorelictids; O:
The first comprehensive system of ciliate classifica- oligotrichs; PI: pleurostomatids; Pt: peritrichs; S:
tion was proposed by Bütschli (1887-1889) in the late suctorians; Se: scuticociliates).
19th century. He, like Kahl (1930-1935) who replaced
his system about 50 years later, used mainly the type Adhesive disc (Fig. 8.39B): basal disc; cup-shaped
and variation in the oral and somatic ciliature as main organelle at the posterior pole used for attachment
characters for distinguishing higher taxa. Subsequent to a substrate.
systems have changed more rapidly. In the mid-20th Adoral zone of membranelles (Fig. 6E; 7A, C, D):
century, after silver impregnation methods became band of membranelles (polykinetids) along left
popular, the scheme was revised e.g., by Fauré- side of oral area.
Fremiet (1950-1970) using infraciliary and morpho- Basal body (Fig. 6A): subpellicularly located basal
genetic data. Fauré-Fremiet's scheme of the ciliates termination of cilia, cylindrical and composed of 9
was later adopted by Honigberg el al. (1964) in their peripheral microtubular triplets; may be non-cili-
major revision of all protozoa. ferous.
Bristle kinety (K) (Fig. 5A): row of modified, often
Since the beginning ofthe 1970's, morphogenetic cri- rather irregularly arranged dikinetids with stiff
teria, particularly the formation of the oral apparatus, cilia, bordering the margins of the glabrous stripe.
and ultrastructural data were employed increasingly to Brosse: see dorsal brush.
distinguish higher categories. De Puytorac el al. Brosse cleft (K) (Fig. 5A): narrow slit in left surface
(1974), for instance, used mainly the fine structure of of head where brosse cilia emerge; produces right
the oral area to construct a system. A more compre- and left lip of oral bulge, possibly covered by pel-
hensive set of characters was utilized by Corliss Iicle of oral bulge.
(1979) in a subsequent attempt, which gained wide- Caudal cilium (Fig. 6C): distinctly longer somatic cil-
spread recognition. With a growing number of taxa, ium at or near posterior pole of body.
more available ultrastructural data, and recognition of Ciliary row: see kinety.
increased ciliate diversity, the classification had to be Ciliature: total assembly of oral and somatic cilia or
altered again. Consequently, the schemes provided cirri; may be arranged in patches, kineties, mem-
e.g., by Levine el al. (1980), Small and Lynn (1985, branes or membranelles.
who used mainly the structure of the somatic kinetid), Cirrus (H) (Fig. 7C): compound somatic ciliature.
de Puytorac el al. (1987), or de Puytorac (1994) intro- Cluster of cilia functioning as single unit but with
duced many new higher taxa, elevated existing groups no common enveloping membrane. Subtypes
to higher ranks, and transferred others to far distant identified by location: buccal, frontal, frontotermi-
categories. nal, frontoventral, midventral, pretransverse,
transverse, marginal and caudal cirri.
In modem systems, a combination of characters, like Contractile vacuole (Fig. 5D; 6B, D): peripherally
details of the somatic and oral infraciliature, fine located expulsion vesicle serving as osmoregula-
structure ofthe cortex, stomatogenesis and other mor- tor in the cytoplasm. Generally pulsates with re-
phogenetical features, macronuclear characteristics, gular frequency emptying contents to exterior via
life cycle information and, if available, molecular bio- one or more pores.
logical data (e.g., subunit rRNA sequences), are used Corte x (Fig. 7F): complex outer layer of body inclu-
in reconstructing phylogenetic trees. But even the lat- ding pellicle and infraciliature.
276 Ciliophora
m
\\1l
extrusomes
(toxicysts) ~ ,1"',."."
preequatorial
ciliary
'~wreath
" 111 ¡..~
,[1///// / / / / / 11 \\\\'\\"-- equatorial
Id 111 \\ \ \ \ \ \ \ ~~~~{h
nematodesmata
Mesodiniid ciliate
contractile
vacuole
circumoral left
l11acronucleus klnety frontal
~ ~ fleldof
fiel<f
right
of '" ::1 ~ ~ klnetles
kineties
contractile
vacuole _-;-~ ~~~~
stylus-_~,
b'~k::'----*"
~~~ ,~
-". "- ¡:~;:'
..
,~ nematodesmata
Ciliophora 277
Cortical granules: smaIl, often refractile granules in tures. Numbering of kineties is clockwise araund
periphery of cell, sometimes corresponding to body when viewed from apical pole with number
extrusomes (e.g., mucocysts). Size, colour, shape 1 being the rightmost postoral row. In a broad
.and arrangement are important species character- sense synonym with ciliary row or basal body row.
istics. Lorica (Fig. 6B): extracellular test, envelope, case, bas-
Cyrtocyst (Fig. 5B): type of extrusome found subpel- ket or sheIl usuaIlY with opening at one, occasion-
licularly in Didinium. aIly both ends, either attached to substrate or carried
Cytoproct (Fig. 6C): cytopyge, ceIl anus; generaIly about (see chapter "Tintinnoinea" in this volume).
permanent slit-like opening through peIlicle near Macronucleus (Fig. 5B, 6B, E): somatic nucleus, ful-
posterior end ofbody. fiIls task of normal ceIl metabolism and is site of
Cytostome (Fig. 6C): ceIl mouth, oral opening; when RNA synthesis; may be multiple, is usually much
present, two-dimensional aperture most common- larger than micronucleus.
Iy permanently open, through which food passes MembraneIle (Fig. 5e, 7A, B): compound structure
¡nto the cytoplasm. composed of many cilia and associated with oral
Dikinetid (Fig. 6C): morphological complex of two apparatus.
neighboring (paired) basal bodies plus associated Micronucleus (Fig. 6B, E): generative nucleus; site of
cilia and infraciliary organeIles. General term for genetic recombination; plays major role in sexual
all kinds of paired basal body situations. phenomena like conjugation. May be multiple;
Dorsal brush (C, G, K) (Fig. 5B, 6A): one or several typicaIly much smaIler than macronucleus.
short, oblique kineties near anterior end of cell. Monokinetid (Fig. 6C): unpaired basal body plus ci-
Endoral membrane (H) (Fig. 7C): row of cilia extend- lium and associated infraciliary organeIles.
ing on bottom and right border of oral cavity. Mucocyst: type of extrusome, sacciform, rod-shaped
Extrusome (Fig. 5B, 7A): membrane-bound, subpeIli- or of paracrystaIline structure, forms mucus-like
cularly located extrusible body ofvarious, probably mass after discharge and very likely has protective
nonhomologous types (e.g., cyrtocysts, mucocysts, function.
pexicysts, toxicysts, trichocysts). Discharge typi- Myoneme (Fig. 6D): contractile fibrils (uniceIlular
cally occurs after chemical or mechanical stimula- equivalent of muscle) easily recognizable in peri-
tion, used to capture prey or for defence. trich stalks.
Frontal plate (Se) (Fig. 6C): nonciliferous, often trans- Nematodesma (Fig. 5B, D, E): rod-like bundle of
versely truncate apical area. microtubules arising fram basal bodies; reinforce
Germinal kinety (Pt) (Fig. 6E): line of nonciliferous waIl of oral basket of gymnostome, hypostome
basal bodies associated with the terminal portion and some other ciliates. AIso termed nematodes-
ofthe paroral membrane (haplokinety). mal rods.
Glabrous stripe (K) (Fig. 5A): more or less wide, Neoformation organeIle (O) (Fig. 7A): permanent
nonciliated area in median of left surface, bor- tube-like structure in which the oral anlagen ori-
dered by bristle kinety and often rippled in con- ginate during morphogenesis.
tracted specimens. Nuclear capsule (K): nuclei enclosed by special mem-
Infraciliature: total assembly of somatic and oral basal brane forming single, tight aggregate.
bodies and their associated subpellicular microfi- Oral basket (Fig. 5E): funnel-shaped cytopharyngeal
brillar and microtubular structures. apparatus consisting of nematodesmata; designat-
Kinetosome: see basal body. ed cyrtos or rhabdos according to origin of micro-
Kinety (Fig. 5E, 6A, C): structuraIly and functionaIly tubules.
integrated row of single or paired basal bodies (not Oral bulge (Fig. 5A): nonciliated, often very hyaline
all necessarily ciliferous) often running longitudi- praminence on anterior end of head surrounding
nally, plus their cilia and associated cortical struc- oral cavity.
4
Fig.5. Main features of a ciliale cell and characlers used for laxonomic identification (basal bodies are usually represented as dots). A: from
Foissner and Dragesco (l996b); B, D, E: from Petz el al. (l995a); C: from BOITor (1972a).
278 Ciliophora
Adult suctorian
ciliate
~stalk
Anterior left and

right side of a litonotid ciliate
frontal
plate~
adoral 1-:--,':",":
adoral
membranelle'
dikinetid
membranelle
~?\""'"
~/,"':
2 ~{:
: ; ~ ",
: ,:/ " ~
"
""""':--- somatic
contractile
vacuole
membranelle 3 ~\" :,: ,: '
paroral
adoral
membrane --r-"í , , ( '\'
,C:?,' : :
'
:,' ! kinety
paroral
membrane
1" .~J
""';"
Cytoproct~\, , , " cytostorne
, , ' .. ' '~monokinetid
C)
.:....=.:~_-~~
~ (haplokinety)
adora I zone of
'~ ••.•
:-- ~ mernbranelles
ru:y¡-::'!'f<' •.•,..,
~~~' •...•..
"'.~ \
,)/~~erminal (polyklnety)
micronucleus '':;,!,~ Ij-r!:¡~ adoral

',':;:] ~~
¿'.:. l' (peniculi)
'membranelles '-3
Ventral view of a scuticociliate ~)'

.~...U~:/
...
-:,,:',~~
~•..~~
\:~,
\
..
.//
anlage of
aboral
, Inety
macronucleus
ciliary wreath
Left (D) and right (E) lateral views of a peritrich ciliate

Ciliophora 279
Oral cavity (Fig. 7A): quite deep, secondarily f1at- Spine: pointed structure, either part of or secreted by
tened, or everted pouch or depression containing cell.
bases of oral ciliature and leading to cytostome- Spica (PI): median suture in anterior body half,
.cytopharyngeal complex. formed by converging of successively shortened
Oral con e (Fig. 5C): everted cytopharyngeal appara- somatic kineties.
tus of didiniids. Stalk (Fig. 6B, D, E): pillar- or thread-like attachment
Oral disc (Fig. 8.39A): apically located oral area, structure at posterior end usually of a single spe-
when conspicuously separated from rest of body. cimen; may be contractile and branching.
e.g., in Licnophora. Stomatogenesis: formation of new oral apparatus in
Oral process (Fig. 5C): tentacle-like projections, e.g., dividing cells.
in Mesodinium (extrusomes?). Stylus (C) (Fig. 5D): podite, styloid process; conical
Papilla (Fig. 6B): pellicular or extrapellicular wart- foot-like appendage projecting from posterior ven-
like bumps or small protuberances on cell surface. tral surface used for temporal attachment to a sub-
Pararal membrane (Fig. 6E, 7C): single or multiple strate.
raw of cilia lying along right side of oral cavity, in Subpellicular platelet layer (O) (Fig. 7A): thickened
hypotrichs along outer right margin of oral cavity pellicle composed mainly of polysaccharide plate-
on cell surface. lets covering posterior body portion.
Pellicle (Fig. 7F): outermost "living" layer compris- Swarmer (Pt, S) (Fig. 8.26B): free-swimrning disper-
ing cell membrane, alveoli and epiplasm. sive stage of sessile adult.
Peniculus (Fig. 6E): type of adoral membranelle. Telokinetal: type of stomatogenesis in which forma-
Perioral kinety (Fig. 6A): ciliature around and in the tion of new oral apparatus occurs principally by
vicinity of the cytostome. direct involvement of basal bodies at the anterior
Peristomial collar (Pt) (Fig. 6D): apical lip circum- extremities of all or some of the soma tic kineties;
scribing retractable peristomial disco the resulting oral areas are essentially apical or
Peristomial disc (Pt) (Fig. 6D): retractable, nonciliat- subapical.
ed, vaulted centre of concentric peristomial field. Tentacle (S) (Fig. 6B): extensible and retractable
Peristomial field: entire oral area with encircling ci- tubular extension of body mostly stiffened by
liature (e.g., adoral zone of membranelles). microtubular bundles, contains extrusomes at its
Pexicyst (Fig. 5B): type of toxicyst-like extrusome in capitate tip, and vesicles for formation of food
didiniids which, on discharge, adheres to pellicle vacuo le membranes. Used in prey capture and as
of prey without subsequent penetration. ingestatory apparatus.
Resting cyst: nonmotile, dormant and resistant (e.g .. Toxicyst: see extrusome.
against desiccation, frost) stage in life cycle con- Trichocyst: see extrusome.
taining condensed cytoplasm; mouthless and sur- Undulating membrane: see paroral membrane.
rounded by membrane. Vestibulum (Fig. 6D): invagination of ventral or api-
Scopula (Pt) (Fig. 6E): organellar complex at posteri- cal cell surface, leading to cytostome-cytopharyn-
or pole, from which stalks and other attachment geal complex.
structures arise. Zooid (Pt) (Fig. 6D): body exclusive of stalk of indi-
Scutica (Se) (Fig. 6C): small field of often barren vidual members in a colony.
basal bodies, from which part of the oral ciliature
develops during divisional morphogenesis; con- Outline classification, order and family-Ievel
sists of only 2-3 basal bodies sometimes. diagnoses
Silverline system (Fig. 7E): system of strip-like or
reticular cortical structures revealed using silver There is no commonly accepted classification scheme
nitrate. for ciliates. The following, rather conservative system
• Fig. 6. Main features of a ciliate cell and charactcrs used for taxonomic identification (basal bodies are usually represented as dots). A: from
Foissner (1984); B: from Batisse (1994); C: original: O, E: from Nagasawa and Warren (1996).
280 Ciliophora
ventral
adoral
membranelles
"",,,,--- macronucleus
Top view of adoral zone of membranelles

of a strobilidiid ciliate
Ventral view of a \ ventral

strombidiid ciliate kinety
@ fro~itr~¡~ ,
¡f" , left
buccal cirrus
~f(
i' ~ /
§ adoral zone of
membranelles
marginal
cirri
Ventral view of an
wri~, l\1 ~~
frontoventral
paroral t ' ~!
~,
~ : urostylid hypotrich
endoral --:--- / , --- left
membrane
membrane
nght ~
f~~', , #~'
, :
'+--\
' I
marginal
clrn
postoral
marginal ~clrn
~: .J?
p~~~~:,~",Clrn v
•
~\::b' .,1---
,~Clrn
transverse
·_··..c.~:-~_····
__ ·,:'F:r••.····_··--1.·ii·- -¿). .._ ....4,.······ ..· , .....
Ventral view of an oxytrichid hypotrich
Schematic section through the cili~t~' cortex
4)
single-vannus double-eurystomus double-patella multiple complex
Types of dorsal silverline systems of fuplotes

Ciliophora 281
is an abridged version based on that given by Corliss Subc1ass Vestibulifera de Puytorac el al., 1974
(1979), centered on representatives from the South Order Trichostomatida Bütschli, 1889
Atlantic Ocean and with some minor additions by Suborder Trichostomatina Bütschli, 1889
Foissner and Foissner (1988) and Petz and Foissner Fami1y Plagiopylidae Schewiakoff, 1896
(1992). Taxa in bold characters are those treated in Family Coelosomididae Corliss, 1961
detail in this chapter. Family Trichospiridae Kahl, 1926
Family Trimyemidae Kahl, 1933
Kingdom Protista Haeckel, 1886 Family Marynidae Poche, 1913
Phylum Ciliophora Doflein, 1901 Family Ba1antidiidae Reichenow in Doflein
Class Kinetofragminophora de Puytorac el al., 1974 and Reichenow, 1929
Subc1ass Gymnostomata Bütschli, 1889 Family Pycnotrichidae Poche, 1913
Order Primociliatida Corliss, 1974 Family Isotrichidae Bütschli, 1889
Order Karyorelictida Corliss, 1974 Family Paraisotrichidae da Cunha, 1917
Fami1y Trachelocercidae Kent, 1881 Family Protocaviellidae Grain in Corliss,
Family Loxodidae Bütschli, 1889 1979
Family Geleiidae Kahl, 1933 Suborder Blepharocorythina Wolska, 1971
Family Protocruziidae Jankowski, 1980 Order Entodiniomorphida Reichenow in Doflein
Order Prostomatida Schewiakoff, 1896 and Reichenow, 1929
Suborder Prostomatina Schewiakoff, 1896 Order Colpodida de Puytorac el al., 1974
Suborder Prorodontina Corliss, 1974 Subc1ass Hypostomata Schewiakoff, 1896
Family Prorodontidae Kent, 1881 Order Synhymeniida de Puytorac el al., 1974
Family Colepidae Ehrenberg, 1838 Order Nassulida Jankowski, 1967
Order Gymnostomatida Bütschli, 1889 Order Cyrtophori'da Fauré-Fremiet in Corliss,
Order Spathidiida Foissner and Foissner, 1988 1956
Suborder Spathidiina Jankowski, 1980 Suborder Chlamydodontina Deroux, 1976
Family Spathidiidae Kahl in Doflein and Family Chilodonellidae Deroux, 1970
Reichenow, 1929 Family Chlamydodontidae Stein, 1859
Family Trachelophyllidae Kent, 1882 Family Lynchellidae Jankowski, 1968
Family Lacrymariidae Fromentel, 1876 Suborder Dysteriina Deroux, 1976
Family Homalozoonidae Jankowski, 1980 Family Plesiotrichopidae Deroux, 1976
Suborder Belonophryina Jankowski, 1980 Family Hartmannulidae Poche, 1913
Suborder Didiniina Jankowski, 1978 Family Dysteriidae Claparéde and Lach-
Family Didiniidae Poche, 1913 mann, 1858
Order Pleurostomatida Schewiakoff, 1896 Incerta sedis
Suborder Amphileptina Jankowski, 1967 Family Kryoprorodontidae Alekperov
Suborder Litonotina Foissner and Foissner, and Mamaeva, 1992
1988 Suborder Hypocomatina Deroux, 1976
Family Litonotidae Kent, 1882 Order Chonotrichida Wallengren, 1895
Family Loxophyllidae Foissner and Leipe, Order Rhynchodida Chatton and Lwoff, 1939
1995 Order Apostomatida Chatton and Lwoff, 1928
Order Pseudoholophryida Foissner and Foissner, Subclass Suctoria Claparéde and Lachmann,
1988 1858
Order Cyclotrichida Jankowski, 1980 Order Suctorida Claparéde and Lachmann,
Family Mesodiniidae Jankowski, 1980 1858
Order Archistomatida de Puytorac el al., 1974 Suborder Exogenina Collin, 1912
l
Fig. 7. Main features of a ciliate cell and characters used for taxonomic identification. A, C: original; B: from Petz and Foissner (1992); D:
from Borror (1 972b); E: from Curds (1975); F: from Lynn and Small (1990).
282 Ciliophora
Family Podophryidae Haeckel, 1866 Family Peniculistomatidae Fenchel, 1965

Family Parapodophryidae Jankowski, 1973 Family Thigmocomidae Kazubski, 1958
Family Urnulidae Fraipont, 1878 Family Conchophthiridae Kahl in Doflein
Family Ephelotidae Kent, 1882 and Reichenow, 1929
Family Spelaeophryidae Jankowski in Suborder Thigmotrichina Chatton and Lwoff,
Batisse, 1975 1922
Family Rhabdophryidae Jankowski, 1970 Order Astomatida Schewiakoff, 1896
Family Stylostomatidae Batisse, 1975 Subclass Peritricha Stein, 1859
Family Ophryodendridae Stein, 1867 Order Peritrichida Stein, 1859
Family Tachyblastonidae Grell, 1950 Suborder Sessilina Kahl, 1933
Family Thecacinetidae Matthes, 1956 Family Vorticellidae Ehrenberg, 1838
Family Phalacrocleptidae Kozloff, 1966 Family Astylozoidae Kahl, 1935
Suborder Endogenina Collin, 1912 Family Epistylididae Kahl, 1933
Family Acinetidae Stein, 1859 Family Operculariidae Fauré-Fremiet In
Family Dendrosomatidae Fraipont, 1878 Corliss, 1979
Family Trichophryidae Fraipont, 1878 Family Opisthonectidae Foissner, 1976
Farnily Endosphaeridae Jankowski in Corliss, Family Scyphididae Kahl, 1933
1979 Family Ophrydiidae Ehrenberg, 1838
Suborder Evaginogenina Jankowski in Corliss, Farnily Ellobiophryidae Chatton and Lwoff,
1979 1929
Class Oligohymenophora de Puytorac el al., 1974 Family Termitophryidae Lom in Corliss,
Subclass Hymenostomata Delage and Hérouard, . 1979
1896 Family Rovinjellidae Matthes, 1972
Order Hymenostomatida Delage and Hérouard, Family Vaginicolidae Fromentel, 1874
1896 Family Lagenophryidae Bütschli, 1889
Order Scuticociliatida Small, 1967 Suborder Mobilina Kahl, 1933
Suborder Philasterina Small, 1967 Class Polyhymenophora Jankowski, 1967
Family Philasteridae Kahl, 1931 Subclass Spirotricha Bütschli, 1889
Family Uronematidae Thompson, 1964 Order Heterotrichida Stein, 1859
Family Cohnilembidae Kahl, 1933 Suborder Heterotrichina Stein, 1859
Family Entorhipidiidae Madsen, 1931 Family Spirostomidae Stein, 1867
Family Entodiscidae Jankowski, 1973 Family Phacodinidae Corliss, 1979
Family Cryptochilidae Berger in Corliss, Family Metopidae Kahl, 1927
1979 Family Condylostomatidae Kahl in Doflein
Family Thyrophylacidae Berger in Corliss, and Reichenow, 1929
1961 Family Reichenowellidae Kahl, 1932
Family Loxocephalidae Jankowski, 1964 Family Climacostomatidae Repak, 1972
Family Cinetochilidae Perty, 1852 Family Stentoridae Carus, 1863
Family Urozonidae Groliere, 1975 Family Bursariidae Dujardin, 1840
Family Pseudocohnilembidae Evans and Farnily Chattonidiidae Villeneuve-Brachon,
Thompson, 1964 1940
Family Thigmophryidae Chatton and Lwoff, Family Peritromidae Stein, 1867
1926 Suborder Clevelandellina de Puytorac and
Suborder Pleuronematina Fauré-Fremiet in Grain,1976
Corliss, 1956 Suborder Armophorina Jankowski, 1964
Family Pleuronematidae Kent, 1881 Suborder Colinophorina Jankowski, 1967
Family Cyclidiidae Ehrenberg, 1838 Suborder Plagiotomina Albaret, 1974
Family Histiobalantidae de Puytorac and Suborder Licnophorina Corliss, 1957
Corliss in Corliss, 1979 Family Licnophoridae Bütschli, 1887
Ciliophora 283
Order Odontostomatida Sawaya, 1940 fragile, often elongate, highly thigmotactic. Oral area
Order Oligotrichida Bütschli, 1887 (see below) apical or ventral. Contractile vacuole probably often
Order Hypotrichida Stein, 1859 lacking. Many obligate interstitial forms, all except
Suborder Stichotrichina Fauré-Fremiet, 1961 one genus marine. Recently elevated to c1ass rank
Family Spirofilidae Gelei, 1929 (Small and Lynn, 1985).
Family Strongylidiidae Fauré-Fremiet, 1961 Family Trachelocercidae
Family Urostylidae Bütschli, 1889 Trachelocercida with simple circumoral ciliature
Family Psilotrichidae Bütschli, 1889 comprising a single row of dikinetids. Recently ele-
Family Kiitrichidae Nozawa, 1941 vated to order rank (Foissner and Dragesco, 1996b).
Family Keronidae Dujardin, 1840 Family Protocruziidae
Suborder Sporadotrichina Fauré-Fremiet, Body ovoid, small, noncontractile. With monokinetid
1961 or dikinetid field on right and dorsal side. 5-8 (usual-
Family Oxytrichidae Ehrenberg, 1838 Iy 6) adoral membranelles. Paroral membrane indis-
Family Aspidiscidae Ehrenberg, 1838 tinctly tripartite. Two barren basal bodies to rear and
Family Euplotidae Ehrenberg, 1838 right of oral region.
Family Gastrocirrhidae Fauré-Fremiet, 1961
Order Prostomatida
A phylogenetic approach using morphologic and mor- Cytostome at or near body surface, apical or subapi-
phogenetic characters resulted in a considerably dif- cal; cytopharyngeal apparatus of rhabdos type.
ferent system for aloricate oligotrichs (Petz and Circumoral infraciliature unspecialized, somatic ci-
Foissner, 1992). As this group dominates the ciliate liation often uniformly holotrichous. Toxicysts com-
fraction in the marine plankton, this improved scheme mono Macronucleus
. . polyploid. Body often large,
is given below. Apomorphies showed that strombi- many specles camlvorous.
diids are more c10sely related to tintinnids and strobi- Suborder Prorodontina
lidiids than to halteriids, which makes the recently Cytostome apical or subapical, round or oval, some-
proposed order Choreotrichida Small and Lynn, 1985 times in sh~lIow atrium. Unique brosse on dorsal sur-
superfluous. It also followed from that study that the face near anterior pole. Somatic toxicysts common.
hypotrichs should be elevated to c1ass rank as later Skeletal plates in one genus. Most species camivores
proposed by Tuffrau and Fleury (1994). or scavengers, a few algivorous.
Family Colepidae
Class Oligotrichea Bütschli, 1887 With characteristics of suborder. Body characteristi-
Subclass Halteriia Petz and Foissner, 1992 cally barrel-shaped, bearing uniquely calcified plates
Order Halteriida Petz and Foissner, 1992 (with smalJ lateral teeth) in longitudinal rows in cor-
Family Halteriidae Claparede and Lachmann, tex; prominent anterior and caudal thoms often
1858 present. Long caudal cilium common. Active scaven-
Subclass Oligotrichia Bütschli, 1887 gers.
Order Strombidiida Jankowski, 1980
Family Strombidiidae Fauré-Fremiet, 1970 Order Spathidiida
Order Oligotrichida Bütschli, 1887 Cytostome apical, round or slit-like, in suborder
Suborder Strobilidiina Jankowski, 1980 Didiniina on top of cone-like proboscis; rhabdos made
Family Strobilidiidae Kahl in Doflein and of 3 microtubular components: transverse ribbons
Reichenow, 1929 originating from the nonciliated basal bodies of the
Suborder Strombidinopsina SmalJ and Lynn, 1985 oral dikinetids, nematodesmal bundles originating
Family Strombidinopsidae Small and Lynn, exclusively from the same source, and bulge microtu-
1985 bules. Somatic ciliation uniform or limited to dense
Suborder Tintinnina Kofoid and Campbell, 1929 bands which, however, rest within longitudinally run-
ning kineties composed of nonciliated basal bodies;
Order Karyorelictida dorsal brush composed of 2 to many kineties.
Nuclear apparatus dual, but macronuclei diploid, non- Toxicysts localized, typically in or near oral area.
dividing, forming by division of micronuclei. Body Free-living.
284 Ciliophora
Suborder Spathidiina Order Cyclotrichida

Cytostome apical, round, oval or slit-like, in some Cytostomal-cytopharyngeal structures apical or anta-
genera covering the "ventral" body margin; somatic pical, in some genera strongly reduced and probably
ciliation usually uniformo functionless; oral basket (rhabdos) without nematodes-
Family Trachelophyllidae mata and bulge microtubules, but probably with trans-
Haptorids with bipolar meridional, rarely spiral1y verse ribbons; somatic ciliation limited to 2 cirri-hke
soma tic kineties, 1 circumoral kinety composed of girdles; dorsal brush absent; toxicysts present; often
basal body pairs from which nematodesmata origi- with incomplete cryptomonad symbionts. Might
nate. Brosse usual1y composed of several rows, belong to another subclass because of morphologic
meridional1y oriented. peculiarities (Foissner and Foissner, 1988).
Family Hornalozoonidae Family Mesodiniidae
Haptorids with ciliation complete on right side and Body ciha of 2 types (l cirrus-like) as 1 girdle that
reduced to 3-rowed brosse on left side. may begin near anterior end ofbody.
Suborder Didiniina
Cytostome apical on conspicuous oral cone; somatic Order Trichostornatida
ciliation bipartitioned in 1 or more ciliated and non- Rather similar to prostomatine gymnostome ciliates.
ciliated girdles (wreaths) which bear 10ngitudinal1y Vestibular ciliature non-complex, derived during fis-
arranged kineties. sion from terminal portions of sornatic kineties. Oral
Family Didiniidae aperture apical in free-living and antapical in endo-
Cytostome apical; cytopharynx eversible in some spe- zooic species. Some produce gelatinous lorica.
cies, equipped with multiple kinds of toxicysts; ci- Mostly endocornmensal in vertebrate hosts.
liation reduced to 1 or more circumferential ciliary Suborder Trichostornatina
wreaths on rotund body; "sensory bristles" (dorsal With characteristics of order. Often with dense soma-
brush) usual1y clearly recognizable. tic ciliation.
Family Plagiopylidae
Order Pleurostornatida Anteriorly located cytostome. Body dorsoventrally
Oral area flattened along ventral margin of lateral1y flattened. Ciliation uniformo Free-living in freshwater,
compressed body, surrounded by toxicysts. Oral bas- marine, salt marsh, and interstitial biotopes; endo-
ket (rhabdos) made of 3 microtubular components: commensals in digestive tracts of echinoids and hip-
transverse ribbons originating from oral dikinetids popotami.
and in suborder Litonotina also from somatic monoki-
netids, nematodesmal bundles originating exclusively Order Cyrtophorida
from oral dikinetids and bulge microtubules; somatic Body frequently dorsoventral1y flattened and flexible;
ciliature with distinct left-right differentiation, dorsal often with adhesive organelle at posterior end.
brush present. Free-living and parasitic on other ci- Somatic ciliature composed of monokinetids and
liates (mainly peritrichs), often large, long voracious restricted mainly to ventral side. Dorsal side naked or
carnivores, widely distributed in freshwater, marine with single ciliary row, viz. the dorsal brush. Several
and interstitial habitats. oral kineties anterior to oral basket (cyrtos), which is
Suborder Litonotina located in first half ofbody. Preoral suture skewed far
Cytostome surrounded by 1 right and 1 left perioral to left. Macronucleus heteromerous (except in one
kinety composed of dikinetids, right kinety accompa- suborder). Stomatogenesis telokinetal (parakinetal
nied by (oralized?) somatic monokinetids forming a according to some workers), but involving extensive
distinct 3rd perioral kinety whose transverse microtu- morphogenetic movements. Mostly marine, free-li-
bular ribbons contribute to oral basket (rhabdos); right ving, ectocommensals, a few parasitic on fish gills.
lateral ciliature with or without dorsolateral kineties, Suborder Chlarnydodontina
ciliary rows successively shortened along perioral and With characteristics of order senso stricto. Ventral
dorsolateral kineties. ciliature thigmotactic, no prominent specialized adhe-
Family Litonotidae sive organelle; body broad, flattened dorsoventrally.
Litonotina without right dorso lateral kineties. Family Chlarnydodontidae
Family Loxophyllidae Thigmotactic zone localized, body nearly ellipsoidal,
Litonotina with dorso lateral kineties. width >2/3 ofbody length.
Ciliophora 285
Suborder Dysteriina Suborder Exogenina

Body fonn less conservative and narrower than in Exogenous budding, most often monogemmic (but
Chlamydodontina. Oral basket often specialized. polygemmic in some species), with no conspicuous
With stylus. Widespread, mainly marine. invagination ofparental cortex. Small pennanent field
Family Hartmannulidae of nonciliferous basal bodies in vicinity of contractile
Adhesive organelle posterior, free from ciliary field vacuole. Swanner typically large or long, the fonner
on left; some uniqueness in infraciliary pattems. with complex ventral ciliature (derived from parental
Family Dysterüdae basal body field), but some of the latter practically
Somatic ciliature reduced, left ciliary field in 2 parts. devoid of cilia, vennifonn and incapable of swim-
Oral basket rods often with prominent teeth anterior- ming. Several species with actinophores, some with
Iy. Stylus may protrude conspicuously. Widely dis- prehensile as well as suctorian tentacles and bodies of
tributed, mainly marine, frequently symphorionts. many of diverse sizes (often large) and shapes. Mostly
marine, typically solitary and free-living or ectocom-
Incerta sedis mensals.
Family Kryoprorodontidae Family Thecacinetidae
Small to medium-sized phyllopharyngiids with ellip- Adult stalked, loricate, with tentacles grouped apical-
soid to spherical body, no distinct dorsoventral diffe- Iy; swanner ellipsoidal, flattened or vennifonn, with
rentiation. Cytostome apical, protruding slightly, with ventral ciliated bando Predominantly marine fonns.
conspicuous oral basket. Somatic ciliation fairly com- Suborder Endogenina
plete, composed of longitudinal and few spiralling Budding endogenous with I or more monogemmic or
kineties. Several short adoral membranelles. No sty- polygemmic swanners produced completely intemal-
lus. Gymno::oum [=Kryoprorodon Alekperov and Iy, becoming free-swirnming in brood pouch before
Mamaeva, 1992; Spiroprorodon Fenchel and Lee, emergence through birth pore. Small pennanent field
1972] was previously included in Enchelyidae or of nonciliferous basal bodies near contractile vacuole
Prorodontidae (Alekperov and Mamaeva, 1992; responsible for swanner ciliature. Swanners small,
Fenchel and Lee, 1972), which is inappropriate. with encircling band(s) of cilia. Adults usually small
Based on an ultrastructural study, Sniezek and Small (but ramified, and enonnous in some groups), often
(1993) proposed that it belongs in the subclass loricate, sometimes colonial. Tentacles frequently in
Phyllopharyngia de Puytorac el al., 1974. This is cor- bundles but not on papillae·("actinophores"). In fresh-
roborated by morphological and morphogenetical water and marine habitats, with ectosymbiotic forms
investigations by Petz el al. (l995a,b), who proposed cornmon and some endocommensals in wide range of
a c10se relationship to cyrtophorids. Species of this hosts.
genus are very likely widespread in the marine plank- Family Acinetidae
ton; some found in the Southem Ocean (Petz el al., Adults nearly always loricate and stalked, with stalk
1995a). persisting in some but not all endosymbiotic forms.
Tentacles in few bundles or reduced to single orga-
arder Suctorida nelle. Swanner small, elliptical. In variety of habitats
Ingestatory suctorial tentacles multiple (few to many, and on/in many hosts.
rarely none), with numerous microtubules, arranged
peripherally, and haptocysts distally at tips. Mature Order Scuticociliatida
form usually sedentary and without cilia. Often with Body often small to medium. Ciliation uniform,
noncontractile stalk. Swanners produced by some sometimes sparse, often including specific thigmotac-
mode ofbudding generally bear cilia but not tentacles tic area(s), somatic kineties composed of single and
or stalk. Without cytoproct. Conjugation frequently paired basal bodies, I or more caudal cilia; paroral
total with unequal conjugants. Body often of conspic- membrane sometimes dominating feature of oral cili-
uous size, attached to variety of substrata. ature, which may lie in extended but shallow oral ca-
Widespread, predominantly symphorionts on diverse vity. Stomatogenesis buccokinetal with complex mor-
marine and freshwater organisms, some endocom- phogenetic movements and unique involvement of
mensals in hosts ranging from ciliates to vertebrates. scutica. Mucocysts prominent, large, elongate, locat-
286 Ciliophora
ed eortieally; fused mitoehondria often eonspieuously marine and freshwater habitats, oeeasionally assoeiat-
presento Cysts widespread; abundant in marine habi- ed with invertebrates as eetoeommensals.
tats, free-living or in symbiotie assoeiation primarily Family Cyclidiidae
with invertebrates (espeeially molluses, eehinoids and Body ovoid to elongate ovoid, generally small to very
annelids), but some edaphie and freshwater speeies. small, with sparse eiliation and distinetive caudal ei-
Suborder Philasterina lium. Cytostome variable in position, oral eiliature less
Body generally small. Infraeiliature of paroral mem- prominent than in Pleuronematidae. Maeronucleus and
brane may show redueed "a" and no "e" segment, large mieronucleus often loeated in anterior 1/3 of
while "b" is always clearly present; seutiea posterior body. Widely distributed, freshwater, braekish,
to paroral membrane, often in anterior part of distinet marine, interstitial, edaphie, eoprozoie.
director meridiano Mueoeysts and mitoehondria very
prominent. Most eommonly in braekish or marine Order Peritrichida
habitats including sand; many free-living; some inqui- Body eharaeteristieally inverted bell- or goblet-
lines in sea urehins or eommensals in molluses, shaped or eonieal-eylindrieal. Conspieuous oral eilia-
eoelenterates, annelids, sipuneulids, and sea horses. ture winding eountercloekwise at apieal poleo With
Family Uronematidae seopula and prominent holdfast derivatives (usually
Body small, elongate ovoid, sparsely eiliated, with eontraetile stalk or eomplex adhesive dise) at antapi-
anterior pole eonspieuously naked (frontal plate) and cal poleo Somatie eiliature redueed to aboral eiliary
1 or more caudal eilia at posterior pole; oral eiliature wreath; eontraetile vacuo le empties into eiliated ves-
relatively ineonspieuous in long and shallow depres- tibulum (infundibulum). Stomatogenesis bueeokine-
sion, adoral membranelle 1 entirely noneiliferous; tal, with plane of fission parallel to main body axis.
seutiea bearing eilia. In marine or freshwater habitats, Dimorphism (with migratory swarmer stage), eolo-
oeeasionally edaphie. nies, lorieae or theeae, and eysts eommon in life eycle
Family Cohnilembidae of many speeies. Conjugation ("total") invariably
Body slender, finger-shaped, tapering to a point ante- involves fusion of a miero- with a maeroeonjugant.
riorly. Long caudal eilium. False "double-mem- Very widespread in aquatie habitats, speeies general-
brane", i.e. paroral membrane plus adjaeent (Ieft side) Iy free-living or oeeurring as symphorionts on diverse
row of soma tic eiliature (nth kinety) most eonspieu- hosts, but some eommensals or parasites on or in
ous in narrow oral depression. Marine habitats. other organisms.
Family Pseudocohnilembidae Suborder Sessilina
Body small, elongate-pyriform, with sparse somatie Adults sedentary or sessile, eommonly stalked (or
eiliature and single caudal eilium. Oral eavity with 2 with ineonspieuous adhesive dise), a few speeies see-
long membranes on right side. Marine, free-living or ondarily mobile; many produce arboroid eolonies;
seavengers, some inquilinie in eehinoids. some groups lorieate. Mueoeysts and pellieular pores
Suborder PIeuronematina universal. Adults generally filter-feeding baeterivores
Paroral membrane often prominent, infraeiliary base (swarmer stage mouthless). In widely ranging habitats
tripartite: short "a", elongate "b" and "e" segment as (freshwater, braekish, marine), with animate substrata
permanent seutiea. Cytostome equatorial or subequa- involving metazoa from many phyla (major exeep-
torial. Caudal eilia eonspieuous in many speeies. tion: eehinoderms); a few speeies endozoie.
Rarely with director meridiano Two types of mueo- Family Vorticellidae
eysts. Body typieally small to very small with oeea- With eontraetile stalk; colonial exeept in 2 genera
sional exeeptions. Widely distributed, free-living (Haploeaulus, Vortieella); zooids not independently
marine speeies; some eommensal in molluses; some eontraetile among colonial forms (exeept in
in freshwater or edaphie habitats, few eoprozoie. Carehesium); all speeies attaehed to inanimate objeets
Family Pleuronematidae or plants, rotifers, erustaeeans, turtles and others; in
With eharaeters of suborder. Paroral membrane is various freshwater and marine habitats.
dominating feature, sometimes as stiff velum, eurling
distinetly around subequatorial mouth; segment "b" Order Heterotrichida
rarely multifragmented. Usually with long, stiff cau- Body generally large to very large, sometimes highly
dal eilia. Some speeies possibly with triehoeysts. In eontraetile. Oral eiliature eonspieuous, adoral zone
Ciliophora 287
composed of many membranelles and often extending Two ovoid to spherical macronuclei. Generally
out of oral cavity onto ancl/or around anterior end of marine, also in salt marshes. In some characteristics
body. l or more undulating membranes on right. rather similar to hypotrichs.
Somatic ciliation frequently holotrichous, with regu- Suborder Licnophorina
lar, even dense rows of cilia. Postciliodesmata com- Body hourglass-shaped, oral disc prominent apically,
mon and prominent. Toxicysts and trichocysts absent. conspicuous fibre-rich and quite complex adhesive disc
Fission usually isotomic. Conjugation temporary, iso- posteriorly. Somatic ciliature reduced. Macronucleus
gamontic. Widely distributed, free-living and symbi- moniliform or fragmented. Stomatogenesis apokinetal.
otic. Ectocornmensals on variety of marine organisms (e.g.,
Suborder Heterotrichina algae, tunicates, coelenterates, annelids, molluscs, echi-
With characteristics of order. Somatic ciliature well noderms).
developed, adoral membranelles numerous and pro- Family Licnophoridae
minent. Body large, often contractile. Single conspicu- With characteristics of suborder.
ous contractile vacuole in posterior end of body.
Mucocysts abundant, pigment vesicles common in Order Strombidiida
many species. No loricae, cysts cornmon. Free-living, Circle of anterior adoral membranelles "open"; somat-
widely distributed. ic ciliature greatly reduced or even absent altogether.
Family Spirostomidae Family Strombidiidae
Large, often elongate, cylindrical forms, some highly Free-swirnming (rarely loricate), pelagic species; pel-
contractile, some pigmented (cortical granules). licle especially in posterior half ofbody made rigid by
Peristomial field long, narrow, sometimes with oral subpellicular platelet layer; perilemma present in
ciliature relatively inconspicuous. Somatic ciliation some species. With girdle of extrusomes ("trichites").
uniform, often dense and typically complete. Predominantly in marine habitats, also ectosymbiotic
Contractile vacuole posterior, frequently large, may on echinoids, several very common planktonic fresh-
have long collecting canal. Macronucleus compact, water forms.
ovoid to elongate-moniliform. Free-living in fresh-
water, marine, or edaphic habitats. Order Oligotrichida (=Choreotrichida)
Family Condylostomatidae Body ovoid to elongate, sometimes tailed, with thick-
Body usually large, contractile, heavily ciliated; ened pellicle. Somatic ciliature cornmonly reduced.
sometimes very elongate or nearly ellipsoidal. Adoral Adoral membranelles extensive and conspicuous, in
zone and paroral membrane prominent. Macronucleus two sections: one inside oral cavity (internal adoral
long and moniliform. Contractile vacuole often with membranelles), other (external adoral membranelles)
long collecting canal. In marine, freshwater and on body surface encircling anterior cell poleo
edaphic habitats. Stomatogenesis in pouch below surface. Mainly
Family Stentoridae marine pelagic organisms, free-swimming (even
Majestic in size and movement. Trumpet-shaped, when loricate); usually free-living, rarely associated
elongate, highly contractile, uniformly ciliated. with echinoids.
Adoral zone spirals nearly 3600 around flared anterior Suborder Strobilidiina
end. Stomatogenesis apokinetal. Often pigmented Peristomial field entirely apical, external adoral mem-
(cortical granules) andJor with symbiotic zoochlorel- branelles form "closed" circle in top view. Somatic
lae. Few species with mucilaginous lorica. ciliature inconspicuous, in longitudinal, spiralled or
Macronucleus commonly moniliform. Typically in transverse rows; originating by intrakinetal prolifera-
freshwater habitats. tion during morphogenesis. In freshwater and marine
Family Peritromidae habitats, rarely inquilines of echinoids.
Body ellipsoidal, dorsoventrally flattened. With regu- Family Strobilidiidae
lar ciliation on ventral surface only. Lateral edges and With characters of suborder.
dorsal surface bear short stiff spines, associated in lat-
ter location with wart-like prominences. Adoral mem- Order Hypotrichida
branelles conspicuous at apical end of body, curving Dorsoventrally flattened, oval or elongate, medium-
down left side to mouth located in anterior 1/3 of cell. sized forms, occasionally tailed. Prominent zone of
288 Ciliophora
adoral membranel1es on left-anterior portion of ven- surface. Dorsal surface often distinctly ribbed, ridged
tral surface. Somatic ciliature represented by rows or or humped. Widely distributed in variety of habitats
groups of cirri on ventral surface and rows of basal including echinoid gut and ascidian branchial cavity.
body pairs on dorsal side; perilemma present in some Family Eup10tidae
species. Stomatogenesis general1y apokinetal, but Marginal cirri absent or greatly reduced in number;
apparently parakinetal in more primitive species. transverse and frontoventral cirri often tremendously
Macronuclear reorganization bands common. developed and very conspicuous. Adoral zone of
Mucocysts plentiful. Widely distributed free-living membranelles also praminent, usually extending over
forms in many and diverse habitats, with few ecto- more than half of body length. Intercirral tracts of
commensal on various invertebrates and rarely inqui- microtubules, kinetodesmata reported. Many species,
linic in echinoids. Data on morphogenesis often need- widely distributed, predominantly marine.
ed to arrange families and genera into proper
taxonomic groupings. Thus, classification is in many
instances provisional. Identification of the species
Suborder Stichotrichina
Body often elongate, sometimes very drawn out pos- There are only a few reports on planktonic ciliates of
teriorly. Ventral cirri general1y smal1 and quite incon- the South Atlantic (Table 2). As these are often out-
spicuous (occasional1y as few as 2-3 cilia per cirros) dated or give only species lists, many identifications
and typically arranged in 3-12 longitudinal (some- have to be regarded as doubtful.
times spiralled) rows. Marginal cirri common, trans-
verse cirri sometimes absent. Stomatogenesis parakin- Although the diagnoses and descriptions that follow are
etal in primitive forms. Several species produce centered on planktonic species, they also include some
loricae, a few ofthese'forming colonies. Free-living in benthic or interstitial fonns (e.g., Tracheloraphis phoe-
diverse habitats, also ectocommensal. nicopterus), as well as a few normally sessile ones (e.g.,
Family Urostylidae Stentor, suctorians). Both types can occur sporadically
Somatic ciliature with 1 or more rows of right and left in plankton samples, especially in nearshore areas, and
marginal cirri ventral1y; frontal ciliature with variously sessile forms are often found on floating debris and
arranged frontal, midventral, frontoterminal, and trans- attached to larger plankters, such as copepods.
verse cirri (latter sometimes absent). Frontal and mid-
ventral cirri differentiate during morphogenesis fram As a10ricate oligotrichs often make up the main por-
longitudinal field of more than 5 oblique cirral streaks. tion ofplanktonic ciliates, a key to common and wel1-
Suborder Sporadotrichina studied marine and freshwater genera is pravided,
Body sometimes elongate, even tailed, but often oval Many of the characters used can be recognized by
to elliptical in outline. Ventral cirri typical1y heavy, careful examination of living specimens with diffe-
conspicuous, arranged in specific localized groups. rential interference contrast but this requires consider-
Marginal cirri often absent or reduced. Stomatogenesis able experience. Thus, additional utilization of silver
apokinetal. In widely diverse habitats, few symbiotic impregnation is preferred. Keys for other groups or
(ectocommensals, inquilines). species are inapprapriate now because many descrip-
Family Oxytrichidae tions ofplanktonic ciliates are superficial according to
Hypotrichs with flexible or rigid body and usual1y 18 modero standards, so correct identification is impos-
frontoventral-transverse (FVT) cirri clustered to 6 dis- sible. But short descriptions ofthe species recorded in
tinct graups. FVT cirri originate fram 6 longitudinal the South Atlantic are given based on life observa-
primordia segregating 1, 3, 3, 3,4,4 cirri. Six or less tions and sil ver impregnation as far as available. If in
parental FVT cirri participate in primordia formation. doubt over an identification, consult the special liter-
At least 1 fragmenting dorsal kinety. ature since many new or hitherto unrecorded species
Family Aspidiscidae are likely to be found in the pelagic realm.
Generally smal1 forms with flattened oval to raunded
body. No marginal cirri but strikingly praminent ven- Species names are fol1owed by common synonyms [in
tral group(s). Reduced number of adoral membra- square brackets] when available but subsequent incor-
nel1es located anteriorly and posteriorly on ventral rect spel1ings are not listed. Movement of individuals
Ciliophora 289
is described, although it alone is not usually sufficient ed in scuticociliatids but recently transferred to kar-
for species identification. Size range (minimum, ma- yorelictids in separate family and order (Small and
ximum) is generally from life and thus often an Lynn, 1985). Poorly known species. Ref.:
approximation. However, it is valuable because dif- Hamburger and Buddenbrock (1911), Kahl (1932).
ferent kinds of fixation lead to variable degrees of
shrinkage and distortion (see above). For each of the Family Colepidae
species treated one or more references ("Ref.") pro- • Tiarilla fusus (Claparede and Lachmann, 1859)
viding a description and illustrations are given. Bergh, 1882 (Fig. 8.3) [=ColepsjitslIs]. Body fusi-
form, tapering posteriorly, sometimes with short
Family Trachelocercidae spines, anteriorly toothed. Cytostome apical,
Trache/oraphis phoellicopterus (Cohn, 1866) invaginated. Five transverse rows of rectangular,
Dragesco, 1960 (Fig. 8.1) [=Traclzelocerca pllOeni- calcareous plates with lateral teeth in cortex. One
coplerus]. Body filiform when fully extended, ante- macronucleus, l micronucleus. About 15 longitu-
rior and posterior portions tapering, glabrous strip e dinal somatic kineties between plates; l elongate
wide, head claviform, oral bulge inconspicuous, caudal cilium. Dorsal brush composed of 3 short
brosse cleft indistinct; highly flexible, contractile, dikinetidal rows slightly oblique to main body
flattened. Contracted specimens banana-shaped, axis. Swims rapidly in sin uso id trajectory. Size in
glabrous stripe distinctly protruding on convex vivo 85-135 x 22-45 /lID. May comprise several
side. Nuclear capsule single, ellipsoidal, 15-30 /lm species (Kahl, 1930). Mass occurrences may cause
long, composed of 4-18, often 12, macronuclei and red tides (Dale and Dahl, 1987b). Ref.: Hamburger
2-8, usually 6, micronuclei. No contractile vacuole. and Buddenbrock (l911), Fauré-Fremiet (1924),
Cortical granules minute, yellowish, between ci- Kahl (1930), Small and Lynn (1985).
liary rows and in glabrous stripe. Soma tic kineties
19-27 in number, composed of dikinetids; nonci- Family Trachelophyllidae
liated glabrous stripe enclosed by bristle kinety, • Chaellea teres (Dujardin, 1841) Kent, 1881 (Fig.
which consists of quite irregularly arranged diki- 8.4). Body elongate cylindrical to fusiform, flex-
netids with rather stiff cilia. Circumoral kinety of ible, slightly contractile; ~ral region set off resem-
dikinetids interrupted at brosse cleft; 2-4, cornmon- bling a "head", often slightly bent but usually
Iy 3 short oblique brosse kineties. Movement glid- indistinct after impregnation. Oral bulge round,
ing and winding. Size in vivo 1000-1500 x 30-50 protruding slightly. Extrusomes 6-10 /lm long, in
flm, fully contracted specimens about 600 /lID. vivo rod-like and fine, in oral bulge and scattered
Usually a mesopsarnmal species which has several in body. Over 100 macronuclear nodules.
synonyms and homonyms (Raikov, 1962). Ref.: Contractile vacuole in posterior end. Cortical gra-
Kahl (1927), Raikov (1962), Raikov and Kovaleva nules colourless, in interkinetal rows. Specimens
(1994), Foissner and Dragesco (1996b). often yellowish-brown, very likely because offood
vacuole contents. Somatic kineties 10-14 in num-
Family Protocruziidae ber, longitudinal, in contracted specimens slightly
Protocruzia pigerrima (Cohn, 1866) Faria el a/., spiralling, on "head" distinctly spiralled. densely
1922 (Fig. 8.2) [=Colpoda pigerrima, Bleplza- ciliated. Dorsal brush typical for genus, composed
rosloma pigerrima, Cryptochilllm fusiforme]. Body of 4 rows of dikinetids. Circumoral kinety indis-
fusiform, tapering anteriorly and posteriorly, later- tinct, very likely composed of 1 dikinetid at ante-
ally compressed, i.e. left side vaulted, right side flat; rior end of each somatic row. Movement slowly
slightly contractile and flexible. 1(2?) macronuclei. gliding, rotating about main body axis when swim-
Contractile vacuole subterminal. About 4 longitudi- ming in straight line. Size in vivo 120-300 x 20-30
nal somatic kineties on right side, bristle-like cilia /lm. Ref.: Kahl (1930), Lipscomb and Riordan
on left side. Adoral zone in anterior 1/3-1/4 ofbody, (1990), Petz el al. (I995a).
composed of few membranelles; paroral membrane
on right of oral area. Movement crawling. Size in Family Homalozoonidae
vivo 30-60 x 10-20 /lID. Frequently confused with • Bllche/yodoll /aevis (Quennerstedt, 1867)
P. adlzaerens (Kahl, 1932). Genus formerly includ- Dragesco, 1960 (Fig. 8.5) [=Lag)'lllls laevis, L.
290 Ciliophora
sulcalus, L. crassicollis, L. ornalus, Enchelyodon 120, usually 50-80 x 30-80 Jlm. Species apparent-
slriallls, Trae/lelocerca sulcata, T. laevis]. Body Iy highly variable, comprising several rather simi-
cylindrical to bottle-shaped, neck not distinctly lar varieties or species (Dingfelder, 1962;
elongate; highly contractile, pellicle usually Foissner, 1979). Ingested phytoflagellates were
appears annulated (from slight contraction?). previously (e.g., Schewiakoff, 1896; Fauré-
Macronucleus elongate ellipsoidal. Contractile Fremiet, 1924) possibly misidentified as zoochlo-
vacuole with collecting canals in posterior end. rellae (Foissner el al., 1995). Ref.: Foissner
About 48 soma tic kineties. Conspicuous extru- (1979), Dragesco and Dragesco-Keméis (1986),
somes (ca. 30 ¡.lm long) and elongate cilia sur- Song and Wilbert (1989), Foissner el al. (1995).
round apical oral aperture. Size in vivo 150-350 x
70-100 Jlm. Encysts readily on slide; resting cysts Family Litonotidae
bott1e-shaped. Usually a mesopsammal species • LitOIlOtllS CygllllS (Müller, 1773) Foissner el al.,
(Dragesco, 1960) but also found on copepods 1995 (Fig. 8.8). Body slender, lanceolate, flexible,
(Hamburger and Buddenbrock, 1911). Redescrip- highly contractile; posterior portion narrowly
tion necessary. Ref.: Hamburger and Buddenbrock rounded to tail-like; neck longer than body, wid-
(1911), Kahl (1930), Dragesco (1960). ened anteriorly and curved dorsally. Anterior and
posterior body portion distinctly flattened, middle
Family Didiniidae section of left side considerably vaulted (especial-
• Didillilllll gargalltlla Meunier, 1910 (Fig. 8.6). Iy in well-fed and contracted specimens).
Body barrel-shaped to globular, with anterior oral Contracted specimens broadly lanceolate, with
cone. Macronucleus single, sausage-shaped. several longitudinal ribs on left side. Contractile
Terminal contractile vacuole very likely. vacuole subterminal; 6-8 Jlm long extrusomes rod-
Extrusomes (cyrtocysts) rod-1ike, scattered in like in top view but distinctly curved in lateral
body, in protargol slide.s often curved; other extru- view, ends tapering; found mainly along oral slit.
somes (developmental stages, pexicysts?) fusi- Ciliary rows 6-13 in number, on right side, gra-
form, 4-6 ¡.lm long. Toxicysts 40-50 f.lID long, dually abutting right perioral kinety anteriorly; se-
between 25-50 Jlm long nematodesmata. Basal veral rows of short cilia on left side. Dorsal brush
body rows 50-75 in number extending from poste- composed of 10-15 dikinetids with clavate cilia.
rior pole almost to oral cone, usually nonciliferous Oral slit extends over 113-1/2 of body along ven-
except 2 ciliary wreaths (1 at anterior end of tral side of neck. Paralleled by three perioral kine-
somatic kineties, I in rear 1/4 of body, about 15 ties, i.e. rows 2 and 3 with normally long cilia on
cilia wide) and dorsal brush (posterior to each ci- right and row I with short cilia on left side; kine-
liary wreath, composed of2-3 short rows ofpaired ties I and 2 composed of dikinetids and row 3 of
basal bodies). Swims moderately fasto Size in vivo monokinetids (Fig. 8.8 B, C). Size in vivo 200-500
70-200 x 50-120 Jlm. Ref.: Meunier(1910), Petz el Jlm (extended), 100-200 f.lID(contracted). Because
al. (1995a). of marked contractility, very variable in shape and
• MOIlOdillilllll balbiallii Fabre-Domergue, 1888 thus easily confused with other species. Litonolus
(Fig. 8.7) [=Didinillm balbianii]. Body oviform, cygnus described by Borror (1972a) belongs to a
with prominent oral cone. Macronucleus horse- different species because it has 4-7 macronuclear
shoe-shaped to reniform. Contractile vacuole in nodules (Foissner el al., 1995). Ref.: Dragesco and
posterior end. Resting extrusomes in oral cone and Dragesco-Keméis (1986), Foissner el al. (1995).
body 5-10 Jlm long; many granular mucocysts in • LitOIlOtllSfasciola (Müller, 1773) Wrzesniowski,
cortex. 54-80( I00) longitudina1 basal body rows, 1870 (Fig. 8.9). Body slightly contractile, posterior
only anterior portions (about 9-17 basal bodies end elongated tail-like, tapering but not pointed.
long) ciliferous, forming single ciliary wreath. Laterally flattened 3-4: l. Without dorsal ribs or
Dorsal brush posterior to ciliary wreath, composed furrows. Extrusomes rod-shaped, about 3-8 Jlm
of 4-5 short rows ofpaired basal bodies with bris- long in protargol slides. Six-eight somatic kineties
tle-Iike cilia. Circumoral kinety immediately ante- on right side (including 2 perioral kineties), 5 on
rior to ciliary band (Foissner el al., 1995). Swims left (including I perioral kinety and dorsal brush).
to and fro, usually very rapid1y. Size in vivo 50- Size in vivo 80-140 Jlm. Foissner el al. (1995) pro-
Ciliophora 291
pose that L. fasGiola actually comprises only equatorial furrow separating conical anterior por-
stressed, more or less contracted specimens of L. tion from globular trunk; colour blue-green.
cygnus and is thus a synonym of the latter. Ref.: Cytostome apical, surrounded by up to 12 retrac-
Kahl (1931), Song and Wilbert (1989). tile, forked processes. 1 to 2 (uncertain) spherical
macronuclear nodules. Possibly 2 contractile vac-
Family Loxophyllidae uoles pre-equatorial in conical portion; posterior
• Kelltrop"ylllllll setigerulll (Quennerstedt, 1867) position very likely mistaken. Crescent-shaped
Petz el al., 1995a (Fig. 8.10) [=Loxophyllum seti- vacuole of unknown function often in posterior
gerllm]. Body broadly lanceolate, variable (depen- end. Without extrusomes (7). 26-49 short, bipar-
ding on nutritional state), broadly rounded pos- tite, longitudinal ciliary rows in area of pre-equa-
teriorly, highly flexible; laterally compressed. torial furrow; anterior portions form 3 pre-equato-
Spines along entire periphery of cell. Macronuclear rial ciliary belts composed of anteriorly, laterally
nodules 4 in number (sometimes 3), 2-3 micro- and posteriorly directed bundles of 14-20 ¡.lmlong
nuclei. 3-7 contractile vacuoles in row along dorsal and stiff cilia ("cirri"), bundles only distally bifur-
margino Extrusomes more or less regularly distri- cate, with 2-3 barbs; posterior portions form equa-
buted along ventral and dorsal margins. No extru- torial belt difficult to recognize because cilia
some warts. About 18-22 somatic kineties on right c10sely adjoin pellicle. Dorsal brush absent (7).
side. Row of papillae along anterior right ventro- Moves in very rapid backward jumps, rests with
lateral margino Size in vivo 70-350 x 55-105 ¡.1m. pre-equatorial cilia rigidly spread. Often attaches
Usually a mesopsammal species. Ref.: Kahl to substrate with oral processes. Size in vivo 15-35
(1931), Dragesco (1960, 1965), Dragesco and x 10-25 ¡.tm. Ref.: Kahl (1930), Borror (1972a),
Dragesco-Keméis (1986). Tamar (1992), Foissner et al. (1995).
• Myr;ollecta rllbra (Lohmann, 1908) Jankowski,
Family Mesodiniidae 1976 (Fig. 8.13) [=Halteria rubra, Mesodinillm
• Mesod;II;1I11l acarus Stein, 1867 (Fig. 8.11) rllbrllm, Cyclolrichillm mellnieri]. Body usually
[=Mesodinillm phialinllm]. Body pyriform, pre- broadly obovoid to barrel-shaped, appearing to
equatorial furrow separating conical anterior por- consist of 2 hemispherical portions; pre-equatorial
tion from globular trunk. 1, possibly 2, macronu- furrow separates wider anterior from narrower and
c1ear nodules in mid-body. Two contractile sometimes slightly tapering posterior portion; very
vacuo les pre-equatorial in conical body portion; fragile. Usually reddish-brown from cryptophy-
position in rear end very likely erroneous. Without cean endosymbionts, sometimes green or colour-
extrusomes (7). Longitudinal ciliary rows 16-25 in less. Macronuclei 1-2 in number, 1 micronucleus.
number, short, bipartite, in area of pre-equatorial Contractile vacuole lacking (7). Around 40 kinety
furrow; anterior portions form 2 pre-equatorial ci- belt rows in area of pre-equatorial furrow com-
liary belts composed of anteriorly and laterally posed 1) of basal body patches at anterior ends,
directed bundles of about 8-12 ¡.lm long and stiff from which anteriorly, laterally and posteriorly
cilia ("cirri"), bundles bifurcate in distal half, directed bundles ~f long, stiff cilia ("cirri") pro-
without barbs; posterior portions form equatorial ject; and 2) of longitudinal rows of paired basal
belt (difficult to recognize because cilia c10sely bodies in posterior portions. Without cytostome;
adjoin pellicle). Dorsal brush absent (7). however, 6-8 bifurcated oral processes (perhaps
Cytostome apical, surrounded by up to 8 retractile, extrusomes) and several basal bodies surround
bifurcated processes ("tentacles"). Moves in very anterior pole in some populations. Movement
rapid backward jumps, rests with pre-equatorial jumping. Size in vivo 10-70 x 20-37 ¡.lm. Very
cilia rigidly spread. Size in vivo 12-20 x 8-15 ¡.1m. likely sometimes confused with other similar-
Foissner el al. (1995) propose that M. acarus, sized reddish species, which already suggests the
which is actually a freshwater species, is possibly variety of shapes and sizes (Kahl, 1930;
a synonym of M. plllex. Ref.: Tamar (1987), Lindholm, 1985; Crawford, 1989; Petz el al.,
Foissner el al. (1995). 1995a). The systematic position of M. rubra is
• Mesod;II;1I11lplllex (Claparede and Lachmann, uncertain: Kahl (1930) synonymized it with
1859) Stein, 1867 (Fig. 8.12). Body pyriform, pre- Mesodinium pulex (which was followed by, e.g.,
292 Ciliophora
Revelante and Gilmartin, 1983), whereas Four to five contractile vacuoles. Patch of orange
Jankowski (1976) established the new but very to yellow-green granules (pigment spot) on ante-
insufficiently diagnosed genus Myrionecta for it. rior left. Right ciliary field composed of 11-18,
Mass occurrences of this ciliate cause red tides. left of 9-20 kineties; 4 postoral ciliary rows. Oral
Ref.: Hamburger and Buddenbrock (1911). Kahl basket composed of 8-13 nematodesmal rods with
(1930), Taylor et al. (1971), Lindholm (1981, a tooth, 1-2 nonciliated dikinetids each associated
1985), Grain et al. (1982). with a rod. One preoral and 2 circumoral kineties
Cyclotrichiul1l gigas Fauré-Fremiet, 19?4 (Fig. (perioral kineties). Thigmotactic. Size 30-150 x
8.14). Body hemispherical to lenticular, anterior 15-70 ~m. Usually a benthic species. Ref.: Kahl
portion nonciliated. Single equatorial girdle of (1931), Deroux (1974), Dragesco and Dragesco-
about 35 ~m-Iong cilia; longitudinal rows of short Keméis (1986), Bardele (1993).
cilia posterior to this. Macronucleus cylindrical,
almost completely encircling cell. One micronu- Family Hartmannulidae
cleus. Movement rapid, rotating about main body • Aegyrialla oliva (Claparede and Lachmann, 1859)
axis. Size about 160 x 250 ~m. Record by Leakey Deroux, 1974 (Fig. 8.17) [=Aegyria oliva, Dysteria
et al. (1994b ) from the Southern Ocean uncertain. oliva, Hartmannula oliva]' Body roughly ellipsoid-
Ref.: Fauré-Fremiet (1924). al, anteriorly obliquely truncate; ventral plate dis-
tinctly smaller than dorsal plate. Dorsoventrally
Family Plagiopylidae slightly flattened. Stylus usually inconspicuous,
• Plagiopyla "asuta Stein, 1860 (Fig. 8.15). Body distinctly subterminal on right ventral side. Two
reniforn1, with distinct nose-like projection at contractile vacuo les. Oral basket wide, composed
anterior margin of oral cleft; with endosymbiotic of many nematodesmal rods which lack a promi-
methanobacteria. Laterally slightly to distinctly nent tooth. Cytoplasm often dark red (from food?).
flattened. Mouth conspicuous in deep, transverse Pigment spot on anterior left of body. Right ciliary
subapical oral cleft. Striated band ("transverse rib- field rather wide. Movement slowly gliding. Size
bon") inversely J-shaped on right side. Contractile in vivo 80-140 x 40- 70 ~m. Third contractile va-
vacuole in rear end. Resting extrusomes rod- cuole found by de Morgan (1925) is very likely an
shaped, about 7 ~ long, forming distinct layer artifact. Ref.: Hamburger and Buddenbrock
underneath pellicle. Ciliary rows 55-90 in number; (1911), Kahl (1931), Deroux and Dragesco (1968),
ventral kineties extending from oral cavity back- Deroux (1974).
ward; dorsal rows turning anteriorly, abutting oral
cavity on anterior ventral and lateral sides. Oral Family Dysteriidae
ciliature formed by densely ciliated anterior por- • Dysteria brasiliellsis Faria el al., 1922 (Fig. 8.18).
tions of somatic kineties. Size in vivo 80-180 x 35- Body elongate triangular, posteriorly tapering and
80 ~m. Indicator of microaerobic or anaerobic with distinct spur-like protrusion dorsally.
conditions. Foissner et al. (1995) considered Laterally distinctly flattened, left and right plates
records from marine and brackish habitats very equally sized. Stylus on posterior ventral side.
likely misidentifications and included the genus in Two contractile vacuoles, ventral in anterior and
Gymnostomatea. Ref.: de Puytorac et al. (1985), posterior 1/3 of cell. Five right field kineties. Left
Dragesco and Dragesco-Kernéis (1986), Sola et equatorial field composed of 7-8 short rows of
al. (1988), Foissner et al. (1995). basal bodies, 1 external right equatorial kinety
extending over about 1/3 of body length. Preoral
Family Chlamydodontidae kinety before left frontal field kineties. Movement
• Chlal1lydodoll I1lllel1losy"e Ehrenberg, 1835 (Fig. slowly crawling, often resting. Size in vivo 100-
8.16). Outline broadly elliptical to slightly renif- 150 x 30-60 ~m. Ref.: Song and Packroff (1997).
orm, anteriorly and posteriorly broadly rounded; • Dysteria cOl1lpressa (Gourret and Roeser, 1888)
dorsoventrally flattened. "Cross-striped band" Hamburger and Buddenbrock, 1911 (Fig. 8.19).
("railroad track") along entire periphery of cell is Body broadly rectangular, spur-like protrusion on
a hollow tube kept open by numerous C-shaped posterior dorsal side may be shifted slightly ven-
structures. One macronucleus, 1 micronucleus. trally. Laterally distinctly flattened, left plate
Ciliophora 293
slightly larger than right. Stylus rather smal!. Two GYlIlllOZOUlIl were found further south in Antarctic
ventral contractile vacuoles. Oral basket narrow. sea ice (Fenchel and Lee, 1972; Corliss and
Movement rapid. Size about 35-45 x 28 ¡..tm. Snyder, 1986; Petz et al., 1995a). However, the
Reinvestigation necessary. Ref.: Faria el al. arrangement of a few somatic kineties differs. Not
(1922), Kahl (1931). having the advantage of silver impregnation,
• Dysteria mOllostyla (Ehrenberg, 1838) Kahl, 1931 Lohmann (1920) might have missed the actual
(Fig. 8.20). Body elongate, posteriorly slightly nar- course of these rows, which bend anteriorly and
rowed. Laterally distinctly flattened, left and right almost encircle the mouth. Therefore, H. pe/agica
plates of equal size, gaping only anteriorly. Stylus very likely belongs to GYlIlnozoulIl Meunier, 1910
on posterior ventral side. Two contractile vacuo les, and is thus tentatively placed in the family
ventral in anterior and posterior 1/3 of body. Five Kryoprorodontidae Alekperov and Mamaeva,
right field kineties. Left equatorial field composed 1992. Ref.: Lohmann (1920).
of about 6-8 short rows of basal bodies, 1 very
short external right equatorial kinety. Preoral kine- Family Thecacinetidae
ty shifted slightly backward relative to left frontal Coryllopllrya campallllla (Schroder, 1906) Kahl,
kinety field, row of 3-4 basal bodies between cir- 1934 (Fig. 8.24) (=Tokophrya campanula]. Body
cumoral kineties and oral basket. Movement slow- thick, disc-like. Apical tentacles thick, distally
Iy crawling; thigmotactic. Size in vivo about 145 x slightly knob-Iike, encircle anterior area in continu-
65 /lm. Ref.: Deroux (1976), Dragesco and ous multi-rowed wreath. Macronucleus horseshoe-
Dragesco-Kernéis (1986), Petz et al. (1995a). shaped, ramified. Lorica cup-shaped, ca. 130 x 140
• Trocllilia sigmoides Dujardin, 1841 (Fig. 8.21). Jlffi, or about 50 x 50 Jlffi (Biernacka, 1963). Stalk
Body slender, ellipsoidal, anteriorly slightly 14-20 /lm thick and short, >200 /lrn (Schroder,
curved leftward, slightly notched on left posterior 1906), or 60 Jlffi (Biernacka, 1963). Polygemmic,
by dorsal grooves. Dorsally vaulted, with 5-6 dis- forming 1-3 apparently internal buds. Body 150-
tinct crests. Stylus long, slightly flexible. Two 200 ¡..tmacross. Swarmer slenderly ellipsoidal, ci-
contractile vacuo les, near base of stylus and in liated end ofbody tapering; size 90-100 by about 30
anterior 1/3 of body on right. Four somatic kine- Jlffi. Epibiotic on copepods. Redescription needed.
ties on right margin (right ciliary field), extend There are controversies concerning the reproduc-
from stylus anteriorly to frontal left margino Left tion of this genus, viz. exogenous or endogenous
ciliary field reduced, composed of 2 short rowS. budding (e.g., Corliss, 1979; Schroder, 1906). Its
Dorsal peripheral ciliary row very short. Two per- systematic position is thus uncertain. Ref.: Schroder
ioral kineties anterior of oral basket. Oral basket (1906), Kahl (1934), Biernacka (1963).
composed of prominent, anteriorly thickened Coryllopllrya interrupta (Schroder, 1906) Kahl,
nematodesmal rods. Size in vivo 20-35 x 12-20 1934 (Fig. 8.25) (=Tokophlya interrupta]. Body
I.un.Ref.: Kahl (1931), Deroux (1976). disc-like. Apical tentacles thick, encircle anterior
area in several rows which are separated into 2
Family Kryoprorodontidae groups each of about 10-40 distally slightly thick-
Holopllrya pelagica Lohmann, 1920 (Fig. 8.23). ened tentacles. Macronucleus variable, horseshoe-,
Body oboval, slightly asymmetrical, colourless; C- or X-shaped, sometimes ramified. One (to se-
cross section circular. Oral opening protruding veral?) micronuclei. Lorica funnel-like. Stalk 20-30
slightly apically. Oral basket wide and long Jlffi thick, short to about 3 times lorica length.
(extending to posterior end), composed of thick Polygemrnic forming 1-4 apparently internal buds.
nematodesmal rods. Macronucleus ellipsoidal Body diameter 100-140 ¡..tm.Swarmer bean-shaped,
(very likely confused with large food vacuole in entire ventral side and I end ciliated; size about 80
original description). About 25 longitudinal x 40 /lm. Epibiotic on copepods. Redescription
soma tic kineties. Length 20-90 /lm. This species is needed. Ref.: Schroder (1906), Kahl (1934).
rather unusual for H%pluya. Instead, body shape
and long oral basket distinctly resemble those of Family Acinetidae
GY11l1l0:::0UlIlspp. (Fig. 8.22). This is further cor- • Acilleta tllberosa (Pailas, 1766) Ehrenberg, 1833
roborated by the fact that several species of .(Fig. 8.26) (numerous synonyrns, for review see
294 Ciliophora
Curds (1985) and Matthes el al. (1988»)". Loriea narrow and shallow, extends along anterior body
highly variable, slenderly to broadly triangular, half. Contractile vacuole in posterior end.
anteriorly transvco:rsely truncate, usually occupied Extrusomes elliptical, in humps between cilia and
by entire cell; distinctly flattened; transversely kineties. Somatic kineties 8-10 in number, kinety
striated in some populations (different species?). left of oral cavity extending very close to adoral
Two bundles of tentacles inserted on hemispheri- membranelle 1, which thus appears to consist of 2
cal papillae on apical ends of cell projecting ciliary rows; 1 elongate caudal cilium. Adoral
through openings on opposite sides of lorica, each membranelle 1 very long, i.e. extends from anteri-
bundle consisting of 4-30 (often 10-20) tentacles, or end almost to cytostome, composed of single
up to 80 IJllllong, with globular ends. No articula- row of cilia; adoral membranelle 2 short, com-
tion between lorica and stalk, which may be very posed of few dikinetids; adoral membranelle 3
short to 5 times lorica length (2-5 ¡lm wide). small, trapezoidal. Undulating membrane short,
Macronucleus globular to ellipsoid, several micro- C-shaped, on posterior right of oral cavity. Scutica
nuclei. One contractile vacuole in anterior body comprises about 3-6 basal bodies. Movement
half. Lorica 23-200, usually 40-100 x 25-80 !lm. jerky. Size in vivo 60-130 x 10-18 ¡lm. Usually a
Swamler ellipsoidal, in vivo 35 x 25-30 !lm, with benthic species. Ref.: Kahl (1931), Evans and
7 long and 1 short oblique ciliary rows which are Thompson (1964), Thompson (1968), Didier and
ventrally interrupted. Ref.: Kahl (1934), Curds Detcheva (1974).
(1985), Matthes el al. (1988), Foissner el al.
(1995). Family Pseudocohnilembidae
• Pselldocoll1lilelllbllS PIISil/IIS (Quennerstedt,
Family Uronematidae 1869) Foissner and Wilbert, 1981 (Fig. 8.29)
Urollellla lIlarilllllll Dujardin, 1841 (Fig. 8.27). (=Lemblls pllsillllS, Pselldocohnilemblls persali-
Body barrel-shaped, anteriorly transversely trun- n liS ]. Body slender, ovoid, anteriorly narrowly,
cate; frontal plate nonciliferous. Without notch in posteriorly broadly rounded. Contractile vacuole
area of oral opening. Oral cavity shallow, extend- slightly subterminal. Longitudinal somatic kine-
ing over anterior 1/3-1/2 of body. Contractile vac- ties 8-11 in number, 1 elongate caudal cilium. Oral
uole in posterior end. Bipolar ciliary rows 12-16 in area extends over almost 1/2 of body length.
number, anterior portions composed of dikinetids, Adoral membranelles 1 and 2 and undulating
posterior of monokinetids; 1 elongate caudal ci- membrane arranged as longitudinal double mem-
lium. Adoral membranelle 1 usually composed of brane; adoral membranelle 3 forming small square
single row of 4-9 basal bodies, rarely with very patch of basal bodies. Swims moderately fast,
short second row; adoral membranelle 2 com- rotates rapidly sometimes, never rests. May form
posed of2-3 basal body rows; adoral membranelle resting cysts. Size in vivo 25-50 x 10-20 ¡lm.
3 formed by small patch of basal bodies. Paroral Usually found in putrid habitats, often occurs
membrane on right in oral cavity, anteriorly termi- together with Uronema marinum. Ref.: Foissner
nating adjacent to adoral membranelle 2. Scutica and Wilbert (1981), Aescht and Foissner (1992),
consisting of few kinetosomes. Swims rapidly. Foissner el al. (1994).
Size in vivo 20-60 x 10-40 ¡lm. Ref.: Kahl (1931),
Dragesco and Dragesco-Keméis (1986), Petz el al. Family Pleuronematidae
(1995a), Song and Packroff(1997). Plellrollellla craSSlIlIl Dujardin, 1841 (Fig. 8.30)
(=Plellronema chlysalis]. Body ovoid, right side
Family Cohnilembidae straight, left convex, anteriorly slightly narrowed
Collllilelllblls verlllilllls (Müller, 1786) Kahl, but both ends broadly rounded; late rally more or
1933 (Fig. 8.28) (=Cohl1ilemblls illfllsiollllm, les s flattened. Oral area extending from anterior
Lemblls infllsionllm, L. verm inLIS,L. elol1gallls]' pole to posterior 1/6 of body. Contractile vacuole
Body elongate cylindrical, anterior half narrowed; dorsal in posterior 1/5 ofbody. Resting extrusomes
highly flexible. Cross section circular. Pellicle rod-shaped according to Kahl's (1931) drawing,
slightly grooved by kineties. Produces hyaline distinct in life. Somatic kineties densely ciliated,
mucilaginous lorica when undisturbed. Oral cavity cilia on posterior pole not elongated. Movement
Ciliophora 295
rapidly to and fro, rests with cilia spread when elongate C- to horseshoe-shaped, transverse to
feeding. Length 70-120 11m. Ref.: Kahl (1931). main body axis. One contractile vacuole at ventral
wall of vestibulum. Colonies with few (about 10)
Family Cyelidiidae zooids on dichotomously branched stalk, up to 550
• CyclidiulII glaucollla Müller, 1773 (Fig. 8.31). Ilill high. Main sta1k long, lateral stalks short (1
Body ellipsoid to barrel-shaped, starving speci- zooid/branch), without 10ngitudinal striations;
mens fusiform; anterior end nonciliferous, nar- myoneme connected at ramifications, extending to
rowed and set off forming frontal plate. distal end and twisting regularly. Zooids in vivo
Sometimes slightly flattened laterally. One sphe- about 80 x 30 11m. Ref.: Hamburger and
rical macronucleus; disintegrated to 2-4 globules in Buddenbrock (1911), Kahl (1935).
a terrestrial population. Micronucleus in indenta-
tion of macronucleus. Contractile vacuole inva- Family Spirostomidae
riably in posterior end ofbody. Resting extrusomes • Alligsteinia clarissima (Anigstein, 1912)
rod-shaped and indistinct. Somatic kineties 9-11, Hirshfield et al., 1973 (Fig. 8.34) [=Blepharisma
usually 10, in number with paired basal bodies in clarissimum]. Body elongate, band-like, flexible,
anterior half. One long caudal cilium distinctly slightly contractile but without torsion, laterally
bent distally. Oral apparatus half as long as body, distinctly compressed. Macronucleus moniliform,
composed of 3 small adoral membranelles and 1 composed of 9-50 nodules. Several micronuclei.
paroral membrane with long cilia (spread sail-like Contractile vacuo le in rear end, with long collect-
in resting specimens). Movement jumping and ing canal extending anteriorly on dorsal side.
gliding, rests with cilia rigidly spread. Size in vivo Cortical granules in interkinetal rows; body
14-30 x 6-16 /lm. There are several rather similar colourless according to Kahl (1932) and Isquith
C)'clidillm spp., which need reinvestigation with and Repak (1974), pink according to Ricci el al.
modern silver impregnation. Ref.: Didier and (1982). About 24-32 somatic kineties. Adoral
Wilbert (1981), Song and Wilbert (1989), Foissner zone extending over about 1/2 of body length.
et al. (1994), Foissner (1996). Inconspicuous undulating membrane on right of
oral area. Size 160-600 x 20-65 Ilill. Usually a
Family Vortieellidae benthic species. Ref.: Kahl (1932), Hirshfield el
Vorticella oceallica Zacharias, 1906 (Fig. 8.32). al. (1973), Isquith and Repak (1974), Ricci et al.
Body conical to bell-shaped. Peristomial disc flato (1982), Dragesco and Dragesco-Kernéis (1986).
Pellicle with 25-34 closely spaced striations. One Gruberia lallceolata (Gruber, 1884) Kahl, 1932
macronucleus vermiform and J-shaped, some- (Fig. 8.35) [=Spiroslomum lanceolatllf7l]. Body
times elongate bean-shaped. One micronucleus. lanceolate, posteriorly narrowed tail-like; flexible,
Single contractile vacuole at ventral wall of ves- contracts without torsion (about 50% of body
tibulum. Epibiotic on the diatom Chaetoceros. length). Macronucleus moniliform, composed of
Size of zooid 18-34 x 22-44 Ilill. Stalk about 80- about 7-27 nodules. Very likely with several
200 /lm long, distally widened (3-5 11macross). micronuclei. Contractile vacuo le in posterior end.
According to Noland and Finley (1931) and Cortical granules between kineties, specimens thus
Warren (1986) possibly conspecific with V stria- perhaps reddish. Somatic kineties 32-38 in num-
ta Dujardin, 1841; according to the redescription ber. Adoral zone extending along 1/3-1/4 of body,
by Nagasawa and Warren (1996) a separa te spe- composed of about 150 membranelles; cytostome
cies. Ref.: Nagasawa and Warren (1996). invaginated. Undulating membrane inconspicuous,
ZoothallllliulII elegalls d'Dudekem, 1864 (Fig. fragmented, composed of about 40 short rows of 3-
8.33). Zooids slenderly campanulate, peristomial 5 cilia. Thigmotactic, attaches to substrate with
disc slightly slanting, constricted beneath peristo- posterior portion. Swims in helicallines. Size 200-
mial collar, pellicle smooth; often bent over or 1100 x 15-55 /lID. There are several rather similar
held at angle to stalk, which should contract in zig- species which differ only in the shape of the undu-
zag manner according to genus diagnosis. lating membrane (Dragesco and Dragesco-
Contracted specimens distinctly curved, with pe- Kérneis, 1986). Ref.: Xahl (1932), Borror (1963),
ristomial collar drawn up snout-like. Macronucleus Dragesco and Dragesco-Kérneis (1986).
296 Ciliophora
Family Condylostomatidae dorsal side very likely without cilia. Adoral zone of
Condylostollla patells (Müller, 1786) Dujardin, membranelles extending along anterior margin
1841 (Fig. 8.36). Body nearly cylindrical, length from mid-way on left to mid-body on right. Size in
to width ratio 4-8: 1, posterior half slightly wid- vivo about 100-130 x 50-70 llm. Kahl (1932) erect-
ened sac-like, rear end broadly to narrowly rounded a new species for the population studied by
ed, contractile; starved specimens dorsoventrally Fauré-Fremiet (1924), P. ¡al/reí (dorsal side
flattened. Posterior end often attached to detritus. uneven, with bristle-Iike cilia). The specimens
Macronucleus moniliforrn, composed of8-25 nod- studied by Wailes (1943) belong apparently also to
ules. Possibly several (about 13-18) micronuclei. P. ¡al/reí (Song and Wilbert, 1997). Ref.:
Contractile vacuóle in posterior end, with collect- Hamburger and Buddenbrock (1911), Kahl (1932).
ing canal extending anteriorly, latter sometimes
originating from confluence of several vacuoles. Family Licnophoridae
Cortical granules in 2-3 irregular interkinetal • LiclIop/lOra allerbachi (Cohn, 1866) Claparéde,
rows. About 40 somatic kineties. Oral cavity trian- 1867 (Fig. 8.39) [=Tríchodína al/erbachi). Body
gular. Adoral zone of membranelles extending hourglass-like in ventral view, contractile; promi-
over 1/4-1/6 of body. Conspicuous undulating nent apical oral disc flattened; stalk short and
membrane on right of oral cavity. Apical membra- thick, adhesive disc circular to elliptical.
nelles (cirri) 4-5 in number, frontal, on right of Macronucleus moniliform, composed of 10-25
oral cavity. Swims rather rapidly. Size in vivo nodules. Somatic ciliature reduced, composed of
about 300-560 x 45-100 llm. Ref.: Kahl (1932). small patch of lateral cilia consisting of few short
ciliary rows, several ventrally and dorsally inter-
Family Stentoridae rupted concentric kineties (ciliary rings) on adhe-
• Stentor polYlllorp/llIs (Müller, 1773) Ehrenberg, sive disco One transverse ciliary row dorsally on
1830 (Fig. 8.37). Extended specimens elongate stalk (dorsal kinety). Adoral zone composed of
cone- to trumpet-shaped, highly contractile; con- >80 membranelles and almost completely sur-
tracted individuals globular to pyriforrn. Sessile, rounding oral disco Paroral membrane extends into
posterior end attaches to substrate with holdfast, oral cavity. Ectocornmensal on marine organisms.
may detach when disturbed. Macronucleus indis- Size 80-120 x 45-70 J.lffi. Ref.: Kahl (1932),
tinctly moniliforrn, composed of about 5-20 no- Villeneuve-Brachon (1940), da Silva Neto (1994).
dules. Contractile vacuole left of oral cavity, with
long, posteriorly directed collecting canal. Key to oligotrich genera
Colourless cortical granules in interkinetal rows. This key covers all the well-known aloricate genera,
About 100 longitudinal somatic kineties. Adoral including those so far only recorded from fresh
zone of membranelles descending in clockwise waters. However, many of these were established as a
spiral into oral cavity, composed of>200 membra- result of splitting the genus Slrombidil/m, which is
nelles; undulating membrane paralleling adoral widespread in marine habitats, suggesting that they
zone. About 50 ciliary rows on peristomial field. may be recorded in marine environments in the future.
Specimens appear green from symbiotic a1gae.
Length of extended specimens in vivo up to 2000 Adoral zone of membranelles forrning closed cir-
llm. Ref.: Dragesco and Dragesco-Kernéis (1986), cle in top view (Strobilidiids, Tintinnids) 11
Foissner el al. (1992). 1a Adoral zone of membranelles forrning open cir-
cle in top view (Halteriids, Strombidiids) 2
Family Peritromidae 2 Several rows of longitudinal soma tic kineties
• Per;trollllls ellllllae Stein, 1862 (Fig. 8.38). Body (Halteriids) 3
broadly elliptical to slightly reniforrn, contractile, 2a Somatic kineties transversely or spirally
posterior dorsal half vaulted; dorsoventrally flat- arranged (Strombidiids) 5
tened. Cytostome in or anterior to mid-body on 3 Soma tic kineties very short, cilia very long and
left. Two macronuclei, 1 micronucleus. Contractile stiff, fused to cirri-like bristles .4
vacuole subterrninal in posterior dorsal portion. 3a Somatic kineties distinctly longer, cilia soft, not
Somatic ciliary rows about 18-24 on ventral side; fused to bristles: Meseres (Fig. 8.42)
Ciliophora 297
4 Somatic kineties without horizontal row close Family Strombidiidae

below each bristle complex: Halteria (Fig. 8.40) • StrolllbidiulIl alltarcticulIl (Busch, 1930) Kahl,
4a Somatic kineties with additional horizontal row 1932 (Fig. 8.56) (=Buehringa anlarclicum,
close below each bristle complex: Pelagohalteria Slrobilidium anlarclicum). Body obovoid to
(Fig.8.41) almost spherical, no anterior protrusion; probably
5 Body with elongate tail: TOlltollia (Fig. 8.43) containing sequestered chloroplasts. Peristomial
5a Body without tail 6 field large, oral cavity separated from ventral SUf-
6 Cytopharynx lined with thick fibres, forming face by cytoplasmic wall. Equatorial kinety con-
basket: Cyrtostrolllbidilllll (Fig. 8.45) tinuous or with very small gap ventrally. Ventral
6a Cytopharynx without basket... 7 kinety usually right of median. Adoral zone con-
7 With permanent neoformation organelle 8 tinuous and of large diameter, composed of 14-17
7a Without permanent neoformation organelle 9 anterior and 6-7 ventral adoral membranelles
8 With ventral kinety: Lilllllostrolllbidilllll (Fig. which invaginate on anterior surface. Extrusomes
8.46) in 1 supraequatorial girdle, about 20 flrn long.
8a Without ventral kinety: Pelagostrolllbidilllll Subpellicular platelet layer in posterior 2/3 of
(Fig.8.44) body. Swims rather slowly, rests briefly and
9 Equatorial kinety transversely arranged, not rotates about main body axis. Size in vivo 35-55 x
extending to posterior body pole, ventral kinety 20-40 flill. Ref.: Petz el al. (1995a).
at about right angle to equatorial kinety: • Strolllbidilllll COlllpresslllll (Leegaard, 1915)
Strolllbidilllll (Fig. 8.49) Kahl, 1932 (Fig. 8.57) (=Laboea compressa].
9a Equatorial kinety distinctly spiralling, extending Body conical, posterior portion longitudinally
to posterior pole, ventral kinety parallel to equa- striped (owing to extrusomes or subpellicular
torial kinety 10 platelet layer) and bluntly pointed. Oral cavity
10 Equatorial kinety with about I turn around body: inconspicuous. Continuous equatorial kinety
Spirostrolllbidilllll (Fig. 8.47) almost in mid-body, composed of monokinetids
lOa Equatorial kinety with several turns around (preparation artifact?). Short ventral kinety in pos-
body: Laboea (Fig. 8.48) terior 1/3 of body. Adoral zone continuous, com-
II With distinct lorica: Tintinnids (not treated here) posed of 12-15 anterior and 6-8 ventral adoral
lla Lorica lacking or indistinct 12 membranelles. One equatorial girdle of about 10
12 External adoral membranelles separated into flill-long extrusomes (after protargol impregna-
outer and inner portions (sometimes with indis- tiori). Size of protargol-impregnated specimens
tinct hyaline lorica): Leegaardiella (Fig. 8.55) 17-28 x 13-24 flill. Ref.: Kahl (1932), Lynn el al.
12a External adoral membranelles not as above .... 13 (1988).
l3 Some somatic kineties form spiral at posterior • Strolllbidilllll cOllicoides (Leegaard, 1915) Kahl,
end of body: Strobilidilllll (Fig. 8.50) 1932 (Fig. 8.58) (=Woodania conicoides). Poorly
13a Somatic kineties not as above 14 studied (specimens considerably deformed from
14 Somatic kineties longitudinal and transversely fixation). Shape of subpellicular platelet layer
arched: Pelagostrobilidilllll (Fig. 8.52) conical. Size of preserved individuals 30-41 x 37-
14a AII somatic kineties more or less longitudinal 44 flill. According to Kahl (1932) possibly conspe-
......................................................................... 15 cific with S. sulcalum. Ref.: Kahl (1932) .
15 Many (usually >10) somatic kineties extending • Strolllbidilllll cOlliclllll (Lohmann, 1908) Wulff,
over entire length of body: StrolllbidillOpsis 1919 (Fig. 8.59) (=Laboea conica]. Body elongate
(Fig.8.53) obconical, with anterior protuberance, posteriorly
15a Few short somatic kineties 16 pointed; often yellow. Oral cavity shallow.
16 Somatic kineties ciliated: Rilllostrolllbidilllll Macronucleus spherical, shape highly variable in
(Fig.8.51) protargol-impregnated cells. Equatorial kinety
16a Somatic kineties nonciliated: Lohlllalllliella supraequatorial, perhaps composed of monokine-
(Fig.8.54) tids. Ventral kinety rather short and consisting of
10-25 monokinetids (preparation artifact?),
298 Ciliophora
extending anteriorly from posterior cell pole. (1932), Fauré-Fremiet and Ganier (1970).
Adoral zone discontinuous, composed of 16 ante- Montagnes et al. (1990).
rior and 10-17 ventral adoral membranelles. • Laboea strobila Lohmann, 1908 (Fig. 8.63)
Extrusomes in girdle about 30 ¡.tm-Iong (in protar- [=Strombidium strobilunz, Strombidiunz strobilus,
gol slides) inserted in anterior 1/3 ofbody. Size of Conoeylis helix]' Body elongate obconical, appears
protargol-impregnated specimens 71-106 x 36-61 screw-like because of interruption of subpellicular
¡.tm; in vivo 20-75 ¡.tm long. Ref.: Kahl (1932). platelet layer by equatorial kinety and extrusome
Montagnes et al. (1988b). girdle ("body whorls"). Macronuclear nodules 36-
• Strolllbidilllll crasslIIlllll (Leegaard, 1915) Kahl, 72 in number. Equatorial kinety composed of
1932 (Fig. 8.60) r=Laboea erassula]. Body obcon- monokinetids with short cilia, originates left of oral
ic, with inconspicuous protuberance on right. Oral cavity and spirals around body in 4-5 helical turns
cavity conspicuous. Equatorial kinety discontinu- towards posterior cell po le. Short ventral kinety
ous ventrally. composed of densely spaced cilia. usually on dorsal side between equatorial kinety
Ventral kinety extending from posterior cell por- and posterior pole, consisting of 12-15 nonciliated
tion almost to equatorial kinety, usually distinctly dikinetids. Adoral zone of membranelles discon-
left of cytostome (artifact?). Peristomial field continuous, composed of 12-18 anterior and 12-22
spicuous. Adoral zone continuous, composed of ventral adoral membranelles. Paroral membrane on
16-19 anterior and 13-19 ventral adoral membra- right in oral cavity. Girdle of short (exact length
nelles. Extrusomes in 1 equatorial girdle about 17 not given) extrusomes parallel to equatorial kinety.
¡.tm-Iong. Subpellicular platelet layer in posterior Swims rather rapidly in zig-zag trajectory. Size of
1/2-1/3 of cell, longitudinally furrowed. Size in fixed specimens 44-110 x 20-60 ¡.tm.Retains func-
vivo 50-70 x 30-40 ¡.tm. Ref.: Petz et al. (1995a). tional chloroplasts from algal food and is thus mi x-
• Strolllbidilllll diverslIlII Busch. 1930 (Fig. 8.61) otrophic (Laval-Peuto and Rassoulzadegan, 1988).
[=Strobilidium diversum]. Body conical, subpel- Ref.: Fauré-Fremiet (1924), Montagnes et al.
Iicular platelet layer(?) with oblique groove (per- (1988b), Lynn and Gilron (1993).
haps equatorial kinety). Cilia of adoral membra-
nelles very long. Size 10-39 x 10-25 ¡.tm. Poorly Family Strobilidiidae
studied species. Ref.: Kahl (1932). • Strobilidilllll typiClllll (Lankester, 1874) Fauré-
• Strolllbidilllll slllcatlllll Claparede and Fremiet, 1924 (Fig. 8.64) [=Torquatella typiea,
Lachmann, 1859 (Fig. 8.62) [=Strombidillm nzinll- Strombidillm typicum]. Body elongate, posterior
tum, Strobilidiunz suleatum]. Body with prominent portion widened sac-like. Oral cavity large, inva-
anterior protuberance. Oral cavity short. ginating on anterior surface. Two macronuclear
Contractile vacuole in mid-body on left. nodules, 1 micronucleus. Contractile vacuole on
Equatorial kinety subequatorial, composed of di- posterior pole. Cytoplasm strongly vacuolized.
kinetids. Ventral kinety extending from equatorial Somatic ciliature lacking(?). Adoral zone almost
kinety across posterior pole a short distance onto as wide as body, forming closed circle in top view,
dorsal surface. Adoral zone continuous, composed composed of approximately 30 membranelles.
of about 12 anterior and 7-8 ventral adoral mem- Size in vivo about 80 x 60 ¡.tm. Re-investigation
branelles. Single extrusome girdle behind mid- with modern silver impregnation needed to con-
body, extrusomes inserted anterior to equatorial firm generic c1assification. Ref.: Fauré-Fremiet
kinety. Subpellicular platelet layer in posterior 1/3 (1924), Kahl (1932).
ofbody. Movement moderately fast by jerks. Size • Pelagostrobilidilllll spirale (Leegaard, 1915) Petz
30-60 x 25-30 )lm. This characterization mainly et al., 1995a (Fig. 8.65) [=Lohmanniella spiralis.
follows Montagnes et al. (1990), who based it on Strobilidium spiralis]. Body subspherical, flattened
Fauré-Fremiet's work and critically reviewed on one side. One C-shaped macronucleus, 1 micro-
many previously published descriptions of this nucleus. Five somatic kineties with short cilia cov-
species. According to these authors, numerous ered by cytoplasmic flap; kinety 1 slightly S-Iike,
populations identified as S. suleatum, e.g., by extending anteriorly from posterior pole; kineties 3
Lynn et al. (1988) and Fenchel and Jonsson and 4 coursing almost longitudinally, shortened;
(1988), be long to a different species. Ref.: Kahl kinety 2, originating at posterior pole, and kinety 5
Ciliophora 299
distinctly arched describing a partial circle. Adoral ellipsoidal, high1y flexible, slightly contractile.
zone composed of33-39 external and 8-20 internal Two macronuclei, 2-3 micronuclei. Contractile
adoral membranelles. Size of protargol-impregnat- vacuole near mid-body on 1eft. Frontoventral cirri
ed specimens 40-60 x 40-52 ¡.Lm.It was not defini- conspicuously shifted backwards, some behind
tively identified by Leakey et al. (1994b) in the leve1 of cytostome. Three frontal, 3 frontoventral,
Southem Ocean; Petz et al. (1995a) found it in the 1 buccal, 3 postora1 ventral, 2 pretransverse and 4-
Weddell Sea. PelagostrobilidiulI1 neptllni 6 transverse cirri. Right marginal cirral row in pos-
(Montagnes and Taylar, 1994) Petz et al., 1995a is terior half of cell, composed of 3-9 cirri, not con-
very similar, but differs by having 2 micronuclei fluent with 1eft marginal cirra1 row. Dorsal
and kinety 5 originates near the rear end and kineties 5-6 in number, with about 10 Jlffi long
extends anteriorly. Ref.: Lynn and Montagnes ci1ia and 3 fine caudal cirri. Adoral zone of mem-
(1988), Montagnes and Taylor (1994). branelles extending over about 20% of body
1ength, composed of 13-18 membranelles of
Family Urostylidae which first 5 appear spine-like, stiff. Movement
• H%sticha {liademata (Rees, 1883) Kahl, 1932 jumping, rests in between. Size 50-100 x 30-40
(Fig. 8.66) [=Amphisia diademata]. Body elongate ¡.Lm. The species was recently transferred to
ellipsoidal, posteriorly broad1y rounded. Two Oxytricha (Song et al., 1991), which is not fol-
macronuclei,2 micronuclei. Contractile vacuo le in 10wed here since its morphogenesis, an essential
mid-body on left. One buccal, 3 frontal, 2 fronto- feature for a generic classification in oxytrichids
terminal and 6-9 (11) transverse cirri in J-shaped (Berger and Foissner, 1997), is not known and its
row. Two midventral rows (midventral cirral dorsal ciliary pattern is rather peculiar. Ref.: Kahl
pairs) extend almost to transverse cirri, composed (1932), Dragesco (1963), Song et al. (1991).
of about 7-15 obliquely arranged cirral pairs. One Trache/osty/a pedicll/ijormis (Cohn, 1866) Kahl,
right and 1 left marginal cirral row, the latter with 1932 (Fig. 8.68) [=Stichochaeta pedicllliformis,
oblique anterior portion. Caudal cirri lacking. Gonostomllm pedicllliforme]' Body elongate rec-
Adoral zone extending over about 40% of body tangular, about anterior 1/3 attenuated (extent
length, composed of 25-28 membranelles of depending on nutritional state), posteriorly broad-
which about first 7 are slightly separated from oth- Iy rounded; flexible, fragile. Dorsoventrally flat-
ers. Undulating membranes about 20 ¡.Lmlong. tened. Macronuclear nodules 16-64 in number; 2
Movementjerking to and fro. Size 45-140 x 20-55 micronuclei. Three frontal, 7-8 ventral, 2 pretrans-
11m. This species differs by the position ofthe con- verse, 5(6) transverse cirri; 1 right and 1 left mar-
tractile vacuole (in mid-body) from the very simi- ginal cirral row not confluent posteriorly. Three
lar H. pllllaster (in posterior 1/3), with which it is dorsal kineties with long and fine cilia, perhaps 3
often confused. Ref.: Kahl (1932), Borror (1963), e10ngate caudal cirri. Adoral zone extending along
Borror and Wicklow (1983), Wi1bert (1986). 1eft margin over 1/3-1/2 ofbody length, composed
• Trichofotaxis crassa (Claparede and Lachmann, of about 40 thickened membranelles; undulating
1858) Kahl, 1932 (Fig. 8.69) [=Amphisia crassa, membranes short, on right of oral cavity. Quickly
Trichotaxis crassa]. Body broadly fusiform, nar- crawling forward and backward, frequently raises
rowed posteriorly, right margin distinctly convex, anterior portion above substrate, thigmotactic.
vaulted dorsally. Macronucleus possibly fragment- Size in vivo 100-250 x 20-40 Jlffi. Usually a
ed. Colour yellow-brown to reddish, sometimes mesopsammal species, but also found in the ben-
colourless. Two left marginal, 1 right marginal, 2 thos and plankton. Very likely closely related to
midventral cirral rows. Adoral zone of membra- oxytrichids. Ref.: Kahl (1932), Borror (1963,
nelles along about 1/3 of body 1ength, extending 1972b), Maeda and Carey (1984).
far onto right side. Size 110-150 x 45-55 Jlffi. Ref.:
Hamburger and Buddenbrock (1911), Kahl (1932). Family Aspidiscidae
• Aspidisca /Yllcells (Müller, 1773) Ehrenberg,
Family Oxytrichidae 1830 (Fig. 8.70). Outline slightly triangular, right
• Acfillotricha sa/fans Cohn, 1866 (Fig. 8.67) margin convex, left almost straight, broadly
[=Tachysoma saltans, Oxyfricha saltans]. Body rounded anteriorly, transversely truncate posteri-
300 Ciliophora
orly, with more or less distinct projection in oral 4 micronuclei. Contractile vacuole in posterior 1/3
area. Dorsoventrally flattened. Dorsal surface con- on right. Five to six frontal, 2 pretransverse, 4-5
vex, smooth, occasionally with several indistinct transverse, 1-3 left marginal cirri. Five to six dor-
longitudinal ribs. Pellicle stiff. Macronucleus sal kineties often composed of about 6-10 cilia
horseshoe-shaped, 1 micronucleus. Contractile each, in some populations interrupted in mid-
vacuole at level of transverse cirri. Seven ventral, body; 3 strong caudal cirri. Adoral zone of mem-
5 transverse cirri. Dorsal kineties 5-6 in number, branelles extending over about 1/2 of body length,
no caudal cirri. Oral apparatus bipartite: 3-4 tiny composed of 30-37 membranelles. Movement
frontal membranelles in deep indentation on ante- moderately rapid. Size 50-120 x 30-70 ¡lm. Ref.:
rior left, slightly' covered by anteriormost frontal Kahl (1932), Curds and Wu (1983), Dragesco and
cirrus; 10-15 adoral membranelles in oval oral Dragesco-Keméis (1986), Hill and Borror (1992),
cavity on posterior left. Six longitudinal dorsal si1- Petz et al. (1995a), Song and Packroff (1997).
verlines, rightmost connects basal bodies arc-like, • Euplotes charoll (Müller, 1773) Ehrenberg, 1830
second from right shortened and only in posterior (Fig. 8.73). Body broadly elliptical to slightly tri-
body half; ventral silverline system wide-meshed. angular; 7-9 distinct longitudinal dorsal ribs. Oral
Size 35-50 x 30-45 ¡lm. Ref.: Wu and Curds cavity large, V-shaped; Iip on right border extends
(1979), Song and Wilbert (1989), Foissner et al. almost to posterior poleo Macronucleus horseshoe-
(1991). shaped, 1 micronucleus. One contractile vacuole.
Ten frontoventral, 4-6 (usually 5) transverse, 2 left
Family Euplotidae marginal and 2-6 caudal cirri. Dorsal kineties 9-12
Urollychia trallsfllga (Müller, 1776) Stein, 1859 in number, with 10-40 cilia in central rowS. Adoral
(Fig. 8.72) (=Uronychia lIncillata, Uron)"chia ant- zone extends arch-like over 2/3-4/5 of body
arctica). Body almost rectangular, posteriorly length, composed of 51-70 membranelles. Dorsal
rounded to slightly tapering, anteriorly distinctly silverline system of double-eurystomus type. Size
convex, spur-like protrusion each on right and left 70-110 x 50-100 11m. Ref.: Kahl (1932), Tuffrau
side; usually 1 spine each dorsally near anterior (1960), Borror (1968), Curds (1975), Song and
end of dorsal kineties. Deeply caved in region of Packroff (1997).
transverse and left marginal cirri. Pellicle rigid. • Euplotes harpa Stein, 1859 (Fig. 8.74). Outline
Two macronuclear nodules connected by short elongate elliptical; 10-12 longitudinal dorsal ribs.
funiculus, 1 micronucleus. Contractile vacuole Oral cavity large; Iip on right border prominent,
anterior and right of transverse cirri. Six fronto- with characteristic protrusion in anterior portion.
ventral, 4-5 transverse, 3 left marginal and 3 Macronucleus C- to slightly 3-shaped, 1 micronu-
strong caudal cirri. Six dorsal kineties. Adoral cleus anteriorly near macronucleus. One contrac-
zone of membranelles bipartite, 11 anterior and 5 tile vacuo le. Ten frontoventral, 5 transverse, 2 left
posterior adoral membranelles. Undulating mem- marginal and 2 caudal cirri. Dorsal kineties 11-13
brane almost surrounding very large oral area. in number, with 36-45 cilia in central rows. Adoral
Movement usually jumping and swimming very zone extends over about 2/3 of body length, com-
fast, rests with adoral membranelles and cirri rig- posed of 52-70 membranelles. Dorsal silverline
idly spread; swims also slow and wobbly. Size in system of double-eurystomus type. Length 120-
vivo usually 80-110 x 50-65 ¡lm. Specimens hav- 260 ¡lm. Ref.: Kahl (1932), Tuffrau (1960), Curds
ing a nodulated C- or horseshoe-shaped macronu- (1975), Dragesco and Dragesco-Keméis (1986).
c1eus very likely belong to U. heinrothi • Ellplotes vallllus (Müller, 1786) Minkjewicz,
Buddenbrock, 1920 (for improved diagnosis see 1901 (Fig. 8.75) (=Monellplotes vannlls). Body
Petz et al., 1995a). Ref.: Kahl (1932), Curds and elongate elliptical to D-shaped, without distinct
Wu (1983), Hill (1990), Petz et al. (1995a). dorsal ribs. Oral cavity narrow, extending over 3/4
Diophrys appelldiculata (Ehrenberg, 1838) of body length. Macronucleus irregularly horse-
Levander, 1894 (Fig. 8.71) (=Stylonychia appen- shoe- to C-shaped. One contractile vacuo le. Ten
diclllata, Diophrys marina). Body outline ellipti- frontoventral, 5 transverse, 2 left marginal and 2-4
cal, left margin sometimes slightly concave, caved caudal cirri. Dorsal kineties 9-10 in number, with
in region of caudal cirri. Usually 2 macronuclei, 2- <16-22 cilia in central rows. Adoral zone extend-
Ci1iophora 301
ing over 2/3 of body length, composed of 53-70 Kahl A. 1932. Urtiere oder Protozoa 1: Wimpertiere
membranelles. Dorsal silverline system usually of oder Ciliata (Infusoria) 3. Spirotricha. Tierwelt
single-vannus type, rarely with irregular ramifica- Deutschlands, 25:399-650.
tions (Song and Packroff, 1997). Movement slow- Kahl A. 1933. Ciliata libera et ectocornmensalia.
Iy gliding. Size in vivo 75-200 x 65-90 ¡.1m. Tierwelt der Nord- und Ostsee, 23:29-146.
lankowski (1979) transferred E. vannus to Kahl A. 1934. Suctoria. Tierwelt der Nord- und Ostsee,
Moneuplotes (as type species), which was recent- 26:184-226.
Iy followed by Borror and Hill (1995). Song and Kahl A. 1935. Urtiere oder Protozoa 1: Wimpertiere
Packroff (1997) synonymized E. mutabilis, E. ros- oder Ciliata (Infusoria) 4. Peritricha' und
eoffensis and E. sharuri with E. vannus. Ref.: Kahl Chonotricha. Tierwelt Deutschlands, 30:651-886.
(1932), Tuffrau (1960), Curds (1975), Song and This series of papers by Kahl ineludes comprehen-
Packroff (1997). sive identification guides to many ciliate taxa
(marine, freshwater and soil) based on fife obser-
vations (no silver impregnation data; in German).
Suggested readings Laybourn-Parry 1. 1992. Protozoan plankton ecology.
Chapman and Hall, London, pp. 1-231. Excellent
Corliss 1.0. 1979. The ciliated protozoa. Charac- review on proto:::oan plankton ecology (also
terization, classification and guide to the literature ineludes freshwater and other proto:::oan groups).
(2nd ed.). Pergamon Press, Oxford, pp. 1-455. Lee 1.1., Soldo A.T. 1992. Protocols in Protozoology.
Exeellent general introduetion to ciliates, review of Society of Protozoology, Lawrence, Kansas.
classifieation systems, characterizes higher-level Collection of various methods for ecological, mor-
taxa and lists all valid genera at that time. phological and cytological studies of ciliates.
Fenchel T. 1988. Marine plankton food chains. Ann. Pierce R.W., Turner 1.T. 1992. Ecology ofplanktonic
Rey. Eco!. Syst., 19: 19-38. Summarizes the roles ciliates in marine food webs. Rey. Aquat. Sci.,
of ciliates in marine pelagic ecosystems. 6:139-181. Concise summGlY on ecology and
Foissner W. 1991. Basic light and scanning electron some methodological aspects.
microscopic methods for taxonomic studies of ci1- Puytorac P. de (ed.) 1994. Traité de Zoologie.
iated protozoa. European 1. Protisto!., 27:313-330. Anatomie, Systématique, Biologie. II. Infusoires
Detailed deseriplions of many useful techniques Ciliés. Masson, Paris, pp. 1-880. Excellent review
for the taxonomic study of ciliates. of ciliate cytology, pro vides a elassification
Hausmann K., Hülsmann N. 1996. Protozoology, 2nd scheme (in French).
ed. Thieme Verlag, Stuttgart, pp. 1-338. Excellent Reid P.e., Turley e.M., Burkill P.H. 1991. Protozoa
overvien' of protozoans (biochemistry, molecular and their role in marine processes. NATO ASI
biology, ultrastructure, morphofogy), profusely Series, G 25: 1-506. Session summaries providing
illustrated. short overviews of various aspects of work with
Kahl A. 1930. Urtiere oder Protozoa 1: Wimpertiere Protozoa (methodology, ecology, taxonomy, ete.).
oder Ciliata (Infusoria) 1. Allgemeiner Teil und Small E.B., Lynn D.H. 1985. Phylum Ciliophora
Prostomata. Tierwelt Deutschlands, 18: 1-180. Doflein, 190 l. In "An illustrated guide to the
Kahl A. 1931. Urtiere oder Protozoa 1: Wimpertiere Protozoa" (1.1. Lee, S.H. Hutner, E.e. Boyee,
oder Ciliata (Infusoria) 2. Holotricha auBer den im eds.), Allen Press, Lawrence, Kansas, pp. 393-
l. Teil behandelten Prostomata. Tierwelt 575. Ciliate elassification with identificalion keys
Deutschlands, 21: 181-398. and examples of common genera.
302 Ciliophora
B
A
(FL)
(FL)
e
(VV, SL)
Tiarina fusus
(RLV, FL)
Pro to cruzia
pigerrima
100
A
(extended
specimen,
FL)
e
(LLV, PI)
Tracheloraphis phoenicopterus
Ciliophora 303
A B
(FL) (DV, PI)
:í~j·~.jL
. .
,, ¡., ,·, ,,.
, ', •· I,
t?:{~
~ :,'
,.Yl·· <J'. i'.~"¿j'~
col>
, , , ,
·, ,'
,\,
:.4)
.
: ~l'
'.01"
. ",'4
I ~..••
~
....l'"i':
'.
,• B
•
;.....
~~·~~i'l'\,~ (resting cyst)
5". 1« ". 11!. ':0
'.~' '. ~~:i
~':,~{i:
~
.. ~, ••.¡';-•.
Chaenea teres
1"
e
(detailof
anterior area,
PI)
• Enchelyodon laevis
A
(FL)
B
(DV, PI)
A
(DV, FL)
Didinium gargantua
B Monodinium
~ (DV, PI) balbianii
304 Ciliophora
A
A
(LLV,
B e
(RLV, FL) (LLV, (RLV,
FL) PI) PI)
~ Litonotus fasciola
'!A!'
LoWil. Litonotus cygnus
A B
(LLV, (LLV,
FL) FL)
e
(IF, SI)
Kentrophyllum setigerum
Ciliophora 305
B
120 (PI)
A
(FU
GD Cyclotrichium gigas
(IJ) Myrionecta rubra
B
(oral area, IF, SI)
f¡ ... ". ", ". '. ~ ". " .
{I/,In' '"",.
(¡'II//// .- . .
liiJt:/~0;;' ". ........
'.
..
LHU.C
•••••••••• o"·· ••••
..........
,·,··.···.·,·.·.,.,
.•tt.····· "O .•• " "
~j~JI/j\\.··········
/(/((: .
Plagiopyla nasuta
I \:<~~~ I
30 :.:::::::.:;:': ..:::::::.::
o,, :.:: : j i::::::':::·':::'j::':':
: : .:.::~.~.:•••••• ~.:~.~.", .'
30
B
(VV,
PI)
~ Chlamydodon mnemosyne ~ Aegyriana oliva

306 Ciliophora
e
(LV,
FL) '\
1
~ Dysteria brasiliensis
A B
(VV, FL) (VV, PI)
(DV, FL)
Dysteria
compressa
,,~7 IF, PI)

sigmoides
~
~
Trochilia
e(oral 5
D Dysteria
area, monostyla ••
Ciliophora 307
B
(RLV,
PI)
A
(FL)
Iso
100
~ ~
~
~ Holop~rya
pelaglca Corynophrya
campanula ü?rynophrya
mterrupta B
(swarmer, FL, SI)
B
(VV,
PI)
'.
120
{) 1:'
....•
10\. ."
~ Acineta tuberosa
~ Uronema marinum
308 Ciliophora
A ,
',:,:
B
(LLV, :,':::, (VV,
FL) ,,::'
,,:,' PI)
":,,.
;: ~~:.
_1"
.:::"
:':'10
e
(IF and
10 ... silverline
system,
DVSI)
Pseudocohnilembus pusillus
D
(resting
~ cyst)
Cohnilembus verminus
,~~
..~:¡;. :: 5(/
'::f,:::: :v
'~y
•.
~,,'t:'::":
"'--,-tí;~ : ~.,;~ ",,/
10 l:':~
/1<. ··~\1~'~·:: ...

~
1~··,:i\L " ('"
B
(VV, PI)
(LLV, FL)
Pleuronema
~ crassum \
\
e
10 (DV, PI)
10
) Cyclidium
glaucoma
Ciliophora 309
~\III~(Zoothamnium
PI)\/§,
~~t
\
- -
e oceanica
Vorticella
B ~
I~~I
. f/J A
,
Gruberia Condylostoma
lanceolata
~ elegans
.(~:I
••• "......... .1"
FL)
(oral PI)PI)
(zooids,
area,
FL)
(LLV,
(RLV,FL)
patens I (VV,
111 BIIIII\~ •
I
' c1arissima
~ I
,
•••
: "l~~c
~\
~
120 I
T
125
An;gste;n;a
1""IWlI'
310 Ciliophora
A
(extended
specimen,
FL)
e
(VV, PI)
B
(contracted
speClmen, D
FL) (resting cyst)
Stentor polymorphus
B
(adhesive
disk,
B SI)
(LV,
FL)
Peritromus emmae
~
Licnophora auerbachi
Ciliophora 311
A
(VV, FL)
.
•
..
.
••
.....
-= ..
B
..
.. (somatic
kinety,
B PI)
(somatic
kinety,
PI)
Pe/agohalteria
~ Halteria
B
(DV, FL)
(VV, FL)
~ A1eseres Tontonia
~
€D ~
Pe/agostrombidium Cyrtostrombidium Limnostrombidium Spirostrombidium

312 Ciliophora
A
(VV, PI)
:::-" '\ 1 7;./
~~~~
B
(top view, PI)
Strobilidium
Strombidium
Laboea
B
(posterior view, PI)
Lohmanniella
B
(top view,
PI)
Strombidinopsis Leegaardiella
Ci1iophora 313
(DV, .
e ~lHr~):u\.
!I!-¡¡r~~
PII -':4~1Yí1%'r:Fniff11jr1f/~
110 lo
Gb
A
(VV,
PI)
B
(tap view, PI)
Strombidium
Strombidium compressum Strombidium
conicoides
comcum
G) Strombidium crassulum
314 Ciliophora
Strombidium diversum Strombidium sulcatum Strobilidium typieum
t,tt
t ....•" e B FL)
, ~PI) (IF, Laboea strobila(VV,spirale
Pelagostrobilidium
Holostieha diademata
,
B
t,
t t
, 110
Ciliophora 315
~j¿A
(VV,
FL)
FLÍ
(LV
§
f ~'\
.
(~y,(¡~?X
V fi.~ .
./
..~,:'.'
!:'·~I·:
• .". l
,
P/,,·-
'
rJf
l.XI'-::Z"
~,
;::::
'
,/ 1.$/
'$,,(~ (
r(
~/\'N ~:jj~W{'/(X\
./
Y.
l·
~',.' .'.. 'i.
...\ ....l~. .. . f ,
~q;·;(!i;:~I··) t.$.(J/.)
?.t~\1~d\~ ,
,.
#
,.
I \: '~~~ )1
{~\~~~~gJ.~\~ , # ,.
..".\' ~\"\~X
r:~~~':f~ 1,~~'~{~~
.~~_.. :r'O"'''~.,.
, I
~
~\
f (I:.Q..
..: •••~((/
t:J.i.{)/
\\;.f;.~Y
.
,.
::.'1
ylY
r".1~""
'~~~.J-
...~... o-O:\.•• )
~ \.• "'(l
-~~~~20l 2°1
(8 Act;notr;cha saltans
A e
(VV, (IF,
FL) PI)
\
~f~
!I.~.
..
~o\f'J=
==t O°a=
~O~~
~ .--
~O~·
120 S
:~~o"oo~,
'" , notice different
. 201
A
(VV,
B
(VV,
arrangernent of FL) PI)
I frontoventral cirri
~ Trachelostyla pediculiform;s ~ Tr;chototax;s crassa

316 Ciliophora
e
(DV,
PI)
E
(IF
o and
(FV, . silverline./
PI)
....
04'. system,
DV, SI)
Aspidisca Iynceus
B
(RLV,
FU
e
(VV,
PI)
/
/ (
~/ (,
l
, ,t/. f \ ..
\ \~
":.\.\
'....~:;;..
(C
•• 18.
11-
I ~,
20
Diophrys appendiculata
Ciliophora 317
A B e
(VV, (VV, (DV,
FL) PO PO
Q,-
a
, ),
a
~
~. .
~;:,
, ~,J
~,J
::~"'~
,
\
.\lJ
,
••
I
''
I
\ I:
I
Uronychia transfuga
e
(FV,
FL)
E
(DV, silverline system, PO
~ Euplotes charon
318 Ciliophora
D
(IF and silverline system,
e DV, SI)
(nuclear
apparatus, PI)
Euplotes harpa
e
(VV,
PI)
B
(LV,
FL)
D
(IF and silverline system,
DV, SI)
Euplotes vannus
Ciliophora 319
• Fig.8. IlIustrations ofthe species. Generally, figures are oriented with the anterior end ofthe organism towards the top ofthe page. Abbreviations:
DV: dorsal view; FL: from life; FV: frontal view; FX: after fixation; IF: infraciliature; LLV: left lateral view; LV: lateral view; PI: protargol impreg-
nation; RLV: right lateral view; SI: wet silver nitrate impregnation; SL: silver impregnation; VV: ventral view.
Figure sources:
From Aescht and Foissner (1992): 8.29A; from AlIgllstin and Foissner (1992): 8.70B, C, D; from Batisse (1994): 8.26A, B; from BOITor (1963):
8.35B, C, 8.66B, 8.68B; from Borror (l972a): 8.12B, C; from Borror (1972b): 8.66C, 8.68C, 8.69B; from Bütschli (1889): 8.IOA, 8.38A, B;
from da Silva Neto (1994): 8.39A, B; from de Puytorac el al. (1985): 8.15C, D; from Deroux (1976): 8.21; from Deroux and Dragesco (1968):
8.17B; from Dragesco (1960): 8.5A; from Dragesco (1972): 8.15A; from Dragesco and Dragesco-Keméis (1986): 8.7B, 8.IOB, 8.16A, B, 8.74A,
B, C; from Fauré-Fremiet (1924): 8.14; from Fauré-Fremiet (1970): 8.64; from Fauré-Fremiet and Ganier (1970): 8.62; from Foissner (1979):
8.7A; from Foissner (1982): 8.70A; from Foissner (1984): 8.8A, B, C; from Foissner (1987): 8.40A, B; from Foissner (1996): 8.31A, B, C; from
Foissner and Dragesco (1 996b): 8.IA, B, C; from Foissner and Wilbert (1981): 8.29B, C; from Foissner el al. (1988): 8.4IA, B, 8.51; from
Gourret and Roeser (1888): 8.19; from Grain el al. (1982): 8.13B; from Hada (1970): 8.61; from Hamburger and Buddenbrock (1911): 8.33B;
from Hoare (1927): 8.29D; from Kahl (1930): 8.3A, B, 8.5B; from Kahl (1931): 8.l7A, 8.28A, 8.30; frOI11Kahl (1932): 8.2, 8.35A, 8.36, 8.39C,
8.69A; from Kahl (1933): 8.37A; from Kahl (1934): 8.24, 8.25; frOI11Kahl (1935): 8.33A; from Klein (1929): 8.70E; from Krainer (1995): 8.44,
8.46; frol11Leegaard (1915): 8.58; from Lindholm (1985): 8.13A; from Lohmann (1920): 8.23; from Lynn and Gilron (1993): 8.45; from Lynn
and Montagnes (1988): 8.54, 8.55A, B, 8.65A, B; from Lynn el al. (1988): 8.57 A, B; from Maeda and Carey (1984): 8.68A; from Montagnes
and Taylor (1994): 8.53; from Montagnes el al. (1988b): 8.59, 8.63B; from Nagasawa and Warren (1996): 8.32A, B, C; from Patsch (1974):
8.37C; from Petz and Foissner (1992): 8.42, 8.50A, B; frol11Petz el al. (1995a): 8.4A, B, C, 8.6A, B, 8.20A, B, C, D, 8.22A, B, C, 8.27B, C,
8.43A, B, 8.47, 8.49, 8.52A, B, 8.56A, B, C, 8.60A, B, C, 8.71C, D, 8.72A, B, C; from Small and Lynn (1985): 8.3C, 8.28B, 8.34B, 8.63C;
from Sola el al. (1988): 8.15B; from Song and Packroff(1997): 8.18A, B, C, D, 8.27A. 8.71A, B, 8.73A, B, C, D, 8.75A, B, C; from Song and
Wilbert (1989): 8.9A, B, C; from Song el al. (1991): 8.67 A, B, C, D; from Stein (1867); 8.37D; from Tamar (1987): 8.11; from Tamar (1992):
8.12A; from Tartar (1961): 8.37B; from Tuffrau (1960): 8.73E, 8.74D, 8.75D; from Wilbert (1986): 8.66A; from Wulff (1919): 8.48, 8.63A;
from Yagiu (1943): 8.34A.
Tintinnoinea
Viviana A. Alder
Introduction 1992; Buck el al., 1992; Pierce and Tumer, 1992;

Garrison and Gowing, 1993).
Tintinnids are lorica-building, planktonic oligotrichid
ciliates ranging in size from 20 to 200 /lm. They consti-
tute a major component of the microzooplankton in General morphology and biology
most manne environments (Beers and Stewart, 1967;
Beers, 1982; Beers el al., 1982). They can also be Morphologic outline
important occasionalIy in estuarine and brackish waters
(Souto, 1973, Barría de Cao, 1981). Short generation The body of tintinnids is typical1y made up of a con-
times, high abundances, and fast reproduction rates, tractile cel1 which is attaphed by a stalk to the poste-
coupled with high grazing impact, enhance the impor- rior portion of the lorica (Fig. 1A, B). In most cases,
tance of tintinnids as a key trophic link between the the celI fllIs a little less than halfthe lorica, its volume
microbial and the metazoan compartments (Heinbokel varying with food availability and the life cycle stage.
and Beers, 1979; Brockel, 1981; Capriul0, 1982; lt has one micronucleus and 2 or more macronuclei,
Capriulo and Carpenter, 1983; Paranjape el al., 1985; and the commonest cytoplasmic inclusions are mito-
Porter el a/., 1985; Laval-Peuto el al., 1986; Paranjape, chondria, vesicular endoplasmic reticulum, lipid and
1987a, b; 1988; Garrison and Gowing, 1993). digestive vacuoles, granules of lorica-building
material (Fig. lE), and endocytoplasmic bacteria.
Tintinnid abundance in plankton samples, widespread Major external features include the ciliary and infra-
geographic distribution, and easily preserved loricae ciliary components 10cated in the somatic and buccal
probably account for numerous publications, particu- regions (Brownlee, 1977; Laval-Peuto et al., 1979;
larly on their taxonomy. Systematic surveys have been Laval-Peuto and Bronwlee, 1986; Sleigh, 1989). The
based almost exclusively on their 10ricae (see below). adoral membranelles (Fig. lA-D), around the buccal
The flrst description of a tintinnid is ascribed to MülIer regíon, commonly forming a closed spiral that com-
(1786;fide Kofoid and CampbelI, 1929), and the flrst prises 2 or 3 longitudinal rows of kinetosomas; the
comprehensive monograph on tintinnids was pub- paroral membrane (Fíg. IC), which consísts of a
lished by Daday (1887b). The studies ofBrandt (1906- single row of kinetosomas. The capsules (=extru-
1907), Laackmann (1907, 1910), Jorgensen (1924) somes), membrane-bound extrusible bodies usualIy
and Kofoid and CampbelI (1929, 1939) subsequently sub-pelicularly located (Corliss, 1979), of variable
provided descriptions and il1ustrations of over 1,000 shape, size and number; these capsules are aligned in
species. Kofoid and Campbell's (1929) work estab- striae (Fig. ID) associated to the adoral membra-
Iished a new classiflcation based on alI previous nel1es, and are probably involved in feeding process-
systematic studies. Thus, their revision has been a es (Laval, 1971, 1972; Laval-Peuto el al., 1979;
major reference for virtualIy alI subsequent publica- Capriul0 el al., 1986). The entire celI, including the
tions, both taxonomic and ecologic. During the last adoral membranelIes, the paroral membrane and the
decades, studies on tintinnids have focused mainly on striae, is enclosed by a continuous membrane known
their abundance in different environments, on their as perilemma (Laval-Peuto, 1975; Fig. lAB). Based
species-speciflc assemblages as related to currents and on their cytological characteristics, tintinnids are
water masses, as indicators of environmental settings, included in the suborder of the Ciliophora in the class
and particularly on their functional role in marine Polyhymenophorea, subclass Spirotrichia (Levine et
trophic chains, (e.g., Souto, 1979; Capriulo and al., 1980).
Carpenter, 1980, 1983; Hargraves, 1981; Verity,
1987; Boltovskoy el al., 1989, 1991; Wasik and The lorica of a tintinnid consists of a proteic bowl
Mikolajczyk, 1990; 1994; Alder and Boltovskoy, which is always open at its anterior (oral) end, and
1991a, b; Barría de Cao, 1992; Boltovskoy and Alder, usualIy (but not always; see Fig. 6.66-74) closed at the
South Atlalltic Zooplanf.ctoll.

edited by D. Boltol'skoy. pp. 311-384
1999 Backhu)'s Publishers. Leidell. The Netherlallds
322 Tintinnoinea
~~~klnety
~ 'dorsolateral
right lateral view

.. ,
...
@)~:~
•
..~~ ..
.
macronuclel
,ventral
in vivo kinety
cell aher protargol

silver impregnatlon
Cymatocylis convallaria
oral membranelles and striae
ISEM micrographs)
protolorica epilorica
paralorica
.. ~.
~ proter
~} ~~
I~
~~
asexual reproduction W
TINTINNID REPRODUCTION p":E O

favella ehrenbergii Different types
of epilonca
E3
E2
El
abnormal
individual nalied
individual
Sexual reproduction of partners

witli different loricae
Tintinnoinea 323
TYPES OF TI NTI N N ID (?) CYSTS
4)
Favella
serrata
0°
Cysts from sediment
trap materials
Cysts from
o
plankton samples
(i)
Fig. 1. General morphology of a tintinnid. A, C: from Petz el al. (1995a); B, H2: from Alder (1995); D: from Capriulo el al. (1986); E, F: modí-
fied from Laval-Peuto (1981) ; G: from Laval-Peuto (1983); H 1: from Laval-Peuto and Bronwlee (1986); H3: from Brandt (1906-1907); l, J: from
Paranjape (1980); K: from Reíd and John (1983); L: from Davis (1985); M. N: from Boltovskoy el al. (1996); O: from Reíd and John (1981).
posterior (aboral) end (Fig. lA, B). The oral end ofthe The lorica is built from posterior to anterior end.
lorica typically presents a collar which may be fenes- Lorical material (a chitin-like substance) is poured out
trated (Diclyocysta, Fig. 6.43-46), annulated through a pore or gutter, and then distributed around
(Codonellopsis; Fig. 6.39), everted (Cyttarocylis and the cell by the beating of the somatic ciliature (Laval-
Salpingella; Fig. 6.117 and 6.61, respectively), or Peuto, 1981). Ifthe first lorica (protolorica) built by
constricted (Codonella; Fig. 6.1-6). The aboral end the mother cell (proter) is lost, the naked mother cell
often bears a horn or a pedicel (a hom provided with (trophont) is capable of building a new, total!y spi-
a knob or a skirt, and a lance; Fig. 6.126, 6.127, ralled one (paralorica = coxlilelJa form) which can
6.129). The characteristics of the oral end are quite only be modified through the addition of helical epi-
constant generical!y, thus an important diagnostic loricae (Fig. lE, F, H; Laval-Peuto, 1977; Laval-
element. On the other hand, the aboral end and al! Peuto and Bronwlee, 1986). The structure of the lori-
structures related to it (knob, lance, skirt) are highly ca wall (continuou~ in most protoloricae; partially to
variable in length, width and general morphology, totally spiralled in paraloricae, epiloricae, and some
even at the intraspecific leve!. The various processes protoloricae; may be highly variable intraspecifically,
involved in building the lorica, and changes in the whereas its texture (which depends on its chemical
lorica during the tintinnid life cycle are thought to composition and the degree ofhardening ofthe lorical
explain, at least in part, the high intraspecific poly- material) is less variable (Laval-Peuto, 1981).
morphism observed in most species (Alder, 1995),
particularly in Antarctic ( van der Spoel, 1986; Some genera may accrete biogenic ancl/or mineral
Boltovskoy el al., 1990; Williams el al., 1994; Alder, particles to their loricae. These particles may cover
1995) and Arctic (Burkovskii, 1973; Davis, 1978, the whole lorica (Tintinnopsis complex; Fig. 6.12-27),
1981) forms. or mainly its bowl (Codonellopsis; Fig. 6.32-39).
324 Tintinnoinea
Although species typically with such an agglutinated Reproduction

\mlca ma)' often have a h)'aline lorica in the open
ocean, genera lacking this capability (Cymatocylis; Reproduction in tintimüds is cornmon\y by binary
Fig. 6.84-91) always remain hyaline, even in waters division (Fig. lE). However, sexual mechanisms (Fig.
with abundant sediment suspended from the sea floor 1G) such as isogamic conjugation or fusion of micro-
ancl/or the continental margino Generally, larger spe- and macrogametes have also been described for a few
cies have a hyaline lorica, while agglutinated ones are species (Gold and Pollingher, 1971; Laval-Peuto,
far more cornmon in the smaller taxa. Apparently, this 1983). According to their proportions in the popula-
phenomenon is an evolutionary adaptation to predation during sexual reproduction, there is a random
tion (Capriulo el al., 1982). association of organisms with different lorical shapes
(and even of naked individuals = trophonts; Laval-
Feeding Peuto, 1983). Field studies have shown that in coastal
and estuarine environments tintinnid reproduction
Because of their role as transferers of matter and ener- rates are 1-2 generations per day (Heinbokel, 1978a;
gy between the microbial and the metazoan cornmu- Burkill, 1982, Stoecker et al., 1983; Verity, 1986b),
nities, tintinnids are an important link in marine plank- exceptionally up to 3 (Verity, 1987). On the other
tonic food webs (Burkill, 1982; Laval-Peuto el al., hand, generation times in oceanic waters are usually
1986; Verity, 1985; Stoecker, 1984; Alder, 1995). longer, up to 42 hours (Heinbokel and Coats, 1986;
While some authors have pointed out that tintinnids Heinbokel, 1987, 1988).
can consume particles only about 40-45% of the
lorica's oral diameter (Spittler, 1973; Heinbokel, Cyst formation has been described for over 30 tintin-
1978a, b), others have observed that larger particles nid species (Reid and John, 1978, 1983; Davis, 1985;
are taken under food stress conditions (Gold and Krisinic, 1987; Fig. U-O). Cysts are generally flask-
Morales, 1975; Capriulo, 1982; Capriulo el al., 1986; shaped, and since they typically remain attached to
Alder, 1995). the inner wall of the lorica, their specific identifica-
tion is easier than for other protists. According to Reid
Tintinnids have been mentioned frequently as impor- and John (1981), highest concentrations of cysts are
tant grazers on detritus, picoplankton, bacterio-plank- found in sediments, their record in the water column
ton, and autotrophic and heterotrophic nannoplankton being only occasional. Recent information from
(Kopylov and Tumantseva, 1987; Hargraves, 1981). sediment trap samples (Boltovskoy et al., 1993a;
Furthermore, tintinnids can also consume microplank- 1996; Uliana, 1997), however, suggests that flux rates
tonic diatoms and dinoflagellates (Verity and of ciliate cysts in the water colurnn at depths up to
Villareal, 1986). Reports from coastal environments 4000 m can exceed 77,000 ind. m-2 d-1 (see Fig. 4
show daily consumption figures to 41 % of the stand- below). Encystment, which is seemingly a common
ing stock of chlorophyll a (Capriulo and Carpenter, phenomenon in the life cycle of tintinnids, probably
1980), and annual consumption up to 62% of the total has a dual function: to provide a mechanism for
5-10 flm chlorophyll a production (Verity, 1986a). repopulation as a response to environmental stress,
Some species show preference in prey (usually dino- and to revitalize the population after a bloom period,
flagellates, including some red tide-forming species; through the nuclear reorganization of the cell
Stoecker el al., 1981, 1984; Stoecker and Guillard, (Paranjape, 1980).
1982). On the other hand, they also constitute an
important food item, mainly for copepods, euphau-
siids, several crustacean larvae (Mysidacea, Methods
Decapoda, Penaeidea, Caridea), fish larvae, chaetog-
naths, dinoflagellates, foraminifers, radiolarians, Tintinnids have traditionally been analyzed on the
acantharians, ctenophores, benthic invertebrates, and basis of water-colurnn samples, the only exceptions
even some other tintinnid species (see Pierce and being a few studies on cysts found in Recent intertidal
Turner, 1992). and offshore marine sediments (Reid and John, 1981),
and some investigations on calpionellids, extinct late
Jurassic to early Cretaceous forms (Colom, 1948;
Tintinnoinea 325
Tappan and Loeblich Jr., 1968; Remane, 1969; 1971; Sediment trap techniques have undergone major
Echols and Fowler, 1973; Corliss, 1979). The most improvements in the last years, thus providing a useful
cornmon sampling techniques for tintinnids are indi- tool for the collection of various microplanktonic
cated below with brief cornments on their perfor- organisms (US GOFS, 1989; Lange and Boltovskoy,
mance (for further information on each type of gear, 1995; see chapter "Radiolaria Polycystina" in this vol-
see Boltovskoy, ed., 1981, and Paranjape, 1991). ume). Yearly abundance and specific composition
cyc1es are also best assessed by means of sediments
Plankton nets have long been the most widely used trap samples, which is of particular importance when
gear for qualitative studies. Improvements in their flux rates are compared with primary production fluc-
design, and the introduction of new ancillary devices tuations in the upper mixed layer (Honjo el al., 1982,
(opening-c1osing mechanisms, flowmeters, etc.) have 1988; Deuser el al., 1983, 1990; Wefer, 1989).
allowed their use in eco10gical studies requiring However, sediment traps also have important short-
assessments of abundance and horizontal and vertical comings for tintinnids: efficient sediment trap sam-
distribution pattems. However, as with several other pling is usually restricted to depths in excess of 500-
microzooplanktonic organisms, standard plankton nets 700 m (US GOFS, 1989; Lange and Boltovskoy,
often fail to yield adequate results because tintinnids 1995), and traps integrate the flux from several biolog-
are small in size, but unlike many comparably-sized ically dissimilar layers. Furthermore, although specif-
phytoplankters, are seldom abundant. Indeed, their ic relative compositions may not change significantly
small size requires the use of phytoplankton-sized due to destruction of the loricae in the process of sink-
meshes (around 20-30 J..llTl). However, these c10g fast, ing (Ling, 1992), overall destruction is very high: on1y
not only complicating the task of filtering enough ea. 1% of the 10ricae produced in the upper layers
water, but also making flowmeter measurements reach 800 m in a recognizable fashion (Boltovskoy el
lmpreclse. al., 1993a). Furthermore, the very significant qualita-
tive dissimilarities between microzooplanktonic
The use of suction pumps with the outlet driven upper-Iayer assemblages and those retrieved in 800-
through a net or another sieving device allows direct 2000 m sediment trap samples, presumab1y due to
control over c10gging even with very small-sized intermediate- and deep-water advection of cells pro-
meshes, very precise estimates of the volumes of duced elsewhere (Boltovskoy el al., 1996), also hin-
water filtered, and under static conditions, very high ders the use of these material s for ecologic and biogeo-
spatial resolution (since no towing is involved). graphic surveys.
Furthermore, damage due to turbulence and pressure
can also be controlled to a certain extent (although Once concentrated, samples can be preserved with 2%
detachment of the cell from the lorica due to mechan- buffered formaldehyde, acid Lugol's iodine solution,
ical disturbance is common). Because tintinnids are mercuric chloride with bromophenol blue (for fixation
usually concentrated in the upper water layers the use and staining), or a solution of 3% glutaraldehyde and
of pumps does not pose major operational problems. 1% osmium tetroxide. Laval (1971, 1972), Laval-
However, vertical profiles to depths in excess of 100- Peuto (1975), Hedin (1975), Gold and Morales (1976),
200 m are increasingly difficult (see Beers, 1981). Laval-Peuto el al. (1979), Gold (1979), Capriulo el al.
(1986) and Wasick and Mikolajczyk (1991, 1992) pro-
Bottle sampling provides another very useful tool for vide useful information on techniques for SEM and
tintinnid studies, especially when vertical profiles of TEM preparations aimed at the analysis of the ultra-
overall abundance and species-specific distributions structure of tintinnid cells and loricae.
are sought. The volume yielded by standard (Niskin,
Go-Flo) bottles (ca. 5-10 1) is usually too low for ade- Usually the thin cytoplasmic stalk (pedunc1e) attach-
quate sample-sizes, for which reason 6-3 bottles from ing the cell to its lorica (Fig. lA) can be easily broken
each level are required. Replicate samples are pooled either by manipulation during sampling (when using
and gently filtered through a 20 ~m mesh. Disturbances nets and pumps), or by the effects of the fixative.
are minimal with these method, which ensures the Moreover, the fixative can also cause subsequent dis-
suitability of the specimens for detailed morphological tortion (shrinkage or swelling) of the cell (Gilron and
and cytological studies. Lynn, 1989). As a consequence, most of the loricae
326 Tintinnoinea
are found empty in fixed samples, only very few Coastal waters. The genera Codonellopsis,
retaining the (distorted) cell inside. This may be a seri- Codonaria, Codonella, Stylicauda, and Tintinnopsis
ous problem not only for taxonomic studies, but also complex are restricted almost exclusively to coastal
for ecologic-biogeographic ones, since one cannot environments; they agglutinate (mainly arenaceous)
ensure whether empty loricae represent individuals particles on their lorica (see Taxonomy, Group 1).
still alive at the time of sampling, or were transported Although tintinnids with agglutinated loricae are
empty from elsewhere by currents. AIso, empty lori- present at virtually all latitudes in the South Atlantic,
cae prevent direct estimation of the cell's contribution from the tropics to the Antarctic, their abundance and
to total tintinnid biomass (Alder, 1990; see Verity and diversity are highest in transitional waters. In the
Langdon, 1984, for altemative methods). Thus, when- Bahía Blanca estuary (Argentina, ca. 39°S, 61 °W), for
ever possible potential biases resulting form the example, the Tintinnopsis group reaches 73% of all
above-described artifacts should be assessed on the specimens (Barría de Cao, 1992). Hyaline neritic tran-
basis of fresh, unpreserved samples (including obser- sitional tintinnids include the genera Favella (F.
vations, counts and measurements in vivo). taraikaensis), Helicostomella, Eutintinnus, Metacylis,
Amphorellopsis, Dictyocysta and Clevea?
Regardless of the collecting method, samples are con-
centrated before storage. Thus, when estimates of abso- Subantarctic waters. Characterized by Acantho-
lute abundance are sought, samples often need to be stomella, Steenstrupiella, Cymatocylis antarctica,
diluted again prior to counting under the inverted Eutintinnus rugosus and Codonellopsis pusilla, all of
microscope. It is strongly recornmended that absolute them inhabiting both neritic and oceanic areas.
abundance assessments be based on counts (Utermohl,
1858, technique) of several dilute counting chambers Antarctic waters. Characterized by the genera
rather than a single one with more concentrated materi- Laaclallanniella and Codonellopsis (see Biogeographic
als, where other particles, especially phytoplanktonic Zonation, Antarctic Oceanic below ) which, together
cells, may strongly interfere. Furthermore, counts of with Cymatocylis, are here the most abundant tintin-
several subsamples yield more precise results (Frontier, nids. It should be noted that the Tintinnopsis complex,
1981), and also permit estimation of counting errors. widely distributed elsewhere, is absent fram the
Southem Ocean.
Geographic and vertical distribution Some tintinnid species characteristic of a given area
may often be found outside its boundaries:
Geographic worldwide patterns Cymatocylis convallaria, forma ajJinis and
Laackmanniella naviculaefera, forma naviculaefera,
Scarcity of information and the overall state of disar- typically Antarctic taxa, have been recorded also in
ray of the classification of marine tintinnids (see subantarctic waters; transitional species, such as
be1ow) precludes a detailed analysis of their distribu- Dictyocysta elegans, were found in typically Antarctic
tion in the South Atlantic. A few broad generaliza- waters; the tropical species Ascampbeliella armilla,
tions, however, seem possible. Table 1 surnmarizes the Brandtiella palliata and Xystonellopsis paradoxa have
distribution ofthe 41 genera and 129 species encluded been recorded also in the Subtropical Convergence.
in this review (see Taxonomy). Most of the surveys This expatriation, attributable to advection by surface
available report only presence or absence. Only a few or deep currents and subsequent resurfacing, or to
supply quantitative species distributions (e.g., Barría entrainment in eddies, is very cornmon in tintinnids.
de Cao, 1992; Alder, 1995). The 4 broad distribution-
al pattems recognized are described as follows. As shown in Fig. 2, although North Atlantic tintinnids
have been intensively studied, large sectors of the
Tropical and sub tropical waters. Characterized by South Atlantic (especially the Central Gyre and the
the genera Rhabdonella, Xystonella, Xystonellopsis, subantarctic latitudes) still remain virtually unknOWll.
Epiplocylis, Climacocylis, Dadayiella, Epiplocyloides, Neritic zones have been more intensively surveyed in
Petalotricha, Cyttarocylis and Amphorides. These gen- the Southwestem Atlantic than off the African coast.
era may be present in both neritic and oceanic waters. According to Pierce and Turner (1993), the 53 tintinnid
Tintinnoinea 327
- \J
re .- \J
Tropical § 'E .'8
and :-e!S; l:
<Il _
Subtropical ~
•.•
~
::::l
l:
<:
1- V"l
--r.~
. .. .~
U U Q).
~ U + I -j
<1l::::l::::l_u_.;:; c·;:; c-;:; c
Uw 'ié
~ 'ié~ .-
U ->. <1l
w .~ ~
u u .~ ~.~
u -~ u ro
U
cLJ.JLJ.J N <1l::::l l· Q) Q) Q) Q)Q) Q)
-- <1l Z tlO Z U Z u Z u
::::l-E-Ea:i áiái O O O
ui50 Vl Z UCC[
,-- ~ ~
Aeanthostomella norv~giea, forma lata •• I
Aeanthostomella norvegiea, forma typiea • ~._ • 4
Albatrossiella? filigera • ~ _ ,
Amphorellopsis quinquealata _ • • !
Amphorides amphora ~. •• 1:
Amphorides ealida
laaekmanni :•
-+- +. ~ f •~
Amphorides
Amphorides
Amphorides
quadrilineata
q}!adrilineata,
quadrilineata,
varo
varo
brasiliensis
minor
_I~I_ I
i.
••
l. re:-
1 I
=- --+---t
-+ 1
-,
*Ampleetella
Ampleetella
Ampleetella
monoeollaria
nuda
trieollaria
...
••
i. -'-~_~
• l- _ ~
I
I
-+
-+ _.
~
+-+--j
Ampleetella tridivisa
*AseampJ2eliella acuta
•
I
I
I • l.
I I
*Aseampbeliella armilla ~~ _ +
Aseampbeliella obscura
Aseampbeliella ureeolata
Brandtiella palliata • I
l.l. • ~ ~
Clevea? melehersi I I l. •
Climacoeylis sealaria • t.
Climaeoeylis sealaroides + • • ,
Codonella amphorella
Codonella
Codonaria sp. aff. C. olla
*Codonella cistellula
aspera
Codonella galea
••
•
j •.- • +
•••
•
+
•
;• h·
I
•.
+
•
• ~ i+I
+
1 ~
+
~
Codonella? perforata ~ • • •
*Codonellopsis baleehi
Codonellopsis brasiliensis ¡..!..
1 +- r - + t +..-+ ~
328 Tintinnoinea
Codonellopsis gaussi, forma C'iJindroconica ••

Codonellopsis gaussi, forma globosa +---,-••
Codonellopsis gaussi, form~Jypica
CodonellQPsis lusitanica
CodonellQPsis morchella,
Codonellopsis morchella,
forma schabi
forma typica • •• • t--t --
Codonellopsis obesa • • --t +---+- t
Codonellopsis orthoceras
Codonellopsis pusilla
Cymatocylis antarctica J· . C1··1· ~.~.J.l-'·L·
· -
•
t1 ~. t -- ~ t--
Cymatocylis calyciformis
••
Cymatocylis convallaria
••
*Cymatoglis cristallina
Cymatocylis drygalskii, forma ovata •• t
Cymatocylis drygalskii, forma typica -¡¡l.,

• r
*Cymatocylis kerguelensis
•
Cymatocylis nobilis
••
Cymatocylis vanhoffeni
• • •·1 • •••• • •
•••
.1.,.
.1 T
Cyttarocylis cassis/plagiostoma
Oadayiella ganymedes
Oaturella datura
•• ·1·, • •:.¡••
• • • • ·f·
group , T T
Oaturella emarginata
*Oictyocysta californiensis
Oictyocysta duplex
Dict oc sta elegans
Dictyocysta elegans, varo lepída
Oictyocysta elegans, varo speciosa
Oictyocysta mitra
Epícancella nervosa
Epiplocylís acuminata
Tintinnoinea 329
- <.1
~ .- <.1
Tropical .~ ~ ''8
and
/
.0;:
ti)
-
t: :;
.•...
Subtropical ~ ~ ~
~ ~
~ r
U. U, Q)
•...U
•
U'iO'iOu >-<tIU u u Uu u
~:J :J_LJ_o,;:¡ c"Z c'';::::; C
Q) ~ ~ .- Q).- <ti .- <tI.- <ti
cWw N <ti=:¡ Q¡ Q) Q¡ Q)Q¡ Q)
.- <ti Z 00 Z u Z u Z u
=:¡...c...ccOcc
O":J't: Q)Q)
O O O
O O
Vl Z U c:o
*Epiploc;j!jis calyx ~ +•
*Epiplocyloides?
Epiplocylis undellaantarctica
Epiplocyloides? reticulata
Eutintinnus apertus
~•.
+ \ .~.
~• + • +
~ + +
t---.:;;.f..
••
•
l.~.
+
+
~ ~
+
+- + •
~
Eutintinnus colligatus + + + •
Eutintinnus fraknoi +
• +
• • • t- +- +
i
t- t ~ +
Eutintinnus medius
Eutintinnus
*Favella lusus-undae
arcuata
--+
• •
¡
• -+--+.
• +
•
~
t.. t..
~• 1
I•
+
j l
*Eutintinnus rugosus + I • +.
•
Favella ehrenbergii • ¡ + + +- +
*Favella serrata +
• + + •
Favella taraikaensis + + •.
• •
Helicostomella subulata + -~
• +
•
Laackmanniella naviculaefera, forma naviculaefera •. + ...,...
• •
Laackmanniella naviculaefera, forma prolongata ~ + • •
Metacyli00rgenseni
Ormosella sp.
Metacylis acantharus
affo Mo mereschkowskyi
Ormosella cornucopia
Ormosella haeckeli
-----+-
----+
t
•
t
•
e ~
~
__..
-~
+
¡.
+.
+
+•
I
+. . .
••
Parundella caudata -- +
I
•.
• •
Parundella messinensis • • .. ..
Petalotricha ampulla • •• +•
Poroecus apiculatussimplex
Protorhabdonella?
Protorhabdonella?
*Rhabdonella amor
curta ·f. • t----
--
• ••
t~·
.. ~ ..I •
+I •
+
•••• t. •
+•
+
f
+
+
*Rhabdonella amor, varo cuspidata •• L

330 Tintinnoinea
*Rhabdonella amor, varo indica

*Rhabdonella amor, varo valdestriata
*Rhabdonella apC2Phxsata
*Rhabdonella chiliensis
*Rhabdonella cornucopia
Rhabdonella spiralis
Rhabdonella spiralis¡ varo henseni
Salpingella acuminata
Salpingella acuminata¡ varo glockentogeri
Salpingella acuminata¡ varo secata
Salpingella acuminata¡ varo undata
Salpingella acuminatoides
Salpingella costata
Salpingella regulata
Salpingella subconica
*Steenstrupiella pozzi
Steenstrupiella steenstrupi i
Stelidiella stelidium
Stenosemella avellana
Stylicauda platensis
Tintinnidium balechi
Tintinnidium spo affo T. semiciliatum
"Tintinnopsis" a'!!Phistoma
"Tintinnopsis" amphora/nucula
"Tintinnopsis" aperta
"Tintinnopsis" baltica
"Tintinnopsis" beroidea
"Tintinnopsis" campanula/butschlii¡ varo butschlii
"Tintinnopsis" campanula/butschlii¡ varo mortensenii
"Tintinnopsis" c0'!!Pressa
Tintinnoinea 331
- \,J
~ .- \,J
Tropical
and
.2
."=
~
-
.'8
'" c: _;¡¡
Subtropical
.=
~.s ::::l
<Jl
~
U U (\)
Urorou
•... ...: roU
~u u u u u
u e·;:;
ro::::l ::::l_lJ_.;:;
(\) e- e- .- - (\) .- e·;:;
ro ro·-
e
ro
.- ro •..• 00 Z u Z Ou Z Ou
lelJ
::::l-5-5cls
UJ UJ N
ee
~ ::::ll(¡¡
O
(\) (¡¡ (\) (¡¡ (\)
::::l
o
Vl
~
o
z U
(\) (\)
cc
"Tintinnopsis" cratera e+-,

"Tintinnopsis" fimbriata
"Tintinnopsis" kar~censis
~+'~~~.,
"Tintinnopsis" kofoidi _
"Tintinnopsis" /obiancoi/brasi/iensis ~,- 'Fe- e
-+-
e f,te
+-t =-lj ::
"Tintinnopsis" nordqvisti
"Tintinnopsis" parva ¡
"Tintinnopsis" prowazeki e
"Tintinnopsis" pusil/a, forma /aevis ~r-· ~e
"Tintinnopsis" radix
"Tintinnopsis" riop/atensis .I! ,e"~ -
"Tintinnop~is" tocantinensis • ~·~t ~
e
"Tintinnopsis" uruguayensis ~-. - +
e
Undella cléparedei grou¡::>
*Undella grandis le~le,:t

L + -+e t • ~~ ~ t + +
Undella hemisphaerica e- tI ~
Undella h)lél/ina te~~e e e
Undellopsis marsupia/is
Xystonella /ongicauda
e+ e e
L e_ele e
, , , ~
Xystonella treforti, forma typica n. formo
etete.e e I
Xystonellopsis cymatica etete
Xystonellopsis favata
Xystonellopsis hastata
Xystonellopsis heros
1:;-'(
e e e e
T t •
II
Xystonellopsis ornata e~ete
Xystonellopsis paradoxa el e
Xystonellopsis pu/chra el e
Xystonellopsis? sp. I e
Table 1. General distributional data for the tintinnid species treated. Dubious genera are denoted with a question mark (?), dubious species with an
asterisk (*).
332 Tintinnoinea
Fig.2. Worldwide distribution oftintinnid studies compared with those for the South Atlantic. Partly based on Pierce and Tumer (1993).
genera recognized in the W orld Ocean can be grouped Geographic patterns in the Southwestern Atlantic
into six overall distribution pattems: warm water
(43.4%); cosmopolitan (30.2%),; neritic (13.2%); Absolute abundances of 40 tintinnid species along a
boreal and austral (3.8% each); and endemic tropical transect of surface oceanic waters in subtropical to
Pacific (5.6%). The authors point out that the tropical Antarctic latitudes of the southwestem Atlantic (30oS
Pacific differs from other oceans in that it hosts three to 60oS, along 55°W; Alder et al., unpublished) sug-
endemic genera: Amplectellopsis, Cricundella and gest four biogeographic zones (Fig. 3) elearly asso-
Codollopsis. However, the former two were synonym- ciated with hydrology, but with seasonally and multi-
ized with Amplectella (Fig. 6.96-99) by Balech (1975), annually variable limits:
a genus well-represented in deep-sea layers of the
Atlantic; while the systematic position of Codonopsis Subtropical Oceanic (Zone 1, north of 300S). These
is dubious, and it ineludes the single species C. ollllla assemblages are strongly dominated (up to 46%) by
(Fig. 3B), originally described by Brandt (1906-1907) Elltintinmls medil/s.
in the genus Cyttarocylis and very rarely found there-
after. Thus, with the exception of the Indian Ocean, the Transitional Oceanic (Zone 2). The largest of all four
largest oceanic areas ofthe world host practically iden- biogeographic zones, encompassing 3 subzones: a
tical generic assemblages. Available specific informa- northem one extending from 34 to 37°S (Subzone 2A);
tion is scarce and contradictory, and does not permit an intermediate one between 37-43°S (Subzone 2B);
precise identification of endemic species. and a southem one located between 45 and 47°S
(Subzone 2e). Species assemblages of the former two
are similar, dominated by the genus Dictyocysta (D.
mitra, D. elegans var. speciosa, D. elegans varolepida),
Tintinnoinea 333
,
Subtrqpical
Ocefnic
30°
...................wi~·;;;·······
o"
•., 35°
.
•••• (lOOOOOOOOOOOO Surnn1er
/
/:: ••••• 2A •••••••••••••••••
..
: 40°
Transitional
.
/28 • Oceanic
f •
\ ~ ... 2C
45°
~/
¡t
. ~ ...
....
....~.:::::::.:::::::.::::::::.:::: :::.::::
Subantarctic
'. .•••·
.' . . 50°
.
3 Oceamc •••.
~1'.. ..
j
~'"o.'-~"
-....--
.
{c\\c\.\((
.'
- \3'
00,,0
•••<............
.'.
Anta«t"
.
o
55°
,,~\3 . e O e~\ •••••

• 4 Oceamc
o"''' ",\
•
gO" ~(o
OOIlOYoO\-;).(
60°
I
70° 60° 50° 40°
Fig. 3. Biogeographic zonation of the southwestem Atlantic based on the absolute abundances of tintinnid specific assemblages.
and the species Epiplocylis aCllminata, Steenstrupiella several hypotheses were offered to explain its disjunct
steenstntpii, Codonellopsis pllsilla and Protorhabdo- distribution (see Pierce and Turner, 1993). However,
nella cllrta. Subzone 2B differs from 2A by the pres- our data suggest a different distribution for this taxon.
ence of AcanthostOlnella Ilorvegica, a typically Arctic A. norvegica is present in the Arctic Sea, but was never
species (Davis, 1985, Boltovskoy et al., 1991), which in found in typically Antarctic waters south of the Polar
surface and subsurface waters occurs only between the Front (=Antarctic Convergence), not even at layers as
Antarctic Convergence (around 56° in this study) and deep as 1000 m (Alder, 1995). In subantarctic lati-
37°S. Subzone 2C defines the boundary between transi- tudes, as already mentioned, A. norvegica is frequent
tional and subantarctic waters; abundant species in sub- within the oceanic domain, but only rarely found in
zones 2A and 2B are scarce here, whereas Protor- neritic waters (see Souto, 1979). Acanthostomella lata
habdollella simplex becomes the sole dominant species. Kofoid and Campbell, 1929, which has been reported
from tropical and subtropical waters (Kofoid and
Acanthostomella norvegica (Fig. 6.75) was never Campbell, 1929: Tropical Pacific; Balech, 1968:
reported (under this name) from tropical-subtropical northem Tropical Atlantic; Souto, 1979: southem
waters. This species is usually considered bipolar, and Tropical Atlantic), is herein considered a form of A.
334 Tintinnoinea
florvegica, differences between the two taxa being centrations so far reported are in the Kiel Fjords: 215
extremely minor, involving only total length and the million ind. 1-1 (Kils, 1993). This extreme value is
degree of development of aboral horns (both traits attributed mainly to Stenosemella nucula, whieh was
without diagnostic value). Their distributional pat- found to form miero-Iayers and micro-patches up to
terns, on the other hand, are different and well-defined: 340 mm thick (Kils, 1989, 1993).
A. norvegica, forma lata (Fig. 6.75d) is restrieted to
warm waters, while A. norvegica, forma typica (Fig. In oeeanie waters off the Argentine shelf (30-600S,
6.75a-e) is a typically cold-temperate organismo 55°W) eoneentrations hardly exeeed 20 ind. 1-1 (Alder
et al., unpublished). Low oceanic tintinnid abundances
Subantarctic Oceanic (Zone 3). Extending from ea. were also registered in the California Current (Beers
49 to 55-56°S, it is characterized both by the presenee and Stewart, 1967) and in the North Paeific Central
of Acanthostomella norvegica and the dominanee of Gyre (Beers et al., 1975), where mean values reach only
Cymatocylis antarctica. The latter is virtually the only 19 and 5 ind. 1-1, respectively. South of the Antaretic
species of Cymatocylis (a typically Antarctic genus) Convergence den sities can be considerably higher: in
whose distribution follows the Circumpolar Antaretic neritic Antaretie waters of the Southern Weddell Sea
Current (Alder, 1995), although it is al so frequently peak tintinnid eoncentrations are approx. 1200 ind. 1-1
found in subantaretic waters. (Boltovskoy et al., 1989; Boltovskoy and Alder, 1992).
High-latitude oceanic waters influenced by sea-ice are a
Antarctic Oceanic (Zone 4). Typical speeies of this unique habitat whieh supports diverse and abundant
zone are Codonellopsis gaussi (with its formae typica, eommunities (Garrison et al., 1983, 1986; Garrison and
globosa and cylindroconica), and Laaclallanniella Buck, 1989a, b; Garrison, 1991a, b; Garrison and
naviculaefera, forma naviculaefera. These are Gowing, 1993; Garrison and Close, 1993). Thus, prob-
extremely polymorphic taxa whose loricae can be ably as a response to enhanced food availability, tintin-
either hyaline or covered to a varying extent with for- nids here are more abundant than in extrapolar oceanie
eign particles (mainly diatoms). The degre~ of agglu- regions. The Weddell-Scotia Confluence (57-62°S,
tination of the lorieae st'ems not to be affected by dis- 49°W), for instanee, hosts coneentrations around 250
tanee from the coast, but is inversely related to ind. 1-1 (Alder and Boltovskoy, 1993; Alder, 1995).
ice-eoverage (Alder, 1995).
Vertical distribution
Comparison of plankton collections from the
Southwestern Atlantie with sediment trap material s Literature on vertical distribution of Atlantic tintin-
from the eastern Tropical Atlantic (unpublished data) nids is less extensive than on geographie distribution,
indieates that tintinnid species assemblages are rough- largely restricted to short remarks on records of vari-
Iy similar on both sides of the ocean. ous taxa in isolated vertical easts. Baleeh (1972) clas-
sified subsurfaee species vaguely into three groups
Abundance patterns (species with dubious depth-preferenees were includ-
ed in a fourth group, omitted herein) aeeording to their
The few data available suggest signifieantly different inereasing degree of preference for deep ("shaded")
tintinnid concentrations between neritie and oceanic waters, as follows:
surface waters. In coastal environments such as the
Bahía Blanca estuary, tintinnids may reach densities Shallow: Ascampbeliella, Codonella acuta, C. aspe-
up to 11,300 ind. 1-1 (Barría de Cao, 1992). Sueh high ra, Xystonella treforti, and others.
densities are comparable to some reported for neritie Intermediate: probably all speeies of Salpingella,
waters of other regions: Narragansett Bay, 14,000 ind. Undella hyalina, Codonaria cistellula, and others.
1-1 (Hargraves, 1981); Long Island Sound, 13,000 ind. Deep: probably all species of Amplectella, Brandtiella
1-1 (Capriulo and Carpenter, 1983); estuary of the palliata, Ormosella, Parundella, Undellopsis, most
Tamar river, SW England, 16,000 ind. 1-1 (Dale and speeies of Xystonellopsis, and others.
Burkill, 1982); Maine Estuary, 7,000 ind. 1-1 (Sanders,
1987); Northern eoast of Denmark, 15,000 ind. 1-1 Quantitative vertical profiles for the extrapolar South
(Cordeiro, 1995). Perhaps the highest tintinnid con- Atlantie are restrieted to subsurface (5-15 m) oeeanic
Tintinnoinea 335
300 00
200 eB1. A frica
Atlantic
Ocean
00 • GBN3
55,239
12,160 -CB1
201,064 3921
8537017
7074
GBN3
GBN3
- 17,350
37,395
27,275
39,434
1,363 m- O
- -24,646
O
125,036
14,632 -77,608 121,971 - 66,712
Min.
Fig. 4. Geographic locations of time-series sediment trap deployments in the eastern equatorial Atlantic, and the corresponding flux rates of
tíntinnids (bold numbers) and ciliate cysts (second figure. not boldened), in ind. m-2 day-l Data from Boltoyskoy el al. (1993a, 1996), and
Uliana (1997).
waters ranging from subtropical to Antarctic latitudes Although the abundance of some species can undergo
(Alder, unpublished). Although of limited value for very significant seasonal fluctuations, often resulting
comprehensive analyses, these data confirm that in shifting winter-surnmer dominance patterns (e.g.
Balech's "true shade species" ("Deep") are indeed off Hope Bay; Alder, 1995), vertical distribution does
totally absent fram the upper 15 m. not seem to be affected by such changes.
AlI available infonnation on the vertical distribution- Several studies on tintinnids in sediment trap samples
al of tintinnid species in the Atlantic sector of the have appeared recently (e.g., Bathmann el al., 1987,
Southem Ocean was surnmarized by Boltovskoy and 1990; Dale, 1989; Nothig and Von Bodungen, 1989;
Alder (1992) and Alder (1995). Based on detailed Ling, 1992; Boltovskoy et al., 1993a, 1996).
quantitative profiles of discrete depth intervals Boltovskoy el al. (1993a, 1996) and Uliana (1997) ana-
between Oand 1000 m, Alder (op. cit.) concluded that lyzed material s from various depths in two nearby,
not all species are restricted to preferential depths. highly productive sites in the eastem tropical Atlantic,
Species with restricted depth ranges, however, reporting flux rates in excess of 200,000 ind. m-2 day-I
decrease in abundance with depth such that deeper- (Fig. 4). Based on indirect comparisons with usual tin-
living species are scarcer than shallower ones. Depth tinnid concentrations in productive surface and subsur-
preferences were c1assified into 3 groups: face tropical waters (around 100.000 ind. m-3), over
99% of the loricae produced at these sites appear to be
O-50 m: Laad71/anniella naviculaejera and Codonel- lost to sedimentation before reaching 800-2000 m. On
lopsis gaussi; the other hand, proportions of deteriorated loricae
40-70 m: Cymatocylis convallaria and Cymatocylis increase only slightly with depth below ca. 800 m, sug-
dl}'galskii; gesting that destructive processes are of minor impor-
80-250 m: Cymatocy/is vanhoffeni. tance at mesopelagic depths. As opposed to overall
336 Tintinnoinea
abundance, species relationships seem to change little Brandt's and Laackmann's studies is lumping species
with depth (Ling, 1992), indicating that susceptibility of very dissimilar morphology in the same genus.
to destruction by grazing and bacterial activity is simi-
lar for the different species. In an attempt to correct this oversimplification,
Kofoid and Campbell (1929, 1939) erected a very
large number of new genera and species, thus falling
Taxonorny into the opposite error. The great majority of these
new taxa (especially the species) are based on minor
Taxonomic studies of South Atlantic tintinnids include and most probably intraspecifically variable morpho-
a few that covered large areas (Brandt, 1906-1907; logical details. This classification erroneously sug-
Laackmann, 1910; Souto, 1979, 1981; Alder, 1995), gests that tintinnid loricae exhibit sharp morphologic
and several focusing on geographically restricted, usu- discontinuities, thus contradicting the increasing body
ally coastal, areas (Da Cunha and Da Fonseéa, 1917; of evidence pointing to smooth transitions between
Faria and Da Cunha, 1917; Balech 1942, 1944, 1945, dissimilar . morphotypes. Because Kofoid and
1947,1948, 1951, 1958~b, 1962a, 1971a, 1973, 1976; Campbell's work has long been a primary reference
Souto, 1970a, b, 1972, 1974; Barría de Cao, 1981, for subsequent studies, subsequent literature is
1986, 1992). Brandt (1906-1907) and Souto (1979, plagued with spurious species names.
1981) provided comprehensive information on tintin-
nid species in tropical and temperate waters, while Although later studies on South Atlantic tintinnids
studies by Laackmann (1906, 1907, 1910) and Alder (Balech, 1942, 1945, 1948, 1951, 1958a, b, 1971a;
(1995) dealt mainly with Antarctic species. Souto, 1979, 1981) placed some of Kofoid and
Laackmann (1910) also described the fauna off South Campbell's species into synonymy, they followed the
Africa frorn the "Gauss" expedition. Finally, Kofoid general guidelines of these authors without major
and Campbell (1929, 1939) described tintinnids from changes (mainly regarding the arrangement of species
the eastern Pacific Ocean (North Temperate, Tropical, into families and genera). Alder's (1995) work on
and South Temperate Pacific, from the Arctic Circle to Antarctic tintinnids also followed the overall se heme
the Tropic of Capricom), re-arranging all genera, spe- proposed by Kofoid and Campbell but, having noticed
cies, forms, and varieties chiefiy from Brandt's (1906, extremely high intraspecific variability, proposed a
1907) and Laackmann's (1910) monographs. Kofoid drastic reduction in the number of species from 70 to
and Campbell also introduced the concept of "series" 11, and invalidated several genera.
within some genera in order to group species of similar
morphology, and indiscriminately erected hundreds of The neglect of intraspecific variability may be attribut-
new species on the basis of previous subspecific cate- able to the scarcity of surveys based on abundant mate-
gories. Besides Kofoid and Campbell's (1929, 1939), rials from geographically large areas. Indeed, most pre-
the only other comprehensive studies of tintinnids are vious t~xonomic studies have covered on1y restricted,
those by Daday (1887b) and Brandt (1906, 1907). geographically iso1ated locations aml/or short time
scales. Such spatial and temporal sample discontinu-
Two very different criteria are currently applied to the ities, often combined with low sample retrieval aml/or
systematics ofTintinnoinea. One ofthem uses the ratio examination, prevented researchers from tracing subtle
of lorica total length to oral diameter as the chief taxo- morphological transitions between the variability
nomic character, and assigns the same diagnostic value extremes within one species. Thus, closely spaced sam-
to all other size parameters and features of the lorica. pIes of neritic to oceanic waters at different times of
This was Kofoid and Campbell's (1929,1939) basis for year show that a single species may encompass several
describing over 1000 species. As opposed to this rather highly dissimilar morphotypes (e.g., Boltovskoy et al.,
simplistic approach, Brandt (1906-1907), Laackmann 1989; Alder and Boltovskoy, 1991a, b; Alder 1995), as
(1910), Alder (1995), among others, stressed the high shown in Fig. 5. These results strongly support
intraspecific polymorphism of tintinnids, recognizing Brandt' s (1906, 1907) emphasis on infraspecific ranks
far fewer species and proposing the use of various like "Formenkreis" and variety. Laackmann (1910),
infraspecific categories, including "Formenkreis", who also had very ample collections at his disposal,
form, and variety. However, a major problem of closely followed the criteria proposed by Brandt.
Tintinnoinea 337
Codonellopsis gaussi
1
i1 ~
P
F;.~~DIJ
L..:=:1
n
g
7
':¡¡C::';;i'
"',1'1'
__ '\
t !
,iff..t,).~.hV
" 11 r---
12
rl0' ')\' I g
cy
11
l~,
cy ~
t, 0"\~
~.~
~4 .1 g I !
'"V"
cy
\J31
Q, e29 0" t-i
e~3
cy cy g r
_._.J
g g g
cy g -
V,,~Av" ~
~ 1\ '-
',- - "'-1
r..-'-.J
, ....
l1 t~#~9)1 \ / re)
/
\ Q
1:=·_-
\ \
I ; • i
~H
\"
" (
,;, ;: ~,<:t¡3
~
¡TI I
( \
1
\ 21 \/16 VIS
\/18 'J 17
\/20
Fig.5. Morphological varialions in Codollellopsis gOl/ssi. Modified from Alder (1995). t: Codollellopsis gOl/ssi, forma t;pico; g:
Codollellopsis gOl/ssi, forma globoso; cy: Codollellopsis gOl/ssi, forma cylilldrocollico; e: Codollellopsis gOl/ssi, fomla coxliello. Figure sour-
ees: from Baleeh (1973): 2, 3-5,11-18,20,22,24,27,33,34; from Baleeh (1947): 8; from Sassi and Melo (1986): 21,30-32; from Hada (l970):
10,19; frol11 Laaekmann (1910): 1,6,9,23,25,26,29; frol11 Laaekmann (1907): 7,28.
The methodology for classifying tintinnid species ticles, were recently found to be highly variable at the
adopted in the present chapter does not differ fram that intraspecific level (Laval-Peuto and Brownlee, 1986;
applied by most specialists, based largely on attributes Alder, 1995). In cases where the structure of the lori-
of the lorica. Its general morphology, presence of a ca is obscured by a dense agglomeration of particles
collar, structure of the oral end, presence and type of (e.g., Tintinnopsis complex; see Identification of the
striae and teeth, number of spiral turns and fenestrae, Taxa); cytological and/or ultrastructural analyses are
and ability to agglutinate particles are the most useful recommended. Most of the morphometric parameters
features for differentiating species. Other commonly regularly used for taxonomic purposes (total length,
used attributes, such as presence of an aboral horn or collar length, bowl length. length of aboral horn and
pedicel, and the type and density of agglutinated par- pedicel, external oral diameter, maximum, minimum
338 Tintinnoinea
and middle diameter of bowl, aboral diameter) were available for a handful of tintinnid species, adopted a
also found to be highly variable intraspecifically. In slightly modified version of Kofoid and Campbell's
agreement with several previous observations (e.g. (1929, 1939) system, which includes 13 families:
Balech, 1959, Souto, 1979, Laval-Peuto, 1981, Laval-
Peuto and Bronwlee, 1986; Alder, 1995), the internal Class Oligotrichea Bütschli, 1887
oral diarneter is considered the only trait with suffi- Order Tintinnida Kofoid and Campbell, 1929
cient intraspecific stability to be useful for taxonomic Family Tintinnidiidae Kofoid and Carnpbell, 1929
identifications. At any rate, because of the very high Genera Tilltill11idium Kent, 1881; Leprotilltin-
specific and supraspecific morphological variability IllIS Jorgensen, 1900
of loricae, it is generally acknowledged that cytologi- Family Codonellidae Kent, 1881
cal studies are needed for a sound c1assification Genera Codollella Haeckel, 1973; Codollaria
system. Kofoid and Campbell, 1939; Codollopsis
Kofoid and Campbell, 1939; Tintblllopsis
Despite efforts to sort out the maze of conflicting Stein, 1867; Poroecus Cleve, 1902
names and morphotypes, the system presented herein Family Codonellopsidae Kofoid and Campbell,
contains undeniable ambiguities, especially where 1929
subtle morphological gradations between species and Subfamily Codonellopsinae Campbell, 1954
even genera are concerned. Keeping these taxa separ- Genera Codollellopsis Jorgensen, 1924;
ate or lumping them into a single taxon is, at this Laackmalllliella Kofoid and Campbell, 1929
point, largely a question of preference. Whenever pos- Subfamily Stenosemellinae Campbell, 1954
sible these details are stressed in order to call the Genera Stellosemella Jorgensen, 1924
reader's attention. Family Dictyocystidae Kent, 1881
Genera Dictyocysta Ehrenberg, 1854
Outline classification Family Metacylididae Kofoid and Campbell, 1929
Genera Metacylis Jorgensen, 1924; Pseudo-
The c1assification system of the subkingdom Protozoa metacylis Balech, 1968; Helicostomella
(kingdom Protista) adopted in this work is that of Jorgensen, 1924; Clilllacocylis Jorgensen, 1924
Levine el al. (1980), in which tintinnids (Tintinnoinea) Family Ptychocylididae Kofoid and Campbell,
are considered a suborder of the phylum Ciliophora: 1929
Genera Ptyclrocylis Brandt, 1896; CYlllatocylis
Phylum Ciliophora Doflein, 1901 Laackmann, 1910; ProtocYlllatocylis Kofoid
Class Polyhymenophorea Jankowski, 1967 and Campbell, 1929; Favella Jorgensen, 1924.
Subclass Spirotrichia Bütschli, 1889 Incertae sedis: Wailesia Kofoid and Campbell,
Order Heterotrichida Stein, 1859 1939
Order Odontostomatida Sawaya, 1940 Family Epiplocylididae Kofoid and Campbell,
Order Oligotrichida Bütschli, 1887 1939
Suborder Oligotrichina Bütschli, 1887 Genera Epiplocylis Jorgensen, 1924; Epiplo-
Suborder Tintinnoinea Kofoid And Campbell, cyloides Hada, 1938; Epicallcella Kofoid and
1929 Campbell, 1929
Family Ascampbelliellidae Corliss, 1960
In Kofoid and Campbell's (1929) work, the suborder Genera Ascalllpbelliella Corliss, 1960;
Tintinnoinea was divided into 12 families, 8 subfami- Acalltlrostolllella Jorgensen, 1927. Incertae
lies, 51 genera, 8 subgenera and over 1,000 species. In sedis: Niemarshallia Corliss, 1960; Luxiella
1939, the same authors expanded the number of fami- Lecal, 1953
lies and genera to 13 and 62, respectively, and reduced Family Cyttarocylididae Kofoid and Campbell,
the number of species to 726. This last revision appears 1929
more consistent, and each taxon is described in a more Genera Cyttarocylis Fol, 1881; Petalotricl,a
exhaustive and precise manner. In her recent review of Kent, 1881
the group, Laval-Peuto (1994), based on subsequent Farnily Rhabdonellidae Kofoid and Campbell,
taxonomic revisions and on the cytological information 1929
Tintinnoinea 339
Genera Rlzabdollella Brandt, 1906; techniques for which were unavailab1e at the time of
Rlzabdollellopsis Kofoid and Campbell, 1929; Kofoid and Campbell (and which so far are partially
Protorhabdonella Jorgensen, 1924; known for only ea. 30 species; cf. Laval-Peuto, 1994).
Epirlzabdollella Kofoid and Campbell, 1939 In the present work we therefore preferred to avoid
Family Xystonellidae Kofoid and Campbell, 1929 using supragenenc categories, stressing generic and
Genera Parlllldella Jorgensen, 1924; specific diversity instead, with special emphasis on
Xystollella Brandt, 1906; Xystollellopsis intraspecific variability which has been shown to
Jorgensen, 1924; Parafavella Kofoid and poses s strong ecological correlates (e.g., Alder, 1995).
Campbell, 1924 The tintinnid species covered are assembled into mor-
Family Undellidae Kofoid and Campbell, 1929 phologically similar groups; however, beeause eriteria
Genera Undella Daday, 1887; Undellopsis for establishing the latter are generally similar to those
Kofoid and Campbell, 1929; A11lplectella used for traditional higher-Ievel systematies, these
Kofoid and Campbell, 1929 groups are closely comparable with the family-Ievel
Family Tintinnidae Claus, 1876 classification outlined above.
Subfami1y Salpingellinae Kofoid and
Campbell, 1939 Identification of the taxa
Genera ElItintilllllls Kofoid and Campbell,
1939; Clevea Balech, 1948; Salpingella The present chapter covers all tintinnnid taxa record-
Jorgensen, 1924; Salpingelloides Campbell, ed so far from the South Atlantic; 41 genera, 129 spe-
1942; Salpingacantlza Kofoid and Campbell, cies, 8 varieties, and 37 formae. These numbers are
1929; Rlzabdosella Kofoid and Campbell, the resu1t of combining ancl/or invalidating many pre-
1929; Epicranella Kofoid and Campbell, viously described species and genera. The revision is
1929; Daturella Kofoid and Campbell, 1929 based on published data and on published and unpub-
Subfamily Tintinninae Kofoid and Campbell, lished personal results from the Southwestem Atlantic
1939 and the Southem Ocean. By necessity they rely to a
Genera Tintinnlls Schrank, 1803; BlIrsaopsis large extent on descriptions and illustrations from the
Kofoid and Campbell, 1929; Cantlzariella literature (some of these taxa were not observed by
Kofoid and Campbell, 1929; Steenstrupiella the author). IJIustrations are considered of special
Kofoid and Campbell, 1929; Amplzorides importance because they summarize practieally all the
Strand, 1926; Odontoplzorella Kofoid and traits used in tintinnid classifieation. For ease of com-
Campbell, 1929; Amphorellopsis Kofoid and parison, a uniform magnification is used for all fig-
Campbell, 1929; Albatrossiella Kofoid and ures within a genus. Ample use has be en made of fig-
Campbell, 1929; Dadayiella Kofoid and ures in Brandt (1906), Laackmann (1910), FarÍa and
Campbell, 1929; BlIsclziella Corliss, 1960. Da Cunha (1917), Balech (1944, 1945, 1948, 1951,
Incertae sedis: PrOa11lpllOrella Kofoid and 1971a, b, 1975), Souto (1979), Laval-Peuto (1981)
Campbell,1939 and Barría de Cao ( 1981, 1986), beeause most of them
Subfami1y Stelidiellinae Kofoid and Campbell, aceompany original descriptions.
1929
Genera Stelidiella Kofoid and Campbell, Most ofthe genera were erected by Jorgensen (1924)
1939; Or11losella Kofoid and Campbell, 1929; and by Kofoid and Campbell (1929, 1939). Because,
Brandtiella Kofoid and Campbell, 1929 as discussed above, polymorphism is common in most
Incertae sedis: Prostelidiella Kofoid and tintinnid species, several morphotypes of the same
Campbell, 1929 species are often included. Short diagnoses for the
genera and many of the species are provided; howev-
A1though this 1atter system is the most widely used er, where illustrations clearly depict all taxonomically
today, it clearly is far from satisfactory. Because it is important traits, diagnoses are omitted. Combined
based on lorical characters only, it cannot be com- names (e.g., "Tintinnopsis" ampllOra/nuclIla) denote
pared to the classification ofthe rest ofthe ciliates. As cases where analysis of the whole spectrum of formae
noticed before, a sol id classification should take into did not reveal limits between the "species" involved.
aecount cyto10gieal and ultrastructural evidence, the Dubious genera and species are denoted with a ques-
340 "'
Tintinnoinea
tion mark and an asterisk, respectively. The designa- 2a Rounded or flat aboral end, always closed 3
tion "group" denotes categories of unclear taxonomic 3 Bowl separated fram the collar by a nuchal con-
affinities, where probably several related taxa are striction (Fig. 6.1 d) .4
lumped. For each of species one or more references 3a Lorica without a differentiated collar.. 5
("Ref.") providing relevant descriptions and/or illus- 4 Oral end without a suboral cone (Fig. 6.7a), or
trations are included. Notice, however, that the names with a smooth one: Codonella (Fig. 6.1-6)
in the literature are not necessarily those utilized here- 4a Oral end with a well-developed suboral cone:
in. For this reason, and because the system used here- Codonaria (Fig. 6.7)
in involves major changes from previous literature, a 5 Bowl without an aboral horn: Tintinnidium
listing with the corresponding equivalences for the (Fig. 6.8-9)
morphotypes treated is provided. 5a Bowl with an aboral horn 6
6 Lorica wall without a visible spiral structure:
Two alternatives are offered as a first approach to the Poroecus (Fig. 6.10)
identification of the tintinnids of the South Atlantic: a 6a Lorica wall with a spiral structure: Stylicauda?
traditional key to generic level, and diagnoses of (Fig. 6.11)
groups and subgroups of morphologically similar 7 Bowl generally with agglutinated particles 8
forms. 1t should be born in mind, however, that both 7a Lorica wall hyaline 14
systems are artificial, insofar as they do not reflect 8 Collar spiralled 9
phylogenetic relationships. 8a Collar absent or non spiralled 11
9 Aboral end cornmonly open: Laackmanniella
Key to the genera of Tintinnoinea recorded in the (Fig. 6.31)
South Atlantic 9a Abora1 end always closed 10
10 Collar short, with only two or three spiral turns:
While identification keys may be useful for rapidly Stenosemella (Fig. 6.42)
pinpointing a given morphotype, extreme care should lOa Collar higher, with more than three turns:
be taken when using this tool. On one hand, geograph- Codonellopsis (Fig. 6.35-39)
ically restricted identification keys become rapidly 11 Lorica without fenestrae; wall covered by atypi-
less applicable as new species are recorded in the area. cal gelatinous external coating: Brandtiella (Fig.
On the other hand, keys by nature must be rigid in 6.49)
their treatment of the morphological traits, allowing 11a Lorica with fenestrae 12
little room for ambiguities. Thus, as long as taxonom- 12 Fenestrae small and inconspicuous, arranged in
ic ambiguities and transitional morphotypes remain few rows on the precollar and in middle of bowl:
exceptional, keys may be an adequate identification Petalotricha (Fig. 6.47)
tool. However, as noted above, transitional features 12a Fenestrae easily visible 13
are the rule for tintinnids, thus seriously restricting the 13 Few, large, quadrilateral fenestrae arranged in
utility of keys. The reader should therefore keep in one or two rows on suboral zone: Stelidiella
mind that the inevitably rather sharp cutoff traits in (Fig.6.50)
the following identification key may not be faithfully 13a Many fenestrae located on both bowl and collar:
represented in actual samples. Dictyocysta (Fig. 6.43-46)
14 Collar spiralled 15
Lorica wall with freely and/or evenly agglutinat- 14a Collar absent or non-spiralled 16
ed particles in highly variable densities, often 15 Collar with a few spiral turns; lorica short and
obscuring the underlying structure; collar absent, wide: Metacylis (Fig. 6.40-41)
non-spiralled or irregularly spiralled 2 15a Collar with many spiral turns. Lorica narrow,
la Lorica wall hyaline or with agglutinated parti- elongate: Helicostomella (Fig. 6.30)
cles, chiefly on the bowl... 7 16 Oral end incomplete; collar with longitudinal
2 Lorica generally with an open or a broken-like facets: Dadayiella (Fig. 6.48)
aboral end; when aboral end is closed, it is not 16a Oral end well-defined 17
rounded nor flat: TintillllOpsis complex (Fig. 17 Aboral end open 18
6.18-29) 17a Aboral end closed 21
Tintinnoinea 341
18 Lorica without longitudinal striae, or with a few out horizontally in spira1 shelf: Climacocylis
tenuous ones: Elltilltinnus (Fig. 6.66-71) (Fig.6.118-119)
18a Lorica with longitudinal striae 19 32a Abora1 end with aboral horn or pedicel 33
19 Abora1 end complete, aperture often not easily 33 Oral margin generally channeled: Xystonella
distinguishable; lorica generally very elongate (Fig.6.120-121)
and nail-shaped: Salpillgella (Fig. 6.61-65) 33a Oral margin not channe1ed: Xystonellopsis (Fig.
19a Abora1 end scalloped, clearly open 20 6.122-129)
20 Oral margin without teeth: Datllrella (Fig. 6.73- 34 Lorica without longitudinal fins; with two easi1y
74) visible laminae 35
20a Oral margin with teeth: Clevea? (Fig. 6.72) 34a Lorica with a few 10ngitudina1 fins 39
21 Lorica with thick-walled aboral horn: Favella 35 Abora1 end tapering posteriorly to very long
(Fig. 6.82-83) aciculate spine: Albatrossiella (Fig. 6.10 1)
21a Abora1 horn absent or thin-walled 22 35a Without an acicu1ate spine 36
22 Oral end differentiated into an ¡nner and an outer 36 Lorica with 10wer cone prolonged as caudal
collar 23 lance: Parlllldella (Fig. 6.102-103)
22a Oral end not differentiated into an inner and an 36a Without cauda11ance 37
outer collar 26 37 Abora1 end never greatly or abruptly expanded:
23 Lorica wall with an anastomosed reticulum: Ulldella (Fig. 6.92-95)
Epiplocyloides? (Fig. 6.114-115) 37a Abora1 end broad1y rounded .38
23a Lorica wall without an anastomosed reticulum 38 Lorica consisting of anterior cylinder and abo-
......................................................................... 24 rally expanded bowl; one or more rings present
24 Oral end without teeth: Ascampbeliella (Fig. on cy1inder: Amplectella (Fig. 6.96-99)
6.76-79) 38a Lorica without rings; with suboral ledge:
24a Oral end with teeth 25 Undellopsis (Fig. 6.100)
25 Teeth 10cated on outer collar: Acanthostomella 39 Longitudina1 fins extending fulllength of lorica:
(Fig.6.75) Amphorellopsis (Fig. 6.51)
25a Teeth 10cated on inner collar: Cymatocylis (Fig. 39a Longitudina1 fins not extending full 1ength of
6.84-91) lorica .40
26 Wall with severa1 easi1y visible, vertical, simple 40 Aboral end acute; lorica with deep nucha1 con-
(dextrogyrous, 1evogyrous or both), branched or striction: Ormosella (Fig. 6.57-58)
anatomosed ribs 27 40a Abora1 end rounded or truncated; [orica without
26a Wall without ribs 28 nuchal constriction .41
27 With oral gutter: Rhabdollella (Fig. 6.104-108) 41 Abora1 end more or 1ess truncated: AmpllOrides
27a Without oral gutter: Protorhabdonella (Fig. (Fig. 6.52-55)
6.109-110) 41 a Abora1 end more or less rounded: Steellstrupiella
28 Wall with pronounced, anastomosed reticu1um (Fig. 6.59-60)
on posterior halL 29
28a Wall without reticulum 30 C1assification scheme based on morphologicalIy
29 With abora1 horn: Epiplocylis (Fig. 6.111-113) similar groups of species
29a Without aboral horn: Epicancella (Fig. 6.116)
30 Lorica wall with regular hexagona1 prismatic South Atlantic species are arranged below in 4 groups
a1veo1i 31 according to characteristics that can be easi1y distin-
30a Lorica wall without a1veoli 34 guished by standard microscope observation.
31 Bowl and collar separated by nuchal constric- However, some genera and species (Helicostomella,
tion: Cyttarocylis (Fig. 6.117) Metacylis and a few species of Dictyocysta and
31a Bow1 and collar not separated by nuchal con- Eutintinnus) are included in groups whose diagnostic
striction 32 features may on1y occasionally (or never) coincide
32 Abora1 end irregular or broken-1ike, without with those ofthe taxa. For instance, Helicostomella has
abora1 horn or pedice1; spira1 structure extending never been reported as having an agg1utinated lorica,
342 Tintinnoinea
but since all other genera with a well-differentiated spi- Subgroup 4C. Species with differentiations at
ralled collar and a bowl fall smoothly into the "agglu- oral end (inner and outer collar, oral band or den-
tinated 10rica group", it was found reasonable to tieles): Acanthostomella, Ascampbeliella, Favella,
include it there as well. Moreover, such exceptions are Cymatocylis.
applicable at infraspecific levels as well: Subgroup 4D. Lorica wall translucid, generally
Laackrnanniella naviculaefera includes 3 different with two conspicuous laminae: Undella,
morphotypes (or formae): forma coxliella and forma Amplectella, Undellopsis, Parundella, Albatros-
prolongata have a hyaline lorica, while forma navicu- siella?
laefera can be densely covered by biogenic particles Subgroup 4E. Lorica wall conspicuous, few to
(Alder, 1995). The "genus" Coxliella fumishes another many vertical, simple (dextrogyrous, levogyrous
vivid example of difficulties with tintinnid systematics: or both), branched, or anatomosed ribs: Rlzab-
several culture- and plankton sample-based studies donella, Protorlzabdonella?
(Laval-Peuto, 1981; Williams et al., 1994; Alder, Subgroup 4F. Lorica wall usually with conspicu-
1995) have demonstrated that at least some of the spe- ous, anastomosed reticulum on posterior half:
cies described in this genus are but transitional forms Epiplocylis, Epiplocyloides, Epicance/la,
of various species belonging to several genera. Cyttarocylis.
Subgroup 4G. Lorica wall with regular, hexago-
Group 1. Species with agglutinated biogenic or nal prismatic alveoli: Climacocylis, Xystonella,
arenaceous particles on their lorica Xystonellopsis.
Subgroup lA. Features of lorica obscured by Miscellaneous (ungrouped): Dadayiella.
densely agglutinated, evenly distributed particles:
Codonella, Codonaria, Tilltillllidium, Poroeclls, Subgroup lA (5 genera, 1 species group, 28 species)
Stylicallda, Tintillnopsis complex).
Subgroup lB. Lorica with clearly differentiated Genus Codollella Haeckel, 1873 emend. Brandt,
bowl and collar. Usually with agglutinated (biogen- 1907, emend. lorgensen, 1924 (Fig. 6.1-5).
ic and/or arenaceous) particles, 10cated almost Globose to ovate bowl, cornnlonly with agglutinated
exelusively on bowl: Laackmanniella, Codonellop- partieles (coccoliths and non-biogenic materials).
sis, Stenosemella. Helicostomella and Metacylis Collar separated from bowl by nuchal constriction.
(tatter never with agglutinated partieles). • Codollella galea Haeckel, 1873 (Fig. 6.1). Ref.:
Balech (1959).
Group 2. Lorica fenestrated • *Codonella aspera Kofoid and Campbell, 1929
Subgroup 2A. Conspicuous fenestrae on bowl (Fig. 6.2). See remark for Codonella amplzorella.
and collar; very exceptionally with agglutinated Ref.: Balech (1959).
(biogenic) partieles: Dictyocysta. Codonella sp. aff. C. olla (Kofoid and Campbell,
Subgroup 2B. Qne or two rows of small, incon- 1929) (Fig. 6.3). Ref.: Balech (1962b).
spicuous fenestrae 10cated on precollar and in • *Codonella natiollalis varo e Brandt, 1906. Ref.:
middle of bowl; lorica without agglutinated parti- Brandt (1906-1907).
cles: Petalotricha. Codonella amphorella Biedermann, 1893 (Fig.
6.5). Resembles *c. aspera. Cytological studies
Group 3. Lorica covered by an atypical gelatinous needed to differentiate one from other. Ref.:
external coating Balech (1962b).
Brandtiella. Codonella? perforata Entz Sr., 1884 (Fig. 6.6).
See remark for Codonaria cistellllla below. Ref.:
Group 4. Hyaline lorica Kofoid and Campbell (1929).
Subgroup 4A. Lorica wall usually with few longi-
tudinal fins: Stelidiella, Amphorellopsis, Ormo- Genus Codonaria Kofoid and Campbell, 1939 (Fig.
sella, Amphorides, Steellstrupiella. 6.7).
Subgroup 4B. Lorica open at both ends: Short pot-shaped 10rica. Structure of bowl and collar
Salpingella, ElItilltillllllS (very rarely with few almost homogeneous. Suboral cone superimposed on
scattered partieles on lorica), Clevea?, Datllrella. collar. Wall covered with coccoliths.
Tintinnoinea 343
Codollaria cistellula (Fol, 1884) (Fig. 6.7). ally belong to various other genera (Eutintinnlls,
Presence of a suboral cone both in C. eistellula Datllrella, Codonella, Stenosemella. Stylieallda,
and in Codonella pelforata (well-developed in the Climaeoeylis, Helieostomella and even Coxliella
former, smooth in the latter; Fig. 6.6) thought to forms) whose aspect is masked by the dense aggluti-
indicate relationship between these two genera. nation of particles due to coastal environmental con-
Ref.: Balech (l962b). ditions (conversely, neritic forms may fail to aggluti-
nate foreign particles when expatriated into the open
Genus Tilltinllidiulll Kent, 1882 (Fig. 6.8-9). ocean). Further studies are needed to confirm this
Lorica irregular in form (mostly as an elongated cyl- observation and to divide the genus Tintil7nopsis into
inder), probably because of variable size and quality morphologically coherent species. Members of the
of agglutinated (arenaceous) particles. Tintinnopsis complex have loricae highly variable in
Tintillllidium sp. aff. T. semiciliatum (Sterki, shape, and always densely covered with particles,
1879) (Fig. 6.8). Ref.: Barría de Cao (1981). mainly of non-biogenic origino The aboral end is
Tilltilllzidiulll balechi Barría de Cao, 1981 (Fig. closed in most species, clearly open in some, and with
6.9). Ref.: Barría de Cao (1981). a broken-like aboral end in a few. Despite the agglu-
tinated particles, spiral lorica wall structure can be
Genus Poroecus Cleve, 1902 emend. Kofoid and seen in some taxa. In some species a nuchal constric-
Campbell, 1929 (Fig. 6.10). tion may be either inconspicuous or well-defined.
Agglutinated lorica (coccoliths), cylindrical bowl,
with or without short aboral hom. "Tilltillllopsis" radix (lmhof, 1886) (Fig. 6.12).
Poroecus apiculatus (Cleve, 1900) (Fig. 6.10). See remark for Tintinnopsis aperta. Ref.: Brandt
Ref.: Brandt (1906-1907). (1906-1907), Balech (1945), Souto (1 970a, b).
"Tilltillnopsis" aperta Brandt, 1906 (Fig. 6.13).
Genus Stylicauda Balech, 1951 (Fig. 6.11). Morphologically similar to T. radix (Fig. 6.12c),
Cylindrical, coxlielliform bowl always with aboral sizes quite different. Cytological studies needed to
hom. differentiate these two species. Ref.: Brandt
• Stylicauda plateJlsis (Da Cunha and Da Fonseca, (1906-1907), Balech (1948).
1917) (Fig. 6.11). Ref.: Balech (1951), Souto "Tilltillllopsis" tocalltillellsis Kofoid and
(1970a). Campbell, 1929 (Fig. 6.14). Ref.: Brandt (1906-
1907).
Tilltinllopsis complex (Fig. 6.12-27). Above 3 species fit into Tintinnopsis because of
This complex encompasses species ascribed to the hom with a clearly broken-like aboral end.
genera Tintinnopsis Stein, 1867 and Leprotil7tinnus However, T. toeantinensis has a globose bowl
Jorgensen, 1899. According to Kofoid and Campbell extended cylindrically to the oral end, whereas T.
(1929), Tintil7l7opsis is the most diversified tintinnid radix and T. aperta usually have a cylindrical, spi-
genus (85 species in total), while Leprotintinnus com- ralled bowl.
prises only 5 species. However, a comparative analy- "TintillllOpsis" Ilordqvisti Brandt, 1906 (Fig.
sis of the lorica of most species commonly regarded 6.15). Broken-like aboral end (without aboral
as belonging to Tintinnopsis (see Kofoid and hom), expanded as in genera Clevea?, Daturella,
Campbell, 1929, Fig. 14-97 therein), and particularly (Fig. 6.72-74) and Climaeoeylis (Fig. 6.119). Ref.:
material s examined for the present work (Fig. 6.12- Brandt (1906-1907).
27), clearly show that their sole cornmon feature is the "TintillllOpsis" kofoidi Hada, 1932 (Fig. 6.16).
capability of accreting particles (mostly of non-bio- Ref.: Balech (1948).
genic origin) in a highly variable number. Other mor- "Tintillllopsis" amphistoma Balech, 1951 (Fig.
phologic traits, difficult to assess since they are usual- 6.17). Ref.: Balech (1951).
Iy obscured by the thick layer of particles adhering to Above 2 species morphologically similar to genus
the lorica, are very variable and do not allow consis- Eutintil7nlls (Fig. 6.66-71) because loricae clearly
tent separation of the morphotypes into reasonably open at both ends.
homogeneous groups. Furthermore, it is probable that • "Tilltillllopsis" ampllOra Kofoid and Campbell,
many of the so called "species" of Tintinnopsis actu- 1929 /Ilucula (Fol) (Fig. 6.18). Bowl amphora-
344 Tintinnoinea
like. Cytological studies are needed to confirm T. "Tilltbl1/opsis" compressa (Daday, 1887) (Fig.
amphora Kofoid and Campbell, 1929 and T. nucu- 6.25). Dubious species in need of additional stud-
la (Fol, 1884) as valid species. Ref.: Brandt (1906- ies. Lorica shows slight nuchal constriction.
1907), Barría de Cao (1981). Il\ustrations provided by Balech (1959) for T.
"TintillllOpsis" baltica Brandt, 1896 (Fig. 6.19). compressa suggest morphological affinities
Ref.: Brandt (1906-1907), Balech (1945, 1948), between this species and T. campanula/butschlii.
Souto (1979). Ref.: Daday (1887b).
"TilltillllOpsis" llruguayensis Balech, 1948 (Fig. • "TilltillllOpsis" beroidea Stein, 1867 (Fig. 6.26).
6.20). Ref.: Balech (1948). Despite wealth of information on this species,
"TilltitlllOpsis" jimbriata Meunier, 1919 (Fig. including cytological studies (Gold and Morales,
6.22). Ref.: Brandt (1906-1907), Balech (1948). 1975), its taxonomic status still dubious, especial-
"Tintinllopsis" campallula Ehrenberg, 1840 Iy regarding its relationships with Tintinnopsis
Ibutschm Daday, 1887 (Fig. 6.21). See T. COnl- parvula J6rgensen (see extensive synonymies in
pressa below. Ref.: Daday (1887b), Brandt (1906- Kofoid and Campbell, 1929, and herein). Ref.:
1907), J6rgensen (1924), Balech (1948). J6rgensen (1912, 1927), Kofoid and Campbell
"Timb/llopsis" campallulalbutscltlii varo blltsch- (1929).
m Daday, 1887 Ref.: Daday (1887b). • "Tilltblllopsis" p"silla Hada, 1937/laevis Wailes,
• "TintillllOpsis" campallulalbutschm varo mor- 1925 (Fig. 6.28). Tintinnopsis pusilla/laevis com-
tellsenü (Schmidt, 1901). Ref.: Balech (1948). prises T. pusilla Hada, 1937 and T. laevis Wailes,
Above four species have in cornmon nuchal con- 1925. Morphological transitions in aboral ends of
striction that can be salient (T. uruguayensis, T. these specimens (Fig. 6.28a-e) do not support stat-
fimbriata) or smooth (T. baltica, T. bütschlii). In us as independent species. However, further cyto-
some cases, oral end seems to be broken-like (T. logical studies are required to confirm this group-
uruguayensis with short hom with broken-like ing. Ref.: Barría de Cao (1981).
aboral end), whereas others show anterior region "Tilltinllopsis" prowazeki Faría and Da Cunha,
ofbowl spiralled, or combination ofboth elements 1917 (Fig. 6.27). Aboral hom as long as collar.
(Fig. 6.19f). Resembles Codonellopsis morphologically (par-
"TilltillllOpsis" parvula J6rgensen, 1912 (Fig. ticularly Codonellopsis gaussi forma typica, Fig.
6.23). Individuals highly variable in morphology 6.3ge). Ref.: Faría and Da Cunha (1917).
and dimensions. In some cases posterior cone can "Tilltillllopsis" eratera (Leidy, 1879).
be distinguished in bowl. In others, bowl consists "TilltitlllOpsis" rioplatellsis Souto, 1970.
of straight cylinder rounded aborally. Latter forms Above two species (not illustrated) are fresh-water
(Fig. 6.23k) relate this species to T. lobiancoi (Fig. inhabitants reported from the Río de la Plata estuary
6.24a). See also T. beroidea below. Ref.: Brandt (Souto, 1979).
(1906-1907), J6rgensen (1924), Balech (1948),
Souto (1970a). Subgroup lB (5 genera, 12 species)
"TilltillllOpsis" lobiallcoi Daday, 1887 Ibrasilien-
sis Kofoid and Campbell, 1929 (Fig. 6.24). This Genus Helieostomella J6rgensen, 1924 (Fig. 6.30).
species encompasses T. lobiancoi Daday, 1887, Elongate, narrow, hyaline lorica. Aboral hom present.
whose typical morphology is illustrated in Fig. Aboral end always closed. Collar and aboral hom
6.24a, and T. brasiliellsis (Brandt, 1906-1907), highly variable in length. Oral end generally continu-
with much shorter lorica. Lorica illustrated in Fig. ous, sometimes with teeth.
6.24e conditionally included in this species . Ref.: • Helicostomella subulata (Ehrenberg, 1833) (Fig.
Daday (1887b), Brandt (1906-1907), Balech 6.30). Ref.: Brandt (1906-1907), Balech (1959),
(1945), Souto (1970a). Souto (1979).
• "TilltillllOpsis" parva Merkle, 1909 (Fig. 6.29).
Individuals like those illustrated in Fig. 6.28g, h, Genus Laackmalllliella Kofoid and Campbell, 1929
show incipient, small collar covered by particles, emend. Alder, 1995 (Fig. 6.31).
latter also present on entire bowl. Ref.: Balech Aboral end usually open. Imbricate collar, variable in
(1945,1948). length. Bowl cylindrical. elongate, either completely
Tintinnoinea 345
hyaline, or partially to totally eovered with particles (Fig. 6.37a, b). Ref.: Balech (1948), Barría de Cao
(mainly diatoms). Monospeeifie genus. (1981).
Laacklllanniella naviclllaejera (Laaekmann, C. lIlorchella (Cleve, 1900) (Fig. 6.33). Collar fre-
1910) (Fig. 6.31). Ref.: Laaekmann (1910), quently showing two fenestrae. Bowl with agglu-
Baleeh (1947), Petz el al. (1995a), Alder (1995). tinated arenaeeous particles. Ref.: Brandt (1906-
L. naviclllaejera, forma llaviclIlaejera stat. nov. 1907), Jorgensen (1924), Balech (1959, 1971a),
(Fig. 6.3la-d). Loriea with agglutinated particles Souto (1979).
in highly variable concentrations. Ref.: C. lIlorchella, forma schabi Brandt, 1906 (Fig.
Laackmann (1910), Alder (1995). 33e-h): Bowl more elongate. Ref.: Brandt (1906-
L. Ilaviculaejera, forma prolongata stat. nov. 1907), Balech (1959).
(Fig. 6.31 e). Lorica non-spiralled, hyaline. Ref.: C. 111 orch ella , forma typica nov. formo (Fig.
Laackmann (1910), Alder (1995). 6.33a-d): Bowl rounded. Ref.: Brandt (1906-
L. llaviclIlaejera, forma coxliella nov. formo (Fig. 1907).
6.31 f). Lorica totally spiralled, hyaline. Aboral Codollellopsis obesa Balech, 1948 (Fig. 6.35).
end closed. Ref.: Alder (1995). Bowl with agglutinated arenaceous particles and
collar without visible spiral structure. Ref.: Balech
Genus Codollellopsis Jorgensen, 1924 (Fig. 6.32-39). (1948,1951).
Aboral end always closed, sometimes bearing well- • Codonellopsis ortllOceras (Haeckel, 1873) (Fig.
developed aboral horno Some species highly polymor- 6.32). Collar and aboral hom highly variable in
phic, with either agglutinated or totally hyaline lori- length. Ref.: Brandt (1906-1907), Jorgensen
cae. Collar sometimes with fenestrae. (1924), Balech (1959).
• Codollellopsis balechi Hada, 1970 (Fig. 6.38). Codollellopsis pusi/la (Cleve, 1900) (Fig. 6.36).
Collar short. Bowl with agglutinated (arenaceous) Collar short, with 4-6 fenestrae. Bowl with arean-
particles. Ref.: Balech (1971a), Alder (1995). ceous and biogenic agglutinated particles. Ref.:
*Codonellopsis brasiliensis (Brandt, 1906) (Fig. Balech (1971 a).
6.34). Dubious speeies. Resembles C. morchella
forma schabi (see Fig. 6.33h and 3.34a), yet Genus Stelloselllella Jorgensen, 1924 (Fig. 6.42)
slightly smaller in size. Collar with two fenestrae Bowl small, ovoidal, with short collar with 1 or 2 spi-
(as in C. morchella). Ref.: Brandt (1906-1907). ral tums. Sometimes with fenestrae at base of collar.
• Codonellopsis gaussi (Laackmann, 1907) (Fig. A lid or "closing apparatus" at oral end.
6.39). Ref.: Laackmann (1910), Balech (1973), • Stelloselllella avellalla (Meunier, 1919) (Fig.
Alder (1995). 6.42). Bowl with agglutinated particles. Lid some-
C. gallssi, forma coxliella nov. form. (Fig. 6.39f). times also with agglutinated particles. Ref.:
Lorica totally spiralled and hyaline. Ref.: Alder Balech (1948), Souto (1979).
( 1995).
C. gaussi, forma cylindrocollica nov. formo (Fig. Genus Metacylis Jorgensen, 1924 (Fig. 6.40, 41).
6.39a-c). Collar variable in length. Short aboral Small loricae, bowl cylindrical or oval-shaped, with
hom. Bowl totally or partially covered with parti- short collar. Wall always hyaline.
cles (mainly diatoms) or hyaline. Ref.: Laackmann • Metacylis jorgenseni (Cleve, 1902) (Fig. 6.40).
(1910), Sassi and Melo (1986), Alder (1995). Ref.: Jorgensen (1924).
C. gaussi, forma globosa nov. formo (Fig. 6.39d). • Metacylis sp. aff. M lIlereschkowskyi (Kofoid and
Collar and aboral hom variable in length. Ref.: Campbell, 1929) (Fig. 6.41). Ref.: Jorgensen (1924).
Laackmann (1910), Balech (1973), Alder (1995).
C. gaussi, forma typica nov. formo (Fig. 6.3ge). Sub-group 2A (1 genus, 4 species)
Long collar and aboral hom. Bowl expanded in
posterior region, always with agglutinated parti- Genus Dictyocysta Ehrenberg, 1854emend. Kofoid
cles. Ref.: Laackmann (1910), Balech (1973), and Campbell, 1929 (Fig. 6.43-46).
Alder (1995). Bowl usually globose. Collar eylindrieal. Collar and
• Codonellopsis lusitanica Jorgensen, 1924 (Fig. bowl (or only collar) with large fenestrae in one to
6.37). Collar often totally covered with particles several rows, sometimes with agglutinated coccoliths.
346 Tintinnoinea
Oral end smooth or denticulated. One of most typical Elongate, subconicallorica, with facets mainly on col-
and clearly distinguishable tintinnids in general. lar. Oral rim entire or crenulate. Aboral hom without
*Dictyocysta calijorlliensis Kofoid and Campbell, knob, or with rudimentary, small knob on terminal
1929 (Fig. 6.46). Oral margin denticulate. Ref.: end.
Balech (1971a). • Dadayiella ganymedes (Entz Sr., 1884) (Fig.
• Dictyocysta duplex (Brandt, 1906) (Fig. 6.45). 6.48). Collar with vertical, short facets projecting
Bowl without fenestrae. Ref.: Brandt (1906-1907), from lorica as "hairs". Ref.: J6rgensen (1924),
Kofoid and Campbell (1929). Balech (1959), Souto (1979).
• Dictyocysta elegans (Ehrenberg, 1854) (Fig.
6.44). Ref.: Brandt (1906-1907), J6rgensen Subgroup 4A. Lorica wall usually with a few
(1924), Balech (1959). longitudinal fins
D. elegans, varo speciosa J6rgensen, 1924 (Fig.
6.44a-f). Ref.: Brandt (1906-1907), J6rgensen Genus Stelidiella Kofoid and Campbell, 1929 (Fig.
(1924), Balech (1959). 6.50).
D. elegalls, varo lepida (Ehrenberg, 1854) (Fig. Collar truncated, smooth or fenestrated (one or two
6.44g-n). Ref.: Brandt (1906-1907), Laackmann rows of quadrilateral fenestrae). Entire or toothed sub-
(1910), Ba1ech (1959). oral bando Bowl with few but well-defined longitudi-
• Dictyocysta mitra Haecke1, 1873 (Fig. 6.43). nal fins.
Collar distinguished by fenestrae, where they are • Stelidiella stelidillm (Biedermann, 1893) (Fig.
larger than on rest of lorica. Ref.: Brandt (1906- 6.50). Collar with one row of 8 quadrilateral
1907), Ba1ech (1959). fenestrae and toothed suboral bando Ref.: Brandt
( 1906-1907).
Sub-group 2B (1 genus, I species)
Genus AmpllOrellopsis Kofoid and Campbell, 1929
Genus Petalotriclla Kent, 1882 emend. Brandt, 1907 (=Odonfophorella Kofoid and Campbell, 1929)
(Fig. 6.47). emend. (Fig. 6.51).
Spheroidal to conical10rica. Bowl separated from col- Amphora-shaped lorica (endirrg in corre) with funnel-
lar and pre-collar by nuchal constriction. Thin collar, shaped oral end. Few but distinguishable vertical
separated from pre-collar by row of irregular fenes- blade-like fins throughout wall or mainly on posterior
trae; frenestrae also on bowl. Oral end smooth, undu- end. Oral rim sometimes toothed.
lated, or toothed. • AmpllOrellopsis qllinqllealata (Laackmann, 1907)
• Petalotriclla ampulla (Fol, 1881) (Fig. 6.47). (Fig. 6.51). Ref.: Laackmann (1910), Balech
Globose bowl, oral margin toothed. Ref.: Brandt (1971a), Alder (1995).
(1906-1907), Balech (1959).
Genus Ormosella Kofoid and Campbell, 1929 (Fig.
Group 3 (one monospecific genus) 6.56-58).
Bell-shaped lorica. Aboral end sometimes acute or
Genus Brandtiella Kofoid and Campbell, 1929 (Fig. with simple spinule. Bowl with none to few vertical
6.49). fins. Collar as hyaline cylinder superimposed on
Collar with suboral ring. Bowl with incipient longitu- bowl.
dinal ribs. • Ormosella cOrllucopia Kofoid and Campbell,
• Brandtiella palliata (Brandt, 1906) (Fig. 6.49). 1929 (Fig. 6.56). Lorica without facets. Ref.:
Presence of fins relates this species to genera Brandt (1906-1907).
Amphorel!opsis, Ornzosella, Steenstrupiella and Ormosella acantllarlls (Kofoid and Campbell,
Amphorides. Ref.: Brandt (1906-1907). 1929) (Fig. 6.57). Bowl commonly with 6-8 deli-
cate facets. Ref.: Balech (1971a).
Group 4 (29 genera, 82 species) • Ormosella IIaeckeli Kofoid and Campbell, 1929
(Fig. 6.58). Bowl with 8-16 delicate facets. Ref.:
Genus Dadayiella Kofoid and Campbell, 1929 (Fig. Kofoid and Campbell (1929), Balech (1971 a),
6.48). Souto (1979).
Tintinnoinea 347
Genus AmpllOrides Strand, 1928 (Fig. 6.52-55). Salpingella aClImillata, varo glockelltogeri Brandt,
Lorica bowl truncated at aboral end. With 3 to 5 lon- 1906 (Fig. 6.61e, f). Aboral region slightly expan-
gitudinal fins mainly on posterior region of bowl. ded next to aboral cylinder. Ref.: Brandt (1906-
Funnel-shaped oral end. See Steel7strupiella. 1907), Jorgensen (1924).
• Amphorides calida (Kofoid and Campbell, 1929) S. acumillata, varo undata Jorgensen, 1899 (Fig.
(Fig. 6.52). Ref.: Brandt (1906-1907), Kofoid and 6.6Ig, h). Oral margin with tooth-like lobules.
Campbell (1929). Ref.: Brandt (1906-1907), Jorgensen (1924).
Amphorides quadrililleata (Claparede and • Salpillgella acumillatoides (Laackmann, 1910)
Lachmann, 1858-1859) (Fig. 6.53). (Fig. 6.62). Ref.: Laackmann (1910), Alder (1995).
Amphorides quadrilineata, varo brasiliensis S. aCllmillatoides, varo secata Laackmann, 1910
(Laackmann, 1910) (Fig. 6.53). Ref.: Laackmann (Fig. 6.62a, b). Oral margin entire. Ref.: Laackmann
(1910). (1910).
• Amphorides laacklllallni (Jorgensen, 1924) (Fig. S. aculIlillatoides, varo ulldata (Laackmann,
6.54). Ref.: Laackmann (1910). 1910) (Fig. 6.62d). Oral margin with lobules. Ref.:
• AlIlphorides amphora (Claparede and Lachmann, Laackmann (1910).
1858-1859) (Fig. 6.55). Ref.: Strand (1928), • Salpingella Sllbcollica Kofoid and Campbell, 1929
Durán (1965), Souto (1979). (Fig. 6.63). Ref.: Kofoid and Campbell (1929),
Balech (1971a).
Genus Steellstrupiella Kofoid and Campbell, 1929 • Salpillgella costata (Laackmann, 1910) (Fig. 6.64).
(Fig. 6.59-60). Bowl with easily visible fins. Ref.: Laackmann
Lorica elongate, slender, normally with two thin fins (1910).
extending along less than half of bowl, with funnel- • Salpingella regulata (Brandt, 1906) (Fig. 6.65).
shaped collar. Aboral end slightly rounded. Genus Bowl with 8-10 longitudinal fins. Longest lorica in
morphologically very similar to Amphorides, requir- genus. Ref.: Brandt (1906-1907).
ing further studies of their systematic relationships.
• Steellstrllpiella steellstrupii (Claparede and Identification of Salpingella species is extremely dif-
Lachmann, 1858-1859) (Fig. 6.59). Ref.: Balech ficult, and consistent assignments do not seem pos-
(1959, 1962b), Souto (1979). sible at present. The fin s, usually present and best
S. steenstrllpii, varo robusta (Kofoid and developed on the posterior region of the bowl (usual-
Campbell, 1929). Ref.: Balech (1962b). Iy extending longitudinally, with or without torsion),
*Steenstrllpiella pozzi Balech, 1942 (Fig. 6.60). can also reach the anterior region, thus suggesting
Ref.: Balech (1942), Souto (1979). morphologic relationships between S. acuminatoides
and S. costata. Although with a much shorter lorica,
Subgroup 4B. Lorica open at both ends the general morphology of S. subconica (Fig. 6.63a,
b) strongly resembles that of S. acuminatoides.
Genus Salpillgella Jorgensen, 1924, emend. Kofoid Further studies on the morphologic variability ofthese
and Campbell (Fig. 6.61-65). species are necessary in order to clarify the systemat-
Bowl thin, with fins mainly on posterior end, (although ics of the genus. Therefore, the present classification
in some cases covering whole surface of bowl), with must be considered provisional.
dextrogyrous torsion (sometimes obscuring aboral end
opening). Funnel-shaped oral end, smooth or denticu- Genus Eutintillllus Kofoid and Campbell, 1939
late (=Salpingacantha Kofoid and Campbell, 1929). emend. (Fig. 6.66-71) (= Eutintinnus, Ceratotintinnus
• Salpingella aClll1linata (Claparede and Lachmann, and Odol7totintinnus, subgenera erected by Kofoid
1858-1859) (Fig. 6.61). Ref.: Jorgensen (1924), and Campbell, 1939).
Balech (1959), Souto (1979). Lorica cylindrical. Oral and aboral ends sometimes
Salpillgella aCllmillata, varo secata Brandt, 1906 funnel-shaped. Oral end frequently smooth, in some
(Fig. 6.61 a-d). Aboral cylinder incipient (Fig. cases with well-defined teeth. Wall usually hyaline;
6.61 a, b) or well-developed (Fig. 6.61 d). Ref.: exceptionally with short, numerous striae throughout.
Brandt (1906-1907). The systematics of this genus is chaotic. Fig. 6.66-71
348 Tintinnoinea
illustrate the most characteristic morphological traits Subgroup 4C. Species with differentiations at the
of South Atlantic species. Exeept for studies with oral end (an inner and an outer collar, an oral
descriptions of new species, no comprehensive work band or denticIes)
can be recornmended for this genus.
• Eutilltill1ll1s jraklloi (Daday, 1887) (Fig. 6.66). Genus Acalltlrostomella J6rgensen, 1927 (Fig. 6.75).
Everted oral and aboral ends. Cup-shaped, subconical lorica. Oral rim toothed.
• Eutintill1ll1S lusus-lIIldae (Entz Sr., 1885) (Fig. Inner and outer collar.
6.67). Oral end wider than aboral end. • Acalltlrostomella llorvegica (Daday, 1887) (Fig.
• Eutilltill1ll1S lIledius (Kofoid and Campbell, 1929) 6.75).
(Fig. 6.68). Loriea a straight cylinder, with an Acalltlrostomella Ilorvegica, forma typica nov.
everted oral end. formo (Fig. 6.75a-c). Globose bowl, sometimes
*EUtilltilllIllS rugosus Kofoid and Campbell, showing smooth middle constriction. With or
1939 (Fig. 6.69). Cylindrieal, striated bowl with without small aboral hom. Ref.: Daday (1887b),
toothed oral end. Balech (1971a).
• Eutilltbl1luS attell11atus (Kofoid and Campbell, Acantlrostomella lIorvegica, forma lata (Kofoid
1929) (Fig. 6.70). Thin bowl with funnel-shaped and Campbell, 1929) (Fig. 6.75d). Bowl bell-
oral end. shaped. Ref.: Kofoid and Campbell (1929),
• Eutilltillll11S apertus (Kofoid and Campbell, Balech (197Ia).
1929) (Fig. 6.71). Small eylinder with slight dila-
tion at middle. Genus Asca11lpbeliella Corliss, 1960 (Fig. 6.76-79).
Small, cup-shaped lorica with two collars. Inner col-
Genus Clevea? Balech, 1948 (Fig. 6.72). lar superimposed on outer one. Oral rim always
Cylindrical, striated bowl; oral end toothed; aboral devoid of teeth.
end sealloped. *Ascampbeliella acuta (Kofoid and Campbell,
• Clevea? melcllersi Balech, 1948 (Fig. 6.72). Only 1929) (Fig. 6.77). Ref.: Souto (1979).
species in genus, erected to accommodate Clevea *Ascampbeliella armilla (Kofoid and Campbell,
melchersi Baleeh, 1948 (Fig. 6.72), which might 1929) (Fig. 6.79). Ref.: Balech (1971b).
represent a transitional form between ElItintillnlls • Asca11lpbeliella obscura (Brandt, 1906) (Fig.
and Datllrella. Despite its smaller size, the gener- 6.76). Ref.: Brandt (1906-1907).
al aspect resembles that of ElItintinnlls rllgoslIs • Asca11lpbeliella ltrceolata (Ostenfeld, 1899) (Fig.
(Fig. 6.69), main difference being the conspicuous 6.78). Ref.: Balech (1971b).
scalloped rim on the aboral end of C. melchersi All records Ascampbeliella species are based on only
(similar to that of Datllrella emarginata, Fig. a few specimens, so morphological descriptions are
6.73). Ref.: Baleeh (1948). still scarce and incomplete.
Genus Daturella Kofoid and Campbell, 1929 (Fig. Genus Favella J6rgensen, 1924 (Fig. 6.80-83).
6.73, 74). Lorica campanulate, cylindrical or subconical, always
Elongate, trumpet-shaped lorica. Hyaline wall, with with aboral horno Oral end entire (sometimes with a
either slender or easily visible, ribbon-like fins. circumoral band), or with small denticles. Suboral
• Daturella elllargillata (Brandt, 1906) (Fig. 6.73). zone sometimes with annuli.
Entire oral end; scalloped aboral end. *Favella arcllata (Brandt, 1906) (Fig. 6.81). Oral
Longitudinal fins located generally on posterior rim entire. Suboral zone with annuli. General
region of bowl. Ref.: Brandt (1906-1907). aspect resembles that of F. ser/·ata. Ref.: Brandt
• Daturella datllra (Brandt, 1906) (Fig. 6.74). ( 1906-1907).
Longitudinal, ribbon-like fins extending all along • Favella elrrenbergii (Claparede and Laachmann,
bowl, sometimes showing levogyrous torsion near 1858) (Fig. 6.82). Cylindrical bowl, rounded ter-
oral end. Ref.: Brandt (1906-1907). minally. Entire oral end. Aboral horn very stout,
with conspicuous pleats. Sometimes with spiralled
Tintinnoinea 349
epiloriea. Ref.: Brandt (1906-1907), Jorgensen followed by thin and short lance. Ref.: Laaekmann
(1924), Baleeh (1959), Laval-Peuto (1981). (1910), Alder (1995).
F. ellrellbergii, forma coxliella Laval-Peuto, C. cOllvallaria, forma coxliella nov. formo (Fig.
1981 (Fig. 6.82e). Spiralled, hyaline loriea with 6.84c). Lorica wall spiralled. Ref.: Laaekmann
spiral turns variable in size and number. Ref.: (1910), Alder (1995).
Laval-Peuto (1981). C. cOllvallaria, forma typica Laackmann, 1910
F. ellrellbergii, forma decipiells Jorgensen, 1924 (Fig. 6.84d). Globose lorica with short aboral
(Fig. 6.682t). Ref.: Laval-Peuto (1981). horno Ref.: Laackmann (1910), Balech (1947),
F. ellrellbergii, forma favella Laval-Peuto, 1981 Alder (1995).
(Fig. 6.82a-d). Ref.: Brandt (1906-1907), • Cymatocylis drygalskii (Laackmann, 1907) (Fig.
Jorgensen (1924), Baleeh (1959). 6.86). Ref.: Laackmann (1910), Alder (1995).
*Favella serrata (Mobius, 1887) (Fig. 6.83). C. drygalskii, forma coxliella nov. formo (Fig.
Toothed oral end. Suboral eonstrietion followed 6.86b): Spiralled lorica bowl. Ref.: Laackmann
by ineipient annulus. Ref.: Brandt (1906-1907), (1910), Alder (1995).
Baleeh (1959). C. drygalskii, forma ovata Laackmann, 1910
• Favella taraikaensis Hada, 1932 (Fig. 6.80). Oral (Fig. 6.86a). Globose to elongate bowl, without
end with eireumoral bando Simple aboral hom. constrictions. Ref.: Laackmann (1910), Balech
Ref.: Baleeh (1948, 1951), Souto (1970a). (1947), Alder (1995).
C. drygalskii, forma typica Laackmann, 1910
Genus Cymatocylis Laaekmann 1910 emend. Kofoid (Fig. 6.86c). Elongate bowl, sometimes with con-
and Campbell 1929 emend. (Fig. 6.84-91). striction at middle. Ref.: Laackmann (1910),
Cylindrieal, elongate, chalice-shaped lorica bowl, Balech (1947), Alder (1995).
highly variable in size. Oral rim entire or toothed; Cymatocylis llobilis (Laackmann, 1907) (Fig.
some species showing an outer, velum-like oral ridge. 6.88). Ref.: Laackmann (1910), Alder (1995).
Wall totally hyaline or, less frequently, partially or Cymatocylis valllloffeni (Laackmann, 1907) (Fig.
entirely striated. With or without an aboral horn of 6.87). Striated wall; striae cornmonly located on
variable length. anterior region of lorica. Oral end everted, without
*Cymatocylis cristallilla Laaekmann, 1910 (Fig. an outer oral ridge. Usually with an aboral horno
6.91). Bowl with ineipient striae irregularly di s- Ref.: Laackmann (1910), Alder (1995).
tributed. Ref.: Laackmann (1910). C. VallllOjJelli, forma cOllica Laackmann, 1910
*Cymatocylis kergllelensis Laackmann, 1910 (Fig. (Fig. 6.87t). Ref.: Laackmann (1910), Alder (1995).
6.90). Lorica wall striated. Ref.: Laackmann (1910). C. valllloffelli, forma coxliella nov. formo (Fig.
• Cymatocylis alltarctica (Cleve, 1901) (Fig. 6.85). 6.87c). Lorica wall totally spiralled. Ref.:
The smallest species in genus. Abnormal individ- Laackmann (1910), Alder (1995).
uals are eommon. Ref.: Laaekmann (1910). C. VallllOjJelli, forma cylilldrica Laackmann, 1910
Balech (l958a, 1971), Alder (1995). (Fig. 6.87g). Ref.: Laackmann (1910), Alder (1995).
C. alltarctica, forma coxliella nov. formo (Fig. C. valllloffelli, forma flava Laackmann, 1910
6.85b). Ref.: Williams el al. (1994), Alder (1995). (Fig. 6.87b). Aboral horn short. Ref.: Laackmann
C. alltarctica, forma typica nov. formo (Fig. (1910), Alder (1995).
6.85a). Ref.: C1eve (1901), Balech (l971a). C. vallllojJelli. forma sllbrotlmdata Laackmann,
• Cymatocylis calyciformis (Laackmann, 1907) 1910 (Fig. 6.87d). Without aboral horno Ref.:
(Fig. 6.89). Oral end with inner rim and outer Laackmann (1910). Alder (1995).
ridge. Abora1 horn long (sometimes twice as long C. vallllOjJelli. forma typica Laackmann, 1910
as the bowl). Ref.: Laackmann (1910). Petz et al. (Fig. 6.87a). Lorica wall entirely striated. Aboral
(l995a), Alder (1995). horn long. Ref.: Laackmann (1910), Alder (1995).
• Cymatocylis cOllvallaria (Laaekmann, 1910) (Fig. Although lorical morphology has been investigated
6.84). Ref.: Laackmann (1910), Boltovskoy el al. extensively. similarities between C. convallaria,
(1990), Petz el al. (l995a), Alder (1995). forma typica and C. drygalskii. forma ovala; between.
C. cOllvallaria. forma affillis Laackmann. 1910 C. drygalskii. forma typiC4 and C. nobilis; and
(Fig. 6.84a, b). Globose and striatect aboral horn between C. vanhoffeni (forma cylindroconica and
350 Tintinnoinea
forma jlava) and C. cristallina, stress the need for Ulldellops;s marsupialis (Brandt, 1906) (Fig.
cytological studies to confirm the validity of these 6.100). '
species and the general systematic organization of the
genus (see Epiplocyloides antarctica below). Genus Parulldella J6rgensen, 1924 emend. Kofoid
and CampbelI, 1929 (Fig. 6.102, 103).
Subgroup 4D. Lorica wall translucid, generally Lorica with upper cylinder and lower cone. Two con-
with two conspicuous laminae spicuous laminae. Differs from Undella in larger size
and presence of aboral horn which, in some species,
Genus UlIdella Daday, 1887 emend. Kofoid and shows wrinkles or striae and short caudal lance.
Campbell, 1929, emend. Balech, 1975 (Fig. 6.92-95). Texture of lorica walI and presence of aboral horn
Lorica cylindrical, globose or cone-shaped. Aboral with caudal lance relate this genus to Xystonellopsis
end expanded, acutiform. or rounded. Laminae with (see Fig. 6.122-125). The latter, however, is clearly
syrnmetric or asyrnmetric suboral dilation or thicken- distinguished from the former by the presence of a
ing (see Parl/ndella below). well developed skirt.
*UlIdella gralld;s (Laackmann, 1910) (Fig. 6.94). • Panmdella caudata (Ostenfeld, 1899) (Fig.
Ref.: Laackmann (1910), Balech (1975). 6.103). Bowl elongate cylinder with thick, striated
Ulldella claparedei group (Fig. 6.95). Because of aboral horn which, on terminal end, may bear
continuous morphologic transitions between U incipient to well-developed caudal lance. Ref.:
claparedei (Entz Sr., 1885), U sllbcalldata sllb- J6rgensen (1924), Balech (1971a), Souto (1979).
cal/data (J6rgensen, 1924), U globosa (Brandt • Partmdella mess;lIells;s (Brandt, 1906) (Fig.
1906-1907), U perpllsilla (Kofoid and Campbell, 6.102). Bowl clearly expanded in posterior region.
1929), and U sllbacllta Cleve, 1901, these species Striated aboral horn, always with caudal lance.
are placed in the U claparedei group. Slightly dif- Ref.: Brandt (1906-1907), J6rgensen (1924),
ferent forms within this spectrum of variability Souto (1979).
(Fig. 6.95a-n) are tentatively attributed to dissimi-
lar environmental settings. Genus Albatross;ella? Kofoid and CampbelI, 1929.
Ulldella hem;sphaer;ca Laackmann, 1910 (Fig. Bowl subcylindrical, tapering distally into long, thin
6.93). Ref.: Laackmann (1910). aboral horn or "spine". Dubious genus comprising
Ulldella hyalilta Daday, 1887 (Fig. 6.92). Ref.: total of 3 species, only one of which has been men-
Brandt (1906-1907), Balech (1975). tioned from the South Atlantic.
• Albatrossiella? jiligera (Laackmann, 1910) (Fig.
Genus Amplectella Kofoid and Campbell, 1929 6.101). Ref.: Laackmann (1910).
emend. Balech, 1975 (Fig. 6.96-99).
Lorica roughly cylindrical, with one or more rings, Subgroup 4E. Lorica wall with conspicuous, few
expanding gradually to globose aboral end. to many, vertical, simple (dextrogyrous, levogyrous
*A mplectella mOllocollaria (Laackmann, 1910) or both), branched or anatomosed ribs
(Fig. 6.97). Ref.: Balech (1975).
• Amplectella Iluda (Kofoid and Campbell, 1929) Genus Rhabdollella Brandt, 1907 emend. (Fig. 6.104-
(Fig. 6.99). Ref.: Balech (1975). 108) (=Rhabdonellopsis Kofoid and CampbelI, 1929).
• Amplectella tr;collaria (Laackmann, 1910) (Fig. Lorica from smalI and stout to elongate, conical, or
6.98). Ref.: Balech (1975). chalice-shaped, usually with gutter between inner and
• Amplectella tridiv;sa (Brandt, 1906) (Fig. 6.96). outer laminae (mainly in larger species). With an abo-
Ref.: Balech (1975). ral horn or pedicel. Lorica walI sometimes with very
small fenestrae located between ribs. Oral rim always
Genus Ultdellops;s Kofoid and CampbelI, 1929, toothless.
emend. Balech, 1975 (Fig. 6.100). *Rhabdollella amor (Cleve, 1900) (Fig. 6.108).
Subcylindrical lorica bowl. Oral end with cylindrical Conical to elongate bowl. Incipient to well-devel-
band; suboral ledge present. Monospecific genus oped aboral hom. Delicate ribs generalIy showing a
results from tentative synonymy of U marsl/pialis levogyrous torsion near the oral end. Ref.: Brandt
(Brandt, 1906) and U subangulata (J6rgensen, 1924). (1906-1907), Laackmann (1910), Souto (1979).
Tintinnoinea 351
*Rhabdonella amor, varo cuspidata Brandt, 1906 oral end. Aboral hom rudimentary or well-developed.
(Fig. 6.108g). Ref.: Brandt (1906-1907). Two laminae well-differentiated. Suboral zone with a
*Rhabdollella amor, var. indica Laackmann, 1910 lip-shaped collar.
(Fig. 6.108a, c, d, h). Ref.: Laackmann (1910). o Epiplocylis acuminata (Daday, 1887) (Fig.
*Rhabdonella amor, varo valdestriata Brandt, 6.111). Coarse reticulum only absent in the sub-
1906 (Fig. 6.108h). Ref.: Brandt (1906-1907). oral zone. Well-developed lip-shaped collar (see
*Rhabdonella apophysata (Cleve, 1900) (Fig. E. calyx? below). Ref.: Daday (1887b), Brandt
6.104). Conical bowl. Pedicel as long as or 10nger (1906-1907), Balech (1959), Souto (1979).
than bowl. Ref.: Brandt (1906-1907), Balech *Epiplocylis calyx (Brandt, 1906) (Fig. 6.112).
(1962b), Souto (1979). Very similar to E. aCl/minata, although slightly
*Rhabdonella chiliensis Kofoid and Campbell, narrower. Ref.: Brandt (1906-1907).
1929 (Fig. 6.106). Ref.: Balech (1971a). o Epiplocylis lllldella (Ostenfeld and Schmidt, 1901)
*Rhabdonella cornucopia Kofoid and Campbell, (Fig. 6.113). Coarse reticulum located only on the
1929 (Fig. 6.107). Ref.: Balech (1962b), Souto posterior zone. Without a differentiated collar.
(1979). Incipient lip formation on the anterior zone. Ref.:
o Rhabdonella spiralis (Fol, 1881) (Fig. 6.105). Ref.: Brandt (1906-1907), Balech (1959, 1962b), Souto
Brandt (1906-1907), Balech (1959), Souto (1979). (1979).
Further studies on the whole spectrum of forms of
Rhabdonella are needed. The only consistent descrip- Genus Epiplocyloides? Hada, 1938 (Fig. 6.114, 115).
tions are those of *R. apophysata, *R. amor and R. Small, cup-shaped lorica with soft reticulum near abo-
spiralis, but the presence or absence of a hom, or even ral end. Incipient aboral hom.
the presence of a pedicel with either a skirt or a knob, *Epiplocyloides? antarctica Balech, 1962 (Fig.
or an incipient knob are not considered here to have 6.114). Oral rim sometimes with denticles.
diagnostic value (see Xystonellopsis below). General aspect resembles that of smallest species
of Cymatocylis. Ref.: Balech (1962a).
Genus Protorhabdonella? Jorgensen, 1924 (Fig. o Epiplocyloides? reticulata (Ostenfeld and Schmidt,
6.109,110). 1901) (Fig. 6.115). Coarse reticulum (similar to that
Conical, short bowl, with few to many vertical ribs. in Epiplocylis) on posterior zone of lorica and
Dubious genus comprising a total of 5 species, only expanded as thin fins towards oral end. Inner collar
two of which - P. simplex (Cleve, 1900) and P. ven- differentiated. Ref.: Brandt (1906-1907).
tricosa (Schmidt, 1901) - should be considered valido
o Protorhabdonella? curta (Cleve, 1901) (Fig. Genus Epicancella Kofoid and Campbell, 1939 (Fig.
6.109). Validity of Protorhabdonella? curta is 6.116).
doubtful because presence, number (approx. 20), Subconicallorica with circumoral band (=Iow collar).
and distribution of striae on lorica wall (main Lorica wall entirely covered with strong, conspicuous
diagnostic feature) may often lead to misidentifi- lattice-like network.
cations, as these delicate structures actually cover o Epicancella nervosa (Cleve, 1900) (Fig. 6.116).
entire wall and are often bifurcated or anasto- Approximately 18 strong longitudinal ribs. Ref.:
mosed and with levogyrous torsion near oral end, Kofoid and Campbell (1939).
as in Rhabdonella amor? var. indica. Ref.: Brandt
(1906-1907), Jorgensen (1924). Genus Cyttarocylis Fol, 1881 emend. Laackmann,
o Protorhabdonella? silllplex (Cleve, 1900) (Fig. 1910, emend. Jorgensen, 1924 (Fig. 6. 117).
6.11 O). Ref.: Balech (1962b). Bowl mostly conical, collar flaring as inverted trun-
cate cone (suboral shelf), generally with nuchal con-
Subgroup 4F. Lorica wall usually with a conspicu- striction; oral rim sometimes denticulate. Whole lori-
ous, aoastomosed reticulum 00 the posterior half ca wall finely sculptured with polygonal reticulum.
o Cyttarocylis cassis (Haeckel, 1873) /plagiostoma
Genus Epiplocylis Jorgensen, 1924 (Fig. 111-113). (Daday, 1887) (Fig. 6.117). This group comprises
Wall with a coarse reticulum covering the posterior C. cassis (Haeckel, 1837), C. euceclyphalus
half to 3/4 of the lorica, gradually fading towards the (Haeckel, 1887) and C. magna (Brandt, 1906).
352 Tintinnoinea
Cyttarocylis cassis/plagiostoma, forma plagiosto- GenusXystonellopsis Jorgensen, 1924 (Fig. 6.122-129).

ma (Daday, 1887) (Fig. 6.117e-h). Lorica an elon- MorphologicalIy similar to Xystonella. Oral rim not
gate to globose cone, shorter than in forma typica. channeled, aboral hom long. Pedicel (skirt+lance)
Aboral end rounded, truncate, or sometimes form- commonly present.
ing a short and thick aboral hom. Ref.: Brandt • Xystonellopsis cymatica (Brandt, 1906) (Fig.
(1906-1907), Balech (1959). 6.125). Subconical, elongate bowl. Suboral zone
Cyttarocylis cassis/plagiostoma, forma typica nov. with one incipient to welI-defined ring. With
formo (Fig. 6.117a-d). Lorica an elongate cone, var- incipient to slender pedicel. Skirt without wrin-
iable in length, with acute (or sometimes truncated) kles. Second ring observed in some individuals
aboral end. Wall sometimes with irregular fenes- (Fig. 6.125e) may be related to the formation of
trae. Ref.: Brandt (1906-1907), Balech (1959). epilorica. Ref.: Brandt (1906-1907), Laackmann
(1910), Jorgensen (1924).
Subgroup 4G. Lorica wall with regular, hexagonal • Xystonellopsis favata (Brandt, 1906) (Fig. 6.122).
prismatic alveoli Pedicel slightly shorter than subconical bowl.
Ref.: Brandt (1906-1907), Jorgensen (1924).
Genus Climacocylis Jorgensen, 1924, emend. Kofoid • Xystonellopsis !lastata (Biedermann, 1893) (Fig.
and CampbelI, 1929 (Fig. 6.118, 119). 6.123). Pedicel short and wrinkled. Oral end
Lorica translucent, generalIy tubular, with inconspic- toothed. Ref.: Brandt (1906-1907).
uous alveoli, partialIy to totalIy surrounded by pro- • Xystonellopsis !leros (Cleve, 1900) (Fig. 6.127).
nounced spiral tums. Sometimes with irregular open- Long, cylindrical bowl ending in thick pedicel,
ing at aboral end. always shorter than bowl. Probably the Xystonel-
Climacocylis scalaria (Brandt, 1906) (Fig. 6.119). lopsis species erected by Kofoid and CampbelI,
Spiral band formed by varying number of spiral 1929 (see p. 242: Fig. 480-485 therein) on the basis
tums extending throughout entire lorica wall. of Brandt (1906-1907) and Laackmann (1910) can
Each spiral tum shows a welI-developed spiral be lumped with X heros. Ref.: Laackmann (1910).
shelf. Broken-like aboral-end. Ref.: Jorgensen • Xystonellopsis omata (Brandt, 1906) (Fig. 6.128).
(1924), Kofoid and CampbelI (1939). Without a pedicel. Suboral area characterized by
• Climacocylis scalaroides mars!lallae CampbelI, two rings with longitudinal ridges between them.
1942 (Fig. 6.118). Lorica anteriorIy elongate; pos- Ref.: Brandt (1906-1907).
terior region an irregular cone. Spiral band located • Xystonellopsis paradoxa (Cleve, 1900) (Fig.
only at anterior region of bowl. Ref.: MarshalI 6.126). Bowl with an incipient ring on suboral
(1934), Kofoid and Campbell (1939). regíon, ending abruptly in two wrinkled skirts fol-
lowed by thick caudal lance. Ref.: Brandt (1906-
GenusXystollella Brandt, 1906 (Fig. 6.120,121). 1907), Laackmann (1910), Jorgensen (1927).
Lorica narrow and conical. Oral rim channeled. • Xystonellopsis p"lc!lra (Kofoid, 1905) (Fig.
Aboral hom or pedicel with knob or skirt, and lance at 6.129). Bowl with three to four rings located on
terminal end. Two welI-developed lamínae (see anterior half of bowl, probably related to forma-
remarks for Xystonellopsis below). tion of epilorica. Thick pedicel with large, wrin-
• Xystollella lOllgicallda (Brandt, 1906) (Fig. 6.121). kled skirt. Ref.: Brandt (1906-1907).
Noticeably smalIer than X treforti, with thin aboral • Xystonellopsis? sp. Balech, 1971 (Fig. 6.124).
hom. Ref.: Brandt (1906-1907), Balech (1959). Status of this species dubious because of absence
• Xystonella treforti (Daday, 1887) (Fig. 6.120). of pedicel; suggesting possible relationship with
Ref.: Brandt (1906-1907). Parundella, or even with Undella. Ref.: Balech
Xystollella treforti, forma coxliella nov. formo (1971a).
(Fig. 6.120b, c). Spiralled lorica. Spiral tums var- As mentioned previously for Rhabdonella, morphologi-
iable in number and size. Ref.: Kofoid (1905), cal gradation between several species of Xystonella and
Brandt (1906-1907). Xystonellopsis (mainly in the aboral hom and pedicel)
Xystonella trefom, forma typica nov. form (Fig. hinders establishment of limits of these two genera.
6.120a, d-f). Lorica long. Oral end denticulate. Brandt (1906-1907) described (Cyttarocylis?)
Pedicel with knob or skirt. Ref.: Brandt (1906-1907). Xystonella lanceolata (Plate 42, Fig. 4-8 therein, not
Tintinnoinea 353
recorded for the South Atlantic), as having a long, slen- Acanthostomel/a graci/is: Kofoid and Campbell, 1929
AcalltllOstomella lIorvegica, forma lata (Kofoid and Campbell)
der pedicel with a marked dilation on its terminal end. . stat. nov.
On the other hand, the alveolar wall texture (from hard- Acanthostomella lata: Kofoid and Campbell, 1929; Souto, 1979
Iy visible, as in X heros, to well-defined) could also Albatrossiella? jiligera (Laaekmann) Kofoid and Campbell, 1929
indicate a relationship between Xystonellopsis and Undel/aji/igera: Laaekmann, 1910
Parundella (see Fig. 6.102, 103). In summary, Albatrossiel/ajiligera: Kofoid and Campbell, 1929
AmpllOrides ampllOra (ClaparMe and Lachmann) Strand
Xystonella and Xystonellopsis are poorly defined gene-
TilltiflllUs ampllora: Claparede and Laehmann, 1858-1859; Brandt,
ra, for which reason the elassification scheme adopted 1906
herewith is provisional. Ampllorella quadri/ineata varo brandti: Jórgensen, 1924
Ampllorella ampllOra: Kofoid and Campbell, 1929; 1939; Baleeh,
Selected synonyrnic references for the taxa treated 1945, 1959
Ampllorella brandti: Kofoid and Campbell, 1929
Ampllorides ampllora: Strand, 1928; Souto, 1970a; 1979
In order to facilitate backtracking the extensive Ampllorides calida (Kofoid and Campbell) Strand
systematic rearrangements proposed in the present Tintinnus ampllora var. c: Brandt, 1906-1907
chapter, all the taxa ineluded in this review are listed Ampllorella ca/ida: Kofoid and Campbell. 1929
alphabetically (in bold characters) followed by the Ampllorides calida: Strand, 1928
Amphorides laackmalllli (Jórgensen) Strand
corresponding names used in selected previous publi-
TintiflllllS ampllora var. dada}'i: Laaekmann, 1910
cations (not boldened). Species and genera of dubious Ampllorel/a laackmanlli: Jórgensen, 1924; Kofoid and Campbell,
validity are denoted with an asterisk (*), and a question 1929
mark (?), respectively. Doubtful synonymic entTÍes are Ampllorides laac1onanlli: Sassi and Mel0, 1986
preceded with a question mark as well. Some previous AmpllOrellopsis quillquealata (Laaekmann) Baleeh
Tilltinllus quinquealatus: Laaekmann, 1907, 1910
names appear under two different species because it
Bursaopsis quinquealata: Kofoid and Campbell, 1929
was considered that the corresponding publications Ampllorel/opsis laevis: Kofoid and CampbelL 1929; Baleeh, 1971a;
incorrectly lumped the species in question. Souto, 1979
Proampllorel/a qllinquealata: Kofoid and Campbell, 1939
This listing is far from comprehensive because it does Ampllorel/opsis quinquealata: Baleeh. 1971a; Souto, 1979
AmpllOrides quadrililleata (ClaparMe and Laehmann) Strand
not cover all the available literature. It does, however,
TilltiflllUs quadri/ineatus: Claparede and Laehmann, 1858-1859
inelude most of the publications dealing with South Ampllorella quadrilineata: Jórgensen, 1924: Kofoid and CampbelL
Atlantic materials, as well as several others from dif- 1929,1939
ferent oceanic regions which were considered of spe- Ampllorides quadri/ineata: Strand. 1928; Souto, 1970a, 1972, 1979
cial significance (in particular Brandt, 1906-1907; Amphorides quadrililleata varo brasiliellsis (Laaekmann)
Tintinnus ampllora var. brasi/iellsis: Laaekmann, 1910
Laackmann, 1910; Jorgensen, 1924; Kofoid and
Amphorides quadrililleata varo millor Jórgensen
Campbell, 1929). Since Kofoid and Campbell's Ampllorel/a quadri/ineata var. minor: JÓrgensen. 1924;
(1929) elassification system has long been a standard Baleeh, 1959
in both systematic and ecologic studies, special emp- Ampllorel/a minor: Kofoid and Campbell, 1929; Baleeh, 1945
hasis was put not only on general nomenelatorial AmpllOrel/a brandti: Hada, 1932b
Ampllorides quadri/ineata var. minor: Souto, 1979
changes, but also on the large number of their species
*Amplectella mOllocollaria (Laaekmann) Kofoid and Campbell
that are considered invalid (see above and Balech, Undel/a monocol/aria: Laaekmann. 1910
1959, 1975; Souto, 1979; Alder, 1995). Brandt's work Amplectel/a monocol/aria: Kofoid and Campbell. 1929; Baleeh.
originally appeared in two volumes published in dif- 1975; Souto, 1970b, 1979
ferent years (illustrations in 1906, descriptions in *Amplectel/a monocollaria: Baleeh, 1975
Amplectella lIuda (Kofoid and Campbell) Baleeh
1907), and is therefore cÍted as "Brandt, 1906-1907".
Undella tricol/aria: Laaekmann, 1910
Undellopsis (Undel/icricos) nuda: Kofoid and Campbell, 1939
Acallthostomella lIorvegica (Daday) nov. combo Amplectel/a nuda: Baleeh, 1975
Acallthostomella lIorvegica, forma typica nov. formo Amplectella tricol/aria (Laackmann) Baleeh
TilltiflllUs sp.: Claparede and Laehmann, 1858-1859 Undella tricol/aria: Laaekmann, 1910
Amphorella norvegica: Daday, 1887b Ulldel/opsis tricol/aria: Kofoid and Campbell, 1929
Tintinnus lIon'egicus varo a gracilis; T nO/TegiclIs varo b minutus: Amplectel/a tricol/aria: Baleeh, 1975
Brandt 1906-1907 Amplectella tridil'isa (Brandt) Baleeh
Acanthostomel/a lIorvegica: Jórgensen, 1927; Kofoid and Undel/a tridivisa: Brandt. 1906
Campbell, 1929; Baleeh. 1971 b: Souto, 1979; Alder, 1995 Amplectel/a tridil'isa: Baleeh, 1975
354 Tintinnoinea
Codonella
50 flm
oral
C. galea
/crest
* C. aspera c. sp. aff. C. olla * C. nationalis

varo e
6.5a
C. amphorella
Codonaria smooth suboral
50 flm / cone
"
C. cistellula C.? perforata

Tintinnoinea 355
Tintinnidium Poroecus Stylicauda

I 50 fJm
I 50 fJm I
.,:.'¿ Ó.~.l.
:.~~:~{
: .. ' 00. ".:
50emI
-~~?~ mI
6.9a 6.9b
6.8a
\:
''b <d
6.8b
T.
T. sp. aff.
semiciliatum
T. baleehi
6.10b
Tintinnopsis complex
I 50 fJm I
."""'y",:,',.""', ..,."',." ..,.."", ..""""""""", P.apieulatus
~.' 'r" .'.

~-:.~:;e:g;;.J r~
~~
:S:::>I)OODO ••
li' t .c::..~.
<)
'~"".a;~
ti ~
tt§1<
tí
. J', '.-
-•~:;:'~~:-".
.~.,¿ ,
o":"e._o
D
6.11a
~~".Ooe S. platensis
.Il·~d"~.o Q D
..
·o'~ 1;) \
~~'~~(;).:=:f:' .-,~.-
,Q~#:~.?:~
.~~~(\
' .:;~~ .',,"
.~
.' D. t de.
v
,.'
::?'o:O o
...
:j\)
·.OObO
;;. :>
, .0 ().
-'
~"
.1
·ob.~• Q
o
'. ba, o q> C)->
spiral t:-
o o t •• structure
•• D
t\.
.~ ~ o~
o
T. radix
o t::J.~ ,)
'<l l> <001
T. aperta D
broken-like / 1/
aboral end , Q
<>
6.14
~
:~
6.12b ~~ v,:,
C>
d '-'
6.17
, [6.15 (detail)]
T. amphistoma
T. kofoidi
T. toeantinensis
----------------------------- :, - id (detailJ(
_-------, T. nordqvisti
:--_._------------- ------- ..-----
356 Tintinnoinea
~~v"
~~b
'. Jl
a
~r,
.~
.-..4
d e f
g
T. uruguayensis
6.20
T. baltiea 6.19
T. amphora/ broken-I ike
nueula 6.18 oral end
[6.22c (detail)J
T. e./b. varo mortensen;; 6.22b

T. fimbriata
T. eampanula/
T. e./b. varo butsch/ii b U tsehl""
" 6.21
a d e f g h k
T. parvula 6.23
T. beroidea
6.26
T. lobianeoi/ brasiliensis
T. pusilla/laevis ~
b .
6.24 ~
T. parva 6.29
~~~v e d e
morphological transition
between aboral ends
b
6.28
6.27
T. prowazeki
Tintinnoinea 357
delailof
oral end Helicostomella 50 pm
6.30h
lorica wall wilh

anaslomosed
W'
slriae
}~' ..
, .. '
Helicostomella subulata
lorica
wall 6.30i
lexlure 6.30d
Laackmanniella
'"
I 50 fJl11 1
.~
.2
'i5. Laackmanniella
6.30c '"
naviculaefera
:§
6030b
6.30a
ro
ou
[6.31dJ 6.31e
\;......
6.31c L. n. forma 6.31f
prolongata L. n. forma
.-------------------- L. n. forma naviculaefera -------------------.----------~ coxliella
358 Tintinnoinea
CodonelJopsis
50 pm
.1
6.32d
C. orthoceras
C. morchella
C. m. forma
typica * C. brasiliensis
Tintinnoinea 359
.. • ~ .,•..•
.•...c.p.~~..,.~ Codonellopsis
I 50 I fJm
'C ",_.~'::::
oq~'b_._.~"'.'"
•• o ¡J':'.G - Q.
. 0: ..)~.. 4
6.35a 6.35b
C. obesa
Metacylis
50Dm
C. pusilla 6.36
M. jorgenseni
6.40
O ~.. 6.41
Metaeylis sp. aff.

M. meresehkowskyi
@
r
6.38a
I
h~
C. baleehi
1,
,,~~I~
I a
~( O
c. lusitaniea 6.37
~=~
~~" ,
i
1-- ..
t-.- _
1
~---
l
=--
~:::.-=--
~-@-.a
1::':.:-.-.. ~.'
" f
¡;-~~~l
n\·: '/. I,¡ (¿
l.' '~:'C.'.".j. I/ \ J t .
c. g. forma
cylindroconica ;"/Y:'~A
; /; ~·"n I
~~I'
'''~I¡•. ·,~rIn ' .
'\~f:=3/
l " I
1/ \,1
I
Stenosemella \ .•.. ,...
{¡ /
I 1
;¡: 1;
-- / / ./
50 fJm 6.39c
~
~
/ 1- )
\ '- /
, II 6.39d
1\ 6.3ge
\
('
6.39f t: (
C. g. forma c. g. forma
6.42b c. g. forma globosa typica
coxliella
S.avellana detail af aperculum C. gaussi
360 Tintinnoinea
Dictyocysta
SOfJm
collar with
quadrangular-
pentagonal
fenestrae
bowl with
subcircular
fenestrae
D. mitra 6.43f
D. elegans
collar with 2
rows of subcircular-
pentagonal fenestrae
6.44c 6.44f
D. e. varo speciosa 6.44e
wall with
subcircular
toreticufation
hexagonal
D. duplex
denticulate oral
margin
D. e. varo lepida
6.44g
* D. californiensis
6.44n
Tintinnoinea 361
Petalotrieha
50 fJm
6.47b P. ampulla
Dadayiella
50 fJm oral end Stelidiella
incomplete 50 fJm
collar d· I ~\ . I
tacets
longitu. Ina ~I I
, I
subequal triangular
slight nuchalj
constnctlon denticles \
slender
pedicel
6.48a
D. ganymedes equidistant
longitudinal
fins
suboral
r-;nn
Brandtiella
50 fJm
." gelatinous external

.~ coating with added
• particles
6.50
S. stelidium
B. paJliata
362 Tintinnoinea
Amphorellopsis Ormosella
~ I 50fJm I
50 fJm
((
1/
II
1
\t6S1d
I
\ I!
\ I I
6.51 a
acute 6.57a \ 11
aboral end A. quinquea/ata
constriction \ IJ
~ nuchal )\ 111
delicate, \ b
sometimes poorly 6.57
Amphorides visible longitudinal
facels
50 fJm O. acantharus
O. cornucopia
Steenstrupiella
6.52 50 fJm
A. ea/ida
A. quadrilineata 1 \
varo brasiliensis
truncated
aboral end 6.60
A. /aackmanni A. amphora S. steenstrupii * S. pozzi O. haecke/i
Tintinnoinea 363
Salpingella
50 IJm
oral margín with

tooth-like lobules
6.61c
6.61b
6.61 h
5. a. varo secata
Salpingella acuminata
6.61e
lobulated !~
5. a. varo 6.61 f
oral margin
6.61g
glockentogeri
5. a. varo
undata
,
I
6.62c
16.62d
6.62a
6.62b
r¡~
~well lo",',"d',,'
defined~'
fins
..
\ '
J \J
6.63a 6.63b
6.64
5. a. varo
undata S. costata 6.65
5. a. varo secata
\1 S. subconica
S. acumimitoides S. regulata
364 Tintinnoinea
Eutintinnus
50 IJm
f----I
6.66d 6.67a 6.67b 6.67c

6.68a 6.68b
f. /usus-undae f. medius
open aboral
end
funnel-shaped
f. fraknoi / oral end

6.66c
6.66a 6.66b
6.69b
Clevea?
50 IJm
f----I
'.~'
6.71
f. apertus
, .•
I 6.70
\ 1 6.69a
f. colligatus
6.72b scalloped
* f. rugosus
6.72a C.? melchersi aboral end
Daturella
50 IJm
f----I
6.74a
op<o ,b",1 eod 1 _ D. datura 6.74c

Tintinnoinea 365
Acanthostomella
50pm
A. norvegica
detai I of teeth
A. n. forma
A. n. forma typica
lata
6.75b
lorica wall
texture
Ascampbeliella
I 50 pm
suboral
+-- ~~~I~rr~ ~. platform
;'i"~'f;'
/' \
A. obscura' *A. acuta A. urceolata * A. armilla
Favella
50 pm
~
annuli
6.81b~
6.80 * F. arcuata aboral hom

F. taraikaensis 6.81a
366 Tintinnoinea
Favella
50 IJm
1-----1
spiral
turns
F. e. forma favella
6.82b
epilorica
F. ehrenbergii
overlapping
splraled wan
F. e. forma favella 6.82h 6.821

F. e. forma decipiens F. e. forma coxliella
* F. serrata
detailoí
denticles
6.83b
Tintinnoinea 367
Cymatocylis 50 IJm
f------i C. drygalskii
. .",
r
\-.
''R~ y--~ wavy to smooth denticulate
outer collar inner collar
\
/
(1 6.84b
6.84a
C. c. forma affinis
C. c. forma
coxliella
6.84c
, 6.84d
f-=-
C. convallaria
C. c. forma
typica
----
l='.--=:= 6.86c
C. antarctica C. d. forma
typica
f:-':''l
! ---1
6.85a " \:::J 6.85b

C. a. forma C. a. forma C. d. forma
ovata C. d. forma
typica coxliella
!
coxliella
~!
! ~
r===-~~_.l
I
~~-------"._-~
everted-
denticulated
oral end
_~ ~
~~~ ... ..•...
I!
"
,¡
1:
¡i
I ¡i :¡
c. v. f?rma
'1
1 ! 6.87a
{lava
,¡ 6.87b
typlCa
C. v. forma
,¡'j
J
c. v. forma
6.87c
--
/6.87d
"
i
I
I
I
"
6.88
coxliella C. v. forma C. nobilis

subrotundat,
J, f
~ 6.87e
C. vanhoffeni
~
i c. v. forma
c. v. forma
conica cylindrica
lorica wall
.,,,~jtexture
whole , 6.89b 6.90
specimen 6.89a
(notto 6.91
scale)
C. calyciformis * C. kerguelensis * C. cristallina
368 Tintinnoinea
Undella
50 fJm
f----I
acute
aboral-
end
rounded
6.93 6.94 6 .95 o aboral
end
U. hemisphaerica * U. grandis Undella c1aparedei group
,
OOOOQQQQQQQmOO
b Undefla c1aparedei gm"p 6.95 "
Amplectella bílamínate
Undellopsis ~
I I
50fJm ¡Wall \ yY ledge ~
suboral
J
~ c::-
~ríngs
6.99
A. tridivisa 6.97 A. trieo/~:r~a A. nuda
* A. monoeollaria U. marsupialis
Albatrossiella? Parundella ~
_ bílaminate
wall
6.103d
P. messinensis 6.103b 6.103c
6.102
6.103a
)' 6.101
A.? filigera P. eaudata
Tintinnoinea 369
-------------,
Rhabdonella
50 ~m
1iI~
fenestrae
R. spiralis
varo henseni :11
1'1 6.105b
* R. amor
pedicel
6.107
knob
6.105a * R. cornucopia
lance R. spiralis
~
* R. apophysata
6.104b
* R. amor
varo
valdestriata
Protorhabdonella?
50 pln 6.108h
7-9
strlae,~ ,,<.
longitudinal _,.
/6.110
P. simplex
P.? curta
6.108f :
370 Tintinnoinea
Epiplocylis
50 fJm
lamina
outer~~
inner
lamina
6.112a
lip-shaped suboral
collar~ zone f. calyx?
f. acuminata
lorica wall
texture
.
[6.111 el
lorica wall texture f. undella

(anterior zone)
6.113a
, 6.113e
6.113c 6.113d
Epiplocyloides? Epican cella

50 fJm
50 fJm
lorica wall
texture
reticu lated
sculpture
6.116
* f.? antarctica f.? reticulata

f. nervosa
Tintinnoinea 371
denticulated oral 50flm suboral

Cyttarocylis f-------l
margin
undulated oral
/' shelf
/.
margin
nuchal
constriction
6.117f
6.117h
c. c./p. forma c. c/p. forma

typica plagios toma
C. eassis/p/agiostoma
Climacocylis
50 flm
spiral
l:iands
smooth oral
margin
spiral
l:iands
6.119b
6.118
C. sea/aroides irreg,ular
aboral broken-like
end
6.119a C. sea/aria
372 Tintinnoinea
outer lamina
Xys ton ella (dentieu lated
ehanneled oral margin
50 ¡Jm oral rim)
inner lamina
(smooth oral
rim)
loriea wall
texture
-+-- knob
-+-- lance
X. 1. forma
6.120a coxliella
X. 1.forma typica
smooth outer undulated inner
lamina ~ lam~
loriea wall X. longicauda X. treforti

6.121 texture
Tintinnoinea 373
Xystonellopsis subequal
50 fJm
-r- teeth
,.A. ~I\ ~ outer
\ ,
J -;¿Iamina
\
LinCiPient
;/ L C1{L 1 I ring
('"1 /
-~ ;ooe"'m;" ~
ti
. lorica wall
texture
I
1
I
\ ~.124 ); XystonelJopsis? sp.
X. favata skirt
~'\l'
v
epilorica? ~
skirt
6.123
ring
~ 6.122 ~
lance~
incipient /\ t
y ring 1I
Ji
ring
1I
!
!
,1
\.~ ¡
1I
¡j
lorica wall
texture
1
\~
\\
l\
I~I 1 6.125e
\
6.125b Ir 6.125c 6.125d
6.125a
X. cymatica
374 Tintinnoinea
Xys ton ellopsis

