CONTRIBUTIONS IN SCIENCE, 3

CUBOZOAN AND SCYPHOZOAN

OF THE CAROUNL
SOUTHEASTE

DALE R^CALDER
Published by the Royal Ontario Museum with the generous
support of the Louise Hawley Stone Charitable Trust
Frontispiece: A. Chiropsalmus quadrumanus. B. Periphylla periphylla. C.Amelia marginalis. D. PhyUorhiza punctata
(photographs A, B, and D by R.J. Larson; C by D. R. Calder).
nrlfl kit CONTRIBUTIONS IN SCIENCE, 3

Dale R. Calder

Cubozoan and Scyphozoan Jellyfishes

OF THE CAROLINIAN BIOGEOGRAPHIC PROVINCE,
SOUTHEASTERN USA

Department of Natural History, Royal Ontario Museum, 100 Queen's Park, Toronto, ON, Canada, M5S 2C6
© 2009 Royal Ontario Museum

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Publication date: September 2009

ISSN 1710-7768

Library and Archives Canada Cataloguing in Publication

Calder, Dale R.
Cubozoan and Scyphozoan jellyfishes of the Carolinian
biogeographic province,southeastern USA / Dale R. Calder.

(ROM contributions in science ; 3)

ISBN 978-0-88854-471-1

1.Jellyfishes—South Atlantic States. I. Royal Ontario

Museum II.Title. III. Series: ROM contributions in
science ; 3

QL377.S4C24 2009 593-5'30975 C2009-904731-4

ROM Science Editorial Committee

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FrontCover: Chrysaora quinquecirrha (photograph © George Grail/National Graphic Image Collection).

Back Cover: Stomolophus meleagris (photograph by R.J. Larson).
Contents

Abstract 7

Introduction 7

Materials and Methods 8

Results 9

Classification 9

Key 10

Systematic Account 11

Class Cubozoa Werner, 1973 11

Alatina alata (Reynaud, 1830) 12

Tamoya haplonema F. Miiller, 1859 14

Chiropsalmus quadrumanus (F Miiller, 1859) 15

Class Scyphozoa Goette, 1887 18

Atolla wyvillei Haeckel, 1880 18

Linuche unguiculata (Swartz, 1788) 19

Nausithoe punctata Kolliker, 1853 21

Periphylla periphylla (Peron and Lesueur, 1810) 23

Chrysaora quinquecirrha (Desor, 1848) 24

 Pelagia noctiluca (Forsskal, 1775) 28

Cyanea capillata fulva L. Agassiz, 1862 30

Cyanea capillata versicolor L. Agassiz, 1862 32

Drymonema dalmatinum Haeckel, 1880 33

Aurelia marginalis L. Agassiz, 1862 35

Phyllorhiza punctata von Lendenfeld, 1884 38

Rhopilema verrilli(Fewkes, 1887) 40

Stomolophus meleagris L. Agassiz, 1860 42

Acknowledgements 46

Glossary 47

Literature Cited 48
 Cubozoan and Scyphozoan Jellyfishes

Abstract

Asynopsis is given of three species ofcubomedusae (Alatina data, Tamoya haplonema, and Chiropsalmus quadrumanus) and
13 species and subspecies of scyphomedusae (Atolla wyvillei, Linuche unguiculata, Nausithoe punctata, Periphylla periphylla,
Chrysaora quinquecirrha, Pelagia noctiluca, Cyanea capillata julva, Cyanea capillata versicolor, Drymonema dalmatinum, Amelia
marginalis, Phyllorhiza punctata, Rhopilema verrilli, and Stomolophus meleagris) reported from the Carolinian Biogeographic
Province of the southeastern United States (Cape Hatteras, North Carolina to Cape Canaveral, Florida). Regional re
cords of eachspecies are documentedby reference to literature reports and to museum specimens. Summaries are given
of the biology, life cycle, abundance, nematocyst complement (where known), toxicity, andreported distribution of each
species. Sampling undertaken each of the four seasons near Charleston, South Carolina, indicated that Cyanea capillata
versicolor was the most prevalent species inshore during colder months, whereas Stomolophus meleagris, Chrysaora quin
quecirrha, and Chiropsalmus quadrumanus were the most abundant medusae during warmer months. The invasive species
Phyllorhiza punctata has now been reported from scattered locations across the study area. A new subclass, Coronamedu-
sae, is established to accommodate the order Coronatae, representing a different clade from the existing nominal subclass
Discomedusae (encompassing the orders Semaeostomeae and Rhizostomeae).The species of Amelia indigenous to the
region, usually identified as A. aurita Linnaeus, 1758 (type locality: Baltic Sea), is distinctly different from that boreal
species and is assigned instead to A. marginalis L.Agassiz, 1862 (type locality: Florida). Nomenclatural matters consid
ered here included authorship and dates of names, and type species of genera.The widely used generic name Periphylla
F. Miiller, 1861 is designated as both valid and a nomen protectum, while its virtually unused senior synonym Cyclophylla
Brandt, 1837 is relegated to a nomen oblitum. No valid type species designation was found for the well-known genus
Cyanea Peron and Lesueur, 1810; Cyanea arctica Peron and Lesueur, 1810 (ajunior subjective synonym of Medusa capillata
Linnaeus, 1758) is hereby so designated. Nomenclatural problems over the name Phyllorhiza L.Agassiz, 1862 exist due to
a longstanding misconception about its type species. Current usage is maintained pending a submission to the Interna
tional Commission on Zoological Nomenclature asking that the accepted concept of the genus be upheld.

Introduction the seasonally abundant cannonball jellyfish (Stomolophus

meleagris) given the considerable demand for jellyfishes
Scyphozoan and cubozoan medusae are an important as food in parts of East Asia (Omori and Nakano, 2001).
component of the plankton in waters of the Carolin These cnidarians have been little studied in the south

ian Biogeographic Province along the southeast coast of
the United States (defined here as Cape Hatteras, North
Carolina, to Cape Canaveral, Florida) because of their
typically large size, high frequency of occurrence, roles
as both predators and prey, and associations with other
taxa (especially fishes and crustaceans). During periods
of abundance they may constitute a hindrance to com
mercial fishing in the region, and especially to shrimp
trawling operations. Moreover, several species occurring
in these waters are venomous to humans and of concern
to public health. Meanwhile, a potential fishery exists for
eastern United States, however, and existing information
on them is widely scattered.
Some 200 species of"true jellyfishes" or acraspedote
medusae (Scyphozoa), and about 40 species of box jel
lies (Cubozoa), are currently recognized worldwide. Of
these, 13 species and subspecies of Scyphozoa and three
species of Cubozoa are known to occur from inshore
waters seawards to the Gulf Stream between Capes Hat
teras and Canaveral. Among these is the well-known in
vasive species Phyllorhiza punctata, originally described
from Australia and recently identified from the study
area (Johnson and Allen, 2005; Lauren Hall, pers. comm.,
2006;Justin Ellenberger, pers. comm., 2006; David Knott,
pers. comm., 2007). Drymonema dalmatinum, another scy-
phomedusa initially thought to be recently invasive in the
western North Atlantic (Larson, 1987a), is now known
to have been present in waters off this coast more than
120 years ago. Three specimens of this medusa, taken
off the coast of South Carolina in 1885, were recently
discovered in collections at the National Museum of
Natural History, Smithsonian Institution (Williams et al.,
2001). Several other cubozoan and scyphozoan medusae
generally limited to the eastern margins of the study area
(Alatina alata, Atolla wyvillei, Linuche unguiculata, Naus-
ithoe punctata, and Periphylla periphylla) are also included
here. As for the number of species of Scyphozoa overall,
Dawson (2004) postulated that diversity may be about
two times the number currently recognized on the ba
sis of morphology. The biodiversity of cubomedusae is
also believed to be substantially higher than previously
thought (Gershwin, 2005).These and other recent stud
ies have provided evidence of extensive crypsis in scy-
phozoans and cubozoans, and the species-level taxono
my adopted herein seems likely to change in the near
term. Thus, the conundrum taxonomists have faced in
distinguishing species and in analyzing distributions of
Hydrozoa (Calder, 1988:2) applies to the Scyphozoa and
Cubozoa as well.
Scientific and public interest in jellyfishes has in
creased in recent decades. Their ecological importance,
particularly as planktonic carnivores, has become more
widely appreciated (Purcell, 1992; Mills, 1995; Arai,
1997). Large medusae are also now recognized as im
portant prey of certainsea turtles, especially leatherbacks
(Dermochelys coriacea) (Bleakney, 1965; Grant and Ferrell,
1993; Lutz and Musick, 1996; Frick et al., 1999). Con
cerns about venomous species have prompted research
on these cnidarians in a number of countries, especially
Australia, the United States, and Japan. An apparent in
crease in the incidence and severity of jellyfish blooms,
and the negative impacts of such swarms on ecosystems
and human activities in various parts of the world, has
also been noted and investigated (Purcell, 2005; Purcell et
al., 2001). In addition, invasive species such as Phyllorhiza
punctata, noted above, are now the subject of numerous
ongoing studies. Finally, jellyfish exhibits have become
popular attractions at institutions such as the Monterey
Bay Aquarium, the New England Aquarium, and the
South Carolina Aquarium.
This study was undertaken to document the species
composition, geographic distributions, seasonal occur
rences, and historical records of scyphozoan and cubo
zoan medusae in the region, and to provide a key to their
identification.
Materials and Methods
The geographic limits of this study extend from Cape
Hatteras, North Carolina, to Cape Canaveral, Florida,
and from estuarine waters inshore to the Gulf Stream
offshore. Known locally as the South Atlantic Bight, that
confusing and inappropriate name is avoided here be
cause it may be mistaken by some outside the immediate
region for a location in the South Atlantic Ocean. The
northern limit of the study area, as defined herein, in
cludes Pamlico Sound, North Carolina, and the southern
limit includes the Banana River, Florida. The region cor
responds with what is commonly known by marine bio-
geographers as the Carolinian Province (Briggs, 1974).
While the northern Gulf of Mexico is also generally
considered part of the same biogeographic province, the
medusae of that region were not considered here.
This account is limited largely to the medusa stages
of Scyphozoa and Cubozoa; the polyp stages of these
cnidarians (with the possible exception of those of Rho~
pilema verrilli and Stomolophus meleagris) are insufficiently
known in the region to discuss in any detail at present.
Data presented herein are based on literature records, on
specimens in collections ofvarious institutions (National
Museum of Natural History, Washington, DC; Royal
Ontario Museum, Toronto, ON; Marine Resources Re
search Institute, Charleston, SC; Grice Marine Labora
tory, Charleston, SC), and on new material obtained
as part of this investigation. Collecting was undertaken
once each season in 2005 during the study (4—12 Febru
ary, 9-20 May, 28 August-2 September, 30 October-4
November) in coastal South Carolina while working at
the Southeastern Regional Taxonomic Center (SERTC)
of the Marine Resources Research Institute, South
Carolina Department of Natural Resources, in Charles
ton. Sampling was undertaken each quarter at stations
in Inlet Creek, Charleston Harbor, and at the entrance
of Charleston Harbor. At each site, qualitative collec
tions were made using an otter trawl (10-minute tow),
a half-metre bongo net (4-5 minute tow), and a one-
metre-wide neuston net (4-5 minute tow). Surveys for
stranded but recently live medusae were also undertaken
each sampling period on Folly Beach, South Carolina.
Abbreviations in the text are as follows: GMBL (Grice
Marine Biological Laboratory [presently known as the
Grice Marine Laboratory], The College of Charleston,
Charleston, South Carolina); ICZN (International Code
of Zoological Nomenclature); ROMIZ (Invertebrate
Zoology collections at the Royal Ontario Museum, To
ronto, Ontario, Canada); SERTC (Southeastern Region
al Taxonomic Center, Marine Resources Research Insti
tute, South Carolina Department of Natural Resources,
Charleston, South Carolina), USNM and NMNH (Na
tional Museum of Natural History, Smithsonian Institu
tion, Washington, DC).
Synonymy lists are limited to the following works:
(1) the original publication in which a given name was
made available nomenclaturally, and (2) references to
works which have provided original records of a given
species in the study area. Matters of nomenclature were
considered with reference to provisions of the ICZN
(International Commission on Zoological Nomencla
ture, 1999). An attempt was made to trace and confirm
authorship and dates of all nominal taxa used herein at
the time they were made available under provisions of
the code. A similar effort was made to determine the
valid type species of each genus, the way in which it was
so designated, and the author and date of that designation
if it was fixed subsequent to establishment of the name.
Scientific names above the family group have been at
tributed to the author who first used them, regardless of
the level in the Linnaean hierarchy at which they were
originally proposed. Thus, the phylum name "Cnidaria"
is credited to Verrill (1865), even though he used the
term at the rank of class.
Several recently proposed changes in the classifica
tion of Scyphozoa have been adopted here. Marques and
Collins (2004) suggested, based on a detailed cladistic
analysis of medusozoans, that the monophyletic clade
comprising Coronatae, Rhizostomeae, and Semaeos-
tomeae (Scyphozoa) is a sister group of another includ
ing the Cubozoa (together with the Stauromedusae and
the fossil Conulatae). They proposed recognizing both
the Cubozoa and Staurozoa as classes within the phylum
Cnidaria, along with Hydrozoa, Scyphozoa, and Antho-
zoa. Moreover, they provided evidence that the scypho
zoan orders Semaeostomeae and Rhizostomeae are more
closely related to each other than to Coronatae. The
name Discomedusae Haeckel, 1880 has been applied to
this subclass (Marques and Collins, 2004; Dawson, 2004).
The order Coronatae, excluded in that work from the
Discomedusae, is herein assigned instead to an equivalent
new subclass, Coronamedusae.
The classification and nomenclature of nemato-
cysts both continue to evolve from the classic work by
Weill (1934a, b). Stability of names of these organelles
is desirable in order to minimize confusion, yet changes
sometimes become necessary when knowledge of them
advances. In this work, I have essentially followed the
nematocyst terminology used earlier (Calder, 1974b;
Mariscal, 1974), with some necessary modifications in
light of more recent work. Data on nematocyst sizes giv
en here are measurements ofmaximum length and width
of undischarged capsules.
The terms bell, umbrella, and disk have been used
interchangeably in the literature for the body of a jel
lyfish exclusive of the manubrium, oral arms, tenta
cles, and other parts that are suspended from it (Cor
nelius, 1997:116—117). The term umbrella has been
used throughout in this work, even for the bell-shaped
cubomedusae. This was done because the terms exum-
brella for the upper and outer side of this structure, and
subumbrella for the lower and inner side, are almost uni
versally adopted.
Results
Classification
CLASS CUBOZOA Werner, 1973
ORDER CARYBDEIDA Lesson, 1843
Family Alatinidae Gershwin, 2005
Genus Alatina Gershwin, 2005
Alatina alata (Reynaud, 1830)
Family Tamoyidae Haeckel, 1880
Genus Tamoya E Miiller, 1859
Tamoya haplonema E Miiller, 1859
ORDER CHIRODROPIDA Haeckel, 1880
Family Chiropsalmidae Thiel, 1936
Genus Chiropsalmus L.Agassiz, 1862
Chiropsalmus quadrumanus (FMiiller, 1859)
CLASS SCYPHOZOA Goette, 1887
SUBCLASS CORONAMEDUSAE, subcl. nov.
ORDER CORONATAEVanhoffen, 1892
Family Atollidae Hickson, 1906 (1880)
Genus Atolla Haeckel, 1880
Atolla wyvillei Haeckel, 1880
Family Linuchidae Haeckel, 1880
Genus Linuche Eschscholtz, 1829
Linuche unguiculata (Swartz, 1788)
Family Nausithoidae Haeckel, 1880
Genus Nausithoe Kolliker, 1853
Nausithoe punctata Kolliker, 1853
 Family Periphyllidae Haeckel, 1880
Genus Periphylla F. Miiller, 1861
Periphylla periphylla (Peron and Lesueur, 1810)
SUBCLASS DISCOMEDUSAE Haeckel, 1880
ORDER SEMAEOSTOMEAE L. Agassiz, 1862
Family Pelagiidae Gegenbaur, 1857
Genus Chrysaora Peron and Lesueur, 1810
Chrysaora quinquecirrha (Desor, 1848)
Genus Pelagia Peron and Lesueur, 1810
Pelagia noctiluca (Forsskal, 1775)
Family Cyaneidae L. Agassiz, 1862
Genus Cyanea Peron and Lesueur, 1810
Cyanea capillatafulva L.Agassiz, 1862
Cyanea capillata versicolor L.Agassiz, 1862
Genus Drymonema Haeckel, 1880
Drymonema dalmatinum Haeckel, 1880
Family Ulmaridae Haeckel, 1880
Genus Amelia Lamarck, 1816
Amelia marginalis L. Agassiz, 1862
ORDER RHIZOSTOMEAE Cuvier, 1799
SUBORDER KOLPOPHORAE Stiasny, 1921b
Family Mastigiidae Stiasny, 1921b
Genus Phyllorhiza L.Agassiz, 1862
Phyllorhiza punctata von Lendenfeld, 1884
SUBORDER DAKTYLIOPHORAE Stiasny, 1921b
Superfamily Rhizostomatoidea Cuvier, 1799
Family Rhizostomatidae Cuvier, 1799
Genus Rhopilema Haeckel, 1880
Rhopilema verrilli (Fewkes, 1887)
Family Stomolophidae Haeckel, 1880
Genus Stomolophus L.Agassiz, 1860
Stomolophus meleagris L.Agassiz, 1860
Key
1. Tentacles present on umbrella 4
— Tentacles lacking on umbrella 2
2. Umbrella almost spherical, with distinct brown band
around margin; appendages lacking on mouth-arms
Stomolophus meleagris
— Umbrella flatter than a hemisphere, lacking brown
band around margin; appendages or filaments on
lower surface of mouth-arms 3
3. Mouth-arms with stout finger-like appendages;
scapulets ("shoulder ruffles") present; exumbrella
without prominent white spots Rhopilema verrilli
— Mouth-arms with long and slender filaments basally;
scapulets absent; exumbrella with numerous promi
nent white spots Phyllorhiza punctata
4. Tentacles on underside of umbrella 5
— Tentacles on margin of umbrella 7
5. Tentacles in a wide band around underside of um
brella; medusae large (up to 1 m in diameter)
Drymonema dalmatinum
— Tentacles in eight U-shaped clusters on underside of
umbrella; medusae smaller than above (up to about
20 cm in diameter) 6
6. Medusa purplish pink to deep red
Cyanea capillata versicolor
— Medusa yellow to yellow-brown, resembling a fried
egg Cyanea capillatafulva

7. Umbrella flat, plate-shaped, with numerous small

tentacles around margin, with four horseshoe-shaped
gonads centrally Amelia marginalis
— Umbrella, tentacles, and gonads not as above 8

8. Umbrella cuboidal; tentacles arising from each of

four interradial corners 14
— Umbrella not cuboidal; tentacles not restricted to
four interradial corners 9

9. Umbrella with distinct circular groove on upper sur

face 11
— Umbrella without distinct circular groove on upper
surface 10

10
 Systematic Account

10. Umbrella with small wart-like clusters of stinging Phylum Cnidaria Verrill, 1865
organelles; margin with 3—5 or more long tentacles Subphylum Medusozoa Petersen, 1979
in each of eight groups; most prevalent in estuarine Class Cubozoa Werner, 1973
and inshore waters Chrysaora quinquecirrha
Order Carybdeida Lesson, 1843
— Umbrella with distinctly elevated gelatinous mounds;
margin with eight long tentacles; usually restricted
Remarks
to offshore waters Pelagia noctiluca
The family name Carybdeidae, from which the name of
this order is derived, has often been attributed to Gegen-
11. Umbrella higher than a hemisphere 12
baur (1857), but it was established prior to that by Lesson
— Umbrella decidedly flattened 13
(1843) as Carybdeae. Louis Agassiz (1862:173) attributed
the name to an even earlier work (Lesson, R. P. 1837.
12. Umbrella small (up to 2 cm high), thimble-shaped,
Prodrome d'une monographic des meduses. Rochefort.
brownish in colour; tentacles short, inconspicuous;
62 pp.). However, only "a very small number of copies"
in surface and near-surface waters offshore
of that document were issued (L.Agassiz, 1860:24), and
Linuche unguiculata
in my opinion it does not meet criteria of publication
— Umbrella large (up to 25 cm high), cone- to dome-
under the code (ICZN Art. 8.1.3). Accordingly, Lesson
shaped, red in colour; tentacles long, conspicuous; in
(1843) is credited here as the author of the order name
deep waters offshore Periphylla periphylla
Carybdeida, and the taxon it refers to is understood in
the sense ofWerner (1984).
13. Umbrella very small (up to 1.5 cm wide); tentacles
eight; gonads forming eight globular masses on un-
dersurface of umbrella Nausithoe punctata
— Umbrella larger (up to 15 cm wide), with thick cen Family Alatinidae Gershwin, 2005
tral lens; tentacles usually 22 (sometimes more)
Remarks
Atolla wyvillei
Gershwin (2005) redefined the family Carybdeidae and
14. Umbrella about as wide or more as high; tentacles established the family Alatinidae to accommodate her
about 7-9 on each of four hand-shaped pedalia new genus Alatina and the earlier described Manokia
Chiropsalmus quadrumanus Southcott, 1967. She assigned Caryhdea alata, a poorly
— Umbrella distinctly higher than wide; tentacles one known but frequently reported species in the western
Atlantic, to Alatina.
on each of four paddle-shaped pedalia 15

15. Medusa with large, deep, cross-shaped stomach; rho Genus Alatina Gershwin, 2005
paliar niche opening a horizontal or crescent-shaped
slit Tamoya haplonema Alatina Gershwin, 2005:504.
— Medusa with shallow, flask-shaped stomach; rhopa
Diagnosis
liar niche opening T-shaped Alatina alata
Medusa with clear, cuboidal umbrella; with four simple
pedalia and four tentacles; stomach shallow, with no mes
enteries; rhopaliar niche ostiaT-shaped, with one upper
scale and two lower scales; velarial canals three per octant;
gastric cirri long, mostly parallel and in a single plane,
forming four crescent-shaped phacellae (modified from
Gershwin, 2005).

