Marine Flora and Fauna of the Northeastern United States. Cnidaria

NOAA Technical Report NMFS Circular 397
Marine Flora and Fauna of
the Northeastern United States.
Cnidaria: Scyphozoa
Ronald J. Larson
August 1976
U.S. DEPARTMENT OF COMMERCE
Elliot l. Richardson, Secretary
National Oceanic and Atmospheric Administration
Rob\~rt
M. White, Administrator
National Marine Fisheries Service
Robert W. Schoning, Director
t'ot Sale by the Superintendent of Documents. U.S. GO'o'crnment Printing Office
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U. S. DEPARTMENT OF COMMERCE
BIBLIOGRAPHIC DATA SHEET
12-73)
NOAA ACCE:;SION NUMBER
NOAA-7610l8l5
4.
2
3
1•
1 •
RECIPIENT'S ACCESSION NUMBER
TITLE AND SL,'ST:TLE
Marine Flora and Fauna of the
Cnidaria: Scyphozoa
7.
NATIONAL OCEANIC AtJD Al'MOSPHERIC ADMINISTRATION
S.
North~astern
United States.
DATE
Au!! 1976
6.
AUTHOR!SI
8.
Ronald J. Larson CU.S. National Museum, Washington. DC)
9.
REPO~T
REPORT NO.
NOAA-TR-NMFS Circular
PERFORMING ORGANIZATION NAME AND ADDRESS
~~7
10. PROJECT/TASK NO.
NOM, National Marine Fisheries Service,
Seattle, WA, 181 CJ S'
Scientific Publications Staff
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12, SPONSORING ORGANIZATION NAME AND ADDRESS
13. TYPE OF REPORT AND PERIOD
COVERED
Same
1'~~h Rent'
14.
15.
PUBLICATION REFERENCE
NOAA Technical Report NMFS Circular 397, August 1976.
16.
21 p, 28 fig, 24 ref.
ABSTRACT
This manual is an introduction to the scyphomedusae found in coastal waters from
Maine to the Chesapeake Bay. It includes a discussion of their identification,
collection, rearing, preservation, and nematocysts. Also included is an introduction
to the natural history of the scyphopolyps and medusa,e, a discussion of stinging
scyphomedusae, a glossary of terms, an illustrated synopsis of ephyrae, an
illustrating key to the scyphomedusae (including the Stauromedusae), an annotated
systematic list, a bibliography of major refer~nces, and an index.
(Author)
~-
17.
17A.
KEY WORDS AND DOCUMENT ANALYSIS
DESCRIPTORS
Coelenterata, Jellyfish, MaTlua1s, Taxonomy, Invertebrates, Descriptions
11e.
IDENTIFIERS/OPEN-SNOED TERMS
Cnidaria, Scyphozoa, Stauromedusae, Scyphistomae, Ephyrae, Scypr.omedusae,
Sea net'tles, Sea wasps, Keys,
U.S. Northeast Coast, Systematic lists
!7C.
COSATI ~IE:::LD/GROUP
6C
18.
19. SECURITY CLASS
(This report)
AVAILABILITY STATEMENT
UNCLASSIFIED
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NOAA FORM 25-13
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NO. OF PAGES
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20. SECURITY CLASS
(This report)
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SUPERSEDES ALL PREVIOUS EDITIONS.
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Synopsis of biolosieal data on Pacific ocean perch, Sebtutodes alutUI.
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FOREWORD
Thj~ isslle of the "Circulars" is part of a subseries entitled "Marine Flora and Fauna uf the
Northeastern United States." This subseries will consist of original, illustrated, modern manuals on
the identification, classification. and general biology of the estuarine and ~'Oastal marfine plants and
animals of the Northeastern United States. Mcnuals will be published tlt irregular intervals on as
many taxa of the 'egion as there are spp-.:ialists available to collaborate in their preparation.