I SOmm I
ring~
x. paradoxa
suboral
zone
6.127 X. pulchra
lance
x. heros
X.ornata
..
Tintinnoinea 375
..
Fig. 6. Tintinnid taxa reeorded for the South Atlantie Deean.
Figures within eaeh genus are reprodueed at the same magnifieation (seale), exeept for those whose number is included in square braekets (e.g.,
[6.1ge]). Speeies and genera ofdubious validity are denoted with an asterisk (*), and a question mark (?), respeetively.
Figure sourees (original names are given in [square braekets]):
From Alder (1995): 39f(as Codonellopsis gaussi, forma coxliella]; 85b [as Cymatocylis antarctica, forma coxliella].
From Baleeh (1944): 30i [as Helicostomella Lemairei]; 109 [as Protorhabdonella curta].
From Baleeh (1945): Ila [as Tintinnopsis platensis]; 12b [as Tintinnopsis radix]; 1ge [as Tintinnopsis baltica]; 23j, k [as Tintinnopsis gracilis];
24a [as Tintinnopsis lobiancoi]; 24d, e [as Tintinnopsis lata]; 36e [as Codonellopsis contracta]; 67b [as Eutintinnus australis]; 80 [as Favella
quequenense] .
From Baleeh (1947); 86a [as Cymatocylis dl)'galskii].
From Baleeh (1948); l3b, e [as Tintinnopsis aperta]; 14a-e [as Tintinnapsis tacantinensis]; 16 [as Tintinnopsis kofoidi]; 19a, d, f[as Tintinnopsis
baltica]; 20a-e [as Tintinnopsis lIrllguayensis]; 21a [as Tintinnopsis biitschlii]; 21b [as Tintinnopsis biitschlii, var. martensenii]; 22a [as
Codonariafimbriata]; 23e, d [as Tintinnopsis gracilis]; 23e-g [as Tintinnopsis parvula]; 29a, b [as Tintinnopsis glans]; 2ge-f [as TintillllOpsis
parva]; 29g, h [as Tintinnopsis turbinata]; 35e-e [as Codonellopsis obesa]; 37e, d [as Codonellopsis lusitanica]; 42a-e [as Stenosemella avella-
na]; 69a, b [as ElIIintinnus rugoslIs]; na, b [as Clevea me!chersi].
From Baleeh (1951): 11b [as Stylicallda platensis]; 17 [as Tintinnopsis amphistoma]; 30e-g [as Metacylis allllLl/ifera]; 35a, b [as Codonellopsis obesa].
From Baleeh (1962a): 114 [as Epiplocyloides antarctica].
From Baleeh (197Ia): 33d [as Codonella morchella]; 36a-d [as Codonellopsis pllsilla]; 38a [as Codonellopsis antarctica]; 46 [as Dictyocysta
californiensis]; 51e [as Amphorellopsis qllinquealata]; 51d [as Amphorellopsis laevis; 57a, b [as Amphorellopsis acantharus]; 58 [as Ormosella
haeckeli]; 60 [as Steenstrupiella pozzi]; 63a, b [as Salpingella subconica]; 75b, e [as Acanthostomella norvegica]; 103a [as Panllldella acule-
ata longa]; 103b [as Parundella caudata]; 106 [as Rhabdonella chiliensis]; 124 [as Xystonellopsis sp.].
From Baleeh (197Ib): 78 [as Ascampbeliella lIrceolata], 79 [as Ascampbeliella armilla].
From Baleeh (1975): 95a-e [as Undella c1aparedei]; 95f-h [as Undella sllbcalldata subcaudata]; 95i, j [as Undella globosa]; 95k, m, n [as
Undella pelpusilla]; 951 [as Undella subacuta].
From Barría de Cao (1981): 8a, b [as Tintinnidium balechi]; 9a, b [as Tintinnidium sp. aff. semiciliatum]; 18b, e [as Tintinnopsis amphora]; 23a,
b [as Tintinnopsis parvllla]; 23i [as Tintinnopsis gracilis]; 24b [as Tintinnopsis karajacensis]; 25 [as Tintinnopsis compren'a]; 28a [as
Tintinnopsis pusilla]; 28b [as Tintinnopsis levigata]; 28e-e [as transitions]; 37a, b [as Tintinnopsis lusitanica].
From Barría de Cao (1986): 41 [as Metacylis sp. aff. mereschkows/"yi].
From Brandt (1906-1907): la-d [as Codonella nationalis]; 2a (as Codonella galea]; 2b [as Codonella galea, var. a]; 3 [as Codonella nationalis, varo
e]; 4 [as Codonella nationalis, var. e]; 5e, d, [as Codonella amphorella]; 5e [as Codonella nationalis, var. b]; 5f [as Codonella amphorella, var. a];
6a [as Codonella peliorata, var. a]; 6b [as Codonella peliorata]; 7a, b [as Codonella cistellula, var. a oceanica]; lOa [as Cyttarocylis apiculata];
12a, e [as Tintinnopsisfracta]; 12d [as Tintinnopsis davidoffi, var. cylindrica]; l3a [as Tintinnopsis aperta]; 14d [as Tintinnopsis aperta, var. a]; 15
[as Tintinnopsis nordqvisti]; 18a [as Tintinnopsis lobiancoi, var.filsiformis]; 18d [as Tintinnopsis nucula]; 19b [as Tintinnopsis baltica, var. a]; 1ge,
g [as Tintinnopsis baltica]; 22b, e [as TintillllOpsis sp.]; 23h [as Tintinnopsis karajacensis, var. a]; 24e [as Tintinnopsis sp.]; 26 [as TintillllOpsis beroi-
dea var. b]; 30a-d [as Tintinnlls sllbulatlls]; 30h [as Tintinnus mediterranells, var. longa]; 32a, d [as Codonella orthoceras, var. a tessellata]; 32b
[as Codonella orthoceras, var. i]; 32e [as Codonella orthoceras, var. h]; 32e, g [as Codonella orthoceras, var. gpura]; 32f, h [as Codonella O/·tho-
ceras, var. fminor]; 33a-e, fEas Codonella morchella]; 33h [as Codonella morchella, var. erythriiensis]; 33g, e [as Codonella morchella, var. scha-
bi]; 34a, b [as Codonella ecaudata, var. brasiliemis]; 40 [as Tintinnlls urceolatus]; 43a, b [as Dictyocysta mitra]; 43e [as Dictyocysta mitra, var. b];
43d, e [as Dictyocysta mitra, var. e]; 44a-e [as Dictyocysta elegans, var. e miilleri]; 44d [as Dictyocysta elegans, var. d]; 44g- i [as Dictyocysta temp-
hlln]; 45 [as Dictyocysta templllln, var. d]; 47a [as Petalotricha ampulla, var. b]; 47b [as Petalotricha ampulla, var. d]; 47e [as Petalotricha amp"l-
la, var. e]; 48a [as Tintinnlls ganymedes]; 48b [as Tintinnus blllbosus, var. a]; 48e [as Tintinnus blllboslls]; 49 [as Tintinnus palliatlls]; 50 [as
Tintinnlls stelidium]; 52 [as Tintinnus amphora, var. e]; 55 [as Tintinnlls amphora]; 56 [as Tintinnus conicus]; 59 [as Tintinnus steenstrupii]; 6la-e
[as Tintilllllls acuminatlls]; 61d [as Tintinnus acuminatlls, var. secata]; 61e, f [as Tintinnlls acuminatus, var. e glockentogeri]; 61g [as TintillllllS
lllldatus]; 61h [as Tintinnlls lllldatus, var. unguiculata] ; 65 [as Tintinlllls regulatlls]; 66b, d [as Tintinnusfraknoi, var. a]; 66e [as Tintinnusfra/,.?lOi];
67 [as Tintinnus lusus-lIndae]; 70 [as Tintinnus IlISlls-lllldae var. e]; 71 [as Tintinnus IlIslls-undae, var. a tublllosa]; 73a [Tintinnus emarginatus];
74a, b [as Tintinnus datura]; 75a [as Tintinnlls norvegicus, var. a gracilis]; 76 [Cyttarocylis obscura]; 81a, b [as Cyttarocylis arcllata]; 82e, d [as
Cyttarocylis ehrenbergi]; 83a, b [as Cyttarocylis serrata]; 92a-e [as Undella hyalina var. a]; 96 [as Undella tridivisa]; 100a, b [as Undella marSll-
pialis]; 102 [as Undella messinensis]; 103e, d [Undella messinensis, var. a]; l04a [as Ptychocylis (Rhabdonella) apophysata, var. b]; 104b [as
Ptychocylis (Rhabdonella) apophysata]; 105a [as Ptychocylis (Rhabdonella) spiralis]; 105b[as PtycllOCylis (Rhabdonella) ~piralis, var. e henseni];
108b, e [as Ptychocylis (Rhabdonella) amor]; 108g [as Ptychocylis (Rhabdonella) amor, var. a cllspidata]; 108h [as Ptychocylis (Rhabdonella)
amor, var. b valdestriata]; lila [as Ptychocylis acuminata, var. e]; 112a [as Ptychocylis calyx, var. a]; 1¡2b [as Ptychocylis calyx, var. b]; 113a [as
Ptychocylis IlIldella, var. b]; ll3b [as Ptychocylis undella, var. a]; 113e [as Ptychocylis lIndella var. f]; 113e [as Ptychocylis lIndella, var. k]; l13d
[as Ptychocylis IlIldella, var. h]; 115 [as Ptychocylis reticulata]; 116 [as Ptychocylis nervosa]; 117a [as Cyttarocylis cassis, var. e magna]; 117b [as
Cyttarocylis cassis, var. d]; 117e [as Cyttarocylis cassis, var. a]; 117d [as Cyttarocylis cassis, var. b conica]; 117e [as Cyttarocylis cassis, var. e];
117f [as Cyttarocylis plagiostoma, var. e]; 117g [as Cyttarocylis plagiostoma, var. a]; 117h [as Cyttarocylis plagiostoma, var. b]; 119a, b [as
Cyttarocylis (Coxliella) scalarills]; 120e [as Cyttarocylis (Coxliella)fasciata, var. procera]; 120d-f [as Cyttarocylis (Xystonella) treforti]; 121 [as
Cyttarocylis (Xystonella) aCIIS,var. b longicauda]; 122 [as Cyttarocylis? (Xystonella)fal'Ota]; 123 [as Cyttarocylis (Xystonella) hastata]; 125a [as
Cyttarocylis (Xystonella) cymatica, var. e ~picata]; 125e [as Cyttarocylis (Xystonella) cymatica]; 125e [as Cyttarocylis (.;'(ystonella) dicymatica];
126b, e [as Cyttarocylis? (Xystonella) paradoxa]; 128 [as Cyttarocylis (Xystonella) ornata]; 129 [as Cyttarocylis (Xystonella) p"lchra].
376 Tintinnoinea
*Aseampbeliella aeuta (Kofoid and CampbeU) CorJiss Codonaria cistellu/a, forma IJ'Pica; C. cistellu/a, forma major:
Craterella aeuta; C. protuberans: Kofoid and Campbell, 1929 Ba1eeh, 1959
Aseampbeliella acuta: Corliss, 1960; Souto, 1979 Codollella ampllOrella Biedermann
*Aseampbeliella armilla (Kofoid and CampbeU) CorJiss Codonella amphorella: Biedermann, 1893; Brandt, 1906-1907;
Craterella armilla: Kofoid and Campbell, 1929 Laaekmann, 1910; Kofoid and Campbell, 1929, 1939; Baleeh,
Aseampbeliella armilla: Corliss, 1960; Baleeh, 1971b; Souto, 1979 1962b; Souto, 1979
Aseampbeliella obscura (Brandt) CorJiss Codonella amphorella varo a; C. Ilationalis varo b: Brandt, 1906-1907
Cyttarocylis obscura: Brandt, 1906-1907 Codonella acerca: Jorgensen, 1924
Craterella obscura: Kofoid and Campbell, 1929 Codonella rapa; C. recta: Kofoid and Campbell, 1929
Ascampbe/iella obscura: Corliss, 1960 *Codollella aspera Kofoid and CampbeU
Aseampbeliella ureeolata (Ostenfe1d) Corliss Codonella ga/ea: Fol, 1884; Brandt, 1906-1907
Tintinnus urceo/atus: Ostenfe1d, I 899a, b Codonella ga/ea var. a; C. ga/ea varo b: Brandt, 1906-1907
Amphorella urceo/ata: Ostenfe1d and Sehmídt, 1901 Codonella aspera: Kofoid and Campbell, 1929; Baleeh, 1959;
Craterella urceo/ata: Kofoid and Campbell, 1929 Souto,1979.
Ascampbeliella urceo/ata: Corliss, 1960; Baleeh, 1971b; Souto, 1979 Codollella e/ongata: Kofoid and Campbell, 1929, 1939
Brandtiella palliata (Brandt) Kofoid and CampbeU Codonella galea Haeekel
Tintinnus palliatus: Brandt, 1906-1907; Laaekmann, 1910 Codonella ga/ea: Haeekel, 1873; Kofoid and Campbell, 1929;
Brandtiella palliata: Kofoid and Campbell, 1929; Souto, 1979 Ba1eeh, 1959.
C/evea? melehersi Baleeh Codonella nationalis; C. nationalis var. a; C. nationalis var. d:
C/evea me/chersi: Baleeh, 1948; Souto, 1979 Brandt, 1906-1907
Climaeoeylis sealaria (Brandt) Jorgensen Codonella inflata; Codonella cuspidata: Kofoid and Campbell, 1929
Cyttaroc)'lis (Coxliella) sca/arius: Brandt, 1906 *Codollella Ilatiollalis varo e Brandt
Climacoc)'lis sca/aria: J6rgensen, 1924 Codonella nalionalis var. e: Brandt, 1906-1907
C1imaeoc)'lis sca/aria: Kofoid and Campbell, 1929, 1939 Codonella pocu/um; C. robusta: Kofoid and Campbell, 1929.
Climaeoeylis sealaroides marshallae CampbelJ Codollella? perforata Entz Sr.
Climacocy/is sca/aroides: Marshall, 1934 Codonella pelforata: Entz, Sr., 1884; Brandt, 1906-1907; Kofoid
Climacoc)'lis sca/aroides varo marshallae: CampbelJ, 1942; Baleeh, and Campbell, 1929; Baleeh, 1962b.
1964; Souto, 1970a, 1970b Codonella perforata var. a; C. perforata var. b: Brandt, 1906-1907
C1imacocy/is sca/aroides marshallae: Souto, 1979 Codonella apicata: Kofoid and Campbell, 1929
Codonaria cistellula (Fol) Kofoid and CampbelJ Codonella sp. aff. C. olla Kofoid and CampbelJ
Cyttarocylis cistellu/a: Fol, 1884; Daday, 1887b Codonella sp. afT. C. olla: Baleeh, 1962b
Codonella cistellu/a: Brandt, 1906-1907; Kofoid and Campbell, 1929 Codonella nationalis var. c: Brandt, 1906-1907
Codonella cistellu/a varo a oceanica; C. cistellu/a varo b; C. ciste/- ? Codonella olla: Kofoid and Campbell, 1929
/ulla var. c: Brandt, 1906-1907 Codollellopsis baleehi Hada
Codonella bengue/ensis; C. oceanica; C. austra/is; C. mucronata; Codonellopsisfi-igida: Baleeh, 1958a
C. lata; C. dada)'i: Kofoid and Campbell, 1929 Codonellopsis ba/echi: Hada, 1970; Souto, 1979; Hermosílla, 1977;
Codonaria cistellu/a; C. australis; C. bengue/ensis; C. lata; C. Alder, 1995
mucronata; C. oceanica: Kofoid and Campbell, 1939 Codonellopsis antarctica: Baleeh, 1971 a; Souto, 1972
••
Figure 6 eaption eontinued.
From Faria y Da Cunha (1917): 27 [as Tintinnopsis proll'azeki]; 82a [as C)'ttarocylis ehrenbergii var. c1aparedii]; 82e [as Cyttarocylis
(Coxliella) helicoideia]'
From Laaekmann (1910): 5b [as Codonella amphorella]; 31a-e [as Leprotintinnus navicu/ae(ems]; 31e [as Leprotintinnus pr%ngatus]; 31 f [as
Coxliella minor]; 39a [as Leprotintinnus forma conica]; 39b, e [as LeprotintillllLls]; 39d, e [as Leprotintinnus gaussi]; 44j [Dictyocysta temp/um,
var. h]; 44k, m [as DiCIJ'ocysta temp/um, var. indica]; 51a, b [as Tintinlltls quinquea/atus]; 53 [as Tilllinnus amphora, var. brasiliellsis]; 54 [as
Tintinnus amphora varo dadayi]; 62a [as Tintinnus acuminatoides]; 62b, e [as Tintinnus acuminatoides, var. secata]; 62d [as Tintinnus unda-
tus]; 64 [as Tintinnus costatus]; 68a [as Tintinnus /usus-undae]; 74e [as Tintinnus datura]; 84a, b [as C)'matocy/is affinis, forma typica]; 84e [as
Coxliellafrigida]; 84d [as Cymatocylis convallaria]; 85a [as Cymatocylis parva]; 86b [as Coxliella intermedia]; 86e [as Cymatoc)'lis dIJ'ga/s-
kii, forma ~'Pica]; 87a [as C)'matocylis vanh6fJeni, forma typica]; 87b [as Cymatocylisflava, forma typica]; 87e [as Coxliellafrigida]; 87d [as
C)'matocylis flava, forma subrotundata]; 87e [as Cymatoc)'/is vanh6fJeni, forma conica]; 87f [as Cymatocylis flal'Q, forma cylindrica]; 88 [as
C)'matocy/is nobilis, forma IJ'Pica]; 89a [as Cymatocylis ca/yciformis, forma typica]; 89b [as Cymatocy/is ca/yciformis, forma brevicaudata]; 90
[as C)'matocylis kergue/ensis]; 91 [as Cymatocylis cristallina, forma IJ'Pica]; 93 [as Undella hemisphaerica]; 94 [as Undella c1aparedei, var.
grandis]; 97 [as Undella monocollaria]; 98, 99 [as Undella tricollaria]; 101 [as Undellajiligera]; 108a, e, d [as Rhabdonella amor, var. indi-
ca]; 108f [as Rhabdonella amor]; 110 [as Rhabdonella amor, var. simp/ex]; ¡llb [as Ptychocylis acuminata var. ?]; 125b [as Xystonella cyma-
liea]; 125d [as X)'stonella dicymatica]; 126a [as Xystonella paradoxa]; 127 [as Undella heros, var. gaussi].
From Lava1-Peuto (1983): 82f [as Favella ehrenbergii, forma coxliella]; 82g, h [as Favella ehrenbergii, forma decipiens].
From Souto (1979): 75d [as Acanthostomella lata]; 77 [as Ascampbeliella acuta].
Original: 5a, 10b, 31d, 38b, 39f, 43f, 44e, f, n, 66a; 67e; 68b; 85b; 950, p; 107; lile; 118; 120a, b.
Tintinnoinea 377
*Codol/ellop.\·is brasiliel/sis (Brandt) Kofoid and Campbell Cymatocylis convallaria Laackmann nov. comb
Codol/ella ecaudata var. brasiliel/sis: Brandt, 1906-1907 CYlllatocylis cOl/vallaria, forma typica Laackmann
Codol/ellopsis brasiliel/sis: Kofoid and Campbell, 1929 Cymatoeylis convallaria: Laaekmann, 1910; Kofoid and Campbell,
Codol/ellopsis gallssi (Laackmann) nov. combo 1929; Ba1eeh, 1947; Sassi and Me10, 1986
Codol/ellopsis gallssi, forma cylil/drocol/ica nov. formo Cymatocylis eOIll'allaria, forma f)pica: Laaekmann, 1910; Alder,
Leprotil/til/I/us glacialis; Leprotil/til/I/us?, forma cOl/ica; Leprotin- 1995
tinnus: Laaekmann, 1910 CYlllatocylis cOl/vallaria, forma affil/is Laackmann stat. nov.
Codonellopsis glacialis: Kofoid and Campbell, 1929 Cyma/ocylis affinis: Laaekmann, 1910; Kofoid and Campbell,
Codonellopsis gaussi, forma cylindroconica: Alder, 1995 1929; Baleeh, 1958b
Codol/ellopsis gallssi, forma coxliella nov. formo Cymatoeylis aJJinis, forma typiea; C. affinis. forma eylindriea; C.
Coxliella mil/or: Baleeh, 1973 affinis, forma umula; C. affinis, forma ven/rieosa: Laaekmann,
Codonellopsis gaussi, forma coxliella: Alder, 1995 1910
Codol/ellopsis gallssi, forma globosa nov. formo Cyma/oeylis gaussi; C. sC)plllls; Cyma/oeylis urnula: Kofoid and
Codonellopsis gaussi: Baleeh, 1973 Campbell, 1929
Codonellopsis gaussi, forma globosa: Alder, 1995 Cymatoeylis convallaria, forma affinis: Alder, 1995
Codol/ellopsis gallssi, forma typica nov. formo CYlllatocylis cOl/vallaria, forma coxliella nov. form
LeprotintillllUs gaussi: Laaekmann, 1910 Cylfaroeylis (Coxliella)ji-igida: Laaekmann, 1907
Codonellopsis gaussi: Kofoid and Campbell, 1929 Coxliella ji-igida: Laackmann, 1910; Kofoid and Campbell, 1929;
Codonellopsis gaussi, forma typica: Alder, 1995 Baleeh, 1958b; Sassi and Melo, 1986
Codol/ellopsis IlIsital/ica Jorgensen Coxliellaji-igida, forma I)pica: Laaekmann, 1910
Codol/ellopsis lusital/ica: Jórgensen, 1924; Kofoid and Campbell, Coxliella eyma/ioeoides: Kofoid and Campbell, 1929; Ba1eeh, 1973
1929; Baleeh, 1948; SoulO, 1979; Barría de Cao, 1981 Cyma/oeylis eonl'Qllaria, forma coxliella: Alder, 1995
Codol/ellopsis morchella (Cleve) nov. combo CYlllatocylis al/tarctica (Cleve) Kofoid and Campbell
Codol/ellopsis morchella, forma typica nov. formo CYlllatocylis al/tarctica, forma typica nov. formo
Codonella morchella: Cleve, 1900b; Brandt, 1906-1907 Amphorella (?) antaretica: Cleve, 1901
Codonella eeaudata: Brandl, 1906-1907 Tintinnus antarctieus: Brandt, 1906-1907
Codonellopsis morehella: Jórgensen, 1924; Baleeh, 1971a; Souto, Cylfarocylis parva: Laaekmann, 1907
1979 Cylfaroeylis an/aretiea: Laaekmann, 1910
Codonellopsis ecaudata: C. turgeseens; C. obeoniea: Kofoid and Cymatoeylis pan'a: Laaekmann, 1910; Kofoid and Campbell, 1929;
Campbell, 1929 Baleeh, 1958a
Codol/ellopsis lIIorchella, forma schabi Brandt stat. nov. Cymatoeylis an/are/ica: Kofoid and Campbell, 1929; Baleeh,
Codonella morehella; C. morehella var. elythriiensis, C. mord1ella 1971a; Souto, 1972, 1979; Williams et al .. 1994
var. sehabi: Brandt, 1906-1907 C)'ma/ocylis an/are/ica, forma typiea: Alder, 1995
Codonellopsis morehella: JÓrgensen. 1924 CYlllatocylis a/llarctica. forma coxliella nov. formo
Codonellopsis sehabi: Kofoid and Campbell. 1929: Baleeh, 1959; Cymatoeylis antare/iea, fonna coxliella: Alder, 1995
Souto, 1970a CYlllatocylis drygalskii Laackmann nov. comb
Codonellopsis el)'thriiensis; C. globosa; C. americana: Kofoid and Cylfaroe)'lis drygalskii: Laaekmann. 1907
Campbell, 1929 CYlllatocylis drygalskii, forma typica Laackmann
Codonellopsis indica: Kofoid and Campbell, 1939 Cyma/ocylis dl)'galskii: Laaekmann, 1910; Kofoid and Campbell,
Codol/ellopsis obesa Ba1ech 1929; Baleeh, 1947; A1der, 1995
Codonellopsis obesa: Ba1eeh, 1948, 1951; Souto, 1979 Cymatocylis dlygalskii, forma f)'piea: Laaekmann, 1910; Alder, 1995
Codol/ellopsis ortllOceras (Haeckel) Jorgensen Cyma/oeylis drygalskii, forma subro/LlIlda/a; C. dlygalskii, forma
Codonella orthoeeras: Haeekel. 1873; Brandl, 1906-1907 flava; C. dl:vgalskii, forma eylindrica: C. dl)galskii, forma \'en-
Codonella orthoeeras var. a tessellata; C. orthoeeras val. b; C. tricosa: Laaekmann, 1910
orthoceras var. e; C. orthoeeras val. d: C. orthoeeras val. e; C. Cyma/oeylis meridiana: C. cylindl'lls; C. t;pica; C. ecaudata;
orthoeeras val. f minor: C. orthoeeras val. k; C. orthoeeras Cymatocylis si/ula: Kofoid and Campbell, 1929
var. I pacifica; C. orthoeeras var. i; C. orthoeeras var. h; C. CYlllatocylis drygalskii, forma ovata Laackmann
orthoeeras var. g pura: Brandt, 1906-1907 Cymatoeylis dl)'galskii, forma ovata: Laaekmann, 1910; A1der, 1995
Codonellopsis orthoceras: Jórgensen, 1924; Kofoid and Campbell, Cyma/oeylis ovata: Kofoid and Campbell, 1929; Baleeh, 1947
1929; Baleeh, 1959; Souto, 1979 Cymatoeylis dl)'galskii: Ba1eeh, 1947
Codonellopsis minor; C. pacifica; C. pura; C. tessellata; C. longa: CYlllatocylis drygalskii, forma coxliella nov. formo
C. meridionalis: C. speeiosa; C. ealijol'lliensis; C. inflata; C. Coxliella intermedia Laaekmann 1910: Kofoid and Campbell, 1929
parm: Kofoid and Campbell, 1929, 1939 Cymatocylis dl)'galskii, forma coxliella: Alder, 1995
Codol/ellopsis p"silla (Cleve) Jorgensen Cymatocylis drygalskii. abnorma1 individuals?
Codonella pusilla: Cleve, 1900b Cymatoeylis dl)galskii: C. dl)galskii, forma coniea: C. dl)galskii,
Codonellopsis pusilla: Jórgensen, 1924; Kofoid and Campbell, forma subro/undata: Laaekmann, 1910
1929; Baleeh, 1971a; Souto, 1970a; 1972, 1979 Cymatoeylis /ubulosa; C. situla: Kofoid and Campbell. 1929
Codonellopsis lagenula: Jórgensen, 1924 Cyma/oe)'lis dl)·galskii, abnornlal individuals?: A1der, 1995
Codonellopsis contracta: Kofoid and Campbell, 1929, 1939; Cymatocylis I/obilis (Laackmann) Laackmann
Ba1eeh, 1944, 1945 Cylfaroeylis nobilis: Laaekmann, 1907
378 Tintinnoinea
Cymatocylis Ilobilis: Laackmann, 1910; Kofoid and Campbell, Cymatocylis calyciformis, fOfila 1)pica; C. cal)"ciformis, forma bre-
1929; Balech, 1958b; A1der, 1995 vicaudata; C. ca~)'cirormis, forma cylindrica: Laackmann, 1910
Cymatoc)'lis Ilobilis, forma tlpica; C. Ilobilis, forma cylilldrica; C. C)'matocylis brevicaudata: Kofoid and Campbell, 1929; Hada, 1970
Ilobilis, forma simplex; C. Ilobilis, fOfila sllbrotulldata: Laack- Cymatocylis cylindroides: Kofoid and Campbell, 1929
mann, 1910 *Cymatocylis kergllelellsis Laackmann
C)"matoc)'lisfollicullls; C. simplex: Kofoid and Campbell, 1929 ? Amphorella norl'egica: Cleve, 1901
Cymatocylis valllroffelli (Laackmann) Laackmann nov. comb Cymatocylis kerguelensis: Laackmann, 1910; Kofoid and Campbell,
Cymatocyli,~ vallllOjJelli, forma typica Laackmann 1929; Hada, 1970
Ptyclrocylis l'allhojJelli: Laackmann, 1907 *C,l'matocylis kerguelellsis: Alder, 1995
Cymatocylis vallhojJeni: Laackmann, 1910; Kofoid and Campbell, *CYlllatocylis cristallilla Laackmann
1929; Balech, 1958b; A1der, 1995 Cymatocylis cristallina: Laackmann, 1910; Kofoid and Campbell,
Cymatocylis l'allhojJelli, forma tlPica: Laackmann, 1910; Alder, 1929; Hada, 1970
1995 Cymatocylis cristallilla, forma cOllica; C. cristallina, forma typica;
Cymatocylis l'allhojJelli, forma cOllica; C. l'allhojJelli, forma sllbro- C. cristallina, forma cylindrica; C. cristallina, forma simplex;
tllndata; C. l'allho.ffelli, forma millor; C. j1am, forma I'elltricosa: C. cristallilla. fOfila ventricosa: Laackmann, 1910
Laackmann, 1910 Cymatocylis Sllbcollica; C. crassa; C. culcullus; C. glans; C. incoll-
Protoc)"matocvlis cOllicoides: Kofoid and Campbell, 1929 ' dita; C. labiosa: Kofoid and Campbell, 1929
Cymatocylis valllroffelli, formajlava Laackmann stat. nov * Cymatocylis cris/allina: Alder, 1995
Cymatocylisflam: Laackmann, 1910; Kofoid and Campbell, 1929; Cyttarocylis cassis (Haeckel)lplagiostoma Daday group
Balech, 1947 Cyttarocylis cassis, forma typica Haeckel, nov. form.
Cymatocylis .Ilm'a, forma 1)pica; C. l'allhojJelli, forma minor; C. DiC1)'oC}'stacassis: Haecke1, 1873
.flam, forma l'elltricosa: Laackmann, 1910 Cyltarocylis cassis: Fol, 1881, 1884; Brandl, 1906; Kofoid and
Cymatocylis l'allhojJelli: Laackmann, 1910; Balech, 1958b Campbell, 1929; Ba1ech, 1959; SoulO, 1979
Cymatocylis l'entricosa; C. millor; C. cylindrica: Kofoid and Cyttarocvlis cassis var. a; C. cassis var. b cOllica; C. cassis var. d:
Campbell, 1929 Brandl, 1906
Cymatocylis l'allhojJelli, formajlal'a: A1der, 1995 C)'ttaroc)'lis cassis var. c maglla: Brandl, 1906; Jorgensen, 1924
Cymatocylis vallllOjJelli, forma sllbrotlllldata Laackmann Cyttarocylis conica: Kofoid and Campbell, 1929
Cymatoc)'lis vallhojJeni, forma sllbrotllndata: Laackmann, 1910; Cyttarocylis magna: Kofoid and Campbell, 1929; Balech, 1959
Alder,1995 Cyttarocylis cassis, forma plagiostoma Daday stat. nov.
Cymatocylis.flal'a, fOfila sllbrotlllldata: Laackmann, 1910 Stethocephallls ellceClyphalus: Haecke1, 1887
Protocymatoc)'lis sllbrotulldata; P. vas; Cymatoc)'lis digitablllum; Cyttarocylis cassis var. plagiostoma: Daday, 1887b
C. digitullls: Kofoid and Campbell, 1929 Cyttarocylis cassis var. e: Brandl, 1906
Cymatocylis valllroffelli, forma cOllica Laackmann Cyttarocylis plagiostoma: Brandl, 1906; Kofoid and Campbell, 1929
Cymatoc)"lis l'allhojJelli, forma cOllica: Laackmann, 1910; Alder, Cyltarocylis plagiostoma var. c; C. plagios toma var. a; C. plagio-
1995 stoma var. b: Brandl, 1906
Cymatocylis flal'a, forma cOllica; C. l'allhojJeni, forma affillis; C. Cyttarocylis ellceCl)phalus: Kofoid, 1912; Kofoid and Campbell,
l'allhojJelli, forma cylilldrica; C. l'allhojJeni, forma calycina: 1929,1939; Balech, 1959; SoulO, 1979
Laackmann, 1910 C,l'ttarocylis 101lga: Kofoid and Campbell, 1929
Cymatocylis cOllica; C. affinoides; C. call'cina; C. el'erta: Kofoid Cyttarocylis brandti: Kofoid and Campbell, 1929, 1939
and Campbell, 1929 Cyttarocylis mllCrDnata: Kofoid and Campbell, 1939
Cymatocy/is valllroffelli, forma cylilldrica Laackmann Dadayiella gallymedes (Entz, Sr.) Kofoid and Campbell
C)'matoc)'lis l'allhojJelli, forma cylindrica: Laackmann, 1910; Tintinllus Ganymedes: Enlz, Sr., 1884; Brandl, 1906-1907
A1der,1995 Amphorella ganymedes: Daday, 1887b; Jorgensen, 1924
Cymatocylis I'anhoffelli, fOfila affillis; C. flava, forma cylindrica: TilltillllllS blllbosus; T. blllbosus var. a: Brandl, 1906-1907
Laackmann, 1910 Amphorella gall)'medes var. telllticallda, forma aCllta; A, ganyme-
Cymatocylis diminuta; C. cylindrella; C. contracta; C. cal)'x: Kofoid des var. tellllicallda, forma obtusa; A. gallymedes, forma acuta;
and Campbell, 1929 A. gallymedes var. blllbosa: Jorgensen, 1924
Cymatocylis valllrojJelli, forma coxliella nov. form. Dadayiella gall)'medes: Kofoid and Campbell, 1929, 1939; Balech,
Coxliellafrigida: Laackmann, 1910; Ba1ech, 1947, 1958b 1959; SoulO, 1970a
Cymatocylis l'allllOjJelli, forma coxliella: Alder, 1995 Dadayiella acuta; D. jorgenselli: Kofoid and Campbell, 1929
Cymatoc)'lis valllroffelli, abnormal specimens? Dadayiella blllbosa: Kofoid and Campbell, 1929, 1939
Cymatoc)"lis l'anhOjJelli: Laackmann, 1910 Dadayiella acutiformis: Kofoid and Campbell, 1939
Cymatoc)'lis l'anho.ffelli, forma robusta: Laackmann, 1910 Datllrella emargillata (Brandt) Kofoid and Campbell
Cymatocylis robllsta: Kofoid and Campbell, 1929 Tintinllus emargillatus: Brandl, 1906-1907
*Cyma/ocylis digitlllus: Balech, 1947 Tintinlllls IlIslIs-lIlldae var. macilentlls: Jorgensen, 1924
C)'matocylis l'Clllhoffeni, abnormal specimens?: Alder, 1995 Daturella emargillata: Kofoid and Campbell, 1929
Cymatocylis calycijorlllis (Laackmann) Laackmann Datllrella datllra (Brandt) Kofoid and Campbell
C)'ttarocylis cal)"ciformis: Laackmann, 1907 Tintinlllls datllra: Brandl, 1906-1907; Laackmann, 1910
Cymatocylis ca~l'cirormis: Laackmann, 1910; Kofoid and Campbell, TilltillllllS tubiflora: Brandl, 1907
1929; Ba1ech, 1962a Datllrella datura; D. gaussi: Kofoid and Campbell, 1929
Tintinnoinea 379
Dietyoeysta mitra Haeekel ? Epiplocylis labiosa: Kofoid and Campbell, 1929; Baleeh, 1962b
Dietyoeysta elegans: Ehrenberg, 1854a, 1854b Epiploeylis IlIIdella (Ostenfeld and Sehmidt) Jorgensen
Dietyoe)'sta mitra: Haeekel, 1873; Brandt, 1906-1907; Jorgensen, Cyttaroeylis undella: Ostenfeld and Sehmidt, 1901
1924; Kofoid and Campbell, 1929; Baleeh, 1959 Ptychocylis Imdella var. a; P. undella var. b; P. Imdella varo e; P.
Dietyoeys/a mi/ra var. a dilG/ata; D. mi/ra var. b; D. mi/ra var. e: undella var. e; P. undella var. f; P. lmdella var. h; P. undella
Brandt, 1906-1907 varo i; P. undella var. k; ? P. undella var. o: Brandt, 1906-1907
Dietyoeysta mitra var. minor; D. mi/ra, forma ob/usa; D. mitra, Epiplocylis ,mdella; E. undella var. blanda: Jorgensen. 1924
forma dilatata: Jorgensen, 1924 Epiplocylis atlantica; E, blanda; E. constricta; E. exquisita: E.
Dietyocysta minor; D. obtusa: Kofoid and Campbell, 1929 impensa; E. dejlexa; E. lineata; E. pacifica; ? Epiplocylis obtu-
Dictyocysta dilatata: Kofoid and Campbell, 1929, 1939 sa: Kofoid and Campbell, 1929
Dietyoeysta elegans Ehrenberg Epiploeyloides? retielllata (Ostenfeld and Sehmjdt) Hada
Dietyoeysta elegal/s Ehrenberg varo speciosa Jorgensen Cyttarocylis reticulata: Ostenfeld and Semidt, 1901
Dictyocysta speciosa: Kofoid and Campbell, 1929 Ptychocylis reticulata: Brandt, 1906-1907
Dictyocysta elegans; D. elegans var. a; D. elegam var. e; D, ele- Epiplocylis reticulata: Jorgensen, 1924
gans var. e mülleri; D. elegans varo d: Brandt, 1906-1907 Epiplocylis brandti: Kofoid and Campbell, 1929
Dictyocysta ampla: Kofoid and Campbell, 1929, 1939 Epiplocyloides re/iculata: Hada, 1938; Baleeh, 1962b; SOlitO,
Dictyocysta inaequalis; D. magna: Kofoid and Campbell, 1929 1970b, 1979
Dictyocysta elegans var. speciosa: Jorgensen, 1924; Ba1eeh, 1959; ? Epiploc)'lis curta, ? E. healdi: Kofoid and Campbell, 1929
1962b; SOlitO, 1970b, 1979 * Epiploeyloides? alltarctiea Baleeh
Dietyoeysta elegal/s varo lepidu (Ehrenberg) Baleeh Epiploc)'lis mira: Baleeh, 1958a
Dictyocys/u lepida: Ehrenberg, 1854b; Jorgensen, 1924; Kofoid Epiplocyloides antarctica: Baleeh, 1962a
and Campbell, 1929 Eutil/t;'lIIus jrakl/oi (Daday) Kofoid and Campbell
Dictyocysta templum: Haeekel, 1873; Brandt, 1906-1907 Tintinnus Fralmóii: Daday, 1887b
Dictyocysta tiara: Haeekel, 1873; Kofoid and Campbell, 1929 Tintinnusfralmóii: Kofoid and Campbell, 1929
Dictyoc)'sta templum var. a; D. templum varo b; D. templum var. f; Tintinnusfraknoi var. a: Brandt, 1906-1907
? Dictyocysta templum var. c grandis: Brandt. 1906-1907 Tin/innus elongatus: Kofoid and Campbell, 1929
Dictyocysta templum var. indica; D. templum var. h: Laaekmann, Eutintinnusfraknóii; Eutintilllllls elongatus: Kofoid and Campbell,
1910 1939
Dictyocysta lata; D. nidulus; ? Dictl'ocysta grandis: Kofoid and Eutil/tillllllS IlIslIs-lIl/due (Entz) Kofoid and Campbell
Campbell, 1929 Tintinnus lusus undae: Entz, Sr., 1885; Daday, 1887b; Brandt,
Dictyocysta mexicana; D. occidentalis; D. reticulata: Kofoid and 1906-1907; Kofoid and Campbell, 1929
Campbell, 1929, 1939 Tintillllus lusus-undae "main speeies": Jorgensen, 1924
Dictyocysta ef. lepida: Baleeh, 1944 ? Eutintinnus lusus-undae: Kofoid and Campbell, 1939; SOlitO,
Dictyocysta elegans varo lepida: Ba1eeh. 1959; SOlitO. 1970b, 1979 1979
Dietyoeysta duplex Brandt Elltintinnus mellills (Kofoid and Campbell) Kofoid and
Dictyocysta templum var. e duplex; D. templum var. d: Brandt. Campbell
1906-1907 Tintinnus medius: Kofoid and Campbell, 1929; Laaekmann, 1910
Dictyocysta duplex: Kofoid and Campbell, 1929 Eutintinnus medius: Kofoid and Campbell, 1939; SOlitO, 1979
* Dietyoeysta calijomiellsis Kofoid and Campbell * Elltillt;'III11S rllgoslIs Kofoid and Campbell
Dictyocysta californiensis Kofoid and Campbell, 1929; Baleeh, Eutintinnus I'IIgosus: Kofoid and CampbelI. 1939: Baleeh. 1944,
1971a; SOlitO, 1979 1945, 1948; SOlitO, 1979
? Dictyoc)'sta pacifica; ? D, ~pinosa; ? D. extensa: Kofoid and Elltintinl/lIs attellllatlls (Kofoid and Campbell) Kofoid and
Campbell, 1929 Campbell
Epieal/eella I/en'osa (Cleve) Kofoid and Campbell Tintinnus lusus-Imdae var. c: Brandt. 1906-1907
Cyttarocylis nerl'osa: Cleve, 1900b Tintinlllls attenuatus; T. colligatus: Kofoid and Campbell, 1929
Ptychoc)'lis nervosa: Brandt, 1906-1907 * Elltintilllllls apertlls (Kofoid and Campbell) Kofoid and
Epiplocylis nerl'osa: Jorgensen, 1924 Campbell
Epiplocylis (Epicancella) nerl'osa: Kofoid and Campbell, 1929 Tintilllllls apertlls: Kofoid and Campbell, 1929
Epicancella nervosa: Kofoid and Campbell, 1939; Baleeh, 1962b Eutintinnus apertus: Kofoid and Campbell, 1939; SOlitO, 1979
Epiploeylis aeumil/ata (Daday) Jorgensen ? Tintinnus lusus-undae var. a tubulosa: Brandt, 1906-1907
C)'ttarocylis acuminata: Daday, 1887b Fuvella turaikael/sis Hada, 1932
Pt)'chocylis acuminata; P. acuminata var. semireticulata; P. acu- Fal'ella taraikaensis: Hada, 1932b, 1937: SOlitO, 1970a, 1979;
minata var. b; P. acuminata var. c: Brandt, 1906-1907 Barría de Cao, 1986
Epiplocylis acuminata: Jorgensen, 1924; Kofoid and Campbell, Favella quequenensis: Baleeh, 1945, 1948
1929; Baleeh, 1959, 1962b; SOlitO, 1970a, 1979 *Fuvella urcuuta (Brandt) Jorgensen
Epiplocylis inconspicuata; Epiplocylis injlata: Kofoid and Cytturocylis arcuata: Brandt, 1906-1907
Campbell, 1929 Fal'ella arcllata: Jorgensen, 1924
* Epiploeylis ea(J'x (Brandt) Jorgensen Favella eltrenbergii (Claparede and Laehmann) Jorgensen
Ptychocylis ca~1'Xvar. a; P. calyx var. b: Brandt, 1906-1907 Favella eltrel/bergii, formajavella Laval-Peuto
Epiplocylis calyx: Jorgensen, 1924; Kofoid and Campbell. 1929 Tintilllllls ehrenbergi: Claparede and Laehmann. 1858-1859
380 Tintinnoinea
Cyttarocylis Márkllsol's=!,}'i; C. Claparedii: Daday, 1887b Metacylis sp. aff. mereselrkolVskyi Kofoid and Campbell
Cyttarocylis ehrellbergi; C. ehrellbergii varo a helgolalldica; Melacylis medilerrallea varo ponlica: J6rgensen, 1924
PI)'choc)'lis markllsol's=kyi: Brandt, 1906-1907 Melacylis mereschkowsÁyi: Kofoid and Campbell, 1929
C)'ttaroc)'lis ehrenbergi varo a c/aparedii: Brandt, 1906-1907; Faría Melacylis sp. atf. mereschkowsÁJ1i: Barría de Cao, 1986
and Da Cunha, 1917 Ormosella cOrllllcopia Kofoid and Campbell
Favella ehrenbergi: Jiirgensen, 1924; Kofoid and Campbell, 1929; TilltinllllS coniclIs: Brandt, 1906-1907
Baleeh, 1959; Souto, 1979 Ormosella corllllcopia: Kofoid and Campbell, 1929
Favella ehrenbergi varo helgolalldica; F. ehrenbergi var. c/aparedei: Ormosella acalltllUrlIS (Kofoid and Campbell) nov. combo
Jiirgensen, 1924 Amphorellopsis acalllhan/s: Kofoid and Campbell, 1929; Baleeh,
Favella marÁ'l/sol's=!»'i: Jiirgensen, 1924; Kofoid and Campbell, 1971a; Souto, 1979
1929 Ormosella Iraeckeli Kofoid and Campbell
Fal'ella helgolandica: Kofoid and Campbell, 1929 Ormosella haeckeli: Kofoid and Campbell, 1929; Baleeh, 1971a;
Fal'ella ehrellbergii, forma fa I'ella: Laval-Peuto, 1981 Souto, 1979
F. ehrellbergii, forma decipiells (Jiirgensen) Laval-Peuto ? Ormosella lrachelilllll; ? O. bresslalli: Kofoid and Campbell, 1929
Cyllaroc)'lis alllllllala: Entz, Jr., 1908 Parlllldella calldata (Ostenfeld) Jiirgensen
Coxliella decipiells: Jiirgensen, 1924; Kofoid and Campbell, 1929 TilllillnllS calldalLls: Ostenfe1d, 1899a, b
Favella ehrellbergii, forma decipiells: Laval-Peuto, 1981 Undella messinellsis varo a: Brandt, 1906-1907
F. ehrellbergii, forma coxliella Lava1-Peuto Ulldella (Panllldella) calldala; U (Parlllldella) aCllleala;
C)'ttarocylis (Coxliella) G/mlllata: Daday, 1887b; Brandt, 1906-1907 U. (Parllndella) aCllleala, forma IOllga: J6rgensen, 1924
Coxliella alllllllala: Brandt, 1906-1907; Laaekmann, 1913; Panllldella calldala: J6rgensen; Kofoid and Campbell, 1929, 1939:
Jiirgensen, 1924; Kofoid and Campbell, 1929 Baleeh, 1971 a; Souto, 1979
Cyttarocylis (Coxliella) helicoidea: Faría and Da Cunha, 1917 Parlllldella aCllleala; P. IOllga: Kofoid and Campbell, 1929
Fa\'ella ehrenbergii, forma coxliella: Laval-Peuto, 1981 Panmdella aCllleala IOllga: Ba1eeh, 1971 a
*Favella serrata (Miibius) Jiirgensen Parlllldella calldala 101lga;P. crassicalldala: Souto, 1979
Tilllinl1l/s serrallls: M6bius, 1887 ? Ulldella pellllcida: J6rgensen, 1899
Cyttaroc)'lis ser/'ala, C. serrala varo a: Brandt, 1906-1907 ? Parundella pellllcida: Kofoid and Campbell, 1929; Souto, 1979
Fal'ella serrala: J6rgensen, 1924; Kofoid and Campbell, 1929; Parlllrdella messillellsis (Brandt) Jiirgensen
Souto, 1970b, 1979 Ulldella messillellsis: Brandt, 1906-1907
Fal'ella attillgala: Kofoid and Campbell, 1929 Ulldella (Panllldella) messillellsis: J6rgensen, 1924
Helicostomella SIIblllata (Ehrenberg) Jiirgensen Panllldella messillellsis: Kofoid and Campbell, 1929, 1939; Souto,
TilllillllllS sllblllallls: Ehrenberg, 1883; Brandt, 1906-1907 1979
Cyttarocylis annllli(era: Ostenfeld and Sehmidt, 1901 Petalotrielra amp"lla (Fol) Kent
Tinlillnlls medilerranells varo IOllga: Brandt, 1906-1907 Tilllillnlls amp"lla: Fo1, 1881
Tinlilllllls palagolliclIs: Brandt, 1906-1907; Faría and Da Cunha, 1917 Pelalolricha amplllla: Kent, 1882; J6rgensen, 1924; Kofoid and
Helicoslomella sllblllala: J6rgensen, 1924; Souto, 1970a, 1979; Campbell, 1929; Baleeh, 1959, 1962b; Souto, 1970a, 1979
Barría de Cao, 1986 Pelalolricha amplllla varo b; P. amp"lla varo c; P. ampLllla varo d:
Helicoslomella kiliellsis; H. ji/si(ormis; H. edelllala; H. IOllga: Brandt, 1906-1907
Kofoid and Campbell, 1929 Pelalolricha amp"lla var. major: J6rgensen, 1924
Melacylis alllllll[(era: Kofoid and Campbell, 1929; Baleeh, 1951; Petalolricha serrala: Kofoid and Campbell, 1929
Souto, 1979 Pelalolricha majar; P. pacifica: Kofoid and Campbell, 1929, 1939
Helicoslomella Lemairei: Baleeh, 1944 PoroeclIs apiclllatlls (Cleve) Cleve
Laackmalll/iella IIaviclllaefera (Laackmann) nov. combo Porella apiclllala: Cleve, 1900b
Laackmalllliella IIaviclllaefera, forma IIaviclllaefera Laaekmann, PoroeclIs apiclllallls: C1eve, 1902b; J6rgensen, 1924; Kofoid and
1910 stat. nov Campbell, 1929.
Leprolinlillnlls naviclllaeferlls; L. prolollgallls; L. prolollgallls, Cyttarocylis apiclllala: Brandt, 1906-1907
forma velllricosa: Laaekmann, 1910 Prolorlrabdollella? simplex (Cleve) Jiirgensen
Laackmalllliella lIal'iClllae(era: Kofoid and Campbell, 1929 C)'ttarocylis silliplex: Cleve, 1900b
Laackmanlliella nal'iclllaefera, forma II01'iclllaefera: A1der, 1995 Rhabdollella amor var. simplex: Brandt, 1907; Laaekmann, 1910
Laackmalllliella IIaviclllaefera, forma prolollgata Laackmann, Protorhabdollella simple.x: J6rgensen, 1924; Kofoid and Campbell,
1910 stat. nov. 1929; Baleeh, 1962b; Souto, 1979
LeprolilllillllllS prolollgallls; L. prolollgatlls, forma I'elllricosa; Protorlrabdollella? curta (C1eve, 1901) J6rgensen, 1924
Leprolinlinlllls nal'iclllaeferlls: Laaekmann, 1910 Cyttarocylis slriala, forma) Cllrla: Cleve 1901
LaacÁ7nallniella prolollgala: Kofoid and Campbell, 1929 Cyttaroc)'lis amor var. cllrla: Brandt, 1907-1907
LaacÁ711alllliella lIal'iclllae(era, forma prolongala: Alder, 1995 Prolorhabdonella cllrla: J6rgensen, 1924; Kofoid and Campbel1,
Laackmalllliella IIaviclllaefera, forma coxliella nov. formo 1929,1939; Baleeh, 1944; Souto, 1979
Coxliella millor: Laaekmann, 1910; Kofoid and Campbell, 1929 * Rlrabdollella apoplrysata (Cleve, 1900) J6rgensen, 1924
LaacÁ71lanniella lIal'iclllae(era, fomla coxliella: Alder, 1995 Cyttarocylis Hebe varo apoph)'sala: Cleve, 1900b
Metacylisjorgellselli (Cleve) Kofoid and Campbell Cyttarocylis Iritoll: Zaeharias, 1906
Codonella J6rgellsellii: Cleve, 1902a Pt)'chocylis (Rhabdollella) apoph)'sala; P. (Rhabdollella) apophy-
TilllillllllS IIrceolallls: Brandt, 1906-1907 sala var. b: Brandt, 1906-1907
Melacl"!isj6rgellsellii: Kofoid and Campbell, 1929; Ba1eeh, 1959 Rhabdonella apopln'sata: J6rgensen, 1924
Tintinnoinea 381
Rhabdonellopsis apophysala: Kofoid and Campbell, 1929; Souto, ? Salpillgella allenllala: Jorgensen; Kofoid and Campbell, 1929,
1970a, 1979 1939
Rhabdonellopsis inlermedia; R. longicalllis: R. Irilon: Kofoid and ? Salpingella gracilis: Kofoid and Campbell, 1929, 1939
Campbell, 1929, 1939 Salpil/gella aCllmil/ata varo IIl/data (Brandt) Jorgensen
Rlzabdol/ella spiralis (Fol) Brandt Tintillnlls lIndallls; T lIndatlls varo lIllglliclllala: Brandt, 1906-1907
TilllillllllS ;piralis: Fol, 1881 TilllillllllS aCllmillallls varo IIl1dallls: Jorgensen, 1899
TinlinllllS slriallls: Biedennann, 1893 Salpillgacalllha IInglliclllala: Kofoid and Campbell, 1929
? Ulldella spiralis: Daday, 1887b Salpillgella aCLIIllillala, fonna lIlldala: Jorgensen, 1924
Cyllarocylis Hebe: Cleve, 1900b Salpillgacallllza IIlldala: Kofoid and Campbell, 1929
Plychocylis (Rlzabdonella) spiralis; P. (Rhabdollella) spiralis vaL d Salpil/gella aClImil/atoides (Laackmann)
slriala; P. (Rhabdonella) spiralis varo e Izenseni; Salpil/gella aClIminatoides varo II/,data Laackmann stat. nov.
P. (Rhabdonella) spiralis varo b chal'esi; ? P~\'chocylis (Rhabdollella) Tinlilllllls lIlldallls: Laaekmann, 1910
spiralis varo a hebe: Brandt, 1906-1907 Salpillgacanlha simplex: Kofoid and Campbell, 1929
Rhabdollella spiralis: Jorgensen, 1924; Kofoid and Campbell, Salpil/gella aClImil/atoides varo secaia Laackmann
1929, 1939 TinlillllUs aCllmillaloides; T aCllmillaloides varo seca la: Laaekmann,
Rhabdollella henselli: Kofoid and Campbell, 1929, Baleeh, 1962b 1910
Rhabdollella chavesi; R. slriala: Kofoid and Campbell, 1929 Salpingella aCllminaloides: Kofoid and Campbell, 1929
Rhabdollella hebe; R. cOllica: Kofoid and Campbell, 1929, 1939 Salpingella laackmanni: Kofoid and Campbell, 1929; Baleeh and
* Rlzabdol/ella clziliel/sis Kofoid and Campbell El-Sayed, 1965; Souto, 1979
Rhabdollella chiliensis: Kofoid and Campbell, 1929, 1939; Baleeh, Salpil/gella costata (Laackmann) Kofoid and Campbell
1971 a; Souto, 1979 Tinlinnlls coslallls: Laaekmann, 1910
* Rlzabdol/ella cOrJIllcopia Kofoid and Campbell Salpillgella coslala: Kofoid and Campbell, 1929
Rhabdollella cornllcopia: Kofoid and Campbell, 1929, 1939; Salpillgella reglllata (Brandt) Kofoid and Campbell
Baleeh, 1962b; Souto, 1970a, 1979 TinlillllUs reglllallls: Brandt, 1906-1907
* Rlzabdol/ella amor (Cleve) Brandt Salpingella regulata: Kofoid and Campbell, 1929.
Cyllarocylis Amor: Cleve, 1900b Salpil/gella Sllbcol/ica Kofoid and Campbell
Plychocylis (Rhabdollella) amor: Brandt, 1906-1907 Salpingella sllbconica: Kofoid and Campbell, 1929,1939; Baleeh,
¡
Rhabdonella amor: Laaekmann, 19 O; Kofoid and Campbell, 1929; 1971a; Souto, 1979; Barría de Cao, 1986
Baleeh, 1962b; Souto, 979 ¡ Steel/stmpiella steellstmpii (Claparede and Lachmann) Kofoid
* Rlzabdollella amor varo clIspidata Brandt and Campbell
Plychocylis (Rhabdollella) amor varo a cllspidala: Brandt, 1906- TinlillllllS steellslrllpii: Claparede and Laehmann, 1858-1859;
1907 Brandt, 1906-1907
Rhabdonella brandli: Kofoid and Campbell, 1929 Sleenslrupiella sleenslrupii: Kofoid and Campbell, 1929; Baleeh,
* Rlzabdol/ella amor varo valdestriata Brandt 1962b
Plychocylis (Rhabdonella) amor varo b I'aldestriala: Brandt. 1906- Steenstmpiella steenstrllpii varo robllsta (Kofoid and Campbell)
1907 Balech
Rhabdollella valdeslriala: Kofoid and Campbell, 1929; Baleeh, Amphorella Sleenslrupii: Daday, 1887b
1962b TilllillllllS sleenslrllpii: Brandt, 1906-1907
Rhabdollella elegalls: Jorgensen, 1924; Kofoid and Campbell, Sleenslrllpiella robllsla: Kofoid and Campbell, 1929; Souto, 1970a,
1929; Baleeh, 1962b 1970b, 1979
Rhabdollella iliflala; R. qllalllllla: Kofoid and Campbell, 1929 Sleenslrllpiella sleenslrupii varo robllsla: Baleeh, 1962b
* Rlzabdol/ella amor varo il/dica Laackmann *Steenstrllpiella pozzi Balech
Rhabdollella amor varo illdica: Laaekmann, 1910 Sleellstrtlpiella po::::i: Baleeh, 1942, 197 1a; Souto, 1972, 1979
Rhabdollella illdica: Kofoid and Campbell, 1929, 1939; Souto, 1979 Stelidiella stelidillm (Biedermann) Kofoid and Campbell
?Rhabdollella exilis: Kofoid and Campbell, 1939 TilllillllUs slelidillm: Biedennann, 1893; Brandt, 1906-1907
Salpil/gella aClImil/ata (Claparede and Laachmann) Jorgensen Slelidiella slelidillm: Kofoid and Campbell, 1929
Salpil/gella aClImil/ata varo secata Brandt Stenosemella avellal/a (Meunier) Kofoid and Campbell
TilllillllllS aCllminallls, T aCllmillalllS varo secala: Brandt. 1906- Tilllinnopsis m'ellana: Meunier, 19 9 ¡
1907 Slellosemella al'ellana: Kofoid and Campbell, 1929; Baleeh, 1948;
Salpingella aCllmillala: Jorgensen, 1924; Kofoid and Campbell, Hada, 1961; Souto, 1970b, 1972, 1979
1929, 1939; Souto, 1979 Stylicallda platensis (Da Cunha and Da Fonseca) Baleeh
Salpillgella secala: Kofoid and Campbell, 1929 TinlillllOpsis platensis: Da Cunha and Da Fonseea, 1917; Kofoid
Salpil/gella aClImil/ata varo glockel/togeri (Brandt) Jorgensen and Campbell, 1929; Baleeh, 1945; Gold and Morales, 1976
TilllillllllS aCllmillalllS: Claparede and Laehmann, Daday, 1887b Stylicallda plalensis: Ba1eeh, 1951; Souto, 1970a, 1979; Barría de
TilllillllllS aCllminallls varo c glockenlogeri: Brandt, 1906-1907 Cao, 1981
Salpillgella aClImillala glockelllogeri varo allenllala; S. aClImillala Tintinl/idillm sp. aff.T. semiciliatllm (Sterki) Kent
glockelllogeri: Jorgensen 1924 TinlillllllS semiciliallls: Sterki, 1879
Salpingella glockelllogeri: Kofoid and Campbell, 1929. 1939: Tilltinnidillm semicilialllm: Kent, 1882; Kofoid and Campbell, 1929
Baleeh, 1959; Souto. 1979 Tinlillnidillm sp. aff. semiciliatllm: Barría de Cao, 1981
Salpingella sinislra: Kofoid and Campbell, 1939 Til/til/I/idill/II baleclIi Barria de Cao
382 Tintinnoinea
Tinlinnidillm balechi: Barría de Cao, 1981 Tintinnopsis karajacensis var. a: Brandt, 1906-1907
"TintillllOpsis" radix (Imhof) Brandt Tinlinnopsis parvllla: J6rgensen, 1912; Kofoid and Campbell, 1929;
Codonel/a radix: lmhof, 1886 Baleeh, 1945, 1948; SOlitO, 1970, 1979; Barría de Cao, 1981
Tilllinnopsis Davidoffii var. cylindrica: Daday, 1887b Tintinnopsis gracilis: Kofoid and Campbell, 1929; Baleeh, 1948;
Tinlillnopsis radix: Brandt, 1906-1907; Kofoid and Campbell, SOlitO, 1970a, 1979; Barría de Cao, 1981
1929; Baleeh, 1945; Souto, 1970a "TilltillllOpsis'" lobial/coi Daday Ibrasiliensis Kofoíd and
Tinlinllopsis davidoffi var. cylindrica; Tilllinnopsis fracla: Brandt, Campbell
1906-1907 Tintinnopsis Lobiancoi: Daday, 1887b
Tinlinllopsis cJlilldrica: Kofoid and Campbell, 1929 Ti/!Iinnopsis sp.: Brandt, 1906-1907
"Til/tinnopsis" aperta Brandt Tintinnopsis lobiancoi: Kofoid and Campbell, 1929; Baleeh, 1945;
Tinlinnopsis aperla: Brandt, 1906-1907; Kofoid and Campbell, Souto, 1979; Barría de Cao, 1981
1929; Baleeh, 1948; SOlitO, 1970a Tilltillnopsis brasiliellsis: Kofoid and Campbell, 1929; Souto, 1979;
"TintÍl/llopsis" tocal/til/el/sis Kofoid and Campbell Barría de Cao, 1981
Tintinllopsis aperla var. a: Brandt, 1906-1907 Tilltillnopsis lala: Baleeh, 1945.
Tinlinnopsis locanlinensis: Kofoid and Campbell, 1929; Baleeh, "TilltillllOpsis" parva Merkle
1948; Souto, 1970a; Barría de Cao, 1981 Tilllinnopsis parva: Merkle, 1909; Kofoid and Campbell, 1929;
"Tintinnopsis" I/ordqvisti Brandt Ba1eeh, 1948; SOlitO, 1979; Barría de Cao, 1981
Tilllinnopsis lIordqvisti: Brandt, 1906-1907 Tilltillnopsis glalls: Meunier 1919; Baleeh, 1945; Souto, 1979;
LeprolilllillllllS nordqvisli: Kofoid and Campbell, 1929; Barría de Barría de Cao, 1981.
Cao, 1986 Tinlinnopsis tllrbinala: Ba1eeh, 1948; SOlitO, 1979; Barría de Cao,
"TintillllOpsis" kofoidi Hada 1981
Tintinllopsis kofoidi: Hada, 1932a, 1932e; Baleeh, 1948; SOlitO, 1979 "Til/tillllOpsis" compressa (Daday)
"Til/tÍl/llopsis" all/pltistoll/a Baleeh Tinlinnopsis beroidea var. compressa: Daday, 1887b
Tinlinllopsis amphistoma: Baleeh, 1951; Souto, 1979 Tilllillllopsis compressa: Laaekmann, 1913; J6rgensen, 1924; Kofoid
"Til/til/nopsis" all/pllOra Kofoid and CampbelVl/llcllla (Fol) and Campbell, 1929; ?Baleeh, 1959; Barría de Cao, 1981
Codollel/a III1Cllla:Fol, 1884 "Tintil/nopsis" beroidea Stein
Tinlinnopsis IIl1cltla: Brandt 1906-1907; Kofoid and Campbell, 1929 Tinlinnopsis beroidea: Stein, 1867; Entz Jr., 1884; Meunier, 1910;
Tilllinnopsis lobiancoi var. filsiformis: Brandt, 1906-1907 J6rgensen, 1924; Kofoid and Campbell, 1929; Durán, 1953;
Tinlillnopsis amphora: Kofoid and Campbell, 1929; Barría de Cao, Baleeh, 1959; Gold and Morales, 1975; Souto, 1979; Barría de
1981 Cao, 1981
"Til/tillllOpsis" baltica Brandt ? Tinlillllopsis beroidea var. b: Brandt 1906-1907
Tinlinnopsis baltica: Brandt, 1896; Kofoid and Campbell, 1929; "TintÍl/llopsis" p"silla Hadal7aevis Waíles
Baleeh, 1945; Gold and Morales, 1975; Souto, 1972, 1979; Tintinnopsis davidoffi var. laevis: Wailes, 1925
Barría de Cao, 1981 Tilltinnopsis pIIsil/a Hada, 1937; Souto, 1979; Barría de Cao, 1981
Tinlinnopsis ballica varo a: Brandt, 1906-1907 Tinlinnopsis levigata Kofoid and Campbell, 1929; Souto, 1979;
Tintinnopsis denticlllala: Kofoid and Campbell, 1929 Barría de Cao, 1981
"Til/tinnopsis" IIrllgllayel/sis Baleeh "TintÍl/llopsis" prowazeki Faría and Da Cunha
Tintinnopsis IIl'l1gllayensis: Baleeh, 1948; SOlitO, 1979 Tintillnopsis prowa=eki: Faría and Da Cunha, 1917; SOlitO, 1979
"Tintinl/opsis" jill/briata Meunier "Tintinl/opsis" crutera (Leidy) Hada
Tinlinnopsis fimbriata: Meunier, 1919; Kofoid and Campbell, 1929 Difllllgia cratera: Leidy, 1879
Tintinnopsis sp.: Brandt, 1906-1907 Codonel/a cralera: Kofoid and Campbell, 1929; Souto, 1979
Tintinnopsis merlllieri: Kofoid and Campbell, 1929 Tinlinnopsis cralera: Hada, 1939
Codonaria jilllbriata: Ba1eeh, 1948 VI/del/a Ityalil/a Daday
"Tintill/lOpsis" campanilla Ehrenbergl blltsclr/ii Daday Vndel/a hyalina: Daday, 1887a; Brandt, 1906-1907; Kofoid and
Tinlinnlls call1pamtla: Ehrenberg, 1840 Campbell, 1929; Baleeh, 1959, 1975
Tinlinnopsis Blltschlii: Daday, 1887b Vlldel/a hyalilla var. a; U. hyalina var. b; U. hyalilla var. c: Brandt,
"Tintinl/opsis" campallltla/blltscltlii varo blltscltlii Daday 1906-1907
TinlillllOpsis biilschlii: Kofoid and Campbell, 1929; Baleeh, 1948; Undel/a iJyalilla, fonna allenllala: J6rgensen, 1924
DlIrán,1953 Undel/a allenllala; U. blll/a; U. declil'is; U. di/ala la; U. parva; U.
TinlillllOpsis bllelsciJlii, varo blletsc1rlii: Souto, 1979 pemana; U. malllmi/ala; U. media: Kofoid and Campbell, 1929
Tinlinnopsis bllelsclriii, var. morlensenii: Souto, 1979, Barría de Vndella Itemispltaerica Laaekmann
Cao, 1981 Undel/a hemisphaerica: Laaekmann, 1910; Kofoid and Campbell,
Tintinnopsis cyatlms: Daday, 1887b; Kofoid and Campbell, 1929 1929; Ba1eeh, 1975
T cyaliJlIs var. annlllala: Daday, 1887b *Ulldella grul/dis (Laaekmann) Baleeh
"TilltillllOpsis" campal/lIla/blltsc/rlii varo II/ortel/sel/ii (Sehmidt) Undel/a c1aparedei var. gralldis: Laaekmann, 1910
Baleeh Proplectel/a gralldis: Kofoid and Campbell, 1929
Tinlinnopsis Mortensenii: Sehmidt, 1901; Brandt, 1906-1907; VI/del/a c1apuredei group
Kofoid and Campbell, 1929; SOlitO, 1979 Tintinnlls sp.: Claparede and Laehmann, 1858-1859 J
Tintinnopsis biitschlii varo lIlortensenii: Baleeh, 1948 Tilllinnlls c1aparMii: Entz, Sr., 1885
"Tintinl/opsis" parvllla Jiirgensen Undel/a ClaparMei: Daday, 1887b; Baleeh, 1975; Souto, 1979
Tintinnoinea 383
Undella subacu/a: Cleve, 1901; Balech, 1975; SOlitO, 1979. Xys/onellopsis cyma/ica var. spica/a: Jorgensen, 1924
Undella c1aparedei varo c; U c1aparedei varo d; U c1aparedei varo Xys/onellopsis dicyma/ica; X inaeql/a/is; X mascarensis; X
e globosa; U c1aparedei varof; U c1aparedei varo g: Brandt, cyclas; X spica/a: Kofoid and Campbell, 1929
1906-1907 Xystol/ellopsis paradoxa (Cleve) Jiirgensen
Undella subcauda/a; U sÍlbacu/a varo subcauda/a: Jorgensen, 1924 Undella paradoxa: C1eve, 1900b
Undella subcauda/a sl/bcauda/a: Jorgensen, 1924 Balech. 1975; Cyl/arocy/is? (Xys/onella) paradoxa: Brandt, 1906-1907
SOlitO, 1979 Xys/onellopsis paradoxa: Jorgensen, 1924; Kofoid and Campbell,
Proplec/ella c1aparedei: Kofoid and Campbell, 1929; Ba1ech, 1959 1929; Balech, 1959; SOlitO, 1979
Proplec/ella pelpusilla; P. praelonga; P. /enuis; P. biangula/a; P. Xys/onellopsis conicacauda: Kofoid and Campbell, 1939
pan'a; P. Jas/iga/a; P. globosa; P. ova/a; P. angl/s/ior; P. sl/b- Xystol/ellopsis Ireros (Cleve) Kofoid and Campbell
cauda/a; P. subacu/a: Kofoid and Campbell, 1929 Undella heros: Cleve, 1900b
Proplec/ella globosa: Kofoid and Campbell, 1939 Undella heros varo gal/ssi: Laackmann, 1910
Undella'globosa; ? U perpl/silla: Ba1ech, 1975; SOlitO, 1979 Xys/onellopsis heros: Kofoid and Campbell, 1929; Ba1ech, 1962b;
UI/dellopsis marsupialis (Brandt) Kofoid and Campbell Souto, 1970a, 1979
Undella marsl/pia/is: Brandt, 1906-1907 Xys/onellopsis gaussi: Kofoid and Campbell, 1929
Undella marsupia/is, forma subangula/a; U marsupia/is, forma Xystol/ellopsis omata (Brandt) Kofoid and Campbell
sllbcy/indrica: Jorgensen, 1924 Cyllarocy/is (Xys/onella) orna/a: Brandt, 1906-1907
Undellopsis (Undellopsis) marsl/pia/is; U (Undellopsis) /inea/a; Cyl/arocy/is orna/a: Brandt, 1906-1907
U (UI/dellopsis) pacifica: Kofoid and Campbell, 1929 Xys/onellopsis O/'na/a: Kofoid and Campbell, 1929
Undellopsis sl/bangula/a: Kofoid and Campbell, 1929; Balech, 1975 Xystol/ellopsis pulcllra (Kofoid) Kofoid and Campbell
Undellopsis cubitum: Kofoid and Campbell, 1929, 1939 Cyl/arocy/is pule/Ira: Kofoid, 1905
Undellopsis marsupialis: Balech, 1975 Xys/onellopsis pulchra: Kofoid and Campbell, 1929
Xystonella treJorti (Daday) Laaekmann
Xystonella treJorti, forma typica nov. formo
Cyl/arocy/is rreJor/i: Daday, 1887b
Cyl/arocy/is? (Xys/onella) /reJor/i: Brandt, 1906-1907
Suggested readings
Xys/onella /rejor/i: Laackmann, 1910; Jorgensen, 1924; Kofoid and
Campbell, 1929, 1939; Balech, 1959; SOlitO, 1970a. 1979
Xystol/ella /reJorti, forma coxliella nov. formo Alder V.A. 1995. Ecologia y sistemática de Tintinnina
Cyllarocy/is Jasciata: Kofoid, 1905 (Protozoa, Ciliata) y microzoopláncteres asociados
Cyllarocy/is (Cox/iella)Jascia/a varo procera: Brandt, 1906-1907 de aguas antárticas. Ph. D. Diss., Univ. Buenos
Cox/iellaJascia/a: Kofoid and Campbell, 1929
Aires, pp. 1-317. Systematics, hori::ontal, vertical
Xystol/ella IOl/gicauda (Brandt) Laaekmann
Cyllarocy/is (Xys/onella) acus varo b longicauda: Brandt, 1906
and seasonal qllantitative and qualitative distribu-
Xys/onella longical/da: Laackmann, 1910; Jorgensen, 1924; Kofoid tion, ec%gy, etc. o/ the tintinnids o/ the At/antic
and Campbell, 1929. 1939; Balech, 1959; SOlitO, 1970a; 1979 sector o/ the Southern Ocean based on e.x:tensive
Xystol/ellopsis Javata (Brandt) Jiirgensen multiannual collections.
Cyl/arocy/is? (Xys/onella) Java/a: Brandt, 1906-1907 Brandt K. 1906. Die Tintinnodeen del Plankton-
Xys/onellopsis Java/a: Jorgensen, 1924; Kofoid and Campbell, 1929
Xystonellopsis Irastata (Biedermann) Kofoid and Campbell
Expedition, Tafelerklarungen nebst kurzer
Tin/innus Iras/a/us: Biedemlann, 1893 Diagnose der neuen Arten. Plankton-Exp.
Cyl/arocy/is (Xys/onella) has/ala: Brandt, 1906 Humboldt Stiftung, Ergebn., 3(L.a.), pp. 1-33, 70
Cyl/arocy/is has/ala: Brandt, 1907 pis. One o/ the most detailed works on tintinnids,
Xystonella has/ala: Brandt, 1907
with excellent illustrations o/ all the species, /or-
Xys/onellopsis has/ala: Kofoid and Campbell, 1929
* Xystol/ellopsis? sp. Baleeh mae and variefies recogni::ed in the author 's 1907
? Fa"ella aciculifera: Jorgensen, 1924 monograph.
? Parlllldella spinosa: Kofoid and Campbell, 1929 Brandt K. 1907. Die Tintinnodeen der Plankton-
? Parundella aciculifera: Kofoid and Campbell, 1939 Expedition. Systematischer Teil. Plankton-Exp.
? Xys/onellopsis aciculifera: Balech, 1968
Humboldt Stiftung, Ergebn., 3(L.a.), pp. 1-499. A
Xys/onellopsis sp.: Balech, 1971a
Xystol/ellopsis cymatica (Brandt) Jiirgensen
cornersfone work on the systematics o/ worlwide
Cyllarocy/is (Xys/onella) cyma/ica; C. (Xys/onella) cyma/ica varo a; marine fin tin nids, with special emphasis on taxo-
C. (Xys/onella) cyma/ica varo b; C. (Xys/onella) cyma/ica varo 110mic categories sllch as Formenkreis, species,
c. ~pica/a; C. (Xys/onella) dicyma/ica; C. (Xystonella) dicyma- /ormae and varieties.
/ica varo a: Brandt, 1906
Brownlee D.C. 1977. The significance of cytological
Xys/onella cyma/ica; X dicyma/ica: Laackmann. 1910
Xl's/onellopsis cyma/ica: Jorgensen, 1924; Kofoid and Campbell,
characteristics as revealed by protargol silver stai-
1929: Balech, 1968; SOlitO, 1979 ning in evaluating the systematics of the suborder
384 Tintinnoinea
Tintinnina. Ms. Sci. Diss., Univ. Maryland, pp. 1- their phylogenetic relationships with other ciliates.
146. A detailed account of tintinnid cytology. Laval-Peuto M. 1994. Classe des Oligotrichea
Daday E. Von. 1887. Monographie der Familie der Bütschli, 1887. Ordre des Tintinnida Kofoid et
Tintinnodeen. Mitt. Zoo!. Sta1. Neapel, 7:473-591. Campbell, 1929. In "Traité de zoologie. ll.
The first mOllograph 011 tilltinnids, with descrip- Infusoires Ciliés" (P. de Puytorac, ed.), Masson,
tions of9 genera, 70 species and 8 varieties (22 of Paris, pp. 181-219. General review of the group,
them belonging to the genus Tintinnopsis). with emphasis on morphology, cytology, lije
Kofoid C.A., Campbell A.S. 1929. A conspectus of cycles and class[fication.
the marine and freshwater Ciliata belonging to the Pierce R.W., Turner J.T. 1992. Ecology ofplanktonic
suborder Tintinnoinea, with description of new ciliates in marine food webs. Rev. Aqua1. Sci.,
species principally from the Agassiz Expedition to 6: 139-181. An overview of the role of planktonic
the Eastern Tropical Pacific 1904-1905. Univ. ciliates (aloricate and tintinnids) in marine pela-
California Pub. Zoo!., 34:1-403. This treatise is gicfood webs, including comments on their abun-
the most widely used revision on tintinnids. It dance, distribution and methodological aspects.
includes descriptions, illustrations and synony- Pierce R.W., Turner J.T. 1993. Global biogeography
mies of al! the species erected until J 929, most of of marine tintinnids. Mar. Eco!. Prog. Ser., 94: 11-
which are renamed and reorganized in a new clas- 26. Worldwide biogeography of tintinnid genera
sification system. based on over J 700 records from J 400 locations.
Kofoid C.A., Campbell A.S. 1939. Reports on the Includes a brief discussion of the physical and bio-
scientific results of the expedition to the Eastern logical factors involved.
Tropical Pacifico The Ciliata: the Tintinnoinea. Reid P.C., John A.W.G. 1978. Tintinnid cysts. J. Mar.
Bul!. Mus. Compar. Zoo!. Harvard Coll., 34:1- Bio!. Ass. U.K., 58:551-557. Review on tintinnid
473. The most complete systematic revision based resting cysts and some forms attributed to
on tropical oceanic materials. Modifies some of Papulijeres Meunier.
the systematic criteria presented in the earlier Souto S. 1979. Tintínidos del Atlántico Sudoccidental
(1929) publication. Includes excel!ent original y Antártida Argentina (Protozoa, Ciliata). Estudio
illustrations for al! the taxa treated. taxonómico, distribucional y ecológico. Ph. D.
Laackmann H. 1910. Die Tintinnideen der deutschen Diss., Univ. Buenos Aires, p. 1-191.
Südpolar Expedition 1901-1902. Deutsch. Souto S. 1981. Tintinnina. In "Atlas del Zooplancton
Südpolar-Exp. 1901-1903, ll(Zoo!. I1I):340-496. del Atlántico Sudoccidental y métodos de trabajo
A complete and detailed systematic treatemet of con el zooplancton marino" (D. Boltovskoy, ed.),
tintinnd species, mainl)' centered on Antarctic Public. Esp. Ins1. Nac. Inv. Desarrollo Pesq., Mar
materials, but also covering oceallic waters oI the del Plata, pp. 353-381. The above 2 papers by
south Atlantic. Souto deal with the systematics and distribution of
Laval-Peuto M. 1981. Construction of the lorica in the tintinnids off the coasts South America III
ciliata tintinnina. in vivo study of Favel!a ehrenber- general, and off Argentina in particular.
gii: variability of the phenotypes during the cycle,
biology, statistics, biometry. Protistologica, 17:249-
272. An excel!ent account oI tintinnid reproduction
with importallt taxonomic implications.
Laval-Peuto M., Brownlee D.C. 1986. Identification
and systematics of the Tintinnina (Ciliophora):
Evaluation and suggestions for improvemen1. Ann.
Ins1. Océanogr., Paris, 62:69-84. Stresses the
importance of cytological data and some relevant
non-mOlphologic traits (biology, physiology, ecolo-
gy, reproductioll, etc.)for tintinnid systematics and
/
Hydromedusae
Jean Bouillon
Introduction With a few exceptions, sexes are separate. Sex cells

generally mature in the ectoderm (Fig. IA, C). The
Hydromedusae, the planktonic, adult stage of the gastrula larva, or planula, resulting from fertilised
Hydroidomedusae, belong to the phylum Cnidaria. eggs, has only two embryonic cell layers, the ecto-
Cnidarians are diploblastic, acoelomate metazoa, they derm and the endoderm. Hydroidomedusae have a
have a tissue level of construction, lacking real organs. simple mouth which opens directly into the gastrovas-
Their digestive system or coelenteron is sack-like or cular cavity without an actynopharynx, and their gas-
branched, with a single orifice serving as both mouth tric cavity lacks septa similar to those of the
and anus. They are usually radially (more rarely bilat- Scyphozoa and Anthozoa (Fig. IA, C, 21). Their
erally) symmetrical with the primary body axis oral- mesoglea is acellular, the cnidocysts being usually
aboral. Cnidaria are fundamentally tentacle-bearing restricted to the ectoderm.
animals, covered with stinging organs or cnidae (cni-
docysts, spirocysts, and ptychocysts), after which the The hydromedusae are essentially carnivorous, but
phylum is named (Fig. 2M, P, R, S). Two basic struc- some may feed on bacteria, protozoans, phyto-
tural types can be recognised among the Cnidaria: the plankton, algae, and even dissolved organic matter.
polyp, which is benthic, sessile, and generally modular Some species harbour symbiotic intracellular algae
(that is, consisting of a series of morphologically sim- from which they may derive some nutrients. They are
ilar structural units), and the medusa, planktonic, free- among the most important planktonic predators.
swimming and solitary. Cnidarian species may be rep-
resented by medusae or polyps, or have both stages in The ca. 700 described species of hydromedusae live
their development. They typically have a ciliated, in all seas, at all latitudes and all depths. Only a few
motile gastrula - the planula larva. have colonised brackish or fresh water.
Symmetry in the Hydroidomedusae is tetramerous,

polymerous, or rarely bilateral. They typically under- Methods
go a polymorphic succession, beginning with the
polyp, an asexual benthic, generally sessile stage, Collection of material
which is followed by the medusa (= hydromedusa), a
sexual planktonic free-swimming stage (Fig. ID, bot- Hydromedusae can be caught by plankton nets tofled
tom). In many forms the medusae are reduced to go- very slowly (1 to 1.5 knots) behind large or small pow-
nophores, which no longer leave the hydroid colonies, ered vessels, or even a rowboat, for about ten to twen-
and in a few others the medusae give rise directly to ty minutes, depending on the concentration of plank-
other medusae (Fig. ID, top). In Hydroidomedusae ton. The mesh of the net should be about 200-250 pm.
the asexual budding of a new medusa or gonophore Larger meshes permit many small specimens to
usually involves formation of a medusary nodule or escape, whereas smaller ones clog easily and damage
entocodon (except in the Narcomedusae) (Fig. IE). the specimens. In coastal waters the mouth-opening of
The hydroids can be solitary but usually form colonies the net should be from 30 cm to 1 m, depending on the
by budding. The colonies, with interconnected power of the vessel. Largest mouths are used in the
coelenteron, often produce individual polyps specia- open sea and in deep water where the fauna is sparser.
lised for different functions (defensive dactylozooids, In areas very rich in plankton, a hand net or even a
reproductive gonozooids, nutritional gatrozooids, bucket may be used. The richest catches are generally
etc.). The hydromedusae are solitary, free-swimming, obtained in the early morning or at dusk, and on rising
and are provided with a velum (except Obelia) tides. For qualitative surface sampling the plankton net
occluding the umbrella aperture (craspedote medusae) is towed 50 m or more behind the vessel to minimise
(Fig. 1A, C, 2A, D, H). turbulence. For sampling a few meters below the sur-

0 1999 Backhuys Publishers, Leiden, The Netherlands
Hydromedusae 387
opening of the
radial canal umbrellar
endodermal or
ring canal
Mm;-
!p
stage
Ca 1
1
QI---, 0 7-
planula\
_,
x
fjj
..-I-%,
fF$/ cyst ,
A /
Direct development
1 (no hydroid stage)
1
I
%- ' -
medusa
- \
m- 7-+.
cyst
stage planula
2 I 1
@ IC /
2 Y
XfL
Development with
hydroid stage
Hydromedusae
ectoblastic
cells
- outline of
future
radial canal
ectoderm
medusar nodule
=entocodon
=glokenkern
- endoderm
oral
tentacle
oral
cavity
gastric
cavity
Fig. 1. A: Diagrammatic optical section across a medusa of Cladonema radiatum (Anthomedusae). Left side a radial section, right side an
interradial one. B: Diagram defining the radii o f a hydromedusa with 4 radial canals. C: Diagrammatic section o f a hypothetical Leptomedusae; left
side a radial section, right side an interradial one. D: life cycle of a hydromedusae with direct development, without hydroid stage (top), and with a
hydroid stage (bottom). E: Section through a medusa bud of Limnocnida tanganyicae. F: Diagram of part of a colony ofLaodicea undulata showing
a hydranth with a typical conical hypostome (order Conica) (left); and a gonotheca with medusa buds (right). G: A diagrammatic view of a hydranth
ofLaomedeaj7exuosa showing the globose hypostome characteristic of the Order Proboscoida.
A: redrawn from Bouillon and Houvenhagel(l970); B: redrawn from Russell (1953); C: redrawn from Kramp (1919); D: redrawn from Boero et al.
(1992); E: redrawn from Bouillon(1957); F: redrawn from Russell (1936); G: redrawn from Khiin(1914).
t
face a buoy can be attached with a rope of known alcohol, going from formaldehyde to a very dilute
length to one side of the mouth-ring and a weight on alcohol solution (less than 10%) and then, step by step
the other side. Sampling between fixed depths requires (10% by 10%) over several days, to the final 70%
closing nets. Quantitative sampling is possible with solution. Polythene containers should be avoided also
plankton nets fitted a little behind the front with a ca- because chemical.precipitates may damage the speci-
librated flow-meter to measure the volume of water fil- mens. For histological studies the best fixative (after
tered (see UNESCO, 1968; Boltovskoy, ed., 1981; anaesthesia) is cold (543°C) acidic Bouin's fixative (3
Omori and Ikeda, 1984; for further details on general parts of saturated aqueous solution of picric acid and 1
zooplankton sampling techniques). part of formaldehyde, just before use 5% glacial ace-
tic acid should be added to this solution).Fixationtime
If it is impossible to sort the medusae immediately, from 2 to 6 hours. This procedure minimises tissue
the catch should be fixed as quickly as possible with maceration in organisms subsequently kept for long
5% formaldehyde in sea water (see Fixation and pre- periods of time in 5% formaldehyde.
servation below). If the material can be brought rapid-
ly to the laboratory, the plankton samples, shielded Another long-term storage method has been deve-
from direct sunlight and kept as cool as possible, loped by Van Impe (1992), in which the medusae are
should be examined under a stereo-microscope, and suspended in a solid agar-agar gel coloured with
the medusae individually transferred with wide- serva-blue, and from which extraction is easy when
mouthed pipettes to finger bowls of clear sea water. required. This method is particularly useful for trans-
After observation they can either be kept for rearing, portation and long term preservation. All holotypes
or anaesthetised and fixed as described below. This should be stored in such a gel, which to a certain
method is of course the most fruitful, allowing obser- degree also prevents drying and nicely stains the pro-
vation of characters of the living animal and perfect teinic medusae tissues blue.
fixation.
In most museum collections the majority of hydro-
Fixation and preservation medusae specimens, including the holotypes, have
been destroyed because label cards were placed in the
Hydromedusae should be anaesthetised before fixa- storage jars. After a few examinations or a single trip
tion. The animals are allowed to extend in a vessel of to a specialist, only the label remains. Cotton or paper
sea water where the anaesthetic substance is added materials in the container caps and lids, whose fibres
slowly, crystal by crystal or drop by drop. The com- adhere to and damage specimens, should also be avoid.
monest anaesthetics for marine medusae are menthol
crystals, propylene phenoxetol, and magnesium chlo-
ride (about 7.5% MgC12 6H20 in fresh water), the last Geographic distribution
being the most recommended. For general taxonomic
purposes hydromedusae can be fixed in 10% buffered In total, 197 species of hydromedusae have been lis-
"formalin" in sea water (40% commercially available ted from the South Atlantic, defined as 0" to 60" lati-
formaldehyde being considered as 100% "formalin"), tude, coast to coast. Of these, 149 are meropelagic
and preserved shortly in 5% formaldehyde. Buffering (neritic, epipelagic, or slope species), and 48 are
the preserving fluid with borax or calcium carbonate holopelagic. The distribution of species among higher
should be avoided, since the medusae may adhere to taxa (shown in Table 1) is rather different from that
any precipitate formed by these chemicals on the bot- observed in other seas, like the Mediterranean and the
tom of the containers. Hexamine should also be avoi- Indo-Pacific (Bismarck Sea). Comparison of the three
ded for this purpose because it destroys the mesoglea. areas shows that while the total number of species is
The best buffer seems to be sodium glycerophosphate. very similar, the number of Anthomedusae is always
Alcohol is to be avoided as a fixative because it cau- higher than that of Leptomedusae, the number of
ses specimens to shrink. Nevertheless, for long term Narcomedusae is highest in the Mediterranean
preservation (for instance in museum collections) for- (10.4%), and the numbers of Limnomedusae and
maldehyde, which causes auto-maceration of tissues, Trachymedusae are proportionally highest in the
is inadequate. It should be replaced gradually by 70% South Atlantic (7.6 and 14.3%, respectively).
Hydromedusae
Geographic area
South Atlantic Mediterranean Indo-Pacific
Subclass No. of No. of 1 No. of
O/O O/o O/o
SPP. SPP-
Anthomedusae 87 45.1 91 45.5
Leptomedusae 61 31.6
Laingiomedusae
Limnomedusae --
Narcomedusae
Trachymedusae
Total species
Table 1. Comparative data on the numbers of species within each subclass recorded in different oceanic areas. Data for the Mediterranean are
from Boero and Bouillon (1993), for the Indo-Pacific (Bismarck sea) from Bouillon et al. (1986).
Other comparisons indicate that: (1) 131 (66.5%) of the The eastern South Atlantic (West African coasts)
South Atlantic hydromedusae are also found in the hosts 109 species, the Western Atlantic (South
North Atlantic, 11 of them being restricted to the American coasts) 141 (135 + 6 Subantarctic endem-
Atlantic; (2) 85 (43.1%) of the South Atlantic hydro- ics). Forty-five (41.2%) of the Eastern Atlantic medu-
medusae are also found in the Mediterranean, and only sae are not known from the western part, and 77
1 (Amphinema rubra) is restricted to these two regions; (54.6%) species of the western fauna have not been
(3) 125 (63.5%) of the South Atlantic hydromedusae are found in eastern waters; 65 (33%) of the total number
also found in Indo-Pacific waters, and 16 of them are of medusae are common to the two areas, 2 of them
restricted to these two areas; (4) 68 (34.5 %) are found (Haliscera alba and Sarsia gracilis) are endemic to
in Atlantic, Mediterranean and Indo-Pacific waters, and the South Atlantic. Among the 45 species found only
may thus be considered cosmopolitan; (5) 50 (25%) in the eastern area, 20 (44%) are present in the North
species are found in Subantarctic or Antarctic waters; 41 Atlantic; 14 (3 1%) in the Mediterranean; 22 (48%) in
of these are also present in other sectors; 6 are endemic the Indo-Pacific; 6 (13%) in the three areas, and 8
to the Subantarctic area, 3 are endemic to the Antarctic; (18%) are endemic to this area. Among the 77 species
and 6 are bipolar; (6) 28 (14.2%) of the total number of found in the western waters only, 55 (71.4%) also
species are endemic to the South Atlantic. inhabit the North Atlantic; 29 (37.6%) the
Mediterranean; 40 (52%) the Indo-Pacific; 16
These results show that, although there are great simi- (20.7%) are found in the three areas, and 17 (22%) are
larities between the hydromedusae of the Northern and endemic to the western area. The western South
Southern Atlantic, the South Atlantic shares as many Atlantic medusa fauna seems thus to have more affin-
species with the Indo-Pacific fauna. All South Atlantic ities with the North Atlantic fauna, and to include
species found in the Mediterranean are also present in more endemics than the eastern one.
the North Atlantic, in the Indo-Pacific, or in both areas,
except for one (Amphinema rubra), which is present The general endemism of the South Atlantic medusa
only in the first two regions. The South Atlantic fauna fauna (14.2%) is lower than, for example, that of the
contains also a rather large Subantarctic and Antarctic Mediterranean (18%), which is one of the most
component, many of these species being epipelagic thoroughly studied areas in the world. The endemic
here and dwelling in deeper waters at lower latitudes. medusa fauna of the South Atlantic certainly contains
Hydromedusae 391
many false endemics. Among the 8 eastern South Navas-Pereira and Vannucci (1994) analysed the eco-
Atlantic "endemic" species for instance, 7 have been logical preferences of the Subantarctic and Antarctic
described only recently (Pagks et al., 1991, 1992), and hydromedusae for different water masses (between
it is very doubtful that they indeed are restricted to this 20°W in the Atlantic region and 145"E in the Pacific
area. Endemism often reflects either the scarcity and sector of the Antarctic). They confirmed earlier obser-
dispersion of observations, many of the species having vations showing that eurybathic, but stenohaline
been reported only once; or is linked to incomplete andlor stenothermal, species may often be found at the
knowledge of the species distribution. Furthermore, in surface in the Antarctic Ocean and at great depths at
some cases it may also be attributable to the question- equatorial latitudes or even north of the Equator; and
able validity of some species, resulting from inadequate that in the Southern Ocean the numbers of both indi-
descriptions (e.g., Haliscera alba, Euphysa monotenta- viduals and species of holoplanktonic forms exceed
culata, Irenium teuscheri, Solmissus atlantica, etc.). those of meroplanktonic ones. They also confirmed the
usefulness of hydromedusae as water mass indicators.
In the western South Atlantic the relationships between
hydromedusan distribution patterns and local water Most literature data confirm that on the average the
masses have been investigated by Vannucci (1957b, Subantarctic and Antarctic fauna is rather poor in
1963) and Goy (1979). Navas-Pereira (1981) studied Leptomedusae (15,7 %) compared to Anthomedusae
the distribution of the hydromedusae of the continental (23.5%), and that the Laingiomedusae and
platform of Rio Grande do Sul (Brazil). Zamponi and Limnomedusae seem to be absent from Antarctic
Genzano (1994) analysed the diversity, seasonality, and waters. As mentioned above, holoplanktonic forms are
abundance of the hydromedusae from Samborombon the most abundant in Subantarctic and Antarctic
Bay (Argentina). They observed a marked seasonality waters, representing 60.8 % of all species, 22.3% for
in the distribution, with peak densities in the summer the Narcomedusae and 38.5% for the Trachymedusae,
months (about 100 ind. m-3), and another important apparently the most characteristic groups in those
maximum in the autumn (>40 ind. m-3). areas.
Pagks et al. (1994) studied the composition and dis-

tribution of macro- and megaplanktonic hydromedu- Morphology, classification and identification
sae in the eastern part of the Weddell gyre (55O35.2'
to 66'53.8's) between 0 and 2000 m. Fourteen hydro- General morphology
medusae were identified, their abundance varying
from 0.21 to 30 specimens per lo4 m3, the most abun- The planktonic stages of the Hydroidomedusae or
dant being Pantachogon haeckeli. They found high hydromedusae are free swimming, solitary animals,
diversity at bathypelagic depths, but medusae were with essentially tetramerous radial symmetry (Fig.
most abundant at mesopelagic depths, reaching 46 1B). Their body, swimming bell, or umbrella has the
specimens per lo4 m3 in the 500-1000 m range. general form of a mushroom, bell, or disk (Fig. 1A, C,
Farther south, Pagks and Kurbjeweit (1994) studied 21). Most of its volume is occupied by a gelatinous
the abundance and distribution of mesoplanktonic mass, the mesoglea, the jelly of the jellyfish, which
medusae from the Weddell Sea (Antarctic Atlantic confers shape and buoyancy (Fig. lC, 21). The convex
sector, about 60°30' to 71°S), reporting 8 hydromedu- upper (aboral) umbrellar surface is called the exum-
sae, among which Pantachogon scotti (to 11.671 ind. brella (Fig. lA, C), and the concave lower (oral) sur-
1000 m-3) and Arctapodema ampla (to 960 specimens face is termed the subumbrella (Fig. lA, C). The space
1000 m-3) were the most abundant. enclosed by the umbrella is the subumbrellar cavity,
which is narrowed by a muscular horizontal dia-
The eastern South Atlantic has been investigated much phragm, or velum, leaving only a central circular aper-
less than the western part, except for the south-eastern ture, the velar opening (Fig. lA, C, 2A, D, H). The
sector, which is one of most productive areas in the velum plays an important role in the locomotion of the
world and where the distribution and ecology of hydromedusae. The free rim of the umbrella bears marginal
medusae have been studied by Pagks (1992), Pages et tentacles and sense organs (ocelli, statocysts, cordyli:
al. (1991), and Pagks and Gili (1991, 1992a, b, c). Fig. 1A, 2A, B, C, E). The tentacles may be solid (Fig.
Hydromedusae
Species
Aegina citrea w w H, E-B

Aeginura grimaldii w w H, B
Aequorea coerulescens M, N
Aequorea conica w M, N
Aequorea forskalea w w M, N
Aequorea macrodactyla I M, N
Aequorea pensilis w w M, N
Aglantha digitale H, E-B
Aglantha elata H, B
Aglaura hemistoma H, E
Aglauropsis agassizi M, N, T-St
Aglauropsis conanti M, N, Sa
Aglauropsis edwarsii
~ -
M, N, Tr
Aglauropsis ka wari M, N, St
Amphinema australis M, N
Amphinema dinema M, N
Amphinema rubra M, B, S
Amphogona apicata H, B
Amphogona apsteini H, E
Annatiara affinis M, B
Arctapodema ampla
Arctapodema antarctica
Blackfordia virginica
Botrynema brucei
Bougainvillia carolinensis M, N
Bougainvillia frondosa M, N
Bouga invillia macloviania M, N
Bougainvillia muscus M, N
Bougainvillia niobe M, N
Bougainvillia platygaster M, N, E
Bythotiara capensis M, N, Tr
Bythotiara murrayi M, B
Calycopsis bigelowi M, S
Calycopsis borchgrevinki M, E-B
Calycopsis chuni
Calycopsis papillata
Calycopsis typa
Chromatonema rubra
Cirrhitiara superba
Hydromedusae 393
Species
I
LY
E I
Cirrholovenra tetranema
Clyt~abrunescens
Clytra d~scorda
-
- w
-
-
-----
w
---
.-A
.
rn
8
1
1 :.i ----
I Clytia hemrsphaerrca -- 1
L 1
. I
Clytra lomae ---- -
Clytra nolrforme
1 I
1 --
.
Clytra ovale --- + M, N
- - - -- -
I i - I 4 I
Clytra phosphoricum w
Clytra simplex rn --
w 1 -- -- -- rn
- --
Colobonema sericeum
Cosmetirella davisi
Crossota alba
Crossota brunnea
Cubaia aphrodite
Cunina duplrcata
.
Cunina frugifera
Cun~naglobosa
i rn -
A
? i: rn I
-
I
l - - H, E
- --
.
Cunina octonarra ¤ H, E -
I
Cun~naperegrrna
Cytaeis sp.
Dipurena baukalion
I: - ----
---
rn
W
-
¤
1 - 1 .
¤
---
I
H,E
M, N
1
Dipurena halterata
Drpurena reesi
Ectopleura dumortieri
Eirene viridula
Eucheilota duodecimalis
- -- --
¤
- --- ---
rn
4
I
1 . i rn -
.
-
I
I I M, N
M,N
1
t
.. . . -4- 1
Eucheilota forest; 1 . M, N, s t - ~ a
r
Eucheilota maculata ---
M, N
- -
...
Eucheilota paradoxica -- -- -- ¤ M,N
Eucheilota ventricularis -- - L --- + - -
M, N-- 1
- --
Eucodonium brownei -
1
- + .
- 1- -r- M, N
--
1
I
Euphysa aurata I , +- W
- +I . 1 M,N ,
Euphysa monotentaculata
4- I I
I 1 M, N, Sa
..
-
Euphysora furcata 1 . 1
1 t - -
Euphysora gigantea
,+-
MLB - -
.. .
Euphysora gracilis M, N
I
Eutima coerulea
Eutima gracilis
--- I M, N
M, N
Eutima mira I . M, N
Hydromedusae
Species
-
- -
Eutonrna scrntrllans
- - --
1
--
7
-
--
' 1- T 1
1
7-
--
-
-
..
- -
-
-- M, N
.
rn rn
Geryonra proboscrdalrs
- -
Gossea brachyrnera
-- -- - - -
+-
I H
rn
- ---- --
¤
w
-
1 -
H, E
M, N
Gotoea srrnrlrs ¤ ~ I M, N, S
.. .
Halrcreas rnrnrrnurn I rn W ¤ rn rn H, B
Halrscera alba I H, B, St-Tr
-- -- ---
t
Halrscera conrca
. .. . .
rn
.. . rn ¤
H, B
H, B
..-.
Halrscera racovrtzae - -
1 . H, B
. ;..
Halitholus interrnedrus , H, E
- -
i
I .. .
Halitiara forrnosa ----- I M, N
-- ---- - -- +
- I
I
Halitrephes maasr
-- I H, B
.. . ..
Halopsis ocellata
-- -
M, N
I
Helgrcirrha schulzer ¤- - - - -
w M, N
Heterotiara anonyrna
-
M, N, E
. . ...
Heterotrara minor W M, N
Hornoeonerna platygonon
- - - - -- -
, L-
?.
---
¤ rn H, B
. . ..
Hybocodon prolrfer - M, N, S
- - - - -
- -- --- --
Hybocodon unrcus 1 rn rn M, N
.;
Hydractrnra rninrrna -- I M, N
--- - - -+-- + - - - - - + --
Hydractrnra rnrnuta rn rn rn M, N
Hydractrnra tenurs
--
t -
II I
I M, N, Sa
I I
I
t
Hydractinia tournrerr rn I M, N, T
-
t i r -
lreniurn teuscheri rn 1 M, N, T
1
Kantrella enrgrnatrca rn rn rn M, N
I
Koellrkerina fascrculata rn rn M, N
Laodrcea indrca
-
w w M, N
Laodrcea rnrnuscula rn M, N, St
Laodicea pulchra - -
M, N
Laodrcea undulata M, N
Leuckartrara adnata M, N, Tr
I Leuckartiara octona M, N
Liriope tetraphylla H, E
Lovenella cirrata
-- -
M, N
Malagazzra carolrnae
- -
M, N
Margalefia rnterrnedra M, N, Tr
Margelopsrs australrs M, N
Merga tergestrna M, N
Merga violacea rn 1 W M, N
Hydromedusae 395
Species *
.
3
-- -
.dm.,
- . 1 - -
-
I Mrtrocoma rnrnervae -- , -- 1 M,N I

Mitrocomella browner w I - M, N ,
Mrtrocomella frigida
-
--
I
I
t --
I M, N
Mrtrocomella grandls
- -
w-- -
I - .
M, N,Tr 1
M~trocomellamrllardae
L --
Modeerra rotunda w rn w
Moerrsia rnkerrnanrca
- 1 - rn
Neoturris prleata
-- --
.
Nrobla dendrotentaculata
-- -- M, N
,
.
Obelra d~chotoma w - -- M, N
.
Obelra longrssrma rn
- 1 - I -
-
Obelra spp.
-
Oceanra armata
- - - -- -- - -
i
Octobulbacea montehermosensrs M, N, Sa-
-
I -
.r '
-
-
Octocanna haeckeli M, N, St
1 Octophralucrum brgelowr
-
- rn rn
+--
M,N
I Olindras phosphorica
t
rn I M, N
Olrndias sarnbaqurensis -
- - I M, N, St ,
Orchrstoma collapsa
--
Orchrstoma prleus
....
I Pandea conica -
Pandea rubra -
Pantachogon haeckelr --
-
-
Pantachogon mrlrtare
- -
w - +-- / H, B ,
-
--
. .
Pantachogon scottr
'- H,
I B
-
N1q
--
I --
Paragotoea bathybra -
- - --
I .
- I --
?. M1
-
. . . -+-. .. .
Pegantha clara - N w - ,I I
-- H, E -
,
- -
- 4 --
Pegantha laevrs
-
Pegantha martagon
- I
1 rn w
- - - -- I I - H, E
, 1 H,E
1 4. . - - 1 .l
t L --
Pegantha rubrg~nosa 1 -- -4
1 H,E I
r . l
.. . ~
-2
Pegantha trrloba rn ¤
tW I -1 - - H, E 1
1 Persa rncolorata
. .. -7
W H, E-B
- 4 --- -
t
Phralella falklandrca --- w M, N -
-- - --- I + --
,
2
Phralopsrs dregensrs
- -
--- -- I I .
I
- t- -- M, N, E
--
Porprta porpita W W ¤
I -- - - - --- - 1 M,N,E-
.. .
Proboscrdactyla rnenonr I M, N , Tr
- 1
.
M, N, Sa
I . . I
I
I . , I - I -
I I M, N
I
- - 1 - - M, N
396 Hydromedusae
c
m
Species
5 me e
1
..
- - - L--
I -T-II'
- - --
7-- -7-
I
Pseudotrara troprca -Q--
- t
Ptychogastria polaris 1
-- - - - - - --
4 1 . 1 rn +
.
Ptychogena antarctica ¤ rn rn -,
1 . M, N
Ptychogena hyperborea ¤ ¤
.
I Ransonra krampr I .
- -- -
Rathkea africana
-
t --
4 - 7
- - - -- + I M, N, T
Rathkea formosissrma -
-- - - - ---- - k r
..
Rhabdoon srngulare
- - ¤ 1 M, B
I
--
Rhacostoma atlantrca
. -
. .
Rhopalonema velatum
+ - - - - ----
Russellra mrrabrlrs ---- + I., ¤ ¤ I M, S
.
Sarsia exrmra M, N
- -- -
i I -
Sarsra gracilis , ¤ I _ M , N , Tr-Sa
.
- - - - - --
..
1 Sarsra producta -
w w - M, N
I Srbogita geometrrca rn rn M, N
1 Smrnthea
-- -
eurygaster t .
- I .
- A-
H, E-B
Solmaris corona
Solmarrs flavescens
Solmissus atlantica
Solmrssus faberi
1
Solmrssus marshal11
-- - - - -- - -- -
1 Solmundella bitentaculata
Staurocladia capensis
Staurocladia hodgsonr
Teracanna octonema
Tetrorchrs erythrogaster
-
1 Thamnostoma tetrella I
, i1 1
.
I M, N, T
- + -
I
..
I
Traricodon coeruleus
Tiaropsidrum roseum
I
w
I . 1 I M, N
I-- ..I I I
1
.
I C--- I I
I. I
I
I
M, N, E
I
M, N , Sa
1 Vannuccia forbesrr
rn
rn I . M I~ M , N, T-St
M, N
Hydromedusae 397
z
L
I 5 53 zs
L
8C u B
s.; ".' 0 U' -2"
1
C U -
b'U g .V
EZ ;W 5 %
I
I
Species s m
5LS;
6 g Z 2s
3
EZC
C:
QS;
o
L~
zz
m
.U
*
<
2
L
.* -
=% 4m.; u
4: ,,-
"r
-0.s
0
I
i V ) < d 3
E -r ds c
Vellela vellela w w w w M, N
Zanclea sp. w I . rn w 1
M, N, B
Zancleopsrs d~chotoma w
i M, N
1 Zanclonia weldoni ¤ w 1 M, N
, Zygocanna vagans w w M, N
I Total species 109 135 39 30 131 16 85 125 35
Ecologic setting
A: Antarctic; B: bathypelagic; E: epipelagic;
Subtropical
E-B: epipelagic-bathypelagic; H: holopelagic;
M: meropelagic; N: Neritic; S: Slope; Sa: Subantarctic; 30'
St: Subtropical; T: Tropical; Tr: Transitional.
Transitional
45"
Subantarctic
! , -
Antarctic
1 60°
Table 2. General distributional data for the species recorded in the South Atlantic (refer to map for general pattern of climatic zones defined)
398 Hydromedusae
2H) or may contain an extension of the circular canal tance as a muscular introduction to the coelenteron
(tentacular cavity: Fig. 1A). The base of each tentacle or actinopharynx.
is commonly swollen into the tentacular bulb (Fig. Adaxial: facing towards the main axis, opposite to
lC, 2C, F, G, J). abaxial; on a marginal tentacle the inner tentacular
surface.
The manubrium, a tubular or quadrangular projection Adnate: having part or all of one side in contact with,
of variable length, hangs from the centre of the sub- or fixed to, another structure (e.g., abaxial side of a
umbrella as the clapper of a bell (Fig. lA, C, 21). The ' marginal tentacle fixed to the exumbrella,
cavity of the manubrium, or gastric cavity, opens dis- Leuckartiara adnata).
tally into the mouth (Fig. lA, C) and extends proxi- Adradial: the axes or sectors lying between the perra-
mally into the radial gastrovascular canals (Fig. lA, dial and interradial ones; in a medusa with 4 radial
C). These canals are generally 4 in number but are canals there are 4 perradial axes; 4 interradial axes,
sometimes more numerous. Through the mesoglea 8 adradial axes, and sixteen sectors (Fig. 1B).
they connect the gastric cavity to the circular canal, Apical knob or chamber: small protrusion of the base
which runs all along the marginal rim of the umbrella of the manubrium into the apical umbrellar meso-
(Fig. lA, C, 2C, G). The gastric cavity, the radial glea (e.g., some Sarsia).
canals, the ring canal, and the tentacular canals (when Apical or umbilical canal: during the development of a
they exist) form the gastrovascular system, which medusa bud, an opening provides continuity and
serves both for digestion and for distribution of food, exchanges between the "mother" gastric cavity and
waste, cnidoblasts, and even gametes. that of the bud (Fig. 1E). Generally this aperture
disappears after liberation, but in some medusae it
The mouth may be simple or provided with lips, remains as a small canal or duct projecting from the
lobes, or tentacles (Fig. 2L). The radii corresponding manubrium into the apical mesoglea, often leading
to the radial canals are termed perradii. The interradii upwards to the outside (e.g., Sarsia products,
lie intermediate between them, and the adradii mid- Coiymorpha nutans).
way between the perradii and the interradii (Fig. 1B). Apical projection or process: a rounded or pointed,
usually roughly conical, mesoglear extension of
The "cathamnal" or endodermal lamella, a unistratified the top of the umbrella (e.g., Amphinema,
membrane crossing the mesoglea, interconnects the Leuckartiara).
radial canals and, like them, connects the gastric cavity Athecata: term applied to the hydroids of the
with the circular canal. It delimits two mesogleal levels, Anthomedusae, all of which lack a proper hydro-
one subumbrellar, the other exumbrellar (Fig. lA, C). theca or chitinous cup partially or entirely surround-
The sex cells may develop and mature either on the ing the hydranths.
manubrium, or the radial canals, or both. Fertilisation is Bell: see umbrella.
usually external, but internal fertilisation may occur in Blind canal: centrifugal canals or radial canals are
a few species. The resulting embryo develops into a pla- termed blind when they are not connected with the
nula larva, which settles and metamorphoses into a new ring canal (e.g., Toxorchis); centripetal canals are
polyp stage, or more rarely directly into a new medusa. called blind when they do not join radial canals or
the manubrium (e.g., some Calycopsis).
The following illustrated glossary of terms most com- Capitate tentacle: tentacle having a knobbed end rich-
monly used in describing hydromedusae covers the con- ly armed with cnidocysts.
cepts and terms used in the keys and diagnoses below. Centrifugal canal (= radial canal): canal issuing from
the manubrium and directed towards the umbrella
Abaxial: away from the main axis or on a site remote margin.
from it; on a marginal tentacle the abaxial side is Centripetal canal: canal issuing from the circular canal
the outer tentacular surface. and directed to the manubrium (e.g., Calycopsis).
Actynopharynx: in Hydrozoa, Cubozoa, and Chordal or chordoid tentacle or structure: formed by a
Scyphozoa the ectoderm and endoderm meet at core of single disk-like or cylindrical cells placed
the mouth rim, in the Anthozoa the ectoderm of end to end in a single row. (e.g., solid tentacles,
the mouth rim is turned in for a considerable dis- Obelia; Fig. 2H).
Hydromedusae
upper endoderm
nerve
ring
exumbrella / sensory
eoithelium
endoderm
I velum
lower nerve
ring
radial gastrovascular
canal
- 1-7 1 endoderm
- - - I- - - of
endoderm - - -the
of
circular canal
the
I
1
p i mented cells
lens of of ectodermal
sensory cells a complex ocelli
of the ocelli ocellum
endoderm
mesoglea
derm
Hydromedusae
Hydromedusae
I Sarsia Clytia Cosrnetira Eirene 1
Turritopsis Hydractinia Lizzia Bougainvillia
Q
open marginal closed marginal
statocyst statocyst
statocyst sairal
I free ecto-edodermal enclosed ecto-edodermal

statocyst ori inating statocyst
from the circiar canal
exumbrellar man~brium
402 Hydromedusae
/
macrobasic eurytele
microbasic
mastigophore
atrichous &
isorhiza T
stenotele
@ ' isorhiza
(Narcomedusae) macrobasic
mastigophore
Hydromedusae
Circular or ring canal: simple canal which runs around this being an important taxonomic trait (Fig. 2M, P,
the umbrella margin linking the ends of the radial R, S). Cnidocysts are used for prey capture,
canals; occasionally the circular canal is not hol- defence, and attachment.
low but consists of a solid core of endodermal Cnidome: entire complement of cnidocyst types in
cells (e.g., Proboscidactyla, Laingiomedusae). In one species or in a genus.
the Narcomedusae, in which the umbrella margin Cnidophore: cnidocyst-filled capsules covered by
is deeply cleft into broad flaps, a circular canal numerous long ciliae and attached to tentacles by
may be present or not; when present the marginal elongated, filiform, and very contractile stalks of
canal follows the edge of the margin of the exum- special structure (e.g., Zanclea, Fig. 2Q), not to be
brellar flaps and is called "peripheral canal confused with branched tentacles.
system", the vertical parts of which are the peroni- Cordylus: minute marginal club-shaped structures si-
a1 canals (Fig. 1A, C, Fig. 2C, G). tuated on the umbrella margin between the tenta-
Cirri: small tentacle-like organs situated on the cles. They have a narrow peduncle and a thick dis-
umbrella margin between the true marginal tenta- tal portion. Cordyli may be hollow or completely
cles, devoid of swollen marginal bulbs and solid. filled by endoderm, with or without cnidocysts.
Two types are generally found: (1) spiral cirri: Their function is unknown, probably sensory (e.g.,
cirri which coil spirally and have a terminal clus- Laodiceidae).
ter of cnidocysts, which are the most common Crenulate: having low rounded cusps or lobes separ-
(e.g., Mitrocomella) (Fig. 2N); (2) flexile cirri: ated by sharp but shallow notches.
straight, not coiled, and having the cnidocysts in Diploblastic: composed of two epithelia, an outer
rings (ex Cosmetira) (Fig. 25). Cirri may be imme- ectoderm and an inner endodem, separated by a
diately adjacent to the marginal bulbs and are then relatively undifferentiated connective layer, the
termed lateral cirri (e.g., Eucheilota maculata) mesoglea (not a true tissue).
(Fig. 2F); they may also occur along the umbrellar Direct development: development where the medusa
margin in the inter-spaces between marginal tenta- stage gives rise to another medusa without passing
cles, in which case they are called marginal cirri through a hydroid phase (e.g.,Trachymedusae, some
(e.g., Phialopsis) (Fig. 25). Narcomedusae), or where a hydroid directly pro-
Clasp: embracing part of a marginal bulb (e.g., duces another hydroid (e.g., Hydra) (Fig.lD, top).
Leuckartiara) (see exumbrellar spur). Ecto-endodermal ocelli: photoreceptors found in the
Cnidocyst (= nematocyst or stinging cell): stinging Tiaropsidae where the cup-shaped mass of pig-
organelle characteristic of the Cnidaria, consisting ment is formed by the endoderm of the circular
of a capsule secreted by a cell called a cnidocyte, canal, the nerve elements being ectodermal. The
containing a refringent fluid and a coiled and fold- ocelli of other Hydroidomedusae are completely
ed tubule (thread) which everts and straightens on ectodermal in origin. In the Tiaropsidae the ocelli
discharge. According the structure of the internal are associated with open ectodermal statocysts
tube different types of cnidocysts are recognised, forming a compound sense organ (Fig. 2D).
4
Fig. 2. A: Section through the bell margin of Aequorea showing a closed statocyst. B: Cordylus of Laodicea. C: part of the bell margin of
Laodicea. D: Open statocyst with ecto-endodermal ocelli from Tiaropsis. E: Simple ocellum of Neoturris (left), and complex ocellum of Sarsia
(right). F: Part of the bell margin of Eutima coerulea. G: Part of the bell margin of Orchistoma pileus. H: Bell margin of the Narcomedusae
Pegantha rubiginosa. I: Side view of Pegantha rubiginosa. J: Portion of the umbrella margin of Cosmetira pilosella showing the flexile mar-
ginal cirri. K: Open statocyst of Mitrocoma. L: Diagrams of the mouth structure of different hydromedusae. M: diagram of undischarged and
discharged desmoneme cnidocyst. N: Portion of the bell margin of Microcomella bvownei showing the open statocyst and the marginal coiled
cirri. 0 : Diagrammatic figures showing the different types of statocysts. P: Microbasic eurytele cnidocysts. Q: Medusa of Zanclea sp. showing
the cnidophores and the exumbrellar cnidocyst bands. R: Macrobasic eurytele cnidocyst of Zanclea sp. S: Various common types of cnidocysts.
A, K: redrawn from Hertwig and Hertwig (1878); B: redrawn from Brooks (1895); C: redrawn from Kramp (1919); D, E (left): redrawn from
Linko (1900); E (right): redrawn from Hyman (1940); F, G, H, I: redrawn from Mayer (1910); J, L, N, 0 : redrawn from Russell (1953); M, P,
R, S (macrobasic mastigophore, microbasic mastigophore, stenotele): redrawn from Weill (1934); Q: redrawn from Allman (1871-1872); S
(apotrichous isorhiza, atrichous isorhiza): redrawn from Bouillon (1995).
404 Hydromedusae
Ecto-endodermal statocyst: club-like organs for orien- pore in contact with the cavity of the bulbs, or of
tation and equilibration, growing out of the the gastro-vascular system. Regarded as having an
umbrella margin in the fashion of a tentacle; each excretory function.
is formed by an endodermal axis originating from Excretory pore: opening of the excretory papillae.
the circular canal and covered by the umbrellar Sometimes there are no papillae and the pores are
ectoderm. In their distal portion there are one or only slits.
more large endoderm cells (lithocytes), each con- Exumbrella: upper, aboral convex surface of the
taining a solid concretion (statolith); these struc- ' umbrella (see umbrella) (Fig. lA, C).
tures are termed free ecto-endodermal statocysts Exumbrellar cnidocyst cluster or band: exumbrellar
(e.g., Narcomedusae, some Limnomedusae and cnidocysts confined to specialised tissue in form
Trachymedusae; Fig. 20, 3.154), but in some spe- of oval, club-shaped, spoon-shaped, or elongated
cies the sensory clubs are entirely enveloped by an patches immediately above the marginal bulbs
ectodermal vesicle and are then called closed ecto- (Zanclea, Fig. 2Q) or between the marginal tenta-
endodermal statocysts (e.g., some Limnomedusae cles (Proboscidactyla).
and Trachymedusae) (Fig. 2H, I, 0 ) . Exumbrellar spur: the marginal tentacular bulbs may
Ectoderm: outermost cellular layer. grow upwards for a short distance and clasp the
Ectodermal statocyst: entirely ectodermal marginal margin of the umbrella forming an exumbrellar
organs sensing orientation and equilibration locat- spur (e.g., Leuckartiara).
ed in depressions or pockets of the velum. The Filiform tentacle: a tentacle which is straight-sided
may remain open (open ectodermal statocysts; throughout, lacking prominent cnidocyst clusters
e.g., Mitrocomidae, Tiaropsidae) (Fig. 2D, K, 0 ) ; along its length and terminally, the cnidocysts
or the velar tissue can seal the sense organ com- being evenly distributed.
pletely (closed ectodermal statocysts; e.g., the Gastric peduncle or peduncle: in some medusae, a
other Leptomedusae) (Fig. 2A, F, 0 ) . Each stato- cone-shaped thickening of the subumbrellar meso-
cyst contains one or more tiny polygonal or spher- glea projecting centrally downwards into the sub-
ical concretions (statolith) (Fig. 2A, F), the closed umbrellar cavity, to the end of which the manubri-
statocyst having a basal cushion of cells with sen- um is attached, the radial canals running down the
sory ciliae (Fig. 2A). peduncle to the manubrium at its end. Variable in
Endoderm: innermost cellular layer, lining the gastro- shape and size (e.g., long and narrow in Eutima
vascular cavities (Fig. lA, C). gracilis; large and pyramidal in Bougainvillia
Entocodon (=glockenkern or medusary nodule): one of macloviana; very short in Pvoboscidactyla stellata).
the most important and characteristic features of Gastrovascular system: the coelenteron or enteron,
Hydrozoan development. In the morphogenesis of comprising the manubrium cavity and the gastro-
medusa buds, of fixed gonophores or of eumedu- vascular canals, i.e. the radial and circular canals
soids, an invagination of the ectoderm of the apical and their derivatives.
budding zone produces a solid multistratified nod- Gonad: there are no real reproductive organs in
ule, the entocodon, which subsequently develops a Hydrozoa, so this term is inappropriate although
cavity: the future subumbrellar cavity. As a gener- widely used. In medusae it indicates the place
al rule the entocodon gives rise to ectodermal com- where the sex cells become mature; this may hap-
ponents only (the manubrial ectoderm; the subum- pen on the manubrium walls andor at the level of
brellar ectoderm and the internal layer of the velum the radial canals. The position of the germ cells
and their striated muscles); the endodermal compo- has considerable value as a diagnostic character.
nents of the buds (manubrium, gastro-vascular When the "gonads" are on the manubrium they
canals) being formed by an evagination of the may completely surround it, being cylindrical, or
"mother" endoderm (spadix) (Fig. 1E). occupy an interradial, adradial, or perradial posi-
Excretory papillae: papillae in some medusae situated tion. When located on the radial canal, they usual-
either between the marginal tentacles, at the base of ly develop on the lateral walls of the canals, but in
some marginal structures (tentacular bulbs, non- some medusae they are continuous also over the
tentacular or rudimentary bulbs, or marginal warts), ventral wall (e.g., Clytia hemisphaerica). Their
or on the radial canals. They provide an opening or position along the course of the radial canals,
Hydromedusae 405
shape and size are often a diagnostic character Marginal wart or swelling: small wart-like swellings
(Fig. 1A, C). of the umbrella margin never carrying tentacles
Gonophore: in many Hydroidomedusae the medusa (e.g., Eutima mira) (Fig. 2F).
stage is reduced to a varying degree and is no Medusa budding: asexual budding forming medusae.
longer liberated, remaining attached to the hydro- This is a common phenomenon in Hydromedusae;
id, in these cases it is called a gonophore or spor- the medusa buds are formed either on the manu-
osac or, since it is not released, a fixed sporosac. brium, the radial canals, the marginal bulbs, or the
Gonotheca: chitinous structure surrounding a gono- subumbrellar rim.
phore. Medusary nodule: see entocodon.
Hollow tentacle: tentacle either with a central cavity Mesentery: in some species a perradial tissue layer
connected with the circular canal cavity, or with- attaching the lateral walls of the manubrium to the
out a cavity but with an endodermal core formed subumbrella.
by several peripheral rows of cells (parenchymat- Mesoglea: the acellular substance lying between the
ic).The basal regions of such tentacles often dis- ectoderm and the endoderm and forming the gelat-
close central cavities (Fig. 2G). inous bulk of the umbrella in the Hydromedusae
Hydranth: the feeding polyp of a hydroid colony. (the jelly of jellyfish) (Fig. lA, C, 21).
Hydroid: the polyp stage in the life cycle of Moniliform tentacle: tentacles with a terminal knob of
Hydroidomedusae (Fig. 1D). cnidocyts and with conspicuous clumps of cnido-
Hydrotheca: chitinous structure surrounding a hydranth. cysts in bands spaced rather regularly along their
Hypostome: terminal region of a hydranth, from length.
which the mouth opens (Fig. IF, G). Mouth arm expansions: dilatations of the mouth rim
Interradial: radial axes lying between the perradii, armed with terminal cnidocyst clusters (e.g.,
between the radial canals. Hydractiniidae) (Fig. 2L).
Lateral cirri: see cirri (Fig. 2F). Mouth: opening of the manubrium to the exterior,
Lip: lobe-like extension of the manubrium margin may be simple and circular, or with simple or
surrounding the mouth opening (see mouth).The complicated lips (see lips) (Fig. lA, C).
lips may be of simple or complicated structure, Nematotheca: small chitinous theca surrounding
crenulated, folded, short or elongated, pointed or defensive polyps or nematophores.
rounded, armed with cnidocysts distributed uni- Non-tentacular marginal bulb: sometimes marginal
formly or in clusters, or devoid of them. In the bulbs without tentacles are developed on the
Rathkeidae the lips are elongate, simple, or umbrella margin. Either they never developed ten-
branched and armed with terminal, and usually tacles, or evolved by reduction of the tentacles. It is
also lateral, cnidocyst knobs (Fig. lC, 2L). necessary to distinguish between bulbs which never
Manubrial or gastric pouch or pocket: lateral perradial have tentacles (permanent non- tentacular marginal
or interradial extensions of the manubrial wall (e.g., bulbs, rudimentary marginal bulbs, e.g., Cirrhitiara
Narcomedusae, Tiarannidae, Gotoea) (Fig. 1A). superba, Aequorea macrodactyla), from those on
Manubrium: median projection of the subumbrella sur- which marginal tentacles will develop later on
rounding the gastric cavity, distally bearing the ter- depending of the growth of the medusae (develop-
minal mouth and proximally leading to the radial ing tentacular marginal bulbs, e.g., Malagazziidae).
canals. The manubrium is very variable in shape Ocelli: photoreceptor found in some Hydroidomedusae,
and size: tubular, cruciform, quadrate, hsifom, bar- most common in Anthomedusae, usually situated
rel-shaped, flask -shaped, short, long, narrow or on the marginal bulbs in abaxial or adaxial position.
very large, etc. (Fig. lA, C, 21). Exteriorly they appear as round, oblong, or elongat-
Marginal cirri: see cirri. ed black, brown, yellow or red spots. They consist
Marginal lappet: one in a series of lobe-like exten- of a small mass or cupule of pigmented cells asso-
sions around the umbrellar margin (e.g., ciated with nerve cells, with or without a lens. Ocelli
Narcomedusae) (Fig. 2H, I). are of ectodermal origin, except in the Tiaropsidae
Marginal tentacle: a tentacle located at the edge of the (see ecto-endodermal ocelli) (Fig. lA, 2C, D, E, G).
umbrella. Octant: an eighth of the umbrella, the space between
Marginal vesicle: see statocyst. the interradii in a medusa with 4 radial canals.
406 Hydromedusae
Oral: near the mouth or related to it (the opposite end to the circular canal, except in a few species where
is termed aboral). growth is centripetal (e.g., Melicertum; the centrip-
Oral arms: outgrowths of the mouth margin armed etal canals) (Fig. lA, C, Fig. 2C, G).
with cnidocyst clusters. Ring canal: see circular canal.
Oral tentacle: in some medusae with a circular mouth Rudimentary bulb: see non-tentacular marginal bulb.
there are oral tentacles arising above the mouth Solid tentacle: tentacle without any a cavity, with an
rim. In the Cytaeididae they are simple and locat- endodermal core formed by a single row of disk or
ed just above the mouth rim, in the ' cylinder-like vacuolated cells placed end to end
Bougainvilliidae they are simple or branched and (see chordal) (Fig. 2H).
situated well above the mouth rim (Fig. lA, 2L). Spadix: the central core formed by an evagination of
Otoporpae: in some Narcomedusae, vertical, elongated, the "mother" endoderm covered by entocodonial
oval or even rounded ectodermal tracts with bristles ectoderm, which forms the manubrium in a medu-
and cnidocysts running up from each statocyst over sa, or on whose surface sex cells ripen in gono-
the exumbrellar margin (Fig. 2H, I, 3.154). phores (Fig. 1E).
Peripheral canal system: see circular canal. Statocyst: see ectodermal statocyst and ecto-endoder-
Perisarc: the chitinous exoskeleton of a hydroid. ma1 statocyst.
Peronia: in Narcomedusae the tentacles leave the Subumbrella: see umbrella.
umbrella at some distance along the margin just Subumbrellar cavity: see umbrella.
above the clefts separating the marginal lappets. At Subumbrellar surface: see umbrella.
the edges of the clefts the subumbrellar and exum- Tentacle: see marginal tentacle.
brellar ectoderm fuse without interposition of meso- Tentaculae: small solid marginal tentacles located
glea, forming the peronial grooves, which are between normal hollow tentacles (e.g.,
invaded by tentacular ectoderm, together making up Amphinema rugosa).
ectodermal strands rich in cnidocysts, muscles, and Tentacular marginal bulb: in most medusae, a dilated
nerves: the peronia. At the base of the peronia the portion of the proximal part of a marginal tentacle,
margin of the umbrella lappets remain curved, giv- next to the umbrellar margin (Fig. lC, 2C, F, G).
ing the umbrella its lobed appearance (Fig. 2H, I). The marginal tentacle bulbs contain a cavity in
Peronial canal: in Narcomedusae the part of the communication with the circular canal and with the
peripheral canal system running vertically along tentacular cavity of the tentacles when they are hol-
the peronia (see circular canal). low. They are variable in shape, most bulbs are
Perradial: the main radial axes of a medusa, cor- simple but in some medusae they are compound,
responding in most species to the radial canals and several tentacles may arise from a single ten-
(Fig. 1B). tacular bulb (e.g., Bougainvillia). During the
Planula: typically the primary free swimming gastru- growth of a medusa a new marginal tentacle is nor-
la larva of the Hydroidomedusae (Fig. ID). mally preceded by the formation of a bulb on
Polyp: basic individual of the hydroids, may be isola- which it develops (see marginal bulb). In some
ted or form colonies; may be of different types, medusae, however, there are no tentacular bulbs
e.g., hydranths, gonozooids, dactylozooids. (e.g., Calycopsidae, Gossea, Proboscidactyla).
Radial canal: canals in medusae leading from the per- Tentacular roots: projection of the endodermal tenta-
radial corners of the manubrium to the circular cular core into the umbrellar mesoglea (Black$ordia,
canal. Usually they are straight and narrow with Obelia, some Narcomedusae) (Fig. 2H).
smooth sides, but in some species they are large, Tentaculiform structure: solid marginal structures
ribbon-like (e.g., Amphinema), and with a jagged resembling tentaculae but without marginal bulbs
outgrowth (e.g., Leuckartiara). Their typical num- and without any contact with the circular canal
ber is four, but can be higher in many medusae, up (exclusively in the Orchistomidae) (Fig. 2G).
to more than a hundred (e.g., Aequorea). They are Theca: chitinous protective sheath of a polyp.
normally simple, but can be branched in some spe- Thecata: name for the hydroid stage of the
cies, the branches sometimes not reaching the cir- Leptomedusae, all of which usually have thecae
cular canal (e.g., Staurodiscus). Generally the radi- protecting their polyps.
al canals grow centrifugally, from the manubrium Umbilical canal: see apical canal.
Hydromedusae
Umbrella: (=bell) main body of the medusa, general- Family Pandeidae Haeckel, 1879
ly resembling a bell, excluding manubrium and Family Protiaridae Haeckel, 1879
tentacles. The outer, usually convex, surface of the Family Russelliidae Kramp, 1957
umbrella is the exumbrellar surface (=exumbrel- Order Capitata Khun, 1913
la), the inner concave surface is the subumbrellar Suborder Moerisiida Poche, 1914
surface (=subumbrella), and the cavity bounded Family Moerisiidae Poche, 1914
by the subumbrellar surface is the subumbrellar Family Polyorchidae A. Agassiz, 1862
cavity (Fig. lA, C, 21). Suborder Tubulariida, Fleming, 1828
Velum: horizontal fold projecting inwards from the Family Corynidae Johnston, 1836
umbrellar margin leaving a central, circular aper- Family Corymorphidae Allman, 1872
ture, the velar opening. It consist of two layers of Family Eleutheriidae Russell, 1953
ectoderm separated by a mesogleal lamella, the Family Euphysidae Haeckel, 1879
inner ectoderm, of subumbrellar origin, with stri- Family Margelopsidae Uchida, 1927
ated muscles. The velum serves in propulsion and Family Tubulariidae Fleming, 1828
orientation of the medusa, acting similar to a pho- Suborder Zancleida Russell, 1953
tographic diaphragm, the diameter of its aperture Family Porpitidae Goldfuss, 1818
is adjustable during swimming, from as wide as Family Zancleidae Russell, 1953
the umbrella to almost closed (Fig. lA, C, 2A, D, Family Zancleopsidae Bouillon, 1978
H, 3.154). Subclass Leptomedusae Haeckel, 1886
Family Aequoreidae Eschscholtz, 1829
Outline classification Family Blackfordiidae Bouillon, 1984
Family CirrholoveniidaeBouillon, 1984
The phylum Cnidaria can be classified as follows Family Eirenidae Haeckel, 1879
(Bouillon, 1985a, 1995; Bouillon et al., 1992): Family Eucheilotidae Bouillon, 1984
Family Laodiceidae Agassiz, 1862
Phylum Cnidaria Verrill, 1865 Family Lovenellidae Russell, 1953
Subphylum Anthozoaria Petersen, 1979 (devoid of Family Malagazziidae Bouillon, 1984
medusa phase) Family Mitrocomidae Haeckel, 1879
Superclass Anthozoa Ehrenberg, 1833 (part); Torrey, 1909
Subphylum Medusozoa Petersen, 1979 (medusa Family Orchistomidae Bouillon, 1984
stage important in the life cycle, although it may be Family Phialellidae Russell, 1953
secondarily lost) Family Tiarannidae Russell, 1940
Superclass Scyphozoa Goette, 1887 Family Tiaropsidae Boero, Bouillon
Superclass Cubozoa Werner, 1973 and Danovaro, 1987
Superclass Hydrozoa Owen, 1843 Order Proboscoida Broch, 1910
Class Siphonophora Eschscholtz, 1829 Family Campanulariidae Jonhston,
Class Hydroidomedusae Bouillon, Boero, 1836
Cicogna, Gili and Hughes, 1992 Subclass Laingiomedusae Bouillon, 1978
Subclass Anthomedusae Haeckel, 1879 Family Laingiidae Bouillon, 1978
Order Filifera Kuhn, 1913 Subclass Limnomedusae Kramp, 1938
Suborder Margelina Haeckel, 1879 Family Olindiidae Haeckel, 1879
Family Bougainvilliidae Liitken, 1850 Family Proboscidactylidae Hand and
Family Clavidae McCrady, 1859 Hendrickson, 1950
Family Cytaeididae L. Agassiz, 1862 Subclass Narcomedusae Haeckel, 1879
Family Eucodoniidae Schuchert, 1996 Family Aeginidae Gegenbaur, 1857,
Family Hydractiniidae L. Agassiz, 1862 emend. Maas, 1904
Family Rathkeidae Russell, 1953 Family Cuninidae Bigelow, 1913
Suborder Tiarida Haeckel, 1879 Family Solmarisidae Haeckel, 1879
Family Calycopsidae Bigelow, 1913 Subclass Trachymedusae Haeckel, 1866
Family Niobiidae Petersen, 1979 Family Geryoniidae Eschscholtz, 1829
Hydromedusae
Family Halicreatidae Fewkes, 1886 Subclass Limnomedusae

Family Ptychogastriidae Mayer, 1910 Medusae with gonads either on manubrium, on manu-
Family Rhopalonematidae Russell, brial wall with perradial continuation along radial
1953 canals, or on radial canals only. Marginal tentacles
peripheral, hollow. Marginal sense organs, when
present, ecto-endodermal closed statocysts. Hydroids
Keys and diagnoses for identification of hydro- solitary or colonial, small, sessile, with or without ten-
medusae of the South Atlantic tatles, without perisarcal thecae.
Class Hydroidomedusae Bouillon, Boero, Cicogna, Subclass Narcomedusae

Gili and Hughes, 1992 Medusae usually flattened, with central lens-shaped
mass of mesoglea and much thinner sides. Margin of
Subclass Anthomedusae umbrella divided by peronial grooves so umbrella
Medusae typically bell-shaped. Gonads confined to margin is lobed. Tentacles solid, inserted on exum-
manubrium, sometimes extending to most proximal brella at some distance from margin, just above pe-
parts of radial canal. Marginal sense organs, if present, ronial grooves; sometimes small secondary tentacles
are ocelli, never statocysts or cordyli. Marginal tenta- on margin itself. Manubrium very broad and short,
cles peripheral, hollow or solid. Hydroids athecate, with entire circular periphery, or with perradial or
body not covered by rigid perisarc. Cnidome normal- interradial peripheral pouches. Generally without radi-
ly with desmonemes (Fig. 2M). al canals; circular canal absent or looped into marginal
flaps to form a "peripheral canal system". Gonads on
Subclass Leptomedusae manubrium walls and/or on manubrial pouches.
Medusae flatter than a typical bell, the umbrella usu- Marginal sense organs free ecto-endodermal stato-
ally hemispherical or discoidal. Gonads confined to cysts. With or without otoporpae. Cnidome atrichous
radial canals, exceptionally extending to proximal part isorhizae and apotrichous isorhizae (Fig. 2s).
of manubrium. Marginal sense organs, when present, Reproduction by direct development, by reduced
ectodermal velar statocysts, rarely cordyli, occasional- polyps, or by complex parasitic larval development.
ly ocelli. Marginal tentacles peripheral and hollow
(except in Obelia). Cnidome often microbasic masti- Subclass Trachymedusae
gophores. Hydroids thecate, generally protected by Medusae with hemispherical or deep bell-shaped
rigid perisarc formed by hydrotheca, nematotheca, and umbrella. Margin entire with thickened peripheral
gonotheca, rarely with naked hydranths. cnidocyst ring. Radial canals and circular canal
present. With solid marginal tentacles or with mixture
Subclass Laingiomedusae of solid and hollow ones. With or without centripetal
Medusae with almost hemispherical umbrella divided canals. Manubrium with or without peduncle. Gonads
by peronial grooves or similar structures so that usually on radial canals. Marginal sense organs as
umbrellar margin is lobed. Four radial canals; no ty- open or closed ecto-endodermal statocysts. Cnidome
pical circular canal, but solid core of endodermal cells generally stenoteles associated with microbasic
around umbrellar margin. Tentacles solid, inserted on euryteles and/or atrichous isorhizae (Fig. 2P, S).
exumbrellar surface well above margin so that tenta- Reproduction by direct development, without polyp
cular bulbs are not in direct contact with endodermal phase.
circular core. Narrow exumbrellar cnidocysts bands
may alternate with tentacles. Manubrium simple, elon- Subclass Anthomedusae
gate, quadrangular or tubular; mouth opening quad-
rangular to circular. Gonads in four masses on proxi- Order Filifera
mal part of manubrium, or as epidermal lining of Medusae with gonads forming separate interradial,
interradial pockets of manubrium. Marginal sense adradial, or perradial longitudinal masses on walls of
organs apparently missing. Cnidome: macrobasic manubrium (exceptionally encircling entire manubri-
mastigophores (Fig. 2s). Reproduction unknown. um). Mouth either with four simple or complex lips,
or with circular mouth surmounted by oral tentacles.
Hydromedusae 409
Marginal tentacles solid or hollow. Cnidome inclu- manubrium in adradial, interradial, or perradial posi-
ding usually desmonemes and microbasic euryteles, tion; with or without ocelli.
never stenoteles. Hydroids with filiform tentacles 1 With well developed peduncle. Umbrella 13 mm
(except in dactylozooids of Ptilocodiidae). wide, 15 rnrn high, cylindrical with quadrangular
margin, with fairly thick walls and rounded top;
Suborder Margelina manubrium short, on broad conical gastric pedun-
Medusae with solid tentacles; ocelli when present cle; oral tentacles with very short trunk, divided 5-
adaxial; with oral tentacles armed with cnidocyst clus- 7 times; gonads slightly folded, extending along
ters, or with mouth bearing arms, knobs, or clusters perradial lobes of manubrium upwards on gastric
armed with cnidocysts. peduncle; marginal bulbs V-shaped, about half as
1 With oral tentacles ................................................2 wide as interradial space; each marginal bulb with
la Without oral tentacles .......................................... 3 35-65 tentacles in double row; ocelli yellow, red
2 Oral tentacles simple, situated on or very near to or brownish-black: Bougainvillia macloviana
mouth rim: Cytaeididae (Lesson, 1830) (Fig. 3.1)
2a Oral tentacles simple or branched, distinctly l a Peduncle absent or weakly developed .................2
inserted above mouth rim: Bougainvilliidae 2 Marginal tentacles with adaxial ocelli .................3
3 Mouth with 4 distinct lips ....................................4 2a Marginal tentacles without ocelli. Umbrella 2 rnrn
3a Mouth with 4 inconspicuous lips, each containing high, higher than wide, dome-like with vertical
a group of about 100 cnidocysts: Eucodoniidae sides, mesoglea fairly thick; manubrium flask-
4 Mouth rim and lips covered with a continuous row shaped, short, cruciform in section; oral tentacles
of cnidocyst clusters along margin: Clavidae with long basal trunk, divided 2-3 times; marginal
4a Mouth lips elongated to form mouth arms with bulbs small, each with 2 long tentacles, without
one or many distinct cnidocyst clusters ...............5 ocelli; 8 adradial gonads, mature females with
5 Marginal tentacles isolated: Hydractiniidae planulae: Bougainvillia frondosa Mayer, 1900
5a Marginal tentacles in 8 groups: Rathkeidae (Fig. 3.2)
3 Basal trunk of oral tentacles short. Umbrella 12 rnrn
Family Bougainvilliidae wide and high, globe-shaped to cubic, with thick
Usually bell-shaped; with short manubrium; with sim- walls, flat top, mesoglea thick; exumbrella with per-
ple circular mouth; oral tentacles simple or dichoto- radial notches; manubrium quadrangular, very flat,
mously branched, inserted distinctly above mouth rim 4 times wider than high; oral tentacles divided 5-6
and armed with cnidocyst clusters; with 4 radial times almost from base; gonads flat, as interradial
canals and circular canal; with solid marginal tenta- pads; marginal bulbs triangular, with 10-13 short
cles either solitary or in clusters, borne on 4, 8, or 16 tentacles; ocelli crescent-shaped; medusa buds pro-
tentacle bulbs; with gonads on manubrium either duced directly from manubrium or from polypoid
forming a continuous ring, or on adradial, interradial, structures developed on manubrium: Bougainvillia
or perradial axes; with or without adaxial ocelli. plafygaster (Haeckel, 1879) (Fig. 3.3)
1 Oral tentacles dichotomously branched; marginal 3a Basal trunk of oral tentacles long .........................4
tentacles solitary: Thamnostoma 4 Manubrium short and broad. Umbrella 2-3.5 rnm
la Oral tentacles dichotomously branched; marginal wide and high, semiglobular, mesoglea fairly
tentacles in 4 or 8 groups .....................................2 thick; oral tentacles divided 1-2 (rarely 3-6) times;
2 Marginal tentacles in 4 groups, all alike: 4 interradial gonads reaching perraddii, globular
Bougain villia in females, prolonged along perradial side of
2a Marginal tentacles in 8 groups, all alike: peduncle in males; marginal bulbs small, with 3-5
Koellikerina (rarely 6-7) long marginal tentacles; ocelli round:
Bougainvillia muscus Allman, 1863 (Fig. 3.4)
Genus Bougainvillia Lesson, 1836 4a Manubrium elongate, narrow .............................. .5
Bougainvilliidae with 4 radially placed clusters of 5 Oral tentacles divided twice, 4 interradial gonads.
solid marginal tentacles; tentacles of each cluster all Umbrella 4 mm wide and high, dome-shaped,
alike; with 4 perradial oral tentacles dichotomously walls very thick; manubrium long and narrow;
branching in normally developed medusae; gonads on marginal bulbs small, bulbous, each with 7-9 slen-
410 Hydromedusae
der, stiff tentacles; ocelli large: Bougainvillia Oceania armata Kolliker, 1853 (Fig. 3.9).
carolinensis (McCrady, 1859) (Fig. 3.5) Umbrella 8-10 mm wide and high, bell-shaped,
5a Oral tentacles divided 4 times; 8 adradial gonads. with flat top, walls uniformly thin; manubrium
Umbrella 5 mm wide, 7 mm high, with vertical flask-shaped, cruciform in transverse section;
sides and flatly rounded apex, walls thick; manu- mouth rim crenulated; 100-200 marginal tentacles,
brium, wide, flask-shaped, cross-shaped in sec- densely crowded, marginal bulbs elongated, alter-
tion, about half as long as bell margin; marginal nately slightly displaced adaxially and abaxially.
bulbs small, oval, each with about 8 tentacles;
with dark ocelli; medusa buds developing from Genus Turritopsis McCrady, 1859
manubrium: Bougainvillia niobe Mayer, 1894 Clavidae with manubrium mounted upon pseudo-
(Fig. 3.6) peduncle formed by highly vacuolated endodermal
cells.
Genus Koellikerina Kramp, 1939 Turritopsis nutricula McCrady, 1859 (Fig. 3.10).
Bougainvilliidae with 8 groups of marginal tentacles, Umbrella 4-1 1 mm high, bell-shaped, higher than
all alike in structure; with 4 oral perradial, dichoto- wide, walls uniformly thick; manubrium large,
mously branched tentacles; gonads on manubrium in cross-shaped in transverse section; 4 radial canals
adradial, interradial, or perradial position; with or that continue into 4 compact vacuolated endoder-
without ocelli. ma1 masses situated above digestive part of manu-
Koellikerina fasciculata (Ptron and Lesueur, brium; 80- 120 closely spaced marginal tentacles;
1810) (Fig. 3.7). Umbrella 12 mm wide, 12 mm gonads interradial, mature females often with
high, barrel-shaped to bell-shaped, with flatly developing embryos and planulae.
rounded apex and thick walls; manubrium on
short, broad gastric peduncle with quadrangular Family Cytaeididae
base; oral tentacles divided 7 times; 4 perradial Anthmedusae with bell-shaped umbrella; manubrium
horseshoe-shaped gonads with transverse furrows; bulbous, with simple, circular mouth; with 4 or more
8 marginal bulbs, each with 10-23 tentacles; with unbranched oral arms on, or very near, mouth rim;
dark red ocelli. with 4 radial canals and circular canal; 4 or 8 margi-
nal solid tentacles; gonads in interradial position or
Genus Thamnostoma Haeckel, 1879 encircling manubrium; without ocelli.
Bougainvilliidae with 4, 8 or more solitary marginal
tentacles and 4 branched oral tentacles above mouth; Genus Cytaeis Eschscholtz, 1829
gonads interradial; with or without ocelli. With characters of family, with only 4 marginal tenta-
Thamnostoma tetrella (Haeckel, 1879) (Fig. 3.8). cles.
Umbrella 4 mm wide, 6 mm high, egg-shaped; Cytaeis sp. (includes several described polyp spe-
manubrium cubical; oral tentacles divided 3 times; cies with very similar medusae) (Fig. 3.11).
4 marginal tentacles; with ocelli. Umbrella up to 5 mm wide, 6 mm high, pear-
shaped to globular, apical mesoglea about twice as
Family Clavidae thick as lateral walls; with or without slight pedun-
Umbrella bell-shaped; short manubrium; gastric gela- cle; manubrium large, pear-shaped; mouth with up
tinous peduncle or vacuolated endodermal cells form- to 32 simple, more or less capitate and adnate oral
ing a pseudo-peduncle; mouth with 4 simple lips, tentacles; with 4 broad radial canals; marginal ten-
armed with continuous row of sessile cnidocyst clus- tacle bulbs large, pyriform to triangular, attached
ters along whole margin; 4 radial canals and circular to exumbrella above tentacles; medusa buds on
canal; solitary solid tentacles, numerous in adults; go- base of manubrium.
nads on interradial walls of manubrium; adaxial ocelli.
Family Eucodoniidae
Genus Oceania Kolliker, 1853 Umbrella bell-shaped, with thickened apex; manubri-
Clavidae with manubrium mounted on short, solid, um cylindrical; with conical gastric peduncle; mouth
pyramidal, gelatinous peduncle without endodermal quadrangular, with 4 inconspicuous lips armed with
vacuolated cells. cnidocysts; with 4 radial canals and circular canal;
Hydromedusae
gonads encircling manubrium; with 4 solid marginal subconical, with thick apex; manubrium very
tentacles with terminal swelling; marginal bulbs extensible; medusa buds on manubrium; gonads
small; without ocelli. undescribed: Hydractinia tournieri (Picard and
Rahrn, 1954) (Fig. 3.14)
Genus Eucodonium Hartlaub, 1907 3 With 4 marginal tentacles. Umbrella 0.3-1 mm
With characters of family. wide and high, globular: Hydractinia minima
Eucodonium brownei Hartlaub, 1907 (Fig. 3.12). (Trinci, 1903) (Fig. 3.15)
Umbrella up to 1 mm high and wide; mouth lips 3a With 8 marginal tentacles. Umbrella 0.5-0.7 mm
containing each a group of about 100 cnidocysts; high, slightly higher than broad, pear-shaped, with
marginal bulbs with blackish pigment granules; apical projection: Hydractinia minuta (Mayer,
cnidocysts along entire tentacle surface and in ter- 1900) (Fig. 3.16)
minal swellings; medusa buds arising from middle
region of manubrium. Family Rathkeidae
Anthomedusae with subglobular umbrella, with slight
Family Hydractiniidae apical process; manubrium short, cylindrical, not
Anthomedusae more or less bell-shaped; with or with- extending beyond umbrella margin; with gastric
out slight apical process; manubrium tubular to sac- peduncle; mouth with 4 lips elongated to form either
shaped, not extending beyond umbrellar margin; with simple or branched oral arms armed with terminal and
or without peduncle; mouth with 4 simple or branched usually also lateral clusters of cnidocysts; 4 to 8 radi-
oral lips elongated into arms armed with terminal al canals and circular canal; gonads completely sur-
clusters of cnidocysts (exceptionally mouth rim sim- rounding manubrium; 8 groups of solid marginal ten-
ple and armed with cnidocyst ring: Kinetocodium); 4, tacles; without ocelli.
8, or more solitary, solid, marginal tentacles; with 4
radial canals and circular canal; gonads on manubri- Genus Rathkea Brandt, 1838
um, interradial, sometimes extending along proximal Rathkeidae with 4 radial canals and 4 oral arms armed
portions of radial canals; with or without ocelli. with clusters of cnidocysts.
Rathkea formosissima (Browne, 1902) (Fig. 3.17).
Genus Hydractinia (van Beneden, 1841) (junior Umbrella with large, solid apical projection; with
synonym: Podocoryna) well developed gastric peduncle; oral arms elon-
Hydractiniidae with 4 or more solid, simple, marginal gated, each with 7-1 1 cnidocyst clusters in double
tentacles, not in groups; usually 4 or 8 simple or row and 1 terminal cluster. Umbrella 2.5 mm wide
slightly branched mouth arms (which are dilations of and 3 mm high, pyriform to bell-shaped, with large
the mouth rim) armed with clusters of cnidocysts; rounded apical projection; manubrium short, quad-
with or without gastric peduncle; gonads on manubri- rangular; perradial bulbs with 3-5 tentacles, inter-
um, interradial, but sometimes extending along prox- radial bulbs with 3 tentacles; gonads interradial in
imal parts of radial canals; with or without ocelli; males, completely covering manubrium in females.
sometimes asexual reproduction by medusa budding Rathkea africana Kramp, 1957 (Fig. 3.18).
on manubrium. Umbrella without apical projection; without gas-
1 Mouth with 4 short lips, each with cluster of cni- tric peduncle, 1.2 mm wide, 1.7 mm high; oral lips
docysts, not prolonged into mouth-arms ..............2 bifurcated, each with 2 broad, terminal clusters of
la Oral arms well developed, simple, undivided; cnidocysts, no lateral clusters; perradial marginal
without ocelli; with gastric peduncle; medusa buds bulbs with 3-4 tentacles of very unequal length,
on interradial side of manubrium .........................3 interradial bulbs with 2-3 tentacles; 8 adradial
2 With 8 marginal tentacles; with gastric peduncle; gonads; medusa buds on gonads.
without ocelli. Umbrella 1.5 mm wide, 2 mm high,
conical with slight constriction below thickened Suborder Tiarida
apex; manubrium cubical: Hydractinia tenuis Medusae with hollow tentacles; ocelli, when present,
(Browne, 1902) (Fig. 3.13) abaxial; without oral tentacles armed with cnidocyst
2a With 4 marginal tentacles; without gastric pedun- clusters, mouth simple, without specialised cnidocyst-
cle; with ocelli. Umbrella 2.5 mm wide and high, armed structures.
412 Hydromedusae
1 Marginal tentacles without basal bulbs or swell- canals bifurcating near point of origin in 8 straight
ings, with a terminal cluster of cnidocysts ...........4 canals joining circular canal (occasionally branch-
la Marginal tentacles with basal bulbs, without ter- ing again); 4 interradial gonads. Umbrella up to
minal cnidocyst clusters or capitations ................2 about 20 mm wide and high, globe-shaped, with
2 With 2 or more simple radial canals; marginal ten- thick walls; manubrium small, barrel-shaped.
tacular bulbs not budding .................................... .3 Bythotiara capensis Pagks, Bouillon and Gili,
2a With 4 radial canals, 2 of them bifurcating; margi- 1991 (Fig. 3.20). With 12 marginal tentacles, all
nal tentacular bulbs developing successively into alike, without secondary tentacles; 4 radial canals
medusa buds: Niobiidae dividing at short but irregular distance into 12 ter-
3 With 4 tentacles in adults: Protiaridae minal branches; 8 adradial gonads. Umbrella 22
3a With 2, 6 or more tentacles in adults: Pandeidae mm wide, 24 mm high, with rounded apex, lateral
4 Marginal tentacles solitary, with terminal swellings mesoglea thick; manubrium quadrangular; margi-
with cnidocysts; no oral tentacles: Calycopsidae nal tentacles leaving circular canal at some dis-
4a Marginal tentacles in groups, without terminal tance above margin, tapering, becoming thread-
swellings; with simple oral tentacles without ter- like before terminal swellings.
minal clusters of cnidocysts: Russelliidae
Genus Calycopsis Fewkes, 1882
Family Calycopsidae Calycopsidae with unbranched radial canals; with
Anthomedusae without apical projection and gastric centripetal canals; gonads transversely folded, often
peduncle; mouth with 4 simple or crenulated lips; forming 8 adradial rows; marginal tentacles of similar
with or without centripetal canals; with simple or structure with cnidocysts only on the terminal knob
folded, adradial or interradial gonads on manubrial and with adnate base.
wall; with 4 or 8 simple or branching radial canals and 1 Umbrella with hnnel-shaped apical depression; 3-
circular canal; with 4, 8, or more hollow marginal ten- 4 centripetal canals in each quadrant; 16-30 mar-
tacles either with highly reduced, or more generally, ginal tentacles. Umbrella up to 40 mm wide, 37
without, basal marginal bulbs, each terminating in mm high, globe-shaped, mesoglea not very thick;
large cnidocyst cluster, their basal portion often manubrium large, broad; mouth with large folded,
adnate to exumbrella; with or without rudimentary or crenulated lips; gonads interradial forming 8 adra-
dwarf tentacles; rarely with abaxial ocelli. dial rows with numerous transverse folds; 16 long
1 With centripetal canals, blind or joining base of marginal tentacles alternating with 16 short ones;
manubrium: Calycopsis cnidocyst knob on adaxial base of each long tenta-
la Without centripetal canals .................................... 2 cle: Calycopsis typa Fewkes, 1882 (Fig. 3.2 1)
2 Gonads smooth, without transverse fold ..............3 l a Umbrella without apical funnel depression .........2
2a Gonads transversely folded ..................................4 2 Gonads in pockets embedded in walls of manubri-
3 With 4 tentacles; gonads on perradial ridges of um, no exterior folds. Umbrella 15-18 mm wide,
manubrium: Pseudotiara 20 mm high, 1 centripetal canal in each quadrant,
3a With 8 to 24 marginal tentacles; gonads interradi- blind or joining base of manubrium; 8- 16 marginal
al: Heterotiara tentacles: Calycopsis borchgrevinki (Browne,
4 Radial canals simple or bifurcated: Bythotiara 1910) (Fig. 3.22)
4a Radial canals branching repeatedly at various le- 2a Gonads in exterior folds ....................................... 3
vels: Sibogita 3 Marginal lobes between marginal tentacles with
conspicuous exumbrellar papillae; 2 adradial cen-
Genus Bythotiara Giinther, 1903 tripetal canals in each quadrant; 8- 12 marginal ten-
Calycopsidae with 4 simple or branching radial tacles, all alike. Umbrella 26 mm wide, 27-33 mm
canals; without centripetal canals; gonads interradial high, mesoglea very thick and rigid: Calycopsis
with transverse furrows, with or without rudimentary papillata Bigelow, 1918 (Fig. 3.23).
or dwarfed tentacles entirely covered with cnidocysts. 3a Marginal lobes without papillae ........................... 4
Bythotiara murrayi Giinther, 1903 (Fig. 3.19). 4 With 1 interradial blind centripetal canal in each
With 8 primary marginal tentacles, some secon- quadrant; with 8 long perradial and interradial ten-
dary tentacles and minute dwarf tentacles; 4 radial tacles and up to 40 smaller marginal tentacles of
Hydromedusae
varying length. Umbrella up to 16 mm wide and Pseudotiara tropica (Bigelow, 1912) (Fig. 3.29).
high, almost spherical, mesoglea thick; 4 radial Umbrella 9 mm wide, 10 mm high, globular, with
canals, connected to manubrium by mesenteries; thick mesoglea; manubrium quadrangular; 4 long
base of tentacles adnate on exumbrella; gonads tentacles with terminal swellings; medusa buds on
interradial, with 8 adradial rows, each with about 16 manubrium. .
transverse folds: Calycopsis bigelowi Vanhoffen,
1911 (Fig. 3.24) Family Niobiidae
4a With numerous centripetal canals, 7 or more in Anthomedusae without gastric peduncle; with 2 sim-
each quadrant (28-56) most of them joining base ple and 2 bifurcating radial canals; without mesente-
of manubrium or upper part of neighbouring ries; with interradial gonads; with marginal tentacular
canals; 16-32 tentacles. Umbrella up to 30 mm bulbs developing into medusa buds.
wide, 40 mm high, mesoglea thick and rigid;
manubrium half as long as subumbrellar cavity; Genus Niobia Mayer, 1900
with fairly short crenulated lips; gonads with 19- With characters of family.
32 folds in each of 8 rows: Calycopsis chuni Niobia dendrotentaculata Mayer, 1900 (Fig.
Vanhoffen, 1911 (Fig. 3.25) 3.30). Umbrella 4 mm wide, watchglass-shaped;
12 marginal tentacles; each tentacle bulb succes-
Genus Heterotiara Maas, 1905 sively developed into small medusa bud.
Calycopsidae with thick walls; with 4 simple radial
canals; without centripetal canals; gonads interradial, Family Pandeidae
without transverse folds; without secondary tentacles; Anthomedusae with or without apical projection;
without ocelli. manubrium quadrate, usually large; with or without
Heterotiara anonyma Maas, 1905 (Fig. 3.26). gastric peduncle; mouth with 4 simple, crenulated, or
With 8 to 12 marginal tentacles, umbrella bell- complexly folded lips; with 4 radial canals (excep-
shaped; 20 mm wide, 22 mm high. tionally 8 as in Octotiara) often broadened, ribbon-
Heterotiara minor Vanhoffen, 1911 (Fig. 3.27). like, or with jagged margin; rarely centripetal canals;
With 16 to 24 marginal tentacles; umbrella globe- with or without mesenteries; gonads either with
shaped; 10 mm wide, 12 mm high. smooth surface or complexly folded, on manubrium
walls in adradial or interradial positions, sometimes
Genus Sibogita Maas, 1905 extending along radial canals; marginal tentacles hol-
Calycopsidae primarily with 4 radial canals that low, mostly with tapering, elongated, conical (almost
branch repeatedly at various levels; without centripe- carrot-shaped) and often laterally compressed bulbs;
tal canals; with transversely folded gonads. with or without rudimentary tentacles, tentaculae, or
Sibogita geometrica Maas, 1905 (Fig. 3.28). marginal warts; with or without abaxial ocelli.
Umbrella 20 mm wide, 38 mm high, globe- 1 Radial canals with long lateral diverticula; margi-
shaped, mesoglea thick; gonads in about 13 trans- nal tentacles numerous with stalked cnidocyst
verse folds in each of the 8 adradial rows; radial knobs along most of their length: Zanclonia
canals with lateral branches of second and third 1a Radial canals without long lateral diverticula; mar-
order at various levels, up to about 40 radial canal ginal tentacles without stalked cnidocyst knobs ..2
branches joining circular canal; tentacles as many 2 With two well developed marginal tentacles:
as-, or half as many as canals, alternating with as Amphinema
many small ones, each flanked by pair of minute, 2a With 4 or more well developed tentacles ............. 3
wart-like protuberances. 3 With marginal cirri, each cirrus located beside
small marginal rudimentary bulb with abaxial
Genus Pseudotiara Bouillon, 1980 ocellus; 4 interradial gonads horseshoe-shaped,
Calycopsidae with 4 marginal tentacles, with very with diverging folds: Cirrhitiara
reduced basal swellings; with 4 small, simple lips; 3a Without marginal cirri.. ........................................4
usually without centripetal canals; with 4 radial 4 Gonads smooth; 4 simple lips: Merga
canals, usually simple; with 8 longitudinal gonads on 4a Gonads reticulate or folded, or both; oral lips more
perradial ridges of manubrium. or less folded or crenulated ..................................5
414 Hydromedusae
5 Gonads reticulate, with isolated interradial pits ...6 warts; manubrium, gonads and lips deep reddish:
5a Gonads reticulate, without isolated interradial pits Amphinema rubra (Kramp, 1957) (Fig. 3.33)
............................................................................. .7
6 Gonads without folds: Pandea Genus Annatiara Russell, 1940
6a Gonads with horizontal adradial folds directed Pandeidae without apical projection; exumbrella with
towards interradii: Neoturris cnidocyst tracks; manubrium short, very broad, cruci-
7 Gonads horseshoe-shaped, with diverging hori- form, with 4 large perradial lobes closely connected
zontal folds, connected by interradial transverse with proximal half or more of 4 radial canals; mouth
bridge ....................................................................8 very broad, cruciform, with folded margin; several hol-
7a Gonads in irregular, mainly vertical folds on entire low marginal tentacles of 2 sizes, regularly alternating.
interradial walls of manubrium, without transverse Annatiara affinis (Hartlaub, 19 13) (Fig. 3.34).
bridge; without mesenteries, but manubrium very Umbrella to 23 mm wide and high, bell-shaped,
broad, cruciform, with perradial lobes closely con- mesoglea uniformly thick; up to 44 large marginal
nected with radial canals: Annatiara tentacles with laterally compressed basal bulbs
8 With well developed mesenteries: Leuckartiara clasping umbrella margin, but without true abaxial
8a Without mesenteries: Halitholus spurs, alternating with as many small tentacles, all
with abaxial ocellus; gonads interradial with se-
Genus Amphinema Haeckel, 1879 veral irregular vertical folds, well developed adra-
Pandeidae usually with considerable apical projec- dially and regularly arranged along sides of perra-
tion; never with more than 2 opposite, hollow, margi- dial lobes.
nal tentacles; with marginal warts or tentaculae; with-
out gastric peduncle; manubrium with broad base; Genus Cirrhitiara Hartlaub, 1913
with or without mesenteries; mouth with 4 simple Pandeidae with large, solid apical projection; with 4
lips; gonads adradial or interradial, occasionally or 8 large hollow marginal tentacles and a number of
extending along radial canals; with or without ocelli. rudimentary marginal bulbs, each carrying a lateral
1 With 8 simple adradial gonads, smooth; with cirrus on one side; all marginal bulbs with ocelli; go-
large, conical, solid, apical projection; with 14-24 nads interradial, horseshoe-shaped, with diverging
small marginal warts. Umbrella up to 4 mm wide folds directed perradially; with long mesenteries.
and 6 mm high, mesoglea of uniform thickness Cirrhitiara superba (Mayer, 1900) (Fig. 3.35).
around top; manubrium cross-like in section, Umbrella 5-7 mm high; manubrium wide; mouth
flask-shaped, almost as long as bell cavity; mouth with 4 recurved, folded lips; 4 broad, large,
with 4 prominent, recurved lips; marginal tenta- smooth edged radial canals entering manubrium
cles with large elongated conical basal bulbs; through 4 wide, funnel-shaped openings; 4 perra-
without tentaculae; without ocelli: Amphinema dial tentacles long, laterally compressed; 12 small
dinema (Ptron and Lesueur, 1810) (Fig. 3.3 1) rudimentary bulbs (occasionally 8 tentacles and
l a With 4 interradial gonads ..................................... 2 rudimentary bulbs).
2 With ocelli, with 4-6 marginal warts. Umbrella 2.5
rnrn wide, 3 mm high, with sharp-pointed apical Genus Halitholus Hartlaub, 1913
projection; manubrium urn-shaped, wide; 4 Pandeidae with large dome-like apical projection;
recurved lips; gonads smooth; tentacles and warts manubrium cubic; gonads more or less horseshoe-
with orange red abaxial ocelli: Amphinema aus- shaped, folded; mouth rim faintly crenulated; radial
tralis (Mayer, 1900) (Fig. 3.32) canals comparatively narrow, not or very faintly
2a Without ocelli, with 5-6 interradial tentaculae. jagged; no mesenteries; 4 or more hollow marginal
Umbrella 4.5 mm wide, 7 mm high, with fairly tentacles; with or without ocelli.
thick walls and fairly pointed apical projection; Halitholus intermedius (Browne, 1902) (Fig. 3.37).
manubrium large, barrel-shaped; mesenteries Umbrella 7 mm wide, 9- 10 mm high; manubrium
long, half length of manubrium; mouth square, 112-213 as long as bell cavity; gonads with an inter-
with 4 simple rccurvcd lips; with broad apical radial horseshoe fold faintly indicated; 4 perradial
chamber above manubrium; marginal tentacles and 4 interradial marginal tentacles; variable num-
with elongate conical bulbs, without marginal ber of minute bulbs, all with conspicuous ocelli.
Hydromedusae 415
Genus Leuckartiara Hartlaub, 1913 cross-shaped in transverse section; mesenteries

Pandeidae usually with apical projection of varying very long; gonads adradial; 4 slightly crenulated
shape; large manubrium connected to radial canals by lips; 8-12 long marginal tentacles and 24-36 rudi-
mesenteries; mouth with extensively folded or crenu- mentary tentacles, all with ocelli.
lated margin; gonads interradial, horseshoe-shaped, Merga tergestina (Neppi and Stiasni, 1911) (Fig.
with folds directed perradially; radial canals broad 3.41). Umbrella with pointed apical projection, 4
and ribbon-like, often with jagged edges; numerous mm wide, 7 mm high, with thin walls; manubrium
hollow tentacles; elongate, laterally compressed basal 112-213 as long as bell cavity; mouth lips faintly
bulbs; often with rudimentary tentacles; with or with- crenulated; gonads adradial, smooth; 4-8 marginal
out ocelli. tentacles with ocelli and few rudimentary bulbs
Leuckartiara octona (Fleming, 1823) (Fig. 3.38). without ocelli.
Marginal tentacles each with pronounced abaxial
spur; with rudimentary peripheral tentacle bulbs. Genus Neoturris Hartlaub, 1913
Umbrella up to 20 mm high, higher than wide, Pandeidae with apical projection strongly variable in
bell-shaped, with generally well-developed coni- shape and size, often reduced; manubrium very large
cal or spherical apical projection, thin lateral and broad, with well-developed mesenteries; gonads
walls; manubrium of varying length, with broad in 8 adradial series with transverse folds directed
base, flask-shaped; gonads typically horseshoe- towards interradii; depressed interradial parts of
shaped on whole surface of manubrium; radial manubrium with isolated pits of gonads; 8 or more
canals with smooth or slightly jagged edges; hollow marginal tentacles with laterally compressed
mesenteries along about half length of manubri- basal bulbs; without rudimentary tentacles or margi-
um; with 12-24, usually 16, long marginal tenta- nal warts; mostly without ocelli.
cles and 16 or more club shaped marginal rudi- Neoturris pileata (Forskal, 1775) (Fig. 3.42).
mentary bulbs, all bulbs with ocelli. Umbrella up to 25 mm wide and 40 rnm high, bell-
Leuckartiara adnata Pages, Gili and Bouillon, shaped, with variable apical projection; mouth
1992 (Fig. 3.39). Marginal tentacles without with very complexly folded and crenulated lips;
spurs; with adnate rudimentary tentacles. mesenteries about half as long as manubrium; ra-
Umbrella 14 mm wide, 15 mm high, bell shaped, dial canals broad with short, sometimes branched
with relatively short conical apical projection; lateral diverticula; up to 90 (usually 60-80) margi-
mouth with short crenulated lips; horse-shoe- nal tentacles, densely crowded, marginal bulbs
shaped interradial gonads, each with two adradial clasping exumbrella but not forming conspicuous
rows presenting seven parallel folds sloping abaxial spurs; without ocelli.
obliquely downwards towards interradii; with
mesenteries about as long as manubrium; radial Genus Pandea Lesson, 1843
canals with sinuous edges, circular canal broad; up Pandeidae with or without apical projection; with or
to 23 marginal tentacles; 3 tentaculae between without longitudinal exumbrellar cnidocyst ribs; go-
each pair of tentacles, completely adnate to exum- nads at first in adradii and eventually encircling manu-
brellar surface, continuing in form of ridges over b r i m , forming complex network; lips wide and fold-
entire exumbrella; probably without ocelli. ed; radial canals ribbon-like; with long mesenteries;
with more than 8 hollow marginal tentacles; without
Genus Merga Hartlaub, 1913 rudimentary marginal tentacles or marginal warts;
Pandeidae with perradial edges of manubrium connect- with or without ocelli.
ed with radial canals by long mesenteries; gonads Pandea conica (Quoy and Gaimard, 1827) (Fig.
smooth; oral lips simple or faintly crenulated; with 4-8 3.43). With longitudinal exumbrellar cnidocyst
or more marginal tentacles and as many or more rudi- ribs; marginal tentacle bulbs laterally compressed.
mentary bulbs or tentaculae; with or without ocelli. Umbrella up to 10 mm wide and 21 (sometimes up
Merga violacea (Agassiz and Mayer, 1899) (Fig. to 30 mm) high, with rounded or conical apex,
3.40). Umbrella dome-like, without apical projec- topped with a peculiar opaque ectodermal thicken-
tion, up to 7 mm wide, 11 mm high, with thick ing; mesoglea fairly thick; with 16-24 longitudinal
walls; manubrium half as long as bell cavity, exumbrellar cnidocyst ribs originating from each
416 Hydromedusae
tentacular bulb; manubrium about half as long as merotrichous isorhizas (a type of cnidocyst restricted to
bell cavity; with short mouth tube and very folded the Leptomedusae families Eirenidae, Eucheilotidae,
lips; radial canals fairly narrow, smooth; mesent- Lovenellidae and Tiaropsidae). Hydroids are known in
eries about 415 of manubrium length; with 16-24 Halitiara injlexa (Bouillon, 1985b), and Halitiara for-
(sometimes up to 44) marginal tentacles, with con- mosa (Brinckrnann-Voss, pers. comm.), and are also
ical, laterally compressed bulbs, but devoid of very different from pandeid polyps, showing some
well-developed abaxial spurs; gonads on entire resemblance to certain thecate Campanulinidae species.
interradial walls of manubrium, forming coarse-
meshed network of ridges with pits in between; Genus Halitiara Fewkes, 1882
with ocelli. Pandeidae with 4 straight radial canals; with 4 perra-
Pandea rubra Bigelow, 1913 (Fig. 3.44). Without dial marginal tentacles and several intermediate solid,
exumbrellar cnidocyst ribs; marginal tentacle cirrus-like tentacles; mouth a simple cruciform open-
bulbs not laterally compressed. Umbrella up to 75 ing; with or without mesenteries; interradial gonads;
mm wide and high, bell-shaped, with rounded without ocelli.
summit, without marked apical process, walls fair- Halitiara formosa Fewkes, 1882 (Fig. 3.36).
ly thick; manubrium broad, 112 bell cavity; Umbrella about 3 rnm high, pear-shaped, with solid
mesenteries about 415 of manubrium length; apical projection, about half as long as bell cavity;
mouth rim cruciform, strongly folded and crenulat- no mesenteries; 4 long, hollow marginal tentacles
ed lips; gonads on entire interradial areas, forming and 24-35 short, solid, cirrus-like tentacles.
very close-meshed irregular and complex network
of ridges with pits in between; up to 25 marginal Family Russelliidae
tentacles, marginal bulbs large, conical, not lateral- Anthomedusae with apical projection; manubrium
ly compressed, clasping margin of exumbrella and mounted upon peduncle; with four unbranched oral
with distinct abaxial spurs; without ocelli; subum- tentacles above mouth margin, without terminal cni-
brella, manubrium, velum and tentacles deep docyst clusters; mouth with four perradial lips; margi-
brownish-red. nal tentacles hollow, without basal swellings, in 8
groups, 4 perradial and 4 adradial, each group with
Genus Zanclonia Hartlaub, 1913 one large and two small tentacles; basal part of large
Pandeidae with 20 long, transverse diverticula at right tentacles sunken into deep furrows of umbrella mar-
angles on both sides of four radial canals; with 24-32 gin; with 8 adradial gonads; with adaxial red ocellus at
marginal tentacles, each with adaxial series of stalked base of free portion of tentacles. Hydroids unknown.
cnidocyst knobs.
Zanclonia weldoni (Browne, 1910) (Fig. 3.45). Genus Russellia Kramp, 1957
Umbrella up to 36 mm high, bell shaped, with With characters of family. One species.
thick walls and round summit; manubrium large Russellia mirabilis Kramp, 1957 (Fig. 3.46). Up
and globular, half as long as bell cavity; mouth to 9 mm wide, 15 mm high.
large with folded lips; gonads in 8 longitudinal
rows of transverse folds. Order Capitata
Medusae with gonads generally completely surround-
Family Protiaridae ing manubrium (with one exception in the South
Anthomedusae with only 4 fully developed marginal Atlantic waters: Zanclea). Mouth simple and circular.
tentacles arising from well developed hollow tentacular Marginal tentacles usually hollow (except in
bulbs; 4 simple radial canals and a circular canal; Margelopsidae and Porpitidae, where they are solid).
mouth with 4 simple lips; Gonads with smooth surface, Cnidome of medusae characterised by presence of
interradial, with or without mesenteries, without rudi- stenoteles. Adult or larval hydroids usually with capi-
mentary bulbs; margin with or without cirrus-like ten- tate tentacles.
tacles, exceptionally with ocelli. Differences between
the diagnoses of the families Protiaridae and Pandeidae Suborder Moerisiida
appear at first rather small, but the cnidome of the Capitata with quadrate manubrium forming radial
Protiaridae is very particular, containing, among others, lobes; mouth cruciform; interradial gonads on manu-
brium and radial lobes, or on radial lobes only, margi- peduncle only; with 4 distinct, crenulated lips;
nal tentacles generally developed along entire circular with 4 stout, tapering perradial tentacles, with
canal; tentacle bulbs usually with abaxial ocelli. Two cnidocysts on proximal part in small rounded
families: Polyorchidae (gonads sausage-like or spi- warts, on middle part forming transverse flaps, on
ral), and Moerisiidae (gonads on manubrium and per- distal parts in rings; with abaxial ocelli.
radial manubrial lobes extending along radial canals).
Suborder Tubulariida
Family Moerisiidae Capitata generally with cylindrical manubrium with cir-
Anthomedusae with prismatic manubrium presenting cular base; mouth usually simple and circular; gonads
radial lobes on proximal parts of the 4 radial canals; normally completely surrounding manubrium; margi-
without gastric peduncle; with simple cruciform nal tentacles developed only at junction between radial
mouth usually with 4 perradial lips; gonads on manu- and circular canals (except in some Eleutheriidae); with
brium and surrounding manubrial lobes; 4 or 16-32 1,4, rarely 8 or 16 marginal tentacles.
moniliform or modified moniliform marginal tenta- 1 Marginal tentacles simple ....................................
2
cles with adnate bulbs; with abaxial ocelli. 1a Marginal tentacles bifurcated: Eleutheriidae
2 Marginal tentacles solitary ...................................
3
Genus Moerisia Boulenger, 1908. With characters of 2a Marginal tentacles in 4 groups: Margelopsidae
family, with moniliform tentacles. 3 Marginal tentacle bulbs with ocelli: Corynidae
Moerisia inkermanica (Paltschikowa-Ostroumova, 3a Marginal tentacle bulbs without ocelli .................4
1925) (Fig. 3.47). Umbrella up to 6.5 mrn wide, 5.5 4 Umbrella without exumbrellar cnidocyst tracks ..5
rnm high, dome shaped, mesoglea very thick; 4a Umbrella with exumbrellar cnidocyst tracks:
manubrium small with radial lobes extending Tubulariidae*
almost nearly to bell margin; gonads on radial lobes 5 With 1-4 marginal tentacles usually unequally
with distal portions sac-like, pendent; up to 32 mar- developed but all of same structure; without apical
ginal tentacles of different lengths, proximal part of projection: Euphysidae*
tentacle narrow, adnate to umbrella; with ocelli; 5a With 1-4 marginal tentacles usually of different
without statocysts. size and structure: Corymorphidae*
* Differences between the Corymorphidae, Euphysidae
Family Polyorchidae and Tubulariidae at the medusa stage are rather slight,
Anthomedusae with gastric peduncle; manubrium separation of these families being based mainly on
prismatic; mouth with 4 oral lips crowded with cni- their hydroid phase.
docysts; with 4 sac-shaped, spiral, or several sausage-
shaped manubrial pouches on peduncle only; gonads Family Corynidae
surrounding manubrial pouches; 4 radial canals with Anthomedusae with bell-shaped umbrella; with sim-
or without blind side branches; 4-260 tentacles with ple circular mouth; 4 radial canals and circular canal;
stout, elongated bulbs; with abaxial ocelli. 2-4 hollow marginal tentacles; gonads completely
encircling manubrium; with abaxial ocelli. Two gene-
Genus Tiaricodon Browne, 1902 ra: Dipurena (gonads divided in one or more rings),
Polyorchidae with 4 radial canals, without diverticu- and Sarsia (gonads undivided).
lae; manubrium with short, sac-like perradial lobes
extending along broad peduncle; mouth with 4 dis- Genus Dipurena McCrady, 1859
tinct frilled lips with cnidocyst band; with 4 perradial, Corynidae with gonads divided into two or more rings
imperfectly moniliform tentacles with stout elongated around manubrium; endoderm of sexual parts diges-
bulbs surrounded by thickened, cnidocyst-studded tive, endoderm of non-sexual parts chordal; manu-
epidermis; gonads on manubrium and perradial lobes; brium usually extending well beyond umbrellar mar-
with abaxial ocelli. gin; with ocelli.
Tiaricodon coeruleus Browne, 1902 (Fig. 3.48). 1 Marginal tentacles with well developed terminal
Umbrella 24 mm wide, 25 mm high, mesoglea knob of cnidocysts and several cnidocyst rings ...2
thick, bell-shaped; manubrium extending almost l a Marginal tentacles with irregular clusters of cni-
to velar opening, with perradial sac-like lobes on docysts, terminal cnidocyst knob inconspicuous or
418 Hydromedusae
absent. Umbrella 3.9 mm wide, 2.5 mm high, bell- Family Corymorphidae

shaped, with globular apical chamber; gonads in 2 Anthomedusae with or without apical process; with-
rings encircling manubrium, one distal, other half out exumbrellar cnidocyst tracks; with simple, circu-
way down manubrium: Dipurena reesi Vannucci, lar mouth; with 1-4 marginal capitate or moniliform
1956 (Fig. 3.49) tentacles; exceptionally with branched or rudimentary
2 Marginal tentacles of uniform diameter, each with tentacles; gonads surrounding manubrium.
large terminal cnidocyst knob and 3 to 6 distinct 1 With 1 marginal tentacle ...................................... 2
rings of cnidocysts just above; middle part of ra- l a With 3 short or rudimentary and 1 long marginal
dial canals with linear swellings. Umbrella up to 6 tentacle of variable structure: Euphysora
mm wide, 8 mm high, bell shaped, with globular 2 Umbrella margin slightly oblique, marginal bulbs
apical chamber; gonads in 2 or more rings, leaving small; marginal tentacle hollow for half its length,
upper third free: Dipurena halterata (Forbes, ending in long and large, oval, ectodermal swell-
1846) (Fig. 3.50) ing bearing numerous cnidocysts: Vannuccia
2a Proximal part of tentacles large, cylindrical, devoid 2a Umbrella margin straight, marginal bulbs large,
of cnidocysts, remaining part becoming abruptly only proximal part of marginal tentacle hollow,
thinner, distal third surrounded by numerous cnido- distal part solid with very thick mesoglea, ending
cyst rings, ending in small terminal cnidocyst knob; in globular cnidocyst knob ...................................3
radial canals without linear swellings. Umbrella 1.8 3 Gonads on manubrium and on 4 sausage-like
mm wide, 3.2 mm high, deeply bell-shaped, apical interradial manubrial pouches: Gotoea
mesoglea thick, with oval or globular apical 3a Manubrium without pouches, gonads simple;
chamber, gonads in 2 rings, one small ring apical, Paragotoea
immediately under and in connection with apical
chamber, other one voluminous on distal part of Genus Euphysora Maas, 1905
manubrium: Dipurena baukalion Pagks, Gili and Corymorphidae usually with 3 short or rudimentary
Bouillon, 1992 (Fig. 3.51) tentacles, and one long tentacle which differs from
others not only in size, but also in structure.
Genus Sarsia Lesson, 1843 1 With only one tentacle, very long and thin, with
Corynidae with undivided gonads. several lateral branches. Umbrella to 26 mm wide
1 Manubrium with distinct conical apical chamber; and high, globular, mesoglea very thick, bell ca-
marginal tentacles with small terminal cnidocyst vity very narrow; manubrium cylindrical or barrel-
knob. Umbrella up to 7 mm wide, 10 mm high, shaped, less than 213 bell cavity, completely encir-
with thick walls; manubrium cylindrical almost as cled by gonads: Euphysora gigantea Kramp,
long as bell cavity; gonads completely surround- 1957 (Fig. 3.55)
ing manubrium: Sarsia producta (Wright, 1858) l a With one long tentacle and 3 small tentacles or
(Fig. 3.52) rudimentary bulbs.. ...............................................2
1a Manubrium without apical chamber; marginal ten- 2 Apical projection almost as long as bell, slender
tacles with distinct terminal cnidocyst knob ........2 and pointed, apical canal as long as apical projec-
2 Mesoglea fairly thin; tentacles long, with large ter- tion. Umbrella up to 5 mm high, including apical
minal knob. Umbrella 3 mm wide, 4 mm high, projection; manubrium about as long as bell ca-
cylindrical, walls moderately thick; manubrium vity; gonads encircling whole length of manubri-
213 of bell cavity, nearly all length surrounded by um; main tentacle very long, moniliform, with
gonads: Sarsia gracilis Browne, 1902 (Fig. 3.53) prominent swellings at irregular intervals and with
2a Mesoglea thick; tentacles fairly short, terminal distinct terminal knob; opposite tentacle short,
knob rather small. Umbrella 3-4 mm high, a little cone-shaped, 2 others reduced to bulbs:
higher than wide, bell-shaped; manubrium cylin- Euphysora gracilis (Brooks, 1882) (Fig. 3.56)
drical, about as long as bell cavity, entirely sur- 2a Apical projection short and broad, no apical canal.
rounded by gonads, eggs few and large: Sarsia Umbrella up to 6.5 mm wide, 8 mm high, bell-
eximia (Allman, 1859) (Fig. 3.54) shaped with pointed apex and fairly thin walls;
manubrium barrel shaped, 213 of bell cavity; radi-
al canals thick, with large vacuolated endodermal
Hydromedusae 419
cells; gonads encircling manubrium, with perradi- Vannuccia forbesii (Mayer,1894) (Fig. 3.60).
a1 longitudinal depressions indicating gonad divi- With characters of genus. Umbrella 3 mm high,
sion (after Schuchert, 1996), or in 4 perradial ellipsoidal, mesoglea evenly thin.
masses separated in interradii (re-examination of
South African material shows perradial gonads); Family Eleutheriidae
main marginal tentacle long, very contractile, its Anthomedusae usually with thickened continuous or
terminal end bifurcated twice in 4 knobs covered broken ring of cnidocysts around umbrella margin;
with cnidocysts; opposite this a fairly long filiform with circular mouth, simple or armed with cnidocyst
tentacle; two short, conical lateral tentacles: knobs; number of radial canals variable, which may or
Euphysora furcata Kramp, 1948 (Fig. 3.57) not branch; gonads on manubrium, on subumbrellar
surface, or in specialised dorsal brood pouches; mar-
Genus Gotoea Uchida, 1927 ginal tentacles hollow, variable in number, bifurcating
Gonads surrounding manubrium and on 4 sausage- in upper branch, armed with cnidocyst clusters or
like pendent pouches of interradial manubrial walls; knobs, and lower unarmed branch terminating in
with 1 tentacle with large terminal knob of cnidocysts; adhesive pad; with abaxial ocelli.
with swollen exumbrellar cnidocyst pad at level of
marginal bulbs. Genus Staurocladia Hartlaub, 1917
Gotoea similis Kramp, 1959 (Fig. 3.58). Umbrella Eleutheriidae adapted for crawling and walking; with-
3 mm wide, 3,5 mm high, cylindrical with flat top out brood pouch above manubrium; gonads around
and thin walls; manubrium flask-shaped with 4 manubrium or developed in interradial ectodermal
interradial pendent pouches, extending slightly manubrial pockets; with 6-1 1 simple radial canals;
beyond velar opening; tentacle very long and of mouth circular or with 5-6 cnidocyst knobs; with up to
same structure as in Paragotoea bathybia; margi- 60 marginal tentacles, dichotomous, upper branch with
nal bulbs of unequal size, the one opposite tenta- several cnidocyst clusters, lower with adhesive organ;
cle much larger. asexual reproduction by medusa budding or by fission.
1 Upper marginal tentacular branch without lateral
Genus Paragotoea Kramp, 1942 (after Kramp, 1961, cnidocyst clusters .................................................2
not Ralph, 1959) l a Upper marginal tentacular branches with lateral
Corymorphidae with 4 radial canals, without gastric cnidocysts clusters. Umbrella 1.5-2 mm wide; 6-1 1
pouches; with circular mouth; with 1 well-developed radial canals; 20-32 marginal tentacles with upper
solid tentacle terminating in large knob of cnidocysts branch as long as lower; with broken and reduced
and 3 very large marginal bulbs without tentacles. marginal cnidocyst ring; with 5-6 lateral cnidocyst
Paragotoea bathybia Kramp, 1942 (Fig. 3.59). clusters on upper branch of marginal tentacles:
Umbrella up to 1.6 mm wide, 2 mm high, with flat- Staurocladia hodgsoni (Browne, 1910) (Fig. 3.6 1)
tened top and thin walls, with conical apex; nume- 2 Upper marginal tentacular branch up to 3 times
rous cnidocysts on exumbrella; manubriurn cylindri- longer than lower branch. Umbrella up to 3.3 mm
cal, proximal part vacuolated, distal part sexual; 4 wide, 6 radial canals; with continuous marginal
large marginal bulbs each with ectodermal abaxial cnidocyst ring; up to 40 marginal tentacles, upper
spur; 1 stiff tentacle, proximal part hollow, distal branch with 2 (rarely 3) cnidocyst clusters on abo-
part solid with very thick mesoglea, with large ter- ral tentacular side, 1 on oral side, and 1 terminal
minal knob of cnidocysts; gonads first interradial, cluster; medusa buds on umbrella margin of small
eventually encircling manubrium with age. specimens: Staurocladia capensis (Gilchrist,
1918) (Fig. 3.62)
Genus Vannuccia Brinckmann-Voss, 1967 2a Upper marginal tentacular branch shorter than
Corymorphidae with slightly asymmetrical umbrella lower branch. Umbrella 3 mm wide, 2 mm high;
margin, with small marginal bulbs; with 1 marginal usually 8 (6-1 1) radial canals; with continuous mar-
tentacle, hollow for half its length, ending in long, ginal cnidocyst ring; up to 30 marginal tentacles,
large, oval cnidocyst swelling; manubrium spindle- upper branch with 2-3 cnidocyst clusters on aboral
shaped, swollen, extending to velum, gonads encir- tentacular side, 1-2 on oral side, and 1 terminal
cling manubrium. cluster; medusa budding at level of marginal ring:
Staurocladia vallentini (Browne, 1902) (Fig. 3.63)
420 Hydromedusae
Family Euphysidae liform tentacles without bulbs; gonads on basal

Anthomedusae with evenly rounded or dome-shaped part of manubrium.
umbrella; without apical canal; without exumbrellar
cnidocyst tracks; 1-4 marginal tentacles unequally Family Tubulariidae
developed but all of same structure, moniliform or Anthomedusae usually with exumbrellar cnidocyst
modified moniliform; manubrium cylindrical, stout; tracks; with 4 radial canals; usually with circular
mouth simple, circular; gonads encircling manubrium. mouth; with gonads completely encircling manu-
brium; with 1-4 marginal tentacles; without ocelli.
Genus Euphysa Forbes, 1848
With characters of family. Genus Ectopleura L. Agassiz, 1862
Euphysa aurata Forbes, 1848 (Fig. 3.64). Tubulariidae with symmetrical umbrella; with 2
Umbrella about 4 rnm high, higher than wide, opposite or 4 simple tentacles, moniliform or with
mesoglea thick, especially in apical region; with abaxial cnidocysts clusters; with 4 radial canals; with
horizontal umbrella margin; manubrium shorter 8 longitudinal exumbrellar cnidocyst rows, in pairs,
than bell cavity; mouth surrounded by cnidocysts; from tentacle bulbs.
gonad encircling almost all manubrium, leaving Ectopleura dumortieri (van Beneden, 1844) (Fig.
upper end of manubrium and mouth free; 1 moni- 3.68). Umbrella 2-3 mm high, bell nearly spheri-
liform tentacle, 3 non-tentacular marginal bulbs cal, mesoglea very thick, especially in apical
smaller than tentacular bulb and with slight spurs. region; apical canal sometimes present; mouth rim
Euphysa monotentaculata Zamponi, 1983 (Fig. with cnidocysts; 4 tentacles with large basal bulbs
3.65). Doubtful species. and prominent round abaxial cnidocyst clusters.
Family Margelopsidae Genus Hybocodon L. Agassiz, 1862

Anthomedusae without apical projection; without cni- Tubulariidae with umbrella margin at oblique angle;
docyst tracks; with simple circular mouth; with gonads without pointed apical process; with or without exum-
surrounding manubrium; 4 radial canals; with solid, brellar cnidocyst tracks; 4 radial canals, 1 short, 2
generally moniliform tentacles in marginal clusters, or medium sized, and one longer; manubrium on short
at different levels on exumbrella; without ocelli. Two peduncle; with 1 simple or compound marginal bulb
genera: Margelopsis (with 4 marginal perradial clusters with 1-3 tentacles; with the 3 remaining perradial
of tentacles), and Octobulbacea (with 8 marginal clus- bulbs rudimentary.
ters of tentacles, 4 perradial and 4 interradial). Hybocodon unicus (Browne, 1902) (Fig. 3.69).
Exumbrella with scattered cnidocysts, no meridian
Genus Margelopsis Hartlaub, 1897 cnidocysts tracks; one moniliform solitary tenta-
Margelopsideae with 4 marginal bulbs, each with cle. Umbrella 2 mm wide, 3 mm high, bell shaped;
cluster of 2 tentacules. umbrella margin slightly oblique, manubrium
Margelopsis australis Browne, 1910 (Fig. 3.66). almost as long as bell cavity, mounted on short
Umbrella 1-3 mm wide and high, almost globular; peduncle; gonads on almost whole length of
manubrium cylindrical, about as long as bell cav- manubrium; tentacle on very small bulb placed
ity; gonad a globular swelling in middle portion of between 2 large swellings of cnidocysts on bell
manubrium; with 4 small marginal bulbs each margin; 3 clasping rudimentary bulbs.
with 2 small (moniliform?) tentacles placed one Hybocodon prolifer L. Agassiz, 1862 (Fig. 3.70).
behind the other. Exumbrella with 5 meridian cnidocysts tracks; 1
tentacular bulb with 1 or more moniliform tenta-
Genus Octobulbacea Zamponi, 1983 cles and with medusa buds. Umbrella 3 mm wide,
Margelopsidae with 8 marginal bulbs, 4 perradial and 4 mm high, bell-shaped, umbrella margin oblique;
4 interradial, each with cluster of two tentacles. 2 meridian cnidocyst tracks issuing from tentacu-
Octobulbacea montehermosensis Zamponi, 1983 lar bulb, 1 track of each 3 reduced non-tentacular
(Fig. 3.67). Umbrella 3 mm high, cylindrical; bulb; manubrium large, cylindrical, mounted on
manubrium short; with 4 radial canals; with 16 short peduncle; gonads completely surrounding
marginal moniliform tentacles issued from per- manubrium; eggs developing in actinulae.
radial and interradial bulbs. and 8 adradial moni-
Hydromedusae 42 1
Genus Rhabdoon Keferstein and Ehlers, 1861 lar mouth, with or without oral tentacles; with 4 radial
Tubulariidae with single hollow marginal tentacle canals, exceptionally bifurcated; marginal tentacles
ending in large cluster of cnidocysts, without marginal when present, 2 or 4, hollow, each with numerous
bulb; with 4 perradial longitudinal exumbrellar cnido- abaxial cnidophores (Fig. 24); gonads usually interra-
cyst tracks, 4 additional interradial and sometimes dial, rarely in single mass around manubrium; without
adradial ones; with vacuolated cells containing refrac- ocelli.
tive droplets along 4 radial canals, at umbrella apex
1
and bell margin; without non-tentacular marginal Genus Zanclea Gegenbaur, 1857
I bulbs; manubrium occupying almost entire bell cavity; Zancleidae with exumbrellar perradial cnidocyst clus-
/i gonads surrounding middle part of manubrium.
Rhabdoon singulare Keferstein and Ehlers, 1861
ters; mouth simple, circular; with 4 simple radial
canals; marginal tentacles when present, 2 or 4, with
(Fig. 3.7 1). Umbrella 1.6 mm wide, 2.1 mm high, numerous abaxial cnidophores; gonads interradial;
i
barrel-shaped, apex dome-shaped, with very thin cnidome generally stenoteles and macrobasic eury-
apex mesoglea, umbrella margin thick; mouth teles, without desmonemes.
simple, circular. Zanclea sp. (Fig. 24, 3.74). Characteristics as
above. Umbrella up to 3 mm high. Medusae of the
Suborder Zancleida several Zanclea species are almost unidentifiable
Capitata with flask-shaped manubrium, with quadrate without detailed study of the structure of their
base and cylindrical mouth tube; gonads interradial; macrobasic euryteles, and a general knowledge of
with or without exumbrellar cnidocyst pouches; their complete life cycle.
marginal tentacles developed only at junction between
radial and circular canals; 2-4 marginal tentacles with Family Zancleopsidae
or without cnidophores or capitate side branches; with Anthomedusae with conical or dome-shaped umbrella,
or without ocelli. without apical chamber; 2-4 capitate marginal tentacles,
1 With ocelli, and marginal tentacles with capitate with or without lateral capitate branches; marginal
side branches: Zancleopsidae bulbs clasping umbrella margin, with adaxial
la Without ocelli .......................................................
2 hemispherical projection studded with cnidocysts;
2 Marginal tentacles terminating in single large manubrium broadly flask-shaped, with quadrate or
spherical cnidocyst knob: Porpitidae cruciform base; mouth square, with or without faint
2a Marginal tentacles with numerous stalked capsules lips; 4 interradial gonads with deep interradial grooves
containing cnidocysts, stalk of capsules threadlike, which may divide them into 8 adradial patches; with or
very extensile (cnidophores, Fig. 24): Zancleidae without ocelli on marginal bulbs or on proximal part of
tentacles.
Family Porpitidae
Genus Zancleopsis Hartlaub, 1907
Genus Porpita Lamarck, 180 1 Zancleopsidae with or without apical projection;
Porpitaporpita (Linnaeus, 1758) (Fig. 3.72). Only either with 2 long, opposed, capitate tentacles with
floating polyp-bearing colonies are known from capitate side branches and 2 opposed, shorter or lon-
South Atlantic, medusae never recorded in this ger, simple capitate tentacles, or with four simple cap-
I
ocean. itate tentacles; marginal tentacle bulbs clasping
umbrella margin, with large hemispherical adaxial
t Genus Vellela Lamarck, 180 1 expansion covered with cnidocysts; manubrium flask-
Vellela vellela (Linnaeus, 1758) (Fig. 3.73). Only shaped; mouth more or less cruciform, with or with-
floating polyp-bearing colonies are known from out simple lips; gonads interradial, with deep interra-
F
1 South Atlantic, medusae never recorded in this dial grooves which may divide them into 8 adradial
ocean. masses; with ocelli.
Zancleopsis dichotoma Harllaub, 1907.(Fig.3.75).
Family Zancleidae Umbrella 2.5 mm wide, 3 mm high, manubrium
Anthomedusae with bell-shaped umbrella; with or flask-shaped, cross-shaped, mouth with 4 small
without exumbrellar oval or clavate patches or elon- lips covered with cnidocysts; in adults 4 capitate
gate tracts of cnidocysts (Fig. 24); with simple circu- tentacles of equal length, 2 opposed with capitate
422 Hydromedusae
side branches and 2 opposed simple; ocelli abaxial 13 Endodermal core of tentacles extending inwards
on marginal bulbs. from bell margin into bell mesoglea:
Blackfordiidae
Subclass Leptomedusae 13a No endodermal tentacular expansions:
Campanulariidae (see under Proboscoida)
Order Conica (Fig. 1F)
Hydranths with conical hypostome, medusae variable. Family Aequoreidae
1 Without statocysts or cordyli; with numerous Leptomedusae with very wide, circular manubrium;
adaxial ocelli; numerous filiform, solid tentaculi- usually without gastric peduncle; with many simple or
form structures without marginal bulbs and not in branched radial canals; gonads on radial canal separa-
connection with circular canal: Orchistomidae ted from manubrium; hollow marginal tentacles; usu-
l a With marginal cordyli or statocysts, with or with- ally with excretory pores or papillae; without marginal
out ocelli; without tentaculiform structures not in or lateral cirri; with closed statocysts; without ocelli.
connection with circular canal ............................2 1 Radial canals branched or bifurcated : Zygocanna
2 With cordyli ........................................................3 1a Radial canals simple, undivided ...........................2
2a With statocysts .................................................... 4 2 Subumbrella with radial rows of gelatinous papil-
3 Manubrium with 4 perradial lobes connected with lae: Rhacostoma
subumbrella; gonads on manubrium and exten- 2a Subumbrella without rows of gelatinous papillae:
ding on perradial lobes: Tiarannidae Aequorea
3a Manubrium without perradial lobes; gonads on
radial canals usually contiguous with manubrium: Genus Aequorea Peron and Lesueur, 1810
Laodiceidae Aequoreidae with numerous simple radial canals;
4 With open statocysts ........................................... 5 subumbrella without rows of gelatinous papillae.
4a With closed statocysts .........................................6 1 Gonads no more than half as long as radial canals.
5 Open statocyts associated with ocelli: Tiaropsidae Umbrella 9 mm wide, 10-12 mm high, conical,
5a Open statocysts without ocelli: Mitrocomidae with very thick mesoglea; manubrium half as wide
6 With distinct gastric peduncle; with 8 or many as umbrella diameter, often broad and flat; mouth
statocysts: Eirenidae with long and slender lips with inward furrow
6a Without distinct gastric peduncle ........................7 which continues along inside of manubrium to
7 Manubrium very broad; with many radial canals; radial canals; about 16 radial canals, twice as many
tentacle bulbs with excretory pores on, or not on, tentacles (20-30), and as many small bulbs, both
excretory papillae: Aequoreidae without excretory papillae or pores; about twice as
7a Manubrium narrow; normally with 4-8 radial many statocysts as tentacles; gonads laterally com-
canals ................................................................... 8 pressed, in proximal half of radial canals:
8 Tentacle bulbs with excretory pores, 4-8 (some- Aequorea conica Browne, 1905 (Fig. 3.76)
times 11) radial canals: Malagazziidae l a Gonads along almost whole length of radial canals
8a Tentacle bulbs without excretory pores ..............9 ..............................................................................2
9 Tentacle bulbs with lateral cirri ........................10 2 About 100 radial canals, 3-6 times as many tenta-
9a Tentacle bulbs without lateral cirri ...................1 1 cles; tentacle bulbs laterally compressed. Umbrella
10 With 8 statocysts (exceptionally 12-14): up to 145 mm wide, usually 60-80 mm, low and
Eucheilotidae thick manubrium, about half as wide as umbrella;
lOa With numerous statocysts (16 or more): gonads extending along almost whole length of
Lovenellidae radial canals; numerous small bulbs; all bulbs with
11 Exumbrella with marginal cirri: Cirrholoveniidae excretory papillae; statocysts numerous, crowded:
1la Exumbrella without marginal cirri ....................12 Aequorea coerulescens (Brandt, 1838) (Fig. 3.77)
12 Gonads divided into two lateral parts separated by 2a Fewer (usually much fewer) than 3 times as many
a median groove; 8 marginal statocysts: tentacles as radial canals ......................................3
Phialellidae 3 About 4-10 or more times as many radial canals as
12a Gonads completely surrounding radial canals; 8 tentacles ................................................................4
or more statocysts ..............................................13 3a With at least half as many tentacles as radial canals.
Hydromedusae 423
Umbrella large, up to 175 rnm wide, saucer-shaped, canals leave periphery of manubrium without
thick in centre, gradually thinning towards margin; branching, but from a cruciform figure in centre of
manubrium half as wide as umbrella; radial canals manubrium the branching ridges pass to point of
usually 60-80, sometimes fewer or up to 160; go- origin of free canals; gonads along greatest part of
nads along almost whole length of radial canals; radial canals; 28-70 tentacles and several small
tentacles with elongate, conical bulbs, generally rudimentary bulbs, with long excretory papillae;
less numerous than radial canals but varying from statocysts very numerous.
half to twice as many; small bulbs few, scattered;
excretory pores on short papillae; 5-10 statocyst Family Blackfordiidae
between successive radial canals: Aequorea forsk- Leptomedusae with narrow, short manubrium; mouth
alea PCron and Lesueur, 1810 (Fig. 3.78) with 4 long, fluted lips; without gastric peduncle; with
4 Tentacle bulbs broad, each with abaxial keel and 80- 100 hollow tentacles; endoderm of tentacles
prominent excretory papillae. Umbrella up to 75 extends inwards from bell margin into bell mesoglea;
mm wide, lens-shaped, central disk thick, margin with 4 radial canals; gonads completely surrounding
thin; manubrium about half as wide as umbrella; radial canals; without permanent rudimentary tenta-
60- 100 (up to 150) radial canals; gonads extending cles; without marginal and lateral cirri; numerous
along almost entire length of radial canals; 10-30 closed statocysts.
(rarely up to 40) tentacles, 6-8 times as many small
rudimentary bulbs; statocysts very numerous: Genus Blackfordia Mayer, 1910
Aequorea macrodactyla (Brandt, 1834) (Fig. 3.79) With characters of family.
4a Tentacle bulbs with long lateral extensions, with- Blackfordia virginica Mayer, 1910 (Fig. 3 23).
out excretory papillae. Umbrella up to 100 mm Umbrella 14 mm wide, higher than hemispherical,
wide, more or less biconvex, with thin margin; with rounded apex; with linear gonads extending
manubrium 112-213 as wide as umbrella; gonads from corners of manubrium to somewhat more
extending along almost entire length of manubri- than half length of radial canals; tentacles with fin-
um; 150-250 radial canals; 10-16 tentacles and as ger-shaped or oval diverticula into bell margin; 1
many small rudimentary bulbs, no excretory papil- (rarely 2) statocysts between successive tentacles.
lae but excretory pores present as slits; statocysts
very numerous: Aequorea pensilis (Eschscholtz, Family Cirrholoveniidae
1829) (Fig. 3.80) Manubrium small; without peduncle; without excreto-
ry pores; 4 simple radial canals; gonads on radial
Genus Rhacostoma L. Agassiz, 1850 canals separated from manubrium; hollow marginal
Aequoreidae with numerous simple radial canals; tentacles; with marginal cirri; without lateral cirri;
subumbrella with radial rows of gelatinous papillae. with 4 or more closed statocysts; without ocelli.
Rhacostoma atlantica L. Agassiz, 1850 (Fig.
3.81). Umbrella up to 300-400 mm wide, 3-4 Genus Cirrholovenia Kramp, 1959
times as wide as high, mesoglea thick in centre; Cirrholoveniidae with 4-40 marginal tentacles; 7-8
manubrium 113-112 as wide as umbrella; 80-100 marginal cirri between successive marginal tentacles.
radial canals; gonads along greatest part of radial Cirrholovenia tetranema Kramp, 1959 (Fig. 3.84).
canals, leaving both ends free; marginal tentacles Umbrella up to 1.5 mm wide and high, mesoglea
slightly more numerous than radial canals, with thin; manubrium small, cruciform; mouth with
elongate, conical bulbs; with excretory papillae; very short simple lips; gonads thick, cylindrical
numerous statocysts. along almost entire length of radial canals; 4 long
perradial marginal tentacles, with broad bulbs; no
Genus Zygocanna Haeckel, 1879 rudimentary bulbs; 7-8 marginal cirri in each quad-
Aequoreidae with numerous radial canals, branched rant; 4 interradial or 8 adradial statocysts.
or bifurcated; subumbrella sometimes with radial
rows of gelatinous papillae. Family Eirenidae
Zygocanna vagans Bigelow, 1912 (Fig. 3.82). Leptomedusae with small manubrium, usually on
Umbrella up to 76 mm wide, flat, thin; manubrium rather well differentiated gastric peduncle; 4-6 simple
113-112 as wide as umbrella; about 30-45 radial radial canals running from circular canal across
Hydromedusae
underside of bell and along peduncle to manubrium; high; with long, slender, tapering gastric peduncle,
with or without excretory pores; with hollow tenta- twice as long as bell diameter; manubrium small,
cles; with or without cirri or marginal warts; gonads flask-shaped; mouth with 4 simple recurved lips; 4
on radial canals separated from manubrium, in each long tentacles with or without lateral cirri; about
species on well defined part(s) of radial canal; 8 to 100 marginal warts, some with lateral cirri; 8 sta-
many statocysts; without ocelli. tocysts: Eutima mira McCrady, 1859 (Fig. 3.86)
1 With more than 8 statocysts .................................2 l a With 4 gonads restricted to peduncle ...................2
la Usually with 8 statocysts (rarely 12); without mar- 2 ' With 2 or 4 marginal tentacles. Umbrella up to 13
ginal cirri ..............................................................
3 mm wide, almost hemispherical, mesoglea thick;
2 Without cirri: Eirene with very long, narrow gastric peduncle extending i
2a With cirri ..............................................................
4 far beyond bell margin and with small conical
3 Without lateral cirri and marginal warts: Eutonina base; manubrium short; mouth with 4 small, sim-
3a With lateral cirri on marginal warts, usually also ple lips; 2-4 (sometimes more) long perradial ten- i
on base of marginal tentacles; without excretory tacles and 40-80 or more marginal warts usually I
pores: Eutima all with lateral cirri; 8 statocysts: Eutima gracilis ,1
4 With lateral cirri; with excretory pores ................5 (Forbes and Goodsir, 1851) (Fig. 3.87)
4a With marginal cirri: Phialopsis 2a With 32 marginal tentacles. Umbrella 10 mm wide,
5 With gonads restricted to subumbrellar parts of a little broader than high, mesoglea thick at apex,
radial canals: Helgicirrha peduncle 112 of umbrella diameter, slender, tape-
5a With gonads on entire length of radial canals: ring; manubrium short; lips slightly fimbriated; 4
Irenium gonads extending from near manubrium along
peduncle and somewhat out on subumbrella; mar-
Genus Eirene Eschscholtz, 1829 ginal tentacles with lateral cirri; about 96 marginal
Eirenidae with distinct gastric peduncle; without mar- warts with lateral cirri; 8 statocysts: Eutima coeru-
ginal or lateral cirri or marginal swellings; with or lea (L. Agassiz, 1862) (Fig. 3.88)
without excretory pores; 4-6 simple radial canals;
gonads on subumbrellar part of radial canals, not Genus Eutonina Hartlaub, 1897
extending onto gastric peduncle; numerous statocysts. Eirenidae with 8 statocysts; without cirri; without
Eirene viridula (Ptron and Lesueur, 1810) (Fig. marginal warts; gonads restricted to subumbrella.
3.85). Umbrella 20-30 mm wide; hemispherical, Eutonina scintillans (Bigelow, 1919) (Fig. 3.89).
middle region fairly thick; with slender gastric Umbrella 10 mm wide, 5 mm high, mesoglea thick;
peduncle with pyramidal base; manubrium rather with short peduncle; manubrium globular; mouth
small but with 4 long pointed lips, with crenulated with 4 simple or crenulated lips; gonads along 114-
margins; with four radial canals; gonads linear, 113 of subumbrellar parts of radial canals.
sometimes slightly sinuous, extending from some-
what beyond base of peduncle to almost bell mar- Genus Helgicirrha Hartlaub, 1909
gin; about 70 tentacles of different sizes, large and Eirenidae with distinct gastric peduncle; with lateral
small frequently alternating; bulbs conical with cirri at base of some or all marginal tentacle bulbs;
distinct adaxial excretory papillae; 50 or more with or without excretory pores; gonads restricted to
statocysts. subumbrellar portion of radial canals; with numerous
statocysts.
Genus Eutima McCrady, 1859 Helgicirrha schulzei Hartlaub, 1909 (Fig. 3.90).
Eirenidae with distinct gastric peduncle; with lateral Umbrella 30-40 mm wide, flatter than hemisphe-
cirri; with marginal swellings; without excretory rical, mesoglea fairly thin; with narrow, elongate,
pores; 4 simple radial canals; gonads on radial canals, conical gastric peduncle extending well beyond
either beneath subumbrella or on gastric peduncle, or bell margin; manubrium small; mouth with 4 sim-
on both; with 8 (exceptionally 12) statocysts. ple, very short upwardly curved lips and folded
1 With 8 gonads, 4 on subumbrella, extending from margins; gonads linear from base of peduncle to
base of peduncle outwards, 4 on greater part of almost bell margin; 30-40 marginal tentacles, with
peduncle. Umbrella up to 30 mm wide, as broad as or without lateral cirri; up to over 100 small tenta-
Hydromedusae 425
cles or rudimentary bulbs each with pair of lateral more rudimentary bulbs all with lateral cirri; go-
cirri; with adaxial excretory pores on all bulbs; 50 nads along middle portion of radial canals; medusa
or more statocysts. buds on gonads; frustules on radial canals:
Eucheilota paradoxica Mayer, 1900 (Fig. 3.93)
Genus Irenium Haeckel, 1879 2a With 12 marginal statocysts. Umbrella 2.5 mm
Eirenidae (?) with numerous statocysts; with nume- wide, higher than hemispherical; manubrium very
rous marginal warts; marginal tentacles and warts short; 4 marginal tentacles with one pair of lateral
with lateral cirri; gonads along entire radial canals. cirri, no rudimentary bulbs; gonads along distal
Irenium teuscheri (Haeckel, 1879) (Fig. 3.91). end of radial canals: Eucheilota duodecimalis A.
Umbrella 40 mm wide, 20 mm high, thick apex; Agassiz, 1862 (Fig. 3.94)
peduncle as long as bell diameter, pyramidal, 3 Manubrium with 4 interradial black spots; 16-30
about 3 times as long as broad; manubrium fairly marginal tentacles. Umbrella up to 13 mm wide
small; mouth with large folded lips; gonads wavy; and 10 mm high; mouth with 4 well-developed
8 large and 40 small marginal tentacles; 60-80 lips; gonads linear on distal 213 of radial canals;
marginal warts and between them 60-80 spiral with 16-20 (up to 30) long marginal tentacles,
cirri; 70-80-statocysts. This species has never each with 1 pair of lateral spiral cirri; 1-3 rudimen-
been recorded since its description by Haeckel. tary bulbs with lateral cirri between successive
tentacles; 8 statocysts: Eucheilota maculata
Genus Phialopsis Torrey, 1909 Hartlaub, 1894 (Fig. 3.95)
Eirenidae with short gastric peduncle; with marginal 3a Manubrium without black spots; 16 marginal ten-
cirri; without excretory pores; with gonads restricted tacles ....................................................................
.4
to subumbrellar portion of radial canals; with nume- 4 With 16 marginal tentacles with normal bulbs, 16
rous statocysts. rudimentary bulbs all with one pair of lateral cirri,
Phialopsis diegensis Torrey, 1909 (Fig. 3.92). and about 24 minute knobs without cirri. Umbrella
Umbrella 20-30 mm wide, 3-4 times as wide as 10 mm wide, hemispherical; manubrium short;
high; mesoglea thin; with very short, conical mouth with 4 prominent lips; gonads linear, along
peduncle reaching about limit of bell margin; middle third of radial canals; with 8 marginal sta-
manubrium very short; mouth with very short tocysts: Eucheilota ventricularis McCrady, 1859
crenulated lips; gonads linear from basal part of (Fig. 3.96)
peduncle almost to bell margin; 16-28 (sometimes 4a With 16 marginal tentacles with very large and
more) tentacles with elongate conical bulbs; elongated bulbs, each with 1 or 2 pairs of lateral
between successive tentacles, 3-9 triangular rudi- cirri, 16 rudimentary bulbs and 32 marginal swell-
mentary bulbs, 3-9 scattered marginal cirri, and 2- ings all without cirri. Umbrella 2.6 wide, 3.5 high,
5 statocysts; no lateral cirri. mesoglea thin; manubrium 113 of bell cavity;
mouth with 4 lips; gonads on middle third of ra-
Family Eucheilotidae dial canals; statocysts not known: Eucheilota
Leptomedusae with short manubrium; without gastric foresti Goy, 1979 (Fig. 3.97)
peduncle; without excretory pores; with 4 straight
radial canal; marginal tentacles hollow; with lateral Family Laodiceidae
cirri, never marginal cirri; gonads on radial canals, Leptomedusae with marginal cordyli (Fig. 2B, C),
separated from manubrium; usually 8 (exceptionally 4 with or without cnidocysts; 4, 8, or more simple or
or 12) statocysts; without ocelli. branched radial canals; marginal tentacles hollow;
with or without marginal cirri; with or without adax-
Genus Eucheilota McCrady, 1859 ial ocelli; without statocysts.
With characters of family. 1 With 6 or more radial canals, some or all branched,
1 With 4 marginal tentacles ....................................
2 all branches joining circular canal: Toxorchis
1a With 16 or more marginal tentacles.....................3 l a With 4 radial canals ..............................................
2
2 With 8 marginal statocysts. Umbrella 4 mm wide, 2 Radial canals open grooves forming large cruci-
higher than hemispherical; manubrium small, form mouth: Staurophora
flask-shaped; with 4 marginal tentacles and 4 or 2a Radial canals closed .............................................3
426 Hydrcomedusae
3 With simple radial canals; usually some or all ten- ral cirri (often lost upon preservation) and one cor-
tacle bulbs with adaxial ocelli: Laodicea dylus between successive tentacles: Laodicea
3a Radial canals with lateral diverticulae; no ocelli: undulata (Forbes and Goodsir, 1851) (Fig. 3.10 1)
Ptychogena
Genus Ptjxhogena A. Agassiz, 1865
Genus Laodicea Lesson, 1843 Laodiceidae with four radial canals giving rise to la-
Laodiceidae with small manubrium; four simple ra- teral diverticula, in which the gonads are located;
dial canals; with simple wavy gonads; with or without manubrium with funnel-shaped perradial lobes; with-
marginal cirri; with or without adaxial ocelli. out cirri; without ocelli.
1 Manubrium with large perradial lobes; about 50 Ptychogena antarctica Browne, 1907 (Fig. 3.102).
marginal tentacles without basal spurs; 3-4 cordy- Umbrella 60-100 mm wide, slightly convex, meso-
li between tentacles. Umbrella 25 mm wide, 15 glea very thick; manubrium wide, with funnel-
mm high; manubrium large, with 4 large perradial shaped perradial lobes almost to terminal portions
lobes extending more than half-way to bell mar- of gonads; gonads along nearly entire length of
gin; mouth with 4 large, slightly folded lips; go- radial canals, in 10-15 lateral folds on either side,
nads from near centre of manubrium almost to cir- each further divided into 2-5 lamellar folds, not
cular canal, with numerous lateral folds; no cirri; attached to subumbrella; about 300 marginal tenta-
adaxial ocelli on each bulb and cordylus: Laodicea cles and as many cordyli with some few cnidocysts.
pulchra Browne, 1902 (Fig. 3.98) Ptychogena hyperborea Kramp, 1942 (Fig.
la Manubrium with short perradial lobes; usually 1 3.103). Umbrella 15 mm wide, 8 mm high, very
cordylus between tentacles or rudimentary bulbs thick; manubrium broad, quadrangular, with 4
............................................................................. .2 large perradial lobes; mouth somewhat folded;
2 Up to 180-600 marginal tentacles; ocelli present gonads in 2-3 pairs of lateral diverticula on either
............................................................................. .3 side of proximal half of radial canals above manu-
2a 4 perradial marginal tentacles and 4 small interra- brial lobes; about 80 marginal tentacles, probably
dial bulbs without abaxial spurs; ocelli not as many cordyli.
observed. Umbrella 1 mm wide; manubrium
small; mouth with 4 simple lips; gonads large, lon- Genus Staurophora Brandt, 1834
gitudinally divided, extending from stomach Laodiceidae with unusual cross-shaped manubrium;
almost to circular canal; 8 cordyli and a few cirri: mouth openings extending along the 4 radial canals
Laodicea minuscula Vannucci, 1957 (Fig. 3.99) transformed for a long distance into open grooves, only
3 Up to 180 marginal tentacles, without basal spurs most distal parts remaining free and closed; mouth arms
in adult specimens; ocelli on about every second slit-like, with strongly folded lips; gonads on diverticu-
tentacle. Umbrella 20-25 mm wide, two or four la in lateral walls of cruciform, enlarged, mouth-radial
times wider than high; manubrium cross-shaped, canal complex; without cirri; with adaxial ocelli.
fairly large; mouth with short, slightly folded lips; Staurophora mertensii Brandt, 1834 (Fig. 3.104).
gonads from corners of manubrium along half part Umbrella 100-200 mm wide, flatter than hemi-
of radial canals or almost to bell margin, sinuous; spherical; numerous hollow tentacles (up to seve-
small abaxial spurs on young tentacles; cirri ral thousand), approximately alternating with cor-
present (often lost after preservation); one cordy- dyli and each with adaxial ocelli.
lus between every two tentacles: Laodicea indica
Browne, 1905 (Fig. 3.100) Genus Toxorchis Haeckel, 1879
3a Up to 200-600 marginal tentacles, with basal Laodiceidae with 4, 6, or more main radial canals,
abaxial spurs; ocelli present usually on each third some or all branching dichotomously one or more
or fifth tentacle. Umbrella, up to 37 mm wide, usu- times, all branches reaching circular canal, primary
ally much smaller, flatter than hemispherical; radial canals not extending to circular canal; gonads
mesoglea thin; manubrium quadrate, short, on outermost branches; numerous tentacles and cor-
attached to subumbrella along arms of perradial dyli; with or without cirri; with or without ocelli.
cross; 4 long sinuous gonads along radial canals, Toxorchis polynema Kramp, 1959 (Fig. 3.105)
contiguous with manubrium; with usually 1-2 spi- (see Bouillon, 1984). Umbrella 17 mm wide, flat;
Hydromedusae 427
manubrium quadrate, broad, flat; mouth with hemispherical; manubrium flask-shaped; 4 short,
broad crenulated lips; 4 groups of radial canals, slightly folded lips; typically 4 radial canals, some-
each bifurcating twice inside cruciform base of times up to 10 but then asymmetrically arranged;
manubrium; 16 (4x4) radial canals leaving manu- gonads linear on distal half of radial canals; 16-36
brium, all running to circular canal; gonads rib- tentacles, with large, conical marginal bulbs; gen-
bon-like, along proximal 213 to 314 of length of erally 1-3 rudimentary tentacle bulbs between each
radial canals, close to manubrium; about 360 mar- pair of tentacles, middle one the largest; all margi-
ginal tentacles with endodermal roots extending nal bulbs with excretory papillae; with 4-6 stato-
into bell mesoglea, and somewhat fewer cordyli; cysts between successive tentacles.
about 80 adaxial ocelli.
Genus Octocanna Haeckel, 1879
Family Lovenellidae Malagazziidae with 8 radial canals; 8 gonads on ra-
Leptomedusae with short manubrium; without gastric dial canals; without ocelli; 4 hollow marginal tenta-
peduncle; without excretory pores; with 4 simple radial cles and 4 small, non-tentacular marginal bulbs.
canals; marginal tentacles hollow, with lateral cirri; Octocanna haeckeli Vannucci and Moreira, 1966
without marginal cirri; gonads on radial canals, not (Fig. 3.108).Umbrella 3.5 mm wide, 2.5 mm high,
reaching manubrium; statocysts 16 or more; no ocelli. almost globular, mesoglea very thick; mouth with
4 long lips; excretory pores on small adradial
Genus Lovenella Hincks, 1868 papillae, gonads at distal end of proximal third of
With characters of family. radial canals; with 8 statocysts.
Lovenella cirrata (Haeckel, 1879) (Fig. 3.106).
k Umbrella 16 rnm wide, hemispherical; manubrium Genus Octophialucium Kramp, 1955
short, urn-shaped; 4 simple lips; 4 spindle-shaped Malagazziidae with 8 or more symmetrical radial
gonads on distal half of radial canals; 8- 16 marginal canals; manubrium with 8 lips.
tentacles with large bulbs, flanked by 5-7 pairs of Octophialucium bigelowi Kramp, 1955 (Fig.
lateral spiral cirri; 3-5 rudimentary marginal warts 3.109). Umbrella, 8 mm wide, 10 mm high, almost
between successive tentacles; about 16 statocysts. spherical; mesoglea very thick; 8 pointed lips; 8
radial canals; gonads swollen, along almost entire
Family Malagazziidae length of radial canals; 8 large tentacles; 1-3 rudi-
Leptomedusae with small manubrium; without gastric mentary triangular bulbs between successive ten-
peduncle; 4-8, sometimes up to 11 radial canals; go- tacles; 3-5 statocysts between tentacles.
nads completely surrounding radial canals, separated
from manubrium; with adaxial excretory papillae; Genus Tetracanna Goy, 1979
rudimentary marginal bulbs not permanent (all bulbs Malagazziidae with 4 radial canals; 8 marginal tenta-
potentially transforming into tentacles); with closed cles; mouth with 4 long lips; gonads on almost entire
statocysts; without ocelli. length of radial canals; 1-3 statocysts between succes-
1 Normally with 4 radial canals (sometimes up to sive tentacles. (Family assignment doubtful; no excre-
10, but then asymmetrically arranged) ............... ..3 tory pores have been mentioned.)
la Normally with 8 or more symmetrically arranged Tetracanna octonema Goy, 1979 (Fig. 3.1 10).
radial canals......................................................... .2 Umbrella 6 mm wide, globular, apical mesoglea
2 Mouth with 4 lips: Octocanna thick.
2a Mouth with 8 lips: Octophialucium
3 With 8 tentacles: Tetracanna Family Mitrocomidae
3a With 16-32 marginal tentacles: Malagazzia Leptomedusae with bases of manubrium attached to
subumbrella along continuation of radial canals; with 4
Genus Malagazzia Bouillon, 1984 or more simple radial canals; marginal tentacles hol-
Malagazziidae typically with 4 radial canals; manu- low; marginal cirri present in some genera; gonads oval
brium with four lips. or linear, only on radial canals; with open statocysts.
Malagazzia carolinae (Mayer, 1900) (Fig. 3.107). 1 With 12-16 radial canals: Halopsis
Umbrella 14-20 mm wide, 6-8 mm high; sub- l a With 4 radial canals .............................................. 2
428 Hydromedusae
2 Without marginal cirri: Cosmetirella 5-8 marginal cirri between successive tentacles:
2a With spiral marginal cirri with terminal clusters of Mitrocomella grandis Kramp, 1965 (Fig. 3.1 13)
cnidocysts .............................................................
3 2 With about 16 marginal tentacles; gonads oval,
3 With 8-16 statocysts: Mitrocomella short, swollen. Umbrella 4-7 mm wide; gonads
3a With numerous (20-160) statocysts: Mitrocoma near distal ends of radial canals; usually 16 (up to
24) marginal tentacles; 6-8 marginal cirri between
Genus Cosmetirella Browne, 1910 successive tentacles; 8 statocysts: Mitrocomella
Mitrocomidae with 4 radial canals; with 8 statocysts; brownei (Kramp, 1930) (Fig. 3.1 14)
without marginal cirri. 2a With 32-72 marginal tentacles; gonads elongated
Cosmetirella davisi (Browne, 1902) (Fig. 3.1 11). .............................................................................
.3
Umbrella up to 60 mm wide, larger in Subantarctic 3 Gonads linear, folded, along greater part of radial
than in Antarctic specimens, almost hemispheri- canals, leaving both ends free; 8 marginal cirri
cal; manubrium small; mouth with somewhat fol- between successive tentacles. Umbrella 13-17 mm
ded lips; gonads linear along 112-213 of radial wide, almost hemispherical, with thin walls; go-
canals; number of tentacles very variable, up to nads hanging down in large vertical folds; 32-72
180; normally 8 statocysts. marginal tentacles: Mitrocomella frigida (Browne,
1910) (Fig. 3.115)
Genus Halopsis A. Agassiz, 1863 3a Gonads laterally compressed, sac-like, pendent on
Mitrocomidae with more than 8 radial canals; with distal half or third of radial canals; 1 (sometimes 0
marginal spirally coiled cirri; with numerous (about or 2) marginal cirrus between successive tentacles.
80) statocysts. Umbrella 6 mm wide, 4.2 mm high, higher than
Halopsis ocellata A. Agassiz, 1863 (Fig. 3.1 12). hemispherical; mesoglea thick; manubrium cruci-
Umbrella 50-65 mm wide, about 4 times as wide as form extending along radial canals; 60 marginal
high, watch-glass-shaped; mesoglea thick toward tentacles: Mitrocomella millardae Pages, Gili and
centre; manubrium broad, flat, 115 of bell diameter, Bouillon, 1992 (Fig. 3.1 16)
circular to star-shaped in outline; mouth with 4
fairly short lips; 12-16 (17) radial canals in 4 Family Orchistomidae
groups branching usually within outline of manu- Leptomedusae with very short manubrium; with large
brium; gonads linear, about 213 of radial canals; up gastric peduncle; mouth with 8-30 sinuous or crenu-
to 450 marginal tentacles; 1 marginal cirrus lated lips; with 8 or more radial canals, simple, rami-
between successive tentacles; about 80 statocysts. fied, or in clusters of 4; up to 64 marginal tentacles,
laterally compressed; no marginal cirri, but numerous
Genus Mitrocoma Haeckel, 1864 filiform tentaculiform structures devoid of marginal
Mitrocomidae with 4 radial canals; with marginal spi- bulbs, not in contact with circular canal, in each inter-
rally coiled cirri; with numerous statocysts (20-100). tentacular space (Fig. 2B); gonads usually on proxi-
Mitrocoma minervae Haeckel, 1879. Doubtfill mal parts of radial canals; numerous (up to 800) adax-
species. Umbrella size dubious; 120-160 marginal ial ocelli; no statocysts or cordyli; without excretory
tentacles; 1 marginal cirrus between successive pores or papillae.
tentacles; 120-160 statocysts.
Genus Orchistoma Haeckel, 1879
Genus Mitrocomella Haeckel, 1879 With characters of family.
Mitrocomidae with 4 radial canals; with marginal spi- Orchistoma collapsa (Mayer, 1900) (Fig. 3.1 17).
rally coiled cirri; with 8, 12, or 16 statocysts. With 16 radial canals in 4 groups of 4. Umbrella 7
1 With 8 marginal statocysts ...................................2 mm wide and high, apex dome-shaped, side walls
l a With 16 marginal statocysts. Umbrella 52 mm vertical; manubrium large; with large, prominent
wide, watch-glass-shaped, mesoglea thin at gastric peduncle; mouth with 8 slightly crenulated
umbrellar margin, thick in central part; manubri- lips; gonads in proximal parts of radial canals; 16
um cruciform, extending perradially; mouth with well developed and 112 rudimentary tentacles, all
4 large, pointed, highly crenulated lips; gonads with adaxial ocelli.
linear along greater part of radial canals, about 220 Orchistoma pileus (Lesson, 1843) (Fig. 3.1 18).
marginal tentacles, all alike, with elongated bulbs; With 32 simple radial canals. Umbrella 30-40 mm
Hydromedusae 429
wide, flat-topped, with vertical sides, apical meso- than hemispherical, with rounded apex; manu-
glea very thick; manubrium wide and short; with brium broad, quadrangular, with 4 perradial
wide, large gastric peduncle; mouth with 32 long, pouches extending 112-213 distance towards bell
complexly crenulated lips; 32 swollen gonads on margin; 4 radial canals; mouth with 4 short, slight-
radial canals near manubrium; 64 short marginal ly crenulated lips; 10-16 sac-like gonads on each
tentacles with large bulbs; about 100 slender ten- side of each manubrial pouch; 20-24 marginal ten-
taculiform structures and about 400 adaxial ocelli. tacles; between successive tentacles 2 (rarely 1)
minute cordyli-like appendages with distal bundle
Family Phialellidae of cnidocysts; cnidome: microbasic euryteles.
Leptomedusae with small manubrium; without gastric
peduncle; 4 radial canals; gonads on radial canals, Genus Margalefia Pagi?s, Bouillon and Gili, 1991
separated from manubrium and divided into two late- Tiarannidae with 4 simple, smooth gonads on the
ral parts by median groove; hollow tentacles; without walls of perradial manubrial pouches.
excretory pores; without lateral or marginal cirri; with Margalefia intermedia Pagks, Bouillon and Gili,
8 closed statocysts; without ocelli. 1991 (Fig. 3.121). Umbrella 26 mm wide, 17 mm
high, higher than hemispherical; mesoglea thick;
Genus Phialella Browne, 1902 manubrium cruciform, elongate, with 4 large per-
With characters of family. radial pouches extending almost to bell margin;
Phialella falklandica Browne, 1902 (Fig. 3.1 19). gonads simple, smooth on manubrial pouches;
Umbrella up to 17 mm wide, semi-globular; mouth with 4 long, folded, slightly crenulated lips;
manubrium short, quadrangular; mouth with 4 with about 150 marginal tentacles; about one cor-
fimbriated lips; gonads along greater part of radial dylus per each 3 marginal tentacles; cnidome:
canals, hanging down in wavy folds; about 60 microbasic euryteles and desmonemes.
marginal tentacles; with 8 statocysts on broad
cushion-like bulbs. Genus Modeeria Forbes, 1848
Tiarannidae with gonads on manubrium and extend-
Family Tiarannidae ing outwards along perradial manubrial pouches.
Leptomedusae with wide, cross-shaped manubrium, Modeeria rotunda (Quoy and Gaimard, 1827)
with 4 perradial pouches joined to subumbrella; (Fig. 3.122). Umbrella 20 mm wide, somewhat
mouth with 4 simple or crenulated lips; with 4 simple less high, hemispherical, mesoglea thick, apex
radial canals; gonads folded on interradial walls of rounded; manubrium short, broad, cruciform, per-
manubrium and/or on perradial manubrial pouches; radial edges of manubrium connected over entire
marginal tentacles hollow; with hollow cordyli bear- length with subumbrella; mouth with 4 large,
ing cnidocysts; without ocelli. slightly crenulated lips; gonads in regular trans-
1 With regularly transversely folded gonads on inter- verse folds on interradial walls of manubriurn,
radial walls of manubrium and extending outwards extending outwards on perradii; 16-28 marginal
along perradial manubrial pouches: Modeeria tentacles; 2-3 minute cordyli-like appendages with
la With gonads on perradial manubrial pouches only distal bundle of cnidocysts between successive
..................................................,...........................2 tentacles; cnidome microbasic euryteles.
2 With 4 simple smooth gonads on walls of perra-
dial manubrial pouches: Margalefia Family Tiaropsidae
2a Gonads in eight adradial rows of 10-16 sac-like Leptomedusae with 4 or 8 (exceptionally up to 16) ra-
invaginations on each side of perradial manubrial dial canals; with one or two types of marginal tentacles
pouches: Chromatonema (long and rudimentary tentacles, both with marginal
bulbs); without marginal cirri; sense organs compound,
Genus Chromatonema Fewkes, 1882 comprising ecto-endodermal ocelli and open statocysts.
Tiarannidae in which the gonads are represented by 8
(10-16) series of sac-like invaginations from the sur- Genus Tiaropsidium Torrey, 1909
face of perradial pouches, separated in interradi. Tiaropsidae with 4 or more (up to 16) simple radial
Chromatonema rubra Fewkes, 1882 (Fig. 3.120). canals; with 8 or 16 (rarely 48) compound statocysts;
Umbrella up to 27 mm wide, 22 mm high, higher with two kinds of tentacles; without marginal cirri.
Hydromedusae
Tiaropsidium roseum (Maas, 1905) (Fig. 3.123). statocysts between successive tentacles: Clytia
Umbrella up to 15 mm wide, flattened; manu- lomae (Torrey, 1909) (Fig. 3.125)
brium short and broad; mouth with 4 very short 3a Gonads elongated, slightly folded along distal 112-
crenulated lips; gonads elongate oval, along some- 314 of radial canals; 60-85 marginal tentacles +
what more than 113 the length of 4 radial canals; 4 young bulbs. Umbrella up to 22 mrn wide, 10 mm
long perradial marginal tentacles; 7 rudimentary high, watch-glass-shaped; manubrium short;
tentacles in each quadrant; 8 complex statocysts. mouth with 4 large fimbriated lips; bulbs globular;
' 1 statocyst between successive tentacles; speci-
Order Proboscoida (Fig. 1G) mens with 3 and 6 radial canals: Clytia simplex
Hydranths with flared to globose hypostome, forming (Browne, 1902) (Fig. 3.126)
a "buccal cavity" beneath the mouth; medusae mor- 4 Gonads oval or linear 112-314 as long as radial
phologically variable. canals, closer to margin than to manubrium; 16-58
tentacles. Umbrella up to 20 mm wide, nearly he-
Family Campanulariidae mispherical, mesoglea fairly thin; manubrium
Leptomedusae with short manubrium; without gastric small, quadrate; mouth with 4 simple lips; 4 (some-
peduncle; typically with 4 radial canals (except in times more, up to 12) radial canals; bulbs globular;
Gastroblasta and Pseudoclytia); with or without 1-3 (usually 2) statocysts between successive ten-
velum (no velum in Obelia); gonads on the radial tacles: Clytia hemisphaerica (Linnaeus, 1767)
canals, completely surrounding them and separated (Fig. 3. 127)
from manubrium; with hollow tentacles (except in 4a Gonads thick, cylindrical, along almost entire
Obelia, where they are solid and with short endoder- length of radial canals; 16 tentacles. Umbrella 4
mic extensions into bell mesoglea); without excretory mm wide, almost hemispherical; manubrium
papillae or pores; without lateral or marginal cirri; small, urn-shaped, with bulging sides; mouth with
numerous (16-200) closed marginal statocysts (only 8 4 recurved slightly folded lips; female gonads with
in Obelia, located on the underside of the basal bulb of very few large eggs; usually 3 statocysts between
some of the marginal tentacles); no ocelli. Two gene- adjacent tentacles: Clytia discoida (Mayer, 1900)
ra: Clytia (with hollow tentacles and normal velum), (Fig. 3.128)
and Obelia (with solid tentacles; without velum).
Clytia noliforme (McCrady, 1859). Only young
Genus Clytia Lamouroux, 18 12 medusae known. Umbrella 0.4 mm wide, 0.3 mm
Campanulariidae with short manubrium; normal high, bell-shaped; manubrium tubular; mouth
velum; hollow marginal tentacles; numerous statocysts. quadrate, 4 radial canals; 4 marginal tentacles;
1 Prominent, brown pigment spot at the base of each slight swellings interradially; 8 statocysts; gonads
one of the about 30 short, thick tentacles; umbrella lacking.
2 mm wide, 0.7 mm high; manubrium very short Clytia ovule (Mayer, 1900) is an abnormal Clytia (see
and broad; mouth with four lips; gonads large, Kramp, 1961); Clytia phosphoricum (Peron and
thick, prominent, on proximal parts of radial canals; Lesueur, 1810) is an old general name, covering all
.......32-40 statocysts: Clytia brunescens (Bigelow, Clytia with alike number of statocysts and tentacles.
1904) (Fig.3.124)
la No brown pigment spots at base of tentacles .......2 Genus Obelia P6ron and Lesueur, 18 10
2 Statocysts in about same number as tentacles + Campanulariidae with short quadrangular manu-
young bulbs ..........................................................3 brium; without velum; with numerous solid, stiff, not
2a Statocysts about twice as numerous as tentacles + extensile tentacles; tentacles with short endodermal
young bulbs ..........................................................4 roots extending into bell mesoglea; 8 statocysts loca-
3 Gonads narrow, elongated along less than 112 of ted on underside of basal bulbs of marginal tentacles.
the distal part of radial canals; about 32 tentacles Obelia spp. (Fig. 3.129). Umbrella 2.5-6 mm wide,
and some young bulbs. Umbrella 9-12 mm (rarely flat, mesoglea very thin; without gastric peduncle;
14 mm) wide, about 4 times broader than high, mouth with 4 simple lips; 4 radial canals; gonads
thin; manubrium short, cruciform; mouth with 4 spherical to ovoid, sac-like, hanging from middle
slightly frilled lips; bulbs elongated; 1 (rarely 2) to end of radial canals. Various nominal species of
Hydromedusae 43 1
Obelia hydroids are common throughout the 2a Tentacles not in groups ........................................3
world, but there are still no characters to distin- 3 Some or all tentacles with adhesive pads .............4
guish their medusa stages with certainty. 3a All tentacles without adhesive pads: Aglauropsis
According to Zamponi and Genzano (1990), the 4 With two kinds of tentacles, with and without
medusae of Obelia dichotoma Hincks, 1868 and adhesive pads.. ....................................................
..5
Obelia longissima (Pallas, 1766), whose hydroids 5 Adhesive pads terminal: Vallentinia
are both present in the South Atlantic, can be 5a Adhesive pads at some distance from distal end of
distinguished by their cnidome, composed of atri- tentacles: Cubaia
chous isorhizas, atrichous anisorhizas and basitri-
chous isorhizas in 0 . dichotoma, and by microba- Genus Aglauropsis F. Muller, 1865
sic mastigophores and macrobasic mastigophores Olindiidae with four radial canals; without centripetal
in 0 . longissima. However, macrobasic mastigo- canals; numerous tentacles of one kind on bell mar-
phores seem improbable in the genus Obelia, as gin, without adhesive pads, not arranged in groups;
well as in any other Leptomedusae (contamina- numerous statocysts. (there is no adequate description
tion?). for Aglauropsis agassizi Muller, 1865, this species is
probably similar to A. kawari).
Subclass Laingiomedusae 1 Gonads wavy, curtain-like, transversally lobed,
along almost entire length of radial canals.
Family Laingiidae Umbrella up to 22 mm wide and 15 mm high;
With characters of subclass. mouth with 4 large folded lips, each with band of
cnidocysts; tentacles about 200, in 2-3 rows; sta-
Genus Kantiella Bouillon, 1978 tocysts over 50: Aglauropsis conanti Browne,
Laingiidae with exumbrellar cnidocyst bands; gonads 1902 (Fig. 3.13 1)
on walls of four manubrial interradial pouches; 4 short l a Gonads smooth, not transversally lobed .............. 2
marginal tentacles with terminal cluster of cnidocysts, 2 Gonads pouch-like, hanging from proximal part of
above peronia-like structures. radial canals. Umbrella 5.5 mm wide and 4.0 mm
Kantiella enigmatica Bouillon, 1978 (Fig. 3.130). high; mouth with 4 small, folded lips; gonads con-
Umbrella 3-4 rnm wide, 2-3 mm high, hemisphe- tinuous with corners of manubrium walls; tenta-
rical, mesoglea thick at apex; manubrium quad- cles 40: 20 above bell margin, 20 on bell margin;
rangular; with short, broad gastric peduncle; 20 statocysts: Aglauropsis kawari Moreira and
mouth margin quadrate or circular, covered by Yamashita, 1972 (Fig. 3.132)
ring of cnidocysts. 2a Gonads curtain-like, straight, extending along half
of length of radial canal (absent proximally), their
Subclass Limnomedusae distal ends pendent; about 110 closely packed ten-
Two families: Olindiidae (with gonads on radial tacles; umbrella 15 mm wide, 11 mm high; mouth
canals only, except the freshwater genus Limnocnida; with 4 small, folded lips, each with large band of
with internal statocysts; without ocelli), and cnidocysts; 46 statocysts: Aglauropsis edwarsii
Proboscidactylidae (with gonads on the manubrium Pagks, Bouillon and Gili, 1991 (Fig. 3.133).
extending over the proximal part of radial canals; 4-6
or more radial canals, usually branched; without sta- Genus Cubaia Mayer, 1894
tocyst and without ocelli). Olindiidae with 4 simple radial canals; without cen-
tripetal canals; with 2 series of tentacles: one series
Family Olindiidae (20) issuing from exumbrella above bell margin, with
Limnomedusae with or without centripetal canals; terminal adhesive disks and about 8 cnidocyst rings,
internal statocysts; simple, unbranched radial canals; the other series (50-60) without adhesive disks, with
gonads on radial canals; without ocelli. 25-30 cnidocyst rings arising from bell margin;
1 With centripetal canals: Olindias numerous statocysts.
la Without centripetal canals ....................................2 Cubaia aphrodite Mayer, 1894 (Fig. 3.134).
2 Tentacles in groups on bell margin; without adhe- Umbrella 12 mm wide, flatter than hemispherical;
sive pads: Gossea manubrium flask-shaped; mouth with 4 lips; go-
432 Hydromedusae
nads papilliform on middle region of radial canals; high as wide; gonads sac-like, folded, on proximal
35 statocysts. part of radial canals; four to eight large tentacles
located at some distance from radial canals, occa-
Genus Gossea L. Agassiz, 1862 sionally also four similar interradial large tenta- ,
Olindiidae with four radial canals; without centripetal cles, all with terminal adhesive pads; 64-75 most-
canals; with one kind of tentacles arranged in groups; ly non-adhesive tentacles with numerous /
without adhesive pads. cnidocyst rings arising from bell margin; 1-2 sta- 1I
Gossea brachymera Bigelow, 1909 (Fig. 3.135). ' tocysts between successive tentacles. I
Umbrella up to 20 mm in diameter; with short,
broad gastric peduncle; gonads extending from Family Proboscidactylidae
base of peduncle to circular canal, wavy, their dis- Lirnnomedusae without statocysts or ocelli; without
tal ends sac-like, pendent; with 4 perradial and 4 centripetal canals; manubrium usually with 4-6 or more
interradial cnidocysts pads, each with one large radial gastric lobes extending along proximal portions
and one minute tentacle; in addition, several iso- of radial canals; with gonads surrounding manubrium
lated tentacles at different stages of development; and usually extending onto gastric lobes; radial canals
without dwarf tentacles; 8 statocysts enclosed in generally branched, obliterated canals may be present;
perradial and interradial pads. with or without exumbrellar cnidocysts clusters or
bands alternating with marginal tentacles.
Genus Olindias Muller, 1861
Olindiidae with 4 radial canals and numerous centri- Genus Proboscidactyla Brandt, 1834
petal canals; numerous tentacles of two kinds: pri- Proboscidactylidae with manubrium with radial gas-
mary tentacles issuing above bell margin, with distal tric lobes; gonads on manubrium and gastric lobes; 4-
adhesive pads and cnidocysts in transverse clasps, and 6 or more branched radial canals, with clusters or
secondary tentacles on bell margin, without adhesive bands of cnidocysts on exumbrella; usually without
pads, with cnidocysts in rings; gonads with papilli- circular canal; marginal tentacles hollow.
form processes; numerous marginal clubs which may 1 Usually with four primary radial canals ...............2
transform into tentacles, statocysts usually in pairs at l a With more than four primary radial canals .........3
base of primary tentacles. 2 With four primary radial canals branching into 16-
Olindias phosphorica (Delle Chiaje, 1841) (Fig. 20 (rarely more) terminal branches and as many
3.136). 1 1- 19 centripetal canals per quadrant; 30- tentacles. Umbrella 5 mm wide, slightly higher
60 primary tentacles; 100-120 secondary tenta- than hemispherical; manubrium with 4 pouches;
cles; 100-170 marginal clubs; 40-60 mm wide. mouth with 4 lips: Proboscidactyla ornata
Olindias sambaquiensis Muller, 186 1 (Fig. (McCrady, 1859) (Fig. 3.140)
3.137). 2 1-27 centripetal canals per quadrant; 80- 2a With four primary radial canals branching into
100 primary tentacles; 200-300 secondary tenta- about 60 terminal branches. Umbrella 9 mm wide,
cles; 100-200 marginal clubs; 50- 100 mm wide. 4.8 mm high; manubrium extending along 8-16
manubrial pouches; mouth with poorly marked lips;
Genus Vallentinia Browne, 1902 up to 60 marginal tentacles: Proboscidactyla meno-
Olindiidae with 4 radial canals, without centripetal ni PagGs, Bouillon and Gili, 1991 (Fig. 3.141)
canals; with 4 to 8 hollow tentacles with terminal 3 Usually 6 primary radial canals; up to 24 terminal
adhesive pads, and numerous tentacles without adhe- branches and tentacles. Umbrella 9 mm wide, 8
sive pads but with numerous rings of cnidocysts; with mm high; mouth with 6 folded lips; marginal ten-
16 or more statocysts. tacles with adaxial basal cnidocyst cushions:
Vallentiniafalklandica Browne, 1902 (Fig. 3.138). Proboscidactyla stellata (Forbes, 1 846) (Fig.
Umbrella 2 rnm wide, 3 mm high; gonads oval, sac- 3.142)
like, on upper part of radial canals. With four large 3a With 5-1 1 (usually 8) primary radial canals; 24-54
perradial marginal tentacles, with terminal adhesive terminal branches and tentacles. Umbrella 6 mm
pads and 24 non-perradial, non-adhesive, tentacles wide and high; usually 8 (but frequently 6) manu-
with numerous cnidocyst rings; 16 statocysts. brial pouches; mouth with closely folded lips:
Vallentinia gabriellae Mendes, 1948 (Fig. 3.139). Proboscidactyla mutabilis (Browne, 1902) (Fig.
Umbrella up to 6 mm wide, about two thirds as 3.143)
Hydromedusae 433
Subclass Narcomedusae Solmundella bitentaculata (Quoy and Gaimard,

1 Without manubrium pouches: Solmarisidae 1833) (Fig. 3.146). Umbrella up to 12 mm; apical
la With manubrium pouches ....................................2 mesoglea very thick; tentacles issuing from
2 Pouches perradial: Cuninidae umbrella near apex, 8-32 statocysts.
2a Pouches interradial: Aeginidae
Family Cuninidae
Family Aeginidae Narcomedusae with perradial and undivided manubrial
Narcomedusae with interradial divided manubrial pouches, with or without peripheral canal system; tenta-
pouches containing gonads; with or without periphe- cles leaving umbrella opposite centre of each manubrial
ral canal system. With perradial primary tentacles pouch and thus equal in number to that of pouches;
leaving umbrella between marginal pouches, number- pouches not extending beyond point of origin of tenta-
ing at least half that of manubrial pouches; with or cles. With or without otoporpae. Two genera: Solmissus
without secondary tentacles on umbrella margin. (without otoporpae) and Cunina (with otoporpae).
Pouches extending beyond point of origin of primary
tentacles. With or without otoporpae. Genus Cunina Eschscholtz, 1829
1 2 tentacles: Solmundella Cuninidae with otoporpae, with or without peripheral
la 4 or more tentacles ..............................................
.2 canal system.
2 8 (or more) tentacles and twice as many manubri- 1 With peripheral canal system ...............................2
a1 pouches; with secondary tentacles: Aeginura l a Without peripheral canal system, manubrial
2a 4-6 tentacles, 4-6 peronia and 8-12 manubrial pouches broad, with parallel sides .......................4
pouches: Aegina 2 Manubrial pouches, tapering from broad base, sep-
arated by wide triangular spaces. Umbrella about 8
Genus Aegina Eschscholtz, 1829 mm wide, dome-shaped; 6-9 tentacles and pouches;
Aeginidae typically with 8 (but occasionally 10 to 12) marginal lappets square, with 4 statocysts, peri-
primary manubrial pouches; with peripheral canal pheral canals broad and flat; otoporpae linear:
system; Usually with 4 (sometimes 5 or 6) marginal Cuninafrugifera Kramp, 1948 (Fig. 3.147)
primary tentacles; without secondary tentacles; with- 2a Manubrial pouches with nearly parallel sides......3
out otoporpae. One species. 3 Manubrial pouches up to 10-14, wide and quadrate,
Aegina citrea Eschscholtz, 1829 (Fig. 3.144). more than twice as wide as septa between them.
Umbrella up to 50 mm, mesoglea thick at apex; 4 Umbrella up to 18 mm wide; conical or almost
tentacles; 4 lappets with numerous statocysts; 5 or globular; manubrium on broad peduncle; 10-14
6-rayed specimens frequent. tentacles arising slightly above margin; peripheral
canals narrow; marginal lappets short and broad,
Genus Aeginura Haeckel, 1879 with 3 statocysts; otoporpae short and oval: Cunina
Aeginidae with 16 primary manubrial pouches; globosa Eschscholtz, 1829 (Fig. 3.148)
peripheral canal system absent or degenerated; 8 pri- 3a Manubrial pouches up to 29, elongate rectangular,
mary tentacles and peronia; with secondary tentacles hardly broader than space between them; lateral
on umbrella margin; without otoporpae. portions of peripheral canals very broad, transverse
Aeginura grimaldii Maas, 1904 (Fig. 3.145). portions narrow. Umbrella up to 58 mm, rather flat;
Umbrella up to 45 mm; manubrial pouches with manubrium pouches increasing with age from 9 in
slight secondary division; 3-5 secondary marginal juvenile specimens; gonads forming continuous
tentacles in each octant, 1 or 2 statocysts between folded band, intermptedly following edge of manu-
each secondary tentacle. Colour of manubrium brium with its pouches; marginal lappets rectangu-
deep chocolate to purplish black. lar, with 2-3 statocysts; otoporpae very small:
Cunina duplicata Maas, 1893 (Fig. 3.149)
Genus Solmundella Haeckel, 1879 4 Usually 8 (7-9) square manubrial pouches, very
Aeginidae with 8 manubrial pouches; without peri- close together. Umbrella 5-7 mm wide, flatter than
pheral canal system; 4 peronia but only 2 long tentacles; hemispherical; no peripheral canals; thick and
without secondary tentacles; without otoporpae. One broad ectodermal pad below base of each tentacle;
species. marginal lappets with 2-5 generally 3 statocysts;
434 Hydromedusae
otoporpae small: Cunina octonaria McCrady, Genus Pegantha Haeckel, 1879

1859 (Fig. 3.150) Solmarisidae with gonads forming diverticula of mar-
4a Usually about 12 (8-14) manubrial pouches, gin of oral manubrium wall; with peripheral canal
square or somewhat rounded distally. Umbrella up system; with otoporpae.
to 14 mm; highly arched; manubrial pouches 1 Exumbrella with deep radiating furrows from ten-
increasing in number with age; no peripheral tacles to apex, surrounded by ribs and supplemen-
canals; ectodermal pad below base of tentacles tary ridges. Umbrella up to 30 mm wide, hemi-
small; marginal lappets short and broad, each with spherical; 12-16 tentacles; gonads with 2-4 lobes
4- 10 statocysts; otoporpae narrow, linear: Cunina projecting into lappet cavities; 12-16 lappets ovate,
peregrina Bigelow, 1909 (Fig. 3.15 1) pointed, each with 20 statocysts; otoporpae long
Cunina mucilaginosa (Chamisso and Eysenhardt, 1821) tapering outwards: Pegantha triloba Haeckel,
and Cunina oligotis Haeckel, 1879, reported from the 1879 (Fig. 3.154)
South Atlantic, are considered unrecognisable species. 1a Exumbrella smooth ............................................. .2
2 Some or all otoporpae long, peripheral canals nar-
Genus Solmissus Haeckel, 1879 row throughout their length.. ............................... .3
Cuninidae without otoporpae, without peripheral 2a All otoporpae short; peripheral canals broad, at
canal system. least in their lateral portions ................................. 4
1 With 8 double umbrellar lappets; 8 subumbrellar 3 12-16 rectangular marginal lappets, each with usu-
tentacles (primary tentacles?); 8 exumbrellar ten- ally 4-6 statocysts; two middle otoporpae long, la-
tacles (secondary tentacles?); 16 manubrial pouch- teral ones shorter. Umbrella up to 16 mm wide,
es; 1-3 statocysts on each lappet; umbrella 3-5 mm dome-shaped; gonads simple, smooth: Pegantha
wide: Solmissus atlantica Zamponi, 1983 (Fig. rubiginosa (Kolliker, 1853) (Fig. 2H, I, 3.155)
3.152). Doubtful species, poorly preserved, pro- 3a 20140 quadrate marginal lappets, each usually
bably an Aeginidae with 3-5 statocysts; with long, linear otoporpae.
la With single lappets ...............................................
2 Umbrella up to 50 mm wide and 20 mm high, len-
2 8-20 (usually 16) rectangular manubrial pouches. ticular; gonads somewhat crenulated: Pegantha
Umbrella up to 62 mm, flat, gelatinous disk, thick clara R. P . Bigelow, 1909 (Fig. 3.156)
and rigid, exumbrella smooth; tentacles long; lap- 4 Umbrella highly vaulted, thick; about 16 square
pet zone very thin, marginal lappets square, up to marginal lappets; lateral portions of peripheral
15 (rarely up to 21) statocysts per marginal lappet: canals broad at base, tapering outwards. Umbrella
Solmissus marshalli Agassiz and Mayer, 1902 up to 30 mm wide and about 20 mm high; margi-
(Fig. 3.153) nal lappets with 5-7 (or 9) statocysts; otoporpae
2a 24 heart-shaped manubrial pouches. Umbrella 20 short and narrow, about twice as long as width of
rnm wide, 7 mm high; biconvex; 24 tentacles; transverse section of peripheral canals; gonads
marginal lappets rectangular, 3 statocysts per mar- simple or lobed sacs: Pegantha martagon
ginal lappet: Solmissus faberi Haeckel, 1879. Haeckel, 1879 (Fig. 3.157)
Doubtful species 4a Umbrella flat, lenticular; 16-26 marginal lappets;
Solmissus bleekii Haeckel, 1879, from the Atlantic peripheral canals very broad throughout. Umbrella
coast of South Africa is regarded as a doubtful, unre- up to 40 mm wide; marginal lappets square, each
cognisable species. with 5-7 statocysts; otoporpae shorter or slightly
longer than width of transverse portions of peri-
Family Solmarisidae pheral canals; gonads sac-shaped when fully deve-
Narcomedusae without manubrial pouches, periphery loped, with oval or papilliform processes: Pegantha
of manubrium circular and unbroken. With or without laevis H. B. Bigelow, 1909 (Fig. 3.158).
peripheral canal system. Gonads on manubrial wall, or
on manubrial wall diverticula; numerous tentacles leav- Genus Solmaris Haeckel, 1879
ing umbrella at level of periphery of manubrium. With Solmarisidae without peripheral canal system; with-
or without otoporpae. Two genera: Pegantha (with out otoporpae, with simple annular gonads.
peripheral canal system; with otoporpae), and Solmaris Solmaris corona (Keferstein and Ehlers, 1861)
(without peripheral canal system; without otoporpae). (Fig. 3.159). Lens-shaped umbrella 12-15 mm
Hydromedusae 435
wide, with 30-36 or more tentacles and lappets; Liriope tetraphylla (Chamisso and Eysenhardt,
lappets rectangular, usually with 2 statocysts 1821) (Fig. 3.162). Umbrella 10-30 rnrn wide,
mounted on large cushion, with long bristles. hemispherical; manubrium on gastric peduncle of
Solmarisflavescens (Kolliker, 1853) (Fig. 3.160). varying size; mouth with 4 lips; 1-3 (or more)
Thick, lens-shaped umbrella 15-23 mm wide, with centripetal canals in each quadrant; 4 long hollow
12-17 tentacles and lappets; usually 13-15 tenta- perradial tentacles with cnidocyst rings and 4
cles; marginal lappets thin, quadrate, each with 2 small solid interradial tentacles with adaxial
statocysts. cnidocyst clusters; gonads variable in shape and
size; 8 statocysts.
Subclass Trachymedusae
1 Numerous tentacles arranged in groups; most with Family Halicreatidae
adhesive disks; manubrium with 8 radial lobes Trachymedusae with wide, circular manubrium; with-
(this diagnosis excludes the genus Tesserogastria, out peduncle; without centripetal canals; 8 or more
not present in South Atlantic): Ptychogastriidae radial canals; numerous marginal tentacles of differ-
la Tentacles without adhesive disks .........................2 ent size, but all structurally alike and arranged in a
2 With centripetal canals: Geryoniidae single series; each marginal tentacle with a flexible
2a Without centripetal canals ................................... .3 proximal portion and a stiff spine-like distal portion;
3 With broad, circular manubrium and broad radial with free ecto-endodermal statocysts.
canals: Halicreatidae 1 With about 16 or more radial canals: Halitrephes
3a Manubrium and radial canals narrow: l a With 8 radial canals .............................................. 2
Rhopalonematidae 2 With tentacles arranged in 16 groups: Botiynema
2a With tentacles in a continuous row ......................3
Family Geryoniidae 3 With perradial gelatinous papillae on exumbrella:
Trachymedusae with gastric peduncle; 4-6 radial Halicreas
canals; with centripetal canals; gonads on radial canals, 3a Without exumbrellar papillae: Haliscera
flattened and leaf-shaped; 2 kinds of marginal tenta-
cles, solid and hollow; ecto-endodermal statocysts Genus Botrynema Browne, 1908
enclosed in mesoglea. Two genera: Geryonia (with 6 With 8 radial canals; 16 groups of 11- 12 tentacles (2
radial canals and gonads, mouth with six lips) and groups with many tentacles in a single row in each
Liriope (usually with 4 radial canals and gonads, some- octant) and 8 solitary perradial tentacles.
times more, see below, mouth with four lips). Zamponi Botrynema brucei Browne, 1908 (Fig. 3.163).
and Genzano (1988) described 3 new genera of Umbrella 25 mm wide; apical mesoglea extreme-
Subantarctic Geryoniidae with mouth with 4 lips, ly thick, terminating in a distinct, sharp knob;
based essentially on the number of radial canals and manubrium short, circular; gonads oval, on proxi-
gonads: Pentarradiata, Heptarradiata, Octorradiata; mal or central halves of radial canals; usually 3
we consider these as abnormal specimens of Liriope. statocysts in each interradial space and 1-2 on
either side of the 8 perradial, solitary tentacles.
Genus Geryonia Ptron and Lesueur, 1810
One species. Genus Halicreas Fewkes, 1882
Geryonia proboscidalis (Forskal, 1775) (Fig. With 8 radial canals; with continuous row of tenta-
3.161). Umbrella 35-80 rnm wide, hemispherical; cles; perradial gelatinous papillae on exumbrella. One
manubrium small, on long, conical, gastric pedun- species.
cle; mouth with 6 lips; up to 7 centripetal canals Halicreas minimum Fewkes, 1882 (Fig. 3.164).
between radial canals; gonads heart-shaped, very Umbrella 30-40 mm wide, thick, disk-like, with
broad above; 6 long, perradial hollow tentacles with small apical projection; 8 clusters of gelatinous
cnidocyst rings, and 6 small, solid interradial tenta- papillae above margin, mouth wide circular open-
cles with adaxial cnidocyst clusters; 12 statocysts. ing; 8 broad, band-like, radial canals; broad circu-
lar canal; gonads flattened, extending along
Genus Liriope Lesson, 1843 almost entire length of radial canals; tentacles up
One species. to 640; 3-4 statocysts in each octant.
436 Hydromedusae
Genus Haliscera Vanhoffen, 1902 Genus Ptychogastria Allman, 1878

With 8 radial canals; with continuous row of tenta- Ptychogastriidae with tentacles in clusters, some with
cles; without papillae on exumbrella (Haliscera alba adhesive disks; manubrium with lateral lobes; with
Vanhoffen, 1902 is considered an invalid species, see eight mesenterial partitions; gonads on manubrial
Kramp, 1961). lobes or on radial canals adjacent to manubrial lobes.
1 Umbrella with fairly thin walls, apex evenly Ptychogastria polaris Allman, 1878 (Fig. 3.169).
rounded; gonads along proximal part of radial Umbrella 18 to 22 mm wide; hemispherical to
canals, close to manubrium. Umbrella 8 mm wide, conical; exumbrella with 16 radiating ridges; 16
4 mm high, hemispherical; 6 tentacles and 2 stato- gonads on sides of 8 manubrial lobes; 48 tentacle
cysts in each octant: Haliscera racovitzae (Maas, clusters each formed by 3 filiform tentacles and
1906) (Fig. 3.165) numerous adhesive ones; 16 statocysts.
la Umbrella with very thick apex, gonads separated
from manubrium.. .................................................2 Family Rhopalonematidae
2 Umbrella up to 18 mm, with very thick, bluntly Trachymedusae with narrow manubrium; with or
conical projection; 64-72 marginal tentacles in without peduncle; usually with 8, rarely more, radial
adults; 8-9 tentacles and 2 statocysts in each canals; without centripetal canals; marginal tentacles
octant; base of tentacles surrounded by broad evenly distributed, sometimes of two kinds, each ten-
thickening of marginal cnidocyst tissue; gonads tacle of uniform structure; gonads on radial canals or
oval, well separated from manubrium in middle hanging in pouches into subumbrellar cavity; free,
portion of 8 broad radial canals: Haliscera conica rarely enclosed, ecto-endodermal statocysts.
Vanhoffen, 1902 (Fig. 3.166) 1 Gonads are a continuous band around manubrium
2a Umbrella 15-17 mm wide, 9-10 mm high, almost extending outwards on radial canals:
hemispherical, with very thick, hemispherical Homoeonema
apex; about 96 marginal tentacles in adults; about l a Gonads isolated, on radial canal, sometimes adja-
12 marginal tentacles and 3 statocysts in each cent to manubrium ............................................. .2
octant; thickening of marginal cnidocyst tissue 2 Without gastric peduncle .................................... 3
less pronounced than in H. conica; gonads broad- 2a With gastric peduncle ..........................................9
ly oval, about 215 as long as radial canals, located 3 With 4 gonads, pendent; 4 large and 24 small ten-
slightly nearer manubrium than bell margin: tacles: Tetrorchis
Haliscera bigelowi Kramp, 1947 (Fig. 3.167) 3a With 8 (rarely more) gonads ...............................4
4 With 8 long, club-shaped, and up to 24 small, cir-
Genus Halitrephes Bigelow, 1909 rus-like, tentacles; gonads elongated along radial
With 16 or more radial canals; with continuous row of canals: Rhopalonema
tentacles; without papillae on exumbrella. One species. 4a All tentacles of one kind .....................................5
Halitrephes maasi Bigelow, 1909 (Fig. 3.168). 5 Gonads on or adjacent to manubrium (sometimes
Umbrella 100 mm wide, mesoglea flaccid; 16-30 also 8 gonads free from manubrium); very nume-
broad, ribbon-like radial canals, some bifurcated; rous tentacles: Arctapodema
100-300 marginal tentacles; gonads and number of 5a Gonads separated from manubrium ....................6
statocysts unknown. 6 Exumbrella with numerous meridian furrows;
gonads sausage-shaped, pendent; very numerous
Family Ptychogastriidae tentacles: Crossota
Trachymedusae with either simple manubrium with- 6a Exumbrella smooth; gonads not pendent.. ......... .7
out mesenteries, or with eight-lobed manubrium, with 7 With 8 tentacles; gonads globular, distal:
eight mesenterial partitions; with marginal tentacles Sminthea
grouped into more or less well defined clusters, some 7a Gonads linear ...................................................... 8
with adhesive disks or with very numerous tentacles, 8 Up to 32 tentacles developing in succession:
not in clusters but inserted at various levels of exum- Colobonema
brella; no centripetal canals or peduncle; with 8 radial 8a With 48 or more tentacles of equal size:
canals; gonads either attached onto manubrium, on Pantachogon
sides of 8 manubrial lobes, or on radial canals adjacent 9 Peduncle short, conical (in young specimens
to manubrial lobe; free ecto-endodermal statocysts. almost invisible) .............................................. 1 0
Hydromedusae
9a Peduncle long, slender ......................................

11 soidal, on distal part of radial canals, unequal
10 With 2 pendent gonads: Persa alternatively 4 larger and 4 smaller; umbrella 4-6
10a With 8 globular gonads: Amphogona mrn wide, lower than hemispheric; mesoglea thin;
11 Gonads linear, on peduncle only: Ransonia manubrium small; small, broad peduncle; 4 short,
11a Gonads sausage shaped, pendent ......................12 simple lips; 5Q-80tentacles; 6-24 statocysts.
12 Gonads attached to peduncle: Aglaura
12a Gonads attached to subumbrellar portion of radial Genus Arctapodema Dall, 1907
canals: Aglantha Rhopalonernatidae without gastric peduncle; with
gonads on manubrium or on radial canals adjacent to
Genus Aglantha Haeckel, 1879 manubrium; with 8 narrow radial canals; numerous
Rhopalonernatidae with long slender gastric pedun- tentacles, all alike, in a single row; free statocysts.
cle; with 8 pendent, sausage-shaped gonads on sub- Arctapodema antarctica (Vanhoffen, 1912) (Fig.
umbrellar portions of 8 radial canals; numerous tenta- 3.176). With 4 interradial gonads encircling manu-
cles, all alike, with free statocysts. brium but interrupted in perradial comers; no go-
Aglantha digitale (Miiller, 1776) (Fig. 3.170). nads on radial canals. Umbrella up to 16 mm wide;
Gonads close to base of peduncle. Umbrella 10-40 manubrium simple, tubular; about 120 tentacles;
mm high, thimble-shaped, subumbrellar muscles statocysts unknown.
strong; with small apical projection; manubrium Arctapodema ampla (Vanhoffen, 1902) (Fig.
small; mouth with 4 simple lips; 80 or more tenta- 3.175). With 8 gonads extending from radial lobes
cles; 8 statocysts. of manubrium outwards onto proximal parts of
Aglantha elata (Haeckel, 1879) (Fig. 3.171). go- radial canals; sometimes also 8 pairs of small go-
nads about midway between peduncle and bell nads on radial canals, separated from manubrium.
margin. Umbrella 10-12 rnm high, narrow, with Umbrella up to 15 mm wide; manubrium with 8
pointed apical projection; 40-48 tentacles; 16 sta- radial lobes; about 100 tentacles; 4-8 statocysts.
tocysts.
Genus Colobonema Vanhoffen, 1902
Genus Aglaura Pkron and Lesueur, 1810 Rhopalonernatidae without gastric peduncle; with api-
Rhopalonernatidae with slender gastric peduncle; with cal outlines of subumbrellar muscular fields forming a
8 sausage-shaped gonads on peduncle, not on subum- star-shaped figure; elongate gonads along 8 radial
brella; numerous tentacles, all alike; free statocysts. canals; tentacles all of one kind, developing in succes-
Aglaura hemistoma Peron and Lesueur, 1810 (Fig. sion; free statocysts.
3.172). Umbrella 4-6 mm high, 3-4 mm wide; apex Colobonema sericeum Vanhoffen, 1902 (Fig.
flat; 48-85 tentacles; 8 statocysts. 3.177). Umbrella up to 45 mm wide, 35 mm high,
slightly conical; manubrium short, tubular; gonads
Genus Amphogona Browne, 1905 linear, along greater part of radial canals; 32 tenta-
Rhopalonernatidae with short conical gastric peduncles, adradial tentacles develop before interradial;
cle, exumbrella smooth; with ellipsoidal or sac- statocysts probably alternating with tentacles.
shaped, pendent gonads on the 8 radial canals; gonads
usually of unequal size; with tentacles all alike, not Genus Crossota Vanhoffen, 1902
densely crowded; with free statocysts. Rhopalonematidae without peduncle; numerous
Amphogona apicata Kramp, 1957 (Fig. 3.173). meridian furrows on exumbrella; 8 or more radial
With a blunt, conical apical projection; gonads canals; pendent sausage-shaped gonads on radial
pendent in the middle part of the radial canals; canals; numerous densely crowded tentacles, all alike;
umbrella up to 7 mm wide and 8 mm high, dome- free statocysts.
shaped with thin walls; a small, broad peduncle; Crossota alba Bigelow, 1913 (Fig. 3.178). Gonads
manubrium small, tubular, about 112 as long as the on 8 radial canals, nearer circular canal than
subumbrellar cavity; 4 short, simple lips; about 64 manubrium. Umbrella up to 42 mm wide, 10 mm
tentacles; statocysts unknown. high; colourless; with 8 sharp longitudinal ridges
Amphogona apsteini (Vanhoffen, 1902) (Fig. separated by 8 broad, flat, furrows; up to 190 ten-
3.174). Without apical projection; gonads ellip- tacles; number of statocysts unknown.
43 8 Hydromedusae
Crossota brunnea Vanhoffen, 1902 (Fig. 3.179). tubular, elongate, mouth with four small, broadly
Gonads on 8 radial canals near base of manu- rounded lips; up to 48 tentacles; 8 statocysts.
brium. Umbrella up to 30 mm wide, 22 mm high;
colour brown; with 8 large, deep longitudinal fis- Genus Ransonia Kramp, 1947
sures and above them 8 similar small invagina- Rhopalonematidae with high conical umbrella (simi-
tions; 600 or more tentacles, very densely crow- lar to Aglantha); with long and narrow gastric pedun-
ded; number of statocysts unknown. cle; 8 radial canals; linear, discontinuous, gonads
along peduncular portions of radial canals, not on sub-
Genus Homoeonema (Maas, 1893). Browne, 1903 umbrella; numerous tentacles, all alike; statocysts
Rhopalonematidae without gastric peduncle; gonads unknown.
forming a continuous band around base of manubrium Ransonia krampi (Ranson, 1932) (Fig. 3.185).
and extending outwards along proximal half of 8 Umbrella 8 mm wide, 15 mm high, with apical
radial canals; numerous tentacles, all alike; vesicular projection; about 88 marginal tentacles.
statocysts.
Homoeonema platygonon Browne, 1903 (Fig. Genus Rhopalonema Gegenbaur, 1857
3.180). Umbrella 1-2 mm wide and high; about 80 Rhopalonematidae without gastric peduncle; with
or more tentacles; 4 statocysts. gonads along radial canals, separated from manubri-
um; tentacles of two kinds: 8 large perradial tentacles
Genus Pantachogon Maas, 1893 with swollen ends, and 16 inter- and adradial short,
Rhopalonematidae without gastric peduncle; apical stiff, cirrus-like tentacles with swollen ends; enclosed
outlines of subumbrellar muscular fields forming statocysts.
entire circle; gonads on 8 radial canals separated from Rhopalonema velatum Gegenbaur, 1857 (Fig.
manubrium; 48 or more tentacles, all alike; free mar- 3.186). Umbrella 8-10 mm wide, with conical api-
ginal statocysts. cal thickening; gonads linear or oval, on middle
1 Umbrella mitre-shaped, with gelatinous apical pro- third of 8 radial canals; one statocyst next to each
jection; 48 tentacles. Umbrella 7-10 mm wide, 6 perradial tentacle and interradial cirrus.
mm high; gonads lancet-shaped, on distal half of 8
radial canals; 4 (or more) statocysts: Pantachogon Genus Sminthea Gegenbaur, 1857
militare (Maas, 1893) (Fig. 3.18 1) Rhopalonematidae without gastric peduncle; with
1a Umbrella without apical projection; 64- 120 tenta- globular gonads on 8 radial canals; 8 perradial tenta-
cles ........................ ................................................ 2 cles; enclosed statocysts.
2 With 64 tentacles. Umbrella about 12 mm wide Sminthea eurygaster Gegenbaur, 1857 (Fig.
and high, with very strong and conspicuous muscu- 3.187). Umbrella up to 6 rnm wide and about half as
lature; gonads along greater part of 8 radial canals; high, with small apical projection; gonads globular
64 tentacles; 64 free statocysts: Pantachogon on radial canals close to circular canal: 8 statocysts.
haeckeli Maas, 1893 (Fig. 3.182)
2a With 120 tentacles. Umbrella about 4 mm wide, a Genus Tetrorchis Bigelow, 1909
little broader than high; with strong musculature; Rhopalonernatidae without gastric peduncle; with 4
gonads linear, extending along proximal two thirds sausage-shaped, pendent, gonads attached to 4 of 8
of 8 radial canals; number of statocysts unknown: radial canals near middle points; 4 large and several
Pantachogon scotti Browne, 1910 (Fig. 3.183) small marginal tentacles.
Tetrorchis erythrogaster Bigelow, 1909 (Fig.
Genus Persa McCrady, 1859 3.188). Umbrella 10-12 mm wide, 8 mm high,
Rhopalonematidae with short gastric peduncle; 2 oval pyriform; manubrium tubular, brilliant carmine; 4
or sausage-shaped gonads, pendent, near middle point large perradial tentacles at level of 4 gonads bear-
of subumbrellar portions of two opposite radial ing radial canals and 16-24 small tentacles; stato-
canals; 8 radial canals; numerous long tentacles, all cysts unknown.
alike, each with terminal knob; free statocysts.
Persa incolorata McCrady, 1859 (Fig. 3.184).
Umbrella 2 mm wide, 3 mm high; manubrium
Hydromedusae 439
Suggested readings Nicely illustrated catalogue of the Mediterranean

Capitate Hydroidomedusae.
Boero F., Bouillon J., Piraino S. In press. Kramp P.L. 1959. The Hydromedusae of the Atlantic
Heterochrony, generic distinction and phylogeny Ocean and adjacent waters. Dana-Rep., 46: 1-283.
in the Family Hydractiniidae (Hydrozoa, General review of Atlantic and Mediterranean
Cnidaria). Modern revision of the family hydromedusae.
Hydractiniidae. Kramp P.L. 1961. Synopsis of the medusae of the
Bouillon J . 1978. Hydromeduses de la Mer de Bismark world. J. Mar. Biol. Ass. U.K., 40: 1-469.
(Papouasie, Nouvelle Guinke). I. Anthomedusae Descriptions of all the marine and freshwater
Capitata (Hydrozoa-Cnidaria). Cah. Biol. Mar., medusae known up to 1961.
19:249-297. Kramp P.L. 1968. The hydromedusae of the Pacific
Bouillon J. 1978. Hydromkduses de la Mer de Bismark. and Indian Oceans. Sect. I1 and 111. Dana-Rep.,
(Papouasie, Nouvelle Guinke). 11. Lirnnomedusa, 72: 1-200. General review ofPacific hydromedusae.
Narcomedusa, Trachymedusa et Laingiomedusa Mayer A.G. 1910. Medusae of the world.
(sous-classe nov.). Cah. Biol. Mar., 19:473-483. Hydromedusae. Vol. I, 11, 111. Carnegie Inst.,
Bouillon J. 1980. Hydromeduses de la Mer de Washington, pp. 1-735. Very nicely illustrated
Bismark. (Papouasie, Nouvelle Guinee). 111. monograph on all the medusae known to that date.
Anthomedusae Filifera (Hydrozoa-Cnidaria). Cah. O'Sullivan D. 1984. A guide to the Hydromedusae of
Biol. Mar., 21:307-344. the Southern Ocean and adjacent waters. ANARE
Bouillon J. 1984. Hydromeduses de la Mer de Res. Notes, 5: 1- 135. Illustrated guide of Antarctic
Bismark.(Papouasie, Nouvelle Guinte). IV. and Subantarctic hydromedusae.
Leptomedusae (Hydrozoa-Cnidaria). Indo- Pagks F., Gili J.-M., Bouillon J. 1992. Medusae
Malayan Zool., 1:25-112. (Hydrozoa, Scyphozoa, Cubozoa) of the Benguela
The above 4 papers contain a review of the most Current (Southeasthern Atlantic). Scientia Marina,
recently described Pacific hydromedusae. 56(Supl. 1):l-64. Recent review of the eastern
Bouillon J. 1985. Essai de classification des South Attantic hydromedusae.
Hydropolypes - Hydromeduses (Hydrozoa- Ramirez F., Zamponi M. 1981. Hydromedusae. In
Cnidaria). Indo-Malayan Zool., 2:29-243. Major "Atlas del Zooplancton del Atlantico
contribution toward a new classzjication o f the Sudoccidental y metodos de trabajo con el zoo-
Hvdroidomedusae. plancton marino" (D. Boltovskoy, ed.), Public.
Bouillon J. 1995. Classe des Hydrozoaires. In "Traite Esp. Inst. Nac. Inv. Desarrollo Pesq., Mar del
de Zoologie", 3(2) (P.P. Grasse, D. Doumenc, Plata, pp. 443-469. General review of western
eds.), Masson, Paris, pp. 29-416. A comprehensive South Atlantic hydromedusae.
treatise covering morphology, histology and Russell F.S. 1953. The medusae of the British Isles.
systematics of Hydroidomedusae. Anthomedusae, Leptomedusae, Limnomedusae,
Brinckmann-Voss A. 1970. AnthomedusaeiAthecatae Trachymedusae and Narcomedusae. Cambridge
(Hydrozoa, Cnidaria) of the Mediterranean. Part. Univ. Press, London, pp. 1-530. Nicely illustrated
I: Capitata. Fauna Flora Golfes Neapel, 39: 1-96. monograph on the medusae arot~ndthe British Isles.
440 Hydromedusae
Bougainvillia frondosa
Bougainvillia macloviana
Bougainvillia carolinensis
Bougainvillia niobe
Koellikerina fasciculata
Hydromedusae
Thamnostoma
@ tetrella
!
@ cytaeis rp.
Turritopsis nutricula
1@ ~ ~ d r l c t i ntenuis
ia Hydractinia tournieri Hydractinia
minima
Hydromedusae
@ Hydractinia mhuta 1 Rathkea formosissima
@ Rathkea africana
part of bell margin

showing
gelatinous warts
Bythotiara murrayi
Bythotiara capensis
I medusa
I
1 " f Calycopsis
I@ ~ a ~ y c o p stypa
is papillata
Hydromedusae 443
Heterotiara anonyma 1
Niobia dendrotentaculata 1@
L----
Sibogita geometrica
Pseudotiara
tropic.
Hydromedusae
Amphinema rubra
a Leuckartiara octona
b J
1 1 Cirrhitiara superba
@ Halitholus intermedius Annatiara affinis
Hydromedusae 445
446 Hydromedusae
Moerisia inkermanica a \
Tiaricodon coeruleus 1
@ Russellia mirabilis
a Dipurena reesi @ Dipurena haltera fa 1 a Dipurena baukalion

Hydromedusae
@ Sarsia producta
Euphysora gigantea
1 Euphysora gracilis
Hydromedusae
Vannuccia forbes,,
Staurocladia capensis
...
a I,
Staurocladia vallentini
@ Euphysa ua>
Margelopsis australis
Hydromedusae
450 Hydromedusae
Hydromedusae
Cirrholovenia \ ( bell margin

@ Blackfordia virginica tetranema Eirene viridula
452 Hydromedusae
Eucheilota duodecimalis a lrenium teuscheri

Hydromedusae 453
Hydromedusae
I
lovenella cirrata
Staurophora Toxorchispolynema
mertensii
6D
Malagazzia
\ carolinae
\ '%
i 1,
@ Octocanna haeckeli
Octophialucium bigelowi
1 m Tetracanna octonema Cosmetirella da visi

Hydromedusae 455
r- I
1- Halopsis ocellafa
1 Mitrocomella millardae 1 Orchistoma collapra

Hydromedusae
Hydromedusae 457
Hydromedusae
Gossea brachymera
1 Olindias phosphor~a
Vallentinia
falklandica
Hydromedusae
--
Cunina octonaria
Hydromedusae
subumbrella h 1 exumbrella
Solmiss~~s
atlantica
U
ecto-
endodermic
_- Solmissus marshalli
velum
otoporpae
/
marginal lappets
Pegantha
rubuginosa
Pegantha martagon
Hydromedusae 46 1
I 1
solmaris flavescens
Botrynema brucei
a Haliscera racovitzae
Halicreas minima Haliscera conica

Hydromedusae
Hydromedusae
Crossota alba
Colobonema sericeum
Crossota brunnea
Homoeonema platygonon
/----
Pantachogon militare aD
- Pantachogon haeckeli
Hydromedusae
Pantachogon scotti
Persa incolorata
Rhopalonema velatum
Sminthea erygaster
Hydromedusae
4
Fig. 3. Species illustrations.
Figure sources: 154: from Bigelow (1909), redrawn; 3, 36: from Bouillon (1995), redrawn; 29: from Bouillon (1978), redrawn; 72b, 105: from
Bouillon (1984), redrawn; 73b: from Brinckmann-Voss (1970), redrawn; 8,89,97, 110: from Goy (1979), redrawn; 2,4-7,9-13, 15-19,22-26,
30-35,37,38,40-49,52-56,59-64,68-70,74,75,78,80,81,83, 85-88, 90-96,98, 101, 103, 104, 106, 107, 111, 112, 115, 116, 118-120, 124-
129, 131, 134-136, 138-140, 142-151, 153-158, 160-185, 187, 188: from Kramp (1959a), redrawn; 28, 58, 66, 84, 100, 102, 126: from Kramp
(1968a), redrawn; 132: from Moreira and Yamashita (1972), redrawn; 1, 3, 20, 21, 27, 39, 50, 51, 57, 71, 72a, 73a, 76, 77, 79, 82, 113, 116,
121, 123, 130, 133, 141, 142, 157, 158, 159, 160, 186: from Pages et al. (1992), redrawn; 137: from Rarnirez and Zamponi (1981), redrawn;
112: from Russell (1953), redrawn; 14: from Vanhijffen (191 I), redrawn; 99: from Vannucci (1957b ), redrawn; 108, 109: from Vannucci and
Moreira (1966), redrawn; 65, 67, 152: from Zamponi (1983).
Siphonophorae
Phil R. Pugh
Introduction Siphonophores are present throughout the World's

oceans, and throughout the water column down to a
Siphonophores are complex, highly polymorphic depth of at least 4500 m, although individual species
hydrozoan cnidarians, whose "colonies" are formed may have restricted latitudinal or depth ranges. The
by many medusoid and polypoid "individual s" that majority are truly oceanic, but a few species are neri-
function physiologically as a single entity. They occur tic, occurring in shallow inshore waters where the
in a bewildering variety of shapes and sizes, from salinity is not greatly reduced. In addition, in regions
about 1 mm to several tens of metres in length. of high salinity, such as the Red Sea, species diversi-
However, because of their fragility they are difficult to ty can be greatly reduced.
study both taxonomically and ecologically. The most
famous, and first to be described, siphonophore is the Unlike most other hydrozoan coelenterates, siphono-
Portuguese Man O'War, Physalia physalis (Linné, phores do not show an altemation of generations
1758), that floats at the surface of the ocean, with its between an attached, asexual polyp stage and a free-
stinging tentacles hanging down many metres into the swimming, sexual medusa. Instead, modified versions
water below. However, this species is exceptional, and ofboth stages are found together, attached to the stem
almost all other siphonophore species are permanent- of the free-floating animal. Totton (1965) considered
ly planktonic, exclusively marine animals. The only the fully grown siphonophore to be an enlarged, larval
other exception is a small group, of the family nurse carrier, which he called a paedophore. This does
Rhodaliidae, that have become benthic attaching not itself be come sexually mature, but buds off the
themselves to the sea-bed like tethered air-balloons. adult sexual medusoids, which may be released, along
with other stem components, to lead a separate life.
There are 3 basic types (orders) of siphonophores that Individual animal s of most species appear to produce
are classified according to whether an apical, gas-fil- medusoids of both sexes, but not necessarily simulta-
led float is present (Cystonectae and Physonectae) or neously. However, for a few species, such as the
absent (Calycophorae). Those with a float are sub- Portuguese Man O'War, each animal produces medu-
divided on the basis ofthe presence (Physonectae) or soids of only one sexo Fertilisation is external and
absence (Cystonectae) of a group of swimming bells development proceeds rapidly over a period of 2 to 3
immediately below the float. About two-thirds of the weeks. A larval nectophore and/or bracts are usually
ea. 150-160 currently recognised species belong to the developed first, and these can be morphologically dif-
Order Calycophorae, and these predominate in samp- ferent from those of the adult animal.
les collected by nets. However, from limited collec-
tions with sUOO1ersibles,about two-thirds of the spe- Little is known about the life span of an individual
cies and three-quarters of the specimens are siphonophore, but it is presumed that some of the
physonects. This is probably because most calyco- smaller species live for only a few months, while
phoran species are small and difficult to see, while some of the larger ones may survive for lOor more
many physonects are large and brightly coloured, and years. The abundance ofthe shorter-lived species thus
their spectacular appearance quickly draws one's varies seasonally, and the period of abundance varies
attention to them. Many of the species collected by from species to species. However, as Mackie el al.
submersible ha ve proved to be new to science. Thus, (1987) discuss, one cannot always as sume that these
even though net collections have already demonstra- variations in abundance are a true reflection of sea-
ted the importance of siphonophores in the marine sonal change, as temporal changes in their horizontal
ecosystem, the true significance of these animals can or vertical distribution may not have been observed.
only be appreciated fully by combining the data from
such collections with in situ observations. Much of the biology of siphonophores has been revie-
wed by Mackie el al. (1987). They are important pre-
Sout/z Atlantic Zooplankton.

edited by D. Boltovsko)', pp. 467-511
'id 1999 Backhu)'s Publis/zers, Leiden, T/ze Netherlands
•
468 Siphonophorae
dators and often occur in such numbers that they form prey item, within a certain size range, that they encoUll-
the dominant group of camivores (Pugh, 1984). They ter, some appear to be highly selective and feed only on
feed primarily on small crustaceans that they ensnare certain taxa (Purcell, 1981). The siphonophore species
by discharging their nematocysts. Some species, that have adopted these various feeding strategies can
however, feed on soft-bodied animals, such as small have different pattems of geographical and vertical dis-
fish, which they capture by injecting poi son from tribution. This has lead Pugh (1986, 1991) to specula-
penetrant nematocysts. Most of the nematocysts ate te that these different distributions should reflect that of
grouped on the animal's tentacles, which can extend their preferred prey items.
to considerable lengths, but other structures also may
possess them. The role of siphonophores in marine food chains is
unclear, but they do not appear to be a dead end bran-
Unlike many jellyfish, which actively pull or push ch. Despite their gelatinous nature, they are preyed
their tentacles through the water, siphonophores are upon by a variety of other organisms, particularly
passive feeders, setting tentacular nets that often form other gelatinous camivores such as medusae, cteno-
complex pattems. Biggs (1977) discussed some ofthe phores and pelagic molluscs. Several species of fish
methods that siphonophores use to capture their prey. also feed on them, as do turtles. In addition, siphono-
He noted that the fishing cycle consisted of two pha- phores have associations with a variety of organisms,
ses: a fishing period, when the tentacles are spread out of which hyperiid amphipods probably are the most
and the animal waits for a prey item to swim into its important. The nature of the association can vary from
net; and a swirnming phase, when the tentacles are simple phoresis, where the other animal simply is car-
retracted and at the end of which the fishing net is ried about, to total parasitism, as with some amphi-
reset. The whole process is highly coordinated and the pods whose juvenile stages develop and gradually
disposition of the tentacles is by no means a tangled consume parts of the siphonophore (see Harbison et
mess. The exact fishing posture depends on several al., 1977). Many of these amphipod associations are
factors, including the animal' s ability to remain neu- species specific, as is also that of the nudibranch,
trally buoyant, and the degree to which the stem can Cephalopyge trematoides, which feeds exclusively on
be extended. Some merely use an extended "long- Nanomia bijuga (Sentz-Braconnot and Carré, 1966).
line" posture, with the tentacles hanging down from Fish also are known to associated with siphonophores.
the buoyant stem. Species using this posture tend to The best known example is the association between
spend long periods fishing, and to capture larger prey the Man O'War fish, Nomeus gronovii, and the
items. Their "sit and wait" strategy, which expends Portuguese Man O'War (see Totton, 1960). In situ
little energy, relies on the greater swirnming speed of observations also have shown associations offish with
the larger prey to increase the chane es of an encoun- siphonophores, including Caristius sp. and a cystonect
ter. The active swirnmers usually set complex nets, species (Janssen et al., 1989). Various parasites have
and some use a "veronica" movement that results in been observed on and within siphonophores (see
the tentacles being spread out from the stem to form a Totton, 1965), but there is little detailed information.
helix of 2 or 3 tums (Mackie and Boag, 1963).
The three-dimensional disposition of the tentacles Methods

allows a greater volume of water to be searched for
prey. Madin (1988) has calculated that the encounter Most of our knowledge conceming the zoogeographi-
volume, within which the tentacles are disposed, may cal distribution of siphonophores has come from ani-
be 0.5 m3, and probably is even greater for some ofthe mals caught in a variety of nets, ranging from simple
large physonect species. Others "squid-jig", periodical- surface dip-nets to sophisticated deep-water trawls.
Iy contracting and relaxing an individual tentacle, or its However, the complexity and fragility of these ani-
side branches, to enhance prey capture. In addition, mals mean that the whole structure can easily be bro-
several species have evolved tentacular structures that ken apart, and typically this is what happens during
appear to mimic other organisms in order to lure in net collection. This also means that many pieces can
their prey (Purcell, 1980; Pugh, 1989). Although many be lost through the meshes so that there can be pro-
siphonophore species are generalist feeders, taking any blems in quantifying their abundance. This is particu-
Siphonophorae 469
larly the case for physonect siphonophores as many of Many supposedly rare, but actually very fragile si-
their potentially myriad parts can be lost. The number phonophore species have been collected by this
of floats, or pneumatophores, can give some indica- means, as well as several new species.
tion of the number of specimens collected but the phy-
sonect pneumatophore is relatively small and is rarely At deeper depths, submersibles with sophisticated coI-
retained in net samples. This problem is further dis- Iecting devices need to be used. Ones that have been
cussed by Pugh (1984). used extensively in the recent past are the Johnson-
Sea-Link I and II because of their ability to collect
The majority of siphonophore species belong to the many delicate specimens intact (see Youngbluth,
Order Calycophorae and these are the ones that are 1984), although they do have a depth limitation of ea.
most frequently caught by nets. These species do not 1000 m. Even so, some gelatinous species stilI cannot
possess a pneumatophore but, fortunately, most of be described because they simply disintegrate "before
them possess other unique "individual" parts that can your very eyes", probably as a result of turbulence
enable a reasonably accurate estimate oftheir abundan- produced by the submersible. Such in situ studies have
ce to be made. This applies to both the polygastric shown that even relatively Iarge physonect species can
(asexual) and eudoxid (sexual) stages. There are a few be quite abundant; for instance Nanomia cara, which
species, in the subfamily Prayinae, that develop two is about 20-40 cm in length, can occur at densities of
identical swimming bells or nectophores. In this case 7-8 per m3 (Rogers et al., 1978).
the numbers of nectophores need to be halved in order
to quantiry the number of specimens. Similarly, in the Personal observations (Pugh, 1989) have shown that
family Hippopodiidae multiple (up to ea. 15) necto- in contrast to nets, where calycophoran species predo-
phores are developed. In this case, the exact number per minate, over 60% of the siphonophore species and
animal can vary, and some of the les s well developed 70% ofthe specimens collected by submersible belon-
ones may be lost, and so quantification is more compli- ged to the order Physonectae. There is an obvious
cated. Again this is discussed further by Pugh (1984). reason for this, in that the physonects are generalIy
Iarger and more highly pigmented than the calyco-
Since nets do not sample all species adequately then phoran species and, thus, much easier to observe.
the importance of the total siphonophore population Nonetheless, it is aIso true that over half of the phy-
can be greatly underestimated. In general, net samples sonect, and a quarter of the calycophoran, species that
indicate that calycophoran siphonophores are more have been collected have proved to be new to science.
abundant in superficial waters and, in the open ocean, Again, species that had been thought to be rare are
may reach den sities of ea. 1 per m3, while in the top shown to be common. For instance, Pugh and
1000 m ofthe water column they average about 10 per Harbison (1986) found that Lychnagalma utriclllaria
1000 m3. However, in inshore waters concentrations (Claus, 1879) was the commonest physonect species
of a single small species can exceed 500 per m3 (e.g. colIected in the vicinity of the Bahamas, despite the
Greve, 1994). fact that there have been no other substantiated
records for this species since it was first described.
The best, and most exhilarating, way to study and col-
lect siphonophores is to use an in situ technique such When siphonophores have been collected in nets it is
as SCUBA diving or submersibles. It is the only way best to fix and preserve the whole sample as early as
by which the full beauty of these animals can be possible in 5% formalin buffered by borax. After a few
appreciated. In the case of SCUBA, the divers take days, the formalin should e changed, but it is best
down wide mouthed jars into which, with a lot of otherwise not to disturb the sample so as to alIow pro-
skill, the animals can be induced to swim. Shipboard per fixation ofthe animaIs. Further details can be found
experiments can then be carried out on the specimens, in the Ostracoda chapter (this volume). The siphono-
using large tanks. Nonetheless, siphonophores like phores inevitably will shrink, and volume loss is rapid,
their freedom of movement and so do not usually sur- with about a 50% reduction within the first 2 hours of
vive long under these conditions. "Blue water" preservation (Pugh, pers. obs.). However, most ofthis
SCUBA diving techniques have been described by is due to a loss of turgor so that the interstitial water
Hamner (1975) and Madin and Swanberg (1984). that previously was retained within the main cavity of
470 Siphonophorae
the nectophore, the nectosac, is lost. Nonetheless, and be a mutual exclusion between them, such that they
in contrast to a previous statement (Kirkpatrick and almost never occur in the same area at the same time
Pugh, 1984), it is best to leave the specimens to fix (Mackie el al., 1987). The oceanic species can be
completely before sorting them from the remainder of roughly divided into boreal, tropical and equatorial
the sample. As formalin is an unpleasant substance, species (Margulis, 1972), although such a scheme
during the sorting of our samples they are transferred does not take account of the depth distribution of
into Steedman's preserving fluid, which contains only some. Thus, some species are clearly restricted to
1% formalin (Steedman, 1976). Nonetheless, some warm, superficial waters, while the deeper-living,
species are so delicate that they disintegrate when pre- bathypelagic species can occur at almost any latitude
served in formalin. In such cases better success has as the temperature at these depths does not vary gre-
been achieved using ea. 3% glutaraldehyde. Alcohol is atly. The zoogeographic distribution of siphonophores
oot recornmended for siphonophores. in the North Atlantic Ocean has been addressed by
Pugh (1977) and Mackie el al. (1987). These studies
It is very difficult to preserve siphonophore specimens • indicate that the species composition of a siphono-
intacto The suggested method is first to anaesthetise the phore population is influenced by the various water
specimen by the slow, dropwise, addition of an isotonic masses presento For instance, there was a clearly defi-
magnesium chloride solution to the sea water, followed ned and very abundant population of siphonophores
by a further dropwise addition of 10-20% formal in. above the permanent thermocline present at lower
latitudes. In addition, different populations were
Once preserved and sorted, the parts of the siphono- found in North and South Atlantic Central Waters. It
phore specimens can be examined under a low mag- is most likely that the sarne situation will pertain in
nification binocular microscope, preferably using the various water masses of the South Atlantic, but
dark background illumination. Many of the compo- there are too few data to demonstrate this. Although
nent parts of a siphonophore species have characteris- there are clear correlations between hydrographical
tic morphologies and, with a practised eye, can be features and siphonophore populations, it is probable
readily identified. They can also be stained using a that this is an indication of other factors that have a
variety of stains, for instance Steedman's Triple Stain more immediate impact on their distribution, such as
or Borax Carmine. Staining can be useful if dark dietary preferences (Mackie et al., 1987).
background illumination is not available, or when
there is a difficulty in identifying the specimen, as the In the North Atlantic maximal species diversity was
stain will help to show up certain critical morphologi- reached at ea. 200N (Mackie el al., 1987) with a slow
cal features. de crease in numbers toward the equator, and a more
rapid one toward higher latitudes. The latter was main-
Iy the result of the disappearance of epipelagic species
Geographical and vertical distribution living above the permanent thermocline. Nonetheless,
there appear to be more species in Antarctic waters than
There are very few, if any, wide ranging studies of the there are in Arctic ones; and several ofthose species are
distribution of siphonophores in the South Atlantic indigenous to that region, for example Pyrostephos
Ocean. The data presented in Table 1 are, thus, based vanhoe./Jeni and Diphyes antarctica. There are also one
on the sparse records in the literature. Only the maxi- or two bipolar species that have never been recorded at
mal latitudinal limits of distribution are given, al- lower latitudes in the Atlantic, these being MlIggiaea
though this may mask some of the finer details, such bargmannae and possibly Gilia reticlllata.
as the presence of certain species in certain water
masses. The data on the main vertical range for each There is also a change in species diversity in the ver-
species is largely based on data in the Southampton tical plane, with fewer species being found in the col-
Oceanography Center database. der, deeper waters. Individual species can occur over
quite wide depth ranges but, in general, they can be
Most siphonophore species are truly oceanic and rare- classified as epipelagic (0-250 m), mesopelagic (250-
Iy appear inshore A few are mainly neritic, such as 1000 m) or bathypelagic (> 1000 m). However, sever-
MlIggiaea atlantica andM. kochi, and there appears to al species have a relatively restricted distribution with-
60QN-37Q5
r -L1 5W
3W
-J7QN- range
60QN-42Q5
60QN-49Q5
42QN-39Q5
42QN-l0Q5
55QN-38Q5
?33Q5-67Q5
39QN-18º5
61
62QN-45Q5
35ºN-31
53QN-33º5
73QN-53Q5
55QN-59Q5
50-65Q5
52-54Q5
QN-44Q5
500-1000
700-1000
700-1400
300-1000
>900
300-600
200-500
35QN-35Q5
5W 0-400*
0-300*
0-300
55QN-56Q5
Africa
Africa
Main 0-100
0-500
0-200
500-1000 ?º5
Latitudinal
Africa
Pleuston
0-500
0-200
(1) 0-100
(1 0-2°L
(3)
(1)
range
45Q5 )
depth 0-200
0-400
0-100 17
21
10
(m) 2.5
1.5
32
25
20
10
13
15
12
1
98
292 -~ ~5W
7
Siphonophorae 37-65Q5
55QN-
50QN-44º5
5W
18QN-36Q5
30QN-19º5
57QN-38Q5
46QN-38Q5
Africa
40º5
40QN-40Q5
58Q5
Africa(1)(1) t 471
rum --¡-
ophore
e
Abyla bicarinata
Apolemia
Physalia physalis
5Wuvaria
0-200*
Africa 0-200
(1)1 J
-----J 0-200
0-20~-i 0-100
500-1000
~ I l8'N-18'S
j 38'N-S9'S
ristata
-+---
_1
-.--- - -l
~+-- 472
>1000 7
36
11
-30
16
(mm)
20_4318
Ma~depthl
15
61
(m)
1 ttr I
42-67º5
-!
J60ºN-33º5
44ºN-40º5
60ºN-67º5
60ºN-63º5
44ºN-33º5
53ºN-34º5
55ºN-37º5
54ºN-38º5
44ºN-
60ºN-60º5
60ºN-66º5
47ºN-43º5
60ºN-57º5
55ºN-39º5
74ºN-67º5
60ºN-42º5
60ºN-33º5
60ºN~63º5
60ºN-65º5
62ºN-48º5
18ºN-57º5
44ºN-2º5
5W59-62º5
50ºN~67º5
47N-67º5
55ºN-8º5
-
1800
500-1000+
400-700
300-800+
500-1500
600-1000
200-1000
Africa
400-600
300-700
69ºN-59º5
35-40º50-200
0-250*
0-200*
0-200
0-100
40º5
range
60ºN-67º5
ºN-45º5
200-500
I60ºN-65º5
60ºN-67º5
-;-
18ºN-39º5
55ºN-39º5
47ºN-45º5
5WI Africa
--+ 60ºN-62º5
5-30º5
0-100
0-200
O~
?(2)
T¡
range
>1000
50º5-67º50-300
0-300* ~ I
~ -.-
II
100 I¡ 400-1000
500-800
100-500*
0-250*
0-600
0-200
500-900
~II[
200-1600
---l
0-600
0-200
---+
200-500
700-1100
-t1800-1500
10
21
35
15
12
20
14
4
95
87
61
JI
j
-+--
Siphonophorae
Height of 'latitudinal
Siphonophorae 473
Height of latitudinal Main depth

nectophore range range
(mm) (m)
Muggiaea kochi 7 48QN-36QS 0-100

Nectadamas diomedeae 60 60QN-62QS 500-1000
NectQpyramis natans 40 44QN-65QS 400-800

Nectopyramis thetis 20 63QN-34QS 300-600
Praya dubia 100 60QN-44QS 100-600
Praya reticulata 60 SW Africa (1) 100-200
Rosacea cymbiformis 30 SW Africa (1), 100-200
Rosacea plicata 30 I
60QN-65QS 200-500
Sphaeronectes gracilis 8
Sulculeolaria biloba 26
Sulculeolaria chuni 8 42ºN-32ºS 0-100..j
Sulculeolaria monoica 18 40QN-29QS 0-100
jSW Africa (1) 0-100 --t
Sulculeolaria quadrivalvis + 20 40QN-31

50QN-41 QS
QS !
1 ~:- 0-100
100
Sulculeolaria turgida 16 43QN-39QS 0-200
Vogtia glabra 30 62QN-56QS
--r-
100-500*
Vogtia pentacantha 40 SW Africa (1) 300-500
Vogtia serrata 40 66QN-65QS 400-800
Vogtia spinosa 30 69QN-59QS 100-300
Table 1. Distribution of siphonophore species fOllnd in the SOllth Atlantic. Height refers to the anterior nectophore in the calycophoran fami-
líes Diphyidae, Clausophyidae and Abylidae. Remarks: (1) Also uncommon in North Atlanlic and/or Medilerranean; (2) Bipolar species also
fOllnd at similar latitudes in North Allantic; (3) Other records trealed as dOllbtful as this is an Indo-Pacific neritic species; * Species known lO
llndergo small-scale diel vertical migrations.
in the mesopelagic zone. In the North Atlantic there Several epipelagic siphonophore species undergo diel
was a marked discontinuity, at ca. 42°N, between the vertical migrations over a depth range ofup to 200-250
more southerly and more northerly siphonophore m (Table 1). However, such migrations may not always
populations in the top 1000 m of the water column. be apparent because, in some species the rate of depth
Although the number of species present on either side change is relatively slow so that there is a sinusoidal
of this discontinuity was very similar, there was a wave of depth change throughout the day and night
change over in the specific population and a dramatic (Mackie et al., 1987). In addition, it may be that the
increase in biovolume to the north, withjust a few spe- depth range of sampling is too coarse to establish any
cies predominating. To the south the siphonophore small-scale diel vertical rnigrations (Pugh, 1984). The
population consisted mainly of small epipelagic spe- factors that might initiate these migrations, and the pro-
cies (chiefly in the families Diphyidae and Abylidae). cesses involved in them are discussed by Pugh (1977,
To the north the deeper living but larger species ofthe 1984). It is presumed that siphonophores undertake such
families Prayidae, Hippopodiidae and Clausophyidae migrations in order to remain in the vicinity of their
became predominant. It is probable that similar chan- prey, which often migrate to superficial waters at night.
ges will occur across the various major fronts, fór Physonect species, with a gas-filled float, are known to
instance the Subtropical Convergence and the Polar be important contributors to deep-scattering layers, and
Front (=Antarctic Convergence), in the South Atlantic; some have a pore to allow release of gas from their float
but again there are no detailed studies. that may facilitate a diel vertical migration.
474 Siphonophorae
Taxonomy phorae, they may also contain large amounts of mes-

ogloea, which contributes to the buoyancy of the ani-
Morphology mal. New or replacement nectophores are developed
at the apex of the nectosome.
As siphonophores are comprised of many types of
structures ("individual s"), each with a different func- The morphology of the nectophores is important for
tion and some found only in these animal s, a unique specific identification. The nectophores of physonect
terminology for these structures has been developed. siphonophores generally conform to a basic design
This can appear somewhat daunting to the non-spe- (Fig. ID, E). They do not possess a somatocyst (see
cialist and necessitates a detailed introduction to it as below) and, with a practised eye, easily can be distin-
it is critical to the identification of the species. In the guished from calycophoran nectophores. The impor-
following section the important terms and features are tant taxonomic features of the physonect nectophores
emboldened. are:
(a) their general shape;
Siphonophores are cnidarians, and so basically consist (b) whether the apico-Iateral ridges have a notch or
of two cell layers, the outer ectoderm and the inner divide toward the abaxial (furthest from the stem)
endoderm, with the latter forming the lining to the gas- end of the nectophore;
tro-vascular cavity. These two layers are separated by (c) the course of the 4 (dorsal, ventral and 2 laterals)
a more or less thickened, amorphous mesogloea. radial canals on the nectosac. The nectosac is
equivalent to the sub-umbrella cavity of a medusa
As noted above, the species in the 3 basic divisions and, by means of contractions of its muscular
(orders) are primarily distinguished on the basis ofthe walls, water is expelled through the narrowed,
presence or absence of either an apical, gas-filled basal opening, or ostium, so that the animal is pro-
float, the pneumatophore, or of swimming bells, pelled by jet propulsion;
nectophores. The latter, numbering from one to fifty (d) the shape of the apico-Iateral processes, or axial
or more, usually are grouped together at or toward the wings, which extend around the stem;
apex of the animal to form the nectosome. The pneu- (e) the structure of the thrust block, which abuts the
matophore (Fig. lA) is produced, during develop- stem and through which passes the gastro-vascular
ment, by an invagination at the aboral end of the larva, canal; and
- which results in its cavity being lined by ectodermal (f) the presence or absence of ridges on the sides of
cells. At the base of the cavity, there is a specialised the nectophore. In this paper two types of these
are a of cells, the gas-gland or pneumadenia, that ridges will be referred to, in continuation of the
secretes the gas to inflate the cavity. In some cysto- scheme adopted by Pugh and Youngbluth (1988).
nect siphonophores this gland may bear processes, Firstly, there are vertical lateral ridges, which are
hypocystic villi, while in the benthic rhodaliids it is closer to the axis than any others, and run from the
greatly expanded to form an aurophore. apico-Iaterals to or toward the infra-Iaterals. The
course of this pair of ridges is not always vertical,
Most siphonophores possess a long stem to which are and in two species, at least, more than one pair is
attached the various "individual s". The latter usually present. Secondly, lateral ridges that usually run
are organised into two zones, the nectosome and sip- from the apico-Iaterals to the ostium. There are
hosome, although only the latter is present in the specific variations in their arrangement.
Cystonectae.
The nectophores of calycophoran siphonophores
Nectosome. The nectosome, which lies immediately generally are more specialised, but there is a great
below the pneumatophore (Physonectae) (Fig. lA) or deal ofvariability amongst the various families and it
is apical (Calycophorae) (Fig. 2), bears, with the is difficult to give an overall picture. Six different
exception of one family, only asexual, medusoid nec- types are illustrated in Fig. 2. The first type (Fig. 2A,
tophores. These act in a co-ordinated way to propel 3.31-3.43) is basically a smooth, rounded structure
the animal through the water. Within the Calyco- containing large volumes of mesogloea, giving buoy-
Siphonophorae 475
r---------
I pneumatophore DISTAl PART OF
I
I ~pneumadenia SIPHOSOME
I
Q)
E
o
o
<J>
U
Q) nectophore
e
I
I
I
I
I
I
~-------- bract---
I
I
I
I
I
I
I
I
I
I tentacle
I
I
I
I tentillum
I
I
I
I
Q)
E
O
<J>
O bract
..r::.
CL
<J>
(i
TENTlllUM
I
I
I
I
I
I
I NECTOPHORE
I
I vertical
I lateral
1 _
ridge
lateral
4)
upper
view
view
Fig. 1. General structure of a physonect siphonophore. A, B, D, E after Totton (1965).