Type Species
Alatina mordens Gershwin, 2005,by original designation.
Remarks
Gershwin (2005) founded Alatinaas a new genus and as
signed to it 11 nominal species,including Carybdea alata
(Reynaud, 1830).

Alatina alata (Reynaud, 1830)

(Fig- 1)

Carybdea (Medusa) alata Reynaud, 1830:95, pi. 33, fig. 1.

Carybdea alata — Larson, 1976a: 15.
Alatina alata — Gershwin, 2005:515.

Common Names
Winged boxjelly,sea wasp.

Type Locality
Atlantic Ocean ("...dans l'ocean Atlantique") (Reynaud,
1830).

Museum Material
North Carolina, Atlantic Ocean east of Cape Hatteras,
35°03' N, 74°41' W, 204-228 m, R/V Sands Station
606-9-T1-C, D, 06 April 1967, one medusa, USNM
53694.

Specific Characters
Medusa with characters of the genus; umbrella up to
6-8 cm high, 5 cm or more wide (up to 23 cm high and
14 cm wide in A. alata forma grandis); exumbrellar sur
face without warts; sensory niches covered by one scale
above and two below; stomach short, mouth surrounded
by four simple lips; gastric filaments forming horizontal
crescents at corners of stomach; pedalia spatula-shaped,
interradial; tentacles four, simple (Mayer, 1910b; Kramp,
1961).

Nematocyst Complement
Medusa, newly metamorphosed from polyp (Puerto
Rico;Arneson and Cutress, 1976):
Exumbrella—
microbasic euryteles (20 jam long x 8 um wide)
holotrichs (spherical) (13 jam in diameter)
Tentacles—
euryteles (30 um long x 7 jam wide)
Gastric cirri—
euryteles (small, no measurements given)

Remarks
The binomen Carybdea alata (recently renamed Alatina Fig. 1. Alatina alata (Reynaud, 1830),after Sterrer (1986) and
alata by Gershwin, 2005) has been applied to certain Mianzan and Cornelius (1999).
boxjellies worldwide, but more than one species is now
thought to exist under that name (Morandini, 2003;
Gershwin, 2005). Uncertainty also exists over the iden
tity of the species upon which Reynaud (1830) based
the name, because the original description was so lack
ing in detail. No type material is known to exist. The
type locality was reported by Kramp (1961) to have been
somewhere in the South Atlantic Ocean. While the spe
cies of the genus from the warm western Atlantic may
be the same as that called C. alata by Reynaud (1830),
the nomenclature of the species needs to be stabilized
by designation of a neotype. Gershwin (2005) expressed
the intention to do this once a suitable specimen could
be located.
Arneson and Cutress (1976) described the life cycle
of a species identified as Carybdea alata based on cul
tures of specimens from Puerto Rico. Spawning of me
dusae was observed there during July (1973) and August
(1974). Fertilization occurred within the ovary, and zy
gotes developed into blastulae before being shed into the
water. These became free-swimming planulae after one
day. Settlement of the planulae occurred about four days
later, and four rudimentary tentacles were apparent after
another couple of days. Some 54 days after fertilization,
with polyps having 12 tentacles, asexual reproduction by
budding began. Metamorphosis commenced two weeks
later in polyps possessing 16 tentacles: this involved re
sorption of the tentacles and hypostome, and gradual
transformation of the polyp into a medusa.A change in
cnidome accompanied this process, with stenoteles oc
curring in the polyp but not the medusa, and spherical
holotrichs occurring in the medusa but not the polyp.
Medusae were liberated one week after metamorphosis
began. A small fragment of the polyp adhering to the
apex of the umbrella was absorbed within 10-13 hours.
This general life cycle,with asexualreproduction by bud
ding and with a single polyp metamorphosing entirely
into a single medusa, is similar to that described in other
species of cubozoans (Werner et al., 1971; Franc, 1995b;
Stangl et al., 2002; Fischer and Hofmann, 2004).
According to Arneson and Cutress (1976), medusae
of Carybdea alata (= Alatina alata) are strong swimmers,
and while neritic are usually confined to cleaner offshore
waters. They rise to the surface at night to feed. From
studies in Puerto Rico, food items included polychaetes,
mysids and crab megalopae (Larson, 1976b). Medusae
of this species are thought to live for more than a year
(Arneson and Cutress, 1975).
Larson (1976a) recorded the occurrence of Alati
na alata (as Carybdea alata) in the study area based on a
specimen at the NMNH (USNM 53694), as cited above.
This species has also been collected immediately east of
the region, in the Sargasso Sea, at 32°33' N, 72° 14' W
(USNM 42017).
The cubomedusa in the western North Adantic
known as Carybdea alata (now Alatina alata) is known to
be venomous to humans (Humann, 2002).The nematocyst
complement of medusae identified as C. alata from Hawaii
incorporated various categories, including those identified as
heterotrichous microbasic euryteles (Yanagihara et al.,2002).
Reported Range
Study area: Cape Hatteras (Larson, 1976a).
Overall: reported to be circumtropical, including Gulf of
Mexico; neritic (Kramp, 1961,1970a; Graham, 1998;
Morandini, 2003).
Family Tamoyidae Haeckel, 1880
Genus Tamoya E Miiller, 1859
Tamoya F. Miiller, 1859:1.
Diagnosis
Medusa with clear, cuboidal umbrella, with height ex
ceeding width; four simple, scalpel-shaped pedalia and
four flattened tentacles; stomach large, deep, cruciform,
with four well-developed mesenteries extending to
subumbrellar wall; rhopaliar niche ostium essentially a
horizontal to crescent-shapedslit; stomach pouches four,
lacking diverticula; gastric cirri in four vertical bands
along interradial walls of stomach; velarial canals
dendritic, numerous; eyes six per rhopalium (two me
dian and four lateral) (Mayer, 1910b; Kramp, 1961;
Cornelius, 1997; Mianzan and Cornelius, 1999; Gershwin
and Alderslade, 2005).
Type Species
Tamoya haplonema F. Miiller, 1859,by subsequentdesigna
tion by Haeckel (1880).
Remarks
The generic name Tamoya was introduced by F. Miiller
(1859) for two new species of cubomedusae from the
coast of Brazil, T haplonema and T quadrumana. L.Agas
siz (1862) referred T quadrumana to a new genus, Chi
ropsalmus, and restricted Tamoya to "...the species with
simple tentacular lobes" (i.e. T haplonema). Such elimina
tion does not qualify as a valid type species designation
(Art. 69.4, ICZN), however, and Haeckel (1880:443) is
i3
taken as the first author to validly designate T haplonema
as the type species of Tamoya.

Tamoya haplonema F. Miiller, 1859

(Fig-2)

Tamoya haplonema F. Miiller, 1859:1, pis. 1-3. — Brooks,

1883:138.— Hoese, 1973:74. — Kraeuter and
Setzler, 1975:67.— Calder and Hester, 1978:88.

Common Name
Glassy boxjelly.

Type Locality
Brazil, Desterro (F. Miiller, 1859).

Museum Material
South Carolina, continental shelf east of Edisto Island,
R/V Dolphin, MARMAP coll. no. 0573410,32°37.0' N,
78°30.0' W, 141 m, 22°C, 36.7%o, %Yankee No. 36 trawl,
9 November 1973, one medusa, ROMIZ B3589. South
Carolina, continental shelfeast ofEdisto Island, 32°33.4' N,
79°25' W, 21 m, trawl, 30 March 1979, two medusae,
GMBL 79-3. South Carolina, Charleston Harbor off
Fort Sumter, R/V Anita, 32°45.5' N, 79°52.9'W, 3-9 m,
21 °C, 20-ft otter trawl, 27 October 1979, two medusae,
GMBL 79-176. South Carolina, continental shelf east of
Edisto Island, R/V Dolphin, Station IS01, 32°28'54n N,
79°42'36" W, 17 m, 17January 1980, one medusa, USNM
61670. South Carolina, continental shelf east of Edisto
Island, 32°29.1' N, 79°41.7f W, 18 m, 14.8 °C, 34.5%>,
27 March 1981, one medusa, GMBL 81-6. Georgia, Fig. 2. Tamoya haplonema F. Miiller, 1859, after Mayer (1910b).
North Ossabaw Sound, 31°51.73' N,81o01.48'W, 12.8 m,
27.13 °C, 31.83%o, bottom sled, 17 June 1998, one
medusa, SERTC No. S2191. Georgia, North Ossabaw tia frown-shaped; stomach large, mouth surrounded by
Sound, 31°51.73' N, 81°01.48' W, 0.5 m, 27.30 °C, four recurved lips;gastric filaments in four vertical bands;
31.70%o, neuston net, 18 June 1998, two medusae, pedalia interradial, flat, spatula to scalpel-shaped, with
SERTC No. S2184. Georgia, North Ossabaw Sound, sharp edges; tentacles four, simple, interradial, flattened
31°51.73' N, 81°01.48' W, 0.5 m, 27.20 °C, 31.97%>, in cross-section; velarial canals 10 per quadrant, forked in
neuston net, 18 June 1998, two medusae, SERTC No. complex patterns (Mayer, 1910b;Kramp, 1961;Gershwin
S2185.South Carolina, seawardofbuoy 17,off Charleston and Alderslade, 2005).
jetties, R/V Silver Crescent, 32°42.07' N, 79°47.29' W,
9 m, 28.7 °C, 31.6%o, trawl, 1 September 2005, one Nematocyst Complement
medusa, ROMIZ B3611. Medusa (Sao Sebastiao, Brazil; Gershwin and Alderslade,
2005; Gershwin, 2006b):
Specific Characters Tentacles—
Medusa with characters ofthe genus; umbrella up to 9 cm club-shaped mastigophores (56.4—59.5 jamlong
high, 5.5 cm wide; exumbrellar surface with small warts x 15.7-17.4 um wide)
bearing batteries of nematocysts; sensory niches covered sub-spherical isorhizas (29.9—36.2 um long
by one scale above and one below; rhopaliar niche os- x 19.7-28.4 jam wide)

14
Remarks
Although indigenous to the region, T haplonema has re
ceived little study here and the biology of the species is
poorly known. Mayer (1910b) noted that the medusa,
with its tough and rigid umbrella, is"exceedingly active."
Tamoya haplonema is "...capable of inflicting a severe
sting to the hand" (Mayer, 1910b:513). Morandini and
Marques (1997) reported the envenomation of a 23-
year-old male in Sao Paulo state, Brazil, by a medusa of
this species. Mianzan and Cornelius (1999) commented
that it is a dangerous species with a sting that is painful
but not lethal. I have handled several medusae taken in
trawls off the South Carolina coast in the past with no
noteworthy sting, but the specimens were damaged to
some extent and many of their nematocysts may have
been lost or discharged.
The nematocyst complement of X haplonema has yet
to be studied in detail. Morandini and Marques (1997)
noted the presence of microbasic mastigophores on
the tentacles of a specimen from Brazil. Gershwin and
Alderslade (2005) and Gershwin (2006a) reported club-
shaped mastigophores and subspherical isorhizas in a
medusa of the species. The specimen they examined was
from the Sao Sebastiao Channel, Sao Paulo state, Brazil,
and the nematocysts examined were from the tentacles
(Lisa-ann Gershwin, pers. comm., 2005).
This cubozoan has been reported every month from
May through December in the study area (Hoese, 1973;
Kraeuter and Setzler, 1975; Calder and Hester, 1978). In
addition, material recorded here included specimens col
lected from offshore waters during the months ofJanu
ary and March. While medusae of T haplonema penetrate
a limited distance into sounds and harbors during the
warmer months (Mayer, 1910b; this study: GMBL 79-
176; SERTC No. S2191; SERTC No. S2184), they ap
pear most prevalent in waters of relatively high salinity
along the coast. Specimens are common in shrimp trawl
catches offshore (personal observations).
John McCrady, well known for pioneering work
on Hydrozoa (Stephens and Calder, 1992), described a
cubomedusa from South Carolina in 1876 that he called
"Cheiropsalmus vitreus" in an unpublished manuscript.
The species he described is almost certainly the same one
that has been called Tamoya haplonema in the southeastern
United States (Stephens, 2000:248), but the name used
by McCrady has never been made available.
Molecular studies comparing medusae of Tamoya
haplonema from Brazil with those identified as the same
species from the Atlantic and Gulf coasts of the United
States are needed.
Reported Range
Study area:Beaufort, NC (Brooks, 1883); off Sapelo Is
land, GA (Hoese, 1973; Kraeuter and Setzler, 1975);
coastal waters from Bull Bay to St. Helena Sound,
SC (Calder and Hester, 1978).
Overall: western Atlantic, from Long Island, New York,
to Argentina; Gulf of Mexico; eastern Atlantic; ner
itic (Mayer, 1910b; Kramp, 1961; Phillips and Burke,
1970; Burke, 1975; Mianzan and Cornelius, 1999;
Pastorino,2001).
Order Chirodropida Haeckel, 1880
Family Chiropsalmidae Thiel, 1936
Genus Chiropsalmus L. Agassiz, 1862
Chiropsalmus L. Agassiz, 1862:174.
Diagnosis
Medusa with clear, cuboidal umbrella; pedalia interradial,
four, hand-shaped, with tentacles inserting at end of each
finger; stomach pouches four, with eight finger-shaped,
unbranched diverticula extending into subumbrellar cav
ity; gonads eight, leaf-shaped, with free margins lacking
grape-shaped swellings (Mayer, 1910b; Kramp, 1961).
Type Species
Tamoya quadrumana F. Miiller, 1859, by monotypy.
Chiropsalmus quadrumanus (E Miiller, 1859)
(Fig. 3)
Tamoya quadrumana F. Miiller, 1859:1, pis. 2,3.
Cheiropsalamus quadrumanus — Brooks, 1883:137
[incorrect subsequent spelling].
Chiropsalmus quadrumanus — Conant, 1898:4. — Mayer,
1910b:515, pi. 57, fig. 3. — Kraeuter and Setzler,
1975:68. — Calder and Peters, 1975:364. — Calder
and Hester, 1978:88.
Chiropsalmus quadrimanus — Pearse et al., 1942:181
[incorrect subsequent spelling].
Chiropsalmus quadramanus — Hoese, 1973:94
[incorrect subsequent spelling].
Common Name
Fourhanded boxjelly.
Type Locality
Neotype from Brazil, Sao Paulo state, Ubatuba, Costao
do Carneiro, off Enseada Beach (Gershwin, 2006a).
15

Fig.3. Chiropsalmus quadrumanus (F. Miiller, 1859), after Mayer
(1910b).
Museum Material
South Carolina, Charleston Harbor, South Carolina De
partment of Natural Resources, Marine Resources Di
vision boat slip, 32°45.2' N, 79°53.9' W, surface, 27 °C,
19%o, dipnetted, 22 July 1971, nine medusae, GMBL
71-121. South Carolina, Charleston Harbor, 32°46.2' N,
79°54.8'W, 8-10 m, 29 °C, otter trawl, 10 August 1971,
one medusa, GMBL 71-139. South Carolina, Inlet Creek
near entrance (no coordinates), 4 m, 30 °C, 34%o, 20-ft
otter trawl, 4 August 1972, one medusa, GMBL 72-190.
South Carolina, Charleston Harbor, off Fort Johnson,
R/V Anita, 32°45.2' N, 79°54.5' W, 26.96 °C, try net,
24 June 1985, one medusa, GMBL 85-25. South Caro
lina, off Morris Island, 32°43.3' N, 79°54.1» W, 28.16 °C,
31.14%o, otter trawl, 24 September 1970, one medusa,
GMBL 70-182. South Carolina, Port Royal Sound, R/V
Anita, 32°16.2' N, 80°43.2' W, 7 m, 20-ft otter trawl, 7
August 1974, one medusa, ROMIZ B1179. South Car
olina, Inlet Creek, R/V Silver Crescent, 32°46.65' N,
79°49.19'W, 5 m, 28.2 °C, 31.6%o, trawl, 1 September
2005,30 medusae, ROMIZ B3612. South Carolina, Inlet
Creek, R/V Silver Crescent, 32°46.65' N, 79°49.19'W,
16
surface, 28.2 °C, 31.6%o, neuston net, 1 September 2005,
one medusa, ROMIZ B3613. South Carolina, Inlet
Creek, R/V Silver Crescent, 32°46.65' N, 79°49.19'W,
surface, 28.2 °C, 31.6%o, neuston net, 1 September 2005,
one young medusa, ROMIZ B3617. South Carolina,
Charleston Harbor seaward of buoy 17 off Charleston
jetties, R/V Silver Crescent, 32°42.31' N, 79°47.77' W,
8 m, 31.6%o, 28.7 °C, 20-ft otter trawl, 1 September 2005,
eight medusae (two in 95% ethanol), SERTC No. S2112.
South Carolina, seaward ofbuoy 17 off Charleston jetties,
32°42.31' N, 79°47.77' W, 8 m, 28.7 °C, 31.6%o, otter
trawl, 1 September 2005, two medusae, ROMIZ B3606.
South Carolina, seaward of buoy 17 off Charleston jet
ties, 32°42.31' N, 79°47.77f W, 8 m, 28.7 °C, 31.6%>,
trawl, 1 September 2005, one medusa (in 95% ethanol),
ROMIZ B3608. South Carolina, Charleston Harbor off
jetties, R/V Silver Crescent, 32°41.63' N, 79°47.68'W,
9 m, 31.6%o, 28.7 °C, 20-ft otter trawl, 1 September
2005, seven medusae, SERTC No. S2113. South Caro
lina, sea buoy off Charleston Harbor, R/V Silver Cres
cent, 32°38.53' N, 79°39.85' W, 16 m, 34.5%o, 28.8 °C,
20-ft otter trawl, 1 September 2005, six medusae, SERTC
No. S2114. South Carolina, Folly Beach at Stono Inlet,
32°38.35' N, 79°58.42' W, 20 °C, stranded on beach,
30 October 2005, one medusa (in poor condition), ROMIZ
B3619.
Specific Characters
Medusa with characters of the genus; umbrella up to
10 cm high, 14 cm wide; exumbrellar surface mostly
smooth; opening of sensory niche triangular; stomach
large, wide, mouth surrounded by four triangular lips;
gastric saccule four, each with two finger-shaped sacs;
gastric filaments in four interradial, crescent-shaped
rows near base of stomach; pedalia with 5—9 irregularly
placed fingers and tentacles (Mayer, 1910b; Kramp, 1961;
Mianzan and Cornelius, 1999).
Nematocyst Complement
Medusa (Georgia and South Carolina, USA; Calder and
Peters, 1975):
Tentacles—
ellipsoidal holotrichous isorhizas (8.4—18.2 um long
x 3.5—5.1 um wide)
ovoid holotrichous isorhizas (4.9-10.9 um long
x 2.8-4.4 jam wide)
microbasic mastigophores (43.4—76.3 urn long
x 8.4-13.8 um wide)
large heterotrichous microbasic euryteles
(21.7-40.0 um long x 11.9-21.1 jamwide)
 medium heterotrichous microbasic euryteles
(11.9-20.9 jam long x 9.1-16.8 jam wide)
Manubrium—
medium heterotrichous microbasic euryteles
(9.8-18.8 jam long x 7.0-14.7 jam wide)
Exumbrellar warts—
ovoid holotrichous isorhizas (4.2-7.0 jamlong
x 2.8—4.2 jam wide)
microbasic mastigophores (63.0—99.6 jamlong
x 9.1-14.7 jam wide)
Gastric filaments—
small heterotrichous microbasic euryteles
(6.3-9.8 jam long x 3.5-7.0 |am wide)
Remarks
This medusa was originally described as Tamoya quadru
mana from Desterro, Brazil,by F. Miiller (1859). Gershwin
(2006a) was unable to locate a holotype, and designated
a specimen from Costao do Carneiro, in front ofEnseada
Beach, Ubatuba, Sao Paulo, Brazil, as the neotype. She
noted that morphological and molecular comparisons of
Brazilian and American forms were warranted, and sug
gested that populations long thought to be Chiropsalmus
quadrumanus may comprise more than a single species.
In particular, she noted differences in the cnidome of
medusae from South Carolina, described by Calder and
Peters (1975), from those of Brazil, describedby Marques
et al. (1997). The name C. quadrumanus is applied here
to the species in the study area pending more detailed
taxonomic study.
John McCrady, an early student of Cnidaria
from Charleston, SC, described this species in 1876 as
"Cheiropsalmus carolinensis" in an unpublished manuscript
(Stephens, 2000:247). The name has never been made
available nomenclaturally.
Kraeuter and Setzler (1975) found Chiropsalmus
quadrumanus to be moderately abundant at times in
Georgia. Moreover, large numbers were reported along
the coast from Daytona Beach to Jacksonville, Florida,
during late July and early August 2002 (Jellyfish Sting
Newsletter, January 2003). In South Carolina they
were relatively abundant both inshore (Inlet Creek)
and nearshore (seaward of the Charleston jetties) in
September 2005 during this study. Medusae have been
reported in the study area fromJune through November
(Hoese, 1973; Kraeuter and Setzler, 1975; Calder and
Hester, 1978).
While common to abundant along the coastbetween
capes Hatteras and Canaveral during the warmer months
of the year, medusae of Chiropsalmus quadrumanus appear
not to have caused serious public health problems in the
study area.Earlier, Kraeuter and Setzler (1975) stated that
there were no reports of the species being a problem in
Georgia. Nevertheless, this cubomedusa, with its painful
sting (Guest, 1959), has been blamed for a human fatality
on the coast ofTexas (Bengston et al., 1991). Caution is
warranted whenever box jellies are encountered.
The cnidome of cubomedusae differs considerably
from that of scyphomedusae,providing evidence that the
two groups represent different clades within the Me-
dusozoa (Calder and Peters, 1975). Nematocysts iden
tified as large heterotrichous microbasic euryteles from
this species are likely the same as those classified as large
trirhopaloids by Oba et al. (2004) in Chiropsalmus quadri
gatus [= Chiropsoides quadrigatus]. It is uncertain whether
nematocysts called medium euryteles and small euryte
les in Chiropsalmus quadrumanus might also be termed
trirhopaloids as well. For an overview of the nematocysts
and cnidome of cubomedusae,see Gershwin (2006b).
Stomach contents of Chiropsalmus quadrumanus from
Mississippi Sound included microcrustaceans (mostly
Lucifer sp.), mantis shrimp larvae, crab zoeae, crab mega-
lopae, and amphipods, as well as small fishes (bumpers,
harvest fish, larval sciaenids) (Phillips et al., 1969; Phillips
and Burke, 1970).
Reported Range
Study area: Beaufort, NC (Brooks, 1883; Conant, 1898;
Mayer, 1910b; Pearse et al., 1942); Doboy Sound
and off Sapelo Island, GA (Hoese, 1973; Kraeuter
and Setzler, 1975); nearshore waters, South Carolina
and Georgia (Calder and Peters, 1975);Winyah Bay,
Price Creek, Port Royal Sound, and Colleton River,
SC (Calder and Hester, 1978).
Overall: western Atlantic, from North Carolina to Brazil
and Gulf of Mexico; eastern Atlantic; other ques
tionable records from warm waters of the Pacific and
Indian oceans; neritic (Mayer, 1910b; Kramp, 1961;
Phillips and Burke, 1970; Burke, 1975; Mianzan and
Cornelius, 1999; Morandini, Soares, et al.,2006).
17
 Class Scyphozoa Goette, 1887
Subclass Coronamedusae, subcl. nov.
Order Coronatae Vanhoffen, 1892
Family Atollidae Hickson, 1906 (1880)

