EDITORIAL COMMITTEE

for this Number

Frederick M. Bayer

Harding B. Owre Robert Austin Smith

Copyright© The University of M iami 1967

Library of Congress Catalogue Card Num ber 67-31694

Printed in the United States of America

P a r k e r P r i n t i n g , C o r a l G a b l e s , F l o r id a
 American
Opisthobranch Mollusks

Decem ber • 19 6 7
P u b l i s h e d by the U n i v e r s i t y of M i a m i
I n s t i t u t e of M a r in e S c ie n c e s
 P reface

In 1963, a series entitled Studies in Tropical Oceanography was estab­

lished at the Institute of Marine Sciences to include major papers in the
marine sciences which cannot be accommodated in journals because of
their size. The editors now take great pleasure in presenting as the sixth
volume in this series a major contribution to the knowledge of the marine
mollusks of American waters. Under the general title “American Opistho-
branch Mollusks,” by Eveline du Bois-Reymond Marcus and Ernest Mar­
cus of the University of Sao Paulo, Brazil, this volume comprises two parts.
The first deals with tropical American opisthobranchs, from Florida to
Brazil and including material from the faunistically related Bay of Panama,
and the second with species from the Gulf of California.
A total of 111 species and subspecies (70 in part I, 47 in part II, with
six common to both) is reported and for the most part described in full
and illustrated. Not until now has knowledge of the opisthobranch fauna
of the Western Atlantic reached the level of that of the shelled molluscan
groups, which were considered to be on the whole well known as early as
1910. The opisthobranchs of the Gulf of California must still be considered
incompletely known even after the present study, as sampling has been
inadequate in that large and environmentally diverse body of water.
The collections which form the basis for this volume were assembled by
various staff members of the Institute of Marine Sciences of the University
of Miami, Dr. Peter E. Pickens and associates of the University of Arizona,
and by colleagues of Professor and Mrs. Marcus in Brazil. New species
and new distributional records will, no doubt, continue to come to light
as further investigations are made, but this volume will remain as a land­
mark in the progress of our knowledge of the faunal areas concerned.
The fact that a large number of opisthobranch species have been based
upon anatomical details of shriveled and faded museum specimens is a
very discouraging aspect of systematic work upon these animals. It makes
tentative field determinations and identifications by non-specialists virtually
impossible. In the present volume, the authors have provided drawings of
many of the living animals made from photographs, which will be useful
to the non-specialist as well as to the systematist. Many species may be
determined with reasonable accuracy on the basis of external form and
color when the necessary illustrations of living animals can be published.
As a step in this direction, we are fortunate to be able to include a plate
of color figures in this book.
We are confident that users of this volume will share our appreciation
of the tremendous effort which its authors have devoted to its preparation.
The detailed descriptions, the significant illustrations, and the exhaustive
comparisons of all the species with previous records combine to enrich
the literature of these neglected mollusks.
The Editors
v
 TABLE OF CONTENTS

P re fa c e ......................................................................................................... v

Part I. Tropical American O pisthobranchs.......................................... 1

A b s tra c t.................................................................................................. 3
Introduction............................................................................................ 3
Explanation of L etterin g ...................................................................... 4
List of Gastropods Classified............................................................... 5
Systematic S ectio n................................................................................. 7
Zoogeographic R e m a rk s...................................................................... 123
Resumo .................................................................................................. 125
Zusammenfassung ................................................................................. 126
Literature Cited ................................................................................... 126

Part II. Opisthobranchs from the Gulf of C alifornia.......................... 139

A b s tra c t.................................................................................................. 141
Introduction............................................................................................ 141
Explanation of L etterin g ...................................................................... 142
List of Gastropods C lassified............................................................. 142
Systematic S ectio n................................................................................. 145
Zoogeographic R e m a rk s...................................................................... 236
Resumo .................................................................................................. 238
Zusammenfassung ................................................................................. 239
List of Type Specim ens........................................................................ 239
Literature Cited ................................................................................... 239

Index to taxa described or mentioned in Parts I and I I ........................ 249

vii
 E x p la n a tio n o f P la t e 1

(See page 2)

1, Chromodoris nyalya, spec, nov.; 2, Felimare bayeri, gen.

nov., spec, nov.; 3, Chromodoris tura, spec, nov.; 4,
Chromodoris neona Marcus; 5, Cyerce cristallina (Trin-
chese); 6, Micromelo undata (Bruguiere); 7, Tridachia
crispata Morch, olive green color phase; 8, Tridachia
crispata Morch, blue color phase; 9, Chromodoris neona
Marcus.
 P art I

Tropical American Opisthobranchs

 T ROPI CAL AME RI CAN OPISTHOBRANCHS
EVELINE a n d ERNST MARCUS
Department of Zoology, University of Sao Paulo, Brazil

A bst r ac t

A total of 70 species are described or listed, 68 opisthobranchs and two

prosobranchs (Titiscania limacina Bergh, from the Pacific side of the Canal
Zone, and a Lamellaria). They were collected chiefly on the coast of
Florida and neighboring seas; some are from the Gulf of Panama, where
faunistic relationships with the Caribbean Sea are apparent, and from
Brazil, Sao Paulo and Sta. Catarina. The opisthobranch faunas in the
intertidal zone of the West Indies and that of Brazil, at least in the rather
well known area Rio-Sao Paulo, have many species in common. Besides
the 2 new subspecies and 18 new species (one of which, Felimare bayeri,
represents a new genus), the following new names are introduced: Risbecia
francoisi (Risbec, 1928) for Risbecia odhneri Risbec, 1953; Anisodoris
rudberghi for A. marmorata Bergh, 1898b (non A. marmorata Bergh,
1881); Aporodoris risbeci for Thordisa hilaris Bergh, 1905, var. Risbec,
1953.

I n t r o d u c t io n

Professor Dr. Frederick M. Bayer sent us for identification a large

collection of gastropods belonging to the Museum of the Institute of
Marine Sciences of the University of Miami. In addition to 60 species of
opisthobranchs, it contains two species of prosobranchs belonging to
families traditionally studied together with opisthobranchs. The majority
of the animals came from Florida and the West Indies, although some were
collected on the Pacific side of the Canal Zone. These and five species
from Brazil expand the ambit of our zoogeographic remarks.
A rich collection of admirable color photos and notes referring to the
colors of the living animals accompanied the material. The photographs
were of very great help in classification and made possible descriptions of
many old species which had been known only in preserved condition.
Part of the material was collected under the auspices of various research
grants. Of particular importance were grants GB 1204, GB 3803, GB
5776, GB 20355, and GP 1363 for biological ship time awarded to the
Institute of Marine Sciences by the National Science Foundation, which
made possible the collection of material from deep water. Likewise, the
support of the National Geographic Society, under its programs for Coral
Reef Ecology and Deep-Sea Biology, was instrumental in the collection
of other material. Finally, the cooperation of numerous private individuals,
students and staff members of the Institute of Marine Sciences, in supplying
many interesting specimens collected personally, has made this collection
even more valuable.
3
 Thanks to arrangements made by Dr. Harald A. Rehder and Dr. Joseph
Rosewater, Division of Mollusks, Smithsonian Institution, United States
National Museum, the U.S. Department of Agriculture shipped the col­
lection to us, and it was returned with the kind help of Mr. W. Garth
Thorburn, Officer of the Foreign Agricultural Service.
We express our cordial thanks to Dr. Erich Schulz, Zoological Institute
and Museum of the University Kiel, who presented us with an excellent
copy of Bergh’s paper on Titiscania limacina (1890).
We thank Professor Dr. Frederick M. Bayer for revising the language
in our manuscript and for reading the proofs.
All specimens and slides were returned to the Marine Laboratory,
University of Miami. The types are deposited in the U.S. National
Museum, Washington, D.C.

E x p l a n a t io n of L e t t e r in g

a— ampulla iz— “reinforcement sac”

ac— pallial caecum j— foot
ai— female atrium k—kidney
am— male atrium m—mouth
an— albumen gland ma— male aperture
ao— female follicle me— mantle cavity
ar— anus mi— male follicle
as— ascus mo— everted genital atrium
av— glandular band ms— left posterior oral gland
b— bursa or spermatheca mu— muscular part of oesophagus
be— fertilization chamber mx— mucus gland
c— genital aperture mz— muscular papillae
ca— caecum n— nephropore
cc— crop nc— pedal nerve
ce— brain ne— capsule gland
cl— seminal groove no— dorsal glands
co— connective hepatic duct nr— prebranchial gland
cr— collar ns— accessory sac
cs— cephalic shield o— oesophagus
cu— ciliated cushion oa— anterior pedal gland
cv— fore end of foot oi— ovotestis
d— male duct on— flagellar oviducal (vaginal) gland
e— muscular part of efferent duct oo— ovary
ei— hermaphrodite duct or— osphradium
eo— pigment patch os— oesophageal gland
es— penial sheath ou— blood (or nephridial) gland
ev— oesophageal pouch ov— oviduct
/ — pharynx p— penis
g— female gland mass q— prostate or prostatic section of d
h—Hancock’s organ R—rhachidian tooth
I—intermediate tooth r—right liver
i— intestine ra—radula
ic— infrapallial ridge re— right hepatic duct
ie— digestive gland ri— rhinophore
is— tentacle rn— mantle border
iu— infrapallial lobe ro— retractor
4
rr—border of sole ve—ventricle
rs—renopericardial canal vn— vaginal gland
sa—salivary gland v.v—flap
se—main hepatic duct vz—spines of vaginal gland
si—penial sac wa—female aperture
so— stomach wo—body wall
sx—sphincter x—appendage of rhinophore sheath
t—seminal receptacle xo— insemination duct
u—caecum of ei y—spermatocyst
ur—transverse furrow of foot yg—hypobranchial gland
v—vagina z—spermoviduct
va—vaginal gland zi—gill
vc—visceral cavity
L i s t o f G a s t r o p o d s C l a s s i f i e d in t h e P r e s e n t P a p e r
Subclass PROSOBRANCHIA ( =Streptoneura)
Order ARCHAEOGASTROPODA ( =Diotocardia)
Superfamily N e r i t a c e a
Family Titiscaniidae
1. Titiscania limacina (Bergh, 1875). Figs. 1-5.
Order MONOTOCARDIA
Suborder M e s o g a s t r o p o d a ( =Taenioglossa)
Superfamily L a m e l l a r i a c e a
Family Lamellariidae
2. Lamellaria perspicua (Linne, 1758). Fig. 6.
Subclass EUTHYNEURA
Superorder OPISTHOBRANCHIA
Order CEPHALASPIDEA ( =Tectibranchia, s. str.)
Superfamily A c t e o n a c e a
Family Hydatinidae
3. Micromelo undata (Bruguiere, 1792). Plate 1, Fig. 6.—Fig. 7.
4. Hydatina vesicaria (Solander, 1786). Figs. 8-9A, B.
Superfamily P h i l i n a c e a
Family Aglajidae
5. Aglaja pusa, spec. nov. Figs. 10-12.
6. Chelidonura hirundinina (Quoy & Gaimard, 1833).
Superfamily B u l l a c e a
Family Atyidae
7. Haminoea antillarum (d’Orbigny, 1841). Figs. 14-19.
8. Haminoea elegans (Gray, 1825). Fig. 13.
Order ASCOGLOSSA ( =Saccoglossa)
Superfamily O x y n o a c e a
Family Oxynoidae
9. Oxynoe antillarum Morch, 1863. Fig. 20.
10. Lobiger souverbiei P. Fischer, 1856.
Superfamily E l y s i a c e a
Family Elysiidae
11. Elysia evelinae Marcus, 1957. Fig. 21.
12. Elysia papillosa Verrill, 1901. Figs. 22-25.
13. Elysia cauze scops, subsp. nov. Figs. 26-27.
14. Elysia tuca, spec. nov. Figs. 28-32.
15. Elysia duis, spec. nov. Figs. 33-37.
Elysia spec.
16. Tridachia crispata Morch, 1863. Plate 1, Figs. 7, 8,.—Fig. 38.
5
 17. Tridachiella diomedea (Bergh, 1894). Figs. 40-47.
Family Caliphyllidae (Phyllobranchidae)
18. Cyerce antillensis Engel, 1927. Fig. 39.
Order ANASPIDEA
Family Aplysiidae (Subfamily Aplysiinae)
19. Aplysia (Varria) dactylomela Rang, 1828.
Family Aplysiidae (Subfamily Dolabriferinae)
20. Dolabrifera dolabrifera (Rang, 1828).
21. Petalifera ramosa Baba, 1959. Fig. 48.
22. Phyllaplysia engeli Marcus, 1955. Figs. 49-50.
Family Aplysiidae (Subfamily Notarchinae)
23. Stylocheilus citrinus (Rang, 1828).
Order NOTASPIDEA
Superfamily U m b r a c u l a c e a
Family Umbraculidae
24. Umbraculum plicatulum (v. Martens, 1881). Fig. 51.
Superfamily P l e u r o b r a n c h a c e a
Family Pleurobranchidae (Subfamily Pleurobranchinae)
25. Berthella tupala Marcus, 1957. Fig. 52.
26. Berthellina quadridens (Morch, 1863). Fig. 53.
27. Pleurobranchus (Pleurobranchus) areolatus Morch, 1863.
Family Pleurobranchidae (Subfamily Pleurobranchaeinae)
28. Pleurobranchaea occidentalis Bergh, 1897. Figs. 54-55A-C.
29. Pleurobranchaea hedgpethi hamva Marcus, 1957. Fig. 56.
30. Pleurobranchaea agassizii Bergh, 1897. Figs. 56A-D.
Order DORIDOIDEA
Suborder E u d o r i d a c e a
Tribe C r y p t o b r a n c h i a
Family Dorididae (Subfamily Chromodoridinae)
31. Cadlina rumia Marcus, 1955.
32. Chromodoris neona Marcus, 1955. Plate 1, Figs. 4, 9.—
Figs. 57-58.
33. Chromodoris nyalya, spec. nov. Plate 1, Fig. 1.—Figs. 62-66.
34. Chromodoris tura, spec. nov. Plate 1, Fig. 3.—Figs. 59-61.
35. Hypselodoris edenticulata (White, 1952). Figs. 67-71.
36. Hypselodoris acriba, spec. nov. Figs. 72-75.
37. Felimare bayeri, gen. nov., spec. nov. Plate 1, Fig. 2.—Figs.
76-79.
Family Dorididae (Subfamily Doridinae)
38. Siraius kyolis, spec. nov. Figs. 80-84.
Family Dorididae (Subfamily Discodoridinae)
39. Anisodoris worki, spec. nov. Figs. 85-89.
40. Anisodoris prea, spec. nov. Figs. 90-93.
41. Peltodoris greeleyi MacFarland, 1909. Figs. 94-98.
42. Discodoris evelinae Marcus, 1955.
43. Discodoris hedgpethi Marcus, 1959. Figs. 98A-E.
44. Discodoris phoca, spec. nov. Figs. 99-101.
45. Discodoris purcina, spec. nov. Figs. 102-104.
46. Discodoris pusae Marcus, 1955. Figs. 105-107.
47. Discodoris aurila, spec. nov. Figs. 108-111.
48. Taringa telopia disay subspec. nov. Figs. 113-114.
49. Taringa aivica, spec. nov. Figs. 115-119.
Family Dorididae (Subfamily Halgerdinae)
50. Aphelodoris antillensis Bergh, 1879. Figs. 120-121.
Family Dorididae (Subfamily Platydoridinae [Arginae])
 51. Platydoris angustipes (Morch, 1863). Fig. 112.
Tribe P h a n e r o b r a n c h i a
Superfamily S u c t o r i a
Family Corambidae
52. Doridella obscura Verrill, 1870. Fig. 122.
Suborder P o r o s t o m a t a
Family Dendrodorididae
53. Dendrodoris krebsii (Morch, 1863).
54. Doriopsilla janaina, spec. nov. Figs. 124-125.
Family Phyllidiidae
55. Phyllidiopsis (Phyllidiopsis) papilligera Bergh, 1890. Fig. 123.
Order DENDRONOTACEA
Family Tritoniidae
56. Tritonia (Tritonidoxa) wellsi Marcus, 1961. Figs. 126-130.
57. Tritonia (Candiella) bayeri, spec. nov. Figs. 131-134.
58. Marionia cucullata (Gould, 1852).
Family Bornellidae
59. Bornella calcarata Morch, 1863. Fig. 135.
Family Dotoidae
60. Doto caramella Marcus, 1957. Figs. 136-137.
Order ARMINOIDEA
Suborder E u a r m i n o i d e a
Family Arminidae
61. Annina miilleri (v. Ihering, 1886).
Order EOL1DOIDEA
Suborder A c l e i o p r o c t a
Family Fionidae
62. Fiona pinnata (Eschscholtz, 1831).
Suborder C l e i o p r o c t a
Family Facelinidae
63. Phidiana lynceus Bergh, 1867. Figs. 138-140.
Family Favorinidae (Subfamily Facalaninae)
64. Godiva rubrolineata Edmunds, 1964. Fig. 141.
65. Austraeolis catina, spec. nov. Figs. 142-148.
Family Favorinidae (Subfamily Favorininae)
66. Cratena kaoruae Marcus, 1957.
Family Aeolidiidae
67. Aeolidiella lurana, spec. nov. Figs. 149-150.
68. Spurilla neapolitana (Delle Chiaje, 1823).
Family Glaucidae
69. Glaucus atlanticus Forster, 1777.
Order SOLEOLIFERA
Superfamily O n c h i d i a c e a
Family Onchidiidae
70. Onchidella floridana (Dali, 1885). Figs. 151-155.
Systematic Section
1. Titiscania limacina Bergh, 1875
Figures 1-5
Reference.— Bergh, 1890b: 3.
Material.—Canal Zone (Pacific coast): Perico Island, Ft. Amador; under
7
rocks at low water. F. M. Bayer and R. W. Bayer, December 25, 1962.
Four females.
Further distribution.— lie aux Fouquets, sea of Mauritius; Philippine Is­
lands, Camiguin, 18° 55' N, 121° 50' E (original locality); Marshall
Islands, Eniwetok Atoll, U. S. National Museum, numbers 1502 (574920)
and 1503 (574923).
Description.— In the present account we will not repeat all of Bergh’s
observations. The length of the preserved animals reaches 9 mm (linear
measurement) to 11 mm (measured over the back), the breadth 3 mm,
the height 3 mm. The tentacles measure 1.5 mm, in life 3 mm (from color
photo); the small eyes lie at their bases. Sections show the eyes to be closed
vesicles containing homogeneous vitreous material. On the photograph
(Fig. 1) the transparent margin of the foot is visible in dorsal view at the
sides of the white body and in front of the small head. The hind end is
rounded.
The opening of the pallial cavity is a curved slit close behind the head.
The convex roof of the mantle cavity occupies about V x of the body length.
Between the forehead and the mouth (m) the right and left margins of the
foot coalesce, forming a veil set off by a deep fold on either side (Figs.
1, 2 ).
On both sides of the back a row of about 12 densely white papillae are
the outlets of big, folded sacs (Fig. 3, no) located in the connective tissue.
These sacs are lined with epithelium containing pink-staining secretion.
When squeezed, the papillae emit clouds of white granules. There are tiny
goblet cells in the epidermis of the back and the sole, the former with
pink-staining granules, the latter with blue-staining threads. Behind the
mouth are the outlets of blue-staining glands, which lie in the connective
tissue of the foot as a compact mass, the anterior pedal gland. The cteni-
dium comprises 35-40 leaflets as in Bergh’s material. In front of the gill
and to its left the inconspicuous osphradium forms a linear ridge in the
pallial roof.
The buccal cavity is ample, the buccal mass (/) longish, with cartilages
(Bergh, 1890b: 8). The radula (ra) comprises about 220 rows (Bergh:
180-205). The rhachidian tooth and two inner lateral teeth are absent.
The third lateral is small, the fourth a little bigger, and the fifth is domi­
nating as in all Neritacea. The marginal teeth are as described by Bergh.
The oesophagus (o) begins flat; its dorsal food channel is limited by two
ample folds. To the sides of the folds there are two broad pouches (ev).
These receive the two short and wide ducts from an unpaired ventral sac
(ay), a spacious glandular organ, Bergh’s buccal gland. The posterior
oesophagus is thick-walled and coated with red-staining glands; it opens
together with the intestinal gland into the stomach as in other Neritacea
(Thiele, 1910a: 353; Fretter & Graham, 1962: Fig. 119 B ). The stomach
8
F i g u r e s 1-5. Titiscania limacina Bergh. 1, living slug, from color photo; 2 ,
side view of preserved head; 3, transverse section 1.5 mm from hind end;
4, diagram of buccal mass and oesophageal gland in ventral view; 5, diagram
of female reproductive organs.— F i g u r e 6 . Lamellaria perspicua (Linne);
animal from color photo.
9
and the intestine contain many loose marginal teeth of the radula. The
intestine passes through the ventricle; there is an auricle on either side, as
already supposed by Bourne (1908: 816).
The anal papilla (Fig. 5, ar) separates the fundus of the pallial cavity
(me) into a right and a left niche. The right niche lodges the nidamental
aperture (wa), the left one the pores of the bursae ( b) on the roof and
the strongly ciliated renal opening (n ) on the floor. The posterior body
cavity is prolonged beyond the ovary into a thin tube 1.5 mm long (Fig.
3, vc).
The ovary (oo) ends about 3 mm from the hind end of the body; the
ovarian follicles are separated by connective tissue. The oviduct (ov) bears
longitudinal folds; its ovarian section is quite short and not ciliated,
whereas the following, presumably renal oviduct is ciliated. We found no
communication with the far distant pericardium. Where the ciliated oviduct
passes to the glandular pallial section, hangs a flat, ciliated pouch (t) with
a broad opening, 0.3 mm long and wide. This pouch is empty in our sec­
tions; it is probably a receptaculum seminis. The pallial oviduct bears
numerous glandular diverticula. The narrower, left ones (an) contain pink
granules as an albumen gland, and the broader right ones (ne) blue
secretion as a capsule gland. In the latter a ciliated ventral channel reaches
the receptaculum. To the nidamental (i.e., outer) part of the oviduct a
bag (iz) is attached. It is 0.2 mm long, 60 fi broad, empty and nonciliate.
Possibly this organ is a crystal (or reinforcement) sac. Its epithelial nuclei
are basal, the upper parts of the cells coarse, pink, and probably secretory.
Near the bag is a spoon-shaped flap (vs) whose concavity is directed
inwards. It is a fold of the oviducal wall. The outer end of this flap,
standing out from the nidamental opening (wa), is fastened to the rectum
(/). A band of red-staining glands (av) begins close to the opening of the
crystal sac (iz), traverses the cavity of the spoon (vs) and ends at its upper
border. Possibly the egg capsules are moulded in the flap.
In the roof of the pallial cavity to the left of the anal papilla (ar) there
are about 5 pores of long ciliated tubes (v). These branch and lead into
about 10 spherical vesicles (b) separated by a small amount of connective
tissue and muscle fibers. Some of the vesicles are lobed or coalesced.
The nuclei of their epithelial cells are giant (30 ^ ) . Their lumina con­
tain unorientated sperms. We did not see any sperm resorption. The
organs are multiple copulatory bursae.
Discussion.— This unique, nonpelagic, free-living prosobranch without
shell was first drawn by Carl Semper during his voyage to the Philippines.
Bergh (1875a: PI. 41, Fig. 10) published this figure with no more text
than: “Titiscania limacina, Bgh. Nebenbei Maassstab”. The figure was
copied by Bergh (1890b: PL 1, Fig. 1), Simroth (1896-1907: PI. 1,
Fig. 3), Pelseneer (1906: Fig. 131), Grasse (1961: Fig. 366C) and
others. It differs from Professor Bayer’s color photograph in that the
10
fundus of the mantle cavity is farther behind, and in the absence of the
pedal veil and of the dorsal glands. Later on, the latter were observed as
white dots (Bergh, 1890b: 4), especially in the ample material from
Mauritius. The pedal veil, however, is not shown in the pertinent draw­
ing (PI. 1, Fig. 2), where the foot is much narrower than in our spe­
cimens. The corresponding text (translated)— “anterior end of head
flattened, thinned out towards the almost fringed border”— though not
explicit, suggests that a pedal veil occurs also in the Indo-West Pacific
slugs. When in 1960 we classified some opisthobranchs from Eniwetok
for the U.S. National Museum, we did not examine the details of the two
specimens mentioned above (Acc. No. 197874). In these, the ctenidium,
evidently very mobile as in other Neritacea (Yonge, 1947: 475), pro­
jected from the pallial cavity as in the original and several other spe­
cimens (Bergh, 1890b: 4).
A comparison of Bergh’s description with ours reveals further differences
concerning the eyes, the osphradium, the oesophageal gland, the paired
auricles, and the appendages of the oviduct. We do not consider these
differences as specific characters, but as results of the different methods
of studying the slugs by anatomical dissection and by reconstruction of
microtomic sections respectively.

2. Lamellaria perspicua (Linne, 1758)

Figure 6
Reference.— Marcus, 1960b: 189.
Material.— Florida: Palm Beach Inlet; under rocks at east side of Peanut
Island. R. C. Work. April 20, 1963. One specimen examined but two
are shown on color photo.
Further distribution.— Dry Tortugas; Virginia Key; Curasao; Southern
middle Brazil (forma mopsicolor du Bois-Reymond Marcus, 1958: 11);
northern Patagonia; Straits of Magellan, and Southern Chile. Eastern
Atlantic Ocean, including the Mediterranean Sea, but not the Arctic. Indo-
West Pacific Ocean. From the intertidal zone to 1287 m.
Description.— The preserved male is 11 mm long, 9.3 mm broad, and
4 mm high. The colorless mantle is transparent and slightly bossed. The
dried shell shows growth lines and is covered with an iridescent, color­
less periostracum. It measures 8 mm in length, 5.3 mm in width, and 3.2
mm in height. It has a little less than three whorls and the columella is
open up to the apex (“patente usque in fundum”— Bergh, 1853: 90).
The left limb of the rhachidian tooth is a little longer than the right
one, and its cusp slants somewhat to the left. There are 3-4 denticles on
either side. The cusp of the lateral teeth bears 3-4 denticles on the upper
edge, 12-16 on the lower edge. The lower or outer denticles vary widely
11
in size from one tooth to the next. The penis exhibits a terminal protrusion
of the male duct, as is often the case in preserved Lamellaria.
The bigger animal on the color photo is vermilion with white blotches
and specks. The blotches are generally longish, of quite irregular outlines
and size, and form a vaguely radial pattern. The smaller one is pink with
dark red dots. Dense marginal bands of the mantle produce a radial pat­
tern of alternate pink and violet zones (Fig. 6), because the animals were
photographed on blue ground.
Remarks.— Though the colors of the present material (red, white, pink,
and violet) are unusual, they all have been recorded for L. perspicua
(Bergh, 1853: 81; 1886: 166-167) from the Mediterranean Sea. Orange
with white occurs more frequently.

3. Micromelo undata (Bruguiere, 1792)

Plate 1, Fig. 6; Figure 7, A-F

References.— Pilsbry, 1893-95: 391-92; Bergh, 1901: 252; Abbott, 1955:

276; Coomans, 1958: 102; Warmke & Abbott, 1961: 141; Jong & Kris-
tensen, 1965: 47.
Material.— 1. Florida: Eastern Dry Rocks, off Key West, under rocks,
1 m. Chris Dermott, May 20, 1967. One specimen.— 2. Bonaire (Hum-
melinck, 1953: 63, locality no. 1055). P. Wagenaar Hummelinck, Sep­
tember 4, 1948. One animal without shell.— 3. Curasao: Piscadera Bay,
outer bay, from Halimeda. Diva Diniz Correa, February 15 and 28, 1966.
Two snails.
Further distribution.— Bermudas; Lower Florida Keys; Antilles from Cuba
and Puerto Rico to Grenada, Bonaire and Curasao; NE Brazil, Per­
nambuco, Alagoas.
Description.— The living snails are transparent white, the borders trimmed
with yellow, opaque white spots on the head shield (cs) and the infra-
pallial lobe (iu), still present in the preserved snails. These spots are
heaps of calcareous granules, 10-20 ^ in diameter. The anterior part of
the head shield is blackish in the preserved animal from Florida. Measure­
ments (in mm) of the preserved snails are: Locality 1: 7 long, 5 wide,
4.3 high; locality 2:7, 3.5, 3; locality 3: (a) 3.1, 2.3, 2.4; (b) 3.1, 2.1,
1.6. The shells are 6 X 4 (Florida) and 2.8 X 2.2 (larger snail from
Curasao). Well preserved animals such as that from Florida are not
twisted out of the shell easily, as Bergh indicated (1901: 242, 252).
The shell was described well by Pilsbry (loc. cit.). In preserved con­
dition, that from Florida is opaque white with 3 reddish brown spiral lines
and 4 rows of axial bows. Between every 2 red lines there are 6-8 spiral
12
F igure 7. Micromelo undata (Brug.): A , shell; B, outside view of freed larval
shell; C, inside view of same; D, side view of preserved snail without shell;
E, anterior part of right side, mantle border lifted; F, diagram of organs in
mantle roof; G, radular teeth; H, combined section of male copulatory organs.
13
rows of pits, and many such occur in front of and behind the red lines.
At the beginning of the body whorl these rows are nearer together than
on the outer lip (Fig. 7,A ). The columellar margin is inflected, and there
is a narrow glaze on its hind part. The large transparent and subimmersed
protoconch is distinct (Fig. 7,B). It is 0.75 mm long and begins with 2
sinistral whorls. At the beginning of the third whorl the shell turns to the
right, and at its end the teleoconch begins, sharply set off from the proto­
conch by a bulge. A broad black spiral line appears approximately at the
beginning of the second whorl on the underside which is tilted towards
the top. At the end of this whorl the line accompanies the turn of the
shell to the right and bifurcates into two axial lines which unite again on
the left side that lies inwards (Fig. 7,C). The red lines of the teleoconch
arise independently from the black ones.
The head shield (c,s) is deeply bipartite (Fig. 7,D). The small eyes
are subepidermal and in clarified snails seen in front of the long and wide
posterior lobes whose ends are rounded. There are two pairs of lateral
appendages, the bulge-shaped so-called tentacles (Fig. 7,E, is) on either
side of the mouth, and the rolled so-called rhinophores (ri) farther be­
hind. Under each rhinophore lies the feathered Hancock’s organ (h ). The
nervous system corresponds to that of Hydatina (Aplustrum) velum (Gm.)
studied by Eales (1938: 79, 80). Her figure completes the earlier descrip­
tions summarized by Hoffmann (1936: 649-50).
From the thickened mantle border (rn) the infrapallial lobe (iu) ex­
tends as a wide frill under the shell to the left side, corresponding to
Bergh’s indication “besonders der Schwanz sehr gross” (loc. cit.). On the
right side the caecum (ac), Bergh’s strong “spiralige Druse,” lies between
the shell and the mantle. In the specimen from Bonaire it is 2.5 mm long,
and its glands form stripes visible through the skin. In the roof of the
ample pallial cavity (Fig. 7,F) the hindmost organ is the anus (ar). The
epithelium lining the cavity and the surface of the mantle are separated
by blood spaces subdivided by strands of muscle. In front of the anus
extends the gill (zi). It is lower and thinner in its inner half than in the
outer, and more similar to that of Acteon (Perrier & Fischer, 1911: 20
ff.) than to the nearly feathery one of Aplustrum (ibid.: PI. 2, Fig. 1).
In front of the ctenidium is the renal opening (n ), not thickened, and
still farther forward the osphradium (or) more to the right, and the well
ciliated renopericardial canal (rs) to the left. The heart separates the
right and the left lobe of the kidney (k ). The foremost part of the right
lobe is connected with a gland (ou) located between the pericardium and
the intestine. Bergh (p. 248) called it “blood gland,” but generally it is
termed nephridial gland in spite of its distance from the kidney (Fretter
& Graham, 1962: 285). Farther to the right lies the flat hypobranchial
gland (yg) richly supplied with blood spaces. The outermost gland of
the mantle edge (rn) is a thick bulge.
14
 The foot (Fig. 7,D, j) is much folded and shorter than the visceral
sac; the so-called parapodia (Hoffmann, 1933: 252) are big.
The black mouth tube was everted in the specimen from Bonaire. The
posterior glands of the mouth tube (ptyalin glands, Eales, 1938: 80) and
the salivary glands are long coiled tubes, the former rather thick, the
latter slender. The jaws are oval plates covered with spiny platelets. The
snail from Bonaire has the radula formula 21X12.0.12. In two or three
rows there was a slight rhachidian thickening. The number of denticles
varies (Fig. 7 ,G ): the 3 outermost teeth have none, the 9th tooth has 1,
the 8th 1-2, and the 7th 1-3 denticles. Teeth 6-2 have 3-5, the innermost
up to 6 denticles. Those of the innermost tooth are hardly smaller than
the small principal cusp; in the specimen from Brazil (Bergh, 1901: 253,
PI. 23, Fig. 9) the difference between the principal cusp and the denticles
of the innermost tooth is much more pronounced. The stomach is thin-
walled; its lining not cuticularized. In the specimen from Bonaire it con­
tained coiled masses of a polychaete. The velvety black intestinal gland
shines through the white flesh of the young snails from Curasao.
The specimen from Florida is in male phase; its ampulla (vesicula
seminalis) contains autosperms. The ovotesticular lobes are completely
surrounded by the intestinal gland. The male copulatory organ (Fig. 7,H)
lies under the right posterior of the lateral cephalic appendages. The
efferent duct is closed. It has two ental glandular portions (q ) staining
differently, followed by a muscular one (d ) which coils through a promi­
nent cutaneous papilla, whose skin is folded longitudinally. The duct ends
with a long narrow tip which is supported by a cushion of vesicular cells.
Probably the tip can be retracted into the hollow outermost part of the
papilla. At the root of the papilla lies the female aperture ( m ) which is
in common for the vagina (v) and the oviduct (ov).
Discussion.— The differences between the shells of M. undata and M.
guamensis (Quoy & Gaimard, 1824) are indicated in Pilsbry (loc. cit.:
393). The radular teeth of the Indo-Pacific species have especially small
denticles (Habe, 1950: PI. 3, Fig. 14). Thiele (1931), Habe, Zilch
(1959-60), and the above-quoted conchologists maintain Micromelo,
while Bergh (loc. cit.) unites it with Aplustrum Schumacher, 1817. Bergh
(p. 241) and Vayssiere (1906: 46) considered Aplustrum and Hydatina
Schumacher, 1817, as one and the same genus, and used the name
Aplustrum, published 22 pages later than Hydatina.
Professor Bayer’s color photo is the first illustration of a living M.
undata. When Pilsbry (loc. cit.: 386) introduced the genus, he wrote:
“animal with two tentacles.” He based this statement upon Quoy’s drawing
of Bullaea guamense which he copied (PI. 59, Fig. 26). Unfortunately
this drawing is not quite clear (Pruvot-Fol, 1934: 28). The two pairs of
lateral cephalic appendages were described by Bergh (1901).
15
 4. Hydatina vesicaria (Solander, 1786)
Figures 8, 9A, 9B
References (to the shell).— Abbott, 1955: 276; Coomans, 1958: 102;
Warmke & Abbott, 1961: 141.
Material.— O ff French Guiana, 6° 00' N, 52° 27' W, 64 m. O re g o n
Station 4192, February 22, 1963. Four snails; the biggest shell 37 mm
long, 29 mm broad, with several barnacles on it.
Further distribution .— Bermuda (Coomans); south half of Florida, and
West Indies (Abbott); Brazil, Bahia (v. Ihering, 1915: 139). Cape Verde
Islands, S. Vicente (Pilsbry, 1893-1895: 387) may be included in the
range of vesicaria.
Remarks .— D’Orbigny (1853: 132) considered his material of H. physis
from Cuba and the Antilles as different from that from the Indo-West
Pacific Ocean, but did not indicate the difference. He maintained the name
physis for the West Indian shells and gave those from the Indo-West Pacific
the name quoyana. Pilsbry (1893-1895: 388) and Kobelt (1896: 133)
were unable to find any constant differences between occidental and
oriental shells, and therefore Johnson (1934: 148) listed H. physis from
West Atlantic warm waters.
We do not know any reference to the anatomy of the occidental Hy­
datina and, therefore, we compared the present material with the descrip­
tions of Bergh (1901: 247), Vayssiere (1906: 50), and Risbec (1928:
37; 1951: 145) of material from the western Indian Ocean, the Gulf of
Aden, and New Caledonia, respectively. Of the latter material, none of
the characters which reveal specific differences between physis and
vesicaria is recorded. These concern the Hancock’s organ, the oral tube,
and the radula. The first, Bergh’s osphradium, has in physis about 15
evidently simple leaflets on either side, and may be mamillate in shriveled,
preserved specimens (Vayssiere, PI. 3, Fig. 45). In the much bigger
Hancock’s organ of vesicaria the high leaflets bear secondary pinnules,
and the surface of the organ becomes meandrous (Fig. 8) by contraction.
The oral tube of vesicaria is longer (3 cm) than in Bergh’s (1-2 cm) and
Vayssiere’s (7 mm) physis. Bergh (p. 248) indicated the salivary glands
and those of the oral tube of H. physis to agree with the corresponding
organs in H. velum (Gmelin, 1791), where both are paired (p. 245).
Vayssiere (p. 54), however, described an unpaired dorsomedian gland
of the oral tube and paired salivary glands in physis, which we found
also in vesicaria.
The radula of vesicaria (Fig. 9A,B) comprises 40 rows and 20 teeth
per halfrow. The teeth of the innermost longitudinal row have up to 5
cusps, those in the middle generally 3, occasionally 4, and the 5-6 outer­
most rows 2 cusps.
16
F ig u r e s 8-9. Hydatina vesicaria (Solander). 8 , left side of head; 9A, longi­
tudinal row of radular teeth; 9B, central part of radula.

The most important difference between physis and vesicaria refers to

the rhachis of the radula: naked in physis, but bearing a tiny pointed,
somewhat irregular, tooth in vesicaria. The long base of the lateral teeth
in vesicaria is probably no more than a gradual difference from physis,
because one tooth of Bergh’s figure 42 (PI. 20) has a similar long base.
A peculiar connection between the teeth of the longitudinal rows in
vesicaria by fibers of the radular membrane was not mentioned by Bergh
nor by Vayssiere, but can be inferred as present also in physis from some
of their drawings of radulae.
The numerous references to H. physis in Hoffmann (1932-1940) were
compared with the corresponding organs of vesicaria, with exception of the
17
central nervous system. They do not furnish further specific differences.
Neither does the male copulatory organ (Bergh, 1901: 249; Vayssiere,
1906: 52; Risbec, 1951: 147).

5. Aglaja pusa, spec. nov.

Figures 10-12
Reference.— Marcus, 1966: 170. Brief descriptive remarks written later
than the present description but unfortunately published earlier.
Material.— Florida: Biscayne Bay, off West Point, Key Biscayne. R. E.
Young. February 28, 1964. One specimen.
Holotype.— USNM 576279.
Description.— The preserved animal is 45 mm long, 19 mm broad, 18
mm high. The color of the dorsal and ventral skin is a greyish black,
interrupted by numerous pigment-free spots of various sizes, some oblong,
some round with a dark center (Fig. 10). Fusion of these light areas
produces local vermiculated designs. The gill is colorless.
The shell is yellow with brown growth lines, longer (15 mm) than
broad (13 mm), thin, but completely calcified (Fig. 11). The inner
margin bears three spine-like dorsal processes, the right one the largest.
The backward-directed wing of the whorl is rather long and pointed, its
inner border thickened. The downward-directed spur on the left posterior
border is pointed also. The outer lip has a concavity on the right side,
on the level of the right spine.
The head shield is nearly rectangular, half the length of the body. On
either side of the mouth there is a bulge with tufts of cilia, the terminations
of sensory cells as described in Chelidonura evelinae Marcus (1955: 96).
In the groove between the cephalic shield and the head-foot lies a pig­
mented longish area, the pinnate (though weakly developed) Hancock’s
organ. An axial, pigment-free furrow emits ventral and dorsal furrows.
The former are transverse, in a right angle to the axis, the latter nearly
parallel to it. The median slit separating the posterior mantle lobes is
deeper ventrally than dorsally. The left lobe has no ffagellum.
The male atrium (am) is folded longitudinally; it contains a few melano-
phores, and is continuous with the loose penial sac (Fig. 12, si). The cili­
ated fundus of the sac lodges the spirally coiled penial papilla (p); its
inner wall bears a ciliated cushion ( c m ), White’s anterior lobe of the
penis (1946: Fig. 4), a cavernous organ built of vertical strands of
muscles separated by blood lacunae. The seminal groove (ci) runs along
the ventral border of the sac (Fig. 12) to the beginning of the coil, where
it receives the duct of the tubular prostate (q). The retractor (ro) inserts
on the dorsal wall of the penial sac.
18
F ig u r e s 10-12. Aglaja pusa, sp. nov. 10, dorsal view from color photo; 11,
shell; 12, male copulatory organ.

Discussion.— Size, color, shell, and male copulatory organ are the principal
elements by which the species of Aglaja are distinguished. The color may
vary, e.g., in A. tricolorata (Pruvot-Fol, 1954: 51) and A. cyanea (Mac-
nae, 1962: 193); in other species it is constant and is preserved rather
well in alcohol (Bergh, 1893: 114, 126, note 1). The shell is easily broken,
and the outer rim (often of conchiolin), adheres to the wall of the shell
cavity and is hardly delimited. Spines on the inner rim of the shell,
as in A. pusa, have not been described in previous species. The male
copulatory organ of A. pusa is similar to that of A. ceylonica White
(1946: 167); this organ is not described for all species. Pruvot-Fol’s
figures of the “penis avec prostate de deux especes d’Aglaja” (1954: 50,
Fig. 8, g) are actually Bergh’s figures (1894: PI. 10, Figs. 13 and 11) of
Navanax inermis (Cooper, 1862) and N. aenigmaticus (Bergh, 1894);
Pilsbry (1895-1896: PI. 15, Figs. 89 and 86) had copied them with the
correct names.
With A. pusa we compare those species of Aglaja whose range is geo­
graphically related to the coast of Florida today or was in the past.
19
West Indian species:
A. gemmata (Morch, 1863: 25). Yellow or dull fleshy with close longi­
tudinal black lines. Shell not described. Unfigured. Possibly identical with
the following species.
A. punctilucens (Bergh, 1893: 131). Outer surface of parapodia and sole
with alternate yellowish and brown longitudinal lines. Head shield with
brilliant emerald-green dots, preserved in alcohol for 30 years. Shell pro­
portionally smaller than that of pusa.
Species from the Atlantic Ocean and the Mediterranean Sea:
A. depicta Renier, 1807 (Bergh, 1893: 123). Cephalic shield and para­
podia edged with an inner orange and an outer sky-blue stripe. Head
shield with two or four stripes of buff in front. Shell similar to that of
pusa in size and shape, but without spines. Penis tubular, not coiled;
generally paired prostates with common duct.
A. tricolorata Renier, 1807 (Bergh, 1893: 111). Edges of head shield and
parapodia frequently as in preceding species; sole velvety black (also in
depicta). Cephalic shield less than half body length. Left posterior mantle
lobe generally with flagelliform filament. Diameter of shell 6.5 mm. Male
organ as in depicta.
A. orbignyana (Rochebrune, 1881; Pilsbry, 1896: 239). Black violet with
yellowish longitudinal lines.
A. maderensis (Watson, 1897: 238). Total length 12 mm; in life, opaque
white with minute specks of crimson.
A. capensis (Bergh, 1907: 29). Greyish white with black marks, possibly
(material preserved in formalin) remains of original color. Shell does not
seem to project into right posterior mantle lobe. Male organ black in front.
A. seurati (Vayssiere, 1926: 125). Total length up to 10 mm; wax-white;
diameter of shell 3 mm.
A. dubia O’Donoghue, 1929: 11. Color as in capensis, but without black
marks. Head shield Vs of back. Shell very little produced on right side.
Left lobe with flagellum.
A. minuta Pruvot-Fol, 1953: 30. Length 5.2 mm preserved. Color as in
tricolorata, but sole brown.
American Pacific species:
A. purpurea (Bergh, 1893: 133; 1894: 209). Blackish purple alive. Right
posterior wing of shell “not long.” Male atrium black.
A. diomedea (Bergh, 1893: 133; 1894: 211). Diameter of shell 5 mm,
its left spur unusually large. Prostate paired (Marcus, 1961a: 6-7; in
PI. 1, Fig. 12, read 67 for 91).
A. ocelligera (Bergh, 1893: 133; 1894: 212). Left posterior lobe with
flagellum. Diameter of shell 4 mm; inner border broadly thickened,
especially on right, rather short wing. End of prostate bifurcate.
A. adellae (Dali, 1894: 73). Color plum with vermiculate golden yellow
spots. End of right wing of shell blunt.
20
A. bakeri MacFarland, 1924: 391. Back with irregular longitudinal lines,
light on dark ground. Head shield V3 of total length. Diameter of shell
4.7 mm; right wing long, narrow, and pointed.
A. nana Steinberg & Jones (1960: 73). Preserved 9.8 mm long. Translu­
cent greyish white with irregular black flecks, small yellow brown dots and
larger opaque white spots. Shell without projecting process. Male organ
not described.

6. Chelidonura hirundinina (Quoy & Gaimard, 1833)

Reference.— Marcus, 1963: 6.
Material.— Florida: Soldier Key, Biscayne Bay; on Halimeda. Sandra
Maxwell, April 12, 1964. One specimen.
Further distribution.—Curasao, from Thalassia; Indo-West Pacific, from
Mozambique to Japan, New Caledonia, New South Wales.
This second Atlantic specimen is 12 mm long in preservation, hence
adult. The length of the brown shell is 4 mm. The median blue line is
incomplete, wanting in the posterior part of the head shield; the lateral
blue lines are interrupted. The opaque white crescent on the mantle is
large and of rather irregular outline in front.

Genus Haminoea Turton & Kingston, 1830

Like many—but not all (e.g., Dolabrifera)— herbivorous groups, the
genus Haminoea comprises a great number of species, contrasting with the
carnivores which are generally of low speciation, e.g., Tylodina and
Umbraculum. The shape of the shell is certainly important for classifica­
tion, but young snails are always relatively broader than adult ones. It is
true that globose or ovate or cylindrical shells occur in anatomically dif­
ferent species. Therefore the shape of the shell can be used for determina­
tion only if species of a restricted area (e.g., the west coast of Florida)
are considered. Material from the Indo-West Pacific reef fauna, with several
species of notoriously wide range, cannot be determined by the shape of
the shell alone. The color of the shell and the periostracum are little
important and of no use for the identification of shells thrown onto the
beach. Of the conchological details, distinct or weak or absent spiral striae
are good characters. The extension of a callus and the insertion of the outer
lip of the aperture to the right or to the left of the apex may depend upon
secondary calcification; we found these criteria less secure than we had
expected from Pilsbry’s Manual and descriptions of Hawaiian species.
The color of the body and the extension of the parapodia are taxonomic
characters of a certain value, if living snails are available. The shape of
the cephalic shield and the position of the eyes must be considered as in
some cases they are specific. Often the Hancock’s organ permits a specific
21
separation. Possibly, its structure implies an ecological separation of the
species whose geographic ranges overlap. Reproductive isolation is prob­
ably brought about by the specifically different male copulatory organs,
which reveal themselves to be very useful for classification.
Of the hard parts in the digestive tract, the gizzard plates are rather
uniform, hence specifically of little use; the number of their ridges increases
with age. The labial armature (jaws) is uniform too. The radula, however,
shows specific details, useful for classification, though probably without
adaptive significance.
The central nervous system still needs comparative studies based upon
uniformly handled, relaxed animals. By such important, although admit­
tedly difficult, work the natural groups of species may become recognizable,
and possibly also their greater or lesser reactive abilities, and their
consequent wider or more restricted range.

7. Haminoea antillarum (d’Orbigny, 1841)

Figures 14-19
References.— Pilsbry, 1893-1895: 358; Kobelt, 1896: 104; Bergh, 1901:
228; Perry & Schwengel, 1955: 194; Warmke & Abbott, 1961: 142.
Material.—Florida: Biscayne Bay, Matheson Hammock, near wading
beach, among Diplanthera. R. A. Smith, March 17, 1964. 20 specimens.
Further distribution.— From West Florida to the West Indies (original
locality: Cuba); West Africa, Gabon. On the west coast of Florida beyond
low tide mark, often in tide pools or about the inlets of creeks and bayous.
Name .— We follow Abbott (1955: 279) and Warmke & Abbott in con­
sidering the name guadaloupensis Sowerby (1868, PI. 3, Fig. 14) as an
intraspecific indication of forms whose shells are larger (18 X 14 mm), more
deeply colored, and often broader in front, or as Pilsbry put it, “more
constricted above.” This name had been introduced without reference to
d’Orbigny’s Bulla antillarum (1853: 124, PI. 4, Figs. 9-12). In giving the
year 1841 for d’Orbigny’s first description we follow Sherborn, and
Warmke & Abbott. H. antillarum and H. guadaloupensis have the same
range. Pilsbry thought guadaloupensis to be a variety of antillarum; Kobelt
put them side by side as neighboring species. Bergh used guadaloupensis,
though his shells, 7X 5 mm, and light yellowish white with greenish hue,
suggest typical antillarum. Johnson (1934: 148), and Perry & Schwengel
write H. antillarum guadaloupensis.
Description.— The shell measures 10.5 X 8 mm; it is whitish with a greenish
hue. Reddish brown spiral lines on the mantle of the preserved specimens
produce a pinkish general color. These lines are not visible in living snails,
whose color (from the photograph) is pepper and salt, with a silky
brownish lustre on the shell, and more dark and fewer white spots than in
22
elegans. The eyes and the cerebral ganglia are farther apart from one
another than in elegans, and the light haloes around the eyes are larger.
The cephalic shield is smaller than in elegans and bilobed behind. The

13. Haminoea elegans (Gray); right Hancock’s organ.— F i g u r e s 14-

F ig u r e
19. Haminoea antillarum (d’Orbigny). 14, right Hancock’s organ; 15, gizzard
plate; 16, radular teeth; 17, male copulatory organ; 18, penial papilla; 19, wart
of same.
23
parapodia of living snails reach the hind end of the shell. The Hancock’s
organ has 10 dorsal and 12 ventral leaflets, hence fewer than in elegans,
yet the organ is much longer (Fig. 14).
The radula has 29 rows and 23 teeth per half-row. The rhachidian tooth
is 73 ^ broad, 52 ^ high, its central cusp 40 ^ long, its base smooth
(rough in elegans). The cusps of the lateral teeth increase in length from
the innermost to the 7th lateral tooth, e.g. from 53 to 78 and decrease
in breadth. Farther outwards these cusps shorten. In the three newest rows
there are four weak inner denticles on the cusp of the innermost lateral
tooth. The gizzard plates have about 24 ridges, and their shape differs
according to the view but is actually not different from those of elegans
(Marcus, 1957: Fig. 14). The same applies to the jaws.
The male atrium is 3 mm in length and folded longitudinally. The penis
consists of a papilla and a collar, together 1.1 mm long, with slightly
cuticularized surfaces. Tiny warts on the papilla and collar were not
mentioned by Bergh (1901: 229), but are visible with high power and
good illumination. In Fig. 18 these warts were drawn with some exaggera­
tion. Though Bergh described the male duct as extended throughout the
papilla, his drawing (PL 19, Fig. 1) shows a subterminal opening, as in
the present material. The ental dilation of the male duct is a seminal vesicle
clothed with prostatic epithelium which is especially high around the ectal,
cylindrical part of the vesicle. The inner appendage of the male duct of
elegans is a compact bipartite prostate (Marcus, 1958: PI. 3, Figs. 16,
17, q).
8. Haminoea elegans (Gray, 1825)
Figure 13
References .— Pilsbry, 1893-1895: 355; Marcus, 1957: 395; 1958b: 35;
Warmke & Abbott, 1961: 141.
Material.—Florida: Biscayne Bay, Matheson Hammock, in part near
wading beach; among Diplanthera. R. A. Smith, November 1963 and
March 17, 1964. About 12 specimens.
Further distribution.— Southeast Florida and the West Indies; Brazil, Rio
de Janeiro and coast of Sao Paulo.
Color alive (from photograph and note).— The shell is transparent. The
body is pepper and salt, with numerous white elements; larger individuals
have a dark greenish cast on the parapodia and head shield. The eyes
stand near together, and are surrounded by small light haloes.
Specific characters.— All over the shell there are engraved straight spirals
of grooves. The cephalic shield is long, without or nearly without a posterior
notch. The parapodia leave the hind part of the shell free. The Hancock’s
organ (Fig. 13) is smaller than in the preceding species, but with more
leaflets: 16 dorsal and 20 ventral. The penis has numerous transverse folds
24
each bordered with a row of cuticular pegs. A groove extends along the
ejaculatory section of the male duct. The ental appendage of the duct
is a bipartite prostate.

9. Oxynoe antillarum Morch, 1863

Figure 20
Synonym.— Oxynoe aguayoi Jaume, 1945: 22.
References.— Pilsbry, 1895-1896: 164; Bergh, 1900: 202; Engel, 1927:
111; Marcus, 1963: 16.
Material.— 1. Canal Zone: Fort Sherman near Colon (Caribbean coast),
among Caulerpa growing on rocks in shallow pools near mangroves.
F. M. Bayer and R. W. Bayer, December 23, 1962. One specimen.— 2.
Florida: Biscayne Bay. From Caulerpa growing on sea wall of Crandon
Marina, Key Biscayne. F. M. Bayer, January 1963. One shell.
Further distribution.— Cuba; Puerto Rico; Virgin Islands, St. Thomas
(original locality); Curasao.
Remarks.— Though the color elements, green, white, black, and brown are
repeated in previous descriptions, their distribution varies. The color photo
of the present animal from locality 1 shows a yellowish green ground color.
The head has a few white spots; the rhinophores are flesh-colored with a
pattern of black. The mantle under the transparent shell is dark green with
many light spots. The borders of the transparent parapodia are set with
high white papillae and have some brown spots. The fore end of the foot
is quite light blue. The dorsal side of the tail has a white mid-line with a
pattern of black.
The shells of Oxynoe are variable, as Pilsbry’s (PL 53, Fig. 90) and
Bergh’s (PI. 14, Fig. 16) figures show, both of the original material.
Therefore Jaume’s species, based on shells which have the same measure­
ments as Pilsbry’s, but are a little narrower at the base, cannot be main­
tained. The present shell (Fig. 20) is 7.3 mm long, 4.5 mm broad; its ratio
of breadth to length approaches Bergh’s rather than Pilsbry’s and Jaume’s
shells, and its shape is similar to Pilsbry’s figure. Engel’s shell (Fig. 27, a)
has the ratio shown by Pilsbry’s and Jaume’s figures.

10. Lobiger souverbiei P. Fischer, 1856

Synonyms (after Kay, 1964b: 194).— L. viridis Pease, 1863: 510;
L. picta Pease, 1863: 75 (quoted from Pilsbry, 1895-1896: 169-170);
L. nevilli Pilsbry, 1895-1896: 168.
References.— Engel, 1927: 112; Marcus, 1957a: 398; 1963: 16; Kay,
1964b: 193.
25
 soy-
F ig u r e 20. Oxynoe antillarum Morch, shell.— F i g u r e 21. Elysia evelinae Mar­
cus, animal, from color photo.— F i g u r e s 22-25. Elysia papillosa Verrill. 22,
animal, from color photo; 23, radular tooth; 24, penis; 25, tip of same.—
F i g u r e s 26-27. Elysia cauze scops, ssp. nov. 26, animal, from color photo;
27, radular tooth.

Material.— Florida: Biscayne Bay. From Caulerpa growing on sea wall of

Crandon Marina, Key Biscayne. F. M. Bayer, January 1963. One speci­
men and three shells.
Further distribution.— Tortugas Keys (Thiele, 1910b: 123); Puerto Rico;
Guadeloupe (original locality); Curagao; Brazil, coast of Sao Paulo.
Indo-Pacific Ocean, from Zanzibar and Ceylon to the Sandwich and
Society Islands (Kay, 1964b).
Remarks.— The present shells are 11.5, 8, and 7 mm long; the largest
complete animal is 17 mm in length and 24 mm in breadth, including the
parapodia. The color photo shows a greenish yellow body and a yellowish
green mantle under the transparent shell. On the mantle some fine black
lines, “as drawn with ink” (Engel), run with interruptions and chiefly
parallel to the longitudinal axis of the body. White papillae are numerous
on the head and the tail and abundant on the parapodia where they nearly
conceal the ground color, greenish with brown spots, and give them a
spinous aspect. Two white lateral stripes run along the head, reaching
the bases of the rhinophores, whose rolled tips are white. Also the sides of
the tail and a mid-caudal patch are white. The parapodia and the tail bear
a brown network.
The Mediterranean L. serradifalci (Calcara, 1840) is evidently a differ­
ent, much bigger species. Pilsbry (1895-1896: 167), it is true, wrote
“length slightly exceeding one inch” (25.4 mm), but actually it reaches a
little more than 40 mm (Vayssiere, 1885: PI. 2, Fig. 48). Gonor’s detailed
description of living serradifalci (1961: 386) does not mention the black
lines of the mantle. The identity of L. pilsbryi Schwengel, 1941 (Perry &
Schwengel, 1955: 195-196, PL 55, Fig. 364; 365 is a misprint) from the
west coast of Florida with souverbiei is probable, but not sure. Its color
elements are slightly different from those in souverbiei; its shell, 12.5 mm
in length, is a little longer than the present one (11.5 mm), the hitherto
biggest shell of souverbiei. The Brazilian specimen of souverbiei had the
first radular teeth lying in the ascus crenulated on both sides of the blade
(Marcus, 1957a: Figs. 21, 22). The same crenulation occurs in the
corresponding teeth of the present material, so that no taxonomic separa­
tion is indicated.
11. Elysia evelinae Marcus, 1957
Figure 21
References.— Marcus, 1957a: 410; 1960b: 153.
Material.— Florida: Biscayne Bay. Among algae from sea wall of Crandon
Marina, Key Biscayne. F. M. Bayer, November 29, 1962. Five specimens.
Further distribution.—Biscayne Bay, between Miami and Miami Beach,
MacArthur Causeway; Brazil, coast of Sao Paulo (original locality).
Remarks.— The preserved animals of the present material are about 3 mm
in length. A dark circumanal band and the thickened tip of the radular
tooth characterize the species.

12. Elysia papillosa Verrill, 1901

Figures 22-25
Reference .— Verrill, 1901: 31.
Material.— Florida: Biscayne Bay, Soldier Key, on Halimeda. Sandra
Maxwell, April 12, 1964. One specimen.
27
Further distribution.— Bermudas.
Description.— The length is 9 mm (tail incomplete), the breadth 2.3 mm
with adducted parapodia. The grey ground color and one or two transverse
brown bands on the rhinophores are almost concealed by many white
papillae and skin glands of different size, giving the slug the appearance of
being sprinkled with sugar. The rhinophores are large; the eyes lie at
some distance behind the rhinophoral bases, as shown in VerriH’s figure
(PI. 4, Fig. 3). The mouth is bordered with pigment cells. The pericardial
eminence is globular, the straight renal sac extends backwards twice as
long as this swelling. The anus is located at the point where the furrow that
rises from the pedal groove meets the right border of the pericardial
elevation. The anterior corners of the foot project.
The radula comprises 6 teeth in the upper, 7 in the lower limb. The
newest tooth is 177 jjl, the biggest functioning tooth 0.2 mm long; older
teeth are smaller. The cusp is pointed, the serrulation coarse, the denticles
2-3 fi broad. The ratio of the total length of the biggest cusp/base = 2.2/1.
The penial papilla is thick, muscular, about 0.7 mm long and 0.3 mm in
diameter; it contains the winding male duct which ends with a triangular
stylet, 60 ^ long, 25 ^ broad at its base.
Discussion.— As far as it is possible to identify material of an Elysia with
the description of a species whose radula and other anatomical details are
not known, the present slug corresponds well to papillosa. Verrill wrote
“head large, neck long; rhinophores large; strongly folded and wide at the
tips. Whole surface of body, head, and outside of parapodia thickly covered
with small, conical papillae.” In the present animal the papillae cover also
the rhinophores, obscuring their transverse bands, all the ground color, and
the undulations of the parapodial edges, which Verrill called deep.

13. Elysia cauze scops, subsp. nov.

Figures 26, 27

References to E. cauze cauze.— Marcus 1957a: 405; 1960b: 153; 1963:

22 .
Material.— 1. Florida: Biscayne Bay. Among algae from sea wall of
Crandon Marina. November 29, 1962. One specimen.— 2. Florida: Bis­
cayne Key, in canal near Still’s shrimp dock. R. C. Work, October 1963.
Three animals.
Syntypes.— USNM 576273.
Distribution of E. cauze cauze.— Miami area, on Caulerpa; Antilles, St.
Martin; Bonaire; Curagao; Aruba; Brazil, coast of Sao Paulo (original
locality). Kay (1964b: 195) reported E. aff. cauze from Hawaii.
28
Description.— The present preserved slugs are no more than 8 mm in
length, hence much smaller than cauze cauze (preserved 13-20 mm). The
color photo of the specimen from the first locality corresponds to the colors
of cauze cauze, in part brownish, in part greenish grey; the collector of the
material from locality 2 noted: “color dark greenish; papillate surface.”
Conical and mamillate papillae and small dark rings around the outlets
of the cutaneous glands are as in cauze cauze. The black anterior border
of the parapodia noted in the typical subspecies occurs only in one of the
present four specimens (locality 2), another has the whole border of the
parapodia black (locality 2), and two have no parapodial color marks
(localities 1 and 2).
The radula shows several characters which suggest a subspecific sepa­
ration from cauze cauze. The size (0.16-0.18 mm) of the teeth, smaller
than in cauze cauze (0.25-0.33 mm), coincides with the smaller size of
the body, and seems to indicate that the smallness of the preserved slugs
of cauze scops is not due to a specially strong contraction in the fixative.
The serrulation of the cutting edge on the inferior side of the cusp is
fine, the denticles are about 1 ^ broad, against 2-3 ^ in cauze cauze.
The ascus is spiral, but in cauze cauze it is straight; this feature must be
considered with caution (Marcus, 1963: 22).
Series of sections prove that cauze cauze and cauze scops are quite
near to one another, because the blood gland (Marcus, 1957a: Fig. 44,
b ), the dorsal crop of the oesophagus (Fig. 43, x ), the broad unarmed
penial papilla (Fig. 42, p ), through which the winding male duct runs,
and the female aperture (ibid., q ) located immediately under the anus
(a) are the same in both subspecies.

14. Elysia tuca, spec. nov.

Figures 28-32
Synonym.— Elysia papillosa (non Verrill, 1901) Marcus, 1963: 21.
Material.— Florida: Soldier Key, Biscayne Bay, on Halimeda. Sandra
Maxwell, April 12, 1964. Five specimens.
Syntypes.— USNM 576286.
Further distribution.— Curasao.
Description.— According to the color photograph, the living animal is
dark green, somewhat clouded by lighter, more yellowish, undefined
areas, especially anteriorly between the parapodia, on the sides of the
body, and on the back and head. The rhinophores are comparatively
small; their distal half bears crowded white papillae. The head shows
a thin net of lighter elements between the rhinophores and an oblique
white cross which begins at the bases of the rhinophores.
29
F i g u r e s 28-32. Elysia tuca, sp . n o v . 28, a n im a l, fr o m c o lo r p h o to ; 29, r a d u la r
to o th ; 30, p e n is; 31, tip o f s a m e ; 32, d ia g r a m o f a lim e n ta r y tr a c t.— F i g u r e s
33-37. Elysia duis, sp . n o v . 33, liv in g a n im a l, fr o m c o lo r p h o to ; 34, r a d u la r
to o th ; 35, d o r sa l v ie w o f p h a r y n x ; 36, te e th in a sc u s ; 37, tip o f p e n ia l p a p illa .