476 Siphonophorae
ancy to the animal. The extent of the nectosac can ventral surface of the nectophore, where it forms the
vary considerably between species. This type of nec- basal facet, also can be of taxonomic importance.
tophore is found in species of the families Prayidae
and Sphaeronectidae. In the latter family only a sing- The structure of the hydroecium is only of taxonomic
le nectophore, the larval one, is developed and is importance in calycophoran siphonophores (Fig. 2A,
retained by the adult animal. In the former the larval C). In species with stream-lined nectophores, the
nectophore may be retained in the adult, and a second, hydroecium ofthe anterior nectophore may be greatly
greatly reduced nectophore developed in addition; or retiuced or absent. However, in the posterior necto-
the larval nectophore can be autotomised and replaced phore it usually extends the full length of the,necto-
by one or, more usually, two adult or definitive nec- phore and the siphosomal stem may be partially or
tophores. Further definitive nectophores may be deve- wholly retracted into it for protection and to reduce
loped in some species. When two definitive nectop- drag during locomotion. In the genus Sulculeolaria
hores are present, they are identical in design and the nectosac of the anterior nectophore may have
form an apposed pair. additional commissural canals connecting the laterals
to the dorsal radial canal (Fig. 2C).
The second type of nectophore (Fig. 2B, 3.55-3.59),
found in the family Hippopodiidae, is flattened in the In calycophoran siphonophores a somatocyst líes
axial-abaxial direction and usually bears spines or above the apical wall of the hydroecium. This may be
protuberances. The nectosac is widely open, but shal- a relatively simple tube, as in certain species of the
low, giving the impression that the animals must be family Prayidae, forming part of the gastro-vascular
weak swirnmers. Gn the ventral radial canal of the canal system, and connecting with the radial canals on
nectosac there is a rete mirabile, a dilation of the the nectosac via a pedicular canal (Fig. 2A). lt may
ventral canal, which is particularly obvious in youn- have basal (descending) or apical (ascending) exten-
ger nectophores. Up to fifteen nectophores may be sions, or branches. In the other calycophoran fami-
developed, which have a tight, biserial arrangement. lies, the somatocyst tends to form a caecal extension
to the gastro-vascular canal, oftaxonomically variable
In the remaining calycophoran families the nectopho- shape and design (Fig. 2C). In those species with nec-
res are more streamlined, and the animals themselves tophores designated anterior and posterior the soma-
usually are active and rapid swirnmers. Generally a lar- tocyst is present only in the anterior one, except for
val nectophore is developed, which is then replaced by the family Clausophyidae where it is present in both.
one or, more cornmonly, two definitive nectophores. The morphological features of importance for the
However, it is believed that the larval one may be retai- taxonomy of calycophoran nectophores are:
ned in certain species. When two nectophores are pre- (a) their basic shape and design;
sent, they differ in structure and do not form an appo- (b) the presence and pattern of ridges; and
sed pair, rather one is wholly (Diphyidae and (c) the structure of the somatocyst.
Abylidae, Fig. 2C, D, F) or partially (Clausophyidae,
Fig. 2E) superimposed above the other. The upper, Siphosome. The siphosome generally is much longer
anterior nectophore, often has a pointed apex, while than the nectosome, and in some physonect species
the lower, posterior nectophore often is truncated api- can extend to several tens of metres. lts zone of pro-
cally so as to fit snugly with the other. These necto- liferation lies irnmediately below the nectosome. The
phores frequently bear patterns of external ridges and siphosome bears a succession of different polypoid
teeth, which are useful diagnostic features. Most of the and medusoid structures, with different functions,
ridges ron longitudinally, from the apex to the base of arranged into a succession of groups or cormidia
the nectophore, but are not necessarily complete. In (Fig. 2A). Each cormidium consists of a single gast-
some species a transverse velar ridge, Iying close to rozooid, the digestive organ or stomach, to the base of
the ostium, is presento In addition, many species have a which is attached a tentacle. The tentacle bears nume-
basal la mella or mouth plate that extends below the rous nematocysts, usually on side branches called
ostium, the opening ofthe nectosac, on its ventral side. tentilla (Fig. 1C), that are discharged on contact with
The angle at which the mouth plate is joined to the a prey item and either paralyse or ensnare it. Many
S'lphonophorae 477
PRAYIDAE
HIPPOPODIIDAE
anterior Subfa m.
DIPHYIDAE
Sulculeolar" IInae
nectophore
radial
canal
posterior
nectophore
basal tenti Ila

DIPHYIDAE
lamellae
Subfam. D'•p hymae
.
478 Siphonophorae
4) l/V J/ElfJ\ \ 4)
ClAUSOPHYI DAE
ABYlIDAE
Fig. 2. General slructure of calycophoran siphonophores from various families. A: after Tollon (1965); C: after Carré (I979).
large nematocysts are often arranged into a cnido- the other cormidial elements. Often they contain large
band that may be covered by an involucrum. amounts of mesogloea and play an important role in
Frequently a terminal filament is present. The struc- flotation. They are absent in cystonects, while in phy-
ture ofthe tentillum can be useful in identifying some sonects each cormidium usually contains several
species, particularly physonect ones. bracts, which are leaf-like with a simple bracteal
canal. In calycophores each cormidium has a single
The cormidia of physonects usually inelude a number bract, except in the family Hippopodiidae where it is
of palpons, which are reduced gastrozooids and bear absent. The bract may have a complex structure, with
a small tentaele or palpacle (Fig. lB). The palpaele a divided canal system or a phyllocyst (equivalent to
probably has a sensory function, while the palpon aids the somatocyst of the nectophore), whose arrange-
in digestion, and accumulates and disposes of waste ment is of taxonomic importance. The bracts of some
products. Palpons usually are attached directly to the species are rounded, while others are angular, conical
stem, but a specialised type, the gonopalpon, may be or helmet-shaped. The latter types usually possess a
budded from a gonodendron. The latter itself may be basal process called the neck shield. Those calyco-
a specialised palpon (Totton, 1965). The gonodendron phores with stream-lined nectophores usually release
bears the developing sexual medusoids, the gono- individual cormidia, which are then known as
phores, and in some species asexual nectophores. eudoxids. Each eudoxid, thereby, comprises a bract, a
gastrozooid and tentaele, a succession of sexual gono-
Another cormidial structure is the bract (Fig. lA, phores and, possibly, an asexual nectophore. The spe-
2A), whose medusoid or polypoid origin is uncertain. cific identification of this stage often is difficult, and
These structures have a protective function, overlying depends largely on the structure of the bract.
Siphonophorae 479
Outline cIassification Family Abylidae L. Agassiz, 1862

Subfamily Abylinae L. Agassiz, 1862
The classification adopted here is basically that used Subfamily Abylopsinae Totton, 1954
by Totton (1965). That taxonomy is well established,
although a few more recent authors (e.g. Alvariño, The only family not dealt with herein is the Rhodaliidae
1981) still use a few outmoded names. Other useful whose species are benthic.
references include Bigelow (1911), Kirkpatrick and
Pugh (1984) and Totton (1954). The only change that
has been adopted here is the re-establishment of the Identification of the taxa
Siphonophorae as a subclass of the class Hydrozoa
(see Bouillon el al., 1992), and the consequent raising Because of their fragility, the identification of species
of the 3 old suborders to order status. Totton (1965) of siphonophores is based largely on the morphology
recognised approximately 130 species and since then of one or more of the "individual s" that comprise the
about 70 other species and subspecies either have whole animal. The structure of the nectophores and
been described or resurrected. However, it is certain bracts is of particular importance, although neither of
that several of Totton's "doubtful" species, and many these, with the exception of asexual nectophores on the
of the subsequent new ones, are not valido This is gonodendra, are present in cystonect species. The fol-
because many aberrant forms have been described as lowing diagnoses refer almost exclusively to the adult
new species, or the reasons given for distinguishing stage of the various siphonophore species. The larval
the new material from extant species are insufficient. nectophore and bracts often have a different morpho-
Nonetheless, it is clear from submersible collections logy, but there is insufficient space to deal with these
that there are still many species yet to be described. in the present text. The species included here are those
that have been recorded from the South Atlantic
Phylum Cnidaria Verril, 1865 Ocean. Because of the dearth of sampling, particularly
Class Hydrozoa Owen, 1843 at deeper depth, it is possible that other species may be
Subclass Siphonophorae Eschscholtz, 1829 found, although the list is quite comprehensive.
Order Cystonectae Haeckel, 1887
Family Physaliidae Brandt, 1835 Order Cystonectae
Family Rhizophysidae Brandt, 1825 Siphonophores with a relatively large, and for the
Order Physonectae Haeckel 1888 Portuguese Man O'War enormous, pneumatophore,
Family Apolemidae Huxley, 1859 whose gas gland may bear hypocystic villi. The pneu-
Family Agalmatidae Brandt, 1835 matophore has an apical pore. No nectosome or bracts
Family Pyrostephidae Moser, 1925 are present, and the cormidia are relatively simple
Family Physophoridae Eschscholtz, 1829 comprising only a gastrozooid and tentacle, and a
Family Athorybiidae Huxley, 1859 gonodendron. The tentac1es, which in some species
Family Rhodaliidae Haeckel, 1888 are unbranched, bear only one type of penetrant
Family Forskaliidae Haeckel, 1888 nematocyst. The gonodendron bears gonopalpons,
Order Calycophorae Leuckart, 1854 gonophores and asexual swimming bells. Each animal
Family Prayinae K611iker, 1853 probably bears gonophores of only one sexo Only ~
Subfamily Amphicaryoninae Chun, 1888 recognised species within two families.
Subfamily Prayinae Haeckel, 1888
Subfamily Nectopyramidinae Bigelow, Family Physaliidae
1911 This family is monotypic for Physalia physalis, the
Family Hippopodiidae K611iker, 1853 Portuguese Man O'War.
Family Diphyidae Quoy and Gaimard, 1827 • Physalia physalis Linné 1758. (Fig. 3.1). This is
Subfamily Sulculeolariinae Totton, 1954 the only pleustonic siphonophore. It has a huge,
Subfamily Diphyinae Moser, 1925 asymmetricpneumatophore, purplish blue in
Subfamily Giliinae Pugh and Pages, 1995 colour and up to 30 cm in length. The top of the
Family Clausophyidae Totton, 1954 pneumatophore is formed into an erectile "sail"
Family Sphaeronectidae Huxley, 1859 that runs diagonally across it either from the right-
480 Siphonophorae
hand or left-hand side. The cormidia are attached species presently distributed amongst 7 families, al-
to one side of the float and their tentacles can though it is clear that some revision is necessary.
stretch down many metres. Species from al! 7 families occur in the South Atlantic,
but as those of the family Rhodaliidae are benthic,
Family Rhizophysidae using their tentacles to tether themselves to the substra-
Only 4 species, in 2 genera, are recognised here. te (Pugh, 1983; Hissmann el al., 1995), they will not be
Totton (1965) also included 3 "species inquirendae" considered here. Riemann-Zürneck (1991) gives details
within this family, of which 2 belonged to the genus ofthe distribution of Rhodalia miranda Haeckel, 1888.
Epibulia Eschscholtz, 1829. In these species there is The other 6 families can be distinguished according to
supposed to be a ring of palpons surrounding the base the arrangement of the nectosome and siphosome:
ofthe pneumatophore: but palpons are absent in other 1 Nectophores present 2
cystonect species. Totton (1965) suggested that these la Nectophores absent 6
alleged palpons might be young gastrozooids and per- 2 Nectophores deeply hollowed axial!y and with
sonal observations have shown this to be the case. The tentacles between them; small delicate bracts:
whole specimen is highly contracted and the young Family Apolemiidae
gastrozooids bear no sign of a tentacular bud. The 2a Nectophores not hollowed axially; nectosomal
genus and its species are, therefore, considered to be tentacles absent. 3
invalid. However, Alvariño (1981) has recorded 3 Nectosome and siphosome elongate, with a nar-
Epibulia ritteriana Haeckel, 1888 from the western row stem .4
South Atlantic. Judging from her illustration, showing 3a Nectosome ami/or siphosome contracted or redu-
a pneumatophore with large hypocystic villi, it is con- ced 6
cluded that her specimens most likely belong to 4 Nectophores bilaterally symmetrical, arranged
Rhi::ophysa filiformis. biserially 5
4a Nectophores dorso-ventrally flattened, usually
Genus RIzizophysa Péron and Lesueuer, 1807 (Fig. 3.2) asyrnmetric in shape, arranged in a spiral: Family
Pneumatophore with hypocystic villi; no wing-like Forskaliidae
processes on gastrozooids. 5 Nectophores with straight dorsal radial canal:
Rhizophysa eysenhardti Gegenbaur, 1859 (Fig. Family Agalmatidae
3.3). Mature specimens pale pink in colour. Hypo- 5a Nectophores with sinuous dorsal radial canal:
cystic villi at base of pneumatophore. Tentacles Family Pyrostephidae _
with filiform tentilla. 6 Nectosome reduced or absent; siphosome reduced
Rhizophysa filiformis (Forskal, 1775) (Fig. 3.4). to solid body or corm: Family Athorybiidae
Mature specimens pale green in colour. Well- 6a Nectosome normal; siphosome shortened into
developed hypocystic villi. Tentacles with side laterally expanded spiral sac bearing enlarged pal-
branches (tentilla) of 3 types. Ref.: Totton (1965). pons: Family Physophoridae
Order Physonectae The following diagnoses will mainly concentrate on

Siphonophores with a relatively small apical pneuma- the distinguishing features of the nectophores or
tophore. A nectosome with an array of identical necto- bracts, as these are the parts most frequently found in
phores is, with two exceptions, present. The number net samples.
of nectophores is specifically variable, but they are
usually arranged biserially, except in the family Family Apolemiidae
Forskaliidae. The cormidia, arranged linearly down Nectophore with a very characteristic shape being
the stem, are comprised of a gastrozooid and tentacle, deeply hollowed axially, forming a pair of large axial
and a specifically variable number ofbracts and gono- wings. The nectosac is extensive and its lateral radial
dendra, the last not bearing asexual swirnming bells. canal s follow an S-shaped course of varying com-
As is always the case there is an exception in that one plexity. Uniquely, amongst the physonects, there is a
species does not have bracts. In addition to these tentacle, or clump oftentacles (7 larval) between each
cormidial elements palpons with palpacles usually are pair of nectophores. Small delicate bracts that preser-
present, but they are absent in some species. About 50 ve poorly. Three species in separate genera have been
Siphonophorae 481
described, 2 since Totton (1965). Material collected Nectosac becoming Y-shaped in larger nectopho-
from submersibles indicates that there might be up to res. Lateral radial canal s distinctly looped. Bract
another ten species (Pugh, pers. obs). However, the 4 with 4 (only 2 in youngest) very characteristic dis-
nectophores in the single record for the area (Pages tal facets. Bracteal canal usually ends before
and Gili, 1992) belong to Apolemia l/varia. reaching the distal extremity, but a vestigial canal
may extend towards it.
Genus Apo/emia Eschscholtz, 1829
Monotypic genus for Apolemia l/varia. Genus Bargmallllia Totton, 1954
Apo/emia llvaria (Lesueuer, ? 1811) (Fig. 3.5, Presently a monotypic genus for Bargmannia elongata.
3.21). 5-6 tentacles between each pair of necto- • Bargma1l1lia elollgata Totton, 1954 (Fig. 3.8) [=?
phores. Nectophores having lateral radial canals Mica micula Margulis, 1982]. Very characteristi-
on nectosac with numerous, short, blind-ending cally shaped, elongate nectophores with large
diverticula on the axial half. Flimsy bracts cover- triangular thrust block and axial wings virtually
ed in opague spots. absent. Apico-Iateral ridges each divide above the
ostium. Highly obligue vertical lateral ridges join
Family Agalmatidae infra-Iaterals close to the ostium. Long, narrow
Nectosome without tentacles between the biserially nectosac with straight radial canals. Bract broad
arranged nectophores. Dorsal radial canal on nectosac and rounded with many patches of cells on its dor-
straight. Siphosome straight, not coiled up to form a sal surface. Totton (1965) included this genus in
sac or corm. This is rather a catch all family, and the family Pyrostephidae. Although there are some
Totton (1965) included in it most of the physonect similarities with the single species in that family,
species. At present about 25 species are included in for the present we follow Stepanjants (1967) and
the family, although the status ofsome remains uncer- include the genus in the family Agalmatidae.
tain. As better material is collected for some of the Nevertheless, it is certain that its status will need
"rarer" species, and many new ones, it is probable that further review. Margulis (1982) described a new
certain species will be removed into new families. species Mica micl/la that appears to be the post-Iar-
Genus Aga/ma Eschscholtz, 1825 val stage of a physoneét siphonophore and, from
Tricornuate tentilla consisting of a coiled cnidoband the shape of the nectophores, it is suggested that
enveloped in an involucrum, plus a terminal vesicle the species to which it belongs is B. elongata.
and 2 lateral processes. When the lateral processes of
the tentillum are relaxed, the whole structure has the Genus Cordaga/ma Totton, 1932
semblance of a copepod. Purcell (1980) has suggested Physonects with heart-shaped nectophores devoid of
that this is a form of aggressive mimicry. lateral or vertical lateral ridges.
• Aga/ma e/egalls (Sars, 1846) (Fig. 3.6, 3.22). Cordagalma cordiforme Totton, 1932 (Fig. 3.9,
Apico-Iateral ridges on nectophores with distinct 3.24). Small delicate physonect reaching 30 cm in
notch toward the ostium. Lateral and verticallater- length. Small characteristically heart-shaped necto-
al ridges only present in larger nectophores, and phores. Lateral radial canals with one ascending
even then the fonner can be indistinct. Nectosac loop but without sigmoid curves. AII radial canals
T-shaped. Lateral radial canals distinctly looped. arise from pedicular canal. Characteristic bracts
Foliaceous bracts, usually tridentate at the distal shaped like a truncated pyramid, with 4 lateral
end, but some small bracts do not show this featu- facets. Short bracteal canal terminating in the mid-
re. The convex dorsal side bears 3 ridges, often dIe of the bract. Very characteristic tentilla.
indistinct in smaller ones, on the distal half. Totton's (1932) description of this species was
Bracteal canal extends to about four-fifths the based only on the nectophores with their very char-
length of the bract, occasionally continuing as a acteristic shape. However, Carré (l968a) was able
very fine canal to the distal tipo to give a full description when the entire animal
Aga/ma okelli Eschscholtz, 1825 (Fig. 3.7, 3.23). was collected. From this description it is apparent
Apico-Iateral ridges on nectophores without dis- that the bracts and, particularly, the tentilla of this
tinct notch. One (young nectophores) or 2 (adult) species resemble very closely those of Anthemodes
vertical lateral ridges. Lateral ridges absent. ordinata described by Haeckel (1888). However,
482 Siphonophorae
the nectophores of the two species appear entirely Youngbluth, 1988 it has become apparent that H
different. Nonetheless, there may be sufficient cllpulifera is not the only species whose tentillum
reason to adopt the name Cordagalma ordinata ends in a cupulate process. The two processes are
(Haeckel, 1888), but for the present Totton' s distinguishable, but with careo It is thus uncertain
specific name is retained, with a modified ending as to what species the records given by Alvariño et
to agree with the gender of the genus. Ref.: Carré, al. (1990) be10ng. Ref.: Pugh and Youngbluth
1968a. ( 1988).
Halistelllllla rllbrlllll (Vogt, 1852) (Fig. 3.11,
Genus Erenna Bedot, 1904 3.26). Nectophores with large axial wings and
Monotypic genus for Erenna richardi. A second spe- extensive thrust block. Vertical lateral ridges run
cies, Erenna bedoti, was described by Lens and van obliquely down from apico-Iaterals, but do not
Riemsdijk (1908), but was synonymised with E. reach infra-Iaterals. Similarly, lateral ridges run
richardi by Bigelow (1911); a procedure followed by obliquely up from ostium but do not reach the
Totton (1965). Margulis (1977a, 1990) be1ieved that apico-1aterals. Bracts fo1iaceous, e10ngate, thin,
there were sufficient reasons to resurrect E. bedoti, distally pointed, with 2 1atero-terminal teeth.
but here the synonymy is retained and only a single Bracteal canal runs almost the entire length of the
species recognised. bract, but is absent from a proximal process.
• Erenna richardi Bedot, 1904 (Fig. 3.10, 3.25) Distally it runs beneath an indistinct dorsal ridge
[=Erenna bedoti Lens and van Riemsdijk, 1908]. in which are imbedded 2 rows of cells, probab1y
Large, but thin, nectophores with a relatively large nematocysts. Tentillum with vestigial invo1ucrum
thrust block bearing 2 digitate processes at its base, at base of cnidoband, and a terminal filament,
on the ventral side. Apico-lateral ridges divide very often tightly coiled when contracted, that has no
close to the ostium. Distinct vertical lateral ridges terminal process. It is probable that the nectopho-
define triangular facets on the outer sides of the res and bracts of other Halistemma species have
large axia1 wings. Nectosac has muscle free region been confused with those of H rllbrum. Recent
axially. All radial canals have a straight course and material colleeted by submersible has allowed the
bear black pigmentation. Long, narrow foliaceous distinction of at 1east one other new species (Pugh
bracts, up to 25 mm in 1ength, with one or 2 pairs and Youngbluth, 1988).
oflateral teeth. Bracteal canal runs the entire length Halistelllma striata Totton, 1965 (Fig. 3.12).
ofthe bract and termina tes below an aggregation of Presently identified on1y by its very distinctive
cells (? nematocysts). Distinctive tentilla with nectophores. These are 1arge, robust structures,
uncoiled, powerfully armoured cnidoband, black in with prominent thrust block and deep axial wings.
co10ur; a secondary, diverticulár canal; and a pair There are 3, slightly ob1ique but complete, pairs of
of spots on the broad terminal filament. vertical lateral ridges. Ostia1 to these there is a pair
of lateral ridges, each of which bifurcates shortly
Genus Halistelllllla Hux1ey, 1859 after leaving the apico-laterals, with the upper
Aga1matids whose tentilla have a single terminal fila- branch running down to the ostium, while the
ment (unicornuate) and only a vestigial involucrum. basal branch heads toward the infra-lateral but
Characteristic sigmoidal courses for radial canal s on peters out before reaching it.
the nectosac of the nectophore, that begins with a
downward sweep. The post-Iarval stage of a Hali- Genus Marrlls Totton, 1954
stemma spp. was described by Haeckel (1888) as a Nectophores with straight radial canals; unicornuate
separate species, Nectalia loligo. Pugh (1974) correct- tentilla on tentacles. The exact status of this genus,
Iy surmised its true status, but A1variño (e.g. 1981) and the 3 species contained within, appears uncertain.
continued to refer to it as a separate species. Marrus antarcticlIs Totton, 1954 (Fig. 3.13,
• Halistemma cuplIlifera Lens and van Riemsdijk, 3.27). Nectophores without a vertical lateral ridge.
1908 (cf. Fig. 1C). The only described feature that In 1954 Totton stated that the apico-lateral ridges
cou1d distinguish this species from others of the bifurcated distally; while in 1965 he mentioned
genus Halistemma is the terminal filament of the pairs of Vk'(apico-lateral) and Vk" (lateral) rid-
tentillum that ends in a cupulate process. With the ges. It is, therefore, not clear what affinities the
description of Halistemma transliratum Pugh and latter ridges have. The bracts were described to be
Siphonophorae 483
flattened, cone-shaped, and truncated distally. The radial canals. Tentacles filiform (?). There is some
bracteal canal runs the entire length of the bract doubt as to the validity of this species, particular-
and ends on a small papilla on the distal facet. ly as Margulis's (1982) description is rather vague
New material of this species, in excellent condi- and it is difficult to pin-point any clear distinguis-
tion, needs to be examined in order that a more hing characters. For this reason the nectophores
detailed description can be given. are not illustrated.
Genus Moseria Totton, 1965 Family Pyrostephidae

Very thin, flimsy nectophores with straight radial Monotypic family for the species Pyrostephos van-
canals. Involucrum covers cnidoband of unicornuate hoelJeni.
tentilla. Very little is known about the species of this Pyrostephos vanhoeffeni Moser, 1925 (Fig. 3.15,
genus. The first species to be described was Moseria 3.29). Nectophores with large thrust block and
con voluta (Moser, 1925) based on some young mate- axial wings that are extensive laterally but not
rial, with a pneumatophore of peculiar construction. axially. Highly looped lateral radial canals on nec-
Totton (1965) enhanced the earlier description, inclu- tosac, whose adaxial wall lacks musculature.
ding a new description ofthe tentillum, as he believed Dorsal radial canal with several pronounced
that that described by Moser (1925) belonged to anot- bends; ventral canal straight. Thickened, dorsally
her species. Margulis (1977b) described a new spe- convex bracts, somewhat triangular in shape, with
cies, M. similis, based mainly on a difference in the irregular teeth on the broad distal end. Canal ends
number of cavities in the pneumatophore. However, some distance from distal extremity. Tentilla with
as this gas-filled structure usually explodes when the diverticular canal. Ref.: Totton (1965).
specimen is brought to the surface, and the gas forced
down the stem, the number of cavities may be totally Family Physophoridae
misleading. It is not clear, therefore, whether M. simi- Monotypic family for the species Physophora hydro-
lis is a distinct species. There also appears to be anot- statica.
her Moseria species in the Southern Ocean (see Pages Physophora hydrostatica Forskal, 1775 (Fig.
et al., 1994; Pugh et al., in press) but it has yet to be 3.16). Flimsy, apparently ridgeless nectophores
described. In the area under consideration there are each with an extensive nectosac, which has charac-
records only for M. similis and M. sp. nov. and these teristically looped lateral radial canals. Both dor-
will be considered together as Moseria spp. sal and ventral canal s are sinuous. Siphosome
compact sac on which the simple, bractless cormi-
Genus Nanomia A. Agassiz, 1865 dia are borne in a spiral. Each cormidium has a
Agalmatids whose unicornuate tentillum has a basal single, greatly enlarged palpon, pinkish blue in
involucrum. Characteristic arrangement of the gono- colour. Ref.: Totton (1965).
dendra in that male and female ones, attached at the
bases of palpons, altemate on either side. Family Athorybiidae
Nanomia bijuga (Chiaje, 1841) (Fig. 3.14,3.28). Relatively large pneumatophore. Nectosome greatly
Nectophores are flattened in the abaxial-adaxial reduced (genus Melophysa) or absent (genus
plane and, in that view, square in shape. L-shaped AthOlybia). Siphosome reduced to a dense corm on
when viewed laterally. Ridges indistinct. Axial which the cormidia are arranged in a spiral. Only 3
wings twisted. Looped lateral radial canals on known species, of which 2 occur in the region under
extensive nectosac. Leaf-like bracts, variable in consideration.
designo Often with 3 processes at distal end, occa- • Athorybia rosacea (Forskal, 1775) (Fig. 3.17,
sionally with a cross ridge. 3.30). Large, red pigmented pneumatophore, with
cormidia arranged in a spiral around it. No necto-
Genus Stepanjantsia Margulis, 1982 some. Elongate, flimsy bracts with 7 inconspi-
Monotypic genus for Stepanjantsia polymOlpha. cuous rows of nematocysts running down the con-
Stepal1jantsia polymorpha Margulis, 1982. Minute vex dorsal side. A very distinctive species. The
nectophores, very variable in shape. Nectosac occu- bracts can be moved in a rhythmic manner, proba-
pying most of nectophore, with looped lateral bly enabling the animal to swim about.
484 Siphonophorae
• Melophysa melo (Quoy and Gaimard, 1827) (Fig. reported in some species, but these may be the first
3.18). Nectosome reduced bearing a maximum of (spent) gonophore retained. In many species the matu-
5 nectophores. The nectophores are boot-shaped, re cormidial units are set free from the stem to lead a
with the nectosac occupying most of the foot, and separate existence, known as the eudoxid or sexual
a long, thick pedicular canal running from its apex stage.
up through the heel. The flattened base has two
facets separated by a central protuberance. The About 100 species distributed amongst 6 families.
dorsal radial canal is straight, while the ventral one The families usually can be distinguished by the basic
makes several curves. The lateral canal s are loop- shape ofthe nectophores and bracts, but the following
ed. The large, gelatinous bracts bear a proximal key cannot take account of every specific variation, or
keel for attachment. The dorsal surface of the all the larval stages.
mature bracts bears several rows of prominent I Up to 15+ similar, closely applied, dorso-ventral-
papillae. A very distinctive species. Only the most Iy flattened nectophores bearing protuberances or
distorted nectophores in any way resemble those spines; large, but shallow nectosac; bracts absent:
illustrated by Totton (1954). Family Hippopodiidae
la Small, flattened nectophores, with vestigial nec-
Family Forskaliidae tosac and reduced somatocyst: Family Prayidae,
Genus Forskalia K611iker, 1853 Subfamily Amphicaryoninae
Cylindrical or cone-shaped nectosome, whose nume- lb Nectophores not dorso-ventrally flattened; bracts
rous nectophores have a multiserial, spiral arrange- present 2
ment. Nectophores flattened dorso-ventrally and often 2 Nectophores and bracts rounded, smooth-walled,
asyrnmetrical in shape. Nectosac restricted to basal with thick mesogloea 3
half; with straight radial canals. The siphosome also is 2a Nectophores and bracts pointed, toothed or of irre-
coiled, particularly in its contracted state, with the gular shape .4
gastrozooids borne on long stalks. Bracts usually 3 Usually 2 nectophores ofapproximately equal size,
gelatinous and of variable shape. There is no known forming an apposed pair; somatocyst simple or
way to distinguish the various species on the basis of branched. Bracts with 5 or 6 branches to the canal
their bracts alone. system: Family Prayidae, Subfamily Prayinae
Forskalia edwardsi K611iker, 1853 (Fig. 3.19). 3a Two nectophores of unequal size, with reduced
Slightly asyrnmetrical nectophores with wo di s- somatocysts; nectosac of smaller nectophore usu-
tinct facets on either side of ostium. Bracts varia- ally reduced or obsolescent. Bracts with 2 bran-
ble in shape. ches to canal system: Family Prayidae, Subfamily
• Forskalia leuckarti Bedot, 1893 (Fig. 3.20). Amphicaryoninae
Asymmetric nectophores with only one pronoun- 3b Single, fragile, larval nectophore, with a simple
ced axial wing, and 2 patches of epidennal cells on somatocyst and narrow hydroecium. Small, fragile
their sides. There is a vertical, disc-like expansion bract with a single canal: Family Sphaeronectidae
of the pedicular canal of the adult nectophores, 4 Two, morphologically different nectophores; one
which is usually red in colour. Bracts variable in (anterior) superimposed over the other (posterior)
shape. . 5
4a Single, usually large, nectophore bearing simple or
Order Calycophorae toothed ridges; somatocyst usually branched. Large
Siphonophores without an apical pneumatophore. The bract without neck shield and extensively branched
nectosome usually consists of one or two, often dissi- canal system: Family Prayidae, Subfamily Necto-
milar, nectophores, although up to ca. 15 are present pyramidinae
in the family Hippopodiidae. The cormidial elements 4b Single, usually small, apically pointed nectophore,
consist of a bract, a gastrozooid, with tentacle, and a with simple caecal somatocyst. Bract conical or
succession of gonophores. Again the Hippopodiidae angular 6
are exceptional in that they do not possess bracts. 5 Posterior nectophore with a somatocyst; anterior
There are no palpons, with the possible exception of nectophore with extensive opening of hydroecium
one species. Asexual swirnming bells have also been onto ventral surface. Small bract with phyllocyst
Siphonophorae 485
and 2 canals extending into neck shield: Family Amphicaryon intermedia Daniel, 1970]. The two
Clausophyidae nectophores, both with a functional flask-shaped
5a Posterior nectophore without somatocyst; hydroe- nectosac, fit together to form a ball-like structure.
cium of anterior nectophore usualIy opens basalIy. The larval nectophore has a large, often swollen
Bracts conical or angular, with phylIocyst and, at somatocyst, while that of the definitive one is
most, one canal 6 minute. The radial canals on the nectosac of the
6 Conical stream-lined anterior nectophore, usualIy larval nectophore are highly branched near their
with shallow hydroecium; posterior nectophore, bases, and those of the definitive one also show
when present, usually apically truncated and of some branching. Small, spherical bract with 2
similar size or smaller than anterior one. Conical recurved hydroecial canals.
bracts: Family Diphyidae
6a Anterior nectophore angular, with inflated soma- Subfamily Prayinae
tocyst and deep hydroecium; posterior nectophore, Two, occasionalIy up to 4, rounded, smooth-walled
when present, larger than anterior one. Rigid, nectophores of similar size. Bracts with 6, occasional-
angular bracts: Family Abylidae ly reduced to 5, canals. CurrentIy there are 8 genera
encompassing 16 species, ofwhich 5 have been recor-
Family Prayidae ded in the are a under consideration.
Subfamily Amphicaryoninae Genus Lilyopsis Chun, 1885
Two nectophores differing in size. The larger, rounded Monotypic genus for Lilyopsis rosea.
one is believed to be the retained larval nectophore. • Lilyopsis rosea Chun, 1885 (Fig. 3.35,3.46). Two,
The first definitive one is smaller (genus Maresearsia) possibly more, very delicate nectophores with
or vestigial (genus Amphicaryon). The bracteal canals large nectosacs. Larval nectophore has a simple
are reduced to 2 long hydroecials. The bracts of the 3 somatocyst, slightIy swolIen at its tip, and straight
species of the genus Amphicmyon cannot be distin- radial canals on the nectosac. Definitive necto-
guished at presento phore has a bifurcated somatocyst and sinuous
• AlIlplricaryon acaule Chun, 1888 (Fig. 3.31,3.44). lateral radial canal s on the nectosac. Bract like a
Nectosac of larval nectophore occupies about half cushion, with characteristicalIy arranged canals.
the height, dorsal canal longer than ventral, and This is a very fragile species and is rarely seen;
laterals slightly, and irregularly, expanded in having not, to my knowledge, been colIected by
region where they bend through 90°. The vestigial SCUBA diverso There is thus some doubt, in my
nectophore is embraced by the larval one. Its redu- mind, as to the records in the area under conside-
ced nectosac has no ostial opening and the radial ration here. Ref.: Carré (1969).
canals are simple, but distinct.
• AlIlplricaryoll erllesti Totton, 1954 (Fig. 3.32). Genus Praya Quoy and Gaimard in BlainvilIe, 1834
Larval nectophore similar to that of A. acaule, but Two large, rounded nectophores (often with addition-
lateral radial canals are more convoluted where al reserve bells) whose somatocysts can be complexly
they bend. Flattened vestigial nectophore not em- branched. Multibranched radial canal s on nectosac.
braced by larval one. Nectosac greatIy reduced Bract laterally flattened with dorsal canal arising from
and of characteristic shape. It's ventral canal end of spur-like right longitudinal canal. Gonophores
forms a network on the ventral wall, while the dor- with characteristicalIy three-pronged mantIe canal.
sal canal is simple. No lateral canals. Ref.: Pugh (1992b).
AlIlplricaryon peltifera (Haeckel, 1888) (Fig. • Praya dubia (Quoy and Gaimard, 1833) (Fig. 3.36,
3.33). Larval nectophore similar to those of other 3.47). Cylindrical definitive nectophores with a
species, but lateral radial canals are simple and baso-ventral extension below the ostium ofthe nec-
usualIy straight. The flattened, plate-like vestigial tosac. Bifurcating canals on nectosac. Somatocyst
nectophore is not embraced by the larval one. Its simple in larval nectophore, but with an ascending
nectosac has disappeared leaving 3, characteristi- (dorsal) and two long, branching lateral canals in
cally arranged, finger-like radial canals. the mature definitive one. Bract with relatively
• Maresearsia praeclara Totton, 1954 (Fig. 3.34, short dorsal canal, which may bifurcate subtermi-
3.45) (=Maresearsia sphaera Stepanjants, 1967; nalIy. Left hydroecial canal not recurved distally.
486 Siphonophorae
• Praya reticulata (Bigelow, 1911) (Fig. 3.37, Subfamily Nectopyramidinae

3.48). Nectophores similar to previous species in Only a single, asymmetrical large definitive necto-
basic designo Highly branched canals on nectosac phore, which bears a vague pattern of ridges. Eudoxid
form a reticulated pattern. Somatocyst, in mature stage resembles nectophore, and a special nectophore
nectophore, with several short lateral and ascen- may be present. Smaller larval nectophore also bears
ding branches. Bract usually with relatively long, ridges, which are spinose in the genus Nectopyramis.
recurved dorsal and left hydroecial canals. Ref.: Pugh (1992a).
Genus Nectada11las Pugh, 1992
Genus Rosacea Quoy and Gaimard, 1827 Definitive nectophore with hydroecium restricted to a
Nectophores with simple somatocyst without side pocket-like structure; branched somatocyst with no
branches. Sinuous lateral radial canals on nectosac. lateral bend at base; radial canals on nectosac arise
Bracts kidney-shaped, but with characteristic arrange- from a single pedicular canal. Ridges of larval necto-
ment of canals. The status of the generic name phore not spinose; same arrangement of radial canals.
Rosacea was briefly discussed by Totton (1965), who No special nectophore present on eudoxid.
noted that the name presently may not be being Nectadamas diomedeae (Bigelow, 1911) (Fig.
applied in the way the original authors intended. 3.40, 3.51). Large rhomboidal definitive necto-
However, so as not to further complicate the involved phore bearing a faint pattern of surface ridges.
nomenclature he proposed to reta in the use of the Small nectosac offset laterally; obliquely slanted,
name Rosacea in the way that Bigelow (1911) applied deep hydroecium with narrow opening. Complexly
it, and he designated R. plicata Quoy and Gaimard divided somatocyst. Large, roughly triangular
sensu Bigelow as its type species. Unfortunately, bract with a vague ridge pattern and a small
Margulis (1994) decided to destabilise the nomencla- hydroecium. Phyllocyst complexly divided.
ture, which is contrary to its very purpose, and erected
a new genus Neorosacea. The issue is too complica- Genus Nectopyramis Bigelow, 1911
ted to be dealt with here, but the destabilisation ofthe Definitive nectophore with extensive hydroecium;
nomenclature is rejected and the generic name somatocyst with lateral bend at base; radial canal s
Rosacea is used in the sense that it has been used for arise separately and directly from somatocyst. Larval
the past 85 years, that is sensu Bigelow (1911). nectophore with spinose ridges. Eudoxid bears a spe-
Ultimately, a submission to the ICZN may be requi- cial nectophore.
red; although it is possible that the description of the Nectopyram;s natans (Bigelow, 1911) (Fig. 3.41,
original type species R. ceutensis Quoy and Gaimard, 3.52). Elongate, bow-shaped definitive nectophore
1827 could be interpreted as that of a larval necto- with a pointed apex and truncated base. Seven
phore of a Rosacea sensu Bigelow species. longitudinal ridges arranged characteristically.
• Rosacea cY11lbifor11lis(Chiaje, 1822) (Fig. 3.38, Simple somatocyst without branches. Larval nec-
3.49). Nectophores relatively flimsy, slightly flat- tophore with functional nectosac and 4 radial canals
tened. Hydroecium extends to base of nectophore. arising separately from the simple somatocyst.
Nectosac opens dorso-basally. Bract with dorsal Bow-shaped eudoxid bract with 5 longitudinal rid-
canal arising from left longitudinal canal, prior to ges. Distinctive lateral branch to dorsal canal.
origin of left hydroecial. • Nectopyram;s thetis Bigelow, 1911 (Fig. 3.42,
• Rosacea plicata sensu Bigelow, 1911 (Fig. 3.39, 3.53). Pyramidal definitive nectophore with indis-
3.50). Cylindrical nectophores with solid meso- tinct ridges. Somatocyst with lateral branches.
gloea. The simple somatocyst extends beyond the Larval nectophore with vestigial nectosac, usually
point of origin of the pedicular canal. Extensive with only 2 radial canals. Pyramidal eudoxid bract
hydroecium, not reaching the base of the necto- with 6 canals; dorsal canal with lateral branch.
phore. Nectosac opens basally. Bract similar to R.
cymbiformis, but dorsal canal arises from left Family Hippopodiidae
hydroecial canal distal to the short longitudinal Diagnosis as briefly summarised in above key. There
canal. The nectophores of these two Rosacea spe- are 2 genera, but as Totton (1965) pointed out they are
cies can be difficult to tell aparto Generally R. cym- not really distinct. However, in order to maintain the
biformis lives at shallower depths than R. plicata. stability of the nomenclature he retained both. The
Siphonophorae 487
genus Hippopodius is monotypic for H. hippopus. extensively looped lateral radial canals. Replacement
Poorly preserved material of certain species can be nectophores of both types are frequently produced,
difficult to specifica11y identify. It appears that hippo- and these may have different characters. Small leaf-
podiids may feed almost exc1usively on ostracods like bracts that may not be released as eudoxids.
(Purcell, 1981), and Pugh (1991) found a very signifi- Anterior nectophQres of a few species of the diphyine
cant correlation between the latitudinal and depth dis- genus Lensia may be confused with this genus. Bracts
tributions of the two groups. of various species are not distinguishable. Ref.: Carré
• Hippopodius hippopus (Forskal, 1776) (Fig. (1979).
3.55). Horse-shoe shaped definitive nectophores • SlIlculeolaria biloba (Sars, 1846) (Fig. 3.60).
with 4 rounded dorsal protuberances of variable Anterior nectophore with obliquely angled soma-
size. The nectophores, which can luminesce tocyst and without ostial teeth. Commissural
brightly, generally tum a milky colour when pre- canals present; mouth-plate divided into two elon-
served. gated lobes. Posterior nectophore without ostial
Vogtia glabra Bigelow, 1918 (Fig. 3.56). More teeth; best characterised by the mouth plate with
broadly horse-shoe shaped than H. hippopus with its two laterallobes and a central protuberance.
2 dorso-lateral protuberances. • Sulculeolaria clllllli (Lens and van Riemsdijk,
• Vogtia pelltacalltha K611iker, 1853 (Fig. 3.57). 1908) (Fig. 3.61) [=? Lensia campanella elongata
Roughly quadrangular nectophores. Some ridges Margulis, 1984]. Anterior nectophore without
bear small protuberances, but these are genera11y ostial teeth. Long, filiform somatocyst reaching to
absent from the facets between them. two to three fifths the height of the nectophore.
Vogtia serrata (Moser, 1925) (Fig. 3.59) [=Vogtia Commissural canals may or may not be present.
kllruae Alvariño, 1967]. Roughly triangular nec- Mouth plate formed by two lateral, non-overlap-
tophore with distinctive ridges and no spines or ping lobes, without a central protuberance.
protuberances. Posterior nectophore without ostial teeth with
Vogtia spillosa Keferstein and Ehlers, 1861 (Fig. slightly emarginated mouth plate.
3.58). Roughly quadrangular nectophore bearing • Sulculeolaria mOlloica (Chun, 1888) (Fig. 3.62).
numerous gelatinous teeth on the ridges and the Anterior nectophore with 5 ostial teeth, 3 dorsal
dorsal and lateral facets. and 2 (smaller) lateral. Commissural canal s pre-
sent; minute somatocyst. Mouth plate divided with
Family Diphyidae pair of dorsal teeth. Posterior nectophore with
Usually two, occasiona11y only one, dissimilar stre- similar ostial teeth; mouth plate variably divided,
amlined nectophores. One, the anterior, usually is but with 2 dorsal teeth.
pointed apica11y and is positioned directly above the • SlIlcllleolaria qlladrivalvis Blainville, 1834 (Fig.
other, the posterior, which usually is apically trunca- 3.63). Anterior nectophore with 2 dorsal and 2
ted. The nectosac occupies the bulk of both necto- lateral ostial teeth, although both may be reduced
phores. The hydroecium of the anterior one usually, or, very occasiona11y, absent. Commissural canals
but not always, is sma11 or virtually absent. Bracts present; bilobed mouth plate without protuberan-
generally helmet shaped with a variably shaped phyl- ceso Long, sinuous somatocyst reaching one third
locyst. Except for one or species, no bracteal canals to two fifths the height of the nectophore. Posterior
are present. It is difficult to give an a11 embracing nectophore with similar arrangement of ostial
diagnosis for the ca. 50 species in this family as there teeth. Deeply divided mouth plate with 2 protube-
is so much specific variability. The family is divided rances, although these can be reduced or absent.
into 3 subfamilies. The third subfamily has been erec- The nectosac has two characteristic constrictions.
ted recently (Pugh and Pages, 1995) to encompass a • Sulculeolaria turgida (Gegenbaur, 1853) (Fig.
single species, Gilia reticulata. 3.64). Anterior nectophore without ostial teeth.
Subfamily Sulculeolariinae Commissural canals may or may not be present.
Monotypic subfamily for genus Sulcllleolaria. Minute ovoid or filiform somatocyst. Bilobed
Genus SlIlcllleolaria Blainvi11e, 1834 mouth plate without protuberances. Posterior nec-
Anterior nectophore with rounded apex, and without tophore without ostial teeth and rounded, undivi-
ridges; posterior nectophore of similar size with ded mouth plate without protuberances.
488 Siphonophorae
Subfamily Diphyinae Genus Dimophyes Moser, 1925

Only a single anterior and posterior nectophore pro- Monotypic genus for Dimophyes arctica.
duced, although the latter may be reduced or suppres- Dimophyes arctica (Chun, 1897) (Fig. 3.67,3.79).
sed entirely. Anterior nectophore usually apically Anterior nectophore without lateral ridges. The
pointed and usually larger than posterior one. The mouth plate is undivided and the prominent
bracts ofthe free-swirnming eudoxids usually are hel- hydroecium is largely open on its ventral side.
met-shaped. The bulk ofthe diphyid species is contai- Carrot-shaped somatocyst reaching to about two-
ned in this subfamily divided amongst about 7 genera, thirds the height of the nectophore. Posterior nec-
with most being included in the catch-all genus tophore reduced and seldom found; with the ope-
Lensia. In several species the posterior nectophore is ning of the nectosac Iying dorso-basally. Very
either not developed or has not been described. Some, characteristic conical bract, with extensive neck
particularly within the genus Lensia, are very difficult shield that is penetrated by a median canal from
to distinguish specifically. Similarly many eudoxid the phyllocyst. The latter has both apical and later-
stages remain unknown. It is, therefore, impossible to al horns.
give a comprehensive description for every part of
every speCles. Genus Diphyes Cuvier, 1817
Genus Chelophyes Totton, 1932 Anterior nectophores with 5 complete longitudinal
Rigid anterior nectophores with 5 ridges, although ridges. There are 3 prominent ostial teeth, although in
dorsal one extends only a short distance up from the one species the dorsal one is obscured. Deep hydroe-
ostium. Divided mouth-plate; hydroecium claw-sha- cium, only a small basal portion of which may open
ped in lateral view extending to about one sixth the ventrally. Posterior nectophore, when developed, also
height of the nectophore and not opening ventrally. with 3 ostial teeth, but again the dorsal may be obscu-
Posterior nectophore apically pointed and the divided red. Long apical process (apophysis) that is inserted
mouth plate forms 2 strong, asyrnmetric teeth. Strong into hydroecium of anterior nectophore. Bracts gene-
serrations on basal parts of ridges, particularly the rally helmet-shaped, although flattened in one spe-
ventral ones. Conical eudoxid bracts with small, roun- cies. There are 4 recognised species, all ofwhich have
ded neck shield, and relatively deep hydroecium. The been found in the area under investigation.
phyllocyst is cylindrical and almost stretches to the Diphyes alltarctica Moser, 1925 (Fig. 3.68, 3.80).
apex. The two species are very similar in form and the Apex of nectosac of anterior nectophore not cons-
eudoxid stages have not been distinguished. tricted, Iying close to apex of nectophore.
Chelophyes appelldiclIlata (Eschscholtz, 1829) Hydroecium to about half the height of the nec-
(Fig. 3.65, 3.78). Only the right lateral and the 2 tophore; relatively short spindle-shaped somato-
ventral ridges reach the apex of the anterior nec- cyst. Dorsal ostial tooth obscured, and smaller
tophore, the former having twisted so as to lie dor- than laterals. Very broad, divided mouth plate.
sally. Long, spindle-shaped straight somatocyst. Dorsal ostial tooth of posterior nectophore simi-
Ventral ridges of posterior nectophore bear an larly obscured. Head piece and neck shield of
obvious notched tooth on a level with the ostium. bract of similar height; spindle-shaped phyllocyst
Chelophyes cOlltorta (Lens and van Riemsdijk, tapering toward its topo
1908) (Fig. 3.66). Anterior nectophore very similar Diphyes bojani (Eschscholt1, 1829) (Fig. 3.79,
to that of C. appendiculata. The main difference is 3.81). Anterior nectophores with variably serrated
that the ventral facet is strongly twisted to the right. ridges. The nectosac gradually narrows toward its
In this case the right ventral ridge does not reach the apex, which lies close to that of the nectophore
apex, and it may merge with the left ventral close to itself. Hydroecium extends to about one-third the
the apex. The somatocyst is similarly affected by height of the nectophore, with the spindle-shaped
the torsion and is twisted to the right. The posterior somatocyst extending up from it to close to the
nectophore does not have a distinct notched tooth apex of the nectophore. The dorsal ostial tooth is
on the ventral ridges. The anterior nectophores of of equal size to or smaller than the lateral ones.
the two Chelophyes species are easily confused, Mouth plate not divided. Posterior nectophore
particularly if the somatocyst of that of C. appendi- with similar arrangement of 3 ostial teeth. Lateral
culata becomes distorted during preservation. teeth, and the baso-lateral margins are serrated.
Siphonophorae 489
"
Eudoxid bract shield-like, lying on the ventral side • Elldoxoides spiralis (Bigelow, 1911) (Fig. 3.73,
of the gonophore. 3.84). Anterior nectophore with characteristically
• Diplryes clramissollis Huxley, 1859 (Fig. 3.70). twisted, serrated ridges. The ventral ridges unite
Anterior nectophore is blunt apically. Nectosac c10se to the apex. Large, divided and c1aw-shaped
with rounded apex lying a short distance from that hydroecium. Carrot-shaped somatocyst extending
of the nectophore. Hydroecium extends to mid- to about halfthe height ofthe nectophore. No pos-
height of nectophore; with relatively short spindle- terior nectophore is developed. Conical bract with
shaped somatocyst extending above it. Ostial teeth lightly serrated ridges. Large neck shield; shallow
of equal size. Mouth plate not divided. Posterior hydroecium; thick phyllocyst reaching almost to
nectophore not developed; eudoxid not found in apex. The gonophore is also characteristically
area. Like Chelophyes conforta, this is largely a twisted.
neritic Indo-Pacific species and has only been
found off South Africa due to t~ intrusions of Genus Lellsia Totton, 1932
water from the Agulhas Current. Small pyramidal anterior nectophores, generally rid-
• Diplryes dispar Chamisso and Eysenhardt, 1821 ged, the number and disposition of the ridges being
(Fig. 3.71, 3.82). Very similar to D. bojani. variable. Small, divided mouth plate, with shallow
Nectosac of anterior nectophore cylindrical basal- hydroecium, rarely extending above ostial level. No
ly but with narrow caecal extension ending c10se to ostial teeth. Posterior nectophore, when developed,
the apex ofthe nectophore. Hydroecium extends to truncated apically with a rounded mouth plate. Bracts
about one-half the height of the nectophore, with helmet-shaped. Shape of phyllocyst generally resem-
the spindle-shaped somatocyst extending up to just bling that of somatocyst of anterior nectophore. This
above the beginning of the nectosacal caecum. The is a catch-all genus, containing over thirty species that
dorsal ostial tooth is considerably larger than the do not show the distinctive characters of the other
lateral ones. Mouth plate not divided. Posterior genera. The species are here grouped according to
nectophore with similar arrangement of ostial whether they have 7 or less ridges (1 dorsal, 2 ventral
teeth. Lateral teeth and baso-lateral margins not and O, 2 or 4 laterals), all running longitudinally, or
serrated. Bract somewhat similar to that of they are multi-ridged.
Chelophyes appendiclllafa, but the phyllocyst is
narrower and tapers towards its apex, which lies Species with 7 or less ridges
further from the apex of the bract.
• Lensia aclrilles Totton, 1941 (Fig. 3.87) [=Lensia
Genus Elldoxoides Huxley, 1859 elfanin Alvariño and Wojtan, 1984]. Anterior nec-
Small, rigid anterior nectophores with 5 ridges; the tophore with 5 complete ridges. The basal parts of
dorsal one being complete. Mouth plate divided; no the lateral ridges bend sharply dorsad. Small,
conspicuous ostial teeth. broad mouth plate, with hydroecium extending to
• Elldoxoides mitra Huxley, 1859 (Fig. 3.72,3.83). ostiallevel. Elongate somatocyst, somewhat varia-
Anterior nectophore not spirally twisted. AII rid- ble in shape, extending to a quarter to a half the
ges complete and serrated basally. Dorsal ridge height of the nectophore. Posterior nectophore and
extended to form basal tooth. Large mouth plate eudoxid stage have not been described.
with serrated wings, the left one being larger than • Lensia campallella (Moser, 1925) (Fig. 3.88,
the right. Hydroecium truncated apically; somato- 3.106) [=Lensia ellgenioi Alvariño and Wojtan,
cyst relatively short and pear-shaped. Posterior 1984]. Anterior nectophore with indistinct ridges
nectophore with distinct notch between apex and that have been subjected to a torsion towards the
apical apophysis. Hydroecium open except near right. This results in a twisting of the apex of the
apex where 2 hydroecial flaps overlap each other. preserved specimens. Oblique baso-ventral facet
Large undivided, asymmetric mouth plate with 2 with mouth plate and hydroecium being virtually
large teeth. Large teeth on ventral ridges about the non existent. Short, oblique, c1ub-shaped somato-
level of the ostium, also asymmetrically disposed. cyst. Bract conical with rounded apex, with minu-
Bract with serrated ridges and a deep hydroecium. te hydroecium. Club-shaped phyllocyst. Ref.:
Phyllocyst pear-shaped. Carré (l968b).
490 Siphonophorae
• Lensia conoidea (Keferstein and Ehlers, 1860) obliquely inc1ined, ovate somatocyst with short
(Fig. 3.89, 3.107). Anterior nectophores with 5 stalk. Posterior nectophore difficult to distinguish
complete straight ridges. Mouth plate shallow but from that of other Lensia species; eudoxid stage
broad; hydroecium minute. Spindle-shaped soma- not described.
tocyst extending to over half the height of the • Lensia lumter Totton, 1941 (Fig. 3.95). Anterior
nectophore. Posterior nectophore with 5 ridges, nectophore with 7 straight longitudinal ridges. The
with a large, asyrnmetric, notched mouth plate. dorso-lateral ridges do not reach the ostium, while
Bract conical with long spindle-shaped phyllocyst. the ventro-Iaterals, which basally form the mar-
• Lensia cossack Totton, 1941 (Fig. 3.90). Anterior gins of the mouth plate, do not reach the apex.
nectophore with very vague ridges, but obvious Oblique mouth plate, occupied by hydroecium,
folds. Minute mouth plate with oblique basal which lies below the ostium and is open ventrally.
facet, and virtually no hydroecium. Ovoid, obli- Short, usually bilobed somatocyst borne on short
quely inc1ined somatocyst reaching up to about stalk. Posterior nectophore and eudoxid stage have
one third the height of the nectophore. This spe- not been described.
cies may not develop a posterior nectophore and • Lensia lelollpi Totton, 1954 (Fig. 3.96). Anterior
the eudoxid has not been described. Alekseev nectophore with 5 straight longitudinal ridges, the
(1984) suggested that this species should be rele- laterals not reaching the ostium. Oblique mouth
gated to a subspecies of Lensia campanella, but plate with hydroecium reaching slightly above
for the present it is retained as a separate one. ostial leve!. Narrow somatocyst stretching to
• Lensiafowleri (Bigelow, 1911) (Fig. 3.91, 3.108). about one third the height of the nectophore.
Anterior nectophore with 5 complete, straight rid- Posterior nectophore and eudoxid stage ha ve not
ges. Mouth plate relatively extensive and contains been described. This species c10sely resembles L.
the globular somatocyst; hydroecium virtually subtiloides (see below) and further studies on the
absent. Hydroecium of posterior nectophore deli- relationship of these two species are warranted.
mited by triangular wings; mouth plate rounded • Lensia meteori (Leloup, 1934) (Fig. 3.97). Fragile
and undivided. Elongate bract, rounded apically, anterior nectophore without any obvious ridges.
with a wide notched neck shield. Phyllocyst glo- Narrow, almost vertical mouth plate with shallow
bular. hydroecium, extending above ostial level and
• Lensia hardy Totton, 1941 (Fig. 3.92). Anterior open ventrally. Somatocyst expanded laterally
nectophore with 5 complete, straight longitudinal with short stalk. Posterior nectophore and eudoxid
ridges. Mouth plate broad but shallow, largely stage have not been described.
occupied by hydroecium, which lies below ostial • Lensia 11l11lticristata (Moser, 1925) (Fig. 3.98).
leve!. Globular somatocyst with short stalk. Anterior nectophore with 7 straight longitudinal
Posterior nectophore and eudoxid stage have not ridges. Neither ofthe pairs oflaterals reach the osti-
been described. This species c10sely resembles L. um, and the ventro-Iateral pair also do not reach the
jowleri. apex. Small mouth plate with shallow hydroecium
• Lensia havock Totton, 1941 (Fig. 3.93). Anterior not extending above ostial leve!. Somatocyst long
nectophore with 7 complete longitudinal ridges. A and thin, extending to one third to a half the height
characteristic feature is that the ventro-Iateral rid- of the nectophore. The posterior nectophore is hard
ges bend ventrad, slightly above the ostial level, to distinguish. The bract has not positively been
and run down onto the mouth plate. Extensive identified, but probably is that of Eudoxia tefluis
hydroecium extending to well above ostial level, Patriti, 1965 (see Kirkpatrick and Pugh, 1984).
and largely open ventrally. Short somatocyst, usu- • Lensia sllbtilis (Chun, 1886) (Fig. 3.99, 3.109).
ally c1ub-shaped, but its outline is often difficult to Fragile anterior nectophore, up to 10 mm in
discern. Posterior nectophore and eudoxid stage height, apparently ridgeless, but with folds. Small
ha ve not been described. mouth plate with oblique basal facet and shallow
• Lensia hotspllr Totton, 1941 (Fig. 3.94). Anterior hydroecium. Somatocyst spherical borne on a long
nectophore with 5, complete, straight longitudinal stalk. Small, rounded bract with small phyllocyst.
ridges. Small mouth plate, with basal facet obli- The posterior nectophore is hard to distinguish. In
quely inclined, and minute hydroecium. Short, general form the anterior nectophore resembles
Siphonophorae 491
that of Lensia meteori, but the shape of the soma- plete, straight ridges that are linked by a velar
tocyst clearly distinguishes them. ridge. Mouth plate rounded ventrally and hydroe-
Lellsia sllbti/oides (Lens and van Riemsdijk, cium, which extends above ostial level, is open
1908) (Fig. 3.100, 3.110). Anterior nectophore ventrally. Short, laterally expanded, mushroom-
with 5 straight, complete ridges. Small mouth like somatocyst. Posterior nectophore and eudoxid
plate and hydroecium with an ovoid somatocyst stage have not been described. The eudoxid stage
borne on an obligue stalk. Rounded bract with described by Alvariño and Wojtan (1984) belongs
club-shaped phyllocyst. The posterior nectophore to Dimophyes arctica.
is hard to distinguish. Like Chelophyes con torta
and Diphyes chamissonis this is largely an Indo- Genus Mllggiaea Busch, 1851
Pacific neritic species that can be carried into the Anterior nectophore with complete dorsal ridge Deep
Atlantic via the Agulhas Current. Other Atlantic hydroecium not open ventrally, although the divided
records should be treated with caution, particular- mouth plate may be obligue. Somatocyst arises very
ly in the light of Totton's (1954) description of a close to wall of nectosac. A posterior nectophore is
very similar species Lensia /eloupi. not developed. Three of the 4 known species are lar-
gely neritic.
Species with more than 7 ridges • Mllggiaea at/alltica Cunningham, 1892 (Fig.
3.74, 3.85). Anterior nectophore with 5 complete,
Lensia ajax Totton, 1941 (Fig. 3.101). Anterior relatively straight longitudinal ridges. Relatively
nectophore with 5 sets of from 2 to 4, usually 3, deep hydroecium, extending to one third the
often incomplete longitudinal ridges. Hydroecium height of the nectophore. Long, thin somatocyst
extends to ostiallevel and is open ventrally. Small, reaching to apex of nectosac. Small conical bract
club-shaped somatocyst. Mouth plate sguarely with an asymmetrical neck shield and a very shal-
truncated basally. Posterior nectophore and low hydroecium. The phyllocyst is club-shaped.
eudoxid stage have not been described. Mllggiaea bargmallllae Totton, 1954 (Fig. 3.75).
• Lellsia exeter Totton, 1941 (Fig. 3.102). Anterior Anterior nectophore with lateral folds in place of
nectophore with several ridges, only 5 of which ridges. Nectosac reaches to almost the apex of the
reach the apex. The dorso-lateral ridges connect nectophore. Obligue mouth plate, with relatively
with a transverse velar ridge. Large mouth plate with shallow hydroecium. Somatocyst sausage-shaped
hydroecium extending up to ostial level and open extending to over halfthe height ofthe nectophore.
ventrally. Long, slender somatocyst extending to The eudoxid stage has not been described. This
one third the height ofthe nectophore. Posterior nec- species can be confused with Dimophyes arctica,
tophore and eudoxid stage have not been described. but the divided mouth plate clearly distinguishes it.
Lensia grima/di (Leloup, 1933) (3.103). Anterior Mllggiaea kochi (Will, 1844) (Fig. 3.76). Anterior
nectophore with a variable number of ridges. Only nectophore with 5 complete longitudinal ridges,
the 2 ventral ridges are complete, and only 5 rid- the laterals bending dorsad toward their bases.
ges reach the apex. The dorso-lateral ridges turn Relatively shallow hydroecium, with somatocyst
dorsal near their bases and bifurcate. Large mouth extending to only half the height of the necto-
plate and hydroecium, which is open ventrally, phore. The eudoxid stage is not distinguished from
extends to slightly above ostial leve!. Short, club- that of Mllggiaea atlantica.
shaped somatocyst. Posterior nectophore and
eudoxid stage have not been described. Subfamily Giliinae
Lellsia hosti/e Totton, 1941 (3.104). Anterior nec- Monotypic subfamily for Gilia reticlllata.
tophore with multiple ridges; several of the later- Gilia reticu/ata (Totton, 1954) (Figs 9.13, 10.9).
als being complete. Extensive mouth plate, and Anterior nectophore with a reticulated pattern of
deep hydroecium, which is open ventrally. ridges and a velar ridge. Mouth plate divided with
Inverted, heart-shaped somatocyst whose true out- hydroecium extending above ostial leve!. Small
line may be difficult to discern. Posterior necto- ovoid somatocyst. Posterior nectophore and bract
phore and eudoxid stage have not been described. also with reticulated pattern of ridges. Phyllocyst
Letlsia /e/ollveteall Totton, 1941 (Fig. 3.105). of bract with swollen centre and a tapering irregu-
Anterior nectophore with numerous, mainly com- lar apico-dorsal process. Two canals ron down into
492 Siphonophorae
the neck shield. This very distinctive species pre- distinct teeth. Hydroecium reaches to only a little
viously had been included in the diphyine genus over half the height of the nectosac. Main body of
Lensia, but recently its status has be come uncer- somatocyst consists of 2 broad wings, of variable
tain. This is because of the presence of 2 canal s in extent, with simple canal running up from it to apex
the neck shield of the bract (see Pugh and Pages, of nectophore. Mouth plate of posterior nectophore
1995), which is a characteristic of the bracts of bearing 2, distinctly asyrnmetrical teeth.
species of the family Clausophyidae. The necto-
phores, however, show typical diphyine characters Genus Clallsophyes Lens and van Riemsdijk, 1908
and so Pugh and Pages (1995) chose to erect a new Smooth-walled, ridgeless, laterally flattened, rounded
diphyid subfamily to encompass the species. nectophores. Anterior nectophore with deep hydroe-
cium in basal half. Lateral radial canals of both necto-
Family Clausophyidae Totton, 1965 phores are looped. Somatocyst long tube of varying
Both anterior and posterior nectophores possess a thickness, part of which is swollen. For most species
somatocyst. The phyllocyst of the eudoxid bract, the eudoxid stage is unknown. The genus currently
when known, has 2 fine branch canals that run down includes 5 species, 2 of which have been described
into the neck shield. This is a small group of ten, only recently. The anterior nectophores of 3 of the
mainly deep living, species divided amongst 4 genera. species are very similar in basic formo The first species
Nine of the species have been found within the area to be described was Clausophyes ovata (Keferstein
under consideration. and Ehlers, 1860), but Margulis (1988) pointed out
Genus CllIlIliphyes Lens and van Riemsdijk, 1908 that since Moser (1925) this name had been wrongly
Nectophores with ridges that end in distinct teeth. applied to another species. The name C. moserae has
Anterior nectophores with 4 ridges (dorsal, ventral been established for this other species. A1though this
and a pair of lateral s) at the pointed apex. Each of destabilises the nomenclature, the species in question
these ridges bifurcates below the apex, the ventral s have not been recorded very often, particularly C.
forming the margins to the hydroecium, so that there ovata, and so this change is accepted here. A table
are 8 ridges at the base. Posterior nectophore with 3 comparing the characters of the 4 species found in the
ridges (dorsal and pair of laterals) reaching apex, all area under consideration is given by Pugh (1995).
ofwhich bifurcate further down. Hydroecium extends • Clallsophyes galeata Lens and van Riemsdijk,
for virtually the whole length of the nectophore and 1908 (Fig. 3.113). Large, robust anterior necto-
has 2 large, asyrnmetrical flaps in its upper half. Bract phore whose mouth plate is partially divided.
flattened; phyllocyst asyrnmetrical with an apical and Deep hydroecium extending to over half the
2 lateral horns in addition to the canals running down height of the nectophore, with thickened walls,
into the neck shield. The two known species are very and a large, central protuberance, where the poste-
similar in design, and the eudoxid stages are, at pre- rior nectophore attaches. Thick somatocyst, irre-
sent, indistinguishable. The anterior nectophore is gularly expanded. Large, robust posterior necto-
usually smaller than the posterior one. phore, with relatively complicated arrangement of
C1l1miphyes lIIoserae Totton, 1954 (Fig. 3.111). lateral radial canal s on nectosac. The most charac-
The lateral ridges bifurcate relatively close to the teristic feature is the presence of a pair of finger-
apex of the anterior nectophore, and the resulting like projections from the walls of the hydroecium,
dorso-Iaterals do not end, basally, in pronounced close to its apex. Ref.: Pugh and Pages (1993).
teeth. The hydroecium stretches up almost to the Clallsophyes laetlllata Pugh and Pages, 1993
same level as the apex of the nectosac. Main body (Fig. 3.114). Anterior nectophore with undivided
ofthe somatocyst globular with a narrow canal, that mouth plate. Deep hydroecium extending to two-
may bear side branches, running up to the apex of thirds the height of the nectophore, and having
the nectophore. In the posterior nectophore the characteristic flaps. Somatocyst expanded only in
mouth plate bears 2, only slightly asymmetric teeth. its apical half. The posterior nectophore and
• C1l1l1liphyes1II11ltidentataLens and van Riemsdijk, eudoxid stage have not been described.
1908 (Fig. 3.112, 3.120). Lateral ridges bifurcate Clallsophyes lIIoserae Margulis, 1988 (Fig. 3.115).
relatively further away from the apex of the anteri- Anterior nectophore up to 20 rnm in height, but
or nectophore, and dorso-Iaterals end, basally, in usually much smaller (7-8 rnm). Mouth plate almost
Siphonophorae 493
completely divided; hydroecium usually extending very similar in basic formo However, Moser (1925)
to half the height of the nectophore, and without initially separated the species into separate genera, the
flaps. The central third ofthe somatocyst is regular- other being Thalassophyes Moser, 1925. Totton
Iy, but variably, expanded. Posterior nectophore (1965) pointed out that they were probably congeneric
with simple somatocyst; no finger-like processes in but retained the two genera, while Margulis (1988)
hydroecium, whose left lateral wing is entire. went as far as synonymising them as a single species.
Clallsophyes tropica Pugh, 1995 (Fig. 3.116). The latter is clearly not the case, but the two species
Nectosac of anterior nectophore relatively small are here treated as congeneric.
occupying less than half the height of the necto- Heteropyralllis crystallina (Moser, 1925) (Fig.
phore. Hydroecium without flaps, but with extensi- 3.118,3.121). No opaque spots present on ridges
ve central process. Somatocyst expanded, to a of anterior nectophore or eudoxid stage. Eudoxid
variable degree for the quarter below its middle. bract with relatively large neck shield.
Posterior nectophore with relatively small nectosac. Heteropyralllis maculata Moser, 1925 (Fig.
Somatocyst expanded in an irregular fashion. 3.119, 3.122) [=Heteropyrarnis alcala Alvariño
Extensive flap in the middle of the left hydroecial and Frankwick, 1983]. Opaque spots present on
wmg. some of the ridges and at the apex of the anterior
nectophore and eudoxid bract, and on the gono-
Genus Crystallophyes Moser, 1925 phore. Neck shield of bract relatively smal!.
Monotypic genus for Oystallophyes arnygdalina.
Crystallophyes amygdalina Moser, 1925 (Fig. Family Sphaeronectidae
3.117) [=Thalassophyes ferraril Alvariño and The small, delieate, rounded larval nectophore is the
Frankwick, 1983]. Anterior nectophore with 5 only one developed. The bract also is small and round-
complete ridges; the pair of laterals being sharply ed. This is a small group of species, rarely found in net
recurved ventrad at their bases. Nectosac extends material, all of which are here treated as belonging to
to three quarters the height of the nectophore, and the genus Sphaeronectes Huxley, 1859. Only one spe-
the shallow hydroecium almost reaches the apex. cies has been found in the area under consideration.
The somatocyst is swollen in its basal two thirds, Sphaeronectes gracilis (Claus, 1873) (Fig. 3.43,
and the canal above may have short side branches. 3.54). Larval nectophore whose nectosac occupies
Posterior nectophore also with 5 ridges, but on the basal half and has 4 straight radial canals. The
each side the laterals and ventrals unite close to the long, narrow hydroecium bends over one side of
apex. Basally the laterals end in distinct teeth. The the nectosac. The narrow, curved somatocyst may
hydroecium extends the entire height of the nec- have a slight terminal swelling. Small hemispheri-
tophore, and has small, pointed flaps. The long cal bract, with minute neck shield and shallow
somatocyst is a simple cana!. The eudoxid stage hydroecium. The long, club-shaped phyllocyst is
has not been described. Margulis (1988) transfer- obliquely angled.
red this species into the genus Chuniphyes, possi-
bly based on Totton's (1954) statement that there Family Abylidae
was an obvious relationship between the two Rigid, angular nectophores, the posterior one, wit-
genera. However, Totton (1954) then noted the hout a somatocyst, usually being much larger, and
clear differences between the genera and retained, bearing serrated ridges and teeth. In all but one spe-
as is followed here, the genus Crystallophyes. cies the somatocyst of the anterior nectophore has
curved over to occupy a ventral position. There are
Genus Heteropyramis Moser, 1925 two subfamilies that are distinguished by the presen-
Anterior nectophore with 5 straight, complete longitu- ce (Abylinae) or absence (Abylopsinae) of a rectan-
dinal ridges; small nectosac and hydroecium restricted gular apical facet on the anterior nectophore.
to central half. Somatocyst thick and slanting basally, However, there is an exception in the former subfa-
with a narrow apical canal extending to close to the mily. Sears (1953) made a detailed study of this
apex of the nectophore. Posterior nectophore not family using material from the "Dana" collections,
described and possibly not developed. Pyramidal bract and described several new species. However, Totton
whose phyllocyst resembles the somatocyst of the (1965) pointed out that many of the descriptions
anterior nectophore. The two species in this genus are were based on deformed specimens. Nonetheless he
494 Siphonophorae
retained some of the new species, whilst noting that it divided by a transverse ridge. Posterior nectophore
was not clear whether they were truly distinct. Here we long and narrow, without wing-like expansions. Short
go further and reject all of Sears' (1953) new species dorsal ridge ends on the dorsal tooth. Bracts with a
and synonyrnise them with previously described ones. median dorsal ridge. Left lateral facet divided by
Several other, more recent descriptions of new species another ridge with a characteristic disposition. Phyllo-
also appear to be based on deformed specimens and are cyst with 2 thin ventro-Iateral branches. Its distal end
also treated as junior synonyms. bends dorsad to form a blind saco
Subfamily Abylinae Ceratocymba dentata (Bigelow, 1918) (Fig.
Genus Abyla Quoy and Gaimard, 1827 3.126, 3.134) Anterior nectophore with triangular
Anterior nectophores with lOor 11 facets. The apical dorsal facet. Its lateral margins are deeply bowed
facet is divided by a transverse ridge, and many ridges and serrated. This facet continues over the top of
¡¡re serrated, particularly basally. Dorsal nectosac and the nectophore and extends into a short peak into
median hydroecium are long tubes extending almost to which the nectosac is extended. Posterior necto-
apex of nectophore. The large oval somatocyst lies vent- phore has a relatively large dorsal tooth and nume-
rally. Posterior nectophore with long, tapering apical rous small teeth on the combo The left lateral ridge
apophysis, has only 4 ridges. The ventral ridges define on the bract joins the apico-Iateral ridge, but not
the hydroecial wings and are heavily serrated basally. the posterior margino
The left hydroecial wing bears a toothed comb or flap. Ceratocymba lellckarti (Huxley, 1859) (Fig.
Five, usually serrated, ostial teeth. Prismatic bracts with 3.127,3.135). Anterior nectophore rectangular in
6 facets, the dorsal one being rectangular. Very large shape with flat apical facet and narrow ventral
phyllocyst, with 2 canals running down toward the vent- one. Apices of hydroecium, nectosac and somato-
ro-Iateral comers of the apical facet. The bracts (Fig. cyst all on the same leve!. Posterior nectophore
3.133) cannot, at present, be identified specifically. with only 5 or 6 teeth on the combo Left lateral
• Abyla bicarinata Moser, 1925 (Fig. 3.123) ridge on bract complete .
[=Abyla brownia Sears, 1953]. Wide anterior nec- •- Ceratocymba sagittata (Quoy and Gaimard, 1827)
tophore with lateral ridges defining wing-like pro- (Fig. 3.128, 3.136) [=Ceratocymba intermedia
cesses, and edges offacets rounded. No transverse Sears, 1953, C. indica Daniel, 1970]. Very charac-
ridge between ventral and apico-ventral facets. teristic anterior nectophore with a long pyramidal
Posterior nectophore as wide as it is long. Comb apical extension that masks the apical facet. Long,
with 4 to 7 teeth; strong ostial teeth. tubular nectosac about twice as long as hydroe-
• Abyla Izaeckeli Lens and van Riemsdijk, 1908 cium. Posterior nectophore with characteristically
(Fig. 3.124) [=Abyla ingeborgae Sears, 1953]. long right ventral tooth. The left lateral ridge on
Anterior nectophore as wide as it is long, without bract joins the posterior margin but not the apico-
wing-like processes. Transverse ridge separates lateral one.
ventral facet from apico-ventral one. Posterior nec-
tophore with up to 5 teeth on combo Lateral ostial Subfamily Abylopsinae
teeth closer to dorsal tooth than to ventral ones. There are 3 genera and 4 species.
Abyla trigona Quoy and Gaimard, 1827 (Fig. Genus Abylopsis Chun, 1888
3.125) [=Abyla carina Haeckel, 1888, Abyla peru- Seven-faceted anterior nectophore, but without an
ana Sears, 1953, Abyla schmidti Sears, 1953, ? apical facet, and pentagonal dorsal and ventral facets.
Abyla tottoni Sears, 1953]. Anterior nectophore as Hydroecium only partially interposed between soma-
broad as it is wide, with most ridges heavily, but tocyst and nectosac. Somatocyst with apical diverti-
irregularly, serrated. Ventral facet not separated, culum. Posterior nectophore with 5 ridges and a short
by a transverse ridge, from apico-ventral one. curved apical apophysis. The left lateral ridge bifur-
Posterior nectophore with from 4 to II teeth on cates close to the apex. Flaps on both wings of
combo Two rows of teeth on basal margin of right hydroecium. Bracts with 7 facets. Phyllocyst with
ventral wing. Ostial teeth heavily serrated. swollen apico-Iateral branches and apical diverticu-
lum, while distally it is a narrow tube. The anterior
Genus Ceratocymba Chun, 1888 nectophores of the two species are very similar in
Anterior nectophore with 7 facets. Apical facet not designo
Siphonophorae 495
Cystonectae
Physalia physalis
I ,
I '
I
1
3.3 L--J
3.1
1 Rhizophysa
L...J Rhizophysa sp. filiformis¡
tentilla
Rhizophysa
eysenhardti¡
tentillum
Upper view of nectophores

or whole specimens of Physonectae
1
5 2 L-..-J
L-..-J
Apolemia uvaria Agalma elegans Agalma okeni
496 SiphQnophorae
Upper view of nectophores

3 0.5
Erenna richardi
Bargmannia Cordagalma cordiforme

erongata
Halistemma rubrum Marrus antarcticus

Halistemma striata
3.14
Pyrostephos vanhoeffeni
Nanomia bijuga
~
Siphonophorae 497
U pper view of nectophores

5
l-...J
Athorybia rosacea
3.16
Physophora hydrostatica
Me/ophysa me/o'
Forska/ia edwardsi
2
Forskalia /euckarti
498 Siphonophorae
Bracts of physonect siphonophores
3.22
1
l.--J
2
2
l--.-J L-!
Agalma elegans Agalma okeni
Apoler!Jia
uvana
3.25
0.2
~
Erenna richardi Halistemma
rubrum
Cordagalma
cordiforme
3.29
3.30
2
L-J 0.5 t2J L-J
1
Marrus L--..J
Pyrostephos Athorybia
antarcticus Nanomia bijuga vanhoeffeni rosacea
Siphonophorae 499
lateral views of calycophoran nectophores

of the families Prayidae and Sphaeronectidae
2
L.-....J 1
L--...J
Amphicaryon acaule Amphicaryon ernesti Amphicaryon
peltifera
Lilyopsis
ros ea
Maresearsia praeclara
Rosacea cymbiformis
3.36 5
L.-....J
Praya dubia
10
L---..J 3.37
Praya reticulata
3.39 Nectadamas
5 diomedeae
L--...J
Rosacea plicata
500 Siphonophorae
lateral views of calycophoran nectophores

of the families Prayidae and Sphaeronectidae
5
L--...J 3.41
2
Nectopyramis natans ~ 3.42 3.43
1
L---l
Nectopyramis thetis
Sphaeronectes
gracilis
Bracts, some with gonophores or special nectophores,

of Calycophorae of the families Prayidae and Sphaeronectidae
1
l---..J Praya dubia
0.5 Maresearsia 1
L-....J
Amphicaryon praeclara
acaule Lilyopsis rosea
Praya reticulata
Rosacea cymbiformis
5
~
Nectadamas diomedeae
3.50 Rosacea plicata c:=:>

Siphonophorae 501
Bracts, some with gonophores or special nectophores,

of Calycophorae of the families Prayidae and Sphaeronectidae
Nectopyramís natans Nectopyramís thetís Sphaeronectes

gracílís
Nectophores of Calycophorae of the family Hippopodiidae
2 Vogtía g/abra
Híppopodíus
hippopus
Vogtía pentacantha
5 5
Vogtía spínosa Vogtía serrata
502 • Siphonophorae
lateral views of anterior nectophores of Calycophorae of the

diphyid subfamily Sulculeolariinae
Suleuleolaria Suleuleolaria Suleuleolaria Suleuleolaria Suleuleolaria

biloba ehuni mono/ea quadrivalvis turgida

diphyid subfamilies Diphyinae and Giliinae,
with the exception of the genus Lensía
Chelophyes
appendieulata Oimophyes
aretiea
Chelophyes Oiphyes.
eontorta Oiphyes
antaretiea ehamissonJ~
Siphonophorae 503

with the exception of the genus Lensía
Eudoxoides Eudoxoides Muggiaea

Oiphyes
dispar mitra spiralis atlantica
Muggiaea Muggiaea Cilia

bargmannae kochi reticulata
504 Siphonophorae
Bracts, some with gonophores, of Calycophorae of the

with the exception of the genus Lensia
1
L--....J Diphyes
Diphyes bojani
antarctica
Chelophyes Dimophyes
appendiculata arctica
3.84
0.5
Muggiaea atlantica
Eudoxoides
sp iralis
1
l.-.---J
Eudoxoides Gilia reticulata
Diphyes
dispar mitra
Siphonophorae 505
lateral views of anterior nectophores of the

diphyid genus Lensia
Lensia
aehilles
2
L--...J
Lensia
Lensia
eampanella eonoidea
Lensia Lensia
fowleri hardy
Lensia
eossaek
Lensia leloupi
Lensia havoek Lensia hotspur Lensia hunter
c=:;>
506 Siphonophorae
lateral views of anterior nectophores of the

diphyid genus Lensia
3.99
1
l..---..J
Lensia
Lensia subtilis Lensia
meteori Lensia
mu/ticristata subti/oides
Lensia ajax
Lensia exeter
3.104 lO
,'l,
3.105
Lensia grima/di Lensia hosti/e Lensia /e/ouveteau

Siphonophorae 507
Bracts, some with gonophores, of the diphyid genus Lensia
Lensia subtilis
Lensia
campanella Lensia fowleri
Lensia subtiloides
Lensia conoidea

family Clausophyidae -
3.113
Clausophyes
galeata
Clausophyes
moserae
Chuniphyes multidentata
Chuniphyes
moserae
Clausophyes laetmata
508 Siphonophorae

family Clausophyidae
Clausophyes Crysta lIoph yes Heteropyramis Heteropyramis

tropica amygdalina crystallina maculata
Bracts of Calycophorae of the families

Clausophyidae and Sphaeronectidae
3.120
~ ~
Heteropyramis
Chuniphyes multidentata maculata
Heteropyramis crystallina
Siphonophorae 509
Ventral (a) and lateral (b) views of anterior nectophores of Calycophorae

of the family Abylidae
2
L......J
Abyla bicarinata Abyla haeckeli
3.127
.. ' ..
' ..
1
L--..l
Ceratocymba ..... . ",":.
Abyla trigona dentata "
::
:':
"
.;.
1
L-.-.....J
Ceratocymba
leuckarti
1
1
~
l.-.-...J
A bylopsis
tetragona
Abylopsis
eschscholtzi
¿
Enneagonum
hyallnum
Ceratocymba
sagittata
510 Siphonophorae
Bracts of Calycophorae of the family Abylidae
3.134
3.135
Abyla sp.
Ceratocymba
dentata Ceratocymba
leuckarti
3.136
3.137
Abyla Abyla tetragona

eschscholtzi
Ceratocymba
sagittata
Bassia bassensis Enneagonum hyalinum

Lensia zenkevitchi
Siphonophorae 511
• Abylopsis eschscholtzi (Huxley, 1859) (Fig. Acknowledgements

3.129,3.137). Dorsal and ventral facets ofanteri-
or nectophore relatively more regularly pentagon- Drs Herrnes Mianzan and Claudia Girola (INlDEP,
al and of nearly equal size. Strongly serrated rid- Mar del Plata, Argentina), and Dr. Mark Gibbons
ges. Lateral radial canals on nectosac without (Univ. of Cape Town, South Africa) kindly allowed
ascending loop. Posterior nectophore less than me to use some of their unpublished data.
twice as long as wide, with relatively large apical
apophysis. Dorsal facet of cuboidal bract forrns a
regular pentagon; apico-Iateral facets rectangular. Suggested readings
• Abylopsis tetragolla (Otto, 1823) (Fig. 3.130,
3.138). Dorsal and ventral facets of anterior nec- Bigelow H.B. 1911. The Siphonophorae. Mem. Mus.
tophore relatively less regularly pentagonal. Ridges Compar. Zool., Harvard ColI., 38: 173-402.
less strongly serrated. Lateral radial canals on nec- Kirkpatrick P.A., Pugh P.R. 1984. Siphonophores and
tosac with ascending loop. Posterior nectophore 3 Yelellids. Synopses of the British Fauna, 29:1-
times as long as wide. Two conspicuous basal 154.
teeth. Dorsal facet of bract elongate and less regu- Mackie G.O., Pugh P.R., Purcell lE. 1987. Siphono-
larly pentagonal; apico-Iateral facets trapezoidal. phore biology. Advances Mar. Biol., 24:97-262.
Pages F., Gili J.-M. 1992. Siphonophores (Cnidaria,
Genus Bassia L. Agassiz, 1862 Hydrozoa) of the Benguela Current (southeastem
Monotypic genus for Bassia bassensis. Atlantic). Scientia Marina, 56 (Supl. 1):65-112.
Bassia bassellsis L. Agassiz, 1862 (Fig. 3.131, Totton A.K. 1954. Siphonophora of the Indian Ocean
3.139). Anterior nectophore without an apical diver- together with systematic and biological notes on
ticulum to the somatocyst, and with the hydroeciwn related specimens from other oceans. Discovery
not extending below the basal facet. Posterior nec- Rep.,27:1-162.
tophore with 4 ridges ending in short basal teeth. Totton A.K. 1965. A Synopsis of the Siphonophora.
Bract with median apical ridge. Phyllocyst is a long British Mus. Natur. Hist., London, pp. 1-230.
tube, swollen apically, without apico-lateral bran-
ches. This species somewhat resembles Abylopsis Note added in proof
eschscholt::.i but there are c1ear differences. There is
usually a bluish tinge to the ridges of all parts. Lellsia zellkevitchi Margulis, 1970 (Fig. 3.141).
Anterior nectophore with variable number of often
Genus EllneagOIl111ll Quoy and Gaimard, 1827 incomplete longitudinal ridges. Hydroecium extends
Monotypic genus for Enneagonum hyalinum. to or above ostial level and is open ventrally. Large
Elllleagolllllll hyalill1111lQuoy and Gaimard, 1827 ovate, often indistinct, somatocyst. Inner margins of
(Fig. 3.132, 3.140) [=Enneagonut1lsearsae Alvariño, mouth plate with distinct but variable coglike serra-
1968]. The large, pyramidal anterior nectophore is tions; outer margins rounded. Posterior nectophore
the only one developed. The conical somatocyst is and eudoxid stage not known. Rare species only
situated above the hydroecium, and extends to a known from tropical waters.
greater height than the nectosac. The bract is
cuboidal, with slightly concave facets. Swollen
somatocyst with 2 lateral and an apical processes .
•••
Fig. 3. Illustrations of the species. All scales are in mm unless otherwise noled.
Figure sources: after Bédot (1895): 3.35. after Bigelow (1911): 3.7,3.23,3.31,3.36,3.37,3.44,3.62,3.72; from Bigelow (1911): 3.17; from
Bigelow (1931): 3.18; after Chun (1885): 3.46; after Leloup (1934): 3.120; after Moser (1925): 3.106, 3.107; after Russell (1938): 3.85; after
Sears (1953), by courtesy ofthe Museum of Comparative Zoology, Harvard University: 3.123-3.140; after Tollon (1965): 3.6,3.13,3.15,3.21,
3.27,3.30,3.32,3.33,3.38,3.39,3.61,3.64,3.66,3.70, 3.74, 3.75, 3.76, 3.78, 3.81, 3.82, 3.83, 3.84, 3.87, 3.88, 3.89, 3.90, 3.92, 3.93, 3.97,
3.102,3.104,3.105,3.109,3.110,3.111,3.112,3.121.
Cubomedusae and Scyphomedusae
Herll/es W Mial/:::an and Paul F. S. Cornelius
Introduction are distinguished by having innumerable mouthlets on

the mouth-arms instead of a single central mouth. The
The Cubomedusae and Scyphomedusae, classes of Cubozoa (Fig. lA) or box jellyfish constitute a dis-
CnidarÍa (formerly Coelenterata), include most of the tinctive group of some 15 species, the Cubomedusae,
larger solitary marine invertebrates. Their large swarms several of which sting Man severely. The medusa
sometimes greatly affect the marÍne environment. Yet, stages ofthe great majority ofboth major groups have
paradoxically, for most regions ofthe world there is no probably by now been described.
reliable taxonomic account ofthem, and little is under-
stood about their place in marine cornmunities. Reproduction is seasonal except in the high tropics.
Although most species are dioecious, a few are her-
All Cubomedusae and Scyphomedusae ha ve a rela- maphrodite, typically being protandrous or protogy-
tively thick mesoglea or middle-layer which gives nous. Some are slightly paedomorphic, becoming sex-
them bulk and provides some skeletal support. The ually mature before all asexual adult characters have
popular name "jellyfish" reflects the prominence of developed (e.g. Chrysaora; see Agassiz and Mayer,
the mesoglea. The name has no accurate zoological 1898; Russell, 1970). Some species, especially the
definition, and though these two groups might be con- larger cubomedusae, and perhaps some deep-water
sidered the "true" jellyfish, the name is sometimes scyphomedusae, evidently live for several years, but
applied also to hydromedusae, siphonophores, and most live for just a few weeks or months. The life
some other gelatinous plankton such as ctenophores cycles typically include a small polypoid benthic
and salps. Both the present groups are characterized stage, the scyphistoma .• Small medusoid larvae, or
by a radial syrnmetry. In fact, the scyphomedusae are ephyrae (Fig. 5.15c and 5.20c), strobilate offfrom its
the largest naturally-occurring perfectly syrnmetrical, upper end. Almost all scyphomedusae and cubomedu-
non-bilateral, objects known from our planet. Their sae have an altemation of stages between this asexual,
syrnmetry and the delicate nature of some of their sessile, benthic stage and a swimming, sexual plank-
structures give some of them great beauty. Many spe- tonic medusastage. The ephyrae each develop into a
cies are deeply or brightly colored, again mostly in a new medusa which is either male or female. A single
radial pattem which adds to their beauty. Color can bé scyphistoma will produce several to many medusae.
helpful in routine identification, but accurate system- Some scyphistomae are known to live for many years,
atics and taxonomy should rely on anatomical details. perhaps a cornmon pattem, and during their lives may
liberate hundreds of ephyrae. It is a feature of some of
The approximately 200 species of scyphozoans are the larger scyphomedusae, and of some cubomedusae,
exclusively marine. There are both pelagic (occurring that those of large size may have been released hun-
at all depths) and benthic species. The Scyphozoa dreds, maybe thousands, ofkm from the area in which
comprises Stauromedusae, mostly small, benthic they eventually reach reproductive sta te. Much pan-
polypoid medusae; Coronatae, mostly mid- to deep- oceanic genetic interchange must occur, and most of
water forms having a deep furrow (coronal grove) the species are extremely widespread in geographic
dividing the exumbrella into inner and outer zones range. Probably most scyphomedusae, and all cubo-
(Fig. lB); Semaeostomeae (Fig. 1C-E), typically medusae of known life-cycle, have a benthic polyp
large, conspicuous, inverted dish-shaped medusae, stage. The life cycles of some coastal forms are
most abundant in shallow coastal waters (the genera known, but those of many are unstudied and the ben-
Desmol/ema and Chrysaora can reach more than 1 m thic stage of most is unknown. In a few coronate scy-
in diameter, and Cyanea was once reported at 2 m); phomedusae the medusa stage is vestigial or entirely
and Rhizostomeae (Fig. lF, G), typically roughly suppressed and the scyphistoma is dominant in th.elife
hemispherical, sometimes large, mostly warm-water cycle, paralleling an evolutionary trend cornmon and
coastal, filter-feeding forms, most species of which well documented in the leptolid hydrozoans (hydroids
South Atlalltic Zooplankton.

© 1999 Backhu)'s Publishers. Leiden, The Netherlands
514 Cubomedusae and Scyphomedusae
and hydromedusae: see reviews in Boero and are suspected of uptaking dissolved organic matter
Bouillon, 1987; Comelius, 1992). In contrast, in at (Ferguson, 1988), processes which ifwidespread may
leas! one species of Pelagia the scyphistoma stage of enable individuals to survive long periods between
the life cycle is suppressed and bypassed, enabling successful prey-encounters.
them to range across the open ocean.
Gelatinous plankton, including Scyphomedusae, have
The anatomy of jellyfish, and of all cnidarians, is been regarded as a "dead end" in pelagic marine
mostly simple (refer to literature cited above; also trophic webs (Greve and Parsons, 1977). However,
Harrison and Westfall, 1991, for histology). This some research has demonstrated the importance of
finds exception in their highly complex subcellular gelatinous prey for several fish groups and some high-
organelles, the nematocysts. Most function to inject er vertebrates (Haedrich, 1967; Arai, 1988; Ates,
toxin, or "envenomate", prey or enemies through long 1988, 1991; Mianzan et al., 1996; Mianzan et al.,
piercing tubular threads, but some threads entangle 1997). Man himself can also be regarded a scyphom-
and some are sticky. The mechanisms of discharge edusa consumer. More than 10,000 tons of
and penetration, and the toxins, are among the most Stomolophus nomurai auct. and 1,000 tons of
complex of subcellular structures of any phylum (- Rhopilema rhopalophora and Lobonema smithii are
review of their effects on Man in Williamson et al., fished for human consumption every year from the
1996). The long survival of the Cnidaria may result China Sea, Japan and several other southeast Asian
largely from the evolution of nematocysts and their regions (Omori, 1978). In 1981, 40 million $ US
powerful toxins, and has given time for a wide range worth of scyphomedusae were imported by Japan
of nematocyst types to evolve. These are widely used alone (Omori, 1981, in Arai, 1997). In Taiwan Aurelia
in systematic diagnosis in some cnidarian groups, but aurita (the moonjelly) is al so eaten (Shulman, 1995),
not so much in the present ones. and various other species are taken in small quantities.
Cubozoa and Scyphozoa are essentially camivorous. Local populations of some Scyphomedusae reach
They feed on swimming animals through a variety of high den sities in many regions ofthe world, occasion-
fishing techniques. Many typically employ the pul s- ally causing problems for Man's interests. During cer-
ing movements of the bell simultaneously for move- tain periods they even dominate the zooplankton bio-
ment and to provide presumably optimal vortices mass (M6ller, 1978, 1980a; Shushkina and Musayeva,
bringing prey into contact with the nematocyst-bear- 1983; Papathanassiou et al., 1987) in both volume and
ing surfaces (Costello and Colin, 1995). These may be organic carbon content. Aggregation· seems to be a
the marginal tentacles, the mouth arms, and even the widespread phenomenon for "jelly plankton" (Arai,
upper surface of the bell in one species or another. 1992). It has even been suggested that blooms might
Some species hang essentially motionless in the be an essential part ofthe life history for some species
water. Medusae of all kinds are typically capable of (Boero, 1991); but populations of high density may
consuming a great variety of prey including copepods, quickly consume all the prey in a volume of water,
small crustacea, fish eggs and larvae (e.g., leading to starvation of the medusae. Nevertheless,
Feigenbaum and Kelly, 1984; Alvariño, 1977, 1985; swarming for gamete release might still be hypothe-
Fancett and Jenkins, 1988); and interspecific and sised, and some cubomedusae certainly accumulate
intraguild predation are al so recorded (Arai and for the copulatory reproductive process that is unique
Jacobs, 1980; Mianzan, 1986b; Purcell, 1991; review to them in the cnidarian phylum.
in Arai, 1997). AIgal symbionts (zooxanthellae),
present in a few species, secrete organic compounds Occasional swarms of great density have a notorious
typically used as metabolic substrates, and which are effect on many human economic activities such as
assimilated by the host medusa (review in Arai, tourism. fisheries and various coastal industries.
1997). A number of species occurs sometimes in Cnidarians are the most frequent marine envenomators
remote island situations, indicating that some may be (stingers). More than one hundred species of cnidar-
long enough lived to cross the great oceans even if ians are now regarded as venomous to Man, including
feeding opportunities are reduced. Some species are some 25 of the larger jellyfish, and the list is continu-
known to metabolise existing tissue (Thill, 1937) and ously expanding (Kokelj et al., 1993; Williamson et
Cubomedusae and Scyphomedusae 515
Cuboid bell Dome-shaped bell

(Cubomedusae) (Coronatae)
~-"'------
Hemispherical bell
(Rh izostomeae)
without scapulets aboral side

exumbrella -
- scapulet
mouthlets
4)
pnmary
mouth
rhopalium
marginal
marginal ~ ~ lappet
cleft ~ ~
C:o. ~ ~J
pillar
~
stomach
~ @
Cubornedusae and Scyphornedusae 517
al., 1996). Swarms of Pelagia noctiluca in the Clogging of the refrigeration-plant intake ducts of
Mediterranean Sea affected tourism for several years ships has required SCUBA divers to cIean the pipes
in the 1970s and 1980s (Rottini-Sandrini and Stravisi, (Mianzan, 1986b, 1989a, Puerto Madryn, Patagonia,
1981; Aubert, 1984; Bernard, 1984; Maretic, 1984; Argentina).
F.E. Russell, 1984; Rottini-Sandrini and Avian, 1991),
but the prevalence of that species seems to wax and Some medusa swarming and its consequences, as out-
wane over the decades. The sea nettle Chrysaora quin- lined above, may result from Man's activities. The
quecirrha is considered a pest in Chesapeake Bay destruction of a major geographic barrier by the build-
(Cargo and Schultz, 1966, 1967; Schultz and Cargo, ing of the Suez Canal, and the transport of scyphozoans
1969). The serious, even letha1, sting of the box-jelly- in ship ballast waters, have be en invoked to explain the
fish Chironex jleckeri of Australia and some regions to Mediterranean invasion by Rhopilema rhopalophora
the north, and the slightly less dangerous (see Come1ius, 1996; Galil el al., 1990; Lotan et al.,
Chiropsalmus quadrumanus which occurs pan-tropi- 1992, as R. nomadica). Cassiopea andromeda entered
cally, are notorious (Williamson et al., 1996; see also the Mediterranean from the Red Sea much sooner after
Williamson, 1985; Burnett, 1991; Fenner, 1991) but the Canal was opened (review in Spanier and Galil,
some other species also sting Man severely. 1991); and populations of Phyl/orhiza punctata have
become briefly established locally at isolated localities
The fishing industry is sometimes affected by jellyfish far from its original Australasian and southeast Asian
swarms (Rottini-Sandrini el al., 1984; M6l1er, 1984a, distribution (Brazil, California, Mediterranean,
b; Mianzan, 1986b; Hay el al., 1990). Jellyfish have Hawaii; review in Corne1ius, Si1veira and.Mianzan, in
probably been preying on fish and their food resourc- prep.). Eutrophication of semi-encIosed areas has also
es for some 200 mili ion years. They still remain com- been considered the cause of some scyphozoan bloom s
petitors for the same plankton resources that are con- (Shushkina and Musayeva, 1983; Papathanassiou et
sumed by many commercial fish (Parsons and Seki, al., 1987; Legovic, 1991).
1995). Overfishing seems to have resulted in the
increase of some scyphomedusa species. In the No cubomedusa or scyphomedusa is known to be par-
Mediterranean Sea, Avian and Rottini-Sandrini asitic, but a few parasitic species of several other
(1988) and Legovic (1991) concIuded that increase of phyla are known to infest them (Laval, 1980).
the Pelagia noctiluca popu1ation followed increased Amphipods of the genus Hyperia can debilitate a
pressure on fish stocks. In the Black Sea, Aurelia 1arge medusa by gradually consuming it from within.
aurita increased from an estimated stock ono million Mostly, however, medusae of these groups are not-
during the 1960s to 400 million during the 1980s, put- ably free of parasites.
ting pressure on pelagic fisheries by competition with
fish stocks for food and by predation on fish 1arvae For the present review, the cIassification adopted
and their food (Shushkina and Musayeva, 1983; broadly follows that of Kramp (1961) with minor
Bingel, 1991; Niermann el al., 1994; Mutlu et al., revisions but is considered provisional and several
1994). In the Benguela Current off Namibia, increase systematic questions await evaluation. Two interest-
of Chrysaora hysoscel/a populations from the 1960s ing and well-founded genera are still best referred to
onward, from 5 million ton s to 30 million tons, has Incerlae sedis at the end (Tessera, Tetraplatia).
been concurrent with a decrease of the pilchard
(Sardinops ocel/ala) (Fea ron el al., 1992).
Methods
Cooling-water intake ducts of power stations in all
parts of the world are occasionally cIogged by swarms It cannot be stressed too strongly that several jellyfish
of large, coastal species (e.g. M611er, 1984a; Werner, sting humans severely. Special care should be taken to
1984, Baltic Sea; Rajagopal el al., 1989, India). avoid this. Being a serious student do es not diminish
•••
Fig. 1. General characters used for separation of the orders of Cubomedusae and Scyphomedusae.
~
the danger - rather, the danger increases since it is groups (see "Ctenopho~a" in this volume), the finding
natural to become blasé. A sting in the eye from a usu- of new species is still possible.
ally harmless species can be extremely serious and
might permanently damage human sight. Techniques Fixation and preservation are not a problem for
for avoiding stings are mentioned below, and should Cubozoa and Scyphozoa. 5% formaldehyde solution
be taken very seriously. is generally suitable for preserving most of the small
medusae, but at least 10% solution is initially needed
Extremely important. With the most venomous spe- fOLlarge specimens owing to the high percentage of
cies, eyes should be protected by spectacles or gog- water in their mesoglea.
gles. Avoid splashes to the eyes. Do not rub your eyes
after handling medusae - even apparently harmless Storage may present a problem. Other small plankton-
species - since the nematocysts can readily penetrate ic groups are well represented in museum collections.
the soft tissues in and around the eyes. Rubber gloves, For Scyphomedusae, typically only small specimens
long sleeves and leg-covering are necessary with the can be found in such institutions. Large species are
most dangerous species, as well as socks and closed often neglected for storage due to the difficulties of
footwear. A gallon (5 litres) of vinegar should be having the appropriate containers, and additional
available for use in emergencies, to kill undischarged space to be kept in collections (Mianzan, 1986b).
nematocysts (but vi negar should not be used with the
siphonophore Physalia sp.). For transport, specimens should be placed in a strong
screw-top plastic jar which is then filled to the top with
Most of the records of Scyphozoa and Cubozoa from fluid (usually, formaldehyde solution). A sink is filled
South America are based on single findings of speci- with tap water. The filled jar is placed in the sink, and
mens on coastlines or on the sea-surface, rather than on the lid quickly placed on it so that only little dilution
plankton sampling surveys (Vannucci, 1951, 1954, occurs and so that no air is included. The jar can then
1957a; Vannucci and Tundisi, 1962; Mianzan, 1986b, be tipped, shaken, and even dropped, without damage
1989; Olagüe el al., 1990). This group is not usually to the specimen inside. If a perfectly shaped specimen
well sampled with the standard available plankton nets. is not needed, or ifweight is limiting (as for postage),
Conventional plankton surveys are inadequate for the following can be done. A well-preserved specimen
obtaining these large animals, since current plankton is placed in a self-sealing plastic bag and excess fluid
nets (Bongo, Nakthai, Multinet) usually sample small tipped off. Some fluid should remain, since air bubbles
volumes qf water (Omori and Hamner, 1982; Larson, under a specimen can hinder later observation. The bag
1986; Arai, 1988, 1997). Estimates of scyphozoan and is then lowered into a sink ofwater slowly, so that the
cubozoan abundance and biomass are also biased for sides of the bag are pressed gently by the water pres-
the same reasons. The use of midwater nets (IKMT, sure, excluding air and gripping the medusa firmly.
RMT, METHOT) makes possible the obtaining of When all the air is out, the top ofthe bag is sealed. The
acceptable information on the distribution and abun- bag can be placed in another, similar bag, to avoid risk
dance of species (e.g., Larson, 1986; Pages ef al., of leaks; a label placed outside this second bag, and the
1992), but still with some restrictions for larger species. whole placed in a third bag, and even a fourth, for near-
complete safety from leaks. The whole package can be
As with other zooplankton groups, analyses of scy- placed in a plastic kitchen-ware container with lid, and
phozoan and cubozoan vertical distribution pattems the lid taped. Material should be unpacked immediate-
are usually made with the aid of vertically stratified Iy after arrival to minimize distortion.
midwater plankton tows. However, the nets usually
used have no opening-closing mechanism so precise
depth data can be missing (Larson, 1986). Geographic and vertical distribution
SCUBA enables some kinds of observation to .be Studies on Cubo- and Scyphomedusae in the South
made on these large animals. For mid-water foons, Atlantic Ocean mainly directed to distribution pattern
submersibles have been used but medusa encounter is and zoogeography are scarce. Kramp (1961) consti-
sometimes rare (Larson ef al., 1991). As in other jelly tutes the most recent available source that included
the study area, and Kramp (1970) presented a review itudes (in particular the families Cyaneidae and
on Rhizostomeae of the world with a zoogeographic Ulmaridae), through the region influenced by the
emphasis. Larson (1986) surnmarized information for Malvinas (=Falkland) Current (cold waters up to 35°S).
the antarctic seas, including the South Atlantic up to The Coronatae, with a few exceptions, are oceanic, so
300S, but based mainly on pelagic species, inhabitants they are not cornmon along the coast. The wide South
of the outer region of the continental shelf and open American continental shelf creates good conditions for
sea. Pages et al. (1992) and Mianzan (l986b, 1989) the development of populations of coastal species.
attempted to clarify the state of formal knowledge in
the southeastern and southwestern Atlantic Ocean, Conditions are different in the southeastern Atlantic
respectively. Ocean. African scyphomedusa surveys include an
important number of coronate species - a conse-
In the present account, data on the distribution of each quence ofthe narrow continental shelf ofwest Africa.
species (Fig. 2) are derived from a compilation of all The influence of the cold waters of the Benguela
available literature records and personal research data, stream might explain the presence of some of the
so some new records are included. semaeostome species observed, even though the low
temperatures and constantly fluctuating physical envi-
As shown in Fig. 2, some of the species have wide ronment nearshore may account for their paucity
distributional ranges, following the continental shelf (Gibbons et al., 1992). The equatorial and oceanic
in the neritic forms or occupying most of the Atlantic waters genera te the necessary environmental stability
Ocean in the oceanic ones. However, most of the spe- to allow high diversity. It is also likely th'Ú some spe-
cies ha ve seldom been found, so instead ofusing lati- cies from the Indian Ocean pass to the Atlantic, like
tudinal range tables, we prefer to present the informa- Tamoya haplonema (see M6ller, 1980b; Cornelius,
tion as distribution maps. 1997b), advected by the Aghulas current (see
"General biological features of the South Atlantic" in
Several species reported from the equatorial Northen this volume).
Hemisphere (Paraphyllina ransoni, Linuche unguicu-
lata and Ceplzea cephea) are included due to the high Abundance and biomass were discussed by Larson
probability of finding them in tropical waters of the (1986). South Atlantic oceanic scyphomedusae, main-
South Atlantic. However, others reported from loca- ly the coronates Periphylla periphylla, Atolla chuni
tions far north of the Equator and not included in the and A tolla vl'yvillei, seem to increase in numbers of
present account, such as Cassiopea frondosa and individuals and biomass with increasing latitude from
Cassiopea xamachana (Caribbean Sea), should be 300S to 800S. This implies that these species are not
mentioned. randomly distributed, but that they tend to occur in
response to factors such as hydrographic conditions
Endemism is rare in Cubo- and Scyphomedusae in the and prey distribution.
South Atlantic, and many ofthe species could be con-
sidered cosmopolitan. Other areas show strong Vertical distribution ofscyphomedusae is not random.
endemism, as the Indo- West Pacific for the It is affected by light, pressure, presence ofprey, stage
Rhizostomeae (Kramp, 1970). The Atlantic, however, of life cycle and temperature, salinity and oxygen gra-
has only a few species recorded exclusively there. dients (Larson, 1986; Fearon et al., 1992; Arai, 1997).
The Coronatae are well studied and the vertical pro-
As with other gelatinous forms (e.g. ctenophores), files of total abundance in antarctic and subantarctic
coastal species from the southwestem Atlantic Ocean waters show that the peaks of their populations are
fit well with local biogeographic schemes proposed in typically between 500 and 1000 m depth (Larson,
the literature (Mianzan, 1989a; Mianzan et al., 1988; 1986). Some species are capable of strong vertical
Ramírez et al., 1990). Warm-water forms, like the displacement. Atolla vanhoe./Jeni migrates at least 200
Cubo medusa e and Rhizostomeae, tend not to penetrate m upward (Arai, 1997), and Periphylla periphylla
higher latitudes (35° and 39°S, respectively). However, seems to have a diel migration range of at least 500 m
the Semaeostomeae dominate the fauna in southem lat- (Mianzan, unpublished observations).
2.1 Wa;¡ •• 2.3

I 2.21
•
\
Tamoya haplonema Carybdea alata Tripedalia cystophora
2.4 2.5 2.6
•
•
Chiropsalmus quadrumanus Chirodropus gorilla
2.7 ••
• •
•
•••
Atolla oarva Atolla vanhoeffeni
2.11 2.12
•
•
•
?
Linuche unguiculata Nausithoe rubra Nausithoe aurea

2.13 2.15
• •
• • ?
?Paraphyllina •. • • •
••• •
• .?
Desmonema gaudichaudi
.? .2.23
ransoni 2.17
2.20
2.21
2.24
Chrysaora
Chrysaora
Drymonema
Chrysaora
quinquecirrha
hysoscella
Diplu/marisdalmatinum
/actea
antarctica
Pelagia noctiluca
••
Deepstaria enigmatica
2.25 2.26 2.27

•• •• •
• ••
•••
• •
I Poralia rufescens
Discomedusa lobata
• Phacellophora camtschatica
2.28 2.29 2.30

• •
•
•
•
• Stygiomedusa gigantea
•••••
•• • Cephea cephea Phyllorhiza punctata
2.31 2.32 2.33
, I ••1
• Stomolophus meleagris
• ••• • II
Rhizostoma pulmo I I
•
Lychnorhiza lucerna I Catostylus tagi
2.35 Tetraplatia volitans
• fupilema inexpectata
2.36
••
•
Tessera gemmaria
Taxonomy Scyphomedusae, including biological and cornmercial

aspects; but taxonomic and most medical aspects and
There have been several reviews of aspects of the bio- the Cubomedusae were omitted. Lack of well-
logy of the two groups covered, but rather few and researched identification literature has held up studies
some of the older ones are still useful taxonomically. of all kinds on the group in most regions of the world,
There is no previous South Atlantic guide. Mayer and this continues to be a problem. We anticipate that
(1910) is the most recent world monograph, though the present synopsis will prove to be accurate for the
Cornelius is currently preparing an updated one. South Atlantic species. The species limits of most of
Kramp (1961) listed, somewhat uncritically, almost all those included here seem adequately understood,
the nominal species and provided an exhaustive com- which could not have been claimed a decade or so ago.
pilation of literature and taxonomic opinion of the pre- Though some species might be added to the faunallist,
vious 50 years. F. S. Russell (1970) reviewed the they may be rather few. Some nomenclatural and taxo-
British scyphomedusa species with great clarity and his nomic revision is inevitably incorporated into this
book has wide relevance despite its narrow geographic chapter and will be explained in future publications.
limitation. Werner (1984) and Franc (l995a, b) both
provided text-book reviews of both groups. Larson Glossary of terms applied to Cubomedusae and
(1986) provided an extensive account of the scypho- Scyphomedusae
medusae of the Southern Ocean. Important collections
of articles on swarming and blooms, especially of For explanations of additional terms for Scypho-
Pelagia in the Mediterranean, were provided in two medusae see Russell (1970), and for Cubomedusae
UNESCO symposia (Anonymous, 1984; UNEP, 1991; Southcott (1967) and Williamson el al. (1996); and
see also reviews on hydromedusa and scyphomedusa for both groups Wemer (1984, in German) and Franc
swarming by Arai, 1992 and 1997, respectively). (1995a, 1995b, in French). For terms conceming
Aspects of envenomation (stings) were surnmarised by hydromedusae see Bouillon (1995, in French; and the
F. E. Russell (1984), Halstead (1988) and Williamson chapter "Hydromedusae" in this volume) and
el al. (1996). The last is a valuable reference source on Cornelius (1995). Many aspects of the biology were
medical aspects and illustrates many species. Comelius covered by Arai (1997). Another glossary was provid-
(l997c) provided a dichotomous key to all the genera. ed by Stachowitsch (1992).
Probably the most useful book on the biology of the
group is that by Arai (1997), which surnmarised an Nematocyst terminology is mostly omitted and the
enormous amount of research data on all aspects of the reader is referred to Hessinger and Lenhoff (1988) for
••
Fig. 2. Geographic distribution of South Atlantic Cubomedusae and Scyphomedusae. Note: in many species the distribution is inadequately
known.
Sources: Carybdea alata: 4,14,25; Tallloya haplonema: 3,12,14,20,23,26,27,31; Tripedalia cystophora: 4,27; Chirodropus gorilla: 5,8,
14,23; Chiropsallllus quadrumanus: 4, 19,20,21,27; Atolla wyvillei: 5,7, 15,26,33,34,39; Atolla chuni: 7; Atolla parva: 15,39; Atolla van-
hoe.ffeni: 5, 7, 15, 39; Linuche unguiculata: 4, 8, 23; Nausithoe !"libra: 4, 7, 34; Nausithoe aurea: 29; Nausithoe prlnctata: 3, 20, 26, 34;
Paraphyllina ransoni: 16; Periphylla periphylla: 5, 7, 11, 14,34; Chrysaora hysoscella: 5, 14,23,25,26; Chl)'saora lactea: 8, 9, 10, 11, 12,
13,20,22,31; CIII)'saora plocamia: 4, 25, 40; CJII)'saora qllinquecirrha: 3, 5, 23; Pelagia noctiluca: 5,7,20,25,26,30,31,33,40; Deslllonema
chierchianllln: 9, 11,22,40; Desmonellla gaudichaudi: 1,7,9, 11,22,32,40; Drymonema dallllatinulll: 4, 9, 10, 11,20,26,31; Aurelia auri-
ta: 3, 9, 10, 11, 12, 13, 20, 22, 23, 24, 31; Deepstaria eniglllatica: 7; Diplullllaris antarctica: 7; Discollledusa lobata: 8, 14, 23, 26;
PhacellopllOra call1tschatica: 5,7,9, 11,23,26,32,34,40; Poralia nifescens: 7,26; St)'giollledusa gigantea: 7,40; Cephea cephea: 6, 23;
Phyllorhi=a punctata: 6, 17, 31; Lychnorhi=a lucerna: 3, 6, 9, 11, 12, 13, 18, 19, 20, 22, 31,40; Catostylus tagi: 26, 35, 36; Eupilellla inexpec-
tata: 14; Rhi=ostollla pulmo: 6,25; Stolllolophus lIleleagris: 6,9, 11, 19,20,22; Tessera gellllllaria: 3; Tetraplatia volitans: 14,37,38.
References: 1: Bellisio and Tomo (1974); 2: Comelius (1973); 3: Goy (1979); 4: Kramp (1961); 5: Kramp (1968b); 6: Kramp (1970); 7: Larson
(1986); 8: Mayer (1910); 9: Mianzan (1986b); 10: Mianzan (1989a); 11: Mianzan (l989b); 12: Mianzan et al. (1988); 13: Olagüe et al. (1990);
14: Pages et al. (1992); 15: Repelin (1964); 16: Repelin (1965); 17: Soares Moreira (1961); 18: Vannucci (1951); 19: Vannucci (1954); 20:
Vannucci (1957a); 21: Vannucci (1966); 22: Vannucci and Tundisi (1962); 23: Kramp (1955); 24: Móller (1980b); 25: Branch et al. (1994); 26:
Thiel (1966); 27: Morandini and Marques (1997); 28: Kramp (1959b); 29: Silveira and Morandini (1997); 30: Stiasny (1934); 31: Comelius et
al. (Ms.); 32: Montsalve (1996); 33: Stiasny (1940); 34: Fagetti (1973); 35: Ranson (1949); 36: Stiasny (1939); 37: Rees and White (1957a);
38: Rees and White (1957b); 39: Kramp and Blanner (1972); 40: this study.
references to their structure and classification. Circular canal (sometimes ring canal): the narrow
Histological terms were covered by Harrison and canal or duct around the perimeter of the umbrella
Westfall (1991). linking the ends ofthe radial canals. Not present in
all genera. See also intermediate canal.
Some of the terms listed, especially some older ones, Circular furrow (obs.): see coronal groove.
are little-used synonyrns, and this is indicated. Since Circular muscle: see muscle field.
the standard monograph on the Scyphomedusae is still Cleft, marginal and rhopalial: see marginal cleft.
Mayer (1910), many of the older terms are still unfor- Club: (1) a club-shaped extension (lateral or terminal)
tunately used by some authors. A few standard zoo- ofthe mouth-arm, typically differing noticeably in
logical and other scientific terms are included if their structure from it. In some genera they may be pad-
application in cnidarians differs from that in literature dle-shaped (Cassiopea), in others three-sided
on other groups. (Mastigias) or circular (some in Rhopilema) in
section; (2) (arch.) for example, as in sensory club,
Abbreviations perradial club - a rhopalium.
abbr.: abbreviation; adj.: adjective; arch.: archaic; n.: Coronal furrow: an unusual synonym for coronal
noun; obs.: obsolete or obsolescent; pl.: plural; sing.: groove.
singular; ssp.: subspecies (singular); sspp.: subspecies Coronal groove: a groove around the exumbrellar sur-
(plural). face of a medusa, typically over a zone of relative
thinness of the underlying mesoglea (jelly). In
Acraspedote (adj., obs.): (medusae) lacking a velum many species it delimits one region of the bell or
(thus the Scyphomedusae were formerly referred to disc from another. Most conspicuous in coronate
as the acraspedote medusae). See also craspedote. medusae.
Adradial: see perradial. Coronal muscle: circular muscle. See muscle field.
Anastomosing (adj.; n.: anastomosis): of a repeatedly Coronatae: one of the three main divisions of
branched structure, such as vessels in a gastrovas- Scyphomedusae.
cular network, in which some of the branchings Craspedote (adj., obs.): of medusae having a velum.
rejoin and fuse with others to form a (typically Thus, hydromedusae were formerly referred to as
irregular) network. This differs from simple the craspedote medusae. See also acraspedote.
branching with a succession of forks, in which Crenate (adj.): having low rounded peripheral lobes,
rejoining does not occur. separated by sharp but shallow notches; with the
Annular furrow (arch.): see coronal groove. profile of several inverted saucers placed edge-to-
Appendage:"a confusing term for a club or filament on edge. Crenulate, more widely used in hydroid lit-
the part of the scyphomedusa indicated. Hence erature, is synonymous.
mouth-arm appendage, mouth-disc appendage, Cubomedusae, cubomedusa(e), Cubozoa: the boxjelly-
terminal appendage, etc. fish. The Cubozoa, usually given the rank of class,
Arm disc: see mouth-arm disco includes just a single order, the Cubomedusae, and
Bell: the dome-shaped body of a medusa, excluding the terms can often be interchanged without altera-
both structures hanging down from it and the mar- tion of meaning. They correspond in rank to the
ginal tentacles, if present. There is no clear dis- Scyphozoa and the two included groups
tinction from disc, which is generally applied to Stauromedusae and Scyphomedusae. The spelling
the somewhat flatter bells, and the two terms are cubo medusa e is the English-Ianguage term, and has
often interchanged especially when species inter- the same scope as the taxon-name Cubomedusae.
mediate in shape are being described. The term Disc: see bell.
umbrella (n.; adj.: umbrellar) covers both. The Ectoderm (n.): see epidermis.
exumbrella, or exumbrellar surface, is that on the Endoderm (n.): see epidermis.
upper side ofthe bell or umbrella, the subumbrel- Entoderm (n.): same as endoderm, but strictly speaking
la the underside. The space enclosed by the bell is termed gastrodermis in the adult. See epidermis.
the subumbrellar cavity, this being filled with sea- Envenomation: the process (or eyent) of one organism
water in continuity with the sea itself, in cubomed- introducing venom into another. Colloquially, a
usae via the velarial aperture. sting.
Epaulette (obs.): see scapulet. outline is visible from above and in some also
Ephyra: the irnmature scyphozoan medusa released from below, and in many its shape is made easier
from the scyphistoma. to see by gonad tissue which lies along its wall.
Epidermis (n.): the outer tissue-Iayer of the adult cni- Gastric septum: in coronate medusae, one of four ver-
darian body-plan, the inner layer being termed tical septa joining the stomach wall to the subum-
gastrodermis. The terms ectoderm and endodenn brella. See also suspensoria.
have also been widely used for these layers but Gastric tentacles (in stomach of scyphomedusa, obs.):
with some danger of inconsistency. In many other see gastric cirri.
animal phyla the embryonic ectodermal and endo- Gastrodermis (n.): see epidermis.
dermal tissues can be shown to give rise to the cor- Gastrovascular: the gastrovascular system comprises
responding tissue layers in the adult, to which the the centrally placed stomach, the gastrovascular
terms ectoderm and endoderm can be validly pouches (=stomach pouches, gastrovascular sinus-
applied. In cnidarians the sexually-produced es), or radial canals and in many species their (gas-
embryo forms a planula, similarly having inner trovascular) anastomoses and the circular canals.
and outer layers which are correctly termed ecto- Genital ostium, g. papilla, g. pillar, g. porticus: in rhizo-
derm and endoderm. But the planula metamor- stome medusae, a genital ostium (pl., ostia) is a gap
phoses into a polyp with loss of identity of the between adjacent mouth-arm bases, proximal to
original tissue layers. Thus, during both metamor- their point of fusiono The basal regions of the
phoses, the integrity of the original layers may be mouth arms are in that region termed "pillars" in
lost, making the homology of the newly produced distinction from the ostia between. -The relative
layers unclear and the terms ectoderm and endo- widths of these pillars and gaps are diagnostic of
derm misleading. The terms epidermis and gastro- certain genera and species. Variously-shaped papil-
dermis are neutral in that they refer just to position lae (genital papillae) project into the ostia in the
of tissue layers in the (adult) animal and not to suborder Daktyliophorae from the subumbrella epi-
their embryonic origino See also mesoglea. dermis, partly closing the aperture. The genital cavi-
Exumbrella (n.), exumbrellar (adj.): see bell. ty onto which the ostia open was formerly known as
Exumbrellar wart: a nematocyst wart on the exum- the genital porticus (e.g. Mayer, 1910, p. 632).
brella. See nematocyst wart. Intermediate canal: a canal intermediate in position
Filament: in some rhizostomes, a narrow filamentous between the (marginal) circular canal and the
extension from the mouth-arm, typically densely stomach, in several genera of rhizostomes. Also
armed with nematocysts and assumed to be used termed intermediate circular canal.
in defence or for the stunning of prey. Both termi- Interradial: see perradial.
nal and lateral filaments occur, this varying with Inter-rhopalar: see rhopalium.
the species. See also club; gastric cirri. Interrhopalar canal: a radial canal not in a rhopalial
Frenulum: in cubomedusae, one of the four vertical radius. See also rhopalial canal.
mesenteries attached just inside the bell aperture Interradius (n.): see perradial.
and supporting the velarium. Intracircular canal network: see radial canal.
Gastral or gastric cavity: an unusual term for the Jelly: see mesoglea.
stomach. See also gastrovascular. Lappets: flaps on the margin of the scyphozoan disc,
Gastric cirri: short filaments, barely longer than wide, somewhat regular in arrangement; much varied in
located in groups (phacellae) on the stomach lin- size but typically from 8 to ca. 100 in number.
ings in some cubomedusae. Reportedly absent in There is confusion in the literature on some species
Caruha; may perhaps decompose or be digested about whether or not the lappets should be counted
away in specimens of other cubomedusae in which in pairs, with each pair comprising a single lappet
the preservative does not penetrate rapidly enough which is cleft in two (as is clearly so in Cyanea).
to prevent it. Gastric filaments occurring in Hence lappet-counts by some authors of some spe-
Scyphomedusae are typically longer. See also fila- cies may be double those of other authors. In at least
mento some rhizostomes the number of lappets increases
Gastric cross (arch.): in many rhizostomes, the cross- gradually with age (perhaps to a more or less con-
shaped outline of the stomach. In many species the stant maxirnum); but in some others, such as some
coronate medusae, it is more fixed, especially once essarily the sexual one (e.g. vertebrates). If just
development is complete. Rhopalial lappets (for- one asexual stage is interposed, as in Leptolida,
merly termed ocular lappets) occur one each side of Cubomedusae and Scyphomedusae, the term dige-
the rhopa1ial niche. In most species they differ in netic (two stages) can be applied. For detailed
size andlor shape to a varied extent from the account see Naumov (1960, 1969).
remaining or inter-rhopalar lappets (=velar lappets). Monodisc scyphistoma: a scyphistoma in which the
Main canal (obs.): same as intermediate, or mid-, cir- ephyrae develop singly, as in rhizostomes. In
cular canal. See radial canal. polydisc scyphistomae, characteristic of semaeos-
Manubrium (n.): a central, tubular, downward projec- tomes, several successively older ephyrae can be
tion ofthe subumbrellar surface bearing the mouth seen developing together, arranged like a pile of
terminally, its lumen being continuous with the plates, the most distal being the oldest. The
gastrovascular cavity (stomach). Greatly varied in spellings monodisk, polydisk, are American.
shape and length between species. Mouth arms: thick tentacle-like structures around the
Marginal cleft: the gap between two lappets, or mouth in scyphomedusae, typically four or a low
between two members of a pair. A rhopalial cleft multiple. In rhizostome medusae the four are fused
is one in which a rhopalium is situated. See also near the base and they immediately fork each in
lappet. two so that there appears to be eight. In semaeos-
Marginal ring canal: same as circular canal, peripher- tomes they are free along their entire length, and
al ring canal. See radial canal. four in all species. Also known as oral arms.
Marginal sense organ: rhopalium. Coronates and cubomedusae lack them.
Marginal tentacle: a tentacle situated on the edge of Mouth-arm disk: the region of the subumbrella to
the bello Not present in rhizostome medusae, nor which the mouth arms are attached.
in about four semaeostome genera. Mouthlet: in rhizostomes, a tiny opening of the gas-
Marginal vesicle: see statocyst. trovascular system on a mouth-arm, typically ca. 1
Medusa: a free-living stage in the cubomedusan and mm in diameter and surrounded by minute tenta-
scyphomedusan life-cycle, in most species swim- cles.
mmg. Mouth lip: a short projection from the rim of the
Medusozoa: the subphylum of Cnidaria comprising the mouth of a medusa.
two classes Hydrozoa (comprising the subclasses Mouth, primary: see primary mouth.
Leptolida, the hydroids, hydromedusae and related Muscle field (circular, coronal, radial): a region ofthe
forms, and Siphonophora, the siphonophores) and subumbrella having musculature. In most genera
Scyphozoa (comprising the Scyphomedusae, that is the fields are regular in shape and arrangement.
the large medusae such as those familiar in coastal Circular or coronal muscle-fields have the muscle
waters, and Stauromedusae, small, stalked, mobile folds (see muscle folds) arranged parpllel with the
but sedentary forms). The Medusozoa and bell margin, and in radial muscle-fields they are
Anthozoa together form the Cnidaria. arranged radially.
Mesoglea (n., formerly spelt mesogloea; adj.: meso- Muscle fold: a fold in the subumbrellar epidermis com-
gleal): in Cnidaria, typically an acellular, ge1ati- prising muscle tissue. Typically, many folds lie
nous but firm layer between epidermis and gastro- parallel and side-by-side, having the general
dermis. In polypoid forms usually functioning as appearance of corrugated paper. See also muscle
an elaborate basement membrane and as a flexible field.
skeleton; in medusoid forms the same in smaller Muscle strand (obs.): same as muscle fold.
organs such as tentacles and manubrium but Nematocyst warts: wart-like structures richly supplied
across most of the bell, of which it forms the bulk, with nematocysts, or "sting capsules", found on
typically much thickened. In most medusae it is the exumbrellae of many medusa species, and in
transparent and forms the jelly of jellyfish. other positions. Some are elongate, almost papil-
Metagenesis (n., adj.: metagenetic, sometimes meta- late (e.g. Pelagia flaveola and Lobonema smithii).
genic): altemation of a sexual stage in the life Cephea has relatively massive exumbrellar struc-
cycle with one or more asexual (vegetative) ones. tures of (somewhat) irregular shape inappropriate-
Monogenetic organisms have a single stage, nec- ly referred to as warts.
Niche, sensory; rhopalial niche: see rhopalium. Phacellus (pl.: phacellae): see gastric cirri.
Octant (n.): see quadrant. Pillar, genital: see genital ostium.
Ocular lappet (arch.): see lappet. Pit, sensory: rhopalial niche. See rhopalium.
Oral arms: see mouth arms. Planula (n.): a simple, ciliated or flagellated, dispersive
Ostium: see genital ostium. phase in the life cycle of most cnidarians; typically
Palmate: shaped roughly like the palm of a hand, i.e. about 1 mm long, roughly the shape of the sole of
comprising a central region and elongate struc- a slipper. Interpreted as a prolonged gastrula.
tures, such as tentacles or pedalia, inserted around Polydisc scyphistoma: see monodisc.
the edge. See also pedalium. Polyp (n.): in Cnidaria, the basic sessile individual,
Papillae: minute to small protrusions from the surface, contrasting with the mobile medusa.
for example fram the exumbrella. See also genital Porticus, genital: see genital ostium.
papilla; nematocyst wart. Primary canal: a confusing term for a radial canal in a
Paramere: see quadrant. perradius, better termed a perradial canal. See
Pedaliwn: (1) in cubomedusae, an extension of the radial canal.
lower edge of the bell at each comer, supporting Primary circular canal: a confusing term for the
the tentacle(s). In chirodropids they are palmate peripheral ring canal or, simply, ring canal. See
with a tentacle on each "finger"; but in carybdeids radial canal.
having more than one tentacle at each comer each Primary mouth (opening): in some rhizostomes, in
pedalium is separate from the others; (2) in coro- addition to the numerous minute mouthlets on the
nates, pedalia are "radial thickenings of the jelly mouth-arms there is a mouth-opening in the centre
on the peripheral zone of the exumbrella between of the subumbrella as in semaeostomes, termed a
the coronal groove and the bases of the marginal primary mouth opening. Whether it developmental-
lappets" (Russell, 1970, p. 27). See also palmate. ly represents the embryonic mouth is not implied.
Perradial, interradial, adradial: the perradial radii are One occurs for example in Stomolophus, and in that
the four main radii ofthe medusa, as though North, genus it may have been secondarily acquired so
South, East and West. In between (SE, NE, etc.) that the term "primary" may be misleading.
occur the four interradii; and in the radii between Primary ring canal: a confusing term for the peripheral
the others (ESE, ENE, etc.), the 8 adradii. ring canal or, simply, ring canal. See radial canal.
Cubomedusae are all tetraradiate and the types of Proboscis (n.): ofa scyphistoma, the mound on which
radii are clear in all species, but in some ofthe three the mouth is situated.
other groups some or many genera and families Quadrant (n.): of a medusa, a quarter of the bell and
have more than four radii and identification oftheir the associated structures. In medusae having four
homologies must be traced carefully. The following radial canals it comprises the segment between the
remarks thus apply essentially to adult forms which centre of one canal and the centre of the next.
are tetraradiate, which in all groups is regarded as Octant similarly refers to an eighth of the bell.
fundamental. The stomach pouches are taken to be Paramere refers to any such fractional segment.
homologous between the groups, and perradial in Radial canal: a canalleading between the stomach and
position. Thus in cubomedusae the stomach pouch- the periphery of a medusa. In a few species, some
es and rhopalia are on the perradii, and the pedalia radial canals originate from the periphery and end
are interradial; in coronates the stomach pouches blindly without reaching the stomach. The branch-
and the comers of the mouth are perradial, and the ing pattems of the radial canals are used in many
rhopalia may be perradial or interradial, depending scyphozoan generic diagnoses. In many species a
on the family; in semaeostomes the gastric pouches circular canal (=ring canal; sometimes termed pri-
and the comers of the mouth (prolonged into the mary circular or peripheral canal) links the outer
mouth-arms) are perradial, and the gonads inter- ends of the radial canal s at or near the margino In
radial; and in rhizostomes the stomach pouches are a few, an intermediate circular (or ring) canal
perradial, the gonads interradial, and the 8 mouth- (mid-circular canal) occurs linking points mid-
arms are regarded adradial. way along the radial canals (also termed the sec-
Perradial canal: a radial canal (see radial canal) lying ondary circular canal). In many species the radial
in a perradius. canals are partially linked laterally by an anasto-
mosing network of branches, more or less orderly in a rhopalar radius, as distinct from a tentacular
in arrangement, the intracircular canal network. In stomach-pouch, in a tentacular radius.
some species (e.g. Rhizostoma) the mid-circular Ring canal: see radial canal.
canal is more developed than the peripheral canal, Ring furrow: an unusual synonym of coronal groove.
and by some authors (e.g. Russell, 1970) has been Scapulet, scapulum (pl.: scapulets, scapulae): append-
confusingly referred to as the "main canal" in ages of the mouth-arms of certain rhizostomes, sit-
view of its probable function. uated near the base on the outer side of each.
Radial sinus: in Cubomedusae, the counterpart of Typically, Iike the rest of the mouth-arms, they
radial canals. support numerous mouthlets. Found in the genera
Radial syrnmetry: symmetry in which any diameter Eupilema, Rhopilema, Rhizostoma and Stomolo-
bisects the object into two identical halves. In phus which together comprise the superfamily
many scyphomedusae, despite the arrangement of Scapulatae (sensu Kramp, 1961). Formerly termed
the radial canals the bell can be just so divided epaulettes.
along any diameter: the division does not have to Scyphistoma (n.): the sessile, polypoid stage in the scy-
be along a radial canal. In cubomedusae, which phomedusan life-cycle. See also scyphomedusa.
have four radial canals, the symmetry is said to be Scyphomedusa (n.): a medusa of the subclass
tetraradiate; and though this is conceptually differ- Scyphomedusae which comprises mainly the
ent it is also radial symmetry since any diame ter majority of large medusae such as most of those
will still produce identical halves. Many scypho- seen at the coast. They and the subclass
medusae have eight radial canals, and the term Stauromedusae or stalked jellies, small, stalked,
octoradiate syrnmetry can be applied. In bilateral mobile but sedentary forms, comprise the class
syrnmetry there is only one plane along which the Scyphozoa. There are three orders, Coronatae,
structure can be cut to produce two mirror-image Semaeostom( e)ae and Rhizostom( e)ae. See also
halves. See also perradial. Medusozoa, Cubozoa.
Radius (n.): see perradial, radial syrnmetry. Scyphozoa: see scyphomedusa.
Rhizostom(e)ae: one of the three main divisions of Semaeostom(e)ae: one of the three main divisions of
Scyphomedusae. Scyphomedusae.
Rhopalar: see rhopalium. Septum (n.): an internal partition. Radial septa separ-
Rhopalial (rhopalar) canal: a radial canal ending at or ate, for example, stomach pouches in some genera
near a rhopalium. Inter-rhopalial canals lie between. of coronates and semaeostomes.
See also radial canal; perradius; rhopalium. Skirt (of umbrella): a peripheral zone in which the
Rhopalial'cleft: a gap in the scyphomedusan margin mesoglea is thin, in many species hanging like a
having a rhopalium, situated between two (rhopa- skirt from the thicker, stiffer central region; and in
lial) lappets. See rhopalium. e.g. swirnming functionally distinct from it.
Rhopalial pit: see rhopalium. Stauromedusa (n.): see scyphomedusa.
Rhopalial lappet: see lappet; rhopalial cleft. Stomach pouch: in coronate and some semaeostome
Rhopalial sinus: in cubomedusae, a sinus connecting medusae, diverticulae from the stoma"h extending
the ring canal to the lumen of the rhopalium. to or nearly to the edge of the bell or disc, per-
Rhopalium (n.), rhopalar (adj.), inter-rhopalar, rhopa- forming the internal transport function done by
lar niche: in both cubomedusae and scyphomedu- radial canal s in other medusae. Peripherally, the
sae, a rhopalium or (marginal) sense organ is a stomach pouches may fork into several much nar-
small, roughly cylindrical, organ typically situated rower diverticulae or sub-pouches, and these may
in a cleft in the margin, or in a niche or pit on the branch. The stomach-pouch in some ways recalls
side of the bell. Most support Iight receptors and the open blood-system found in arthropods.
posture- and movement-detecting organs. They Rhopalar stomach-pouch: one in a rhopalar radius;
occur on or near the margin of the bell in scypho- tentacular stomach-pouch: one in a tentacular radi-
medusae, and in pits in the exumbrellar surface in uso Gastrovascular pouch is a less appropriate term
cubomedusae. The radii in which they occur are since the pouches are of the stomach alone. See
termed the rhopalar radii, those between being the also gastrovascular.
inter-rhopalar radii. Rhopalar stomach-pouch: one Strobilation (formerly, also strobilization): the succes-
sive vegetative cutting-off of a sequence of new rather than include a lengthy discussion we prefer to
individuals from one end of a parent individual. In mention none of the higher divisions here.
scyphomedusae, ephyrae are strobilated from a scy- Genera in bo1d characters are those reported from
phistoma. See also ephyra; monodisc scyphistoma. south Atlantic waters, and are therefore treated in
Subumbrellar cavity, subumbrellar surface: see bell. detail here.
Suspensoria: in some cubomedusae, vertical mesent-
eries joining the perradial edges of the manubrium Phylum Cnidaria Hatschek, 1888
to the subumbrellar surface of bell. See also fren- Class Cubozoa Wemer, 1975
ulum; gastric septum. arder Cubomedusae Haeckel, 1877
Tentacle field: a region of the subumbrella in which Family Carybdeidae Gegenbaur, 1856
tentacles are grouped (e.g., Cyanea, Desmonema). Genera Carybdea Péron and Lesueur,
Tentacle, marginal: in cubomedusae and most scy- 1810; Tamoya Müller, 1859; Tripedalia
phomedusae, a tentacle actually or essentially sit- Conant, 1897; Caruláa Southcott, 1967
uated on the edge of the bell. Family Chirodropidae Haeckel, 1892
Tentacul~r radius: a radius in which there is a tentacle, Genera Chirodropus Haeckel, 1880;
as distinct from a rhopalar radius in which there is Chironex Southcott, 1956; Chirop-
a rhopalium. salmus L. Agassiz, 1862
Terminal appendage, terminal club, terminal filament: Class Scyphozoa Goette, 1887
see club; filamento Appendage is the general term, Subclass Scyphomedusae Lankester, 1877
indicating either or both clubs and filaments. arder Coronatae VanhOffen, 1892-
Tetraradiate symmetry: see radial symmetry. Family Atollidae Bigelow, 1913
Umbrella, umbrellar: see bell. Genus Atolla Haeckel, 1880
Velar lappet (obs.): also known as velar lobe. A con- Family Atorellidae VanhOffen, 1902
fusing term indicating all the lappets of a scyphom- Genus Atorella VanhOffen, 1902
edusa excepting the rhopalial lappets: that is, most Family Linuchidae Haeckel, 1879
ofthem. Nothing to do with a velarium or a velum. Genus Linuche Eschscholtz, 1829
Velarium, velarial canal: in cubomedusae, the velari- Family Nausithoidae Bigelow, 1913
um is a mesentery-Iike horizontal ring of tissue Genera Nausithoe K61liker, 1853;
partially constricting the bell aperture (or velarial Palephyra Haeckel, 1880
aperture). It corresponds functionally with the Family Paraphyllinidae Maas, 1903
velum of hydromedusae, but is not homologous Genus Paraphyllina Maas, 1903
with it. From the ring canal short canals enter the Family Periphyllidae Haeckel, 1880
velarium, typically several per quadrant, somewhat Genera Nauphantopsis Fewkes, 1885;
contorted in outline and branching in some species. Pericolpa Haeckel, 1880; Periphylla
Wart: see nematocyst wart. Haeckel, 1880; Periphyllopsis
VanhOffen, 1900
Classification and species Iist arder Semaeostomeae L. Agassiz, 1862
Family Pe1agiidae Gegenbaur, 1856
The following classification is largely based on the Genera Chrysaora Péron and Lesueur,
work of Kramp (1961). The position of the 1810; Pelagia Péron and Lesueur,
Cubomedusae remains problematic. Some authorities 1810; Sanderia Goette, 1886
regard the group as equivalent in rank to the Family Cyaneidae L. Agassiz, 1862
Scyphozoa, calling them Cubozoa, while others Genera Cyanea Péron and Lesueur,
. include them within the Scyphozoa, as the 1810; Desmonema L. Agassiz, 1862;
Cubomedusae. Arguments are balanced, and the ranks Drymonema Haeckel, 1880
given to the taxa ofthe Hydrozoa have also to be con- Family Ulmaridae Haeckel, 1879
sidered. Kramp (1961) and several authors before and Subfamily Aureliinae L. Agassiz, 1862
since have recognized newly proposed groupings Genus Aurelia Péron and Lesueur, 1810
within the Rhizostomeae at various ranks above fam- Subfamily Deepstariinae Larson, 1986
ily level. The validity of some has been debated, and Genus Deepstaria Russell, 1967
Subfamily Poraliinae Larson, 1986 Class Cubozoa (Fig. 3) (after K.ramp, 1961; Wemer,
Genus Poralia VanhOffen, 1902 1984)
Subfamily Sthenoniinae Mayer, 1910 Medusa: bell roughly 4-sided, about as tall as wide
Genus Phacellophora Brandt, 1835 (not dish-shaped); bell margin without lappets; tenta-
Subfamily Ulmarinae K.ramp, 1961 cles peripheral, interradial, 4 or in 4 groups of sever-
Genera Diplulmaris Maas, 1908; al, eaeh tentacle on an simple (undivided) pedalium or
Discomedusa Claus, 1877 several on one palmate pedalium. Sense organs eom-
Subfamily Stygiomedusinae Russell, plex, 4, perradial, in niehes in sides of bell; stomaeh
1959 with 4 wide perradiallobes (saes) extending into sub-
Genus Stygiomedusa Russell, 1959 umbrellar eavity, the lobes ineompletely separated by
Order Rhizostomeae Cuvier, 1799 radial septa; mouth with single opening; gonads leaf-
Suborder Kolpophorae Stiasny, 1921 shaped, in 4 pairs, attaehed along interradial septa;
Family Cassiopeidae L. Agassiz, 1862 opening of bell eavity partly closed by annular dia-
Genus Cassiopea Péron and Lesueur, phragm (velarium). Polypoid stage: small, seeming
1810 solitary but in faet, in speeies deseribed, eomprising a
Family Cepheidae L. Agassiz, 1862 large developing polyp whieh metamorphoses totally
Genera Cephea Péron and Lesueur, into a young medusa (i.e., no strobilation, nor ephyra
1810; Cotylorhiza L. Agassiz, 1862 stage) and a mueh smaller resting polyp (Restkorper
Family Mastigiidae Stiasny, 1921 in German) whieh after medusa release enlarges to
Genera Mastigias L. Agassiz, 1862; form the next developing po1yp; without exoske1eton.
Mastigietta Stiasny, 1921; Phyllorhiza
L. Agassiz, 1862; Versuriga K.ramp, Order Cubomedusae
1961; Thysanostoma L. Agassiz, 1862 With the eharaeters of the class.
Suborder Daktyliophorae Stiasny, 1921
Superfamily Inscapulatae Stiasny, 1921 Family Carybdeidae (Fig. 3.1-3.3)
Family Lychnorhizidae Haeekel, 1880 Cubomedusae with 4 tentacles, or in 4 groups of 3 or
Genera Anomalorhiza Light, 1921; 2; eaeh tentacle inserted separately on its own simple,
LycJmorhiza Haeekel, 1880; Pseudo- unbranehed pedalium. Generie divisions are provi-
rhi=a von Lendenfeld, 1882 sional.
Family Catostylidae Gegenbaur, 1857
Genera Acromitoides Stiasny, 1921; Genus Carybdea
Acromitus Light, 1914; Catostylus L. Stomaeh not suspended from mesenteries; gastrie eirri
Agassiz, 1862; Crambione Maas, 1903; in brush-like bundles at eomers of stomaeh or in eres-
Crambionella Stiasny, 1921 eentie areas extending horizontally;. only 2-3(4)
Family Lobonematidae Stiasny, 1921 branehing velarial canal s in eaeh oetant; aperture to
Genus Lobonema Mayer, 1910 rhopalial niehe roughly T-shaped, 3-lipped.
Superfamily Scapulatae Stiasny, 1921 Carybdea alata Reynaud, 1830 (Fig. 3.1) (mostly
Family Rhizostomatidae Cuvier, 1799 after Mayer, 1910; K.ramp, 1961; Dehida, 1970).
Genera Eupilema Haeekel. 1880; Largest earybdeid when full grown, up to 230 mm
Rhizostoma Cuvier, 1799; Rhopilema bell height reeorded, but also fertile 1/4 as high.
Haeekel, 1880 Bell nearly round in eross-seetion; elongate, height
Family Stomolophidae Haeekel, 1880 variously reported as 1.2 to 1.7 times width, even
Genus Stomolophus L. Agassiz, 1862 twiee, proportionaly higher when large; hyaline (-
glassy), exumbrella lacking nematocyst warts at
Ineertae sedis least when young (some doubt); apex bluntly flat-
Genus Tessera Haeekel, 1880 topped. Pedalia long, sealpe1- or laneet-shaped;
Genus Tetraplatia Buseh, 1851 pedalial canal bent through right-angle at top with-
out divertieulum. Contraeted tentacle thiek, band-
In the deseriptions that follow, useful diagnostie ehar- ed, up to about 1.5 times bell height, tapering for
aeters are highlighted in bold. basal ea. 1/6. Rhopalial niehe deep, entranee T-
shaped, conspicuous, slightly more than 1/4 up milky-yellow; rhopalial photoreceptors brown.
from bell edge; entrance with one bluntly pointed Dangerous, sting painful but not lethal.
flap above and none or two shallow below, their The somewhat similar Carybdea has no exumbrel-
bases together forming an equilateral triangle with lar warts (except perhaps when small) and a T-
the point downwards; possibly shorter and not shaped entrance to rhopalar niche. Carybdea
meeting in young individuals, longer and meeting atlalltica (Barattini and Ureta, 1960) may be con-
together in larger ones, but this perhaps an artifact specific with Tamoya haplonema.
of preservation; possibly some variation in lip-
shape. Velarium with 24 short, branching, non- Genus Tripedalia
anastomosing velar canals. Manubrium short, lips With 12 tentacles, three at each comer, each on a dis-
slightly recurved. Gastric cirri in 4 deeply cres- tinct pedalium.
cent-shaped areas; a stomach pouch in each gap • Tripedalia cystopllOra Conant, 1897 (Fig. 3.3)
between leading into radial sinus. Qne half-gonad (after Conant, 1898; Mayer, 1910; and Wemer,
on each side of each radial sinus. 1984). A cuboid medusa l cm or more across, with
Color: jelly notably clear, endoderm translucent, three tentacles at each comer. Bell slightly lower
milky white; gastric cirri and flexible parts of ten- than square in side view, flattened apically,
tacles pink to yelrow-pink; sensory knobs of rho- exumbrella covered sparsely with small nemato-
palia dull ochre, the ocelli deep brown to nearly cyst warts. Velarium about 1/6 width of base of
black. bell, with 6 unbranched velar canals per quadrant,
Dangerous, sting painful, causing lasting problems triangular in outline. Rhopalia on hort stalks in
but not lethal. wide niches bordered by rim, about 1/4 to 1/5 up
bell. Stomach short, with 4 wide gastrovascular
Genus Tal1loya pouches leading into radial canals; manubrium
Stomach connected to subumbrella by substantial per- reaching nearly to base ofbell; it and mouth cross-
radial mesenteries; gastric cirri in interradial bands shaped in section. Gastric cirri small, tapering,
extending vertically along walls of stomach; aperture "brush-shaped", arising from a short stalk in
to rhopalial niche narrow, the slit inverted crescent- each interradial comer of stomach. Three sep-
shaped or horizontal. arate pedalia on each comer of bell, length of
Tal1loya ltaplollel1la Müller, 1859 (Fig. 3.2) each about half height of bell, laterally narrow,
(mainly after Mayer, 1910, and Kramp, 1961). each with a single tentacle about 2.5 times its
Bell elongate-cuboid, glassy-transparent in tife, length when contracted. Gonads 4, lamellar, each
somewhat rigid, with flat top; reported to 220 mm spanning both sides of each radial sinus, extending
long; length: breadth ratio possibly increasing with vertically from bell margin nearly to apex and
size. Exumbrella, pedalia and velarium typical- when mature each reaching neighbour laterally;
Iy, perhaps always, covered with nematocyst broods planula larvae. Color: Medusa including
warts. Velarium with ca. 10 much-branched but gonads a uniform light yellowish-brown.
not anastomosing canal s in each quadrant. Pedalia
4, length approx. 2/5 of bell height, scalpel- Family Chirodropidae (Fig. 3.4-3.5)
shaped, sharp-edged. Rhopalia 4, complex; Tentacles in 4 clusters, each cluster on one palmate
entrance to niche a horizontal slit. Tentacles pedalium, several tentacles in each cluster (except in
when contracted approx. as long as bell height, youngest individuals); stomach pouches 4, each with
with regularly spaced nematocyst bands; extend- two diverticula. Distinctions between the genera are
ing to ca. 3 m. Stomach occupying up to 1/3 length under revision. What follows is based on current pub-
of bell; gastric cirri in 4 vertical interradial lished diagnoses.
groups (in radii of bell comers). Qne half-gonad
on each side of radial sinus; lamellar with frilled Genus Cltirodrop"s
edges, when mature each extending to meet next. With 4 stomach pouches each having 2 branched or
Coloration: Bell glassy-transparent; exumbrellar feathered sac-like projections into bell cavity; gonads
warts translucent, spaces between translucence 8, their free margins having grape-like swellings.
forming characteristic lace-pattern; tentacles Cltirodrop"s gorilla Haeckel, 1880 (Fig. 3.4)
Class Cubozoa
Order Cubomedusae
single pedalium palmate pedalium

Family Carybdeidae Family Chirodrop.idae
1 1
rhopalium
rhopalium
() J. Corbera
Carybdea alata
3.4a
1 Chirodropus gorilla
rhopalium
Tamoya haplonema
rhopalium
Chiropsalmus quadrumanus
Tripedalia cystophora
(after Mayer, 1910; Kramp, 1955). A large, poorly bell edges; from each stomach pouch hang two
known, cuboid medusa. Bell reported up to ea. 120 smooth finger-shaped sacs nearly reaching bell
rnm high and wide, nearly flat-topped, probably margino Gonads 4, each a thin sheet either side of
without prominent vertical flutings on sides and a radial canal; when mature each extending to
comers; jelly thick, rigid. Lower part of bell thin, meet next laterally. Coloration transparent except-
comprising 16 gelatinous lappet-Iike structures ing tentacles, outer 1-2 typically pale mauve,
connected by velarium like a web. Velarium wide remainder colorless to pale whitish-yellow.
(1/6 bell height), thin, velarial canals 20+ per quad- Dangerously venomous, lethal to children. C.
rant, tortuous, some with major forkings, all with quadrigatus Haec~el, 1880, may be conspecific.
short side-branches. Rhopalia 1/4 way up from bell
margin, slit narrow, horizontal to downcurving, Class Scyphomedusae
lips small. Pedalia 4, inserted along 1/3 of bell
height, palmate, one proximal and several paired Order Coronatae (Fig. 4, 5.1-5.10)
digitate processes each bearing a tentacle; up to 12 After Kramp (1961), Larson (1986), Wemer (1984).
tentacles recorded on each bell comer. Gastric Medusa: umbrella with coronal groove on upper sur-
pouches 4, each bearing 2 conspicuous finger- face; margin cleft into stout lappets with thick meso-
shaped processes hanging down into subumbrellar glea; tentacles solid, situated in the clefts, appearing
space, each with numerous digitate to filiform away from edge ofbell but in fact periphera1; stomach
processes on one side (length differing between wall joined to subumbrella by four gastric septa;
specimens?). Gonads 4, attached beneath top of mouth-opening single, with undivided lips.
subumbrellar cavity, Iike bunches of grapes in Exumbrella radially thickened between coronal
shape. C. palmatus (Haeckel, 1880) may be con- groOYe and marginal 1appets (pedalia). Polypoid
specific. stage: those known having Stephanoscyphus form,
Virulence unknown. strobilating ephyrae which develop into adults.
Atorella subglobosa might be expected in the South
Genus Chiropsalmus Atlantic, but has not yet been reported.
With 4 stomach pouches each with two finger-shaped,
unbranched, diverticula projecting into bell cavity; Fami1y Atollidae (Fig. 4, 5.1-5.4)
gonads 8, laminar, margins entire, undivided. Coronatae with marginallappets 16 or more, the clefts
Chiropsalmus quadrumallus (Müller, 1859) (Fig. between having altemately a rhopalium or a tentacle.
3.5). Fist-sized, cuboid, transparent, with several
tentacles at each comer. Bell robust, nearly as high Genus Atolla
as wide (1: 1.3), slightly rounded but flat on top; Atollidae with the characters of the family, to which
diameter to ea. 140 rnm; surface smooth. Velarium may be added: gonads 8, adradial; rhopalia 8; disk
with numerous branched canals. Rhopalia 4, on quite flat, mesoglea of central region lens-shaped;
sides of bell, in triangular niches ea. 1/6 up from coronal groove deep. The four accepted species are
bell margino Tentacles 24 to ea. 40, each extending notably uniform in appearance. Just the cornmonest
up to ea. 3-4 m, banded when contracted, each species is therefore described in detail, and the dis-
bom on short finger-shaped process inserted on criminating characters of the other three species sum-
one of 4 palmate pedalia; pedalia robust and marised afterwards.
inserted on lower 1/3 of bell, bilateral along mid- • Atolla wyvillei Haeckel, 1880 (Fig. 5.1) (mainly
line. Manubrium 1/2-113 bell height, with 4 trian- after Russell, 1970). The following description
gular lips. Stomach wide, round, prolonged into 4 applies to adult or near-adult individuals. Disc flat,
pouches which extend into radial sinuses along notably circular; diameter up to ea. 150 rnm; meso-
•••
Fig. 3. Cubomedusae: schematic external views and details of diagnostic characters of the adults of the species recorded in the South Atlantic.
Scale bars equal 10 cm unless otherwise indicated.
Sources: from Cornelius (1997c), redrawn: 3.1b, 3.2b. 3.5b; from Mayer (1910), redrawn: 3.2a, 3.5a; from Pagés el al. (1992), some redrawn:
3.la, 3.4a, b (copyright © 1. Corbera); from Werner (1984): 3.3a, b.
-
an
cles
aeach
th 16 J:
in
nts wide
N from
534 '" Fig.
oQJOe.c
Linuchidae
marginal
V'l
QJ
l'l:l
O
stomach
bell
terminal
Rhizostomatidae
Deepstariinae
Stygiomedusinae
Cepheidae,
groove
arising
Stomolophidae
Tentacles
Canal network
More than 16
j
~]J.J::C:I~
Without
l'l:lQJ Fig. 5.32
QJ
5.28-5.29
r(
pouches I'¡
,
,----,radial
;( ••••
clubs
near ~~(~~t'
\~~/IJJfl~
stomach
Fig.
~Fig. ~
I
mouth
V{r,1i~~
Small;
5.30-5.31
5.24-5.25
~(with
12
V
t.'ir;r
Mastigiidae
With
ll
NetworkWith
Tentacles
Fig.5.10
Sthenoniinae
Ulmarinae
With
Wlthout
tentacles
arising
Fig.5.9
~.
I
Periphyllidae
'h,,~T:>
canals)
from
arms
terminal~(\1
canal
subumbrellar
adult
groove
tentacles
middle
Cyaneidae
surface
from
1"coronal
Large;
Catostylidae
Lychnorhizidae,
NauslthOldae~
Poraliinae
Aureliinae
fused
8tentacles
'
marginal
near
Fig.5.16-5.18
O> Fig.
mouth
-~
basall~
11
c of
ring
coronal ubs I
5.26-5.27
arms
I}JI~~
In
(".and
L
.adu!t
'
ICubomedusaein adult
' and
Paraphyllinidae
Fig.5.11-5.15
Fig.
Fig.5.8
Fig.5.5-5.7
'ir
5.19-5.23
(/111
'°1
Scyphomedusae
, ¡ Fig.5.1-5.4 1\
~
glea of central region lens-shaped, its edge with 2l • Atolla parva Russell, 1958 (Fig. 5.3). Small, diam-
(not 22) notches, coronal groove slightly closer to eter typically ea. 10-30 rnm. Marginal tentacles
periphery than to centre; jelly thick, unbending; 20-26; outline of stomach recalling Ace of Clubs
tentacles short. Marginal lappets typically 44, but with tbe four lobes equal; the single "hyper-
shortly elongate, rounded. Rhopalia 22, alternating trophied" tentacle in exactly same radius as sep-
with 22 marginal tentacles which are short, stout tum between lobes of stomach; no papillae on
and sharply tapered; one "hypertrophied" tentacle marginal lappets; peripheral radial septa straight,
(noticeably larger than the rest), not in exactly parallel, not diverging centripetally, terminating
same radius as septum between lobes of stom- at level of inner edge of coronal muscle.
ach. Each tentacle inserted onto prominent basal • Atolla val/hoejjeni Russell, 1957 (Fig. 5.4).
pedaliurn. Other nurnbers of marginal tentacles Diameter typically up to ea. 30 rnm, occasionally
occur, but recorded somewhat rarely. Coronal larger; marginal tentacles typically 20; base of
musculature robust, an annulus typically 5-10 rnm stomach in shape of square-sided cross with
wide, whitish, visible when pigmented epidermis right-angled corners and straight sides, and with
sloughed off; outer half of annulus thicker than total of 8 dark pigment spots next to outer cor-
inner. Manubrium wide, projecting below disc, lips ners; the single "hypertropbied" tentacle in
somewhat pointed, each thickened by bulging, lon- exactly same radius as septum between lobes of
gitudinal, mesogleal ridge. Stomach outline of 4 stomach (i.e., opposite an inner comer of stom-
rounded lobes; 4 vertical gastric septa, triangular ach-cross); no papillae on marginal lappets;
in horizontal section; 4 perradial ostia from stom- peripheral radial septa not or hardly diverging at
aeh leading into annular gastrovascular sinus. inner ends, and not or hardly extending beyond
Gastrovascular sinus typically divided by 22 pairs inner margin of coronal muscle.
of closely adjacent radial septa, between which
narrow channels run each to base of one rhopali- Family Linuchidae (Fig. 4, 5.10)
um; septa diverging at inner ends and extending Rhopalia 8, marginal tentacles 8, minute; marginal
beyond inner margin of coronal muscle. Gonads lappets minute, 16; subumbrella having 2-3 rows of
8, elongate-oval with rounded ends, forming brok- large, conspicuous warts; stomaeh pouehes 16, radiat-
en annulus just internal to tentacle bases. ing, breaking up into numerous ragged-edged branch-
Coloration: reddish-brown; stomach deeply pig- es in the marginal lappets; coronal groove just below
mented, epidermis over central mesogleal "Iens" top ofbel!.
pale to dark; coronal muscle when exposed white
to greenish-yellow; tentacles uncolored. Thin, col- Genus Lilluche
ored epidermis often lost, revealing transparent Linuchidae with the characters ofthe family, to which
mesoglea. may be added: bell apex flat, sides vertical; marginal
See Russell (1970) for diserimination of the four lappets bluntIy oval; gonads 8 (i.e. 4 double ones).
speeies. A. gigantea Maas, 1897, has been held dis- Linuche ullguiculata (Schwartz, 1788) (Fig. 5.10).
tinct by some recent authors (e.g. Larson, 1986). One of the smallest scyphomedusae. Appearing
• Atolla c1llllli VanhOffen, 1902 (Fig. 5.2). mostly brown, thimbled-shaped, 10-20 rnm; a
Diameter up to ea. 70 mm' marginal tentacles 24; few short, almost invisible, tentacles around mar-
shape of stomaeh 4 rounded lobes; the single gin; active, bell pulsating rapidly. Bell tiny, stiff-
"hypertrophied" tentacle in exactly same radius sided; top nearly flat, with coronal groove imme-
as septum between lobes of stomach; with 7-9 diately below; sides straight, with 16 vertical
small, papillae on upper surface of each margi- flutings (elongate pedalia); no exumbrellar warts.
nal lappet, typically one in centre and remainder Marginallappets very short, 16; tentacles rninute,
in two lateral rows; peripheral radial septa diver- 8, adradial, only ea. 1.5 times as long as lappets.
gent centripetally. Manubrium extending to 2/3 ofbell height, 4-sided,
•••
Fig. 4. General classification and main diagnostic characters of South Atlantic Scyphomedusae.
mouth with 4 large recurved lips. From tloor of known only from laboratory-reared Brazilian mate-
stomach pouches 48 dark warts project down rial. Central disc recorded to 10.5 rnm, exumbrellar
slightly into subumbrella; arranged in outer ring of surface flat, smooth; coronal groove ca. 2/5 from
32 and two concentric inner rings of 8. Gonads centre, gently crenate. Zooxanthellae in gastroder-
elongate, slightly crescentic. Coloration: colorful in mis. Lappets 16, bluntly pointed to truncate; alter-
brownish patches. Gonads, subumbrellar warts, and nate clefts between lappets having rhopalium or fine
irregularly polygonal areas in gasrtic pouches tentacle (to 5.5 mm in life); all clefts similar in
brown; 8 rows of dark brown spots along sides of depth. Peripheral region without radial grooves.
manubrium; rest colorless, transparent. Rhopalia 8, conspicuous, on short hemispherical
L. aquila is said to replace L. llllguiculata in the projections. Subumbrellar musculature unde-
Pacific, and to differ by having a ring canal and scribed. Mouth cross-shaped. Stomach 4-lobed;
two rows of subumbrellar warts, not three, the 16 gastric filaments typically 3 in each quadrant,
inner ones being in a single row, not two rows of one larger tlanked by two smaller laterals. Gonads
eight. If the geographic and taxonomic distinctions spherical, spaces between about equalling their
are valid, still either form might conceivably occur diameter. Coloration diagnostic: transparent
in the south of either ocean. There are reports of except for a yellow spot centrally on each lappet,
intermediate material (Mayer, 1910; Comelius, yellow to pale brown radial patches peripheral to
pers. obs.). each gonad, and circularly-orientated patches over-
Iying spaces between gonads; gonads brown when
Family Nausithoidae (Fig. 4, 5.5-5.7) mature.
Rhopalia 8, tentacles 8, marginal lappets 16; without Nausithoe p'lllctata K6l1iker, 1853 (Fig. 5.7) (after
sac-like pouches on subumbrella; stomach pouches tvtayer, 1910; Larson, 1986.) Tiny, discoid, thick,
16, radiating, not divided. with finely warted surface and crenate coronal
groove. Disc tlat, diameter recorded to ca. 15 mm;
Genus Nausithoe central region 2.5 wider than high; it and lappets
Gonads adradial, 8. Several nominal species await with numerous tine nematocyst warts; coronal
validation. groove prominent, crenate, delimiting central
NausitllOe rubra VanhOffen, 1902 (Fig. 5.5, based half of disco Marginallappets 16, with altemating 8
on N atlantica illustrations in Russell, 1970) thick tentacles and 8 prominent rhopalia in clefts
(after Russell, 1970; Larson, 1986.) Small, lower between; radial grooves running to lappets, separ-
than hemispherical, with smooth surface and ating thick pedalia. Tentacle clefts and rhopalial
evenly Circular coronal groove. Disc diameter clefts about equal in depth. Circular muscle
recorded to 35 mm, height half the width, coro- narrow, less than 1/3 lappet length in radial
nal groove prominent, circular (not crenate), extent. Gonads 8, spaced apart, oval to rigidly
delimiting central half of disco Marginal lappets elongate in outline, (?always) centripetal to cir-
16, with altemating 8 tentacles and 8 rhopalia in cular muscle. Coloration varied: jelly translu-
clefts between, shallow radial grooves in radii of cent-milky, greenish or light brownish; gcinads
lappets, delimiting gradually raised thickened (male) bright yellow or (female) brown to red.
radial pedalia. Tentacle clefts deeper than rho- Pigment patches between gonads reported by Thiel
paliar ones. Circular muscle broader than in N. (1966). Small crystalline spots on exumbrella.
pUl/ctata, more than half lappet-Iength in radi-
al extent. Gonads 8, contiguous in pairs, U- Family Paraphyllinidae (Fig. 4, 5.8)
shaped initially, one arm of U growing out- Rhopalia 4, perradial; marginal tentacles 12.
wards to underly circular muscle. Coloration:
endodermis chocolate-red throughout; mesoglea Genus Paraphyllil/a
lightly yellowish-brown throughout; stomach Paraphyllinidae with the characters of the family, to
walls deeply colored, as is mesoglea in lappets. N which may be added: tentacles 12, marginal lappets
atlantica Broch, 1913, may be conspecific. 16; gonads interradial, in 4 pairs.
Nausithoe aurea Silveira and Morandini, 1997 Paraphyllilla rallsol/i Russell, 1956 (Fig. 5.8)
(whence details taken) (Fig. 5.6). Medusa minute, (after Russell, 1970). Bell transparent, dome-sha-
ped, with prominent coronal groove; up to 7.5 cm single, lips 4, typically with 4 large mouth-arms SUf-
in diameter, exumbre11a with nematocyst warts, rounding. Polypoid stage: a polydisk scyphistoma.
mesoglea fairly thick. Pedalia 16. Marginallappets
16, rounded. Rhopalia 4, perradial. Tentacles 12, 3 Family Pelagiidae (Fig. 4, 5.11-5.15)
between each rhopalium and next, 2/3 length of Stomach giving rjse to radiating pouches which are
bell. Subumbre11ar coronal muscle continuous, separate and unbranched; lacking a ring canal; tenta-
prominent. Stomach attached to subumbre11a by 4 cles arising from umbre11a margin in clefts between
interadial gastric septa, forming 4 pockets that sep- lappets (except secondarily some in Chrysaora quin-
arate subumbre11a from manubrium. Phacelli 8, quecirrha from underside of lappets); mouth arms
each with ca. 20 gastric filaments. Perradial ostia 4, long, pointed, much folded.
connecting stomach with gastrovascular sinus; dis- The sting of most species is painful, and medical
tally divided by 16 centrifugal septa. Manubrium attention may be needed.
with 4 mesogleal perradial reinforcements and cre-
nate margino Gonads 8, W-shaped. Color of stom- Genus Chrysaora
ach deep chocolate-red, rest colorless. Marginallappets 32-48 (sometimes secondarily more
in some specimens); rhopalia 8; tentacles per octant 3-
Family Periphyllidae (Fig. 4, 5.9) 9; stomach pouches 16; in marginal zone, typically
Rhopalia 4, interradial; marginal tentacles 4 or more having the 8 rhopalar stomach pouches much narrow-
(up to 28). er than the 8 inter-rhopalar pouches. Notes on colora-
Genus Periphy/la tion and pattem are taken from a variery. of literature
Tentacles 12 (4 perradial, 8 adradial); marginal lap- sources, and may describe the range of the species
pets 16; gonads 8. rather than that of any one population. Some, perhaps
Periphy/la periphy/la (Péron and Lesueur, 1810) a11, species cornmonly reach sexual maturity before
(Fig. 5.9) (after Mayer, 1910; Russell, 1970; all the tentacles have developed, confusing identifica-
Mianzan, 1989). Be11 steeply conical to domed, tion based solely on tentacle number and distribution.
apex pointed; coronal groove prominent; height In addition, it seems that al! species may have no
to 35 cm, diameter to 25 cm; height:diameter ratio exumbrella color pattern, or may be variously colored
variable. Mesoglea thick, rigid. Exumbrella with a range of density, so color can be misleading in
smooth. Marginallappets 16 (8 rhopalar + 8 ten- identification.
tacular), uniform, elongate-rounded, thick basally,
thinning to margino Rhopalia 4, in deep clefts Several authors have described Chrysaora spp. from
between lappets. Tentacles 12, three between west Africa, including C. fulgida Reynaud, 1830 and
each rhopalium and next, thick, tapering; in life C. africana Vanh6ffen, 1902, but for a variety of rea-
held pointing either aborally or ora11y but all in sons we are as yet unable to relate them to the taxa
same direction, more or less straight. Coronal described here. C. blossevillei Lesson, 1832, based on
muscle broad, longitudinally folded. Stomach Brazilian material, is also problematic. It is also impor-
large, occupying whole of upper part of central tant to note that there seem to be no valid records of C.
dome; communicating with peripheral gastrovas- hysoscella and C. quinquecirrha from South America
cular sinus through 4 perradial openings, numer- south of the equator. C. quinquecirrha records from
ous large gastric cirri projecting through each one. west Africa await validation.
Gonads 8, disposed in 4 U-shaped groups looping • Chrysaora hysoscella (Linné, 1766) (Fig. 5.12).
across lower 2/3 of be11, straddling interradial Diameter up to ca. 30 cm; exumbre11a smooth; ten-
septa. Color: stomach and manubrium deep red- tacles 24 in adult, 3 of uniform size per octant, all
dish-brown to brown; exumbrella and sub-surface arising from edge of disc in cleft between lappets.
tissues colorless. Clefts between tentacular lappets uniform in depth;
length of oral arms up to ca. 5 times bell diameter,
Order Semaeostomeae (Fig. 4, 5.11-5.25) narrow, V-shaped in section, edges friUed, extreme
Medusa: umbre11a typica11y flatter than a hemisphere, terminal region (may be lost) spirally coiled. Radial
with margin cleft into lappets; tentacles (if present) septa (as seen in optical horizontal section) typical-
hollow, peripheral or on subumbrella; mouth opening Iy narrow, becoming wider centripetaUy only
gradually, inner end roughly truncate. Coloration spiral, lacking such structures. Radial septa (as
greatly varied, but lappets typically brown; rest of seen from below) slightly thicker than in other
bell colorless, or variously colored, or with varied Chrysaora spp., constant in width except at inner
pattem of radiating lines or patches, some lines end where they widen abruptly with pear-
meeting near-centrally to form isosceles triangles, shaped outline. Coloration greatly varied. Different
some specimens having the "triangles" filled in. color varieties each seldom recorded, their relative
Ground color varied from colorless through white abundance in space and time being unknown.
to yellow, orange and brown, radial lines varied Known varieties are: (1) nearly (?actually) colorless
from red through orange to chocolate. Mouth-arms and translucent, with faint orange radial stripes; (2)
white, yellowish to orange, perhaps also brown. same, with triangles infilled with similar pigment;
Sting harmless to severe, possibly environmental- (3) triangles more densely infilled with finely retic-
ly dependent (Comelius, pers. obs., U.K.). ulate dark red-brown pigment, with pale bands
Chrysaora lactea Eschscholtz, 1829 (Fig. 5.15). between each "triangle" and next; (4) with ground
Diameter recorded up to ca. 25 cm. Exumbrellar color uniforrnly distributed across bell, both within
surface thickIy covered by minute truncate triangles and over bands between. Ground color
papillae. Tentacles up to 40 in aduIt, each may also be orange, or white; bell may have dark
octant having 5 tentacles, a central primary purple edge; tentacles may be dark purple; mouth-
tentacle (forming earliest and being largest until arms may be white/translucent-colorless. Marginal
adult size is reached), two secondaries slightly tentacles white (recent photos) to red with yellowish
smaller or same size (?depending on age) later- bases (type description). Sting recorded as irritating
ally, and two, smaller, tertiaries in the spaces but not severe, lasting 30-60 mins.
between the primary and secondaries, all aris- Chrysaora qllillqllecirrha (Desor, 1848) (Fig.
ing from clefts between lappets. Clefts between 5.14). Diameter to ca. 25 cm; exumbrella smooth;
tentacular lappets not uniform in depth, those of tentacles typically 40 in adult (56 and 72 rarely
tertiary marginal tentacles being shallower than reported), each octant having a central primary
those of secondaries, and those of primaries still tentacle, two secondaries of intermediate size
shallower (see Fig. 5.l5b); oral arms V-shaped in either side, and a minute tertiary near each side
section, edges slightly frilled, terminal region of octant arising beneath rhopalar lappet (i.e. not
possibly spiral (data scant). Stomach pouches uni- from a cleft between the lappets); clefts between
form in width. Radial septa narrow, some widen- tentacular lappets uniform in depth; oral arms nar-
ing at inner end either gradually (as in C. hysoscel- row, V-shaped in section, edges frilled, terminal
la), or ábruptly with pear-shaped outline (as in C. region not spirally coiled (?); very long, length up
plocamia but not so widely). Coloration little to ca. 15 times bell diameter. Radial septa (as
known, so far recorded as uniform, ranging from seen from below) thin, constant in width, not
pale milky-white, or with weak purplish tint, or wider at inner end; outer 1/3 to 1/4 following broad
pale lilac. Brownish-red radial lines, spots and curve (rather than merely bending as in other spe-
patches on exumbrella recorded in some popula- cies). Coloration greatly varied. Some specimens,
tions (M ayer, 1910; Cornelius, pers. obs.). and/or populations, temporarily or permanently
Marginal tentacles colorless, once pale reddish- entirely lack radial lines on exumbrella, but others
purple (Eschscholtz, 1829). Mouth arms and sub- have them. Some are colorless, transparent, whitish
umbrellar ostia spotted dark brown. Sting mild, yellowish or orange, with mouth-arms of the same
lasting 30-60 minutes (Comelius, pers. obs) .. colors but not in all specimens the same color as
Chrysaora plocamia (Lesson, 1832) (Fig. 5.13). the bell; lappets same color as ground color of
Sometimes huge, diameter reported to max. ca. 1 bell. Others have variously developed radial
m, oral arms to ca. 5 m or more, but in many pop- streaks, from small radial dashes near centre, to
ulations reported smaller, more typically to ca. 50 nearly entire radial stripes (? not meeting near cen-
cm diameter. Exumbrella smooth. Tentacles in tre to fonn isosceles triangles); pattem possibly
aduIt 24, 3 per octant; clefts between tentacular never so strongly developed as in some C. hysos-
lappets roughly uniform; oral arms much frilled cella and C. plocamia. Marginal tentacles color-
distally; terminal region of mouth-arm sinuous to less, yellow, red, or of intermediate color. Sting
painful to severe, medical attention may be needed.
Genus Pelagia pa1ial lappet; in some a1so a few tentacles arising

Marginal lappets 16; rhopalia 8; tentacles 8, altemat- from underside of centre of lappet, the number
ing with rhopalia; stomach pouches 16, all alike, each increasing with age; typically ea. 30-250 tenta-
terminating in pair of unbranched canals or sub- des per octant, perhaps more in large specimens.
pouches which enter marginal lappets. Circular subumbrellar musculature near-circular
Pelagia Iloctiluca (Forskal, 1775) (Fig. 5.11). in alignment, in 16 altemating wide and narrow
Diameter to ca. 12 cm, bell square-hemispheri- muscle-fields corresponding to tentacular lappets
cal with numerous conspicuous, in some speci- and pairs of rhopaliallappets; direction of muscle-
mens or popu1ations elongate, stinging warts; folds bowed inwards in centre of each field (no
marginal tentades 16; mouth arms to about 5 radial muscle-fields); mouth-arms comprising four
times bell height, simply frilled, a1so with numer- much-folded curtains approximately same width
ous warts. Co10ration somewhat varied: ground as bell, hanging down in life. Gastrovascular
color of bell transparent to milky, warts system comprising 16 broad pouches; terminating
red/magenta to brown and giving color to bell; in narrow branching canal s in lappets, with some
mouth-arms and marginal tentacles colorless or anastomoses; gonads hanging conspicuously
red/magenta to brown. below umbrella. Coloration much varied, but
Sting painful, severe. analysis fraught due to some previous confusion
between the two species recognised here.
Family Cyaneidae (Fig. 4, 5.16-5.18) Probably not all variants yet known: apparent
Stomach with radiating pouches divided peripherally color may partly follow recent diet. (]¡-ound color
into numerous branching canals ending blindly in of bell variously recorded as pale milk-white to
marginallappets; lacking a ring canal; gonads in com- brownish, but many individuals (varied within a
plex, folded, downward eversions of subumbrellar local population) are variously strongly marked on
wall; tentacles arising from subumbrella, at some dis- exumbrella with 16 broad dark radial lines,
tance from margino extending from near periphery to near centre of
bell. Tentacles pale milk-white to brownish.
Genus Deslllonellla Mouth-arms in individuals with pale !>ells also
Tentacles either numerous, in 8 clusters each of sev- pale, milk-white to pale yellow; in those with
eral rows, or each cluster of a single line of ca. 5-10 darker bells, also darker, brownish. Gonads
tentacles; no radial muscle strands in subumbrella. brown.
Goy (1979) repeated a possible record of Cyanea capil- Deslllollellla gaudichaudi (Lesson, 1832) (Fig.
lata (Linné, 1758) from Cape Hom and the Bay of 5.16). Disc up to 1.5 m. Similar to D. chierchia-
Concepción originating from manuscript notes by C.H. num, but tentacles per octant up to only ca. 20,
Mertens and discussed by Brandt (1838). Brandt typically ca. 2 to ca. 10, broad, flattish-oval in
recorded that Mertens saw certain medusae in the cross-section, up to ea. 10 times as wide as
plankton but it seem~ likely that he saw Desmonema sp. deep; inserted in a single line (not in a fie1d
since there have been no further reports, and C. capil- extending radially). Co10ration: differences from
lata was the most similar species known at the time. D. chierchianutll, if any, not well documented.
• Deslllollellla c/zierc/zialluIII Vanh6ffen, 1888 Ground color white, white with dark spots, dark
(Fig. 5.17) (partly after Mianzan, 1986b). Disc up centre with light margin and dark spots, or uni-
to ca. 1.5 m in diameter, flat, thickest in central formly dark brownish or purp1e, or maroon-violet
region; margin with 8 very wide, deeply cleft ten- (part1y after Larson, 1986; Mianzan, 1986b;
tacular lappets, and 16 narrow, semi-circular rho- Montsalve, 1996).
paliallappets flanking the rhopalia in pairs; tenta-
des fine, numerous, roughly circular in Genus DrYlllonellla
cross-section, almost entirely in 8 groups inserted' Tentacles not grouped in isolated clusters, but arising
in straight, narrow fields typically 2-5 tentades from a wide annu1ar zone of the subumbrella at some
in radial extent, the groups extending whole distance from the edge.
width of each tentacular lappet, but in many spec- DrYlllonellla dallllatillulII Haeckel, 1880 (Fig.
imens a1so extending onto adjacent regio n of rho- 5.18). Umbrella to ca. 1 m in diameter; bell rather
flat, peripheral skirt regio n of exumbrella hav- Genus AlIrelia

ing numerous (up to 144?) furrows, some fork- Aureliinae with the characters of the sub-family, to
ing; lappets numerous, probably increasing in which may be added; umbrella margin with 8-16
number with age, up to ca. 176?, (16 rhopalar plus broad, shallow marginal lobes.
20 groups of 8 tentacular), elongate and fused lat- AlIrelia allrita (Linné, 1758) (Fig. 5.20). Disc flat,
erally along entire length to form skirt ofbell, end typically up to ca. 30 cm in diameter, rarely
shallow-circular and crenate. Rhopalial niches 8, recorded up to 40, 50 or even 100+ cm; flexible,
conspicuous, beneath skirt, on underside, at level transparent; marginal 10bes 8, broad, shallowly
of inner end of lappet; border a raised annulus, almost imperceptibly - cleft between; tentacles
flanked either by two sausage-shaped, curving filiform, about as long as disc diameter, very
lips, with small gap on outer and inner sides, or by numerous, up to a few hundred to more than 1000
continuous raised margin without gap (both seen in large specimens, arising nearly at bell edge.
by present authors). Tentacles arising over wide Subumbrella musculature inconspicuous. Mouth-
zone of subumbrella, from outer edge of stomach arms elongate, tapered, V-shaped in section, in life
to inner edge of lappets, seemingly irregularly typically held horizontally when young (when
scattered; outermost narrow, those of inner ca. 2/3 shorter than bell radius), dangling down in older
of tentacle zone wide, up to 10 mm wide, nearly specimens (when longer than radius). From stom-
flat in section, longer than outer tentacles, exten- ach lead 16 unbranched radial canals and 16 which
sile to nearly 2 m. Oral arms (not preserved in all typically fork roughly di chotomou sIy into branch-
specimens) thin sheets, much folded when cones which connect approximately straight to ring
tracted, extending downward greatly during feed- canal. Gonads in 4 (nearly) complete rings, con-
~ ing to a length slightly greater than bell diameter. spicuous due to their color and transparency of
Gastrovascular pouches as numerous as marginal bello Coloration: transparent, some specimens col-
lappets, narrow, elongate, forking in basal region. orless throughout, in some gonads colored red
Coloration (after Haeckel, 1880; Stiasny, 1940; through magenta to blue, in others extreme periph-
Mianzan, 1986b, 1989; Comelius, Silveira and ery of bell same color as gonads; in still others,
Mianzan, in prep.): ground-color of bell probably entire bell washed with same hue. Radial canal s
somewhat pale in all specimens, variously report- and tentacles whitish.
ed as reddish-white, milk-white, light rose to vio-
let, yellowish brown to pale golden yellow. Many, Subfamily Deepstariinae
but not all, specimens have a star pattem compris- Umbrella thin, wide, shallow; gastrovascular canals
ing 16 darker radiallines (of same general color as anastomosing; marginal tentacles absent.
bell but darker, or brown) extending from or near
centre towards periphery, the distance differing Genus Deepstaria
between specimens. Tentacles translucent, color- With the characters of the subfamily.
less, but also noted as white with inner canal red- Deepstaria eIligmatica Russell, 1967 (Fig. 5.25).
dish, or pale golden-yellow. Imperfectly known, many specimens damaged.
D. gorgo Müller, 1883 is conspecific. Two nominal species, information from both com-
bined here based on Russell (1967), Larson
Family Ulmaridae (Fig. 4, 5.19-5.25) (1986), Larson el al. (1988). Bell remarkably
Unbranched and/or branched radial canals and a ring thin, broad, delicate, a rather flat inverted bowl
canal; with or without subgenital pits. when relaxed; up to ca. 60 cm diameter; mesoglea
1-2 cm thick, tapering towards edge; flexing down
Subfamily Aureliinae in "pursing" manner, margin closing to enclose
Tentacles numerous, in some specimens more than large subumbrellar volume. Rhopalia reported 8 to
1000, filiform, sub-marginal; oral arms fundamently ca. 20, number perhaps increasing during growth;
V-shaped in section; gonads invaginated, with exter- rhopalar lappets minute to prominent, rectangular
nal subgenital pits; radial canal s forked to anasto- with circular ends. Marginal lappets not recorded,
mosed (after Kramp, 1961; Larson, 1986). apparently very shallow or entirely lacking.
Marginal tentacles lacking. Subumbrellar mus- longer than tentacular; probably increasing in
culature diffuse, probably extending over entire number to a maximum recorded 64. Tentacles
subumbrella. Coronal muscle well developed (but maximum recorded 48, arising at non-uniform
thin) comprising large number of circular, striated intervals around bell so that intervening gaps are
muscle fibres near bell margin; distal portion of non-uniform till last is added; laterally com-
muscle bending over toward subumbrella. pressed with row of wart-Iike structures along
Numerous fibres extending from coronal muscle to one side; stout basally but probab1y extensile to
subumbrella. Stomach small, about 1/6 oftotal bell ca. twice disc diameter, in life held remarkably
diameter, circular; protruding beneath subumbrella straight and at varied, irregular angles. Mouth
as short, thick-walled manubrial tube; oral arms 4- arms 4, sides much prolonged and curtain-Iike,
5, narrow, elongate, basal 1/3 heId approximately somewhat folded, about as long as disc diameter.
horizontally, with near right-ang1ed bend and Gastrovascular system comprising circular stom-
remainder hanging down, terminating in curious ach in centre 1/3 of disc; radial canals about
hook-shaped organ. Base of each has on each side a same width as gap between each and next, up to
"plaque" in which are numerous pits containing the 96 reported, comprising 16 rhopalial canals and in
gastric cirri, 3-10 rnm long, up to 8 per pito each intervening paramere 1-5 inter-rhopalial
Gastrovascular canals fin e, somewhat irregular- canals; all essentially unbranched for inner 1/2-2/3
edged, forming reticulate network across most of length, somewhat regularly anastomosed in
of bell, anastomoses more numerous closer to outer 1/3; some forking of rhopalial radial canals
periphery; the spaces in the meshwork elongate, peripherally; both rhopalia and tentacles receiving
aligned radially. Radial canals 8 to ca. 20, both a radial canal. When mature, the 4 gonads forming
rhopalar and (7 when >8) inter-rhopalar reported as single wide ring around stomach, eventually
discernible in some, but not in all, specimens, protruding through; no subgenital cavity or ostia.
approximately twice width of those in main canal Coloration: possibly partly reflects feeding histo-
meshwork, running somewhat tortuously from ry. Recorded as transparent excepting canal
stomach to a peripheral ring canal; ring canal noted system yellowish-brown, tentacles dull purple
as cornmunicating with meshwork canals at broad (Mayer, 1910); gonads and gastrodermis of stom-
intervals. Gonads reportedly 4-5, seen in only a few ach and oral arms orange (from photographs in
specimens, short-stalked and lobed, attached near Rybakov, 1976); gonads bright orange, rest color-
base of oral arms. Coloration: most specimens less (BBC, 1993).
reported colorless but deep brown exumbrella and
stomach lining a paler brown recorded once Genus Discomedllsa
(Larson et al., 1988, as D. reticulwn); more obser- Rhopalia 8; tentacles 24; marginal lappets 32; the
vations needed before value ofthis as species char- eight perradial and interradial canals branched, the
acter can be evaluated. eight adradial canals unbranched; with ring canal.
• Discomedusa lobata Claus, 1877 (Fig. 5.23).
Subfamily Ulmarinae Composite description based on Haeckel, 1880 (as
Tentacles marginal, arising from clefts between lap- Undosa undulata, from Fernando Po, Guinea,
pets; gonads sac-like, protrusive; subgenital pits West Africa, best description), and partly on
absent (after Kramp, 1961; Larson, 1986). Mayer (1910), Thiel (1966), and Pages et al.
(1992). Poorly known, only Haeckel providing a
Genus Dipllllmaris reliab1e account of undamaged adult material.
Rhopalia 16; tentacles 16; marginal lappets 64; rhop- Umbrella quite flat, jelly moderately thick,
alar canals 16, branched; tentacular canals 16, clear; 120 rnm by 40 mm recorded from life
unbranched. (Haeckel, 1880). Rhopalia 8. In a 32-tentacled
Dipllllmaris alltarctica Maas, 1908 (Fig. 5.22). specimen, rhopaliallappets 16, each c1eft in two,
Disc flat, flexing during swirnming; mesoglea each half-Iappet semi-circular to pointed; tentacu-
thin, transparent; maximum diameter reported 180 lar lappets each as broad as a full rhopalial lappet
mm; marginal lappets elongate, rounded to shal- but not cleft, semi-circular (and so its semi-circle
lowly pointed, rhopalial lappets slightly wider and twice the width ofthat ofthe half-rhopaliallappet)
to pointed; in a specimen having three tentacles PhacellopllOra camtschatica Brandt, 1838 (Fig.
between one rhopalium and next, a primary tenta- 5.19). Disc flat (when resting), typically to 60 cm,
cle in the centre of the paramere was flanked by some to ca. 1.5 m diameter; jelly moderately thin,
one secondary on each sid~. Thus between one flexible. Marginallappets 16, broad, curved, outer
rhopalium and next are, in sequence, a double rho- margin shallowly divided into sub-lappets (each
paliallappet, secondary tentacle, tentacular lappet, supplied by a radial canal) in some individuals.
primary tentacle, tentacular lappet, secondary ten- Rhopalia 16, in deep clefts. Rhopalial lappets
tacle, and a double rhopaliallappet, then next rho- typieally in pairs forming Y, arms of Y distal
palium. Tentacles 48. Haeckel recorded (in a 120 and free, rhopalium situated in angle between.
rnm diameter individual, evidently from life) the Marginal tentacles in 16 linear to outwardly-con-
16 primary tentacles 2-3 times bell radius, 16 sec- cave groups, 5-25 per group, eaeh tentacle ellip-
ondaries half "the size of' ("so gross als") the 16 tical in section, with longitudinal canal and 2
tertiaries, unclear if 3 times narrower or shorter rows of nematocyst warts on one side.
(A. Svoboda, pers. cornm.). In specimens having Subumbrellar musculature weak, not prominent.
48 tentacles, possibly the 16 tertiary tentacles Mouth-arm in length about 1/3-1/2 disc diameter,
develop from tissue at base of clefts in rhopalial sides in form of convoluted curtains which may be
"double" lappets. Subumbrellar musculature (?) lost. Stomach occupying slightly more than 1/3 of
not described. Mouth arms thick, broad, "resemble disc diameter, circular. Radial canals broad, typ-
those of Aurelia aurita, but margins more curtain- ically wider than gaps between, margins irreg-
like, more folded, with numerous tentacular fila- ularly sinuous in plan view; 16 rhopalial radial
ments; gutters of the mouth-arms very deep" canals running to peripheral canal, forking several
(Mayer, 1910, after Haeckel). Stomach circular, times, the branches continuing near-parallel with
wide, occupying central 1/2 of disco Adradial original canal to margin; inter-rhopalial canal s
eanals 8, unbranehed nearly to margin, the 8 typically 3-5, increasing in number during growth,
perradial and interradial eanals producing straight but with sinuous sides, all reaching ring
basally branehes which run at ea. 45° towards canal. Ring canal same width, sub-peripheral, in
distal point of adra di al eanals (at periphery), many specimens straight to bowed inwards in each
mueh anastomosing peripherally but leaving 16 of the 16 inter-rhopalial parameres. Peripheral to
near-triangular patehes of mesoglea internally ring canal, rhopalial radial canals fork around
without gastrovaseular eanals (see diagram, Fig. rhopalia; inter-rhopalial radial canals continue
5.23). Marginal ring-canal following edge of disc in nearly to margin of lappet. Occasional anastomo-
16 shállow to nearly flat curves, in sub-crenate pat- sis in both rhopalial and inter-rhopalial canals, (?)
temo Gonads 4, in much-folded ribbon Iying near especially in older individuals. Coloration varied,
stomach periphery, sac-like when mature, project- perhaps partly consequent on diet. Recorded as
ing from floor of subumbrella. No subgenital ostia. milk-white disc and tentacles, with oral arms and
Coloration: 16 radial exumbrellar streaks reported gonads milky-white to lemon-yellow (Larson,
(Haeckel), comprising numerous dark brown pig- 1986; Montsalve, 1996); radial canal s orangelbuff
ment spots and nematocyst warts arranged in star- (Comelius, pers. obs., Bay ofBiscay); in formalin,
pattem aligned with ridges in underlying mesoglea. exumbrella yellowish-milky, canal system
milky/transparent, gonads dull orange to ochre,
Subfamily Sthenoniinae mouth-arms lighter (M ayer, 1910); disc milky-
Tentacles arising from the subumbrella in 8 or 16 lin- yellowish, gonads sepia (Mianzan, 1986b); disc
ear c1usters; mouth-arms broad, curtain-like; radial colorless to bluish, canals and mouth-arms yellow,
canal s both branched and unbranched; gonads sac- tentacles light violet, gonads reddish-brown
like, protrusive; subgenital pits absent (after Kramp, (Naumov, 1961); stomach "brilliant orange, rest of
1961; Larson, 1986). disc colorless excepting yellow warts on lappets
and on adjacent exumbrella" (Mills el al., 1996).
Genus Phacellophora
Rhopalia 16; tentacles in 16 groups, altemating with Subfamily Poraliinae
the rhopalia; radial canal s in rhopalial radii branched, Tentacles fewer than 50, spaced apart; mouth-arms
but in tentacular radii unbranched; gonads 4. more than 4, short, narrow, tapered, slightly folded;
gonads arranged in continuous ring around stomach; Subfamily Stygiomedusinae

radial canals unbranched (after Larson, 1986). Radial canal system much branched and anasto-
mosed; with ring canal; marginal tentacIes absent.
Genus Poralia
Poraliinae with the characters of the sub-family. Genus Stygiomedllsa
• Poralia rllfescens VanhOffen, 1902 (Fig. 5.21) Stygiomedusinae with the characters of the family, to
(partly after Bige10w, 1909; RusselI, 1962; which may be added: mouth-arms longer than disc
Larson, 1986). Disc flat with short, vertical diameter; altemating branched rhopalial and
peripheral skirt. JelIy thin, in life disc flexing unbranched interrhopalial radial canals a11anastomos-
from flat, through hemispherical to fulIy spherical ing towards periphery; young brooded intema11y.
(with opposite sides of margin meeting) in overa11 Stygiomedllsa gigalltea (Browne, 1910, as
shape; the skirt supported by strong ribs with Diplulmaris) (Fig. 5.24) (mainly after a 50 cm
equal intervening gaps; one extra-circular radial diameter specimen described by Russell and Rees,
canal and one marginal tentacle per rib; 1-2 1960. S. fabulosa Russe11, 1959, and S. stauchi
lappets and ribs between adjacent rhopa1ia11appets Repelin, 1967, are conspecific (Comelius, 1973;
(? depending on stage of growth). Exumbrellar Larson, 1986). A deep-sea species, sometimes
rhopalial pits somewhat conspicuous, situated upwe11ed. Diameter recorded up to 140 cm, most
near upper margin of skirt, rhopalium short, specimens 50-100 cm. A large, flat, deep brown to
stubby. Rhopalial lappets long, pointed, straight- maroon medusa; lacking marginal tentacles;
sided, without gap between; tentacular lappets with very long, narrow mouth-arms; with annu-
shorter, but arising from position of end of thick lar exumbrellar groove around central half of
ridge-like extension of mesoglea so terminating at disc, approximately outlining stomach; and, peri-
same level as rhopalial lappets, similarly pointed. pheral to this, prominent anastomosing gastrovas-
Marginal tentacles ca. 24, altemating in ones and cular canals altemate ones of which in many, but
twos with the 16 rhopalial lappets (with some not all, specimens form a back-to-back-B partero.
inconsistencies in sequence in most individuals), Disc wide, raised in centre; in life, sloping out to
easily lost through damage. Marginal tentacIes perimeter with annular groove halfway to margin;
fine, in life approx. same length as disc diameter, when out of water, flattening under own weight;
arising from exumbre11a surface 1/5 of distance surface smooth. Je11y thick and firm, thickest
between margin and ring canal. Mouth-arms 8, beyond margin of stomach (i.e. immediately peri-
arranged in pairs, tapering gradua11y to blunt pheral to annular groove), tapering periphera11y.
point; length in life approx. 1/2 be11 diameter; Marginallappets shallow, rounded, 40 rhopalial and
fused sub-basa11y for short length, basal spaces 20 inter-rhopalial. Rhopalia 20, simple, situated at
between forming 8 heart-shaped genital ostia. ends of "B-pattem" radial canals. Subumbre11ar
Stomach circular, occupying central 1/3 of disco musculature annular, weakly developed, in area of
Radial canal s ca. 48 (depending on stage of thickened je11y.Mouth cross-shaped, small. Mouth-
growth), wide, slightly sinuous, irregular, _ arms 3 or more times as long as disc diameter, uni-
leading to wide peripheral ring canal. The 16 ted basally to form a tube round in section and
per-, inter- and adradiaJ canals continuing beyond somewhat massive; distal ca. 3/4 comprising two
ring canal for short distance then forking on either thin, flat flanges with groove between; the margins
side of each rhopalium; between two rhopalial finely sca11oped, diameter of crenations ca. 5 rnm.
radii 1-2 radial cana1s extending into tentacu1ar Stomach large, much modified to encIose four
lappets. Gonads 8 (abnormal numbers recorded), quadrant-shaped brood chambers each having
arranged in (near) continuous ring around thick-bordered ca. 15 mm diameter aperture
periphery of stomach, diameter of gonad ring through stomach floor. Numerous gastric cirri, ca. 5
approx. 1/3 of disc diameter. Ground-co10r of epi- rnm long, armed with nematocysts, surrounding
dermis of disc paJe orange-translucent, radial each brood pouch. Radial canals 40, conspicuous,
cana1s whitish, mouth-an11S pale lilac, marginal straight to variously tortuous; gastrovascular net-
tentacIes colorless. Red pigment granules reported work in two zones; altemate radial canals crossing
throughout mesoglea. inner zone unbranched (inter-rhopalial canals) and
without lateral connections, those between (rhopa- central mound bearing 10-50 large irregular
lial canals) having broad, looping side-branches warts, surrounded by a moat; only peripheral
typically in back-to-back-B pattern (this irregular region of bell flexing. Disc wider than high, to ca.
and not discemible in some specimens). In outer- 50 cm; varied in shape from filamentous to thick, in
most zone all 40 radial canals distinct, and all simi- some specimens angular, projections, pointed to
larly connected by anastomosis of irregular mesh rounded; peripheral flexible skirt region without
about 1/2 as large as inner "B-pattem" zone. Ring marks, grooved radially. Marginal lappets 50-70,
canal following the divided margino Gonads 4, in rounded, clefts between shallow, forming crenate
stomach pouches, gametes perhaps developing in edge; rhopalia 8, set in from margino Mouth-arms 4,
germinal line on outer surface of brood pouches. each forking basally making 8 main groups of
Embryo brooded in unique cyst, probably a modi- branches; much branched, extending slightly
fied scyphistoma, with umbilical connection to gas- beyond disc margin; mouthlets numeroU's, 10-50
trovascular system; young released viviparously at short clubs and long filaments between the branch-
large size. Coloration: generally "a deep brown-red es, filaments up to 2-3 times disc diameter. Radial
or plum color typical of deep-sea animals", pigment canals 32 (8 main plus 8 groups of 3), joined
cells present right through jelly (Russell and Rees, peripherally by anastomosing network; no ring
1960); radial canals and anastomoses whitish, con- canal. Coloration: exumbrella b1ue through lilac to
trasting and conspicuous. mauve, translucent peripherally; mouth-arms
brown, short clubs white, filaments colorless.
Order Rhizostomeae (Fig. 4, 5.26-5.32) Variation: proportional extent of central exumbrel-
Margin with lappets; without tentacles; with rhopalia lar region, and shape, size and number of the warts,
between some marginal clefts. Mouth typically having greatly varied. Related Cotylorhiza (Mediterranean;
not a single central opening but numerous, in some spe- ? Canaries, possibly Africa south of the equator)
cies thousands, of minute mouthlets on eight (4 x 2) lacks exumbrellar projections, dominant coloration
adradial, fleshy, branched, arm-like appendages arising yellow.
from centre of subumbrella (central "normal" mouth-
opening also present in some species) (after Kramp, Farnily Mastigiidae (Fig. 4, 5.27)
1961). Subumbrellar muscles annular; rhopalar canals 8; with
ring canal; genital porticus continuous; subgenital ostia
Suborder Kolpophorae broad; arm-disk quadratic, with four primary canals.
Mouth-arms dichotomous, in cross-section triangular
or three-winged; peripherally from stomach, a net- Genus PhyIlorhiza
work of anastomosing cana1s cornmunicating direct Mouth arms broad, laminar, J-shaped, with large win-
with stomach, altemating with radial canals; rhopalial dow-like openings in lateral membranes; with numer-
pits smooth, radial folds lacking; subgenital ostia ous fi1aments in lower portion. Rhopalar canal s with-
without papillae. out comunication with intracircular meshwork of
canals.
Family Cepheidae (Fig. 4, 5.26) PhyIlorhiza punctata von Lendenfeld, 1884 (Fig.
Numerous (more than three) inter-rhopalar radial 5.27) (partly after Mayer, 1910). A medium-sized
canals per paramere; ring canal lacking; rhopalia 8; medusa slightly flatter than a hemisphere when
subgenital cavities 4 or more, more or less separated; mature, brown with conspicuous white spots,
subgenital ostia small, round; stomach octagonal, mouth-arms large, bushy. Bell diameter to 50 cm,
with eight primary radial canals. surface finely granular, shallowly grooved radially
between lappets; jelly thick, firm but flexible, to ca.
Genus Cephea 50 mm. Subumbrella with numerous filaments,
Three or more interrhopalar canal s in each octant; gastrula-stage embryos adhering between them.
exumbrella with I~rge, conspicuous warts in centre; Subgenital ostia oval, more than twice width of
mouth-arms with long narrow filaments. pillars between. Marginallappets typically in each
Cephea cephea (ForskaI, 1775) (Fig. 5.26) (after octant 4 truncate or notched ones centrally, flanked
Mayer, 1910; Comelius, pers. obs.) Massive; with on each side by two narrower pointed ones and a
single rhopalar lappet on each side; the centrallap- Lycllllor!liza lucerna Haeckel, 1880 (?=
pets partly fused basally by membranous web. Catostylus cruciatus) (Fig. 5.28). Umbrella hemi-
Rhopalia 8. Circular subumbrellar muscle fields spherical when young (up to ea. 7 cm diameter),
interrupted over radial canals. Mouth-arms J- becoming gradually flatter with age, and at known
shaped, 8, 3-bladed, 2/3 longer than bell diameter; maximun of 45 cm is inverted saucer-shaped, ca.
the 3 leaves bearing mouthlets, pinnately branched 1/6 times as high as wide. Jelly flexible, thin.
but shallowly; distally bearing blunt, tapering fila- Exumbrellar surface with numerous, closely-
ments, extensile up to length of mouth-arm. packed low conical projections ca. 1 rnm wide,
Stomach cross-shaped; radial canals 8, connected barely visible and barely noticeable to finger-tip
subdistally by circular canal; peripheral to circular touch. Marginallappets steep-sided triangular,
canal, a finely anastomosed network connecting clefts between rounded; ends of lappets in older
radial canal s and extending into lappets; centripe- specimens (above ca. 25 cm) possibly dividing to
tally, a similar zone connecting with the 4 perradi- form additional lappets, but 4 inter-rhopalar lap-
al radial canal s and the stomach, but not with the pets per octant in younger specimens. Rhopalar
interradial canals. Coloration: jelly of umbrella lappets tiny, parallel-sided, pointed. Rhopalia 8,
clear except for many embedded yellowish-brown white, peripheral, in deep clefts in margino
zooxanthellae sub-epidermally giving pale to mid- Subumbrellar musculature a continuous ring
brown color to bell (? to greyish-blue), darkest (unbroken over radial canals) of ca. 20 circular
peripherally; also many groups ofminute refractive muscle-folds, in zone between ring canal and
whitish bodies giving spotted appearance; mouth- stomach. Mouth-arms in 4 pairs, laterally flattened
let regions deep brown, filaments colorless. basally, length ea. 2/3 disc diameter; proximal half
Formerly probably just Indo-Pacific, spread to iso- laterally flattened, with mouthlets on inner side
lated N and S Atlantic localities by introductions, only; distal half 3-vaned, central vane and two
some persisting several years; several South laterals, mouthlets in band along the much-folded
American records. Mastigias scintillae Soares edges. Numerous thick filaments among the
Moreira (1961) is conspecific. mouthlets, deciduous. Mouth-arm disc massive,
4 pillars supporting it, each broad-based, tapering
Suborder Daktyliophorae slightly downward; the gaps between twice
Mouth-arms three-winged; network of anastomosing breadth of pillars. Stomach with outline of
canals issuing from primary ring canal (i.e. not from Maltese cross, the arms and comers rounded and
stomach), cornmunicating with radial canals and only broadened slightly peripherally; gonads rim-
through them ultimately to stomach; subumbrellar ming the stomach, cream to white; no central
muscles annular; rhopalar pits with radial folds; sub- mouth. Eight rhopalar radial canals, all eonnecting
genital ostia narrowed by papillae. to margin (contrary to Mayer, 1910); joined sub-
peripherally by circular canal 1/3 of radial dis-
Superfamily Inscapulatae tanee from margin (see diagrams in Vannucci,
Scapulets lacking; with permanent ring canal; radial 1951; Mianzan, 1986b). From ring canal (in older
canals 16 or 32, not all extending to umbrella margin; specimens) typically 4 broad centripetal canals
with a continuous genital porticus. ron part-way to stomach but do not connect with
it; spaces between them slightly narrower than
Family Lychnorhizidae (Fig. 4, 5.28) width of canals, some of which dilate just centrip-
Radial canals 16, between them centripetal canals etal to ring canal. Extra-circular network fine, uni-
which typically end blindly and which do not anasto- formly anastomosed; in some radii, continuation
mose; mouth arms broad, much folded. of rhopalar canal to periphery not irnmediately
clear but typically just diseemible; rhopalar radial
Genus LYc!lIlor!liza canals broader intemally to circular canal than
Mouth-arms lacking terminal clubs, with or without between it and margino Color: exumbrella color-
filaments; eight radial canals reaching to margin, less to uniform very pale brown (36°S, Argentina),
eight only reaching ring canal; in each ofthe 16 spac- or with varied density of elongate, variegated
es 2-4 centripetal canals. patches quite asymmetrically arranged, brown
to pale brown (southem Brazil), in one published Family Rhizostomatidae (Fig. 4)

photograph (Caribbean, Humann, 1992) shown Mouth-arms coalesced in proximal portion only;
deep blue. Lappets slight1y darker brown than without primary mouth-opening; manubrium with a
exumbrellar patches, deep blue (even if exumbrel- complex canal system; distal portion of arms 3-
lar patches brown). Mouth-arms colorless; regio n winged, usually with a terminal club.
of mouthlets pale brown in some specimens, col-
orless in others. Color variation is great. There is Genus ElIpilellla
some variation in the centripetal canals, occasion- No fi)aments on mouth-arms, clubs, or any other
al ones being forked or joined to a neighbour. appendages.
There is so me variation also in the shape of inter- ElIpilellla ;llexpectata Pages, Gili and Bouillon,
rhopalar lappets (see description), some apparent- 1992 (whence details taken) (Fig. 5.31). Large,
ly dividing from the four reported as usual in the hemispherical, recorded to ca. 400 mm diameter;
species to form additional ones in older individu- infrequently reported. Umbrella slightly higher
als, which have perhaps not often been examined than wide, jelly thickest centrally; exumbrellar
in detai1. surface finely granular. Marginal lappets typi-
eally 4 double ones (8 in total) in eaeh oetant,
Family Catostylidae (Fig. 4, 5.29) triangular, pointed; rhopalar lappets 1/3-1/2.as
Radial canals 16, between each and next an intracircu- long, narrow, pointed. Rhopalia 8. Mouth-arms
lar network of anastomosing canals communicating fused basalIy for "somewhat more than" halftheir
with the ring canal; rhopalar radial canals 8, extending length; distal part of mouth-arms on three vanes,
to umbrella margin; the interrhopalar canal s also 8, two outward and one inward, all with mouthlets on
extending only to ring canal; mouth-arms pyramida1. inner faces, directed downwards; distal regions of
each bearing "several inwardly-direeted,
Genus Catostyllls hooked-shaped, solid terminal appendages"
Broad intracircular anastomosing network of canals in (Haeckel, 1880, as E. scapulare; Pages et al.,
direct connection with ring canal and interrhopalar 1992) bearing many mouthlets; no mouth-arm
canals (not with rhopalar canals) and ring canal; clubs or filaments. Radial canal s 16, continuing to
mouth-arms without appendages. margin (some perhaps indistinctly through net-
Catostyllls tag; (Haeckel, 1869) (Fig. 5.29) (main- work), with circular canal some distance from
ly after Mayer, 1910.) Large, hemispherical, liUle margin; finely anastomosed network peripheral,
known. Bell massive, flexible but firm; recorded and broader network centripetal, to circular canal,
to 65 ém; surface smooth, except for shalIow eonneeted to radial eanals along whole length
forking grooves in peripheral ea. 1/6 (?) of radi- external to mouth-arm bases. Coloration: "uni-
uso Marginal lappets up to 8 per oetant, trian- formly transparent excepting gonads and mouth-
gular, plus 2 narrow rhopalar lappets approxi- lets" (Pages et al., 1992), or with large blue patch-
mately same length; rhopalia 8. Mouth-arms 8, es about 1/5 from bell edge (specimens deposited
approximately as long as bell diameter, mouthlets in The Natural History Museum, London). E.
on distal 3/4; tapering neatly to somewhat point- clallstra, Haeckel, 1880, and E. scaplllare
ed end; without filaments or clubs. Gonads rim- Haeckel, 1880, based on lndo-Pacific material,
ming the cross-shaped stomach. Color: recorded await modem assessment.
with exumbrella bluish-white through off-white to
pale cream, also brown; peripheral exumbrellar Genus Rh;zostoma
grooves reported "reddish or brownish purple"; Scapulets small, manubrium short; mouth-arms lack-
gonads outlining stomach-cross once reported red. ing lateral clubs and filaments, but each with single,
c1ub-like terminal appendage; typically with ring
Superfamily Seapulatae canal; intracircular network of canals with few and
Scapulets present, eight pairs on upper arms; with or large meshes.
without ring canal; radial canals 16, all extending to Rhizostoma pllllllO (Macri, 1778) (=R. octopus)
umbrella margin; with 4 separate subgenital cavities. (Fig. 5.30). The "football jellyfish". Bell hemi-
spherical, diameter typically to ca. 40 cm, excep-
tionally to ca. 90 cm. Exumbrella surface finely dish-brown or blue. All other structures translu-
granular, jelly thick, stiff over central 2/3, thin- cent, colorless to pale greyish-white. Validity of
ner and flexible in outer 1/3; translucent. Marginal Rhi::.ostoma ¡u/eum (Quoy and Gaimard, 1827) is
lappets 8-12 per octant. Rhopalia 8; rhopaliallap- unclear.
pets smaller than inter-rhopalar, pointed.
Subumbrellar musculature in eight distinct periph- Family Stomolophidae (Fig. 4, 5.32)
eral muscle fields. Manubrium short, massive, Mouth-arms coalesced along whole length; with per-
translucent; concealed by 16 scapulets upon it. manent primary mouth-opening; manubrium with
Each scapulet inverted Y-shaped in section, bear- limited canal system; lower arms dichotomously or
ing numerous mouthlets. Mouth-arms inverted Y- irregularly branched, without terminal clubs.
shaped in section, supporting two long, massive,
outwardly-directed blades also bearing numerous Genus StolllolophllS
mouthlets. Terminally on each is a large, trans- Ring canal lacking; rhopalar canal s wide; scapulets
lucent terminal club. Stomach occupying central large, canal system elongate, small-meshed, some-
1/3 of bell, roughly square with concave sides; what separate from radial canals but joined in places.
from it 16 substantial radial canals connect to bell StolllolophllS lIleleagris L. Agassiz, 1862 (Fig.
edge; younger specimens have narrow ring canal 5.32) (part1y after Mayer, 1910). The "cannonball
which follows closely outline of each marginal jellyfish" of U.S.A. Bell slightly more than a
lappet; in many older specimens ring canal appar- hemisphere and slightly steeper-sided, solid, up
ently absent in places and perhaps in some is com- to ca. 185 mm diameter; mouth-arms hort, fused
pletely lacking; an intermedia te ring-canal about together, lacking clubs or filaments. Jelly thick,
1/3 of radius in from margin, broad; centripetal to rigid. Exumbrellar surface smooth, with minute
this a coarse, irregular anastomosing network of nematocyst warts. Marginal lappets up to ca. 14
canals, connecting only with intermediate ring- per octant, short, rounded; rhopalar lappets up to 3
canal and not with radial canals. Peripheral to times longer, pointed. Rhopalia 8, in conspicuous
intermediate ring-canal a similar but finer mesh- pear-shaped pits on margino Subumbrellar circular
work, its branchings becoming increasingly more musculature interrupted over radial canals; radial
fine towards perimeter. Gonads 4, each a much musculature in 16 outward-pointing triangular
convoluted lobe fundamentally forming most of a patches, the bases against manubrium. Mouth-
circle but this not obvious due to its convolutions. arms fused along most of length to form solid
In older animals surface of gonad bearing grooves "manubrium", grooved externally, projecting
extending to its edge. Coloration: bell translu- slightly beyond perimeter; free ends bifurcating,
cent, surface recalling frosted glass; typically flared; unusually in rhizostomes, a central mouth
pale-grey to colorless but some specimens or present permanently; basally on each mouth-arm
populations also variously described as washed two scythe-shaped scapulets (16 in all) bearing
with pale milk-white, opalescent, and rose-red. In mouthlets. No clubs or filaments. Stomach occu-
near-mature and older specimens the largely white pying central 1/5 ofbell; radial canals 16, connect-
gonads show diffusely through as four large inter- ed by broad anastomosing network over most of
nal patches. Marginal lappets neatly and nar- bell; no ring canal. Coloration greatly varied:
rowly edged with ultramarine blue, violet, or exumbrella brown, cream, milky bluish to yellow-
brown Hne of dense color, constant in width and ish (lavender to pink reported); some specimens
following margin; rhopalar lappets less strongly with patches and bands of color, especially a
colored, or colorless. "Frills" of scapulets and peripheral band; gastrodermis reported dull yel-
mouth-arms pink off west Scotland, also bluish, low; exumbrella surface may al so be reticulated
violet, yellowish and reddish recorded; and in ter- brown, especially densely in lower 1/3 of bell
minal clubs of mouth arms canals may be bright where also occur numerous white to yellowish
blue, yellowish-brown, pink, deep purple-brown, spots; mouthlet regions brownish to pale pinkish-
or colorless. Maturing gonads tinged brown to red- fawn.
Incertae sedis South Atlantic (Rees and White, 1957b), has been
separated on lacking extemallongitudinal ridges.
Family Tesseridae
Genus Tessera (after Haeckel, 1880; Mayer, 1910). Acknowledgernents

Body conical; with 8 tentacles, 4 perradial and 4 inter-
radial. We are indebted to Prof. M. Arai (Canada) for infor-
Tessera gellllllaria Goy, 1979 (Fig. 5.34) (after mation on Atolla spp.; to Drs. F. Lang da Silveira and
Goy, 1979). Body conical, having prominent api- A. Carrara Morandini (Brazil) for help with
cal peduncle with nematocyst warts. Length Lycnorhiza lucerna, Chrysaora lactea and Nausithoe
reported 2.2 mm. Gastric cavity divided by 4 septa aurea; and to Drs. A. Marques and A. Carrara
each with 2-4 gastric filaments. Tentacles 8 (4 per- Morandini (Brazil) for data on Tripedalia cystophora,
radial, 4 interradial) in a primary circle, ending in Tamoya hap/onema and Chiropsalmus quadrimanus.
nematocyst wart; with secondary inner-circle of 4 Dr. F. Pages (Spain) offered useful information on
bulbs. Gonads 4, reniform, hanging from gastric Desmonema sp. and Aurelia sp. from Tierra del Fuego
septa. Color not recorded. The species taxonomy (Argentina). Drs. F. Ramírez (Argentina) and Lisa-
of this genus needs revision. ann Gershwin (D.S.A.) kindly commented on the
manuscript. Mr. 1. Corbera (Spain) kindly gave per-
Family Tetraplatiidae mission to redraw several ofhis excellent illustrations
from Pages et al. (1992). Mrs. 1. Outumuro and R.
Genus Tetraplatia González (Argentina) kindly provided photos and
Body bipyramidal, with ciliated outer surface. Gonads video tapes of Chrysaora plocamia from Puerto
4, each with marked oral and aboral bilobed portion; Madryn (Argentina). Dr. R. Larson (U.S.A.) allowed
bilobed lappets 4, each lobe with a statocyst. use of so me illustrations from Larson (1986), and the
Tetraplatia volitans Busch, 1851 (Fig. 5.33) (after American Geophysical Union, Scientia Marina
Russell, 1970; Pages et al., 1992). Tiny, aberrant, (Spain), Masson (France), Gustav Fischer (Germany),
information scant; anatomical homologies SPB (Holland), Brazilian Joumal of Oceanography,
obscure. Body greatly flexible, elongate-subocto- and Cambridge University Press (U.K.) released
hedral when swirnrning, shape like two 4-sided copyright permission for some of the images repro-
near-pyrarnids base-to-base; length reported to duced or redrawn here. Dr. R. Elías (Argentina) drew
ea. 15 mm; equatorial groove demarcating oral many of our illustrations. H.W.M. deeply appreciates
porfion from aboral, oral noted as 2-5 times length help from crews of the RJVs. "Meteor", "Evrika",
of aboral (but see below). Four long ridges on "Oca Balda" and "E. Holmberg" during cruises from
edges along whole length, continuous across equa- which much of the information presented in this chap-
torial groove; prominence varied, bearing nemato- ter was obtained. This is contribution N°I004 of
cyst tracts, with other tracts and scattered nemato- INIDEP.
cysts betweeen ridges. Marginal lobes (? lappets)
16,2 pairs between adjacent ribs; digitate distally,
8 final branches per quadrant. Sense organs (rho- Suggested readings
palia?) 8, statocysts without ocelli, 2 on each side,
one between each pair of marginallobes. No ten- Systernatics
tacles or mouth-arms. Mouth opening single, sim- Franc A. 1995a. Classe des Scyphozaires. In "Traité de
ple; gastrovascular system a continuous cavity Zoologie", 3(2), Cnidaires, Ctenaires (D. Doumenc,
leading to one canal within the length of each lon- dir., 1. Bouillon, ed.), Masson, Paris, pp. 597-884.
gitudinal ribo Gonads 4, equatorial, each with 4 Franc A. 1995b. Classe des Cubozoaires. In 'Traité de
sausage-shaped lobes aligned longitudinally, two Zoologie", 3(2), Cnidaires, Ctenaires (D. Doumenc,
directed orally, two aborally. Coloration: recorded dir., 1. Bouillon, ed.), Masson, París, pp. 885-922.
as whitish to bluish-white. Nominal species T Kramp P.L. 1961. Synopsis of the rnedusae of the
chuni Calgren, 1909, alone recorded from the world. 1. Mar. Biol. Ass. U.K., 40:1-469.
Orde.r Coronatae
Famlly Atollidae
tingle lenlacle
rger lhan resl
In all species) ~ ,-
E
u
o
@ 5.1 e
Atolla wyvillei
I S.2b
subumbrellar view
Atolla chuni
S.2e
"
S.3b S.3e
subumbrellar view
Atolla parva
slomach pouch
S.4a
S.4b
subumbrellar view
~T~
~
S.4e
Atolla vanhoeffeni
Order Coronatae
Family Nausithoidae
S.Sa
.-",,\. .' (-. \
~-\; \;: \'"... ~~-(-. !

t--'.-:-:
• i
, '::: , .~---'.
-;~gi2:~; .••..
S.6b
('"'"
Nausithoe aurea .-.-.< "l~", - ,,"w_, subumbrellar view
S.7b
Nausithoe punctata subumbrellar view
Family Paraphyllinidae
I
S.8a
S.8b
subumbrellar view
Paraphyllina ransoni
Order Coronatae
Family Periphyllidae Family Linuchidae
coronal
groove
~I
5.10
5.9 Linuche unguicuJata

Periphylla periphylla
Order Semaeostomae
Family Pelagiidae
Chrysaora hysoscella
I~
Chrysaora plocamia
D
Chrysaora quinquecirrha
E
E
5.15c
Chrysaora lactea
ephyra
Order Semaeostomae
Family Cyaneidae
Desmonema gaudichaudi
.Desmonema chierchianum Drymonema dalmatinum

Order Semaeostomae
Family Ulmaridae
5.20a
5.19a
Phacellophora
camtscnatica
5.21 b
5.21 a
Poralia rufescens
E
u
o
5.23a
\
Discomedusa lobata
Diplulmaris antarctica
Order Semaeostomae
Family Ulmaridae
~I
~I
1
"-
..
5.25b
Deepstaria enigmatica
~
Order Rhizostomeae
Family Cepheidae I Family Mastigiidae
Cephea cephea
Phyllorh;za punctata
Family lychnorhizidae
Family Catostylidae
I~
~
Lychnorh;za lucerna Catostylus tag;
Order Rhizostomeae
Family Rhizostomatidae
~I 5.30a
Rhizostoma pulmo
Family Stomolophidae
o .
51
Eupilema inexpectata
5tomolophus meleagris
Incertae sedis
E
E
E
E
o
Tetraplatia volitans Tessera gemmaria
Fig. 5. Seyphomedusae: sehematie external views and details of diagnostie eharaeters of the adults of the speeies reeorded in the South
Atlantie. Seale bars equal I cm unless otherwise indieated.
Sourees: from Cornelius (1997e), redrawn: 5.10, 5.22a, 5.26a; from Goy (1979, I!:) Masson, Paris): 5.7b, 5.34; from Larson (1986), some
redrawn: 5.2b, e, 5.3a and 5.4a (eomposite by R. Elías), 5.4b, 5.17a (redrawn from photograph), 5.21a, 5.21 b, 5.22b, 5.24a (eomposite by R.
Elias), 5.24b, 5.25a, b: from Mayer (1910), redrawn: 5.la, 5.2a, 5.7a, 5.lla, 5.14a, 5.15a, 5.30a; from Mianzan (l986b): 5.15e, 5.20e, 5.28a,
5.32a; fr;m Mianzan (1989): 5.9; from Grenaeher and Noll (1876), redrawn: 5.29a; from Pages el al. (1992), so me redrawn: 5.12a, 5.23a, b,
5.3Ia, b, 5.33 (copyright I!:) J. Corbera); from Repelin (1964): 5.1 b, 5.3b; from Russell (1970): 5.le, 5.3e, 5.4e, 5.5a, b; 5.8a, b; from Silveira
and Morandini (1997): 5.6a, b; from Soares Moreira (1961): 5.27a, b; from Stiasny (1921): 5.28b, 5.32b; from Stiasny (1923): 5.26b, 5.29b,
5.30b, 5.3lb; original: 5.11b; 5.12b, 5.13a (from photos and video by J. Outumuro and R. Gonzáles), 5.13b, 5.14b, 5.15a, b, 5.16a, b, 5.17b,
5.18a, b, 5.19a, b, 5.20a, b, 5.27a, 5.27b (illustrated by R. Elías and H. Mianzan).
Lal'son R. 1986. Pelagic Scyphomedusae (Scyphozoa: Plathelminthes, Nemertini, Entoprocta, Nemathel-

Coronatae and Semaeostomae) of the Southem minthes, Priapulida. G. Fischel', Stuttgart, pp. 11-3.
Ocean. Biol. Ant. Seas 14, Ant. Res. Ser. 41(3),
Amer. Geophys. Un ion, pp. 59-165. Biology
Mayel' A.G. 1910. Medusae ofthe Wol'ld. Volume III. Al'ai M.N. 1997. A functional biology of Scyphozoa.
The Scyphomedusae. Carnegie Institution, Chapman and Hall, London, pp. 1-316.
Washington D.C. Pp. i-iv, 499-735. [Facsimile Russell F.S. 1970. The medusae of the Bl'itish Isles.
edition published in 1977 by A. Ashel' and Co. n. Pelagic Scyphozoa. Cambl'idge Univ. Pl'ess,
RV., Keizel'sgl'acht 526, Amstel'dam]. Cambridge, pp. 1-284.
Wemel' R 1984. Stamm Cnidal'ia, Nesseltiel'e. In:
"Lehrbuch del' speziellen Zoologie" (Cool'dinated by Ecology
A. Kaestnel'), Band I: Wil'bellose Tiel'e H.E. (Grunel', Al'ai M.N. 1988. Intel'action of fish and pelagic colen-
ed.), 2 Teil: Cnidal'ia, Ctenophol'a, Mesozoa, tel'ates. Canadian J. Zool., 66:1913-1927.
Histology Venoms and stinging

Harrison F.W., Westfall lA. 1991. Microscopic anat- Williamson l.A., Fenner P.E., Bumett lW., Rifkin
omy of invertebrates. Vol. 2. Placozoa, Porifera, l.F. (eds). 1996. Venomous and Poisonous Manne
Cnidaria and Ctenophora. Wiley, New York, pp. Animals: a medical and biological handbook. Surf
1-436. Life Saving .Queensland Inc. and Kensington
N.S.W., University of New south Wales Press,
Brisbane, pp. 1-580.
í
Ctenophora
Hermes W Mian::.an
Introduction large oral lobes and auricles (Fig. 1B), and by the
absence oftentacular sheaths. Members ofthe order
Ctenophores are a small, well defined phylum of bira- Cestida have a ribbon-like body and modified tenta-
dially symmetrical animals with about 150 species dis- cles (Fig. 1G).
tributed world wide. They are exclusively marine,
planktonic or benthic (one order), solitary organisms Most of the ctenophores are simultaneous hermaphro-
with gelatinous soft bodies. The general body plan is dites, capable of self fertilization. Only the genus
closely akin to, but more advanced than, that of Ocyropsis has been reported as dioecious (Harbison
Cnidaria (Harbison and Madin, 1982; Harbison, 1996). and Miller, 1986). The Platyctenida, as an exception
to the rule, are protandrous, and asexual reproduction
The locomotory apparatus is composed of eight sym- has also been recorded (Harbison, 1985).
metrical tracks (subtentacular and substomodeal
comb rows), distinguished by fused ciliary plates Bioluminescence is a common feature in most of the
(ctenes) on the body surface (Hernández-Nicaise and species, occurring generally along the comb rowS. In
Franc, 1993) (Fig. lA, D). Metachronal beating ofthe some cases, clouds of bioluminescent substance are
ctenes produces locomotion in all but creeping forms. generated, probably as a means to avoid predation
Ctenophores are the largest animals using cilia for (Harbison, 1996).
locomotion. Swimming is coordinated by the apical
sense organ or statocyst (Fig. lA). Tentacles, when All ctenophores are carnivorous, feeding on zoo-
present, usually bear adhesive structures called collo- plankton of various sizes, including copepods,
blasts (Fig. 1C); colloblasts are designed to capture euphausiids, fish eggs and larvae. The Beroida also
prey, and are another unique feature ofthe phylum. feed on gelatinous forms such as other ctenophores,
medusae and salps (Fraser, 1962; Swanberg, 1974;
The gastrovascular system (Fig. 1E) can be divided Kremer, 1979; Alvariño, 1985; Purcell, 1985;
into two portions: axial and peripheral. The axial por- Mianzan and Sabatini, 1985; Monteleone and
tion consists of a large stomodeum (pharynx), the Duguay, 1988; Hoffmeyer, 1990). Their role in
infundibulum (gut), infundibular canal, and the marine trophic webs is more complex than previously
anal canals (excretory) and pores. The peripheral thought. Long considered a carbon dead-end in
portion consists of perradial, interradial, adradial marine food webs, recent studies indicate that the
and meridional canals, the tentacular canal and the group can be consumed by various fish species
paragastric canals. Except for the adradial and (Oviatt and Kremer, 1977; Arai, 1988; Ates, 1988).
meridional canal s, any of them may be missing Recent information from the Argentine continental
(Harbison, 1985). shelf (35-47°S) stresses the seasonal trophic signifi-
cance of ctenophores for 69 fish species between the
Ctenophores comprise two classes: Nuda (without coast and 200 m depth (Mianzan el al., 1996). More
tentacles during their entire life), with the single order complex trophic relationships also imply a more
Beroidea (Fig. lF); and Tentaculata, with five orders. active role in the transmission of parasites; the first
The most primitive Tentaculata, the Cydippida, keep record of a metacercarian encysted in ctenophores
their tentacles during their entire life (Fig. lA). The was recently published (Martorelli, 1996).
order Platyctenida is, with few exceptions, a benthic
group, having lost the ctenae as an adaptation to bot- Ctenophores are among the most difficult marine ani-
tom condition. The order Thalassocalycida is mono- mals to study (Harbison, 1986), and represent the
generic; it was proposed for forms that did not fit the greatest challenge to conventional oceanographic
definitions of the other orders (Harbison, 1985). The sampling (Harbison el al., 1978). Standard "quantita-
order Lobata is easily recognizable by the presence of tive" samplings give the impression that ctenophores
SOllfh Atlantic Zooplankton.

1999 Backhuys Publishers, Leiden, The Netherlands
562 Ctenophora
Type Cydippida Type lobata
mouth
(; gastrovascular system
4)
Type Cestida subtentacular
substomodeal
~ combrow
~ combrow
tentacle
7~"omOdeum
mouth
Ctenophora 563
represent an insignificant fraction of the open ocean pileus and Beroe eucumis are common to both
fauna (Harbison, 1986). Of the few specimens that (Gibbons el al., 1992).
can be collected with nets, only a small fraction can be
preserved in recognizable fashion (Harbison, 1986), This chapter is aimed mainly at students and plancto-
mainly Cydippida and Beroida. In situ techniques like logists working with net samples. The information is
diving or submarines, on the other hand, have proven based on local and historical papers, and on personal
to be very effective for collecting these delicate ani- collections. A diagnosis and schematic figures of the
mals, resulting in the discovery of a number of new external appearanee (based on photographs, living
speCles. specimens or published information) is presented for
each taxon considered herein.
It is well known that ctenophores occur from time to
time in vast numbers in coastal and estuarine regions
of all oceans (Fraser, 1962; Harbison el al., 1978); and Methods
a number of authors have concluded that neritic spe-
cies are important predators that can regulate the Neritic ctenophores can be collected with nets, prefe-
abundance of certain herbivores (Harbison el al., rably with large mouth areas. Species of Beroe,
1978; Deason, 1982; Feigenbaum and Kelly, 1984; Hormiphora, Pleurobrachia and Mnemiopsis are stur-
Mianzan and Sabatini, 1985). Recently, new immi- dy enough to survive trawling aetion. A gentle plank-
grant species, sometimes referred to as biological pol- ton net vertical haul is the best method as it causes
lution, have invaded environments like the Black Sea less damage to the ctenophore. Plankton nets towed
that are not prepared for such organisms, leading to horizontally or obliquely at low speed (1 knot) are
the collapse of local pelagic fisheries (Shushkina and also suitable. As with other gelatinous organisms
Musayeva, 1990; Vinogradov et al., 1989; Zaika and (medusae, salps, etc.), it is preferable to collect at
Sergeyeva, 1990; see GESAMP, 1997, for review). night, when the animal s are more abundant near the
sea surface.
Coastal species are sturdier than oceanic ones, easily
eolleeted by plankton nets, so they are well known in Pelagic ctenophores should be collected by divers
several regions of the world. In South Atlantic waters, using jars to capture the specimens (Harbison et al.,
however, eoastal faunas are still poorly known. Only 1978), or by submarines (Harbison, 1985, 1986), as
a few papers on ctenophores from the South Atlantic they are more delicate, and trawling destroys them.
have appeared (Moser, 1910; O'Sullivan, 1986; These collection methods are also suggested for stud-
Hernández-Nicaise and Franc, 1993). Recent local ies of feeding, growth, and metabolismo
inforn1ation shows that eoastal forms are an important
macroplanktonic group in the Southwestern Atlantic. Volumetric measures, as those used for the abundant
Seasonally, they can dominate other planktonic pred- eoastal species (Deason, 1982; Feigenbaum and
ators both in abundance and in biomass (Mianzan el Kelly, 1984), are recommended for quantitative stud-
al., 1996). A few species, including Mnemiopsis leid- ies. Purcell (1988) proposed the use of the tentacle
yi, M. maceradyi, Pleurobaehia pileus and Beroe bulbs, which persist intact after preservation in 5%
ovala, have been mentioned in localliterature (Balech formaldehyde, for quantification of Mnemiopsis.
and Motti, 1941; Ramírez, 1973b; Hoffmeyer, 1983;
Mianzan, 1986a, b; Mianzan and Sabatini, 1985; Immediately after collection specimens should be
Ramírez el al., 1990). separated from other organisms and carefully trans-
ferred to glass jars filled with fresh, clean seawater
Three species of ctenophores are known from the using a perforated spoon or a glass pipette.
Southeastern Atlantic and the Benguela Current Examination of live specimens is recommended. If
system (O'Sullivan, 1986), but only Pleurobraehia the specimens have to be preserved for later examina-
•••
Fig. 1. General characters used for identification of ctenophores types. E: redrawn from Chun (1880).
564 Ctenophora
tion, photographs should first be taken with the aid of The data in Fig. 2 are derived from a compilation of
a microscope in order to record structures of taxono- all literature records, and from personal research
mic value which may disappear later due to damaged records. This is an initial attempt to assemble all the
or improper preservation. It is also advisable to available information.
remove specimens of the genus Mnemiopsis from the
plankton samples with a sieve (1 cm mesh size), as As pointed out above, the distributional pattems pre-
they rapidly disintegrate after fixation leaving a gelat- sented here apply mainly to coastal ctenophores.
inous, difficult-to-handle plankton sample, unfit for Oceanic ones are still poorly known and special tech-
further zooplankton quantitative studies (Mianzan, niques are necessary for their study. Because of the
1990). extreme scarcity of ctenophore studies based on sub-
manne surveys, the biogeographic data summarized in
Ctenophores are extremely fragile animals, difficult to Fig. 2 had to draw heavily on reports based on plank-
fix and preserve, improper fixation producing a mass ton nets. Most ofthe data are based on personal obser-
of amorphous jelly. Even with great care some distor- vation and shipboard identifications following the pro-
tion and shrinkage of specimens is to be expected. cedures suggested by O'Sullivan (1986) (see Methods).
Formalin solutions are inadequate to fix ctenophores.
Early methods using solutions containing osmic or The distributional ranges presented (Fig. 2) include
chromic acid caused discoloration and sometimes dis- some record s from equatorial waters of the Northem
integration ofthe specimens. The following procedure Hemisphere; they are included because of the high
(Adams el al., 1976; O'Sullivan, 1986) is recommen- probability of finding some of these organisms in
ded for adequate preservation. Specimens must be tropical waters of the Southem Atlantic.
poured into ajar of fixative (1 g of trichloracetic acid
or I g of p-toluenesulphonic acid in 99 mI of seawa- In spite of the widespread distribution of most of the
ter) for thirty minutes. Slight changes in body color, species, which occur in several oceans, the geographic
from transparency to translucency, will be observed. ranges of neritic ctenophores in Southwestern
Then, animals must be kept in 1 mi of stock preserving Atlantic Ocean fit well with local biogeographic
solution (0.5 mI propylene phenoxetol, 4.5 mi propy- schemes proposed in the literature (Balech, 1954;
lene glycol, 5.0 mi formaldehyde 40%, and 90 mi sea- Boltovskoy, 1970; Dadon and Boltovskoy, 1982).
water) and 99 mi sea water, for five to seven days. For Warm water species off southern South America
storage, 5 mi stock solution and 95 mi sea water must include Ceslum veneris, Lampea paneerina,
be used. Never store ctenophores with Crustacea as Mnemiopsis maecradyi and Beroe ovala. They occur
the sharp spines will cut them, leading to disintegra- from southern Brazil, Uruguay, and Buenos Aires
tion. Storage temperatures should be between 5 and province (Argentina) down to 42°S. Cold water forms
20°C. Avoid knocking or shaking the jars. are represented by Callianira anlareliea,
Pleurobraehia pileus, Mertensia sp., Mnemiopsis
leidyi and Beroe el/eumis. They occur from 50-55°S to
Geographic distribution 34-35°S, associated with the subantarctic waters of
Malvinas (=Falkland) Current (Fig. 2).
Major works on ctenophore taxonomy were usually
focused on restricted geographic areas. As an exam- The wide shelf off Argentina and the presence of semi-
pie, Chun (1880) described the ctenophores from the permanent hydrographic features like the Río de la
Gulf ofNaples, Italy; and Mayer (1912) presented an Plata front (freshwater outflow), the north-Patagonian
important contribution for the Northwestem Atlantic fronts (tidal) (Brandhorst and Castello, 1971; Carreto
waters. Moser (1910) produced the first world-wide el al., 1986; Glorioso, 1987; Guerrero et al., 1997), and
distributional study, but the South Atlantic remained coastal semi-enclosed areas (Bahía Blanca) (Mianzan
with only a few species-specific records. Recently, and Sabatini, 1985; Mianzan, 1986b), generate espe-
Hemández-Nicaise and Franc (1993) published an cially adequate conditions for the development of
updated geographic distribution of Ctenophora, and it important populations of coastal ctenophores, in par-
is clear that a very noticeable gap still remains in the ticular of the genus Mnemiopsis. Members of this
South Atlantic. group preserve very poorly in net plankton samples
Ctenophora 565
Order Cydippida
* :* ' ,
.* .
-_
¡••
. ....
*..1 : ". ..: {)O
[*
.
. .... :...
------
•• * :* *:,
........ ,..............•...... * - -- --
"'f* *
---------~----- .~ 400
J: * -*'
.... '1'
* Callianira antarctica 200
* Tinerfe cyanea * Pleurobrachia pileus • Mertensia. sp.

1_ • Lampea pancerina ...................• Hormiphora plumosa • .. .: 600
Order lobata
:~ * !
.......•. ¡ , .
...........
*..~
,, ,,
.. ; 200
,, ,,
: : :
............ J.?:*:II.?
, ..... ,
* Leucothea spp . ........... , 1" + ¡ 400
• Bolinopsis spp. :* Eurhamphaea vexilligera * Ocyropsis maculata

~ Mnemiopsis maccradyi ....... j • Ocyropsis crystallina ~ Mnemiopsis leidyi 600
Order Cestida Order Beroida

? *? 1*
.......................• *':7
..................... .. : 200
, , *:
......... : ..... *-It .. L~ .....: 400
:* * * ¡ ,
*:
*: * * *: *
:• Beroe forskalii
* Cestum veneris
Beroe cucumis
600
• Velamen parallelum ......... ? ~ ~ • Beroe ovata
600 300 00 300 600 300 00 300
Fig. 2. Geographic distribution of South Atlantic Ctenophora, Sources: T cyanea: 1; L. pancerina: 2, 10; P. pilells: 1, 3,4, 10; H. plllmosa:
2; C. anlarclica: l, 3; Merlensia sp.: 5, 10; Lellcolhea spp.: 2,4,7; Bolinopsis spp,: 2,4; M. maccradyi: 7, 8,10; E, I'exilligera: 2; o. cl)'slal-
fina: 1,2,7; o. macl/lala: 2,7; M, leidyi: 8,10; C. veneris: 1,2, ,7, 9,10; V parallelllm: 2, 7; B, cl/cl/mis: 1,2,4,6,10; B. ovata: 2, 7,10; B,
forskalii: 1. l: Moser (1910); 2: Harbison el al. (1978); 3: O'Sullivan (1986); 4: Gibbons el al, (1992); 5: Richter (1996); 6: Pages (1996); 7:
Humann (1992); 8: GESAMP (1997); 9: Comelius (l997a); 10: This study,
(Mianzan et al., 1996), and because of this limitation Vertical profiles are of course scarce for south Atlantic
information on their distribution, and particularly their ctenophores. Gibbons (pers. cornm., 1996) is currently
abundance, is very scarce in the area, even though they working on Pleurobrachia pileus, and has found great-
are one ofthe most representative and abundant cteno- est densities during the night in Sto Helena Bay (South
phores in coastal southwestem Atlantic waters. Present Africa). Mianzan (unpublished data) observed diel
knowledge of ctenophores off South Africa is insuffi- migratory movement of Mnemiopsis leidyi in the north
cient for drawing a general pattem. patagonian front (42°S) from 70 m to the surface.
566 Ctenophora
Taxonomy are obtained. Genera in bold underlined characters

are those reported from SE and SW Atlantic waters,
Specific identification of ctenophores is a difficult task. and therefore treated in detail here.
Because ofthe problems outlined above, many original
descriptions were based on small individuals (juve- Phylum Ctenophora Eschscholtz, 1829
niles), often distorted or damaged by collection and Class Tentaculata sensu Milis, 1998
preservation. As a result, descriptions of local races or Order Cydippida Gegenbaur, 1856
varieties as new species are cornmon. For example, at Family Pleurobrachiidae Chun, 1880.
least 50 species were assigned to the genus Beroe, but Genera Plellrobrachia Fleming, 1822; Hormi-
most of them are synonyms (Tiffon, 1993). Care must phora. L. Agassiz, 1860; Tblerfe Chun, 1898
also be taken with juveniles of several species in the Family Mertensiidae L. Agassiz, 1860.
c1ass Tentaculata. Even adults without tentacles (Fig. Genera Calliall;ra Péron and Lesueur, 1808;
3.ge) pass through a "tentacular stage" during develop- Mertensia Lesson, 1836
ment, resembling Cydippida larvae (Fig. 3.9c), which Family Lampeidae Krumbach, 1925.
further confuses proper identification. Genus Lampea Stechow 1921
Family Bathyctenidae Mortensen, 1932.
In spite of the time elapsed since these earlier works, Genus Bathyctena Mortensen 1932
development of a classification system for Family Haeckeliidae Krumbach, 1925.
Ctenophora has been very restricted. Efforts have Genera HaeckeLia Carus, 1863; AlILacoclella
been scarce due to the expensive equipment and meth- Mortensen, 1932
ods required for an authoritative taxonomic revision Family Euplokamidae MilIs, 1987.
of Ctenophora, particularly for pelagic and bathype- Genus ElIpLokam;s Chun, 1879
lagic forms. In one submarine cruise, Harbison (1986) Family Dryodoridae (sensu Harbison, 1996).
found over 22 mesopelagic species, of which only 5 Genus Dryodora L. Agassiz, 1860
had been described previously. Thus, it appears likely Order Platyctenida Mortensen, 1912
that most of the evolution within the group has taken Family Ctenoplanidae Willey, 1869.
place in the deep ocean, where high concentrations of Genus CtenopLana Korotneff, 1886
ctenophores are found (Harbison el al., 1978) . Order Thalassocalycida Madin and Harbison, 1978
Family Thalassocalycidae Madin and Harbison,
Because Ctenophora are a soft-bodied phylum and do 1978.
not preserve in the fossil record, their phylogenetic Genera ThaLassocaLyce Madin and Harbison,
relationships are still not weJl understood. 1978
Order Lobata L. Agassiz, 1860
Harbison el al. (1978) surnmarized our present knowl- Family Bolinopsidae Bigelow, 1912.
edge ofthe group; however, as the list ofknown spe- Genera Bol;llops;S L. Agassiz, 1860; Mnem;ops;s
cies is continuously expanding, their classification is L. Agassiz, 1860
changing accordingly (Harbison and Madin, 1982; Family Leucotheidae Krumbach, 1925.
Harbison, 1985, 1996; MilIs, 1998). Genus Lellcothea Mertens, 1833
Family Eurhamphaeidae L. Agassiz, 1860.
Outline c1assification Genera ElIrhamphaea Gegenbaur, 1856;
Kiyoh;mea Komai and Tokioka, 1940; Deiopea
The following synopsis is based on the scheme pro- Chun, 1879
po sed by Harbison (1996), in particular for adult epi- Family Bathocyroidae Harbison and Madin, 1982.
pelagic forms from the North Atlantic, and Milis Genus Bathocyroe Madin and Harbison, 1978
(pers. comm.). The main diagnostic characters are Family Ocyropsidae (Les son, 1843).
those suggested by Mayer (1912), Liley (1958), Greve Genus Ocvrops;s Mayer, 1912
(1975), Harbison (1985,1996), O'Sullivan (1986) and Order Cestida Gegenbaur, 1856
Harbison and Madin (1982). Many descriptions are Family Cestidae Gegenbaur, 1856.
complemented with personal observations. As pointed Genera Cestllm Lesueur, 1813; Velamen
out above, this classification is provisional, very like- Krumbach, 1925
ly to change as more species and phylogenetic data Class Nuda sensu Milis, 1998
Ctenophora 567
Order Beroida Eschscholtz, 1829 ated from stomodeum. Tentacles with tentilla sim-
Family Beroidae Eschscholtz, 1829. ple. Color: comb rows milky opaque; ectomeso-
Genus Reroe Browne, 1756 derm glassy transparent; tentacles, sheaths and sto-
modeum milky or in some specimens dull orange.
Order, family and species level diagnoses Length 10-25 rom. P. pigmentata Moser, 1903, a
related species, was mentioned from Natal (South
Class Tentaculata (with tentacles) Africa) by Moser (1910), based on a single speci-
Order Cydippida (Fig. 3.1-3.6) men ofsmall size (2.5 mm). Extension ofthe range
Body simple, rounded, ovoid or cylindrical, in some of this species to the area considered herein should
species laterally compressed in the stomodeal plane; be based on clearly identifiable specimens.
tentacles retractable into sheaths; paragastric and Recently, specimens resembling P. baehei (accord-
meridional gastrovascular canals ending blindly at oral ing to Tyffon, 1993) were collected off Argentina
pole. down to 45°S, these records need further validation.
Family Mertensiidae (Fig. 3.1-3.2) • Horllliphora pllllllosa L. Agassiz, 1860 (Fig. 3.4).
Body strongly compressed along stomodeal axis, with Pear-shaped body with no lateral compression; ci-
aborally projecting wing-shaped keels. Openings of liary comb rows equal in length, starting some dis-
tentacle sheaths near aboral pole. Tentacles with ten- tance below aboral pole and extending two thirds
tilla. Tentacle bulbs supplied with perradial canals of distance to mouth. Tentacle sheaths pressed
from which subtentacular adradial canals arise. close to stomodeum. Long tentacles with two
• Mertellsia Lesson, 1836 (Fig. 3.1). Body laterally types of slightly yellowish tentilla and cock-
compressed, tentacular axis wider than stomodeal shaped expansion on upper side ofbase. Length up
axis. Four subtentacular rows of cilia longer than to 15-20 rom.
the 4 substomodeal, and arising from higher up. Tillerfe eyallea (Chun, 1898) (Fig. 3.5). Body
With aboral keels. Color of tentacles, comb rows slightly flattened in stomodeal plane. Gelatinous
and pole plate pink. Length up to 55 rom. tubercles around statocyst. Ciliary comb rows
Calliallira alltarctica Chun 1897 (see Callianira shorter than meridional canals. Tentacles with
sp., Fig. 3.2). Body long and narrow, quite strong- simple tentilla. Mouth expandable, stomodeum
Iy compressed in the stomodeal planeo With pro- widely eversible. Color blue. Up to 20 rom.
nounced aboral keels. AII comb rows end at level
of keels, uneven; substomodeal ones somewhat Family Lampeidae (Fig. 3.6)
longer than the subtentacular ones. Meridional Body cylindrical. Openings of tentacle sbeaths
canals extending into keels. Tentacle bases long between mouth and oral end of comb rowS. Tentacles
and narrow, located at the level of the infundibu- with tentilla. Mouth widely expandable. Pharynx
lum. Tentacle sheaths open at base of keels. widely eversible. Tentacle bu lbs supplied by separate
Stomodeum voluminous, quite long. Infundibulum tentacular canals. Adradial canals arising directly
short. Color: body clear, tentacles and stomodeum from infundibulum.
reddish. Length to 60 rom. • Lalllpea pallcerilla (Chun, 1879) (Fig. 3.6). Body
cylindrical, only a little narrower towards the oral
Family Pleurobrachiidae (Fig. 3.3-3.5) end. Mouth opening wide. Lips dilatable to a wide
Body slightly compressed in stomodeal plane, spheri- sole on which the animal moves over solid surfac-
cal, ovoid or cylindrical in shape. Tentacle sheaths es. Color transparent. Length up to 70 rom.
open aborally to infundibulum, close to apical sense
organ. Tentacles with tentilla. Tentacle sheaths close Order Lobata (Fig. 3.7-3.13)
to stomodeum or at some distance from it. Body compressed in the tentacular plane, expanded on
• Plellrobrachia pilells (O. F. Müller, 1776) (Fig. each side of mouth to forro a pair of oral lobes.
3.3a, b). Egg- to spherical-shaped body; tentacular Subtentacular comb rows giving rise to four flap-like
diameter slightly wider than sagittal. Rows of ci- auricles, usually shorter than substomodeal comb roW.
liary combs equal in length, starting near aboral Meridional canals much longer than comb rows,
pole, extending more than three quarters of distance extending into oral lobes. Paragastric canals join sub-
towards the mouth. Tentacle sheaths widely separ- tentacular meridional canals orally. Statocyst usually
568 Ctenophora
deeply sunk. Tentacles migrate orally as lobes develop, Family Ocyropsidae (Fig. 3.11-3.12)
tentacle sheaths are lost, and tentacles become reduced, Well developed oral lobes arising between mouth
surrounding mouth and extending to oral lobes. margin and statocyst. Connections of canals as in
Bolinopsidae. Tentacles absent in adults. Oral lobes
Family Bolinopsidae (Fig. 3.7-3.9) large, muscular; auricles short.
Tentacle bulbs close to mouth margin and tentilla Ocyropsis crystallina (Rang, 1828) (Fig. 3.11).
along it. Oral lobes arise above mouth margin leve!. Body laterally compressed. Oral lobes large.
• Bolinopsis vitrea (L. Agassiz, 1860) (Fig. 3.7). Comb rows short, ciliated plates few in number.
Oral lobes arising between mouth and statocyst. Four subtentacular rows only about two-thirds as
Color transparent. Length up to 80 rnm. Gibbons long as 4 substomodeal ones. Stomodeum flat; its
et al. (1992 ) observed lobate ctenophores resem- broad side constricted near middle. Color milky
bling Bolinopsis in Table Bay (Africa). translucent. Length up to 45 rnm.
• Mnellliopsis leidyi A. Agassiz, 1865 (Fig. 3.8). Ocyropsis lIlaclllata (Rang, 1828) (Fig. 3.12).
Body laterally compressed, with large lobes aris- Similar to o. crystallina. Four conspicuous brown
ing near stomodeum, generating 4 deep noticeable spots on large, prominent oral lobes. Body com-
furrows. Aboral apex not as acute as in Bolinopsis. pressed. Color of lobes whitish, translucent.
Outer surface smooth. Color usually transparent or Length up to 100 rnm.
slightly milky translucent. Length up to 100 rnm.
Alcinoe rosea from North of the Malvinas Family Eurhamphaeidae (Fig. 3.13)
(=Falkland) Islands is probably a species of Orallobes arising at mouth margin leve!.
Mnemiopsis (O'Sullivan, 1986). • Ellrllalllphaea vexilligera Gegenbaur, 1856 ( Fig.
• Mllellliopsis lIlaccradyi Mayer, 1900 (Fig. 3.9a-e). 3.13). Body resembles Bolinopsidae, with two
Body laterally compressed, similar to M. leidyi. conical processes on aboral end, terminating in
Outer surface sprinkled with numerous small warts long flexible filaments. Subtentacular comb rows
(Fig. 3.9b), well developed on sides of orallobes, extend onto these processes. Color: body transpar-
but absent between subtentacular comb rows. ent; with red vesicles dispersed on the body, pro-
Color glassy transparent. Length up to 110 mm. ducing a reddish-brown ink with bioluminescent
Balech and Motti (1941) recorded this species as properties. Length up to 100 rnm.
Bolinopsis elegans, off Necochea, Argentina. M.
leidyi and M. maccradyi have been provisionally Order Cestida (Fig. 3.14-3.15)
cited as subspecies of a single species (M. leydyi) Ribbon-shaped body strongly compres sed in tentacu-
(Harbison, pers. cornm.; GESAMP, 1997). Seravin lar axis. Subtentacular comb rows rudimentary or
(1994) considered M. maccradyi as a synonym of absent. Oral margin greatly elongated, extending
M. leidyi. along entire width of body, fringed with tentilla.
Family Cestidae (Fig. 3.14-3.15)
Family Leucotheidae (Fig. 3.10) Characters those of the order.
Body laterally compressed. Oral lobes ansmg • Cestlllll velleris Lesueur, 1813 (Fig. 3.14a, b).
between mouth and statocyst, large, delicate, with Body flat, ribbon-shaped, strongly compressed in
complex windings of substomodeal meridional the tentacular axis, with the sides strongly exten-
canals. Auricles long and serpentine. Two long pri- ded in the stomodeal axis. Subtentacular comb
mary tentacles. Tentilla forming dense barb along rows reduced, represented by 4 short tracts close to
labial ridge extending onto oral lobes. Body surface sides of sense organ. At a short distance, 4 long
covered with numerous papillae. stomodeal comb rows cornmence, extending along
• Lellcotllea 1Il11lticornis (Quoy and Gaimard, sides of arched aboral edge of anima!. Tentacles
1824) (Fig. 3.10). With the characters ofthe fam- with tentilla, closely set in a double row, extending
ily. Color of oral lobes and papillae yellow or along sides of oral edge of animal. Subtentacular
brown. Length up to 200 rnm. Gibbons el al. mesodermal canal s arise under the subtentacular
(1992) observed lobate ctenophores resembling comb rows. Color: young animals transparent,
Leucothea in Table Bay (Africa). older ones often becoming violet with greenish-
blue or ultra-marine fluorescence. Length along the
stomodeal axis: up to 2 m .
Ctenophora 569
Ve/amen paraUe/um (Fol, 1869) (Fig. 3.15a, b). Body strongly compressed. Fusions of diverticu-
Resembles a small Cestum. Without subtentacular lae of meridional cana1s very dense. Mouth open-
comb rows. Subtentacu1ar meridional canals arise ing wide, with large, half circle-shaped 1ips. Color:
from adradial canals at edge ofwings. Color trans- a1most transparent in young specimens, becoming
parent. Length up to 150 mm. slight1y pinkish with age. Height to 20 cm. Cited
as a warm water species, so its presence in
Class Nuda (without tentacles) Antarctic waters must be confirmed.
Order Beroida (Fig. 3.16-3.18)
Conical forms; with very wide mouth and stomodeum;
meridional gastrovascu1ar canals usually with numerous Acknowledgements
diverticu1ae. Without tentacles. Row ofbranched papil-
lae in form of a figure 8 around the aboral pole. On1y I am indebted to C. Milis (Seattle, U.S.A.) for taxo-
one family. Many species in these genera are probably nomic advice and checking and completing the list of
synonymous. Milis et al. (1996) concluded that much taxa, F. Pages (Barcelona, Spain) and M. Gibbons
confusion still persists in the specific identification in (South Africa) for information on ctenophores from
these genera, and that further revision is necessary. In Tierra del Fuego and South Africa, respectively. To F.
the present review I follow the nomenclature used by Ramírez (Argentina) for critically reading the manu-
Mayer (1912), since the materials ana1yzed agree well script; to P.F.S. Comelius (Nat. Hist. Museum,
with his descriptions. Although it has been suggested London, UK) for his help in obtaining the bibliography.
that Pan dora mitrata (Moser, 1910) is probably a juve- To N. Scarlatto for many translations from the German,
nile form (O'Sullivan, 1986), recent information indi- and to E. Giolitti, M. Scelzo and M. Tobio for the pho-
cates that it could be a val id species (Beroe mitrata, tographs. I a1so deeply appreciate the METEOR
Milis, 1997, pers. com.). Beroe Izyalina, is also doubtful, (SARP), EVRIKA and OCA BALDA crews for ship-
since its only description is based on one small individ- board he1p. This is contribution N°1003 ofINIDEP.
ual of 15 rom, having few projections ofthe meridional
canals with no anastomoses (Moser, 1910).
Suggested readings
Family Beroidae (Fig. 3.16-3.18)
Characters those of the order. Arai M. 1988. Interactions offish and pelagic coe1en-
o Beroe cucum;s Fabricius, 1780 (Fig. 3.16). Body terates. Canadian 1. Zool., 66:1913-1927. General
sac- or vase-shaped, often compressed in the ten- review o/ the role o/ gelatinolls plankton in marine
tacular plane. Ciliary comb rows equal in length, eeosystems.
extending from the aboral pole to about three Ates R.M. 1988. Medusivorous fishes, a review. Zool.
quarters the distance toward the mouth. Four Mededelingen, 62:29-42. Short review o/ trophie
meridional canals of each broad side connected relationships between fishes and gelatinous plank-
orally by means of oral forks of the paragastric ton.
canal. Meridional canal s with numerous diverticu- Harbison G.R. 1996. Ctenophora. In "Introducción al
lae, no anastomoses. Color: mature specimens estudio del Zoop1ancton Marino" (R. Gasca, E.
pink, especially along meridional canal s and comb Suarez, eds.), El Colegio de la Frontera Sur (ECO~
rowS. Up to 150 rom in height. SUR)/CONACYT, México, pp. 101-147. General
o Beroe ovata Chamisso and Eysenhardt, 1821 (Fig. review of systematies to fami/y level inclllding
3.17a-c). Body mitten-shaped. Lateral compres- exeellent photographs of the most representative
sion very marked. Four meridional canals of each species.
broad side connected orally by oral forks of para- Mayer A. 1912. Ctenophores of the Atlantic coast of
gastric canal. Eight meridional canals interconnec- North America. Camegie Inst., Washington, 162,
ted by loose network of numerous diverticu1ae, pp. 1-58. Useful deseriptions and illllstratioJls of
with a few anastomoses forming a wide meshwork. North Atlantie etenophores.
Young B. ovata cannot be distinguished from B. Moser F. 1910. Die Ctenophoren der Deutschen
eueumis. Color dull milky, pink or reddish brown. Sudpolar Expedition 1901-1903. Deutsche
Height: up to 115 rom, usually 60-70 rom. Südpolar-Exp. 1901-1903, 11:1-192. First world-
o Beroe /orskaiii Milne Edwards, 1841 (Fig. 3.18). wide distribution review on etenophores.
570 Ctenophora
Order Cydippida
(with tentacles during the entire life)
Tentacles exit near the aboral pole

With aboral keels Without aboral keels
Fam. Mertensiidae Fam. Pleurobrachiidae
1 ~
.~
Mertensia sp. ~ Pleurobrachia pileU5

1 juvenile
3.3b
Tentacles exit near the mouth

Fam. lampeidae
/) )) I
1)
/
3.4
1
Hormiphora plumosa
Lamp~a
3.6 pancenna Tinerfe cyanea
Ctenophora 571
Order lobata
(with two large oral lobes)
Oral lobes arise between the mouth and the statocyst

Adult with tentacular bulbs With tentacles
Fam. Bolinopsidae Fam. Leucotheidae
warts
3.9a
Bolinopsis vitrea Mnemiopsis leidyi Mnemiopsis maccradyi 3.10
Leucothea
multicorn;s
Adult without tentacles bulbs
Fam. Ocyropsidae
3.9c
live juvenile in
"tentacular stage"
live juvenile with oral

Oral lobes arise at the
lobes developed level of the mouth margin
Fam. Eurhamphaeidae
Eurhamphaea
3.ge vexilligera
live adult
3.13
572 Ctenophora
Order Cestida
(ribban-shaped bady)
10 cm
Cestum venerís
(reduced subtentacular comb rows)
3.15a
~
Velamen parallelum
(subtentacular comb rows absent) 3.15b
Order Beroida
(canical bady)
Beroe ovata
(meridional canals 3.18
with anastomoses forming Beroe forskalíí
Beroe cucumís an open meshwork)
(meridional canal s (meridional canal s
without anastomoses) with anastomoses
forming a dense
mesnwork)
Ctenophora 573
Fig.3. Schematic external views ofthe species recorded in the South Atlantic (gastrovascular system is simplified). Seale bars are 1 cm, unless
otherwise indicated.
Figure Sources: from Vanhoffen (1903), redrawn: 1; from Chun (1880), redrawn: 2, 10, 18; from Chun (1898), redrawn: 5; from Mayer (1912),
redrawn: 4, 7,11,13, 14b, 15; from Rang (1828), redrawn: 12; original: 3a, 3b, 6, 8, 9, 14a, 16, 17.
Nemertina
Ray Gibson
Introduction boseis apparatus, the mesopsammic anoplan Riserills

plIgetensis Norenburg, 1993, whieh can only be identi-
The phylum Nemertina (=Nemertea or Rhynchocoela) fied as probably belonging in the anoplan order Hetero-
contains more than 1100 species currently recognised nemertina, or the entocommensal genus Malacobdella,
as valid (Gibson, 1995), although many are inade- whose members are leech-like with a posterior ventral
quately deseribed. The worms are almost exclusively sucker, are at present unknown or subject to discussion,
marine, with only a small number of terrestrial or but the enoplan group Hoplonemertina (Hoplone-
freshwater species known. Nemertines are typically mertea) is divisible into the Monostilifera (proboscis
defined as unsegmented, bilaterally symmetrical, acoe- armed with a single central stylet borne on a eylindrical
lomate invertebrates, although studies by Turbeville basis) and the Polystilifera (proboscis armed with a
and Ruppert (1985) and Turbeville et al. (1992) have shield- or pad-like basis bearing numerous small sty-
suggested that they may be more closely related to pro- lets). The Polystilifera, in turn, eomprises two ecologi-
tostome coelomates than to any aeoelomate group, and cal groups, the benthie species being placed in the
some of the interstitial taxa exhibit pseudosegmenta- Reptantia and the pelagic forms in the Pelagica.
tion (Norenburg, 1988). The principal anatomical
features of nemertines are that they possess a gut with The earliest reference to a pelagic nemertine, although
separate mouth and anus, a closed blood system con- it was reported as a mollusc, was probably the account
sisting of distinct vessels or lacunae, and an eversible of Pterosoma plana Lesson, 1830, found in large num-
muscular proboseis contained, when retracted, in a bers between the Moluccas and Papua New Guinea.
fluid-filled ehamber (the rhynchocoel) extending Moseley (1875a: 382) conc1uded after consulting
above the alimentary canal. They also possess a well Lesson's (1830) original illustration of Pterosoma that
developed nervous system with lobed cerebral ganglia "On the whole ... there seems little doubt that the ani-
and paired longitudinal nerve cords, a ciliated epidermal seen and figured ... was a Nemertine and not a
mis and gut epithelium and, in most taxa, a protoneph- mollusk; but it seems to have been a distinct form, with
ridial excretory system. a pair of eyes and an unbranched digestive tract."
Hubrecht (1887), who reproduced Lesson' s original
The identification, and thus classification, of nemerti- illustrations, concurred with Moseley's eonc1usion,
nes rests largely upon histological studies oftheir ana- but Gibson (1995: 501) was more cautious, stating that
tomy, yet for almost half of the named species their "if this form is a nemertean, as appears probable, its
internal morphology is eompletely undescribed (Riser, taxonomic affiliations rema in unknown." The first cer-
1989). Many systematic changes have been made or tain description of a pelagic nemertine was given by
proposed during recent years, at almost all taxonomic Moseley (1875b) when he reeorded Pelagonemertes
levels from the species upwards, and to a large extent rollestoni Moseley, 1875, from the Indian Ocean. The
both the classification and phylogeny of the phylum internal anatomy ofthis specimen was later partly elu-
rema in open to discussion. At present there is no uni- cidated by Hubrecht (1887), who was the first worker
versally aecepted classification for the group. to both affiliate pelagic nemertines with the
Hoplonemertina and examine them histologically. In
Nemertines can broadly be divided into two major his second paper (Moseley, 1875a), in which he descri-
morphological groups on the basis of their proboscis bed what he believed to be a juvenile specimen of
morphology: those with no stylet apparatus constitu- Pelagonemertes rollestoni, Moseley proposed a new
ting the Anopla, and those in which the proboscis is family, the Pelagonemertidae, to accommodate the
armed with one or more stylets forming the Enopla. species; Bürger (1895) regarded this individual as a
The higher systematic affinities of certain taxa, notably different taxon and renamed it Pelagonemertes lI1ose-
the highly aberrant New Zealand species, Arhynchone- leyi Bürger, 1895, a distinction maintained by sub se-
mertes axi Riser, 1988, which completely laeks a pro- quent authors.
SOllth Atlantic Zooplankton.

/999 Backhll)'s PlIblishers, Leiclen, The Netherlands
576 Nemertina
During the next two decades many other pelagic proboseis

nemertines were described from the Atlantic (Verrill, pore
1892; Joubin, 1904, 1906), Pacific (Woodworth, 1899; brain
Cravens and Heath, 1906) and Indian Oceans (Laidlaw, lobe
1906; Bürger, 1909), Bürger's article containing the
frrst report of a pelagic nemertine from the South
Atlantic, a specimen of Nectonemertes mirabilis lu:
.ir J ')
Verrill, 1892, being obtained in an open net hauled ver- _::'~.:: ..
:JU .'.0.
tically to the surface from a depth of 3000 m. .I!J
lil<
It was, however, a number of papers by August Brink- ,Í/e
lI{
li •.
mann (1912, 1915, 1917a, b, 1920) which led to the ":11.'
E' proboseis
most significant advances in the systematics of pelagic tt (',.:--"
nemertines and, indeed, formed the basis for the classi- Í[l (\1.
llL -.~ Iti

IÁ \ intestinal
fication of the group still in use today. Brinkmann \ll divertieulum
(1917a) (republished verbatim in 1932) was the first to .u--\ In
propose a classification for the pelagic nemertines, alt-

.\1\
"111
( Llj
(~\
hough he noted in his Introduction that the "demand
- rQ
for space altogether transcending the limits assigned to lateral
., :'01~. \
~<., rhynehoeoel
blood
[his] chapter" prevented him from fully describing the
anatomy of the many species established; these were rrí ) vessel
extensively covered in a separate monograph (Brink-
•,'/í
'1 /~ fl n
lateral
mann, 1917b). Brinkmann (l917a) distinguished , t; ., lateral
between the monostiliferous and polystiliferous hoplo- nerve eord
nemertines as separate suborders; he also divided the mid-dorsal
Polystilifera into two tribes, which he called the blood
vessel
Reptantia (bottom dwelling Polystilifera with cerebral
caudal
sensory organs, an excretory system and lateral rhyn- fin
chocoel diverticula) and Pelagica (pelagic Polystilifera
with no cerebral organs, excretory system or rhyncho- Fig. I. Mature male Nectonemertes mirabilis in dorsal view,
coel diverticula). In this paper he established the seven drawn to emphasise morphological features which may be seen eit-
families Arrnaueriidae, Bathynemertidae, Buergerielli- her in stained whole mount or transparent live bathypelagic nemer-
tines. Redrawn from Coe (1926).
dae, Chuniellidae, Dinonemertidae, Phallonemertidae
and Planktonemertidae, emended the definitions of the
Nectonemertidae and Pelagonemertidae, and described
nine new genera (Armalleria, Bathynemertes, Bllerge- cal studies made by Brinkmann enabled him to dis-
riella, Chuniella, Crassonemertes, Natonemertes, cuss the comparative anatomy of the pelagic nemerti-
Pendonemertes, Phallonemertes, Plotonemertes) and nes, particularly their epidermis, body wall and other
eleven new species. musculature, parenchyma, alimentary tract, sensory
organs, proboscis apparatus and blood, nervous and
Brinkmann' s (1917b) monograph, using the same clas- reproductive systems; anatomical features which can
sification as that given in his 1917a paper, was the first be used to distinguish between pelagic species are
major treatise on pelagic nemertines, most of the 37 illustrated in Fig. I and 2.
species he recognised being both described and illu-
strated in detai1. He defined the Pelagica (Brinkmann, Brinkmann' s (191 7b) discovery of a small crustacean
1917b: 145) "Pelagische Polystilifera. Cerebralorgane, in the intestine of a Nectonemertes mirabilis was the
Exkretionsorgane, Rhynchocoelomdivertikel und first evidence on the food of bathypelagic nemertines
metamere Gefasskommissuren fehlen. d Gonaden nur and, though few data exist on their diets, it is general-
in der Kopfregion entwickelt." The detailed histologi- Iy believed that Crustacea and other pelagic inverte-
~.,~~
Nemertina 577
~~ ..•
~•.•.
, rhynchocoel
:.--'>'.''''''~~o::'0,~ muscu lature
~ ~~\
'(.~/b:1
~D ~-fnl\
~...:~·~ZiV ~JiJ¡ rhynchocoel
~:ffy
~~'~-:-?¿y
Planktonemertidae
~~ circular
Proa rma ueria :.:.. . E~ muscle
Dinonemertes Plionemertes '. . "..~)\ laye~.

E ,::::J\& longltudlnal
::::'~'ll muscle
..:'::/: VIi; layer
::;,::;~
~--
Pelagonemertidae
lateral
blood vessel
intestinal
rhynchocoel Nectonemertes Pelagonemertes
Fr;¡;;;
Pelagonemertes joubini
diverticulum pelagica brinkmanni
Cuneonemertes gracilis
:: : ~ ,,,,,,1
.
" ..,:~}~ \ nerve
spermary
Phallonemertes murrayi
Fig. 2. Analomical fealures of balhypelagic nemertines. A-C: diagrams lO show variations in rhynchocoel length in representative genera; D-
E: diagrams to show differences in the organisation of the rhynchocoel musculature, comprising interwoven circular, longitudinal and spiral
libres (D), and primarily with outer circular and inner longitudinal muscle layers but with a few spiral libres running among the circular mus-
cles (E); F-I, transverse sections through part of the intestinal regions drawn schematically to show differences in the arrangement of the later-
al intestinal diverticula relative to the position of the lateral nerves; ]-K: the organisation of the blood system, with mid-dorsal blood vessel
extending the full body length (1), and with a short, rudimentary mid-dorsal blood vessel (K); L, ventral view of the head of a mature specimen
showing the slender muscular penes which extend through the body surfaee from the spermaries. All redrawn from Coe, 1926.
brates probably constitute the food of these worms; sual features ofthe nervous system, zoogeographic dis-
nemertines as a phylum are typically carnivorous tribution and the means whereby these animals are dis-
(McDermott and Roe, 1985). persed in the oeeans. Coe's (1926) monograph was the
second major treatise on pelagic nemertines to be
It was in 1920 that the first of Coe's many articles published. In this work Coe followed the familial
dealing with pelagic nemertines appeared. During the arrangement proposed by Brinkmann (1917a, b) but
next 36 years Coe added much to the knowledge of established several new genera as well as eight new
bathypelagic nemertines, his various articles covering species. The introduetion to this extensive work inclu-
not just descriptions of many new taxa and the syste- des a history of the subject, with reference to the
matics of the group but also the adaptations shown by various expeditions during whieh pelagie nemertines
pelagic nemertines to their way of ¡ife, details of unu- had been colleeted, and is followed by a detailed eOffi-
578 Nemertina
parative discussion ofwhat he called the "morphologi- serve as spermatophores by being retained within the
cal peculiarities", commenting (p. 18) that "The special female ovaries after mating.
adaptations of these worms to a pelagic existence far
beneath the surface of the ocean have led to numerous Three of Coe's publications (Coe, 1935, 1936, 1945)
modifications of the organ-systems as found in their were principally concerned with collections of bathy-
littoral relatives. Most ofthese modifications ... are due pelagic nemertines made within the Bermuda region.
to the loss or reduction of structures which are found in His 1935 paper included a discussion of the distribu-
a more highly organized condition in the tion of pelagic species in relation to water temperature,
Drepanophorus-like forms from which the pelagic density, salinity, oxygen levels, phosphate content and
nemertines are thought to have been descended." It was pH with the view of showing how stable conditions
in the section summarising the ecology of pelagic permit a great vertical range and almost unlimited
nemertines that Coe (p. 91) commented that at the geographical dispersal of species below 1200 m depth.
depths the nemertines live they "swim slowly to and In the last of these three articles on the pelagic nemer-
fro or float idly with a very slight muscular effort", alt- tines of Bermuda Coe (1945) extended his discussion
hough later (p. 93) he refers to the violent undulatory ofthe means whereby these animal s were distributed to
swimming movements earlier noted for Pelagone- include south Atlantic species, later publishing this dis-
mertes and Nectonemertes by Moseley and Brinkmann cussion as a separate article (Coe, 1946). He explained
respectively. This supposed feeble locomotion has the dispersal of pelagic organisms in terms both of
been widely restated in subsequent literature, but video superficial and deep currents. During this period
film of several north Pacific species shown at the 1995 Wheeler (1934, 1937) described new species from the
international nernertine conference in Monterey, South Atlantic and Indian Oceans, later (Wheeler,
California, by Drs. Pam Roe and Jon Norenburg 1940) reporting new records of pelagic nemertines
demonstrated that many of the pelagic species can from Antarctic waters.
swim actively at least for short periods of time.
By 1954 57 species of pelagic nemertines had been
Coe (1926) also commented on the great variation in described worldwide, of which 12 were known from
size between species of bathypelagic nemertines, the Pacifico Coe (1954a) added a further 17 species to
noting that whilst some species of, for example, the known Pacific fauna, seven of which were pre-
Balaenanemertes, attained sexual maturity at lengths of viously known only from the Atlantic (Coe, 1945).
only 6-10 mm, other forms were much larger. He cited Coe's (l954a) classification of the Pelagica listed 11
one specimen of Dinonemertes investigatoris Laidlaw, families, with the Balaenanemertidae and Pachy-
1906, which was 203 mm long, 56 mm wide and 15 nemertidae being the only two not included in
mm thick, although other species of Dinonemertes and Brinkmann's (l917a, b) publications. Coe established
Nectonemertes may atta in lengths of 60-85 mm. The four new genera (Chunial7na, Plenanemertes, Tono-
largest South Atlantic form yet discovered is 11 emertes, Tubonemertes) and ten new species for the
Calonemertes hardyi (Wheeler, 1934). Pacific, providing keys to both families and genera.
Many of the taxa he recorded in this paper carne from
Although virtually nothing is known about the repro- collections made in the submarine canyon off
ductive biology of pelagic nemertines, Coe (1926) Monterey Bay, California, and Coe's (l954b) article
extensively discussed the structure ofthe gonads, sper- was principally concerned with this region. His last
matogenesis and ovogenesis in the Pelagica. Except for paper on pelagic nemertines (Coe, 1956) gave keys to
Proarmaueria pellucida Coe, 1926, which appears to the families and genera of pelagic nemertines known
be irregularly hermaphroditic, bathypelagic nemertines to occur in the eastern and central north Atlantic
are dioecious, with both the position and morphology Ocean. Apart from Brinkmann and Coe, only one
of their gonads modified in comparison to those found other zoologist, the Russian Korotkevich, has made
in benthic nemertines. In one pelagic form, Phallone- major contributions to our present knowledge of
mertes murrayi (Brinkmann, 1912), the spermaries are bathypelagic nemertines. Korotkevich (l955a) listed a
prolonged to form slender, muscular penes (Fig. 2L), total of 73 species of pelagic nemertines from Russian
regarded as true copulatory organs by Coe, which may seas, 16 of which were described as new. Korotke-
Nemertina 579
vich's classification of the group, however, was very System (MOCNESS) (Wiebe et al., 1985) provide the
different from that given by Brinkmann and Coe in most accurate figures for depth and other hydrographic
that she recognised only three families (Armaueriidae, data for pelagic organisms as well as improving the
Nectonemertidae, Pelagonemertidae) and seven gene- condition ofthe specimens obtained. The small size of
ra (Armaueria, Dinonemertes, Mesarmaueria, Necto- many pelagic nemertines, which particularly as juveni-
nemertes, Pelagonemertes, Planktonemertes, Proar- les may be les s than 10 rnm long, indicates that an
malleria). Other articles by Korotkevich (1955b, appropriately sized mesh should be employed. P.
1956,1960,1962,1963,1964,1967) dealt with diver- Schalk (jide Van der Spoel, 1988) used a combined
se topics relating to pelagic nemertines, including lists rectangular mid-water trawl RMT 1 + 8 with two nets,
of far eastern and Antarctic species. a discussion of one with a mesh of 0.32 rnm and the other of 4.5 rnm,
the polyphyletic origin of the group, and the sugge- at depths of 0-500 m and obtained large numbers of
stion that the Middle Cambrian Burgess Shale fossil meso- and bathypelagic species from several taxono-
AmisÁwia sagittiformis Walcott, 1911, originally mic groups, including nemertines. Recently Roe and
described as a chaetognath, was a pelagic nemertine; Norenburg, in a paper presented at the Fourth
she affiliated it with the family Pelagonemertidae, but International Conference on Nemertine Biology in
Conway Morris (1977) fully redescribed this fossil 1995, demonstrated that the use of a slow net towing
and concluded that its phyletic position could not be speed (of less than about 2 knots) was essential for the
determined. recovery of living pelagic nemertines in excellent con-
dition.
In her most recent discussion on the systematics of the
Pelagica Korotkevich (1977a) recognised only three As with benthic nemertines, the identification of pel-
families and eight genera; these families, with their agic species depends ultimately upon a study of their
contained genera listed in parentheses, were the internal anatomy by means of histological sections,
Armaueriidae (Armalleria, Mesanl7aueria, Proarma- although specimens should be carefully examined
ueria), the Nectonemertidae (Dinonemertes, Necto- alive prior to preservation. Certain external features,
nemeftes, Planktonemertes) and the Pelagonemertidae such as the tentacles of male Nectonemertes (Fig. 1),
(Obnemertes, Pelagonemertes). are oftaxonomic significance whilst particular intern-
al structures, such as the gut diverticula, gonads or
Comparatively few new taxa of pelagic nemertines nervous system, can sometimes be distinguished in
have been described since about the middle 1960's; whole mount specimens (Fig. 1) stained with borax or
Friedrich (1969b) and Van der Spoel (1988) described alum carmine and then cleared (Coe, 1954a; Van der
new species from the Gulf of Guinea and Indonesia Spoel, 1985); smaller sized and transparent worms
respectively, whilst Chernyshev (1992a) used suppo- can often be studied as whole mounts without the
sed cases of homonymy, unavailability of names and need for staining, or other appropriate stains, such as
/lomina nuda to propose new names for a number of Mayer's haemalum, can be employed.
nemertine taxa, including some pelagic forms, later
(Chernyshev, 1992b) reclassifying the family Histological studies should be made on carefully fixed
Armaueriidae with the suggestion that this group was specimens embedded in a suitable wax; harder waxes
derived from primitive Nectonemertidae. Cherny- (e.g., 56°C m.p. paraffin wax) are generally more sui-
shev's proposals have not met wide acceptance. table than lower melting point media, and serial sec-
tions of 5-8 /lm are ideal. The choice of fixative is to
some extent governed by which staining protocol will
Methods be followed; some authors have used formalin or alco-
hol fixation, with the sections subsequently stained in
Most of the earlier data on the vertical distribution of haematoxylinleosin, but Bouin's solution, formol-cal-
bathypelagic nemertines, based on the use of non-clo- cium or Susa's fluid followed by staining with a tri-
sing nets, were inaccurate, whereas closing nets such chrome method (e.g., Mallory or Masson) work well
as modified Tucker trawls with thermally protecting with benthic nemertines and normally yield a better
cod ends (Childress et al., 1978) or a Multiple differentiation between body organs and tissues; these
Opening/Closing Net and Environmental Sensing techniques can equally be applied to pelagic species.
580 Nemertina
600 40° 20° 0° 20°
0°
0°
·_1
•
T::!
20° I ~ 10°
T::!
T+
10°
20°
. ~.
. 30°
30°
~
•• ~
40°
40°
•
• 50°
50°
\l Calonemertes hardyi
• Nectonemertes minima
• Nectonemertes mirabilis
60° 40° T Nectonemertes primitiva
• Pelagonemertes joubini
+ Pelagonemertes korotkevitschae
T::! Pelagonemertes moseleyi
• Pelagonemertes rollestoni
t0. Planktonemertes vanhoeffeni
O Probalaenanemertes irenae
O Protopelagonemertes hubrechti
Fig. 3. Locations in lhe South Atlantic from where pelagic nemertines have been collected.
Geographic and vertical distribution 10cations such as the submarine canyon occurring off
Monterey Bay, California (Coe, 1954a) or close to
Geographic patterns Bermuda (Coe, 1935, 1936).
The 97 species of bathypelagic nemertines at present The North Atlantic, with representatives from 26 out of
recognised (Gibson, 1995) have mostly been found in the 40 genera (65%), has been investigated more fre-
off-shore deeper oceanic waters; none has yet been quently than any oceanic region; 30% of the known
found in shallow bays or in semi-landlocked seas such species are also known only from the North Atlantic
as the Mediterranean or North Sea, although several (Table 1), largely due to the publications of Joubin
individuals have been recovered fram nearer-shore (1906), Brinkmann (1917a, b), Coe (1935, 1936, 1945)
Nemertina 581
«c:
-zQ..~.:
'ü
< ~
<.:
.:c:
'ti c:
<.J
:t~ :t
'ü vi vi
;i .~
Q..
.~ ~ ~
<.J<.J
•..
<.J
••---• •~ • •
~-- • • +~._-~
•• • •
~l·
954
kmann,
oe,
ata
ae,
er,
15
1906)
oubin,
4n,
92
1945
nn,
906)
917
5
1954 1917
n,1964
1906)
1926)
1906)
1937
Brinkmann,
aidlaw,
1936
ann,
dii
1906
~ich, 1917
1934)Brinkmann,
1906)
1917
1917
1906
Korotkevich,
Bürger, (Joubin,J206)
1977
1909
1917
1917
1955
~ --r---
~andronemertes
Mononemertes
ductor Chernyshev, 1992
Balaenanemertessargassicola (Joubin, 1906)
grandis Brinkmann, 1917 .t·
~-+t~
·1¡
k,,',h, I964)
582 Nemertina
...
.<: tí'"
:o ~ e i
:('"...
e... ~·0
¡¡::
c:
¡¡::
...
Q.
~i..!: .~ :(
·'"0vi Q.
,Nemertobus maximovi (Korotkevich, 1960)

Neoarmaueria angusta (Korotkevich, 1955)
Neoarmaueria crassa (Korotkevich, 1955)
Neoarmaueria
Neoarmaueria
Neoarmaueria
divaricata (Korotkevich,
laticeps (Korotkevich, 1955)
tenuicauda (Korotkevich, 1955)
Neuronemertes aurantiaca
Obnemertes latilobata
Coe, 1926
Korotkevich,
1955)
1960
Ilr-
••
• I :
Obnemertes ramosa Korotkevich, 1960

Obnemertes sol ida Korotkevich, 1964 •
•
,Pachynemertes obesa Coe, 1936
Parabalaenanemertes fusca Brinkmann, 1917 •
Parabalaenanemertes zonata (joubin, 1906) •
•
Paradinonemertes drygalskii Brinkmann, 1915
•
Paradinonemertes macrostomum Coe, 1954
•
Pelagonemertes brinkmanni Coe, 1926
•
· :, ..
Pelagonemertes excisa Korotkevich, 1955
Pelagonemertes joubini Coe, 1926
Pelagonemertes korotkevitschae Friedrich, 1969
Pelagonemertes laticauda Korotkevich,_1_9_5_5 _
·1· ·
±l~
Pelagonemertes moseleyi Bürger, 1895
Pelagonemertes oviporus Korotkevich, 1955
· .:.;':
Pelagonemertes parvula Korotkevich, 1964 ¡-
Pelagonemertes robusta Korotkevich, 1955
Pelagonemertes rollestoni Moseley, 1875
. •
Pendonemertes levinseni Brinkmann, 19_1_7 _
•
Phallonemertes murrayi (Brinkmann, 1912)
•
Planktonemertes agassizii Woodworth, 1899
•
Planktonemertes curvice~a
Planktonemertes vanhoeffeni
Korotkevich, 1964
Brinkmann, 1915
----+---
• • • •
Planonemertes labiata Coe, 1936 •

Planonemertes lobata Coe, 1926
Plenanemertes rubella Coe, 1954
Plionemertes constricta Coe, 1954
Plionemerte~plana Coe, 1926
• •
Plotonemertes adhaerens Brinkmann, 1917
Plotonemertes aurantiaca
Proarmaueria korotkevitschae
Coe, 1936
Proarmaueria pellucida Coe, 1926

Cherny;;hev, 1992
Proarmaueriella caudata (Korotkevich, 1955)

~~
F:
• 1: •:
.••
•
Probalaenanemertes irenae Wheeler, 1~
•
Probalaenanemertes wijnhoffae Brinkmann, 1917
• •
Protopelagonemertes beebei Coe, 1936
• •
•
t-•• •
Protopelagonemertes hubrechti (Brinkrnann, 1917)
Protopelagonemertes jocula tori Van der Spoel, 1988
Tononemertes pellucida Coe, 1954
Tubonemertes aureola Coe, 1954
Tubonemertes wheeleri (Coe, 1936) •
Xenoarmaueria acoeca (Korotkevich, 1955)
f-
•
Zinarmaueria platonovae Chernyshev, 1992 •
Nemertina 583
and Van der Spoel (1985). North Pacific records, prin- cies have now been collected; Brinkmann (1917a), for
cipally by Coe (1926, 1954a) and Korotkevich (1955a), example, reported a total of 117 individuals collected
indicate that although the number of genera is less than by expeditions in the North Atlantic, Coe (1945) lis-
for the North Atlantic, with only 21 (52.5%) represented 44 specimens from the Bermuda area and Van der
ted, the number of species indigenous to the North Spoel (1985) found 248 examples in collections made
Pacific is identical at 30% (Table 1). Collections in the by the AMNAPE cruises.
South Pacific and Indian Ocean have yielded 25% and
22.5% ofthe genera respectively, with similar numbers
of indigenous species (9 from the South Pacific, 8 from Vertical profiles and hydrographic data
the Indian Ocean) (Table 1) known only from one or
the other region. Many of the bathypelagic nemertines There are few accurate data on the depth distribution
reported from these two oceans are from Antarctic lati- of bathypelagic nemertines, principally because so
tudes (Brinkmann, 1920; Wheeler, 1934, 1940; many of the early records were made from specimens
Korotkevich, 1960, 1963, 1964; O'Sullivan, 1983). collected in nets which were hauled open to the sur-
Representatives of only 6 genera (15%) and 11 species face. Brinkmann (1917a) concluded that Nectone-
have so far been recovered from the South Atlantic mertes mirabilis lived in a fairly well defined layer of
(Fig. 3) (Wheeler, 1934; Coe, 1946; Friedrich, 1969b), water where the temperature was 6°C or less and the
whilst only a single species and genus is known from salinity did not exceed 35%0, this layer occurring at
the Arctic Ocean (Korotkevich, 1977b) (Table 1). depths of 500-2000 m between Newfoundland and
Ireland in the North Atlantic where this species was
By far the greater number of bathypelagic species collected (Fig. 4).
(80%) are known from only one oceanic area (Table
1). Coe (1945, 1946, 1954a, b) argued that, as a con- Collections ofpelagic nemertines made between 1980-
sequence of the circulation of deep oceanic currents, 1983 by the Amsterdam Mid North Atlantic Plankton
pelagic nemertines may over a long period oftime be Expeditions (AMNAPE), however, both yielded
expected to drift from one part ofthe world to another hydrographic data and enabled Van der Spoel (1985)
and thus possess wide zoogeographic distributions. to further analyse the vertical distribution of
The recorded data on these animals does not, howe- Nectonemertes mirabilis in detai1. He found that
ver, support Coe's hypothesis; very few species are Nectonemertes mirabilis at latitudes north of 41°N
known from more than a single oceanic region (Table lives at rather shallow depths of about 300 m during
1) and of the 6 (6.2%) the most widespread form the winter, whereas at other times of the year it des-
appears to be Pelagonemertes rollestoni. Van der cends to 500 m or deeper. At depths of less than
Spoel (1983) believes that there are comparatively approximately 1750 m the species reaches further
few planktonic taxa with a widespread zoogeographic south in spring than in summer, whereas its northem-
distribution and other authors, such as Van der Spoel most abundance is highest during the auturnn. Near
and Heyman (1983), have shown that significant pro- 55°N its uppermost distribution appears to follow the
portions of the bathypelagic fauna from one oceanic 9°C isotherm. Shallow (approximately 25-100 m
area or another relate to endemic taxa. With so many depth) summer night samples may illustrate vertical
of the pelagic nemertines known from only single or, migration. Van der Spoel (1985: 21) concludes that
at best, a small number of specimens, identifications pelagic nemertines "as a group, are bathypelagic ani-
of conspecific forms from widely separated geograp- mals with an upper limit of distribution between 500
hic regions, such as Dinonemertes investigatoris from and 1000 m depth." Depth data for species recovered
the North Atlantic and Indian Ocean, must be open to during the AMNAPE cruises are summarised in Fig. 5.
uncertainty. Among all the reported bathypelagic
nemertines one species, Nectonemertes mirabilis, Data on the nemertines found in the South Atlantic are
appears to be by far the most common and abundant sparse; they are accordingly listed with the species
(Coe, 1954a). More than 400 specimens of this spe- descriptions .
•••
Table 1. The recorded world-wide distribllti~n of bathypelagic nemertine genera and species.
-
Q ~ ::1
-""l
e> _.
VI
00
~
0..(JQ
C1l :::eJ;Cl
=:: ~. ~ Depth ranges (m) ~.
o f..
"'lE
t~~ (Xl
o V1
o oo V1
o ~::r::
~'<
~ g g.
o o o o o Sal."/" Ternp.·C lE
e> o
E¡-
Depth (m)
N
O> O>
3 'O
- ;:¡ q:¡
e>
o..a d. Armaueria rubra 35.59 10.25 (OÍ

3 -
oo O>
'O
::- oo
o oo
Vl oo
o oo
Vl
o
· .
~2l ~ ñ'
.
~ ~ Calonemertes hardyi ~ 35.73 9.15 '"O
(!)
~ e>'"
e>
....•
§ CY l:lJ ~ g.
<:>.. •..•
Chuniella sp. 35.10 6.90 e

•....• e> o
¡¡;-",
g ~
!l: Z
:::o Dinonemertes investigatoris 35.55 6.15
....•
(!)
:::r. cr'
<
~ ~
e>
/
1::::::::1:::::::¡:::::::::::::¡:::::::::::::::¡:¡:¡:::::::::::¡¡¡::rrr:::1:¡:¡:¡:¡:¡:¡:::::
~
:! ~ - "...
D •
...
~. g .•.• .... ""
~ > ~I\ Dinonemertes mollis ~ Z

~V1 35.12-14.35-
35.55 8.20 ::-e>
e:: '"
fr
~'V'l
l:lJ
::l e> lE
3 8'
•
-
Ill-o
~ g. <Ill o... '" e
0"0" 0..-.
Illn
Nectonemertes minima 35.12 4.35 V> ~'"
0..0..
'< ~
n~ 3 l:lJ O>
<n
_
::-",
O>
::-'0
e> e>
III
::J ::l e> o..
,~
CJ> 0..0>
• • •,•,, ¡i,"•
:3 ¡¡;- •....• =-::3
'< II
'" (JQ_.
::-0
"'<
, ..• ,,,, /
.•.•
, ~I'"
e>
o..
I I 1fII 00
e>
< '"
e>
~ O' Nectonemertes mirabilis ;--- ¡¡;-
O> 5o ~ /"I IIaIIII.
o..
''"@
"
.Vi
lE (5 /
I o \ I,\\IIIf
()I"
............
I
f
Z ~~
~
S'
~3 Ill, 34.96-13.45-
35.55 9.70 5. ~,. S
::+
"':l
\~
- e>
'" _.
-
'"
IV", ~.~ n g.
~e> <CJ>
Ill-o O :o-
• <n
::l s'
-oii
" o
Q..Ill
n o... e> ~
~ (D'
<:>..Y'
3 =:!'. '"
'(;;'e>
So
,,' O"
III
::J
O ~o..
e'" CJ> Nectonemertes primitiva ::l _. e>
'"
O>
'"
e>
34.96-16.70-
35.70 11.90 V> 0..'0
=:~
::: o.. '" '"
~.§
- o
~ ~
~ ~v """-0
'"
~~
~.V1
Paradinonemertes wheeleri 35.03 3.45 s:: 3
~ g-
:::-~ o lE
0..::-
1llCJ>
4.35-
'"
~(JQ
'"~ _.
<
O>
<-o
Illlll
o.. n Pelagonemertes joubini
35.12-
35.14 9.35 ~&
o.. '"
...:.
lE
('il
::. 3 35.41 8.15

~]
o o
~ '" III Pelagonemertes moseleyi 3 -
....<
e> O>
::J
CJ> ttl g
'"
3
e>
0..C/J
'"
g-
•..•
Pendonemertes levinseni 35.10-16.15-

35.87 10.45
~i
§ ~
'"Q
~g ~c
§;:: ~ \O~
~ \O ::i::
::: 00 O" !l:
'"
..., ~
V>
PhaJlonemertes murrayi 35.12 I 4.35 ~~'" illl¡:II¡III!I¡I~II:IIII~II¡:¡¡I~:·¡:'¡I[¡¡::¡~::"::::::::·.:::·:·:::::~::::¡~¡::::::::::::r::::::: ..
~b Planktonemertes vanhoeffeni 10.20 ==
~
'"
~.
'"
~.
"'"
'" '"
d. 4.45
::.
~::
cr-rt'"
'<
~
Plotonemertes adhaerens
s~
;-
Nemertina 585
Classification Family Balaenanemertidae Coe, 1954

Genus Balaellallelllertes Bürger, 1909
The systematics of pelagic nemertines have still not Family Buergeriellidae Brinkmann, 1917
been satisfactorily resolved, with the two conflicting Genus BlIergeríella Brinkmann, 1917
elassifications (Coe's system, based upon Brinkmann's Family Chuniellidae Brinkmann, 1917
pioneering work, and Korotkevich's system) differing Genera C/lllllíalllla Coe, 1954; Chllllíella
widely in the number of families and genera that they Brinkmann, 1917
recognise. One ofthe major difficulties with this group Family Dinonemertidae Brinkmann, 1917
of nemertines is that for so many of the taxa only indi- Genera Alexandrollelllertes Chernyshev,
vidual, or at best a very small number, of specimens 1992; Díllollelllertes Laidlaw, 1906; Para-
has been found; this problem is compounded by the dillollelllertes Brinkmann, 1915; Plallolle-
lack of anatomical data for many of them, especially lIlertes Coe, 1926; Pliollelllertes Coe, 1926;
the older established species, and insufficient knowled- TlIbollemertes Coe, 1954
ge of either intraspecific variation or possible sexual Family Nectonemertidae Verrill, 1892
dimorphism. Recent collections of pelagic nemertines emend. Brinkmann, 1917a
from the north Pacific have yielded many specimens, Genus Nectollelllertes Verrill, 1892
several of which appear to be new (Drs. Pam Roe and Family Pachynemertidae Coe, 1954
Jon Norenburg, pers. cornm.), and it is to be hoped that Genus PachYllelllertes Coe, 1936
with the possibility of molecular techniques being Family Pelagonemertidae Moseley, 1875b
applied to systematic studies on these pelagic animals emend. Brinkmann, 1917a
that future studies will resolve the conflicting elassifi- Genera ClIlleollelllertes Coe, 1926; Gelalle-
cations of Coe and Korotkevich. The elassification lIlertes Coe, 1926; Lorallelllertes Cherny-
used in the present chapter follows the Brinkmann/Coe shev, 1992; NallllOllelllertes Wheeler, 1937;
system, but Korotkevich's elassification is also shown Natollelllertes Brinkmann, 1917; Nelller-
to emphasise the differences between these two sche- toblls Chernyshev, 1992; Obllelllertes
mes. Gibson (1995) lists 40 genera and 97 species of Korotkevich, 1960; Parabalaellallemertes
pelagic nemertines as currently valid, of which 11 spe- Brinkmann, 1917; Pelagollelllertes Mose-
cies have now been recorded from the South Atlantic. ley, 1875; Probalaellallelllertes Brinkmann,
1917
The following elassification is based upon Coe's Farnily Phallonemertidae Brinkmann, 1917
(1954a) work, but with the higher taxonomic categories Genus Phallollelllertes Brinkmann, 1917
named after Sundberg's (1991) revised terminology for Family Planktonemertidae Brinkmann,
the phylum. Families shown in bold underlined print 1917
inelude genera and species which have either been Genera Crassollelllertes Brinkmann, 1917;
recorded from the south Atlantic or are likely to occur Mergollelllertes Brinkmann, 1917;
within the region; these taxa are covered in detail in the Monollelllertes Coe, 1926; NellrOllelllertes
present work. Coe, 1926; Plallktollelllertes Woodworth,
1899; Plellallelllertes Coe, 1954;
Phylum Nemertina Schultze, 1853 TOIlOllelllertes Coe, 1954
Class Enopla Schultze, 1853 Family Protopelagonemertidae Brinkmann,
Subelass Hoplonemertina Hubrecht, 1879 1917
Superorder Polystilifera Brinkmann, 1917 Genera Calollelllertes Coe, 1945; PelldOlle-
Order Pelagica Brinkmann, 1917 lIlertes Brinkmann, 1917; Plotollelllertes
Family Armaueriidae Brinkmann, 1917 Brinkmann, 1917; Protopelagollelllertes
Genera Arlllalleria Brinkmann, 1917; Neo- Brinkmann, 1917
arlllalleria Chernyshev, 1992; Proarlllalle-
ría Coe, 1926; Proarlllalleriella Cherny- Korotkevich's (1977a) classification, which is not
shev, 1992; Xenoarlllalleria Chernyshev, used in the present chapter, recognised the following
1992; Zillarlllallería Chernyshev, 1992 families and genera:
586 Nemertina
Family Armaueriidae. Armaueria. Mesarmaueria Bathypelagic nemertines have been relatively little
(Chernyshev, 1992b, synonymised this genus with investigated from the South Atlantic in comparison
Proarmaueria), Proarmaueria. with other oceanic regions; for example, representati-
Family Nectonemertidae (Korotkevich, 1955a, inclu- ves of all 11 families have been recorded from the
ded in this family the Buergeriellidae, Churuellidae, North Atlantic, with a total of 44 known species from
Phallonemertidae, Planktonemertidae and Proto- 26 genera (45% ofthe world's known species), where-
pelagonemertidae). Dinonemertes (Dinonemertes, as in the South Atlantic the 11 reported species are
Planonemertes and Plionemertes were synonymi- from 6 genera and only 4 families. No examples from
sed with Dinonemertes by Korotkevich, 1955a), the families Armaueriidae, Balaenanemertidae,
Nectonemertes (Korotkevich, 1955a, included Buergeriellidae, Chuniellidae, Dinonemertidae, Pachy-
Buergeriella. Chuniella (partim) and Drepano- nemertidae or Phallonemertidae have yet been found in
phonls (partim) as synonyms of Nectonemertes), the South Atlantic.
Planf..:tonemertes (Korotkevich, 1955a, included the
following genera as synonymous with Planktone- In the section which follows a key is provided to the
mertes: Calonemertes. Chuniella (partim), Crasso- families, genera and species of pelagic nemertines
nemertes, Mergonemertes. Mononemertes, Neuro- recorded from the South Atlantic and brief diagnoses
nemertes. Pachynemertes, Paradinonemertes, for each taxon are given. Other species will undoub-
Pendonemertes, Phallonemertes, Plotonemertes, tedly be discovered during future oceanographic sur-
Protopelagonemertes). veys. Where the name of the species in current use
Family Pelagonemertidae (The family Balaenane- differs from that recognised by Korotkevich (l955a,
mertidae was included in this taxon by Korotke- 1977a) the alternative name is indicated.
vich, 1955a). Obnemertes, Pelagonemertes
(Korotkevich, 1955a, included as synonymous
with Pelagonemertes the following genera: Key to the South Atlantic families, genera and spe-
Balaenanemertes, Cuneonemertes, Gelanemertes, cies of bathypelagic nemertines
Nannonemertes, Natonemertes, Parabalaenane-
mertes, Probalaenanemertes). Mid-dorsal blood vesse1 extends to posterior end
of body 2
la Mid-dorsal blood vessel ends b1indly in rhyncho-
Identification of pelagic nemertines coel close behind brain (Pelagonemertidae) .4
2 With distinct and posteriorly bilobed caudal fin
Although histological studies are ideally required for clearly demarcated from body: (Nectonemertidae;
the certain identification of pelagic nemertines, indi- this fami1y contains on1y the single genus,
viduals can often be placed in families on the basis of Nectonemertes) 8
either external features or internal structures visible in 2a Without distinctly demarcated caudal fin, alt-
whole mount specimens, especially when stained and hough posterior end of body may be f1attened
cleared. Coe (1954a, 1956) provided keys to the (two genera of Planktonemertidae not yet recor-
Pacific and North Atlantic families respectively in ded from the South At1antic do possess an
which he distinguished between them by such featu- obvious caudal fin) 3
res as the presence or absence of anterior tentacles, the 3 Body slender, narrowed at both ends, not f1atte-
length ofthe mid-dorsal blood vesse1, the number and ned (Protopelagonemertidae) 10
arrangement of the intestinal diverticula and whether 3a Body broad and very f1at: Planktonemertidae.
or not they possess distinct ventral branches, the Only one genus, Planktonemertes, and one spe-
shape of the body (i.e., broad and f1attened or slender cies Pla"kto"emertes vallhoejJe,,¡ (Fig. 6.9), has
and rounded), and the development of external phalli so far been recorded from the South Atlantic
on the male gonads (spermaries). Anatomical features 4 Body with well deve10ped caudal fin, intestinal
of pelagic nemertines which can be used in their iden- diverticula separated by extensive amounts of
tification are illustrated in Fig. 1 and 2. gelatinous parenchyma (Pelagonemertes) 5
Nemertina 587
4a Body with well developed caudal fin, intestinal Diagnoses of the families, genera and species
diverticula closely appressed and not widely
separated by gelatinous parenchyma: Proba/ae- Family Nectonernertidae
nanemerfes; only one speeies so far recorded Body rather slender, rnoderately flattened; posterior
from the South Atlantic, Proba/aenanemertes end with well dev.eloped caudal fin, often posteriorly
irenae (Fig. 6.8) bilobed; rhynchocoel nearly as long as body; adult
5 With 2-8 pairs of long, widely separated intestin- males with well-developed tentacles, spermaries
al diverticula 6 numerous, in two groups beside and behind brain;
5a With more than 12 pairs of long but fairly close- intestinal diverticula without ventral branch.
Iy distributed intestinal diverticula 7 Genus Nectonemertes
6 With 4-5 pairs of distinct intestinal diverticula, and Body rather slender, moderately or very flattened,
group of short intestinal lobes posteriorly; posteri- with distinct and posteriorly bilobed caudal fin; rhyn-
or of body tapering to end in well differentiated chocoel nearly as long as body; intestinal diverticula
caudal fin: Pelagonemertes mose/eyi (Fig. 6.6) without ventral branches; mid-dorsal blood vessel
6a With 7-8 pairs of distinct intestinal diverticula, extends full length of body; anterior tentacles present
and 3-4 short intestinallobes posteriorly; posteri- in sexually mature males; spermaries in two groups
or of body tapering only slightly to end in blunt alongside and posterior to brain on each side ofbody.
tail without caudal fin: Pelagonemertes korotke- • Nectollemertes minima Brinkmann, 1915 (Fig.
vitschae (Fig. 6.5) 6.1). Body moderately slender, posteriorly with
7 Proboscis with 30 nerves: Pe/agonemertes jOllbi- distinct caudal fin, mature males with pair of large
ni (Fig. 6.4) tentacles up to about 4 rnm long on the head; deve-
7a Proboscis with 16-22 probo seis nerves: Pelago- lopment of tentacles correlates with sexual matu-
nemertes rol/estoni (Fig. 6.7) ration of the spermaries, with immature males
8 Rhynchocoel musculature in front of brain with showing only rudimentary tentacle formation;
inner circular and outer longitudinallayers but this lengths of 11-19 rnm, widths of < 4 rnm, maxi-
sequence is reversed post-cerebrally; intestinal mum thickness 1.25 rnm; living individuals yello-
caecum with 3 pairs of diverticula; males with 5-7 wish-red, with darker red alimentary tract; rhyn-
pairs of spermaries, females with about 25 pairs of chocoel reaching almost to posterior ofbody, with
ovaries: Nectonemertes minima (Fig. 6.1) wall anteriorly containing separate inner circular
8a Rhynehocoel musculature throughout arranged as and outer longitudinal muscle layers but with
outer circular and inner longitudinallayers 9 muscle layer sequence reversed behind brain; 18
9 Males with 4-6 pairs of spermaries, females with proboscis nerves; stylet armature simple, with
8-10 pairs of ovaries; intestinal caecum with 4 very few stylets; intestinal eaecum with 3 pairs of
pairs of diverticula: Nectonemertes primitiva lateral diverticula, intestine with about 60 pairs
(Fig. 6.3) which are unbranched; females with about 25
9a Males with 15-20 or more pairs of spermaries, pairs of ovaries located dorsomedially to lateral
females with 20 or more pairs of ovaries; inte- nerves; males with 5-7 spermaries situated in a
stinal caecum with 6-8 pairs of diverticula: single row on each side of head and reaching far
Nectollemertes mirabi/is (Fig. 6.2) behind brain to anterior base of tentacles.
10 Mouth and proboscis pore opening separately: Distribution: Obtained at various depths between
Ca/ollemertes, only a single species is known in 3500 m and the surface. Known from the North,
this genus and is recorded from the South Tropical and South Atlantic between latitudes
Atlantie, Calonemertes hardyi (Fig. 6.10) 57°N and 35°S, mostly in the eastern regions of
lOa Mouth and proboscis pore opening through same the Atlantic.
aperture: Profope/agollemerfes, only a single spe- Nectollemertes mirabilis Verrill, 1892 (Fig. 1,
cies from this genus has been recorded from the 6.2). Body very flattened, with posterior lateral
South Atlantic, Protopelagollemertes hllbrechti margins developed as thin horizontal fins and pos-
(Fig.6.11) terior end forming a caudal fin; lengtIYup to 60
rnm, width to 10 mm; rhynchocoel nearly as long
as body, with wall containing separate circular and
588 Nemertina
longitudinal muscle layers; 30-60 pairs of intestin- Genus Pelagollemertes

al diverticula as well as 6-8 pairs of diverticula on Body flat, with extensive gelatinous parenchyma
the intestinal caecum, diverticula without ventral between the organs; body walls very thin, muscle lay-
branches, dorsal branches much lobed; sexes ers consisting primarily of separate bundles; 4-25
dimo!phic, females with 20 or more pairs of ova- pairs of intestinal diverticula; mid-dorsal blood vessel
ries along sides ofbody; mature males with pair of rudimentary; testes in males anterior to brain.
muscular tentacles anteriorly, with compact clus- • Pelagollemertes joubilli Coe, 1926 (Fig. 2F, 6.4).
ters of 15-20 or more spermaries on each side of Body elongate, oval, flattened, broadly rounded at
brain; colour in life flesh-pink to red, orange or front and tapering gradually posteriorly; length 10-
bright scarlet, juveniles pale grey or colourless. 24 rnm; rhynchocoel nearly as long as body; stylet
Distribution: Obtained from virtually all depths armature comprising a slightly curved basis with
down to 3000 m or more, living at shallower about 8 conical stylets; 30 proboscis nerves; 18-24
depths in winter (Van der Spoel, 1985). This spe- pairs of intestinal diverticula; females with 10-18
cies has one of the widest distributions recorded pairs of ovaries; males with compact cluster of 4-6
for any pelagic nemertine and has been found in spermaries on each side of body in front of brain.
the North Atlantic from Cuba to southern Distribution: Taken between depths of about 600-
Greenland and above both eastern and western 1750 m, although nets drawn to the surface from
continental slopes, through the equatorial and 4800 m have contained specimens. Known from
South Atlantic to the latitude of South Africa, in the North Pacific (south ofthe Aleutian Trench, in
the North Pacific within a wide area off the coast the submarine canyon at the entrance to Monterey
of California, off Japan and the Kuril Islands, and Bay, California, and in the Coronado Trough off
in the Bering Sea east of Kamchatka, and in the southern California), South Pacific (south of the
South Pacific to 14°50'S, 85°03'W (Coe, 1954a). East Pacific Ridge), North Atlantic (between 45°-
• Nectollemertes primitiva Brinkmann, 1917a (Fig. 500N and 29°-300W) and the South Atlantic
6.3). Body with rounded head, male with slight (Angola Basin).
constriction anterior to tentacles, caudal fin clear- • Pelagollemertes korotkevitschae Friedrich, 1969
Iy demarcated and posteriorly bilobed; length 12- (Fig. 6.5). Body broad, flattened, anteriorly blunt-
20 rnm, width 3-5 rnm; 40 or more pairs of inte- Iy rounded, posteriorly tapering only slightly to
stinal diverticula, 4 pairs on intestinal caecum, end in blunt tail without caudal fin; length 22 mm,
most diverticula distally forked or lobed; females width 12.5 mm; mouth ventral, separate from pro-
with 8-10 pairs of ovaries; males with single row boscis pore; ll pairs of long, slender intestinal
of 4-6 spermaries on each side of head irnmediate- diverticula which are either unbranched or only
Iy behind brain; colour in life translucent with yel- branch towards their distal ends, posterior 3-4
lowish-red internal organs. pairs shorter; females with 7-9 ovaries on each
Distribution: Obtained from depths of 200-1 000 m, side of body situated above or close to lateral
but also taken in open vertical hauls from 3000 m to nerve cords; males unknown; rhynchocoel extends
the surface. Found in the North Atlantic between almost to posterior end of body.
31°-55°N and 29°-36°W, South Atlantic (off the Distribution: Taken from a depth of 1000 m.
west coast of the French Congo) and North Pacific Known only from the tropical South Atlantic (off
(Baja California and off the coast of California). the coast ofGabon, West Africa, at 3°36'S, 9°12'E).
• Pelagollemertes moseleyi Bürger, 1895 (Fig. 6.6).
Family Pelagonemertidae Body sturdy, wide and flat, broadly rounded ante-
Body usually broad and flattened anteriorly, posterior riorly. tapering posteriorly to end in a well diffe-
end often distinctly narrowed and in some genera rentiated caudal fin; length 13-17 rnm, width 8.5-
ending in broad caudal fin; dorsal blood vessel ends 11 rnm, thickness 1-5 rnm; rhynchocoel nearly as
blindly in rhynchocoel a short distance behind the long as body; stylet armature a sickle-shaped basis
brain; mouth and proboscis pore separa te; intestinal bearing about 7 conical stylets; 22 proboscis ner-
diverticula usually with ventral branch; myofibrillae ves; 4-5 pairs of distinct intestinal diverticula, with
present in lateral nerves; spermaries in two groups group of smaller intestinallobes posteriorly; fema-
behind, alongside or in front of brain. les with 6-8 pairs of ovaries; males not known.
Nemertina 589
Distribution: Taken from depths of about 800- Family Planktonemertidae

1500 m. Found in the North Atlantic (approxima- Body broad and generally much flattened; posterior
tely 45°N, 300W), Tropical Atlantic (off the coast end usually with caudal fin; rhynchocoel musculature
of Gabon, West Africa) and the North Pacific off comprising meshwork of interwoven circular and lon-
the southem coast of Japan. gitudinal fibres; dorsal vessel extends almost full
Pelagollelllertes rollestolli Moseley, 1875a (Fig. length of body; dorsal anastomosis of cephalic blood
6.7). Body broad and oval, narrowing posteriorly vessels present; intestinal diverticula much divided,
to a pointed tail with no obvious caudal fin; usually with ventral branch; spermaries in two longi-
lengths of 11-45 mm, widths of 3.5-23 mm and a tudinal groups or rows immediate1y behind brain.
thickness of 3 mm have been recorded; rhyncho- Genus Plallktonelllertes
coel near1y as long as body; stylet armature a sick- Body broad and very flat; mouth and proboscis pore
le-shaped basis bearing about 9 minute conical united; rhynchocoel extends almost to hind end of
stylets and 6 or more accessory stylet pouches; 16- body, with wall containing meshwork of interwoven
22 proboscis nerves; 12-20 or more deeply bran- circular and longitudinal muscle fibres; intestinal
ched and lobed intestinal diverticula; females with diverticula numerous, much branched and with ventr-
4-13 pairs of ovaries; males with cluster of 5-8 al branch extending be10w lateral nerve cords; sper-
spermaries each side of body in front of brain. mari es in two elongated groups or in irregular rows
Distribution: Mostly trawled from depths between behind brain.
350-1500 m, some individuals have been found in Plallktollelllertes vanhoe./felli Brinkmann, 1915
non-closing nets drawn to the surface from depths (Fig. 6.9). Body widest anterior1y, gradually narro-
of 3500-3600 m. A widespread species, known wing posteriorly; after preservation length 20.5
fram the Atlantic (off the west coast of Africa rnrn, width 8.5 rnrn, thickness 4.5 rnrn, but Coe
between the Gulf of Guinea and the Cape of Good (1926: 116) cornrnents that "The worm in life was
Hope), Pacific (off Japan, east ofthe Kuril Islands, doubtless much flatter ... for the figures ... show
east of the Emperor Seamount Chain, in the sub- that the dorsal musculature was violently contrac-
marine canyon off Monterey Bay, California, and ted during preservation"; Van der Spoel's (1985)
near the Galapagos Islands), Indian Ocean (near example was 13 rnrn long and 6 rnrn wide; 24 pro-
Sri Lanka) and south of Australia where the boscis nerves; rhynchocoel nearly as long as body
Pacific, Indian and Antarctic oceans converge. (Van der Spoel, 1985, states two-thirds of body
length), with wall containing interwoven mesh-
Genus Probalaellallelllertes work of circular and longitudinal muscle fibres;
Body small, much flattened, with distinct caudal fin; stylet armature a sickle-shaped basis bearing very
without tentacles; intestinal diverticula with dorsal small stylets; mouth and proboscis according to
and ventral branches. Coe (1926: 116) "open together in a single atrium",
Probalaenallelllertes irellae Wheeler, 1934 but Van der Spoel (1985: 17) states "Mouth and
[=Pelagollemerfes ¡renae in Korotkevich (1955a, proboscis pores separate"; intestinal caecum with 5
1977a)] (Fig. 6.8). Body rounded anterior1y, tape- pairs of branched diverticula, intestine with nume-
ring slightly posteriorly to end in distinctly flatte- rous diverticula, each with dorsal and ventrallobes;
ned tail; length 9-17 rnrn, width 3-3.5 mm, thick- female with 14 pairs of ovaries; males unknown.
ness 1.7-2 mm; colour in life scarlet with orange Distribution: The original specimen was taken in a
spots along the lateral margins; rhynchocoel near- vertical haul between 3000 m and the surface, but
ly as long as body, with wall containing separate a second individual identified as belonging to this
longitudinal and circular muscle layers; proboscis species carne from 515-995 m depth. Known from
slender, with 17 nerves; females with 5-14 ovaries the South Atlantic (west of the Cape of Good
on either side of body; males unknown. Hope) and North Atlantic (approximately mid-
Distribution: Taken in a vertical haul from a depth way between the Canary Islands and the Mid-
of 1000 m to the surface. Known from only two Atlantic Ridge); anatomical differences recorded
individuals found in the South Atlantic (west of between the North and South Atlantic records sug-
the Cape ofGood Hope). gest that the North Atlantic form may not belong
to this taxon.
••
590 Nemertina
Family Protopelagonemertidae stylets, with 6 pouches of accessory stylets; 23-29

Body rather slender, not much flattened; posterior end proboscis nerves; 30-40 pairs of profusely bran-
without caudal fin; rhynchocoel musculature an inter- ched intestinal diverticula, branches extending late-
woven meshwork of longitudinal and circular fibres; rally above and below lateral nerve cords; females
intestinal diverticula with ventral branch beneath with about 30 pairs of ovaries; males unknown.
lateral nerve cord; spermaries in two simple rows Distribution: First taken in non-closing nets drawn
behind brain. to the surface from a depth of 2000 m, this species
Genus Calollemertes . has been subsequently trawled from between
Body rather slender and not much flattened, narrowed 1000-2500 m; one specimen obtained in a net
at both ends and without a caudal fin; intestinal diver- where the water depth was 4400-4800 m but "it is
ticula with ventral branches; mouth and proboscis probable [this) specimen was caught nearer the
pore separate; rhynchocoel extending to posterior end surface" (Coe, 1954a: 236). Recorded from the
ofthe body; North Atlantic (West European Basin, southwest
Calollemertes hardyi (Wheeler, 1934) [=Plankto- of Ireland, and in the Bermuda region), the South
nemertes hardyi in Korotkevich (l955a)) (Fig. Atlantic (in an area between 33°-34°S and 4°-16°E
6.10). Body rounded at both ends, little tapered near the Tristan da Cunha Islands) and the North
anteriorly and posteriorly, distinctly rounded in Pacific (south of Alaska between 48°37'-48°58'N
cross section; colour in life scarlet with irregular and 157°29'-157°50'W).
black markings; length 80-110 mm, width 22-25
rnm, thickness 13 rnm; probo seis stout, 8 rnm in
diameter, with 24 nerves; rhynchocoel extends to Suggested readings
posterior end of body, with wall composed of
meshwork of interwoven longitudinal and circular Brinkmann A. 1917. Pelagic nemerteans from the
muscle fibres; mid-dorsal blood vessel extends to "Michael Sars" North Atlantic Deep-Sea
rear end ofbody; numerous small intestinal diver- Expedition 1910. Rep. Sci. Res. "Michael Sars"
ticula; female with gonads located towards ventral North Atlantic Deep Sea Exp., 1910,3, Part 2:1-
surface. 18. Tlzefirst major systematic treatment o/ bathy-
Distribution: Obtained from depths of 570-1 000 m. pelagic nemertines, establishing most o/the higher
Originally described from the South Atlantic taxonomic categories that remain in Cllrrent llsage.
(33°25'S, 6°31 'E), a second specimen was found at Brinkmann A. 1917. Die pelagischen Nemertinen.
27°02.5'N, 20017.7'W in the North Atlantic (Van Bergens Mus. Skr., 3: 1-194. Comprehensive and
der Spoel, 1985). /ully illllstrated descriptions o/ most o/ the North
Atlantic bathypelagic nemertines; this monograph
Genus Protopelagollemertes complements BrinÁ711aml's 1917a article which,
Body anteriorly and posteriorly narrowed and barely beca l/se o/ editorial restrictions, cOl/Id not be as
flattened; mouth and proboscis pore united; rhyncho- detailed.
coel extends nearly full length of body, with walls Bürger O. 1895. Die Nemertinen des Golfes von
containing a meshwork of interwoven circular, spiral Neapel und der angrenzenden Meeres- Abschnitte.
and longitudinal muscle fibres; intestinal diverticula Fauna Flora Golfes Neapel, 22: 1-743. A magn(fi-
with both dorsal and ventral branches; mid-dorsal cent monograph which, thOllgh oId, contains a
blood vessel almost full body length. wealth o/ historical, anatomical. distriblltional
Protopelagollemertes Izubrechti (Brinkmann, and ecological data on nemertines and is illllstra-
1917a) (=Planktonemertes hubrechti in ted with several superb colol/r plates as well as
Korotkevich, 1955a) (Fig. 6.11). Body large when plates o/ histological drawings.
mature, without caudal fin, more slender and thic- Coe W.R. 1926. The pelagic nemerteans. Mem. Mus.
ker than most bathypelagic nemertines; lengths of Compar. Zool., Harvard Col!., 49: 1-244. The
7-105 rnm, widths up to 35 rnm and a thickness of second major treatise on bathypelagic nemertines.
10 rnm are reported; rhynchocoel nearly as long as inclllding a history o/ the subject and illustrated
body; stylet armature comprising a slender but descriptions o/ many taxa and accol/nts o/ their
sharply curved basis bearing many sharply conical biology. distriblltion and ecolog)'.
Nemertina 591
O'••
"'"
'" '"(Pt7
a>
~
.,.
""
"O•••
';:,
<;>
""
"\)
"l\
o
'"
(r
G'
(j>
@€P
O'
rhynchocoel
~
..• '"
6.2a
Ó
Nectonemertes mira bilis Pelagonemertes joubini

Nectonemertes
minima
¿ .-
~;~~~:J
.~=~C~:
,:::~. 'S"
.;:.. ;-
'-;:-
6.3c
<1
Nectonemertes
Pelagonemertes korotkevitschae primitiva
592 Nemertina
Pelagonemertes
moseleyi
anterior .. ~
proboseis
stylet
basis
posterior
proboseis
Probalaenanemertes
irenae Pelagonemertes rollestoni
6.10b 6.11
9
~
Calonemertes hardyi Protopelagonemertes
Planktonemertes vanhoeffeni hubrechti
Nemertina 593
Coe W.R. 1927. Adaptations of the bathype1agic Gibson R. 1982. Nemertea. In "Synopsis and c1assifi-
nemerteans. Amer. Naturalist, 61:345-352. A cation of living organisms", 1 (S.P. Parker, ed.),
general discussion of various morphological McGraw-Hill, New York, pp. 823-846. An illu-
features of bathypelagic nemertines related to the strated c!assification ofthe phylum Nemertina, lis-
habitat in which they live. tillg all the taxonomic levels down to genera and
Coe W.R. 1945. Plankton of the Bermuda many of the known species.
Oceanographic Expeditions. XI. Bathypelagic Gibson R. 1995. Nemertean genera and species ofthe
nemerteans of the Bermuda area and other parts of world: an annotated checklist of original names
the North and South Atlantic oceans, with eviden- and description citations, synonyms, current taxo-
ce as to their means of dispersa!. Zoologica (New nomic status, habitats and recorded zoogeographic
York), 30:145-168. A useful illustrated general distribution. 1. Natur. Hist., 29:271-562. A world-
accollnt of the bathypelagic nemertines of the wide checklist of all the nemertine genera and spe-
North and South Atlantic, with keys to the families cies; an essential reference work for the current
and genera. systematic nomenclature of the phylum.
Coe W.R. 1954. Bathypelagic nemerteans of the Korotkevich V.S. 1955. Pe1agicheskie nemertiny dal·
Pacific Ocean. Bul!. Scripps Inst. Oceanogr., nevostochnykh morei SSSR [Pelagic nemertines of
6:225-285. A comprehensive illustrated accoUllt oi the Far Eastem seas of the USSR]. Akad. Nauk.
pelagic nemertines found in the Pacific Ocean, SSSR, Moskva, pp. 1-131. A detailed account o/
including keys to families and genera. the bathypelagic nemertines ji-om the north-west
Coe W.R., Ball S.e. 1920. The pelagic nemertean Pacific, including descriptions o/ several new taxa.
Nectonemertes. J. Morpho!., 34:457-485. A detail- Korotkevich V.S. 1964. Pe1agicheskie nemertinui
ed and illllstrated account of the anatomy of this antarkticheskikh i umerennuikh vod yuzhnogo
genus of bathypelagic nemertines. polushariya [Pe1agic nemertines of Antarctic and
Gibson R. 1972. Nemerteans. Hutchinson, London, temperate waters of the southem hemisphere].
224 pp. A /ulZv illllstrated comprehellsive account Issledovaniya Fauny Morei, Nauka, Leningrad,
of all aspects of nemertille biology, including their 2:132-167. An illustrated account ofthe bathypel-
anatomy, physiology, reproduction, ecology, agic lIemertines of Antarctic and southern ¡lidian
::oogeography and phylogenetic relationships. Oceall waters.
Fig. 6. Illustrations of the speeies.