Remarks
Hickson (1906) is believed here to have been the first
author to use the name Atollidae. Fewkes (1886) com
mented on the marked similarity between the nomi
nal genera Atolla Haeckel, 1880 and Collaspis Haeckel,
1880. Bigelow (1909) regarded the two as congeneric,
and acted as First Reviser (ICZN Art 24.2) in assigning
precedence to the generic name Atolla. He also adopted
the name Atollidae in place of the family group name
Collaspidae Haeckel, 1880.The nameAtollidae, although
founded after Collaspidae, is validunder the current code Fig.4. Atolla wyvillei Haeckel, 1880, after Haeckel (1881b),
(ICZN Art. 40.2), having come into prevailing usage. Mayer (1910b),and Mianzan and Cornelius (1999).
For nomenclatural purposes it takes the date of priority
of the replaced name (i.e., Collaspidae Haeckel, 1880),
Museum Material
and is deemed to be a senior synonym of it (ICZN Art.
South Carolina, continental slope east of Georgetown,
40.2.1).
R/V Oregon, 33°24' N, 76°29' W, 30 January 1972,
552 m, two medusae, GMBL 72-423. South Carolina,
Genus Atolla Haeckel, 1880
continental slope east of Georgetown, R/V Oregon,
33°24' N, 76°29'W, 30 January 1972,552 m, one medusa,
Atolla Haeckel, 1880:488.
ROMIZ B3596.

Diagnosis
Medusa decidedlyflattened,disc-shaped, with deep coro
nal furrow on exumbrella;rhopalia and tentacles alternat
ing, equal in number, exceeding eight; marginal lappets
double the number of tentacles; gonads eight, adradial;
subgenital ostia four, interradial; coronal muscle well de
veloped (Mayer, 1910b; Kramp, 1961; Russell, 1970).
Type Species
Atolla wyvillei Haeckel, 1880, by monotypy.
Atolla wyvillei Haeckel, 1880
(Fig. 4)
Atolla wyvillei Haeckel, 1880:488.
Atolla bairdii Fewkes, 1886:936, pi. 1, pi. 2, pi. 3, figs. 1,2.
Common Name
Wyvilles crownjelly.
Type Locality
Antarctic Ocean ("Antarktischer Ocean [indischer und
atlantischerTheil]") (Haeckel, 1880).
Specific Characters
Medusa up to 15 cm in umbrella diameter, with thick
central lens and wide coronal groove; base of stomach
four-leaf-clover-shaped; tentacles and rhopalia alternat
ing, often 22 but as many as 36 in number; radial septa
diverging at proximal ends, extending beyond margin
of coronal muscle; lappets spatula-shaped with rounded
ends, usually 44 in number, lacking warts (Kramp, 1961;
Russell, 1970; Mianzan and Cornelius, 1999).
Nematocyst Complement
Undescribed.
Remarks
A detailed account of Atolla wyvillei, including a syn
onymy Hst, was given by Russell (1970). He considered
it to be a deep-sea medusa, havinga widespread distribu
tion in all oceans. From literature records, he concluded
that it occurs in maximum abundance at 500—1500 m
during the day, but may be found nearer the surface at
night and in areas of upwelling.The medusa is known to
be bioluminescent (Nicol, 1958; Russell, 1970; Herring
andWidder, 2004).One of the tentacles in medusae of A.
wyvillei is hypertrophied and used in prey capture (Hunt
and Lindsay, 1998), but it may be missing or unrecogniz
able in preserved material.
Fewkes (1886) documented the occurrence of Atolla
wyvillei (as A. bairdii) in the region off Cape Hatteras.
Mayer (1910b) observed that the species (again assigned
to A. bairdii) was widely distributed in deep waters of
the North Atlantic, but that it was also occasionally
found at the surface in the Gulf Stream off the Carolinas.
Bigelow (1928) reported it from waters immediately out
side of the study area, some 210 miles SE of Cape Hat
teras (34°27' N, 71°36'W),m collections of the Arcturus
Oceanographic Expedition. Material examined here was
collected from mesopelagic waters in the region of the
Gulf Stream off Georgetown, SC.
Bigelow (1938) observed that gonads develop early
in Atolla wyvillei, and he believed that medusae of the
species are fertile throughout much of their develop
ment. He also considered it likely that A. wyvillei is holo-
planktonic, based on its bathymetric distribution.
Although the cnidome of Atolla wyvillei has yet to be
characterized, "nematocyst cushions" with dense aggre
gations of nematocysts occur along the peripheral walls
of the gastrovascular sinuses in medusae of this species
(Russell, 1970).
Genus Linuche Eschscholtz, 1829
Linuche Eschscholtz, 1829:91.
Diagnosis
Medusa thimble-shaped, with coronal furrow incising ex
umbrella; rhopalia and tentacles alternating, both eight in
number; tentacles short, inconspicuous; marginal lappets
16, bluntly oval; gonads eight, in four crescent-shaped
pairs; stomach pouches giving rise distally to 2—3 whorls
of wart-shaped protuberances; symbiotic zooxanthellae
present (Mayer, 1910b; Kramp, 1961).
Type Species
Medusa unguiculata Swartz, 1788, by monotypy.
Linuche unguiculata (Swartz, 1788)
(Fig. 5)
Medusa unguiculata Swartz, 1788:199, pi. 6, figs. la-e.
Linuche unguiculata — Calder, 1974b: 170. — Larson,
1976a: 15.
Common Names
Thimble jellyfish; button jellyfish; sea thimble.

Reported Range Type Locality

Study area: Gulf Stream off Cape Hatteras, NC, West Indies, near Jamaica ("Vest-Indifka Sjon nara Jamai
35°12'10' N, 74°57'15" W, surface, Albatross Station ca...") (Swartz, 1788).
2110 (Fewkes, 1886, as A. bairdii).
Overall: widely distributed in all oceans; deep-sea
(Kramp, 1961; Russell, 1970).

Family Linuchidae Haeckel, 1880

Remarks
Haeckel (1880:494) included two subfamilies in his new
family Linergidae Haeckel, 1880: Linuchidae Haeckel,
1880 and Linanthidae Haeckel, 1880. The family group
name Linuchidae has subsequently been adopted in place
of Linergidae or Linanthidae, and it's type genus (Linuche
Eschscholtz, 1829) is a now held to be a senior synonym
of Linerges Haeckel, 1880 (Mayer, 1910b; Kramp, 1961).
The status of Linantha Llaeckel, 1880 is uncertain but was
included in Linuchidae by Kramp (1961). Linuchidae is
thus upheld here as the valid name of the family.
Fig. 5. Linuche unguiculata (Swartz, 1788), after Mayer (1910b).

19
Museum Material
North Carolina, continental slope east of Cape Hatteras,
R/V Sands, 35°06' N, 74°41' W, 6 April 1967, 229-307 m,
two medusae, USNM 53696. North Carolina, continen
tal shelf east of Cape Hatteras, R/V Eastward, between
35°21.6' N, 74°50.0» W and 35°30.0' N, 74°41.2' W,
25 April 1973, four medusae, ROMIZ B2331.
Specific Characters
Medusae up to 2 cm high, 1.6 cm wide; subumbrellar
protuberances in three whorls; ring canal absent (Mayer,
1910b; Kramp, 1961; Mianzan and Cornelius, 1999).
Nematocyst Complement
Medusa (North Carolina, USA; Calder, 1974b):
Tentacles, lappets, manubrium—
holotrichous a-isorhizas (6.8-9.3 um long
x 4.8—6.4 jam wide)
small heterotrichous microbasic euryteles
(10.1-13.2 jam long x 7.6-9.8 jam wide)
large heterotrichous microbasic euryteles
(13.2-16.5 um long x 9.6-11.9 um wide)
Remarks
Kramp (1970a) maintained that the genus Linuche
Eschscholtz, 1829, was monotypic, with the three nomi
nal species sometimes assigned to it (Medusa unguiculata
Swartz, 1788, Linerges draco Haeckel, 1880, and L. aquila
Haeckel, 1880) being identical. Following Mayer (1910b),
the Atlantic L. unguiculata is held here to differ from the
Indo-Pacific L. aquila in lacking a ring canal and in hav
ing three rows, rather than two, of subumbrellar protu
berances,although Bigelow (1928) had earlier discounted
such differences.
The complete life cycle of Linuche unguiculata was
observed by Ortiz-Corp's et al. (1987), who studied
specimens from Puerto Rico. Planula,scyphistoma, stro-
bila, ephyra, and medusa stages were observed and de
scribed. The colonial but unbranched scyphistoma stage
corresponded with polyps known as Stephanoscyphus
komaii Leloup, 1937, a name now regarded as a junior
synonym of L. unguiculata (Swartz, 1788). Silveira and
Morandini (1998a, b) added additional observations on
the life cycle based on studies in Brazil on scyphistomae
from nature and from laboratory cultures. In subtropi
cal waters around Sao Sebastiao, Sao Paulo state, asexual
reproduction by planuloid formation was more prevalent
than medusa formation (Silveira and Morandini, 1998b).
They also discovered the existence of dormant stages in
polyps of L. unguiculata (Silveira and Morandini, 1998a).
Medusae of Linuche unguiculata, highly varied in
morphology and brownish in colour due to symbiotic
zooxanthellae (Mayer, 1910b), are widespread in shallow-
waters of the tropics and subtropics.Vast swarms extend
ing over many miles are frequent in the warm western
Atlantic (Mayer, 1910b), with aggregations attributed to
a combination of Langmuir circulation and a circular
swimming behaviour (Larson, 1992). Most of the me
dusae in the study area are likely transported there from
regions to the south by the Gulf Stream and its eddies.
Medusae of Linuche unguiculata are mildly toxic
(Humann, 2002). Moreover, the species has been implicated
as a cause of"seabather's eruption" (SBE) (Segura Puertas
et al., 1999). Planula larvae,as well as ephyrae and medusae
of this species, can cause this skin ailment. An outbreak
of SBE from Jacksonville to the Florida Keys during
1995 was documented in the Jellyfish Sting Newsletter
(January 1996).
The nematocysts of the scyphistoma, ephyra, and
planuloid stages of Linuche unguiculata, as described in
Brazil by Silveira and Morandini (1998b), are similar to
those given above for the medusa. A detailed energy bud
get for the species was characterized by Kremer (2005).
Specimens in collections at the Royal Ontario Museum
(ROMIZ B2331) are from slightly north of Cape Hat
teras, but have nevertheless been considered here.
Reported Range
Study area: Cape Hatteras (Calder, 1974b; Larson,
1976a).
Overall: amphi-Atlantic, in warm waters; neritic
(Mayer 1910b;Kramp,1961;Ortiz-Corp'setal.,1987;
Mianzan and Cornelius, 1999; Morandini et al., 2005;
Morandini, Soares, et al., 2006).
Family Nausithoidae Haeckel, 1880
Genus Nausithoe Kolliker, 1853
Nausithoe Kolliker, 1853:323.
Diagnosis
Medusa with coronal furrow dividing exumbrella into a
dome-shaped to flattened central disk and a peripheral
margin bearing 16 spade-shaped lappets; rhopalia and
tentacles alternating, both eight in number; gonads eight,
adradial; stomach pouches 16, simple (Mayer, 1910b;
Kramp, 1961; RusseU, 1970).
Type Species
Nausithoe punctata Kolliker, 1853,by subsequent designa
tion by Mayer (1910b).

Remarks
The generic name Stephanoscyphus Allman, 1874a, fre
quently applied to the scyphistoma stages of coronate
scyphozoans, is a junior subjective synonym of Nausithoe
Kolliker, 1853. Stephanoscyphus mirabilis Allman, 1874a,
type species of Stephanoscyphus by monotypy, has been
shown to be the polyp stage of the medusa known as
Nausithoe punctata Kolliker, 1853 (Jarms, 1997).The latter
is the type species of Nausithoe Kolliker, 1853 by sub
sequent designation by Mayer (1910b). Stephanoscyphus
was described as new in three separate papers by Allman
(1874a, 1874b, 1875).

Nausithoepunctata Kolliker, 1853

(Fig. 6)
Fig. 6. Nausithoe punctata Kolliker, 1853, after Mayer (1910b),
Nausithoe punctata Kolliker, 1853:323. Sterrer (1986), and Mianzan and Cornelius (1999).

Common Name
None known.
Type Locality
In Messina, infrequent ("In Messina ziemlich selten")
(Kolliker, 1853:323).
Museum Material
North Carolina, Atlantic Ocean east of Cape Hatteras,
35°27' N, 73°14'W, 100-0 m, Grampus (Bache) Station
10161, 28 January 1914, one medusa, det. H. B. Bigelow,
USNM 41722.
Specific Characters
Medusae small with umbrella flatter than a hemisphere,
up to 1.5 cm in diameter; central disk with thick me-
soglea, without radiating furrows; exumbrellar surface
with numerous fine nematocyst warts; coronal groove
lobed; periphery of umbrella with 16 deep radial grooves
extending to lappets; lappets linguiform, with rhopalar
and tentacular clefts nearly equal in depth; stomach
short, with four lips around mouth; gastric cirri in four
interradial groups; gonads large, globular, eight, adradial,
equally spaced (Mayer, 1910b; Kramp, 1961; Mianzan and
Cornelius, 1999).
Remarks
Originally described from the Mediterranean by Kollik
er (1853), the medusa of Nausithoe punctata was described
by Mayer (1910b) as a surface form that is common "in
all tropical or warm seas." He noted that there were no
records of it on the east coast of the United States north
of the Carolinas, although it was abundant at theTortugas
and Bahamas during summer.The specimen collected off
Cape Hatteras by the Bache (USNM 41722), from a tow
made between 100 m and the surface, was identified by
H. B. Bigelow and reexamined here. Although a neritic
species (Bigelow, 1938), medusae of N punctata are most
likely to occur at the seaward limits of the study area. It is
not known if the full life cycle is completed in the study
area, or if medusae are carried into the region by the Gulf
Stream from areas to the south.
The life cycle of Nausithoe punctata has long been
known to be metagenetic, having both a benthic scyph
istoma and a planktonic medusa stage.The names Stepha
noscyphus mirabilis Allman, 1874a and Spongicola fstularis
Schulze, 1877, applied to the scyphistoma, are junior sub
jective synonyms of N punctata Kolliker, 1853 (Lo Bianco
and P.Mayer, 1890; Mayer, 1910b;Jarms, 1997).