On the neck, an approximately triangular white spot contains a few

pale pink elements. The anterior border of the foot is dark; its round
lateral corners project only slightly in the extended animal. The para-
podial edges are straight, not undulate, with no specially colored band,
but with an irregularly broad and interrupted zone of white glands and
papillae. Some pale pink marks lie between the papillae. There are
scattered white papillae on the lateral parts of the parapodia.
The preserved specimens are about 8 mm in length, 4.8 mm in breadth;
the head is rounded, not carinated on the sides. The parapodia leave the
neck and head free and reach the tip of the tail; they have only slightly
30
undulate margins. All papillae have vanished. The green color and white
marks have faded out nearly completely, so that the dark pigment be­
comes visible on the tip of the tail and in the middle of the rhinophores.
The radula has 10 teeth in the upper limb, 9 in the lower, and a
number of teeth heaped up in the ascus. The teeth are small, 90-105 /x
in total length, the base of a 1 0 5 tooth measured 47 p. The serrulation
of the under edge is fine; there are 12 denticles on 13 ^ of length. The
buccal bulb is 0.6 mm long.
A blood gland is absent; the renal pore lies on the right side of the
reno-pericardial swelling, behind its middle. The wall of the oesophagus
is muscular in the posterior section, but without a crop. The penis is
very long (Fig. 30), in one specimen 1.3 mm and in another 1.9 mm in
length, with a diameter of 0.2-0.3 mm according to the level. Its tip is
narrowed, about 0.4 mm long, 80 ^ in diameter. The outlet of the male
duct contains a cuticular structure consisting of about three needles 52 fi
long which are pinnate on their inner end (Fig. 31). The male opening
lies at the base of the right rhinophore. The anus opens into the furrow
that continues the anterior pedal fold upwards. The female aperture lies
immediately below the anus.
Remarks.— The radular teeth and the penis show that our former papillosa
from Curagao is the same species as the present one. Both differ from
papillosa Verrill (see Figs. 23-25) by the absence of transverse color
marks on the smaller rhinophores, and in the nearly straight borders of
the parapodia, which contrast with Verrill’s description.

15. Elysia duis, spec. nov.

Figures 33-37
Material.— 1. Florida: Bear Cut, Biscayne Bay. Warren Burgess. January
1963. One specimen, only color photo seen.— 2. Florida: Key Biscayne;
rocks at north end of island at Bear Cut. Sandra Maxwell, May 2, 1964.
One specimen.
Holotype.— USNM 576284.
Description.— The living slugs, on the photos, have a light green ground
color, especially light on the head and neck. Black are the mouth region,
the tips of the rhinophores, and a ring that separates the outer third of
the rhinophores from the base. This outer third is red. The anterior border
of the foot is red, followed by black and blue bands. The same colors in
the same sequence appear on the parapodial margins. The red band con­
tains some white spots and is irregularly broad, wider where the para­
podial folds are broader. The black band is prolonged downwards; a
brilliant blue one fades out into the green ground color. The underside
is dark greyish green with white spots.
31
 The preserved slug is greyish green. It is 10 mm long, 7 mm broad with
abducted parapodia. The latter leave head and neck free and reach the
tip of the tail; their borders are undulate and have about six larger waves.
The rhinophores are strong. The corners of the foot are prominent; its
anterior border is grooved. The surface of the body and parapodia is
smooth.
The radula comprises eight teeth in the upper, nine in the lower limb,
and 14 in the spiral ascus. The length of the tooth is 52 its base 29
the under edge of the cusp bears 1 ^ broad denticles. The epithelium of
the buccal bulb (pharynx) contains two patches of brown pigment,
forming a broad V in dorsal view (Fig. 35). In the specimen examined,
the tip of the penis is everted, a short point on an egg-shaped papilla.
Remarks .— The only species of Elysia from the western Atlantic warm
waters with similar bright colors is the Bermudian E. picta Verrill (1901:
30). Its head and back are reddish brown, the neck bears a yellow cross;
a purplish spot lies between the parapodia in front. These are edged with
a red border and a band of bright blue below it. A colored anterior region
of the foot is rare in Elysia but occurs, e.g., in E. gracilis Risbec (1928:
278), a very slender form with reddish sides of the body, and in E.
splendens Baba (1949: 36, 132). The latter has a complicated color
pattern consisting of 3-4 yellow lines on the outside near the parapodial
margins and a vermilion tip of the tail; it has rather prominent denticles
of the radular cusp. E. livida Baba (1955: 12, 43) must be compared
with duis. It is recorded from the western and eastern coast of Middle
Japan, but in view of the gap in our knowledge of elysias from tropical
Pacific America its possible further distribution is not known. E. livida
has a deep blue ground color with no special pattern on the rhinophores,
and the three colored bands on the edges of the parapodia are black,
orange, black.
Elysia spec.
Material.— Canal Zone: Ft. Sherman (Caribbean coast). Among Caulerpa
growing in rocky pool near mangrove. F. M. Bayer and R. W. Bayer,
December 23, 1962. One specimen.
Description.— The length is 6 mm (tail wanting), the breadth 8 mm; the
surface is minutely papillose. The collectors noted a pale yellow-green color
alive; the preserved slug is yellow and has dark glands with dark ringlets
around them. Glands occur also on the reno-pericardial eminence and on
the middle of the back, where such are absent, e.g., in cauze. The rhino­
phores are slightly blackish; there are pigmented cells on the labial borders,
no color marks on the edges of the parapodia. The anterior border of
the foot is grooved; the sole has a transverse furrow.
The buccal mass is large, nearly 2 mm in length. The size of the radular
32
teeth in the spiral ascus is from 17 to 90 increasing to 240 ^ in the
functioning teeth; the youngest one is smaller, 210 /x. There are eight
teeth in the upper, 17 in the lower limb, and about 15 in the ascus. The
base of the 240 /x-tooth measures 80 the denticles of the cusp are fine.
Discussion.— An immature specimen of the genus Elysia can be classified
specifically only if it has striking color marks, or belongs to a restricted,
well-inventoried fauna. E. flava Verrill (1901: 30) from the Bermudas
resembles the present slug by its very minute papillae and the light yellow
head, neck, rhinophores, back, and foot. But the color of the outer side of
its parapodia is olive-green, speckled with white, and their inner surface
is almost black, due to the diverticula of the intestinal gland.

16. Tridachia crispata Morch, 1863

Plate 1, Figs. 7, 8; Figure 38
Reference.— Marcus, 1963: 23.
Material.— 1. Florida: Biscayne Bay, Matheson Hammock, near wading
beach, among Diplanthera. R. A. Smith, March 17, 1964.— 2. Florida:
Elliott Key. Margot Fish Shoal (reef) off middle of island; on “artificial
reef.” R. C. Work, November 2, 1962.— 3. Florida: Long Reef, off Elliott
Key; under rocks, depth 3 m. Warren Burgess, July 11, 1964.— 4. Florida:
Key Largo; Garden Cove. Sheldon Dobkin, March 21, 1964.— 5. Florida:
Long Key Bight. E. Iversen, D. Tabb and A. C. Jones, October 10, 1960.—
6. Florida: Grassy Key, Florida Bay side, just below Tom’s Harbor, Via­
duct No. 2; bottom, calcareous fill with Thalassia and scattered algae;
depth, less than 5 ft.; water temp. 28° C. Warren Burgess, October 12,
1962.— 7. Haiti: St. Marc Bay, 1/4 mile north of St. Marc; coral reef
with Thalassia; depth 3-9 m. J. Randall et al., December 22, 1959 (CRR-
C ar-2).— 8. Los Roques Islands (E. of Bonaire): Dos Mosquises; south
end of southern island; rocky bottom with Acropora, depth 4-5 ft. R. C.
Work, August 2, 1963 (D. de Sylva No. 450).— 9. Jamaica: northeast
coast, at Wilkes Cove about 4 mi. E of Port Antonio, 18° 10' 00" N,
76° 22' 40" W. D. de Sylva, Sept. 26, 1963 (D. de Sylva No. 485). (A
total of 18 specimens.)
Further distribution.— West coast of Florida; Dry Tortugas; Miami area;
Bermuda; Antilles, St. Croix (original locality) and other islands; To­
bago; Bonaire; Curasao; Aruba; coast of Venezuela, and British Honduras.
Remarks.— The animal from locality 6 was about 10 cm long when fully
extended. The specimen from locality 3 is blue instead of the usual green.
The parapodia are edged with a fine white line as in the other specimens,
but the next zone is pale blue instead of colorless. Parallel to the edge
follows a row of yellow dashes which are light green in the other slugs.
33
The adjacent broad band is pinkish lavender in this and pink in the other
animals. The lower part of the parapodia is bright blue with a few light
blotches instead of dark green with many white spots. The sole is light
blue instead of white or yellowish. One of the two specimens from
locality 4 has the parapodia less folded than usually; its digestive diverticula
seem to be empty.
Bermuda was included in the survey of occurrences, because we think
that Verriirs slug (1900: 547, PL 66, Fig. 4) called “Elysia crispa Morch”
is probably T. crispata Morch (1863: 40), though the parapodia were
not drawn united in front. Thus the names Elysia (Elysiopterus) verrilli
Pruvot-Fol (1946: 39) and E. pruvotfolae Marcus (1957a: 415) should
be discarded.
17. Tridachiella diomedea (Bergh, 1894)
Figures 40-47
References.— Bergh, 1894: 194; MacFarland, 1924: 406; Steinbeck &
Ricketts, 1941: 544.
Material.— 1. Canal Zone (Pacific side): Deale Beach (Ft. Kobbe Beach).
F. M. Bayer and R. W. Bayer, December 1962.— 2. Panama: Venado
Island, off Ft. Kobbe. F. M. Bayer and R. H. Chesher, December 30,
1963. (A total of three specimens.)
Further distribution.— Gulf of California, from tidal zone to 18 m; Bay
of Panama.
Remarks.— Thanks to Dr. Bayer’s photographs the color of the living
slug, which deserves Steinbeck & Ricketts’ qualification as “one of the
most spectacular and brilliantly colored small animals captured on the
trip,” can now be described for the first time. The ground color is olive
green, dark on the sides, light on the back. Brilliant blue dashes and
streaks occur on the sides and back. The inner side of the rhinophores is
black, the outer side black with white longitudinal lines. The frilled
margin of the parapodia has a coral red edge and below it a row of
triangular black spots alternating on the outer and inner sides corre­
sponding to the frills (Fig. 40). The spots are interspaced with pink.
Black and pink marks reach into a blue zone which fades out into the
olive ground color. In the preserved specimens the black epithelial pigment
is retained on the rhinophores, on the border of the parapodia and, in one
animal, on the sole, where it becomes more intense backwards.
Prof. Bayer who observed living specimens from Panama in his aquari­
um in the Institute of Marine Sciences supplied the following color notes:
rhinophores deep olive green, nearly black, with pale stripes. The under­
side of the head and the anterior fourth of the sole are pale pea green.
Behind a sharp boundary, which corresponds to the shallow transverse
groove of MacFarland’s description (1924: 407), the color is darker pea
34
green with paler discontinuous stripes. A median darker band runs along
the sole. The groove between head and sole lodging the mouth opens and
closes, as the animal crawls on the glass, evidently testing the surface.
The largest specimen is 35 mm long. The alimentary system is essentially
the same as in Tridachia crispata, with a rich net of hepatic ramifications.
These reach the peripheral regions of the parapodia (Fig. 41), in contrast
with crispata (Yonge & Nicholas, 1940: 294). Also the radular teeth are
different, as the lower borders of their cusps are smooth in diomedea .
The ridges which communicate with the pericardial swelling are branchial
veins and do not “contain ramifications of the liver tubules” (MacFarland,
1924: 408). Symbiotic algae, as known of crispata, occur in diomedea
too, but do not lie in the conjunctive tissue as a band near the margin of
the parapodia. In diomedea they are located in the cells of the hepatic
diverticula (Fig. 41), also in their central parts, e.g., on the underside of
the pericardium. The heart is situated in front of the kidney which is a
U-shaped sac, whose anterior concavity is occupied by the heart. The
renal pore opens on the right side of the middle of the reno-pericardial
elevation. The anus is located as in crispata (Marcus, 1960b: 154),
under the right parapodium, in the fold that continues the anterior pedal
fold towards the dorsum.
Reproductive organs.— The reproductive organs were studied in a single
series of transverse sections stained with haematoxylin-eosin. Evidently
the follicles of the ovotestis are separate, male (mi) and female (ao),
as in Elysia maoria (Reid, 1964: 368). The peripheral follicles are in
an initial male phase, but already contain a little ripe sperm. The contents
of the central follicles are purplish amorphous masses, possibly remains
of ova. The epithelium of these follicles does not show any ovogonia but
consists of high cells with acidophilous granules. The ciliated ducts leaving
the follicles have slender nuclei and a thin coat of muscles; they unite to
form a single hermaphrodite duct (Fig. 46) which has a folded widening,
the ampulla (Fig. 42, a) behind and to the left of the mucus gland (mx).
At the outlet of the ampulla male and female ducts separate. The latter,
the inner oviduct, receives the albumen glands (an) and enters the mucus
gland.
Into the male duct (d) open the thin-walled prostatic ducts. The prostatic
glands (q) are big, club-shaped cells staining light blue (Fig. 43). They
cluster loosely around very narrow ducts which form a ramified layer.
This lies nearest to the skin, chiefly on the dorsal side of the parapodia,
in part also ventrally. The collecting prostatic ducts, whose wide ends
open into the male duct, have no more gland cells.
Ectally to the entrance of the prostate the male duct (d) becomes mus­
cular, curves around the mucus gland (mx), and pierces the big muscular
penial papilla (p). This lies tucked into the male atrium with its tip
35
F i g u r e 38. Tridachia crispata Morch, head.— F i g u r e 39. Cyerce antillensis
Engel, dorsal aspect of a preserved ceras.— F i g u r e s 40-47. Tridachiella dio-
medea (Bergh). 40, head; 41, section of parapodium; 42, diagram of repro­
ductive organs; 43, transverse section of prostatic gland; 44, transverse section
of albumen gland; 45, transverse section of albumen duct; 46, transverse
section of hermaphrodite duct; 47, section of albumen gland passing to duct.
36
directed backward. The ejaculatory duct in the papilla forms a slight
widening. The male opening (ma) lies in a fold at the base of the rhino-
phore, near the right eye.
The richly branched albumen glands (an) stain dark blue. Their sub-
epithelial gland cells (Fig. 44) are smaller than those of the prostate and
more densely packed. The ducts are wider. Where they come near to the
middle (Fig. 42, 45, 47), the granular contents of the gland cells stain
bright red.
A little outwards from the point where the inner oviduct passes into
the mucus gland (m x ), the latter is connected with a long and narrow
duct, the bursa canal, running to the left. The canal dilates into a vesicle,
the bursa copulatrix or spermatheca (b ), which lies on the left side of
the body, laterally to the oesophageal pouch. The mucus gland opens into
a ciliated, non glandular outer oviduct (ai). This duct accompanies the
rectum (/) ventrally and opens (wa) a little below the anus (ar) behind
the male aperture.
The genital organs of Tridachiella diomedea are sufficiently different
from those of Tridachia crispata (Marcus, 1960b: Fig. 43) for a generic
separation, together with the fact that the parapodia are not united.
Bergh’s short description of the reproductive organs of diomedea (1894:
195) is compatible with ours, but his figures 5 and 6 show two joined
seminal receptacles, a spermatheca and a spermatocyst, without any con­
nection with the other organs. We suppose that one of these vesicles is
actually the ampulla. This interpretation is rather forced, but the ampulla
is the only globular organ that might be drawn connected with the sperm-
storing bursa.
18. Cyerce antillensis Engel, 1927
Figure 39
References.— Engel, 1927: 117; Marcus, 1963: 17.
Material.— Florida: 1. Virginia Key, in Duck Lake, on algae. F. M. Bayer,
January 30, 1963. One specimen.— 2. Elliott Key, Margot Fish Shoal
(reef) about 2 miles offshore at middle of island. R. C. Work, June 17,
1963. One specimen.
Further distribution.— Curagao; Tobago (original localities).
Remarks.— The small specimen (5 mm) from locality 1 was greenish when
collected, faded to yellowish tan with small brown spots (Dr. Bayer’s note).
The color photo shows it diaphanous, powdered with opaque white. The
head, tentacles, and rhinophores have a little brown pigment, leaving two
areas over the big eyes free. The tan viscera shine through, and some
brown spots lie in the cerata. The projected penis is colorless.
The big animal (about 35 mm) from locality 2 has cerata up to 11
mm long. The color photo shows a wine-red head and rhinophores, white
37
tentacles, and colorless longish spaces over the small eyes. The red color
of the head extends to the opaque white reno-pericardial eminence, in
front of which the light orange anal papilla stands out. The rest of the
back is black, the cerata are diaphanous with a broad red band along
the mid-line, and an opaque white border, due to glands. Back and cerata
are strewn with orange papillae.
As in Cyerce jheringi Pelseneer, 1892 (1894: 51), now C. cristallina
(Trinchese, 1881), the intestinal gland does not enter the cerata (Engel,
1927: 117). These contain two ventral vessels and one dorsal branch which
runs towards the heart. The penial stylet is straight (Marcus, 1963:
Fig. 24).
Remarks.— During the long editorial delay in publishing this paper, we
have received from Florida and Curasao three specimens of another Cyerce.
These represent the first Western Atlantic records of the Mediterranean
Cyerce cristallina (Trinchese, 1881). The illustrated specimen (Plate 1,
Fig. 5) came from Florida: Margot Fish Shoal, off Elliott Key; 5 m, coral
reef. R. C. Work, April 20, 1967. The other records are: Cape Florida
Flats, Biscayne Bay. Chris Dermott, June 25, 1967.— Curasao, Piscadera
Bay. Diva Diniz Correa, January 28, 1966.

19. Aplysia (Varria) dactylomela Rang, 1828

References.— Engel, 1936a: 5; Eales, 1960a: 307; Marcus, 1963: 10.
Material.— Los Roques Islands (E of Bonaire): Isla Larga, west end of
island, north side; reef and rock, 1-5 ft. R. C. Work, August 8, 1963.
(Work No. 6). One middle-sized specimen.
Further distribution.— Worldwide in warm seas, with exception of the
Mediterranean Sea. Panama is the only record from the American Pacific
coast.

20. Dolabrifera dolabrifera (Rang, 1828)

References.— Engel, 1936a: 29; Kay, 1964a: 184.
Material.— 1. Bahamas: Rabbit Keys just off South Bimini; rocky undercut
shore of coralline limestone. C. R. Robins, March 17, 1957 (BWI-8-9).
Two specimens.— 2. Florida: Palm Beach Shores, north inlet of Lake
Worth; rock jetty, Singer Island. Thomas Fraser and Robert Gore, July 6,
1965. One specimen.— 3. Florida: Biscayne Key, Bear Cut. Marine
biology class, October 1961. Two specimens.— 4. Florida: Plantation
Key, east side. Nancy Voss, February 28, 1960. Two specimens.— 5. East
of Curasao. Los Roques Islands (11° 50' N, 66° 40' W ): Dos Mosquises,
38
north side of southern island; rocky, 1-4 ft. Robert C. Work, August 5,
1963. One specimen, mottled grey-green in life.
Further distribution .— Circum-tropical and circum-subtropical, but not yet
recorded from the American Pacific coast.
Remarks.— The species is common in the West Indies (Engel, 1927:
93-94; 1936a: 32-35); locally, e.g. at Curasao, it is more frequent than
species of the Aplysiinae and Notarchinae (Marcus, 1963: 11). In
western Atlantic warm waters it ranges from the Bermudas (32° 19' N,
64° 50' W) to Cabo Frio (22° 57' S, 42° 01' W ).
Professor Bayer informed us that the living animal from locality 2 was
108 mm long, 48 mm wide. These measurements are among the largest
recorded for this species. Preserved the compact, greenish specimen meas­
ures 60 mm in length, 47 mm in width, and 30 mm in height. The notum
is knobby. The shell is covered by a brown periostracum; it is heavily
calcified and its inner side irregularly callused.

21. Petalifera ramosa Baba, 1959

Figure 48
Reference.— Marcus, 1962b: 45.
Material.— Florida, Biscayne Bay, at Marker 28, depth 8-10 ft.; mud with
patches of algae. M. Roessler and R. C. Work, May 16, 1963. Water
temp. 25.5° C; salinity 33.2 ppt. Swims actively by flexing body dorso-
ventrally. One specimen.
Further distribution.— Southern Japan, Amakusa Bay, from low tide mark
to a few meters in depth; Middle Japan, Sagami Bay; in both these original
localities on Zoster a marina. Florida, Monroe County, 2 miles SSW of
Alligator Reef, 40-50 m.
Remarks.— The animal shown in the color photo is 30 mm in length,
diaphanous, with yellowish green viscera. There is a mid-dorsal white
streak, and a delicate pattern of white spots and lines is concentrated on
the slightly ragged tentacles and rhinophores. Most of the fringed papillae
are retracted and wart-like in the photographed living animal. The warts
have a white center and a white ring around the base.
When crawling the animal is longish, the sides of its body are parallel,
the head is broad. The dorsal slit extends from 15 to 19 mm behind the
anterior end, hence lies behind the middle. The shell (Fig. 48) is more
elongate than that of the Japanese specimens (Baba, 1959: Fig. 1 B; Eales,
1960b: Fig. 2) and widely different from that of Petalifera bonairei Engel
(1936a: Fig. 22b). The two known Floridian animals are considerably
smaller than those from Japan which measured 50-70 mm alive, 40 mm
preserved.
39
 22. Phyllaplysia engeli Marcus, 1955
Figures 49, 50
References.— Marcus, 1955: 105; 1957b: 53.
Material.— Florida: Biscayne Bay. R. A. Smith, March 23, 1964. On
Thalassia. Seven specimens.
Further distribution.— Curasao, Piscadera Bay, from Thalassia in the
mangrove region. Brazil, coast of Pernambuco, Rio de Janeiro, and Sao
Paulo (original locality), especially on Padina.
Remarks.— The color photos show the living small animals bright green
with light longitudinal lines; one specimen has more, the other has less
numerous white specks. On the rhinophores the specks coalesce, forming
a longitudinal streak or three transverse bands. There are further concen­
trations of the white elements on the tentacles and a longitudinal streak
to the left of the dorsal slit. Some cutaneous villi and blue spots are
recognizable.
The anterior border of the foot is slightly grooved, as mentioned in our
second description. The cusps of the mandibular cones are less regular than
in typical engeli, somewhat oblique and nodular; the radular formula is
typical, 30 X 19.1.19, but the 2nd lateral (Fig. 20) has a small innermost
and outermost denticle, hence is quadricuspid. The same condition con­
tinues to the 14th or 15th tooth. The three or four following teeth are
tricuspid due to the absence of an outermost denticle. The outermost
tooth has no denticles. In all teeth the denticles can be split, especially
the big inner one. This irregularity, not drawn in Fig. 50, appears here and
there, not in a whole longitudinal row as similar ones do in other species.
The gut in a clarified whole mount is less looped than in the previously
drawn snail (Marcus, 1957b: Fig. 12).
The collar of the penis bears 2 spines, the base of the papilla 4, its
ectal part 3 (Fig. 49). The latter had no spines in the typical material,
but the number of penial spines was stated as variable (1957b: 62).
Ph. engeli is near to Ph. lafontii (P. Fischer, 1870), redescribed by
Engel (1936b: 200-202), and called Ph. depressa (Cantraine, 1835) by
Pruvot-Fol (1954: 106). The radulae of the Floridian engeli and that from
Curasao (Marcus, 1963: 11) agree; both have more complicated cusps
than the Brazilian specimens, hence are still more different from lafontii
than the latter. The specimens from Florida and Brazil agree with regard
to the size; those from Curasao were up to twice their length.

23. Stylocheilus citrinus (Rang, 1828)

References.— Rang, 1828: 71; Marcus, 1962a: 16.
Material.— Brazil: Coast of Sao Paulo, Sao Sebastiao (23° 50' S, 45° 20'
40
W ), on floating lumps of pitch. A. S. Ferreira Ditadi, September 19-21,
1964. Many specimens.
Further distribution.— North and South Atlantic; East Indies; Western
Pacific Ocean, on floating algae and wood.

F i g u r e 48. Petalifera ramosa Baba, inner side of shell.— F i g u r e s 49, 50.

Phyllaplysia engeli Marcus. 49, penis; 50, radular teeth.— F i g u r e 51. Umbracu-
lum plicatulum (v. Martens), inner side of shell.— F i g u r e 52. Berthella tupala
Marcus, radular teeth.— F i g u r e 53. Berthellina quadridens (Morch), living
animal, from color photo.

Remarks.— The absence of dark striations on the lighter ground color and
of “eye spots” (ocelli) of different color (Engel, 1936a: 59) distinguish
St. citrinus from St. longicauda (Quoy & Gaimard, 1824). Most of the
present living slugs were translucent yellow, as were the first slugs from the
same locality and Rang’s material. Some specimens, however, were more
reddish, approximately sienna. The radula is highly variable. Fine denticles
occur on the outermost lateral teeth in citrinus and sometimes in longi-
41
cauda; in citrinus they number up to eight. The length of the gut, which
we had found very much longer in longicauda from the Lesser Antilles
(Marcus, 1963: 14-15) than in citrinus, is not much longer in material
from Eniwetok, Marshall Islands. The penial spines of citrinus are smaller
and more numerous than those of longicauda, but this difference is of
degree, and, moreover, the armature of the penis becomes stronger with
age. As in our first material as well as in the present one, the secretion of
the mantle gland was purplish, not yellow as in Rang’s specimens. The
color of this secretion is not a constant specific character in Stylocheilus
(Marcus, 1962a: 18). The present animals have black contents of the gut
due to particles of pitch ingested together with the rasped algae.
One animal of the collection from Florida (Virginia Key, beach facing
Bear Cut, Biscayne Bay, J. Ewald, May 11, 1960) is enigmatic. It shows
remains of yellow color without striae or ocelli as citrinus; its seven outer­
most radular teeth are smooth (hitherto only seen in longicauda), the gut
is short (citrinus), the penial spines are large and few (longicauda). We
mention this specimen in order to stress the difficulties in classifying a
deficient, possibly faded, example of Stylocheilus, but do not intend to
unite citrinus with longicauda.

24. Umbraculum plicatulum (v. Martens, 1881)

Figure 51
References.—v. Martens, 1881: 104; Pilsbry, 1896: 178; Sarasua, 1945:
26; White, 1952: 106; Marcus, 1967a: 614.
Material.— West coast of Florida, SW of Sarasota, 106 m. J. Moore,
Summer 1960. One specimen.
Further distribution.— South of Dry Tortugas, 73 m; Florida (Maxwell
Smith, according to Sarasua, 1945); Cuba, Matanzas (original locality),
and La Habana; Caribbean coast of Colombia (Marcus, 1967a).
Remarks.— The preserved animal is 95 mm long, 83 mm broad, and about
50 mm high, hence very big. The back is dark reddish brown to purple with
small blotches of black which are not correlated with the large cutaneous
warts. The shell measures 66X51 mm; it is white with an oblique apex
slanting backwards, 38 mm from the front, 28 mm from the rear, 34 mm
from the right and 17 mm from the left side, hence excentric. The internal
scar of the columellar muscle is complete (Fig. 51), as is characteristic
for the Umbraculidae (Burn, 1959: 28).
In order to preserve this rare animal intact, it was not dissected.
Possibly U. bermudensis (Morch, 1875) from the Bermudas and the
Straits of Florida, mentioned by Pilsbry (1896: 178) and Johnson (1934:
153), is a young stage of plicatulum (Sarasua, 1945: 26). Burn (1959)
gives weighty reasons for reducing to the synonymy of U. umbraculum
42
(Gmelin, 1791) all other species of the circum-tropical and circum-sub-
tropical genus Umbraculum Schumacher, 1817. But as the results of his
anatomical studies have not been published yet, it is preferable to apply
the name of a well-described shell collected not far from the West coast of
Florida.
25. Berthella tupala Marcus, 1957
Figure 52
Reference.— Marcus, 1957a: 416.
Material.— 1. Florida: Key Largo. Under rocks in Largo Sound near mouth
of canal. F. M. Bayer and R. C. Work, November 1962.— 2. Florida:
Biscayne Bay; Southwest Point, Key Biscayne. R. C. Work, January 1963.
(A total of three specimens.)
Further distribution.—Brazil: coast of Sao Paulo (original locality) and
Rio de Janeiro.
Remarks.— The color photograph of the specimen from the first locality
represents a yellowish animal with white spots. Comparison with our
original material shows that the jaw elements (0-5 blunt denticles on
either side of the tip), radula, and shell agree. The innermost lateral tooth
has a small denticle near its base (Fig. 52), not mentioned in the first
description, but present in our original material. The five outermost teeth
decrease in size; on the ten preceding ones there is a small second cusp
as drawn in the original Figure 66. Such cusps are absent in the likewise
yellow B. patagonica (d’Orbigny, 1837: 204). That species is further
distinguished from tupala by the pointed denticles on the border of the
jaw elements, and the rectangular broad foot extending beyond the
notum on all sides. (d’Orbigny, 1846: PI. 14, Fig. 4; reproduced by
Pilsbry, 1896, and Vayssiere, 1898).

26. Berthellina quadridens (Morch, 1863)

Figure 53
Reference.— Marcus, 1962b: 463.
Material.— 1. Haiti: St. Marc Bay, Va- mile north of St. Marc; coral reef
with Thalassia. Depth 3-9 m. John Randall et al., December 22, 1959
(CRR-Car-2). Three specimens.— 2. Los Roques Islands (E of Bonaire):
Isla Larga, west end of island, north side. Reef and rock, 1-5 ft. R. C.
Work, August 9, 1963 (Work No. 6). Three specimens.— 3. Canal Zone
(Pacific side): Perico Island, Ft. Amador. Under rocks at low water.
R. W. Bayer, December 30, 1962. Two specimens.
Further distribution.—Virgin Islands (original locality: St. Thomas), St.
John; Guadeloupe (Deshayes, 1857: 142).
Remarks.— The original indication (Morch, 1863: 29): orange alive,
43
young animals vermilion, is now confirmed by the note “bright orange” for
the animals of the second locality, and by the color photo from the third.
The photo enables us to give the first figure of the entire animal. The dots
in the no turn are epidermal glands; they appear clearly in the preserved
animal and were added to the drawing of the living specimen. The pre­
served animals in the present collection are 13-27 mm in length.
Prof. Dr. F. M. Bayer informed us in a letter of February 13, 1963, that
the animals collected at the third locality continued alive in an aquarium
at the Institute of Marine Sciences in Miami, feeding readily upon sea
anemones.

27. Pleurobranchus (Pleurobranchus) areolatus Morch, 1863

Synonyms.— Pleurobranchus crossei Vayssiere, 1896: 353; 1898: 332;
Pleurobranchus (Susania) atlanticus Abbott, 1949: 73; 1955: 286; Susania
gardineri White, 1952: 106.
References .— Morch, 1863: 29; Marcus, 1962b: 466; 1963: 25.
Material.— 1. Florida: Biscayne Bay, under rocks at east end of County
(MacArthur) Causeway. R. C. Work, March 1962. Two specimens.— 2.
Florida: Biscayne Bay: Tide flats near Southwest Point, Key Biscayne.
Sandra Maxwell, March 25, 1964. One specimen.— 3. Florida: V2 mile
SSW of Alligator Reef Light, Monroe County; at ledge, 4.5-6 m. W. A.
Starck and H. A. Feddern, April 30, 1961 (CRR-F-296). Two speci­
mens.— 4. Los Roques Islands (E of Bonaire): Isla Larga, west end of
island, north side; reef and rock, 1-5 ft. R. C. Work, August 9, 1963
(Work No. 6). One specimen.— 5. Canal Zone (Pacific side): Deale
Beach (Ft. Kobbe Beach); under rocks at low water. F. M. Bayer and
R. W. Bayer, December 30, 1962. Two specimens.
Further distribution.— From the east coast of Florida (vicinity of Miami,
Dade County) throughout the Florida Keys; Virgin Islands (original
locality: St. Thomas); French Antilles; Islote Aves (W of Dominica);
Barbados; Curasao.
Remarks.— P. crossei is evidently the same species. Its type specimen after
preservation for 70 years still showed areolae similar to those of the present
young (24 mm long) individual from the third locality, described as cream
with orange-brown spots. The measurements, the shell with a broad con-
chinous border (Vayssiere, 1898: Fig. 153), the sculpture of the notum,
and the elements of the jaws and radula of crossei agree with areolatus.
The number of leaflets of the ctenidium, 26 in crossei, lies within the range
of variation, 16-27, in areolatus (Marcus, 1962b: 468).
The living animal from the second locality, photographed with scale, is
75 mm long, 55 mm broad, the biggest hitherto recorded specimen. Its
brownish shell measures 2.5 X 3.6 mm. Individuals from the fifth locality are
44
somewhat darker than the others, but the sculpture of the notum, the jaws
and the radula agree with areolatus. Due to preservation in formalin
the calcareous shell was dissolved.
Thanks to the photographs and notes on specimens from most localities,
the indications of color and sculpture can be amplified. The notum is
brownish-orange to brick-red; the rhinophores are reddish brown; the upper
side of the foot is orange, mottled with white. The wart-like tubercles are
round with slightly stellate borders, the lateral warts even smaller than the
small central ones. The center of the warts may be white or dark, or of
the same color as the rest. Chalk-white specks are irregularly strewn on
the warts with variable density. Single warts or groups of them which are
light or dark produce a corresponding blotched pattern.
The number of denticles on the free edge of the jaw elements varies
from none to eight on either side (Bergh, 1897: 112; 1898a: PI. 9, Fig.
34; Abbott, 1949: 75); it may also be different from side to side, and does
not serve for the separation of genera or subgenera in the Pleurobranchinae
(Burn, 1962a: 131).

28. Pleurobranchaea occidentalis Bergh, 1897

Figures 54, 55 A - C
Reference.— Pleurobranchaea Meckelii Leue (Var. occidentalis) Bergh,
1897: 28-30, PL 7, Figs. 16-18.
Material.— 1. Florida: off Miami, 25° 40.5' N, 80°03.5' W, 144 m; grey
mud. R /V G er d a sta. 334. H. Owre, F. M. Bayer, and marine biology
class, July 21, 1964. One specimen.— 2. Northern Straits of Florida, from
26° 13' N (off Pompano Beach) to 26° 21.5' N (off Boca Raton), 80°
00.5'-80° 03.5' W, 110-185 m. G er d a stations 413, 415, 417, 419.
September 22, 1964. A total of 34 specimens.
Further distribution.— S of Cuba, 22° 09' N, 82° 21' W, 324 m; near
Martinique, 870 m , B la k e Expedition 1877-1878.
Description.— The length of the living animal from the first locality was
about 18 mm, the width 9 mm; preserved it is 12 mm long, 11 mm high.
The specimens from the second localities measure, preserved, 13-22 mm
in length. The color of the living slug was lighter than in meckelii; it was
diaphanous with irregular brown rings and white dots, especially on the
snout, tentacles, head, margin of the notum, and back of the foot. The
latter has no spur. The preserved specimens are colorless or of a uniform
pinkish brown.
The foot gland is weakly developed. The smooth edge of the frontal
veil is no specific character (Vayssiere, 1901a: 44). The foot is as broad
as the dorsal part of the body. The mantle is smooth; it leaves the gills
free (as shown in a photo); there is no siphonal mantle fold over its tip.
45
 54, 55A—55C. Pleurobranchaea occidentalis Bergh. 54, right side
F ig u r e s
view of anterior region; 55 A, tips of mandibular elements; 55 B, radular teeth;
55 C, diagram of reproductive organs.— F i g u r e 56. Pleurobranchaea hedgpethi
hamva Marcus; insemination duct and adjacent organs.

46
The gill (Fig. 54) bears 12-15 pairs of pinnate leaflets, about 5 of which
are free from the branchial membrane. The anus (ar) lies over the end of
this membrane, and the prebranchial gland opens (nr) at the root of the
gill.
The elements of the jaw (Fig. 55 A) are high rodlets in the middle of
the mandible, low platelets near the borders. Their maximum height is
seven times their breadth, their free borders have 5-16 uniform denticles,
while Bergh figured 2-6 irregular ones. In the present material the radula
(Fig. 55 B) has 34 rows and 60-70 teeth per half-row; no fold or minute
tooth was seen on the rhachis. The second cusp is absent in the outermost
20 or fewer teeth; this reduction was mentioned also for m eckelii (Vays-
siere, 1885: 132). The connection of the basal region of the teeth with
the radular membrane is distended. In certain positions it appears like a
denticle. Hydrozoa, polychaetes, and crustaceans were found in the gut.
There is no flap on the collar-like membrane that surrounds the genital
apertures. The male atrium (Fig. 54, mo ) is everted in most of the present
specimens. The hermaphrodite duct (Fig. 55 C, ei) widens into a long,
winding ampulla (a) full of sperm. The exit of the ampulla is connected
with the gland mass (g) by a thin strand and is continued into a common
efferent and insemination duct ( d + xo) . Farther outward these ducts
separate. The efferent duct ( d) enters the prostate ( q) and then forms as
in P. capensis (Vayssiere, 1901a: Fig. 232) a long loop, whose ectal limb
is differentiated into a penial sheath (si) and a cuticularized penis papilla
( p ) . The papilla projects into the wrinkled male atrium ( am) .
The vagina (v) leaves the nidamental or outer oviduct (ov) as a wide
tube with folded epithelium and runs straight forwards to the thin-walled
bursa or spermatheca ( b) . The eggs in the spermatheca are evidently a
surplus of female germ cells which are digested in this organ. The insemi­
nation duct ( xo) begins in the ectal third of the vagina, loops inwards and
forms a series of widenings which represent a spermatocyst (y). The
innermost of these widenings is joined with the insemination-efferent duct
by a narrow canal, halfway between prostate and ampulla. Evidently the
alien sperms rising through the insemination-efferent duct reach the eggs in
the glandular oviduct or gland mass. Studies of living material may reveal
whether the inner oviduct, the communication of the hermaphrodite duct
with the gland mass is wider in other phases of the reproductive cycle.
Discussion.— The most important difference between meckelii and occi-
dentalis, which leads us to separate them specifically, resides in the penial
sac. This is voluminous in m eckelii and lodges many circumvolutions of
the efferent duct. The sac of m eckelii is separated from the male atrium by
a closed septum. Less important differences are the lesser size of occi-
dentalis, its less numerous branchial leaflets, and its naked rhachis of the
radula. P. m eckelii has a longitudinal fold (Bergh, 1897: 18) or a median
47
tooth (Vayssiere, 1901a: 35) on the rhachis. The mandibular elements
stressed by Bergh (p. 30) as peculiar in occidentalis agree in the present
material better with the regularly dentate ones of m eckelii (Vayssiere:
PL 3) than with those of Bergh’s specimens of occidentalis (Bergh: PL 7,
Figs. 16, 17).
P. morosa (Bergh, 1892b: 28) from the western Atlantic Ocean has the
gill free for half of its length, hence is a different species.
The caudal spur figures in the diagnoses of the genus Pleurobranchaea
given by Bergh (1897: 4), Vayssiere (1901a: 22), and O’Donoghue
(1929b: 48). This is not correct, as it is absent in tarda, m aculata ,
generally called novaezealandiae (see Burn, 1958: 6), and japonica Thiele
(1925: 283). The spur is inconstant in capensis Vayssiere, 1900 (Bergh,
1907: 31, 32: absent; O’Donoghue, 1929b: 49: present), melanopus
Bergh (1907: 34: absent; O’Donoghue, 1929b: 51-52: present), and
probably in meckelii (Pruvot-Fol, 1954: 215).

29. Pleurobranchaea hedgpethi hamva Marcus, 1957

Figure 56
References. — Abbott, 1952: 1 (hedgpethi); Marcus, 1957c: 21 (hamva);
1961b: 141 (hedgpethi hamva).
M aterial. — 1. Florida: shrimp grounds of Dry Tortugas, 51 m, R. B. Man­
ning, August 4-7, 1958. One specimen.— 2. Off Savannah Beach, 32° 08'
N, 79° 16' W, 70-95 m. One specimen.
Further distribution.— P. hedgpethi hamva: North Carolina, point of Cape
Hatteras; Brazil, coast of Sao Paulo (original locality) and Rio de Janeiro.
P. hedgpethi hedgpethi: Gulf of Mexico from Port Aransas (original local­
ity) to the Bay of Campeche; Surinam (Nijssen-Meyer, 1965: 144).
R em arks. — The flap of the membrane around the genital apertures directed
upwards, not forwards, distinguishes hamva from typical hedgpethi. How­
ever, Nijssen-Meyer (I.e.) describes a slug from Surinam whose flap points
sideways and slightly upwards. This is intermediate between P. h. hedg­
pethi and P. h. hamva, and makes the distinction of the subspecies doubt­
ful. The smooth, minutely serrated or tuberculated frontal veil, the num­
ber of radular teeth, and the shape of the bursa (spermatheca) are not
specific characters.
The preserved slug from locality 2 is 38 mm in length, including the
pharynx, extruded as frequently in preserved animals of this genus. The
radula contains 35 rows and 67-68 teeth per half-row.
Our former figure of the reproductive organs (Marcus, 1957c: Fig. 52)
needs a correction with regard to the inner oviduct. This is a short duct
( o v ) which arises, together with the male duct ( d ), from the ampulla (a)
48
and enters the albumen gland (an), the folded, inner part of the female
gland mass. What we previously called “inner oviduct” is really the
insemination duct (xo). This begins near the entrance of the vagina (v)
into the spermatheca or bursa copulatrix ( b ), but independently from it.
The outer section of the insemination duct is thin and coiled, the middle
part consists of two muscular bulbs, topographically a spermatocyst (y),
and the innermost course coincides with the male duct between ampulla
and prostate ( q ). Obviously the same correction holds for our figure of
P. hedgpethi hedgpethi (Marcus, 1959b: Fig. 6)..

30. Pleurobranchaea agassizii Bergh, 1897

Figures 56A - 56D
Reference.— Bergh, 1897: 48, PI. 7, Fig. 28-32.
Material.— 1. Southern Straits of Florida, 24° 17-18' N, 82° 48-56' W,
350 m, G e r d a station 413. January 25, 1965. One specimen.— 2.
Northern Straits of Florida, off Jupiter, Fla., 27° 00' N, 79° 49' W,
287-262 m. G e r d a station 655. July 16, 1965. Two specimens.
Further distribution .— Region of the Great Bahama Bank, 24° 08' N,
82° 51' W (B lake Expedition, 1877), 620 m.
Description .— The largest preserved slug is 10 cm long, 4 cm wide, hence
four times the length of that of the original description. The buccal mass is
extruded in one specimen, in the others the everted oral cavity forms a kind
of muzzle. The outer part of its blackish cuticle is smooth, the inner one
beset with nodules (Fig. 56 A ). Some black pigment is maintained in the
skin behind the rhinophores and in the folds of the sole.
The veil bears about 20 warts of different sizes in one irregular line.
As in other species of the genus the groove of the rhinophores and that of
the tentacles contain transverse folds which correspond to the Hancock’s
organ. The notum is smooth and laps over the body on both sides, espe­
cially on the branchial side; the tail stands out behind. The spur is colorless
and 2-3 mm long. The anterior border of the foot is thickened, not
distinctly bilabiate. Over its middle opens the deep anterior foot gland.
The posterior pedal gland varies, in the first slug it is weak, 3 mm in
length, in the others well developed, 10-15 mm long.
On either side of the smooth rhachis of the gill there are about 40
pinnules, the hindmost of which are quite small. The branchial membrane
ends at the 7th or 8th pinnule, so that more than half of the gill is free.
The anus lies in the edge of the membrane. The prebranchial gland opens
at the beginning of the branchial rhachis, the kidney farther ventral, and
the genital apertures farther in front. There is no flap over the latter.
The mandibles measure 20 X 12 mm each; their prismatic elements are
up to 0.2 mm long, their free surface is about 50X 20 ^ and bears 2-12
49
denticles of very variable size (Fig. 56 B ). The radula is 22 mm in length
and 16 mm in width. It consists of 32 rows and 98 teeth in the half-row.
In the middle of the half-row the teeth reach 0.7 mm. Up to 17 of the
outer teeth are without secondary cusps; in the younger rows the cutting
edge of the outer teeth is slightly knobbed (Fig. 56 C ), in the older rows
it is smooth.
The reproductive organs (Fig. 56 D) are more similar to occidentalis
than to hedgpethi. The tubular ampulla (a) divides near its outlet. One
branch, the inner oviduct (ov), enters the albumen gland (an), the other,
the efferent duct (d) , courses to the prostate (q). As in the two preceding

F ig u r e s 56A —56D. Pleurobranchaea agassizii Bergh. 56 A, mouth region; 56 B,

mandibular elements; 56 C, outer tooth of young row; 56 D, diagram of
reproductive organs.
50
species the ental section of the male branch, between ampulla and prostate,
functions as efferent duct of its own sperm and insemination duct of alien
sperm ( d + x o ), conveying the latter to the inner oviduct.
Ectally to the prostate the muscular efferent duct runs free and is only
slightly curved before it enters the penis ( p ). Here it begins serpentining
and ends straight. The muscular sheath (si) around the stout penial papilla
is continuous with the wall of the male atrium ( a m) .
The female aperture behind that of the male atrium is the outlet of the
glandular oviduct ( ov) together with the entrance of the vagina (v). The
vagina begins in the anterior wall of the outer oviduct (nidamental duct)
and leads into a bursa copulatrix ( b) or spermatheca. An at first straight,
then looping insemination duct ( xo) leaves the bursa, dilating into a
spermatocyst ( y ) or receptaculum seminis. The ingoing course of the
alien sperm passes through the efferent duct. The ental part of the
glandular oviduct, the albumen gland ( an) , is meandrous, the walls of the
ectal part, the mucus gland (m i), are richly folded.

31. Cadlina rumia Marcus, 1955

References.— Marcus, 1955: 119; 1963: 25.
M aterial.— Florida: Southwest Point, Key Biscayne, Biscayne Bay. R. C.
Work, December 26, 1962. Two specimens.
Further distribution. — Lesser Antilles, St. Martin and Curasao; Brazil,
coast of Sao Paulo (original locality).
Rem arks. — The preserved specimens are 7 and 8 mm in length; they have
nine rhinophoral leaves and five gills; the radular formula is 71X18.1.18.
The first lateral tooth bears 5 outer and 2 inner denticles.
Corresponding data of C. scabriuscula (Bergh, 1890a: 160) found
between Florida and Cuba, 24° 44' N, 83° 26' W, 68 m, are: length 12
mm, rhinophoral leaves about 20, gills 9, radula 108 X 30.1.30, 8-10 outer
and 4-5 inner denticles.
In a color photograph the living animal of the present collection is
transparent white with minute yellow dots on the tips of small warts cover­
ing erect bundles of spicules. The sides and the post-branchial part of the
notum bear a total of 8-12 larger, light yellow, circular spots. The rhino­
phores are yellow, the retracted gills evidently yellowish.
The color of living scabriuscula is not known. According to Bergh’s
notes the preserved specimens have color features similar to those of
rumia , so that the above-mentioned morphological details are necessary
for determination.
32. Chrom odoris neona Marcus, 1955
Figures 57-58A; Plate 1, Figs. 4, 9
References.— Marcus, 1955: 124; 1958a: 419; marcus, 1967a: 614.
51
Material.— 1. Florida: Palm Beach Shores, north inlet of Lake Worth;
rock jetty, Singer Island. Thomas Fraser and Robert Gore, July 6, 1965.
One specimen.— 2. Florida: Biscayne Bay, from rocks along eastern end
of County (MacArthur) Causeway. F. M. Bayer and R. C. Work, May 2,
1962. One specimen.— 3. Same locality. R. C. Work, April 1963. One
specimen.
Further distribution.— Brazil, coast of Sao Paulo (original locality) and
Cabo Frio (22° 57' S, 42° 01' W). Caribbean coast of Panama Canal Zone.
Remarks .—A color photo of the specimen of April 1963 refers to an
animal which died and disintegrated and was not preserved. The photo

57-58A. Chromodoris neona Marcus. 57, specimen from April 1963

F ig u r e s
(Biscayne Bay); 58, specimen from May 1962 (Biscayne Bay); 58 A, specimen
from July 1965 (Lake W orth).— F i g u r e s 59-61. Chromodoris tura, sp. nov.
59, living animal, from color photo; 60, labial hooks; 61, radular teeth.
52
proves that it belongs to Ch. neona, as Professor Bayer had already
presumed. Its pattern (Fig. 57; PI. 1, Fig. 4) differs little from Brazilian
neona; the band on the tail is white, not red, the light blue color is paler,
and the eosin-red elements are darker.
The specimen of May 2, 1962 has the same colors, but its pattern
(Fig. 58) differs farther from Brazilian animals by its red net with light
blue meshes. Rhinophores and gills have purplish bands only. The number
of rhinophoral foliations and gills, and the details of the radula agree
with the original material.
The photo of the slug from locality 1 (PI. 1, Fig. 9) shows a relatively
broad, white notal border interrupted by red lines. The red band on the
tail and the rhinophores purplish all over are as in Brazilian neona.
Otherwise, the pattern (Fig 58A) is similar to that of the animal of May
2, 1962, and we confirm Professor Bayer’s classification. The experience
with Indo-Pacific Chromodoridinae whose numerous color variations, first
separated as species but now either definite or doubtful synonyms,
recommends cautiousness with further species.
The animals at hand (July 1965 and May 1962) are 10-12 mm in
length, 3.5-5 mm in width, and 3.5-4 mm in height. The photos show that
they have shrunken little by preservation. The photo of the slug of April
1963 reveals an animal whose length (23 mm) and width (7 mm) exceed
the measurements of the original material (15-16, 3-4). The aspect of the
two preserved slugs is hardly chromodoridan, because the notum is broadly
expanded, possibly by the loss of turgor, and the foot is concave and
narrow. The tentacles are relatively long and pointed in one slug (July
1965) and so much contracted in the other (May 1962) that they appear
as grooved lobes. The club of the rhinophores bears 17 foliations on either
side in both specimens, and both have nine unipinnate gills.
The radulae of both specimens have more rows, 46-48 (1965), 50
(1962) and teeth in the half-row, 36 (1965), 38 (1962) than were
indicated in the original description, 42 rows, 29 teeth per half-row.
Similar differences are frequent in chromodoridids of one and the same
species. The rhachidian plate of the Floridian slugs is 16 ^ high and has
a 6 fi long cusp. As it is only half as high as the innermost lateral tooth
(34 ^ ) , it is a typical “false” rhachidian tooth, not a true one as in
Felimare bayeri (Fig. 78).

33. Chromodoris nyalya, spec. nov.

Figures 62-66; Plate 1, Fig. 1
Material.— 1. Florida: Elliott Key; Margot Fish Shoal (reef), about 2 miles
offshore at middle of island. R. C. Work, May 12, 1963. One slug.— 2.
Florida: Long Reef, off Elliott Key, under rocks, depth 3 m. Warren
Burgess, July 11, 1964. One animal.
53
H olotype .— USNM 576277.
Description .— The color of the whole animal, from color photos, is a
brilliant blue, with the rhinophores and the axes of the branchiae a little
darker than the other parts. A bright red, uninterrupted band runs around
the notum. The colors have faded in the preserved specimens; the larger
one is 10 mm long, 4 mm high, 3 mm broad. The tentacles are rolled in;
the rhinophores have 11 leaves; there are six (locality 1) and seven
(locality 2) unipinnate gills; the anal papilla completes their circle behind.
The gonopore lies 3.5 mm behind the fore end. The foot is narrow, its
anterior border grooved; the tip of the tail is blackish. The notal border
bears glands (Fig. 63); two antero-lateral groups and a postbranchial
semicircle of seven glands appear sepia in the slug preserved in Bouin
(locality 2), not in that kept in formalin.

photo; 63, preserved animal, showing sub-notal glands; 64, labial hooks; 65,
teeth from middle of half-row, in situ; 66, radular teeth.
54
 The labial cuticle is covered with pointed hooks bearing secondary
cusps; the central hooks are 34 ^ high, the peripheral ones minute. The
formula of the radula is 60X27.0.27. The rhachis is naked. The innermost
lateral tooth is 41 ^ high, its base 24 ^ broad (Fig. 66), the edge has 2-6
inner and 4-5 outer denticles. The height of the second and third teeth
measures 42 ^ the width of the base 16 The cutting edge has one
pointed cusp and four outer denticles, the foremost of which is stronger
than the rest, so that a bicuspid aspect results. The same holds for the
further teeth, whose number of denticles generally decreases outwards.
Some teeth with only one strong cusp and several small denticles must be
defined as unicuspid. The marginal teeth are short, 24 ju,; their upper prong
is longer than the lower one and followed by about 4 weak denticles. The
lower prong is often cleft in any of the teeth.
Discussion. — The combination of blue with red is rare among the Chromo-
doridinae, while blue with yellow or orange is frequent. Therefore we also
compared the species with the latter colors, but did not find the same
pattern in any species with the peculiar radula of the present one. We even
pondered possible allocation to N oum ea Risbec (1928: 165), due to the
broad innermost lateral tooth whose form is similar to that of Noumea
flava Risbec (1928: 169; 1953: Fig. 47A). However, the different width
of the first and second lateral tooth is less pronounced in nyalya than in
Risbec’s species. Pruvot-Fol (1951a: 126, 139) questions the validity of
N oum ea , with exception of the type species. Even if Noumea is maintained
in the sense of Risbec (1953: 85-90) and Baba (1949: 54-55, 144) none
of its species agrees with nyalya, whose lateral teeth are quite peculiar.

34. Chromodoris tura, spec. nov.

Figures 59-61; Plate 1, Fig. 3
M aterial. — Canal Zone (Pacific side): Deale Beach (Ft. Kobbe Beach).
F. M. Bayer and R. W. Bayer, December 1962. One specimen.
H olotype.— USNM 576266.
Description. — Alive, from a photo, the color of the notum is dark violet,
nearly black, surrounded by a broad, bluish white margin. The latter bears
one orange line, the dark center, scattered, fire-red spots and dashes, and
a widely interrupted row of yellowish streaks on its border. The rhino-
phores are bluish black, the branchiae white with black tips. The tail is
dark violet with a brick-red midline.
Preserved the slug is 4 mm long, 2 mm broad, 1.5 mm high. The rhino-
phores have about 10 leaves; the tentacles are broad and rounded; the
anterior border of the narrow sole is grooved. There are seven unipinnate
gills.
The labial armature (Fig. 60) is composed of hooklets with bifid tips
and of different shape and size. The radula (Fig. 61) comprises 42 rows
55
with 31 teeth in each half-row. The rhachis is naked. The innermost lateral
tooth is twice as broad (10 ^ ) as the following tooth (5 ^ ). The edge of
the innermost tooth is denticulated on both sides of the cusp; the four inner
denticles are minute, the four outer ones of different size. That beside the
cusp is nearly as long as the cusp itself but the others are smaller. The
second and following lateral teeth have outer denticles, the foremost of
which is nearly as long as the cusp in the second tooth, so that this appears
bicuspid. Most of the lateral teeth are similar to the sixth, unicuspid with
three or more outer denticles; the latter are smaller in the marginal teeth.
Discussion.— The species has some features in common with the Pana­
manian C hrom odoris agassizii Bergh (1894: 182), that was described upon
one preserved specimen. Most of the lateral teeth of agassizii are bicuspid,
of the H ypselodoris- type. Also the jaw elements are different. It is true
that in species such as nyalya and the present one the length of the first
denticle behind the cusp makes the decision between Chrom odoris and
H ypselodoris difficult. One or two innermost teeth are not sufficient to
define such a species generically. In Chromodoris most teeth are unicuspi-
date, hook-shaped, and serrate on the outer side; in H ypselodoris most are
bicuspidate or bifid. Baba’s species (1949: Figs. 51-55, 56-59) show
both types with clear-cut differences.
The color pattern of Chrom odoris californiensis Bergh (1879a: 112)
is similar to, though not identical with, that of tura, but calif orniensis has
the H ypselodoris- type of the radula.

35 . H ypselodoris edenticulata (White, 1952)

Figures 67-71
Reference. — White, 1952: 113.
M aterial. — 1. Florida: Monroe County, 1 mile SW of Alligator Reef Light,
30-34 m. W. A. Starck II and Henry Feddern, November 26, 1959. Two
specimens.— 2. Florida: Miami Beach. Mosier, 1935. One specimen
(UMML-30-159).— 3. Florida: Lake shore of Palm Beach, on sea wall
just S of Palm Beach inlet, 8-9 ft. R. C. Work. May 12, 1962. One speci­
men.— 4. Florida: Palm Beach Shores, north inlet of Lake Worth. Rock
jetty, Singer Island. Thomas Fraser, Robert Gore. July 6, 1965.
Further distribution. —Dry Tortugas, 18 m.

D escription. — The preserved slug from the third locality is 50 mm long,

23 mm high, 15 mm broad. Its sole is 7 mm broad. The color in life
(collector’s note) is “deep indigo blue, almost black, with chrome yellow
markings, opaque white in formalin. Sole bright ultramarine blue.” The
traces of the yellow marking in the preserved specimen (Fig. 67) are: the
rims of the rhinophoral sheaths, of the branchial pocket, of the notum and
56
the sole; about 5 rows of streaks on the notum, a line along the afferent
and the efferent branchial vessels; 4-5 series of partly confluent rings on
the sides of the foot; a median line on the tail. The yellow pattern, white
in preserved condition, is raised above the surrounding skin.
The notal border of the tail has 6-7 thick glandular knobs, continued a
little forwards as smaller granules. There are ten unipinnate gills; some
branchiae have several tips (Hoffmann, 1940: 46).
The anterior border of the foot is transversely grooved without a median

67-71. Hypselodoris edenticulata (W hite). 67, preserved slug; yellow

F ig u r e s
markings drawn black; 68, hooklet of jaw; 69, radular teeth of slug from
Alligator Reef Light (A ); 70, same from Palm Beach (P ); 71, diagram of
reproductive organs.
57
notch. The tentacles are folded at the tip. The rhinophores have 28 leaflets.
The mandibles are yellow and composed of basally stratified, single-pointed
hooklets (Fig. 68).
The radula is yellow, its formula is 116X246.0.246. The first lateral
tooth has a big upper cusp and a stalked, bifid, lower cusp on both sides of
the rhachis, probably an individual malformation (Fig. 70, P I). The
second and following teeth are bicuspid with a big upper and a narrower
lower cusp, 15 ^ in length. The upper cusp increases rapidly outwards
from 15 to 40 and keeps its length to around the 190th, then it
shortens and becomes blunt. The outermost about 25 teeth bear a row
of up to 10 blunt denticles on their under edge, most numerous and
most distinct from 230-240.
The preserved specimens from locality 1 are much swollen, not quite
40 mm long, about 25 mm high. Their color in life was “indigo blue with
yellow lines” (collector’s note). The tail is much contracted; its border
bears glands. The rhinophores have 33 leaflets. The radula is colorless,
except for the youngest rows which are yellowish; its formula is 92X 208.
0.208. The shape of the teeth is somewhat different from those in the slug
from locality 3. The first lateral tooth has a 25 ^ long upper and a 12 ^
long lower cusp, in the second tooth the corresponding measurements are
28 and 17, in the middle of the half-row, 46 and 16. From around the
186th tooth the lengths decrease. From the 150th tooth outwards there are
few weak and blunt denticles on the under edge, up to 10; these are more
distinct from about the 190th tooth. The outermost teeth have no denticles.
The specimen from locality 2 is swollen, colorless, with traces of blue in
its folds. It is 45 mm long, 22 mm high, 30 mm broad, and its sole is up
to 8 mm wide. There are about 30 rhinophoral leaves, nine unipinnate dark
gills, each with a light stripe along the efferent vessel; one gill has four tips.
The flat folded tentacles, the grooved anterior border of the foot without a
notch, the radula of 110 rows with 210 teeth per half-row, justify the
identification. The innermost lateral tooth corresponds to that from the first
locality. The labial cuticle bears pegs 20 /jl in length, hence much shorter
than the 40 ^ high stratified hooklets of the other specimens. The uniform­
ity of the small pegs suggests that they are not worn high ones, but
regenerated after a shedding of the old cuticle.
The slug from locality 4, already identified by Professor Bayer, is only
13 mm in length, 4 mm in width, and 3.5 mm in height. It was examined
one month after preservation in Bouin’s liquid and was still blue with light
longitudinal streaks on the notum. The sides of the foot bore three rows of
light round spots and rings. The notal glands mentioned for Ch. nyalya
which, preserved in Bouin, appear sepia (Fig. 63), occur also in Ch.
edenticulata from locality 4. It has nine gills.
The genital organs (Fig. 71) are as in White’s Figure 15, but her
explanation of spermatocyst and spermatheca must be transposed. A
58
vestibular gland drawn but not explained by White is not developed in the
examined slug from locality 1; only the nidamental duct is widened a little
near its end. The male duct ( d ) and female duct separate at the outlet of
the fusiform ampulla (a). The spermatheca (b ) is large, spherical; the
elongated spermatocyst (y) is located farther distally than the spermatheca,
the insemination duct (xo ) still a little farther outwards.
Discussion. — White compared her species with H. californiensis (Bergh,
1879a: 112), not with the similar and geographically much nearer H.
sycilla (Bergh, 1890a: 165) from the coast of Yucatan. Both calif or niensis
and sycilla bear denticulations on the lower prong of the cusp on all teeth;
those of sycilla are less developed than in calif or niensis. Such denticles are
altogether absent in White’s species. Our material has them on the teeth of
the external part of the half-row, namely in the outermost fourth (locality
1) or tenth (locality 2). In this respect our material is intermediate
between sycilla and White’s species. Nevertheless we call it edenticulata,
as previously (Anderson & Lane, 1963: 265), because it is quite close to
White’s species, and farther distant from sycilla.
The jaw elements of californiensis (Bergh, 1894: 181) and sycilla have
secondary points beside the hook; for sycilla these were only drawn
(1890a: PI. 3, Fig. 5), not described. In White’s and our specimens the
hooks are simple. The minute structure of the jaw elements furnishes a
more clear-cut distinction between californiensis and sycilla on one side
and edenticulata and ours on the other.
The relationship of californiensis, often mentioned in the literature
(O’Donoghue, 1926: 211) and sycilla, known by a single slug described
only once, is beyond the scope of the present discussion, because californi­
ensis with its strong denticulation of the radular teeth is sufficiently separ­
ated from edenticulata. For a color photo of californiensis we are indebted
to Dr. James R. Lance, who has found this beautiful species again (1961:
66); its pattern differs widely from edenticulata.
The 290 teeth in the half-row of the radula of sycilla outnumber not
only those of White’s small specimen, but even those of White’s and our
much bigger ones. White’s and our specimens have 25, 28 and 33 rhino-
phoral leaves respectively; sycilla has 40-45. The sole of sycilla whose color
was well preserved is dirty yellowish, that of White’s and our animals blue.
Whether the light pattern on blue ground color consists of continuous or
interrupted lines, rows of separate rings, or rings joined together to form an
interrupted band, cannot be used for systematics. White’s notes “in life
deep Prussian blue with brilliant yellow markings” which become light in
preserved condition, agree better with the present material than with
Bergh’s “blue with chalk white marks.” The substitution in life of white by
yellow with age is known for chromodoridids (Pruvot-Fol, 1951a: 135,
no. 157, Haefelfinger, 1959: 311).
59
 Without knowing its jaw elements and radula we cannot discuss the
Bermudian Chromodoris zebra Heilprin (1889: 327), and its possible
relations to Hypselodoris gracilis (Rapp, 1827) from the eastern Atlantic
Ocean and the Mediterranean Sea. The possibility that edenticulata is a
synonym of zebra cannot be excluded. Heilprin indicated 12-13 gills for
zebra; Smallwood (1910: 139) gave 12-14.