6.la: Mature male in dorsal view, drawn to show the cerebral ganglia and lateral nerves, rhynchoeoel and the extreme posterior portion ofthe
blood system;
6.1 b: Female with partially everted proboseis;
6.2a, 6.2b: Mature mal e and female drawn to emphasise the distribution of the gonads; both individuals viewed from the dorsal aspeel;
6.3a: Male with four pairs of spermaries and short eephalie tentaeles from ventral aspeet;
6.3b: Mature male with fully developed eephalie tentaeles and outline of the alimentary traet;
6.3e: Female in dorsal view, drawn to show the cerebral ganglia and lateral nerves, rhynehoeoel and ovaries;
6.4a: Female in ventral view, drawn lo show the alimentary traet and lateral divertieula, rhynehoeoel, cerebral ganglia and lateral nerves and ovaries;
6.4b: Stylet basis and stylets;
6.5: Female in ventral view, drawn to show the partially everted proboseis, rhynehoeoel, alimentary traet and lateral divertieula, cerebral gan-
glia and lateral nerves and ovaries;
6.6a, 6.6b: Female viewed from ventral (a) and dorsal (b) aspeet, drawn to emphasise the distribution ofthe ovaries, cerebral ganglia and later-
al nerves, lateral blood vessels, rhynehoeoel and arrangement of the alimentary traet;
6.7a, 6.7b: Male (a), and female (b) with proboseis partially everted, viewed from the ventral surfaee, drawn to emphasise the cerebral ganglia
and lateral nerves, rhynehoeoel, gonads and arrangement of the intestinal divertieula;
6.7e: Longitudinal seetion through the middle region ofthe proboseis to show the stylet basis with its minute stylets;
6.8a, 6.8b: Outline drawings of a preserved individual, with proboseis partially everted, in dorsal (a) and lateral (b) aspeet to show the general
body shape and distribution of the ovaries;
6.9a-e: Ventral (a) and lateral (b) views ofthe type speeimen; the stylet basis and stylets (e);
6.IOa, 6.10b: Female viewed from ventral (a) and dorsal (b) aspeets, drawn to show the irregular blaek markings;
6.11: General body shape and appearanee of a female.
Figure sourees: from Brinkmann (1917b), redrawn: 6.la, 6.1 b, 6.3a, 6.9a-e, 6.11; from Bürger( 1895), modified: 6.6a; from Coe (1945), redrawn:
6.2a, 6.2b; from Coe (1954a), redrawn: 6.3b, 6.3e; from Coe(I926), redrawn: 6.4b, 6.7b, 6.7e; from Friedrieh (1969b), redrawn: 6.4a, 6.5; from a
photograph in Friedrieh (1969): 6.7a; from Kato and Tanaka( 1938), redrawn: 6.6b; from Wheeler (1934), redrawn: 6.8a, 6.8b, 6.1 Oa, 6.1 Ob.
Polychaeta
María Ana Fernández-Alamo and Erik V. Thuesen
Introduction A few studies have been undertaken on the physiolo-

gy and biochemistry of pelagic polychaetes (Ikeda,
As some other marine phyla that comprise mostly ben- 1974; Thuesen and Childress, 1993; Childress and
thic forms, the Annelida contain a small minority of Thuesen, 1993). The active forms of pelagic poly-
species that have evolved to spend all their life in the ehaetes have very high metabolic rates, leading
pelagic realm. These holoplanktonic Polychaeta have Thuesen and Childress (1993) to suggest that they
developed a number of special adaptations to live in may have the highest rates of all deep-sea pelagic ani-
the planktonic environment including small size, long mals. High levels of activity by a1ciopid and tomop-
setae, enormous and complex eyes, flattened or gelati- terid polychaetes are supported by in situ observations
nous bodies, a high degree oftransparency, sperm sto- (Video Archive Library, Monterey Bay Aquarium
rage in females, etc. (Fig. 1). Pelagic polychaetes are Research Institute, Monterey, California).
found mainly in the open sea, but they also occur in
neritic regions. They inhabit the entire water column As in almost every other pelagic marine phylum of
from surface layers to abyssal depths. Most research animals, bioluminescence has been observed in pela-
has been undertaken on their distribution and taxono- gic species of the Annelida. In the Alciopidae,
mic relationships, and very little is known about other Alciopina and Krohnia are bioluminescent, but this
aspects of their biology and ecology. ability is probably present in other genera as well.
Clark (1970) describes the histology of the mucus
The holoplanktonic species are mainly active predators glands of Rhynchonerella angelini, and he mentions
which attack their prey with the rapidly everted probos- the possible relationship of these organs with the bio-
cis. However, filter-feeders or phytophagous forms are luminescence in this species. In the Tomopteridae,
also known, for example some species of the family there are photogenic organs located on the parapodia
Iospilidae (Day, 1967). Judging from their behavior which give off an unusual yellow light. This pheno-
and their strong proboscides, the A1ciopidae are active menon is present in Tomopteris nisseni. Qne of the
predators, but there are no direct observations of a1cio- biological functions of bioluminescence is likely the
pid feeding. They are assisted in their search for prey by attraction between males and females, which would
telescopic eyes, which enable them to orient them- be of great importance for pelagic forms since they
selves in different directions, as well as to recognize the usually do not form dense populations. Dales (1971)
size and form of the prey. The tomopterid polychaetes has reviewed this vital function in pelagic worms.
have very short pharynges, and can either consume
their prey whole or suck out the internal body fluids. Qne of the most unique aspects of pelagic polychaetes
The long cirriform appendages of the second segment is the evolution of eyes in the family Alciopidae.
in the tomopterid polychaetes may playa role in prey These complex eyes are very large and telescopic, and
(or predator) detection. Remains of herring fry, frag- each eye has a lens that is controlled by muscles which
ments of siphonophores, chaetognaths and appendicu- can change the direction of its axis. The eyes can
larians have be en found in their digestive tracts apparently determine not only the intensity and direc-
(Rakusa-Suszczewski, 1968). Many other species have tion of light, but also the outline and size of objects.
no obvious prey-catching organs, and possibly they This must be of great advantage to the species in this
feed on microscopic prey or eggs in the plankton (Day, pelagic family since it enables them to survey their
1967). The aberrant polychaete Poeobius meseres cap- surroundings in different directions. The first studies
tures falling detrital matter using a mucous web (Uttal- of these complex structures were undertaken by Greeff
Cooke, 1992). Members of the family Typhloscoleci- (1876) and Demoll (1909), and their function has been
dae rnay be neotenic forms in connection with their described by Hermans and Eakin (1974) and Wald and
transition to ectoparasitism, and their pharynx is modi- Rayport (1977). The eyes ofthese annelids are remark-
fied into a characteristic suctorial organ. ably similar to those of cephalopods and vertebrates,
SOllth Atlalltic Zooplankton.

1999 BackhllYs Publishers, Leiden, The Netherlands
596 Polychaeta
Basic body plan of a

pelagic polychaete
(Pe/agobia /ongicirrata)
antenna
eyes
tentacular _
cirrus
setigerous
Anterior region in
ventral vlew
m
lobe
(A/ciopina parasitica)
E
::J
E notopodial
ramus
.9
VI
l'Il
.•....•
Q)
E __ -:.~ "¡,'he,
septentnonalf5, ¡,-",," , ...4"~. "'.r
70:'''' r1 '~~~ g ,ed
'""~ /~ :~.~',,~~"
neuropodial
chromophil
gland
ramus
~
compound
setae ~7Wl ~~~
,'4
~ ,>
\ tlrst two pair~ afussetae

Lopadorhync/J on theof
uncmatus
parapodia
\ ,simple acic I
acicular-cornpound seta
of Rhynchonerella petersi
Types of setae <i Types of par~podial

tomoptend polydlaetes
glands in 'i'
\!I
Polychaeta 597
and beca use of their different developmental origin, Geographic distribution

they have been cited as an extreme example of conver-
gent evolution (Salvini-Plawen and Mayr, 1977). Since the middle part of the twentieth century, strong
Wald and Rayport (1977) have emphasized the correlations between the distributions of planktonic ani-
extraordinary evolutionary convergence represented mals and the movements of water masses in the oeeans
by the possession of accessory retinas in alciopid poly- have been established. Tebble (1960) has demonstrated
chaetes, cephalopods and deep-sea fishes. Recent find- such eorrelations for some species of pelagic poly-
ings of a small multi-gene family of packed box-con- ehaetes in the South Atlantic Ocean, and Tebble (1962)
taining genes (Pax gene s) in different metazoan phyla, and McGowan (1960) for pelagic polychaetes in the
which play an important role in embryonic develop- North Paeific Ocean. St0p-Bowitz (1948) found differ-
ment in the nervous system, has revealed that their ences between the pelagie polyehaete fauna in the west-
expression is structurally similar in different visual em and eastem parts of the North Atlantic Ocean, the
systems, yet ontogenically distinct (Zuker, 1994). westem part being richer, both in species and abun-
dance. In addition, some speeies oecur only in the east-
Uschakov (1972) pointed out, correctly, that in the ge- em half while they are absent in the westem half, and
neral economy ofthe sea, the role ofthe 8000+ species vice versa. Orensanz and Ramírez (1973) have studied
of Polychaeta is tremendous because they represent a the distribution ofpelagic polychaetes in the southwest-
very important link in all processes ofthe production of em Atlantic Ocean. They recorded 27 species from the
living matter, and in particular, they are the basic food, families Lopadorhynchidae, Iospilidae, Alciopidae,
rich in calories, of numerous fishes. Although Typhloscolecidae and Tomopteridae, and also some
Uschakov was referring to benthic or meroplanktonic larval and epitokous forrns. Orensanz et al. (1974)
species, it is clear that holoplanktonic polychaetes play described the distribution of nine polychaete speeies
an important role in the pelagic ecosystem. from antarctic waters near Argentina. St0p-Bowitz
(1992) has documented the distribution of pelagic poly-
chaetes in the eastem South Atlantie offthe eoast ofthe
Methods Africa between the equator and about 200S.
Mero- and holoplanktonie species of polychaetes are The distributions of some holoplanktonic species have
eommon in plankton cornmunities, and they are col- been mapped by Tebble (1960) in the South Atlantic
leeted by eornmonly used methods, such as standard Oeean. He discusses their zoogeographical pattems,
and horizontal hauls with open or opening-closing and describes endemic, eosmopolitan and tropical-
nets of different mesh sizes (0.2-0.5 rnm). Many spe- subtropical species. Tebble (1960) concluded that
cies of pelagic polyehaetes are very fragile and are although there were three endemic species of antarctic
often damaged during colleetion. Nets with large vo- pelagie polyehaetes, there were no endemic speeies of
lume: mouth ratios, large eod ends and slow towing pelagic polychaetes in the subantaretic Atlantic. He
speeds improve the condition of the fragile species. also noted that the Subtropieal Convergence was not a
Samples are best preserved immediately in 4-5% for- barrier to the distribution of any planktonie polychaete
malin in filtered seawater neutralized with a borax- speeies. Five speeies (Tomopteris planJ..1onis, T. sep-
saturated solution. Although, long-terrn preservation tentrionalis, Typhloscolex mul!eri, Travisiopsis levin-
in an ethanol solution is not recommended for gelati- seni and Pelagobia longicirrata) were found in all the
nous worrns, "working specimens" may be kept in a investigated hydrological zones of the South Atlantie
70% ethanol solution. The glands in the parapodia of Ocean. The Discovery samples that Tebble (1960)
the tomopterid polychaetes can be seen with the naked analyzed also included some specimens eollected with
eye, but this requires care and some praetice. The opening-closing nets, and he provided some depth dis-
chr0IT!.0phil glands stain deeply with haematoxylin tribution data. However, much more work is needed in
which helps to observe these structures. order to properly elucidate the bathymetric ranges of
•
598 Polychaeta
pelagic polychaetes. Distribution records of 63 species Fig. IC) and simple acicular setae (stout, similar to the
of pelagic Polychaeta in the South Atlantic Ocean are aciculum, Fig. 1C). Compound setae consist of a basal
shown in Fig. 2. and distal parts. The distal part may be flattened to forro
a blade, and may have serrations (Fig. 1C). Compound
setae may also be acicular, as in some species of
Morphology, identification and systematics Rhynchonerella (Fig. IC). Fauchald (1977) has
remarked on the importance of accurate examination of
Morphology and identification these structures through precise microscopic observa-
tions. Each parapodium may bear dorsal and ventral
The body plans of polychaete worros often reflect their structures named parapodial cirri that are variable in
habits and habitats. This is shown in the great diversity forro and size. In addition, some glands are associated
of structural forros that deviate from the basic metame- with the parapodia, and they have taxonomic impor-
ric body. The basic polychaete body plan ineludes the tance in the holoplanktonic families Alciopidae and
folIowing: (1) a head consisting of a prostomium and a Tomopteridae. In the Alciopidae, the segmental glands
peristomium; (2) a segmented body or metastomium; or organs are located lateralIy, dorsally or ventralIy
and (3) the anal segment or pygidium. Variations ofthe between the parapodial lobes on most body segments
morphological characteristcs of these parts of the body (Fig. 3.6b, 3.7b, 3.12b, 3.15a, 3.16b, 3.20a, b). UsualIy
are important for the identification of polychaetes. The they are pigmented and their forro and pigmentation
holoplanktonic species are highly specialized for pela- pattem are used in the identification of some species
gic life, and the main morphological characters used for (Fig. 3.17a). In the Tomopteridae, there are various
their identification will be described briefiy. types of parapodial glands located in the notopodial and
neuropodial pinnules which are membranous structures
The basic body plan of a pelagic polychaete is illustra- adapted for swirnming (Fig. ID). Chromophil glands
ted in Fig. l. The head presents a prostomium with a are always present in all species, and occur only in the
variable number of antennae (1, 2, 4, 5, 6), eyes with ventral or distal regions of neuropodial pinnules, from
different degrees of development, and nuchal organs the third parapodia on toward posterior (Fig. ID). They
which appear as ciliated pits, grooves or lobes with dif- are often swollen and can be stained by haematoxylin.
ferent forros in typhloscolecid polychaetes (Fig. 3.53a, Spur glands are present in some species. They are alone
3.54a, 3.55a, 3.56a, 3.57b, 3.59b). The peristomium of in the first two or three parapodia, but associated with
the head is usually integrated by fusion of some seg- chromophil glands in the subsequent parapodia (Fig.
ments with welI developed food-gathering appendages ID). In some species, rossete glands are present in the
referred to as tentacular cirri (Fig. lB). There is often trunks ofthe first two parapodia (Fig. ID) and in the
an eversible buccal cavity or proboscis that may have notopodial and neuropodial pinnules of later parapo-
chitinous jaws (Fig. 3.24b). Proboscides are variable in dia (Fig. ID). Hyaline glands occur near the apex of
size and in the distribution of their papilIae (Fig. 3.9a, the pinnule as elear areas around a yellowish spot
3.15a, 3.18a, 3.19a, 3.20a). In the metastomium each (Fig. ID). These structures are very difficult to see
segment of the body bears a pair of lateral parapodia. because the spot is often missing. Day (1967) recom-
These structures are typicalIy biramous with the para- mends a brief staining with haematoxylin to make the
podial trunk dividing into a dorsal ramus, the notopo- elear area readily' visible. This basic plan of the para-
dium, and a ventral ramus, the neuropodium. This bira- podia can be modified whereby any part of the para-
mous condition is present in tomopterid polychaetes podium may be suppressed. All the setae may be
(Fig. ID), but in the other holoplanktonic families the lacking (achaetous) as in tomopterid polychaetes. The
parapodia are uniramous. Each ramus consists of a setigerous lobes may not be present, leaving only dor-
setigerous lobe supported by a stout intemal chitinous sal and ventral cirri (apodous), as the first parapodia
rod or aciculum, and a bundle of chitinous setae which in the genus Vanadis (Fig. 3.15, 3.20a). The cirri may
show an infinite variety offorros (Fig. 3.1b, 3.4b, 3.29b, be lost, such as the ventral cirrus in the first parapodia
3.27b, 3.32c). Setae are also referred to as chaetae by in some species of the genus LopadorhyncJzus (Fig.
some zoologists. Holoplanktonic families present two 3.32b, 3.33b), or the cirri may be well developed, as
main kinds of setae: simple and compound. Simple in alI species of the family Typhloscolecidae (Fig.
setae are capilIary setae (slender hair-like structures, 3.5Ia,3.54a).
Polychaeta 599
r-----.,-, ----c,.--~-.,-,----,
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, Pseudalciopa ----~--.---.---:----- 600
,
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, Naiades cantraini -,----------r------- modesta , , Plotohelmis a/ata
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o;:::::::: Plotohelmis capitata

___ ~.- - - - - - - - - - r - ~
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'\·-·-------r··---- Torrea candida 600
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2.10------:----·----:-----···-:---- 2.11 ------:---------:---------:----

• 2.1 2 ------~------.--~---.--
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, . bongraini f- 400
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Rhynchonerella
____ .0. __ .
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Watelio gravieri -,------·---r----- angelini -::

~;~;:~~~:~S~~~~}
¡~~~~~:::: i¡::::::::;:
600
600 400 200 00 _ 200 600 400 200 00 200 600 400 200 00 200
600 Polychaeta
2.13-·----~-·····---·-······--"·-·· 2.1 4 ....--~-----

..--~-.----.-."-.-. 00
• 2.1 5 ------~---------.---------:---~
---~-.- ... -----------------.-
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400
____ J , _
Rhynchonerella _ ____
;_________ Rhynchonerella -1600
,----------r----- gracílis ;__________ Rhynchonerella.
longicirrata ----------,-._--- moebii
12.1 6 ~-------h~---------j-h- 112.1 7 ------~---------i---------j---· 112.1 8------~---------

-~--.-~--¡-.: 100
............ -.- ---.-~- 11 - ; ---- - 11 --~----- --------+)~-

--------- --------- ,
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---- .... - .._---_.- -------- Vanadis ,
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•••
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Rhynchonerella _ ~~E
~~~~~~~ ___
~;.• .. __.,_.. . Vanadis _ 600
____ J
,
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_
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-:._--.--_.;..... _-_... !~--' , ; .. ~--------- longissima
2. 1 9------~····-·--·~-··-----·~-··· 2.20------:------·--- -- - . 2.21 ---..-~---- ~-- ~.... 100
-------- - - - - - - -•
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"';::::::::T:::::: Vanadis minuta -,----------,------ Vanadis studeri ---- __________

--------- v.anad,s.. vlOlacea .1600
2.22----·-j-········· -- .-• .
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ti. :
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---------- •... -._------ ---------- •... --------- ----_ ..... _-----_ .. - - - .. - - - - - - - - - -... ..- - - - - - ~- - - - - - - - - ,- - - - - - - - - -.-,
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~----------:----------~-
_ Vanadis 600
~--.-------~---------;--
brevirostris .----:----------¡---
-,----------r-- Vanadis crystal1ina- ..-.;::::::::::::::: Vanadis formosa
600 400 200 00 200 600 400 200 00 200 600 400 200 00 200
Po1ychaeta 601
2.25 : . 2 .26 ~ ~ . 2.27-·····~········· j . 00

•
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400
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····;::::::::T:::: Vanadis tagensis ............ ,._---- lospilus affinis . ~···;::~~~:::.Iospilus

phala~roides· 600
,-------,-,------.,,---,-, ----, , , , , , ,
2.28-·····:····· Phalacrophorus
pictus
2 .29 : : ~ . 2.30- :
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.... l .:.. Phalacrophorus. Lopador~ynchus. 600

..~.:~::~:
:::::r:::::::::f: .;.~:
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Pedinosoma curtum 600
,.------ ••• r- ••• ~----------~---------¡----~: : -,--------_.
600 400 200 00 200 600 400 200 00 200 600 400 200 00 200
602 Polychaeta
2.3 7 ------i----------~---~··---i---- 2.38------:----------;---------

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Tomopteris kefersteini ----;~~~~~~~~
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600 400 200 00 200 600 400 200 00 200 600 400 200 00 200
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coniceps : , •...
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1
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600 400 200 00 200 600 400 200 00 200 600 400 200 00 200
604 Polychaeta
12.61 ------~-------<---------i----
__Travisiopsis _
112 .62 ------~----~----
---------j-.:~
. . ~...~. 112 .63 ------~----------~---------~--- 100
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levinseni ••
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---- ----------:---------- Travisiopsis -1600
--- ---------- I
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~~:~~:~::::
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----f ---------:- ----;:::::::::::_ Travisiopsis lobifera" _ _ ~-------- lumbricoides
600 400 200 00 200 600 400 200 00 200 600 400 200 00 200
Fig. 2. Charts of the horizontal distribution of pelagic Polychaeta species in the South Atlantic.
Finally, the pygidium which is the last segment ofthe Caruncle: protuberance from the posterior section of
body that bears the anus, only conserves a pair of cirrÍ the prostomium carrying nuchal organs (Fig 3.54a).
named anal or pygidial. They are taxonomically Chaeta (pl. chaetae): see seta.
important only in the family Typhloscolecidae (Fig. Chaetiger: see setiger.
3.53b, 3.54b, c, 3.55b, 3.56b, 3.59a). In the Chromophil gland: a densely staining gland in the
Tomopteridae, the pygidial region may be elongated ventral pinnule of Tomopteris (Fig. 1D).
bearin-g rudimentary parapodia and receives the name Chromatophore: a special cell or group of cells carry-
oftail (Fig_ 3.36a). ing pigment (Fig. 3.12b).
Cirrus (pl. cirri): sensory appendage which is usually
The following list of terms and definitions is given to slender and cylindrical (Fig. lA, B).
assist the reader with identification of pelagic poly- Compound seta: seta having a terminal portion which
chaetes. Several sources of polychaete terminology articulates with the shaft (Fig. IC).
were used for definitions, including Fauchald (1977), Distal end-piece: the tellninal section of a compound
Pleijel and Dales (1991), and Glasby (1996). seta (Fig. IC).
Eye: Iight sensing organ which usually occurs on the
Achaetous: see asetigerous. prostomium but may occur elsewhere on the body
Acicula (pl. aciculae): supportive chitinous rod found or appendages (Fig. lA, B).
intemally in projecting parapodial rami (Fig. 3.30c). Eversible proboscis: part of pharynx capable of being
Acicular setae: stout, projecting setae (Fig. IC). extended by tuming the inner part outwards (Fig.
Anal cirrus (pl. anal cirri): cirrus projecting from the 3.9a).
pygidium (Fig. 3.56b). Genital papilla: projection on which a reproductive
Antenna (pl. antennae): sensory projection arising duct opens.
from the dorsal, lateral or frontal surface of the Head: collective term used for a structure formed by the
prostomium (Fig. lA). fusion of peristomium, prostomium and in many
Achaetous paddles: wide parapodia without setae cases, one or more cephalised segments (Fig. lA).
(Fig. 10). Hyaline gland: gland in the pinnules of certain species
Apodous: segments without parapodia. of Tomopteris; usually translucent, sometimes
Asetigerous: segment without setae (Fig. ID). with a yellow spot in the center (Fig. 1D).
Biramous parapodium: parapodium with two bran- Intermediate cirrus: a cirriform structure attached to
ches, a notopodium and a neuropodium (Fig. 1D). the body wall between the notopodium and neuro-
Blade: distal, f1attened portion of a compound or sim- podium.
ple seta (Fig. 1C). Jaws: simple or complicated hardened apparatus with
Capillary setae: long slender setae tapering to afine opposable units situated in the eversible pharynx.
point (Fig. IC). Lobe: conical process of a parapodium (Fig. lA).
Polychaeta 605
Metastomium: segmented portion of the body Pygidium: post-segmental terminal part of the body
between the prostomium and pygidium (Fig. 1A). bearing the anus (Fig. lA).
Neuropodium (pl. neuropodia): ventral branch (ramus) Ramus (pl. rami): branch, used about the dorsal or
of a parapodium ventral branch of a parapodium (Fig. ID).
Notopodium (pl. notopodia): dorsal branch (ramus) of Rosette gland: yellowish star-shaped gland found in
the parapodium. the two first rami and in the pinnules of some spe-
Nuchal: pertaining to the neck. cies of Tomopteris (Fig. ID).
Nuchal lappet: nuchal organs forming posterior out- Segment: one of the serially repeated units comprising
growths from the prostomium. the trunk, separated intemally or extemally by septa.
Nuchal organs: paired chemosensory structures usual- Segmental organ: organs occurring in segmental
Iy situated postero-1aterally on the prostomium, arrangement, specifically used about the nephridia
variously developed as pits, grooves, lappets, or and nephridial ducts.
elongated ridges (Fig. 3.53a, 3.55a). Segmental glands: glands occurring in segmental
Nuchal papilla: dorsal papilla found on the posterior arrangement (Fig. 3.7b).
part of the prostomium. Seminal receptacles (= receptacula seminis): recepta-
Palps: paired sensory or feeding structures arising cles formed from the parapodiallobes of the most
from the ventro-Iateral surface ofthe prostomium. anterior parapodia which receives and sto res sper-
Palpode: an anterior papilla-like prolongation of the matozoa in some female alciopids (Fig. 3.16a,
prostomium in typhloscolecids (Fig. 3.52). 3.18a, 3.19b, 3020b).
Parapodium (pl. parapodia): one of a pair of lateral Seta (pl. setae): chitinous bristle protruding from a
appendages (sometimes greatly reduced) from a pocket in the body wall (Fig. 1C).
body segment, often with a main trunk and two Setigerous lobe: the projection or part of the notopodium
branches (rami), usually bears setae (Fig. lA, D). or neuropodium which bears the setae (Fig. lA).
Peristomial cirri (= tentacular cirri): paired elongated Shaft: proximal part of a compound or simple chaeta
cirri on the peristomium which may be derived (Fig. IC).
from both pre-segmental and segmental sources Setiger: segment bearing setae (Fig. 3.1 b).
(Fig. lA, B). Spathulate: blade-shaped, like a spatula.
Peristomium: strictly, the pre-segmental region ofthe Spur glands: gland with a spur-like projection, situat-
body surrounding the mouth. In practice also ed alone on the ventral side ofthe first pair ofneu-
inc1udes segments fused to the rear of the peristo- ropodial pinnules and distally to the chromophil
mium. Sometimes bears cirri and food-gathering gland in posterior parapodia (Fig. iD).
tentac1es. Spiniger: seta that tapers to a fme point, usually refer-
Parapodial cirrus: appendage, often sensory, of the ring to a compound seta (third seta from left in
parapodium (Fig. lA, B). Fig. lC).
Parapodial rami: branch ofthe parapodium (cf. neuro- Setigerous (= chaetigerous): bearing setae.
podium, notopodium). Tentacular cirrus: dorsal aml/or ventral cirri of para-
Parapodial trunks: the portion of the parapodium aris- podia of cephalised segments, often more elongate
ing from the body segment (Fig. iD). than those of more posterior parapodia; have been
Peristomium: the pre-segmental region of the body confused in the literature with peristomial cirri
surrounding the mouth. (Fig. lA, B).
Pharynx: anterior portion of the digestive tract modi- Tentacular segment: segment bearing tentacles (Fig.
fied for feeding. lB).
Pinnules: membranous projections of the neuropodia Unarmed pharynx: pharynx without chitinous hooks
and notopodia of Tornopteris used as swimming (Fig.3.10a).
paddles and bearing glands (Fig. iD). Uniramous parapodium: parapodia with only one
Proboscis (pl. proboscides): eversible, non-muscular branch, usually the neuropodium (Fig. 3.34b).
part of the pharynx (strictly); often used inter-
changeably with pharynx. Systematics
Prostomium (pl. prostomia): anteriormost, pre-seg-
mental region ofthe body, may bear eyes, antennae Overviews of the systematics of the pelagic polychaeta
and palps, and enc10ses part ofthe brain (Fig. lB). are given by Greeff(1876), Bergstrom (1914), Fauvel
606 Polychaeta
(1916), Wesenberg-Lund (1935, 1939), St0p-Bowitz Chiaje, 1830; Ploto/rell1lis Chamberlin, 1919;
(1948), Dales (1957), Day (1967), Uschakov (1972), Pselldalciopa Step-Bowitz, 1991; R/ry"c/ro"erella
Rice (1987), and Pleijel and Dales (1991). Uschakov Costa, 1864; Torrea Quatrefages, 1850; Valladis
(1972) suggested that the pelagic families, despite their Claparede, 1870; Watelio St0p-Bowitz, 1948
biological differences and external morphologies could Family Iospilidae Bergstrom, 1914
be placed in the suborder Phyllodociformia using the Genera IospilllS Viguier, 1886; Phalacrop/rorlls
name proposed by Levinsen in 1882 without systema- Greeff,1879
tic rank. This suborder belongs to the order Family Lopadorhynchidae Claparede, 1870
Phyllodocemorpha (Livanov, 1940, fide Uschakov, Genera Lopadorhy"chus Grube, 1855; Mallpasia
1972), which inc1udes the marine polychaetes with the Viguier, 1886; PedillOSOl1la Reibisch, 1895;
most primitive characters. Pleijel and Dales (1991) Pelagobia Greeff, 1879
have considered that holoplanktonic families of uncer- Family Pontodoridae Bergstrom, 1914
tain phylogenetic affinities belong to the superfamilies Genus POlltodora Greeff, 1879
Phyllodocoidea (Alciopidae, Lopadorhynchidae and Family Tomopteridae Grube, 1848
Pontodoridae), Iospiloidea, Typhloscolecoidea and Genus TOl1lopteris Eschscholtz, 1825
Tomopteroidea in the order Phyllodocida. Family Typhloscolecidae Uljanin, 1878
Genera Sagitella Wagner, 1872; Travisiopsis
Fauchald (1974) reviewed the morphological and ana- Levinsen, 1885; Typhloscolex Busch, 1851
tomical features used in the taxonomy of recent poly- Family Yndolaciidae St0p-Bowitz, 1987
chaetes, and discussed some principies of their phy- Genus Ylldolacia St0p- Bowitz, 1987
logeny. He proposed that the evolution ofthe different Order Fauvelopsida Fauchald, 1977
polychaete families was a rapid radiation from several Family Flotidae Buzhinskaya, 1996
ancestors adapted to life in the semi-consolidated Genus Flota Hartman, 1967
detritus layer above the sea bottom. Step-Bowitz
(1984) has discussed the systematic problems in the Identification
families Alciopidae, Tomopteridae and
Typhloscolecidae. Pleijel (1991) suggested that Order Phyllodocida
holoplanktonic families are derived from benthic Family Alciopidae
phyllodocid polychaetes. The systematic study of The main diagnostic character of the members of this
some of these families are inc1uded in major works, family is a single pair of large and complex eyes.
such as Greeff (1876), Bergstrom (1914), Uschakov Prostomium small with four, five or six antennae.
(1972), and Pleijel and Dales (1991). Three, four or five pairs oftentacular cirri and an ever-
sible proboscis, with short papillae or long terminal
Eight families of holoplanktonic polychaetes have horns. Body slender and elongate with numerous seg-
been recorded from the South Atlantic Ocean. ments, almost always broken into fragments in zoo-
Pontodora pelagica has not yet been recorded in this plankton samples. Parapodial glands are present, late-
region, and it belongs to a monogeneric and monospe- rally, dorsally or ventrally between the parapodial
cific family, Pontodoridae. The phylogenetically iso- lobes on most body segments; these are referred to as
lated polychaete Poeobius meseres has only been "segmental glands". The glands range in size from
found in the North Pacific Ocean. The diagnostic small oval projections in some species to elongated
characters of the families and recorded species from kidney-shaped structures extending from the dorsal to
the Atlantic Ocean are given below. the ventral cirrophore. A pair of anal cirri is present,
but it is usually difficult to see due to body damage. It
Outline c1assification is assumed by the presence of the eversible muscular
proboscis, sometimes provided with a pair of lateral
Order Phyllodocida Clark, 1969 projections for grasping prey, that alciopid poly-
Family Alciopidae Ehlers, 1864 chaetes are predators; however, the nature of the prey
Genera Alciopa Audouin and Milne Edwards, is unknown (Day, 1967). By the presence ofthis struc-
1833; Alciopilla Claparede and Panceri, 1867; ture, their complex eyes, and their behavior, Uschakov
KroJlIlia Quatrefages, 1866; Naiades Delle (1972) suggested they are active predators. Rice
Polychaeta 607
(1987) deseribes the behavior, in situ, of some speeies 7 Prostomium large and bulbous, divided into
ofthis family from the Bahama Islands in the Westem hemispheres by a median groove. Segmental
Atlantie Oeean, and gives new information on mor- organs as lateral bars behind the basis of the pa-
phology using eleetron mieroseopy and diseusses rapodia. Males without genital papillae:
aspeets of reproduetion, and systematies. He added Plotohelmis tenllis (Apstein, 1900) (Fig. 3.14)
two new taxonomie eharaeteristies (sperm type and 8 Aeieular setae simple 9
body transpareney), and eonstrueted an evolutionary 8a Aeieular setae eompound .10
tree of the aleiopid polyehaetes. In general, he agrees 8b Two kinds of eompound setae ll
with St0p-Bowitz (1949) in the generie relationships, 9 Segments 4-6 with only 2-7 simple aeieular
dividing the genera of this family into two somewhat setae and their dorsal eirri mueh larger and more
distinet groups on the base of the type of setae. rounded than later ones: Rhynchonerella moebii
(Apstein, 1893) (Fig. 3.10)
Key to genera and species 9a Simple aeieular setae and eompound eapillary
1 Setigerous lobe without a eirriform appendage setae present from the first parapodia. The first
........................................................................... 2 few dorsal eirri not obviously different from
1a Setigerous lobe with a single eirriform appendage later ones: RhyncllOnerella gracilis Costa, 1861
.............................................................................. 3 (Fig.3.9)
lb Setigerous lobe with two eirriform appendages 10 Distal end-pieees of aeieular setae small and
.......................................................................... .4 simple. Prostomium and dorsum strongly pig-
2 Setae eapillaries only: Naiades calltrailli Delle mented: Rhy"chonerella angelini (Kinberg,
Chiaje, 1830 (Fig. 3.1) 1866) (Fig. 3.7)
2a Setae eompound spinigers only: Torrea calldida lOa Distal end-pieees of aeieular setae large and clear-
(Delle Chiaje, 1841) (Fig. 3.2) Iy serrated. Head not pigmented: Rhy"cilollerella
2b Setae eapillaries and simple aeieulae: Alciopilla petersi (Langerhans, 1880) (Fig. 3.11)
parasitica Claparede and Paneeri, 1867 (Fig. 3.3) 11 Compound setae with smooth or dentieulate
2e Setae eompound spinigers and simple aeieulae: end-pieee on the first seven or eight parapodia.
Plotoilelmis 5 Cirriform appendages on the pedallobe are very
3 Setae eapillaries and simple aeieulae: Kroilnia smal!. Segmental glands are seldom pigmented:
lepidota (Krohn, 1845) (Fig. 3.5) Rily"cho"erella bOllgraini (Gravier, 1911) (Fig.
3a Setae eapillaries and eompound spinigers: 3.8)
Watelio gravieri (Benham, 1929) (Fig. 3.4) 12 Prostomium with 4 antennae 13
3b Setae eompound spinigers only: Vanadis 12 12a Prostomium with 5 antennae 15
3e Setae eompound spinigers and simple or eom- 12b Prostomium with 6 antennae 18
pound aeieulae: Rily"cilo"erella 8 13 Proboseis with a pair of lateral homs. Females
4 Setae eapillaries and eompound spinigers: with a pair of seminal reeeptacles .14
Alciopa reYllalldi Audouin and Milne Edwards, 14 First 6-7 p¡lirs of parapodia modified, aehaetous
1829 (Fig. 3.6) with very small dorsal and ventral eirri.
5 Segmental glands with ventro-Iateral ehromato- Segmental glands lightly pigmented before seg-
phores 6 ment 12, if at all: Valladis minuta Treadwell,
5a Segmental glands without ventro-Iateral ehro- 1906 (Fig. 3.18)
matophores 7 14a First 5-6 pairs of parapodia modified. Segmental
6 Chromatophores ventro-Iateral branehing on the glands as vertieally pigmented bands before seg-
ventrum. Males with four pairs of genital papil- ment 7: Vanadis studeri Apstein, 1893 (Fig. 3.20)
lae on ventral region of segments 12 to 15: 15 Proboseis terminated by 12 equal small papillae.
Plotoilelmis capitata (Greeff, 1876) (Fig. 3.12) Segmental glands and body strongly pigmented
6a Chromatophores ventro-Iateral but no bran- at intervals 16
ehing. Males with ten pairs of genital papillae on 15a Proboseis terminated by 12 triangular marginal
ventral region of segments 13-22: Plotohelmis papillae. Segmental glands present from seg-
alata Chamberlin, 1919 (Fig. 3.13) ment 5, never pigmented and body uniform vio-
let when fresh: Valladis violacea Apstein, 1893
608 Polychaeta
15b Proboseis terminated by two long horns and a 3 First two ehaetigers redueed, only with small
varying number of srnall papillae 17 lobes and few short setae: Phalacrophorlls pic-
16 Dark bars around body every five or ten seg- tllS Greeff, 1879 (Fig. 3.25)
ments: Valladis IOllgissima (Levinsen, 1885) 3a First seven ehaetigers redueed only with small
(Fig. 3.17) lobes and few short setae: PhalacropllOrllS IIlli-
16a Dark bars aeross dorsum at interva1s of two or forlllis Reibiseh, 1895 (Fig. 3.24)
three segments: Valladis alltarctica (McIntosh,
1885) (Fig. 3.21) Family Lopadorhynchidae
17 Parapodia well developed and with setae from Worms with flattened body. Prostomium broad with
the third foot posteriorly: Vanadis formosa four antennae, no palps and poorly developed eyes.
Claparede, 1870 (Fig. 3.16) Eversible proboseis, simple and unarmed. With three
17a Parapodia well developed and with setae from tentaeular eirri on the first one or two segments whieh
the seventh to tenth parapodia posteriorly: may laek setae. Parapodia uniramous with dorsal and
Valladis crystallina Greeff, 1876 (Fig. 3.15) ventral eirri. Setae mainly eompound with serrated
18 Proboseis with 12 terminal papillae, ofwhieh two end-pieees. The serrations are of taxonomie impor-
lateral pairs are longest. Four pairs of tentaeular tanee and must be examined in lateral view (Pleijel
eirri: Vanadis tagensis Dales, 1955 (Fig. 3.19) and Dales, 1991). Pygidium without eirri.
In addition to those included in the key above, alciopids Praetieally nothing is known of the biology of this
reported from the South Atlantie include three more group. None have ever been found with food in the
speeies deseribed reeently from the Gulf of Guinea by gut, but eertainly they do not feed on phytoplankton
Dr. Step-Bowitz (1991), most of them with only one (Day, 1967). Usehakov (1972) speeulated that the
speeimen: Pselldalciopa modesta Step-Bowitz, 1991, simple hook-shaped setae of the first modified para-
Rhynchonerella longicirrata Step-Bowitz, 1991, and podia of Lopadorhynchlls might hold the prey.
Vanadis brevirostris Step-Bowitz, 1991.
Key to genera and species
Family Iospilidae 1 Two pairs of tentaeular eirri on the first segment
Small worms with prostomium rounded, without ........................................................................... 2
antennae. Two minute palps and two eyes. Proboseis 1a Two to three pairs of tentaeular eirri on the first
eversible, with or without a pair of lateral ehitinous segment. Next two to three segments with para-
hooks. First two or ten segments of the body with podia modified, with only eurved, simple aeieu-
redueed parapodia. Later parapodia uniramous with lar setae: Lopadorhyllchlls 3
short dorsal and ventral eirri, a longer setigerous lobe 1b Three pairs of tentaeular eirri, on two first seg-
and eompound spinigerous setae. Day (1967) men- ments; eaeh segment with setigerous lobes and
tions that speeies of the genus Iospilus include some few setae: Mallpasia 6
of the few holoplanktonie polyehaetes whieh feed on 2 Prostomium triangular with four small antennae
diatoms, and speeies of Phalacrophorus whieh have and a pair of eyes. Tentaeular segment with a
ehitinous hooks are possibly predators. Usehakov few setae and two long, tapered eirri. Later pa-
(1972) suggested that these jaws are developed as rapodia with a eonieal setigerous lobe and longer
adaptive struetures to pelagie life, for eatehing rapid- dorsal and ventral eirri essentially similar, more
ly moving planktonie organisms. or less eylindrieal. Compound setae with basal
joints smooth: Pelagobia longicirrata Greeff,
Key to genera and species 1879 (Fig. 3.35)
1 Proboseis unarmed: lospillls 2 2a Body small and flattened with few segments.
la Proboseis armed withjaws: PhalacropllOrlls 3 Round prostomium with two pairs of lateral .
2 First two ehaetigers without eirri, and small antennae. Tentaeular eirri without setae. All sub-
lobes with few setae: Iospillls affillis (Viguier, sequent parapodia with eordate dorsal eirri, eo-
1911) (Fig. 3.23) nieal setigerous lobes and laneeolate ventral
2a First two ehaetigers with eirri: Iospilus phala- eirri. Setae all fine and eompound: Pedinosollla
croides Viguier, 1886 (Fig. 3.22) cllrtlllll Reibiseh, 1895 (Fig. 3.26)
Polychaeta 609
3 First two pairs of parapodia modified, with very small eyes. Two pairs of tentacular cirri presento
curved and simple acicular setae ..4 Uniramous parapodia with slender, elongated pedal
3a First three pairs of parapodia modified, with lobes that are extended until the long composite setae;
curved and simple acicular setae 5 dorsal and ventral cirri are oval and small.
4 First two pairs of parapodia are large and stout POlltodorapelagica Greeff, 1879 (Fig. 3.29) has not
with strong acicular setae which are rounded at yet been recorded in the South Atlantic Ocean. It
the base by a collar-like expansion: Lopador- has been reported from warm waters in the Atlantic
"Y"Cllls lI11cillatlls Fauvel, 1915 (Fig. 3.30) Ocean (Dales and Peter, 1972), the Agulhas and
4a First two pairs of parapodia are stout, but not lar- Mozambique Currents (Day, 1967), and from warm
ger than subsequent ones, with a linear series of and cold waters in the Pacific Ocean (Treadwell,
stout simple, hooked acicular setae and lack ven- 1943; Uschakov, 1955; Berkeley and Berkeley,
tral cirri: LopadorllYllchlls krolllli (Claparede, 1960; Femández-Álamo, 1983, 1992).
1870) (Fig. 3.31)
4b First two pairs of parapodia stouter and shorter Family Tomopteridae
than subsequent ones, with simple acicular setae. Polychaetes characterized by parapodial reduction to
The ventral cirri are partially fused to the inferior biramous achaetous paddles with notopodia and neu-
margin of the setigerous lobes: Lopadorhy"clllls ropodia modified for swimming into membranous
lIellseni Reibisch, 1893 (Fig. 3.32) structures named pinnules. These structures may carry
4c First two pairs of parapodia are Sh0l1 with incon- various types of glands including chromophils, hya-
spicuous, hooked setae. The ventral cirri are lines, rosettes and spurs. Dales (1957) gives a good
present partially fused to the setigerous lobes. description ofthese organs. Prostomium with one pair
Ventral cirri on the four and subsequent parapodia of antennae and simple eyes. First two segments fused
with a long filiform tips: Lopadorhy"chllS with the prostomium and carrying cirri supported by
appelldiclllatlls Southem, 1909 (Fig. 3.33) aciculae. Second segment with long cirriform appen-
5 Setigers 1-3 shorter and stouter than subsequent dages. Short and unarmed pharynx.
ones, they have only simple acicular setae and
lack ventral cirri: Lopadorlly"cllllS brevis Very little information about the biology ofthe mem-
Grube, 1855 (Fig. 3.34) bers of this family is available. In general, nothing is
5a Setigers 1-3 shorter and stouter the subsequent to be seen in the transparent gut. Uschakov (1972)
ones, with simple acicular setae. They have the believes that they are voracious predators.
ventral cirri partly fused to the distal inferior
margin of setigerous lobes: Lopadorhy"chllS Key to species
Ilatiollalis Reibisch, 1895 (Kim, 1967, has con- Abbreviations used in key, Pin: Pinnules; Hy: Hyaline
sidered this species synonymous with L. brevis glands; Ros: Rosette glands; Chr: Chromophil glands;
Grube, 1855). Sp: Spur glands.
6 Posterior parapodia with dorsal cirri foliaceous 1 Body is prolonged into a tail bearing rudimen-
and ventral cirri lanceolate: Mallpasia coeca tary parapodia 2
Viguier, 1886 (Fig. 3.27) 1a Without tail 5
6a Posterior parapodia with dorsal cirri digiform or 2 First pair of cirri present.. 3
subulate and ventral cirri blunt 7 2a First pair of cirri absent .4
7 Dorsal tentacular cirri of segment 2 longer than 3 Pin restricted to the parapodial rami. Ros present
twice the body width: Mallpasia gracilis in the trunks of the first two parapodia and at
Reibisch, 1893 (Fig. 3.28) apices of both rami of later parapodia. Chr large
7a Dorsal tentacular cirri of segment 2 less than and swollen from the third parapodia and sub se-
twice the body width: Mallpasia isochaeta quently on all parapodia. Sp present from second
parapodia onwards (fram the third are associated
Family Pontodoridae with Chr): TOlllopteris nationalis Apstein, 1900
A monogeneric and monospecific family with a small (Fig. 3.36)
and slender pelagic polychaete, whitish and fragile 3a Pin extended on the distal ends of parapodia
body. Rounded prostomium with four antennae, and trunks. Hy present in the apices on dorsal and
610 Polychaeta
ventral Pin from the third parapodia. Chr volu- fourth parapodia: TOl1lopteris e/egalls Chun,
minous situated in the inferior half ofthe ventral 1887 (Fig. 3.39)
Pin from 3-4 parapodia onwards: TOlllopteris 6a Pin rounded and restricted to parapodial rami.
kral1lpi Wesenberg-Lund, 1936 (Fig. 3.37) With brownish pigment at the tips of both dorsal
3b Ros present in the trunks of the first two parapo- and ventral rami. Chr situated ventrally in the Pin:
dia and in both pinnules of the following parapo- TOl1lopteris kefersteini Greeff, 1879 (Fig. 3.41)
dia. Chr large from the fifth parapodia and sub- 6b Pin oval. Ros in the trunks of the first two para-
sequently on a11 parapodia. Sp well developed podia and near the tips of both rami thereafter.
from the third parapodia located in the inferior Chr rounded situated ventra11y from the third
part of the ventral Pin: TOl1lopteris duccii Rosa, parapodia: TOl1lopteris ro/asi Greeff, 1882 (Fig.
1907 (Fig. 3.38) 3.42)
4 Pin broad with an inner clear area and a narrow 7 Pin oval and restricted to parapodial rami. Chr
fri11y outer margin containing the glands. we11 developed, from fourth parapodia, with a
Glandular outer edge not extending between the triangular form, and situated at the proximal end
rami in juveniles. Hy present towards the distal of the ventral region of the Pino Hy and Ros
end of the inferior part of the ventral Pin from absent: TOl1lopteris p/allktonis Apstein, 1900
the third foot onwards. Chr swollen from fourth (Fig. 3.48) ..•
parapodia: TOl1lopteris euchaeta Chun, 1887 7a Pin oval the dorsal extending onto the distal end
(Fig. 3.43) of the parapodial trunk. Hy present at the apices
4a First pair of cirri present in juveniles. Pin restrict- of ventral Pin from 2-4 parapodia. Chr not com-
ed to the parapodial rami. Ros in the trunks of the pact situated between the ventral and apical
first two parapodia and near the tips ofboth rami region ofPin from the fourth parapodia onwards:
thereafter. Sp present from 2-3 parapodia. Chr TOl1lopteris septelltriollalis Quatrefages, 1865
large and start on the third parapodia: TOl1lopteris (Fig. 3.49)
dunckeri Rosa, 1908 (Fig. 3.44) 7b Body very large, Pin oval, fri11yand extend on to
4b Body large and flaccid. Nuchal organs we11 the distal ends ofthe parapodial trunks. Hy deep-
marked. Parapodia swo11en with short, conical Iy pigmented from the third parapodia in the
rami. Black spots present on the dorsal and ven- superior half of ventral Pino Chr large and volu-
tral Pin from the third to subsequent parapodia. minous from the fourth parapodia, in inferior
Chr we11 marked from the third parapodia: half of Pin: TOl1lopteris carpellteri Quatrefages,
TOl1lopteris kel1lpi Momo, 1930 (Fig. 3.45) 1835 (Fig. 3.50)
4c Body large, taillong. Pin reduced to a fringe bor- 7c Pin continuous with dorsal and ventral ridges
dering the parapodia. Hy in variable position along the parapodial trunks. Hy in a dorsal posi-
very ventral and dorsal Pino Chr prominent, tion from the third ventral Pino Chr ventra11y
present from the fourth parapodia posteriorly. from the fourth parapodia: TOl1lopteris /igu/ata
Cirriform appendages of second segment may Rosa, 1908 (Fig. 3.40)
exceed length of body: TOl1lopteris nisseni
Rosa, 1908 (Fig. 3.46) In addition to those listed in the key, Tomopteris circu-
4d First cirri present in juveniles, absent in adults. losa Smp-Bowitz, 1992, T congo/ana Smp-Bowitz,
Pin oval. Ros present from the first parapodia on 1992, T gracilis Step-Bowitz, 1992, T guineensis
the inner side of the Pin, next to the apex of each Smp-Bowitz, 1992, T lutea Step-Bowitz, 1992, and T
ramus. Chr inconspicuous from the first parapo- simplex Step-Bowitz, 1992, have been described
dia: TOl1lopteris he/go/alldica Greeff, 1879 (Fig. recently from the Gulf of Guinea by Dr. Step-Bowitz
3.47) (1992), most of them from only one specimen.
5 First cirri present 6 According to Day (1967), T apsteini Rosa, 1908 is
5a First cirri absent.. 7 probably synonymous with T nationalis Apstein,
6 Parapodial trunks long and slender with oval 1900. Tomopteris cavalli Rosa, 1907 was not
Pino Chr well developed from fourth parapodia, considered valid by Tebble (1960). Day (1967) put
situated towards the apical ventral region of Pino Enapteris euchaeta within the genus Tomopteris on the
Hy restricted to the dorsal Pin of the third and basis of a careful description of the parapodia.
Polychaeta 611
TOlllopteris ofliscijormis Eschscholtz, 1825, the 4a Nuehal organs extended backward as lobes sim-
type species of the genus, has been considered as ple and digiform. Anal cirri long and ovate: Tra-
indeterminable; it was reported by Step-Bowitz visiopsis lanceolata Southem, 1911 (Fig. 3.54)
(1992), but its characteristics are very similar to T 4b Nuchal organs extended baekward as lobes digi-
rolasi Greeff, 1882 (cf. Orensanz and Gianuca, 1974) form and paralIel. Anal eirri subtriangular:
in the same paper, and it is important to clarify if the Travisiopsis lobifera Levinsen, 1885 (Fig. 3.56)
latter is synonymous with the first. 4c Nuchal organs extended backward as lobes se-
micircular. Anal cirri spathulate: Travisiopsis
Family Typhloscolecidae levinseni Southem, 1911 (Fig. 3.55)
Worms highly specialized, with fusiform and trans- 4d Nuchal organs extended backward as branehed
parent body. Prostomium indistinct from the rest of lobes. Anal cirri spathulate: Travisiopsis coni-
the body and projected forward into a small papilla. ceps (Chamberlin, 1919) (Fig. 3.52)
Without eyes, with welI developed nuchal organs
which often form free posterior projections. Pygidium In addition to the above, Step-Bowitz (1991, 1992)
with a pair of flattened anal cirri. reeorded only one specimen of Travisiopsis lumbri-
coides Reibisch, 1895 (Fig. 3.57) in the tropical
These worms have be en considered to be ectopara- waters (10m depth) of the Gulf of Guinea.
sites of gelatinous animals, such as chaetognaths and
medusae. The charaeteristie suetion apparatus that Family Yndolaciidae Step-Bowitz, 1987
replaces the proboscis may indicate that they are Worms with short and flat body; redueed prostomium
semi-parasites (Uschakov, 1972). Feigenbaum (1979) without antennae, palps or eyes. Proboseis short,
has regarded the possibility that typhloseoleeids are unarmed with deep ventral slit. Peristomium and fol-
predators of ehaetognaths. lowing segment with a pair of large tentacular cirri;
body segments with biramous parapodia with dorsal
Key to genera and species and ventral cirri in needle form; compound setae, each
1 Prostomium with large transverse ciliated lobes with a long slender blade.
dorsatly and ventralIy: Typltloscolex 2 Yndolacia lopadorrhyncllOides Step-Bowitz, 1987
la Prostomium with nuehal organs as curved ridges (Fig. 3.58) is the only species in the family. It is
without freely projeeting lobes. No projeeting bathypelagic (800-900 m depth), and found in the
earunele: Sagitella 3 equatorial region off the eoast of Africa.
1b Prostomium with nuchal organs extended baek-
ward as free processes. A earuncle usualIy Order Fauvelopsida
present Travisiopsis .4 Family Flotidae
2 Prostomium blunt and bearing a small asymmet- Nine segments, parapodia biramous, with setae eha-
rical palpode. Ciliated lobes almost as broad as racteristicalIy cross-barred. Body and parapodia com-
the head, the dorsal one with a pair of small pletely eneased in transparent gelatinous sheath.
rounded nuchal lappets at the sides: Bathy- to abyssopelagie.
Typltloscolex lIlulleri Busch, 1878 (Fig. 3.59) • Flotajlabelligera Hartman, 1967 (Fig. 3.60). At a
3 Prostomium with a small palpode. Head with a total length of 25 mm, it has been found off the
pair of semicircular nuchal ridges on dorsal coast of southwestem Chile and off Cape Hom
region but no free nuchal lobes: Sagitella kowa- (Hartman, 1967). It will most likely be found at
lewskii Wagner, 1872 (Fig. 3.51) abyssal depths in the southern seetion ofthe South
4 Nuehal organs extended backward as rounded Atlantic Ocean. One other species in this family
lobes. Anal cirri oval: Travisiopsis dubia Step- has be en described, Flota vitjasi Buzhinskaya,
Bowitz, 1848 (Fig. 3.53) from the western North Pacific Ocean.
612 Polychaeta
Suggested readings Pleijel F., Dales R.P. 1991. Polychaetes: British phyl-
lodocoideans, typhloscolecoideans and tomopte-
Dales R.P. 1955. The evolution ofthe pelagic alciopid roideans. Synopses of the British Fauna (New
and phyllodocid polychaetes. Proc. Zool. Soco Series), 45, Linnean Society of London and
London, 125:411-420. Contains some ideas about Estuarine and Coastal Sciences Association, pp. 1-
phyletic relationships between pelagic pol)'- 195. The most recent review ofthe species ofsome
chaetes, based on comparison of structures holoplanktonic families ji-om the British Islands;
derived from the stomodeum. inc/udes remarks on biology, ecology, methods of
Dales R.P. 1957. Pelagic polychaetes of the Pacific research and ~ystematics of the British species
Ocean. Bull. Scripps Inst. Oceanogr., 7:99-168. A belonging to the superfamilies Phyllodocoidea,
general review ofthe systematics and distribuliolls Typhloscolecoidea and Tomopteroidea.
of the pelagic species ji-om the Pacific Ocean. St0p-Bowitz C. 1948. Polychaeta. Rep. Sci. Res.
Inc/udes descriptions with illustrations, synonyms "Michael Sars" North Atlantic Deep Sea Exp.,
and charts of distriblltion ofthe species, and iden- 1910,8:1-77. An excellent revietl' ofthe distribu-
tification keys. tion of pelagie species in the North Atlantic
Dales R.P., Peter G. 1972. A synopsis of the pelagic Oeean; includes illustrated descriptions, syno-
polychaeta. 1. Natur. Hist., 6:55-92. Inc/udes nyms and general remarks on geographical and
information of the species belonging the holo- hydrographical factors, :!oogeography, hori::.ontal
planÁ10nicfamilies Lopadorhynchidae, Iospilidae, and vertical distribution of the species.
Pontodoridae, Alciopidae, Tomopteridae and St0p-Bowitz C. 1981. Polychaeta. In "Atlas del
Typhloscolecidae. A complete list of synonyms, Zooplancton del Atlántico Sudoccidental y méto-
taxonomic comments, general distribution and dos de trabajo con el zooplancton marino" (D.
location of the type of each species, and bibliogra- Boltovskoy, ed.), Publico Esp. Inst. Nac. Inv.
phy. Desarrollo Pesq., Mar del Plata, pp. 471-492.
Day 1.H. 1967. A monograph on the Polychaeta of General review ofthe systemalics and distriblltion
southem Africa. Parto 1 Errantia. British Mus. of the pelagic species ji-om the sOllthwestern
Natur. Hist., London, Publ. 656, pp. 1-458. Atlantic Ocean. II7c/udes briefremarks on biology,
Illllstrated descriptions and distribution of the methods of research, history of study, vertical dis-
polychaete fauna, including the Indian and tribution, keys of identifieation, ete.
Atlantic Oceans. Inc/udes the species of holo- St0p-Bowitz C. 1984. A short note on the problem
planktonic families with a complete list of valid with systematics of some pelagic polychaetes.
species. synonyms, records in the survey area, Proc. I Intemat. Polychaete Conf., Sydney,
::.oogeographic provinces, depth ranges, etc., and Australia. A review ofthe main diagnostic charac-
general distriblltiol7 in the world ocean. ters in the exc/usively pelagic families.
Orensanz J.M., Ramirez F.C. 1973. Taxonomía y St0p-Bowitz C. 1992. Polychetes pélagiques des cam-
distribución de los poliquetos pelágicos del pagnes de "I'Ombango" dans les eaux équatoriales
Atlántico Sudoccidental. Bol. Inst. Biol. Mar. et tropicales ouest-africaines. ORSTOM Ed., Coll.
(Mar del Plata), 21: 1-86. Very good review of the Etudes et Thesis, Paris, pp. 1-115. Inc/udes the
systematics and distribution of the pelagic poly- systematies and vertical and geographical distri-
chaetes ji-om the Southwestern Atlantic Ocean; butions of the species collected during twelve
inc/udes illustrated descriptions and distribution crllises in the Gulf of Guinea.
data. Tebble N. 1960. Distribution ofpelagic polychaetes in
Orensanz J.M., RamÍrez F.C., Dinofrio E.O. 1974. the South Atlantic Ocean. Discovery Rep.,
Resultados planctológicos de la Campaña 30: 161-300. An excellent review of the distribu-
"Oceantar 1" -II: Poliquetos.'Contrib. Inst. Ant. tions of pelagic species and their relationships
Argentino, 184: 1- 41. Describes the distribution of with the Subtropical Convergence, Antarctic
the planktonic polychaetes collected in the Conl'ergence and other water masses in the survey
Atlanlic Sector ofthe Southern Ocean. Inc/udes an area; inc/udes descriptions of the species and
illustrated chart for identification of the species. remarks on ::.oogeographieal aspects.
Polychaeta 613
J ]1' ~
~ iae,dV:.t:2
Naiades cantraini •
~ iae,dv)
1iIO'~
Torrea candida
b Ipl
•
I----l
0.25
b (p)
Alciopina
d
parasitica
a
(ae, dv)
~~~iae,t
~~~ :4b1P1
>------l
1
Watelio gravieri ••
Krohnia lepidota •
a
b (p)
lae,dv~I'
~~:-:_-~... _~ ~
Alciopa reynaudl ,0.5 ~
(ae, dvi /f!

~&I'
§¿~~
(ae~~
I----l
5 8)
N-~ 1--------<
I~
5
Rhynchonerella bongraini
Rhynchonerella moebii CD>
a
~ (ae, dv) b (p)
-_.-.
~=.:: ~- Rhynchonerella
gracilis
n~~~~
8)
•.. -. - 1 1---------1
<2:r.-
.,. ---?=
. 1 0.5
·-=-·:I~ ~
Rhynchonerella petersi ~ Plotohelmis capitata
614 Polychaeta
a b
a
o" a 5
(ae~dV) I
~b? Plotohelmis a/ata -G!) ~~~:\S:
-~ GD
Plotohelmis 1-----<
0.5
tenuis
a Vanadis
(9ae,dv) crystallina
Vanadis ~
formosa ~
Vanadis
longissima GfI C)
b
(9ae, dv)
12
-- - -'P Vanadis tagensis
e (p)
~ I
0.51
••• Vanadis antarctica e>

Polychaeta 615
k-
0.5
>------< b (p)
~
b (p) I------l
0.1
lospilus phalacroides G&
C!J) lospilus affinis
i.~
#C~~I
__.. ~
iwa,dv)
I
~(p)
~~ ~(P)
./ ~ 1------< >---l
'c(P)~ "'-:-:k 0.25
0.1
~ ~ Pedinosoma
Phalacrophorus uniformis curtum ~
~
Phalacrophorus pictus CD
a
(wa, dv)
1--1
0.1
Maupasia coeca
Maupasia gracilis ~
1,
Pontodora pelagica
•
616 Polychaeta
~B.
~~--- ~~
~ L....t 11 ~Lj~ 3
5
I f~.
~.~~-~
"'_
,-
~ - =
-,
.
3
(wa, dv)
(ae, vv)
b
e (p) (ae'dvi:~~
(ae, vv)
b
~
~ 1
e (p)
Lopadorhynchus uncinatus Gn>

~ ~9~'~ ~
~~ ~0.5
~ ~ ~ ~n~ ~
L
~
Lopadorhynchuskrohni ~
~ ~ ~'I~
~ b ~~ 7 c(p)
t------i
0.5
(ae,3d v ) (ae, vv) 3
Lopadorhynchus henseni
el iwa,dvlC
Lopadorhynchus
appendiculatus ~
j¡L~ ~~ ~Il
~
cF.b~~r¡
(p) (ae, vv)· ~GJ)
'IiI-/
6'«< -_ .>
~;>
~~~ ~ ~
~-
Q- ?-
~t-~
Is
~ ~ ¡¡;-~~
~
~b
-- ~~-
(ae, dv)
12
/ ~
¡;/I
(e)
ae, vv
~
I-----i
0.5
GD
Lopadorhynchus brevis
11
t-------<
3 1
(wa, dv) Pelagobia longicirrata ..,

Polychaeta 617
"'""'" ,...'" '.. .••.... -

,,.
r-:::1 í ~ ~..::.-~.
- ..... '. _-<o,.
~I_._-~ ~ a .' b(p)

t------<
1 •
(ae, VV)
Tomopteris krampi CIiI
Tomopteris
ligulata ~I
~
a
(ae, dv) ~~
f1/~ .0
~f)~
~I'b~ I I
(ae, dv)
Tomopteris roJasi ~
Tomopteris kefersteini 0.5 CID
~I)~
~
Tomopteris
(ae, dv)
euchaeta
~¡--' b(p)~
~
a
(ae,dV~'~~
. ~__~
;]) ;:-_.~~ ~
b(p) ~
Tomopteris dunckeri CID

b (p)
a(p) ~-----
(~
~ kemp,
Tom01!teris
t------<
,
Tomopteris nisseni '1
n~ Tomopteris heJgoJandica ~
Tomopteris
pJanktonis ~~í,~~
Tomopteris septentrionalis ~ ~
•
618 Polychaeta
b (p)
~\~)
~~~
~ (a~,.dV)
~~dV)
Il
Tomopteris carpenteri •• ~-~\'
'.'
¡;.;-
1
:::
/--I~I
(j;/
SagiteJla kowalewskií CID

~
Travi~iopsis ~
comceps
a
(wa, dv)
-- l'
~I
Travisiopsis dubia
.;::::::..
1___
:::::
(ae, dv)
b
1
I
Travisiopsis
lanceolata ~
Travisiopsis
levinseni
CD
l~'Q
I~ /)::) 1
b (pe)
-:;:-8
-.--:
¿ 3
a ~
(wa, dv)
!
11 Travisiopsis
lumbricoides ~
~
Travisiopsis lobifera
a
a (wa, dv)
(ae, dv) animal in
gelatinous
sheath
11
b
(ae, dv) )5
C)
Yndolacia lopadorrhynchoides Typhloscolex muJleri CB> G>
Polychaeta 619
•••
Fig.3. Illustrations ofthe species. All scales are in rnm. lp: first parapodium; ae: anterior end; dv: dorsal view; Iv: lateral view; p: parapodi-
um; pe: posterior end; vv: ventral view; wa: whole animal. All figures redrawn; sources: after Benham (1929): 8, 50; after Chamberlin (1919):
52; after Fernández-Álamo (1983): 1,3-7,9-20,22,24-37,39,43,44,46,48,49,51,53,54, 59; after Hartman (1967), composite by M.T.
Biondi: 60; after Malaquin and Carin (1922): 42; after Mclntosh (1885): 21; after Monro (1930): 45; after Monro (1936): 56; after Reibisch
(1895): 57; after Rosa (1908): 38,40; after Step-Bowitz (1987), composite by M.T. Biondi: 58; after Viguier (1911): 23; after Wesenberg-Lund
(1935): 41, 47; after Wesenberg-Lund (1936): 55; after Wesenberg-Lund (1939): 2.
Heteropoda
Gotthard Ric/¡fer and Roger R. Seapy
Introduction distinctly broadened. The larvae are planktotrophic,

feeding on small phytoplankton and zooplankton;
The Heteropoda are holopelagic prosobranch gastro- changing to carnivory after metamorphosis. Duration
pods that are found in moderately low to low abun- of the larval stage is unknown.
dances throughout the world's oceans, mostly at tro-
pical to subtropical latitudes. AII heteropods are Three families (Atlantidae, Carinariidae and
carnivorous, utilizing image-forming eyes to locate Pterotracheidae) comprise the superfamily Heteropoda.
prey and a protrusible radula with elongate, hook-sha- The Atlantidae, which exceed by far the other two
ped teeth to capture their prey. families in number of species and abundance, are gene-
rally considered to be the most primitive family. Their
Heteropods exhibit a series of striking adaptations to soft parts can be fully retracted into the dextrally (right-
the pelagic environment. The foot (orparts ofit), sole- hand) coiled shell and sealed off at the shell aperture by
like in bottom-dwelling gastropods, is laterally com- a chitinous operculum. The shell of atlantids has
pressed and enlarged as a swimming fin. In normal undergone striking modifications, compared with the
position all heteropods swim with their ventral side shell of their presumptive bottom-dwelling ancestors.
(and the ventral fin) turned upward (conseguently the The adult shell (teleoconch) develops directly, and
dorsally-positioned shell or visceral nucleus hangs without a sharp dividing line, from the larval shell (pro-
below the body). Gn the postero-ventral edge of the toconch). The last body whorl is greatly enlarged and
fin a sucker is situated, which is large in the laterally flattened, bearing a keel along its outer edge.
Atlantidae and smaller in the Carinariidae and Both the flattened body whorl and keel function to sta-
Pterotracheidae. The eyes are highly developed, with bilize the body in the water and counterbalance the
large, spherical lenses and narrow retinas. The tae- skulling movements of the large, laterally-compressed
nioglossate radula (tooth formula = 2-I-R-I-2, see swimming fin. Swimming movements appear aw-
below) is characterized by enlarged lateral and mar- kward, being clearly hampered by the short body. The
ginal teeth with elongated shafts and hook-shaped necessity to retract the animal completely into the shell
ends. Like all taenioglossate radulae, the marginal leaves little room for alterations. Nonetheless, there are
teeth are swivel-mounted and their long shafts spread remarkable differences between the various species of
out laterally when the radula is protruded from the Aflanfa and the monotypic Oxygyrlls. In the former
mouth, forming a highly effective means of snaring genus the shell seems far too large for the animal.
prey. The bodies and shells of heteropods are largely When completely retracted into the shell, almost one-
transparent; only the eyes and viscera are opague. Al! half of the enlarged, outermost shell whorl remains
heteropods are dioecious and show sexual dimorp- unoccupied. This can be attributed, at least in part, to
hism (e.g., large penis and penial appendage in males, the highly-functional increase in shell diameter (in-
radular differences in some atlantids, absence of suc- crease in lateral surface area to increase stabilization
ker in female pterotracheids). Internal fertilization fol- during swimming and sinking). In Oxygyrlls the shell
lows spernl transfer by spermatophores during copu- lumen is almost completely filled with the retracted
lation. Fertilized eggs are layed singly or in animal. The body is elongated and, in conjunction with
egg-strings; the latter either free-floating or attached the bilateralized shell, swimming motions are much
to the female (Firoloida). Larvae are free-swimming swifter and more accurate than in Aflanta. In fact, the
veligers, with right-coiled, calcareous shells. The lar- swimming behavior of OA)'gyrlls resembles that of
val velum is small and bilobed in newly-hatched lar- Carinaria much more than Aflanfa.
vae; later deeply fissured with six (Atlantidae,
Carinariidae) or four (Pterotracheidae) long, slender The Carinariidae and Pterotracheidae consist of ani-
lobes. The velum is bilaterally symmetrical, except mals whose bodies are greatly enlarged, drawn out in
for one species of pterotracheid in which one lobe is the anterior-posterior axis and basically cylindrical.
South Atlalltic Zooplallktoll.

edited by D. Boltol'skoy, pp. 621-647
© 1999 BackllllYs Publishers, Leidell, The Netherlands
622 Heteropoda
The elongated esophagus connects the buccal region the number of species rose to about 100 before Tesch
with the viscera, which are compacted into a structure (1906) started to revise the chaotic taxonomy of the
termed the visceral nuc1eus. At a location posterior to group. For the complete list of references on
the eyes the esophagus is modified as a kind of maw Heteropoda up to 1974, we refer the interested reader
which holds prey organisms during a preliminary to van der Spoel (1976). Reviews ofthe biology ofthe
phase of digestion (attributed to secretions of the sali- Heteropoda were given by Thiriot-Quiévreux (1973)
vary glands; Gabe, 1952). The relatively small visceral and Lalli and Gilmer (1989).
nuc1eus inc1udes the true stomach, intestine, digestive
gland, kidneys, heart, gonads and gills. In Carinariidae
the calcareous shell is reduced to a cap-like covering Methods
over the stalked visceral nuc1eus (Carinaria and
Pterosoma), or is represented by the protoconch plus a Studies on heteropods have been based almost exc1u-
rudimentary teleoconch embedded in the mantle tis- sively on animals collected using plankton nets and
sues ofthe visceral nuc1eus (Cardiapoda). The body of trawls or observations on animals surviving in aquaria
the carinarids is more rigid than that of the pterotra- for relatively short periods of time. Limited in-situ
cheids, resulting in propulsion exc1usively by skul!ing observations (Hamner et al., 1975; Lalli and Gilmer,
movements of the fin (like in the atlantids). The 1989) have provided our meager knowledge on the
Pterotracheidae lack a shell in the adults (larvae shed natural behavior of heteropods (e.g., swimming, fee-
the shell at metamorphosis). The visceral nuc1eus has a ding, mating, buoyancy, escape responses).
distinctive pyriform shape that is largely enveloped by
the cylindrical body. In the Pterotracheidae, with Because heteropods range widely in size a variety of
longer and slimer bodies, the swimming movements of different types and sizes of nets (ranging from small,
the fin are reinforced by lateral undulations of the fine-meshed plankton nets to large, coarse-meshed
body, especially in stressful situations. trawls) have been employed by researchers. Like
many other planktonic animals, heteropods are able to
A number of evolutionary trends are suggested among avoid relatively slow-moving plankton nets with nar-
the shell-bearing species (Atlantidae and Carinariidae) row openings (McGowan and Fraundorf, 1966;
that involve reduction in shell weight: 1) a decrease in Seapy, 1990a); a factor that is rarely appreciated or
the number of larval and adult shell whorls (e.g., taken into account.
Atlanta lesllellri, A. oligogyra and all species of
Carinaria); 2) reduction in the thickness (range = 3-40 Fixation and preservation of heteropods can best be
/lm; Richter, 1973a; Batten and Dumont, 1976) of the accomplished using 2% formal in in sea water, buffered
shell walls (e.g., A. lesuellri, A. Fagilis, A. meteori, A. to pH 8.3 with sodium borate (borax). Use of a pH
gibbosa and al! species of Carinaria); 3) a secondary buffer is critical because the aragonitic shells of
dissolution of the inner shell walls, so that only a thin Atlantidae and Carinariidae are extremely sensitive to
organic membrane remains to divide the whorls (e.g., reduced pH and will dissolve rapidly. Shell dissolution
A. inflata, A. helicinoides and A. calijorniensis); and is a significant problem for shelled molluscan
4) replacement of the aragonitic shell material in zooplankton in general (Tumer, 1976). If heteropods
Atlanta by conchiolin (e.g., the keel in Protatlanta and are maintained for long periods of time in liquid pre-
both the keel and shell in Oxygyrlls). servative, the pH should be checked every three
months. Specimens in rich plankton samples are even
The literature on Heteropoda is diverse and wide- more endangered beca use the gradual deterioration of
spread, dating from the first species description proteins increases the acidity of the solution; even
(Pterotrachea coronata) given in 1775 (Niebuhr, ms. when buffered. Because of health hazards associated
ForskiH). However, sound heteropod taxonomy can be with formaldehyde exposure, many researchers trans-
dated from the year 1852, when Eydoux and Souleyet fer formalin-fixed specimens to 80-95% ethanol
published their famous paper on the Heteropoda ofthe (Thiriot-Quiévreux, cited in Turner, 1976, recom-
Bonite Expedition. Due to uncritical investigations mends 90-95%). Below a concentration of 80%
based on damaged or badly-preserved specimens or ethanol requires buffering to neutralize acids soluble in
the overvaluation of single morphological characters, alcohol. A major drawback to the usage of ethanol,
Heteropoda VLJ
however, is that it causes dehydration and bleaching of Nearly all heteropods are epipelagic (Tesch, 1949;
tissues, in addition to shrinking the body; a particular Thiriot-Quiévreux, 1973), although two species of
problem in Pterotracheidae and Carinariidae. Although Pterotrachea (P. coronata, P. scutata) and one spe-
a variety of Iiquid fixation and preservation procedures cies of Carinaria (c. lamarch) appear to be mesopel-
have been developed, none offer the ideal solution. agic and undergo nocturnal vertical migration into the
epipelagic zone (Pafort-van Iersel, 1983). Vertical
By far the best method for preserving atlantid and cari- distributions of heteropod 1arvae in the Mediterranean
narid shells is to dry them and store them in acid-free were reported by Thiriot-Quiévreux (1968) and
glass tubes or gelatine capsules. As a testament to this Richter (1968, 1973b). In the latter study, vertical
method, the invaluable type specimens of atlantids migration by heteropod 1arvae was recorded. Seapy
collected and described by Souleyet (1852a) are main- (1990a) reported on day-night distributional patterns
tained dry in the British Museum and are still extant, of atlantids (11 species) and pterotracheids (2 species)
whereas the type specimens are stored in ethanol in from the epipelagic zone off Hawaii. While diel diffe-
the Musée d'Histoire Naturelle de Paris and the shells rences could not be distinguished for upper epipe1agic
are completely dissolved. species, lower epipelagic species cornmon1y migrated
toward the surface at night. Michel and Michel (1991)
reported on vertical distributions of heteropods from
Geographic and vertical distribution the Florida Straits. Low abundances of large atlantids
(size range = 4 to 8.5 rnm) were obtained, which can
General patterns be attributed to the coarse mesh (3 mm) of the MOC-
NESS trawl. Nonethe1ess, their results for the atlantid
Although we are uncertain about the occurrence of a Atlanta peroni supported the conclusion of Seapy
number of species in the major oceans, our present (1990a) that this is a lower epipelagic species that
understanding of global distribution patterns (Table 1) migrates into the upper epipelagic at night.
is that most species are cosmopolitan; 12 of 21 species
of Atlantidae, 4 of 9 Carinariidae, and all Pterotrachei- Heteropods occur in moderately 10w to 10w abundan-
dae. These distributional data are provisional, insofar ceso Highest abundances have been recorded for atlan-
that they are valid only for the species recognized tid species, although several pterotracheids (particular-
today. The latest descriptions of species new to scien- Iy Firoloida desmaresti) can be as abundant as many of
ce (van der Spoel and Troost, 1972; Richter, 1990, the atlantids in a given location. Generally, however,
1993; Seapy and Richter, 1993) show us that we can- atlantids are numerically dominant, followed by pte-
not be certain whether or not we really know all extant rotracheids and carinarids. Quantitative abundance data
heteropod species. Moreover, large areas ofthe oceans (numbers per unit volume ofwater filtered) are limited,
are still poorly investigated (among these the South and the following examples are illustrative of this pat-
Atlantic), and intensive studies in these regions may tern. In Hawaiian waters the mean densities of eleven
lead to the detection of still more species. atlantid species ranged from less than one to a maxi-
mum of 25 per 1000 m3, and two pterotracheids ranged
The Pacific (33 species) and Indian Oceans (32 spe- from less than one to 7 per 1000 m3 (Seapy, 1990a). In
cies) have a higher species richness than the Atlantic the California Current the carinarid Carinariajaponica
Ocean (23-25 species) (Table 1). Only one species occurred at densities of 1ess than one animal per 1000
(Atlanta ji-agilis) is Iimited to the Atlantic Ocean, m3 during most of the year, increasing during surnmer
while two species (Carinaria japonica and Atlanta months to 1-10 per 1000 m3 (Seapy, 1974). In the mid
californiensis) are restricted to the Transition Zone North Atlantic Pafort-van Iersel (1983) reported 0.2 to
fauna 1 province in the North Pacific Ocean (Seapy, 3.3 individuals per 10,000 m3 for two species of
1974; Seapy and Richter, 1993). Five atlantids Carinaria and 0.2 to 9.8 per 10,000 m3 for five species
(Atlanta echinogyra, A. gibbosa, A. meteori, A. plana, of Pterotrachidae. From the Florida Straits, Miche1 and
A. tllrriclllata) and four carinarids (Carinaria cithara, Michel (1991) found that two pterotracheid and two
C. cristata, C. galea, Pterosoma plan 11m)have Indo- carinarid species ranged from less than one to a maxi-
Pacific distributions. mum of 6 per 10,000 m3 and less than one to 3 animals
per 10,000 m3, respectively.
i -++++-
---r-
624 +
j
+- +
I
+-
+ + ++ Oce~cean t +--
+?
Heteropoda Ocean +
+ -t
775) + -¡ 1
t + I ]1 ~-
í j
+-+ +++
I
~
:?
:J
I Atlantic pacifiCllndi:~J
Table 2. Nllmbers of individllals of carinarid and pterotracheid species colleeted from obliqlle depth intervals by opening-closing MOC ESS
and RMT net tows from Profile ITI, February 1979 dllring RIV Meteor Crllise 51. COllnts of animal s were not correeted for the volllme ofwater
filtered during the tows. Dlle to poor preservation C. richardi cOllld not be clearly distingllished from c. placen/a. Since C. richardi appears to
be rare in the SOllth Atlantic (Magaldi, 1977, 1984), all specimens were considered 10 be C. placen/a.
Distribution in the South Atlantic Ocean keraudreni, Atlanta peroni, A. gaudichaudi, A. fusca,
A. lesuellri, A. inclinata, A. inflata, A. meleori, and A.
Records of heteropods from the South Atlantic Ocean helicinoides) and three Pterotracheidae (Pterotrachea
are very limited, and do not enable us to make any sta- hippocampus, P. minuta, and Firoloida desmaresti).
tements about biogeographical pattems. Although No Carinariidae were recorded, although he subse-
South Atlantic waters were sampled during the Bonite quently (Magaldi, 1984) reported a single specimen of
Expedition, Souleyet (1852a) did not give exact station Cardiapoda richardi collected by the RN Atlantis II
locations for his species. Tesch (1949) reported eight in March, 1971 at 37°48'S, 52°14'W.
species from the South Atlantic (Protatlanta souleye-
ti, Carinaria lamarcf..:i, Cardiapoda placenta (only In the numerous plankton samples taken by the RN
near the equator), C. richardi, Pterotrachea coronala, Meteor during Cruise 51 in the equatoria1 Atlantic
P. hippocampus, P. scutata, and Firoloida desmare- Ocean (South Equatorial Current) between January
sti). These records are of limited value, however, and June 1979 (unpublished), all species ofheteropods
since only a few stations were included in the South described in this paper were found (see Table 1). The
Atlantic during the Dana Expeditions. From nearsho- same collections provided data on the vertical distribu-
re waters in the eastem South Atlantic, Frontier tion and vertical migration pattems for one carinarid
(1968) named 13 species; Atlanta leslleuri, A. galldi- and four pterotracheids (Table 2). Cardiapoda placen-
chaudi, A. inclinata, A. helicinoides, A. fusca, A. tur- ta and Firoloida desmaresti were limited essentially to
'-iculata, Oxygyrus keraudreni, Protatlanta souleyeti, the upper 100 m of the water column, and the latter
Carinaria lamarcki, Cardiapoda placenta, Pterotra- species was about four times as abundant as the for-
el/ea coronata, and P. scutata. From the westem mer. Pterotrachea coronata was collected in all depth
South Atlantic Magaldi (1977) examined 107 station intervals above 250 m, but was most abundant below
samples collected by the R/V El Austral during 100 m, while P. scutata was only captured below 100
December 1966 and January 1967 off the coasts of m, as was P. hippocampus (with the exception of a
Brazil and Uruguay, between the equator and 35°S. single individual between the surface and 30 m).
He reported on nine species of Atlantidae (Oxygyrus Because these data are from a single vertical profile,
Table 1. Distriblltion of cllrrently recognized species of Heteropoda in the lhree major oceans of the world. Presence denoted by + symbols,
absence by - symbols.
626 Heteropoda
l
Species Station No. Time Number 0-250 m
Neuston o0P-ecimensj
I
-'- (males) (mixed)
Atlanta oligogyra
Protatlanta souleyeti
-l
-----r
96
108
100
116
I
L
J--06:3-?-t.
16:55
~
J 92
39 (92)
~
(39)
---l-17:º--º--¡ 1020 (80)--r------ 19_9_
~ 1
t
+-
2?--l ~
~7:~
~ ~--~- 69 (69) _. _ 35~ ,
Firóloida desmaresti
96
--l--06:3L 220 (82) 3 +- +
1 98 L12:0U 126 (126) , 21 J
Table 3. Comparison of the numbers of 2 atlantid and 3 pterotracheid species captured by surface neuston net tows and oblique ring net tows
to 250 m depth at selected stations occupied during Cruise 51 of the RJV Meteor in the central equatorial Atlantic Ocean. AJl specimens exa-
mined from lhe neuston tows were mature males, while lhose in the 0-250 m tows were a mixture of males and femaJes. Numbers in paren-
thesis refer to specimens identified beyond all doubt as males. A more cursory examination of the remaining animal s showed that they were
al so males.
they should not be used to indicate ranked abundances the specimens from the oblique tows were a mixture of
of carinarid and pterotracheid species. For example, in larval stages and males and females of different sizes.
February 1979 323 specimens of F. desmaresti were Seapy and Richter (1993) reported comparable neuston
captured (13 stations and 64 samples), which contrasts tow records for A. ealiforniensis off Southern
with the two F. desmaresti collected in June 1979 from California. Surface aggregation of animals of not only
a comparable number of stations and samp1es. one species, but also of only one sex and age points to
Examination of these samp1es for atlantids revealed a purposeful act. The digestive tracts of the animals
that about 70% were found between the surface and 50 were empty, so there is no evidence that males assem-
m, and more than 60% were between the surface and ble at the surface for feeding purposes. Although we
30 m. About 25% were caught between 50 and 100 m, know literally nothing about the sexual behavior of
and on1y 5% be10w 100 m. An exception to these heteropods, we speculate that such swarms are some-
results was Atlanta fragilis, which lives deeper than how involved in mating. Hypothetically, a female
other at1antids and appears to undergo a nocturna1 ver- swims into an all-male swarm at the surface, couples
tical migration. In 27 tows (14 during the day and 13 with a male, and drops beneath the surface to mate
at night) a total of 68 adult specimens was caught. (similar to the mating behavior seen in mosquitos; dis-
During the day 30 were captured at depths between cussed in Seapy and Richter, 1993). We have no know-
100 and 250 m and none were present in the 0-30 m ledge ofhow long males remain at the surface; swarms
samples. At night 33 specimens were present in the 0- may form intermittently or they may break up more or
30 m samples, while 5 were in the 100-200 m samples. less passively with the slow movements and mixing of
water masses of different origin and composition.
Evaluation of neuston tows during Cruise 51 of the RJV Adult male atlantids were found by Seapy (1996) to
Meteor suggests that heteropods belonging to different serve as the exclusive substratum for egg attachment by
families, genera and species form hugh swarms at the the pleustal water skater Ha/abates sericeus off
surface, consisting exclusively of mature males of com- Hawaii. The only logical explanation for this selectivi-
parable size. The numbers of individuals belonging to ty is that only adult males were available to the water
three different species (Atlanta oligogyra, Protatlanta skaters at the ocean surface. AII-male swarming beha-
souleyeti and Firoloida desmaresti) from neuston tows vior is not comparable with any other form of reported
and oblique (0-250 m), open net tows are compared in aggregation behavior (whether in heteropods or other
Table 3. For each species all of the specimens exami- pelagic taxa), since they include both males and fema-
ned from the neuston tows were mature males, while les, and could include larval stages.
Heteropoda 627
The R/V Meteor Cruise 51 data also provide an indi- rids and pterotracheids, larval shel1s are often distinc-
cation of the high densities that atlantids can achieve tive in size, shape and whorl omamentation.
in the tropical Atlantic Ocean. Al1 recognized 1arval
stages are included in these data and account (in most In Atlantidae the overall shape and surface omamenta-
cases) for 80-99% of the totals: Atlanta injlata (171 tion on the larval shel1 and the spire of the adult shell
per 100 m3), Protatlanta sOllleyeti (80 per 100 m3), A. are of great taxonomic importance (Thiriot-Quiévreux,
peroni (72 per 100 m3), Oxygyrlls keraudreni (68 per 1973, 1975; Richter, 1986, 1987, 1990, 1993; Seapy,
100 m3), A. helicinoides (67 per 100 m3), and A. oli- 1990b; Seapy and Thiriot-Quiévreux, 1994). Presence
gogyra (41 per 100 m3). These maxima shou1d only be and expression of surface omamentation, depth of the
used to suggest an approximate ranking and magnitu- suture (separating the shel1 whorls) and depth of the
de of difference in abundances. Actual abundances umbilicus can best be observed using a scanning elec-
can vary in relation to location, seasons, tow depths, tron microscope. General shape of the spire and spire
etc. It is wel1 known that the thecosome pteropods, whorls are best investigated using transmitted light.
which overlap spatially with the atlantids in the epi-
pelagic zone, are far more abundant. In the above The number of shel1 whorls comprising the spire is an
samples the highest maxima among the thecosomes important taxonomic character in Atlantidae. Unlike
were 2,041 per 100 m3 (for Limacina ¡nj/ata) and 302 most bottom-dwel1ing gastropods, however, the exact
per 100 m3 (for Creseis virgllla). Even a group ofneri- number of spire whorls can be difficult to determine
tic origin, the pelagic larvae of Architectonicidae, beca use the transition between the larval shel1 (proto-
showed a higher frequency (395 per 100 m3). conch) and adult shel1 (teleoconch) is often difficult to
determine. Nonetheless, even an approximation ofthe
number of spire whorls is infinitely more reliable for
Taxonomy taxonomic purposes than the total number of shel1
whorls, since the latter increases with shel1 growth.
Characters of taxonomic importance Here we have approximated the number of spire
whorls by determining the location where whorl width
Four morphological characters (shel1, eyes, opercu- begins to increase rapidly (Fig. 1), to accomodate the
lum and radula) are of great importance in the identi- increased size of the post-metamorphic animal.
fication ofheteropods, particularly Atlantidae, and are
discussed below. In Carinariidae the protoconch is compact, thick-wal1ed
and helicoid. Unlike the atlantids, the border between
..
Shell the protoconch and teleoconch is quite distinct. Larval
shel1s of carinarids are transparent, but some species
The shel1s of heteropods are calcareous (composed of have a yellow, brown or pink coloration, which is visi-
aragonite) with the exception of the kee1 in ble even on the protoconchs at the apex of the adult
Protatlanta and the keel and shel1 in Oxygyrus (com- shells. The teleoconch is transparent and colorless, with
posed of a proteinaceous, cat1ilaginous-like material, thin and brittle, undu1ating wal1s. The adult shel1ranges
conchiolin). Because the adult shel1 (teleoconch) of in size from a large, cap-shaped or f1attened structure
atlantids and carinarids is produced directly from the covering the visceral nucleus (in Carinaria and
larval shel1 (the protoconch), the larval shel1 is present Pterosoma, respectively) to a greatly-reduced shel1,
as the inner, helical part of the adult shel1 in atlantids only slightly larger than the larval shel1, which is retai-
and at the apex of the adult shel1 in carinarids. The ned at the apex ofthe visceral nucleus (in Cardiapoda).
first shel1 material secreted forms the embryonic shel1
(referred to as protoconch I), which is dome-shaped In Pterotracheidae only a larva1 shel1 develops, and it is
and is produced by the mantle of the developing thrown off during metamorphosis. Three Pterotrachea
embryo while it is stil1 inside the egg capsule. 1arvae have been distinguished from the Mediterranean
Although the embryonic shel1 is uncalcified in some (Krohn, 1860; Richter, 1968; Thiriot-Quiévreux,
groups of gastropods, it is always calcified in the 1973). Here we give short descriptions of the shel1s of
heteropods. The remaining portion is the true larva1 these species. 1) Shel1 transparent and smooth, with a
shell, referred to as protoconch II. In atlantids, carina- perfectly round diameter and strong wal1s. Whorls
628 Heteropoda
t t
At/anta fusca At/anta inf/ata Atlanta /esueuri
••
t At/anta gaudichaudi t
At/anta peroni Atlanta tokiokai
0.5 mlTl
Fig.1. Determination ofnumber ofwhorls eomprising lhe shell spire in Allanlidae. Spire skelehes made al a 90° angle lo lhe plane oflhe spire.
Arrows indieale approximale areas where whorl expansion begins. Speeies of Al/al/la wcre selected lhat were represenlalive of six of lhe allan-
lid speeies grollps (see Table 4). Modified from Seapy (1990b).
(about 1 and 112) enlarge slowly, and the last half deta- Eyes
ches fram the first, leaving a wide gap between the first
and second whorls. 2) SheIl extremely thin, so that it The eyes are enclosed in a gelatinous capsule and con-
coIlapses when dried. Whorls (also about 1 and 1/2) sist of a distal spherical lens, intermediate region that
enlarge rapidly and are not separated by a deep, wide is partiaIly to completely pigmented, and a praximal,
suture. Entire shell wall undulated parallel to growth strip-like retina. The pigmented layers shield the
lines. 3) SheIl f1at and helicoid, consisting of about two space between the lens and retina from extraneous
whods that enlarge rapidly. The walls are thinner than light. The eyes of different species can vary in shape
in the first species, but are stiIl strong and completely (from triangular to cuboidal to rectangular) and the
caIcified. pigment layers can differ in coverage and disposition
Heteropoda 629
transverse slit in distal
0.2 mm
Type b e
I 1
Type a Type
Fig. 2. The three types of eye morphology (a, b and e) recognized among the species of Atlantidae (type e eye from A. helicinoides). From
Seapy (1990b).
Type a Type b Type e

(macro-ol igogyre) (micro-ol igogyre) (monogyre)
Fig. 3. The three types of opercular m rphology recognized in A/lan/a and Pr%/Ion/a. From Seapy (1990b).
in the eye. In the Atlantidae, three basic eye types Operculum

(Fig. 2) were characterized by Richter (1961), which
he subsequently (1974) refered to as types a, b and c. The operculum of all Heteropoda is cartilaginous and
Distribution of the eye types among the species of flexible. It is present in all larval stages. In Atlantidae
atlantids is surnmarized in Table 4. the larval operculum grows spirally, but after meta-
morphosis it grows in a straight line. Thus, the nuc1eus
In Pterotracheidae eye shape is effected by the width (or gyre) is situated excentrically in the narrower
of the retinal base, resulting in eyes that range in upper portion of the operculum. Richter (1961) distin-
shape from rectangular to triangular when viewed guished three different types of opercula in the atlan-
dorsally. Bonnevie (1920) and subsequent workers tids, which he characterized as macro-oligogyre,
ha ve emphasized the importance of eye shape in dis- micro-oligogyre and monogyre; he subsequently
tinguishing between the species of Pterotrachea. For (1974) designated these as types a, b and c, respecti-
the most part, eye morphology has not been used as a vely (Fig. 3). Only Oxygyrus does not tit into one of
taxonomic character in Carinariidae. these three types. Its operculum is broadly triangular
630 Heteropoda
- - eIe ~Operculum
3-1/4 **
I ?-- ~I
L
,>---- *
--
1--
ni
1/4
--"
4-1/2
~pecies group
nta gaudichaudi species
-+---
11
3-1/2
-'-
-A--tokiokai
~- 5-3/4
~
group L ~ t=-+-J
r-t-
b+-eatype
-LT1tb
1
t keraudre~
I
b
Eye
type ~- a ab --'--
aeRadula
t-~~,__ b_ I 4-1/2 I L_L--
--+-- b 4-1/2
1
346
3-1/2
4-1/4
3-3/4
2-1/2
3-1/2
Spire
5-1/2
1-2-1/2 -b-
t
t-
11
11 abe,
b
11
r- ¡-?-
, 11
«,>up
_
Oxxgyrus
A peroni
gibbosa
b b f ~
11
b_t- 1
C-=l=_1I
Table 4. Occurrence offour taxonomic characters among the currently recognized species of Atlantidae, arranged into species groups (see text).
See Fig. 3 for characterization of operculum types. *: Involute spire negates whorl count; **: Operculum broadly triangular, not comparable
with those of Prolal/anla and Al/al/la.
Heteropoda 631
(almost trapezoidal), lacks a spiral part and shows rows may be of taxonomic value. The radula types of
growth lines somewhat like those on the shell of atlantid species are summarized in Table 4. All
Mytilus (Bonnevie, 1920). In three species of Indo- Carinariidae and Pterotracheidae have type II radulae.
Pacific atlantids (Atlanta turrieulata, A. eehinogyra
and A. plana) the nucleus is decorated by spines or In each tooth row the central, rachidian tooth is flanked
hooks, arranged in a specific pattem (Tokioka, 1961; on either side by a lateral and two marginal teeth (Fig.
Richter, 1972). Opercular types for the species of 4A). The outward spreading of the swivel-mounted
Atlantidae are surnmarized in Table 4. marginal teeth mentioned in the Introduction can be
seen in this photograph as those teeth that flare outward
In Carinariidae and Pterotracheidae the operculum is at the tuming point of the radula. The shapes ofthe cen-
present only in the larvae, since it is thrown off during tral, lateral and marginal teeth are taxonomically impor-
metamorphosis. The carinarid operculum consists of tant. At the family level the central tooth differs notice-
two layers, connected only in the central parto The ably. Among the Atlantidae this tooth has a single
outer layer is larger than the inner layer and is larger median cusp in Atlanta (Fig. 4B) andProtatlanta, but is
than the shell aperture. When the larva withdraws into tricuspid in Oxygyrus. In Carinariidae three long and
its shell, the inner layer enters the aperture while the prominent cusps are present (Fig. 4C), while in
fringe of the outer layer bends outward, forming a per- Pterotracheidae a well-developed median cusp is flan-
fect closure. The same type of operculum is present in ked by numerous short cusps that decrease in size late-
the pterotracheid Firoloida desmaresti, a species with rally (Fig. 4D).
a larva 1 shell quite similar to that of carinarids. All
other species of pterotracheids (in the genus
Pterotraehea) have an extremely thin operculum, Heteropoda species from the South Atlantic
which looks from the outside like a set of stacked dis-
hes of different sizes (Richter, 1968). The following scheme outlines the classification ofthe
Heteropoda; species denoted with bo1d underlined
Radu1a characters are those recorded in the South Atlantic and
treated in detail in the following section.
Radu1a shape and radu1ar tooth morphology were rejec-
ted as taxonomic characters by Buchmann (1924) and Superfamily Heteropoda
Tesch (1949) as a result of their 1ack of understanding Family Atlantidae
of radu1a morphogenesis. Shapes of the radular teeth Oxygyrlls keraudreni species group
change dramatically during ontogenesis and depend on Genus Protatlanta
the position on the radula ofthe tooth row investigated. Protatlanta souleveti
This morphogenesis ofteeth continues in Atlantidae for Genus Oxygyrlls
a great part of radula growth, whereas in Carinariidae Oxvgyrus keraudreni
and Pterotracheidae (as well as in many other proso- Atlanta fusea species group
branch gastropods) it is restricted to the late larval and Atlanta (usca
first postlarva1 stages. For radula growth and tooth Atlanta turriclllata
morphogenesis and function see Richter (1961, 1963). Atlanta injlata species group
Atlama inflata
Here we distinguish two types of radulae. In type 1radu- Atlanta helicinoides
lae the teeth that are produced first are never cast off Atlanta califomiensis
from the anterior end of the radula as they outwear their Atlanta lesuellri species group
use; the number oftooth rows increases continuously as Atlanta lesueuri
the animal grows and all morphogenetic stages of teeth Atlama olir:ogyra
are present in mature animals (i.e., unlimited number of Atlanta peroni species group
tooth rows). In type II radulae teeth are cast offthe ante- Atlanta peroni
rior end of the radula as soon as the radula has reached Atlanta (ragilis
its specific number of tooth rows (i.e., limited number Atlanta rosea
of tooth rows). In the latter case the number of tooth Atlanta frontieri
632 Heteropoda
Fig.4. Scanning electron micrographs of heleropod radulae. A: low magnificalion view al turning poinl of Carinaria lalllarcki radula, B-O:
high magnification views of mature seclions of Atlanta lesuellri, C. lalllarcki, and Pterotrachea hippocalllplIs radulae, respectively. Scale bars:
A = 200 11m,B = 10 11m,e and o = 20 11m.Richler, original.
Atlanta gaudichaudi species group Carillaria cithara

Atlanta ~audichaudi Carinaria eristata
Atlanta plana Carillaria japoniea
Atlanta eehinogyra Genus Cardiapoda
Atlanta inclinata species group Cardiapoda placenta
Atlanta inciinata Cardiapoda riehardi
Atlanta tokiokai Genus Pterosoma
Atlanta gibbosa species group Pterosoma plan 11m
Atlanta ~ibbosa Family Pterotracheidae
Atlanta meteori Genus Firoloida
Unassigned species Firoloida desmaresti
Atlanta peresi Genus Pterotraehea
Family Carinariidae Pterotraehea eoronata
Genus Carinaria Pterotracl,ea seutata
Carinaria lamareki Pterotraehea hippoeamp"s
Carinaria ehallell~eri Pterotraehea milluta
Heteropoda 633
Representatives of all three families of Heteropoda Genus Protutlallta Tesch, 1908

occur in the South Atlantic, and include 14-16 species Protatlullta sOllleyet; (Smith, 1888) (Fig. 5A).
of Atlantidae, four Carinariidae and five Pterotrachei- Adult shell calcareous, with strongly truncated
dae. Descriptions of these families and species are keel composed of conchiolin. Shell small (less
given below. than 2 mm) and helicoid. Spire slightly elevated
and composed of about 3 whorls. Radula large,
Family Atlantidae Wiegmann and Ruthe, 1832 triangular. Central tooth monocuspid, with a large,
Members of this family are small (Iess than 10 mm) low basal plate. Lateral teeth monocuspid with a
and possess a dextrally-coiled shell into which the low process on its inner side. Marginal teeth
body can retract and close offwith a chitinous opercu- monocuspid. Eyes type a, and operculum type a.
lum, which is attached to the dorsal side of the foot. A In the Atlantic Ocean shells with two different
single gill and osphradium are positioned dorsally wit- spire morphologies are encountered, often in the
hin the mantle cavity. A pair of well-developed tenta- same plankton tow samples. In the first morph the
cles arise ahead of the eyes. In all atlantids a large suc- spire whorls are smooth; in the second a low spi-
ker is located on the posteroventral margin of the fin. ral ridge is present, which appears just after the
It is present in both sexes and serves to hold prey tight- beginning of the second whorl, is centered on the
Iy while the strong radula tears off and ingests pieces whorl, and ends after about three-fourths of a tum
of prey tissue (Richter, 1968; Thiriot-Quiévreux, in tile third whorl. A less conspicuous, secondary
1969). The radula is of two types (I and II), and can spiral ridge is present inside of the primary ridge
show distinct sexual dimorphism. Since males and on the second whorl. In sediment trap samples
females feed on the same prey, we know nothing about from the Walvis Ridge in the eastem South
a possible functional basis for this dimorphism. Atlantic, specimens of Protatlanta souleyeti with
a much stronger spiral sculpture prevail. This
The identification of most heteropods, but especially sculpture matches precisely that described by Issel
of species in the genus Atlanta, can be very difficult. (1911) for his Protatlanta sculpta. Since the spire
To facilitate identification, we have distinguished sculpture is as variable in P. sOllleyeti as in a num-
eight species groups in the Atlantidae, each of which ber of other atlantids, we conclude that Tesch
is named after a representative species (Table 4). (1949) was correct in not accepting P. sClIlpta as a
(Similarly, van der Spoel, in his 1976 text, arranged valid species.
the various species of Atlanta that he recognized at the
time into a series of six species groups.) It is our hope Genus Oxygyrus Benson, 1835
that even the inexperienced student will be able to dis- Oxygyrlls kerulldreni (Lesueur, 1817) (Fig. 5B).
tinguish between these species groups. This accom- Among the largest species of atlantids, attaining a
plished, the probability of identifying the specimens diameter of 10 mm. Entire adult shell composed of
at hand to the species level should be greatly incre- conchiolin. With growth, adult shell overgrows
ased. Table 4 includes the species of atlantids current- calcareous larval shell and keel eventually SUf-
Iy recognized from the worlds oceans, arranged by rounds the last shell whorl. Truncate anterior edge
species groups. Those species that we consider to be of keel ends at shell aperture. Protoconch with
present in the South Atlantic are characterized below. prominent, evenly-spaced, zigzag spiral ridges
Papers containing descriptions (more or less comple- that cover the surface. Shell spire involute (two
te) of all these species are found in the list of referen- whorls of larval shell helicoid). Operculum trian-
ceso For synonymy lists see van der Spoel (1976). gular to trapezoidal and lacks a spiral parto Only
one-halfofthe shell opening (aperture) covered by
Oxygyrlls kerUlldrell; species group operculum when animal retracts into shell. Type c
eyes, with large pigmented base, which reaches
Shell with a glass-like, transparent keel, composed of beyond the distal part of the lens. Radula very
conchiolin. Leading edge of keel truncate, sloping large and triangular in shape. Median tooth tricus-
abruptly and ending at the shell opening. Although all pid; in juvenile specimens with three pointed
or part of the adult shell is constructed of conchiolin, cusps, in adult specimens with one central cusp
the larval shell is calcareous. Radula type 1. flanked by blunt flattened lobes. Lateral teeth with
634 Heteropoda
Fig.5. Scanning electron micrographs ofthe shells of A: Prolallallla sOllleyeli and B: Oxygyrus keralldrelli. Shells are viewed from right side
at 90° and 60° to the shell plane. Scale bars are 0.5 mm for low magnifications and 0.1 mm for high magnification. From Seapy, 1990b (except
for P. sOllleyeli spires which are from Richter, original).
a strong process on the inner side, bearing a poin- type a, operculum type a. Radula type 1, small and rib-
ted accessory cusp. Marginal teeth monocuspid. bon-shaped. Central tooth (like in all Atlanta species)
monocuspid, with one posterio-lateral process on each
All remaining species of Atlantidae are members of side. Lateral teeth with a strong rectangular process on
the genus Atlanta Lesueur, 1817 in which the coiled the inner side (like in all Atlanta species) and bicuspid,
adult shell is produced directly from the calcareous even in adult specimens. Marginal teeth monocuspid.
larval shell. Both the adult shell and keel are calcare- • Atlanta fusea Souleyet, 1852 (Fig. 6E, lA). Keel
ous. Keel height varies from low to high, and the lea- high and colorless, and inserts between fifth and
ding edge is not trunca te, but slopes towards the shell sixth shell whorls in animals larger than about 1.5
mouth and ends in a narrow fissure. Those species mm. Spire elevated and cone-shaped, of about 4
whose distributions are either limited to the Pacific whorls, and with a complex pattern of ornamenta-
Ocean or the Indo-Pacific region are not described, tion. Spire coloration ranges from yellowish brown
but are listed under the species group in which they to dark brown. Prominent spiral ridge located
belong (and in the outline c1assification above). along the outer margin of spire whorls. Spire orna-
mentation ends on the protoconch and is replaced
Atlanta fusea species group by rows of small punctae on the teleoconch.
Shell small (less than 2.0 mm maximal diameter) and • Atlallta turriculata d'Orbigny, 1836. This species
pigmented (yellowish-brown to brown, or reddish has been reported from the eastern South Atlantic
brown). Keel tall and rounded. Spire strongly sculptu- by Frontier (1968) and the Mediterranean by
red and shaped either as a tall cone or a turret. Eyes Thiriot-Quiévreux (1973) and Vatova (1974).
Heteropoda 635
Fig.6. Seanning eleetron mierographs ofadult shells, larval (or juvenile) shells, and radulae of Atlanta speeies. A: Atlanta helieinoides, B: A.
inflata, C: A. lesllellri, O: A. oligogyra, and E: A. jilsea. Mierographs of righl side of adull shell al 90° and abolll 60° lO lhe shell plane; lar-
valljuvenile shells orienled faeing aperture and pholographed wilh transmilled lighl. Radula SEM mierographs from animals of idenlieal size.
Riehler, original.
However, Tesch (1949) did not record A. fllrriclIla- At/anta inflata species group
fa from Atlantic waters or from the Mediterranean, Shell small (diameter less than 2.0 mm) and inflated
and Richter (1968) did not find this species in the (width about 40% of shell diameter). Spire low, with
central Mediterranean. Thus, it seems probable that shallow sutures and surface sculpture ranging from
the records mentioned here refer to juvenile or aber- none (smooth) to low and more or les s distinct, even-
rantly-shaped A. /lIsca; a species with world-wide Iy-spaced spiral ridges. Inner walls of spire gradually
distribution. decalcified after metamorphosis so that only a thin
636 Heteropoda
cartilaginous membrane separates the whorls. (Note with an intact keel, the keel is very tall and nearly
that this membrane is visible using transmitted light or truncate along its leading edge. No pigmentation at
when the spire is broken). Operculum type c, radula keel base. Eyes type b, with extremely large lenses
type I (up to 120 tooth rows observed). in adults. Operculum thin. Radula large (largest in
• Atlanta inflata Souleyet, 1852 (Fig. 6B, lB). the genus Atlanta) and triangular, with monocuspid
Shell very small (maximal diameter = 1.5 mm) lateral teeth. Due to the thinness of the shell, the
and flat (body whorl flat-oval in cross section), eyes and buccal mass with the contained radula can
with a high keel. Spire of about 4 and 1/2 whorls, be clearly seen through the transparent shell in well
bearing low spiral ridges that are highly variable, preserved or freshly collected specimens.
but generally more prominent than in animals • Atlanta oligogyra Tesch, 1908 (Fig. 6D). Shell
from the Mediterranean Sea or the Indian Ocean moderately small (less than 3 mm) and transpa-
(Richter, 1968, 1987). Pigmentation varies from rent. Shell walls relatively strong and thick, with a
colorless to faintly yellowish. Inner surface of faint yellowish or pink hue. Keel moderately ele-
shell at aperture may have a brownish hue. Eyes vated and rounded, without a truncate leading
Type a. Radula relatively large, with monocuspid edge. Keel base generally brownish. Shell surface
lateral teeth and strong sexual dimorphism (larger smooth. Spire somewhat elevated and rounded,
and broader, with large lateral and marginal teeth with shallow sutures. Eyes type a, with lenses of
in males; more narrow and ribbon-like in females). average size (distinctly smaller than in adult A.
This species has been confused with the following lesllellri of comparable size). Radula small, rib-
species in the past, but the differences between the bon-like. Lateral teeth monocuspid or with a very
two are clear (Richter, 1987). small accessory cusp. Although difficult, young
• Atlanta helicinoides Souleyet, 1852 (Fig. 6A). specimens of A. oligogyra can be distinguished
Shell small (les s than 2.0 mm), with the outer from young A. lesuelll'i by the slightly lower and
body whorl notably more inflated (almost circular rounded spire, with a shallow suture. Confusion
in cross section) than the previous species. Keel about the validity of this species arose after Tesch
relatively low, thin and seemingly less strongly (1949) combined it with A. lesllelll'i, thus rever-
calcified than in A. injlata. Spire of about 4 and sing his previous position (1906) when he recog-
1/2 whorls, with evenly-spaced spiral sculpture nized the two species as distinct. Richter (1974,
that is variable in expression but, as a rule, is coar- 1986) showed that Tesch was correct in his 1906
ser and more prominent than in A. injlata. paper and that A. oligogyra is a valid species.
Coloration constantly yellowish. Eyes type c (the
only species in the genus Atlanta with this eye Atlanta peroni species group
type). Radula very similar to that of A. inflata, Shells large (to 10 mm diameter) and flat, with
with a similar sexual dimorphism. smooth surfaces and a small spire. Colorless or faint-
• Atlanta californiensis Seapy and Richter, 1993 ly tinted in pink or yellow. Eyes type b, operculum
(not recorded in the South Atlantic). type b. Type 11 radula, large but with a low growth
angle. Number of tooth rows limited (up to 50-70). In
Atlanta lesllellri species group full-grown specimens lateral teeth with a small but
Shell small to medium size (less than 4-6 mm), with distinct accessory cusp on the posterior side close to
an extremely small spire of about 2 and 1/2 whorls. the main cusp. Considerable confusion has surroun-
After metamorphosis whorl size increases rapidly, ded the species comprising this group. Subsequent to
and shells of the largest adults consist of only 3 and Tesch (1949) and prior to Richter (1993) only a sing-
1/2 whorls. Operculum type b. Radula type 1; number le species (A. peroni) was recognized. The latter study
of tooth rows not limited (> l 00 observed). recognized four va lid species, resulting in the re sur-
• Atlanta lesuellri Souleyet, 1852 (Fig. 6C, 1C). rection of A. ros ea (discussed below) and the descrip-
Shell entirely transparent and colorless, of mode- tion of two new species; A. frontiel'i, represented in
rately large size (up to 6 mm), with thin, brittle the Indian Ocean by specimens previously identified
walls. Shell surface smooth. Spire somewhat eleva- by Richter (1974) as A. peroni, and A. fragilis, which
ted, with prominent and deep sutures. In specimens appears to be limited to the Atlantic Ocean.
Heteropoda 637
Fig.7. Seanning eleetron micrographs ofadult shells, larval (or juvenile) shells, and radulae of Atlanta speeies. A: A.ji-agilis, B: A. peroni, C: A.
rosea, D: A. gaudichaudi, E: A. tokiokai, F: A. inclillata. Mierographs of righl side of adull shell al 90° and 60° lO lhe shell plane; larvaVjuvenile
shells orienled faeing aperture and pholographed wilh transmilled light. Radula SEM mierographs from animals of idenlical size. Riehler, original.
638 Heteropoda
• At/anta peroni Lesueur, 1817 (Fig. 7B, 1D). Suture separating first 2 and 1/2 whorls extremely
Among the species in the genus At/anta, this shallow, so that whorls are difficult to distinguish.
attains the largest size (to 10 rnm shell diameter). It These spire features, visible best using transmitted
is also the type species in the genus. Shell flat, with light on larval shells or the isolated spire, are so
a small, 10w spire of about 3 and 1/2 whorls. Full- characteristic that they suffice to distinguish this
grown shells with up to 6 whorls. Keel moderately species. However, it often requires the preparation
elevated, rounded in profile, with a brown base in (and partial destruction) ofthe shel!' The larvae of
larger specimens. Shell walls strong, relatively this species can be easily distinguished from those
thick. Shells are colorless, except in larger speci- of other species in this group by the bright pink
mens that are faintly yellowish. Eyes relatively tinge to their shells.
smal!. Radula large, with high lateral teeth; number Based on our current information, this species
of tooth rows limited (to about 60). Distinctive appears to be extremely rare but is distributed cir-
sexual dimorphism in the radula. In adult males cumglobally. In 1908 Tesch united A. rosea with
marginal teeth are sharply-bent hooks, much shor- A. perolli, probably because the holotypes in the
ter than the respective lateral teeth. In females mar- Musée d'Histoire Naturelle de Paris were already
ginal teeth are longer and less sharply curved. badly damaged (shells dissolved) when he had the
This species has been confused with other species opportunity to study them. Comparing "aberrant"
in the past due to the absence of distinctive shell shells of A. peroni from the Indian Ocean and the
characters; e.g., comparability of shell shape, tropical Atlantic Ocean with the type shells of
absence of surface sculpture, and spires not notably Souleyet, preserved dry in the British Museum,
elevated or inclined (Richter, 1993). These pro- London, Richter (1993) found that they belonged
blems are understandable when, as is generally the to this very species.
case, one has only juveniles and young adults to • At/anta frontieri Richter, 1993 (not recorded in
work with. Species having shells similar in general the South Atlantic).
appearance to A. peroni include A. jragilis, A.
rosea and A. gaudichalldi in the Atlantic, and A. Atlallta galldichalldi species group
jrolltieri and A. plana in the Indo-Pacific. Although Shell small to intermediate size (maximal diameter
Atlanta peroni is quite cornmon in the Atlantic, it 2.5-5 mm), with small, but distinctly helicoid spire.
is rare in the Indian Ocean, where the related spe- Radula type 1, with unlimited numbers of tooth rows
cies A. jronlieri is abundant (Richter, 1993). and monocuspid lateral teeth. These radular charac-
• Atlanta fragilis Richter, 1993 (Fig. 7A). Shell ters distinguish this species group from the A. peroni
large (up to 9 mm diameter), transparent and com- species group, in which the number of tooth rows is
pletely colorless, smooth, thin-walled and fragile limited and the lateral teeth are bicuspid.
(e.g., in the fifth whorl, the shell thickness is about • At/anta galldichalldi Souleyet, 1852 (Fig. 7D,
10 ¡.un- compared with about 25 flm in A. peroni). 1E). Shell moderately small (maximal diameter
Keel high; higher and deeper penetrating between about 3 rnm), with smooth surface, and a small,
the last whorls than in A. peroni. Keel base not low conical spire of about 3 and 1/4 whorls.
pigmented. Spire of about 3 and 1/2 whorls. Eye Whorls of teleoconch much flatter than in the A.
lenses very large in adult specimens. Radula large, peroni species group. Shell colorless, but keel
number of tooth rows limited (to about 70). generally with a brown base. Eyes type b, opercu-
Radula shows sexual dimorphism (like A. peroni). lum type b. This species differs from the other two
Larvae of A. jragilis and A. peroni cannot be dis- in the group in that the shell spire lacks spiral
tinguished by shell shape. If it has not been over- sculpture and the operculum lacks spirally-arran-
looked in the Pacific or Indian Oceans, A.jragilis ged spines or hooks (Richter, 1972). The shell
appears to be restricted to the Atlantic Ocean, characteristics given above may seem insufficient
where it is rare by comparison with A. peroni. to enable one to distinguish all but fully-grown
• Atlanta rosea Souleyet, 1852 (Fig. 7C). Shell of specimens of this species from A. peroni or A. jra-
moderate size (to about 5 mm) and flato Coloration gilis. However, comparison of the size and shape
faintly yellowish. Spire almost globular, of about ofthe spires enables one to identify A. galldichall-
3 and 1/2 whorls, and with a smooth surface. di, even ifthe radulae are not compared.
Heteropoda 639
• Atlallta plalla Richter, 1972 (not recorded in the tubercula arranged in distinct spiral lines, which
South Atlantic). occasionally are elevated sufficiently to almost
• Atlallta echillogyra Richter, 1972 (not recorded in form spiral ridges. Radula small and ribbon-like
the South Atlantic). (growth angle about 9°). Lateral tooth with an
accessory cusp.
Atlallta illclillata species group It is clear that the two species in this group are clo-
The two species comprising this group share a series sely related, twin species. One could debate
of characters that are so striking that it seems extre- whether such species should be taken as subspe-
mely difficult to distinguish them. They are: 1) large, cies, but this would not resolve the underlying
beehive-shaped, inclined spire; 2) decalcified inner taxonomic problems. On the other hand, it is quite
walls of the spire (like in the A. inflata group); 3) clear that the two species in the A. gibbosa species
internal wall structure ofradially-arranged lines (visi- group, both of which have calcified inner spire
ble only using transmitted light); 4) surface sculpture whorls and a radula with a very different shape,
of small tubercula (or punctae) on the spire and, to a cannot belong in this group.
lesser degree, on the body whorl; 5) type b eyes; and
6) type c operculum. This species group and the next Atlanta gibbosa species group
one include four species with large and inclined spi- Shells of medium size (up to about 4 mm diameter),
res. Van der Spoel (1976) grouped all species with completely transparent and colorless, with thin walls
inclined spires in an A. inclinata species group. and a high keel that penetrates between the last
However, this distinctive character has concealed whorls. Eyes type b, with lenses that are conspicuous-
basic, but less striking, differences between the spe- ly large (in adult specimens), thus suggesting that
cies (Richter, 1990). Comparisons of these four spe- these species prefer deeper daytime depths (Richter,
cies indicate that an inclined spire has developed 1974). Operculum type b. Type 11 radula, with low
along two quite different lines in the genus At/anta, growth angle. Lateral teeth straight (not curved as in
and the species in each lineage appear to be more clo- most species) and with strong accessory cusps. The
sely related to species with straight spires than they two species in this group appear to be found exclusi-
are with the species in the other lineage. vely in the Indo-Pacific region, although A. gibbosa
• Atlallta illclillata Souleyet, 1852 (Fig. 7F). Shell was reported by Souleyet (1852b) from an Atlantic
large (to 6-7 mm), colorless or light yellowish. plankton sample. Thus, we have included the
Spire of 4 and 1/2 whorls, with shallow suture. On characterization of this species group and A. gibbosa
the underside of the spire, a low and indistinct below.
ridge marks the line where, on the following • Atlallta gibbosa Souleyet, 1852. Shell spire of
whorl, the keel develops. Area between this ridge about 6 whorls, with up to 7 and 1/2 whorls in full-
and the umbilicus rounded like other whorls. Tiny grown adult shell. These numbers of spire and
tubercula scattered on the spire surface, and rarely total shell whorls are greater than in any other spe-
on the first whorl of the teleoconch; sometimes cies of atlantid. Spire in lateral view with a broad
fOlming irregular, spirallines. Radula large, with a base and a pointed apex. Spire least strongly incli-
growth angle of about 16°. Number of tooth rows ned among species in this and the previous group.
limited to about 60. Lateral teeth monocuspid. Shell surface smooth, and umbilicus wide. Radula
• Atlallta tokiokai van der Spoel and Troost, 1972 shows a distinct sexual dimorphism, in principIe
(Fig. 7E, 1F). Shell moderately small (3 mm maxi- resembling that described for A. peroni and A. fra-
mal diameter), light yellow-brown color, and gilis, but still more conspicuous. Marginal teeth of
brown inside the outer lip. Spire of 5 and 1/2 adult males are sharply-curved, short hooks;
whorls, suture very shallow and difficult to distin- leaving a characteristic gap between the marginal
guish in side view. Keel tall and rounded. Ridge teeth and the cusps of the lateral teeth. Central
on underside of spire strongly developed; area tooth with a broad, low basal plate and a long,
between this and umbilicus flattened. Internal slender cusp.
structure like that described for A. inclinata, but Atlallta meteori Richter, 1972 (not recorded in the
hardly visible because of the much more strongly South Atlantic).
developed surface sculpture. Spire whorls with
640 Heteropoda
Unassigned species fin sucker is present in both sexes, but is much smaller
• Atlanta peresi Frontier, 1966. Shell moderately than in the Atlantidae. The location ofthe sucker on the
small (type specimen = 2.7 mm diameter), flat, posteroventral margin of the foot and its remoteness
colorless, and with a smooth surface. Spire small, from the mouth make its use in feeding improbable. It
of 3 and 1/2 whorls. Keel high, penetrating deep may be used during mating to hold the male and female
between the whorls in large individuals. Diameter together, however such behavior has not been obser-
of teleoconch whorls increase almost continuously ved. The radula in all Carinariidae is broadly triangular
and quite slowly, beginning in the third whorl; a and has a relatively low and limited number (34 to 44)
feature not known from other atlantids. Operculum of tooth rows (type II radula). The central tooth has a
type b. Radula and eye types unknown. broad, low base with well-developed posterio-Iateral
Because Thiriot-Quiévreux (1968, 1970, 1973) and processes and three long cusps (Fig. 4C). The lateral
Vatova (1974) reported A. peresi from the teeth have a smaller, curved cusp on their inner acces-
Mediterranean, we have inc1uded it here. Frontier sory plate (in addition to the main cusp). This cusp is
(1966) described this species from the westem more conspicuous in the genus Carinaria than in the
Indian Ocean. However, no specimens from the other two genera. The marginal teeth are monocuspid
1964/1965 Meteor Expedition in the central nor- (Fig. 4A). According to Tesch (1949) the radulae show
them Indian Ocean examined by Richter (1974) few differences among the different genera and species
could be identified as this species. In addition, we of Carinariidae, whereas Vayssiere (1904) and
are not aware of any records indicating its presence Bonnevie (1920) obviously overvalued its taxonomic
in the Pacific Ocean. Van der Spoel (1976) placed value in the family. (For morphogenesis of radular
this species in the Atlanta leslIellri group because teeth in the Carinariidae see Richter, 1968).
ofthe low number ofwhorls that make up the spire.
Unlike A. leslIeuri and A. oligogyra, however, the Genus Carillaria Lamarck, 1801
shell whorls ofthe teleoconch enlarge quite slowly The visceral nuc1eus is stalked and located dorsally
and the keel inserts between the last shell whorls above or immediately posterior to the ventral swim-
(as in most species in the A. peroni species group). ming fin. The nuc1eus is covered by a cap-shaped
Since A. peresi shares shell characters of both the shell. Following metamorphosis the teleoconch grows
A. lesueuri and A. peroni species groups, we have directly from the protoconch, which is retained at the
not placed it in either group. Not having specimens shell apex. In contrast with the Atlantidae, there is a
of A. peresi available for examination, we have had c1ear junction between the protoconch and teleoconch.
to rely on the species description ofFrontier, which A pair of tentac1es are located ahead of the eyes (like
is incomplete. Frontier did not name any type mate- in Atlantidae), but they can be ofunequal size, with the
rial or a type specimen, and the reports of this spe- right tentac1e often reduced or vestigial. Eye morpho-
cies by Thiriot-Quiévreux and Vatova from the logy is a taxonomic character only in distinguishing C.
Mediterranean Sea did not give a more thorough cithara (cylindrical shape) from the other species of
description. Carinaria (triangular shape) (Okutani, 1961). The tail
has a dorsal crest, developed to varied degrees.
Family Carinariidae Fischer, 1883 Carillaria lamarcki Péron and Lesueur, 1810 (Fig.
This family differs most conspicuously from the 8). Body size up to 220 mm (in the Mediterranean
Atlantidae by the size and shape ofthe body. The body Sea; Tesch, 1949), although specimens collected
is basically cylindrical and is greatly enlarged and elon- by Pafort-van Iersel (1983) from the mid North
gated, covered by a cutis of varied thickness. The body Atlantic attained a maximal length of only 114
is divided into three regions; proboscis, trunk and tail. mm. Shell depressed and broadly triangular in
The proboscis is more or less distinct from the head lateral view; height about one-half of basal length.
region, located at the anterior end of the trunk. A stal- Shell height to length ratio is lowest among the six
ked visceral nuc1eus is located at the posterior end of species of Carinaria, except for C. challengeri.
the trunk, and is followed by a tail. Although a shell is which is comparable. Trunk covered by thick cutis,
present, it is reduced to either a cap-like structure over with many tuberc1es on its surface. Thickness of
the visceral nuc1eus or a microscopic remnant of the cutis and size of tuberc1es increases with age. Two
larval shell, located dorsally on the visceral nuc1eus. A tentac1es present; right smaller than left.
Heteropoda 641
Fig.8. Carillaria lamarcki. A: female from right side, in-situ (courtesy of R. Gilmer). B: visceral nllclells and shell (Seapy, original). Scale
bars, A: 10 mm, B: 5 rnrn.
Carillaria challengeri Bonnevie, 1920 (Fig. 9). Carillaria cristata (Linnaeus, 1766) (not recorded
This species is much smaller than C. lamarcki, in the South Atlantic).
attaining a maximal size of 40 mm (Bonnevie, Carillaria japollica Okutani, 1955 (not recorded
1920). Shell depressed; similar to that of C. in the South Atlantic).
lamarcki. Pair of darkly-pigmented, hemispherical
structures (termed claspers by Bonnevie, 1920) Genus Cardiapoda d'Orbigny, 1836
located posteriorly on either side of the ventral The shell is greatly reduced and located on top of the
midline of the tail. Other differences between this visceral mass. In contrast with Carinaria, the cutis is
species and C. lamarcki cited by Pafort-van Iersel much thinner and the tai1 1acks a dorsal crest.
(1983) include: 1) visceral mass more darkly pig- • Cardiapoda placenta (Lesson, 1830) (Fig. lOA, B).
mented and on a smaller stalk; 2) cutis thinner, Larger of the two species in the genus, exceeding
with more numerous and smaller tubercles; 3) 110 mm in length (Lalli and Gilmer, 1989). Larva1
retinal base of eyes more darkly pigmented; 4) shell consists of three whorls. After metamorphosis
tentacles more strongly developed, right one dis- a small teleoconch forms, which is represented by a
tinctly smaller than left one. Bonnevie reported narrow fringe at right angles to the larval shell aper-
that the claspers were present on sexually mature ture. With growth the shell becomes imbedded in
specimens of 30-40 mm length; thus, this structu- the dorsal surface of the digestive gland, with on1y
re could not be a juvenile feature that is lost in the top of the protoconch visible. The numerous
adults. Reference to these structures as claspers, gills (>20) form a distinctive, fan-like arc encircling
however, is probab1y inappropriate. As Tesch the visceral nucleus. The tail terminates in 12, fing-
(1949) noted they have little to no musculature er-like reddish-brown to b1ack extensions or fila-
and probably could not function in mating. This ments, which form a fan-like structure that can be
species was originally described from the North rapidly contracted when the animal is disturbed
Atlantic as C. lamarcki varo challengeri by Bonne- (Lalli and Gilmer, 1989). Both sexes with reddish-
vie (1920). However, Pafort-van Iersel (1983) brown spots covering the trunk and proboscis; the
identified the above mentioned differences spots are higWy contractile, changing in size from
between it and C. lamarcki, and showed that its 0.5 mm to 2 mm diameter (Lalli and Gilmer, 1989).
spatial distribution overlapped widely with that of Of the two species of Cardiapoda, Tesch (1949)
C. lamarcki (i.e., it was sympatric). She concluded reported that C. placenta was collected at stations
that C. challengeri should be recognized as a va lid close to land, whi1e C. richardi was collected from
speCles. open ocean stations. In agreement with Tesch,
Carillaria cithara Benson, 1835 (not recorded in Pafort-van Iersel (1983) termed the former species
the South Atlantic). neritic and the latter oceanic.
642 Heteropoda
Carinaria challengeri
e
',' .
¡¡;<~~~;\:
2 mm
Low magnification (x8), High magnification (x20),

including visceral nucleus showing structure of c1asper
Fig.9. Sketehes ofposterior end of Carinaria cha/lengeri, right side ofbody. Redrawn from Bonnevie (1920).
• Cardiapoda ric/zardi Vaissiere, 1904 (Fig. 10C). nearly flat, divided down the midline by a low
Small species; maximal body length 20 mm. Shell ridge, and located on the dorsal surface of the vis-
is imbedded in the tissues ofthe digestive gland, and ceral nucleus. While swimming, the visceral
consists of a protoconch and teleoconch represented nucleus is elevated and the gills are exposed. When
as a single whorl, which is mostly incomplete. Eight probed, the nucleus is retracted into a depression in
gills present; much smaller than in C. placenta and the dorsal surface of the disk. This species is not
located at the entrance to the mantle cavity. Body present in the Atlantic, but is included here for
covered by scattered, opalescent pigment spots. A comparison with the other genera of Carinariidae.
large and darkly-pigmented, fan-shaped structure
arises ventrally from the end of the trunk. This con- Family Pterotracheidae Gray, 1843
spicuous structure immediately distinguishes this An adult shell is lacking in this family; the larval shell
species form C. placenta. Tail short, with a very is thrown off during metamorphosis. Like the carina-
long filamentous extension. Sucker on swimming riids, the body is elongate, basically cylindrical, and
fin only present in males (as in Pterotrachea). consists of a proboscis, trunk and tail:--The body,
however, is more slender and streamlined (Fig. lla).
Genus Pteroso11la Lesson, 1827 The visceral nucleus is compacted into a fusiforrn
• PterOS011la planu11l Lesson, 1827 (not recorded in structure, oriented at approximately a 90° angle to the
the South Atlantic). Body length to 80 mm. Trunk long axis of the body. This orientation serves to gre-
expanded as large, flattened and oblong disk. atly reduce the surface area of this opaque structure
Proboscis arises ventral1y beneath the frontal mar- when viewed from beneath, thus potentially reducing
gin ofthe disk, and the tail extends from the poste- the visibility of the animal to upward-searching pre-
rior end, tapering down to a narrow filament. Shell dators (Seapy and Young, 1986). The swimming fin is
Heteropoda 643
Fig. 10. Photographs of Cardiapoda speeies and P/erosoma plalltllll. A: C. placen/a, femate, right side. B: C. placen/a, female, left side. C: C.
richardi, female, left side. D: P. plallllm, maleo left side. Seale bars, A, D: 10 mm, B, C: 5 mm. A-C: eourtesy ofR. Gilmer; D: Seapy, original.
located about midway between the head region and servation. Thus, measurements of body length are
visceral nucleus. A fin sucker is present only in males often not representative of a healthy, relaxed animal.
and has been assumed to function in mating, although To solve this problem, one can calculate body length
this function is uncertain. Harbison (cited in Lalli and from the diameter of the lens using a regression
Gilmer, 1989) observed copulation behavior in equation for the relationship between lens diameter
Pterotrachea hippocampus; while the male and fema- and body length taken from undamaged animals
le were paired ventrally and head to head, the male's (Seapy, 1985). An example of net damage is the holo-
sucker was in contact with the female but could not be type specimen of Pterotrachea minl/ta, which was
seen to actually hold the female. The radula in examined in the above study and found to be severely
Pterotracheidae (type Il) is comparatively smaller stretched; measuring 38.8 rnm totallength. Based on
than in Carinariidae and has a much lower growth the specimen's lens diameter, however, the true body
angle. Also, the number oftooth rows (24-30) is lower length was estimated to be 20.5 rnm.
than in Carinaridae. The central tooth is basically
polycuspid with a prominent central cusp (Fig. 4d); Genus Firoloida Lesueur, 1817
the lateral and marginal teeth are monocuspid (for This genus is monotypic, consisting of a single spe-
morphogenesis of radula and teeth see Richter, 1968). cies with a cosmopolitan distribution.
Pterotracheids are particularly vulnerable to damage • Firoloida desmaresti Lesueur, 1817 (Fig. 12A).
during net capture (particularly stretching) and to Smallest of the pterotracheids; maximal body
shrinkage and the distortion ofbody shape during pre- length about 40 rnm. Terminal visceral nucleus, fol-
644 Heteropoda
Pterotrachea hippocampus
t:~~;;;;a
.\
...•...
.
"\. ....
'!"4'~.'"
~.
~.,
v-==..
-:;:-;¡;~:'"
'':~:.
Male from right side
in normal
swimming orientation
10 mm
Dorsal view of head region

4]) ,, ,
,, , "
"
~~ 1>."
,,
, V-?;. -/s
,
" ~~
~r
,~- - - - - -1 "
~,' 1/
,f
i::::(;'~ , I
, I
,,,0~" , I
,,
, ~
00
Q)
e
,, Q)
Q)
>-
1 mm
Fig, 11. Sketches of Plerolrachea hippocamplls. From Seapy (1985).

Heteropoda 645
Fig. 12. Photographs ofPterotraeheidae. A: Firoloida desmaresli, female, right side. B: Plerolrachea coronala, female, right side. C: P. scula-
la, male, ventral view. D: P. hippocampus, mal e, right side. Seale bars: 10 mm. A, C: eourtesy of R. Gilmer; B, D: Seapy, original.
lowed by a very short, ventral tail and either a per- re. Yentrolateral folds of cutis present on the trunk,
manent egg string (in females) or a filamentous extending posteriorly to near the anterior end of fin.
extension (in males). Body highly transparent (the Tentac1es lacking in males and females. No perma-
only pterotracheid that lacks pigmentation), with a nent posterior egg string in females. Pyriform viscer-
thin cutis and without tuberc1es. Prominent tenta- al nuc1eus ranges in shape from squat to nan'ow and
c1es present in males; lacking in females. Radula elongate. Eyes range in shape from rectangular to
easily distinguished from that of other Pterotrachei- triangular, depending upon the width of the retinal
dae by the shape of the central tooth, which bears base (Bonnevie, 1920). Morphometric analyses ofthe
two long, sickle-shaped and pointed posterior pro- eyes (see Fig. llb for orientation of measurements)
cesses on its basal plate. This species is the most and the visceral nuc1eus by Seapy (1985) revealed
transparent of all heteropods; it is very difficult to species-specific pattems, which are included below.
pick out from a fresh plankton sample. It has a cir- Central tooth of radula with a broad rectangular basal
cumtropical distribution and is extremely abundant plate; more similar to the basal plate in atlantids than
in all oceans; sometimes equal in numbers to in carinarids. The genus differs conspicuously from
Atlantidae (Tesch, 1949). all other heteropods insofar as it bears two rows of
additional palatine teeth on the roof of the buccal
Genus Pterotrachea Niebuhr (ms. Forskiíl), 1775 cavity (MacDonald, 1871).
Body elongate, with prominent tail. Caudal tail fila- Pterotrachea coronata Niebuhr (ms. Forskiíl),
ment present, but usually broken off during net captu- 1775 (Fig. 12B, 13A). Largest of all pterotra-
646 Heteropoda
... ~'~' .._- .
juvenile
" @ o
0.5 mm
1.0 mm
I 1.0 mm I
Pterotrachea
Pterotrachea minuta
coronata Pterotrachea holotype
scutata
Pterotrachea
hippocampus
Fig.13. Sketehes ofright eyes (in dorsal view) offour speeies of Pterotrachea. From Seapy (1985).
cheids; reaching a body length of 330 mm (Lalli and streamlined. Visceral nucleus shape highly
and Gilmer, 1989). Body basically cylindrical in variable; length ranges from two to four times
shape and elongate; the most streamlined in the width; the length to width ratio showing a general
genus. Visceral nucleus shape narrow and elonga- pattem of decrease with increased body length.
te (length ranges from four to seven times width). Eye shape a function of body size; changes from
Eye shape tubular; rectangular in dorsal view, with narrowly triangular in small individual s (length
length averaging 2.2 times retinal width, although about twice retinal width) to broadly triangular in
(like in P. hippocampus) the length to retinal large specimens (length about one and 1/2 times
width ratio decreases with age (from about 2.5 in retinal width).
small individuals to 1.9 in large ones). Pterotrachea minuta Bonnevie, 1920 (Fig. 13E).
Pterotrachea scutata Gegenbaur, 1855 (Fig. 12C, This is the smallest species of Pterotrachea, with a
13B). Second largest pterotracheid; maximal size maximal body length of 36 mm (Pafort-van Iersel,
200 mm. Proboscis, posterior part oftrunk and tail 1983). Eye and visceral nucleus intermediate in
cylindrical. Lateral folds of cutis on anterior one- appearance between P. hippocampus and P. coro-
third of trunk expanded as an oval disk; diagnostic nata. In comparison with P. hippocampus, the eyes
for the species. Visceral nucleus length about three are more narrowly-triangular and the visceral
times width. Like P. coronata, eyes rectangular in nucleus is longer and narrower (length about four
dorsal view, with length averaging 2.2 times retin- times the width). The eye and visceral nucleus
al width. In contrast to the other species in this morphologies used by Bonnevie (1920) to characte-
genus, P. scutata bears a wreath of two or three rize P. minuta were found by Seapy" (1985) to
irregular rows of tiny peribuccal teeth along the match those of young P. hippocampus collected
entrance of the mouth (Vayssü::re, 1904). from Hawaiian waters. Other taxonomic features
• Pterotrachea hippocampus Philippi, 1836 (Fig. used by Bonnevie to separate the two species were
11, 12D, 13C, D). This species reaches a maximal also evaluated in the latter study. Regardless of the
length of80 mm (Bonnevie, 1920). Body cylindri- characters examined, the two species could not be
cal, similar to that of P. coronata, but less narrow resolved and all specimens were concluded to
Heteropoda 647
belong to P. hippocampus. Whether these same Tesch ].J. 1949. Heteropoda. Dana Rep., 34:1-54.
relationships hold true for Atlantic specimens assig- Report on the heteropods collected during the cir-
ned to these two species, and whether or not P. cumg!oba! Dana Expedition. Beginning with his
minuta is, in fact, a valid species, remains uncertain. detailed 1906 report on the Izeteropods of
theSiboga Expedition and concluding with this
report, Tesch brought order out of taxonomic
Acknowledgements chaos for the group. reducing significantly the
number of recognized species and setting the stage
We thank G. Fischer, Department of Geology, for subsequent taxonomic work.
University of Bremen, F. R. G. for making available Thiriot-Quiévreux C. 1973. Heteropoda. Oceanogr.
to us the sediment trap samples from the Walvis Mar. Bio\. Aun. Rev., 11:237-261. Thefirst review
Ridge. We also extend our gratitude to C. Lalli and R. paper on the group, covering a variety of aspects
Gilmer for providing the in-situ carinariid and pterotr- of heteropod biology. Noteworthy for its breadth of
acheid photographs. topical coverage, inclusion of scanning electron
micrographs and detailed treatment of develop-
mental biology and larva! morphology.
Suggested readings Van der Spoel S. 1976. Pseudothecosomata, Gyrnno-
so mata and Heteropoda (Gastropoda). Bohn,
Lalli C.M., Gilmer R.W. 1989. Pelagic snails. The Scheltema & Holkema, Utrecht, pp. 1-484.
biology of holoplanktonic gastropod mollusks. Comprehensive treatment of taxonomy, lists of
Stanford Univ. Press, Stanford, pp. 1-259. The synonymies, and relevent literature through 1974.
book includes a chapter on the Heteropoda, which The material in this text was recently updated and
serves as the most recent review on the biology of published in CD-ROM by van der Spoel et al.
the group. Many unpublished field observations, (1997).
data and ill-situ photographs are i/lcluded.
Pteropoda
Siebrecht van del' Spoel and José R. Dadon
Introduction Pteropods are protandric hermaphrodites and cross

fertilization is the rule, although self fertilization
Pteropods, an informal name for planktonic molluscs probably also occurs. For most species the entire life
of the Orders Thecosomata (shelled pteropods) and cycle is presumed to take about one year. The eggs are
Gyrnnosomata (naked pteropods), are cornmon in all delivered in gelatinous ribbons or spheres. A veliger
marine environments from the poles to the equator, larva hatches, metamorpho~es, and develops into a
and from the surface to bathypelagic depths. juvenile stage. This simple scheme is va lid for
Pteropods are typically open-ocean organisms, and Cavolinia uncinata, Clio cuspidata and Limacina heli-
although many may be encountered in neritic waters, cina, among others. Ovoviviparity is observed in some
there are no real coastal species. species. In Limacina he/icoides and C/io chaptali this
is thought to be an adaptation to the deep-sea environ-
The first important monograph on the group, Boas ment, where protection ofthe young generation is cru-
(1886), concentrated on taxonomy, distribution and cial. However, the epipelagic species Hydromyles
variability. Pteropoda of the major oceanographic globulosa and Limacina inflata are also ovoviviparous.
expeditions were described in the following publica-
tions: Quoy and Gaimard (1824, 1832), d'Orbigny Antarctic pteropods produce one generation per year.
(1836-1846, 1841-1855), Eydoux and Souleyet Subantarctic pteropods show different generation
(1840), Souleyet (1852b), Gould (1861), Pelseneer modes and seasonal differences in growth rates. For
(1866, 1887, 1888a, b, 1906), Jeffreys (1877), Dall example, in the Southwestern Atlantic Limacina ret-
(1885, 1908), Knipowitsch (1902, 1903), Vayssiere roversa reproduces twice ayear. One generation, bom
(1902, 1904), Hedley (1903), Sykes (1904, 1905), in early spring, shows high growth rates and preco-
Tesch (1904, 1910), Meisenheimer (1905b, 1906), cious sexual maturation, and reproduces in late sum-
Schiemenz (1906), Broch (1910), and Bonnevie mer. The second generation grows during autumn but
(1913). Starting with these results, Tesch (1913) pro- shows no growth at all during the winter, reproducing
duced a major monograph encompassing almost all in the spring. This annual cycle matches the general
recent taxa. Two recent monographs (van der Spoel, primary production cycle of the Subantarctic zone,
1967, 1976) and a handbook (Lalli and Gilmer, 1989) with two maxima, the main one in spring and a secon-
surnmarize current knowledge of pelagic molluscs. dary one in early autumn (Dadon, 1989; Dadon and
LaurÍa de Cidre, 1992).
The Thecosomata comprise the shelled species (Fig.
lA-K), of special interest for ecological and geologi- Asexual reproduction has been described for two spe-
cal studies (Biekart, 1989). They contribute signifi- cies: Clio pyramidata and C. po lita , but the reproduc-
cantly to the carbonate cycle of the ocean, leaving a tive cycle ofthese species is not yet complete1y under-
fossil record that can be important in palaeoclimatic, stood. Schizogamy was observed in both (van der
palaeoceanographic, and palaeocologic studies Spoel, 1979): the adult specimen divides transversely
(Herman and Rosenberg, 1969; Diester-Haass, 1972; at the middle so that an upper soft part with all the
Buccheri and di Stefano, 1984). The Gymnosomata vegetative organs becomes detached and swims out of
(Fig. 1L-O), are shelled only in an early tife stage, the shell. The lower half, with gonad tissues, remains
usually do not form swarms and do not contribute in the shell and develops wings. Once these are func-
much to the zooplankton biomass except for some tional, it also lea ves the shell and starts a free swim-
species in the polar seas. They are carnivorous and ming periodo Probably this part delivers eggs only
most species are specialized feeders. when the numerous ova in the body are fertilized with
the sperm already present before schizogamy, but this
has not yet been observed.
SOl/th Atlantic Zooplanktoll.

edited by D. Bolto\'skoy. pp. 649-706
1999 Backhl/Ys Pl/blishers, Leiden, The Netherlands
650 Pteropoda
EUTHECOSOMATA
-5
eO
u
O
Q)
~ mantle
second
flank¡
IOCk~,
columella spine~
,
I
surface
I
________ 1
Limacina in oral view
median lip depression

centralrib
(dorsal) dorsa I l'Ip Diacavolinia in ventral view
(d~~~~í)lrib 1- _lliP,rims
ventral
I'Pl/
1/ ~tl
I \ \,;"
'-l' \ - \ lIt :~ ./ lock
~~nG~a¿e
y
,pertuz.¿ ~~, /"e,
k I\. i
,ec,~~~
e\e •/\
W
~
te'eow"h
(ventral side)
~
\
1
ventral
db,
lock
lateral ~ tubercles
c.
VSPine
L
ventral view dorsal view

o
t~! J..~ Cavoliniid lock system
,"ud,'
lateral line ~
'pme - ...---'"
protoconch <i
Cavolinia in ventral view
Pteropoda 651
EUTHECOSOMATA
nose
lip radula
1 angle plate
teleoconch
¡,
'~
// outer
hump
,7" I
"'''\ dorsal
~ nbs
/.~\\1" ',-=-ril~~eral
lock
@
/11 '\ \ -
Radula teeth
.ir i 1 nb
\ ~slde
I
I
1
li
11
i\ /
-
~
J '\.
d.orsal
'p;oe
surface
protoconch 11~
,. Cuvierina with
detaehed protoeoneh
1
1
I
I
Diacavolinia in lateral view 4) m

PSEUDOTHECOSOMATA
pseudoconch
1
lip
. Al!Vl o ~¡/
eplpodial
tentacle ~L
- \;: "'"
Desmopterus in oral view

median footlobe
o Cymbulia in oral view o

652 Pteropoda
PSE U DOTH ECOSOMAT A GYMNOSOMATA
'---- hooks
median
tubercle
anus
median
footlobe
Peraclis in oral view
crest
fringed
Pneumodermopsis in ventral view
~MC¡¡
1j!JP/~
median lateral
plate teeth
Spongiobranchaea
Clione
upper half in ventral view aradula
ustra lis ,.,
~
Pteropoda 653
Development of the shell and the soft parts may be Pteropod shell micro-structure is simpler than that of
parallel (e.g., in Limacina), or may differ in speed, as the average gastropod shell, being composed mainly
in Cavoliniidae. Some Pseudothecosomata have no of a single layer (Bé el al., 1972).
shell, developing only a pseudo-shell, and in the
Gymnosomata shell development ceases after the Gyrnnosomata are hunters, whereas Thecosomata are
embryonic stage. The development in Limacinidae, ornnivorous mucus feeders. The solitary Gyrnnosomata
primitive Cavoliniidae, and Peraclididae is the most are usually specialized feeders, the extreme example
"typically molluscan". After the embryonic stage (i.e., being that of Clione limacina that, in the North
after protoconch 1 is formed) the juvenile produces the Atlantic, feeds only on Limacina helicina, and in the
protoconch II and the soft parts grow regularly to South Atlantic on L. helicina and L. relroversa
maturity, when the teleoconch is formed. Seasonal (Conover and Lalli, 1972). Most other species are less
growth rates have been studied for Limacina relrover- specia1ized in their diet but all feed either exclusively or
sa in neritic waters. During the cold season the te leo- main1y on a few Thecosomata species (Lalli, 1972).
conch shows liUle or no growth, whereas in spring
growth rates are high (Dadon, 1989). Thecosomata consume microplankton that is collect-
ed in a mucous trap (Gilmer, 1974; Gilmer and
In Styliola, Clio, and Cavolinia growth proceeds nor- Harbison, 1986). All Pseudothecosomata produce
mally to the end of the juvenile stage. Then shell free-floating mucus webs (Gilmer, 1972) of enormous
growth continues while soft parts stop growing, on1y size, with diameters lOto 100 times that of the animal
e10ngating to reach the shell rim. This produces a thin itself. The mucus with microplanktonic organisms
teleoconch which contains a very tiny soft body, known adhering to it is eaten and a new web is produced.
as the minute or skinny stage. Not until the teleoconch These webs have a1so been described for so me
reaches full size and shape do the soft parts grow fur- Cavoliniidae. The existence of such webs is contro-
ther to maturity. In adults shell growth usually does not versial in the Limacinidae, which al so feed by means
stop entire1y but continues very slow1y at the aperture of a mucous apparatus. The mucus is secreted on the
border, also involving some thickening of the she11. wing surface by glands. The food of Thecosomata
consists chiefly of phytoplankton, including cocolith-
In Hyalocylis, Cuvierininae and Cavoliniinae (except ophorids, silicoflagellates, diatoms and dinoflage1-
Cavolinia) development is the same, but the shell is lates, and also foraminifers, radiolarians, and other
actively broken during the minute or skinny stage. protists.
The teleoconch is separated from the protoconch and
the junction is closed by a septum. In Diacavolinia The Limacinidae, Cavoliniidae, Peraclididae, Cymbulii-
this is accomp1ished by fusion of hind dorsal and ven- dae, Pneumodermopsidae, Notobranchaeidae, TWipto-
tral shell sides to form a joint. Maturation is reached dontinae, Cliopsidae, Clionidae, Hydromylidae, are all
only after loss of the juvenile shell parts. monophy1etic as based on apomorphies. The evolution-
ary status of the Laginiopsidae, which have been stud-
In the Cymbuliidae shell development ceases after the ied much less, is unclear. Based on the apomorphic
juveni1e stage (with left-coiled shell), and metamor- character of development of the hooks, the Gyrnno-
phosis involves the production of a pseudoconch. somata, except for H)'dromyles, can be grouped togeth-
er as a monophyletic group. For the Thecosomata, in
In Gymnosomata the embryonic shell, formed in the which real apomorphies are difficult to find, further
egg, is lost and a typical juvenile naked stage with grouping is prob1ematic. They may be monophy1etic,
three ciliated bands around the body emerges (Fig. but there is still no real proof.
10). With age these ciliated bands are lost and gonads
develop. Paedoclione is an exception in that the cili- Pteropods are usually placed in the gastropod subclass
ated juvenile normally shows sexual maturity, the Opisthobranchia. The Gyrnnosomata were considered
genus thus being clearly neotenic. to be close to the Anaspidae, and the Thecosomata
4
Fig. 1. General morphology and characters used for identification in Pteropoda.
654 Pteropoda
close to the Pyramidellidae or Philinoglossoidae. The species are very fragile. The tows should be of short
fully developed operculum in Limacina and Peraclis duration to prevent damage. The net should be opened
indicates that this classification of the Thecosomata gently and cleaned of its contents with extreme careo
may be questionable. The cylindrical tentacles with Collecting Cymbuliidae requires special care beca use
eyes on top in a11 pteropods, ornamented shells in pseudoconchs easily detach from the soft parts. Scuba
Peraclididae, and high spiral she11s with fully deve- diving and sampling/observation from submersibles
loped operculum in Limacinidae and Peraclididae, are modern developments which offer unique oppor-
show that pteropods sti11 ha ve close links to the tunities to collect the fragile animals.
Prosobranchia. The Pyramide11idae also represent a
group between the Opisthobranchia and the Preservation and collections
Prosobranchia.
Net samples should be sorted before fixation. Naked
The Thecosomata, however, may be polyphyletic, and species are best preserved in formalin, preferably after
are certainly not monophyletic with the Gyrnnosomata. narcotization (Heyman, 1981). Alcohol is also ade-
The frequently proposed relation of pteropods (espe- quate, and is especially recommended for shelled spe-
cially thecosomatous ones) with the Bu110idae by con- cies in museum co11ections. Specimens should be han-
sidering their anatomy neotenic (resembling bulloid dled carefully with sma11 forceps (e.g. entomological
larvae) is not generally accepted (cf. Lalli and Gilmer, aluminium forceps) as the bodies and shells are frag-
1989). Though there may be a few neotenic characters ile. A dissecting microscope suffices for observation
in pteropods, there is no indication that pteropods, of most characters, although a compound microscope
except for Paedoclione, are in any sense reproductive is required for the buccal organs.
larval organisms. Lalli and Gilmer (1989) suggest that
the feeding behaviour and buoyancy strategy shared by Alcohol-preserved, formalin-preserved, and even dry
all Thecosomata indicate their common ancestry. collections should be kept in the dark. Specimens
Inasmuch as the Thecosomata seem to ha ve more cha- should not come in contact with wood or cardboard
racters in common with the Prosobranchia than do the (Moolenbeek, 1994). Glass tubes and jars can be used
Gyrnnosomata, the latter may be considered hypothe- but, especially for dry collections, glass should be
tically more highly evolved. avoided as it affects the shells through a process of re-
crystalization. AII shelled specimens should be proper-
ly cleaned with fresh water before storage. Cotton wool
Methods is used to cap tubes in dry co11ections and alcohol-fi11ed
bottles, but such stoppers should be wrapped in soft or
Sampling rice-paper in order to prevent the tiny organisms from
becoming entangled in the cotton wool. Whenever pos-
The tiny shells of pelagic molluscs can be found along sible labels should not be placed in the jar together with
the beach. However, they are not easily visible because the animals, as even a paper label may crush the shells.
of their small size, and are usually overlooked unless Dry shells can be handled with a wet brush to which the
occurring in mass accumulations. Because the she11s specimens adhere upon contact, or with fine forceps.
are very fragile only fragments are usua11y found.
Dissection of the radula, sometimes needed for iden-
For quantitative and biogeographic studies the most tification, is achieved by making a cross cut in the
widespread co11ection method is by nets with a mesh ventral side of the neck region and picking out the
size of 0.3 to 4.0 mm, although 0.05-0.2 mm mesh esophagus at the level of the ganglia. This section of
openings are required for retrieval of juveniles. the mouth parts is placed on a slide, the tissues are dis-
Whenever possible opening-closing nets which col- solved with KOH so that the chitinous plates become
lect animals from pre-determined depth intervals are visible under the microscope. Almost any medium
preferred. Non-closing nets, however, which were can be used for mounting. Hooks and buccal cones
used in many collections, sti11offer a good opportu- can be accessed by making a cut in the hood over the
nity to study horizontal distributions. Handling nets buccal mass, and the organs can be mounted on
requires special care as the she11ed as we11 as naked microscopic slides as described above for the radula.
Pteropoda 655
Geographic and vertical distribution specific forms are related to climatic belts, including
cold-temperate (C p. forma pyramidata), warm-water
World wide distribution patterns (C p. forma lanceolata), polar (C p. forma su/cata),
and subpolar (C p. forma antarctica) representatives.
Pteropods are most abundant in Subarctic and
Subantarctic waters. The Thecosomata (Limacina) are Species restricted mainly to particular water masses,
sometimes dominant components of the plankton in like Limacina lesueuri, represent another distribution
these areas, and may represent an important food type, not related to climatic belts but restricted to
source for whales and large fish such as codo As water masses. 1. leslleuri is found in the Central water
Thecosomata are largely phytoplankton feeders, the masses of al! oceans, thus comprising five discrete
effects of grazing by species like Limacina he/icina populations. However, since this species is easily
antarctica can be very important in polar seas. The expatriated to waters outside the gyres, its pattern may
swarming behaviour in most Thecosomata and a few resemble a warm-water belt pattern. Diacria major is
Gyrnnosomata like C/ione /imacina antarctica results even more restricted to the five Central water masses,
in very patchy distribution patterns and varying inasmuch as it is not strongly expatriated.
effects on the local ecosystem. Limacina inflata,
Creseis and Diacavo/inia are considered to have nerit- In the genus Diacria both types of distributional pat-
ic preferences; they may occur in very high abundan- terns apply, serving as an example ofthe main biogeo-
ces in coastal areas. Limacina injlata, however, is graphic trends in pteropods. There are two species
usual!y absent from inner neritic waters, where groups in the genus, the "qlladridentata group" with
Creseis acicu/a forma acicula is most abundant. nine taxa, and the "trispinosa group" with seven taxa
Though meso- and bathypelagic species are known to (see Fig. 2). In the qlladridentata group D. danae
exist, most pteropods are epipelagic, and high abun- "Subtropical form" is the most widespread, living in
dances are found only in the epipelagic realm. all oceans roughly in latitudinal belts at 1O-30oN and
S; and D. danae "Equatorial form" is restricted to the
Pteropods are geologically a rather recent group, the equatorial waters of all oceans. The distribution of this
oldest fossils dating from the Miocene. They probably species and its forms clearly follows climatic belts.
originate from an Atlantic centre of speciation (van der
Spoel and Heyman, 1983). This is still visible in the dis- D. quadridentata is an Indo-Pacific warm-water spe-
tribution patterns of some groups, such as the mesope- cies that does not enter the Atlantic Ocean because the
lagic genus Perac!is, whose species have different African continent interrupts dispersal of really warm-
ranges but are all found in the Northeastern Atlantic off water species. D. costata "Equatorial form", an Indo-
Dakar. It is self-explanatory that these deep-living spe- Pacific organism, shows a range comparable to,
cies do not reflect the influences of climate and shallow though narrower than, the preceding one. In the
water mass configurations, inasmuch as their distribu- Pacific it gives rise to a second form, D. costata
tion patterns probably reflect older ocean climatic pro- "Central Water form" that is especially adapted to the
vinces. In bathypelagic species, like Limacina he/i- Central Water Masses. D. schmidti schmidti is
coides, climatic influences are undetectable. Restriction restricted to the Eastern Pacific Ocean oxygen mini-
of this bathypelagic species to the Atlantic Ocean and mum area. This is a well-established zoogeographic
its deep water outflow may indicate that it originated in province with many endemic taxa. D. schmidti occi-
the oldest (late Cretaceous) deep-sea basin, the North dentalis is restricted to western Pacific coastal areas.
Atlantic Ocean. The bathypelagic C/io andreae has a D. erythra elythra is a Red Sea and western Indian
much wider distribution, which may be explained by ocean endemic species that probably originated in the
the abundance of young stages in shallow layers. Red Sea and subsequently invaded part of the Indian
Ocean. D. erythra crassa is endemic to the Red Sea.
The general zoogeography of pteropods is illustrated
below with some typical patterns. Species cornmon in In the trispinosa group (Fig. 2) D. trispinosa fonna tri-
the epipelagic layers are influenced by climate and cli- spinosa is the most widespread taxon, living in all
mate changes in addition to currents and water masses. oceans between 400N and 40oS. D. trispinosa atlantica
In Clio pyramidata, a cosmopolitan species, many infra- "typical form", found in the North Atlantic between
200
400
0°
200
Ew
656 40°
00
•••
cria trispinosa
E
E;]D
6maculata
atlant. upw.
maJor
rampali
trispinosa
piccola
ooojI
[!]
I fOO
1600
Pteropoda •
erythra costata
D
occidentalis quadridentata crassa
D
schmidti
l~~
danae
Fig. 2. Distribution of the Diaeria Irispillosa and the Diaeria quadridelllala groups (the "equatorial" and "central" water forms are not indi-
cated).
700N and 40oN, inhabits the transitional waters. D. tri- found in shells preserved in sediments south of Africa.
spinosa atlantica "upwelling form" is especially adapt- D. mandata is found in northwestem boundary cur-
ed to the upwelling area off Northwest Africa. rents of the Atlantic and Pacific Oceans, seemingly
Upwelling areas also host isolated populations of many adapted to coastal water masses with a tropical origino
other planktonic and nektonic groups. Diacria rampali D. piccola has been collected so far only from sedi-
lives in all oceans between 300N and 30oS, maintaining ments. D. major appears only in the Central Water
this broad distribution by expatriation around the south- masses of all oceans, and is therefore a typical water
em tip of Africa. Proof for such expatriation can be mass-linked species.
Pteropoda 657
Endemism is rare in pelagic molluscs, although a few Limacina helicoides, Clio chaptali, and C. andreae,
cases have been recorded. The oxygen minimum layer and the Gymnosomata Massya and Schizobrachium are
of the eastem tropical Pacific hosts some endemic bathypelagic. Clio andreae and Limacina helicoides
species such as Desmopterus pacificus. This area is undergo ontogenetic vertical migration, juveniles
not physically isolated, but adaptations to the special occurring higher in the water colurnn than the adults.
conditions have resulted in endemism. The
Mediterranean is more or less isolated as the Strait of Distribution in the South Atlantic
Gibraltar functions as a partial barrier, and special
endemic forms of a few species, such as Cavolinia tri- According to their affinities with water masses (Fig. 3,
dentata forma tridentata, are known here. A compar- 4), pteropods of the South Atlantic can be divided into
able situation is found in the Red Sea where endemics warm water, cold water, Subantarctic and Antarctic
like Diacavolinia jlexipes occur. species. No transitional forms are known in the area.
Vertical patterns and migration The warm water fauna is the most diverse, with more
than 80 taxa of Thecosomata and Gyrnnosomata. It
Many pteropods undergo more or less pronounced inhabits equatorial and subequatorial waters, reaching a
diel vertical migrations, feeding at night near the SUf- southem distributional limit at the Subtropical Front.
face and migrating to greater depths during the day The Pseudothecosomata are restricted to this region.
(Stubbings, 1938; Humphreys and Myers, 1968; van
der Spoel, 1973b; Kobayashi, 1974; Wormuth, 1981). Thoügh poor in diversity, Euthecosomata and
Reverse migrations, with shallower daytime OCCUf- Gymnosomata are abundant in cold waters. The com-
rences, have been reported in a few cases (Stubbings, monest cold water species is Limacina helicina an-
1938). Most species are influenced by specific hydro- tarclica, its geographic range extending from the
graphic conditions so that day and night levels are dif- Subtropical Front to the Antarctic coast. L. lzelicina
ferent from place to place. Migration ranges vary in antarctica f. antarctica is the dominant pelagic gas-
different species; for example, Diacavolinia species tropod in Antarctic waters. Swarms of this species
migrate daily only over some ten metres, whereas with densities over 10,000 ind. per 1000 m3 were
Clio pyramidata migrates over hundreds, even up to found south the Antarctic Convergence near 65°S
1500, metres (van der Spoel, 1973b). (Chen, 1968a). L. helicina antarclica f. rangi is very
frequent in Subantarctic waters. In the Argentine Sea
Irregular bottom topography is often responsible for and surrounding areas, large numbers of that species
remarkable deviations from the normal vertical pat- inhabit the slope and penetrate the shelf water up to
temo For example, deep-sea species may occupy shal- the 100 m isobath, more exceptionally up to the 50 m
lower layers above sea mounts and in other shallow isobath (Dadon, 1989). Here the swarms seem to be
waters. Neritic species like Creseis acicllla may smaller (Dadon, op. cit.).
descend to the bottom at night in shallow areas where
intermediate depths are lacking. Deviations from nor- Other cold water species are the Thecosomata
mal vertical distributions may also be induced by pro- Limacina retroversa allstralis and the Gyrnnosomata
ductivity and temperature changes (van der Spoel and Clione limacina antarctica and Spongiobranchaea
Schalk, 1988; Schalk and van der Spoel, 1988). When australis. L. retroversa australis is clearly a subantarc-
temperatures at depth are slightly higher than the usual tic species, whereas both Gyrnnosomata are present
0-4°C, as in the Banda Sea, normally bathypelagic spe- from the Subtropical Front to the Antarctic coast.
cies may appear in the epipelagic zone. Furthermore,
diel vertical migrations may be suppressed altogether Table 1 lists the world wide biogeographic affinities
when productivity is above normal (Schalk, 1988). by species; further details are given below.
Although most species are epipelagic, several are typi- Limacina bulimoides (Fig. 5.4). The dominant ptero-
cally mesopelagic (see Table 1): e.g. Clio recurva. pod in the Benguela Current (Morton, 1954),
Peraclis spp., and some Gymnosomata, such as where its abundance may exceed 350 ind. per m3
Thliptodon and Ceplzalobrachia. The Thecosomata (Dadon and Masello, in press).
658 Pteropoda
30' o"
Eastern o'
Tropical
Atlantic
20'
South Atlantic
Tropical Gyre
Falkland
(=Malvinas)
Subantarctic
60"
Antarctic
Austral
Polar
Fig.3. Surface water masses (after Peterson and Stramma, 1991) and biogeochemical provinces (after Longhurst, 1995) in the South Atlantic.
Limacilla helicilla alltarctica forma all1arctica (Fig. Limacilla /eslIeuri (Fig. 5.8). Most abundant in the
5.1). Cold water stenothermic. Present in the Central Atlantic, with densities to 24 ind. m-3
Southem Ocean from Antarctic coastal waters to (Dadon and Masello, in press), and frequent but
the southem part ofthe West Wind Drift. not abundant in the boundary currents.
Limacilla he/icilla alltarctica forma rallgi (Fig. 5.2). Limacilla retroversa allstralis (Fig. 5.3). The most
Present to the northem limit of the West Wind abundant pelagic mollusc in Subantarctic waters,
Drift, but extending with the Malvinas with up to 67 ind. per m3 (Dadon, 1990a, b),
(=Falkland) Current as far north as the Brazil- exceptionally to 1,000 ind. per m3 (Boltovskoy,
Malvinas (=Falkland) Convergence. Near South 1971 a, b). Its geographic range in surface waters
Africa it occurs to 30oS. North ofthe Weddell Sea extends from the Subtropical Front to 3-10
the southem boundary of this form bends south- degrees south of the Antarctic Convergence
wards. A chne seems to develop, with the forma (Chen, 1968a; Dadon, 1993). Off southem South
rangi at the northem extreme and the forma an- America it is very abundant in the slope and shelf
tarctica at the southem end (Dadon, 1993). water up to the 100 m isobath (south of 41°S;
Limacilla illflata (Fig. 5.7). Very abundant in the Brazil Dadon, 1990a). The shelfbecomes narrower to the
Current (reaching 38 ind. per m3), but its numbers north, and here swarms drift to the coast and the
decrease in the Central Atlantic and in the Benguela species is recorded up to the 50 m isobath (Dadon,
Current (Dadon and Masello, in press). 1989). L. retroversa occurs near Cabo Frio
Pteropoda 659
Antarctic
Surface
20" 40" 80"5
1000
---- 3000 AtlanticDeep

Q.
Water
O .r. 20001
••••
Q)
4000
Fig.4. Vertical water-mass structure ofthe South Atlantic. Modified from van der Spoel and Heyman (1983).
(Massy, 1920), but this may be the result of fre- lation of C. i. forma labiata is found between 0°
quent local upwellings (see "General biological and 400S, and a smaller population occurs in the
features of the South Atlantic"). Swarming in central and southem Arabian Sea. The South
neritic waters is associated with retention areas on Indian and Atlantic populations are connected by
the outer shelf, but even these areas cannot prevent the population in the Agulhas Current, composed
a massive seasonal migration of stocks into the of a mixture of the forms injlexa and labiata (Fig.
oceanic zone (Dadon, 1990a). 5.29, 5.30). In the Pacific the species does not
Li",acilla trochiformis (Fig. 5.5). Abundant in the occur north of 400N, in contrast to the Atlantic
Brazil Current (up to 1.8 ind. per m3 off Cabo population which reaches 55°N. C. i. forma inflexa
Frio; Dadon and Esnal, 1995), but absent east of is the most cornmon form in the North Atlantic. C.
100W (van der Spoel, 1967). i. forma imitans is found in the Atlantic and near
Cavolillia illjlexa formae illjlexa, i",itans and labia- Zanzibar. In the Atlantic it occurs between 55°N
ta. C. inflexa is a warm water species. C. i. forma and 45°S, and is absent in the Benguela Current
labiata shows a tendency for bisubtropical distri- and off northem South America.
bution, more abundant in the South Atlantic, Clio alldreae (Fig. 5.15). Only 3 small populations
South Indian, and Pacific Oceans, but also present are recorded south of SON: off Cabo Blanco, off
in the Mediterranean. In the Indian Ocean a popu- Cape of Good Hope, and in the Argentine Basin.
660 Pteropoda
5.1
Limacina helicina antarctica Limacina helicina antarctica Limacina retroversa australis Limacina bulimoides
forma antarctica forma rangi
Limacina trochiformis Limacina helicoides Limacina inflata Limacina lesueuri
Creseis acicula Creseis acicula Creseis virgula Creseis virgula

forma acicula forma clava forma conica forma virgula
Hyalocylis striata 5tyliola subula Clio andreae Clio chaptali

Pteropoda 661
C1io cuspidata C1io piatkowskii C1io pyramidata C1io pyramidata

forma antarctica forma excisa
C1io pyramidata C1io pyramidata C1io recurva Cuvierina co/umnella

forma lanceo/ata forma su/cata forma at/antica
Diacria quadridentata s.1. Diacria rampali Diacria trispinosa

forma trispinosa Cavolinia gibbosa
forma flava
Cavolinia inflexa Cavolinia inflexa Cavolinia tridentata Cavolinia tridentata

forma inflexa forma labiata forma atlantica forma australis
662 Pteropoda
Cavolinia uncinata uncinata Diacavolinia limbata Diacavolinia longirostris Diacavolinia strangulata

forma uncinata forma limbata
Peraclis apicifulva Peraclis bispinosa Peraclis depressa Peraclis moluccensis
Peraclis reliculata Peraclis triacantha Peraclis valdiviae Cymbulia peroni

morpha minor
Cymbulia peroni Cymbulia sibogae Cymbulia parvidentata Corolla calceola

morpha peroni
Pteropoda 663
Corolla intermedia Corolla spectabilis Gleba cordata Desmopterus papilio
Clione limacina antarctica Thliptodon diaphanus Cliopsis krohni Notobranchaea macdonaldi

morpha krohni morpha macdonaldi
Notobranchaea tetrabranchiata Pneumoderma atlanticum Pneumoderma atlanticum Pneumoderma atlanticum

atlanticum forma atlanticum atlanticum forma eurycotylum atlanticum forma pygmaeum
••••• i
5.64
Pneumoderma atlanticum Pneumoderma mediterraneum Pneumodermopsis paucidens Pneumodermopsis macrochira

forma souleyeti forma paucidens
664 Pteropoda
5.67
-
Schizobrachium
polycotylum
documented
range
inferred
range
seasonalor
incidental
occurrences
Spongiobranchaea intermedia
interpolated
seasonalor
incidental
occurrences
Fig. 5. Sketches of the horizontal distribution of pteropod species in the South Atlantic.
Clio cllspidata (Fig. 5.17). Found east of 400W Antarctic Convergence to the Antarctic coast. lt is
between 65°N and 200S. In the southem part ofthe not abundant; dense patches were observed only in
Eastem Atlantic, influences of Benguela Current the South Sandwich Trench, reaching 10,000 ind.
are evident. 1000 m-3 (Massy, 1932; Chen, 1968a; RamÍrez
Clio piatkolVskii (Fig. 5.18). Recorded in the Weddell and Viñas, 1983).
Sea at great depth. Creseis acicllla forma aciCllla (Fig. 5.9). Coastal
Clio pyralllidata forma alltarctica (Fig. 5.19). This areas off Brazil.
form occurs irnmediately south of the area of Clio Creseis virgula forma virgula (Fig. 5.12). Frequent in
pyramidata forma lanceolata (Fig. 5.21). It is Brazilian neritic waters, and some vague records
found in the Westem Atlantic between 200S and in the Caribbean Area, in addition to those of a
55°S, south of the Agulhas Current south of small population in the Gulf of Darien.
Australia and New Zealand at 500S, and west of Diacavolillia lilllbata forma lilllbata (Fig. 5.34).
Chile at 500S. It normally inhabits Antarctic Present in the upwelling area offNorthwest Africa
Intermediate Waters and Subantarctic and the tropical westem Atlantic.
Intermediate Waters (Chen, 1966). It is frequent, Diacavolinia strangulata (Fig. 5.36). Originally
but not abundant, and widely distributed between described from the West Pacific Ocean, but pro-
40° and 600S. It is rare at the surface, with maxi- bably has a wider distribution.
mum densities reaching about 200 ind. 1000 m-3 Styliola subida (Fig. 5.14). Frequent in the Brazil
(Chen, 1968a). Never found in neritic areas off Current (to 60 ind. 1000 m-3; Dadon and Esnal,
South America (Dadon, 1989). 1995). One exceptional record at 64°S is due to
Clio pyralllidata forma sulcata (Fig. 5.22). Recorded expatriation by local currents.
between the area of Clio pyramidata forma an- CYlllbulia perolli morpha perolli (Fig. 5.45). Present in
tarctica and the Antarctic Continent. South of the Mediterranean and occasionally in the Atlantic.
Australia, between 1800W-1600W and 1400W- Peraclis bispillosa (Fig. 5.38). Recorded from 600N
900W. the forma sulcata seems absent. lt is char- (northwest of Iceland) to 25°S (off the West
acteristic for Antarctic water, and found from the African coast).
Pteropoda 665
Peraclis depressa (Fig. 5.39). Typical for the eastem became very important (van der Spoel, 1967).
Atlantic, occurring in the eastem and southem Isolation is rare and problema tic in the marine pelagic
basins. realm where all water masses tend eventually to mix
Peraclis valdiviae (Fig. 5.43). In the Pacific, isolated and isolated basins are essentially lacking. So geo-
records from the Gulf of Panama, the Tasman Sea graphically isolated subspecies are only rarely found,
and the Indo-Malayan archipelago. Numerous though the Mediterranean and Red Sea are relatively
records from the northem and southem Indian isolated areas where independent taxa may develop. In
Ocean and the entire Atlantic Ocean. It c1early the late sixties the cladistic and phylogenetic approach
occupies shallower depths in colder areas than in was stimulated by the theory of Hennig.
warmer areas, with pronounced subtropical sub-
mergence at 40-45°N and 40-45°S. Variation is cornmon in planktonic animals with
Cephalobrachia macrochaeta. Bathypelagic (below broad distributions ranging over different climatic
1000 m) between 65°N-I0oS, in colder areas also belts and ocean basins. Species with a large north-
between 400 and 750 m. Probably a cosmopolitan south range tend to reflect the influences of c1imate,
deep-sea species. and species that occur in various water masses usual-
Clione limacina antarctica (Fig. 5.53). This is the Iy differ in the different environments.
dominant Gyrnnosomata in Antarctic waters (Chen
and Ericson, 1967), extending to the Subtropical Clio pyramidata exhibits shell-shape variations: nar-
Front. In neritic water it is present on the outer shelf row pyramidal in polar seas, broad lanceolate in the
up to the 100 m isobath (Dadon and Chauvin, tropical waters. Its embryonic shelJ differs in size: the
1998). warmer the water the smaller the volume of protoconch
SpolIgiobrallchaea australis (Fig. 5.66). The most fre- 1. Thus, two different trends of variation are involved,
quent Gyrnnosomata in subantarctic waters, also the protoconch showing a purely eco-phenotypical
present in the Antarctic; it is found in c10se associ- temperature-dependant variation, and shell-shape
ation with its prey, Limacina helicina antarctica showing clinal variation depending on selective pres-
and L. retroversa australis. In neritic waters it sure in different water masses and restricted gene-flow
occurs on the outer shelf, eventually reaching the between populations. Restricted gene-flow smooths out
80 m isobath (Dadon and Chauvin, 1998). strong differences, producing gradual c1ines, whereas
SpolIgiobrallchaea intermedia (Fig. 5.67). Probably absence of gene-flow yields stepped clines. In Clio
a eurythermic, cosmopolitan species. Known from steps in the cline occur at boundaries between major
only four 10calities; Panama, Dakar, Argentine water masses (van der Spoel, 1967). Thus, shelJ shapes
waters and the Tasman Sea. of C/io pyramidata can be used as hydrographic indica-
Thliptodon antarcticus. Its Antarctic record is dubi- tors, and embryonic shell dimensions can serve as eco-
ous as it was reported by Meisenheimer from a logical indicators (Fumestin, 1978, 1979).
single sample from the Valdivia expedition in
which many tropical organisms were present at an Intraspecific variability in South Atlantic specimens of
extremely southem loca lity (incorrect labeling?). L. helicina has been discussed extensively (Tesch,
1946; van der Spoel, 1967, 1976; Dadon, 1984a, b,
1989, 1993). Associated with climatic belts (van der
Taxonorny Spoel, 1967), an ecocline is found along its latitudinal
range in the South Atlantic, affecting differences in
Variation both morphology and life cycle strategy (Dadon, 1993).
At the tum of the century the typological species con- Diacria costata is a typical example of water mass-
cept and morphological studies dominated taxonomy. related variation (Van Leyen and van der Spoel,
Pfeffer (1880) and Boas (1886), however, already 1982). In Central waters the shells are smaller than in
payed attention to the concept of variability in ptero- Equatorial waters. An example of more locally
pods. Many species are represented by one or more restricted variations is in the shell-color of Diacria tri-
subspecific taxa. With the advent ofthe theory of geo- spinosa, with a pattem different in upwelling areas
graphic speciation (Mayr, 1963), geographic variation from that in the open ocean.
666 ~
++u
c:
.¡;o
.;=
+
.D
++
<ii
~
Q).D
-oa¡
.~
oU'"u
,;::
a¡
o+Q)
± co
::J
.;=
~ S u
~
'"+
o-'"~00
Q.
c:
-::J++
eQ)a¡c:~'".¡;o
Q.::J
'6- ± '" +-z
eNU+a¡ua¡~..c:co(§~+~VlQ.a¡G±.-'"co3:VluU
~ s L.U E
::J ±
u ~Uo~'" .D0c..+'"Q.~·13+±::Jcoo-+~c: e+~+++2±+>-U+eu.'=!+ Q.uu+'" .¡;o
+'"c: '" + + ++ + ++ + + +
++Pteropoda
rl
3: <u
+ + ± - 1
macina
avolinia Cavolinia
Suborder :1Euthecosomata
helicina
Limacina
Cavolinia
Diacavolinia
gibbosa -1
Family antarcticaretroversa
inflexa
inflexa
helicina
tridentata
uncinata
u Limacinidae angulosa australis
antarctica
J
--1 j
Pteropoda 667
'" '" u u u
u U .:t= ro t:lO u '00
__ ttI
'00
u -e
1"0
~
e ~ 0;::
O ~ u ~~~ .~ u
E~g-g~~'ñc c.. ro •... o
._
ClJ
::J '"
~ ClJ
'''::;e >- e 0;:;
~ ~ ~ S
~I
ClJ O
::J .- ~ ~
.~ e -§
.~ .~
~U/-""'COr"'«O ~
.6-1 J!!
-g
.'ª ~u .~ eO:¿ ~ S Ue ClJ o.
::J
3: 1
."
«~l·~"D
.E: ~0.1 ~ ..c
~
V>
O
u
Q) o;::
zClJ
Diacavolinia limbata
+ + + + + +
forma limbata
Diacavolinia longirostris + + + + + + + + + ±
Diacavolinia ovalis + + + + + + + ±
Diacavolinia robusta + + + + + + +
Diacavolinia strangulata + + + + + ±
+-1 -t-j-
Diacria danae + + + + + I - + + + + + +
- I
Diacria major + + + + + + + + +
Diacria rampali I~ + + + +1 -I± + + + I + + + +
Diacria trispinosa + + + + + ± - I ± + + + I + + + +
forma trispinosa
Clio andreae + + + + + + + + + + + + + +
Clio chaptali + + + + + + + + + + +
Clio cuspidata + + + ± + ± + + + + + + +
Clio piatkowskii + + + +
Clio pyramidata + + + + + + + + +
forma antarctica
Clio pyramidata forma excisa -1 + + + + +
-1
Clio pyramidata + + + ± + + I - ± + + + + + + +
forma lanceolata
Clio pyramidata + + + +
forma martensi
Clio pyramidata ± ++ - ± + + + + +
forma sulcata
Clio recurva + + + + + + + + + + +
+ ± + + + + +
Creseis acicula forma clava + + + +
Creseis acicula forma acicula
+
+ ¡ ++ ± ±
----+-
+ - - + +-G
+ I + + +
Creseis chierchiae + + +
+
+
+
+
+
4+1~'~:
+
+
- -
± +
+
+
+
+
+
+
+
+
+
+ +
Creseis virgula forma conica + +
Creseis virgula forma constricta + + + + + + + + +
Creseis virgula forma virgula + + + + ++ + + + + + +
Hyalocylis striata + + + + + ± + + + +
Styliola subula + + + + + ++ + + + + + + +
Cuvierina columnella
+ + ± + I - + + +
forma atlantica -[
Suborder Pseudothecosomata
Family Cymbuliidae
Cymbulia parvidentata + + + + + + + + + +
668 Pteropoda
Cymbulia peroni + + + + + +
morpha peroni -
[
-1
Cymbulia peroni 1- +
+ + + + + + + +
morpha minar
Cymbulia sibogae + + + + + + + + + +
Corolla calceola + + + + + + + + + +
Corolla cupula + + + +
+
Corolla intermedia + + + + + + + + + +
Corolla ovata + + + + ± ± + + + + + +
Corolla spectabilis + + + + + + +
Gleba cordata + 1: + + + + + +
Family Desmopteridae
Oesmopterus papilio + + + + + + + + +
Family Peraclididae
Peraclis apicifulva + + + + + ± + + +
Peraclis bispinosa + + + + + +
Peraclis depressa + + + + + +
Peraclis moluccensis + + + + + + +
Peraclis reticulata + + + + + + + +
Peraclis triacantha + + + + ±
:1:
+
: + + +
Peraclis valdiviae + + + + + + ± + + + + + +
Order Gymnosomata
Family Clionidae
Clione limacina antarctica + ++ - + + + + + + + +
Paraclione longicaudata + + + + + + + + + ±
Cephalobrachia macrochaeta + + + + + + + + + + +
Massya longicirrata + + + + + + + +
Thliptodon antarcticus + + + + + + + + + + +
I
Thliptodon diaphanus + + + + -1 + + + + + + + + + +
Family Cliopsidae
C1iopsis krohni + + + + + ++ +++ + + +
morpha grandis
C1iopsis krohni + + + + + ++ +++ + + +
morpha krohni
C1iopsis krohni + + + + + ++ +++ + + +
morpha modesta
Family Notobranchaeidae
Notobranchaea macdonaldi
+ + + + + ± + + + + + + +
morpha macdonaldi
Pteropoda 669
l'O
..D
_
ra.~
n3
Q)
U
.~ u....;
U ..:t: re tl.O u
.~
tl.O 0.0
.~
~ .~ Q.. (U ~ u . Vl ~ 'i: ~ e '00 ~ ~

::J .- •..... ..o Q. ra ••.. O .- > tl.O ;:; e n:l ~ Q) O >- e .•...
O~~~~~~~
E G
u g- g E
Q. 15[:l a> I .D
~ ~
"13 eU I ~~~~
.~
N -[ ~
c::
s~5
oS ~ ~ c:: ~
~]~
"'·_·üe I
.g ~ :fr~~
~Q. ~ ~ O~
u.~
Q).¡:
v>..c::
Notobranchaea
+++ + + ± ±+ +++ ++ +
tetrabranchiata
Family Pneumodermatidae
Pneumoderma atlanticum
+++-± + + ++ +++ ++ +
atlanticum forma atlanticum
atlanticum forma + + + ± + + + +
Pneumoderma atlanticum I
eurycotylum ++ I + -+ -± - + + I +-1- +
atlanticum forma pygmaeum
+- + -±-- + + ++ +
boasi +
++ ± + ++ ++ + +
souleyeti •
Pneumoderma mediterraneum + + - + - + + + + + + +
Pneumodermopsis
(Crucibranchaea) macrochira +++±++ + I ±±¡- +++ ++ +

Pneumodermopsis
(Pneumodermopsis) I + + + - I - +
brachialis I
Pneumodermopsis
(Pneumodermopsis) paucidens + + ± + + + + + + + + + +
forma paucidens I
Platybrachium antarcticum - + + + +
Schizobrachium polycot~
SfJonjjiobranchaea australis
++
+
+
+
~+
- T
+
abundant
- +
± ++
+
+ +-- +
~ +
+- ~ +,
-~ -
- +
+ r- - +
+ -
+
+.
+ +
+ +
+
+] + + t --
+
-
+
+ -
+ present
± moderate presence
traces
absent
Table 1. World wide biogeographic affinities of pteropod species.

670 Pteropoda
In Cymbulia peroni size variations mark the difference 1), on top of which a conical section develops;
between the Atlantic and Mediterranean populations soon afterwards the conical section broadens and a
(Fumestin, 1979). In soft-bodied Gymnosomata these transverse slit starts to separate the dorsal from the
relatively minor variations are more difficult to assess. ventral sides. All soft parts migrate into this
bivalve-!ike shell and the embryonic shell is shed;
the opening left at the base ofthe bivalve-like sec-
Glossary of terms used in descriptions of tion is closed by bending the ventral and dorsal
Pteropoda (see Fig. 1) walls and secreting additional shell material, but
no septum is formed; this fusion gives rise to the
Aperture (=shell aperture): the opening of the shell caudal joint (Fig. lB).
through which the animal can expand (not to be Caudal septum (=caudal spine mark, closing mem-
confused with the mouth of the soft body) (Fig. brane): the membrane closing off the adult shell
lA, C, H, K). when the protoconch is discarded (Fig. 1H).
Apex (apical): the top of the shell formed by the Oral cavity: the depression in which the mouth is
embryonic shell (Fig. lA, K). located (see also buccal mass, buccal cone, sucker
Apex angle (=top angle): the ang1e between the late- arm).
ral shell sides enclosing the embryonic shell, in Central depression: see median lip depression.
side view (Fig. 1K). Central dorsal rib: undular elevation ofthe dorsal shell
Arm (=acetabulous or acetabu!iferous arm, sucker side usually running from the rostrum to close to
arm): in the Pneumodermatidae, two lateral and the caudal joint, mark or embryonic shell (Fig. 1C).
sometimes one median arm on the wall of the buc- Ciliar band (=ciliary band): transverse circular band
cal cavity, covered with stalked suckers. Arms can around the body in juvenile Gymnosomata; usual-
retract completely into the hood over the head and Iy one band surrounds the head parts, one lies in
proboscis (Fig. 1L). the middle of the body, and one around the poste-
Balancer: organ with uncertain function at the mantle rior end (Fig. 10).
border in Thecosomata (Fig. lB). Closing mechanism: see lock system.
Basal plate: underside of a radular tooth from which Closing membrane: see caudal septum.
the cusp or dent of the tooth projects (see radula) Columella: the axis of the coiled shell of the
(Fig. 1G). There can be one, two, three, or more Limacinidae to which the columellar muscle is
cusps on one basal plate, defining unicuspoid, attached and that also forms the inner aperture bor-
bicuspoid, tricuspoid or multicuspoid teeth. der where a thin membrane, the columellar mem-
Bicuspoid: see basal plate. brane, is formed in some species (Fig. 1A, K).
Buccal cone: wall structure in the buccal cavity with- Cone filaments: filaments at top of the buccal cone.
out suckers, with a sticky glandular epithelium; Constriction: usually applied to (a) narrow area
these eversible cones can be covered by the hood between protoconch I and II (Fig. IR); (b) the nar-
over the head (Fig. 1M). rowing ofthe aperture by two dents or lateral folds
Buccal mass: the radula, jaws, hooks and musculature of the dorsal !ip near the base of the rostrum in
of the mouth or buccal cavity (Fig. 1L). Cavo!iniidae.
Bursa seminis: sack-like evagination of the sexual Crests: (a) ridges on the soft body, especially those
duct storing autosperm. forming the gills (Fig. 1L); (b) calcareous ridges
Callus (=calcareous ridges): extra calcareous secre- on the shell or ridges on the whorl attached by
tion, usually of irregular nature, over the teleo- means of broad bases to the preceding whor!.
conch near the aperture or near the caudal spine, as Ctenidium: see gil!.
in Cavolinia (Fig. 1C). Dorsal !ip: see rostrum.
Caudal fold: part of the caudal margin of the shell Dorsal ribs: longitudinal elevations of the dorsal side
not occupied by spines or the caudal joint, of the shell in Cavoliniidae; the central one is fre-
formed by an uninterrupted shell wall where the quently referred to as central rib (Fig. 1C, F).
ventral and dorsal sides merge gradually, espe- Dorsal side (=dorsal valve): in molluscs, the side
cially in Cavoliniidae (Fig. lB). opposite to the side where the mouth or foot are
Caudal joint (=closing fold): Diacavolinia first deve- located (Fig. 1F).
lops a bowl-shaped embryonic shell (protoconch Embryonic shell: see protoconch.
Pteropoda 671
Epipodial tentacle: the tentacle inserted into the rim of Lateral line: line between the lateral side of the aper-
the wing disc (e.g., Desmopterus) (Fig. 1I). ture and the protoconch or caudal septum; line
Epipodium: tissue or organs directly surrounding the between spine and lock area visible in ventral
ontogentic foot. view (Fig. 1C).
Flank: in Cavoliniidae, the somewhat flattened lateral Lateral rib (latero-dorsal rib): undular elevations of
side of the ventral surface of the shell (Fig. lB). the dorsal shell side at both sides of the central rib
Fold: width of the fold is defined as the distance (Fig. IC, F).
between joint and gutter comer in Cavoliniidae Lateral spine: shell projections at the comers of the
(Fig. lB). shel! aperture, projecting strongly laterally in
Foot (=pedal fin, wing): in pteropods the molluscan some species (Fig. 1C).
foot is transformed into two broad flaps used in Lip angle: in cavoliniid shells the angle between the
swirnming, called wings, plus the lips and small line through the tip ofthe dorsal aperture lip ofthe
footlobes (Fig. lB, 1, J, L, M). shell and lock tubercle and the line through the
Footlobe (posterior footlobe, lateral footlobe): parts of two lateral spine tips of the shell (Fig. IF).
the molluscan foot that are transformed into wings, Lip: (a) folds around the mouth ofthe Thecosomata in
situated around the mouth (Fig. lB, 1, J, L, M). Pseudothecosomata these lips form a proboscis
Fringed: bordering ofthreads either loose or formed into structure (Fig. lJ); (b) also used for the ventral and
tassels and twists, especially used for describing the dorsal shell parts at the aperture in Cavo!iniidae
appearance ofthe gil! in Gyrnnosomata (Fig. IL). (Fig. IC).
Gil!: in Gyrnnosomata, lateral and posterior appen- Lip belly: outward bulge of the dorsal !ip m
dages function as gills, real ctenidia are absent in Cavo!iniidae (Fig. lB).
Pteropoda (Fig. lL). Lip flap: inwardly bent part of the dorsal !ip 1ll
Gizzard plate (=gizzard tooth): chitinous plates on the Cavoliniidae (Fig. lB).
wal! of the gizzard of some Opisthobranchia. Lip gutter: in Cavoliniidae, the longitudinal groove of
Gullet bladders: a pair of closed bladders in the buc- the dorsallip continuous with the dorsal rib (Fig.
cal mass looking like inflated, curved sausages. lB).
Gutter comer: in Cavoliniidae, the posteriormost cor- Lip rim: margin ofthe lips ofthe shell aperture, either
ner of the shell aperture through which the balan- sharp or thickened; usually the ventrallip is thick-
cers can emerge (Fig. lB). er than the dorsal one (Fig. I C).
Gutter: a hollow groove (see gutter comer, lip gutter) Lip shoulder: part of the dorsal side of the lip visible
(Fig. lB). when the shel! is in ventral view, as in Fig. 1C.
Head: in molluscs the head bears the buccal mass, Lock angle: in cavoliniid shells the angle between the
eyes, and tentacles (Fig. IL). line joining the right end of the joint and right lock
Hook (hook sac): chitinous teeth located in evaginable tubercle and the line through the two lateral spine
sacs (hook sack) at both sides of the radula, under tips (Fig. lB).
the hood (probably modified parts of the jaw), Lock area: in Cavoliniidae, the area near the comers
present in most Gyrnnosomata (Fig. IL). of the ventrallip, where the dorsal and ventral side
Hump: in profile, the strong bending ofthe dorsal side are connected with the main closing mechanism
where it goes over into the dorsal !ip in (Fig. IC).
Cavoliniidae; the outside is sometimes called the Lock link: hook on the inner ventral shell surface, fit-
outer hump, the bend visible on the inside of the ting between the lock tubercles in Cavo!iniidae
aperture is the inner hump; both are frequently (Fig. ID).
used as diagnostic characters (Fig. IF). Lock rib: rib on the dorsal side above the lock area,
Hypobranchial gland: see mantle gland. usually not extending far posteriorly and com-
lnner hump: see hump. posed mainly of the thickening under the lock
Jaw: chitinous plate in the oesophagus opposite the tubercles (Fig. IF).
radula (Fig. IL). Lock system: system of one (primary) or two (when a
Joint: width of the joint is defined as the distance secondary system is present) pairs of locking
between the two fold areas in Cavoliniidae (Fig. structures fixing the position of ventral and dorsal
lB). shel! valves in Cavoliniidae (Fig. ID, E).
672 Pteropoda
Lock tubercles: knobs on the inside of the shell wall, Periostracum: a chitinous layer of material covering
visible from the outside; a total of three tubercles the calcareous shell.
may be present (main, lunar, minor tubercles visible Pharynx: sometimes used in molluscs to designate
from inside), they constitute the closing mechanism mouth and esophagus with adjacent organs.
(Fig. lE). Posterior footlobe: see median footlobe.
Mantle gland (=hypobranchial gland): mucus-secreting Proboscis: usually elongated part of the mouth and
gland present in the mantle lining of many molluscs. esophagus with or without buccal organs (Fig. lL).
Mantle (mantle band): extension ofthe body wall that Protoconch 1 (protoconch n, embryonic shell): proto-
secretes the shell and lines most of the shell conch 1 is the shell formed in the egg; protoconch n
(absent in Gyrnnosomata). is the shell formed during the veliger stage, together
Median footlobe (=posterior footlobe): the central they comprise the embryonic shell (Fig. lA, C, H).
part of the foot, located between the lateral foot- Pseudoconch (=slipper): in Cymbuliidae, the calcare-
lobes (Fig. lB, 1, J, L, M). ous shell is lost and replaced by a permanent carti-
Median lip depression (=central depression): the con- laginous, slipper-shaped structure: the pseudoconch
cave central part ofthe ventrallip in Cavoliniidae. (Fig. 1J). Lalli and Gilmer (1989); also described as
Median tubercle (=median footlobe tubercle): in pseudoconch formations in Euthecosomata, but it
Gyrnnosomata, the knob between the lateral foot- seems not to be a permanent structure in this group.
lobes, probably used for fixation during copula- Radula plate (=radula teeth): components ofthe radu-
tion (when a median tubercle is present the penile la, sometimes "first plate" or "second plate" are
sucker is usually absent) (Fig. lL). referred to, in which case numbering starts in the
Minute stage: a developmental stage in several centre of the radula (Fig. 1G).
Thecosomata in which the shell is fully grown Radula: series of chitinous teeth in the buccal mass used
while the soft parts are still very small; when the in catching prey (Fig. lG, L, M, N). The first, or the
soft parts are very slender and long it is called additional last lateral teeth are sometimes different
skinny stage (Fig. IR). from the others. The additional (a), lateral (1), modi-
Nose: small thickening visible in lateral view near the fied lateral (mI) and median (m) teeth are expressed
tip of the dorsallip in some Cavoliniidae (Fig. l F). in a radula formula as: (a)-I-(ml)-m-(rnl)-I-(a); some
Odontophore: stiff structure supporting the radula examples of radula formulae are: Limacina: 1-1-1;
(Fig. l L, M). Cavolinia: 1-1-1; Peraclis: (1)-1-1-1-( 1); Pneumo-
Operculum (operculum formula): plate closing offthe dermopsis: 6-1-6; Pneumoderma: 1-0-1; Thlipto-
shell aperture when the animal is retracted; it con- don: 3-(1)-1-(1)-3; and Clione: 15-1-15.
sists ofwhorls around a nucleus or gyre (Fig. lK). Rhinophore (=oral tentacle): pair of tentacles on the
The operculum formula is LG:H:W:D:E, the let- head; when an additional pair is present it is called
ters standing for greatest operculum diameter : oral tentacle (Fig. 1M).
greatest gyre diameter : greatest last whorl diame- Rostrum: (a) anterior part of the dorsal shell Iip pro-
ter : smallest operculum diameter : smallest gyre jecting as a snout in Thecosomata (Fig. lB). In
diameter : smallest last whorl diameter. some species it is clearly separated from the rest of
Oral cavity: the depression in which the mouth is locat- the dorsallip; this part ofthe shell hosts many dis-
ed (see also buccal mass, buccal cone, sucker arm). criminating characters, especially in Cavoliniidae;
Oral tentacle: see rhinophore. (b) also used for the basal extension of the colu-
Outer hump: bulge of dorsal side at the base ofthe lip mella in coiled forms (Limacina, Peraclis) (Fig.
in Cavoliniidae (Fig. lF). lA, K). The length of the rostrum is equal to the
Palps: finger-like extensions. length of the dorsal lip between rostrum tip and
Pedal: related to the foot (e.g., pedal ganglion, pedal inner hump (in Cavoliniidae). In species without a
gland, pedal wing). hump the length cannot be measured exactly,
Penile sucker: sucker at the base ofthe penis (see also though the rostrum may be presento
median tubercle) (Fig. 1M). Second lock: usually a simple extra lock mechanism
Peribuccal (=additional) denticules: teeth placed between the ventral and dorsal shell sides, at the
around the mouth as found in Notobranclzaea upper side of the lateral spines in Cavoliniidae
(probably related to the jaw). (Fig. lB, C).
Pteropoda 673
Side angle: in Thecosomata, angle between the line Ventrallip base: ridge between the ventrallip and the
through the lateral spine tips and the line through ventral shell surface in Cavoliniidae (Fig. lC).
the right spine tip that contacts the left lip belly Ventral lip: prolongation of the ventral side of the
(Fig. lB). shell over the aperture in Cavoliniidae (Fig. l C).
Skinny stage: see minute stage. Ventral rib: transverse sculpture lines covering the
Spine (lateral spine, caudal spine): lateral spines are ventral side, usually congruent with, but not
lateral prolongations of the shell near the aperture homologous with, the growth lines; they are com-
comers. They differ in shape among species and posed of tubercles that may coalesce into a single
are often bent dorsally. The caudal spine projects ridge or adopt a comb-like pattem.
posteriorly, it is straight or bent and in some spe- Visceral nucleus (=visceral mass): liver, gonad, heart,
cies discarded in the adult (Fig. IC, F, H). sexual glands and kidneys grouped together (Fig.
Spine mark: marks left on the adult shell after the 11).
spines are shed off (Fig. IH). Whorl (body whorl, embryonic whorl): a coiled shell
Spire (=spira): (a) narrower whorls forming the centre is made of one or more coils, each coil is called a
of the shell and surrounded by the body whorl, whorl; the last one is the body whorl, the first ones
usually including the protoconch whorls plus a are the embryonic whorls or protoconch whorls
few small teleoconch whorls (Fig. lA); (b) narrow (Fig. lA, K).
whorls of the operculum.
Striae (striation): sculpture lines on the surface of the Outline c!assification
shell; very prominent striae are sometimes called
ribs (Fig. IC). Striae can be due to growth lines The listing below covers all Recent pteropods
(transverse striation) or to sculpture (longitudinal described to date down to the genus leve!. Taxa
or sometimes transverse striation) (Fig. lA). recorded in the South Atlantic and included in the
Sucker (lateral sucker, median sucker): suckers present review are underlined.
present on the acetabuliferous arms, those on the
lateral arms are lateral suckers, those on the Phylum Mollusca
median arm are median suckers; used for captu- Subphylum Aculifera
ring prey (Fig. lL). Sometimes the penis has a Class Gastropoda
sucker, when absent, a median footlobe tubercle Subclass Euthyneura (=Opisthobranchia)
takes its function. Order Thecosomata Blainville, 1824
Suture: the boundary line between two whorls, as seen Suborder Euthecosomata Meisenheimer,
on the surface of the shell (Fig. I A, K). 1905
Teleoconch: shell section formed after the juvenile Family Limacinidae Gray, 1847
developmental stage (Fig. lA, B, C, H). Genus Limacina Bosc, 1817
Tentacle (dorsal tentacle, anterior tentacle): usually the Family Cavoliniidae Fischer, 1883
tentacles are finger-shaped outgrowths ofthe body; Subfamily Cavoliniinae van der Spoel,
the eyes in pteropods are located on the dorsal or 1972
nuchal tentacles in the neck region. The term tenta- Genus Cavolinia Abildgaard,1791
cle is also used for the two tubercles found on top (emend. Philippi, 1853)
of the hood covering the head (anterior tentacles). Genus Diacavolinia van der Spoel,
Umbilical keel: the ridge running around the umbili- Bleeker and Kobayasi, 1993
cus in some limacinid species. Genus Diacria Gray, 1847
Umbilicus: hollow canal or opening on the underside Subfamily Clioinae van der Spoel, 1972
of the shell in the centre of the spire (Fig. lA). Genus Clio Linnaeus, 1767
Umbo: top ofthe (coiled) shell, related to the top (Fig. Genus Creseis Rang, 1828
lA). Genus Hvalocvlis Fol, 1875
Valve angle: in lateral view of Cavoliniidae, angle Genus Styliola Gray, 1850
between the dorsal and lateral shell sides. Subfamily Cuvierininae van der Spoel,
Veliger: larval stage, following the trochophore, cha- 1967
racterized by presence of a velum. Genus Cuvierina Boas, 1886
674 Pteropoda
Suborder Pseudothecosomata Meisenheimer, Genus Spollgiobrallchaea d'Orbigny,

1905 1836
Family Cymbuliidae Cantraine, 1841
Subfamily Cymbuliinae Cantraine, 1841 Diagnoses and identification of species
Genus CVlIlbulia Peron and Lesueur,
1810 Adaptations of pelagic molluscs to floating behaviour
Subfamily Glebinae van der Spoel, 1976 hinder construction of taxonomic identification keys
Genus Corolla Dall, 1871 for the Pteropoda. Taxa included below (listed alpha-
Genus Gleba (Forskal, Ms.) Niebuhr, betically within each family) are those effectively
1776 recorded or expected to occur in the South Atlantic
Family Desmopteridae Chun, 1889 Ocean. Characters and measurements cited refer to
Genus Deslllopterus Chun, 1889 adult individuals, unless otherwise noted. A compu-
Family Peraclididae Tesch, 1913 terized identification program for pelagic molluscs is
Genus Peraclis Forbes, 1844 (emend. available from ETI (ETI-e/o Institute of Taxonomic
Pelseneer, 1888) Zoology, P. O. Box 4766, 1009 AT Amsterdam, The
Order Gymnosomata de Blainville, 1824, Netherlands; van der Spoel et al., 1997).
emend. van der Spoel, 1976
Suborder Gymnoptera van der Spoel, 1976 Kingdom Animalia
Family Hydromylidae Pruvot-Fol, 1942 Phylum Mollusca
Genus Hydrolllyles Giste!, 1848 Subphylum Aculifera
Family Laginiopsidae Pruvot-Fol, 1926 Class Gastropoda
Genus Lagilliopsis Pruvot-Fol, 1922 Subclass Euthyneura (=Opisthobranchia)
Suborder Gymnosomata de Blainville,
1824, (senslI van der Spoel, 1976) Order Thecosomata
Family Clionidae Gray, 1840 Usually shelled pteropods.
Subfamily Clioninae Pruvot-Fol, 1924
Genus Cliolle Palias, 1774 Suborder Euthecosomata
Genus F owlerill a Pelseneer, 1906 Always shelled, radula formula 1-1-1.
Genus Paedocliolle Danforth, 1907
Genus Paracliolle Tesch, 1903 Family Limacinidae
Genus Tltalassopterus Kwietniewski, Shell coiled, sinistral, with operculum, mantle-cavity
1910 on the dorsal side. Only one genus.
Subfamily Thliptodontinae Pruvot-Fol,
1926 Genus Lilllacilla
Genus Cepltalobracltia Bonnevie, 1913 Three subgenera: Limacina with 2 species, Munthea
Genus Massva Pruvot-Fol, 1924 with 3 species, and Thilea with 3 species.
Genus Thliptodoll Boas, 1886 o Lilllacilla (Lilllacilla) helicilla (Phipps, 1774) all-
Family Cliopsidae Dall, 1889 tarctica (Woodward, 1854) forma alltarctica Wood-
Genus Cliopsis Troschel, 1854 ward, 1854 (Fig. 6.1). Spire very depressed, about 6
Family Notobranchaeidae Pelseneer, 1886 colorless whorls. No umbilical keel, umbilicus wide.
Genus Notobrallcltaea Pelseneer, 1886 Transverse striae prominent, best visible on last
Family Pneumodermatidae Pelseneer, 1887 whorl. Faint spiral striation on first whorls. Wing
Genus Abrallchaea Zhang, 1964 protrusion presento Maximum diameter: 6.5 rom.
Genus Platybracltiulll Minichev, 1976 o Lilllacilla (Lilllacilla) Itelicilla (Phipps, 1774) all-
Genus PlleulIloderllla Dumeril, 1806 tarctica (Woodward, 1854) forma rallgi
Genus PlleulIloderlllopsis Keferstein, (d'Orbigny, 1836) (Fig. 6.2). Spire depressed, with
1862 about 6 colorless whorls, higher than forma antarc-
Genus Platybracltiulll Minichev, 1976 tica. Umbilicus wide, umbilical keel, spiral striation
Genus SchizobracltiulIl Meisenheimer, and transverse striation faint or lacking. No wing
1903 protrusion. Embryonic shell with rough omamenta-
Pteropoda 675
tion of irregularly rarnified, flat ridges (Boltovskoy, (involute). Last whorl swollen, aperture heart-
1974b). Shell diameter: 3.5 rnm, height: 2.0 rnm. shaped with blunt tooth extending on last whorl as
o Lilllacina (Lilllacina) retroversa (Fleming, 1823) faint keel or ribo Rostrum, or aperture tooth, vari-
allstralis (Eydoux and Souleyet, 1840) (Fig. 6.3). able in shape. Umbilicus deep. Embryonic shell
Spire highly coiled, more than 6 colorless whorls. with rough granulated surface both on proximal
No umbilical keel. Umbilicus wide, but may be zone and first whorl (Boltovskoy, 1974b).
narrow in young specimens. Transverse striae Maximum diameter: 1.5 rnm.
presento Wing protrusion presento Embryonic shell o Lilllacina (Thilea) lesllellri (d'Orbigny, 1836)
with rough ornamentation of irregularly ramified (Fig. 6.8). Spire depressed, broader than high, body
flat ridges (Boltovskoy, 1974b). Shell height: 2.5 whorl increasing very rapidly in diarneter. Up to 4.5
rnm, maximum diameter: 1.5 mm. colorless whorls. Umbilicus narrow but deep, spiral
o Lilllacina (Munthea) blllillloides (d'Orbigny, striation around umbilicus, rest of shell smooth.
1836) (Fig. 604). Blunt spire highly coiled, more Inner border of aperture straight, suture very well
than 6 colorless whorls. Thicker parts of shell near marked. Wing protrusions absent. Embryonic shell
aperture brownish, smooth except for faint growth with rough granulated surface (Boltovskoy, 1974b).
lines. Conspicuous suture sometimes light-brown. Shell height: 0.8 rnm, diameter: 13 rnm.
Thickened inner aperture chestnut-brown.
Umbilicus very small. Small rostrum more point- Family Cavoliniidae
ed in older than in young specimens. Protrusions Shell not coiled, primitive forms with straight tubular
on anterior wing border absent. Embryonic shell shells, more highly evolved ones with pyramidal
with rough ornamentation of irregularly ramified shells or shells composed of two valve-like halves;
flat ridges (Boltovskoy, 1974b). Shell height: 2 soft parts show secondary bilateral syrnmetry; colu-
rnm, diameter: lA mm. mellar muscle is attached to inner surface of proto-
o Lilllacina (Mllnthea) trochiforlllis (d'Orbigny, concho
1836) (Fig. 6.5). Spire high, body whorl increasing
rapidly in diameter. With up to 5, rapidly growing Subfamily Cavoliniinae
colorless whorls. Narrow deep umbilicus. Thicker Dorsal and ventral surfaces different and well separat-
parts along columella and upper aperture lip, pur- ed, dorsal aperture border curved.
ple-brown, smooth except for faint growth lines.
Clear suture. Columellar aperture border slightly Genus Cavolinia
con cave. Wing protrusions absent. Embryonic shell Members of this genus either retain the protoconch or
with spiral ornamentation of irregularly ramified do not form a closing septum if the latter is shed.
flat ridges, depression separating protoconch 1 and Cavolinia gibbosa (d'Orbigny, 1836) formajlava
11(Boltovskoy, 1974b). Protoconch I smooth, pro- (d'Orbigny, 1836) (Fig. 6.9). Shell globular, trans-
toconch 11 with growth lines. Shell height: l rnm, parent, with dark brown color patches. Ventral
width: 0.8 rnm; apical angle: 75-96°. side vaulted, width <70% of length. Lateral spines
o Lilllacina (Thilea) helicoides Jeffreys, 1877 (Fig. small and slightly curved caudally. Caudal spine
6.6). Shell large, thick, dark chestnut brown, curved dorsally in posterior parto Aperture narrow,
depressed, with 3-4 whorls ofwhich the last one is with faint harnmered sculpture on the ventral sur-
swollen and composes most of the shell. Narrow face. Embryonic shell usually present, its junction
umbilicus, sometimes absent. Inner aperture mar- with rest of shell smooth. Shell length: 5.0 rnm,
gin curved outwards. Spiral punctuated lines on width: 4.5 rnm.
first whorl and embryonic shell, transverse growth Cavolinia gibbosa (d'Orbigny, 1836) formagibbo-
lines on last whorl. Body dark brown. Wings sa (d'Orbigny, 1836) (Fig. 6.10). Shell globular,
fleshy. Small juveniles slightly lighter in color and transparent, relatively wide (about 68% of length);
occurring higher in the water column than adults. ventral vault strongly protruding (width about 71%
Protoconch 1 covered with pyramidal warts, proto- of length). Regularly convex ventral vault does not
conch 11smooth. Shell height: 11 mm. show forma jlava-like edge with sharp topo
o Lilllacifra (Thilea) injlata (d'Orbigny, 1836) (Fig. Anterior part of ventral side perpendicular to shell
6.7). Shell with up to 3 whorls coiled at one level axis, and may even be slightly elevated. Lateral
676 Pteropoda
spines small, caudal spine and upper shell lip pro- Shell margin between caudal and lateral spines
nounced. Juvenile shell starts development as blunt concave, its upper stretch more or less straight and
oval protoconch 1 with fine transverse striations. horizontal. Growth lines faint, transverse striation
Only an irregularity in growth separates proto- poorly developed. Tips of lateral spines directed
conch 1and n. Shelllength: 8 rnm, width: 5.5 rnm. slightly caudally. Caudal spine short and straight.
Cavolinia inflexa (Lesueur, 1813) forma imitans Well-developed arch-shaped rib at base of dorsal
(Pfeffer, 1880) (Fig. 6.11). Shell transparent with lip. Shell length: 16 rnm, width: 10.5 rnm.
flat dorsal side and slightly convex ventral side. Cavolinia tridentata (Niebuhr, 1775) forma allS-
Caudal portion of shell behind lateral spines short- tralis (Péron, 1807) (Fig. 6.15). Shell hyaline or
er than in forma inflexa and more strongly curved opaque, with flat dorsal side and moderately deve-
dorsally; otherwise quite similar to the latter. loped ribs. Ventral side moderately vaulted, with
Sculpture of faint growth lines. Lateral spines sculpture offaint growth lines and very faint trans-
well-developed, projecting from middle of shell, verse striation. Shell margin between caudal and
not bending as far caudally as in forma labiata. lateral spines irregularly concave. Strong, whitish
Caudal spine long. Upper aperture lip rather large, keels along lateral sides. Tips of lateral spines
separated from rest of ventral side by conspicuous directed caudally. Caudal spine straight. Dorsal
incision. Shelllength: 8 mm. aperture lip separated from dorsal side by promi-
Cavolinia inflexa (Lesueur, 1813) forma inflexa nent arch-shaped rib between closing mechanisms,
(Lesueur, 1813) (Fig. 6.12). Dorsal side flat, ven- strongly truncated, usually ending in two anterior
tral slightly convex; posterior section of shell projections. Shell aperture trapezoidal, with brown
bending dorsally. Small vault of ventral side ends to dark brown or purple pattem over the ventral
in transverse groove under ventral lip. Transverse and dorsal sides; dorsal aperture lip always lighter
ribs absent on ventral side. Sculpture of faint in color. Shelllength: 16 mm, width: 10.5 rnm.
growth lines present. Lateral spines poorly deve- Cavolinia uncinata (Rang, 1829) lmcinata (Rang,
loped, located well above middle of shell. Caudal 1829) forma lmcinata (Rang, 1829) (Fig. 6.16).
spine very long, undifferentiated from rest of Colorless shell with flat dorsal side provided with
shell. Embryonic shell sometimes present; when five longitudinal ribs and two smaller ribs border-
absent shell closed by callus resembling a rounded ing posterior slits of aperture. Ventral side clearly
embryonic shell. Embryonic shell defined by faint vaulted, with transverse ribs, especially near aper-
constriction which separates it from teleoconch. ture. Ventral side depressed medially near aperture;
Juveniles dorso-ventrally compressed, pyramid- this depression is also found ventrally under the
shaped in ventral view, with oval aperture. Shell curved lip. No keel along lateral sides. Sculpture of
length: 7 rnm, width: 5 rnm. faint growth lines and transverse striation near aper-
Cavolinia inflexa (Lesueur, 1813) forma labiata ture. Ventral surface with faint harnmered sculp-
(d'Orbigny, 1836) (Fig. 6.13). Shell broader than ture. Lateral spines almost straight. Caudal spine
in other forms (width 80% of length). Dorsal side retained in adults, curved, relatively long. Dorsallip
flat, ventral side somewhat convexo Section curves strongly over aperture resembling a hood.
behind lateral spines is smaller than in other forms Protoconch usually broken off, leaving open place
(about 36% of the total shelllength; vs 50% in C. of rupture. Shelllength: 7.5 rnm, width: 6.6 rnm.
inflexa forma inflexa, and 40% in C. inflexa forma
imitans). Sculpture of faint growth lines. Lateral Genus Diaeavolinia
spines well-developed and distinctly bent caudal- These species discard the protoconch at a very early
ly. Caudal spine long. Upper lip continuous with stage and close the shell, merging the ventral and dor-
dorsal side of shell, rather long, distinctly tapering sal shell sides.
into sharp tipo Shelllength: 8 rnm, width: 5.5 rnm. • Diaeavolinia angulosa (Eydoux and Souleyet, Ms.)
Cavolinia tridentata (Niebuhr, 1775) forma atlan- (Gray, 1850) (Fig. 6.17). Shell transparent, triangu-
tiea van der Spoel, 1974 (Fig. 6.14). Shell with flat lar in outline. Ventral side vaulted, with 18 thin and
dorsal side and moderately developed ribs, ventral straight ventral ribs; anterior part without ribs.
side moderately vaulted. Light-brown hue over Dorsal side convex, with well-developed central rib
nearly entire shell. No keel along lateral sides. and latero-dorsal ribs; lock ribs small but rather
Pteropoda 677
thick, lock area well-developed. Characteristic nose absent. Broad dorsallip with notch and gutter, but
in middle of the rostrum; owing to strong inner no constriction. Both ventral and dorsal lips with
hump, which is continuous with underside of lip thickened rim, ventral lip moderately sized with
shoulder, central dorsal rib not continuous with ros- weak median depression. Traces of lip flaps some-
trum. Gutter comers small. Sculpture offaint growth times present; well-developed lip bellies, lip
lines and faint transverse striation. Lateral spines shoulders very smal!. Hump absent. Protoconch II
perfectly straight, directed laterally, hooked and area not strongly projecting. Shelllength: 7.8 mm
with sharp tips (in some populations lateral spines and width: 7.8 mm.
may be reduced). Caudal spine absent, no caudal Diacavolillia constricta van der Spoel, Bleeker
spine mark since ventral and dorsal sides merge. and Kobayasi, 1993 (Fig. 6.20). Shell hyaline,
Dorsal lip with notch, gutter with inner hump, no subquadrangular in outline, with brown hue over
constriction. Ventrallip moderately sized with clear entire surface. Ventral side vaulted. Nineteen con-
median depression. Shelllength: 4.8, width: 3.7. spicuous, comb-shaped ventral ribs. Dorsal side
Diacavolinia atlantica van der Spoel, Bleeker and flat with moderately developed central and latero-
Kobayasi, 1993 (Fig. 6.18). Shell hyaline, triangu- dorsal ribs; lock ribs moderate, lock area large
lar in outline. Brown hue over entire shell surface. with large main tubercle, large link, large lunar,
Ventral side vaulted, 39 thick, comb-shaped ven- and small minor tubercle. Second lock system
tral ribs, flanks not pronounced, upper part ofven- present. Laterallines slightly convex, flanks clear-
tral side flat in outline. Dorsal side slightly con- ly developed. Growth lines typical; sculpture ribs
vex, with well-developed central and lateral ribs; on dorsal side. Lateral spines small, not bent,
lock ribs small but strongly developed. Lateral placed relatively far anteriorly, hooked but not
lines convex; lock area well-developed, with large very sharp at tipo Gutter corners smal!. Caudal
main, lunar, and small minor tubercle and link. spine and caudal spine mark absent. Ventral lip
Second lock system present at upper level of large with weak median depression. Strongly bent
attachment of spines to lateral lines. Sculpture of dorsal lip with notch in relatively broad rostrum,
weak growth lines and very faint transverse stria- constriction and gutter present. No lip flaps; large
tion. Lateral spines well-developed, slightly bent, lip bellies and strong lip shoulders, aperture
relatively long, sharp, hooked; gutter corners extremely narrow. Hump absent. Protoconch II
large. Caudal spine and caudal spine mark absent. area not projecting, with moderate caudal joint.
Dorsal lip with notch and constriction in broad Shelllength: 5.6 mID, width: 5.3 mID.
gutter-shaped rostrum. Relatively small ventrallip Diacavolinia deblainvillei van der Spoel, Bleeker
with distinct median depression. No lip flaps; lip and Kobayasi, 1993 (Fig. 6.21). Shell colorless,
bellies well-developed, lip shoulders moderately hyaline. Dorsal side convex, central rib and latero-
developed. Hump absent. Central part of wings dorsal ribs well-developed, lock ribs well-deve-
intensely brown. Protoconch Il projecting slightly. loped but smal!. Ventral ribs, 23, strongly deve-
Shelllength: 10.9 mm, width: 9.6 mID. loped, comb-shaped. Flanks present. Lock area
o Diacavolinia bicornis van der Spoel, Bleeker and small, main and lunar tubercles normal, minor
Kobayasi, 1993 (Fig. 6.19). Shell hyaline, triangu- tubercle absent. Second lock system present.
lar. Brown hue especially on lower ventral and Sculpture of faint growth lines and faint transverse
dorsal sides. Ventral side vaulted. Thin ventral striation. Short lateral spines bent dorsally, hooked
ribs straight. Dorsal side convex, with well-deve- and sharp. Caudal spine and caudal spine mark
loped central and lateral ribs. Laterallines slightly absent. Dorsal lip with notch but no constriction;
convex, flanks not very clear. Lock rib moderate- gutter present. Characteristic ventral lip with
Iy developed, lock area and main tubercle and link median depression, ventral side convexo No lip
medium sized; lunar and minor tubercles rather flaps; lip bellies pronounced, lip shoulders smal!.
small, all tubercles close together. Second lock Aperture wide, rim sharp, ventral rim slightly
mechanism present. Sculpture of faint growth thicker. No hump. Protoconch II are a projecting
lines and faint transverse striation. Lateral spines moderately. Shelllength: 6.8 mm width: 5.4 mID.
large, not bent, sharp and lanceolate. Gutter cor- Diacavolillia deshayesi van der Spoel, Bleeker
ners smal!. Caudal spine and caudal spine mark and Kobayasi, 1993 (Fig. 6.22). Shell hyaline,
678 Pteropoda
subtriangular in outline. Faint brown to pink hue moderately developed, lateral lines slightly con-
over entire shel\. Ventral side vaulted; 20-40 (usu- vex. Lock area small, lock mechanism relatively
ally 27) distinct comb-shaped ventral ribs. Dorsal large. Second lock mechanism at upper level of
side convex, with strong central rib and latero-dor- attachment of spines to lateral line. Sculpture of
sal ribs; lock ribs small, lock area, main tubercle faint growth lines and faint transverse striation.
and link large. Lunar and minor tubercles typical. Lateral spines well-developed, slightly bent dor-
Tiny second lock system presento Lateral lines sally, gutter comers well-developed. Caudal spine
slightly convex, flanks poorly developed. and caudal spine mark absent. Broad dorsal lip
Sculpture of faint growth lines and faint transverse with small notch and gutter, but no constriction.
striation. Growth lines well-developed on dorsal Broad rostrum with small nose at tip of lip.
side. Well-developed lateral spines slightly bent, Ventral lip medium-sized, with median depres-
sharply pointed and hooked. Gutter comers large. sion. No lip flaps; lip belly well-developed and
Caudal spine and caudal spine mark absent. low, lip shoulders smal\. No hump. Protoconch II
Ventral lip relatively large with clear median area projecting moderately. Shell length: 10.4
depression. Slender dorsallip with notch, constric- mm, width: 8.6 mm.
tion and gutter. Traces of lip flaps present; mode- • Diacavolinia limbata (d'Orbigny, 1836) forma lim-
rate lip bellies and well-developed lip shoulders. bata (d'Orbigny, 1836) (Fig. 6.25). Shell triangular
Hump absent. Protoconch II area projects mode- in outline, hyaline, brown-purple. Ventral side con-
rately. Shelllength: 8.2 mm, width: 6.8 mm. vex, without flattened area as in forma africana.
• Diacavolinia e/egans van der Spoel, Bleeker and Twenty-five ventral ribs, in anterior group of 15
Kobayasi, 1993 (Fig. 6.23). Shell slightly triangu- strong ribs, and a posterior group of weak ribs.
lar, colorless, hyaline. Dorsal side slightly convex, Dorsal side slightly convex, with well-developed
central dorsal rib well-developed, latero-dorsal central ribs, lateral ribs and lock ribs. Lateral lines
ribs moderately developed, lock ribs smal\. slightly convexo Flanks not pronounced. Lock area
Ventral side vaulted, 18-24 ventral ribs weakly well-developed with a large main tubercle and link.
developed. Flanks inconspicuous; lateral lines No separate lunar or minor tubercles. Second lock
slightly convex, lock area visible, lock mechanism mechanism at upper level of attachment of spines to
medium-sized. Main tubercle hooked, lunar tuber- lateral side. Sculpture of faint growth lines and faint
cle small, minor tubercle absent. Second lock transverse striation. Lateral spines slightly bent,
system rudimentary. Sculpture of faint growth sharp, hooked; gutter corners large. Caudal spine
lines and very faint transverse striation. Hooked and caudal spine mark absent. Broad dorsallip with
lateral spines bend slightly dorsally and projecting small notch and gutter but no constriction. Ventral
laterally. Caudal spine and caudal spine mark lip moderately sized with well-developed median
absent. Small dorsal lip without notch or constric- depression. No lip flaps; low and strong lip bellies
tion. Rostrum short and narrow, concave but wit- and small lip shoulders. Hump absent. Protoconch
out a clear gutter. Ventral lip medium-sized with II area projecting moderately, the relatively small
moderate median depression. Caudal fold 0.84 caudal joint is about 1.36 mm, caudal fold, about
mm long, moon shaped, left and right folds form a 1.68 mm long, moon-shaped. Shell length: 12.7
small angle in ventral view. No lip flaps; lip bel- mm, width: 10.4 mm.
lies weak; lip shoulders small. No hump. • Diacavolinia /ongirostris (de Blainville, 1821)
Protoconch II area projects moderately. Shell (Fig. 6.26). Shell hyaline, triangular, brown, espe-
length: 6.2 mm, width: 5.4 mm. cially near ventral lip border and centre of dorsal
• Diacavo/inia limbata (d'Orbigny, 1836) forma and ventral sides. Ventral side vaulted. Seventeen
africana van der Spoel, Bleeker and Kobayasi, thin, linear ventral ribs composed of small tuber-
1993 (Fig. 6.24). Shell roughly triangular, with cles. Dorsal side slightly convex, with well-deve-
brown-purple hue. Dorsal side convex, central rib loped central rib, lateral ribs and lock ribs. Lock
and latero-dorsal ribs well-developed, lock rib area small with blunt or sometimes sharp main
strong. Ventral side vaulted, upper half of ventral tubercle and smalllink. Lunar and minor tubercles
side flattened in outline. Twenty-seven ventral typica\. Second lock mechanism presento Lateral
ribs strongly developed, comb-shaped. Flanks lines slightly convex, flanks poorly developed.
Pteropoda 679
Growth lines strong on both ventral and dorsal and latero-dorsal ribs; lock ribs weak. Lock area
sides. Sharp lateral spines slightly bent, protruding well-developed with small main, lunar and minor
laterally and slightly hooked; gutter corners large. tubercles, and link. Second lock system absent.
Caudal spine and caudal spine mark absent. Dorsal Lateral lines convex, flanks well-developed.
lip with notch and gutter but no constriction. Lateral spines .not bent, with small, sharp hooks;
Ventrallip moderately sized with distinct median gutter corners smal!. Caudal spine and caudal spine
depression. Only traces of lip flaps; lip bellies and mark absent. Dorsal lip without notch or constric-
lip shoulders small. Aperture oval, large, lip angle tion but gutter presento Depressed dorsal lip with
about 135°, lock angle 77°, side angle 66°. faint nose. Rostrum slender, moderately long.
Aperture forms two long lateral slits directed pos- Moderate ventrallip with small median depression.
teriorly, aperture rim sickle-shaped in dorsal view. No lip flaps; strong lip bellies and strong lip shoul-
Caudal fold about 1.60 rnm long, nearly straight, ders. Hump very small or absent, inner hump
left and right folds form angle with each other. always absent. Protoconch Il area projecting
Hump absent. Protoconch Il area projecting mod- strongly. Shelllength: 5.4 rnm, width: 4.5 rnm.
erately, with curved caudal joint, of about 1.28 Diacavolillia strallgulata (Deshayes, 1823) (Fig.
mm. Juveniles dorso-ventrally compressed, 6.29). Shell hyaline, triangular, brown. Brown
spoon-shaped in ventral view with straight shell color especially noticeable along centre of dorsal
except for posteriormost section that bends slight- side. Ventral side vaulted. Eighteeen strong, linear
Iy dorsally. In older juveniles caudal spine is dis- ventral ribs composed ofrows oftubercles. Dorsal
carded and rounded shell composed of two val ves side slightly flattened, with well-developed central
is left, initially open caudally. Shell length: 7.2 and weaker latero-dorsal ribs; lock ribs smal!.
mm, width: 6.8 rnm. Lock area, main tubercle and link well-developed.
Diacavolillia ovalis van der Spoel, Bleeker and Lunar tubercle relatively small, minor tubercle
Kobayasi, 1993 (Fig. 6.27). Shell hyaline, color- strongly reduced. Extremely small second lock
less triangular. Ventral side vaulted. Dorsal side mechanism marked by little curve on lateral bor-
convex, central rib and latero-dorsal ribs well- der of dorsal side. Ribs on dorsal side. Lateral
developed, lock ribs well-developed but small, 19- lines nearly straight, flanks well pronounced.
23 ventral ribs, strongly developed, comb-shaped. Lateral spines only slightly bent, sharp and
Flanks present; lateral lines convexo Lock area hooked; gutter corners smal!. Caudal spine and
small with moderately developed main and lunar caudal spine mark absent. Dorsal lip with notch,
tubercle and small minor tubercle. Second lock gutter, pronounced constriction, and inner hump.
system presento Lateral spines well-developed. Abrupt lateral constriction in rostrum gives rise to
Long hooked lateral spines bend dorsally. Caudal tubercle distally, separated from rest of dorsal
spine and caudal spine mark absent. Dorsal lip side. Ventral lip moderate in size with normal
with notch, constriction, gutter. Ventrallip normal median depression. No lip flaps; moderate lip bel-
sized with median depression. Small nose present lies and well-developed lip shoulders. Moderate
at base of rostrum. Lip flaps weakly developed; lip hump present but no inner hump; rostrum projects
bellies strong and low, lip shoulders well-devel- anteriorly. Protoconch Il area projecting moder-
oped. Aperture narrow. Dorsal aperture rim sharp, ately, provided with a relatively broad caudal
ventral rim slightly thickened. Inconspicuous joint. Shelllength: 4.6 rnm, width: 4.2 mm.
hump present but inner hump absent. Protoconch
Il area projecting moderately. Shell length: 6.1 Genus Diacria
mm, width: 5.1 mm. These species discard the protoconch and form a clos-
Diacavolillia robusta van der Spoel, Bleeker and ing septum.
Kobayasi, 1993 (Fig. 6.28). Shell hyaline, subtrian- • Diacria dallae van der Spoel, 1968 (Fig. 6.30).
guiar, brown. Brown coloration especially notice- Shell red-brown, globular with rounded ventral and
able along aperture rim, joint, and ventral side, dorsal sides and well-developed ribs. Lateral sides
except for its margins. Ventral side vaulted. curved and thickened between lateral spines and
Nineteen conspicuous, thin, linear ventral ribs. corners of caudal spine mark. Shell sculpture con-
Dorsal side somewhat flattened, with weak central sisting of faint growth lines and 3 dorsal, mode-
680 Pteropoda
rately developed striae. Lateral spines small, cau- conch 1 globular, no sculpture on protoconch 1 or
dal spine absent as protoconch is discarded leaving 11, lateral sides of protoconch 11 thickened.
oval spine mark. Dorsal lip arch-shaped bordering Teleoconch length: 8 mm, width: 10 mm, ratio
very narrow shell aperture. Aperture lips thick- upper lip-spine tip to spine tip-membrane: 1.3,
ened. Shelllength: 2.1 mm, width: 1.7, diameter of spine mark width: 1.5 mm, aperture height: 0.9
caudal spine mark: 0.56-0.80 mm. Subtropical and mm. The hydroids Laomedea striata (Clarke,
equatorial forms of this species (Van Leyen and 1907), Kinetocodium danae Kramp, 1922 and
van der Spoel, 1982) differ only in size. Campaniclava clionis are often found as epibionts.
• Diaeria major (Boas, 1886) (Fig. 6.31). Shell with
long, often broken off, caudal spine. Surface Subfamily Clioinae
almost entirely white, hyaline, with exception of Pyramidal shells.
brown-red dorsal aperture rim and ventral side,
along aperture. Ventral side flat. Differs from D. Genus Clio
trispinosa f. trispinosa chiefly in lateral spines Aperture flattened or triangular, faint transverse stria-
directed more caudally. Aperture lips well-deve- tion or no striation, mostly a slight dorso-ventral dif-
loped. Juveniles with long dorso-lateral flattened ferentiation.
needle-like shell with lateral sides diverging ante- C/io andreae (Boas, 1886) (Fig. 6.34). Shell
riorly only. Protoconch 1 globular, no sculpture on straight except for posterior section which curves
protoconch 1 or 11; lateral sides of protoconch 11 abruptly dorsally, transparent, triangular in out-
thickened, discarded in adults. Teleoconch length: line, oval in cross-section, dark body and visceral
13 mm, width: 11 mm, spine mark width: 1.2 mm, mass visible through wall. Lateral ribs well-deve-
aperture height: 1.0 mm. loped, straight, regularly diverging, not excavated;
• Diaeria rampali Dupont, 1979 (Fig. 6.32). longitudinal ribs on dorsal and ventral sides and
Slender shell with long caudal spine, sometimes transverse striation faint or absent, both sides flat-
broken off. Rim of aperture and middle section of tened. Aperture width about one half to one third
dorsal ribs brown, color pattero ofventrallip con- total shell length. Embryonic shell spherical with-
nected with dark spot on ventral side anterior to out terminal cusp, separated from rest of shell by
lateral spines. Lateral spines white, lateral ribs clear constriction. Shelllength: 20 mm, 5 times as
occasionally light brown. Teleoconch, posterior to long as broad.
lateral spines, slender, caudal spine long (65% of Clio e/zaptali (Eydoux and Souleyet, Ms.) Gray,
the teleoconch length). Juvenile has long, dorso- 1850 (Fig. 6.35). Shell very broad, regularly
ventrally flattened, needle-like shell, with lateral curved dorsally, roughly triangular in outline, oval
sides diverging anteriorly only. Protoconch 1 glo- in cross-section, transparent, dark purple-brown
bular, no sculpture on protoconch 1 or 11. Lateral body visible through wall. Sharp lateral ridges
sides of protoconch 11thickened. Small ring may concave. Width near shell aperture about 3/4 total
be present between protoconch 1 and 11. shell length. Lateral sides rounded in cross sec-
Teleoconch length: 9.5 mm, width: 9 mm, spine tion. Three dorsal ribs present, medio-dorsal rib
mark width: 0.95 mm, aperture height: 0.95. broadest, ventral side without ribs or swellings.
• Diaeria trispinosa (de Blainville, 1821) forma tri- Wavy transverse striation on ventral and dorsal
spinosa (de Blainville, 1821) (Fig. 6.33). Shell sides, growth lines faint. Blunt ending, obtuse
strongly compressed dorso-ventrally, with long embryonic shell with transverse and longitudinal
caudal spine, sometimes broken off. Rim of slit- striae, not clearly delimited from teleoconch.
like aperture chestnut brown, rest of shell general- Constriction between protoconch 1 and 11 absent.
ly transparent and chestnut brown on thicker areas Shell length: 19 mm, width: up to 16 mm.
such as lateral ribs and tops of longitudinal ribs. Clio ellspidata (Bosc, 1802) (Fig. 6.36). Shell di s-
Three ribs ventrally, five dorsally. Lateral spines tinctly curved dorsally, pyramid-shaped, transpa-
are long extensions of lateral ribs, perpendicular to rent, triangular in outline and in cross-section.
shell axis. Embryonic shell spherical. Juvenile Elongated dorsal and lateral gutter shaped ribs
with long dorso-ventrally flattened needle-like enhance this shape. Greatest width of shell
shell, lateral sides diverging anteriorly only, proto- between tips of spines, at or behind middle.
Pteropoda 681
Ventral surface regularly convex, ventral rib C/io pyramidata Linnaeus, 1767 forma excisa van
appears as small swelling in middle disappearing der Spoel, 1963 (Fig. 6.39). Shell pyramid-shaped,
near the shell aperture. Two latero-dorsal sides, transparent, roughly triangular in cross-section.
delimited by dorsal rib, each with two inconspicu- Shell generally broader and stronger than in other
ous ribs. Growth lines prominent, lateral sides forms. LateraL sides round in cross section. Lateral
rounded in cross section. At aperture, dorsal side ribs concave at base, convex in middle, and straight
of shell protruding above ventral side. Embryonic and parallel to body axis near aperture. Four longi-
shell in adults spherical with blunt cusp, sharp tudinal ridges on each latero-dorsal side more
cusp at tip of embryonic shell in juveniles. developed than in most other forms. Growth lines
Constriction separates protoconch 1 from 1I, the prominent. Shelllength: 20 rnm, width: 11 rnm.
latter cylindrical over whole length. Shell length: Clio pyramidata Linnaeus, 1767 forma /allceo/ata
20 rnm, width: 30 rnm. Hydroids of Campaniclava (Lesueur, I813)(Fig. 6.40). Shell transparent, py-
cleodora (Gegenbaur, 1854) sometimes on shell ramid-shaped. Greatest width near middle of shell.
of this species. Circular in cross section near base, triangular at
C/io piatkowskii van der Spoel, Schalk and Bleeker, middle and in anterior half. Ventral surface con-
1992 (Fig. 6.37). Shell pyramidal, extremely broad, cave anteriorly, concavity of ventral side resulting
opaque. Lateral sides strongly diverging, top angle from lateral ribs which bend ventrally in upper
about 70. Proximal part of lateral sides concave, parto Three longitudinal ribs at both latero-dorsal
distal part convexo Ventral side slightly convex, sides, one on ventral side. Dorsal rib straight, pro-
with one central longitudinal ribo Convex dorsal truding distinctly above shell aperture; lateral ribs
side with 3 ribs. Transverse section through shell very long, thickened, strongly diverging but never
narrow oval and slightly asyrnmetrical as dorsal ending in free processes. Growth lines prorninent.
side is vaulted higher than ventral side. Lateral sides Transverse striation and growth lines distinct. The
sharp, not gutter shaped, with intricate pattem of oval embryonic shell is smaller than in other forms.
growth lines. Besides growth lines, there is wavy Shelllength: 20 rnm, width: 15 rnm.
transverse striation on ventral and dorsal sides. C/io pyramidata Linnaeus, 1767 forma martel/si
Dark purple footlobe and lips, large dorsal tentacles, (Pfeffer, 1880) (Fig. 6.41). Shell transparent, elon-
wings black, large, thick and fleshy, posterior foot- gate-triangular in outline, slightly curved ventral-
lobe small. Dentition on the median plate strong. Iy. Middle part of shell convex in outline; anterior
Protoconch 1 blunt and continuous with protoconch section with sharper lateral ribs, with parallel or
II and teleoconch without visible boundary. Shell somewhat diverging lateral sides. Poorly devel-
length: 13.5 rnm, width: 16 rnm. oped lateral ribs do not project laterally. Dorsal
C/io pyramidata Linnaeus, 1767 forma alltarctica surface usually with five ribs, transverse striation
Dall, 1908 (Fig. 6.38). Shell transparent, straight, on both surfaces distinct. Shell length: 17 rnm,
conico-cylindrical in shape. Cross-section near width: 9 rnm.
embryonic shell circular; fully grown specimens C/io pyramidata Linnaeus, 1767 forma su/cata
have depressed ventral side near aperture, so (Pfeffer, 1879) (Fig. 6.42). Shell transparent, tri-
cross-section here is more half-moon shaped. angular in outline, lateral sides not gutter shaped.
Lateral sides only developed in uppermost part of Cross-section near apex circular, triangular near
shell. With faint dorsal and broad ventral ribs aperture. Slight dorsal curvature present. A pro-
c10se to aperture, lateral ribs bordering the ventral nounced curving of shell top to left is common.
side, developed only near shell aperture. Growth Ventral side with one rib, dorso-lateral sides with
lines and prominent and strong transverse wavy five large longitudinal ribs, sometimes seven.
sculpture. Transverse striae cause shell to resem- Transverse striation regular, conspicuous growth
ble that of Hyalocylis striata (Fig. 6.50). Dorsal lines present. Embryonic shell droplet-shaped,
aperture border slightly higher than ventral. pointed, separated from adult shell only by faint
Reddish to dark-brown visceral mass visible but evident constriction. Shell length: 20 rnm,
through shell wall. Embryonic shell pointed, sep- width: 10 mm.
arated from rest of shell by constriction. Shell C/io recurva (Childem, 1823) (Fig. 6.43). Shell
length: 17 mm. transparent, straight except for posterior part, tri-
682 Pteropoda
angular in outline, cross section oval. Three con- Protoconch I is conical. Shell length: 1.6 (excep-
spicuous dorsal ribs present, middle one broadest. tionally up to 2.7) mm, width: 0.2 mm, embryonic
Ventral rib so broad that it covers nearly whole shell 0.4-0.5 mm long.
ventral side. Lateral ribs double, with f1at lateral Creseis virgula (Rang, 1828) forma conica
surface between each pair of ridges, which may be Eschscholtz, 1829 (Fig. 6.47). Shell transparent,
concave distally, gutter-shaped. Sculpture of con- slender, tube-shaped, curved, cross-section circu-
centric undulations produces wavy transverse stri- lar, surface smooth. With wing protrusions. Most
ation. Embryonic shell subspherical with clear typical is the rounded embryonic shell which does
cusp which may be wom off in older specimens. not show rings or constrictions, and the strong cur-
Radula formula 1-1-1, sometimes (1)-1-1-1-( 1). vature (angle between teleoconch and protoconch:
Shelllength: 30 mm, width: 17 mm wide, aperture 30°). Anterior part of shell straight, posterior sec-
1/2 or more of total shell length. Sometimes tion curved dorsally. Shelllength: 7 mm, aperture-
hydroids of Campaniclava clionis VanhOffen, diameter: 1 mm.
1910 attach to the shell. Creseis virgula (Rang, 1828) forma constricta
Chen and Bé, 1964 (Fig. 6.48). Shell transparent,
Genus Creseis short, narrow, tube-shaped, slightly curved, cross-
Aperture rounded, no longitudinal groove over dorsal section circular, surface smooth. Well defined
side. constriction separates protoconch from adult por-
Creseis acicula (Rang, 1828) forma adeula tion; protoconch with a distinct bulge, gradually
(Rang, 1828) (Fig. 6.44). Shell transparent, cross- increasing in size toward anterior part of adult
section circular, surface smooth, extremely long shell. Shell length: 0.54 mm, diameter: 0.12 mm.
and narrow, tube-shaped, not curved, aperture Creseis virgula (Rang, 1828) forma virgula
rounded. Columellar muscle runs free through (Rang, 1828) (Fig. 6.49). Shell transparent, cross-
shell lumen over great distance (between attach- section circular, surface smooth, long, narrow,
ment point and point where muscle disappears into tube-shaped, curved in posterior parto With wing
visceral mass). With wing protrusions. Embryonic protrusions. Embryonic shell without rings above
shell differs in shape from that of forma clava by rounded posterior part, delimited only by faint
caudal end consisting of oval-shaped embryonic constriction. Angle between teleoconch and proto-
part with one or two ring-shaped swellings crani- conch: >30°. Length: 6 mm, diameter: 2 mm.
ally. Rear angle of shell 13-14°, shelllength: 33
mm, width: 1.5 mm. Genus Hyalocylis Fol, 1875
Creseis acieula (Rang, 1828) forma clava (Rang, • Hyalocylis striata (Rang, 1828) (Fig. 6.50). Shell
1828) (Fig. 6.45). Shell transparent, cross-section completely transparent, fragile, cross-section
circular, surface smooth, short and narrow, tube- round, surface with transverse undulations, long,
shaped, not curved. Body connected by columellar tube-shaped, slightly curved dorsally at end.
muscle to caudal tip of shell; this muscle disap- Embryonic shell discarded in adults, punctuated
pears quickly above attachment to shell into vis- closing membrane formed near place of rupture.
ceral mas s, only small part of it running free in Rear angle of adult shell: 24°, shelllength: 8 mm.
shell lumen. With wing protrusions. Embryonic
shell tip rounded; outline of caudal section shows Genus Styliola
two or more swellings caused by transverse rings • Styliola subtda (Quoy and Gaimard, 1827) (Fig.
below concave portion ending in perfectly spheri- 6.51). Shell transparent, long, tubular, straight,
cal embryonic end. Shell length: 6 mm, diameter needle-like. Cross-section round. Surface with
at the aperture: 1 mm. faint growth lines. In upper 2/3 of shell a grooved
Creseis chierehiae (Boas, 1886) (Fig. 6.46). Shell rib projecting from aperture rim like cusp, rib
transparent, cross-section circular, surface with twisting slightly, so its position is medial at cranial
conspicuous transverse rings. With wing protru- end but caudal end is displaced left. Posterior shell
sions. Embryonic shell with anteriorly convex, end pointed, with two consecutive constrictions.
posteriorly concave sides, blunt tipo Clear con- Rear angle of shell: 11°, shelllength: 13 mm.
striction separates embryonic from adult shells.
Pteropoda 683
Subfamily Cuvierininae • CYlllblll;a peroll; de Blainville, 1818 morpha pe-

rOIl; de Blainville, 1818 (Fig. 6.55). Pseudoconch
Genus ClIV;er;lla covered with spines arranged in rows, some spines
ClIV;er;lla collllllllella (Rang, 1827) forma atlallt;- irregularly dispersed between rows. Spines near
ca van der Spoel, 1970 (Fig. 6.52). Shell complete- border ofshoe.-shaped cavity ofpseudoconch larg-
ly transparent, bottle-shaped, greatest width at 1/3 er than rest and asymmetrica!. Colorless heart-
shell length from base, constricted below reniform shaped swirnming disc, in middle of posterior
aperture. Surface smooth or with very faint growth margin a wing lobe with long filamentous tentacle
lines. In juvenile stage shell with needle-shaped projecting outwards. At outer wing border two
protoconch (Fig. 6.52c, d, e), closed off by convex transparent spots, without muscles; muscle fibres
septum; protoconch 1 ova!. Shelllength: 10 rnm. fused. Radula teeth degenerate, jaw relatively
smal!. Veliger with velum with 4 lobes, larva 1
Suborder Pseudothecosomata shell with 1.5 whorls (maximum diameter: 0.72
With or without shell, radula formula (l )-1-1-1-( 1). rnm), last whorl increasing rapidly in width.
Length of pseudoconch: 65 rnm.
Family Cymbuliidae Cymblllia s;bogae Tesch, 1903 (Fig. 6.56).
Primary shelllost completely, visceral mass surround- Pseudoconch without medial constriction.
ed by secondary gelatinous slipper-like shell, the Anterior end slightly hollow and provided with
pseudoconch. small spines. Spines near opening of cavity absent
or extremely smal!. Two rows of spines ending at
Subfamily Cymbuliinae ventral points not constricted in middle, almost
straight throughout their length. Wings disc
Genus Cymblllia shaped. Pseudoconch length: 24 rnm.
With long, pointed, slipper-like pseudoconch. Well-
developed buccal mass. Visceral mass forms dark Subfamily Glebinae
nucleus embedded in perfectly transparent pseudo- Reduced buccal mass; organisms collect food with
conch covered with cusps. free floating mucous web secreted by mucus glands
Cymblllia parvidelltata Pelseneer, 1888 (Fig. on wmgs.
6.53). Pseudoconch dorsally sharp pointed, late-
rally compressed in middle, no large spines at bor- Genus Corolla
der of excavation. Spines less developed than in Pseudoconch with many distinct tubercles on aboral
Cymbulia peroni, with five rows of tubercles on side.
oral side; on aboral side dorsally 3 and ventrally 4 Corolla calceola (Verrill, 1880) (Fig. 6.57).
rows of tubercles. Ventral end of pseudoconch Pseudoconch oval, transparent, with shallow,
hollow without spines. Rows of spines at under- broad cavity and blunt anterior margino Small
side are not straight and parallel but sinuous. tubercles regularly distributed, relatively few in
Wings disc-shaped. Pseudoconch length: 35 mm. numbel. Aperture more than half of length of
Cymblllia peroll; de Blainville, 1818 morpha pseudoconch. Swirnming dise oval, not reaching
m;llor van der Spoel, 1976 (Fig. 6.54). far anteriorly over pseudoconch. Proboseis short
Pseudoconch smaller than in morpha peroni, with and broad, with two syrnmetrical tentacles, free
less regularly curved underside. Sides have only 1 from disc over most of length. Mantle gland asym-
or 2 rows of spines, rather than 3. Spines small, metrieal, with two oblique, completely transparent
even near aperture, and without asymmetrical bands. Right comer of gland is anterior to left eor-
development of spines along aperture. Two knobs nel. Pseudoeoneh length: 40 rnm, swirnming dise
at dorsal side of pseudoconch cavity small (with diameter: 70 rnm.
broad bases in morpha milZor; with large, narrow Corolla cllpllla Rampal, 1996. (Fig. 6.58).
bases, usually with two or more spines, in morpha Pseudoconch undescribed. Swirnming dise folded
perolZi). Wings and visceral mass as in morpha on itself in posterior part, semicireulal. Proboseis
perolZi. Radula teeth degenerate, jaw relatively free only along half of its length; rest covered by
smal!. Pseudoconch length: 42 mm. parapodial coi!. Mucus glands along border of
684 Pteropoda
swimming disk, in latero-posterior position; 20 in Genus Gleba

number. Oval, blunt and very flattened pseudoconch. Few
Corolla intermedia (Tesch, 1903) (Fig. 6.59). tubercules on aboral side. Proboscis very long.
Pseudoconch transparent, yellowish, oval, both Gleba eordata (Forskal, Ms.) Niebuhr, 1776 (Fig.
poles nearly equal in shape. Aperture of cavity not 6.62). Pseudoconch covered with few warts, rather
reaching middle of pseudoconch. Small tubercles flat and broad, posterior end rounded, anterior end
on pseudoconch irregularly distributed, absent on nearly straight and ending abruptly. Cavity large
ventral side. At aboral side tubercles are smaller but shallow, clearly visible only in thicker posterior
than elsewhere. Margin of swimming disc situated parto Swimming disc characterized by straight pos-
close to ventral pseudoconch border. Proboscis terior border and semi-circular shape. Numerous
prominent. Mantle gland with two complete and well- developed mucus glands along wing border.
one incomplete transparent transverse bands; Muscle bands as in Cymbulia, much closer together
asymmetrical; left part is more developed than than in Corolla and difficult to see separately.
right. Pseudoconch length: 39 mm, swimming disc Proboscis very long, free over most of length,
diameter: 30 mm. reaching over 2/3 of swimming disco At base of pro-
Corolla ovata (Quoy and Gaimard, 1832) (Fig. boscis two small, symmetrical tentacles. Visceral
6.60). Pseudoconch transparent, with broad cavity mass relatively small. Mantle gland with two com-
occupying 1/2 or more of dorsal half. Tubercu- plete and one incomplete oblique transparent bands.
lation rather coarse, irregular, densest on dorsal Gland twisted to left. Diameter of swirnming disc:
side. Swimming disc broadly oval, not reaching far 60 mm, length of pseudoconch: 45 mm.
beyond border of pseudoconch in preserved speci-
mens (may reach far over this border when alive), Family Desmopteridae
with two symmetrical tentacles and two homs of
moderate length. Wing musculature consisting of a Genus Desmopterlls
few opaque broad muscle bands forming 3 separate Shell and pseudoconch absent.
crossing systems visible in well-preserved speci- • Desmopterlls papilio Chun, 1889 (Fig. 6.63). Wings
mens. Proboscis free from disc surface over nearly disc-shaped composed of five lobes, transparent
total length. Mantle gland slightly asymmetrical, except for a few muscle fibres. Two incisions in
with 4 broad bands and 3 narrow transparent swimming disc border define square-shaped median
bands, of which one is incomplete. No glands parto Two wing tentacles arising between lateral
along wing border. Pseudoconch length: 40 mm, lobes ofwings. Visceral mass with opaque appendix
diameter of swimming disc: 80 mm. curved around swimming disco Besides median
• Corolla speetabilis Dall, 1871 (Fig. 6.61). lobe, disc has two long tentacles. No proboscis.
Pseudoconch rounded at both ends, covered with Anterior part of rounded cylindrical body large, pos-
large warts which are smaller and more closely terior part small, only extreme end showing some
spaced towards postero-dorsal side. Aperture coiling. Head parts curved ventrally in right angle to
wider than in C. ovata, nearly identical to that of plane of swimming disco Integument with small red-
C. ealceola. Swimming disc translucent and yel- dish gland cells. Glandular, ciliated fold at posterior
lowish, broadly rounded to quadrangular, some- body pole. Buccal appendages lacking, tentacles
times with two depressions in posterior margin; small with rudimentary eyes. Body length: 2.0 mm,
protruding far behind pseudoconch border. Small diameter of swimming disc: 4 mm.
mucus glands along border of swimming disco
Proboscis free from upper surface of swimming Family Peraclididae
disco Lateral foot appendages, forming comers of
proboscis, rounded. Mantle gland shows one com- Genus Peraclis
plete and two incomplete transverse transparent Left coiled shell, sculpture usually irregular reticulate.
bands, gland slightly asymmetrical. Pseudoconch Peraclis apieijlllva Meisenheimer, 1906 (Fig.
length: 40 mm, width: 25 mm, diameter of swim- 6.64). Shell elongate, with 4 whorls; white near
ming disc: 80 mm. aperture, dark red near apex. Spire rather high
from rapid increase in whorl size. Sculpture a sub-
Pteropoda 685
regular reticulum, with fine rows of tubercles • Peraclis lIIoluccellsis Tesch, 1903 (Fig. 6.67).
arranged in spiral lines forming reticulate pattern Hyaline shell with 5 whorls. Body whorl increas-
on upper half of body whorl. Keel without aper- es very rapidly, spire strongly depressed. First
ture tooth. Radiating crests between keel and deep whorls coiled in one plane, last whorl very large,
suture, size decreasing from aperture towards may be higher than apex, obscures other whorls
apex. Large aperture with rounded outer border, except in apical view. Last whorl, at dorsal margin
upper aperture border tilted slightly toward apex, with keel ending in aperture tooth directed slight-
lower margin points to rostrum with very well- Iy toward apex and curving spirally with shell.
developed columellar membrane. Operculum with Between keel and deep suture a triangular field
6 whorls without radial striation. Shell heighí: 4 with about 7 strong, irregular crests, undulating
mm, diameter: 3.3 mm. and placed closer together than in P. bispinosa and
Peraclis bispillosa Pelseneer, 1888 (Fig. 6.65). P. apicijulva. Shell sculpture absent. With growth
Shell elongate, with 4 to 5 whorls. Spire relatively lines, with perpendicular striae sometimes visible
high from rapid increase in of whorl size. under high magnification. Aperture wide, inflated
Sculpture composed of irregular reticulum; disap- in upper part and pointed towards rostrum.
pearing abruptly before aperture border. On last Rostrum not curved, but twisted, with narrow co-
half of body whorl strong keel ends in distinct lumellar membrane. Umbilicus absent. Operculum
aperture tooth. Radial crests between deep suture glassy, sinistral, with 4-5 whorls and radial striae.
and keel resemble those in P. apicijulva, distinct Embryonic shell with rough ornamentation of
but more regular and thinner than in latter. irregularly ramified flat ridges (Boltovskoy,
Aperture wide near tooth, at end of keel; triangu- l 974b). Shell heighí: 3 mm, diameter: 2.5 mm.
lar portion of aperture border turned upwards Peraclis reticulata (d'Orbigny, 1836) (Fig. 6.68).
forming second tooth. Base of aperture pointed Shell reddish or yellow, relatively high, horny,
toward keeled, spirally twisted rostrum with very with 4-5 whorls and deep suture without radial
weIl-developed, broad columeIlar membrane. crests. Spire not depressed. Shell covered with
Operculum with about 5 whorls, no radial striae. reticulate sculpture of hexagonal (sometimes
Soft body dark brown to black. Embryonic shell square or pentagonal) cells, sculpture disappearing
almost smooth; faint, irregular ornamentation vi- at short distance from aperture border. Reticulated
sible only under high magnification (Boltovskoy, layer is not a periostracum as it is composed of cal-
1974b). SheIl heighí: 7.5 mm, diameter: 6 mm. careous material. Young specimens of about 2 mm
• Peraclis depressa Meisenheimer, 1906 (non have 3.5 whorls totally covered by reticulum. Keel
Vayssiere, 1913) (Fig. 6.66). Slightly elongate present. No aperture tooth, aperture wide oval,
sheIl with 3 whorls and deep suture without radial base of aperture pointed toward keeled rostrum.
crests. Color of sheIl white, only large specimens Depending on age, faint radiating crests may
with yellowish apex, keel brownish to yellow. develop along suture. Rostrum present with narrow
Spire depressed, whorls increasing quickly in size. columellar membrane. Umbilicus very narrow or
Short keel ends in distinct aperture tooth, aperture absent, operculum with 5.5 whorls without, radial
wide, base of aperture pointed towards keeled ros- striation. Shell heighí: 6 mm, diameter: 3 mm.
trum. Shell surface with regular structure of spiral Peraclis triacantha (Fischer, 1882) (Fig. 6.69).
lines, sometimes forming reticulate pattern. About 4 whorls, body whorl covering all others.
Tooth-like protrusion visible in inner angle of Deep suture without radial crests. Overall color of
upper aperture border, turned upwards and Iying shell white. Spire strongly depressed. Faint sculp-
close to apex. Aperture tooth with keel runing over ture of spiral lines with cross connections resem-
only short distance, along upper whorl. Outer bling reticulate pattern. Two aperture teeth, one
aperture border regularly rounded. Columella pro- supported by strong rib, aperture wide and round-
vided with very strong keel which continues into ed. Ribs and rostrum brown. Spirally twisted ros-
rostrum; columellar membrane relatively well- trum, with keel and very large columellar mem-
developed, but not broad. Umbilicus clearly vi- brane. Umbilicus well-developed. Rostrum and
sible and wide. Operculum, with 5 whorls, no radi- second aperture tooth both visible in apical view.
al striae. Shell heighí: 3 mm, diameter: 2.5 mm. Operculum with subcentral nucleus, with 5 whorls
686 Pteropoda
and suture bordered by many transverse lines. Genus Paracliolle

Shell height: 5 mm, diameter: 5 mm. Two pairs of buccal cones, hooks presento
• Peraclis valdiviae (Meisenheimer, 1905) (Fig. Paracliolle IOllgicaudata (Souleyet, 1852) (Fig.
6.70). Four whorls separated by deep suture with 6.72). Body slender, oval; visceral mass visible
radial crests. Spire strongly depressed, body whorl through body wall. Body and wings dark. Posterior
constituting most of shell. Spiral ribs on body footlobe absent in some specimens, small when
whorl, stronger near aperture. Sculpture of hexago- presento Median tubercle in animal s with and with-
nal or quadrangular reticulation on first whorls, out posterior footlobes. Buccal cones of same type
similar to P. bispinasa. Aperture wide and oval.· as in Cliane, but larger. Median radula plate with
Strong keel runs along suture. Columellar mem- one long spine on free border, sometimes two
brane well-developed. Operculum with 5 whorls reduced lateral ones. Jaw reduced. Hook sacs
characterized by double lines along spire, with small, transparent, shallow, each with fewer than
transverse striae between double lines. Median 15 short curved hooks. Row of rugosities between
radula tooth monocuspoid with numerous lateral hook sacs represents jaw. Lateral gill absent, its
denticules. Shell height: 6.2 mm, diameter: 5.0 mm. site marked by transparent lacunar spot. Radula
formula 2-1-2 to 6-1-6. Body length: 10 mm.
Order Gymnosomata
Naked molluscs. Their identification based chiefly on Subfamily Thliptodontinae
anatomical details of buccal mass. Buccal cones absent.
Suborder Gymnosomata Genus Cephalobrachia

Molluscs without shell or mantle; body usually cylin- Hook sacs very large and muscular; intermediate
drical; in some families with gill-like appendages on radular plate always absent.
lateral ami/or posterior sides of the body; all species Cephalobrachia macrochaeta Bonnevie, 1913
with radula and hooks; some families with acetabuli- (Fig. 6.73). Body oval to barrel shaped, largely
ferous arms, others with buccal cones. transparento Mostly blackish visceral mass fills
entire body. Small lateral footlobes coalesce in
Family Clionidae front, fonning single lobe, fixed anteriorly. Posterior
Characterized by complete absence of gills. Body footlobe, small when present, pointed; all footlobes
usually smooth, in most genera the only appendages united. Median tubercle absent. Wings large, narrow
are those of the buccal mass; always with hook sacs; at base, tapering toward free tips. Hook sacs very
wings located beneath mouth at both sides of body, large, with large gold-colored hooks. When hook
with lateral and posterior footlobes; in some species sacs invaginated, both body poles are rounded;
footlobes strongly reduced. when evaginated, forming two powerful arms,
diverging from buccal cavity. Hooks, about 40 on
Subfamily Clioninae each side, of different sizes, largest as long as 1/4
Buccal cones presento body length. Jaw replaced by lip of glandular cells,
ventral to radula; lip forming bow across ventral
Genus Cliolle side of proboscis at base of odontophore between
Three pairs of buccal cones. hook sacs. Radula on large conical odontophore,
Cliolle limacilla (Phipps, 1774) alltarctica formula 3-1-3, with about 12 transverse rows; teeth
(Smith, 1902) (Fig. 6.71). Body largely transpa- extremely large. Lateral radula teeth hom shaped, in
rent, with orange to red pigmentation in tail, tenta- some laterals basal plate obscure or absent. Median
cles, and mouth organs. Reddish-brown visceral plate with blunt cusp and number of small denticleE
mass visible through body wall. Body robust, with at free border. Young specimens with circular cilia·
glandular skin. Three pairs of buccal cones of ted bands. Body length: 10 mm.
equal size but dorsal pair with bases free from
bases of central pair of cones. Median radula plate Genus Massya
only in very first rows, lateral teeth simple. Radula No intermediate plate, no gullet bladders. Only on(
formula 3-0-3 to 10-1-10. Body length: 40 mm. specles.
Pteropoda 687
Massya longicirrata (Massy, 1917) (Fig. 6.74). sometimes 2 to 4 small cusps near topo Body
Body largely transparent, rounded, unpigmented, length: 8 rnm.
often with ciliary bands. No buccal arms. Anterior
tentacles very long, trailing behind body, exceeding Family Cliopsidae
its length; tentacles homologous with head palps. With posterior gill, no buccal cones, posterior foot-
Base of tentacles muscular and broad, tapering lobe reduced or lacking.
abruptly above base. Wings very broad and round-
ed distally, left one always smaller than right one. Genus Cliopsis
At posterior margin of wings, where they join, No lateral gill; posterior gill hexagonal in adults.
small projection (posterior footlobe) is apparent. Cliopsis kro/l1li Troschel, 1854 morpha grandis
Hooks, 20 on each side, minute, triangular. sub- Boas, 1886 (Fig. 6.77). Visceral organs visible
equal in size. Jaw composed ofrow ofsmall spines. through the body wall. Integument not as transpa-
Radula formula: 3-1-3, very large, with large arcu- rent as in other morphs, but with same distribution
ate, dentated median plates with 40 to 50 small of chromatophores. Body broad with tlat posterior
cusps at free border, lateral teeth with broad bases. pole, balloon-like. Broad circular posterior gill with
Around 11 to 16 transverse rows of median plates 4 broad but short, fringed crests. Hook sacs shallow,
and 8 to 9 rows of laterals. Body length: 3.5 rnm. each with about 60 hooks. Head parts relatively
smal!. Lateral lobes of foot long, projecting over
Genus Thliptodon body. Median foot tubercle split into longitudinal
With intermediate plate and gullet bladders. halves. Proboscis long, evaginable to twice length of
Thliptodon antarcticlIs Meisenheimer, 1906 (Fig. body. Radula formula: 5-1-5. Body length: 40 rnm.
6.75). Body largely transparent, rounded to oval, Cliopsis krohni Troschel, 1854 morpha krohni
visceral mass visible through body wal!. Wings Troschel, 1854 (Fig. 6.78). Body oval, completely
small, implanted laterally in middle of body. Head transparent, faintly colored or colorless. Visceral
and body parts of nearly same size. Three foot- organs yellowish-brown, filling almost whole
lobes usually fused. Median footlobe short and body; visible through body wal!. Posterior gill
pointed. No median foot tubercle. Hook-sacs with 4 rays, without fringes. Posterior tentacles
about 3 rnm deep, each with 30 to 50 hooks. Penis retractable, anterior very smal!. Proboscis, when
with accessory sucker. Radula formula 3-1-1-1-3, evaginated, twice as long as body or longer.
with about 9 to 13 rows. Median plate very slen- Digestive tube in proboscis may reach length of
der, band-like, with fine dentition, or an undulated 110 rnm. Length of esophagus not permitting buc-
anterior border. Intermediate plate differing dis- cal organs to become evaginated like proboscis.
tinctly from that in other species by slightly curved Median footlobe lacking, well-developed median
spine at tip; this spine with dentition resembling tubercle with longitudinal grooves. Hook sacs
that of median plates of Cephalobrachia. Three shallow, with up to 60 hooks. Radula formula: 6-
lateral plates slender with straight margins, com- 1-6 to 4-1-4. Median plates tricuspoid. About 20
posed by articulated basal and top parts. Body transverse rows of teeth. First lateral teeth differ
length: 4 rnm. from other laterals by denticle on medial side in
Thliptodon diaphanlls (Meisenheimer, 1903) middle of head cusp. Body length: 24 rnm.
(Fig. 6.76). Body largely transparent, visceral • Cliopsis kro/lIli Troschel, 1854 morpha modesta
mass visible through body wal!. Shape variable, (Pelseneer, 1887) (Fig. 6.79). Body oval, com-
from droplet-shaped with extremely large head to pletely transparent, visceral organs visible through
barrel-shaped with head section about as large as body wal!. Dark chromatophores near posterior
body section. Wings implanted laterally in middle gill and in middle of body. Visceral mass not
of body. Posterior and lateral footlobes separate. reaching the posterior poleo Posterior gill ring-
Median foot tubercle present. Radula formula: 3- shaped or hexagonal, with 4 thin, short rays, with-
1-1-1-3, with 8 rows. Median plate very fragile, out fringes. Lateral footlobe broad, no posterior
thin, transparent; borders drawn out on each side. lobe; median foot tubercle present. Proboscis long.
Dentition usually absent, sometimes very faint. Buccal organs similar to those in morpha kro/mi.
Intermediate plates with a short top cusp and Body length: 3 mm.
688 Pteropoda
Family Notobranchaeidae Radula formula 6-1-6 or 5-1-5. Thirty transverse

With posterior gill only, sometimes with buccal rows of teeth. Median plate semilunar, with 1 or 3
con es, posterior footlobe distinct. small central cusps and dentition on free border.
Central cusps very small, represented by central
Genus Notobranchaea group of denticles between two lateral groups.
Resembling Clioninae by usually possessing buccal Body length: 16 mm.
cones (two pairs); in some species jaw modified and
rows of isolated peribuccal teeth occur at both sides of Family Pneumodermatidae
jaw; hooks and hook sacs normally developed. Either two lateral sucker arms, or one median and two
Notobranchaea l1Iacdonaldi Pelseneer, 1886 mor- lateral arms. Buccal mass with well-developed radula
pha l1Iacdonaldi Pelseneer, 1886 (Fig. 6.80). Body and pair of hook sacs filled with large hooks. Gills
opaque, brown/yellow to grayish, visceral organs usually at posterior body pole and in middle of right
visible through body wall. Anterior tentacles lateral side. In juvenile specimens bands of cilia are
small. Two pairs ofbuccal cones. Lateral gills rep- present where gills will develop and around hood,
resented by simple protrusions of skin. Posterior covering proboscis and buccal organs. Penis, absent
gill with 3 crests, very broad in living specimens, during female stage, is evaginable and in some spe-
dorsal one with 8 to 10 fringes on each side. Head cies provided with an extra sucker; in species without
rounded, neck contracted. Wings large. FootIobes sucker a special median tubercle appears at base of
well-developed; posterior one long and pointed, posterior footlobe. Ski n usually with chromatophores.
lateral ones triangular, attached to body along 1/3 With acetabuliferous arms. With at least one gill.
of their base. Median tubercle presento Penile
sucker absent. Penis extremely large, with Genus Pnelll1loderllla
unarmed chitinous plates. Long and flexible fla- With both lateral and posterior gills, usually fringed.
gellum at base of evaginated penis, above its PnelllllOderl1la atlanticlll1l (Oken, 1815) atlanti-
attachment penis broadening and its walls becom- Clll1l(Oken, 1815) forma atlanticlll1l (Oken, 1815)
ing lined with transverse thickenings in pairs. Jaw (Fig. 6.82). Body oval to cylindrical, purple to
composed of one line of cusps, 13 to 14 in num- brown, rather transparent, visceral mass visible
ber. Hook sacs shallow, with about 20 short hooks. through body wall. Two lateral arms, each with 80
Radula formula: 8-1-8, median plate unicuspoid. suckers, with short stalks, subequal. Posterior gill
Body length: 10 rnm. hexagonal, with long fringed branches, each with
Notobranchaea tetrabranchiata (Bonnevie, about 7 ramified fringes. Lateral gill short, without
1913) (Fig. 6.81). Body opaque, visceral organs fringes. Posterior footIobe short. Lateral footlobes
visible through body wall. Visceral mas s not horseshoe-shaped, with median foot tubercle.
reaching caudal body poleo Skin smooth, unpig- Penís without sucker. Radula formula: 4-1-4 or 4-
mented. Anterior tentacles small, greyish-green. 0-4. Tricuspid median plate may be absent in older
Buccal cones absent. Posterior gill with 4 undulat- specimens. Hook sacs relatively long. Body
ing but not fringed crests, which converge into a length: 13 rnm.
terminal knob. Head parts short and thick, no clear Pnelll1loderlllQ atlanticlll1l (Oken, 1815) atlanti-
indication of a neck region. Foot composed of two Clll1l (Oken, 1815) forma ellrycotyllllll
lateral lobes with broad bases attached horizontal- Meisenheimer, 1905 (Fig. 6.83). Body oval to
ly near neck region; and rather long, narrow point- conical, semi-transparent, visceral mass visible
ed posterior lobe. Wings narrow at base, broadly through body wall. Lateral and posterior footlobes
truncated distally. Very shallow vestibulum and well-developed, lateral ones long, posterior one
buccal cavity partly surrounded by thick circular rounded. Lateral gill with long lateral and median
muscles. Hooks short, robust, curved, hollow, folds, but without fringes. Posterior gill with 4 re-
pointed, about 14 in each saco Hook sacs shallow latively long branches, with clear fringes. Each
with weakly developed musculature. With "clear acetabuliferous arm bears 16 to 20 suckers. In
cells" dorsal to hook sacs. Jaw composed of row larger specimens 25 to 30 suckers on long stalks,
of conical, somewhat triangular denticles, about arranged in two rows along armo Basal suckers
16 in number. Penis forms a broad folded lobe. larger than lateral ones. Radula formula: 4-1-4.
Pteropoda 689
Median plate seems to persist in older specimens. spirally coiled appendage. Radula formula: 7-0-7.
Body length: 6 mm. Body length: 20 mm.
Pneumoderllla atlanticulIl (Oken, 1815) atianti-
cum (Oken, 1815) forma pygmaeum (Tesch, Genus Pneumodermopsis
1903) (Fig. 6.84). Body rounded, slightly transpa- With two lateral acetabuliferous arms and median
rent, skin usually pigmented. Wings small, posteri- arm, or suckers implanted on buccal wal!. Two sub-
or footlobe long. Two short lateral arms, each with genera: Crucibranchaea and Pneumodermopsis.
up to 8 subequal suckers. Rays of posterior gill o Pneumodermopsis (Crucibranchaea) macrochi-
with two small fringes at their median side. Lateral ra Meisenheimer, 1905 (Fig. 6.88). Body largely
gill small, with fringed, lateral somatic crests. transparent, visceral mass filling whole body, vi-
Rook sacs well-developed, each with up to 100 sible through body wal!. Purple-gray chromato-
hooks. Jaw with 6 spines. Median footlobe blunt. phores may appear on head and foot. Posterior gill
Radula formula: 4-0-4. Lateral teeth subequal with composed of 4 crests radiating from body poleo
square basal plates, outermost ones with larger Median footlobe relatively long, pointed. Median
basal plates. Body length: 3 rnm. tubercle present. Two very distinct lateral arms, at
Pneumoderllla atlanticum (Oken, 1815) boasi top of each a terminal sucker larger than all other
(Pelseneer, 1887) (Fig. 6.85). Body oval, semi- suckers, though similar in structure. Total ofup to
transparent, skin usually pigmented. Wings small, 55 suckers on each arm, in 2 or 3 rows; suckers in
posterior footlobe long. Two lateral arms, each rows sometimes in groups, viz. proximal group of
with 40 suckers. Lateral gill short, with fringes, about 30 suckers and distal group of about 20
lateral somatic crests also fringed. Posterior gill behind single large distal sucker. Median sucker
with short radiating crests with fringes, but not arm completely reduced, represented only by five
subdivided. Rook sacs rather short. Body length: 7 relatively long-stalked suckers. Jaw resembling
rnm. that of P. paucidens. Rook sacs shallow, with 16
Pnellmoderllla atlanticlllll (Oken, 1815) souleye- to 20 rather short hooks. Radula formula: 7-1-7,
ti (Pelseneer, 1887) (Fig. 6.86). Body round, semi- with about 16 transverse rows of teeth. Median
transparent, with numerous glandular tubercles, plate tricuspid, central cusp is smallest. First late-
skin usually pigmented. Visceral mass fills nearly rals with basal cusp projecting toward middle of
whole body. Wings small relative to footlobes. radula. Body length: 8 rnm.
Read relatively large, clearly separated from body. Pneulllodermopsis (Pnelllllodermopsis) brachia-
Median footlobe rather long, lateral lobes well- lis Minichev, 1976 (Fig. 6.90). Body rounded oval
developed, nearly parallel. Median foot tubercle with two circular folds, transparent, visceral mas s
present. Lateral sucker arms of average length. visible through body wal!. Skin with chromato-
Eighty suckers on each arm. Most distal suckers phores. Median arrn with 5 suckers, top sucker
smaller than others, sucker stalks normal in length. larger than other four. Two lateral arms, each with
Lateral and posterior gills present, but rather sim- 12 suckers. With posterior and lateral gills.
ple in shape, not fringed. Posterior reduced to Posterior footlobe short. Rook sacs long, 40 hooks
small ring with 4 short rays. Rooks short, robust, in each saco Radula formula: 4-1-4, 22 transverse
in slender, deep sacs, about 80 in number. rowS. Lateral teeth hook-shaped. Median teeth tri-
Tentacles extremely smal!. Jaw consisting of 30 cuspid with extremely small median cusp. Body
slightly curved denticles. Radula formula: 3-0-3 or length: 1.2 mm.
4-0-4. Body length: 6 mm. Pnellmodermopsis (Pneullloderlllopsis) paucid-
o Pneullloderllla mediterraneulll (Van Beneden, el/s (Boas, 1886) forma pallcidens (Boas, 1886)
1838) (Fig. 6.87). Body oval, skin semi-transpa- (Fig. 6.89). Larger specimens usually thickly spot-
rent, usually pigmented. Wings well-developed, ted with dark chromatophores over whole body
posterior footlobe long. Rather long body with except for wings. Visceral mass visible through
posterior gill with well-developed long fringes, body wall. Posterior gill absent. Lateral gill
and short lateral gill with fringes. Long sucker extremely short. Posterior footlobe short. Median
arms, each with 6-7 suckers, central 4 largest. arm with five suckers, top sucker largest, two
Rook sacs long with 40 slender hooks. Rooks on lower pairs small, attached to long stalks. On buc-
690 Pteropoda
cal wall ten to twelve suckers on short stalks in Genus Spollgiobrallchaea

single row. Penis armed with chitinous style. With 2 lateral and no median arms.
Hooks, only 6 on each side, short and broad. Spollgiobrallchaea allstralis d'Orbigny, 1836
Radula formula: 2-1-2, with 12-22 transverse (Fig. 6.93). Body semi-transparent, long, oval,
rows. Median tooth bicuspid, without central cusp. with numerous dark brown or violet chromato-
Body length: 5 mm. phores which are lacking on gills, footparts, and
wings. Wings oval, narrow at base, pointed distal-
Genus Platybrachilllll ly. Lateral gill more or less surrounds body in mid-
Only one species. die region; posterior gill circular; both gills sim-
Platybrachilllll alltarcticllm Minichev, 1976 (Fig. ple. Anterior tentacles long. Lateral footlobes
6.91). Body conical with two circular folds, semi- attached over entire length. Median footlobe ra-
transparent, visceral mass visible through body ther long and pointed. Two strong lateral arms,
wall. Median arm absent. Two broad lateral arms, median arm absent. Lateral sucker arms bear 7 to
each with 40 suckers. Lateral suckers with short 10 suckers on short peduncles, proximal 3 or 4 re-
stalks. Posterior footlobe narrow and long. latively small. With dorsal gland patch. Hook sacs
Tentacles short. Hook sacs short. Radula formula: small compared to those of Pneumoderma. Hooks,
6-1-6. Lateral teeth claw-shaped, broad-based, about 20 in number, rather long. Jaw composed of
median teeth with two dentated main cusps, two cusps in two groups on median projection of ven-
blunt cusps and rounded comers. Body length: 14 tral side of buccal wall. Radula formula: 8-1-8 in
mID. last rows, 35 rows; with reddish basal membrane
and red median teeth. Median plate developing
Genus Schizobrachilllll later than lateral ones, lacking in first transverse
Only one species. rows. Median plate strong, with one central cusp
Schizobrachilllll polycotylulIl Meisenheimer, and one or two smaller cusps on each side, near
1903 (Fig. 6.92). Body semi-transparent. Two margino First lateral teeth with additional cusp.
dichotomously branching lateral arms, with small, Body length: 22 mm.
very numerous (up to over 1,000) suckers. Median Spollgiobrallchaea i1lterllledia Pruvot-Fol, 1926
arm absent. Posterior gill absent (though small (Fig. 6.94). Body semi-transparent, with chroma-
crests sometimes remain), lateral gill reduced but tophores. Two strong lateral arms; on each arm 18
present. Posterior footlobe, wings and lateral foot- to 20 large subequal suckers (twice the number in
lobes of average size. With dorsal gland patch. S. australis); median arm absent. Posterior gill of
Jaw supported by thin membrane, composed of S. australis type, lateral gill reduced or absent.
numerous slender spines with open space in mid- Posterior end of body cylindrical. Head parts rela-
die. Globular cells under jaw; at right, under radu- tively large. Wings small, median footlobe short.
la, group of smaller cells (probably secretory). Lateral lobes rounded. Median foot tubercle
Hook sacs large, with 20 slender hooks. Radula absent. Dorsal gland patch well-developed. On
formula: 8-1-8. Radula teeth variable in size and each side about 34 robust hooks. Strong and clear-
shape. Median tooth with large basal plate, two ly bent hooks resemble those of Notobranchaea.
cusps, its hollow anterior border ends at both sides Jaw well-developed, composed of numerous rows
in more or les s pronounced comers. Cusp of first of spines, basal spines longer than those on top of
lateral tooth straight or bending slightly toward the odontophore. Radula formula: 7-1-7. Median
median line, whereas cusps of other laterals point radula plate quadri- or tricuspid. First lateral tooth
anteriorly. Body length: 32 mm. structure providing firm connection with prece-
ding and following laterals. Each first lateral tooth
fitting into slit of next one. Denticle, found in S.
australis, strongly reduced or absent. Body length:
20mID.
Pteropoda 691
Limacina (Limacina) helicina antarctica

~ forma antarctica F
a
Limacina (Munthea)
bulimoides
~ Limacina (Limacina) helicina antarctica

forma ran i
(1;
Limacina (Munthea) trochiformis
~ Limacina (Limacina) retroversa australis
Limacina
(Thilea)
lesueuri
Limacina (Thilea) inflata

692 Pteropoda
ventral lateral
view view
ventral oblique view

lateral from left
view
view (notice upper lip)
Cavolinia gibbosa forma gibbosa <In>
(1)
Cavolinia gibbosa forma flava
lateral
view
ventral
view ,11
embryonic
juvenile shell
Cavolinia inflexa
forma imitans <ID Cavolinia inflexa forma inflexa <ID
Pteropoda 693
ventral
view
ventral
view
Cavolinia inflexa forma labiata
Cavolinia tridentata
forma atlantica
ventral
view
Cavolinia tridentata forma australis
f 1 '
,- . 1
.-?ff'\ J
.:>
,
~r(, '~.
g"
juvenile
embryonic
shell
e lateral view
dorsal view
Cavolinia uncinata uncinata forma uncinata <D Diacavolinia angulosa

694 Pteropoda
ventral
ventral view
view <D ventral
view <8> Diacavolinia constricta
Diacavolinia atlantica
Diacavolinia bicornis
ventral
view
<ID ventral
view <D
Diacavolinia deblainvillei Diacavolinia deshayesi
ventral
view <IJ) ventral
view
Diacavolinia e/egans Diacavolinia limbata
forma africana
lateral
ventral view view
Diacavolinia limbata forma limbata <D

Pteropoda 695
d
b dorsal view e lateral view
ventral view
Diacavolinia longirostris
Diacavolinia ovalis
Diacavolinia robusta
detail of rostrum
in ventral view
Diacavolinia strangulata
b;
ventral
view
embryonic
Diacria danae shell
ventral view Diacria major

696 Pteropoda
ventral
vlew
ventral view
e>"
oo'
.'
..... -.'
"
embryonic
.-"
'. d
shells
ventral
view
Diacria rampali
t
·'Ir.'.·.· ..·..'.·
h
",
;
t, e
embryonic
shell
/ holotype
(embryonic
shell)
juvenile
Diacria trispinosa forma trispinosa <D
ventral view
IClio andreae e Clio chaptali e

Pteropoda 697
ventral
view embryonic
shell
Clio cuspidata Clio

piatkowskii
, d~
embryonic
shell
embryonic
Clio pyramidata forma antarctica shell
Clio pyramidata forma excisa
paralectotype
Clio pyramidata forma lanceo/ata Clio pyramidata forma martensi
698 Pteropoda
l' I
d \/
embryonic b
shell
view
embryonic
shell
a Creseís
Clío pyramídata forma sulcata <D Creseís acicula acicula
forma acicula forma
<O clava
<D
c--1
e
dorsal view
\mb'YOO"
shell
Clio recurva
<ID
Creseís
chíerchíae <O
.' I
embryonic
shell a
C
Creseís vírgula forma coníca
G Creseís vírgula
forma constricta
Pteropoda 699
· .~".'
specimen with
embryonic shell
and juvenile caudal section
anached
Creseís vírgula forma vírgula e Hya/ocylis stríata
b
embryonic
shell
embryonic
shell juveniles
a
adult (juvenile
shell broken oíO
Cuvíerína columnella forma atlantica (1))
Stylio/a subula <ID
Cymbulia parvídentata
pseudoconch pseudoconch
I
Cymbulía peroní morpha mínor Cymbulia peroní morpha peroní
700 Pteropoda
a
b entire
specimen
(schematic)
)~.,~/
Corolla ealceo/a <I}I
\ .
'.
oral ."""\.------~. :".'." d

lateral aboral (1''.'\
view entire :'<1
view view r' I ./
specimen 1'.
pseudoconch
@'/
Cymbulia sibogae ~
"~
, .. . -:
.\""..
~,. .' .:~.

•• r .:.~
Corolla eupula •
':"
a b
aboral oral
view view
pseudoconch
........
Corolla intermedia e
whole e
specimen
Corolla ovata Corolla spectabi/is G

Pteropoda 701
whole
specimen Gleba cordata
Desmopterus papilio G
Peraclis bispinosa
Peraclis depressa
Peraclis
moluccensis
Peraclis
triacantha
Peraclis reticulata
702 Pteropoda
Peraclis valdiviae <&>
Paraclione
longicaudata ~
ventral
view
lr '~\"!
r::-
dorsal
view
~V _
Clione Iimacina antarctica
~~
c\""f9~~~~,~
Cephalobrachia
macrochaeta
::;:~0~t~
b '
. -:1<
.:~
",;
.,
f
<8)
~~" ~--r'?:: ..
~ 1':"
~~'";;,..,' ..:;r.~t-·,-\
,r ~.)
):'"
b .f<1~?:1~;.~.:··.'.,
y) : ).~¡ ,\.
;.L~
t.~/·'~ "\ .
'Ü (,:;" '\.~~.
, .,......"t]:';';'.;,
1;. !J ',' "\"?r>-.•..,,.,
'
",,,.,.;
. e
~ l·t~.::.'~.
... ':>,. / .
••. .:- .. ::=.' -- :~ i't.l~~~·•
"" .. ...:..;; .
,
Massya
longicirrata
J~"~ v
/b
e '~"¿:;"'):"",i
~,.~,;
'..
Thliptodon antarcticus
I
contracted
specimen Thliptodon diaphanus
J
e
Pteropoda 703
,; :.:.
);:.:
:'{:" ...
'.
~-:.' '."1 . . •. ~', ",

":.:.;;.
. '1
;,..;-.
", ..•.
.
Cliopsis krohni
lectotype morpha krohni
C1iopsis krohni morpha grandis
C1iopsis krohni morpha modesta <&>

Notobranchaea macdonaldi <!iD>
morpha macdonaldi
Notobranchaea tetrabranchiata <!iD
atlanticum forma
atlanticum
<e
704 Pteropoda
e e
Pneumoderma atlanticum Pneumoderma atlanticum
Pneumoderma atlanticum atlanticum forma boasi
atlanticum forma pygmaeum
eurvcotvlum
·~~"B·.
,,~~~)
C-"
~\;~;j7:í
souleyeti
f, . ')~~
• Pneumoderma mediterraneum
•
fá~
¡t;;¿i;\(
• '~jo'
' "~. '
>-)¡,,'"
,{~',I',,""A "",
1r
P,',',",,".,
1",'.... ,-'~.
--4
Pneumodermopsis
~~.~~;;;\~q;'j
•
•
(Pneumodermopsis)
brachialis
Pneumodermopsis
(Crucibranchaea) cV~~:~.' juvenile
macrochira Pneumodermops;s (Pneumodermopsis)

paucidens forma paucidens
Pteropoda 705
a
~~f.·.f
~. ::.:-'::.".)
Platybrachium
antarcticum
b\/
-t
Spongiobranchaea
Schizobrachium
polycotylum e australis
Fig. 6. IlluslTations of pteropod speeies. Figure sourees:

From Boltovskoy (1973b): 2b, 5b, 7e, 12a, 36d; from Boltovskoy (1975e): 16b, 16e; from Gilmer and Lalli (1990): 71e; from MeGowan (1968):
61e; from Newman and Greenwood (1987): 34d; from Rampal (1996): 58a; from Riehter (1976): 46b; Riehter (1979): 17b, 17e, 17d, 26b, 26e,
26d, 55e, 64e; from van der Spoel (1967): lb, 9a, 9b, 10e, Ila, Ilb, 11e, 13b, 13e, 29d, 38b, 38e, 39b, 3ge, 40b, 40e, 42a, 43d, 43e, 47e, 48b,
49b, 50b, 50e; from van der Spoel (1969): 16d, 16e; from van der Spoel (1973b): 75b; from van der Spoel (1976): 3e, 25b, 25e, 30b, 30e, 41b,
52a, 53a, 53b, 54b, 56a, 56b, 56e, 56d,57b, 59a, 59b, 5ge, 62a,64b,65b,67b,67e, 68b, 72b, 73b, 74b, 76b, 77b, 78, 79b, 80b, 8Ib,82b,86b,
87b, 88e, 89b, 8ge; from van der Spoel (1981): 12e, 12d, 12e, 12f, 14b, 14e, 15b, 15e, 16f, 16g, 29b, 2ge, 31e, 33e, 33d, 33e, 33f, 34a, 34b,
36e, 38d, 39d, 40d, 42d, 43e, 44b, 47a, 4ge, 49d, 51 b, 52b, 52e, 52d, 60a; from van der Spoel (1992): 70b, 70e; from van der Spoe1 and
Boltovskoy (1981): 31 b, 32b, 33b, 34e, 35b, 35e, 42b, 42e, 43b, 55b, 63b. 69b, 71b, 88b, 92b, 93b, 94b.
706 Pteropoda
Suggested readings Holkema, Utrecht, pp. 1-484. An updated revisioll o/

these pelagic molluscs, with detailed data on iden-
Bé A.W.H., Gilmer A.W. 1977. A zoogeographic and tification. mOlphology. anatm. and distribution.
taxonomic review of euthecosomatous Pteropoda. van der Spoel S., Boltovskoy D. 1981. Pteropoda. In
In "Oceanic Micropaleontology" (A.T.S. Ramsay, "Atlas del Zooplancton del Atlántico Sudoccidental
ed.), Acad. Press, London, pp. 733-808. Excellent y m-étodos de trabajo con el zooplancton marino"
review o/ the world-wide distribution and c!assifi- (D. Boltovskoy, ed.), Publico Esp. Inst. Nac. Inv.
cation o/ Euthecosomata. Desarrollo Pesq., Mar del Plata, pp. 493-524.
Lalli C.M., Gilmer R.W. 1989. Pelagic snails. The General review o/ the systematics and distribution
biology of holoplanlctonic gastropod mollusks. o/ the pteropods o/ the Southwestern Atlantie
Stanford Univ. Press, Stanford, pp. 1-259. All Ocean. Includes brie/remarks on biology, methods
aspects o/ pelagic mollllscs. inclllding the ptero- o/research, histoy o/study. etc. (in Spanish).
pods, are covered in this manual. van der Spoel S., Newman L., Estep K.W. 1997.
van der Spoel S. 1967. Euthecosomata, a group with Pelagic molluscs of the world. World Biodiversity
remarkable developmental stages (Gastropoda, Database, CD-ROM Series. Expert Center for
Pteropoda). 1. Noorduijn and Zoon, Gorinchem, pp. Taxonomic Identification (ETI), Amsterdam, The
1-375. A major monograph on the biology, morphol- Netherlands, UNESCO, Pans. An electronic gllide
ogy. taxonomy. and distribution o/ Ellthecosomata. /01' the identification and distribution o/the pelagic
van der Spoel S. 1976. Pseudothecosomata, Gymno- mollllscs o/ the world.
somata and Heteropoda. Bohn, Scheltema and
Cephalo·poda
Kir N. Nesis
General characteristics of planktonic cephalopods fol' fish larvae and juveniles. It is best to tow nets at
low ship speeds. Vertical plankton tows are not rec-
Early developmental stages of various coleoid cephalo- ornmended because the low volumes of water usually
pods may be found in the plankton, including: squids filtered normally result in only very small numbers of
(Teuthida), both neritic (Myopsida) and oceanic larvae, and fail to capture even the juveniles of active-
(Oegopsida); some cuttlefish (Sepiida: Spirula and some Iy swimming cephalopods in sufficient quantities. For
members of the family Sepiolidae); some benthic octo- larvae, the Bongo net 01' its various analogues may be
pods of the genera OCfOpuS, Scaeurgus, ElIGXOCfOPllS, recornmended. Juvenile stages of active nektonic
Eledone (Octopoda, Octopodidae); and all developmen- squids, and almost all stages of semiplanktonic squids
tal stages of Vampyromorpha and pelagic octopods (7 and pelagic planktonic octopuses, may be caught by
families of the Octopoda). Most adult squids belong to various midwater trawls, such as the Isaacs-Kidd
the nekton (some deepwater forms are semiplankton- Midwater Trawl (IKMT) and similar nets. Fine-
ic/seminektonic), cuttlefish to the nekton 01' nekto-ben- meshed nets (5 01' 10 rnm knot-to-knot) should be
thos, and octopodid octopuses to the benthos. inserted either in the cod-end 01', in some types of
IKMT, along the whole length of the trawl. For sub-
Newbom cephalopods are termed either "juveniles", if adult and adult pelagic cephalopods the best devices
they resemble small adults, 01' "paralarvae" (in are large cornmercial pelagic otter-trawls (01' pelagic
malacologists' jargon) 01' "larvae" (in ichthyologists' versions of bottom otter-trawls) with a fine-meshed
jargon), if they are different from the adults and have section at the cod-end. When using fine-meshed nets
some larval features. The terms "paralarva" and "larva" in a commercial trawl, permission from the local fish
are equivalen!. malacologists using "paralarva" simply protection agency should be requested.
to distinguish cephalopods from those molluscs that
undergo full metamorphosis (Nesis, 1979b; Young and For better preservation and easier subsequent identifi-
Harman, 1988). 1 will use the term larva in the same cations, irnmediate sorting of the catch prior to fixa-
sense as it is used in ichthyology. tion is recommended, although this is rarely possible
when dealing with standard plankton catches.
Identification of larvae and early juveniles to species
level is difficult, 01' even impossible in the case of very Fixation and preservation
early larvae, because species-specific characters usual-
ly develop only in the postlarval stage. The use of iden- Usual methods of fixation and preservation of cepha-
tification keys for adults is impossible, and keys speci- lopod larvae and juveniles are the same as those for
fically for planktonic stages are problematic because larval and juvenile fishes. Fixation of freshly caught
taxonomically important characters develop in specimens with 4% formalin and subsequent transfer
sequence. Thus, different keys would be required for to 70% ethyl alcohol is normal. But specimens may be
representatives of the same species depending on their kept in formalin for an unspecified time. Sorted and
size. Usually, precise identification to only the generic fixed larvae may be kept in 2% formalin. When work-
level is possible. ing with juveniles and subadults it may be better to
spread specimens immediately after capture and fix
them after spreading, but this operation is impossible
Methods when working with plankton samples containing lar-
val cephalopods. Detailed instructions for fixation,
Collection preservation, and preparation of cephalopods are
described by Roper and Sweeney (1983).
Methods of capture of planktonic life stages of
cephalopods do not differ from those recornmended
SOl/lh AIlanlic Zooplankton.

edited by D. Boltovskoy. pp. 707-795
1999 Backhll)'s PlIblishers, Leiden, The Nelherlands
708 Cephalopoda
Geographic and vertical distribution Okhotsk and Bering seas. But rich comrnercial squid
fisheries are known on both sides of the South
Planktonic cephalopods, and planktonic developmen- Atlantic: off Argentina and the Malvinas (=Falkland)
tal stages of nektonic, nektobenthic, and benthic Islands in the west, and offNamibia in the east. Larval
cephalopods, are distributed through the World and juvenile cephalopods are generally at maximum
Ocean, from the North Polar Basin to the Antarctic. abundance in epipelagic layers, decreasing at greater
But, except for larvae of nearshore benthic octopuses, depths. Adults are usually most abundant in mesope-
they are nearly absent over the inner shelf and rare lagic layers, particularly where they contact the bot-
over the outer shelf. Their main habitat is oceanic tom along slopes and rises. In terms of the concentra-
waters. Meridional ranges of most species of neritic tion of cephalopods under one square kilometer of
and nerito-oceanic (living over the continental slope) ocean surface, maximum abundances always occur
cephalopods extend along shelves and slopes, some- over the shelfbreak and continental slope, and ocean-
times across climatic belts. On the other hand, most wards from trade wind upwelling areas, while they are
oceanic cephalopods are zonally distributed, their 10w over the inner shelf and in the central part of anti-
ranges elosely associated with the large oceanic circu- cyelonic gyres (Nesis, 1985).
lation systems. Their range boundaries in many cases
coincide with global oceanic fronts and convergences, Two typical examples illustrate cephalopod abun-
such as (in the South Atlantic) the Southern Tropical dance in pelagic areas. In large Bongo net (mouth
Front, the Southern Subtropical Convergence, and the diameter 60 cm, mesh 0.7 mm) samples from the
Antarctic Convergence. The geographic distribution upper epipelagic ¡ayer (O to 50-55 m) ofthe open Gulf
of oceanic cephalopods and their types of ranges are of Guinea during June-August of 1985 there were, in
listed and mapped by Nesis (1985). different water masses, from 10 to 36 cephalopod lar-
vae per 1000 m3 (Arkhipkin et al., 1988). In midwater
Planktonic cephalopods are distributed from the sur- trawl samples from the outer shelf and slope off
face to abyssal depths, and inelude epipelagic, meso- Namibia (l7-2rS), the biomass (wet weight) of
pelagic, and bathypelagic species, as well as various cephalopods was 80.1 g per trawling hour in auturnn
intermediate types. Most speciesperform diurnal ver- (April-June), and 31.5 g h-I in summer (January), or
tical migrations, and many descend during ontogeny. 8.1 and 2.4% of the total biomass of fish, cephalo-
Larvae of almost all species live on the average elos- pods, decapod crustaceans and euphausiids, respec-
er to the surface than adults. Even late larvae may per- tively (5-10 times less than fish biomass). These were
form diurnal vertical migrations (usually within the Isaacs-Kidd and Aseyev-Samyshev trawls towed ho-
epipelagic realm). Diurnal migrations are ofthe usual rizontally at levels from 50 to 1000 mover bottom
type, with nighttime ascent. Only the juveniles (not depths of 105-4700 m, 15 to 200 nautical miles from
larvae nor adults) of some oceanic ommastrephid shore (Nezlin and Nesis, 1988).
squids (mostly Ommastrephes and Sthenoteuthis) per-
form reverse migrations, with daytime ascent. The The biomass of cephalopod larvae is always very sig-
vertical distribution of oceanic cephalopods is dis- nificantly lower than that of, say, planktonic crusta-
cussed by Roper and Young (1975) and Nesis (1985). ceans, and usually may be considered insignificant. But
the biomass of adult cephalopods in mesopelagic layers
Quantitative distributional data on planktonic cepha- in some very productive areas ofthe World Ocean may
lopods are scanty, first because no device catches all be comparable to that of fish (Clarke, 1996).
developmental stages with equal efficiency, and second Furthermore, the life span of an average cephalopod is
because trawls that effectively catch adult cephalopods much shorter than that of an average fish, and the
are non-quantitative. The only available information is growth rates, food rations and production:biomass
estimates of relative abundance (biomass) of pelagic (P:B) coefficients are correspondingly much higher
cephalopods, and of their relative distribution in the (Nesis, 1985). Thus, the role of cephalopods in total
World Ocean (Clarke, 1987; Nesis, 1985). production and consumption of food in the ocean is
much more important than their share of biomass may
The South Atlantic is not as rich in cephalopods as, suggest (Boyle, 1983-1987; Nesis, 1985; Clarke,
for example, the subarctic North Pacific and the 1996).
Cephalopoda 709
Table l summarizes the known horizontal and vertical ing their juvenile or maturing stages, but the larvae
distribution ranges of all planktonic cephalopods in always have tentacles.
the South Atlantic.
The mouth, with a parrot-like beak, is located in the
centre of the crown of extremities. A funnel connect-
Taxonomy and identification ing the mantle cavity with the environment is located
under the head (Fig. lE). The mantle is connected
Morphological characteristics of planktonic with the head and funnel by three locking cartilages:
Cephalopoda (refer to Fig. l and the Glossary below) one nuchal (occipital) and two mantle/funnel ones
(Fig. lE, G). In Octopoda the mantle is fused with
The body of coleoid cephalopods is divided into the both head and funnel, leaving only one ventral split
head, with extremities, and the body covered by the under the funnel, or two splits on the lateral sides of
mantle (Fig. lE). The dorsal mantle length (ML), the head (Amphitretidae). In the squid family
from anterior to posterior end along the dorsal mid- Cranchiidae the mantle is also fused but lea ves three
line, is the standard measurement of cephalopods. In splits, on the lateral and ventral sides of the head. In
octopuses lacking a clear boundary between head and some Sepiolidae, and in the squid family
mantle, the ML is measured from the midpoint of a Promachoteuthidae, the mantle is fused with the head
line connecting the centers of the eyes. in the occipital region only, and in postlarval squids
Grimalditeuthis (fam. Chiroteuthidae) it is fused with
The head bears two large, well developed eyes (Fig. the funnel. The funnel-locking cartilage is a cartilagi-
l E, K, M) which may be covered by a transparent skin nous pad with variously shaped furrows, grooves, or
(Sepiida except Spirulidae, Myopsida, Octopoda), or pits. It may be simple, with straight or slightly curved
freely comrnunicate with the environment (Spirulidae, furrow, widened posteriorly, or of more complex
Oegopsida, Vampyromorpha). The extremities (8 form: U-like, ear-shaped, triangular, etc.; the mantle-
arms in octopods, 8 arms and 2 tentacles in squids and locking cartilage is a cartilaginous ridge, knob or
cuttlefishes) are located on the anteriormost body end swelling that locks into a corresponding furrow in the
as a crown around mouth (Fig. 1E). Arms bear suckers funnel cartilage and thus looks like its negative. A
along their entire length; in some adult squids some glandular funnel organ (Fig. IH) is located inside the
suckers may be transformed into hooks. Tentacles con- funnel, on its dorsal and lateral surfaces. It consists of
sist of a more or less cylindrical stem and a widened a medial component (in the form of an inverted V)
and flattened club (Fig. l C, S). OnIy the club, in adult- and two lateral components. The medial components
hood, bears suckers, and sometimes hooks. Larvae are may bear ridges and papillae, important features in
always devoid ofhooks. The suckers in squids and cut- the taxonomy of some squids (Histioteuthidae,
tlefishes are seated on short thin stalks and equipped Cranchiidae). Its form (W, VV or IIII-like) is also
with a horny ring, usually with a dentate rim. In vam- important in the taxonomy of octopods. The funnel
pyromorphs and octopods there are no stalks or horny valve, a semilunar muscular flap, is located on the
ring. Arms and tentacles of squids and some cuttlefish- dorsal funnel surface near its opening; it may be
es bear, on the outer sides, longitudinal skin keels absent in some squids.
(swimming keels), and on the inner sides, besides
suckers, thin protective membranes supported by mus- A pair of fins is located on the posterior end of the
cular trabeculae (Fig. IC, O). In some cephalopods the body (absent in incirrate octopods). The juveniles of
arms are joined by a film (umbrella) along their bases Vampyromorpha have two pairs of fins at certain
(Fig. 2.25D). The arms are numbered from the first to developmental stages, one larval (which disappears
the fourth pair beginning from the dorsalmost paie. during growth), and one definitive (Fig. 2.60D, E).
Tentacles in squids and cuttlefishes are located between
the third (ventrolateral) and the fourth (ventral) pairs The inner shell (gladius, or pen, Fig. ID), which may
(Fig. lE). In some species one or both arms of mature be calcified (Spirulidae, Sepiidae) or chitinous, is loca-
males are modified into a section called the hectocoty- ted under the mantle on the dorsal side. It is extremely
lus (Fig. lB, J, N). Some, mostly deepwater, squids reduced or lacking in Octopoda, some Sepiolidae and
lose their tentacles at the end ofthe larval stage, or dur- allied families, and in the Promachoteuthidae.
35ºN-30º5 -~ ~ +
"Adults ~ ~~
ooomp-bp
35ºN-? ~45-50º5-70º5 Ecologic ,. ~
I
mp-bp
no
710 boundaries)
53ºN-70º5
ep oob¡::>-ap
o/no
mp-bpno
lep-mp-b
lep-mp
ler-mp oosb
usb
o/no o45ºN-45º5
45º5-60º5
35-43ºN-35-43º5
ep-mp
ep 40ºN-30º5
o35-40ºN-35-40º5
ono 45ºN-30º5
35ºN-30º5
40ºN-40º5
43ºN-35º5
20-30ºN-40-45º5
35-45ºN-35-40º540ºN-10º5
35ºN-20º5
55ºN-23º5
40º5
20ºN-5º5?
62ºN-25º5
20ºN?-40-45º5
40ºN-30º5
nmp-bp-b
nlep-mp-bp 10º5-35-40º5
30-35º5?
35-42ºN-25º5
50-65ºN-30-45º5
o10º5-35º5
ep-m¡:>
63ºN-40-45º5
mp-b
?mp-bp
e¡::>-mp 8ºN-50º5
27º5-33º5
ep19º5-35º5
(approximate
?ep-mp-bp
?ep
ep-mp
mp
ep-mp
ep-mp
er
ep-mp
epep-mp
e~mp
+
t-st
sst t-st
bst
epandI sst-ntnt-an
t-nstan AIWP
WA-EA
WA
CG
t-st-bor-nt
t-st-nt
t-st-bor-nt
t-st5EA
5WAfr
t-st
CGCG
WA-EA
AO
Vertical CG
CG
IWP,5WAfr
AO,t-st
spAO, CG 55-60ºN-45-50º5
distribution
IWP
tlWP Cepha1opoda
¡I+l- 1ep-mp
l
+
lep-mp
bipCG ep-mp-b
setting
-+ bp lep-mp-bp
e~mp
lep-mp-bp
bp
mp-bp lep-mp
lep-mp-bp
mp-bp
Worldwide
juveniles
larvae
aeq/bst
eq-sst-sp
an CG
ep-mp
~G CG
CG + 55º5-70º5 ¡
cta l-
jI r me:bp r
c
I
orizontal distribution
ep swn Isb-b
Isb-ubsb
usb Adults Ecologic
-+
uth Atlantic
nermis
mp-bp ep-mp?
ep-mp-b
mp-sb-b
o/no
setting
-+
56-60º5-70º5
dn
mp-bp
nmp-bo35-40ºN-30º5
no
omp-b
o/no
an CGep-mp-Is-b
50ºN-21
40º5-70º5
25-30ºN-35º5
45-50º5-70º5
nomp-bp
20-30º5-45º5
50-55ºN-30-35º5
40ºN-?
40-50º5-70º5
45ºN-?
5º5-43º5
20-30º5-47º5
50ºN-30-35º5
30-35ºN-25º5
o45ºN-36º5
25ºN-35º5
37ºN-35º5
58ºN-45º5?
37º5-60º5
nolep-mp-bp
50ºN-30-35º5
ep-mp-b
no obp 28ºN-56ºN
60ºN-22-23º5
60-65ºN-40º5
40ºN-30º5
10-25ºN-5-1
mp-b
(approximate
o/no
mp-bp
lep-mp ep-mp
mp-bp ep
epep
ep
ep-mp
ep,s
ep-mp
mp-bp
lep-mp-bp
bp
ep-mp-bp
mp-bp-b
mp-bp ep-mp
ep-my-bp
mp-bp
ep-mp-bp
e2-
ep?
ep-11'1Jl
35-40º5-50-60º5 I
mpº5 5º5
35º5-54º5
28º5-46º5
10º5-35º5
35ºN-22º5
35ºN-35º5
23º5-54º5
20º5-45º5
and1
nst-bor
Worldwide
larvae
juveniles
t-nst-bor
t-st
sst-nt
tt-st
sst-ntAIWP?
5WA-5EPO
t-st-bor
nt-an
t-st
sst-nt
t-st-bor-nt
an
tAO
5EA-5WIO
5WA
t-st
nt WA-EA
WA-EA
AIWP
EA AO
5WA
AIWP
Vertical
t-st(bet)
bet CG
AO
WA
CG
AO,PO
ep CG
CGAO
AOdistribution 711
Cephalopoda
1
~ Adults
~ bp-b
712mp-bp-b no ;
rlep-mp
usb e¡:>-mp-Is-b
oSO-60ºN-SO-SSºS
oo34-36ºN-3SºS
noolep-mp
Isb-ub
sb-ub
usb
o/no
mp-bp lep-mp-bp-b
lep-mp-bp
lep-mp-b dn
dn ooo4SºN-2S-30ºS
nmp-bp-b
bnep
nmb-mp
obp-ap?
ep-mp ns4SºN-40ºS
4S-S0ºN-3S-4SºS
40-4SºN-3S-40ºS
SS-60ºN-4S-S0ºS
SO-SSºN-40-4SºS
mp-bp 43ºN-3SºS
32ºN-l0ºS
40ºN-30ºS
o27ºS-28ºS
30ºN-3SºS
40ºN-30ºS
?36ºN-33ºS
20-2SºS-40-4SºS
30-3SºN-3SºS
3SºS-SOºS
40-4SºN-4S-S0ºS
1 Isb-ub,s
no no
obp
mp ep?ep-mp
SWAfr
SOºN-2SºS
40ºN-3SºS
42ºN-2S-3SºS
32-40ºN·30ºS
20ºN-20ºS
4SºN-3SºS
40ºN-37ºS
30ºS-4SºS
lSºS-28ºS
32ºN-1OºS
42ºN-3SºS
Is-ub
oep-mp
(approximate
ap
mp-bp ep
ep
ep-mp
ep-mp-bp
lep-mpep-mp
epep,s
ep
mp-bp
Ecologic
t-st
t-st
+
tt-st
WA-EA
t-st
t-stAIWP
?WA
Vertical
t-st
sst-nt
sst-nt
t-st
bst
nt
bct/bst
aeqtAO IW
EA
WA-EA
AIWP
CG
EAO
CG
AO
CGCGdistribution
SEA
CG
CGCG Cephalopoda
tic
¡t
setting Worldwide
juveniles
larvae and
ep-mp
Horizontal distribution
Cephalopoda 713
The tissues encircling the mouth in squids and cuttle- of short and rapid restructuring which may be accom-
fishes are connected with the arm bases by the buccal panied by rejection or reduction of some provisional
membrane, with ray-like buccal connectives, the rays larval structures (Gonatopsis, many Chiroteuthidae).
numbering 6, 7, or 8 (Fig. lA). Connectives leading to In general, larvae differ from juveniles and subadults
the arms of the 1st and 2nd pairs are attached to the by their short and narrow fins, short arms, sometimes
dorsal arm surfaces, those ofthe 3rd pair to the ventral by the absence of some arms in newbom larvae, by
ones (except in the enoploteuthid genus Enigmoteufis), lack of arm and tentacular hooks and of photophores,
and those ofthe 4th pair can connect either dorsally or and by presence of stalked eyes, arm crown pillar, etc.
ventrally. The last character is taxonomically impor-
tant. Among squids the buccal connectives are attached
to the dorsal sides of the 4th pair of arms in 8 families, Glossary
and to their ventral sides in 16 families.
The following glossary of most commonly used terms
Many cephalopods, particularly squids and for the description of cephalopods is based chiefly on
Vampyromorpha, have photophores, sometimes very Sweeney et al. (1992) after Roper et al. (1984), with
numerous, complicated and variable in structure. modifications (refer to Fig. 1).
Their distribution on the body is strongly specific and
may be a very valuable (sometimes almost the only) Antitragus: a small knob-like cartilaginous projection
taxonomic character. During ontogeny some photo- fram the posterior wall of the funnellocking-car-
phores, or groups of photophores, develop at different tilage in some squids of the families
times (i.e., at different body sizes), and this may be Chiroteuthidae and Mastigoteuthidae (see tragus)
useful for identification. (Fig. 2.42G).
Arms: the eight circumoral appendages in coleoid
The color of cephalopods is determined by chroma- cephalopods (Fig. lE, V). One pair of modified
tophores (Fig. IV-X) which may be black or brown, appendages called tentacles lies between the ventro-
red, orange, or yellow. Their distribution on the body lateral (3rd pair) and ventral (4th pair) arms in cut-
of larvae is sometimes stable and characteristic, but tlefishes and squids; one pair of modified, greatly
light-colored chromatophores fade quickly in forma- reduced appendages, called retractile filaments, lies
lin and be come indistinguishable. between the dorsal (1st) and dorsolateral (2nd) arms
in Vampyromorpha.
Ontogenetic changes in cephalopods proceed continu- Arm crown: all circumoral appendages (arms and ten-
ously and gradually, without sudden metamorphosis. tacles).
In some genera the larvae resemble small adults Arm-crown pillar (=arm-crawn stalk, brachial pillar,
(Loligo, Bafhyteufhis, Japetella). In others, however, snout): an elongation ofthe head between the eyes
they are very different, and ontogeny includes stages and the arm crown in many larval and
•••
Table 1. Geographic and vertical distribution of the species recorded in the South Atlantic (the latitudinal boundaries indicated are approxi-
mate). Abbreviations:
Horizontal distribution, worldwide:
aeq: anti-equatorial; an: Antarctic; bct: bicentral; bip: bi-peripheric; bor: boreal; bst: bi-subtropical; eq: equatorial; nst, sst: northem, south-
ern subtropical; nt: notalian (subantarctic); sp: southern peripheric; st: subtropical; t: tropical.
Horizontal distribution, South Atlantic:
AIWP: Atlanto-Indo-West Pacific; AO: Atlantic Ocean; CG: circurnglobal; EA: Eastern Atlantic; 10: Indian Ocean; IWP: Indo-West Pacific;
PO: Pacific Ocean; SEA, SWA: Southeastern, Southwestern Atlantic; SWAfr: Southwest Africa (Namibia and Western Cape); WA: Western
Atlantic.
Vertical distribution:
ap: abyssopelagic; b: bathyal; bp: bathypelagic; ep: epipelagic; lep: lower epipelagic; Is: lower sublittoral; mb: mesobenthic (bathyal and near-
bottom bathyal layers); mp: mesopelagic; s: surface and near-surface; sb: sublittoral; ub: upper bathyal; usb: upper sublittoral.
Ecologic setting:
dn: distant-neritic (o ver and near slopes and seamounts); n: neritic (shelfareas); no: neritic-oceanic (slopes and nearby areas); o: oceanic; swn:
shallow-water neritic.
714 Cephalopoda
arm 1 (dorsal)
lappet~\; .. Q 8.
buccal ,,\ ,.filM~Q~~g arm 2
, •• ~~. .. ~fI .. 7hOO"
Iccal G
<1
,.~ "
.. "'. arm 3
~.~.._~ ~l'6",k ,
:,, •• 1, \
\'.; ..
stalk of
attac~~~ /7f;r~.J arm
l. 4
club buccal (ventral) buccal
connective suckers
(ventrally
attached)
arm crown (squid)

8
Ilgula calamus sucker
¡,J~~t-
-.,~.' ~
@
@~@
;~
(i) sperm groove

t=--
hectocotylus (octopUS)
r ,"",1 m,mb,,",
-o
'"
Q)
Le
rachis~ 1
v
vane~
j-T
-v
S owmg malO
r 11
•• 11-.......",
I tail diagnostic
1_1features,
gladii (squids)
hy~oth.etic sq!Jid
t
f «)
in ventral view
ú)
Cephalopoda 715
Todarodinae o funnel-Iocking cartilages
o funnel groove
Ommastrephidae
o gill lamellae
funnel organ
photophores
of squid eye
orbital sinus
(squid) f) hectocotylus (squid)
lamellae
(modified
trabeculae)
hectocotyl ized
male arms
(squid)
o photo~hore~ on squid
Intestlne
arm suckers and
protective membrane
716 Cepha1opoda
od~~ó~~~ ~~I¡¡!¡:::: ¡iljflf;¡¡¡¡Iii~
G ctenodont radula
O spermatophore
(!)
uk~6~q heterodont radula
web
suckers in
longitudinal
\
rows 8suckers
in (oblique)
transverse
rows
~":';"';""":""""' .. ,
squid
t'~:~::
teeth
" club suckers

o sucker rings octopus
pores
water pores
o
primaryarm
suckers
Dosterior
llcapll
(octopus, 3 or 4)
G (mantle
end)
dorsal view ventral view
octopodid chromatophore patterns
primaryand
first secondary
o arm suckers
(octopus)
Cephalopoda 717
juvenile squids, particularIy Brachioteuthidae, tophores located well beneath the integument,
Chiroteuthidae and Cranchiidae (Fig. 2.42A-F, usually covering the ¡nner organs of the digestive
2.49A, B, 2.59A-C). system (Fig. 1V).
Branchial heart: a globular thick-walled muscular pro- Chromatophore, tegumental: the chromatophores locat-
pulsive organ located at the base of each gil!. ed in or near the surface integument (Fig. IV-X).
Buccal connectives: muscular rods connecting the Circumoral appendages: the 8 arms (squids, cuttle-
supports of the buccal membrane to the bases of fishes, and octopuses) and 2 tentacles (squids and
the arms (Fig. lA). cuttlefishes) that arise from the head and encircle
Buccal lappet: small subtriangular flap formed by the the mouth of cephalopods (Fig. lA).
tip of the buccal membrane support and the Club: see tentacular club.
adjoining buccal membrane; may bear suckers Cone (=conus): the spoon- or cup-shaped conical pos-
(Fig. lA). terior end of the gladius or cuttlebone; homologous
Buccal membrane: thin web of tissue that encircles to the phragmocone of fossil teuthoids (Fig. ID).
the mouth, reinforced by six to eight buccal sup- Comeal membrane: the very thin, turgid, transparent
ports (Fig. lA). membrane that covers the eyes of myopsid and
Buccal membrane supports: longitudinal muscular sepiid cephalopods (Fig. lE).
rods fused with the buccal membrane (Fig. lA). Dactylus: the distal terminal section of the tentacular
Buccal suckers: small suckers on the buccal lap- club, often characterized by suckers of reduced
pets/membrane (Fig. lA). size (Fig. lE).
Calamus: the conical papilla on the octopod hectoco- Fins: the muscular flaps located along the lateral or
tylus at the distal end of the sperm groove, distal dorso lateral surfaces of the mande of sepiid, teu-
to the last sucker and proximal to the ligula (Fig. thid, vampyromorph, and cirrate octopod cephalo-
lB) (see ligula). pods; used for locomotion, steering, and stabiliza-
Carpal cluster (=carpus, carpal pad, fixing apparatus): a tion (Fig. lE).
usually distinct group of suckers with smooth ring Fin lobe: the portion of each fin that protrudes anterior-
and knobs on the carpus ofthe tentacular club (Fig. Iy from the anterior point of attachment of the fin to
IC, E). the mande (Fig. lE).
Carpal knobs: small rounded hemispherical protube- Fixing apparatus (=carpal cluster): a usually distinct
rances on the carpus, to which carpal suckers from group of suckers with smooth ring and knobs on
the opposite club adhere during locking of the the carpus of the tentacular club (Fig. 1C).
clubs (Fig. 1C). Foveola: transverse membranous fold of ski n that
Carpal suckers: smal! suckers with smooth rings on the forms a pocket in the funnel groove of some
carpus of the club, which usually adhere to knobs ornmastrephids (Fig. IF) (see side pockets).
on the opposite carpus during locking of the clubs Funnel: the subconical tube on the ventral side of the
(Fig. IC). head through which water is expelled from· the
Carpus: the proximal zone of suckers and/or knobs on mande cavity during locomotion and respiration;
the tentacular club (Fig. 1C, E). reproductive and waste products, and the ink also
Cartilage: a solid concentration of connective tissue- pass through the funnel (Fig. lE, V).
derived material occurring in the funnel-mantle Funnel groove: the depression in the posteroventral
locking apparatus, nuchal attachment, integumen- surface of the head, in which the anterior portion
tal "scales", cranium, etc. ofthe funnellies (Fig. IE-F).
Chromatophore: an organ consisting of a pigment- Funnel cartilage (=funnellocking cartilage): the carti-
filled sac with associated muscles and nerves that laginous pad that contains variously shaped
provides much of the background color, color grooves, pits, pockets, or depressions on each ven-
pattems, and color changes of cephalopods (Fig. trolateral side of the posterior part of the funnel,
IV-X). which joins with the mande component to lock the
Chromatophore, extrategumental: the larval chroma- funnel and mande together during locomotion
•••
Fig.1. Morphologic details and characters used for the identification ofCephalopoda. From Sweeney et al. (1992).
718 Cephalopoda
(Fig. lE, G) (see mantle cartilage, mantle locking includes the calamus in basal part and usually a
cartilage ). series of transverse ridges and grooves on the oral
Funnel organ: the glandular structure inside of the surface (Fig. lB) (see calamus, hectocotylus).
funnel, generally a single W-shape form in octo- Mantle: the fleshy muscular, tubular, or sac-like body
pods, and a dorsal inverted V -shape component of cephalopods; contraction provides propulsion
with opposed ventral oblong components in deca- through jet-like expulsion of water, as well as res-
pods (Fig. lH). piration; contains the viscera (Fig. lE, V).
Funnel val ve: the semi lunar muscular flap, a one-way Mantle length (ML): in decapods measured dorsally
val ve, on the inner, dorsal surface near the distal from the anteriormost point of the mantle to the
opening of the funnel (Fig. lH). Absent in some posterior apex of the mantle, or to the tip of the
squids. fused fins, whichever is longest. In octopods
Gill lamellae: the leaf-like convoluted individual measured dorsally from the midpoint between the
components of the gill through which gas eyes to .the posterior end of the mantle.
exchange occurs (Fig. 11). In decapods and octo- Mantle cartilage (=mantle locking cartilage): the car-
pods (other than cirrates) the inner and outer de- tilaginous ridge, knob, or swelling on each side of
mibranchs of the gills are attached to a narrow the ventrolateral, internal surface of the mantle
central axis and are typically arranged vertically in near its anterior end that locks into a correspond-
two diverging rows, or oriented parallel to the gill ing funnel cartilage during locomotion (Fig. lE)
axis (lamellae perpendicular to axis). In cuttle- (see funnel cartilage, funnellocking cartilage).
fishes the gills have free lamellae (not attached at Manus: central portion of the club between the dacty-
the tip) and no branchial canal. lus distally and the carpus proximally (Fig. lE).
Gladius (=pen): the feather or rod-shaped chitinous Nidamental glands: a pair of large tongue-like (in
supporting struCture in the dorsal midline of squids) or oval (in cuttlefishes) female glands, in
squids and non-sepiid cuttlefishes; homologous to mature females occupying the whole middle ofthe
the shell of ancestral forms (Fig. ID). ventral side of the mantle cavity. They secrete the
Hectocotylus: in males, an arm or arms modified for external egg envelope either surrounding each egg
transferring spermatophores to the female; modifi- or uniting eggs in gelatinous capsules. Absent in
cations may involve suckers, sucker stalks, protec- Octopoda, Vampyromorpha and Enoploteuthinae.
tive membranes, trabeculae, and arm shape (Fig. Nuchal cartilage: the oblong cartilaginous plate on the
lB, ]). Not all species have a hectocotylus (see posterodorsal surface of the head of most squids
calamus, ligula). and cuttlefishes that articulates with a complemen-
Hooks: chitinous claw-like structures ontogenetically tary struCture on the inner surface of the antero-
derived from the suckers on the arms ami/or clubs dorsal part of the mantle.
of some squids (Fig. 1C). Nuchal folds: a series of longitudinal folds or pleats of
Ink sac: the organ that manufactures and stores the ink skin on the nuchal region, the dorso lateral area
of cephalopods; gene rally lying along the intestine around the posterior part of the head, normally
(sometimes imbedded in the digestive gland) and covered by the anterior mantle wall.
emptying via a duct into the rectum. Olfactory organ: a pit, bump, or a finger-like protu-
Iridophores: cells located in the skin, containing lam- berance on the posterolateral surface of each side
inar crystals readily reflecting, refracting and pola- of the head; of presumed olfactory function.
rizying incident light. Orbital pore: minute pore in the skin, anterior to the
Keel: a flattened, muscular extension along the aboral cornea of cuttlefishes (except Spirula) and myop-
surface of some arms or the tentacular club to ren- sid squids; remnant of the primary eyelids.
der them more hydrodynamic (Fig. le, E) (see Orbital sinus: an anteriorly directed indentation in the
also swimming membrane). eyelid of oegopsids (Fig. 1M).
Koelliker organs: minute bristle-like structures that Pen: see gladius.
cover the body of planktonic octopod larvae. Photophore (=light organ): an organ of greater or less-
Lanceola: the expanded portion of the gladius vane. er complexity that produces and distributes biolu-
Light organ: see photophore. minescence, either intrinsically through biochemi-
Ligula: the spatulate to spoon-shaped terminal struc- cal reaction or extrinsically through luminescent
ture of the hectocotylus of octopods, which bacteria (Fig. lE, K, L).
Cephalopoda 719
Protective membrane: thin fold of integument along the Suckers, primary: arm suckers in larval or juvenile
lateral angles of the oral surface of the arms and octopuses developed before hatching; either 3 or 4
clubs lateral to the suckers, usually supported by arranged in one row (Fig. 1Y).
muscular rods called trabeculae (Fig. 1C, O) (see Suckers, secondary: arm suckers injuvenile octopuses
trabeculae ). developed after hatching; arranged in one or two
Rachis: the thickened central axis that usually extends rows distal to, and smaller than, primary suckers
the entire length of the gladius. Free rachis is the (Fig. 1Z).
portion that does not support vanes (Fig. ID) (see Swimming membrane (=keel): an elongate muscular
gladius, vane). vane along the aboral surface of some arms that
Radula: the chitinous ribbon-like band in the mouth of helps streamline and support the arms during
cephalopods containing numerous transverse rows swimming (Fig. 1C, E).
of teeth (Fig. IP, Q). Absent in Spirllla and most Tail: the posterior extension generally of the gladius
cirrate octopods. and mantle epithelium, frequently elongate. Fins
Radula, ctenodont: a radula in which the rachidian may extend posteriorly along the tail (Fig. 1E),
(central) and lateral teeth are similar in appearance; and the tail may be swollen by the inclusion of
each has multiple cusps (more than three). The vacuolated tissue.
central cusp (mesocone) on the rachidian tooth Tentacles: elongate circumoral appendages of cuttle-
may be inconspicuous (Fig. IQ). fishes and squids used to capture prey; located
Radula, heterodont: a radula in which the teeth on between ventrolateral (3rd) and ventral (4th) arm
each side are different from one another in appear- pairs, divided into a proximal stalk and a distal
ance; the rachidian tooth has one or two cusps on club; clubs generally expanded with arrangement
either side of a distinct central cusp (mesocone), of suckers (or hooks), stalks in adults commonly
and each lateral tooth bears one or two cusps (Fig. devoid of suckers. In cuttlefishes tentacles can
IP). retract into pockets on the head, or (in squids)
Radula, homodont: a radula in which all teeth appear merely contract (Fig lE).
similar in size and shape; each rachidian and late- Tentacular club: terminal portion of a tentacle; armed
ral tooth has one simple cusp. with suckers (sometimes also hooks), used for
Rostrum: see spine. capturing prey (Fig. 1C, E, S).
Side pockets: small membranous folds of the integu- Tentacular pocket: an open depression in the anteroven-
ment that form pockets lateral to the foveola (Fig. tral surface ofthe head of cuttlefishes into which the
IF) (see foveola). ejectable tentacles are retracted when not in use.
Snout: see arm-crown pillar. Trabeculae: muscular rods that support the protective
Spermatophore: a tubular structure manufactured by membranes on the arms and clubs of cephalopods;
male cephalopods for packaging sperm; each holds occasionally membranes are reduced ami/or tra-
millions of spermatozoids and is transferred intact beculae are elongated, forming papilla-like out-
and attached to the female until fertilization begins growths that extend beyond the edge of the mem-
(Fig. IR). brane (Fig. 10).
Spine (=rostrum): the sharp spike-like extension on the Tragus: a small cartilaginous knob-like projection
posterior tip of the gladius or cuttlebone (Fig. 1D). from the inner wall of the funnel locking cartilage
Sucker ring: chitinous, often serrated or denticulate in some squids of the families Chiroteuthidae and
ring that encircles the opening of suckers of squids Mastigoteuthidae (see antitragus) (Fig. 2.42G).
and cuttlefishes (Fig. 1T). Vane: a thin lateral expansion ofthe gladius that ari-
Suckers: muscular suction-cup structures on the arms ses from the rachis (Fig. ID) (see rachis).
and tentacles (rarely on the buccal membrane) of Water pores: small orifices at the base of the web of
cephalopods; some are stalked, placed on muscular some pelagic octopuses, for example Tremoctopus
rods that contract (squids and cuttlefishes); some and Ocythoe (Fig. 1U, 2.75D).
are sessile, embedded (without stalks) on the oral Web: a thin muscular fold of skin extending between
surface of the arms (octopuses). They are usually the arms of many octopuses and a few squid (for
counted either in longitudinal or in transverse example, Histiotellthis), giving an umbrella-like
(oblique) rows (Fig. 1S). appearance when the arms are spread (Fig. 1U).
720 Cephalopoda
Outline c1assification Family Sepiidae Keferstein, 1866

Family Sepiolidae Leach, 1817
Classification of the class Cephalopoda at the supra- Subfamily Rossiinae AppellOf, 1898
generic level is as unstable as that ofmost other classes Subfamily Sepiolinae Leach, 1817
of invertebrates. For non-specialists in cephalopod mol- Subfamily Heteroteuthinae AppellOf, 1898
luscs it is reasonable to compromise between tradition- Family Sepiadariidae Naef, 1912
al systems as presented in university textbooks (mostly Family Idiosepiidae AppellOf, 1898
outdated), and modem improvements and amendments Otder Teuthida Naef, 1916
not yet adopted universally. 1 use the system of Suborder Myopsida d'Orbigny, 1845
Cephalopoda described in my previous books (Nesis, Family Loliginidae Steenstrup, 1861 (including
1982, 1985, 1987), which basically coincides with the Pickfordiateuthidae Voss, 1953; subfamilies not
one used in many other manuals and textbooks (Clarke, listed)
1996; Clarke and Trueman, 1988; Guerra Sierra, 1992; Suborder Oegopsida d'Orbigny, 1845
Mangold, 1989; Roper, 1983; Roper et al., 1984; Family Lycotheuthidae Pfeffer, 1908
Sweeney et al., 1992; and others). A critical (skeptical) Subfamily Lycoteuthinae Pfeffer, 1908
analysis of the improvements and new tendencies in Subfamily Lampadioteuthinae Berry, 1916
cephalopod systematics, primarily elevating the rank of Family Enoploteuthidae Pfeffer, 1900, s.l.
some taxa one level (subfamily to family, family to Subfamily Enoploteuthinae Pfeffer, 1900
suborder/order, etc.) was published by Nesis in 1996. Subfamily Ancistrocheirinae Pfeffer, 1912
Subfamily Pyroteuthinae Pfeffer, 1912
The class Cephalopoda includes two Recent sub- Family Octopoteuthidae Berry, 1912
classes: Nautiloidea (one order, family, and genus: Family Onychoteuthidae Gray, 1849
Nautilus, having no pelagic stage and absent from the Family Walvisteuthidae Nesis and Nikitina, 1986
Atlantic), and Coleoidea (all other modem cephalo- Family Gonatidae Hoyle, 1886
pods). The Subclass Coleoidea includes four orders: Family Chtenopterygidae Grimpe, 1922
Family Bathyteuthidae Pfeffer, 1900
Sepiida (cuttlefish), divided by some authors into two Family Histioteuthidae Verrill, 1881
or three orders, with two suborders: Spirulina (with Family Psychroteuthidae Thiele, 1921
only family, genus and species: Spirula spirula), Family Architeuthidae Pfeffer, 1900
and Sepiina (with 4 families); Family Neoteuthidae Naef, 1921
Teuthida (squids) also with two suborders: Myopsida Family Brachioteuthidae Pfeffer, 1908
(neritic squids, one or two families), and Oegopsida Family Ommastrephidae Steenstrup, 1857
(oceanic squids), 23 to 26 families, depending on Subfamily lIIicinae Pfeffer, 1912
the author; Subfamily Todaropsinae Nigmatullin, 1988, Ms
Vampyromorpha one family, genus and species: Subfamily Todarodinae Pfeffer, 1912
Vampyroteuthis infemalis; and Subfamily Ornithoteuthinae Nign1atullin, 1979
Octopoda (octopuses), with two suborders: Cirrata Subfamily Ommastrephinae Pfeffer, 1912
(with 3 families), and Incirrata (with 3 superfa- Family Thysanoteuthidae Keferstein. 1866
milies and 8 families). Family Pholidoteuthidae Adam, 1950
Family Lepidoteuthidae Naef, 1912
The following outline surnmarizes the classification Family Batoteuthidae Young and Roper, 1968
system adopted herein. Families and subfamilies Family Cycloteuthidae Naef, 1923
treated in this review are underlined. Family Chiroteuthidae Gray, 1849 (including
Grimalditeuthidae Pfeffer, 1900)
Class Cephalopoda Cuvier, 1797 Family Mastigoteuthidae Verrill, 1881
Subclass Coleoidea Bather, 1888 Family Joubiniteuthidae Naef, 1922
Order Sepiida Naef, 1916 Family Promachoteuthidae Naef, 1912
Suborder Spirulina Stolley, 1919 Family Cranchiidae Prosch, 1849
Family Spirulidae Owen, 1836 Subfamily Cranchiinae Pfeffer, 1912
Suborder Sepiina Naef, 1916 Subfamily Taoniinae Pfeffer, 1912
Cephalopoda 721
Order Vampyromorpha Pickford, 1939 South Atlantic, such as the Psychroteuthidae,

Family Vampyroteuthidae Thiele, 1915 Lepidoteuthidae and Walvisteuthidae, are omitted
Order Octopoda Leach, 1818 from this review because their larvae and juveniles
Suborder Cirrata Grimpe, 1916 rema in to be discovered.
Family Cirroteuthidae Keferstein, 1866
Family Stauroteuthidae Grimpe, 1916 As mentioned above, identification of cephalopod lar-
Family Opisthoteuthidae Verrill, 1896 vae and juveniles is usually feasible on1y to the generic
Suborder Incirrata Grimpe, 1916 level beca use most key diagnostic characters appear in
Superfamily Bolitaenoidea Chun, 1911 the subadult-adu1t stage. Identification of the young is
Family Bolitaenidae Chun, 1911 further hampered by the fact that for many species
Family Amphitretidae Hoyle, 1886 (including these stages are still undescribed. In view of this, diag-
Idioctopodidae Taki, 1962) noses in this review are provided usually for on1y one
Superfamily Octopodoidea d'Orbigny, 1845 species of each genus known to occur in the South
Family Octopodidae d'Orbigny, 1845 Atlantic, even when more than one species has been
Subfamily Octopodinae d'Orbigny, 1845 recorded in this basin. The remaining species of the
Subfamily E1edoninae Grimpe, 1921 genus are only listed and, occasionally, commented
Subfamily Bathypo1ypodinae Robson, 1932 upon briefiy, but their known distribution is detai1ed in
Subfamily Graneledoninae Voss, 1988 Table l. On the other hand, more than one species per
Family Vitreledonellidae Robson, 1932 genus is included when their 1arvae ancl/or juveniles
Superfamily Argonautoidea Naef, 1912 have been adequately diagnosed and are morpho10gi-
Family Alloposidae Verrill, 1882 cally different enough to allow positive recognition.
Family Ocythoidae Gray, 1849
Family Tremoctopodidae Brock, 1882 The diagnoses of the species are based primarily on
Family Argonautidae Naef, 1912 subadult and adu1t stages, if these stages inhabit the
pelagic realm. Larval and juvenile characters, when
available, are presented as well. On the other hand, for
Identification those cephalopods whose adults are benthic only, the
planktonic larvae aml/or juveniles are diagnosed.
In the South Atlantic there are representatives of 3 out
of the 5 families of Sepiida, as well species of the Key to young stages of coleoid cephalopods up to
families ofTeuthida, Vampyromorpha and Octopoda. the family level (modified from Sweeney et al., 1992)
Species of all families of co1eoid cepha10pods may be
caught in the plankton, at least in their larval stages, Fins absent: order Octopoda, suborder Incirrata
except Cirrata, although some of these peculiar ani- ............................................................................ 33
mals may be caught as juveniles or adults in deep 1a Fins present 2
tows with large trawls (not considered in this review). 2 Fins attached to the mantle laterally 3
Representatives of three strictly benthic subfamilies 2a Fins attached to the mantle terminal1y or dorsal1y
are lacking in the plankton: Rossiinae (Sepiida, .............................................................................. 4
Sepiina, Sepiolidae), Bathypolypodinae and 3 Eight subequal arms joined by a deep web; in
Grane1edoninae (Octopoda, Incirrata, Octopodidae). late juveniles cirri developed on arms from the
sides of suckers: order Octopoda, suborder
The commonest planktonic cephalopods are oceanic Cirrata. Very rare in the plankton, not included
squids (Oegopsida), primarily of the families in the present review
Enoploteuthidae, Onychoteuthidae, Histioteuthidae, 3a Eight (in earliest 1arvae sometimes 4 or 6) arms
Ommastrephidae, and Cranchiidae. Representatives of and 2 tentacles between ventro1ateral (3rd) and
the planktonic octopod family Bolitaenidae are rather ventral (4th) arm pairs; no cirri; suckers stalked
conunon in deep tows (meso- and bathypelagic). . 5
4 Fins near dorsal midline of posterior mantle; 2nd
Planktonic stages are known for most but not all ge- pair of fins developed on the mantle anterior to
nera and species. Some families known to occur in the first pair at mantle length (ML) approx. 15 mm,
722 Cephalopoda
later first pair reduced; suckers not stalked; in lOa Eye not entirely covered by transparent me m-
early larvae arms only with cirri, without suck- brane (cornea), eye lens in contact with water in
ers, in older larvae suckers begin far from the living animal when eyelid not contracted ... 11
mouth; retractile filaments without suckers and ll Fins in juveniles and adults similar to those of
cirri present between dorsal (1st) and dorsolate- fish, with muscular rays; tentacles in larvae short,
ral (2nd) arms: order Vampyromorpha, with spatulate round clubs; aboral side of club
Valllpyrotellthis illjemalis (Fig. 2.60) with one chromatophore: family Chtenoptery-
4a Fins insert on mantle dorsolaterally or laterally; gidae, Clltellopteryx siCllla (Fig. 2.22)
suckers stalked; neither cirri nor filaments; only Ila Fins without muscular rays 12
one pair of fins, if two (Chiroteuthidae), the se- 12 Larvae with long neck 13
cond located not on the mantle but on continua- 12a Larvae without long neck 14
tion of gladius 6 13 Larval neck with dorsal hump behind head, with
5 Fins long and narrow, not paddle-shaped, extend single subdivision chamber; no conspicuously
laterally from near anterior to posterior end of long tail; arm-crown pillar absent: family
mantle, not fused posteriorly; long wide calcare- Brachioteuthidae, Brachiotellthis (Fig. 2.29)
ous internal shell (sepion) under the skin on dor- 13a Larval neck without hump, subdivided into many
sal mantle side: family Sepiidae. Absent in the irregular chambers; tail very long, sometimes
plankton, not included in the present review. In with secondary fin behind primary one (com-
the South Atlantic present only off Africa monly broken off); base of arm-crown elongated
5a Fins paddle- or ear-shaped 7 and forming an arm-crown pillar: family
6 Chambered calcareous shell in the form of ram's Chiroteuthidae (including Grimalditeuthidae)
horn under the skin in posterior mantle (2-3 cham- (Fig. 2.42, 2.43)
bers in newborn larvae); fins tiny, petaloid or kid- 14 One round (inconspicuous) light organ at the
ney-shaped, on posterior mantle, widely separated base of each arm of 1st-3rd pairs; tentacles long
and perpendicular to body axis; beaks in larvae and thin; fins small, kidney-shaped; body color
large, protruding beyond short arms; only 2 arm uniformly deep-brown: family Bathyteuthidae,
pairs in newborn larvae: order Sepiida, suborder Bathytellthis abyssicola (Fig. 2.23)
Spirulina, family Spirulidae, Spirula spirula 14a No light organs at arm bases; complex of other
(Fig.2.1) features different.. 15
6a Coiled chambered shell absent; beaks not pro- 15 Funnel locking cartilage in the form of a tilted
truding beyond arms 8 "T"; larvae with barrel-like mantle densely co-
7 Fin length (along antero-posterior axis) exceeds vered by large purple chromatophores; juveniles
its width: family Sepiadariidae. Indo- West with wide protective membranes on 2nd-3rd
Pacific, absent in the South Atlantic arms: family Thysanoteuthidae, Thysallotellthis
7a Fin width equal to or larger than its length: fa- rholllblls (Fig. 2.37)
mily Sepiolidae (Fig. 2.2, 2.3) 15a Funnellocking cartilage straight, round or subtri-
8 Tentacles not developed at hatching, arms com- angular; chromatophores less dense, not purple ..
plete (4 pairs): family Idiosepiidae. Indo-West .......................................................................... 16
Pacific, absent in the South Atlantic 16 Mantle fused with funnel, no funnellocking car-
8a Tentacles developed at hatching, not shorter than tilage 17
longest arms 9 16a Mantle not fused with funnel, funnel locking
9 Tentacles in larvae fused into a "trunk" (probos- cartilage developed 18
cis), divided in juveniles; funnellocking cartilage 17 Mantle free in nuchal region: family
in the form of an inverted "T": family Chiroteuthidae, Grilllalditellthis bOllplandi
Ommastrephidae (Fig. 2.30-36) (only postlarval stages - juveniles and adults)
9a Tentacles never fused; funnel locking cartilage (Fig. 2.43)
not in the form of an inverted "T" 10 17a Mantle fused with head in nuchal region: family
10 Eye entirely covered by transparent membrane Cranchiidae (Fig. 2.47-59) 17b
(cornea): suborder Myopsida, family Loliginidae 17b Two (longitudinal) or four (diverging in the
(inc1uding Pickfordiateuthidae) (Fig. 2.4, 2.5) form of a V) hyaline stripes with cartilaginous
Cephalopoda 723
tubercles on ventral mantle beginning at the sites 25 Tentacles in larvae long but with very short club
of mantle-funnel fusion; eye photophores (less than 12 suckers), rudimentary in juveniles,
numerous, round, in 1-2 arches; funnel valve absent in adults: family Octopoteuthidae (Fig.
present (except Leachia): subfamily 2.15,2.16)
Cranchiinae (Fig. 2.47-49) 25a Tentacles well developed, with more than 12
17c No hyaline stripes with cartilaginous tubercles suckers: family Cycloteuthidae (Fig. 2.40, 2.41)
on ventral mantle, except in the area of mantle- 26 Left eye usually much larger than right; skin
funnel (or also mantle-head) fusion in some ge- photopores present around eyelid: family
nera; one large crescent-shaped and one (rarely Histioteuthidae, Histiotellthis (Fig. 2.24, 2.25)
2) small, usually rod-shaped eye photophores; 26a Eyes symmetrical; no photophores around eyelid
funnel valve absent (except in Megalocranchia ......................................................................... 27
and Egea): subfamily Taoniinae (Fig. 2.50-59) 27 Photophores present on ventral side of eyeball
18 Mantle fused with head in nuchal region; mantle andJor on mantle or viscera 28
thick; fins large, broad, terminal, posteriorly 27a Photophores absent 29
fused; no photophores: family 28 Arms and tentacles without hooks: family
Promachoteuthidae, Promachoteuthis (Fig. Lycoteuthidae (Fig. 2.6, 2.7)
2.46) 28a Arms (in all genera) and tentacles (except
18a Mantle not fused with head, not thickened in lar- Pterygioteuthis) with hooks and suckers (in lar-
vae and juveniles; fins different; photophores vae with suckers only): family Enoploteuthidae
usually present in juveniles and adults 19 s.l. (Fig. 2.8-14) 28b
19 4th am1S absent or rudimentary in early larvae 28b Photophores present on ventral side of eyeball
......................................................................... 20 ....................................................................... 2ge
19a 4th arms developed even in early larvae 21 28c Photophores on eyeball absent: subfamily
20 Tentacular club suckers in 4 rows, arm suckers in Ancistrocheirinae, Ancistrocheirus /esllellrii
2 rows: family Onychoteuthidae (Fig. 2.17-20) (Fig.2.12)
20a Tentacular club suckers in more than 4 rows, 28d Photophores present on viscera, absent on man-
arms in juveniles and adults with 2 rows ofsuck- tle surface: subfamily Pyroteuthinae (Fig. 2.13,
ers and 2 rows of hooks: family Gonatidae, 2.14)
Gonatlls alltarcticus (Fig. 2.21) 28e Photophores present on mantle surface, no photo-
21 Body elongate with long pointed tail; tentacles phores on viscera: subfamily Enoploteuthinae
very long and thin 22 (Fig. 2.8-11)
21 a Body without long pointed tail; tentacles not 29 Tentacular club with more than 4 rows of suck-
concpicuously elongated 24 ers, at least in the carpal part; no papillae or
22 Arms very long, thin, flexible, with minute suck- "scales" on mantle 30
ers in 8-12 rows; funnel locking cartilage oval, 29a Tentacular club with 4 rows of suckers; minute
without tragus and antitragus: family papillae or "scales" on mantle 32
Joubiniteuthidae, JOllbillitellthis portieri (Fig. 30 Numerous minute suckers or buds form a cluster
2.45) in the carpal part of the tentacular club; club
22a Arms not very long, suckers in 2 rows; funnel suckers of both central rows equal in size; fins
locking cartilage either straight or oval with at attached dorsolaterally, diverging anteriorly
least tragus 23 ......................................................................... 31
23 Funnel locking cartilage simple, straight: family 30a No' numerous minute suckers on carpus; club
Batoteuthidae, Batoteuthis skolops (Fig. 2.39) suckers of both central rows very unequal in
23a Funnellocking cartilage oval, with tragus, usual- size; fins attached dorsally, not divergent: fami-
Iy also antitragus: family Mastigoteuthidae Iy Psychroteuthidae, Psychrotellthis g/acia/is
(Fig.2.44) Thiele. Present in the South Atlantic but not
24 Eyes in larvae stalked, tubular, directed antero- included in the present review because early
laterally 25 developmental stages are not described
24a Eyes in larvae not stalked and not anterolateral- 31 Carpal group of suckers and buds very short in
Iy directed .26 comparison with long central part bearing 4 rows
724 Cephalopoda
of suckers: family Architeuthidae, Architeuthis 37 Specialized funnellocking cartilage absent; males

(Fig.2.26) not dwarfed; hectocotylized arm (3rd left or right)
31 a Carpal group of suckers and buds longer than or in males modified only at the end, hectocotylus
comparable in length to rather short central part developed only during sexual maturation: family
bearing 4 rows of suckers: family Neoteuthidae Octopodidae (Fig. 2.64-69) .37b
(Fig. 2.27, 2.28) 37a Specialized funnel locking cartilage present;
32 Tentacles long, club suckers numerous, round in males dwarfed; hectocotylized arm (3rd left or
larvae but laterally compressed in juveniles and right) in males modified totally and developed
adults; ski n papillae with cartilaginous tubercles into a sac, thus giving the impression that irnma-
(at least in juveniles), neither adjoining nor ture (even newborn) males have 7 arms: super-
overlapping; fins rhomboidal or heart-shaped, family Argonautoidea 38
reaching mantle end; tail needle short or absent: 37b Arm suckers first in one row (3 or 4), then in two
family Pholidoteuthidae, Pholidoteuthis (Fig. rows: subfamily Octopodinae (Fig. 2.64-68)
2.38) 37c Arm suckers in one row throughout: subfamily
32a Tentacles present only in larvae, clubs short, Eledoninae, Eledone nigra (Fig. 2.69)
with 6 sucker-like structures; juveniles and 38 Dorsal and ventral arms much longer than the
adults without tentacles; skin papillae leathery, rest; newly born not enclosed in gelatinous coat-
without cartilaginous teeth, tightly adjoining or ing; ventral mantle side in juvenile and adult
overlapping each other (as fish scales); fins females reticulate with interconnecting tubercles
round or suboval, not reaching mantle end, man- and ridges: family Ocythoidae, Ocythoe tuber-
tle tapering into long tail (acute needle in juve- culata (Fig. 2.72)
niles): family Lepidoteuthidae, Lepidoteuthis 38a Arm length relationships different; hatchlings
grimaldii Joubin. Present in the South Atlantic partly or entirely enclosed in gelatinous coating;
but not included in this review because larvae ventral mantle side in females not reticulated
are poorly known ......................................................................... 39
33 Body gelatinous, transparent (at least in larvae 39 Dorsal arms in larvae greatly elongated, robust;
and juveniles), often covered with jelly-like in juveniles dorso-lateral arms are somewhat
coating 34 shorter or longer than dorsal; bases of both dor-
33a Body firm, muscular, opaque (except sal and dorsal and dorso lateral arms joined by
Tremoctopus gelatus, Fig. 2.74) 37 web; no flag-like flaps on ends of dorsal arms in
34 Eyes large, spindle-shaped in side view; suckers juvenile and adult females; ventro-Iateral arms
in one row within web and in two rows outside very short in larvae; only head and arm bases
web; adults not transparent: family Alloposidae, enveloped by a cuff-shaped membrane in larvae;
Alloposus mollis (Fig. 2.71) no secondary shell; right ventro-lateral arm
34a Eyes not spindle-shaped; suckers in one row modified i.n males: family Tremoctopodidae,
throughout; all developmental stages transparent Tremoctopus (Fig. 2.73, 2.74)
or semi-transparent 35 39a Dorsal arms slightly elongated in postlarval stages,
35 Eyes tubular, located on the head top and directed equal to tbe otbers in larvae; arm bases not joined
anterodorsally: family Amphitretidae (including by web; large flag-like flaps on ends of dorsal anns
Idioctopodidae), Amphitretus pelagicus (Fig. in juvenile and adult females; ventro-Iateral arms
2.63) not short; early larvae completely enclosed in
35a Eyes rectangular or elliptical, located on the gelatinous coating; juvenile females build a secon-
head sides and directed laterally 36 dary calcareous shell, Iive and (later) incubate eggs
36 Eyes rectangular; digestive gland very long and in it; left ventro-lateral arm modified in males:
slender, posteriorly pointed, its length exceeds family Argonautidae, Argonauta (Fig. 2.75)
mantle length: family Vitreledonellidae,
Vitreledonella richardi (Fig. 2.70) The family Walvisteuthidae Nesis and Nikitina, 1986
36a Eyes elliptical; digestive gland round to oval, its (recorded in the South Atlantic) is not included in the
length less than half the mantle length: family key because only two mature males are known.
Bolitaenidae (Fig. 2.61, 2.62) Morphologically it is close to the Onychoteuthidae.
Cephalopoda 725
Diagnoses of the taxa direct. Some species may be in the plankton as juve-
niles or adults in the period of mating. Six genera in
Most of the diagnoses below are complemented with the South Atlantic, 3 in the plankton.
brief distributional data (see also Table 1), general bio- ROlldeletiola minor (Naef, 1912) (Fig. 2.2). Body
logical remarks, and relationships with related species. short, rounded. Mantle fused with head dorsally,
Mantle length (see Glossary) is abbreviated as ML. its anterior end on the ventral side not produced
anteriorly and not covering funne!. Fins kidney-
Order Sepiida like, short, much shorter than the ML. 1st and 2nd
Suborder Spirulina arm pairs not connected by membrane. Arm suck-
Family Spirulidae ers not modified. Tentacular suckers in 16 rowS.
One monospecific genus in the World Ocean. 3rd arm pair in males usually bent inwards toward
Spirllla spirllla (Linné, 1758) (Fig. 2.1). An inter- mouth. A large photophore, bilobed in juveniles,
nal calcareous shell in the form of a decoiled (as in compact in adults, with two lenses, in anterior part
a ram's hom) spiral, functioning as a fioat, on pos- of the ink sac, deeply imbedded into its tissues; in
terior end ofbody in sagittal plane under thin skin, females connected with accessory nidamental
consisting of isolated chambers connected by a glands. A very small species, ML up to 25 mm,
hollow tube (siphuncle). Number of chambers usually less than 20 mm.
from 3 (in smallest known juvenile) to 40 (in Dwells chiefiy on the bottom in the lower sublitto-
adults). Mantle thick, not fused with head. Eyes ral and upper bathyal, but reproductive adults
large, protruding, covered by transparent skin. commonly rise to midwater and may be caught
Fins small, kidney-shaped, near very end of body near the surface.
oblique to longitudinal body axis. Large round Heterotellthis (Heterotellthis) dispar (Rüppell,
photophore between fins at end of body. Bases of 1845) (Fig. 2.3). Body short, egg-like. Mantle mus-
all arms joined by membrane. Arms with 2 rows of cular, not fused with head. Anterior mantle margin
suckers. Tentacles retractable with thin stem and in larvae and early juveniles straight, in later juve-
small widened club. Club suckers in 16 rows. niles and adults strongly protruding forward on
Color creamy-white with maroon-red. Life posi- ventral side, reaching level of anterior eye margin
tion vertical, head down. Development almost and almost covering funnel from below. Fins in
direct. Egg size 1.5-1.7 mm, ML of smallest juve- larvae andjuveniles tongue-like, wide (fin width in
nile 1.5 mm; ML of adults up to 5 cm. newboms exceeding mantle width), located in the
Occurs mainly over continental and insular slopes middle of mantle; in adults kidney-shaped and
and in the open ocean not far from the slope. Range shifted to posterior half of mantle such that anteri-
disjunct and associated with closed circulations of or fin margin is near midline ofventral mantle side,
intermediate water masses. In the South Atlantic and posterior margin reaches or exceeds end of
found only off Namibia and Westem Cape, pro- body. Arms in larvae and juveniles very short, in
bably expatriated from the SW Indian Ocean, where adults moderately long, arm suckers in 2 rowS. lst-
a known population exists. Spawns on the bottom at 3rd arm pairs connected at bases with deep mem-
bathyal depths. Early juveniles ascend to midwater. brane. Arm structure differing greatly in adult
Late juveniles and immature subadults migrate from males and females. In males right 1st and 2nd arms
approx. 100-300 m at night to 50Ó-IOOOm at day. connected from inside by muscular band up to half
their lengths; suckers extend up to arm tips; on 3rd
Suborder Sepiina arms 'there are 5 strongly enlarged suckers, includ-
Family Sepiolidae ing 2 (in central part of arm) very large ones, at
Mantle short, broad, rounded posteriorly. Gladius not right angle to one another. In females arms not con-
calcified, chitinous, or lacking. Fins round or kidney- nected with membrane, without enlarged suckers,
shaped, not joined posteriorly. Funnel cartilage sim- ends of 1st and 2nd arms without suckers. Longest
ple, straight. Eye covered by transparent skin. Three arms in males are 3-4th, in females 2-3rd.
subfamilies, 14 genera. Mostly benthic or nerito- Tentacles retractable, very long, whip-like, with
oceanic animals. Eggs laid on bottom, development unwidened short club bearing 8 rows of micro-
726 Cephalopoda
scopically small suckers. Gladius absent. Large Dwells on the inner shelf, shallow bays and
oval bilobed photophore on ventral side of ink sac, estuaries; a very euryhaline and euryoxic species.
noticeable even in newborns. Luminous mucus Larvae distinguishable from those of Loligo and
with luminescent bacteria may be ejected through related genera by pattern of red chromatophores
funnel by muscular contraction so that animal may on dorsal side (none at ML 2.0-2.5 mm, 6 on head
"shoot with fire". Color very bright, vivid, with and 4 on mantle at ML 3.5-5.5. mm, 9-15 at ML
white fin bases and metallic iridescence on head 6.5-7.5 mm), and by 3 dark chromatophores
and body. Egg size in ovary 1.5-2.0 by 1.3-1.8 rnm, ventrally on head along sides of funnel. Lolligun-
before hatching 2.5 rnm. ML in adult females up to eula mereatoris Adam, 1941, included by some
25 rnm. Adult female originally described as H. authors in a separate genus Afrololigo, is distribu-
atlantis G. Voss, 1955. ted in nearshore waters but not in bays and estua-
Lives predominantly over the slopes, spawning on ries. It has been reported to have an unpaired rec-
the bottom at bathyal depths. Juveniles and sub- tal photophore in adult females only. The larvae
adults perform ample diurnal migrations. are distinguished from those of L. brevis only by the
Populations of the Western and Eastern Atlantic precocious development of dorsal mantle chromat-
are probably isolated. ophores (Vecchione, 1982; Vecchione ahd Lipinski,
1995).
Order Teuthida • Loligo vulgaris Lamarck, 1798 (Fig. 2.5). Mantle
Suborder Myopsida cylindrical in the anterior half, conical in the pos-
Family Loliginidae (including Pickfordiateuthidae) terior half, its width in adults 3-6 times less than
Mantle elongated, its anterior margin with mid-dorsal ML. No tail. Mid-dorsal mantle projection tongue-
projection. Eye covered by transparent corneal mem- like, noticeable even in larvae. Fins transversely
brane with only minute pore at anterior end. Tentacles tongue-like in larvae, oval in juveniles, elongate
not retractable. Arms with 2, tentacles with 4 rows of diamond-shaped in adults, fin length equal to or
suckers, without hooks. Minute suckers may be longer than width. Arm length in adults >25%
present on buccal membrane. Buccal membrane with ML. Central club suckers much larger than margi-
8 lappets, connectives to 4th arms attached ventrally. nal ones. Buccal membrane with small suckers.
Funnel cartilage straight, elongated. Gladius feather- Lateral sides of gladius bent arch-like, without
like, with short free rachis. Two (rarely one) photo- marginal ribs. No photophore .. Egg size: length
phores present on ink sac in some species. Three to 2.3-2.7, width 1.8-2.2 mm. Newborn larvae (ML
five subfamilies, 8-15 genera. Neritic nektobenthic approx. 2.2 mm) with 9 red chromatophores dor-
animals with bottom egg-masses, eggs surrounded by sally on mantle, 4 on dorsal side of head and 2
capsules of thick tough jelly. Larvae not very differ- patches ("cheek patches") of 4 chromatophores
ent from adults. Larvae of different species very sim- each on ventral side ofhead. ML in adults up to 50
ilar, distinguished mostly by number and disposition cm. Two subspecies; only L. v. reynaudii
of chromatophores (which may fade after fixation). In d'Orbigny, 1845 recorded from the SE Atlantic.
the South Atlantic 6-7 genera. Three related species have be en mentioned for the
• Lolliguncula brevis (Blainville, 1823) (Fig. 2.4). SW Atlantic: Loligo (or Amerigo according to
Mantle short, wide spindle-like, rounded posteri- some authors) sanpaulensis Brakoniecki, 1984
orly, without tapering tail. Mantle length 2-3 times (=L. brasiliensis sensu Castellanos and Menni,
mantle width. Mid-dorsal mantle projection short, 1969, L. emmalána Gray, 1849); L. (or Amerigo)
blunt. Fins egg-like in larvae, kidney-like in juve- patagoniea Smith, 1881 (=L. gahi Brakoniecki,
niles, oval in adults, length approximately half of 1984, partim, ?non d'Orbigny, 1835), the deepest,
ML, width 1.5 times length. Arm length in adults approx. 80-800 m, and coldest-living 10liginid;
approx. half of ML. Arm suckers toothed only on and Doryteuthis plei (Blainville, 1823). Larvae of
distal half. Central club suckers noticeably larger D. plei (McConathy et al., 1980), L. sanpaulensis
than marginal ones. Buccal membrane with small (Leta, 1987a), and L. patagoniea (Rodhouse et al.,
suckers. Lateral sides of gladius bent arch-like. No 1992) are hardly distinguishable from those of
photophore. Egg size approx. 2 mm. ML up to 10- Loligo (Sweeney et al., 1992; Vecchione and
12 cm. Lipinski, 1995).
Cephalopoda 727
Order Teuthida bottom layer on the slope. Adult males much rarer
Suborder Oegopsida than females.
Family Lycoteuthidae • Selenotelltlris scintillans G. Voss, 1958 (Fig. 2.7).
Mantle wide, robust, cylindrical in anterior, conical in Fin in larvae tongue-like or oval, in adults broadly
posterior half. Arms with 2 tentacles with 4 rows of diamond-shaped, not reaching midpoint ofmantle.
suckers, hooks absent. Buccal membrane with 8 lap- Mantle not tapering into tail. AII arms approxi-
pets, connectives to 4th arms attached dorsally. mately equal in length. Photophores: 5 in one row
Funnel cartilage straight, elongated. Gladius narrow, on ventral side of eyeball, central one largest; one
with two widenings, in the middle and near posterior large, globular on ends of 2nd and 3rd arms (only
end. Large diverse photophores on ventral side of eye- in adult males); 2 on tentacular stem (at base of
ball, inside of mantle cavity, sometimes on arms, ten- tentacle and at base of club); 10 in mantle cavity
tacles and body surface. 2 subfamilies, 4 genera, 5 on internal organs, 2 near rectum, 5 at level of
species. Mostly nerito-oceanic squids. Two species in branchial hearts (3 large along midline, 2 smaller
the South Atlantic. at sides); 3 closely spaced, but not merged, in cau-
Lycotelltlris diadema (Chun, 1900) dal part of mantle cavity; one large, globular on
[=?Onychoteuthis (?) longimanus Steenstrup, 1857; caudal end of mantle (developed at ML >5 rnm).
?Onychoteuthis (?) lorigera Steenstrup, 1875] In early larvae only central eye photophore is
(Fig. 2.6). Fin· in larvae oval or triangular, with developed, then three, also a photophore at base of
straight posterior margin; in adults diamond- or club; later 4 photophores in mantle cavity, but all
broadly heart-shaped, reaching midpoint ofmantle visceral photophores present only at ML approx. 9
anteriorly, posteriorly not tapering into a tail. AII mm. Small species, ML up to 4.5 cm.
arms in larvae and juveniles approximately equal, A nerito-oceanic (?) species, distributed in the
short, in subadult and adult males (ML >80-90 equatorial and western part of central water ma-
rnm) ends of 2nd and 3rd arm pairs very elongated sses, or bi-centrally. Ascends at night to epipelag-
(2nd arms may be 2-3 times as long as the mantle, ic layers.
3rd much shorter), tapering, whip-like and devoid
of suckers. In females the ends of 2nd-3rd arms are Family Enoploteuthidae s.1.
filiform but not lengthened. Photophores: 5 in one Mantle conical (or cylindrical in anterior and conical
row on ventral side of eyeball, central one largest in posterior half). Arms with 2, tentacles with 4 rows
even in early larvae; 2 on tentacular stem (at base of suckers, many of them transformed into hooks in
and at middle); 10 in mantle cavity on internal adulthood (usually 2 rows on arms, 1 or 2 on tentacu-
organs: 2 near rectum, 5 at level ofbranchial hearts lar club). Buccal membrane with 8 lappets, connec-
(3 large along midline, 2 smaller at sides); 3 tives to 4th arms attached dorsally. Funnel cartilage
merged together in caudal part ofmantle cavity. In straight, elongated. Gladius with one widening in pos-
larvae only 6 noticeable: 2 round near rectum, 2 terior half. Large, diverse, and very numerous (some-
oval near midline, and 2 round at sides at level of times many hundreds) photophores on ventral side of
branchial hearts (latter lacking in earliest larvae). eyeball, on the ventral mantle or inside of mantle ca-
On 2nd and 3rd arms in adult males, one (on 2nd) vity, sometimes on arms and tentacles. Three subfam-
or two (on 3rd) long rows of small photophores ilies, considered as separate families by many authors;
along base of lateral keel, through whole length of 7-8 genera. Oceanic and nerito-oceanic pelagic squids.
2nd and in distal half of 3rd ann (lacking in juve- Six genera and 13 species in the South Atlantic.
niles). No large globular photophores at arm ends,
nor caudal photophore. ML in males up to 19 cm, Subfamily Enoploteuthinae
females smaller, up to 11 cm. Larvae were Enoploteutlris (paraenoploteutlris) anapsis Roper,
described as Asthenoteution planctonicum Pfeffer, 1964 (Fig. 2.8). Mantle wide in larvae and juve-
1912. "Onychoteuthis lorigera" was described niles, later narrow and with soft tail in adults. Fin
from an adult male, Lycoteuthis diadema from an tongue-like in larvae, oval in juveniles, in adults
irnmature female. transversely diamond-shaped, not reaching tip of
A nerito-oceanic species, very abundant off mantle. Arms long, with 2 rows of hooks in adults.
Namibia and over Walvis Ridge. Adults in near- Tentacles strong, robust, much longer than anns,
728 Cephalopoda
club in larvae and juveniles with 4 rows of suckers (Asterotellthis) veranyi (Rüppell, 1844), was
(in juveniles central ones larger than marginal), in !
described as Enoploion ellstietum Pfeffer, 912. A.
adults with 2 rows of very unequal hooks (5-7 in a (Pygmabralia) grimpei G. Voss, 1958, was found
row), without marginal suckers. Photophores in 6 in the Gulf Stream off Florida. A. (Heterabralia)
straight longitudinal rows anteroventrally on man- siedleekyi Lipiñski, 1983, was found over the
tle; 8Y2 longitudinal rows on ventral side of head Schmitt-Ott Seamount in the SE Atlantic.
(central row only partial, in anterior half, 1st and • Abraliopsis (Abraliopsis) pfefferi (Joubin, 1896)
2nd lateral rows joined at the bases of ventral arms (Fig. 2.10). Mantle conical, without tail. Fin
and anterior to the margin of funnel pit); 8-10 close- tongue-like or transverse-oval in larvae and juve-
ly spaced photophores on ventral side of ,eyeball, niles, heart-shaped in adults, wide and long, reach-
marginal rather large, others approx. equal; 3 longi- ing tip of mantle. Arms rather long, with 2 rows of
tudinal rows on ventral arms; one incomplete row suckers in larvae; 2 rows of hooks and 2 rows of
on 3rd arms, reaching only midlength. Early larvae small suckers on tips in adults. Ventral arms
not easily distinguishable from Abralia, later ones noticeably longer than others. Tentacles long,
identified by the pattem of photophores and long robust, club very faintly widened, with elongate-
robust tentacles without marginal suckers. Egg size oval carpal membrane and longitudinal keel; in
supposedly about 1.0 rnm. ML up to 8 cm. larvae and juveniles its central part with 4 rows of
Another Atlantic species, E. (Enoploteuthis) leptll- suckers, in adults with 2 rows of hooks (all 4-5
ra (Leach, 1817), is distinguished from E. anapsis ventral hooks large, crescent-shaped; dorsal hooks
by its thin, rather short tentacles and the photo- very small), and one row of very small suckers
phore pattem (7 longitudinal rows on the antero- between small hooks. In juveniles ventral hooks
ventral mantle and 10 on the ventral side of the appear earlier than dorsal. Buccal membrane deep
head, 1st and 2nd not joined). purple. Photophores on ventral mantle in approx. 6
• Abralia (Pygmabralia) redfieldi G. Voss, 1955 longitudinal rows in juveniles, diffuse in adults
(Fig. 2.9). Mantle rather narrow, conical, without except bare mid-ventral strip; on ventral head also
tail. Fin tongue-like or oval in larvae and juve- diffuse in central part and the 2 rows between eyes
niles, heart-shaped in adults, reaching tip of man- and funnel opening; in 3-4 longitudinal rows on
tle. Arms of moderate length, with 2 rows of suck- base (2 in juveniles) and 2 on tips of ventral arms.
ers in larvae; 2 rows of hooks and 2 rows of small On eyeball, ventrally, 5 round photophores, 1st
suckers on tips in adults. Tentacles long, robust, and 5th somewhat larger than others. Three round
club in larvae and juveniles with 4 rows of suck- black photophores on ends of ventral arms, in lar-
ers, in adults with 3 hooks in ventralmost row and vae first 1 (at ML approx. 3 rnm), then 3, but
2 rows of suckers in its central parto Buccal mem- appearjng as white balls, becoming black only
brane pink. Photophores on ventral mantle in later. Egg size 1.0-1.5 mm, average 1.0 by 1.3
approx. 10 transverse rows in juveniles, diffuse in mm. ML up to 40 mm.
adults; in 9-10 longitudinal rows on ventral sur- Closely related to a complex ofvery similar Indo-
face of head; in 4 longitudinal rows on base and 3 West Pacific species: A. hoylei (Pfeffer, 1884); A.
on tips of ventral arms. On eyeball, ventrally, 5 paeifieus Tsuchiya and Okutani, 1990; A. tui
photophores, all round, 1st, 3rd and 5th larger than Riddell, 1985.
2nd and 4th. In larvae one (central) appears first, • Abraliopsis (Micrabralia) gilchristi (Robson,
then 3, then all 5. Larvae distinguished from those 1924) (Fig. 2.11). Mantle conical, without tail. Fin
of Abraliopsis by short ventral arms without large tongue-like or transverse-oval in larvae and juve-
terminal photophores; ventral mantle photophores niles, heart-shaped in adults, long but not wide,
in transverse rows, and thick tentacular stems are reaching tip of mantle. Arms rather long, with 2
also characteristic. Egg size supposedly about 0.8 rows of suckers in larvae; 2 rows of hooks and 2
rnm. ML up to 35 rnm. rows of small suckers on tips in adults. Ventral
A nerito-oceanic species, distributed mostly over arms noticeably longer than other. Tentacles long,
the slope and not far from islands, but also in the robust, club almost not widened, with elongate-
open ocean in equatorial waters. The larva of a oval carpal membrane and longitudinal keel; in
related nerito-oceanic Atlantic species, A. larvae and juveniles its central part with 4 rows of
Cephalopoda 729
suckers, in adults with 2 rows of hooks (4 ventral of small photophores on ventral sides of mantle,
hooks very large, dorsal very small); in juveniles, head, 4th arms, and dorsal sides of mantle and
one row of very small suckers between small fins. No photophores on eyeball. Larvae and juve-
hooks. In juveniles ventral hooks appear first. niles distinguishable by very wide, elliptical fin,
Buccal membrane deep purple in color. photophores on tentacular stem (at ML approx. 7
Photophores on ventral mantle in longitudinal mm), lacking on eyeball, and by characteristic pat-
rows in juveniles, diffuse in adults except bare tern of large mantle and head photophores (from
mid-ventral strip; on ventral head in 8 longitudinal ML approx. 5-6 mm). Tentacular hooks developed
rows with some dispersed between two parallel at ML approx. 9 mm. ML up to 40 cm.
central rows; in 3 longitudinal rows on base and 2
on tips of ventral arms (2 rows in juveniles). On Subfamily Pyroteuthinae
eyeball, ventrally, 5 round photophores, 1st and Pyroteutltis margaritijera (Rüppell, 1844) (Fig.
5th somewhat larger than other. Three round black 2.13). Mantle narrow, cup-like, with acute tail.
photophores on ends ofventral arms, in larvae first Fins in larvae andjuveniles tongue-shaped or oval,
1, appearing as a white ball, then 3, becoming in adults semicircular, not joined and not reaching
black. Usual egg size 1.2-1.3 mm. ML up to 6.0- mantle end. Arms rather long and very strong, all
6.5 cm. pairs, incl. 4th, bear hooks in 2 rows, almost to
A very common species through all its range, tips. Tentacles long, thin; c1ub elongated, not wid-
sometimes found even over the shelf. A third trop- ened, in larvae with 1 row of large and 3 of small
ical-subtropical Atlantic species, A. suckers; in adults with 1 central row of 3-5 hooks
(PjejJerifeufhis) aflanfica Nesis, 1982, differs and 2 rows of suckers (suckers of ventral row
from the two described above by ventral photo- located between hooks). Gladius caudally nar-
phores on head and mantle forming longitudinal rowed into acute needle. Photophores: 10 in the
stripes (7 on head, 6 on mantle), and club without mantle cavity (2 large near anus; 3 c10sely spaced
carpal membrane and keel. Juveniles hardly dis- at level of brachial hearts; 2 on sides near gill
tinguishable. bases; 2, next to each other, at the level of sto-
mach; and 1 very small ventrally on the conus of
Subfamily Ancistrocheirinae gladius); 12 on ventral and caudal parts ofthe eye-
• Allcistroclteirus lesueurii (Férussac and ball (small and large, of different structure, located
d'Orbigny, 1842) (=Thelidiofeufhis alessandrinii in strict order, some large ones with green metal-
Vérany, 1851) (Fig. 2.12). Mantle rather wide, lic glitter); 6 or 7 under ski n on tentacular stem.
conical, in adults with short, soft tail. Fin tongue- Photophores in mantle cavity begin to develop at
like in larvae, very wide transverse-elliptical in ML approx. 5 mm, all developed at ML 20 mm.
juveniles, diamond-shaped, reaching anterior First eye photophores appearing in early larvae
mantle margin in adults. Arms long and strong, (ML 2-3 mm). Their distribution, as well as the
with 2 rows of suckers in juveniles and hooks in semicircular fins, acute tail end, and 2 rows of
adults. Tentac1es robust, central part of club in lar- suckers/hooks on 4th arms, are characteristic for
vae with 2 rows of suckers, in juveniles 2 rows of larvae and juveniles. In freshly caught larvae 2
hooks (median) and 2 rows of very large suckers pink patches: at base of tentac1e and at base of
(marginal), and in adults only hooks (8-9 in a club. Egg size 1.0-1.3 mm. ML up to 5 cm.
row), without suckers. Photophores: the large ones This species represented by series of local forms
not numerous, regularly arranged, the small very of subspecific, or may be even specific rank: P.
numerous. Twenty-two large hemispheric photo- addolux Young, 1972, and P. serrata Riddell,
phores in altemate groups of 4 and 2 on ventral 1985.
side of mantle (4"groups of 4, and 3 groups of 2 Pterygioteutltis giardi Fischer, 1895 (Fig. 2.14).
photophores); 16 photophores on head (5 around Mantle narrow, cup-like, with acute tail. Fins in
each eye orbit at varying distances from it, 1 each larvae and juveniles tongue-shaped or oval, in
near funnel and near base of 2nd and 4th arms); adults semicircular, not joined and not reaching
20-30 large and small photophores along tentacu- mantle end. Arms rather short but strong, 1-3 pairs
lar stem; one under skin near stem base; hundreds with 2-5 pairs of hooks, 4th pair barren, without
730 Cephalopoda
hooks (except in mature males). Tentacles long and broken upon collection. Arms in larvae with suck-
thin; club compact, barely widened, in larvae with ers only, hooks appearing at ML approx. 2.5 mm.
1 row of large and 3 of small suckers; in adults with Suckers in adults only on very tips of arms.
4 rows of suckers; no hooks. Gladius narrowed cau- Tentacles in larvae long, thin, with bunch of rath-
dally into acute needle. Photophores: 8 in mantle er large suckers on tip (8 in 2 rows), club reduced
cavity (2 large round near anus; 2 oval near brachial at ML approx. 12 mm, stems at 25 mm. Qne or
hearts, 3rd between the latter; 2, next to each other two photophores on ventral mantle, anterior to
at level of stomach; and 1 very small ventrally on body end; one elongate brown photophore on end
conus ofgladius); 15 (rarely 14) on ventral and cau- of each arm; some poorly noticeable (except in
dal parts ofthe eyeball (small and large, ofvariable live animals) photophores on head and in mantle
structure, located in strict arder, some large ones cavity. No large unpaired photophore on ink saco
with green metallic glitter); 4 under skin on tentacu- ML up to 30-50 cm.
lar stem. Anal and brachial-heart photophores begin The commonest species in the South Atlantic is O.
to develop at ML approx. 2.0-2.5 and 1.5 mm resp., megaptera (Verrill, 1885).
all photophores in the mantle cavity developing at Taningia danae Joubin, 1931 (=Cucioteuthis
ML > 10 mm. First eye photophores appearing at unguiculata) (Fig. 2.16). Mantle broadly conical,
ML 2 mm, at 5 mm they number 9-10. Their distri- tapering into short tail. Fin in larvae transverse-oval
bution, as well as the semicircular fms, acute tail (its width exceeding largely its length, equal to or
end, tentacles with a compact bunch of very small longer than ML), in juveniles and adults diamond-
suckers at the tip, and naked 4th arms, are character- shaped, barely short of anterior (but not posterior)
istic for larvae and juveniles. No pink patches on mantle margino Eyes in early larvae on short wide
tentacle in larvae. Egg size 1.0 mm. ML up to 25, stalks, subsequently not stalked. Arms short, thick.
rarely to 30 mm. Arm hooks appearing rather early (at ML >5 mm),
A circumglobal species, represented in the Atlantic and then suckers remaining only on very tips of
by the subspecies P. giardi giardi. Another Atlantic arms. Tips of 2nd arms devoid of suckers, with one
species, P. gemmata Chun, 1908, distinguished by very large, complex photophore, whose central yel-
the presence of hooks (2-8, mostly 3-7) only in a lowish luminescent part may be covered by two
ventral row on Ist-3rd arms, 2 rows of very small black lobes ("lips"); no photophores on other arms.
suckers on 4th arms (noticeable in juveniles in side Tentacles in early larvae longer than arms, then
view), and 14 (rarely 15) eyeball photophores; also shorter, with bunch of 8 rather large suckers on tip,
circumglobal but predominantly anti-equatorial club reduced at ML approx. 40-45 mm, but stems
species, represented by a series of local forms. remaining until much later. Large unpaired photo-
phore on ink sac, developed at ML approx. 5 mm.
Family Octopoteuthidae Gladius with parallel lateral sides. Easily distin-
Body conical, acute posteriorly. Rins very long and guished by extremely wide fin and photophore on
wide, reaching or barely short of anterior margin and tips of 2nd arms developing at ML approx. 4.5-5
posterior end of mantle, joined dorsally along longitu- mm. A very large squid, ML up to 170 cm. The
dinal midline. Arms with two rows of hooks covered only species ofthe genus Taningia.
by skin sheath. Tentacles only in larvae and early
juveniles, lacking in adults. Buccal membrane with 6 Family Onychoteuthidae
lappets, connectives to 4th arms attached ventrally. Mantle narrowly conical or cylindrical, muscular.
Funnel cartilage straight, elongated, widened posteri- Fins on posterior half, transverse-oval, heart- or dia-
orly. Gladius with wide vane, beginning near anterior mond-shaped. Anns with two rows of suckers with
end. With photophores. Two genera, at least 6 species smooth rings. Tentacles long and strong, club with 2
in the Atlantic Qcean. rows of hooks and 2 rows of marginal suckers, which
Octopoteuthis sp. (Fig. 2.15). Mantle tapering into may be reduced in adulthood. Fixing apparatus often-
short tail. Fin tongue-like in larvae, transverse- tacles well developed, oval. Buccal membrane with 7
oval in juveniles, diamond-shaped with rounded lappets, connectives to 4th arms attached ventrally.
angles in adults. Eyes in larvae on short wide Funnel cartilage straight, elongated. Gladius with nar-
stalks. Arms rather short, weak, their tips usually row vane and chitinous or cartilaginous end conus.
Cephalopoda 731
Photophores may appear on eyeball and near hindgut. Filippova, 1972, in the Antarctic to South Georgia;
Six genera, at least 9 species in the South Atlantic. and M. aeqllatorialis Thiele, 1920, supposedly
01lykia carriboea Lesueur, 1821 (Fig. 2.17). tropical species of unclear taxonomic status.
Mantle in larvae and juveniles barrel-shaped, in Kondakovia /ollgimana Filippova, 1972 (Fig.
adults cylindrical, narrowing posteriorly. Head 2.19). Mantle. soft, not very muscular, conical,
usually not retractable into mantle. Fins tongue- with short tail in adults. Fins transverse-oval in
like in larvae, transverse-oval with almost straight larvae and juveniles, widely heart-shaped without
posterior margin in juveniles and adults. In larvae tapering end in adults; fin length in adults approx.
3rd, and particularly, 4th arms developing much 40%, width approx. 50-60% ML. Arms and tenta-
¡ater than 1st and 2nd. Larval tentacular club with cles very long and weak, arrn length approx. equal
4 straight longitudinal rows of suckers, decreasing to ML. Club in adults with 30-40 (usually 32-33)
from ventral to dorsal row, later marginal suckers hooks and 6-10 rows of suckers on dactylus.
becoming larger than central ones, after that (at MI Surface of mantle and head with soft longitudinal
approx. 8 rnm) central suckers transformed into skin ridges or papillae, even in early larvae (ML 3-
hooks. In adults 12-14 pair of hooks, marginal 5 mm). Gladius not visible from dorsal, with short
suckers absent or only in small numbers on pro xi- (usually not more than 10% ML), triangular, carti-
mal and distal part of club. No nuchal folds on laginous rostrum posteriorly. No nuchal folds, no
head. Gladius narrowly lancet-shaped, not visible photophores. ML up to 100 (1 15?) cm.
from dorsal, with acute chitinous needle behind, Onyc!lotellt!lis banksii (Leach, 1817) (Fig. 2.20).
not protruding through the skin upon fixation. Skin Mantle narrow, cylindrical in anterior, conical in
smooth. Large chromatophores covering both dor- posterior half. Head can be wholly retracted into
sal and ventral sides of mantle, color intense, from mantle. Fins tongue-like in larvae, heart-shaped in
red-brown to deep-bluish (on dorsal mantle side). juveniles, diamond-shaped with short tapering tail
No photophores. Barrel-shaped mantle, straight in adults. Arrns rather short, strong. Only 1st and
posterior margin offin, intense pigmentation (from 2nd arrns in early larvae, 3rd appearing later only
ML approx. 3 rnm), and club armament are cha- at ML approx. 5 rnm. Tentacles long. Larval ten-
racteristic features. ML up to 10 cm. tacular club with 4 rather irregular rows of suck-
One species in the Atlantic; larvae epipelagic, ers, 2 central rows transformed later into hooks,
juveniles (but not adults) live at the surface (even those ofthe ventral row first (at ML approx. 12-15
during the day). mm). In adults 19-23 hooks, marginal suckers
Morotellt!lis robsoni Adam, 1962 (Fig. 2.18). absent in juveniles. Nuchal folds on head well
Mantle muscular, cylindrical in anterior, conical in developed. Gladius very narrow, with strong lon-
posterior part, with long tail in adults. Fins trans- gitudinal dorsal rib easily visible from dorsal, with
verse-oval in juveniles, heart-shaped with tapering acute chitinous needle behind, directed diagonally
end in adults; fin length in adults approx. 65%, dorsally and protruding through skin upon fixation
width approx. 50% ML. Arrns and tentacles long in larvae. Skin smooth. Larvae almost transparent,
and strong. Club in juveniles with 9, in adults with with sparse chromatophores, covering predomi-
26- 30 hooks; marginal suckers absent at ML > 15 nantly dorsal mantle side in juveniles and adults,
rnm; 4-5 rows of suckers on dactylus. Surface of red-brown. One elongate photophore on ventral
mantle, head and arms with fiat tubercles of irre- side of eyeball, 2 near the hindgut (one small near
gular form, smooth in early larvae. Gladius not vi- anus, one large and round in an arch-like bend of
sible from dorsal, with long cartilaginous rostrum the gut near its beginning, developed at ML
posteriorly, triangular in transverse section, semi- approx. 9-11 rnm). Degenerating mature females
translucent in juveniles; adult length 20-35% ML. described as Challnotellthis mollis AppellOf, 1890.
No nuchal folds, no photophores. ML up to 75 cm. Larvae easily distinguishable by acute needle pro-
Larvae and juveniles may be caught at night at the truding from posterior end, light in color, and usu-
surface. At least 3 other species in the South ally with head retracted. ML up to 17 cm.
Atlantic: M. illgens (Smith, 1881) in Subantarctic Of the approx. 5 species known in genus, only O.
and Antarctic areas, including the Patagonian shelf banksii, a species complex represented by some
and slope, usually at the bottom; M. knipovitchi local forms, is known from the Atlantic. Juveniles
732 Cephalopoda
may leap out of the water at night; adults widely Only one species in the South Atlantic. Larvae in
eurybathic. some places very common in the upper layers.
Family Gonatidae Family Chtenopterygidae

Mantle cylindrical in anterior, conical in posterior One genus in the World Ocean.
parto Fins heart- or diamond-shaped. Arms with 4 Chtellopteryx sic lila (Vérany, 1851) (=CtenopteIYx
rows of suckers, in juveniles or adults two inner rows sicula) (Fig. 2.22). Mantle short, wide (approx. 40-
on 1st-3rd arms transformed into hooks. Tentacles 60% ML), slightly depressed dorsoventrally, round-
long and strong, club with many rows of suckers, in ed behind. Head in larvae half-retractable (up to
some species a1so with one or more hooks in central midline of eye). Fins tongue-like in larvae, resem-
parto Fixing apparatus of tentacles either 1acking or bling fish fins in juveniles (ML >3.5 mm) and
represented by short but strong transverse ribs and adults, consisting of a series of flexible muscular
grooves. Bucca1 membrane with 7 1appets, connec- "rays" joined by thin transparent film. Fin in juve-
tives to 4th arms attached ventrally. Funnel cartilage niles much shorter, in adults a little shorter, than
straight, elongate. Gladius with narrow vane with 1-2 mantle. Arms short, 4th pair longer and wider than
widenings, without cariilaginous end conus, usually others. Arm suckers small, in 2 rows in proxima1, 4-
not reaching mantle end in adults. Photophores lack- 6 rows in distal part of 1st-3rd arms, in 1-2 zig-zag
ing (present in one species on eyeball). rows on 4th arms. Tentacles in larvae «4 mm)
Three genera. Only 1 species in the South Atlantic. short, with very distinctive round club with 6 irre-
GOllatlls (Gollatlls) alltarcticlIs Lonnberg, 1898 gular rows of suckers; in juveniles (>6 mm) and
(Fig. 2.21). Mantle narrow, adult width 20-25% adu1ts long, retractable, with a short narrow club
ML. Head in larvae can be wholly (including bearing 8-14 rows of very small suckers; no fixing
arms) retracted into mantle. Fins petaloid in lar- apparatus. Buccal membrane with 7 1appets, with
vae, transverse-oval in juveniles, heart-shaped 12-15 minute suckers in 2 rows on lappets, connec-
with tapering tail in adults. Arms rather short, in tives to 4th arms attached ventrally. Funne1 carti-
adults approx. 25% ML. Arm suckers trans- lage straight, elongate. Gladius with long rachis and
formed into hooks at ML approx. 25 mm. Nine wide vane, without end conus. Photophores on eye-
to ten transverse rows of suckers on basal halves ball (one elongated) and ink sac (large, round,
of 4th arms. Tentacular club rather short, in resembling a fried egg on a black frying pan); in
adults approx. 15-24% ML, moderately wide. In adult males also dorsocaudally on the mantle. Ink
adults central part with one large central, one sac photophore appearing in late larvae. Egg size
smaller distal, and 3-5 small proximal hooks, approx. 1.5 mm. ML up to 9, usually to 5-6 cm. One
surrounded by naked space; 4 rows of small species in the Atlantic.
suckers on dactylus and on ventral side of cen-
tral part, 2-4 on dorsal side. Fixing apparatus Family Bathyteuthidae
represented by short strong transverse ribs (each One genus in the World Ocean.
with 1 sucker), and grooves. In laryae whole ten- Bathytellthis abyssicola Hoyle, 1885 (Fig. 2.23).
tacular stem with 4 rows of suckers, zone of Body cy1indrical, rounded behind. Head wider
growth occupying distal half of larval club. At than mantle, eyes directed somewhat anteriorly.
ML approx. 15-20 mm, 2 large buds appear in Fins in both 1arvae and adults small, kidney-
central part club, later transformed into hooks. shaped, 10cated near mantle end, not joined. Arms
Central, distal, and proximal hooks developed at short, conica1, not tapering, joined by web near
ML approx. 20-25, 25-30 and >30 mm, respec- bases, 4th pair in adults 10nger and wider than
tively. Gladius narrowly lancet-shaped, with two others. Suckers small, on lst-3rd arms in 2 rows
widenings. Radula with 5 longitudinal rows of proximally, 4 distally, on 4th arms in 1-2 rows.
teeth. No photophores. Egg size supposed1y Protective membranes of anns well developed,
about 5 mm. Larvae distinguishable by thin thick, with entire margin, without finger-like lap-
mantle with sparse, large chromatophores, and pets. Tentacles long, thin, retractable, club not
by large spindle-shaped midgut gland; juveniles widened, without fixing apparatus, with very
by club structure. ML up to 35 cm. small suckers in 6-8 rows. Buccal membrane with
Cephalopoda 733
7 lappets, with 1-2 minute suckers on lappets, con- each with 9 photophores; 4 photophores along
nectives to 4th arms attached dorsally. Funnel car- midline of ventral side of head; 4 longitudinal
tilage straight, elongate. Gladius with thin rachis rows of photophores (3 rows of large, 1 short row
and oval vane, without end conus. Photophores of small) at the base of 4th arms, 2-3 on lst-3rd
visible only in late larvae and juveniles (on basal arms; no elongate black photophores on arm ends;
parts of 1st to 3rd arms), obscure in adults. Color 17 large and 1 small photophore around right eye;
uniformly deep-maroon, tentacles whitish. an arch of 7 large photophores anterior to, and 10-
Position in life vertical, head down. Larvae and 14 small ones lateral and posterior to left eye. Skin
juveniles adult-like, but with protruding eyes, also not papillate. Larvae with large eyes, globular arm
deep-maroon, arm photophores developing at ML suckers, mantle and head photophores develop at
approx. 5-6 mm. ML up to 75 mm. ML approx. 5 mm. Larvae covered by many large
One species in the Atlantic. A widely distributed chromatophores. Egg size: length 1.8 mm, width
species, circumglobal in Antarctic and polar-subpo- 1.5 mm. ML up to 18-19 cm.
lar zones; absent in the Gulf of Mexico, Caribbean H. rever.,sa does not occur in the Gulf of Mexico,
and Mediterranean seas, as well as the equatorial Caribbean Sea and South Atlantic Central Water
and central zones ofthe Westem Atlantic. Mass, but is cornmon in the Mediterranean Sea,
reaching 22-23°S offNamibia.
Family Histioteuthidae Histioteuthis bOllllellii (Férussac, 1834) (Fig.
One genus in the W orld Ocean. Mantle conical, head 2.25). Mantle short, cup-like. Fins in adults
usually much wider than mantle. Fins small, round, extending beyond mantle end. Arms in larvae as
located on mantle end, joined. Left eye usually much long as mantle, in adults 1.5-3 longer. Inner arm
larger than the right. Arms long, conical, with 2 rows web in juveniles a third, in adults 50-60% of arm
of suckers, in some species joined by web near bases length, outer web poorly developed. Arm suckers
from outer and/or inner side. Tentacles very long, club rather small in juveniles, globular, with low, bare-
with 4-8 rows of suckers commonly of strongly ly noticeable teeth. Tentacles very long. Club
unequal size. Fixing apparatus long, consisting of suckers in 5-6 rows, equal in larvae but in adults
knobs and suckers. Buccal membrane with 6 or 7 lap- median suckers in 2 central rows much larger than
pets, connectives to 4th arms attached dorsally. others. Buccal membrane with 6 lappets, connec-
Funnel cartilage straight, slightly curved and widened tives to 4th arms attached to dorsal parts of 4th
caudally. Gladius with thin rachis and wide vane, arms and to middle of web joining 3rd and 4th
without end conus. Photophores very numerous, arms. Large ventral mantle photophores distribu-
large, with different and complicated structure, cover- ted in 7-8 diagonal rows, each with 7 photophores,
iRg ventral (rarely also dorsal) side of mantle, head, gradually diminishing posteriorly; 4-5 photo-
and arms; only fins and tentacles are devoid of photophores along midline of ventral head surface; 3
phores; also lacking on eyeball and inside mantle ca- longitudinal rows at the base of all arms; one long
vity. The only genus, Histioteuthis, is represented in dark photophore on ends of lst-3rd arms, short
the South Atlantic by 9 species. one on 4th arms; 17 large photophores around
Histiotelltizis reversa (Verrill, 1880) [=H. elonga- right eye; an arch of 7 large photophores anterior
ta (N. Voss and G. Voss, 1962)] (Fig. 2.24). to, and approx. 10 small ones lateral and posterior
Mantle short, cup-like. Fins in adults extending to left eye; 3 conspicuous large round black pho-
beyond mantle end. Arms in larvae shorter than tophores posterior to left eye on ventral side of
mantle, in adults arm length not more than 1.5 head. Skin not papillate. Larvae with large eyes,
times ML. Outer arm web absent, inner vestigial. relatively short arms, globular arm suckers, and
Arm suckers rather large in juveniles, globular, in thick tentacles. Mantle photophores are developed
adults with smooth or serrated margino Club suck- at ML approx. 5 mm, arm end photophores on lst-
ers in 4 rows on manus, equal in larvae, of diffe- 3rd arms at 7 mm; larvae covered with many large
rent size in adults, median 3 much larger than oth- chromatophores. Six-lappet buccal membrane and
ers. Buccal membrane with 7 lappets. Ventral arm.-end photophores are particularly characteris-
mantle photophores intermixed large and small, tic. Egg size 2.3 rnm. ML up to 33 cm.
large ones distributed in approx. 7 diagonal rows, H. bonnellii is distributed in the Atlantic from SW
734 Cephalopoda
Greenland and Iceland to the Southem Subtropical Fins semicircular or kidney-like in larvae and
Convergence, including the Mediterranean, but juveniles, extending somewhat beyond mantle
not the Gulf of Mexico, Caribbean Sea and both end, longitudinal-oval in adults, usually shorter
Central Water Masses. Also present in the south- than half of ML. Arms short. Suckers on Ist-3rd
em subtropical and temperate belt (20-30° to 40- arms transformed into hooks in adults. Tentacles
45°S) of the S Indian and SW Pacific Ocean. short (shorter than arms in larvae and juveniles),
thin, club slightly widened. Carpal part with very
Family Architeuthidae many small suckers, central part with 2 rows (3-4
One genus in the W orld Ocean. pairs) of very large ones and 2 marginal rows of
• Architeuthis dux Steenstrup, 1875 sensu lato (Fig. smaller, on dactylus small suckers in 3 rows.
2.26). Giant squids, largest of all invertebrates. Fixing apparatus: 2 close rows of altemating suck-
Mantle narrow, cylindrical, tapering into short tail. ers and knobs along entire (in juveniles) or distal
Fins small, transverse-oval in larvae, longitudinal- part of stem. Kidney-shaped fin s, short thin tenta-
oval in juveniles, semicircular or semioval, not cles, and large central club suckers are characte-
reaching mantle end in adults. Head moderately ristic for juveniles. ML up to 27 cm.
wide. Arms very long, tentacles extremely long. A. antarcticus rarely occurs north of the Antarctic
Arms with 2 rows of suckers. tentacular club with Convergence.
a group of small suckers in many rows in carpal Neoteuthis thielei Naef, 1921 (Fig. 2.28). Mantle
part and 4 rows of suckers in long central part; narrowly-conical, narrower than head, long, poin-
suckers oftwo central rows much larger than mar- ted behind. Fins in larvae and juveniles narrow
ginal. Fixing apparatus stretched along almost and long, beginning in anterior third of mantle,
entire stem. Funnel cartilages simple, straight. widening behind; in adults like narrow strip along
Connectives from buccal membrane attached to most of the mantle, not joined posteriorly. Arms
4th arms dorsally. Gladius with short wide rachis, short, particularly in juveniles, with 2 rows of
acute behind, with small end conus, not reaching rather large suckers, not transformed into hooks.
mantle end. No photophores. ML up to 5 m, usu- Tentacles short and thin in juveniles, long in
ally to 1-2 m. adults, very small suckers in many rows occupy-
Supposedly 3 subspecies or vicariant species with ing most of widened club, abruptly changing into
circumglobal bi-subtropical or anti-equatorial dis- 4 rows of large suckers only in distal part, of
tribution: A. dllx dllx Steenstup, 1875, in the North which the central ones are not larger than the mar-
Atlantic; A. dux martensi (Hilgendorf, 1880) in the ginal. No fixing apparatus. ML up to 17 cm.
North Pacific; and A. dux sanctipauli (Vélain,
1877) in the Southem Hemisphere. Adults com- Family Brachioteuthidae
monly caught near the bottom in the upper One genus in the World Ocean.
bathyal. An undescribed small species of this Brachioteuthis riisei (Steenstrup, 1882) (Fig.
genus was recorded off Florida and Madeira. 2.29). Mantle narrow, cylindrical or becoming
conical ahead of the end of the anterior fin. Head
Family Neoteuthidae narrow, but not retractable into mantle. Neck in
Mantle conical. Fins widened posteriorly. Arms with larvae long, sometimes grotesquely so, eyes below
2 rows of suckers, sometimes converted into hooks in midline of head. Fins petaloid in larvae, diamond-
adults. Tentacular clubs with many small suckers in shaped in juveniles and adults, straight or weakly
the carpal section, 4 rows of large suckers in the cen- tapering behind, fin length less than width, not
tral section, 3-4 smaller ones on dactylus. Buccal exceeding half ML. Arrns short, with 2 rows of
membrane with 7 lappets, connectives to 4th arms suckers. Tentacles in larvae and juveniles much
attached dorsally. Funnel cartilage simple, straight. longer than arms, club wide, with 2-4 rows of
Gladius with thin rachis and wide vane. No photo- rather large suckers in carpal, 6-8 rows of knobs in
phores. 3 genera and supposedly 3 species in the distal part, in adults relatively shorter, with many
South Atlantic. rows of small suckers in carpal, 4-6 rows of larger
• Alluroteuthis alltarcticus Odhner, 1923 (Fig. ones in distal parto Transition from larval to adult
2.27). Body cylindro-conical, head not retractable. pattem at ML > l O mm. Fixing apparatus: 2 rows
Cephalopoda 735
of small suckers along stem. Buccal membrane teristic, often called rhynchoteuthis or rhynchoteuth-
with 7 lappets, connectives to 4th arms attached ion. They are bom with both tentacles fused together
ventrally. Funnel cartilage straight, simple. and forming a narrow "trunk" (proboscis), as in a
Gladius with thin, very long rachis and narrow baby elephant. On the tip of this trunk a diamond-
vane. In some forms a photophore on ventral eye- shaped snout plate stretches laterally, with 8 suckers:
ball. Eggs oval, 1.2-1.5 mm in greater axis. Larvae 3 on the each dorsal and ventral sides, of the same
and juveniles easily distinguishable by narrow size, and 2 on the lateral angles, larger in some gene-
head, long neck, and many rows of carpal club ra than others. In late larvae the proboscis begins to
suckers, light in color, with sparse chromatophores. divide at the base, the cleft gradually moving forward,
Juvenile appearance at ML approx. 15 mm. ML up lateral angles of the snout elongating, and finally the
to 17 cm in Antarctic, up to 8 cm in other areas. proboscis divides fully and the tentacles separate. The
A complex of species with unclear relationships. lateral angles become tentacular clubs and snout suck-
B. riisei (Steenstrup, 1882) is supposedly a North ers - rudiments of the fixing apparatus. In some ge-
Atlantic and southem circumglobal bi-temperate nera, at the time of proboscis division the lateral
species with a tropical-subtropical circumglobal angles of the proboscis tip begin to elongate, forming
counterpart [B. behni (Steenstrup, 1882)] and smalllobes resembling "ears", rudiments of club ends
another, as yet unnamed, circum-Antarctic species. (dactyli). The mantle in larvae is at first barrel-shaped,
Eye photophores are reported in adult B. riisei but later cylindrical, fins are short, petaloid. Relative
not in two other. The larvae of B. be/mi are charac- length of the proboscis, size relationships of lateral to
terised by a very long neck and eyes directed at other suckers, and presence/absence of gut and eye-
right angles to the longitudinal head axis; those of ball photophores are characters used for the identifica-
B. riisei and of the Antarctic form by a moderate- tion of larvae. ML in newbom larvae is usually about
ly long neck and eyes directed anteriorly at acute 0.8-1.0 mm. Five subfamilies, 11 genera. Nine genera
angles to the longitudinal head axis. Another spe- and 11-12 species in the South Atlantic.
cies in the South Atlantic - B. picta Chun, 1910,
has a photophore ventrally on the eyeball. Subfamily IIIicinae
One genus in the World Ocean.
Family Ornrnastrephidae • IIlex argelltilllls (Castellanos, 1960) (Fig. 2.30).
Large or medium-sized active nektonic squids. Mantle narrow, its width 1/4-1/6 ML. Fin heart- or
Mantle muscular, cylindrical or conical. Fin at poste- narrowly diamond-shaped, width 45-60% ML.
rior mantle end, transversely diamond- or heart- Eight transverse rows of small suckers at club end.
shaped. Head large. Arms long, with strong swim- Teeth of large tentacular suckers approximately
ming keels and two rows of suckers with toothed equal in size. Carpal group of tentacular suckers
rings. Tentacles strong, with 4 rows (8 rows at the short, compact, fixing apparatus without knobs.
club end in JIlex) of unequal suckers, also with Funnel pit without foveola and side pockets. No
toothed rings, and a short fixing apparatus usually photophores. ML up to 40 cm. Larvae: probo seis
consisting of knobs and suckers. In many genera the short, usually 1/4-1/2, rarely to 3/4 ML, usually
anterior part of the funnel pit is separated from the shorter than arms. AII suckers on the proboscis tip
posterior part by a crescent-like skin fold (foveola); in equal in size. Proboscis division beginning at ML
some genera, posterior to it, at both sides of the fun- approx. 4-5 mm, finished at 6-8 mm. Arms of 3rd
nel, some skin pockets are developed. Funnel carti- pair beginning to develop (first suckers appear) at
lage with two deep furrows crossing in the form of an ML approx. 1.5 mm, of 4th pair at 3.5 mm.
inverted "T"; mantle cartilage with two strong crossed Tentacular ends in juveniles from ML 10-15 mm
ridges. Funnel and mantle cartilages merged together with more than 4 rows of suckers.
in the postlarvae of two species. Buccal membrane Two species in the South Atlantic: Illex argentinus
with 7 lappets, connectives to 4th arms attached ven- (Castellanos, 1960) distributed in the SW Atlantic,
trally. Many species have diverse photophores on a very common and important commercial spe-
mantle, arms, eyeball ancl/or gut. Gladius narrow, cies; and a close species 1. coindetii (Vérany,
without fan, with short conus. Larvae are very charac- 1837), distributed on both sides of tropical
736 Cephalopoda
Atlantic. Both inhabit near-bottom layers on the Subfamily Ornithoteuthinae

shelf and slope and pelagic layers over the shelf, One genus in the W orld Ocean.
slope, and "nearshore abyssal" areas. Larvae and Ornithoteuthis alltillarum Adam, 1957 (Fig. 2.32).
juveniles seem indistinguishable. A third species, Mantle narrow, conical, tapering into thin tail.
I. illeeebrosus (Lesueur, 1821), also an important Head wider than mantle. Fin heart-shaped, tapering
commercial squid, inhabits the N Atlantic. posteriorly, its length more than 50% ML. Six-
seven pairs of large tentacular suckers, rings with-
Subfamily Todarodinae out especially enlarged teeth. Carpal group of
Three genera, 2 in the South Atlantic. suckers short. Fixing apparatus weakly developed,
Todarodes angolensis Adam, 1962 (Fig. 2.31). without knobs. Funnel pit with foveola with longitu-
Mantle robust, cylindrical. Fin heart-shaped, dinal skin ridges and weakly developed side pockets.
length approx. 50% ML. Four transverse rows of Fhotophores in mantle cavity: one round near anus
small suckers at club end. Fifteen to eighteen pairs (large), one oval near beginning of gut (small), and
of large central tentacular suckers, diameter one narrow strip of luminous tissue from the latter
approx. 2.5% ML, with teeth of approximately the to very end of the body. One elongate photophore
same size. Carpal group of 2 pairs of tentacular on ventral side of eyeball. ML up to 30 cm. Larvae:
suckers. Length of sucker-bearing part is 2/3-3/4 proboscis short, 1/4-1/2 (rarely to 3/4) ML, thick,
of total tentacle length. Fixing apparatus with longer than arms in very early larvae, then equal or
knobs. No rows of finger-shaped lappets on ten- shorter. Two lateral suckers on tip are up to 2 times
tacular stem, no cirrus-like outgrowths of protec- larger than others. Proboscis beginning to split
tive membranes on arms and tentacular club. early, at ML approx. 4-4.5 rnrn, finishing at approx.
Funnel pit with foveola with small longitudinal 5-6 rnrn. Arms of 3rd and 4th pairs beginning to
skin ridges, without side pockets. No photophores. develop at ML approx. 1.5 and 2.5-3 rnrn. Anterior
ML up to 44 cm. Larvae: proboscis rather short, (anal) gut photophore yellowish, developed at ML
usually 1/2 ML in early larvae, 1/3-1/4 ML in 3-3.5 rnrn, posterior photophore whitish and deve-
older larvae, usually not longer or not much longer loped at approx. 8 rnrn, eyeball photophores at ML
than arms. AII suckers on the proboscis tip more or 4-4.5 rnrn, visceral luminous strip in juveniles> 30
less equal in size. Proboscis division beginning at rnrn. Larval mantle with many large brown chro-
ML approx. 4.5-5 rnrn, finishing at approx. 8 rnrn. matophores. Early larvae may wholly retract into
Fourth arms beginning to develop at ML approx. mantle.
2.5-3 mm; first suckers appearing on arms of 3rd Two close species, distinguishable only by adult
and 4th pair at ML approx. 1.5 and 6-7 rnrn, males: a tropical Atlantic O. antillarllm and its
respectively. Tentacular ends (dactyli) in juveniles Indo-Pacific counterpart O. volatilis (Sasaki,
with 4 rows of suckers. 1915). Both reported off Namibia (O. antillarllm
T. angolensis is known from the SE Atlantic, the by Laptikhovsky, 1989; O. volatilis by Villanueva
Tasman Sea and around New Zealand, and in and Sánchez, 1993). These nerito-oceanic species
some parts of the Southem Indian Ocean; it inha- live mostly in midwater over the slope and sea-
bits the pelagic realm and near-bottom layers over mounts (or not far from these), and near the bot-
the shelf and slope. A closely related open-ocean tomo
subantarctic species, T. jilippovae Adam, 1975,
has 12-14 pairs of large (diam. 3-4.5% ML) cen- Subfamily Ommastrephinae
tral club suckers and 4-5 pairs of carpal suckers; Five genera, 4 in the South Atlantic.
ML up to 56 cm. Its larvae are not described but Ommastrephes bartramü (Lesueur, 1821) (Fig.
probably are not distinguishable from those of T. 2.33). Mantle cylindro-conical, narrowing behind,
angolensis. Another genus, Martialia (one spe- rather wide in larvae and adults, very narrow in
cies, M. hyadesi Rochebrune and Mabille, 1887), juveniles. Head not retractable. Fin broadly dia-
also a subantarctic circumglobal species, is very mond-shaped, with very slightly tapering end. Fin
common in the SW Atlantic, both in midwater and width 2/3-3/4 ML. Arm ends not tapering. Ventral
at the bottom. Its larvae are not described. protective membranes on 3rd arms widened, in
Cephalopoda 737
adult females form a wide triangular lobe. Tentacles 4th arms, grouped to form large oval, orange lumi-
rather short and narrow. Rings oflarge club suckers nous spot on dorsal mantle muscles in its anterior
with 4 long teeth located crosswise, others smaller. part; one eyebal1 photophore, two nearly equal gut
Fixing apparatus with knobs and suckers; photophores, near anus and near beginning of gut.
4- 7 small dentate suckers proximal to first knob on Dorsal photophore developed only before sexual
tentacular stem. Funnel pit with foveola, longitudi- maturation, at ML approx. 8-9 cm. No silvery
nal ridges, and side pockets. Funnel and mantle car- reflecting cigar-like stripe along ventral mantle,
tilages never fused together. Many small photo- wide dark stripe along dorsal midline not bordered
phores dispersed under skin of mantle, head, and 4th by thin ones. ML up to 65 cm. Larvae: proboscis
arms. No dorsal, eyeball, or gut photophores. A sil- long, thin, usual1y not less than 2/3 ML and 2-3
very, reflecting, cigar-like stripe along ventral man- times as long as arms; lateral suckers on tip larger
tle from anterior part to anterior fin margin, wide than others. Proboscis beginning to split at ML
dark stripe bordered by thin reddish ones along dor- approx. 4.5-5 mm, finishing at 9-10 mm. Third
sal midline. ML up to 86 cm in the N Atlantic, to 65 and 4th arm pairs developed at ML approx. 1.5-1.7
cm in the S Atlantic. Larvae: proboscis rather long, and 2.0-2.5 mm, respectively. Mantle in larvae
thin, 1/2-2/3 ML, may be equal to ML in early lar- barrel-shaped, in juveniles narrow, pencil-like.
vae, usually longer than arms. Lateral suckers on tip Head not retractable into mantle. Eyeball photo-
noticeably (up to 2 times) larger than others. phores developed at ML approx. 3-3.5 mm, ante-
Proboscis beginning to split at ML approx. 5 mm, rior (anal) and posterior gut photophores at 3-4
fmishing at 7-9. Third and 4th arm pairs developed and 5-5.5 mm, respectively. Mantle in larvae with
at ML approx. 1.5 and 2.5-3 mm, respectively. dispersed large brown chromatophores, in juve-
Mantle in larvae barrel-shaped, in juveniles very niles with crimson and brown chromatophores.
narrow, cylindrical, pencil-like. Head not retract- One species in the Atlantic Ocean. Foraging range
able into mantle. Mantle in larvae with dispersed between approx. 34-36°N and S, reproductive
large brown chromatophores, in juveniles with range between approx. nON and 200S (off
crimson and brown chromatophores. Namibia southern boundary of reproductive range
Only species with disjunct bi-subtropical range: near 15-18°S, of foraging range usual1y near 23-
North and South Pacific, North and South Atlantic 24°S). Epipelagic oceanic species, very common
(in the latter from 13-14°S to 50-55°S in the SW and potentially commercial.
and to 35°S in the SE Atlantic), and Southern Eucleoteuthis lumillosa (Sasaki, 1915) (Fig.
Indian Ocean. A very common species, subject to 2.35). Mantle cylindro-conical, narrowed behind,
strong commercial exploitation in the N Pacifico head not retractable. Fin heart-shaped, with taper-
Sthelloteuthis pteropus (Steenstrup, 1855) (Fig. ing end. Ventral protective membranes on 3rd
2.34) (=Ommastrephes pteropus, Sthenoteuthis arms not widened. Rings of large club suckers with
pteropus). Mantle cylindro-conical, narrowing one long tooth in the centre of distal side, others
behind, rather wide in larvae, robust in adults, nar- smal1er. Fixing apparatus with knobs and suckers.
row in juveniles. Head not retractable. Fin broad- Funnel pit with foveola, longitudinal ridges, and
ly diamond-shaped, not tapering into tail. Fin usual1y side pockets. Funnel and mantle cartilages
width 3/4 ML or more. Arm ends not tapering. fused. Photophores: 1 each on ventral side of eye-
Ventral protective membranes on 3rd arms wid- bal1; 1 each in middle and near base of 4th arms;
ened but not forming a wide triangular lobe, width two long narrow luminous stripes on ventral man-
approximately equal to arm width. Tentacles rath- tle along the sides of ventral midline from anterior
er narrow but strong. Rings of large club suckers to posterior end, interrupted near anterior end and
with 4 long teeth located crosswise, others small- near line of maximal fin width; many tiny simple
er. Fixing apparatus with knobs and suckers; not photophores dispersed under skin of mantle; one in
more than 2 small dentate suckers proximal to first mantle cavity on gut at boundary between middle
knob on tentacular stem. Funnel pit with foveola, and posterior third of length. No dorsal photo-
longitudinal ridges and side pockets. Funnel and phore. ML up to 22 cm. Larvae: proboscis rather
mantle cartilages not fused. Many small photo- long, 1/2-2/3 ML, in early larvae longer, in later
phores dispersed under skin of mantle, head, and shorter than arms; lateral suckers on tip up to 1.5
738 Cephalopoda
times larger than others. Proboscis beginning split Family Thysanoteuthidae

at ML approx. 5-5.5 mm, finishing at 9-10. Fourth One monospecific genus in the World Ocean.
arm pair developed at ML approx. 2 mm. Mantle Thysanoteuthis ,hombus Troschel, 1857 (Fig.
narrow. Funnel and mantle cartilages fused at ML 2.37). Large muscular nektonic squids. Mantle in
approx. 8-11 mm. Eyeball photophores at ML larvae sac-like, in juveniles broadly cup-like, in
approx. 3.5 mm, gut photophore at 3-3.5 mm, adults nearly cylindro-conical. Head wider than
luminous stripes on the mantle at ML approx. 20 mantle and may retract into it only in newborn lar-
mm. Single gut photophore and luminous stripes vae. Fins petaloid in larvae, kidney-shaped in
are characteristic features. Mantle in larvae with juveniles, in adults longitudinal diamond-shaped,
dispersed large brown chromatophores, in juve- reaching anterior and posterior mantle ends but
niles with crimson and brown chromatophores. not fused. Arms short but strong, with 2 rows of
One species with disjunct range of bi-peripheral suckers, in juveniles relatively longer than in both
type: N and S Pacific, South Atlantic, and S Indian larvae and adults. Third arms are longest, 4th wid-
Ocean. ened at base. All anns bear wide protective mem-
Hyaloteuthis pelagica (Bosc, 1802) (Fig. 2.36). branes with long cirrus-like trabeculae. Tentacles
Mantle narrow, cylindrical, head not retractable. al so short and strong, club widened, with 4 rows
Fin diamond-shaped, not tapering behind, posterior of suckers. Fixing apparatus: 2 rows of alternating
margins straight. Second arms with some enlarged knobs and suckers along stem. Buccal membrane
suckers. Ventral protective membranes on 3rd arms with 7 lappets, connectives to 4th arms attached
not widened. Club narrow. Rings of large club ventrally. Funnel cartilage in the form of a tilted
suckers with one long tooth in centre of distal side, "T", with narrow projection in upper and rectan-
others smaller; marginal suckers minute. Fixing gular in lower parto Nuchal cartilage with 2 knobs
apparatus with knobs and suckers. Funnel pit with fitting 2 pits near anterior margin of the mantle.
foveola, longitudinal ridges, and usually side pock- Gladius is like tip of·arrow with short stem. No
ets. Funnel and mantle cartilages not fused. photophores except probably one on the ink sac in
Photophores: 1 each on ventral side of eyeball; 3 juveniles, lacking in larvae and imperceptible
round patches near end, middle, and at base of 4th (probably not functional) in adults. The characte-
arms; 19 round patches on ventral mantle side dis- ristic egg spawn is a mucous stocking-like cylinder
tributed in altemating groups oftwo and four, cau- 1-2 m long, 10-30 cm in diameter, floating
dalmost one not paired; many tiny simple photo- obliquely with one end on the very surface. A
phores dispersed under ski n of mantle; one in string of viscous mucus containing 2 parallel rows
mantle cavity on gut at boundary between middle of rose-violet eggs approx. 1.5 mm in diameter
and posterior third oflength. No dorsal photophore. wound around the cylinder. Larvae and juveniles
ML up to 9 cm. Larvae: proboscis rather long, 1/2- also very characteristic: larvae have barrel-like
3/4 ML, in early larvae longer, in late larvae short- mantle densely speckled by large violet chromato-
er than arms; lateral suckers on tip up to 1.5 times phores (fading after prolonged fixation), small
larger than others. Proboscis beginning to split at head and protruding eyes (ML <5 mm); juveniles
ML approx. 3.5-4 mm, finished at approx. 6-6.5 (> 1O mm) - widened bases of 4th arms and wide
mm. Fourth arm pair beginning to develop at ML protective membranes on 2nd-3rd arms. ML up to
approx. 3 mm. Mantle narrow. Larvae have only 1 m.
eyeball and gut photophores (at ML approx. 1.5 Larvae and juveniles epipelagic, adults rise to very
mm), luminous patches on mantle beginning to surface at night, descend at day to mesopelagic
develop at ML approx. 9 mm, quite distinct at 15 layers. Living in pairs (male + female) beginning
mm. Single gut photophore and round luminous from the late juvenile stage.
patches are characteristic features. Mantle in larvae
with dispersed large brown chromatophores, in Family Pholidoteuthidae
juveniles with crimson and brown chromatophores. One genus in the World Ocean.
One bi-central oceanic species, known from the Pholidoteuthis boschmai Adam, 1950
Atlantic and Pacific oceans but not reported from (=Tetronychoteuthis dussumieri Pfeffer, 1912,
the Indian. non d'Orbigny, 1839) (Fig. 2.38). Mantle cylindri-
Cephalopoda 739
cal, in juveniles and adults covered by fiat poly- Family Cycloteuthidae

gonal papillae or scales, usually not contacting one Mantle broadly-conical. Fin large, round, its length
another, bearing cartilaginous denticulations on >2/3 ML. Arms with 2 rows of suckers. Tentacles
angles, which resemble papillae of sea stars. In long, club compact, widened, with 4 rows of suckers,
adults denticulations usually broken off and papil- fixing apparatus weakly developed. Buccal membrane
lae arranged in oblique rows so that skin becomes with 7 lappets, connectives to 4th arms attached ven-
cobble-road-like. Fin diamond-shaped, reaching trally. Funnel cartilage triangular or irregular-oval,
mantle end. Fin length 35-45%, width 50-70% with deep pit for fitting nose-shaped or tubercular
ML. Arms with 2 rows of dentate suckers; maxi- mantle cartilage. Gladius with widened lateral vane,
mal arm length in adults 30-60% ML. Tentacles posteriorly either rounded or tapering into conus.
long, club almost not widened, with 4 rows of Photophores located on eyeball, head, and mantle SUf-
suckers, round in larvae andjuveniles (ML <20-25 face, ink sac, etc. Two genera.
mm), elongate and compressed from sides in late Cyclotellth;s s;rvellt; Joubin, 1919 (Fig. 2.40).
juveniles and adults. Acute teeth located on lateral Mantle sac-like in larvae, cup-like in juveniles,
sides of chitinous lining of slit-like opening of narrow cup-like, tapering into thin tail in adults.
sucker ring. Buccal membrane with 7 lappets, con- Head wider than mantle, eyes in larvae and juve-
nectives to 4th arms attached ventrally. Funnel niles on short stalks, protruding. Fins in larvae pe-
cartilage simple, straight. Gladius with narrow la- taloid, in juveniles transversely-oval, located on
teral vane, constricted in posterior third, and short posterior mantle end, in adults widely transverse-
needle-like rostrum on the tip of conus. oval, located in mid-mantle, fins do not join dor-
Photophores unreported. ML up to 60 cm. sally. Fin length 60-80% ML. Arms long, tentacles
The other South Atlantic species, P/¡. adami very long. Widened club with prominent aboral
G.Voss, 1956 (=Ph. urllguayellsis Leta, 1987), is keel. Central suckers in mid-part of club very large
bathyal, with heart-shaped fin (length 65-75% in larvae but not larger than marginal ones injuve-
ML), and long arms (maximum length 65-75% niles and adults. Funnel cartilage of characteristic
ML). ML up to 78 cm. Juvenile stages unde- form with one deep central pit from which two
scribed. Status ofN Atlantic P/¡. massyae (Pfeffer, grooves extend, one wide anterior and a narrow
1912), based onjuveniles, is unclear but it may be posterior. Mantle cartilage is an L-like crest with
a form of Ph. boschmai. knob on summit. Gladius lancet-like with long
acute end conus. A large photophore on ink sac,
Family Batoteuthidae noticeable in early juveniles, and a series of 15-20
One monospecific genus in the World Ocean. small ones, of different form and size, on eye iris
Batotellthis skolops Young and Roper, 1968 (Fig. (only in adults). No photophores on body surface.
2.39). Small squids with narrow spindle-like man- Larvae may be distinguished by protruding eyes
tle tapering into long thin tail which in juveniles and large central club suckers. ML up to 60 cm.
mayas long as other part of mantle. Head very Another species, C. akimllshki.ni Filippova, 1968,
small and narrow. Fins short and narrow, heart- with narrow retractable head and fin length
shaped or transverse-oval, located before begin- approx. 50% ML, was reported from the subtropi-
ning of tail. Arms short, thin, with 2 rows of suck- cal South Atlantic. Juveniles undescribed.
ers. Tentacles thin, long, club not widened, Discoteuth;s discus Young and Roper, 1969 (Fig.
suckers located in 6 rows along almost whole 2.41). Mantle widely-conical, "hot tapering into
length of tentacle. No fixing apparatus. Buccal tail, its anterior margin without finger-like fiaps.
membrane with 7 lappets, connectives to 4th arms Head wider than mantle, eyes in juveniles protrud-
attached ventrally. Funnel cartilage simple, ing anteriorly and sideways. Fin wide, transverse-
straight, slightly curved. Gladius narrow, stick- oval, its length in juveniles approx. 60% ML, in
like, with long conus. Photophores unreported. adults equal to ML. Arms rather short, commonly
Larvae undescribed, juveniles characterized by broken off at ends. Tentacles very long, club wide,
narrow mantle, very small fin, long tail, and shape with prominent aboral keel. Central suckers (4-5
of club. ML up to 27 cm. pairs in mid-part of club) very large, globular.
740 Cephalopoda
Funnel cartilage elongate-triangular with rather Chiroteuthis veranyi (Férussac, 1835) (Fig. 2.42).
shallow pit located near short side of triangle and Larvae are known as Doratopsis vermicularis
one narrow anterior groove. Mande cartilage: a (Rüppell, 1844). Mantle narrowly conical, of se-
low crest with knob on end. Gladius thick, paddle- migelatinous consistence, tapering into a thin tail.
like, with short rachis and parallel sides, without Fin oval or almost round. Head long, wider than
conus. A large unpaired p,b.otophore on outer man- mantle, eyes near middle ofhead. First arms short-
tle surface ventrally near end ofbody, some others est, 4th arms far longest, much wider than others,
around iris. No photophores on anterior mande, with widened ventral protective membrane. Arm
head, or ink saco Juveniles may be distinguished suckers on lst-3rd arms in two rows, on 4th in two
by very large fin, large central club suckers, and rows only in basal half, then zigzag-like and in one
tail photophore. Known ML up to 13 cm. row near the end. Arm suckers with acute teeth on
Another species, D. laciniosa Young and Roper, distal margino Suckers on 4th arms slighdy small-
1969, differs in having 2 pairs of photophores on er than on first three. Tentacles in adults 5-6 times
the ventro-anterior mande margin and on the head ML. Definitive club wide, without keel, with 4
between the bases of 3rd and 4th arms, a series of rows of very peculiar suckers (nearly 20 in a row)
finger-like flaps on anterior mande margin, no tail on thick, long, dark-colored pedicels, widening at
photophore, funnel cartilage with 2 grooves, large the middle, located so that pedicels of inner rows
central club suckers flattened. appear to grow from those of marginal rowS. Club
suckers with long acute median tooth and some
Family Chiroteuthidae smaller ones. Funnel cartilage ear-shaped, mande
Mande conical or sac-like, in larvae tapering into long cartilage nose-like. Funnel valve present. Two large
thin tail, usually broken off in adults at level of poste- round photophores on ink sac, two parallel elongate
rior end offin. Fin small, round or oval. Arrns of 1st to luminous stripes, two large oval and one small
3rd pairs with 2 rows of suckers, 4th usually much round photophore on ventral eyeball, a longitudi-
wider ancl/or longer than others, with very small suck- nal row of round photophores along ventral pro-
ers in 1 or 2 rows, sometimes only in basal part of armo tective membrane of 4th arms, one large complex
Tentacles extremely long, with thin stem, club wid- photophore on end of tentacular club, some small
ened, with 4-6 rows of suckers, with well developed ones on stalk. Adult squids yellowish-brown. ML
protective membranes, usually with keel. No fixing up to 16 cm. Larvae are very different from adults,
apparatus. Structure of club in larvae of some species almost transparent, elongate, rod-like, with cylin-
differs widely from that in adults, adult club developed drical mantle. On conus of gladius behind fin are 1-
in proximal part oflarval club, the latter then degener- 3 small additional fins. Primary fin butterfly-like,
ating or autotomizing. Buccal membrane with 7 lap- then transverse-oval. Head very narrow, with
pets, connectives to 4th arms attached ventrally. extremely long, transparent neck, subdivided by
Funnel cartilage ear-shaped, bottle-shaped, elongate- many transverse septa, and long (but shorter than
triangular, or oval, usually (but not always) with 2 pro- neck) narrow "snout" (arm-crown pillar). First 3
jections: tragus (on ventral side) and antitragus (on arms very short. Tentacular club narrow, keeled,
posterior side). Mande cartilage nose-like. Funnel club and part of the stem with 4 rows of ordinary
valve absent in some genera. Gladius very thin, rod- suckers on short pedicels, resorbed during transi-
like, with long conus, reaching far behind posterior tion to juveniles which occurs at ML approx. 50
end of fin. In some species larvae and juveniles have mm, although some larval features discemible at
another fin, different in shape from primary one and ML approx. 75 mm.
located in the conus of gladius. Photophores located on Five species, including 3 in the South Atlantic. C.
eyeball, 4th arms, tentacular club and ink sac, but may veranyi (incl. C. V. lacertosa Verrill, 1881) distri-
be absent. Transition from larval to juvenile and sub- buted in the tropical and subtropical Adantic and
adult stage is gradual and protracted; thus, specimens circumglobally in southem subtropical and sub-
with larval features may be rather large, in some cases antarctic waters from off South Africa to Peru and
the second fin and tail being even larger than in subad- Chile.
ults. Four genera plus some undescribed ones. For Grimalditeuthis bOllplandi (Vérany, 1837) (Fig.
most South Alantic species larvae undescribed. 2.43). Larvae are known as Doratopsis sagitta
Cephalopoda 741
Chun, 1908. This species is usually cited as the rarely with 3 or none. Mantle cartilage nose-like or
only member of a separate family, consists of 2 short ridges forming a right angle.
Grimalditeuthidae, but belongs to Chiroteuthidae Gladius narrow, with long conus, reaching far behind
according to Young (1992) because larvae are chi- posterior end of fin and forming a tail thread.
roteuthid-like. Mantle narrow-conical, soft, gelati- Photophores in various positions, may be on eyeball,
nous, transparent, tapering into long tail. Fin round on eyelid ventral to eye sinus, on ventral side of man-
or transverse-oval. Behind it another fin located on tle, fin, funnel and 4th arms, absent in some species.
conus of gladius, approximate1y equal in length but Development gradual, without clear larval stage.
heart-shaped; broken off in most subadult and adult Three genera.
specimens. Head long and narrow. Funnel in adult Mastigoteuthis spp. (Fig. 2.44). Mantle elongate,
squids fused with mantle, head not fused with it in smooth. Head wider than mantle. Eyes usually
nuchal region. Arms long, thin, subequal in length, large, protruding in larvae. Fin wide, heart-shaped
with 2 rows of suckers except at very ends. Sucker or oval, both halves fused along dorsal side of
sta1ks grow out of centre of thick three-topped mantle. In so me species fin continues beyond
tubercle. Qne elongate dark photophore on end of mantle end. Arms subequal in larvae, in adults 4th
each armo Tentacles present only in larvae, then pair much longer than Ist-3rd. Tentacles in larvae
resorbing, usually only stalk bases remaining in with 4-6 suckers, in adults with more than 15-20
juveniles, lacking in adults. Gladius very thin, with rows of suckers along almost entire circumference
long conus. ML up to 23 cm. Larvae transparent, of club. Approximately 10 species, about 7 in the
with long "snout" (arm-crown pillar) and even South Atlantic. Mostly bathypelagic, some also
much longer septate neck. Transverse line of dark bathyal, larvae predominantly mesopelagic.
chromatophores on ventral side of snout imme- Larvae distinguishable by long tail and club with
diately ahead of eyes. Fins petaloid. Funnel not minute suckers. Larvae of most species rema in
fused with mantle. Funnel carti1age bottle-shaped, undescribed.
mantle cartilage nose-like. Arms lst-3rd very
short, 4th pair much longer than others. Third arm Family Joubiniteuthidae
developed later than 1st-2nd, at ML approx. 4-5 Qne monospecific genus in the W orld Qcean.
mm"but soon (at approx. 15 mm) becoming equal JOllbinitellthis portieri (Joubin, 1912)
to 2nd. Tentacles well deve1oped, club short, (=Valdemaria danae Joubin, 1931) (Fig. 2.45).
keeled, with 4 rows of suckers (2 in very early Mantle narrow, cylindrical or conical, semige1ati-
larvae); suckers on distal part of stalk in 3 diagonal nous, tapering into very long (longer than mantle
rows. Transition to juvenile stage occurring at ML itself) acute needle-like tail, sometimes broken off
approx. 10-15 mm, but fusion of funnel to mantle in adults but present in larvae and juveniles. Fin
and disappearance of tentacles occurring much very small, oval or round. Head narrow, with ra-
later, between 20 and 40 mm (without second fin). ther long neck and small eyes. Arms lst-3rd
extremely long, thin, flexible, with thread-like
Fami1y Mastigoteuthidae ends and very small suckers in 6 rows; 4th arms
Mantle cylindrical in anterior and conical in posterior much shorter, with suckers in 4 rows. Tentacles in
part, semigelatinous. Fin large, round, oval, heart- or juveniles shorter than 1st-3rd arms, with thread-
diamond-shaped, usually longer than half of mantle. like stalk and long, laterally compressed club with
Arms with 2 rows of suckers, 4th usually much long- minute suckers in 8-12 rows; protective mem-
er than others. Tentacles extreme1y long, whip-like, branes present only on distal half of club. No fix-
with thin stem and barely widened club bearing mi- ing apparatus. In adults tentacles fully reduced.
nute suckers in many rows. No keel nor fixing appa- Funnel cartilage oval, without tragus and antitra-
ratus. In some species tentacles autotomized during gus, mantle cartilage an elongated tubercle.
capture. Buccal membrane with 7 lappets, connec- Buccal membrane with 7 lappets, connectives to
tives to 4th arms attached ventrally. Furmel cartilage 4th arms attached ventrally. No photophores.
ear-shaped or oval, usually with 2 projections: traglls Gladius needle-like, with very long conus. Known
(on ventral side) and antitraglls (on posterior side), ML up to 10.5 cm. Larvae with spindle-shaped
742 Cephalopoda
semitransparent mantle, very small petaloid fins Two rows of suckers on arms but many rows in
and needle-like taillonger than mantle. Neck rath- median or distal part of some arms in males of some
er short, without septa. "Snout" (arm-crown pillar) genera; in Mesonychoteuthis part of suckers in mid-
absent. 3rd arms appear at ML approx. 8 mm. parts of arms converted into hooks. Arms with protec-
Tentac1es thick and short, c1ub with 4 (at ML 6-9 tive membranes and weak keels. Tentac1es long, c1ub
mm), then 8 (ML 18 mm) rows of suckers. Thick short, not wide, with 4 rows of denticulate suckers, in
tentac1es and very long acute tail are characteris- 3 genera central suckers converted into hooks. Fixing
tic. ap'paratus well developed, stretched along the stalk.
Tentac1es autotomized in mature females of most ge-
Family Promachoteuthidae nera. Buccal membrane with 7 lappets, connectives to
One genus in the W orld Ocean. 4th arms attached ventrally. Gladius thin, rod-like,
• Promac!lOteuthis sp. (Fig. 2.46). Mantle wide, with diamond-shaped caudal widening (Ianceola), in
cup-like, thick-walled, semigelatinous, fused with some genera lanceola tapering caudally into a long
head in nuchal region. Head small, narrower than tai!. Rear end of mantle may not reach tail end. Fins
mantle. Only anterior part of nuchal cartilage attached to rear sides of lanceola, in some genera their
present. Eyes very smal!. Fin very large, widely anterior ends may reach in front of lanceola on man-
heart-shaped or transversely-oval, fin width tle sides. Lanceola reduced in Bathotlzauma. Funnel
approx. twice fin length. Arms short, subequal, valve present or absent. Photophores always present,
with 2 rows of toothless suckers. Tentac1es long, located usually on ventral side of eyeball, on arm ends
thick, robust, narrowing toward the end. Club not (in adult females of some genera), in Megalocranchia
differentiated, without protective membranes and also on ventral side of digestive gland. The coelom is
keel, beset with minute suckers in many (8-20) very voluminous, closed and filled with a weak solu-
rows. No fixing apparatus. Buccal membrane tion of NH4Cl, forming a float like in a bathyscaph
weakly developed, with 7 lappets, connectives to ("bathyscaphoid squids"). Mantle cavity subdivided
4th arms attached ventrally. Funnel cartilage oval, by thin horizontal membrane into two chambers,
without tragus and antitragus. No photophores. upper (inc1uding coelom) and lower, both connected
Gladius strongly reduced, with rather wide fan by two round openings over gills. Nidamental glands
and, in one species, thin rachis. ML up to 18 cm. subdivided into two parts and united with oviductal
Larvae with very short conical arms, tentac1es glands and branchial hearts into one complex on sides
long, with thick stalks, also conical, fin shorter of mantle cavity. Larvae with very short arms and
than half of ML, tentacular suckers in approx. 6 long tentac1es, usually with characteristic stalked
rows. eyes, commonly with ventrally directed rostrum,
The only described species, P. megaptera Hoyle, without photophores. Larval development protracted.
1885, is known from the Northwestem Pacifico An Larvae are elear, adults usually purple, brown or yel-
undescribed species reported for the subantarctic low, but in some genera transparent. Two subfamilies,
South Atlantic and the SE Pacifico All species pre- 14 genera. Approx. 25 species in the South Atlantic.
dominantly abyssopelagic.
Subfamily Cranchiinae
Family Cranchiidae Two (longitudinal) or four (in the form of an inverted
Slow-moving p1anktonic and semiplanktonic squids V) hyaline stripes with cartilaginous tuberc1es on ven-
of small to gigantic size. Mantle fused with head and tral mantle beginning at sites ofmantle-funnel fusiono
funnel, thin, membranous, semigelatinous or leathery, Funnel fused with head laterally. Funnel valve present
smooth or bearing small cartilaginous tubercles either (except in Leachia). Eye photophores numerous, in 1-
circumscribed in some areas or distributed through 2 arcs. One of the ventral arms hectocotylized in
most of the body. Mantle covered by gelatinous coat males. Four genera. Three genera and approx. 7 spe-
in some genera. Fins of very variable form and size, cies in the South Atlantic.
from small petaloid to large lancet-like or round, Cranchia scabra Leach, 1817 (Fig. 2.47). Mantle
length up to 60% ML. Head short and narrow. Eyes as elongate balloon, posterior end subacute in
very large, stalked and tubular in larvae and juveniles adults. All mantle surface and upper side of fins
of most genera, non-stalked and globular in adults. covered with very numerous cartilaginous tuber-
Cephalopoda 743
cles with multifid tips, formed in larvae at ML 4- inner eyeball photophores formed at ML approx. 8
5 rnm. Four hyaline stripes on ventral mantle near and 12 mm, respectively. One species in the
mantle-funnel fusion, in the form of an inverted V, Atlantic Ocean.
apparent in larvae at ML approx. 10-15 rnm. Head Leachia (Pyrgopsis) atlantica (Degner, 1925)
may retract into mantle up to eyes. Eyes not (Fig. 2.49). Mantle ofleathery consistence in juve-
stalked, even on larvae. Fins in larvae petaloid, in niles, gelatinous' in adults, almost transparent,
juveniles as a horizontal figure 8, short, longer and cylindrical in anterior and narrowly-conical in
wider in adults but not extending beyond mantle posterior half, acuminate to needle-shaped posteri-
end and not fused. Arms short. Tentacular stalks orly. On anterior-ventral side of mantle 2 parallel
bear suckers along entire length in larvae, only on straight hyaline stripes, bearing cartilaginous
club in adults. Club with 4 rows of suckers. tubercles, their length approx. 14-15% ML; with
Tentacles not autotornized in adults. Funnel valve 7-8 large multifid rosette-shaped tubercles,
presento 14 round eye photophores: 6 in a ring approx. 5 small multifid ones, and some simple
around pupil, 7+ 1 in an arc on ventral side. Large conical between the large ones. Second large
elongated photophore on ends of all arms in matur- tubercle from anterior end not displaced laterally.
ing and mature females. In males right 4th arm hec- Head narrow, with rather long "snout" (arm-crown
tocotylized and ends of 3rd arms modified (minute pillar) in larvae. Eyes in larvae and juveniles
suckers in 4 rows). Light-colored. ML up to 15 cm. stalked, with short rostrum (slipper-shaped), in
Larvae easily distinguishable by cartilaginous adults not stalked. Fins in larvae petaloid, in adults
tubercles. Outer and inner eyeball photophores transversely-oval, not extending beyond mantle.
developed in larvae at ML 15-17 and la mm, Anterior margin of fin arcuate, posterior almost
respectively. One species in the World Ocean. straight, its maximum width behind the middle.
Liocrallclzia reinhardti (Steenstrup, 1856) (Fig. Fin width 1.3-1.6 times fin length. 3rd arms long,
2.48). Mantle almost globular in larvae, oval in others rather short. Tentacles long, fixing appara-
juveniles, spindle-shaped, acuminate behind in tus stretched along stalk, club with 4 rows of suck-
adults. On anterior-ventral side of mantle 4 hya- ers, of which the 4-5 central pairs are enlarged.
line divergent stripes in the shape of an inverted V, Funnel valve absent. 6 oval eye photophores: 1
bearing cartilaginous tubercles, long (to 'mid-man- near pupil, 6 in an arc on ventral side. Large elon-
tle) in juveniles, relatively shorter in adults. gate photophore on ends of 3rd arms in maturing
Another similar strip along dorsal midline of man- and mature females. In males right 4th arm hecto-
tle over gladius, reaching anterior end of fin. Head cotylized and midportions of all other arms modi-
narrow, eyes not stalked, even in larvae, large, fied (suckers enlarged, with reduced apertures).
round in adults. Fins petaloid in larvae, resembling Squids semitransparent with only some dispersed
a horizontal figure 8 in juveniles, in adults oval or large dark chromatophores. ML up to 10 cm.
almost round, extending far beyond mantle end Larvae distinguishable by tubercles on anterior-
and fused here. Arms rather short, particularly in ventral hyaline stripes (developed at ML approx. 7
larvae and juveniles, 3rd arms longest. Tentacles mm), cylindrical mantle, and stalked eyes, but spe-
long, fixing apparatus stretched along stalk up to cies characters not distinguishable before late
its middle, club with 4 rows of suckers. Tentacles juvenile stage. First eyeball photophores deve-
autotomized in mature females. Funnel valve loped at ML <3 cm, full pattem of photophores
presento 14 round eye photophores: 6 in a ring and reduction of eyestalks at >5 cm.
around pupil, 7+ I in an arc on ventral side. Large The system of the genus Leaclzia remains unclear,
elongated photophore on ends of 3rd arms in the number of species and their distribution
maturing and mature females. In males one of 4th unknown. Supposedly 5-6 species in the South
arms hectocotylized, ends of 3rd arms modified. Atlantic, some in tropical-subtropical, others in
Color light-yellow or brown, larvae and juveniles southem subtropical and southem peripheral areas.
semitransparent. ML up to 20-25 cm. Larvae dis-
tinguishable by tubercles on hyaline stripes but not Subfamily Taoniinae
on body, developed in larvae on ventral side at ML No hyaline stripes with cartilaginous tubercles on
approx. 4-6 rnm, on dorsal side at 8 mm. Outer and ventral mantle except at sites of mantle-funnel fusion
744 Cephalopoda
(or also of mantle-head fusion) in some genera. anteriorly, reaching tip of lanceola or exceeding it
Funnel not fused with head laterally. Funnel valve by a bit, either joined there by a thin membrane or
absent (except in Megalocranchia and Egea). Eyes remaining divided by a narrow split. In larvae
with one large crescent-shaped and usually one (rare- mantle end may exceed end of lanceola. Arms
ly two) much smaller, usually rod-shaped photo- short. Suckers in 2 rows, some enlarged suckers in
phores. No hectocotylization. Ten genera. All genera distal part of 3rd arms. Tentacles long, strong, not
and approx. 17 species present in the South Atlantic. autotomized. Stalk in larvae bears 4 rows of suck-
HelicocrulIchiu pfefferi Massy, 1907 (Fig. 2.50). ers, in juveniles suckers absent in proximal third,
Mantle leathery, smooth, cup- or spindle-shaped, in 2 rows in central and in 4 in proximal part of
conical and rounded behind. No cartilaginous stalk. Central club suckers with sharp teeth.
tubercles and hyaline stripes. Head small, ventral Medial component of funnel organ with three
side completely covered by large funnel, which papillae, one central conical and two lateral spatu-
exceeds arm bases. Eyes small, elongate, with late. No funnel val ve. Two eyeball photophores, a
short stalks, in adults barely stalked, directed ante- small elongate one in concavity of a large cres-
riorly. Fins at all stages small, paddle-shaped, cent-shaped one. No arm photophores. Light co-
attached to rear sides of short diamond-shaped lored, with sparse dispersed chromatophores. ML
lanceola, extending slightly beyond mantle end up to 25 cm. Larvae and juveniles distinguishable
and joined there by a thin membrane. Arms very by fins attached only to sides of lanceola. Two or
short, 3rd longest. Suckers in 2 rows, enlarged in 3 species, one in the South Atlantic.
mid-parts of arms in males (but not in females). SUlldulops melullcholiclls Chun, 1906 (Fig. 2.52).
Tentacles long, strong, not autotomized. Suckers Mantle membranous, sac-like in larvae, in adults
stretching throughout stalk in larvae. Fixing appa- leathery and slimy, broadly cup- or barrel-shaped,
ratus in adults consisting of altemating suckers rounded behind, mantle width 40-60% ML.
and knobs, al so stretching throughout stalk. No Cartilaginous tubercles absent. Head small, in lar-
enlarged club suckers. Medial component of fun- vae with long "snout" (arm-crown pillar). Funnel
nel organ with three conical papillae. No funnel large, covering arm bases. Eyes in larvae on long
valve. Qne large eyeball photophore. No arm pho- stalks, elongate, rostrate, slightly curved (slipper-
tophores. Ends of all arms of adult males modified like); in juveniles not stalked, cylindrical; in adults
(with small suckers in many rows). Light colored, globular, directed anteriorly. Lanceola wide, dia-
oval brown or orange chromatophores form trans- mond-shaped, fiat in anterior half, bent roof-like
verse rows on mantle..sides, irregularly disposed and with longitudinal keel in posterior half. Fins in
on mid-dorsal and mid-ventral mantle. No small larvae paddle-shaped, in juveniles and adults se-
black dots. ML up to 8 cm. Larvae and juveniles mioval or kidney-shaped, attached to rear sides of
distinguishable by small, paddle-shaped fins, huge lanceola, not reaching its tip, which protrudes
funnel (noticeable in larvae at ML 2.5 mm), and between fins. Arms very short, with wide protec-
transverse rows of chromatophores on sides of tive membranes. Suckers in 2 rows, small, smooth
mantle. Three species, 2 in the South Atlantic. H. or weakly dentate. Tentacles in larvae and juve-
pfefferi may be a species complex. niles long, thin, in adults usually autotomized.
• Liguriellu podophtulmu Issel, 1908 (Fig. 2.51). Stalk in larvae with 2 rows of suckers. Central club
Mantle semigelatinous, smooth, slimy, widely suckers with small teeth. Medial component of
spindle-shaped, round behind in larvae, subacute funnel organ with central conical papilla and two
in adults. Two small conical cartilaginous tuber- wide triangular lateral fiaps. No funnel val ve. Two
cles each at site ofhead-to-mantle fusion (ML >30 eyeball photophores, a small oval one in the con-
rnm). Head small, with "snout" (arm-crown pillar) cavity of a large one. No arm photophores. All
in larvae, large in adults. Funnellarge. Eyes in lar- arms modified in males, bases wide, ends of Ist-
vae on long stalks, rostrate (i.e., with an acute ven- 3rd pairs narrow and tapering, suckers in basal
tral outgrowth), in adults not stalked, large, globu- parts of all arms enlarged, with reduced apertures.
lar. Fins in larvae (ML <15 mm) paddle-shaped, in Squid almost colorless. ML up to 11 cm. Larvae
adults semioval, attached to rear sides of diamond- and juveniles distinguishable by elongate, and
shaped lanceola, not extending onto mantle sides slightly curved eyes and shape of lanceola and fins.
Cephalopoda 745
• Teuthowellia pellllcida (Chun, 1910) Three species in Atlantic Ocean, 2 in the South
(=Megalocranchia pardus Berry, 1916; Atlantic. T. pellucida distributed around the
Anomalocranchia impennis Robson, 1924; Southern Subtropical Convergence in the Atlantic,
Megalocranchia richardsoni Del!, 1959) (Fig. southern part of the Indian Ocean and SW Pacifico
2.53). Mantle thin, membranous in larvae and T. maculafa (Leach, 1817), in the tropical East
juveniles, leathery in adults, cup-like, slightly Atlantic, differs by having only one tubercle at the
rounded behind in larvae, subacute in juveniles mantle-to-funnel fusion site, gladius tail length
and adults. Two-4 (rarely 1 or 5) smal!, conical 23-29% ML, and large club suckers with 19-26
cartilaginous tubercles at sites of mantle-to-funnel teeth.
fusion (at ML >25-30 mm, one tubercle at 10-20 Taollills pavo (Lesueur, 1821) [=Belonella belone
mm). Head smal!, short. Eyes in larvae on short (Chun, 1906); Belonella pacifica pacifica Nesis,
stalks, in juveniles semi-settled, elongate, at ML 1972; Galifellthis friluminosa Lu and Clarke,
>5 cm not stalked, globular. Eye diameter in 1974] (Fig. 2.54). Mantle thin, membranous in lar-
adults 14-18% ML. Fins petaloid in larvae, invert- vae and juveniles, leathery in adults, cylindrical in
ed-heart-shaped in juveniles, elongated-oval or anterior and conical in posterior half in larvae and
lancet-shaped in adults, fin length to 60% ML. At juveniles, spindle-shaped in adults, posteriorly
ML >25-30 mm anterior margins of fin go onto drawn into long, thin, acute tail. Mantle not nar-
mantle sides anterior to lanceola; posterior end of rowed in front of anterior fin margino No cartila-
fins do not extend beyond end of lanceola. Arms ginous tubercles at sites ofmantle-to-funnel fusiono
short, with 2 rows of suckers, 3rd arms longest. In Head smal!, short and narrow, in larva e with short
mid-parts of 2nd and particularly 3rd arms some "snout" (arm-crown pillar). Eyes in larvae on
enlarged suckers with smooth or barely dentate long, in juveniles on short stalks, in adults not
rings. Tentacles long in larvae, rather short in stalked, globular. Fins in larvae «18 mm) dia-
adults, with thick stalks, not autotomized. mond-shaped, in juveniles and adults long (30-
Tentacular stalk in specimens ML <25-30 mm 40% ML at ML >4-5 cm) and narrow, oval,
with 4 rows of smal! suckers along en tire length, stretched along mantle sides, not extending
in adults proximal third without suckers, rest with beyond end of lanceola. Arms short in larvae, long
3 oblique rows. Four rows ofsuckers on club, cen- and strong in adults, with 2 rows of suckers, 3rd
tral one somewhat larger than marginal, rings den- arms longest. Width of arm protective membranes
tate, with 26-32 teeth. Rear sides of lanceola approx. equals sucker diame ter. Large suckers in
straight, taillength from site of maximal lanceola adults with broad blunt teeth. Tentacles long and
width to gladius end 20-22% ML. Medial compo- thin in larvae, rather short and strong in adults,
nent of funnel organ with three papil!ae: one cen- with thick stalks, not autotomized. Fixing appara-
tral conical and two lateral spatulate. No funnel tus: an oval carpal group of knobs and suckers,
valve. Three eyeball photophores, a small sickle- and 2 rows of alternating knobs and suckers on
shaped one in the concavity of a large semi lunar dista12/3 of stem. Four rows of stalked suckers on
one, and the smallest elongate, near acute end of club, central much larger than marginal; their rings
sickle. Large arm photophore on ends of al! arms (beginning from ML approx. 5-6 cm) transformed
in mature females. Ends of l-2nd arms modified in into hooks with 2 (rarely one) large teeth, without
males, with 2-4 rows of specialized suckers with small side-hooks; base of ring not widened, ring
reduced apertures. Color in larvae clear, semi- opening not reduced, hook not covered by skin
transparent, with many oval dark chromatophores hood. Funnel moderately long. Medial component
forming a longitudinal stripe along dorsal mantle. of funnel organ with three conical papillae. No
Adults dark, with numerous moderately spaced funnel valve. Two eyebal! photophores, one smal!
smal! oval chromatophores. ML up to 20 cm. rod- or crescent-shaped, one large semilunar. No
Larvae may be distinguished by many oval chro- arm photophores in mature females. Ends of lst-
matophores forming longitudinal stripe along the 3rd arms modified in males, with 2 rows of abrupt-
back. In larvae first eye photophore appears at ML ly reduced suckers. Larvae clear, semitransparent,
approx. 5 mm, second at 15-20 mm, third at adults purple-violet. Epithelium easily abraded,
approx. 50 ffiffi. exposing a reticulate subcuticular pattern. ML up
746 Cephalopoda
to 54 cm. Larvae distinguished by very thin man- fusion sites in larvae ML 7, 9, 15, 18, and approx.
tle, long narrow fin, long eyestalks. Three species, 30 rnm there are 1, 2, 4, 5, 6 tubercles, respective-
2 in the South Atlantic. Iy. Club hooks begin to form at ML 55-65 mm.
Galiteuth;s glacialis (Chun, 1906) Five species, 3 in the South Atlantic. G. glacialis
(=Crystalloteuthis glacialis Chun, 1906; is one of commonest Antarctic squids, rarely found
Teuthowenia antarctica Chun, 1910; Galiteuthis northward of the Antarctic Convergence. Two
aspera Filippova, 1972) (Fig. 2.55). Mantle mem- other species have smooth mantle, no tubercles at
branous in larvae, tough and leathery in juveniles mantle-to-head fusion sites, and smooth arm suck-
and adults, semigelatinous in mature females, er rings: an Atlantic tropical-subtropical species G.
spindle-shaped. In juveniles (ML > 12-13 rnm) and armata Joubin, 1898 (2-4 small single-pointed car-
adults anterior half of mantle, dorsal head side, tilaginous tubercles at sites of mantle-to-funnel
and arm bases covered by numerous tiny skin fusion at ML >25 rnm), and a southern subtropical-
teeth, shagreen-like. At each mantle-to-funnel subantarctic species G. suhmi (Hoyle, 1885) (one
fusion site, 2 compound cartilaginous tubercles tubercle with 5-6 tips at sites of mantle-to-funnel
usually with 5-6 tips; at mantle-to-head fusion site fusion in adults).
2 short cartilaginous ridges, usually with 3 tips, Mesollychoteuthis hami/ton; Robson, 1925 (Fig.
beside anterior end of gladius. Head narrow, with 2.56). A gigantic squid. Mantle membranous in
short "snout" (arm-crown pillar) in larvae. Eyes on larvae, tough and leathery in juveniles and adults,
short stalks in larvae, not stalked at ML >20 rnm. semigelatinous in mature females, spindle-shaped
Fins in larvae petaloid, very small, later (ML 20- in juveniles and cup-shaped in adults, thick (up to
25 mm) round, then rounded diamond-shaped, and 4-5 cm), smooth. At each mantle-to-funnel fusion
in late juveniles and adults lancet-shaped. Length site 4-6 conical cartilaginous tubercles; at mantle-
in adults 35-45% ML, width 15-20% ML, anterior to-head fusion site l conical tubercle beside ante-
fin margin at site of maximum width of lanceola. rior end of gladius; all usually poorly visible in
Arms short, approx. 25% ML, in juveniles and adults. Head wide, without "snout" (arm-crown
adults (ML >25-35 mm), 3rd-4th arms longer than pillar). Eyes on short stalks in larvae, not stalked
1st-2nd. Suckers in 2 rows, rings with some blunt and globular injuveniles and adults. Fins in larvae
or acute teeth. Tentacles long in larvae (Ionger petaloid, very small, in juveniles round, in adults
than mantle at ML <10 rnm), in juveniles and broadly heart-shaped, length in adults up to 60%
adults approx. half of ML. Club short, with 4 rows ML, width somewhat less, anterior fin margin at
of suckers in larvae but in adults central rows with site of maximum width o~ lanceola. Arms long,
5-6 pairs of large acute hooks with widened base, robust, ca. 50% ML, subequal. Suckers in 2 rows,
covered by ski n hood, and small marginal suckers. with dentate rings. Some suckers on mid-section
Fixing apparatus: an oval carpal group of knobs of all arms, from 3-5 pairs on 1st to 8-10 pairs on
and suckers, and 2 rows of alternating knobs and 4th arms converted into strong hooks, covered by
suckers on distal2/3 of stalk. No arm photophores skin hood. Transformation begins at ML approx.
in mature females. Ends of 1st-2nd arms modified 45 rnm with central tooth becoming elongated,
in males bearing 2 rows of abruptly reduced suck- overgrowing others and downturned, two others
ers. Medial component of funnel organ with three remaining as small teeth beside the longest.
conical papillae. Funnel valve absent. Two photo- Tentacles not shorter than mantle. Club short, nar-
phores on eyeball, one small narrow arc-like, one row, with 4 rows of suckers in larvae, in adults
large crescent-shaped. Larvae and juveniles semi- central rows with 11-12 pairs oflarge acute hooks
transparent with dispersed red-brown chromato- with widened base, covered by skin hood, and
phores grouped in irregular transverse rows on same number of small marginal suckers. Hooks
mantle. ML up to 50 cm. Larvae identifiable by appearing at ML approx. 50 mm. Fixing appara-
short eyestalks and fin, juveniles by cartilaginous tus: an oval carpal group of knobs and suckers,
tubercles and shagreen mantle. First tubercles at and 2 rows of alternating knobs and suckers on
mantle-to-head fusion sites appear in larvae at ML distal two thirds of stalk. No arm photophores in
5-6 mm, 2nd and 3rd tips at ML 10-12 mm and mature females. Ends of all arms elongated in
approx. 25 rnm, respectively. At mantle-to-funnel mature males, ends of 2nd-3rd arms modified,
Cephalopoda 747
with 2 rows of abruptly reduced suckers. Medial photophore on ends of 3rd (or lst-3rd) arms in
component of funnel organ with three conical mature females. Ends of lst-3rd arms modified in
papillae. Funnel valve absent. Two photophores mature males, with minute suckers in 2 rows.
on eyeball, a small elongate-oval anterior and a Larvae and juveniles semitransparent and almost
large crescent-shaped posterior. Larvae semitrans- colorless. ML.up to 81 cm. Larvae identified by
parent with dispersed red-brown chromatophores, conical "snout" and long eyestalks, juveniles by
adults dark purple-brown. ML up to 200-225 cm, ink sac photophores and fins divided by tail needle.
weight up to 150 kg. Larvae identifiable by wide In larvae caudal pair of ink sac photophores deve-
spindle-shaped mantle. Tubercles at mantle-to- lops first, at ML approx. 25-30 rnm. Juveniles
head and mantle-to-funnel fusion sites appear in become adult-like at ML 40-60 rnm. Probably 3
larvae at ML approx. 15 rnm, eyes not stalked at species, one in the South Atlantic.
ML >40-45 rnm. Egea illerlllis Joubin, 1933 (=Phasmatopsis lllci-
One species, rarely found northward of the fer G. Voss, 1963) (Fig. 2.58). Mantle elongate,
Antarctic Convergence but occasionally reaching narrowly-spindle-shaped, acute behind, smooth,
approx. 40oS. membranous in larvae and leathery in adults. No
• Megalocrallchia oceallica (G. Voss, 1960) (Fig. cartilaginous tubercles. Head small, without
2.57). Mantle narrowly cup-like or semi-spindle- "snout". Eyes in larvae on short stalks, elongate, in
shaped, subacute behind, smooth, membranous in adults (>40 rnm) not stalked, globular, looking
larvae and leathery in adults, slimy. No cartilagi- anteriorly. Fins in larvae petaloid, attached to rear
nous tubercles. Head small, in larvae with conical sides of lanceola, not fused, in adults very narrow,
"snout" (arm-crown pillar). Eyes in larvae on long anterior margins extending onto mantle sides at
stalks, elongate, with rostrum, in adults not stalked, ML >20-25 rnm. Fin length approx. 30% ML.
globular. Fins in larvae petaloid, attached to rear Arms short, protective membranes narrow.
sides of lanceola, not fused, in adults egg-shaped, Suckers in 2 rows, none enlarged, rings with low
divided by tail needle. Anterior fin margins blunt teeth, no suckers with acute teeth. Tentacles
attached to anterior sides oflanceola, not extending rather long, not autotomized. Four rows ofsuckers
onto mantle sides. Fin length approx. 50% ML. along tentacular stalk in larvae, 2 in proximal and
Arms short, in adults with wide protective mem- 4 in distal part in juveniles, in adults two rows of
branes. Suckers in 2 rows, on lateral arms some alternating knobs and suckers; no carpal group of
suckers on mid-portions enlarged, their rings with suckers and knobs. Central club suckers not larger
some blunt teeth; sucker rings on distal end of arms than marginal ones, rings with 8-12 acute teeth on
in adult squids with long, wide, acute teeth on dis- distal margino Lanceola with straight rear margins,
tal margino Tentacles rather short but strong, not acute but without needle-like tai!. Funnel smal!.
autotomized. Four rows of suckers along tentacular Medial component of funnel organ with 2 low tri-
stalk in larvae, 2 on proximal and 4 on distal part angular lateral flaps. Funnel valve presento Two
in juveniles, and in adults two rows of alternating narrow C-shaped eyeball photophores, anterior
knobs and suckers; no carpal group of suckers and with hook-like bend. No ink sac photophores. A
knobs. Central club suckers somewhat larger than large photophore on ends of 3rd arms in mature
marginal ones, with 6-8 large acute teeth on distal females. Ends of all arms modified in mature
and some small ones on proximal margino males bearing minute suckers in 4 rows. Larvae
Lanceola with con cave rear margins, tapering pos- and juveniles semitransparent and almost color-
teriorly into thin needle-like tai!. Funnel smal!. less. ML up to 42 cm. Larvae identified by a nar-
Medial component of funnel organ with 2 low co- row spindle-shaped mantle and anteriorly directed
nical lateral papillae without central one. Funnel eyes. Juveniles became adult-like at ML 35-40
valve presento Two eyeball photophores, anterior mm. One species in the W orld Ocean.
ice-hockey-stick-like, strongly bent, posterior BatllOthallllla Iyrolllllla Chun, 1906 (Fig. 2.59).
elongated, narrow. Two dumb-bell-shaped photo- Mantle cylindrical, rounded behind, smooth,
phores with round lenses on ends located on ven- membranous (resembling sausage skin). No carti-
tral side of digestive gland, on enlarged ink-sac. In laginous tubercles. Head small, in larvae with very
live squids they appear as bright green spots. Large long "snout" (arrn-crown pillar). Eyes in larvae on
748 Cephalopoda
very long (in so me cases equal to ML) stalks, approx. halfway between it and web margin, thread·
elongate, with rostrum, in juveniles eyestalks like arrn ends naked. Along both sides of each arrn
shorten, in adults (> 100 mm) eyes not stalked, glo- two rows of cirri, one pair near each sucker, begin-
bose, directed lateral-anteriorly. Fins in larvae and ning closer to mouth than first suckers and extend-
adults small, padd1e-shaped, on short wide stalks, ing almost to arrn ends. Between 1st and 2nd arrns
attached not to lanceola but to mantle sides far pair of thin retractile suckerless filaments usually
from one another. Arms short, approx. 30% ML in coiled inside of special pouches on outer side of
adults. Suckers in 2 rows, none enlarged, rings web. Tentacles absent. Two large complex evertible
smooth. Tentacles very long (longer than ML), not photophores located on dorsal mantle side between
autotomized. 2 rows of suckers along tentacular bases of definitive fins and rudiments of larval ones
stalk. Central club suckers somewhat larger than (present in larvae ML 6-7 rnm), and very many
marginal ones, rings with long acute teeth on dis- small photophores dispersed on arrns, head, ventral
tal margino Lanceola reduced. Funnel smal!. mantle and fins. Two large light oval spots, suppos-
Medial component of funnel organ with 2 conical edly extraocular photoreceptors on head between
lateral papillae. No funnel valve. One large half- eyes (present in larvae ML > 1O rnm). Radula
round eyeball photophore. No ink sac photo- present, all teeth runicuspid. No ink saco Gladius
phores, no arrn photophores in mature females. 1st very thin, wide, transparent. Body exterior dark
arrns modified in mature males through most their purple-violet, arrns and web inside intensely velvet-
length, bearing minute suckers in 2-4 rowS. Larvae black. Color not variable (chromatophores are non-
and juveniles semitransparent and almost color- functional). Egg size 3.8 rnm. ML up to 13 cm.
less. ML up to 20 cm. Larvae may be identified by Larvae are almost adult-like except fins.
their cylindrical mantle, rounded behind, reduced
lanceola, very long eyestalks, "snout", and small Order Octopoda
fins. Eyeball photophores appear at ML >25-30 Suborder Incirrata
rnm. One species in the W orld Ocean. Superfamily Bolitaenoidea
Two families.
Order Vampyromorpha Family Bolitaenidae
One family. Body gelatinous, pigmented. Arrns shorter than man-
Family Vampyroteuthidae tle, 3rd pair the longest. Arrn tips not tapering. Web of
One monospecific genus in the World Ocean. moderate length. Suckers in one row. Mantle opening
Vampyroteuthis infeTllalis Chun, 1903 (Fig. 2.60). single, under head, wide. Stomach behind digestive
Mantle conical, rounded behind, semigelatinous. gland. Radula - multidentate, comb-like. Ink sac
Head fused with mantle, poorly separated from present, moderate in size. A lemon-yellow circumoral
body and arrns. Eyes very large, diameter up to bioluminescent ring around mouth in mature females
40% ML, open (not covered by corneal membrane). only. Eggs connected by their stalks in large clusters,
Mantle opening moderately wide, reaching mid- females hold the egg-mass on their arrns near the
point between eye and funne!. Funnel sunken into mouth within the web. Three genera, 3 or 4 species, 3
head tissues almost to its opening. Mantle locking- in the South Atlantic.
apparatus lacking. Pair of long paddle-shaped fins Japetella diaphalla Hoyle, 1885 (=Dorsopsis
without cartilaginous base on sides ofposterior part taningi Thore, 1949) (Fig. 2.61). Eyes large, ellip-
of mantle. In smallest larvae larval fins on very end tic, their relative length (% ML) gradually
of mantle. At ML approx. 9 rnm, pair of definitive decreasing during growth. Larger eye diameter is
fins beginning to develop anterior to larval one, at 19-27% ML at ML <20 rnm, 15-25% ML at ML
ML 20-25 rnm larva with two pairs of fins of sub- 20-60 mm, and 12-20% ML at ML >60 mm.
equallength, then definitive fins grow more rapidly Lateral and ventral surfaces of eye covered by sil-
and at ML approx. 30 rnm larval fins gradually very reflecting layer. Optic nerves short, optic
resorbed. Arms short, subequal, twice as long as ganglia near the brain (easily visible because of
mantle, with tapering ends. Web deep, approx. half transparency of tissues). Mantle opening reaching
of arm length. Suckers small, unstalked, without level of pupil. Suckers located near one another,
horny rings, in one row, beginning far from mouth, distance between them not more than one sucker
Cephalopoda 749
diameter, large, diameter averaging 6% ML. Family Amphitretidae

Suckers in middle part of 3rd right ann in adult Body gelatinous, covered by jelly coating. Eyes tele-
males enlarged, urn-shaped. Hectocotylus absent. scopic, tubular, directed upwards, their bases drawn
Maximum web depth 25-33% of length of longest together on dorsal side of head. Anns long, j oined by
ann. Funnel sunken into head tissues almost to its deep web. Ann tips not tapering. Suckers in one roW.
opening, located under anterior eye margino Mande fused with head and funnel. Mande opening
Digestive gland large, elongate-ovoid, covered by represented by two split-like or oval apertures on
silvery reflecting tissue. Gills with 8-11 lamellae per sides of head lateral to funnel. Funnel very long.
demibranch (half of gill). Egg length approx. 3 rnm. Stomach in front of digestive gland, ovary above it.
Color mostly light-brownish in juveniles, dark- lnk sac present, on posterior-dorsal side of digestive
brown, not transparent in adults. ML up to 10 cm. gland. Inner demibranch of gills reduced. No photo-
Larvae with narrow head, elliptical eyes, short arms phores. Radula multidentate, comb-like. 3rd right ann
(3-4 times shorter than ML) with 5-6 suckers (in hectocotylized in males, with narrow and very long
early larvae), wide mande and large ovoid digestive ligula and short triangular calamus. Eggs large, length
gland. Larvae almost totally covered by thick sticky approx. 5 rnm, width 2 rnm, not held by female. One
jelly. Optic nerves not detectable in early larvae or two genera and species, only one in the South
(ML <6 rnm). One species in the World Ocean. Adantic.
• Amphitretus pelagicus Hoyle, 1885 (Fig. 2.63).
• Eledollella pygmaea Verrill, 1884 (Fig. 2.62). Animal almost nonpigmented, colorless, wholly
Eyes small, elliptical, widely separated, relative covered by jelly coating overlayed by a thin epi-
length decreasing gradually from 12-17% ML at thelium with some sparse chromatophores. Only
ML <20 mm, to 11-14 % ML at ML 20-60 mm, eye lenses, ann ends and funnel tip emerge from
and 7-11 % ML at ML >60 mm. Lateral and ven- coating. Mantle elongate, narrowly-oval (in
tral surfaces of eye covered by silvery reflecting Atlantic subspecies). Anns subequal, 1.5-3 times
layer. Optic nerves long (not less than brain ML. Web depth up to 2/3 maximum ann length.
width), optic ganglia far from brain (easily visible Suckers small, widely separated. No enlarged
through transparent tissues). Mande opening suckers. Digestive gland oval-subtriangular. Tip
reaching level of pupil. Suckers not numerous, of hectocotylized ann in mature males finely
located far apart, distance between them one or pointed and curved hook-like, set off from basal
two sucker diameters, small, diameter averaging part of ann by angular swelling. ML up to 9 cm.
4% ML. Three suckers in middle part of 3rd right Larvae like miniature adults, easily distinguish-
ann enlarged in adult males, one or two very large, able by telescopic eyes and two lateral mande
urn-shaped. Hectocotylus present, the end of 3rd openmgs.
left ann modified in mature males. Maximum web One species widely distributed in the World
depth approx. 50% of longest ann length. Funnel Ocean with two subspecies: A. p. pelagiClls in the
sunken into head tissues almost to opening, loca- tropical Indo-Pacific, and A. p. thielei Robson,
ted under anterior eye margino Digestive gland 1930 in the southern subtropical and subantarctic
large, ovoid, covered by silvery reflecting tissue. Adantic.
Gills with 4-8 lamellae per demibranch (half of
gill). Egg length approx. 2 mm. Color mostly red Superfamily Octopodoidea
or reddish-brownish in juveniles, adults very light- Two families.
ly colored, almost transparento ML up to 20-25 Family Octopodidae
cm. Larvae with narrow head, with widely sepa- Bottom octopuses, some species with planktonic lar-
rated (about 3 eye widths) small, elliptical eyes; val stage. Body neither gelatinous nor transparento
anns short, subequal, with 4 suckers in early lar- Anns muscular, much longer than body, suckers in 1
vae; wide mantle and large ovoid digestive gland. or 2 rows. Web usually not longer than half maximum
Larvae almost totally covered by thick sticky jelly. ann length. Mande opening not reduced. Radula not
Optic nerves visible even in the smallest larvae. comb-like. Digestive gland in front of stomach and
One species in the W orld Ocean. gonad, ink sac (ifpresent) on its ventral side. Right or
(in a few species only) left 3rd ann hectocotylized in
750 Cephalopoda
males. Eggs small or large. Small eggs entwined with head with 2, 3 small over each eye, golden irido-
their stalks into clusters glued by female to the sub- phores around eyes. Digestive gland with 13 large
strate; large eggs glued to the substrate singly. dorsal visceral chromatophores. Large larva ready
Females watch eggs during entire incubation period to settle on bottom at 10 mm ML. Arms with >30
(up to one year). Small eggs (ca. 1.5-5.0 mm) give rise suckers. Dorsal mantle with >80, dorsal head with
to swimming planktonic larvae; whereas large eggs >50 chromatophores, ventral head with 4, ventral
(ca. 10-40 mm) produce crawling benthic juveniles. mantle and funnel as in hatchlings. Web vestigial.
Intermediate-sized eggs yield either, depending on 8-11 gilllamellae per demibranch. Egg length 2.2-
ratio egg size/female size. In some species females 2.5 mm. Adults inhabit mostly the inner shelf,
retaining egg string on arms up to hatching. Most spe- sandy and muddy bottoms.
cies with planktonic larvae belong to the subfamilies OctOPllS defilippi Vérany, 1851 (=Macrotritopus
Octopodinae (5 genera, some species with benthic Grimpe, 1922) (Fig. 2.65). Hatchlings 1.3-1.5 mm
devclopment as well), and Eledoninae (some species ML. Mantle round, head wide. Arms subequal,
of the genus Eledolle). AII members of the other two approx. 50% ML. Three primary suckers on each
./ subfamilies, Bathypolypodinae and Graneledoninae, armo At ML 1.5-1.7 mm 3rd arms becoming long-
have benthic juveniles. Planktonic larvae of bottom er (as long as ML at 2.5 mm), in later larvae much
octopuses are all rather alike, distinguished mainly by longer than others (3-4 times ML). Second arms
the distribution of chromatophores, the number of 1.5-3 times shorter than 3rd. Arms thick, their ends
suckers per arm in newboms, and the arm length ratio. in late larvae tapering and fragile. Outer surface of
It should be noted that light (red and yellow) chroma- arms with one row of dark chromatophores, plus
tophores fade shortly after fixation in formalin or one chromatophore at base of each sucker. At ML
ethyl alcohol, only the brown ones remaining. At ML approx. 3 mm dorsal mantle surface clear in mid-
2-8 mm hatchlings of all octopuses with planktonic region, 2 chromatophores in posterior area; ventral
larvae have short rounded mantle, smooth skin, cha- with 6-8 in mid-region, forming approximately
racteristically covered by bunches of short bristles transverse rows; funnel with 2+2, dorsal head with
(Koelliker's bristles), large prominent eyes, and short 8 (2+4+2), ventral head with 2, one small over
arms not joined by a web, with 3-4 primary suckers in each eye and one large at base of each arm; gold-
one row. During subsequent growth and settling on en iridophores around eyes. Digestive gland with
the bottom the mantle elongates, bristles are lost, arms 11 large dorsal visceral chromatophores. Large lar-
become long and unequal, with many suckers in l or vae ready to settle on bottom at 9-11 mm ML, rare-
2 rows, a web develops, and color becomes variable. Iy to 15 mm ML. Body elongate, tubular. Third
Planktonic stages are undescribed in many species; in arms very long, >40 chromatophores in one row.
some cases the larvae described are uncorrelated with Second and 4th arms somewhat shorter. Dorsal
their bottom-dwelling adult counterparts. Thus, mantle clear in the mid-region, with >20 chroma-
despite the large number of genera and species of tophores in the posterior area. Web vestigial.
adult octopods described for the South Atlantic, only Eleven (rarely 12) gill lamellae per demibranch.
6 species are included here. Egg length approx. 2 mm.
Larvae may be found as far north as the southem
Genus OctOPlIS Cuvier, 1797 slope of Georges Bank. Adults live on the entire
Body muscular. Arm suckers in two rows. Ink sac shelf and uppermost slope, on sandy and muddy
present. Right 3rd aml hectocotylized in males, with bottoms. There is also a closely related species
well-differentiated ligula and calamus. No color rings (still undescribed) which inhabits the tropical and
or stripes on mantle and arms in adults. subtropical Indo-West Pacific, the Arabian Sea
OctOPllS bllrryi G. Voss, 1950 (Fig. 2.64). and South Africa to Hawaii and Tonga.
Hatchlings approx. 1.5 mm ML. Mantle round, • OctOPllS macrop"s Risso, 1826 (Fig. 2.66).
head narrow. 4 primary suckers on each armo Outer Hatchlings undescribed. Early larvae (ML 2.4 mm)
surface of arms with 8-10 dark chromatophores in with short round body, wide head, short subequal
2 rows. Dorsal mantle surface clear, ventral with arms, each with 4-6 chromatophores in one row.
26-30 chromatophores (6 across mantle), funnel Three primary suckers on each arm, 1st smaller
with 2+2, dorsal head with 10 (2+4+4), ventral than 2nd-3rd. Suckers arranged in 2 rows begin-
Cephalopoda 751
ning from 4th armo Web rather deep. Dorsal man- Genus Scaellrglls Troschel, 1857
tle with approx. 12 chromatophores on anterior (2 As De/opus, but 3rd left arm hectocotylized in males.
lateral diagonal patches), and >20 on posterior Scaeurgus unicirrhus (delle Chiaje, 1830) (Fig.
area; ventral mantle with >35 chromatophores on 2.68). Hatchlings approx. 2.0 mm ML. Body squat,
anterior area and along midline (2 longitudinal globose, head. wide, eyes prominent. Arms short,
rows), and >20 on posterior area; dorsal head with subequal, 4 primary suckers on each, gradually
14, funnel with 4. Subsequently the body becomes decreasing in size from 1st to 4th. Outer surface of
ovoid and somewhat pointed, the head narrows, arms with 3-4 dark chromatophores in one row.
arm ends get pointed, 1st arm grows somewhat Dorsal mantle surface clear, posterior part with
longer than others, then 2nd (evident only in latest patch of about 10, ventral mantle with 35-40 chro-
larvae), outer side of arms first with one, then with matophores on middle and posterior parts, funnel
two rows of chromatophores, number of chroma- with 2+2, dorsal head with 14 (2+4+4+4), ventral
tophores on dorsal mantle to >30, on dorsal head head with 2 large, 1 each at arm bases. lridophores
to > 18; ventral mantle in mid-region with many around eyes not visible. Digestive gland with
chromatophores. ML of planktonic larvae up to approx. 20 large dorsal visceral chromatophores.
15-20 mm; 9-13 lamellae per demibranch. Egg Web vestigial. Seven gilllamellae per demibranch
size 1.8-2.0 mm. in early larva, 11-14 in adults. Egg length 2.2-2.5
A shallow-water species, more common at is1ands mm.
than near continents, inhabit mostly coral reefs. Lower sublittoral and upper bathyal. A characte-
Even juvenile octopuses may rise at night to the ristic species of oceanic islands and shallow-sum-
surface. There are several very similar species in mit seamounts.
the Indo- West Pacifico
OctOP"S vlIIgaris Cuvier, 1797 (Fig. 2.67). Genus Eledolle Leach, 1817
Hatchlings 1.5-2.0 mm ML. Mantle elongate, co- Body muscular. Arm suckers in one row. Ink sac
nica1, head wide, eyes protruding. Arms short, sub- present. Right 3rd arm hectocotylized in males, hecto-
equal, with tapering tips. Three primary suckers on cotylus very short, ligula and calamus almost undif-
each arm, then suckers abruptly becoming much ferentiated. Ends of all other arms in males with mo-
smaller, rearranged into 2 rows. Outer surface of dified suckers.
arms with 2 dark chromatophores in one row. Eledolle Iligra (Hoy le, 1910) (Fig. 2.69). Only a
Dorsal mantle surface clear in mid-region, with 2 larva ML 6.9 mm described. Mantle egg-shaped;
small chromatophores in posterior part, ventral sur- arms subequal, somewhat shorter than ML; arm
face with 12-18 scattered large chromatophores, chromatophores in one row (> 16); dorsal mantle
funnel with 2+2, dorsal head with 8-10 (2+4+4), densely covered with large chromatophores (>50;
ventral head with 2, later 4, one small over each 8-9 across mantle), ventral mantle with >50 small
eye, blue-green iridophores around eyes. Digestive ones, dorsal head with 6 (4+2) chromatophores,
gland with 6-8 large dorsal visceral chromato- ventral head with some small ones, funnel with 3;
phores. Large larvae ready to settle on bottom at 12 visceral chromatophores on digestive gland;
7.5-11, usually 8-10 mm ML, settling normally at silvery iridophores around eyes. Ten lamellae per
the age of 7-8 weeks. Arms subequal, somewhat demibranch in adults. Egg size 3.5 mm.
longer than ML, with approx. 25-30 suckers. Outer
surface with 20-30 chromatophores in 1 row prox- Family Vitreledonellidae
imally and 2 rows distally. Dorsal mantle in mid- One monospecific genus in the World Ocean.
region clear, ventral mantle with 25-30 chromato- Vitreledollella richardi Joubin, 1918 (Fig. 2.70).
phores, dorsal and ventral head and funnel as in Body gelatinous, transparento Mantle elongate-oval,
hatchlings. Web vestigial. Eight-l0 gill lamellae rounded behind. Head small, narrow, fused with
per demibranch. Egg length 1.8-2.5 mm. mantle, poorly differentiated from arms. Eyes
Predominantly OIl, inner shelf, but also on outer small, subrectangular, directed laterally. Optic gan-
shelf and uppermost bathyal. Very common and glia far from brain. Mantle opening single, wide,
commercially important. A very close species reaching approx. level of eyes. Funnel of moderate
dwells in waters off Japan, China and Korea. length, not fused with head. Arms Ist-3rd pair sub-
752 Cephalopoda
equal, in juveniles nearly as long as ML, in adults bom, have only 7 arms. Hectocotylus detached dur-'
2-3 times ML, 4th slightly shorter. Arm tips tape- ing mating. Adults Iight-colored on the outside, body
ring. Web very deep, in juveniles almost reaching covered by small chromatophores. Inside of web
arm ends, in adults approx. 40-50% maximum arm intensely purple-brown. Eggs stalked, incubated in
length. Suckers small, not numerous, um-shaped, large cluster on female's arm crown. Larvae gelati-
sunken into arm tissues, widely spaced at the base, nous and semitransparent. Animals distinguished by
converging and slightly enlarged beyond web mar- short mantle of semigelatinous consistency, very
gin, edge-to-edge at the very tips. Digestive gland large eyes, and characteristic arrangement of suckers.
long, slender, cigar-shaped, pointed; stomach in Larval males also distinguishable by showing only
front of it. Ink sac on dorsal side of digestive gland. seven arms. ML up to 50 cm, totallength to 2 m.
Inner demibranch of gills lacking. No photophores. Qne species. Larvae Iive at middepths, adults at
Radula heterodont, not comb-like, rachidial teeth both mid-depth and near the bottom on the slope,
multidentate, others unidentate. End of 3rd left:arm seamounts, and submarine rises.
hectocotylized in adult males, shorter than 3rd right
armo Female ovoviviparous, eggs (many hundreds, Family Ocythoidae
approx. 4 mm in length) incubated in widened dis- Qne monospecific genus in the W orld Qcean.
tal oviducts. Adults nearly colorless, with sparse, OcytllOe tllberclllata Rafinesque, 1814 (Fig.
small, red-violet chromatophores. ML up to 11 cm, 2.72). Body firm, muscular. Mantle in larvae,
totallength to 45 cm. Newbom larvae (ML approx. juveniles, and males widely-ovoid, in females nar-
2.2 mm) almost transparent, encased in jelly coat- rower. Mantle fused with head, connected with
ing, easily identifiable by small deeply pigmented funnel by well-developed, strong connective appa-
eyes, short arms (half ML), widely set basal suck- ratus. Ventral mantle in females covered by set of
ers, and very long (nearly equal to ML), narrow, skin tubercles connected by ski n ridges in a reti-
pointed digestive gland. culate pattem, other parts of body and entire body
in males smooth. Head wide, weakly separated
Superfamily Argonautoidea from mantle and arms. Eyes large. Mantle opening
Four families. wide, reaching level of upper eye margino Funnel
Family AIIoposidae long, conical, extending beyond bases of 4th arms,
Qne monospecific genus in the World Qcean. not sunken into tissues. Skin pores (water pores)
AlloposlIs mollis Verrill, 1880 (=Haliphron atlanti- beside funnel near bases of both 4th arms. Arms
el/S Steenstrup, 1859) (Fig. 2.71). Mantle short, se- short in larvae, long and thin in juveniles and
migelatinous, ovoid, width equal to length. Mantle adults, 1st and 4th subequal, much longer than 2nd
fused with head, connected with funnel by wealdy and 3rd. Web and arm membranes almost absent.
developed connective apparatus. Head wide, weakly
separated from mantle and arms. Eyes very large, Suckers
one row, small,
others numerous,
in 2 rows. 3Ink
primary ores small,
sac present, photo-in
diameter 35-40% ML. Mantle opening wide, reach- phores absent. Males significantly smaller than
ing level of upper eye margino Funnel exceeding females. Third right arm hectocotylized, in larvae
bases of 4th arms, sunken into tissues almost to tipo and juvenile males appearing as a stalked saco
Arms thick, round in section, approximately equal to Hectocotylus develops inside sac, released at
ML in larvae and juveniles, 2-4 times ML in adults; maturity, detached during mating. Females have a
1st pair by far the longest, 4th the shortest. Web very unique buoyancy apparatus inside the dorsal man-
deep, in juveniles to 85%, in adults 40-65% maxi- tle resembling a fish swimming bladder. 19-20
muro arm length. Suckers nuroerous, small, in one lamellae per demibranch. Females ovoviviparous,
row, set far apart inside web, in two rows and crowd- incubate eggs (>100,000, 1.8 mm in length, 1.0
ed from web edge to arm ends. Ink sac present, pho- mm in width) up to hatching. In adults upper side
tophores absent. Digestive gland large, in front of ofbody deep-blue, lower side rose-violet with sil-
stomach. Males smaller than females, but not very metallic iridescence. Larvae distinguishable
dwarfed. In males 3rd right arm fully modified, by long 1st and 4th, short 2nd and 3rd arms, males
developed inside a special sac under skin, thus it also by characteristic appearance of hectocotylus.
appears as though immature males, even the new- Reticulate pattem on ventral mantle evident in
CephaJopoda 753
juveniles at ML approx. 10 mm, water pores at all row along dorsal (inner) side of2nd arms; one row
stages. Ventral mantle in larvae clear. ML of of elongate yellowish (in daylight) bodies along
females up to 31 cm, of males to 3 cm. margin of 1st arm film is luminescent. Larvae, juve-
Adults live near the surface, males commonly in niles, and males semitransparent with dispersed
empty tests of salps or doliolids. small chromatophores. Males and juveniles (to ML
approx. 15 mm) usually with pieces ofburning ten-
Family Tremoctopodidae tacles of Physalia, the Portuguese Man O'War in
One genus in the World Ocean. the 1st-2nd arm suckers, as an additional method of
TremoctopllS violacells delle Chiaje, 1830 (Fig. defense. Hatchlings ML <2 mm. Larvae without
2.73). Mantle semiovoid or subtriangular in juve- jelly coating but head and arms enveloped by a
niles and males, ovoid in adult females, fused with cuff-shaped membrane. Some suckers on the mid-
head and connected with funnel by connective die of 1st amlS enlarged (last ones in one row),
apparatus. Body firm, muscular, mantle surface suckers on 1st arms larger than those on 2nd-4th.
smooth. Head wide, eyes large. Mantle opening Web on 1st arms begins to form at ML approx. 10
wide, its upper edges reaching level of upper eye mm. One row of chromatophores along outer side
of 1st arms in larvae, 2 in juveniles. Larvae distin-
margino funnel long, its anterior end reaching ante-
rior eye margino Funnel free. Two pairs of skin guishable by long 1st arms. ML of females up to
pores (water pores) on the head, dorsally between 30-45 cm, of males to approx. 1.5 cm.
bases of 1st and 2nd arms and ventrally beside fun- Two subspecies, one in the Atlantic (T. v. viola-
nel opening. Arms in larvae short, 1st longest, 2nd ceus). Other, T. v. gracilis (Souleyet, 1852), in the
and 4th shorter, 3rd very short. Web very short in Indian and Pacific oceans. Juveniles rise to the
larvae, deep between 1st and 2nd arms in juveniles, surface at night; adult males and females Iive con-
these two arm pairs bordered by a film narrowing stantly near the surface.
toward the arm ends. In inmlature females ends of Tremoctop"S gelatlls Thomas, 1977 (Fig. 2.74).
1st arms tapering into thin whip, their film widened Body gelatinous, so transparent that a newspaper
into wide ragged film-like tissue on 1st arms, in can be read through the body of a living female.
adults gradually tom off, becoming shorter than on Eyes very large. Ink sac and water pores reduced.
2nd arms. Web vestigial between 3rd and 4th arms. Eight-ll lamellae per demibranch in females, 7-8
Three primary suckers in one row, enlarging from in males. Adult females pale orange-red. Other fea-
1st to 3rd, then 2 rows of small suckers on all arms, tures as in T. violaceus. Larvae encased by a gela-
but on ends of 1st arms suckers arranged in 1 row tinous coating ovelayed by a very thin epidermis
and very far aparto Ink sac present, photophores with Koelliker' bristles. Ends of 1st arms emerge
present on 1st arm film in one subspecies (T. v. from the coating, some 2 or 3 suckers on the mid-
gracilis). Males dwarfed, their 3rd right arm whol- dIe of 1st amlS are enlarged, thereafter splitting
Iy modified and enclosed in sac under skin, thus into into 2 rows. ML of females up to 33 cm.
appearing that immature males (even newboms)
have 7 arms. During maturation hectocotylus everts Family Argonautidae
and becomes longer than specimen itself, detaches One genus in the W orld Ocean.
during mating. Ends of 1st and 2nd arms in males Argollallta argo Linné, 1758 (Fig. 2.75). Mantle in
thin, without film. 13-16 lamellae per demibranch larvae and males semiovoid, in females egg-like,
in females, 9-11 in males. Eggs (3.0-3.6 by 1.8-2.1 caudal end curved slightly upwards. Mantle fused
mm) connected with stalks, egg cluster (> 100,000) with head and connected with funnel by well-
bom by female on its 1st arms. Color of females developed connective apparatus conspicuous even
bright-blue to wine-red from above, silver tinged in larvae. Body firm, muscular, mantle surface
with rose or Iight gold from below. Females of the smooth. Head wide, eyes large. Mantle opening
Atlantic subspecies T. v. violaceus with row of ocel- reaching level of anterior eye margino Funnel free.
lar dark spots along ventral (outer) side of film of Skin pores (water pores) lacking. Arms in larvae
1st arms; females ofthe Indo-Pacific T. v. gracilis short, 1st longest, others subequal, arm ends taper-
with these spots in 2 rows (or in a double row) ing. In males arms 1st, 2nd and 4th subequal, 3rd
along ventral (outer) side of 1st arm film, and in 1 shortest; in females arms long, 1st much longer
754 Cephalopoda
than others, 4th and 2nd shorter, 3rd are the short- Males with many small chromatophores.
esto Fourth arms 10-20% longer than 2nd, 1.5-2 Hatchlings ML 0.85-0.9 rnm. Early larvae (up to 3
times the length of 3rd. In maturing and mature rnm) covered by jelly coat with only arm ends pro-
females distal section of 1st arms with very exten- truding. Eyes directed somewhat antero-ventrally.
sible wide membranous ski n flap which secretes Larvae clear, with some (23-24 in hatchlings)
and holds the shell (secondary shell). Web vesti- sparsely distributed large chromatophores. Flaps
gial. Three primary suckers in one row (two first on 1st arms detectable at ML 3, developed at >4
larger than 3rd), then 2 rows of small suckers on all rnm. ML of females up to 30 cm, of males to 1 cm.
arms. Ink sac present, photophores absent. Twenty Dwells in coastal areas, rare in the open ocean.
eight lamellae per demibranch in females. Female Juveniles and adult females rise to the surface du-
builds characteristic thin, fragile calcareous shell ring the night. Another Atlantic species, A. hians
where it lives and incubates eggs. Female stays Solander, 1786, is also tropical-subtropical, but
within shell with 1st arms extended along the sides oceanic; with small shell (up to 10 cm), usually
of the shell and other arms retracted inside. Shell smooth, not flattened sideways, keel wide (10-15%
formation begins at ML approx. 5-7 rnm; during of shell length), ribs not numerous «35), set far
subsequent life, beginning from approx. 8 rnm, apart, nodules brown. Fourth arms much shorter
shell is enlarged, in large females reaching 20-30 than 2nd-3rd, length in large females approx. 60%
cm in length. Shell elongated, flattened sideways, of 2nd. Egg size 0.7-0.8 mm. 20 lamellae per de-
with very narrow (4-5%, rarely 6% of shelllength) mibranch. A third species, A. nodosa Solander,
keel and thin, slightly bent, outside earlets; with 1786, dwells in the tropical Indo- West Pacific; it
many (>50 in large shells) smooth ribs, drawn was often recorded off Namibia and South Africa
close to one another, cornmonly bifurcated and (Sánchez, 1988; Laptikhovsky, 1989; Villanueva
ending in small, acute, triangular nodules on the and Sánchez, 1993); it is a large species with wide
keel. In young females the shell is small, ribs are keel (l 0-15% of shelllength), and ribs (up to 30-40)
few, set far apart, keel wide, up to 10% shell beset by chains of separate tubercles or nodules.
length. Eggs very small (0.6-0.8 rnm), laid and
incubated in rear part of shell between caudal end
of mantle and posterior shell wall. Newbom larvae Suggested readings (see also Table 2)
liberated by female probably every night. Female
begins to reproduce shortly after building shell and Boletzky S. von. 1974. The "larvae" of Cephalopoda:
continues to spawn eggs during entire life. Males a review. Thalassia Jugoslavica, 10:45-76. General
dwarfed, their 3rd left arm totally modified and description o/larval stages o/ cephalopods.
enclosed in sac on stalk (from ML approx. 1.5 Boyle P. (ed.). 1983-1987. Cephalopod life cycles.
rnm), giving the impression that irnmature males V.l: Species accounts; V.2: Comparative reviews.
(even newboms) have 7 arms. During maturation Academic Press, London, V.1, pp. 1-475; V.2, pp.
the hectocotylus everts and becomes much longer 1-441. Descriptions o/ paralarvae o/some repre-
than male itself, detaches during mating. Part ofthe sentative species.
hectocotylus with sperm crawls into female's man- Clarke M.R. (ed.). 1996. The role of cephalopods in
tle cavity and remains there; male dies after mating the world's oceans. Phl Trans. Roy. Soco
and is probably devoured by female. Up to 3 hec- London, B, 351:979-1112. General data on the
tocotyli may be found in one female. Color of rale o/ cephalopods in the World Ocean as COI1SU-
females purple-blue to wine-red from above, light mers and as prey.
from below; extended 1st arm flap purple-red. Clarke M.R. 1966. A review of the systematics and
Fixed females red-brown. Shell porcelain-white, ecology of oceanic squids. Advances Mar. Biol.,
early section and nodules on the keel dark brown. 4:91-300. A general review o/ reproduction and
-
Table 2. Seleeled referenees on Soulh Allanlie eephalopods and lheir early slages of developmenl (papers devoled lo isolaled genera and spe-
eies are nol ineluded).
Cephalopoda 755
southern important
open ocean
Tropical
subtropical
Namibia
and
South
Africa and
Angola,
Subantarctic
and
Antarctic
Malvinas
(=Falkland)
Argentina an Geographic areas
I Brazil, Uruguj'
•
Adam (1952) • • - +
• • •
+ • • + •
f
756 Cephalopoda
distribution of all oceanic squids and Spirula system. Zool. Zhurnal, 75(3):335-349. A disclls-
Á710wnto date. sion of new propositions for the systematics of
Fioroni P. 1978. Cephalopoda, Tintenfische. ceplzalopods (in Russian).
Morphogenese der Tiere, Lief. 2, G5-1. G. Fischer, Roper C.F.E., Sweeney M.J., Nauen C.E. 1984.
Stuttgart, pp. 1-181. A general description of Cephalopods of the World. FAO Species .....
cephalopod ontogeny. Catalogue, 3, FAO Fisheries Synopsis, 125(3): 1-
Guerra Sierra A. 1992. Mollusca Cephalopoda. In 127. Illustrated glossary ofimportant terms.
"Fauna Iberica, 1", Mus. Nac. Cienc. Natur., Roper C.F.E., Voss G.L. 1983. Guidelines for taxo-
Madrid, pp. 1-327. A good introductory work. A nomic descriptions of cephalopod species. Mem.
brief review of cephalopod systematics, with keys Nat. Mus. Victoria, 38:48-63. Lists of taxonomi-
in Spanish. Glossary of important terms (withollt cally important characters of variolls cephalopod
illustrations). orders.
Mangold K. (ed.). 1989. Céphalopodes. In "Traité de Roper C.F.E., Young R.E., Voss G.L. 1969. An illus-
Zoologie", 5(4) (P.P. Grassé, ed.), Masson, Paris, trated key to the families of the order Teuthoidea.
pp. 1-804. Most important review work, inel. Smithsonian Contrib. Zool., 13: 1-32. Illustrations
cephalopod development, ontogeny, taxonomy and of representative genera and species with keys to
distribution. the family leve!.
Naef A. 1921-1923. Die Cephalopoden. Fauna Flora Sweeney M.J., Roper C.F.E., Mangold K., Clarke
Golfes Neapel, 35(1): 1-863. Descriptions and fig- M.R., Boletzky S. von (eds.). 1992. "Larval" and
ures of larval and juvenile stages of most juvenile cephalopods: A manual for their identifi-
Mediterranean cephalopods. cation. Smithsonian Contrib. Zool., 513:1-282.
Nesis K.N. 1985. Oceanic cephalopods: distribution, Illllstrated glossaty of important terms, keys of
life forms, evolution. Nauka, Moskva, pp. 1-287. late larval andjuvenile cepha!opods to family and
A list of all oceanic cephalopods known to date, genus leve/s, and illustrations oIthe larvae oImost
with distributional data and list of genera of those species having a larval stage.
inhabiting the shelves (in Russian). Young R.E. 1972. The systematics and areal distribu-
Nesis K.N. 1987. Cephalopods of the World. TFH tion of pelagic cephalopods from the seas off
Publications, Neptune City, pp. 1-351. Keys for all Southern California. Smithsonian Contrib. Zool.,
oceanic cephalopods of the world, and shelf 97: 1-159. A good, well-illustrated description of
cephalopods ofmany areas, inel. South Atlantic. oceanic cephalopods and larva! stages of gonatid
Nesis K.N. 1996. The system of recent cephalopods: .§!!lIids off California.
The critical analysis of newly proposed improve-
ments using the characters of the reproductive
Cephalopoda 757
A B e D E
(EJ)
changes during growth
F
(A 9)
Spirula spirula
Rondeletiola minor
Heteroteuthis
(Heteroteuthis)
dispar
E
(maturing ó)
B
(L)
outer ~
/... eye
:;.,--- funnel
yolk sac ~~ arms
statocyst~~
Pho<oPho,e~~)<)
(Em
_Abefore~inner ~Ik~c F G
hatching) (Aá, dv) (Aó, vv)
758 Cephalopoda
el.
A
vv (Em before dv
hatching)
t'~~
~B
changes in fin
e
Loligo vulgaris
outllne from (A9)

larva to adult
Lolliguncula brevis
e
(Al
Lycoteuthis diadema
Cephalopoda 759
Selenoteuth·
s..cmt,lIans ,s
(Paraenoploteuthis)
fnoploteuthis
anapsis
dv vv
760 Cephalopoda
A
(L)
D
e (A 9,
(J) mantle photophores
not figured)
8) Abralia (Pygmabralia) redfieldi
A
(J; ML=13 mm)
....
•
(Te)
A
(J; ML=13 mm) B
(Aó)
G Abraliopsis (Abraliopsis) pfefferi

Abraliopsis (Micrabralia)
gilchristi
Cepha1opoda 761
A
(U
D
(A, T)
•
B(U
Ancistrocheirus lesueurii
E (U
E
(Imm9)
F
(Ieft eyeballl
Pyroteuthis
margaritifera
F
(Aól
•
762 Cephalopoda
A
(L,
ML=4.5 mm)
e
(U
B
e
o, ML=8 mm)
(L, ML=6 mm)
Octopoteuthis Sp.
F
(Sa)
A
(U E
O)
• •
e
(2nd arm
E
photophore)
(A, schematized)
B G G G
(U (TC, (TC, (TC,
ML=5 mm) ML=11.5 mm) ML=32 mm)
Taningia danae Onykia carriboea

• Cephalopoda
F
763
'-':'(" (skin sculpture)
B
E
(L, Moroteuthis sp., 0, TC)
ML= 6 mm)
A
(L, Moroteuthis sp., A
ML= 6 mm) (L, ML=5 mm)
D
e 0, ML=14.7 mm)
0, ML= 14.7 mm)
Moroteuthis robsoni
\* ML=8 mm
B
(L) it ML=5.5 mm
e ML=3 mm
(L)
G
(sehematie
representation of
arms and tentacles)
0'
.,.,
., e
I
•
., (A)
,
:>
F B
(L TC) (J)
~ (L, ~C)
Onychoteuthis banksii Kondakovia longimana

764 Cephalopoda
.
•• 'y;
J"\
)1~
..
E
(L, Te;
ML= 10 mm)
B
(L)
G
(L, TC;
ML= 18 mm)
I I i.
(A, TC)
A
(L, ML= 5.7 mm) e G. fabrícií, a North Atlantic
(J) counterpart of G. antarctícus
'- - - - - - - - - - - - - ~~~- - - - - - - - - - - - - - - - - --
Gonatus (Gonatus) antarcticus
~ñ
(~IU
B
A
(L, ML=3 mm)
o
(J, ML=12 mm)
(L, ML=2.5 mm) e
(J, ML=6 mm)
•
o
(J)
F
(A, SV)
E
(J) E
Chtenopteryx sicula • (A)

Bathyteuthis abyssicola
Cephalopoda 765
.¡¡
A B
(EL, vv) (EL, dv)
Histioteuthis sp.
F e
0, ML=6 mm)
(photophores
reft eye)around
Histioteuthis re versa o E
(Imm)
0, ML=9 mm)
.,.- ...•
",:'~;~:-::.
o F
(A) (photophores around
Histioteuthis bonnellii left and right eye)
766 Cephalopoda
11\
~~\~
l'
¡
i~
A "
(L, ML= 1Omm) ~ t
• ..
• I
e
0, Te) Architeuthis dux
A ;1
O) ;~
dv
}~~
ft \&~\ B
•
i~\
•
0, Neoteuthis sp.)
1 ~~~ \
! ~;?- ! E
(A, T)
e D e
0, T) (j, T)
(A, Neoteuthis sp.)
Alluroteuthis antarcticus Neoteuthis thielei

Cephalopoda 767
--------------------------------------------------------------:
D
A B (U
(L) (U
____________________
~~~T~:~~~~~~~iCS _
J
(L, TC,
ML=lO mm)
....
.. K
U, TC)
F
H U)
(L, Te. from the Scotia Sea
ML=6.S mm)
Brachioteuthis riisei
768 Cephalopoda
(r\ 'l-f.T
,rt.~~
~,x-~
i'. • ,~\
t;
~d1· :,.;:)••. .
~,/dv
A { i' • .• ~
(L, ML=l.4 mm) tt ~"!
¡:. • Iii
¡, j ,,' t
(. ., j t'
,; ~.'. ,:i \.
;;, ~ .e 04.. '.'1
i••."';..•••.,'••"~.y·I e) "~o
.
'.
E
dv vv.1'\
~,.,...." " .)J.• .•/'./'\.,.J
'.~I..'
'. O. ML=ll mm)
~. ;~:.
¡f • :.,\\i \I !
'1:.1
:~'\\!~.. ~/
~.r;;~
"\ ...w~.,'7;
~~ ~
~
'~I ~\\J~~,
~'j
! , •
/" ¡) "
't ;'. J .
:, ','"i
¡'.'.. i j
:'; . •
..'.-\.
.~' '. j'} ; • I t •
f 1,,' ~ , • • ~~
~
~.'
¡
\.' ', ••• J
j
\11 • ~
\ ft ••• '~'.'
, •• ~ f; .• t '.
"
,l~":··"'···)
' '
, • ~ dv vv
~.."~ r
\... \:,V )
\. ".'l
..•...
, :..~J..: - ....,.~\j..."..,-
/. eoindetii
I l' ,
e
-
(L, ML=6 mm)
. . I~i.~
. .
';~"""";'%
~;l
. '<1 '.
.
§,¡
~:).
¡;<',
l·
.
fr.~· ~
f' 11 ". ~ 1 ..•... i /:',/1
¡. • .\ i• il \
~.. .i ., .•.
, '\': :ji ~f
.> . ~
..
-i{: -~i'
f"· .... P1
.
~ l. \ . '1
'1 •
r. ," ':.1
.' , ",
' •• ':
'1'·
•
",
y," ":r
vv .J .••
,,~11lf\'" ' .,t.' '~
:b:.~~,
•.
i.1
dv ¡
'r;j:'
vv .
\~.
( ,·,1 ) .~.~
.. :(~
,
~::..;:.:
'lli/'
:~..;~ ... \/ ".,.,.,,~>
t;@\,
t," . W' ';¡' D ' ,1.;: G o.
. "::1Te)'.', ~r;-:I'
..:;'.•·f ..
(L. ML=8 mm) o, ML=30 mm)
O , TC)
J: ,~~
~.; .•.•
~~
:.
I/Iex argentinus /. illeeebrosus