Nematocyst Complement Reported Range

Undescribed. Study area: east of Cape Hatteras (this record).
 Overall: circumglobal in warm waters; surface and
shallow-water (Mayer, 1910b; Kramp, 1961; Russell,
1970; Burke, 1975; Mianzan and Cornelius, 1999).
Family Periphyllidae Haeckel, 1880
Genus Periphylla F. Miiller, 1861
Periphylla E Miiller, 1861:311.
Diagnosis
Medusa with cone-shaped to dome-shaped central disk
above a deep coronal furrow on exumbrella; mesoglea
thick, rigid; exumbrella peripheral to coronal groove
with 16 radial furrows, extending to lappets; rhopalia
four, interradial; tentacles 12, in four groups ofthree each;
marginal lappets 16, linguiform; gonads eight, U-shaped,
adradial (Mayer, 1910b; Kramp, 1961; Russell, 1970;
Mianzan and Cornelius, 1999).
Type Species
Carybdea periphylla Peron and Lesueur, 1810, by
monotypy.
Remarks
There has been longstanding confusion over authorship
and date of the familiar and widely used generic name
Periphylla.Early authors,such asHaeckel (1880) and May
er (1910b), attributed the name to J. Steenstrup and to
his "Acta et Catalogus Musei Hafniensis," dated as 1837
and 1842. In a detailed review of questions surrounding
the nomenclature of the genus, however, Kramp (1947)
discovered that this work by Steenstrup was never pub
lished. He therefore concluded that the name Periphylla
was first made available by Haeckel (1880). Earlier, Neave
(1940a) had cited both F. Miiller (1861) and Haeckel
(1880) as authors of the name. Indeed, E Miiller (1861) is
confirmed here as having made the name Periphylla avail
able (Art. 12.2.5, ICZN) before Haeckel (1880), with
Carybdea periphylla Peron and Lesueur, 1810 as its type
species by monotypy.
Recognizing F. Miiller (1861) as author of Per
iphylla instead of Haeckel (1880) removes a threat to this
well-known name from its subjective synonym Quoyia
L. Agassiz, 1862, a seldom-used name established for
Carybdea bicolor Quoy and Gaimard, 1833. The latter is
now considered a junior subjective synonym of Carybdea
periphylla Peron and Lesueur, 1810 (Kramp, 1961;Russell,
1970; Morandini, 2003), the type species of Periphylla.
Questions have also arisen about Dodecabostrycha Brandt,
1837 as a possible threat to Periphylla. Kramp (1947)
showed that the type species of that genus is Chrysaora
gaudichaudii Lesson, 1830 (= Desmonema gaudichaudii),
now classified as a semaeostome (Kramp, 1961; Corne
lius, 1997). Thus, Dodecabostrycha and Periphylla are not
congeneric.
An additional nomenclatural problem remains be
cause Periphylla is a junior subjective synonym of the
virtually unknown Cyclophylla Brandt, 1837. That name
was established by Brandt as a subgenus to accommodate
Carybdea periphylla and C. bicolor. Kramp (1947) recog
nized the unfortunate synonymy between the two ge
nus-group names and stated "It seems...most deplorable
if Periphylla should be replaced by a name [Cyclophylla],
which has appeared only once in the literature and never
revived...and I most entreatingly propose that the famil
iar name Periphylla be retained for this genus." No action
has ever been undertaken to have the name conserved,
and nomenclatural stability remains threatened. Under
the current code (International Commission on Zoo
logical Nomenclature, 1999), Reversal of Precedence
(Art. 23.9, ICZN) can be applied in this case, with the
junior subjective synonym Periphylla being designated
as both valid and a nomen protectum. The older name
Cyclophylla thereby becomes a nomen oblitum. Firstly,
Cyclophylla has not been used as a valid name in zool
ogy since before 1899 (Art. 23.9.1.1). Secondly, Periphylla
has been used in at least 25 works by more than 10 au
thors in the past 50 years (Art. 23.9.1.2) (Kramp, 1961,
1968a; Naumov, 1961; Russell, 1970; Larson, 1976a; van
der Spoel, 1987; Larson et al., 1991; Fossa, 1992; Pages
et al., 1992; Franc, 1995a; Arai, 1997; Cornelius, 1997;
Sheiko and Stepanjants, 1997; Phleger et al., 1998; Eiane
and Aksnes, 1999; Jarms et al., 1999; Gorsky et al., 2000;
Mills, 2001; Shimomura et al, 2001; Jarms et al., 2002;
Bamstedt et al., 2003; Morandini, 2003; Dawson, 2004;
Herring and Widder, 2004; Lindsay et al., 2004). Thus,
the well-known name Periphylla is assigned precedence
over Cyclophylla under current provisions of the code,
and is maintained as the valid name for this widespread
genus.
Kramp (1961), Russell (1970), and others have treat
ed this genus as monotypic. It differs from the morpho
logically similar Paraphyllina Maas, 1903 in having inter
radial rather than perradial sense organs (Russell, 1970;
Mianzan and Cornelius, 1999).
 Periphylla periphylla (Peron and Lesueur, 1810)
(Fig. 7)
Carybdea periphylla Peron and Lesueur, 1810:332.
Periphylla hyacinthina — Fewkes, 1886:933.
Common Name
Merchant-cap.
Type Locality
In equatorial Atlantic Ocean ("..l'Ocean Atlantique
equatorial") (Peron and Lesueur, 1810).
Museum Material
Florida, continental slope east of Flagler Beach, R/V
Oregon, 29°34' N, 80°00' W, 6 October 1953, 1518-
1701 m,two medusae, USNM 58477. Florida, continental
slope east of St. Augustine, R/V Silver Bay, 29°56' N,
80°05' W, 29 April 1961, 375 m, five medusae, USNM
58722.
Fig. 7. Periphylla periphylla (Peron and Lesueur, 1810), after
Haeckel (1881b) and Mayer (1910b).
Specific Characters
Medusae up to 25 cm in umbrella diameter, usually as
high or higher than wide, with cone- to dome-shaped
central disk; exumbrella smooth; manubrium large,
not extending beyond umbrella margin (Kramp, 1961;
Russell, 1970; Mianzan and Cornelius, 1999).
Nematocyst Complement
Medusa (Lurefjorden, Norway; Jarms et al., 2002):
Tentacles, lappets, mouth lips, gastric filaments,
exumbrella—
holotrichous isorhizas (11.3-15.6 |am long
x 7.3-9.8 um wide)
small holotrichous isorhizas (7.9 um long
x 6.1 jam wide)
round heterotrichous microbasic euryteles
(17.6-21.8 jam long x 13-15 um wide)
long ellipsoid heterotrichous microbasic euryteles
(16-62 jam long x 13-16 jam wide)
giant ellipsoid heterotrichous microbasic euryteles
(38-100 jam long x 24-27 jam wide)
short ellipsoid heterotrichous microbasic euryteles
(21.8-25.8 jam long x 16.3-18.5 jam wide)
Remarks
Synonymy lists for Periphylla periphylla exist in works
such as Russell (1970) and Morandini (2003), and a de
tailed review of the nomenclatural history of this me
dusa was given by Kramp (1947).The binomen Periphylla
hyacinthina had often been applied to this species during
the first half of the 20th century, with J. Steenstrup being
credited as author (Mayer, 1910b). Kramp (1947) discov
ered that the supposed publication by Steenstrup ("Acta
et Catalogus Musei Hafniensis,') containing the specific
name was never printed. Authorship of P. hyacinthina was
traced by Kramp even earlier to Faber (1829), but he
nevertheless demonstrated that the oldest name for the
species was Carybdea periphylla Peron and Lesueur, 1810.
That specific name has now gained almost universal use,
in combination with the generic name Periphylla (see
comments under the genus Periphylla above).
Details of the anatomy, morphology, and coloura
tion of Periphylla periphylla were given by Russell (1970).
Considerable research interest has focused recently on
huge populations of this coronate medusa in certain
Norwegian fjords (Fossa, 1992;Jarms et al., 1999; Gorsky
et al., 2000;Youngbluth and Bamstedt, 2001; Jarms et al.,
2002; Bamstedt et al., 2003).The life cycle was described
for the first time by Jarms et al. (1999) from that environ
ment. Periphylla periphylla was found to be holopelagic,
23
 with developmentprogressing directly from an unusually
large egg (about 1.5 mm in diameter) to a spherical em
bryo to a young medusa.There was no planula,no sessile
polypoid stage, and no typical ephyra stage in the life
cycle. Populations were not seasonal in occurrence.Jarms
et al. (1999) hypothesized that medusae of this species
might be unusually long-lived, perhaps capable ofsurviv
ing for several decades, and that mortality in the Luref-
jorden where their study was carried out was low.Major
prey items in the fjord were two species of copepods
(Calanus finmarchicus, Euchaeta norvegica), although a total
of 39 prey species were found (Fossa, 1992). Periphylla
periphylla is known to be bioluminescent (Nicol, 1958).
The cnidome of Periphylla periphylla was described
in populations from Norway by Jarms et al. (2002). Two
categories of nematocysts (holotrichous isorhizas and
heterotrichous microbasic euryteles) were described and
illustrated, both of them occurring in more than one size
class and shape. Those described as "giant ellipsoid het
erotrichous microbasic euryteles" are the largest nema
tocysts known to date from the Scyphozoa, reaching to
100 jamin length.They were present in small numbers on
the tentacles and exumbrella. Much more abundant on
these parts of the medusa were long ellipsoid euryteles.
Both kinds of euryteles were shown to become progres
sively larger as the medusae grew. Little or no change in
size with medusa growth was noted in the other nema
tocyst types.
Periphylla periphylla was first reported from the study
area by Fewkes (1886), and specimens from the region
exist in collections at the NMNH. Bigelow (1928) re
ported this species (as P.hyacinthina) from the vicinity of
the study area (100 miles SE of Cape Hatteras, 34°47' N,
73°41'W) in collections of the Arcturus Oceanographic
Expedition. Elsewhere, P periphylla appears to be a wide
spread mesopelagic and bathypelagic medusa, having
been reported from the Atlantic, Pacific, and Southern
Oceans (Bigelow, 1938; Kramp, 1947, 1961; Russell,
1970; van der Spoel, 1987; Mianzan and Cornelius, 1999;
Morandini, 2003).
Reported Range
Study area: Gulf Stream off Cape Hatteras, NC,
35°12.17' N, 74°57.25'W, surface, Albatross Station
2110 (Fewkes, 1886, as Periphylla hyacinthina).
Overall: reported to occur in all oceans except
the Arctic; predominantly deep-sea and consid
ered predominantly mesopelagic (Kramp, 1947;
van der Spoel, 1987; Mianzan and Cornelius, 1999;
Morandini, 2003).
24
Subclass Discomedusae Haeckel, 1880
Order Semaeostomeae L. Agassiz, 1862
Family Pelagiidae Gegenbaur, 1857
Genus Chrysaora Peron and Lesueur, 1810
Chrysaora Peron and Lesueur, 1810:364.
Diagnosis
Medusa with dome-shaped umbrella; exumbrella with
out conspicuous warts; manubrium large; oral arms four,
long, frilly, simple, surrounding a mouth; rhopalia eight;
exumbrellar sensory pits deep; tentacles three or more
per octant (but one per octant in postephyrae), arising
from clefts between lappets on umbrella margin; mar
ginal lappets 32 or more; stomach pouches 16, with
eight rhopalar pouches narrower than eight tentacular
ones; septa terminating closer to tentacles than to rhopa
lia; gonads four, much folded, interradial (Mayer, 1910b;
Kramp, 1961; Russell, 1970; Gershwin and Collins, 2002;
Morandini, Silveira and Cornelius, 2006).
Type Species
Chrysaora mediterranea Peron and Lesueur, 1810, a junior
subjective synonym of Medusa hysoscella Linnaeus, 1767,
by subsequent designation by Mayer (1910b).
Remarks
Louis Agassiz (1862:125) stated that the type species of
Chrysaora Peron and Lesueur, 1810 was Medusa hysoscel
la Linnaeus, 1767, but it was not one of the originally
included nominal species. Therefore, that designation,
followed by Kramp (1961), is nomenclaturally invalid
(ICZN, Art. 67.2). Chrysaora mediterranea Peron and
Lesueur, 1810, one of the nominal species originally
included in Chrysaora, was validly designated as its type
species by Mayer (1910b). Chrysaora mediterranea is now
considered a junior subjective synonym of C. hysoscella
(Kramp, 1961; Russell, 1970). A revision of the genus is
currently in preparation (Andre Morandini and Antonio
Carlos Marques, pers. comm., 2007).
Chrysaora quinquecirrha (Desor, 1848)
(Fig. 8)
Pelagia quinquecirrha Desor, 1848:76.
Dactylometra quinquecirrha — Brooks, 1883:137.
— Mayer, 1910b:585, pi. 64A.
Chrysaora quinquecirrha — Calder, 1972a:41. — Hoese,
1973:74. — Schwartz and Chestnut, 1973:60. —
Miller, 1974:5. — Kraeuter and Setzler, 1975:69.
 — Calder and Hester, 1978:88. — Rountree,
1983a:25.

Common Names
Sea nettle; stinging nettle.

Type Locality
USA, Massachusetts, Nantucket Bay (Desor, 1848:76).

Museum Material
South Carolina,Town Creek andjones Creek intersection,
33°19.9' N, 79°10.2' W, 25.8 °C, 35%o, hand collecting,
22 July 1970, one medusa, GMBL 70-134. South Carolina,
off Moms Island, 32°43.3' N, 79°54.1' W, 28.16 °C,
31.14%o, otter trawl, 24 September 1970, one medusa,
GMBL 70-182. South Carolina, Isle of Palms near De-
wees Inlet, 32°49' N, 79°43' W, 0-1 m, 28 °C, 35%.,
dipnetted, 01 August 1972, one medusa, GMBL 72-182.
South Carolina, Inlet Creek, 32°46.9' N, 79°49.5' W, 4 m,
28 °C, 31%o, 20-ft otter trawl, 28 September 1972, one
medusa, GMBL 72-373. Florida, continental shelf east of
Jacksonville, R/V Dolphin, MARMAP coll. no. 0575242,
30°16.5' N,80°58.6.0'W,24 m, 26 °C, 36.3%., 3A Yankee
No. 36 trawl, 11 September 1975, three fragmentary me
dusae, ROMIZ B3591. Georgia, continental shelf east of
Cumberland Island, R/V Dolphin, MARMAP coll. no.
0575056,30°59.5' N,80°52.0'W,24 m, 14.8 °C,%Yankee
No. 36 trawl, 27 January 1975, seven fragmentary medu
sae, ROMIZ B3588. South Carolina, Charleston Harbor
off Fort Sumter, R/V Anita, 32°45.5' N, 79°52.9' W, 3-
9 m,21 °C, 20-ft otter trawl, 27 October 1979, one medu
sa, GMBL 79-176. South Carolina, Charleston Harbor,off
James IslandYacht Club, 32°45.5' N, 79°55.0'W, 7 m, 28 °C,
24%o,otter trawl, 13 June 1981, four medusae, GMBL 81-
34. South Carolina, Charleston Harbor, Anchorage, R/V
Anita, 32°45.91' N, 79°53.35' W, 28.3 °C, 27.4%., otter
trawl, 10 August 2004, one medusa, SERTC No. S2186.
Georgia, Ogeechee River, 31°50.92' N, 81°04.72' W,
Fig. 8. Chrysaora quinquecirrha (Desor, 1848), from a photo
5.3 m, 26.29 °C, 24.48%.., bottom sled, 10 June 1998,
graph by R.J. Larson.
four medusae, SERTC No. S2190. Georgia, Ogeechee
River, 31°51.00' N, 81°04.62' W, 0.5 m, 28.30 °C,
23.90%o, neuston net, 14June 1998,12 medusae, SERTC 1 September 2005, two medusae, ROMIZ B3607. South
No. S2188. Georgia, Ogeechee River, 31°51.00' N, Carolina, Inlet Creek, R/V Silver Crescent, 32°46.65' N,
81°04.62' W, 0.5 m, 28.24 °C, 25.89%,, neuston net, 79°49.19' W, surface, 28.2 °C, 31.6%o, neuston net,
14 June 1998, eight medusae, SERTC No. S2189. 1 September 2005, 10 medusae, ROMIZ B3615. South
South Carolina, Stono River, Bird Key, 32°38.299' N, Carolina, Inlet Creek, R/V Silver Crescent, 32°46.65' N,
79°59.095' W, 5 m, dip net, 23 August 2005, five me 79°49.19'W, surface, 28.2 °C, 31.6%., neuston net, 1 Sep
dusae, SERTC No. S2111. South Carolina, seaward of tember 2005, two postephyrae, ROMIZ B3618. South
buoy 17 off Charleston jetties, R/V Silver Crescent, Carolina, off jetties at entrance of Charleston Harbor,
32°42.31' N, 79°47.77' W, 8 m, 31.6%.., 28.7 °C, trawl, R/V Silver Crescent, 32°43.02' N, 79°48.56'W, surface,