36. Hypselodoris acriba, spec. nov.

Figures 72-15
Material.— Los Roques Islands (E of Bonaire): Dos Mosquises; south end
of southern island. On living Acropora palmata. R. C. Work, August 11,
1963 (Work No. 7). Four specimens.
Syntypes .— USNM 576290.

Description .— The color of the living animals, from photos and collector’s
notes, is blue, hidden by yellow pigment on the notum. The area around
the mouth is blue, the anterior border of the foot yellow. The thin margin
of the notum (Bergh’s “Mantelgebrame” ) is brimmed with a fine white
line, closely accompanied by a pale red one. The inwards following yellow
color of the margin darkens gradually and ends on a bright yellow band,
visible also on the hyponotum, due to the thinness of the margin. On the
inner edge of the yellow band there is a row of about a dozen blackish
brown spots on either side, and next to them a row of irregular brilliant
blue blotches. The middle area of the back is stippled with yellow tubercles
and blue dots in the spaces of a yellow network. The same pattern occurs
on the sides of the foot, and further two rows of blue blotches as well as
one of violet spots on the limit towards the lavender-blue sole. The rhino­
phores are lavender, the bases of the gills brick-red, their tips yellowish.
The preserved slugs are up to 60 mm long, 18 mm broad, 15 mm high.
The tentacles are small and folded. The rhinophores have about 30 leaves
of which 20 are complete. The anterior border of the foot is grooved, its
lateral angles projecting and pointed. The sole is narrow, 4 mm in width.
Thick whitish glands lie along the border of the notum. There are eleven
bipinnate gills, in one specimen unipinnate with several bifid tips. The anal
papilla completes the circle of the branchiae behind.
The labial elements (Figs. 73, 74) are unicuspid. The radular formula
is 83X 175.0.175. The rhachis is naked. The innermost and lateral teeth
(Fig. 75) are of the same type, bicuspid with a stronger upper and a
considerably smaller lower prong. In about ten marginal teeth there are
two blunt cusps of equal size and a weak denticulation on the lower edge.
One specimen had an aberrant innermost tooth with a single hook widely
distant from the apex and a second tooth with one cusp in similar position
and a lower much smaller one.
60
Discussion.— The teeth in the center of the half-row agree with White’s
Figure 16 A; but there the innermost teeth (Figs. 16 B, C) are a little
shorter. White had no record of the living animal and classified it as
Glossodoris aegialia (Bergh, 1905b: 70) from the Gulf of California. As
that species has two lateral rows of eye-spots bearing pupillae and a
similar row in the anterior half of the back, it can hardly be identical
with White’s species which is smooth. The elements of the labial plates
and the radula of the present species agree sufficiently with Bergh’s, but
both, as well as White’s species, are of the regular H ypselodoris- type, so
that the agreement is of little weight. The arrangement of the pupillae in
aegialia excludes a nearer relation to acriba.

F i g u r e s 72-75. Hypselodoris acriba, sp . n o v . 72, liv in g a n im a l, fr o m c o lo r

p h o to ; 73, ja w e le m e n ts ; 74, s in g le r o d le t o f ja w ; 75, ra d u la r t e e th .

It is much more difficult to separate acriba from the Bermudian papillose,

broadly oval Chrom odoris roseo-picta Verrill (1900: 549; 1901: 33),
whose labial armature and radula are not known. Verrill’s species is much
smaller (25 X 10 mm) than acriba, but has more numerous (16) gills. The
distribution of the papillae and the color do not agree in both species, but
61
there may be intra-specific variations. The new name introduced here may
in future prove to be a synonym, but it is less troublesome to unite identical
species than to separate wrongly united ones.
The Mediterranean H. coelestis (Deshayes, 1866), also with notal
tubercles, is blue with a widely different color pattern (Pruvot-Fol, 1951b:
PL 1, Fig. 17).
Felimare, gen. nov.
Diagnosis.— Chromodoridinae (Odhner, 1926: 53) with a rhachidian plate
nearly as high as the neighboring teeth, provided with a smooth cusp. The
lateral teeth are bicuspidate. The labial armature consists of simple
hooklets. The penis is unarmed; the spermatheca has a single duct, the
spermatocyst lies farther distal.
Type-species.— Felimare bayeri, spec. nov.
R em arks. — The new genus is related to the Indo-West Pacific genus Ris-
becia Odhner (1934: 249), whose three species, the type frangoisi (Risbec,
1928: 159; 1953: 94; unnecessarily called odhneri), versicolor (Risbec,
1928: 147; 1953: 93), and reticulata (Quoy & Gaimard, 1832; Risbec,
1956: 10) have a denticulate cusp of the rhachidian plate and unicuspidate
lateral teeth.
37. Felimare bayeri, spec. nov.
Figures 76-79; Plate 1, Fig. 2
M aterial. — Florida: 1. Virginia Key, from rocks off east side (Sewage
Beach). May 15, 1962.— 2. Biscayne Bay, along south side of Ricken-
baeker Causeway, west of Virginia Key. March 1963.— A total of two
specimens.
H olotype.— USNM 576270.
Description. — The color alive (based on a photograph) is: notum blue with
6-10 longitudinal anastomosing orange yellow lines edged with black, the
more lateral ones often interrupted. The margin of the notum is white,
indistinctly set off from the blue ground color, containing black spots of
different size, and bordered by a yellow line externally. There is a yellow
circle around the blue rhinophores. The gills are colorless with a black line
along the afferent vessel and a yellow with black one along the efferent
vessel. The tail is white with orange yellow lines and black spots, edged
with blue. The sides of the foot bear yellow stripes.
The length of the preserved specimen (locality 1) is 21 mm, its height
and breadth 9 mm; the damaged animal from the second locality is 15 mm
long. The anterior border of the foot is not grooved. The shape of the
tentacles is not well preserved, evidently reaching the border of the notum
in front and that of the foot behind. The rhinophores have about 20
foliations. There are eight unipinnate gills. Six (locality 1) and four
62
F i g u r e s 76-79. Felimare bayeri, gen. nov., sp. nov. 76, living animal, from
color photo; stippled, yellow; hatched, blue; 77, hooklets of jaw; 78, radular
teeth; 79, diagram of reproductive organs.

(locality 2) prominent yellowish glands lie around the posterior end of

the no turn.
The labial cuticle has simple hooks (Fig. 77). The radula (Fig. 78)
comprises 90 rows with a central tooth and 106 lateral teeth in the half-row.
The rhachidian tooth is 80 ^ high, 24 ^ broad, its cusp is 10 ^ long; the
neighboring teeth are 85 ^ high, 16-18 ^ broad, with an about 25 n long
upper and an 8 p long lower prong. The teeth in the middle of the half-row
are 65-80 ^ long, 20-24 ^ broad; the upper prong reaches 40 ^ in length,
the lower prong 14 /x. The length of the cusps decreases outwards, about
63
9 of the marginal teeth are very irregular in one and the same longitudinal
row. The prongs are less unequal, but the lower one is cleft and followed
by blunt denticulations, so that the whole cusp appears almost pectinate.
The genital organs are as in other Chromodoridinae: the hermaphrodite
duct (ei), the large ampulla (a), and the male duct ( d ) with an inner
glandular and an outer muscular section. The ejaculatory duct ( p ) is
separated from the long, tubular atrium by a strong sphincter.
The vagina (v) leads to the lobular bursa ( b ). The sperms leave the
latter by the vagina and enter the longish spermatocyst (y). Opposite to
the connection of the spermatocyst with the vagina the sperms reach the
fertilization chamber (be) by the insemination duct (xo). The female gland
mass (an, mx) opens by the outer oviduct separated from the common
atrium.
The species is named for Professor Dr. Frederick M. Bayer.

38. Siraius kyolis, spec. nov.

Figures 80-84
Material.— Florida: Biscayne Bay; MacArthur Causeway, under rocks.
R. A. Smith, February and October 22, 1962. Seven specimens.
Syntypes .— USNM 576280.

Description .—According to the collector’s notes, the living slugs are “dirty
yellow with blackish spots” and “tan with greenish black spots.” The
preserved specimens are white, in one animal with yellow gills. The biggest
slug is 12 mm in length, 6 mm in breadth, 4 mm in height. The width of
the hyponotum is 2 mm, that of the foot 3 mm. The body is rigid. The
center of the notum is sculptured with low bosses of different sizes stiffened
by perpendicular spicules which do not project from them (Fig. 81). The
bosses on the sides of the notum are smaller and disappear near the edge.
There is a horizontal layer of larger spicules in the connective tissue under
the bosses, around which they lie radially. The underside is smooth, also
with horizontal spicules; a band of glands lies around the border of the
hyponotum.
Four lobes surround the pits of the rhinophores, whose clubs bear about
15 foliations. The tentacles are barrel-shaped with an outer furrow. There
are up to 11 unipinnate gills. The border of the branchial pocket is circular
with a lobed outline, or contracted to semilunar or reniform shape. The
anal papilla lies in the center of the plumes. Often some plumes are bitten
off; in one slug, only three remain. The foot is bilabiate and notched in
front, pointed behind; it nearly reaches the posterior border of the notum.
The labial cuticle is smooth, without markings. The radula (Fig. 82)
comprises 47 rows with 2.35.0.35.2 teeth. The inner teeth are strongly
64
F i g u r e s 80-84. Siraius kyolis, sp . n o v . 80, p r e s e r v e d s p e c im e n ; 81, sk in ;
82, ra d u la r te e th ; 83, d ia g r a m o f a lim e n ta r y tra ct; 84, d ia g r a m o f r e p r o d u c tiv e
org an s.

curved hooks; in the middle of the half-row the teeth become straighter;
the marginal teeth are pectinate with stronger denticles on inner and outer
edges. The teeth of the oldest rows are bigger than those of the younger
ones, suggesting that growth continues after the formation of the tooth in
the radular papillae. In the oldest row (compared with the 40th row), the
height of the teeth is: innermost tooth 44 ^ (30 ^ ); middle tooth 80 ^
(68 /x); 34th tooth 95 ^ (80 ^ ). The size of the outermost hook-shaped
and pectinate teeth decreases suddenly. The salivary glands are ribbon-like,
the left is longer than the right one. The stomach projects from the
intestinal gland; the right liver (r) has about half the breadth of the
stomach (so). One stomach contained part of a dorididan radula com­
prising three rows of teeth.
The thick winding ampulla (a ) divides at its outlet into the spermatic
65
duct and the inner oviduct. The latter is quite short, merging into the
large female gland mass (g). The former becomes glandular immediately.
This bent prostatic section ( q ) is followed by a narrow and short efferent
duct ( d ), whose ectal half is contained in a muscular sheath; its tip
projects into the male atrium.
The vagina (v) begins wide, narrows gradually, and enters the sperma­
theca ( b ) with a thin canal. The dark contents of the spermatheca shine
through the skin as a blackish spot. The spermatocyst ( y ) lies near the
spermatheca; all sperms are fastened with their heads to the fundus of the
spermatocyst. This seminal receptacle is connected with the insemination
duct ( xo ) by its own canal. The insemination duct opens into the gland
mass near the entrance of the inner oviduct.
Discussion .— Since the type-species of Siraius, S. ilo Marcus (1955: 135),
was described, 19 specimens of it have been seen, all greyish yellow without
dark spots. From the coast of Sao Paulo the range of ilo was extended to
Cabo Frio (1958b: 57). It has 22 branchiae and more teeth per half-row
than kyolis, but only 37 radular rows. In Siraius as in Austrodoris (Odhner,
1934: 269), the shape of the salivary glands cannot be maintained as a
generic character. In S. ilo they are sac-like, short and broad, in kyolis
ribbon-like, similar to those of Doris bovena Marcus (1955: Fig. 117, r).
They are still longer in Doris bicolor Bergh (1884b: 655), a small
Mediterranean species which probably belongs to Siraius. With respect to
the number of gills and of teeth per half-row, bicolor is near kyolis but it
has a maximum of 34 radular rows, up to six pectinate teeth, and a con­
siderably longer right liver than kyolis.
The seminal receptacles, called seminal vesicles by Bergh, were called
vaginal in Siraius. If, however, those of Homoiodoris Bergh (1882: PI.
6, Fig. 16) were qualified as semiserial (Odhner, 1926: 54), the vesicles
of S. ilo and S. kyolis would be semiserial too. In the vaginal type (Odhner,
1926: 51), the two vesicles lie quite near to one another at the inner end
of the vagina, while in the semiserial and in the serial type the spermatocyst
is connected with the insemination or uterine duct. In the former type, the
insemination duct originates from the entrance of the vagina into the
spermatheca, as in Siraius; in the latter it leaves the spermatheca inde­
pendent from the vagina, as in Peltodoris greeleyi (Fig. 98). The true
vaginal arrangement occurs, e.g., in Austrodoris Odhner (1926; 1934),
but it is rare in the doridids. In Doris, another genus with vaginal seminal
vesicles, these do not lie so near together (Marcus, 1955: Fig. 106, the
type-species) as in Austrodoris. In Neodoris Baba (1938: 13) the semi­
serial and the serial types occur (Marcus, 1959a: 37 ff).
39. Anisodoris worki, spec. nov.
Figures 85-89
Material.— Florida: Biscayne Bay; under rocks, east end of County
66
(MacArthur) Causeway. R. C. Work, August 7, 1964. One specimen.
Holotype .— USNM 576289.
Description .— The color photo shows a slug 28.5 mm long, 16 mm broad.
This is evidently the natural size, as the preserved specimen is 20.5 mm
in length, 11.5 mm in breadth, and 6.5 mm in height. The breadth of the
hyponotum is 4 mm; the foot is 3.5 mm broad but rolled in.
The color alive is yellowish; the papillae, rhinophores, and gills are
darker yellow. Dark spots in the connective tissue are interrupted by the
papillae, which are lighter than the spots. The middle of the back is darker
yellow. The papillae are smaller on the margin of the notum. The hypo­
notum and the foot show small black specks; there is a row of eight
larger, longish spots on either side of the inner border of the hyponotum
(Fig. 86). In the preserved animal there are small brown chromatophores
on the tips of the round papillae, in the rhinophores, and in the gills.
The notal sculpture is cauliflower-like (Fig. 86), as the interspaces
between the larger papillae (0.8 mm) are filled with small ones (0.15 mm).
The pits of the rhinophores and branchiae are surrounded by middle-sized
papillae. The spicules are often blunt and do not project above the surface.
The tentacles are rather long, flattened clubs without furrows; there are
about 25 foliations on the rhinophores, and six multipinnate gills with
warts on their outer sides. The anterior border of the foot is transversely
grooved, the upper lip notched in the middle.
The labial cuticle is smooth. The radula has 27 rows and 38 hamate
teeth per half-row (Fig. 87). The size of the innermost tooth is 40-47
from the second to the third tooth the size increases from 50 to 80
The teeth in the middle of the half-row measure 160 the outermost
tooth 45-52 [A, the penultimate and the antepenultimate teeth 85 and 100 ^
respectively. As the fourth tooth from the outer border is 150 ^ long, the
fourth from the middle 95 the pararhachidian teeth are smaller than the
outer ones. The stomach is not covered by the intestinal gland.
The ampulla (a) is long and winding, the female gland mass (g) large,
wrapping the seminal reservoirs. The male duct leaves the gland mass,
widens into a folded prostate ( q ), and runs looping ( d ) towards the
muscular, strong penis ( p ) lodged in the atrium. The vagina opens
together with the penis, the nidamental duct farther ventrally. The vagina
(v) is narrow; it enters the spermatheca ( b ). The short insemination duct
( x o ) leaves the spermatheca at this entrance. The spermatocyst (y) is
inserted on the duct which continues into the gland mass. The ectal section
of the vagina bears a flattened gland ( va ), connected with the vagina by a
short, muscular duct (vz). The wall of this duct (Fig. 89) contains a large
papilla with a spikelet and many smaller papillae, also with spikelets,
around the bigger one.
67
 The species is named for the collector.
Discussion .— In his original diagnosis of Anisodoris, Bergh (1898b: 508)
included “tentacles as in Archidoris;,> that is: “low, fold-like” (Bergh,
1878: 616). In the type-species, Archidoris tuberculata, they bear an

F i g u r e s 85-89. Anisodoris worki, sp . n o v . 85, liv in g a n im a l, f r o m c o lo r p h o to ;

86, u n d e r s id e o f f o r e e n d o f p r e s e r v e d slu g ; 87, r a d u la r te e th ; 88, d ia g r a m
o f r e p r o d u c t iv e o r g a n s; 89, d u c t o f v a g in a l g la n d .

68
external furrow. When Eliot (1910: 95) selected the type-species of
Anisodoris, A . punctuolata (d’Orbigny, 1837: 187), he did not consider
its pointed tentacles; in other species there are furrowed lobules (Marcus,
1959a: 48). Nor can “tentacles finger-shaped” (Pruvot-Fol, 1954: 239)
be used as a character of Anisodoris, and even less “vestibular gland
present” (ibid.). The smooth labial cuticle is not mentioned in Pruvot-Fol’s
diagnosis, though it is the only character that separates A nisodoris from
Discodoris with labial rodlets. Sometimes these are very short (MacFarland,
1909: 75) or absent (Marcus, 1955: 148) in Discodoris, possibly worn
or in regeneration. Often the notal papillae of A nisodoris are stronger than
the granules of the generally softer Discodoris, while the rigid notum of
Peltodoris becomes velvety by densely standing, uniform caryophyllidia.
Therewith the general aspect of most species of these genera is different. In
preserved specimens, however, the notum may be swollen or shrunken
and the sculpture modified. The characters of sculpture are difficult to
describe unequivocally in different languages, and, moreover, exposed to
subjective preferences, e.g., “pustules” (Burn, 1962c: 156) and “Nopp-
chen” (Bergh, 1905a: 98).
The species nearest to Anisodoris worki is Archidoris marmorata Bergh
(1881: 86) which belongs to the Discodoridinae due to its prostate
(Odhner, 1926: 64-65), and to A nisodoris due to its smooth labial cuticle
and notal sculpture. The Anisodoris m arm orata Bergh (1898b: 515) from
northern and middle Chile (Marcus, 1959a: 45, 88) must be re-named
as A. rudberghi, nom. nov.
Bergh’s Archidoris m arm orata came from Trieste. It differs from w orki
by the absence of spines in the duct of the vestibulo-vaginal gland (Bergh,
1881: 92), by the reddish brown back with white nodules and violet black
clubs of the rhinophores, by the size of the body (living 47, preserved
nearly 40 mm), and by eight gills.
Vayssiere’s A rchidoris marmorata Bergh (Vayssiere, 1901b: 16) from
Marseille cannot be the same species, because its labial cuticle bears
rodlets, nor can it be an Archidoris. It is not known whether it has a
prostate and a vestibular gland. Both features, labial rodlets and undes­
cribed prostate and accessory gland, also make it difficult to define A rchi­
doris stellifera Vayssiere (1903: 82) from the Gulf of Lions and Naples.
Certainly it is not an Anisodoris. Vayssiere (1904: 129; 1913: 314-315)
considers the radular teeth of his marmorata Bergh and stellifera as
specifically different.
Pruvot-Fol (1954: 240) unites marm orata Bergh, 1881, marmorata
Vayssiere, 1901, and stellifera Vayssiere, 1903, under the name A nisodoris
stellifera Vayssiere, 1904. Her figure 90 combines details from the publi­
cations in 1881, 1901, and 1904, though only the reproductive organs of
marmorata Bergh (her figure 90, e) are known. It is possible that she had
material of marmorata Bergh, 1881, as she writes: “labial cuticle unarmed,
69
folded, sometimes pierced by some hairs, which do not constitute an
armature.”
Accessory vaginal glands are frequent in the Dorididae (Bergh, 1892a:
1086); if not combined with other structures, they are no more than
specific characters.
The vaginal appendage of Anisodoris fiindersi Burn (1962c: Fig. 11,
sp) is considered as a spermatheca by the author. The sculpture of the
notum of fiindersi differs by two rows of high tubercles from that of worki.
Also A. timorensis Bergh (1905a: 97), whose reproductive organs are
not known, differs by the tubercles of its back, 80-120 ^ in diameter,
from worki.
40. Anisodoris prea, spec. nov.
Figures 90-93
Material.—Florida: Biscayne Bay; Matheson Hammock; Thalassia and
sand flats. F. M. Bayer, February 21, 1948. Four specimens.
Syntypes .— USNM 576278.
Description .— The biggest animal is preserved 28 mm in length, 19 mm in
breadth, and 8 mm in height. The hyponotum is 4 mm broad. The smallest
specimen is 17 mm long, 10 mm broad. The color is a creamy white,
without traces of pigment; the body is stiff. The notum is covered with low,
flat knobs of different size, each big one surrounded by a number of small
ones. The middle of the back is nearly smooth, sharply delimited on either
side by a row of large tubercles which form a kind of crest. Towards the
sides the knobs are smaller. The spicules are partly dissolved, but sections
(Fig. 91) show that they were radially dispersed in the knobs and
projected slightly from them.
The pits of the rhinophores bear a bigger tubercle on either side. There
are about 30 rhinophoral leaflets containing spicules. The tentacles are
flat, broader than long, contracted. The border of the branchial pocket is
retracted; there are seven multipinnate gills. The foot is bilabiate, its upper
lip notched.
The labial cuticle is smooth. The radula is 2.3 mm in length, 2 mm in
width. Its formula is 25X29-30.0.29-30; the last 5 rows are still weak.
The rhachis is broad. All teeth (Fig. 92) are hook-shaped, the inner ones
bigger (0.13 mm) than the outer ones (50 ^ ). The maximum height in
the middle of the half-row is 0.2 mm. The stomach is free, covering the
spherical right liver. The salivary glands are long.
The ampulla ( a ) is winding, its outlet separates into the spermatic and
the oviduct. Where the latter enters the female gland mass (g), lies the
small albumen gland. The male duct passes immediately into the folded
prostate ( q ), which is much longer than wide. The following section of the
efferent duct ( d ) is coiled and ectally enclosed in a muscular sheath. A
70
short conical penial papilla projects into the atrium. The vagina (v) begins
wide, narrows farther inwards, forms a loop, and enters the spermatheca
( b ). As in the other species of the genus the entrance into and the outlet
from the spermatheca is established by a short single duct; the seminal
reservoirs are arranged semiserially. The insemination duct (xo ) or com­
munication between the receptacles is angled in a similar way as in A.
nobilis (Marcus, 1961a: Fig. 57), and also its continuation, the insemi­
nation duct of the mentioned figure, is quite short in the present species;
it goes out from the spermatocyst ( y ).

F i g u r e s 90-93. Anisodoris prea, sp. nov. 90, dorsal view of preserved slug; 91,
section of skin; 92, radular teeth; 93, diagram of reproductive organs.

Discussion .— The species of Anisodoris with ectal appendages of the female

organs are mentioned in the preceding discussion of A. worki. All the
other species are treated by d’Orbigny (1837: 186-189); Bergh (1898b:
509-520; 1905a: 96-98); MacFarland (1905: 38; 1906: 116); Marcus,
71
(1961a: 17); Odhner (1926: 85-88); and Marcus (1959: 48-50). None
of them has the peculiar sculpture of the notum as prea, whose nearly
smooth middle is flanked by big tubercles.

41. Peltodoris greeleyi MacFarland, 1909

Figures 94-98
References. — MacFarland, 1909: 84; Marcus, 1955: 137.
M aterial .— Florida: Virginia Key, from red rocks (riddled with Cliona )
off east side (Sewage Beach). F. M. Bayer, H. B. Owre. May 1963. Two
specimens.
Further distribution.— Brazil, coasts of Alagoas (original locality), Cabo
Frio, and Sao Paulo.
D escription. — The animal shown in a color photo is 35 mm in length,
29 mm in breadth. The preserved specimens are 29 and 26 mm long,
18 mm broad, and 7 mm high. The length of the foot is 23 mm, its width
9.5 mm, so that it projects over the 4-5 mm broad hyponotum.
In life the color is a very light brown with more or less radially arranged
dark brown blotches. These are small in the center of the back, bigger and
transverse in one row on either side between the rhinophores and the
branchial pocket, and smaller around the rhinophoral sheaths and the notal
border, where they are rather close to one another. The rhinophores and
gills become increasingly darker brown upwards; the tips of the gills are
nearly bluish black. The rim of the rhinophoral pockets, the tips of the gills,
and the border of the notum are sparsely frosted with white secretion.
The body is depressed, hard and rigid, its border undulate. The surface
of the notum is villous and velvety due to densely set caryophyllidia (Fig.
97). These are supported by around seven spicules, 0.18 mm in length,
which end with a basal knob. The numerous further spicules in all parts
of the skin, up to 0.6 mm in length, 30 ^ in diameter, are often rough.
The rhinophores have about 20 foliations, their sheaths are high with
crenulate rims. Also the border of the branchial pit is prominent. As in
Brazilian greeleyi (Marcus, 1955: 138, Fig. 128) and possibly in P.
crucis (ibid.: 139) there is a pad in front of the mouth, probably originated
by the bulging walls of the mouth cavity. The leaflike, pointed tentacles
are apposed to the pad ventrally. The anterior border of the foot is bila­
biate, the upper lip notched. The 14 unipinnate gills surround the anal
papilla in a circle.
In the examined specimen the labial cuticle is smooth, but shows
quincuncial pits such as occur also in the previously described greeleyi
(Marcus, 1955: 138) and P. noumeae Risbec (1956: 8). The radula is
4.7 mm long, 2.5 mm broad; its formula is 75 X 98.(1) .98. The false
rhachidian plate (Fig. 96) has no cusp; the lateral teeth bear cusps, also
72
the outermost ones. The rhachidian plate is 24-34 ^ high, the third lateral
tooth 65 fi, and the following teeth increase slowly to the maximum height
of 0.16 mm in the middle of the half-row. From there outwards they de­
crease very little, so that the penultimate tooth is still 0.13 mm, and

F ig u r e s 94-98. Peltodoris greeleyi MacFarland. 94, living animal, from color

photo; 95, underside of fore end of preserved slug; 96, radular teeth; 97, cary-
ophyllidium; 98, diagram of reproductive organs.
73
only the outermost one is small, 60-80 The large, strongly muscular
stomach lies free.
The winding ampulla ( a ) is set off from the hermaphrodite duct (ei).
The male and the female duct separate at the outlet of the ampulla. The
spermatic duct enters the broad and flat, lobate prostate ( q ) immediately.
The following coiled section of the efferent duct ( d ) is sheathed by silky
muscle fibers and ends with a thick, soft papilla ( p ) in the atrium. Near
the entrance of the oviduct into the female gland mass lies the small
yellow albumen gland ( an). The mucus gland (mx) passes into a wide
nidamental duct ( o v ) which is still glandular close to the outer opening.
Between the penis and the nidamental duct the straight vagina (v) courses
inwards. Its entrance into the spermatheca ( b ) is far distant from the
origin of the insemination duct (xo). This duct receives the short canal
of the spermatocyst ( y) near its opening into the female gland mass.
Discussion.— As the genus D iscodoris comprises more than 40 species, it
is practical to separate the subdepressed species with a rigid, minutely
granular notum and labial cuticle without rodlets or hooklets under the
name Peltodoris Bergh (1880a: 41). A subdivision of Peltodoris according
to unipinnate (greeleyi) or multipinnate gills (the eight remaining species)
may be pondered, but in the original specimen of P. noumeae Risbec
(1937: 159; 1953: 40; 1956: 8) most of the branchial plumes are
bipinnate, and the posterior ones simply pinnate. That specimen was small
but evidently mature. The species of Austrodoris Odhner (1926: 25 ff;
1934: 255 ff.), a genus well delimited geographically and anatomically,
have different types of branchiae. In some, large and small animals have
quadripinnate and unipinnate gills respectively, in others the plumes of
one and the same branchial circle are differently pinnate.
The present specimens cannot be separated specifically from the
Brazilian P. greeleyi, while P. hummelincki Marcus (1963: 27) from
Aruba and Curagao has neither caryophyllidia nor unipinnate gills. Dif­
ferences of little weight between the Floridian and the Brazilian greeleyi
lie in the shape of the body and the color pattern. The much greater
numbers of rhinophoral foliations, radular rows, and teeth per half-row
have probably no systematic value, because the present specimens are
bigger than those from Brazil.
The same holds for the false rhachidian plate, which O’Donoghue
(1929b: 78) also found in the bigger animal of another doridid, but not
in the smaller specimen of the same species. The basal knobs of the spicules
in the sensory papillae (Fig. 97) of the present material were not men­
tioned in the descriptions of the Brazilian animals, but they occur there
too, at least in the material from Sao Paulo, re-examined with regard to
this character.
74
 42. D iscodoris evelinae Marcus, 1955
Reference.— Marcus, 1955: 143.
M aterial.— Florida: 1. Palm Beach Inlet; east side of Peanut Island; on
rocks, 1-2 ft. R. C. Work, May 12, 1962.— 2. Same locality; on rocks in
2 ft. of water at low tide. R. C. Work, June 9, 1962. (A total of two
specimens.)
Further distribution. — Brazil, coast of Sao Paulo.
Rem arks. — The color of the first animal in life was described as dull, that
of the second as pale rose orange with dark rose orange spots. The two
Brazilian specimens were olivaceous brown with darker blotches above, and
lighter with large dark spots below.
The size of the second of these specimens was about 9 cm in life; it has
six gills, whereas the other animal has seven. Many spicules have rough
ends. The radula formula of the second slug, 38X71.0.71, is a little, but
certainly not specifically, different from that of the original material,
35 X 77-80.0.77-80. The longitudinal serpentine fold along the rhachis
between the alternate rows of teeth is a peculiar character, but occurs
also in the following species.
The male copulatory organ is spiny also; it is systematically very im­
portant. The two other species with strong penial spines occurring in
western Atlantic warm waters, D. branneri MacFarland (1909: 66) from
northeastern Brazil and D. hedgpethi Marcus (1959b: 254) from the
northwestern Gulf of Mexico, are immediately distinguishable from eve­
linae by the maximum of 48 teeth in the half-row.
As in the Brazilian material and in other species of the genus, to which
names such as fragilis and notiperda were given, autotomy of pieces of the
notal or pedal brim had happened in the second of the present specimens
when handled by the collector. He saw the slug swimming actively by
undulations of the margin. Another big species of the Discodoridinae,
Sebadoris crosslandi (Eliot, 1903: 368), when dropped into a few feet
of water, finds its way to the bottom with movements like those of a
flat-fish (id., 1910: 19).

43. D iscodoris hedgpethi Marcus, 1959

Figures 98A-98E
Synonym . — D iscodoris spetteda Marcus & Marcus, 1966: 181. Nomen
nudum.
Reference.— Marcus, 1959b: 254.
M aterial.—Florida: Biscayne Bay, Bear Cut, Key Biscayne. March 21,
1965. One specimen.
Further distribution. — Texas: Port Aransas.
75
Description .— The preserved slug is 55 mm in length, 45 mm in width,
and about 20 mm in height. The sole is 45 mm long, 25 mm broad. The
notum is greyish brown with irregular darker spots, smaller to the sides,
and with blackish blotches in about three rows. The brown spots on the
sole are smaller than the dorsal ones, and those on the hyponotum are
darker than the spots on the sole. The broad border of the notum is
strongly undulated. The anterior border of the foot is bilabiate, with a
notched upper lip.
The sculpture of the notum consists of densely set tubercles (Fig. 98 A ),
about 0.8 mm in height and 0.4 mm in diameter. The tubercles are mamil-
late, tipped with white, and stiffened by parallel spicules which are dis­
solved; probably some of them had projected from the surface. The white
tentacles are conical and 4 mm in length (Fig. 98 B). The rhinophores
stand 10 mm apart from one another and from the anterior border; their
club has 23 leaflets. There are six multipinnate gills. Their color is that
of the notum, but is peppered with brown dots. The rims of the rhino-
phorial and branchial pockets are beset with tubercles of the same type
as the remaining notum.
The two triangular areas of the labial cuticle consist of 0.2 mm high
rods, 7-8 n in diameter. The radula is colorless, 7 mm long, 4.5 mm broad.
It comprises 35 rows and 47-53 teeth in the half-row. All teeth are hooks
with smooth cusps, which are 250 ^ long in the middle of the half row,
decreasing inwards and outwards. A serpentine fold runs along the rhachis
(Fig. 98 C) as in the preceding species. To the right or to the left of the
rhachis a rudimentary tooth occurs in a few rows. The salivary glands are
short and broad, white ribbons. The gut contains tissues of a red sponge.
After leaving the massy prostate the efferent duct runs straight to the
11 mm long penis (Fig. 98 D) lodged in a spacious male atrium. The base
of the penis is 1.7 mm in diameter and bears 0.25 mm long cuticular hooks.
From there the papilla tapers gradually towards the 0.55 mm wide tip.
The papilla is covered with smaller, scale-like hooks, set on muscular warts
which touch at their bases (Fig. 98 E ).
The female organs are as in other species of the genus; the vagina is
unarmed.
Discussion .— The present specimen does not have the denticle on the
innermost tooth as mentioned in the original description (Marcus, 1959b:
Fig. 8), but this is an anomaly, as we verified by examination of the other
specimens of the original material. Therefore, the name spetteda Marcus,
1966 (Stud. trop. Oceanogr. Miami 4: 181) must be suppressed.
In a key the West Atlantic species of Discodoris with armed penis can
be arranged in the following manner:
1. Notal projections are caryophyllidia ................................... mortenseni
1. Notal projections are conical tubercles or broad w a r ts ......................2
76
 98A —98E. Discodoris hedgpethi Marcus. 98 A, sculpture of notum;
F ig u r e s
98 B, ventral view of fore end; 98 C, rhachis and first lateral teeth; 98 D,
penis; 98 E, surface of same.

77
2. Half-row of radula with a minimum of 71 teeth ............... evelinae
2. Half-row of radula with a maximum of 53 teeth ............................ 3
3. Rhachis of radula deeply grooved longitudinally; penis of adult
4 mm long, its hooks up to 27 p in h e ig h t................................. branneri
3. Rhachis of radula with longitudinal serpentine fold; penis of adult
11 mm long, its hooks up to 250 ^ in h e ig h t.......................... hedgpethi

44. Discodoris phoca, spec. nov.

Figures 99-101
Material.—Florida: Key Biscayne, among mangroves, west side of island;
feeding on sponges. G. Y. Hendrix, June 16, 1963. One specimen.
H olotype .— USNM 576272.
Description .— The color photo shows the slug 55 mm long, 27 mm broad;
the notum is purplish brown with a net of white dots, mainly to the sides
of the center. The preserved animal is grey with pigmented spots and
blotches between a pigmentfree net. The dark markings consist of finely
branched melanophores. These are densely set on the sides of the foot,
scarce on the hyponotum and sole, and lie deep in the connective tissue.
The tentacles are spotted, the rhinophores black with white tips; the gills
are black and the branchial pocket is white.
Measured over the back the slug is 70 mm long, its linear length is
42 mm, the breadth 32 mm, the height about 25 mm. The hyponotum is
10 mm in wddth; the rhinophores lie 10 mm behind the fore end, the
branchial pocket 13 mm in front of the rear.
The notum is beset all over with caryophyllidia which often contain
melanophores on their top. The caryophyllidia also attain the contracted
borders of the rhinophoral pits. The rhinophoral clubs have about 20
leaves. The tentacles are finger-shaped, cylindrical, with pointed tips. The
borders of the branchial pocket are frilled due to contraction. There are
six tripinnate gills. The anterior border of the foot is bilabiate with a
notched upper lip.
The yellow labial cuticle consists of two triangular areas; the rodlets are
50 /x high, 5 [jl in diameter, and composed of round layers as in D. aurila
(Fig. 109). The radula is yellow, 4.3 mm long and 3.7 mm broad; its
formula is 27X42.0.42. All teeth are smooth hooks (Fig. 100). From the
innermost tooth whose base is 85 ^ in length, the size increases gradually
towards the middle (0.23 mm) and remains so to the outermost sixth.
The base of the 39th tooth is still 0.183 mm long. Outwards the size
decreases rapidly. The base of the outermost tooth is 60-85 ^ long, and
it has a cusp 0.11 mm long.
The peritoneum contains some melanophores; over the blood glands it
is black. The lining of the oral tube is black too; the buccal mass is white.
The salivary glands are long and thin. The oesophagus is dilated, crop­
78
like, and folded longitudinally. The stomach which contains soft masses,
no spicules, projects from the liver; the globular right liver is visible behind
the stomach. The intestine courses winding on the right side of the
intestinal gland.

F ig u r e s 9 9 - 1 0 1 . D iscodoris phoca, sp . n o v . 9 9 , liv in g a n im a l, f r o m c o lo r

p h o to ; 1 0 0 , r a d u la r te e th ; 1 0 1 , d ia g r a m o f r e p r o d u c t iv e o r g a n s.

The outlet of the ampulla (a) divides into the male and female ducts.
The spermatic duct immediately enters the massive prostate ( q ) whose
transparent ental part consists of small cells, the yellow ectal part of large
ones. Also the following tubular section is glandular; the next coiled stretch
of the efferent duct ( d ) is narrow, and its short ectal portion runs within
a muscular sheath. The strongly muscular male atrium lodges the small
penial papilla. Neither atrium nor papilla bears spines, as was determined
by the examination of sections.
The rather long vagina (v) leads to the spermatheca ( b ) which is
surrounded by the coils of the efferent duct. The insemination duct ( xo )
begins at the entrance of the vagina. The spermatocyst ( y ) has its own
canal; behind its connection with the insemination duct the latter merges
into the gland mass ( g ) near the ampulla and the quite short inner oviduct.
79
Within the gland mass the insemination duct forms a thin-walled, globular
vesicle, the fertilization chamber ( b e ) , filled with sperm, similar to that
of Thordisa diuda (Marcus, 1955: Fig. 140, r ).
Discussion — Of the Atlantic species of Discodoris, four have a spiny penis
by which they differ from phoca. All of these occur on the American coast.
They are: (1) branneri MacFarland (1909: 66) with a longitudinal row
of five or six larger black spots along either side of the notum; (2) evelinae
Marcus (1955: 143) has mamillate warts, not caryophyllidia; (3) hedg-
pethi Marcus (1959b:254) differs from evelinae by more radular rows,
fewer teeth per half-row, and longer and stronger spinous male organ; and
(4) mortenseni Marcus (1963: 30) is similar to phoca, but different by
minute penial spines, small size, in life up to 20 mm, a white foot, the
breadth of the radula little inferior (92.3 per cent) to its length, one
fourth of the outer teeth decreasing in size, and often the two outermost
teeth are dentate.
The remaining Atlantic species of D iscodoris have no penial spines, as
phoca, or the armature of the male organ is not described. American
species are: (5) alba White (1952: 113), in which the last third of the
half-row decreases in size, and whose black spots were only described for
the back; (6) m uta Bergh (1877b: 532), with the base of its innermost
tooth (PI. 62, Fig. 17) split, and without black spots on the sole; (7)
notha Bergh (1877b: 530) with a muscular and cuticularized cone beside
the aperture of the male atrium; (8) purcina of the present collection,
whose radula is broader than long; (9) pusae Marcus (1955: 147) with
8-12 marginal teeth developed as thin lamellae; and (10) voniheringi
MacFarland (1909: 73), whose six gills are bipinnate and whose labial
cuticle consists of three fields, a median and two lateral ones.
The Central and Eastern Atlantic and Mediterranean species of D isco­
doris differ from phoca; (11) cavernae Starmiihlner (1955: 217) by two
ramified vestibular glands; (12) edwardsii Vayssiere (1902: 232) by
2-3 rudimentary innermost teeth; (13) erubescens Bergh (1884b: 662)
by its color, white with reddish hue alive and 12 bipinnate gills; (14)
indecora Bergh (1881: 108) by its small size (alive 21 mm) and color,
olive brown with whitish spots; (15) maculosa Bergh (1884b: 658) by
its rhachidian platelets; (16) rubens Vayssiere (1919: 65) by its 7-8
lamellate marginal teeth; and (17) tristis Bergh (1899: 11) by its tentacles
grooved on the outer side.
The Indo-Pacific representatives of Discodoris were only cursorily
compared with phoca, with exception of the two species from the American
Pacific coast, heathi MacFarland (1905: 39; 1906: 118) and aurila in the
present collection. The former— as well as D. fulva O’Donoghue (1924:
27), evidently a juvenile heathi— has 12-16 outermost teeth in form of
thin, concave plates. D. aurila has a very differently colored notum and
no melanophores on the underside.
80
 45. Discodoris purcina, spec. nov.
Figures 102-104
Material.— Florida: Southwest Point, Biscayne Key. F. M. Bayer and E.
Deichmann, 1962. One specimen.
Holotype .— USNM 576281.
Description .— The length of the preserved animal is 25 mm, its breadth
18 mm, the height 7.5 mm. The foot is curved, 17 mm long, 4 mm broad.
The notum is transparent greyish with fine black granules, swollen, not
well preserved, probably bearing small papillae that had been stiffened by
spicules which are dissolved. The border of the notum is even, not
undulated. The hyponotum is broad, the underside colorless. A general
sulphureous hue may be due to preservation.
The tentacles are rather thick and pointed, not grooved. The rhinophores
are blackish owing to melanophores, and they have about 20 leaflets.
The anterior pedal border is bilabiate, the upper lip notched. The six
black bipinnate gills are of small size. The borders of the rhinophoral and
branchial pockets are smooth.

F ig u r e s 102-104. Discodoris purcina, sp . n o v . 102, la b ia l fie ld s; 103, r a d u la r

te e th ; 104, d ia g r a m o f r e p r o d u c t iv e o r g a n s.

81
 The labial cuticle bears two triangular fields of yellowish, stratified
rodlets, up to 50 ^ high and 5 ^ in diameter. The radula is colorless, very
short, broader (3.3 mm) than long (2.5 mm), and comprises 16 rows
and 43 teeth per half-row. All teeth are hook-shaped with smooth cusps.
The innermost tooth is 90 f± long, the six following teeth are also small.
Then the size of the teeth increases to 300 ^ and decreases in about six
of the outer teeth. The stomach is free, not enclosed in the intestinal gland.
The ampulla is fusiform; the spermiduct and the female duct separate
from one another in the female gland mass. The prostate is sausage-shaped.
The male atrium contains crystals, probably precipitates resulting from
the “hexamine” buffering of the formalin (letter of Prof. Bayer of May 12,
1964), which hinder the examination of the copulatory organ. The vagina
has no special gland; the insemination duct leaves the spermatheca to­
gether with the entrance of the vagina into the latter. The insemination
duct is very long, and its entrance into and outlet from the spermatocyst
are united.
D iscussion.— The principal characters of D. purcina are the unicolored
notum with black gills, and the exceptionally short radula which is broader
than long. By its radula D. purcina comes near to only two of the nine
species of the genus that were recorded from West Atlantic warm waters,
D. alba White (1952: 113) from Dry Tortugas, and D. mortenseni
Marcus (1963: 30) from Curasao and Tobago. In both of them the
radula is a little longer than broad. The innermost tooth of alba is bluntly
hamate, and the teeth in the center of the half-row have rather short
square bases. D. mortenseni has an expansion near the base of the inner­
most tooth, and also the basal expansions in the middle of the half-row
are more pronounced than in purcina. Outermost teeth with denticles
are frequent in mortenseni.
A small number of radular rows occurs also in the Californian D. heat hi
MacFarland, 1905. Moreover, its notum is similar to that of purcina in
color and sculpture. However, it differs from purcina by its 8-10 bran­
chiae and the quite different shape of all radular teeth (Marcus, 1961a:
Fig. 63).
46. D iscodoris pusae Marcus, 1955
Figures 105-107
R eference.— Marcus, 1955: 147-150, Figs. 151-165.
M aterial. — 1. Florida: Biscayne Bay. Under rocks at east end of County
Causeway. R. C. Work, April 11, 1963. One specimen.— 2. Florida:
Biscayne Bay, off Matheson Hammock. March 6, 1965. Shrimp trawler.
Three specimens.
Further distribution .— Brazil, northern coast of the State of Sao Paulo,
shallow water.
82
Description .— As the specimens from the two mentioned localities are
different in size and some inner characters, they are described separately.
The animal from the first locality, accompanied by a color photo, has a
pinkish red notum, darker in the middle than on the sides. There are
nearly contiguous violet red blotches along the mid-line and more spaced

F ig u r e s 105-107. Discodoris pusae Marcus. 105, living animal, from color

photo; 106, radular teeth; 107, diagram of reproductive organs.

smaller ones on either side between the rhinophores and the branchiae.
Small dark and blunt knobs of irregular sizes and intervals are scattered
all over the notum, those near the margin are smallest. The tip, shaft,
and anterior side of the rhinophorial club are whitish; basal and posterior
parts of the rhinophores dark. The outer side of the gills is chalk-white,
their inner side violet with rare white stipples, probably secreted matter.
83
Also the red anal papilla is frosted with white. Long white spicules form
a horizontal lattice under the surface and around the border of the notum.
The length of the preserved animal measured over the back is 30 mm,
lineal 20 mm, the breadth 18 mm, the height 10 mm. The width of the
hyponotum is 7 mm, equal to that of the sole. The anterior border of the
foot is bilabiate, its upper lip notched. There are about 20 rhinophoral
foliations; the border of the pit is slightly lobate. The tentacles are flat,
with broad base and pointed tip. Of the six gills the anterior ones have
small, bipinnate plumes; the posterior ones are much bigger and tripinnate.
This character is probably due to a regeneration of lost gills. The border
of the branchial pocket is smooth. The spicules were dissolved.
The labial armature consists of two nearly triangular fields. The
stratified rodlets measure 60 /a in height, nearly 10 ^ in diameter. The
radula comprises 26 rows, each half-row contains 36-38 lateral and 15-18
marginal teeth. The rhachis is naked, the innermost tooth is often rudi­
mentary. The marginal teeth are soft, blunt, and concave lamellae; their
edges bear minute irregular denticles, hardly discernible. The lateral teeth
are smooth hooks whose size increases outwards, only three laterals pre­
ceding the marginals are smaller.
The ampulla (a) is long, winding. At its ectal end the male duct enters
the prostate ( q ) directly, and the inner oviduct enters the female gland mass
(g). The inner massive portion of the prostate consists of small cells.
The middle part is also massive, but composed of large cells; the outer­
most section is tubular. The sheathed muscular, efferent duct ( d ) ends
with a short penial papilla.
Between the outer oviduct or nidamental duct (ov), that is the outlet
of the gland mass (g), and the penial papilla an uncommonly narrow
vagina (v) runs inwards and opens into the longish spermatheca ( b ).
Close to the vaginal entrance the insemination duct ( x o ) originates, so that
the arrangement of the seminal receptacles is serial. The considerable
width of the insemination duct is exceptional, and the same holds for the
position of the spermatocyst (y) quite near to the gland mass. A long
sausage-shaped gland ( vn ) with cuticular spines in its duct opens into the
female vestibule.
The specimens from the second locality were red alive; preserved they
are whitish with brown spots. Their length is 11 mm, the width 6 mm,
and the height 4 mm. The breadth of the sole is 2 mm. As far as the skin,
which is not well preserved, is recognizable, it resembles that of the original
description (Marcus, 1955: 147). The rhinophoral clubs have about 16
foliations on either side. The five to six gills are bipinnate, as frequently
those of small doridids of a species are, whose larger specimens have a
greater number of branchiae and more ramified pinnules.
The labial rodlets are 50 high and 8 ^ in diameter. The radula is
1.6 mm in length, 1.17 mm in width, hence half the size of that of the
84
specimen from the first locality. In Brazilian D. pusae the corresponding
measurements are 2.3 and 1.5 mm. The number of radular rows is 23,
that of the hooks 15, and that of the marginal lamellae 10. The innermost
lateral tooth bears one or two inner denticles, absent in the specimen
from the first locality. In the original material there is sometimes one den­
ticle. The lamellae are smooth, as in the specimens from Brazil; while
they bear minute irregular denticles in the slug from the first locality.
This minor character observed in a single specimen cannot be considered
as taxonomically significant.
The reproductive organs combine characters of the animal from the
first locality with such from Sao Paulo. The section of the insemination
duct between spermatocyst and female gland mass is short as in the former,
and its width is the same as that of the vagina, as in the latter. Cuticular
spines in the duct of the vestibular gland are wanting as in the small
specimens of the original material (Marcus, 1955: 149). Both Floridian
materials have simple, not stratified, labial rodlets.
Remark .— The original material, of which we have about 20 specimens,
is united with the animal of April 1963 by the slugs of March 1965.

47. Discodoris aurila, spec. nov.

Figures 108-111
Material.— Canal Zone (Pacific side): Deale Beach (Ft. Kobbe Beach);
F. M. Bayer and R. W. Bayer, December 1962. Two specimens.
Syntypes .— USNM 576268.
Description .— The color in life is light brown, sprinkled with white, evi­
dently glandular, stipples, and a dark band in the midline. This band
consists of melanophores which lie deep in the connective tissue. Further
melanophores in higher layers of this tissue and in the epidermis form a
reticulum. There is a zone of larger, vaguely delimited dusky spots between
midline and margin, and small, quite black dots between this zone and
the middle. The clubs of the rhinophores and the gills are uniformly brown,
the rhinophoral shafts as light as the notum. The underside of the pre­
served animals is colorless, without melanophores.
The preserved animals are 20 and 16 mm in length, 10 and 10.5 mm
in width, and 6.5 mm in height. The foot is 5.5 mm broad. The back of
the smaller specimen is somewhat rubbed off. The notum is swollen in
the preserved specimens. It is beset with caryophyllidia, that is, tubercles
with an inflated tip and projecting spicules. The borders of the pockets
of the rhinophores and gills are smooth.
The tentacles are triangular lobes; the rhinophores have 18 foliations.
The six branchiae are multipinnate. The anterior border of the foot has a
transverse furrow, the upper lip a central notch. The two blood glands
are dark.
85
 The labial plates are two areas with irregular outlines, 230 ^ long. They
bear slightly moniliform rodlets 30-40 ^ in length, 5 /x in diameter. The
radula is 2 mm long, 1.8 mm broad; its formula is 24X47.0.47. All teeth
are hook-shaped, from the highest tooth in the center of the half-row
(230 fx) they decrease evenly to the innermost (80 and outermost
(70 fi ) teeth. The stomach lies free over the intestinal gland.

F i g u r e s 108-111. Discodoris aurila, sp. nov. 108, living animal, from color
photo; 109, labial armature and its elements; 110, radular teeth; 111, diagram
of reproductive organs.— F ig u r e 112. Platydoris angustipes Morch, living
animal, from color photo.

The ampulla ( a ) is long; its outlet divides immediately into male and
female ducts. The male duct enters the voluminous prostate ( q ) whose
nuclei are small in the inner, large in the outer part. Ectally to the
86
prostate the efferent duct ( d ) begins with a muscular bulb, narrows, and
projects into the vestibule with a minute papilla.
The female gland mass (g) is conspicuous, its nidamental duct ( o v )
is glandular to near its aperture. The vagina (v) courses inwards beside
the penial papilla. The vaginal entrance into the large spermatheca ( b )
and the origin of the insemination duct ( xo ) are neighboring but separated.
The insemination duct is long, looping; the duct of the small, silky
spermatocyst (y), enters it near its opening into the gland mass.
Discussion .— The range of most doridids is little known, because many
of them were described only once, and based upon very few, often a single
specimen. The two descriptions of the Antillean Discodoris notha, for
example (Bergh, 1877b: 530; 1904: 53), refer to a total of three specimens
collected in 1860 and 1899. A doridid of the almost unexplored opistho-
branch fauna of Panama requires comparison with many congeneric species,
so much the more, as the present collection reveals obvious relationships
between the Pacific side of the Canal Zone and the Caribbean. Therefore
we went through the descriptions of the 28 species from the Indo-Pacific
Ocean and the 18 from the Atlantic Ocean and the Mediterranean Sea.
The geographically least distant D. heathi Mac Farland (1905: 39; 1906:
118), whose southernmost record on the American Pacific coast is Laguna
Beach (33° 32' N ), is not related with aurila. Its 12-16 outermost radular
teeth are reduced to thin, concave plates. Though it is not considerably
bigger than aurila, it has 8-10 branchial plumes.
Among the Indo-Pacific species we found D. pallida Baba (1937: 305)
from Amakusa (32° 30' N) similar to aurila. It differs by its nine gills,
the radular formula 15 X 14-17.0.14-17, and the larger labial plates. Also
D. modesta Bergh (1877b: 534) from the Palau Islands, Western Caro­
lines, resembles aurila. Its rhinophores have 50 leaflets, the ends of the
labial rodlets are thickened and often fringed; the radular formula is
29X48.0.48. The Atlantic D. voniheringi MacFarland (1909: 73) from
Alagoas, Brazil (9° S) has bipinnate gills, and the labial armature consists
of a median and two lateral plates.

48. Taringa telopia disa, subsp. nov.

Figures 113-114
References (to telopia telopia). — Marcus, 1955: 152; 1958b: 57.
Material.— Florida: Biscayne Bay, MacArthur Causeway; under rocks.
R. A. Smith, February 1962. One specimen.
Holotype .—USNM 576282.
Further distribution (of telopia telopia). —Brazil, Cabo Frio (22° 57' S,
42° 01' W ); coast of Sao Paulo (original locality).
Description .— The preserved animal is 18 mm long, 10 mm broad, 5 mm
87
high, the sole 4 mm, the hyponotum about 3 mm in width. According to
the collector’s note the living animal was “brown with dark blackish brown
spots;” the preserved specimen is olivaceous above and spotted below.
The sole is colorless; some caryophyllidia are topped with black. As in
the Brazilian specimens the caryophyllidia are biggest on the sides of
the notum, smaller in the middle and on the borders.
The rhinophores which contain dark spots, the gills, and the anterior
border of the foot agree with those of telopia telopia; the tentacles are
somewhat irregular cones.
The labial cuticle is smooth. The formula of the radula is 42 X 6.53.0.53.6.
The innermost tooth has up to 6 tiny inner and 4 outer denticles, but these
numbers vary; it measures 34 jjl. The lateral teeth increase from 52 ^ the
second to 0.16 mm; the 4-5 outermost ones decrease.
The reproductive organs agree with those of the Brazilian material.
Discussion .— Examining further material from Brazil we found 45 lateral
teeth as a new maximal number, still much less than in the Floridian spe­
cimen (53). The latter is not quite as long as the biggest Brazilian slugs
(preserved 22 mm), but has more radular rows (42) than telopia telopia
(32). These two characters lead us to separate the animal from Florida
as a subspecies. The more numerous denticles of the innermost tooth in
telopia disa are less important, because this number varies within the only
available radula. Also the slight difference of six, compared to 4-5, marginal
teeth is insignificant.
Since the type of Taringa was described, Swennen (1961: 64) found
T. armata in the Eastern Mediterranean. Living slugs of this species were
24 mm in length against 30 mm in telopia. In mature doridids the size is
a specific character. The numbers of the rhinophoral foliations and of the
lateral teeth in armata are distinctly inferior to those in telopia. The penial
papilla of armata is as long as broad, while it is nearly twice as long in
telopia telopia and telopia disa. Swennen mentioned the triangular, flattened
tentacles in telopia and the finger-shaped ones in armata. These differences
may be produced by contraction of the tentacular muscles. The variability
of the tentacles is attested by Bergh’s description (1894: 197) of them in
a preserved specimen of A porodoris millegrana (Alder & Hancock, 1855)
as “abgeplattet fingerformig.”
All doridids have a labial cuticle (Bergs, 1892c: 114). Our indication
for the genus, “lips unarmed” (Marcus, 1955: 151) which corresponds to
Odhner’s “Lippen unbewaffnet” (1926: 54), refers to the absence of
thickened plates with hooks or rodlets. In the description of T. telopia
this was detailed: “the cuticle of the oral cavity is colorless and smooth”
(1955: 154). Consequently the labial structures of telopia and armata are
the same. An amplification of the diagnosis of the genus is not needed;
such would not even be advisable with respect to the highly significant
labial structure.
88
 49. Taringa aivica, spec. nov.
Figures 115-119
Material.— Canal Zone (Pacific side): Deale Beach (Ft. Kobbe Beach).
F. M. Bayer and R. W. Bayer. December 1962. One specimen.
Holotype .— USNM 576267.
Description .— The notum is beset with many minute caryophyllidia and
a few long, conical papillae, whose bases bear the same small caryo­
phyllidia as the remaining surface. The spicules are dissolved.
The color alive is yellowish, mottled with grey; the conical papillae
are darker yellow. The rhinophores and gills are brownish. The preserved
specimen is olivaceous with small brownish-black dots. These melano-
phores are especially numerous around the bases of the conical warts. The
hyponotum and the sole have no pigment and are smooth. The inner
organs, especially the intestinal gland, are pinkish.
Measured over the back the length is 22 mm; the breadth is 15 mm,
the height 7 mm. The width of the hyponotum is 4 mm. The rim of the
rhinophoral sheath is undulate, the club bears 16 foliations. The tentacles
are flat, triangular, 0.8 mm long. The foot is bilabiate, its upper lip notched.
There are six tripinnate gills.
The labial cuticle is smooth. The radula is 3 mm long, 2.8 mm broad,
and comprises 32 rows with 48 teeth per half-row. The two or three outer­
most teeth are flat, leaf-like, their borders pectinate. All other teeth have
a row of 4-12, generally eight, pointed denticles on the outer side of
their cusp (Fig. 117). Also the innermost tooth has denticles only on
the outer side. The height of the innermost lateral tooth is 95 fi, it
increases outwards to about the 40th tooth, whose base is 0.2 mm, the
cusp 0.16 mm in length. The approximately five following denticulated
teeth decrease gradually; the marginal pectinate ones are still smaller. The
salivary glands are long, the two blood glands dark; the stomach is free.
No right liver was seen; it is very small in the type species.
The ampulla (a) is long, coiled, coursing under the anterior genital
complex. The spermatic duct and oviduct originate at the outlet of the
ampulla. The male duct immediately enters the massive, broad prostate
( q ) consisting of small acini. A lobe of the mucus gland touches the
prostate. From the right end of the prostate the narrow and relatively
short efferent duct ( d ) runs to the penis, a conical papilla ( p ) located in
the globular male atrium. The papilla is coated with a thick, smooth
cuticle; the wall of the atrium is beset with big conical papillae with a
thick, smooth cuticle over peculiar glands (Fig. 119).
An inconspicuous albumen gland (an) lies on the mucus gland (mx),
whose glands extend along the nidamental duct (ov) to near its opening.
Between this aperture and that of the male atrium the vagina (v) begins
with a muscular bulb containing cuticularized epithelial folds. Farther
89
F i g u r e s 113-114. Taringa telopia disa, ssp . n o v . 113, p r e s e r v e d s p e c im e n ;
114, m a le c o p u la to r y o r g a n .— F i g u r e s 115-119. Taringa aivica, sp . n o v . 115,
liv in g a n im a l, f r o m c o lo r p h o to ; 116, sk in ; 117, r a d u la r te e th ; 118, d ia g r a m o f
r e p r o d u c t iv e o r g a n s; 119, s e c tio n o f c u t ic u la r iz e d a tria l p a p illa .

90
entally the vagina narrows and enters the spermatheca ( b ). Beside this
entrance, but with an independent origin, the narrow insemination duct
( x o ) leaves the spermatheca. This duct forms two loops and is connected
with the large, sausage-shaped spermatocyst (y) by a short canal before
it enters the gland mass. The entrance of the insemination duct is far
distant from that of the inner oviduct.
Discussion. — Since the introduction of Thordisa Bergh (1877b: 540), the
diagnosis of that genus has been reduced too much (Bergh, 1892c: 132-
133). The characters of the type species must be considered, in order to
define Thordisa : notum villous, labial cuticle smooth, lateral radular teeth
without denticles, marginal teeth pectinate, penis unarmed. The dorsal
sculpture of densely set soft villous papillae should not be considered in a
restricted sense, and Th. diuda Marcus (1955: Fig. 136) with conical
tubercles, longer than broad and of different length, can be retained in
the genus. The tentacles, pointed conical knobs in the living animal (Fig.
134), round buttons in preserved specimens (Fig. 137, /), are difficult
to judge.
E tidoris Ihering (1886: 234) has sparsely set long tubercles mixed with
smaller ones, hence is not villous (Marcus, 1958a: 25). More important
than this difference from Thordisa is the glandular section of the efferent
duct in E. ladislavii, which contrasts with the massive prostate of Thordisa.
Species whose marginal radular teeth are not pectinate, e. g., Th. clan-
destina Bergh (1884a: 106) and Th. maculosa Bergh (1905a: 123), must
be excluded from Thordisa.
For species with denticles on the outer side of the lateral teeth we adopt
A porodoris Ihering (1886: 238). This character was stressed by Eliot
(1910: 106-107); it was not seen in the type species by Alder & Hancock
(1855: 42, D oris m illegrana) . Even Thiele (1931: 438) left the denti-
culation unmentioned. Thordisa dubia Bergh (1894: 178) is identical with
A porodoris millegrana. Bergh’s description of a voluminous prostate and
an unarmed penis completes Eliot’s. Further species of A porodoris are
Thordisa souriei Pruvot-Fol (1953: 66) and Thordisa hilaris Bergh, 1905,
variety Risbec (1953: 55), which we name A porodoris risbeci, nom. nov.
A porodoris rubra Bergh (1905a: 94), whose cusps of the lateral teeth
are smooth, does not belong to A porodoris. The cones in its efferent duct
suggest a Baptodoris, but the type-species of this genus has bipinnate gills
(Bergh, 1884b: 671), while rubra seems to have unipinnate ones.
Taringa Marcus (1955: 151) is separated from the preceding genera by
an armed penial papilla; the radula is as in Aporodoris. The three species
differ by their dorsal sculpture and the shape of the male organ. The
typical T. telopia, telopia disa, and T. armata Swennen (1961: 64) have
only caryophyllidia, while these are mixed with longer tubercles in aivica.
In the Atlantic and the Mediterranean species the penis is approximately
91
bell-shaped, in the Pacific one conical. The innermost tooth of the latter
has no inner denticles.