Cephalopoda 769
••
A B
(U (U
e
(U
Todarodes angolensis
Ornithoteuthis antillarum
B
(U
G
(U
O. volatilis
770 Cephalopoda
o F
(E]) (9, vv)
•
(L)
G
(TC)
~ A
E (L)
(Te, carpal part)
• o
(A,9)
Sthenoteuthis pteropus
B
(L, mantle opened
lo show 2 gul
pholophores)
e
(L, manlle opened
lo pholophore)
show one gul
Cephalopoda 771
"::',
D F
(L, manlle opened " .
lo show gul
(A,9,
small dispersed
pholophore) skin pholophores
indicaled)
•
(L, manlle
opened
show gullo G
pholopnore) (A,9)
fucleoteuthis luminosa
A e
(L, ML=2 mm) (L, ML=3.7 mm)
F
Ul
E
(posllarva, manlle
opened lo show gUI
pholophore)
D
(posllarva, manlle
opened lo show gUI
phOlophore)
Hyaloteuthis pelagica
772 Cephalopoda
~~@~@G)@~~@~ ~@
~(ó)(J@) @~(!)~~ ~ ~(6)
A
(detai I of part of an egg mass)
e
(L)
B
(egg with embryo)
.. '
D E
(L, ML=3 mm) (L, ML=4 mm)
F
U, ML=6 mm)
G
U, ML=10 mm)
J K
(funnel locking (mantle locking
cartilage) cartilage)
H
U, ML=18 mm) Thysanoteuthis rhombus
• Cephalopoda 773
A
(j, ML=24 mm)
~
.; o !)
o
Cl
•}U
D E
J
(skin papillae, (Te, ML=60 mm)

ML=20 mm) (ski n papi Ilae,
ML=24 mm)
e
(A)
00 G
(club sucker,
ML=20 mm)
H
(club sucker,
ML=24 mm)
F I
(skin papillae, (club sucker,
ML=60 mm) ML=60 mm)
• B
0, manlle)
e
(Lj, ML=51 mm)
dv
A
vv
0, ML=30 mm)
Batoteuthis skolops
(TC)
D
Pholidoteuthis
boschmai
774 Cephalopoda
c~ .•.
{"O' ""._- :-
.•.
·c....
-.""
i, F
(funnel and
'1: mantle cartilages)
A
(L)
• c~~f
E
(ink sac photophore)
Cycloteuthis sirven ti
E
(funnel and
mantle carti lages)
e D
u W ~ro
~
~ {J} Discoteuthis discus
Cephalopoda 775
A
(L, ML=4 mml
'C
(L, ML=17 mm)
B
(L, ML=7 mm)
F
(Sal
E
(transitional stage,
o notice ink sac
(L, ML=35 mm) photophores)
G
(funnel cartilage)
H • I
(A, club suckersl
(L, T)
Chiroteuthis veranyi
776 Cephalopoda
-+-
dv
vv
A
(L, ML=4 mm)
dv
B e
(L, ML=7 mm)
(L, ML=17 mm)
D
(L, ML=35 mm)
'!J
I~,
1,
(VIV <,
l'·:"
t 0 F
(L, Fe)
E
(L, TC)
o
H
(j, lai I broken off)
Grimalditeuthis bonplandi
Cephalopoda 777
B
I (L, eye with photoph
, M . (Mastigopsis) ore)
d. hJorti
e
(L, arms)
M. (MfastigopSiS)
c , hJorti
H
U, small body
photophores
Indlcated)
(L ,ML=4.6
F mm)
/
>( .:~,/
,'\ .. ~.
.. ~
- - :. . . ~ {:::::.-
""F:
~.,; .:,
.
A
(L, ML=6 mm)
E
M L=20
(LL,mm)
~ I
i·:;~·
:.;·
M. (Mfastigopsis)
c. hJorti M. (Mastigoteuthl' s) sp.' . (H SV'
notlce lar ~ '
on eyelidg photophore
~ear eyesinus)
D
(L, Te) .
M. (Mastigopsis) M. (Mastlgoteuth )
.. . . ..... IS gnmaldll
d. hJorti
o
Mastivoteuth' IS sp.
•
778 Cephalopoda
l~
~:
dv vv
B e
(L, ML=7 mm) (Sa)
A
o E
(A) (3rd arm)
(L, ML=7 mm)
/oubínífeufhís porfíer;
l~
't:'~
...
~t~;
,1.,;
['Lo,
1~
i:.~~
.
r;-
;:
. "-.l
-:1
.'....•;...~
•
A
(L, vv,
ML=10.5 mm)
:-,.
E B
F G 0, TC) e (L, dv,
(L, Te) ML=10.5 mm)
(4th arm) (T) Promachofeufhís Sp.
Cephalopoda 779
A
(L, ML=5 mm,
anterior part of body, sv)
e
(cartilaginous tubercle)
F
B (A,é)
(L, ML=ll mm)
Cranchia scabra
A
(EL, H)
e o
(LL) (J)
'•• 1,-
G
(eye) _.----
-~ "-
--'_.' F
E
(A,9vv)
(A,9dv)
Liocranchia reinhardti
(AS 'f) (1)
O J
l!:J!JUl!/ll!
(S!Sd09JÁd) l!!t¡:Jl!íJ7
(WW 9 =lW '13)
V
(WW lE=lW '11)
9
Bpodo¡tn.¡d;}:) 08L
Cephalopoda 781
E
(Iarval eye)
/
ír
F
\\ (adult eye)
dv vv \
B
(L)
G
Liguriella podophtalma (mantle-funnel fusion)
8f
V'M~8mm)
I
(A, eye) vv
,~ H
',~.J.\
~-.: U, eye,
ML=25 mm)
vv
':-' , i
, í f
'i #
j,,,,,,~/
-... ¡;:-
e
(L, H)
'1 .-,... -.
~~ \yo~\j)
( i h:, \-: J
E
B (Aó)
(L, ML=12 mm) Sandalops melanc h o1"
ICUS (funnel organ)
(WW SIl=lW 'aÁa) (WW L=lW 'aÁa)
~ :l
(WW LS=lW 'f)
O
(WW SIl=lW 'es)
3
(WW OL=lW '1)
8
(WW a=lW '1)
:) (WW L=lW '1)
V
BpodoIBqd;}:) Z8L
Cephalopoda 783
A
(L, ML=5 mm)
C
(L, ML=16 mm)
B
(L, ML=10 mm)
e
(Sa) o
A (L, ML=24 mm)
(L, ML=2 cm)
D
(eye)
(rings of large
cluD suckers)
Taoniuspavo
•
G
(L, Te,
ML=26 mm)
(L, ML=51 mm)