The :nanuals are an outgrowth of t:.e widely used "KeyS to Marine Invet1ebrates of the Woods Hole
Region," edited by R. I. Smith, published in 1964, p.nd produced und'er the auspices of the Systematics-Ecolo~ Program, ~,hrine Biological Laboratory, Woods Hole, Mass. Instead of revising the
"Woods Hole Keys," the slaff of the Systematics-Ecology Program decided to expand the geographic
coverage and bathymetric ra"ge p.nd produce the keYf in 'In entirely new set of expanded pub.
lications.
The "Marine Flora and Fauna orthe Northeastern United States" is being prpareu in, :ollaborati':m
with systematic specialists in the United States dnd abroad. Each manual will be based primarily on
recent and ongoing revisionary systematic ,'eser,rch and a fresh examination of tht plant~ and
animals. Each major taxon, treated in a separst,; manual, will include an introduction, :Ilustrated
glossary. uniform originally i'lustrated keys, annotated check list with information when available on
distribution. habitat, life history. and re!at'ld biology, references to the major literature oithe group,
Bnd a systematic index.
These manuals are intended for use by biology students. hiologists, biological oceanographers, infermed l~ym~n, and others wishing to identify coastal organisms for this region, In many instances
the malluals will serve as a guide to additional information about the species or the group.
r.eogra»h.c coverage of the "Marine Flora and Fauna of the Northeastern United States" is plan.
p"d to ir.c1ude organisms from the headwaters of estuaries seaward to approximately the 200·m depth
.;., the c:mtinental shelf from Maine to Virginia, but may vary somewhat with each major taxon and
the inter~sts of collaborators. Whenever possible representative specimens dealt with in the manuals
will be deposited in the reference collections of major museums in the region.
After a sufficient number Df manuals of related taxonomic groups have been published, the
manuals will be revised. grouped, and issued as special volumes. These volumes will thus consist of
compilations of individual manuals within phyla such as th:l Cnidaria, Arthropoc\a, and Moiiuse.., or
of groups of phyla.
CONTENTS
Introduction . .. .
.
Description of orders of 3cyphozoa .
Methods
Collecting . .
Rearing . . .
Pre3crvation
Nematocysts . . .
Nalural historv
Stauromedusae
Scyphistomac .
Ephyrae . . . .
Scyphomedusae
Stinging medu!;ae .
Glossary . . . .
introduction to the keys to tho.! Scyphozoa
Kev tn the Stauromedusae of the northeast coast of the United States
Sy~opsis of ephyrae of Chrysaora, Cyanea. and Rhopi/ema
.
Key to the pelagic; scvphomedusae of Ihe northeast coasl of the United States
Annotated systematic list
Selected bibliography ..
Index to scientific names
Acknowledgments . . . .
Coordinating eJitor's comments
The National Marine Fisheries Service (NMFS) does not approve, recommend or endorse any proprietary product or proprietary materiai
mentioned in this publication. No reference shall be made to NMFS, or
to thb publication furnished by ~MFS, in any advertising or sales promotion which would indicate or imply that NlIIFS approves, recommends
or endorses any proprietary product or proprietary material mentioned
herein, or which has as its purpose an intent to cause directly or indirectly
the advertised product to be used or purchased because of this NMFS
pul>1ication.
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.IR
Marine Flora and Fauna of the Northeastern
United States.
Cnidaria: Scyphozoa
RONALD J. LARSON'
ABS'l"RACT
This manual is an introduction to the sC}llhomedusae found in coastal walers rrom Maine to the
Chesapeake Bay. It includes a discussion oCtheir identifi~ation., collection, rearing, preservation. and
nematocysts. Also included is an introduction to the natural history of th" scyphopolyps and medus.e,
s discussion of stinging 8cYDhomedusac, n glo8~ary or terms, an illustrated synopsis 01 ephyrBe, 1\'D'il·
lustrating key to the scyphomoousae (including the Stauromedusael, an annotated systematic list, a
bibliography of major references, and finally an index.
INTRODUCTION
The class Scyphozoa of the phylum Cnidaria or
Coelenterata as it once was called, are commonly known
as "jellyfish," "sea nettlt-s," and "sea wasps."