25
28.7 °C, 31.6%o, neuston net, 1 September 2005, one
medusa, SERTC No. S2109. South Carolina, off jetties
at entrance of Charleston Harbor, R/V Silver Crescent,
32°42.31» N, 79°47.77' W, 8 m, 28.7 °C, 31.6%o, 20-ft
otter trawl, 1 September 2005, three medusae, SERTC
No.S2108.
Specific Characters
Medusa up to 25 cm in umbrella diameter; exumbrella
with small wart-like clusters of nematocysts; tentacular
lappets U-shaped; abrhopalar clefts of rhopalar lappets
mere notches; septa between gastric pouches slightly
curved over adaxial 2/3 of their length; tentacles usually
3-5 per octant, but as many as nine reported; colour var
ied, from entirely whitish to variously pigmented, with
red or brownish patterns on exumbrella and along oral
arms (Mayer, 1910b; Kramp, 1961; Calder, 1972a; Mian
zan and Cornelius, 1999; Gershwin and Collins, 2002).
Nematocyst Complement
Medusa (Virginia, USA; Calder, unpublished data):
Tentacles, exumbrella—
holotrichous a-isorhizas (7.6-9.9 urn long
x 4.0-5.0 um wide)
holotrichous A-isorhizas (19.4-28.9 um long
x 13.3-18.8 jam wide)
heterotrichous anisorhizas (15.8-25.3 jam long
x 13.9-22.8 jam wide)
heterotrichous microbasic euryteles
(12.7-16.5 jamlong x 7.7-9.7 um wide)
Remarks
The sea nettle (Chrysaora quinquecirrha) is perhaps the sec
ond most familiar jellyfish in the southeastern United
States, after the abundant cannonball jelly (Stomolophus
meleagris). A well-known stinger, it ranges from euhaline
coastal waters far up into mesohaline regions of estuaries.
In Chesapeake Bay, the scyphistomae of C. quinquecirrha
are most prevalent in mesohaline waters of creeks and
tributaries,and ephyrae and young medusae appear there
before being observed in the bay proper (Cargo and
Schultz, 1966,1967). Similar distribution patterns might
be expected to occur in the study area,although no com
parable studies have yet been undertaken.
The life cycle of Chrysaora quinquecirrha is known
to be metagenetic, passing through several stages from
medusa to zygote to planula to scyphistoma to strobila
to ephyra and back to medusa (Littleford, 1939; Cargo
and Schultz, 1966, 1967; Calder, 1972c, 1974a). Podo-
cyst formation by scyphistomae is one of the means by
26
which asexual reproduction occurs, and these cysts are
also resistant to adverse environmental conditions. Me
dusae are almost always dioecious, with the ripe gonads
of the male being pink and those of the female yellowish
to ashy gray or grayish brown (Mayer, 1910b; Littleford,
1939). Growth is rapid and maturity is reached within
two months under favorable conditions (Calder, 1972c).
While the scyphistoma stage is potentially perennial, me
dusae are of seasonal occurrence in the study area, and
the life span of a given individual appears to be no more
than a few months. Medusae have been reported in the
region every month from April to October (Hoese, 1973;
Schwartz and Chestnut, 1973; Miller, 1974; Kraeuter and
Setzler, 1975; Rountree, 1983a). Material examined here
included fragmentary specimens from shelf waters east
of Cumberland Island, Georgia, collected in late January
(ROMIZ B3588).
The medusa of this species has been reported to pass
through stages having as few as one tentacle and two lap
pets per octant to as many as nine tentacles and 10 lappets
per octant during growth (Calder, 1972a). Maturity is
commonly reached at the 24-tentacle stage.This supports
the contention of Kramp (1955, 1961) that Dactylometra
L. Agassiz, 1862 and Kuragea Kishinouye, 1902, distin
guished on the basis of numbers of tentacles and lap
pets, merely represent developmental stages of Chrysaora
Peron and Lesueur, 1810 and are junior synonyms ofit. A
preliminary phylogenetic analysis of the Pelagiidae sug
gested that specieswith 40 tentacles (fiveper octant) may
form a separate group, however, and more study is clearly
needed to establish whether Dactylometra may be valid
(Gershwin and Collins, 2002). Earlier, Thiel (1978) had
argued for recognition of Dactylometra as distinct from
Chrysaora, based on supposed reproductive differences.
Stages in the life cycle of Chrysaora quinquecirrha,
as described by Littleford (1939), Cones (1969), Calder
(1972a), Morales-Alamo and Haven (1974), and Cargo
and Rabenold (1980), resemble those of Chrysaora lactea,
described by Morandini et al. (2004).While medusae of
the two species are much alike, morphological charac
ters distinguishing them include the following: (1) the
abrhopalar clefts of the rhopalar lappets are deep in
C. lactea but mere notches in C. quinquecirrha (Mayer,
1910b; Kramp, 1961; Mianzan and Cornelius, 1999);
(2) septabetween gastricpouches are slighdy curved over
the adaxial 2/3 of their length in C. quinquecirrha, but
straight in C. lactea (Mianzan and Cornelius, 1999; Ger
shwin and Collins, 2002); (3) the exumbrella of C. quin
quecirrha is covered with wart-like clusters of nematocysts
while that of C. lactea is densely covered with minute
truncate papillae (Mayer, 1910b;Mianzan and Cornelius,
1999); and (4) the peripheral pair of tentacles in each
quadrant are tertiaries in C. quinquecirrha and secondaries
in C. lactea (Morandini, Silveira, and Cornelius, 2006).
While medusa size is highly varied, umbrella diameters of
fully-grown C.quinquecirrha tend to be largerthan those of
C. lactea (Mayer, 1910b; Kramp, 1961; Gershwin and
Collins, 2002).
Medusae assigned to Chrysaora quinquecirrha occur in
several colour varieties (Mayer, 1910b:586-588). Some
are essentially milky whitish, while others have conspicu
ous red or brown patterns of pigmentation on the um
brella and oral arms. Pigmentation intensity and patterns
vary widely, as shown in colour illustrations of the spe
cies by Mayer (1910b). Earlier, Mayer (1898) reported
that the colour of pigmented medusae faded if they were
kept in an aquarium for a few days.He found that when
maintained in an aquarium without food for about six
weeks, the umbrella shrank and all colour faded away.
The different colour forms have been treated by Mayer
(1910b) and subsequent authors as variants of the same
species. However, Papenfuss (1936) reported differences
in nematocyst size in the two forms and assigned the
milky-white estuarine form to "Dactylometra quinquecir
rha var. chesapeakeiV Calton and Burnett (1978) reported
differences in the effects of venom from the two forms
on mice, with the white phase causing more pain than
the red.The possibility that cryptic species may be repre
sented in medusae identified as C. quinquecirrha from the
east coast of the USA needs to be explored.
Chrysaora quinquecirrha has been little studied in the
southeastern United States, but it has been the subject of
extensive research elsewhere, particularly in Chesapeake
Bay to the north of the study area. Pioneering investiga
tions on the biology of C. quinquecirrha were undertaken
in the Chesapeake system, where it is normally abundant
each summer (Littleford andTruitt, 1937; Littleford, 1939;
Truitt, 1939; Cargo and Schultz, 1966, 1967; Cones and
Haven, 1969). Mansueti (1963) investigated associations
of these medusae with small fishes in the bay, as well as
fish-medusa associations elsewhere in the world. Several
authors investigated the natural history and ecology of
C. quinquecirrha in Chesapeake Bay (Gatz et al., 1973
Cargo, 1974, 1977, 1979; Baird and Ulanowicz, 1989
Cargo and King, 1990; Purcell, 2005; Purcell et al., 1999
Keister et al., 2000; Condon et al, 2001). Other stud
ies of the life cycle of the sea nettle were carried out
(Cones, 1969; Calder, 1972a, 1974a; Morales-Alamo and
Haven, 1974; Cargo and Rabenold, 1980). Many bio
chemical, physiological, and ultrastructural studies have
been undertaken on the species (Perkins et al., 1970;
Burnett and Gould, 1971; Black, 1972, 1981; Blanquet,
1972a, b; Crawford and Webb, 1972; Loeb, 1972, 1973,
1974a, b; Mangum et al., 1972;Webb et al., 1972; Black
and Webb, 1973; Lin and Zubkoff, 1973, 1976a, b, 1977;
Bynum and Black, 1974; Loeb and Gordon, 1975; Black
et al, 1976; Gardner and Zubkoff, 1978; Joseph, 1979;
Loeb and Hayes, 1981; Quensen et al, 1981; Black
and Riley, 1985; Wright and Purcell, 1997). Numerous
studies have been published on food, feeding behavior,
and food chains in relation to sea nettles (Herman et
al., 1968; Miller and Williams, 1972; Loeb and Blanquet,
1973; Clifford and Cargo, 1978; Feigenbaum and Kelly,
1984; Purcell et al., 1991; Cowan and Houde, 1992,1993;
Purcell, 1992; Nemazie et al., 1993; Purcell, Nemazie et
al, 1994; Purcell, White et al., 1994; Purcell and Cowan,
1995; Olesen et al., 1996; Ford et al., 1997; Kreps et al.,
1997; RiUing and Houde, 1999; Matanoski et al., 2001;
Ma and Purcell, 2005). Indeed, medusae of C. quinquecirrha
were considered keystone predators in the Chesapeake
Bay because of their substantial effects on food webs in
the plankton (Purcell and Decker, 2005). Investigations
on nematocysts of the species include those of Papenfuss
(1936), Sutton and Burnett (1969), and Calder (1971,
1977). Finally, a massive literature on chemistry, toxicol
ogy and medical aspects ofjellyfish stings, some of it un
dertaken on C. quinquecirrha from the Chesapeake Bay,
has been reviewed in works such as those of Burnett
and Calton (1977, 1987), Williamson et al. (1996), and
Burnett (2001).
As one of the more frequently occurring jellyfishes
in the region, Chrysaora quinquecirrha must be considered
a significant predator in coastal planktonic systems of the
southeast. These medusae are known to be important in
overall plankton dynamics in tributaries of the Chesa
peake Bay (Purcell, 1992). Purcell reported that prey of
the species was varied, but consisted primarily of cope-
pods and copepod larvae. Other prey included cteno-
phores, fish eggs and larvae, and polychaetes. Rotifers,
cladocerans, barnacle larvae, and bivalve larvae were also
occasional food items. By feeding heavily on the abun
dant ctenophore Mnemiopsis leidyi, sea nettles may reduce
mortalities in zooplankton and ichthyoplankton in Ches
apeake Bay (Purcell and Cowan, 1995). In turn, medusae
of C. quinquecirrha are preyed upon by various species
including spider crabs (Libinia dubia), harvestfish (Pepri-
lus alepidotus), and orange filefish (Alutera schoepfi) (Cargo
and Schultz, 1966; Calder, 1972c). Certain nudibranchs
(Coryphella sp.) feed on the scyphistoma stages (Cargo
and Schultz, 1967).
27
 Variations in the abundance of Chrysaora quinquecir
rha from year to year in the Chesapeake estuary that cor
related with salinity and temperature were documented
by Purcell et al. (1999). From studies in mesocosms, me
dusae of this species appear to be negatively phototactic
(Schuyler and Sullivan, 1997).
The cnidome differs from one stage to another in
the life cycle of Chrysaora quinquecirrha. Only two nema
tocyst categories are known to occur in the scyphistoma,
holotrichous a-isorhizas and heterotrichous microbasic
euryteles (Calder, 1971). Heterotrichous anisorhizas and
holotrichous A-isorhizas appear during the strobilation
process and, together with a-isorhizas and euryteles, form
a part of the nematocyst complement in both ephyrae
and medusae (Calder, 1977). Although the cnidome of
ephyrae and medusae is the same, the sizes of all the
nematocyst types are much larger in the medusa (Calder,
1977; this study).
Chrysaora quinquecirrha is known to range from
southern New England to the Caribbean Sea in the
western Atlantic (Kramp, 1961). According to Mian
zan and Cornelius (1999), no valid records exist of
C. quinquecirrha from south of the equator in South
America. Moreover, they maintained that reports of the
species from the southeastern Atlantic, off the west coast
of Africa (Kramp, 1961), require validation. Although I
have examined medusae from the western Pacific (Ma
laysia, ROMIZ B2821) that resembled descriptions of
C. quinquecirrha, records of this species outside the Atlan
tic Ocean (Kramp, 1961) are considered doubtful. The
specimen from Malaysia at the ROM is likely referable
instead to Chrysaora chinensis Vanhoffen, 1888 (Antonio
Carlos Marques, pers. comm., 2007).
Reported Range
Study area: common in baysand shelfwatersfrom spring
through autumn across the entire region (Mayer,
1910b; Calder, 1972a; Schwartz and Chestnut, 1973;
Hoese, 1973; Miller, 1974; Kraeuter and Setzler,
1975; Calder and Hester, 1978; Rountree, 1983a).
Overall: New England to the Caribbean Sea and Gulf
of Mexico; neritic (Hedgpeth, 1954; Kramp, 1961;
Burke, 1975; Larson, 1986).
28
Genus Pelagia Peron and Lesueur, 1810
Pelagia Peron and Lesueur, 1810:349.
Diagnosis
Medusa with dome-shaped umbrella; exumbrella with
conspicuous warts in the form of elevated mesogleal
mounds; manubrium large; oral arms four, long, frilly,
simple, surrounding a mouth; rhopalia eight; exumbrellar
sensory pits shallow; tentacles eight, alternating with rho
palia and arising from umbrella margin; marginal lappets
16; stomach pouches 16, all alike; septa terminating mid
way between tentacles and rhopalia; gonads four, much
folded, interradial (Mayer, 1910b; Kramp, 1961; Russell,
1970; Gershwin and Collins, 2002).
Type Species
Medusa noctiluca Forsskal, 1775, by subsequent designa
tion by Mayer (1910b:570).
Pelagia noctiluca (Forsskal, 1775)
(Fig. 9)
Medusa noctiluca Forsskal, 1775:109.
Pelagia noctiluca — Kraeuter and Setzler, 1975:68.
— Calder and Hester, 1978:88.
Pelagia cyanella — Hoese, 1973:94.
Common Names
Oceanic jelly; purple jelly; mauve stinger; warty jellyfish.
Type Locality
Mediterranean Sea ("Mari Mediterraneo") (Forsskal, 1775).
Museum Material
Florida, continental slope east of Daytona Beach, R/V
Silver Bay, 29°25' N, 79°51' W, 3 October 1961, 732-
549 m, five medusae, USNM 54369. Florida, continental
shelf east of Ponce de Leon Inlet, R/V Dolphin,
MARMAP coU. no. 0576061, 29°00.5' N, 80°04.0' W,
154 m, Va Yankee No. 36 trawl, 31 January 1976, one
medusa ROMIZ B3586. Georgia, continental shelf east
ofOssabaw Island, R/V Dolphin, 31°46.3* N, 79°12.8'W,
22 °C, 36%o, surface, dipnet, 20 March 1976, seven
medusae, GMBL 76-13. South Carolina, continental
shelf east of Charleston, R/V Dolphin, MARMAP coll.
no. 0576325, 32°50.9' N, 78°14.6' W, 104 m %Yankee
No. 36 trawl, 29 August 1976, one fragmentary medusa,
ROMIZ B3593. Florida, continental shelf east of Jack
sonville, R/V Dolphin, MARMAP coll. no. 0576378,
30°10.0'N,80°35.5'W,34m,%YankeeNo.36trawl,17Sep-
tember 1976, 16 fragmentary medusae, ROMIZ B3594.

Specific Characters
Medusa up to 6.5 cm in umbrella diameter; exumbrella
with numerous wart-like clusters of nematocysts; tenta
cles one per octant; colour varied, often purplish (Kramp,
1961; Mianzan and Cornelius, 1999).

Nematocyst Complement
Medusa (Mediterranean Sea; Avian et al., 1991):
Tentacles, oral arms—
holotrichous a-isorhizas (4.35—7.88 um long
x 3.31 —5.05 um wide)
holotrichous A-isorhizas (12.19—21.65 urn long
x 7.38—11.75 um wide)
heterotrichous anisorhizas (15.20—30.14 um long
x 13.87-28.78 um wide)
heterotrichous microbasic euryteles
(10.11-16.87 um long x 8.17-12.98 um wide)
undetermined nematocyst category
(no measurements given)

Remarks
Pelagia noctiluca is one of the few purely oceanic species
of Discomedusae (Semaeostomeae and Rhizostomeae).
A benthic scyphistoma stage in the life cycle is bypassed
entirely, with the planula larva developing directly into
an ephyra, and the ephyra to a medusa. A detailed re
view of the ontogeny of P. noctiluca was given by Russell
(1970).This species is common offshore in and near the
Gulf Stream, and it is sometimes carried inshore by ed
dies. Specimens reported from the coast of Georgia by
Kraeuter and Setzler (1975) were thought to have been
transported shorewards from the Gulf Stream.
I have persisted here in using the name heterotric
hous anisorhizas for the nearly spherical haplonemes of
scyphozoans such as Pelagia noctiluca, Chrysaora quinquecir
rha, and Cyanea capillata. These nematocysts were called
holotrichous isorhizas by Weill (1934b), and O-isorhizas
by Ostman and Hydman (1997).The superbmicrographs
of Ostman and Hydman show a distinct tapering of the
tubule distally, with the spines more developed proxi-
mally. In my opinion, these characters conform with the
diagnosis of heterotrichous anisorhizas. The so-called
a-isorhizas and A-isorhizas may also be more accurately
called heterotrichous anisorhizas, but the taper of the tu
bule is much less distinct and the long-established name
for them is retained for the present. I am also uncon
vinced that so-called "birhopaloids" and "trirhopaloids"
Fig. 9. Pelagia noctiluca (Forsskal, 1775), after Mayer (1910b)
and Stephens and Calder (2006).
of cubozoans and scyphozoans are anything more than
varieties of heterotrichous microbasic euryteles.
Kramp (1961), Russell (1970), and others have been
followed here in regarding this oceanic medusa as cir-
cumglobal in warm and temperate waters. However, the
taxonomy of these medusae needs re-examination given
the broad questioning now underway about the likeli
hood of extensive crypsis in Scyphozoa.
Larson (1987b) reported that medusae of Pelagia noc
tiluca from inshore waters of Puerto Rico appeared to be
opportunistic predators. A wide variety of zooplankton
prey, including hydromedusae, siphonophores, cteno-
phores, polychaetes, copepods, ostracods, cumaceans, iso-
pods, amphipods, chaetognaths, and fish eggs, were found
in the gastrovascular cavities of field-collected medusae.
In turn, these medusae are an important item in the diet
of both North Atlantic loggerhead (Caretta caretta) and