J
K H
(tubercles
in the (tubercles in the I 0, Te,
mantle-to-funnel mantle-to-head ML=84 mm)
(A, TC)
fusion sites)
fusion sites) Galiteuthis glacialis
•
784 Cephalopoda
'h
.....
.
..:.:..:.
....
IfI/'¡:' "\ ,-:¡~\
:/:>?: \,
,
f:····:··J~f~t'
/:.~'.
A
(L)
(
"
:' .~
e ~,~
u, arms)
:.J í·~:'.,')
:r 'ir" ..>
•
1\ 'tg./
Jp/
t
i::: ~ B ~
lo:, , D
';;: U)',', Mesonychoteuthis hamiltoni (A, «, eyes damaged)
W W. / ,:\
~.::
','
"'"
¡:::
"
" '¡
A ~ /. 11
t~:·
~.-~
:,:., .....
\~\ >"!
Ul,,~.t
~~
""-~t.,.,
\!I¡,~t /l].,q,'7
! ~'~.:: ,.,.' :;:'.";.;;.•:;.
': ••, ;;:¡:
E
F
(eye in 2 positions)
(digestive
with gland
photopnores)
e
A (A)
(L, ML=17 mm) B
(j, ML=54 mm) Megalocranchia oceanica
• Cephalopoda 785
A B e
(L, ML=7 mm) (L, ML=16 mm) (L, ML=22 mm) D
(L, ML=32 mm)
E
O, ML=48 mm)
I J K F
H (L) (L) O) L (Aó)
leye photophores)
(funnel organ
and valve)
(Al Egea inermis
i,
!
, J
,\.
~ .•. ol., • .-./
:l.
/
A B
e
(L)
(L) 1L) F
(L, eye)
~_.....
--==3€X=
G
lA, eyel
Bathothauma Iyromma
786 Cephalopoda
f
A
(L)
'1
B
(L)
o
Iíght spots (L)
aoove eyes
large
photophore
F
(Aó, dv)
G
(Aó,sv)
• H
(Aó,
arms inside)
Vampyroteuthis infernalis
Cephalopoda 787
:;----
\
......:...!i
.' !
·:~t)\\r!91~;'}} e
(L without vv
A dv
(L with mueous mueous eoating)
B
eoating) (L without
dv mueous eo at'ing) vv
E
(A)
]apetella diaph ana
e D
A (countour of
(L)
submature9)
Eled onella pygmaea (mature ó)
•
788 Cephalopoda
e
(eyes) D
(eye)
B
A (contour of a
(U submature ó)
Amphitretus pelagicus
A B
(HI, ML=1.5 mm) (hatchling, brown
(chromatophores contracted,
open circles- yellow, c1osed- brown) chromatophores
ML=1.5 mm)
expanded
• e
(hatchling,
ML=lSmm)arms
Octopus burryi
D
(LL, ML=4.5 mm)
Cephalopoda 789
e
(hatchling,
3rd arm)
• (A
F
á)
Octopus defilippi
E
(LL, dv)
E
(LL, VV)
A
(EL)
Octopus macropus
790 Cephalopoda
A B e
(Em immediatly beíore hatchling,
in 3 positions)
E
(L, ML=3.8 mm)
1)
h'J'
O
..
~
f.ff
la &' "
G
,> .:1I
. I
'\
•
,
I
I
~V:-'.<'
J
Q
.
•
, ,.•.....
i\;:. 1 G
(arm)
o H
F (L, ML=2 mm) (recently settled J)
(L, ML=6 mm)
~~~
.•..
• Octopus vulgaris
I
(AÓ)
Cephalopoda 791
dv
A
(L, ML=2 mm)
sv vv e
(arm) D
(A él
B
(L, ML=2.5 mm) Scaeurgus unicirrhus
•
A
(L, ML=6.9 mm, vv)
B
(L, ML=6.9 mm, dv)
Eledone nigra
•
792 Cephalopoda
~
)
A
(newborn L) B
(older L,
schematized, showing
brain and suckers)
~~ '
./
\ ',-.-
/
(
o
(ó ,contour)
Vitre/edonella richardi
:¡
;,.
\~\fi
•.. ...~
~. .l~
"4:~~4-~
;.' ,
,.'",. !
" .. e
~IWi !Ir
".;" ,.,;,:-:~, ."'. .~
• A
(L)
Alloposus mollis
(J
B
9) e
(Aó)
(Aówith opened
o
sac of hectocotylus)
Cephalopoda 793
-.---
•
.".~.:"
...
'">.~.-
Ocythoe tuberculata
B
A (L newborn Ó) F
(Saó)
(L newborn 9)
H
(Imm 9with 1st-2nd
between expanded film
arms)
G
e (Imm 9with contracted fill'l1
between 1st-2nd arms)
(O,
:¡: tentacles
notice on of
pieces arms
Phrsa'ia
e-E: developmental stages of 9

Tremoctopus violaceus
794 Cephalopoda
• Argonauta hians
(JmmÓ)
B
dv
(A<jl)
e
l\
/1
'\, f . 1'1' '~ Tremoctopus ge/atus
A
l' .\ \ (ó, vv) / a:1'] \
B
If{'
f¡, '"
/ \ /,
)-'\
'" y,,.
\
" 11,(,'
'\
I .'!: .• ~,i ( "'""í . 'i\"",.,

:~J~
~¡1~ ) \ ;' ! I lWiIJ-':i>,,¡~:~\
r- , \',,~_.~-'
'\
A, ~ ~"
! ,. ~(':,-'",~'
¡II itl• ~---', ",'-"\i( \\¡ l'(1P
/ -I \... ,/'' • ,
'.'. ,
'~\J) /
J
{\, '~-!y'
11,
\,; ~
<.t ",. "l~ ,~
~~ ~I/
~"\'-,J'( {.:' /"-.:.,:]
r' ._.~)
" \
I
\'l. " : l' rf'J··~~ .
r~1\
" \ y\ \, ir '!' :;¡, 1
~,.'"
i' "'~----.,(:"
/ \ I laj';, . 'n". "1-
,j ~I ~
" J'
,i
i·
~
_~
~il
', ¡
'\\
I ~S\,.
:',7-'-;."','\ \ \\'1 I :'".'
'", >1 j h
I \', c,;... 1/
(
'l.lf
\l' ''''',."
\ '1','r:'Y, ,":,' I
I '\\'
1\
I '.'.\
I I \\'
J'I / I~\\:", ,
~
",,,)!
., I ',J
\ jsv
':\ ,i!
'
'
..,
~\' 1',' •. ~I!'N
',~-,,-";/;l;
";'.r,~"",
..••
-s..•:,
\l'
.J
1\ \
I~,.
',,,,:!,
j \~.;
'~~,.~
".'
.' ,:~
..-
/.'
.-
,-------/ --.."
dV"{'gtíl.'f
o
\', .----~/
':-/-;;/r' "..;,;;¿. .
t. j.:t:;:0~,%,\.
','Sc-..'/ e
(ó, vv)
(LL<jl)
(O~d~)
:¡: ~i~'~~Z~
.~,~ ,- ..
~'\'F!j..,-
•:~:.tit.~fi~ '~!~~~~~;~'~)f:>G',
~" ..;,~ d~'~'~'-'Q/.J!J~
""'~'1,J"
H
rJ:- LL:iJ\l.
'-~'<~~"'~~~~S;~~'"<:~:<~~~~~"~Y'
Í,",.-'tJ./I
'Ii:!/'!.11/1/1
"\'\ \ \ '\"\~""""~
\"'"\\\\" \\\\ ' \'~-I

¡: \ \ \. " i l\'i \\ \ \ \"
!"¡r,'f"".~_,,._~-,,:. ... \\~
"'~ • '.'-.
F
". ,
p
~{j¡!!,lil!I! I\ ,¡ ,i, \\" \, \\ ',o. ,;~" (A9in the shell, E
Argonauta argo
--~L..L!lr:i\ii,\\\\\\~\-v_ '\).... .•..--~,J......l,..L-4,_."'.............. arm sliberated)
U?)
Cephalopoda 795
•••
Fig. 2. Illustrations of cephalopod species. Abbreviations:
A: adult
dv: dorsal view
EJ: early juvenile
EL: early larva
Em: embryo
FC: funnel cartilage
fv: frontal view
H: head
Imm: irnmature
J: juvenile
L: larva
LL: late larva
MC: mantle cartilage
ML: mantle length
PHl: posthatchling
Sa: subadul t
sv: side view
T: tentacle
TC: tentacular club
vv: ventral view
Figure sources: 18 (part): from Dubinina (1980); 21 (part), 43 (part): from Nesis (1973); 31 (part): from Nesis (1979a); 10 (part), 15 (part), 16
(part), 18 (part), 19 (part), 26 (part), 43 (part), 45 (part): from Nesis (1982); 46: from Nesis (1984), after Roper and Young (1968); 1-9, 12-14,
16 (part), 17,20,21 (part), 22, 23, 24 (part), 25, 27, 28, 29, 30H-K, 32-35, 36 (part), 37, 38 (part), 39, 40, 42, 44E, H, 1, 46-48,50-52,54 (part),
55 (part), 58-60, 61 (part), 62, 63, 65 (part), 66, 67, 70-73, 74 (part, as A/lopslIs mo/lis), 75: from Nesis (1984), after various sources; 56, 57
(part): from Nesis (1984), after Voss (1980); 41: from Nesis (1984), after Young and Roper (1969); 65 (part): from Nesis 'and Nikitina (1981);
38 (part): from Nesis and Nikitina (1990); 11: from Riddell (1985); 31 (part): from Roper et al. (1984); 53, 57 (part): from Sweeney et al. (1992),
after Voss (1985); 10 (part), 15 (part), 16 (part), 18 (part), 19 (part), 24 (part), 26 (part), 36 (part), 44A-D, 49 (part), 54 (part), 55 (part), 61
(part), 64, 68, 69, 74 (part); from Sweeney et al. (1992); 30A-G: from Vidal (1994); 49 (part): from Voss (1980); 43 (part), 44F-G, 45 (part):
from Young (1992).
Ctenopoda and Onychopoda (=Cladocera)
Takashi Onbé
Introduction Reproduction and Iife history
Of the approximately 600 species of so-called Since the reproductive biology of marine cladocerans
Cladocera (Schram, 1986; Korovchinsky, 1996) only has be en reviewed extensively by Egloff el al. (1997),
8 species are marine, and Bosmina marilima occurs only some basic features will be mentioned here. The
exclusively in the Baltic, the largest brackish water life cycle of marine cladocerans, as in freshwater
system ofthe world. Fryer (1987a, b) abandoned the forms, consists of alternating parthenogenetic and
order Cladocera as a taxonomic unit, and assigned its gamogenetic generations. However, because of a lack
3 superfamilies and 1 suborder to 4 separate orders: of data from control!ed laboratory studies, most of the
Ctenopoda, Anomopoda, Onychopoda and Haplo- biological factors in population growth remain to be
poda. This system is adopted here (see Taxonomy determined. Although in field populations gamoge-
below). These are epiplanktonic animals, seasonal!y netic individuals usually occur irnmediately before
abundant in coastal, continental shelf, and oceanic population maxima, and coincide with decreasing par-
waters of temperate to tropical regions, with a few thenogenetic reproduction (typically shown by
species also in colder arctic seas (Dolgopolskaya, decreasing brood size) (cf. Bainbridge, 1958; Onbé,
1958; Della Croce, 1964, 1974; Mordukhai- 1968, 1974; Fofonoff,1994), the important factors or
Boltovskoi and Rivier, 1987). As in their freshwater combination of factors triggering the transition from
counterparts, the !ife cycle of marine cladocerans is parthenogenesis to gamo gene sis in marine environ-
heterogonous, characterized by an alternation of par- ments remain unknown. However, various biological,
thenogenetic and gamogenetic generations. In favor- chemical, and physical processes, such as overcrowd-
able environments they reproduce parthenogenetical- ing (interactions among individuals), scarcity of food
Iy in the absence of males. By virtue of extremely supp!ies, and !ight-temperature combinations may be
high reproductive potential in parthenogenetic gener- associated with the initiation of gamogenesis (see
ations, populations often increase explosively. During Egloff el al., 1997).
such phases populations are usually composed exclu-
sively of females bearing parthenogenetic embryos RamÍrez and Pérez Seijas (1985) described some
(2n). When densities reach their peak, fecundity ofthe aspects of the reproductive biology of Evadne nord-
parthenogenetic females usually declines, and among manni and Podon inlermedius in Argentine waters.
the still prevalent parthenogenetic females, males and They noted a general trend in the decrease in the aver-
females producing haploid gametes (n) appear, giving age size of parthenogenetic females, and showed a
rise to resting eggs (2n) by gamogenesis (bisexual distinct seasonal decrease in brood size before gamo-
reproduction). genesls occurs.
Most studies of marine cladocerans have involved Pedogenesis, a special kind of parthenogenesis, has
mid-Iatitudinal northern hemisphere waters. By con- been known for Evadne nordmanni since the turn of
trast, comparatively little is known of the southem the century (Kuttner, 1911). This is a phenomenon in
hemisphere, including the South Atlantic, though al! which embryos developing in a mother's brood pouch
eight marine species have been recorded there. Marine come to have their own eggs in their brood space (see
cladocerans in the South Atlantic to 1980 have been Fig. 6E, F). Subsequent studies revealed that such a
reviewed by Ramírez (1981a). Egloff el al. (1997, unique pattern of reproduction is prevalent throughout
Table 4 therein) recently summarized studies on abun- the marine and Caspian Podonidae (Bainbridge, 1958;
dance cycles of neritic and oceanic cladocerans. Gieskes, 1970; Mordukhai-Boltovskoi and Rivier,
1971; Onbé, 1974). This phenomenon was also indi-
cated by RamÍrez and Pérez Seijas (1985) in South
South Atlalltic Zooplallkton.

edited by D. Boltol'skoy, pp. 797-813
1999 Backhuys ~ublishers, Leiden, The Netherlands
798 Ctenopoda and Onychopoda (=Cladocera)
Atlantic Evadne nordmanni. Although they did not ni and Pseudevadne tergestina there was no clear-cut
describe it, their Fig. 13g shows a "late" stage embryo, diel rhythmicity in feeding throughout a 24 hr. cycle.
dissected from a mother animal, bearing a few pedo- It should be noted, however, that food of animal ori-
genetic eggs (or embryos) in its brood space. Pleopis gin, if any, might be largely underes!imated in such
schmackeri was thought to be non-pedogenetic (Onbé, experiments.
1983; Da Rocha, 1985), until Kim and Onbé (1989a)
pro ved the opposite. A parthenogenetic female of Although several aspects of the feeding ecology of
Evadne spinifera from coastal waters of southem marine cladocerans have be en investigated (see
Brazil studied for this review contains pedogenetic detailed review in Egloff et al., 1997), an adequate
embryos, the first such report for this species (Fig. 6E, understanding of their role in trophodynamic pathways
F). Apparently, this is an excellent strategy for these of the pelagic realm requires much additional work.
animals enabling them to adapt to ever-changing envi-
ronments by maximizing their reproductive potential Prospects for future work
for efficient resource exploitation.
Because of the notorious difficulty in rearing them in
In inland and marine waters of the northem hemi- captivity, owing mainly to attachment to the water
sphere, podonid species show a distinct diel cycle in surface film, particularly in podonid species, or to our
development of embryos and release of neonates lack of knowledge of their food preferences, the pop-
(Rivier, 1969; Mordukhai-Boltovskoi and Rivier, ulation dynamics of marine cladocerans and their pro-
1971; Onbé, 1974, 1977, 1978a; Bryan, 1979; Mullin duction in the local planktonic communities have
and Onbé, 1992). As a rule, Evadne nordmanni received little attention. Recent success in laboratory
(Onbé, 1974) and Pseudevadne tergestina (Onbé, cultures of some species, e.g. Penilia avirostris
1974; Bryan 1979; Mullin and Onbé 1992) liberate (PaffenhOfer and Orcutt, 1986; Tumer et al., 1988)
their neonates only in total darkness around midnight. and Pleopis polyphemoides (Fofonoff, 1994), promis-
In the South Atlantic, Perotti (1988) obtained a simi- es future elucidation of the population dynamics of
lar result for Evadne nordmanni in the laboratory. field populations (see Egloff et al., 1997, for review).
Food and feeding habits Whether South Atlantic marine cladocerans differing
morphologically (e.g. in SEM images) or biochemical-
The endopods on the thoracic limbs of Penilia aviros- ly (e.g. by DNA analysis) from those of other oceans
tris have minute, finely arranged setae capable of represent isolated populations remains unknown. In
retaining particles as small as 2 11m (Tumer et al., view of recent re-assessments of cosmopolitanism in
1988). Recent research indicates that Penilia must freshwater cladocerans (Frey, 1982, 1987), detailed
play an important role in microbial food webs, grazing morphological and biochemical surveys of marine cla-
on bacteria and minute protists (Tumer et al., 1988), docerans aimed at species concepts and possible geo-
and SEM observations show diatom frustules and graphic (local) variants are apparently needed.
dinoflagellates in the gut (Kim et al., 1989). Marine
podonids prey on minute microplankton of both ani- No attempts have hitherto been made to explore the
mal and plant <;>rigin(Jagger et al., 1988; Kim et al., distribution and abundance of the resting eggs of
1989, 1993; Kleppel et al., 1988). Some authors sug- marine cladocerans in sediments of the South
gest that they even suck body fluids of larger animal s Atlantic. From previous findings in temperate waters
(Mordukhai-Boltovskoi and Rivier 1971; Fofonoff, in the northem hemisphere (see Onbé, 1991, for
1994), which are grasped by powerful thoracic limbs review), it has become increasingly evident that the
specifically modified for raptorial behavior. annual life cycle of marine cladocerans is initiated
fram hatching of resting eggs deposited on the sea-
By measuring in-situ gut pigment concentrations in floor. The seasonal distribution and abundance of
time series experiments, Uye and Onbé (1993) dem- resting eggs, in relation to their planktonic popula-
onstrated a clear diel feeding rhythm in Penilia avi- tions, must be known to estimate the dependence of
rostris and Podon leuckarti: both exhibiting elevated cladocerans on these benthic eggs to resume their
night-time gut-pigment contents. In Evadne nordman- planktonic life. Resting egg distributions may al so be
Ctenopoda and Onychopoda (=Cladocera) 799
useful for predicting what species can establish plank- For quantitative estimation of resting eggs, multiple
tonic populations, or estimating whether the eggs can undisturbed sediment samples from at least 6-10 cm
establish seed banks in local environments (see deep should be carefully collected with a gravity
Marcus, 1996, for copepods, and references therein). corer. Samples are then sieved through a 100 11m
mesh. If necessary, ultrasonic treatment can facilitate
sieving by dispersing particles evenly. The sieved res-
Methods idue is transferred to centrifuge tubes with a dense
solution of sugar (1 kg sucrose in 1 liter water) and
ColIection and preservation of plankton samples centrifuged at 3000 rpm for 5 mino Eggs are then
skimmed from the surface of the fluido Examinations
Methods of collection and preservations should fol- show that almost all eggs present in a sediment sam-
low those described in detail for zooplankton by pie can be retrieved after two centrifugations without
Omori and Ikeda (1984). A plankton net with a mesh apparent loss of egg viability (Onbé, 1978c).
size as fine as 100 11mis recornmended for collecting
marine cladocerans. Conventional zooplankton nets Measurement
with meshes of 200-330 11mor more are too coarse to
reta in the small, soft-bodied cladocerans, in particular A conventional scheme adopted by Onbé (1974), part-
the neonates. Because of rapid population growth, and Iy modified from Baker (1938), Bainbridge (1958), and
sudden disappearance from local planktonic cornmu- Onbé (1968), can be used for measurements of body
nities (resulting largely from altemating parthenoge- size ofthe three principal morphotypes (Fig. 1). During
netic and gamogenetic generations), quantification of embryonic development, the total length of the mother
seasonal population fluctuations requires intensive increases greatly with increasing size of embryos in its
spatial and temporal collecting. At least weekly inter- brood pouch, in particular in podonids. Thus, the body
vals are necessary for assessment of such population length, or morphological length as defined by Baker
changes. Estuarine species, such as Pleopis polyphe- (1938), is the recornmended standard.
moides, should be sought in neritic, coastal, and
inland waters of low salinity (Della Croce and
Angelino, 1987). Live specimens for laboratory Geographic and vertical distribution
experiments can be collected by nets with large cod-
end buckets, like the ones developed by Reeve (1981) The worldwide distribution of the ctenopod Penilia
or its modified versions. avirostris has been reviewed by Della Croce (1964,
1974), and that of 6 of the 7 marine podonid species
For preservation 70% alcohol should be used, as for (excluding Pleopis schmackeri) by Mordukhai-
other planktonic crustaceans (Omori and Ikeda, 1984). Boltovskoi and Rivier (1987).
However, cladocerans may also be preserved in buf-
fered formalin (2-5%) for at least a couple decades, Western South Atlantic
adequate for species identifications (Della Croce and
Angelino, 1987; Onbé, personal observations). Cladocerans off Brazil (around 18-25°S) were first
reported by Gibitz (1922) (Penilia avirostris, Evadne
ColIection and sorting of benthic resting eggs spinifera and Pseudevadne tergestina).
Since the discovery of abundant cladoceran resting Valentin et al. (1986) carried out a thorough 6-day (23-
eggs in sea floor sediments (Purasjoki, 1945, 1958; 29 Nov. 1975) study on the spatio-temporal micro-dis-
Onbé, 1972, 1973, 1974, 1978b), resting eggs ha ve tribution of plankton at a fixed station in the Cabo Frio
been identified for six species: Penilia avirostris, region near Rio de Janeiro (ca. 23°S) during upwelling
Evadne nordmanni, Pseudevadne tergestina, Podon of Central Atlantic waters. They found that Pleopis
leuckarti, Podon intermedius and Pleopis polyphe- polyphemoides (maximum densities ca. 500 ind. m-3)
moides (see Onbé, 1985, 1991). The following proce- and Pseudevadne tefgestina (up to 600 ind. m-3) were
dures have been used successfully for extracting rest- closely associated with warm, less saline surface waters
ing eggs from sediments (Onbé, 1978c, 1985, 1991): ofhigh chlorophyll content. Another less abundant spe-
\\' 1\l,0/,,-r\
,v .
.. O~\e{\~ ...
\~'oo \.
_____ total length (TU ----\

"
Penilia
:_~----- TL -: , TL----~
1
BL
y
:""
: ~ ~81. ~~'- :'./ ~
~
Evadne or
(j
Pseudevadne
~ ~.:
,/ Podon or Pleopis
Fig, 1. Measurements in the three main morphotypes of marine cladocerans (modified from Onbé, 1974; Egloff el al., 1997).
cies, Penilia avirostris (up to ea. 100 ind. m-3), Shelf Water/Coastal Water (temperature variable,
occurred only sporadically throughout the period sur- >35.0WlO).
Penilia avirostris showed no definite pattem.
veyed. Valentin et al. (1987) found high abundances In general, c1adocerans avoided waters under oceanic
(max. 45% of total zooplankton numbers) of Penilia influences (Tropical Water, >20°C, 36%0). Da Rocha
avirostris both in March and in June in their 4 surface (1985) recorded the first South Atlantic occurrence of
zooplankton surveys in coastal waters between Cabo the Indo-Pacific species Pleopis schmackeri, near the
Frio and Río de Janeiro in June, October 1984, January mouth of Rio Una do Prelado, southem Brazil, along
1985, and March 1986. Evadne spp. (probably also with Penilia avirostris, Pseudevadne tergestina and
inc1uding Pseltdevadne tergestina) were also found in Evadne spinifera. Pleopis schmackeri was subse-
significant numbers in March (max. 8%). quently collected off the southem Brazilian coast
(Resgalla and Montú, 1993). The specimen used for
At a station off Santos (ea. 24°S), Da Rocha (1982) Fig. 5D, E was collected in coastal waters of southem
found 4 species, in order of decreasing abundance: Brazil by Charrid Resgalla. Apparently, the high-tem-
Penilia avirostris, Pseudevadne tergestina, Evadne perature, high-salinity preference of this species,
spinifera and Podon intermedius. The last 3 showed already noted in the northwestem Pacific (Onbé,
distinct preferences for four local water masses: 1983; Kim and Onbé, 1989a, b) also applies here.
Podon intermedilts was most abundant in Subtropical
Water (lO-20°C, 35-36%0), Pseudevadne tergestina in Resgalla and Montú (1993) studied the winter
ShelfWater (>20°C, 35-36%0), and Evadne spinifera in (September 1988) and surnmer (February 1990) varia-
tions of cladocerans collected from 5 discrete layers February 1979 in shelf waters off Mar del Plata,
(0-25,25-50, 50-100, 100-200 and 200-500 m) at 37 Argentina in some detai1. Twenty-six (of the 200)
stations during two cruises in southem Brazilian shelf samples collected in September through January con-
waters. In the winter cruise, 3 species, Pleopis poly- tained cladocerans. Of the 6 species recorded, Podon
phemoides, Evadne nordmanni and Podon intermedi- infermedius was .the most abundant throughout the
us were identified, whereas 5 species (Penilia aviros- period, extending from coastal to middle shelfwaters,
tris, Pseudevadne tergestina, Pleopis polyphemoides, followed by Evadne nordmanni, which was much less
Pleopis schmackeri and Evadne spinifera) occurred in abundant. Pleopis polyphemoides occurred chiefly in
summer. Interestingly, cladoceran den sities were high- September near Mar del Plata. Penilia avirosfris and
er in winter (max. 3316 ind. m-3) than in summer Pseudevadne tergestina (not reported but illustrated
(max. 1056 ind. m-3). Several species showed clear in their Fig. 2) were found only in summer (January)
affinities with certain hydrographic conditions, thus at the northem end of the area surveyed (38°S); and
serving as indicators ofthe corresponding water mass- Podon leuckarti, a first record for the region, occurred
es. Pleopis polyphemoides and Evadne nordmanni in spring (September-November) south of 41 oS.
occurred in cold coastal waters and waters influenced
by the subantarctic Malvinas (=Falkland) Current Eastern South Atlantic
«13°C and <33%0);Penilia avirostris and Pseudevadne
tergestina in tropical shelf waters (salinity around 35- Gibitz (1922) recorded Penilia avirostris in the Gulf
3(jlÁlo~
Pleopis schmackeri in coastal warm, high salin- of Guinea (BOS, 12°E), Pleopis polyphemoides and
ity «36%0) waters; and Evadne spinifera in tropical Evadne nordmanni off southem Africa (18-20 to
oceanic waters (Brazil Current) of highest tempera- 35°S, 12°E) and near Cape Town (35°S, 18°E), and
tures and salinities (24°C and -35%0) (see Fig. 11 in Evadne spinifera off southem Africa (18°S, 100E).
"General biological features of the South Atlantic" in
this volume). Seguin (1970, fide Raymont, 1983), in his seasonal
study of zooplankton off Abidjan, Cóte d'Ivoire,
Based on 54 surface zooplankton samples collected in where a warm season from October to June altemates
spring (November, 1986) from the Argentine- with a colder upwelling period from June to October,
Uruguayan Common Fishing Zone (offthe Río de La observed moderately abundant cladocerans in
Plata estuary), Femández Aráoz et al. (1991) record- September and January, although they were never as
ed Evadne nordmanni, Pleopis polyphemoides, Podon important as copepods.
infermedius and Penilia avirOSfris in coastal and shelf
waters influenced by Río de la Plata runoff. Bainbridge (1972) studied the zooplankton off Lagos,
Nigeria, where the hydrographic conditions are stable
Penilia avirostris, Pseudevadne fergestina, Evadne over most of the year, with a warm uppermost layer
nordmanni, Podon intermedius andJor Pleopis poly- over the continental shelf disturbed in some years by
hemoides were recorded off Argentina (Buenos Aires upwelling during August and September. Only in .
Province to the Beagle strait) by Rammner (1933), September, at the end ofthe upwelling period, did cla-
Olivier (1954, 1962), and Ramírez and De Vreese docerans briefiy dominate copepods with high num-
(1974). Off Buenos Aires, Pleopis polyphemoides and bers of Penilia avirostris and Pseudevadne tergestina.
Evadne nordmanni increase substantially during the
austral summer, while Penilia avirostris seems to be In his account of the worldwide distribution of zoo-
restricted to this season, when the SST is around 19- plankton, Raymont (1983) reviewed the distribution
20°C (Ramírez and De Vreese, 1974). This species of cladocerans along the west coast of Africa. Della
(a long with several other zooplankters) is probably an Croce and Angelino (1968-69) and Angelino and
indicator of the sporadic influence of subtropical Della Croce (1975) reported the year-round presence
Brazil Current waters on the northem Argentine shelf of Penilia avirostris in Agulhas Bank waters off Cape
(Ramírez, 1977). Town and its neighboring lagoon. On the Agulhas
Bank, Della Croce and Venugopal (1972) also record-
Ramírez and Pérez Seijas (1985) studied cladoceran ed Penilia avirostris and Pseudevadne tergestina
distribution and reproduction between May 1978 and (September-March). In January, Evadne spinifera and
"
GUlA
(Guiana
Current
Coastal)
GUIN
(Guinea
Current
Coa sta 1)
SATl BENG
(South Atlantic (Benguela
Tropical Gyre)
Coastal)
I Cunent
EAFR
~(East
Africa
FKLD Coastal)
(Southwest ~
Atlantic
Continental GUlA BRAZ FKLD SATL EAFR BENG GUIN
Shelf)
Podon
Penilia leuckarti
avirostris I J.
t.~
• ••
l.. •..
Podon intermedius ~
Pleopis polyphemoides+
Pleopis schmackeri
Evadne nordmanni l.
t f •
Evadne spinifera
Pseudevadne tergestina
-+
r. •
+. --:~~
••
~ ~.--: ~.
•
Fig.2. Recorded distribution ofmarine cladoceran species in Longhurst's (1995) Soulh Allanlic biogeochemical provinces.
Pleopis polyphemoides were found in associatlOn 1971 b), Indian Ocean (Della Croce and Venugopal,
with Pseudevadne tergestina. An abundance of 1972), eastern Pacific (Longhurst and Seibert, 1972),
Penilia avirostris was also documented by De Decker northwestern Pacific (Kim and Onbé, 1989b, 1995),
(1975) in the Agulhas Bank, where it outnumbered all and the Gulf of Mexico and Caribbean Sea (Della
other zooplankters, especially in surnmer. Croce and Angelino, 1987). However, little is yet
known of the oceanic distribution of cladocerans in
In surnmary, cladocerans have been recorded so far in the South Atlantic (Fig. 2).
the South Atlantic mainly from coastal shelf waters,
which fall within the following "biogeochemical Some species may be transported offshore into the
provinces" of Longhurst (1995): Guiana Current open ocean by currents. For example, parthenogenet-
Coastal, Brazil Current Coastal, Southwest Atlantic ic females of Pseudevadne tergestina collected far off
Continental Shelf, East African Coastal, Benguela the coast of central Japan contained only a few or fre-
Current Coastal, and Guinea Current Coastal. Only quently no embryos in their brood pouch, thus sug-
two species have so far been recorded around the gesting that they represent expatriate populations
periphery of the western part of the South Atlantic (Kim, 1989; Kim and Onbé, unpublished).
Tropical Gyre (Fig. 2). Expatriation may also affect cladoceran distributions
on both sides ofthe South Atlantic, asjudged from the
Oceanic distribution surface circulation systems (e.g., Peterson and
Stramma, 1991) and biogeochemical provinces
Although marine cladocerans were earlier believed to (Longhurst, 1995). Boltovskoy (1986) reviewed expa-
be restricted to neritic areas, it is increasingly evident triation events in the transition area of the southwest-
that some species range into the open ocean as well. em Atlantic, where displaced subantarctic and sub-
The following oceanic occurrences have been report- tropical organisms comprise 70-80 to >90% of the
ed: North Atlantic (Wiborg, 1955; Gieskes, 1970, overall zooplankton assemblage. Samples collected in
this area are reported to have high proportions of juve- no distinct pattem of diel vertical migration through-
niles (Pteropoda), sexually irnmature (Chaetognatha), out 24-72 hr periods (Checkley et al., 1992; Mullin
and deformed (Foraminifera) specimens, suggesting and Onbé, 1992; Onbé and Ikeda, 1995).
that the expatriates do not reproduce there.
Species collected off Santos, southem Brazil, seem to
Vertical distribution and rnigrations contradict the observations above (Da Rocha, 1982).
Pselldevadne tergestina and Evadne spinifera
Marine c1adocerans dwell mainly in the upper layers, occurred predominantly in surface layers at all times,
particularly in stratified environments (Lochhead, whereas Penilia avirostris was found to undergo a
1954; Bainbridge, 1958; Trégouboff, 1963; Onbé, "normal" diel vertical migration, residing in midwater
1974, 1978a; Onbé and Ikeda, 1995). Da Rocha (1982) during the day and migrating upward to the surface at
reported on the vertical distribution of Penilia aviros- night. Podon intermedills was always between 20 and
tris, Pseudevadne tergestina, Evadlle spinifera and 50 m, below the thermocline. Differences in local
Podon intermedius at three depths four times a year off hydrographic conditions, as well as biological factors
Santos, Brazil: surface (l m), midwater (20-25 m), and such as predators, may affect the migratory behavior
near bottom (45-47 m) . While Pseudevadne tngesti- of these species.
na and Evadne spinifera were in the surface zone con-
sistently throughout the period, Penilia avirostris and Surnrnary of distributional patterns of the species
Podon intermedius resided between O-50 and 20-50 m, covered (modified from Della Croce, 1974)
respectively, the former preferring the upper 25 m.
Resgalla and Montú (1993) reported on vertical distri- Penilia avirostris. World Ocean, particularly abun-
bution pattems of species from two winter and surnmer dant in so me warm-temperate eutrophic embay-
cruises with 37 stations offRio Grande (Brazil). Their ments or inland seas.
results also demonstrated the typical surface prefer- Podoll leuckarti. Baltic Sea, North Sea, English
ence of c1adocerans. Two of the 3 winter species, Channel, Irish Sea, North Atlantic - northeastem
Pleopis polyphemoides and Evadne nordmanni (but North America, Iceland-Greenland Seas;
not Podon intermedius) were restricted in their distri- Norwegian Sea, White Sea, Barents Sea, Kara
bution to the upper 50 m, but 4 ofthe 5 predominant- Sea, eastem Siberian Sea, Chukchi Sea, Bering
Iy surnmer species, Penilia avirostris, Pseudevadne Sea, Japan Sea, northwestem Pacific; southwest-
tergestina, Pleopis polyphemoides and Pleopis em Atlantic. Coastal, cold water species.
schmackeri, extended their distribution down more or Podon interllledius. Baltic Sea, North Sea,
less to depths of 50-100 m. Only Evadne spinifera was Norwegian Sea, English Channel, Irish Sea,
distributed in the 25-50 m depth range. Atlantic Ocean from Ireland to Iceland, northwest-
em Atlantic, Mediterranean, Black Sea, south-
In temperate waters of the northem hemisphere, a westem Atlantic, southeastem Australia.
slight but distinct diel rhythm in the vertical migration Temperate coastal species.
has been noted for several podonid species. For exam- Pleopis polyphellloides. Baltic Sea, North Sea,
pIe, in the southem Sea of Japan (depth 1000 m) English Channel, Irish Sea, Norwegian Sea, north-
Evadne nordmanni was distributed within the upper westem Atlantic, eastem and westem Pacific,
125 m. During the nighttime, most individuals Mediterranean Sea, Black Sea, eastem and west-
(-83%) were concentrated in a discrete layer at 30 m, em Atlantic, South African waters, Australian
whereas during the day alrnost all animals (99.9%) waters, New Zealand waters. Temperate. coastal,
occupied the upper 20 m (Onbé and Ikeda, 1995). A embaymental, estuarine species.
similar "reverse" diel vertical migration has been Pleopis schlllackeri. Mainly in waters of the Far East
observed for Pleopis polyphemoides in the Chesapeake from South China Sea through northem Honshu,
Bay, USA (Bosch and Taylor, 1973) and in the Inland Japan; sporadic records from Aqaba Bay (Red
Sea of Japan (Onbé, 1974, 1978a), and for Pleopis Sea), Madagascar (Indian Ocean). Recent1y
schmackeri, Evadne spinifera and Pseudevadne ter- recorded from coastal waters of southem Brazil,
gestina in the southem Sea of Japan (Onbé and Southwestern Atlantic. Warm water, coastal,
Ikeda,1995). By contrast, Pellilia avirostris exhibited oceantc speCles.
Evadne nordmanni. Atlantic Ocean (mainly north of Classification

400N), Greenland Sea, North Sea, Baltic Sea,
Norwegian Sea, Barents Sea, Kara Sea, eastern Traditionally, marine cIadocerans (order Cladocera)
Siberian Sea, Chukchi Sea, Bering Sea, were cIassified into 8 species belonging to 3 genera,
Mediterranean Sea, Black Sea, eastern and west- Penilia, Evadne and Podan: the first was assigned to
ern Pacific Ocean, southeastern Australian waters, the superfamily Sidoidea (Brooks, 1959), family
southwestern Pacific, southeastern and southwest- Sididae, and the latter two to the superfamily
ern Atlantic. Coastal, oceanic cold-water species. Polyphemoidea (Brooks, 1959), family Polyphemidae.
Evadne spinifera. Coastal, oceanic waters of all Based on a comparative study of the Caspian and
oceans approximately between 600N and 40oS. marine Polyphemoidea, Mordukhai-Boltovskoi (1968,
Coastal, oceanic warm-water species. 1978) proposed a new family Podonidae within the
Pseudevadne tergestina. Occurring widely from tem- superfamily Polyphemoidea, in which he placed 7 gen-
perate to tropical waters in both northem and era: Pleopis (incIuding "Podan" polyphemoides, as
southern Hemispheres. Coastal, oceanic warm also proposed by Gieskes, 1971a, and the redescribed
water species. "Podan" schmackeri); Pseudevadne (incIuding
"Evadne" tergestina, as suggested earlier by Di
Caporiacco, 1938); Podan, Evadne, Podonevadne,
Taxonomy Cornigerius and Caspievadne (the last 3 genera being
endemic to the Ponto-Aral-Caspian Basin). Based on a
General morphology numerical analysis of many morphological characters
of European podonids (but not incIuding "Podan"
The external morphology of marine cIadocerans dif- schmackeri), Meurice and Dauby (1983) cIaimed that
fers considerably between the 2 orders, Ctenopoda and removal of Pleopis polyphemoides from the genus
Onychopoda. In the former, a bivalved carapace cov- Podan was not justified. However, Mordukhai-
ers the thoracic region and the thoracic limbs (thoraco- Boltovskoi and Rivier (1987) used this taxonomy
pods, or trunk limbs, or legs) completely; whereas in throughout their thorough review of the world preda-
the latter only the dorsal part of the body (the brood tory cIadocerans (Podonidae, Polyphemidae, Cerco-
chamber) is enveloped by the carapace, the thoracic pagidae and Leptodoridae), incorporating the marine
limbs being exposed. Thoracic limbs also differ great- Pleopis schmackeri and many Ponto-Aral-Caspian
Iy. Ctenopoda have 6 pairs of indistinctly segmented specles.
thoracic limbs, while Onychopoda bear only 4 pairs of
thoracic limbs, which are modified into segmented, Fryer (l987a, b) recently rejected the order Cladocera
prehensile, appendages (the last pair being reduced). as a taxonomic unit in the belief that it is not a mono-
phyletic group, and divided it into 4 orders:
The distribution of setae on the segments of the anten- Ctenopoda (formerly superfamily Sidoidea Brooks,
nae and the exopods of the thoracic limbs is an impor- 1959), Anomopoda (formerly superfamily
tant diagnostic character, also known as the "setal for- Chydoroidea Brooks, 1959), Onychopoda (formerly
mula" (see Brooks, 1959). This pattem is expressed as superfamily Polyphemoidea Brooks, 1959), and
follows: a, a', a" ...Ib, b', b" ..., where a is the number of Haplopoda (formerly suborder Hap10poda G. O. Sars,
setae on the successive exopod segments (from proxi- 1865). The former 3 orders had previously been
mal to distal), while b is that on the endopod segments. placed in the suborder EucIadocera Eriksson, 1934. In
For example, the antennal setal formula of the genus a recent review on the reproductive biology ofmarine
Evadne is O, 1,1,4/1, 1,4; thus, the 4-segmented exopod c1adocerans (Egloff et al., 1997), we adopted
has no setae on the first segment, one on each ofthe fol- Mordukhai-Boltovskoi's taxonomy and Fryer's
lowing 2, and 4 on the last one while the endopod has 3 system for marine cIadocerans, but, as suggested by
segments bearing 1, 1, and 4 setae, respectively (Fig. Fryer (l987a, b), for the sake of convenience the old
6A, B, E). The setal formula of the exopod of the the- established name "Cladocera" is retained here as a
racic limbs of Podon ;ntermed;us is 2, 1,I,2; that is, 2 general descriptive term for these animals.
setae on the exopod of the flISt limb, 1 on each of the
following 2, and again 2 on the last one (Fig. 4D).
Outline c1assification (marine species only) 7a Exopods ofthoracic limbs 1-4 with 2,2,2,1 setae,
antennal elevator museles diverging dorsally:
Subelass Branchiopoda Latreille, 1817 Evadne spinijera (Fig. 6E-G)
Order Ctenopoda G. O. Sars, 1865 (formerly super-
family Sidoidea) Diagnoses of the taxa
Family Sididae Baird, 1850
Genus Penilia Dana, 1852 Subelass Branchiopoda
Order Onychopoda G. O. Sars, 1865 (formerly According to Fryer's (1987a, b) elassification system,
superfamily Polyphemoidea) the subelass Branchiopoda consists of 10 orders, of
Family Podonidae Mordukhai-Boltovskoi, 1968 which 4 orders comprise the so-called Cladocera:
Genus Podon Lilljeborg, 1853 Ctenopoda, Anomopoda, Onychopoda, and Haplo-
Genus Pleopis Dana, 1852 poda. Marine eladocerans inelude Ictenopod and 7
Genus Evadne Lovén, 1836 onychopod species. The diagnostic features of these
Genus Pseudevadne (Claus, 1877) new orders are described in detail by Fryer (1987b).
Only the external features of Ctenopoda and
Onychopoda, as defined by Fryer (1987b), are sum-
Identification marized below.
Key to species Order Ctenopoda (=superfamily Sidoidea) (surnmar-

ized from Fryer, 1987b)
6 pairs of thoracic limbs (trunk limbs), covered Body short, with few segrnents, tagrnosis mostly
with bivalved carapace: genus Penilia, Penilia obscure, terminating in postabdomen (telson) armed
avirostris (Fig. 3) with a pair of terminal elaws. Postabdomen some-
la 4 pairs ofthoracic limbs, exposed, carapace cov- times tilted ventrally but never reflexed and never
ering only dorsal brood pouch 2 articulated to trunk. Trunk and its appendages, but not
2 Head and trunk separated by a distinct cervical head, enelosed by functionally bivalved, but hingeless
groove; dorsal pouch more or less rounded 3 and uncalcified, carapace. Head short. No headshield.
2a Head and trunk continuous without distinct cervi- Labrum fleshy. Compound eyes fused to give single,
cal groove; dorsal pouch tapering, triangular.. ....4 median sessile eye with many lenses. Ocellus presento
3 Exopods of thoracic limbs 1-3 with 1, or 2 setae: Antennules tubular in female. Antennae biramous,
genus Podon (Fig. 4) 5 natatory (secondarily uniramous in female of
3a Exopods of thoracic limbs 1-3 with 3 or 4 setae: Holopedium). Exopod and endopod each with 2 or 3
genus Pleopis (Fig. 5) 6 segments, between them armed with 13 (Penilia) to
4 Exopods of thoracic limbs 1-3 with 2,2,1, or 27 natatory setae. Six pairs oftrunk limbs (or thorac-
2,2,2 setae; dorsal pouch ending in spine: genus ic limbs), all pre-genital and showing much serial
Evadne (Fig. 6) 7 similarity, but 6th pair reduced.
4a Exopods of thoracic limbs 1-3 with 2,3,3 setae;
end of dorsal pouch always rounded: genus Family Sididae
Pseudevadne, Pselldevadne tergestina (Fig. 7) Antennules movable, short in female and long in male.
5 Exopods ofthoracic limbs 1-4 with 1,1,1,2 setae: Antennae long, consisting of a large basipod with 2
Podon lellckarti (Fig. 4A, B) branches (endo- and exopods). Carapace oblong,
5a Exopopds of thoracic limbs 1-4 with 2,1,1,2 covering 6 pairs of thoracic limbs and postabdomen.
setae: Podon interllledills (Fig. 4C-E)
6 Exopods ofthoracic limbs 1-4 with 3,3,3,2 setae: Genus Penilia
Pleopis polyplzellloides (Fig. 5A-C) Diagnosis as for the species below. This genus was
6a Exopods ofthoracic limbs 1-4 with 4,4,4,2 setae: considered to contain several species (Richards, 1895;
Pleopis sclzlllackeri (Fig. 5D, E) Kramer, 1895), all of which were subsequently
7 Exopods of thoracic limbs 1-4 with 2,2,1,1 setae; lumped into Penilia avirostris Dana (Calman, 1917;
two antennal elevator museles running parallel Steuer, 1933). In the light of modern taxonomy, how-
to each other: Evadne nordlllanni (Fig. 6A-D) ever, all specimens throughout the World Ocean
Penilia avirostris
@
female antennule
" ~ caudal
seta
head of female
4]) in ventral view
o
should be re-examined (Korovchinsky, 1983, 1986, Single, large, median, compound eye with many
1996; Onbé, 1997). ommatidia. No ocellus. Labrum large. Antennules
• Pe"ilia av;rostr;s Dana, 1852 (Fig. 3). Body trans- tubular and uniramous. Antennae biramous and nata-
parent; rostrum sharply pointed in female, rounded tory, with 3-segmented endopodite and 4-segmented
in maleo Antennules short with several distal aes- exopodite bearing long natatory setae. Four pairs of
thetascs (small sensory setae covered by delicate segmented, grasping thoracic limbs, generally with
cuticle, often projecting from antennule, also exopodites and gnathobases but no branchial epipo-
known as olfactory hairs) in females, but elongated dites.
and almost reaching posterior end of carapace in
males. Antennae consisting of a large basipod, and Family Podonidae
2-segmented exo- and endopods. Exopods bearing Antennules very short, fixed undemeath head and
2 swirnming setae on first segment and 6 on the immovable. Antennae short, biramous; exopod 4-seg-
second, endopods with 1 (on first segment) and 4 mented and endopod 3-segmented with setal formula
(on the second) swirnming setae (setal formula: 0,1,1 or 2,4/1 ,1,4. Four pairs ofthoracic limbs, exopod
2,6/1 ,4). Carapace with a prominent spine (the so- with 1 to 4 setae. Caudal furca either strongly develop-
called "Schalenstachel", shell spine) on posterior ed (long), or less developed (short). Carapace cover-
end (carapace spine in Fig. 3A), with row of min- ing only dorsal brood chamber, trunk and thoracic
ute spines on both ventral and postero-dorsal edges, limbs exposed. Cervical organ (Fig. 4A) present at
and much finer spinules between these spines. dorsal edge above base of antennae. Males differing
Postabdomen large, ending in a pair of long termi- from females by strong terminal claw or hook of endo-
nal claws with 2 spines (distal one much longer pod of thoracic limb 1 (Fig. 4B), and a pair of penes
than proximal) in postero-basal portion. Dorso-pos- situated anterior to postabdomen (Fig. 4B). Head and
terior comer of carapace depressed, bearing a pair compound eye relative to body much larger than in
of long setae (so-called "Schwanzborsten", tail females. All species inhabiting surface layers of estu-
setae or caudal setae, see Fig. 3A). Six pairs oftho- arine, coastal, or open waters ofthe World Ocean.
racic limbs (thoracic legs, or thoracopods), last one
short and reduced; all limbs biramous, endopod Genus Podo"
indistinctly segmented with many minute plumose Head and carapace always separated by distinct cervi-
setae directing posteriorly, exopod consisting of a cal groove (Fig. 4A). Brood pouch oval to hemispher-
flat lamella with several thick plumose setae. ical. Postabdomen ended with pair of long caudal fur-
Endopod of male thoracic limb 1 with a sharp cae. Exopods of thoracic limbs 1-3 with 1,1,1 or 2,1,1
recurved claw on distal end (terminal hook in Fig. setae. Resting eggs: spherical, membrane furnished
3E). In males a pair of copulatory appendages with omamentations; perimeter indistinct. Male: dor-
(penes) located posterior to thoracic limb 6. Resting sal part small, triangular, with a pair oftestes; head and
eggs: usually produced by larger females in popu- eye large relative to female; terminal seta of endopod
lation; grayish, ovoid and somewhat flattened, may of first thoracic limb transformed into recurved hook
vary in size geographically, in the Inland Sea of (Fig. 4B), a pair of copulatory appendages (penes)
Japan 0.21-0.29 rnm (mean: 0.25 rnm) long, 0.14- located posterior to the last pair of thoracic limbs.
0.20 mm (mean 0.18 mm) wide, ca. 0.10 rnm in • Podo" leuckart; G. O. Sars, 1862 (Fig. 4A, B).
depth (Onbé 1972). Totallength, females: 0.4-1.2 Body large. Brood pouch small, hemispherical.
rnm; males: 0.7-0.9 rnm. Caudal furca large, elongate. Antennae: segments
1-4 ofexopods with 0,1,1,4 setae. Exopods oftho-
Order Onychopoda (=superfamily Polyphemoidea racic limbs 1-4 with 1,1,1,2 setae. Resting eggs:
Brooks, 1959) (summarized from Fryer, 1987b) spherical, large, 0.21-0.23 mm in diameter, light
Body short. Thorax short; segmentation obscured by brownish in color; outer membrane densely pored.
fusiono Carapace reduced to a dorsal brood pouch. Totallength (standard length = body length): 0.8-
Fig.3. Penilia al'irostris. A: original; S-E: from Onbé (1997), redrawn.

808
Ct
Podon intermedius,
parthenogenetic female
I-IV: 1st to 4th

thoracic limbs
o
Podon leuckarti,
parthenogeneti e
female
111 IV
I/~Iíf terminal
hook
:',: I
Podon intermedius,
exopods of 1st-4th
\.~'.':''::.''"''''''
.L
~<t ~::"''',,,,":
\':'
\~
I
,/
~
:;',,'1'
V
\"
~
caudal
furca
~
caudal
process
caudal
seta
O> thoracic limbs
Podon intermedius,
4} caudal part
1.0 mm (0.5-0.9 mm) in female, <1.0 mm (0.5-0.8 Genus Evadne

rnm) in male. Body oval or triangular. Carapace: posterior tip more or
• Podon inter11ledillS Lilljeborg, 1853 (Fig. 4C-E). less pointed. Cervical groove indistinct, head directly
Body large. Brood pouch small, oval to hemi- connecting to brood chamber without distinct depres-
spherical. Caudal furca large, elongated. Caudal sion in between (but often depressed when brood
process larger than that of any other species of chamber contains many fully developing embryos).
Pleopis and Podon. Antennae: exopod of segments Antennae: smaller than those of Podon and Pleopis,
1-4 with 0,1,2,4 swirnming setae. Exopods oftho- exo- and endopods 4- and 3-segmented, respectively;
racic limbs 1-4 with 2,1,1,2 setae. Resting egg (a setal formula 0,1,1,4/1,1,4. Thoracic limbs: 4 pairs;
specimen from off Woods Role, Massachusetts, exopods of thoracic limbs 1-3 with 2,2, 1, or 2,2,2 setae.
USA; Onbé and Grice, unpublished): spherical, Postabdomen thick, short, ending with a pair of quasi-
large, 0.27 rnm in diameter; outer membrane with equitriangular caudal furcae. Resting eggs: spherical,
complex array of groove structures. Total length without any noticeable omamentation on egg surface;
(standard length): 1.1-1.2 rnm (1.0 rnm) in female, perimeter clearly definable. Male characteristics simi-
0.6-1.0 mm (0.9-1.0 mm) in male. lar to those of other podonid species.
• Evadne nord11lanni Lovén, 1836 (Fig. 6A-D).
Genus Pleopis Body triangular, brood pouch variable in shape
Exopods of thoracic limbs 1-3 with 3,3,3 or 4,4,4 depending on developmental stages of embryos.
setae. All other features similar lo Podon. Carapace always pointed at posterior end. Two
Pleopis polyphe11loides (Leuckart, 1859) [=Podon antennal elevator muscles lying parallel to each
polyphemoides (Leuckart, 1859)] (Fig. 5A-C). other. Exopods of thoracic limbs 1-4 with 2,2, 1,l
Body small. Brood chamber varying from hemi- setae. Resting eggs spherical, ca. 0.20 rnm in dia-
spherical to spherical. Caudal furca short, in out- meter; egg membrane smooth, with clear perimeter;
line almost an equilateral triangle. Antennae: exo- cytoplasm roughly granulated, light brown. Total
pod segments 1-4 with 0,1,2,4 swirnming setae. length (standard length): 0.4-1.4 rnm (0.2-0.7 rnm)
Exopods of thoracic limbs 1-4 with 3,3,3,2 setae, in female, 0.4-0.8 rnm (0.2-0.7 rnm) in maleo
respectively. Resting eggs: small, spherical, 0.15- • Evadne spb,ijera P.E. Müller, 1867 (Fig. 6E-G).
0.21 mm in diameter (mean 0.18 mm); greenish- Body oval. Carapace ending in a long spine.
brown; numerous small pores scattered throughout Antennal elevator muscles diverging dorsally.
outer egg membrane (pores crater-shaped as seen Exopods of thoracic limbs 1-4 with 2,2,2,1 setae.
in SEM); perimeter obscured with the omamenta- Resting eggs unknown. Total length (standard
tion of membrane. Totallength (standard length): length): 0.7-1.4 mm (0.5-0.7 rnm) in female, 0.6-
0.3-0.7 rnm (0.2-0.6 rnm) in female, 0.4-0.7 mm 1.3 rnm (estimated values, 0.5-0.7 rnm) in maleo
(0.4.-0.7 mm) in male.
Pleopis sch11lackeri (Poppe, 1889) (=Podon GenusPselldevadne
schmackeri Poppe, 1889) (Fig. 5D, E). Body hem- Posterior tip of carapace always rounded. Four pairs
ispherical, with shallowest cervical groove in gen- of thoracic limbs; exopods of thoracic limbs 1-3 with
era Podon and Pleopis. Caudal furca slender, 2,3,3 setae. Lateral sides of carapace (crista of fomix,
sharply pointed. Antennae: exopod of segments cf. Mordukhai-Boltovskoi, 1983) partly overhang,
1-4 with 0,1 ,2,4 setae. Exopods of thoracic limbs covering basipods of thoracic limbs. Other characters
1-4 with 4,4,4,2 setae, respectively, the most similar to Evadne.
numerous in the marine Podonidae. Resting eggs • Pselldevadne tergestina (Claus, 1877) (=Evadne
unknown. Total length (standard length): 0.34- lergeslina Claus, 1877) (Fig. 7). Body oval.
0.87 mm (0.30-0.65 mm) in female, 0.43-0.46 mm Carapace always with rounded posterior end, lack-
(0.42-0.43 mm) in male (based on only 2 indivi- ing spine; varying in shape from elongate-oval to
duals) . hemispherical. Elevator muscles of antenna
•••
Fig.4. Podan lellckarti and P. inlermedills. From Onbé (1997), redrawn.
810 Ctenopoda and Onychopoda (=C1adocera)
Pleopis
111 IV
.~
Pleopis polyphemoides,
@ exopods of 1st-4th
thoracic limbs
"
Pleopis polyphemoides
4])
Pleopis
polyphemoiiJes
male
\' \
\'
\'
111 IV
Pleopis schmackeri,
@ Pleopis schmackeri 4) exopods of 1st-4th
thoracic limbs
Cten~poda and Onychopoda (=Cladocera) 811
diverging dorsally. Exopods of thoracic limbs 1-4 Proposal for assigning the Cladoeera to 4 separate
with 2,3,3,1 setae, respectively. Several rows of orders beeause of their polyphyletie origins.
hypodermal cells present in brood pouch (often Fryer G. 1987b. A new classification ofthe branchio-
disappearing in preserved specimens). Resting pod Crustacea. Zool. J. Linn. Soc., 91:357-383.
eggs: spherical, 0.19-0.24 mm (mean 0.20 mm) in Formal deseription of a new classifieation system
diameter, egg membrane smooth (but SEM obser- of the Branehiopoda, including the Cladoeera.
vation shows fine mat of microvillar structures), Gieskes W.W.C. 1970. The Cladocera of the North
with clear perimeter, cytoplasm thickly granulat- Atlantic and the North Sea: biological and ecolog-
ed, light-brownish. Totallength (standard length): ical studies. Ph. D. Diss., McGill Univ., pp. 1-205.
0.3-1.3 mm (0.3-0.9 mm) in female, 0.5-0.8 mm A comprehensive survey of the distriblltion,
(0.3-0.6 mm) in male. growth, and reprodllction, of marine cladocerans
based on specimens collected mostly by CPR
(Continuous PlanÁ.'fonRecorder).
Acknowledgments Kim S.W. 1989. Studies on the ecology ofmarine cla-
docerans in the northwestern Pacific Ocean. Ph. D.
Dr. Geoffrey Fryer kindly reviewed the manuscript Diss., Hiroshima Univ., pp. 1-180. A detailed
offering several valuable suggestions. 1 thank accollnt of the distribution and abundance of
Professor Demetrio Boltovskoy for persuading me to marine cladocerans based on specimens collected
write this chapter, and for offering valuable sugges- during the Cooperative Study of the Kuroshio and
tions and comments on the manuscript. Prof. Charrid Adjacent Regions (CSK), with biological notes on
Resgalla kindly placed at my disposal specimens of some species.
marine cladocerans from coastal Brazilian waters. Lilljeborg W. 1901. Cladocera Sueciae, order
Beitrage zur Kenntnis der in Schweden lebenden
Krebsthiere von der Ordnung der Branciopoden
Suggested readings und der Unterordnung der Cladoceren. Nova Acta
Reg. Soco Sci. Uppsala, 3(1-6):1-701. Classical
Baker H.M. 1938. Studies on the Cladocera of descriptions of cladocerans with many sllperb
Monterey Bay. Proc. California Acad. Sci., Ser. 4, illllstrations.
23:311-365. Some of the most detailed deserip- Mordukhai-Boltovskoi Ph.D., Rivier I.K. 1987.
tions and illllstrations of marine podonid speeies Khishchnye vetvistousye Podonidae, Polyphemi-
.from California. dae, Cercopagidae i Leptodoridae fauny mira .
DeBa Croce N., Venugopal P. 1972. Distribution of Fauna SSSR, 148, Akad. Nauk SSSR, Zool. Inst,
marine cladocerans in the Indian Ocean. Mar. Biol., Nauka, Moskva: pp. 1-182. A comprehensive
15: 132-138. Distriblltion of 4 species of lIlarine review of almost all aspects of the world predato-
cladoeerans eolleeted during the Jnternational ry cladocerans, including marine podonid species.
Indian Oeean Expedition (lIOE). Biology, ecology, world distribution, descriptions
Egloff D.A., Fofonoff P.W., Onbé T. 1997. and illustrations of species.
Reproductive biology of marine cladocerans. Onbé T. 1991. Some aspects of the biology ofresting
Advances Mar. Biol., 31:79-167. A review of all eggs of marine cladocerans. In "Crustacean Egg
aspeets of the reproduetive biology, with eoneise Production" (A. Wenner, A. Kuris, eds.),
aeeounts on distribution, growth and nutrition of Crustacean Issues 7, A. Balkema, Rotterdam, pp.
marine cladoeerans, including substantial eom- 41-55. A review ofthe resting eggs ofmarine cla-
parative information on jreshwater speeies. docerans: collection and sorting, morphology and
Fryer G. 1987a. Morphology and the classification of identification, distribution and ablllldance in sea-
the so-called Cladocera. Hydrobiologia, 145:19-28 . floor sediments, etc.
••
Fig.5. Pleopis polyphemoides and P. schmockeri. A-C: from Onbé (1997), redrawn; D, E: original.
,~
Evadne
~ \~I ~ terminal
hook
@~~
~
, f:S I seta
terminal
Evadne nordmanni,
male, 1st thoracic limb
distal part of endopod
I~C'::J ~~
~
. .I\~~~\;
\ ~ ~caudal
emb,yo
CD
"
caudal ~ process
I~\~~ ~ouch
Evadne nordmanni,
furca ~~ exopods of 1st-4th
caudal
seta Evadne nordmanni , thoracic limbs
narthenogenetic female
1'"'"
terminal
hook
4) sn copulatory Evadne nordmanni,

male
o
appendage
(penis)
111 IV
Evadne spinifera,
4) ~----~ E~ooe~m~~
parthenogenetic female with advanced I
emryos ana a batch of eggs of next brood
@ exopods of 1st-4th
thoracic limbs
Cteoopoda and Onychopoda (=Cladocera) 813
Pseudevadne tergestina
~.. ~
/í
copulatory
appendage
(penis)
male
hook
exopods of 1st-4th
thoracic limbs
Fig.7. Pselldel'adne tergestina. A, C: original; B: from Onbé (1997), redrawn.
RamÍrez F.C. 1981. C1adocera. In "Atlas del Smimov N.N., Tirnms B.V. 1983. Revision of the
Zooplancton del Atlántico Sudoccidental y méto- Australian Cladocera (Crustacea). Rec. Australian
dos de trabajo con el zooplancton marino" (D. Mus., Suppl. 1: 1-132. Comprehensive deserip-
Boltovskoy, ed.), Publico Esp. Inst. Nac. Inv. tions and illustrations offamilies ofinland cladoe-
Desarrollo Pesq., Mar del Plata, pp. 533-542. A erans, with speeies of marine Sididae (by N. M.
eoncise review of systematies, bi%gy, distribu- Korovehinsky) and Podonidae (by Ph. D.
(ion, and researclz methods, including the distribll- M ordllkhai- Boltovskoi).
(ion of the Southwestern At/anlie species.
Fig.6. Evadne nordmGnni and E. ~pinirerG. A-D: from Onbé (1997), redrawn; E-G: original.
Ostracoda
Martin V. Angel
J-l.¿...oY O
7 g- O . 4 hettacra 200 Cli
.•....•
Introduction Alada 400
5/ 200
400
@óQ'
eL
CDW --------_
1 2a 3..-0 w........•. -------e"~-O+O
.... Cli
..r:. ShelfooAd
..r:.
E
ñ.
O CDW e a
O
4° belgicae
W
Alada ----------
Shelf ----
Ostracoda are a sub-class of relatively small crusta-
Q
ceans which occur(JAASW
predominantly ~in aquatic habitats.
(JAASW ~
The majority of species are benthic, but in oceanic
waters most members of the family Halocyprididae
and a few species of the family Cypridinididae are
holoplanktonic. In neritic waters many species of
Cyprinidinidae and other myodocopid families are
meroplanktonic. Adult sizes of the oceanic planktonic
species range from lengths of 0.5 to 30 mm, but most
are in the range of 0.8 to 4 rnm. Apart from the large
bathypelagic species of Gigantocypris, planktonic
ostracods have received scant attention from oceanic
ecologists despite their relatively high numerical abun-
dance (often outnumbered only by copepods) and more
or less ubiquitous occurrence at all oceanic depths.
Planktonic ostracods, unlike their benthic counter-

parts, have carapaces with little if any calcification, so
they are poorly represented in the fossil record. Since Fig. 1. Models of the life cycles of Alacia hettacra and A. belgi-
the majority of ostracodologists are palaeontologists, cae in the region of the Antarctic Peninsllla. Adlllts (Ad), while
rather than zoologists or ecologists, the planktonic spe- occllpying Antarctic Sllrface Water (AASW) in the llpper 100 m of
cies have been neglected despite their importance in the water column are advected offshore. They migrate down 500 m
into Circllmpolar Deep Water (CDW) in autumn (a) where they
pelagic ecosystems. The earliest taxonomic descrip-
release their eggs (e). The eggs hatch and develop throllgh sixjllve-
tions to meet modern standards were by G. O. Sars nile stages (1-6) during the winter (w) and early spring (sp), while
(1865) who described three species from Norwegian being advected back in over the shelf. The later stages migrate back
waters. Much of the pioneering taxonomic work was up towards the surface where they develop into adlllts as they move
carried out by C. Claus (1874,1890, 1891, 1894) and back ¡nto the upper layers. Redrawn from Kock (1992).
G. W. Müller (1894, 1906, 1912), who described

material collected in the Mediterranean and during the
major exploratory expeditions of the late 19th and although Cannon (1940) had earlier described the )
early 20th centuries on many vessels, notably the internal anatomy of Gigantocypris muelleri from this
Valdivia, Belgica and Siboga. Many of these early material.
expeditions sampled in the South Atlantic. Other expe-
ditions that made major contributions to planktonic There is relatively little information available about
ostracod studies were the Swedish Antarctic expedi- the Iife histories of most myodocopid species. Kock
tion (see the exhaustive treatise by Skogsberg, 1920), (1992) developed a model of the Iife-cycles of two
and the Dana expedition (see the series of papers by Alacia species endemic to the Southern Ocean. The
Poulsen, 1962, 1969, 1973, 1977). I1es (1961) adults occur near surface in Antarctic Surface Water
described some of the ostracods collected in the during the early surnmer and so are alJvected off the
Benguela Current region by William Scoresby, but dis- shelf over deep water (Fig. 1). Before hatching the
appointingly never completed the analysis ofthe copi- eggs sink to depths of 400-500 m, where the early
ous material collected during the Discovery Expedi- juveniles stages stay within the Circumpolar Deep
tions (some of this material has been used herein), Water which slowly moves them back over the shelf.
Soulh AI/anlic Zoop/ankton.

edited by D. Bo/tovskoy, pp. 815-868
1999 Backhu)'s Publishers, Leiden, The Nether/ands
816 Ostracoda
Paramollicia rhynchena
44" N 13"W
April'1974
V'>
n:l
~ 200
>
IJ
e
'<- -nffi1hp."
O
•...
Q) Male
...o 100 N=110
E Mean=2.34
::::l
1.6
2.0
2.4IV
Instar
N=609
N=587
Instar
0.81.2 V Instar
N=442
Mean=1.24
N=199
VI
Mean=1.79
Mean=2.44
Female
m m)
Fig. 2. Size classes of a population of Paramollicia rhy"che"a sampled at a depth of 500-600 m, showing how the size ranges of!he four juve-
ni le instars and !he adults are quite discreteo The mean carapace lengths show that the carapace increases in length by a factor of 1.3-1.4 at each
instar. From Angel (1993).
During the winter the last two juvenile stages migrate Halocyprids will eat almost any dead material offered
back Up towards the surface ready to restart the cycle to them, but do not appear to be active predators. In
when they mature in the spring. The development has fresh net samples the halocyprids can be seen feeding
been described for a Southern Ocean species P. serru- actively on gelatinous, damaged organisms and parti-
lata by Ramirez and Perez Seijas (1981). Length cles. They are clearIy able to handle "particles" which
tends to increase by a factor of 1.3-1.4 at each moult are considerably larger than themselves, manipulating
with little or no overIap in carapace lengths between them with the mandibles and the fifth and sixth limbs,
successive instars (Fig. 2, 3). rotating them and finally rejecting them using the cau-
dal furca. Dnder normal conditions they are probably
Some species (e.g., Conchoecilla daphnoides and O. opportunistic feeders and probably actively feed in the
haddoni) show similar seasonality in their reproduc- net while being caught. Analyses of gut and faecal
tion; at the end of March only small juvenile instars pellet contents have shown that the food taken varies
were present at 44°N, 13°W (Angel, 1977), whereas widely between species. Some species feed on detri-
later in the Spring adults appeared (Angel, 1984). tus, since some bathypelagic species have large num-
Similar seasonality can be expected at high latitudes bers of phytoplankton tests in their guts, which are
in the South Atlantic. However, in most species, spe- probably derived from the sedimenting rain of detrital
cimens at all developmental stages can be found at a aggregates. Some species have guts containing large
locality throughout the year. There is a general, but by quantities of folded membranes which appear to be
no means unversal trend, for the earIy juvenile stages moulted carapaces of other crustaceans. In others, the
to occupy shallower depths than the adults, and for contents are hard and resinous and have an amber-like
any species which undertakes seasonal and/or ontoge- consistency; their food source remains a mystery.
netic migrations a false impression of absence may be Planktonic ostracods seem likely to be playing an
gained if the sampling regime fails to cover the important role in the re-cycling of material within the
species's full range. body of the deep ocean. AII the myodocopids that
have be en studied, have proved to be carnivorous.
Ostracoda 817
Boroecia antipoda Alacia belgicae

Fig.3. Superimposed outlines of juvenile and adult female instars of two species. Modified from Kock (1992).
Gigantocypris species take a wide variety of small sampling micranekton. In open oceanic waters abun-
micronektonic prey including mysids, chaetognaths, dances are low, so about 2500 m3 of water need to be
copepods and medusae. Their prey can be much the filtered to give quantitatively meaningful samples,
same size as the ostracod itself. Macrocypridina spe- particularly fram depths >500 m, i.e. a 1 m2 net needs
cies are also predators of plankton, consuming large to be towed for one hour at 2 knots (1 m.s-1). This
numbers of copepods. towing speed is the compromise most widely adopted
which balances the needs to minimise avoidance, to
All non-selective planktivores will inevitably include filter enough water, and to collect material in identifi-
some ostracods in their diets, since the ostracods are able condition. At depths of 100-400 m where the
often the second-most abundant group ofmacraplank- ostracods are usually most abundant, >25,000 speci-
ton. Cyclothone species which are by far and away the mens are Iikely to be collected. Vertically hauled nets
most abundant fishes at mesopelagic depths, myc- pravide good material but seldom pravide enough
tophid fishes, decapod crustaceans, and heteropod specimens to describe the cornmunity structure ade-
molluscs have all been reported as being non-selective quately. Ostracod concentrations decline exponential-
feeders which take ostracods. There are a few species ly with depth, so if adequate numbers are to be col-
which specialise in ostracod predation, the hatchetfish lected, the nets need to be towed longer. The
Argyropelecus aculeatus predominantly eats Mikro- deeper-living species are also more fragile, so it is dif-
conchoecia curta and the siphonophore Hippopodius ficult to achieve the optimal balance between collect-
hippopus feeds almost exclusively on halocyprids, ing a large enough sample by towing for a long time
and Diphyes dispar and Chelophyes appendiculata and collecting material that is not too damaged to
also prefer to feed on the ostracods. The vertical ran- identify. Sampling the benthopelagic realm requires
ges of• these predators match those of their prey. the nets, either to be attached above bottom trawls, or
fitted with echo-sounding devices to continuously
monitor in real-time the height of the net above the
Methods bottom. Collections fram research submarines and
deep-towed instrument packages collect good qualita-
Representative samples of adults and the later juvenile tive, but poor quantitative, materia!. Results of repeat-
stages are collected using plankton nets with 200-300 ed sampling of a community at a depth of 1000 m sug-
j..Lmmeshes, but larger species are also taken in trawls gest that an accumulative sample of araund 10,000
818 Ostracoda
specimens is needed to provide representatives of all 30% in the numbers of halocyprid species caught at all
the species presento depths down to at least 2000 m (Fig. 4). At the same
time, dominance of the assemblages by a very few spe-
Few efforts have been made to culture oceanic ostra- cies witWn each depth stratum increased. Similar trends
cods (e.g., Ikeda, 1992). Specimens are easily kept occur in other pelagic taxa and Angel (1991) postulat-
alive onboard ship for a few days - long enough to ed that the changes are associated with the southem
investigate experimentally many outstanding ques- limit to which winter-mixing extends deep enough to
tions concerning their life-histories, diet, assimilation replenish surface nutrients to result in a well-developed
efficiencies, and ecological roleo spring bloom in the phytoplankton (i.e., the Subtropical
Convergence in the South Atlantic). Recently mosaics
Catches for taxonomic and distributional studies should of remotely sensed images of ocean colour have vis-
be fixed as soon as they come onboard to avoid dete- ualized pattems of chlorophyll concentrations in the
rioration. Sea water-formalin is the most widely used surface waters and clearly show this boundary, particu-
fixative. A stock preserving solútion is made up by larly in the boreal summer (Campbell and Aarup, 1992).
adding one part of concentrated (40%) formaldehyde
solution (neutralised with 5 g borax per litre) to nine In the North Atlantic the subtropical circulation is
parts of filtered sea water. Put the sample into a con- dominated by the subtropical gyre involving the Gulf
tainer, add filtered sea water until it is half-full, then top Stream and the Canary Current. Peterson and Strama
up with the stock preserving solution. Seal the contain- (1991) have provided a detailed description of the
er hermetically, trapping as little air as possible to mini- upper circulation and the distribution of major ocean-
mise oxidation and mechanical damage when the ship ographic fronts in the South Atlantic. These fronts
rolls. Ideally the catch should only occupy only 10% of coincide with the boundaries of biogeochemical pro-
the container. Change the preservative after 24-48 h. vinces (Longhurst, 1995) within which there are cha-
For long-term storage transfer it into either Steadman's racteristic seasonal cycles of productivity.
preserving fluid (10% propylene glycol, 0.5% propy-
lene phenoxetol, 1% concentrated (40%) formalde- Poulsen (1977) noted that most epipelagic and meso-
hyde, and 88.5% distilled water), or 80% ethanol. pelagic species occur in all oceans, whereas many
bathypelagic species have more restricted geographi-
cal ranges. This may be an artefact of poor sample
Zoogeography and vertical distribution coverage since deeper-living species described from
the Southern Ocean (i.e., Archiconchoecia versicula
Few wide-ranging zoogeographic studjes have been and Metaconchoecia arcuata), and the North Pacific
conducted on halocyprids in the South Atlantic; the (i.e., L. acutimarginata and P. vitja:::i) have subse-
data in Table 1 are derived from a compilation of allli- quently been found in the North Atlantic. However,
terature records, and so may give a slightly false impres- species of Mollicia and Paramollicia do have restric-
sion of the normal distributional ranges of many spe- ted ranges, and there are examples of bipolar sibling
cies. In the South Atlantic the zoogeographic patterns species pairs (Boroecia borealis and B. maxima in the
are likely to be analogous to those described by Angel Arctic and B. antipoda in the South; Obtusoecia obtll-
and Fasham (1975) along the 200W meridian in the sata in the Arctic and o. antarctica in the South).
North Atlantic. There changes in the community com-
position were related to boundaries associated with the Some species have small low latitude and large high
main hydrographic circulation. At high latitudes (Le., latitude forms, sometimes showing clinal changes (e.g.,
>500N) the vertical stratification of the populations Discoconchoecia elegans), or sometimes disjunct dis-
inhabiting the water colurnn down to depths of 2000 m tributions (e.g., Archiconchoecia cucllllata). There is a
is weakly subdivided into three depth zones. Whereas similar size and bathymetric separation between
at lower latitudes, where thermal stratification is better Paraconchoecia manzillata and P. nanomamillata.
established, the ostracods were also more clearly Angel (1982), when redescribing the two species of
organised into five bathymetric zones. A widely-spaced Halocypris inflata and H. pelagica (which had been
latitudinal transect along 200W revealed that a major synonymised as H. brevirostris), suggested that the size
faunal boundary occurs between 40-42°N. Sampling and depth ranges ofthe co-occurringjuveniles could be
across this boundary showed a poleward reduction of interpreted as exemplifying character displacement.
Ostracoda 819
..~ ~~
::::l _ .~
~tlll
.t:
.•••
O·E
-lIS
Morphology
~._
..r:::
lIS
~t':~O
tlllE
~"t:l
L.l.
oS---
.••• ·v~
tlll~
E~r:::
::::l
r:::Q.
r:::.J: ••
.....••
tlll___
~ =ti r:::
.•.. .;; '2 Z
Distribution
~r::: v lIS~~E O
Alacia alata • 1.9-2.4 1.7-2.0 38ºN-35º5 100-500 ye~.25-j

Alacia belgicae 2.32-2.96 2.20-2.86 50º5-78º5 200-500 no 9.26
Alacia hettacra 2.06-2.45 1.72-2.10 43º5-70º5 50-500 no 9.27
Alacia leptothrix 3.1-3.4 2.8-3.1 25ºN-46º5 800-2000 ? 9.28
Alacia valdiviae 5.08-6.00 -- 4.92-5.58 --- 28ºN-70º5 1000-3000 no 9.29
Archiconchoecia aff. cucullata 2.04-2.22 2.22-2.30 18ºN >3000 no 9.11
Archiconchoecia bidens 1.7 71º-57'5 1400-1600 ? 9.5
Archiconchoecia bifurcata 1.64-1.88
Archiconchoecia bimucronata
Archiconchoecia bispicula
1.00-1.15
0.96-1.02
1.50-1.60~ºN-62º5 t 1000-3700 f no 9.6
0.87-0.96
0.90-0.96 60ºN-0º
30ºN-40º5 800-1500 +I ~ no
700-2000 + 9.7
9.8
Archiconchoecia cucullata (small) 1.58-1.84 1.62-1.88 32ºN-0º 400-800 yes 9.10
Archiconchoecia cucullata 5.5 2.01-2.57 2.12-2.64 60ºN-69º5 400-1250 ?no 9.9
Archiconchoecia cuneata 0.76-0.86 0.67-0.73 40ºN-19º5 500-1000 ~ 9.12
Archiconchoecia longiseta 1.27-1.30 32ºN-0º 500->3500 no 9.13
Archiconchoecia simula 1.10-1.17 1.12-1.14 32ºN-69º5 2000-4000 no 9.14
Archiconchoecia striata 0.54-0.60 0.62 40ºN-36º5 0-200 .:les 9.15
Archiconchoecia ventricosa 0.78-1.04 0.74-0.87 49ºN-?0º 100-800 no 9.16
1
Archiconchoecia versicula 0.78-0.92
3.00-3.32 0.80-0.92
2.80-3.06 30ºN-43~
7ºN-71 º5 1000-3000
300-700 no
? 9.17
9.30
Boroecia antipoda
Conchoecetta acuminata • 2.80-3.60 2.00-2.52 43ºN-45º5 0-500 yes 9.31
Conchoecetta giesbrechti 2.16-2.34
1.68-1.90 34ºN-40º5 0-1250 yes 9.32 ~
Conchoecia hyalophyllum 1.50-1.86 1.44-1.66 64º-48º5 0-700 yes 9.33
Conchoecia lophura
Conchoecia macrocheira
2.28-2.72
3.40-3.64
2.16-2.52
3.00-3.20
60ºN-49º5
50ºN-44º5
100-2250
500-1500
---
no
no
9.34
9.35
Conchoecia magna • 1.70-1.96 1.56-1.80 50ºN-55º5 0-700 yes 9.36
Conchoecia magna, large form 2.00-2.30 1.92-2.10 Equatorial 0-700 ?
Conchoecia parvidentata 2.52-2.76 2.12-2.32
--- ?- 60º5 -+-- 0-2000 9.37
Conchoecia subarcuata 1.92-2.36 1.84-1.96 45ºN-56º5 0-600 yes 9.38
Conchoecilla chuni
Conchoecilla daphnoides'
Conchoecissa ametra __
4.17-5.25
2.0-2.4
3.32-4.44
2.40-3.04 62ºN-47º5
1.40-1 .55 ~N-63º5
3.16-4.16
0-1250
64ºN-50º5 ~ 400-2000
100-3500
yes
n~
no
1 9.40---:
9.39
9.41
Conchoecissa imbricata' 2.72-3.24 2.28-2.72 65ºN-72º5 0-2000 yes 9.42
Conchoecissa plinthina 4.92-5.83 4.67-5.50 31ºN-43º5 500-2500 no 9.43
Conchoecissa squamosa 4.28 3.88 4ºN-35º5 1000 no 9.44
Conchoecissa symmetrica 3.6-4.84 3.72-4.25 ?- 68º5 500-3300 no 9.45
Discoconchoecia discophora 1.25-1.5 1.1-1.3 24ºN-50º5 0-2000 ? 9.46
Discoconchoecia elegans' 1.10-1.85 1.00-2.00 81ºN-65º5 0-2000 yes 9.47
Euconchoecia chierchiae
Fellia bicornis
Fellia cornuta
3.28-3.60
1.7-1.9
3.3 3.16-3.48
1.5-1.8
3.3 18ºN-45º5
32-44º5 I ~~~-2000
40º5-? ~-500
750-2000 •? 9.23
9.22
Fellia dispar 1.08-1.30 1.16-1.26 18ºN-35º5 0-300 r¡s ? 9.24
9.18
Gaussicia edentata 1.58-1.73 1.40-1.52 49ºN-71º5 500-1000 no 9.48
820 Ostracoda
:l~ -~~'-"
.. .~~.s.--
:l~~
.••• !llc::
~E~ .•••~c:: u u. _~"C
:co'E c:: e~ '-" Z
'';:Distribution
u. _ '-" !ll c:::!::!ll
c::E~~ u ~ !ll~;E
Morphology
c::.J: .-o.•....
j~~o Q. ••
9.69
9.54
9.59
9.68
9.51
9.52
9.53
9.55
9.56
9.57
9.58
9.60
9.61
9.62
9.63
9.64
9.65
9.67
9.71
9.72
9.74
9.73
9.76
9.82
9.81
9.19
9.20
9.21
9.75
9.77
9.78
9.79
9.1
9.4
9.49
9.50
9.70
9.2
9.3 +
Gaussicia9.80 3.84-4.12
32ºN-42ºS
3.44-3.48
49ºN-4ºS
2.95-3.33
1.82-1.91
40ºN-?40ºS
20ºN-?30ºS
65ºN-70ºS
44ºN-47ºS
1.94-2.18
44ºN-35ºS
60ºN-46ºS
60ºN-35ºS
60ºN-61ºS
31ºN-37ºS
30ºN-40ºS
13.50-15.83
2.8-3.4
1.06-1.20
1.13-1.21
1.31-1.62
11.33-13.33
1.60-1.92
2.84-3.28
4.80-7.30
0.86-0.98
0.90-1.12
0.98-1.06
0.97-1.14
0.80-0.95
0.72-0.82
0.96-1.16
0.88-
1.12-1.28
1.32-1.60
0.90-1.16
0.74-0.86
0.80-0.94
0.72-0.86
0.94-1.20
0.76-0.86
0.76-0.90
0.70-0.80
3.00-3.68
3.44-3.60
>2000 2.28-2.52
49ºN-50ºS
700-1500
60ºN-50ºS
800-1500
1000-2000
32ºN-?Oº
800-
400-800
60ºN-?49ºS
100-300
60ºN-
60ºN-31
64ºN-45ºS
60ºN-35º5
60ºN-47ºS
700-2700
18ºN-20ºS
40ºN-35ºS
60ºN-57ºS
18ºN-64ºS
18ºN-35ºS
53ºN-40ºS
39ºS-50ºS
50ºS-74ºS
1.10-
1000-2500
0-1000
700-1000
600-1000
250-1500
400-2000
200-500
47ºS-60ºS
no
no ?60ºS
1.08-1.20
36º-50ºS
250-750
900-1500
7.5-9.4
38-64ºS
60ºN
?0-800
300-800
200-500
100-400
400-800
200-400
32ºN-?
60ºN-?
31ºN-?23ºS
2500-3500
no400-700
0-2000
0-
50-300
>2000
>1250
>2500
.no
no
no0-400
25-50
1.36-1
yes 1.00
-1500
>35001000-ºS
1.30
40ºS
1.5
200-400
yes
yes
gaussiyes ? 12500-2000
9.66
.56-1-
__ 60ºN-35ºS
400-700
>2000
no
64ºS-50ºN
slight 3.56-3.84
3.37-3.55
2.46-2.84
15.50-18.67
13.83-15.50
1.00-
1.76-2.10
3.08-3.52
4.70-7.60
4.60-7.60
0.85-1.02
0.94-1.16
0.99-1.08
0.95-1.18
0.90-
0.76-0.86
1.26-1.44
1.40-1.80
1.10-1.28
0.98-
0.76-0.88
0.80-1.02
0.68-0.84
0.96-1.18
0.74-0.92
0.76-0.90
0.75-0.80
0.92-1.06
3.16-3.72
3.84-4.16
1.30-
1.34-1.64
1.10-1.26
8.0-10.6
2.0-2.43
3.2-3.9
not 1.42
1.12
1.24
1.02
1.60
3.84-4.44
known
1.00-1.2ti
j '.32-'.60
Ostracoda 821
:l(lJ
"" ¿~'-'
¡s
0.0
(lJ0.0
u. -o.oE~~~O
::: -""
t':lo.o~
•.•
:l e::
'E(lJ
¿-c""'-'
Morphology e:: \,I~
e:: •.•.
:co•..e::'~'2 Z
Distribution
e::
t':l
\,1
e::.l:
(lJ.s~
t':l
._ Q.E(lJ0.0
t':l(lJ;E
1.66-2.10
1000-3000
no
no9.119
9.84
9.112
9.115
9.120
9.101
9.106
9.107
9.100
9.102
9.108
9.85
9.109
9.104
9.93
9.94
9.95
9.99
9.98
9.89
9.90
9.97
9.113
9.83
9.110
9.118
9.92
9.87 ---j
yes 9.111
9.105
9.91
9.114
9.~
----i9.1~
----,
----j 9.~
Mol/icía
10ºN-37ºS
-+-
1.40-1.55
no
1.05-1.4
43ºN-28ºS
--l46ºN-49ºS
24ºN-70ºS
40ºN-40ºS
34ºN-47ºS
53ºN-30ºS
32ºN-37ºS
60ºN-43ºS
26ºS-65ºS
1ºS-65ºS
2.88-3.16
2.51-2.92
2.30-2.64
1.42-1.52
1.72-1.88
3.48-3.88
2.76-3.00
3.08-3.28
2.20-2.46
2.56-2.72
no
1.58-1.78
1.50-1.78
1.36-1.56
1.40-1.80
1.56-1.72
1.06-1.36
20º-47ºS
no
48ºN-?
50ºN-?
no
no
40ºN-47ºS
es
1.54-2.02
49ºN-47ºS
0.98-1.18
0.84-1.08
1.34-1.40
1.35-1.40
1.62-1.86
es 3.04
49ºN-?
32ºN-?
28ºN-?
34ºN-?
?0-300
2.12-2.36
30ºN-?
400-1200
100-500
49ºN-40ºS
53ºN-40ºS
no
0.76-0.90
1.10-1.22
>2000
0-300
100-400
100-1000
1.30-1.50
1000-2000
1000-1500
2.20-2.40
2.44-2.52
1.20-1.37
1.88-2.02
1.54-1.86
0-300
700-1500
600-1250
100-1000
800-1500
700-2000
no
100-500
100-700
300-600
100-500
0-500
0-2000
18ºN-?
I+9.860-600
50-400
?-70ºS
>1500
>1250
0-500
yes
yes 1
1000-2000 >1500
40ºN-43ºS
47ºN-70ºS
52ºN-46ºS
34ºN-37ºS
45ºN-23ºS
60ºN-55ºS
18ºN-40ºS
32ºN-34ºS
60ºN-44ºS
18ºN-29ºS
1000-2000
32ºN-50ºS
1500-3000
2.78-3.00
2.48-2.80
1.60-1.94
>2500
1.20-1.40
2.20-2.88
2.40-2.56
1.42-1.62
yes ~_9.103
I~es~17
~ºN-56ºS53ºN-?0º
18ºN-31
+ka!!!pta
deepnoform
no
800-1500
ºS +--700-1500 1 -+-
0.86-1.04
1.46-1.70
1.08-1.30
1.50-1.70
1.26-1.36
1.06-1.26
1.80-2.02
2.28-2.96
2.72-3.16
3.44-3.92
2.40-3.28
2.48-2.72
3.08-3.48
2.28-2.52
3.32-3.60
2.68-2.88
2.20-2.66
2.88-3.04
3.12-3.60
3.28-3.48
1.30-1.85
2.94-3.32
3.08-3.20
2.97-3.44
1.80-2.04
1.98-2.20
1.86-2.20
1.52-1.84
1.88-2.32
1.50-1.86
1.72-1.90
1.20-1.52
1.56-1.72
1.88-2.10
1.92-2.34
1.24-1.48
1.5-1.8
1.2-1.7
for the genus.

822 Ostracoda
In the oceanic waters at high latitudes, planktonic Number of species

ostracod numbers are generally low in the surface 100 o 10 20 30 40
L I , ! J
m, except in late surnmer. They occur at maximum O
abundance (5-10 ind. per m-3) at depths of200-300 m.
Below 300 m, along with other taxa, their abundance
cteclines with increasing depth. However, ostracods
are usually ranked second in numerical abundance in
the macroplankton throughout the water column. But
because of their relatively small size, their contribu- 400
tion to the total standing crop in the water column is
relatively les s important. Their abundances increase
quite sharply within 100 m ofthe sea bed in the ben-
thopelagic zone (i.e., in the benthic boundary layer). E
Within the water column the halocyprids are bathy- -5

Q. 800
(j)
metrically zoned. Vertical ranges tend to be smaller at O
shallow depths than in deep water. Epipelagic species
inhabit the euphotic zone and the top of the seasonal
thermocline extending down to 250-300 m; the depth
at which the light field becomes totally symmetrical
1200
regardless of the sun's elevation. The shallow meso-
pelagic zone extends down from 250-300 m to the
depth at which the down-welling daylight becomes
dim relative to the bioluminesence produced by the
organisms themselves, at about 700 m in the clearest
oceanic water. The deep mesopelagic zone extends 1600
from there down to where daylight ceases to become
detectable by the animals, usually at around 1000 m.
The bathypelagic zone extends from 1000 m to around
2500-2700 m, where it gives way to the abyssopelagic
Fig. 4. Balhymelric profiles of lhe numbers of halocyprid species
zone. The boundary between the bathypelagic and
collecled along a transecl of slalions in lhe Northeaslem Allanlic
abyssopelagic zone does not seem to be related to a from [ION lO 60oN, approximalely along 20oW. Nole lhe sudden
specific physical para meter, but may be related to the decrease in species richness lhrougholll lhe waler column north of
depth at which fishes cease to be a major component 40oN; a similar pallem is lo be expecled in lhe SOlllh Allanlic.
ofthe midwater community (Angel, 1983). The lower Slalions norlh of 400N have open symbols (from Angel, 1993).
boundary ofthe abyssopelagic zone is at the top ofthe

benthopelagic zone about 100 m above the bottom in
the North Atlantic, but its height above the bottom in cornmon feature in several species is for adult females
some parts of the South Atlantic may be higher as a to have tails to their vertical distributions which extend
result of the greater incidence of benthic storms in the into far deeper water than either juveniles or adult
Argentinian Basin. There are few regions where males. For example, M. curta is predominantly a shal-
soundings exceed 6000 m where novel hadal cornmu- low mesopelagic subtropical species often migrating
nities may occur. These deep near-bottom zones have up into the neuston at night, but adult females are regu-
hardly been studied in the South Atlantic. lady caught in small numbers as deep as 2000 m.
Many species have bathymetric ranges that only rough- Angel (1979) analysed the spectra of carapace lengths
Iy correspond to this zonation. An individual species of halocyprid assemblages throughout the water co-
may migrate vertically between these zones either lumn offBermuda. By day, as the sampling was extend-
daily or seasonally. Ontogenetic migrations result in a ed into deeper water, the size range of the ostracod
species changing zones during its development. A assemblage progressively extended to include larger
Ostracoda 823
and larger animals, while the numbers caught declined. DORSAL

At night, vertical migration resulted in the bathymetric
E;:=3
distribution of sizes becoming more uniformo
Some species always appear to be non-migrants (e.g.,

left
Paraconchoecia inermis). Several species undertake
assymetric adductor shoulder
extensive diel vertical migrations, synchronised with gland muscles vault rostrum
the cycle of day and night. for example Paraconchoe-
cia spinifera and Conchoecissa imbricata migrate up male
gland
300-400 m at dusk - equivalent to 2xlO5 body
lengths. Along 200W in the northeastem Atlantic, the
greatest expression of diel vertical migration occurred
in the oligotrophic waters around 30oN, and become
less evident at both higher and lower latitudes.
Repeated sampling throughout much ofthe year in the right . LATERAL
assymetnc
southem Bay of Biscay indicated that both the extent
"~i3
of diel migrations and the number of animal s partici-
pating varied markedly with season (Angel, unpub-
lished). There are al so major changes occurring
throughout the year in the bathymetric ranges of the
species as a result of seasonal migrations.
VENTRAL
Fig. 5. Diagram of a "typical" halocyprid ostracod, showing the

Taxonomy main features of the carapace used in identification.
Morphology
The best general accounts of the morphology of

planktonic ostracods are by !les (1961) for the halop-
cyprid Boroecia (Conchoecia) antipoda and Cannon
(1940) for Gigantocypris (both based on Southem
Ocean material). This description focuses on halocy-
prids, drawing heavily on Iles (1961).
Fig. 6. Diagram of the respiratory flow through the carapace of

The ostracod body is completely enclosed within a Paraconchoecia ~pinifera, showing the location of the vibratory
bivalved carapace which is hinged along the mid-dor- plates and the inhalant and exhalant flows.
sal margino In most species, the carapace is developed
anteriorly into a rostrum, below which is a notch
known as the incisure, through which the exopodites of The carapace may also be omamented with sculpturing
the second antennae extend during swirnming (Fig. 5). which ranges from linear or concentric striations to
While the animal is swirnming and behaving normally, heavy reticulations (Fig. 9.56). Dorsally, the anterior of
the carapace is slightly agape, but it can be clamped the carapace is often vaulted over the powerfully mus-
shut by adductor muscles inserted close to the midpoint cular protopodites of the second antennae which pro-
of each valve. The shapes of planktonic ostracods range vide the propulsive power for swirnming; in a few spe-
from subglobular to subcylindrical. In many species the cies the vaults are sharp-edged (Fig. 9.104), developed
carapace is compressed laterally and raked anteriorly. into wings (Fig. 9.25), or armed with spines. An in):lal-
Spines or tubercles may omament the posterior dorsal ant respiratory flow of water enters through the gap
comers ancl/or the posterior ventral comers (Fig. 9.42). between the carapace val ves at the incisure, passes ayer
The rostrum may be extended as a spine, or spines may the body and discharges through a siphon on the poste-
be developed elsewhere on the carapace (Fig. 9.23). rior margin (Fig. 6).
824 Ostracoda
Gn the carapace are several large glands whose po si- are five terminal setae; a long terminal e-seta and four
tions are taxonomically important (Fig. 5). In halocy- shorter, thin-walled sensory seta e (a-d setae). In
prids there are "asymmetrical" glands which typically males, the second segment carries a seta which curls
open close to the posterior dorsal hinge on the left ca- around and braces the shaft of the capitulum; this seta
rapace val ve, and at the posterior ventral comer on the is probably analagous to the anteriorly pointing dorsal
right valve. However, in some of the genera the seta of females. The terminal setae include a- and c-
glands are displaced well away from these positions. seta e which are usually short, thin-walled and senso-
For example in the genus Metaconc!lOecia (Fig. 9.58) ry, and the b, d and e-setae are long and relatively
the gland on the left valve is on the rostrum and in thick walled. Typically the e-seta carnes an armature
Conchoecilla the gland on the right valve is just of spines or tubercles the number and arrangement of
beneath the incisure (Fig. 9.39). In males there is a which is species specific. The d and b setae may also
group of glands situated just below the posterior dor- carry some armature. During swimming, these setae
sal comer close to where the tips of the long setae of are carried splayed out in front of the animal so that
the sixth limb protrude through the carapace (see they interlock (Angel, 1970). In species of the sub-
below). The ventral margins ofthe carapace are lined family Halocyprinae the terminal setae on the first
with mucus-secreting glands. In many species there antennae also consist of one long and four shorter but
are accessory glands which produce biolumines- show little or no sexual dimorphism. In the
cence, located either on the anterior margin just below Archiconchoecinae there are six subequal terminal
the incisure, besides the exhalant siphon, or at the setae (Fig. 9.10 female D), whereas in the
tips of cara pace spines or tubercles. Euconchoecinae there are 20-250 terminal setae (Fig.
9.18 female D).
Adult halocyprids have seven pairs of segmented
limbs. The nomenclature for the setation of the limbs The second antennae are biramous (Fig. 7D). The
derived by Skogsberg (1920) is followed herein (Fig. large muscular protopodite occupies the anterior third
7). The most anterior pair of limbs are the first anten- or so of the carapace and provides the propulsive
nae or antennules. These lie either side of the frontal power for swimming. The exopodite usually consists
organ (or organ ofBellonci). The frontal organ is sex- of a long basal segment and eight short segments each
ually dimorphic in most species (Fig. 9.84D), and is carrying a long, feathered swimming seta. The endo-
usually divided into two sections, the stem (segment- podite is inserted on the inner face of the protopodite
ed in some species) and the capitulum, but the capi- just posteriorly to the insertion of the exopodite. In
tulum and stem are fused in females of some species Conchoecinae the endopodite is sexually dimorphic
(Fig. 9.109 female D). In the genus Bathyconchoecia (Fig. 7D, E). In females it is composed of two seg-
(not treated herein because this bathypelagic genus ments. The basal segment has an anterior swelling
has seldom been reported from the South Atlantic) the known as the processus mamillaris (Fig. 7D, E)
frontal organ is totally lacking in all species but one. close to its base on its anterior edge - a process which
In living specimens, the tip of the capitulum can usu- is characteristically absent in the Halocyprinae.
ally be seen either projecting forward just below the Distally is another process on which two setae are
rostral opening, or down-tumed across the inhalant inserted (a- and b-setae). The second segment carries
flow ofwater. In females the stem is unsupported, but five setae, the longest being the f-seta, with the h-, i-
in males the stem is firmly anchored by a seta from and j-setae being shorter, subequal, thin-walled and
the second segment of each the first antenna. sensory. In males the second segment of the endopo-
dite usually carries two short setae (c- and d-) near its
The first antennae are uniramous and sexually dimor- base, and five much longer setae subtenninally. At the
phic. In females of some species the two basal seg- base of the f-seta there is sometjmes an extremely
ments are fused (Fig. 9.115 female D), but in males short stubby e-seta. This segment also carries a hook
the segmentation is always clear. In the females of appendage (or c1asping organ). Examination of spe-
most species there is an anteriorly pointing dorsal seta cjes of Bathyconchoecia has made it clear that the
on the second segment (Fig. 9.36 female D). The two hook appendage is a modified thjrd segment with the
very short terminal segments carry a number of long shank of the hook carrying one of the longer setae.
setae. In females ofthe subfamily Conchoecinae there There is a marked asymmetry in the size and elabora-
Ostracoda 825
Adult female,
frontal organ and
first antenna
Arrangement of the limbs

within the carapace
of an adult female
Female, second Male, endopodite

antenna of second antenna Male,
(notice hook appendage) frontal organ and
first antenna
Mandible Fifth
limb
Male, sixth limb
o o
Male, caudal furca
with copulatory Female,
appendage sixth limb
Endopodite of
the maxilla
Fig.7. Diagram ofthe limbs ofhalocyprid ostracods also illustrating the nomenclature ofthe setae. Modified from I1es (1961).
826 Ostracoda
tion ofthe hook appendages between the left and right relative lengths can be useful in distinguishing species
antennae; the left hook appendage is larger and more (especially between some Archiconchoecia species).
elaborate than the right. This asymmetry is probably Laterally on an epipodial appendage on the base of
related to the offset of the copulatory organ on the both the fifth and the sixth limbs are three groups of
caudal furca (Fig. 7 J) and the asymmetry of the cara- long plumose setae. These setae lie along the flanks of
pa~e glands. In myodocopids the copulatory organ is the body within the carapace, and beat continuously
placed symmetrically and there is no asymmetry of driving the respiratory stream of water through the
the antennal endopodites. carapace. These groups of setae are sometimes
referred to as vibratory plates.
The third pair oflimbs, the mandibles (Fig. 7F), show
no sexual dimorphism. The exopodite is reduced to a The sixth pair of Iimbs once again show marked sex-
small segment often carrying a plumose seta. The ual dimorphism in the Conchoecinae (Fig. 7H, L).
endopodite is large and three-segmented, and armed The exopodite is usually four-segmented. Dorsally the
with a number offilamentous setae and terminal c1aw- segments may carry a few short setae, and ventrally
Iike setae. It is used in the manipulation of large food the first segment has several short setae. In females
items at the gap e of the carapace valves. The pro topo- the Iimb's three terminal setae are usually subequal
dite consists of two segments, the coxale which has a and only a little longer than the third segmento
toothed edge (termed the pars incisa) and the basa le Whereas in males the terminal setae are long and plu-
which carries three Iines of teeth (the tooth-lists) and mose, and their tips protrude from between the cara-
a masticatory pad. During feeding, these pads and pace val ves just below the posterior dorsal comer
tooth Iists of the two maxillae pivot against each where the male carapace glands discharge. Mating
other, reducing food items to a size that can be swal- behaviour has not been described in halocyprids, but
lowed. Poulsen (1973) based his revision of the secretions released onto these terminal setae may play
"super-genus" Conchoecia on characters of these some role in courtship. A useful character for distin-
mandibular tooth-Iists, but even with good oil-immer- guishing males from females is the very muscular first
sion microscopy these characters are difficult to dis- segment of the male's sixth Iimb, which is c1early vi-
tinguish. Moreover, some of the species Poulsen con- sible through the carapace of undissected specimens.
sidered to be congeneric differ markedly in other
characters. Consequently, attempts to identify species The seventh Iimbs of halocyprids are vestigial, each
via genera can, in several cases, be frustratingly con- consisting of two short segments with two unequal
fusing, and so for routine identifications it will often terminal setae. Most female myodocopids brood eggs
be pragmatic to ignore the genera. and developing young inside the carapace, to the side
and dorsal to the body, so their seventh limbs are well
The fourth pair of limbs, the maxillae (Fig. 7K), are developed. In Gigantocypris, for example, they are
not sexually dimorphic and show few useful interspe- multi-segmented and carry numerous setae. In live
cific differences. The terminal claw-like setae are also specimens they are constantly writhing around inside
used to manipulate food. The strong spinous setae on the carapace, probably keeping the inside of the cara-
the anterior margin of the protopodite of the fifth pair pace and the brood of eggs or larvae c1ean.
of Iimbs also assist feeding by pushing food up
towards the mouth. The caudal furca (Fig. 7J) in halocyprids is a paired
structure which is probably homologous with the telson
The exopodite of the fifth Iimb consists of three seg- of other crustacean groups (it has also been interpreted
ments (Fig. 7G). The first, in halocyprid genera like as being a uropod, and hence a true appendage).
Bathyconchoecia, carries an extremely long seta on its Typically an adult halocyprid has eight pairs of hook
dorsal surface. At this position in the cave-dwelling setae on its furca. The first instar has just two pairs of
genera, Deeveyae and Spelaeoecia (see Komicker and these hook setae and a new pair is added at each instar.
Iliffe, 1989) is a small articulated segment which car- However, in Archiconchoecia adults of some species
ies two or three long setae. So these dorsal setae prob- have fewer pairs and these species may have undergone
ably represent a vestigial endopodite. The third seg- neoteny and so may show a reduction in the number of
ment terminates in three long, c1aw-like setae whose juvenile instars. Recently Ikeda (1992) and Ikeda and
Ostracoda 827
Imamura (1992) have reported that Discoconchoecia pace (Fig. 8D, E). Those on the anterior margin
pseudodiscoplzora reared in culture undergoes seven release their secretions into the inhalant stream of
rather than six larval instars (the number of paired hook water entering the carapace. The secretion initially
setae on the caudal furca remains at two for the initial lights up the animal, but is then left behind in the
two instars after hatching). The furca is used in the exhalant plume as a c10ud which is about the same
manipulation for food items, particularly by arching size as the animal. The glands on the posterior margin
forward and forcing material away from the carapace fringe the enlarged gap between the carapace val ves
gape. It may also function as a means of levering the through which the exhalant respiratory outflow pass-
animal out of the grasp of a predator. es, and they discharge directly into the exhalant flow.
In a few species the luminescence is retained within
In the halocyprids the male copulatory appendage is the carapace usually within elaborate structures, such
located ventrally on the base of the furca on the left- as the long posterior dorsal spines, ventral tubercles
hand side (Fig. 7 J). It consists of an elongated flattened and prolongations of the rostra in Conclzoecissa
tube containing several obliquely arranged muscles. It imbricata, and the prolongations of the rostra and the
terminates in a hollow spine through which the semi- postero-dorsal corners in Conchoecilla daphnoides.
nal vesicle discharges. Adult females appear to mate Similarly, in Paraconchoecia spinifera luminescence
immediately on maturation because every mature is retained along the sharp-edged shoulder vaults.
female has a store of sperm at the base of the oviduct,
which appears as a white spot at the base of the furca. Outline c1assification
Colour is not used in the taxonomy of the planktonic The same c1assification of Ostracoda has been adopt-
ostracods beca use most identification is carried out on ed here as used by Athersuch et al. (1989) in a synop-
preserved material in which all colour has been lost. sis of British marine Cypridacea and Cytheracea. The
As in most pelagic groups, their coloration is correlat- subclass Ostracoda is divided into six orders:
ed with their day-time depth of occurrence. Shallow- Myodocopida, Platycopida, Podocopida, Bradoriida,
living species are generally translucent and virtually Leperditicopida, and Palaeocopida. The first three are
colourless. Many mesopelagic species have subtle extant, the others are extinct. The Myodocopida are
yellow, orange or red coloration associated with the the only ostracods considered here (bold text denotes
mouthparts and some of the carapace glands. The gut the taxa considered here; genera listed are those treat-
is often opaque orange-brown to black, and is the ed in the present chapter):
most obvious feature in daylight. In large deep-living
species red pigmentation becomes more pervasive Order Myodocopida Sars, 1866
(e.g. Alacia valdiviae and Conchoecissa anzetra); Suborder Halocypridina Dana, 1849
although juveniles still tend to be translucent. In deep- Family Thaumatocyprididae Mueller, 1906
living crustaceans such red coloration is produced by Family Entomoconchidae Brady, 1868
a carotenoid pigment originating from phytoplankton Family Halocyprididae Claus, 1891
and passed along the food-chain. The Macrocypridina Subfamily Deeveyinae Kornicker and Iliffe,
species are exceptional in that the adults are choco- 1987
late-brown, a feature which shared with a very few Subfamily Halocyprinae Dana, 1849 (genera
other pelagic taxa (e.g the mysid LongithoraxJuscus). Fellia, Halocypria, Halocypris)
Gigantocypris muelleri is also well coloured, appear- Subfamily Conchoecinae Müller, 1912 (genera
ing like a translucent orange-red ball; its deeper-living A lacia, Boroecia, Conchoecetta, Conchoecilla,
congener G. dracontovalis is much paler and tinted Conchoecia, Conchoecissa, Discoconchoecia,
more with purple. Gaussicia, Loricoecia, Macroconchoecia, Meta-
conchoecia, Mikroconchoecia, Mollicia,
Most halocyprids produce bioluminescence. Obtusoecia, Orthoconchoecia, ParaconcllOe(:ia,
Luminous secretions are released into the water from Paramollicia, Porroecia, Proceroecia, Pseudo-
glands situated either along the anterior edge of the conchoecia)
carapace just below the rostral incisure (Fig. 8A, B, C, Subfamily Archiconchoecinae Poulsen, 1969
G, H), or halfway up the posterior margin ofthe cara- (genus Archiconchoecia)
828 Ostracoda
A B
Conchoecia subarcuata Conchoecia magna Mikroconchoecia curta
D
~~JE Paramollicia rhynchena
Orthoconchoecia haddoni
Discoconchoecia elegans
~_:;;:t
Conchoecilla daphnoides
Conchoecissa imbricata
Paraconchoecia spinifera
Fig. 8. Skelehes (nol lO sea le) oflhe siles ofbiolumineseenee in various haloeyprids. From Angel (1993).
Subfamily Euconchoecinae Poulsen, 1969 Identification

(genera Bathyconchoecia, Euconchoecia)
Suborder Cladocopina Sars, 1866 Pelagic ostracods are collected live, and their taxonomy
Family Polycopidae Sars, 1866 is based extensively on their limb morphology, unlike
Suborder Myodocopina Sars, 1866 many benthic podocopids whose taxonomy, developed
Family Cypridinidae Baird, 1850 by palaeontologists, is based extensively on carapace
Subfamily Cypridininae Brady and Norman, characteristics (as for subfossil or fossil material). The
1896 (genera Macrocypridina, Gigantocypris) use of carapace characters gives continuity between the
Subfamily Azygocypridinae Komicker, 1970 systematics of living and fossil taxa, but confusion can
Family Philomedidae Müller, 1912 arise between species which are either close siblings or
Subfamily Philomedinae Müller, 1912 show marked convergence. In the planktonic species,
Subfamily Pseudophilomedinae Kornicker, the sole use of carapace characters can lead to confu-
1967 sion between some closely related species, but an expe-
Family Cylindroleberidae Müller, 1906 rienced field ecologist will be able to identify most
Subfamily Cylindroleberidinae Müller, 1906 adult specimens on the basis of external characters such
Subfamily Cyclasteropinae Poulsen, 1965 as size, shape, omamentation and the positions of the
Family Sarsiellidae Brady and Norman, 1896 gland openings. Identification of juvenile instars is
Family Rutidermatidae Brady and Norman, 1896 much more of a problem, and for the earliest three or
four stages it is largely guesswork.
Ostracoda 829
Distinguishing the halocyprids from other myodoco- not find this a simple task without resorting to dissec-
pids is readily done in shallow-living species on the tion, especially when dealing with species which are
basis of the absence of obvious light receptors. In most opaque (e.g., Halocypris injlata). Any specimen with
myodocopids living at depths shallower than about clear male secondary sexual characters is adult: i.e.,
2000 m, compound eyes are clearly visible through the first antenna with armature (Fig. 7C), well developed
transparent carapace, situated close to the insertion of hook appendages on the endopodites of the second
the adductor muscles. The adults of the mesopelagic antennae (Fig. 7E), sixth limb with a strongly deve-
genus Macrocypridina are immediately recognisable loped basal segment visible through the side of the
on the basis of the dark chocolate-brown colour of the carapace (Fig. 7L), and a fully-developed copulatory
carapace, with an unpigmented window in each valve organ (Fig. 7J). Many adult females have well-deve-
overlying the compound eyes. The large globular loped ovaries, which show up as white opaque patches
bathypelagic species of Gigantocypris lack compound on either side of the abdomen, and an iridescent patch
eyes, but have large and elaborate naupliar eyes with of stored sperm at the base of the caudal furca. A cha-
mirrored reflectors (Land, 1978). Many abyssal and racter that is definitive for most species is the caudal
trogloditic (cave-dwelling) myodocopids have also lost furca carrying eight pairs of hook setae, however, some
both compound and naupliar eyes. Halocyprids lack adults of some Archiconchoecia species have fewer
obvious visual receptors, but must have some light sen- pairs. Juvenile characteristics incIude: a lack of
sitivity because they undertake diel vertical migrations development of the fifth and sixth limbs, and small
synchronised with the day/night cycle. The frontal but distinct swellings on the tips of the sensory setae
organ, which is generally considered to be chemosen- of the first and second antennae, and in most species
sory, has nerve tissue with an ultrastructure that is sug- the occurrence of fewer than 8 pairs of hook setae on
gestive of some light sensitivity (Anderssen, 1977). the caudal furca. Specific identification of immature
stages is difficult because they lack many ofthe diag-
Order Myodocopida nostic characters and some species undergo large allo-
Carapace often relatively large, up to 30 mm, with or metric changes in carapace.
without an anterior rastral incisure. Ventral margin
often convexo Calcification of outer lamellae of the Although the majority of species can be identified
carapace usually weak or even absent. Omamentation without resorting to dissection, there are several com-
usually weak or lacking. Adductor muscle scars var- plexes of sibling species, including some of the more
ied. Between five and seven pairs of appendages. abundant taxa, for which accurate identification
Antenna biramous; exopodite with 7-9 articulated requires some dissection. Before attempting any dis-
podomeres, endopodite reduced with few podomeres. sections measure the carapace length, height and
Maxilla without a branchial plate. Fifth limb with an breadth, and sketch the outline. Note any special cha-
epipodial appendage (brachial plate, Fig. 7G). Sixth racteristics ofthe carapace, and most importantly try to
and seventh limbs varied (Myodocopina and locate whether the carapace glands are asymmetrical
Halocypridina) or absent (Cladocopina). Furca trian- and where exactly they open. Identification of some
gular, lamelliform and armed with strong marginal species requires recognition of whether there are
chelate setae (Fig. 7J). The anus opens anteriorly and accessory glands at various positions around the cara-
ventrally to the furca (Komicker, 1975). pace valves, but in most species these can only be
Marine organisms, many of which are truly pelagic in seen under high power. Dissection is simple. Lay the
the open ocean, while others are essentially benthic specimen on its back and, with afine needle, slice
swimmers and may be encountered in shallow sublit- through the body wall and adductor muscles down
toral waters. each side. The main part ofthe body with all the limbs
Only two suborders of Myodocopida have oceanic can then be hooked out. The carapace can be mount-
planktonic representatives (see outline classification ed either permanently or temporarily, with or without
above). staining, for closer examination ofwhere the carapace
glands are situated. The limbs can then be picked off
Suborder Halocypridina in order, but for most purposes only the first antennae
When identifying halocyprids the first task is to deter- together with the frontal organ and the left-hand se-
mine whether the specimen is adult. The novice will cond antenna will be needed.
830 Ostracoda
Family Halocyprididae men may be10ng. A good initia1 guide is provided by

The four subfamilies of Halocyprididae can be distin- measuring the carapace 1ength, as is the upper depth
guished on the basis of the numbers of terminal setae of occurrence (lower depths can be misleading
on the first antennae and the presence or absence of a because most c10sing nets willleak after c10sure).
Qrocessus mamillaris on the endopodite of the second
antenna. The Conchoecinae have 5 terminal setae on a. Carapace with posterior dorsal corner acutely
the first antennae and a processus mamillaris on the angled and developed into a long wedge, asym-
endopodite of the second antennae (Fig. 7D, E). metrical gland on right valve opens on the ante-
Halocyprinae also have 5 terminal setae on the first rior margin just below the rostral incisure.
antennae but lack a processus mamillaris. Archi- Genus Conchoecilla. Two species: C. daphnoides
conchoecinae have 6 subequal terminal setae and no C1aus, 1890 (Fig. 9.40), one of the longest species
processus mamillaris, whereas the Euconchoecinae which has a widespread distribution, and C. chuni
have >20 terminal setae and no processus mamillaris. (Müller, 1906) (Fig. 9.39), which is restricted mostly
to high latitude waters. Müller (1906) described a
Subfamily Halocyprinae small form (c. daphnoides minor), but since there
The subfamily Halocyprinae consists of three genera have been no further reports of it, its status remains
erected by Poulsen (1969): Fellia, Halocypria and uncertain.
Halocypris. Fellia inc1udes three species with spine
andJor rounded processes on the carapace. F. cornuta b. Carapace with highLy reticulate sculpture with
(Müller, 1906) (Fig. 9.23) is a 1arge species with large spines at the posterior dorsal corner of both
lateral spine processes on the shoulder vaults. F. valves and tubercles at the posterior ventral
bicornis (Müller, 1906) (Fig. 9.22) is a smaller spe- corner, the armature of the male antennular e
cies with a rounded swelling developed just below seta are paired needle-Iike spines (except in C.
and anterior to the midline of each carapace valve. F. imbricata).
dispar (Müller, 1906) (Fig. 9.24) is another large, but Genus Conchoecissa. Five species: C. imbricata
rarely caught, species originally described as a sub- (Brady, 1880) (Fig. 9.42) which is a common middle
species of F. cornuta, with only lateral protuberances latitude mesope1agic diel vertical migrant with long
developed on the shoulder vaults. Halocypria, mono- symmetrical extensions to the rostrum. C. ametra
specific for H. globosa Claus, 1874 (Fig. 9.19), has a (Müller, 1906) (Fig. 9.41) - a deep mesopelagic spe-
globous carapace 1acking spines and protuberances cies with asyrnmetrical extensions to the rostrum. C.
and has a c1ear1y developed rostrum and incisure. plinthina (Müller, 1906) (Fig. 9.43) - a very 1arge
Halocypris inc1udes two species: H. injlata (Dana, deep-living species with small symmetrical develop-
1852) (Fig. 9.20); and H. pelagica C1aus, 1890 (Fig. ments of the rostra. C. symmetrica (Müller, 1906)
9.21), in which the rostrum is barely deve10ped so (Fig. 9.45) - a high latitude deep-living, deep-bodied
they lack an obvious incisure. The two species are mesopelagic species with symmetrical rostra. C.
only readi1y distinguishab1e on the basis of size. squamosa (Müller. 1906) (Fig. 9.44) - a poorly
known deep mesopelagic species with normal rostra.
Subfamily Conchoecinae
The subfami1y Conchoecinae was treated as being c. Large deep-Iiving species with highly orna-
monogeneric by Müller (1906) and Skogsberg (1920), mented, relatively elongate carapaces, lacking
but subdivided into species-groups. Although the tubercles at the posterior ventral corners.
recent trend has been to recognise these species- Genus Macroconchoecia. There are three large bathy-
groups as genera, 28 of which are known from the pelagic species, the largest M. macroreticulata (Ellis,
South Atlantic waters, some are still c1early heteroge- 1984) (Fig. 9.55) being the deepest bodied. A species
neous and probab1y need further subdivision before with rows of spines omamenting the reticulations
the systematics ofthe subfamily can be considered to around the posterior margins of the shoulder vaults -
be stab1e. So, for several of the genera it is impracti- M. spinireticulata (Ellis, 1984) (Fig. 9.57), and a spe-
cal to give a meaningful description of the generic cies with the ventral margin of the carapace almost
characteristics. Here a series of characters is used to parallel with the dorsal margin - M. reticulata
narrow down the range of species to which a speci- (Müller, 1906) (Fig. 9.56).
Ostracoda 831
d. SmaIl globous species with reticulate cara pace large abyssopelagic species has been found in the
sculpturing distinguished by the thin-waIled Southem Ocean, although in very deep samples from
setae of the antennae being bifurca te. the North Atlantic female specimens which may
Genus Mikroconchoecia. Four species the commonest belong to the same species have been found. The
being M. curta (Lubbock, 1860) (Fig. 9.78) which is unique feature ofthis species is the trifid antennular a
<1 mm long, has the asymmetrical gland on the right seta. There are three small «1 mm) globous species.
carapace valve opening only just above the posterior The shallow mesopelagic to epipelagic tropical spe-
ventral comer. The males have about eight small pegs cies, M. kyrtophora (Müller, 1906) (Fig. 9.66) in ven-
as armature on the antennular e seta and a normal tral profile has clearly visible shoulder vaults, and the
sized copulatory appendage. M. echinulata (Claus, male antennular e seta has a unique armature of peg
1891) (Fig. 9.79) is slightly smaller, with the right spines. M. nasotuberculata (Müller, 1906) (Fig. 9.69),
asymmetrical gland opening midway up the posterior also a tropical to subtropical mesopelagic species, is
margino The males have only five pegs as armature on distinguished by having the left gland opening close
the antennular e seta and an extremely large intromit- to the tip ofthe rostrum, but most readily by the deve-
tent organ. M. acuticosta (Müller, 1906) (Fig. 9.77) is lopment of protuberances near the posterior dorsal cor-
a large species (> 1 nun) shaped similarly to M. curta ner of the carapace. M. teretivalvata (Iles, 1953) (Fig.
but with humped shoulder vaults. The male antennu- 9.75) is a shallow mesopelagic species occurring
lar e seta has an armature of 12 pegs. M. stigmatica often abundantly in subtropical to temperate waters, it
(Müller, 1906) (Fig. 9.80) is also > 1 mm in length and has a broad carapace which curves smoothly in ven-
tapers anteriorly. The right asymmetrical gland opens tral aspect. The male has 10 pairs of spines in the
midway up the posterior margino armature of the antennular e seta. M. arCl/ata
(Deevey, 1978) (Fig. 9.59) is another small deep
e. SmaIl species with the gland on the left valve bathypelagic species which is deep-bodied relative to
opening on the rostrum or on the anterior third the other species and the carapace does not taper ante-
of the dorsal carapace margino riorly. The thin-walled setae on the male second
Genus Metaconchoecia. This difficult genus contains antennal endopodite have side branches. AIl the
many sibling species which are distinguished mostly remaining species will be difficult to identify without
by subtleties of carapace shape. There are four species experience, good material and reference to Gooday
in which the asymmetrical gland on the right carapace (1981). The largest species (> 1.3 mm) is M. skogsber-
valve opens on a prominence towards the top of the gi (Iles, 1953) (Fig. 9.74), a mesopelagic species
dorsal margino M. pusilla (Müller, 1906) (Fig. 9.71) is abundant in the Southem Ocean. M. discoveryi
a small «1 mm) deep mesopelagic species which is (Gooday, 1981) (Fig. 9.61) has the opening of the left
often abundant at middle to high latitudes. M. mac- asymmetrical gland > 15% of the carapace length
romma (Müller, 1906) (Fig. 9.68) is larger (> 1 mm) behind the tip of the rostrum. It is al so a slim species
with the gland on the left valve opening further back with the carapace breadth <40% its length, and the
along the dorsal surface and tends to be more abun- antennular e seta armature consists of 11 pairs of
dant at low latitudes. M. isocheira (Müller, 1906) spines. M. fowleri (Gooday, 1981) (Fig. 9.62) is
(Fig. 9.65) is a small Southem Ocean endemic, occur- slightly longer and broader than the previous species,
ring at mesopelagic depths in which the mal e is imme- and has the left asymmetrical gland opening 12-15%
diately recognisable by having only a single row of ofthe length behind the rostrum. The male antennular
stout spines as armature on the antennular e seta. A e seta has an armature of 13 pairs of spines. There are
much larger (> 1.5 mm) deep bathypelagic species, M. two species with carapaces that strongly taper anteri-
glandulosa (Müller, 1906) (Fig. 9.63) also has the left orly. The largest (1.06-1.26 mm) known only from
asymmetrical gland set well back, but the carapace is southem latitudes is M. wolferi (Gooday, 1981) (Fig.
almost parallel-sided in ventral aspect and it uniquely 9.76) and has 13 pairs ofspines in the armature ofthe
has a group of large gland cells just below the rostral male antennular e seta. M. obtusa (Gooday, 19&1)
incisure on both val ves. Another species that might be (Fig. 9.70) is shorter (0.91-1.06 mm) with a rather
confused with M. macromma in that it has the left bluntly rounded anterior edge to the carapace below
asymmetrical gland set well back is M. /llllata the rostrum (see M. acuta below) and has 12 pairs of
(Deevey. 1982) (Fig. 9.67). Only the male of this spines in the armature of the male antennular e seta.
832 Ostracoda
Species that are less tapered anteriorly inelude a very with characteristically shaped anterior margins to the
broad species M. inflata (Gooday, 1981) (Fig. 9.64) in carapace below the incisure and numerous small
which the carapace breadth is often >50% the length. spines around the posterior dorsal comer. Alacia alata
Ihe smallest species «0.87 mm) M. rotundata (Müller, 1906) (Fig. 9.25) has the shoulder vaults
(~üller, 1906) (Fig. 9.72) has only 8 pairs ofspines in developed into winged extensions and is abundant at
the armature of the male antennular e seta. M. austra- mesopelagic depths, especially in upwelling regions.
lis (Gooday 1981) (Fig. 9.60) is superficially like C. Loricoecia acutimargil7ata (Chavtur, 1977) (Fig.
teretivalvata, but is slimmer and has II pairs of spines 9.52) is a large abyssopelagic species originally
in the arrnature of the male antennular e seta. M. sub- described from the North-east Pacific but appears to
inflata (Gooday, 1981) (Fig. 9.73) is anteriorly broad- be cosmopolitan in very deep water. It is distin-
ened, particularly in the male, which also has 11 pairs guished from the other species with sharp shoulder
of spines in the arrnature of the antennular e seta. M. vaults by the depth of its occurrence and by having
acuta (Gooday, 1981) (Fig. 9.58) is shorter and has a accessory glands probably secreting bioluminescence
distinctive acute edge to the anterior edge of the cara- overlying the right asymmetrical gland.
pace below the rostrum. It has 10 pairs of spines in the Paraconchoecia spinifera (Claus, 1890) (Fig. 9.104)
armature of the mal e antennular e seta. Ihese species is a medium-sized, vertically migrating species which
can be consistently sorted on the basis of their cara- is abundant at shallow mesopelagic day-time depths.
pace shape, and despite some overlap, show differen- It has a spine at the posterior dorsal comer of the right
ces in their vertical ranges at any one position. carapace valve only and the female has a row of long
hairs along the leading edge of the antennular e seta.
f. Species with large and distinctive compound Paraconchoecia dasyophthalma (Müller, 1906) (Fig.
accessory glands opening about midway along 9.94) and P. mesadenia (Ellis, 1985) (Fig. 9.101) are
the ventral edge of the cara pace. a1so readily distinguishable because the right asym-
Genus Gaussicia. Ihis genus was revised by Gooday metrical glands open well anterior of the posterior
(1976). Ihe largest (3 mm) species G. gaussi (Müller, ventral comer. In the latter, an abyssopelagic species,
1908) (Fig. 9.49) is bathypelagic and the accessory the gland opening is displaced more anteriorly than in
glands open posteriorly to the midpoint. Ihe ventral the former, deep bathypelagic species. Juveniles of
carapace margin is nearly parallel to the dorsal edge. Pselldoconchoecia concentrica (Müller, 1906) (Fig.
Whereas in the next largest species G. incisa (Müller, 9.119) have a fringe of spines around the shoulder
1906) (Fig. 9.50) the carapace tapers anteriorly in the vaults which are lost in the late juvenile and adult
female. Ihere is a marked sexual disparity in carapace instars.
length (female 2.45-2.8 mm, males 1.8-1.9 mm). Ihe
female antennular a-d setae are <50% the e seta, and h. Species with a c1ear row of accessory glands
in the males the a seta is relatively long but the c seta c10se to the posterior ventral comer of the left
is short. G. edentata (Müller, 1906) (Fig. 9.48) is carapace valve.
smaller and is notable for all the female antennular A unique feature of COllchoecia lophllra Müller, 1906
setae being subequal and the male antennular c and e (Fig. 9.34), a medium-sized deep mesopelagic spe-
setae being exceptionally long. At low latitudes, a still cies. These accessory glands are clearly seen in juve-
smaller species occurs - G. subedentata (Gooday, nile instars making them readily identifiable.
1976) (Fig. 9.51) which has similarly long antennular
setae but the e seta armature in the male consists offar i. Species with the asymmetrical gland on the left
fewer pairs of spinules. cara pace valve opening well anterior ofthe pos-
terior dorsal comer.
g. Species with sharply edged shoulder vauIts. Ihe largest of these species, which has the gland
Ihere are six species belonging to different genera opening almost half-way along the carapace, is
which have this elear characteristic. Boroecia antipo- Paraconc1lOecia dorsotllberculata (Müller, 1906)
da (Müller, 1906) (Fig. 9.30) is one of a sibling pair (Fig. 9.96), a medium-sized bathypelagic species
of bipolar species, and indeed originally described as which is unusually slim in ventral aspecto Another
a subspecies of the Arctic species B. borealis. It is an smaller mesopelagic species that also has the gland
abundant large mesopelagic species at high latitudes opening well forward is Proceroecia convexa
Ostracoda 833
(Deevey, 1977) (Fig. 9.114). The female has carapace larger species - Paraconchoecia mamillata (Müller,
val ves with well rounded posterior dorsal comers and a 1906) (Fig. 9.100) tends to occur deeper and reaches
rather blunt frontal organ. In the male the armature of higher latitudes than its smaller sibling P. nanomamil-
the antennular e seta consists of about 18 pairs of spine lata (Deevey and Brooks, 1980) (Fig. 9.102). Both
teeth. Porroecia pselldoparthenoda (Angel, 1972) (Fig. these species have a criss-cross partern of striations
9.111) is deeper-bodied and is shallower-living. Note omamenting the carapace.
the well rounded posterior dorsal comer of the female
carapace and the square comer in the male. The arma- I. Species with the posterior dorsal carapace cor-
ture on the male antennular e seta has pairs of small ners forming an acute angle but not developed
stout spines distally that become altemate proximally into spines, the asymmetrical gland on the right
and then become a single roW. This species is quite valve opens posterio-ventrally.
similar to the shallow mesopelagic species Porroecia Conchoecetta aCllminata Claus, 1890 (Fig. 9.31) is a
parthenoda (Müller, 1906) (Fig. 9.109) in which the medium-sized shallow mesopelagic species restricted
gland opens closer to the posterior dorsal comer, but is geographically to latitudes <35°. The female is readi-
otherwise quite similar to the previous species. Finally, Iy distinguished from its more tropical congener C.
in this group is Proceroecia brachyaskos (Müller, giesbrechti (Müller, 1906) (Fig. 9.32) by its pointed
1-906) (Fig. 9.113) which is a rather small deep meso- rostra. The males are easily distinguished on the basis
pelagic species with a carapace that tapers anteriorly. In of the armature of the antennular e seta. A bluntly
the female the thin-walled sensory setae are over half acute posterior dorsal comer also occurs in the large
the length of the e seta, and in the male the a to d anten- rugged species Orthoconchoecia atlantica. A charac-
nular setae are uniquely short relative to the e seta. teristic of its genus is having a c or d seta on the anten-
nal endopodite which is as long or longer than the seg-
j. Species with carapace with welI rounded poste- ment on which is inserted.
rior dorsal comer and curved posterior margino
Obtllsoecia antarctica (Müller, 1906) (Fig. 9.86) is an m. Other species with long c or d setae on the
abundant shallow mesopelagic cold-water species antennal endopodite.
occurring almost exclusively south of the Antarctic Genus Orthoconchoecia. Three other species of the
Convergence. Like Boroecia antipoda, this species is genus occur in the South Atlantic. Two have small but
one of a bipolar pair of sibling species and was origi- distinct spines at the posterior dorsal comer of the cara-
nally described as a subspecies of the Arctic species pace valves. o. secernenda (Vavra, 1906) (Fig. 9.90)
Obtllsoecia obtllsata (Sars, 1866). The carapace and o. bispinosa (Claus, 1890) (Fig. 9.88) are readily
lengths of the two sexes are very disparate. A unique distinguished by their different colouring when live.
character of the males is that the hook appendage on The larger o. secernenda being rather opaque and more
the left antennal endopodite is very large but the right intensely coloured than the smaller translucent paler-
hook appendage is reduced to a straight process. This coloured o. bispinosa. Juveniles of o. secernenda have
species also has an unusually large copulatory much larger shoulder vaults and even in the adults the
appendage. Paraconchoecia inermis (Müller, 1906) vaults tend to be more powerfully developed. The third
(Fig. 9.99) is a much more cylindrical species with the species, o. haddoni (Brady and Norman, 1896) (Fig.
one of the largest length:height ratios. This medium- 9.89) lacks the spines at the posterior dorsal comer and
sized species seldom occurs outside the depth range of shows a large sexual disparity in size, the females being
300-600m and is a non-migrant. The armature of fine much larger. Otherwise the limb structure is very simi-
hairs on the male antennular e seta is a character lar between all three species. There are twO races of o.
shared only by Discoconchoecia spp., Paraconchoe- haddoni, with the large form occurring at high latitudes
cia dorsotllberclllata, and the P. mamillata siblings. and a smaller form at low latitudes where it is often
associated with upwelling regions.
k. Species with the right asymmetrical gland
opening on a tubercle over half way up the pos- n. Species with the asymmetrical gland on the right
terior margino valve opening some way up the posterior margino
There are two deep mesopelagic to bathypelagic spe- Two ofthese species have spines at the posterior dor-
cies which are most readily distinguished by size. The sal comer ofthe right carapace valve and are medium-
834 Ostracoda
sized mesopelagie speeies. Males of Paraconchoecia o. Species with a spine at the posterior dorsal cor-
aequiseta (Müller, 1906) (Fig. 9.91) are only readily ner of the cara pace with the asymmetrical
distinguishable from P. hirsuta (Müller, 1906) (Fig. glands in the normal positions and without
9.98) on the basis of the rather longer antennular e other elaborations of the carapace.
s~ta and the hirsute antennal b seta in the latter spe- In three speeies of Paraconchoecia: P. allotheriu/11
eies. No adequate eharaeter for separating the females (Müller, 1906) (Fig. 9.92), P. oblonga (Claus, 1890)
has yet been established. Two ofthe other species lack (Fig. 9.103), and P. echinata (Müller 1906) (Fig.
spines at the posterior dorsal comer but have deep 9.97) there is a relatively small anterior taper in the
longitudinal striations running parallel to the ventral shape of the earapaee. Females have fine hairs along
edge of the carapaee. In Paraconchoecia dentata the anterior margin ofthe antennular e seta and a fron-
(Müller, 1906) (Fig. 9.95), the smaller speeies, there tal organ with a clearly differentiated capitulum whieh
are 2 to 4 striae, and in Paraconchoecia cophopyga is slightly down-tumed. The armature of male anten-
(Müller, 1906) (Fig. 9.93) there are 5 or more striae. nular e seta consists of a double row of spines. P.
In both speeies the eapitulum of the frontal organ is echinata is distinguished by its larger size, the rather
poorly differentiated from the stem and is only very square end of its earapace, and in the males by the
slightly down-tumed. The males are uniquely differ- pateh of large spikes on the antennular b seta. P. allo-
entiated by the b seta on the first antenna being long- therium is readily distinguished from P. oblonga by
er than the other setae, espeeially the e seta. The arma- its mueh greater slirnness and being more parallel-
ture of the e seta is a double row of blunt, flattened sided both in ventral and lateral aspeet. The other two
spmes. speeies - Discocol1choecia elegans (Sars, 1866) (Fig.
9.47) and D. discophora (Müller, 1906) (Fig. 9.46)
There is a trio of similar speeies: Paramollicia dicho- have carapaces which strongly taper anteriorly. The
toma (Müller, 1906) (Fig. 9.105), P. plactolycos frontal organs are straight with little differentiation
(Müller, 1906) (Fig. 9.107), and P. major (Müller, between stalk and capitulum. There are no anterior
1906) (Fig. 9.106) that have this eharaeteristic. They hairs on the female antennular e seta. Males are
lack spines at the posterior dorsal comer, and have the instantly attributable to this genus on the basis of the
posterior margin smoothly curving into the ventral antennular e seta armature eonsisting of an oval pad of
margin and a strong anterior taper. The striations at hairs. Females of D. discophora are deseribed in the
the anterior end of the carapaee intereept the ventral literature as having a longer frontal organ and a dorsal
margin obliquely. The genus Paramollicia is typified seta on the first antenna. However, D. elegans, which
by having two groups of aecessory glands on both is often one of the most abundant speeies and ranges
val ves of the earapace, one group opening at a level from high to low latitudes, shows clinal shifts in these
just dorsal to the opening of the asymmetrieal gland eharaeters. At present the best distinction between
on the right val ve, the other close to the posterior dor- these speeies are differences in carapace shape, but
sal comer. The eapitulum of the frontal organ is dif- thorough investigation of this genus will no doubt
ferentiated from the stem and down-tumed. The end show that is D. elegans is a species eomplex in which
of the male first antennal e seta is flattened into a D. discophora may be just one of a number of sib-
sword-shape, and the armature eonsists of pairs of lings.
winged, stubby spines. P. major (Fig. 9.106), original-
Iy deseribed as a large subspecies of P. plactolycos p. Species with the asymmetrical gland on the
(Fig. 9.107), is readily distinguished on the basis of its right valve opening at a notch just anterior to
larger size, the shape of the anterior margin of the ca- the posterior ventral corner.
rapace when viewed ventrally, the eurve of the hook These are all fairly small speeies and three are often
appendage and the hairs on the b seta on the endopo- abundant at shallow mesopelagic depths. Three eom-
dite of the male second antenna. P. dichotoma (Fig. pose the "procera" complex, with the female having
9.105) has simi lar sized glands cells in the posterior and almost straight and undifferentiated frontal organ
dorsal group of aeeessory glands, whereas in P. plac- with the sensory setae on the female first antenna
tolycos half of the glands eells are twiee as large as the being over half the length of the e seta. These three
others. speeies are best differentiated on the basis of size.
Proceroecia macroprocera (Angel, 1971) (Fig.
Ostracoda 835
9.115) being the 1argest, P. procera (Müller, 1894) leading edge of the female antennular e seta, and
(Fig. 9.117) being interrnediate, and P. microprocera arrnature of the male antennular e seta consisting of
(Angel, 1971) (Fig. 9.116) the smallest. Where they paired sharp spines.
co-occur their centres of abundance are arranged in
size order, with the largest species occurring deepest. s. Speeies with earapaees tapering anteriorly,
P. vitjazi (Rudjakov, 1962) (Fig. 9.118) is exception- laeking postero-dorsal spines, and typified by
al in that it is a larger, abyssopelagic species and in having an aeeessory gland opening on the side
some specimens there is a clear v-shaped scu1pturing. of the right asyrnmetrieal gland.
It is unlikely to be encountered at depth <2000 m. Genus Loricoecia. The carapace shape of these two
species is characteristic, but the diagonistic character
q. Species with large aeeessory glands arranged - the lateral accessory gland, is difficult to see. The
along the posterior rnargins of left earapaee two species 1. loricata (Claus, 1894) (Fig. 9.54) and
valve. 1. ctenophora (Müller, 1906) (Fig. 9.53) are often
Genus Alacia. One species in this heterogeneous subdominants at mesopelagic depths. 1. loricata has
genus has already been described - A. alata, which is oblique striations in the anterior region of the cara-
distinguishab1e on the basis of the winged extension pace, whereas the larger species 1. ctenophora is
of the shou1der vault. It also includes the largest halo- almost totally smooth.
cyprid species which is A. valdiviae (Müller, 1906)
(Fig. 9.29). When alive and for some time after pres- 1. Species with large, well developed shoulder
ervation this species is coloured bright red. It has sin- vaults whieh in juvenile stage earry a fringe of
gle accessory glands postero-ventrally and postero- spines.
dorsally on each valve. It has few other characters in Pseudoconchoecia concentrica (Müller,1906) (Fig.
common with the three other species with these addi- 9.119) often has a clear pattero of concentric striations
tional accessory glands. Two of these species are which render the cara pace more opaque than most
Southero Ocean endemics: A. hettacra (Müller, 1906) other species. The shape of the carapace viewed ven-
(Fig. 9.27), and A. belgicae (Müller, 1906) (Fig. 9.26). trally is characteristic for this species. Poulsen (1973)
A. hettacra is the smaller and more commonly linked this species with P. serrulata (Claus, 1874)
encountered species which has two accessory postero- (Fig. 9.120) on the basis ofthe structure ofthe basale
ventral glands on the left valves only. Poulsen (1973) of the mandible, but otherwise these two species have
describes A. belgicae (Fig. 9.26) as having four acces- little in common.
sory postero-ventral glands on the left valve, but Kock
(1992) illustrates up to nine. Both species have unusu- u. Species with double row of triangular teeth.
al peg-like teeth as arrnature on the male e seta. The Pseudoconchoecia serrulata (Claus, 1874) (Fig.
remaining poorly-known species ascribed to this 9.120) is another Southero Ocean endemic, which can
genus by Poulsen (1973) is A. leptothrix (Müller, be the dominant at shallow depths at high latitudes. Its
1906) (Fig. 9.28). Its carapace shape and longitudinal rather stout carapace shape is diagnostico
striae are more reminiscent of Paramollicia, but it has
one postero-dorsal and two postero-ventral glands on v. Large species (ea. 3 rnrn) with c1ear oblique stri-
each carapace val ve. ations with ventral edge either parallel to the
dorsal or the rnaxirnurn height near the rniddle.
r. Species with c1ear longitudinal striae along the Genus Mollicia. The species in this genus are deep
ventral region of the earapaee. mesopelagic to bathypelagic. M. tyloda (Müller,
In addition to various species described above on the 1906) (Fig. 9.85) is uniquely characterised by its
basis of other characters viz.: Alacia leptothrix (Fig. sculpturing pattero with a deeply incised striation
9.28), Paraconchoecia cophopyga (Müller, 1906) cutting across many of the obliques striae. M. kampta
(Fig. 9.93) and P. dentata (Fig. 9.95); Paramollicia (Müller, 1906) (Fig. 9.83) is the smallest of the spe-
rhynchena (Müller, 1906) (Fig. 9.108) is an abundant cies and the female can be distinguished by the rather
mesopelagic species at mid-latitudes which is distin- straight oblique ventral edge of the carapace irnme-
guished from the others by the clearly differentiated diately anterior to the opening of the right syrnmetrical
capitulum ofthe frontal organ, the lack ofhairs on the gland, and towards the end of the b seta of the male
836 Ostracoda
antennule is a small bulging pad. M. mollis (Müller, magna, but larger and without the square ending to
1906) (Fig. 9.84) is slightly larger and the b seta on the rostrum, is C. parvidentata Müller, 1906 (Fig.
the male antennule is bare, whereas in both the other 9.37). This species appears to be most abundant at
two species it carries long hairs. The e seta of the middle latitudes in the South Atlantic at mesopelagic
female antennule has quite long spinules along its depths. Like C. hyalophylium it has two accessory
I~ading edge. M. amblypostha (Müller, 1906) (Fig. glands adjacent to the right asymmetrical gland. In the
9.81) has a very differently shaped hook appendage female the frontal organ is blunt rather than pointed,
and shares with M. kampta the characteristic of hav- anu the male antennular e seta is armed with pairs of
ing a square end to the copulatory appendage, which small pointed spines. C. subarcuata Claus, 1890 (Fig.
is rounded in the other species. M. eltaninae (Deevey, 9.38) is relatively slimmer than the previous three
1978) (Fig. 9.82) has been rather arbitrarily placed in species and the shape of the asyrnmetrical gland on
this genus. This large abyssopelagic species lacks the the right valve is characteristically much broader. The
striations typical of the rest of the species, and the armature of the male antennular e seta consists of
capitulum of the female frontal organ is not clearly pairs of long, sharp spines, similar to that of C. ma-
differentiated from the stem. The anterior margin of crocheira Müller, 1906 (Fig. 9.35), which is a very
the carapace distinguishes this species from others much larger bathypelagic species with a much nar-
with which it co-occurs. rower rostrum. The female frontal organ is blunt and
is curved rather than bent downwards, also the left
w. Species with rather rectangular outlines to the hook appendage is bent through two right angles.
carapace lacking carapace spines and in most
species any accessory glands. Subfamily Archiconchoecinae
These include some of the dominant to subdominant The subfamily Archiconchoecinae at present is mono-
species at mesopelagic depths and include several generic. Archiconchoecia can be subdivided into
strong diel migrators. Females of the smaller species those species with:
Porroecia spinirostris (Claus, 1874) (Fig. 9.112) and
P. porrecta (Claus, 1874) (Fig. 9.110) share the cha- (a) conspicuous reticulate striations wruch probably
racteristics of having a straight undifferentiated frontal includes three species: a deep mesopelagic species
organ and having a fringe of fine hairs along the lead- A. cuculiata Brady, 1902 (Fig. 9.9), a small form
ing edge of the antennular e seta. The armature of the of A. cucullata (Fig. 9.10) which occurs at shal-
male antennular e seta consists of paired spines distal- lower depths and is the more abundant from in
Iy which become altemate and finally a single row tropical and subtropicallatitudes, and a distinctive
proximally. Besides being the larger, P. porrecta undescribed abyssopelagic species referred to here
males are distinguished by having a greater number of as A. aff. cucullata (Fig. 9.11);
spines on the e seta and some long hairs on the b seta
of the antennal b seta. Females have the antennal b (b) only six pairs of caudal furca claws: A. striata
seta fringed with fine spinules which are absent in P. Müller, 1894 (Fig. 9.15), a shallow mesopelagic
spinirostris. Two species Conchoecia magna Claus, species, and A. versicl/la Deevey, 1978 (Fig.
1874 (Fig. 9.36) and C. hyalophylium Claus, 1890 9.17), a deep bathypelagic species;
(Fig. 9.33) have the same type of e seta armature as
the previous two species. Both have long hairs on the (c) a small mesopelagic species with very short stub-
male antennal b seta. The frontal organ of the females by terminal claws on the fifth and sixth limbs: A.
are similar in that they are down-tumed. C. hyalophyl- Cl/neata Müller, 1908 (Fig. 9.12);
lum is readily distinguished in ventral view by the ros-
tra being more rounded and the longitudinal striae in (d) an abyssopelagic species with a clear lens-like
the anterior region of the carapace. It also has two structure on the posterior ventral comer of the ca-
accessory glands next to the right asymmetrical gland rapace, one terminal seta on the first antenna long-
- a character needing high magnification. C. magna er than the others, and very short seta on the endo-
occurs in two size forms in the equatorial Atlantic, but podite ofthe second antenna: A. longiseta Deevey,
the biogeographical range of the large form is 1978 (Fig. 9.13), of which only the female is
unknown. Somewhat similar in carapace shape to C. known;
Ostracoda 837
ÓA
5
ÓB
Gigantocypris muelleri
•• Gigantocypris dracontovalis
~ ÓA
9BE9B
Macrocypridina castanea
9B
Macrocypridina poulseni
ÓB
ÓA
Archiconchoecia bidens
838 Ostracoda
-- 1 ÓA
E__3sc=?
0.5
ÓB ---
•
\/ ÓC
Archiconchoecia bifurcata
• Archiconchoecia
bimucronata
?A~lCJ ~--~ ÓA -.rCJ: ---
~~ ;E3>~
9° 0.1
" ~·t
•
9C
0.1 0.1
Archiconchoecia bispicula Archiconchoecia cucullata

Ostracoda 839
y
~~---
::::,;:,,~~-... ~-.::::::-:
9B -
Archiconchoecia cucullata
small form Archiconchoecia aff. cucullata
deep water form
CJC_J
9A
C_3€3
0.5 , áA
0.5
+ áB
0.1 0.1
Archiconchoecia cuneata
Archiconchoecia longiseta
840 Ostracoda
E=JE3
:¡: d'B ee
Do.sQ
• Archiconchoecia simula e Archiconchoecia striata
900S0A
~€9
9B d'B
900SC]
/fé:> 8
e Archiconchoecia ventricosa
. ~~
Archiconchoecia versicula
Ostracoda 841
Euconchoecia chierchiae Halocypria globos a
Halocypris inflata
e ge
Halocypris pelagica
842 Ostracoda
0.1
9AO OA
9B9 2~B
e
0.1
Fellia bicornis • Fellia cornuta

de
9° I 9A
0.2 9B eB
e Fellia dispar
• Alacia alata
Ostracoda 843
~ 0.2
e Alacia hettacra
• Alacia belgícae
?ADDA
?BC9C3t ::
0.4
de
Alacia leptothríx Alacia valdívíae

844 Ostracoda
?ACJ,CJdA ?~~c=JdA
?B€3E3dB ?~E3dB
• Boroecia antipoda
<¡?c
e
0.2
Conchoecetta acuminata
?A~,CJdA ?A~10A
?BC03JE3J dB?BE38B
<¡?c
• •
dc
0.1 0.1
Conchoecetta giesbrechti Conchoecia hyalophyllum

Ostracoda 845
0.1
Conchoecia lophura Conchoecia macrocheira
Conchoecia magna Conchoecia parvidentata

846 Ostracoda
9CJ,OA
~<=J:,B
ÓH
0.05
9Cf:j
• Conchoecia subarcuata
• 0.2
Conchoecilla chuni
9B=:::i3i=~ 1
9A~ 2c=JÓA
ÓA \ill>')~~ ~ ÓB
ÓD
9~€3JÓB
ÓD
• Conchoecilla daphnoides
e Conchoecissa ametra
Ostracoda 847
9A ~
'C];A
9B~~B
Conchoecissa imbricata Conchoecissa plinthina
Conchoecissa squamosa Conchoecissa symmetrica

848 Ostracoda
dA
C~7 0.5
9AC7,CJÓA
9R=»~~B
ÓB ~ iD
dD
«D
• Oíscoconchoecia díscophora • Oíscoconchoecia elegans
9c;J,c-~A Jl= =t
9~Q
i¿_3J C3ÓB
dD
9BE:)E3ÓB
e
e (~iIb
• Gaussícia edentata e
t¿ Gaussícia gaussí
Ostracoda 849
0.1
Gaussicia incisa Gaussicia subedentata
Loricoecia acutimarginata Loricoecia ctenophora

850 Ostracoda
?ACJ,CJdA C(A - ~,~ -- dA
?03E3dB ?E:3~B
de
e Loricoecia loricata • Macroconchoecia macroreticulata
1 ---"'es$' -n:qMP""
C(A -- ~- dA C(A -~ -'- dA
%J~B do
~E3,B
0.05 ~
• Macroconchoecia reticulata • Macroconchoecia spinireticulata

Ostracoda 851
Metaconchoecia acuta Metaconchoecia arcuata
Metaconchoecia australis Metaconchoecia discoveryi

852 Ostracoda
90,
90_3
CJdA
E_ ~~,
9CJ,OA
ié-_3 c=g,
9D~, - 0.05
9cG
•
0.05
0.05 9C~
.~\ ,1
0.1
G Metaconchoecia fowleri
Metaconchoecia glandulosa
se,
9(---)50A
ÓH
90050A
9B~C)dB
9~~
G Metaconchoecia inflata o 0.1
Metaconchoecia isocheira
Ostracoda 853
«A 0.5
J2~eB 0.5
'-----Q.1'
G Metaconchoecia lunata
0.02 0.02
Metaconchoecia kyrtophora
de
0.05
Metaconchoecia nasotuberculata
~ Metaconchecia macromma
854 Ostracoda
?DsC=L ?C1
?BtE3<c3dB ?B€3
ÓD
• Metaconchoecia obtusa • Metaconchoecia pusilla
?AC :J,Q e
?C3 E 3t :~~SUA ÓD
ÓH
0.1
0.05 '\: 0.02 --===r-- dB
Jt!?\
• Metaconchoecia rotundata
• 0.05
Metaconchoecia subinflata
Ostracoda 855
9°
0.1
0.05
Metaconchoecia skogsbergi
• Metaconchoecia teretiva/vata
de
Metaconchoecia wo/feri Mikroconchoecia acuticosta
856 Ostracoda
9AO, _ 9005
9B§§ ÓB
9B5 GÓB
9D
• Mikrocanchaecia curta
es> Mikrocanchaecia echinulata
9AO UÓA 0.5

(-~A
9A --- 2
9B88ÓB
ÓD
9BC3 E3óB
9D
9D
• •
9~
0.1 QJ
Mikroconchaecia stigmatica Mal/icia amblypastha

Ostracoda 857
( :;0
<¡lA
?BE0--~B
1
0.4
ÓI
b 0.1
0.1
Mollicia eltaninae
Mollicia kampta
Mollicia mollis Mollicia tyloda

858 Ostracoda
9~~7, OA 9AL;OA
Ei)-
9B
E9ÓB nE3~
+ ÓB
0.1
• Orthoconchoecia atlantica
~ Obtusoecia antarctica
_, 19c=3$B
9fC09 c=3?ÓBI9D
óe
~ Orthoconchoecia bispinosa e Orthoconchoecia haddoni

Ostracoda 859
9A~ICJ;A
9BE3 C3ÓB ÓD
• Paraconchoecia aequiseta
Orthoconchoecia secernenda
~ -~IC,>~A
E=~-~~ÓB
0.1
Paraconchoecia allotherium Paraconchoecia cophopyga
860 Ostracoda
~A~ 1 c:=t ~A
t-7~ÓA
9~~~
~~~~ ~ ÓB
9B - ~-~ ÓB
9D
• Paraconchoecia dasyophthalma o Paraconchoecia dentata
~Ac=J,CJÓA
~~~B
~A~ l_ _
:JÓA
~BE3€3ÓB
ÓI
• Paraconchoecia dorsotuberculata e Paraconchoecia echinata

Ostracoda 861
12A~~[ 1A 2ALJ~ I
J:A
2B~C3dB 9B <::;=_]2) E3dB
0.4 ÓD
• Paraconchoecia hirsuta
e Paraconchoecia inermis
• Paraconchoecia mamillata
•• Paraconchoecia mesadenia
862 Ostracoda
9 AC7 c=J:A 9~- _~,c=JdA

'9~E7dB ÓD
9E3C3dB ÓD
0.2
0.05
J!j 0.1
• Paraconchoecia nanomamillata
~ Paraconchoecia oblonga
(9E3
:J,~
C3dB
-~dA
9[---7 c=J:A
9B 9~-- ~B
•• Paraconchoecia spinifera • Paramollicia dichotoma

Ostracoda 863
JLJ,~do/2A
9BC3 ÓD
0.1
Paramollicia major
Paramollicia plactolycos
Paramollicia rhynchena • Porroecia parthenoda

864 Ostracoda
9A(-=2c=J:A 9AC ) 1
UÓA
'9BC3C9óB
90
9Bé???) ~ÓB
óo óo
8>
éc • 0.05
Porroecia porrecta
0.1
Porroecia pseudoparthenoda
9C~O 9ADC~A
9~E3
1 ÓA
ÓB
9E3 E3ÓB
90
óo
0.05
9b iSo
• 0.1
Porroecia spinirostris
• 1
Proceroecia brachyaskos
Ostracoda 865
9AC~C~A
dD
9E3E;)óB
«D 0.1
• Proceroecia convexa • b
~~\d
0.05
Proceroecia macroprocera
Proceroecia microprocera
• 0.05
Proceroecia pro cera

866 Ostracoda
?AC70S C=:L OÓA

E9E3óB
9A
?B Q~
~1
9B 9ÓB
• Proceroecia vitjazi G>

0.2 0.2
Pseudoconchoecia concentrica
JL:J,Cl
?BGEt
~
~ Pseudoconchoecia serrulata
Ostrq.coda 867
(e) another small deep bathypelagie speeies with a resentative oeeurs in the South Atlantie, E. chierchiae
poorly developed rostrum - A. simula Deevey, Müller, 1891 (Fig. 9.18). Bathyconchoecia speeies
1982 (Fig. 9.14); have eurtains of ea. 200 setae on the first antennae and
laek a frontal organ (with one exeeption). No speeies
(f) a speeies with two long terminal and one very of this genus have yet been reported from the South
short (shorter than the last segment) setae on the Atlantie but they are Iikely to oeeur there either in the
fifth and sixth limbs - A. bimucronata Deevey, bathypelagie zone or in very deep water.
1978, ofwhieh only the male is known (Fig. 9.7);
Family Cypridinidae
(g) speeies showing marked sexual dimorphism in Speeies of eypridinids are readily reeognised in hav-
earapaee shape in whieh the female earapaee ing eyes, either eompound (Macrocypridina) or nau-
length is 1.0-1.1 mm and the mal e is ea. 0.9 mm pliar (Gigantocypris). Adult females that are >2 mm
with the frontal organ terminating either in a sin- in length and are earrying embryos within the dorsal
gle needle-Iike point - A. ventricosa Müller, 1906 posterior region of the earapaee are immediately
(Fig. 9.16), or a double point A. bispicula Deevey, attributable to being eypridinids. When sampling
1978 (Fig. 9.17); inshore, speeies not deseribed here may be eneoun-
tered. Adult Macrocypridina are irnmediately identifi-
(h) larger species ea. 1.6 mm in length - A. bifurcata able beeause of their ehoeolate-brown eoloration that
Deevey, 1978 (Fig. 9.6) whieh has a bifid end to persists after long periods of preservation. There are
the frontal organ, and A. bidens Deevey, 1982 two mesopelagie speeies, Macrocypridina castanea
(Fig. 9.5), whieh has a Y-shaped end to the frontal Brady 1897 (Fig. 9.3) and M. poulseni Martens 1979
organ of whieh only the male is known. (Fig. 9.4) distinguished by the shapes of their earapa-
ees: the latter speeies was originally designated M.
Subfamily Euconchoecinae castanea var. rotunda by Poulsen (1962). The two
The subfamily Eueonehoeeinae eonsists of two gene- Gigantocypris speeies are also immediately reeogni-
ra: Euconchoecia and Bathyconchoecia. The former, sable by their large size and rotundity. G. dracontoval-
the only haloeyprid genus in whieh the female broods is Cannon 1940 (Fig. 9.1) is either abyssopelagie or
eggs and juveniles within the earapaee, has about 20 benthopelagie whieh is smaller than the more abun-
terminal setae on the first antennae. Only a single rep- dant bathypelagie speeies G. muelleri Skogsberg 1920
•••
Fig.9. IlIustrations ofthe species. Abbreviations:
A: carapace in lateral view;
B: carapace in ventral view;
C: Endopodite of second antenna (Ieft for males);
D: Frontal organ and first antenna;
E: Details of the setation around the rostrum;
F: Detail oftip of frontal organ;
G: Detail of the sculpturing and openings of the carapace glands at the posterior dorsal comer;
H: Armature of e seta of first antenna;
1: Armature of b and e setae of first antenna;
J: Armature of e seta of first antenna with close up of spines;
K: Armature of d and e setae of first antenna;
L: Armature of b, d and e setae of first antenna;
M: Detail of capitulum of frontal organ;
N: Detail of the last segment of the first antenna;
P: Hook appendage of right endopodite of second antenna;
Q: Lateral view of carapace showing pattem of striations.
AIl scales are in mm.
Figure sources. 121: from Deevey (1977), redrawn; 14,20: from Deevey (1978a), redrawn; 12,74: from Deevey (1982), redrawn; 63, 64: from
Ellis (1984), redrawn; 35: from Müller (1906), redrawn; all others are original.
868 Ostracoda
(Fig. 9.2), and the ventral gape of the carapace Suggested readings
extends its full length. The setation around the ros-
trum is also distinctive. Very large colourless speci- Angel M.V. 1993. Pelagic Marine Ostracoda.
mens occur occasionally in very deep water and pro- Synopses ofthe British Fauna (New Series), (D.M.
bably belong to a third species. In the North Atlantic Kermack, R.S.K. Barnes, J.R. Crothers, eds.), 48,
Gigantocypris muel/eri is somewhat smaller than published for Linnean Society of London and The
specimens collected in the Southern Ocean, but where Estuarine and Coastal Sciences Association by the
their size changes and if this is of taxonomic signifi- Field Studies Council, pp. 1-239.
cance needs to be determined. Iles E.J. 1961. The appendages of Ralocypridae.
Discovery Rep., 31 :299-326.
Poulsen E.M. 1977. Zoogeographical remarks on
marine pelagic Ostracoda. Dana Rep., 87: 1-34.
Skogsberg T. 1920. Studies on marine ostracods. Part
l. (Cypridinids, halocyprids and polycopids).
Zool. Bidrag Uppsala, suppl. 1:1-784.
Copepoda
Janet M. Bradford-Grieve, Elena L. Markhaseva, Carlos E. F. Rocha and Bernardo Abiahy
Introduction orders, as well as data on feeding and feeding modes,

reproduction, development, distribution and taxono-
Marine copepods are small crustaceans usuall y my are given throughout the following text.
between 0.2 - J 2 rnm in length. More than J0,000
species of free-living and parasitic copepods are
known to inhabit fresh, brackish, and marine waters Mo r phology and biology
and terrestrial localities (Huys and Boxshall, 1991 ).
Marine copepods are known to be pelagic (inhabiting External morphology
the water column), benthopelagic or hyperbenthic
(living in the near-bottom water layers), benthic The classification of the copepods is largely based on
(living on the bottom or in the sediments) or in asso- body tagmosis (division into functional regions) and
ciation with other animals. Their vertical range is segmentation and armature of the various limbs. The
from the surface to abyssal depths and they are known terminology adopted here for the description of the
fro m all the biogeographical zones of the World external morphology of copepods follows that of
Ocean in neritic to oceanic waters. Being numerous Huys and Boxshall (1991). The major body articula-
and abundant marine organisms, they may sometimes tion subdivides the body into an anterior pati - the
for m up to 90-97% of the biomass of marine prosome (Pr) - and a posterior part - the urosome
zooplankton, therefore copepods are an important lin k (Ur). The anterior region of the prosome, covered by
in marine food webs and the marine economy. the dorsal cephalic shield, is the cephalosome (Ce)
which comprises the five cephalic somites bearing the
Milne-Edwards ( 1840) established the Copepoda as a antennules (A l), antennae (A2), mandibles (Md),
separate taxon. Since that time different classification maxiUules (Mxl ) and maxillae (Mx2), respectively,
schemes have been proposed for copepods. Up to now and the first thoracic segment bearing the m axillipeds
I 0 copepod orders are recognized: Platycopioida, (Mxp) (Fig. 1). In many copepods the second thoracic
Calanoida, Misophrioida, Harpacticoida, Monstrilloi- somite is also fused to the cephalosome, forming a
da, Mormonilloida, Gelyelloida, Cyclopoida, Siphono- cephalotho rax. This somite bears the first pair of
stomato ida and Poecilostomatoida. Since 1990 detail- swimming legs (Pl) and is called the ftrst pedigerous
ed phylogenetic ana lyses of the Copepoda were somite (Pd l).
carried out by Ho (1990), Dahms (199 1) and Huys and
Boxshall (199 1). The latter authors give an excellent There are seven postcephalic thoracic somites: the
review of existing phylogenies and the history of cope- maxiUiped-bearing somite, five pedigerous somites
pod studies since the eighteenth century up to the early (Pdl-5), each bearing a pair of swimming legs (P I -
1990s. The scope of the present work is the known PS) (Fig. 2), and the genital somite. The seventh thor-
pelagic and benthopelagic fauna of South Atlantic acic or genital somite (Gns) bears the sixth pair of
which contains four of the known copepod orders: legs (P6) w hich are highly modified and fom1 the
Calanoida, Cyclopoida, Poecilostomatoida and opercula that close ofT the paired female genital
Harpacticoida. For more detailed infom1ation on apertures (Huys and Boxshall, 1991). Each pair of
copepod phylogeny and morpho logy the reader is swirruning legs is joined by an intercoxal sclerite
referred to Huys and Boxshall (1 99 1), on intemal ana- which may be fused to the coxae. The swimming legs
tomy to Boxshall (1992), and for a bibliography of usually have two basal segments: coxa (C) and basis
literature on copepods to Vervoort's publications (B). Two rami are articulated to the basis: an outer
(1986a, 1986b, 1988). exopod (Exp) and inner endopod (Enp). The exopod
and endopod are boardered by spines and/or setae
References for fllliher reading on external morpholo- which are denoted in the family desctiptions accor-
gy, anatomy, phylogeny and definition of the copepod ding to the fo tmula system of Sewell ( 1949) (Fig. 2).

edited by D. Boltovskoy, pp. 869- 1098
870 Copepoda
A2
(antenna)
A1
(antennule)
Mxp
(max il liped)
Mx1
(maxillule)
Pc
(praecoxa)
Mx2
(maxilla)
Copepoda 871
Within the Copepoda there are two major plans of Structure of calanoid
body organization (tagmosis) into an anterior prosome swimming leg
and a posterior urosome, separated by the major body
articulation (asterisks in Fig. 3). In the gymnoplean
intercoxal
tagmosis (Platycopioida and Calanoida) the major sclerite
body at1iculation is located between the fifth pedige-
rous somite (Pd5) (=sixth thoracic somite), primitively \I
p~
bearing the P5, and the genital somite. In the podople-
an type oftagmosis (all remaining copepod orders) the
major articulation primitively lies between Pd4 and
Pd5 (between fifth and sixth thoracic somites).
In gymnopleans, the urosome includes the anterior

somite corresponding to the seventh thoracic somite
End 1
(=genital somite). In females ofCalanoida it is usually
fused with the first abdominal somite, forming a geni-
tal double-somite. The abdomen is the limbless post-
genital region of the body. In podopleans the urosome
comprises the sixth thoracic somite bearing the P5, the
seventh thoracic somite and five abdominal somites. In
the majority of Harpacticoida, Poecilostomatoida and
Cyclopoida the fema le second and third urosomites
(=seventh thoracic segment and first abdominal
somites) are usually fused to fonn a genital double-
somite. The difference between gymnopleans and pod-
opleans may be observed even in the early copepodite
development (Fig. 3). The last somite of the urosome
is the anal somite bearing a pair of caudal rami (CR).
A detailed review of the literature on the copepod

internal anatomy may be found in Blades-Eckelbarger
(1986) and Brodsky eta/., ( 1983). The original stu-
dies on the internal anatomy of Calanus flnmarchicus
and Epilabidocera amphitrites were made by Lowe
(1935) and Park ( 1966), respectively. Comparative coxa basis exopod endopod
studies of copepod skeletomusculature were publis- 1 I 2I 3 1I 2 I 3
hed by Boxshall (1985, 1990). /o- 1 1-1 11-1 / 1-1/111,1, 5
Reproduction and development Fig. 2. Structure of calanoid swimming leg. Basic copepod swim-
ming leg, showing the maximum setation of a second leg (fro m
Calanoids, cyclopoids, poecilostomatoids and harpac- Huys and Boxshall, 1991). Sewell ' s (1 949) system of spine and
ticoids are sexually dimorphic. Fertilization is by cop- setal description, used in the family descriptions, is g iven in the
box.
ulation when the male transfers one or more spennat-
ophores to the female's genital somite. Then spenn is
transferred through the female ' s copulatory pore and
stored in seminal receptacles. Fertilized eggs are
released through the gonopores that represent the ovi-
fig. 1. Structure of calanoid body and mouthparts. For further details see "Use of the keys" in the lntroduction.
872 Copepoda
Copepodid stages
Cl Cll c Ill CIV cv Adult
c
C'V<Jl
<1> ""0
-0...0...
o
0(1)
Co_
Eo
>-u
lJ
c <Jl
ro-o
~0
o_O...
Q<l>
-oO...
oo
o._ U
Fig. 3. Comparison of the developmental pattern in podoplean and gymnoplean copepods. C l-CY: first copepodid to fifth copepodid deve-
lopmental stages: solid stars indicate flexure planes, hollow star indicates poorly defined flexure plane, heavy arrows indicate the stage at which
the definitive tagmosis is attained and specialisation of the joint commences. From Huys and Boxshall (1991).
duct openings. Eggs are held together by outer mem- copulation (Huys and Boxshall, 1991). It is consid-
branes and form single or paired egg sacs. In harpac- ered that upon chemical stimulation the male per-
ticoids, calanoids, cyclopoids and poecilostomatoids forms a distinctly different swimming behavior -
true egg sacs are present, however the majority of "searching movements" or "mate-seeking behaviour",
calanoids free-spawn their eggs into the water column and these movements bring him into close proximity
(Huys and Boxshall, 1991; Huys et al., 1996). to a potential mate. The review of literature on mating
Precopulatory mating before copulation is usual. The behaviour of calano ids is given by Blades-
male grasps the female with the prehensile Al (har- Eckelbarger (1991). Recently the mating behaviour of
pacticoids, calanoids) and also usually with the PS cyclopoids and calanoids was observed using video-
(calanoids). In cyclopoids the male has both A1 computer motion analysis techniques. It was proposed
geniculated, therefore it grasps the P4 ofthe female in that mating behaviour (encounter-pursuit-capture and
a ventral to ventral orientation. In poecilostomatoids copulation) can be compared to predatory behaviour
the Al are non-geniculated, but Mxp are sexually ( encow1ter-attack-capture and ingestion) (Buskey,
dimorphic and used for holding the female during 1996).
Copepoda 873
The fecunclity of a female depends on the species, the vious accounts of copepod feeding (Alcaraz et a/.,
amount and quality of food eaten, season, hydrograph- 1980; Koehl and Strickler, 1981; Strickler, 1982;
ic conditions and latituclinal zone. The developmental Price et al., 1983; Turner eta/., 1993). Copepods in
rate of eggs depends on temperature and may prolong their environment are dominated by viscous forces
from l day to several months. Diapause eggs are and the flow around them is laminar. Their appenda-
known for calanoids (e.g. Uye, 1985; Marcus, 1996). ges behave more like solid paddles than open rakes
Nearly all free-living copepods have 12 developmental (Koehl and Strickler, 1981). They flap their A2, Mdp,
stages: 6 naupliar and 6 copepodite stages, the sixth Mx1 , and Mxp to produce a pulsing stream of water
being the adult. The fust nauplius has 3 pairs of which brings food particles near the stationary Mx2
appendages: AI, A2 and Md. T he sixth naupliar stage (Fig. 4). The Mx2 then fly ap art so that water and the
has all appendages up to P2 present although the pos- food particles flow into the basket formed by the Mx2
terior limbs are vestigial. The duration of each naupliar wh ich close over the algae and water. The endites of
stage is very short (from few hours to few days). The Mxl then pushes the algal cells into the mouth. A lgal
period between moults from one copepodite (or cope- cells are usually redirected without actually being tou-
podid) stage to another may be much longer. The fifth ched by the feecli ng appendages. Similarly, particles
copepodite stage of some calanoid species may live can be rejected by movements of the Mx2. The captu-
longer than others. They may feed intensively and store re of small cells produce a different behaviour. The
lipids. especially in those calanoids whose adult stages Mx2 move continuously wi th a low amplitude inter-
have reduced oral pans. Some species of Calanus may rupted by "combing" of the appendages by scraping
ha,·e one generation per year in the northern areas. but their setae against the endites ofMx1. In these studies
the same species in warmer waters may ha,·e two gen- it was also suggested that herbivorous and carnivo-
er.nioos per year. Continuous breeding is characteristic rous feeding are similar processes in calanoids and
():- ~icaJ copepod.s: in temperate regions breeding that predators catch their prey by grasping it with the
cycles strongly Yary (Da\is. 198-t ). Generally. copep- tips of :\lx2 and Mxp setae or capture a parcel of
ods exhibit a wide variety of life histories. water containing the prey.
Feeding and feeding modes Carnivorous calanoids differ from suspension-feeders

in that the Md has a strongly chitinized Gn with teeth
Copepods are recognized as indise~iminate or selecti- narrow and saber-shaped on the cutting edge. Usually
ve suspension or particle feeders. They may be herbi- the ventral teeth are sharper in raptorial predators. The
vorous, predatory feeders (carnivores), mixed-mode fewer the number of teeth, the greater the degree of car-
feeders (omnivores) (Arashkevich, 1969; Davis, nivory (Arashkevich, 1969; Ohtsuka, 1991 ; Ohtsuka el
1984; Schnack, 1989), saphrophages or coprophages al., I 996). Also the Mx l is usually characterized by
(Ohtsuka et al., 1996). However some copepods are strongly developed Lil -L i3, with setae of Li 1 spine-
apparently able to switch from one feeding mode to and dagger-shaped, and the setae of Enp strong, claw-
anoth er according to circumstances (Lowndes, 1935; like, and serrated along their whole length. The setae of
Kleppe!, 1993). The type offeeding in general is usu- Lel are usuall y plumose. Th e Mx2 has the majority of
rsid-
ally reflected in the structure of oral parts and the type Li bearing stout, spine-like setae, and sometimes those
:;>er-
!1£-
of locomotion. The main food items for suspension- on the distal Li are highly chitinized and claw-like, and
F.!T-. feeders are phytop lankton and mic rozooplan kters often serrated. The Mxp is massive with spine- and
city (ciliates, tintinnids, copepod nauplii). Suspension fee- claw-like setae. Many setae on the oral parts lack
ders usually move slowly and smoothly when feeding setules. The Mx2 and Mxp may be developed in differ-
nng
producing water streams around the body by beating ent ways: for example in Candacia the Mx2 is more
rles-
r-of of the oral pa11s. In general suspension-feeding powerfully developed than the Mxp. On the other hand,
calanoids have A2, Mdp, Exp, and Enp as well as in Euchaetidae and Pseudeuchaeta (Aetideidae), the
deo-
Mxl Le1 (coxal epipodite), Exp and Enp supplied Mxp is more powerfully developed. However in both
llSed
w ith long, strong, densely plumose setae impo1tant in cases they are both prehensile appendages used for
~
the c urrent production process. High-speed photogra- grasping. The recent study of the Md of carnivorous
phy (500 frames per second) of dye streams around heterorhabdids showed that an anaesthetic is injected
feeding calanoid copepods has led to a revision of pre- into a groove or tubular lumen of the ventralmost tooth
Fig. 4. Copcpod feedin g. Feeding currents bypass Mx2 (A-B) until an alga nears them. An alga is captured by an outward fling (C-D) and an
inward sweep (E-F) ofMx2. Diagrams are traced from high-speed fi lms of a feeding Subeucalanus pileatus. Heavy arrow indicates movements
of Mx2 (and of Mxl in F). Small circles represent position of and line arrows indicate the movements of algae observed during similar appen-
dage motions in other frames o f fi lms. Tn the first column, the copepod is viewed from its left side and Mxl has been left out for clarity. In the
second column, the animal is viewed from its anterior end (from Koehl and Strickler 198 1). Right-band panel indicates relative appendage
movements of Subeucalanus pileatus during the capture of a Prorocentrum micans cell at concentrations of 1.0 mm3 litre-1 or 120 cells mr 1.
Dots indicate cell movement. From Price et a/. ( 1983).
of the manctible and rel.eased into prey from a hypoder- harpacticoids are regarded as surface feeders (Huys et
mic needle-like tip of the tooth (Nishida and Ohtsuka, al., 1996). Cyclopoids more probably are carnivorous,
1996), confirming the earlier proposal that the hetero- capturing food actively, but some (Oithona simi/is) are
rhabdids are sucking predators based on the structure of mainly herbivores (Davis, 1984). A recent study of sap-
their oral parts (Arashkevich, 1969). phirinid copepods leads to the proposal that these
copepods are free living during daytime, but attach at
The morphology of omnivore oral parts is, in general, night to other animals and feed on their tissues (Nishida
intermediate between that of suspension-feeders and and Chae, 1996); and oncaeaid copepods were
predators. Scolecitrichids, with oral parts close in observed to be active predators on larger zooplankters,
structure to those of mixed-mode feeders, possess such as Sagitta and Oikopleura (Davis, 1984), or have
sensory setae on the distal part of Mx2 and feed on a surface mode of feeding (Boxshall, 198 1) having
detritus, discarded appendicularian houses, and exu- been observed grazing on appendicularian houses.
viae (Ferrari and Steinberg, 1993; Ohtsuka et al. ,
1996). The mechanism of detecting food particles or
prey is speculated to be chemosensory. Methods
In harpacticoids direct carnivory of metazoan organ- Copepods inhabit ctifferent environments and, depend-
isms is relatively unknown, some are known to be sus- ing on what kind of copepods are collected and the final
pension- and mucus trap-feeders, but the majority of purpose of the study, ctifferent gear is utilized. For the
Copepoda 875
purposes of quantitative and/or qualitative study of the More than 70 species are apparently restricted to the
pelagic copepod fauna, standard plankton nets are used. Atlantic Ocean although some of these represent
There are a variety of such nets (Juday's, Nansen's, 0.5 infrequently recorded species which may be revealed,
m2, I m2 cone nets, Bongo nets, plankton traps, etc.). in the future, to be more widespread.
The nets may have closing systems so they sample at
known depths (for stratified sampling), or be without Typical assemblages of South Atlantic epipelagic
such systems for integrated hauls to the sea surface. As copepods, their horizontal distribution, and connection
well as thi s standard equipment, special gear is with water masses are given in Bj0rnberg (198 1); and
l
employed for example for ecological studies of ben- in the set of subsequent publications (Can1paner, 1985;
thopelagic copepods as sampled from the D.S.R.Y. Carola, 1994; De Decker, 1984; Brenning, 198 l a,
"Alvin", or the study of the distribution of sound scat- 198 1b, 1982, 1983 , 1985a, 1985b, 1986, 1987).
terers (Wishner, 1980; Gowing and Wishner, 1986, etc.). evertheless, many of the species dealt with in this
When copepods are not collected for in vivo experi- chapter have been recorded so infrequently that we
p ments and live observation, but for the later study of cannot, with certainty, make a statement about their
preserved material (taxonomy, distribution, gut-content vertical and horizontal distribution and their relation-
analysis, etc.), they are usually preserved with 2.5-5% ships. if any, with the hydrographic circumstances in
formaldehyde, glutaraldehyde or 70-75% ethanol (the the South Atlantic. Although it is commonly thought
latter not commonly used as it makes copepods fragile). that the Atlantic is the best studied ocean, it is clear
as well as other less commonly used preserYati\·es. from a re\iew of our knowledge of the pelagic copepod
fauna that large areas of the central South Atlantic have
The identification of copepods is often impossible nor been sampled. or if sampled, not comprehensively
without the previous dissection of specimens. reported on. Thus the southern boundaries for many
Dissection is perf01med with fine dissecting needles warm and temperate \Yater species are hardly known.
and forceps. Dissected limbs are placed on a slide and
are examined in lactic acid, glycerin (usually mixed Data on the little-known near bottom South Atlantic
with distilled water), or are mounted permanently in copepod fauna may be found mainly in taxonomic
euparal, polyvinyl lactophenol, Berlese ' s F luid, publications (Alvarez, 1984; l 985a, b: 1986;
Canada Balsam or some other mounting medium. Andronov, 1976; 1979; Bj0rnberg, 1975 ; Campaner,
There are a variety of mounting techniques with corre- 1977, 1978a, b; 1979, 1986). Benthopelagic copepods
sponding preferences according to the respective pur- are poorly known world wide which probably
pose (for instance for the observation of integumental explains the apparent level of endemism in the South
rs et organs: Koomen and Von Vaupel Klein, 1995). Prior Atlantic near-bottom fauna.
005.. to dissection, clearing and staining of the specimen is
l :!re recommended to make small morphological structures Horizontal distribution
more easi ly observed under the light microscope. A
brief but extensive review of the methods for fixation Many species are found over a range of depths
and preservation, mounting and sealing media, slide (Vinogradov, 1968) and the geographical distribu-
preparation, and scanning electron microscopy may be tions of species with different bathymetric distribu-
ere found in Huys and Boxshall (1991 , pages 446-454). tions varies. Cosmopolitan distri butions are more
lefS. The illustrative material accompanying copepod stud- typi cal of deep water species (e.g. Gaetanus tenuispi-
~ e ies varies. Usually line drawings, made using a came- nus, Pseudochirella obtusa, Paraeuchaeta barbata,
ring ra Iucida or other drawing apparatus, are used. Oncaea englishi, 0 . mediterranea). Apparently bipo-
Scanning electron micrographs also illustrate some lar species also tend to be deep water species (e.g.
taxonomic descriptions and cinematographic observa- Aetideopsis minor, Chiridius polaris).
tions of feedi ng and reproductive behaviour are recent
additions to available techniques. Epipelagic copepods are rarely cosmopolitan due to
their environment which is strongly influenced by cli-
Distribution mate and hydrography. However, quite a few epipelag-
ic species have wide distributional ranges (see Table
The pelagic copepod fauna of the South Atlantic 1). For example Acrocalanus species, Mecynocera
includes more than 500 free-living copepod species. clausi, Eucalanus hyalinus, Pareucalanus sewelli,
876 Copepoda
1600 120° 80° 400 oo 40° 80° 120° 1600
800
60°
' I
I .<• .~ 300
• ......
• •• oo
• ··-
• •••• ~
'"'I'
•
300
••• "' If '

600
• 60° @{
700
Sapphirina nigromaculata Pontellina platychela
600 200
oo
,
"
....
.:
y0. + + + + +
:
'
+
:
:
'
...'
' ...
'
+ + + + + + 1• • · - - · - - - - - - . .
:
:
:'
'
:
--------.o;
0
zoo
.. ..
.•: - ~-- - · · · · ·· ·· -~ -- ----·····-~-- ~
........ ............ ; ... 400
::• . '
• P. marshi
.
• P. acutus
+ P. richardi
o P. hessei
D Area
surveyed
Suggested
distribution II Area of high
abundance "® ~
'
'
'
............
. ......
~
o P. nudus/serricaudatus
0 P. cha rteri
60°
' -• :
Macrosetella gracilis Pseudodiaptomus spp.
Fig. 5. Geographic distributions. A: widespread epipelagic tropical-subtropical species. B: an epipe lagic Atlantic species. C: results based on
continuous pump samples; D: distribution of the mostly estuarine genus Pseudodiaptomus. A: from Lehnhofer ( 1929); B: from Fleminger and
Hiilsernann (1974); C: modified fi·om de De Decker (1984); D: from Wright (1936), Grindley (1963).
most Oithonidae, Sapphirinidae (Fig. 5A) and bution. Examples of such species are Euchaeta marina,
Corycaeidae have a widespread circumglobal tropical- E. paraconcinna and Pontellina platychela (Fig. 5B).
subtropical distribution. It appears that only those spe- The distribution of these species exhibits another fea-
cies that are able to breed at latitudes that extend to ture of low latitude distributions in the South Atlantic.
40°S have such a widespread distribution. Those spe- These distributions are not symmetrical north and
cies which are endemic to the Atlantic Ocean are more south of the equator. It appears that the coastal
usually epipelagic and have distributions which do not Brazilian Current carries such species further to the
extend very far south, such that the South American south on the eastem border of the South Atlantic.
and African continents present a barrier to their distri-
Copepoda 877
FemaleI Male Vertical Inshore/

Species I
length length
(mm)
distrib. offshore latitudinal distribution
_ L (mm) distr.
Order Calanoida
Family M~acalanidae~---
Megaca/anus princeps 8.50-11.50 7.90-10. 10I B o w, a
Bathycalanus eltaninae
Bathycalanus inflatus__
Bathyca/anus princeps
Bathycalanus richa.'-'rdc.:.i_ __
Bradycalanus Qg:<Jig<=a=-s_ _
_ __:_: 15:::_:.00-16.00
13.60-13.85
10.75-13.50
-----'8::_:_..25-13.50
12.50-14.90
12.70
unknown
unknown
7.80-8.15
unknow
u ~? B
B t iSWAt, SEP, a
B o
o
o
o
SWAt, a
w, SAt (at 572 S)
w, a
SAt, I
l
-j
J
Bradyca/anus pseudotypicus 12.50-13.75 unknown B o SAt (at S42 S), IWP, I, SEP, a
J!radycalanus typicus 9.00-15.00 unknown B o SAt (at 562 S), IWP, I
Calanoides acutus
Calanoides carinatus
Calanoides
- - - macrocarinatus
- --
Calanus agulhensis
Calanus australis
Family Calanidae
_ _ _ 3.20-3.90
I
_ _ _ 2.25-2.85
3.50-5.70
2.45-2.95
2.60-3.59
I 1
2.90-3.90
2.74-3.00
2.90-3.47
5.50
2._
E-~0
7 0'----_cc..
E-M
E-M
E
E
o
o
c
At (472 N-372S), P, I
a j
:.:..,._____
-
- --
Calanus propinquus 4.75-6.00 4.75-4.95 E-M j o a, sa
Calanus simillimus 2.65-3.80 2.62-3.42 E-M _j_ o sa, a 1
Canthocalanus pauper 1.30-1.60 1.30-1.50 E o t, SEAt, P
Cosmocalanus darwinii 1.60-2.40 1.63-2.05 E 0 t,st~
Mesocalanus tenuicornis
~ +~
1.80-2.40 1.70-2 .20 t, st, w
Nannocalanus minor 1.80-2.25 1.20-1.80 t, st, w
Neocalanus gracilis 2.43-4.00 2.30-3 .10 E o t, st, temp, w
Neoca/anus robustior 3.42-4.32 2.84-3.30-H 0 t, st, w
Neoca/anus tonsus 3.40-4.10 3.30-4.40 E-M _ _o_ a, sa, STr _ __
--- --
Undinu/a vulgaris 2.25-3.25 2.04-2.50 E J n t,w
Family Paracalanidae
Acrocalanus andersoni _ _ _ ...J..I_ 1.:...:..1.:..=.5-1.3o 0.99-1.10 L_L eqP, eq l, SAt (72 S), NAt _ _ j
Acrocalanus gibber 0.93-1.13 0.94-1.40 EE t,w
Acrocalanus gracilis
Acrocalanus longicornis
1.20-1.32
1.00-1.26 1 0.88-1 .00
0.95 -1.25 E 0
t,w
t, st, -"w'----- I
Acroca/anus monachus o t, st, P, I, SEAt
Calocalanus at/antic_u-'-
Calocalanus contractus
s _ __
0.90-1.05 unknown
0.80-0.9:...:.1---l__:u:.::nknown
0.68-0.76 0:.:.._49'---+ E
E
E ~~ (at 262 5), Med
0 t, st, P, A_t _ __
_J
S:aloca/anus equalicauda 1 .10 unknown Brazil, NAt
Calocalanus lomonosovi 0.78~0 . 82 0.78 I SAt (at 262 Sl
Caloca/anus m-'-'in.:.:o:.r:._ _
Caloca lanus minutus
Ca/oca/anus namibiensis
0.50-0.57
0.66-0.70
0.43 unknown
0.~0-0.57
O..J0-0.52
j E o
SAt(at2~
t, st, At, I, SWP, SAl (at 202 S)
....:o SWAfr, SWP
Calocalanus pavo:.:.__ __
Calocalanus plumulosus
0.88-1.20
0.93-1.20
0.91-1.04
0.83-0.90
I =-
E _
E
E
_
I
o t, st, temp, w
o t, st, temp, w, Med
Calocalanus styjiremis ' 0.59-0.72 0.55 t, st,w _ _ __
+r!
---
Ca/oca/anus tenuiculus 0.59-0.7 : r r0.48-0.55 SWAfr (1 72 -2425)
Calocalanus tenuis 1.18-1.31 unknown
J
De/ius nudus 0.52-0.70 0.42~ E 0
De/ius sewelli 0.53-0.57 0.43 l E -+ 0
Paracalanus aculeatus 0.80-1 .36 0.92-1.36 I E o? 1, sl, w
Paracalanus campaneri _ 1 .OS unknown E -"--=- o:...?--'off Brazil
878 Copepoda
Female Male Vertical Inshore/

Species length length distrib. offshore Latitudinal distribution
(mm) (mm) distr.
Paraca/anus denudatus - - --0.70-0.93 unknown E o? t, st, temp, w

Paracalanus indicus ---
0.85-0.95- - -
0.85-1 .02 E c w?
Paraca/anus nanus 0.60 0.57 E w
Paracalanus parvus 0.70-1.30 0.74-1.40 E c w?
Paracalanus tropicus 0.74-0.90 unknown E SAt (at 1 8"5)
Paracalanus quasimodo 1.00 unknown E c off Brazil, NAt (at 33 2 Nl
Parvocalanus crassirostris 0.47-0.55 0.35-0.39 E c? w
Parvoca/anus scotti 0.64-0.67 unknown E e, c SWAt
Famil Mecynoceridae
Mecynocera clausi I
o.92-1.21 0.94-1. 12 E o t,st, tem~
Family Eucalanidae
Eucalanus hyalinus 5.10-7.10 4.70-6.25 E-M 0 At (65"N - sa), w
Pareucalanus Jangae 5.61-7.22 6.12-6.59 E-M 0 STr, ci rcumpo lar
I
Pareucalanus sewelli 3.86-6.10 2.89-4.58 E 0 t,st,w_ _ __ I
Rhincalanus cornutus 2 .90-3.80 unavailable E 0 At
-t
Rhincalanus gigas 6.46-9.30 6.90-7.20 E-M 0 a (to 50 2 5)
Rhincalanus nasutus 3.90-5.30 2.70 -4.30 E 0 t, st, w
Subeucalanus crassus 2.80-3.70 2.90-3.10 E 0 t, st, w
Subeucalanus /ongiceps 4.20-4.90 3 .20 E 0 sa
Subeucalanus monachus 2.13 -2.35 2.20 E 0 t, st, At
Subeucalanus pileatus 1.95-2.50 1.80-2 .25 E 0 t, st, w
Subeucalanus subtenuis 2.65-3. 10 2.75 E 0 t, st, w
Family Spinocalanidae
Mimocalanus cultrifer 1.00-1.70 unknown M-B 0 w
Monacil/a gracilis f1.8o-2.2s unknown M-B o At (0"-12 2 5), NWP?
Monacil/a typica 1.95-2.50 1.59-2.30 E-B 0 w
:
Mospicalanus schie/ae 2.1 1-2.18 unknown B o a
Spinocalanus abyssalis 0.85-.:. .:1·.: : . o23::..._____:_:
1 .06-1 .20 E-B 0 w
(
Spinoca/anus brevicaudatus 1.40-1.95 1.30-1 .72 M-B 0 w
(
Spinocalanus horridus 2.00-2.47 unavailable B o a, w
(
Spinocalanus magnus 1.87-3.10 1.80-2.43 M-B o a, w
(
Clausoca/anus arcuicor~~smily Clauso.c a. lanid_ae l 1. 15-1.62 0.97-1.1 7 E o t, st, w

(
(
Clausocalanus brevipes 1.24-1.62 1.12-1.20 E o sa
(
Clausocalanus furcatus 0.94-1.3 1 0.70-0.92 E o t, st, w
(
Clausoca/anus ingens 1.44-1.90 0.87-1.07 E o STr
(
C/ausoca/anus jotJ_ei 1.01-1.56 0.87-1.0 7 E n, o t, st, w
Clausocalanus /aticeps _ _ _..,_1.25-1.67 1.01 -1.10 E o a, sa
Clausocalanus lividus 1.26-1.77 1.13-1 .45 E o st, w
Clausoca/anus mastigophorus 1.23-1.84 1.05-1 .45 E o t, st, w
C/ausocalanus m:.:.:i.:..:n.Or
:::.:...__ 1.08-1.26 0.79-1 .04 E o off S Afr
Clausocalanus parapergens
C/ausoca/anus pau/ulus
0.97-1.38
0.66-0.80
0 .97-1.14
0.47-0.56
E
E
o t, st, w
o st, w
•
Clausoca/anus pergens 0.70-1.10 0.52-0.67 E o st, w
Ctenocalanus citer 0 .80-1.41 0.99-1.45 :::. :::.:.ff Brazil?, SAfr ?, SWP
E_ ____,o:..___o
Ctenocalanus vanus _ _ __:0:..:·.::..
92-1. 16 1.20-1.26 E o t, st, w?
Orepanopus forcipaws 1.84-2.74 1.66-1 .94 E ~a , SWAt, SEP
I
· Farrania frigida - -- 2 .25-2. 35 2.34 B o eqAt
Copepoda 879

Species length length distrib. offshore latitudinal distribution
(mm) (mm) distr.
Family Aetideidae
Aetideus acutus 1.48-1.80 1.22-1.58 E-M 0 1, st, w
Aetideus armatus 1.60-2.00 1.30-1.53 E-M 0 t, Sl , W
Aetide us australis 1.78-2.04 1.60-1.74 E-M 0 a, sa
Aetideus giesbrechti 1.80-2.20 1.52-1.60 E-M 0 I, st, w
A etideus pseudarmatus 1.65-1.80 unknow n E-M 0 I, st, SEAt, P
A etideopsis carinata 2 .66-2.96 2.4 M 0 SE At
Aetideopsis minor 2 .87-3.90 u nknown M-B 0 bipol ar
A etideopsis rostrata 3.80-4 .60 3 .13 -4.20 8 0 a, ar, NAt, NP, tP
Br~idius plinoi_ 2.45-2.52 unknow n BP c Brazil
Chiridiel/a atlantica 2.30 unknown M 0 eqAt
Chiridius gracilis 2 .45-2 .80 1.96-2.32 M 0 t, st, sa, a, w
Chiridius polaris 3 .30-4 .25 3 .55-3 .80 M-A 0 bipola r
Chirid~ppei 1.59-2 .20 1.50-1.58 E-M 0 t, st, w
Chirundina streetsii 4 .1 .5-5.55 3.80-5.20 M? 0 t, st, a, w
0 runcrne a magna -.44-8 .10 6.70 M 0 t, w
Corr.a~~e:rma cuttisetosa 2.20 un known BP c Brazil ...,
E_-cmre/la amoena 2 . -~ .00 3 .00-3.8.5 E 0 t, w
- - --1
=xmre/la btrumida .!. - 0- - .10 .!.80-6.10 E-B 0 At, P, IWP

f::Jchirella curtica:xia 3. .5()....! . .5.5 3.1 ~ .30 E-\1 0 At (66°N-11 °$), P, tl
Euchirella i"ormosa .!.80-.5 ..50 .!.1.5·.5.20 E 0 t. st, \\
Euchirella maxima 6..50-8.-o 6.1 0-7.3.5 E- \1 0 ~~ sa, w, At (66°N-3 5°5)
Euchirella messinensis 4.40 -6. 20 2.80-.5.46 \I 0 tAt, \led
Euchirella pulchra 3.04-4 .40 3 .06-4.15 E-M 0 1, st, temp, w
Euchirella rostrata 2.9 5-3.95 2.5 0-3 .1 0 E-M _ ~ cosmo_es>litan except ar
Euchirella simi/is 4 .00-5.41 unknown M 0 1, st, sa, w
Euchirella sp lendens 3.88-5.05 3.40-3 .76 E-M 0 tAt
Euchirella truncata 5.20-6.80 4.50-5.60 E-M 0 t, w, NAt
7.60-9 .08 7.60-9.08 M-B
rl
Caetanus antarcticus 0 t,__g, w, a ,~
Caetanus brevicornis 3.70-4.98 4.2 5
~
M 0 t, w
Caetanus brevispinus 3 .60-4.90 2.30-4. 10 E-8 0 cosmopolitan except I -1
Caetanus kruppii 4.70-5.70 4.50-5.20 M-B At (65°N-39°S), w
J
0
J
Caetanus miles 3.40-3 .80 3.00 E-M 0 t, w
+- -
Caetanus minor 1.70-2.40 unknown E-8 0 t, st, sa, w
[ Caetanus pileatus 4.90-6.70 4.44-5.08 M 0 t, w, sa, NAt, stAt
Caetanus robustus
tc'aetanus tenuispinus
Lutamator elegans
- 7.50-9.20
5.1 5
7.00 __J_
3 .00-3 .80_1 2.00-3.4 3
unknown
8-A
E-8
BP
0 __!, st, w, NP, N~
0 cosmQe91itan
c j8razil
-·
Meso comantenna spinosa 1.90 unknown 8P c Brazil
Paivella inaciae I .42-1 .46 1.1 5 E c? Nige ria
Paivella napora i 1.27-1.31 unknown E-A 0 SAt
I Paraco mantenna gracilis 1.40 unknown BP c Brazil
Paracomantenna magalyae 1.80-1.90 unknown BP c Brazil
Pseudochire lla~tillipa 5 .58-6.10 5.40 8-A 0 bipola.!:_
Pseudochirel/a dubia 4.80-6.10 4.00 B-A + 0 cosm_QQolitan exceJ::ll ar
Pseudochirella hirsuta 8.50-9.41 7.20 M-8 0 a, sa
Pseudochirella mawso ni 5.42-6.75 5.20-5.66 M 0 a, sa
J Pseudochirel/a obtusa_ 5.25-6.50 4.80-5.50 M-B 0 cosm_Qpo litan excee!.._clr
880 Copepoda
I
Species length length distrib. offshore Latitudina l distri but ion
(mm) (mm) distr.
Pseudochirella pustulifera 6.60-7.40 6.20-6.50 M-B 0 ,a, sa, w, NAt

Pseudoch irella spectabilis 7.17-8.00 6.00-6.66 M 0 bipolar
Pseudochirel/a spinosa 6.20 unknown M? 0 At (182 N-282 S)
Undeuchaeta incisa
Undeuchaeta major
5.70-6.60
4.15-5.50
4.08-5.58
3.90-6.60
M-B
M-B
0
0
At (652 N-352 5), P, sa, a
a, sa, w, At (63gN-52°S), NP
..
Un deuchaeta plumosa
Family Euchaetidae
3.00-4.20 2.85-3.90 E-M 0 w, At (62°N-35 2S) "
Euchaeta acuta 3.60-4.28 3.36-4.08 E 0 t, st, temp, w
? Euchaeta concinna 2.1 0-2.52 2.30 E 0 tiP, tAt
Euchaeta marina 3.40-3.64 2.88-3 .20 E 0 At (52 2 N-37 2 S), Med
Euchaeta media 3.64-4.58 3.56-3.88 E 0 t, st, tem , w
!.A
Euchaeta paraconcinna 2.38-2.56 2.36-2.60 E? 0 At
!.A
Euchaeta pubera 3.90-4.20 2.86-3 .48 E 0 t, st, temp, w, At (370N-09)
!:.A
Euchaeta spinosa 6.20-7.00 5.80-6.60 M 0 t, st, temp, w, At (52°N-37DS)
Paraeuchaeta aequatorialis 4.80-5.40 4.60-4.90 M? 0 t, st, w, SAt (15°-35°S)
Paraeuchaeta antarctica 8.40-9.90 6.60-7.30 M 0 a, At (at 352 S), I (at 452 S)
Paraeuchaeta barbata 7.50 - 11.80 6.70-9."10 M? 0 cosmopolitan
Paraeuchaeta biloba 5.30-6.40 5.1 0-5.40 E-M 0 a, sa, At (S of 35°S), SEP, W I
Paraeuchaeta bisinuata 5.30-5.70 4.70-5.20 B 0 t, st, sa, w, At (602 N-32 2S)
Paraeuchaeta calva 7.40-8.40 6.80-7.90 M? 0 t, st, temp, w
Paraeuchaeta comosa 7.80-10.00 7.50-8.80 B 0 t, st, sa, w
Paraeuchaeta dactylifera 8.90-10.00 unknown B 0 sa, At (at 352 S)
Paraeuchaeta exigua 6.40-7.30 5.30-6.00 B 0 st, sa, P, I, At {30°-36°S)
Paraeuchaeta gracilis 6.40-6.90 5.20-5.80 M? 0 At (62°N-32°S)
Si
Paraeuchaeta hansenii 8.10-9.70 8.10-8.90 M 0 sa, w, At (602 N-262 S)
s.
Paraeuchaeta kurilensis 5.83-7.90 5.91-6.90 B 0 a, w, At (49°N-32°S)
Si
Paraeuchaeta parvula 7.50-8.50 6.40-6.80 B 0 a, sa, At (S of 262 5), SEP, W I
Paraeuchaeta pseudotonsa 5.80-7.30 5.70-6.80 B 0 a, sa, At (65 2 N-36 2S), SP, 51 _____.
~
Paraeuchaeta rasa 5.80-6.40 5.00-5 .80 M-B 0 a, sa, At (S of 312 5)
Si
Paraeuchaeta regalis 8.80-9.60 8.00-8.80 B 0 a, sa, SAt (3 12 -362 S), SEP ;5ij
Paraeuchaeta sarsi 7.80-11 .30 6.80-9.40 B 0 t, st, sa, a, w, At (61 2 N-57 2S)
~
Paraeuchaeta scotti 5.70-6.60 5.40-6.00 B t, st, w, At (62 2 N-342 S)
0
s.
Paraeuchaeta sesquipedalis 8.90-9.80 8.50 B? 0 SEAt, NEP
5I
Paraeuchaeta vervoorti 8.30-10.80 unknown B? 0 SEAt, P
Valdiviel/a brevicomis 6.83-7.25 5.00 B? 0 a, w
0
Valdiviella insignis 10.00-12.00 8.7-10.00 B? 0 a,w ti)
Valdiviella minor 5.41-5.44 4.96 B 0 a,w
!ZJ
Valdiviel/a oligarthra 9.33-9.75 8.25-9.08 B 0 a,w
Family Phaennidae
Brachyca/anus bjombergae 1.60-2.00 unknown BP c Brazil
Comucalanus chelifer 7.41-8.70 6.35-6.66 M-B? 0 a, sa, w
Cornucalanus robustus 6.75-7.66 6. 16-6.75 B 0 a, sa, SAt (at 342 5), SEP
Neosco/ecithrix caetanovi 3.20-3.90 3.40 BP c Brazil
Onchocalanus magnus 8.30-9.08 7.83-8.33 B 0 a, sa, I, SEP
Onchocalanus subcristatus 7.00 unknown B 0 a
O nchoca/anus trigoniceps 7.83 -9.16 6.50-6 .75 B 0 a, sa, w
Onchoca/anus wolfendeni 6.00-6.91 5.33-5.66 B 0 a, sa, At (at 342 5)
Phaenna spinifera 1.80-2.90 1.80-2 .50 M 0 t, st, w
Copepoda 88 1
---,
Fe male Male Vertical Inshore/

Species length length distrib. offshore Latitudina l distribution
(mm) (mm) distr.
Xanthocalanus agilis 2.40 2.58 M? 0 At, IW P?

1
Xanthocalanus m arlyae 3.30-3.75 unknown BP c Brazi l ~
Xan!hoca/anus soaresmoreirai unknown 1.58 B? 0 Brazi l
Fa mily Scolecitrichidae
Amal/oth rix dentipes 2.36-2.84 2.72-3.00 M? 0 a, sa, SAt (at 36°S)
Am al/othrix emarginata 3.65-5.60 3.75-4.25 M? 0 sa, w, At (65°N-552 S)
Amal/othrix hadrosoma 5 .00 -5.66 unknown M-B? 0 a, sa, SEP (at 34 2 S)
A m allothrix o btusifrons 5.25-6.3 5 un known ? M-B? 0 a, At
A mal/othrix valida 2.10-4.40 4.00-4.64 B 0 a, w
Landrum ius gigas 7.58-8.58 unknown M-B 0 SWAt, 5&WP
Lophothrix frontalis 5 .83-7.25 5 .58-5.91 B 0 sa, w, At (652 N-35 2 S)
Lophothrix humilifrons 5.50-7.33 6. 35 B? 0 a, sa, te mp, w
Lophothrix latipes 2.96-3.19 unknown ? M o . t, st, te mp, w
Lophothrix simi/is
L.EE_hothrix quadrispinosa
5 .50
5.60
u nknown M- B?
unknowl"l___j_M?
rlSAt,
o
SEP
SAt, I
Scaphoca/anus antarcticus 5.00-5.83 5.16-5.75 M? o a, SAt (at 342 S)
Scaphocalanus brevicornis 1.90-2.84 2.00-2.50 M? .2______a r, w
Scaphocalanus curtus 0.93-1.30 1.22-1.35 E-M _()_______J' s t' w
Scaphoca/anus echinacus 2.16-2.56 2.12-2.36 E-M 0 t, st, te mp, w
Scaphoca/anus elongacus 2.-6-3.20 3.16-3.-t-4 Vl-8? 0 a, sa, At, IW P, NWP
Scaphocalanus farrani 1.40-1.91 2.68-3.28 \11 0 a, sa
Scaphocalanus m agnus 4.25-4.75 3.50-4.7-t VI? 0 w
Scaphoca/anus parantarcticus 5.08-5.83 4.83-.5 .08 w 0 a, sa
Scolecithricella abyssalis 1.87-1.92 1.45 E-t'v\? 0 w
Scolecithricella dentata 1.21-2.07 1.3-1.85 E? 0 t, st, w
Scolecithrice lla m inor 1 .0 8-1.46 1 .2-1.46 E 0 a, sa, w
Scolecithricel/a ovata 1 .70-2.39 1 .38-1.80 E-M? _<:>_____<~, sa, w
Sco /ecithrice lla profunda --~ .08-2.04 unava ilab le M? 0 sa, w?

<< 5co lecithricella pse udoc ulata» 1.71 unknown BP c Bra zil
Scolecithricella tenuiserrata .00-1.1 5 1.19-1.33 E C, 0 w
Sco /ecithricella vittata 1.40-2.0 0 1 .62-1.72 E-M 0 t, st , t~ , w
Scolecithrix bradyi 1. 10-1.61 1.30-1.56 M? 0 t, st, w

Scolecithrix danae 1.80-2.40 1 .9 7-2.21 E 0 t, st, w
Scottoca/an us securifrons 3.38-4 .60 4 .50-5.30 B 0 t, st, te mp , w, At (63 2 N-362 S)
Fa mily Diaixidae
Diaixis helenae 0.77-0.88 0 .70 BP c SEAt
Diaixis hib ernica tropica 0 .94-1.04 u nknown BP c SEAt
Diaixis truno vi 0.84-0.98 0 .76-0.84 BP c SEAt
f•m;ly Tl•••~
Tharx.b is asymmetrica 1.12 unknown BP c? SEAt (at 1 7°S)
Tharybis megalodactyla 1.24 1.06 BP c? SEAt (at 1 7°S)
Fa mily Mesaioke ratidae ,----------
Mesaiokeras mikhailini unknown 0 .54 0 -780 m c? SEAt (a t 18°S)
Mesaiokeras semiplenus _ , _0 .40-0.50 0.40-0.42 BP c SE&SWAt
Mesa io keras tantillus 0. 32-0.33 - - -0 .29 BP? c SEAt
---
Family Arie tellidae
---
Ariete l/us-aculeatus 4.63 At, P, I
-- --5-.00- 3 .60-4.08 M 0
A rietellus giesbrechti 4 .73-5 .70 5 .28-5.35 M 0 w
882 Copepoda

Species length length distrib. offshore Latitudinal distribution
(mm) (mm) distr.
Arietellus minor 3.30 unavai lable M 0 Ascension Is.

Arietel/us pavoninus 4.80 unknown M 0 P, At ~
Arietel/us plumifer 4.55-5.60 5.25-5.90 M 0 w :=-:i
Arietel/us setosus 4.60-4.75 3.93-4.45 M 0 w
Arietellus simplex 4.75-6.90 4.85-6.02 M-B 0 w
Paramisophria gise/ae 2.55-2.60 unknown BP c Brazi l
Paraugapti/us buchani 3.25-3.63 2.80-3.08 M 0 w
Pilarella /ongicornis 1.53-1.73 unknown BP c Brazil
Family Augaptilidae
Augapti/us anceps 3.80-5.00 4.23 M 0 N&tAt, P n
Augaptilus glacialis 4.10-5.30 4.10-5.30 M 0 a, ar, w
Poi
Augaptilus longicaudatus 3 .50-6.10 3.25-3.80 M 0 t, st, temp, w, At (52 2 N-35"5)
Augaptilus megalurus 4.30-6.10 4.00-5.00 M 0 a, w, At (42~N -55 "S)
A.ugaptilus spinifrons 3.00-3.55 unavai lable E-M 0 N&tAt, NP
Centraugaptilus cucullatus 5 .10-6.70 unknown M 0 At, P
Centraugaptilus horridus 5 .60-1 0.00 8 .33-8.40 M 0 N&tAt, 1-P
Centraugaptilus rattrayi 4.70-6.18 4.98 M 0 a, sa, temp, w, At (65"N-55"5)
Euaugaptilus affinis 5.40 unknown M 0 N&SAt, I
Euaugaptilus aliquantus 5.9 1-6.41 unknown M 0 a, SWAt
Euaugaptilus angustus 6. 10-7.9 unknown M 0 w, At (392 N-02 )
Euaugapti/us antarcticus 9.16-10.33 9. 16-9 .83 M 0 a
Euaugapti/us austrinus 8.35-9.20 8.59 M-B 0 a, SWAt
Euaugaptilus bullifer 4.90-6.08 4.70 M 0 w
Euaugaptilus c/avatus 3.70-4.00 unknown M 0 N&eqAt
Euaugaptilus e/ongatus 5.30-6.70 unavailable M 0 w, At (692 N-02 )
Euaugaptilus facilis 4.00-5.90 5.00 M-B 0 w, At (62 2 N-02 )
w, At (63"N-0°) ~
Euaugapti/us fi/igerus 4.50-6.80 4.10-5.30 M 0 I
2
w, At (54 N-12 S) 2
Euaugaptilus gibbus 2.70-3.53 3.80 B 0
Eu~ tilus hecticus 2.45-2.75 2.40 E-M 0 w

Euaugapti/us /aticeps 6.09-7.91 6.30-6.80 M 0 a, w, At (63 2 N-02 )
Euaugapti/us /atifrons 4.70-5.60 unknown M 0 At, I ~
Euaugaptilus /ongimanus 5.30-5.80---+- 6.30 M 0 w, At (41 UN-11 2 5) !!!I
Euaugaptilus mag!!_US 7.33-8.58 8.08-8.25 M-B 0 w !.I
Euaugaptilus maxillaris 3.27-6.40 5. 12 M-B 0 At, I !.1
Euaugaptilus nodifrons 4.85-8.75 4.80-8.08 M 0 w, At (47 2 N- 322 5) !..1
Euaugaptilus oblongus 5.40-7.40 5.40-5.70 M 0 w, At (63 2 N-0 2 ) :..11
Euaugaptilus pachychaeta 7.00 unknown M 0 eq At a
Euaugaptilus p alumbii 2.25 1.95 M 0 w u
Euaugaptilus perasetosus 7.08-7.75 unknown M-B 0 a, SWAt u
Euaugaptilus rigidus 4.30-5.50 4.30 M 0 NAt, EP, NWP, G. of Gu inea u
Euaugapti/us squamatus 5.40-6.30 5 .90-6.40 M 0 N&SAt, P u
Euaugaptilus tenuispinus 4.50-5.30 unknown M 0 At, I u
Ha/Qe_tilus acutifrons 2.60-3.00 2.10-2.40 M 0 w ~
Haloptilus angusticeps 3.10-3 .60 3.04-3. 12 B 0 w, At (43 2 N-0") !J

Haloptilus austini 3.30 unknown E 0 N&tAt, NP !J
Haloptilus chierchiae 4.80-5.10 unknown E 0 N&eqAt, P, I !J
Haloptilus fertilis unknown 2.85-3.20 E 0 tAt & P
Haloptilus fons 4 .80 unknown M 0 N&eqAt, NP G
Copepoda 883
,~

Species length length distrib. offshor e latitudinal distribution
(mm) (mm) distr.
IH afoptilus fongicirrus
~~titus fo~orni£_ _
H a}_Qpt ilus majOI_ _ _ _
J
2.84-3.10
1.94-2.50
7.35
unknown
1.18
j unknown _
t
_
M
M _
M__
_j
o
_2
__j__
N&tAt, NP
st,J_em&_w, At (502 -32 N_L
~~qAt _ _ _ _
Hafoptilus mucronatus 3.10-3.60 2.17-2.28 E o t,st, w 1
H afoptilus ace/latus 7.90-8.80 4.1 0 E o a
Halof!iilus ornatu2._ 4.3 0-4.80 2.75-3.05 E _ .Q_ w, ~_(52 2N - 02)_ _
- f
Haloptilus oxycepha/us
H afoptilus pfumosus
3.31 -3.80
4.00-4.70
2.50
unknow n?
1 E-M E-M
o a, sa, w
o t, st, At, P
Hafoptilus spiniceps 3.80-5.45 2.55 E o It, st, w
Pachyptifus abbreviatus 3.90-6.1 unknown M o N&tAt,J -P
Pachyptilus longimanus 6.72 unknown I M -B o SAt
- _]
PonJQptilus ova l is 4.1 6-5.62 j_ unknow_r1_ _ M -B_ _ j __Q J N&tAt, I ____
I
_ _ _
l oisseta palumbi!_
_ _Fami ly Heterorhabdidae _
Heterorhabdus abyssalis
Heterorhabdus austrinus
_ _ _ _
_ _
__?.59-7.50
2.70-3.5 5
3.06-3.65
--f-
_§ .4 5 -7 .~
2.60-3.40
2.84-3.42
M-B
M
M
L O
0
0
a, ~w, At (562 N-552$)
At, I, P
a, sa
Heterorhabdus cfausii 2.40 -2.65 2.20-2.40 M-B 0 It, st, w
Heterorhabdus compactus 2.23-3.35 2.20 B 0 At, NP
j Heterorhabdus fobatus 2.10-2.60 2.00-2.50 M 0 st, sa
H eterorhabdus norv~cus 2.80-4.20 2.65-4.00 M-B N&SAt
H eterorhabdus papilliger 1.62-2.35 1.60-2.30 M
0
0 w
---1
H eterorhabdus robustus 3A8-3.70 3.36-3.60 M 0 &tAt, P
j Heterorhabdus~inifrons _ 2.80-3.40 2.85-3.40 M 0 _0t, p
H eterorhabdus tanneri 4.01 3.75-3.85 M 0 _iA~
IHeterorhabdus viper;- - 2.80 2.60-3.70 M 0 SAt, P
H eterostyfites fongicornis 2.30-3.40 2.70-3.68 M 0 t,st, w
Heterostylites m~r 4.00-5.40 3.80-5.09 M 0 a, sa, w
I Mesorhabdus angustus 5.80 unknown 1 B 0 At
Neorhabdus falciformis 5.60 unavailable B 0 SAt
Family lucicutiidae
I
Lucicutia aurita
Lucicutia bicornuta
Lucicutia cfausii
- --
7.30-8.70
6.80-8.40
~0- 2.10 _
I _.z..10-8.40_
6.90-7.70
1.60-1.90
+
B
M~ _
M-B _.L 0
o
o
w
w
w
Lucicutia curta 1.90-2.60 1.80-2 .40 M-B o w

Lucicutia flavicornis ~30-2 .00 j !.]0-1 .70 _ - +--- M o - w
Lucicutia g_aussae 1.30-1.50 1 .20-1 .40 M o w
Luc icutia gem ina 1.40-1.90 1.30-1.70 E-M 0 w
I Lucicutia grandis 4.40-6.50 3.90-4.90 M 0 w
J Lucicutia fongicornis 1.50-2.00 1.40-1.80 M 0 w
Lucicutia Iucida 3.50 3.30-3.50 M 0 w
Lucicutia macrocera
- -- 3.30-4. 10 3.60-4.30 l
M 0 w
Lucicutia magna 3.40-3.90 3.10-3.40 M 0 w
Lucicutia major 8 .00-8.20 unknown I
B 0 At
Lucicutia maxima 7.80-9.30 7.70 -7.80 B 0 w
Lucicutia ovalis 1.50-1.80 1.20-1 .50 M o w
I Lucicutia wolfendeni 6.00-9.80 6.00-8.30 M -B o w
Fa mily Metridinidae
IGaussia- asymmetrica
-
9.00-11 .00 unknown M-B o -1.SW At
884 Copepoda

Species le ngth le ngth distrib. offshore latitudinal distribution
(mm) (mm) distr.
Caussia_princeps
Metridia brevicauda
------
10.50-12 .00 9.40-10.40
1.80-2.25 1 .50-1.65
M-B
!±L...... 010 w
Metridia calypsoi
Metridia curticauda
Metridia discreta
2.95
2.25-3.60
3.35
l unavailable
1 .80-2.30
unknown
M?
M
B
o
o
o
Braz il
P,At
N&tAt, I
Metridia gerlachei 3.38-4.25 2.16-3.00 M o a
Metridia longa 4.1 0-4.l_()__3.50-3.70 E- B o w
Metridia lucens 2 .39-2.93 1.62-2.30 M o a, sa, w, At (81 2 N-55 2S)
-u
Metridia princ.!!0.__ 7.65 -9.00 5.80-8. 15 M-B o a, sa, w, At (652 N-35 2S)
Metridia venusta - - - - - 2 90-3.30 2.50-2.60 M-B o w
Pieuromamma abdomina/is 40-3.70 2.75-3.50 E-M o t, st, w
4
Pleuromamma E-M o t, st, w
- - -borealis
-- - - - .80-2.25 1.72-2.25
Pieuromamma gracilis I 1.70-2.15 1.60-1.90 ( E-M o t, st, w •
Pleuromamma pisek_i_ _ 1. 7 0-2 . 0~ .60-1 ~ E-M o~,w •
Pleuromamma q uadrung!!_lata _ _3.30-4.9 8 3.06-3 .88 E-M o t,s~
Pleuromamma robusta 3.00-4.30 3.00-4 .00 E-M o t, st, sa, w
Pleuromamma xiphias 4.10-4.90 4.00-4.70 E-M o t, st, w
Family Phyllopodidae
Phyi!Qe_us aequalis 3.00 unknown M
__:J,At
FJ2yllopus bidenta ws_ _ _ 2.40-3.00 2.25 E-M w
Phyllopus helgae 2.25-2 .46 2.10-2 .46 E-M 0 a, sa, w, NAt
Phyllopus impar 2.20-3.00 2.62-2.95 B 0 w
Family Centropagidae
Centropages brachiatus 1.73-2.30 1.58-1.90 E n _ 5W&SAfr, SAm
1
Centropages bradyi _ _ 2.00-2 .50 1.90-2.40 E 0 stAt, P
Centropages calaninus 1.90-2 .00 1.80 E 0, c IW P, SEAt
Centropages chierchiae_ _ 1.60-2.26 1.54-2.02 E c t, st, At, I
--~
Centropages elongatus 1.50-1.60 unavailable E o, c IW P, SEAt
Centropa~s furcatus 1.60-1.92 1.5S-1.92 E c,o t, st, w
Centropages gracilis 1.85-1.90 1.80 E ___s_Q__ t, st, w
---
Centropages kroyeri 1.25-1.35 1.20 E 0 t, st, w
Centropages longicorn~ 1.97 unknown E o? NW P, SAfr
-·-
E
Centropages orsinii
Centropages typicus
1.50-1.60
1.60-2.00
1.25-1 .30
---
1.40-1.90 E
o? IWP, SEAt
n Med, stAt, WAfr
-j
Centropages violaceu_s_ _ 1.76-1.92 1.77-1.86 E 0 t, s~ _l
Family Pseudodiaptomidae
Pseudodiaptomus acutus 1.00-1 .28 0 .89-0.99 E e eq Braz il
Pseudodiaptomus chartetj__ ~-1 .~ 1 .25-1 .30 E e SAfr
Pseudodiaptomus hessei 1 .20-1.80 1.05-1.45 E e W&SAfr
Pseudodiaptomus marshi 1 .30-1 .53 0.96-1.07 E e~ Brazil
Pseudodiaptomus nudus 1 .30-1.40 1.1 5-1.20 E n,_ c ~SAfr
Pseudodiaptomus richardi__ 1 .22-1.36 0.92-1.05 E e SAm (02 -302 5)

Pseudodiaptomus serricaudatus 1.12 1.1 5 E e, c tWAfr, I, Red Sea
Family Temoridae
Temoropia ~umbae~ 0.70-1 .05 0.84-1. 11 M 0 w
---r--
Temora discaudata 1.70-2 .00 , 1.70-1.9 0 E c,o t,st, IP
~
___
Temo ra longicornis "1.00-1.50 1.00-1.35 E c, 0 t & NAt
Temora stylifera 1 .60-1.85 1.65-1 .80 E C,.2__ t, st, w
---
Copepoda 885
Female Male
Vertical Inshore/
• Species length length distrib. offshore Latitudinal distribution
(mm) (mm) distr.
ITemora turbinata 1.05-1.61 0.93-1.56 E C,O t, st, w (not EP)

Family Candaci idae
Candacia armata 2.50-2.75 2.70- + - E 0 w
Candacia bipinnata 2.35-2.65 2.15-2.50 E-M 0 l, Sl, w
Candacia catula 1.45-1.60 1.40-1.55 T
E? 0 IWP, SEAt
- - --
Candacia cheirura 2.25-3.00 2.03-2.35 E-M 0 sa
Candacia curta 1.82-2.70 1.90-2.65 E? 0 t, st, w
--,
Candacia elong_ata 3.00-3.25 3.15-3.30 E-M 0 w
Candacia ethiopica 2.15-3.00 2.05-2.35 E-M 0 t, st, w
1
Candacia longimana 2.90-3.90 1.40-3.50 E-M 0 t, st, w
Candacia magna -U().....t.-la 3.90-4.36 M 0 G. of Guinea, I
Candacia norvegica 3.10 3.10 E 0 w
Candacia pachydactyla 2.30-2.90 2.30-1.- 5 E-M 0 t, st, w __j
Candacia paenelongimana :?.5-1-2.9:? 2 ..!9 E 0 tAt
Candacia cenuimana :?.05 2.10 \.1 0 w
Gmdacia \aricans 2.15-2.50 1.10-1 ..!0 E-\.1 0 t, st, w
ParaG3Ildacia bL-piflOSa 1.80-1.82 o.-o E 0 w
~~-a~a ; ·mpoe>.. 1.80-2.30 1.80-2.05 E 0 t,st,w
Paracandacia rruncata 1.10 2.10 E? 0 IWP, SEAt
Family Pontellidae
Calanopia americana 1.30-1.50 1.20-1.40 E n tAt
Calanopia elliptica 1.40-1.90 1.80 E 0 t, st,w
Calanopia minor 1.40 1.20 E C, 0 t, w
Labidocera acuta 3.05-3.40 2.80-3.30 E n IWP, SEAt
Labidocera acutifrons 3.41-4.25 3.43-4.04 E 0 t, st, w
Labidocera darwinii
Labidocera detruncata
2.00
2.25-2.80
1.50-2.00
2.15-2.50
E
E j n
0
SAm (25°-38°S)
IWP, SEAt
_j
Labidocera fluviatilis 2.50 2.40-2.50 E n SAm (20°N-36°S)
I
Labidocera minuta 1.95-2.05 1.65 E 0 IWP, SEAt
Labidocera nerii 2.85-3.70 2.65-3.56 E 0 eqAt
Labidocera scotti 2.40-3.05 2.15-2.55 E n tAt
Labidocera wollastoni 2.20-2.30 2.20-2.30 E 0 t, st, At
Pontella atlantica 5.80-5.90 4.30-5.70 E 0 t,w
Pontella diagonalis 4.00 unknown E 0 I, SAfr
Pontella fera 2.70 2.50-2.60 E 0 1t,w
Pontel/a gaboonensis 2.70-2.95 2.65-2.68 E n tEAt
Pontel/a lobiancoi 3.95-4.20 3.30-3.75 E o?
!At?
Pontella marplatensis 2.50 2.70 E n Mar del Plata
Pontella securifer
Pontel/a spi!!i£es
Ponte//ina platychela
3.65-4. 55
4.70
1.54-1.96
3.42-4.20
unknown
1.41-1.74
+- E
E
E
0
0
0
t,w
t, st, w
eqAt, WAt to 40°S
Pontellina plumata 1.44-1.94 1.34-1 .92 E 0 t, st, w
Pontellopsis brevis 2.00-2.58 1.99-2.15 E 0 t, st, At, SAfr, P? I?
Pontellopsis perspicax 3.00 2.50 E 0 t, st, At, P
Pontellopsis ~lis 4.00-4.40 3.40-3.50 E 0 t, st, w
I
Pontellopsis villosa 2.20-3.00 1.80-2.83 E 0 t, st, w
Family Acartiidae
IAcartia bifilosa 1.02-1.10 1.00-1. 10 E o? NAt, SAfr, NWP
--- -
886 Copepoda
~ T
Fema le Male Vertical Inshore/
Species length length ~~ distrib. offshore Latitudinal distribution
(mm) (mm) distr.
________j__ - - -
-
r - E-
- - - - 1.20-1.24 -+ 1 .03 o? tSAt c
\ Acartia centrura r--- --
Acartia danae 1 .05-1 .34 0. 72-0.86 E o Is, st, w
_!lcartia ljjjjebQ!Sj_ _ _:!..]3-1 .40 I 1.10 E
E
e, cL!l3 r<li!.!_
o _J, st, w
Acartia negfigens 1.04-1.27 0.80-1.05
Acartia plumosa 0.87-1.21 0.89-1.13 I E e }EAt, NWP l
Acartia tonsa 0.90-1.50 1.00-1.10 E e, c, o t, st,'!:!...._
I
Paracartia africana 1.20-1 .45 1.12-1.30 E c SWAfr
Family Tortanidae
Tortanus (Tortanus) gracilis _ _ _ 1.5_
2-_2 .1 0 1.50 l E _ oJ WP, SE~
Family Bathypontiidae
5.20-5.60 4.90-5.53 M-B 0 tAt
Bathypontia elongata
Bathypontia sp. 2.81 unknown B 0 \tAt
I
Order Cyclopoida
Family Oithonidae (J
Oithona atlantica 1.00-1.43 0.82 E 0
lst, w (J
0.50-0.90 0.40-0.53 E c, o eq l&P, eqEAt, off SAfr
Oithona attenuata
0.44-0.51 0.43 -0.4 5 E e North of Braz iI
Oithona bjornbergae (J
Oithona brevicornis 0.58-0.72 0.47-0.57 E e jc ongo R, tWP, tNI (J
Oithona fa/lax 0.85-0.96 0.67-0.70 E n, o ti&P, eqAt, saAt (J
Oithona frigida 1.20-1.24 0.84 E 0 a

1 .06-1.35 unknown 1 E 0 eq, t,w ()
Oithona hamata
Oithona hebes 0.4 7-0.55 0.45-0.57 E c, e eq, tWAt & EP ()
Oithona m inuta 0.45-0.65 0.42-0.50 E c, e Gulf of Guinea

Oithona nana 0.49-0. 72 0.44-0.53 E n, c, e jeq, t, st, w
0.6 2-0.80 0.62-0.65 E n, c, e t, st, w
O ithona oculata
- I c, e eq, t, st, WAt
r·
O ithona oswaldocruz i 0.51-0.71 0.46-0.65 E
1.06-1.51 0.59-0.68 E C, 0 t, st, w
Oithona plumifera
Oithon~seudofrigida
Oithona rigida
O ithona robusta
1.22-1.30
0.75-0.83
1.42-1 .59
unknown
0.55-0.66
1.20- 1.24
t- M
E
E
c,e
c, o____1_sl, WP, I, EAt
t, I&P, Rod S.•, offSAft
c, o t, st, l&P, tAt
Oithona setigera 1 .14-1.90 0.90 E ~ c,o t,~w _ - - -
- - c, o w, a, sa, b1polar
Oithona simi/is
Oithona si'!!E!_ex _
0.68-0.96
0.36-0.41
0.67-0.70
0.37-0.41 I E
E _ _c, _()__eq, t, st, w
Oithona tenuis
Oithona vivida
Paroithona flemingeri
1.10-1.25
0.69-0.83
0.40-0.45
un known
unknown
unknown
1 E
E
E
C, 0 eq, t, st, l&P, eqEAt,SWAt
c,o eq, t, st, l&P, NEAt, WAt
c,~ _ eq, t, st, WAt _
O rder Harpactico ida

Fam i!r.!ctinosomatid~ ~ -
Microsetel/a norv~ca 0.35-0.53 0.33-0.42 E o, c w

0.64-0.85 un known? E o w
M. rosea
Family Miraciidae_ _
Macrosetel/a gracilis 1.40-1.50 1.13-1 .1 6 I E 0 t, st, '!'!...__
IM.
M iracia efferata
minor
1.45-2.00
0.93
1.40-1.60
unavai lable
E
E
w 0
o_ \l,At
Ocu losetel~acilis 1 .20-1.35 1 .1 5-1.30_ ] E _ o...J.!,st,w - _j
Family Euterpin idae

Euterpina acutifron_s _ - - - - 0.50-0. 75 0.50-0.56 E c ....Lw _
Copepoda 887
Species
r Female
length
Male
length
I Vertical Inshore/
distrib. offshore latitudina l distribution
(mm) (mm) distr.
_j_ _j _ _j
Family Clytemnestridae
Qytemnestra rostrata 0.60 - 1.00 0.80-0.90 E-M o t, st, w
C. scute/lata 1.00-1.24 1.05-1.30 E-M o t, st, w
Family_Aegisthidae
Aegisthus aculeatus 1.85 unknown M o w
A. mucronatus 2.55 unavailable? M-B J o w
Order Poecilostomatoida
~mi ly Oncaeidae
Conaea_!!Jg)ressa 0.67 0.60 M tSAt
Conaea rapax 0 .97-1.14 0 .87-1. 00 M 0 w
Epicalymma exigua 0.48-0.52 0.42-0.44 M o tSAt, NAt, M
Lubbockia aculeata 2.21-2.30 2 .35 E-M o t, st, w
Lubbockia squi/limana 1.45-1 .60 1.80-2. 1 E-M o t, st, w
Oncaea antarctica 1.1 5-1.26 0.59-0.63 E-M o a
•oncaea" atlantica 0 .25-0.26 0.24 E 0 tAt
Oncaea clevei 0.68-0.76 0 .46 E o P, At
Oncaea curvata 0.59-0.67 0.48-0.49 E-M o a
Oncaea eng!._ish t_·_
Onccea lacinia
Oncilea /acimana
Oncaea media
_ 0.95-1.1 5
0 .40-0.48
0 .50
0.61 -0.96
0.89-0.90
0.38-0.46_.
0.42
0.59-0.67
E-B
E-B
M
E-M
o
o
o
0
win P& Med
w in P
tSAt
w
--1
Oncaea mediterranea 1.14-1.26 0.74-0.97 E-M 0
Oncaea pari/a 0 .61-0. 73 0.50-0.58 M? 0
Oncaea prolata 0 .64-0.72 0.50 E-B 0
Oncaea scottodicarloi 0.59-0.78 0.53-0 .60 E 0 w
Pachos tuberosum 2.00 unknown E o _ At,P
Familv ~rinidae
Copi ia hendorifi 3.90-5.10 5.50-8.30 E 0 t. st. I. SEAt
C.Q{Ji "a ar.a 3.20..5 ..!0 .!.5o..-.oo E 0 t,w
Caoifu r:JeO ·~ 3.20-! ..!0 .!.50..6.i0 E-\1 0 t, st. w
Capwa ~lrabilli 2.20..-UO 3.20..6.10 E- 'v\ 0 t, st, "'
C!XX a q-..:.aara:a 2.20-4 ...!0 3.:>o..s.;-o E-.'v\ o t, st,w
Cop ilia ~ 'liTed 3.20-5.40 5.50-9.00 E-M o t, w
~l
Sapphmna angusra 2.50-5.50 4 .00-7.60 E o t, st, w
Sapphirina auronitens-sinuicauda 1.42-2.81 1 .42-2.75 E o t, st, w
S~phirina bicu~ida ta 2.26-3 .00 2.58-3.42 E o _ t.~
Sapphirina_gastrica _ 2.33-2.68 2.20-2.65 E o _ t, '!'!._
Sapphirina intestinata 1.62-2.78 1.65-2.87 E o t, st, w
Sapphirina iris 5 .20-7.40 5 .90-7.50 E o t, st, w
Sapphirina metal/ina 1.68-2.52 1.61 -2.58 E o t, st, w
Sapphirina nigromaculata 1.55-2 .8 1 1.71 -3 .04 E o l, st, w
Sapphirina opalina-darwinii 2.13-4.17 2 .42-4.36 E o t, st, w
Sapphirina ovatolanceolata-gemma . 2 .1 3-4 .1 7 2 .42-4.36 E o t, st, w
Sapphirina scarlata 3.30-4. 65 3 .40-4.85 E o t, st, w
Sapphirina stellata 2.00-2.75 2.00-2.75 E o t, st, w
Family Corycaeidae
Corycaeus (A~us) fla ccus _ 1.49-1.85 1.28-1.68 E --j 0 t, s!,_w _
Corycaeus (A~s) /imbatus _ L 1.35-1 .40 _ 1.15- 1.20_ E 0 t, st, A1_!'
Corycaeus (Agetus) typicus 1.62- 1.65 1.27-1 .62 E 0 t,w
888 Copepoda
I
t
0.95-0.97
otSAm
E
E
J.
¡t,E__w
tlP,
¡tP,
t,
It, st,
offshore
SAfr
5Afr
st,At?
IP,5Afr !P,
w
W
st, ocI! At,
At, 1~5Afr
IP,Ilatitudinal
SAfr NEP,
Pt,0.70-0.90
P? EE SAfr
E distr.
1.35-1.44
1.49-1.85
E0.83
st,
MaleAt, 1 E E EE
P?distribution ".
----;---- (mm)
t
2.02-2.08
0.80-1.18I 1.65 1 ~t,E
,<;_1
t---S?- At, length
I PI tAt
distrib.
50Vertical Inshore/
0.70-0.90 ---1Ej
E
(mm) 1.09 74
2.50-2.57
1 1'.<;_1
0.80-1.02
0.92-0.98
0.72
1.76-2.04 2.08-2.16
0.72-0.80 length0.66-0.73
0.70-0.77
0.64
1.20-1.32
1 I
emale
1 0.84 unknown __
1.57-1.74
------l 1 I0.70-1.17
~ 0.80
2.75-2.94 2.09-2·U
Species
1.04 0.84
Corycaeus (CorycaeusJ c1ausi

Corycaeus (CorycaeusJ crassiusculus .. ~ .._._.~~_._. ~._. ~~~~~_~_
Corycaeus (CorycaeusJ speciosus
Corycaeus (DitrichocorycaeusJ affinis ~_
Corycaeus (Ditrichocorycaeusl amazonicus
Corycaeus (MonocorycaeusJ robustus
Corycaeus (OnychocorycaeusJ agilis
Corycaeus (OnychocorycaeusJ giesbrechti
Corycaeus (OnychocorycaeusJ latus
Corycaeus (OnychocorycaeusJ ovalis
Corycaeus (OnychocorycaeusJ pacific_u_s _
Corycaeus (UrocorycaeusJ furcifer
Corycaeus (UrocorycaeusJ lautus
Corycaeus (UrocorycaeusJ longistylis
Farranula concinna
Farranula curta
Farranula gracilis
Farranula rostrata
Abbreviations:
Vertical distribution
E = epipelagic, M = mesopelagic, B = bathypelagic, A = abyssopelagic, BP = benthopelagic.
Inshore/Offshore distribution
e = estuarine, n = neritic, c = coastal, 0= oceanic.
Latitudinal distribution
a = Antarctic, Afr = Africa, Am = America, ar = Arctic, At = Atlantic, E = eastern, eq = equatorial, I = Indian, IWP = Indo West Pacific,
Med = Mediterranean, N = northern, P = Pacific, S = southern, sa = subantarctic, SH = Southern Hemipshere, st = subtropical,
t = tropical, temp = temperate, Tr = Transition Zone, W = western, W= widespread in tropical and subtropical waters of the
Atlantic, Pacific and Indian Oceans.
Table 1. Morphometric and distributional data for lhe species treated.

Copepoda 889
In the regions between the shore and neighbouring mus, Neocalanus tonsus, Rhincalanus gigas and
oceanic waters there are more restricted habitats Metridia gerlachei. Ca/anoides acutus, Calanus pro-
occupied by coastal species. Coastal regions are high- pinquus and Rhinca/anus gigas are the main members
ly influenced by the land and are inhabited by species of the Antarctic copepod community although not
which are variously adapted to the variability contri- completely restricted by the Antarctic Convergence.
buted by freshwater runoff and coastal upwelling. Neocalanus tonsus plays a dominant role in the sub-
These species may de defined as coastal (e.g. some antarctic zone and north of the Subtropical
species of genera Centropages, Acartia, Paracalanus, Convergence (Bradford-Grieve and Jillett, in press).
Ca/anus, Temora, Oithona). They are restricted to All above-listed dominant copepods are epi- to meso-
varying degrees to the coast. Other species are more pelagic species and are found in the Southern
closely associated with the shore or low salinity water Hemisphere only; Calanoides acutus, Rhincalanus
and may be defined as neritic or brackish water spe- gigas, and Metridia gerlachei are restricted to
cies (e.g. all or some species of Pseudodiaptomus, Antarctic waters, whereas others have Antarctic-sub-
Acartia, Labidocera, Pontella). Whereas other spe- antarctic distributions (see Table l ). They survive
cies such as Ca/anoides carinatus thrive in upwelled winter by dispersing to deep depths and vary in the
water (e.g. Mensah, 1974). The distribution of the timing of their ascent to the upper water layers. In bio-
mostly estuarine genus Pseudodiaptomus illustrates logical spring Calanoides acutus is most abundant in
the degree of local geographic restriction that can be the upper water layers, whereas at this time
found among such species (Fig. 5D). It is amongst Rhincalanus gigas is more abundant in deeper layers.
some of these coastal species that resting eggs are Later Ca/anus propinquus occupies the upper water
known to occur which allow them to survive unfa- layers and is followed by R. gigas at slightly deeper
vourable conditions (e.g. Uye, 1985). depths. In the subantarctic surface waters Neocalanus
tonsus dominates in the north and Calanus simillimus
In addition to the already noted extension of tropical ·in the south. Rhincalanus gigas also dominates in
species down the coast of Brazil, other regions where southern subantarctic waters but in the deeper water
strong currents dominate usually differ from neigh- layers (Voroni na, 1984: 185).
bouring areas in that species penetrate with the cur-
rents. For example, the presence of Macrosetella gra- Although the evidence is not complete, it appears that
cilis and Oithona rigida in the southeastern Atlantic in the South Atlantic from the equator to 60°S. the
indicates the Agulhas Current and its retroflection frontal zones of the various water masses have an in-
south of South Africa (De Decker, 1984), and also fluence on the distributional patterns of many pelagic
demonstrates an extension of the Indian Ocean distri- oceanic copepods. This pattern is often obscured
bution of the species (Fig. 5C). In the Gulf of Guinea when a species has a vertical distribution which
and in the Benguela current Calanoides carinatus and crosses the vertical boundaries of a water mass.
.\ferridia lucens dominate, whereas the neighbouring
tropical Atlantic pelagic conununity is dominated by Vertical distribution
Subeucalanus subtenuis and Nannocalanus minor
(see Fig. 12 in Heinrich, 1993). The main reason for The geographic distribution of pelagic animals is close-
the temporal changes in species composition and dif- ly linked with their bathymetric distribution. The habi-
ferences in biomass dominance in these tropical com- tat of pelagic copepods may be subdivided into the fol-
munities is the response to upwelling rather than lowing vertical zones: 0-200 m: epipelagic; 200-750
being the result of horizontal advection. (1000) m: mesopelagic; 750 (1000) - 3000 m: bathype-
lagic. and deeper than 3000 m: abyssal (Vinogradov,
In the polar and subpolar regions in the spring-sum- 1968). Species and even taxa of higher rank may be
mer season species diversity is low and the biomass of strongly connected with a certain layer. For example, in
one to three species in the upper layers may reach the South Atlantic Nannocalanus minor, Undinula vul-
80%, whereas in the tropical oceanic zooplankton garis, Subeucalanus subtenuis , S. pileatus,
diversity is much higher. Calanoid species common in Paracalanidae, all Clausocalanus species, Euchaeta
the Atlantic sector of the Antarctic-Subantarctic are: marina, Pontellidae, Temora turbinata are conspicuous
Calanoides acutus, Calanus propinquus, C. similli- members of the epipelagic copepod fauna. They remain
890 Copepoda
in the uppermost layers all their life. On the other hand, check. If this fails then we refer you to the revisions
Megacalanidae, Bathypontiidae, Pseudochirella mentioned in the text and original descriptions.
(Aetideidae), Paraeuchaeta, Valdiviella (Euchaetidae),
the majority of Spinocalanidae and other meso- or Abbreviations used in the figures and keys (see Fig.
bathypelagic species avoid the uppermost layers. 1 and 2)
However, our knowledge of vertical distributions is A I : antennule
often incomplete and those that are better known do not A2: antenna
conform exactly to particular depth classifications. B: basis 0
Many copepods migrate vertically and may be found at C: coxa
different depths at different times of the day, in differ- Ce: cephalosome (the head region when this includes -.J
ent seasons, at different latitudes, and at different devel- only the somite of the maxillipeds)
opmental stages. CR: caudal rami
Enp: endopod; Enpi -3 - endopod segments I-3
A number of species are recorded from Brazilian coast- Exp: exopod; Exp I-3- exopod segments 1-3
al and slope waters which are interpreted to live in Gn: gnathobase
association with the sea floor. These species belong to Gns: genital somite
the families Aetideidae, Phaennidae, Scolecitrichidae, Md: mandible
Diaixidae, Tharybidae, Mesaiokeratidae, and Arie- Mdp: mandibular palp
tellidae (Table 1). A few epipelagic copepod species Mx I: max itlule
inhabit the neustonic environment and live in close Le 1: coxal epipodite- first external lobe
association with the thin fi lm at the very sea surface Le 2: basal exite- second external lobe
(Champalbert, 1980). These species almost exclusive- Li I: praecoxal arthrite - first internal lobe
ly belong to the Pontellidae (e.g. Turner et a/. , 1979; Li 2: coxal endite- second intemallobe
Champalbert, 1985; Ohtsuka, 1985) although other Li 3, 4: basal endites - third and fourth intem al
copepods may be found in this environment (Hattori et lobes
a/., 1983; Nair et al., 1989). Such copepods may be Mx 2: maxilla
fu rther subdivided according to their degree of attach- Li I, 2: praecoxal endites - first and second lobes
ment to the sea surface (Matsuo and Marumo, 1982). Li 3, 4: coxal endites - third and foutth lobes
Li 5: basal endite - fifth lobe
Li 6: (if present) - sixth lobe
Identification Mxp: maxilliped
Li I: praecoxal endite (not figured)
Use of the Keys Li 2-4: syncoxal endites (not figured)
Pl -5: swimming legs 1-5. In the keys PI -P4 segmen-
Keys are given here to orders, families, genera, and tation is denoted as, e.g., 3/3 (number of Exp seg-
species although, in a few cases, keys cover more than ments/number of Enp segments). The spine (Roman
one of these taxonomic levels where there is only one numerals) and seta (Arabic numerals) formula for the
or a few species in the group. These keys have been swimming legs is given in the mode of Sewell (1949)
constructed from sometimes inadequate information (see Fig. 2). T he outer elements are given fi rst and are
on the morphology of each species and its variability. separated by a hyphen from the inner elements. The
Also it is likely that the fauna of the South Atlantic is terminal segment of each ramus has 3 components,
incompletely known and that not all species have been separated by conunas, with the sequence: outer, ter-
included in the keys. Only those families which have minal, inner. P5 of Euchaetidae males (see Fig.
been recorded from the South Atlantic have been 6.7M): the following abbreviations are used, dp: digi-
included here. There are bound to be cases where the tiform process, dpsl: distal poorly sclerotized lobe, ht:
keys do not appear to work. In these instances where hairy tubercle, ppsl: proximal poorly sclerotized lobe,
a particular character appears to be at fault or is ambi- sl: serrate lamella.
guous to interpret, take the alternative of a couplet to Pc: praecoxa
see if this brings you to a credible identification. T he Pes: praecoxal sclerite (not figured)
size of species (Table I) is often a useful additional Pd i-5: pedigerous somites 1-5
Copepoda 891
!"'isions Pr: prosome (the anterior region of the body limited Family Miraciidae Dana, 1846 (p. 967)
by a major articulation) Family Euterpinidae Brian, 192 1 (p. 968)
Sc: sy ncoxa (fused praecoxa and coxa) Family Clytemnestridae A. Scott, 1909 (p. 968)
~Fig. Url-5: urosomites 1-5, last somite being the anal Family Ae~:isthidae Giesbrecht, 1892 (p. 968)
somite Order Poecilostomatoida Thorell, 1859 (p. 969)
ven .: ventral v iew Family Oncaeidae Giesbrecht, 1892 (p. 969)
Family Sapphirinidae Thorell, 1859 (p. 971)
Outline classification Family Corycaeidae Dana, 1852 (p. 973)
Order Siphonostomatoida Thorell, 1859
octudes Taxa covered in the present chapter are denoted with Order Monstrilloida Sars, l ~03
bold underlined characters.
Keys and diagnoses of the taxa
-~ Subclass Copepoda M ilne-Edwards, 1840
Order Platycopioida Fosshagen, 1985 Subclass Copepoda
Order Calanoida Sars, 1903 (p. 901) Copepods at some stage in their life-cycle possess at
Family Megacalanidae Sewell, 1947 (p. 905) least 2 pairs of swimming legs with rami connected by
Family Calanidae Dana, 1849 (p. 906) an intercoxal scle rite; aCe into which the Mxp bearing
Family Paracalanidae Giesbrecht, 1892 (p. 908) somite is incorporated; uniramous A 1 with up to 27
Family Mecvnoceridae Andronov, 1973 (p. 9 11 ) segments, although in many copepods (harpacticoids)
Family E ucalanidae Giesbrecht, 1892 (p. 9 11 ) the segm entation of A 1 is secondarily reduced; pos-
Family Spinocalanidae Vervoort, 195 1 (p. 9 13) sess egg-sacs, although some important groups lack
Family Clausocalanidae Giesbrecht, 1892 (p. 914) true egg-sacs and they are secondarily lost in some
Family Aetideidae Giesbrecht, 1892 (p. 9 17) highly derived parasitic forms (Huys eta!., 1996).
L::rernal Family E uchaetidae Giesbrecht, 1892 (p. 923)
Family Phaennidae Sars, 1902 (p. 927) Key to orders of Copepoda (adapted fro m Huys et
Family Scolecitrichidae Giesbrecht, 1892 (p. 930) a!., 1996)
Family Diaixidae Sars. 1902 (p. 933) I Body showing gymnoplean tagmosis, with Pr-
Family Tbarvbidae Sars. 1902 (p. 934) Ur boundary located between the Pd5 and Gns
family \lesaiokeratidae Matthews. 1961 (p. 935) (Fig. 6.1 A): C alanoida (p. 901)
f....,·~ Arietellidae Sars. 1902 (p. 936) Ia Body showing podoplean tagmosis. w ith Pr-Ur
F~_ .-.. ~aaptilidae Sars. 1905 (p. 937) boundary located between Pd 4 and 5 (Fig. 6 .1 B)
f _ Hererorhabdidae Sars. 1902 (p. 9-l2) ...........................................................................2
l w ic miid.ae Sars. 1902 lp. 9-C l ~ :\1 Exp usually \\;th 2 or more segments (Fig .
~en _ Ico:il?::rid:!e 5.!rs.. l9C~ p. o_:- 6.1 C . rarely }-segmented or absent, in which
IP seg- ;af5rl~ B:OCsl.~·- : o~: p. 9-q case baseoendopod of P5 \\'ith 2 or more setae
Roman Fig. 6.1D ): Harpacticoida (p. 967)
ibr the 2a .-\.2 Exp ! -segmented or absent (Fig. 6.1 E), P5
19-l9) ...!...."-""'""~~ (;:~~.:.=:_ :~: (~ .. l..i~~ baseoendopod represented by at most 1 seta
!:::::d are Farrul~ Can~e G 5..~.:~ :.::c.:: ~ '-'5- !Fig. 6.1F. H) ............... .................. .. ................ .. 3
ts.. The Family Pontellidae ::>:ma.. :.::5;. _p. o:;- 3 ~lale AI typica lly geni c ulate (Fig. 6.11-1).
ooents, Family Acartiidae Sa.rs. i~;;3 UJ. 961 Female A I with up to 26-segmented. Mxp usu-
le':". ter- Family Tortanidae Sars. 190: lp. 9631 ally similar in both sexes. Female Gns with sing-
:e Fig. Family Bathpontiidae BrodsJ...;·. 1950 lp. 963) le midventral copulatory pore (except in
p: digi- Order \1isophrioida Gurney. 1933 Oithonidae): Cyclopoida (p. 964)
OOe.ht: Order \1ormonilloida Boxshall. 1979 3a Male A l non-geniculate. Female AI a t most 7-
:C lobe, Order Gelyelloida Huys, 1988 segmented (Fig. 6.11). Mxp always sexually
Order Cvclopoida BUtme ister, 1834 (p. 964) dimorphic (Fig . 6.1 G, J). Female Gns with copu-
Family Oithonidae Dana, 1853 (p. 964) latory pores located within dorso-lateral genita l
Order Harpacticoida Sars, 1903 (p. 967) apertures (Fig. 6.1F) (except in Erebonasteridae):
Family E ctinosomatidae Sars, 1903 (p. 967) Poecilostomatoida (p. 969)
892 Copepoda
Microsetella atlantica
gymnop lean podoplean

tagmosis tagmosis
Microsetella
rosea
0
/
Oncaea scottodicarloi
OMxp l!
Oithona simi/is
Fig. 6.1.
Copepoda 893
l
=np
Centrf!pages Euaugaptilus
hptcus hulsem anae
Pleuromamm~
abdom ina Its
anterior ~od y,
latera l view
Tortanu S angularis
_ -{
amerior~\ ,
lateral 're\\
. tom us
Pseudo d tafichardi
0 I
\~\
\i\
0 J1l
•\1
]L o ~-.
r.r}i
.
rt:r-·1
~~
~ r .
P3
<(PS
Phyllopu_s
I I·
Calanus aequalts Candacia

australis varicans
Megacalanus
princeps Temora
A cartia clausi discaudata
Fig.6.2.
894 Copepoda
0 0
9
lateral view
a l
Lu c ic utia
fla vicornis
A rietellus
9 setosus
H a loptilu s
Rhincalanus oxycephalus
nasutus
0
I
Tharybis
tt asymmetrica
do rsa l view vipera
Mecynocera I
clausi
\
....
(..-
\
Ph yllopus P1 P2
t
<;?PS-
N eoscolecithrix aequalis
caetano vi Lucicutia Paracalanus Ca lanus
/ongiserrata indicus australis
Fig. 6.3.
Copepoda 895
~
-:::-
J Mxp2
Mx2
Sco lecithrix
Clausocalanus
arcuicornis
Onchoca/anus danae
cristatus
ucia
Y'lis
~
~ Spinocalanus
longicornis
---
iemale anterior body
lateral view Chiridiella atlantica J
L
Paraeuchaeta biloba
- - - - - - - - -- -- - -- ---- - - -- -
genital
prominence
0 <(Gns 0 <(Gns
or swelling
•otecio' lobe>
of genital --.,.;.
flange "y,.
P2
posterior lobe /
of genital
flange Scolecithrix
danae
schematic lateral view of schematic ventral v iew of

female euch aetid gen ital somite female euchaetid genital somite
Fig. 6.4.
896 Copepoda
0 0 ~
CPS CPS
Heterorhabdus lobatus Haloptilus ornatus
Chirundina
~
streetsii
0
1/) r ~
CPS
CPS
Pseudodiaptomus richardi Phyllopus helgae
Lucicutia aurita
y
CPS CPS
Arietellus plumifer Centropages typicus
Ca ndacia cheirura
Fig.6.5.
Copepoda 897
O PS
Acartia tonsa
OPS O PS
OPS
Tortanus (Tortanus) Temora turbinata
gracilis
Labidocera acutifrons
OPS
OPS OPS
Bathycalanus Tharybis
Oiaixis m ega lodactyla
eltaninae gambiensis
Temoropia
mayumbaensis
0 O r.fl
~ OPS
~- \ Cornucalanus
che lifer
our
Subeucalanus longiceps OPS
Amallothrix
r1g. 6.6. gracilis
898 Copepoda
Mesaiokeras tantillus
OPS
0
Megacalanus OPS
pnnceps
Pseudochirella obtusa
OPS
Calanus
australis
Undinula vulga ris
O PS
I
Cosmocalanus
OPS darwinii
OPS
Clausocalanus Aetideus
bre vipes armatus
, distal, poorly
;:~ sclerotized lobe
(dpsl)
P4
Exp
OPS
Spinocalanus OPS schematic diagram
of male swimmmg leg
longicornis Aetideopsis left exopod
rostrata
Fig.6.7.
Copepoda 899
OPS
Mecynocera clausi
OPS
Calocalanus pavo
Oncaea media
0
0
<(A2
OPS
Corycaeus
(Corycaeus) clausi
Paracalanus indicus
F~g. 6.8.
900 Copepoda
Sapphirina nigromawlata
Copilia hendorffi
Fig. 6.9.
Fig. 6. illustrations of characters used in the key to orders and the families Calanoida and Poecilostomatoida. Figure sources: from Alvarez
( 1985a): 6.3J: from Andronov ( 1973a): 6.4 D, 6.7C; from Andronov ( 1976): 6.3 1, 6.6H; from Andronov (1979): 6.6G; from Bj!irnberg (1968): 6.6F;
from Bradford ( 1971 a): 6.58, 6. 7G; from Bradford ct al. ( 1983): 6.4A, B, K, 6.6L, M, 6.71; from Bradford-Grieve ( 1994): 6.2K, L, 6.3A, B, C, M,
6.3K, 6.61 , J, K. 6. 7A. B. F, H, K, 6.88, D, G; from Brady (1883): 6.6B; from Corral ( l972b): 6.8A; from Dahl ( 19 12): 6.8E, f; from Dakin and
Colefax ( 1940): 6. 7E; from Damkaer (1975): 6.4G; from Deevey ( 1972): 6.6E; from Farran ( 1929): 6.5G; from Frost and Flemingcr ( 1968): 6.4C;
from Giesbrecht (1892): 6.9C. D, 6.1 C, D, H , 6.2C, D, E , I, N, 6.3E, F, H, K, 6.5C, 6.6A, C, D ; from Heron and Bradfo rd-Grieve ( 1995): 6.1 E, F, G, T,
J, 6.8C, D; from Hiilsemann (1966): 6.50 ; from Huys and Boxshall (1991): 6. 1A, B: from Lehnhofer (1926): 6.9E, F, G; from Lchnhofer ( 1929):
6 .9A, B; from Markhaseva (1996): 6.7D, L; from Matthews 1972: 6.2B; from Ohtsuka ( 1992): 6.2F; from Ohtsuka et al. (1994): 6.5H ; from Rose,
1933: 6.2M; from Sars ( 1903): 6.2A, G, J, 6.4H, 6.5F, I; from Sars ( 1924): 6.3 D, G; from Sars, (1924-25): 6.71; from Tanaka ( 1957b) : 6.5A; from
Vcrvoort ( 1957), by courtesy of the Mawson Antarctic Collection, University of Adelaide, Australia: 6.4E; from Wolfenden ( 191 1): 6.4F; from
Wright ( 1936): 6.2H, 6.5E; from Original: 6.41, 1. 6.7M;
Copepoda 901
Order Calanoida Sars, 1903 4 P5 Exp2 with finger-like projection instead of

The Calanoida are defined by the combination of the internal seta (Fig. 6.2A). Posterior corners of Pr
gymnoplean tagmosis, the presence of only 1 spine on often prolonged into points. Oral parts not redu-
the outer margin of P2-P5 Expl , the presence of a ced. Distal setae of Mxp without small shield-
-_) coxal epipodite (Le i), but not a lobate basal exite on
~x 1, and the presence of a seta on the inner margin of 4a
shaped appendages: Centropagidae (p. 950)
P5 Exp2 segment without finger-like projection
A2 C. The presence of a maximum of 2 setae on the internally (Fig. 6.2B). Posterior comers of Pr
j terminal segment of Mx2 (Enp4) is an apomorphy
(derived character) of the Calanoida (Huys and
usually rounded. Oral parts strongly reduced to
different degrees. Distal setae of Mxp strong,
Boxshall, 1991 ). curved and often with small shield-shaped
appendages: Augaptilidae (Augaptilus, Centrau-
Order Harpacticoida Sars, 1903 gaptilus, Euaugaptilus) (p. 937)
The Harpacticoida are defmed by their podoplean tag- 5 Inner margin of P2 Enpl with excavation which
mosis, the presence of short A 1 with at most 9 seg- has one or two hooks on its inner margin (Fig.
ments in females and 14 in males, the fusion of the 6.2C). R with two filaments. Pr in Pleuromamma
Enp and B into baseoendopod on P5 of both sexes of with small black rounded organ on one side of the
most species, those with a separate Enp having a body. Segmentation ofP 1-P4: 3/3: Metridinidae
maxi mum of 2 setae on it (Huys et al., 1996). (p. 947)
5a Inner margin of P2 Enpl without excavation or
Order Cyclopoida Bunneister, 1834 hook. Pl-P4 with various segmentation which
The Cyclopoida are defined by their podoplean tagmo- often differ from 3/3 ....... ..................... .... ... ....... 6
sis, by the possession of a combination of digeniculate 6 Segmentation of P1-P4: 311 -3, 3/2, 3/2, 3/2 ...... 7
A1 bearing a sheath on segment 15 in males with the 6a Segmentation ofP1-P4: 1-311 -2, 3/2-3, 3/3, 3/3
lack of a defined A2 Exp. The A2 Exp is represented by .. ..... ................. ..... .... ................. ....................... ]]
up to 3 setae, but no Exp segment remains. They are 7 Anterior part of head with (sometimes without)
also characterized by the fusion of Enp 1 and Enp2 of one or two pairs of cuticular lenses, often lens is
~ and by a 4-segmented Md Exp derived by loss of present in base of bifurcate R (Fig. 6.2D, E):
the ancestral Expl (Huys and Boxshall, 199 1) . Pontellidae (p. 957)
7a Anterior pmt of head without cuticular lenses. R
Order Poecilostomatoida Thorell, 1859 not bifurcate; as 2 filaments or absent .. ..... ....... 8
The Poecilostomatoida are defined by their podoplean 8 Anteri or part of head with single pigmented eye
tagmosis, by their strongly sexually dimorphic Mxp .. ...... ..... ....... ....... .. .... ..... .... .. .. .... ... ...................... 9
and by the structure of bilobate Mx l (Huys and Sa Anterior part of head without single pigmented
Boxshall, 199 1). eye .................................................. ........... ... ... . lO
9 Anterior to upper labrum a rounded, densely hai-
red plate present (Fig. 6.2F). R absent. P2-P4
Order CALANOIDA Exp with external sp ines: Tortanidae (p. 963)
9a Densely haired plate anterior to labmm absent
Key to the families of Calanoida (modified from (Fig. 6.2G). R present. P2-P4 Exp without extern-
Brodsky eta!. , 1983) al spines, only unarticulated spinifo rm protrusion
_-\b.=z Females of segment present (Fig. 6.21): Acartiidae (p. 961)
o.oF: l Ur with no more than 3 free somites ................. 2 I0 Anterior part of head usually rectangular (Fig.
3.. C .\l. l a Ur with 4 free somites ........... .... .................... .. 13 6.2I). Mx2 enlarged, with strong distal setae.
:3::: .!:ld
oAC:
2 P5 present ......... ........... .............. ....... ........... .. .. ..3 Mxp strongly reduced. Outer margin of P2-P4
:..?..G.L 2 P5 absent: E ucalanidae (Eucalanus, Subeu- Exp segments finely serrated. Some species with
- i0291: calanus, Pareucalanus) (p. 911) body partly coloured by dark pigm ent.
s:::3.o:;e_
3 P5 similar in structure to PI-P4. Segmentation of Swimming legs usually heavily pigmented black
'·"-=~
:;:- :"::om p l-P4: 3/3, 3/3,3/3,3/3 ....... .................. ............. .4 or dark-brown: Candaciidae (p. 954)
3a P5 very different in structure from P1-P4. P I-P4 I Oa Anterior part of head rounded (Fig. 6.2N). Mx2
segmentation may be different from 3/3 ...... .... .5 not enlarged, Mxp not reduced. Outer margins of
902 Copepoda
P2-P4 Exp segments not serrated. Species not l9a Anterior surface of P l B adjacent to Enp without
dark-pigmented: Temoridae (Temora, Temoro- cylindrical projection. CR only 3-5 times longer
p ia mayumbaensis deviates in having UR of 4 than wide; usual ly asymmetrical, their distal ends
free somites) (p. 953) cut obliquely (Fig 6.3H): Heterorhabdidae
11 Segmentation of P1 : 2-3/1-2. P2: 3/3 .... ........ .. 12 (p. 942)
ll a Segm entation of Pl: 1-3/ 1-2, P2: 3/2-3 : Para- 20 P I Enp l-2-segmented, PI Expl lacking outer
calanidae (Calocalanus, Parvoca/anus) (p. 908) spine, P3 B with outer spine: Bathypontiidae
12 A l twice as long as body. Sperrnathecae nearly (p. 963)
spherical (Fig. 6.3A; 6.3B). Segmentation ofP I: 20a PI Enp 3-segmented; P 1 Exp 1 with outer spine.
3/ 1. Posterior margins of Pd2-4 and Gns w ithout P3 B withou t outer spine ............ ... .......... .. .. .... 2 1
spines: Mecynocerida e (Mecynocera clausi 2 1 Copepods usually with characteristic eye com-
Thompson, 1888) (Fig. 7.53) (p. 9 11) posed of pigmented spot surrounded by refracti-
12a AI not more than 1.5 ti mes longer than body. le lens. Mx l Lel with 10 setae. Mx2 Li 1-Li2
Sperrnathecae not spherical. Segmentation of with 4 and 3 setae respecti vely (see also Fig.
PI: 2/2. Posterior margins som e or all Pd and 6.2H): Pseudodiaptomidae (p. 952)
Gns with two dorsal spines each and spine-like 2 1a Copepods usually without eye. Mx l Lel with 5-
projections laterally (Fig. 6.3C): .E ucalanidae 9 setae or without setae. Mx2 Li 1 with 1-3 setae,
(Rhinca/anus) (p. 9 11) L i2 with 1-2 setae .. ............... .. ........ .. .... .... .......22
13 P 5 present ............... ............... ................. ......... 14 22 Ce rounded or pointed at apex (Fig. 6.30 ) . Mdp
l3a P5 absent... ............................. .. .......... ..... .. ....... 29 w ith rudimentary Enp or without Enp. P5 3-seg-
14 Segmentation ofPl -P4: 3/3; rarely P 1 Exp or P l mented penultimate segment w ith inner projec-
Enp 2-segmented .... .... .. .......................... ........ . 15 tion bearing 1-3 setae (rarely P5 1- or 2-segmen-
14a P l Enp l-3-segmented, P2 Enp 2-3-segmented, ted), P5 may be biramous. Setae of oral parts and
segmentation o fP3 -P4: 3/3, 3/3 ......... .... .... .. .. .23 CR long, hairy and brightly coloured:
15 P5 similar in structure to P l-P4 (Fig. 6.3M) ... 16 A rieteUidae (p. 936)
15a P5 strongly differing in structure from P 1-P4 22a Ce rounded (Fig. 6.3G). Mdp with well devel-
(Fig. 6.2H, M) .. .. ................ .......... ... ..... ....... .... .20 oped Enp. P5 5-segmented, un iramous, w it h dis-
16 Mx 1 Le2 present, with a single seta ................ l 7 tolateral teeth (Fig. 6.2M): Phyllopodidae (p.
16a Mx I lacking Le2 with seta .............................. 18 949)
17 P2-P4 Exp3 with 3 outer spines (Fig. 6.2L). P5 23 Segmentati on of P l-P2: 3/2, 3/3. Terminal spi-
C always smooth on inner mar gin. Large-sized nes of Exp of P2-P4 smooth (Fig. 6.3L):
copepods (8-1 7 mrn): Megacalanidae (p. 905) P aracalanidae (De/ius, Paracalanus) (p. 908)
17a P2-P4 Exp3 with 2 outer spines (Fig. 6.2K). PS C 23a P I Enp ]-segmented. P2 Enp 2-segmented .... 24
often serrated on ilU1er margin (Fig. 6.3M). Body 24 Mx2 with setae mod ified into sensory appenda-
length less than 10 mm: C alanidae (p. 906) ges (Fig. 6.4A, B) ........................ .. .. .. .............. 26
18 CR long, usually 3-5 times longer than wide, if 24a Mx2 without sensory setae .. .............. ..... ......... 25
shorter then asymmetrical .. ................ ............. 19 25 R of two thin long fi laments or two-pointed (Fig.
18a CR short, symmetrical. Md with elongated, nar- 6.4C): Clausocalanidae (Clausocalanus, Cteno-
row gnathobase. Oral parts strongly reduced to calanus) (p. 914)
different degrees. Dista l setae of Mxp powerful, 25a R, if present, one-pointed, thin and ha rd ly visi ble
strongly curved. Usually very transparent cope- (Fig. 6.4D): Mesaiokeratidae (p. 935)
pods. Sometimes frontal spine p resent (Fig. 26 Mx2 w ith L worm-like and 7 brush-li ke sens01y
6.3E): A ugaptilidae (Haloptilus) (p. 937) appendages distally (Fig. 6.4A). Mx2 Li l usual-
19 Anterior surface of P l B adj acent to Enp with ly with 5, rarely with 4 setae: P haenni dae
short cylindrical projection with seta (Fig. 6.3K). (Cornucalanus, Onchocalanus, Brachycalanus,
CR long, 3-10 times longer than wide, symme- Xanthocalanus) (p. 927)
trical or asymmetrical, their distal ends with 26a Mx2 usually with 3 (rarely 4) worm- like and 5
parallel borders (Fig. 6.3F): L ucicutiidae (p. 945) (rarely 4) b rush-like sensory appendages distally
(Fig. 6.4B). Mx2 L il with 3-4 setae (5 setae in
Landrumius)........ ................. ....... .. .... .. ......... ....27
Copepoda 903
~::bout 27 Posterior corners of Pr extended into 2 points 35a Posterior corners of Pr often prolonged into
kmger (Fig. 6.3J): Phaennidae (?) (Neoscolecithrix) (p. points, lobes if rounded, usually with spines. R 1-
:!lends 927) pointed, 2-pointed without filaments or, rarely
bdidae 27a Posterior corners ofPr not extended into 2 points absent. Ce sometimes with frontal spine
OOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOO OOOO OOOOOOooooooO OOOOOOOOO oo•o28 (Gaetanus); 111 some species of Euchirella,
! outer 28 P l -P4 surfaces strongly spinulose (Fig. 6.4K). Chirundina, Undeuchaeta with crest (Fig. 6.SA):
niidae Mxl Li1 not extendi ng beyond Enp. Mx2 LiS Aetideidae (p. 917)
usually with sensory appendage (Fig. 6.4B): 36 Mx2 with setae on Lil -Li5 strongly transformed
i spine. Scolecitrichidae (p. 930) (Fig. 6.4F). Segmentation ofP I-P4: 111, 2-3/ 1-2,
- ...21 28a Pl -P4 surfaces weakly spinulose. Mx 1 Lil com- 3/1 -2, 3/ 1-3. R absent: Aetideidae (Chiridiella)
e comparatively large, extending or reaching beyond (p. 917)
efracti- Enp. Mx2 LiS without sensory appendage (see 36a Mx2 without strongly transformed setae on L i ]-
j f -Li2 also Fig. 6.31): Tharybidae (p. 934) LiS. Segmentation of PI-P4:2-3/l , 2-3/1-2, 2-
5<l Fig. 29 Segmentation ofPl-P4: 2-311, 3/1, 3/3,3/3 or 2- 3/2-3, 2-3/2-3. R two-pointed: E uchaetidae
3/ 1' 3/2, 3/3, 3/3 00000000000000000000 00 00000000000000000000000030 (Valdiviella) (p. 923)
lrirh 5- 29a Segmentation of P l-P4: 1/1, 2-3/J -2, 3/1 -2, 3/ 1-
: 5e"'.ae. 3 or 2-311, 2-3/ 1-2, 2-3/2-3, 2-3/2-3 000000000000000036 Males
")'') 30 Segmentation of P 1-P4: 2-3/1 , 311, 3/3, 3/3. 1 A1 on one side geniculate (Fig. 6.4H) 000000000000002
.\ldp Supralabrum (anterior to labrum) is highly pro- Ia Neither A l geniculate .oo.oo ooooooo .. ............. oo .. .... .. l6
.3-sa!:- minent stmcture. Mxp strong with long, strong- 2 Inner margin of P2 Enpl without excavation or
_iec- ly built, slightly curved setae. Many species with hooks ...... oo ........................... ..... oo·······oo··············-3
!'~en
asymmetrical genital field (Fig. 6.4E, T, J): 2a Inner margin of P2 Enpl with excavation and
r.s :md E uchaetidae (p. 923) hook, or pair of hooks (Fig. 6.2C): Metridinidae
lol!:'ed: 30a Segmentation of P I-P4: 2-3/ 1, 3/2, 3/3, 3/3, (p. 947)
(sometimes P2: 3/1 - Euchirella). Supralabrum 3 Left A I geniculated .. ....... oooo .... ..... .. ...... 00 .. ... oo ... A
<!e' el- (anterior to labrum) not highly prominent. Mxp 3a Right Al geniculatedoo ... .. ...... oo .. oo oo ....... oo .. oooo ..... 8
E± dis- moderately developed. Genital field usually 4 P5 of sitnilar segmentation to P l -P4; distal seg-
he !p. symmetrical 00 000 oooooo ·. 00.00 000.0000000 00 .00 00000 00000 000 00.00.3 1 ment modified compared with those of P2-P4
31 Mx2 with terminal setae modified into sensory (Fig. 6.5B, C, D) .. oo .......... oo .. oooo .... ....... oo ...... oooo ... 6
~ spi- worm-like and brush-li ke appendages (Fig. 4a P5 very different from Pl-P4, with 3-segmented
- .3L): 6.4A, B) ooooooooooooooo oo oooooooooooo oooooooooooooooooooooooooooo .. 32 Exp and strongly reduced Enp, sometimes Enp
) 31a Mx2 without tenninal sensory appendages 00 00.34 of one leg absent (Fig. 6.5F, H) ........ .. ....... .... ... S
c _ ~ -t 32 Mx2 with 1 worm-like and 7 brush-like appen- S Right P5 without Enp. In both P5, B with long
.penda- dages (Fig. 6.4A). R 2-pointed: Phaennidae outer seta (Fig. 6.5F): PhyUopodidae (p. 949)
---~6
(Phaenna) (p. 927) 5a Right P5 with Enp. Outer seta of right P5 B usu-
- ~5
32a Mx2 with more than I worm-like appendage. R ally longer than left one; latter may be absent (Fig.
!C Fig. one-pointed, or with 2 fi laments 00000000000000000000033 6.5H): Arietellidae (Arietellus, Paraugaptilus ; in
C2no- 33 Small copepods, l.ess than 1 mm in total length: Paramisophria right P5 Enp absent) (p. 936)
Diaixidae (p. 933) 6 Anterior surface ofP I B adjacent to Enp without
,isJble 33a Copepods more than I mm 111 total length: sh01t cylindrical protrusion, but seta present .. 007
Scolecitrichidae (Scolecithrix, Scaphocalanus 6a Anterior surface of P I B, adjacent to Enp with
;e:::.sory curtus) (p. 930) short cylindrical protrusion bearing seta (Fig.
.3Ual- 3-l P3-P4 Exp3 with 4 inner setae. Surfaces of P 6.3K; see also Fig. 6.5D): L ucicutiidae (p. 945)
11Jlidae usually smoothoo .oo ooooOOOOoo oooooo oooo ooOOoooo.oo ooo ooOOoooooo3S 7 Both P5 usually 3-segmented but differ from P 1-
3-la P3-P4 Exp3 with 5 inner setae. Anterior and usu- 4 as Exp modified and B has inner projection
ally posterior surfaces of P usually spinulose (Fig. 6.SB): Heterorhabdidae (p. 942)
~ :md 5 (Fig. 6.4G): Spinocalanidae (p. 913) 7a Both P5 2-3-segmented, nearly like P4, but
E...~y 35 Posterior corners of Pr always rounded. R of 2 differs as distal Exp setae short, or transforn1ed
~in filame nts: Clausocala nidae (Microcalanus , into spines (Fig. 6.5C): Au gaptilid ae (in
Pseudocalanus) (p. 914) Euaugaptilus (part.) right A I geniculate) (p. 937)
904 Copepoda
8 One ofP5 legs c helate (Fig. 6.5E, G, I, 6.6A, B, 17 P5 similar in structure to Pl -P4 (Fig. 6.7 A, B)
D) .. ............ ....... ............. .. ....... ... ............... ....... ...9 ..... .... ... .......... ...... ............. ........... .......... ..... ...... 18
8a Neither P5 chelate .. ...... .. ... .......... ... ...... ........... 15 17a P5 strong ly differing from Pl -P4 ..... .. ... .......... 19
9 P2-P4 Enp 2-segmented, Exp 3-segmented .... 11 18 P2-P4 Exp3 with 2 outer spines (Fig. 6 .2K) :
9a P2-P4 Enp and Exp 3-segmented ..... ........ ...... . I 0 Calanidae (Calanus, Canthocalanus, Mesocala-
10 P3-4 Exp with 3 outer spines: Centropagidae nus, Neocalanus, Nannocalanus) (p. 906)
(p. 950) 18a P2-P4 Exp3 with 3 outer spines (Fig. 6.2L):
I Oa P3-4 Exp with 2 outer spines: Pseudodiaptomi- Megacalanidae (Megacalanus) (p. 905)
dae (p. 952) 19 Pl -P4 segmentation: 2-3/ 1, 3/2, 3/3,3/3 ......... .20
11 Cu ticular lenses or pigmented eye present on 19a P 1-P4 segmentation different .. .. .... ........... ....... 27
anterior part of Ce ... ........................................ 12 20 Distal part of Mx2 with sensory appendages
II a Cuticular lenses and p igmented eye absent on (Fig . 6.4A, B) .... .................. ................. ... ........ 21
anterior part of Ce .. .... ............ .... ......... ..... ..... ..14 20a Distal part of Mx2 without sensory appendages
12 Anterior part of Ce sometimes with 1 or 2 pairs .............. ...... ............. .............................. .......... 24
of cuticular lenses, often lens present near base 21 P l -P4 surface of segments usually highl y spinu-
of strong R (Fig. 6.2E): Pontellidae (p. 957) lose .. .... ................... ........... .... .... ....................... 22
12a Anterior part ofCe always lacks cuticular len ses, 2la P l -P4 surface of segment p oorly spinulose, or
single p igmented eye present. R is not strong or smooth .. .......... .................... ...... ....... ..... ...........23
absent ... .. ............. ......... ............. ....................... l 3 22 P 5 usually biramous on both sides, asymmetri-
13 Anteriorly rounded hairy p late present, anterior cal; C and B narrow and elongated on left, sh ort
to upper labrum (Fig . 6 .2F). P 5 uniramous, mas- o n right, w ith B usually very swollen (Fig.
sive, asymmetrical; 3-segmented on right w ith 6.6M) : Scolecitrichidae (p. 930)
large c hela, 4-segmented on left, curved (Fig. 22a P5 usu ally uniramous, if Enp present they a re
6.6B): Tortanidae (p. 963) rudimentary; asymmetrical, longer on left (Fig.
13a Rounded hairy p late, ante rior to upper labrum, 6.6L): Phaennidae (p. 927)
absent (F ig. 6.2G). P5 uniramous, right 4-seg- 23 Left P5 with long Enp, right usually unira mous
mented, distal segment of right, longer leg, c ur- (not in Parundinella) (Fig. 6.6H): Tharybidae
ved, left 3-4-segmented (Fig. 6.6D) : Acartiidae (Tharybis megalodacty la Andronov, 1976) (p.
(p. 96 1) 934)
14 Posterior corners of Pr always asynm1etrical. 23a Both P5 uniramous (Fig. 6.6G): Diaixidae (p. 933)
Gns is almost always asymmetrical, bearing pro- 24 P2-P4 Exp3 with 4 se tae ...... ............... ............ 25
cess on right side: Candaciidae (p. 954) 24a P2-P4 Exp3 w ith 5 setae (Fig . 6.7K): Spino-
l4a Posterior corne rs of Pr and Gns usually synmle- calanidae (p. 913)
ttical. Gns usually w ithout process on right side: 25 R !-pointed, 2-pointed, or absent, never wi th
Temoridae (in Temoropia P2-4 Enp m ay be fi laments .. .. .. ...... ..... .................... .................... .26
incompletely 3-segmented, Fig . 6.6C, E) (p. 85) 25a R of 2 filaments or rudimentary, rarely absent.
15 P5 d iffers in structure from P l-P4. Med ium P 5 elongated, uniramous, very asymmetrical
sized and s ma ll calanoids ( 1.0-5.6 mm): (right leg sometimes rudi mentary) (Fig. 6.7F):
Bathypontiidae (p. 963) Clausocalanidae (p. 914)
15a P5 close in structure to Pl-4 (Fig. 6.6F). Large 26 P5 bira mous with Enp variously developed, or
calanoids (8- 10 mm): Megacalanidae (Bathy - uniramous, or w ith only one leg present.
calanus) (p. 905) Calanoids usually longer than Inm1 (Fig. 6.7D,
16 Ur of 5 free somites. Anterior part of head w it- G , L): Aetideidae (p. 9 17)
hout elongated prominence a nterior to A 1 (Fig. 26a P5 always uniramous, asymmetrical: 3-segmen-
6.61) .... .... ..... ....... .......... ...... ................... .. .... ... .. 17 ted on right, short, nearly as long as first segment
I 6a Ur of 4 free somites (anal somite fused to CR) of 5-segmented left leg. Calanoids usually less
(Fig. 6.6J, K) . Anterior part of head triangular, than 1 mm (Fig. 6.7C): Mesaiokeratidae (p. 935)
sometimes with elongated promine nce anterior 27 P l -P4 segmentation: 2-3/1 , 311 , 3/3, 3/3. P5
to A l : Eucalanidae (p. 9 11) powerful, nearly as long as Ur; asymmetrica l.
Exp 3-segmented on left, Exp2 with (or without
Copepoda 905
-A B) - Valdiviella) serrated lamella, sometimes its tip 3, 4 and 4 setae respectively (except in Bathycalanus).
____.18
prolonged into thin long seta (Fig. 6.71, J, M): P I Exp3 w ith 1 external spine in Bathycalanus richar-
_ ____19
Euchaetidae (p. 923) di. P2-P3 B with articulated external spine. Spine and
6-lK): 27a P1-P4 segmentation: 3/1-3, 3/3, 3/3, 3/3 .........28 seta formula:
ocala- 28 PI Enp 3-segmented (see also Fig. 6.7E, H):
Calanidae (Cosmocalanus, Undinula) (p. 906) coxa basis exopod segments endopod segments
6.2L) : 28a P1 Enp 1-2 segmented ..... .... .... ... ...... ..... .......... 29
Leg l 0- l 0-l 0/I- l ; 011-l; JJII,I,4 0- J; 0-2; I ,2,3
29 Both P5 5-segmented of nearly same length (Fig. Leg2 0-1 1-0 1-l; 1- 1; 111,1,5 0-1; 0-2; 2,2,4
_ ____ 20
6.8A). P1 Enp always !-segmented. A2 not Leg 3 0-1 I-0 l-1 ; I-1 ; III,I,5 0- l; 0-2; 2,2,4
- --··27 described: M ecynocer id ae (Mecynocera clausi Leg4 0- 1 1-0 T-1; T-1; TTT,T,5 0- 1; 0-2; 2,2,4
~ges Thompson, 1888) (Fig. 7.53) (p. 9 11) Leg 5 0-0 011-0 1-0; l-1; II,l,4 0-1; 0- 1; 2, 1,3
----··21 29a If both P5 present, only one 5-segm ented;
:xlages strongly asymmetrical in length (Fig. 6.8B, G). P5 similar to P2-4.
_ ....24 P 1 Enp 2 -segmented, rarely ! -segmented
spmu- (Paracalanidae: Calocalanus minutus, C. nami- Males
_ ____22
biensis. C. renuiculus. Parvocalanus scotti). A2 Ur of 5 somites. A l 25-segmented, segments 7-9 may
)Si'. or terminal segment very short \\itbout 3 terminal be fused. Right A l is geniculated in Bathycalanus.
_ ... .23 setae. (Fig. 6.8H): Paracalanidae (p. 908) A2-Mx2 similar to that of female. P l -P4 as in female.
::netri- P5 with 3-segmented rami similar to female P5; Enp l
~ short Fz:::;:Uly 2\lega.cabn.idae Sc<relL l ~ - \\i thout inner setae; Exp3 with different armam ent,
: Fig. R;:;_..::o.:~ ~C:!i:C.-G:ie- e (199-!: Brodsk~· er al. also left Exp2 with inner distal projection, or seta.
I -~
~- are Fem:Ues The family includes 3 genera: Megacalanus Wolf-

r F ig. C:;:- ~c Pc: :::::::...~ hoe ~-"Cor sepanue. Pd-l and 5 sepa- enden. 190-t: Bathyca/anus Sars. 1905 and Brady-
:-are. L r of ~ somires. R \\itb 2 thick sausage-like calanus A. Scott. 1909: all are found in the South
2IDOUS appendages. or 2 sharp slightly tapering spines. or Atlantic.
rbidae filaments which narrow terminally. Al 25-segmented,
"6 (p. 8-9 segments partly fused (here and below only free Key to genera of Megacalanidae
:\1 segments are mentioned). A2 Exp 7-segmented; Females (adapted from Brodsky et a!., 1983)
[). 933) Expl with 1 seta or without setae; Exp2 with 3 with 1 R with 2 strong, sausage-like filaments. P I Exp I
___ __ 25
or without setae. Mdp Expand Enp well developed, B and Exp2 lacking external spines. Mx 1 Li2 wit-
-pino- with 4 setae, Enp l with 4 setae and internal lobe-like hout setae: Bathycalanus
projection (may be absent), Enp2 with 9 terminal and l a R with 2 moderately thin, tapering filaments. P l
f" \\ith 2 posterior setae. Mx 1 Li 1 with 9 marginal spines, 1 Exp l and Exp2 with external spine each. Mx l
- .... 26 anterior and 4 posterior setae (Megacalanus), or 12- Li2 w ith setae ... ... ................ .......... .. ... .... ....... .... 2
ilisent. 13 setal elements (Bathycalanus , Bradycalanus); Li2 2 A l longer than body by tetm inal 7-9 segments.
m-ica! with 4 setae (setae absent in Bathycalanus); Li3 with Mx 1 Enp2 with 4 setae. P l B with hook: Mega-
0. -F): 2 (Bathycalanus, Bradycalanus), or 4 setae (Mega- calanus princeps Wolfenden, 1904 (Fig. 7.1)
calanus); Li4 w ith 2 to 4 setae; Enp l with 3 setae (or 2a A 1 shotier than body. Mx 1 Enp2 with 2 setae.
...00.. or with 1-2 setae in Bathycalanus and Bradycalanus), P l B without hook: Bradycalanus
:-esent. Enp2 with 4 setae, and Enp3 with 7 setae (Enp2 with
6.- D, l and Enp3 with 4-6 setae in Bathy calanus, Males
Bradycalanus); Exp with 11 setae; Le2 with ] seta; 1 Right A l geniculated: Bathycalanus
gmen- Lei w ith 9 setae.Mx2 Lil-Li5 with 5-6, 3, 3, 3, and 3- I a Right A l not geniculated .. ........ ......... .......... ... ...2
~ent 4 setae, respectively; terminal setae may be very long, 2 P 1 Exp 1 lacking outer spine; P I B without hook:
t· less curved at tips looking like ribbon-like lamella Bradycalanus
il- 935) (Bathycalanus); outer seta on coxa present.Mxp C 2a P l Expl with outer spine; Pl B with hook:
3. P5 with 1 proximal seta, then with groups of 2, 4 and 3- M egacalanus p rinceps (Fig. 7.1)
~cal. 4 setae from proximal to distal; Enp2-E np6 w ith 4, 4,
'Tihout
906 Copepoda
Key to species of Bathycalanus B with an external articulated spine, on P4 and P5 this

Females and males may be reduced to a small seta. P2 Exp1 sometimes
I Forehead with pair of chitinous projections. A1 with recurved spine (Neocalanus), or Exp2 with deep
extends beyond CR by about 4-6 segments: invagination (Undinula) . Spine and seta formula:
Bathycalanus richardi Sars. 1905 (Fig. 7.5)
Ia Forehead rounded or with crest.. ............... ...... ..2 coxa basis exopod segments endopod segments
2 Forehead with crest: Batlzycalanus princeps
Leg I 0- 1 0-1 1-1; 1-1 ; 11, 1,4 0- 1; 0-2; I ,2,3
(Brady, 1883) (Fig. 7.4) Leg 2 0-1 1-0 1- 1; 1- 1; 11,1,5 0-1; 0-2; 2,2,4
2a Forehead without crest ............... ............... ........ 3 Leg 3 0-1 T-0 1-1 ; l- 1; ll,1,5 0-1 ; 0-2; 2,2,4
3 Posterior comers of Pr rounded: Bathycalanus Leg4 0-1 1-0 I- I; 1-1 ; ll,J,5 0- 1; 0-2; 2,2,3
eltaninae Bjmnberg, 1968 (Fig. 7.2) Leg 5 0-0 011-0 1-0; 1-1 ; n ,1,4 0-1 ; 0-1 ; 0/112,2,2
3a Posterior comers of Pr triangular: Bathyea/anus
injlatus Bj0mberg, 1968 (Fig. 7.3) (males un- P5 similar to P2-4; C sometimes with inner edge teeth
known) (Calanus , Nannocalanus, Cosmoca/anus).
Key to species of Bradycalanus Males

Femal es (male is known only fo r B. pseudotypicus Ur of 5 somites. A l 25-segmented, with segments 1
enormis Bj0mberg, 1968, and is absent from the South a nd 2 always fused, segments 3-5, 7 and 8, 9 and 10,
Atlantic) and 24 and 25 may also be fused. A2-Mx2 - either
I Posterior com ers of Pr rounded: Bradycalanus exactly as in female, or reduced in size and setation.
gigas Sewell, 1947 (Fig. 7.6) Mxp with reduced inner setae but enlarged outer setae
1a Posterior comers of Pr with d istinct points: on terminal segments. PI-P4 as in female. P5 with both
Bradycalanus typicus Scott, 1909 (Fig. 7.8); rami usually 3-segmented, right leg similar to other P,
Bradycalamts pseudotypicus Bj0mberg, 1968 left leg variously modified; Enp sometimes reduced
(Fig. 7.7) (probably these two species are syno- and devoid of segmentation on one or both sides.
nyms, cf. M ichel, 1994).
There are now 8 genera in the family: Ca lanoides
Family Calanidae Dana, 1849 Brady, 1883; Calanus Leach, 1816; Cosmocalanus
Females Bradford and Jillett, 1974; Canthocalanus A. Scott,
Ce and Pd I may be fused or separate, Pd4 and 5 1909; Mesocalanus Bradford and Jill ett, 1974;
always separate. R of2 fi laments. CR with 4 subequal, Nannoca/anus Sars, 1925; Neocalanus Sars, 1925;
terminal setae and 1 outer seta. Ur of 4 somites. A1 25- and Undinula A. Scott, 1909; all of whic h are found
segmented, generally segments 8-9 partly fused; seg- in the South Atlantic.
ments 23 and 24 with elongate seta. A2 C w ith I seta,
B with 2 setae; A2 Exp 7-segmented; Expl and Exp2 Key to genera of CaJanidae (adapted from Bradford
with 2 setae each, Exp3-6 with I seta each, Exp7 usu- and Jillett, 1974)
ally with 1 seta nearly at midlength and 3 tenninal Females
setae: Enp l with 2 setae, Enp2 with 14- 16 setae. Mdp 1 P5 C inner edge with teeth ...... .... .... .. ....... ......... 2
B with 4 setae; Enp1 with 4 setae and prominent lobe; Ia P5 C inner edge without teeth (naked) ..... ........ .4
Enp2 totally with 10- 11 (8 or 9 tenninal and 2 posteri- 2 P2-3 B with posterior surface spinules: Cosmo-
or) setae. Exp and Enp of approximately equal length. calanus darwinii (Lubbock, 1860) (Fig. 7 . 17)
Mxl Li I with 9 termi nal spines, 4 posterior and 1 ante- 2a P2-3 B without surface spinules (naked) .......... 3
rior setae; Li2 and Li3 with 4 setae each; Li4 with 4 3 Ce and Pd I fused: Nan11ocalanus minor (Claus,
setae, Enp with 14- 15 setae; Exp with II setae; Le1 1863) (Fig. 7. 19)
with 9 setae; Le 2 with 1 seta. Mx2 Lil-Li5 with 4-6, 3a Ce and Pdl partly fused: Calanus
3, 3, 3 and 4 setae, respectively. Terminal part with 4 P2 Exp l with recurved spine (hook) adjacent to
about 10 setae. Mxp C with 1 proximal seta, then with outer spine: Neocalanus
groups of 2, 4 a nd 4 setae from proximal to distal; 4a P2 Expl w ithout recurved (hook) adjacent to
Enp2-6 with 4, 4, 3/4, 4, 4/5 setae respectively. P2-P3 outer spine ............. .............................. ......... ..... 5
Copepoda 907
. P5 this 5 Posterior comers of Pr pointed. P2 Exp2 with Key to species of Calanoides

:rtimes outer proximal edge invaginate: Undinula vulg- Females
~dee p aris (Dana, 1849) (Fig. 7.23) A1 extends beyond CR. Specimens 3.50-5.70 mm
13: 5a Posterior corners of Pr rounded. P2 Exp2 outer in total length: Calanoides acutus (Giesbrecht,
edge without invagination 0000000000 00 0000 0000000000 00 0000.6 1902) (Fig. 7.9)
~ems 6 P1 Enp1 without inner seta. P5 Enp with 7 setae 1a AI shorter. Specimens 2.25-3.90 mm in total
oooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooo ooo0oooooo ooo7 length ................................................... ... ......... ..2
6a P1 Enp1 with 1 inner seta. P5 Enp with 6 setae: 2 Spermatheca consists of 2 parts, proximal part is
-- -.
~ Calanoides massive, in lateral view spermatheca short:
I Mx2 Li1 with 6 setae. P1 without modification: Calanoides macrocarinatus Brodsky, 1972
L.., '>.2 Mesocalanus tenuicornis (Dana, 1849) (Fig. (Fig. 7.11)
7.18) 2a Spermatheca is of different shape, in lateral view
~teeth a Mx2 Li1 with 4 setae. PI C antetior margin ter- spermatheca long: Calanoides carinatus (Kroyer,
minates in well-defined projection; B with inner 1849)(Fig. 7.10)
marginal seta modified into proximally thicke-
ned spine: Canthocalanus pauper Giesbrecht, Males
:xfltS 1 1888 (Fig. 7.16) 1 Right P5 Enp 1 and Enp2 naked oo. 0000000000000000000002
~ 10. 1a Right P5 Enp 1 and Enp2 with 1 inner seta each:
-either \ lales Calanoides acutus (Fig. 7.9)
ei2Iion. I Left P5 highly modified, of complex structure 2 Distal border of right Exp3 extends about 0. 75 of
~setae (prehensile) 00 00 00.00 00 00 .... 00.00 .... 00 00.00 ... 00 00.00 00 00 00.00 00 002 way along left Exp2. Left Enp 2-segmented:
;;:h both Ia Left P5, if modified, then slightly (not prehensile) Calanoides macrocarinatus (Fig. 7.11)
JOher P. OOOOOOOOOOOOOOoOOOOOoOOOoO oooooooooo oooooooooooooO oOoO o•o oooo ooooooo o•oooooooo3 2a Distal border of right Exp3 reaching, or excee-
:reuced 2 P5 C inner edge with teeth. P2 Exp2 without ding distal border of left Exp2. Left Enp !-seg-
s.. invaginated outer proximal border: Cosmo- mented: Calanoides carinatus (Fig. 7.10)
calanus darwinii (Fig. 7. 17)
rwides 2a P5 C inner edge without teeth. P2 Exp2 with Key to species of Calanus
:alanLIS invaginated outer proximal border: Undinula Females
. Scott. vulgaris (Fig. 7.23) 1 Posterior comers ofPr triangular, pointed distally
19- ~: 3 PS C inner edge \Vith teeth ................... ............. 4 000000000000000000000000000000000000000000000000000000ooo0ooooooooooo ooooooooo2
i925: 3a P5 C inner edge without teeth 0000000000000 00000000000 00 .5 1a Posterior comers of Pr rounded: Calanus agul-
~ :OUUd ~ Right P5 Exp3 with inner setae: Nannocalanus hensis De Decker, Kaczmaruk and Marska, 1991
minor (Fig. 7.19) (2.45-2.95 mm) (Fig. 7.1 2); Calanus australis
~a Right P5 Exp3 without inner setae: Calanus Brodsky, 1959 (2.60-3 .59 mm). In practice these
XIord 5 P2 Exp1 with recurved spine (book) adjacent to species are very difficult to distinguish, as pre-
outer spine: Neocalanus viously mentioned (Bradford-Grieve, 1994).
5a P2 Exp1 without recurved spine (hook) adjacent 2 Total length 4.75-6.00 mm. Seta of A1 segment
_ .... 2 to outer spine 0000 0000000000 00 0000 0000000000 00 000000 000000000000 00.6 23 longer than last 8 segments: Calanus propin-
__ ,,,4 6 Both P5 Enp with 7 setae each: M esocalanus quus Brady, 1883 (Fig. 7.14)
:#smo- tenuicornis (Fig. 7.18) 2a Total length 2.65-3.80 mm. Seta of A1 segment
-_l - ) 6a P5 Enp with different number of setae 00000000000007 23 shorter than last 7 segments: Calanus similli-
- ...
.... .) Mouth parts not very reduced. Right P5 Enp mus Giesbrecht, 1902 (Fig. 7.15)
Oaus. with 8 setae, left with 2 setae: Canthocalanus
pauper (Fig. 7 .16) Males
- a Mouth parts very reduced. Right P5 Enp with 1 Right P5 Exp extends beyond distal border of
tO!DI to different number of setae (not more than 6), left Exp1 .. .ooooooooooooooooooooooooooooooooooooooo.ooooooooooooooo .. 2
often left Enp lacking setae: Calanoides 1a Right P5 Exp does not extend as far as distal bor-
:ew: to der ofleft Exp 1: Calanus simillimus (Fig. 7 .15)
- ... 5
908 Copepoda
2 Right P5 Exp extends more than half way along elements (4 posterior and I anterior setae, and 9 ter-
left Exp2; rig ht P5 about 1.5-I .6 times shorter minal spines); Li2 and Li3 with 3-4 setae each; Li4
than left P5: Calanus australis (2.90-3.47 mm) with 4 setae; Enp with 13-14 setae; Exp with 11 setae;
(Fig. 7. 13), Calanus agulhensis (2. 74-3.00 mm) Le2 with 1 reduced seta; Le 1 with 7-9 setae. Mx2 Li 1- I
(Fig. 7 .12). These t\vo species are difficult to Li5 with 6, 3, 3, 3 and 3-4 setae respectively; term inal
distinguish. part wi th 8 setae, 1 of them may belong to developed
2a Right P5 Exp extends less than half way along Li6. Mxp C with 1 proximal seta, then 3 groups of 2,
left Exp2; right P5 about t\vice shorter than left: 3 and 4 setae from proximal to distal; Enp2-Enp6 with
Calanus propinquus (Fig. 7 .14) 3, 4, 3, 3+ 1 and 4 setae respecti vely. P l -P4 Exp usu-
ally 3-segmented but PI Exp 2-segmented in some
Key to species of Neoca/anus Calocalanus species. P2-P4 Enp usually 3-segmented
Females but 2-segmented in Calocalanus minor, posterior sur-
I PI B with large spine at base of inner seta. P2- faces of some segments ornamented with spines. P2-
P5 B naked............. ..... ........ ............... .......... ... ...2 P4 Exp3 external .borders serrated, or smooth in
1a PI B without large spine at base of inner seta. P2- De/ius and Calocalanus. Terminal spines of P2-P4
P5 B with conspicuous, posterior surface spinu- Exp smooth. Spine and seta formula:
les: Neoca/anus tonsus (Brady, 1883) (Fig. 7.22)
2 Spermathecae of typical structure; Gns slightly coxa basis exopod segments endopod segments
swollen ventrally: Neocalanus gracilis Dana, I
Leg I 0-011 0-0/1 0/T-0/ 1; 0- 1; I!,1,4 0- 1; 0/ 1,2,112 or
1849 (Fig. 7.20) or 0-0; 1,1,5 0,2,3
2a Left and right spermatheca enclosed under com- Leg2 0-1 0-0 I- 1; 1-l; 11,!,5 0- 1; 0-2; 1/2,2,3
mon cover; Gns very swollen ventrally: Neo- Leg 3 0-1 0-0 T- 1; I-I ; II,I,5 0- 1; 0-2; 2,2,3
calanus robustior (Giesbrecht, 1888) (Fig. 7.2 1) Leg4 0-1 0-0 1-1 ; 1-1 ; 11,1,5 0- 1; 0-2; 2,2,3
Males P5 uniramous, absent, or vestigial (Acrocalanus), pre-

1 P5 only slightly asymmetrical: Neocalanus ton- sent on left only (De/ius). If both P5 present, then sym-
sus (F ig. 7.22) metrical, presented by coxae (Bestiolina), 2-segmented
1a P5 significantly asymmetrical, with left leg (or 3-segmented) (Paracalanus, Parvocalanus), or 3-4
modified ....................... ......................................2 segmented (Calocalanus).
2 Setation of left P 5 Enp: Enp i and Enp2 with l
inner seta each, Enp3 with 6 setae: Neocalanus Males
gracilis (Fig. 7 .20) Ur of 5 somites. Geni tal opening o n left ( in
2a Setation ofleft P5 Enp highly reduced (5 setae in Calocalanus on left or right). Cephalic hump present
total): Neocalanus robustior (F ig. 7.2 I) (Acrocalanus, Bestiolina and Paracalanus), or absent.
A I 25 segmented, with some o r all of segments 1-6
Family Paracalanidae Giesbrecht, 1892 fused and swollen, segments 9- 10 may also be fused.
References: Giesbrecht (1892), Vervoort ( 1963), A2 Exp 7 very short without 3 terminal setae which
Andronov ( 1972, 1977), Bradford-Grieve (1994) are present in female. Expl and Exp 2 without setae.
Fem ales Other oral parts reduced compared with female. Right
Ce and Pdl usually fused. Pd4 and 5 fused, or separa- P5 may be absent (Acrocalanus, De/ius), or present as
ted. Ur of 2 to 4 somites. Anal somite usually longer C (Bestiolina), 2-3-segmented (Paracalanus, Parvo-
than any somite between it and Gns. R of 2 fi laments calanus), or 3-4-segmented (Calocalanus).
(Acrocalanus, Calocalanus, Paracalanus); 2-pointed,
solid (De/ius, Bestiolina); or massive (Parvocalanus). The family includes the genera Acrocalanus Gies-
A I usually 25-segmented, generally with segments l brecht, 1888; Bestiolina Andronov, 199 1; Calocalanus
and 2, also 8 and 9, partially fused; in De/ius first 8 Giesbrecht, 1888; De/ius Andronov, 1972; Para-
segments fused. A2 Exp 7-segmented with Expl and calanus Boeck, 1864; and Parvocalanus Andronov,
Exp2 each bearing 2 setae, segment 7 elongate. Md 1970. Of these only Bestiolina has not been recorded in
Enp 1 wi thout prominent lobe. Mx 1 Lil with I4 setal the South Atlantic.
Copepoda 909
-
~
~- Key to genera of Paracalanidae and male of Parvo- 4 Ur short, Pr:Ur = 3.3. Ce with moderately deve-
• ~H
T • .
calanus loped dorsal hump: Acrocalanus andersoni
~= Females Bowman, 1958 (Fig. 7.24)
l P 1 B with inner marginal seta ... ............... ......... 3 4a Ur relatively longer, Pr:Ur = 3.6-4.0. Ce is even-
ia P 1 B usually without iooer marginal seta ......... 2 ly rounded without any dorsal hump: Acro-
2 R of 2 slender filaments. P5 3-4-segmented: calanus gracilis Giesbrecht, 1888 (Fig. 7.26)
Calocalanus
2 R bifurcate, short, solid. P5 2-segmented: Males (unknown for A. monachus, incomplete descrip-
Parvocalanus tion for A. andersoni; see Vervoort, 1963)
3 R of 2 slender filaments. P3-P4 Exp3 external 1 P4 Exp3 with about 10-12 teeth on distal part of
edges at least partly serrated ............................ .4 segment: Acrocalanus gibber (Fig. 7.25)
-~- 3a R biramous, short. P3-P4 Exp3 external edges 1a P4 Exp3 with more than 10-12, usually about 15-
- p __ not serrated: De/ius 20 teeth on distal part of segment: Acrocalanus
.! Both P5 developed, symmetrical 2-3 segmented: gracilis (Fig. 7.26) and Acrocalanus longicornis
Paracalanus (Fig. 7.27) (these species are very difficult to dis-
~ P5 absent, or vestigial: Acrocalanus tinguish)
= )Iales Key to species of Calocalanus

:. a Cephalic hump present (lateral view) ........... .... 2 Females
b Cephalic hump absent (lateral view) ................. 3 1 Ur of2 somites: Calocalanus pavo (Dana, 1849)
Right P5 absent, or !-segmented: Acrocalanus (Fig. 7.36)
Right P5 2-3-segmented: Paracalanus la Ur of 3-4 somites ............................................... 2
Right P5 absent: De/ius 2 Ur of 3 somites ................. ...... ......... ....... ........ .. .4
-~ Right P5 present, 2-4 segmented ......... .. .. .... ..... .4 2a Ur of 4 somites .............................. .................... 3
-!. Right P5 4-segmented: Calocalanus 3 P5 terminal segment with 4 spines: Calocalanus
~ Right P5 2-, or 3-segmented (Parvocalanus) ; P5 at/anticus Shmeleva, 1975 (Fig. 7 .29)
2-segmented, does not extend as far as distal 3a P5 terminal segment with 3 spines: Calocalanus
border of left leg segment 1: Parvocalanus cras- equalicauda (Bernard, 1958) (Fig. 7.31)
sirostris (Dahl, 1894) (Fig. 7.51) (males unk- 4 P5 3-segmented ................................................. 5
nown for P. scotti) 4a P5 4-segmented ................................................. 9
5 P5 terminal segment with 2 long terminal setae
m to species of Acrocalanus (based on Bowman, (longer than terminal segment) ......................... 6
1958; Tanaka, 1956; Bradford-Grieve, 1994) 5a P5 terminal segment with 1 terminal seta ......... 7
-:anales 6 Terminal spines of P5 terminal segment longer
I Body of square appearance in lateral view: than leg itself: Calocalanus contractus Farran,
Acrocalanus monachus Giesbrecht, 1888 (Fig. 1926 (Fig. 7.30)
- .28) 6a Terminal spines of P5 terminal segment as long
Body not of square appearance in lateral view .2 as leg itself: Calocalanus lomonosovi Shrneleva,
.-\1 exceeds beyond end ofCR by 2 tenninal seg- 1975 (Fig. 7.32)
ments. Dorsal outline of Ce significantly hum- 7 Terminal spine ofP5 terminal segment nearly as
ped: A crocalanus gibber Giesbrecht, 1888 (Fig. long as, or longer than leg itself.. ..... .............. ... 8
- .25) 7a Terminal spine ofP5 terminal segment is not lon-
.-\1 exceeds end ofCR by 3-6 terminal segments. ger than terminal segment itself: Calocalanus
Dorsal outline of Ce moderately humped, or not minor Shrneleva, 1975 (Fig. 7.33)
humped ................. ................. .... .......... .... ...... .... 3 8 Terminal spine ofP5 terminal segment nearly as
A 1 exceeds end of CR by 5-6 terminal segments. long as ieg itself: Calocalanus styliremis
P~ Exp3 with 15-20 fine teeth: Acrocalanus Giesbrecht, 1888 (Fig. 7.38)
longicornis Giesbrecht, 1888 (Fig. 7.27) 8a Terminal spine of P5 terminal segment is longer
.-\ I exceeds end ofCR by 3-4 terminal segments. than leg itself: Calocalanus minutus Andronov,
P~ Exp3 with 11 -15 coarse teeth ...................... .4 1973 (Fig. 7.34)
910 Copepoda
9 P5 terminal segment with I setal element: Calo- Males

calanus namibiensis Andronov, 1973 (Fig. 7.35) 1 Specimens 0.43 mm tn length: De/ius sewelli
9a P5 terminal segment with 2-3 setal elements .. ! 0 (Fig. 7.42)
I 0 P5 terminal segment w ith 2 spines (1 shorter Ia Specimens 0.42-0.52 mm in length: De/ius
than other): Calocalanus tenuiculus Andronov, nudus (Fig. 7.41)
1973 (Fig. 7.39)
I Oa P5 terminG!I segment with 3 setal elements ( 1 Key to species of Paracalanus
spine and 2 plumose setae) .... .. ........................ 11 Females
II CR asymmetrical: Calocalanus plumulosus 1 A I longer than body. Presence of setae on pos-
(Claus, 1863) (Fig. 7.37) terior Pr is frequent.. .......... .......... .............. .. .. .. .. 6
ll a CR symmetrical: Calocalanus tenuis Farran, Ia A l shorter than body. Presence of setae on pos-
1926 (Fig. 7.40) terior Pr is rare ...... ............................................. 2
2 Gns usually with posterodorsal spinules along
M ales (unknown for C. at/anticus, C. minor, C. ten- posterior border .... .. ....................... .................... 3
uis, C. tenuiculus; C. equalicauda known after fifth 2a Gns without posterodorsal spinules along poste-
copepodite stage only) rior border ...... ..................... ... ...... ......................4
1 Right P5 reaching distal border of left leg seg- 3 Pr:Ur = 3.S: Paracalanus tropicus Andronov,
ment 1 ..... ....... .... .............................. .. ...... .. ........ 2 1977 (Fig. 7.49)
I a Right P5 exceeding distal border of left leg seg- 3a Pr:Ur = 3.0: Paracalanus indicus Wolfenden,
ment 1 ........... .......... .. ............................... .... ......4 1905 (Fig. 7.46)
2 Terminal segment of right P5 with 3 small spi- 4 External distal edge of P3-P4 Exp 3 serrate:
nes: Calocalanus lomonosovi (Fig. 7.32) Paracalanus quasimodo Bowman, 197 1 (Fig.
2a Terminal segment of right P5 with 1, or 2 small 7.50)
unequal spines ................................................... 3 4a External distal edge ofP 3-P4 smooth ............... S
3 Ce and Pd I separated by constriction. PS 5 A1 extends to posterior border of Ur3: Para-
extends well beyond CR : Calocalanus minutus calanus parvus (Claus, 1863) (Fig. 7.48)
(Fig. 7.34) Sa A I extends to posterior border of Gns: Para-
3a Ce and Pdl are not separated by constriction. P5 calallus nan us Sars, 1907 (Fig. 7.47)
extends slightly beyond CR : Calocalanus nami- 6 P4 Exp2 has leaf-like spines: Paracalanus acu-
biellsis (Fig. 7.3S) leatus Giesbrecht, 1888 (Fig. 7 .43)
4 Total length less than 0.8 mm .......... .......... .. ..... 5 6a P4 Exp2 lacking leaf-like spines: Paracalanus
4a Total length over 0. 8 mm .......................... .. ...... 6 campaneri Bj0rnberg, 1980 (Fig. 7.44); Para-
S Terminal segment of PS left with l , or 3 spines: calanus denudatus Sewell, 1929 (Fig. 7.45)
Calocalanus styliremis (Fig. 7.38) It is very difficult to distinguish between Paracalanus
Sa Terminal segment of P5 left with 2 setae: aculeatus, P. camp aneri and P. denudatus. Bj0mberg
Calocalallus contractus (Fig. 7.30) (1980) refers to the difference in the shape of the
6 Right PS leg does not extend as far as distal bor- receptaculum serninis in P. campaneri and P. acule-
der of segment 2 of left leg: Calocalanus pavo atus as the main difference between these two species.
(Fig. 7.36)
6a Right P5 leg extends beyond segment 2 of left Males (unknown for P. campaneri, P. tropicus; P.
leg: Calocalanus plumulosus (Fig. 7.37) denudatus and male P. quasimodo are not included
because of a lack of detailed description)
Key to species of De/ius 1 Specimens over 0.60 mm in total length ........... 2
Females I a Specimens less than 0.60 mm in total length:
I P5 3-segmented. PI Exp3 with 1 outer spine : Paracalanus nanus (Fig. 7.47)
De/ius sewelli Bj0rnberg, 1980 (Fig. 7.42) 2 Right P5 extending as far as proximal third of left
l a P5 2-segmented. P 1 Exp3 with 2 outer spines: leg segment 3: Paraca/anus aculeatus (Fig. 7.43)
De/ius nudus (Sewell, 1929) (Fig. 7.41 ) 2a Right PS not extending as far as distal border of
left leg segment 2 .......... .................................... 3
Copepoda 91 1
3 P2-P4 B with many posterior surface spinules: P4 similar to female, but PI -P4 C with inner seta
Paracalanus indicus (Fig. 7.46) each. P5 uniramous, 5-segmented, asymmetrical, lon-
3a P2-P4 B without posterior surface spinules: ger on right.
Paracalanus parvus (Fig. 7.48)
Monotypic fam ily contammg M ecynocera clausi
Key to species of Parvocalanus Thompson, 1888 (Fig. 7.53).
Females
l PI Enp 2-segmented. P5 terminal segment just Famil y E ucalanidae Giesbrecht, 1892
pos- under 0.5 times as wide as long with 2 subequal References: Bradford-Grieve ( 1994), Brodsky et a/.
+----6 strong terminal spines: Parvocalanus crassiro- (1983), Fleminger (1973)
pos- stris (Da hl , 1894) (Fig. 7.51) Females
.., Ia P 1 Enp !-segmented (Vervoort, 1963). P5 termi- Body elongate, especially anterior to A2, more or less
~g nal segment about 0.3 times as wide as long with transparent, often with triangular anterior part of Ce.
- ____3 inner tenninal spine longest: Parvocalanus scot- Ce and Pdl fused, Pd4 and 5 partly fused.CR someti-
ti (Fruchtl, 1923) (Fig. 7 .52) mes asymmetri cal; usually left ramus slightly and
second seta much longer than others. R of 2 long,
Family Mecynoceridae Andronov, 1973 slender filaments. Ur of 3 or 4 sornites w ith CR usu-
References: Andronov ( 1970), Bradford-Grieve ( 1994) ally completely fused to anal somite. A 1 23-segmen-
F emales ted, generally with segments J-2 and 8-9 fused. A2
Ce and Pdl , as wel l as Pd4 and 5 separate. R with 2 Exp shotter than Enp; Expl and Exp2 w ith 2 setae
e::'ZR': long filaments. Ur of 3 somites with articulated CR. each, and either fused or separate.
Fig. AI very long, 23-segmented, segments 1-2, 8-9 fused; Md Enp reduced and B and Exp elongate (except in
carrying elongate setae on segments 7, 14, 16, 18, 21, Rhincalanus). Mx I Li 1 with 14 tenninal spines and
23 and 24. A2 Exp 8-segmented, shorter than Enp; setae including 4 posterior and I anterior setae; Li2
Exp l and Exp2 with 2 setae each, terminal segment may be absent or without setae, B and Enp elongate;
short. Mx I L i l with 12 terminal setae including 4 Exp with reduced number of setae; Le2 with l seta;
posterior setae; Li2 and Li3 w ith 3 setae each; Li4 Lei w ith 9 setae. Mx2 Lil -LiS with 6, 3, 3, 3 and 3-4
\\ith 4 setae; Enp with 14 setae; Exp with It setae: setae, respectively. Mxp Enp2-Enp6 with 3, 4, 3, 4
rs acu- Le2 small, lacking setae; Let with 9 setae. Mx2 Li 1- and 4 setae respectively. Tenninal Exp spines of
Li5 with 5, 3, 3, 3 and 4 setae respectively. Mxp C swim ming legs smooth. Spine and seta fonnula:
-.ltutJIS '\\ith groups of 2, 3 and 4 setae from proximal to dis-
PIITtl- tal: B with prox imal row of long teeth; Enp2-6 with 4, coxa basis exopod segments endopod segments
-5) -t 3, 4 and 4 setae respectively. PI Enp !-segmented
Leg I 0-0 0-1 0- 1; 0-1 ; 1,1,4 or 0- 1; 0,2,2
~ with an outer lobe. Spine and seta formula: l-1 ; !1,!,5
ac::..~ Leg 2 0-1 0-0 l-1 ; I- I; lll.l.5 0-1; 0- 1; 1,2,2
o:· ihe Coxa basis exopod segments endopod segments Leg 3 0-1 0-0 l-1 ; l- 1: 111,1,5 0- 1; 0- 1; 1,2,2
.. a...-:..:e- Leg 4 0-1 0-0 J- 1; I- I ; Ill,l,5 0-1 ; 0- l ; l ,2,2
leg 1 0-0 0-1 0- 1; 0- 1; 1,1,4 0,1,2
leg2 0-0 0-0 1- 1; 1-1 ; ni ,l,5 0-l ; 0- 1; 1,2,2
leg 3 0-0 0-0 1-1; 1- 1; 111,1,5 0-1; 0-1 ; 1,2,2 P5 absent (Eucalanus, Pareucalanus, Subeucalanus),
leg 4 0-0 0-0 1- l ; 1-1 ; Ill,1,5 0-1 ; 0-1 ; l ,2,2 or present, 3-segmented, symmetrical.
P5 uniramous, 5-segmented, symmetrical; no outer Males

.., edge spines on each Exp segment but l inner edge Ur of 5 somites, with CR usually completely fused to
---·--
seta on Exp2 and 1 tenninal spine and 4 inner edge anal somite. AI 24-segmented. Oral patts and P similar
setae on Exp3. to females. P5 uniramous on both sides (Eucalanus,
c a:· ~en: Pareucalanus), biramous on left (Rhincalanus), or
:: -.~3 ) :\!ales absent on tight (Subeucalanus).
___
c:-...c:r of
...;,
~
Pr as in female. Dorsal cephalic hump present. Ur of
5 so mites, with articu Iated CR. A l 25-segmented (21 The fami ly includes the genera Eucalanus Dana,
free segments) with segments 1-2, and 3-6 fused. PI- 1852; Pareucalanus Geletin, 1976; Rhincalanus
912 Copepoda
Dana, 1852 and Subeucalanus Geletin, 1976, which Ia P5 tem1inal segment with plumose outer setae
are all found in the South Atlantic. (or spine), much longer than its segment.. ........ 2
2 Dorsal spines absent from Pd3 and 4 : Rlzinca-
Key to genera of Eucalanidae lanus gigas Brady, 1883 (Fig. 7.58)
Females 2a Dorsal spines present on Pd3 and 4: Rhinca-
1 P5 present. Dorsal surface of at least some of lanus nasutus Giesbrecht, 1888 (Fig. 7.59)
Pd2-4 and Gns with 2 spines: l on right and l on
left. Posterior comers of at least some Pd2-4 Males (poorly described for R. cornutus)
extended into spines: Rhincalanus I Specimens are over 6.5 mm: Rlzincalanus gigas 0
Ia P5 absent. Dorsal surface of Pd2-4 and of Gns (Fig. 7.58)
without spines. Posterior comers of Pd2-4 lac- la Specimens are less than 6.5 mm length:
king spines ............................................. .. .... ...... 2 Rhincalanus nasutus (Fig. 7.59)
2 Female Ur of 3 free somites (somite 4 fused with
CR): Eucalanus hyalinus Claus, 1866 (Fig. 7.54) Key to species of Subeucalanus
2a Female Ur of2 free somites (somite 3 fused with Females
CR) ..................................................................... 3 1 Anterior part of Ce triangular (dorsal view) ..... 2
3 A2 Expl and 2 fused. Mxl with 2 inner lobes: la Anterior part ofCe rounded (dorsal view) ...... .. 3
Subeucalanus 2 Anterior part ofCe triangular and elongated (dor-
3a A2 Exp l and 2 separate. Mxl with 3 inner lobes: sal view). Mdp B wi th 3 setae. Mxl Li4 with 5
Pareucalanus setae: Subeucalanus longiceps Matthews, 1925
(Fig. 7.6 1)
Males 2a Anterior part of Ce triangular, but not elongated
I At least Gns with 2 dorsal spines. Left P5 bira- (dorsal view). Mdp B with2 setae. Mxl Li4 with
mous: Rhincalanus 4 setae: Subeucalanus subtenuis (Giesbrecht,
1a Gns without dorsal spines. P5 uniramous ......... 2 1888)(Fig. 7.64)
2 R ight P5 4-segmented: Eucalanus hyalinus 3 Gns much wider than long and onion-shaped
(Fig. 7.54) (dorsal view): Subeucalallus crassus (Gies-
2a Right P5 3-segmented, or absent ....................... 3 brecht, 1888) (Fig. 7.60)
3 Right P5 absent: Subeucalanus 3a Gns nearly as wide as long and of different shape
3a Right P5 present, 3-segmented: Pareucalanus (dorsal view) ..................................................... .4
4 In lateral view distal part of seminal receptacles
Key to species of Pareucalanus bilobed: Subeucalanus pileatus (Giesbrecht,
Females 1888) (Fig. 7.63)
I Mdp B with 4 setae: Pareucalanus lcmgae 4a 1n lateral view distal patt of seminal receptacles
(Fleminger, 1973) (Fig. 7.55) not bilobed: Subeucalamts monachus (Gies-
Ia Mdp B with 2 setae: Pareuca/anus sewelli brecht, 1888) (Fig. 7.62)
(Fieminger, 1973) (Fig. 7.56)
Males
Males I Terminal seta on left P5 longer than terminal
I Total length over 6 mm; Pareucalanus langae segment: Subeucalanus subtenuis (Fig. 7.64)
(Fig. 7.55) Ia Terminal seta on left P5 as long as, or shorter
la Total length less than 6 mm: Pareucalanus than terminal segment ....................................... 2
sewelli (Fig. 7.56) 2 Males of Subeucalanus are poorly known morp-
hologically, and their morphology is also highly
Key to species of Rhincalanus variable. The following males cannot be separa-
Females ted in the identification key, but their sizes and
I P5 terminal segment with smooth outer terminal figure references are as follows: Suheucalanus
spine, shorter than its segment: Rhincalanus pileatus (total length 1.80-2.25 mm) (Fig. 7.63);
cornutus (Dana, 1849) (Fig. 7 .57) Subeucalanus monachus (total length 2.2 mm)
Copepoda 9 13
a- setae (fig. 7.62); Subeucalanus longiceps (total and Sognocalanus. Pl Expl without outer spine in
1________ 2
length about 3.2 mm) (Fig. 7.61); Subeucalanus Mimocalanus, and Teneriforma; both Exp l and Exp2
illlinca- crassus (total length 2.9-3.1 mm) (Fig, 7.60). without external spines in Jsaacsicalanus; Enp with
external lobe. Surfaces of some segments of swim-
tllrinca- i="arn.ily Spinocalanidae Yervoort, 195 1 ming legs may be spinulose. Spine and setae formula:
iteferences : Damkaer (1975), Brodsky et al. (1983),
Sdmlz ( 1989, 1996) coxa basis cxopod segments cndopod segments
Females
!Iigas Leg I 0-0 0-1 O/I-O; on-1 ; 1,1,4 0,2,1/2/3
Ce and Pd I separate, Pd4 and 5 separate or partly Leg 2 0- 1 0-0 1- 1; 1-1 ; III,l,5 0- 1; Oil ,2,2
~sed. (!saacsicalanus), may be prolonged into lobes Leg 3 0-1 0-0 1-1 ; 1-1 ; lll,I,5 0-1 ; 0-1 ; 112,2,2
:ength: Honacilla, some species of Spinoca/anus). Pd 1-4 Leg4 0- 1 0-0 1- 1; 1- 1; III,l,5 0- 1; 0-1; 1/2,2,2
~erimes with lateral spinules. Ur of 4 somites, Gns
~-.mnetrica l or asymmetrical (Monacilla). R 2-pointed PS absent.
~Honacilla), !-pointed (Damkaeria); single long blunt
=one (Teneriforma, Kunihu/sea, Tsaacsica/anus); or Males
~ ) .....2 ..2senr (Spinoca/anus, Mimoca/anus Mospica/anus) . Ce and Pd l partly fused or separate, Pd4 and 5 separate.
I"------·3 --~1 23 -24-segmented, 8-9 segments fused. A2 Exp of Ce in Spinocalanus usually expanded dorsally to nearly
a: dor- '!~..,.~or greater length than the Enp; Exp 7-segmented; cover Pd I. R a reduced knob, long cone, or absent. Ur
- ith 5 E.·cpl w ith 0-2 setae; Exp2 with 2-3 setae (majority of of 5 somites. Anal somite usually reduced
IS. 1925 ~alanids), or I seta (Jsaacsicalanus), seta absent (Spinocalanus) or not reduced (Teneriforma). A1 with
(!;" reduced (Mimocalanus, Spinocalanus part. and segments 8-9 fused, segment I0 separate from or fused
locgared .? .i'lomaxillaris). Exp2 and Ex p3 separate partly or completely with segment 9; segment 20 merg-
U-\\ith \firzocalanus, Tsaacsica/anus), patily fused, or coming w ith 21 , with noticeably wrinkled margins, on right
:s.xecht. ~ely fused (Spinoca/anus, Teneriforma). Enp I with side in species with left-handed PS and on left in species
.: setae. Md Enpl with 2-4 setae, Enp2 with 10-11 with right-handed PS; segments 24 and 25 separate, seg-
t-s1uped S-~e (9 in lsaacsicalanus). Mx l Li1 with 9 terminal ment 25 may be reduced. A2, Md, Mx I, Mx2 and Mxp
1 (Gies- ~ and 0-4 posterior setae, with 1 mid-marginal reduced. PI-P4 generally similar to those offemale. PS
..-crior surface seta, and with or without 1 small dis- simple, B uneven, usually longer on left (left-handed):
a shape ~ anterior surface seta; Li2 with 4-6 setae (3 setae in PS biramous, present on both sides, Exp usually 3-seg-
-------·.4 !J.xnkaeria, 2 setae in Kunihulsea and I seta tn mented, Enp styliform (majority of Spinocalanus spe-
~des lscL:Icsicalanus); Li3 w ith 4 setae (3 setae m cies); or one of Exp not styliform (Monacilla); m1ira-
:S:mcht. J..lJmihulsea, 2 setae in Teneriforma, seta absent in mous, sometimes present on right side only,
fuacsicalanus); Li4 with 5 setae(4 setae m 4-segmented (Damkaeria), or uniramous and well
~des !_qJicalanus, 3 setae inKunihu/sea); Enpl plus Enp2 developed on left side only, 5-segrnented, but reduced
J G1es- -=ies from 3 to 7 setae (10 setae in Kunihulsea); Exp on right (some Spinocalanus species, Teneriforma).
r--;-..h 9(?}-11 setae; Le2 small with (Monacilla), or wit-
,_.. minute seta (Spinocalanus pati., Mimocalanus) , The family contains the genera Damkaeria
oc absent (Spinocalanus part.); Lei with 9 setae. Fosshagen, 1983; Tsaacsicalanus Fleminger, 1983;
~al Se:arion of Mx I deviates strongly in Rhinomaxillaris. Kunihulsea Schulz, 1992; Mimocalanus Farran, 1908;
- .6.!) 1\t~ Li l -LiS with 5-7, 3, 3, 3 and 2-4 setae respecti- Monacilla Sars, 1905; Mospica/anus Schulz 1996;
r shorter ~~-= external seta present (Monacilla, Spinocalanus Rhinomaxillaris Grice and Hi.ilsematm, 1967;
------·2 ,__L or absent; sometimes Li6 is quite well-developed Sognocalanus Fosshagen, 1967; Teneriforma Grice
:norp- 1ii::h setae (2 setae in Monacilla, Kunihulsea and and Hulsemann, 1967; and Spinocalanus Giesbrecht,
c ;:ighly ~"lCa!anus). Mxp C wi th I proximal seta (someti- 1888. Of these, only Mimocalanus, Monacilla, and
! 5.-.
~ c:rs absent), and usuall y with groups of 2, 3 (l seta in Spinocalanus have been found in the South Atlantic.
IZ...
~ and - eneri(orma, absent in Jsaacsicalanus) and 3 (4 in
!Oihurus fimoca lanus, 2 setae in Teneriforma and 1 seta in Key to genera of Spinocalanidae
c- - .03 ):
-sica/anus) setae from proximal to distal; C and B Females
:=.....: mm) SXiletimes with transverse spine-comb. Enp1-Enp5 1 PI Exp 1 without outer spine. Posterior surface
-~Y with 4, 2-4, 2-3, 3- 4 and 4 setae respectively. of P2-P4 Enp2 without row of spines:
~on of Mx2 and Mxp deviate in Rhinomaxillaris Mimocalanus cultrifer Farran, 1908 (Fig. 7.65)
914 Copepoda
la PI Expl with outer spine. Posterior surface of 3a Pdl -4 without lateral spinules: Spinocalanus
P2-P4 Enp2 with 2 rows of spines .................... 2 hrevicaudatus Brodsky, 1950 (Fig. 7.69)
2 R absent. Mxl Le2 absent, if present lacking
seta: Spinocalanus Males (doubtful description for S. horridus)
2a R in form of 2 strong points. Mx I Le2 small 1 P2 Enp2 with 1 outer seta ............ ................. ....2
with minute seta: Monacilla la P2 Enp2 without I outer seta: Spinocalanus
ahyssalis (Fig. 7.68)
Males (undescribed for Mimocalanus cu/trifer, un- 2 CR nearly as long as wide. P5 not extending to
known for Monacilla gracilis) posterior border of Ur3. Pl Enp with 2 inner
1 P5 uniramous, present on both sides, usually 5- setae: Spbwcalanus brevicaudatus (Fig. 7.69)
segmented. PI Exp I without outer spine: Mimo- 2a CR nearly twice as long as wide. PI Enp with 3
calanus inner setae. P5 nearly reaching posterior border
la P5 biramous, present on both sides. Exp usually ofUr3: Spinocalanus magnus (Fig. 7.7 1)
3-segmented (if uniramous, strongly asymmetri-
cal); P 1 Exp 1 with external spine (in Monacil/a Family C lausocalanidae Giesbrecht, 1892
typica external spine absent) ............................. 2 References: Frost and Fleminger (1968), Bradford-
2 Anal somite usuall y reduced. P5 Enp styliform Gri eve (1994)
on right and on left: Spinocalanus Females
2a Anal somite half length of CR. P5 Enp not styli- Pdl and Ce usually fused, sometimes separate, Pd4
form, at least on right: Monacilla typica Sars, and 5 usually fused. R usually with 2 filaments, two-
1905 (Fig. 7.67) pointed, or absent. Posterior corners of Pr usually
The new genus and species Mospica/anus schielae symmetrical, rOtmded, sometimes extended into trian-
Schulz, 1996, described recently from 60°26' S, gular lobes (Farrania). Gns usually symmetrical. A 1
50°48'W is not included in the above key. generally with segments 8-9 fused and sometimes
with segments 1-2 and 24-25 fused; extremely long
Key to species of Monacilla setae absent. A2 Exp 6-segmented (Clausocalanus,
Females (male known only forM. typica) Ctenoca/anus, Drepanopus, Microcalanus) or ?-seg-
1 Gns symmetrical, not protruding on right. P4 C mented (Farrania); Expl with 1 or 2 setae, or setae
inner posterior surface with transverse row of spi- are absent; Exp2 with 2 medial setae, or setae are
nules, left and right leg of equal length: absent. Coxa with 1 seta, B with 2 setae; Enp 1 with 2
Monacilla gracilis (Wolfenden, 191 1) (Fig. 7.66) setae. Mdp B with 4 setae; Enp 1 with 4 setae and
Ia Gns asymmetrical, protruding on right. P4 C Enp2 with 9 terminal and 1-2 appendicular setae
inner posterior surface with transverse row of (C/ausocalanus, Drepanopus), or 5 (Farrania pacifi-
spinules, longer on left, sometimes absent on ca) setae. Mxl Lil with 9 terminal, l anterior and 4
right: Monacilla typica Sars, 1905 (Fig. 7.67) posterior setae; Li2 with with 5 setae, Li3 with 4
setae, Li4 with 5 setae; Enp with up to 16 setae; Exp
Key to species of Spinocalanus with II setae; Le l with 9 setae (Clausoca/anus). Mx2
Females Li l-Li5 with 3-5, 3, 3, 3 and 4 setae respectively; Li6
I Pl Enp with 3 inner setae. Mxp B without trans- and terminal part oflimb usually with 6 setae, or up to
verse spine-combs: Spinocalanus magnus 7 setae. Mxp C with 1 proximal setae, then with
Wolfenden, 1904 (Fig. 7.7 1) groups of 2, 3 and 3 setae from proximal to distal;
la PI Enp with 2 inner setae. Mxp B usually with Enp2-6 with 4, 4, 3, 4 and 4 setae respectively, B with
transverse spine-comb ......... .............................. 2 inner proximal row of fine spinules (in Drepanopus
2 P2 Enp2 without outer seta: Spinocalanus abys- setation is reduced). Pl Enp with external lobe, or
salis Giesbrecht, 1888 (Fig. 7.68) may be absent (Microca/anus, Drepanopus bungei).
2a P2 Enp2 with outer seta .... ...... ......... .... .... .......... 3 fn Drepanopus P2 Enp may be !-segmented and Enp
3 Pd 1-4 with spinules on right and left sides: of P4 not completely 3-segmented (D. bungei).
Spinocalanus horridus Wolfenden, 1911 (Fig. Posterodistal border of B of P2 and 3 may be orna-
7.70) mented with spines, Enp of P2-4 usually narrow.
Copepoda 915
.Jimus ..~ of legs with or without spinules. Spine and 1a P3-P4 Exp spines without comb-like serration.
~ f.lx:u::;..:!a: P5 synunetrical, both left and right legs present
......................................... .. ...... .. ........................ 2
______ 2
= basis exopod segments cndopod segments 2 P2-P3 B w ith spiniform processes distally:
Clausocalmms
0.1 0 1-0; 1-1; I,I,3 0,2,3/2
.Jim us 2a P2-P3 B without spiniform processes ............... 3
0.0 1- 1; 1-1; In,l,4 0-1 ; 1/0,2,2
0.0 1-1 ; I-I ; Ill,l,4 0-1 ; 0- 1; 1,2,2 3 Gns with late ral swellings anteriorly (dorsal
6r!g to ~- (U) 0.0 1-1; 1-1; m ,I,4 0-1 ; 0- 1; 1.2,2 view). Posterior comers of Pr rounded: Drepa-
: ::=mer nopus forcipatus Giesbrecht, 1888 (Fig. 7.86)
- .09)
~ih
3
: -=-~ (.\licrocalanus, Pseudocalanus), or present. 3a Gns without anterolatera l swellings (dorsal
~L uniramous on both sides, symmetri cal, 3- view). Posterior comers of Pr extended into
:xmler ~--d terminally produced into short bifid, poin- tria ngular lobes: Farrania frigida (Wolfenden,
a::::: ;rocesses (Clausocalanus); or with 2-3 terminal 191 I ) (Fig. 7.87)
- Farrania); or consisting of vety sma ll appen-
"""'='- &\·eloped only on left and of 2-4 segments Males
.nord- ::ca/anus): or symmetrical, 2-segmented with I P5 uniramous .. .. ................................................. 2
----;=.:::n ed terminal spine, pectinated along its distal la P5 biramous; Enp, although rudimentary, pre-
...::: ::.aif (Drepanopus). sent in both legs; left leg moderately longer, or
e. Pd~ shorter than right leg: Farrania frigida (Fig.
5.. i'WG- 7.87)
5ualiy -.-.b segments 1-2 and 8- 10 always fused; several 2 Left P5 5-segmented, distinctly longer than right
) aian- segments may be also fused. A2, Md, Mx l and leg which is 1-3-segmented ............................... 3
....... .-\1 :L :::,-ually reduced in size and setation compared 2a Left P5 4-segmented, shorter than right leg
et!:nes ;ana]e. P l -P4 as in female, although there may which is 4-segmented: Drepanopus forcipatus
.ong '!nne sexua l dimorphism in the propmtions of legs (Fig. 7.86)
rltrzZJS. s:pinulation. P5 present on both sides, but of 3 Right P5 with tetminal spinules on distal seg-
--sa!:-
c:::e,_~ length, uni.ramous; longer leg nearly always ment. Left P5 segment 5 attached to segment 4
r :;etae f:.. 5-segmented, shorter leg 1-3 segmented, less suba pically: Clausocalanus
~are
balf length of C of other leg (Clausocalanus, 3a Right P5 is shott stump which lacks spines and
Jllri-.h 2 CJJn:OCalanus), or longer (Microcalanus); or both legs spinules. Left P5 segment 5 attached to segment
t.! and - siu:J.ilar length (Pseudocalanus), sometimes even 4 apically: Ctenocalanus
s...'"tae - :-.2dimentary Enp (some species of Drepanopus,
r-~ti F.::n-=nia). Key to species of Clausocalanus (from Frost and
:md 4 F leminger, I 968, with minor changes)
ri-.il ...1 ~ are at least 6 gene ra in the family: the species Females
~E.~ - genera, Clausocalanus Giesbrecht, 1888; 1 G ns length about equal to length of Ur3:
~~b2 Ctzr'Oea/anus Giesbrecht, 1888; Drepanopus Brady, Clausocalanus jitrcatus (Brady, I 883) (Fig.
• Li6 ~= and Farrania Sars, I 920 occur in the South 7.74)
rt;> to ~c. Other genera: Microcalanus Sars, 1901 , Ia Gns more than 1. 5 times as long as Ur3 ........... 2
l llli"ith Psn.docalanus Boeck, 1873, and Spicipes Grice and 2 In lateral view, ventral profi le of Gns conspi-
~= • s....omann, 1965 have not been recorded in the South cuously protuberant ante rior to genital pore:
~v.ith -\~tic. The genus Clausocalanus was revised by Clausocalanus jobei Frost and Fleminger, 1968
r::opus - and Fleminger (1968). (Fig. 7.76)
be. or 2a In lateral v iew, ventral profile of Gns not cons-
rgei). • • to genera of Clausocalanidae picuous ly protuberant anterior to genital pore
cEnp F lm3.les ................................................................... ........ 3
~~i). P3-P4 Exp outer spines w ith comb-like serration 3 Third segment of P5 less than 2.0 times as long
OG13- along inner marg in. P5 asymmetrical, single leg as first segment ofP5 .................................... ... .4
r;:;ov.·. ;:>resent: Ctenocalanus 3a Third segment ofP5 more than 2.0 times as long
as first segment ofP5 ...................... .................. 7
916 Copepoda
4 Forehead usually strongly vaulted. Distance (L) 9a In lateral view, portion of seminal receptacle
between spiniform processes 2 and 3 of P3 B ventral lobe visible anterior to dorsal lobe; semi-
equal to or more than 2.0 times p roximal width nal receptacle dorsal lobe usually bulb-shaped
(D) of process 3; process 2 and 3 uniformly tape- .............. .............. ............................................. 10
ring from base. CR more than 1.65 times as long 10 In lateral view, seminal receptacle dorsal lobe
as wide: Clausocalanus laticeps Farran, 1929 digitiform or slightly constricted in region of
(Fig. 7.77) attachment to seminal receptacle ventral lobe. In
4a Forehead not vaulted. Distance (L) between spi- lateral view, posterior comers ofPr rounded. A I
niform processes 2 and 3 of P3 B less than 1.5 segment 2 Jess than 0.95 times as long as seg-
times proximal width (D) of process 3 of P3; ment 24. P4 Exp 3 less than 1.30 times as long
process 2 and 3 not uniformly tapering from as terminal spine of P4: Clausocalanus pergens
base. CR Jess than 1.65 times as long as wide Farran, 1929 (Fig. 7.83)
........................................ ................................... 5 I Oa In lateral view, seminal receptacle dorsal lobe
5 In lateral view, R slender and curved. In lateral bulb-shaped, constricted in region of attachment
view, forehead broadly rounded, protuberant to seminal receptacle ventral lobe. In lateral
above R; greatest anterior extension of forehead view posterior corners of Pr somewhat angular.
well dorsal to frontal organ: Clausocalanus Al segment 2 more than 0.95 times as long as
ingens Frost and Fleminger, 1968 (Fig. 7.75) segment 24. P4 Exp3 more than 1.30 times as
Sa In lateral view, R thick, usually curved ventra- long as terminal spine o f P4 .... ..................... ... 11
posteriad. In lateral view, forehead broadly II CR more than 1.52 times as long as w ide. Al
rounded but not markedl y protuberant above R; segment 2 more than 1.40 times as long as seg-
greatest anterior extension offorehead at level of ment 24: Clausocalanus brevipes Frost and
frontal organ or rarely slightly dorsal to frontal Fleminger, 1968 (Fig. 7.73)
organ ............. .... ............... .................................. 6 11 a CR less than 1.52 times as long as wide. A I seg-
6 In lateral view R usually straight, tapering uni- ment 2 less than 1.40 times as long as segment
formly in either lateral or anterior view; in later- 24: Clausocalanus parapergens Frost and
al view R usually directed ventrad or, rarely, cur- Fleminger, 1968 (Fig. 7.81)
ved ventroposteriad: Clausocalanus lividus
Frost and Fleminger, 1968 (Fig. 7.78) Males
6a In lateral view, R usually curved ventroposte- 1 In lateral view, seminal vesicle extending within
riad; R not tapering uniformly in either lateral or Pr anterior to level of P l articulation with Pr.
anteroventral views: Clausocalamts mastigop- Spermatophore extending by at least 1/4 of its
horus (Claus, 1863) (Fig. 7.79) length anterior to articulation of Pd I with Pd2
7 Distance (L) between spiniform processes 2 and .... ......................... ....... ... ........... ........... ...... ........ 2
3 of P3 B more than 2.0 times proximal width Ia In lateral view, seminal vesicle not extending
(D) of process 3 of P3: Clausocalanus paululus within Pr anterior to level of Pl articulation with
Farran, 1926 (Fig. 7.82) Pr. Spermatophore not extending or extending
7a Distance (L) between spiniform processes 2 and by less than 118 of its length anterior to articula-
3 ofP3 Bless than 1.5 times proximal width (D) tion of Pd I w ith the Pd2 .. .............. ................... .5
of process 3 of P3 ............................ .................. 8 2 Total length less than 0.70 mm: Clausocalanus
8 In lateral view, ventral profile of Gns with pro- paululus (Fig. 7.82)
minent step posterior to genital pores: 2a Total length greater than 0.70 mm .................... 3
Clausoca/anus minor Sewell, 1929 (Fig. 7.80) 3 Pr less than 5.7 times as long as Ur2: Clausoca-
8a In lateral view, ventral profile of Gns w ithout lanus arcuicomis (Fig. 7.72)
prominent step posterior to genital pores ......... .9 3a Pr more than 5.7 times as long as U r2 .... ...... ... .4
9 In lateral view, dorsal lobe of seminal receptac le 4 Left PS segment 5 armature includes 2 thick,
arising from anterior edge of ventral lobe; dorsal curved spiniform setae; PS segment 2 length
lobe of seminal receptacle not bulb-shaped: more than 1.45 times as great as width of Ur2:
Clausocalanus arcuicornis (Dana, 1849) (Fig. Clausocalanus jobei (Fig. 7. 76)
7.72)
Copepoda 917
~cle Left P5 segment 5 armature consists of slender, Family Aetideidae Giesbrecht, 1892
usually straight setae; P5 segment 2 length less Reference: Markhaseva ( 1996)
~d than 1.45 times as great as width of Ur2: Females
____ IQ
Clausocalanus minor (Fig. 7.80) Pd4 and 5 fused in majority of genera. R, if present, 2-
.2 lobe Longer leg of P5 much shorter than Ur: pointed, or in form of blunt plate, or conical and !-
!POD of Clausocalanus brevipes (Fig. 7.73) pointed. Posterior corners of Pr symmetrical or asym-
:roe. In Longer leg of P5 longer than Ur ....................... 6 metri cal, rounded or pointed, sometimes extended into
Ded.. .-\.1 R in lateral view, not knob-like and not protru- spines of various size and configuration or in lobes of
:.:5 seg- ding ventrally. Longer leg ofP5 usually on right various shapes. Gns symmetrical or asymmetrical,
:.:5 long side: Clausocalanusfurcatus (Fig. 7.74) sometimes with 1-2 spines; ventral swelling more or
rlleT!!ens R in lateral view, knob-like and protruding vent- less pronounced. Al 23-24-segmented, of various
rally; longer leg of P5 on left side ..................... 7 lengths. A2 Exp 7-segmented; Expl with 1-2 setae or
~lobe Total length less than 0.80 mm: Clausocalanus without setae; sometimes Exp l and Exp2 (Euchirella)
rlmtent pergens (Fig. 7.83) fused completely or incompletely, or Exp2 and Exp3
:ate raJ Total length greater than 0.80 mm .................... 8 (Comantenna , Mesocomantenna) pattly fused; Exp2
r::.gular. P3 B wi th spiniform processes 1-3 slender and with 1-3 setae, or setae absent ( Chiridiella,
~gas rapering uniformly from base. CR more than 1.6 Comantenna, Euchire/la). Mdp B with 2-3 setae;
I:IlleS as rimes as long as wide: Clausoca/anus laticeps sometimes with I seta, or without setae; Enp I w ith 1-
______ } !
F ig. 7.77) 3 or without setae; Enp2 with 9 terminal and 1-2 pos-
~.AI P3 B with spinifonn processes 2-3 usually stout terior setae, sometimes this number differs; Exp 5-seg-
and not tapering unifonnly from base. CR less mented, Expl-4 with I long seta each; Exp5 with 2
than 1.6 times as long as wide ..........................9 term inal setae. Mx I Li I with 9 terminal setae, I shor-
P: segment 2 less than 4.4 times as long as wide: ter anterior seta and 4 setae on posterior surface (some-
..-.: seg- Clausocalanus ingens (Fig. 7.75) times number of posterior setae is fewer); Li2 with 3-
~em P5 segment 2 more than 4.4 times as long as 5 setae, rarely with 2 or 1 seta in Chiridiella this lobe
::- and wide ................................ ....... .......................... 10 may be reduced; Li3 with 3-4 setae; Enp with 14-16
t:r2 more than 1.35 times as long as segment 2 setae, or fewer: 3-5 or 7 in Chiridiella, I 0 or 11 in
of P5: C/ausoca/anus lividus (Fig. 7 .78) Paivella and 3 or 5 in Euchirella; Exp in most genera
l:r2 less than 1.35 times as long as segment 2 of with II, rarely with 8- 10 setae, but in Chiridiella this
~ irhin P5: Clausocalanus mastigophorus (Fig. 7.79);? number is 3-6 and Exp may be reduced. Mx2 usually
•.!! Pr. Clausoca/anusparapergens (Fig. 7.81) with 5 well-developed Li (6 Li in Jaschnovia,
~ of irs Senecella and some species of Aetideopsis), Enp with
. Pd1 • to species of Ctenocalanus 6 long setae, sometimes with additional small setae;
---···2 - MLales Lil-5 with 2 long and I short plumose seta; one of two
tt..~g .-\ I reaching posterior border of CR. P4 Exp3 long setae on LiS (rarely also on Li4) thickened and
il_,c ,rh outer spines not expanded at base: Ctenocalanus transfonned into claw-like spine. Two genera have
~g citer Heron and Bowman, 1971 (Fig. 7.84) Mx2 strongly deviating from typical structure: in
r::ri~J.!a- .-\1 exceeding CR by several segments. P4 Exp3 Pseudeuchaeta setae on Li4 and Li5 and on distal part
---····5 oute r spines robust, expanded at base: of Enp are more sclerotized and sabre-like; and in
awUcnUS Crenocalanus vanus Giesbrecht, 1888 (Fig. Chiridiella Mx2 highly transformed. Mxp C with 1
- .85) proximal seta (sometimes absent), with groups of 2, 3
...
---···-' and 3 setae from proximal to distal. P I Enp outer lobe
JDZSOCa - well developed, with small spinules terminally, rarely
Right P5 present, !-segmented: Ctenocalanus with hairs instead of spinules (Pseudeuchaeta); outer
_ ..... A cirer (Fig. 7.84) lobe absent in Chiridiella (except C. kuniae). Posterior
.:: ::rick. ~ Right P5 absent: Ctenocalanus vanus (Fig. surfaces of Enp and Exp sometimes with fine small
! :~gt.h - _ 5) spinules, larger spinules on lateral borders of C and B
--
c: ....-r_:
")
(Bradyidius, Jaschnovia). Majority of genera without
spines or spinules on P4 C, but sometimes with spines:
918 Copepoda
2 parallel transverse rows of 4-6 spines (Paivella); or The re are 26 genera in the family: Aetideus Brady,
bush-like spines near base of inner seta (Gaetanus, 1883; Aetideopsis Sars, 1903; Azygokeras Koeller and
Pseudochirella, Euchirella), or only few small spinu- L ittlepage, 1976; Batheuchaeta Brodsky, 1950;
les (some species of Aetideus and Undeuchaeta). Bradyidius Giesbrecht, 1897; Chiridiella Sars, 1907;
Typical spine and seta formula: Chiridius Giesbrecht, 1892; Chirundina Giesbrecht,
1895; Chirundinella Tanaka, 1957; Comantenna
coxa basis exopod segments en do pod Wi lson, 1924; Crassantenna Cole et al., 1972;
segments Euchirella G iesbrecht, 1888; Gaelanus Giesbrecht,
Leg I 0-0 0-1 /0 0'1-0; 1-1 ; 1,1,3 0,2,3
1888; Jaschnovia Markhaseva, 1980; Lutamator
TI-l; 1,1,3 (Euchirella) Bradford, 1969; Mesocomantenna Alvarez, 1986;
0-0 (Chiridiella) ]/]1,1,4 (Chiridiella) Paivella Vervoort, 1965; Paracomantenna Campaner,
l-1; 1,1,3 (Gaetanus part.) 1978; Pseudeuchaeta Sars, 1905; Pseudochirella Sars,
Leg2 0-1 0-0 1-1 ; 1-1: lll,l,4 0-1 ; 1,2,2
1920; Pterochirella Schulz, I 990; Senecella Juday,
or 1,2,3
Leg 3 0-1 0-0 1-1 ; 1-1; 111,1,4 0-1; 0-1;
1923; Sursamucro Bradford, 1969; Undeuchaeta
1,2,2 G iesbrecht, 1888 and poorly described Gaidiopsis A.
Leg 4 0- 110 0-0 1-1; l-1 ; lll,1,4 0-1; 0-1; Scott, 1909. The Aetideidae were recently reviewed
1,2,2 by Markhaseva (1996) and all keys are adapted from
this review. Sixteen of these genera are known to
P5 absent (except for weak P5 in Comantenna recur- occur in the South Atla ntic an d are given below.
vata and vestigial P5 found in some specimens of
Sursamucro and Aetideopsis). Key to genera and some species of Aetideidae
Females
M ales 1 Posterior somite of Pr and Gns symmetrical. ... .2
Pd4 and 5 usually fused, sometimes incompletely 1a Posterior somite of Pr or Gns, or both, more or
(Aetideopsis, Azygokeras). R, if present, two-pointed, less symmetrical. ......... .. ................................ .. I 6
or coni cal (1-pointed). Crest may be present or absent. 2 R present, not reduced ..... ... ........... ... ................. 3
Posterior corners of Pr symmetrical, rarely somewhat 2a R absent or reduced to blunt plate ................... I I
asymmetrical (some Pseudochirella and Undeuchaeta 3 R 2-pointed .............. ....... .................................. .4
sp ecies), of various shape. Ur of 5 free somites; U r5 3a R !-pointed, coni cal ........ .... ........... ...... ........... .. ?
sharpl y reduced; genital opening on left. A I usually 4 P 1 Exp l without outer spine. Mx1 Li2 with 3,
23-24-segmented, but sometimes 20-21-segmented Li4 with 4 setae, Exp with 11 setae .................. 5
(Aetideus); in all genera, except Azygokeras, not 4a P1 Exp l with outer spine. Mx1 L i2 with 5 (rare-
genicul ated; with aestetasks in most species. Left and ly 4) setae, Li4 with 5 setae, Exp with 8-10 setae
right A 1 may have d ifferent number of segments. A2 .......................... ................................................. 6
as in female, but with fewer setae on Exp 1, Exp2 and 5 P4 C near base of inner seta with 2 parallel rows
Enp; some setae of Mdp reduced in size. Mx1 and of 4-6 spines on posterior surface: Paivella
Mx2 very reduced . Mxp less well equipped w ith setae 5a P4 C w ithout 2 parallel rows of 4 -6 spines on
than in females. P 1-P4 segmentation more pro- posterior surface: Aetideus
nounced than in females (i.e. when female has indis- 6 Pl Exp2 outer spine thicker than on Exp1, usu-
tinct division, male has complete division). P 1 Exp ally more sclerotized and more obtuse. P2 Enp
spines and setae as in female, but sometimes external always 2-segmented: Bradyidius plinoi
spines reduced, occasionally absent. P4 C without Campaner, 1978 (Fig. 7.96)
spines. Right and left P5 usually present, sometimes 6a P 1 Exp2 outer spine not thicker than on Exp 1
right P5 absent (Aetideus). P5 biramous or uniramous. and not differing from it in its sclerotization and
lf biramous, with Exp 3-segmented on left and 2-seg- shape. P2 Enp 1- or not clearly 2-segmented:
mented on right (in Azygokeras, Chiridiella and Aetideopsis
Pseudeuchaeta right Exp 3-segmented). Both left and 7 P4 C wi thout spines near inner seta .................. 8
right Enp !-segme nted (left Enp 2-segmented in 7a P4 C with spines near inner seta .... .......... ....... .. 9
Bradyetes matthei, Bradyidius arnoldi and Coman- 8 Mxp B with lateral blade along its outer border.
tenna brevicornis). Mx l Li2 with 5, L i3 with 4 setae, Li4 with 5,
Copepoda 919
3rady. Enp with 15, and Exp with II setae: Chinmdina l6a P4 C without spines near inner seta. P1 Exp with
~crand streetsii Giesbrecht, 1895 (Fig. 7.101) 2 outer spines: Undeuchaeta
: 9 50: Sa Mxp B without lateral blade along its external 17 A2 Enp less than half, usually 1/3 or Y.. length of
5.. :90- : border. Mx I Li2 with 4, Li3 with 5, Li4 w ith 4 , Exp; Expl and Exp2 completely or incomplete-
s.Xecbt. Enp with II, Exp with LO setae: Chirundine/la ly fused. Mdp B and Enp Iwithout seta (rarely
.;enna magna (Wolfenden, 19 11) (Fig. 7. 102) w ith small seta). Mx 1 Enp with 3 or 5 setae. P 1
. I9- 2: 9 A2 Enp less than half length of Exp. Mx I Enp Exp 2-segmente d: E uchirella (part.)
s.Xecbt. with no more than 5 setae: Euchirella (part.) 17a A2 Enp more than half length ofExp; Exp1 and
~ator 9a A2 Enp more than half length of Exp. Mx 1 Enp Exp2 completely sep arated. Mdp B with 3, Enpl
. ~986: with no less than 14 setae ... .. ..... .... .................. I 0 with 2-3 setae. Mx 1 Enp with 15-16 setae. P 1
r:::;paner, 10 Pd4 and 5 separated (except P. hirsuta). Mdp B Exp 3-segmented: Pseudochirella (part.)
C:7 Sars. with 3 setae. Anterior part of head without fron-
Juday. tal spine. Mxp C without lateral p late. P1 Exp Males (unknown for Lutamator, Mesocomantenna,
rna era with 3 outer spines: Pseudochirella (part.) Paracomantenna, Bradyidius p linoi, Chiridiella atlan-
r.:ru A. lOa Pd4 and 5 fused. Mdp B with 1-2 setae. Anterior tica and Comantenna curtisetosa)
!"\-:.ewed part of head often with frontal spine. Mxp C 1 R present .............. ....... ........ ............................... 2
eci from often with lateral plate. PI Exp with 2 or 3 l a Rabsent ........................................................... J4
lJ'IInl to external spines: Gaetanus 2 R 2-pointed ........................................................ 3
I~ Mxp C without finger-like appendage near d istal 2a R !-pointed ........................................................ 5
group of 3 setae ............................................... 12 3 Posterior comers of Pr rounded: Paivella ina-
lla Mxp C with fmger-like appendage near distal ciae Vervoort, 1965 (Fig. 7. 126)
group of 3 setae ......... ...... ..... .... ....................... 14 3a Posterior comers of Pr pointed ............ .. .. ......... .4
~..... 2 12 PI Exp !-segmented. Mx2 with moctified distal 4 P5 uniramous : Bradyidius (part.)
c.oce or setae. Mx l Enp with less than 7 setae. P 1 Enp 4a P5 biramous: Aetideopsis, Bradyidius (part.).
_
- ... 16
......).,
- .... 11 Ca
witho ut outer lobe: Chiridiella atlantica
Wolfenden, 1911 (Fig. 7.97)
P 1 Exp 3-segmented. Mx2 of typical structure.
5 Anterio r part of head with fron ta l spine:
Gaetanus (part.)
5a Anterior part of head without frontal spine ....... 6
_ .... A Mx1 Enp w ith 11 - 16 setae. P 1 Enp with an outer 6 Right P5 Exp2 and Enp in form of chela:
lobe ..... .......... ................. ......... ."........................ 13 Euchirella (part.)
lrim 3. 1: Posterior comers of Pr with spines directed 6a Right P5 Exp2 and Enp not in form of chela .... 7
___ .... 5 straight backward. R absent. Md Enp2 with 8-9 7 Anterior Ce with crest ............................. .. ........ 8
5 rare- terminal (sometimes with 7 termi nal) and I pos- 7a Anterior Ce without crest ................................ 1 I
C s.erae terior seta; Mdp Enp I and B with 2 setae each: 8 PI Exp 2-segmented : Euchirella (part.)
-·····6 Chiridius 8a PI Exp 3-segmented ...... .................................... 9
lcl rows 1:3 Posterior comers of Pr without spines (or 9 Left P5 Exp3 bilobated: Chirundinella magna
points), triangular or slightl y pointed (in lateral (Fig. 7. 102)
c::es on ,·iew). R present, reduced, in form of blunt plate. 9a Left P5 Exp3 non bilobated ............ ...... ........... l 0
~dp Enp2 with 4-5 setae; Enpl and B without 10 Left P5 Exp3 distally stylet-like, elongated:
tL t:.:>-u- setae: Lutamator elegans Alvarez, 1984 (Fig. Undeuchaeta
1?: Enp - .124) 1Oa Left P5 Exp3 distally not stylet-like, obtuse-
plinoi .! :\2 Exp terminal segment not longer tha n 2 pre- triangular: Chirundina streetsii (Fig. 7. J 01 )
,·ious segments together ... ... ........... .. .. ............. 15 11 Left P5 without teeth on Exp2, segments of
~::!. •.U Exp tem1inal segment much longer than 2 si mple configuration: Gaetanus (part.)
previous segments together: Paracomantemza 11a Left P5 with teeth on Exp2, or segments of com-
15 ..U Exp tem1inal segment w ith 3 setae: Com- plex configuration ............... ..... ....... ................. l2
aJrtenna curtisetosa A lvarez, 1986 (Fig. 7.103) 12 PI Exp 2-segmented: Euclzirella (part.)
--····8 5.J. ..U Exp terminal segment with I seta: Meso- 12a P 1 Exp 3-segmented ........................................ 13
-···9 comantenna spinosa Alvarez, 1986 (Fig. 7.125) 13 P l Exp 1 with outer spine: Pseudochirella
croer. P-l C with spines near inner seta. P 1 Exp w ith 3 l3a PI Exp I without outer spine: Undeuchaeta
~ 5. spines ..... ............................................... ........... 17 (part.)
920 Copepoda
14 P5 with left leg only, uniramous: Aetideus Key to species of Aetitleopsis

14a P5 with both left and right legs, uni- or biramous Females
......... ....................................................... ......... 15 1 Posterior corners ofPr wing-like, divergent ( dor-
15 P5 uniramous, left P5 5-segmented, right 4-seg- sal view). Gns (dorsal view) with lateral swel-
mented: Chiridius lings: Aetideopsis carinata Bradford, 1969 (Fig.
15a P5 biramous .................... ................................. 16 7.93)
16 Right P5 Exp 3-segmented: Chiridiella 1a Posterior com ers of Pr not wing-like, not diver-
16a Right P5 Exp 2-segmented: Comantenna gent (dorsal view). Gns (dorsal view) barrel-like
without lateral swellings .......... .. ........ .......... .. ... .2
Key to the species of Aetideus 2 Proxin1al outer part of Mxp C with plainly visi-
Females ble projection: Aetideopsis rostrata Sars, 1903
1 Posterior comers of Pr prolonged into wing-like (Fig. 7.95)
lobes pointed terminally, reaching at least mid- 2a Proximal outer part of Mxp C without projec-
dle of Ur2. Anterior part of head with crest... .. .2 tion: Aetideopsis minor (Wolfenden, 1911) (Fig.
Ia Posterior comers of Pr pointed, not prolonged 7.94)
into wing-like lobes, just reaching or exceeding
posterior border of Gns. Anterior part of head Males (unknown for A. minor).
\vithout crest .............. ................... ................... ..3 1 Rostral points more or less divergent. Total
2 Total length 1.8-2.2 mm. Duct between ventral length over 3 mm: Aetideopsis rostrata (Fig.
and dorsa l part of spermatheca strongly narro- 7.95)
wed. Base of R not visible in dorsal view: l a Rostral points widely spaced, divergent. Total
Aetideus giesbrechti (Cleve, 1904) (Fig. 7.91) length about 2.4 mm: Aetideopsis carinata (Fig.
2a Total length 1.48-1.80 mm. Duct between ventr- 7.93)
al and dorsal parts of spermatheca not narrowed.
Base of R visible in dorsal view: Aetideus acu- Key to species of Chiridius
tus Farran, 1929 (Fig. 7.88) Females
3 Gns widest at midlength (dorsal view): Aetideus 1 Mx I Exp with 8 setae. Posterior corners of Pr
australis (Vervoort, 1957) (Fig. 7.90) with points reaching at least midlength of Gns.
3a Gns widest anterior to midlength .......... ...... ..... .4 Specimens 1. 59-2.80 mm in length. Mdp Enp2
4 Spermatheca with short duct connecting its dor- commonly with 9 setae ... ................................... 3
sal and ventral parts: Aetideus armatus 1a \tlx I Exp with I 0 setae. Posterior comers of Pr
(Boeck,l872) (Fig. 7.89) with points reaching 113 of way along Gns, rare-
4a Sperrnatheca with long duct connecti ng its dor- ly reaching, or even exceeding its middle. Mdp
sal and ventral patts: Aetideus psemlarmatus Enp2 with 8 setae. Specimens 3.30-4.25 mm in
Bradford, 1971 (Fig. 7.92) length: Chiridius polaris Wolfenden , 1911 (Fig.
7.99)
Males (unknown for A. pseudarmatus) 2 Specimens 1.59-2.20 mm in length. Pr 2.7-3.2
I Points of posterior comers of Pr not exceeding times longer than Ur. P 1 Exp I , 2 and 3 outer
posterior border ofUrl (in dorsal view) .......... .2 spines not reaching base of next spine. P2 Enp !-
I a Points of posterior corners of Pr exceeding pos- segmented, exceeding P2 Exp2-3 border:
terior border of Ur i (in dorsal view) .. ...... ......... 3 Chiridius poppei Giesbrecht, 1892 (Fig. 7.1 00)
2 Second P5 segment 3.35-4.0 times as long as 2a Specimens 2.45-2.80 mm in length. Pr 3.2-4.2
wide: Aetideus giesbrechti (Fig. 7.91) times longer than Ur. Pl Expl and 2 outer spines
2a Second P5 segment 2.3-3.1 times as long as reaching or exceeding base of next spine. P2 Enp
wide: Aetideus armatus (Fig. 7.89) 2-segmented (sometimes separation is obscure),
3 CR 1.64-1 .83 times as long as wide: Aetideus mostly not reaching border of P2 Exp2-3:
acutus (Fig. 7.88) Chiridius gracilis Farran, 1908 (Fig. 7.98)
3a CR are about 3 times as long as wide: Aetideus
australis (Fig. 7.90)
Copepoda 921
Wes 8 Left and right sides ofGns (dorsal view) without

Points of Pr posterior comers not longer than swellings. Projection on dorsal surface of Gns
::<dor- Gns. Right P5 nearly as long as left. Pl Exp l and very large, exceeding posterior border of Gns
5"\·el- ~ outer spines very short, not reaching middle of and covering 2 following Ur somites. Md Enpl
59 Fig. next joint: Chiridius poppei (Fig. 7 .I 00) with 1 small seta: Euchirella messinensis mes-
2. Points of Pr posterior corners usually reaching sinensis (Claus, 1863) (Fig.7.109)
- CiYer- posterior border of Gns .................. .... ............... 2 8a Left side of Gns (dorsal view) with swelling, or
~~-like P 1 Exp 1 outer spine reaching and exceeding base bilobated, or non-lobated projection. Md Enpl
-······2 of following spine: Chiridius polaris (Fig. 7.99) without seta ............................... .. ...................... .9
ll!- \isi- P1 Exp 1 outer spine not reaching base offollow- 9 Posterodorsal surface of Gns with 3 projections
,._ 1903 ing spine: Chiridius gracilis (Fig. 7.98) (left lateral view) A2 Enp2 lobes with less than
6 setae each: Euchirella formosa Vervoort,
trojec- r to species of Euchirella
_. I 949 (Fig. 7.I 07)
1 Fig. f:ana.les 9a Posterior pmt of Gns without projections (left
I Gns symmetrical.. ................ .............................. 2 lateral view). A2 Enp2 lobes with not less than 6
a Gns asymmetrical ................................. ..... ...... ..4 setae each: Euchirella pulchra (Lubbock, 1856)
Crest present. R rudimentary: Euchirella curti- (Fig. 7. 110)
. Total cauda Giesbrecht, 1888 (Fig. 7.106) 10 Projection on Gns ear-like, on anterior half:
• Fig. Crest absent. R not rudimentary ................ .... .... 3 Euchirella simi/is Wolfenden, 1911 (Fig. 7.112)
.\fdp with chitinous projection. P4 C with 5 very I Oa Projection on Gns blunt, triangular, on its poste-
'- ~oral poorly visible small spines. A2 Enp2 with 3 rior half: Euchirella splendens Vervoort, 1963
Fig. setae on outer lobe and 1 seta on inner lobe. R (Fig. 7.113)
small: Euchirella amoena Giesbrecht, 1888
Fig. 7.104) Males (unknown for E. simi/is)
.\Idp without chitinous projection. P4 C with 8- 1 P5 always biramous. Left P5 Enp well-devel-
9 clearly recognizable spines. A2 Enp2 with 6 oped, nearly as long as P5 Exp I , or equal to half
~ oi Pr setae on outer and 8 setae on itmer lobes. R its length; right Exp2 and Enp not elongate and
o.f Gns. large: Euchirella rostrata (Claus, 1866) (Fig. not sharpened distally; not forming obvious
f Enp2 - .Il l ) grasping organ ....... ..... ............. ................ ...... .... 2
___ , ,j
., Crest present ... ... ................... ....... .......... ... .... ..... 5 1a P5 not always biramous, if left P5 with Enp, then
-s of Pr Crest absent ......... ................... ................. ... .. ..... 6 Enp rudimentary; right Exp2 and Enp elongate
& rare- Posterior corners of Pr slightly asymmetrical, not and sharpened distally forming grasping organ
~- .\hlp rounded (dorsal and lateral view). A2 Enp with 5 ........................................................ ..... .. .... ....... .4
u:::n ill setae on each (inner and outer) lobe: Euchirella 2 Crest absent. R large: Euchirella rostrata (Fig.
! Fig. maxima Wolfenden, 1905 (Fig. 7.1 08) 7.111 )
~ Posterior corners ofPr symmetrical, rounded, or 2a Crest present. R not large .................. ................ 3
- - - -..,j __")
- rounded-rectangular (dorsal and lateral view). 3 Terminal part of right P5 Exp2 with teeth:
~ cmer .-\2 Enp2 with 6 setae on each lobe: Euchirella Euchirella curticauda (Fig. 7.106)
. L--pi - bitumida W ith, 191 5 (Fig. 7.105) 3a Terminal part of right P5 Exp2 without teeth:
JOroer: Gns with left lateral swelled (dorsal view), or Euchirella maxima
\\ith large projection on left half of somite dor- 4 Crest present ...................................................... 5
sally; if right projection also present, it is signif- 4a Crest absent ....................................................... 8
· .::-:nes icantly smaller than left projection (dorsal view) 5 Crest high: Euchirella bitumida (Fig. 7.1 05)
?.::Enp ............. ................... ........................................... 7 5a Crest low ............................................................ 6
~:...rre). Gns with large projection on rig ht side (dorsal 6 Left P5 8 shorter than projecting pa1t of right P5
:xp:-3: \iew). left side without swelling or projection C........................................................ .................7
............ .................. ..................... ................. ... .. 10 6a Left P5 8 nearly as long as projecting part of
P-1 C with 1 spine : Euchirella truncata Esterly, right P5 C: Euchirella formosa (Fig. 7 .I 0 7)
191 1 (Fig. 7. 114) 7 Right P5 Exp l with less than 4 tooth-like projec-
-~ P~ C \Vith 2 spines ............................................. 8 tions: Euchirella pulchra (Fig. 7.11 0)
922 Copepoda
7a Right P5 Expl with 4 tooth-like projections: Males (unknown for G. minor)

Euchirella messinensis messinensis (Fig. 7.1 09) 1 Head with frontal spine ...... .... ................. ..........2
8 Robust spine present on right P5 B at base of 1a Head without frontal spine .................. ...... .... ... .4
Exp: Euchirella amoena (Fig. 7. 104) 2 Left P5 Exp3 bilobate. Spines on posterior cor-
8a No spine present on right P5 B at base of Exp ners of Pr short, not reaching posterior border of
................ ...... .... .............................. ...... ..... .... ....9 Gns: Gaetanus antarcticus (Fig. 7.115)
9 Left P5 without rudimentary Enp; Exp chela, 2a Left P5 Exp3 prolonged, stylet-like, never bilo-
when closed, compact, rounded, club-like: bate. Spines on posterior comers of Pr long,
Euchirella truncata (Fig. 7. 114) exceeding posterior border of Gns .................... 3
9a Left P5 with rudimentary Enp; Exp chela, when 3 Left P5 Enp longer than half length of left P5
closed, compact, oblong, not club-like: Exp1 , or even longer: Gaetanus kruppii (Fig.
E uchirel/a splendens (Fig. 7 .1 13) 7.118)
3a Left P5 Enp shorter than half length of left P5
Key to species of Gaetanus Expl: Gaetanus pileatus, G. brevicornis (detail-
Females ed description and figures of G. brevicornis are
I Anterior head with frontal spine ........... ............ 2 not available, therefore convincing differences
Ia Anterior head without frontal spine ........... ... .... 7 between males of these species could not be
2 Mx1 Li2 with 5 setae. Pl Exp with 3 outer obtained) (Fig. 7 .116)
spines: Gaetanus antarcticus Wolfenden, 1905 4 Left P5 Exp3 bilobate. Specimens about 7 mm in
(Fig. 7.115) size: Gaetat~us robustus (Fig. 7 .122)
2a Mx l Li2 with 4 setae. P1 Exp always with 2 4a Left P5 Exp3 slylet-like. Specimens not more
outer spines ............................ .... ........................ 3 than 4 mm in length ...... .... ................................ .5
3 A2 Exp2 with 1 seta ........... ...... ............... ......... .4 5 Posterior comers ofPr without spines: Gaetanus
3a A2 Exp2 with 2 setae ... ....................... ...... ........ 5 miles (Fig. 7.11 9)
4 F rontal spine straight, not curved towards R dis- Sa Posterior comers of Pr with spines ................... 6
tally: Gaetanus pileatus Farran, 1903 (Fig. 6 Spines on posterior corners of Pr exceeding pos-
7.12 1) terior border ofGns: Gaetanus tenuispinus (Fig.
4a Frontal spine curved towards R di stally: 7. 123)
Gaetanus miles Giesbrecht, 1888 (Fig. 7.119) 6a Spines on posterior corners of Pr significantl y
5 Spines of Pr posterior comers curved dorsally shorter: Gaetanus brevispinus (Fig. 7. 117)
(lateral view): Gaetamts brevicomis Esterly,
1906(Fig. 7.116) Key to species of Paivella
Sa Spines of Pr posterior comers directed straight Females
backwards (lateral view) ....... .................. .......... 6 1 Gns without lateral projections: Paivella inaciae
6 P 1 Exp 2-segrnented; P2 Enp !-segmented. Vervoort, 1965 (Fig. 7. 126)
Specimens Jess than 3 mm in size ( 1. 70-2.40 Ia Gns with lateral projections: Paivella naporai
mm): Gaetanus minor Farran, 1905 (Fig. 7.120) Wheeler, 1970 (males unknown) (Fig. 7.127)
6a P 1 Exp 3-jointed; P2 Enp 2-segmented.
Specimens more than 4.5 mm in size: Gaetanus Key to species of Paracomantenna
kruppii Giesbrecht, 1903 (Fig. 7 .118) Females (males unknown)
7 A2 Exp1 with I seta. Mx1 Li2 with 5 setae. P1 I Mdp B with 1 small seta; Enp2 with 5 setae.
Exp 3-segrnented. Exp P1 with 3 outer spines: Mxp C distally with tube-like appendage: Para-
Gaetanus robustus Sars, 1905 (Fig. 7.122) comantenna gracilis Alvarez, 1986 (Fig. 7.1 28)
7a A2 Exp1 without setae. A2 Exp2 with 2 setae. Ia Mdp B without seta; Enp2 with 4 setae. Mxp C
Mx l Li2 with 4 setae. Exp P1 with 2 outer di stally with finger-like appendage: Para-
spines ...... ................... ................ ....................... 8 comantenna magulyae Campaner, 1978 (Fig.
8 Mxp C with lateral plate: Gaettmus brevispimts 7. 129)
(Sars, 1900) (Fig.7 .117)
8a Mxp C without lateral plate: Gaetanus tenuispi-
nus (Sar s, 1900) (Fig. 7 .123)
Copepoda 923
to species of Pseudochirella 4 Left P5 Exp2 with 3 teeth distally. Posterior cor-

-······2 '=""ftllales ners of Pr without spines: Pseudochirella dubia
___ __..1
I Gns symmetrical .......... .. ........................... ....... ..2 4a Left P5 Exp2 with 2 teeth distally .... ......... .... .... 5
nor- cor- Gns asymmetrical ................ ...... ................ ........ 6 5 Posterior corners of Pr with spines ................... 6
o:rier of :\2 Exp2 with 3 setae ......................... ............... 3 Sa Posterior comers of Pr without spines: Pseudo-
:Z 6.U Exp2 with 1 seta .................................. ........4 chirella pustulifera
F-'r" bilo- Pd4 a nd 5 distinctly separated. Ventral swelling 6 Left P5 Exp3 2 times longer, than wide: Pseudo-
"J;- !ong. o f Gns strongly protruding: Pseudochirella chirel/a mawsoni (Fig. 7.133)
..,
-·····.) dubia (Sars, 1905) (Fig. 7.I31) 6a Left PS Exp3 more than 3 times longer than
'lef. P5 .:.a Pd4 and 5 indistinctly separated. Ventral swe ll- wide: Pseudochirella obtusa (Fig. 7.134)
rii Fig . ing of Gns slightly protruding: Pseudochirella
hirsuta (Wolfenden, 1905) (Fig. 7. I 32) Key to species of Undeuchaeta
' left P5 Posterior comers of Pr with spines reaching Females
derail- about midlength ofGns: Pseudochirella spinosa 1 Head with crest. Gns with spine on genital fie ld
n::; are Wolfenden, 191 I) (Fig. 7.137) ............. ... ................... ....................................... .2
mnces Posterior com ers of Pr without spines, rounded, 1a Head without crest. Gns without spine on genital
~tbe or prolonged into triangular, or rounded lobes .5 field (late ra l view) : Undeuchaeta plumosa
Posterior comers of Pr protruding into rounded (Lubbock, 1856) (Fig. 7 .140)
wmin lobes: Pseudochirella obtusa (Sars, 1905) (Fig. 2 Posterior corners ofPr nearly equally prolonged,
- . 134) triangular (lateral and dorsal views):
more Posterior comers of Pr protruding into triangular Undeuchaeta major Giesbrecht, 1888 (Fig.
-- ..... 5 lobes : Pseudochirella batillipa Park, 1978 (Fig. 7 .139)
ll!UIIUS - .130) 2a Posterior corners of Pr not equally prolonged,
Right posterior comer ofPr with spine, left with- one of the m shorter and rounded (lateral view):
.... 6 out spines: Pseudochire/la spectabilis (Sars, Undeuchaeta incisa Esterl y, 19 11 (Fig. 7.138)
~pos 1900) (Fig. 7.136)
Fig . Both left and right posterior comers of Pr with- Males
out spines, symmetrical ................. .................... ? 1 Head without crest: Undeuchaeta plumosa (Fig.
i~tly
-, Projection present on right posterolateral part of
Gns: Pseudochirella mawsoni Vervoort, 1957
7.140)
I a Head with c rest ..... ............. ............ ............... .....2
Fig. 7. 133) 2 Left P5 Exp2 twi ce, or more, longer than wide :
Projection present on right anterolateral patt of Undeucheta incisa (Fig. 7.138)
Gns: Pseudochirel/a pustulifera (Sars, 1905) 2a Left P5 Exp2 1.5, or less, longer than wide:
Fig. 7 .135) Undeuclweta major (Fig. 7.139)
rai bles (unknown for P. spinosa) Family Euchaetidae Giesbrecht, 1892

Left P5 Exp2 without tooth, or with 1 tooth on References: Bradford eta/. (1983), Park (1995)
Its distal part ........... .. ................... ... ........... ........ 2 Females
Left P5 Exp2 with 2-3 teeth on distal part of seg- Pd1 separated from Ce by faint line representing par-
ment ... .................... ........................................... .4 tially fused joint, Pd4 and 5 completely fused. R sin-
Posterio r corners of Pr with small teeth on right gle, spiniform. Supralabrurn highly prominent with
.PIITa- and left: Pseudochirella spectabilis (Fig. 7 .136) rows of long stiff hairs, lies anterior to labrum. Ur of
-_1 .:-) Posterior comers of Pr without teeth, rounded 4 somites. Gns symmetrical, or asymmetrical.
xpC ............... ............ .... ... ......................................... 3 Appendicular caudal setae highly developed, straight,
PIITil - Distal part of right P5 Exp blade-like: smooth ly curved, or geniculated, and usually longer
Fig. Pseudoclzirella batillipa (Fig. 7.130) than marginal caudal setae. A l with extremely long
Distal part of right P5 Exp not blade-l ike: seta on each of segments 3, 7, 9, 14, 18, 21 and 24;
Pseudoclzire/la hirsuta (Fig . 7 .132) segments 8 and 9 fused. A2 C with 1 seta, B with 2
setae, Enpl with 2 setae and Enp2 with 8 setae on
inner and 6 setae on outer lobe; Exp 7-segmented,
924 Copepoda
Exp1lacking setae, Exp2 with 1 (or lacking seta) mar- R; the length of setae on A1 (setae on corresponding
ginal seta. Md Enpl with 1 seta, Enp2 with 9 terminal segments shorter), A2 (C without seta), Mx I Li2-Li3
and I appendicular seta (sometimes this seta absent). with 2-3 setae; Mx2 Li5 with I of setae stronger than
Mx1 Lil with 9 tenninal setae, 1 anterior seta and others, spine-like; also subdivision of Exp and Enp
from 1 to 3 posterior setae, Li2 has 2 setae (sometimes segments of P2-P4 may be incomplete. P5 of males
1 seta, or both setae absent), Li3 with 1 seta, Enp 1 similar to these of Undeuchaeta in the Aetideidac; ser-
(consisting of 2 fused segments) has 1-7 setae, Enp2 rated lamella and digitiform process absent in left P5
with 3 setae, Exp with II setae and Lel with 5-9 Exp2. Until a complete revision of Valdiviella is made
setae. Mx2 very strongly built; Li1-Li5 with 1 short the genus is placed temporarily in the Euchaetidae.
and 2 long setae each: terminal setae of Li 1-3 or Li 1-
4 armed with long spinules in addition to dense rows Key to genera of Euchaetidae
of short spinules; terminal part of Mx2 with 6 long Females
setae. Mxp very strongly built; C with 1 seta (this seta I R 2-pointed. P2-P3 Exp incompletely 3-seg-
may be absent), then with groups of 2, 3 and 3 setae mented, or 2-segmented: Valdiviella
from proximal to distal; Enp2-6 with 4, 3, 2, 3 and 4 I a R single, !-pointed. P2-P3 Exp3-segmentedoooo .2
setae; medial edge of B armed either with short spi- 2 Appendicular caudal seta greatly developed,
nules only, or with both short and long spinules. PI straight, and much thicker and longer than distal
End with usually well-developed outer lobe; two first marginal setae of CR: E uchaeta
P 1 Exp segments usually unseparated, Exp 1 with 2a Appendicular caudal setae either geniculated or
outer spine that is usually very small or missing, Exp2 smoothly curved, are relatively small , and dis-
with outer spine and Exp3 with outer spine. Spine and ti nctly thinner but usually longer than principal
seta formula: marginal setae of CR: Paraeuchaeta
coxa basis cxopod segments endopod segments Males

1 R 2-pointed. Left P5 Exp2 without sl and dp:
Leg I 0-0 0- 1 I-I ; 1,1,3 0,2,3
Leg 2 0- 1 0-0 T-1 ; 1- 1; fll,l,4 1,2,3
Vahliviella
Leg 3 0- 1 0-0 1- 1; 1-1 ; 111,1,4 0-1 ; 0- 1; I ,2,2 I a R single, 1-pointed. Left P5 Exp2 with sl and dp
Leg4 0- 1 0-0 T- 1; T-1; JII,I,4 0-1 ; 0-1 ; 1,2,2 000 00000000oooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooooo2
2 Distal Exp segment on both right and left P5
P5 absent. tapering into long spine: E ucltaeta
2a Distal Exp segment of left P5 tapers distally to
Males end bluntly bearing minute terminal spine, but
Similar to female, but appendicular caudal setae much never tapers distally into long, strong spine:
less developed than in female. A 1 with 8-10 segments Paraeuchaeta
completely fused; segments 12 and 13 partially fused;
long seta fo und on each of segments 3, 9, 14, 18, 2 1, Key to species of E uchaeta (adapted from Park,
and 24. Setation on A2, Md, Mx 1, and Mxp and 1995, with changes)
sometimes even limb itself (Mx2) reduced compared Females
with female. P5 large and complicate. Male left P5 I Mx I Li 1 with 2 posterior setae; Mx I Enp I with
Exp2 extends distally into serrated lamella (sl) and 4 setae: Euchaeta spinosa Giesbrecht, 1892
with digitiform process (dp); Exp3 with hairy tubercle (Fig. 7.147)
(ht) proximally and distal groups of hairs; segment I a Mx I Li I with 1 posterior seta; Mx 1 Enp I with I
extended beyond these spines into spine-like process seta ...... .................................. ................. ......... ...2
of various lengths or vestigial. 2 Mxp B with short spinules only along proximal
half of medial margin 00 0000000000 00 0000 0000000000 00 00 0000 00003
The family includes genera Euchaeta Philippi, 1843, 2a Mxp B with long marginal spinules and short
and Paraeuchaeta Scott, 1909. Recently a revision marginal spinules along proximal half of medial
was made of this family (Park, 1995). The genus margin 000000000000000000000000000000000 ooooooooooooo·····•oooooooo .... 5
Valdiviella Steuer, 1904 is usually also included in this 3 Dorsally Gns nearly symmetrical: E ucltaeta
family, but is distinct from Euchaetidae: two-pointed pubera Sars, 1907 (Fig. 7.146)
Copepoda 925
~ding Dorsally Gns asymmetrical ............................. ..4 6a On left P5 Exp2 s l without a gap between medi-
Li2-Li3 G:JS \\ith right side step posterior to middle ( dor- al and distal poti ions of distal part of lamella:
~ihan sa: \iew): Euclzaeta media Giesbrecht, I 888 Euchaeta concinna (Fig. 7.142)
:-£ Enp r1g. - .144)
:a:Jales Gns without step on right side (dorsal view): Key to species of Paraeuchaeta
he: ser- Euchaeta acuta Giesbrecht, 1892 (Fig. 7. 141 ) Females (adapted from Park, 1995, with changes; for
~ P5 T"·o setae ofMx2 Enp armed with both long and key to female Gns morphology see Fig. 6.41; 6.4J)
:.: L:lade shon spinules. Mx1 Enp with 2 spinulose setae: 1 Appendicular caudal setae geniculated ............. 2
~- Euclzaeta marina (Prestandrea, I 83 3) (Fig. la Appendicular caudal setae smoothly curved
- .1-B) ......................................................................... 19
One seta of Mx2 Enp armed with both long and 2 Genital flanges elongated along length of genital
shoner spinules. Mx I Enp with 1 spinulose seta field in form of tongue, and usually subdivided
.......................................................................... 6 into anterior and we ll-developed posterior lobes
Gns right side with moderately developed pro- ...... ............................................... ... ............ ....... 3
. ..,
r;;ec_ ____ _
Jection on posterior half of somite: Euchaeta 2a Genital flange neither elongated along length of
~ paraconcinna Fleminger, 1957 (Fig. 7.145) genital field nor produced into posterior lobe; of
Cbtal Gns right side with strongly developed projec- different shape ....................................... .......... 16
non at about midlength: Euchaeta concinna 3 Supra labrum pointing obliquely forward. Mx I
~or Dana. 1849 (Fig. 7.142) Lil w ith 1+9+3 setae .... ...... .............................. .4
oc dis- 3a Supralabrum pointing ventrad. Mx1 Lil with
u::ripal bles based on Park, 1995, with changes; for key to l +9+ 2 setae ...................................................... 14
- -----e·iarions of male left Exp see Fig. 6.7M) 4 Mx 1 Le 1 with 7 large distal setae extending and
sl of left P5 Exp2 not extending as far as tuft of 2 medium sized, densely plumose proximal
snff hairs .......... ..... ................................... ....... ... 2 setae: Paraeuchaeta hansenii (With, 1915)
me dp: 2 sl of left P5 Exp reaching or extending beyond (Fig. 7.158)
mft ofstiffha irs ................................................ .5 4a Mxl Lel with different number and/or type of
:md dp Left P5 Exp much shotier than right. Terminal setae ........................... .............. ....................... ... 5
______ 2
spine of left P5 Exp relatively short: Euchaeta 5 Mx I Lel with 5 long setae in addition to some
~ P5 spinosa (Fig. 7. 147) very small setae proximally ............................... 6
Both P5 Exp similar in length. Terminal spine of Sa Mxl Lei with 6 or 7 long setae in addition to 1
y to left P5 Exp very long ......................................... 3 or 2 small proximal setae pointing laterad and/or
~:e.. but dp of left P5 Exp2 terminates in sharp point: some minute setae proximally ................. ........ 13
~me: Euchaeta pubera 6 In lateral view anterior edge of genital swelling
dp ofleft P5 Exp2 is smoothly rounded terminal- usually slightly oblique, high ... ....... ........ .......... 7
ly ..... ........................................ .... .......... ..... ....... .4 6a In lateral view anterior edge of genital swelling
1 Park. sl of left P5 Exp2 bilobed: Euchaeta media (Fig. usually significantl y oblique, low ..................... 9
- . 144) 7 In lateral view genital flange strongly emargi-
sl of left P5 Exp2 not bilobed: Euchaeta acuta nate with posterior lobe prominent ................... 8
l v.idl Fig. 7.141) 7a In lateral view genital flange slightly emarginate
- l ~92 Left P5 Exp3 with dpsl next to tuft of stiff ha irs: without prominent posterior lobe: Paraeuchaeta
Eucha eta marina (Fig. 7.143) sarsi (Farran, 1908) (Fig. 7. 164)
liith I Left P5 Exp3 without dpsl next to tuft of stiff 8 In lateral view posterior lobe of genital flange
.., hairs ........................ ......... ........................... ..... ..6 d irected posteroventrad; total length 7.4-8.4
On left P5 Exp2 sl distal margin with relatively mm: Paraeuchaeta calva Tanaka, 1958 (Fig.
_____3 large teeth in lateral portion and fine teeth in 7.153)
medial potiion, these two potiions are separated 8a In lateral view posterior lobe of genital flange
from each other by gap between medial and dis- directed ventrad; total length 8.8-9.6 mm:
I •••••5 tal part of lamella: Euclraeta paraconcinna Paraeuchaeta regalis (Grice and Hulsemann,
Fig. 7. 145) 1968) (Fig. 7.163)
926 Copepoda
9 In lateral view R very small: Paraeuchaeta par- 19 Mxl Li2 with 2 setae: Paraeuchaeta antarctica
vula (Park, 1978) (Giesbrecht, 1902) (Fig. 7.149)
9a In lateral view R well-developed ....... ............. ! 0 19a Mx I Li2 with 1 setae: Paraeuchaeta hisinuata
10 In lateral view dorsal edge of Gns with distinct (Sars, 1907) (Fig. 7 . 152)
hump at midlength or posterior to midlength .11
1Oa In lateral view dorsal edge of Gns without dis- Males (unknown for P. dactylifera and P. vervoorti)
tinct hump ....... ......................... ............... ........ . 12 (for key to abbreviations of male left Exp see Fig.
II In lateral view gential swelling with 2 conspicu- 6.7M)
ous "semicircular" ridges on each side: I P5 right Exp2 spiniform: Paraeuchaeta hilobtt
Paraeuchaeta rasa Farran, 1929 (Fig. 7.162) (Fig. 7.151)
!Ia In lateral view genital swelling with low ridge Ia P5 right Exp2 not spiniform .. ..................... .. ..... 2
close to anterior margin on each side: 2 Left P5 Exp with short dp (shorter than half
Paraeuchaeta aequatorialis Tanaka, 1958 (Fig. length of sl). Right P5 Enp with lobe ................ 3
7.148) 2a Left P5 Exp with dp longer. Right P5 Enp with-
12 Gns with process on left: Paraeuchaeta barbata out lobe ...... ....................................................... .4
(Brady, 1883) (Fig. 7.150) 3 sl of P5 Exp2 short and bordered with small
12a Gns without process on left: Paraeuchaeta kuri- number of large teeth; ht rounded without distal
lensis Heptner, 1971 (Fig. 7.159) spiniform process: Paraeuchaeta bisinuata
13 1n lateral view dorsal edge ofGns without hump; (Fig. 7.152)
low ridge present anterodorsally on left of 3a sl of P5 Exp2 not short, ht not rounded but taper-
somite: Paraeuchaeta comosa Tanaka, 1958 ing into long spiniform process: Paraeuchaeta
(Fig. 7.154) antarctica (Fig. 7 .149)
13a In lateral view dorsal edge of Gns with conspic- 4 sl of left P5 Exp2 tapering into spiniform process
uous hump; low ridge absent anterodorsally on with large marginal teeth .. ......... .......... ......... ..... 5
left of somite: Paraeuchaeta scotti (Farran, 4a sl of left P5 Exp2 not tapering into spinifonn
1908) (Fig. 7.1 65) process and without large marginal teeth .......... ?
14 Genital flange without finger-like or tooth-like 5 Left P5 Exp I with flat-top hump on posterior
projection: Paraeuchaeta vervoorti Park, 1995 margin: Paraeuchaeta pseudotonsa (Fig. 7. 161)
14a Genital flange with finger-like or tooth-like pro- Sa Left P5 Expl with conspicious conical process
jection ............................. .. .. .. ........................... 15 midway along posterior margin ......................... 6
15 Mx 1 Li4 with 3 setae. R very small: Para- 6 Anterior margin of R flat (lateral view). On left
euchaeta dactylifera (Park, 1978) (Fig. 7 .155) P5 Exp I conical process on posterior margin
15a Mx 1 Li4 with 5 setae. R longer: Paraeuchaeta rounded. sl of left P5 Exp2 bordered with teeth on
sesquipedalis Park, 1995 (Fig. 7.166) both edges: Paraeuchaeta ex.igua (Fig. 7.156)
16 Mx I Li 1 with l +9+3 setae, Li2 with I setae, and 6a Anterior margin of R smoothly curved (lateral
Li4 with 4 or 5 setae ........................................ 17 view). On left P5 Exp1 conical process on poste-
16a Mx 1 Li 1 with 1+9+ I setae, Li2 without seta, and rior margin triangular. sl ofleft P5 Exp2 tapering
Li4 with 3 setae: Paraeuchaeta biloba Farran, into very large spine, curved strongly inward and
1929 (Fig. 7.151) with only its medial edge bordered with teeth:
17 In lateral view Gns with large cylindrical genital Paraeuchaeta gracilis (Fig. 7 .157)
swelling: Paraeuchaeta gracilis (Sars, 1905) . 7 Left P5 Exp2 medial margin without isolated
(Fig. 7.157) tooth-like process close to proximal end ........... 8
17a In lateral view Gns with bilobed or rounded tri- 7a Left P5 Exp2 medial margin with isolated tooth-
angular genital swelling ................................... 18 like process close to proximal end .................. ! 0
18 In lateral view genital swelling rounded, triangu- 8 R terminated with small spiniform process. dp
lar: Paraeuchaeta pseudotonsa (Fontaine, 1967) with massive base and tapering distally into
(Fig. 7.161) slender, more or less spiniform process:
18a In lateral view genital swelling bilobed: Para- Paraeuchaeta sesquipedalis
euchaeta exigua (Wolfenden, 1911) (Fig. 8a R longer than in P. sesquipedalis. dp not taper-
7.156) ing distally, but massive and rounded ............... 9
Copepoda 927
r:.!lTClica sl of left P5 Exp2 terminated with 2 large teeth. Ia Gns without conspicuous ridge crossing dorsal
PI Exp2 outer spine about 2/3 length of Exp3 surface. Mx I Le 1 with more than 4 setae ......... 2
spine: Paraeuc/zaeta rasa 2 Posterior corners of Pr (dorsal v iew) slightly
sl of left P5 Exp2 lacking 2 large teeth terminal- diverging. Mxl Lei w ith 7 setae: Valdiviella oli-
ly. PI Exp2 outer spine about as long as Exp3 garthra Steuer, 1904 (Fig. 7.171)
n-oorri) spine: Paraeuchaeta aequatorialis (Fig. 7.148) 2a Posterior corners ofPr (dorsal view) not diverg-
see f ig. 10 dp of left P5 Exp2 distally pointed ................. 11 ing, but obliquely c ut off or as round lappet.
lOa dp of left P5 Exp2 distally rounded ... ............. 12 Mxl Lei with 6 setae .................................... .... 3
r biloba I~ Distal extension of sl narrow, not rectangular, 3 Pr about 2.5 times as long as Ur: Valdiviella
less than half width of proxima l pa1t: insignis Farran, 1908 (Fig. 7 .169)
--······2 Paraeuchaeta scotti (Fig. 7.165) 3a Pr about twice or less than twice length of Ur:
~half lla Distal extension of sl slightly nan·ower than Valdiviella brevicornis Sars, 1905 (Fig. 7.168)
- ...... 3 proximal portion, nearly rectangular:
9 \loith- Paraeuclzaeta barbata (Fig. 7 .150) Males
_____.4 C Distally end of sl of left P5 Exp2 with 2 widely I Posterior corners of Pr rounded, both without
- small diverging spiniform processes: Paraeucltaeta tooth ................................................................... 2
distal regalis I a Posterior corners of Pr elongated into small
m uara ::a Distal end of sl of left P5 Exp2 without 2 wide- tooth .................. ................................................. 3
ly diverging spiniform processes ........... .......... 13 2 Left P5 Enp shotter than half length of left Exp l.
u r:!peT- 13 Posterolatera l margin of Pr with small tooth-like Right P5 Exp2 not tapering into spiniform p ro-
Jidw.eta process ............................................................. 14 cess, but obtuse: Valdiviel/a brevicornis (Fig.
13a Posterolateral margin of Pr without tooth-like 7.168)
process ............................................................. 15 2a Left P5 Enp longer than half length of left Exp I.
1.! R elongated. s l of left P5 Exp2 consists of elon- Right P5 Exp2 tapering into spiniform process:
~orm gate main body with small spiniform distal pro- Valdiviella minor (Fig. 7 . 170)
l-... ..... 1 cess along lateral margin: Paraeuch.aeta calva 3 Left P5 B longer than corresponding Exp seg-
IC:Slcrior (Fig. 7 .1 53) ment. Left Enp exceeding well beyond middle of
:.. - :61) l.!a R not elongated. sl of left P5 Exp 2 consists of left Exp I: Valtliviella oligarthra (Fig. 7.171 )
process more or less rectangular base and dagger shaped 3a Left P5 B nearly of same length as conespond-
______ 6 extension straight along lateral margm: ing Exp segment. Left Enp slightly exceeds mid-
0:: left Paraeuchaeta sarsi (Fig. 7.164) dle of left Exp l: Valdiviella insignis (Fig.
~ dp markedly enlarged proximally: Paraeuc/zaeta 7. 169)
eemon hansenii (Fig. 7.158)
.156) ~ .:a dp is nearly of same width over its whole length Family Phaennidae Sars, 1902
......................................................................... 16 References: Bradford (1973), Bradford et a/. (1983),
posre- 16 sl consists of rectangular basal part and spini- Park ( l983b), Fenari and Markhaseva (1996).
~g form distal part extending along external margin Females
~:rod of basal part: Paraeuchaeta parvula Ce and Pdl usually fused, or separated, Pd4 and 5 fused
- ;:eeili: ~ 6a sl is of different shape .. ........... ........................ 17 completely, or incompletely. R present, of 2 filaments
~- sl tapering in dagger shape. dp di stally enlarged. (Xanthocalanus, Brachycalanus), or 2-pointed
R relatively long: Paraeuchaeta comosa (Phaenna), or as a bifurcated plate with slight
- ...... 8 !-a sl in form of triangular hollow scoop. dp not (Neoscolecith.rix), or strong (Corn.ucalanus,
c. ~"lOth enlarged distally. R relatively small: Onchocalanus) bifurcation. Posterior corners ofPr and
___ lO Paraeuclzaeta kurilensis Gns symmetrical. Poste1ior corners of Pr extended into
::ess. dp 2 points each (Neoscolecithrix), prolonged into lobes
into Key to species of Valdiviella (Xanthocalanus), or not prolonged into lobes, rounded
lrnC~:5: Females (adapted from Park, 1978, with changes) or triangular.Ur of 4 sornites. Al 21-25-segrnented, 8-
Gns with conspiciuos ridge crossing dorsal sur- 9 segments fused. A2 Exp nearly equal or longer than
~
face. Mx l Lei with 4 setae: Valdiviella minor Enp, 6-segmented; Exp I without setae, Exp 2 without
___ 9 Wolfenden, 1911 (Fig. 7.170) medial setae (in Neoscolecithrix Exp 1-Exp2 with 2
928 Copepoda
setae each). Mdp B usually with 3 setae. Mx 1 Li I usu- Males

ally with 8-9 terminal and 4 posterior setae R similar to that of females. Oral pa1ts reduced com-
(Xanthocalanus, Cephalophanes, Neoscolecithrix), 7 pare d with those of female. PS un iramous, legs almost
terminal and 3 posterior setae (Brachycalanus), or only equal in length, left leg S-segmented, slightly longer
9 terminal setae in Phaenna; and with total number of than 4-segmented right leg (Phaenna), or strongly
I 0- 12 setae in Cornucalanus and Onchocalanus; Li2 asymmetrical, unequal in length, with long S-segment-
usually with 2 setae, sometimes with 1 seta (Phaenna), ed left leg, and sh01t 3- or 4-segmented right leg;
3 setae (Neoscolecithrix, some species of sometimes right PS 5-segmented and very shott, reach-
Xanthocalanus ), or 4-5 setae (some species of ing only first segment of left leg (Xanthocalanus).
Xanthocalanus , Talacalanus); Li3 with 3-4 setae; Li4
usually with 5 setae, rarely with fewer setae (3 setae in There are 6 genera in Phaeru1idae: Brachycalanus
Phaenna): Enp has from 6 (Phaenna) up to 13 setae Farran, !90S; Cephalophanes Sars, 1907; Cornu-
(Brachyca/anus); Exp usually with 10 setae (in calanus Wolfenden, l90S; Onchocalanus Sars, 190S;
Phaenna 5 setae ): Lei with 9 setae. Mx2 Lil-LiS with Phaenna Claus, 1863; and Xanthocalanus Giesbrecht,
S (rare ly 4). 3 . 3 (2 in Phaenna), 3 (2 in Phaenna) and 1892. Neoscolecithrix Canu, 1896 is of unclear taxo-
4 setae respectively; one seta of LiS may be trans- nomic status (see Schulz and Beckman, l99S) and is
formed into sensory worm-like appendage (Phaenna, placed conventionall y here in the Phaen nidae.
some species of Brachycalanus, Xanthocalanus har- Talacalanus Wolfenden, 1911 has been redefined by
pagatus, in latter species one of setae on Li3 also Campaner ( 1978b) to accomodate Xanthocalanus
worm-like); LiS has strong claw-like spine in greeni Farran, !90S, and X. maximus Brodsky, 19SO,
Cornucalanus and Onchocalanus; terminal part of limb but this genus seems very c lose if not identical to
usually with l worm-like and 7 brush-like sensory setae Xanthocalanus. Representatives of a ll genera except
(in some species of Xanthocalanus and Brachycalanus Cephalophanes have been taken in the South Atlantic.
2 wonn-like appendages and 6 brush-like appendages
have been recorded), rarely nwnber of brush-like Key to Phaennidae genera
appendages greater (8 in Brachycalanus bjornbergae); Females
number and composition of sensory appendages in I PS absent. Mx l Exp with S setae, Li2 with l seta
Neoscolecithrix diffe rs (3 worm-like and 4-S brush- and Li3 with 3 setae. Pr is ovoid in dorsal view:
like), similar to that of scolecitrichids. Mxp C with 1 Phaenna spinifera C laus, 1863 (Fig. 7 .180)
proximal seta, then with 2 (rarely I ) setae, 2 setae (usu- la P5 present. Mxl Exp with more than S setae, usu-
ally I brush-like and 1 sclerotized seta) and 3 setae ally w ith 10 setae, Li2 with more than I seta a nd
from proximal to di stal; in Neoscolecithrix setal com- Li3 usuall y with more than 3 setae. Pr not ovoid
position differs: l proximal seta, then 2 setae ( I worm- in dorsal view .............................. ...................... 2
like and I scierotized), 2 setae ( 1 brush-like and 2 sci e- 2 Posterior corners of Pr extended into 2 points
rotized) and 3 setae from proximal to distal; Enp2-Enp6 each: Neoscolecithrix caetanovi Alvarez, 198S
usually with 4, 4, 2-4, 3-4 and 4 setae; sometimes (Fig. 7. 175)
Enp2-3 each with strong, spine-like attenuations 2a Posterior corners ofPr not extended into 2 poin ts
(Cornucalanus). P 1 Enp with well developed external ................................. ... ................. .................. .... 3
lobe. P2-P4 wide and flat, surfaces may be covered by 3 R is sh01t plate, or w ith 2 filaments. Mx2 has 1
spines and spinules, sometimes strongly (Brachy- sto ut, spine-like seta, but never claw-like ........ .4
calanus). Spine and seta formula: 3a R is large plate subdivided d istally into 2 large
ram i (bifurcate). Mx2 has strong claw-like spine
coxa basis exopod segments endopod segments on L i5 ................................................................. 5
4 A l short, nearly reaching last somi te of Pr, w ith
Leg 1 0-0 0-0/ 1 I-0: I- 1; 1,1,3 0,2,3
Leg 2 0- 1 0-0 1- 1; 1-1; 111.1,4 0-1 ; 1,2,2
knife-shaped aesthetasks. Ur and posterior and
Leg 3 0-1 0-0 1-1 ; 1- 1; lll,1,4 0- 1; 0- 1; 1,2,2 anterior surfaces of PI-PS densely spinulose:
Leg4 0-1 0-0 1-1 ; 1- 1; 111.1,4 0-1 ; 0- 1; I ,2,2 Brachycalanus bjornbergae Campaner, 1978
(Fig. 7 .172)
PS usually 3-segmented, sometimes 2-segmented 4a Al longer, aesthetasks, if present, of different
(Xanthocalanus in pa rt), or absent (Phaenna). shape. Ur and posteri or and an terior surfaces of
Copepoda 929
Pl-P5, if spinulose, not densely so: Xantho- Key to species of Onchocalanus

::rl oom- calanus Females
iaJnost Mxp Enp2-3 each with strong, spine-like attenu- I Forehead w ith crest: Onchocalanus subcrista-
• :Onger ations. Pl B with inner marginal seta: tus (Wolfenden, 1906) (Fig. 7. 177)
s::ongly Comucalanus Ia Forehead without crest .............................. ... ..... 2
eg:;:nent- 5a Mxp Enp2-3 without attenuations, but with typ- 2 In lateral view Gns without well pronotmced
;:.... leg: ical setae. P 1 B w ithout inner marginal seta genital swelling: Onchocalanus trigoniceps
::.. .reach- (excluding Onchocalanus magnus): Oncho- Sars, 1905 (Fig.7.178)
m). ca/anus 2a In lateral view Gns with well pronounced genital
swelling .... ..........................................................3
crzlanu.s :'\lales (unknown for Brachycalanus) 3 R with wide base, rami strong, tapering into fil-
Comu- R bifurcate ......................................................... 2 aments. Posterior corners of Pr angular (lateral
i.. 1905: l3 R not bifurcate, with two filaments: Xantho- view). T hird segment of P5 much longer than
silrecht. ca/anus second segment: Onchocalanus magnus
!r i:a.."i:o- Mx I Exp with 5 setae. P5 legs almost equal in (Wolfenden, 1906) (Fig. 7.176)
:!nd is length (left leg 5-segmented, slightly longer than 3a R with narrow base, rami strong, with minute
Clidae. 4-segmented right leg): Phaenna spinifera (Fig. hair terminally. Posterior corners of Pr rounded
iDee by 7.180) (lateral view). Third segment of P5 nearly equal
:e...arzu.s Mxl Exp with more than 5 setae, usua lly with 8, in length (or slightly longer) than second seg-
- :950. 10 setae. P5 very unequal in length, with long 5- ment: Onchocalanus wolfendeni Vervoort,
~to segmented left leg, and shott 3- or 4-segmented 1950 (Fig. 7.179)
~cep t right leg ........... ................................................... 3
.:::.~,tic. Posterior comers of Pr extended into 2 points Males (unknown for 0. subcristatus)
each: Neoscolecithrix caetanovi (Fig. 7 .175) Left P5 about twice as long as Ur. Right P5 much
Posterior comers ofPr not extended into 2 points shmter than left leg segment 1. Penultimate seg-
each ............... ................................................... ..4 ment of left leg with row of strong spinules:
:sera .! Ylx Li4-5 each have large spiniform setae: Onchocalanus magnus
._ --:.ew: Cornucalanus Ia Left P5 shorter. Right P5 longer than left leg
Ylx Li5 has large spiniform seta: Onchocalanus segment 1. Penultimate segment of left leg with-
~ !bll- out row of strong spinules ................................ .2
ei:! and io.::ey to species of Cornucalanus 2 Rostral plate narrowed at its base. Left P5 with
lm·oid females distal segment of about 1/3 length of penultimate
___ .1 Forehead with spiniform process. Mxp B with segment: Oncltocalanus wolfendeni
;:c::ms large marginal teeth. P5 w ith single terminal 2a Rostral plate not narrowed at its base. Left P5
~ :985 spine; Cornucalanus chelifer (Thompson, with distal segment shorter than 1/3 length of
1903) (Fig. 7.173) penultimate segment: Onchocalanus trigonic-
a Forehead without spiniform process. Mxp B eps (Fig. 7.178)
with relatively small marginal teeth. P5 with 2
- has 1 term inal spines: Cornucalanus robustus Key to species of Xantlzocalanus
____ _.t
YervoOlt, 1957 (Fig. 7.1 74) Females (unknown for X. soaresmoreirai)
- 1.:!..-ge 1 P5 3-segmented: Xauthocalanus agilis
~spine hJes Giesbrecht, 1892 (Fig. 7. 181)
Anterior part of head (lateral view) with deep, Ia P5 2 -segemented: Xanthocalanus marlyae
wide gap between R and forehead. P4 C with Campaner, 1978 (Fig. 7.182)
seta: Comucalanus chelifer (Fig. 7. 173)
L .-\nterior part of head (lateral view) without any Males (unknown for X. marlyae)
gap between R and forehead. P4 C without seta: 1 Rostral filaments are sausage-like. Mxl Exp
Cornucalanus robustus probably with 4 setae: Xanthocalanus soares-
~I moreirai Bj0111berg, 1975 (Fig. 7.183)
t::es of
930 Copepoda
1a Rostral fi laments are not sausage-like. Mxl Exp number and composition of setae in th is group dif-
with 10 setae: Xan thocalanus agilis (Fig. 7. 181) fe rs), and finally 3 distal setae; Enp2-Enp6 usually
with 4, 2-4, 2-3, 3-4 and 4 setae respectively. P l Enp
Family Scolecitrichidae Giesbrecht, 1982 with, or rarely without outer lobe. P2-P4 with Enp
References: Bradford (1973) Bradford et a!. (1983), flattened and usually broad; posterior surface of both
Park ( 1980, 1982, 1983a), Vyshkvartseva (1989a, rami with conspicuous spines. Spine and seta formula:
1989b), Ferrari and Markhaseva ( 1996).
Females coxa basis exopod segments endopod segments
Ce and Pd1 fused, or separate (Mixtocalanus), Pd4
Leg I 0-0 0- 1 0/I-0; 1-1; f,T,3 0,2,3
and 5 usually fused. sometimes separate (Scolecithrix, Leg 2 0-1 0-0 I-I ; l-1 ; TTI,l,4 0-1 ; 1,2,2
Scopalatum part., etc.). Posterior com ers of Pr round- Leg 3 0-1 0-0 l- 1; l- 1; 111, 1,4 0-1 ; 0-1 ; I ,2,2
ed, or extended in rounded or triangul ar lobes, may be Leg4 0-0/ 1 0-0 1-1 ; I-I; !Tl,l,4 0-1; 0-1; I ,2,2
pointed (Puchinia). Forehead low, rounded, or highly
produced in form of triangle (lateral view), crest P5 present or absent, if present, uniramous, 1-3 seg-
present or absent. R of 2 filaments, biramous, or as mented, usually symmetrical, with 1-4 terminal spines.
plate, bifurcate or not, with or without filaments. Ur of
4 somites. A1 19-23 segmented; further study of A I Males
morphology needed before more detailed description Pd4 and 5 may be partly or completely separate
can be given. A2 Exp usually 6-segmented (7-seg- (Amallothrix, Lophothrix, Mixtocalanus, Scapho-
mented in Scottocalanus); Exp usually longer than calanus, Scolecithrix, Scottocalanus). R and Pl -P4
Enp, o r may be eq ual in length (Lophothrix, usually similar to that of fema le. Mouthparts usually
Landrumius, Undinothrix), Enp may even be longer or reduced compared with females. P5 asymmetrical, - .!
more powerful (Archescolecithrix, Heteramalla, usua lly biramous on both sides (excluding
Mixtocalanus, Scaphocalanus); Exp1 usually lacking Scolecithrix, Scolecithricella ovata, etc.); with moder-
setae (1 seta in Puchinia), Exp2 lacking medial setae ately narrow and elongated C and B on left; with B
(this segment composed of2 fused segments and bears short and usually very swollen on right.
1 distal margina l setae that commonly belongs to
Exp3). Mdp usually with 1-3 setae, Enp 1 with 0-2, The fa mily includes 19 genera: Amallothrix, Sars,
Enp2 with 9 setae. Mx1 Li1 with 1-4 posterior setae, 1925; Archescolecithrix Vyshkvartseva, 1989; Hetera-
7-9 terminal setae and sometimes I anterior seta malla Sars, 1907; Landrumius Park, 1983; Lophothrix
present; L i2 us ually with 2-3 setae (absent in Giesbrecht, 1895; Macandrewella Scott, 1909;
Heteramalla sarsi, in some Mixtocalanus species - 1 Mixtocalanus Brodsky, 1950; Parascaphocalanus
setae, in Puchinia 5 setae); Li3 with 3-4 setae; Li4 Brodsky, 1955; Pseudophaenna Sars, 1902; Puchinia
usually with 4-5 setae; Enp usually 1-2 segmented, Vyshkvartseva, 1989; Racovitzanus G iesbrecht, 1902;
with 3+5 or 3+6 setae respectively, sometimes this Scaphocalanus Sars, 1900; Scolecithricella Sars, 1902;
number is different (6, 7 or 8 setae); Exp with 5-10 Scolecithrix Brady, 1883; Scolecocalanus Farran,
setae; Lel with 9 setae. Mx2 Li I with 3-4 setae (5 in 1936; Scopalatum Roe, 1975; Scottocalanus Sars,
Landrumius), Li2-Li4 with 3 setae each, Li5 with 4 1905; Xantharus Andronov, 1981; Undinothrix
setal elements of which usually 1, and rarely 2 may be Tanaka, 1961. Seven of these genera (dealt with below)
transformed into worm-like sensory appendages; ter- have been recorded in the South Atlantic.
minal part of limb with 3 worm-like and 5 brush-like
sensmy appendages (in Racovitzanus 4 worm-like Key to Scolecitr ichidae gener a
plus 4 brush-like and in Parascaphocalanus 4 worm- Females
like plus 2 brush like respectively). Mxp C usually I R is large strongly chitinized plate: with terminal
with I proximal seta (or 1 worm-like appendage excavation, bifurcate termi nally, or deeply
instead of this seta); then from proximal to distal: 2 div ided into 2 strong ram i each terminated with
setal elements, usually represented by I sclerotized small conical point (in Lophothrix latipes termi-
seta plus 1 worm-like appendage (but there may be nated with filaments). P5 3-segmented, if 2-seg-
also 2 sclerotized setae, or 1 sclerotized setae, or I mented, then with very long sub-apical spine
worm-li ke appendage), 1 brush-like appendage (rarely directed backwards (reaching Ur3) ................... 2
Copepoda 931
~ dif- Ia R less developed, usually of 2 filaments, rarely 3 posterior setae: Scottocalanus securifrons
:!5llally 2-pointed or bifurcate plate with 2 thick fila- (Fig. 7.213)
?: Enp ments (separated or not from plate). P5 1-2-seg- 5a R with filaments. Mxl Lil with 1-2 posterior
"iit Enp mented, rarely absent ... ............. ...... ... ............ ... .4 setae .. .... ................. .... ................................ ... .. ... 6
o:-both 2 P 1 B without i1mer marginal setae. Mx2 Li l with 6 PI Expl with outer spine: Amallothrix
~ula: 5 setae. Mxl Exp with 10 setae: Landrumius 6a Pl Expl without outer spine: Scolecithricella
gigas (A. Scott, 1909) (Fig. 7 .189)
2a P l B with inner marginal seta. Mx2 Li I with 3- Key to species of Amallothrix
4 setae. Mx 1 Exp with fewer setae ................... 3 Females (partly adapted from Park, 1980, with chang-
3 P5 3-segmented, terminal segment with 3 or 4 es)
spines. A2 with Exp and Enp of about equal 1 R with long rami tapering into short filaments
length. Mx 1 Li 1 with 4 posterior setae: ...................... .. ................. ............. .... ..... .......... .. 2
Lophothrix 1a R with short rami tapering into filaments ......... 3
-3 seg- 3a P5 2-segmented, tenninal segment with 2 spines, 2 Spennatheca with round distal sac separated by
spmes. one very long (reaching Ur3) sub-apical spine narrow neck: Amallothrix valida (Farran, 1908)
directed backwards. A2 Exp longer than Enp. (Fig. 7.188)
Mxl Lil with 3 posterior setae: Scottocalanus 2a Spennatheca without such separate distal sac,
;;eparate securifrons (T. Scott, 1894) (Fig. 7.213) but thin and finger-like: Amallothrix dentipes
~apho -t P5 absent, or vestigial. PI Enp produced distally (Vervoort, 1951) (Fig. 7.184)
• P : -P-t into prominent spiniform process along outer 3 Mx2 Li5 with 2 sensory worm-like appendages.
~y margin: Scolecithrix Mdp B with 3 setae: Amallothrix obtusifrons
::c:rical. .!a P5 present, 1-2-segmented (absent in Scapho- (Sars, 1905) (Fig. 7 .187)
:biing calanus curtus, 3-segmented in Scolecithricella 3a Mx2 Li5 with 1 senso1y worm-like appendage.
r::::oder- pseudoalata). PI Enp without prominent spini- Mdp B with 2 setae ....................... ..... .......... .....4
w:th B form processes along outer margin ........ ........ ... 5 4 Total length more than 5 mm. P5 distal spine
P5 !-segmented, flattened, plate-like. Mx I Li3 almost 2/3 as long as inner spine: Amallothrix
with 3 setae: Scolecithricella hadrosoma (Park, 1980) (Fig. 7.186)
[. Sals. 5a P5 2-segmented, not flattened, not plate-like. 4a Tota1length less than 5 mm. P5 inner spine more
u erera-
•• Mx l Li3 usually with 4 setae, rarely with 3 setae than twice as long as distal spine: Amallothrix
Yto;J;rix ......... .................................................................. 6 emarginata (Farran, 1905) (Fig. 7 .185)
l909: " PI Expl usually without outer spine. Mx1 Enp
r:alanu.s usually with 3+6 setae: Scaphocalanus MaJes (unknown for A. hadrosoma, doubtful descrip-
";inia 5a Pl Expl usually w ith outer spine. Mxl Enp usu- tion for A. obtusifrons)
L.. IQ02: ally with 3+ 5 setae: Amallothrix I P5 right B only slightly swollen: Amallothrix
i. !902: emarginata (Fig. 7 .1 85)
~laJes (unknown for Landrumius) I a P5 right B very swollen ........... .......................... 2
w Sa:rs. P5 biramous on both sides (except for 2 Right P5 Exp2 strongly curved: Amallothrix
:.":rir: Scolecithricella ovata) ..................................... .. 2 dentipes (Fig. 7. 184)
below) L Right P5 uniramous: Scolecithrix 2a Right P5 Exp2 slightly curved: Amallothrix vat-
Left P5 Enp longer than Exp ... ....... ...... .... ......... 3 ida (Fig. 7.1 88)
Left P5 Enp shorter than Exp ........................... .5
D istal segment of right P5 Exp is triangular plate. Key to species of Lophothrix
R with strongly chitinized thick rami: Lophothrix Females
D istal segment of right P5 Exp of different 1 Anterior part of Ce with crest ..... .......... ............ 2
shape. R of 2 filaments .......... ..... .. .... .............. .. .4 1a Anterior part of Ce without crest ..... ..... ... ..... .. ..4
Right P5 B only slightly swollen: Amallothrix 2 P5 terminal segment with 4 spines: Lophothrix
-~~ emarginata (Fig. 7.185) quadrispinosa Wolfenden, 1911 (Fig. 7.194)
:- .:-s._~- Right P5 B strongly swollen: Scaphocalanus 2a P5 tetminal segment with 3 spines .................... 3
1 spine R in fonn of strongly chitinized plate distally 3 Forehead highly produced in form of triangle
, di\·ided into 2 thick rami. Mxl Lil usually with (lateral view); posterior corners ofPr rounded. R
932 Copepoda
without filaments: Lophotlzrix frontalis (these two species are difficult to distinguish)
Giesbrecht, 1895 (Fig. 7.190) (Fig. 7.196, 7.200)
3a Forehead not highly produced, rounded (lateral 6a Inner spine of P5 extends far sho1t of distal end
view); posterior corners of Pr with tooth-like of term inal spine: Scaphocalanus echinatus
distal process. R with filaments: Loplzothrix lat- (Farran, 1905) (Fig. 7.198)
ipes (T. Scott, 1894) (Fig. 7. 192)
4 P5 terminal segment with 4 spines: Loplwthrix Males
simi/is Wolfenden, 1911 (Fig. 7.193) I PI B inner marginal seta moderately developed
4a P5 term inal segment with 3 spines: Lophothrix ....... .................................................. .................. 4
humilifrons Sars. 1905 (Fig. 7. 191) Ia PI B inner marginal seta very small or reduced
......................... .... ......... ..................... ............. ...2
Males (unknown for L. simi/is and L. quadrispinosa) 2 Mx l Lil with 2 posterior setae, Enp with 6, Exp
I Anterior forehead with low crest: Lophothrix with 4 setae respective ly: Scaphocalanus curtus
latipes (Fig. 7 .192) (Fig. 7.197)
Ia Anterior forehead without crest.. .. ..................... 2 2a Mxl Lil with 3 posterior setae, Enp with 3+6
2 Forehead triangu larly produced forward, but setae, Exp with 7 setae ...................................... 3
without its anterior part set off as small triangu- 3 Total length more than 2.6 mm; Ur3 with a
lar lobe (dorsal view): Lophothrix frontalis length: width ratio of about 100:76: Scapho-
(Fig. 7.190) calanus farrani (Fig. 7 .200)
2a Forehead with its anterior part set off as small 3a Total length less than 2.4 mm; Ur3 with a
triangular lobe: Lophothrix humilifrons length:width ratio of about 100:86: Scapho-
calanus echinatus
Key to species of Scaphocalanus (adapted from 4 Ur less than half length of Pr ............................ 5
Park, 1982, with changes) 4a Ur longer than half length of Pr: Scaphocalanus
Females elongatus
I P5 absent. Pl Enp without outer lobe; B without 5 Ur2 with straight sides; Ur3 longer than wide.
inner marginal seta: Scaphocalanus curtus Mx I Li4 often with 6 setae: Scaphocalanus par-
(Farran, 1926) (Fig. 7.197) antarcticus (Fig. 7.202)
la P5 present. PI Enp with outer lobe; B with inner 5a Ur2 with inflated sides; Ur3 wider than long.
marginal seta ........... ..... ... ............. ...................... 2 Mxl Li4 often with 5 setae: Scaphocalanus
2 Forehead with crest .. .... ........ ........................ .. ... 3 a11tarcticus
2a Forehead without crest ...................................... 5 Males of S. magnus and S. brevicornis are not included
3 In dorsal view forehead broadly rounded, with in the key because their descriptions are not detailed.
low crest: Scaphocalanus antarcticus Park,
1982 (Fig. 7. 195) Key to species of Scolecithricella (based on Park,
3a In dorsal view forehead produced into triangular 1980, and Campaner, 1984)
form, with conspicuous crest.. .................... ...... .4 Females
4 Posterior comers of Pr produced into elongated I Posterior corners of Pr h·iangular, extending
lobe reaching almost to spermatheca: Scapho- almost to distal part of Gns. Mx2 Li5 with
ca/anus magnus (T. Scott, 1894) (Fig. 7.201) coarsely sen·ated spine: Scolecithricel/a pseudo-
4a Posterior comers of Pr produced into relatively culata Campaner, 1979 (Fig. 7.208)
short lobe, extending far sh011 of spermatheca: I a Posterior corners of Pr rounded, short, covering
Scaphocalanus parantarcticus Park, 1982 not more than anterior half of Gns. Mx2 Li5
5 Digitiform extension of spermatheca extraordi- spine not coarsely serrated .... ............ .... ............ 2
narily long, with irregular outline: Scapho- 2 P 1 Exp I with outer spine: Scolecithricella ovata
calanus elongatus A. Scott, 1909 (Fig. 7. 199) (Farran, 1905) (Fig. 7 .206)
5a Digitiform extension of spermatheca not 2a PI Expl without outer spi ne .............. ............... 3
extraordinarily long, with smooth outline ......... 6 3 Posterior corners of Pr emarginate on postrior
6 Inner spine of P5 extending beyond distal end of edge (lateral view): Scolecithricella dentata
terminal spine: Scaphocalanus brevicornis (Giesbrecht, 1892) (Fig. 7.204)
(Sars, 1900), Scaphocalanus farrani Park, 1982 3a Posterior corners of Pr not emarginate ........ ...... 4
Copepoda 933
QgUisb)
. Y1x1 Li4 with 3 setae .................. ....................... 5 Ia Right P5 Exp terminal segment not divided into
~3 Yix 1 Li4 with 5 setae ..... .................................. .. 6 branches: Scolecithrix danae (Fig. 7.2 12)
L~end ' R with short points: Scolecithricella minor
~arus (Brady, 1883) (Fig. 7.205) Family Diaixidae Sars, 1902
:3. R with long filaments: Scolecithricella tenuiser- References: Bradford et a/. (1983), Andronov ( 1979),
rata (Giesbrecht, 1892) (Fig. 7.209) Ferrari and Markhaseva (1996).
P5 inner spine longer than distal spine .......... ... 7 Females
~eloped P5 inner spine much shorter than distal spine: Ce and Pd I fused or separate, Pd4 and 5 separate,
------· A Scolecithricella vittata (Giesbrecht, 1892) (Fig. rounded or pointed, symmetrical, or asymmetrical. R
.OOuced 7.2 10) is divided plate with, or without filaments, or non
---····1 Yix l Enp with 8 setae: Scolecithricella proftm- divided plate with filaments. Ur of 4 somites; Gns
6.Exp da (Giesbrecht, 1892) (Fig. 7.207) symmetrical, or asymmetrical. A 1 24-segmented, 8-9
r:s curtus :! Y1xl Enp with 7 setae: Scolecithricel/a abyssal- segments fused. A2 Exp more than twice as long as
is (Giesbrecht, 1888) (Fig. 7.203) Enp; Exp 7-segmented, Expl without setae and Exp2
rr.:!J. 3-6 with 3 setae (D . durani, D. helenae) . Mxl Lil with 8
- ........ .3 lales (unknown for S. pseudoculata; the description terminal setae; Li2 with 2 setae; Li3 with 3 setae; Li4
\Olili a :::5. profunda is not enough complete to include the with 3 setae; Enp with 9 setae; Exp with 8 setae; Le
Satpho- ~i es in the key) with 8 setae (D. hibernica). Mx2 terminal part with
P5 uniramous: Scolecithrice/la ovata (Fig. 7.206) sensory filaments: with 3 worm-like and 5 brush-like
irh a ;! P5 biramous ..................................................... ..2 sensory setae (D. hibernica and D. helenae); Li I-Li4
'Salpho- Left P5 Exp3 spiniform distally with swollen each with 4, 3, 3, 3 setae; LiS with 4 setal elements: I
base, longer than combined length of Expl plus sensory appendage, I spine and 2 setae (D. hiberni-
Exp2: Scolecithrice/la minor (Fig. 7.205) ca). Mxp C with 1 proximal worm-like seta with
~ us Left P5 Exp3 of diffe rent shape and much shorter setules along its length, with 3 groups of setae: I
............. ............. ..... .......................... ..... ............. ...3 worm-like plus 1 sclerotized setae, 1 worm-like plus 2
r=. \\ide. Right P5 Exp terminated by characteristic sclerotized setae and 3 distal setae (from proximal to
par- grooved structure: Sco/ecithricella vittata distal); B with medial row of denticles and 3 medial
Right P5 Exp not terminated by characteristic setae; Enp2-Enp6 with 4, 4, 3, 4 and 4 setae (D. hiber-
c::. long. grooved structure, but in spiniform projection nica and D. helenae). PI Enp with reduced outer lobe.
rllllrnus ... .. ................. .................................... ............ .... .4 P 1-P4 surfaces with or without spinules. Spine and
P2 Exp 1 outer spine straight: Scolecithricel/a seta formula:
tenuiserrata (F ig. 7.209)
P2 Exp I outer spine curved ........................ ....... 5 coxa basis cxopod segments cndopod segments
Left P5 Exp3 with long ha irs terminally :
Leg 1 0-0 0-1 1-0; 1-1; 1,1,3 0,2.3
~:t Park. Scolecithricella abyssalis (Fig. 7.203) Leg 2 0- 1 0-0 1- 1; 1- 1; Jll,T,4 0- 1; 1,2,2
Left P5 Exp3 nude: Scolecitlrricel/a dentata Leg 3 0- 1 0-0 1- 1; 1-1 ; lll.I,4 0-1 ; 0-1 ; 1,2,2
Fig. 7.204) Leg4 0- 1 0-0 1- 1; I-1 ; fTl.1 ,4 0- 1; 0- I; 1,2,2
to species of Scolecithrix P5 absent.

f"anales
Posterior corners of Pr synunetrical. P5 absent: Males
:JJ\mng Scolecithrix danae (Lubbock, 1856) (Fig. 7.212) Ce and Pdl separate, Pd4 and 5 separated completely,
L~ Lt5 Posterior comers of Pr asymmetrical, prolonged or incompletely. R is plate with filaments. A 1 left 24-
, on right into lobe extending nearly to CR. P5 ves- segmented, right 23-segmented. Oral parts reduced
---··-
tigial, lamelliform, one-segmented: Scolecithrix compared with femal e. P l -P4 as in females. P5 unira-
bradyi Giesbrecht, 1888 (Fig. 7.211) mous, large, complicated, with large B; right leg 3-
--·--3 segmented, terminal pa11 with digitiform process, left
.wes leg 4-segmented .
Right P5 Exp te1minal segment di vided into 2 sub-
equal branches: Scolecithrix bradyi (Fig. 7.211 )
---·-•
934 Copepoda
This family contains the genera Diaixis Sars, 1902 with 3-4 setae; Li4 usually with 3-4 setae; Enp usual-
and Anawekia Othman and Greenwood, 1994. Only ly with 6-7 setae; Exp with 2 setae ( Undinella), or 3-
Diaixis is recorded in the South Atlantic. 7 setae (5 setae in Tharybis macrophthalma); Le I
usually with 6-7 setae (with 9 setae in Tharybis mac-
Key to species of Diaixis rophthalma). Rythabis strongly deviates in setation of
Females Mx I. Mx2 Lil with 4 setae; Li2-Li5 with 3, 3, 3 and
1 Posterior comers of Pr symmetrical, rounded 4; sensory setae may be present on Li5 (Parundinella ,
......................... .. ............................. ... ............... .2 Rythabis); Enp (terminal part) of limb in type spec ies
Ia Posterior corners of Pr asymmetrical, extended with 3 worm-like plus 4 brush-like setae plus 1 scle-
into triangular pointed lobe on left and curved rotized setae (Tharybis macrophthalma), or with 3
point on right: Diaixis hibernica tropica worm -like and 5-6 bmsh-like sensory setae (there are
Andronov. 1979 (Fig. 7.2 15) deviations in composition and number of sensory
2 Gns without spines both on left and right lateral setae in Tharybidae; correct setal f01mu la for family
swellings : Diaixis helenae Andronov, J 979 can not be given till revision of all its species). Mxp C
(Fig. 7.214) with 1 proximal seta, then with groups of 2, 3 and 3
2a Gns with spines on lateral swellings (dorsal and sclerotized setae (Tharybis macrophthalma) from
ventral view): Diaixis trunovi Andronov, 1979 proximal to d istal, in other species some setae may be
(Fig. 7.2 16) replaced by sensory setae; B with medial row of den-
tides followed by 3 setae; Enp2-Enp6 with 4, 4, 3, 4
Males (unknown for D. hibernica tropica) and 4 setae respectively (Tharybis macrophthalma).
I Left P5 dista l segment with 2 projections, separ- PI Enp with well-developed outer lobe. Posterior sur-
ated from segment: one is curved, other with faces of P2-P4 may be slightly spinulose. Spine and
excavation distally; d istal part of penultimate seta formula:
segment comb-like: Diaixis trunovi (Fig. 216)
I a Left P5 distal segment with 2 finger-li ke projec- coxa basis exopod segments endopod segments
tions; distal part of penultimate segment not
Leg I 0-0 I(vestigial)- I 1-0; I- I; 1,1,3 0,2,3
comb-like: Diaixis helenae (Fig. 7.2 14) Lcg2 0-1 0-0 1-1; 1- 1; lll,J,4 0- 1; 1,2,2
Leg 3 0-1 0-0 1- 1; 1- 1; 111,1,4 0-1 ; 0-1; 1,2,2
Family Tharybidae Sars, 1902 Leg 4 0- 1 0-0 I- I; l- 1; U!,l,4 0-1 ; 0-1 : 1,2,2
References: Bradford et a/. (1983), Schulz and
Beckmann ( 1995), Ferrari and Markhaseva ( 1996). P5 present, uniramous, consists of basal segment plus
Females one segment (Parundinella); or 3-segmented (Undi-
Ce and Pd1 fused or separate, Pd4 and 5 fused or sep- nella, ThGiybis, Rythabis), terminal segment with 1 or
arate, posterior corners rounded or pointed, symmetri- 2 terminal spines as well as articulated inner edge spine
cal or asymmetrical, may be prolonged into lobes (Tharybis), or with 2-4 short terminal spines and some-
( Undinella). R is bifurcate plate with filaments times outer edge spine ( Undinella ), or with 2 strong ter-
(Parundinella, Undinel/a), or plate with or without minal articulated spines and 2 strong subterminal arti-
filaments. Ur of 4 somites, Gns symmetrical or asym- culated (1 outer and 1 inner) spines (Rythabis).
metrical. AI 23-24-segmented, segments 8-9 fused.
A2 Exp incompletely 7-segmented (6-segmented in Males
Rythabis); slightly more than twice length of Enp Al 21-23-segmented. Oral parts similar to female,
(Tha1ybis, Undinella), about twice length of Enp nevertheless setation may be reduced.
(Parundinella spinodenticulata), B and Enp I with 2 P l-P4 as in females. P5 in Parundinella biramous,
setae; Exp l and Exp2 without setae (Exp2 with 2 asymmetrical; left leg Exp 3-segmented, first segment
setae in some species of Parundine/la). Md with with inner proximal process, Enp 1-2-segmented;
strong gnathobase. Mxl Li I large and extending right leg Exp 2-segmented, Enp in part fused with 8 ;
beyond remainder of appendage (Tharybis), or not in other tharybids P5 usually biramous on left, unira-
enlarged (Undinella) , with 11 -14 terminal spines (9 mous on right; left leg with 2 basal segments, Exp 3-
terminal and 4 posterior setae in the type species segmented, shorter than !-segmented Enp; P5 right
Tharybis macrophthalma); Li2 with 2-3 setae; Li3 leg 3-segmented.
Copepoda 935
~ i"Elil~ Tharybidae includes the genera Thatybis absent. Oral pmts identical to that of fema le. A 1
5.!=5. !902: Undine/fa Sars, 1900, Parundinella slightly asymmetrical, left 21 -23-segmented; 8-9 and
?l:-:-·nger. 1957 and Rythabis Schulz, 1995. 10- 12; 13- 14 segments may be fused, right 20-23-seg-
mac- ~phaenna Sars, 1902 and Neoscolecithrix Canu, mented; 8-9, 20-21 segments may be fused; well
l:!tioo of - ":\ere tentatively placed in the family (Bradford equipped with aesthetascs on both sides. A2 B with 2
:. 3 and l!!i - - ; 983 ). Recently Schulz and Beckmann ( 1995) setae; Enp 1 with 2 setae, Enp2 without distinct inner
r:dinella . =-:sed more likely placement of Neoscolecithrix lobe; Exp 6-segmented, Expl wi th I seta, Exp2 with
~~ies ~ Phaennidae, or even more probably, the 2-3 medial seta p lus I distal marginal seta.
~ : scle- 5....--okcirrichidae. Only Tharybis is recorded in the Mdp Enp 1 with 3 setae, Enp 2 with 9 te rminal and 1
r -,th 3 _ .....n Atlantic. appendicular setae. Mx1 Li2 with 2 setae, Li3 with 3
.:.ere are setae; Exp with 8 setae. Mx2 Li1 w ith 5 setae, Li2-Li4
s.ensory ro species of Tharybis with 3 setae each. PI Enp without outer lobe. Left P5
frunjJy es greatly elongated, uniramous, 5-segmented. (M. kauf
l. Mxp C Gos symmetri cal. Subapical atticulated spine of mani, M. nanseni) (Fosshagen, 1978; Andronov.
3 .and 3 P5 extending well beyond remaining terminal 1973a); right leg shott, not extending beyond Exp I of
u) from .spines; distal end not bifurcate: Tltarybis mega- left leg.
!'Gl:!Y be Iodactyla Andronov, 1976 (Fig. 7 .2 18)
rof den- Gns asymmetrical, more swo ll en on right. Monotypic family containing Mesaiokeras Matthews,
- .!_ 3. 4 Subapical articu lated spine does not extend bey- 1961.
:alma). ond remaining terminal spines; distal end bifur-
~sur- cate: Tltarybis asymmetrica Andronov, 1976 Key to species of Mesaiokeras
~and males unknown) (Fig. 7 .2 17) Females (unknown forM. mikhailini)
1 Inner marginal seta present on P 1 B. Posterior
::=-._.·Jy :.\lesaiokeratidae Matthews, L96 J comers of Pr equal in length, but slightly asym-
'="anales metrical in shape. R present, small, thin:
-.:- :md 5 fused or sepa rated, posterior corners of last Mesaiokeras semiplenus Andronov, 1973 (Fig.
sxcire of Pr symmetrical, or slightly asymmetrical. R 7.220)
•'-
.
~ ...
::&.~t. or present as sing le small rostral fi lament. Gns
~==netrica l , or asymmetrical. Ur of 4 somites. A I
la Inner marginal seta absent on PI B. Posterior cor-
ners of Pr symmetrical. R absent: Mesaiokeras
.:.=-: .!..segmented, short, not reaching the end of Pr, tantillus Andronov, 1973 (Fig. 7.221)
•:• one may be longer than left. Oral parts and P 1-
- - ~ee in most details with those of male (these are Males
. '
.~,or ~-ribed in more detail below (Fosshagen, 1978, for 1 P 1 Exp2 with outer spine. Left P5 penultimate
~spine Jl. ~.::11~finani) . PI Enp without outer lobe, sometimes segment (segment 4) with 19 thin comb-like spi-
1r. some- ..-c.:h reduced number of setae on external margin. nules; 3-segmented on right, distal segment with
~;~er iJL;;;aJ parts ofP2-P4 Enp may be covered with spinu- 2 apical spines: Mesaiokeras mikhailini
.::::.=.. ani- ""' on posterior surfaces. Spine and seta formula: Andronov, 1995 (Fig. 7.2 19)
~ 1a PI Exp2 without outer spine. Left P5 without
coxa basis exopod segments endopod segments comb-like spinul es on penultimate segme nt
(segment 4); P5 !-segmented on right, tap ering
~~ 0-0 0- l/0 0/l-0; 0/1-l ; l/ll,J,3 0,2.3
~
0-1 0-0 1- 1; 1-1 ; 111,1,4 0-J; 1,2,2

with I small internal spinule ................. ............ 2
_.::; -
~3 0-1 0-0 I-I; 1- 1; 111,1,4 0- 1; 0-1 ; 1,2,2 2 P5 penultimate segment (segment 4) with shott
~us. ~; - 0-l 0-0 1-1; I-I ; lll,1,4 0-1; 0-l; 1.2,2 stout spine distally; d istal end of segment 5 with
~ent long finger-like projection with small spinule
~ted: ~ .small, symmetrical, 2-segmented. sometim es lac- and 2 adjacent apical spi nes: Mesaiokeras
fwr.b B: ~g on right. semiplenus (Fig. 7.220)
tL.. enira- 2a P5 penultimate segment (segment 4) without any
'- E-q~3- lales spine distally; d istal end of the segment 5 wit-
~ ri;ht ~-somite ofPr symmetrical, or slightly asymmetri- hout finger-like projection, but with 2 long spi-
~ R present, in form of small thin fi.lament, or nes: Mesaiokeras tantillus (Fig. 7.22 1)
936 Copepoda
Family Arietellidae Sars, 1902

coxa basis cxopod segments endopod segments
References: Ohtsuka et a/. ( 1994).
Females Leg I 0-1 l-1 I- 1;1-1; ll/1,1,4 0- 1;0-2; I ,2,2
Total length 0.80 - 7.00 mm. Body relatively robust, Leg 2 0-1 0-0 T- 1;1-1 ;Ill,1,5 0-1;0-2;2,2,3/4
Ce and Pd1 separated or weakly fused, Pd4 and 5 Leg 3 0-1 0-0 l-1 ;1-l;lll,I,S 0-1;0-2;2,2,3/4
fused, sharply or weakly produced, with or without Leg 4 0-0/1 1-0 1-1;I-I ;111,1,5 0-1;0-2;2,2,2/3
dorsolateral and/or ventrolateral processes. Ce roun -
ded or pointed anteriorly, R produced ventrally with P5 variable, not natatory, almost symmetrical ; C and
pair of filaments. Ur relatively short, of 4 somites; intercoxal sclerite separate or fused; B and Enp sepa-
Gns with single paired gonopores and copulatory rate or fused; Enp with 0-4 setae; Exp 1-3-segmented
pores; gonopore(s) located ventrolaterally or ventral- or completely fused with B, carrying 0-5 spines and
ly, with or without opercular plate; copulatory pore setae.
sharing common opening with gonopore or separate
with gonopore located ventromedially or posteriorly, Males
rarely ventrally on right side; seminal receptacles usu- Body similar to female, but U r of 5 somites. Left Al
ally paired, rarely left receptacle entirely lacking. geniculate, 16-20-segmented, proximal segments
Egg-sac present or absent. CR symmetrical or slight- often with row of setules along the posterior margins.
ly asymmetrical, relatively short with up to 7 setae. Mouthparts and Pl -4 s imilar to female or with slight-
A I symmetrical or asymmetrical, longer on left than ly different armature of A2 Enp2 and Md Exp 1. P5
on right, 16-22-segmented, sometimes differing in variable, not natatory, almost symmetrical to strongly
fusion pattern and armature. A2 C without seta; B asymmetrical; C and intercoxal sclerite fused to form
with outer edge seta; Expand Enp separated from B; common base or separate; right B sometimes fused
Enp 2-segmented, segment 1 with 0- 1 seta at mid- with C; right Enp 1-segmented, bulbous or absent;
length, segment 2 with 1-3 inner setae medially and 5 right Exp distinctly or indistinctly 3-segmented, seg-
or 6 setae terminally; Exp indistinctly 6- 10-segmen- ments I and 2 each with 1 seta on outer margin (rare-
ted. Md gnathobase well chitinized, with 3 or 4 sharp ly segment 1 unarmed), segment 2 with tuft of setules
teeth; Enp rudimentary, (-segmented with 1 or 2 setae on inner distal angle of segment 2, segment 3 with 0-
terminally or completely absent; Exp 5-segmented, 3 spines and setae terminally; left Enp 1-2-segmented,
segment 1 with normal or reduced setae, segments 2- unarmed or completely absent; left Exp distinctly 3-
4 with 1 seta each, segment 5 carrying 2 setae, one of segmented, segments 1 and 2 each with seta on outer
which is sometimes vestigial. Mxl Li I 0-6 spines and margin, segment 3 with 1-3 spines and setae terminal-
setae; Li2 with 1 seta or una1med; Li3 represented by ly.
I seta which may be vestigial or absent; Enp bulbous,
(-segmented, with 0-3 setae or completely incorpora- There are now eleven genera in this family. Arietellus
ted into B; Exp lobate, with 3 setae; Lel with 5-9 Giesbrecht, 1892; Campaneria Ohtsuka, Boxshall,
setae. Mx2 well developed; Li I with 1-3 setae (I ves- and Roe, 1994; Crassarietellus Ohtsuka, Boxshall,
tigial), Li2 with 1-2 setae; Li3 and 4 w ith 2 setae each; and Roe, 1994; Metacalanus Cleve, 190 1;
Li5 with stout spine, spinulose or bare; Enp 4-seg- Paraugatiloides Ohtsuka, Boxshall, and Roe, 1994;
mented with I, 3, 2, 2 setae respectively. Mxp elon- Paraugaptilus Wolfenden, 1904; Pilarella Alvarez,
gate, C with I medial and 2 terminal setae; B with 1985; Paramisophria T. Scott, 1897; Rhapidophorus
patches of setules or spinules and 2 setae medial ly; Edwards, 189 1; Sarsarietellus Campaner, 1984; and
Enp 6-segmented, segment I almost fully incorpora- Scutogerulus Bradford, 1969. Ariete!Lus,
ted into B with 1, 4, 4, 2-3, 2-3 , 4-2 setae, outermost Paramisophria, Paraugaptilus, and Pilarel/a occur in
terminal setae of segment 6 may be reduced. P l-4 dis- the South Atlantic.
tinctly 3-segmented, rarely with Enp segments of PI
incompletely fused. Spine and seta formula:
Copepoda 937
~ecently an excellent revision has been made of this 3 Posterior corners of PdS symmetrical .... .. ........ .4
:3mily which describes new genera, some accommo- 3a Posterior corners of PdS asymmetrical; that on
dating already known arietellids, which treats charac- left larger: Arietellus giesbrechti Sars, 190S
~ rransformations of the genital system and appenda- (Fig. 7.223)
ges in detail, and includes phylogenetic relationships 4 Caudal setae very long, plumose; posterior cor-
::med on a cladistic analysis (Ohtsuka et al., 1994). ners of PdS divergent... ................ .. ................ .... S
~e above definition excludes Rhapidophorus 4a Caudal setae short; posterior comers PdS not
J: C and ~dwards, 1891 which is inadequately described. divergent ..... .. ........................................... .......... 6
~ sepa- _-ifietellids are widely distributed from neritic (inclu- S Ce with short spine: Arietellus setosus
~ted 2mg cave-dwelling species) to oceanic waters and ran- Giesbrecht, 1892 (Fig. 7.227)
~and ging vertically from the epipelagic to bathypelagic Sa Ce with long spine: Arietellus aculeatus (T.
~-perbenthic layers (Ohtsuka et al., 1994). Scott, 1894) (Fig. 7.222)
6 Pr elongate, almost 3 times as long as wide; A1
~ey to Arietellidae genera (and some species: extends to posterior border ofPr; CR longer than
~.=iL-\l •-::J:JTamisophria, Paraugaptilus, Pilarella) anal somite : Arietellus plumifer Sars, 190S (Fig .
~ems Females 7.226)
::zgms. Left and right Al of equallength ...................... 2 6a Pr squat, 2.2S times as long as wide; Al extends
::. sligbr- b Left and right Al ofunequallength ...... ............ 3 at least to posterior border of anal somite; CR
x;;~. P5 P4 with vestigial inner coxal seta; PS Exp s horter than anal somite: Arietellus minor
so:ongly absent: Paraugaptilus buchani Wolfenden, Wolfenden, 1911 (Fig. 7.224)
IOrnl 1904 (Fig. 7.230)
!'S fused P4 w ithout inner coxal seta; PS Exp present but Males
~m: unsegmented: Arietellus 1 Posterior corners of Pd5 rounded; caudal setae
a:_ s.eg- P5 w ith Enp; Enp and Exp apparently fused with not spread out: Arietellus simplex (Fig. 7.228)
1:1 :are- B, Enp with 2 setae: Paramisophria giselae I a Posterior corners of Pd5 pointed ....................... 2
~s..~..Iles (Campaner, 1977) (Fig. 7.229) 2 Posterior corners of PdS symmetrical ............... 3
.<h 0- P5 without Enp, represented by 1 seta: Pilarella 2a Posterior corners of PdS asymmetrical; that on
~ned. longicornis A lvarez, 198S (Fig. 7.231) left larger: Arietellus giesbrechti (Fig. 7.223)
~~- 3- 3 Ce, viewed dorsally, prolonged into long spine
LO'.Iter _ Wes (unknown for Pilarella longicornis) .......................................................................... .4
~- Right PS Enp L-segmented ................................ 2 3a Ce, viewed dorsally, conical, w ithout spine ...... S
L Right PS Enp absent: Paramisophria 4 Ce with short spine: Arietellus setosus (Fig.
P5 Exp2 on both sides enlarged; left leg with 7.227)
-ze;zlfus strong pincer-like spine and weaker process of 4a Ce with long spine: Ariete/lus aculeatus (Fig.
crxsaall. same length, bifid at tip; Enp not distinctly hi- 7.222)
Ehall. lobed: Paraugaptilus buchani (Fig. 7.230) 5 Pr elongate, almost 3 times as long as wide; CR
1901: P5 Exp2 only slightly enlarged; left leg termina- longer than anal somite: Arietellus plumifer
~ 199~: ting in 2 ordinary subequal spines; left Enp usu- (Fig. 7.226)
ally distinctly bilobed: Arietellus Sa Pr squat, 2.25 times as long as wide; CR shorter
YJI:oru.s than anal somite: Arietellus minor (Fig. 7.224)
¢: :md r.- • ro the species of Arietellus
c.zlf':JS. 'Temales Family Augaptilidae Sars, 1905
x:cmin Posterior corners of PdS rounded ...................... 2 Females
Posterior corners of PdS pointed ....................... 3 Total length 2.1-9.6 mm. Body usually slender, Pr
Caudal setae spread out in all directions; anteri- oval, Ur relatively small proportion of total length, Ce
or Ce w ithout point: Arietellus pavoninus Sars, and Pdl separated, Pd4 and S fused, posterolateral cor-
1905 (Fig. 7.225) ners and anterior Ce usually rounded but sometimes
Caudal setae not spread out; anterior Ce, viewed pointed. R filaments slender or absent, R sometimes
laterally, with small point: Arietellus simplex represented by small !mob. Ur usually short, of 3-4
Sars. 1905 (Fig. 7.228) somites; Gns with ventral swelling, CR syttu11etrical
938 Copepoda
with 6 setae. Al symmetrical, 24-25-segmented. A2 C Males

with 1 outer seta; B usuall y elongate with 1-2 outer Body similar to female, but Ur of 5 somites. Left Al
edge setae; Exp and Enp separated from B; Enp 2-seg- geniculate, ?2 1-22-segmented sometimes geniculate on
mented, usually but not always much longer than Exp, tight Mouthpatts and P 1-4 similar to the female. P5
Enp I wi th 1-2 setae placed towards the distal end, with 3-segmented Enp and Exp on left and right, slight-
Enp2 with 4-7 i1111er setae medially and 6-7 setae ter- ly asymmetrical; right Exp2 often with inner expansion
minally; Exp 5-8-segmented, with varying numbers of of variable shape but usually in form of thick spine.
setae: from 4 tetminal setae, to every segment with at
least 1 seta. Md gnathobases more usually symmetri- The following genera are now in this family:
cally developed on each side with 2-7 (or more) short Augaptilina Sars, 1920; Augaptilus Giesbrecht, 1889;
or long teeth set either obliquely or more or less at Centraugaptilus Sars, 1920; Euaugaptilus Sars, 1920;
right angles to axis of gnathobase, in Pachyptilus and Haloptilus Giesbrecht, 1898; Heteroptilus Sars, 1920;
Heteroptilus gnathobases asymmetrica lly developed; Pachyptilus Sars, 1920; Pontoptilus Sars, 1905; and
Enp usually 2-segmented, sometimes 0- or !-segment- Pseudaugaptilus Sars, 1907 of which Augaptilina,
ed, with varying number of setae; Exp usually 4-5-seg- Heteroptilus and Pseudaugaptilus have not been
mented with varying numbers of setae. Mx 1 variable recorded from the South Atlantic.
from full complement of lobes, with reduced setation
distally in Haloptilus, Pontoptilus, Pachyptilus, and Matthews ( 1972) made an excellent review of the spe-
Heteroptilus, to extreme but variable reduction in at cies of Euaugaptilus, Augaptilus, Haloptilus, and
least outer and terminal lobes. Mx2 Li 1-4 small and Pseudaugaptilus known to that date. Data on the dis-
usua lly furnished with long setae, carryi ng characteris- tribution of setae on Mx l in particular, used in the
tic shield-shaped appendages or, in case of keys, are from his Table 5 and might not always agree
Augaptilina, Li5 and Enp carries dense elongate with the figures reproduced here.
bunches of setae resembling brush. Mxp C with 3-10
setae; 8 slender or squat with 2 setae medially; Enp 6- Key to genera and species of Augaptilidae
segmented, sometimes elongate, Enpl almost fully Females
incorporated into B with 1-2 setae, Enp2-5 with 3-4, 3- I Ur of 4 somites .......... ........................................ 2
4, 2-4, 1-4 setae respectively, Enp6 with 2-4 setae the la Ur of 3 somites ............ ..... .. ............ .................. .4
outermost 1-3 of which are reduced. P l -4 with Exp 3- 2 P5 Enp 1-segmented: Pachyptilus (p. 942)
segmented, Enp usually 3-segmented except for P5 in 2a P5 Enp 2-3-segmented ....................................... 3
Pontoptilus and few Euaugaptilus where it is 2-seg- 3 P5 Enp 2-segmented; R absent: Pontoptilus ova-
mented and in Pachyptilus and Heteroptilus where it is lis Sars, 1907 (Fig. 7.282)
!-segmented; occasionally P 1 Enp 2-segmented (some 3a P5 Enp 3-segmented; R present: Haloptilus (p.
Euaugaptilus, Heteroptilus, Pachyptilus, and Augapti- 941)
lina). Spine and seta formula: 4 Mx1 reduced to a 3-segmented straight limb:
Augaptilus (p. 939)
coxa basis exopod segments endopod segments 4a Mxl better deve loped ........................................ 5
5 Md gnathobase with numerous teeth often cut
Leg I 0-l 1-0 1- l ;OII- I; 0/1/ ll,l,4 0-1;0-1 /2; 1/2,2,112
Leg 2 0- l 0-0 1-1 ;I-1 ;IU ,1,5 0-1 ;0-2;2,2,3 obliquely. R more usually small or absent, occa-
Leg 3 0-1 0-0 OII-1;1- l;Tn,I,S 0-1 ;0-2; 1/2,2.3/4 sionally in form of 2 long fine filaments:
Leg 4 0-1 0-0 1-l ;T-l;lll,l,5 0-1 ;0-2;2,2,3/4 Euaugaptilus (p. 939)
Leg 5 0-0 0 1-0 0/1-0;0/T-0/ l ;0/1/11,1,3/4 0-0/1;0- l/2; 5a Md very elongate, with 2 thin, curved teeth eit-
1/2,2, 112/3
her side of smaller tooth. R strong, bifurcate,
projecting forwards: Centmugaptilus (p. 939)
P5 natatory, symmetrical, very similar to other swim-
ming legs but smaller than P2-4; i1111er edge seta on Males (unknown for Pontoptilus ova/is)
Exp2 often spine-like and articulated with its seg- 1 Setae on Mx2 and Mxp coiled distally, and car-
ment. rying special shield-shaped appendages. R
strong, bifurcate, projecting forwards. Body
squat: Centraugaptilus (p. 939)
Copepoda 939
This combination of characters absent.. ............ 2 2 Female P5 Enp3 w ith 7 setae; A I extends be-
l.s.:.-\ 1 Right P5 Exp2 without inner spine; without yond posterior border of Pd I. In both sexes
t:k:>OD shield-shaped appendages on setae of Mx2 and bifurcate R relative ly small: Centraugaptilus
~e. p: :\lxp: Haloptilus (p. 94 1) cucullatus (Sars, 1905) (Fig. 7.23 7)
:;:_ shght- Right P5 Exp2 with at least l inner spine, some- 2a Female P5 Enp3 with 6 setae; A 1 does not
?'"""SlOll rimes small; shjeld-shaped appendages frequent- extend to posterior border of Pd I . In both sexes
~- ly on the setae of the Mx2 and Mxp ... ............. .3 bifurcate R relatively large: Centraugaptilus
:\1xl reduced to 3-segmented straight limb: horridus (Farran, 1908) (Fig. 7 .238)
izm.ily: A ugaptilus (p. 939)
- I 89: :\lx I well developed: Euaugaptilus (p. 939) Key to species of Euaugaptilus
s. 19~0: Females
s. :920: ~ ~ to species of Augaptilus 1 A2 Exp with 5 or fewer setae ......... ................... 2
5: and F emales 1a A2 Exp with 7 or more setae ............................. 9
y:ilina. Anterior Ce pointed: Augaptilus spinifrons 2 Ce in lateral view with prominent hump associa-
x been Sars, 1907 (Fig. 7.236) ted with musculature of A2: Euaugaptilus gib-
Anterior Ce not pointed ..................................... 2 bus (Wolfenden, 1904) (Fig. 7.250)
Gns very long, equal to at least half length of U r: 2a Ce in lateral view without such hump ............... 3
~spe Augaptilus megalurus Giesbrecht, 1889 (Fig. 3 Mxl Exp bearing 2-4 setae .............................. .4
llS". and - .235) 3a Mx l Exp bearing 6 or 7 setae ........................... 7
.:..e dis- Gns shorter................ ................................ .. ....... 3 4 Mxl Exp wi th 2 setae, Lel with 7 setae:
: i::i the CR covered in small spinules distally: Augaptilus Euaugaptilus oblongus (Sars, 1905) (Fig.
"S :!giCe longicaudatus (Claus, 1963) (Fig. 7.234) 7.258)
CR smooth .................. .... .. ................... ............. .4 4a Mx1 Exp with more than 2 setae, Lel with a dif-
CR 3 times longer than wide, not divergent, anal ferent number of setae ...................................... .5
somite equal in length to preceding somite: 5 Mx 1 Exp with 3 setae, Le 1 with 5 setae; P1
A ugaptilus anceps Fanan, 1908 (Fig. 7.232) Exp2 and 3 with rounded expansion at base of
CR 5 times longer than wide, divergent, anal outer edge spines: Emmgaptilus longimanus
---·-·..1 somi te much longer than preceding somite: (Sars, 1905) (Fig. 7.254)
Augaptilus glacialis Sars, 1900 (Fig. 7.233) 5a This combination of characters absent.. ............ 6
- .......3 6 Mx J Exp with 3 setae, LeI with 8 setae; distal
m Ol·a- bJes end of Md gnathobase cut obliquely with long
Right P5 Exp2 with 2 dissimilar inner points: teeth: Euaugaptilus magnus (Wolfenden,l904)
(p. A ugaptilus megalurus (Fig. 7.235) (Fig. 7.255)
This segment w ithout points .............................2 6a Mx I Exp with 4 setae, LeI with 9 setae; distal
limb: Ri ght P5 Exp3 terminated by long stylet: end of Md gnathobase hardly cut obliquely and
Augaptilus longicaudatus (Fig. 7.234) with short teeth: E uaugaptilus austrimts Park,
- .... 5 Terminal spine of this segment norma1. ............ 3 1993 (Fig. 7.244)
~em Left P5 Exp3 with tenninal spine eq ual to 1/3 - 7 Ce in dorsal view w ith obvious indentation poste-
1 .t length of subapical spine: Augaptilus gla- rior to A2; Md palp almost as long as gnathobase;
cia/is (Fig. 7.233) g nathobase expanded d istally: Euaugaptilus
Left P5 Exp3 with terminal spine equal to 1/5 - antarcticus (Wolfenden, 1911) (Fig. 7 .243, see
r6 eir- l 6 length of subapical spine: Augaptilus spini- also Park, 1993)
~e- frons (Fig. 7.236) 7a Ce in dorsal view without obvious indentation
39) posterior to A2 ; Md palp shorter than gnatho-
ro species of Centraugaptilus (female and male) base; gnathobase tapers distally ........................ 8
In lateral view anterior Ce in form of triangular 8 Pr more than 3 times as long as wide; R with 2
projection; R not visible in dorsal view ............ 2 fine fila ments: Euaugaptilus laticeps (Sars,
In lateral view anterior Ce round in form ; cons- 1905) (Fig. 7.252)
picuous, bifurcate R projecting forwards and 8a Pr less than 3 times as long as wide; R 2-pointed:
'iSible in dorsal view: Centraugaptilus rattrayi Euaugaptilus squamatus (Giesbrecht, 1889)
T. Scott, 1894) (Fig. 7.239) (Fig. 7.263)
940 Copepoda
9 R without filaments ......................................... 10 20 PI Enp 2-segmented: Euaugaptilus palumbii

9a R with pair of filaments or fine points ............ 12 (Giesbrecht, 1889) (Fig. 7.260)
10 Mxl Enp with 2 setae, Exp with 3 setae, Lei with 20a PI Enp 3-segmented ..................... ................... 21
8 setae: Euaugaptilus nodifrons (Sars, I905) 21 Mx I Li4 with I or 0 setae ............................... 22
(Fig. 7.257) 21 a Mx I Li4 with 3 setae: Euaugaptilus tenui-
I Oa Mxl Enp without setae, Exp with different num- spinus Sars, I920 (Fig. 7.264)
ber of setae, Le 1 with 4 setae .......................... 1I 22 Mxi Li4 without setae ......................... ...... ...... 23
11 Mx1 Exp with 2 setae: Euaugaptilus latifrons 22a Mxl Li4 with I seta ............ .... .................. ...... 24
(Sars, 1907) (Fig. 7.253) 23 Mx I Exp with 2 setae, Le 1 with 3 setae:
11a Mx1 Exp with 8 setae: Euaugaptilus angustus Euaugaptilus rigidus (Sars, 1907) (Fig. 7.262)
(Sars, 1905) (Fig. 7 .242) 23a Mx1 Exp with 3 setae, Lei with 5 setae:
12 P3 and 4 Exp2 and 3 extending distolaterally Euaugaptilus clavatus (Sars, 1907) (Fig. 7 .246)
into outer rounded lobes: Euaugaptilus facilis 24 Mxl Lel with 2 setae, Li2 with I seta:
(Farran, 1908) (Fig. 7.248) Euaugaptilus ajjinis Sars, 1920 (Fig. 7 .240)
12a P3 and 4 Exp2 and 3 not extending distolateral- 24a Mx 1 LeI with 4 setae, Li2 absent: Euaugaptilus
ly into outer rounded lobes .............................. 13 bullifer (Giesbrecht, 1889) (Fig. 7.245)
13 P5 B distolateral outer seta long, extending well
beyond distal border of Exp3 ............... ...... .. ... 14 Males (unknown for E. affinis, E. aliquantus, E. cla-
13a P5 B distolateral outer seta short, not extending vatus, E. latifi'ons, E. pachychaeta, E. perasetosus, E.
beyond distal border of Exp3 ................. .. ....... 16 tenuispinus)
14 P3 Expl without outer spine; Mxl Enp absent, I Prehensile AI on right.. ..................................... 2
Lei with 6 setae: Euaugaptilusfiligerus (Claus, I a Prehensile AI on left (presumed, for some spe-
1963) (Fig. 7.249) cies, which have not been described completely)
I4a P3 Expl with outer spine; Mxl Enp present, Le 1 ........................................................................... 3
with 9 setae ...................................................... 15 2 P5 right Exp3 with l very long terminal seta:
15 In lateral view, genital swelling placed on ante- Euaugaptilus hecticus (Fig. 7.251)
rior part of Gns; CR slightly asymmetrical; Mx I 2a P5 right Exp3 with 4 spines: Euaugaptilus
Enp with 3 setae: Euaugaptilus aliquantus squamatus (Fig. 7.263)
Park, 1993 (Fig. 7.241) 3 P5 right Exp2 with multi-pointed inner projec-
15a In lateral view, genital swelling placed towards tion: Euaugaptilus filigerus (Fig. 7.249)
middle of Gns; CR symmetrical; Mxl Enp with 3a P5 right Exp2 with only I inner projection which
2 setae: Euaugaptilus perasetosus Park, 1993 may be bifurcate at its tip ................................. .4
(Fig. 7.261) 4 P5 right Exp2 and 3 fused: Euaugaptilus
16 Mxp B with 2 proximal, unusually developed palumbii (Fig. 7.260)
setae which are short, thickened and spine-like: 4a P5 right Exp2 and 3 separate ........................ ..... 5
Euaugaptilus pachychaeta Matthews, 1972 5 P5 right Exp3 terminal spine as long as or longer
(Fig. 7 .259) than its segment.. ............................................... 6
16a Mxp B with proximal setae normal... .. ............ 17 5a P5 right Exp3 terminal spine shorter than its seg-
17 P5 Exp and Enp 2-segmented: Euaugaptilus ment ................................................................... 7
hecticus (Giesbrecht, 1889) (Fig. 7.251) 6 P5 right Exp2 with prominent inner proximal
17a P5 Exp and Enp 3-segmented ......................... 18 pointed projection: Euaugaptilus elongatus
I8 Mx1 Enp with setae ......................................... l9 (Fig. 7.247)
18a Mxl Enp absent ............. .. ....... ......................... 20 6a P5 right Exp2 with inner proximal rounded pro-
19 Mx 1 Li4 with 5 setae, Enp with 5 setae, Exp jection: Euaugaptilus bullifer (Fig. 7.245)
with 8 setae, Lei with 9 setae: Euaugaptilus 7 P5 right Exp3 with only 1 terminal spine; also
maxillaris Sars, 1920 (Fig. 7.256) with 2 outer edge spines: Euaugaptilus nodi-
19a Mx I Li4 with 3 setae, Enp with 3 setae, Exp frons (Fig. 7.257)
with 6 setae, Lei with 8 setae: Euaugaptilus 7a P5 right Exp3 with 2 tem1inal spines (one may
elongatus (Sars, 1905) (Fig. 7.247) be slightly subterminal) and 1 outer edge spine
........................................................................... 8
Copepoda 941
wbu.bii P5 right Exp3 w ith 1 inner edge spine ....... .. ..... 9 2a Mx l Enp with 2 setae, Exp w ith 10 setae, Lel
P: right Exp3 without inner spine ................... II w ith 9 setae: Haloptilus ornatus (Giesbrecht,
___ .11 Right P5 Exp3 2 terminal-most spines subequa l: 1892) (Fig. 7 .276)
Euaugaptilus facilis (Fig. 7 .248) 3 Anterior Ce without cephalic spine, rostra l fila-
lDIUi- Right P5 Exp3 2 terminal-most spines unequal ments present ........ ...... .. .... ................................ .4
..... .... .......... ............. ............ ............ ................. 10 3a Anterior Ce with cephalic spine and rostral fila-
..,., Right P5 Exp3 subterminal inner edge spine 2.5 ments ...... .......................................................... ! 0
--·- -'
___ .2..! times longer than terminal spine: Euaugaptilus 4 Mx I Enp with more than I 0 setae .................... 5
setae: longim anus (Fig. 7.254) 4a Mx I Enp with less than I 0 setae ....... .... .... ........ 6
- .262) Right P5 Exp3 subterminal inner edge spine 5 Mxl Enp with II setae, Exp with 8 setae, Lei
setae: more than 3 times longer than terminal spine: w ith 8 setae; CR twice as long as wide:
. - ..2..!6) Euaugaptilus rigidus (Fig. 7 .262) Haloptilus angusticeps Sars, 1907 (Fig. 7.266)
C seta: P5 right Exp3 with 2 terminal spines of more or 5a Mxl Enp with I 3 setae, Exp with II setae, Lei
!ess equal length .............................................. 12 with 9 setae; CR 1.5 times as long as wide:
~us _ P5 right Exp3 with 2 terminal spines unequal in Haloptilus fons Fa rran, I 908 (Fig. 7.270)
!ength .............. ................ .... ................... .......... 13 6 Pr le ngth:width ratio 5 .I; Mx I Enp with 7 setae:
P5 left Exp3 termi nal seta small, segment roun- Haloptilus chierchiae (Giesbrecht, 1889) (Fig.
E. cia- ded: inner point of right Exp2 simple: 7 .268)
ISm. E. Euaugaptilus mag11us (Fig. 255) 6a Pr length:w idth ratio less than less than 5; Mx1
P5 left Exp3 tenninal seta about 1/3 length of its Enp with less than 7 setae .............. ................... 7
segment, segm ent oval; inner point of right Exp2 7 Pr about 4 times as lon g as wide .................. .. ...8
divided terminally: Euaugaptilus gibbus (Fig. 7a Pr about 3 times as long as w ide ....................... 9
- .250) 8 Mxl Enp with 5 setae, Exp w ith 11 setae, Lei
_ ....3 1..3 ~lx l with Enp with 5 setae: Euaugaptilus with 9 setae: Haloptilus austini Grice, 1959
l!J s.era: ma.:~:illaris (F ig. 7.256) (Fig. 7.267)
_.:-:! ~1xl Enp absent ................ .. .................... ......... l4 8a Mx 1 Enp with 2 setae, number of setae on Exp
rrq:Wus ~lx l Exp with 6 or more more setae ............... I 5 and LeI unknown: Haloptilus major Wolfenden,
~ ~1x l Exp with 4 or fewer setae ....................... 17 I 9ll (Fig. 7.273)
- ~1x.I Exp with 8 setae; Li3 absent: Euaugaptilus 9 Mx I Exp with 9 setae; P5 B outer edge seta
angustus (Fig. 7.242) more than twice length of Exp: Haloptilus lofl-
t•illch 5:;. ~1xl Exp with 6 setae; Li3 present, with setae g icirrus Brodsky, I950 (Fig. 7.27 1)
_ .•....! ........................ ... .. ............................................ 16 9a Mx l Exp with 7 setae; P5 B outer edge seta
aprilus ~td gnathobase widened distally: Euaugaptilus much shorter than Exp: Haloptilus longicornis
antarcticus (Fig. 7.243) (Claus, 1863) (Fig. 7.272)
o;: ~Id gnathobase tapers distally: Euaugapti/us IO Pr more than 8 times as long as wide: Haloptilus
'lc::ger /aticeps (Fig. 7 .252) ocel/atus Wolfenden, I905 (Fig. 7.275)
- .. 6 ~1x l Li3 present, with 2 setae: Euaugaptilus lOa Pr less than 4.5 times as long as wide ......... .... 11
~ s.eg- oblongus (Fig. 7.258) 1I AI extends beyond CR by 5 or more segments
.:;. ~1x I Li3 present but without setae: ........ .................. .......................................... ..... 12
Euaugaptilus austrinus (Fig. 7 .244) ll a AI extends beyond CR by 3 or fewer segments
....... ......... ......................................................... 13
iLry to species of Haloptilus 12 Ce with relatively short cephalic spine; Mx 1 Exp
'"anales (unknown for H.fertilis) with II setae, Lei with I 0 setae: Haloptilus
:\nterior Ce without cephalic spine or rostral acutifrons (Giesbrecht, 1892) (Fig. 7 .265)
filaments ................ .................................... .... .... 2 l 2a Ce with long cepha lic spine; Mx I Exp with 9
I IUNii- Anterior Ce with another combination of these setae, Lei with 8 setae: Haloptilus oxycephalus
characteristics ..... .. ................. .... ....... ................. 3 (Giesbrecht, 1889) (Fig. 7.277)
~lxi Enp with 4 setae, Exp with 8 setae, Lei 13 A l extends beyond CR by 3 segments; Mx. l Exp
with 6 setae: Haloptilus plumosus (Claus, 1863) with 8 setae; Mx2 proximal Enp segment very
(Fig. 7.278) strongly develop ed compared with remaining
942 Copepoda
setae: Haloptilus spiniceps (Giesbrecht, 1892) 10 Mx I Enp with 3 setae, Exp with 8 setae, Lel
(Fig. 7.279) with 8 setae: Haloptilus spiniceps (Fig. 7.279)
13a AI extends beyond CR by I segment; Mxl Exp 1Oa Mx 1 Enp with 4 setae, Exp with 11 setae, Le 1
with II setae; Mx2 proximal Enp slightly better wi th 10 setae: Haloptilus acutifrons (Fig. 7.265)
developed than remaining setae: Haloptilus
mucronatus (Claus, 1863) (Fig. 7.274) Key to species of Pachyptilus
Females (males unknown)
Males (unknown for H.fons, H. austini, H. major and 1 A l does not extend to CR; distal setae of Mx2
H. longicirrus) and Mxp thickened and spine-like: Pachyptilus
l P5 C with inner triangular projections on both longimanus (Wolfenden, 1906) (Fig. 7.281)
sides .......... ...... ............ ....................................... 2 Ia Al extends to posterior border of CR; none of
I a P5 C with inner triangular projections on left setae on Mx2 and Mxp thickened more than
side only, or without any projections (H. plumo- remaining setae: Pachyptilus abbreviatus (Sars,
sus?) ................................................. ................. .4 1905) (Fig. 7.280)
2 P5 C each with similar-sized projection ..... ...... 3
2a Right P5 C inner projection twice as long as that Family Heterorhabdidae Sars, 1902
on left: Haloptilus chierchiae (Fig. 7.268) Females
3 P5 right Exp3 term inal spine as long as its joint; Total length 2.0-8.0 mm. Body usually slender, Pr
C furnished with hairs on inner border as well oval, Ce and Pd 1 separated, Pd4 and 5 fused, postero-
triangular projections: Haloptilus angusticeps lateral corners rounded. Ce rounded anteriorly; rostral
(Fig. 7.266) fi laments slender. Ur elongate, of 4 somites; Gns with
3a P5 right Exp3 terminal spine shorter than its ventral process sometimes protruding greatly and
joint; C naked on inner border apatt from trian- sometimes occupying whole length of somite; CR
gular projections: Haloptilus mucronatus (Fig. usually asymmetrical (largest on left) with varying
7.274) degrees of fusion between them and anal somite, with
4 P5 C apparently without inner triangular proces- 7 setae, one of terminal setae often slightly or extra-
ses: Haloptilus plumosus (Fig. 7.278) ordinarily elongate on the left side. A I symmetrical,
4a P5 C with irtner projection on left side ............. 5 25-segmented. A2 B with 2 outer edge setae; Exp and
5 PS Enp hardly extending beyond distal border of Enp separated from B; Enp 2-segmented, Enpl with 2
Exp l on both sides: Haloptilus ocellatus (Fig. distally placed setae, Enp2 with 6-8 inner setae
7.275) medially and 6-7 setae terminally; Exp 7 -9-segmen-
Sa P5 Enp extend beyond distal borders ofExp1 on ted, proximal 2 segments sometimes without setae.
one side at least ........ .................... ......... ........ .... 6 Md gnathobases usually asymmetrically developed on
6 PS right Exp3 terminal spine longer than its joint each side with 3-5 (or more) long, sharp teeth, in
........................................................................... 7 Heterorhabdus, Hemirhabdus, and Neorhabdus one
6a P5 right Exp3 tem1inal spine equal to or shorter tooth set far apart from remaining teeth; Enp 2-seg-
than its joint ................................ ....................... 8 mented, Enp l with 2-4 setae, Enp2 with 6-9 setae ter-
7 P5 left inner coxal projection blunt terminally; minally; Exp 1-4 with 1 seta each, Exp5 carrying 2
right Exp3 proximal spine half length of termi- setae. Mx l tends to have Li2-4 and Enp reduced; Li 1
nal spine: Haloptilus onwtus (Fig. 7.276) with 5-1 3 spines and setae (most reduced setation in
7a P5 left inner coxal projection pointed terminally; Hemirhabdus, least reduced in Disseta); Li2 small or
right Exp3 proximal spine equal in length to ter- absent with 0-2 setae; Li3 small or absent with 0-2
minal spine: Haloptilus fertilis (Fig. 7 .269) setae, Li4 with 1-4 setae; Enp small, !-segmented,
8 Left P5 Exp3 with 2 equal terminal spines: with 4-11 setae; Exp large, with 5- 11 setae (least redu-
Haloptilus longicornis (Fig. 7.272) ced setation in Disseta and Mesorhabdus); Lel with
8a Left P5 Exp3 with subequal spines ................... 9 3-8 setae. Mx2 usually modified as a grasping, pier-
9 Left P5 Enp extends almost to distal border of cing limb (least modified in Disseta and most modi-
Exp2: Haloptilus oxycephalus (Fig. 7.277) fied in Hemirhabdus); Li l and 2 with 3-5 and 0-3
9a Left P5 Enp does not extend beyond middle of reduced setae respectively; Li3 and 4 with 2-3, 2-3
Exp2 ........ ......................................................... 10 usually small setae respectively; LiS with 1-3 setae, I
Cop e poda 943
or 2 of which may be greatly enlarged and spine-like; Key to Heterorhabdidae genera (and some species:
Enpl usually with an enlarged spine-like seta; remain- Disseta, Mesorhabdus, Neorhabdus)
~- J-el der of Enp with greatly redu ced setae. Mxp C with 3- Females
- setae proximal one of which is greatly enlarged and 1 Mx2 with strong spine-like setae; setae of right
spine-like in Heterorhabdus; B slender with 3 small CR subequal.. ..... ............ ................ ................ .... 2
s.:=rae medially; Enp 6-segmented, segment 1 almost Ia Mx2 without strong spine-like setae; one seta on
:cliy incorporated into B with 2 setae, segments 2-5 right CR much longer than others: Disseta
o•-~h2
-ith 3, 3, 2-3, 2-3 setae respectively , segment 6 with palumbii Giesbrecht, 1889 (Fig. 7.283)
iypri!us 3-l setae outermost 1-2 of which are reduced. P 1-4 2 One tooth on Md gnathobase apart from other
., l J ~rinctly 3-segmented, Exp of P2-4 flattened and teeth, is much larger and falciform .......... ... .. ... .3
1100e of
.eaf-like. Spine and seta formula: 2a Teeth on M d gnathobase subequal.. ............. ... ..4
~ dlan 3 Mx.2 with more than 2 spine-like setae; Md gna-
Sars. coxa basis exopod segments endopod segments thobases asymmetrical; Mx l with Li2 but without
Li3; Lei with 5 setae: Heterorhabdus (p. 944)
~-g I 0-1 110-1 1-1 ;1-1 ; Jl. TTJ,l,3/4 0-1 ;0-2; I,2,2
=..:; ~ 0-1 0-0 l-1 ;l-1 ;111,1 ,5 0-1 ;0-2;2,2,3
3a Mx2 with only 2 spine-like setae; Md gnathoba-
:...~ 3 0-1 0-0 l-1;1-I ;Tn,l,S 0- 1;0-2;2,2,3/4 ses almost symmetrical; Mx1 without Li2 and
~-g-t 0-1 1-0 l-1 ;l-1 ;lll,l,S 0- 1;0-2;2,2,3 Li3; Le 1 with only 3 setae: Neorhabdus falci-
de:r. Pr -"'"; 5 0-0 1-0 T-O; l-1;11,1,4 0-1 ;0-1;2,2,2 formis (Wolfenden, 1911) (Fig . 7.298)
JQ:,-..crt)-
4 Proximal lobes of the Mx2 and their setae little
;osnal
?5 natatory, symmetrical, very similar to other swim- developed; P5 Exp2 with posterodistal, toothed
cs "\\ith =::ng legs but smaller; inner edge seta on Exp2 modi- flange: Heterostylites (p. 945)
and Eed into long slender spine articulated with its seg- 4a Proximal lobes of the Mx2 and their setae well
~: CR .:::::ent: Exp2 postetior distal border expanded into developed: Mesorhabdus angustus Sars, 1907
~mg
ro.Jthed flange in Heterostylites. (Fig. 7 .297)
~."\\ith
r :!'Gra-
l ales Males
~~
3ody similar to female, but Ur of 5 somites. Left A I 1 One setae on right CR longer than others:
!xp and ~culate, 2 1-23-segmented. Mouthparts and legs 1- Disseta pa/umbii (Fig. 7.283)
v.rJ:i 2 - sunilar to female. P5 with a 3-segmented Enp and Ia Setae on right CR subequal. .. .... ..... ........ ........... 2
E."tp on both sides, right B usually w ith inner exten- 2 P5 right B with long, narrow internal appenclix,
'S ~ of variable shape and usually fringed by hairs or parallel to Enp, extending almost to distal border
~ules, sometimes with inner extension on left; right of Exp2; right Exp2 very enlarged; both Exp3
Fxp2 w ith inner expansion of variable shape but usu- term inated by long spines: Heterostylites (p. 945)
-- in fmm of thick spine; left Exp3 often but not 2a These characters absent .... ......... ........ .... ............ 3
one ays terminated by an elongate spine. 3 One tooth on Md gnathobase very set apart from
=-~- other teeth, is much larger and falciform ......... .4
il:is family was revised by Heptner (1972) and the 3a Teeth on Md gnathobase subequal: Mesorhabdus
~ redefined based on the structure of the mouth- angustus (Fig. 7.297)
~~la -' .-rc-,s which are re lated to feeding type. Later Grice 4 Mx2 w ith more than 2 spine-li ke setae; Md gna-
1 ~ - 3) added a fmther genus, although it has been thobases asymmetrical; P5 right Exp3 more than
_ provisionally placed in this family pending dis- twice length of Exp2, left Exp 3 tenninated by
~- of the male. As currentl y defined the fami ly very long, curved spine: Heterorhabdus (p. 944)
<!des the following genera: Alrhabdus Grice, 1973; 4a Mx2 w ith only 2 spine-like setae; Md gnathoba-
;;: .-eCu.- fJ!:swra Giesbrecht, 1892; Hemirhabdus Wolfenden, ses almost symmetrical; P5 right Exp3 hardly
: v.i~ I t. Heterorhabdus Giesbrecht, 1892; Heterostylites longer than Exp2, left Exp3 term inated by short
~ .,:~- Sl:s.. 1920; Mesorhabdus Sars, 1905; Microdisseta spine: Neorhabdus falciform is (Fig. 7 .298)
;::;.oCi-
S;;:ner. 1972; and Neorhabdus Heptner, 1972 of
~ Q.-3 _:_-n Alrhabdus, Hemirhabdus, and Microdisseta
~ not been taken in the South A tlantic.
~-1
944 Copepoda
Key to the species of Heterorhabdus 10 Left side of Gns with undulating surface bearing
Females patches of smal l sp inules : Heterorhabdus tan-
I Anterior Ce pointed in dorsa l view : Hetero- neri (Giesbrecht, 1S95) (Fig. 7.293)
rhabdus spinifrons (Claus, 1S63) (Fig. 7.292) 1Oa Left side of Gns without such characteristics; in
1a Anterior Ce row1ded ...... ........... ........ ..... .... ........ 2 dorsal view Gns with both lateral borders undu-
2 PS Exp 1 with inner seta: Heterorhabdus clausii lating, more pronounced on left: Heterorhabdus
(Giesbrecht, 1SS9) (Fig. 7.2S6) austrinus Giesbrecht, 1902 (Fig. 7 .2SS)
2a PS Exp 1 without inner seta ............................... 3
3 Mxp C with median seta short and slender. ..... .4 Males F l!
3a Mxp C with median seta long and thick .. ..... .. .. 6 I Anterior Ce pointed in dorsa l view: Hetero-
4 Body sh01t and squat; U r hardly half length of rhabdus spinifrons (Fig. 7 .292)
Pr; Al does not extend beyond posterior border 1a Anterior Ce round in dorsal view .............. ...... ..2
of Pr; longest terminal seta on left CR shorter 2 Left PS Exp3 oval with short terminal spine .. .. 3
than U r: Heterorhabdus compactus (Sars, 1900) 2a Left PS Exp3 extended terminally into styliform
(Fig. 7 .2S7) process ............................................................... S
4a These characters not combined ....................... ..S 3 Right PS Exp2 with inner rounded projection
s Body oval; Ur longer than half Pr; AI longer bordered by hairs: Heterorhabdus vipera (Fig.
than Pr; longest terminal seta on left CR more 7.294)
than twice length of Ur: Heterorhabdus robus- 3a Right PS Exp2 with inner pointed projection .. .4
tus Farran, 190S (Fig. 7.29 1) 4 R ight P5 Exp2 with broadly based inner pointed
Sa Body elongate; Ur larger than halfPr; A 1 almost projection; right B with inner expansion bordered
longer than Pr; longest terminal seta on left CR by hairs: Heterorhabdus robustus (fig . 7.291)
very short: Heterorhabtlus vipera (Giesbrecht, 4a R ig ht PS Exp2 w ith narrowly based inner poin-
1SS9) (Fig. 7 .294) ted projection; right B without inner expansion:
6 Mx2 LiS with unequal setae ...... .... ........... ....... .. S Heterorhabdus compactus (Fig. 7 .2S7)
6a Mx2 LiS w ith both setae of equal length .......... 7 S PS right Exp2 with large internal lamella ......... 6
7 ln lateral view, Gns ventral marg in with distinct Sa PS right Exp2 with sma ll internal proj ection .... 7 ;::-'!':
step posteriorly; PS terminal spine < 112 length of 6 Right PS Exp2 lamella with distal ridge situated Fee
Exp3 except in some large specimens (2.S mm): obliquely, so that adjacent rounded lobe occu-
Heterorhabdus lobatus Bradford, 197 1 (this pies most of termina l part of lamella:
species has not been recorded in the South Heterorhabdus papilliger (Fig . 7.290)
Atlantic; it has been included here because it is 6a Right PS Exp2 lamella with distal serrated ridge
easily confused H. papilliger) (Fig. 7 .2SS) situated centrally and parallel to axis oflamella so
7a 1n lateral view, G ns without conspicuous step in that rounded lobe situated on each side of ridge
posterove ntral margin; PS terminal spine > 112 terminally: Heterorhabdus lobatus (Fig. 7 .2SS) 8
the length of Exp3 except in some large speci- 7 Right PS Exp 1 with small inner distal knob; B
me ns ( 1.9 mm): Heterorhabdus papilliger with angu lar inner distal marg in distal to large
(Claus, 1S63) (fig. 7.290) inne r lobe: Heterorhabdus tanneri (Fig . 7.293)
S Ventral flap of Gns large, extending almost to 7a Right PS Exp I w ithout small inner knob; B w ith-
posterior border ........ ...................... .................. .9 out angular inner distal margin distal to large 'Rl
Sa Ventral flap ofGns small; in lateral view, ventral inner lobe ........... ....... ....................... ............ ... ... S 'Sie
posterior profile of Gns straight: Heterorltabtlus S Left PS B with small inner projection: Hetero- 0!:
abyssalis (Giesbrecht, 1SS9) (fig. 7.2S4) rhabdus clausii (Fig. 7.2S6) ·~
9 In dorsal view, Gns lateral borders smoothly Sa Left PS B without inner projection ..... ........ ...... 9 ~
rounded; in lateral view , left side of Gns projects 9 Right PS B inner lobe directed anteriorly so that
into rounded lobe, posterior to point where geni- w he n mounted it is twisted at its base:
tal flap hinges : Heterorhabdus norvegicus Heterorhabdus abyssalis (Fig . 7.2S4) '!..::
(Boeck, 1S72) (Fig. 7.2S9) 9a Right PS B inner lobe directed more or less into
9a In dorsal view, one or both Gns lateral borders midline, spine patch on inner edge broad distally l=:rJ
undulating ....... .. ... ...... ..... .... ..................... ........ 10 ............. ........ ..... ......... .... ...... ..................... ....... 10
Copepoda 945
emng lC Right P5 B inner lobe arises from proximal part with 4 setae the outermost 3 of which are reduced. P 1-
IS ;.an- of segment by narrow stalk: Heterorhabdus 4 distinctly 3-segmented, Enp segments ofP I and 5 2-
austrinus (Fig. 7 .285) 3-segmented, inner distal setae on B of P 1 inserted on
-: lD lOa Right P5 B inner lobe arises from central part of conical projection. Spine and seta formula:
~ ~ segment by wide stalk: Heterorhabdus norvegi-
~ cus (Fig. 7.289) coxa basis cxopod segments endopod segments
Leg I 0-0 0-1 1- J ;1- J; 11,1,4 0-1 ;0-2; 1,2,2

Key to the species of Heterostylites Leg 2 0- 1 0-0 1-1;1-1;11 1,1,5 0- I ;0-2; 1,3,2
Females Leg 3 0-J 0-0 1-1;1-1 ;J JI,l,5 0-1 ;0-2; I ,3,2
fGDo- A1 extends beyond CR by 6 segments. Total Leg4 0-1 1-0 l- 1;1-l;lll,l,5 0-1 ;0-2; I ,3,2
length 4.0-5.4 mm: Heterostylites major (F . Leg 5 0-0 1-0 1-0;I-l;Il,l,3 0-1;0-1;1,3, 1
..,
Dahl, 1894) (Fig. 7.296)
0:! ••.. 3 1:! A I extends beyond CR by 5 segments. Total P5 natatory, symmetrical, very similar to other swim-
ifi:""orm length 2.3-3.4 mm: Heterostylites longicornis ming legs; inner edge seta on Exp2 modified into long
- ·-···5 (Giesbrecht, 1889) (Fig. 7 .295) slender spine at1iculated with its segment.
i~on
I Fig. ~!ales Males
Left P5 Exp3 with 2 unequal term inal spines, Total length 1.2-8.6 mm. Body similar to female, but
.m_A longest about 3 times length of shot1er, and 1 Ur of 5 somites. Left Al geniculate, 21-segmented.
I!Oiffied shorter subterminal spine: Heterostylites longi- Mouthparts and P 1-4 similar to female. P5 wi th 3-seg-
~ cornis (Fig. 7.295) mented Enp and Exp on left, B usually modified with
__) I!! Left P5 Exp3 with 2 very unequal terminal spi- one or more inner extensions which may bear spines;
r ;x»innes, longest 14 times longer than shottest, and 1 Exp and Enp usuall y 2-segmented on right, last Exp
1::5!00.: longer subtenninal spine: Heterostylites major segment recurved; C on either side may sometimes be
(Fig. 7.296) modified with extensions.
_ _6
oo._.. / ~.!Ill.ily Lucicutiidae Sars, 1902 There is only one genus (Lucicutia Giesbrech t, 1898) in
• ......7ed Females this family which was reviewed by Hiilsemann ( 1966).
oa:u- :ooi length 1.3-9.8 mm. Body slende r, Pr oval, Ce
~lla: :!:3d Pd 1 separated, Pd4 and 5 fused, posterolateral Key to Lucicutia species (adapted from Hiilsemann,
~ers rounded. Ce rounded anteriorly, sometimes 1966)
2r=~e .:h antero lateral horns; rostral filame nts slender. Ur Females
~50 '!"...ongate, of 4 somites. CR symmetrical, relatively 1 P l Enp 3-segmented .......................................... 2
-::~e ~g. with 7 setae. AI symmetrical, 25-segmented. A2 1a PI Enp 2-segmented .................................. .. .... 13
L::-: 3 .... ;th 1-2 outer edge setae; Exp and Enp separated 2 Ce with 2 pairs of spine-like protrusions .......... 3
__.._: B
aum B: Enp 2-segmented, Enp1 with 2 setae a t about 2a Ce with only I pair or without protrusions ...... .4
11 t!r"ge c!dlength, Enp2 with 7 inner setae medially and 6 or 3 Spine-like protrusions on anterior edge of Ce
- setae terminally; Exp 8-segmented. Md gnathobase strong, directed forward; sides of anal somite
~---=n tri:h 9 sharp teeth; Enp 2-segmented, Enp I with 4 straight; CR longer than Ur: Lucicutia bicomu-
t~e se'..:>.e. Enp2 with 8 setae termina lly; Exp 1-4 with 1 ta Wolfenden, 1905 (Fig. 7.300)
!K..a each, Exp5 carrying 2 setae. Mx 1 Li1 with 13 spi- 3a Spine-like protrusions on anterior edge of Ce
1400- ~ and setae; Li2 with 3 setae; Li3 with 3 setae, Li4 smaller, directed downwa rd; ana l somite swol-
.......z....h ~ setae; Enp !-segmented, with 4+ 5 setae; Exp len; CR shorter than Ur: Lucicutia maxima
t --9 -=-~e. with 11 setae; Le 1 with 5-6 setae. Mx2 Lil and Steuer, 1904 (Fig . 7.312)
.'5.:1~ .: ~;th 3-4, 3 setae; L i3 and 4 with 3 setae each; Li5 4 Anal somite long, almost as long as CR ....... .... 5
Ca_'-e: 'aT.h 3 setae one of them stouter than other 2, spinulo- 4a Anal somite much shorter than CR ................... 6
~- Enp with 7 setae. Mxp C with 3-5 setae; B with 5 Anal somite slender: Lucicutia Iucida Farran,
liS c::o p!:ehes of setules or spinules and 3 setae medially; I 908 (Fig. 7 .308)
- --'\.
E.;:p 6-segmented, Enp I almost fully incorporated into Sa Anal somite swollen: Lucicutia grandis
·- I O 3 vith 2 setae, Enp2-5 with 2 setae each, segment 6 (Giesbrecht, 1895) (Fig. 7 .306)
946 Copepoda
6 P5 Enp 2-segmented: Lucicutia ova/is Males (unknown fo r L. major)

(Giesbrecht, 1889) (Fig. 7 .313) I P l Enp 3-segrnented ...................................... .... 2
6a P5 Enp 3-segmented ......................................... .7 1a PI Enp 2-segmented ................................... ..... l 2
7 A I reaches 3-4 segments beyond CR; sma ll 2 Ce with 2 pairs of spine-like protmsions ... ....... 3
form , total length 1.5-2.0 mm: Lucicutia longi- 2a Ce with only I pair of spine-like protrusions or
comis (Giesbrecht, 1889) (Fig. 7 .307) without protmsions .............. ..... ........................ .4
7a Characters not in this combination ...... .............. 8 3 Spine-like protrusions strong, lateral pair poin-
8 Each side of Ce with 1 strong spine-like point: ted backwards: Lucicutia bicornuta (Fig. 7.300)
Lucicutia aurita Cleve, 1904 (Fig. 7 .299) 3a Spine-like protrusions small, lateral pair pointed
8a Each side of Ce with I rounded protrusion or downwards; anal somite slightly shmter than
without protrusion .............. .. ... .. .......... .............. 9 preceeding somite; CR about 7 times longer than
9 Each side of Ce with 1 rounded protrusion ..... I 0 wide: Lucicutia maxima (Fig. 7.3 12)
9a Each side of Ce without protrusions ... .... ........ ll 4 Each side of Ce with l strong spine-like point
10 Anal somite shorter than preceeding somite; CR .. .................... ......... ...... .... .. ......... .... .... ............... 5
longer than rest of Ur: Lucicutia major 4a Each side of Ce with 1 obtusely pointed or roun-
Wolfenden, 1911 (Fig. 7.311) Wolfenden's, ded protrusion or without protrusion ................. 6
( 1911) figure of L. grandis appears to fit his 5 P5 B on left and right protruded, bearing teeth:
description of L. major so is used here Lucicutia aurita (Fig. 7.299)
lOa A nal somite as long as preceeding somite; CR Sa PS B on left protruded, bearing teeth; rig ht B
shorter than rest of Ur: Lucicutia wolfendeni protrudes proximally, w ithout teeth: Lucicutia
Sewell, 1932 (Fig. 7.3 14) grandis (Fig. 7.306)
ll P5 terminal spine about 1/2 length ofExp3; total 6 Each side ofCe with 1 obtusely pointed or roun-
length less than 2 mm ..................................... .1 2 ded protrusion: Lucicutia wolfendeni (Fig.
11 a P5 terminal spine about length of Exp3; CR 3-4 7.314)
times longer than wide; total length 1.9-2.6 mm: 6a Sides of Ce without protmsions ........... ...... ....... 7
Lucicutia curta Farran, 1905 (Fig. 7.302) 7 Right A I extends 3-4 segments beyond CR; CR
12 CR separate, divergent; anal somite shorter than abo ut 6 times longer than wide: Lucicutia longi-
preceding somite; genital swelling large: comis (Fig. 7.307)
Lucicutia jlm•icornis (Claus, 1963) (Fig. 7 .303) 7a Right A I reaches at most 2 segments beyond
12a CR parallel and touching; anal somite as long as CR; CR at most 7 times longer than wide ....... .8
preceding somite; genital swelling small: 8 P5 right C inner margin with spiny process:
Lucicutia gemina Farran, 1926 (Fig. 7 .305) Lucicutia Iucida (Fig. 7.308)
13 A 1 extends 4-7 segments beyond CR ............ . 14 8a P5 right C inner margin smooth ......... ... ...... ... ...9
13a A I extends at most 2 segments beyond CR. ..... l5 9 P5 right and left B with prox imal portion of its
14 A I extends 6-7 segments beyond CR: Lucicutia inner margin extended into pointed projection;
macrocera Sars, 1920 (Fig. 7.309) right Enp 3-segmented: Lucicutia ova/is (Fig.
l4a Al extends 4-5 segments beyond CR; CR shor- 7.313)
ter than rest of Ur, 5 times longer than w ide; P5 9a P5 right and left B w ithout proximal pointed
terminal spine as long as or slightly shorter than projections; right Enp 2-segrnented ................. 10
l/2 Exp3; tota l length about 3.5 mm: Lucicutia l 0 CR 3-4 times longer than wide: Lucicutia curta
magna Wolfenden, 1903 (Fig. 7.3 10) (Fig. 7 .302)
15 A 1 extends as far as posterior border of CR; ana I 1Oa CR more than 4 times longer than wide; total
somite longer than preceding somite; in lateral length less than 2 mm .. ... ................................. ll
view Gns w ith large ventroposterior projection; I I Al extends beyond middle of CR; anal somite
Ce with 1 strong spine-like proj ection or roun- shorter than preceding somite; P5 left B w ith
ded protmsions on each side: Lucicutia clausii inner distal com er protruded with 4 (-8) small
(Giesbrecht, 1889) (Fig. 7.30 1) teeth: Lucicutia jlavicornis (Fig. 7.303)
15a A I extends 2 segments beyond CR, anal somite II a A l extends to middle of CR; anal somite as long
about as long as preceding somite; G ns wi thout as preceding somite; P5 left B with inner distal
ventroposterior protrusion; Ce evenly rounded: comer protruded, ending in I point: Lucicutia
Lucicutia gaussae Grice, 1963 (Fig. 7.304) gemina (Fig. 7.305)
Copep oda 947
12 P5 left B inner proximal corner with I sp ine-like nes small. P l B inner seta situated on its anterior sur-
~
protrusion, inner distal corner of same segment face and passing across face of Enp segment l . Enp
__ 12 with small teeth: Lucicutia macrocera (Fig. segment l of P2 typically incised and ornamented
-··--3 7.309) wi th 1 or more hook-like spinous processes; Enp typi-
ions or 12a P5 left B anned d ifferently ....... ....................... 13 cally small, cylindrical. Spine and seta formula typi-
-:-·-·-r • 13 P5 left B middle of i1mer margin with 2 pointed cally as follows but sometimes reduced.
r porn- projections, right B with I spine-like proj ection:
Lucicutia magna (Fig. 7.3 I 0) basipod 1 bas ipod 2 exopod segments endopod segments
~ted l 3a P5 left B armed differently ............... .... ... ........ l4

Leg I 0-1 1-1 I-1; T-1; 11,1,4 0-1; 0-2; I ,2,2
:r dlan 1-t In dorsal view Ce smoothly rounded; right AI Leg 2 0-1 0-0 I-1; I- I ; 111,1,5 0-0; 0-2; 2,2,4
~jmn extending beyond CR; P5 right B inner margin Leg 3 0-1 0-0 I- I; 1-1; III,1.5 0-1; 0-2; 2,2,4
smooth, left B inner distal corner ending in I Leg4 0-1 1-0 1-1; I-1; TTI,l,5 0-1; 0-2; 2,2,3
~ pomr point: Lucicutia gaussae (Fig. 7.304)
1-la Each side of Ce with 1 strong spine-like pro- P5 u niramous, very small, symmetrical 2-4-segmen-
£ iOilll- trusion or shallow protrusion; right A1 extending ted, C and intercoxal sclerite f used; B and Exp seg-
-· ___ 6 to end ofCR; P5 right B inner margin with small ment I usually with outer seta and spine respectively;
; teelh: proximal tooth-like projecti on, left B inner distal Exp segment 2 with 2-4 setae, terminal Exp segments
corner variable: Lucicutia clausii (Fig. 7.30 I) may be fused.
c.;r:: B
riaaia fami ly Metridin idae Sars, 1902 Males
Reference: Boxshall (in prep.) Ur of 5 somites, sometimes strongly asymmetrical,
r-roun- Females single genital aperture located ventrolaterally at pos-
fig. 3ody elongate. Ce and Pdl separate, Pd4 and 5 fused, terior margin of Gns, usually on right side, sometimes
posterolateral corners of Pr usually rounded, expan- on left. A I usuall y prehensile on left side only, occa-
ded in Gaussia; Pdl with dark-pigmented spot on one sionally on right side only. Mouthparts similar to
:P'"" CR S!de in Pleuromamma. Ur of 3 somites, CR often those of female. P5 asymmetrical, attached to plate
~- ::..:,-ymmetrical with up to 6 setae; genital apparatus fonned from fusion of C and intercoxal sclerite; right
.:ompris ing a common genital aperture located leg comprising B with an outer seta and ?2-3-segmen-
beyond medially on ventral surface of Gns; copulatory pore ted Exp, Exp segment 2 with an inner spinous process
rontained with in median genital ape1ture. R with pai- in some genera, Exp segment 3 with 1 or 2 minute dis-
-as: :-ed fila ments. A1 23-24-segmented; segments I and 2 tal setae; left leg comprising B with outer seta and 2-
:::sed. segments 3 to 7 separate, segments 8 and 9 or 3-segment Exp: Exp segment l bearing curved
.-------9 :':!sed. segments 10-22 separate, apical segment dou- inner process, distal segment swollen, often curved or
1:1of its :;re. A.2 with C and B separate with 1 and 2 setae claw-like.
~ non: r-e:,-pectively; Enp 2-segmented, bilobed di stal seg-
[s (Fig. .:uem with 1-2, 8-9+6-7 setae; Exp 7-segmented with This family contains three genera: Gaussia Wolfenden,
I. ~- L 1, 1, 1, 4 setae respectively. Md with well- 1905; Metridia Boeck, 1864; and Pleuromamma
-ed ~,-eJoped gnathobase, palp B with 4 setae; Enp 2- Giesbrecht, 1898, all of which occur in the South
_10 ~'"gl11e nted with 4 and l 0 setae respectively; Exp 5- Atlantic.
crzna ~'"gl11ented with 1, I, I, 1, 2 setae respectively. Mxl
~:: '' ith about 15 sp ines and setae, Li2-4 with 4-5, 4 , Key to genera and species of Metridinidae
e: Uk:l1 5-- setae; Enp 2-segmented with 6, 9-12 setae; Exp Females and males
__ il l Pd l with anterolateral dark-pigmented spot on
·ith 11 setae; Le I and 2 with 9 and 1 setae respecti-
.somire \"el:. Mx2 Lil-5 with 5 (9 in P. xiphias), 3, 3, 3, 4 one side: Pleuromamma (p. 948)
B wiih ~'"me respectively; Enp 4-segmented with 3-4 , 3, 2, 2 1a Pd I without anterolateral dark-pigmented spot
:imJll ::>e..:!e. or reduced. Mxp 7-segmented, C with I, 2, 4, 4 ................. ............... ......... ............... ................. ..2
s.e-..:!e: B with 3 setae plus 2 setae on the incorporated 2 Posterior corners of Pr produced into long
'="::rp segment 1; free Enp 5-segmented with 4, 4, 3, 3, points; Ur of female symmetrically or asymme-
- 5et:ae respectively . P 1-4 with both rami 3-segmen- trically swollen; anal somite with very long pos-
riauitr ;..,_~ Exp of P2-4 very large and flat, outer margin spi- terolateral processes: Gaussia (p. 948)
948 Copepoda
2a Posterior comers of Pr not produced into long 7 P5 segment 4 1/3 length of segment 3; segment
points; Ur of female symmetrical, not swollen; 3 often without outer edge spine: Metridia cur-
anal somite with short posterolateral processes: ticauda Giesbrecht, 1889 (Fig. 7.319)
Metridia (p. 948) 7a P5 segment 4 more than 3/4 length of segment 3;
segment 3 with outer edge spine: Metridia longa
Key to species of Gaussia (Lubbock, 1854) (Fig. 7.322)
Females and males
1 Forehead flat anteriorly, in lateral view without Males (unknown for M. discreta) (right- or left-han-
anterodorsal triangular projection; Pr posterior dedness of P5 variable)
comers asymmetrical, left comer shorter and Terminal segment of right PS with long appen-
curves outwards whereas right corner is longer dix, originating proximally and lying parallel to
and divergent. Male unknown: Gaussia a:.ym- outer border: Metridia princeps (Fig. 7.324)
metrica Bj0rnberg and Campaner, 1988 (Fig. Ia Terminal segment of right PS without long
7 .315) appendix ... ......................................................... .2
la Forehead pointed anteriorly, in lateral view with 2 Right Expl of P5 without long, pointed inner
anterodorsal triangular process; Pr posterior cor- extension: Metridia venusta (Fig. 7 .325)
ners more or less symmetrical, divergent. Male 2a Right Expl ofP5 with long, pointed inner exten-
P5 left segment 3 with blunt and short, proxi- sion ...... ..... .................. ... .................... ................. 3
mally directed spine; terminal segment of right 3 Left P5 narrowed distally; left P5 rounded dis-
P5 with prominent undulating border, and 4 tally: Metridialonga (Fig. 7 .322)
setae; distance between setae 1 and 2 and 2 and 3a Both P5 rounded distally ... ..... ...... .... ............... ..4
3 almost equal: Gaussia princeps (T. Scott, 4 Posterior comers of Pr rounded ............ ... .. .. ...... 5
1894) (Fig. 7.3 16) 4a Posterior comers of Pr pointed or with small
knob ... ............. ......... .......... ........ ........... ..... .... .... ?
Key to species of Metridia (unknown for M. calypsoi) 5 Right P5 with 1 thick, short inner spine on penul-
Females timate segment: Metridia brevicauda (Fig. 7.317)
1 P5 2-segmented, asymmetrical: Metridia venu- Sa Left P5 with more than 1 short spine on penulti-
sta Giesbrecht, 1889 (Fig. 7.325) mate segment ..................................................... 6
l a P5 3- or 4-segmented ........................ .... ........... ..2 6 Left P5 with 3 inner spines on penultimate seg-
2 P5 3-segmented ........ ................... ... .. .... .... .... ... ..3 ment: Metridia gerlachei (Fig. 7.321)
2a P5 4-segmented ....................... ........ ......... ....... ..5 6a Left P5 with 2 inner spines (one thick and one
3 P5 segment 3 very small with only 2 terminal fine) on penultimate segment: Metridia calypsoi
setae; posterior Pr rounded: Metridia brevicau- Gaudy, 1963 (Fig. 7 .318)
da Giesbrecht, 1889 (Fig. 7.317) 7 Posterior comers of Pr bearing minor knob:
3a P5 segment 3 larger and with 4 setae; posterior Metridia curticauda (Fig. 7.319)
Pr rounded or pointed ... .... ... ............ ..... .......... .. .4 7a Posterior com ers of Pr pointed: Metridia lucens
4 Posterior Pr comers pointed; CR twice as long as (Fig. 7 .323)
wide: Metridia lucens Boeck, 1864 (Fig. 7.323)
4a Posterior Pr comers rounded; CR about 3.5 Key to the species of Pleuromamma (adapted from
times as long as wide: Metridia gerlachei Steuer, 1932)
Giesbrecht, 1902 (Fig. 7.32 1) Females
5 P5 C with row of long hairs; anterior Ce without 1 Anterior Ce prolonged into point: Pleuromamma
anterior horns; CR about 5 times as long as wide: xiphias (Giesbrecht, 1889) (Fig. 7.332)
Metridia princeps Giesbrecht, 1889 (Fig. 7.324) la Anterior Ce not pointed ..................................... 2
Sa P5 C naked ......................................................... 6 2 Proximal segments of A 1 with 2-4 large spines
6 P5 B with outer distal plumose seta; CR more ............................. .... .............. .... ... ........... .......... 3
than twice as long as wide .............. ... .......... .. .... 7 2a Proximal segments of Al w ith small points ... . .4
6a P5 B with outer distal seta naked; CR less than 3 Proximal segments of A I with 2 spines, one
twice as long as w ide: Metridia discreta Farran, straight, other curved: Pleuromamma abdomi-
1946 (Fig. 7.320) na/is (Lubbock, 1856) (Fig. 7.326)
Copepoda 949
~em 3a Proximal segment of Al with 3-4 large spines: Family Phyllopodidae Brodsky, 1950
cur- Pleuromamma quatlrungulata (F. Dahl, 1893) Reference: Boxshall (in prep.)
(Fig. 7.330) Females
Dei:.i 3: 4 P5 terminated by setae: Pleuromamma robusta Total length 2-3 mm. Body relatively robust, Ce and
aa (F. Dahl, 1893) (Fig. 7.33 1) Pd1 separated, Pd4 and 5 fused with posterolateral
4a P5 termined by 3 spines ............................ ........ 5 corners of different shapes, sometimes asymmetrical.
5 P5 2-segmented, apical spines long and pointed, Ce rounded anteriorly, R short and bifurcate; labrum
c-furri- equal to about half their segm ent: Pleuromamma large and greatly produced. Ur relatively long, of 4
borealis (F. Dahl, 1893) (Fig. 7.327) somites; Gns usu ally asymmetrical with single genital
llppen- Sa P5 !-segmented, apical spines vety short ... ...... 6 aperture located ventrolaterally on posterior right bor-
a.:...:o!I U> 6 Anal so mite with lateral borders divergent, der. CR symmetrical, relatively short, with up to 6
somite dilated posteriorly; Gns symmetrical: setae. A l symmetrical, not extending beyond the pos-
t long Pleuromamma gracilis Claus, 1863 (Fig. 7 .328) terior border of the Pr, 24-segmented. A2 C with 1
, 6a Anal somite with lateral borders parallel, somite seta; B with 2 outer edge setae; Exp and Enp separa-
cmer not dilated; Gns flattened on left, genital pore ted from B; Enp 2-segmented, Enp I with 1 seta distal
with l large black spot: Pleuromamma piseki to midlength, Enp2 with 7-8 inner setae and 6 setae
~- Farran, 1929 (Fig. 7.329) terminally; Exp 8-segmented, proxima l 2 segments
without setae and paitially fused with Exp3. Md gna-
~- .VIales (right- or left-handedness of P5 variable) thobase longer than pa1p, w ith numerous sharp teeth
1 Anterior Ce prolonged into point: Pleuromamma and one inner seta; Enp 2-segmented with 2 and 6
... ·-- xiphias (Fig. 7.332) setae respectively; Exp 5-segmented, segments 1-4
with I seta each, segment 5 carrying 2 setae. Mx I Li 1
s.::=.1:
' la Anterior Ce not pointed ..................................... 2
2 Outer edge of Expl on long narrow P5 (maybe large, remainder of limb reduced; Li 1 with 15 spines
right or left), with only I spine ........ ................. 3 and setae; Li2 with 2 seta; Li3 and 4 with I seta each;
~ 2a Outer edge ofExpl on long narrow P5 (left) also Enp !-segmented, with 4 setae; Exp small with 7
~.3rt) with chitinous peg ........... .................................. 5 setae; Lel in the form of a small lobe, without setae;
::i- 3 Ur more or less asymmetrical; segment 17 of A I Le2 absent. Mx2 well developed; Li I and 2 with 3 and
5 with distal pointed extension: Pleuromamma 2 setae respectively; Li3 and 4 with 3 setae each; LiS
~ seg- llbdominalis (Fig. 7.326) with 4 setae; terminal part 4-segmented with 10-1 1
3a Ur not very asymmetrical; segment 18 of A 1 setae. Mxp C short with 2 proximal, 2 medial and 1
i! roe without distal pointed extension ........... .... ........ .4 distal setae; B w ith 3 setae dista lly; Enp 5-6-segmen-
~ Proximal segments of A I with 4 teeth: ted, Enp I almost fu lly incorporated into B, with 2, 4 ,
Pleuromamma quadrungulata (Fig. 7 .330) 4, 3, 1+2 setae respectively, each segment with one of
,,: -la Proximal segments of A1 with degenerate teeth: setae small or vestigial. Spine and seta formula:
Pleuromamma robusta (Fig. 7.33 1)
la!I!IS 5 Chitinous peg on left P5 Exp l w ithout distal coxa basis exopod segments endopod segments
knob: Pleuromamma borealis (Fig. 7.327) ---
Leg l 0- 1 1-1 1-1 ;I-I ; 11,1,4 0-1 ;0-2; 1,2,2
5a Chitinous peg on left P5 Exp I with distal knob Leg 2 0-1 0-0 1-1;1-1 ;TTT,l,5 0- 1;0-2;2,2,4
~ .... .. ..... ............................... ....................... .......... 6 Leg 3 0-1 0-0 1-1 ;1-1 ;Tll ,1,5 0-1 ;0-2;2,2,4
6 Left A I w ith segrhent 17 naked, double tooth Leg4 0- 1 0-0 1-1 ;I- I ;111,1,5 0-1 ;0-2;2,2,3
row on segment 18 as we ll as on proximal part
of segment 19-2 1; left P5 Exp I process with P5 uniramous, not natatoty, C and intercoxal sclerite
knob separated from process by slight notch: fused; B w ith an outer edge seta; Exp 3-segmented with
Pleuromamma piseki (Fig. 7.329) I-0; 0-1 ; 0,0,1 spines and setae; outer and distal borders
~ 6a Left A I with toothed ridges on segments 17, 18 of Exp segments 2 and 3 expanded and toothed.
and 19-2 1; serrations a re very variabl e, teeth can
be pointed or mo re or less blunt, long and thin or Males
short and thick; left P5 Exp I process ve1y varia- Body similar to female, but Ur of 5 somites. Left AI
ble: Pleuromamma gracilis (Fig. 7 .328) geni culate, 20-segmented. Mouthparts and P 1-4 simi-
lar to female. P5 C and intercoxal sclerite fused, B
950 Copepoda
with long outer edge seta on each side; left leg with !- asymmetrical spiniform posterior corners. Ur usually
segmented, foliaceous Enp, Exp 3-segmented with of 3 somites, Gns often with asymmetrical spine
last 2 segments in form of chela; right leg without armature; Ur2 may also bear processes. AI 24-25-
Enp, Exp 3-segmented, segments 2 and 3 more or less segmented. A2 Exp and Enp almost equal in size; C
fused, segment 3 expanded and truncate distally. with 1 seta, B with 2 setae; Enp 2-segmented carrying
2 and 13-15 setae respectively; Exp 7-segmented car-
Brodsky (1950) separated Phyllopus Brady, 1883 rying I, 3, I, 1, 1, I, and 4 setae respectively, although
from the the remaining Arietellidae by placing it in a further fusion of proximal segments may take place in
separate subfamily; later Campaner (1977) removed it some genera so that Exp is 5-segmented. Md gnatho-
from the Arietellidae and placed it in a family of its base with well developed teeth, B with 4 setae, Enp 2-
own. Only one genus is placed in this family. segmented with 3-4, and 6-8 setae respectively; Exp
4-segmented with 1, 1, 1, and 2-3 setae respectively.
Key to species of Phyllopus Mxl variable, Lil with 14-15 spines and setae, Li2
Females with 1-4 setae, Li3 with 2-4 setae, Li4 with 2-5 setae,
1 Posterior borders ofPr very asymmetrical... .. ... 2 Enp with 2-4 + 4-13 setae, Exp with 7-11 setae, Le2
1a Posterior borders of Pr symmetrical or only with 1 small seta, Lei with 8-9 setae, 2 of them short.
slightly asymmetrical... ....................... ... ............ 3 Mx2 well-developed, Lil-5 with 4-5, 3, 3, 3, and 3-4
2 In lateral view right prosomal corner pointed; setae respectively, terminal part with 6-8 setae. Mxp
Gns with lateral swellings in dorsal view; A I relatively small, C usually with 3 pronounced endites
extends almost to posterior border of left prose- bearing 2, 3, and 3-4 setae respectively; B with 5 dis-
mal corner: Phyllopus impar Farran, 1908 (Fig. tal setae and sometimes with proximal row of inner-
7.336) edge spinules, Enp 5-segmented with 2-4, 1-4, 1-3, 2-
2a In lateral view right prosomal corner more or 4 (1 usually on outer edge), and 3-4 setae respectively.
less obliquely truncate; Gns without lateral swel- P 1-5 Exp and Enp 3-segmented; apical spine of Exp
lings in dorsal view; A1 extends to posterior bor- strongly serrate; sometimes Enp segments 1 and 2
der of Pd2: Phyllopus bidentatus Brady, 1883 fused. Spine and seta fonnula:
(Fig. 7.334)
3 Gns asymmetrical, swollen on right: Phyllopus coxa basis cxopod segments endopod segments
helgae Farran, I908 (Fig. 7.335)
Leg I 0-1 0-1 TIO- I ; flO- l ; 0-1; 0-1/2; 0/ 1,2,2/3
3a Gns symmetrical: Phyllopus aequalis Sars, I 920 ll/O,T,3/4
(Fig. 7.333) Leg 2 0-1 0-0 I-I; l- 1; ll/Ill,l,5 0-1 ; 0-2; 0/2,2,3/4
Leg 3 0-l 0-0 T-l; T-1 ; TTT,T,5 0-1 ; 0-2; 0/2,2,3/4
Males Leg 4 0- l 0-0 I- I; I- I; lll,l,S 0- l ; 0-2; 0/2,2 ,2/3
1 P5 left Exp3 straight... .... .. ............ ..... ... ... .......... 2
1a P5 left Exp3 curved sharply at its base: Phyllopus P5 biramous, natatory, C and B without setae, Exp
bidentatus (Fig. 7.334) (Wilson, 1950, figures a segments 1 and 2 with or without 1 outer edge spine,
male P. aequalis, although this appears to be very segment 2 with strong inner spine-like process, Exp
similar to the male of P. bidentatus) segment 3 with or without 2 outer edge spines, 1 ter-
2 P5 left Enp very expanded distally; right Exp3 minal serrate spine and 2 or 4 inner edge setae; Enp
slightly expanded distally: Phyllopus helgae segments 1 and 2 with 0-1, 1 inner edge seta respecti-
(Fig. 7.335) vely, Enp segment 3 with 4-6 setae.
2a P5 left Enp slightly expanded distally; right
Exp3 rectangular in shape: Phyllopus impar Males
(Fig. 7.336) Mouthparts and P 1-4 identical to female apart from
male AI , and sometimes P2. Ur of 4-5 somites. AI
Family Centropagidae Giesbrecht, 1892 prehensile on right. P5 asymmetrical, usually resem-
References: Gurney ( 1931 ), Bayly and Arnott (1969). bling other swimming legs, rami 2 or 3-segmented,
Females right Exp variously transformed into grasping organ,
R always present, with 2 filaments. Ce and Pdl , Pd4 sometimes Enp atrophied.
and 5 separate, latter somite rounded or often with
Copepoda 951
b........,ll~ There are a number of genera in this famil y many of 9 Ur2 in dorsal view with convex right lateral bor-
~ ~....::le which live in freshwater. Centropages Kroyer, 1848, der and straight left border bearing small ventro-
~-::5- lives in marine habitats, Gippslandia Bayly and lateral spine just posterior to midlength; posteri-
~:C
Amott, 1969, li ves in estuarine environments in or prosomal corners and Gns very asymmetrical:
~wg
south-eastern Australia, Gladioferens Henry, 1919, Centropages brachiatus (Dana, 1849) (Fig .
~car- li ves in brackish, nearshore habitats in Austral ia and 7.337)
_gh ;\ew Zealand, Isias Boeck, 1864, lives in nearshore 9a Ur2 in dorsal view wi th left border without oma-
·-~in marine habitats and is possibly benthopelagic, mentation; posterior prosomal corners a nd Gns
~
Parathalassius Dussart, 1986, lives in brackish water not very asymmetrical in shape ....................... lO
~2-
near the mouth of the La Foa River, New Caledonia, 10 Ur2 in dorsal view with slight, wide, right side
and Sinocalanus Burckhardt, 1913 , and Limnocalanus anterolateral swelling; A I extends to posterior
~-= E.xp
m ·ely. Sars, 1863, live in northern hemisphere brackish a nd border of CR: Centropages typicus Kroyer,
~.Li2 freshwater habitats. The remaining genera are found 1849 (Fig. 7.347)
~-e.
m fres hwater habitats. Only Centropages has been lOa Ur2 in dorsal view w ith conspicuous, na rrow,
taken in the South Atlantic. right side process just posterior to midlength,
--...
D. T
-~
;:..-,.
plus more anteroventrally placed swelling; A1

i 5ho;t.
oa: :....! Key to the species of Centropages does not extend to posterior border of CR:
:. r hp Females Centropages chierchiae Giesbrecht, 1889 (Fig.
~...;a..... _..-..CS I Posterior comers of Pr rounded ......................... 2 7.340)
15&- Ia Posterior corners of Pr pointed .......................... 7 11 Posterior Pr points directed straight posteriorly;
c::::lC'r- CR with peg-like projection on posterior border: Gns with ventral spine; A I does not extend to
-·- .... Centropages bradyi Wheeler, 1899 (Fig. 7 .338) posterior border of CR: Centropages orsinii
m·eJ~· . 2a CR w ithout such projection ............................... 3 Giesbrecht, 1889 (Fig. 7.346)
fE._~
3 Ur2 ornamented ................................................ .4 lla Posterior Pr points divergent; Gns with spinous,
2::A:!. ...:.
. 'l 3a U r2 unomamented ............................................. 5 asymmetrical, anterolateral swelling on each
.! Ur2 with spi nous lateral knob on each side; Gns side and patch of dorsal spinules; AI extends to
with naked lateral borders: Centropages gracilis posterior border of CR: Centropages kroyeri
-=s (Dana, 1849) (Fig. 7.343) Giesbrecht, 1892 (Fig. 7.344)
.!a Ur2 with conspicuous ventral process, centrally
placed; Gns with a patch of spinules on right side: Males
Centropages violaceus (Claus, 1863) (Fig. 7.348) 1 Posterior corners of Pr rounded ......................... 2
P5 Exp 2 inner spine does not extend as far as Ia Posterior corners of Pr pointed .......................... 6
distal border of Exp3: Centropages longicomis 2 Right P5 Exp3 terminated by 2 thick spine-like
Mori, 1932 (Fig. 7.345) ex tensions: Centropages elongatus (Fig. 7.341)
~- :::xp 5a P5 Exp2 inner spine extends to or well beyond 2a Right P5 Exp3 tapering to single point.. ........... 3
1~.
di stal border of Exp3 ......................................... 6 3 Right P5 Exp3 sharply curved at an acute angle
LUp 0 P5 Exp2 inner spine extends as far as or slightly about 1/3 of its length from proximal end:
. I ~-
beyond distal border of Exp3: Centropages Centropages calaninus (Fig. 7.339)
~f::;p
elongatus Giesbrecht, 1896 (Fig. 7.341) 3a Right PS Exp3 not acutely bent... ..................... .4
~<!eLi-
Q.! P5 Exp2 inner spine extends well beyond distal 4 Right P5 Exp3 very broadly thickened proximal-
border of Exp3: Centropages calaninus (Dana, ly, with pointed projection on inner border:
1849) (Fig. 7.339) Centropages gracilis (Fig. 7.343)
Segments l , 2, and 5 of AI w ith conspicuous 4a Right P5 Exp3 only slightly thickened proximal-
I ~
anterior spine ......................... ............................ 8 ly without pointed projection on inner border
-z Proximal AI segments without large spines ... 11 ............... ...... ...................... ............... ................. 5
s . .-\1
~-
Gns without setae; posterior corners ofPr with 2 5 Right P5 Exp2 inner process arising narrowly at
i!::::..'"\!.. points on each side: Centropages furcatus about midlength from its bulbous segment; Exp3
t:.: . (Dana, 1849) (Fig. 7.342) curved in an S-shape and is at least 3 times longer
Gns w ith setae; posterior comers of Pr with I than Exp2 inner process; CR without distal peg-
point on each side .... .......................................... 9 li ke process: Centropages violaceus (Fig. 7.348)
952 Copepoda
Sa Right PS Exp2 inner process, which is truncate asymmetrical. A 1 symmetrical, usually 22-segmented
distally, arises broadly fro m about midlength; (24-segmented in Archidiaptomus, 2 1-segmented in the
Exp3 curved in an S-shape and is just greater Hyalinus group of Pseudodiaptomus). A2 Exp slightly
than length of Exp2 inner process; CR w ith dis- longer than Enp, Enp with fusion between segments
tal peg-like processes: Centropages bradyi (Fig. and with B. Md gnathobase with numerous small teeth.
7.338) Mxl Lil usually with with 15 spines and setae, Li2 and
6 Posterior comers of Pr w ith pair of points on 3 with 3 setae each, Li4 w ith 5 setae, Enp segments 1-
each side: Centropagesfurcatus (Fig. 7.342) 3 with 4, 4, 5 setae respectively, Exp with I 0 setae, Lcl
6a Posteri or corners of Pr with single point on each with I 0 setae. Mx2 Li 1-5 w ith 4, 3, 3, 3, 3 setae respec-
side ..................................................................... ? tively. Mxp C squat with 2, 3, 4 setae, one of terminal
7 Posterior comers of Pr with degenerate points; group enlarged into spine; B bordered by spinu les and
left PS Exp2+3 pointed: Centropages orsinii 3 setae; Enp 4-segmented, often carrying 4 characteris-
(Fig. 7.346) tic setifonn bristles (2 on segment 1 and 1 each on seg-
7a Posterior corners of Pr with distinct points; left ments 2 and 3). Pl-4 biramous, with 3-segmented rami;
PS Exp2+ 3 oblong ............... .... .......................... 8 C and B often decorated with spinules on anterior, pos-
8 Right geniculate AI segments 15 and 16 both terior and lateral surfaces, also sometimes on anterior
with large spine .... .. ..................... ....... .... ...........9 surfaces of Enp and Exp. Spine and setae formu la:
8a Right geniculate A I segment 16 with large
spine, segment 15 without spine: Centropages coxa basis cxopod segments endopod segments
typicus (Fig. 7.347)
Leg 1 0- 1 0-0 1-l ;0-1 ;Ofll,1 ,3 0- 1;0- l ; l ,2,2/3
9 Right PS Exp2 inner process gently curved at Leg 2 0- 1 0-0 l-1 ;1-l ;ll,l,S 0- l ;0-2;2,2,4
less then right angle: Centropages kroyeri (Fig. Leg 3 0- 1 0-0 1-1;T- l;H,l,5 0-1 ;0-2;2,2,4
7.344) Leg4 0- 1 0-0 1-1;1-1 ;TI,l,5 0- 1;0-2; 1/2,2,4
9a Right PS Exp2 inner process abruptly curved at
right angles ...................................... ............... . 10 PS uniramous or biramous; not natatory; with 3 Exp
10 Right PS Exp2 inner process abruptly curved at segments (the terminal 2 may be fused); may be
distal 1/3, w ithout teeth along distal part; spines slightly asymmetrical. Ovisacs present.
on A 1 segments 15 and 16 with their bases about
1/3 length of segments: Centropages brachiatus Males
(Fig. 7.337) Body of similar fo rm to female, U r of 5 somites. A l
1Oa Right PS Exp2 inner process abruptly curved at asymmetrical; left 22- or 24-segmented; right with 20-
about midlength and bearing teeth along distal 21 segments, geniculate between segments 18 and I 9;
part; spines on A l segments 15 and 16 with their terminal part beyond joint with 2-4 segments.
bases about l /2 length of segments: Centropages Mouthpa11s similar to female. PS uniramous or bira-
chierchiae (Fig. 7.340) mous, very asymmetrical; Exp 2- or 3-segmented; spi-
nes, Exp segments, and Enp, if present, variously
Family Pseudodiaptomidae Sars, 1902 modified into a form which appears to be adapted for
References: Madhupratap and Haridas (1978), Wa lter clasping.
(1986).
Females Sars ( 1902) proposed the family Pseudodiaptom idae
Small copepods slightly over 1 mm. Usually w ith to include Pseudodiaptomus H errick, 1884, and
characteristic eye composed of pigmented spot sur- Poppella Richard, 1888 (junior synonym of
rounded by refractile lens. Ce and Pd 1 fused or sepa- Calanipeda) , but did not define it. The above defin-
rate, Pd4 and 5 fused or separate; posterior metasom al tion has been created from a defi niti on of
corners may be rounded or variously decorated with Pseudodiaptomus (Madhupratap and Haridas, 1978;
large posterior spines or rows of spinules. Ce with Walter, I 986). Th is family conta ins the genera:
pair of rostral filaments. Ur of 2-4 somites, may be Archidiaptomus Madhupratap and Haridas, 1978;
asymmetrical; Gns very variable (symmetrical, asym- Calanipeda Kritchagin, 1873; and Pseudodiaptomus
metrical, ornamented with spines and/or spinules, or Herrick, 1884, of w hich only Pseudodiaptomus has
undecorated), CR tend to be elongate and may be been taken in the South Atlantic.
Copepoda 953
;-.emed ~ to the species of Pseudodiaptomus 4 P5 left Enp worm-like and elongate, extending
:C~tbe ~rmales beyond proximal thickening of Exp2+3:
~y Posterior comers of the Pr without spines .. ...... 2 Pseudodiaptomus serricaudatus (Fig. 7.355)
~G Posterior comers of the Pr w ith spines ....... ..... .4 4a P5 left Enp relatively short, extending as far as
;:ee-JL L:"r of3 somites, separation between somites 2 and proxima l thickening on Exp2+3: Pseudo-
~and 3 indistinct; Gns with ventral hyaline keel which tliaptomus uudu.s (Fig. 7.353)
~1 - ~nds beyond distal margin: Pseudodiaptomus 5 Left Exp2+ 3 oblong with small terminal spines:
i!t!..Lel lllarshi Wright, 1936 (Fig. 7.3 52) Pseudotliaptomus marshi (Fig. 7.352)
respec- -:..,. ~r of -+ somites; Gns otherwise formed ..... ....... 3 Sa Left Exp2+3 broad and with thick terminal spi-
=inal Gns asymmetrical with right side slightly swol- nes .......... ................ ...... ..................... ................ .6
b:md :en and left side depressed: Pseudodiaptomus 6 Left Enp nanow and elongate, extending to dis-
~ nudus Tanaka, 1960 (Fig. 7.353) tal part of Exp; distal thick Exp spin es unequal:
IJC seg- Gns symmetrical in dorsal view: Pseudo- Pseudodiaptomus richardi (Fig. 7.354)
i! !2II11: diaptomus serricaudatus (T. Scott, 1894) (Fig . 6a Left Enp spatulate, extending only half way
llr. pos- - .355 ) along Exp2; distal thick Exp spines al most equal
r::.=rior L"r highly asymmetrical, Gns almost equal in in length: Pseudodiaptomus acutu.s (Fig. 7.349)
h: :ength to remaining somites, asymmetrical in
dor::.al view, convex on left and concave on right, Family Temorid ae Giesbrecht, 1892
~ nght side with hairy conical prominence; Ur2 Females
·-·- ::regula r: Pseudodiaptomus richardi (F. Dahl,

IS9.f ) (Fig. 7 .354)
Body of varying form, in some cases rather short and
stout, in other cases comparativel y slender; antetior
~.! C r more or less symmetrical ................... .. ........ 5 Ce tends to be widest part of body. Ce separate from
- ~
Gns about as long as remaining 3 somites com- Pd I, anterior Ce unarmed or provided with 2 soft
bined, hardly expanded laterally, with minute rostral filaments or R 1-2-pointed (Temoropia); Pd4
:. E~D spines and tubercles at prox imal and di stal ang- and 5 fused, partly fused or separate. Ur of 3-4 somi-
~- Qc :es and on dorsal surface; P5 B without inner tes, CR of different structure in different genera,
process: Pseudodiaptomus acutus (F. Dahl, sometimes elongate. AI 24- or 25-segmented. A2 Enp
I 89-f) (fig. 7 .349) more or less equal in length to Exp, Exp 6-7-segmen-
),'~ Gns not as long as fo llowi ng 2 somites combi- ted. Md with broad gnathobase with I la rge tooth set
:.s .'\ l ned: P5 B with itmer processes ........ ..... .... .. ...... 6 slightly apatt fro m remaining teeth, Enp segment 1
i::.2(}- One of setae on CR thickened: Pseudodiaptomus often fused to B. Mxl Lil with 12- 15 spines and
c !Q_ hessei(Mnlzek, 1894) (Fig. 7.35 1) setae, Li2 and 3 usually with 3 setae each (with 2 and
~~- t-<S ~one of setae on CR enlarged: Pseudodiaptomus 4 setae respectively in Temoropia setosa), B and
~ charteri Grindley, 1963 (Fig. 7.350) (may be a proximal Enp segment often fused, Exp with 8- 10
bpi- ;unior synonym of P. stuhlmanni Poppe and setae, Le2 with l or no setae, Le l with 8-9 setae. Mx2
Mrazek, 1895) w ith Li I and 2 w ith 4-5, 2-3 setae respectively, Li3
~iO£ and 4 with 3 setae each, Li5 w ith 2-4 setae. Mxp of
Wes moderate size, may be slightly modified with recur-
Right P5 w ith !-segmented Enp ................... ... ..2 ved Enp or wi th fusio n between Enp segments; C with
ud3e Right P5 w ithout Enp .............. .......................... 5 2, 2-3, 2-3, setae, B with 0-3+2 setae, Enp with 2-4, 1-
~ Right P5 Enp small, parallel-sided with short ter- 4, 1-3, 0-3+1, 4 setae respectively. P l-4 with the Enp
i llf minal seta ..... .............. ... ..................................... 3 1-, 2- or 3-segmented; exopod segments 1 and 2 may
~- 2::! Right P5 Enp distally expanded with many ter- be fused. Spine and setal formula typically:
1 of minal and surface spi nes ................... ............... .4
Right P5 Exp2 outer distal spine is strong and coxa basis exopod segments endopod segments
extends beyond proximal thickening on Exp3:
Leg I 0-0/ 1 0/ 1-0 1-1 ;I- I ;l!ll,1.3/4 various
Pseudodiaptomus hessei (Fig. 7.351) Leg 2 0-1 0-0 l-1 ;1-1; ll/111,!,4/5 various
3-.!. Right P5 Exp2 outer distal spine is fine and does Leg 3 0-I 0-0 1-l ;l- 1; Tl/IJ I,l,4/5 various
:as nor extend beyond proxi mal thicken ing o n Exp3: Leg 4 0-1 0-0 I- 1;I-I ; li/JTT,l,4/5 various
Pseudodiaptomus charteri (Fig. 7.350)
954 Copepoda
P5 not natatory, 2-3 -segmented; usually without Enp; Males

both C fused with intercoxal sclerite to form common 1 Anterior Ce widest part of body, Pd4 and 5
basal segment. Ovisac present in some cases. fused; CR elongate; P5 largest on left, segment 2
enlarged; terminal spine of right leg much shor-
Males ter than its segment: Temora ...... .......................2
Segmentation of body and legs as in female except Ur 1a Pd2 is widest part of body; Pd4 and 5 separate;
of 5 somites. Right A I distinctly geniculate. P5 not CR short; P5 smallest on left, segment 2 not
natatory, usually without Enp; larger than those of enlarged; terminal spine-like segment of right
female and prehensile, often pincer-like on one side, leg longer than previous segment, small spine
2-4-segmented with common basal segment. inserted adjacent to it: Temoropia mayumbaen- ~
sis (Fig. 7.356)

It appears that this family now contains Epischura 2 Posterior corners of Pr pointed .......................... 3
Forbes, 1882; Eurytemora Giesbrecht, 1881; 2a Posterior corner of Pr rounded ......................... .4
Heterocope Sars, 1863; Lamellipodia Schmeil, 1897; 3 P5 inner appendix on left B, narrow; bend in s::::m
Temora Baird, 1850; Temoropia T. Scott, 1894. right terminal hook does not extend to distal bor- ~
Temorites has been removed to the Bathypontiidae. der of right segment 2, bend at about proximal
Of these, the first two genera (Temora and 1/5: Temora stylifera (Fig. 7.359)
Temoropia) are exclusively marine and are known to 3a P5 inner appendix on left B, broad; bend in right
occur in the South Atlantic; Eurytemora is a brackish terminal hook extends beyond distal border of
genus; whereas Epischura, Heterocope, and left segment 2, bend at about proximal l/3: ~
Lamellipodia are found in lakes. Temora discaudata (Fig. 7.357)
4 P5 left terminal segment (including stout termi-
Key to Temoridae genera and species nal spine) longer than penultimate segment;
Females armed with 2 stout terminal spines and 2 subter-
1 Anterior Ce is widest patt of body, Pd4 and 5 minal spines (not evident in figure): Temora tur-
fused; Ur of 3 somites, Gns hardly protuberant binata (Fig. 7.360)
ventrally, CR elongate; P5 uniramous, 3-seg- 4a P5 left terminal segment (including stout tem1i-
mented: Temora ... ... .......................................... 2 nal spine) shorter than penultimate segment,
1a Pd2 is widest part of body; Pd4 and 5 separate; armed with I stout terminal spine, 2 small inner
Or of 4 somites, Gns very protuberant ventrally, distal spines, and I terminal plumose seta:
CR short; P5 with !-segmented Exp and with Temora longicornis (Fig. 7.358)
seta-like Enp which not longer than Exp:
Temoropia mayumbaensis T. Scott, 1894 (Fig. Family Candaciidae Giesbrecht, 1892 ~
7.356) Females
2 Posterior comers of Pr pointed ......... ............... .. 3 Total length 2-4 mm. Body relatively robust, anterior
2a Posterior corners of Pr rounded ......... ............... .4 Ce rectangular in dorsal view with conspicuous later-
3 CR symmetrical: Temora stylifera (Dana, 1849) al constriction anteriorly. Many species possess dark
(Fig. 7.359) pigment which gives dark brown or black colour to
3a CR asymmetrical: Temora discaudata some patts of body. Ce and Pd 1 separated, Pd4 and 5
Giesbrecht, 1889 (Fig. 7.357) fused and extended into pointed, often asymmetrical,
4 One terminal caudal seta conspicuously thicke- processes, rarely rounded. R atrophied. Or of 3 somi-
ned; inner spine on terminal segment of P5 con- tes; Gns often spinose or asymmetrically swollen; Or2
siderably shorter and thinner than both terminal is sometimes asymmetrical. CR short w ith 6 setae. AI
spines: Temora turbinata (Dana, 1849) (Fig. symmetrical 24-segmented, segments 24 and 25
7.360) fused. A2 C with 1 seta; Band Enpl fused; Enpl with
4a One caudal seta longer than other but not espe- with 1 distal seta, Enp2 with 8+7 setae; Exp 6-seg- ~
cially thickened; inner spine on terminal segmented, separated from B, narrow and short, with
ment of P5 shorter than both terminal spines: Exp2 long and distal segments short with 0, 2, 1, 1, 1,
Temora longicornis (Muller, 1792) (Fig. 7.358) 2 setae respectively; Md gnathobase narrow with few
teeth, basal tooth usually divided into one or more
Copepoda 955
~lined cusps or may be simple; B large; Enp 2-seg- Key to Candaciidae genera (adapted from Grice
. :md 5 ~~ted with 0 and 6 setae respectively; Exp 4-seg- 1963)
gment 2 =errted with 1, 1, 1, 2 setae respectively. Mxl with Females
± shor- ;::::.o:,< lobes small except for very elongate Li2; Li I 1 P5 terminal segments end in processes at least one
-·-····1 -rill 6-10 spines and setae; Li2 and 3 with 3 setae of which is finger-like process and may be finely
~te: 2:h: Li4 with 2 setae; Enp 2-segmented, with 0 and serrated on one or both margins: Paracandacia
:< ~ not 5 s...."fae respectively; Exp small, with 7 setae; LeI and I a P5 terminal segments end in one or more spine-
J: :ight .: "';tbout setae. Mx2 very large; Li1 and 2 rudimen- like processes: Candacia (p. 956)
: spine ~· \\;th 3 and 2 setae respectively; Li3 and 4 with 2
.WUen- S...-"'2e each; LiS with 2 stout spines of variable length Males
~ thickness and very small spinule or proximal 1 P5 is not chelate on right but ends in long plu-
--······3 ~e may be one-halflength and considerably thinner mose seta; right A1 has no teeth in geniculate
- .... A ;:."':::m distal spine; Enp with 3 stout spines and 4 small region: Paracandacia
~in S...-"'2e. Mxp small, C with l medial and 2 terminal 1a P5 is chelate on right; right A1 has teeth on one or
~bor s..~e: B with patches of setules and 3 setae medially; more segments in geniculate region: Candacia
'Urimal E;:p 6-segmented, segment 1 almost fully incorpora-
~ into B, with 1, 2, 2, 2, 2, 2, setae respectively. Pl- Key to species of Paracandacia
~right .: ~ith 3-segmented exopods with serrated outer mar- Females
~of _:r-;s_ especially Exp2 and 3, and 2-segmented Enp I In dorsal view, Gns with spine-like protrusions
~ :3: s....~ents 1 and 2 fused). Spine and seta formula: arising from each side and directed posteriorly:
Paracandacia bispinosa (Claus, 1863) (Fig.
: :=rmi- coxa basis exopod segments endopod segments 7.375)
~nt: la In dorsal view Gns without spine-like protrusions
::....~: 0-1 0-0 1-l;l-1 ; U,I,4 0-3;1 ,2,3
~er- .............................................................................. 2
::..---; ~ 0-1 0-0 I-1 ;1-1 ;lll,l,5 0-3 ;2,2,3
cur- -"""; 3 0-1 0-0 l- l;l-l;ill,l,5 0-3;2,2,4 2 P5 tenninal finger-like process finel y seiTate
:_"'g .! 0-0 l-0 l-l ;l-l ;lli ,I,5 0-3;2,2,3 proximally; distal seta on inner margin approxi-
~zmi - mately twice as long as proximal seta; points of
glli.C1lt. ?5 uniramous, not natatory, usually symmetrical; C posterior corners of Pr directed backwards :
:.;:mer =C intercoxal sclerite fused; B and !-segmented Exp Paracandacia simplex (Giesbrecht, 1889) (Fig.
~ seta: :;cparate, B with 1 distolateral spinule; terminal seg- 7.376)
W.."'lt longest, either may end in one or more spine- 2a P5 terminal fmger-like process finely serrate dis-
~ -e processes, finger-like process, or single long seta; tally; setae on inner margin subequal, distal seta
s...."tae may or may not be present on inner margins. slightly longer than proximal seta; points at pos-
terior comers of Pr directed ventrally, scarcely
E:::::,._~Of _ IaJes visible in dorsal view: Paracandacia truncata
:s b::..~ 3ody similar to female, but Ur of 5 somites; posterior (Dana, 1849) (Fig. 7.377)
!i,."5 .f ,rk romers of Pr always asymmetrical; Gns is almost
w.iO _:':':-ays asymmetrical, bearing process on right side Males
.,. .~5 5Jllletimes with complex apex. Right A 1 geniculate, 1 Right Al segments 16 and 19-20 w ithout roun-
-
~.cal ..
' ~~-segme nted, segments 17-18 and 19-20 fused, with ded or elongate protrusions: Paracandacia sim-
.iOilll- ~ ,,;rhout denticulate segments at bend. Mouthparts plex (Fig. 7.376)
:L: l Fr2 '!:Hi P1-4 sin1ilar to the female. P5 not natatory, 4-seg- la One or both of right A1 segments 16 and 19-20
::ae. AI .wented on left and 3-segmented on right; may be che- with rounded or elongate protmsions ................ 2
~ 25 h:e on right or ending in a long feather-like seta; C, 2 Al segment 16 with rounded protrusion distally,
·! -T.:h =:.or counted in segmental complement, are small and segment 19-20 produced distally: Paracandacia
o-s...."g- ::l.~ with intercoxal sclerite. bispinosa (Fig. 7.375)
~ .._;...;..
......... 2a Al segment 16 with elongate protrusion distally,
~ fami ly contains only two genera: Candacia segment 19-20 not produced: Paracandacia
~~ ::>ana. 1846, and Paracandacia Grice, 1963 (Grice, truncata (Fig. 7.377)
- Oo:Jre ·063. revised the family).
956 Copepoda
Key to species of Candacia (adapted from Grice, Sa 4 or S spines or spine-like points present on ter- .:.:!,
1963) minal segment of PS .................................. ....... 9
Females 9 Apex of terminal segment of PS with 3 subequal
1 Posterior corners of Pr broadly rounded: spine-like points: Candacia longimana Claus, .!.
Candacia elongata (Boeck, 1872) (Fig. 7.366) 1863 (Fig. 7.368) ~
1a Posterior corners of Pr pointed or otherwise pro- 9a Apex of terminal segment of PS with I spine-
duced ................. ... ... ... ............ .. .......................... 2 like point or 2 unequal spine-like points ......... lO ~~
2 Ur2 with ventral protrusion, lamella, or spine- 10 Terminal segment of PS with 2 outer spines and
like process arising from ventral surface ..... ..... 3 2 distal spines: Candacia varicans (Giesbrecht,
2a Ur2 without lateral or ventral protrusion and 1892) (Fig. 7.374)
without spine-like process arising from ventral lOa Terminal segment of PS with 3 small outer spi-
sw·face ...................................................... .......... 6 nes, and 2 distal spines (outer spine very small,
3 In dorsal view, lateral margins ofGns somewhat inner spine large): Candacia magna Sewell,
triangular and pointed; lamella on ventral sur- 1932 (Fig. 7.369)
face of Ur2: Candacia bipinnata (Giesbrecht, I1 No spines or spine-like processes present on
1888) (Fig. 7.362) Gns; in dorsal view, Gns with distinctly convex
3a Tn dorsal view, lateral margins of Gns not poin- protrusion on each side; in lateral view, Gns with -z.
ted ......................... .. ... ................. ... .... ............... .4 ventral, rounded protmsion di rected posteriad:
4 In dorsal view, protrusion on ventral surface of Candacia catula (Giesbrecht, 1889) (Fig. 7.363)
Ur2 visible on right side of somite; in dorsal II a Spines or spine-like processes present on dorsal,
view, lateral margins of Gns with knob-like pro- lateral or ventral margins of Gns ..................... 12
trusions; apex of leg S ends in long saber-li ke 12 In dorsal view, one robust spine-like process
point: Candacia armata (Boeck, 1872) (Fig. extending obliquely posteriad from left side, and
7.361) one robust spine extending posteriad fro m right
4a In dorsal view, protrusion on ventral surface of side of Gns; both surpass posterior margin of
Ur2 not visible; apex of leg S ends in 2 or more Gns: Candacia pachydactyla (Dana, 1849) (Fig.
points .............................................. ................... S 7.37 1)
S Tn lateral view, ventral protrusion of Ur2 direc- 12a In dorsal view, spines from left and right sides of
ted obliquely anteriad; terminal segment of PS Gns smaller and not reaching posterior margin
with 2 small , outer, spine-like points and 2 ofGns .......... ..................... ....... ................ .... .... l 3
small, subequal, spine-like points at tip: 13 fn lateral view, small protuberance arising from
Candacia paenelonginuma Fleminger and ventral side of Gns near posterior margin; distal
Bowman, 19S6 (Fig. 7.372) 2 inner setae on terminal segment of PS, coarse
Sa In lateral view, ventral protrusion of Ur2 direc- and unequal in length: Camlacia ethiopica
ted obliquely posteriad; terminal segment of PS (Dana, 1849) (Fig. 7 .367)
with 2 small, outer, spine-like points and 3 13a In lateral view, no protuberance on ventral sur-
spine-like points distally, middle one of which is face of Gns; distal 2 inner setae on terminal seg-
longest: Candacia cheirura Cleve, 1904 (Fig. ment of PS, thin and approximately equal in
7.364) length: Candacia norvegica (Boeck, 1864) (Fig.
6 No inner edge setae present on tetminal segment 7.370)
ofPS .................. .. .......... ......... ............ ....... .........7 J
6a 2 or 3 inner edge setae present on terminal seg- Males
ment of PS ....................................................... 11 I Left posteri or comer of Pr rounded: Candacia [3
7 Spine-like process present on ventral side of elongata (Fig. 7.366)
Gns: Candacia curta (Dana, 1849) (Fig. 7.36S) I a Left posterior corner of the Pr pointed .............. 2
7a No spine-like process present on ventral side of 2 ln dorsal view, Gns without process or prot-
Gns ..................................................................... 8 rusion ................................................................. 3 .]
8 3 spine-like points present on terminal segment 2a Tn dorsal view, Gns with process or protrusion 4
of PS: Candacia tenuimana (Giesbrecht, 1889) 3 Proximal spine on Mx2 LiS considerably thicker
(Fig. 7.373) than distal spine: Candacia catula (Fig. 7.363)
Copepoda 957
Le'r- 3a Proximal spine on Mx2 LiS not notably thicker 14 In dorsal view, process on Gns divided into two
___Q
than di stal spine: Candacia varicans (Fig. lobes: Candacia norvegica (Fig. 7.370)
~-ua] 7.374) L4a In dorsal view, process on Gns knob-like:
Claus.. 4 Right Al with segments 2 and 3 fused ... .......... 5 Candacia longimana (Fig. 7.368)
4a Right A 1 with segments 2 and 3 separate ....... II
spoc- 5 Right Al with segments 17 and 18 fused ......... 6 Family Pontellidae Dana, 19S3
__ w Sa Right A I with segments 17 and 18 separate ..... ? References: Boxshall (in prep.)
lli!S and 6 In dorsal view, process on right side of Gns Females
i.Techi.. small, consisting of rounded knob in front of Ce and Pd 1 separate, sometimes with hook on lateral
which is pointed projection: Candacia ethiopica borders, anterior Ce sometimes has anterior crest; Pd4
bspi- (Fig. 7.367) and 5 fused or separate, usually with asymmetrical
small 6a In dorsal view, process on right side of Gns expansions. R bifurcate, sometimes with widened
Sen·ell. large, consisting of single broad and rounded base incorporating lens. Eyes usually large, with I or
projection: Candaciapachydactyla (Fig. 7.371) 2 pairs of dorsal chitinous lenses and I ventral lens.
i:%1: on 7 Right A I with segments 19 and 20 fused or par- Ur of 1-3 somites, often asymmetrical; genital appa-
roo,· ex tially fused ......................................................... 8 ratus comprising a common genital pore contained
\\itb 7a Right A 1 with segment 19 and 20 separate .... I 0 within a median genital aperture, copulatory pore con-
~ 8 In lateral view, distal end of rig ht posterior cor- tained within median genital ape11ure; CR with up to
- .353) ner of Pr truncate, extending beyond posterior 6 setae. A 1 16-24-segmented. A2 C and B separate
·~ border of Gns: Candacia bipimzata (Fig. 7.362) with I and 2 setae respectively; Enp much larger than
8a In lateral view, distal end of right posterior cor- Exp; Exp usually 5-segmented, segment 2 elongate;
ner of Pr not truncate, may or may not extend Exp with 1, 3, 2, 2, 4 setae in Anomalocera patersoni,
beyond posterior border of Gns ......................... 9 terminal segment atrophied or rudimentary; proximal
9 In lateral view, right postetior Pr comer not segment of Enp with 2 setae, more or less fused to B,
~.;::::;. of reaching posterior border of Gns and turned compound distal segment bilobed with 9, 7 setae or
fig. upwards: Cantlacia curta (Fig. 7.36S) with setation reduced. Md gnathobase with 5-7 teeth,
9a In lateral view, right posterior Pr corner reaching B with 4 setae, Exp 5-segmented with 1, 1, 1, 1, 2
~of posterior border of Gns and turned slightly setae and Enp 2-segmented with 3 and 6 setae and of
~ downwards: Candacia armata (Fig. 7.361) similar size. Mx 1 Li I large with 8-1 0 spines and
.1..~ l0 Tip of "thumb" of chela on right P5 reaching to setae; Li2 as long as Li I and with 3 setae; Li3 short
g fnxn about midlength of "finger": Candacia armata with 3 setae; Li4 with 2 setae; proximal Enp segment
: 64:3.1 (Fig. 7.36 1) fused to B, Enp segments with 2, 2, S setae or redu-
iCOOrSC lOa Tips of "thumb" and "finger" of right P5 sube- ced; Exp relatively small, sometimes atrophied with 9
riDpiL:a qual: Candacia magna (Fig. 7.369) setae; Le2 with I seta; Le1 with 7-8 setae. Mx2 with
II In dorsal view, distal end of process on right side long, strong setae although sometimes rudimentary on
ofGns endi ng in point.. ................................. .. l2 proximal part of limb; Li 1-S usually with 4, 3, 3, 3, 2
~ Ia In dorsal view, distal end of process on right side setae; Enp setation 1, 1, 2, 2. Mxp small, 5- or 6-seg-
~ of Gns rounded or lobate ......................... ........ 14 mented with C large with inner border lobed and with
~2 Distal segment of left PS longer than penultima- 0, 2, 2, 2/3 long setae; B and Enp short, B with 2 or 3
te segment: Candacia cheirura (Fig. 7.364) setae, free Enp 3- or 4-segmented with 2, 2, 2+ I, 3
l 2a Distal segment of left PS sho1ter than penultima- setae or with setation reduced. P 1-4 Exp 3-segmented.
te segment ................ ...... ................................ .. l3 Enp of P 1, 2- or 3-segmented; of P2-4 2-segmented.
ruliJcill l3 In dorsa l view, process on Gns di rected out- Spine and seta formula typica lly of the following
wards with distal end curved posteriorly; right fo rm, although setation may be reduced:
posterior corner of Pr notched at tip: Candacia
r~- tenuimana (Fig. 7.373) coxa basis exopod segments endopod segments
l3a In dorsal view, process on Gns not curved and
~
Leg I 0-1 0-0 1-1 ; 1-1 ; ll, I ,4 0-1 ; 0-2 ; I ,2,3
directed somewhat obliquely posteriad; right Leg 2 0-1 0-0 1-1 ; 1- 1; ll1 ,1,5 0-3; 2,2,4
posterior Pr corner not notched at tip: Candacia Leg 3 0- 1 0-0 1-1 ; 1- 1; 111 ,1,5 0-3; 2,2,4
paenelongimana (Fig. 7.372) Leg 4 0-1 0-0 1-1 ; 1-1 ; 111,1,5 0-3 ; 2,2,3
958 Copepoda
PS not natat01y, C fused with the intercoxal sclerite; B lens absent; dorsal cuticular lenses and ventral
with l outer seta; Exp 2-segmented, segment 1 with 2 lens.feeb ly developed ....................................... .4
outer spines and inner distal spinous process, segment 4 A2, Md palp, and P5 B with normal setae; CR dis-
2 with 1 or 2 spines plus an apical sp inous process, or tinctly separated from anal somite: Pontellopsis
Exp may be !-segmented and anned with spinous pro- (p. 960)
cesses or tapering to a point; Enp !-segmented with 4a A2, Md palp, and P5 B with well developed,
simple or bifid apex, or absent. long plumose setae; right CR indistinctly separa-
ted from anal somite: Pontellina (p. 960)
Males 2.
Ur of 4-5 somites, genital open ing on the left; free Ur Males
somites sometimes w ith asymmetrical processes. A I I Anterior Ce w ith 2 lenses on dorsal surface ..... 2
prehensile on the right, middle section swollen, termi- la Anterior Ce without lenses or lateral hooks ..... A
nating in 2-4 segments distal to the joint. Mouthparts 2 Anterior Ce without lateral hooks: Labidocera
identical to those of the female. P5 asymmetrical, C (part) (p. 959)
and intercoxal sclerite usually fused to form transver- 2a Anterior Ce with lateral hooks .......................... 3
se plate, sometimes with C free on right side only; 3 Prehensile A I with terminal part 2-segmented;
right leg subchelate, comprising unarmed C, B with 1 left P5 without Enp: Pontella (p. 960)
seta, and 2-segmented Exp: Expl-2 swollen with 3a Prehesile A1 with term inal part 4-segmented;
outer process and spine, Exp3 forming subchela. Left left P5 w ith rudimentary Enp: Labidocera (part)
PS comprising B with 1 seta; Exp I elongate with 1 (p. 91)
seta; Exp2 and 3 partly fused with up to 4 outer and 4 Ur3 asymmetrical on right: Pontellopsis (p. 961)
distal spines on apical segment. 4a Ur3 symmetrical ................................................ 5
5 Outer seta on P5 B long and very plumose,
The fami ly includes Anomalocera Templeton, 1937; extending at least half way along Exp l:
Calanopia Dana 1853; Epilabidocera C. B . Wilson, Pontellina (p. 960)
1932; lvellopsis Claus, 1893 (see Wickstead and Sa Outer seta on P5 B short: Calanopia (p. 958)
Krishnaswamy, 1964); Labidocera Lubbock, 1853;
Pontella Dana, 1846; Pontellina Dana, 1853; Key to species of Calanopia
Pontellopsis Brady, 1883. Of these genera Females
Anomalocera, Epilabidocera, and lvellopsis have not l P5 with I segment distal to B ........................... 2
been recorded from the South Atlantic. Ia P5 with 2 segments distal to B: Calanopia ellip-
tica (Dana, 1849) (Fig. 7.379)
Key to Pontellidae genera (from Silas and Pillai, 2 P5 terminal segment bifurcate distally, inner
1973) spine long and spinulose: Calanopia minor A.
Females Scott, 1902 (Fig. 7.3 80) --:~
I Mxp with 6 distinct segments; P 1 Enp 2-seg- 2a P5 terminal segment trifid distally, midd le point
mented; Ur of2-3 somites ................................. 2 longest: Calanopia americana F. DaW, I 894
Ia Mxp with 5-7 segments; PI Enp 3-segmented; (Fig. 7.378)
U r of 1-3 somites ............................................... 3
2 One pair of dorsal cuticular lenses and 1 ventro- Males
median lens present on Ce: Labitlocera (p. 91) 1 Distinct " thumb" present on right P5 Exp l -2
2a Dorsal cuticular lenses and ventromedian lens ........................................................................... 2
absent on Ce: Calanopia (p. 90) Ia "Thumb" on right P5 Expl-2 rudimentary or
3 Body relatively large; Pd4 and 5 usually separate; absent: Calmzopia elliptica (Fig. 7.379)
Pr more or less of same width throughout; lateral 2 Distal border of right P5 B extends beyond dis-
cephalic hooks prorninant; one pair of dorsal cuti- tal border of left leg B: Calanopia americana
cular lenses and rostral lens well developed; (Fig. 7.378)
medioventral eye prominant: Pontella (p. 91) 2a Distal border of right P5 B does not extend much
3a Body relatively small; Pd4 and 5 fused; Pr more beyond middle of left B: Calanopia minor (Fig.
broadened in mid-lateral margin and narrowed 7.380)
anteriad; lateral cephalic hooks absent; rostral
Copepoda 959
~~ Key to species of Labidocera lb Left P5 B with short process (Fig. 7.385) ......... 2
Females 2 Right P5 Exp3 length about half width of Exp 1-
'CRCis- Ce with lateral hooks .. ....................................... 2 2: Labidocera scotti (Fig. 7.388)
JI!IIIJpsis ia Ce without lateral hooks .................. ... .......... .....3 2a Right P5 Exp3 m uch longer than half width of
, Anterior Ce tri angular in dorsal view; posterior Exp l-2: Labidocerajluviatilis (Fig. 7.385)
..:loped. prosomal corners extending into points on both 3 Left P5 B with broadly-based, triangular, inner
sepzra- sides: Labidocera wollastoni (Lubbock, 1857) distal process which extends just beyond distal
(Fig. 7.389) border ofExpl: Labidocera danvinii (Fig. 7 .383)
2a Anterior Ce rounded in dorsal view; posterior 3a ' Left P5 B with this process extending beyond
prosomal corners rounded on left and with an distal border of who le Exp and decorated with
lee _ .. ...2 elongate, ventrally directed spine on right: spinules .... .......... ............... .. ............................... 4
5-.~-~ Labidocera minuta Giesbrecht, 1889 (Fig. 7.386) 4 Left P5 Exp 1 a nd 2-3 almost equal in length;
i:!!:IKua Anterior Ce with crest, visible in dorsal view as right Exp l -2 with very long " thumb" and trian-
an anterior spine ...... ......................................... .4 gu lar projection with constriction a t midlength:
~3. Anterior Ce w ithout crest, rounded in dorsal Labidocera wollastoni (Fig. 7.389)
ce-..ili!'d.: Yiew ................................................................... 5 4 a Left P5 Expl more tha n twice as long as Exp2-
.!. Posterior Pr comers divergent; Ur2 w ith right 3; right Expl -2 with very short "thumb" and
~eti side, posteriorly directed, posterodorsal spine broad rounded projection: Labidocera acuti-
p2.ri which covers some of right CR : Labidocera frons (Fig. 7.382)
acutifrons (Dan a, 1849) (Fig. 7 .382) 5 Right P5 B long, more than twice length of
~ c· , ~3. Posterior Pr corners directed posterio rly; Ur2 Expl: Labidocera acuta (Fig . 7.381)
.:;
without any dorsa l projections: Labidocera 5a Right P5 B relatively short, shorter than or only
osc. acuta (Da na, 1849) (Fig. 7.381) slightly longer than Exp1 .................................. 6
~"{p:: .:;
Posterior Pr comers produced into large rounded 6 Left P5 term inal segment with 3 distal rounded
lobes w ith lateral spines in dorsal view: lobes: Labidocera minuta (Fig. 7.386)
5S) Labidocera detnmcata (Dana, 1849) (Fig. 7.384) 6a Left P5 terminal segment with 3-4 distal spines
5:! Posterior Pr corners produced into points, with- ........ ..... .............. ... .................................. ........... 7
out posterior lobes ............................................. 6 7 Right P5 Exp 1-2 without process between
Posterior Pr corners produced into very short, "thumb" and distal border of segment:
slightly divergent points: Labidocera nerii Labidocera detruncata (Fig. 7 .384)
~ (Kroyer, 1849) (Fig. 7.387) 7a Right P5 Expl -2 with elongate, distally rounded
--
;,a Posterior Pr comers produced into long points process w hich originates very close to " thumb":
r mner .............. ... ................. ......... .. .... ......... ... ....... ....... 7 Labidocera nerii (Fig. 387)
_-\_ P5 Exp terminal segment with bifurcate tip ........ 8
P5 Exp with 3 terminal points, innermost smallest Key to species of Pontella
~ and middle one largest: Labidocera scotti Females
·~.! Giesbrecht, 1897 (Fig. 7.388) 1 Ur of 3 so mites .................................................. 2
'"' In dorsal view, Gns w ith w ide, rounded projec- 1a Ur of 2 somites ...................... ........ .................... 4
tion on right: Labidocera darwinii Lubbock, 2 P5 Enp tapering to point: Pontella marplatensis
1853 (Fig. 383) Ramirez, 1966 (Fig. 7.395)
Expl -2 .,. In dorsal view, Gns without projection on right, 2a P5 Enp bifurcate d istally ... ............. ................ ... 3
symmetrical: Labidocera jluviatilis F. Dahl, 3 P5 Enp inner distal spine much larger than outer
~' OC 1894 (Fig. 7.385) spine; Gns w ith right side ventrolateral, narrow,
process which extends to posterior border of
rc Cis- ... wes Ur2: Pontella atlantica (Milne-Edwards, 1840)
riauul left P5 B with one of distal corne rs produced (Fig. 7.390)
into large process w hich extends at least beyond 3a P5 Enp inner distal spine smaller than outer
=h distal border ofExp 1 (Fig. 7.3 82) .................... 3 spine; Gns with small left ventral process which
r (~Ig 1:! left P5 B not especially produced distally (Fig. is not visible dorsally: Pontella lobiancoi (Canu,
- .387) ............. ................ ....................... ............. 5 1888) (Fig. 7.394)
960 Copepoda
4 CR subequal... ................ ................... .... .............5 5 Chelate right P5 Exp1-2 with the I large projec-
4a CR unequal, enlarged on right side ................... 6 tion in form of broad, obliquely truncate lobe;
5 Gns with sides almost parallel in dorsal view; P5 Exp3 expanded proximally into broad, obliquely
Exp without inner edge spines: Pontella truncate lobe: Pontella atlantica (Fig. 7.390)
gaboonensis T. Scott, 1894 (Fig. 7.393) 5a Chelate right P5 Exp1-2 with 1 large projection
5a Gns with widest part anterior in dorsal view; P5 in form of broad rounded lobe; Exp3 not proxi-
Exp with 2 long inner edge spines: Pontellafera mally expanded: Pontella lobiancoi (Fig. 7.394)
Dana, 1849 (Fig. 7 .392)
6 Gns dorsal surface covered in irregular ridges Key to species of Pontellina
and raised areas; left comer of Pr extends to pos- Females
terior border of Gns: Pontella spinipes 1 Posterior Pr corners extend more than 1/2 way
Giesbrecht, 1889 (Fig. 397) along Gns; P5 B seta extends well short of base
6a Gns is not covered in irregular ridges but has of first outer edge spine on terminal segment:
specific processes; left corner of Pr does not Pontellina plumata (Dana, 1849) (Fig. 7.399)
extend to posterior border of Gns ............... .. .... 7 1a Posterior Pr corners very short, extending only
7 Gns with 2 rounded lobes on left, posterior most about 1/3 of way along Gns; P5 B seta extends
of which is broad and covers much of left CR; beyond base of first outer edge spine on terminal
right side with rounded lobe and posterolateral segment: Pontellina platychela Fleminger and
projection which is broad in lateral view: Hiilsemann, 1974 (Fig. 7.398)
Pontella diagonalis Wilson, 1950 (Fig. 7.391)
7a Gns left side with small dorsolateral projection Males
at midlength and posterodorsal , narrow, hooked I Posterolateral corners of Pr, in lateral view,
projection covering part of left CR; right side rounded and lacking denticle; right chela with
with small dorsal projection just posterior to strong distal expansions, in anterior-posterior
midlength: Pontella securifer Brady, I 883 (Fig. direction, of both Exp 1-2 and Exp3: Pontellina
7 .396) platychela (Fig. 7.398)
Ia Posterolateral corners of Pr, in lateral view,
Males (unknown for P. diagonalis and P. spinipes) angular and bearing denticle; rig ht chela Exp 1-2
1 Chelate right P5 Exp 1-2 with 1 or 2 large pro- extending distally as relatively slender, digiti-
jections besides "thumb" ..... .............................. 3 form process opposing apex of narrow, curved
Ia Chelate right P5 Expl-2 without projection or Exp3: Pontel/ina plumata (Fig. 7.399)
with low, triangular raised area at midlength
between tip of "thumb" and distal border of seg- Key to species of Pontellopsis
n1ent ..... ................... ................. .......................... 2 Females
2 Left P5 terminating in curved spine, Exp2-3 with 1 Ur apparently of 1 somite, in dorsal view ......... 2
2 proximal patches of hairs: Pontella marpla- 1a Ur distinctly of 2 somites ........ ..... ..................... 3
tensis (Fig. 7.395) 2 Gns with large, hairy left side projection directed
2a Left P5 terminating in straight spine, Exp2-3 posteriorly, indistinctly separated from somite 2
with entire inner border hairy : Pontella in dorsal view: Pontellopsis villosa Brady, 1883
gaboonensis (Fig. 393) (Fig. 7.403)
3 Chelate right P5 Exp 1-2 with 2 large projections 2a Gns lateral borders not noticably swollen, right
besides "thumb", one long and triangular, the side with 2 unequal spine-like processes at about
other a smaller rounded lobe: Pontella securifer midlength: Po1ttellopsis brevis (Giesbrecht,
(Fig. 7.396) 1889) (Fig. 7 .400)
3a Chelate right P5 Exp l-2 with I large projection 3 Gns with naked, globular projection on right;
besides "thumb" ................................................ .4 anal somite projecting posteriorly as far as pos-
4 Chelate rightP5 Expl-2 with I large projection in terior border of CR: Pontellopsis perspicax
form of elongate, narrow lobe, rounded distally; (Dana, 1849) (Fig. 7.401)
Exp3 expanded disally: Pontellafera (Fig. 7.392) 3a Gns asymmetrical swollen on each side, each
4a This combination of characters absent .............. 5 swelling bearing 3 spines; anal somite not pro-
Copepoda 961
~ projec- jecting posteriorly: Pontellopsis reg{l/is (Dana, and 2 w ith 8-9 and 1-0 setae respectively. Mx2 L i 1-5
c- lobe: 1949) (Fig. 7.402) with 3-4, 2-3, 2-3, 1-3, 1-2 setae respectively; Enp
~liq-;..tely with 4-5 setae. Mxp reduced and highly modified, C
. 390) . lales with 5-6 long setae; B with 1 short thick setae; Enp
~ection Posterior Pr comers almost symmetrical, both with fused segments with 4-5 short thick setae. Pl-4
Ill proxi- points long, extending almost to posterior border slender and delicate with long natatory setae; di stola-
~· /39-l) ofUr3: Pontellopsis villosa (Fig. 7.403) teral borders of Exp of P2-4 expanded into conspi-
1::! Posterior Pr comers asymmetrical. .................... 2 cuous tooth; articulated spine not present in th is posi-
-
.. Left posterior Pr corner extending almost to pos- tion. P 1 B may have very small outer edge spine or
rerior border of Ur3, right corner extending half seta; terminal Exp spine of P2-4 with outer edge teeth.
:! way way along anal somite; Ur somites slightly Spine and seta formula:
--ba_~ asymmetrically developed: Pontellopsis brevis
~t: , Fig. 7 .400) coxa basis exopod segments cndopod segments
- _: QQ I Left posterior Pr comer extending to posterior
Leg I 0-0 1/0-0 1- 1; 1-1 ; 11,1,4 0-1 ; 1,2,3
g only border of Ur2, right comer extending to posteri- Leg 2 0-0 0-0 0- 1;0- 1; 0,1,5 0-2; 1,2,4
~.ends or border of a nal somite; Ur3 asymmetrically Leg 3 0-0 0-0 0- 1; 0-1 ; 0,1.5 0-2; 1,2,4
~ developed on right ........ .. ....... ........................... .3 Leg 4 0-0 1-0 0-1 ; 0- 1; 0,1,5 0-2 ; l ,2,3
~and ~
Cr3 with large, naked protruberance on right
side: Pontellopsis perspicax (Fig. 7.401) Leg 5 usually uniramous, very small, symmetrical, 3-
~ Cr3 with spinulose swelli ng on right: segmented, C and intercoxal sclerite may be fused;
Pontellopsis regalis (Fig. 7.402) both B may be fused to each other or separate, B with
If '1.ew. 1 outer seta; Exp in the form of spine or elongate seta;
~ \\;th f~;l y Acartiidae Sars, 1903 small Enp present in Acartiella.
110S4-"'rior !Fanales
mdlina ,~ .. n. more or less slender copepods. Single eye pre- Males
~ Ce and Pd I separate, Pd4 and 5 always fused, Ur of 5 somites. A 1 usually prehensile on right side
~ ,;ew. s:erolatera l corners ofPr rounded or pointed. Ur of only, with 12 - 18? segments; middle segments may be
I E.'Ql-2 :: smnires. anal operculum may be on anal somite or greatly inflated. Mouthparts more or less as in female.
;., ciigiti- ,......., may open between last two urosomites into dor- P5 uniramous, asymmetrical, attached to plate formed
. c::r.·ed po,·e on anal somite; CR generally slightly asym- from fusion of C and intercoxal sclerite; B usually
~cal. short or long, sometimes fused with ana l asymmetrically developed with 1 outer dista l seta; left
s:x::.ire. w ith 6 setae. Without R and with or w ithout P5 with 2-segmented Exp (3-segmented in Paracartia
~ed filaments. Upper lip large, prominent and trilo- africana), Exp2 w ith variously decorated tip; right P5
re-.:.. Al 17-22-segmented, many segments with long usually with 3-segmented Exp, Exp 1 and 2 often with
.._ .... .2 R't:!e• .-\2 Enp 3-segmented; B (wit h I seta) and Enp 1 inner processes; Exp3 opposed to process on Exp2 to
--- ___ 3 C:s_~ resulting allobasis bears 9 setae arranged in f01m type of subchela.
C:ec-..ed ·..._..,. . ximal group of 8 and isolated d ista l seta, 8 proxi-
g:::::e ~ setae interconnected at their bases by tiny tendi- This fami ly is now considered to include Acarfia
_ ISS3 ._s strands extending from single muscle inserted Dana, I 846; Acartiefla Sewell, 19 14; Paracartia T.
.::;~em ro last seta enabling the 8 setae to be moved Scott, 1894; and Paralabidocera Wo lfenden, 1908
!C..::_ght zs v-ogle unit (Acartia); Enp2 with 6-9 distal setae; (see Bradford, 1976), of which Acartia and
a:~ so:::::=: rerminal segment with 7 terminal setae; Exp Paracartia have been recorded from the South
~echL. -:....:. reduced number of segments: 4-segmented with Atlantic .
.:... .:.. 3 setae respectively. Md w ith well-developed
:!ghi: :o-=•=:...Obase with large separate tooth on one border, Key to Acartiidae genera and species of Paracarfi{l
:zs i)OS- ;. B wirh 1-2 setae; Enp !-segmented with 2+8-9 Females
nFar ~; Exp 5-segmented with 1, I, I, I, 2 setae respec- 1 R absent or present as 2 filaments, posterior cor-
~-. ~lxl with reduced lobes and setation; Lil short ners of the Pr rounded or with small points, Gns
~. e:!L"t :iliout 6-8 spines and setae, Li2-4 w ith 0, 3, 1 only slightly wider than following Ur somite:
~ ~ Enp appa rently absent; Exp with 2+5 setae; Le I Ac{lrfi{l
962 Copepoda
la Rand filaments present, posterior comers of Pr about 4 posterodorsal spines on Gns and 2 on :t
enlarged into wings which may be asymmetrical, Ur2; A1 segment 1 with coarse anterodistal
Gns swollen: Paracartia africana Steuer, l 91S spine: Acartia (Acartia) danae Giesbrecht, 1889
(Fig. 7.4 11) (Fig. 7.406)
6a PS terminal spine-li ke segment with small, blunt,
M ales outer protrusion just proximal to midlength;
1 Left PS Exp 2-segmented: Acartia about 2 posterodorsal spines on Gns and 2 simi-
la Left PS Exp 3-segmented; Exp2 with outer dis- lar sized spines on Ur2; A1 segment 1 without
tal projection which extends 2/3 along Exp3; coarse anterodistal spine: Acartia (Odontacartia)
Exp3 with outer patch or hairs: Paracartia afri- centrura Giesbrecht, 1889 (Fig. 7.40S)
cana (Fig. 411)
Males
Key to the species of Acartia 1 Posterior comers of Pr rounded ............ ........ .... .2
Females Ia Posterior comers ofPr pointed ...................... .. .. S
1 Posterior corners of Pr rounded ........................ .2 2 Left PS Exp2 with 3 terminal spines as well as
1a Posterior comers of Pr pointed .......................... 4 another plumose spine inserted just distal to
2 PS B with conspicuous inner lobe, terminal midlength; Acartia (Acartia) negligens (Fig.
spine-like segment with coarse spinules for short 7.408)
distance at about midlength: Acartia 2a Left PS Exp2 with at most 1 terminal spine ..... 3
(Acanthacartia) tonsa Dana, 1849 (Fig. 7.410) 3 Left PS Exp2 with 1 very small terminal spine
2a PS B without inner lobe, terminal spine-like seg- and very long inner spine, inserted at midlength,
ment without coarse spinules ............................ 3 and which extends almost to dista l border of its
3 Surface of Gns and Ur2 naked, but with nume- segment; posterior corner of Pr and Ur2-4 bor-
rous coarse spines along posterodorsal border: dered by spines: Acartia (Acantlracartia) plu-
Acartia (Acanthacartia) p lumosa T. Scott, 1894 m osa (Fig. 7 .409)
(Fig. 7 .409) 3a This combination of characters absent.. .. .......... 4
3a Surface of Gns and Ur2 covered in rows of very 4 Right PS Expl without inner edge spine; left
small spinules: Acartia (Acanthacartia) bijilosa Exp2+3 with 2 fine spines inserted subterminal-
(Giesbrecht, 188 1) (Fig. 7.404) ly; Uri and 2 decorated with rows of fine hairs:
4 Posterior corner of Pr with very small point; Acartia (Acanthacartia) tonsa (Fig. 7.410)
outer distal plumose seta on PS B, S times as 4a Right PS Exp1 with inner edge spine; left
long as terminal spine-like segment which has Exp2+3 with 1 fine terminal spine and 1 broad,
coarse spinules fo r short distance at about mid- truncate spine inserted at about midlength:
length: Acartia (Acartia) negligens Dana, 1849 Acartia (Acanthacartia) bijilosa (Fig. 7.404)
(Fig. 7 .408) S Posterior comers of Pr extend into long points
4a Posterior corner of Pr with large point; outer dis- which reach more than half way along Ur2 :
tal plumose seta at most, 3 times as long as ter- Acartia (Odontacartia) lilljeborgii (Fig. 7 .407)
minal spine-like segment.. ................................. S Sa Posterior corners of Pr either do not reach poste-
S Point on posterior corner of Pr long, extending rior border ofUrl or extend just beyond this bor-
3/4 of way along Gns; Gns and Ur2 bordered der ................ ................ ........ .................... .......... 6
with numerous posterodorsal spinules; PS term i- 6 Posterior border of Ur2-4 with few large coarse ..........~
nal spine-like segment bordered distally with spines; posterior border of Pr wi th pair of addi- · )
small spinules: Acat1ia (Odontacartia) lilljebor- tional accessory spines as well as larger spines
gii Giesbrecht, 1889 (Fig. 7.407) which extend j ust beyond posterior border of
Sa Point on posterior corner of Pr extending 1/3 of Ur 1; left PS Exp2+3 without terminal spines:
way along Gns; Gns and Ur2 bordered by at Acartia (Odontacartia) centrura (Fig. 7.40S)
most 4 posterodorsal spines; PS terminal spine- 6a Posterior border of Ur somites almost naked;
like segment otherwise decorated ..................... 6 spines on posterior corner of Pr do not extend to
6 PS terminal spine-like segment with coarse spi- posterior border of Url ; left PS Exp2+3 wi th 3
nules for short distance just distal to midlength; terminal spines as well as pai r of subterminal
Copepoda 963
=:L .2 on spines, one of wh ich is plumose: Acartia Males

~12.1 (Acartia) danae (Fig. 7.406) Posterior corners of Pd5 with small rounded or poin-
i;o:_ 1889 ted extremities or without protrusions. Ur of 5 somi-
~amily Tortanidae Sars, 1902 tes, often asymmetrical; genital aperture located ven-
alunL References: Brodsky ( 1950), Boxshall (in prep.) trolaterally on left posterior border of Gns, Ur2
Gffigth: Females sometimes with posterolateral process. Right A I pre-
. :simi- Small copepods with one median eye; no rostrum; a hensile, its middle pari widened and sometimes equip-
~Ul 5elllicircular plate, thickly covered with short bristles, ped with denticulate plate. P5 asymmetrical, unira-
w:ania) ;.:.l3 before the labrum. Ce and Pd1 separate; Pd4 and 5 mous; 3-segmented on right, segment 1 unarmed,
~d or separate, posterior comers of Pd5 w ith round- segment 2 often expanded into medial or distal lamel-
=rl or pointed projections which are often asymmetri- late process armed with up to 3 setae, segment 3 for-
.-::31 Ur of 2-3 somites; Gns often asymmetrical, bear- ming subchela typically curved and armed with 3 or 4
I :::g common genital aperture located medially on the small setae. Left P5 4-segmented; segment I unar-
- ·· --·--·-
- ...... 5 ·erma] smface, copulatory pore contained within med, segments 2 and 3 elongate each w ith inner and
~n as :::ledian genital aperture. CR sometimes asymmetrical, outer seta, segment 4 curved, typically armed with 2
vJ ro .me larger than other, often fused to anal somite, with setae on inner margin, I apical seta and 2 setae on
o- Fig. ':l setae. A l 12- to 15-segmented with proximal seg- outer margin; inner seta on segment 3 often carried on
:::Jents fused. A2 C and B separate, Enp and B often slender process. Eggs probably released into water.
~----.3 :!sed: with Exp indistinctly 3-segmented, typically
:=.!spine ~-:-a.h segment I unanned, long middle segment with up This family is monotypic. Tortanus has been review ed
.::ength. ~ 3 setae, and short apical segment with 2 setae; Enp (Ohtsuka, 1992) and has been divided into the subge-
;:::- o;- its .::-segmented with 6 terminal setae. Md with elongate nera Tortanus Giesbrecht, 1898; Eutortanus Smirnov,
~~ oor- ~p: unarmed B; Enp 2-segmented with 0 and 6 setae; 1935; Acutanus Ohtsuka, 1992; and Atortus Ohtsuka,
~ plu- .::_~ indistinctly 1-4-segmented with 5 setae in total. 1992. Only one species, Tortanus (Tortanus) gracilis
hl with very much reduced number of lobes (Lil (Brady, 1883), has been taken in the South Atlantic
___A 7-d distal segment present); Li 1 with up to 13 spines (Fig. 7.412).
=..c: leir ...-d setae, distal segment elongate bearing 3 powerful
'r.;"inal- ·:law-like setae around apex, and may have up to 7 Family Bathypontiidae Brodsky, 1950
r~: ·,;ymes subapically. Mx2 with Li 1-4 reduced, Li5 with References: Sars (1924-25), Brodsky ( 1950), Deevey
: s....~e. 2 of them claw-like; Enp with 6 claw-like setae ( 1973).
r: teft ~:-::s 1 or 2 smaller setae. Mxp reduced, with 2-5 long Females
i:road. :-;inuJate setae on first segment; the distal part of limb Ce and Pdl separate, Pd4 and 5 separate, often pro-
~~rill: :.-segmented, bearing 3 or 4 setae on inner margin and duced backward with rounded or acute p oint. R usual-
--=-'-- on outer margin of distal segment. Swimming legs ly without filaments, in form of blunt or short process,
f""'!TTl 3-segmented Exp; Enp of PI 2-3-segmented, P2- can be bifurcate at its tip. Ur of 4 somites, Gns with
--r;xi.:ILS
-::;.
··-").
'-- .i...- - .!... .::-segmented. Spine and seta formula: slightly protruding ventral process; CR short with 4-5
setae. AI 23-segmented, segments 8-9 and 24-25
- pos:-r- coxa basis cxopod segments endopod segments fused, 24-segmented in (Ternorites discoveryae) with
,~ numerous aesthetascs proximally, may be asymmetri-
_ _6 0- 1 0-0 0- 1; 0-1; H*,T,4 0-3; 1,2,3
- 0-3; 1/2,2,3/4
cally developed (A lloiopodus p inguis). A2 (based on
0- 1 0-0 T- 1; 1-1 ; 111*,1,5
ax!:"',e
- 0- l 0-0 1-1 ; l- 1; IIT,l,S 0-3 ; 2,2,3/4 Bathypontia) rami almost equal in length; C and B
~
li zrid.i- ~- 0- 1 1-0 !-I ; l-1 ; Ill,l,5 0-3; 1,2,3 with 1 and 2 setae respectively, Enpl with 1-2 setae
- ...._ ""'
~ just distal to midle ngth (l rather small), Enp2 with 7+9
~n sometimes reduced on segments indicated( *)
oier o:· setae; Exp 7-segmented with 0, 1, 1, 1, 1, 1, 1+3 setae;
sv:::.es~ other genera have various reductions in segment num-
-~ :=~e P5 not nata tory, uniramous, 2-3-segmented; C bers and setation (Exp 6-segmented in Alloiopodus,
~a~- inrercoxal plate fused; B with outer seta; 1 free
a::;C Ternorites discoveryae, and Zenkevitchiella. Md
~....;m ~ segment present, sometimes in form of curved (based on Bathypontia) gnathobase with large tooth
~= ~g process, typica lly w ith 1 outer margin spine often separated by some distance from numerous small
~. - .:~• ~ 3 spinous processes arranged along margins. teeth; B very large with 3-4 setae; Enpl and 2 with 2-
964 Copepoda
3 and 8-11 setae respectively; Exp 5-segmented with This family contains Alloiopodus Bradford, 1969;
I, 1, 1, 1, 1-2 setae respectively; other genera with Bathypontia Sars, 1905; Temorites Sars, 1900, and ~·
~!:1.2
reduced setation on Enp (Alloiopodus End with 0, 3 Zenkevitchiella Brodsky, 1955 (Grice and Hl.ilsemann bS
setae; Temorites discoveryae 0, 1 setae). Mx1 (based 1967; Fosshagen 1967; Bradford 1969; Schul z, 1996). ~
on Bathypontia) Li 1 with 9-10 long straight spines; Schulz (1996) considers that the assignment of ~I
Li2-4 reduced in size and setation with 1 seta each; Foxtonia Hiilsemann and Grice, 1963, to th is family is
Enp absent, Exp well-developed with I 0 setae; Le 2 not clear. It is possible that Bradyetes jlm·ens Grice
absent, Le I with 3-4 setae; other genera and species and Hiilsemann, 1967, may also be refened to this
with variously reduced setation (Temorites discover- family. Only the genus Bathypontia has been recorded
yae has the greatest reduction: Li I with 4 spines and from the South Atlantic. :'l::r.~
setae, Li2-4, Enp, and Le1 and 2 devoid of setae, Exp ~~

"'
~·
with 6 setae). Mx2 (based on Bathypontia) large, Lil- Key to Bathypontia species
5 with 0-1 , I, 2, 2-3, 3-4 setae respectively (one of Females and males
them enlarged), Enp with 6 enlarged, sometimes flat- 1 Mature female and male over 4 mm long; last
tened, distally serrate setae; other genera with more or segment of fe male P5 little longer than prece-
less modified setae and in some genera (e.g. ding segment; male P5: Bathypontia elongata
Alloiopodus) B elongated. Mxp (based on Sars, 1905 (see Fig. 7.413) t=·c~'
Bathypontia) with with relatively short setae; C Lil-3 1a Mature female less than 3 mm long; last segment z::c J
with 1-2, 1-2, 3 setae respectively from proximal to of female P5 3 times as long as preceding seg-
distal; B with 3 setae at about midlength; Enp 6-segment; male unknown: Bathypontia sp. Wheeler,
mented, Enp I almost completely fused to B with 1-2 1970 (Fig. 7.414)
seta, Enp2-5 with 4, 3-4, 2, 2, 2-4 setae respectively.
PI C with inner distal seta. Pl-4 with 3-segrnented
Exp; Enp of P I 1-2-segrnented, of P2-4 3-segrnented. Order CYCLOPOIDA
P 1 Exp segments I and 2 with or without outer edge
spines; Exp segment 3 of P2-4 with 3 outer edge Family Oitbonidae Dana, 1853 ~-
spines. Basis of P3 with a long thin gently curving Reference: Nishida ( 1985).
outer spine extending to the distal border of Exp seg- Females
ment 1. Seta and spine fmmula of Bathypontia as fol- Total length: 0.36 mm-1.90 mm. Pr short oval to long
lows: fusiform in dorsal view, of 5 somites. Anterior end of
head quadrate, rounded or produced into pointed R.
coxa basis exopod segments endopod segments Ur slender, of 5 somites; Gns longest. CR frequently
with 1 dorsal, 4 apical and 1 outer marginal setae; 2 of
Leg I 0- 1 0- 1 0- 1; 0- 1; ll,J,4 0-2-3/4
Leg 2 0-1 0-0 I- I; I-1 ; 111,1,5 0- 1; 0-2; 2,2,3
apical setae may be absent in Paroithona. A I slender,
Leg 3 0- 1 J-0 I-I ; I-I ; lll,J,5 0- 1; 0-2; 2,2,3 with proximal pa1t indistinctly segmented making it
Leg 4 0-0 1-0 1- 1; I- I; ITI,l,5 0- 1; 0-2; 2,2,3 difficult to determine precise number of segments. A2
uniramous, bent outward at medial geniculation;
P5 of female uniramous, 3-segmented, with two termi- proximal part !-segmented, with 1-3 inner and 0-2
nal spines, no other armature present (Bathypontia, outer marginal setae; distal part 1- or 2-segmented.
Temorites); or biramous with 2-3-segmented Exp and Proximal segment of distal patt short, with 2 medial
!-segmented Enp (Zenkevitchiella, Alloiopodus). and 3 distal inner marginal setae; distal segment long,
usually with 7 terminal setae. Md (Fig. 7.416) consis-
Males ting of gnathobase with 1-2 transverse rows of setules
A I 19-20-segmented, prehensile on right; terminal on anterior surface near distal margin. B transversally l..J::;a
part composed of 4 nan ow elongate segments, proxi- elongated, with 1 inner seta; fused to Enpl which has
mal segment characteristically curved. P2 may be 2-3 elements varying in shape and size; Enp2 with 2-
symmetrical or asymmetrically developed on each 5 setae; Exp 4-segmented, with 5 setae. Mx I (Fig.
side with Exp2 outer edge spine greatly elongate on 7.42 1) Li 1 with max imum of 11 inner setae; Li2 seta
one side. P 5 of male 3-5-segmented, Enp may or may present or absent; Li3 with 2-3 setae distally; Li4 with
not be present. 0-2 setae; Le 1 seta present or absent; Enp 1-segmen-
Copepoda 965
~ 1%9: ~ with 0-4 setae; Exp ! -segmented, with 3-4 setae. Oithonids of marine and estuarine waters are restric-
• and . h 2 with praecoxa and C fused, plane of fusion mar- ted to the sub-family Oithoninae Kiefer, 1928, wh ic h
5emann ed by incomplete transverse line, best observed on contains Oithona Baird, 1843 and Paroithona Farran,
~ 1996). :.=ner surface; Lil with 3 setae; Li2 with 1 spinulose 1908. Limnoithoninae Keifer, 1928 was created for
Dent of seta: Li3-Li4 each with 3 setae; B produced into inner the freshwater genus Limnoithona Burckhardt, 1912.
i!r.cily is .obe (LiS) bearing 2-3 setae or with 1 claw fused to Fine details of the morphology of Oithonidae as well
11 Grice -~- and 2 setae; Enp I with I inner marginal proxi- as keys have been produced by Nishida ( 198S) and
this ::'.3.1 seta and 3 inner marginal di stal setae; Enp2 w ith Wellershaus (1970). Rocha (198S, 1986) helped to
eoorded .:: strong, curved setae; Enp3 minute, with l strong elucidate the taxonomic details of common species in
:m-Yed seta and 2-3 more or less reduced setae. Mxp estuaries along the Brazilian coast both on systematic
- segmented; praecoxa and C incompletely separated; and biological grounds. In the South Atlantic Ocean,
~'"ta formula: 1-3-2 ( 1-2-2 in Limnoithona); B with I considering inlet and embayment environments, and
;:;.,_-"dial and 1 distal inner marginal setae; 1 midmar- neritic and oceanic provinces together, 22 species
c.g:lasr ::•al row and/or I distal posterior surface row of (Table I) have been reported.
1 p:-ece- c::ri::lute setae present or absent; Enp 1 with I or 3 setae;
-~aza E.;:p~ with 3-4 setae. P 1-4 usually with 3-segmented Readers interested in knowing more about the distri-
q and Exp except for Paroithona where P 1 Exp2 bution and taxonomy of Oithonidae on a broad basis
,egmenr ~ 3 and P1-4 Enp2 and 3 are partly or completely should refer to Giesbrecht (1892), Rosendorn ( 1917),
.2 seg- =::...,-'"d. In most species P4 Enp2-3 with 1-2 modified Wellershaus (I 970), Ferrari and Bowman ( 1980), and
-~ler. ~e which are thick and slightly curved towards tip, Nishida (198S).
widl distal flange. Seta and spine fonnula:
Key to Oithonidae species (based on Rosendorn,
coxa basis cxopod segments e ndopod segments 191 7; Ferrari and Bowman, 1980; Nishida, 1985;
Rocha, 1985, 1986)
..:; ! 0- 1 1-0/ 1 l-0/ 1; 0/1-1 ; lUlll,l,4 0- I ;0-1 ; I ,2,3
l-0; (UI,I,4/5)* 0-1 ; ( 1/2-2-3/4)*
Females
,_._-;: 2 0- 1 0 1-0 T-0/ I ; 0/ l-I ; 1/11/111,1,5 0-011; 0-1 /2; I ,2,3 I P 1 Exp and Enp 2-segmented; CR with 4 setae,
0-0; ( 1/2-2- 1/2/3)* longest two setae similar to apical spine ofP 1-P4
~;.: 0- 10 1-0 l-0/ 1; 0/T-1 ; 1/ H/11 1,1,5 0-0/ 1; 0- 1/2; I ,2,3 Exp; P1-P4 Exp outer spine formula: !-III, I-I-II,
iong 0-0; (1-2-1 / 2)*
I-0-II, l-0-I: Paroithona jlemingeri Ferrari and
~;! 0-10 1-0 0/T-0/ 1; 0/ l-1 ; O/ llll/lll,l,5 0-1 ; 0-2; 1,2,2
rend of Bowman, 1980 (Fig. 7.436)
0- 1; (1 -2-1/2)*
.erlR. Ia P1 Expand Enp 3-segmented; CR with 6 plu-
qoently • ""czr!)!;hona mose setae; Pl-P4 Exp outer spine formula
l!e: 2 of otherwise arranged ( Oithona) ............................ 2
sk:Dder. ~. :epresented by 2 lateral processes on Url ; its dor- 2 Forehead produced into pointed R curved down-
cin~ it ~ process small, with 1 terminal seta and ventral pro- wards (Fig. 7.4 1S) ............................................ .3
21G• •-\2 cess :1:>-ually a free segment with 1-4 tenninal setae. P6 2a Forehead not produced into pointed R (Fig.
:an: ~ed into lateral process with 1-2 setae on anterior 7.4 16, 7.42S) .................................................... l 4
t:rl 0-2 ..-..of genital segment. 3 P2 Exp2 with outer spine ................................. .4
~ed. 3a P2 Exp2 without outer spine ............................. 8
waiiaJ ·!.11es 4 P3-P4 Exp2 with outer spine/seta (Fig. 7.418)
.a:!g. -=:length: 0.37 mm-1.24 mm; usually smaller than ............ .............................. .......... ....................... S
I~ f--...,' e. Pr longer than Ur, of S somites. Forehead 4a P3-P4 Exp2 without outer spine (Fig . 7.4 15)
;seroles ~"ti in lateral view, tnmcate in dorsal view. ........................................................................... 7
~ posterior margin of Ce usually produced into 5 Forehead rounded in dorsal view; R wide, gra-
~· " irh well-developed rows of pores. Ur of 6 somi- dually tapering towards tip; Md B with 2 thick,
t:!S. ~.:-luding CR. A1 with 2 geniculations. A2, Md, blunt curved spines; Md Enp armed with 4 setae:
tl Ftg. l . :\h2 and Mxp similar to those offemale in basic Oithona brevicornis Giesbrecht, 189 1 (Fig.
Ll.Z~~ s:::x::=:cre however A2, Md, and P6 sexually dimorphic 7.418)
•irh ~ ~e species. P 1-4 Exp generally with I,I,II-UI Sa Forehead pointed in dorsal view (Fig. 7.430);
~ ~marginal spines. PS as in female. Md B with 2 pointed spines (Fig. 7.430) .......... 6
966 Copepoda
6 P3 Exp3 with 3 serrate spines on outer margin; 14 P4 Exp3 with 2 outer spines (Fig. 7 .416) ... .... 15
Md Enp with 5 setae; Mx1 with well developed 14a P4 Exp3 with different number of outer spines
Li4 armed with I seta: Oithona robusta (Fig. 7.417, 7.433) ... ..... .............. ........ ............. 21
Giesbrecht, 1891 (Fig. 7.430) 15 Free segment of P5 with 2 tenninal setae (Fig.
6a P3 Exp3 with 2 outer marginal spines, proximal- 7.429) ............................................. .... .......... .... 16
most reduced and smooth; Md Enp with 4 setae; 15a Free segment of P5 with only 1 terminal seta or
Mxl without Li4: Oithona vivida Farran, 1913 1 tenninal seta and I marginal spinule (Fig.
(Fig. 7.435) 7.426) .............. ............ ... .................................. 18
7 P2-P4 Exp 1 with short inner seta; P3 Exp3 with 16 Forehead broadly truncate in dorsal view (Fig.
2 rudimentary spinules on outer margin: Oitlzona 7.425); Md B with 2 spinulose spines (Fig.
pseudofrigida Rosendom, 1917 (Fig. 7.428) 7.425) ........ ...... ....... ...... .... ................................ 17
7a P2-P4 Expl without inner seta; P3 Exp3 without 16a Forehead triangular with tip blunt in dorsal ~ ..
such spinules on outer margin: Oithona .frigida view; Md B with 2 thick, blunt curved spines:
Giesbrecht, 1902 (Fig. 7.420) Oithona minuta Scott, 1894 (Fig. 7.423)
8 P4 Exp3 with minute outer seta/spine, always 17 Anterodorsal region of Ce with pair of ocular ~
shorter than Exp3 (Fig. 7.421) .... ............. ......... 9 lenses; CR 2.0-2.2 times longer than wide; Mx 1
8a P4 Exp3 with outer seta/spine longer than Exp3 Enp with 4 setae: Oithona oculata Farran, 1913
(Fig. 7.415) ......... .... ... .... .......... ........................ 11 (Fig. 7.425)
9 Mxl Li4 consisting of short protuberance with 17a Anterodorsal region of Ce without pair of ocular
seta; Mxl Enp with I seta; R claw-like, with lenses; CR 3.0-3.3 times longer than wide; Mxl -=
constriction at base in lateral view: Oithona Enp with 2 setae: Oithona rigida Giesbrecht,
hamata Rosendom, 1917 (Fig. 7.421) 1896 (Fig. 7 .429)
9a Mxl Li4 absent (Fig. 7.419); Mxl Enp without 18 Md B with 1 strong spinulose spine and 1 fine
seta (Fig. 7.419); R pointed, but without cons- seta on distal end (Fig. 7.424); Mxl Enp with 4
triction at base in lateral view (Fig. 7.419) ..... 10 setae (Fig. 7 .424) ................... ...... ....... ............. 19
10 Md Enp with minute seta and 3 setae subequal in 18a Md B with 2 thick, blunt curved spines with
length; Mx1 Li2 present and armed with 1 seta; short marginal spinules (Fig. 7.422); Mxl Enp
Mx1 Le1 represented by 1 seta: Oithona simi/is with 3 setae (Fig. 7.422) .................. ... ............. 20
Claus, 1866 (Fig. 7.432) 19 CR about 4 times longer than wide; Mxl Li I dis-
IOa Md Enp with 4 setae gradually increasing in tal marginal seta about twice longer than other
length from innermost to outermost; Mxl Li2 setae: Oithona attenuata Farran, 1908 (Fig.
and Le 1 absent: Oithona fa/lax Farran, 1913 7.416)
(Fig. 7.419) 19a CR 2.0-2.7 times longer than wide; Mx1 Li1
II Md Enp with 3 setae (Fig. 7.427); Mxl Enp with distal marginal seta similar to other setae:
without seta (Fig. 7.427) ............................... ..12 Oithona nana Giesbrecht, 1892 (Fig. 7.424)
11a Md Enp with 4 setae (Fig. 7.415); Mx1 Enp with 20 Forehead triangular with tip blunt in dorsal view,
1 seta (Fig. 7.415) .. ............... ....... ......... ...... ..... 13 and quadrate and dorsally humped in lateral
12 Md Enp with 3 setae similar in length; CR 3.5-4.0 view; CR 1.6-1.9 times longer than wide; Md
times longer than wide; R curved downwards: Enp with 4 setae: Oithona lzebes Giesbrecht,
Oithona plumifera Baird, 1843 (Fig. 7.427) 1891 (Fig. 7.422)
12a Md Enp with 2 long setae and 1 minute seta; CR 20a Forehead rounded to broadly pointed in dorsal
about 2.5 times longer than wide; R curved view, and from quadrate to rounded in lateral
downwards and backwards: Oithona tenuis view; CR about 3 tin1es longer than wide; Md
Rosendorn, 1917 (Fig. 7.434) Enp with 5 setae: Oithona oswaldocruzi
13 P4 Exp3 with outer spine slightly longer than Oliveira, 1945 (Fig. 7.426)
segment (1.2:1); Mx1 Li2 and Lei present, each 21 Pl -4 Exp outer spine formula: T-1-II, 1-T-TT, 0-1-I,
with 1 seta: Oithona setigera (Dana, 1849) (Fig. 0-0-I; middle apical setae ofCR short, thickened
7.431) and densely plumose; Md B with 2 thick, blunt
13a P4 Exp3 with outer spine twice as long as seg- curved spines; Md Enp with 4 setae; Mx l with-
ment; Mx1 Li2 absent; Mxl Lei represented by 1 out Enp, Lel and Li2: Oithona bjornbergae
seta: Oithona atlantica Farran, 1908 (Fig. 7.415) Ferrariand Bowman, 1980(Fig. 7.41 7)
Copepoda 967
----' -
,, ~L PI -t Exp outer spine fotmula: I-I-III, I-I-III, I-I- 12a P1-P4 Exp1 without itmer seta .... ......... ........... l5
sp:::::~es m. I-I-III; Md B with inner seta implanted near 13 Mx I Enp with 4 setae; Md B with 1 thick spinu-
-~ 1 :o 2 apical setae of this segment; Md Enp with 5 lose spine and 1 fine seta on distal end: Oithona
:e f::-ig. setae: \lt:x I Li4 absent: Oithona simplex FatTan, nana Giesbrecht, 1892 (Fig. 7.424)
__ l6 19I3 (fig. 7.433) 13a Mx1 Enp with 3 setae (Fig. 7.422) ................. 14
:;;...~ or 14 Mx1 Le1 represented by 1 seta; Md B with I
~Fig. thick, curved blunt spine and 1 pointed spinulo-
__ IS PI Exp3 with 2 outer spines ........ .. .................... 2 se spine on distal tip: Oithona brevicornis
r tg _ P I Exp3 with 3 outer spines .............................. 6 Giesbrecht, 1891 (Fig. 7.4 18)
-~ ~-
.-Jg.. P3 Exp3 with I outer spine: Oithona bjornber- 14a Mx 1 LeI absent; Md B with 1 spinulose spine
,- gae Ferrari and Bowman, 1980 (Fig. 7.417) and 1 weak spine anned with short spinules:
.:or-sal P3 Exp3 with a different number of outer spines Oithona hebes Giesbrecht, 1891 (Fig. 7.422)
~: ........... ............ .... ................... .... ............... .... ...... 3 15 Mxl Enp with 4 setae; Md B with 1 thick spinu-
:: P3 Exp3 with 2 outer spines ............................. .4 lose spine and 1 fine seta on distal end: Oithona
OL--ubr =:..! P3 Exp3 with 3 outer spines .............................. 5 attenuata Fa nan, 1908 (Fig. 7.416)
!: ~hi P2 Exp3 with 2 outer spines: Oithona simi/is 15a Mx I Enp with 1 seta (Fig. 7 .420); Md B with 2
:... l9B Claus, 1866 (Fig. 7.432) similar spinulose spines (Fig. 7.420) ....... ........ 16
-~ P2 Exp3 with 3 outer spines: Oithona fallax 16 Anal somite longer than CR: Oithona frigida
"OL-clar Farran, 1913 (Fig. 7.419) Giesgrecht, 1902 (Fig. 7.420)
~= ~b:l Penultimate segment of Al with semicircular 16a Anal somite shorter than CR: Oithona setigera
~tL. protrusion: Oithona plumifera Baird, 1843 (Fig. (Dana, 1849) (Fig. 7.431)
- A27)
I ::!ne 5.2 Penultimate segment of A l without semicircular
11--:± ~ protrusion: Oithona atlantica Farran, 1913 (Fig. Order HARPACTICOIDA
_ .. i9 - .-H5)
:s: \\ii:h P4 Exp3 with 3 outer spines: Oithona simplex Family Ectinosomatidae Sars, 1903
~~ Farran, 1913 (Fig. 7.433) Reference: Rose (1933).
-~0 P4 Exp3 with 2 outer spines .............................. 7 Body smooth, spindle-shaped; without distinct demar-
iicis- Free segment of P5 with 2 terminal setae (Fig. cation between Pr and Ur. R prominent, fused to Ce .
. c6er - .429) .... ....................................... ................ ...... 8 Pr prolonged laterally by well-developed epimeral
: !.g. Free segment of P5 with 1 tetminal seta or 1 ter- plates, except for last somite. Female Ur of 4 somites;
minal seta and 1 marginal spinule (Fig. 7.426) m ale Ur of 5 sornites; last somite small and more or
1:1 Lii .......... ......... ..................................... .. .... ........... 10 less cleft posteriorly. CR divergent; its 2 median setae
~~= .-\nterodorsal region of Ce with pair of ocular len- elongate. A I small. A2 with terminal spines denticu-
ses: Oithona oculata Farran, 1913 (Fig. 7.425) late; Exp 2-3 -segmented. Rami of P1-4 3-segmented.
,'le'\\•. &. .-\nterodorsal region of Ce without pair of ocular P5 lamellar with strong spines. Ovisac simple.
~'"nl lenses .. ...... ............ .................. ............ ......... ...... 9
e.: Me ~ fd B with 2 spinulose spines on distal end: Two species (Microsetella norvegica and M. rosea),
;;:r;!'.::iu.. Oithona rigida Giesbrecht, 1896 (Fig. 7.429) of the only pelagic genus in the family, are known
:\ld B with 2 thick, blunt spines on distal end : from the South Atlantic.
-.h-~ Oithona minuta Scott, 1894 (Fig. 7.423)
~ ..... ::!.! :\fd Enp with 5 setae (Fig. 7 .426) ................... 11 Family Miraciidae Dana, 1846
e: Me ~ :\Id Enp with 4 setae (Fig. 7.422) .............. ..... 12 Reference: Wells (1970).
·~ :\fd B with 2 similar spinulose spines on distal Ce and Pdl fused. CR about as long as last 2 Ur somi-
end: Oithona robusta Giesbrecht, 1891 (Fig. tes. P1 Exp 3-segmented, Enp 2-segmented. P2-4 with
_0--I-L - .430) both rami 3-segmented. P2 Enp of male modified, 2-
~ II:! :\id B with 1 spinulose spine and 1 smooth spine: segmented. P5 basal segment expanded medially. Ce
. -..'1...--
_._._.__... Oithona oswaldocruzi Oliveira, 1945 (Fig. usually with pair of large cuticular lenses .
wct.- -.426)
~ P I -P4 Exp l with short inner seta .................... 13
968 Copepoda
There are four genera in the family: Macrosetella A. of P wide; P 1-4 rami 3-segmented. P5 rudimentary,
Scott, 1909; Distioculus Huys and Bottger-Schnack, elongate, !-segmented. P6 relatively large, !-segmen-
1994; Oculosetella Dahl, 1895; and Miracia Dana, ted.
1846 (Huys et a!., 1996). Four pelagic species
(Macrosetella gracilis, Miracia efferata, M. minor, Both species (Aegis thus aculeatus and A. mucronatus)
and Oculosetella gracilis) in this family are known in the only genus (Aegisthus Giesbrecht, 1891) in the
from the South Atlantic. family are known from the South Atlantic.
Family Euterpinidae Brian, 1921 Key to pelagic Harpacticoida species (from Wells
Reference: Rose ( 1933). 1970)
Body variable in shape. Al shOLt, prehensile in male. 1 Pdl free; there are in total 9-10 (Ur 2-3 fused in
A2 Exp well-developed. Md with biramous palp. P J A. mucronatus) free body somites in female and
not prehensile, similar to P2; all with 3-segmented I 0 in male (family Aegistbidae) ...................... .2
Exp and Enp. P5 lamelliform, 2-segmented. Ovisac I a Pd l fused with Ce; there are in total 8 free body
simple. somites in female and 9 in male .......... ..... ......... 3
2 Female A l 6-segmented. Pr without network of
One species (Euterpina acutifrons), of the on ly pela- ripples in chitin: Aegistlms mucronatus
gic genus (Euterpina Norman, 1903) in the family, is Giesbrecht, 1891 (Fig. 7.447)
known from the South Atlantic. 2a Female Al 7-segmented. Pr with network of rip-
ples in chitin: Aegistlms aculeatus Giesbrecht,
Family Clytemnestridae A. Scott, 1909 1891 (Fig. 7.446) ~
Reference: Rose ( 1933). 3 Both rami of P1 2-segmented (family
Prof 4 so mites, its posterolateral comers flattened and Euterpinidae): Euterpina acutifrons (Dana,
prolonged into points. Ur of 5 somites in female and 1847) (Fig. 7.443)
of 6 somites in male. CR short, setae very short. A I 7- 3a At least one ramus of P 1 3-segm ented ............. .4
8-segmented, with short setae and long sensory 4 Exp of PI !-segmented, P5 elongate, not flatte-
organs; male AI prehensile on both sides. A2 3-seg- ned (family Clytemnestridae: Clytemnestra)
mented; Exp rudimentary, represented by 1-2 setae. (Fig. 7.444) .................... ................... .......... ....... 5
Mxp 2-segmented, elongate, with terminal claw; male 4a Exp of P 1 3-segmented, P5 of typical harpacti-
differs from female. P J Enp 1-segmented, Exp 3-seg- coid form with wide baseoendopod (Fig. 7.439)
mented. P2-4 with both rami 3-segmented. P5 rudi- ......... ..................................................................6
mentary, with 2 long segments; symmetrical or almost 5 Female AI 8-segmented, Exp ofP I with 4 setae:
in male. Clytemnestra scutellata Dana, 1847 (Fig. 7.445)
5a Female AI 7-segmented, Exp of Pl with 3 setae:
Two species (Clytemnestra rostrata and C. scutellata) Clytemnestra rostrata (Brady, 1883) (Fig. 7 .444)
in the only genus (Clytemnestra Dana, 1848) in the 6 Enp ofPl 2-segmented (Family Miracidae) ..... 7
family are known from the South Atlantic. 6a Enp ofPl 3-segmented (Microsetella) ........... 10
7 Cuticular lenses absent, body narrowly spindle-
Family Aegisthidae Giesbrecht, 1892 shaped: Macrosetella gracilis (Dana, 1847)
References: Rose ( 1933), Wells ( 1970). (Fig. 7.439)
Female 7a Cuticular lenses present, body more or less
Pr wide and Ur narrow. Pr of 5 somites, Ur of 5-6 cylindrical .......................................................... 8
somites; Url and 2 either separated only on dorsal 8 A2 Exp absent: Oculosetella gracilis (Dana,
surface or completely around. Ce prolonged into 1852) (Fig. 7.442)
point. CR short, fused to last Ur somite and fused in 8a A2 Exp !-segmented, with 2 setae (Miracia) .. .9
median line; carrying 2large setae. A I 6-7-segmented 9 PS basal segment with 5 setae in female, 3 setae
with long sensory organs and segment 3. Md reduced in male: Miracia efferata Dana, 1849 (Fig. 7.440)
to I toothed blade. Mxl without Enp. Mx2 elongate; 9a P5 basal segment with 4 setae in female, 2 setae
Li5 with 1 thickened pickaxe-like seta. Mxp small, 2- in male: Miracia minor T. Scott, 1893 (Fig.
segmented. Mx2 and Mxp rudimentary in male. Basis 7.441)
Copepoda 969
~ IC Setae of CR not longer than body: Microsetel/a out into process armed with 3 elements. Mxp 4-seg-
e;wen- norvegica (Boeck, 1864) (Fig. 7.437) mented, C and B with or without setae; Enpl small,
lOa Setae of CR at least twice as long as body: Enp2 in the form of claw and may bear proximal setae
J1icrosetella rosea (Dana, 1847) (Fig. 7.438) (Lubbockia and Pseudolubbockia ). P 1-4 biramous
=a;us) with both rami 3-segmented; inner coxal and outer
i:l ilie basal seta present; inner seta sometimes present on B
Order POECILOSTOMATOIDA of PI (Lubbockia and Pseudolubbockia). Spine and
seta fmmula:
• v·elli • -~ to families of Poecilostomatoida
:\2 uniramous, 3-segmented, C and B fused, Enp coxa basis cxopod segments endopod segments
iCsed in 2-segmented, subprehensile to prehensile, termi-
Leg l 0-l /0 1-1 1-0; 1-1; 11 1/11,1,4 0- 1; 0-1 ; 0/1/ I ,U2,3/4/5
cle and nal part with 6-7 setae or spines: Oncaeidae Leg2 0-1 /0 1-0 1-0; 1-1; 111/11,1,5 0-1 ; 0-2; 0/U11,11/1,3
-····-2 (Fig. 6.8C, 6.8D) (p. 969) Leg 3 0-1/0 1-0 1-0; 1-1 ; lllll lll,J,S 0-1 ; 0-2; 0/1/ll,ll/1,3/2/1
eebody h .-\2 uniramo us, 5-segmented, C and B separate, Leg 4 0-110 1-0 1-0 ; OII-I; n rr,1,s 0-1 ; 0-2; 0/llll,ll/1 ,2/1 / 1
...
-·····-' Enp differently arranged .................................... 2
r~of .-\2 uniramous, 5-segment, prehensile, C and B P5 absent or uniramous, with single segmented proto-
-.arus large and wide each w ith thick long seta, each pod (C+B); inner coxal seta absent; outer seta on B
Enp segment with a few strong, thick spines, may be present; Exp !-segmented bearing 1-4 ·setae;
•ofrip- Enpl short (Fig. 6.8£). Body of cylindrical Enp absent. P5 may be represented by 1-2 setae alone
aecht. shape; Corycaeidae (Fig. 6.8F) (p. 973) (Conaea , Epicalymma). P6 represented by opercular
:\2 uniramous, 5-segmented, elongate, prehensi- plates bearing 1-2 setae or spines closing off genital
(family le. C and B separate not particularly thickened, openings.
fr>...na.. Enpl well-developed bearing several moderate
spines, terminal segment claw-like (Fig. 6.9A, Males
_____ _4
6.9B, 6.9F). Body dorsoventrally flattened in Ur of 6 somites, genital apertures paired, located dor-
l ;'i..orrre- male at least: Sapphirinidae (Fig. 6.9C, 6.9D, solaterally at posterior border of large Gns. A I never
JSe.r.Tal 6.9G) (p. 971) genicu late, last 3 segments often fused; 3-6 segmen-
---·-5 ted. A2 and mouthparts rarely reduced. Mxp B usual-
~- ~:!::llily Oncaeidae Giesbrecht, 1892 ly more robust and muscular in male than female, 5-
- !!39) ...=:erence: Heron and Bradford-Grieve ( 1995). segmented in Pseudolubbockia. P6 represented by
____ 6 F~ales paired flaps at posterior of Gns.
- 5-.."'tae: r.- dongate to elongate-oval, or cyclopiform. Ce and
- _!5 ~.:! always separate. Ur generally slender, of 5 somi- This family contains five genera Conaea Giesbrecht,
:. X'Oae: 1:5.. genital apc1tures paired, located dorsally or late- 189 1; Epicalymma Heron, 1977; Lubbockia Cla us,
":": ,. ,# .
~- on anterior pa1t of Gns; seminal receptacles pai- 1863; Oncaea Phil ippi, I 843; and Pseudolubbockia
r. __ r:C.. CR with 6 setae, dorsal seta may or may not be Sars, 1909; of which all but Pseudolubbockia have
-· _10 ~ed into dorsal expansion on ramus (Conaea and been taken in the South Atlantic. Membe rs of this
- .-r1.~- ~.:J(.-nzma) . Eggs carried in paired sacs. R fused to family in the South Atlantic are badly in need of revi-
~~/) c~: aauplius eye present; labrum well-developed, sion, especially Oncaea; many previous records can-
~y incised. A l short 6-7-segmented. A2 unira- not be confirmed against current knowledge of spe-
.Less ._s_ 3-segmented, subprehensile to p re hensi le; with cies because inadequate illustrations are provided .
=~""gllented Enp, terminal part with 6-7 setae or spi- The following keys are tentative guides only. If a spe-
D:ma.. r-es-=-~ lacking; C and B fused. Md complex with 3- cies cannot be identified fi·om these keys, then a wider
5 5ri:>tenninal elements; gnathobase sometimes with suite of species w ill have to be examined (see e.g.
Cia' a of scale-like denticles on median or distal edge; Heron and Bradford-Grieve, 1995).
-=~ ~; absent. Mx I small (most reduced in
Plc-Jt.iolubbockia), bilobed; Lil w ith 2-3 setae; outer Key to genera of Oncaeidae and one species of
~ 5-..""'Z.:!e :cpresenting reduced palp, armed with 2-4 setae. Epicalymma
-· ~:E !:L .::-segmented; proximal segment representing C, The key includes species not yet taken in the South
~ ~ large; distal segment representing B, drawn Atlantic (Heron and Damkaer, 1978). Pachos tubero-
970 Copepoda
sum (Giesbrecht. 189 1) has also been included here Males

although it is not now considered to be an Oncaeid I Mxp terminal claw strongly curved, inner edge
(Heron and Damkaer 1978). with sharp step just proximal to bordering spinu-
Females and males les and adjacent to distal end of claw: Lubbockia
1 P 1-4 with inner coxal seta .............. ................... 2 aculeata (Fig. 7.451)
I a P 1-4 without inner coxal seta .. ......................... .4 la Mxp terminal claw slightly curved, inner edge
2 P5 fused w ith U rl , with 2 setae and 2 basal setae: without sharp step proximal to bordering small
Pachos tuberosum Giesbrecht, 189 1 (Fig. 7.465) spinules: Lubbockia squillimana (Fig. 7.452)
2a P5 either absent or a free segment .... ................ 3
3 Md with narrow serrate tip; P5 a free segment Key to species of Oncaea
armed with 4 elements: Pseudolubbockia Females
3a Md terminating in lash or lashed element; P5 1 In lateral view, Pr with conspicuous dorsoposte-
absent ( 1 species) or an elongate free segment rior projection ............ .............. .......................... 2 A:
aimed with 2 elements: Lubbockia (p. 970) 1a In lateral view, Pr without dorsoposterior projec-
4 CR with conspicuous expansion on dorsal sur- tion .................. .......... .... .............. .. ..... ................ 3
face, surrounding insertion of dorsal seta ......... 5 2 Gns about 1.8 times as long as wide; P5 globu- 2
4a CR without expansion on dorsal surface: Oncaea lar-shaped: Oncaea antarctica Heron, 1977
(p. 970) (Fig. 7 .453)
5 P 1-3 Exp3 with II, III, III outer spines: 2a Gns about 1.3 times as long as wide; P5 parallel-
Epicalymma exigua (Farran, 1908) (Fig. 7.450) sided: Oncaea clevei Fruchtl, 1923 (Fig. 7.455)
5a P1-3 Exp3 with III, II, II outer spines: Conaea 3 P4 Enp3 without inner edge spines; "Oncaea"
(p. 970) atlantica Schmeleva, 1966 (Fig. 7.454)
3a P4 Enp3 with at least 1 inner edge spine ......... .4
Key to species of Conaea 4 P4 Enp3 with l outer edge spine: Oncaea eng-
Females lishi Heron, 1977 (Fig. 7 .457)
1 Gns in dorsal view indented laterally just anteri- 4a P4 Enp3 with 2 outer edge spines ..................... 5
or to midlength; Mxp B distal inner setae much 5 CR as long as combined length of 2 preceding ~
larger than proximal seta: Conaea rapax somites ....................................................... ........ 6

Giesbrecht, 1891 (Fig. 7 .449) 5a CR shorter than combined length of 2 preceding
1a Gns in dorsal view with straight lateral borders; somites ............................................................... 8
Mxp B distal inner setae only slightly larger than 6 Gns length about double that of Ur3-5 plus CR:
prox imal seta: Conaea expressa (Gordejeva, Oncaea media Giesbrecht, 1891 (Fig. 7.460)
1973) (Fig. 7.448) 6a Gns length less than double that of Ur2-5 plus
CR ........................................ ...... ... .............. ....... 7
Males 7 P5 terminal setae about equal in length: Oncaea
I Anal somite long, longer than previous 3 somi- scottodicarloi Heron and Bradford-Grieve, 1995
tes combined: Comzea expressa (Gordej eva, (Fig. 7.464)
1973) (Fig. 7.448) 7a P5 terminal setae unequal in length: Oncaea
la Anal somite short, of approximately same length mediterranea (Claus, 1963) (Fig. 7.46 1)
as each of previous 3 somites combined: Conaea 8 P5 terminal setae with 1 of them half length of
rapax Giesbrecht, 1891 (Fig. 7 .449) other ...... ........... ............. ..... ............. ................... 9
8a P5 tenninal setae with both setae equal or sube-
Key to species of Lubbockia qual in 1ength ............ ....................................... IO
Females 9 P5 elongate, delimited fro m Uri: Oncaea prola-
I In dorsal view terminal spines ofP5 do not extend ta Heron, 1977 (Fig. 7.463)
as far as posterior border of Gns: Lubbockia acu- 9a P5 short, not delimited fro m Ur i: Oncaea /acinia
leata G iesbrecht, 1891 (Fig. 7.45 1) Heron, English and Damk.aer 1984 (Fig. 7.458)
1a In dorsal view one terminal spine of P5 extends 10 P5 elongate, at least 4 times as long as wide:
to posterior border of Gns: Lubbockia squilli- Oncaea pari/a Heron, 1977 (Fig. 7 .462)
mana Claus, 1863 (Fig. 7 .452) I Oa P5 short, at most 1.5 times as long as wide .... 11
Copepoda 971
l~ In dorsal view, anterior part of Ce is broadly 11 a Mxp segment 2 with several anterior rows of
er OO.ge rounded, widest part anterior to posterior border setules at midlength: Oncaea scottodicarloi
~~!IlU of Ce: Oncaea latimana Gordej eva, 1975 (Fig. (Fig. 7.464)
kia 7.459)
ll.a In dorsal view, anterior part of Ce na tTowed, Family Sapphirinidae Thorell, 1859
~~e widest part at posterior border of Ce: Oncaea Females
E sm.ill curvata Giesbrecht, 1902 (Fig. 7.456) Body more or less flattened, often leaf- like. Two ocu-
-5:, lar lenses present on anterior Ce. Ce and Pd I fused
"lales (Copilia) or separate; Pd3 and 4 may be expanded
1 P4 Enp3 w ith conical projectio n between termi- laterally into rounded lateral plates. Ur may be slender
nal and subterminal spines: Oncaea antarctica or tapering gradually from width of posterior Pr, of 3-
~-re- (Fig. 7 .453) 5 somites (in Sapphirina fused Ur2 and 3 have
..,
a P4 Enp3 without conical projection .......... ........ 2 obvious demarcation between sornites); Url (Pd5)
r .... -Jec- P5 with 1 seta; P4 Enp3 without outer edge spi- small or may be fused to Ur2; genital apertures paired,
-·-··--3 nes: "Oncaea" atlantica (Fig. 7 .454) located laterally or dorsolaterally on anterior part of
g:obu- - With an other combination of characters ...... .... 3 Gns; CR leaf-shaped or in fonn of narrow, elongate
l : -- - P5 with 1 term inal seta and I basal seta; P4 Enp3 rod, with 4-5 setae. Eggs caJTied in paired sacs. AI
''ith I outer edge spine: On caea englishi (Fig. short with at most 6 segments. A2 uni ramous, 5-seg-
~_:re! - - A 57) mented, prehensile, terminated by claw. Md w ith I
!~5 5 P5 with 2 terminal setae; P4 Enp3 with 2 outer large terminal toothed element usually with 1-2 small
lillauoa" edge spines ............................. ............................4 setae at its base. Mxl small with ! lobe, with 4 spines.
P5 separated from Ur L.. .. ............ ...... ................. 5 Mx2 claw-like, ! -segmented; drawn out into process
_..! P5 not separated from Ur i .. .. .......... ........ .......... 7 armed with 3 major elements: I long spinous seta and
!'1:1' t!7fa- ~lxp tenninal claw extends beyond proximal 2 variously spinulose additional setae. Mxp claw-like
border of segment 2: On caea prolata (Fig. 3-segmented, B with 2 inner edge spines; Enp 1 fused
:; -..f63) with Enp2 and i.n form of claw which bears 2 proxi-
~g ~ lxp termina l claw does not extend as far as mal setae. P 1-3 biramous with both rami 3-segmen-
- . 6 proximal border of segment 2 ........................... 6 ted; P4 with 3-segmented Exp and Enp !-segmented
~g CR about same length as anal somite: Oncaea (Copilia 0, 1, 1), 2-segment ( Vettoria 0-1 ; 0,1, 1) and 3-
/arimana (Fig. 7.459) segmented (Sapphirina). Spine and seta formula typ i-
-CR.: CR shorter than anal somite: Oncaea pari/a cally:
- k--~
Fig. 7.462)
,-5 p:..JS CR longer than anal somite .. ... ........................ .. 8 coxa basis exopod segments endopod segments
CR about same length as ana l somite ... .. ........ 10
Leg I 0-1 1-0 1-0; I-I ; ll/lll,L4 0- 1; 0-1 ; 1,2,3
~-.reG In dorsal view, Gns widest anteriorly; P5 with- Leg 2 0-1 1-0 1-0; 1-1 ; ll/111.1,5 0-1; 0-2; 11,1,3
~ l""S out basal seta: Oncaea c/evei (Fig. 7.455) Leg 3 0-1 1-0 I-0; 1- 1; TJ/I ll.l,5 0-1; 0-2; 1/11,1,2/3
in dorsal view, Gns widest at midlength; P5 with Leg 4 0-1 1-0 1-0; 1- 1;11/111,1,5 0-1 ; 0-2; !10,1,0
~ ~ seta ........................... ......... ......... ....... ...... .. 9
_·\nteroventral profile of Gns in lateral view, P5 !-segmented with 2 small term inal setae and small
~of gently curving: On caea m edia (Fig. 7.460) basal seta on each side, or 1 spine and seta, or respre-
_-\nreroventral profi le of Gns in latera l view, sented by 3 setae alone (Vettoria). P6 represented by
~!::.:le- :mgular: Oncaea mediterranea (Fig. 7.461) 3 small rounded appendages (Sapphirina), or single
-. _JO : hp inner border w ith small distal setules and seta (Vettoria);
I j:TO/:IJ- ::OOg setules at midlength: On caea lacinia (Fig.
- A 58) Males
_.1xp inner border with setules of similar size Body shape usually different from that of female, usu-
·--...................................................................... ! I ally more leaf-shaped with no distinct break in width
' W"!de: • :_-.;:p segment 2 with 2 anterior, proximal, semi- between the Pr and Ur. Ur of 6 so mites, Ur l and ana l
!::.-cular rows of setules: Oncaea curvata (Fig. somite often covered by adjacent Ur somites; genital
fe_)f - ..!56 apertures paired, located ventrolaterally near posterior
972 Copepoda
border of the Gns; CR leaf- or rod-shaped. A I never Males

geniculate. A2 more slender than in female. Md may I Mxp B without obvious thicken ing: Copilia
be reduced. Mxp final hook is longer in male than quadrata (Fig. 7.470)
female, 4-segmented. P2 Enp3 dista l part with modi- 1a Mxp B with thickening obvious ........................ 2
fied spines in Sapphirina. P6 represented by 3 setules 2 Mxp B with thicken ing proximal to midlength:
in Sapphirina. Copilia vitrea (Fig. 7.471)
2a Mxp B thickening more distally placed ............ 3
This family contains three genera Sapphirina J. V. 3 Mxp B thickening in form of rounded dista l
Thompson, 1829; Copilia Dana, 1849; and Vettoria expansion: Copilia mirabilis (Fig. 7 .469)
Wilson, 1924; of which only the first two have been 3a Mxp B thickening set back from dista l border of
taken in the Atlantic Ocean and have been reviewed segment .. .............. ...... ................ .................. ... ...4
by Lehnhofer ( 1926, 1929). 4 Mxp B with prominent rounded lobe just di sta l
to midlength: Copilia hendorffi (Fig. 7.466)
Key to genera of Sapphirinidae 4a Mxp B not prominent ........................................ 5
1 Body flattened, leaf-like or like thin scale; often 5 A2 Enp2 more than twice length of terminal seg-
iridescent when alive. Ur somites enlarged late- ment; Mxp B squat, length/width = 3: Copilia
rally; head with or without ocular lenses .......... 2 lata (Fig. 7.467)
1a Ce square; Ur narrow; frontal ocular lenses 5a A2 Enp2 less than twice length of terminal seg-
remarkable and set far apart: female Copilia (p. ment; Mxp B elongate, length/width= 4: Copilia
972) mediterranea (Fig. 7.468)
2 Head with ocular lenses; CR in form of 2 broad D
plates: male and female Sapphirina (p. 972) Key to the species of Sltpplririna (female and male)
2a Head w ithout ocular lenses; CR in form of 2 (from Lehn hofer 1929; Rose 1933)
parallel batons: male Copilia (p. 972) CR with 2 small points on inner distal border:
Sapphirina bicuspidata Giesbrecht, 1891 (Fig.
Key to the species of Copilia (from Lehnbofer, 1926; 7.474)
Rose, 1933) la CR with only I or no projection on inner distal
Fem ales border ....... .............. ....... ... ............. ..................... 2
1 Ocular lenses separated by distance of less than 2 CR without inner dista l projection: male
their own length; P5 with 1 spine besides 2 setae: Sappltirina iris Dana, 1849 (Fig. 7.477)
Copilia vitrea (Haeckel, 1864) (Fig. 7.471) 2a CR with I inner distal projection ...................... 3
1a Ocular lenses separate by distance of at least 1 3 CR with 2 terminal setae lance-shaped:
l/2 their length; P5 without spine ..................... 2 Sappltirina metal/ina Dana, 1849 (Fig. 7.478)
2 A2 C with numerous small spines; large spine on 3a CR with 2 terminal setae normal... ................... .4
B with 4-6 lateral spines: Copilia mirabilis 4 CR with inner distal projection large and exten-
Dana, 1849 (Fig. 7.469) ding posterior to level of insertion of apical setae
2a A2 without such ornamentation ................. .. .....3 ............ .................. .. ......... .. ... ............................. 5
3 A2 Enp 1 about equal in length to B .. ................4 4a CR with inner distal projection small and usual-
3a A2 Enp I much shorter than B .......... ..... ... .. .. ..... 5 ly on inner border ........................ .. .................... 6
4 A2 Enp2 slightly longer than B: Copilia quad- 5 A1 5-segmented; CR alm ost twice as long as
rata Dana, 1849 (Fig. 7.470) wide: S(tpphirina angusta Dana, 1849 (Fig .
4a A2 Enp2 about twice as long as B: Copilia lata 7.472)
Giesbrecht, 1891 (Fig. 7.467) 5a AI 3-segmented; CR nearly as long as wide:
5 A2 Enp I terminal spine about as long as its seg- Sappltirina opalina-darwinii Dana, 1849 (Fig.
ment: Copilia melliterranea (Claus, 1863) (Fig. 7.480)
7.468) 6 A I segments 3 and 4 with short spines instead of
5a A2 Enp 1 term inal spine much shorter than its setae: Sapphirina stellata Giesbrecht, 1891 (Fig.
segment: Copilia ltendorffi (Dahl, 1892) (Fig. 7.483)
7.466) 6a A I segments 3 and 4 with norma l setae ........... 7
Copepoda 973
I P4 Enp longer than Exp: Sapphirina ovatolan- ly abse nt; PI C with or without inn er seta. Spine and
f"llpilUI ceolata-gemma Dana, 1849 (Fig. 7.48 1) seta fo rmula:
:Ia P4 Enp shorter than Exp ........ ............. ...... .........8
P4 Enp nearly as wide as Exp ............ ...... .........9 coxa basis exopod segments endopod segments
~T.h: Sa P4 Enp much narrower than Exp ........... ......... II

Leg I 0-0/ 1 1-0 1-0; 1-1 ; l/Ill,l,4 0- 1; 0-1 ; 0,2,3
9 C R inner border straight: Sapplririna gastrica Leg 2 0- 1 110?-0 1-0; I- 1; IJJ,I,5 0- 1; 0-2; 0,1,3
Giesbrecht, 1891 (Fig. 7.475) Leg 3 0-0 1-0 1-0; 1-1 ; lll,1,5 0- 1; 0-2; 0,1,1
I tfuol 9a CR with convex inner border .......................... I 0 Leg 4 0-0 1-0 l-0; 0- 1; I,l,S 0, 1,0/ 1
lO A2 Enp2 hardly longer than Enp I: fema le
~of
. Sapphirina iris Dana, 1849 (Fig. 7.477)
iOa A2 Enp2 much longer than Enpl : Sapphirina
P5 in form of 2 small setae on each side. P6 represen-
ted by opercular p la tes closing off genital openings .
'6-ral auronitens-sinuicauda Claus, 1863 (Fig. 7.473)
! 1 P4 Enp3 with L terminal seta: Sapphirina in- M ales
.:; testinata Giesbrecht, 1891 (Fig. 7.4 76) Ur of 2-3 somites, genital apertures paired, located
5-..~- : 1a P4 Enp3 w ith 2 terminal setae ............ .......... ... 12 ventrolaterally near posterior border of Gns. A 1 never
Lopilill :2 Female 1.55-2.8 1 mm, male 1.7 1-3.04 mm. gen iculate. A2 terminal hook longer in male than
Proportion of A2 segments 3-5 to segments 1-2 female. Mxp ftnal hook is longer in male than female,
s-.~- in male I : 1.00-1.46, in female I: 1.00- 1.31: 4-segmented; B w ith or without inner edge spine. P6
f"t;tpi/UJ S applzirina nigromacu/ata Claus, 1863 (Fig . represented by I plumose seta on flaps closing off
7.479) gen ita l apett ures.
i2a Female 3.33-4.62 mm, male 3.97-4.85 mm .
f~e) Proportion of A2 segments 3-5 to segments l -2 This family contains two genera C01ycaeus Dana,
in male 1:1. 72- 1.95, in fe male I : 1.60- 1.90: 1849, and Farranula Wilson, 1932, both of which
..:e. S apphirina scarlata G iesbrec ht, 1891 (Fig. have been taken in the South Atlantic. T he family was
II Fi; 7.482) reviewed by Dahl (19 12). The re has been no really
critical exam ination of the identity of members of this
r&'UJ ~:mlily Corycaeidae Dana, 1852 family in the South Atlantic since this review, so
Fema les so me distri bution records may be suspect.
rr...,Je 3ody cylindrical to conical, tapering poste riorly.
8:Lll1ar lenses close together, sometimes contiguous. Key to gene r a of Corycaeidae a nd subgenera of
c~ and Pd1 fused or separate; Pd3 and 4 prolonged Corycaeus (females and males) (after Rose 1933 and
~: ~o lateral points; Pd2, 3 and 4 may be fused or sepa- Dahl1 9 12)
.- ,,
3e. Ur generall y slender, of 2-3 somites, U ri (Pd5 ) A2 with 2 proximal, plumose setae; Ur of I
~ : genital ape1t ures paired, located dorsolaterally somite in both sexes. Female Ce with beak-sha-
~~ anterior part of Gns; seminal receptacles paired; ped, posteriorly directed process on ventra l keel
-~~
.x:- CR elongate with 4 setae. Eggs carried in paired sacs. (Fig. 7.502). Exp of P l -3 with 0, 0, I outer spi-
.:;
A: short, 6-segmented w ith naked setae; without sen- nes. P4 w ithout Enp in both sexes and B wi th 1
IS:..!.- ,~- organs. A2 uni ramous, 5-segmented, prehensile; nude seta: Farranula (p. 975)
- 6 C :md B large, each with thick, long seta; Enp I ve ry I a A2 with 2 proximal nude setae in female; in
..
-=-
, ~n: Enp with strong, hooked spines. Md with 2 ter- male 1 seta ftnely plumose along one edge. Ur
.... . ;-""""lal elements in form of spino us teeth. Mxl small generally of 2 somites (except females of s.g.
-ili I lobe; with 4 sp ines. M x2 !-segmented; drawn Agetus). Ventral projection on Ce rounded in
:r~- mto process armed with 3 major elements: 1 long both sexes. Exp P 1-3 w ith I, I, III, spines. P4
F:g. ;!:::mose seta, 2 variously spinulose spines. Mxp 3- (besides basa l seta) with small, peg-like Enp
:o-~ented, B wi th l inner edge spine; E np1 fused w ith l-2 setae: Corycaeus ................ .... ............. 2
~o:" '""i.ll Enp2 and in the form of claw which bears 2 2 CR of male longer tha n remainder of Ur; of
II (Fi; p;n"UIDal setae. P 1-3 biramous with both rami 3-seg- female nearly as long and not divergent: s.g.
~ted. Exp longer than Enp; P4 with 3-segme nted Urocorycaeus (p. 975)
Exp: Enp !-segmented with 1 or 2 se ta, or complete-
974 Copepoda
2a CR of male shorter than remainder of Ur; of external border. Male anal somite, viewed dor-
female very rarely slightly longer (speciosus) sally, slightly longer than wide, and together
and strong ly divergent.. .. ................... ....... ......... 3 with CR about equal to Gns; in lateral view pos-
3 Peg-like Enp ofP4 with 2 setae and small poste- terior borders Gns distinctly arched dorsall y and
rior projection; coastal forms: s.g. Ditrichocory- ventrally: Corycaeus (Agetus) typicus (Kl·oyer,
caeus (p. 974) 1849) (Fig. 7.486)
3a Peg-like Enp of P4 with J seta, without posteri- 2a Ur of female strong ly expanded and in lateral
o r projection .. ............... .................................. ... .4 view wi th peg-like dorsal projection; corners of
4 Small forms at most 1.18 rrun. Females with Pd4 not projecting dorsally; A2 B not toothed on
very wide Ce; Pdl is generally 2/3 as wide as external border. Male anal somite, viewed dor-
long. Male A2 distal hook longer than C and B sally, 1 1/3 times as long as wide; and together
together: s.g. Ollyclwcorycaeus (p. 975) wi th CR, longer than Gns; posterior border of
4a Large forms (except Agetus limbatus). Female Gns not projecting in lateral view: Corycaeus
Ce narrow; Pd 1 not 2/3 as wide as long. Male A2 (Agetus) jlaccus Giesbrecht, 189 1 (Fig. 7.484)
distal hook does not pass proxi mal border of B
.......................... ...................................... ........... 5 Key to species of Corycaeus subgenus Corycaeus
5 In both sexes 2 terminal hooks of A2 almost (from Dahll912)
equal; 2 basal setae almost equal. Female Gns, in Females and males
lateral view, overhangs anal somite. CR short in 1 CR approximately as long as 2 preceeding Ur
both sexes and sometimes a little divergent: somites together, sometimes longer in male; 2:2
Corycaeus (Monocorycaeus) robustus (Gies- female A2 B with 2 pointed teeth on inner distal
brecht, 1891) (Fig. 7 .492) border. Male anal somite in dorsal view more
5a In male, large terminal hook of A2 at least twice than 2 times as long as wide: Corycaeus
length of smaller hook. In female, Gns does not (Cmycaeus) speciosus Dana, 1849 (Fig. 7.489)
overhang anal somite .... ............................ ........ .6 1a CR at most 2/3 as long as 2 preceeding Ur somi-
6 Ur of 1 somite in female. CR usually not diver- tes; inner border of A2 B with only I po inted
gent. In both sexes, A2 with both basal setae tooth ................................................................... 2
almost equal; male A2 without tooth on inner 2 Anal somite in dorsal view in fema le scarcely
distal border, often with fi ne hairs: s.g. Agetus longer than its width, in male at most I 1/4 times
(p. 974) as long as wide; in female Gns, in lateral v iew,
6a Ur of 2 somites in female. CR strongly diver- does not project over anal somite. Female Pd3 in
gent. Female A2 with 1 basal seta at least twice dorsal v iew broader than Pd2. In male, CR only
length of other seta. Inner dista l border of male 3/4 as long as Gns: Corycaeus (Corycaeus)
A2 tenninated by 1 large pointed tooth: s.g. clausi F. Dahl, 1894 (Fig. 7.487)
Corycaeus (p. 974) 2a. Anal somite in dorsal view in female 1 1/3 times
as long as w ide, in male more than I 1/2 times as
Key to species of Corycaeus subgenus Agetus (from long as wide; in female Gns, in lateral view, pro-
Dahl 1912) j ects over anal somite. Female Pd3 in dorsal
Females and males view as broad as Pd2. ln male, CR as lo ng as
1 Female CR, in dorsal view, about 1/2 length of Gns: Corycaeus (Corycaeus) crassiusculus
Ur. Male anal somite I 112 times as long as wide Dana, 1849 (Fig. 7 .488)
and 3/4 length of CR; eyes close together: Cory-
caeus (Agetus) limbatus Brady, 1883 (Fig. 7.485) Key to species of Corycaeus subgenus Ditrichocory-
la Female CR, in dorsal view, longer than 1/2 caeus (from Dahl 19 12)
length of Ur. Male ana l somite at most 1 1/3 Females and males
times longer than w ide and not 2/3 length ofCR; 1 Female anal somite about as long as Gns and
eyes always clearly separate .............................. 2 also as long as CR. Male CR less than I l/4
2 Female Ur slender and in lateral view dorsal times as long as ana l somite and 3/4 as long as
hump flattened; corners ofPd4, viewed laterally, Gns: Corycaeus (Ditrichocorycaeus) am azoni-
angular and projecting dorsally; A2 B toothed on cus F. Dahl, 1894 (Fig. 7.49 1)
Copepoda 975
X: dor- Ia CR in both sexes I 1/4 times as long as anal sexes with only 1 plumose seta: Corycaeus
~ somite and 1/2 as long as Gns: C01ycaeus (Urocorycaeus) furcifer Claus, 1863 (Fig.
~- pos- (Ditrichocorycaeus) ajjinis McMurrich, 1916 7.498)
lly:mC (Fig. 7.490) la Female CR at most 1 1/4, in male not I 112,
r.~-:er. times as long as remain ing part of Ur. A2 B in
Key to species of Corycaeus subgenus Onyclw- both sexes outer border not toothed ........ ...... .... 2
~"r3.1 corycaeus (from Dahll912) 2 Female anal somite slightly narrower than Gns
~of Females and males and in dorsal view anterior part with rounded
~on Female anal somite more than twice as long as expansion; seta on A2 B less than half length of
~Co::- wide; male Gns at most half as wide as long .... 2 seta on C; P4 Enp wi th 2 long setae. Male Ur of
~~.hcr Ia Female anal somite not twice as long as wide; 2 somites; P4 Enp with I long and I very short
reer of male Gns more than half as wide as long and in rudimentary seta: Corycaeus (Urocorycaeus)
~ dorsal view genital swelling projects strongly lautus Brady, 1883 (Fig. 7.499)
--~5:-! - postcrolaterally .................................................. 3 2a Female anal somite much narrower than Gns and
Female CR longer than Gns; in male at least 4/5 cylinder-shaped; setae on A2 B and C only
~ length of Gns; female body in lateral view slightly differing in length; P4 Enp with 1 long
strongly vaulted; male Gns ventral border has no and 1 rudimentary seta. Male Ur of I somite; P4
median hook: Corycaeus (Onychocorycaeus) Enp with only I seta: Corycaeus (Urocorycaeus)
eg
__Ur
,..,_ agilis Dana, 1849 (Fig. 7.493) longistylis Dana, 1849 (Fig. 7.500)
......1.;!.1<• • .:3 CR in both sexes not 2/3 length of Gns; female
rCi..'2l body in lateral view scarcely vaulted; male Gns Key to species ofFarran uta (from Dahl 1912)
r more ventral border has small median hook: Females and males
!~US Corycaeus (Onychocorycaeus) giesbreclrti F. 1 Female U r in dorsal view narrowing evenly
""'..!.S9) Dahl, 1894 (Fig. 7.494) towards posterior, in lateral view syl1ll1letrically
rx-.:m- Female Gns not longer than combined length of expanded proximally both dorsally and ventra l-
~:ed anal somite and CR; male ana l somite about 2 ly. Male CR thick, at most 1/3 length of rem ai-
I 3 times as long as wide: Corycaeus (Onycho- ning Ur ............ .... .............. .............. ... ............ .... 2
.....,._.....,~,-
~~~ - corycaeus) latus Dana, 1849 (Fig. 7.495) 1a Female Ur in dorsal view with undulating later-
~ ·~ Female Gns considerably longer than combined al borders; in lateral view with broadly humped
~~~
length of anal somite and CR; male anal somite dorsal surface. Male CR long and slender, about
! ?~~ at most twice as long as wide .. .... ............... .......4 112 length of remaining Ur ............. ............... .... 3
only Female Pd4 in dorsa l view projecting posterior- 2 Female Ur 4 times as long as CR which are near-
JDIDZS) ly into point; male Gns short, 4/5 as broad as ly 2 times as long as wide in dorsal view. Male
long: Corycaeus (Onychocorycaeu!.) ovalis Ur more than 3 times as long as CR which are at
G- ~ Claus, 1863 (Fig. 7 .496) most only 2 times as long as wide in dorsal view:
t::::::les as Female Pd4 in dorsal view usually projecting Farranula rostrata (Claus, 1863) (Fig. 7.504)
.-.;ro.- posteriorly into rounded lobe; male Gns with 2a Female Ur less than 3 times as long as CR which
~ expanded patt elongate, 2/3 as broad as long: are about 3 times as long as wide in dorsal view.
= ""'
- Corycaeus (Onyclwcorycaeus) pacijicus Male Ur about 3 times as long as CR which are
rurz1 Giesbrecht, 1891 (Fig. 7.497) about 3 times as long as wide in dorsal view:
Farranula curta (Farran, 1911) (Fig. 7.502)
to species of Cmycaeus subgenus Urocorycaeus 3 In lateral view female Ur with anterior thickened
~ Dahll9 12) part about same length as narrow posterior sec-
Fftnales and males tion: Farranulagracilis (Dana, 1849) (Fig. 7.503)
Female CR I 1/2, in male I 3/4, times as long as 3a In latera l view female Ur with anterior thickened
~~ remaining part of Ur. A2 B, in both sexes, outer part longer than narrow posterior section:
I { l. .! lx>rder with row of small teeth. P4 Enp in both Farranula concinmt (Dana, 1849) (Fig. 7.501)
g ~
riii01JU-
976 Copepoda
Acknowledgements Suggested readings
Responsibilities for compilation, Introduction: Elena Available literature on copepod morphology, anatomy,
Markhaseva and Janet Bradford-Grieve; Calanoida: phylogeny, life histories ecology and biology is very J
Elena Markhaseva for Calanidae to Mesaiokeratidae, extensive. Copepod specialists are usually special ized
Janet Bradford-Grieve for Arietellidae to in more or less narrow aspects of the study of these
Bathypontiidae. Cyclopoida (Oithonidae): Carlos animals; thus, those involved in studies of thei r biolo-
Rocha and Bemado do Abiahy. Harpacticoida : Janet gy, life cycles or, for example, molecular biology, are
Bradford-Grieve. Poecilostomatoida (Oncaeidae, usually not taxonomi sts, whereas taxonomists may
Sapphirinidae, Corycaeidae): Janet Bradford-Grieve. employ a limited range of teclmiques, often excluding
We would like to thank the following individuals for such approaches as DNA analysis, statistical methods,
their assistance: Knud Schulz (Zoological Institute etc. However, sooner or later every student has to deal
and Zoological Museum of the Univerisity of with taxonomic problems which he (or she) is often
Hamburg) for checking the sections Key for calanoid not trained to solve, at which point the availability of
:::
families and famil ies Spinocalan idae, Thmybidae and information aimed at helping to tackle these difficul-
Bathypontiidae and Rony Huys (Zoology ties becomes crucial. Publications containing basic
Department, the Natural History Museum, London) information are mentioned in this chapter in the
checked the section on external morphology. Gayle Introduction. The term inology adopted here for the
Heron (University of Washington, Seattle) assisted description of the external morphology of copepods
with the compilation of the South Atlantic Oncaeidae follows Huys and Boxshall (1991). This book is re-
fauna and checked the text. Nevertheless, any etTors commended for its modern views on copepod mor-
which occur are entirely our responsibility. We thank phology and contains a historical review of the main
the National Institute of Water and Atmospheric publications on copepods. Classical publications on
Research (NIWA) in New Zealand for providing the classification of copepods, either original or of a
facilities while we combined sections and for assisting review nature are: Giesbrecht ( 1892), Sars (190 1-
one of us (E.M.) through the NIWA Visiting Scientist 1903, 1924, 1924-25), Scott A. (1909), Wolfenden
Programme. This work was pattly supported by con- (1911 ), Rose (1933), Lang (1948), Sewell (1947), -
,.--
tract C0 1421 from the New Zealand Foundation for Brodsky (1950), Owre and Foyo (1967). During the
Research Science and Technology. Bernardo do last decades several manuals on regional copepod fau-
Abiahy's contribution was sponsored by FAPESP - nas or revisions of selected copepod fami lies have
Fundayao de Amparo a Pesquisa do Estado De Sao been published (Vervoort, 1949, 1951 , 1957, 1963,
Paulo, Brazil, Process Number: 96/2579-9. We are 1965; Tanaka, 1956, 1957a,b, 1958, 1960, 1960,
grateful to Demetrio Boltovskoy for his kind help. 1964a, b, 1969; Tanaka and Omori, 1969, 1970;
Heptner, 197 1; Matthews, 1972; Damkaer, 1975; Roe,
1975; Park, 1978, 1980, 1982; 1983a, b, 1995;
Bradford and Jillett, 1980; Bjemberg, 1981; Bradford et
a/., 1983; Heron eta/., 1984; Nishida, 1985; Bradford-
Grieve, 1994; Hernandez and Suarez Morales, 1994;
Ohtsuka et a!., J 994; Heron and Bradford-Grieve,
1995; Hi.ilsemann, 1966; Markhaseva, 1996), among
others.
Copepoda 977
~
~
lS -rer;
ri!tozed
-~
I'" .:""'c.O~ir
~
=-·· ue
E may
::riing
~-
deal ·I
h oiten
~·of
"
liifiru.I-
= '-'4:)(C
c iilc
•
<(Mx2
iile
Bathyca lanus inflatus
pepods
l:: r.s rc-
irl >::lOr- <(P2
II:! :::na:m
Megacalanus princeps
iJcs Ofi
,a . of :1
1901-
& •.;er.
~ !Q.!-=.
c;tle
c.::~ <( Pl
:s mve
1%3. Bathycalanus
1960. princeps
1910:
5: R...~.
99 ~:
e;
•
a:Srd-
~ 1~: Bathycalanus eltaninae
~ -~
~ Brady.calanus
~ g1gas Bathyca/anus richardi
smn:>e
5<1P!OUE?fE?:J
. su~6 su~6
r::::::::::::::-, \~
\ ~~,~J;.'' - ---------·::
\ ~
I
I '
su~6
Sd p '~-------·--·_;
51D!dAJ snuete:>ApeJg •
Ld6
su~6
J1 o
8L6
Copepoda 979
6l
9Gns
e Calanus agulhensis
~
9Gns
CD Ca lanoides macrocarinatus
1
I
OUr
1 i
[ .., ?
1
~
r\~
"7 \~
I
9P5
tl
OPSC
O PS
I
e 9P1 9 P2 O PS
•
Ca lanus propinquus
Ca lanus austra lis
J/UfMlep SnUT!fT!:JOWSO:J
Jadn-ed snuete:JOlj.JUe:J G
SdP
SdO Ldo
I
'I
:..-
---- \I
=
-
086
Copepoda 98 1
/ c'r.i.l:....~l
i~
- ~q
A'
u-
lr-114
~----- ~~
......
"'"" ~Go. ~ff
~ !~,
~q
r r~ I qJ~
<_j? Gns ..o.
O PS 4!-:.-rtf!.
,.,.-<"'A'.
'"/' <_j?PS
e Nannocalanus minor
e Neocalanus gracilis _j
-
<_j? Gns
A'
~ OGns
• !:'o
'
.
1\\
=-==~
<_j? Gns
OPS
-
Enp
OPS <_j?P2 <_j?PS
e Neocalanus robustior e Neocalanus tonsus

982 Copepoda
·: J
~
.B
lr.,...l, 9
>------1
~~~\
\:\
::
;I
//
::
!/
::
fD Undinula vulgaris
!/
!f
:•
((
OPS
$
e Acrocalanus andersoni
I \
9P4
e Acrocalanus gibber
9 OP4
Exp 3
9P4
OPS
fD Acrocalanus longicornis
e Acrocalanus gracilis
Copepoda 983
9Pl
e Acroca/anus monachus
9ur
9P3
8 Calocalanus at/anticus
I
I
(
~tf 9 PS
9
fJ!;
O PS d
s_m 9P5 9P5 e Ca localanus equalicauda
e Ca localanus contractus
--~
_./';do \
f
I
I
'
f iiOSOU OW Of SnU f?f FDOf f?:J
JOUfW snueje:JOjf?:J
1786
Copepoda 985
I
cf rs
• cf rs Calocalanus pavo
• Calocalanus plumulosus
---....1'
-----
----
-;....'
~ OPS
G•?5 cfrs OPS
• Ca localanus styliremis
snpnu sn!faa
986
snJ-epnu~p snu l?f I?:JI?ll?d ---
IJ~ U I?d'w-e::> snur?f i?:Jf?Jf?d
/
SUC)Q
•
~
L
!1/~M~S Sn!f ~Q
L86 BpOd;;)dO:)
988 Copepoda
~ <j? PS
't
~ 9P5
~
OGns
+ OPS
• Paracalanus indicus
• Paracalanus nanus
'
9 II 9
<j?Gns
•
O PS
Paraca lanus parvus
e Paracalanus tropicus
Copepoda 989
~'
,.: ;·
•
i
.
'tl....~
~
-/ ?l 'cI. t;r:~
l
\ c ~.;
O~
PS U OGns
-
I
~~~ • ·~ j
"!o-.,.J'; ,.... f
~
_,.,\'
5\
_
J,.·
·~
\'
I
e Parvocalanus crassirostris
~
)? P4
e Paracalanus quasimodo
)?Mx2
~~-.\
- )?P3 I . )?Mxp
?j-J
l
~
I
.;;! )?P3
...r-
~.
..
...,::::::::;iQ
)?PS
- ::r-;
)?P4
)?P2
e Parvocalanus scotti e Mecynocera clausi

O PS
990 Copepoda
9Mdp ~
e
•
Pareucalanus langae
Eucalanus h yalinus
I
I
I)~
9Mdp
• Pareuca lanus sewelli fD Rhincalanus comutus

Copepoda 991
I
I
I
I
~~
C-'-. ~
?C~~
• Rhinca lanus g igas
OUr
9Mdp
0 Subeucalanus crassus
--~
~P5
.- --' ;~
·~
J~t)_
9ur OPS
e Rhincalanus nasutus
992 Copepoda
~ l i OUr
<j?Pl
?Md) ~ ·
\ YJ{\ / <j?Mxp
?A2~
9
?"
1~ <j?Mx2
G Subeucalanus longiceps 0 Subeucalanus monachus
<j?Ce
OPS
<j?Gns
G Subeucalanus pileatus
G Subeucalanus subtenuis
Copepoda 993
I
t
~
'
~
<j?P2
<j? Pl
• M onaci//a gracilis
0 Mimocalanus cultrifer
O PS
G Monacil/a typica
G Spinoca lanus abyssa lis
994 Copepoda
~
-- ,\~
/
""
OPT
9Mx2
0 Spinocalanus brevicaudatus
•
~ c?: ,)
9
'vY'(R /) \ ·~
<J ·-
ri\,
J•
G~ 9Gns
OPS
• Spinoca lanus magnus Clausocalanus arcuicornis

Copepoda 995
~~"
9R ..<= !
/
'"
\
I '\
~ i
!
' 9 \I
trv I
! II
9R ! i
I
I
1
I
!' - - · - ;I
lA
t
~--------;
/.
',.
,:__
{:
.
I
__ /
-----.J
'\:,_x~
.. ·l
( \
·------ '-fl
i---J
::~to I I
::?
t;
::.---
6Jj
~
. .
OPS
9P5
9Gns
• O Pl e
--.......
Clausocalanus furcatus
0\)~ {f-//( f\ I
(
I
\~
i ._.,
I
'
P,
! \ r '----) J a \ ~ f/P
I
R I ()
II
0
9 {l r ·- \ ' +
I
•
+
I
\ ·,_, I
I
lq
')
I '\ l___ )~ .I 1n
lt i
. 9Gn;-------J c-.. \ ~n
- - . J\ "-~'-'/\\ lQ·,\'- •,__ft
~
(
~l
l t
·- -y---1 ~P3 B \ ·H·:_·
\\~
~} ~
. ---- I
_/ / 1
W, )) l! rvv\ /,! ~ OPS
o/ KL-1
9P5
~ +
dPS 9P5
• Clausocalanus ingens
• Clausocalanus jobei
996 Copepoda
rr· 9 l /\
9'~
9P3 B OPS
• Clausocalanus lividus
8
9P3 B
Cla usocalanus /aticeps
9
\Vf
J f~
I~ \ ) 9P3B
I
OPS
I \
~~/fr 10 Cla usocalanus minor
8 Clausocalanus mastigophorus
Copepoda 997
(\ ~
9R
d
5
~~I
y~ \dPJB 9P5
OPS
\:0'(P5 ', '[ c\ dP5 1
e Cla usocalanus parapergens • Cla usocalanus paululus
_..;,·
~~ 9R
I r-~~
I 9Gns
~--- I
~~
v
i"""'c,. _
.... " j
L-----! r~ I
~ ~ \'<PJ), ' I
/vi;',\ 9P4 Exp 2-3 OPS
,w \\
(p art.)
/ -( 8 Ctenocalanus citer
OP3 B
8 Clausocalanus pergens
snue11 snueJ e:>ouao
("lJed)
r-zdx3 vd6
I
I
~
snJedt:JJO) sndoued<uo 8
866
Copepoda 999
9
OPS
05~ 9
9Gns
~
~
~
Aetideus acutus
9Gns
~ OPS
&d
'--
- Vi
~"'
9 P4C
..--,...
OPS
OUr
~ Aetideus australis
e Aetideus armatus
Copepoda
9
9
(
~
~ (;; (
~ 9Gns
v;J
~~
9Gns
e Aetideus pseudarmatus
• Aetideus giesbrechti
9Gns
9MxpC
9MxpC
e Aetideopsis minor
• Aetideopsis carinata
Copepoda 1001
p 9ce ~
::J~ w
9Mxp c Aetideopsis rostrata O PS
-----------------------
9 ?J
~
lu.'.
9 P2
~"'
Chiridiella atlantica
• Chiridius gracilis
1002 Copepoda
9
9
e Chiridius polaris
• Chiridius poppei
~
• Chirundina streetsii
I ~) Chiwndinel/a magna
• Comantenna curtisetosa
..
Copepoda 1003
I~
OPS
• Euchire/la amoena
• Euchirella bilumida
OPS
OPS
~ OPS
• Euchirella curticauda 0 Euchirella formosa

1004 Copepoda
~. )?Ur
• Euchirella max ima

OPS
A • Euchirella m essinensis
messinensis
)?Ce
.~w N' I lOPS
~"' 1~;. J \.5) ~P4 (
:e . Euchirella pulchra
~
S Euchirella rostrata
Euchirella simi/is
Copepoda 1005
~ ~
9Gns
OPS
OExp 3 P 5
• Euchire lla splendens
• Euchirella truncata
9
~
9
!'i 9MxpC
(part.)
Ia.
> 9 Pl
0 9P4 C
Caetanus antarcticus
• Caetanus brevicornis
1006 Copepoda
9
A7\
9
'
9P4 C
{p art.)
«~:f.,c
0 Caetanus krupii
• Caetanus brevispinus
d rs
I I\)~
9
f.:J 9Pl
Caetanus miles
• Caetanus minor
Copepoda 1007
0
9
9
9Mx pC
(p art.)
\1 +- · IG Gaeta nus robustus

• Gaetanus pileatus
~R
MV!o$ 9
• Gae tanus tenuispinus 0 Lutam ator e legans

1008 Copepoda
•
<_(Mdp
Mesocomantenna spinosa
~
9
<_(Pl
• Paracomantenna gracilis
• Paivella inaciae • Paivella naporai

Copepoda 1009
.,..,
~
OPS
~ \-
~\\dp
C(A2
G>
g([\
Paracomantenna maga lyae
• Pseudochire lla batillipa
...
C(Ur
~
\ 9~;a~.)
G Pseudochirella dubia
• Pseudochire lla hirsuta
1010 Copepoda
f! (part.)
OUr
~~~L-
lcrr4c
• Pseudochire lla obtusa
• Pseudochire /la mawsoni
~H ~:~·
OUr
~-'
~ u 9Gns I 9
::J H ~\)
9 tJ
9'~
(part.)
• Pseudochirelfa spectabilis
~ (part.)
9Gns
~
\'rf, C(P4 C
•
(part.)
Pseudochirella p ustilifera
0 Pseudochirella spinosa
Copepoda 1011
~~
9 ~
l!;f· lr
9Pl l OPS
9Gns
~~
OUr
~-~:~' ~ ;r ' ~\\
~?Go.~
,.
OPS
hp ·'!
::..,..,? -
~--·
G Undeuchaeta incisa 9Gns
:J
Undeuchaeta major
~
;.-...,.
~/
--.:
11~ I I
.. .0
c- 9 RJ
~
_.. __
I
,-
9G ns
._... I:";;
IE
G Undeuchaeta plumosa
1012 Copepoda
<(Ur
dPS
<(Gns
Exp2
1a t:~al
~
<(Gns 9 1
9 ve~tral
V IeW
V IeW
<( Gns
G Euchaeta concinna
--------- -- -- --------
• Euchaeta acuta
It\
.~tdpsl
Exp 2-3
~ dp
'"'f''''
dExp 2-3
<(Ur (right
d view)
Enp & Li3 &