50. Aphelodoris antillensis Bergh, 1879

Figures 120-121
References.—Bergh, 1879c: 107; 1880c: PI. 16, Figs. 12-18; Marcus,
1963: 32.
Material.— Florida: 1/2 mile SSW of Alligator Reef Light, Monroe
County; at ledge, 4.5-6 m. W. A. Starck and H. A. Feddern, April 30,
1961 (CRR-F-296). One specimen.
Further distribution .— Virgin Islands, St. Thomas
(original locality);
Puerto Rico (kodachrome by Mrs. Germaine L. Warmke, identified by
the present authors); Curasao.
Description .— The preserved specimen is 10 mm in length, 5.5 mm in
width and 5.5 mm in height. It is white with blotches of ramified mela­
nophores, which leave white circles free around glandular openings. The
rhinophoral leaves are dark. The gills are dotted with black. The sides
of the body and the tail have scattered black spots; the sole is white.
 The border of the notum is thin, the tail pointed. The foot is rounded
in front, entire, not bilabiate. The tentacles are grooved on their outer
side. The sheaths of the rhinophores are high, their borders smooth. The
retracted clubs bear 10 foliations on either side of the rhachis. The five
gills are tripinnate.
The labial cuticle is smooth. The formula of the radula is 38X67.0.67.
All teeth, including the innermost, are hook-shaped and have no denticles,
as in the specimen from Curasao (Marcus, 1963: Fig. 42). The salivary
glands are very long.
The reproductive organs were described by Bergh (1879c: 109, 112-
113) but not figured. The genital aperture (Fig. 120, c) lies far behind.
The curved, compact ampulla (a) divides into the inner oviduct and the
spermatic duct at its exit. The incipient male duct which is not quite
short enters the acinous prostate ( q ). The following inner section of the
efferent duct ( d ) courses in coils; the middle portion is coated with a thick
layer of annular muscles. The outermost narrow part winds within a muscu­
lar sheath, and opens on the truncate tip of the penial papilla (p). This
cylindrical organ is lodged in a globular atrium lined with a thin, wrinkled
cuticle. The outlet of the male atrium is long, narrow, and richly provided
with longitudinal muscle fibers.
The vagina (v) begins beside the male opening. Its outer, cuticularized
section has strong annular muscles. The cuticle is continued into the fol­
lowing dilated ampulla-like part. Also the innermost portion of the vagina,
that opens into the spermatheca ( b ), is wide, contrasting with the much
narrower insemination duct ( x o ). The latter leads the sperm to and recei­
ves them from the spermatocyst (y) by a long canal. From the point where
canal and duct meet, a rather long stretch of the insemination duct con­
veys the sperms to the female gland mass (g), near the entrance of the
inner oviduct into the mass.
Discussion .— The color pattern of the typical material is not known. The
above-mentioned color photograph of two living slugs shows that the color
of the present specimen is well preserved. Therefore the black bands and
stripes of the animal from Curasao (Marcus, 1963: Fig. 40) and the
stippled blotches of those from Florida and Puerto Rico are intra-specific
variations.
51. Platydoris angustipes (Morch, 1863)
Figure 112
Synonym.— Platydoris rubra White, 1952: 118.
References.— Bergh, 1877b: 503; Marcus, 1957a: 422.
Material.— 1. Florida: vicinity of Pacific Reef. R. C. Work, August 1962.
Two specimens.— 2. Florida: 1/2 mile SSW of Alligator Reef Light,
Monroe County; at ledge, 4.5-6 m. W. A. Starck and H. A. Feddern,
April 30, 1961 (CRR-F-296). Two specimens.
93
Further distribution. — Dry Tortugas; Jamaica, Montego Bay (Haas, 1920:
142); Virgin Islands, St. Thomas (original locality); Brazil, Bahia.
R em arks .— The first description and drawing (Fig. 112) of a living animal
are made possible by color photographs of the slugs from locality 1. The
bigger animal is orange red, the smaller one vermilion. The former is
slightly mottled and the latter has darker blotches. The marginal zone of
the mantle is darkened by diffuse, fine pigment granules. An opaque white
line along the edge of the notum emits short white triangles inward. The
rhinophoral clubs are as dark as the margin but frosted with white in
the smaller specimen. Also four irregular white areas appear on either side
of the smaller specimen, and scattered white dots between them. The gills
of both slugs are ivory, richly frosted with opaque white.
The biggest of the preserved specimens (locality 1) is 40 mm long,
28 mm broad, 9 mm high. The hyponotum is up to 12 mm broad. The
foot is curled, narrow, 23 mm in length, 4 mm in width. The tentacles
are grooved laterally. The edge of the rhinophoral pocket is crenulated;
the club has 20 leaves in the smaller, (25 mm long) slug. The outline of
the branchial cup is stellate; there are six tripinnate gills and two black
blood glands.
The specimens from locality 2 measure in length 30 and 15 mm, in
breadth 22 and 9 mm.
The cuticle of the labial disc is smooth; the radula is as previously
drawn (Marcus, 1957a: Fig. 84), with smooth cusps of the marginal
teeth, in contrast with P. argo (Linne, 1767) from the Eastern Atlantic
and the Mediterranean Sea.

52. D oridella obscura Verrill, 1870

Figure 122
Synonym .— Corambella baratariae Harry, 1953: 1; Marcus, 1959b: 256.

References.— Verrill, 1870: 405; Franz, 1967: 73 ff.

M aterial. — Florida: Biscayne Bay, on floating Sargassum thrown upon the

beach of Virginia Key. Louise Bush, April 1964. One specimen.
Further distribution. — Massachusetts; New Jersey; Virginia; North Caro­
lina; Gulf coast from Mississippi, Louisiana and Texas.
R em arks. — The synonymy adopted here was established by Franz (1967).
As shown in a color photograph, the living animal is transparent with a
white network in the notum (p. 79) and over the dark liver. It forms a
radial pattern and surrounds the dorsal black spots mentioned in Harry’s
description (1953, Fig. 1), which are much more numerous, as also in the
Texan material (Marcus, 1959b: Fig. 12). The rhinophores are colorless.
94
 53. D endrodoris krebsii (Morch, 1863)
Synonym s.— Doriopsis krebsii (Morch), var. pallida Bergh, 1879b: 44;
D. atropos Bergh, 1879b: 49.
References. — Morch, 1863: 34; Bergh, 1875c: 87; Marcus, 1957a: 443;
1963: 35; 1967b: 203; Collier & Farmer, 1964: 389.
M aterial. — 1. Florida: Biscayne Bay. Southwest Point, Key Biscayne;
under rocks. R. C. Work, January 17, 1962.— 2. Same locality. Galen
Perry, February 25, 1962.— 3. Biscayne Bay, Bear Cut, Key Biscayne.
G. L. Voss, June 20, 1959.— 4. Biscayne Bay, among rocks, County
(MacArthur) Causeway, Miami Beach. R. C. Work, December 27, 1963.
Kept in the Laboratory to March 24, when it died and was preserved.—
5. Virginia Key, from red rocks (riddled with Cliona) off east side (Sewage
Beach). F. M. Bayer and H. B. Owre, May 1963.— 6. Bahamas: Exuma
chain. Sand beach on Joe Cay at Cut between Joe and Thomas Cays.
Feddern, Staiger, Devany and Pierce, August 24, 1963 (CRR-BW I-53).—
7. Los Roques Islands (E of Bonaire): Dos Mosquises; south end of
southern island. Rocky bottom with A cropora, depth 4-5 ft. R.C. Work,
August 2, 1963 (D. de S. no. 450). (A total of 10 specimens.)
Further distribution.— Virgin Islands, St. Thomas and St. Croix (original
localities); St. Kitts; Curasao; Brazil, coast of Rio de Janeiro and Sao
Paulo, from Cabo Frio (22° 57' S, 42° 01' W) to Cananeia, 25° S. East
Pacific, Gulf of California.
Rem arks. — The color photo of the specimen from locality 1 shows a slate-
black notum, a beige frilled margin, and blackish-brown blotches and spots
in the transitional zone. The wrinkled skin bears puckered warts of irregular
outlines, size, and distance from one another; the biggest warts are 5 mm
in diameter. Most of them stand out paler than the dark ground color,
some are uniformly slate-grey, others whitish and frequently capped with
a dark brown spot. The tips of the rhinophores are light, the club and the
shaft dark. The efferent side of the gills is light, the inner side dark. Accord­
ing to the collector’s note the animal from locality 4 had the same color,
that from locality 7 when alive was mottled dark grey and black.
Referring to the specimens of locality 5, Prof. Bayer observed: “color
evidently variable; some examples dark grey, often blotched with black,
or almost totally black. Red ones become dark after a time in the labo­
ratory.” The two color photos of this material show a purplish red and a
brown slug. The color of the former is brightest in the anterior half. The
finely frilled margin of this animal is light, the back rather smooth with
scattered low warts. Where the skin is scraped off, the subcutaneous tissue
appears white. The rhinophoral clubs are nearly black, their small tips
white. The same holds for the transparent gills. The brown slug has yellow­
ish brown areas and small black irregular blotches; its rhinophores are
95
black, topped with white. The brownish gills have black afferent vessels
and many white tips. The anal papilla contains orange faecal matter.
The preserved specimens from locality 2 are black with a yellow border;
also their inner organs are nearly black.
The slug from locality 1, whose color photo is accompanied by a scale,
was evidently creeping and stretched, as its length, 17 cm, exceeds the
hitherto known maximum (12 cm), while the width, 5.5 cm, is inferior
to such (7 cm). The consistency and the sculpture, gelatinous, swollen
and nodose, correspond to Bergh’s description of the original material
“aufgeblasen-knotig” (1875c: 87). The sculpture described as “soft,
blisterlike pustules” in other species of Dendrodoris occurs in the present
material too (locality 5).
The number of the rhinophoral foliations varies in the Antillean krebsii
and its Brazilian synonym atropos from about 20 to 40; there are nearly
always six gills in adult animals, but also eight occur (Collier & Farmer,
1964: 389). The salivary glands are rather big as in our first figure
(Marcus, 1957a: Fig. 149, q ) . In the central nervous system of one
specimen from locality 5 the territories of the ganglia are delimited by
shallow furrows. The stained and clarified nerve ring shows these areas also
defined by the different size of their nuclei. The smallest are those of the
nearly globular rhinophoral bulbs which are contiguous in the midline, the
biggest are in the visceral region as often is the case in opisthobranchs
(Hoffmann, 1936: 863). Thus the contiguous but individualized subintes-
tinal and supra-intestinal centers become visible on either side of the
midline behind the cerebro-pleural masses. On the right side, over the
supra-intestinal ganglion, the abdominal center is marked, as Bergh stated
(1875c: 88), by a huge cell, 0.13 mm in diameter with a nucleus of
65 /a. Also the pedal ganglia contain some giant nuclei; those of the
cerebro-pleural ganglia are medium-sized.
The length of the hook-bearing portion of the male organ, 4.7 mm in
a specimen from locality 2, exceeds what we have previously observed,
but not the maximum known, 6 mm (Bergh, 1879b: 63).
The only other West Indian D endrodoris, D. subpellucida (Abraham,
1877: 265-266, PI. 30, Fig. 36), cannot be recognized, unless new material
from its original locality, St. Vincent, proves to be beyond the intra­
specific range of variation of D. krebsii.

54. Doriopsilla janaina, spec. nov.

Figures 124-125
M aterial .— 1. Canal Zone, Panama (Pacific side): Perico Island, Ft.
Amador, under rocks at low water. F. M. Bayer and R. W. Bayer,
December 25, 1962. One specimen.— 2. Canal Zone. Panama (Pacific
side): Venado Beach, Ft. Kobbe; in rocky tide pools on point opposite
96
Venado Island. F. M. Bayer, R. W. Bayer and R. H. Chesher. December
31, 1963. Four specimens.
H olotype .— USNM 576269.

Description .— The slug from the first locality is 25 mm long and 14 mm

broad according to the color photo. In preserved condition it is 22 mm
long, 13 mm broad, and 5 mm high, broader in front than behind. The
largest of the preserved specimens from the second locality is 21 mm in
length, 11 mm in width, and 5 mm in height. The notum is beset with
small tubercles, less distant from one another than their maximum breadth,

122. Doridella obscura Verrill, living animal from color photo.—

F ig u r e
123. Phyllidiopsis papilligera Bergh, dorsal view of preserved speci­
F ig u r e
men.— F ig u r e s 124-125. Doriopsilla janaina, sp. nov. 124, living animal,
from color photo; 125, diagram of reproductive organs.

0.5 mm. The skin is leathery; the notum contains large spicules, up to 1
mm in length and 0.1 mm in diameter. The hyponotum is 2.5 mm broad,
reticulated, with dense white strands and transparent interspaces.
The color alive, from the photo, is light red. Two blackish bands, which
have a common origin between the rhinophores, end under the branchial
97
tuft, and leave the tubercles free. Minute white dots on the blunt tips of the
latter are cutaneous glands. Around the base of every tubercle, except the
quite marginal ones, there are about five black spots, evidently single
melanophores. There are two pairs of bigger white tubercles and many
small, dark red spots, the latter especially in the middle. The rhinophores
are dark red with small white tips, and white dots occur on the rim of the
rhinophoral pits. The black pigment is retained after preservation; the
white glandular secretion and the red color have disappeared. The absence
of black pigment in the inner organs contrasts with several other dendro-
doridids.
The tentacles are quite short, the rhinophores have around 18 leaflets on
either side; the sheaths of the rhinophores are high, with smooth rims and
small tubercles on the outer side. The anterior pedal border is bilabiate; the
sole is 7 mm in breadth. There are five multipinnate gills; the anal papilla
lies to the left, between the anterior and posterior plumes.
The central nervous system is smooth when examined from below.
A constriction shows between the pedal ganglia, and apposed to their
hinder surface lie the buccal ganglia, far from the oral vestibule. The latter
is gradually dilated into a pharynx, whose anterior part is invaginated into
the vestibule, as in our figure of D. areolata Bergh (Marcus, 1962b: Fig.
18). Two retractors insert at the point of this invagination.
The ampulla ( a ) is longish; the spermoviduct enters the female gland
mass ( g ) where the oviduct and the spermiduct separate. The latter
emerges from the mass as a short duct and forms the prostate ( q ), the
uppermost organ of the anterior genital mass. The prostate consists of
branched diverticula. The male duct ( d ) passes through it and is in its
farther ectal course sheathed by muscles. Before the efferent duct enters
the male atrium, it becomes spinous ( p ). The spines are numerous and
pointed, about 50 ^ in length.
Beside the male aperture lie the openings of the vagina (v) and the
oviduct. The terminal part of the vagina is widened and heavily ciliated.
The vagina runs to a spherical spermatheca (b). A long duct connects the
spermatheca and the spermatocyst ( y ). This duct leaves the spermatheca
at the entrance of the vagina, and the insemination duct ( xo ) starts from
the same point, coursing to the gland mass, which it enters near the
emerging male duct.
The egg ribbon is about 2 mm broad and forms a spiral of a little more
than three coils, 11 mm in diameter; unrolled it would be 5 cm long.
The diameter of the eggs is 0.14 mm, the length of the capsule 0.23 mm.
There are about 200 eggs per mm, hence a total of ca. 10,000 in the
present ribbon.
Discussion.— Doriopsilla albopunctata (Cooper, 1863: 58), known from
Monterey to Point Eugenia, 27° 51' N (Lance, 1961: 67), is related to
the new species by the white-tipped notal tubercles and the lobed prostate.
98
The latter character was described in Doriopsis fulva MacFarland (1905:
45; 1906: 131), now united with albopunctata (Steinberg, 1961: 57 ff.).
Steinberg describes the color variations of albopunctata as bright yellow,
orange, and warm brown. Cockerell & Eliot (1905: 41) found a deep
chestnut colored living animal of their D oriopsis reticulata which is synony-
mized with albopunctata too. In reticulata 30 rhinophoral leaflets were
indicated, but if this species is identical with fulva, which has 15-20 leaflets
on either side of the rhachis (Marcus, 1961a: 30), Cockerell & Eliot must
have summed them from both sides.
Red and pink tones and blackish bands are not mentioned in the descrip­
tions referring to albopunctata and its synonyms. The branchiae are
yellowish white in living albopunctata , as the descriptions of fulva and
reticulata show, and the rhinophores are dark yellow.
The position of the buccal ganglia apposed to the pedal ganglia, and the
absence of a ptyaline gland (Marcus, 1962b: Fig. 18) distinguish D oriop-
silla Bergh, 1880, from D endrodoris Ehrenberg, 1831 (ibid.: Fig. 19).
The location of the anus to the left of the gills cannot be considered
as a generic character of D oriopsilla as compared with a median position
in Dendrodoris. The records referring to the type-species of Doriopsilla do
not agree in this respect, as Bergh (1880b: 319) found the anus symmet­
rical between the two posterior gills of a total of four, Pruvot-Fol (1954:
337) and we (Marcus, 1962b: 472) to the left of five and four gills re­
spectively. MacFarland described the anal papilla of Doriopsilla fulva (now
called albopunctata ) at the right of the center of the circle of branchiae
(1906: 131); we found the anus surrounded by the gills in the same species.

55. Phyllidiopsis (Phyllidiopsis) papilligera Bergh, 1890

Figure 123
References.— Bergh, 1890a: 176; Marcus, 1962b: 475.
M aterial.— Jamaica, East Palisadoes, 25-30 m. T. F. Goreau, by diving,
December 20, 1964. One specimen.
Further distribution. — Gulf of Mexico, 185 m (original locality); Virgin
Islands, St. John, 18 m; Grand Bahama Island; Jamaica (letter from Dr.
Malcolm Edmunds, of Accra, Ghana, September 8, 1964).
Rem arks. — At all mentioned localities only single specimens were gathered.
The present one is big, 22.5 mm in length, 15 mm in width, very well
preserved, and has a distinct symmetrical color pattern. There are numerous
white papillae.
56. Tritonia (Tritonidoxa) wellsi Marcus, 1961
Figures 126-130
R eference .— Marcus, 1961b: 146.
M aterial. — 1. Beaufort, N.C. Shark Shoal Jetty, on Leptogorgia virgulata.
99
W. K. Patton, June 7, 1961. Three specimens.— 2. Brazil: Sao Sebastiao,
23° 50' S, 45° 20' W, on gorgonians. Jorge Petersen, March 1962. Three
specimens.— 3. Near Ubatuba, 23° 27' S, 45° 06' W. Liliana Forneris,
August 1963. One young specimen.

F ig u r e s 126-130. Tritonia (Tritonidoxa) wellsi Marcus. 126, dorsal view of

living slug; 127, jaw; 128, radular teeth of a young slug; numbers indicate rows;
129, masticatory border of jaw; 130, half-row of adult radula; numbers indicate
position in row.
100
Description. — The adults measure up to 15 mm in length when living.
They are transparent dark pink with whitish gills. The back and the sides
are smooth; the hind end is long, tapering to a pointed tip. The veil is
entire, not bilobed; the outermost of its 6 appendages are the spoon-shaped
tentacles. These rather long appendages and the produced foot corners give
the present living slugs a different aspect from the preserved specimens
(1961b: Fig. 22). The sheaths of the rhinophores are high, their rims
smooth with a point on the outer side. There are 9-10 branchiae, richly
ramified dichotomously. The genital aperture is wide; it lies under the 2nd
right gill, the anus below and a little in front of the fourth.
The masticatory border of the jaw bears up to 10 rows of denticles. The
radular formula is 20X10.1.1.1.10. The rhachidian tooth is tricuspid,
flanked by the inner lateral or intermediate teeth. These are concave on the
inner side of the cusp, with about 10 denticles on the outer side. The
innermost denticle is stronger than the following ones. The radula of the
young slug was 0.47 mm long, 0.1 mm broad, with 30 rows, about 20 of
which are preradular, the rest juvenile (Fig. 128). The first row has only
the rhachidian tooth, 10 ^ high and 5 V broad. Its median cusp is simple,
but in the following rows more differentiated than in the definitive ones.
Rows 2-7 have 1.1.1 teeth, rows 8 and 9 have 2.1.2, rows 10 and 11 have
3.1.3, and rows 12-17 have 4.1.4 teeth. The intermediate tooth appears
in row 18. The juvenile rows of the radula have only five lateral teeth,
the innermost of which bears outer denticles, absent in the corresponding
adult tooth.
The reproductive organs are as in our first description; the spermatheca
(loc. cit., Fig. 24, s) is more elongated than in the sections.
Discussion.— The range of Atlantic warm-water species from Cape Hatteras
to southern middle Brazil is not exceptional.
In subdividing Tritonia we isolate Tritonidoxa by its flagelliform penis
(Odhner, 1926: 35). If the shape of the velum is applied (Odhner, 1963:
51), the type-species and two more out of a total of four species (Marcus,
1961b: 148) are not covered by the key. As to the genus M yrella Odhner,
1963, we remain skeptic (Marcus, 1959a: 65), and suppose that the notal
“pores” actually are openings of cutaneous glands.

57. Tritonia (Candiella) bayeri, spec. nov.

Figures 131-134
M aterial. — 1. Florida: Elliott Key; Margot Fish Shoal (reef), about 2 mi.
offshore at middle of island. R. C. Work, June 17, 1963. One specimen.
— 2. Biscayne Bay; Bear Cut, off north point of Key Biscayne. Sam Stout
and J. J. McClelland, July 9, 1964. Five specimens, living on Briareum
asbestinum (Pallas), feeding after the manner of Cyphom a. — 3. Biscayne
101
Bay, Soldier Key; on Pseudopterogorgia. September 30, 1964. One speci­
men.
H olotype — USNM 576276.
D escription .— The length of the preserved animals is 4.5-11 mm; the
biggest is 6 mm broad with gills, 4 mm without them. Its height is 3 mm,
the breadth of the sole 2 mm. The living slugs are diaphanous with an
opaque white diffuse network on the notum (Fig. 131), not regular as in
T. reticulata Bergh (1882: 239). The branchiae are white, the velar
processes and those of the rhinophoral sheath are peppered with white. The
pinkish (locality 1) or dark (2) intestine shines through the notum.
The back is smooth; the foot is rounded in front, its corners are not
produced, and the hind end tapers. The cephalic veil is entire with a total
of four digitate appendages, which are triangular lobes in the preserved
animals. The two outermost are the tentacles characterized by ventral
grooves. The rim of the rhinophoral sheath is smooth, but bears a long
process (x) directed obliquely outwards, which is similar to the appendages
of the veil. The clubs of the rhinophores have pinnate plumes as in the
other tritoniids. There are up to 15 branchial tufts on either side of the
notum; larger and smaller ones alternate more or less regularly. The gills
dichotomize up to five times, when they have 32 tips. The genital apertures
lie in the interspace between the second and third branchial tufts, the anus
between fifth and sixth tufts, approximately in the middle of the body.
The renal pore lies a little dorsal to and in front of the anus.
The masticatory border of the jaws is set with several rows of conic
teeth with a longitudinal sculpture. The teeth are smaller in front, large
behind. The radula has 34 rows; it is horn-yellow, the rhachidian tooth
brown. The latter is tricuspid, the pointed median cusp is a little longer
than the rounded lateral cusps. The half-row has 11 lateral teeth. The
innermost lateral (intermediate) tooth has an inner concavity and about
15 thin outer denticles. The second lateral tooth is hook-shaped as the
following teeth, but broader and with 4-5 denticles between the cusp and
the base, located on the underside, hence difficult to see. The third lateral
tooth is the smallest of the half-row, its cusp is smooth as in the following
teeth. These increase in size, only the outermost tooth is smaller. There
are no gastric plates.
The pink ovotestis (oi) is located dorsal to the intestinal gland. The
hermaphrodite duct (ef) forms a globular ampulla (a), whose outlet divides
into sperm duct and oviduct (ov). A quite short muscular section connects
the ampulla and the long, winding prostatic section ( q ). This continues to
the male atrium containing the short, conic penis ( p ). The inner section of
the oviduct (ov) between ampulla and female gland mass (g) is long. The
outlet of the gland mass, the nidamental duct, is wide and short, and opens
into the female atrium (ai). From the dorsal wall of the atrium arises the
vagina (v) or bursa canal, about as long as the vesicle of the bursa ( b ).
102
In the evaginated phase the male opening ( ma) lies in front of the female
apertures, of which the vaginal pore is dorsal to the nidamental orifice.
The species is named for Professor Dr. Frederick M. Bayer.
Discussion .— The digitiform process on the smooth rim of the rhinophoral
sheath is the most important character of T. bayeri. In tritoniids with jagged
rims one of the lobules or points may project beyond the others (Bergh,
1907: 85, T. indecora; Vayssiere, 1912: Fig. 81, Tritoniopsis elegans,
according to Odhner, 1936: 1080), but these structures differ widely from
that in the present species. A further peculiar feature of T. bayeri is the
denticulation of the second inner lateral tooth. In Tritoniopsis, whose name
has to be maintained (Mattox, 1955: 12), and Paratritonia, denticulate

F i g u r e s 131-134. Tritonia (Candiella) bayeri, sp. nov. 131, living animal, from
color photo (locality 2 ), the white lines drawn black; 132, masticatory border
of jaw; 133, radular teeth; 134, diagram of reproductive organs.
103
lateral teeth occur (Vayssiere, 1912: 94-95; Baba, 1949: Fig. 103 B, b;
106 c), but the combination of a differentiated innermost lateral tooth with
a second one distinguished from the remaining teeth is a novelty.
Only four appendages of the veil, two outer tentacles and two inner
processes, as in T. bayeri, are rare, at least in adult tritoniids. During
growth these appendages as well as the branchial tufts increase in number
(Haefelfinger, 1963: 64; Tardy, 1963: 5). Whether they are short papillae,
pointed lobes, or finger-shaped, is systematically significant only when the
slugs are observed alive; otherwise, the lengths can be widely different. One
species with an undifferentiated innermost lateral tooth (Pruvot-Fol, 1937:
69) and with Duvaucelia-like openings of the intestine and reproductive
organs has four lobes. Four digitiform processes occur in Duvaucelia lineata
(Alder & Hancock, 1848: 191; 1851: Fam. 2, PI. 4 ), a true Duvaucelia.
With Odhner (1963: 51) we separate Duvaucelia and Tritonia as genera
by the position of the anus and the reproductive apertures. The species
classified correctly by Haefelfinger (1963) and Tardy (1963) show that
the distinction between Duvaucelia and Tritonia is sound: Haefelfinger’s
three species belong to Tritonia, and that of Tardy to Duvaucelia. It is
true that Tardy in his comparative notes does not assign the species called
Candiella villafranca by Vayssiere (1901b: 101) to the correct genus, as
it belongs to Tritonia. Furthermore, Duvaucelia insulae Baba (1955: 25,
50) is a Tritonia .
Tritonia bayeri belongs to the subgenus Candiella Gray, 1850, due to
its conic male copulatory organ and the entire rather than bilobed veil
with a small number of appendages.
There are no tritoniids from Western Atlantic warm waters which
deserve a special comparison with the new species. T. palmeri Cooper,
1862 (Cockerell & Eliot, 1905: 33), from the warm temperate area of
California, and the more northern T. festiva (Stearns, 1873) (Marcus,
1961a: 31) do not have the mentioned features of the rhinophores, the
second lateral tooth, and the veil characteristic of bayeri. Also the male
copulatory organ of / estiva (not known of palm eri ) is different.

58. M arionia cucullata (Gould, 1852)

Synonym .— M arionia occidentalis Bergh, 1884a: 49.
References.— Gould, 1852: 308; 1856: PI. 25, Figs. 403 a-f; MacFarland
1909: 9; Odhner, 1926: 40; 1934: 294; Carcelles, 1944: 263.
M aterial. —Brazil: Sta. Catarina, 30 km N of Florianopolis, shallow water.
Jorge Petersen, November 11, 1962. Eight specimens.
Further distribution.— Rio de Janeiro, in the market, among shrimps
(original locality); off Rio de Janeiro, 73 m, sandy bottom; Argentinean
coast, between 35° and 38° 50' S, from intertidal rocks with M ytilus
to 146 m.
104
R em arks .— The present specimens preserved in formalin are up to 75 mm
long, measured over the back, 50 mm in linear extension. The breadth of
the large slugs is 18 mm, and their height is 15 mm. The general color of
the dorsal side is a glossy greyish green with a brick red net which faded
out in the course of a fortnight. The areas in the meshes of the net are
darker olivaceous low bosses. The hyponotum is rough, lighter than the
back; the sole is smooth and still lighter.
The border of the notum is frilled and bears 12-14 tufts of gills, often
different numbers on the two sides. The genital apertures lie under the
third, the anus under the fifth tuft. The anterior border of the foot is
bilabiate and notched in the middle. The front margin of the veil has about
8 processes on either side of a slight median incision, each composed of
3-4 tubercles. The tentacles are furrows in the sides of the veil; the rhino­
phores have long smooth sheaths and 7-8 longitudinal tripinnate crests.

59. Bornella calcarata Morch, 1863

Figure 135
References. — Bergh, 1874b: 289; Marcus, 1958a: 28.
M aterial.—Brazil: off Cabo Orange, 4° 46' N, 51° 21' W, 59 and 66 m,
bottom temp. 23° C. O r e g o n Stations 4207 and 4209, February 26 and
27, 1963. R. C. Work. Two and six specimens, respectively.
Further distribution .— Virgin Islands, St. Thomas (original locality);
Surinam (Nijssen-Meyer, 1965: 146); Brazil, State of Pernambuco, Recife.
R em arks .— Only few specimens of B. calcarata had hitherto been reported:
the type, re-examined by Bergh, and one from Brazil, and four from
Surinam. The present material is accompanied by color notes of the living
animal: ground color beige-orange-pink; broken red lines on the sides
becoming a reticulation on dorsum and lower part of dorsal appendages;
area around mouth orange red. The clubs of the rhinophores and the gills
are still red in the preserved slugs.
The biggest animal is 45 mm in length, 14 mm high, and 8 mm broad.
The breadth of the veil is 12 mm. The cerata of the rhinophores are 9 mm
long, those of the notum 6 mm. There are 7-8 processes on the coalesced
rhinophoral sheath and first ceras of the biggest as well as of the smallest
animal (30 mm). The number of the velar processes or tentacles beside
the mouth is different on the right and left side, viz. 13 and 11 in the
biggest animal, 8 and 12 in the smallest.
The second (II) to seventh (V II) right and left tufts of dorsal appen­
dages have the following number of cerata (c) and gills (g) in the biggest
and the (smallest) animal:
II right 4 c, 5 g (3 c, 5 g), left 3 c, 5 g (4 c, 6 g).
III right 4 c, 4 g (3 c, 4 g), left 2 c, 4 g (3 c, 4 g).
105
 IV right 2 c, 4 g (2 c, 5 g), left 2 c, 3 g (2 c, 4 g).
V right 3 c, 3 g (2 c, 2 g), left 2 c, 3 g (2 c, 3 g).
VI right 2 c, 2 g (1 c, 1 g), left 1 c, 3 g (1 c, 1 g).
VII right 2 c, 1g (1 c, 0 g), left 1 c, 2 g (1 c, 0 g).
The radula is remarkably small 2.2 mm long, 0.6 mm broad. Only the
rhachidian tooth is strong; the 14 laterals are delicate. The scaly cuticle
of the muscular labial pads, the thick jaws, whose shape Bergh (1874b:
294) compared with the shells of Isognomon (formerly Perna), and the
cuticular spines of the second stomach contrast with the weak radula.
The Bornellidae “are said to creep briskly about over the surface of
fucoids” (Bergh, 1884a: 35). Probably they feed upon the animals growing
on the algae. One cannot correlate the contents of the stomach of the
present slugs, masses of nematocysts, with the equipment and the subdivi­
sion of the gut. Vayssiere (1912: 99) had a similar opinion. As food of the
Bornellidae, nematocysts and hydroids (in one case Tubularia) are
mentioned, as well as copepods which we think were ingested casually.
Besides, unrecognizable animal masses are reported, and structures which
cannot be classified, though they were described in detail (Bergh, 1874b:
297) and figured (PI. 37, Figs. 4).

60. D oto caramella Marcus, 1957

Figures 136-137
R eference. —Marcus, 1957a: 458.
M aterial .— Brazil: coast of Sao Paulo, intertidal zone of Ilhabela. October
1956. Two specimens.
R em arks .— Material found at the original locality after the conclusion of
the first description requires some complementary notes. The present slugs
have the anterior border of the rhinophoral sheath slightly prolonged; this
character is used for the separation of species in D oto. The same holds for
the number of cerata. There are five pairs in the new animals of caramella,
whereas four were recorded in the original description. D. pita Marcus,
1955, described with four to five pairs, was later found with seven. D.
caramella shows the following generic characters of its alimentary system:
thin jaws, a globose left posterior oral gland (Fig. 136, ms ) , up to 0.16 mm
in diameter, much bigger than the right one, and a concentration of hepatic
tissue chiefly in the cerata (Odhner, 1936: 1065). Also the ducts (co)
which connect the main liver duct and the diverticula in the cerata contain
hepatic tissue in caramella. In D. pita the main duct consists entirely of
hepatic tissue. As is characteristic for the family Dotoidae (Marcus, 1961b:
148, M iesea), the gonad lies over the main liver duct, though the ovotestis
may extend so far to the sides that the duct becomes central.
The hermaphrodite gland of D oto caramella fills the entire body cavity.
106
Few voluminous masses of sperm lie in the gonadal center, big ovocytes
on the periphery. The gonad of D. pita consists of separate follicles, each
with central male and peripheral female germ-cells. The ampulla of
D. caramella (Fig. 137, a ) is fusiform, its outlet, the strongly ciliated
spermoviduct (z), bifurcates into a quite short inner oviduct and a male

F ig u r e 135. Bornella calcarata Morch, right side view, left appendages

stippled.— F i g u r e s 136-137. Doto caramella Marcus. 136, ramification of
digestive gland; 137, diagram of reproductive organs.
107
duct. The latter becomes prostatic ( q ) at once. The cells of the inner
prostate are big and stain blue; the finely granular plasma of the outer ones
is orange. A section formed by high cells connects the prostate and the
muscular part ( e ) of the efferent duct, a short, winding tube. The conical
penis ( p ) is IV2 times as long as thick.
A ciliated infundibular valve leads the eggs into the fertilization chamber
( be ) . The heads of the alien sperms in this chamber are all directed
inwards. Also in the following seminal receptacle (t ) the sperms are
parallel. From here the eggs pass through the vagina (v) and on to the
mucus and albumen gland. Hence the vagina functions as inner oviduct;
it is provided with a strong sphincter O x). The outer oviduct forms the
female gland mass ( g) . The fundamental structure of the female organs
in D. caramella is the same as in D. uva (Marcus, 1959a: Fig. 160),
D. divae (Marcus, 1960b: Fig. 57), and others (Marcus, 1961a: Figs.
129, 134, 138, 140, 146), while the inner oviduct of D. pita (Marcus,
1957a: Fig. 175) does not communicate with the single sperm-storing
vesicle.

61. Arm ina miilleri (v. Ihering, 1886)

Reference. — Marcus, 1960b: 170.
M aterial. — Brazil: 1. Coast of Sao Paulo, Ilha Anchieta. Liliana Forneris,
February 1964. Two specimens.— 2. Coast of Sao Paulo, off Cananeia.
Luiz Tommasi, August 1964. Three specimens.— 3. Coast of Sta. Catarina,
30 km N of Florianopolis; shallow water. Jorge Petersen, November 11,
1962. One specimen.
Further distribution. — Brazil, near Cabo Frio, 22° 57' S, 42° 01' W; Sta.
Catarina, Florianopolis (original locality).
R em arks .—By the finding of the species on the coast of Sao Paulo its
Brazilian range becomes nearly continuous.
Of the 9 slugs of this species now known, 3 were found at one locality,
on the continental coast opposite the Island of Sta. Catarina. The present
preserved specimen from there is 40 mm long, measured over the back,
22 mm broad, and about 12 mm high. Its color corresponds to that of the
original material; it has a black notum with light ridges, a yellowish
hyponotum, and a bright pink sole. Though the slug was kept in formalin,
the colors faded out considerably in the course of a month.
Nijssen-Meyer (1965: 147) classified a specimen from Surinam as
A . sem peri (Bergh, 1861) hitherto known from the Red Sea to Japan and
the Philippines. She compares her slug thoroughly with A . miilleri. It has
well-developed caruncles like miilleri , while these are minute in semperi.
The first lateral tooth of the radula has at least six inner denticles in the
specimen from Surinam, which distinguishes it from miilleri .
108
 62. Fiona pinnata (Eschscholtz, 1831)
References .— Marcus, 1961a: 50 (principal references); Bayer, 1963:
460 (biology).
M aterial .— Florida: Virginia Key, beach at Marine Laboratory; from
Velella drifting past the laboratory dock or stranded on the beach. G. Y.
Hendrix and Jon C. Staiger, March 1, 1963. F. M. Bayer, March 7, 1964.
Six specimens.
Further distribution. — Pelagic and gregarious in warm and temperate seas.
R em arks .— The branchial fold on the large cerata of about four of the
inner rows (Marcus, 1961a: Fig. 174) is the best character by which this
species can be recognized.
Bayer’s specimens fed upon Velella, avoiding its lower surface, and upon
Lepas anatifera. They did not accept L. pectinata, hydroids, and sea
anemones grown on Sargassum, nor Physalia.

Notes on the American species of Phidiana

In the species of Phidiana from the Atlantic coasts of America the
anterior region of the foot is distinctly widened. The dilatation present in
living animals is seen in d’Orbigny’s (1846: PI. 14, Figs. 4-6) and
Edmunds’ figures (1964: Fig. 11 A ). In preserved material a broadly
rounded, disc-like or crescent-shaped configuration results from contrac­
tion (Bergh, 1874a: PI. 9, Fig. 4; Engel, 1925: Fig. 4; Marcus, 1955:
Figs. 249, 252, f). For the Atlantic species of the genus, lynceus (Bergh,
1867), selencae (better selenkai ) Bergh 1879, and brevicauda Engel,
1925, the name lynceus has to be used (Engel, 1925: 67; Marcus, 1963:
47-48). It ranges from Florida, Jamaica, and the Lesser Antilles to
southern middle Brazil.
The name Ph. patagonica (d’Orbigny, 1837: 192) may possibly replace
lynceus. Ph. patagonica occurs from the coast of Patagonia, Bahia San
Bias, 40° 36' S, to Tierra del Fuego, Hoste Island, Orange Bay (Odhner,
1926: 25; Carcelles & Williamson, 1951: 318). The substitution of
lynceus by patagonica is not possible for the moment, because patagonica
is known only externally. Its antero-middorsal color marks are light red,
not white as in lynceus.
In order to facilitate a future comparison of new material of patagonica
with lynceus, we summarize the taxonomic characters of the latter: foot
dilated in front; anus generally located between anterior and posterior
group of the posterior or right hand left liver, but sometimes between the
branches of the anterior or of the posterior group. Masticatory border of
jaw with 14-30 denticles; radula containing 15-19 teeth with 4-6 denticles
beside and 2-6 denticles on the cusp on either side; penial stylet a hook­
shaped funnel, bent away from the male opening or towards it, or forward.
109
 The fore end of the foot is evenly rounded, not dilated or only slightly
so, in the species of Phidiana from the Pacific coasts of America. In his
comparison of inca with patagonica d’Orbigny (1837: 194) mentions this
difference unequivocally. We agree with Engel (1925: 63) that a part of
the body whose contraction results in a determined shape, e.g. the disc
recorded for lynceus, selenkai, and brevicauda (Marcus, 1957a: 470), can
be a specific character. The Pacific American species of Phidiana are: inca
(d’Orbigny, 1837: 193; 1846: PI. 13, Figs. 1-7), exigua Bergh (1898b:
559), and pugnax Lance (1962: 157). The range of inca extends from
southern Chile, 41° 50' S, to Peru, Callao, 12° S; the single specimen of
exigua was found together with inca at Coquimbo, 30° S, and pugnax is
recorded from Pacific Grove, 36° 38' N, to San Diego, 32° 42' N, and the
neighboring Los Coronados Islands, 32° 24' N.

F ig u r e s 138-140. Phidiana lynceus Bergh. 138, dorsal view of fore end o f

specimen from Pacific side of Canal Zone, from color photo; 139, ventral view
of same, preserved; 140, ventral view of preserved fore end from third locality.
F i g u r e 141. Godiva rubrolineata Edmunds, living animal, from color photo.

The Pacific species cannot be united under a common name. In Engel’s

key (1925: 65) referring to the radulae, inca and exigua are separated by
the absence of denticles on the cusp and the presence of 2-3 there, respec­
tively, as Bergh’s descriptions indicate (1898b: 554, 559). Further
material of inca however showed two denticles on either side of the cusp
(Marcus, 1959a: Figs. 185, 186). As the remaining characters considered
110
as systematically important agree in inca and exigua, we think that both
may be united.
Ph. pugnax is said to have six major groups of cerata, the first three of
which are easily distinguishable. If we understand the kodachrome of the
living slug correctly, the anterior liver of pugnax comprises two groups.
Such occur in Atlantic Phidiana too (Marcus, 1963: 46). Lance’s color
photo shows two further distinct groups on the right side, which belong
to the posterior (or left) liver. The anterior of these is evidently Lance’s
“third major group of cerata,” and within this group he found the anus.
In inca it lies behind this group; in lynceus both positions, that of pugnax
and that of inca, occur.
The radula of pugnax comprises 19 teeth ( inca 19-23; lynceus 15-19);
there are evidently 2-3 denticles beside the cusp and 4 on it ( inca 3-7
beside, up to 3 on the cusp; lynceus 4-6 beside, 2-6 on the cusp). The
number of denticles on the masticatory border, 14, is small for the big
slug, 37 mm long, of the description of pugnax. For lynceus 14-30 denticles
were recorded, for inca 20-23. The penial hook of pugnax offers no specific
characters. On the whole, Ph. pugnax, whose foot decidedly agrees with
the not-widened mca-type, shows some features more similar to lynceus ,
and cannot be identified with either of them.

63. Phidiana lynceus Bergh, 1867

Figures 138-140
References .— Marcus, 1963: 45; Edmunds, 1964: 16.
M aterial .— 1. Canal Zone (Pacific side) : Deale Beach (Ft. Kobbe Beach).
F. M. Bayer and R. W. Bayer, December 1962. Three specimens.— 2.
Florida: Biscayne Bay, Southwest Point, Biscayne Key. R. A. Smith,
October 31, 1962. Two specimens.— 3. Florida: Biscayne Bay, Bear Cut;
Thalassia flats. King Tsue, March 12, 1964. One specimen.—4. Florida:
Virginia Key, beach facing Bear Cut, Biscayne Bay. J. Ewald, May 11,
1960. One specimen.
Further distribution. — Jamaica; Lesser Antilles; Brazil, Rio de Janeiro and
coast of Sao Paulo.
Rem arks. — The material from the first locality was identified with the
species hitherto only known from the Atlantic coast, due to the fore end
of the foot, only slightly prominent in the dorsal view of the color photo
(Fig. 138), but crescent-shaped and thickened in the preserved animals
(Fig. 139), similar to our former drawing (Marcus, 1955: Fig. 252).
The color agrees with Edmunds’ description, only the red blotch on the
tentacles is little distinct. The crimson markings around the mouth and the
reddish coloration of the buccal mass and oesophagus, traces of which are
retained in the preserved material, are noteworthy with respect to the
111
antero-middorsal red marks of Ph. patagonica, mentioned in the preceding.
The species of Phidiana are cannibalistic (Marcus, 1955: 471; Lance,
1962: 159; Edmunds, 1964: 18), and therefore frequently have short tails
and missing or regenerating cerata as in the present specimens. The smallest
slug, 6 mm long in preservation, has no stylet yet; the second specimen, 7
mm long, has a stylet whose circular base shows that it is still in develop­
ment; the stylet of the biggest slug, 8 mm long, is torn off.
The ratio of length to breadth shown in the photo, of a specimen with a
regenerating tail is 8/1. There are about 18 denticles on the masticatory
border of the jaw, and 16 radular teeth with 4-6 denticles beside the cusp
and 4 on it.
The color of the animals from the second locality was noted alive by the
collector as: “back orange, with blue median stripe and orange along
sides.” The same colors appear on the photo of the slug from the third
locality. The fore end of the foot of this specimen (Fig. 140) resembles
that of Bergh’s figure (1874a: PI. 9, Fig. 4).

64. G odiva rubrolineata Edmunds, 1964

Figure 141
R eference.— Edmunds, 1964: 23.
M aterial.— Florida: Biscayne Bay; among algae from seawall, Crandon
Marina, Key Biscayne. F. M. Bayer, November 29, 1962. One specimen.
Further distribution. — Biscayne Bay, near the Institute of Marine Sciences,
University of Miami (original locality); Jamaica, from mangrove roots at
Port Royal.
R em arks. — The colors are as in the material from Virginia Key; that from
Jamaica was slightly different. The first right liver has 15 cerata, 11 in
the original material. The hepatic diverticula in the cerata are smooth. The
denticles of the masticatory border are rough. The radular cusp is flanked
by 4-6 denticles on either side. The curved prostatic section of the efferent
duct is visible through the skin.

65. Austraeolis catina; spec. nov.

Figures 142-148
M aterial. — Florida: Biscayne Bay; among algae from seawall, Crandon
Marina, Key Biscayne. F. M. Bayer, November 29, 1962. Two specimens.
Syntypes. — USNM 576274.
D escription. — Preserved the slugs measure about 5 mm in length, the
smooth tentacles nearly 3 mm, the rhinophores 1.5 mm. In the color photo
we counted about 12 rhinophoral rings in the live animal; in preserved
112
condition, due to contraction there are about 18. In this state the anterior
foot corners are 1 mm long; the groove of the anterior foot border is
continued to the tip of the angles. The tail is pointed and slender in the
living slug. The ground color on the photo is transparent greyish; there
are brown marks on the sides of the head and in the middle of the rhino­
phores. The diverticula of the intestinal gland in the cerata are brown.
The skin is irregularly frosted with white, densely in the middle of the
head, on the tentacles, the distal part of the rhinophores, and on the foot

F i g u r e s 142-148. Austraeolis catina, sp . n o v . 142, liv in g a n im a l, fr o m c o l o r

p h o to ; 143, la te r a l v ie w o f p r e s e r v e d s lu g , w ith in s e r tio n o f c e r a ta ; 144, ja w ;
145, d e n tic le s o f m a s t ic a to r y b o r d e r ; 146, r a d u la r to o th ; 147, e v e r t e d m a le
c o p u la to r y o r g a n ; 148, te r m in a l d is c o f sa m e .

113
corners, not around the big eyes. No color marks are retained in the
preserved specimens.
The pointed cerata of the preserved slugs measure up to 1.5 mm in
height. They are arranged in five groups: the three first are arches, the two
last simple, unarched. The cerata on the arches stand in more than one
row; their number is respectively 19, 10, 12, 2, 2. The genital opening
lies near the hind limb of the first arch, the renal pore in the interhepatic
space, the anus within the second arch.
The mandible is light yellow, its masticatory border bears around 30
conical, knobbed denticles. The radula comprises 23 teeth; the cusp of
the tooth is prominent, flanked by 4-6 slightly curved denticles. The
diverticula of the digestive gland in the cerata are nodular, the cnidosacs
indistinctly set off from the wall of the ceras.
The penis is everted in one specimen (Fig. 147); it is strong, cylindrical
and curved upward. It ends with a disc (Fig. 148) whose edge is beset
with 10 broad warts, each bearing a tiny spine. There are two further
warts on the surface of the disc, also with spines. Between them lies the
opening of the thick, winding male duct. There is no penial gland.
Discussion.— Edmunds’ useful survey of the penial characters of the
Facalaninae (1964: 26) leads to the Australian genus Austraeolis Burn
(1962b: 120) for the present species, especially if one adds “or warts”
to “minute fleshy filaments” in the original diagnosis. The type-species of
Austraeolis, A . ornata (Angas, 1864) differs from catina by the absence
of cuticular hooks on the six filaments of the penial glans, 9 rhinophoral
rings, about 20 denticles on the masticatory border of the jaw, and 20-22
rows of radular teeth. A . fucia Burn (1962b: 122), whose penial armature
is not known, has six rhinophoral annuli and only the two first arches of the
liver have distinct anterior and posterior limbs. Its radula contains 19 teeth.

66. Cratena kaoruae Marcus, 1957

R eferences .— Marcus, 1957a: 472; 1959b: 257.
M aterial .— Florida: Whitewater Bay; Clearwater Pass, sta. 7. Salinity
28-33%0; taken among hydroids. Durbin Tabb, April 16, 1961. Four speci­
mens.
Further distribution .— North Carolina, from Cape Hatteras southwards;
Louisiana; Texas; Brazil, 200 km southwest of Santos in a lagoon of
variable salinity, 14-16%0 (original locality).
R em arks. — The length of the preserved slugs is 6, 6, 4, and 3 mm, their
breadth 2.5 mm, their height nearly 3 mm. The cerata are up to 3 mm
long. A slug 6 mm long has three horseshoe-shaped arches of cerata and
three rows. The former bear 12, 13, and 9 cerata, the latter 3, 2, and 2.
The genital openings lie within the first horseshoe, the anus within the
second.
114
 The rhinophores are smooth; the antero-lateral angles of the foot are
produced. The masticatory border of the jaw bears small denticles. The
yellow radula has 29 teeth with 5-6 denticles on either side of the median
cusp.
Lemche (Bull. zool. Nomencl. 21 (1 ): 50-51) proposed the suppression
of Cratena published in 1864 and 1870, in Danish and German, with a
recognizable type-species. As long as this proposal is not accepted, we
continue to use Cratena for Favorinidae (Favorininae) with prominent
lateral denticles flanking the radular cusp. Also, Burn (Bull. Nomencl.
21 (6 ): 410) and Heppell (ibid.: 415) wish to maintain Cratena.

67. Aeolidiella lurana, spec. nov.

Figures 149-150
M aterial. —Brazil: Bay of Santos, Urubuquegaba Island, under stone in
shallow water. Walter Narchi, October 19, 1964. One specimen.
H olotype .— USNM 576293.
D escription .— The length alive is 8.5 mm. Preserved the slug is 6.5 mm
long, 2.0 mm broad, 1.6 mm high. The width of the foot in front is 2.2
mm; the cerata measure up to 1.4 mm, the cnidosac 0.2-0.35 mm.
The color is bluish white, transparent, suffused with pink on the tentacles
and the sides of the head, and with yellow on the sides of the back, the
middle of the head, and the upper part of the rhinophores. There are
orange marks on the head, the rhinophores, and the back, as indicated in
Figure 149. The cerata are orange with dark brown nodular diverticula
of the intestinal gland and a white transverse band in the upper third.
The tentacles with regenerating tips are as long as the rhinophores;
if complete, they certainly would surpass this length. The tentacles and
rhinophores are smooth, the foot is broader than the back, its anterior
border is not grooved and its antero-lateral angles are prolonged into
short processes. The tail is rather long, though in the present specimen
it is in regeneration. The cerata are thickest in their upper third, the bases
slender and the tips conical.
The rows of cerata number 16 on the right side, 15 and one ceras on
the left. They leave a wide space in the middle of the back free. The
disposition of the cerata is in life obscured by the narrow interspaces
between the rows. The anterior liver has five branches on either side
with 3-5 cerata each. The genital apertures lie laterally under the in­
terspace between the second and third row. The first three branches of
the posterior liver form arches; the renal pore is located under the
space between the limbs of the first arch, the anus between the first
and second arches, farther dorsally than the nephroproct. The remaining
branches were not analyzed in order to damage the holotype as little
as possible.
115
 The border of the jaw is smooth. The radula comprises 16 complete
teeth, each pectinate and widely emarginated in the middle. The oldest
tooth is 0.155 mm broad; each wing bears 18 tapering denticles. The
corresponding numbers for the youngest tooth are 0.17 mm and 21
denticles. The central denticle is slightly broader than the neighboring
ones, but not broader than any of the more lateral ones.
D iscussion .— Engel (1925: 38 ff.) and Macnae (1954: 35) gave the
bases for the discussion of A eolidiella Bergh, 1867, and Pruvot-Fol
(1954:428) set forth her opinion as to the synonymy of the European
species. The name Eolidina Quatrefages, 1847, which she used, had been
rejected by Odhner (1939: 83). To Macnae’s list A . hulli Risbec (1928:
262; 1953: 123) and A . m acleayi (Angas, 1864; Burn, 1962b: 127)
must be added. A . alba Risbec (1928: 261; 1953: 124) with bossy

149-150. Aeolidiella lurana, sp. nov. 149, dorsal view of living slug,
F ig u r e s
from a colored figure by Sergio Teixeira da Silva, orange pattern stippled;
150, youngest and oldest tooth.
116
rhinophores and A . ? takanosimensis Baba (Risbec, 1956: 31, 32)
with foliated ones cannot be included in Aeolidiella (Marcus, 1961a: 56).
The only Aeolidiella known from the West Atlantic warm waters is
A . occidentalis Bergh (1875b: 397) from St. Thomas. As only three very
badly preserved slugs have been examined, this species remains problem­
atic. The radula and the location of the anus, somewhat farther in front
than in most true Aeolidiidae, disagree with the characters known for
Aeolidiella. The double arch of the tooth occurs in this genus and in
Spurilla (Marcus, 1955: Figs. 283, 284). In the latter the double arch
is chiefly brought about by the different length of the denticles, in A eo ­
lidiella also by widenings of the basal plate. The tooth of A . occidentalis
is rather of the Spurilla- type. When the smooth rhinophores of A e o ­
lidiella, in some species provided with a spiral groove (Vayssiere, 1888:
109, PL 1, Fig. 8b), are contracted in preserved animals, the circular or
oblique folds that Bergh described of occidentalis may appear. Bergh’s
ventral view of the fore end (1875b: PL 8, Fig. 18) agrees with that of
the present Aeolidiella in preserved condition, not with that of Spurilla
(Marcus, 1955: Fig. 278). The white lines of occidentalis (1875b: Pl.
8, Fig. 9) disagree with Spurilla, whose white dots or spots on the cerata,
head and back will hardly appear as lines in preserved material. In any
case, lurana differs widely from occidentalis.
The 11 valid species which result from Macnae’s list and our additions,
as well as the synonyms (Engel, 1925: 42) or near relatives (Pruvot-Fol,
1954: 429-431) of A . glauca (Alder & Hancock, 1845), were compared
with A . lurana. A more detailed comparison is needed for A . saldanhensis
Barnard (1927: 201) from Southwest Africa; A . takanosimensis Baba
(1930; 1949: 111, 183) from the Pacific coast of Japan from about 35° N
southwards; and A . multicolor Macnae (1954: 36) from False Bay,
South Africa.
Macnae compared his species with A . indica Bergh (1888: 781) from
Mauritius. As he did not see the figure of the radula (PL 78, Fig. 2),
he could not note its less developed wing-like expansions. Evidently,
indica is a distinct species closely allied to multicolor. Risbec (1956:
31) thought that m ulticolor and takanosimensis are one and the same
species. However, the three first groups of cerata of the posterior liver
stand in arches in m ulticolor, while these groups are many-branched in
takanosimensis. A slightly different distance between the nephroproct
and the anus in these species is taxonomically insignificant, as we know
from Spurilla and Berghia (Marcus, 1957a: Figs. 236, 245). As the three
first groups of cerata of the posterior liver in lurana agree with multicolor,
not with takanosimensis, our species approximates multicolor.
The insufficiently described A . saldanhensis differs from m ulticolor and
from lurana by its rounded, not prolonged, antero-lateral angles of
the foot.
117
 The radulae of multicolor and lurana differ by the central denticle,
much broader than the adjacent lateral denticles in the former and only
slightly broader than these in the latter. The number of denticles is
different too, but this character is highly variable, at least in certain
species, e. g., A . japonica Eliot, 1913 (Baba, 1949: 183). The tail of
living m ulticolor (Macnae, 1954: PI. 2, Fig. 4), is 1/10 of the total
length, that of lurana 1/4. The foot of multicolor is narrower than the
back, that of lurana broader. The anterior right liver and its left partner
have 6-7 branches in multicolor, five in lurana.
The differences between the species of A eolidiella are slight, and the
same holds for those between m ulticolor and lurana. Nevertheless, as far
as one dares compare an ample material whose average length is 20 mm
(m ulticolor) with a single slug 6.5 mm long (lurana), the differences are
sufficient for a separation.

68. Spurilla neapolitana (Delle Chiaje, 1823)

References. — MacFarland, 1909: 91 (braziliana); Marcus, 1955: 184;
1957a: 475 ( neapolitana var. braziliana ); 1959b: 258; 1960b: 187;
1962b: 480.
M aterial .— 1. Florida:
Biscayne Bay; Matheson Hammock; Thalassia
and sand flats. F. M. Bayer, February 21, 1948. Five specimens.— 2.
Florida: Biscayne Bay, on Sargassum at Belle Isle, Venetian Causeway.
H. Boehme, July 1947. Two specimens.— 3. Florida: Ragged Key No. 4.
March 29, 1957. One specimen.
Further distribution. — Mediterranean Sea; Atlantic coast of France and
Morocco; Canary and Cape Verde Islands; Sargasso Sea; Florida, Biscayne
and Florida Bay; Texas; Brazil southwards to Sao Paulo.
R em arks. — The largest of the preserved specimens is 18 mm in length,
hence full-grown. No color marks are retained. As the examined specimens
from the first and second localities have a smooth masticatory border of
the jaw as in all hitherto studied American individuals, we have once
more pondered the possibility to maintain MacFarland’s separation. How­
ever, the nearly 3 ^ high denticles (Bergh, 1877a:761) of Mediterranean
neapolitana may be almost completely worn (Bergh, 1883: 14) or
present only for a short extension in front and behind (ibid.: 17).
Vayssiere (1888: PI. 5, Fig. 71) gave a drawing of the mandible of his
biggest specimen completely without denticles (ibid.: 115); also Pruvot-
Fol (1954: 433, Fig. 169, e) indicates the absence of denticles.
Zoogeographically it is meaningless to stress the absence of mandi­
bular denticles in American neapolitana by applying a special name, be­
cause Sp. sargassicola (Bergh, 1861) from the Sargasso Sea and Arcachon
has denticles 12-13 /x high (Bergh, 1877a: 761, note 1), and this species
118
is considered merely as a synonym of neapolitana (Engel, 1925: 46, 48;
Pruvot-Fol, 1954: 433).

69. Glaucus atlanticus Forster, 1777

References. — Bergh, 1864: 105 (monograph); Simroth, 1895: 174
(records from the tropical Atlantic).
M aterial.— Florida: Virginia Key, beach at Marine Laboratory. December
9, 1961. One specimen.
Further distribution. — Circumtropical, pelagic.

70. Onchidella floridana (Dali, 1885)

Figures 151-155
Principal references.— Dali, 1885: 288; Binney, 1890: 203; Hoffmann,
1928: 95; Abbott, 1955: 274.
M aterial. — Florida: Bear Cut, Key Biscayne. R. C. Work, March 13,
1964. Three specimens.
Further distribution.— West coast of Florida, Knights Key, north to
Charlotte Harbour (original locality); Dry Tortugas; Lower Keys; Cuba;
? Bermuda.
D escription. — The notum of the three preserved slugs examined is of a
uniform dark greenish grey. The underside is light grey, the upper border
of the sole dark. Dali described the living animals as uniformly slate blue
to dark grey with a bluish white underside and a greenish tinge to the
sensory lobes.
The length measured over the back (and linear) is in mm: 25 (16),
29 (13), and 29 (12); the breadth 18 (11), 20 (13), and 17 (11). The
breadth of the sole is 6, 5, and 4 mm; the height of the body 7, 8, and
7 mm. The hyponotum of the smallest slug is as broad as the sole on the
sides, much narrower on the hind end. The greatest breadth lies behind
the middle of the body. The hyponotal line is about in the middle of the
hyponotum, its inner half is concealed by the sole in ventral view. The
foot is broad in front, short and pointed behind; the sensory lobes are
well developed; the length of the tentacles cannot be evaluated in pre­
served animals, though they are everted in one slug.
The warts of the notum are rather uniformly middle-sized with some
quite small ones between them. About 100 middle-sized tubercles accom­
pany the frilled perinotum. These warts are more or less prominent accor­
ding to their place on the frills. Twenty big mantle glands are recognizable
by dissection (Fig. 151), not by larger tubercles, because those containing
outlets of smaller glands are not smaller. This contrasts with the disposition
119
 I
F i g u r e s 151-155. Onchidella floridana (D a li). 151, piece of mantle border
with papillae and opened glands; 152, radular teeth; 153, diagram of alimentary
tract; 154, diagram of reproductive organs; 155, male copulatory organ.
120
first described of O. celtica (Joyeux-Laffuie, 1882: 244), and generally
observed in Onchidella.
The pneumostome, the opening of the mantle cavity (anus), and the
female aperture lie close together; they are concealed by the tip of the
foot. The peritoneum is not pigmented; there is a black membrane around
the central nervous system. The latter is much more concentrated than
in Stantschinsky’s species of Onchidium (1907: PI. 12, Figs. 16-19).
The alimentary tract (Fig. 153) furnishes no specific characters. The
radula (Fig. 152) has 77 rows and 79 teeth per half-row. The central
cusp of the rhachidian tooth is weak, though not shorter than the lateral
cusps. The lateral teeth are obtuse and spatulate as in most species of
Onchidella; the innermost tooth is, as usual, smaller than the following
ones. The intestine shows Plate’s type 4 (1893: PI. 8, Fig. 32), as in
the other known species of the genus. In front it loops around the root
of the pigmented aorta, which divides into an anterior and a posterior
trunk. The details of the intestinal course and the pyloric caecum are as
in O. indolens (Marcus, 1956b: P1.23, Figs. 4, 6). The posterior lobe
of the intestinal gland is small. The lung is slightly pigmented; the surface
of the kidney is smooth, its inner wall lamellate, the recurrent renal limb
is folded under the right half of the kidney.
The hermaphrodite gland is brown; it lies over the bursa. The her­
maphrodite duct begins with a short ascending and descending spiral,
followed by a tubular caecum ( u ), much bigger than that of O. celtica
(Fretter, 1943: Fig. 4, c), and the seminal vesicle of O. steindachneri
(Hoffmann, 1928: PI. 3, Fig. 4), but similar to that of O. indolens
(Marcus, 1956b: PI. 24, Fig. 10, u ). The genital complex resembles that
of steindachneri (Hoffmann, 1928: 40-41): the prostate, Hoffmann’s
“appendicula,” is compact; the section of the oviduct containing the mucus
glands is coiled; the albumen glands are arborescent; the brown, globular,
ample bursa has a rather long duct; the vagina has peculiar retractors,
stressed by Hoffmann (1928: 41), and a flagelliform gland.
The male duct enters the body cavity near the right tentacle, describes
some loops, and enters the penial retractor. The latter originates at the
hind end of the body cavity, hence corresponds to Plate’s type III (1893:
148, 170, note 1) as in steindachneri (Hoffmann: 41). Also the penis
is similar to the latter, though shorter. The male duct opens between two
muscular papillae (Hoffmann’s “Reizkorper” ) in the fundus of the penial
pouch. Farther in front the pouch becomes broader; its foremost, atrial
part is narrowed again. The lining of the pouch is a thin cuticle thrown
into longitudinal spiral folds, connected with one another by numerous
fine transverse ones.
Discussion .— In classifying species of Onchidella the male organ should
be considered. Of the external characters the color is the least important.
In O. indolens (Gould, 1852) it varies in living slugs at the same locality,
121
not correlated with age and substratum (Marcus, 1956a: 88), and the
sculpture of the notum appears different in wet and dry, crawling and
quiet slugs. Insufficiently preserved material may exhibit strange textures
of the notum (Hoffmann, 1928: 96). The sculpture of specimens collected
in 1852, examined 76 years later (Hoffmann, 1928), and that of fresher
ones (Stearns, 1894) appears different. The marginal tubercles bearing
the outlets of the large mantle gland may be more or less salient, as they
are pressed outward by the more or less intense contraction of the cutaneous
muscles. Hoffmann (1928: 33) counted about 20 bigger glands in O.
armadilla (Morch, 1863: 43), but reported (p. 97) that the border is
actually nearly smooth, and that the glands open on small, hardly
projecting prominences. O. indolens has 14-30 large mantle glands, ac­
cording to the size of the body, O. borealis 20-28 (Marcus, 1961a: 3),
so that these numbers are diagnostically of little use.
Whether the large mantle glands can be counted according to more
prominent perinotal tubercles or do not appear without dissection seems
to be a significant character, which makes the name of the Bermudian
Onchidella problematic. Heilprin (1889: 327, PI. 16, Fig. 4), who had
not seen Dali’s description (1885), introduced an Onchidium trans-Atlan-
ticum, certainly an Onchidella, from Bermuda, without any specific charac­
ters. It might be florid ana, as Pilsbry asserted (1900: 503)* but this is
by no means certain. On the contrary, Arey & Crozier’s observation
(1921: 446) that 14 of the perinotal tubercles belong to large mantle
glands makes Pilsbry’s opinion doubtful, because in floridana the large
mantle glands cannot be evaluated from the size of the perinotal tubercles.
In any case, the occurrence of O. floridana on the Bermuda Islands needs
confirmation.
Slugs from areas where the ranges of two species approximate one
another or possibly overlap should be dissected. Cuba can be included
in the range of floridana, because Clench & Aguayo (1939: 6) mention
that the penes of floridana and armadilla are different. The record of
floridana from Puerto Rico (Dali & Simpson, 1901: 307), however, is
doubtful, though Dali is the author of floridana. Hoffmann (1928: 96)
thinks that the Puerto Rican floridana is actually armadilla, due to the
proximity of St. Thomas, the original locality of armadilla. White’s
Onchidium sp. (1952: 106) is certainly floridana by its occurrence at
Dry Tortugas, whence Thiele reported floridana (1910b: 125).
O. indolens and O. armadilla differ from O. floridana by their long
penial papilla pierced by the efferent duct, by their penial pouch devoid
of spiral folds, and by the origin of the penial retractor approximately
on the level of the central nervous system, corresponding to Plate’s type
I (1893: 148).
O. floridana differs widely from the neighboring Atlantic species, and
agrees in most characters with O. steindachneri (Semper, 1882: 280;
122
Stearns, 1894: 384; Hoffmann; 1928: 38) from the Galapagos Islands.
They are separated by the regularly diversified perinotal tubercles of
steindachneri (Hoffmann, p. 39) and the caecum of the hermaphrodite
duct. The ratio of length to breadth of this appendage is 2/1 in stein­
dachneri, and 5/1 in floridana.