Scyphomedusae, as these jellyfish are termed, are the
most conspicuom of the jellylike animals. Some species
reach 1 m or more in diameter and most are very colorful. All have stinging organs, hence the names "sea
nettle" anct "sea wasp." 8cyphomedusae are seasonal;
most species are seen during the warmer months. They
represent the sexual stage of most scyphozoan species.
The asexual stage is a small benthic (bottom-living)
po:yp (scyphopolyp) wh;,:h is perennial. The
scyphopolyp generally buds larval scyphomedusae
(ephyrae) during the spring. The medusae are large by
midsummer.
SCYP!:lOmedusae have a muscular saucer-shaped or
hemispherical bell or umbrella which propels the
medusae through the ','!ilter by contracting and expelling
water behind. Long threadlike tentacles often hang from
the umbrella and are covered by stinging organs
(nematocysts) which are us"d to capture prey. Surroul1.~ing the mouth are ribbol'.like, curtainlike, or
gelatinous arms which transport prey to the mouth.
8cyphopolyps are minute !lnd rarely seen. They are
saclike with a circle of tentaclr;s around the oral end and
are attached to the bottom by a stalklike peduncle.
Seyphozoans usually have a scyphomedusa and a
scypht)polyp stage in their life history but one group
remains as a'polyp only. The 8tauromedusae or stalked
medusae lack a medusa stage. 80me scyphomedusae
lack a polyp stage; Pelagia, an oceanic medusa, has eggs
which, when fertilized, transform directly into a juvenile
medusa and bypass the normal polyp stage.
'Department of Marine Sdence,. University or Puerto Rico. Mayal!Uez,
Puerto Rico 00708; present address: Division of Echinoderms, U.S.
National Museum, Washir,gton, D.C. 20560.
Scyphomedusae are generally regarded as pests
because of their irritating stings, but many are
ecologically important. Chrysaora, well known as the sea
nettle, consumes large numbers of ctenophores which
might otherwise be very detrimental to oyster and clam
populations by feeding on their larvae. Other
scyphomedusae prey on jellyfish which have few
predators and feed on commercially important invl'rtebra(~ and fish larvae.
FiVl' ;,,'ders of Scyphozoa are recognized. The
Stauromedut·.:. or "stalked jellyfish," are not well
know,' even ~hough they may be abundant. They are inactive and cryptically pigmented and are difficult to see
when attached to algae. The Cubomedusae, or sea wasps,
are also infrequently seen but arc infamous because of
their potent sting. The Coronatae, or "crown medusae,"
are mostly deepwater (bathypelagic). The
Semaeostomeae, commonly known us sea nettles, are the
most fam.iliar order, and most shallow-water
scyphozoans belong to this group. The Rhizostomeae,
which lack tentacles, are mostly tropical and are uncommon in New England .....aters.
DESCRIPTION OF
ORDERS OF SCYPHOZOA
Order Stauromedusae-Small benthic scyphozo'lns
which lack a free-swimming medusa stage. Stauromedusae are urn- or funnel-shaped and are attached to the
substrate by a Dtalklike peduncle. In some forms, the
margin is divided into eight arms, the knobbed tentacles
project from the arm tips; in others the tentacles project
from the margin. Modified tentacles, which may act as
anchors or sense organs, occur on the margin hetwpA'1 the
groups of tentacles in some species. The pnads ;,re
paired, leaflike, or folded structures extending along t.:~
length of the calyx. A wl)rmlike planula larva develops
from the zygote and creeps ever the substrate, eventually attaches itself and becomes a juvenile
stauromedusa. Stauromedusae occur mainly in cold,
shallow marine waters. Adult Stauromedusae usually occur only during the summer, and the wintering juveniles
may be difficult to find because of their small size.
Tropical and deepwater forms are rare.
Order Cubomedusae (Fig. I)-Small- to medium-sized
medusae with & somewhat cuboidal or rectangularshaped transparent umbrella. A velarium, or shelflike
projection, encircles the inner margin of the umbrella.