Li4 2-3
<(Mxl
!]! OPS
left (dorsal (left

ventral
lateral view view) view)
view
IS Euchaeta marina IS Euchaeta m edia
Copepoda 1013
~
9ce 9ce
9Gns
u 9Gns
9Gns 9ur
•
\ entral v iew 9ur
Euchaeta paraconcinna
G Euchaeta pubera
[j
~
9 Gns
---
2j
9Gns
O PS
a
9Gns
9ur
Exp 2-3
O PS
G Euchaeta spinosa
- Paraeuchaeta aequatorialis
1014 Copepoda
~
,,
'I
--,
,, <j:>Gns <j:>Gns <j:>Ce
~\
,,,, dorsal
~
left lateraI
view
view
<(Gns
tl
\ij OPS
d'Ps Exp 2-3

OPS
• Paraeuchaeta barbata
• Paraeuchaeta antarctica
<j:>Gns Exp2-3
right view OPS
cf
OPS
<j:>Gns <_(Gns 9Gns

OPS
left view ventral dorsa l view
• •
view
Paraeuchaeta biloba Paraeuchaeta bisinuata

Copepoda 1015
fl?5
cl-3
~~~.
I
9Gns 9Gns
......_
,. 9Gns
.q
• Paraeuchaeta
comosa
•
Exp2-3
- Paraeuchaeta calva
O PS
;r---
; \
Exp2-3 H. 9Gns
~.
O PS
I
·!
/
~
• Exp2-3
O PS ~
~
Paraeuchaeta exigua
9Gns
OPS
f\)V,
9Mxl
(part.)
(schematic
diagram)
Paraeuchaeta
G dactylifera OPS
• 9Gns
Paraeuchaeta gracilis
1016 Copepoda
<?Gns
~
~~-- ;/
1/
{
9ce
1
F
' -
::
Exp 2-3 '--..
<?Gns
OPS
9ur
• Paraeuchaeta hansenii
G Paraeuchaeta kurilensis
<?Mxl
(schemati c
diagram)
~
• Paraeuchaeta parvula
<?Gns
Exp2-3
OPS
O PS
• Paraeuchaeta pseudotonsa
Copepoda 1017
9Gns
•
9Gns
>
Paraeuchaeta rasa
.
,.
'
)
I
I
9
.,
.,
• Paraeuchaeta regalis
...,
Exp2-3
OPS
9Gns
Exp2-3
OPS
,,
9Gns • Paraeuchaeta scotti
• Paraeuchaeta sarsi
PS
1018 Copepoda
<(Mxl <(Mxl
(schematic
(schematic
diagram) d iagram)
• Paraeuchaeta sesquipedalis • Paraeuchaeta vervoorti
v;: J~ ~
? nNw ':( ?C• ?C•
I I Ill/
7\ O PS
dee
<(Ur
OPS
Q ~
•
OGns
•
<(Pl den
Va ldiviella brevicornis Valdiviella insignis
Copepoda 1019
OPS
or.:
•
OUr
9ur
•
Valdiviella o ligarthra
Valdiviella minor
y
~
£ 9A1
~~
M
9 Enp Mx2
?"
~
r1 ?C· \
~
/1
ZJ -- \ !¥(
~ Midi~
_J
n
I9 \
V
9R
e
~
.:
~- - }
! ?" . G 9P5 0Mx2

OPS
L Cornuca lanus chelifer
~
,
j
~,.--
. ~
~.r
~?PS
• Brachyca lanus bjornbergae
1020 Copepoda
yy
dR
__ , <;? R
0 ~
~'
<;?Pl
<;?PS
G
•
Neoscolecithrix caetanovi
Cornucalanus robustus
<;? Gns
<;?PS
Onchocalanus
0 magnus
G Onchoca lanus subcristatus

Copepoda 1021
H<_(R ;f:\
<_(Pl
<_(PS
-
~
c:l?i <_(PS
• Onchocalanus wolfendeni
• O nchocalanus trigoniceps
d'PS
9M'/ ~1%
~
<_(PS ,
~PS
S Xanthocalanus agilis
~
<_(Pl
'-(
• Phaenna spinifera <_( P2
1022 Copepoda
;~
<( ~ ~
~~
'/ ){f/) '\
<(P2
• Xanthocalanus soaresmoreirae
• Xanthocalanus marlyae
..
;
h
dPS
Exp3 (left)
<(Enp P2
~\
dEnp P2 I
• •
drs
Amalloth rix den tipes Amallothrix emarginata
Copepoda 1023
LiS
Mx2
(part.)
(schematic d iagrams)
Amallothrix hadrosoma
<(P3
~
""~
Amallothrix va lida ·c· fl
liiS:? ~ <(PS
•
<(R
-~ I •,
Amallothrix obtusifrons OPS
\ ',
; '
. ,
·7
~u
~ ':
0 /
<(R
D
·'
.:,.:
!) .
<(Enp P2
I'.L.-{
(7 <(PS
G Lophothrix frontalis
• <(PS
Landrum ius g igas
5!f!Wf5 Xfl}Ol.{dO?
5U01J!f!WI1l.{ Xfll.{JOL{d07
tZ:OI
Copepoda 1025
-,?'
0
9C• \~
OR
w OPS
!'i
0 Scaphoca lanus brevicornis
\_ <:j?Pl
• Scaphocalanus curtus
I 1) ~
<:j?Gns
9 <:j?R
<:j? PS
• Scaphocalanus echinatus
G Scaphocalanus elongatus
1026 Copepoda
9Enp P2
....•...
· ~··
O PS
• Scaphocalanus farrani
G Scaphocalanus magnus ___j
r;f;PS
OMxl
(schematic
diagram) 9Mx l
•
(part.)
Scaphocalanus
parantarcticus
OP2
• Scolecithricella abyssalis
Copepoda 1027
<(Enp P2
~ ~t
_. -· ' '
,·· .. . · -
9 ..· ;·
) ......... / d
.'
......
...' ·.~
•'
.·:
., ::
·'
7 ,---J
-- , :>-
Q~1
<(Gns <( Gns
0 Sco/ecithricella dentata
?
,\
~
:::::>
"I
).''
~ '
G
<(PS <(PS
Scolecithricella minor
dPS
~
_, ·----1
--------.}} ::
- - - -..___y
.1' ::
~®
____,.;
-·-·
""""'
<(Gns
<(Enp P2
cfps <(PS <(Mxl
(p art.)
•
<(Enp P2
e Scolecithricella ovata
Scolecithricella profunda
1028 Copepoda
-
8 Scolecithricella pseudoculata
<(Gns
• Scolecithrice lla tenuiserrata
'l
I
9Ur
<(PS
8 Scolecithricella vittata
~
<(PS
I uv dPS
~ Scolecithrix bradyi
Copepoda 1029
.......-..._
s~
'\
'
;=j
~
~/
'e
~
9Gns
<(P3
,....--
!i,. ~~
-' · ·::
..\
~ •'
' 1"\ '•

0,
..
---
---~
~.:
..... -.;
--........_ ::' •
.,
=::::..
~
!!
- v '•
--~~ \ / ;.
'•
OMx2 ·:•,
OPS '•
=
5i3'
'•
• Scolecithrix danae
~
<(R
I
:::;:=l
<(PS
)_)Enp P2
•
O PS
~\__9A2
•
Scottocalanus securifrons
5
Diaixis helenae
1030 Copepoda
Pedigerous
4&5
I
Pedigerous
4&5
<(Gns
G Diaixis hibernica tropica
<(P2 OP5
Oiaixis trunovi
Md
:j
• Tharybis asymmetrica
<(P5 ·OP5
• Tharybis megalodactyla
Mesaiokeras mikhailini
Copepoda 103 1
~- v<C:>r-
.,\~
~ -.J
.._,;
...
~9Pl
"9
d
9P5
~ <?Mx2
u /~:?
9 <?Mxl cfPs
~e
<? Gns
Mesaiokeras semiplenus
• Mesaiokeras tantillus
f:
V' ~ ~b4 l v~;: OAl
i "'\ - ~I
~-
OP5
OP5
<?R
e Arietellus aculeatus
• Ariete llus giesbrechti
1032 Copep oda
1(\ R
• Arietellus
minor
G Ariete llus pa voninus
•
OPS
Arietellus plumifer
1f\R
I End
OPS u)?PS
I End
drs
G Arietellus setosus
• Arietellus simplex
Copepoda 1033
- _:__·~.·.;
[I J"
.'
n~
-
;:;;;;;;:r-.
/
c::~I
j;' ~'
u .
? PS
?G ns
:;
• Paramisophria giselae
• Paraugaptilus buchani
~
~ R
Mxl
?PS
'--
.........
?PS
5
G Pilarella long icornis
G Augaptilus anceps G Augaptilus glacialis

1034 Copepoda
9c•f'0
d(?
• Augaptilus longicaudatus
G Augaptilus m egalurus
•
OPS
Augaptilus
spmrfrons
~/\
w
<(Ce
' ·.
n R
R . : ..
..
•.
~ ... ~'
<(PS
OPS
• Centraugaptilus cucullatus
• Centraugaptilus horridus
Copepoda 1035
,,
---- ------:~ \
k1R
: ~·
~l
<2 ·'::: 9
·.. :.
' . •.·. Md
i:
<(PS
--
'1: 2;) Pl
•
Mxp O PS
G Centraugaptilus rattrayi Euaugaptilus affinis
tO
~
Y} <(Ce
Exp
Mx2
<(Mxl
(schematic d iagrams)
5 <(A2 Exp
•
O PS
e Euaugaptilus a/iquantus Euaugaptilus angustus

1036 Copepoda
Exp Exp
<(Mxl
(schematic (schematic d iagrams) <(A2 Exp
diagram)
• Euaugaptilus antarcticus
• Euaugaptilus austrinus
n R
A~ Exp
<( Pl
Mxl r
• Euaugaptilus bullifer
OPS
G Euaugaptilus clavatus
Copepoda 1037
rr
R
)? PS OPS
G Euaugaptilus elongatus
P3 OPS
~
• Euaugaptilus facilis
0
n
R
4\d ~) )?Ce (
Mx l
A2 Exp
.....
r • Euaugaptilus gibbus
O PS
• Euaugaptilus filigerus
1038 Copepoda
,_
___,
-
<(PS
<(PS
<(Mx1
e Euaugaptilus hecticus
Mx1
OPS
0 <(Ce
• Euaugaptilus laticeps
P1
Mx1
• Euaugaptilus latifrons
~
I Exp
OPS
•
<(PS
Euaugaptilus /ongimanus
Copepoda 1039
0ctce
M<~
=
K
i;;:f,
~~' '1'\~
R
-...:=/
.J '--'
<_( PS
OPS
e Euaugaptilus magnus
•
O PS
Euaugaptilus maxillaris
~ <_(Ce 'M
R
Mxl
-
..1
...,... A2
~
r::.
• Euaugaptilus nodifrons
OPS
OPS
• Euaugaptilus oblongus
1040 Copepoda
9
h)
v <;? PS
~
Mxp
Mxl
I
~
OUr
• Euaugaptilus pachychaeta
•
Mxp
O PS
Euaugaptilus palumbii
A2 Exp O PS
G Euaugaptilus rigidus
<;?A2 Exp
(sc hematic di agrams)
8 Euaugaptilus perasetosus
Copepoda 1041
~
KR
rr
R
<(P5
::::.?5
~ Mxl
Pl
•
....._
~': . j:
~::~ll_:r
• Euaugaptilus tenuispinus
•
O P5
Euaugaptilus squamatus
I
f)\
- <(Ce
~
dee
<(P5 t;
~
::::es
"
i.---,.
--
--
Q· :q ,{\
I
--{ ~9
•
),
OP5
Haloptilus angusticeps
O P5
e Haloptilus acutifrons
1042 Copepoda
<j?Ce
cfl <j?Ce
•
Gn <j?PS
OPS
H aloptilus chierchiae
-
OPS
IS>
• Haloptilus fertilis
?(\
------
1
-
9ce
~
Mxl
Haloptilus longicirrus
~ H aloptilus fans
Copepoda 1043
9 11'\
9R
(schematic d iagram)
G Hafoptilus major
~~-OP5
C>l"
Hafoptilus longicornis
1;\ 9ce Mx1
I 1§\
• Haloptilus mucronatus
O PS 9ce
Mx1
'(
:xe\
?"" ~
.
k
·~
·~ 9Ur
~ ~
\\xl
C'!?3 9 Mx2
OPS
~
G Haloptilus ornatus
Md
e OPS
Haloptilus ace/latu s
1044 Copepoda
0 lour
Mx2
7~
~~~
•
Mxp
OPS
t\ • Halopti/us plumosus
~ 9Ce
R \~
Mxl
9P5
• Haloptilus spiniceps
• Pachyptilus abbreviatus
Copepoda 1045
9
~~l
\f
. 1:
Gn.
llf\ 1M AUA I I
~
\~
~
Gn
~~ Mxl
C(PS
Mxp
G Pachyptilus long imanus
• Pontoptilus ova/is
I
d' PS
~
Disseta palumbii
>\
Mx2
• Heterorhabdus abyssa lis
\
i
O PS
Utf?J
C(Gns C( Gns C( Gns
• Heterorhabdus clausii
• Heterorhabdus austrinus
1046 Copepoda
--
.......
OPS
OPS
• Heterorhabdus compactus
L'ftt.\
~ Heterorhabdus lobatus
aw
<:(Gns <:(Gns
zc"'&td Mx 2
• <:(PS
OPS
Heterorhabdus no rvegicus
• Heterorhabdus papilliger
Copepoda 1047
w 17\R
<j>Gns
OPS
e Heterorhabdus robustus
• Heterorhabdus
spinifrons
r-....
_.;r \
[r.s
rt
r Gn
OPS
• •
Mxp
Heterorhabdus tanneri Heterorhabdus vipera

1048 Copepoda
~
OPS
<:(PS Exp2
• H eterostylites longicornis
OPS
9
• Heterostylites major
Gn
Mx2
Mxl
• M esorhabdus angustus
• Neorhabdus falciformis
Copepoda 1049
~'K
~~
Cc .
•' <(Gns
..._...,./
-
._;;;-;:::::::_.-
Ce
_ ::;/
I
v
.:- -==-<
-
-....-
.....
~'
.:::.:.::::'
.-
~, '
<(Ur
Q
d'Lr <lu.;
Lucic utia aurita

cf PS • Lucicutia bicornuta
',\
•'
•\
.,
:'•' 0
~~
•'•'
•'
,,"
ol
"I"
'•
\',
•',
\ cfPS
~!I ~
K d Ps
G
Pl
<( PS lcfPS
Lucicutia clausii
• Lucicutia curta
1050 Copepoda
C(Gns
OPS
...
•
C(Ur
Lucicutia flavicornis • Lucicutia gaussae
OUr
C(Ur
OPS
Ce Ce
• Lucicutia gemina
• Lucicutia grandis
Copepoda 1051
r - - - - - --,
;---- --
-----:
OPS
~
!-I' I
~
l!
I
9ur
h 9ur
• Lucicutia long icornis

• Lucicutia Iuc ida
-l 9
9
9ur
OPS OPS
t• Lucicutia magna
"G Lucicutia macrocera
~ G Lucicutia maj or
1052 Copepoda
<j?Ur
C\ 0 <j?Ce
<j?Gns
OUr
<j?Ur OPS
Pl <j?PS
OPS
0 Lucicutia ova/is
• Lucicutia maxima
,,------...
L--
------,
• <j?U r
Lucicutia wo /fendeni • Causs ia
asymmetrica
Copepoda 1053
~ ~\, ~
~~
G Gaussia princeps
• Metridia brevicauda
~
~-
<(Ur
OPS
OPS
• Metridia calypsoi
• Metridia curticauda
1054 Copepoda
C(
CR
t:J
C(P2 End 1
• Metridia discreta
G
C(PS
Metridia gerlache i
OPS
C(
v Pd 4 +5
d' Ps
C(PS
• Metridia tonga
• Metridia lucens
Copepoda 1055
'~
:.....
C( PS
• Metridia princeps
• Metridia venusta
~
Cj>Ur
!L
G
Cj>PS
'5
rd'Al
OU r Ven
G Pleuromamma abdomina/is G Pleuromamma borealis

1056 Copepoda
-~-~
~~ <(PS
<(Ur
dPS
<(PS
lo'A1
18
17
18
19-21
• Ple uromamma piseki
• Pleuromamma gracilis
dPS
OPS . lo'A1 ~
·~l'::~
17 17- - - .
- 18 19-21
18
19-21
• Pleurom amma quadrungulata

• Pleuromamma robusta
Copepoda 1057
_,...,
;:" .--; I'

1-
/,
..
........___ PS
If
'
• 9P5
Pleuromamma xiphias
G Phyllopus aequalis
.- --
--
y
I
--.;
I:;?d 4+5
G Phyllopus bidentatus
• Phyllopus he lgae
1058 Copepoda
<j?PS
• Phyllopus impar
• Centropages brachiatus
<(PS
Exp
G Centropages bradyi
•
_ _ ____Centropages calanin~
Copepoda 1059
---l..
-
rcfPS
~
Cj>Ur
rOAl
0 Centropages chierchiae
<j?PS
<j?Ce
• Centropages elongatus
- <j?Al <j?PS Exp2, 3
-
-......
,.......
~ <j?Ur
rO Al
:~
cfur
<j?PS
O PS
Centropages furcatus
• Centropages gracilis
1060 Copepoda
<(PS
dorsa l
view
OPS
ventral
view
G Centropages longicornis
G Centropages kroyeri
<(Pl
t1
1?r±t r OAl
19
I• I
• Centropages orsinii
Centropages typicus
Copepoda 1061
cfPS
<j?Ur
c
<j?PS Exp
Centropages violaceus
• Pseudodiaptomus acutus
-.-J
__.
t,'
r
v <j?PS
<j?PS
cfPs O PS
Pseudodiaptomus charteri G Pseudodiaptomus hessei
1062 Copepoda
<(Ur
<(Ur <(PS
K OPS
w <(Gns • Pseudodiaptomus nudus
<(PS
• Pseudodiaptomus marshi
O PS
~
<(Ur
•
<(PS
OPS
Pseudodiaptomus serricaudatus
<(PS
G Pseudodiaptomus richardi
Copepoda 1063
9
0
,..,
Gns
ventral v iew
.........___
__/
4J Gns
lateral view
9 P5
OPS
G Temora discaudata
OPS
'i)pS
G Temoropia mayumbaensis
OPS
PS Temora longicornis
9
•
9P5
• Temora stylifera
OPS
Temora turbinata
1064 Copepoda
c=_)
I
o' Gns
i,
,-.,
yPS
• 9ur
9ur
Candacia bipinnata
o'Gns
~
s 9ur
9Pd 4+5
19
• Candacia armata
18
cfPS
~ 14
8 Candacia curta
rOA1
9ur
9ur
• Candacia catula
9ur
r;j
• Candacia cheirura
OPS
Copepoda 1065
o'U r
=:_j
8 Pd 4+5
0~"')?ens
• Candacia elongata
•
<j?P5
Candacia ethiopica
dens
15 [)
)?ens
dA1
e
~
Candacia longimana <j?P5
/
• Candacia magna
19-20
)?ens
~~
dens
<j?P5
i----
_/' <j?P5 0- 1rd A1

dA1
dens
~
• •
<j?Ur \
Candacia
Ca ndacia norvegica Candacia paene/ongimanaiG pachydactyla
1066 Copepoda
G Candacia varicans
• Candacia tenuimana
rd' A1
17-18
19-20
G Paracandac ia bispinosa
<j?PS
• Paracandacia simplex
<j?
16
15
• •
rd' A1 OPS
Paracandacia truncata Calanopia am ericana
Copepoda 1067
;/
<j_>PS
..
• Calanopia elliptica
• Calanopia minor
our
~
<j_>Ur
• Labidocera acuta
1068 Copepoda
<(Ur OPS
•
OPS
• Labidocera acutifrons
Labidocera darwinii
<(Ps
co
rOAl
OPS
• Labidocera fluviatilis
• Labidocera detruncata
~
~
~·
(
690I
1070 Copepoda
r~
,;.II
f\ <(~Ce,
~~
<(Ur
<(Ce OUr O PS g
•
OUr
•
+ven
Labidocera wollastoni
Ponte/fa atlantica __.....,
OS
~
~
<(PS
8 Ponte/fa diagonalis
• Ponte/fa fera
Copepoda 1071
?PS~ N ~h ;~ \ ~ D~
~
.o ; )
~~ f.~
''='
""""
:~:, ~~~·'A If m~~
\_r O PS
6
/l J
~ '~ / / \ '. 1 ~B!Ul~~ <(PS
• Pontella gaboonensis
• Pontella lobiancoi
---
9 ~~ 0
<(Ce .
~
~ <(Ur
~)
~ ref Al
<(PS
G Pontella marplatensis
<(Ur
dorsal
view
<( PS
<(U r
•
ventral
view
Pontella securifer
1072 Copepoda
()0
<( Gns
•
<(Ur
ventral
view
Pontella spinipes •
<(Gns ra ps
• cfPs
Pontellina plumata
• Pontellopsis brevis
I?JnJJUa:> l? tJJ I?:JI;I I?SOIJ!A SfSd01Ji3JU0d
I
Sd p
r---
1---
····~
£LOI
Sdp
aeuep f?!JJ f?:JV G
SdP
!2Joqafll!/ f?!JJf?:JV
Sd O
•
~
'--"
'---""
I
.
.
I
epod~do:) tLOI
Copepoda 1075
·'
________..,
~
~
.:..:=. ~ O PS - -
OPS i ·'
dur
Acartia tonsa
G Tortanus gracilis
~
? .,A R
i'
I I OPS
G Paracartia africana
1\ R
.. d Ps
)>Ur
)>PS
PS
• P3 Bathypontia e/ongata
0 Bathypontia sp.
1076 Copepoda
((j ~Lf
~-Q
<(Ce <(Pl Exp
'•11'\
<(P4 Exp
G O ithona atlantica 0 Oithona attenuata
~~\
1/<(C~ <(Mxl
~~\~CR~ ~
,\
l '(Mdp '(M' I
<(Mdp
<(Pl
•
<(Pl
<(P3 <(P4
G Oithona bjornbergae Oithona brevicornis
saqaLf eUOLfJ!O eJeWeLf euo Jl
dx:HdO 4.0
dp~
ep!2u; euOLIJ!Q
dx3 ( dO
LLOI
E?J;}j!WnJd eU04J!Q
dx3 ldO •
G eueu eu04J!O
8L01
Copepoda 1079
~
~'~
~c{
~
.--:-:--,_.
~ ·~
0
-~ ~
~
_________,
--- -------.:
------..... 9ce 9Mx1
~
-=---
8~
--!-"\
-
.... - . 9~ ~ - 9~
~~
!i
~.
~V/
• ,_'4 ~
't)t'
/or 1
- Exp
G Oithona pseudofrig ida
G Oithona rigida
~
~
~
v~
____) 9Mdp
..-.)
J- \
-=1
~
~~
OP 1
- Exp
Oithona robusta G Oithona setigera

1080 Copepoda
<i)p4
9P3
~ Oithona simplex 9 P4
~
9Pl o·h 9 Pl
0 tt ona simi/is
I
~s
~ 9Mxl
9Pl
9P2 Exp 9 P3 Exp 9P4 Exp
0 Oithona tenuis
9P2 9P3
G Oithona vivida
Copepoda 1081
-
::.
~
_J I Y.~ \\ f, :£ IP I: v, i I !\. I '+' \ flY )\ l f //1\\ OPS
I r-~ it'~i
JiG ~ II
I
,.. ,--'
____J '~i'l• I II \
\
I
~~
(~
.__, I
~ \ -A II
-~-·- ..... , IIi ____;,.{) 1------f-_. I
'
I
<jlP2
\
U ---9"/ <jlMdp
f t
-· <jlCR
:--,...... <jlP3
.__/
• 9
Paraoithona flemingeri
G Microsetella norvegica
11.
\'
\llicrosetella
<jlPS
rosea G Macrosetella gracilis I 0 Miracia efferata

1082 Copepoda
u
Ce + Pdl
~
C? PS
I
0
I
Oculosetella gracilis
0P1 ~PS
G Miracia m inor
G Clytemnestra rostrata
Mxp
G Euterpina acutifrons
0 Clytemnestra scute llata

Copepoda 1083
Pi O PS
!I
S?PS
IIj,
ji
1:
l'
II
!I
19~P
i
e Aegisthus aculeatus
y
~~- G Aegisthus mucronatus

~. ./ '
/'
........
_..___.
~I
.~-~
~ :\t\p
C-
,
• Conaea expressa
G Conaea rapax
t80 T
Copepoda 1085
....---...
, .....
~
I
-~-'
,.--,
,...,_,.......
~
~
<(Ur
OUr
OUr
ID' ~
v /W', I 9 \
OMxp OPS
P3 Enp +
G Oncaea clevei
"Oncaea" atlantica
~
·f.-~
I
<(P4 A \
~u, ] ~rs
OP5
c:
_..
.
~
\\
.,_
~ ~ ~PS
~
'
~
h ~ ~
... /)
b OMxp
<(Mxp O Mxp
e Oncaea curvata G O ncaea englishi

Copepoda
<j>Mxp
~
OMxp
G> Oncaea /atimana
<j>Mxp
dMxp
0 Oncaea lacinia
t< JJ
a ''
dMxp
'-l
l OMxp 0 Oncaea mediterranea
0 Oncaea media
Copepoda 1087
~
.• •r
\
·-:: 1·
!
:---1
:_ _ j
P? .
~~
U)
<j?Mxp
~~
[5J
~"
0 Oncaea prolata
cfMxp
<j?Mxp OMxp
1.---
e Oncaea pari/a
:..___-iL' 0
~ <j?
t::j
~ 1 e ~
'1 ~
?"
<j?P4
0 Pachos tuberosum
~ OMxp
<j?Mxp
O ncaea scottodicarlo i
I
dxwp
•
-upod;:ldO:) 8801
Cop epoda 1089
...._______.~.
~,..,.
,
~
~"' .
- ) /
-
~--
-
-
~ ~ ~-----
OA 2 OMxp I
<(U r
- Enpl ~
G Copilia vitrea
0 A2
~
~-
• Copilia q uadrata
-·
·--
----
)_,
O P2 Enp 3
•
2
<(A
Sapphirina angusta
O P2 Enp 3
G Sapphirina auronitens-sinuicauda
1090 Copepoda
?Q _V,J OA2
Sapphirina gastrica
•
OP2 Enp 3
d P4 Enp
Sapphirina bicuspidata
OP4 Enp 3
G Sapphirina iris
~OAl
OP2
Enp 3
G Sapphirina intestinata
G
Copepoda 1091
"""'!'--..
OP2
dP2 Enp 3
OA2 G Sapphirina opalina-darwinii
<j_)A2
G Sapphirina nigromaculata
c"'
OAl
~ O P2
Enp 3
~ .... G Saphirina scarlata
---'
~
;;:::::::::L
-
--
~
f ,, OA2
J~.
<j_)A2 OP2 ~
0
Enp 3 OP4
<j_)Ur (It) oCR
Saphirina ovatolanceolata-gemma
/frrAl<j_)CR
<j_)Al OP2
Enp 3
0 Sapphirina stel/ata
1092 Copepoda
G Coryca eus (Agetus) limbatus

~OA2
0 Corycaeus (Agetus) flaccus
/fj
0
• Corycaeus (Agetus) typ icus
Corycaeus (Corycaeus) clausi
S?A2
G Corycaeus (Corycaeus) crassiusculus

Copepoda 1093
r-
<(A2
9P3 )
~
~OA2
0 Corycaeus (Corycaeus) spec iosus
OA2
G Corycaeus (Ditrichocorycaeus) affinis
<(A2
;------
e Corycaeus (Oitrichocorycaeus) amazonicus OA2

<(A2
0 Corycaeus (Monocorycaeus) robustus

1094 Copepoda
. , OA2
• 0. • • Corycaeus (Onchocorycaeus) giesbrechti

_ cf A2
Corycaeus (Onychocorycaeus) agilis
9A2
cf A2
0 Corycaeus (Onchocorycaeus) latus

G Corycaeus (Onchocorycaeus) ova /is
Copepoda 1095
..........
OA2
G Corycaeus (Onychocorycaeus) pacificus
0 Corycaeus (Urocorycaeus) furcifer
.,. cf A2
Corycaeus (Urocorycaeus) lautus

~ 0 Corycaeus (Urocorycaeus) longisty lis
Copepoda 1097
F~ i. Illustrations of the species (sec text, Identification, for abbreviations used).