Z o o g e o g r a p h ic R em arks

Already more than fifty years ago Thiele (1910b: 109) called the
Malacofauna of the West Indies “on the whole well known,” mentioning
that nudibranchs and cephalopods were not considered. In 1906 (p. 132)
Eliot found the knowledge of the West Indian nudibranchs very imperfect.
We think that now Thiele’s opinion can be generalized for the opistho­
branchs. The tropical West Atlantic species of the present collection
gathered between Florida and Cabo Orange (4° 46' N) can be grouped
according to their entire range or systematic affinities as follows:
1. Species only known from the West Indies: Oxynoe antillarum, Tri-
dachia crispata, Cyerce antillensis, Umbraculum plicatulum (perhaps
better in group 3 or even in 7), Pleurobranchaea occidentalis and P.
agassizii, H ypselodoris edenticulata, Discodoris hedgpethi, D. purcina,
A phelodoris antillensis, and Phyllidiopsis papilligera.
2. Species of the western Atlantic Ocean and the American Pacific
coast: Berthellina quadridens, Pleurobranchus areolatus, Phidiana lynceus
and, not in the present collection, Tritonia exsulans Bergh, 1894 (Marcus.
1961a: 32), occur on either side of the Isthmus of Panama and are
probably testimonies of the intercommunication between the western
Atlantic and the eastern Pacific Oceans during part of the Pliocene epoch
(Ekman, 1953: 30-38; Weisbord, 1962: 8). Collier & Farmer (1964:
380, 389) recorded Cadlina evelinae and D endrodoris krebsii from several
localities in the Gulf of California. The Tertiary connection of the tropical
opisthobranch fauna in the West Atlantic and East Pacific Oceans explains
the affinities of Onchidella floridana with O. steindachneri from the Gala­
pagos Islands. Other cases were previously recorded (Marcus, 1961a: 57).
3. Species from the West Indies and the Bermudas: The original de­
scriptions of the Bermudian species are so incomplete that only Elysia
papillosa can be allotted to this group. Elysia tuca seems to be related to
a species from the Bermudas, and Umbraculum bermudense is probably a
young U. plicatulum. Whether H ypselodoris edenticulata and H. acriba
are identical with Bermudian species, cannot be decided yet.
4. West Indian and Brazilian species: This most numerous group evi­
dences the unity of the littoral opisthobranch fauna of the warm West
Atlantic Ocean. M icrom elo undata, H ydatina vesicaria (belongs also to
group 5), Haminoea elegans, Lobiger souverbiei (see also group 6),
Elysia evelinae, E. cauze scops (also group 6), Phyllaplysia engeli,
123
Berthella tupala, Pleurobranchaea hedgpethi hamva, Cadlina rumia,
Chrom odoris neona, Siraius kyolis (a new species with Brazilian affinity),
Peltodoris greeleyi, D iscodoris evelinae, D. phoca, D. pusae, Taringa telopia
disa, Platydoris angustipes, D endrodoris krebsii, Bornella calcarata, Phidi-
ana lynceus (belongs also to group 2), Cratena kaoruae, and Spurilla nea-
politana (belongs also to group 5).
5. West Indian species occurring also in the eastern Atlantic Ocean
including the Mediterranean Sea or related to species of this region:
H ydatina vesicaria (also group 4), H aminoea antillarum, Anisodoris w orki
(a new species related to the Adriatic A . m arm orata ), and Spurilla
neapolitana (also group 4).
6. West Indian species which also occur in the Indo-West Pacific Ocean
(the three first) or have systematic relationships with inhabitants of this
region (the three la st): Chelidonura hirundinina, Lobiger souverbiei,
Petalifera ramosa, Felimare bayeri, G odiva rubrolineata, and Austraeolis
catina. Elysia aff. cauze is reported from Hawaii.
7. Species of the West Indies and of most warm seas: Dolabrifera
dolabrifera, A plysia dactylom ela, Stylocheilus citrinus on flotsam, and
the two pelagic slugs Fiona pinnata and Glaucus atlanticus.
8. In all seas with exception of the polar regions: Lamellaria per-
spicua.
9. New West Indian species whose zoogeographic affinities cannot yet
be recognized: Aglaja pusa, Elysia duis, Chromodoris nyalya, A nisodoris
prea, and Tritonia bayeri.
10. Of the nine species collected on the Pacific side of Panama three,
which are also known from the tropical West Atlantic Ocean, were
mentioned in the preceding. Also two further species, Tridachiella diom edea
and Taringa aivica have Atlantic relationships. The former is the represen­
tative of a monotypical genus, only known in the Bay of Panama and in
the likewise tropical Gulf of California, and is vicarious for the Caribbean
Tridachia crisp at a, also the only species of its genus. Taringa aivica
belongs to a well characterized genus which is recorded from Brazil,
Florida, and the eastern Mediterranean Sea.
The four remaining Pacific species do not evidence Atlantic relations.
Those of D oriopsilla janaina are distinctly Californian, as the most similar
species reaches its southern limit at 27° 51' N. Two further new Pacific
species belong to the circumtropical and circumsubtropical genera
Chrom odoris and D iscodoris. Their geographic affinities cannot be defined
positively; only the absence of Californian and Caribbean similarities can
be stated. It must be stressed that the opisthobranchs of the tropical
Pacific coast of Central and South America are very little known.
The most enigmatic range is that of Titiscania limacina. It is known
from the Pacific coast of Panama (present collection), the Gulf of Califor­
124
nia (Dr. Peter E. Pickens, Tucson, Ariz., coll.), the West Pacific Ocean
(Marshall Islands), the Eastern (Philippines) and the Western Indian
Ocean (Sea of Mauritius). Even if the anatomical differences between ours
and the previously described material resulted in a specific separation, the
Indo-West Pacific and the East Pacific tropical forms would be closely
related. As long as the biology of T. limacina is not known, not even a
reasonable hypothesis can be ventured to explain how Titiscania has over­
come the East Pacific barrier.
11. In the present paper, five species are treated that occur on a section
of the coast of southern middle Brazil, but are not recorded from the
West Indian region. The benthic opisthobranch fauna of the upper littoral
is reasonably well inventorized for the referred section, the coast of
Sao Paulo, which, however, comprises less than one tenth of the Brazilian
coast. Therefore a subdivision of the Brazilian opisthobranch fauna,
perhaps an inner tropical local fauna north of Cabo Frio and an outer
tropical one south of it, would be premature. A generalized qualification
of the Brazilian opisthobranch fauna as an impoverished West Indian fauna
with a number of endemic species is justified, because the relatively well
known area lies not far from the southern limit of the western Atlantic
warm-water fauna. This limit can be assumed to lie in northern St.
Catarina, hence farther south than indicated by Ekman (1953: 47).
The mangrove ends at about 27° S, not at Rio, and also large blocks of
stony corals, it is true, no reefs, occur south of Rio. The continental
shelf of St. Catarina is relatively narrow. Therefore the high temperature
and salinity of the Brazilian Current running along the edge of the shelf
reach the littoral of St. Catarina, which is little influenced by conti­
nental inflows.
One of the five Brazilian species, Tritonia wellsi, can be expected to
occur also in the West Indian region, because it is recorded from North
Carolina, south of Cape Hatter as. In contrast with this warm-water species
Marionia cucullata is eurythermal; its range extends from the upper littoral
at 38° 50' S to that of Rio de Janeiro. A third zoogeographically in­
teresting species of this Brazilian group is Aeolidiella lurana. A single
specimen of it was gathered on a place we frequently visited, 100 m in
front of the beach of Santos. The species is allied to a South African
one from False Bay, and is the first case of a southern transatlantic
relationship in the Brazilian opisthobranch fauna. In that from Florida
a parallel case, G odiva rubrolineata , is known.

R esu m o

O trabalho contem 70 especies, descritas ou brevemente mencionadas,

68 opistobranquios e dois prosobranquios, Titiscania limacina Bergh, da
costa Pacifica da Zona do Canal do Panama e uma Lamellaria. O material
foi coletado, principalmente, na costa da Florida e nos mares vizinhos;
125
algumas especies sao do Golfo do Panama, cujas relagoes faumsticas com
o mar dos Caraibas sao evidentes; outras provem do Brasil, uma do Cabo
Orange e algumas das costas de Sao Paulo e de Sta. Catarina. A fauna dos
opistobranquios da zoma das mares das Indias Ocidentais e a do Brasil,
pelo menos no trecho razoavelmente bem conhecido, entre Rio e Sao Paulo,
tem muitas especies em comum. Alem de 2 novas subespecies e 18 novas
especies das quais uma, Feiimare bayeri, e considerada como represen-
tante de um novo genero, os novos nomes seguintes sao introduzidos:
Risbecia jrangoisi (Risbec, 1928), em vez de Risbecia odhneri Risbec,
1953; A nisodoris rudberghi, nom. nov., em vez de A . marmorata Bergh,
1898 (non A . marmorata Bergh, 1881); e A porodoris risbeci, nom.
nov., em vez de Thordisa hilaris var. Risbec, 1953.

Z U SA M M E N F A SSU N G

Die Arbeit enthalt 70 beschriebene oder kurz erwahnte Arten, und

zwar 68 Opisthobranchier und 2 Prosobranchier, Titiscania limacina
Bergh von der pazifischen Seite der Kanalzone und eine Lamellaria. Das
Material stammt hauptsachlich von der Kiiste von Florida und aus benach-
barten Meeren; weitere Arten wurden im Golf von Panama gefunden,
dessen faunistische Beziehungen zum karaibischen Meer deutlich sind, und
einige Arten wurden an der Kiiste von Sao Paulo und Sta. Catarina
gesammelt. Die Gezeitenzonen Westindiens und Brasiliens, wenigstens in
dem einigermaassen erforschten Abschnitt Rio-Sao Paulo, haben viele
Opisthobranchier-Arten gemeinsam. Ausser 2 neuen Unterarten und 18
neuen Arten, von denen eine, Feiimare bayeri, als Typus einer neuen
Gattung angesehen wird, werden folgende neue Namen eingefiihrt: Risbecia
jrangoisi (Risbec, 1928), fur Risbecia odhneri Risbec, 1953, Anisodoris
rudberghi , nom. nov., fur A . marmorata Bergh, 1898 (non A . marmorata
Bergh, 1881), und A porodoris risbeci, nom. nov. fur Thordisa hilaris var.
Risbec, 1953.
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137
 P art II

Opisthobranchs from the G u lf of California

 OPISTHOBRANCHS FROM T H E GULF OF
CALIFORNIA
EVELINE a n d ER N ST MARCUS
,
Departm ent of Z oology University of Sao Paulo, Brazil

A bstr ac t
The paper contains a total of 47 species and subspecies collected by
Dr. Peter E. Pickens (University of Arizona, Tucson) in the intertidal
zone of the northeastern part of the Gulf of California. Of the 20 new
forms lnuda luarna, sp. n., and Tayuva ketos, sp. n., represent new genera
of the Dorididae. Several species recently collected on the Pacific coast of
Panama by Dr. Frederick M. Bayer occur also in the Gulf, among them
the Indo-West Pacific neritacean Titiscania limacina. About two thirds of
the present collection are warm-water opisthobranchs; relationships with
the tropical West Atlantic fauna are evident.

I n t r o d u c t io n
Dr. Peter E. Pickens of the University of Arizona, Tucson, sent us 44
species of opisthobranchs for classification, adding three prosobranchs
without shell (Titiscania) or with an internal one (Lamellaria), traditionally
collected and studied together with opisthobranchs. The very well preserved
material was accompanied by photographs of the living animals in natural
colors.
Except for the so-called nudibranchs, the malacofauna of the Gulf of
California is well known. Contrasting with the numerous references to the
prosobranchs (Parker, 1964: 151-156) and the Cephalaspidea (Keen,
1958: 496-499) those to the nudibranchs are poor (Steinbeck & Ricketts,
1941: 538-547). Since then, the range of several Californian species was
extended to the Gulf (Winkler, 1958; Lance, 1961; Farmer & Collier,
1963), and new species were described by Farmer (1963), Collier &
Farmer (1964), and Lance (1962b).
The inventory of the opisthobranchs is certainly still incomplete for
this sea extending over nine degrees of latitude. According to Parker
(1964: 3, 27) “the configuration of the shoreline of the Gulf is so variable
that a tremendous number of ecological niches are present for marine
and intertidal invertebrates. The great diversity of species in any one
environment is the outstanding feature of the shallow water assemblages
in the Gulf of California.”
For a fine set of kodachromes of opisthobranchs, chiefly from the area
of La Jolla, we are indebted to the generosity of Dr. James R. Lance.
To Professor Dr. Frederick M. Bayer we are very much indebted for
revising the language in our manuscript and for reading the proofs.
The material seen by the authors will be sent to the United States
National Museum. Dr. Peter E. Pickens has kept duplicates of most species
in the Department of Zoology, University of Arizona, Tucson.
141
 E x p l a n a t io n of L e t t e r in g

a— ampulla ne— pore of notal gland

ac— common atrium ni— nidamental duct
am— male atrium no— notal gland
an— albumen gland nu— loose tissue
ar— anus nx— pharynx
as— anterior section of efferent duct o— inner oviduct
aw — female aperture oa— genital aperture
b— gill oe— oesophageal pouch
bi— appendage of rhinophorial sheath or— oral glands
c— caecum os— oesophagus
ca— seminal groove ov — ovotestis
cc— buccal ganglia p — penis
ci— cuticular tube q — prostate
co— oral tube qu— ectal portion of prostate
cu— spicules r— retractor
d— efferent duct ra— parapodium
e— ejaculatory duct —
ri right liver branch
ei— brain —
rs rhinophoral sheath
eo — ciliary groove s— spermatheca
/—flap sa— salivary gland
g— female gland mass sc— spermatocyst
h— hermaphrodite duct se— septum
i— intestine si— siphon
ia— inner liver branch so— stomach
in— insemination duct —
ss sheathed part of male duct
io— rhinophore sv— seminal vesicle
is— pigment granules t— tentacle
iz — gizzard u— rhinophoral plumes
k— kidney v— vagina
I— right branch of posterior left liver ve— ventricle
lu— lung vn— vestibular gland
vo— velar appendage
m — mouth vs— vessel
ma— male aperture w— body wall
mo— mantle foramen x— auricle
mu— mucus gland y — eye
n— renal pore zo— foot corner

L ist of G a st r o po d s C l a s s if ie d in t h e P r e se n t P aper

Subclass PROSOBRANCHIA ( = Streptoneura)

Order ARCH AEOGASTROPODA ( =D iotocardia)
Superfamily N e r it a c e a
1. Titiscania limacina Bergh, 1875. Fig. 1.
Order MONOTOCARDIA
Suborder M e s o g a s t r o p o d a ( =T aenioglossa)
Superfamily L a m e l l a r i a c e a
Family Lamellariidae
2. Lamellaria perspicua perspicua (Linne, 1758), Figs. 2-3.
3. Lamellaria perspicua mopsicolor du Bois-Reymond Marcus,
1958. Fig. 4.
 Subclass EUTHYNEURA
Superorder OPISTHOBRANCHIA
Order CEPHALASPIDEA (Tectibranchia 5. str.)
Family Bullidae
4. Bulla gouldiana Pilsbry, 1895.
Family Atyidae
5. Haminoea virescens (Sowerby, 1833). Figs. 5-7.
Family Aglajidae
6. Navanax inermis (Cooper, 1862). Fig. 8.

Order ASCOGLOSSA
Family Hermaeidae
7. Hermaea hillae, spec. nov. Figs. 9-11.
Family Elysiidae
8. Tridachiella diomedea (Bergh, 1894).

Order A N A SPID EA
Family Aplysiidae (Subfamily Aplysiinae)
9. A plysia (Pruvotaplysia) parvula Morch, 1863.
10. A plysia (Neaplysia) californica Cooper, 1863. Fig. 12.
11. Aplysia (Aplysia) juliana Quoy & Gaimard, 1832. Figs. 13-15.
Family Aplysiidae (Subfamily Notarchinae)
12. Stylocheilus longicauda (Quoy & Gaimard, 1824). Figs. 16-17.

Order NOTASPIDEA
Family Pleurobranchidae
13. Berthellina engeli ilisima, subsp. nov. Fig. 18.
14. Pleurobranchus (Pleurobranchus) areolatus (Morch, 1863).
Fig. 19.

Order D ORIDOIDEA
Suborder E u d o r id o id e a
Tribe C r y p t o b r a n c h i a
Dorididae (Subfamily Conualeviinae)
15. Conualevia mizuna, spec. nov. Fig. 20.
Family Dorididae (Subfamily Chromodoridinae)
16. Cadlina evelinae Marcus, 1958.
17. Chrom odoris norrisi Farmer, 1963. Figs. 21-24.
18. Chromodoris banksi sonora subsp. nov. Figs. 25-29.
19. H ypselodoris californiensis (Bergh, 1879). Figs. 30-33.
20. Casella sedna, spec. nov. Figs. 34-37.
Family Dorididae (Subfamily Inudinae, subfam. n ov.).
21. Inuda luarna, gen. nov., spec. nov. Figs. 38-44.
Family Dorididae (Subfamily Doridinae)
22. D oris pickensi, spec. nov. Fig. 45.
Family Dorididae (Subfamily Discodoridinae)
23. D iscodoris mavis, spec. nov. Fig. 46.
24. Diaulula sandiegensis (Cooper, 1862).
25. Taringa aivica timia, subsp. nov. Figs. 47-51.
26. Tayuva ketos, gen. nov., spec. nov. Figs. 52-56.
143
 Tribe P h a n e r o b r a n c h i a
Superfamily N o n s u c t o r ia
Family Gymnodorididae
27. N em brotha eliora, spec. nov. Figs. 57-58.
Family Polyceridae
28. Polycera atra MacFarland, 1905.
29. Polycera gnupa, spec. nov. Fig. 59.
Superfamily S u c t o r ia
Family Onchidorididae
30. Acanthodoris pina, spec. nov. Fig. 60.
Suborder P o r o s t o m a t a
Family Dendrodorididae
31. Dendrodoris krebsii (Morch, 1863). Fig. 61.
32. Doriopsilla albopunctata (Cooper, 1863).
33. Doriopsilla janaina Marcus, 1967.
34. Doriopsilla rowena, spec. nov. Fig. 62.

Order D EN D RO NO TO ID EA
Family Tritoniidae
35. Tritonia (Candiella) pickensi, spec. nov. Fig. 63.
Family Dendronotidae
36. Dendronotus nanus, spec. nov. Figs. 64-65.
Family Dotoidae
37. D oto amyra Marcus, 1961.
38. D oto lancei, spec. nov. Fig. 66.

Order A R M INO IDEA

Suborder P a c h y g n a t h a
Family Dironidae
39. Dirona picta Cockerell & Eliot, 1905. Fig. 67.

Order AEOLIDOIDEA
Suborder P l e u r o p r o c t a
Family Flabellinidae
40. Coryphella iodinea (Cooper, 1862). Figs. 68-69.
41. Coryphella cynara, spec. nov. Figs. 70-75.
42. Flabellina telja, spec. nov. Figs. 76-82.
Suborder C l e i o p r o c t a
Family Facelinidae
43. Hermissenda crassicornis (Eschscholtz, 1831).
Family Aeolidiidae
44. Spurilla chromosoma Cockerell & Eliot, 1905.

Order SOLEOLIFERA
Superfamily O n c h id i a c e a
Family Onchidiidae
45. Onchidella binneyi Stearns, 1893. Fig. 83.
46. Onchidella hildae (Hoffmann, 1928). Figs. 84-86.
47. Hoffmannola hansi, spec. nov. Figs. 87-95.
 S y s t e m a t ic S e c t io n

1. Titiscania limacina Bergh, 1875

Figure 1
M aterial.— Puerto Lobos, Sonora, Mexico. In an area of small boulders.
P. Pickens, April 16, 1965. One female.
Further distribution. — lie aux Fouquets, sea of Mauritius; Philippine
Islands, Camiguin, 18°55'N, 121°50'E (original locality); Marshall Is­
lands, Eniwetok Atoll, U.S. National Museum nos. 1502(574920) and
1503(574923), classified by the authors; Canal Zone, Pacific coast, Perico
Island, Ft. Amador, under rocks at low water. F. M. Bayer and R. W.
Bayer, December 25, 1962; four females.
R em arks .— The species has been described in Part I. Therefore, we now
only report its occurrence in the Gulf of California. The measurements
(in mm) of the present preserved specimen are: linear length 6, over the
back about 12, breadth 3, height 3, length of tentacles 1.5. For recognition
of this unique shell-less, nonpelagic, free-living prosobranch we mention
its principal outer characters: white, slug-shaped; two thin, long tentacles;
an anterior oval area about one fourth of the body length, which is the
roof of the mantle cavity; and a row of dense white glands on both sides
of the back. The ctenidium may project from the pallial cavity.

2. Lamellaria perspicua perspicua (Linne, 1758)

Figures 2-3
References. — Marcus, 1959: 13 (literature); 1960a: 189.

M aterial.— Puerto Penasco, Sonora, Mexico. In the rocky intertidal zone.

February 13, 1965. P. Pickens.
Further distribution. — In all seas, except the Arctic and Antarctic; in­
tertidal to 1287 m. Original locality: Mediterranean Sea.
Description. — The following refers to one male (collection no. 30) and
one female (no. 31). The photos show the highly arched mantle, the
respiratory siphon about in the middle of the front, the protruded tentacles,
the hind end slightly drawn out and, in the male, the opaque white
anterior border of the foot. The animals are more or less translucent.
Both are whitish, the male with a pinkish, the female with a brownish
hue. Glandular inclusions of the skin are opaque white in the male, vitreous
in the female. In the preserved male black pigment spots stand out from
the grey ground color; the female is more uniform. The sole is grey, the
head lighter, and the penis white. The roof of the shell chamber is black,
the floor white.
145
 The notum is rather smooth. In the male there is a minute external
scar in the middle of the mantle, which examined from the inside is not
visible. Most measurements of the male are smaller than those of the fe­
male (Marcus, 1959: 13). The lengths are 15 and 18 mm, the breadths
10 and 19 mm, but the heights are 9 and 8 mm.
The shell of the male (Fig. 2) is 12 mm long, 9 mm broad, and 6 mm

F ig u r e1. Titiscania limacina Bergh; side view of preserved specimen.—

2-3. Lamellaria perspicua perspicua (L inne). 2, shell, outer surface
F ig u r e s
(a ); inner surface (b ); right side view (c ); 3, radula.— F ig u r e 4. Lamellaria
perspicua mopsicolor d.B.-R. M arcu s; r a d u la .— F i g u r e s 5-7. Haminoea
virescens (Sowerby). 5, (a) shell, (b) outer lip of shell; 6, right Hancock’s
organ; 7, shell sculpture.
146
high. That of the female is 15 mm long; it is broken. The shell is white,
thin; its growth lines are distinct. The inside is smooth, shiny. There are
about three whorls; the apex is little prominent. The columella is open,
and so the apex can be seen from below.
The jaws are very weak in both specimens. The radula of the male
(Fig. 3) has 60 rows, that of the female 79 rows, and the teeth (in
micra) are a little smaller in the male (lateral tooth 304) than in the
female (310). The difference between the left (210, 220) and the right
(200, 187) legs of the rhachidian tooth is more pronounced in the female.
The cusp of the rhachidian tooth in the male is narrower (94) than in the
female (140). In the male it has 4-5 strong denticles on the right side
and 6-7 finer ones on the left, where they are difficult to see, because
the cusp is inclined to the left. The corresponding numbers of the female
are 5-8 and 8-10. The lateral tooth of the male has 4-8 strong inner and
7-12 fine outer denticles, that of the female 5-8 and 4-11, less diffe­
rent in size than those of the male.
The tip of the penis, whose shape is known to vary, is distinctly set
off from the base, nearly as in Bergh’s figure (1907: PI. 14, Fig. 9).
Discussion .— We do not know any reference to L. perspicua from the
west coast of North America; it is, however, recorded from southern
Chile (Marcus, 1959: 13). Oldroyd (1927: 737-738) described the fol­
lowing species: 1. L. stearnsii Dali, 1871, whose shell is small, 6.6 mm.
long, 5.1 mm broad, and 3.0 mm high. In a specimen, which Bergh
(1886c: 170) received from Dali, these measurements were 9, 7, and
3.6 mm. Ghiselin (1964: 123) saw a pore in the mantle in a species that
he calls L. stearnsii. When the pore is enlarged, the shell becomes exposed.
Such a pore occurs in Marsenina (Bergh, 1886a: 166; 1886b: 5; 1887:
255), a quite different genus. As Ghiselin noted, in the first edition of
Rogers’ “Shell Book” the pore is mentioned for L. rhombica Dali, 1871,
but not for stearnsii. In the edition of 1951 (p. 147) the description of
rhombica contains the passage “the reflexed mantle does not cover the
shell.” In fact, L. rhombica was allotted to Marsenina many years ago
(Bergh, 1886b: 14; 1886c: 185) and figured with this name in Oldroyd
(1927: 739) and Rehder’s list (Rogers, 1951: 492).
2. L. orbiculata Dali, 1871, as Bergh (1886c: 170) defined it, is
recorded from Alaska to the Gulf of California, e. g., from Puerto Penas-
co (MacGinitie, 1949: 372). Its shell is about the same size as that of
stearnsii and broader than that of perspicua. It has 2.5 whorls.
3. L. diegoensis Dali, 1885. The shell is high, the texture of its surface
malleated. The growth lines are obscure, slightly elevated, transverse
waves (Keen, 1958: Fig. 279). The soft parts (Bergh, 1886c: 172; 1905b:
111) are blackish, in part velvety black (1887: 249); Keen (1958: 325)
found them bright red. In L. perspicua the color is not a good specific
character (Marcus, 1960a: 190).
147
 Keen (p. 326) recorded the rather solid shell of L. inflata (C. B.
Adams, 1852) from Panama.
3. Lamellaria perspicua m opsicolor du Bois-Reymond Marcus, 1958
Figure 4
References. — du Bois-Reymond Marcus, 1958: 11; Marcus, 1959: 14;
1963: 51.
M aterial.— Cholla Bay, Puerto Peiiasco, Sonora, Mexico. June 27,
1965. P. Pickens. The single male was found on the oxystomatous crab
Speloeophorus schmitti, rare in Cholla Bay. As Dr. Pickens supposed,
this association was certainly fortuitous.
Further distribution.— Curasao; Brazil, coasts of Rio de Janeiro and Sao
Paulo (original locality). Magellanic region and southern Chile. Intertidal
and subtidal.
R em arks .— The animal was dark bluish grey (Dr. Pickens). The alcohol
of preservation had stained blue. As the centers of the cutaneous bosses
are dark and surrounded by lighter rings, the similarity of the animal to a
colony of compound ascidians is striking. This “behavioural concealing
adaptation” (Ghiselin, 1964) to sponges and incrusting tunicates was
frequently noted (MacGinitie, 1949: 372; references in du Bois-Reymond
Marcus, 1958: 11). As crustaceans, fish, and turtles are known to eat
synascidians, the biological significance of the similarity can hardly be
understood as protective and resulting from selection. For Lamellaria
living on sponges and feeding upon them, a protective aspect is more likely,
as sponges are seldom eaten by other animals.
The shell of the present animal is middle-sized, 4.5 mm long, 3.5 mm
broad, and 1.5 mm high. The left limb of the rhachidian tooth (Fig. 4) is
longer than the right one, and its cusp slants to the left. It has 4-5 denticles
only on the right side, as is characteristic of mopsicolor. The lateral teeth
bear 4-6 coarse, sharp denticles on the inner and eight small ones on
the outer side.
4. Bulla gouldiana Pilsbry, 1895
Reference.— Marcus, 1961a: 4 (literature).
M aterial.— Puerto Penasco, Sonora, Mexico. January 23, 1965. P. Pickens.
“These animals can be seasonally abundant. Most are found in the winter.
They occur in the sand flats of Cholla Bay near Puerto Penasco.”
Further distribution. —From southern California, Sta. Barbara, through the
Gulf of California (original locality: Guaymas) to Ecuador.
R em arks. — The photograph shows a light yellowish-orange head shield
mottled with white. The shell is as described and figured by Pilsbry
(1893-95: 340, PI. 36, Figs. 22, 23). The biggest shell at hand is 32 mm
in length, although examples 59 mm long have been recorded.
148
 5. H aminoea virescens (Sowerby, 1833)
Figures 5-7
Reference .— Marcus, 1961a: 5.
M aterial. — Puerto Penasco, Sonora, Mexico. March 13, 1965. Robert
Taylor. Abundant material has been found in the intertidal zone in spring
at Puerto Penasco, Puerto Libertad, Puerto Lobos, Tiburon Island, and
San Esteban Island, hence at all visited localities from about 29° N to
the North (Dr. Pickens).
Further distribution. — From Puget Sound to the Gulf of California and
Mexico (Oldroyd, 1927: 42; Abbott, 1955: 279).
R em arks. — Steinbeck & Ricketts’ bubble-shells (1941: 546-547) were
classified by Hanna. Baker & Hanna (1927: 131) doubted the validity of
virescens from the oceanic side of Lower California. They called the shells
from the Gulf, from San Esteban Island to Guaymas Bay, H. strongi
(p. 130). Keen (1958: 496) does not include virescens in her synopsis of
the shells of tropical West America.
We compared the present animals with snails from Point Loma (32°
40' N, 117° 14' W ). The radulae, the Hancock’s organ (Fig. 6), and
the male copulatory organ are the same. The shells from both localities
have distinct growth lines and fine spiral striae (Fig. 7). The callus
(Fig. 5A) is smooth, not rough and covered with small conical pro­
jections as in strongi , and there is “a decided compound, concave curve”
(Fig. 5B) in the apical third of the outer lip, absent in strongi. This
character, already stressed by Baker & Hanna, is here considered as
decisive for determination. The posterior contraction of the shell (Pilsbry,
1893-95: PI. 40, Fig. 5; Marcus, 1961a: Fig. 5) appears less marked,
when the shell is turned more to the right, as evident in Pilsbry’s figure 19
(PL 43). H. virescens rosacea Spicer (1933: 53) has only a moderate
constriction.
The two present shells measure 14X 9.0 and 10X7.5 mm. The animal
in the photograph is dark green with yellowish marks: dots on the head
shield, mottles on the parapodia, and a rather dense area on either side of
the tail. In preservation the shells were brownish, the mantle flecked with
red and black, and the foot reddish.

6. Navanax inermis (Cooper, 1862)

Figure 8
References. — Marcus, 1961a: 7 (literature, food, parasites); MacFarland,
1966: 9.
Material. — (1) Punta Sargento, near Tiburon Island, Sonora, Mexico.
Dorothea Caskey, January 10, 1965. “Seasonally abundant at Pta. Sar-
149
gento, but has also been found at Puerto Penasco and at Kino Bay, south
of Tiburon Island. In the latter localities it was rare. N. inermis feeds on
Haminoea primarily, but has also been known to eat other opisthobranchs
as well. Most of the specimens from Pta. Sargento had clinging copepods
on the parapodial flaps” (Dr. Pickens). (2) Puerto Lobos, between

sc q aw rncL
Navanax inermis (Cooper); male copulatory organ.— F i g u r e s 9-11.
F ig u r e 8 .
Hermaea hillae, sp. nov. 9, diagram of alimentary organs; 10, radular tooth;
11, diagram of reproductive organs.
150
Tiburon Island and Puerto Penasco. Donald A. Thomson, February 25,
1966. “Intertidally among the rocks.”
Further distribution.— From Monterey Bay to southern California (original
locality: San Diego) and Lower California, Mexico (Lance, 1966: 71).
Rem arks. — The photograph (material 1) shows two specimens, 75-mm
long alive, of dark brown to black ground color. One of them is covered
with golden yellow dots, the other with longitudinal stripes of the same
color. The parapodia are edged with a yellow line followed by a row of
bright blue dots. In front of the tail there is a white transverse band, and
the inner side of the caudal flaps is bordered with white.
Dr. Pickens states that the color patterns are various. This is shown
also by the four preserved specimens of material (1), which are more or
less densely dotted or striped. One of them has stripes on the back and
dots on the ventral side. The body of the largest specimen is 40 mm long;
the left caudal flap is 15 mm long and the right one is shorter. In this
animal the head shield is longer than the mantle shield; in the three others
it is shorter.
We give a figure of the protruded penis (Fig. 8) of an animal 36 mm
long and nearly black. There are two flaps (/) of a collar-like dilatation
at the penial base, and the two prostates ( q ) are closely apposed to one
another (Bergh, 1894: 217).
Under the parapodia and on the gill, which is white with brown stipples,
many flat copepods were found. Those known from N. inermis on the
oceanic coast are Pseudomolgus navanacis Wilson, 1935.
The two preserved specimens (material 2) from Puerto Lobos are 60-70
and 37 mm long. They have yellow circular spots on a black ground; those
of the longer specimen are larger than those of the shorter one, but even
in the latter they are bigger than in the material collected in 1965 (1).
In the photograph there are also some blue dots on the border between the
yellow spots. The material collected in 1966 agrees with Bergh’s figures
(1894: PI. 11, Figs. 2-4) based upon Cooper’s original color sketches
(p. 232) better than does that obtained in 1965. One pharynx is everted
as frequently occurs in preserved Aglajidae. The length of the penis is
not correlated with that of the body.
As previously, we use the name Navanax, though a clear separation of
Navanax and Chelidonura is impossible.

7. Hermaea hillae, spec. nov.

Figures 9-11
M aterial. — Puerto Penasco, Sonora, Mexico, among some intertidal hy-
droids (Aglaophenia). Mary Anne Hill, March 5, 1966. Five specimens.
D escription .— Alive the slugs are up to 12 mm in length and have cerata
151
3 mm long. They are translucent with black eyes and mahogany digestive
annexes in the body and diverticula in the cerata. Also the oesophageal
pouch (Fig. 9, o e ) has this color.
The head is rounded in front. On either side it bears a supra-oral lobule,
provided with clusters of sensory cells, which are connected with the
cerebro-pleural ganglia. Sections of H. coirala Marcus show the same in­
nervated bosses which represent vestigial labial tentacles. The rhinophores
are rolled in, open on the outer side, and about 1.8 mm long in preserved
condition. The upper, inner tip is longer than the under, outer one, but
the terms “anterior” and “posterior” cannot be used, because the rhino­
phores turn forwards and backwards. As the intestinal diverticula do not
reach the rhinophores, these are colorless. In front, the foot is a little
prolonged on both sides. Preserved, the sole is concave throughout its
length, hence narrow, and provided with glands. The pointed tail of the
preserved specimens is about 2 mm long.
The cerata stand in a single row of 20-30 on either side; many of them
had fallen off. Their height varies, reaching a maximum of 3.3 mm, which
is thrice the diameter of the body (1 mm). They are fusiform and have
many knobs, especially in their distal part. Each knob corresponds to a
hepatic branch given off at a right angle from the straight diverticulum
running along the axis of the ceras. Large white glands open on the surface
of the ceras, principally in the upper half and on the bosses. The absence
of the albumen gland in the cerata is a character of the genus.
The anus (Fig. 9, a r ) is dorso-median, in front of the heart; the male
genital pore (Fig. 11, m a) lies behind the right rhinophore, on the level
of the brain, and the female pore ( aw) farther behind.
The radula contains 23 teeth (Fig. 10). Four of them are new teeth
in the ascending limb, 14 older ones in the descending limb, and 5 lie in
the slightly projecting ascus. The largest teeth are 90 ^ in length, more than
V3 of which belongs to the base (38 ^ ). The laminae flanking the under­
side of the cusp bear fine, pointed denticles.
Clusters of oral glands (Fig. 9, or) surround the mouth tube. A pair of
very thin salivary glands ( s a ) open from behind into the dorsal food
channel of the pharyngeal bulb, and accompany the oesophagus on its
passage through the nerve ring. Also in H. coirala they were now found in
the sections. The oesophagus is narrow. Before it widens into the dorso-
ventrally flattened stomach it forms an unpaired, knobby pouch (oe)
extending forwards to the pharynx. This pouch is continued into two blind
tubes to the sides of the pharynx. Near the entrance of the oesophagus
the intestine, 0.1 mm in length, leaves the stomach, running upwards. The
dorsal lining of the stomach (so) is thrown into longitudinal folds; its
lumen communicates with the left liver duct and farther behind with the
right one (ri). The stems of the digestive gland extend from the level of
the anus to the beginning of the tail, where they are confluent. Each stem
152
emits a straight tube into each ceras, and a few short, irregular, ramifying
canals (ia) towards the midline, where they do not meet.
In the follicles of the ovotestis the growing germ cells are peripheral,
the ripe ones central. The ampulla (Fig. 11, a) which is constricted in the
sectioned specimen emits a spermoviduct. As in other Elysiacea the bifur­
cation of the latter is difficult to see, because the inner oviduct is short and
narrow.
The prostatic part ( q ) of the sperm duct, immediately ectal to the
bifurcation of the spermoviduct, bears a diverticulum as in H. coirala, and
also the penis is tucked into the male atrium (am) in the same way.
The inner oviduct leads into the duct of the spermatheca (s) or bursa,
recognizable by its contents, unorientated sperm; in the spermatocyst (sc)
or seminal receptacle the spermatozoa are fastened to the wall by their
heads. The common dorso-median duct of the albumen glands (an) enters
the oviduct (o) quite near its entrance into the mucus gland.
“The egg case consists of a small incomplete, transparent circle laid on
the surface of the water rather than being attached to the side of the
aquarium. The egg case will float at the surface unless the surface film is
disturbed. Then it will sink” (Dr. Peter E. Pickens). Similar egg strings
of related genera were drawn by Baba & Hamatani (1952: PL 7, Fig. 5),
Hamatani (1960: Fig. 2C), and others.
The species is named for Dr. Mary Anne Hill.
Discussion .—The most similar species is H. coirala Marcus (1955: 108)
from southern middle Brazil, not recorded from the Caribbean yet, but to
be expected from there. The radular teeth of H . coirala are smooth and
more slender, and the cerata are much shorter, only as long as the diameter
of the body. H. hillae and H. coirala have the character of “twin species.”
The oldest family name of the Stiligeridae as understood by Thiele
(1931: 413) and Pruvot-Fol (1954a: 180) seems to be Hermaeidae
Alder & Hancock (1855: XVI). Six species of this family are recorded
from the Pacific coast of North America: (1) Phyllobranchopsis entero-
morphae (not enteromorphea, MacFarland 1966: 46) Cockerell & Eliot
(1905: 52). Only the leaf-like cerata distinguish this species from those
of the genus Hermaeina, so that Phyllobranchopsis is generally considered
as a subgenus of Hermaeina (Thiele 1931: 413; Macnae 1954: 62, 63;
Marcus 1961a: 13).
(2) Hermaea vancouver ensis O’Donoghue (1924: 19). Though the
author correctly indicated auriform rhinophores as a character of Hermaea ,
he described them as spindle-shaped (p. 20) in vancouverensis. Hence
this species must be allotted to Stiliger.
(3) Hermaeina smithi Marcus (1961a: 12; Gonor 1961) from Zostera.
Generally the species of Hermaeina live on Enteromorpha (M acnae
1954: 53).
153
 (4) Stiliger fuscovittatus Lance (1962c: 33). Differs from no. 2 by
its radula.
(5) Hermaea ornata MacFarland (1966: 38). Because of the extension
of the albumen gland into the cerata, this species is defined as a member
of the subgenus or genus (Burn 1966: 105) Placida.
(6) Hermaeina oliviae MacFarland (1966: 43). According to the
pointed radular tooth this species belongs to Hermaea, not to Hermaeina.
As the reproductive organs are not described, one cannot decide, whether
it is a true Hermaea or a Placida. The shape of the radular teeth, not their
number (39 in an 11.4-mm slug), is similar to that of H. hillae. The
opaque cerata are set in transverse rows of 3-5 beside each other, and
their surface is smooth (PI. 4. Figs. 4, 5), not knobby.
8. Tridachiella diomedea (Bergh, 1894)
References.—MacFarland, 1924: 406; 1966: 54; Steinbeck & Ricketts,
1941: 544.
Material.— 1. Puerto Penasco, Sonora, Mexico. P. Pickens, June and
August, 1964. “Quite common the year round at Puerto Penasco in rocky
intertidal habitats. It is also found at Puerto Lobos. It is one of the few
opisthobranchs that may be seen moving around during the day.”—2.
Guaymas, July, 1965.
Further distribution.—Throughout the Gulf, from the intertidal to 18 m
(original locality: near Ceralvo Island); Pacific coast of Panama.
Remarks.—In comparison with the Panamanian slugs photographed by
Professor Dr. Frederick M. Bayer the present ones show small differences.
There are light blue and yellow lines on the outer sides of the rhinophores,
and the edge of the parapodia is light orange. The four present specimens
are 14-30 mm in length; the frills of the parapodia are more elaborate in
larger than in smaller slugs.
9. Aplysia (Pruvotaplysia) parvula Morch, 1863
Reference.—Eales, 1960: 287-291.
Material.—Puerto Penasco, Sonora, Mexico. Mary Anne Hill, June 28,
1965. A single specimen found in a tide pool.
Further distribution.— In all warm and warm-temperate seas from approxi­
mately 40° N to 40° S. Original localities: St. Thomas, Virgin Islands,
and St. Vincent, Lesser Antilles.
Remarks.—The first finding in the Gulf is San Marcos Island (MacFar­
land, 1924: 398), on the coast of Lower California. The ground color
shown in the photograph is greenish grey, vitreous and of different shades.
Numerous scattered white dots are present, many of which are concen­
154
trated forming white blotches. Also the tips of the tentacles and rhino­
phores are white. Some brown tones occur in front and on the tip of the
tail. The short parapodia are dark, due to the scarcity of white dots, and
rimmed with blue followed by faint orange. In preservation the borders of
the parapodia and the mantle foramen are black; the shell is orange.
10. Aplysia (Neaplysia) californica Cooper, 1863
Figure 12
References.—Eales, 1960: 294; Marcus, 1961a: 9; MacFarland, 1966: 12.
Material.—Puerto Penasco, Sonora, Mexico. P. Pickens, April 30, 1965.
“In both sand and rocky intertidal habitats at Puerto Penasco, Puerto
Lobos, Puerto Libertad, Tiburon Island, and San Esteban Island. At Puerto
Penasco, it appears in the intertidal in December, spawns in January, after
which some of the larger specimens appear to die, and occurs in the inter­
tidal, or at least small to middle sized specimens do, until July, after which
they disappear. Quite often they are found in the rocky intertidal with
A. (A.) juliana. In winter and early spring they are quite abundant”
(Dr. Pickens).
Further distribution.—From the Gulf, also in its southern portion (Winkler,
1958: PI. 35), to Humboldt Bay (40° 45' N, 124° 14' W). Intertidally
and subtidally to 12 m (Lance, 1961: 65; Steinberg, 1963b: 69). Wright
(1960: 63) observed a spawning aggregation between the rocks of the
jetty at the mouth of Bodega Bay (38° 14' N) in the second half of
September, hence in rather cool water. Original locality: San Pedro.
Remarks.—The rectangular plate on the ventral side of the apex of the
shell (Fig. 12) characterizes the species and its subgenus. The photo shows
a greenish-grey ground color with dark pigment spots and lines and white
skin glands. The bars of dark and light areas on the inner side of the para­
podia shine through. The sole of the preserved animals is darker than the
rest of the body. The two specimens at hand are 130 mm long, 70 mm
high, and 60 mm wide. The breadth of the sole is 28 mm. The extracted
shell measures 40X25 mm.
11. Aplysia (Aplysia) juliana Quoy & Gaimard, 1832
Figures 13-15
Reference.—Eales, 1960: 363-369.
Material.—San Esteban Island, SW of Tiburon Island, Sonora, Mexico.
P. Pickens, February 26, 1964. Abundant in an area of small boulders and
sand. A few specimens were found in the large tide pools at Puerto
Penasco during the summer. The species occurs together with A. (N.)
calif ornica (Dr. Pickens).
155
Further distribution.—World-wide in warm seas, including the Mediter­
ranean. Original locality: Mauritius.
Description .—This refers to three specimens of San Esteban. Alive they
were dark purple to black. Preserved they are black, especially on head
and sole. The outer side of the parapodia shows a pattern of grey and white
spots; the inner side is black above, grey below. The upper side of the
mantle is black, the inside of the cavity greyish. The upper side of the gill
has traces of pigment, the under side is white. The purple glands are white.
“The animals put out a milky white secretion when they are disturbed”
(Dr. Pickens). The seminal groove is white (Fig. 13, ca); its bordering
ridges are black. The inner side of the penis sheath is also pigmented,
leaving the white warts free.
The animals are 100, 100, and 110 mm long, about 45 mm broad, and
38 mm high. The shape is compact (Fig. 13), the visceral hump is set
rather far back. The parapodia (rd) are 30 mm apart from one another in
front and behind joined high up (30 mm); their borders are short and
straight. The dorsal area is exposed in two specimens; the parapodia are
closed over the mantle region in one. The head is short, though the mouth
is protruded so that the jaws are visible. The rolled tentacles are joined
to the mouth by oral lobes (co). The stout rhinophores are 7 mm in
length. They stand 18 mm behind the fore end and 12 mm apart from one
another. In front of them the eyes are visible through small pigmentless
areas. The skin appears leathery.
The foot is as broad as the body. Its thin anterior border is slightly
concave. The adjacent part of the sole shows thin longitudinal folds; 2 cm
behind it becomes irregularly wrinkled. The hind end is rounded, wavy, and
projects a little beyond the hump; the edge is pigmentfree. A sucker, a
temporary structure in the subgenus, is not recognizable.
The mantle foramen (mo) is wide (7X 6 mm), medium sized (4X2.5
mm), or small (about 2X 2 mm), due to the different state of contraction.
From the deeply concave hind end of the mantle the short anal siphon (si)
projects with a frilled border. The genital aperture lies to the right of the
anterior mantle rim, not covered by it. The shell measures 33X22 mm.
Radial lines and growth lines are visible; the calcareous layer is dissolved.
The shape corresponds to Eales’ broad type (1960: Fig. 46 d). The mantle
cavity begins 65 mm behind the fore end. The multiporous opaline glands
occupy an area of 24 X 8 mm. In an anterior area of 8 X 6 mm the fundi
of the glands are so long that they hang into the body cavity; the greater
part, however, lies between the fibers of the muscles in the floor of the
pallial cavity.
The cerebral ganglia within the common connective sheath are contigu­
ous, not fused; the buccal ganglia are separate, connected by a short
commissure.
156
 The jaws have a narrow anterior field in front, continuous with the
principal area near the middle, separated laterally. The formula of the
radula (Fig. 15) which is 15 mm in length and 12 mm in width, is
63 X 35.1.35. The colorless rhachidian tooth has a laterally expanded base
and a short cusp with a few faint denticles. The bases of the brown lateral
teeth turn outwards, the heads are short and broad; the cusps are shorter

F ig ure 12. Aplysia (Neaplysia) californica Cooper; inside view of decalcified

shell.—F ig ures 13-15. Aplysia juliana Quoy & Gaimard. 13, dorsal view of
preserved specimen; 14, ventral view of fore end; 15, radular teeth of 50th row
and rhachidian tooth of 25th row.—F ig ures 16-17. Stylocheilus longicauda
(Quoy & Gaimard). 16, color pattern on right parapodium; 17, part of penis.
157
on the inner and longer on the middle teeth of the half-row. Head and cusp
together make up the total height in the middle lateral teeth. The cusps
bear small denticles, often in different numbers on the two sides of one
cusp. One cusp may have a total of two to six denticles. Several of the
outermost teeth are oblong plates without cusps. The palatal rods are well
developed.
The two salivary glands are 50 mm long and 7 mm broad behind and
fixed to the sides of the gizzard. The latter contains 10 big teeth, some
smaller ones in front of them, and the bases of many still smaller teeth
behind. These pointed teeth were found free in the stomach. The long
caecum is directed backward and ends with a double spiral.
The penis is pointed and laterally flattened. The sides are pigmented, the
edges light. The penial base bears a narrow, 1.7-mm-long area of warts on
the edge not occupied by the spermatic groove. These warts are simple
pegs, about 100 fi long and 35 ^ in diameter; they have no spines. The
same holds for the compound warts up to 240 ^ high in a 5.5-mm-long
area of the widened inner portion of the sheath. Its ectal, cylindrical
portion is lined with longitudinal ridges. The absence of spines and the still
little differentiated glandular oviduct show that the animals are immature.
Discussion.—Eales (1960:364) included California in the range of A. juli-
ana, and therefore Lance (1961: 65) listed it. She later stated in a letter
(Beeman, 1963: 146) “that this record of A. juliana could not be traced.”
The species is the first representative of the subgenus in the Gulf.
A. (A.) reticulopoda Beeman (1960: 144), a seventh valid species of the
subgenus Aplysia (see Eales, 1960: 272, 354 ff.), has tentacles not con­
nected with the mouth as A. (A.) cedrosensis Bartsch & Rehder (1939: 2),
from the Pacific coast of Lower California, not from the Gulf, as Eales
(1960: 354) and Beeman (1960: 146) said. Branched penial warts occur
sometimes also in cedrosensis. The light color and other characters indi­
cated by Beeman allow for a specific separation of reticulopoda.
The ranges of A. dura, juliana, and nigra (Eales, 1960: 360, 363, 369)
show that geographic criteria cannot be decisive for the classification of a
sea hare, at least not yet. Therefore we have compared the present speci­
mens with the other species of the subgenus. A. depilans Gmelin, 1791, is
brown to greenish brown with large white or grey blotches, the mantle
speckled with brown, and the inner side of the parapodia blotched with
dark and light patches. The radula is of a simple type as in our animals,
but the cusps of the lateral teeth have a more elaborate denticulation.
A. (A.) dura Engel & Eales (1957: 87, 106; Eales, 1960: 360) is similar
to cedrosensis. The broad head and long neck, small parapodia, which do
not cover the mantle and anal siphon, distinguish A . dura from the present
material.
A. vaccaria Winkler (1955: 5) has fimbriated tentacles (Eales, 1960:
371) and at least 100 radular rows. Even considering that our specimens
158
are not quite adult, the difference between 100 and 63 cannot be ignored.
The cusps of the lateral teeth of vaccaria are longer than in any other
species of the subgenus. All pairs of ganglia, even the buccal ganglia, are
completely fused. The short pointed tail, some mottling on the inner side
of the parapodia, a black and white striped ctenidium, and a crested anal
siphon are characters of vaccaria, absent in our material, but their con­
stancy in vaccaria must be established by numerous specimens.
The type-specimen of A. nigra d’Orbigny (1837: 209) was not found
(Engel & Eales, 1957: 101), and no further specimen from the area of
Callao has been examined; Eales’ description (1960: 369) is based upon
material from Sydney and New Zealand. The species is darker than juliana,
and in this respect more similar to our present specimens, but its visceral
hump is more centrally placed than in juliana. The base of the rhachidian
tooth of nigra is very little expanded, and the cusps of the lateral teeth in
the middle of the half-row reach IV2 times the length of the tooth. Also
the formula 100 X 60.1.60 suggests that our material is better called juliana
than nigra. In Eales’ figures (1960: Figs. 46 e, f) the rhachidian tooth of
juliana is much more expanded than that of nigra, but this expansion varies
in juliana (Macnae, 1955: Fig. 7 a-c). The penis of A. nigra was not
drawn, so that its spatulate shape cannot be compared with the conical
penis with a spoon-shaped tip of juliana; the latter description (Eales,
1960: 366) agrees better with our material.
Due to its dark brown, almost black color and the black secretion of the
whitish purple glands a sea hare from the State of Rio de Janeiro (Marcus,
1958b: 43, juliana) has been called nigra by Eales (1960: 270). The color
of the secretion of the original material of nigra was milky white or slightly
violaceous; that of juliana is white. We have now re-examined the specimen
from Rio. Its radular formula is 83 X 41.1.41. The base of the rhachidian
tooth and the cusps of the lateral teeth conform with those of juliana. The
tip of the penis is pointed, not spatulate. As the penial warts have no
cuticular spines, the specimen, though 120 mm in length in preserved
condition, is not quite adult. Therefore the radula possibly does not have
its maximum numbers. The cerebral ganglia are contiguous, not fused as in
nigra. As long as juliana and nigra are maintained separated, we prefer
our first classification of the Brazilian specimen in question.
12. Stylocheilus longicauda (Quoy & Gaimard, 1824)
Figures 16-17
References.—Engel, 1936: 55; Marcus, 1963: 11; Farmer, 1967: 341.
Material.—Puerto Penasco, Sonora, Mexico. P. Pickens, July and August,
1964. “Found in the large tide pools during July and August only. They
appear just before Aplysia californica and A. juliana disappear from these
tide pools. During July and August they may be quite abundant, and are
usually found among the fronds of Sargassum.”
159
Further distribution.—Circumtropical; Farmer’s is the first record from
the west coast of the Americas. Original locality: New Guinea.
Remarks .—The photograph shows the specifically characteristic brown
longitudinal lines (“striae,” Engel), well compared with pencil strokes on
rough paper, and the blue ocellar spots surrounded by orange circles
(Fig. 16). The six present specimens have their colors preserved to a
different degree. The cutaneous appendages (villi) are, as usually, more
developed in the larger (23 mm) than in the smaller (8 mm) animals.
The radula varies with age and also among adult specimens. The radula
removed from a 22-mm animal has the formula 26X30.1.30, and agrees
with Engel’s figures.
Both the penis and its sheath are spiny. The spines of the sheath are
set on warts, as previously described and figured (Vayssiere, 1906: 80,
Fig. 12 bis; Engel, 1936: 55, Fig. 41 A; Eales, 1938: 90, Fig. 9). At
the ectal end of the basal dilated part of the sheath there is a number of
especially big spiny warts. In a 14-mm animal the sheath contains but
one row of warts bearing spines and some warts without spines. On the
penis of our larger specimens some of the spines are also borne on distinct
warts (Fig. 17), not yet recorded in the literature. Allusive warts occur
also on the penis of an animal from Eniwetok Island, Marshall Islands
(Dr. J. B. Burch coll., April 1960). In the above mentioned 14-mm slug
the penial spines are not borne on warts.
13. Berthellina engell ilisima, subspec. nov.
Figure 18
Reference.—MacFarland, 1966: 70 (B. engeli).
Synonym.— Pleurobranchus plumula (non Montagu) Bergh, 1894: 197,
PI. 9, Figs. 12-14, PI. 10, Figs. 1-8.
Material.— (1) Puerto Penasco, Sonora, Mexico. P. Pickens, November
21, 1964. “One of the most common opisthobranchs at Puerto Penasco;
only Aplysia californica may be seasonally more abundant. In late summer
there are no adults in the intertidal zone, just juveniles. The adults are
most common during the winter and spring and are almost always paired
during the spring, so that an overturned rock will usually reveal two side
by side.”— (2) Tiburon Island.— (3) Guaymas, July 1965.
Further distribution .—Gulf of California, 24° 11' N, 109° 55' W (original
locality), i.e. near Ceralvo Island, 18 m. Several localities in the Gulf of
California between tide marks (MacFarland, 1966: 71).
Description .—Alive the “apricot slugs” (Dr. Pickens) are up to 60 mm
long. They are deep orange colored; preserved they are white with traces
of pink. The digestive gland is brown. The notum contains white nodules,
probably glands.
160
F ig ure 18. Berthellina engeli ilisima, ssp. nov. A, shell; B, head; C, jaw
elements; D, radular teeth from middle row and near margin.
Of the four preserved specimens at hand the largest is 18 mm in length,
11 mm in width, and 13 mm in height. The sole is 12X8 mm; the tail is
covered by the mantle in the preserved animals; alive it projects behind.
The shell (Fig. 18, A) lies near the hind end. It is 4-4.5 mm long,
2-2.5 mm broad, longer and relatively narrower than that of Bergh’s speci­
men (2X 1.25 mm). It is flat, transparent, light brown, with an iridescent
periostracum. The larval shell stands out over the apical border as a white
knob. The growth lines are coarse. The posterior half of the shell bears
longitudinal radiating rows of dots which produce a spiral sculpture. There
are no spiral lines on the anterior half of the shell. In one of our four shells
the longitudinal lines reach farther forwards, but not to the anterior border.
The sides of the veil are auriculate. The rhinophores are joined for about
half their length, rolled in, and have the eyes at their bases (Fig. 18, B).
In one well extended specimen the ctenidium is about V3 of the body
length. Its smooth rhachis bears about 20 leaflets on either side, nine of
which lie behind the insertion of the branchial membrane. As the posterior
pinnules are smaller than the anterior ones, the membrane fastens about
2A of the length of the ctenidium. The anus lies on the edge of the
membrane. The orifice of the Bourne’s, or prebranchial, gland opens
between the genital apertures and the root of the gill.
The tips of the mandibular platelets are pointed scales without denticles
(Fig. 18, C). The radula (Fig. 18, D) comprises about 90 rows with
about 250 elongated teeth to the half-row. The highest teeth (0.3 mm)
stand in the middle of the half-row and bear 12-16 denticles on their
161
posterior edge; the outermost teeth are smaller and have less denticles.
For the reproductive organs we refer to MacFarland 1966: PI. 16,
Fig. 9.
Discussion.—The Pleurobranchacea contain the shelled Pleurobranchidae
and the shell-less Pleurobranchaeidae. The former comprise the Pleuro-
branchinae with a gland at the hind end of the sole and a tuberculated or
granulated ctenidial rhachis, and the Berthellinae without a pedal gland at
the hind end and with a smooth or transversely grooved rhachis of the gill.
The branchial leaflets are placed alternately in two genera of the Berthel­
linae: in Berthella Blainville, 1825, and in Berthellina Gardiner, 1936.
In Berthella the teeth are short, hook-shaped, (unciform) and smooth, in
Berthellina elongated like the blade of a knife (lamelliform) and serrated
on their posterior edge. To this basis (Odhner, 1939: 15-20) Burn
(1962b) added Berthellinops with paired branchial pinnules, unciform
teeth and transversely grooved rhachis.
Two species of the Berthellinae, one with unciform, the other with
lamelliform teeth occur in British and Norwegian waters (Engel, 1934:
584; Odhner, 1939: 16-17), so that Pruvot-Fol’s restriction of Berthellina
engeli to the Mediterranean Sea (1954a: 227) cannot be accepted.
Bergh’s above-cited specimen, the four animals at hand, and MacFar-
land’s B. engeli evidently belong to one and the same species, though
Bergh could not indicate the live color and did not detail the sculpture of
the shell. His name, Pleurobranchus plumula Montagu, 1803, has been
accepted by MacFarland (Steinbeck & Ricketts, 1941: 542). Today
Bergh’s and Vayssiere’s Pleurobranchus or Berthella plumula must be
given the name Berthellina engeli Gardiner, 1936. If also Bergh’s specimen
from the Gulf belonged to engeli, the range of this species would reach to
the eastern Pacific Ocean. Bergh himself (1894: 199; 1898: 123) was
astonished at such a discontinuous distribution, and Pilsbry (1895-96:
195) added two question marks to this reference.
Our material differs from the East Atlantic and Mediterranean engeli
by the shape of the shell and by the spiral sculpture. In engeli the shell is
convex and auriculate, in our animals flat and spatulate. The longitudinal
rows of dots reach the anterior border of the shell in engeli, while they do
not in our shells. Generally the longitudinal lines are restricted to the
posterior half in our shells.
MacFarland (1966: 72) evidently had partially decalcified shells at
hand; his drawing (PI. 13, Fig. 14) is a copy of the original figure (Bergh,
1894: PI. 9, Fig. 12). MacFarland found the longitudinal lines much
less clearly defined than the lines of growth, but traceable to the shell
margin, and did not mention that they actually are rows of dots.
We must also consider, whether really B. engeli or perhaps another
species is closest to ours. B. edwardsii (Vayssiere, 1896: 122; 1898: 265),
subsequently spelled edwardsi by Vayssiere (1902: 223) and others, has
162
been united with Bergh’s and Vayssiere’s plumula, the B. engeli of present
usage (Odhner, 1932: 29). Odhner did not return to this idea when he
looked (1939: 18) for a name prior to engeli. Both species are recorded
from the warm temperate eastern Atlantic Ocean; farther north, and in the
Mediterranean, only engeli has been found. The color of living edwardsi
is not known; that of engeli varies from pale yellowish and lemon to
vermilion and apricot (Gantes, 1956: 262). The auriculate and convex
shell is the same in both species (Odhner, 1932: 28), and in both the
entire shell has longitudinal striae, though these are a little weaker in
edwardsi (Vayssiere, 1898: PL 17, Fig. 35) than in engeli (Vayssiere, 1880:
208, PL 7, Fig. 2, 2a; 1898: 18, Pl. 17, Fig. 17; 1913: 217; 1932: no
page number; Odhner, 1939: 18). The shells of edwardsi are 6-15 mm
long, 3.5-11 mm broad; those of engeli 3.8-7 mm long, 2-4.5 mm broad.
This is not a clear-cut difference, but seems to be sufficient for the separa­
tion of edwardsi and engeli. Our animals and Bergh’s specimen from the
Gulf of California correspond better to the measurements of engeli.
White (1955: 174) classified a specimen from the coast of Angola
(7° 16' S, 12° 47' E, off Ambrizette) as edwardsi due to the fact that the
rhinophores were joined for about half their length. Our material shows
the same character, but Odhner’s discussion (1932: 29) reduces its value.
Also the number of radular teeth would favor an approximation of our and
Bergh’s material from the Gulf to edwardsi, if Burns’s key (1962b: 134)
is considered. But there are also records of 200, 225, and 230 teeth in the
half-row of engeli (Bergh, 1892b: 23; Odhner, 1932: 29; Gantes, 1956:
262).
Vayssiere (1902: 226) observed a concentration of the central ganglia
more pronounced in his plumula (— engeli) than in edwardsi, but this
difference is difficult to distinguish precisely.
The Caribbean species B. quadridens (Morch, 1863), B. circularis
(Morch, 1863), and B. amarillia (Mattox, 1953), which we recognize as
distinct (Marcus, 1962: 465), have denticulated mandibular platelets, and
so are more easily distinguished from engeli ilisima than the species from
the eastern Atlantic Ocean. B. quadridens was recently collected on the
Pacific coast of Panama by Professor Dr. Frederick M. Bayer.
The valid Indo-Pacific species of Berthellina, whose range extends from
the Gulf of Suez to Hawaii, were briefly diagnosed by Burn (1962b: 134).
One or more of the characters mentioned for each of them show that
engeli ilisima cannot be one of these species.
14. Pleurobranchus (Pleurobranchus) areolatus (Morch, 1863)
Figure 19
References .—Bergh, 1897: 111; Abbott, 1949: 73-78 (atlanticus); Marcus,
1962: 466.
Material.—Puerto Penasco, Sonora, Mexico. P. Pickens, November 1963
163
and 1964. “The species is not very common, but two or three animals may
be found in the same locality. They give off a very strong iodine odor when
kept in aquaria.”
Further distribution.—Florida; Caribbean Islands southward to Barbados
and Curasao. Canal Zone, Pacific coast of Panama. Original locality:
Virgin Islands, St. Thomas.
Description .—The two photos show rather flat animals, 85 mm long, whose
brick-red notum covers the body. The dark red rhinophores project from
the anterior notch. Pointed tubercles, larger in the middle of the notum
than on the sides, are lighter than their interspaces. Chalk-white specks
are irregularly strewn over the notum, chiefly on the tubercles. The foot
is rather uniformly pink.
The biggest of the three preserved animals is 63 mm long. It is the
largest specimen hitherto known. The sculpture consists of round, promi­
nent reddish areas with a light papilla in the middle. These areas are 3 mm
in diameter in the middle of the back, decreasing outwards to 1 mm. The
skin between the areas is pigmented with black; it contains vestiges of
calcareous inclusions. The gill has 25 leaflets on either side; the branchial
membrane extends to the 17th leaflets, dorsally to which lies the anus. The
brown shell is 5.5 X 3.5 mm; the growth lines are very coarse.
The mandibular platelets have 1-5 denticles on either side. The radular

F ig ure 19. Pleurobranchus areolatus (Morch); diagram of reproductive organs.