29
leatherback (Dermochelys coriacea) sea turtles (Lutz and
Musick, 1996).
Cyclic blooms of this species have been documented
in the western Mediterranean region at approximate in
tervals of 12 years (Goy et al., 1989). By contrast, massive
blooms have occurred at irregular intervals in the Adri
atic Sea, raising concerns over the causes as well as the
explosions (Mills, 2001).
Medusae of Pelagia noctiluca are venomous to hu
mans (Williamson et al., 1996; Humann, 2002).
Reported Range
Study area: Gulf Stream at the eastern margin of the
study area; off Sapelo Island, GA (Hoese, 1973;
Kraeuter and Setzler, 1975).
Overall: carried northwards from the tropical and subtropi
cal Atlantic to Europe in the Gulf Stream and North
Atlantic Drift; Gulf of Mexico; warm Pacific and
Indian oceans; pelagic (Kramp, 1961; Russell, 1970;
Burke, 1975,1976; Mianzan and Cornelius, 1999).
Family Cyaneidae L. Agassiz, 1862
Genus Cyanea Peron and Lesueur, 1810
Cyanea Peron and Lesueur, 1810:363.
Diagnosis
Medusa with flattened umbrella; oral arms four, sur
rounding a mouth; rhopalia eight; tentacles numerous,
arising from subumbrella proximal to umbrella margin,
occurring in several rows within eight adradial clusters;
stomach pouches terminating in numerous branching,
blind canals within the marginal lappets; gonads four,
much folded, interradial; subumbrella with both circu
lar and radial muscle bands (Mayer, 1910b; Kramp, 1961;
Russeil, 1970).
Type Species
Cyanea arctica Peron and Lesueur, 1810, ajunior synonym
of Medusa capillata Linnaeus, 1758, designated herein.
Remarks
The name Cyanea was applied earlier to a dipteran by
Meigen (1800), but that work was suppressed for nomen
clatural purposes by the International Commission on
Zoological Nomenclature (1963). Cyanea Peron and Le
sueur, 1810 is therefore a valid name for this genus of scy
phozoans, and it is not threatened by a senior homonym.
Peron and Lesueur (1810) established the genus
30
Cyanea with C. arctica Peron and Lesueur, 1810, C. baltica
Peron and Lesueur, 1810, C. borealis Peron and Lesueur,
1810, C. brittanica Peron and Lesueur, 1810, and C. lus-
itanica Peron and Lesueur, 1810 as included species. The
type species of Cyanea has been stated to be Medusa capil
lata Linnaeus, 1758 by authors including Mayer (1910b)
and Kramp (1961). However, Medusa capillata was listed
as a synonym of all five of the nominal species included
by Peron and Lesueur (1810) in their new genus, and
not treated as a valid species. Because it is not one of the
originally included valid species, it is ineligible to be type
species of the genus (Art. 67, ICZN). I was unable to
find any earlier designations of C. arctica or of any of the
other four nominal species included by Peron and Le
sueur (1810) in the genus, as type species of Cyanea. The
choice of C. arctica as type species was made because it is
the most familiar of the five nominal species originally
included in the genus.
Cyanea capillatafulva L. Agassiz, 1862
(Fig. 10)
Cyaneafulva L.Agassiz, 1862:119.
Cyanea capillata var.fulva — Mayer, 1910b: 600.
Common Name
Tawny lion's mane.
Type Locality
USA, New York, Long Island Sound (L.Agassiz, 1862).
Museum Material
None seen from study area.
Specific Characters
Medusa up to 20 cm in umbrella diameter; endoderm
of gastrovascular system and bases of oral arms yellow to
yellow-brown (Mayer, 1910b).
Nematocyst Complement
Medusa (Virginia, USA; Calder, unpublished data):
Tentacles, oral arms—
holotrichous a-isorhizas (6.9—10.7 jam long
x 4.6—6.2 (am wide)
holotrichous A-isorhizas (15.8-20.8 urn long
x 8.0-14.0 jam wide)
holotrichous a-isorhizas (6.6—10.9 jam long
x 2.9—5.6 jam wide)
heterotrichous anisorhizas (11.8—14.5 jam long
x 9.1-11.9 urn wide)
 Fig. 10. Cyanea capillata fulva L. Agassiz, 1862, modified from L. Agassiz (1862).
heterotrichous microbasic euryteles
(12.7-15.4 um long x 8.3-9.2 um wide)
Remarks
Louis Agassiz (1862) believed that medusae of Cyanea
from the mid-Atlantic region of the United States were
referable to a distinct species (C.fulva), marked primar
ily by their yellowish colour. Haeckel (1880), how
ever, relegated C.fulva to the synonymy of C. capillata.
Mayer (1910b) considered it a local race of that species
(C. capillata var. fulva), differing from the rich-brown
form from boreal •waters north of Cape Cod, Massachu
setts (C. capillata capillata). Besides their colour difference,
Mayer noted that medusae of C. capillata vtLi.fulva were
much smaller (rarely exceeding 20 cm in diameter, com
pared with 50—120 cm or more in the northern form).
He added that the lappet notches were more uniform
in C. capillata var.fulva, that the tentacles were much less
numerous and that the oral arms were "less voluminous
and by no means so complexly folded." It is currently
unclear whether the latter characters might be a function
solely of the smaller size of these medusae. Kramp (1961)
included C.fulva in his synopsis of the medusae of the
world, but regarded it as a doubtful species.
The likely existence of taxonomically distinct popu
lations of Cyanea along the Atlantic coast has been sus
pected for decades from jellyfish research conducted at
the Virginia Institute ofMarine Science in the early 1970s
(personal observations), but the question was never stud
ied in any detail. Subsequently, Brewer (1991) concluded
that two populations of medusae of Cyanea in northeast
ern Long Island Sound, USA, apparently corresponding
with C.fulva and C. arctica (= C. capillata),"might be sepa
rate species." Medusae of the two populations differed in
3i
(1) colour, (2) umbrella diameter of mature individuals,
(3) size of the oral folds, and (4) numbers of tentacles.
The two forms were reportedly isolated reproductively by
season of occurrence, as well as by geographic location,
and they were also said to differ in the severity of their
sting. Based on such reported differences, this scyphozoan
is regarded here as a distinct subspecies of C. capillata, and
it might even prove to be a distinct species.
Rice and Powell (1972) reported that the venom of
medusae from Chesapeake Bay (here regarded as Cyanea
capillatafulva) is scarcelydetectable,ifat all,to humans. Brew
er (1991) also reported that medusae corresponding with
C.fulva were non-venomous, or only mildly so,to humans.
No specimens of Cyanea capillata fulva were seen in
the study area during this project. Mayer (1910b) report
ed them "southward to the Carolinas," but he provided
no collection data to document the location or seasonal
ity of these medusae in the region.
Reported Range
Study area:"Carolinas" (Mayer, 1910b).
Overall:"Cape Cod...to the Carolinas;" neritic (Mayer,
1910b).
Cyanea capillata versicolor L. Agassiz, 1862
Cyanea versicolor L. Agassiz, 1862:119. — A. Agassiz,
1865:46.
Cyanea capillata var.versicolor — Mayer, 191Ob:600,
pi. 65, figs. 1, 2.
Cyanea capillata — Hoese, 1973:74. — Kraeuter and
Setzler, 1975:69. — Calder, Boothe, and Maclin,
1977:20.— Calder and Hester, 1978:88.
Common Names
Variegated lion's mane; winter jellyfish.
Type Locality
USA, coast of South Carolina (L.Agassiz, 1862).
Museum Material
South Carolina, off entrance of North Edisto River,
32°31' N, 80°08" W, 9 m, 16 °C, 30.2%o, 11 April 1970,
one medusa, GMBL 70-25. South Carolina, Charleston
Harbor, Rebellion Reach (no coordinates), 12 m, 13 °C,
23%o, otter trawl, 18 March 1971, one medusa, GMBL
71-36. South Carolina, Charleston Harbor, Marine Re
sources Center boat slip,32°45.2' N, 79°53.9'W, surface,
11 °C, 15.1%o, dipnetted, 27 March 1971, one medusa,
GMBL 71-42. South Carolina, Charleston Harbor, Ma-
32
rine Resources Center boat slip, 32°45.2f N, 79°53.9f W,
surface,15 °C, 13.5%o, dipnetted, 11 April 1971, two me
dusae, GMBL 71-46. South Carolina, Charleston Harbor
off Fort Johnson, 32°45.6' N, 79°54.4»W, 8 m, trawl, 21
April 1971, seven medusae, GMBL 71-52. South Carolina,
Charleston Harbor, Marine Resources Center boat slip,
32°45.2' N, 79°53.9'W, surface, 18 °C, 20.5%o, dipnetted,
25 April 1971,36 medusae, GMBL 71-58. South Carolina,
Charleston Harbor, Marine Resources Center boat slip,
32°45.2» N, 79°53.9'W, surface, 18 °C, 20.5%o,dipnetted,
25 April 1971,11 medusae, GMBL 71-59. South Carolina,
shelf waters off Charleston, 32°37.8' N, 79°39.8' W,
14.5 m, 15 °C, 32%o, % Yankee trawl, 27 March 1981,
eight medusae, GMBL 81-5. South Carolina, Folly River,
marina near Folly Road, surface, collected by hand,
17 March 1988, two medusae, ROMIZ B944. South
Carolina, off Charleston Harbor, R/V Anita, 32°43.0' N,
79°48.19' W, 32.8%o, 8.93 °C, bongo net, 9 February
2005, two juvenile medusae, SERTC No. SI746. South
CaroHna,offCharlestonHarbor,R/VAnita,32°42.701'N,
79°47.664' W, 32.8%o, 8.93 °C, bongo net, 9 February
2005 one juvenile medusa, ROMIZ B3595. North Caro
lina, Ocracoke Island, stranded on the beach, 35°05.53' N,
75°58.57 W, 27.5%o, 10 May 2005, one adult medusa,
ROMIZ B3598. North Carolina, Ocracoke Island, Silver
Lake Harbor, National Park Service dock, surface, dip-
netted, 35°06.93' N, 75°59.17'W, 22.5%o, 10 May 2005,
one adult medusa, ROMIZ B3599. South Carolina, off
Charleston Harbor, R/V Anita, 32°41.86' N,79°47.27'W,
30.9%o, 22.3 °C, trawl, 12 May 2005, two medusae,
ROMIZ B3597. South Carolina, off Charleston Harbor,
R/V Anita, 32°24.19' N, 79°47.27' W, 30.9%o, 22.3 °C,
trawl, 12 May 2005, two medusae, SERTC No. S1763.
Specific Characters
Medusa up to 15 cm in umbrella diameter; endoderm
of gastrovascular system and bases of oral arms purplish-
pink to deep red (Mayer, 1910b).
Nematocyst Complement
Undescribed.
Remarks
Although founded as a distinct species (Cyanea versicolor)
by L. Agassiz (1862), Mayer (1910b) considered this me
dusa to be no more than "a well-marked local race of
Cyanea capillata? It was said by him to be smaller than
C. capillata var.fulva,with mature specimens measuring up
to 11 cm in diameter. Mayer reported seeing thousands
of these medusae off the southeast coast of the United
States. While he did not say when or where he encoun
tered them, it was probably during the late fall and early
winter of 1904—05 as he was piloting the yacht Physa-
lia of the Carnegie Institution ofWashington to Florida
from New England for the 1905 opening of the new
Tortugas Laboratory (Stephens and Calder, 2006). Only
a few of the medusae he observed lacked the pinkish
colouration characteristic of the taxon, with those few
having the pale yellow colouration of C. capillata wax.fulva
instead. Even the ephyrae were described as pink, instead
of yellow as in the more northerly form.
Opinions have differed on the status of this taxon.
Haeckel (1880) noted that it was inadequately described,
but nevertheless treated it as a valid species. Like Mayer
(1910b) earlier, Bigelow (1928) regarded Cyanea versicolor,
as well as C. arctica and C fulva,as varieties of C. capillata.
Stiasny and van der Maaden (1943) regarded C. versicolor
as a colour variety ("Farbenvarietat") of C. capillata, while
Kramp (1961) regarded the two as fully conspecific. Rus
sell (1970) noted that a detailed comparison of C. capil
lata, C.fulva, and C. versicolor was needed to sort out their
relationships. Over the past half-century, many authors
have simply used the name C. capillata for these scypho
zoans in the western North Atlantic. Cyanea capillata
versicolor is treated here as a distinct subspecies based on
characters already noted by L. Agassiz (1862) and Mayer
(1910b). Nevertheless, a re-examination of the taxonom-
ic status of these southern medusae, and their relation
ships to more northerly forms of Cyanea along this coast,
is warranted. Notably, medusae of Cyanea in southeastern
Australia, commonly assigned to C. capillata, have recent
ly been shown to constitute two morphologically and
molecularly distinct clades, C. annaskala and C. rosea, that
differ from C. capillata of the North Sea (Dawson, 2005).
Molecular studies are needed to determine whether
C. versicolor from the southeastern United States might
likewise be a valid species, as originally maintained by
L. Agassiz (1862).
Several large specimens of this medusa, pinkish to
orangy-red in colour, were collected from North and
South Carolina during May, 2005. One of these, from
Ocracoke Island, North Carolina (ROMIZ B3599), was
15 cm in diameter.
Cyanea capillata versicolor is one of the most common
scyphomedusae in the study area, although it is gener
ally less familiar than Stomolophus meleagris and Chrysaora
quinquecirrha because of its occurrence during the colder
months of the year.Sometimes known as the "winter jel
lyfish"in the region, it has been reported every month
from December through May (Mayer, 1910b; Hoese,
1973; Kraeuter and Setzler, 1975; this study).This medusa
is reported by local surfers to be mildly venomous.
The scyphistoma stage ofthis species was reported in
the field (as Cyanea capillata), one mile inside the mouth
of the North Santee River, South Carolina, by Calder,
Boothe, and Maclin (1977).
Reported Range
Study area: in bays and shelf waters during winter
throughout the region (L. Agassiz, 1862; A. Agassiz,
1865; Mayer, 1910b; Hoese, 1973; Kraeuter and
Setzler, 1975; Calder and Hester, 1978).
Overall: "in swarms off the coast between Cape Hat
teras...and Cape Canaveral..." (Mayer, 1910b:600);
Gulf of Mexico (Phillips et al., 1969); neritic.
Genus Drymonema Haeckel, 1880
Drymonema Haeckel, 1880:633.
Diagnosis
Medusa with flattened umbrella; oral arms four, long,
frilly, surrounding central mouth; rhopalia eight; tentacles
numerous, arising in a wide band from subumbrella proxi
mal to umbrellamargin;stomach pouches terminating in
numerous dichotomously-branched ramuli; gonads four,
much folded, interradial (Mayer, 1910b; Kramp, 1961).
Type Species
Drymonema dalmatinum Haeckel, 1880,by monotypy.
Drymonema dalmatinum Haeckel, 1880
(Fig. 11)
Drymonema dalmatina Haeckel, 1880:642.
Drymonema dalmatinum —Williams et al., 2001:128.
Common Names
Purple seamane;big pink jelly;stinging cauliflower, pink
meanie.
Type Locality
Mediterranean Sea;Dalmatian Coast, Hvar Island ("Mit-
telmeer; Kiiste von Dalmatien, Insel Lesina") (Haeckel,
1880).
Museum Material
South Carolina, in or near the GulfStream east ofCharles
ton, Albatross Station 2626, 32o27'30" N, 77°20f30" W,
21 October 1885, three medusae, USNM 61670.
33
 Fig. 11.Drymonema dalmatinum Haeckel,1880,after Mayer (1910b) and Haeckel (1881b).
Specific Characters
Medusa up to 1 m in umbrella diameter; exumbrella
with >100 radial furrows; colour pinkish to light purple
(Kramp, 1961).
Nematocyst Complement
Undescribed.
Remarks
Drymonema victoria Haeckel, 1881a, originally described
from the vicinity of Gibraltar, and D.gorgo F. Miiller, 1883,
from Brazil, are generally considered conspecific with D.
dalmatimim (Mayer, 1910b; Stiasny, 1940; Kramp, 1961; Lar
son, 1987a; Mianzan and Cornelius, 1999). Medusae of this
species reach as much as a metre in umbrella diameter.
Drymonema dalmatinum was present in the warm
western North Atlantic more than a century ago (Wil
liams et al., 2001), and it is not a recent invasive species in
the region as sometimes thought.The specimens collected
off South Carolina in 1885 (see MUSEUM MATERIAL
34
above) were likelybeing transported northwards in the Gulf
Stream.While it is improbable that D. dalmatinum is indig
enous to nearshore waters of the study area, eddies could
transport medusae of the species towards the coastline.
In addition to the three medusae of this species tak
en from waters off South Carolina in 1885, collections
at the National Museum of Natural History, Smithso
nian Institution, include another 10 specimens collected
in 1898 from the Bermuda area (Williams et al, 2001).
Moreover, I personally observed this scyphomedusa in
Bermuda during studies on hydroids of the region be
tween 1976 and 1997. One specimen was stranded on
rocks at Flatts Inlet during September 1977, and another
was observed in the water column during a submers
ible (SDL-1) dive on Challenger Bank (5 March 1997).
Acquaintances familiar with D. dalmatinum in Bermuda
report that it inflicts a painful sting and Humann (2002)
described it as highly toxic. In contrast, Williams et al.
(2001) reported that envenomations by the species do
not appear to be severe.
 Larson (1987a) provided accounts of swimming and
feeding behavior, growth, and associations with fishes
and hyperiid amphipods in Drymonema dalmatinum from
the Caribbean. Medusae of Amelia aurita appeared to be
a major prey item of the species.A bloom of D. dalmati
num around Puerto Rico during summer and autumn of
1999 was described by Williams et al. (2001).
Reported Range
Study area: east of Charleston, SC (32°27,30n N,
77°20'30" W) (Williams et al., 2001).
Overall: South Carolina to Argentina, Gulf of Mexico,
eastern Atlantic including the Mediterranean; nerit-
ic (Kramp, 1961; Larson, 1987a;Williams et al.,2001;
Morandini et al., 2005).
Family Ulmaridae Haeckel, 1880
Remarks
Although the family group name Aureliidae L. Agassiz,
1862 predates Ulmaridae Haeckel, 1880, the former has
long been held to be a subfamily of the latter, and pre
cedence of Ulmaridae over Aureliinae is maintained here
(ICZN Art. 35.5).
Genus Amelia Lamarck, 1816
Aurellia Peron and Lesueur, 1810:357 [name suppressed
under the plenary powers by the International
Commission on Zoological Nomenclature (1958,
Opinion 515); name number 1157 on the Official
Index of Rejected and Invalid Generic Names in
Zoology].
Amelia Lamarck, 1816:512 [name validated under the
plenary powers by the International Commission
on Zoological Nomenclature (1958, Opinion 515);
name number 1268 on the Official List of Generic
Names in Zoology].
Diagnosis
Medusa with plate-shaped to shield-shaped umbrella;
oral arms four, unbranched, surrounding a mouth; rhopa-
lia eight; lappets and tentacles numerous, arising from ex-
umbrella just above margin; radial canals branching and
sometimes anastomosing, extending outwards towards
margin from central stomach; ring canal present; gonads
four, much folded, interradial (Mayer, 1910b; Kramp,
1961; Russell, 1970).
Type Species
Medusa aurita Linnaeus, 1758, by designation under the
plenary powers by the International Commission on
Zoological Nomenclature (1958, Opinion 515).
Amelia marginalis L. Agassiz, 1862
(Fig. 12)
Amelia marginalis L. Agassiz, 1862:86.
Amelia aurita— Hoese, 1973:94. — Kraeuter and Setzler,
1975:69. — Calder and Hester, 1978:88. — Roun-
tree, 1983a:25 [not Amelia aurita (Linnaeus, 1758)].
Amelia aurata — Schwartz and Chestnut, 1973:60
[incorrect subsequent spelling]
[not Amelia aurita (Linnaeus, 1758)].
Common Name
Southern moon jelly.
Type Locality
"...reefs of Florida" (L.Agassiz, 1862).
Museum Material
South Carolina, continental shelf east of Edisto Island,
R/V Dolphin, MARMAP coll. no. 0573410,32°37.0' N,
78°30.0'W, 141 m, 22 °C, 36.7%o, ^Yankee No. 36 trawl,
9 November 1973, one fragmentary medusa, ROMIZ
B3590. South Carolina, seaward ofbuoy 17 offCharleston
jetties, R/V Silver Crescent, 32°42.31' N, 79°47.77'W,
8 m, 28.7 °C, 31.6%o,otter trawl, 1 September 2005, one
medusa, ROMIZ B3605.
Specific Characters
Medusa flatter than a hemisphere, up to 30 cm in um
brella diameter; oral arms usually less than radius of um
brella; genital pouches nearly half as wide as umbrella
radius (Mayer, 1910b). Scyphistoma with about 16 ten
tacles; mouth round or irregular in shape, seldom cru
ciform; oral region with polyspira nematocysts; asexual
reproduction by budding, not by podocyst formation.
Nematocyst Complement
Medusa (Virginia, USA; Calder, unpublished):
Tentacles—
holotrichous a-isorhizas (5.6—6.9 um long
x 3.8-4.7 um wide)
holotrichous a-isorhizas (6.8—9.0 jamlong
x 2.7—3.5 um wide)
heterotrichous microbasic euryteles
(10.0-11.8 um long x 6.0-6.9 yon wide)
35
Remarks
Mayer (I910a:4) mused "with the exception ofthe spong
es and corals, there is no phylum of the animal kingdom
more difficult to classify than the medusae." Taxonomic
confusion and uncertainty extends to species recognition
even in the familiar, widespread, and much-studied ge
nus Amelia. Kramp (1968a) commented on problems that
have confounded differentiation of species within the ge
nus. He highlighted characters that he considered useful
in the taxonomy of Amelia (e.g., branching of the canals,
formation of secondary notches on the margin, folding
of the edges of the oral arms) and listed others that he
believed held no value (e.g., colouration, mesoglea thick
ness, extent of vaulting of the exumbrella, shape and size
of the genital pits, the ratio of gonadal to umbrella diam
eter, number of canals arising from genital sinuses). Dif
ficulties involving the identification of species of Amelia
persist,as is evident in recent revisions of nominal species
of the genus from the Pacific coast of the United States
and Canada (Gershwin, 2001). As noted by Gershwin,
there is little agreement to date over which characters are
of taxonomic value in these medusae. Moreover, Daw
son and Jacobs (2001) and Dawson (2003) demonstrated
the existence of several cryptic species of Amelia and en
couraged the application of molecular and phylogenetic
methods to sort out the taxonomy and phylogeny of the
genus. Schroth et al. (2002) provided additional evidence
of the existence of cryptic species in Amelia based on
molecular methods.
Taxonomic uncertainties extend also to the identity
of species of the genus occurring along the Atlantic coast
of North America, and the existence of cryptic species
here has to be considered possible or even likely. In pio
neering work, L.Agassiz (1862) maintained that south
ern populations of Amelia were morphologically differ
ent from those of the northeastern states. He assigned
medusae from New England to Amelia flavidula Peron
and Lesueur, 1810, and established the name A. marginalis
for a species he encountered in southern Florida.Mayer
(1910b) dismissed this classification after comparing the
meristics of medusae from theTortugas, Florida, and from
Eastport, Maine, and he considered both A. flavidula and
A. marginalis as synonyms of A. aurita (Linnaeus, 1758).
Later workers, including Kramp (1961), followed Mayer,
and the name A. marginalis was treated as a synonym of
A. aurita (type locality: Baltic Sea).
Considerable evidence now exists to support the
contention of Louis Agassiz that southern populations
of Amelia are taxonomically distinct from those of bo
real waters to the north. Justification for recognizing a
Fig. 12. Amelia marginalis L. Agassiz, 1862, from a photograph
of a specimen by the author.
taxonomically distinct species group taxon for southern
Amelia in the present work is based more on differences
in scyphistomae from the two geographic areas than on
their medusae. Of particular note are biochemical and
morphological comparisons of scyphistomae identified as
"Amelia aurita" from Chesapeake Bay,Virginia, and from
locations further north in New England. Differences in
the free amino acid (FAA) composition of polyps from
Virginia and Massachusetts were documented by Webb
et al. (1972). Whereas (3-alanine constituted a major frac
tion of FAA in those from the former location, it was
lacking in scyphistomae from the latter. Differences in
nematocyst complement have also been noted, with scy
phistomae of Amelia from Virginia, Delaware, and Texas
all having polyspiras (a variety of holotrichous isorhizas),
and those from Massachusetts lacking them (Webb et al.,
1972; Calder, 1983; Calder, unpublished data). In terms
of basic morphology, Morales-Alamo and Haven (1974)
found that mouth shape was predominantly cruciform
in fully developed scyphistomae from Massachusetts and
from New Brunswick, Canada, and round in those from
Chesapeake Bay and Delaware Bay as well as in those
from Mississippi and Texas in the Gulf of Mexico. They
also observed that scyphistomae of Amelia from Massa
chusetts formed podocysts, while those from Chesapeake
Bay did not. Finally, Zubkoff and Lin (1975) compared
isozyme (malate dehydrogenase and tetrazolium oxidase)
patterns in scyphistomae of Amelia from several regions.
A number of geographic groupings of scyphistomae

36
were recognized, based on differences in their isozyme
patterns: (1) northern northwestAtlantic (Massachusetts,
USA, and New Brunswick and Prince Edward Island,
Canada), (2) middle northwest Atlantic (Chesapeake
Bay, Delaware Bay), and (3) northeast Pacific (Washing
ton, USA). Scyphistomae from Great Britain were most
like those from the northern northwest Atlantic. Earlier,
Spangenberg (1964) observed several characters in scyph
istomae of Amelia from Texas that differed from previous
descriptions of specimens from boreal waters. These in
cluded asexual reproduction by active budding from the
lateral walls of scyphistomae instead of by stolon forma
tion, the absence of any podocyst development, the abil
ity to glide rather rapidly over the substrate rather than
being more sedentary, the occurrence of the polyspira
variety of nematocysts, and the ability of young speci
mens to swim. The same attributes have been noted in
scyphistomae of Amelia fromVirginia that were cultured
in the laboratory (personal observations).
As for the medusa, physiological differences have
long been known to exist between those assigned to
Amelia aurita from Halifax, Nova Scotia, and Tortugas,
Florida (Mayer, 1914). Pulsation rates in relation to water
temperature were shown by Mayer to differ considerably
in the two populations. Moreover, upper lethal tempera
tures were 26.8—28 °C for medusae from Halifax ver
sus 40 °C for those from Tortugas.These observations have
been attributed in the past to acclimation and not to taxo
nomic differencesbetween the populations. Finally, Dawson
(2003) suggested on the basis of molecularand phylogenetic
researchthat A. aurita is likely endemic to the boreal Adantic
Ocean, a region to the north of the current study area.
Taken together, such evidence is seen as sufficient
justification for recognizing here a distinct species group
taxon of Amelia occurring at least from Delaware Bay
southwards to Florida and the West Indies (Mayer,
191Ob:621). The old species group name marginalis L.
Agassiz, 1862 is resurrected for this taxon here. Admit
tedly, more research and deliberation is needed to con
firm that the medusa assigned the name Amelia margin
alis from reefs of south Florida by L. Agassiz (1862) is
the same species as that occurring from the southeastern
states northwards to Delaware Bay and possibly beyond.
Indeed, the possibility that more than one species exists
in the region should be explored.While A. marginalis was
only briefly defined and never illustrated by L. Agassiz
(1862), and its identity remains somewhat questionable,
use of that name seems preferable than establishing a new
name for the warm-water semaeostome treated here.
Type material of the species (MCZ 352) from Key West,
Florida, is housed in the Museum of Comparative Zool
ogy, but I have not examined it. Finally, more research is
necessary to determine whether the species of Amelia
found in cold waters off northeastern North America is
A. aurita, originally described from the Baltic Sea (Lin
naeus, 1758). The isozyme studies of Zubkoff and Lin
(1975) indicated an affinity between scyphistomae from
boreal waters of the northwest and northeast Atlantic.
Medusae of Amelia marginalis (reported as A. aurita)
are of sporadic occurrence in the region. Records are
limited but indicate a summer and fall seasonality of the
species here. Hoese (1973) reported it from waters off
Sapelo Island, Georgia, during the month of September.
Kraeuter and Setzler (1975), also working in the Sapelo
Island area, collected seven specimens during July 1971.
In North Carolina, Schwartz and Chestnut (1973) found
medusae of this species in small numbers during August
and September in the Newport River and along the
coast from Cape Lookout to Cape Fear. Medusae up to
457 mm in diameter were collected in their samples.Also
in North Carolina, Rountree (1983a) observed medu
sae of A. marginalis (as A. aurita) in moderate numbers
offWrightsville Beach during October and November,
1982. No medusae of this species were seen by me dur
ing eight years (1973—1981) of field work in estuaries
of South Carolina, although Calder and Hester (1978)
reported an ephyra collected in Bull Bay on 23 July
1973. An adult medusa was collected in a trawl during
the present study at the entrance of Charleston Harbor
on 1 September 2005 (ROMIZ B3605).
Species of the genus Amelia have infrequently been
reported asvenomous to humans, and most accounts have
described them as harmless (Williamson et al., 1996).
Reported Range
Study area: Newport River, and nearshore waters be
tween Cape Lookout and Cape Fear, NC (Schwartz
and Chestnut, 1973); off Sapelo Island, GA (Hoese,
1973); Hudson Creek, Doboy Sound, GA (Kraeuter
and Setzler, 1975); Bull Bay, SC (Calder and Hester,
1978);Wrightsville Beach, NC (Rountree, 1983a);
all as Amelia aurita.
Overall: Delaware Bay to the West Indies, and Gulf
of Mexico; neritic (L. Agassiz, 1862; Hyde, 1894;
Mayer, 1910b; Burke, 1975, as Amelia aurita).
37
 Order Rhizostomeae Cuvier, 1799
Suborder Kolpophorae Stiasny, 1921b
Family Mastigiidae Stiasny, 1921b
Genus Phyllorhiza L. Agassiz, 1862
Phyllorhiza L. Agassiz, 1862:158.
Diagnosis
Medusa with nearly hemispherical umbrella; oral arms
eight, adradial, gelatinous, dichotomous, fused basally,
broad in lateral view, 3-winged,J-shaped terminally, with
large fenestrae in lateral membranes, with numerous fila
ments and with many small mouth openings surrounded
by minute tentacles; subumbrellar muscles annular; scapu-
lets ("shoulder ruffles") absent on mouth-arms; marginal
tentacles lacking; rhopalia eight; radial canals eight, ex
tending to umbrella margin and connecting with a ring
canal; centripetal canals from ring canal joining central
stomach (Mayer, 1910b; Kramp, 1961; Cornelius, 1997).
per the diagnosis of the genus by Stiasny (1924). Kramp
(1965) considered this diagnosis to be somewhat vague,
and noted that it was"almost equal" to that of the genus
Mastigias. Nevertheless, he considered it advisable to re
tain Phyllorhiza for P. punctata, a common species in Aus
tralian waters.
In order to maintain current usageofthe generic name
for this well-known and widelydistributed invasive species,
and to avoid possiblenomenclatural confusion, a caseshould
be submittedto the ICZN requesting that Phyllorhiza punc
tata be designated as type species of Phyllorhiza under the
plenary powers. The diagnosis given above is in accor
dance with the current concept of Phyllorhiza.
Phyllorhiza punctata von Lendenfeld, 1884
(Fig. 13)
Phyllorhiza punctata von Lendenfeld, 1884:296, pi. 4.
Phyllorhiza punctata —Johnson and Allen, 2005:104,105.