Tentacles vary from four to many and are on bladelike
projections of the bell, termed pedalia. Four rhopalia are
located about midway between the corners of the um-
brella near the margin. The mouth is at the end of a
flask-shaped stomach which hangs within the umbrella.
The life cycle of the Cubomedusae is unlike that of other
scyphozoans. The cubopolyp has several features in common with hydrozoan polyps, e.g., solid tentacles, stenotele nematocysts. Cubopolyps metamorphose directly
and totally into a single medusa, other scyphopolyps
undergo transverse fission producing one to many ephyrae. Werner (1973) has placed the Cubomedusae in ~,;,·~ir
own class, Cubozoa, because of this group's unique characteristics. Cubomedusae are tropical or subtropical me-
/b.,,_~~ ...----gastric
1l!I'+----gonad---------.H-!1fI
, /pedalia----.
velarium
2
gastric cirri
\------~rt_oral a r m - - - - -
3
~;gure
Figure 3.- Semaeoslome medus••
l.-Cubomedusa.
Figure 2.-Coronalemedu68.
FilCW'O 4.- RWzostome med.us",
2
c-.:,',-.
.j.
dusae but are occasionally transported inti) temperate
regions by warm currents. Cubomedusac ure known as
sea wasps because of their virulent sting.
Order Coronatae (Fig. 2)-Small- or medium-sized
medusae with flattened or dome-shaped umbrella which
is divided into two regions by a circular (coronal) furrow.
The tentacles occur between marginal lappets. Radial
thickenings of the margin, termed pedalia, I'ive rise to
the tentacles, lappets, and rhopalia. The gonads are on
four septa which project into the coelenteron. The
Coronatae are mostly bathypelagic and darkly pig·
mented. The coronate polyp, all known species being
placed in the genus Stephanoscyphus, is covered by a
chitinous cuticle wpich has numerOlIS circular annulations (Kramp 1959).
Order Semaeostomeae (Fig. 3)-Large medusae,
usually saucer-shaped. The mouth has four long oral
arms. These may be folded once as in Pelagia, or folded a
number of times as in Cyanea. Tentacles occur between
marj{inal lappets as in the Coronatae, or on the subumhrella. Gonads occur on folds of the gastrodermis. The
polyp stage is small and naked or has a thin cuticle
around its base and can encyst under certain conditions
/Fig. 7). A typical ephyra (Fig. 5) is produced, usually by
polydisc strobilation (Fig. 9). Semaeostomes Ilrc the
most common scyphomedusae in temperate coastal
waters.
Order Rhizostomeae (Fig. 4)-Large medu~ae, usually
with a hemispherical umbrella. Marginal tentacles are
ab~ent, but eight oral arms serve both to capture and
transport food to the ge.stric cavity by Nay of a canal
system. A central mouth is usually absent; instead the
11lllut h is formed by a large number of bifurcating grooves
which lie on the ventral and lateral sides of the oral arms.
The oral Elrms may have terminal appendages which
have a high concentration of n£>matocysts at their tip.
The glmads e,re foids of the gastrodermis as in the
sel11ucostome~;. The polyp stage has a long stalklike
peduncle ',vhich is partially covered with a thin cuticle.
J<:phyrae ,Irf, usually produced by monodisc strobilation
(Fig. 10). Rhizostomes are mostly tropical, with only a
few species extending into temperate waters.
rhopolium
cirri
mouth
5
7
6
gastric
9
8
10
Figure 5.- Ephyra, newly released.
Figure 7.-Semaeostome polyp.
Figure 9.-Semaeostome slrobila.
Figure H.- SfepMnoscyphus.
Figure 8.-Stauromedusa.
Figure 10.- Rhi,oslorne slrobila.