164
formula is 90 X 230.0.230. This is the highest number of teeth recorded for
the species and tallies with the large size of the present specimen.
Of the previous descriptions of the reproductive organs (Fig. 19)
Lloyd’s (1952: 80, Fig. 15) for P. tuberculatus Meckel, 1808 ( mem -
branaceus Montagu, 1815) agrees best with our findings. The coiled
ampulla (a) divides at its outlet. The male branch becomes folded and
prostatic ( q ) immediately at its beginning. An external furrow separates
the ental third of the prostate from the two ectal thirds, as drawn by
Abbott (1949, PI. 5, Fig. 6, PR). Bergh (1897: 113) said “without
prostate,” but the thickening of the male duct due to the prostatic glands
varies, hence the occurrence of an intercalary prostate is disputed in
many opisthobranchs. In the present specimen the soft duct that comes from
the folded prostate is glandular too. These glands end abruptly, and the
next section ( d ) is narrower and muscular. Its coils straighten in the
outermost portion, the thick-walled ejaculatory duct. This pierces the
conical and slightly curved penis (/?), whose wall is loosely muscular and
more or less swollen at its base. In all three specimens the penis projects
from the cutaneous flap (/) which surrounds the genital apertures.
The female opening (aw) is that of the nidamental duct which communi­
cates with the vagina a little farther entally. The entrance of the vagina
(v) is thickened by strong musculature, and also its farther inward course
is muscular. Before the vagina bifurcates it emits the heavily ciliated in­
semination duct (in), the “conduit” or “canal de Cuvier” in Pruvot-Fol’s
terminology (1960:188). The vaginal bifurcation leads into two wide,
ovoid vesicles, the spermatheca (s) and the spermatocyst (sc). The latter
is defined by the orientated spermatozoa which it contains; the former
contains debris as usual. The insemination duct enters the outlet of the
ampulla (a) which is fixed to the gland mass (g). The pervious canal,
the inner oviduct (o) of Figure 19 is interpreted. We cannot affirm that
the eggs pass from the ampulla to the gland mass, the glandular oviduct.
We found many eggs, about 70 ^ in diameter, in the insemination duct.
This may be their normal way out to the gland mass. It might also
represent a surplus of female germ cells conveyed to the spermatheca for
being digested.
Discussion .—Steinbeck & Ricketts (1941: 542-543) discuss the literature
referring to the Pleurobranchacea of the Gulf and their own collecting
results. Their first item “Berthella plumula (Montagu) 1803” is MacFar-
land’s Berthellina engeli, here called B. engeli ilisima.
Their next paragraph refers to “Pleurobranchus digued Rochebrune
1895.” This is a pleurobranchid, scarlet above, buff-white below, which
the author compared with P. patagonicus d’Orbigny, 1837, whose northern­
most record is from Middle Chile (20° 19' S). As this species was allotted
to Bouvieria Vayssiere, 1896 ( = Berthella Blainville, 1825), Odhner
(1926: 22) assigned digueti to this genus. Vayssiere (1898: 345-46)
165
mentioned digueti, but did not examine the typical material. Pilsbry
(1895-96: 201) received an animal from the original locality of digueti,
the area of La Paz, and described it under the name proposed by Roche-
brune. It is, in fact, a Pleurobranchus with a tuberculated rhachis of the
ctenidium. Oldroyd (1927: 52) repeated Rochebrune’s and Pilsbry’s
descriptions, but repeated the original description (p. 738) under the
name Lamellaria digueti Rochebrune, 1895. The words Utentaculis canali-
culatis” exclude Lamellaria. MacFarland (1966: 75) described as P.
digueti Rochebrune, 1895, two preserved specimens up to 33 mm in
length. Steinbeck & Ricketts had taken them at Cape San Lucas and near
the original locality of digueti. The location of the anus is not quite the
same in Pilsbry’s and MacFarland’s specimens, and also the platelets of
the jaw, smooth (Pilsbry) and denticulate (MacFarland), disagree. In
our report on a large material of P. areolatus in Part I, none to eight
denticles on either side of the jaw platelets are indicated.
A d d it io n s to t h e S y s t e m o f t h e D o r id id a e
When we compared the two new genera of this family with Thiele’s
synopsis (1931: 430-439), we had also to consider the modifications and
additions that have been published since then. An extract of our notes,
even without arguments, may be useful for other workers too:
Awuka Marcus, 1955: 155, probably Aldisinae, not Kentrodoridinae, as
originally indicated.
Boreodoris Odhner, 1939: 31, Aldisinae ( Thorunna, eliminated).
Cadlinella Thiele, 1931: 431, Cadlinellinae Odhner, 1934: 249.
Conualevia Collier & Farmer, 1964: 381, Conualeviinae, id. ibid.
Erythrodoris Pruvot-Fol, 1933: 131, possibly Gruveliinae.
Felimare Marcus, 1967, Chromodoridinae.
Glossodoridiformia O’Donoghue, 1927a, 87: placed in the Doridinae, but
reproductive organs unknown.
Guyonia Risbec, 1928: 102 (1953: 41), Doridinae; in 1928 Risbec called
the radular cartilage “machoires” (Risbec, 1953: 35, 41).
Inuda Marcus, present paper, Inudiinae.
Jorunna, Kentrodoridinae Odhner, 1939: 26.
Kentrodoris, Kentrodoridinae Odhner, 1939: 26.
Lissodoris Odhner, 1934: 245, Cadlinellinae, ibid.: 248.
Neodoris Baba, 1938: 13, between Doridinae (Doris) and Discodoridinae
(Anisodoris).
Otinodoris White, 1948: 203, Archidoridinae.
Risbecia Odhner, 1934: 249, Cadlinellinae, id. ibid.
Rosodoris Pruvot-Fol, 1954b: 23, Chromodoridinae.
Sebadoris Marcus, 1960b: 904, Discodoridinae.
Siraius Marcus, 1955: 134, Doridinae.
Spongiodoris Pruvot-Fol, 1933: 131, incompletely known.
166
Taringa Marcus, 1955: 152, Discodoridinae.
Tayuva Marcus, present paper, Discodoridinae.
Thorunna, Actinocyclinae Odhner, 1939: 26.
15. Conualevia mizuna, spec. nov.
Figure 20
Material.—Puerto Penasco, Sonora, Mexico. Mary Anne Hill, February
5, 1966. Found under a rock in the rocky intertidal zone. One specimen.
Description .—The living animal in the color photograph is 16 mm in
length, white with quite light yellow rhinophores and gills. The measure­
ments of the preserved specimen are: 11 X 8.5 X 5 mm. The sole is 4 mm
in width, the hyponotum 2-2.5 mm.
The notum is smooth, somewhat swollen; on both sides scattered trans­
parent glands of different size are seen in the stained, clarified specimen
(Fig. 20, A) but not in the photograph, contrary to Collier & Farmer’s
Plate 3 (1964: 387). The rhinophores are smooth (Fig. 20, B), not
perfoliated and not ridged around the pits. Traces of spicules are not
recognizable. The eight gills are pluripinnate and also without a ridge
around the pit. The tentacles are broad and short (Fig. 20, C), not set off
from the oral area. The foot is bilabiate in front, not notched. The
gonopore lies slightly in front of the middle of the right side.

F ig ure 20. Conualevia mizuna, sp. nov. A, clarified slug; B, rhinophores of

preserved slug; C, fore end of same; D, diagram of reproductive organs.
167
 The labial cuticle is thin and smooth. The radula measures 2.7 X 2.0
mm; it comprises 35 rows with 64 teeth per half-row. The rhachis is naked.
The pleural teeth are all simple hooks; the innermost is 50 ^ in height,
the biggest 85 the outer teeth decrease again. A voluminous free
stomach lies over the intestinal gland.
The long and coiled ampulla (a) divides within the female gland mass
(g). The emerging male duct immediately becomes thick and glandular
( q ), followed by a sheathed section (ss)? and ends with an acrembolic
penis. The vagina begins wide and narrows inwards. The location of the
large spematheca (s) and the smaller spermatocyst (sc) corresponds to
the vaginal type (Odhner, 1926: 51).
Discussion.—The smooth rhinophores and the rhinophoral and branchial
pits without a surrounding ridge characterize the species as belonging to
Conualevia Collier & Farmer (1964: 381). The type species, C. marcusi,
from the Gulf of California, has 16 unipinnate gills and 97 teeth per
half-row of the radula. The second species of the genus, C. alba, from
Newport Bay to Bahia Tortuga (Lance, 1966: 75), is similar to C. mizuna
by its 8 tripinnate gills and 56 teeth in the half-row. However, its gills and
rhinophores are white, the latter ringed by hundreds of small black dots;
the notal edge bears a single row of opaque white glands, and the “prostatic
part of the vas deferens is poorly differentiated and is almost nonglandular.”
16. Cadlina evelinae Marcus, 1958
References .—Marcus, 1958a: 18; Collier & Farmer, 1964: 380.
Material.— 1. Puerto Penasco, Sonora, Mexico. P. Pickens and Mary Anne
Hill, November 1964 and May 1965.—2. Guaymas, San Carlos Bay.
August 1964. “Not very common on the fronds of Sargassum , usually in
rocky intertidal tide pools.”
Further distribution .—Brazil, coast of Sao Paulo, under stones in the
upper littoral (original locality). Gulf of California, Coloradito, 25 km
north of Puertecitos, and Angel de la Guarda Island. Pacific coast of
Lower California, Sebastian Viscayno Bay.
Descriptive notes.—The animal of the photo is white with orange dots on
the dorsum which have faded out in preservation. In this condition brown-
staining glands in the border of the notum appear irregularly depending
upon their degree of fullness.
The five slugs at hand are 5-7.5 mm in length, 3-5 mm in width. Antero-
external grooves of the stout tentacles show that the tips are contracted.
The rhinophoral clubs have about 16 leaves, the rim of the pits is slightly
undulate. There are 6-9 gills, essentially unipinnate, but in part bipinnate.
The edge of the branchial pocket is smooth. The tail is pointed, the anterior
pedal border bilabiate, its upper lip entire or slightly notched.
168
 The labial elements are hooklets with bifurcate tips. The radula has
74-81 rows with 76-80 lateral teeth. The cusp of the rhachidian tooth
bears generally four, sometimes five or six denticles. In the middle of the
half-row there are 1-3 fine denticles, farther outward three stronger ones.
The ejaculatory duct is not cuticularized. Also the two convoluted
accessory glands agree with the original material.
Remarks.—The occurrence of C. evelinae in the Gulf of California, and
that of C. sparsa Odhner, 1921, from Juan Fernandez and the north coast
of Chiloe at San Diego (Marcus, 1961a: 15) make it advisable to survey
the nine species of Cadlina that occur in American Atlantic and Pacific
warm-temperate and temperate waters. Not all their color patterns in life
are known, and could not be used in the following key. The yellow tones,
frequent in Cadlina, fade out in preservation. Collier & Farmer (1964:
380) stressed the variability of the denticles on both rhachidian and lateral
teeth. Also the ramification of the branchial plumes and the notch in the
upper lip of the anterior foot border are variable characters in Cadlina.
Even the hooks of the ejaculatory duct appear, at least in C. sparsa
(Odhner, 1926: 56), only in fully mature specimens. These restrictions
should be considered when the key is used for identification. Living animals
from the Pacific coast of North America can be determined with Lance’s
list (1962a: 157) and Collier & Farmer’s description and figure (1964:
380) of C. evelinae.
1. Notum with a single series of marginal spots.......................................... 2
1. Notum without spots, or with scattered spots, the marginal ones
often larger than those in the cen ter........................................................4
2. Notum rough; rhinophores dark, conspicuous against the ground
color..................................................................................................................3
2. Notum smooth; rhinophores of the same color as the ground or a
shade darker.................................................................juvenca Bergh, 1898
3. Teeth in outer half of half-row saw-like......................................................
.................................................................flavomaculata MacFarland, 1905
3. Teeth in outer half of half-row hooks with pectinate tip s ......................
....................................................................................... rumia Marcus, 1955
4. Notum with blackish-brown dots or dark ocular spots with light
center............................................................................sparsa Odhner, 1921
4. Color of notum opaque white, yellow or orange, no ocular spots.........5
5. Rhinophores and gills dark brown, almost b lack ......................................
................................................................................ limbaughi Lance, 1962
5. Rhinophores and gills of the same color as the dorsum; the bases
of the rhinophores sometimes darker........................................................6
169
6. Ejaculatory duct with cuticular hooks 7
6. Ejaculatory duct unarmed...................... 8
7. Rhinophores with 16-18 leaves; six gills; 40-50 lateral teeth per
half-row........................................................ marginata MacFarland, 1905
7. Rhinophores with 25-30 leaves; 12 gills; 31 lateral teeth per
half-row....................................................................... berghi Odhner, 1926
8. Radular rows 108; 53 lateral teeth per half-row......................................
.............................................................................. scabriuscula Bergh, 1890
8. Radular rows 74-81; 65-80 lateral teeth per half-row.............................
...................................................................................evelinae Marcus, 1958
17. Chromodoris norrisi Farmer, 1963
Figures 21-24
Reference.—Farmer, 1963: 81.
Material.— 1. Puerto Penasco, Sonora, Mexico. P. Pickens and Robert
Taylor, August 1964 and March 1965.—2. Guaymas, July 1965.—3. Ibid.
Alex Kerstitch, March 12, 1966. Found crawling about on gorgonians at
a depth of 20 m.
Further distribution.—Pacific coast of Lower California. Cedros Island
and Puerto Rompiente. West coast of the Gulf of California, Ceralvo Island
(original locality), Bahia de los Angeles and San Luis Gonzaga Bay.
Descriptive notes.—The photograph of a specimen from locality 1 shows
a whitish ground color which is darker in front of the gills, evidently due
to the underlying viscera. The notum bears dark red spots, larger and less
numerous than in Farmer’s figure (PI. 1 a), three rows of apricot spots,
also less numerous than the yellow ones of the original material, and a
few light red dots (Fig. 21). Three of the four slugs received, which do
not include the animal of the photo, correspond to its color pattern, the
fourth (locality 2) resembled the original figure, but faded out later on.
The notum, edged with cadmium-orange dashes and blotches, and the tips
of the rhinophores and gills of the same color, agree in the present and the
original material. Also the often branched tips of the gills occur both in
the original and the present specimens.
The anterior end of the foot is bilabiate, the short hind end pointed.
The length of the slugs at hand is 6, 8, 9, and 20 mm. The width of this last
specimen (locality 2) is 9.5 mm, its height 8 mm; the breadth of the sole
2.5 mm. The border of the notum contains big glands, not only behind.
The buccal armature of one of the large-spotted specimens was exam­
ined, in order to compare it with that of the small-spotted animals of
Farmer. The labial elements (Fig. 22) are curved hooks with bifid tips.
The radula (Fig. 23) has 65 rows and 44 teeth per half-row. The rhachis
170
F ig ures 21-23. Chromodoris norrisi Farmer. 21, living slug from locality 1,
drawn from color photo; 22, labial rodlet; 23, radular teeth.
bears a short pseudo-tooth with a minute cusp. The innermost lateral tooth
has five outer and five inner denticles. In the middle of the half-row the
teeth bear 8-12 outer denticles. Also the outermost teeth have denticles
which make their cusps pectinate. By this radula also the large-spotted slugs
prove to be norrisi.
An accessory gland on the genital vestibulum is not developed in the
20-mm slug.
In the color photos of specimens from locality 3, three slugs are seen
which were about 46 mm in length and 20 mm in width (Fig. 24, A).
They differ from the original figure by a broad, irregularly undulated notal
edge, which seems to be permanent. The color elements are similar to those
of the original material, but the pattern is different. The borders of the
notum and the foot are milky white. The center of the notum is light yellow,
irregularly sprinkled with red dots. Larger red spots and an interrupted
yellow line occur along the notal border and on the sides of the foot. The
white rhinophores have a red band near the tip. The tips of the white gills,
and longitudinal marks on the branchial stems, are dark red.
The two specimens received in formalin had preserved the red spots on
the notum, the hyponotum, the foot, and on the upper lip of the anterior
pedal border. The center of the notum is covered with yellow specks.
In alcohol the red elements are dissolved, while the yellow ones are
preserved. As in the original material the yellow areas are somewhat
swollen in all specimens of Dr. Pickens’ collection.
171
 Further characters of the preserved material from locality 3 are: Length
40 mm; width 28 mm; gelatinous, without spicules; hyponotum about
8 mm; upper lip of foot border slightly notched; genital aperture rather
far in front, about 12 mm from the tip.
Bases of tentacles broad, tips long and pointed, grooved outside; border
of rhinophoral pits smooth; number of foliations 25; gills 18, about 9 on
either side, unipinnate, but many tips branched. Anus in the middle of the
posterior circle of the gills.
Bifid labial hooks; radula 82 rows, 65 teeth, rhachis naked (Fig. 24, B).
Innermost lateral tooth 3-4 inner, 4-6 outer denticles; all following teeth
with outer denticles, 4-5 increasing to 8-10; even the penultimate tooth
bears denticles. Stomach covered by liver.
Inner part of efferent duct glandular (Fig. 24, C, q ); its convolutions

Figure 24. Chromodoris norrisi Farmer. Material from locality 3: A, living

slug drawn from color photo; B, radular teeth; C, diagram of reproductive
organs.
172
tightly packed together forming a voluminous prostatic mass. Female
vestibulum with an annexed gland.
An undulated notal border, a greater number of gills, a naked rhachis,
and a vestibular gland separate the minutely spotted animals from locality
3 from the specimens from localities 1 and 2. The width of the notum of
the original material is difficult to judge; its vestibular gland was not
examined. In order to make a future separation possible, perhaps by
different spawns (Gohar & Aboul Ela, 1957: 220-221), the material from
locality 3 was described separately, but a specific name is considered as
premature.
Remarks.— Chromodoris amoena Cheeseman, 1886, from warm waters of
North New Zealand, erroneously transferred (Powell, 1937: 121) to
Ceratosoma , is related to norrisi, but its minimum of 77 teeth per half-row
(Odhner, 1934: 249) considerably exceeds Farmer’s and our maximum of
65, and the denticles of the teeth are less numerous.
18. Chromodoris banksi sonora, subspec. nov.
Figures 25-29
Material.— 1. Puerto Penasco, Sonora, Mexico. P. Pickens, March, June,
August 1964, 1965; April 2, 1966. “Other specimens have been found in
November. This is the second most common doridid found at Puerto
Penasco. It is found in the lowest part of the rocky intertidal zone, quite
often in pairs.”—2. Guaymas, July 1965.
Further distribution.— Chr. banksi banksi Farmer (1963: 84) was collected
in December, January, and March in three different localities along the
west coast of the Gulf, from Puertecitos (30° 24' N, 114° 40' W; original
locality) to Angel de la Guarda Island (29° 33' N, 113° 35' W).
Description of material of 1964-65. —The photographs of a slug 30 mm
long show a brownish-greyish ground densely set with dark spots of differ­
ent sizes (Fig. 25). Especially on the sides many spots are larger. Red and
yellow notal papillae are glands, as the preserved material proves. Also
the hyponotum is spotted. One photograph and some of the preserved
specimens have several yellow hyponotal spots among the black ones. The
borders of the rhinophoral and branchial pockets, the tips of the rhino­
phores and the gills, the borders of the notum and the sole are orange.
The marginal orange stripe is accompanied by yellow on either side.
In preservation the colors are as follows: ground color grey, with black
dots and yellow glands, reddish rhinophores with a yellow border of the
rhinophorial pocket, grey gills with black and yellow elements and their
pouch with a yellow rim; notal border with two lines, the outermost
colorless and the inner a yellow stripe.
The biggest of the 13 slugs at hand is 25 mm long, 11 mm broad, and
173
10 mm high; the smallest is 13 mm in length, 6.5 mm in width, and
3.5 mm in height.
The tentacles are grooved on their outer side, evidently due to contrac­
tion of their tips. The rhinophores bear about 20 leaves. The unipinnate
gills, whose total is 27-34, surround the high anal papilla. There are 14-15
large plumes, whose size decreases backwards, where a number of smaller

F ig u r e s 25-29. Chromodoris banksi sonora, ssp. n ov. 25, living slug from
color photo; 26, gills; 27, jaw elem ent; 28, radular teeth; 29, diagram o f
reproductive organs.
174
ones form two spirals (Fig. 26). Asymmetrical defects occur, though only
nine gills (Farmer, 1963: 84) were not observed in the present material.
The anterior border of the foot is bilabiate. The sole is narrow. The hind
end is pointed and projects backwards.
The yellow labial plates (Fig. 27) bear curved hooklets with bifid tips.
The height of the labial elements reaches 50 the diameter 4 The
radula (Fig. 28) is 6X1.5 mm; it has 150 rows. The half-rows contain
38-41 lateral teeth. A rhachidian pseudo-tooth is present; its base measures
34 /A, the cusp 20-26 fi. The innermost lateral tooth, whose base is
50 n in height, bears a 32 ^ long cusp with 3-7 inner and 7-8 outer denti­
cles. In the following lateral teeth the single cusp reaches 80 ^ and bears
7-10 outer denticles. In the outer third to fourth of the half-row the teeth
have no denticles.
The spermoviduct (Fig. 29, h) bifurcates within the female gland mass
(g). The male duct ( d ) begins straight, then forms thin, glandular coils
( q )y has a muscular sheath in the following section (ss), and bears a
broad, penial papilla ( p ) in its ectal course. Two retractor muscles (r)
insert on this papilla. The common atrium (ac) receives the nidamental
duct (ni) beside the ejaculatory duct ( e ). The vagina (v) begins farther
inwards. It leads to the thin-walled spermatheca (s) and the tubular, curved
spermatocyst (sc). These seminal receptacles lie one beside the other at
the inner end of the vagina, hence corresponding to the vaginal arrange­
ment described by Odhner (1926: 51). Also the insemination duct (in)
goes out from the inner end of the vagina and enters into the gland mass
near to the ingress of the spermoviduct. A vestibular gland (vn) is
connected with the common atrium by a wide duct.
Material of 1966 .—In life the 15-mm-long slug “lacked the red trim on
the border of the mantle and the red dots on the body. The rhinophores
and gills were red-tipped, however” (Dr. P. Pickens). The preserved animal
was a brilliant purplish red with numerous black and some orange spots on
the notum. The tips of the rhinophores and gills were orange red. There
is a total of only 12 gills; probably the branchial tuft is in regeneration.
The radula agrees with that of the material of 1964-65.
Discussion .—The general character of the color pattern, the radular for­
mula, and the shape of the rhachidian pseudo-tooth approximate the
present material to Chr. banksi, whose reproductive organs, however, were
not described. The gills of banksi banksi and banksi sonora differ, as reveal­
ed in the preceding description. A second, possibly more weighty difference
refers to the first lateral tooth of b. banksi, figured without denticles
(Farmer, 1963: Fig. 1, j). Perhaps the figure is a back view of this tooth.
Less important differences are: the cusps of the inner lateral teeth of
b. banksi are shorter and bear fewer denticles than in b. sonora; the borders
175
of the branchial and rhinophoral pockets of b. banksi are not colored;
and its ground color is pure white.
Relatively short cusps of the inner lateral teeth as in Chr. b. banksi occur
in Chr. dalli Bergh (1879a: 109), a related species from Puget Sound.
The total of 15 branchial plumes, and the rhachidian pseudo-tooth divided
into two halves (PL 13, Figs. 11, 12) by a longitudinal groove, separate
dalli from banksi sonora; the rhachidian tooth differs also from banksi
banksi.
The radula and the gills of banksi sonora agree well with Chr. punctilu-
cens Bergh (1890a: 163) from the Straits of Florida and the Canary
Islands (Odhner, 1932: 33). Along the notal border, around and on the
rhinophores and gills, and on the hyponotum punctilucens has more black
elements than the present slugs. Its insemination duct originates far from
the inner end of the vagina (Odhner, 1932: Fig. 5), and a vestibular
gland is not developed (1934: 251).
19. Hypselodoris californiensis (Bergh, 1879)
Figures 30-33
References.—Bergh, 1879a: 112; 1894: 181; O’Donoghue, 1926: 211
(literature); 1927a: 90; MacFarland, 1966: 157; Lance, 1966: 72;
Farmer, 1967: 341.
Synonyms.— Chromodoris universitatis Cockerell, 1901b: 79, according
to Cockerell & Eliot, 1905: 37; Chromodoris agassizii Bergh, 1894: 182.
Material.— 1. Puerto Penasco, Sonora, Mexico. P. Pickens, March, June,
July, and November 1964-1965. “It is the third most abundant doridid,
after no. 3 (Chromodoris banksi sonora) and no. 4 (Casella sedna), at
Puerto Penasco. It is a rocky intertidal form usually found singly.”—2.
Puerto Libertad, Sonora, Mexico. Joy Cooper, May 30, 1965. One
specimen.— 3. Guaymas, July 1965.
Further distribution.—Monterey to Angel de la Guarda Island (Farmer
& Collier, 1963: 62), Punta Penasco (MacFarland, 1966: 162), and the
Bay of Panama; rare at northern end of range (MacFarland, 1906: 130;
Steinberg, 1963b: 69). Original locality: Santa Catalina Island, California.
Description.— In life, the slugs (Fig. 30) are 32 mm long. They are dark
ultramarine blue with golden yellow spots on the dorsal surface and the
hyponotum, and have a lighter blue, not spotted, sole. In the specimen from
the second locality (Fig. 31) the notum is set off from the foot by an outer
whitish and an inner light blue line. A labeled photo received by courtesy
of Dr. James R. Lance from LaJolla shows one light blue line. In Dr.
Lance’s specimen the golden yellow elements are a few longish streaks;
in front and behind they are shorter, more spot-like. The rhinophores of
the Sonoran slugs are dark blue. The gills are dark blue, sometimes with
yellow dots, in the animals from the first locality; in the specimen from
176
F ig ures 30-33. Hypselodoris californiensis (Bergh). 30, living slugs from
color photo, locality 1; 31, living slug from color photo, locality 2; 32, jaw
elements; 33, radular teeth.
Puerto Libertad they are whitish and tipped with dark blue. In Bergh’s type-
specimen the roots of the branchial leaves were silver-white.
In preservation part of the blue pigment has passed into the surrounding
wet cotton and the label, as was observed in the original material, in
MacFarland’s (1966: 158), and in other blue Chromodoridinae (Mac­
Farland, 1906: 129; Anderson & Lane, 1963). The yellow spots of the
smallest present specimens have faded out to white.
The largest of the six slugs from Puerto Penasco is 30 mm long, 10 mm
high, and 8 mm broad; the corresponding measurements of the smallest
animal from this locality are 17.5, 4, and 4 mm. The tentacles are rolled in
(locality 2) or appear as short cylinders (locality 1), which end with a
groove due to the contraction of the tip. The rhinophoral clubs have 17
leaves. There are 8-10 unipinnate gills. Of the semiglobular glands in the
mantle border, frequently mentioned in descriptions of the Chromodoridi­
nae, six or more under the hind end of the notum are most conspicuous,
but there are also smaller ones on the level of the rhinophores. The anterior
border of the foot is transversely grooved.
177
 The labial plates (Fig. 32) are yellow triangles; their elements are
unicuspidate hooklets. In the material from locality 1 they are about 50 ^
in length and occasionally have secondary points. Near the borders of the
labial plates they are shorter and broader. The latter type of labial hooklets
is the only one in the slug from the second locality. The radula (Fig. 33)
is blue and composed of bifurcated hooks, whose outer or posterior prong
is denticulated, especially in the middle of the half-row. The anterior prong
of the innermost tooth bears a single denticle on the inner side, present in
the material from Puerto Penasco and Puerto Libertad, but not mentioned
in the previous descriptions of califor niensis and the synonymized species.
The outer prong of the hook is shorter than the inner one, especially in the
middle of the half-row. The radula from locality 1 consists of 70 rows
with 93 teeth per half-row. The innermost tooth, 66 /x in height, has cusps
about 30 n long. In the middle of the half-row the inner cusp reaches
57 /jl\ the ten outer teeth are shorter. The small slug from the second
locality, 12 mm long in preservation, had 51 rows and 85 teeth in the
half-row.
Remarks.— The naked mid-line of the radula is limited by the converging
bases of the adjacent teeth. The illusion of a triangular rhachidian pseudo­
tooth (O’Donoghue, 1927a: 91) is produced by focusing only the deepest
layer of the radula.
Chromodoris agassizii from the Bay of Panama (Bergh, 1894: 182) has
the same color pattern and radula as calif or niensis. The size of the single
specimen is the same as that of the present slug from the second locality,
and also the labial elements of both agree. Therewith no discriminatory
character remains.
Pruvot-Fol (1951: 152) approximates Chr. macfarlandi Cockerell,
1901, to Chr. universitatis Cockerell, 1901 ( —californiensis). Chromo­
doris macfarlandi, however, has a radula of what we call the Chromodoris-
type (MacFarland, 1966: PL 34, Fig. 1-5), hence differs widely from
Hypselodoris californiensis.
In the two similar, though striped, Caribbean species of Hypselodoris,
H. edenticulata (White, 1952) (see Marcus, 1967b) and H. sycilla (Bergh,
1890a: 165), the half-row of the radula has 169-246 and about 290
teeth, respectively. In californiensis the maximum is 132.
20. Casella sedna, spec. nov.
Figures 34-37
Material.— 1. Puerto Penasco, Sonora, Mexico. Rocky intertidal zone.
P. Pickens, July, August, and November 21, 1964. 20 specimens.—2.
Guaymas, July, 1965. “The species is the most common nudibranch at
Puerto Penasco. It is present throughout the year, but may be locally
abundant during August.”
178
Description .—The color alive, from a photo of a 38-mm slug (Fig. 34) is
transparent white on the back and on the sides of the foot. The notum and
foot are bordered by an opaque white zone. That of the notum is thickened
and bears voluminous glands with broad openings. Outward to the white
zone of the notum follow an inner red and an outermost yellow line, while
that of the foot has only a yellow line. The clubs of the rhinophores and
the tips of the gills are red; the rhachis of the rhinophores is opaque white,
and probably also the quite narrow outer side of the gills. The opaque white
and yellow elements are still distinct in the preserved specimens.
The preserved slugs are 12-20 mm in length; the biggest is 11 mm broad
and 8 mm high. The notum is smooth, and so are the slightly protruding
rims of the rhinophoral and branchial pockets. The rhinophores of the
living animal lie 1/4-1/5 of the total length behind the anterior border;
their clubs bear 25-30 foliations. The distance between the gills and the
hind end of the notum is about the same as that between the rhinophores
and the fore end. The branchiae are unipinnate, the anterior ones longer
than the posterior ones. Their bases are arranged in two spiral lines to the

F ig u r e s 34-37. Casella sedna, sp. nov. 34, living anim al from color photo;
35, jaw elem ents; 36, radular teeth; 37, diagram o f reproductive organs.
179
sides of the anus. In the smallest present specimen there are 18, in the
largest 34 gills. Dr. Pickens observed that the animals continuously rotate
the gills back and forth, clockwise and counterclockwise. The border of the
mantle is undulated and about 2 mm broad. Also the edges of the foot are
undulated and projecting. The yellow border line ends where it reaches
the anterior border of the foot which is grooved transversely.
The labial cuticle is strengthened in a yellowish ring-shaped area
consisting of pegs about 65 ^ long (Fig. 35), 4 ^ in diameter, and cleft
into two pointed tips. The radula (Fig. 36) is 4 mm long, 1.6 mm broad;
its formula is 130 X 55.(1).55. The rhachidian pseudo-tooth is 30 ^ high
and has a cusp 16 ^ long; the corresponding measurements of the first
lateral tooth are 53 ^ and 32 p. The cusp of the second lateral tooth is
40 fx long; the cusps in the middle of the half-row measure 55 jjl. The cusp
of the innermost lateral tooth bears 1-2 inner and about 5 outer denticles.
The following teeth have only outer denticles, up to 7 in number, on their
cusps. The outer 25-35 teeth have smooth cusps; the outermost teeth are
small. The stomach and the small left caecum are covered by the digestive
gland. The strongly curved intestine courses first to the left side, then
forward, then to the right and backwards, and then curves to the anus in
the mid line.
The ramified ovotestis lies upon the liver. The quite thin hermaphrodite
duct (Fig. 37, h) opens into a spacious ampulla ( a ), whose outlet merges
into the female gland mass (g). Here the male and female ducts separate.
The spermatic duct emerges from the gland mass far ectally. Rather near
to its origin it widens, becomes glandular and much convoluted ( q ). When
these coils are spread out, they prove to be a prostatic section of the male
duct, not an annexed, massy prostate. The following part ( d ) of the
efferent duct is a thin, muscular tube. Entally it is coiled, ectally it
straightens. Then the muscular sheath is thickened, and the thin duct itself
winds through the sheath functioning as an acrembolic penis ( p ). In the
outermost section of the ejaculatory duct ( e ) the muscular sheath is thin­
ner. The male opening ( ma ) is the foremost aperture of the common
genital vestibule. Immediately behind the male opening the thin vagina
(v) begins, describes a loop and opens entally into a spherical spermatheca
( s ) and a longish spermatocyst (sc). So the arrangement of the “vesiculae
seminales” (Odhner, 1926: 53) corresponds to the vaginal type recorded
for Casella and the other genera of the Miamirinae. The insemination
duct (in) goes out from the vagina at some distance from the seminal
vesicles and reaches the gland mass (g) near the origin of the spermatic
duct. The nidamental duct (ni), the outlet of the female gland mass, is
the hindmost canal that opens into the common atrium.
Discussion .—A slug shaped like a Chromodoris, but with a strongly undu­
lated edge of the mantle and foot (Engel & Eeken, 1962: 25) can easily
be allotted to Casella. But a moderate undulation as in the present species
180
differs only by degree from what occurs in certain species of Chromodoris,
such as Chr. nona (Baba, 1955: PL 9, Fig. 23). The decreasing of the
denticles on the outer radular teeth belongs to the characters of the
subfamily Miamirinae, in contrast with the Chromodoridinae (Odhner,
1926: 53). If the pectinate outer teeth of a Chromodoris (Farmer, 1963:
Fig. 1, b) are compared with the smooth ones of the present Casella, the
difference is obvious. However, smooth outermost teeth occur in Chromo­
doris too (ibid., Fig. 1, g), and denticulate ones in Casella (Vayssiere,
1912: 58; O’Donoghue, 1929: 825).
The tips of the labial rodlets, entire or cleft, and the presence or absence
of a rhachidian thickening or a pseudo-tooth are variable characters in
Casella.
So high a number of gills as in the present species is exceptional in
Chromodoris. But this character is not reliable; of Casella atromarginata
from the Red Sea there are indications of 15 (Vayssiere, 1912: 57),
25 (White, 1951: 248), or 22-32 gills (Gohar & Abul-Ela, 1959: 49),
and in specimens from the eastern Indian Ocean 12 (Bergh, 1905a: 162)
and about 30 (Risbec, 1956: Fig. 35) were recorded.
The color of the dorsal surface of the mantle varies in certain species of
Casella. In the original material of C. rufomarginata Bergh (1890b: 943)
from Amboina it is darker than the border-line, in a specimen from the
Red Sea lighter (Engel & Eeken, 1962: 25). Therefore we separate
C. sedna from the other species by a short discussion, not by an inclusion
in our previously published key of the genus (Marcus, 1960b: 904), where
C. rufomarginata was separated according to its original description. In
that species as well as in C. atromarginata, atromarginata pallida, and foxi
only the notum, not the foot, bears a colored border-line. In C. cincta and
C. sedna both are colored, blue in cincta, red and yellow and yellow in
sedna. C. cincta has 11, sedna 17 gills on either side. C. rubra is uniformly
yellowish red without border-lines.
The hitherto known range of the species of Casella extends from Lake
Timsah, 81 km north of Suez, to middle Japan, New South Wales, the
Solomon Islands, and Tahiti.
Inuda, gen. nov.
Diagnosis.—Dorididae with cleft hooks on the labial cuticle, multidentate
radula with bicuspid rhachidian tooth and denticulate lateral teeth.
Prostate well developed; male copulatory organ acrembolic, with spines;
vagina unarmed.
Type species.— lnuda luarna, spec. nov.
Apart from the shape of the rhachidian tooth, Inuda resembles Cadlina
but the well-developed prostate makes it impossible to allocate these
genera in the same subfamily.
181
 We also pondered an affinity of lnuda with Carminodoris Bergh, 1889,
and therewith its systematic place in Odhner’s Baptodoridinae (1926: 54).
But the radula and the labial armature are less differentiated in all species
of Carminodoris (Bergh, 1889: 818; 1905a: 111; Risbec, 1928: 109;
1930: 283; 1937: 159; 1953: 47), so that it appears artificial to assign
lnuda to the same subfamily.
It requires a subfamily of its own, Inudinae, subfam. nov., to be placed
between the Chromodoridinae and the Cadlinellinae.
21. lnuda luarna, gen. nov., spec. nov.
Figures 38-44
Material.—Puerto Penasco, Sonora, Mexico. P. Pickens. Found under
boulders in a large tide pool, together with Taringa aivica timia.
The two specimens at hand were preserved together with the mentioned
discodoridine which differs externally from I. luarna by its notal sculpture.
Description.—The preserved animals are whitish mottled with brown
(Fig. 38), due to numerous dots of dark brown pigment (Fig. 40, is)
lying in different layers of the notum. The rhinophores are brown, and
also the gills are darker than the notum. The measurements are: 27 (20)
mm long, 20 (18) mm broad, 11 (14) mm high. Measured over the back
the lengths are 37 (40) mm. The foot is 20 mm long, 8 mm broad, its
anterior border is bilabiate, slightly concave in the middle (Fig. 39). The
antero-lateral corners of the foot are distinct.
The rather stiff notum is smooth; the vertical bundles of spicules ( cu)
do not project over the surface, but produce some slight and irregularly
distributed elevations of the skin. The spicules are up to 0.6-0.8 mm
long and 30 ^ in diameter. Most of them lie in the deeper layers of the
body wall. The tentacles are broad and grooved on their outer side. The
rhinophoral clubs bear about 16 leaves. There are seven tripinnate gills.
The rims of the rhinophoral and branchial pockets are smooth.
The labial cuticle bears a broad yellow ring of stratified hooks (Fig.
41). These are cleft for half their length, up to 50 ^ long, and 10 ^
broad. The radula (Fig, 42) is 3.1 mm long, 1.8 mm broad. It comprises
145-150 rows, has a well-developed rhachidian tooth and 50 teeth per
half-row. The rhachidian tooth (r) is 50 ^ in height, 16 ^ in breadth,
and bears two 16 /x long cusps. Its base is tripartite. The innermost
lateral tooth has a secondary cusp on either side of the middle cusp
(50 fx). In the following teeth there is no inner secondary cusp, and the
outer one diminishes in size, constituting a denticle. Behind it smaller
denticles appear, increasing in number outward to about ten. In the different
rows the denticulation varies much. The length of the main cusp increases
outwards to 65 ^. The outermost teeth are imperfectly developed. The
stomach is covered by the intestinal gland.
182
F ig u r e s 38-44. Inuda luarna, gen. nov., sp. n ov. 38, sculpture o f notum ; 39,
fore end in ventral view ; 40, transverse section o f notum , clarified; 41, labial
rodlets; 42, radular teeth; 43, diagram o f reproductive organs; 44, tip o f penis.
 The exit of the ampulla (Fig. 43, a) divides into the short inner
oviduct (o) which merges into the female gland mass (g), and the male
duct. The latter begins as an inner prostatic part ( q ) forming broad folds,
and continues as a tubular portion with high cells and large nuclei. The
muscle layers of the following ciliated efferent duct ( d ) separate into an
outer and an inner layer, the former is a sheath 0.7 mm long. The ectal
section of the duct bears about 20 longitudinal rows of spines up to
18 fji long, which end where the ejaculatory duct coalesces with the
atrium (ac). In Fig. 43 about half the penis ( p ) is everted, as is shown by
the spines on its outside and inside. Some of the spines are slightly
curved (Fig. 44).
The vagina (v) begins in the fundus of the common atrium (ac); it
is folded but not cuticularized. The spermatheca (s) and the spermatocyst
(sc) are both connected with the vagina by long canals. This arrangement
corresponds to that in Cadlina laevis (Odhner, 1939: Fig. 13); Odhner
calls it the semiserial type. The nidamental duct (ni) opens (aw) imme­
diately behind the male aperture (ma).
22. Doris pickensi, spec. nov.
Figure 45
Material.— 1. Puerto Lobos, Sonora, Mexico. P. Pickens, April 16, 1965.
“Fairly common under boulders in the intertidal zone; also found in the
spring at Puerto Penasco.” We received nine specimens together with one
of Taringa aivica timia.—2. Puerto Penasco, Sonora, Mexico. Mary Anne
Hill, February 5 and March 5, 1966. February: “These are probably a dif­
ferent color phase of Doris pickensi.” March: “A very small orange form
(7 mm) found among some gorgonians.”
Description .—In the photographs, the animals are 25 mm long, white
with a translucent border of the notum and more opaque white notal
papillae (Fig. 45, A). Dr. Pickens states “that some specimens have tinges
of brown or are speckled with brown.” The rhinophores, gills, and the
anterior border of the foot are light yellow. Evidently also Dr. Lance
collected this species in the Gulf, to judge from a kodachrome that he
kindly sent us.
The largest, much contracted, specimen is 23 mm long, 15 mm broad.
The breadth of the lateral borders of the notum is about 3 mm. The gills
are 5 mm from the hind end. The length of a smaller, straighter animal
is 12.5 mm, its width 8.5 mm, and the height 3.5 mm. The hyponotum is
2 mm broad; the foot measures 11X4.5 mm.
Of material from collection 2 we have three specimens collected in
February. The largest measures 19 X 13X7 mm. In the color photograph
the slugs are a gelatinous light orange with brick red rhinophores and gills.
The specimen collected in March was 7 mm long alive and preserved.
184
 The notum bears small, mamillary warts whose tips are about 300 ju,
distant from one another in the central parts of the notum, 200-250 ^ on
the sides. Each papilla has a big spicule (Fig. 45, B), 400 ^ long and 35 ^
in diameter, and several smaller ones surrounding it. The spicules may
project on the surface. A mat consisting of thick spicules lies in the back,
extending into the hyponotum.
The tentacles are short, broad and grooved on their outer side. The
clubs of the rhinophores have 10 leaves. There are 13-14 unipinnate
gills. As far as the rims of the much contracted rhinophoral and branchial
pockets can be seen, they are smooth, not papillose. The anal papilla is
high and lies somewhat behind the center of the branchial circle. The
anterior border of the foot is bilabiate, sometimes notched, sometimes
entire.
The labial cuticle is smooth. The radula of a smaller animal has 35,
that of the largest 40 rows. The half-rows contain 50 and 64 teeth res­
pectively. All teeth are hook-shaped, without denticles; their size increases
in the two inner thirds of the half-row, decreasing in the outer third. The
innermost teeth converge over the narrow rhachis. The salivary glands are
quite short and broad sacs which envelop a great part of the pedal com­
missure. The free stomach is large; the roundish caecum lies beneath it.

F ig ure45. Doris pickensi, sp. nov. A , sketch of living slugs from color photos
1 and 33; B, transverse section of notal papillae; C, diagram of reproductive
organs.
185
 The narrowed outlet of the sausage-shaped ampulla (Fig. 45, C, a)
enters the female gland mass (g) from which the male duct emerges. It
begins narrow but widens immediately and becomes prostatic ( q ). The
following portion ( ss ) runs winding within a muscular sheath, which is
entally continued into a retractor (r) and ectally opens into the atrium.
The efferent duct does not form a papilla. From the atrium the vagina (v)
courses apposed to the male sheath and leads first to the large spermatheca
(s) and then, with a circular loop, to the small spermatocyst (sc). Near
the spermatocyst the insemination duct (in) connects the seminal recepta­
cles with the glandular oviduct (g). The nidamental oviduct (ni) opens
(aw) immediately behind the male aperture.
The species is named for Dr. Peter E. Pickens.
Discussion.— Though the acrembolic ejaculatory duct may project into
the male atrium (“preputium,” Bergh), it does not form a true penial
papilla (glans) as in Archidoris. The absence of a prostatic part of the
efferent duct is the principal character of Austrodoris; a muscular sheath
of the efferent duct occurs in Austrodoris and in Doris . As Hoffmann
(1938: 1070) said, thick and short salivary glands ( Austrodoris) and
long, band-like ones (Doris) cannot be used as generic characters, because
in A. nivium Odhner (1934: 269) both types were observed in one and
the same animal. In African material of D. verrucosa, Bergh (1907: 47)
found the salivary glands nodular and shorter than usually in this type
species of Doris. Papillae on the rims of the rhinophoral and branchial
pockets, which protect them like valves, are specific, not generic characters
(Eliot, 1903: 557).
Doris echinata O’Donoghue (1922a: 149) has brown spots on the
opaque white notum, but differs from Doris pickensi by six gills, four
anterior unipinnate and two posterior bipinnate ones, and 16-18 radular
rows of 13-15 teeth per half-row. It was found at San Juan Island (Wash­
ington) and Vancouver Island (British Columbia), and called Doridigitata
maculata O’Donoghue (1926: 206) and Doris odonoghuei Steinberg
(1963a: 63). The reproductive organs of this species are not known, so
that its true generic place is not settled. In his family Doridigitatidae
O’Donoghue (1927a: 87-89) included a white slug from Laguna Beach,
for which he created the new genus Glossodoridiformia (species: alba). Its
notum is villous, there are five unipinnate gills, 17-18 radular rows, and
25-26 teeth per half-row.
Doris (Asteronotus) alabastrina Cooper (1862: 204) from San Diego
is alabaster white, opaque, and has 12 unipinnate gills. These characters
agree with the present species, while Conualevia alba Collier & Farmer
(1964: 383) which occurs abundantly at Newport Bay, hence one degree
north of San Diego, has eight tripinnate gills. But the description of
Cooper’s species is so poor that its name, indeed a “nomen dubium”
(Steinberg, 1961: 59), cannot be used for the present species.
186
 23. Discodoris mavis, spec. nov.
Figure 46
Material.--Puerto Penasco, Sonora, Mexico. P. Pickens, March 5, 1966.
The slug was found crawling under a granite boulder. “When pulled olf
a rock it has a tendency to fold the edge of the mantle ventrally, so that
it covers the foot” (P. Pickens).
Description .—The living specimen was 24 mm long, 17 mm broad. Its
color was orange pink with scattered brownish spots (Fig. 46, A), denser
towards the sides. The color has completely disappeared in the preserved
specimen; only some dark rings around the openings of the cutaneous
glands are recognizable.

F ig ure 46. Discodoris mavis, sp. nov. A, pigment pattern of living slug from
color photo; B, spicule pattern on border of same; C, section of notum; D, end
of half-row of radula; E, diagram of reproductive organs.
187
 There are no traces of spicules, but the color photo shows small
tubercles with white tips (Fig. 46, B). These were evidently supported by
vertical spicules. Cutaneous glands lie between the tubercles (Fig. 46, C).
The preserved specimen, that of the color photo, measures 19 X 13X8
mm. The underside is pigmentfree. The thin edge of the notum is curled
inwards. The foot is 7 mm in width and projects beyond the 5-mm-broad
hyponotum. The tentacles are digitiform; the anterior pedal border is
bilabiate, and the upper lip notched. The rhinophores have a white tip,
a pink shaft, and a brown club with 17 foliations. The rim of the
rhinophoral pocket is smooth. The 8-10 orange-pink gills are multi-
pinnate; the border of their pit is smooth.
The labial armature consists of two areas with stratified rodlets up to
100 n high. The radula measures 2.6 X 1.6 mm and comprises 25 rows.
Each half-row contains 15 marginal lamellae and 21-22 hooks (Fig. 46, D),
up to 230 /A in height. The innermost hook is slightly angled on the
rhachidian side under the cusp. The edges of the lamellae are irregularly
dentated, probably worn, not provided with distinct denticles.
The ampulla (Fig. 46, E, a ) is long and curves around the prostate,
which is composed of three parts. The acini of the inner part ( q ) have
small cells. Apposed to it lies the outer part ( qu ), whose acini are big-
celled. Then follows a smooth tube ( d ) which begins glandular, as third
prostatic section of the efferent duct, and continues muscular. The sheathed
muscular part (ss) of the efferent duct leads to a rather long penial
papilla ( p ) which projects from the male atrium (am ).
Behind the atrium lie the vagina (v) and the protruded nidamental
opening (aw). The vagina is muscular and seems to be coated with a thin
cuticle; its ectal part bears a large gland (vn) lined by a high epithelium
with basal nuclei. Entally the vagina narrows and opens into the longish
spermatheca (s). Near its entrance, but independent from it, the in­
semination duct (in) leaves the spermatheca. The spermatocyst (sc) has
its own connection with this duct, which merges into the gland mass (g)
near the nidamental duct (ni).
Discussion.— D. mavis is related to D. heathi MacFarland, 1905, from
Vancouver Island to northern Lower California (Farmer & Collier, 1963:
62; Farmer, 1967: 341) and to D. pusae Marcus from the coast of S. Paulo,
Brazil, and from Miami, Florida. D. heathi is yellow ochre to raw umber
(MacFarland, 1966: 193), D. pusae orange or pinkish red. The darker
flecks occurring in both species are especially dense in the mid-region.
The skin of D. heathi (Marcus, 1961a: Fig. 62) is rather similar to that
of mavis, but the spicular tubercles are larger and more densely set in
heathi. The skin of D. pusae (Marcus, 1955: Fig. 154) has mainly
horizontal spicules, so that it is less similar to the skin of D. mavis.
The labial rodlets are about 42 ^ long in D. heathi, those of D. pusae
reach 60 In D. heathi the number of radular rows, 20-22, is inferior
188
to that of D. mavis; in D. pusae it is 21-26. The number of marginal
lamellae and lateral hooks per half-row does not furnish palpable dif­
ferences; D. heathi has 9-17 lamellae and 20-31 hooks; D. pusae 8-18
lamellae and 15-27 hooks. The denticle-shaped shoulder of the innermost
hook and the irregular denticles of the lamellae noted for D. mavis occur
in D. pusae as well.
The prostates of all three species agree; the penial papilla of D. mavis
and D. heathi is longer than that of D. pusae. In large specimens of the
latter the duct of the vaginal gland is provided with cuticular spines; the
gland of D. heathi is angled, and so its shape is farther distant from that
of D. mavis than that of D. pusae.
24. Diaulula sandiegensis (Cooper, 1862)
References.—MacFarland, 1906: 122; 1966: 190; Marcus, 1961a: 18.
Material.—Puerto Penasco, Sonora, Mexico. P. Pickens, February and
May, 1965. “Rare. Collected in the rocky intertidal.”
Further distribution.—From Lower California, Cape San Lucas (Lance,
1961: 66) to the Aleutian Islands and northern Japan. Original locality:
San Diego.
Descriptive notes.—The present specimens are small, 20 and 12 mm in
length. They have a series of brown rings on either side of the back.
Many paler rings are scattered chiefly on the sides of the white notum,
but also in the middle. The radula of the larger slug has 17 rows with
29 teeth per half-row.
Probably the “large, seal-brown nudibranch” from the western coast of
the Gulf (Steinbeck & Ricketts, 1941: 543) is this species. According
to Lance (loc. cit.) D. sandiegensis is uncommon intertidally, common
subtidally to 37 m.
25. Taringa aivica timia, subsp. nov.
Figures 47-51
Material.—Puerto Penasco, Sonora, Mexico. P. Pickens, January, March,
August 1964, January 15, and April 16, 1965. Generally found under
rocks in large tide pools together with Inuda luarna, Doris pickensi, and
Doriopsilla albopunctata.
Description.—The photographs of three of the five specimens (collection
No. 20, 20 mm alive; 61, 62) show a purplish brown ground color of the
notum with light marks around the back, the branchial pocket and the
bases of the papillae. These are brownish yellow, the rhinophores dark,
their tips white. In preservation the hyponotum and the sole have dark
brown flecks, especially numerous along the dorsal border of the foot.
In another photograph (no. 19, one specimen 35 mm long) the animal
is brownish pink with brown dots, light stellate marks around the bases of
189
the light orange papillae, light violet gills, dark brownish violet rhinophores
tipped with white, and with white papillae on the rims of their pockets.
Some of the notal papillae appear branched. In a third photograph (no.
14, six specimens) the notum is yellowish grey (Fig. 47). The rhinophores
are dark brown, the gills of the ground color, and the cutaneous papillae
yellowish. Around their bases they have dark brown blotches.
Measured over the back, the largest of the preserved specimens, one
of lot no. 14, is 70 mm long, its linear length is 35 mm, the height 26 mm,
the breadth 43 mm; that of the hyponotum is 10 mm. The much folded
foot measures 25 X 20 mm. The brown dots of the hyponotum are pre­
served. A 17-mm slug (no. 1) has uniform caryophyllidia. Evidently it
had been quite white alive, because it was put into the vial with Doris
pickensi. In all other specimens the notum bears a few long, smooth
papillae between the uniform, densely set caryophyllidia. The papillae are
much contracted in the preserved specimens and only sometimes bicuspid.

F ig ures47-51. Taringa aivica timia, ssp. nov. 47, sketch of living slug from
color photo 14; 48, radular tooth from middle of row; 49, marginal tooth; 50,
diagram of reproductive organs, prostate simplified; 51, penial cuticle.
190
The tentacles are pointed, sometimes widened under the tip. The rhino­
phoral clubs have 16 leaves. The rim of their sheaths bears about eight
lobular papillae. There are six multipinnate gills; the border of the
branchial pouch is undulate. The anterior border of the foot is bilabiate
and notched.
The labial cuticle is smooth. The radula of a 29-mm slug, belonging to
lot no. 20, comprises 29 rows with 48 teeth per half-row. The 44 inner
teeth are hook-shaped, the four outer ones pectinate. The inner teeth
increase in size to the 30th (Fig. 48), continue high to the 40th, and
decrease to the 44th. The cusp of the hook-shaped teeth bears a minutely
knobby ridge on the outer side, which corresponds to the distinct denticles
in aivica aivica. The radula of the largest specimen is 7 mm long, 6 mm
broad, has 41 rows, and 75-76 teeth per half-row. Of these the 70 inner
ones are hook-shaped, the 5-6 outer ones pectinate. The inner teeth have
only a ridge on the outer side of the cusps, but no knobs. The 17-mm slug
had no knobs at all; its radular formula is 30X45.0.45. In all examined
radulae the flat, pectinate teeth (Fig. 49) have a broad uppermost spine,
and the rest of the edge is feathery.
The prostate is a massy dilatation of the efferent duct; in the diagram
(Fig. 50, q) it is simplified. The distal part of the duct ( d ) has a folded
epithelium. The penial papilla is coated with a beaker-shaped thick cuticle
(Fig. 51). The wall of the male atrium ( am ) bears spineless papillae.
Between the male atrium and the glandular nidamental duct ( ni ) the
narrow vagina (v) begins tubular, not bulbar. The vaginal epithelium is
thrown into longitudinal folds. The entrance of the vagina into the
spermatheca (s) and the exit of the insemination duct (in) are neighboring,
but independent from one another. The entrances of the inner oviduct (o)
and the insemination duct into the female gland mass (g) almost meet.
Discussion.—The present animals represent a new subspecies of T. aivica
Marcus, 1967. Both have long papillae and small caryophyllidia on the
notum. The denticles of the lateral teeth are strong in aivica aivica, and
its marginal teeth have not the broad spine of aivica timia. This spine
resembles that of the pectinate teeth of Dictyodoris tesselata Bergh
(1880b: PI. C, Fig. 11), a doubtful doridid, whose prostate is not known.
The muscular bulb of the vagina with cuticularized folds in aivica aivica
and the absence of spines on the atrial papillae of a. timia might be due
to different states of reproductive development. The shape of the cuticular
coat is different in both subspecies. Alive, the color of the previously
described form is yellowish mottled with grey.
Tayuva, gen. nov.
Diagnosis.—Discodoridinae with pointed tentacles, labial plates with
rodlets, hook-shaped radular teeth, stout penial papilla, large vestibule
(atrium) stiffened by spicules and lodging the penial papilla and the
191
vaginal aperture. Nidamental opening independent from that of atrium.
Type species.— Tayuva ketos, spec. nov.
The anatomy of the present species evidences Bergh’s terms “preputium
or penial sac” as inappropriate for the terminal section of the male duct.
There are two genera of the Discodoridinae, Phialodoris and Audura,
with aberrant outer genital organs, but neither of them can receive the
present species.
Tayuva ketos is one more example to reveal that the classification
of many doridids is impossible without dissection of the reproductive
organs; the extraction of the radula is often insufficient.
26. Tayuva ketos, gen. nov., spec. nov.
Figures 52-56
Material.— 1. Puerto Penasco, Sonora, Mexico. P. Pickens, November 2,
1963.—2. Puerto Lobos. P. Pickens, April 16, 1965. Under rocks in
large tide pools.
Description.—In a photograph of the specimen from locality 1 (Fig. 52)
the notum has brown blotches on a whitish ground, and between them
single brown papillae. These are more numerous towards the sides where
the blotches are smaller. The rhinophores and gills are brown. The blotches
of the slug from locality 2 are greyish brown, and several especially large
ones occur in the marginal zone. The rhinophores are brown with light
tips, the gills greyish.
The specimen from the first locality was 47 mm long alive; preserved,
it is 30 mm in length, 22 mm in width, and 9 mm in height; the second
slug measures 50, 40, and 8 mm. The sole of the smaller animal is
18 mm long, 9 mm broad. The hyponotum of both specimens is white with
small dark flecks. Those on the light sole are larger.
The papillae (Fig. 53) of the smaller slug are rather pointed, those of
the larger, evidently more contracted animal, blunt. The tips of the
papillae are ciliated. The spicules, which are up to 350 ^ in length and
30 /x in diameter, do not reach the tips of the papillae.
The white tentacles are 2.5 mm long, carrot-shaped, and have some dark
spots. The rhinophores have 24 leaves; the borders of their pits and that of
the branchial pocket bear papillae of the common type. There are six
pluripinnate gills. The anterior border of the foot is bilabiate and notched
in the middle.
The yellow labial cuticle bears up to 100 ^ long striped rodlets (Fig.
54) which are 10 /x in diameter at their thickened free end. The radula
of the smaller animal measures 2.7 X 2.4 mm, that of the larger one
4.0 X 4.0 mm. There are 24 (locality 1) and 29 rows (locality 2), and 38
and 41 teeth per half-row. The teeth (Fig. 55) are hooks whose cusps are
up to 220 jui long in the middle of the half-row and smaller to the sides.
192
 The blood glands are dark. The oesophagus enters the completely free
stomach from the ventral side. The stomach is large; its dorsal, strongly
folded and glandular wall projects into the gastric lumen. The intestine
leaves the stomach on the left side, in front and dorsally, and curves to
the right and backwards. The pylorus conceals the small caecum.
Near the outlet of the winding ampulla (Fig. 56, a) the short inner
oviduct and the spermatic duct ( d ) separate. The former merges into the
female gland mass (g), the latter becomes prostatic. The longish ental
portion of the prostate ( q ) is soft and consists of radially disposed
glandular sacs. The ectal globular portion ( qu ) is more compact. Farther
outwards follows the efferent duct ( d ) containing sperm in its folded
lumen. The duct ends with a stout, crooked penial papilla (p), which
projects into the large vestibulum ( ac ). The vestibular wall is stiffened by a
layer of longitudinal spicules, 3 mm in length and 10 jjl in diameter.