Type Species Common Names

Phyllorhiza chinensis L. Agassiz, 1862, by monotypy. Australian spotted jellyfish; spotted jellyfish; whitespotted
jellyfish; blue-tinted jellyfish.
Remarks
Significant nomenclatural problems emerge with the rec Type Locality

ognition here that Phyllorhiza chinensis L.Agassiz, 1862 is Port Jackson, Australia (von Lendenfeld, 1884).
the type species of this genus.The concept of Phyllorhiza
Museum Material
over the last century has been based on the erroneous
South Carolina, off BuU Island, 33.846° N, 79.593° W,
assumption, asserted in influential works such as those
6 m, 16.9 °C, 35.94%o, trawl, 8 November 2007, one
of Mayer (1910b) and Kramp (1961), that Phyllorhiza
medusa, SERTC #2921. South Carolina, off North Is
punctata von Lendenfeld, 1884 is the type species of the
land, 33.220° N, 79.129° W, 8 m, 18.8 °C, 35.72%o, trawl,
genus. Phyllorhiza punctata was not an originally included
7 November 2007, one medusa, SERTC #2923. South
nominal species when Phyllorhiza was established, and it
Carolina, off North and South Santee rivers, 33.062° N,
is therefore ineligible to be fixed as the type species of
79.170° W, 8 m, 18.9 °C, 36.41%o, trawl, 7 November
the genus (ICZN Art. 67.2). As for Phyllorhiza chinensis
2007, one medusa, SERTC #2924. South Carolina,
L. Agassiz, 1862, the only originally included species in
off North Island, 33.292° N, 79.130° W, 9 m, 19.0 °C,
the genus, and its type species by monotypy (ICZN Art.
36.20%o, trawl, 7 November 2007, one medusa, SERTC
68.3), its identity is uncertain. Mayer (1910b) regarded
#2925. South Carolina, off Litchfield Beach, 33.457° N,
it as "too imperfectly described to be recognizable," but
79.012° W, 9 m, 18.7 °C, 36.26%o, trawl, 7 November
suggested that it was a probable synonym of Cephea cephea
2007, one medusa, SERTC #2926.
(Forsskal, 1775), an Indo-Pacific rhizostome assigned to
a different family from P. punctata. Kramp (1961) men
Specific Characters
tioned P. chinensis only incidentally, referring to Mayer's
Medusae with more or less hemispherical umbrella, up to
(1910b) conclusion that it was "imperfectly described."
50 cm in diameter; exumbrellar surface granular; oral arms
Problems over the type species of this genus were noted
eight, thick, transparent, each with a long and ribbon-
almost a century ago by Cockerell (1911), but his con
shaped appendage; coronal muscles in eight separate fields,
cerns have been overlooked or ignored. He believed that
interrupted at radial canals;colour bluish brown, with nu
a new generic name was warranted for Phyllorhiza sensu
merous white spots on exumbrella; zooxanthellae present
von Lendenfeld (1884) and Mayer (1910b).
or absent (Kramp, 1961; Mianzan and Cornelius, 1999).
The current concept of Phyllorhiza is essentially as

38
Nematocyst Complement
Medusa (Lake Illawarra, NSW, Australia; Peach and Pitt,
2005):
Oral arms—
holotrichous isorhizas, oval
(5.1 urn long x 3.7 urn wide)
rhopaloids, medium (7.6 um long x 5.4 um wide)
rhopaloids, large (14.4 um long x 10.7 um wide)
birhopaloids (14.5 uxn long x 10.2 um wide)
Umbrella margin—
holotrichous isorhizas, round
(5.7 u.m long x 4.9 um wide)
holotrichous isorhizas, oval
(5.5 um long x 3.5 urn wide)
rhopaloids, medium (7.4 urn long x 4.8 um wide)
rhopaloids, large (13.3 urn long x 9.1 um wide)
birhopaloids (13.5 um long x 10.5 um wide)

Remarks
Phyllorhiza punctata von Lendenfeld, 1884 has generally
been regarded as an invasive speciesin the western Atlan
tic.Although recently reported in the Gulf of Mexico, at
times in substantial numbers (Graham et al., 2003; Bol
ton and Graham, 2004), the medusa is known to have
been present in the Atlantic for more than three decades.
Cutress (1973) reported it from Puerto Rico as early as
1969, and noted that it was abundant in most bays and
harbors on the north and west coasts of the island in Fig. 13. Phyllorhiza punctata von Lendenfeld, 1884, after
1971. He stated that it had also been found in Jamaica. Mianzan and Cornelius (1999) and Johnson and Allen
This species had been reported even earlier, as Mastigias (2005).
sciutillae, by Moreira (1961) from Brazil in the South At
lantic (Migotto et al., 2002). Hypotheses as to possible
mechanisms of dispersal of this species were reviewed by Aquatic Species webpage of the United States Geological
Bolton and Graham (2004). Survey (http://nas.er.usgs.gov/queries/collectioninfo.asp?
Medusae of Phyllorhiza punctata were first collected SpeciesID=119, last accessed on March 10, 2009). As
within the study area from the Banana River, Florida noted above, five specimens from the region are housed
(shoreward of Cape Canaveral), in 2001 (Johnson and Al in collections at the Southeastern Regional Taxonomic
len, 2005; Lauren Hall, pers. coram., 2005, 2006). Hall Center, Marine Resources Research Institute, Charleston,
reported that specimens were observed again during South Carolina.

July 2006 in Guana Lake, Florida, between St. Augus
tine and Jacksonville. More specimens were seen dur
ing August 2006 at the same location (Justin Ellenberger,
pers. comm., 2006). Hall noted that specimens from the
east coast of Florida differed from specimens in the Gulf
of Mexico in containing algal symbionts in their tissues.
Zooxanthellae are present in populations of the species
from Australia (Pitt et al., 2005).
By 2007, Phyllorhiza punctata had been reported
This medusa is known to have a scyphistoma stage
in its life cycle (Cutress, 1973; Hofmann and Crow,
2002), but little has yet been published on these polypoid
stages.
Published reports indicate that Phyllorhiza punctata
is only mildly venomous to humans (Williamson et al.,
1996; Humann, 2002).
Reported Range

from scattered locations throughout the study area from Study area: North Carolina to Florida (Johnson and
Florida to North Carolina: see NAS—Nonindigenous Allen, 2005; Lauren Hall, pers. comm., 2005, 2006;

39
 Justin Ellenberger, pers. comm., 2006; David Knott,
pers. comm., 2007).
Overall: North Carolina to Brazil, Gulf of Mexico,
Mediterranean Sea, Pacific Ocean; neritic (Kramp,
1961; Garcia and Durbin, 1993; Mianzan and Cor
nelius, 1999; Mills, 2001; Graham et al., 2003;John-
son and Allen, 2005; Morandini et al., 2005; Mo-
randini, Soares, et al., 2006; NAS—USGS webpage
[http://nas.er.usgs.gov/queries/collectioninfo.asp?
SpeciesID=1192, last accessed on March 10,2009]).
Suborder Daktyliophorae Stiasny, 1921b
Superfamily Rhizostomatoidea Cuvier, 1799
Remarks
Scapulatae Stiasny, 1921b (established as "Stamm Scapu-
latae"), sometimes adopted as the name of this super-
family, is invalid as a family group name under the code
(ICZN Art. 11.7.1.1) because it was not based on a ge
neric name in the taxon that was used as valid at the time
ofits establishment. Likewise, the name Inscapulatae Stia
sny, 1921b is invalid as a family group name for another
taxon at the same rank in the suborder Daktyliophorae.
Both could be used as names of taxa at ranks above the
family group.
Family Rhizostomatidae Cuvier, 1799
Genus Rhopilema Haeckel, 1880
Rhopilema Haeckel, 1880:596.
Diagnosis
Medusa with dome-shaped umbrella; oral arms eight,
adradial, gelatinous, fused basally, 3-winged distally, each
with numerous appendages or filaments and with nu
merous small mouth openings surrounded by minute
tentacles; scapulets ("shoulder ruffles") borne on manu
brium; marginal tentacles lacking; rhopalia eight; radial
canals 16, extending to umbrella margin, connecting
with a mesh of anastomosing canals between each adja
cent pair; gonads four, much folded, interradial (Mayer,
1910b; Kramp, 1961).
Type Species
Rhopilema rhopalophorum Haeckel, 1880, by monotypy.
40
Rhopilema verrilli (Fewkes, 1887)
(Fig. 14)
Nectopilema verrilli Fewkes, 1887:120, pi. 4.
Rhopilema verrillii — Mayer, 1910b:707,pi. 74, fig. 1.
— Schwartz and Chestnut, 1973:60 [incorrect
subsequent spelling]
Rhopilema verrilli — Hoese, 1973:74. — Kraeuter and
Setzler, 1975:69. — Calder and Hester, 1978:88.
Common Names
Mushroom jelly; mushroom cap jelly.
Type Locality
USA, Connecticut, New Haven Harbor, "at the mouth
of the Quinnipiac River" (Fewkes, 1887).
Museum Material
South Carolina, Charleston Harbor off Fort Johnson,
32°45.6' N, 79°54.4' W, 8 m, trawl, 21 April 1971, one
medusa, GMBL 71-52. South Carolina, Folly Beach,
32°39.12' N, 79°56.76'W, 22 °C, stranded on beach, 13
May 2005, one medusa, ROMIZ B3600.
Specific Characters
Medusa reaching 45 cm or more in umbrella diameter;
umbrella thick, firm; exumbrella smooth; each octant
with six velar lappets; oral arms with up to 60 spindle-
shaped appendages, a large, terminal, tapering one on
each (Kramp, 1961; Calder, 1972b).
Nematocyst Complement
Medusa (Virginia, USA; Calder, 1972b):
Gastric cirri, appendages, oral tentacles—
holotrichous a-isorhizas (5.3—6.9 um long
x 3.3-4.5 um wide)
heterotrichous anisorhizas (6.9—8.9 um long
x 5.4-7.4 jam wide)
heterotrichous microbasic euryteles
(7.6-10.1 jam long x 5.0—7.1 jam wide)
Remarks
The taxonomic status of Rhopilema verrilli has been a
matter of some debate. Stiasny (1921a) considered de
scriptions of the medusa by Fewkes (1887) and Mayer
(1910b) to have been in error, and Kramp (1970b) con
sidered it to be an invalid species. Calder (1972b) ex
amined new material, and concluded that R. verrilli was
a distinct species. However, the generic affinities of this
species need to be reviewed.

Fig. 14. Rhopilema verrilli (Fewkes, 1887), after Mayer (1910b).
Rhopilema verrilli has been reported along the east
coast of the United States from Long Island Sound
(Fewkes, 1887; Mayer, 1910b) to Georgia (Hoese, 1973;
Kraeuter and Setzler, 1975; Harper and Runnels, 1990).
It also occurs along the northern coast of the Gulf of
Mexico, although it is apparently rare west of the Missis
sippi Delta (Harper and Runnels, 1990).The medusa has
never been reported in any notable abundance anywhere,
and it has received little scientific study across its entire
range.
Medusae of Rhopilema verrilli have been reported in
the study area during the colder seasons of the year, hav
ing been recorded each month from November through
May (Mayer, 1910b; Hoese, 1973; Schwartz and Chestnut,
1973; Kraeuter and Setzler, 1975). Fishermen in Pam
lico Sound, North Carolina, informed Mayer (1910b)
that the medusa was seen there quite frequently during
autumn. In South Carolina, large specimens often wash
ashore on beaches in spring (personal observations). At
the northern end of its range, medusae seem to appear
during late summer or autumn. Mayer (1910b) provided
records of the species from several locations in the Long
Island Sound area during August, September, and Octo
ber. Of six specimens from Virginia examined by Calder
(1972b), four juveniles (3-9 cm in diameter) and one
mature female (24 cm in diameter) were taken in No
vember, and one mature male (45 cm in diameter) was
taken during January. Medusae of R. verrilli have been
found from December to May in the Gulf of Mexico. A
single specimen was reported from the Gulf during July,
1971 (Burke, 1976; Harper and Runnels, 1990), but it
seems possible that the medusa may have been misidenti-
fied. During the winter of 2006,83 specimens of various
sizes were observed stranded on beaches in southwestern
Florida (Fort Myers Beach; Sanibel Island; Lover's Key
State Park) between 17 January and 27 February (per
sonal observations).
The life cycle of Rhopilema verrilli is metagenetic.
Cargo (1971) briefly described the scyphistomae of a
medusa identified as this species from the Patuxent Riv
er, Maryland. They were obtained following isolation
of gonadal tissue in cultures from a medusa captured at
Solomons, Maryland. Calder (1973) established labora
tory cultures of scyphistomae from a medusa collected
on the eastern shore ofVirginia, near Wachapreague, and
provided descriptions and illustrations of the planula,
scyphistoma, strobila, ephyra, and young medusa stages.
Strobilation was usually monodiscous, although two to
three ephyrae were produced by a given strobila on oc
casion. Podocyst formation was the only observed means
of asexual reproduction in the cultures.
The nematocyst complement of the medusa was de
scribed by Calder (1972b), and that of the planula, scy
phistoma, strobila, and ephyra stages by Calder (1973).
Additional information on the cnidome of the ephyra
stage was provided by Calder (1977).This species is not
considered venomous to humans (Calder, 1972b).
Rhopilema verrilli is the largest medusa normally en
countered in inshore waters between Capes Hatteras and
Canaveral.The purple sea mane (Drymonema dalmatinum)
may attain a larger size, but it has seldom been reported
in the study area.
The scyphistoma stage of Rhopilema verrilli was re
ported in nature from Little River Inlet, South Carolina,
in dredge collections taken 21 and 22 April 1976 (Calder,
Bearden, et al, 1977).
Reported Range
Study area: Pamlico Sound, NC (Mayer, 1910b;
Schwartz and Chestnut, 1973); Sapelo Island, GA
(Hoese, 1973; Kraeuter and Setzler, 1975); numerous
locations across the coastal zone of South Carolina
(Calder and Hester, 1978).
Overall: Long Island Sound to Georgia, northern
4i
 and southeastern Gulf of Mexico; neritic (Mayer, Type Locality
1910b; Hedgpeth, 1954; Kramp, 1961; Burke, 1975; USA, Georgia, "Warsaw Island" (Wassaw Island), and
Harper and Runnels, 1990; personal unpublished South Carolina, Charleston (L.Agassiz, 1862).
observations).
Museum Material
Georgia (without location or depth data), 16 May 1924,
Family Stomolophidae Haeckel, 1880 one medusa, USNM 42164. North Carolina (without lo
cation or depth data), 17 April 1927, one medusa, USNM
Genus Stomolophus L. Agassiz, 1860
42194. South Carolina (without location or depth data),
9 July 1945, five medusae, USNM 53597. North Caro
Stomolophus L.Agassiz, 1860:pl. 14, figs. 1-8.
hna, near Cape Fear,R/V Silver Bay,33°53' N, 78° 18' W,
depth and date not given, one medusa, USNM 53608.
Diagnosis
South Carolina, Inlet Creek, 32°47.8' N, 79°49.0' W,
Medusa with nearly hemispherical umbrella; mouth-
5 m, 29 °C, 32.4%o, 20-ft otter trawl, 24 September 1970,
arms eight, adradial, gelatinous, 3-winged distally but
five young medusae, GMBL 70-183. Florida, continen
fused throughout, bearing numerous small mouth open
tal shelf east ofJacksonville, R/V Dolphin, MARJVLAP
ings surrounded by minute tentacles; appendages and
coU. no. 0575242,30°16.5' N, 80°58.6.0' W, 24 m, 26 °C,
filaments lacking; scapulets ("shoulder ruffles") borne on
36.3%o,3A Yankee No. 36 trawl, 11 September 1975, three
manubrium; marginal tentacles lacking; rhopalia eight;
medusae, ROMIZ B3587. Georgia, continental shelf east
radial canals 16, extending to umbrella margin, con
of Doboy Sound, R/V Dolphin, MARMAP coll. no.
necting with a mesh of anastomosing canals between
0576364, 31°26.8' N, 80°51.8' W, 16 m, % Yankee No.
each adjacent pair; gonads four, much folded, interradial
36 trawl, 12 September 1976, one medusa, ROMIZ
(Mayer, 1910b; Kramp, 1961).
B3592. South Carolina, newly-liberated ephyrae from
cultures based on material collected in Murrells Inlet,
Type Species
Stomolophus meleagris L.Agassiz, 1860, by monotypy. 30 July 1981, ROMIZ B961. South Carohna, James Is
land, Fort Johnson, Marine Resources Division boat ba
sin, 32°75.25' N, 79°89.90» W, surface, dipnet, 1 October
Stomolophus meleagris L. Agassiz, 1860
2002, coll. D. Knott, one young medusa, SERTC No.
(Fig. 15)
S744. South Carohna, Charleston Harbor, Anchorage,
R/V Anita, 32°45.91' N, 79°53.35' W, 28.3 °C, 27.4%o,
Cephea rhizostoma? — Gibbes, 1848:xxiii [not Cephea
otter trawl, 10 August 2004, five young medusae, SERTC
rhizostoma Lamarck, 1816 = Rhizostoma pulmo
No. S2187. South Carolina, Folly Island at Stono Inlet,
(Macri, 1778)].
32°38.52' N, 79°58.48'W, 22 °C, stranded on beach, 13
Stomolophus meleagris L. Agassiz, 1860:pl. 14, figs. 1-8;
May 2005, one medusa, ROMIZ B3601. South Caro
1862:139, 151. — A. Agassiz, 1865:40. — Brooks,
hna, Inlet Creek, R/V Silver Crescent, 32°46.65' N,
1883:136. — Mayer, 1910b:710, pi. 75, figs. 1-3.
79°49.19' W, surface, 28.2 °C,31.6%o,neuston net, 1 Sep
— Corrington, 1927:347. — Gutsell, 1928:358. —
tember 2005, nine young medusae, SERTC No. S2107.
Pearse et al., 1942:181. — Schwartz and Chestnut,
South Carohna, seaward of buoy 17 off Charleston jet
1973:60. — Kraeuter and Setzler, 1975:70. — Calder
ties, R/V Silver Crescent, 32°43.02' N, 79°48.56» W, sur
and Hester, 1978:88. — Calder, 1982:149, figs. 1-4.
face, 28.7 °C, 31.6%o, neuston net, 1 September 2005,
— Calder, 1983:1185. — Rountree, 1983a:19;
25 young medusae, ROMIZ B3609. South Carolina,
1983b:172. — Huang, 1988:341. — Shanks and
seaward of buoy 17 off Charleston jetties, R/V Silver
Graham, 1988:81. — Grant and Ferrell, 1993:77.
Crescent, 32°43.02' N, 79°48.56» W, surface, 28.7 °C,
— Costello and Colin, 1995:399. — Grant et al.,
31.6%o, neuston net, 1 September 2005, two young
1996:123.— Frick et al., 1999:165.
medusae, ROMIZ B3610. South Carohna, Inlet Creek,
Stomolophus — Smith, 1907:224.
R/V Silver Crescent, 32°46.65' N, 79°49.19'W, surface,
Stomolopus meleagris — Shanks and Graham, 1987:159,
28.2 °C, 31.6%o, neuston net, 1 September 2005, seven
[lapsus calami].
young medusae, ROMIZ B3614. South Carolina, Inlet
Creek, R/V Silver Crescent, 32°46.65' N, 79°49.19'W,
Common Names
surface, 28.2 °C, 31.6%o,neuston net, 1 September2005,
Cannonball jelly; cabbagehead jelly;jellyball.