3
with crustaceans being the major part of the food. Berrill
(1962) noted that Lucernaria fed on the amphipc>d
Amphithoe and the small gastrupod Lacuna. At Woods
Hole, Mass., I found mostly harpacticoid copepods and
gammarid amphipods, and a few chironomid fly larvae in
gastric cavities of Craterolophus. Prey, caught by the
tentarles, cause one or all arms to fold over the mouth
which then expands and engulfs the food.
Stauromedusae, like scyphistomae, occur year-round.
Berrill (1962) found that Stauromedusae, along the coast
of Maine, spawn during the summer and then die.
.Juveniles appear during the fall and become sexually
mature by summer.
Apparently Stauromedusae are very sensitive to en·
vironmental CO'lditions and have been difficlllt to raise in
aquaria. Berri11 (1962) remarked that they are becoming
rare in Massachusetts Bay due to pollution.
METHODS
Collecting
To obtain medusae in the best possible condition they
shadd be dipped from the water surface usinp.: a bucket
or other large container. Deepwater species can be
caught in plankton nets equipped with large sample
uuckels.
Scyphopolyps can be obtained by dredgin!:, collecting
at low tide, or diving. Polyps commonly attach to shells,
oysters being a favorite substrate. Polyps will also settle
on "lass slides placed in areas where medusae spawn.
Rearing
Scyphozoans can be raised from embryos to maturity
in the laboratory (Spangenberg 1965). Polyps can be
reared in small dishes and fed newly hatched Artemia.
Planktoni<: .. cages are more difficult to raise, requiring a
large volume of circulating water and suspended food.
Medusae should be fed daily, preferllbly more often.
Artemia can be fed to such plan:<ton-feeding forms as
Aurelia and rhizostome medusae. Ch'J'saora can be fed
ctenophores, Artemia, or other medusae. (See Russell
Scyphistomae
Scyphistomae are the polyp stages of scyphomedusae;
in the semaeostome (Fig. 7) and rhiir.ostome medusae,
they are small, about 2-4 mm high, and naked. or with a
chitinous theca around the peduncle. They are flaskshaped with a single whorl of tentacles and a cruciform
mouth. They are often found in shallow water attached
to shells or other substrates. Those of the coronate
medusae are completely covered by a chitinous theca
and are found in deeper water.
Scyphistomae feed on small crustaceans and cal: he
reared in the laboratory on a diet of Artemia nauplii.
They undergo asexual reproduction to form more polyps.
Some pre·Juce res:stant stage" known as pedal cysts
which can resist temperature extremes at which medusae
are not found (Cargo and Schultz 1967). Scyphistomae
also undergo strobilation, a process which produces from
one to many ephyrae by transverse constrictions of the
polyp's oral region (Figs. 9, 10).
Scyphistomae are difficult to identify. Calder (1971)
used nt:matocvsts to aid in identifying the polyps of
Aurelia, Ch,,'s~ora, and Cyanea. They can a1;:0 be reared
in the laboratory until they strobilate &nd then the
ephyre.e or postephyrae can be identified. Increased
feeding and/or change in waleI' temperature or addition
of thyroxin may induce strobilation.
1970.)
P.·eservation
Scyphozuans should be fixed in a 20% formaldehyde
solution and stored in a 5~c neutralized formaldehyde
solutio:!. Alcohol dehydrates specimens, making them
distorted. Ordinary paper disintegrates in Formalin.
thus labels should be made from high-rag content and
water-resistant paper.
NEMATOCYSTS
Nematocysts occur in all cnidarians. They are
produced by specialized cells, cnidoblasts. A great
amount of work has been done on the cnidoblast and how
the nematocvst functions (Picken and Skaer 1966). Weill
(1934) sho\V~d that the cnidome is taxcl10mically important; and'eeently Calder (l97l) used nematocysts as a
means of identifying polyps of ChrysaoriJ, Cyanea, and
Aurelia. (Also see Calder 1974.)