F ig u r e s 52-56. Tayuva ketos, gen. nov., sp. nov. 52, living slug from color
photo; 53, spicules and m elanophores in notum papillae; 54, labial rodlet; 55,
radular tooth from m iddle o f half-row ; 56, diagram o f reproductive organs.
193
 The vagina (v) whose epithelium is folded begins in the ental third of
the atrium. Quite near to its entrance into the spermatheca (s’) the insemi­
nation duct (in) emerges from the latter, runs curved towards the female
gland mass (g), and receives the canal from the spermatocyst (sc) short
before it reaches the mass. The disposition of the sperm-storing vesicles
is of the serial type (Odhner, 1926: 51). The insemination duct opens
into the gland mass near to the entrance of the inner oviduct. The
aperture (aw) of the outer oviduct, the outlet of the gland mass, is inde­
pendent from that (ma) of the penial and vaginal atrium.
27. Nembrotha eliora, spec. nov.
Figures 57-58
Material.— 1. Puerto Lobos, Sonora, Mexico. Paula Vreeland, April 16,
1965. Three specimens in a small boulder-strewn area in the intertidal
zone.—2. Same locality. Alex Kerstitch, February 25, 1966. One specimen
subtidally in about 10 m of water, on a rock.
Description .— (Material 1). A photograph shows that the slugs are black
with many green and blue longitudinal stripes (Fig. 57). To the sides of
the median blue line bordered with black, the green lines form two triangles
between the black rhinophores. The paramedian green stripes end in front
of the gills which are black with a green line along the efferent and afferent
vessels; the more lateral green stripes are confluent behind the gills. Each
of the two next pairs of green stripes is subdivided by a black streak which
begins in the anterior third. The most ventral stripes are undivided and

F ig u r e s 57-58. Nembrotha eliora, sp. nov. 57, living slug from color photo;
58, half-row o f radula.
194
 uninterrupted from the anterior to the posterior end. In preservation the
blue color has disappeared from the black ground, but the green is pre­
served. Ten light and 15 black stripes are distinguishable. The light
stripes unite on the back of the tail which is smooth, not cristate.
The two present specimens are 30 and 12 mm in length, 10 and 8 mm
in height, and 6 and 4 mm in width. The sole is folded. The anterior
border of the notum is somewhat thickened. The border of the rhinophoral
pits is light in one, dark in the other specimen. The clubs have about 22
leaves. The broad black tentacles have a light outer groove. There are five
and seven pluripinnate gills. Extra-branchial processes (as in Tambja
Burn, 1962) are not developed. The anus lies between the gills, the
gonopore at the end of the anterior fourth of the body. The anterior pedal
border is green, thick, neither grooved nor notched. The sole is white,
its border black.
The cuticle of the labial disc is folded and smooth, not composed of
rods. The buccal mass is 6 mm long in the larger specimen. The radula
(Fig. 58) is yellowish, except for the pale first lateral tooth. There are
15 rows of teeth and four outer lateral teeth per half-row. The rhachidian
tooth is a transversely striate plate, divided in the middle, 0.35 mm in
width and 0.29 mm high. Its anterior edge is thickened and smooth. The
inner lateral tooth is 0.8 mm or more in length, its pointed cusp 0.6 mm;
a strong spur projects in the middle. The outer teeth decrease in size
outwards; the first has a strongly crested inner edge.
In order to keep one specimen intact, the larger one with everted genital
atrium was dissected, because only the male organs are systematically
important. The spermoviduct which leaves the large and wide ampulla
divides within the female gland mass as in N. morosa Bergh (Macnae,
1958: Fig. 14). The prostatic part of the efferent duct begins with a thin
layer of muscles, its ciliated epithelium is high; its wide lumen contains
sperm. Farther outward the muscles become thicker and the lumen narrows;
the diameter of the duct continues the same. In the next section the duct
is thicker. Here its muscle layers are divided into that of the duct proper
and that of its sheath. The terminal part of the duct in the sheath bears
cuticular hooks and opens into the fundus of the atrium, whose epithelium
is greenish. In the present everted vestibulum the occurrence of a vestibular
gland, as indicated for morosa by Bergh (1907: 70), not by Macnae
(1958), could not be examined.
(Material 2). The living animal was about 80 mm long (Dr. P. Pickens).
Preserved it is 40 mm long, 20 mm high, and 12 mm broad. In the photo­
graphs the ground color is green. The lateral crests which separate the
notum from the sides and unite behind the gills into a caudal crest are more
salient than in the specimens from lot no. 1. Their color is yellow, trimmed
with a thin black line. Also the yellow bulges on the notum which
correspond to the stripes of material 1 are seamed with black. The tentacles
195
are yellow; the clubs of the rhinophores violet, their sheath yellow with a
black basal rim. Two dark blue spots appear behind the rhinophores. The
gills are violet, their inner (afferent) and outer (efferent) vessels yellow.
The margin of the foot is broad and bright ochre.
Discussion.—This is the first Nembrotha from the Pacific coast of the
Americas. The two American Atlantic species are N. gratiosa Bergh
(1890a: 172) and N. divae Marcus (1958b: 47). The latter, found at
Cabo Frio (22° 57' S, 42° 01' W), in life is scarlet with white dots and
white gills, and therefore differs widely from the present species, as it does
also by its radula. The color of N. gratiosa, from off Key West in 52 m
depth, is similar to that of specimen 2 from Puerto Lobos, but the radulae
are different. The rhachidian plate of N. gratiosa is undivided, its anterior
edge is thin, and there are only three outer lateral teeth. The color of the
slugs of lot no. 1 differs from that of N. gratiosa. Among the four species
recorded from Japan (Baba, 1960: 72-73) N. sagamiana Baba (1955: 16,
44) has a similar divided rhachidian tooth; due to its tuberculate notum,
Burn ascribes sagamiana to Tambja (1962a: 98). N. eliora is similar to
N. morosa Bergh (1877: 457), recorded from South Africa to the Philip­
pines, in Macnae’s concept of its synonymy (1958: 360). This species has
up to five light stripes and an undivided median tooth. Though the cleft tip
of the first lateral or intermediate tooth of morosa is absent in capensis,
Macnae united it with morosa; the simple cusp of capensis (Bergh, 1907:
PI. 12, Figs. 10-12) is much shorter than in eliora.
Bergh’s term (1883: 165) “prostata discreta nulla” (“gesonderte
Prostata fehlt” Thiele, 1931: 425) refers to the type-species ( nigerrima
Bergh) and others, such as morosa (Macnae, 1958: 361), divae Marcus,
1958, and eliora. In their intercalary prostates the widening of the prostatic
section of the efferent duct or an equal diameter of the efferent duct
throughout its length depends on its physiological state. An independent
prostatic gland annexed to the efferent duct as in N. lineolata (Eales, 1938:
Fig. 16) is a different type and can lead to a subdivision of the genus.
28. Polycera atra MacFarland, 1905
References .—MacFarland, 1906: 142; 1966: 115; Odhner, 1941: 17.
Material.—Puerto Penasco, Sonora, Mexico. P. Pickens, May 2, 1965 and
March 5, 1966.
Further distribution.—From Tomales Bay, Marin County (Marcus, 1964:
131), to Los Coronados Islands, Lower California (Lance, 1961: 66).
Original locality: Monterey Bay.
Remarks .—The pale grey ground color bears narrow dark stripes in
MacFarland’s specimens (1966: PL 18, Figs. 2 & 3). In a photograph
(no. 58) which we owe to Dr. Lance, the dark stripes are broader. So
196
they agree with the original figure (MacFarland, 1906: PI. 29, Fig. 22),
which was “redrawn from an incomplete color sketch” (page opposite to
PL 22). Still broader are the dark stripes in another photograph (no. 40)
received from Dr. Lance, and even broader than these in Pickens’ speci­
men of 1966. The darkest color pattern is shown by Pickens’ specimen of
1965, a dark brown animal not showing the light ground color.
The last-mentioned slug, as well as that of Pickens (1966), and that of
Lance’s photo No. 40 have red spots between the dark pigment, still
recognizable in the preserved material.
The tips of the velar processes and the rhinophores as well as the points
of the lateral tubercles are more or less intensely powdered with yellow.
The tubercles stand on the light stripes, except in Pickens’ specimen of
1965, which lacks these stripes. The maximum of 4 tubercles on either
pallial ridge (Marcus, 1964: 131) is exceeded by 5 (Pickens 1965 and
1966) and more (MacFarland, 1966: 116). The number of gills varies
from 7 (Lance, photos 58 and 40; Pickens, 1966) to 11 (MacFarland
1966: 116).
The determination of preserved specimens of polycerids is far from easy.
Certain colors, especially yellow tones, are generally not preserved. Con­
tracted rhinophores and partially retracted ones are indistinguishable. The
dorso-lateral ridges or pallial margins and thickenings on them are some­
times nearly evened in preservation. The color of the living animals is
valuable, but its intraspecific variation and change with age must be
considered.
The jaws and the radula are important and must be compared with the
published figures for every identification; for the jaw of P. atra one of
the original figures (MacFarland, 1906: PL 20, Fig. 67) should be used,
not the last one (id., 1966: PL 31, Fig. 29).
Until now, none of the species is known to occur on both the Atlantic
and Pacific coasts of the Americas, but as geographic criteria should not
be overrated in the classification of opisthobranchs, the following artificial
key refers to the American Atlantic and Pacific species of Polycera Cuvier,
1817, Greilada Bergh, 1894, and the farther distant Galacera Risso-
Dominguez, 1960.
1. Extrabranchial processes conspicuous lobes, or digitiform, club-
shaped or carrot-shaped appendages..................................................... 2
1. Extrabranchial processes in form of small tubercles, bosses, or
papillae, much shorter than the veil digitations....................................9
2. A single extrabranchial appendage on either side...............................3
2. More than one pair of extrabranchial appendages...............................4
3. Pallial ridge indistinct; rhinophores with 12 leaves.................................
.............................................................. marplatensis (Franceschi, 1928)
197
 3. Low pallial ridge between veil and extrabranchial appendages
present; rhinophores with 25 leaves.................. chilluna Marcus, 1961
4. Jaws with upper wing-like expansion................................................... 5
4. Jaws without wing-like expansion........................ priva Marcus, 1959
5. Three gills................................................... zosterae O’Donoghue, 1924
5. Seven to nine gills.......................................................................................6
6. Extrabranchial processes shorter than gills, .hedgpethi Marcus, 1964
6. Extrabranchial processes longer than gills.......................................... 7
7. Extrabranchial processes carrot-shaped......... aurisula Marcus, 1957
7. Extrabranchial processes nearly cylindrical with pointed tips.........8
8. Rhinophores with 14-15 leaflets; foremost gill especially long;
3-4 extrabranchial processes on either side. . . .hummi Abbott, 1952
8. Rhinophores with 9 leaflets; foremost gill not especially long, 2
extrabranchial processes on either side.................... gnupa, spec. nov.
9. Second tooth of radula with basal spine.......................................... 10
9. Second tooth of radula without basal spine...............................................
..................................................................... alabe Collier & Farmer, 1964
10. First tooth half the size of the second............. herthae Marcus, 1963
10. First tooth smaller than second, but larger than half its size........... 11
11. Pallial ridges with closely set mamillary bulbs between rhino­
phores and gills and behind gills; oral tentacles distinct........................
.................................................................................. odhneri Marcus, 1955
11. Pallial ridges with few, generally no more than five on either
side, compressed, pointed tubercles; oral tentacles very short.............
.............................................................................. atra MacFarland, 1905
29. Polycera gnupa, spec. nov.
Figure 59
Material.—Cholla Bay, Puerto Penasco, Sonora, Mexico. P. Pickens,
March 5, 1966. This species was found, together with P. atra “in a
branching bryozoan colony, growing on the sandstone reef in the middle
of Cholla Bay below the zero tide level.”
Description .—The two present specimens were 14 mm long in life, 7 and
8 mm preserved. The ground color is pale grey. This appears on the foot,
the pallial ridges, along the caudal crest, and on scattered tubercles on the
back and the sides. For the most part the dorsal and lateral skin is densely
stippled with brown pigment, which is concentrated to blackish brown
around the gills and on the head.
198
 The two velar processes on either side are long, slender and pointed.
They are white with a bluish and an orange band around the middle
(Fig. 59, A). The rhinophores have a dark shaft, 9 lighter brown perfoli-
ations, and a white tip. The oral tentacles (Fig. 59, B, t) are quite short
and broad lobules. The two extrabranchial appendages are nearly cylin­
drical with pointed tips. The tip is light and followed by a bluish and then
an orange ring. Both rings lie above the middle of the appendage. In the
larger specimen the outer, anterior, appendage is shorter, more pointed
and thinner than the stout inner posterior one; in the smaller specimen
it is longer. Both appendages are longer than the gills.
There are 7 unipinnate gills, the foremost of which is not much longer
than the others. The tip and the inner side of the gills are lighter than the
outer side. The knobbed median crest behind the gills ends near the tip
of the tail which bears an anterior orange and a pale blue posterior mark.
The antero-lateral corners of the foot are finger-shaped.

F ig ure 59. Polycera gnupa, sp. nov. A, right side view of preserved slug; B,
fore end of same in ventral view; C, jaws; D, half-row of radula, and one
innermost tooth in a different view.
199
 The jaws (Fig. 59, C) are yellow and have broad, upper, wing-like
expansions. Their inner border is stronger than in P. hedgpethi (Marcus,
1964: Fig. 1). The radula (Fig. 59, D) com prises 9 rows of
3-4.1.1.0.1.1.3-4 teeth. The height of the teeth in micra is from the middle
outwards: 220, 300, 140, 120, 100, 50. Old teeth are reddish brown. The
innermost or first lateral tooth has a pointed cusp, and in its middle two
unequal spurs; the second lateral tooth has a basal spur. The innermost
marginal tooth has a rudimentary cusp, the outer marginal teeth have none.
The eversible part of the efferent duct is provided with very numerous
(about 1000) cuticular bristles.
r
Discussion .— P. gnupa is similar to two West Atlantic species, but cannot
be united with either of them. However, its taxonomic separation is
possibly not definitive, and more material of gnupa may diminish the
differences between it and the neighboring species. The color pattern of
adult specimens (Marcus, 1958b: 56) of the Brazilian-Floridian P. auri-
sula, not that of young slugs which are brighter (id., 1957: 432), agrees
with that of P. gnupa, but the carrot-shaped extrabranchial appendages
of aurisula separate it from gnupa.
In P. hummi Abbott, 1952, the rhinophores and the extrabranchial
appendages bear two brilliant blue rings, one above and one below the
orange ring. The rhinophores of 10-mm-long living slugs have 14-15
leaflets. Mrs. Sally D. Kaicher (St. Petersburg, Fla.), graciously sent us an
excellent color photograph of P. hummi, determined by Dr. R. Tucker
Abbott. This photograph shows that the differences between the anterior
and posterior gills, a general feature in Polycera, is accentuated in
P. hummi, so that in lateral view the branchial cup appears slanting
backwards, as if it were cut obliquely, even more than in Abbott’s figure
(1952: PI. 2, Fig. 1). In a single specimen of P. hummi from Beaufort,
N. C., two denticles were found at the base of the unworn cusp of the
second radular tooth (Marcus, 1961b: Fig. 5). This character must be
considered with prudence in comparing P. hummi with the only two
specimens from the Gulf of California.
The big P. hedgpethi Marcus, 1964, from Tomales Bay, California, is
grey with small black dots and yellow-orange marks on the velar and
extrabranchial appendages, on the pallial ridges, caudal crest, upper border
of the foot, and body tubercles. Blue elements were not recorded. This
feature and the black dots distinguish the color pattern of P. hedgpethi
from P. gnupa. As we strive for a key to preserved material, whose colors
may vary in several specimens, we separated P. hedgpethi by its short
extrabranchial appendages. Its tripinnate gills and more radular rows might
be correlated with its larger size, and perhaps with age. A short cusp of the
first tooth as in P. hedgpethi (Marcus, 1964: Fig. 3) can be seen also in
P. gnupa in the corresponding view (Fig. 59, D).
200
 30. Acanthodoris pina, spec. nov.
Figure 60
Material.— 1. Puerto Penasco, Sonora, Mexico. Paula Vreeland, February
13, 1965. One specimen from the rocky intertidal zone.—2. Same locality.
Two more specimens found on February 5, 1966, by P. Pickens and Mary
Anne Hill.
Description .—As shown in a color photograph, a slug in movement is
rather flat (Fig. 60A); another animal, at rest, is higher. In life, the
length is about 18 mm; preserved, the animals are 8-11 mm in length,
7-8.5 mm in width, and 5-7 mm in height. The black dorsum is beset with
soft conical papillae, large red ones and numerous small white ones in
between. In preservation the bases of the large papillae are white and 1 mm
in diameter. The papillae are up to 2 mm in height. The notum, the
frontal veil, and the foot are bordered with red. The red stripe of the foot
is accompanied by an outer white line. In the photographs the rhinophores
are dark brown with white tips; the gills are brick red. The hyponotum is
sprinkled white and black; the sole is black.
The foot is rounded behind, where it stands out beyond the notum when
the slug is gliding. The anterior border of the foot is bilabiate due to a

F ig ure 60. Acanthodoris pina, sp. nov. A, living slug from color photo; B,
half-row of radula.— F ig ure 61. Dendrodoris krebsii (Morch). A, male duct,
specimen from Puerto Penasco (the section between the asterisks bears hooks);
B, male duct, specimen from Sao Sebastiao, Brazil.
201
transverse furrow; the upper lip is white. The veliform head has no median
notch. The clubs of the long rhinophores have 20 leaves; the sheaths bear
red papillae similar to those of the dorsum. In one case their number
differs on the two sides (3 and 5); in the other specimens there are 4 and 5
on both sides. There are 5-7 pluripinnate gills of equal size, one of them
in front. The anal papilla projects from behind into the center of the
branchial circle which is free from cutaneous papillae. One papilla com­
pletes the circle of the gills. The peritoneum, correctly called pseudo­
peritoneum by McFarland (1906; 1925-26), is colorless.
The labial armature consists of two lateral areas and a median plate of
thickened cuticle as in most species of the genus. The labial elements form
about 18 irregular rows. Their surface is rough, and they are widened
above and cleft into a variable number of lobules. The crop (ingluvies
buccalis) is less than half the volume of the buccal mass. The radula
(Fig. 60B) has 28 rows. There are 5-6 outer lateral teeth whose size
decreases outwards. The innermost lateral or intermediate tooth is 0.12 mm
high and 0.285 mm long. Its cusp is 0.12 mm in length, pointed and
curved. Its inner ridge bears 1-3 denticles. Sometimes there are up to
six denticles, and on some teeth none at all.
In the reproductive system the unarmed copulatory organ is a conspicu­
ous feature, though penial hooks as well as their absence have been
related for A. pilosa, A. rhodoceras, and A. falklandica (Eliot, 1910: 16,
67; Marcus, 1961a: 27). The prostatic section of the male duct forms two
loops and is little shorter than the following muscular part, which runs
within a muscular sheath. The muscular atrium, Bergh’s “preputium,”
is distinctly set off from the efferent duct and lodges its everted, cylindrical
tip.
The genital aperture lies at the level of the anterior pedal border. The
external openings of the male and the two female canals are united.
The long vagina runs looping inwards and opens into a big, globular
spermatheca. From this point a long, thin duct leads to the narrow
spermatocyst filled with orientated sperm, and a short duct, the insemi­
nation duct, to the female gland mass.
Essentially the genital organs agree with MacFarland’s figures (1925-26:
PI. 3, Figs. 1 and 6), except that the glandular part of the efferent duct
and the duct of the spermatocyst are longer in the present slug.
Discussion .—Leaving aside possible synonyms (Steinberg, 1963a: 64),
we have compared our specimens with the 12 species from the Pacific
coast of North America (O’Donoghue, 1927b: 6-7), two of which have
red elements in their color pattern. The back of A. pilosa purpurea Bergh
(1880a: 98) is purple brown, preserved reddish brown, not black, and
the shape of the first lateral tooth with its short and blunt cusp differs
from the present material. The dorsum of A. lutea MacFarland (1925-26:
12) is orange red with dots of lemon yellow between the papillae; there
202
are 34-39 radular rows, and the labial armature, especially the ventro-
median plate, is rudimentary.
Extending our comparison to the 7-10 further species of Acanthodoris,
we found it difficult to separate our slug from the geographically widely
distant A. metulifera Bergh (1905b: 98) from Tasmania. The different
characters of metulifera are: 40 radular rows; 3-4 denticles on the first
lateral tooth; head and foot yellowish; peritoneum with many violet-brown
flecks on the dorsal side. Possibly (Eliot, 1907: 350) Bergh’s species is
identical with the older A . globosa which has seven outer lateral teeth, and
whose labial elements are not divided.
31. Dendrodoris krebsii (Morch, 1863)
Figure 61
References.—Bergh, 1875: 87; Marcus, 1962: Fig. 19; 1963: 35; Collier
& Farmer, 1964: 389, Fig. 2 G-H, PI. 5; Marcus, 1967b: 95.
Synonyms.— Doridopsis Krebsii var. pallida Bergh, 1879b: 44; D. atropos
Bergh, 1879b: 49; Dendrodoris atropos Marcus, 1957: 443.
Material.— 1. Puerto Penasco, Sonora, Mexico. Rocky intertidal zone. In
1964 and part of 1965, April, June, July, and November.—2. Puerto
Lobos and Puerto Libertad. P. Pickens. “They are found infrequently.”—
3. Guaymas, Sonora, Mexico. Alex Kerstitch, March 12, 1966. Intertidal.
One specimen.
Further distribution.—Atlantic Ocean: Florida; Bahamas; Virgin Islands
(St. Thomas, St. Croix: original localities); Antilles; Curasao; coast of
Rio de Janeiro (original locality of atropos ) to Sao Paulo, Cananeia
(25° S).—East Pacific Ocean: Lower California (oceanic side); Gulf of
California from Puertecitos (30° 25' N) to the Bay of La Paz; mainland
of Mexico, Jalisco, Bay of Tenacatita (about 19° N, 105° W).
Remarks .—The color as shown by a photo of a 5 5-mm slug is velvety
black with a red-trimmed border and white tips of rhinophores and gills.
The latter are six in number, but as the two hindmost tufts are subdivided
from their bases, one might as well count eight branchiae. The largest of
the four preserved animals was 29 mm in length, 16 mm in width, and
10 mm in height.
Collier & Farmer stressed the thickened base of the penis in their East
Pacific material. In two specimens examined from Puerto Penasco we also
found this part thicker (Fig. 61A) than in two Atlantic specimens, one
from St. Kitts, Lesser Antilles, the other from Sao Sebastiao, coast of
Sao Paulo (Fig. 61B). These Atlantic slugs agree with one another.
For this comparison only completely retracted penes can be used. The
figures of Western Atlantic Dendrodoris, referring to D. krebsii (Bergh,
203
1875: PL 11, Fig. 15, 16) cannot be discussed, because the penes of the
three examined slugs (p. 90) were protruded in different degrees. In our
animals from the Gulf of California the ental penial hooks are up to 40 ^
high, while their maximum is 30 ^ in the Caribbean and Brazilian animals.
Neither this difference nor that of the diameter of the penial base is
sufficiently clear-cut to justify a subspecific separation. The length of the
section of the male duct between the prostate and the base of the penis is
variable. It measured 4 mm in one of the animals from Puerto Penasco
(Fig. 61 A), 2.5 mm in the other, 2.6 mm in the slug from St. Kitts, and
10 mm in that from Sao Sebastiao.
By re-examination of our Atlantic material and the study of that from
the Gulf of California we did not find more than individual differences,
e. g., the length of the cerebro-buccal connectives, and a shallow furrow
between the right and left cerebro-pleural ganglia.
Dr. Pickens observed that these slugs and a flatworm, we presume
Pseudoceros bajae Hyman, 1953 (Bull. Am. Mus. Nat. Hist., vol. 100,
art. 2, p. 365), have very similar colors. The polyclads generally hide
under stones during the daytime, and prey upon other animals in the dark,
while the dendrodoridids feed principally on sponges. So one can hardly
expect any biological significance of this resemblance. Presumably the
number of adult Polycladida, Opisthobranchia and other invertebrates with
planktonic stages depends greatly on the environmental factors, mainly
predation, acting upon the larvae.
32. Doriopsilla albopunctata (Cooper, 1863)
References.— O’Donoghue, 1927a: 93; Steinberg, 1961: 58; MacFarland,
1966: 196.
Synonyms.— Doriopsilla fulva MacFarland, 1905: 45; 1906: 130; 1966:
194 ( Dendrodoris fulva). Doriopsis reticulata Cockerell & Eliot, 1905: 41.
Material.—Puerto Penasco, Sonora, Mexico. P. Pickens, January 15,
1965. A single specimen collected in the same tide pool as Taringa aivica
timia.
Further distribution .—California, from Salt Point, Sonoma County, about
38° 25' N, to Lower California, Point Eugenia (27° 51' N, 115° 05' W).
Gulf of California, Puertecitos (30° 25' N, 114° 40' W) (Farmer &
Collier, 1963: 62). Original locality: Santa Barbara.
Remarks.—In the photograph the animal is chestnut brown with a
purplish hue, the crenulate notal margin and the rhinophores are orange,
the gills and the tail light orange. Small opaque white dots are densely
set all over the notum, with exception of the margins. Those in the mid­
line are quite small, the lateral ones a little larger, and those lying farther
outwards small again. The dots leave numerous round areas of the ground
204
color free, whose size is distinctly greater than that of a dot. In the
preserved slug these areas are slightly elevated; they correspond to sub­
cutaneous glands. Alive, the specimen was 60 mm long; preserved, its
height, 8.5 mm, is small in comparison with the length (40 mm) and
breadth (21 mm). As in other Porostomata, the mouth is minute. The
contracted pharyngeal bulb together with the oral tube is only 4.5 mm
long.
33. D oriopsilla janaina Marcus, 1967
Reference.— Marcus, 1967b: 96.
M aterial.— Puerto Lobos, Sonora, Mexico. P. Pickens, April 16, 1965.
Two specimens in the intertidal zone.
Further distribution.— Panama, Pacific side of the Canal Zone (original
locality).
D escriptive notes.— The present preserved slugs are about 20 mm long,
14 and 10 mm broad, and 4-5 mm high. In the photograph the pink color
of the notum and the generally white, but sometimes dark red tips of the
tubercles are most conspicuous. The bases of the tubercles are surrounded
by black dots, especially large in one median and two lateral bands be­
tween gills and rhinophores, and confluent in front. Two pairs of white
blotches, evidently consisting of secretion, lie on the sides. The lateral
tubercles are smaller than those in the middle. The rhinophores are red,
the gills orange, the borders of the rhinophoral pits and the branchial
pocket are white. The hyponotum is transparent, showing the bundles
of spicules.
34. Doriopsilla rowena, spec. nov.
Figure 62
M aterial.— Puerto Penasco, Sonora, Mexico. Mary Anne Hill, April 2,
1966. Seven specimens.
Description. — Alive, the animals are 8-10 mm in length. The biggest pre­
served specimen is 10 mm long, 5 mm broad, and 3 mm high. The hypo­
notum is about 2 mm in width, the foot 2.5 mm, pointed in front and
behind. The anterior pedal border is entire.
The spicules are dissolved except for traces in the hyponotum. Evi­
dently the spicules are large and lie diagonally crossed in the connective
tissue of the back, parallel to the course of the cutaneous muscle fibers.
Those in the hyponotum are mostly radial, a few parallel to the border.
The color of the living animals varies; the collector noted whitish yel­
low, pinkish orange, and yellow orange as fundamental colors. In all
specimens, small blackish brown melanophores, still present in the pre­
served material, are uniformly scattered over the smooth back. In the
205
color photographs there are also white dots. Of these, especially large
spots occur in two lateral lines from the rhinophores to the gills, and
a few big spots externally to these lines. The white elements, evidently
secretion, are not preserved.
The ground color is more intense on the rhinophores and gills; these
organs are light yellow in the whitish yellow slugs.
The tentacles are coalesced in front, forming a small rim around the
mouth. The club of the rhinophores bears 8 thick, in part incomplete,
foliations; the brim of the inconspicuous sheath is smooth. There are 5-6
pluripinnate gills; the thick anal papilla and the renal pore in front of
it lie to the left of the branchial tuft.
The eyes (Fig. 62, B, y ) are uncommonly big: the pigment cup is 80-
100 ^ wide, the lens 60 ^ in diameter. The rhinophoral ganglia are
distinctly set off from the cerebro-pleural mass, whose two halves are
separated by a furrow. The buccal ganglia (cc) lie immediately behind
the nerve ring. The pharynx ( n x ) is, in the dissected slug, similar to that
of D. pharpa Marcus (1961b: Fig. 19, h )\ it begins wide with thin,

F ig u r e62. Doriopsilla rowena, sp. nov. A, living slug from color photo; B,
anterior part of alimentary tract; C, diagram of reproductive organs.
206
glandular walls, then becomes narrow and muscular, passes through the
nerve ring, and continues narrow in its post-cerebral course. The following
wider oesophagus (os) enters a large gizzard (iz), which is as long as the
next gastric section, the stomach (so), which is especially muscular in the
present species.
The pear-shaped ampulla (Fig. 62, C, a) receives the hermaphrodite
duct (h ) not quite at its inner end, but subapically. The bifurcation of
the spermoviduct lies in the female gland mass.
The prostatic part (q ) of the thin efferent duct bears many digitiform
glandular appendages. These are much longer than in D. janaina. The
following muscular part is sheathed by muscle layers (ss) and ends with
inconspicuous spines.
The vagina (v) leaves the atrium. The further female organs are rather
similar to those of D. janaina, though the proportions are somewhat dif­
ferent. In the present species the insemination duct (in ) is much longer
than the canal which connects it with the spermatocyst ( s c ) ; in D. janaina
the insemination duct is shorter, and the stalk of the spermatocyst leaves
the spermatheca at the entrance of the vagina into the latter.
Discussion.— Besides the mentioned differences in the copulatory organs,
the size, the notal tubercles, the color, and the 18 leaflets of the rhi­
nophores separate D. janaina from D. rowena.
Another species which must be mentioned in connection with D. rowena,
due to its range and the lobate prostate, is D. albopunctata. This large
species is quite different from D. rowena. Three species of Doriopsilla
from West Atlantic warm waters (areolata Bergh, leia Marcus, and
pharpa Marcus) have the prostatic part of the efferent duct thickened,
but not beset with digitiform appendages.

35. Tritonia (Candiella) pickensi, spec. nov.

Figure 63
M aterial.— 1. Puerto Penasco, Sonora, Mexico.— 2. Guaymas, Sonora,
Mexico. P. Pickens and Mary Anne Hill, June 27 and July 11, 1965, and
July 1, 1966. Ten specimens, from gorgonians living intertidally and sub-
tidally. “Each gorgonian had at least four of these opisthobranchs” (Dr.
P. Pickens).
Description.— In the photographs on a ruler, the living animals reach a
length of 7 mm and a maximum width of 2 mm at the anterior border of
the foot, including the gills about 3 mm. The rhinophores are 2 mm long.
The length of the preserved slugs is 2-5 mm.
The animals are transparent white with the pinkish intestinal gland
shining through the skin. In one photograph, the sides of the back are
blue. A broad opaque white band runs along the middle of the back,
bifurcates towards the rhinophores, and expands towards the base of each
207
gill (Fig. 63, A ). In the preserved material this band is not retained; only
the opaque white gonad and the yellowish liver are recognizable.
The notum is smooth, and its border is webbed between the bases of
the gills. This may be a consequence of contraction, because it does not
appear on the photographs. The foot is round in front and pointed behind.
The veil is rounded, without a median notch, and bears 2-4 digitate ap­
pendages (vo) besides the auriculate tentacles ( t ) . In one specimen, 3
mm preserved, 6 velar appendages were found.
A process (Fig. 63, B, bi) on the outer side of the rhinophore sheath,
basally connected with the notal web, corresponds to a gill, as occurs on
the rhinophores of Dendronotus and Bornella. The rhinophore sheath
( rs ) widens distally forming a cup with a slightly ragged rim. The 5
plumes ( u ) around the rhinophoral club (io) are simple in young, pin­
nate in older slugs. There are 7-11 gills ( b) on either side, without exact
symmetry. The gills dichotomize and may have 8-12 tips, but some of them
do not branch much.

F igu re 63. Tritonia (Candiella) pickensi, sp. nov. A, living slug from color
photo, dorsal view; B, lateral view of right rhinophore; C, cones of masticatory
border; D, radular teeth; E, copulatory organs.
208
 The genital apertures are located between the second and the third right
branchial tuft; the anus lies between the fourth and fifth gills, hence
behind the middle of the body in small animals but slightly in front of
the middle in larger ones. The renal pore is a little in front of the anus.
The masticatory border of the jaw (Fig. 63, C) bears about four rows
of smooth, pointed teeth. The radula (Fig. 63, D) has the formula 25-
28X10-11.1.1.1.10-11. The teeth are colorless or yellow. The tricuspid
rhachidian tooth is only a little broader than high, its middle cusp a little
longer than the lateral cusps. These fit into the concave inner side of the
first lateral teeth. The cusp of this intermediate tooth bears 6-8 smaller
denticles; one stronger point in front of them is not always distinct. All
following teeth are smooth. A malformed third row of two coalesced teeth
was noted in one of the radulae examined.
The ovotestis overlies the intestinal gland. The ampulla (Fig. 63, E, a)
is slender, fusiform. As systematically important organs of the reproduc­
tive system, the copulatory organs were especially considered. The inner
muscular section ( d ) of the sperm duct is rather long, though shorter
than the following prostatic part ( q ). The penial part ( p ) of the male
duct ( e ), which lies in the atrium ( am) , is not quite as long as the pro­
static section. In two preserved 5-mm slugs the prostates were 0.68 and
0.45 mm long, the penes 0.51 and 0.3 mm. In correlation with the
slender, relatively long penis the bursa canal (vagina, v) is longer than
the ampulla (s) of the bursa (spermatheca).
The species is named for Dr. Peter E. Pickens.
Discussion. — The editors of MacFarland’s posthumous work (1966)
intentionally left it in its original form (p. V III). Therefore Odhner’s
(1963) names of the Tritoniidae were not considered. In Odhner’s nomen­
clature Duvaucelia is restricted to species with the anus far in front of the
middle and the genital apertures near the first gill. So MacFarland’s species
of Duvaucelia would be called Tochuina (tetraquetra) and Tritonia (the
3 other species).
Of the three subgenera of Tritonia, only Tritonidoxa Bergh, 1907, is
well characterized by its flagelliform penis (Odhner, 1926: 35). The dis­
tinctly bilobed veil with numerous short processes of Tritonia and the
evenly rounded veil with at most a slight median notch and up to 10 digi­
tate appendages of Candiella separate the type species well (Larsen,
1925: 12), principally because their sizes are very different (Tritonia
hom bergi up to 200 mm, Candiella plebeia up to 30 mm). However,
species of intermediate size, specimens not full grown, and preserved
material cannot always be allotted safely to one or other of these sub­
genera, as already stated by Eliot (1905: 17; 1910: 146).
Tritonia pickensi is an indubitable Candiella. The only Candiella among
MacFarland’s species (1966) is Tritonia festiva (Stearns, 1873). Its in­
nermost lateral tooth may bear denticles (MacFarland, 1966: 222).
209
However it differs from T. pickensi by reticulate lines on the sides and
dorsum; absence of an appendage on the rhinophore sheath; 10, ac­
cording to MacFarland even 12, velar processes; and a completely dif­
ferent penis (Marcus, 1961a: Fig. 112; MacFarland, 1966: PL 44, Fig. 2).
Another Californian tritoniid might belong to Candiella, the little known
(Marcus, 1961a: 32) Tritonia palm eri Cooper, 1862. If the comparison
is based upon Cockerell & Eliot’s specimen (1905: 33), the rugose back
with small warts and the roughly 10 velar appendages separate palmeri
from pickensi.
The species most nearly related to T. pickensi is T. (Candiella) bayeri
from Florida, also with an appendage of the rhinophore sheath. It has a
diffuse net of opaque white lines on the notum; the velar and rhinophoral
appendages are peppered with white; the mandibular cones have a longitud­
inal sculpture; the rhachidian tooth is much broader than high; there is a
denticulate second lateral tooth besides the first, and the copulatory organs,
penis and vagina, are shorter than in the species from the Gulf of California.

36. Dendronotus nanus, spec. nov.

Figures 64-65
M aterial.— Puerto Penasco, Sonora, Mexico. Dorothea Caskey, February
13, 1965. “They were found floating at the surface in about 18 inches of
water. They swim with lateral bending movements. They are rare.”
For the swimming of D . iris (Cooper, 1863) see Agersborg, 1922: 261-
262.

D escription.— The expanded and the contracted slug in the photos are
semitransparent greyish with a slight reddish hue, few scattered opaque
white dots, and an opaque white line along the edge of the foot. The
tips of the outermost ramifications of the appendages are white; then
follows a blackish brown ring and an orange base (Fig. 64, A ).
The skin of the preserved slugs contains a little brown pigment. By local
contraction darker zones are brought about, e.g., on the edge of the foot,
where an inner dark line accompanies the outer white one. The larger of
the two animals is 13 mm long, 4 mm high without the cerata, and 3 mm
broad. The corresponding measurements of the smaller slug are 10, 3, and
2 mm. The fore end is semilunar, the body not sharply set off from the foot,
the tail end pointed; the anterior border of the foot is bilabiate and evenly
rounded to the sides. The rhinophores have about 15 leaves; their tips
are light.
The frontal veil bears three rows of papillae (Fig. 64, B ). That nearest
to the mouth comprises about 20 white nodules. Then follow about 10
papillae of different size. They are decorated with a dark ring and bear a
few small white knobs. The dorsal-most row of the frontal veil consists of
210
about three larger compound papillae of almost equal size on each side.
The white tips and orange bases have faded out; only the brown rings are
preserved. The rim of the rhinophoral sheath bears about five papillae,
some simple, some compound. The papilla on the outer posterior side of
the rim is highest. Two or three compound papillae occur along the back
of the sheath between rim and base (Fig. 64, A ). Immediately behind the
outer side of the base there is a fan-shaped appendage with a broad base
and about six compound papillae on its free border. The male opening
lies behind this appendage on the right side.
The body bears six pairs of flabellate appendages (cerata), the foremost
of which are the largest. The broad trunk emits about five branches
terminating with 4-10 brown bulbs beset with numerous light papillae. The
ramification of the cerata is not dichotomous. The female opening lies
under the first right ceras. Farther dor sally than the reproductive apertures
the anus is located in the pericardial interspace between the first and
second ceras. The second cerata lie a little behind the middle. Of the four
branches which go out from the thick trunk, the second from the middle is
a trifle longer than the others, and the same holds, though still less
pronounced, for the foremost ceras. The following cerata stand near to

F igure 64. Dendronotus nanus, sp. nov. A, lateral view of fore end, combined
from color photo and preserved specimen; B, head of larger preserved speci­
men in ventral view.
211
one another and decrease in size backwards; the hindmost of the right side
is represented only by one bulbar papilla. This description refers to the
preserved 13-mm slug; in the smaller one the number of appendages,
especially on the veil and the rhinophores, is much smaller.
Thompson (1960) had only quite small (2 mm) and full-grown (90
mm) specimens of D. frondosus for the examination of the intestinal
diverticula in the cerata. In our 9- and 13-mm-long animals the liver
branches do not extend into the cerata.
The yellow mandibles of the larger specimen are 4.0 mm long, 1.3 mm
broad behind and 3 mm in front, where 1.7 mm belong to the superior
process, 1.3 mm to the masticatory border. The latter bears about 90
crested pointed denticles. The radula (Fig. 65, A) comprises 26 rows.
There are 9-14 lateral teeth per half-row. The rhachidian tooth has about
10 strong denticles on either side of the cusp. The lateral teeth increase in
size to the sixth and decrease outwards. The innermost lateral tooth is
shorter than the following ones. Its broad cusp bears up to 12 denticles.
In the old rows of the radula the second to sixth or even eighth lateral teeth
have up to six denticles; in the newer rows two or three, and in the newest
ones only a single denticle or none at all may appear. The salivary glands
are long and voluminous as in D. frondosus (Bergh, 1879a: PI. 3, Fig.
12). The posterior and the right and left anterior livers open by separate
ducts into the stomach, as drawn by Odhner (1936: 1064, Fig. 4).
The reproductive organs of the larger specimen (Fig. 65, B) are juvenile
but not primordial. The male duct ( d ) is very long and convoluted, with
beginning glandular differentiation of the ental half, hence similar to adult
D. iris (Odhner, 1936: Fig. 40). Also the cylindrical copulatory organ
agrees with iris. The rudimentary bursa copulatrix (Fig. 65, B, s) is the
same as in frondosus and iris; a distinct differentiation of the vagina into
vagina proper and vesicula seminalis is not developed, but in any case the
type is more like iris than like frondosus, robustus, and MacFarland’s
species (1966: PI. 50, Figs. 2-4), whose vesicle (MacFarland’s “sperma­
theca” ) communicates with the vagina by a duct. The great length of the
ectal, non-glandular, efferent duct is peculiar in the present species, and
the difference between our Figure 65, B, and that of a young specimen of
D. iris (30 mm; Odhner, 1936: Fig. 41) is obvious.
R em arks. — If Odhner’s synopsis is followed, D. gracilis (Baba, 1949: 87,
167) and D. nanus must be assigned to the first group with “obsolete”
veil, hence not protruding laterally as in D. robustus (Bergh, 1894: 142).
The second character of group I, “velar processes about 3-4, largest to­
wards the median line of the body, smaller laterally” does not apply to
D. nanus. The cerata of the hitherto described species of Dendronotus,
including also MacFarland’s (1966: 265-279), are arborescent, each
containing one or two main stems and looking very different from the
212
F igu re 65. Dendronotus nanus, sp. nov. A, innermost teeth of radula; B,
diagram of reproductive organs, the efferent duct drawn as a simple line.

flabelliform cerata of D. nanus. The comparison, it is true, needs living

material or photos; in our preserved specimens the cerata were more
pedunculate than in the photographs. We compared especially a color
photo of D. iris which we owe to Dr. James R. Lance, and Agersborg’s
figures (1922: Figs. 1-4), because that species agrees with nanus in the
rhinophoral appendages, the white line on the edge of the foot, and has
similar reproductive organs. As preserved specimens of iris reach the size
of 210 mm (O’Donoghue, 1921: 188), one might ask whether nanus was
a dwarf form of iris. The number of lateral teeth does not exclude this
possibility. According to Odhner, iris has “6-12” teeth, less than nanus
with 9-14. However, Odhner inverted O’Donoghue’s “ 12-6”, certainly a
213
lapsus for “ 12-16.” In an about 9-mm-long specimen of D. iris the cerata
have already “the characteristic form and arrangement” (O’Donoghue,
1921:91), i.e. “the first three or four pairs arise as practically two main
trunks right from the back, and each soon subdivides.” This description
matches Lance’s and Agersborg’s figures, but not the present animals.
We do not consider the details of the color, e.g., the blackish brown
subapical ring of the white-tipped papillae, and the denticulation of the
older lateral teeth as differential characters in view of their notorious
intraspecific variability in Dendronotus.

37. D oto amyra Marcus, 1961

R eference.— Marcus, 1961a: 38.
M aterial.— Puerto Penasco, Sonora, Mexico. Mary Anne Hill, February 5,
1966. One specimen.
Further distribution.— Monterey Bay; Point Pinos, Pacific Grove.
R em arks.— The present specimen, ivory in both living and preserved
condition, was 7 mm long in life; preserved, it measures 4.3 mm. Its six
pairs of cerata bear 4-6 rings of elongate, white-tipped tubercles. The
rhinophoral sheaths are wide, their border is smooth. The gills are short;
gills of considerably different lengths were noted in the original description.
The remarkably high anal papilla is also developed in the present specimen.
The other species of the genus recorded from the coast of California, as
well as the recently published D. varians MacFarland (1966: 289), differ
from D. am yra by shorter, more or less hemispherical tubercles of the
cerata.
38. Doto lancei, spec. nov.
Figure 66
M aterial.— Puerto Penasco, Sonora, Mexico. Mary Anne Hill, February 5,
1966. Five specimens crawling on Aglaophenia.
D escription.— Alive the largest slug is 7 mm in length, preserved 5.2 mm;
the height of the cerata reaches 3 mm in the photographs, 1.2 mm
preserved. To the light brown ground color of the living specimens blackish
brown pigment is added on the veil and the rhinophoral sheaths, but the
borders of the latter and the veil are light. There are up to six circlets of
ceratal tubercles. In the three basal ones the tubercles are light with white
dots, the cutaneous glands. In the farther distal ones they bear a black
subapical ring, and most of them a black apical spot. In the cerata that have
fallen off (Fig. 66, D) a black spot is seen on the inner side of the base.
Its pigment lies in the parenchyma of the ceras and that of the notum.
The cephalic veil projects laterally beyond the anterior border of the
foot and bears a ridge in front of each rhinophore. The rim of the
214
rhinophore sheaths is smooth and scalloped in the two smallest slugs; in
the middle-sized one it is slightly scalloped and a little projected in front.
In the two large slugs the rim is scalloped and forms a spout in one animal,
while it is uniformly thick, without expansion, in the other.
Three of the slugs have seven pairs of cerata, the smallest specimen has
six, the biggest eight. The cerata in the middle are the highest, but the
difference is not considerable. The conical apical tubercle and those in the
3-6 circlets are of the same type; in the preserved slugs they are blunt. The
number of tubercles on the cerata varies widely. The biggest cerata of the
largest slug (Fig. 66, B) have 20, while those in the second bear 35, up to
9 in one circlet (Fig. 66, C ). The 2 basal circlets are interrupted on the
inner side by the oblique insertion of the ceras and the 2 following ones by
the broad, branched gill (Fig. 66, D ).
The radula (Fig. 66, E) comprises 86 complete and some young rows.
The position of the denticles of the teeth alternates in succeeding rows,
and they are asymmetrical on the sides of the central cusp. The left-sided
posterior oral gland, a common character of D oto and M iesea Marcus
(1961b: 148), is also much bigger than the right one in D. lancei.
The reproductive organs are essentially the same as in other species of the
genus; a specific character is the straight course of the ejaculatory duct
through the penis, which is 0.4 mm long and 0.095 mm in diameter
(ratio: 4.2:1).

F i g u r e 66. Doto lancei, sp. nov. A, living slug from color photo; B, ceras
with few tubercles; C, ceras with many tubercles; D, same seen from inner
side; E, radular teeth.
215
 The species is named for Dr. James R. Lance of Scripps Institution of
Oceanography, La Jolla, California, who sent us a beautiful kodachrome
transparency (his No. 34) of a slug from the Gulf of California which
evidently belongs to the species described here.
Discussion.— D oto columbiana O’Donoghue (1921: 204) from Vancouver
Island and Dillon Beach (38° 15' N) agrees with D. lancei regarding the
subapical ring around the ceratal tubercles, an uncommon character in
D o to , and the shape of the gills (Marcus, 1961a: Fig. 128). In D. colum ­
biana, however, neither apical spots of the tubercles occur, nor black spots
on the inner side at the insertion of the cerata. Already the classification of
the specimens from Dillon Beach, which are smaller than those from
Vancouver and have more cerata, may be criticized. A further amplification
of the concept of D. columbiana to include slugs with still more cerata than
occur in the animals from the Vancouver region and from Dillon Beach,
with color marks not recorded from the original material and absent in all
specimens from Dillon Beach also, and with a different border of the
rhinophoral sheath, is in our opinion untenable. The ejaculatory duct
winds through the penis in D. columbiana (ratio: 3.5:1), while its course
is straight in that of D. lancei (ratio: 4.2:1).
D. kya Marcus (1961a: 39) from Pacific Grove (36° 38' N) has eight
pairs of cerata and black spots at their insertions, but neither rings nor
spots on the cerata. The latter insert axially in kya, obliquely in lancei
and columbiana.
Another species with a basal ceratal or notal spot and with a range
reaching the Caribbean is D. cinerea Trinchese, 1881 (Marcus, 1963: 38).
Its gills are indistinct. The cerata have light tips, and the ceratal tubercles,
which are set somewhat irregularly, have neither rings nor apical spots.
The variable shape of the border of the rhinophore sheath in D. lancei
cannot once and for all invalidate this character in all other species of D oto,
but it is not always reliable (Marcus, 1961a: 41).

39. Dirona picta Cockerell & Eliot, 1905

Figure 67
R eferences.— Cockerell & Eliot, 1905: 46; MacFarland, 1912: 517; 1966:
296; Marcus, 1961a: 44.
M aterial.— 1. Cholla Bay, Puerto Penasco, Sonora, Mexico.— 2. Puerto
Penasco. Mary Anne Hill, February 5, 1966. “Found crawling among
hydroids (Aglaophenia) in a tide pool on the rocky shore.”
Further distribution. — From Dillon Beach to the Pacific and Gulf coasts
of Lower California, north to Puertecitos, 30° 25' N (Farmer & Collier,
1963: 62). Farmer (1967: 342) found it at Puerto Penasco. The occur­
rence in the Sea of Japan (Volodschenko, 1955) is taxonomically not
certain (Marcus, 1961a: 46).
216
Descriptive notes .— The living animals were 37 mm (1) and 10 mm (2) in
length. The adult one was orange, the juvenile (Fig. 67, A) white. The latter
shows the preradula passing into the definitive one (Fig. 67, B). The
rhachidian tooth (R ) and the inner lateral tooth (I) of the preradula of
D. picta are similar to the rhachidian tooth and the single lateral tooth in
the adult radula of M adrella (Vayssiere, 1903: PI. 3, Fig. 31; Baba,
1949: Figs. 99-100, 102), a neighboring genus of the Arminoidea Pachy-
gnatha Odhner (1939: 48).
During the transport to Tucson, the full-grown slug laid an egg string,
about 40 cm in length, consisting of interrupted, coiled sausage-shaped
sections, lighter and darker brown in the photograph, white in preserved
condition. Each section is about 5 mm long and 1 mm in diameter. In one
plane it comprises 10 egg capsules each about 0.2 mm in diameter. There
are about 10 eggs, 80 ^ in diameter, per capsule. Each cm of the string
contains about 10 sausages, so that 800,000 to 1,000,000 eggs can be
computed for the whole string. The egg mass is much more like that of
D. albolineata (O ’Donoghue, 1922c: 132, Fig. 2) than that of D. aurantia-
ca Hurst (1966: 9, PI. 2, Fig. 4 ).
Nom enclatural note. — We follow Farmer & Collier (1963: 62,) Lance

F i g u r e 67. Dirona picta Cockerell & Eliot. A, young animal; B. rhachidian

teeth (R) of rows 1, 10, 16; inner lateral teeth (I) of rows 2, 16; outer
lateral teeth (O) of rows 9, 16.
217
(1961: 68), O’Donoghue (1926: 224), and Steinberg (1963: 71) indi­
cating Cockerell & Eliot, 1905, as authors of the present species. In view of
the history of the discovery (MacFarland, 1912: 515 “aided by memor­
anda” ), article 50 of the Code of Zoological Nomenclature of 1961 (“some
other person is alone responsible” ) is not applicable to the use of Mac-
Farland’s name as author of Dirona, D. picta, and D. albolineata, all
contained in and characterized by Cockerell & Eliot, 1905. The first author
of the family name Dironidae is Eliot (1910: 69).
40. Coryphella iodinea (Cooper, 1862)
Figures 68-69
References. — Cooper, 1862: 205; Cockerell, 1901a: 121; O ’Donoghue,
1927a: 103; MacFarland, 1966: 308 (Flabellinopsis iodinea); Farmer,
1967: 342.
M a teria l — Puerto Penasco, Sonora, Mexico. Robert Taylor, June 1, 1965.
A single specimen was found moving around some tunicates.

F ig u r e s 68-69. Coryphella iodinea (Cooper). 68, half-row of radula; 69,

diagram of reproductive organs.

Further distribution. — Monterey area (Smith & Gordon, 1948: 181);

San Pedro; Laguna Beach; San Diego (original locality); Lower California,
Los Coronados Islands (Lance, 1961: 68), and Ensenada (Farmer &
Collier, 1963: 62). The locality “Puget Sound” is doubtful (see Remarks).
218
Description. — The photographs show purplish, red, and orange cerata
with the outer fifth creamy white, and the pinkish viscera shining through
the violet skin. The rhinophores are brick red and have a cream stripe
along their anterior mid line. The long tentacles are blue with white ends.
The corners of the foot are short. The short tail is violet.
The colorless preserved specimen is compressed. It is 35 mm in length,
14 mm in height, and 4 mm in width. The tail is pointed; the groove of the
anterior border of the foot continues along the pointed anterolateral
corners. The tentacles are wrinkled due to contraction. The rhinophores
are shorter; their clubs bear the uncommonly high number of 80 leaves on
either side. The genital pore lies at the end of the anterior fourth, the anus
almost in the middle of the body length, under the most lateral cerata, in
typical pleuroproctic position.
The flabby cerata form about eight indistinct groups, which leave the
middle of the back free. They occupy a narrow band, the hindmost cerata
forming a single row. In front their arrangement is branched, fan-like,
similar to Odhner’s figure of C. rufibranchialis (1929: Fig. 5). The largest
cerata, 10 mm long, are the innermost ones. The cerata are not united on
peduncles; each is inserted directly on the back. The digestive diverticulum
in the cerata is smooth and passes gradually into the canal which leads to
the cnidosac. The cnidosacs are 0.35-0.47 mm long, hence one-twentieth
of the cerata.
The thin jaws are colorless; their masticatory process bears many rows of
blunt denticles. The minute buccal bulb is 2.7 mm in length and 2 mm in
diameter; the ribbon-shaped salivary glands are rather long. The radula
(Fig. 68) comprises 22 rows. The rhachidian cusp is flanked by 14-16
denticles on either side; the inner edge of the lateral teeth has 11-16
denticles.
The racemose hermaphrodite gland lies ventral to the gut. The her­
maphrodite duct (Fig. 69, h ) dilates into a coiled ampulla ( a ) which has
a black stripe. The spermoviduct divides into the inner oviduct (o)
entering the female gland mass (g) and the male duct. The latter is
prostatic ( q ) in its long, convoluted course. It ends in a thick, muscular
penis papilla ( p ) which is not cuticularized. The epithelium on the papilla
and that of the male atrium ( am) is ciliated. The ejaculatory duct opens
in the center of the papilla, not into an external furrow (Marcus, 1961a:
48). The vagina is not separated from the nidamental duct ( n i ) the
termination of which is strongly muscular. On one side the female duct
bears an ample pouch, the spermatheca (s').
Rem arks. — The five groups of cerata originally described are not always
recognizable. Cockerell said: “separate tufts not obvious.”
Bergh (1879c: 559, note 2) examined the original specimens of
Cooper’s A eolis (Phidiana?) iodinea and assigned them to Flabellina
Cuvier, 1830. The principal character of this genus is the insertion of
219
groups of cerata on simple, bifurcate or trifurcate sockets, pedaments or
foot-stalks (Alder & Hancock, 1855: XXI; Fischer, 1887: 539; Vayssiere,
1913: 294; Thiele, 1931: 451; Pruvot-Fol, 1954a: 413). In fact, Bergh
(1879a: 80) recorded in a single, “most badly preserved” specimen from
Puget Sound, “the groups of papillae firmly affixed on the edge of foot­
stalks, whose form and number could not be determined.” Neither in our
specimen, nor in a color photograph classified by Dr. James R. Lance,
which he graciously presented to us, nor in the above-mentioned descrip­
tions does such an insertion of the cerata appear. For the moment it seems
best to maintain the name iodinea for a Californian violet Coryphella with
orange cerata and without peduncled supports, applying Cockerell’s
generic allocation. The “low, crescentic, ridge-like elevations of the
dorso-lateral body margin,” which bear the cerata in the present species,
led MacFarland (1966: 308) to introduce a new genus for it. Though this
proceeding stresses the elevated areas of the ceratal bases in a convenient
manner, it duplicates the difficulty to separate between Coryphella and
Flabellina. If one compares MacFarland’s painting (PI. 58, Fig. 1) with
Dr. Lance’s and Dr. Pickens’ photographs, the ridge appears much more
accentuated in the former. In preserved material the natural height of these
ridges cannot be evaluated, while common foot-stalks of the cerata appear,
when these appendages are plucked. A separation of two families, Flabelli-
nidae and Coryphellidae (Pruvot-Fol, 1954a: 413, 421) is not natural. The
single family must be called Flabellinidae according to its oldest genus.

41. Coryphella cynara, spec. nov.

Figures 70-75
M aterial .— Cholla Bay, Puerto Penasco, Sonora, Mexico. L. Pardy, June
29, 1965. “Only a single specimen of this swimming nudibranch was found.
It was lying on the sand at low tide line. It swims with a ‘breast stroke,’
moving the cerata in horizontal plane along the side of the body in unison
and extended. On the return stroke the cerata are collapsed, but return
along the same plane.”
D escription .— The photographs show a semitransparent pigeon-blue animal
with a short, pointed tail, produced but not very long anterior angles of the
foot, long tentacles, strong rhinophores, and high, slender cerata (Fig. 70).
The border of the foot is accompanied by a blue line, the back of the tail
is dark blue and crested with white; a white ogee arched line begins be­
tween the rhinophores and extends on to the bases of the tentacles. There is
a white zone under the blue tips of the tentacles. Also the tips of the foot
angles are white. The cerata are orange brown with a red ring in their distal
fourth and end with a white point. The cerata conceal the ground color of
the back. There is a large reddish patch on the genital region and smaller
violet blotches near the tips of the rhinophores, on the bases of the foot
corners, and two on either tentacle.
220
 The blue color has passed into the alcohol; the rings near the tips of the
cerata, the patch, and the rhinophoral blotches, and a band on the tail
are cherry red.
The length of the preserved specimen is 12 mm, the breadth 4 mm, the
height 3 mm without the 4-mm-long cerata which had fallen off. The
tentacles are 5 mm in length, the rhinophores 3 mm. The latter are perfoli-
ated and have about 30 leaves. The gonopore lies at the end of the first
third of the body length. The anus ( a r ) in front of the first right side
group of the left posterior liver branches (/) is more dorsal than in the
preceding species. The heart lies far back, between the first rows of the
posterior liver. The foot, rather distinctly set off from the flanks in the
photographs, passes to the sides gradually in the preserved slug. The
anterior border of the foot is bilabiate.
There are six slanting cushions of cerata (Fig. 71), farther ventral in
front, and more dorsal behind, each with an irregular series of caducous,
thin cerata. The intestinal diverticula in the cerata are smooth; the cnidosac
occupies about one-tenth of the ceras.
The breadth of the bluish grey jaw (Fig. 72) is 68 per cent of its length;
the masticatory process is strong. Its border bears denticles about 50 /x
high in one or two rows (Fig. 73), which work into those of the other jaw.
Sidewards to them there is an area of smaller tubercles, decreasing in size
and arranged in 15 irregular rows.
The radula (Fig. 74) comprises 16 rows. The rhachidian tooth is narrow,
0.4 mm in length, 0.22 mm in breadth. It bears 7-9 denticles on either side
which nearly reach the tip of the median cusp, where they are smallest.
The lateral teeth are slender, obtuse-angled triangles with 12-15 denticles
on one of the short sides. The ratio of length (290-330 /x) to breadth
(74-82 fi) is 4:1. The pharynx is 3.2 mm long, hence rather large.
The gonad is ventral to the main liver duct. The reproductive organs
(Fig. 75) are in the male phase; the female gland mass (g) is small. The
coiled, black ampulla ( a ) enters the gland mass, from which the winding
prostatic section ( q ) of the efferent duct emerges. This looped part is
knobbed due to its glandular sacs. The short ectal muscular efferent duct
( d ) enters the large, muscular male atrium. The penial papilla ( p ) which
projects from the male opening ends nearly globular and is beset with
epithelial warts.
The glandular oviduct (g) opens into an ample, muscular female atrium
which egresses (aw) immediately behind the male aperture. Within the
muscular wall of the atrium lies a spacious pouch, the spermatheca (s).
Discussion — A similar narrow lateral tooth, although a much shorter one,
occurs in C. falklandica Eliot, 1907, widely distributed in the southern
hemisphere (Odhner, 1944: 19; Marcus, 1959: 71). Its anus lies under the
third to fourth row of the second group of cerata. C. cooperi Cockerell
(1901c: 85) has a green patch on the left side of the heart. Its lateral
221
F i g u r e s 70-75. Coryphella cynara, sp . n o v . 70, liv in g s lu g f r o m c o lo r p h o to ;
71, r ig h t s id e v ie w w it h o u t c e r a ta ; 72, ja w ; 73, m a s t ic a to r y b o r d e r o f jaw ;
74, r a d u la r te e th ; 75, d ia g r a m o f r e p r o d u c t iv e o r g a n s.

tooth was compared with that of a European species, “except that the
lower corner is very much more produced.” As the general color differs
widely from that of cynara, and only the white tips of the cerata agree,
Cockerell’s description without figures is far from sufficient to approach
cynara to cooperi. As Steinberg said (1961: 62), only further collection
in the San Pedro area can establish the identity of C. cooperi. The Carib­
bean C. dushia Marcus (1963: 41) is morphologically rather different
from cynara.
Steinberg united (1963a: 66) C. piunca Marcus (1961a: 47) with
C. trilineata O’Donoghue (1921: 197; 1924: 25). The single examined
specimen of the latter had a U-shaped median tooth, and the longer side
of the triangular lateral tooth (the hypotenuse, as O’Donoghue called it)
is denticulate. The jaw was not figured, only compared with that of
C. fusca O’Donoghue (1921: 195), in which the breadth is 76 per cent of
the length. The median tooth of piunca (loc. cit.: Fig. 165) is not U-
222
shaped, the lateral tooth bears denticles on one of the shorter sides of the
triangle, and the breadth of the jaws is 63-70 per cent of its length. If these
characters were found to be connected by intermediate types, and inhabi­
tants of the type localities, the Vancouver region and Dillon Beach, agreed
in their reproductive organs, the proposed synonymy could be accepted.
Many species of Coryphella have similar buccal armatures. The color pat­
tern varies intraspecifically in some cases, and in others that of different
species agrees.
In Bergh’s later lists (1891; 1892a), which comprise several problemat­
ical species of Coryphella and Flabellina, cooperi and iodinea are not
included.
From the coasts of central and northern California two new species of
Coryphella were recently published (MacFarland, 1966: 313-322). Only
one of them, C. pricei, must be compared with C. cynara, because both
have high, slender cerata. The short conical copulatory organ of C. pricei
(loc. cit.: PL 66, Fig. 8) differs widely from the nearly globular one of
C. cynara.
42. Flabellina telja, spec. nov.
Figures 76-82
M aterial. — 1. Cholla Bay, Puerto Penasco, Sonora, Mexico. Dorothea
Caskey, June 30, 1965. Two specimens about 3 m from one another “were
found in about 3 feet of water. They swim with lateral bending movements
of the body in much the same manner as D endronotus nanus.”— 2. Puerto
Penasco, Sonora, Mexico. Mary Anne Hill, June 18, 1966. One specimen
on some algae in the rocky intertidal zone.
Description. — The color in the photograph of the specimen taken in 1965
is similar to that of Coryphella cynara. The body is bluish, the back and
the area between the rhinophores and tentacles bluish grey; the cerata
contain orange brown hepatic diverticula and have white tips. The knobs of
the rhinophores, the two distal thirds of the long tentacles, and the tips of
the basally broad, not much produced corners of the foot are white, and
a white crest covers the dark blue back of the tail which ends white.
The blue color has stained the alcohol in which the animals are pre­
served. The preserved specimens are 11 and 8 mm in length; in the larger,
the body is 3 mm in breadth, the sole 2 mm. The further measurements
are: cerata up to 4 mm; foot corners ( z o ) 1 mm; tentacles (t) 3 mm;
rhinophores (io) 2.5 mm, with 30 leaves.
The slug of 1966 is white with a bluish tone, especially on the tentacles,
and with red hepatic diverticula in the cerata. Alive it was about 5 mm in
length, and the rhinophores are as long as the tentacles or a little longer.
The foot is set off from the sides, its anterior border is bilabiate and
notched. The gonopore lies under the right anterior group of the cerata; the
anus (ar) farther backwards between the socket bearing the hind-most cerata
223
of the right anterior, and that of the first row of the posterior liver.
The cerata stand in oblique rows (Fig. 76) which are more dorsal in
front, more ventral behind. The cerata are firmly inserted in groups on
common sockets (Fig. 77).
The following numbers refer to the large slugs taken in 1965. The
anterior livers (ri) have each 4 groups with 6, 5, 3, and 2 cerata; the
median cerata are, as usually, highest. The posterior liver (/) comprises
nine groups on either side, the two first each with 6, the two following each
with 5, the five last with 4, 3, 2, 1, and 1 cerata. Buds of cerata, which
were not counted, appear on the outer ends of the sockets as in many
Eolidacea. The cnidosacs are up to 0.3 mm long, hence less than 1/10 of
the highest cerata.
The colorless, delicate jaws (Fig. 79) are 1.0 mm in length, 0.75 mm in
breadth. The masticatory process is small; it bears about 10 rows of
denticles, the largest about 20 jjl in height, which stand along the border
(Fig. 80). There are 28 radular rows (Fig. 81). The rhachidian tooth is
100 fx high and 80 ^ broad. Its median cusp is flanked by 8-11 denticles.
The large uppermost of these meet over the median cusp; the following
ones are slightly narrower. The lateral teeth are 130 ^ in length, 42 ^ in
breadth. Their longest side is straight, the shortest provided with 6-8
denticles, the middle-sized side is concave. The pharynx is minute, 1.5 mm
in length in the 11-mm slug.
The mandibles and the radula of the small specimen collected in 1966
were compared and found to agree with the corresponding organs of the
larger slugs; the denticles flanking the cusp of the rhachidian tooth are
only 5-8 in number.
The gonad is dorsal to the main liver duct. The coiled, colorless ampulla
(Fig. 82, a) enters the female gland mass (g), from which the winding
prostatic section ( q) of the male duct emerges. The following portion, not
glandular, opens into the atrium ( am) . Its muscular folded walls can be
understood by the everted copulatory organ in the smaller specimen (Fig.
78). There the atrium forms a collar with a folded, warty peduncle; a
roundish penial papilla bearing the ejaculatory duct projects from the collar.
The male and female openings are united ( o a ) . A stalked spermatheca
(s) goes out from the nidamental duct (ni ). Far entally, immediately at the
entrance of the ampulla and the exit of the prostate, there is a second
seminal vesicle (sv) embedded in the mucus gland (g), similar to Odhner’s
finding (1939:61) in Coryphella pellucida (Alder & Hancock, 1846).
Discussion. — The insertion of the groups of cerata on peduncled supports
characterizes the genus Flabellina, This feature is obvious in Vayssiere’s
frequently copied figure (1888: PI. 2, Fig. 1) of the type species, F. affinis
(Gmelin, 1791). The presence of such supports led Baba (1955: 29, 52)
to transfer Coryphella ornata Risbec (1928: 266; 1953: 143; Baba, 1936:
44) to Flabellina. If this assignment is maintained, Risbec’s species must
224
F i g u r e s 76-82. Flabellina telja, sp . n o v . 76, r ig h t s id e v ie w w it h o u t c e r a ta ;
77, f o u r th r o w o f c e r a ta o f r ig h t liv e r ; 78, a n te r io r p a r t w ith e v e r te d c o p u la to r y
o r g a n ; 79, ja w ; 80, m a s t ic a to r y b o r d e r o f ja w ; 81, r a d u la r te e th ; 82, d ia g r a m
o f r e p r o d u c t iv e o r g a n s.

receive a new name, because F. ornata Angas (1864: 67) has priority.
While the peduncled supports of Risbec’s species are distinct in Baba’s
figures of 1955 and in Abe’s (1964: PI. 32, Fig. 114), they are not in the
earlier ones. Even F. affinis may exceptionally seem to have sessile cerata
(Odhner, 1939: 62). Possibly the peduncles are obscured by contraction
of the cutaneous muscles of the back.
225
 Neither perfoliate rhinophores nor anterior corners of the foot produced
into tentaculiform appendages (O ’Donoghue, 1922b: 139) are peculiar
characters of Flabellina. Coryphella californica Bergh (1904: 6) has the
front corners of the foot longer than the tentacles or the rhinophores.
Perfoliation of the rhinophoral clubs occurs in C. trophina (Bergh, 1894:
134), C. fusca, C. trilineata (O’Donoghue, 1921: 196, 197), and C. piunca
Marcus (1961a: 47).
When Pruvot-Fol (1954a: 421, note 2) called the lateral teeth different
in Flabellina and Coryphella, she probably meant the approximately rec­
tangular base of these teeth in F. affinis (Vayssiere, 1888: PL 4, Fig. 48),
from which the triangular cusp is set off. This is a specific character of
F. affinis, different only by degree from certain species of Coryphella, e.g.,
C. lineata (Loven, 1846) (Bergh, 1876a: PI. 15, Figs. 18, 20).
It is an open question whether a penial stylet is characteristic of
Flabellina (O’Donoghue, 1922b: 139). Coryphella ornata Risbec, 1928,
allotted to Flabellina by Baba (see above), has an unarmed penis (Risbec,
1953: Fig. 110, I). For F. affinis, and vaguely also for his F. iodinea
(1879a: 81), Bergh indicated a stylet (1876a: 651; 1886d: 50). Vayssi­
ere, who knew Bergh’s papers (1888: 11), called the penis of F. affinis
unarmed (p. 79, 84; 1913: 294; 1928) without any reference to Bergh’s
observation, and Bergh repeated the occurrence of a stylet in his diagnoses
of the genus (1891: 39; 1892a: 1034). Bergh’s figure (1876b: PI. 16,
Fig. 4) is not incompatible with Vayssiere’s (1888: PI. 4, Fig. 49); pos­
sibly the thick wall of the penial papilla shown by Vayssiere becomes cutic-
ularized in completely mature slugs of F. affinis. In Samla annuligera Bergh
(1900: 237) the penis is unarmed; the species seems to be a Flabellina.
The above-mentioned Flabellina from Puget Sound differs from the
present species by its iodinea-Ykt color, viz., violet purple body, orange
rhinophores, orange red cerata, and foot angles longer than the tentacles.
The lateral tooth of Flabellina telja is similar to that of Coryphella fusca
O’Donoghue (1921: 195). Along with other characters this species, in
preservation up to 19 mm in length, has 19 rows of radular teeth. The
median cusp of the rhachidian tooth bears 4-6, exceptionally 7, lateral
denticles which leave a great extension of the cusp free.
The teeth of Coryphella fisheri MacFarland (1966: PI. 65, Figs. 16-18)
are similar to those of F. telja. The color pattern, the annulate (not per­
foliate) rhinophores which are longer than the tentacles, and the absence of
a groove in the anterior margin of the foot distinguish C. fisheri at once
from the present species.
43. H ermissenda crassicornis (Eschscholtz, 1831)
R eferences .— Marcus, 1961a: 52 (literature); MacFarland, 1966: 358.
M aterial. — 1. Puerto Penasco, Sonora, Mexico. P. Pickens, January 15,
1965.— 2. Puerto Lobos. P. Pickens, April 16, 1965. “In the rocky
226
intertidal zone, quite common at the second locality.”
Further distribution. — From Sitka, Alaska (original locality), to Lower
California, Point Eugenia, and in the Gulf at Angel de la Guarda Island
(Farmer & Collier, 1963: 62); northern part of the Gulf of California
(Lance, 1966: 79).
D escriptive notes. — The known color variation of this species is evidenced
by the present photographs which show a transparent bluish and a more
opaque orange yellow slug. The tips of the cerata are white, the intestinal
diverticula in the cerata are brown.