F.~ . :>Ources: from From Alvarez ( 1984): 124; from Alvarez (I 985a): 175; from Alvarez ( 1985a): 231 ; from Alvarez (1986): I 03, I 25, 128;
t= -\ndronov ( I 972): 41; from Andronov (1973a): 220, 221; from Andronov ( 1973b): 34 (all except male, male P5), 35 (all except male, male
2'5 ~ 3'". from Andronov ( 1976): 2 17, 218; from Andronov ( 1977): 49; from Andronov ( I 979): 2 14, 215 , 216; from Andronov ( 1995): 219; from
~ (1958): 3 1; from Bj0rnbcrg (1968): 2, 3, 7; fi·om Bj0rnberg ( 1975): 183; from Bj0mberg ( 1980): 42, 44; fi·om Bj0rnberg and Campaner
. with kind permission from Kluwer Academic Publishers: 3 15, 3 16 (all except male P5 , fern. P5); from Bowman (1958): 24 (all except
~ =le P5); from Bowman (1971): 46 (male P5), 50; from Boxshall ( 1977): 45 1, 452 (male, fern.) ; from Boxshall and Bottger (1 987), by
:>:ilOD of Oxford University Press: 454 (fern.); from Bradford (1969): 93 (male); from Bradford (1970): I I (male Ce, male Ur, male P 5),
- 2.... except fern . Ur, fern . Gns, fern. Mx I); from Bradford ( 197 1a): 89 (male), 92; from Bradford (1971 b): 149 (fern. Gns, left lateral view;
: - 6. 179, 200; from Bradford (197lc): 275 (male), 284. 285,288,289 (fern. Gns), 290 (male P5, fem. Ur); from Bradford ( 1974): 222
cale P5, male A I); from Bradford and Jillctt (1980): 90, 100 (fem.), 104 (male), 105 (male), 106 (male), 109 (male); from Bradford et
~31: 141. 144, 147 (fem.), 149 ((fern. Gns, dorsal view; fern. total view), 151, ! 52 (fem.), 154, 156, 158 (fern.), 161 , 164, 173 (all except
~ Ce. R). 180, 184 (P5 fem.), 185, 190, 198, 20 I, 204, 205, 206, 207 (all except fcm. Mx I), 210, 211 , 212, 213; from Bradford-Grieve (I 994):
J=.. l:r. fem. Md, fern. P5 , fern. PI , fem. P2), 11 (fern., fem. P5), 13, 14 (fcm. Ur, fern. P5), 15, 17, 18, 19,20 (all except male, male P5,
•= ..
~ 22 (all except male, male P5), 23 (all except male, male P5), 27 (all except male P4 Exp3, male P5), 34 (male, male P5), 35 (male,
?5 '· 3- (all except fcm. Ur, fem. P4, fem. P5), 46 (all except male P5), 53 (all except male), 55 (all except male), 56 (all except male), 58
~--ept male P5), 59, 60 (all except male, male P5), 6 1, 73, 74 (fcm .), 75, 76, 77, 78, 85 (all except fem. P4 Exp2-3), 334, 406 (male), 449
~fern.); from Brady ( 1883): 412 (all except fem. Ur); from Brodsky ( 1950): 188, 27 1; from Brodsky (I 972): 9 (fem. Ce, fem. Gns), I 0 (all
=::;:t fern.). II (fem. Gsn, fem. Ce), 20 (fem. Gns), 21 (all except male P5); from Brodsky and Zvereva (1976): 174; from Brodsky et a/.
>t : male, male PIS), 5 (all except male; as Bathycalanus bradyi), 48, 65, 67 (fcm.), 68 (all except male P5), 69, 70, 71 (all except male,
~ .. 184 (all except fe rn. lateral view, fem. R, male lateral view, male P5), 185; from Campaner (1977): 229 (fem. , fem. P5); from
~ (I 978a): 96, 129; from Campaner (1978b): 172, 182; from Campancr (1979): 208; from Campaner (1984): 203 (fcm. Ur, fcm. Gns,
lob I). 107 (fcm. Mx I), 209; from C laus ( I 863): 278 (male P5, Mx I), 452 (all except male, fem.); from CotTa! (1972b): 30, 36, 37 (fcm.
i=. P-1. fem. P5), 38, 40, 53; from Dahl (1912): 484, 485, 486, 487, 488, 489, 49 1, 492, 493, 494, 495, 496, 497, 498, 499, 500, 501 , 502,
::.:'"-. from Dakin and Colefax (1940): 23 (male P5); from Damkaer ( 1975): 71 (male, male P5); from De Decker eta/. ( I 99 I): I 2; from
~ I9- 2): 356; from Deevey (1973): 230; from Fanan ( 1908): 165 (fern.); from Farran ( 1929): 162, 195, 364 (male); from Farran and
-.;;:u; I 95 1): 4, 9 (fem., fem. P5, male, male P5); from Ferrari and Bowman ( 1980): 417, 436; from Fleminger ( 1957): I 45; from Flcmingcr
5J (all except fern. R, fem. Mdp, fem. PI), 55 (Ur and P5 male), 56 (Ur and P5 male), 60 (male, male P5), 62.63 (fem.); from Flcmingcr
3'>3: from Fleminger and Hiilsemann (1974): 398, 399; from Frost and Fleminger ( 1968): 72, 74 (male), 79, 80, 81, 82, 83; from Fliichtl
~55 (P4Enp); from Gaudy (1963): 3 18; from Giesbrecht ( 1892): 10 (fem.), 20 (male, male P5), 21 (male P5), 43 (fern. P5), 54 (fcm. R.
q.. 63 (male), 64, 142 (fem.), 143, 18 1 (all except fcm. , fern. P5), 203 (fem. P5), 234 (male, male PS, fern. P5), 25 1, 265 (Mxl), 269,
=-: c.=..e. 27-1 (male), 276 (male P 5, Mxl), 279 (male P5), 286, 290 (Mx2), 292 (Mx2), 294, 3 17, 337,339, 340,341 ,342,344, 346,348,
- _35~. 360. 38 1, 382, 384, 386, 387, 389, 390, 392, 394, 396, 397, 400,401 ,404, 405,406 (fcm.), 407,408, 410, 438 (fem.), 445 (fem. A I),
...,.:, -=-3- male), 470 (male), 47 1 (male), 472 (male, fem., fem. CR), 473 (all except fcm. A2, male P4, male P2 Enp), 474 (male, male P4 cnp,
.!-5 (fem., fem. C R), 476 (male, male P4Enp, male CR, fem.) , 477 (all except male P4 Enp3, fern. A2), 478 (male, male Ur, male P4),
~e. fern. , fern. C R), 480 (male, fem.), 481 (male Ur, fem., fem . Ur), 482 (male, fcm.), 483 (male, fem. , fern . CR , fem . A I), 427 (male
C:xn Giesbrecht ( I 902): 32 I, 456 (fem.); from Gonzilles and Bowman ( I 965): 5 I, 425 (male, fem., fcm. Ce); from Gordejeva (I 973): 448;
&lrdeje,·a ( 1975): 459; from Grice (1959): 267; from Grice ( 1962): 28,374 (male); from Grice ( 1963): 36 1 (all except fcm .), 362,363,
:!e= '· 365, 366, 367, 368, 369, 370, 37 1, 372, 373, 374 (fem.), 375, 376, 377; from Grice and Hiilsemann ( 1967): 87 (all except fem. P5,
- t : ,. 11 5 (male); from Grice and Hiilsemann (1968), by pennission of the publisher, copyright 1968 Uni versity of Hawaii Press: 163, 189;
<r.nd.ley (1963): 350, 35 1, 353; from Heptner ( 197 1): !59; from Heron ( 1977): 449 (all except male, fem. ), 453 (fem.), 456 (all except
=le ~Lxp, male P5, fern. Ur), 457, 46 1 (fem.), 462 (fcm.) , 463 (all except male); from Heron and Bowman (I 971 ): 84, 85 (fcm. P4 Exp2-
::::= Heron and Bradford-Grieve (1995): 449 (male, fem.), 453 (male), 456 (male, male Mxp, male P 5, fem. Ur), 460, 464; from Heron et
S- -ISO. 458,46 1 (male), 462 (male), 463 (male); from Hiilscmann ( 1966): 299, 300 (all except male PS), 301 , 302 (fem. Ur, fcm. PS),
-·"-'-· 305. 306, 307, 308, 309 (male P5), 3 I 0 (male PS), 3 12, 3 I 3, 3 14; from Hiilsemann ( 199 I): 86; from Jespersen ( 1934): 296 (male P5);
G: 1968): 22 (male, male P5); from Johnson ( 1938): 54 (fcm. PI ); from Lang (1948): 441 (male. male PS), 444, 446; from Lchnhofcr
.!M. -167, 468, 469 (all except male), 470 (all except male). 47 I (all except male); from Lehnhofer ( 1929): 472 (all except male, fem.,
CR). .!-3 (fem. A2, male P4, male P2 Enp), 4 74 (all except male, male P4 cnp, fem.), 475 (all except fern., fem. C R), ), 476 (all except
J:Le P-!Enp, male CR, fcm.), 477 (male P4 Enp3, fem. A2), 478 (male A I, fem . A2), 479 (all except male, fern. , fcm. CR), 480 (all except
!'::=. '· -181 (all except male Ur, fern., fem. Ur), 482 (all except male, fcm.), 483 (male A2, male C R, male P2Enp, fem. A2); from
~-a (1996): 88 (fem.), 89 (fern.), 91 (fem.), 93 (fem.), 94, 95 , 98, 99, 101 (all except male), 104 (fcm.), 105 (fem .), 106 (fcm.), 108,
:i:=.l. I 10 (all except male Ce, male P5), I ll , 11 2, 113 (fem .), 11 4 (fern.), I I 5 (fern.), I I 6 ((all except male P5), I I 7, 118, I 19, 120, 12 1,
- r:.:. 130. 13 1, I 32, 133, 134, 135, 136, 138, I 39, 140; from Matthews ( 1972): 24 8 (male), 259,263 (male); from McMurrich (1 9 16): 490;
~ I 937): 16, 23 (male), 338, 343, 345; from Nishida ( 1985): 4 I 5, 416, 418, 41 9, 420, 421, 424, 425 (fem. Mdp, fern. Mx I, fcm. P5),
.~ !:.2S., .!~9. 430, 43 1, 4 32, 43 3 (redrawn), 434 (redrawn), 435 (redrawn); fi·om Ohtsuka el a/. (1994): 222 (male P5 E nd), 226 (male P5), 227
?: End). 228 (male P5 End), 229 (fern. Gns); from Owre and Foyo ( 1967): 57, 18 1 (fem. , fem. P5); from Park ( 1968): 11 4 (male); from
-..~ ,. 88 (male), 9 1 (male); from Park ( 1975a): 100 (all except fcm. , male P5); from Park (1975b): I 13 (male); from Park (1976): I 10
Ce. :nale P5); from Ramirez (I 966): 385 , 395; from Rocha ( 1986): 422, 426; from Roc ( 1975): 68 (male P5), 170 (male), 192 (male), 256
2:0: (male), 266 (male); from Saraswathy ( 1973): 316 (male P 5, fcm. P5); from Sars (1903): 272 (fem.), 322, 323, 325, 33 I (all except
3-!-. 358, 36 1 (fem .), 413 ; from Sars (1924-25): I (fem. R, fem. Mx2), 5 (male), 67 (aJJ except fern., male P5), 146, 147 (male), 150,
1098 Copepoda
Fig. 7. continued.
152 (male), 157, 168 (fem.), 169 (fem.), 173 (fem. Ce, R), 178, 187, 19 1, 192 (fem.), 223, 225, 226 (all except male P5), 227 (all except mak
P5 End), 228 (all except male P5 End), 232, 233, 234 (fern., CR), 23 5, 236, 237, 238, 239, 240, 242, 245 (fem.), 246, 247, 248 (fem.), 249, 25C.
252, 253, 254, 255, 256 (fem.), 257, 25 8, 260 (fem.), 262 (fem.), 263 (fem.), 264, 265 (fern.), 266 (fcm.), 268 (fern.), 270,274 (fcm.), 276 (fern.. ..
277 (fem.), 279 (all except male P5), 280, 282, 283 , 287,289 (all except fern. Gns), 291 , 292 (all except Mx2), 295, 296 (fern.), 297,309 (fern.. ,
320, 324, 332, 333, 335, 336; from Schmaus and Lehnhofer (1927): 58 (male P5), 297; from Scott (1894): 278 (Mx2, Mxp), 355, 388, 423; fro;:;
Scott (1 909): 8, 142 (male), 168 (male), 199, 222 (fem. R), 378, 379, 380; from Sewell (1929): 25, 26, 17 1; from Sewell ( 1947): 6, 268 (male .
fro m Shmeleva (1967): 454 (male); from Shmeleva ( 1975): 29, 32, 33; from Silas and Pillai {1 973): 402, 403; from Stephen and Saradb 'i
(1973): 265 (male); from Steuer {1 91 5): 4 11; from Steuer {1926): 4 12 (fern. Ur); from Steuer (1 932): 326, 327, 328,329, 330,33 1 (male AI r..
from Tanaka (1 957a): 100 (male P5); from Tanaka (1957b): 101 (male), 102 (female); from Tanaka (1958): 148, 153; from Tanaka ( 1960): r .
275 (fe rn.), 455; from Tanaka (1961): 196; from Tanaka (1964a): 293; from Tanaka ( 1964b): 245 (male), 260 (male); from Tanaka (1969): 10:
(male Ce); from Tanaka and Omori (1969): 107; from Tanaka and Omori ( 1970): 169 (male); from Ueda and Hiromi ( 1987): 409; from Vcrvoo;:
(1949): I02 (male P5); from Vervoort {1 95 1): 9 (all except fe m. C and fem. Gns), 14 (all except fe m. Ur and fem. P5), 3 19; from Vervoor.
(1 957), by courtesy of the Mawson Antarctic Collection, University of Adelaide, Australia: 184 (fern. lateral view, fern. R, male lateral vie\\ .
male P5), 277 (male), 302 (male P5); from Vervoort (1963): 24 (male, male P5), 43 (all except fem. P5), 45, 52; from Vcrvoort ( 1965) : 126.
393; from Vyshkvartseva ( 1993): 197; from Wells (1 970): 437, 438 (fern. P5), 439, 440, 44 1 (all except male, male P5), 442, 443, 445 (ai:
except fern. A 1), 447; from Wheeler (1970): 127, 4 14; from Wilson (1950): 67 (male P5), 116 (male P5), 391 ; from With ( 19 15): 158 (male ~
165 (male); from Wolfenden (1906): 177; from Wolfenden {19 11): 27 (ma le P4 Exp3, male P5), 66, 87 (fern. P5, fern . Ur), 97, 137, 170 (fern. ,_
193, 194, 222 (fern.), 224, 281 , 298, 300 (male P5), 310 (fern. Ur, fern.), 3 11; from Wright ( 1936): 349, 352, 354.
Mysidacea
Masaaki Murano
Introduction arctic species, varies from 7 to 15 days depending on

water temperature (Murano, 1964). On the other hand,
The order Mysidacea belongs to the phylum a deep-sea gigantic mysid, Neognathophallsia ingens
Arthropoda, subphylum Crustacea, class Malacostraca, was estimated to have a period of larval development
superorder Peracarida (Bowman and Abele, 1982). in the marsupium of 530 days (Childress and Price,
The superorder Peracarida consists of 6 orders, includ- 1978). In N. intermedia, release from the marsupium
ing Spelaeogriphacea, Amphipoda, Isopoda, Tanaida- starts about one hour after sunset. One or two indi-
cea and Cumacea, in addition to Mysidacea. With these viduals at a time are released at intervals of 20-30
groups, Mysidacea have in common that adult females seconds. Liberated juveniles begin to swim and feed
are provided with a marsupium between pairs of pereo- immediately after release. The female mysid, after
pods and keep their embryos in it until they grow to releasing all the juveniles, undergoes a sequence of
juveniles. Hence, Mysidacea are also known as "opos- molting, copulation, and subsequent spawning with-
sum shrimps", as well as "mysids". in a night. Spawning is carried out even in the
absence of males, but these non-fertilized eggs are
The first mysid species was described by O. F. Müller abandoned after one or two days. Released mysids
in 1776 as Cancer jlexuosus, now known as Praunus need about a month to reach the adult stage at about
jlexUOSllS, which occurs abundantly along coasts and 20°C (Fig. 2).
estuaries of northem Europe (see Tattersall and
Tattersall, 1951). Many discussions on the taxonomic The life-span of N. intermedia is estimated to be five
position of mysids were made by carcinologists during months for a winter generation, two months for spring
the 19th century. Comprehensive classification systems and autumn generations, and one and a half months
currently used for this order were established by Boas, for a summer generation. On the other hand, in N. ing-
Calman, Hansen and several others. Historical develop- ens the life-span of females was estimated to be 2950
ment of the taxonomic status of Mysidacea was days, with the onset of reproduction at 2400 days
described in detail in "British Mysidacea" by Tattersall (Childress and Price, 1978).
and Tattersall (1951). Recently, it was proposed that
the suborder Lophogastrida should be elevated to the Mysids seem to prefer animal-matter as food, but
status of order (e.g. Schram, 1986). In this book, how- stomach contents of those collected in coastal envi-
ever, 1 follow the older classification. ronments are in most cases dominated by detrital,
unidentifiable materials with small amounts of diatom
Mature females spawn their eggs inside their own shells, tintinnid loricae, bodies and appendages of
marsupium. The brood size varies according to spe- small crustaceans, etc. In general terms, mysids are
cies, and to season and body size within the same spe- sea-bottom scavengers. Many reports on the relation-
cies. In most cases, the number of eggs spawned is ships between mysids and fish indicate that mysids
fewer than 100, but Mysis stenolepis breeds as many can be one of the most important food organisms for
as 140-245 eggs (Amaratunga and Corey, 1975). the latter, especially in coastal regions where they
Brood size in mysids was summarized by Mauchline occur in abundance. Besides fish, birds (Prince, 1980,
(1980). At the time of spawning, a female copulates Schneider, 1981) and whales (Hudnall, 1983) have
with a male, so that "eggs" produced into the marsu- been reported as predators of these crustaceans.
pium are actually embryos. Hatching to larvae an~
subsequent growth occur in the marsupium. In East Asian countries, mysjds are frequently caught
commercially for human consumption and as food for
Morphological changes of embryos and larvae in cultured fishes. In Japan Neomysis intermedia and N.
Neomysis intermedia are shown in Fig. l. The breed- japonica are used for a popular dish known as "tsuku-
•
ing period in N. intermedia, which is a temperate-sub dani", a food boiled in soy sauce, and in southeastem
SOLlthAt/anlie Zoop/an/...1on.
edited by D. Bo/tovskoy, pp. 1099-1140
1999 BaekhLl)'s PLlblishers, Leiden, The Nether/ands
1100 Mysidacea
ti-
0.5 mm 0.5 mm 1.0mm
Day 1 Day 2 Day 3
Day 5 Day6
Day 7
Fig.1. Growth of Neom)'sis intermedia embryos in the marsupium at 21-22°C. After Murano (1964).
() y = 0.064 X + 0.819
e :¡: T T
S ~U~reru0:0e(])
Cl..
3.0
1.5
2.5
2.0 +, ±
~rl i/ /~
..c 1.0
1'-
E
0.5 O
spawning
first ') 1¿ll----r T
t'-
ó Y=0.029X+1.621
y = 0.060 X + 0.694
5 10 15 20 25 30 35 40
Days
Fig. 2. Growth of Neomysis intermedia at 20°C after release. After Murano (1964).
Mysidacea 1101
Asia some species (name unknown) are used in the ish-water lakes and rivers. The most typical fresh-water
preparation of a particular type of sauce. species is Mysis relicta, which is found in lakes of
northern North Ametica and Europe. It evolved from
Mysis oculata as a result of isolation in fresh waters
Sampling methods during glacial times (Tattersall and Tattersall, 1951 ).
Besides M. relicta, there are about 30 species of mys-
Mysids live in various habitats in the sea. Therefore, ids occun·ing in fresh-water lakes, rivers, and fish
collection methods must be chosen according to the ponds. They include An.tromysis in the Amazon basin,
habitat in which mysids live. An.tromysis and Taphromysis along the coasts of the
Caribbean Sea and the Gulf of Mexico, Paramysis and
Pelagic species. Mysids are usually scarce in the water Diamysis in eastern Europe, Mesopodopsis and
column, so vertical hauls with ordinary small plankton Nan.omysis in southeastern Asia, and Neomysis in
nets are not effective. For conical nets, large mouth northern North Pacific coastal lagoons. Most of them
diameters (1 m or more) are required. The author has inhabit more or less brackish waters.
collected mysids successfully with oblique or horizon-
tal hauls with an ORI-net with a mouth area of 2 m2 In the oceans, mysids live in various environments,
(Omori, 1965). For micronektonic mysids, such as such as the water colwnn, on and just above the sea-
Eucopia, Gn.athophausia and some others, the Isaacs- floor, within bottom substrata, in caves, and as com-
Kidd Midwater Trawl (e.g. Pequegnat, 1965), and the mensals of other animals. They occur most abundant-
RMT (e.g. Hargreaves, 1984, 1985, 1989) are useful. ly in shallow coastal waters, in the near-bottom layers
just above the sandy or muddy bottom, rocky shores,
Epibenthic and near-bottom species. Mysids are and coral reefs, sometimes in dense swarms. Their
usually much more abundant on and just above the sea biomass is high, but estimates of its magnitude have
floor, than in the water column. Devices for collecting hardly ever been made because of practical limita-
them have been designed by Bossanyi ( 1951 ), tions. In the water column of the open sea mysids are
Wickstead ( 195 3), Beyer (1958), Macer ( 1967), rather rare, although in tropical and subtropical
Tokioka (1968), and Omori (1969). These are plank- regions, at the surface, several species, such as Siriella
ton nets mounted on sleds towed on the sea floor. thompson.i, are sometimes abundant during the night.
They are equipped with opening-closing or only clos- There are no epipelagic species in cold waters, includ-
ing devices for precluding contamination of the catch ing the Arctic and the Antarctic. Mesopelagic species
with organisms from the upper layers. are also rare, although their numbers are greater than
those in the surface layers. On the contrary, at depths
Species along rocky shores and coral reef lagoons. greater than 700 or 800 m, mysid biomasses exceed
Carleton and Hamner (1987) designed a diver-operat- those in the upper layers. Murano et al. ( 1976) esti-
ed device for quantitative sampling in coral reef mated the average density of mysids in a 0-1000 m
lagoons where deployment of boat-towed devices is deep water colmnn at a station off middle Japan at 0.7
extremely difficult. The device is a 10 m2 trap exploit- mg m-3 during the day, and 0.6 mg m-3 at night; these
ing the escape behavior of motile epibenthic animals, values represented 0.3% and 0.5% of the total bio-
such as mysids, that can be herded. Visual collection mass, respectively (Fig. 3). Below 500 m the biomass
with a scoop net during SCUBA diving is also useful was dominated by Eucopia and Boreomysis.
for collecting mysids in cavities, m spaces among
rocks, and in pelagic swanns. Vertical distribution and migrations
Many species of deep-sea mysids have been found on

Geographical and vertical distribution and just above the ocean-floor at various depths, but
our knowledge on these organisms is very poor, both
Habitats and geographic patterns taxonomically and ecologically. The deepest recorded
abyssal mysids areAmblyops magna (from 7 120-7260
Mysids are originally marine animals, but a few species m in the Kurile-Kamtchatka Trench; Birstein and
are known as marine relicts inhabiting fresh- and brack- Tchidonova, 1958), followed by Mysimen.zies hadalis
1102 Mysidacea
(from 6146-6354 m m the Peru Trench, Bacescu, g (wet weight) per 1000 m 3
1971), and A. aequispina (from 5760 m in the Kurile- 0. 2 0.1 0 0.1 0.2
Kamtchatka Trench, Birstein and Tchidonova, 1958). 0 c
Most sand-burrowing and coastal mysids perfonn a
die! vertical migration, ascending and dispersing into 200
G
the water column during the night, and descending to
Day Night
deeper layers towards dawn. In meso- and bathypelag-
ic species, however, it seems that clear evidence of 400
migration is generally lacking (Fig. 3), although a few
]
..c
I
reports on vertical migrations of deep-sea mysids have
been published (Waterman et al., 1939; Murano, 1977).
a<II
0 600 I
Tal
South Atlantic Mysids
800
Mysid collection records in general are considerably B.

fewer than those of other zooplankton because mysids
are difficult to collect with ordinary plankton nets, and 1000
South Atlantic records are especially limited. In this

region, the most important contribution was made by Fig. 3. Day and night vertical distribution of mysids. After
0. S. Tattersall (1955). She examined materials col- Murano et al. ( 1976).
lected by the RIV "Discovery" and other vessels dur-
ing the period 1925 to 1950, and reported 95 species, tion map. The geographic and vertical distributions of
including 28 new species. Species from the west coast each species, as well as the correponding body
of Africa and around the Falkland (=Malvinas) Islands lengths, are shown in Table 3.
covered herein owe a great deal to this report. Species
from the Brazilian coasts were examined taxonomical- Epipelagic species (0-200 m) in open seas.
ly by da Silva (1970a, b, 1971a, b, 1972, 1974, 1979), Anchialina typica and SirieLla thompsoni. Both are
dos Reis and da Silva ( 1987), and Bacescu (1968a, b, distributed world-wide in warm water regions. A. typ-
c, 1969, 1984, 1986); and ecologically by Almeida ica from the Atlantic is known as A. typica typica and
Prado ( 1973 , 1974). There are no reports of mysids that from the Indian and Pacific Oceans as A. typica
along the coast from southern Brazil to Argentina, orienta/is (Nouvel, 1971).
except for Mysidopsis rionegrensis, which was
described by Hoffineyer (1993) based on specimens Mesopelagic species (200-800 m) in open seas. 11
from San Matias Gulf, and species around the Arachnomysis /euckarti, A. megalops, Caesaromysis
Falklands (=Malvinas), which were collected by the hispida, Echinomysis chuni, Euchaetomera glyphi-
"Discovery" expeditions. Taxonomic studies on the dophtha/mica, E. intermedia, E. tenuis, E. typica, E.
west coast of Africa were made at two localities, the zurstrasseni, Euchaetomeropsis merolepis, Katery-
southernmost area (0. S. Tattersall, 1952, 1958, 1962, throps oceanae, Longithorax capensis, Meterythrops
1969; Nouvel, 1973; Wooldridge and McLachlan, picta. All these species belong to the tribe Ery-
1987), and around the equator (0. S. Tattersall, 1957, thropini. Many of them are cosmopolitan, but E. zur-
1961; Wittmann, 1986, 1992). Ecological studies were strasseni has been recorded from the southern hemi-
performed on the southwest coast of Africa by sphere, including the Antarctic, and E. intermedia and
Wooldridge (1981 , 1983, 1986, 1989), Webb and L. capensis have not been reported from the Pacific.
Wooldridge (1990), and Webb et al. (1987).
Bathypelagic species (deeper than 800 m) in open
Nearly 100 species of mysids are known from the seas. Chalaraspidum a/atum, Lophogaster spinosus,
South Atlantic Ocean to date, but many of them have Gnathophausia gracilis, G. zoea, Neognathophausia
only been recorded once or a few times. There are no ingens, N. gigas, Eucopia australis, E. grimaldii, E.
species with enough records for compiling a distribu unguiculata, E. scu/pticauda, Boreomysis acuminata,
Mysidacea 1103
Mysids Euphausiids
Not fused with 4th to 8th
Carapace Fused with all of thoracic somites
thoracic somites
Endopods of 1st & 2nd pereopods Modified into gnathopods Not modified
Absent or if present, covered
Gil ls Developed on bases of thoracic limbs
by carapace
Statocyst ___1Jsually p resent on end2£Sld of uropods j
Not present on endopod of uropod2.__
Female pleopods Usually reduced to unjointed, single lobe Developed into bilobed, natatory organ
Usually developed on 3rd or 4th, Developed on inner lobes of
IMale copulatory organ or on both 3rd & 4th pleopods 1st & 2nd pleopods
T able I. Differences between the external morphology ofmysids and euphausiids.
B. atlantica, B. bispinosa, B . il/igi. The first nine spe- African coastal species. Lophogaster challengeri,
cies, except L. spinosus, are cosmopolitan. Some Boreomysis insolita, Siriella africana, S. day i, S. wolffi,
Boreomysis species have also been recorded from the Anchialina truncata, Gastrosaccus brevifissura, G.
North Atlantic or the Indian Oceans. kempi, G. namibensis, G. psammodytes, G. olivae,
Erythrops africana, E. bidentata, Hyperamblyops meg-
Coastal species along northern South America alops, Katerythrops resimora, Afromysis hansoni,
(Brazilian coast). Siriella chierchiae, S. melloi, Leptomysis capensis, L. megalops, Mysidopsis bispino-
Bowmaniella atlantica, B. brasiliensis, B. dissimilis, sa, M. came/ina, M. eremita, M. major, M. schultzei, M.
B. inarticulata, B. recifensis, Katerythrops brasiliana, simi/is, M. suedafrikana, Nouvelia valdiviae, Paralep-
Brasilomysis castroi, Metamysidopsis elongata atlan- tomysis dimorpha, Mesopodopsis wooldridgei. Some of
tica, M macaensis, M. munda, M. sw!fti, Mysidopsis these species also inhabit the European and Mediter-
coelhoi, M. juniae, M. sankarankuttyi, M. scintilae, ranean coasts, and off Africa in the Indian ocean.
M. tortonesei, Promysis atlantica, Mysidium gracile, Gastrosaccus, a genus related to Bowmaniella, is dis-
Heteromysis mureseanui. Some of these a lso occur tributed in Europe, Africa, India, Australia, and New
along the Central and North American coasts, but Zealand, but is absent along North and South America.
there are no species in common with the African coast
and other regions. All the Bowmaniella species are Species of undefined distribution. Eucopia lingui-
known from the coasts of the American continents cauda, Heteroerythrops purpura. Collection records
and the West Indies, except for B. banneri, which has are too scanty to infer their general distribution pattern.
been recorded off California.
Coastal species along southern South America Taxonomy

(Uruguay to South Georgia). Lophogaster muranoi,
Hansenomysis falklandica, Amblyopsoides obtusa, General morphology and main diagnostic features
Pseudomma calmani, P. magellanensis, P. minutum,
Mysidopsis acuta, M. eremita, M rionegrensis, The Mysidacea are characterized morphologically as
Arthromysis magellanica, Neomysis monticellii, N follows (Fig. 4 and 5). Almost all of the cephal on and
patagona, Mysidetes anomala, M. brachylepis, M eras- the thoracic somites are covered by the carapace. The
sa, M. intermedia, M. macrops, M. patagonica. Lopho- carapace is shield-like, fused dorsally with the head
gaster muranoi and M. rionegrensis were collected off region and not more than four anteri or thoracic
the coast of central Argentina. The other species have somites. The eyes are usually stalked and movable, the
been recorded from the Subantarctic regions around the cornea generally developed with visual elements. The
Falkland (=Malvinas) Islands, in the Strait of Magellan, cephalon bears five pairs of appendages; antennule,
and the southern part of the Patagonian Shelf. antenna, mandible, maxillule and maxilla. The thorax
has 8 pairs of pereopods, all of which are divided into
1104 Mysidacea
cervical
sulcus carapace
abdom inal sornites ------ --- ------
inner antenna!
flagellum of flagellum
antennule
Fig. 4. A typical mysid in lateral view.
two branches, endopod and exopod. Endopods of the "Schizopoda" for some time in the past. Differences
first and second pereopods are usually transformed between their adults are shown in Table 2.
into gnathopods, which differ considerably from the
remaining limbs. All exopods are natatory. Females Features of importance for identification of the ani-
bear a marsupium composed of less than seven pairs of mals include:
lamellae, in which the embryos grow. The abdomen
consists of six somites, generally of the same form, Integument. External skeleton covering the body. It
although the last somite is longer than the others. Each is usually thin and usually poorly calcified, but cal-
abdominal somite, except for the last, is provided with cification is conspicuous in species of the families
a pair of pleopods. Pleopods have a flagelliform exo- Lophogastridae and Gnathophausiidae. Usually the
pod and endopod, frequently reduced in the female and surface of the integument is smooth, but in some spe-
sometimes in the male, often sexually modified in the cies the body is partly or totally covered with minute
male. The posterior extremity of the body is a tail fan spinules, bristles or scales. In rare cases the carapace
formed by the telson and a pair ofuropods. A statocyst and the abdominal somites can be armed with long
is usually present in the endopod of the uropods. spines (Fig. 6.42, 6.49), or a row of short spines on the
posterior somites of the abdomen.
Mysids are similar in their general shrimp-like
appearance to the euphausiids. Actually, these two Carapace (Fig. 4 and SA). The anterior margin is pro-
animal groups were combined in the order duced frontally in various degrees to form a rostrum.
Mysidacea 1105
antenna!
rlagellum
~
antenna !~
peduncle
rostrum~
t-----1 ~ cervical
su lcus
~ carapace
rostrum
2
4 abdominal
somites
5
incisor
process '\.
lacin ia ~
mobili s ~
antenna!
molar
peduncle process-..,..
Typical mysid in Mandible with

dorsal view 3-segmented palp
1&
" ~apica l
lobe
margm
0
Antenna! scale
Maxillule Maxilla
1106 Mysidacea
dactylus
\ :-----,<;~rpopropodus exopod
endopod
I
claw
epipod
0
Typical male
First pereopod pleopod
8th male pereopod
0
Typical female Cross section
pleopod Telson
of abdomen
Uropods
Fig. 5. Morphological details in mysids. F, G, H, M: after Tattersall and Tattersall (1951).
The posterior margin is usually emarginate and some- stalk and visual elements (Fig. 6.37). Some species
times furnished with spines (Fig. 6.34) or lobes (Fig. have two corneas in a single eye (Fig. 6.45).
6.3 1). In a few cases humps or protuberances are
present along the mid-dorsal line (Fig. 6.72). Antenna! scale (Fig. 5B, C). The antenna! scale con-
sists of a pair of thin lobes, which are modifications of
Eyes (Fig. 4). The eye is usually composed of two por- the exopod of the antenna. The shape varies variously
tions, the visual part (cornea), and the stalk. The stalk is according to species, such as oval, lanceolate, styli-
frequently provided with a digitate to papilliform pro- form, triangular, etc., with rounded or pointed apex.
cess on the dorsal surface. The comea is developed to The inner margin is usually setose. The outer margin
varying degrees, tending to be reduced with increasing varies specifically: naked, setose, dentate, or spinose.
habitat depth. In deep-sea mysids the eyes are frequent- Sometin1es this scale is divided distally into two seg-
ly modified into plate-like structures without a definite ments by a more or less distinct suture.
Mysidacea 1107
Outline classification and diagnoses of supra-

Number of
Decade generic taxa
Species .
<1900* ----1- 135 Table 2 shows the number of new species erected
1 each decade during the 20th century (species
1901-1910 107
described before 1901 are lumped). As the table
19 11-1 920 45
shows, more than half of the known species were
192 1-1930 54 described during the last 50 years, and there still is a
193 1-1940 61 steady tendency for the number of species to increase.
1941 -1950
1951-1 960
1961-1970
I 33
108
144
The classifica tion system given below may be revised
as new species are added in future. Underlined taxa
are those treated in detail in the present chapter.
1971-1980 11 6
198 1-1 990 131 Phylum Arthropoda
Subphylum C rustacea Pennant, 1777
Class Malacostraca Latreille, 1806
Table 2. Number of new mysid species erected each decade during
t:e ::Oth cenntry (*species described before 190 I are lumped). Superorder Peracarida Caiman, 1904
Order Mysidacea Boas, 1883
Suborder Lopho~astrida Boas, 1883
£odopods of pereopods (Fig. 50, H). Those of the Family Lopho~astridae Sars, 1870
C:st and second, and sometimes third, pereopods are Family Gnathophausiidae Bacescu, 1984
;:::::()(jified into feeding appendages, called gnathopods Family Eucopiidae Sars, 1870
« maxillipeds. In the third to eighth pereopods, the Suborder Mysida Boas, 1883
::!:pUS and propodus are usually fused with each other Family Petalophthalmidae Czemiavsky,
- form a carpopropodus. The carpopropodus is sec- 1882
tr.C.arily divided into subsegments. Sometimes the Family Mysidae Dana, 1850
=nber of subsegrnents is specifically diagnostic. In Subfamily Boreomysinae Holt and
50!:le species the second or third endopod has a sub- Tattersall, 1905
=Zlelare termi nation (Fig. 6. 15). Subfamily Siriellinae Norman, 1892
Tribe Siriellini Murano, 1986
l"ropods (Fig. 5L). These are composed of two Tribe M etasirieJiini M urano, 1986
~hes, endopod and exopod. The endopod is usu- Subfamily Rhopalophthalminae Hansen,
- shorter than the exopod, setose all around, and 1910
f.:::llshed with a few or many spines on part of, or Subfamily Gastrosaccinae Notman, 1892
t::tmghout, the inner margin; it may also lack spines. Subfamily Mysinae Hansen, 1910
-:.c outer margin of the exopod is armed with spines, Tribe Aberomysini Biicescu, 1986
setae. or, in rare cases, with both. Tribe Erythropini Hansen, 1910
Tribe Leptomysini Hansen, 1910
Tdson (Fig. 5K). The telson is the most important diag- Tribe Mancomysini Bacescu, 1986
li:I:Sl<: character. Its shape is triangular or trapezoidal, Tribe Mysini Hansen, J9 J0
w.....:. broad or narrow apical margin, or linguiform with Tribe Heter omysini Hansen, 1910
~y or narrowly rounded apicaJ margin. Its lateral Subfamily Mysidellinae Hansen, 19 10
c::=;ins are naked, or entirely or partly armed with Family Lepidomysidae Clarke, 1961
r=es which may be dense or sparse. The distal margin Family Stygiomysidae Car oli, 1937
_ =ned with one or several pairs of spines and some-
~ a pair of apical plumose setae. A deep or shallow
::..5:31 cleft is frequently present. The cleft is unarmed or
.I::::led with spines on the margin, and sometimes bears
tz:r of plumose setae at the anterior end of the cleft.
1108 Mysidacea
[ Species
Female
length
(mm)
M ale
length
(mm)
Depth range (m)
and onshore-
offshore
distribution
Geographic
distribution in the
South Atlantic
World wide
geographic
distribution
Chalaraspidum alarum 38 35 > 1000 Oceanic

~
Off Caee Town _ _World-wide, 402 N-502S
Off Cape Lopez,
1
Lophogaster chal/engeri 24 50-300
off Cape Town
Lophogaster spinosus 300-700 Off Rio de Janei ro W Atlantic, 342 N -25°S
Lophogaster muranoi 13.4-19
30 16-
3019 170 San Matias Gulf
Cnathophausia gracilis up to 138 up to 134 >700 Ocea nic TroJJi ca l World-wide, 35 2 N-20°S
Cnathophausia zoea up to 85 up to 70 >700 Oceanic Tropica l-Temperate World-wide, 62°N-35 2 S
Neognathophausia ingens up to 350 up to 160 >600 Oceanic Trop ica 1-Temperate World-wide, 402 N-402 S
Neognathophausia gi~s up to 160 ~~1 64 >600 Oceanic Temperate-Subantarctic World-wide, 602 N-692 S
Eucopia sculpticauda ~ to 60 up to 55 >700 Oceanic Tropical-Tempe~ World-wide, 602 N -402 S
Eucopia australis up to 53 up to 54 >600 Oceanic Whole area World-wide, 602 N-65°S
Eucopia unguiculata up to 38 up to 32 >600 Oceanic Whole area World-wide, 602 N-60°S
Eucopia grimaldii up to 38 up to 40 >800 Oceani c W hole area World-wide, 602 N-602 S
Eucopia linguicauda 41 unknown 1000-1600 Off Cape Town O ff S Africa
Around Falkland
Hansenomysis fa lklandica 15.5 15 40-500
(=Malvinas) lsi.
Boreomysis bispinosa 26 22 700-1400 Oceanic Whole area?
Boreom ysis acuminata 19 18.2 1400-2000 Ocean ic Tropical Atl a nt ic~ 52 N-152 5
Boreomysis atlantica 23 22(imm.) 1500-2500 O cean ic Whole area? Atlantic, 382 N-502 S
Off Ango la, E Atlantic, W Indian,
Boreomysis illigi 16 0-1000
off Cape Town 302 N-342 S
Boreomysis insolita 8.2 9 50-250 Walv~
Siriella chierchiae 9.5 9.6 Coastal Off Brazil north of 252 S W Atla ntic, 302 N-252 S
Siriella wolffi 9.4 8 (irnm.) Coastal Off Luanda
Siriella dayi unknown 10 Coastal Off Cape Town
Siriella m elloi 9? Coastal Off Natal
- - -
Siriella thompsoni 5.5- 10.2 5.5-11 .5 Surface- Oceanic
- -
Tropical-Temperate World-wide, 402 N -402 S
Off Cape Lopez, SE Atl antic, SW Indian,
Anchialina truncata 9 9 Surface
off Cape Town 02-352 5
Anchialina typica 5.3-5.7 4.6-7.2 Surface Oceanic Tropical World-w ide, 352 N-30°S
Tropical W Atlantic,
Bowmaniel/a brasiliensis 9-10 9-10 Coastal Off Rio de Janeiro
35~N -30°S
Coasts of Rio de Janeiro
Bowmaniel/a dissimilis 8.8-12 7.5 Coastal W Atlantic, 402 N-252 S
to Recife
Bowmaniella atlantica 5.5 9 Coastal Off Rio de Janeiro
--+--
Bowmaniella inarticulata
-tI
unknown 9-11 Coastal Off Natal
Bowmaniella recifensis unknown 10-11 Coastal Off Rec ife
Castrosaccus brevifissura 10.5-12 11 -11.5 Coasta l O ff Cape Town
S Atlantic, Indian,
Castrosaccus kempi 9 9 Coastal Off Cape Town
22 2 N-352 S
Castrosaccus namibensis 10-11 9-1 0 Coastal Off Namibia
Castrosaccus psammodytes 14.2-14.5 11.6 (imm.) Coastal Off Cape Town
Castrosaccus olivae 9- 10 9-10 Coastal Off Caee Town
Pseudomma m~ellan en sis 9.4 9.25 300 , Around Tierra del F~1 Southern S America
Around Falkl and
Pseudomma calmani 6-8 7-7.5 94-219 Southern S America
(=M alvinas) lsi.
Pseudomma minutum 4.2-4.6 4.0-4.4 100-1so Around Tierra del Fuego Southern S America
Mysidacea 1109
Depth range (m)

Female Male Geographic World wide
length and onshore-
Species length distribution in the geographic
(mm) (mm) offshore
distribution South Atlantic d istribution
ji:"''5 Around T. del Fuego,

4mblyopsoides obtusa 20.8-22.2 20 300-550 Southern S America
Falkland (=Malvinas) lsi.
Caesaromysis hispida 6.3-8.2 8.0-8.2 200-700 Ocea nic W hole area World-wide, 55°N-40°S
=x Atlantic, Indian,
=chinomysis chuni 7.2-11 7-9 340-2500 Oceanic 382 S,30"W
15°N-40QS
Euchaetomera tenuis 6.9-9.7 5.8-7.5 250-550 Oceanic W hole area World-wide, 402N-40QS
~ Euchaetom era typica up to 8.9 up to 7.0 100-300 Oceanic Who le area World-wide, 402 N-402 5
-""'5 Circumantarcti c,
Euchaetomera z urstrasseni 6.8-10 6-8 100-2500 Ocean ic Cold water
ir?"S 302 S-65 2 S
Euchaetomera Off Cape Verde,
5.9-8.7 6.0-6.6 100-550 Oceanic World-wide, 452 N-302 S
~
- phidophthalmica Orange R. mouth
...,_ E.xhaetomera intermedia 6-9 6-7 100-200? Ocean ic Whole area World-wide, 202 N-38°S
..---..s Exhaetomeropsis m erolepis 5.4-7.4 5.7-5.9 100-600 Oceanic Whole area World-w ide, 40°N-402 S
~rachnomys is leuckarti 5.3-7.7 6.3-6 .7 50-500 Oceanic Whole area World-w ide, 402 N-35QS
~rachnomys is megalops 6.4-9.5 7.1-8.5 200-500 Oceanic Whole area World-wide, 402 N-402 S
£r,Throps africana 3.8-4.6 4.4-4.6 Coastal Off Cape Lopez
=-- mrops bidentata 5.5 unknown 50-100 Off Cape Town
S Atlantic, NW Pacific,
~eroerythrops purpura 5.8-6.2 6.3 500-600 Oceanic? Off Angola
352 N-232 5
---'--
J:.a:erythrops brasiliana 2.8-3 2.8 Coastal Off Natal
Atlanti c, W Pacific,
tU;erythrops o ceanae up to 9.3 up to 10.8 300-700 Oceanic Whole area
Indian, 50°N-40°5
5'5 K.i:erythrops resimora 9.8 1000? Off Cape Town
Atlan tic, W Pacific,
!l.fderythrops picta up to 14.4 up to 14.0 500-800 Oceanic Off Cape Town Indian, 65°N-35°5
- .oeramblyops megalops unknown 6 (imm.) Off Cape Town
S Atlantic, Indian,
~ithorax capensis 8.2-13 9- 10 450-550 Oceanic W hole area
Ea: 52 N-40°S
SE Atlantic, W Indian,
,;,;
-crxn~·sis hansoni 8.5-13 10.4- 13 Coastal W coast of Africa
02-352S
k_ Ua5 · o mysis castroi 7.5-9 .0 8.0-9.0 Coastal Off Rio de Janeiro W Atl antic, 35°N-252 S
"W:JL .elia valdiviae 5.5 6.5 Coastal Off Congo
---
511$ fh:nl\ sis atlantica 4.5-5.5 4.0-5.5 10-35 Coastal Off Rio de Janeiro
35°N-23°S
.. 9dopsis acuta 10.8-12.6 10.5-11.5 0-250 Coastal Around Tierra del Fuego
Southern S America,
322S-53°S
">odopsis coelhoi 5.5 5.5-6.0 Coastal Off Rio de Janeiro
" Trop ica l W Atlantic,
•- tdopsis tortonesei 6.0-7.0 5.0-7.5 Coastal Off Rio de janeiro
15°N-25S2
---
4.0-4. 1
up to 12
4.2-4.8
13
I Coastal
Coastal
Off Rio de janeiro
Off Cape Town, False Bay
~
5.2 ---r- Coastal Off Rio de janeiro

_. dopsis rionegrensis 5.6-7.2 5.6 20-50 Coastal San Matias Gulf
sdopsis schultzei 10 9-9.8 0-10 Coastal Off Cape Town, Angola
sdopsis came/ina 6.2 unknown 1-50 Coastal False Bay I
_ dopsis bispinosa 3.5-5.0 unknown 0-10 Coastal O ff Cape Town
12.5 unknown Coastal O ff Cape Tow; ; - - [
1110 Mysidacea
Depth range (m)

Female Male Geographic World wide
and onshore-
Species length length distribution in the geographic
offshore
(mm) (mm) South Atlantic distribution
distribut io n
1
Mysidopsis sim i/is 1 8-11 10 10-50 Coastal Off Cape Town, Ango la
Mysidopsis suedafrikana 4.2 (imm.) 5.0(imm.) 0- 10 Coastal O ff Cape Town
Mys idopsis sankarankuttyi 2.6 unknown Coasta l Off Natal
O ff Cape Lopez, SE Atlanti c, SW Indian,
ILeptomysis capensis 8.0-8.6 7.6-8.4 29-77 Coastal
Ca[:>e Town 02 -35 2 5
ILepto mysis megalops 8.2-10.2 7.2- 11 50-150 Along coasts of Africa
M editerranean, E Atl antic,
402 N -382 S
1
Paraleptomysis dimorpha 5-8 5-6 58-67 Coasta l Off Gabon, Luanda
Metamysidopsis elongata
4.5-8.0 4 .5-7.0 Coastal Off Rio de janeiro
atlantica
Metamysidopsis macaensis unknown Coastal? Off Rio de j aneiro
Metamysidopsis munda 6 6 Coastal Off Rio de j aneiro
302 N -282S
Metamysidopsis swifti 6- 7 5.5-6.5 Coasta l Off Rio de j aneiro
282 N-262S
Arthromysis mage/lanica up to 31 up to 30 20 -30 Coastal Around Tierra del Fuego
1
Mesopodops is wooldridgei 8- 13 7-11 Coastal O ff Namibia, Cape Town

Magell an Str.,
Neomysis monticellii unknown up to 10.4 50-70 Coastal
o ff Puerto Deseado
I Around T. del Fuego, Chi le, Austral SW
1 Neomysis patagona up to 24 up to 19.5 20-40 Coastal Atlantic, 41~s- 55 QS
Fa l kland (=Ma lvi nas) lsi.
IMysidi um gracile 5 Coasta l Off Rio de j aneiro
~emmy>;, mwe<e•nu;
sidetes anomala
6
4
20
3.5
20
~
30 Coastal
Coasta l
202S, 402 W
Magell an Str.
r Southern S America
Magellan Str., around
1 Mysidetes patagonica 16.4-21 16-20 150-300 Coastal Southern S America
Fa lkland (=M alvinas) lsi.
l Mysidetes intermedia 13.8 14.8 100-200 Coastal

Magel lan Str., around
Fal kland (=Ma lvinas) Is_/._
Southern S America,
S02S-53 2 S
A round Falkland Southern S America,
Mysidetes macrops 15- 16 15-16.6 100-500
(Ma lvin as) lsi., S. Georgia 502S-53 2 S
I
A round Falkland Subantarctic, A ntarctic,
Mysidetes brachylepis 17 17-20 100-550 (Ma lvin as) lsi., S. Georgia 502S- 78~s
Around Falkland
Mysidetes crassa 10 10 40-300
(=Malvinas) lsi.
Table 3. Body length, depth range, and geographic distribution of South Atlantic Mysidacea.
Mysidacea 1111
~rder Lophogastrida lies, Petalophthalmidae, Mysidae, Lepidomysidae and

..!:Sllpium consisting of 7 pairs of lamellae. All pie- Stygiomysidae. Two of these, Lepidomysidae and
~ in both sexes, but none of the modified setae in Stygiomysidae, are members of stygian faunas dwell-
- ~- are biramous. Statocyst absent. Generally ing in marine caves.
;::eso- to bathypelagic animals, or epibenthic forms at
;;rderable depths. Includes 3 families. Family Petalophthalmidae
First pereopod without exopod. Carpopropodus of
7
.__,•1y Lop hogastridae endopods of 3rd to 8th pereopods not joined. Female
--="'WDent calcareous or not. Anterior margin of car- with 7 pairs of oostegites. No statocyst on endopod of
~e produced into a broad and short rostrum. Second uropods. Includes 4 genera and about 28 species.
~d developed as a gnathopod; 3rd to 8th pereo- Most species bathypelagic or epibenthic at consider-
ocs normal. Lateral margin of tel son with less than 10 able depths. Only one species has been recorded from
~. Includes 5 genera: Chalaraspidum, Lopho- the South Atlantic.
-~er. Ceratolepis, Paralophogaster and Pseudo-
~aspidum. Chalaraspidum and Lophogaster are Family Mysidae
~nted in the South Atlantic. Exopod of I st pereopod, as a rule, well-developed.
Carpopropodus of endopods of 3rd to 8th pereopods
-..,......·!y Gnathopbausiidae divided into subsegments. Marsupium of 2 or 3 pairs
~;ument strongly calcareous. Rostrum elongate, of oostegites, except in the subfamily Boreomysinae,
~. spear-shaped, denticulate, 3-edged in cross which has 7 pairs of oostegites. Statocyst present on
~ Posterior margin of carapace usually produced endopod ofuropods. Most mysid species (over 90%)
5trong. median, dorsal spine. Abdominal somites belong to this fami ly. Includes 6 subfamilies.
bilobed pleural plates (Fig. 6.7); 6th somite divid-
e: CIO 2 parts by a transverse groove (Fig. 6.7). Subfamily Boreomysinae
E::=pod of uropods 2-segmented. Telson large, with 2 Female with 7 pairs of oostegites. Endopods of 3rd to
x:; '·eels on dorsal surface; lateral margin with spines 8th pereopods with 3- or 4-jointed carpopropodus.
~ed in series; apex with crescent-shaped lobe. Outer margin of antenna! scale naked, tern1inating in
-: and appendages usually red. Deep water forms, a thorn. Exopod of uropods unjointed; outer margin
--:;pelagic or epibenthic. Includes 2 genera, with I or 2 spinules marking end of short naked prox-
-· tphausia and Neognathophausia. imal part. Telson cleft distally; cleft anned with teeth,
without plumose setae. Includes only one genus,
fl::::::::y Eucopiidae Boreomysis, with about 40 species. Most species
~ce very large, membranous. No pleural plate on bathypelagic.
r=O:xninal somites. Antenna! scale with 2 segments.
6:rlopods of 2nd to 4th pereopods relatively short, Subfamily Siriellinae
~:oped as gnathopods; those of 5th to 7th very long, Cornea of eyes globular. Antenna! scale with a thorn
~. subchelate; that of 8th normal. Exopod of uro- marking end of naked outer margin. Endopods of 3rd
:- "'ith distal suture. Only one genus, Eucopia, to 8th pereopods with tuft of long, peculiarly serrated
~ species are poorly defined and difficult to dis- setae surrounding dactylus. Exopod of uropods 2-
-~ Eucopia is most common below l 000 m in jointed; outer margin of basal segment with spines
"1'orld Ocean. Body and appendages red. which are usually restricted to distal part. Apex of tel-
son rounded or truncate, armed medially with 3 small
Sc:..."'Ilier :\1ysida spines, and a pair of plumose setae. Includes 2 tribes,
6:rlopods of 1st and 2nd pereopods modified as 3 genera, and about 65 species. Most species inhabit
~· No branchiae on pereopods. Marsupium coastal regions, but a few are epipelagic, open-ocean
- ~e usually composed of 2 or 3, sometimes 7, forms.
of oostegites. Female pleopods generally rudi-
~· . .\1ale pleopods usually biramous, often one Tribe SirieUini
:::xwe pairs modified into accessory copulatory Male pleopods biramous, multi-jointed except for
s;;:::::c::::nes. Includes 900 species belonging to 4 fami- endopod of 1st pleopod (which is Jacking), with well-
1112 Mysidacea
developed, bilobed, coiled or straight pseudobranchia Key to the South Atlantic genera of Mysidacea
(Fig. 6.23).
Pleopods in both sexes with both rami well
Subfamily Gastrosaccinae developed and natatory, without modified setae.
Female with 2 pairs of oostegites. Both rami of uro- Branchiae present on some or all thoracic limbs.
pods undi vided; outer margin of exopod with 1, 2 or Marsupium composed of 7 pairs of oostegites.
numerous spines, without setae. Endopods of 3rd to Endopod ofuropods without statocyst. (Suborder
8th pereopods with carpopropodus divided into a few Lophogastrida) .. ..................... ........................... 2
or many subjoints. Male 3rd pleopod developed and la Female pleopods reduced to unjointed, single
modified. Telson with apical cleft. Includes 8 genera, lobes. Male pleopods variable. Branchiae absent.
about 85 species, of which 3 genera and 12 species are Marsupium composed of 2-7 pairs of oostegites.
known from the South Atlantic. Endopod of uropods with or without statocyst.
(Suborder Mysida) ........................................... .6
Subfamily Mysinae 2 Integument calcareous. Pleural plate distinct on
Marsupium composed of 2 or 3 pairs of oostegites. abdominal somites ........ ....... ..................... ........ .3
Male pleopods variable. Both rami of uropods undi- 2a Integument not calcareous. No pleural plate on
vided; outer margin of exopod setose throughout its abdominal somites: Family Eucopiidae, Genus
whole length. Most mysid species (more than 90%) Eucopia
belong to this subfamily. Divided into 6 tribes. 3 Rostrum elongate, slender, denticulate, spear-
shaped, 3-edged in cross section. Exopod of uro-
Tribe Erythropini pods 2-jointed. (Family Gnathophausiidae)
Outer margin of antenna! scale usually naked, with ...........................................................................4
terminal thorn, rarely dentate or setose. Male plea- 3a Rostrum produced into broad short plate; anteri-
pods biramous, 1st pair with multi-jointed exopod and or margin smooth, denticulate or tridentate.
unjointed endopod; 2nd to 5th pairs with multi-joint- Exopod of uropods entire or 2-jointed (family
ed exopod and endopod; 4th or 5th pair sometimes Lophogastridae) ............................................... 5
with modified setae. Telson variable. 4 Antenna! scale with suture. Outer margin of
antenna! scale produced anteriorly into long.
Tribe Leptomysini acute spine: genus Gnathophausia
Antenna! scale lanceolate, setose all around. Male ple- 4a Antenna! scale without suture. Distal end of
opods biramous, 1st pair with multi-jointed exopod outer margin of antenna! scale not as long: genus
and unjointed endopod, 2nd to 5th pairs with multi- Neognathophausia
jointed exopod and endopod; exopod of 4th pleopod 5 Integument strongly calcareous. Anterior margin
lengthened with modified setae. Telson variable. of carapace tridentate: genus Lophogaster
Sa Integument less calcified. Anterior margin of
Tribe Mysini carapace sen·ate: genus Chalaraspidum
Antenna! scale usually setose all around. Male plea- 6 Marsupium composed of 7 pairs of oostegites.
pods: 1st and 2nd pairs always rudimentaty; 3rd pair Statocyst absent: family Petalophthalmidae.
rudimentary or biramous; 4th pair always biramous, genus Hanseuomysis
endopod unjointed, exopod more or less elongate and 6a Marsupium composed of 7 or less pairs of oos-
divided, usually into several segments. Telson vari- tegites. Statocyst present in endopod of uropods
able. (family Mysidae) ...............................................7
7 Marsupium composed of 7 pairs of oostegites:
Tribe Heteromysini subfamily Boreomysinae, genus Boreomysis
Antenna! scale setose all around. Endopod of 3rd per- 7a Marsupium composed of2 or 3 pairs of oostegites
eopod sometimes enlarged and robust. Pleopods rudi- .............................................................................. 8
mentary in both sexes, except long 3rd pleopod of 8 Exopod of uropods 2-jointed: subfamily
male Harmelinel/a. Penis long, cylindrical. Telson Siriellinae, genus Siriella
with or without apical cleft. 8a Exopod ofuropods unjointed ............................ 9
Mysidacea 1113
Exopod of uropods with outer margin armed with 21a Telson more than 1.5 times as long as broad
spines but no setae (subfamily Gastrosaccinae) -·-··-·--·-·- ·-·-·-··-··-··---·-·-·-·--·--·--·····-·-···--·-·······-·- ·-·23
----·-·······-····---····--··-·-··-·-·--···-···-····-····---···-·-····-··-·-10 22 Eyes depressed dorso-ventrally. Telson trape-
Exopod of uropods with outer margin armed zoidal, with 2 or 3 pairs of spines and a pair of
with setae but no spines (subfamily Mysinae) plumose setae on distal margin: genus Erythrops
--·-··-·- ·· ·· ·-·-·-·· ·- ······- ·- ·-·-··· ····-· ···· ···-···- ·· ·-·· ···· -···-·12 22a Eyes globular. Telson triangular, with a pair of
lC Exopod of male 3rd pleop od slightly longer than slender spines and 1 or 2 plumose setae on apex:
endopod: genus Anchialina genus Heteroerythrops
lOa Exopod of male 3rd p leopod much longer than 23 Telson with a pair of apical plumose setae: genus
endopod ·--·-·-·-·-·-··-··-··-·-·-·---·-·····--···---·-·-·----·····11 M eterythrops
It Exopod of male 3rd pleopod modified terminal- 23a Telson without a pair of apical p lumose setae:
ly as complicated copulatory organ. Female 1st genus Kate1ythrops
p leopod uniramous: genus Bowmaniella 24 Last thoracic somite extremely lengthened.
:a Exopod of male 3rd p leopod styliform. Female Lateral margin oftelson distally armed with sev-
lst pleopod biramous: genus Gastrosaccus eral pairs of spines: genus Longithorax
Eye plate-like, without cornea, lacking distinct 24a Last thoracic somite normaL Lateral margin of
stalk. ................ ............ .... ...... ·-·-·-·-·- ...... ........... 13 telson distally densely armed with about 10 slen-
Ca Eye stalked, w ith cornea more or less developed der spines: genus Hyperamblyops
·-······ ·-·-·-·-·--·-······-·--·-·-·--·-··-···-·--·-·-·--····--··-··-·-·-14 25 All pleopods of male biramous; exopod of 4th
Eyes fused in single p late: genus Pseudomma pleopods with modified seta(e) (tribe Lepto-
.=a Ey es not fused in single p late: genus mysini) ........................... ...... ...... ....... ... ....... .... -26
Amblyopsoides 25a At least 1st and 2nd p leopods of male uniramous
Cornea of eye divided into anterior and lateral -·-·-·· ······ ····-·-·-·-·- ····-·-···-·-·-·--··-········-·-·-···-········.34
portions; lateral cornea not developed to special- 26 Telson with apical cleft -·-·-···-········-·-·- ···· ··· ·· ···-27
ized lens __ ._._._._ ... __ ._ .. ._._._. _... _._ .. .. .. _._._._. _.. ..... .. _.15 26a Telson withou t apical cleft ................... ...... .... .31
Cornea of eye not divided into two portions _.. . 18 27 Second segment of endopod of maxilla greatly
Rostrum produced frontally into long spear-like expanded dista lly: genus Afromysis
process extending beyond distal end of eyes 27a Secon d segment of endopod of maxilla not
--- ·--····-····-· ····-···-·-···--·-···- ·-·-·-·-·-·-··· ···-·· ···-·- ····· ·· 16 expanded distally , longer than broad ......... ..... 28
~?. Rostrum not extendi ng beyond distal end of eyes 28 Antenna! scale extremely long and narrow.
---·· ··········· ·········-·-····-···---·-·-·-·--·········-·-·-·-·--·····-17 Carpus of endopod of 8th pereopod with 10-15
Exopod of 1st pereopod reduced to small protu- characteristic hooks on one side and 4-6 strong
berance: genus Caesaromysis spines on the other side: genus Brasilomysis
~ Exopod of 1st pereopod developed into natatory 28a Carpus of endopod of 8th pereopod with setae
organ: genus Echinomysis but no hooks or spines ..... ............................... .29
Outer margin of antenna! scale naked, ending in 29 Endopods of 3rd to 8th pereopods with carpo-
a thorn: genus Euchaetomera propodus divided into 2 subsegments: genus
_- :=. Outer margin of antenna! scale setose, lacking Nouvelia
distal thorn: genus Euchaetomeropsis 29a Endopods of 3rd to 8th pereopods w ith carpo-
_-\ntenna l scale vestigial or absent: genus propodus divided into 3 or more subsegments
A.rachnomysis ········-·-·-·-···-· ······--··--·-·-·----·····-··--·- ···-···· ·- -·-··--·- ·30
_...._ _-\ntennal scale well developed ...... ...... ............ l9 30 Endopod of uropods with row of spines along
_-\ntennal scale with a thorn marking end of inner margi n, some of these sp ines very large
naked outer margin ..... .. ....... ... ... ...... ...... .......... 20 and conspicuous. C left of telson without plu-
_-\ntennal scale setose all around -·-·--··--······ ·-·-·25 mose setae and spines: genus Promysis
Telson with lateral margin naked .................... 21 30a Spines on endopod ofuropods less conspicuous.
I elson with lateral margin armed with spines Telson cleft, if present, with spines: genus
-----· ···-········-···-··· ·-- ···--·-·- ·-··--· ···-·····-·- ·-·-· ··· ··-· -·-·24 Mysidopsis (part)
Telson shorter than or as long as broad .... ...... 22 31 Cornea of eye with enlarged facets in posterior
part: genus Paraleptomysis
1114 Mysidacea
3la Cornea of eye without specialized facets ........ 32 Antenna! scale not jointed, with acutely pointed
32 Maxilla without exopod: genus Metamysidopsis apex; outer margin serrate. 6th abdominal somite
32a Maxilla with exopod .......... ................... ........... 33 not divided into 2 sections. Uropods with both
33 Exopod of 4th pleopod of male with a modified rami not jointed. Telson oblong-triangular, with
seta at apex of distalmost segment: genus narrowly truncate apex and a couple of dorsal spi-
Mysidopsis (part) nose keels. Ref.: Fage (194 1), as Chalaraspis
33a Exopod of 4th pleopod of male with modified alata; B irstein and Tchindonova (1962).
setae on distal 3 segments: genus Leptomysis
34 At least 2nd pair of male pleopods rudimentary Genus Lophogaster
and uniramous. 4th or both 3rd and 4th pleopods Integument firm. Anterior margin of carapace triden-
of male elongate and modified. Endopod of 3rd tate.
pereopods normal (tribe Mysini) .................... 35 • Lophogaster challengeri Fage, 1941 (Fig. 6.2).
34a All pleopods in both sexes rudimentary. Endopod Carapace with microscopic tubercles, especially in
of 3rd pereopod with or without enlarged distal central dorsal area; median tooth of frontal plate
segments (tribe Heteromysini) ...................... .. .38 short, barely exceeding lateral teeth; pair of post-
35 Telson with deep apical cleft, armed with a pair orbital spines lacking. Lobe of inner distal margin
of plumose setae: genus Arthromysis of antennular peduncle evenly rounded at anterior
35a Apex of tel son entire, incised or emarginate .... 36 end, smooth, with a small bristle. Apex of anten-
36 Body generally slender. Telson with large spine na! scale curved inwardly. Ref.: Fage (1942), 0. S.
marking end of lateral margin, distal margin Tattersall (1955).
semicircular; lateral and distal margins with • Lophogaster spitwsus Ortmann, 1906 (Fig. 6.3).
spines. 3rd and 4th pleopods of male biramous: Carapace without microscopic tubercles; median
36a
genus Mesopodopsis
Telson without spine marking end oflateral mar-
process of frontal plate very elongate. Antenna!
scale very elongate, rather lanceolate; apex straight:
• •
gin. 4th pleopod of male biramous ........... ....... 37 outer margin with 9-10 teeth. Ref.: Fage (1942).
37 Exopod of 4th pleopod of male 3- or 4-segment- • Lophogaster muranoi Fukuoka, Hoffmeyer and
ed, ultimate and penultimate segments with a Vinas (Fig. 6.4; the description of this species was 1
stout seta. Antenna! scale with rounded apex: in press at the time of submission of the present .. J
genus Mysidium work). Carapace with microscopic tubercles ante-
37a Exopod of 4th pleopod of male 2-segmented; rodorsally; median process of frontal plate rela-
dista l segment with 2 equal or subequal terminal tively long; a pair of post-orbital spines present.
setae. Antenna! scale with acutely pointed apex: Lobe of inner distal margin of antennular peduncle
genus Neomysis rounded with slight median emargination; margin
38 3rd thoracic endopod strongly built: genus finely serrate, 2 unequal setae present. Antenna!
Heteromysis scale with outer margin slightly convex, with 5
38a 3rd thoracic endopod almost equal to following denticles; apex straight.
limbs: genus Mysidetes
Family Gnathophausiidae
Identification of the genera and species Genus Gllathophausia - I
.
For each of the species treated one or more references Antenna! scale divided into 2 parts by articulation:
("Ref.") providing detailed descriptions and illustra- outer margin ending in strong spine.
tions are included. • Gnathophausia gracilis Willemoes-Suhm, 1875
Family Lophogastridae
(Fig. 6.5). First 2 abdominal somites with 1 and 2
(respectively) prominent denticles on dorsal
-
Genus Chalaraspidum median line. Posterolateral angles of carapace with
Integument soft. Anterior margin of carapace very spines. Ref.: Sars (1885), Fage (194 1).
broad, truncate and serrate. • Gnathophausia zoea Willemoes-Suhm, 1873
Chalaraspidum alatum Willemoes-Suhm, 1874 (Fig. 6.6). Supra-ocular spines present. First 2
(Fig. 6.1 ). Anterior margin of rostrum serrate. abdominal somites without prominent denticles.
Posterolateral wmgs of carapace rounded. Posterolateral angles of carapace rounded; poste-
Mysidacea 1115
rodorsal spine very long, denticulate. Ref.: Sars antennule enormously thickened and densely hirsute.
1885), Fage (1941). Antenna! scale lanceolate; outer margin with spines
among setae. Male pleopods biramous, without mod-
Genus Neognathophausia ified setae. Exopod of uropods 2-jointed.
A=ltennal scale without articulation; outer margin end- Htmsenomysis falklalldica 0. S. Tattersall, 1955
:; in small spine. (F ig. 6.14). Carapace with well defined upturned
• Seognathophausia ingens (Dohm, 1870) (Fig. ri m. Antenna) scale 6.5 times as long as broad;
6. 7). Rostrum rather short, broad at base, indis- outer margin with 9-l 0 strong spines among setae.
tinctly denticulate. Supra-orbital spines wanting. Exopod of uropods with proximal joint anned
_-\ ntennal scale subovate, apex truncate, distal half with 9 spines among setae on di stal 3/4 of outer
of its outer edge minutely serrate. The largest margin. Sides oftelson almost parallel in proximal
mysid species. Ref.: Sars (1885); Fage ( 194 1); half, then narrowing; lateral margin with dense
Clarke (196 1), as Gnathophausia ingens. row of spines arranged in series on distal4/5; apex
• Seognathophausia gigas (Willemoes-Suhm, with 3 spines, middle one short; no plumose setae
1873) (Fig. 6.8). Rostrum slender, distinctly den- on apex. Ref.: 0. S. Tattersall (1955).
ticulate. Supra-orbital spines distinct but small.
.-\.ntennal scale somewhat tapering toward apex, Family Mysidae
outer margin with 4 strong teeth. Ref.: Sars Subfam ily Boreomysinae
. 0~ 1885); Fage ( 1941), as Gnathophausia gigas .
Lateral margin of te1son clearly arranged alter-
f:!:Ilily E ucopiidae nately with a longer spine and several shorter
Genus E ucopia spines. E ndopod of 2nd pereopod with subche1ate
• Eucopia sculpticauda Faxon, 1893 (Fig. 6.9). termination. Telson cleft with rhomboidal space
~lore robustly built than other Eucopia species. • Boreomysis bispi~tosa 0. S. Tattersall, 1955 (Fig.
Dorsal surface of telson ornamented with a series 6. 15). Anterior margin of carapace rounded, wi th
- of honeycomb ridges. Ref.: Tattersall and
Tattersall (1951 ).
extremely short upturned projection. Endopod of
2nd pereopod with 2 spines on carpopropodus.
• Eucopia australis Dana, 1852 (Fig. 6.1 0). Distal Ref.: 0. S. Tattersall ( 1955).
segment of antenna! scale asymmetrical. Distal
5egment of exopod of uropods about as long as Lateral margin of telson not arranged alternately
broad. Lateral margin of telson arranged with a with a longer spine and several shorter spines.
long spine and alternately several short and equal Endopod of 2nd pereopod without subchelate ter-
spines. Ref.: Fage ( 1942), Tattersall (1955). mination. Te1son cleft without rhomboidal space
• Eucopia uflguiculata (W.-Suhm, 1875) (Fig. • Boreomysis acuminata 0. S. Tattersall, 1955
o.l l ). Closely allied to E. australis. Distal seg- (Fig. 6.16). Rostrum with narrow, very sharply
ment of antenna! scale nearly sym metrical. Distal pointed apex. Eyestalk expanded distally, forming
segment of exopod of m·opods clearly wider than large triangu la r projection; cornea small, restricted
:tS length. Ref.: Tattersall and Tattersall (1951). to distal outer area of eyestalk. Ref.: 0 . S.
• Eucopia grimaldii Nouvel, 1942 (Fig. 6.12) . Tattersall (1955).
Dorsal surface of telson n ot ornamented. Spines • Boreomysis atlantica Nouvel, 1942 (Fig. 6.17).
on lateral margins of telson not a rranged regular- Anterior margin of carapace broadly rounded with
ly. Ref.: Nouvel (1943). median small triangular projection. Cornea large,
Eucopia linguicauda 0. S. Tattersall, 1955 (Fig. somewhat flattened; eyestalks with small papilla
b.1 3). Telson linguiform with broadly rounded on dorsal surface. Ref.: Nouvel (1943).
:rpex; lateral margin armed on distal half with • Boreomysis illigi 0. S. Tattersall, 1955 (Fig.
many small spines. Ref.: 0. S. Tattersall ( 1955). 6.1 8). Rostrum triangular, apex acutely pointed,
lateral margins convex. Eyes large; papilla very
~'Enily Petalophtha1midae long, extending forward beyond anterior margin
6e:::u.s Hansellomysis of cornea. Ref.: 0. S. Tattersall ( 1955).
E~ reduced to small plate with projecting distal cor- • Boreomysis insolita 0. S. Tattersall, 1955 (Fig.
~= \isual elements absent. Outer flagellum of male 6.19). Anterior margin of carapace evenly round-
1116 Mysidacea
ed and uptilted to form delicate, transparent, Ref.: Sars (1 885), as Anchialus typicus; 0. S.
almost vertical rim. Eyes re latively large, globu- Tattersall (1955).
lar; ocular papilla small, inconspicuous. Ref.: 0. • Anchialina typica (Kroyer, 1861) (Fig. 6.26).
S. Tattersall (1955). Frontal margin of carapace produced into short
trapezoidal plate with rounded angles, plate bent
Subfamily Siriellinae strongly downwards, as a result of which true
anterior margin is invisible from above. Exopod of
Outer margin of antenna! scale naked. 4th or 3rd male 3rd pleopod with L-shaped, naked setae on
and 4th pleopods of male with modified setae several middle segments. Ref.: ouvel ( 1971 ).
.
• Siriella chierchiae Coifrnann, 1937 (Fig. 6.20).
Male 3rd and 4th pleopods with modified setae on Genus Bowmaniella
ultimate and penultimate joints of both endopod
and exopod. Ref.: W. M. Tattersall ( 1951),
Male 2nd pleopod with multi-jointed exopod and
unsegmented endopod. Male 3rd pleopod with exo-
-
Brattegard (I 970a). pod ending in complex copulatory organ; endopod
• Siriella woljfi 0. S. Tattersall, 1961 (Fig. 6.2 1). unjointed, with 2 curved spines at tip. All female ple-
Male 4th pleopod with terminal setae modified. opods biramous. Telson with 2 spines on each apex.
Ref.: 0. S. Tattersall (1961). • Bowmaniella brasiliensis Bacescu, 1968 (Fig.
6.27). Posterior margin of carapace with lateral
Outer margin of antenna! scale naked. Male pleo- lobes not reflexed. Endopod of uropods with 4-7
pods without modified setae spines on inner margin; exopod with 13-18 spines
• Siriella dayi 0. S. Tattersall, 1952 (Fig. 6.22). on outer margin. Telson shallowly cleft (1:0.08);
Pseudobranc hiae of male 2nd to 4th pleopods C- lateral margin armed with 6-7 small spines. Ref.:
shaped, not spirally coiled. Lateral margin of tel- Bacescu (1968a).
son with 4 strong spines in basal part. Ref.: 0. S. • Bowmaniella dissimilis (Coifmann, 1937) (Fig.
Tattersall ( 1952). 6.28). Posterior margin of carapace with lateral
• Siriella melloi da Silva, 1974 (Fig. 6.23). lobes not reflexed. Endopod of uropods with 5-6
Pseudobranchia of male 4th pleopod nearly long spines along inner margin; exopod with 18- J
straight. Exopod of uropod shorter than endopod, 20 spines on outer margin. Telson shallowly cleft
with 3 spines in distal part of outer margin of prox- ( 1:0.11 ); lateral margin with 6-8 fairly evenly 9
imal segment. Ref.: da Silva (1974). spaced spines. Ref.: Coifrnann ( 1937). l
• Siriella thompsoni (Milne-Edwards, 1837) (Fig. • Bowmaniella atlantica (da Si Iva, 1971) (Fig. II!
6.24). Pseudobranchiae of male 2nd to 4th pleo- 6.29). Posterior margin of carapace with lateral
pods spirally coiled. Exopod of uropods shorter lobes reflexed. Inner margin of endopod of uro-
than endopod. Lateral margin of telson with 2 pods with 3 spines, 2 in proximal portion and 1 in
strong spines in basal part. Ref.: Coifmann (1937), middle portion; exopod with 11 -16 spines on outer
Ii (1964). margin. Telson with lateral margin armed with 8
spines including 2 longer spines at apex. Distal
Subfamily Gastrosaccinae cleft of telson with 10 spines on each lateral mar-
Genus Anchialina gin. Ref.: da Silva (1970a), as Gastrosaccus bras-
Posterior margin of carapace straight or feebly emar- iliensis; da Silva ( 197la).
ginate. Antenna! sympod with strong spiniform pro- • Bowmaniel/a inarticulata da Silva, 1972 (Fig.
cess furnished with subsidiary spinules. Endopod of 6.30). Posterior margin of carapace with lateral
male 2nd pereopod with merus more or less expanded lobes reflexed. Endopod of uropods with 3 long
and produced posteriorly. Exopod of male 3rd pleo- spines and 9 minute spines on inner margin; exo-
pod longer than endopod, with modified setae and/or pod with 14 spines on outer margin. Telson deep-
characteristic protuberances. ly cleft ( 1:0.4); lateral margin armed with 13 short • G
• Anchialina truncata (G. 0. Sars, 1884) (Fig. spines. Ref.: da Silva (1972).
6.25). Rostrum abruptly truncate at tip. Exopod of • Bowmanie/la recifensis da Silva, 197 1 (Fig.
3rd pleopod with short and straight setae on sever- 6.31). Posterior margin of carapace with lateral
al distal segments. Probably a coastal species. lobes reflexed. Endopod of uropods with 4 spines 3
Mysidacea 1117
)S.. and 7 minute spines on inner margin; exopod w ith wi th 8 long, slender, somewhat irregularly spaced
14- 16 spines on outer margin. Telson deeply cleft spines and with a row of 6 small, graduated, close-
( I :0.42); lateral margin of telson w ith about 14 ly set spines at extreme proximal end of inner mar-
short, unequal spines. Ref.: da Silva (197 lb). gin. Telson less than 3 times as long as broad; lat-
eral margin armed with 7 strong, somewhat
Genus Gastrosaccus irregular spines, term inal one very long and
_ laJe 2nd pleopod with multi-segmented exopod and strong; cleft less than 115 oftelson length. Ref.: 0 .
~opod. Male 3rd pleopod with exopod styliform, S. Tattersall (1958).
~ding in 2 simple setae; endopod multi-segmented, Gastrosaccus olivae Bacescu, 1970 (Fig. 6.36).
uo curved spines at tip. Female pleopods: I st pair Posterior marg in of carapace with a well devel-
.mamous, 2nd to 5th pairs uniramous. oped reflexed lobe on each s ide. Exopod of male
Gastrosaccus brevifissura 0. S. Tattersall, 1952 3rd pleopod 3 times as long as endopod, distal seg-
(Fig. 6.32). Posterior margin of carapace with flat, ment with 2 unequal strong setae terminally.
rectangular, backwardly directed lobe, median Endopod of uropods with 6 long irregularly
- - :ns... portion between lobes transverse. Male 2nd pleo- spaced spines. Telson less than 3 ti mes as long as
pod with 9-segmented exopod twice as long as 7- broad; lateral marg in armed with 6 large, irregu-
segmented endopod. Exopod of male 3rd pleopod larly spaced spines, of which the terminal one is
\\;th distal segment armed with 2 short subtermi- the longest, distal cleft 1/9 of telson length. This
nal spines and 2 spiniform terminal setae. Telson species has previously been known as G. sanctus.
about twice as long as broad; lateral margin w ith 7 Ref.: Bacescu (1970).
spines, apical one particularly long; apical cleft
,·ery shallow. Ref: 0. S. Tattersall (1 952). Tribe Erythropini
Gastrosaccus kempi W. M. Tattersall, 1922 (Fig. Genus Pseudomma
6.33). Posterior margin of carapace with small, Eyes reduced to a single, broad, flattened plate, with
inwardly directed lobe on each side. Male 2nd ple- median notch on anterior margin.
opod w ith 8-segmented exopod 1.5 times as long • Pseudomma magellanensis 0. S. Tattersall, 1955
as 8-segmented endopod. Endopod of ma le 3rd (Fig. 6.37). Eyeplate with serration on antero-lat-
pleopod with distal segment anned with strong eral corners; each half with a marked ridge along
sera on outer margin and 2 terminal unequal setae. middle line. Antenna! scale 3 times as long as
Telson 3 times as long as broad; lateral marg in broad; apical lobe slightly longer than large tooth
\\;th 9 large spines, spaces between third to termi- at end of unarmed outer margin. Telson 1.5 times
nal spines occupied with 3 to 6 small, regularly as long as broad; lateral marg in with 8-10 small
graduated spinules. Ref. : 0. S. Tattersall ( 1952), spines; apex broad, truncate, ve1y slightly emargi-
35 Gastrosaccus gordonae. nate, with 3 pairs of slender spines. Ref: 0. S.
Gastrosaccus namibensis Wooldridge and Tattersall (1 955).
~lclachlan, 1987 (Fig. 6.34). Postero-dorsal mar- • Pseudomma calmani 0 . S. Tattersall, 1955 (Fig.
gin of carapace deeply emarginate, bearing 8 slen- 6.38) Eyeplate without serrations on antero-lateral
der spine-like filaments. Male 2nd pleopod with corners. Antenna! scale 3.5 times as long as broad;
!0-segmented exopod slightly longer than 10-seg- apical lobe occupying 115 of scale length. Telson
mented endopod. Endopod of uropods with 9 slen- 1.5 times as long as broad; lateral margin with 5-6
der. irregularly spaced spines. Telson nearl y 2.5 small spines; apex arched, with 3 pairs of long
ames as long as broad; lateral margin with 6 or 7 spines. Ref.: 0. S. Tattersall (1955).
spines including terminal one, which is not mark- Pseudomma minutum 0. S. Tattersall, 1955
edly longer than others. Ref. : Wooldridge and (Fig. 6.39). Eyeplate w ithout serration on antero-
_ fclachlan, (1987). latera l corners. Antenna! scale ovate; apical lobe
• Gastrosaccus psammodytes 0 . S. Tattersall, 1958 occupying more than half of scale length. Telson
Fig. 6.35). Adult male undescribed. Anterior por- triangular, as long as broad; lateral margin with
::on of rostrum curved downward and, as a result, 2-3 very small spines; apex narrowly rounded,
i::l dorsal view appearing truncate and very short. with a pair of long spines. Ref. : 0. S. Tattersall
Endopod of uropods armed along inner margin (1955).
1118 Mysidacea
Genus Amblyopsoides as long as broad. Rather common in the South

Eyes reduced to 2 quadrangular, immovable plates Atlantic, including the Southern Ocean. Ref.: Illig
without visual elements and definite eyestalks. (1930).
Amblyopsoides obtusa 0. S. Tattersall, 1955 (Fig. • Euchaetomera glyphidophtlwlmica Tllig, 1906
6.40). Antenna! scale with apical lobe very long, (Fig. 6.46). Rostrum small, narrow, with acute tip.
occupying nearly 112 of its length. Telson trape- Postero-lateral comea largely expanded laterally.
zoidal; apex 113 as broad as at base, with 2 pairs of larger than anterior one. Antenna! scale 4 times as
long spines, a pair of minute spines and a pair of long as broad. Lateral margins of telson naked.
median plumose setae; lateral margin with dense Ref.: Illig ( 1930), Murano (1977).
row of spines. Ref.: 0. S. Tattersall (1955). Euchaetomera intermedia Nouvel, 1942 (Fig.
6.47). Rostrum narrow, with acute apex. Postero-
Genus Caesaromysis lateral cornea somewhat expanded laterally, equal
Cornea of eyes divided into anterior and lateral por- to anterior one in size. Antenna! scale more than
tions. Rostrum produced frontally into long spear-like 4.5 times as long as broad. Lateral margins of tel-
process extending beyond distal end of eyes. Exopod son unarmed. Ref.: Nouvel (1943).
of 1st pereopod reduced to small protuberance.
Caesaromysis hispida Ottmann, 1893 (Fig. 6.4 1). Genus Euchaetomeropsis
Carapace and abdominal somites covered with Allied to Euchaetomera, except for antermal scale
many long spines. Antenna! scale in male small, without distal spine on outer margin.
lanceolate without spine-like process at outer dis- • Euchaetomeropsis merolepis (Illig, 1908) (Fig.
tal end; in female, reduced to short conical pro- 6.48). Postero-lateral cornea slightly expanded lat-
cess. Ref.: Illig (1930), Murano (1977). erally. Antenna! scale setose all around . Telson
with lateral margin unarmed; apex truncate, with 1
Genus Echinomysis spine at each comer. Ref.: Illig (1930), Casano\"a
Allied to Caesaromysis, except for 1st pereopod with (1968).
multi-segmented exopod.
• Echinomysis chuni Illig, 1905 (Fig. 6.42). Genus Arachnomysis
Antenna! scale lanceolate, without thorn terminat- Anterior margin of carapace and posterior margin of
ing naked outer margin. Lateral margin of telson abdominal somites with acute spines. Antenna without
armed with spines. Ref.: Illig (1930). scale. Eyes conical. Telson very small; lateral margins
smooth; apex nanow, with a pair of very small spines.
Genus Euchaetomera Arachnomysis leuckarti Chun, 1887 (Fig. 6.49).
Cornea of eyes divided into anterior and lateral por- Number of facets across the largest diameter of
tions. Rostrum not extending beyond distal end of eye 10-14. Ref.: Murano (1977).
eyes. Antenna! scale with dista l spine at end of naked • Arachnomysis megalops Zimmer, 19 14 (Fig.
o uter margin. 6.50). Number of facets across the largest diame-
Euchaetomera tenuis G. 0. Sars, 1884 (Fig. ter of eye 18-25 . Ref.: Murano (1977).
6.43). Postero-lateral cornea not expanded lateral-
ly, much smaller than anterior one. Antenna! scale Genus E1ythrops
neatly 5 times as long as broad. Telson with later- Telson trapezoidal, shorter than or as long as broad.
al margin naked. Ref.: Ii (1964). with lateral margins unarmed.
• Euchaetomera typica G. 0. Sars, 1884 (Fig. 6.44). • E rythrops africana 0. S. Tattersall, 1955 (Fig.
Rostrum narrow, with acutely pointed apex. 6.5 1). Rostrum low, triangular, with pointed apex.
Postero-lateral cornea expanded laterally. Antenna! Antenna! scale 8 times as long as broad. Telson
scale 3 times as long as broad. Telson slightly short- shorter than broad, apex with 2 pairs of spines.
er than broad; lateral margin with 5-6 small slender Ref.: 0. S. Tattersall (1955).
spines; apex a little wider than 1/3 of broadest patt, • Erythrops bidentata Nouvel, 1973 (Fig. 6.52).
with 1 slender spine at each comer. Ref.: Ii (1964). Rostrum low, triangular, with rou nded apex.
Euchaetomera zurstrasseni (Illig, 1906) (Fig. Antenna! scale nearly 5 times as long as broad.
6.45). Allied to E. typica. Antenna! scale 5 times Telson slightly shorter than broad; apex with 2
Mysidacea 1119
pairs of spines. Endopod of uropods with inner • Hyperamblyops megalops (0. S. Tattersall, 1955)
margin ftnely serrated. Ref.: Nouvel ( 1973). (Fig. 6.58). Antenna! scale without terminal thorn
at end of naked outer margin. Telson linguiform,
Genus Heteroerythrops twice as long as broad; lateral margin armed along
:-~Lson triangular, as long as broad, with latera l mar- distal 1/3 with 11 small spines which are length-
:-s naked. Antenna! scale without terminal thorn at ened distally; apex narrow, with a pair oflong slen-
~ of unarmed part of outer margin. der spines and 3 plumose setae. Ref.: 0. S.
• Heteroerythrops purpura 0. S. Tattersall, 1955 Tattersall (1955), as Gibberythrops megalops.
F ig. 6.53). Rostrum short, triangular, with bluntly
rounded apex. Eyes large, globular, set widely Genus Longithorax
apart. Antenna! scale narrow, shotter than anten- Last thoracic somite extremely long, occupying about
na! pedunc le; inner margi n and distal l/3 of outer 1/3 of cephalothorax. Eyes normal or somewhat
margin with plumose setae. Apex of tel son with a reduced, with ftnger-like process on eyestalks. Telson
pair of long spines and a pair of feeble, median triangular, with spines on apex and distal part oflater-
plumose setae. Ref.: 0. S. Tattersall (1955). al margins.
• Longithorax capensis Zimmer, 1914 (Fig. 6.59).
6e:::ns Katerythrops Finger-like process on eyestalk lengthened, curv-
~.:son triangular, longer than wide; lateral margin ing outward in fro nt of cornea. Antenna! scale 3
00; apex with 2 pairs of spines, without a pair of times as long as broad; apex extending far beyond
~se setae. thorn at end of naked outer margin. Telson trian-
• Kar.erythrops brasiliana Bacescu, 1984. (Fig. 6.54) gular, longer than wide; apex narrow, with a pair
_-\ntennal scale 3 times longer than broad. Telson of spines; lateral margin with 1 spine near distal
slightly longer than wide; apex with inner spines 7 end. Ref.: 0. S. Tattersall (1955).
nmes longer than outer. Ref.: Bacescu (1984).
• lf.trterythrops oceanae Holt and Tattersall , 1905 Tribe Leptomysini
Fig. 6.55). Antenna! scale short, narrow, some- Genus Afromysis
\Ohat outwardly curved, more than 7 times as long Distal segment of endopod of maxilla strongly
~ broad. Telson 1.5 times as long as wide. Ref.: expanded distally, armed with spines, in addition to
:-anersall and Tattersall (195 1), Ii (1964). setae, on distal margin.
• Kmerythrops resimora 0. S. Tattersall, 1955 (Fig. • Afromysis hansoni Zimmer, 19 16 (Fig. 6.60).
-;,.56). Anterior margin of rostrum almost straight Outer margin of distal segment of endopod of
:!:xi rurned upward; lateral margins also bent maxilla extremely long. Exopod of male 4th pleo-
-=9ward. Antenna! scale small, slender, arcuate, pod more than twice as long as endopod, its last
~g:htly longer than antenna! peduncle. Telson 1.5 segment very long, more than 4 times as long as
ones as long as broad; apex with inner spines preceding segment. Lateral margin of telson with
~ce as long as outer. Ref.: 0. S. Tattersall (1955). spines throughout. Ref.: Zimmer (19 16), 0 . S.
Tattersall (1955).
' Jfeterythrops
"IT!:-'«!.triangu lar, longer than wide, with lateral mar- Genus Brasilomysis
=- cled: apex narrow, with 2 pairs of spines and a Carpopropodus of endo pod of 8th pereopod with sim-
of plumose setae. ple spines on outer margin and flagellate curved
• _'J.nerythrops picta Holt and Tattersall, 1905 (Fig. spines on inner margin. Eyestalk, antennule, and
- 5- ). Antenna! scale large; outer margin coasely antenna extremely lengthened.
~e. Eyes globular and big. Ref.: Tattersall and • Brasilomysis castroi Bacescu, 1968 (Fig. 6.61 ).
-:!llersall (195 1), Ii (1964). Antenna! scale nearly styliform, 14 times as long
as broad. Telson tongue-shaped; lateral margin
- Hyperamblyops naked along proximal 1/3, then armed with a
~- convex-lens-shaped in dorsal view, facets
__ series of small spines of the usual type, gradually
~rly developed. Lateral margin oftelson dense- becoming very peculiar. Ref.: Bacescu (1968c),
~ \\ith about l 0 slender spines in distal part. Almeida Prado (1 974).
1120 Mysidacea
Genus Nouvelia long as broad. Telson linguiform, 1.5-1. 7 times as

Endpods of 3rd to 8th pereopods with 2-segmented
carpopropodus. Telson with distal cleft armed with
long as broad; lateral margin with about 20 spines
along whole length; apex with 3 pairs of long
•
1
spines on margins and with a pair of plumose setae at spines. Ref.: Biicescu (1968b), Almeida Prado
anterior end. Exopod of male 4th pleopod with 2-3 (1974).
modified setae. • Mysidopsis tortonesei Biicescu, 1968 (Fig. 6.66).
• Nouvelia valdiviae (Illig, 1906) (Fig. 6.62). Antenna! scale lanceolate with acutely pointed - J
Rostrum low, triangular. Antenna! scale lanceo- apex, 8 times as long as broad. Telson linguiform,
late, 6 times as long as broad. Endopod of uropods 1.8 times as long as broad; lateral margin with 3
with about 40 spines along inner ventral margin. spines in basal part, followed by short unarmed
Telson 1.6 times as long as broad; lateral margin section, and then with 14-20 spines; apex rounded.
naked on proximal half except for one spine near with 2 pairs of spines. Ref.: Biicescu (1968b).
base, distal half with 8 short spines; each apex of Almeida Prado (1974).
distal lobes with 2 spines, outer one more than • Mysidopsis juniae da Silva, 1979 (Fig. 6.67).
twice as long as inner; apical cleft 116 of telson Antenna! scale lanceolate with rounded apex, 6.5
length, with 8 small spines on each margin. Ref.: times as long as broad. Telson 1.2 times as long as
Illig (1930), as Doxomysis valdiviae. broad; lateral margins naked; apex truncate with
median projection, armed with 1 spine at each cor-
Genus Promysis ner. Ref.: da Silva (1979).
Telson cleft at apex, margin of cleft smooth, pair of • Mysidopsis major (Zimmer, 1912) (Fig. 6.68).
plumose setae present or absent at its anterior end. Rostrum very long, with pointed apex. Eyes
Antepenultimate and penultimate joints of endopod of extending laterally beyond width of carapace.
male 4th pleopod with a single strong spinose seta Antenna! scale linear with narrowly rounded apex.
each, ultimate joint terminating in 2 short spinose Telson twice as long as broad, with broadly round-
setae. Endopod of uropods with row of spines along ed apex, armed throughout with small equal
inner margin, some of them very large and prominent. spines. Ref.: Zimmer (1912), as Paramysidopsis
• Promysis atlantica W. M. Tattersall, 1923 (Fig. major; 0. S. Tattersall (1955).
6.63). Endopod of uropods slightly shorter than Mysidopsis scintilae dos Reis and da Silva, 1987
exopod; inner margin armed along its length with (Fig. 6.69). Rostrum short, triangular. Eyes not
spines, distal 3 of which are very long, stout and extending laterally beyond width of carapace.
inwardly curved. Telson 1.5-1.9 times as long as Antenna! scale lanceolate with rounded apex, 3.7
broad; lateral margin with about 25 spines on dis- times as long as broad. Telson linguiform with
tal 2/3; pair of plumose setae lacking. Ref.: W. M. broadly rounded apex, with 25-35 spines along
Tattersall (1923), Clarke (1956), Almeida Prado margin. Ref.: dos Reis and da Silva (1987).
(1974). • Mysidopsis rionegrensis Hoffmeyer, 1993 (Fig.
6. 70). Antenna! scale lanceolate with rounded
Genus Mysidopsis apex, 5 times as long as broad. Telson linguiform.
Exopod of male 4th pleopod ending in single stout, 1.5 times as long as broad; lateral margin with 8-
long, spinose seta. Telson usually short, usually trun- 11 small spines; apex rounded, with about 14
cate or rounded, rarely split at apex, without apical longer spines. Ref.: Hoffineyer (1993).
plumose setae; lateral margins usually spinose, some- • Mysidopsis schultzei (Zimmer, 1912) (Fig. 6. 71 ).
times unarmed. Antenna! scale lanceolate with rounded apex; 8
• Mysidopsis acuta Hansen, 1913 (Fig. 6.64). times as long as broad. Telson elongate lingui-
Antenna! scale lanceolate with acute tip, 5.5 times form, more than twice as long as broad, lateral and
as long as broad. Telson linguiform, twice as long as apical margins with about 90 subequal spines.
broad; lateral margin with about 30 short subequal Ref.: Zimmer (1912), as Paramysidopsis schult-
spines; apex with a pair of long spines. Ref.: Hansen zei; 0. S. Tattersall (1955).
(1913), 0. S. Tattersall (1955), Holmquist (1957). • Mysidopsis camelina 0. S. Tattersall, 1955 (Fig.
• Mysidopsis coelhoi Biicescu, 1968 (Fig. 6.65). 6.72). Body robust, sigmoid in lateral view.
Antenna! scale lanceolate with acute tip, 8 times as Carapace with 2 prominent protuberances along
Mysidacea 1121
rnid-dorsalline postetior to cervical sulcus, anteri- pleopod with long, modified setae on distal 3 seg-
or one considerably larger than posterior. ments. Telson entire, lateral margin with numerous
Antenna! scale oval, 2.5 times as long as broad. longer and shorter spines.
Telson linguiform; lateral margin wi th 9-10 small • L eptomysis cap ens is Illig, 1906 (Fig. 6. 78).
spines; apex with 2 pairs of short spines. Ref.: 0. Whole integument densely covered with fme short
S. Tattersall(l 955). bristles. Ref.: Illig ( 1930), P illai ( 1973).
• Jfysidopsis bispinosa 0. S. Tattersall, 1969 (Fig. • Leptomysis megalops Zimmer, 191 5 (Fig. 6.79)
6. 73). Antenna! scale oval, twice as long as broad; Rostrum triangular, with bluntly pointed apex.
outer distal angle of antenna! sympod produced Antenna! scale bluntly pointed distally, 9 tim es as
into conspicuous, strong, finger-like process with long as broad. Margin of telson densely atm ed
rounded apex extending to more than l/3 of scale with many spines arranged rather irregularly. Ref.:
length. Telson small, triangul ar, 1.5 times as long Lagardere and Nouvel ( 1980).
as broad, lateral margins unarmed; apex with a
pair of spines; small emargination between spines. Genus Paraleptomysis
Ref.: 0. S. Tattersall (1969). Allied to Leptomysis except for comea of eye with
• Jfysidopsis eremita 0 . S. Tattersall, 1962 (Fig. enlarged facets in posterior pottion.
6.74). Carapace with well developed hump on • Paraleptomysis dimorpha Wittmann, 1986 (Fig.
median dorsal region just behind cervical sulcus. 6.80). Antenna! scale lanceolate with rounded
Antenna! scale oval, twice as long as broad; outer apex, 7 times as long as broad. Telson twice as
distal angle of antenna! sympod with strong spine. long as broad, expanded distally; lateral margi n
Telson linguifotm, slightly less than twice as long with 3-4 spines in proximal half and 14-20 spines
as broad; whole lateral and apical margins with 19 in distal half, these spines consisting of larger and
evenly spaced slender spines; apex with a pair of smaller ones, tending to increase in length distally;
long slender spines. Ref.: 0. S. Tattersall (1962) . apex ttuncate, wi th 3 pairs of long spines, longer in
• J.fysidopsis simi/is (Zimmer, 19 12) (Fig. 6.75). females than in males. Ref.: Wittmann (1986) .
.-\ntennal scale lanceo late with rounded apex,
more than 3 times as long as broad. Telson lingui- Genus Metamysidopsis
form except for pointed apex, nearly tw ice as long Allied to Mysidopsis, except for maxilla without exo-
as broad; lateral margin with about 30 spines pod. Telson entire, linguiform.
rbroughout its length; apex with 2 spines conspic- • Metamysidopsis elongata atlan tica Bacescu,
uously longer than the lateral ones. Ref.: Zimmer 1968 (Fig. 6.8 1). Antenna! scale 5 times as long as
19 12), as Paramysidopsis simi/is; 0. S. Tattersall broad. Telson 1.5 times as long as broad, distal
1952), as Leptomysis tattersalli. half much narrower than proximal half, with 30-46
• .Uysidopsis suedafrikana 0. S. Tattersall, 1969 spines. Endopod of uropods with about 35 large
Fig. 6.76). A ntenna! scale oval with rounded apex, and small sp ines along inner margin. Ref.:
~.25 times as long as broad. Telson trapezoidal, 1.5 Biicescu (l968b), Almeida Prado (1974).
times as long as broad; lateral margin nearly • Metamy sidopsis macaensis da Silva, 1970 (Fig.
straight, unarmed; apex bluntly rounded, with 3 6.82). Antenna! scale 3.5 ti mes as long as broad.
pairs of spines. Ref.: 0 . S. Tattersall ( 1969). Telson as long as broad, triangular with truncate
• J.fysidopsis sankarankuttyi Bacescu, 1984 (Fig. apex, slight constriction near distal end; apical and
6. 77). Telson linguiform, slightly longer than lateral margi ns with 26 spines, of which the ones
\\ide; lateral margin with 2 pairs of short spines on the distal lobe are very strong. Endopod of uro-
near distal end; apex with 4 large su bequal spines. pods wi th 19 spines along inner margin. Ref.: da
Ref.: Biicescu (1 984). Silva (1970b).
• Metamysidopsis munda (Zimmer, 1918) (F ig.
(.ieoaus Leptomysis 6.83). Antenna! scale narrowly lanceolate, 5-6
Cc:::J.ea of eye without enlarged lateral facets in peste- times as long as broad. Telson broadly triangular,
portion. Distal segment of endopod of max illa about 1.3 times as long as broad; distal 1/3 of lat-
ger than broad. Endopods of 3rd to 8th pereopods eral margin with about 18 short acute spines.
-=b 3-segmented carpopropodus. Exopod of male 4th Endopod of uropods with about 46 spines along
1122 Mysidacea
inner margin. Ref.: Zimmer (19 18), da Costa al; apex entire, narrowly rounded or truncate; lateral
(1964). margin with spines throughout its length.
• Metamysidopsis swijti Bacescu, 1969 (Fig. 6.84). • Neomysis monticellii Colosi, 1924 (Fig. 6.87).
Antenna! scale narrowly lanceolate, about 5 times Rosh·urn short; apex narrowly rounded and bent
as long as broad. Telson entire, broadly ling ui- downward. Telson elongate triangular, 2.5 times
form, with rounded apex, 1.3 times as long as as long as broad; lateral margin with about 28
broad at base, lateral margin naked, ending distal- short, subequal spines; apex with 2 pairs of spines.
ly in small tooth; apex with 11 -15 shott spines on Exopod of male 4th pleopod terminating in 2
each side of longer mid-apical pair. Ref.: Bacescu unequal setae. Ref.: Colosi (1924), 0. S. Tattersall
( 1969), Brattegard (1970a). ( 1955).
• Neomysis patagona Zimmer, 1907 (Fig. 6.88).
Tribe Mysini Rostrum relatively long, divided into 2 lobes by
Genus Arthromysis conspicuous incision. Telson somewhat less than 3
Eyes extremely long and narrow. Male 3rd, 4th and times as long as broad; lateral margin with a large
5th pleopods biramous. Telson deeply cleft; cleft with number of small spines; apex very narrow, with 2
spinules on margin and pair of plumose setae at ante- pairs of spines. Ref.: 0. S. Tattersall (1955).
rior end.
• A rthromysis magellanica (Cunningham, 1871) Genus Mysidium
(Fig. 6.85). Antenna! scale long, setose all around. Male 4th pleopod biramous; endopod small, unj oint-
3rd pleopod with multi-jointed exopod and ed; exopod 3 or 4-jointed, with strong seta on each of
unjointed endopod; 4th pleopod with extremely distal 2 segments. Telson quadrangular; apex evenly
long, multi-jointed exopod which is armed with rounded, transverse, emarginate, o r incised; proximal
very long modified setae on distal segments, and half of lateral margin unarmed, distal half and apex
unjointed endopod; 5th pleopod with both rami densely armed with small spines.
multi-jointed. Lateral margin of telson with row of • Mysidium g racile (Dana, 1852) (F ig. 6. 89).
subequal spines. Ref.: 0. S. Tattersall ( 1955). Antenna! scale 4 times as long as broad. Telson
with apex slightly emarginate; lateral margins
Genus Mesopodopsis slightly concave; number of spines of lateral and
Eyestalks elongate. Anterior margin of carapace with apical margins is about 50. Exopod of 4th male
a pair of post-orbital spines. Distal joint of male pleopod with distal 3 segments of equal length.
antennular peduncle with accessory flagellum, in Ref.: da Costa ( 1964), Brattegard ( 1969).
addition to male lobe. Male 3rd pleopod biramous,
endopod unjointed, exopod 2 to 3-jointed. Male 4th Tribe Heteromysini
pleopod biramous, endopod 2 to 3-jointed, exopod Genus Heteromysis
elongate, 3 (5?)-jointed, dista l joint with 2 modified Eyes with cornea occupying full width of distal part of
setae. Each lateral margin of telson ending in a stro ng eyestalk. Endopod of 3rd pereopod with some distal
spine; distal margin between strong spines produced segments enlarged. Abdomen without well-developed
into semicircle. pleura. Telson with apical cleft lined with denticles.
• Mesopodopsis wooldridgei Wittmann, 1992 (Fig. • Heteromysis mureseanui Bacescu, 1986 (Fig.
6.86). Eyestalks 3-4 times as long as cornea. 6.90). Rostrum short, obtusely triangular. Eyes
Lateral margin of telson with 5-7 spines; apical large; sta lks ending antero-medially at edge.
lobe with 3 1-39 spines. Ref.: Wittmann ( 1992). Antenna! scale not extending to distal end of
antennular peduncle. Endopod of 3rd pereopod
Genus Neomysis not so swollen, terminating in strong claw; carpo-
Antenna! scale long and narrow, tenninating in ac ute- propodus with 2 dentiform flagellated spines and 4
ly pointed tip, setose on both margins. Endopods of plumose setae on inner margin. Endopod of uro-
3rd to 8th pereopods with carpopropodus divided into pods without spines. Telson slightly longer than
many subj oints. Exopod of male 4th pleopod elon- broad, with sinus armed with about 20 spines;
gate, 2-jointed, terminating in 2 stout barbed setae; each apex with 2 spines; lateral margin armed with
endopod unjointed. Telson subtriangular or trapezoid- only 4 spines on distal 1/3. Ref. : Bacescu (1986).
Mysidacea 1123
T,pmll A
Chalaraspidium alatum C, posterolateral
angle
part Y1'
Lophogaster challengeri
erA'
(--..) lobe
AP[I] -~~.
3rd segment
C, posterolateral
angle
T
Lophogaster spinosus
® Gnathophausia gracilis
9ws
AN [I]
Gnathophausia zoea
1124 Mysidacea
transverse
(5
biB-0
bilobed
pleural
plate
tl- Neognathophausia
ingens
AN[I]
Ows
Neognathophausia gigas
AN[r]
T
UR[ r]
Eucopia sculpticauda
UR[r] and T Eucopia unguiculata
ab
UR [r] and T
UR[r] and T
Eucopia australis Eucopia grimaldii
Mysidacea 1125
ab
UR[I) and T
<:j? PR3[r)
Eucopia linguicauda
UR[r)
and T
d(adu ll) ANN[!)
AN[r)
Hansenomysis falklandica
<:j?(adult) ab
PR2 En
UR[r) and T
~ Boreomysis bispinosa
<:j?(adult) ab
Boreomysis acuminata
C (anterio r part)
AN[I)
AN[r) UR[r) and T
$ Boreomysis illigi Boreomysis atlantica T

1126 Mysidacea
)_)(adu lt) ab
AN[rJ UR[rJ and T
En
d PL3 (dista l part) cf PL4 (dista l part)
Siriella chierchiae
Boreomysis insolita
PL3 PL4
UR [r) and T
cfPL3 (pseudobranchial lobe)

Siriella dayi
~ AN[I] U R[I] and T

~ Siriella wolffi
cfPL3
pseudobranch ia l
lobe
UR [I] and T
Siriella melloi ab (left half)

Mysidacea 11 27
Ca m
.,.·< ~R [ I]Ex T OPL3 O PL3

AN[I) (Ex, distal part)
pseudobranchia l
Anchialina truncata
dab lobe
AN [I]
Sirie/la thompsoni
d'ws
UR[r]
<D Bowmaniella brasiliensis
T O PL3
(Ex, distal part)
UR[I] En
s Anchialina typica
OPL3
ab and C
Bowmaniella
atlantica Bowmanielfa dissimilis
1128 Mysidacea
~ \. C (dorsal pm) :'
I UR[r) En
T
T (left posterior part)
dab (right half)
C (dorsal pm)
T
UR[r)
8 Bowmaniella inarticulata
Gastrosaccus brevifissura
O PL3
AN[r)
T
OPL2
T
Gastrosaccus kempi
Gastrosaccus namibensis
Mysidacea 1129
EY am ofC C (dorsal pm,

lateral v iew)
~
T
C (dorsal pm, dorsal view)
Cpm
Gastrosaccus psammodytes
UR[r) En
T
Gastrosaccus olivae
J ~r - - - eyeplate UR[I] and T
,- dab l
Pseudomma magallanensis
UR and T
UR[r) and T
AN [I]
Pseudomma minutum Pseudomma calmani
1130 Mysidacea
T OAN [I]
Amblyopsoides obtusa Caesaromysis hispida
postero-lateral
cornea
AN[I] AN [r]
\)
<j?ws T
Echinomysis chuni <j? ws T
~ fuchaeto.mera
~ tenws fuchaetomera typica
T (posterior end)
AN[I]
<D ' fuchaetomera zurstrasseni fuchaetomera glyphidophtha/mica

Mysidacea 1131
oostero-
la;c=ral
cornea
AN[r]
AS[r]
Euchaetomera intermedia
Euchaetomeropsis merolepis
dab
EY [I]
Arachnomysis leuckarti
<j>ws
• Arachnomysis megalops
Oab
AN[r] AS[I) T and UR En
Erythrops bidentata
Erythrops africana
1132 Mysidacea
AN[r]
UR[r] and T
• Heteroerythrops purpura e T
Katerythrops
brasiliana
Katerythrops oceanae
AS[ Ij T
• dab
Meterythrops picta
AN[I] T
8 Katerythrops resimora
dab
d (immature) ab
Hyperamblyops megaiops Longithorax capensis

Mysidacea 1133
AN[r]
MX OPL4
Afromysis hansoni
-----
T
Brasilomysis castroi
dws
<~!) Nouvelia valdiviae
UR [r] and T
Promysis atlantica
dab
oPL4
UR[r] and T T UR [I] En 0 PL4 AN[r]

Mysidopsis acuta Mysidopsis coelhoi
1134 Mysidacea
AN[I]
Oab
AN[I]
UR[I] En
Mysidopsis tortonesei
T 0 ab and C
v T
Mysidopsis juniae
UR[I] and T
T Oab
Mysidopsis major AN[r] T
8 Mysidopsis scintilae
T
AN[r]
UR]r]
AN[I]
• Mysidopsis rionegrensis Mysidopsis schultzei
Mysidacea 1135
AN[r]
Mysidopsis came/ina
<( ws UR[I]
MD and MP <D Mysidopsis bispinosa
Mysidopsis eremita
UR[rl En T
Mysidopsis simi/is
AN[r]
UR[r] En
ab (left half)
T
Mysidopsis suedafrikana Mysidopsis sankarankuttyi

1136 Mysidacea
AN[I ]
Leptomysis capensis UR[I] En T

_ _.
OPL4
UR(r] En
8 Leptomysis megalops
AN[r]
Paraleptomysis dimorpha
UR[I]
or
MX (distal end)
Metamysidopsis
• ANIII elongata atlantica
AN[I]
UR[I] En T
Metamysidopsis macaensis Metamysidopsis munda

Mysidacea 1137
UR[I) En
AN(r]
Metamysidopsis swifti
"--
OPLS
OPL3
LjJT
cD Arthromysis magellanica
ANN [r]
0 Mesopodopsis wooldridgei
cfpb
cf pb
(latera l view)
a ~
OPL4 (distal end)
Neomysis monticellii
AN[I)
T AN[I]
T
e UR[r]
Neomysis patagona
~
~
Mysidium gracile
1138 Mysidacea
U R[I] and T
Heteromysis mureseanui
ANI II
<JP Mysidetes anomala
UR[r] and T
AN[r]
CD Mysidetes intermedia Mysidetes macrops
n
AS[r]
\ I
pb T (d ista
.. l end)
\j\;'
T
Myside tes crassa Mysidetes brachylepis
Mysidacea 1139
Genus Mysidetes Telson 2.5 times as long as broad, cleft for 1/3 of
::es large; cornea globular w ith short broad eye- telson length. Ref.: W. M. Tattersall (1923).
;::o'ks. Endopod of3rd p ereopod w ithout enlarged dis- • Mysidetes crassa Hansen, 191 3 (Fig. 6.96).
_: segments. Endopod ofuropods usually with row of Rostrum short, with acute tip. Antenna) scale
~es on inner margin. Telson with apical cleft lined about 2.5 times as long as broad. Telson 1.5 times
.til denticles. as long as broad; lateral margin naked in proximal
• Jfysidetes anomala 0. S. Tattersall, 1955 (Fig. half and with 9 spines in distal half; apical cleft
t>.9 1). Rostrum acutely pointed. Antenna! scale 0 .25 as long as telson , with 28 spines in proximal
very slender, more than 7 times as long as broad. half. Ref.: Hansen ( 1913).
Telson less than twice as long as broad; apical
cleft less than 1/4 of telson length . Ref.: 0. S.
Tattersa ll (1955). Suggested readings
• Jfysidetes patagonica 0. S. Tattersall, 1955 (F ig.
6.92). Rostrum short, triangu lar, with acutely Brattegard T. 1969. Marine biological investigations
pointed apex. Antenna! scale more than 6 times as in the Bahamas. I 0. Mysidacea from shallow
:ong as broad. Telson less than twice as long as water in the Bahamas and southern Florida. Part I .
l:>road, cleft for 1/3 of telson length. Ref. : 0. S. Sarsia, 39:17-106.
Tattersall (1955). Brattegard T. 1970. Marine biological investigations
• .\Jysidetes intermedia 0. S. Tattersall, 1955 (Fig. in the Bahamas. 11. Mysidacea from shallow
:3.93). Rostrum short, triangular, with narrowly water in the Bahamas and southern Florida. Part 2.
rounded apex. Eyes large, set close together, not Sarsia, 41: 1-36.
extending laterall y beyond width of carapace. Brattegard T . 1970. Mysidacea from shallow water in
.\ntennal scale 4 times as long as broad. Telson the Caribbean Sea. Sarsia, 43:111- 154.
""ice as long as broad, cleft fo r 0.3 of its le ngth. Brattegard T. 1973 . Mysidacea from shallow water on
Ref.: 0. S. Tattersall ( 1955). the Caribbean coast of Colombia. Sarsia, 54: 1-66.
• .\Jysidetes macrops 0. S. Tattersall, 1955 (F ig. Brattegard T. 1974. M ysidacea from shallow water on
"-9~). Rostrum very sho1t, with pointed apex. the Caribbean coast of Panama. Sarsia, 57:87-1 08.
Eyes very large, extending laterally beyond width Brattegard T. 1974. Additional M ysidacea from shal-
:· carapace. Antenna! scale 4 times as long as low water on the Caribbean coast of Colombia .
.:road; antenna! sympod with 2 spines on outer Sarsia, 57:47-86.
6.:.-r.al angle. Telson twice as long as broad, cleft The six references above are major contributions to
·or 0.38 of its length. Ref.: 0. S. Tattersall (1955). the mysidacean fauna of the Atlantic coasts of
• JJ_rsidetes brachylepis W. M. Tattersall, 1923 Central America, including many species which
Fig. 6.95). Angle of rostrum acute, tip bluntly may quite possibly occur along the Atlantic coasts
:ronded. Antenna! scale 3 times as long as broad. of South America.
IL Illustrations of the species.

~ons used : ab : anterior part of body; am : anterior margin; AN: antenna; ANN : antennule; AP: antennular peduncle; AS: antenna!
C carapace; En: cndopod; Ex: exopod; EY: eye; F: femal e; Ill: left; M : male; MD: mandible; MP: mandibular palp; MX: maxilla; pb:
:a pan of body; PL: pleopod; pm : posterior margin; PR: pereopod; Jrl: right; R: rostrum; T: telson; UR: uropod; ws: whole specimen.
- -=-- ;.oorces: after Bacescu ( 1968a): 27, 28; after Bacescu ( 1968b): 65, 66, 81; after Baccscu (1968c): 61; after B~cesc u (1969): 83 (except
:0:,. 84: after Biicescu (1970): 36; after Bacescu ( 1984): 54, 77; after Bacescu (1986): 90; after Brattegard ( 1969): 89; after Brattegard
~0: after da Silva ( 1970b ): 82; after da Silva (1971 b): 29, 31 ; after da Silva ( 1972): 30; after da Silva ( 1974): 23; after da Silva ( 1979):
- ~ dos Reis and da Si lva (1987): 69; after Fage ( 1942): 3; after Hansen (1913): 96; after Hoffmeyer ( 1993): 70; after li ( 1964): 24, 26
~male ab, UR[l) En, T, 43 , 44, 55; after Illig (1930): 42, 45 62; after Murano (1977): 41 , 46,48-50, 57 AS[ I); after Nouvel (1943): 17,
- .0=- '\ou\'el (1971): 26 male PL3; after Nouvel (1973): 52; after 0. S. Tattersall ( 1952): 22, 32 (modified), 33 (modified); a fter 0 . S.
- 1955): I, 2, I0- 16, 18, 19, 37-40, 5 1, 53 , 56, 58, 64, 71, 72, 75, 85, 87, 88, 9 1-94; after 0 . S. Tattersall (1 958): 35; after 0 . S. Tattersall
:L after 0. S. Tattersall (1962): 74; after 0. S. Tattersal l ( 1969): 73, 76; after Pillai ( 1973): 78; after Sars ( 1885): 5, 6, 7 (modified), 8
- . :!5 (modified); after Tattersall and Tattersall (1951 ): 9, ll ; after W. M. Tattersall (1923): 63, 95; after Wittmann ( 1986): 80; after
1992): 86; after Wooldridge and McLachlan ( 1987): 34; after Zimmer ( 19 12): 68 (modified); after Z immer ( 19 14): 79; after Zimmer
:~. 60: after Zimmer (1918): 83 female ab (modified).
11 40 Mysidacea
Page L. 1941. Mysidacea Lophogastrida. I. The Muller H.-G. 1993. World catalogue and bibliography
Carlsberg Foundation's oceanographical expedition of the recent Mysidacea. Lab. Trop. Ecosystems
round the world 1928- 1930 and previous "Dana"- Res. Jnf. Serv., Wetzlar, 1-49 1. A list of9 79 mysid
expeditions under the leadership of the late species in 149 genera, including taxonomic refer-
Professor Johannes Schmidt. Dana Rep., 19:1-52. ences, geographic distribution, depth range, habi-
Page L. 1942. Mysidacea Lophogash·ida. II. The tat preferences, and a bibliography of 3490 titles.
Carlsberg Foundation's oceanographical expedi- Nouvel H. 1943. Mysidaces provenant des campagnes
tion rou nd the world 1928-30 and p revious du Prince A lbert Ier de Monaco. Res. Camp. Sci.
"Dana"-expeditions under the leadership of the late Accomplies sur son Yacht par 1e Prince Albert Ier.
Professor Johannes Schmidt. Dana Rep., 23: l -67. Prince Souverian de Monaco, I 05: 1-128. A mono-
These two works are detailed reports on members of graph based on mysid materials collected in the
the suborder Lophogastrida collected by the North Atlantic Ocean.
Danish RJV "Dana", including materials from the Sars G.O. 1885. Report on the Schizopoda collected
South Atlantic Ocean (in French). by H. M. S. Challenger during the years 1873-76.
Illig G. 1930. Die Schizopoden der Deutschen Rep. Sci. Res. Voyage H.M.S. Challenger 1873-
Tiefsee-Expedition. Wiss. Erg. Deutsch. T iefsee- 76, 13: 1-228. A monograph on the Mysidacea and
Exp. Dampfer "Valdivia" 1898-1 899, 22:397-625. Euphausiacea collected from the World Ocean by
A monograph on the Mysidacea and Euphausiacea the British RJV "Challenger ".
collected by the German RJV "Valdivia", includ- Tattersall O.S. 1955 . Mysidacea. Discovery Rep ..
ing mysid materials from the South Atlantic Ocean 28: 1-190. The most important monograph for the
(in German). South Atlantic Ocean, including many species
Mauchline J. 1980. The bio logy of mysids and fi'om the area treated here, collected mainly by the
euphausiids. Part I. The biology of mysids. RJV "Discovery".
Advances Mar. Bioi., 18: 1-369. A useful summary Tattersall W.M. 195 1. A review of the Mysidacea of
of mysid biology; includes keys to genera, and the United States Nationa l Museum. Smithsonian
information on reproduction, vertical distribution Inst., U.S. Nat. Mus. Bull. , 20 I, I-X, pp. 1-292. A
and migration, food, feeding, chemical composi- monograph on the mysids deposited at the U. S.
tion, internal anatomy, physiology, behaviour, National Museum, including South Atlantic spe-
population dynamics, geographic distribution, cies.
predators, parasites, and economic importance. Tattersall W.M., Tattersall O.S. 195 1. British Mysida-
Mauchline J., Murano M. 1977. World list of the cea. Ray Soc. Pub!. , London, pp. l-460. A mono-
Mys idacea, Crustacea. J. Tokyo Univ. Fish., graph on the Mysidacea around the British Isles.
64:39-88. A list of 120 genera and 765 species, including an historical account, general morpholo-
with latitudinal ranges, body lengths, geographic gy, and bionomics, in addition to detailed descrip-
region, habitat, and sources ofgood descriptions. tions and illustrations of76 species.
Amphipoda
Georgyi Vinogradov
Dedicated to the memory of my mother, the marine and rarely salps. Even species with free-living adults
biologist Nina Vinogradova often deposit their young on gelatinous animals
(Laval, 1980; Madin and Harbison, 1977). Symbiotic
hyperiideans often act as ectoparasites or commensals,
Introduction but they sometimes penetrate the bodies of j elly-fish,
and may eat their mesogloea and gonads (Hyperia,
The Amphipoda (Peracarida, Malacostraca, Crustacea) Lestrigonus). The free-living predator Themisto uses
are a crustacean order that has successfully colonized je lly-fish as a substrate for moulting and copulation
the pelagic ocean. Their success can be explai ned to a (Sheader, 1977). Phronimidae are famous for pre-
great extent by their high diversity - over 6,000 spe- paring barrel-like houses from the bodies of salps and
cies, exhibiting a large variety of life-forms and occu- the nectophores of calycophore siphonophores by eat-
pying diverse biotopes. Most amphipods are benthic, ing parts of them away. They fmd shelter and rear
but the approximately 400 species in the pelagic their young in such gelatinous houses.
domain inhabit all depths and latitudes of the world
ocean. Some hyperi ids, among them Scinidae, Scypho-
lanceola, Streetsia, Parapronoe crustulum, and prob-
The order Amphipoda includes 4 suborders: ab ly some others (as well as the gammarid
Gammaridea (encompassing the majority of species), Cyphocaris challengeri) are luminescent (Bowman,
Hyperiidea, Lemodipoda (=Caprellidea) and 1967; Vinogradov et a/., 1982).
lngolfiellidea. These are easily distinguished by such
features as the morphology of mouth parts, peculiar- In tropical regions, high concentrations of hyperii-
ities of segment fusion, number of gills, etc. But phy- deans rarely occur. However, even in the tropics their
letic relationships within the Amphipoda are poorly biomass can reach 5 g. m·3 in isolated swarms of
known, such that, for example, some branches within Anchylomera, Phronima or Primno (Vinogradov and
&be gammarid phylogenetic tree may rank with the Rudjakov, 1971). On the other hand, the diversity of
hyperiids or caprellids, which are treated as "subor- tropical hyperiids is high, although precise data on
ders" (see Bousfield and Shih, 1994). species compositions and geographic variability are
still scarce. Such lack of i11formation may result from
.\11 recent pelagic amphipods o riginated from benthic long-standing poor taxonomy in a number of families
~cestors, but representatives of the various suborders (e. g., Hyperiidae, Phronimidae and some others), and
aave occupied the pelagic realm at various times and the difficulty of assessing faunal distributions based
:n various ways. The most ancient immigrants are the on small standard plankton nets (see Methods below).
Hyperiidea. They have lost their connection with the
:x>rtom completely, and none of their known 260 spe- In cold-water regions, hyperiid species diversity is
=ies is benthic (thoug h in rare cases some individuals fairly low, but the ir biomass may be very high, to the
.::13\·e non-obligatory secondary contacts with benthic extent that whales and seals occasionally feed on
:ommunities). Hyperiid amphipods are significant hyperiid swarms. There are even records of ships hav-
:omponents of epi- and mesopelagic communities. ln ing to stop because swarms of Themisto clogged the
:::;e meso- and the macroplankton they are subsidiary engine water-intake pipes.
~ abundan ce only to copepods, euphausiids, chaetog-
c:uhs. and gelatinous animals. As opposed to the hyperiids, representatives of the
Gammaridea are much newer to the pelagic realm:
Hyperiids occupy various pelagic ecological niches. only about 150 of their over 5,000 marine species are
....pproxirnately half of them are parasites on such strictly pelagic, and the majority of these have close
~latinou s zooplankton as medusae, siphonophores, relatives on the sea bottom. On the other hand, only a
~h Atlantic Zooplankron.
<..-.Ied by D. Boltovskoy, pp. 1141-1240
:999 Backhuys Publishers, Leiden, The Netherlands
1142 Amphipoda
few benthic gammarids (wood-burrowing, and some itats. It seems that their predecessors lived on soft
highly specialized epiphytic animals) fail to rise deep-sea mud, which requires thin, long legs for effi-
above the bottom. In many benthic gammarid species cient locomotion. This morphologica l trait, wb1ch
a considerable part of the population (sometimes proved so successful and occurs repeatedly and inde-
males only) may rise above the sea-floor at night. pendently in vari ous groups, allowed the gammarids
Such short excursions require no special anatomical to leave the bottom. Other groups seem to have adapt-
adaptation, and the morphology of the animals is fu lly ed to the plankton through different pathways (see
adapted to their benthic habitat. These migrations usu- Vinogradov, 1995).
ally occur in littoral regions, and these crustaceans are
only seldom carried by currents into the open ocean, As mentioned above, strictly pelagic garnmarids are not
in which case they are bound to perish. Such animals usually abundant, but inhabit all latitudes and depths of
are not included in this review. the ocean. As a rule, planktonic gammarids are signifi-
cantly less numerous in samples than hyperiids.
On the other hand, a totally different assemblage of
upward-migrating gammarids occurs in the near-bot- Various shallow-water, and even intertidal, benthic
tom layer. These are strictly carnivorous animals, pro- gammarids sometimes may be found in the open sea
vided with special mouth parts, moving over the sea sitting on drifting sea-weed. A few species are asso-
bottom in search of carrion or easy live prey. The hab- ciated with the pelagic seaweed Sargassum, but they
its of these bentho-pelagic scavengers, widely distri- are not truly planktonic animals, and therefore are not
buted at all depths, and including dozens of genera, included in this review.
are somewhat analogous to those of terrestrial scav-
enging insects. This group formed the basis for ga m- Caprellids are generally benthic animals, inhabiting
marid expansion into the pelagic zone. Some bentho- littoral areas with sea-weeds and hydroids. A fe w
pelagic gammarids still stay close to the bottom, caprellids have limited swimming capabilities, wrig-
whereas others rise thousands of meters up into gling li ke bloodworms. One benthopelagic species.
mesopelagic depths and may be found in plankton Phtisica marina, often ventures into the water-column
samples (Eurythenes, Paralicella, etc.). and can therefore be considered hemi-planktonic.
Another large group of caprellid amphipods is the
Benthic amphipods that venture high in the water co- cyamids, or whale-lice, which dwell on the skin of
lumn consume not only animal remains from the bot- w hales. Whale-lice are highly-specialized parasites
tom , but also feed in the pelagic zone, consuming that have lost the ability to swim. Although they occur
dead organisms or preying on live ones (Smith and in the pelagic domain, they are not planktonic animals
Baldwin, 1984). This leads directly to the next, strict- and therefore are not included in this review.
ly pelagic, gammarid life-forms (like Cyphocaris) .
Some of the latter (the lys ianassoids) became either The last amphipod suborder, the Ingolfiellidea, have
parasites of gelatinous animals (Danaella, Thoriella), no pelagic representatives and are therefore also omit-
or ectoparasites of large planktonic crustaceans (my- ted here.
sids and decapods) (Paracyphocaris), and fish (espe-
cially sharks; Thrischizostoma). The last g roup has Among other crustaceans, Amphipoda are most close-
not been reported from the South Atlantic. ly related to the Isopoda or Mysidacea (see Bousfield
and Shih, 1994, for a detailed discussion of phyloge-
Gammarid amphipods with a hypertrophic lateral netic relationships of the Amphipoda).
shield (Stegocephalidae) also arrived at their present
pelag ic milieu through a near-bottom phase.
Methods
Plankton ic gammarids with very long appendages,
such as those in the fam ilies Pardaliscidae and Pelagic amphipods can be collected with any plankton
Vitjazianidae (not reported from the south Atlantic net, but collections not specifically designed for them
Ocean), as well as some E usiridae, seem to be basical- may be poorly representative. Planktonic amphipods
ly best adaptated to deep, sluggish-water pelagic hab- usually have well-developed eyes, are active swim-
Amphipoda 1143
mers, and can therefore dodge standard nets quite effi- hyperiid suborder Physocephalata mouth parts are
ciently. Thus, for most species, large nets w ith mouth taxonomically useful only at the level of families and
openings above 1 m2 are required for quantitative fau- some small species of Hyperiidae.
nal analysis. Large high-speed nets (Bongo, Isaacs-
Kidd midwater trawls with a mouth area about 5 m2, Dissections are performed with dissecting needles in a
etc.) have made it possible to acquire representative drop of water with glycerine on a glass plate or crys-
materials, yielding a more complete picture of species tallising dish under the binocular microscope. The li-
compositions, including rare species (Shulenberger, quid is renewed at suitable intervals. For a detailed
1977; G. V inogradov, 1991 , 1993b). The mesh of examination, the appendages from one side of the
these nets, notably Isaacs-Kidd trawls, shou ld prefer- body should be removed (taking care not to disturb
ably be small enough (e.g., 5 mrn in the Samyshev- their order). Mouth parts are usually removed one by
.-\seev modification) to retain the animals efficiently; one, and it may be easier to isolate the entire buccal
larger mesh-openings, while inadequate for quantita- mass and dissect it separate ly, starting with the max-
tive purposes, can still retrieve qualitatively interesting illipeds (see General Morphology below). If perma-
samples. Despite avoidance problems, even smaller nent slides are not necessary, the appendages may be
plankton nets may collect enough specimens to furnish placed in a cavity slide in g lycerine and covered w ith
3 general picture of the di versity at a given site. a cover slip for subsequent analysis with a high-pow-
ered compound microscope.
~t should be stressed that some amphipods often
swarm, with the result that a single tow may strongly Glycerine-gelatine slides are usually the most useful.
over- or underestimate their overall abundance. In The med ium is prepared by soaking 7 g of gelatine in
.Jddition, some hyperiids may enter a sampler w ith 42 cm3 of disti lled water for 2-3 hours, then adding 50
.10sts (gelatinous planktonic animals), thus resulting g of glycerine and 0.5 g of crystalline phenol (carbo-
:n unrealistically high numbers of amphipods. lic acid) or, preferably, thymol. The mixture is
wam1ed in a water bath wi th continuous stirring, after
Bentho-pelagic gammarid scavengers may be collect- which it is filtered and cooled, turning it into a dense
ed by means of baited traps. Of course trap samples gelatinous substance. Slides are prepared by placing
.!re only qualitative. amphipod appendages in a small drop of glycerine on
the glass slide. A tiny amount of glycerine-gelatine is
:\nimals must be preserved in 70% alcohol or in 4% dropped on the cover slip and heated until it melts
~e red formaldehyde. (care should be taken to avoid letting the mixture boil;
if this happens the process must be started over). The
:dentification of amphipods usually requires dissec- cover slip is then turned and placed on the material on
:mn, or at least manipulation under a binocular micro- the slide. During this procedure the appendages may
s.cope. Before detailed examination and dissection of move. If some of them overlap or trap air bubbles,
~e specimen, the characters wh ich can be destroyed carefully pressing the cover slip with a needle under
~- subsequent handling should be observed: e.g. (1) the dissecting microscope may correct the problem. If
furm and comparative length of the head, (2) form and bubbles appear in th e g lycerine-gelatine near the mar-
:olour of the eyes (if any), (3) shape of the rostrum gins of the cover slip, small amounts of glycerine-gel-
•·hen present) and of the epistome (in gammarids), atine may be picked up with a dissection needle, melt-
.! ) presence or absence of depressions or cairns and ed, and forced under the cover glass. The slide is then
!pines on the body, (5) presence or absence of fused allowed to cool, avoiding further manipulation until
xgments in the pereon and the urosome (see General the medium has completely hardened.
lorphology below). Subsequently, glycerine or
;lycerine-gelatine slides must be prepared with the Such slides may be used for a long time; and if the
~eopod s (taking care not to lose the coxae), uro- cover glass was mounted on a " ridge" of water-proof
?Ods, both antennae, telson, epimera (at least epimera varnish enclosing the medium with the appendages,
:::). and the mouth parts (it is preferable to place the they become practically permanent. Once mounted in
~-r on a separate slide) . In Hyperiidea epimera are the glycerine-gelatine slide, appendages cannot be
ery rarely used for diagnostic purposes, and in the removed without risking damage or loss. Canada bal-
1144 Arnphipoda
sam-mounted materials last even longer, but thi s tech- subtropical areas (down to the Subtropical Conver-
nique is more laborious and time-consuming, and is gence). Some species, however, are restricted fairly
not recommended for field conditions and/or with well to the tropics. Most warm-water hyperiids are
abundant materials. circumtropical. These include representatives of all
hyperiid families and practically all genera (except
Cyllopus, Hyperiella, and probably some abyssal
Geographic and vertical distribution Physosomata). Some families (mainly Oxycephali-
dae, Platyscelidae, Parascelidae, Lycaeopsidae, Ana-
Hyperiids pronoidae, but also Pronoidae, Brachyscelidae,
Lycaeidae) are entirely warm-water. Some hyperiids
Although records of hyperiids in nearshore areas are (especially several deep-water Physosomata) have
not uncommon, they are essentially pelagic animals. pan-oceamc or circum-oceanic distributions (see
(only Lestrigonus bengalensis tends to inhabit coasta l Table 1).
waters; Bowman, 1973), though they do not particu-
larly avoid them. Based on their thermal relationships, Large high-speed nets came into common use in the
hyperiids can be divided into cold-water and warm- 1970' s. Thus, detailed data on the hyperiid fauna of
water species. various parts of the ocean have become available only
recently, particularly from the tropical and subtropical
In the southern hemisphere, cold-water hyperiids are areas of the Pacific and Indian Oceans (Shulenberger.
associated with circum-Antarctic, bipolar (antitropical) 1977; Siegel-Causey, 1982; Young and Anderson.
or pseudo-antitropical areas. Bipolar and circum- 1987; Barkhatov and Vinogradov, 1988; Young, 1989:
Antarctic species have similar distributions here, but Vinogradov, l990a, 199 1, 1993b). These studies show
the former are present also in boreal and Arctic regions. that species compositions are relatively homogeneous
Sometimes it is not clear if a given species occurs in within the vast tropical anticyclonic gyres, and domi-
only one hemisphere or both (see Themisto gaudichau- nant species are also more or less similar in the various
di, Primno macropa), and distinction between circum - collections. Although the most abundant species may
Antarctic and bipolar species is partly formal. There vary somewhat between areas, they invariably include
are, however, some clearly circum-Antarctic species Phronima atlantica, Phronimella elongata, Phrosina
with no representatives in the northern hemisphere (e. semilunata, and one species of Primno. Also, Scina
g., Cyllopus). The northernmost distributional boun- crassicornis, Anchylomera blossevillei, Hemityphis
dary of these species is usua lly the Antarctic tenuimanus, Vibilia armata, and a representative of
Convergence (=Polar Front, 45-60°8), but they range Brachyscelus are often among the first ten.
occasionally to the Subtropical Convergence (35-40°S, Unfortunately, large collections have not been taken or
e.g., Hyperiella antarctica). Aided by northerly cold published from the Atlantic Ocean.
currents, some of these species may at times reach even
lower latitudes (e. g., Themisto gaudichaudi). The third Hyperiids inhabit the entire water column, but the ver-
distributional pattern includes the pseudo-antitropical tical distribution of the two infraorders differs strong-
species, characterized by many species of the genus ly. The more primitive Physosomata inhabit mostly
Hyp eria. These animals are abundant in cold-water deeper layers, including the hadal, only a few (some
regions and very rare in subtropical and tropical waters. Lanceola, especially Lanceola felina , as well as some
As a rule, they inhabit vety broad depth-ranges, at the Scinidae) having colonized the surface and subsurface
surface in cold-water areas, deeper in warm-water layers. Adult females of Mimonectes with eggs are
regions, avoiding the warmer surface layers. In the found near the surface. Most common Atlantic Scina
South Atlantic this type of distribution is well exempli- are usually fou nd at 500 m (Thurston, l 976b). Strictly
fied by Hyperia spinigera and Hyperia medusarum. In abyssal subspecies of Scina are also known (e.g ..
cold-water regions, Themisto is by far the most widely Scina wagleri abyssalis) , and many Scina species
disttibuted genus. (sometimes those caught at 100-200 m) inhabit depths
of 4000 m or more (Vinogradov et a!. , I 982). Other
Warm-water hyperiid species may also have fairly Physosomata are almost exclusively deep-water ani-
broad ranges, covering not only the tropical, but also mals, inhabiti ng depths mainly below 3000 m.
Amphipoda 1145
World-wide South Atlantic Vertical

Species
distribution distribution distribution (m)
Acanthoscina acanthodes Circumtropical 0-30°S 200-500
~ Amphithyrus bispinosus Circumoceanic 0-302 S, E Atlantic 0-300
Amphithyrus sculpturatus Circumoceanic 0-302S, E Atlantic 0-100
-\mphithyrus simi/is Atlantic and Pacific Warm water 0-200
A napronoe reinhardti Circumoceanic 362 S, 82 E 25-500 1A
-~
£:'z_ Anchylomera blossevillei Circumoceani c Tropical-Subtropi cal 0-300 2, 3
Warm w. and Boreal E
~ Andaniexis abyssi E Atla ntic 200- 1000 4
Atlantic,
Around South Afri ca
~ -\ndaniexis australis Cape Basin 1000-5000 (0-1000) 4
(Atlantic an d Indian)
-\rchaeoscina steenstrupi Circumocean ic 2 2
43 20'5, 46 02'W 200-500 1A
Circumocean ic in the
Bathystegocephalus globosus Tropical-Subtropical 100-1500 5
S Hem isphere
Brachyscelus crusculum Circu moceanic 0-32 2 5 0-400
Brachyscelus globiceps Circumtropica l Warm water 0-500
Brachyscelus rapax Circumoceanic Around Cape of Good Hope 0-500
Calamorhynchus pellucidus Circumtropical 0-402 5 0-100 (>100)
: t= -
Cleonardo longipes Cosmo12olitan? W hole area? 1000-2000 (4000) Rare, 6
Crano cephalus scleroticus ___,___ Circumtropical 0-402 S Epipelagi c
Ctenoscina brevicaudata C ircumoceanic A round Cape of Good Hope 700-2000 (5000?) Rare
y llopus lucasi Circumantarctic Antarctic 0-1000 2
~
Cyllopus magellanicus Circumantarctic Antarctic 0 -1000 (2000?) __2_______.
Ci rcumocean ic,
Cyphocaris anonyx Whole area 0-2500 (adults >500)
except polar seas
Ci rcumoceanic,
Cyphocaris chal/engeri Whole area 200-500 (0-200)
except pol ar seas
Ci rcumoceanic, except
Cyphocaris faurei Whole area 500-1 000 (0-2000)
borea l and Arctic
Circumoceanic, except
Cyphocaris richardi Whole area 300-5600 (7800?)
Arctic Basin
Cy stisoma fabricii Circumoceanic 0-502S 200-1000
Cystisoma longipes Circumocean ic 0-35°S 200-1 000
Cystisoma magna Circumtropi ca l Warm w ater Deep water Rare, 1A
Cystisoma pellucida Circumocean ic 0-352 S 200-1000
Atlantic, Pacific,
Dairella latissima 0-382 S 0 -1000 Rare
Mediterranean
Danaella mimonectes Bipo lar 2
58" 11'S, 61 00'W 500-3200 1A
~
Euandania gigantea Circurnoceani c extrapolar W hole area 1000-4000
Eupronoe armata Circu mtro Jical 0 -302 S E Atl antic 0-100
Eupronoe maculata Warm waters 0 -302S, E Atlantic 0-200
s::::SI:: Eupronoe minuta Circumtropical 0-302 S, E Atlantic 0-100
Eurythenes gryllus Circumoceanic Whole area 500 -4000, (0-500, 6500) 7
=- ~
Eurythenes obesus Circumoceanic 0-5225 500 (5500)
Eusirel/a elegans Atlanti c Ocean Whol e area 0-3000
Eusirella flagella Weddell Sea Weddell Sea
Eusirella multicalceola Ci rcumoceanic below 602 Whole area l 500 -2000-(4000)
Eusiroides stenopleura Circumantarctic South of 432 S 700-2000 (0)
Eusiropsis riisei Circumtropica l 0-45°S 0-500
E.usirus antarcticus Antarctic, partly Subantarcti c South of 54-602 S 0-1800
Antarctic (Scotia,
Easirus perdentatus Antarctic 0 -2 000
Weddell Seas)
1146 Amphipoda

Species distribution distribution (m) Remarks
distribution
Eusirus properdentatus Antarctic Antarctic (Scotia Sea) 0-1200
Euthamneus rostrat us Circumocean ic 0-62 S 0-500
C /ossocepha /us
Circumtropica l 0-409 S 0-200 (>200)
milneedwardsi
Circumocean ic,
Halice macronyx Whole area 500-2000
including Antarctic
Circumoceanic,
Halice secunda Whole area 500-2000 (3000)
- -"1-
j_ncluding Antarctic
Circumoceanic, I
Harcledo curvidactyla 67°S, whole area? 500-2000 (3000)
including Antarctic 1-
Hemiscelus diplocl]~latus 152 45'S, 332 11 'W
? Epipelaglc? _
~
H~m ityphis ~nuima n us Circumocea nic Around Cape of Good H ope 0-200
H yperia gassa _ _ Trqpical Atlantic Gulf of- Guinea
-
_Q-300 (1 000?)
-r Ra re
Hyperia curticephala
r Upwellings off Peru
-
Off Namibia
-
surface layer 8
and N<!!!l ibia
Hyperi'!_macroce@ala
H yperia medusarum
- --t
Antarctic
Circumoceanic
p~ud<:>ant i tro p_i c<li_
_!.rou nd So~h Georgi'!_
Coasts of S Africa,
_ Nami bia,_2 A rgent i n~
0-300 I 9
Ci rcumoceanic,
H yperia spinigera W hole area 600-900 (25-2000) 10
~xcept Arct.k Basin
South of the Subtropica l
Hyperiella antarctica Circumantarctic 0-500 11
(~vergen ce_
-+-
Hyperiel/a dilatata Ci rcumantarctic South of 55 S 2 0-500
H yperiel/a macronyx_ Antarctic Around South Gerogia I
fi:1yperietta steb bingi
ljyperietta stephenseni
Warm waters
Circumtropical _
Warm water
Tro£_ica l
0-200
0-200
T
/jjlperietta vosseleri 1 _ CircumtrQgi cal Tropical 0-200
H yperioides /ongip_es
Hjperionyx mayoda~!Yius
Hypf!!._oc;f!_e cryf!I_odacty/us
Hyp_eroche luetkeni~s _
Circumoceanic
_ CircumtrQgical
?
Circumantarctic
0-382 S
Around South Africa
O ff Ca pe of Good H op~
Around South Georgia
Q:3Q9 (600)
0-200
_!:.pi pelagic
Epipelagic
-t
/-j_yp~E[oche martinezi ... Atlantic Off Brazil _ ~pipe l~ic_
H yperoche med usarum
lufs!pis lovenj_
BJB.olar
_i:irc u mtro~ca l
South of 25-35 2S
_ TrQPica l _
0-300
Epi pelagic
1
Circu mocean ic,
Lanceola clausi W ho le area 1000-3000 (200-5000)
i nclud i ng_£hE£f.rctic
Lanceola felina
Lanceola loveni
Circumtropical_
Ci rcumoceanic
0-372 S
-Warm water
--
0-200
_l000-2000200-400QL
i=-
3000-4000 (>4000),
Lanceola pacifica Ci rcumoceanic Whole area except Antarctic
_juveniles !90-20_Q_ _
Lanceola sayana Ci rcumoceanic
Circumoceanic,
W hole area _! 000-200Q_(0-3000)
I
Lanceola serrata 0-52 2 S, rare in the tropics 750-1500 (300-2000)
except Arcti ~as in
Lep_to co tis ten uirostris L Circu mtropical 0_00°S, abse_!!_t off Afric<!__ 0-500 12
Lestrigonus bengalensis
1 Lestrigon us
crucipes
Ci rcumtropical
Ci rcumtropical
Tropical, coastal
Warm water
0-200
0-200
l
Lestrigon us latissimus Tropical E tropica l Atlantic 0-100 I
Lestrigonus macrophtha/mus Circumtropical Equatorial 0-100
Lestrigonus schizogeneios Circumtropical 0-452 S 0-200 13
Lycaea nasuta Circumocea ni c Around South Africa 0-300
Lycaea pachypoda Ci rcumtropical Tropical 0-300
Amphipoda 1147

Species
Lycaea pulex complex Circumtropical Tropica l 0-500
Lycaea serrata Circumtropical Equatorial 0-300
Pacific, Atlantic, Red Sea, Very
Lycaeopsis themistoides Off Namibia Epipelagic
Mediterranean rare
'v!egalanceola remipes Atlantic and Indian 41 °43'S, 42 2 20'W 0-2000 1A
'-'letacyphocaris helgae Circumocea nic extrapolar Whole area 200-1000 (0-5000)
\1imonectes gaussi Circumoceani c Around South Africa >500
\1imonectes /oveni Circumoceanic (N hemisphere) Gulf of Guinea 200-1000
\1imonectes sphaericus Ci rcurnoceanic Tropical (rare- whole area?) (0)-200-2000
Orchomenella abyssorum Circumoceanic Off Buenos Aires (at 3475 rn) 200-8300
Austral lndowestpacific,
Orchomenella distincrus Tropica l 2000-5000 (0 -2000)
Pacific, Atlantic
Austra l lndowestpacific,
aro= Orchomenella gerulicorbis Tropical 750-6000
Pacific, Atla ntic
Oxycepha/us clausi Circumtropica l Tropical 0-100 (200)
Oxycepha/us /atirostris Circumtropical Tropical 0-200 Rare
O'Cycephalus piscator Circumtropical Warm water 0-100 (2000?)
Paracallisoma alberti Circumoceanic, mainly bipolar Whole area 1000-4500 (0-1000)
Circumoceanic, chiefly N
Paracyphocaris praedator Equatori al 300-3500
and tropica l Atlantic
Paralycaea gracilis Circumoceanic Warm water 0-500
Circu rnoceanic,
Parandania boecki Whole area 500-6000 (0) 14
except Arctic Basin
Paraphronima crassipes Circumoceanic extrapolar 0° to Pol ar Front 0-500 (500-1500)
Pa!EfJhronima gracilis Circurntropical Warm water 0-500
Parapronoe campbe/li Circumoceanic O ff Namibia and South Africa 25-500
r>arapronoe crustulum Circurnoceanic Warm water 50-700
0 arapronoe parva Circumtropi cal 0-302 5, E Atlantic 25-300
:>a•ascelus edwardsi Circumtropical 0-3 9S 0-200 15
0 arascelus typhoides Circumtropi cal 0-362 5 0-200
"ari!JYphis maculatus Circumoceanic Around Cape of Good HoQ_e 0-500
:>aratyphis parvus Circumtrqpical Gulf of Guinea 0-500
Paratyphis promontori Circumoceanic Around Cape of Good Hope 0-500
i'egohyperia princeps Circumoceanic 33°07'S, 432 00'E in haul 0-2000 Rare, 1A
:><, ppsiella nipoma Circumocean ic extrapolar W hole area, mainly WW 3000-6300
P..•onima atlantica Circumoceanic Warm water 0-1 000 (1500)
Phronima colletti Circumtropical _!~Qf!.ical 0-300, mainly epipelagic
f'::-onima curvipes Circumtropica l Warm water 0-300
"""-•anima pacifica Circurntropica l Warm water 0- 100 (100-400)
"'"-"'nima sedentaria Circurntropical Warm water 0- 11 00
=--.'anima solitaria Circumtropica l Warm water 0-1000
P"-•onima stebbingi Circumtropica l Warm water 0-500, mainly epipelagic
"""-"DD1imel/a e/ongata Circumtropica l Warm water 0-50 (50-200)
-onimezpsis spinifera Circumtropical Tropical, off Namibia - - - t - 0-500
~ina semiltmata Circurnoceanic Warm water 0-500 (1 000) 2, 3
W Atl.: 0-202 S,
?.: s1ca marina Atlantic Coasta l, shallow water 4
E Atl.: off South Africa
-:-.see/us armatus Circurnoceanic 0-25°5 (E Atlantic) 0-800
--~see/us crustulatus Circumtropical Equatorial 0-200
~see/us ovoides Circumtropical 0-309 5, E Atlantic 0-200 (200-800)
~;,ce/us serratulus Circumoceanic extrapolar 0-30°5, E Atlantic 0-50 (0-300)
1148 Amphipoda

Species distribution Remarks
distribution distribution (m)
Primno brevidens Circumoceanic extrapolar Gulf of Guinea 0-500 (500-900)
Primno latreillei Circumtropical Warm water 0-1000
South of the Polar Front,
Primno macropa Circumantarctic 0-1000 16
oceanic
Pro noe capita Circumoceanic extrapolar Warm water 0-200 Rare
Rhabdosoma armatum Ci rcumtropical Tropical 25-50
Rhabdosoma brevicaudatum Atlantic and Pacific Equatori L 0-50 (50-500)
Rhabdosoma minor Ci rcumtropical Tropica l 0-300
Rhabdosoma whitei Circumtropica l 0-402 S ~0- 2 00 (>200)
Schizoscelus ornatus Circumtrop ical 0-142 S 0-200
Scina borealis Ci rcumoceanic Whole area 200-1000 (50-3000)
Scina crassicornis Mainly circumtropical 0-662 S 0-500 (>500) Common
Scina curvidactyla Mainly circumtropical 0-502 S 100-1 500 Common
Scina excisa Circumoceanic, mainly tropical 0-352 S 200-500
Circumoceanic,
Scina incerta Whole area except Antarctic 500-75 0 (200-1000)
except Arctic Basin
Scina /anghansi Circumtropical Equatorial 0- 1000
Very
Scina lalifrons Circumtropica l Tropical
rare
Scina lepisma Warm waters 0-302 S 200-1000
Scina marginata Circumtropi cal 0-202 S 40-1000
Scina nana Circu mtropical Tropica l 100-2000 17
Scina oedicarpus Circumtropical 0-352 S 200-1000
Scina pubera Atlantic and Indian Warm water 0-1000
Scina pusilla Circumoceanic? Gu lf of Guinea 500-1000 Rare
Scina rattrayi Circumocea nic 0-33 2 S 150-700
Scina simi/is Atlantic and Indian, tropica l Equatorial 0-500
Scina spinosa Circumoceanic 0-552 5 200- 1000 (1000-4000)
Scina stebbingi Atlantic and Pacific 0-402 S 40-1000
Scina stenopus Warm waters 0-362S 100-500 (500-1000)
Scina submarginata Circumocea nic 0-55 2S 40-4000
Scina tullbergi Ci rcumtropical Tropical 0-700
500-1800,
Scina vosseleri Warm waters
juveniles 50-100
Scina wolterecki Circumoceanic Tropical and SE area 1000-2000 (200-4000)
Scopelocheiropsis ab yssalis Circu moceanic? Whole area? 0-3000
Scypholanceola aestiva Circumoceanic 0-35 2S 1000- 1500
Simorhynchotus antennarius Ci rcumtropical Tropica l 0-500
Stenopleura atlantica Ci rcumtropical 0-352 S 0-400
Streetsia challengeri Circumoceanic 0-452 S 0-300 (300-1 000) - -
l
Streetsia mindanaonis Ci rcumtropica l 0-302 S 0-500
Streetsia porcella _ _ Warm waters 0-35 2 S 0-300
Streetsia steenstrupi Circumtropica l 02 to Subtropical Convergence 0-300
Synop ia gracilis Tropical Atlantic Tropical water_s_ _
Tropica l (especially Caribbean) 0-50
Synopia schee /eana Tropical waters 0-50
Atlantic and tropica l Pacific
Synopia ultramarina Circumtropica l Tropical waters 0-50
Tetrathyrus forcipatus Circumoceanic 02 to Cape of Good Hope Subsurface
Themistella fusca Tropical Gulf of Guinea Epipelagic (0-200)
Themisto gaud ichaudi Bipolar or circumantartic South of 352 5 0-500 2, 3, 18
Thoriella islandica Circumocean ic W hole area? 1000-3000 (500?) Very rare
Amphipoda 1149

~ Species
Thyroeu s se_haeroma Circumtropical 0-36°5 0-200
Tryphana malmi Anti equatorial Off Namibia and South Africa 0-200
\'ibilia antarctica Circumantarctic Antarctic 0 -1000 19
\ 'ibilia armata Circumoceanic 0-50°5 200-600 (0-1 000)
.'ibilia australis Circumtropical 0-50QS 0-700
\'ibilia chuni Ci rcumtropical Tropical 0-500 Rare
.'ibi!ia cultrie_es Circumoceanic Tropical 0 -300 Rare
.'ibilia gibbosa Atlantic and Pacific 0 -20QS 0 -500
Atlantic, Indian,
•' bilia jeangerardi Warm water 0-500
Mediterranean I
.'ibilia propinqua Ci rcumocean ic 0° to Subtropica l Convergence 50-1000
•'bilia pyripes Circumoceanic Warm water 0-500
"bi/ia robusta Circumocean ic 0-3 0°5 0-500
-bilia stebbingi Warm waters Warm water 0-1000 Rare
:bilia viatrix Circumoceanic Warm w., but ranges to 50°5 0-150
-bilioides alberti 1 Circumoceanic Off Rio de j aneiro 200-1 000 (3000?) Rare
Lzbl.e 1. Distribution of the species (species listed in alphabetical order).

~:
Only record of this species;
..._ Only record from the South Atlantic;
~l.igrates into the upper layers at night;
Often forms swarms;
Bentho-pelagic, on the bottom associated w ith abundance of weeds, sponges, and hydroids;
- Often present in the near-bottom layer;
Found in the North Atlantic, northwestern and South Pacific and indian sector of the Antarctic;
b the water colunm £. g1yllus is represented as a rule by comparatively large specimens (length to 90 nm1). Juveniles and large(> I 00
=> females inhabit near-bottom layers, where they can even burrow into the ground for shelter. They are bentho-pelagic scavengers and
:!!<! frequently recovered in baited bottom traps (Thurston, 1990; Vinogradov et al. , 1996; Christiansen, 1996);
_-\ssociated with jelly-fishes of the genus Chrysaora;
Chiefly bipolar, in the South Atlantic extending northward with cold-water currents;
~~ the coast of southern Africa this species shows a clear preference for colder upwelled waters of the neritic region (Siegfried, 1963, as
H galba");
::; Argentine shelf waters found almost exclusively in oceanic waters, not penetTating into wam1er neritic waters; bouudary is correlated
"""1th I ooc surface isotherm (Ramirez and Vinas, 1985);
- "Sot found at water temperatures below 18.5°C;
- .-=usociated mainly with the jelly-fish Phialidium;
- =orded in the Nof\vegian Sea ;
_ ~:be . orth Atlantic recorded as far north as 34°N, suggesting that it may also inhabit mid-latitudes of the South Atlantic;
:=!A.-ords from tl1e nortl1crn hemisphere represent P abyssalis (Bowman, 1985). 1n Argentine shelf waters found almost exclusively in
.xcmic waters, not penetTating into warmer neritic waters, boundary is correlated with 11- 12°C surface isotherms (Ramirez and Vinas,
. =S5):
- Cl:::ie specimen found in the Antarctic Lndian Ocean;
:: g;nulichaudi can range into middle to low latitudes along South America and Africa carried by cold northbound coastal currents, and
~ward to the Antarctic Convergence inhabitating mainly cold Antarctic Intermediate Water. Animals cited for the northern hemisphere
=:;belong to a separate species: T compressa (see Schneppenheim and Weigmann-Haass, 1986).
~ specimens may be found in the tropics.
1150 Amphipoda
The Physocephalata are restricted entirely to surface even in the Notth Atlantic. Some of these species ha,·e
layers (with the exception of the family Cystisomati- restricted ranges; for example, abyssal and hada!
dae, which occurs to depths of 3000 m or more). The animals strictly associated with a fe w deep-sea trench-
lower limit of most Physocephalata is about 200-500 es, (like Hirondellea gigas from the Indian-Pacific
m, only a few venturing deeper (for example, some region). But others do not seem thus limited. The inYen-
Vibiliidae or Primno macropa, to depths of 1500- tory of South Atlantic pelagic gammarids probably is
2000 m). However, isolated specimens of subsurface, still far from complete. Coastal plankton samples rna~
and even epipelagic, species may also be found much contain many bottom-dwelling gammarids, practicall~
deeper than usual, occasionally to 1000-2000 m. For all of which venture high up in the water column ic
example, during the 1990 Russian expedition to the their diel migrations, but these species are not included
hydrothermal vent fields of the tropical Pacific Ocean, in the present review.
a slurp-gun from the deep-sea manned submersible
"Mir" caught a few Lycaeopsis zamboangae and Pelagic gammarids (including those which have nor
Parascelus edwardsi near the bottom at depths of lost connection with the bottom completely) inhabit all
1500-2300 m (Vinogradov, 1993a, 1994). depths at all latitudes. Most of them are meso- and
bathypelagic, but the mesopelagic animals often mo,·e
It is generally assumed that hyperi ids decrease sharp- to subsurface layers (especially Cyphocaris challenge-
ly in the thin near-bottom layer (e.g., Vinogradov, ri), or sink to abyssal and even hadal depths. Strictly
1990c). However, from the "Mir" submersible I have epipelagic gammarids are also known: in the South
observed swarms of Themisto abyssorum near the bot- Atlantic Stenopleura atlantica, Eusiropsis riisei, and
tom of the Norwegian Sea, with isolated animals even all species of Synopia (the latter practically hypo-neu-
contacting the sea-floor for short periods of time. stonic animals). And a few pelagic gammarids ha,·e
been recorded only from abyssal and hadal samples
Many Physocephalata (and some Physosomata, like (e.g., Scopelocheirus shellenbergi, several locations.
Scina) have die! activity cycles and limited vertical including the Orkney Trench). Bentho-pelagic sca-
migrations. The range of these migrations is usually venging gamma1ids are a major component of abyssa:
less than 100 m, but Scina crassicornis, Vibi/ia arma- epibenthic communities. Some never venture far
ta and sometimes Primno macropa, may rise as much above the bottom (many lysianassoids), and are there-
as 400 m from their daytime levels (Thurston, 1976b), fore not included in this review. Others, however, are
and Themisto has well marked die! vertical migrations often recorded in p lankton samples (Eurythenes.
(Bowman et al., 1982; Hiroki, 1988). Land (1992) Paracallisoma, Orchomenella).
reported some very interesting investigations on loco-
motion of hyperiids as a function of light. He showed Caprellids
that some species (Phronima, Platyscelus) cease
swimming or swim downwards under high ambient Bentho-pelagic caprellids may be found in plankton in
light levels, and statt swimming upwards as the light the upper 1000 m of water, mainly in neritic areas.
level falls below a given threshold, thus staying with-
in a selected lighting regime. In other hyperiids
(Brachyscelus, Phrosina), however, this mechanism Taxonomy and identification
has not been observed, suggesting that vertical move-
ments are triggered by other factors. General morphology
Gammarids Unfortunately, amphipods (especially the gammarids

have a bad reputation among planktologists and are
The geographic distribution of epipelagic gammarids often poorly identified, frequently referred to in the
is similar to that of the warm-water hyperiids literature on ly as undifferentiated "Amphipoda··_
(Synopia is more closely associated with the tropics However, despite their remarkable diversification, the
s.s.). Deep-water gammarids of the South Atlantic are identification of planktonic amphipods is not all ~
mainly pan-oceanic or circum-oceanic. On the other complicated. With a little practice major families anc
hand, records of deep-water pelagic gammarids are many genera may be easily distinguished, and species
very scarce in the Indian and the Pacific oceans, and identifications are not insurmountable.
Amphipoda 11 51
Pe reon
Cephalon... :-·...
PIeon
2
4
·· ·. ·· ··
: Urosome 5
F~ I. Diagram ofhyperiid amphipod, based on male of Hyperia (le ft; a fter V inogradov el a/., 1982), and typical pereopod with articles num -
~(right).
?d.ag:ic amphipods vary in length from 2 to 150 nun. is one of these seven segments fused to the head. In
:.C overall shape of the animals is globular to needle- some hyperiids, 2 to 5 of the first pereonites may be
"'- commonly elongate. In many p arasitic hyperiids fused to each other (especia lly in females). Amphipods
~ ia lly Physosomata and Physocephalata) and a differ from all other Malacostraca in having pereonal
=- garrunarids (also parasitic, like Danael/a or legs an·anged in two distinct groups: the first four pairs
- J:it:Jriella) the male body is elongate and compressed, directed forward (with the dactyls directed backwards),
;:;aea.s females are g lobular (Fig. 4 .34, 4.45). Females the last three pairs backwards (dactyls forward) (Fig.
~sometimes larger tha n males. Rarely (especially in 1). Their abdominal appendages comprise 3 pairs of
~i ds) the amphipod integument bears sharp or biramous swimming legs (pleopods) and 3 pairs of
.:;.':::n:t outgrowths, eithe r isolated (spines) or as ridges thrusting legs (uropods). The last ch aracter is unique -
arin ae). Such outgrowths are usually on the dorsal all other similar groups of crustaceans have 5 pairs of
~- rarely lateral) side of the animal, sometimes only pleopods and I pair of w-opods. In caprellids the abdo-
~e posterior body (see, for example, Eusirus, Fig. men and its appendages are strongly reduced.
- - In Calamorhynchus (Hyperiidea) two horizontal
~are on the lateral surface of the head. The head of an amphipod bears two pairs of antennae.
The first 3 articles of the first pair, and the first 5 arti-
~am phi pod head has 5 pairs of appendages, and ses- cles of the second pair are known as the peduncles. In
£r eyes (if an y). The thorax has 8 pairs of appenda- hyperiids some of the peduncular articles may be
~ The first thoracic segment, bearing the maxilli- absent, especia lly in antennae II, but the two first arti-
is fused with the head, and other segments have cles of the peduncl e of antenna II are always present
u.iramous legs. The abdomen has 6 p airs of bira- and fused to the head. The remaining smaller mticles of
legs. Lastly, there is a tenn inal supra-an al telson. the antennae form the flagellum. In many gammarids
-=phipods are an order of the Peracarida lacking a ca- there may be an accessory flagellum forking from the
~ covering the thorax, so seven distinct thoracic end of the third p eduncular article of antenna I, either
~nts (pereonites) are v isible. Only in the caprellids well developed (Fig . 4.1 8) or reduced. Hyperiids never
1152 Amphipoda
palp
Maxilla I
Maxilliped Mandible
Maxilla II
Fig. 2. Typical gammaridean mouth parts (after Barnard, 1969). ln different Gammaridea these may be reduced to various degrees. In
..
Hyperiidea, all palps, inner plates of the rnaxillipcds, and entire maxillae may be strongly reduced (sec Fig. 4.48, 4.99, and 4.104 for examples
of hyperiidcan maxillipeds).
possess this accessory flagellum. Sometimes the anten- dosoma, Fig. 4.1 51), rarely rounded or hood-like pro-
nae (especially in gammarid males) have calceoli - tiny cess tenned rostrum is sometimes present on the ante-
globular sense organs attached to the antenna (absent in rodorsal margin between the peduncles of antem1ae I.
hyperiids).
Eyes, when present, are very variable both in shape
In Hyperiidea, flagellae of the antennae ~re
often more (round, oval, elongate, irregular) and in size (from
or less reduced. In many species only the proximal very large, to a few separate ommatidia, to a simple
conical article is developed (or even hypertrophic), pigmented spot). In hyperiids the eyes can cover the
although a few rudimentary distal articles may also be entire head (in the Physocepha lata). Sometimes
present (Fig. 4.37). In Hyperiidea Phronimoidea and hyperiid eyes are strongly modified, having lost not
Platysceloidea antennae are strongly reduced in only the pigment (as in the Cystisomatidae), but even
fema les but well-developed in males. In Phronimoidea the refracting elements (Scypholanceola). In many
both pairs have multi-articulate flagellae. In Platy- Physocephalata the eyes are double structures, with a
sceloidea antenna II (Fig. 3d) is long, irregular, 5- specialized upward-pointing region covering a narrow
articulated (3 elongate peduncular articles + 2 elon- field of view (Land, 1989). In some cases eyes may be
gate flagellar atticles). In Lycaeopsidae, antennae are partly reduced, inconspicuous externally, and partly
short in both sexes, but in females antennae II are covered by the integument; in th is case they are
absent. In Hyperiidea Physosomata both antennae termed "glandular", referring to their gland-like
may be reduced in different combinations, but mor- appearance. The presence/absence and characteristics
photypes with gigantic proximal articles of the flagel- of the eyes are more important as a d iagnostic charac-
lum of antennae I also occur. ters in hyperiids than in gammarids.
The gland cone (or antenna! gland) is located in the The mouth parts are highly variable and their mor-
proximal patt of the peduncle of antennae II; its shape phology is imp01tant for classification. All mouth
is an important taxonomic trait in some hyperiids (Fig. parts are concentrated in a buccal mass beneath the
4.114). A triangular, spine-like or needle-like (in Rhab- head. They include the following structures (F ig. 2):
Amphipoda 1153
l..:pper lip: a single lobe or flap antetior to the mouth. pmtly or completely separated in most of the
The frontal part of the head above the upper lip is Physosomata, but completely fused into a single
termed the epistome; normally quite indistinct, median lobe in other hyperiids, or reduced. Outer
but occasionally protruding anteriorly as a laminar lobes may be broad or narrow, in some cases fused
rounded lobe. into a single plate (Paraphronimidae). Maxillipeds
Lower lip: a bilaterally symmetrical structure form- were originally the first pair of thoracic legs, but
ing a partition behind the mouth. It consists of one they (with their body' s segment) have become
pair of lateral or outer lobes, provided with later- completely incorporated into the cephalic complex
al projections, and one pair of inner lobes, which and are therefore included in the mouth parts, rath-
are often reduced or fused. Lips, especially the er than among the legs of the pereon.
lower one, are important taxonomic characters for
the gammarids, but are of little use in hyperiids. Pereon (or thorax): this section bears seven pairs of
:)landibles: a pair of appendages attached laterally to uniramous legs called pereopods I-VII. Pereopods I
the mouth. The mandibles have powerful muscles and fT also are termed gnathopods I and II. Pereopods
and often solid anterodistal processes called are 7-articulated: the first article, attached to the body,
incisors which are often strongly serrate. Right is termed coxa (or coxal plate), atticle 2 - basis, arti-
next to the distal "teeth" may be an articulated ser- cle 3 - ischium, article 4 - merus, article 5 -carpus,
rate accessory plate, the lacinia mobilis, which article 6 - propus or metacarpus, and article 7- dac-
may be present on only one of the mandibles. A tyL However, coxa and dactyl are often (including
molar, with or without a grinding surface, often this review) the only terms used, other articles being
occurs on the media-ventral smface of the man- referred to by their numbers.
dible, but it may be reduced or absent. Most
arnphipods have a 3-articulated palp attached to The coxa is an immovable flat plate inserted into the
the dorsolateral surface of the mandible. In lateral surface of the pereonite (pereon segment), and
Hyperiidea this palp is often absent, or may be sometimes fused to it (in hyperiids). Jn gammarids the
present in the male and absent (or, more rarely, coxa may be enlarged, forming a lateral shield (Fig.
reduced to 1 or 2 articles) in the female of the 4.29), while in other suborders it is always small.
same species. Other articles may broaden in different ways or be
la.xilla 1: In Gammaridea this pair of appendages is elongated, more or less narrow and with parallel mar-
small, each bearing an inner lobe (=plate) (some- gins. Atiicles 5-7 or 6-7 sometimes form the chela or
times with setae), an outer lobe with spines, and subchela, especially on the gnathopods, but some-
anached to the latter, a palp (reduced or absent in times on other pereopods as well. However, prehen-
some cases) comprising 1 or 2 articles. In sile pereopods are not always present, and all pereo-
Hyperiidea Physosomata (but not in the pods are simple in many amphipods. Prehension is
Physocephalata), the maxilla I has an inner lobe, usually accomplished in byperiids by pressing article
and the palp, when present, is formed by a single 6 (provided with a small dactyl) against an expanded
micle. In some physocephalata (Lycaeidae, miicle 5; whereas in gammarids (and in some hyper-
Oxycephalidae) the maxilla T can be rudimentary iid cases as well) the strong dactyl presses against an
or absent. expanded article 6. In chelate pereopods the paJm, the
Yn:illa II: This pair of appendages bears simple part of surface or margin of the article on which sub-
outer and inner lobes (=plates). Tn gammarids a sequent ariicle(s) close(s) for the purpose of prehen-
:ow of setae on the dorsal face of the inner lobe (or sion, is located on the distally produced carpal pro-
~eir absence) is important for systematics. In cess (Fig. 4. 147). In subchelate pereopods the carpal
~-peri ids the maxilla IJ may be rudimentary or process is Jacking, and the palm located on the distal
~sent (Lycaeidae, Oxycephalidae). or posterior margin of the article (Fig. 4.9, 4.129).
lh:rillipeds: Maxillipeds are formed by an inner and
::::t outer lobe (=plate). In gammarids the outer When the carpal process is small, and it is difficult to
::.roe has a 4-articulated palp (rarel y reduced), decide whether the pereopod is chelate or subchelate,
~ereas the palp is always absent in hyperiids. In the term weakly chelate is used (Fig. 4.20 P I).
~ latter the inner lobes of the maxi llipeds are Weakly subchelate means that the palm of the sub-
1154 Amphipoda
chela is not distinct. Sometimes it is impossible to are hatched and the young develop (amphipods lack
draw a clear line between simple and weakly subche- planktonic larvae and new-boms resemble miniature
late pereopods. In hyperiid families Lanceolidae and adults). These structures are located on pereopods 2-
Chuneolidae (the latter not recorded in the South 5, medial to the gills, and present only in mature
Atlantic Ocean) the apex of article 6 is expanded in females. In Garrunaridea oostegites have marginal
some pereopods into a hood surrounding the base of setae in the latest stages of development, whereas in
the dactyl. The dactyl may be flexed posteriorly so Hyperiidea these setae are absent. In Rhabdosoma
that it lies within the spoon-shaped cavity within the (Hyperiidea) the oostegites are reduced in size, and
hood (also called retractive dactyl; Fig. 4.42). their function taken over by the gills.
Gnathopods (pereopods) I and II: are commonly Abdomen: comprising 6 segments, 3 segments bear-
smaller and more delicate than other pereopods, more ing pleopods corresponding to the pleon sensu stricto.
or less prehensile, and rarely simple. Often the gnath- and 3 segments of the urosome (bearing uropods).In
opods are hidden behind other pereopods. The shape hyperiids mosomal segments (urosomites) II and III
of the distal articles of the gnathopod is an important are fused. In caprellids the abdomen is reduced.
taxonomic character. In gammarids, gnathopods II are Ventrolateral plate-like extensions of the pleon seg-
usually sexually dimorphic, with the chela or subchela ments are called epimera or epimeral plates. The
strongly enlarged in males. Such dimorphism is espe- shape of the free margin of the epimera III and II is
cially noticeable in benthic species, whereas in the imp01tant in gammarid taxonomy.
planktic ones it is weak, or absent altogether. In hyper-
iids only Phronimopsis has sexually dimorphic Pleopods: paired biramous swimming appendages on
gnathopods II, its chela larger in females than in males. the segments of the pleon (first three abdominal seg-
ments). The rami are multisegmented and strongly
Pereopods Ill and IV: these are usually slender and setose.
their dactyls are directed posteriorly.
Uropods: the last tlu·ee abdominal segments bear uro-
Pereopods V-VII: are often more robust and elongate, pods. Uropods are paired biramous appendages
and their dactyls directed anteriorly. Article 2 of these attached laterally to the body. They comprise the basi-
pereopods is often inflated. In Hyperiidea the distal podite and two branches or rami: an exopodite (outer
articles of pereopods VI and VII may be reduced (or and endopodite (inner). The rami are uni-articulate.
completely absent in pereopod VII). Pereopod VII may except in gammarids, where the exopodite ofuropod ill
be much smaller than the rest. Pereopods III through may be 2-atticulated. In hyperiids the endopodite is
VII are usually simple, but a few may be prehensile in sometimes fused with the basipodite, and the exopodite
some cases. In Phronimidae and Phrosinidae reduced (Cystisomatidae, Oxycephalidae, Scinidae). In
(Hyperiidea) pereopods V are extremely subchelate other cases, parts of the uropod are completely fused or
(even almost chelate in Phronima). In Phronima the reduced, and the an imals have only one petal-like arti-
subchelae of pereopod V are sexually dimorphic. cle, rather than a "n01mal" uropod (Phrosinidae).
In some species pereopod articles may bear conspicu- T elson: a flap attached to the last urosomal segmenL
ous glands, in which case they are termed "glandular" above the anus. In Gammaridea it may be divided
in the keys and diagnoses that follow. (=cleft) (Fig. 4.23, 4.32) or tmdivided (=entir e) (Fig.
4.9, 4.30); whereas in Hyperiidea it is always undivid-
Gills: hollow sacs inserted into the medial surfaces of ed (Fig. 4.93 , 4.173). It may be longer or shotter than
the coxae. Typically there are 6 pairs of gills in gam- the basipodites of uropods III. In hyperiids it is some-
marids, 5 in hyperiids, and 3 in caprellids, but some times fused with the urosome.
pairs may be reduced. Gills are always lacking on the
first pereopods. All appendages (from mouth parts to uropods) may
bear setae. The number and distribution of these setae
Oostegites: thin plates which overlap to form a pouch (especially on the mouth parts) are important taxo-
beneath the pereon (marsupium) in which the eggs nomic characters, especially in gammarids, where
Amphipoda 1155
~ey may be pinnate or simple. In Hyperiidea they are In general, only taxa recorded from the South Atlantic
2.'ways simple. plankton are included in the section below, and only
their characters are taken into account. A few excep-
: or more detailed description of amphipod morpholo- tions, of special interest for various reasons, are
g:· see Gu1janova (1951); Bowman and Gruner denoted with an asterisk. Also, additional information
:973); Lincoln (1979), Vinogradov et al. (1982), and of potential usefulness in several systematically sensi-
i3amard and Karaman (1991). tive cases is often included in square brackets.
Abbreviations used in keys, figures and diagnoses are
as follows:
Outline classification ANT: antenna,
ART: article(s),
- ~e classification of the planktonic amphipods of the CX: coxa,
South Atlantic Ocean is surnn1arized as follows (this MD: mandible
~• includes all the planktonic or partly planktonic taxa MX: maxilla,
orld-wide, with South Atlantic taxa underlined): MXP: maxillipeds,
P: pereorod(s),
~:Uer Amphipoda T: telson,
Suborder Gammaridea UR: uropod(s); in figures also refers to the entire uro-
Families Eusiridae, Hyperiopsidae, Iphi- some.
mediinae, Lysianassidae s. l. , Pardaliscidae,
Stegocephalidae, Stilipedidae, Synopiidae,
Vitjazianidae (may include displaced benthic I dentification : keys a nd diagnoses
representatives of other families in shallow-
water plankton, especially at night - see Intro- Key to the suborders and families of Amphipoda
duction) 1 Gills exceeding 3 pairs, head free from pereonite
Saborder Hyper iidea I, body compressed or flattened, rarely needle-
Infraorder Physosomata like, abdomen well developed ............ .............. 2
Superfamily Archaeoscinoidea Ia Gills not exceeding 3 pairs, pereonite I fused to
Family Archaeoscinidae head, body skeletal, segments tubular.
Superfamily Lanceoloidea [Abdomen vestigial, not segmented; MD with
Families Chuneolidae, Lanceolidae, Micro- fe w lacinia mobilis]: Suborder Lemodipoda
phasmidae (=Caprellidea): Family Phtisicidae (Fig. 4.184)
Superfamily Scinoidea 2 Palps ofMXP present (but may be rudimentary),
Families Mimonectidae, Proscinidae, Scinidae urosome with 3 segments. ANT I may bear
::nfraorder Physocephalata accessory flagellum. Setae on appendages may
Superfamily Vibilioidea be pinnate. T may be cleft. (Suborder
Families Cystisomatidae, Paraphronimidae, Gammaridea) .......................... ........ .................3
Vibiliidae 2a Palps of MXP absent, urosome with only 2 seg-
Superfamily Phronimoidea ments (urosomites II and III fused). ANT I
Families Dairellidae, Hyperiidae, Phronimi- always without accessory flagellum. Setae on
dae, Phrosinidae appendages are never pinnate. T never cleft
Superfamily Lycaeopsoidea (Suborder Hyperiidea) ......... .... .. ..... .......... ... ... .. 8
Family Lycaeopsidae 3 Gnathopod II is elongate, slender, minutely che-
Superfamily Platvsceloidea late or subchelate, ART 3 elongate, ART 5 and 6
Families Anapronoidae, Brachvscelidae, densely setose, dactyl minute (Fig. 4.18). ANT I
Lvcaeidae, Oxycephalidae, Parascelidae, with broad, barrel-like ART 1 of peduncle:
Platvscelidae, Pronoidae, Tryphanidae Lysianassoidea (=Lysianassidae sensu lato)
Sc.:xlrder Lemodipoda (=Caprellidea) 3a Gnathopod II and ANT I of another type ........ .4
Family Phtisicidae 4 Pereon high, laterally compressed, lens-like.
Subfamily Phtisicinae Coxal plates I-IV high, forming broad lateral
1156 Amphipoda
shield. ex v small, located in posterior excava-

tion of ex IV (Fig. 4.29). MD lacking palp and
molar: Family StegocephaJidae
4a Pereon, coxae and MD of another type ............. 5
5 Mandibles lacking molar (palp present). [Inner
plates ofMXP strongly reduced. T cleft]: Family
Pardaliscidae
Sa Mandibular molar well developed, cylindrical Vibilioidea Lycaeopsoidea
...........................................................................6
6 Mandibular palp extremely tumid. [Both T and
peduncle of UR III very short, subequal in
length. Accessory flagellum of ANT I multi-
articulate]: Family Synopiidae
6a Mandibular palp more or less slender.. ........ .....7
7 Mouth parts symmetricaL Palps of both MX I of
common type, more or less flat P I and li sub-
chelate. [ Acessory flagellum of ANT I absent or
small (1-2 articulate). Pleon well-developed]:
Family Eusiridae
7a Mouth parts asymmetricaL Palp of right MX I Phronimoidea Platysceloidea
bent, spoon-like. P I and li simple. [ART 4 of P
lli-IV greatly elongate relative to ART 5. P V-Vll Fig. 3. Diagrams of male heads of Hyperiidea Physoccphalata (a ftc
(including ART 2) long, slender, thin]: Family Bowman, 1973).
Hyperiopsidae*. Representatives of this family
have not been observed in the South Atlantic, but
are common in the plankton of the World Ocean. 10 P VI-VII without spoon-shaped ART 6 anc
Two genera: Hyperiopsis with globular head, hooded dactyL [MD with narrow serrate incisor.
small coxae IV, broadly expanded ART 4 ofP III- well-deve loped lacinia mobilis]: Fam il~
IV, Parargissa with large coxae IV (which cover- Archaeoscinidae
ing coxae IT-III), slender ART 4 of P III-IV. lOa At least P VI and VII with spoon-shaped ART 6
8 Head small, usually shorter than pereonite I. and hooded dactyl: Family Lanceolidae
Eyes small, composed of few facets, often com- II P V longer than other P, ART 2 usually with ser-
pletely absent MD without molar. MX I with rate margin(s). Endopodites of all UR fused with
inner lobe. MXP often with inner lobes separate. basipodites. Inner lobes of MXP complete!~
Both male and fema le ANT I with strong conical fused, more or less reduced, or completely lack-
proximal flagellar ART, 0-3 other reduced distal ing: Family Scinidae
a11s (Fig. 4.37, 4.45). (Infraorder Physosomata) l l a P V not longer than other P, ART 2 with smooth
.. ....................... .................................................. 9 margins. UR with free endopodites and exopo-
8a Head large, longer than pereonite I. Eyes large (or dites. Inner lobes of MXP well developed, sep-
small to moderately large in Vibilia and arate. [Anterior pereonites of female much
Vibilioides) , occupying most of head surface inflated; male ANT II at most one-third as long
(excluding rostrum). MD commonly with molar. as ANT I]: Family Mimonectidae
MX I without inner lobe. MXP with inner lobe 12 ANT I inserted on ventral surface of head; fUS1
completely fused. If proximal flagellar ART of ART of male flagellum curved. Male ANT I!:
male ANT I is strong, it is oval, or flagellum with folded in zigzag form (Fig. 3d). (Superfamily
>4 ART (Infraorder Physocephalata) .............. 12 Platysceloidea) .......... ..................... .............. .. .2C
9 MD with 3-articulated palp ................ ............. 10 12a ANT 1 inserted on anterior surface of head. Male
9a MD without palp, or with rudimentary !-articu- ANT II not folded in zigzag form ................... 13
lated palp. [ANT I much longer than head]. 13 Male ANT I and II with multiarticulated flagel-
(Superfami ly Scinoidea) .................. ........... .... ! ! lum (Fig. 3b), but ART I of ANT 1 flagei!UI!:
Amphipoda 1157
may be gigantic (in Phronimidae, Fig. 4.119). 22a P VII with all ART MD without palp in female
(Superfamily Phronimoidea) ....... .................. 17 ..... ...... ..... .............. ......... ..................... ............ .23
13a Male ANT I with flage llum composed of large 23 Female ANT II short, composed of a few arti-
proximal ART, sometimes a few rudimentary cles. MD without palp in both sexes: Family
distal ART ....... ........... ..................................... 14 Anapronoidae
ANT I not inserted in groove on head; first ART 23a Female ANT II rudimentary or absent. MD with
of male flagellum lanceolate or conical; distal palp in male ................................... .................. 24
ART inserted terminally or absent (Fig. 3a). 24 P II more or less subchelate. Dactyl of P II sim-
(S uperfamily Vibilioidea) ............................... 15 ple .............. ........ .... ....... ........ .... ............. .... ..... .25
1- a ANT 1 inserted in groove on head ; first ART of 24a P II not subchelate. Dactyl of P II covered by
male flagellum conical, distal ART inserted sub- numerous setae: Family Tryphanidae
terminally (Fig. 3c): Fami ly Lycaeopsidae 25 P I simple or with smooth subchela. P V much
~ Cuticle thin and weak, musculature very weak. longer than VI UR III (and someti mes II) with
Head very large. rounded dorsally, ventral sur- endopodites fused with basipodites: Family
face flat, with dentate border. UR II absent; UR Lycaeidae
I and Ill endopodites fused with basipodites: 25a P I with strong serrate subchela. P V - VI of
Family Cystisomatidae approximately same length. Endopodites of all
:5a Cuticle dense, musculature strong. Head without UR free: Family Brachyscelidae
dentate border, rounded or sub-quadrang ular. 26 Mouth parts in form of a broad, short cylinder. P
UR all present, endopodites free ..................... 16 VII consisting (except CX) only of large ART 2,
Body not transparent. MD with palp. P II che- sometimes with 1 or 2 rudimentary distal arts:
late: Family Vibiliidae Family Platyscelidae
- Body transparent. MD with palp in male, not in 26a Mouth parts in form of a cone. P VII with all
female. P II simple: Family Paraphronimidae ART present (except in Thyropus): Family
P V without large subchela ............................. 18 Parascelidae
P V with large subchela .. ........ ..... ... ................. 19
P II chelate: Fami ly Hyperiidae Suborder Gammaridea (descriptions of gammari-
P II simple, body flat: Family Dairellidae dean genera modified partly from Barnard and
UR with free exopodites and endopodites. Head Karaman, 199 1)
subconical, narrowing, prolo nged ventrally:
Fami ly Phronimidae Family Eusiridae
3. UR composed each of single leaflike segment Body often with dorsal spines and carina (especial ly
only. Head globular: Fami ly Phrosinidae on pleon), sometimes smooth. Pleon well-developed.
Body not conglobate. P V and VI with ART 2 Rostrum variable, large, or absent. ANT I longer or
somewhat broadened, but never operculate. P VI shorter than ANT II, accesso1y flagellum absent or
with distal ART inse11ed on margin of ART 2, or uni-articulate, scale-like or elongate, someti mes fused
on medial surface a short distance back from with ART 3 of peduncle. Accessory flagellum of
margin .............................. ........... ..................... 2 1 ANT I in eusirids, when present, often poorly visible,
Body conglobate. P V and VI with ART 2 deve- requiring high magnification. ANT often with nume-
loped into broad operculum. P VI with distal rous calceoli in both sexes. Coxae medium-large or
ART inserted on m edial surface of ART 2, far small. MD with 3-articulated palp, molar usually tri-
back from margin ............................................ 26 turative, sometimes reduced or simple. Lower lip with
Body elongate or needle-like; head produced or without inner lobes. Shape of P I and II often dis-
mto distinct rostrum anterior to eyes: Family tinctive: strongl y subchelate, ART 5 very narrow,
Oxycephalidae bag-like ART 6 attached to very narrow margin, and
Body fairly compact; head globular. ...... ......... 22 thus ART 6 strongly flexible relative to axis of ART
P VII consisting (except CX) on ly oflarge ART 5; palm opposite articulation of ART 5 and 6; ART 5
::! •• 1 or 2 rudimentary distal ART MD with palp with narrow outgrowth in middle of posterior margin
in both sexes: Family Pronoidae (so-called "eusirid gnathopod", see Fig. 4.6, 4.7). PI
1158 Amphipoda
and II normal in other cases, strongly or weakly sub- ART 3 not produced, in males twice as short as ART
chelate; in former case, ART 6 often enlarged and 1 of main flagellum; accessory flagellum uni-articu-
broadened, lanceolate or elongate-oval, with palmar late or absent. Epistome not produced. Mandibular
margin almost equal to posterior margin of ART, molar columnar, triturative, ART 2 of mandibular
ART 5 with more or less well-developed posterior palp without lobes. Inner lobe of MX I with 2 apical
lobe. P V-VII often elongate. Exopodites of UR I and setae (I seta in Pacific species C. macrocephala).
II usually shorter than endopodites. T cleft or entire Inner lobe of MX II subequal in length to, but broad-
(in the calliopiids). Sixty five genera, mainly benthic. er than (sometimes slightly), outer lobe. Lower lip
Seven genera in plankton of South Atlantic. with small inner lobes. ex l not strongly produced
anteriorly. P I and II large, subchelate, with distinct
Key to the genera of Eusiridae palmar margins and palmar spines, not of eusirid
1 Mandibular molar nontriturative, conical... ....... 2 form; ART 5 shorter than 6, with strong posterior
Ia Mandibular molar triturative, columnar ............ 3 lobes. P III-VII elongate, simple, ART 4-6 each much
2 P I and II of eusirid form (but ART 5 w ithout longer than ART 2, dactyls long. Epimeral plate III
posterior outgrowth): Eusiropsis with smooth posterior margin, sharp posteroventral
2a PI and II not of eusirid form. [Inner plate ofMX corner. Exopodites of UR I-III shorter than endopo-
II broader than outer plate]: Eusirella dites. T cleft. Eight species, cold- or deep-water. Still
3 T entire (but apically dentate). Inner plate ofMX not found in the South Atlantic, but widely distribut-
II not broader than outer plate. [P I and II with ed in world ocean. C. longipes, which may be found
large posterior lobe on ART 5. Body smooth; in the region, is described here.
coxae short]: Stenopleura • Cleonardo longipes Walker, 1906 (Fig. 4.1 ).
3a T cleft (or incised). Inner plate of MX II broad- Length to 10 mm. Body transparent. Rostrum
er than outer plate ............................... .............. .4 slightly less than one half length of ART 1 of
4 P I and II of strong eusirid form. [P I not larger peduncle of ANT I. ART 2 of peduncle of ANT 1
than PIT. Accessory flagell um of ANT I present, 1-1.5 times as long as ART 1; ART l without any
elongate]: Eusirus teeth. Subchela in P I shorter, moderately broader
4a P l and II not distinctly eusirid ....... ..... .............. 5 than in P II. Posterior lobes of ART 5 ofP I-II tri-
5 P III-VII not elongate: Eusiroides angular, slightly narrower than main pa11 of ART
Sa P III-VII elongate ....... ....................................... 6 in P I, as broad as it in P IT, " heig ht" smaller than
6 CX I weakly produced anteriorly, body smooth length of base. ART 6 of P HI-IV l-1.5 times as
........................................................................... 7 long as dactyl. ART 6 of P 6 1.1-1.5 times as long
6a ex l strongly produced anteriorly, body carinate as dactyl. T very long, top reaching distal ends of
or toothed: Rhachotropis* [Pelagic and bentho- rami of UR III, deeply cleft. Live animals with
pelagic representatives of this genus have not small red eyes, which may be obscure in fixed
been reported from the South Atlantic, but are specimens .
common in the world ocean]
7 Palmar margin of subchela of P I and 11 faint, Genus Eusire/la Chevreux, 1908
straight, or slightly concave, w ithout spines (but Body slender, compressed, smooth. Rostrum small.
with setae). (Accessory flagellum of ANT I Eyes absent. ANT with numerous calceoli in both
never present): Harcledo sexes. ANT I longer than II. ART I of peduncle of
7a Palmar margin of subchela of P I and II distinct, ANT I as long as head; ART 2 longer than 1, ART 3
slightly convex, with palmar spines. (Acessory very short; accessory flagellum scale-like, uni-articu-
flagellum of ANT I !-articulate or absent): late, or absent. Flagellum of ANT Il much shorter
Cleonardo than peduncle. Molar poorly triturative or simple, sub-
conical. Inner lobe of MX I without setae, palp short.
Genus Cleonardo Stebbing, 1888 ART l longer than 2. Inner lobe ofMX TI broader bm
Body smooth. Rostmm small to large. Eyes very not longer than outer. Lower lip with inner lobes.
small or absent. ANT I longer than or subequal to Coxae very short, CX IV without posterior lobe, nm
ANT II; calceoli present. ART 1 of peduncle of ANT excavated . P Tand II not of eusirid fonn, strongly sub-
I subequal to head, ART 2 not shorter than ART l , chelate, ART 5 without posterior lobe. ART 6 of PI
Amphipoda 1159
, .. - lanceolate. P II larger than P 1, ART 6 elongate, Genus Eusiroides Stebbing, 1888

:llmost linear. Palms of P I and II almost as long as Body compressed, smooth, or weakly carinate. Eyes
posterior margins of ART 6, with palmar spines. PIll- reniform, sometimes surrounded by additional large
\ li elongate, simple. Epimeral plate III with smooth pigment spot. ANT I longer than 11. Accessory flagel-
;:nargins. Exopodites of UR 1-lll shorter than endopo- lum uni-articulate, from elongate to scale-like.
dites; rami with lateral and dorsal spines. T elongate, Mandibular molar strong, triturative, columnar. Inner
:left. Purely pelagic genus. Five species, 3 in the lobe of MX I with 1 medial, l apical seta, or fewer.
South Atlantic. Inner plates of MX II much broader, but not longer,
than outer. Lower lip with small inner lobes. CX 1 pro-
~ey to the species of Eusirella duced anteriorly, expanded ventrally (sometimes
ART 6 of PIT 1.5-2 times as long as ART 5 ...... 2 slightly). CX IV excavated, with posterior lobe. P T
la ART 6 of P II only slightly longer than ART 5: and II subchelate, not of eusirid fo1m, ART 5 with pos-
Eusirel/a multicalceola terior lobe, ART 6 broad. P III-VII simple, not elon-
CX I broadened distally. ART 6 subequal in PI gate. Epimeral plate Ill serrate or smooth. Exopodites
and P II: Eusirella elegans of UR I-ll slightly shorter than endopodites. T cleft.
~ CX I not broadened distally. ART 6 ofP II clear- Fifteen species, one in the South Atlantic plankton.
ly larger than ART 6 ofP I: Eusirel/ajlagella • Eusiroides stenopleura K. Barnard, 1932. Length
to 25 mm. Rostrum short. Faceted eyes reniform,
• Eusirella elegans Chevreux, 1908 (=Eusirella val- but within cuticle, surrounded by round patch of
diviae Schellenberg, 1926) (Fig. 4.2). Length redd ish pigment occupying nearly whole side of
about 5-l 0 mm. ANT II with 8-1 2 flagellar ART in head, dorsally almost contiguous with its counter-
adults. CX I broadened distally, anterodistal comer part (K. Barnard, 1932). Accessory flagellum of
projecting anteriorly. ART 6 of P I and 11 wider ANT 1 scale-like, fused with peduncular ART 3,
than ART 5. ART 6 of P II 1.5-2 times as long as almost unnoticeable. ART 2 of mandibular palp
ART 5. Distal part of anterior margin of P I and II expanded. All coxal plates not as high as the per-
convex. ART 6 almost equal in PI and P 11. Palmar eonites; coxae 1-IV wider than high. CX I sub-
::nargin of P I and II with spines and setae. quadrangular, not projecting below ventral margin
• Eusire/la flagella Andres, 1982 (Fig. 4.3). Length of head, slightly produced anteriorly. ART 2 of P
:!bout l 0 mm. ANT II with more than 15 flagellar VII wide, with broad posterior lobe, posterior mar-
.-\RT in adults, flagellum with only few calceoli gin smooth or finely serrate. Epimeral plate III
important character!). CX I subquadrate, not with smooth margin, posteroventral comer at
~adened distally. ART 6 of P 1 and TI scarcely almost 90, ventrally with row of submarginal
;der than ART 5. ART 6 of P II about J .5 times spines. T 50-70% cleft. Live animals red.
..:.5 long as ART 5. ART 6 of P II clearly longer
;ban ART 6 of P l. Very similar to, possibly syn- Genus Eusiropsis Stebbing, 1897
on~mous with, E. heterochela Birstein and M. Rostrum small. Eyes absent. Coxae very short. ANT
·:mogradov (1964) from the Indian Ocean (distin- subequal, bearing numerous calceoli. Accessory fla-
;-.rished almost alone by form ofCX I; see Birstein gellum scale-like. Mandibular molar weak, not
=nd Vinogradov, 1964 and Andres, 1982). columnar. Inner lobe of MX I without setae. Inner
• Eusirella multicalceolu (Thorsteinson, 1941) lobes of MX II broader but not longer than outer,
~1g. -+.4). Length to 15 mm. ANT II with 6-9 short without facial row of setae. Lower lip with inner
~ellar ART in adults. Both ANT 1 and II with lobes. P 1 and II subequal, of eusirid fo1m but with
~ia lly numerous calceoli (important charac- ART 5 lacking posterior lobe, moderately short, broad
z-~ . CX I not broadened distally. ART 6 of P I (only slightly longer than ART 4). PIll-VII elongate,
a&d II scarce ly wider than ART 5. ART 6 of P II simple; dactyls simple but setose on inferior margins;
ccl~· slightly longer than ART 5. Distal pat1 of ART 2 without anterior lobes. Exopodites ofUR I-III
~erio r margin of P I and II slightly concave. shorter than endopodites. UR III extended well
-.....~T 6 almost equal in P I and P II. Palmar mar- beyond UR I. T elongate, weakly cleft. Two species,
;= of P I and II with spines, without setae. one in the South Atlantic.
1160 Amphipoda
• Eusiropsis riisei Stebbing, 1899 (Fig. 4.5). Length • Eusirus antarcticus Thomson, 1880. Length to 15
to 12 mm. Body smooth. Mandibular molar not mm, in Antarctic waters to 36 mm. Pereonite VII,
triturative; ART 3 of mandibular palp longer than all pleonites, and urosomite T carinate; pereonite
ART 2. Dactyl of palp of MXP not spinose along Vll with weak dorsal tooth, pleonites I and II (but
inferior (inner) margin. Coxae very short, progres- not Ill) with strong dorsal teeth. Lateral lobes of
sively longer toward CX VII; CX IV not twice as head broadly rounded (as in E. perdentatus). Eyes
long as CX I. Live an imals with rose rings on P. pear-like (oval in young), nearly fused dorsally,
eyes larger in males than in females . ANT pedun-
Genus Eusirus Kreyer, 1845 cles calceolate. ANT I slightly longer than II,
Body often dorsally carinate or toothed. Rostrum reaching pleon. ART 2 of peduncle of ANT I as
small to large. Eyes ren iform or absent. ANT sub- long as, or slightly shorter than, ART 1; ART 1
equal, or ANT I longer than II. Coxae short or medi- with 3 sharp distal denticles; ART 2 with 3-5 den-
um-large. CX I not produced anteriorly, broadened tides. Accessory flagellum shorter than ART 3 of
ventrally. CX IV more or less excavated, with poste- peduncle of ANT I. ART 4 of P ITT-IV twice as
rior lobe. Mandibular molar strong, columnar, tritura- long as ART 5. CX IV as high as pereonite IV.
tive. Inner lobe of MX 1 with 1-2 apical setae. ltmer Posteroventral comer of epimeral plate ITT at
lobes of MX li broader but not longer than outer, approximately 90°, rounded; margins finely ser-
often with strong medial setae. Dactyl of MXP palp rate. T twice as long as wide; with exception of
not spinose along inferior (inner) margin. P I and TT large antarctic specimens, cleft a third of length or
alike, subchelate, of classic eusirid form. P III-VII more, in antarctic animals cleft one quatter or less.
elongate, simple. Posterior margin of epirneral plate • Eusirus perdentatus Chevreux, 1912 (Fig. 4.6).
III serrate or smooth. Exopodites of UR I-II shorter Length 30-67 mm. Head subequal to pereonites I-
than endopodites. UR Ill not extended beyond UR 1. II combined, lateral lobes broadly rounded.
T variable: elongate or short, cleft, incised or emargi- Pereonites V-VII, pleonites I-TTI with dorsal
nate. Twenty two species, mainly benthic, but a few spines, urosomite I carinate. ART 1 of peduncle of
deep-water planktonics. In cold Antarctic waters, ANT I with strong distal teeth on inner margin.
some benthic species also rise into the pelagic realm. small distal denticle on ventral margin. Ventral
A few of them were found in the Atlantic sector of the margin of ART 4 of peduncle of ANT II serrate.
Antarctic, at or near the southern border of the region Eyes large, pear-like, in males occupying main
(60-62°S), one species reaching subantarctic waters. part of lateral surface of head (often even fused
dorsally), in females not so large. ART 4 of P III-
Key to the species of Eusirus IV longer than 5 or 6. ART 5 of P V-VII only
1 Pleon segment lii with dorsal tooth .................. 2 slightly longer than ART 4.
I a Pleon segment lii without dorsal tooth (but may • Eusirus properdentatus Andres, 1979 (Fig. 4.7).
be carinate) ......................... ................... ............ 3 Length to 55 mm. Similar in appearance to E. per-
2 Flagellum of ANT II not much longer than ART dentatus. Pereonites V -VII, pleonites I-III with dor-
5 of peduncle. ART 5 of P V-Vll only slightly sal spines, urosomite I carinate. Eyes large, pear-
longer than ART 4: Eusirus perdentatus like. Lateral lobes of head subrectangular, rounded.
2a Flagellum of ANT II much longer than ART 5 of slightly produced. ANT I slightly longer than ANT
peduncle. ART 5 of P V-VII twice as long as H; accessory flagellum uni-articulate. Peduncle of
ART 4: Eusirus properdentatus ANT II reaching three quarters of length of pedun-
3 Anterior margins of ART 5-6 ofP (gnathopod) I cular ART 2 of ANT T. Flagellum of ANT 11 3.5
equal in length. (T cleft less than one fifth of times as long as peduncular ART 4 and 5 com-
length): Eusirus microps* [Inhabits Antarctic bined. Ventral part of ART of ANT I, dorsal part of
waters. May be found on southern border of flagellar ART, 4-5 peduncular ART of ANT II cal-
region. Length to 42 mm]. ceolate. Mouth parts as in E. perdentatus but with-
3a Anterior margins of ART 5 of P (gnathopod) I out distal dentic les on ART 2 ofpalp ofMXP, inner
almost twice as long as ART 6. (With exception margin of that ART nearly smooth. P I and II sub-
of large Antarctic specimens, T cleft a th ird of equal. ART 4 and 6 in P III-IV subequal, ART 5
length or more): Eusirus antarcticus sh01ter. Length of ART 4, 5, and 6 of P V-VII i.r:.
Amphipoda 1161
ratio 1:2:3. Epimeral plate III weakly denta te. Ra mi Stenopleura atlantica Stebbing, 1888 (Fig. 4.9).
of UR III subequal. T about 20% cleft, incised , top Length to 7 mm. With characters of genus .
reaching middle oframi of UR III.
Superfamily Lysianassoidea (=family Lysianassidae
Genus Harcledo Barnard, 1964 sensu lato)
3ody smooth, compressed. Rostrum medium. Eyes Peduncle of ANT I short, stout, ART 1 ofte n barrel-
~ -al or reniform. ANT subequal. ART 1 of peduncle like, ART 2 and 3 usually much shorter than I .
~:· A T I shorter than head , longer than ART 2 (and Accessory flagellum usually prese nt, more than 3-
-\.R.T 2 longer than ART 3); ART l of main flagellum articulated, rare ly absent. Pereon lacking processes,
:-;ce as long as ART 3 of peduncle; accessory flagel- urosomal segment I often with prox imal dorsal
.::m absent. Mandibular mo la r columnar, triturati ve; depression, di stal dorsal process. Body usually com-
-\.RT I of mandibular palp without any teeth, ART 2 pact (especially in benthic and necto-benthic forms),
=:lobed, ART 3 shorter than 2. Lower lip with inner chitin smooth, often dense. Mouth parts very variable,
..:ci>es. Inner lobe of MX 1 with I pinnate apical seta. often with more or less reduced mandibular molar and
==ner lobes of MX II broader but not longer than variable position of mandibular palp, some lobes or
- er. Coxae very short and slightly enlarged in row l- palps of maxillae and MXP also may be reduced. PI
"'11. CX I not produced anteri orly nor expanded dis- and II (gnathopods) usually smaller than others, P I
t:illy. P I like P II, not of eusirid form, subchelate simple or p rehensile, P II specifically elongate, slen-
.mout distinct palmar margin or p almar spines, ART der, minutely chelate or subchelate (rarely simple),
: much shorter than ART 6, ART 6 broadly lanceo- ART 3 elongate, ART 5 and 6 densely setose, dactyl
-re. P Ill-VII elongate, simple, ART 2 not anteriorly minute. Coxae usually long, coxae I to III occasional-
· te. ART 4-6 ofP V-Vll each 1. 5 times as long as ly reduced in size and/or hidden by fo llowing coxae.
"4'{T 2. Epimeral plate III smooth. Exopodites ofUR CX IV with indentation on posterior margin (rare in
-ill slightly shorter than e ndopodites. T elongate, other sp ecies), anterior margin of ex v occupying
- mted, cleft. Purely pelagic genus. Monotypic. this indentation, CX V often bilobate. Rami of UR Ill
• Harcledo curvidactyla (Pirlot, 1930) (Fig. 4.8). lanceolate, exopodite often 2-articulated, endopodite
- Length to 20 mm (Atlantic specimens to 15 mm).
With cha racters of genus. Barnard (1 964 a)
may be reduced in size. T cleft, emargi nate or entire,
long or short, ra rely absent. A bout 170 genera, main-
described the new genus Harcledo especially for ly benthi c. Ten genera in the South Atlantic plankton.
.\feteusiroides curvidacty la (= M . plumipes) (see "The Lysianassidae have been and are being divided
also Barnard and Karama n, 199 1) , but in many into groups by various taxonomists but no compre-
recent pape rs it is still assigned to Meteusiroides. hensive synoptic classificati on has appeared as yet"
(Barnard and Karaman, 199 1, P. 42 1).
Genus S tenopleura Stebbing, 1888
y s lender, smooth. Rostrum medium, thin . Key to the genera of L ysianassoidea
::;;eroventral margin of head not produced . Eyes 1 T absent. [Body globular· in females, more or
~fo rm. All coxae sho rt, CX I sharply produced less elongate in males. Integument weak] ......... 2
.E::::.."'tiorly. ANT I slightly longer than or equal to la T present ............ ......... .... ............... .......... .......... 3
-.L. "T II. Accessory flagellum a bsent. Mandibular 2 Maxilli pedal palp completely a bsent: Danaella
:ar co lumnar, triturative, ART 3 of mandibular 2a Max illipeda l palp reduced, but present.
-P almost as long as ART 2. Inner plate of M X TI [U rosome composed of three segments]:
_ broader than outer p late. Lower lip with inner Thoriella
-. P I and II (gnathopods) not of eusirid form, sub- 3 Both coxae I a nd IT (and occasionally coxae III
.::e:are. with large posterior lobe on ART 5; ART 5 too) distinctly smaller than following coxae,
~ 6 together not elongate but ART 5 much shorter largely hidden by them ........... .. ................... .. .. ..4
6: ART 6 not bent; dacty l subequal to ART 6. P 3a Coxae l may be small, but neither coxae II nor
-._' ll elongate. Epime ral pla te III smooth. III di stinctly reduced, both visible ................ .... 6
:podites of U R I-II shorter than endopodites. T 4 CX III small, partially covered by enlarged CX
=:.re. apically toothed. Pure ly pelagic genus. IV. ART 2 of P V with posterodistal sword-like
~typ ic . tooth, or strongly dentate: Cyphocaris
1162 Amphipoda
4a CX Ill large, not covered by CX fV. ART 2 ofP la Posterior margin of ART 2 of P V without
V without long tooth, not dentate ................ .... ..5 sword-like tooth but with posterodistal serrate
5 All P III-VII more or less subchelate. T deeply lobe: Cyphocaris richardi
cleft. Endopodite of UR III shorter than exopo- 2 Both margins of sword-li ke tooth of ART 2 of P
dite but not reduced : Paracyphocaris V smooth .. ............................................... .... ...... 3
5a P VI and VII not subchelate. T no more than 2a Both margins of sword-like tooth of ART 2 of P
25% cleft. Endopodite of UR III strongly V deeply dentate. [Sword-like tooth relatively
reduced: Metacyphocaris short, not longer than ART 3-5 of P V com-
6 CX I distinctly reduced in size, largely hidden by bined]: Cyphocaris anonyx
CX II, but d istal frontal corner may be free .... ..7 3 Sword-like tooth clearly shorter than ART 3-7 of
6a CX I not distinctly reduced in size, not hidden by P V combined ... .............................. ... ..... .......... A
ex n ..................................................................8 3a Sword-like tooth almost as long as ART 3-7 ofP
7 Inner plate of MX I with five or more setae. V combined: Cyphocaris challengeri
[Setae of inner plate of MX I mostly terminal; 4 Sword-like tooth not reaching posterior margin
molar subconical but extremely broad; MD with of coxae V: Cyphocaris faurei
deepS-shaped furrow between palp and incisor]: 4a Sword-like tooth clearly overreaching posterior
Eurythenes margin of coxae V: Cyplwcaris bouvieri*
7a Inner plate ofMX I with one or two immense fal- [Deep-water animals, in Atlantic Ocean found
cate setae: Hirondellea* [present in deep-water only in northem part, but also present in Indian
of northern Atlantic Ocean: H. brevicaudata and Pacific Oceans]
Chevreux, 1910, with short, weakly cleft T].
8 ART 7 (dactyl) of P I minute, shrouded in large Cyphocaris an onyx Boeck, 1871 (Fig. 4.1 0 ).
tuft of long setae or cirri attached to ART 6 and 7 Length to 14 mm. Pereonite I overlapping head.
........................................................................... 9 produced anteriorly, rounded or (rarely) sharp-
8a ART 7 of P I clearly visible, not shrouded. [P I pointed, curved. Eyes absent. Convex anterior mar-
subchelate. Coxae I-IV high. T cleft]: Orchomene gin of CX IV covering CX III and posterior half of
group CX II. P III-V weakly subchelate. ART 2 of P \"
9 MD lacking molar: Scopelocheiropsis with posterior sword-like tooth; both margins of
9a MD bearing molar. [P II clearly subchelate. this tooth deeply serrate, tooth relatively short, not
ART 2 of P I linear, its tissue not glandular]: longer than ART 3-5 of P V combined. Posterior
Paracallisoma margins of P V-VII dentate, each with 8-1 2 denti-
cles. Posteroventral comer of epimeral plate III at
Genus Cyphocaris Boeck, 1871 ca. 90. Live animals red, orange in alcohol.
Body slender, smooth. Integument weak. Pereonite I • Cyphocaris chal/engeri Stebbing, 1888 (Fig.
more or less produced anteriorly, overlapping head. 4.11 ). Length to 13 mm. Pereonite I produced
ANT long. Coxae I-III small, pattly covered by CX anteriorly, overlapping head, form changing from
IV. Epistome not strongly produced. Mandibular cone-like (in fi rst turn in juveniles) to subrectan-
molar of medium size, columnar, often triturative; gular or even rounded. Eyes large, reniform. CX
palp 3-articulated, strong, attached at level of molar. P IV large, with strongly convex anterior margin.
I and II weakly subchelate. ART 2 of P V often with covering all coxae I-III. CX V wider than high.
sword-like tooth on posterior margin. P III-V simple ART 2 ofP V with posterodistal sword-like tooth.
or weakly subchelate, P VI-VII simple. Posteroventral both margins of tooth smooth, length subequal to
comer of epimeral plate III with small, sharp to distal part of P (ART 3-7 combined). In live ani-
approximately right-angled process. Endopodite of mals "swords" projecting laterally. Beside;;
UR III well-developed. T long, deeply cleft. Purely sword-like tooth, posterior margin of ART 2 of P
pelagic genus. Ten species, 4 in the South Atlantic. V with 2-3 denticles. ART 2 of P VI-VII with
strongly serrate posterior lobe, broader in P \1
Key to the species of Cyphocaris than in P VII. Posteroventral corners of epimeral
1 ART 2 of P V with posterodistal sword-like plates I-III produced posteriorly, sharp. Live ani-
tooth ................. ................. .......... .... ................... 2 mals pale-brown.
Amphipoda 1163
Cyplwcaris faurei Barnard, 1916 (Fig. 4. 12). Genus Eurytltenes Smith in Scudder, 1882
Length to 30 mm. Pereonite I inflated, overlapping Coxal plate I small, rounded, much shorter than coxal
posterior part of head, but not covering all dorsal plate II, but anteroventral corner not covered by coxal
surface of head. Eyes pear-like, broadened ventral- plate II. ANT I much shorter than ANT II, ART 1 of
ly. CX TV large, moderately narrow, but with peduncle moderately small, weakly inflated, accesso-
strongly convex anterior margin, covering coxae II- ry flagellum long, approx. I 0-articulated. Epistome
III but not I. CX V significantly wider than high. large, globular (inflated) but not longer than upper lip.
ART 2 of P V with posterodistal sword-like tooth, Eyes large, expanded ventrally. Mandibular molar
both margins of tooth smooth, clearly shorter than strong, pyramidal, triturative, distal rib covered with
ART 3-7 of P V combined, not even reaching pos- setae; palp attached on level of molar. Inner lobe of
terior margin of CX V. Besides sword-like tooth, MX I with ca. I 0 pinnate setae, palp weak, with nar-
posterior margin of ART 2 of P V smooth. ART 2 row ART 2. lnner lobe of MX II much shorter than
of P VI-VII with finely sen·ate posterior lobes. outer, pointed, with row of pinnate setae on inner mar-
Posteroventral comers of epimeral plates I-III sharp- gin. Outer lobes ofMXP broad, reaching distal end of
ly produced posteriorly. Live an imals red-rose. ART 2 of palp; palp 4-articulated. P I with small but
Cyplrocaris richardi Chevreux, 1905 (Fig. 4.13). clear subchela formed by dactyl and distal margin of
Length to 56 mm (female), 40 mm (male). ART 6. Epimeral plate III with rounded posteroven-
Pereonite I overlapping head, produced anteriorly, tral corner. Rami of UR III broadly lanceolate, with
,·ery vatiable: sometimes anterior end with long numerous long setae on inner margins, exopodite with
narrow hom-like process, sometimes (especially small ART 2. Telson large, triangular, deeply cleft.
in large specimens from Antarctic waters) like Gills without transverse folds. Two species.
short cone with rounded top. Eyes small, oval,
faint. Convex antetior margin of CX IV covering Key to the species of E urythenes
CX III and distal part of CX IT. CX V wider than 1 Dactyls of P III-V short (about 30% of length of
high. Posterior margin of ART 2 of P V without ART 6); ART 3-7 on P Til, combined, twice as
sword-like tooth, with posterodistal lobe, posterior long as ART 2: Eurythenes gryllus
margin of ART and both margins of lobe strong or la Dactyls of P III-V long (about 60% of length of
finely serrate. Posterior margin of ART 2 ofP VI- ART 6); ART 3-7 on P III, combined, five times
\ "Il serrate. Posteroventral corner of cpimeral as long as ART 2: Eurytlrenes obesus
plate III at nearly 90, slightly produced posterior-
ly. Live animals red-orange. • Ewythenes gryllus (Lichtenstein in Mandt, 1822)
(Fig. 4.15). Length to 154 mm. Very like E. abe-
~us Danaella Stephensen, 1925 sus but distinguished from it by characters given
~e body more or less slender, female globular, with in key. Live animals red, reddish-brown, or pale,
~~ nites lll-VI especially enlarged (IV and V with white eyes. In water column E. gryllus as a
~ _emely enlarged). Accessory flagellum of ANT 1 ru le represented by comparatively large specimens
~nt. Epistome not prominent. MD lacking molar (length to 90 mm). Juveniles and large(> 100 mm)
· palp. Outer lobes of MXP large, opercular, cover- females inhabit near-bottom layers, where they
-: small curved inner lobes; palp completely absent. can even burrow to avoid danger. They are ben-
• simple, with very short dactyL P II simple. Coxae tho-pelagic scavengers, common in baited bottom
--small, hidden by subsequent coxae. Urosome con- traps. See also latest atticlcs on E. gryllus in bait-
s:s::ng of two large segments with vestigial urosomite ed traps at various heights above bottom in north-
:nner rami ofUR I-II shmt, spiniform; UR III like ern Atlantic and adjacent waters: Thurston, 1990;
.:::.-ngial microscopic bud. T absent. Purely pelagic Vinogradov eta/., 1996; and Christiansen, 1996 .
==.IS. Monotypic. • Eurythenes obesus (Chevreux, 1905) (Fig. 4. 16).
Danaella mimonectes Stephensen, 1925 (Fig. Length to 60 mm. Very like E. gryllus but distin-
-.14). Length to 20 mm. With characters of genus. guished from it by characters given in key. Live
Parasitic on gelatinous planktonic an imals. animals red, with white eyes.
1164 Amphipoda
Genus Metacyphocaris Tattersall, 1906 margin with large curved tooth on posteroventral cor-
Integument thin. Head deformed, short, with frontal ner. Urosomite I with dorsal carina. T varying from
process. ANT short, ANT I attached to ventral surface triangular to rounded, from deeply cleft to entire.
of frontal process of head. Accessory flagellum of About 90 species, mainly benthic or near-bottom
ANT I uni-articulate. MD with small smooth molar; scavengers. A few bentho-pelagic or pelagic species
mandibular palp rudimentary, 2-atticulated, or absent. (e.g. 0. pelagica, see Birstein, Vinogradov, 1960) .3- 1
Coxae I and II small , partially hidden by CX Ill. P I have been reported from plankton samples as well as
(gnathopod I) simple, with subequal ART 5 and 6. P II bottom traps (see Thurston, 1990 and Low ry.
simple. P Ill-V subchelate, P VI-VII clearly simple. Stoddart, 1994). Only one such species was reported
Exopodite ofUR II elongate, 2-atticulated; endopodite from the South Atlantic. Two others in northern trop-
very short, scale-like. T elongate, weakly (no more ical Atlantic waters may well cross the equator.
than 25%) cleft. Purely pelagic genus. Monotypic.
Metacyphocaris helgae Tattersall, 1906 (Fig. Key to species of the Orchomene group
4.17). Length to 17 mm. With characters of genus. The generic status of the species treated here follows
Animals orange or red in alcohol. Parasitic animals, Lowry and Stoddart (1994). They belong to a group of
even on "hard" (not gelatinous) pelagic animals. about 14 similar species recognized as follows: later-
al lobes of the head with rounded tops; ART 5 of P I
Orchomene group with narrow posterodistal lobe; ART 5 of P II stout:
This group of taxa has been the subject of conside- ART 2 of P III ordinary, not proboscoid; posterior
rable controversy and instability. The lysianassoid margin of epimeral plate liT smooth, posteroventraJ
genera Orchomene, Orchomenella, Orchomenopsis, corner rounded or at approximately 90°; dorsal carina
Allogaussia, Tryphosa are similar, but subsequent
description of many intergradational species led
of urosomite I hump-like, proboscoid, partly ovelap-
ping urosomite IT; rami of UR III setose (even weak-
•
Barnard ( 1964b) to regard all these genera as syno- ly) in females, exopodite 2-arriculated; T spinose.
nyms of Orchomene. Since then, some taxa have been deeply cleft (see Shulenberger and Barnard, 1976).
judged invalid, some "old" orchomenoid species
groups have been elevated to subgeneric or generic Chela of P II with palmar margin of ART 6
rank, new genera and subgenera have been described deep ly excavated, hook-like, fo rming clear.
(Abyssorchomene, Orchomenyx), vari.ous species rounded-tr iangular opening between it and dac-
have been redistributed among old and new genera tyl, distal part of posterior ("inner") margin of
and subgenera, and the process continues to the dactyl with setule-like long serrations, dactyl
present (see Barnard and Ingram, 1990; Barnard and reaching top of process of palmar margin of
Karaman, 199 1; Lowry and Stoddart, 1994). ART 6. Inner plate of MX II 4 times as long as
wide, only slightly shorter than outer lobe. (Eyes
The following characters d istinguish the gro up. All easily visible): Orchomenella gerulicorbis
coxal plates large and high. Accessory flagellum of la Chela of P 11 of another type. Inner plate of MX
ANT I present, approximately 5-articulated. Epistome ll about twice as long as wide, 1.5 ti mes as short
blunt, produced anteriorly. Head with triangular later- as outer lobe. (Eyes visible in live animals but
al lobes. Mouth parts bundle not projecting conically may be colou rless and inconspicuous in alcohol
below head. Mandibular mo lar weak, rounded, poorly ............................... ............................................2
but distinctly triturative; palp thin, attached proximal 2 ART 6 of P II 1.5 times narrower than ART 5.
to molar. Palp of MX I 2-articulated, inner lobe with posterodistal process pointed, produced axially.
2 strong ap ical setae. Lobes ofMX II subequal, long, palmar margin not excavated; dactyl of P II as
narrow. Palp of MXP 4-articulated . Upper lip never long as, or longer than, palmar margin of ART 6:
projecting conspicuously in front of epistome. P I Orcltomenella abyssorum
with small, clear subchela formed by dactyl and distal 2a ART 6 of P II as broad as ART 5, posterodistaJ
margin of rectangular A RT 6. ART 2 of P III not corner at approxi mately 90, dactyl of P II short-
deeply indented. Epi meral plate III with posterior er than palmar margin of ART 6: Orchomenella
margin serrate to smooth, or with concave posterior tlistinctus
Amphipoda 1165
• Orchomenella abyssorum (Stebbing, 1888) (Fig. shorter than pereonite I; lateral lobes well-developed.
4.18). Length to 13 mm. Eyes long, lunate, colour- Coxal p lates I-III shorter than pereonites I-III.
less in alcohol, glandular, onunatidia not evident; Mandibular molar small, attached prox imally; palp
about 40% as high as head. ART 1 of mandibular attached at level of molar; incisor smooth. Inner plate
palp short, subquadrate; ART 2 long, linear; ART ofMX I strongly setose; palp 2-articulated, large. ART
3 lanceolate, slightly less than half length of proxi- 2 of P !linear, its tissue not glandular. P IT clearly sub-
mal ART combined. Epimeral plate TU smooth, chelate (not chelate!). T cleft. Three or fo ur similar spe-
with posteroventral corner at approximately 90° cies. Only one species in southern Atlantic plankton.
(posterior margin very weakly excavated distally, • Paracallisoma alberti Chevreux, 1903 (Fig. 4.21 ).
with weak, blunt tooth on corner). Dorsal process Length to 26 mm. Eyes absent (but live animals
of urosomal segment I weakly overlapping next may have faint pigment spots). Epistome slightly
segment. Endopodite ofUR III reaching top of, but convex anteriorly. CX I variable (I to 2 times as
not exceeding, main (fust) ART of exopodite. See hig h as broad), may narrow slig htly distally.
also characters given in key and in preface to key. Posteroproximal excision of CX IV less than half
• Orchomenella distinctus Birstei n et M . Vinogra- width of coxal plate. ART 6 and 5 of P 1 subequal.
dov, 1960 (Fig. 4.19). Length to 18 mm. Eyes Posterodistallobes of ART 2 ofP VI-VII exceeding
long, flask-shaped, very pale pink, mostly glandu- ART 3. T cleft, with sub-apical outer denticle on
lar, ommatidia not evident; about 60% as high as each half. L ive animals orange to d irty-white.
head. ART 1 of mandibular palp shoti, subquad- Typical bentho-pelagic scavengers, conunon in
rate; ART 2 long, linear; ART 3 lanceolate, half baited traps. But animals also rise into water colwnn
length of proximal ART combined. Epim eral plate and may be fatmd in ordinary plankton samples.
III smooth, broadly rounded posteroventrally.
Dorsal process of urosomal segment I weakly Genus Paracyphocaris Chevreux, 1905
overlapping next segment. Endopodite of UR Ill Body spindle-li ke, smooth. Pereonite I overlapping
exceeding length of ma in (first) ART of exopo- shott, high head, pointed anteriorly. ANT I shoti er
dite. T with 0-2 pairs of dorsal spines. See also than peduncle of ANT II; accessory flagellum uni-
characters given in key and in preface to key. For articulate. MD lacking mola r; mandibular palp
a more detailed description of this species refer to attached medially, in Atlantic animals palp 3-mi icu-
Barnard and Ingram (1990). lated (but reduced in Pacific species P. distinctus).
• Orchomenella gerulicorbis (Shulenberger et Coxae I and II small, partially hidden by CX III. P I
Barnard, 1976) (Fig. 4.20). Length to 12 nun. Eyes (gnathopod) simple or very weakly subchelate, P II
\\ithout ommatidia, composed of pigment granules, nearly simple, ART 6 shorter than ART 5. P III-VI
faint, reddish-brown in life, magenta in preserva- clearly subchelate, P VIT weakly subchelate.
tiYe, fonning irregular crescent paralleling antero- Endopodite of UR III shorter than exopodite, but not
dorsal margin of lateral lobe of head; about third as reduced. T elongate, deeply cleft. Purely pelagic
high as head. ART 1 of mandibular palp short, subgenus. Three species, one in South Atlantic.
quadrate; ART 2 long, linear; ART 3 lanceolate, • Paracyplwcaris praedator Chevreux, 1905 (Fig.
half length of proximal ART combined, or less. 4.22). Length to 11 mm. ANT slender. ART 4 of
Epimeral plate III smooth, with rounded postero- peduncle of ANT TI shmter than ART 5.
,-entral corner at ca. 90°. Dorsal process of uroso- Mandibular palp 3-alticulated, shmter than body
mal segment I weakly overlapping next segment. of MD. ltmer plate of MX I with 3 strong pinnate
Endopodite of UR ill exceeding length of main setae. Both ART 5 and 6 ofP I about twice as long
first) ART of exopodite. T with 3 pairs of dorsal as wide. ART 2 ofP V-VU with rounded postero-
;;pines. See also characters given in key and in pre- distal lobes. Posteroventral corner of epimeral
face to key. For a more detailed description of this plate III blunt. Endopodite of UR III shmier than
species refer to Shulenberger and Barnard ( 1976). ART I of exopodite. T 2 times as long as broad,
deep ly cleft. Live animals red (no colour loss in
Ge::us Paracallisoma Chevreux, 1903 alcohol), with large drops of oil inside of pereon.
~:- - (dactyl) of PI minute, slu·ouded in a large tuft Anin1als may act as parasites of large pelagic crus-
- .ong setae or cini attached to ART 6 and 7. Head taceans. For example, P. praedator can feed upon
1166 Amphipoda
eggs of shrimp marsupium (see Bowman and (but see Halice macronyx). ANT I-II well-developed.
Wasmer, 1984). ANT I with atticulated accessory flagellum, often
sexually dimorphic; proximal ART of main flagell um
Genus Scopelocheiropsis Schellenberg, 1926 occasionally fused. Mouth patts under the head col-
ART 7 (dactyl) ofP I minute, shrouded in a large tuft lectively form a cubical, only exceptionally a conical.
of long setae or cirri attached to ART 6 and 7. MD body. MD lacking molar, mainly asymmetrical, palp
lacking molar (important character), palp attached variously absent to 3-articulated. Seventeen genera.
somewhat distally; incisor smooth. Inner plate of MX mainly benthic, only one represented in the South .. J
I strongly setose (9-1 0 pinnate setae); palp 2-articulat- Atlantic plankton.
ed, with apical row of strong spines. Posterodistal cor-
ner of ART 6 ofP II at approximately 90°. Exopodite Genus Halice Boeck, 1871
of UR III 2-articulated. T deeply cleft. Only one spe- Rostrum well-developed. Eyes absent. Coxae I-IV
cies in South Atlantic plankton. (Another, still unde- broader than long. Urosomites with dorsal spines
scribed, species of Scopelocheiropsis was found in (short and blunt on urosomite I, sharp on urosomite
Mozambique Channel, Indian Ocean). II). ANT 1 shorter than II. ART 2 of peduncle of ANT
• Scopelocheiropsis abyssalis Schellenberg, 1926 I shorter than ART 1, ART 3 shotter than ART 2:
(Fig. 4.23). Length about 5 mm. Mandibular incis- proximal flagellar ART fused in males, may be free in
or smooth, narrow, almost straight. In Atlantic females; accessory flagellum well-developed, 3-attic-
specimens, ART 4 of maxillipedal palp described ulated. Mouth parts grouped quadrately. MD with
as vestigial, but in Antarctic specimens this ART smooth or slightly serrate incisor, both with /acinia
nonnal, dactyl-like (in second species of genus, mobilis, mandibular palp present, 3-atticulated. MX I
see above, this ART also normal). Coxal plate I with both lobes, 2-articulated palp. MX II with espe-
only slightly widened distally. In P I and II (gna- cially long, narrow Jobes, inner lobe somewhat short-
thopods) ART 5 equal to ART 6; ART 6 of P II er than outer. MXP with reduced inner lobes, with
twice as long as wide. ART 5 ofP III and IV with- apical setae, palp 4-atticulated, nearly as long as inner
out anterodistal process; ART 6 longer than ART margin of outer lobe. PI and fT simple, weak, without
4 and 5 combined; P sub-prehensile. ART 2 of P posterior lobes in ART 5, dactyls simple, or rarely
V more or less slender. with their distal section spinose and the base unarmed.
ART 5 ofP II expanded. P V-VII long, slender, dac-
Genus Thoriella Stephensen, 1915 tyls slender and styliform. Rami of UR Ill elongate-
Body more or less slender in males, probably globular oval with setae on inner margin, exopodite with small
in adult females. Accessory flagellum of ANT I distal ART. T cleft 40% or more. Twenty two species,
absent; flagellum of ANT II with especially inflated mainly benthic; 2 in the South Atlantic plankton.
ART. Epistome broadly rounded, produced anteriorly.
Mandibular molar reduced, palp absent. MXP opercu- Key to the species of Halice
lar with inner plates apparently large and subrectangu- I Head formed into blunt anterior hood covering
lar, outer plates narrower and triang ular; palp appar- base of ANT I. ART 6 ofP I longer than ART 5:
ently composed of sing le large opercular ART with Halice macro11yx
deep lateral incision. P I simple, short; P II simple. 1a Head normal in appearance, with projecting lat-
Coxae small, short, not touching serially; CX IV with- eral lobes. ART 6 of P I subequa l to ART 5.
out posterior excavation. Urosome with all segments. [ART 3 of mandibular palp longer than half of
UR III lacking rami. T absent. Purely pelagic genus. ART 2. Inner plate of MX I with sharp comers.
Monotypic. with single pinnate seta): Halice secunda
• Thoriella islandica Stephcnsen, 19 15 (Fig. 4.24).
Length to 20 mm. With characters of genus. • Halice macronyx (Stebbing, 1888) (Fig. 4.25).
Parasites of gelatinous planktonic animals. Length 9-10 mm. Head unique for Halice, formed
into blunt anterior "shark nose" (term of Barnard.
Family Pardaliscidae 1969). ART I of main flagellum of ANT 1 con-
All coxae short. Rostrum developed, head usually joint (i.e., composed of several fused ART), coni-
normal in appearance, with projecting lateral lobes cal, equal in length to at least five of following
Amphipoda 1167
ART. ART 3 of mandibu lar palp longer than half 4a T cleft. [ART 5 of peduncle of ANT II very elon-
of ART 2, but shorter than whole ART. ART 6 of gate. Mandibular incisor smooth]: Euandania
PI longer than ART 5. Dactyls ofP I-II with their
distal section spinose and the base unarmed. ex Genus Andaniexis Stebbing, 1906
IV large, trapezoidal. Posteroventral corner of epi- Body smooth or carinate. Head with eye spots. ART 1
meral plate III acute. Urosom ite II with spme. of flagellum of ANT I almost as long as peduncle.
Peduncle ofUR I long and narrow. ART 4 of peduncle of ANT II much shorter than ART
• Halice secunda (Stebbing, 1888) (Fig. 4.26). 5. Mandibular incisor smooth. Palp of MX I 2-articu-
Length 12-13 mm. ART 1 of peduncle of ANT I lated. Outer plate of MX II not geniculate or gaping.
without outgrowth; accessory flagellum nearly ART 2 of palp of MXP not produced. ART 2 slender
equal in length to ART I of main flagellum; ART in P III, broad in P IV and V. ART 2 of P VI not
1 of main flagellum composed of several fused expanded, or slightly expanded. T entire. Nine spe-
ART, conical, equal in length to at least five offol- cies, two in plankton of the South Atlantic.
lowing ART. ART 3 of mandibular palp longer
than half of ART 2, but sh01ter than who le ART. Key to the species of A ndaniexis
Inner lobe of MX I with sharp corners, with single 1 ART 6 ofP II shorter than ART 5. Posterior mar-
pinnate seta. Coxal plates tetragonal. ART 5 and 6 gin of ART 2 of P VII serrate Andaniexis abyssi
of P I subequal. ART 4 and 5 of P ITT and IV 1a ART 6 ofP II longer than ART 5. Posterior mar-
expanded, flatly oval. ART 2 of P V-Vlll broadly gin of ART 2 ofP VII smooth Andaniexis aus-
oval. Urosom ite II with spine. Peduncle of UR 1 tralis
long and narrow. T cleft no farther than half\vay.
Live animals yellow-orange. • Andaniexis abyssi (Boeck, 1871) (Fig. 4.27).
Length about 7 mm. Rostrum absent. Head with
Family Stegocephalidae rounded lateral lobes, large quadrate white eye
Animals much higher than wide, lens-like. CX I never spots. Accessory flagellum of ANT I as long as
hidden by CX II. Coxae forming lateral shield; CX I- ART 1 of main flagellum. Ventral margin of CX
III high and narrow, CX IV broad, with posteroproxi- IV almost straight, posteroventral corner rounded.
mal excavation, CX V-VII much smaller, practically Posterior margin of ART 2 of P V and VI serrate.
whole CX V situated in excavatio n of CX IV. Epimeral plate III with a smooth prolonged poste-
Integument smooth, often dense. Mouth patts project- rior edge; posteroventral corner angular, not pro-
ing in a conical bundle below head. MD lacking palp duced. T with quadrate proximal, triangular distal
and molar. Accessory flagellum of ANT I 1- or 2- parts, smooth, without setae. Bentho-pelagic.
articu lated. P I and II (gnathopods) feeble, simple or • Andaniexis australis K. Barnard, 1932 (Fig.
weakly subchelate. Nineteen genera, mainly benthic; 4.28). Length 4-9 mm. Resembling A. abyssi
5 in plankton of South Atlantic. except in fo llowing particulars: ART 6 ofP II not
very slender, almost as wide as, and scarcely long-
Key to the genera of Stegocephalidae er than ART 5, subequal in length to ART 6 of P
Outer plate of MX II absent. [Mandibular incis- I; posterior margin has a very slight concavity dis-
or strongly dentate]: Bathystegocephalus tally which is defined by fairly long, stout spine
Ia Outer plate ofMX II present... .... ............. ......... 2 against which dactyl impinges. Posterior margin
1 Outer plate of MX II attached to lateral process of ART 2 of P IV and V perfectly smooth, entire.
from base of inner plate. [T cleft. Palp ofMX I 2- White eye spot present, apparently even larger
articulatc. Mandibular incisor strongly dentate]: than in A. abyssi. Bentho-pelagic.
Phippsiella
2a Outer plate of MX II appressed to inner plate .... 3 Genus Bathystegocephalus Schellenberg, 1926
3 Palp of MX I 2-articulate. [Mandibu lar incisor Integument dense, smooth. Eyes absent. ART I of fla-
smooth. P IV simple]: Andaniexis gellum of ANT I scarcely sh01ter than peduncle. ART
3a Palp of MX I 1-atticulate ....... .......................... .4 4 of peduncle of ANT II shorter than ART 5.
.! T entire. [ART I of main flagellum of ANT I Mandibular incisor dentate. Palp of MX l uni-articu-
much longer than peduncle. Mandibular incisor late. MX II completely lacki ng outer plate. ART 2 of
smooth]: Parandania palp ofMXP not produced. ART 2 slender in P III-IV,
1168 Amphipoda
broad in P V. ART 2 of P VI not expanded. T cleft one ART 2 s lender in P III, broad in P IV and V. ART ~
third. Monotypic. of P VI expanded. Posterior lobe of ART 2 of P \11
• Bathystegocephalus globosus (Walker, 1909) rounded below. T cleft more than 50%. Ten species.
(Fig. 4.29). Length to 12 mm. With characters of Only one in the South Atlantic plankton.
genus. Live animals intensely dark. • Phippsiella nipoma J. Barnard, 1961 (Fig. 4.3 1).
Length to 20 mm. Left MD Jacking lacinia mobi-
Genus Euandania Stebbing, 1899 lis. Rostrum poorly developed. Epimeral p late III
Body smooth. Eyes absent. ART I of flagellum of ANT with a smooth, prolonged posterior edge; postero-
I longer than peduncle, ART 4 of peduncle of ANT II ventral comer angular, not produced, lacking evi-
twice as short as ART 5. Mandibular incisor smooth. dence of a notch.
Palp of MX I uni-articulate. Outer plate of MX II not
geniculate or gaping. ART 2 slender in P III, broad in Family Synopiidae
P IV and P V. ART 2 ofP VI expanded. T as broad as Head as tall as, or taller than long, except rarely almost
long, cleft one third. 2 species. Only one species, E. as long as pereonites I-III combined, galeate or with
gigantea, in plankton of region: similar species, E. plow-shaped protuberance on forehead, with down-
nonhiata Andres (1985), found near South Shetland turned rostrum. ANT I with multi-articulate accessory
Islands (63°25'S, 64°36' W, see Andres, 1985). flagellum, base of primary flagellum composed of se-
• Euandania gigantea (Stebbing, 1883). Length to vera l fused ART in males. MD ordinarily with weak
50 mm. ANT I flagellar ART I much longer than palp, but tumid in Synopia. Inner lobes of lower lip
peduncle. ART 5 of peduncle of ANT II longer separate, outer lobes frequently notched. Mandibular
than ART 4. Inner (medial) margin of lower lip molar well-developed. PI and II weakly subchelate or
with small protuberance near distal apex, ART 2 simple. Coxae more or Jess long, CX IV never much
ofP VI broad. ART 2 ofP VII broad, tapering dis- larger than CX HI. T elongate, ra rely short (for exam-
tally. Urosomite III shorter than basipodite of UR p le, in Synopia). Fifteen genera, mainly benthic. Only
III. T widely cleft. As preserved, integument very one genus in the South Atlantic plankton.
flimsy, colour olive-brownish.
Genus Synopia Dana, 1852
Genus Parandania Stebbing, 1899 Lateral lobes of head not sharp. Eyes large. Small sec-
Integument moderately weak. Head very short. Eyes ondary eye on each side, at base of ANT I. MD of
absent. ANT 1 as long as peduncle of ANT II. First medium size, with medium-large molar, triturative, or
ART of flagellum of ANT I longer than peduncle of swollen and pillowlike, but poorly triturative.
ANT I. ART 5 of peduncle of ANT 2 twice as long as Mandibular palp extremely tumid. CX III strongly
ART 4. Mandibular incisor smooth. Palp ofMX I uni- expanded distally, with strong posteroproximal exca-
articulate. Outer plate of MX II not geniculate or gap- vation; CX IV small, situated in posterior excavation
ing. ART 2 of palp of MXP not produced. P I and Il of ex Ill (an unusual character, commonly observed
simple. ART 2 slender in PIll, broad in P IV-V. ART in coxae IV and V in some other gammarideans).
2 of P VI not expanded. T entire. Purely pelagic ART 6 of P 1 and II slender or tumid, subovate, not
genus. Monotypic. linear, may bear marginal setae, lacking strong spines.
• Parandania boecki (Stebbing, 1888) (Fig. 4.30). Dactyl of P II small. Pleonites 1-flT not denticulate.
Length to 26 mm. With characters of genus. Live Basipodite of UR lli short, subequal to T. T short.
animals brown, ordinarily do not lose colour in cleft or entire, in the latter case distally trifoliate.
preservatives. Feeds on jelly-fish. Purely pelagic (epipelagic) genus. Nine species, 3 in
the tropical South Atlantic. Genus poorly repotted in
Genus Phippsiella Schellenberg, 1925 literature.
Body smooth. Eyes absent. ART 1 of flagellum of
ANT I longer than peduncle in males, shorter in Key to the species of Synopia
females. Mandibular incisor dentate. Palp of MX 1 2- 1 ART 2 of P lll-IV rectangular, or with quadrate
articu lated. ART 4 of peduncle of ANT II scarcely posteroventral corners: Synopilrgracilis
longer than ART 5. Outer plate of MX II geniculate Ia ART 2 ofP III-IV ovate, or with rounded poste-
and gaping. ART 2 of palp of MXP not produced. roventral corners ........... ... .............. .................... 2
Amphipoda 11 69
2 Flagellar ART 2, and all following of ANT T, with Vinogradov et al., 1982). ln this volume we follow
long hairs: Synopia caraibica* [Known from Vinogradov el al. ( 1982). Family and genus keys are
Caribbean Sea] modified from Bowman and Gruner (1973) and
2a Flagellar ART 2, and all follow ing of ANT T, Yinogradov et al. (1982), species keys from
without long hairs .............................................. 3 Vinogradov et a/. ( 1982), with a few exceptions men-
3 Telson scarcely longer than broad, apico-lateral tioned below. Family and genus diagnoses are modi-
margins smooth, each apex with one stout spine: fied partly from the same authors.
Synopia scheelana
3a Te lson much longer than broad, ap ico-lateral Infraorder Physosomata
margins serrate, each apex with at least 2 setae: Head small, usually shorter than pereonite I. Eyes
Synopia ultramarina small, composed of few facets, or absent. Integument
weak. MD without mo lar, MX I with inner lobe, MXP
• Synopia gracilis Dana, 1853. Length 5-6 mm. often with separate inner lobes. ANT I with strong,
Body slender, without convexity along back. Head conical proximal flagellar ART, 0-3 rudimentary api-
triangular. Eyes more or less round. ANT I with cal ART in both males and females. Mainly deep-
accessory flagellum equall ing length of ART 1 of water animals. All Physosomata are associated with
main flagellum (the latter composed of several various gelatinous plankton, mainly with deep-water
fused ART). Segment r ofpereon shorter than seg- jellyfish and siphonophores.
ment II, VI as long as VII. ART 2 of P V and VI
truncate below. P VI as long asP VII. T cleft. Live Superfamily Archaeoscinoidea
animals more or less enti rely intense blue, colour Body of male slender (as in Scina), body of female
deepest along venter. For discussion of validity of inflated to spherical. Cuticle transparent. MD with
S. gracilis see (Barnard, 1972). long thin 3-articulate palp, ART 3 longer than ART 1
• Synopia scheelana Bovall ius, 1886 (Fig. 4.32). and 2 combined. Incisor very narrow. MXP with large
Length 4-6 mm. Body slender. Head triangular. outer lobes. All P simple. T triangu lar. One family.
Eyes elongate. ANT I with secondary flagellum
shorter than ART I of main flagellum (the latter Family Archaeoscinidae
composed of several fused ART). Segment I of Small animals (2-6 mm). With characters of superfami-
pereon as long as segment II, VI shotter than Vll, ly. Two genera. Single genus in the South Atlantic.
which is longest. ART 2 of P V ovate, of P VI
almost circular. T cleft (see key). Live animals Genus Archaeoscina Stebbing, 1904
blue or hyaline. Body smooth. Head sho1t and high. Eyes small.
• Synopia ultramarina Dana, 1853 (Fig. 4.33). Peduncle of ANT I 3-articulated in females, 2-articu-
Length 5-6 mm. Body strongly arched. Head trian- lated in males. ln females this ANT shorter than two
gular. Eyes more or less round. ANT I with acces- first pereonites, in males equal to first three pereo-
sory flagellum as long as or longer than ART I of nites. ANT 11 wi th conical ART of flagellum in
mai n flagellum (the latter composed of several female, with rod-like ART of flagellum in males.
fused ART). Segment I of pereon is longer than F lagellum of ANT II not whip- li ke in males. P V-VII
segment II, VI is longer than VII, which is longest with long, strong, non-retractile dactyls. Two species
of all. ART 2 of P V and VI rounded, nearly circu- (one of them doubtful), only one in the South Atlantic
lar. P VI longer than P Vll. T cleft (see key). plankton.
Ordinary li ve animals and eggs intense blue, but • Archaeoscina steenstrupi (Bovallius, 1885) (Fig.
may be hyal ine (see Bovallius, 1886). 4.34). Length 3.5-4 mm (females), 2.5-3.5 mm
fo r discussion of two more Synopia species found (males). ART 3 of mandibular palp curved, 1.5
-'nly in the northern part of the tropical Atlantic times as longer as ART I and 2 combined. ART 5
~ean, and not treated here, see (Andres, 1984). ofP I and IT broad but not expanded distally, ART
6 more oval than conical. Rami ofUR narrow, lan-
Suborder Hyper iidea ceolate, with serrate margins. T reaching middle
8assification of the Hyperiidea is treated variously by ofbasipodite ofUR III .
.:rfferent authors (see Bowman and Gruner, 1973;
1170 Amphipoda
Supetfamily Lanceoloidea 4a Rostrum practically absent: Lanceola pacifica

Pereons equal, weakly inflated both in males and 5 Sm all distal ART of ANT I fused (Fig. 4.39):
females. MD palp 3-articu late, ART 3 shorter than Lanceola sayana
ART l and 2 combined; incisor may be wide or nar- 5a Small distal ART of ANT I separate (Fig. 4.36):
row. Three families, l in the South Atlantic. Lanceo/a felina
Family Lanceolidae • Lanceola c/ausi Bova llius, 1885 (Fig. 4.35).

Medium to large animals (6-70 mm). Body inflated, Length I 0- 16-(20) mm. Body smooth. Rostrum
usually semi-transparent. Eyes small or absent, or absent. Distal (small) ART of ANT I well-deve-
transformed into reflector organ (in Scypholanceola). loped, free, cylindrical, apical longer than others.
Coxae separate from pereonites. ANT I first flage llar P moderately short; P V-VII almost subequal, P V
ART large, conical; apical ART small. MD w ith laci- shorter than pereon, P VI slightly longer than V.
nia mobilis usually poorly developed. MXP with Hooded dactyls of P V-VU strong. T triangular,
inner lobes fused proximally, separate distally. P V- 0.5-0.7 times as long as basipodite ofUR Ill. Live
Vll with spoon-shaped ART 6 , hooded retractile dac- animals pink or colourless, sometimes orange. L.
tyl. Includes 6 genera, 3 in the South Atlantic. clausi can be recognised easily by their general
appearance: small size and moderately short ANT
Key to the genera of Lanceolidae and P. Very variable species, especiall y in body
I ART 6 of P l conical ................................... ......2 shape, and in some details ofP morphology.
Ia ART 6 of P I w idening distally: Prolanceola* • Lanceola felina Bovallius, 1885 (Fig. 4.36).
[rare animals, still not found in Atlantic Ocean, Length to 33 mm. Resembles L. sayana. Rostrum
but widely distributed) short, slightly curved, obtuse. Body smooth, w ith-
2 Left MD with lacinia mobilis minute. [Palp of out carina. Distal (small) ART of flagellum of
MX T not shorter than outer plate] .................... 3 ANT I flat but separate, apical longer than other
2a Left MD with lacinia mobilis nearly as w ide as two. P VI relatively shorter than in L. sayana, sub-
incisor: Megalanceola equal to pereon. T subequal to, or slightly shorter
3 Lateral surface of head with may be small, but than, basipodite of UR III, with rounded top.
eyes common, without concavities: Lanceola • Lanceola loveni Bovallius, 1885 (Fig. 4.37).
3a Lateral surface of head with 2 concavities separ- Length 15-29 mm. Integument weak. Rostrum
ated by an oblique ridge. Eyes transformed into poorly developed. Eyes small. Body with weak
reflection organs : Scypho/anceola dorsal carina. Carina forming small mid-dorsal
teeth in pereonites III-IV, pleonites I-III and
Gen us Lanceola Say, I 818 urosomite l. D istal (small) ART of ANT I free,
Inter-ANT lobe rounded, rud imentary to well-deve- cylindrical. P VI-VII thin, VI slightly longer than
loped. Eyes small, round to oval. MX I with broad V. Hooded dactyls (P V-Vll) thin. T triangular.
inner lobe. MD with very smalllacinia mobilis, broad sometimes w ith produced top; top usual ly sharp. T
incisor. PI with ART 5 expanded distally. Includes 12 0.3-0.5 times as long as basipodite ofUR III.
species, 6 in the South Atlantic. • Lanceola pacifica Stebbing, 1888 (Fig. 4.38).
Length 22-38 mm (females), 26-34 mm (males).
Key to the species of Lanceola Rostrum absent. Eyes large, kidney-like. Body
I T subequa1 to basipodites of UR III .................. 2 without carina. 3 distal (small) ART of A T I
la T 1.5-2 times as short as basipodites of UR III free, cylindri ca l; apica l setae short in ANT 1. P VI
..... .............. .................................... ................. .. .3 much longer than V. Hooded dactyls of P V-VII
2 P VI much longer than P IV: Lanceola loveni thin, curved. T pointed, wi th rounded top, slightly
2a P VI subequal to P IV: Lanceola c/ausi longer than basipodite of UR III. In deep-water
3 Dorsal part of pleon and pereonites VI-VII Lanceola pacifica, two morphological forms are
smooth ....... .... ............... ..... ........................... ... .4 known (typica and robusta), probably associated
3a Pleonites and pereonites VI-VII produced poste- symbiotically with "weak" medusae and " muscu-
riorly onto mid-dorsal spines: Lanceo/a sen·ata lar" siphonopbores (Woltereck, 1909).
4 Rostrum well-developed .................................... 5
-
Amphipoda 1171
• Lanceola sayana Bovallius, 1885 (Fig. 4.39). cesses. Coxae II-V sharp anteriorly. Dactyls of P
Length 19-42 mm. Rostrum sharp, triangular, V-VII strongly curved in mature animals, barely
sometimes long and straight, but often short and curved in young. T with sharp or rounded top.
curved. Eyes we ll pigmented. 3 distal (small) ART
of ANT T fused. Integument more or Jess dense. Superfamily Scinoidea
Weak carina on dorsal side ofpereon and pleon . P Males always with non- inflated body. ANT I in males
VI strong, longer than P V and longer than pereon. (and often in females) long, not shorter than five first
P VU twice as short as VI. Hooded dactyls of P V- pereonites combined. MD with rudimentary !-articu-
VII especially strongly curved. T with rounded late palp, or completely lacking palp. Three families
top, subequal to or longer than basipodite of UR in total, 2 in the South Atlantic.
III. Colour of live animals from dark-red to
orange, or colourless, sometimes anterior patt of Family Mimonectidae
body pigmented and posterior colourless, some- Animals medium-large (25 -30 mm). Pereon expand-
~ -- - times pereon colourless but pleon and mouth parts
orange or carmine-red.
ed, in mature females balloon-like with pereonites I-V
strongly inflated, pleon slender. Eyes small or absent.
• Lanceola serrata Bovallius, 1885 (Fig. 4.40). ANT I with long narrow conical proximal ART of fla-
Length to 34 mm. Integument weak. Pereonites V- gellum, 3 small distal ART. Female ANT I longer than
VII, pleonites (especially), and urosomitc I with pereonite I, in males subequal to first five pereonites
strong dorsal teeth. Rostrum short, broad, flat, combined. MXP with oval outer lobes, separate inner
poorly developed, sometimes absent. Eyes small, lobe. Coxae free. P simple. P III-VII subequal. T short.
oval. Distal (small) ART of ANT I free. P VI Two genera. Single genus in the South Atlantic.
much longer than V. P VII twice as shott as VI. T
narrow, triangular, with sharp top, reaching (or Genus Mimonectes Bovallius, I 885
almost) distal end of basipodite of UR Til. MD completely lacking palp, with well-developed
lacinia mobilis. Five species, 3 in the South Atlantic.
~nus Megalanceola Pirlot, I 935
:..arge animals. Head short. Rostrum very small. Inter- Key to the species of Mimonectes
-....::: "\...'\T lobe fairly narrow. Eyes oval, without pigment. I ART 6 of P 1 and II with sharp procession on dis-
ID with narrow incisor and lacinia mobilis nearly as ta l end (over base of dactyls): Mimonectes gaus-
~ad as incisor. PI with ART 5 only slightly expand- si
!rl distally. Two species in genus. Single species in Ia ART 6 ofP I and II without sharp process on dis-
~ South Atlantic. tal end ................ ................................................ 2
• .l1egalanceola remipes (Barnard, 1932) (Fig. 2 ART 6 of P II with concave palmar margin
-l.4 1). Length to 73 mm (females), to 48 mm (especially in males), dactyl strong, curved:
(males). Head with small straight rostrum . ART 4- 1l1imonectes sphaericus
6 ofP III-V very broad, oval or conical. Dactyls of 2a Palmar margin of ART 6 of P II straight or con-
P III-IV very small, retractile. ART 2 of mandibu- vex, dactyl short, almost straight: Mimonectes
lar palp shorter than ART 3. loveni
~us Scypholanceola Woltereck, 1905 • Mimonectes gaussi (Woltereck, 1904) (Fig. 4.43).
~reral surface of head with 2 concavities separated Length 11-18 mm in females, males to 9 mm. PI
-- ridge nmning obliquely posteroventral of inter- and II with conical, distally nanowed ART 6, on P
"'-"-"T lobe. Eyes without refracting elements or pig- I, ART bearing numerous lo ng setae. ART 6 of P
;;;;:cnt. in form of band with ends lying in concavities, 1 subequal, or no more than in 1.3 times as short
:-.:nnected across ridge. Two species in genus. Single as, ART 5. Dactyls of P long, thin, barely curved.
-;recies in the South Atlantic. • Mimonectes loveni Bovallius, 1885 (Fig. 4.44).
• Scyplzolanceola aestiva (Stebbing, 1888) (Fig. Length 24-28 mm, males to 13 mm. Mandibular
-l.42). Length to 61 mm, mainly 25-45 mm. Head incisor more or less wide, lacinia mobilis 0.5
with small but well-developed, sharp rostrum. times as broad as incisor. MXP without any exci-
Distal (small) ART of A T I without apical pro- sions on tops of outer plates. ART 2 of PI and II
1172 Amphipoda
broad, but almost linear. ART 6 ofP II conical, in Coxae V sharp both anteriorly and posteri orly. UR
females broader than in males. P VI-VII with short straight, very long, thin.
curved dactyls, ART 3 twice or more as short as
ART4. Genus Ctenoscina Wagler, 1926
• Mimonectes sphaericus Bovallius, 1885 (Fig. Pereon and pleon segments with strong mid-dorsal
4.45). Length to 18 mm in females, males to I 0 spines. Pereonites I and II separate. ANT II of males
mm. Mandibular incisor narrow. MXP with small spine-like. MXP without inner lobe; outer lobes well-
excision on top of outer plates. Excision on distal developed. PI chelate, chela formed by ART 5 and 6.
end of ART 6 of P II stronger in males than in P V with all ART free. P III-VII with slightly spoon-
females, sometimes weak, sometimes very strong shaped ART 6. Three species. Single species in South
(article broadened distally in forma va/diviae). P Atlantic.
VI-VII as in M. loveni. Ctenoscina brevicaudata Wagler, 1926 (Fig. 4.4 7).
Length to 4 mm. Pereonites I and II without mid-
Family Scinidae dorsal spines. Anterodistal comer of ART 2 of P :
Small or medium-large animals (2.5-30 mm exclud- with two thin, slightly curved processes (one may be
ing ANT). Body slender, flattened dorsoventrally. weak). Anterior margin of this ART slightly serrate.
Head shott and wide. Eyes small or absent. ANT l Coxae 11-V sharp both anteriorly and posteriorly.
horn-like, as long as pereon or as pereon and pleon
combined, with 1-atticulate peduncle, very long ART Genus Scina Prestandrea, 1833
2 (first ART of flagellum), without ART 3 (reduced Body without mid-dorsal spines, sometimes with 4
ART 3 may be present in juveniles). MD without weak dorsal carina. ANT II of male well-developed. I
palp, lacinia mobilis, or molar. Inner lobes of MXP Pereonites I and II separate. MXP with inner lobes
completely fused or absent. All P long and thin. P V completely fused into a single lobe, outer lobes well
much longer than others, ART 2 very long, often
broadened, with marginal teeth. All UR with endopo-
developed, sharp or oval. P I simple. All P without
spoon-shaped ART 6. All UR with endopodites fused
•J
dites fused to basipodites, exopodites free, short, spi-
niform on URI and II, usually III as well. T small, tri-
to basipodites, exopodites free, short, spiniform.
Thirty two species, 22 in the South Atlantic.
•~
angular. Four genera, 3 in South Atlantic.
Key to the species of Scina
Key to genera of Scinidae I ART 4 of P V subequal to, or slightly shorter
I Body segments without mid-dorsal spines: Scina than, or no more than 25% longer than, ART 5.
")
la Body segments with mid-dorsal spines .......... ... 2 ·································· ······ ···································-
2 Pereon segments I and II fused. ART 4-6 ofP V Ia ART 4 ofP V much longer than ART 5 ......... 1-t
fused. Outer plates of MXP reduced to small 2 ART 6 of P V thin, weak, short, more than twice
narrow rectangular appendages: Acantlwscina as short as ART 5. Medium-size animals with
2a Pereon segments I and II not fused. ART 4, 5 ANT I longer than pereon ................................. 3
and 6 of P V separate. Outer plates of MXP 2a ART 6 ofP V less than twice as short as, or sub-
broad: Ctenoscina equal, or slightly longer than ART 5. Usuall~
Genus Acanthoscilza Vosseler, 1900

small species with different proportions of A 'T
1 and pereon ...................................................... .5
•
Pereon and pleon segments with strong mid-dorsal 3 ART 2 of P V with whole anterior and posterior
spines. Pereonites I and II fused. ANT II of males margins weakly serrate .................................... ..~
spine-like. MXP without inner lobe, outer lobes small, 3a ART 2 of P V with anterior and posterior mar-
finger- like. PI simple. P V with fused ART 3-5. Two gi ns smooth, or antetior margin with few denti-
species. Single species in South Atlantic. cles near distal process: Scina stebbingi
• Acanthoscina acanthodes (Stebbing, 1895) (Fig. 4 Dactyl ofP VI very shott, curved, with ART 4 1.5
4.46). Length 4-6 mm. Lobes of maxillae II fused
proximall y. Dactyls of P I and II in 5-6 times as 4a
times as long as ART 5 or 6: Scina curvitiactyla
Dactyl of P VI moderately long, a lmost straight
•s
short as ART 6. ART 2 ofP V in 1.5 times as long with ART 4 subequal to ART 6, slightly longer - (]
as ART 2 ofP IV. ART 5 and 6 ofP VI subequal. than ART 5: Scina crassicornis
Amphipoda 1173
' ART 6 of P I (sometimes I and II) with pointed much shatter, more robust than V: Scina oedi-
process on anterodistal corner, over dactyl .... ... 6 carpus
.::a ART 6 ofP I and II without anterodistal process 16 ART 6 ofP VI shorter than, or subequal to, ART 5
...................................... ..................................... 8 ... ..... .......... ..... ............ .... ............. ... .................. 17
6 Outer lobes ofMXP oval, less than twice as long 16a ART 6 of P VI essentially longer than ART 5
as inner lobe. ART 6 shorter than, or subequal to ..................... ...... ......... ...... ........... ..... ............ .. . 19
ART 5 in P VI-VII ........ .... ... ........ .... ............. ... .7 17 P and UR not very long and thin . ART 4 ofP VI
6a Outer lobes of MXP pointed, more than 5 times shorter than, or subequal to, ART 5 ................ 18
as long as inner lobe. ART 6 longer than ART 5 17a P and UR very long and thin. ART 4 of P VI 1.5
in P VI-VII: Scina incerta time as long as ART 5: Scina stenopus
Anterodistal processes of ART 6large in PI and 18 Anterior margin of ART 2 of P V dentate. ART
II. P III and IV long, delicate: Scina marginata 6 of P VII much longer than ART 5: Scina lan-
-a Anterodistal process of ART 6 small in P I ghansi
(especially in males), absent in P II. P III and IV 18a Anterior margin of ART 2 ofP V smooth, some-
robust: Scina submarginata times with few denticles near distal process. ART
P VII with broad ART, ART 6 of this P only 6 ofP VII shorter than ART 5: Scina tullbergi
twice (or less) as broad as wide, dactyl small, 19 Outer lobes of MXP 3 times or more as long as
nail-like .. ...... ......... .... .... ..... .. ........ .... ............. .....9 broad, lanceolate or pointed ..... .... ... ........ ........ 20
Sa ART 6 ofP VII more than twice as long as wide, 19a Outer lobes ofMXP twice as long as broad, oval,
dactyl ordinary, not nail-like ........................... 10 with excavated top: Scina latifrons
P V-VII with numerous long setae on surface of 20 ART 5 of P V subequal to, or only slightly lon-
distal parts of ART: Scina pubera ger than, ART 6 ... ... ... .......... ... ... ......... ..... ........ 21
"-3 P V-VII without numerous long setae on surface 20a ART 5 ofP V twice as long as ART 6: Scina rat-
of distal parts of ART: Scina spinosa trayi
IC ART 2 ofP VI shorter than whole P VII. Dactyl 21 Dactyls ofP V-VII long, slightly curved: Scina
of P V moderately short .................................. 11 simi/is
-Oa ART 2 of P VI longer than whole P VII. Dactyl 21 a Dactyls of P V-VII very short: Scina nana
ofP V long and thin: Scina lepisma
l' Inner margin of UR 1 dentate .......................... 12 • Scina borealis (G. 0. Sars, 1882) (Fig. 4.48).
B a Inner margin ofUR I smooth .......................... 13 Length 7-8 mm. Body smooth. ANT I slightly
~.~ MXP with long (more than 3 times as long as longer than pereon. Outer lobes of MXP with
wide) pointed outer lobes. Dactyl of P V sharp top. Inner lobes about half as long as outer.
unarmed: Scina borealis P V with strong denticles on both margins of ART
.:a MXP with short (less than twice as long as 2. Distal process of ART 2 well-developed. ART
broad) oval outer lobes. Dactyl of P V with row 5 of P V much longer than ART 6. P VI slightly
of setae on anterior margin: Scina vosseleri sh01ter than V, with ART 5 shorter than ART 6. P
- Inner margin of UR li with step-like excision on VII weak. UR I with strongly dentate inner mar-
level of exopodite: Scina excisa gins. Live animals red.
. Inner margin of UR II without step-like exci-
~
Scina crassicornis (Fabricius, 1775) (Fig. 4.49) .
sion: Scina pusilla Length 12-29 mm. Pereon elongate, oval; pleon
- - ~
'-'- Inner margin ofUR I with a single large spine on and urosomite I with weak dorsal and two lateral
level of exopodite, a few smaller spines on basi- carinas. ANT I longer than pereon. Outer lobes of
-- podite ... .......... .... .... .......................................... 15 MXP elongate, pointed. Inner lobes 3 times as shott
~ Inner margin of UR I without single large spine as outer. ART 6 ofP V in 3-4 times as shott as ART
on level of exopodite .. .............. ............. .......... 16 5, dactyl of P VI long, slightly curved. P VI much
.;:
Outer lobes of MXP 8 times as long as wide, shorter than V. P VII longer than half of P VI .
inner lobe very short. P V and VI subequal: • Scina curvidactyla Chevreux, 1914 (Fig. 4.50).
~
Scina wolterecki Length 13-23 mm. Body with well-developed dor-
.;:
-- Outer lobes of MXP 3 times as long as wide, sal and lateral carinas. ANT I longer than pereon.
inner lobe about 3 times as short as outer. P VI Outer lobes of MXP oval with pointed top. Inner
1174 Amphipoda
lobes 2-2.5 times as short as outer. ART 6 of P V Some specimens may have delicate triangular pro-
thin, as long as quarter of ART 5, dactyl very cess on anterodistal corner of ART 3 ofP V. Ratio
small. P VI (as opposes to S. crassicornis) only of length of ART 4, 5 and 6 ofP V variable and can
slightly shorter than P V; dactyl very small, curved. change at least from 10:15:14 to 10:22:15. Dactyl
ART of P in adults broader than in juveniles. Live ofP V extremely long, slightly curved. P VI almoSt
animals transparent, with pink rings on P. subequal toP V; dactyl short, curved. P VIJ more
• Scina excisa Wagler, 1926 (Fig. 4.51). Length 4- than twice as short as VI. Live animals red.
7 mm. Body smooth. ANT I strong, not longer • Scina marginata (Bovallius, 1885) (Fig. 4.56).
than pereon. Outer lobes ofMXP elongate, broad- Length 4.5-6.5 mm. Body robust, smooth. ANT I
ly oval. Inner lobe 4-5 times as short as outer. as short as pereon. Outer lobes of MXP short
Anterior margin of ART 2 ofP V strongly dentate, broad, oval. Inner lobes 1.5-2 times shorter than
posterior margin finely serrate; dactyl long, outer. P short and strong. ART 6 of P I with ser-
straight. P VI slightly shorter than V; anterior mar- rate posterior margin, with well-developed, sharp
gin of ART 2 weakly serrate. P VII moderately anterodistal process nearly subequal in length to
long. Inner margin ofUR II with step-like excision dactyl. ART 6 of P II with serrate anterior margin
on level of exopodite. and distal part of posterior margin; with shorr
• Scina incerta Chevreux, 1900 (Fig. 4.52). Length anterodistal process. P VI and VII strong, ART
9-16 mm. Pereon elongate, oval, with well-deve- shorter and broader than in P V. Dactyls of P V-
loped dorsal carina. ANT I thin, equal to, or long- VII short, strongly curved.
er than, pleon and pereon combined. Outer lobes • Scina nana Wagler, 1926 (Fig. 4.57). Length 2~
of MXP elongate, pointed, broadened in proximal mm. Very like S. tullbergi. Body smooth. ANT I
part. Inner lobes 6-8 times as short as outer. ART thick, not longer than pereon. MXP as in S. tull-
6 of P I with small triangular sharp anterodistal bergi. In P III-IV ART 2, 4, 5 and 6 in ratio
process. P V longer than other, ART 6 slightly 10:4:7:5. P V long, strong, posterior margin of
shorter than ART 5. ART 6 of P VI longer than ART 2 serrate, anterior margin smooth with 1-2
ART 5. P VII much shorter and weaker than P VI. denticles in distal part; ART 4 almost twice as
Live animals carmine-red, with tops of A T I long as ART 5; ART 6 shorter than 5; dactyl very
transparent. short, curved. P V I shorter than V; ART 4 longer
• Scina langhansi Wagler, 1926 (Fig. 4.53). Length than 5 ( 13:9), subequal to ART 6; dactyl very
8- 10 mm. Pereon broad, oval, smooth. PIeon with short, curved. ART 6 of P VI narrower than ART
weak carina. ANT I subequal to pereon. Outer 5. P VII short, weak, dactyl very short.
lobes of MXP not large; strongly pointed. Inner • Scina oedicatpus Stebbing, 1895 (Fig. 4.58).
lobes 5 times as short as outer. ART 4 ofP V long- Length 7 -8 mm. Body with weak carinas.
er than ART 5 and 6 combined. ART 4-6 of P VI Pereonites I-V broad, VI and VII narrow. ANT I - 3
subequal. P VII short. longer than pereon. Outer lobes of MXP lanceo-
• Scina latifrons Wagler, 1926 (Fig. 4.54). Length late. Inner 4 times as short as outer. P long and
to 6 mm. Body smooth. ANT I subequal to pereon. thin. P V much longer than other; both margins o:
Outer lobes of MXP short, oval, sometimes with ART 2 dentate; ART 4 longer than ART 5, ART 6
excavated top. Inner Jobes 4 times as short as 2-3 times as short as ART 5, dactyl thin. P \ 1
outer. P long and thin. Posterodistal corner of much shorter, stronger than 5. P VII short and thin.
ART 6 of P I with small rounded process, with UR long and thin. Live animals red.
strong seta on top. ART 2 of P V with smooth • Scina pubera Wagler, J926 (Fig. 4.59). Length to
anterior margin, finely setTate posterior margin; 9 mm. Body robust, smooth. ANT I strong, short-
ART 4 twice as long as 5 or 6. ART 6 of P VI er than pereon. MXP with elongate oval outer
longer than 4 or 5. P VII short and weak. lobes. Inner lobes less than half as long as outer. P
• Scina lepisma (Chun, J 889) (Fig. 4.55). Length 7- short and strong. Surface of P V and VI covered
10 mm. Body smooth. ANT I slightly longer than with numerous thin setae (unique character! ). P
pereon. Outer lobes of MXP short, oval, with VII as in S. spinosa.
uneven distal margin. Inner lobes twice as short as • Scina pusilla Chevreux, 1919 (Fig. 4.60). Length
outer. ART 2 of P V with both margins serrate. 3-4 mm. General appearance as in S. latifrons.
Amphipoda 1175
MXP as inS. latifrons, but inner lobes moderately ly smooth (unique character!). P V slightly longer
shorter. ART 2 of P V broad, with both margins than VI, last slightly longer tan VII. Dactyls of P
strongly serrate; ART 6 longer than 5 or 4 . P VI V-VII short, strongly curved.
slightly shorter than V; ART 4 longer than 6, 6 • Scina stenopus Stebbing, 1895 (Fig. 4.65). Length
longer than 5; dactyl short, curved . T extr·emely 5-9 mm. Body smooth. Head with dorsal carinae,
long, reaching middle of basipodite of UR TIL large lateral spines below attachment point of
• Scina rattrayi Stebbing, 1895 (Fig. 4.61). Length ANT I. ANT I as long as pereon and pleon com-
2.5-6 mm. Pereon broadly ov al, body smooth, bined. Outer lobes of MXP elongate, broad lance-
compressed dorsoventrally. ANT I subequal to olate or oval. P and UR extremely long and thin.
pereon. Outer lobes of MXP elongate, subconical, In P V ART 2 with both margins entirely dentate;
w ith rounded top. Inner lobes reduced, more than ART 4, 5 and 6 in ratio 10:7:4.5; dactyl short. P VI
=~- 10 times as short as outer. P long and thin. ART 2 slightly shorter than V, ART 2 smooth, ART 4, 5
of P V with dentate posterior margin and distal part and 6 in ratio I 0:6:6.
of anterior margin; ART 4 thin, slightly shot1er • Scina submarginata Tattersall, 1906 (Fig. 4.66).
than ART 2 or subequal to it; ART 6 twice or more Length 5-8 mm. Body robust, smooth. ANT I
times as short as ART 5; dactyl very short. P VI strong, shorter than pereon. Outer lobes of MXP
slightly shorter than V; ART 6longer than 5. P VII short, broad, oval. Inner lobes 1.3-1.25 times as
very short and weak. UR I and III with spines on short as outer. P short and strong. Anterodistal
inner margins, UR IT with spines on outer margin. process of ART 6 triangular, small in PI (especial-
• Scina simi/is Stebbing, 1895 (Fig. 4.62). Length ly in males), absent in P II. Anterior margin of
2.5-3.5 mm. Very much like S. tullbergi. Body ART 6 smooth in P I, weakly serrate in P II. P III-
smooth. ANT I thick, not longer than pereon. VII as in S. marginata.
Outer lobes of MXP elongate, lanceolate. Inner • Scina tullbergi (Bovallius, 1885) (Fig. 4.67).
lobe more than 10 times as shot1 as outer. In P III- Length 2-4.5 mm. Body smooth . ANT I thick, not
IV ART 2 , 4, 5 and 6 in ratio 12:6(7):8:7. P V longer than p ereon. Outer lobes of MXP elongate,
long, strong, posterior margin of ART 2 serrate, lanceolate. Inner lobe 8- 12 times as short as outer.
anterior m argin smooth, with 1-2 denticles in dis- P V long, strong, posterior margin of ART 2 ser-
tal part; ART 4 almost twice as long as ART 5; rate, anterior margin smooth with 1-2 denticles in
ART 6 subequal to 5; dactyl moderately long, distal part; ART 4 twice as long as ART 5; ART 6
slightly curved. P VI shmter than V; ART 4 and 6 subequal to 5 or 1.5-2 times as short as it; dactyl
subequal, ART 5 shorter than 4 or 6, dactyl long, short. P VI shorter than V. ART 6 of P VI sub-
slightly curved. ART 6 of P VI narrower than equal to or slig htly shorter or longer than ART 5;
ART 5. P VTT short and weak, dactyl long. as broad as, or narrower than, ART 5. P VII short
Scina spinosa Vosseler, 1901 (Fig. 4.63). Length and weak. UR weak ly armed. Live animals trans-
6- 11 mm. Body robust, smooth. ANT I clearly parent-red.
shorter than pereon. Outer lobes of MXP short, • Scina vosseleri Tattersall, 1906 (F ig. 4 .68).
broad, more or less narrowing distally. Inne r lobes Length 9- 15 mm. Body smooth. Head with weak
~-4 times as shm1 as outer. P shot1 and strong. carina. ANT I thin, subequal to w ho le body . MXP
Posterodistal parts of ART 3 and 4 ofP III and IV with short, oval outer lobes with obtuse top. Inner
more or less concave (especially in adu lts). ART 2 lobes 3 times as short as outer. ART 2 of P V with
of P V w ith both margins serrate; dactyl short, strong anterodistal process, almost reaching distal
slightly curved. P VI slightly shot1er than V. P VII e nd of ART 4 , thi s process a quarter as long as
slightly shorter but more robust than P VI, w ith main body of ART 2; both margins of process
strong broad ART; dactyl short, nail-like, curved. dentate. Dactyl ofP V with many setae on anterior
Species characters widely variable. margin. P VI shorter than V, VII slightly shorter
• 5cina stebbingi C hev reux, 19 19 (Fig. 4 .64). than VI.
:...ength 8-13 mm. Body smooth. ANT I subequa l • Scina wolterecki Wagler, 1926 (Fig. 4.69). Length
;:o or shorter than pereon. Outer lobes of MXP 7-10 nun. Body smooth. ANT I on ly slig htly
dongate, oval. Inner lobes 2-3 times as short as shm1er than pereon and pleon combined. MXP
outer. ART 2 of P V with both margins complete- w ith unusually narrow and long outer lobes,
1176 Amphipoda
strongly reduced inner lobes. P V long, but mod- basipodites and endopodites. Deep-water animals
erately shotter than in S. oedicarpus. P VI sub- Only one genus in family. Cystisomatidae show spe-
equal to V. UR I with one (rarely a few) strong cific development of physosoma stage in young (,,;th
denticle(s) on inner margin opposite exopodite. long, but weak, parchment-like spines on body. see
Fig. 4.70); length of physosoma: to 15 mm.
Infrao rder Physocephalata
Head large, longer than pereonite l. Integument weak Genus Cystisoma Guerin-Meneville, 1842
or strong. Eyes usually gigantic, occupying entire lat- With characters of family. About 5 species, 4 in the
eral surface of head, except rostrum (if any). Only in South Atlantic. Species level characters very variable.
Vibilia and Vibilioides (Vibiliidae), and in Bougisia only a few very specific.
(Hyperiidae, not recorded in the South Atlantic) eyes
small to moderately large. MX I without inner lobe, Key to the species of Cystisoma
MXP always with inner lobes completely fused. Epi- I MD with only one central denticle ................. ..._
and mesopelagic animals. la MD with one central, l -2 lateral denticles ..... ...-
2 Only "top" and "glandular" denticles present in
Superfamily Vibilioidea ventral denticles of head (see family descrip-
ANT I insetted on anterior surface of head, flagellum tion): Cystisoma fabricii
with large straight ART 1, one or a few rudimentary 2a Beside "top" and "glandular" denticles, a fe\l
distal ART, inserted terminally. ANT II straight, other pairs of ventral denticles of head .. ........ .. ..::
inse1ted on anterior or ventral surface of head, sh01t or 3 Distal parts of ART 6 of P III-VII glandular in
rudimentary. Only PI and II may be chelate or subche- adult females. Urosoma and UR combined long-
late. P VII may be partly reduced. T free. 3 families. er than pleon: Cystisoma pellucida
3a Distal patts of ART 6 ofP III-VII not glandular
Family Cystisomatidae in adu lt females. Urosoma combined with l."R
Large animals (to 140 mm). Cuticle thin, parchment- not longer than pleon: Cystisoma magna
like, completely transparent; musculature very weak. 4 P long and thin; ART 2 of P Vll almost linear
Head very large, rounded dorsally, ventral surface flat Cystisoma longipes
with dentate marginal carinas. Two other rows of 4a P with broadened ART; ART 2 of P Vll pear-
"ventral" denticles forward of mouth parts. "Top" like : Cystisom a latipes* [Found in Nonh
(anterior) denticle in these rows largest, present in all Atlantic Ocean]
species; the second ("glandular") denticle is also
present in all species. This denticle is a mdimentary • Cystisoma .fabricii Stebbing, 1888 (Fig. 4. 7 1
ANT ll, as indicated by the presence of a gland open- Length to 92 (females) - 50 (males) mrn. Heac
ing at its base (Stebbing, 1888). Other ventral denticles with 12-1 6 marginal denticles on each side, only:::
may number 0-6, sometimes different in right and left pairs of ventral denticles ("top" and "glandular"· .
rows. Eyes very large, oval, practically without pig- MD with only one curved denticle. Posterodistal
ment, situated on dorsal surface of head. ANT I with process of ART 5 ofP II finel y serrate.
reduced peduncle ART, 2-articulate flagellum, with • Cystisoma longipes (Bovallius, 1886) (Fig. 4.72 _
first ART long and slender, second strongly reduced. Length to 80 (females) mm. Head with 15-1 8 mar-
ANT II attached to ventral surface of head, strongly ginal denticles on each side, and 5-6 pairs of ventra:
reduced, spine-like (see above). MD without palp, denticles. MD with 2-3 denticles (central + l -2lare-
with one (central) or few (central and lateral) denticles ral). Posterodistal process of ART 5 of P II strong-
on its body. Pereon ites and pleonites with short mid- ly setTate. P long and thin; P V subequal in lengrt
dorsal spines/carinas. First two to fo ur pereonites may to entire body; ART 2 of P Vll almost linear
be fused. Coxae fused with pereonites. P with serrate • Cystisoma magna (Woltereck, 1903) (Fig. 4.73
margins, P I and II chelate. In adult females distal part Length to 140 (females) - 70 (males) mm. Head
of P VII developed into "eggs pincers": distal end of with 10- 14 marginal denticles on each side, 5-6
elongate ART 6 expanded with rounded excavation pairs of ventra l denticles. Pereon with especiall_
(egg size), dactyl strongly curved, able to fix egg in strong dorsa l carina. MD with only one centra:
this excavation. UR II absent, other UR with fused denticle. Posterodistal process of ART 5 of P r:
Amphipoda 1177
strongly serrate. Distal patts of ART 6 ofP III-VII Family Vibiliidae

not glandular in adult females. Urosoma combined Medium-large animals (5-23 mm). Body compact,
with UR not longer than pleon. moderately slender; head small or moderately large,
Cystisoma pellucida (Willemoes-Suhm, 1874) subquadrangular. Eyes small, oval, separate; rarely
(Fig. 4.74). Length to 85 mm. Head with 13- 15 represented by a few rows of isolated ommatidies. All
marginal denticles on each side, 4-7 pairs of ven- pereonites separate. Coxae free. ANT I with short
tral denticles. MD with only one central denticle. peduncle, large, straight, oval, lance-shaped, or coni-
Posterodistal process of ART 5 ofP II strong ly ser- cal fi rst ART of flagellum, remaining ART rudimen-
rate. Spines on CX I and II especially long, sharp. tary, hardly visible. ANT II thin, subequal to ANT I.
Distal parts of ART 6 of P III-VII glandular in MXP with short rounded inner lobe. Only P TT chelate
adult females. Urosoma combined with UR longer (with exception Cyllopus lucasi). P VII may be with
than pleon. Poorly described C. neptunus Guerin- reduced distal ART. Rami of all UR free. 3 genera.
Meneville, 1842, mentioned in many older publi- All Vibiliidae are associated with gelatinous plankton,
cations, may be senior synonym of this species. mainly with salps.
?:unity Paraphronimidae Key to the genera of Vibiliidae

Small or medium-large animals (6-15 mm). Body 1 Eyes small or moderately large, never occupying
- ender, transparent. Head large, globular, or almost most of head surface. ANT I with spatiliform
-::bical. Eyes large, occupying most of head surface. flagellum ... .... ............. ............... ......................... 2
'W pereonites free. All coxae fused with pereonites in 1a Eyes occupying most of head surface. ANT I
::;ales, coxae 11-V free in fema les. MD without palp with conical flagellum: Cyllopus
.:.d molar. MXP with fused outer and inner lobes. P I 2 MX I with rounded inner plate. MX II well-
=d II shorter than other, P I weakly subchelate. ART developed. ART 3-7 of P VII combined longer
:of all P narrow, linear. UR with free rami. Only one than ART 2 (except in Vibilia chuni): Vibilia
~us in family. 2a MX I lacking inner plate. MX II strong ly
reduced or absent. ART 3-7 of P VII combined
Genus Paraphronima Claus, 1879 much shorter than ART 2: Vibilioides
illl characters of fami ly. Two species.
• Paraphronima crassipes Claus, 1879 (Fig. 4.75). Genus Cyllopus Dana, 1852
Length 20-25 mm, rarely to 30 mm (females). Head large. Eyes occupying most of head surface.
~fuch like P. gracilis but body moderately more ANT I with slender conical flagellum. ANT II insert-
robust. Pereonites I-IV subequal. P stronger, with ed on ventra l surface of head. P VII with well-devel-
slightly broadened ART 2. ART 5 of P I less oped broad ART 2, reduced 3 or 5 distal ART Two
expanded distally than in P. gracilis. P VIT only specJes.
slightly shorter than VI, or subequal to it. Cyllopus lucasi Bate, 1862 (Fig. 4.77). Length to
• Paraphronima gracilis Claus, 1879 (Fig. 4.76). 23 mrn. Head longer than pereonites I-III com-
:...ength 10-17 mm (males to 11 mm). Head very bined. P I weakly chelate. ART 2 of P I subequal
:.arge and robust in comparison with slender deli- to distal ART combined; ART 6 expanded distal-
cate body and thin P, longer than pereonites I-IV ly, with serrate triangular posterodistallobe reach-
::ombined, more than twice as high as pereonites. ing one quarter length of ART 6; ART 6 conical-
Pereonites I and II clearly smaller than pereonite oval, shotter than ART 5; dactyl as long as two
dl. ART 4 ofP I with spoon-like process in poste- thirds of ART 6. Posterodistal process of ART 5 of
rodistal pati, P II simple, with narrow linear ART, P II reaching distal end of ART 6; ART 6 shorter
? Ill-VI subequal. P VII shorter than P VT (reach- than 5. P V and VI long, with ART 6 as long as
::Jg d ista l end of ART 5 of P VI). Rami of all UR ART 4 and 5 combined; dactyl thin, 8 times as
ID<>ner than basipodites. Live animals glass-like, short as ART 6. Proximal part of ART 2 of P VII
c-ansparent, with two rows of red face ts in eyes, strongly broadened, ART 2 sharply narrowing dis-
5mall subsurface glitter dots near base of each P; tally, with concave posterior margin. Distal ART
~t5 glow golden-orange in light. combined shorter than ART 2.
1178 Amphipoda
Cyllopus magellanicus Dana, 1852 (Fig. 4.78). 6 Adult animals longer than 15 mm. ART 2 ofP II
Length 10-1 7 mm. Head as long as pereonites 1-III broadly oval: Vibilia robusta
combined. PI simple. ART 2 ofP I slightly short- 6a Adult animals shorter than 15 mm. ART 2 of P
er than distal ART combined; ART 5 and 6 elon- II linear or only slightly expanded .................... -
gate, 5 slightly broader than 6; ART 6 linear; pos- 7 Dactyls ofP V-Vl shotter than quatter of ART 6:
terior margin of ART 6 distally serrate; dactyl Vibilia jeangerardi
twice as short as ART 6. Posterodistal process of 7a Dactyls of P V-VI about third length of ART c
ART 5 of P II not reaching middle of ART 6. P V
and VI much longer than lll and IV, with thin dis- 8 Integument dense. Eyes occupying about third o:-
tal articles; ART 5 longer than 4; 6 much longer lateral surface of head in females. Peduncle of
than 5. P VII reaching only middle of ART 4 of P ANT I as high as ART 1 of flagell um
VI; ART 2 almost oval; 3-5 distal arts reduced. Basipodite of UR I twice as long as rami
Endopodite of UR III expanded and elongate iL
Genus Vibilia Milne-Edwards, 1830 males: Vibilia propinqua
Animals 5-15 mm. Head fairly small. Rostrum small 8a Integument weak. Eyes occupying about quaner
(if any). Eyes never occupying most of head surface.
ANT I with broad, laterally flattened flagellum. ANT
of lateral surface of head in females. ART I of
flagellum of ANT I clearly higher than peduncle
•
II inserted on anterior surface of head. MD with serrate Basipodite of UR I 1.5 times as long as rami
incisor, strong cylindrical molar. ART 3 of mandibular Rami of UR III subequal in males: Vibilia gib-
palp longer than ART 2. MX I with well-developed bosa
outer lobe, inner lobe small and rounded. MX II small, 9 Basipodite ofUR III longer than exopodite ...... l:
with both lobes. P VII with complete number of ART. 9a Basipodite ofUR III subequal to, or shorter than..
Seventeen species, 12 in the South Atlantic. exopodite ......................................................... 1i
10 ART 2 of P VII longer than distal ART com-
Key to the species of Vibilia bined. Distal process of ART 5 of P II reachin~
I Posterior lateral corners of last urosomite not middle of ART 6. ART I of flagellum of ANT :
produced rearward ............................................. 2 pointed, with parallel dorsal and ventral mar-
l a Posterior lateral corners of last urosomite pro- gins: Vibilia chuni
duced rearward ........................ .. ................ ........ 9 I Oa ART 2 of P VII shorter than distal ART com-
2 Distal process of ART 5 of P II reaching top of bined. Distal process of ART 5 of P II reaching
ART 6. [Eyes present) ....................................... 3 base of dactyl. ART J of flagellum of ANT :
2a Distal process of ART 5 of pereopod II 0.75 sharp, rhomboidal: Vibilia armata
times as long as ART 6, or shorter.. .................. 4 11 ANT 1 subequal to head and pereonite I com-
3 P III-VI thin and long, ART 4linear, dactyls not bined. Distal process of ART 5 of P II reaching
longer than third of ART 6; P V-VI at least quar- base of dactyl. Anterodistal corners of ART 5-6
ter longer than P III-IV; endopodite of UR Ill of P VII with rounded processes. Distal ART o;-
expanded and elongate in males: Vibilia antarc- p VII knife-like: Vibilia cultripes
tica lla ANT I longer than head and pereonite I com-
3a P HI-IV with robust, broad ART 4, dactyls sub- bined. Distal process of ART 5 of P II reaching
equal to or slightly sho1ter than ART 6; P V-VI middle of ART 6. Anterodistal corners of AR-:-
only slightly longer than P III-IV, rami ofUR III 4-5 of P VII without processes. Distal ART ofP
subequal in males: Vibilia viatrix VII linear with rounded top: Vibilia pyripes
4 Adult animals 5 mm or shorter ......................... 5
4a Adult animals longer than 5 mm ....................... 6 Vibilia antarctica Stebbing, 1888 (Fig. 4. 79
5 Eyes transformed into 3 rows of separate facets; Length to 14 mm. Head slightly higher than long.
flagellum of ANT I conical; ANT II 2-4 articu- Eyes small. ANT I slig htly shorter than head and
late, much shorter than ANT I: Vibilia australis first pereonite combined in females, longer ffi
Sa Eyes compact; ANT I with pointed but broad fla- males. First ART of flagellum with parallel dorsa:
gellum; ANT II 6-8 articulate, subequal in length and ventral margins. ART 6 of P I slightly longer
to ANT I: Vibilia stebbingi than 5, narrowing distally. Posterodistal process of
Amphipoda 1179
ART 5 of P n reaching distal end of ART 6 or reaching middle of ART 6. ART 4 and 6 sub-
more; strong dactyl equal to half or third of ART equal, ART 5 shorter in P ITJ and IV; dactyls
6. P V and VI subequal, longer than HI and IV ; strong, length equal to third of ART 6. P VII mo-
dactyls almost straight, equal to third of ART 6. P derately short; ART 2 longer than distal ART
VII twice as short as VI; ART 2 with straight ante- combined (unique character among Vibilia, but
rior margin, posterior margin convex, with distal see Vibilioides), with rounded posterior lobe pro-
lobe reaching middle of ART 4; ART 5 and 7 sub- duced to middle of ART 4; ART 5 and 6 subequal;
equal, slightly shorter than ART 6. T triangular, ART 7 narrow, finger-like, shorter than ART 6. T
with rounded top. triangular, with rounded top, not reaching midd le
Vibilia armata Bovallius, 1887 (Fig. 4.80). Length ofbasipodite ofUR III.
5-11 mm. Delicate animals. Integument semi-trans- • Vibilia cultripes Vosseler, 1901 (Fig. 4. 83).
parent. Head as high as long. Eyes dark, moderate- Length to 15 mm. Body robust. Integument dense.
ly large, especially in males, where they may occu- Head slightly higher than long. Eyes large. ANT I
py half of lateral surface of head. Flagellum of equal in length to head and pereonite I combined.
ANT I as long as head and pereonite I combined, ART I of flagellum egg-like laterally. ART 2 ofP
more or less rhomboid, sharp. ART 5 and 6 sub- I and 11 broadened. ART 6 of P l subequal to 5;
equal in P I, dactyl longer than half of ART 6. dactyl strong, slightly curved, as long as half of
Posterodistal process of ART 5 of P lT reaching ART 6. Posterior margin of ART 6 and dactyl ser-
base of dactyl. Anterior margins of ART 4-6 of P rate. ART 5 and 6 subequal in P II. Posterodistal
VI with numerous, very fine, setae; dactyl shorter process of ART 5 reaching middle of ART 6, ante-
than half of ART 6. P VII moderately long, reach- rior margin irregularly serrate. P VII as long as P
ing distal end of ART 6 of P VI. ART 5 and 7 of P IV. ART 2 of P VII subequal to ART 3-5 com-
VII subequal; ART 6 narrower than, and 1.5 times bined. ART 5-7 subequal. Anterodistal comers of
as short as, ART 5. T triangular, with rounded top, ART 5-6 of P VII with rounded processes with
reaching middle of basipodite of UR III. scales on their surfaces (unique character!). Distal
Vibilia australis Stebbing, 1888 (Fig. 4.81 ). ART of P VII knife-like, pointed distally, broad-
Length 3-5 mm. The smallest species of Vibilia . ened in middle, also with scales on surface.
Head small. Eyes transformed into three vertical Epimeral plate Ill with weak posterodistal denti-
rows of separate facets (unique character!). ANT I cle. T oval, broader than long.
subeq ual to head and pereonite I combined, • Vibilia gibbosa Bovallius, 1887 (Fig. 4.84). Length
peduncle broader than flagellum. Flage llum elon- to 7 (males) or 8 (females) mm. Integwnent thin.
gate, conical, two rudimentary apical ART easily Head higher than long, narrowing ante1iorly. Eyes
\isible. ART 2 of P I broadened; ART 6 oval, occupying about quarter of lateral surface ofhead in
longer than 5, dactyl slightly sh01ter than ART 6. females, about third in males. ANT I subequal to
Posterodistal process of ART 5 of P II not reach- head and pereonite I combined. ART I of flagellum
ing dista l end of ART 6. ART 4 of P m and IV of ANT I oval. ART 6 of P I with convex anterior
subequal to ART 6, ART 5 1:\vice as short. Dactyls margin, longer than ART 5; dactyl twice as short as
of P V and VI 3 times as short as ART 6. P VTI ART 6. Posterodistal process of ART 5 of P II
mice as short as VI, posterodistal lobe of ART 2 reaching half or two third of ART 6. ART 2 of P V
reaching middle of ART 4; ART 7 equal in length and VI linear; ART 4 and 5 subequal, 1.5 times as
tO 1:\vo thirds of ART 6, surface covered with fine short as 6; dactyl slightly longer than quarter of
setae. T triangular, with rounded top, reaching ART 6. Posterior lobe of ART 2 of P VII broadly
::niddle of basipoditc of UR lll. rounded, distal part slightly longer than ART 3;
~lb i/ia clumi Behning and Woltereck, 1912 (Fig. ART 7 finger-like, 1.5 times as short as ART 6. T
~.82). Length to 7.5 mm. Head higher than long, triangular, with rounded or broadly rounded top,
.:yes moderately small. ART 1 of flage llum of reaching middle of basipodite of UR III .
.-\..'\'T I broad with parallel dorsal and ventral mar- Vibilia jeangerardi Lucas, 1845 (Fig. 4.85) .
gins. distally pointed. ART 5 and 6 of P I sub- Length to ll mm. Head slightly higher than long.
~ua l, dactyls twice as short as ART 6. ART 5 of Eyes dark, moderately small, in males larger than
P ll longer than 6, posterodistal process of ART 5 in females. First ART of flagellum of ANT I very
1180 Amphipoda
broad, oval, higher than, and twice as long as, Vibilia robusta Bovall ius, 1887 (Fig. 4.88 .
peduncle. In P 1, ART 5 shorter than ART 6; dac- Length to 20 mm . The largest species of Vibilia.
tyl strong, slightly curved; twice as short as ART Integument very dense. Head higher than long.
6. In P IT, posterodistal process of ART 5 reaching Eyes very large, occupying most of lateral surface
3 quarters of length of ART 6, ART 6 subequal to of head. ART 1 of flagellum of ANT I not longer
5. P V and VI subequal, ART 2 moderately broad, than head, with parallel dorsal and ventral mar-
oval; ART 4 slightly longer than 5; ART 5 shorter gins, anteriorly rounded. Pereon as long as pleoc
than ART 6; dactyls short (shot1er than quarter of and urosoma combined. ART 6 of P r subequal to
ART 6). P VI and VII in ratio 9:7. ART 2 ofP VTI ART 5, with convex anterior margin. ART 2 of P
broad, oval; ART 7 slightly shorter than 6. Pleon II strongly broadened (width two thirds length):
longer than last 4 pereonites. Posterior corners of ART 5 subequal to 4, posterodistal process reach-
last urosomite rounded, not produced rearward. T ing only middle of ART 6; ART 6 twice as shon
broad, rounded, reaching middle of basipodite of as 5; dactyl twice as short as ART 6. ART 2 of P
UR III. III and IV broadened, posterior margins more con-
Vibilia propinqua Stebbing, 1888 (Fig. 4.86). vex than anterior; ART 4 subequal to ART 5, ART
Length to 12 mm. Int

(Demetrio Boltovskoy) South Atlantic Zooplankton

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(Demetrio Boltovskoy) South Atlantic Zooplankton

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South Atlantic

Edited by Demetrio Boltovskoy

Edited by Demetrio Boltovskoy

Volume 1: pages 1 - 868

© Backhuys Publishers, Leiden, The Netherlands 1999

Printed in The Netherlands

Bernardo Abiahy Demetrio Boltovskoy

José R. Dadon Ray Gibson

Fernando L. Medina Mantelatto Takashi Onbé

Roger Seapy Erik Thuesen

Vassily Spiridonov Siebrecht van der Spoel

Introduction now been more or less well established for over 50

South Atlantic Zooplankton.

South Subtropical •••• --- ••• -+

Biogeochemical provinces according to

Chlorophyll a Primary production Zooplanktonic biomass

10 General biological features of the South Atlantic

Percentage of total species recorded in the South Atlantic

Tropical-Subtropical Transition Subantarctic-Antarctic

200 300 400 500

Fig. 9. Biogeographic divisions of coastal South America according to various authors.

I-uu ul >50% Tropical Water

• >50% Subantarctic Water

~ >50% Coastal Water

Winter-Spr ing 1977

16 General biological features of the South Atlantic

Subtropical D Krohnitta subtilis

Surface salinity fields in the RIO de la Plata influence area

Typical salinity profile along the estuary (0/00)

t</·, I area sampled

AC: Angola Current

'"-- -------~----- ------------------

Chi a (mg m-3)

Planktonic biomass (rng ash-free dry weight m-2)

40 60 80 100 120 140 S

o 2 3 4 5 Prim. prod. (mg C m-3 h+)

0, Chi a (mg m-3)

Distance offshore (nautical miles)

100 " A"""""""""""":'"\",,C,,"'::"'"'

Great Warm Season

Temperature at 17 m A 0.9 Phosphate at 5 m c

Chlorophyll a B Primary production D

30 General biological features of the South Atlantic

Great Cold Season

plankton biomass to the edge of the shelf is related to Zooplankton communities

2000'S 1030' 1000' 0030' 0000' 0030'N

60 70 80 90 100 110 120E

34 General biological features of the South Atlantic

x: • ·\~__---.--.. f.~·~~· dinoflagellates

:;.I~/iiliiriliil·~_"'·'!!1,= --------.------ ...---

ments indicates considerable deposition of carbon. Part

Zooplankton studies in the Benguela Current have

and in the vicinity of Walvis Bay (23°S). Within these CJ <10

region. The mean biomass of zooplankton was esti- D<0.5

ONDJ FMAMJ J ASONDJ FMAMJ J AS ONDJ FM

the water column. It also serves to seed shelf commu- Phytoplankton

Chlorophyll (rng m-3)

Nitrates (rnrnol rn-')

Introduction 4 !lffi) and micropores (<1 !J.m) occur in non-spinose

South Atlantic Zooplankton.

So far, only sexual reproduction (gamete release) has

aooo oooo o o oooooo o ooo o o ooo o a o oo o o oo o oo oooo o Do o

Specimens per m-3

Dic_t_yacanthatabu_'a_ta . • -+I • t--- • -j- -+- ._ _. __