42

Fig. 15. Stomolophus meleagris L. Agassiz, 1860, after Mayer
(1910b).
44 postephyrae, ROMIZ B3616.South Carolina, seaward
of buoy 17 off Charleston jetties, R/V Silver Crescent,
32°43.02' N, 79°48.56' W, surface, neuston net, 1 Sep
tember 2005, six young to maturing medusae, SERTC
No. S2110. South Carolina, seaward of buoy 17 off
Charlestonjetties,outboardboat,32°43.17' N,79°48.16' W,
surface, 19.1 °C, 32.5%o, neuston net, 2 November 2005,
two medusae, SERTC No. S2327. South Carolina, sea
ward of buoy 17 off Charleston jetties, outboard boat,
32°43.17' N, 79°48.16' W, surface, 19.1 °C, 32.5%<>, dip-
net, 2 November 2005, three medusae, ROMIZ B3620.
Specific Characters
Medusa globular and resembling a cannonball,up to 18 cm
in umbrella diameter; mesoglea thick, rigid; exumbrella
appearing quite smooth except for minute nematocyst
warts; velar lappets about 14 per octant; scapulets reach
ing to or beyond umbrella margin; margin of umbrella
with a conspicuous brownish band (Kramp, 1961; Mian-
zan and Cornelius, 1999).
Nematocyst Complement
Medusa (South Carolina, USA; Calder, 1983):
Manubrium, scapulets, gastric cirri—
holotrichous a-isorhizas
(3.8—4.8 urn long x 2.4—3.1 um wide)
small heterotrichous microbasic euryteles
(6.4—10.1 urn long x 4.5—6.5 um wide)
medium heterotrichous microbasic euryteles
(10.5-12.6 urn long x 7.3-9.0 urn wide)
large heterotrichous microbasic euryteles
(13.2-17.8 um long x 8.0-10.3 urn wide)
Remarks
There has been confusion over the date of authorship
of Stomolophus and its type species, Stomolophus meleagris.
Louis Agassiz (1862) provided the first description of the
genus and species, and 1862 is frequently cited as the
date of publication of the names (Neave, 1940b; Kramp,
1961). However, both generic and specific names were
made available earlier by L. Agassiz (1860:Plate 14), who
used the binomen Stomolophus meleagris in association
with illustrations of the species (ICZN Art. 12.2.7).
As noted by Kramp (1970b), the family Stomolophi-
dae, containing the single genus and species Stomolophus
meleagris, is endemic to the Americas. This scyphozoan
has been reported from southern New England to Brazil
in the Atlantic, and from California to Peru in the Pa
cific. This distribution is noteworthy zoogeographically
because the scyphozoan order Rhizostomeae is predom
inantly an In do-west-Pacific group.
Stomolophus meleagris is the most conspicuous me
dusa in the southeastern United States, and it has long
been known to be abundant in the region. Louis Agassiz
(1862:139) reported seeing "myriads" of these medusae
stranded on the beach in April at "Warsaw Island" (Was-
saw Island, Georgia). An even earlier report of the species
from the region was listed as "Cephea rhizostomaT' from
South Carolina (Gibbes, 1848). Mayer (1910b) described
it as "very common" along the coast of the Carolinas and
Georgia, and stated that mature medusae were abundant
during winter and spring in coastal waters from Florida
to South Carolina. It continues to be seasonally abundant,
often washing ashore on beaches in considerable num
bers.These medusae also periodically create problems for
the commercial shrimp fishery of the region by clog
ging nets, damaging catches, and impeding the sorting
process. On the other hand, 5. meleagris has been consid
ered a fishery resource (Huang, 1988; Hsieh et al., 2001).
Harvesting of cannonballs for food has been undertak
en in both Florida and Georgia (Griffin and Murphy,
http://www.dnr.sc.gov/cwcs/pdf/Cannonballjellyfish.pdf,
last accessed on March 10, 2009).
Medusae of Stomolophus meleagris are likely to be sig
nificant predators of plankton in the southeastern United
States, given their abundance in the region. The diet of
the species, as observed by Larson (1991) in the Gulf of
Mexico, consisted of bivalve veligers (mostly oyster lar
vae), copepods, gastropod veligers, larvaceans (Oikopleura
43
sp.), and tintinnids. They may also prey on larvae of red
drum (Sciaenops ocellatus) (Duffy et al., 1997). In turn,
these medusae are preyed upon by sea turtles (most nota
bly the leatherback, Dermochelys coriacea, but also Kemp's
ridley, Lepidochelys kempi, and loggerhead, Caretta caretta)
as well as sunfish (Mola mola) and sharptail mola (Mola
lanceolata) (Grant and Ferrell, 1993; Frick et al, 1999).
Species associated with these medusae as symbionts in
clude the spider crabs Libinia dubia and L. emarginata, as
well as the fishes Monacanthus hispidus, Chloroscombrus chry-
surus, Nomeus gronovii, Peprilus burti, P.triacanthus, Aluterus
schoepji, Caranx bartholomaei, C.fusus, C. hippos, and Hem-
caranx amblyrhynchus (Corrington, 1927, Gutsell, 1928;
Hildebrand, 1954;Thiel, 1976; Rountree, 1983a; Shanks
and Graham, 1988;Tunberg and Reed, 2004). Rountree
(1983a) observed that such associations were dependent
on the population dynamics of both the host medusa and
of its associates.Moreover, numbers of symbionts per me
dusa were related to season and medusa population size.
The life cycle of the cannonball jelly is known from
studies on laboratory cultures of planulae, scyphistomae,
podocysts, strobilae,and ephyrae reared in South Carolina
(Calder, 1982). Information on development of meteph-
yrae and medusae was provided earlier by Mayer (1910b)
on specimens from Charleston Harbor, South Carolina,
and by Stiasny (1922) on material from the Pacific coast
of Panama.
The seasonal population dynamics and movements
of Stomolophus meleagris in the study area appear to be
complex and not well understood. The presumed annual
cycle outlined here is as proposed by Kraeuter and Set
zler (1975), working in Georgia, and Rountree (1983a),
in North Carolina, supported by observations made dur
ing this study.Kraeuter and Setzler suggested that medu
sae trapped inshore do not survive the winter. However,
Rountree (1983b) intimated that there was evidence of
overwintering by adults. After largely or entirely disap
pearing from the region in early to mid-winter, fully-
grown medusae in substantial numbers abruptly appear
in nearshore waters along the coast during late winter or
early spring.These medusae are thought to be either sur
vivors from the previous year that move in from offshore
waters, or individuals that migrate up from the south in
coastal water currents. Lending support to the former
hypothesis is the fact that seasonal patterns of occur
rence appear similar near Fort Pierce, Florida (Tunberg
and Reed, 2004), well to the south of the study area.
During summer there is unmistakable local recruitment,
with postephyrae and very small medusae appearing in
inshore sounds, bays, harbors, and inlets (ROMIZ B3614;
44
ROMIZ B3616). As they grow, juvenile medusae evi
dently migrate out of the estuaries and into waters along
the coast. Indeed, medusae collected at the entrance of
the Charleston jetties on 1 September 2005 (ROMIZ
B3609; SERTC No. S2110) were larger than those col
lected the same day in Inlet Creek (ROMIZ B3614;
SERTC No. S2107). Meanwhile, older and larger medu
sae are thought to move inshore into more sheltered and
less saline waters during summer and autumn. Towards
the latter part of fall and early winter, numbers of me
dusae wane again. It is unclear if this late-year decline in
abundance is due to mortality, or to migrations out of the
area, or a combination of the two. However, field work
conducted during this study in South Carolina (using a
trawl, bongo nets, neuston net, dipnet, and visual obser
vations) provided evidence that medusae move seawards
and out ofthe estuaries in autumn. Only one medusa was
observed inshore during sampling in Inlet Creek (one
station) and Charleston Harbor (two stations) on 2 No
vember 2005, while more than a dozen specimens were
observed at a station seaward of the Charleston jetties
on the same day.The hypotheses of Kraeuter and Set
zler (1975) and Rountree (1983a, b) about the seasonal
distribution and abundance patterns of this ecologically
important species need further testing and refinement. A
lack of knowledge about the distribution and biology of
the sessile stages (scyphistomae, strobilae, podocysts) of
S. meleagris in nature further hinder the understanding of
the population dynamics of the species in the area. How
ever, with postephyrae and very young medusae being
observed in greatest numbers during this study in estuar
ies (e.g., Inlet Creek), it seems probable that the polyps
will eventually be found there as well.
In addition to seasonal changes in the population
size of Stomolophus meleagris, abundances of these me
dusae also vary widely from year to year (Kraeuter and
Setzler, 1975). While the causes of these annual variations
are currently unknown, such variations are not unusual
for jellyfish species (Purcell et al., 2001). A substantial
decline has been noted in counts of cannonball jellyfishes
in the study area in recent years. During surveys to assess
abundances of cannonballs and leatherback sea turtles by
the South Carolina Department of Natural Resources,
mean numbers of medusae per tow fell from 198 and
339 in 2001 and 2002, respectively, to 132 and only two
in 2003 and 2004, respectively (Griffin and Murphy,
http://www.dnr.sc.gov/cwcs/pdf/Cannonballjellyfish.pdf
last accessed on March 10, 2009). When especially
abundant, medusae are a major pest to local fisheries,
especially those of penaeid shrimp. The variability and
unpredictability of medusa numbers are also negative
factors in the potential development of a commercial
fishery based on the species.
Cannonball jellyfishes are active, strong, rapid swim
mers, and capable of directional movement as shown by
Shanks and Graham (1987).The course of these medusae
was described as steady, having little pitch or yaw, and
they rolled counter-clockwise at a constant rate. Move
ments of individuals within a local group were observed
to be non-random, with such medusae tending to swim
in the same direction. Cannonballs often occur in dense
aggregations, and Shanks and Graham suggestedthat ori
entated swimming might play a role in the formation and
maintenance of such swarms.
The cnidome of the planula, scyphistoma, and eph
yra stages, as well as of the medusa, has been described
(Calder, 1983). In the medusa, all of the nematocyst cat
egories noted above were present on the manubrium
and scapulets,but only holotrichous a-isorhizas and small
euryteles were present on the gastric cirri. In that study,
dense aggregations of medium euryteles were observed
in pockets on both the manubrium and scapulets. These
were released into the water in large numbers, often in
aggregates of two to four, by medusae held in buckets
and other containers upon collection. Analogous "nema
tocyst cushions" were reported in the coronate medusa
Atolla wyvillei by Russell (1970), and "nematocyst res
ervoirs" are known to occur also in the pleustonic hy-
drozoans Porpita porpita and Velella velella (Calder, 1988).
Such "nematocyst magazines," distinct from common
"nematocyst batteries," may be more widespread in the
Cnidaria than currently realized. Shanks and Graham
(1988) noted that when disturbed, medusae of Stomolo
phus meleagris released mucus containing discharged and
undischarged nematocysts. They noted that potential
predators (small fish) were driven off by toxins in the
mucus, which created a kind of chemical defence for the
species. Phillips et al. (1969) had noted earlier that "nem
atocyst rich mucous strands" were secreted by medusae
of S. meleagris. Medusae of Cassiopea, another rhizostome
genus, are known to release nematocysts into the water
when stimulated by passing prey such as small crustaceans
(Smith, 1936). Moreover, "nematocyst bags" (ectodermal
vesicular structures heavily armed with nematocysts) are
released into the water by medusae of this genus when
disturbed (Jensch and Hofmann, 1997). From personal
experience in Bermuda and Belize, swimmers can be
painfully stung by floating nematocyst aggregations dis
charged from Cassiopea xamachana without ever having
been touched by a medusa.
Medusae of Stomolophus meleagris are generally
considered nonvenomous to humans (Humann, 2002).
Nevertheless, Burnett and Calton (1985) reported an en
venomation of a 32-year-old woman attributed to this
species on a beach atAvon, North Carolina, in 1984.This
case initially seemed somewhat surprising, if indeed the
responsible organism was S. meleagris. However, Shanks
and Graham (1988) later reported that a diver was stung
after pinching bell (umbrella) margins of these medusae
with forceps in the water. Disturbed medusae released
clouds of nematocysts into the water that were respon
sible for the envenomation, just as described above for
Cassiopea. Under normal circumstances, however, the
species seems unlikely to represent a significant stinging
threat to people.
Reported Range
Study area: common to abundant in estuaries and
nearshore waters across the entire region (Gibbes,
1848; L. Agassiz, 1862; A. Agassiz, 1865; Brooks,
1883; Smith, 1907; Mayer, 1910b; Corrington, 1927;
Gutsell, 1928; Pearse et al., 1942; Schwartz and
Chestnut, 1973; Kraeuter and Setzler, 1975; Calder
and Hester, 1978; Calder, 1982, 1983; Rountree,
1983a, 1983b; Huang, 1988; Shanks and Graham,
1987, 1988; Grant and Ferrell, 1993; Costello and
Colin, 1995; Grant et al, 1996; Frick et al., 1999).
Overall: southern New England to Brazil, Gulf of
Mexico, eastern Pacific; neritic (Hedgpeth, 1954;
Kramp, 1961, 1968b; Kramp, 1970b; Burke, 1975,
1976; Larson, 1976a; Mianzan and Cornelius, 1999;
Morandini et al., 2005; Morandini, Soares,et al.,2006).
45
Acknowledgements
I am grateful to Rachael King, David Knott, Betty
Wenner, Joe Cowan, Nadia Meyers, Susan DeVictor,
and other staff members of the Southeastern Regional
Taxonomic Center, Marine Resources Research Insti
tute (MFJU), South Carolina Department of Natural
Resources, Charleston, SC, for their encouragement
and help in undertaking this study. Data on materials in
collections of the National Museum of Natural History
(NMNH), Smithsonian Institution, are reproduced here
with permission of the museum. I thank Stephen Cairns
and Allen Collins for their cooperation in accessing and
citing those records. The loan of a specimen of Nausithoe
punctata from the NMNH was made possible by Allen
Collins. Information on the occurrence of Phyllorhiza
punctata in northeast Florida was provided by Lauren Hall
of the St.Johns River Water Management District, Palat-
ka, Florida, and Justin Ellenberger of the Florida Fish and
Wildlife Conservation Commission, Ponte Vedra Beach,
Florida. David Knott of MRRI alerted me to records
of P. punctata off the southeastern United States in 2007.
Illustrations herein were prepared by Patrice Stephens-
Bourgeault, of Studio Stephens-Bourgeault in Toronto,
I thank her for her artwork. Appreciation is likewise-
expressed to Ronald J. Larson for granting permission
46
to use several photographs of medusae taken by him.
Librarian Champa Ramjass of the Royal Ontario Mu
seum assisted by obtaining a number of obscure refer
ences on interlibrary loan. Pearce Webster of MFJU
provided data on occurrences of Stomolophus meleagris
and Phyllorhiza punctata from SEAMAP cruises off the
southeastern United States. Sincere thanks are due to Al
len Collins and Lisa-ann Gershwin for constructive criti
cisms of an earlier draft of this manuscript, and to Gaelle
Chevalier for editorial help. Gratitude is also expressed
to Virginia Morin for her work in production of this
report. Funds in support of this research were provid
ed by the Southeastern Regional Taxonomic Center
through NMFS Grant No. NA16FL1490, the National
Science Foundation (Partnerships for Enhancing Ex
pertise in Taxonomy Program), the Natural Sciences
and Engineering Research Council of Canada, and
the Royal Ontario Museum. I am grateful to the Lou
ise Hawley Stone Charitable Trust within the ROM
Foundation for funds to cover costs of publication. This
publication is Contribution Number 627 of the Ma
rine Resources Division, South Carolina Department
of Natural Resources.
Glossary
Appendages — finger-shaped projections present on
the lower surface of the mouth-arms of certain
jellyfishes (e.g.,Rhopilema verrilli). Homologous
with filaments.
Acraspedote medusae — medusae that lack a velum.
Bell — see umbrella.
Disk — see umbrella.
Ectoderm — outermost layer of tissue (epidermal layer
of cells).
Endoderm — innermost layer of tissue, lining gastrovas-
cular cavities (endodermal layer of cells).
Ephyra(e) — the initial and youngest medusoid stage in
the life cycle of a scyphozoan.
Filaments — slender and elongate tentacle-like structures
occurring on the lower surface of the mouth-
arms of certain jellyfishes (e.g., Phyllorhiza punc
tata). Homologous with appendages.
Interradial (interradii, pi.) — located midway between
the four primary radii of a medusa. If the four
primary radii are taken as equivalent to N, E,
S, and W on a compass, interradial regions
occur on the NE, SE, SW, and NW radii. In
cubomedusae, pedalia are interradial while
stomach pouches,lips,andsense organs (rhopalia)
are perradial (on the four primary radii)
(Cornelius, 1997).
Manubrium — the tube that extends from the mouth or
mouths of a jellyfish to the stomach.
Mouth-arms — structures for feeding, usuallyconspicu
ous, suspended from the central gastric region
on the underside of the umbrella. They vary
from four long, frilly, curtain-like structures
surrounding a single central mouth (e.g., in
Chrysaora quinquecirrha) to a thick, fused, firm
gelatinous mass supporting many small mouths
(e.g., in Stomolophus meleagris).
Oral arms — see mouth-arms.
Pedalia — in cubomedusae, the four interradial exten
sions of the lower umbrella supporting the
tentacles. Cubozoan pedalia vary from paddle-
shaped to hand-shaped.
Perradial (perradii, pi.) — Located on the four primary
radii. See interradial.
RJiopalium — a sensory structure, usually comprising
organs for equilibrium and light reception.
Rhopaliar niche — in cubomedusae, a pit containing a
sense organ (rhopalium) located above the rim
ofthe umbrella, with one present on each ofthe
four perradii.
Scapulets — outgrowths from the vertical walls of the
fused mouth-arms ofsome rhizostome medusae
(e.g., Rhopilema verrilli), and bearing numerous
mouth openings like those on the undersides
of the mouth-arms. Also known as "shoulder
ruffles."
Stinging organelles — microscopic organelles used in
food capture and defence, and especially abun
dant on tentacles. They consist of an outer cap
sule and an inner thread that discharges when
triggered. Certain kinds contain venom and
cause the sting of a jellyfish.Upwards of 30 dif
ferent categories have been recognized, named,
and classified based on differences in morphol
ogy and function, and they have proven to be
useful characters in taxonomy. Also known as
nematocysts, cnidocysts, or cnidae.
Tentacles — thread or cord-like strands,often highly con
tractile, bearing numerous stinging organelles.
The term usually refers to those on the margin
of the umbrella of a jellyfish (i.e., marginal ten
tacles), although they are sometimes present on
the undersurface (e.g., in Cyanea capillata fulva;
C. c. versicolor, Drymonema dalmatinum).
Umbrella — the dome- to disk to cuboidal-shaped gelat
inous main body part of a jellyfish, from which
are suspended structures including mouth-arms
and marginal tentacles (absent in some species).
Also known as the bell or disk.
Velarium — a shelf of tissue projecting inward from the
margin of the umbrella in cubomedusae.
Velarial canals — extensions of the gastricpouches into
the velarium of cubomedusae, the number and
shape of which are useful taxonomically.
Velum — a shelf of tissue projecting inward from the
margin of the umbrella in hydromedusae.
47
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