44. Spurilla chrom osom a Cockerell & Eliot, 1905

Reference and synonym y. — Marcus, 1961a: 54.
M aterial. — Puerto Penasco, Sonora, Mexico. Dorothea Caskey and Robert
Taylor, December 1964 and March 1965.
Further distribution.— California, from San Pedro (original locality) to
Point Loma; Lower California, Ensenada (Farmer & Collier, 1963: 63);
common in the whole Gulf of California (Lance, 1966: 79).
D escriptive notes. — The photographs show two differently colored slugs.
One is white with a purplish tinge and darker intestinal diverticula in the
white-tipped cerata. Spots on the back and the antero-lateral corners of
the foot are opaque white; the tentacles and the rhinophores are light
violet. The other animal, 23 mm long alive, is yellowish orange with
opaque white spots. The cerata, foot angles, and tentacles are greenish
grey. The rhinophores are yellowish orange, their tips are frosted with
white. Some white flecks occur on the cerata.
The five preserved specimens at hand, two with white, three with dark
intestinal diverticula, correspond to the two types in the photographs.
Already the previous descriptions comprise these color variations. The
present animals are 6-8 mm in length, hence not full-grown. The long
and stout tentacles, the few (in the preserved material 7-8) leaves of the
thick rhinophores, and the tubercles visible in the everted buccal cavity
are characters also recorded by Eliot (Cockerell & Eliot, 1905: 51).
The masticatory border of the jaw is smooth. The up to 29 radular teeth
with 39 denticles on either side of the central cusp exceed the hitherto
known maxima, 21 and 32, respectively. As the present material contains
also animals with 21 radular teeth and 28 denticles, the new maxima
have no systematic importance.

45. Onchidella binneyi Stearns, 1893

Figure 83
References. — Stearns, 1878: 399 ( O . carpenteri ); 1893: 342; Semper,
1882: 281 ( Onchidium carpenteri) .
227
M aterial .— Puerto Penasco, Sonora, Mexico. P. Pickens, April 2, 1966.
Abundant in May, June and July on boulders and flat pieces of rock; at
night in the midintertidal zone.
Further distribution .— Gulf coast of Lower California, from 28° 26' to
29° N (Stearns, loc. cit.; Parker, 1964: 157, station 109, map, p. 49).
D escription .— The biggest of the seven slugs was 30 mm long measured
over the back (24 mm lineal), 15 mm broad, and 8 mm high. The
maximum breadth of the present specimens is in the middle or a little
behind it. The hyponotum of either side and the sole amount each to V3
of the width of the underside; the hyponotal line separates the two outer,
rough, thirds from the smooth innermost third.
The animals are evenly rounded in front and behind. The head is
small, and the two separate sensory lobes are apposed to one another in
the preserved slugs; the color is olive brown to black brown alive (Dr.
Pickens); the preserved animals are greenish to brownish grey, clay-
colored above, and lighter, greenish-yellowish, below. When slightly
magnified the dorsal warts are lighter than the ground color.
The warts of the notum are of three types. The largest and the medium
sized ones are beset with small warts, the big ones with up to 30, the
intermediate ones with up to 5. Between the large and medium sized warts
occur the single small warts. On either side there are 8-9 perinotal warts
on the rim which bear the ducts of the so-called repugnatorial glands.
Between every two of these warts there are 3-8 smaller ones.
The male aperture lies behind the right tentacle; the distance between
respiratory (pneumostome) and pallial (anus) openings is one-half or
less of the hyponotal breadth. In well stretched specimens the anus and
the adjacent female aperture are not covered by the tip of the foot.
The peritoneum is unpigmented; also the tissue around the nerve ring
is colorless. The connection of the salivary glands with the brain, and the
course of the intestine (Plate, 1893: PI. 8, Fig. 32) correspond to the
scheme of the genus. The radula consists of about 100 rows, each with
170-180 lateral teeth per half-row. The rhachidian tooth is about 50 p in
height, has a long, pointed, median cusp and blunt lateral cusps, which
are shorter and narrower. The first lateral measures 40 the second,
55 fi. The third, 70 ^ in length, reaches nearly the maximum size,
80 /x, which occurs farther outwards. The shape of the pleural teeth is
the same as in other Onchidella,
The ovotestis has more than 10 follicles. Of the female reproductive
organs the very wide duct of the bursa is remarkable, and the very long
and coiled vaginal gland, which justifies the name “flagellum.”
The efferent duct (Fig. 83, d ) enters the body cavity to the left of
the male aperture. It runs straight and transversely to the left side, then
bends back, enters the penial retractor (r), and opens into the inner third
228
of the penial sac ( p ) at the insertion of the muscle. The origin of the
latter in the posterior angle of the body cavity corresponds to Plate’s third
type (1893: 170). The penis sac is divided into a wider outer and a
narrower and longer middle part by a constriction. The epithelium is
high and folded. The end of the middle part bears a U-shaped caecum
(c) whose epithelium contains calcareous concretions. The length of this
caecum is 2 mm, that of the two parts of the sac together, 7 mm.

F ig u r e 8 3 . Onchidella binney i Stearns, male copulatory organ.— F i g u r e s 8 4 -

8 6 . Onchidella hildae (Hoffmann). 8 4 , living slug from color photo; 8 5 , radular
teeth; 8 6 , male copulatory organ.

Discussion.— Onchidium carpenteri Binney (1860:154) was uniformly

smoke colored, up to 5 mm in length, and 3 mm broad. The material
came from the Juan de Fuca Strait. Only by re-examination of the original
material (Smithsonian Institution) might one decide whether it perhaps
belongs to the species generally called Onchidella borealis (Dali, 1871),
a small, dark Onchidella from the Vancouver Island region. When Binney
some years later (1876: 184-185, PI. 6) studied O. borealis, he did not
mention O. carpenteri, but many years later a 14-mm-long Onchidella
carpenteri with Dali as author, without locality, appears in Binney’s work
(1890: 214, PI. 6, Figs. D & E ).
229
 Stearns (1878: 399) applied the name carpenteri Binney to specimens
with the average 17.2X12.2 mm from the western part of the Gulf of
California, San Francisquito Bay, Las Animas Bay, Angeles Bay, collected
by Mr. William J. Fisher of San Francisco. Stearns’ figures (PI. 7, Figs.
7, 8) are not specific and the description is not informative. Fortunately,
however, Semper (1882: 281) received 17 specimens from one of these
localities from the same collector who had provided Stearns with material.
Stearns (1893a: 342) replaced the name carpenteri by binneyi, and pub­
lished two good figures (PI. 50, Figs. 1, 2). These give a general idea of
the O nchidella which occurs “throughout the Gulf of California” (Keen,
1958: 512, Fig. 1037), though Keen’s indication “black” and a width
of 18 mm for a slug 20 mm long is not quite typical for O. binneyi. In
any case, Semper’s anatomical data together with ours and with Stearns’
figures of 1893 settle O. binneyi. There are slight differences between
Semper’s and our findings in the number of small warts between every two
of the perinotal ones with glandular openings. Semper found at least 8
smaller warts, we at most 8; we consider this difference as an intraspecific
variation. The same holds for the about 400 pleural teeth per radular row
in Semper’s material and 340-360 in ours. For the intraspecific variability
of the number of the pleural radular teeth in Onchidella several examples
are known: O. celtica (Joyeux-Laffuie, 1882: 253, note 1); three species
from Chile, united today (Wissel, 1898: 112); juvenile specimens have
considerably fewer teeth than fully adult ones (Marcus, 1959: 17); in
O. indolens the number per half-row varies from 75 to 100 (Marcus,
1956b: 78); and in O. borealis from 45 to 88 (Marcus, 1961a: 4).
Whatever O. carpenteri Binney, 1860, from “De Fuca Strait” may be,
the last available figures (Binney, 1890: PI. 6, Figs. D & E) show that
it is not binneyi Stearns, 1893 (PI. 50, Figs. 1, 2).

46. Onchidella hildae (Hoffmann, 1928)

Figures 84-86
R eference .— Hoffmann, 1928: 35.
M aterial .— Puerto Penasco, Sonora, Mexico. P. Pickens, April 2, 1966.
Found together with the preceding species.
Further distribution.— Ecuador, Puna Island; Panama, Pacific coast (Hoff­
mann, 1929: Fig. 1).
D escription .— The slug (Fig. 84) shown in a color photo is 18 mm long,
14 mm broad, with the maximum width in the middle, and furnished with
a total of 24 long, digitiform marginal papillae. Hoffmann found 16 such
papillae. Preserved, the length of the same animal is 26 mm (measured
over the back), the width 13 mm, and the height 7 mm. The sole is 3 mm
broad, the tuberculate hyponotum 5 mm; the hyponotal line runs near the
foot and is extended to the tentacles. The oral lobes are well divided.
230
 The ground color is greenish white; the large warts in the center of
the notum contain some brown pigment and are basally surrounded by
pigment-free stellate areas. These areas are separated from one another
by brown pigment, Hoffmann’s “dickes, braunes Netzwerk.” Small warts
are located on the large notal warts and on the pigmented stripes between
them. These warts are bigger (200 in the present species than in O.
binneyi (60 ^ ). Also the 24 perinotal warts are beset with small warts.
In the living animal the perinotal warts project on the border. Between
every two of them 4-6 (in Hoffmann’s material 2-4) small light warts lie
on the frilled perinotum. The underside of the animal is light.
The position of the male aperture to the right of the right tentacle,
and that of the female orifice to the right of the pallial opening, are
as in other Onchidella, The pallial opening is close behind the tip of the
foot; the distance of the pneumostome, also median, from it is V3 of the
hyponotal breadth.
The radula (Fig. 85) comprises 70 rows with 66 pleural teeth per half­
row. The median cusp of the rhachidian tooth is straight and pointed; the
lateral cusps are curved and blunt. As in other congeneric species the two
innermost lateral teeth are smaller (26 and 42 ^) than the others (up to
60 /j,). The mesocone of the third and the following teeth in the two
inner thirds of the half-row bear fine denticles, suggestively present also
in the preceding species. The outer third has no denticles, and the teeth
diminish in size. The oesophageal crop is extremely distended by its
contents, as in one of Hoffmann’s specimens. The intestine courses as
usually in Onchidella.
The ovotestis comprises about 8 follicles. The coils of the hermaphrodite
duct which act as a seminal vesicle, and its caecum, a ciliate diverticulum,
are big. The bursa canal is wide, the bursal ampulla relatively small.
The vaginal gland is short and thick, very different from that in the
preceding species. The efferent duct (Fig. 86, d ) enters the body cavity
as in O. binneyi, but its course is very different. It accompanies the male
atrium, forms a free spiral, descends, enters the retractor (r), ascends, and
opens into the slender penial sac (p) in front of the origin of the caecum
(c). The male organ itself is similar to that of binneyi, but the caecum is
straight, 0.7 mm in length, the glandular folded sac is not distinctly divided,
and together with the atrium only 3 mm long. The penial retractor corre­
sponds to the third type, as indicated by Hoffmann (1928: 37).
Discussion.— As Hoffmann (1928: 38) said, O. hildae is closely related
to O. binneyi. He based the difference between them upon the penial
caecum, curved in binneyi, straight in hildae; now also the course of the
efferent duct in the body cavity, and the sizes of the vaginal gland can
be added as considerable differences. Concerning a plain dark color and
one with light areae (Marcus, 1956a: Figs. 1, 2), a smooth (Figs. 4, 5),
prickly (Fig. 7), or puffy skin (Fig. 3) we are cautious; but the size of
231
the smallest warts, 60 ^ in binneyi, and 200 ^ in hildae, is obviously
different. The narrow sole and the broad hyponotum, already observed by
Hoffmann, but not used as differential feature, would alone be weighty
reasons for a specific separation.
Though Hoffmann called the penis “rather short,” the little penis of
the present specimen and the small number of radular teeth, not mentioned
by Hoffmann, might possibly suggest a juvenile state of our single slug.
But its body size is the same as that of Hoffmann’s largest animals (25
mm measured over the back), and also the sperms in the coils of the
hermaphrodite duct and the volume of the mucus gland define the present
animal as adult. Therefore the size of the penis and the number of teeth
can be considered as further specific characters.

47. Hoffmannola hansi, spec. nov.

Figures 87-95
M aterial .— El Sahuaral, a few miles south of Kino Bay, Sonora, Mexico.
Mary Anne Hill, June 22, 1966. Found at night on the higher intertidal
rocks, where they were quite numerous. Also at the same locality, Novem­
ber 12, 1966, P. Pickens. Fourteen specimens sent.
Further distribution .— Probably Angeles Bay and Puerto Refugio on the
Angel de la Guarda Island, during day-time, “exposed to the killing sun”
(Steinbeck & Ricketts, 1941: 220, 226, 545). April 1, 1940.
D escription .— The biggest slug is 48 mm long measured over the back (26
mm lineal), 20 mm broad, and 16 mm high. In the least contracted
specimen the corresponding measurements are 34 (27), 23, 11. The
width of the hyponotum is 5 mm, the sole measures 18 X 15 mm.
The animals are nearly truncate in front, rounded behind; the maximum
breadth lies a little behind the middle. The sensory lobes form a continuous
veil with an entire rim in front of the mouth. The notal border is smooth,
without frills or projecting warts. Also the hyponotum is not warty. The
innermost fourth of the underside of the notum becomes darker than the
outer part when the surface is stained weakly with carmine. By this treat­
ment a few minute pores appear in this dark-staining stripe (Fig. 88),
whose limit against the light outer part corresponds to a hyponotal line.
Outside this line the epidermal cells of the hyponotum bear cuticular cones;
the cells of the innermost, darker staining fourth are not cuticularized.
The color is rather uniform black in the center of the notum, brownish
toward the margins. The whole underside is light (Fig. 88). Scattered over
the notum are large, flat warts on lighter blotches (Fig. 87) with shallow
pits in the middle, which are not glandular openings. The bases of the large
warts are separated from one another by several smaller dark warts.
Besides, the whole notum bears a great number of scattered pigment-free
wartlets, about 20 ^ in diameter. A single row of about 12 light apertures
232
( n e ) of big subepidermal glands ( no) on either side lies about 3 mm
above the notal border. The thick white mucus, exuded by the slugs when
disturbed, as the collector observed, is produced by these glands. The
above-mentioned pores are the openings of ducts which come from parcels
of large gland cells lying between the big notal glands and the body cavity.

F i g u r e s 87-91. Hoffm annola hansi, sp . n o v . 87, d o r sa l v ie w o f p r e s e r v e d slu g ;

88, v e n t r a l v ie w o f s a m e ; 89, situ s v is c e r u m ; 90, t r a n s v e r s e s e c tio n o n le v e l
o f h e a r t; 91, r a d u la r te e th .

233
These cells contains granular and gelatinous secretion and open into cel­
lular ductules which unite into wider ducts leading to the pores. The parcels
of these gland cells are embedded in diagonal fibers of the body muscula­
ture and blood spaces. From the main duct a few branches go into the
perinotum, where they drain clusters of small gland cells.
The openings of the body are as in H. lesliei (Stearns); the male
aperture lies in the middle of the forehead hidden by the veil; the opening
of the mantle cavity (anus) behind the posterior end of the foot; the
female opening to the right of the anus; and the pneumostome on a
papilla between anus and notal border.
The body cavity of Hoffmannola is divided into a wider left compart­
ment and a smaller right one by a septum (Fig. 89, se) formed by a fold
of the annular musculature protruding from the body wall. The septum
begins at the hind end of the body cavity and ends in front of the level of
the stomach (fe). On the ventral side it extends farther forward than
dorsally. The peritoneum is colorless, but the pericerebral conjunctive
tissue and that along some blood vessels is pigmented.
The salivary glands ( sa ) are branched; their fundi are fastened to the
nerve ring as in Onchidella (Plate, 1893: 112). The radula (Fig. 91)
comprises about 170 rows, each with about 600 teeth per half-row. The
tricuspidate rhachidian tooth, the shorter innermost lateral tooth, and the
long rectangular bases of the outer lateral teeth, their large, blunt mesocone,
and the small, pointed ectocone agree with H. lesliei (Hoffmann, 1928:
Text-fig. 12). Also the lobes of the intestinal gland, whose size decreases
from the right to the left and to the hind one, and the course (/) of the
intestine (ibid.: Text-Fig. 13) are the same.
The asymmetry of the reno-pulmonary complex ( k ) , much longer on
the left than on the right side, is fundamentally the same as in H. lesliei,
but the organs are much bigger in the present species.
The ovotestis ( o v ) consists of more than 10 follicles, whose efferent
ductules unite into the hermaphrodite duct. Its fine coils, the seminal
vesicle (Fretter, 1943: 648-649), and the caecum, Hoffmann’s vesicula
seminalis (1928: Text-fig. 14, ves. sem.), as well as the following parts,
the pouches of the mucus gland ( mu) , and the prostate in the middle,
the lobulate albumen glands on both sides, are the same as in Onchidella.
The brownish, globular bursa inserts far ectally, the coiled vaginal ap­
pendage, Fretter’s flagellum (1943: 703), is quite short and inserted still
farther ectally.
The male duct (Fig. 92, d ) enters the body cavity beside the right
tentacle. It begins thick, 0.18 mm in diameter, and glandular; in the
preserved specimen this section is green. It loops on the bottom of the
visceral cavity backwards to the fore end of the septum {se), turns for­
wards, becomes thin (80 /x), colorless, with single pigment cells on the
walls, and enters the penis, a straight muscular tube between the male
234
opening and the insertion of the penis retractor (r), which originates deep
between the two muscle layers of the septum. In some cases there is a
pigment spot at the insertion of the retractor. A loose outer layer of
muscles sheathes the central one around the coils of the male duct ( d ),
still 80 fi in diameter, which first descends and then ascends again (Fig.
95). The anterior efferent section (Fig. 94, as) of only 25 diameter
opens into a cuticular tube (ci) produced by a thin epithelium and sur­
rounded by the outer muscle layer. A dilatation with conical glandular
villosities (Hoffmann, 1928: PI. 4, Fig. 16, dr. a.) is not developed. A
loose tissue, rich in nuclei ( nu) , lies between the outer muscle layer
and the inner one containing the male duct. The foremost section, the
male atrium (Fig. 93, a m ), is a muscularized invagination of the skin,
whose noncuticularized epithelium is thrown into longitudinal folds.
The species is named in remembrance of Hans Hoffmann (Jena).

F ig u r e s 92-95. Hoffmannola hansi, sp . n o v . 92, m a le c o p u la to r y organ;

93-95, th r e e s e c tio n s o f s a m e w ith h ig h e r p o w e r .

Discussion .— Steinbeck & Ricketts’ findings on the western coast of the

Gulf of California are included in the “Further Distribution” of the present
species, though Dr. G. Dallas Hanna had classified them (Steinbeck &
Ricketts, 1941: 545) as Onchidium lesliei Stearns, 1892. This species
from Albemarle Island, Galapagos Islands (Stearns, 1893b: 383), can be
235
separated from the present one only by dissection. Stearns’ species is well
known since Hoffmann (1928: 56-67) examined it and found it different
from Onchidium and Onchidella, creating the genus Watsoniella for it.
Hoffmannola Strand (1932: 193) is a nomenclatural replacement of
Watsoniella Hoffmann.
Specific characters of H. lesliei which separate it from H. hansi are: the
notal glands are much smaller, much more numerous, and open on different
levels, the reno-pulmonary complex is smaller, the radula has 120 rows
and 360 teeth per half-row, the penial retractor originates under the nerve
ring, and the penial atrium contains glandular villosities. The glands
opening with ventral pores occur in both species. Hoffmann saw the pores,
but could not define the glands in his very old material. They are compar­
able to Wissel’s middle-sized multicellular mantle glands (1898: 596, 497).
In his figure (PI. 34, Fig. 9) and in sections of Onchidella evelinae Marcus
& Burch, 1965, they are globular groups of cells with simple, not ramified
ducts. Wissel noted that their occurrence varies individually; in our material
of O. marginata (Gould 1852) they were missing (Marcus 1959: 17 ff.).
Steinbeck & Ricketts (loc. cit.) called the Onchidiacea “common in
Arctic and Antarctic waters.” The contrary is correct. The group occurs on
the coasts of all latitudes except the Arctic and Antarctic zones. The
latitudes of 60° N and 55° S are given as limits of the range in Hoffmann’s
survey (1929: 253).
ZOOGEOGRAPHIC REMARKS

The lists of the macro-invertebrate assemblages and environments in

Parker’s monograph (1964) show that the shell-less opisthobranchs of
this sea are largely unknown. Parker’s fundamental work which is based
upon six years of studies contains abundant data and explanations. With a
small list of species, as the present one, collected in a restricted area of the
Gulf, we cannot supplement Parker’s work with notes on the opistho­
branchs, e.g., we cannot evince a faunistic relation between the northern
extremity of the Gulf and its merely warm-temperate (not tropical) climate
(Parker, 1964: 39-41, 46). The subregions of the Panamic Province
(Ekman, 1953: 45) might have different limits for the various groups of
marine animals, so that Berry’s (1959) inclusion of the northern part of
the Gulf in the Panamic Province may be definitive for the molluscs.
We can make only an attempt to explain the geographic zoology for
most of our slugs. Even the labels “Panamic” or “Californian” are
provisional, because the level of systematic knowledge regarding the opis­
thobranchs of the coasts adjacent to the Gulf (that of Sinaloa and of the
oceanic side of Baja California) is quite different. The knowledge of the
latter is much more advanced, especially with regard to its warm-temperate
continuation, southern California from San Diego to Point Conception.
For a number of species found in the Gulf of California it is difficult to
236
indicate foreign relationships: lnuda luarna, Tayuva ketos, Coryphella
cynara, and Flabellina telja. Also the affinities of some additional species
are vague: Casella sedna which belongs to an ill-defined Indo-West Pacific
genus, N em brotha eliora and Acanthodoris pina, both comparable with
Indo-West Pacific species of their well-defined genera, and D oris pickensi,
whose geographically nearest and definitely congeneric species is the mor­
phologically distant Atlantic D. verrucosa known from the west coast of
Florida (Bergh, 1894: 161).
For all other species of the present collection, some geographic com­
ments are possible. Of the two new subspecies, Chrom odoris banksi sonora
and Berthellina engeli ilisima, the former is near to Chr. banksi banksi from
the west coast of the Gulf, while the latter testifies for the Central American
Sea, still existent in the lower Pliocene (Ekman, 1953: 37). The latter
type of distribution holds also for Hermaea hillae, near to H. coirala, for
Tridachiella diom edea, allied to Tridachia crispata, for Pleurobranchus
areolatus and Cadlina evelinae, for the well-defined genus Taringa, for
Polycera gnupa, related to the tropical West Atlantic P. hummi and P.
aurisula, D endrodoris krebsii, and Tritonia pickensi, similar to T. bayeri
from Florida.
On the sandy bottom of the intermediate shelf (27-65 m) and the clayey
bottom of the outer shelf (66-120 m ), a very great number of shelled
Mollusca live as almost indistinguishable “twin-species” in the Gulf of
California and the Gulf of Mexico (Parker, 1964: 117, 120-121).
Concerning the intertidal opisthobranchs of the Gulf of California the
comparison should consider rather the Caribbean Sea and its extension to
the South than the monotonous sandy and muddy beaches of Texas and
Louisiana.
Tridachiella diom edea, Pleurobranchus areolatus, and a subspecies of
Taringa aivica, similar to that of the Gulf, occur also on the Pacific coast
of Panama, and Dendrodoris krebsii on the coast of Jalisco, Mexico.
Due to our scant knowledge of the East Pacific tropical opisthobranchs,
the list of purely East Pacific warm-water species in the Gulf is small;
Bulla gouldiana, the genus Conualevia, Chrom odoris norrisi, Doriopsilla
janaina and the closely related D . rowena, Spur ilia chromosoma, Onchidella
hildae and O. binneyi, two species similar to one another, and Hoffmannola
hansi, a relative of H. lesliei from the Galapagos Islands can be mentioned;
the range of B. gouldiana and Sp. chrom osom a extends to warm-temperate
southern California.
Of the species occurring in the Gulf and on the Pacific coast of North
America, three groups may be distinguished: (1) those restricted to the
coast south of Point Conception; (2) those occurring also farther north,
but not beyond the 10° C isotherm for March, approximately at Cape
Blanco; and (3) those which live also farther north. To the first group
belong the above-mentioned Bulla gouldiana, the genus Conualevia,
237
Chrom odoris norrisi, and Spurilla chromosoma, as well as the testimonies
of the Tertiary Central American Sea Cadlina evelinae and D endrodoris
krebsii. In the second group we have Navanax inermis, Aplysia calif ornica,
H ypselodoris californiensis, Polycera atra, Doriopsilla albopunctata, D oto
amyra, Dirona picta, and Coryphella iodinea. The third group includes
Ham inoea virescens, Diaulula sandiegensis, and Hermissenda crassicornis.
It would be interesting to know whether the species of groups 2 and 3
reproduce also in the true tropical environments of the Gulf (Parker,
1964: 41-42), the protected waters of La Paz and Cape San Lucas,
similar to a tropical Pacific island.
D iscodoris m avis from the Gulf is intermediate between the eurythermic
D. heathi from the North American west coast and D. pusae, a tropical
species from the West Atlantic. Dendronotus nanus and D oto lancei have
relationships to North American Pacific species.
Besides the mentioned A plysia californica the collection contains the
Anaspidean A . parvula, already known from the Gulf, and two species not
yet recorded from the North American Pacific coast, A . juliana and
Stylocheilus longicauda. The range of A . parvula includes as East Pacific
localities the Juan Fernandez Islands and the Gulf, so that it may have
come from the West Pacific or from the Caribbean by the Central Ameri­
can way. A . juliana has been recorded from Peru and the Galapagos
Islands, and so its occurrence in the Gulf is that of an East Pacific warm-
water species such as Onchidella hildae and others. The observation of the
stenothermic Stylocheilus longicauda at Puerto Penasco during July and
August, among the fronds of Sargassum, is significant. The species has been
recorded from floating algae and pumice (Engel, 1936: 67), and may have
reached west America from the west, e.g., from the Ryukyu or the
Sandwich Islands, where it occurs.
Regarding the three prosobranches of the collection, Titiscania limacina
was recently found on the Pacific coast of Panama. The east Pacific
occurrences of this Indo-West Pacific species cannot be explained. The
two forms of Lamellaria perspicua are eurythermic. The antiboreal temper­
ate water of the Magellanic region is no barrier for them.

R esu m o

O trabalho contem 47 especies e subespecies coletadas por Dr. Peter

E. Pickens da Universidade da Arizona (Tucson) no litoral superior do
nordeste do Golfo da California. Das 20 formas novas lnuda luarna, sp. n.,
e Tayuva ketos, sp. n., representam novos generos das Dorididae. Varias
das especies recentemente encontradas por Dr. Frederick M. Bayer na
costa pacifica do Panama ocorrem tambem no Golfo, entre estas Titiscania
limacina dos Neritacea, do Indo-Pacifico ocidental. Cerca de dois tergos
da colegao presente sao opistobranquios da agua quente; re la te s com a
fauna tropical do Atlantico ocidental sao evidentes.
238
 ZUSAMMENFASSUNG

Die Arbeit enthalt 47 Arten und Unterarten, die Dr. Peter E. Pickens
von der Universitat von Arizona (Tucson) im oberen Litoral der nordost-
lichen Kuste des Golfes von Kalifornien gesammelt hat. Von den 20 neuen
Formen stellen lnuda luarna, sp. n., und Tayuva ketos, sp. n., neue Gat-
tungen der Dorididen dar. Mehrere der Arten, die Dr. Frederick M. Bayer
vor Kurzem an der pazifischen Kuste von Panama gefunden hat, kommen
auch im Golf vor, unter ihnen die zu den Neritacea gehorige, im ubrigen
indowestpazifische Titiscania limacina. Gut zwei Drittel der festgestellten
Opisthobranchier sind Warmwasserformen. Beziehungen zur westatlan-
tischen Warmwasserfauna sind deutlich.

L is t of T ypes

The type specimens of the new species described in this paper have been
assigned catalogue numbers in the U.S. National Museum. As it was impossible
to insert these in the text, they are given below in the order in which the
species appear in the text.
7. Hermaea hillae, 678400 27. N em brotha eliora, 678410
13. Berthellina engeli ilisima, 678401 29. Polycera gnupa, 678411
15. Conualevia mizuna, 678402 30. Acanthodoris pina, 678412
18. Chromodoris banksi sonora, 34. Doriopsilla rowena, 678413
678403 35. Tritonia (Candiella) pickensi,
20. Casella sedna, 678404 678414
21. lnuda luarna , 678405 36. Dendronotus nanus, 678415
22. Doris pickensi , 678406 38. D oto lancei, 678416
23. Discodoris mavis, 678407 41. Coryphella cynara, 678417
25. Taringa aivica timia, 678408 42. Flabellina telja, 678418
26. Tayuva ketos , 678409 47. Hoffmannola hansi, 678419

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248
 INDEX*
To Taxa Described or Mentioned

Acanthodoris falklandica, 202 alba, Aeolidiella, 116

globosa, 203 Conualevia, 186
lutea, 202 Discodoris, 80
metulifera, 203 Glossodoridiformia, 186
pilosa, 202 albolineata, Dirona, 217
pilosa purpurea, 202 albopunctata, Doriopsilla, 98, 99, 189, 204,
pina, 201, 237 238
rhodoceras, 202 amarillia, Berthellina, 163
acriba, Hypselodoris, 60, 123 amoena, Chromodoris, 173
Acteon, 14 amyra, Doto, 214, 238
adellae, Aglaja, 20 angustipes, Platydoris, 93, 124
aegialia, Glossodoris, 61 Anisodoris, 68, 69
aenigmaticus, Navanax, 19 flindersi, 70
Aeolidiella, 116 marmorata, 69, 124
alba, 116 nobilis, 71
glauca, 117 prea, 70, 124
hulli, 116 punctuolata, 69
indica, 117 timorensis, 70
japonica, 118 worki, 66, 124
lurana, 115, 125 annuligera, Samla, 226
macleayi, 116 antillarum, Bulla, 22
multicolor, 117, 118 antillarum, Haminoea, 22, 124
occidentalis, 117 Oxynoe, 25, 123
saldanhensis, 117 antillensis, Aphelodoris, 92, 123
takanosimensis, 117 Cyerce, 37, 123
affinis, Flabellina, 224, 225, 226 Aphelodoris antillensis, 92, 123
agassizii, Chromodoris, 56, 176, 178 Aporodoris, 91
agassizii, Pleurobranchaea, 49, 123 millegrana, 88, 91
Aglaja adellae, 20 risbeci, 91
bakeri, 21 rubra, 91
capensis, 20 souriei, 91
ceylonica, 19 Aplysia (Neaplysia) californica, 155, 238
cyanea, 19 (Aplysia) cedrosensis, 158
depicta, 20 (Varria) dactylomela, 38, 124
diomedea. 20 depilans, 158
dubia, 20 (Aplysia) dura, 158
gemmata, 20 (Aplysia) juliana, 155, 238
maderensis, 20 nigra, 159
minuta, 20 (Pruvotaplysia) parvula, 154, 238
nana, 21 (Aplysia) reticulopoda, 158
ocelligera, 20 vaccaria, 158
orbignyana, 20 Archidoris, 68, 186
punctilucens, 20 stellifera, 69
purpurea, 20 tuberculata, 68
pusa, 18, 124 areolata, Doriopsilla, 98
seurati, 20 areolatus, Pleurobranchus (Pleurobranchus),
tricolorata, 19, 20 44, 123, 163, 237
aguayoi, Oxynoe, 25 argo, Platydoris, 94
aivica, Taringa, 89, 124 armadilla, Onchidella, 122
aivica aivica, Taringa, 191 armata, Taringa, 88, 91
aivica timia, Taringa, 182, 184, 189, 204 Armina miilleri, 108
alabastrina, Doris (Asteronotus), 186 semperi, 108
alabe, Polycera, 198 atlanticus, Glaucus, 119, 124

*Names in italic type are currently rejected for nomenclatural or taxonomic reasons; names
in roman type are in current use. Responsibility for this index rests with the editors,
not with the authors of the work.

249
atlanticus, Pleurobranchus (Susania), 44, 163 limbaughi, 169
atra, Polycera, 196, 198, 238 marginata, 170
atromarginata, Casella, 181 rumia, 51, 124, 169
atropos, Dendrodoris, 96, 203 scabriuscula, 51, 170
Doridopsis, 203 sparsa, 169
Doriopsis, 95 Cadlinella, 166
Audura, 192 calcarata, Bornella, 105, 124
aurantiaca, Dirona, 217 californica, Aplysia (Neaplysia), 155, 238
aurila, Discodoris, 80, 85 Coryphella, 226
aurisula, Polycera. 198, 237 californiensis, Hypselodoris, 59, 176, 238
Austraeolis, 114 Candiella, 104, 209
catina, 112, 124 plebeia, 209
fucia, 114 capensis, Aglaja, 20
ornata, 114 Nembrotha, 196
Austrodoris, 66, 74, 186 Pleurobranchaea, 47
nivium, 186 caramella, Doto, 106
Awuka, 166 carpenteri, Onchidella, 227, 229, 230
Onchidium, 227, 229, 230
Casella atromarginata, 181
bakeri, Aglaja, 21 cincta, 181
banksi banksi, Chromodoris, 173, 237 foxi, 181
banksi sonora, Chromodoris, 173, 237 rubra, 181
Baptodoris, 91 rufomarginata, 181
baratariae, Corambella, 94 sedna, 178, 237
bayeri, Feiimare, 62, 124 catina, Austraeolis, 112, 124
Tritonia (Candiella), 101, 124 cauze, Elysia, 28
berghi, Cadlina, 170 cauze scops, Elysia, 28, 123
Berghia, 117 cavernae, Discodoris, 80
bermudense, Umbraculum, 42, 123 cedrosensis, Aplysia (Aplysia), 158
Berthella, 162, 165 celtica, Onchidella, 121, 230
patagonica, 43 ceylonica, Aglaja, 19
plumula, 162 Chelidonura, 151
tupala, 43, 124 evelinae, 18
Berthellina, 162 hirundinina, 21, 124
amarillia, 163 chilluna, Polycera, 198
circularis, 163 Chromodoris, 56, 124
edwardsi, 162 agassizii, 56, 176, 178
engeli ilisima, 160, 237 amoena, 173
quadridens, 43, 123, 163 banksi banksi, 173, 237
Berthellinops, 162 banksi sonora, 173, 237
bicolor, Doris, 66 dalli, 176
binneyi, Onchidella, 227, 231, 237 macfarlandi, 178
bonairei, Petalifera, 39 neona, 51, 124
borealis, Onchidella, 2.29, 230 norrisi, 170, 237, 238
Boreodoris, 166 nyalya, 53, 58, 124
Bornella, 208 punctilucens, 176
calcarata, 105, 124 roseo-picta, 61
Bouvieria, 165 tura, 55
bovena, Doris, 66 universitatis, 176
branneri, Discodoris, 75, 78, 80 zebra, 60
braziliana, Spurilla, 118 chromosoma, Spurilla, 227, 237, 238
brevicauda, Phidiana, 109, 110 cincta, Casella, 181
Bulla antillarum, 22 cinerea, Doto, 216
gouldiana, 148, 237 circularis, Berthellina, 163
Bullaea guamense, 15 citrinus, Stylocheilus, 40, 124
clandestina, Thordisa, 91
Cadlina, 169 (key), 181 coelestis, Hypselodoris, 62
berghi, 170 coirala, Hermaea, 152, 237
evelinae, 123, 168, 170, 237, 238 columbiana, Doto, 216
flavomaculata, 169 Conualevia, 166, 237
juvenca, 169 alba, 186
laevis, 184 mizuna, 167

250
cooperi, Coryphella, 221, 222, 223 branneri, 75, 78, 80
Corambella baratariae, 94 cavernae, 80
Coryphella, 220, 226 edwardsii, 80
californica, 226 erubescens, 80
cooperi, 221, 222, 223 evelinae, 75, 78, 80, 124
cynara, 220, 222, 237 fragilis, 75
dushia, 222 heathi, 80, 82, 87, 188, 189, 288
falklandica, 221 hedgpethi, 75, 78, 80, 123
fisheri, 226 indecora, 80
fusca, 222, 226 maculosa, 80
iodinea, 218, 238 mavis, 187, 238
lineata, 226 mortenseni, 76, 80, 82
ornata, 224, 226 muta, 80
pellucida, 224 notha, 80, 87
piunca, 222, 226 notiperda, 75
pricei, 223 phoca, 78, 124
rufibranchialis, 219 purcina, 80, 81, 123
trilineata, 222, 226 pusae, 80, 82, 124, 189, 238
trophina, 226 rubens, 80
crassicornis, Hermissenda, 226, 238 spetteda, 75, 76
Cratena, 115 tristis, 80
kaoruae, 114, 124 voniheringi, 80, 87
crispa, Elysia, 34 diuda, Thordisa, 91
crispata, Tridachia, 33, 35, 37, 123, 124, 237 divae, Doto, 108
crossei, Pleurobranchus, 44 Nembrotha, 196
cristallina, Cyerce, 38 Dolabrifera dolabrifera, 38, 124
cucullata, Marionia, 104, 125
dolabrifera, Dolabrifera, 38, 124
Cyerce antillensis, 37, 123
cristallina, 38 Doridella obscura, 94
jheringi, 38 Dorididigitata maculata, 186
cynara, Coryphella, 220, 222, 237 Doridopsis atropos, 203
Krebsii var. pallida, 203
dalli, Chromodoris, 176 Doriopsilla, 99
dactylomela, Aplysia (Varria), 38, 124
albopunctata, 98, 99, 189, 204, 238
Dendrodoris, 99 areolata, 98
atropos, 203 fulva, 204
fulva, 204
janaina, 96, 124, 205, 207, 237
krebsii, 95, 123, 124, 203, 237, 238 leia, 207
subpellucida, 96
pharpa, 206, 207
Dendronotus, 208
rowena, 205, 237
frondosus, 212
Doriopsis atropos, 95
gracilis, 212
fulva, 99
iris, 210, 212, 213
krebsii var. pallida, 95
nanus, 210, 238
reticulata, 99, 204
robustus, 212
Doris, 186
depicta, Aglaja, 20
(Asteronotus) alabastrina, 186
depilans, Aplysia, 158 bicolor, 66
depressa, Phyllaplysia, 40
bovena, 66
Diaulula sandiegensis, 189, 238
echinata, 186
Dictyodoris tesselata, 191 odonoghuei, 186
diegoensis, Lamellaria, 147 pickensi, 184, 189, 237
digueti, Lamellaria, 166 verrucosa, 186, 237
Pleurobranchus, 165 Doto, 215, 216
diomedea, Aglaja, 20 amyra, 214, 238
Tridachiella, 34, 124, 154, 237 caramella, 106
Dirona albolineata, 217 cinerea, 216
aurantiaca, 217 columbiana, 216
picta, 216, 238 divae, 108
disa, Taringa telopia, 87, 124 lancei, 214, 216, 238
Discodoris, 69, 74, 76 (key), 124 pita, 106
alba, 80 uva, 108
aurila, 80, 85 varians, 214
dubia, Aglaja, 20 Noumea, 55
Thordisa, 91 flavomaculata, Cadlina, 169
duis, Elysia, 31, 124 flindersi, Anisodoris, 70
dura, Aplysia (Aplysia), 158
floridana, Onchidella, 119, 123
dushia, Coryphella, 222
Duvaucelia, 104, 209 foxi, Caselia, 181
insulae, 104 frangoisi, Risbecia, 62
lineata, 104 fragilis, Discodoris, 75
frondosus, Dendronotus, 212
fucia, Austraeolis, 114
echinata, Doris, 186 fulva, Dendrodoris, 204
edenticulata, Hypselodoris, 56, 123, 178 Doriopsilla, 204
edwardsii, Berthellina, 162 Doriopsis, 99
Discodoris, 80 fusca, Coryphella, 222, 226
elegans, Harminoea, 23, 24, 123 fuscovittatus, Stiliger, 154
Tritoniopsis, 103
eliora, Nembrotha, 194, 237 Galacera, 197
Elysia cauze, 124
gardineri, Susania, 44
cauze scops, 28, 123
gemmata, Aglaja, 20
crisp a, 34
glauca, Aeolidiella, 117
duis, 31, 124
Glaucus atlanticus, 119, 124
evelinae, 27, 123
globosa, Acanthodoris, 203
flava, 33
gracilis, 32 Glossodoridiformia, 166, 186
alba, 186
livida, 32
papillosa, 27, 123 Glossodoris aegialia, 61
papillosa, 29 gnupa, Polycera, 198, 237
picta, 32 Godiva rubrolineata, 112, 124, 125
pruvotfolae, 34 gouldiana, Bulla, 148, 237
sp., 32 gracilis, Dendronotus, 212
Elysia, 32
splendens, 32
tuca, 29, 123 Hypselodoris, 60
(Elysiopterus) verrilli, 34 gratiosa, Nembrotha, 196
greeleyi, Peltodoris, 66, 72, 124
engeli, Phyllaplysia, 40, 123
Greilada, 197
engeli ilisima, Berthellina, 160, 237
guadaloupensis, Haminoea antillarum, 22
enteromorphae, Phyllobranchopsis, 153 guamensis, Micromelo, 15
Eolidina, 116 Guyonia, 166
erubescens, Discodoris, 80
Erythrodoris, 166
Etidoris, 91 Haminoea, 21
ladislavii, 91 anlillarum, 22, 124
evelinae, Cadlina, 123, 168, 170, 237, 238 antillarum guadaloupensis, 22
Chelidonura, 18 elegans, 23, 24, 123
Discodoris, 75, 78, 80, 124 strongi, 149
Elysia, 27, 123 virescens, 149, 238
Onchidella, 236 virescens rosacea, 149
exigua, Phidiana, 110, 111 hamva, Pleurobranchaea hedgpethi, 48, 124
exsulans, Tritonia, 123 hansi, Hoffmannola, 232, 237
heathi, Discodoris, 80, 82, 87, 188, 189, 238
falklandica, Acanthodoris, 202 hedgpethi, Discodoris, 75, 78, 80, 123
Coryphella, 221 Polycera, 198, 199, 200
Felimare, 62, 166 hedgpethi hamva, Pleurobranchaea, 48, 124
bayeri, 62, 124 hedgpethi hedgpethi, Pleurobranchaea, 48, 49
festiva, Tritonia (Candiella), 104, 209 Hermaea coirala, 152, 237
Fiona pinnata, 109, 124 hillae, 151, 237
fisheri, Coryphella, 226 oliviae, 154
Flabellina, 219, 220, 224, 226 ornata, 154
affinis, 224, 225, 226 vancouverensis, 153
iodinea, 226 Hermaeina, 153
ornata, 225 oliviae, 154
telja, 223, 237 smithi, 153
Flabellinopsis iodinea, 218 Hermissenda crassicornis, 226, 238
flava, Elysia, 33 herthae, Polycera, 198

252
hilaris variety, Thordisa, 91 digue ti, 166
hildae, Onchidella, 230, 237, 238 orbiculata, 147
hillae, Hermaea, 151, 237 perspicua, 11, 124, 238
hirundinina, Chelidonura, 21, 124 perspicua mopsicolor, 11, 148
Hoffmannola, 234 perspicua perspicua, 145
rhombica, 147
hansi, 232, 237
stearnsii, 147
lesliei, 234, 237
lancei, Doto, 215, 216, 238
hombergi, Tritonia, 209
leia, Doriopsilla, 207
Homoiodoris, 66
hulli, Aeolidiella, 116 lesliei, Hoffmannola, 234, 237
hummelincki, Peltodoris, 74 limacina, Titiscania, 7, 124, 125, 145, 238
hummi, Polycera, 198, 200, 237 limbaughi, Cadlina, 169
Hydatina physis, 16 lineata, Coryphella, 226
quoy ana, 16 Duvaucelia, 104
lineolata, Nembrotha, 196
(Aplustrum) velum, 14
Lissodoris, 166
vesicaria, 16, 123, 124
livida, Elysia, 32
Hypselodoris, 56
Lobiger nevilli, 25
acriba, 60, 123 picta, 25
californiensis, 59, 176, 238 pilsbryi, 27
coelestis, 62
serradifalci, 27
edenticulata, 56, 123, 178
souverbiei, 25, 123, 124
gracilis, 60 viridis, 25
sycilla, 59, 178
longicauda, Stylocheilus, 41, 42, 159, 238
luarna, Inuda, 181, 182, 189, 237
ilisima, Berthellina engeli, 160, 237
ilo, Siraius, 66 lurana, Aeolidiella, 115, 125
inca, Phidiana, 110, 111 lutea, Acanthodoris, 202
indecora, Discodoris, 80 lynceus, Phidiana, 109, 110, 111, 123, 124
Tritonia, 103
indica, Aeolidiella, 117 macfarlandi, Chromodoris, 178
inermis, Navanax, 19, 149, 238 macleayi, Aeolidiella, 116
indolens, Onchidella, 121, 122, 230 maculata, Dorididigitata, 186
insulae, Duvaucelia, 104 maculosa, Discodoris, 80
insulae, Tritonia, 104 maderensis, Aglaja, 20
Inuda, 166, 181 Madrella, 217
luarna, 181, 182, 189, 237 marginata, Cadlina, 170
Inudinae, 182 Onchidella, 236
iodinea, Coryphella, 218, 238 Marionia cucullata, 104, 125
Flabellina, 226 occidentalis, 104
Flabellinopsis, 218 marmorata, Anisodoris, 69, 124
iris, Dendronotus, 210, 212, 213 marplatensis, Polycera, 197
Marsenina rhombica, 147
janaina, Doriopsilla, 96, 124, 205, 207, 237 mavis, Discodoris, 187, 238
japonica, Aeolidiella, 118 meckelii, Pleurobranchaea, 45, 47, 48
jheringi, Cyerce, 38 membranaceus, Pleurobranchus, 165
Jorunna, 166 metulifera, Acanthodoris, 203
juliana, Aplysia (Aplysia), 155, 238 Micromelo guamensis, 15
juvenca, Cadlina, 169 undata, 12, 123
Miesea, 215
kaoruae, Cratena, 114, 124 millegrana, Aporodoris, 88, 91
Kentrodoris, 166 minuta, Aglaja, 20
ketos, Tayuva, 192, 237 mizuna, Conualevia, 167
krebsii, Dendrodoris, 95, 123, 124, 203, 237, mopsicolor, Lamellaria perspicua, 11, 148
238 morosa, Pleurobranchaea, 48
krebsii var. pallida, Doriopsis, 95 Nembrotha, 195
Doridopsis, 203 mortenseni, Discodoris, 76, 80, 82
kyolis, Siraius, 64, 124 mulleri, Armina, 108
multicolor, Aeolidiella, 117, 118
ladislavii, Etidoris, 91 muta, Discodoris, 80
laevis, Cadlina, 184 Myrella, 101
lafontii, Phyllaplysia, 40
Lamellaria diegoensis, 147 nana, Aglaja, 21

253
nanus, Dendronotus, 210, 238
Navanax, 151
aenigmaticus, 19
inermis, 19, 149, 238
neapolitana, Spurilla, 118, 124
Nembrotha capensis, 196
divae, 196
eliora, 194, 237
gratiosa, 196
lineolata, 196
morosa, 195
nigerrima, 196
Neodoris, 66, 166
neona, Chromodoris, 51, 124
nevilli, Lobiger, 25
nigerrima, Nembrotha, 196
nigra, Aplysia, 159
nivium, Austrodoris, 186
nobilis, Anisodoris, 71
norrisi, Chromodoris, 170, 237, 238
notha, Discodoris, 80, 87
notiperda, Discodoris, 75
Noumea flava, 55
noumeae, Peltodoris, 72
nyalya, Chromodoris, 53, 58, 124
obscura, Doridella, 94
occidentalis, Aeolidiella, 117
Marionia, 104
Pleurobranchaea, 45, 123
ocelligera, Aglaja, 20
odhneri, Polycera, 198
Risbecia, 62
odonoghuei, Doris, 186
oliviae, Hermaea, 154
Hermaeina, 154
Onchidella, 121, 228, 231, 234
armadilla, 122
binneyi, 227, 231, 237
borealis, 229, 230
carpenteri, 227, 229, 230
celtica, 121, 230
evelinae, 236
boridana, 119, 123
hildae, 230, 237, 238
indolens, 121, 122, 230
marginata, 236
steindachneri, 121, 122, 123
Onchidium, 121
carpenteri, 227, 229, 230
trans-Atlanticum, 122
orbiculata, Lamellaria, 147
orbignyana, Aglaja, 20
ornata, Austraeolis, 114
Coryphella, 224 226
Flabellina, 225
Hermaea, 154
Otinodoris, 166
Oxynoe aguayoi, 25
antillarum, 25, 123
254
pallida, Casella atromarginata, 181
Doriopsis krebsii var., 95
palmeri, Tritonia, 104, 210
papilligera, Phyllidiopsis, 99, 123
papillosa, Elysia, 27, 123
papillosa, Elysia, 29
Paratritonia, 103
parvula, Aplysia (Pruvotaplysia), 154, 238
patagonica, Berthella, 43
Phidiana, 109, 110, 112
pellucida, Coryphella, 224
Peltodoris, 69, 74
greeleyi, 66, 72, 124
hummelincki, 74
noumeae, 72
perspicua mopsicolor, Lamellaria, 148, 238
perspicua perspicua, Lamellaria, 11, 124, 145
Petalifera bonairei, 39
ramosa, 39, 124
pharpa, Doriopsilla, 206, 207
Phialodoris, 192
Phidiana, 109, 112
brevicauda, 109, 110
exigua, 110, 111
inca, 110, 111
lynceus, 109, 110, 111, 123, 124
patagonica, 109, 110, 112
pugnax, 110
selencae, 109
selenkai, 109, 110
phoca, Discodoris, 78, 124
Phyllaplysia depressa, 40
engeli, 40, 123
lafontii, 40
papilligera, 99, 123
Phyllidiopsis (Phyllidiopsis)
papilligera, 99, 123
Phyllobranchopsis enteromorphae, 153
physis, Hydatina, 16
pickensi, Doris, 184, 189, 237
Tritonia (Candiella), 207, 237
picta, Dirona, 216, 238
Elysia, 32
Lobiger, 25
pilosa, Acanthodoris, 202
pilosa purpurea, Acanthodoris, 202
pilsbryi, Lobiger, 27
pina, Acanthodoris, 201, 237
pinnata, Fiona, 109, 124
pita, Doto, 106
piunca, Coryphella, 222, 226
Platydoris angustipes, 93, 124
argo, 94
rubra, 93
plebeia, Candiella, 209
Pleurobranchaea agassizii, 49, 123
capensis, 47
hedgpethi hamva, 48, 124
meckelii, 45, 47, 48
morosa, 48
occidentalis, 45, 123
Pleurobranchus (Pleurobranchus) areolatus,
 44, 123, 163, 237 rufibranchialis, Coryphella, 219
(Susania) atlanticus, 44, 163 rufomarginata, Casella, 181
crossei, 44 rumia, Cadlina, 51, 124, 169
digueti, 165
membranaceus, 165 sagamiana, Tambja, 196
plumula, 162 saldanhensis, Aeolidiella, 117
tuberculatus, 165 Samla annuligera, 226
plicatulum, Umbraculum, 42, 123 sandiegensis, Diaulula, 189, 238
plumula, Pleurobranchus, 162 sargassicola, Spurilla, 118
Berthella, 162 scabriuscula, Cadlina, 51, 170
scops, Elysia cauze, 28, 123
Polycera, 197 (key)
Sebadoris, 166
alabe, 198
sedna, Casella, 178, 237
atra, 196, 198, 238
selencae, Phidiana, 109
aurisula, 198, 237
selenkai, Phidiana, 109, 110
chilluna, 198
semperi, Armina, 108
gnupa, 198, 237
serradifalci, Lobiger, 27
hedgpethi, 198, 199, 200
seurati, Aglaja, 20
herthae, 198
Siraius, 66, 166
hummi, 198, 200, 237
ilo, 66
marplatensis, 197
kyolis, 64, 124
odhneri, 198
smithi, Hermaeina, 153
priva, 198
zosterae, 198 sonora, Chromodoris banksi, 173, 237
souriei, Aporodoris, 91
prea, Anisodoris, 70, 124
souriei, Thordisa, 91
pricei, Coryphella, 223
priva, Polycera, 198 souverbiei, Lobiger, 25, 123, 124
sparsa, Cadlina, 169
pruvotfolae, Elysia, 34
spetteda, Discodoris, 75, 76
pugnax, Phidiana, 110
splendens, Elysia, 32
punctilucens, Aglaja, 20
Spongiodoris, 167
Chromodoris, 176
Spurilla, 117
punctuolata, Anisodoris, 69
braziliana, 118
purcina, Discodoris, 80, 81, 123
chromosoma, 227, 237, 238
purpurea, Aglaja, 20
neapolitana, 118, 124
pusa, Aglaja, 18, 124
sargassicola, 118
pusae, Discodoris, 80, 82, 124, 189, 238
stearnsii, Lamellaria, 147
steindachneri, Onchidella, 121, 122, 123
quadridens, Berthellina, 43, 123, 163
stelliferca, Archidoris, 69
quoyana, Hydatina, 16
Stiliger fuscovittatus, 154
vancouverensis, 153
ramosa, Petalifera, 39, 124 strongi, Haminoea, 149
reticulata, Doriopsis, 99, 204 Stylocheilus citrinus, 40, 124
reticulata, Risbecia, 62 longicauda, 41, 42, 159, 238
Tritonia, 102 subpellucida, Dendrodoris, 96
reticulopoda, Aplysia (Aplysia), 158 Susania gardineri, 44
rhodoceras, Acanthodoris, 202 sycilla, Hypselodoris, 59, 178
rhombica, Marsenina, 147
Lamellaria, 147
risbeci, Aporodoris, 91 takanosimensis, Aeolidiella, 117
Risbecia, 62, 166 Tambja sagamiana, 196
frangoisi, 62 Taringa, 91, 167, 237
odhneri, 62 aivica, 89, 124
reticulata, 62 aivica timia, 182, 184, 189, 204
versicolor, 62 armata, 88, 91
robustus, Dendronotus, 212 telopia disa, 87, 124
roseo-picta, Chromodoris, 61 Tayuva, 167, 191
Rosodoris, 166 ketos, 192, 237
rowena, Doriopsilla, 205, 237 telja, Flabellina, 223, 237
rubens, Discodoris, 80 telopia disa, Taringa, 87, 124
rubra, Aporodoris, 91 telopia telopia, Taringa, 87, 88
Platydoris, 93 tesselata, Dictyodoris, 191
rubra, Casella, 181 Thordisa, 91
rubrolineata, Godiva, 112, 124, 125 clandestina, 91

255
 diuda, 91 tuca, Elysia, 29, 123
dubia, 91 tupala, Berthella, 43, 124
hilaris variety, 91 tura, Chromodoris, 55
souriei, 91 Tylodina, 21
Thorunna, 167
timia, Taringa aivica, 182, 184, 189 Umbraculum, 21
timorensis, Anisodoris, 70 bermudense, 42, 123
Titiscania limacina, 7, 124, 125, 145, 238 plicatulum, 42, 123
trans-Atlanticum, Onchidium, 122 undata, Micromelo, 12, 123
tricolorata, Aglaja, 19, 20 universitatis, Chromodoris, 176
Tridachia crispata, 33, 35, 37, 123, 124, 237 uva, Doto, 108
Tridachiella diomedea, 34, 124, 154, 237
trilineata, Coryphella, 222, 226 vaccaria, Aplysia, 158
tristis, Discodoris, 80 vancouverensis, Hermaea, 153
Tritonia, 101, 104, 209 vancouverensis, Stiliger, 153
(Candiella) bayeri, 101, 124 varians, Doto, 214
exsulans, 123 velum, Hydatina (Aplustrum), 14
(Candiella) festiva, 104, 209 verrilli, Elysia (Elysiopterus), 34
hombergi, 209 verrucosa, Doris, 186, 237
indecora, 103 versicolor, Risbecia, 62
insulae, 104 vesicaria, Hydatina, 16, 123, 124
palmeri, 104, 210 villafranca, Tritonia, 104
(Candiella) pickensi, 207, 237 virescens, Haminoea, 149, 238
reticulata, 102 virescens rosacea, Haminoea, 149
villafranca, 104 viridis, Lobiger, 25
(Tritonidoxa) wellsi, 99, 125 voniheringi, Discodoris, 80, 87
Tritonidoxa, 101, 209
Tritoniopsis, 103 Watsoniella, 236
elegans, 103 wellsi, Tritonia (Tritonidoxa), 99, 125
Tochuina, 209 worki, Anisodoris, 66, 124
trophina, Coryphella, 226
tuberculata, Archidoris, 68 zebra, Chromodoris, 60
tuberculatus, Pleurobranchus, 165 zosterae, Polycera, 198

256