NATURAL HISTORY
Ephyrne
Ephyrae (Fig. 5) are small, 1-4 mm in diameter, flattened larv,,1 medusae which usually have eight arms. At
the tip of each arm there is a pair of blunt or pointed
lappets, and between the lappets, a rhopalium or sense
organ. The mouth is cruciform and a few gastric cirri
may occur in the coelenteron. Ephyrae feed on plankton
and grow rapidly. They soon appear like small scypho·
medllse.e, at which time they can usually be identified
with certainty. (See synopsis of Ephyrae.)
Stauromedusae
Stauromedusae, contrary to what their name implies.
are really polyplike. Some are permanently attached.
Others can move in a somersaulting motion by adhering
to the substrat.e with the oral eO!d and releasing the pedal
disc, then reattaching the disc at a new location; but
none have been observed to swim.
They attach to algae, sea grass (Zostera), and other
substrates in shallow areas which have adequate water
circulation. They may be abundant, but because of their
inconspicuous coloration and inactivity, they are rarely
seen. Most of the diet consists of other benthic animals,
Scyphomedusae
By being planktonic, scyphomedusae may find food,
4
which is unavaHable to the polyp, and can also di,'·
tribute gametes over an extenilive area thereby in·
creasing the distribution of the species and also insuring
tllat uninhabited substrates are utilized if conditiom are
favorable for the polyp.
Scyphomedusae are usually short-lived. They gr'lW
rapidly and later die due to changes in water temperature or other factors, which for the most part are not
I ..lderstood. Chrysaora ephyrae from the Chesapeake
Bay are released in early June. By July these medusae
arc mature, and most have died by mid-September.
Aurelia and Cyanf!a may live somewhat longer.
The occurrence of some scyphomedusae is related to
water temperature. Cyanea in the Gulf of Maine are first
seen in April or May and are mature by early summer
(Bigelow 1926). Cyanea is found in the Chesapeake Bay
early in the winter and matures by February or March.
Chrysaora occurs only at higher temperatures (about
~O°C) in the Chesapeake Bay in early May, and slightly
later at Woods Hole, Mass. Because of this dependence
on high temperatures, it apparently cannot strobilate in
colder water and is not found north of Cape Cod. Aurelia,
whose range overlaps' the two previous species. occurs
during the wP.rmer months in both the Gulf of Maine and
in the Chesapeake Bay, but appears earlier further
south. In the Gulf of Mexico and off Florida. both Aurelia
and Chrysaora can occur at almost any time. Rhopilema
and Stomolophus also may occur nearly year-round.
Medusae feed on a variety of prey. These include
ctenophOl'p's, planktonic microcrustacea (i.e., copepods,
amphipods, and larvae), larval and small fishes, pelagic
polychaetes, siphonophores, and even other medusae.
Fraser (1969) calculated that a Cyanea less than one-half
meter in diameter would eat over 1.5 million copepods of
the genus CalantL< in its lifetime. One can only guess at
the amount of food eaten by a 2·m Cyanea. Chrysaora
feeds mostly on ctenophores and other medusae. Studies
which I have made in Puerto Rico (unpublished) indicllte
that a Chrvsaora medusa would have to consume about
15 kg of ~tenophore Mnemiopsis in order to attain a
diameter of 150 mm. In Aurelia, food is trapped on the
umbrella and by the short tentacles, and is transferred,
not dire.::tly to the lips, as in most medusae, hut to a food
groove formed by the small velarium. Cilia move the food
which is mixed with mucus to the adradial canals where
it is then transferred to the lips (Southward 1955).
Rhizostomes are filter feeders and many are active
swimmers. In forms such as Rhopilema, water is forced
around the oral arms by the contracting urn brella during
swimming. In Stomolophus, water is forced through the
keellike appendages. Minute tentacles, which line these
structures, filter the suspended food from the water. The
food is then carried to the stomach by the cilia-lined
canals. Stomolophus collected at Beaufort, N.C .. fed
mostly on bivalve veligers and harpacticoid copepods
(author, unpublished).
The gonads of scyphozoans are always near the site of
digestion. In semaeostomes and rhizostomes the gonads
are infoldings of the sl1bumbrella which also bear the
gastric cirri. Some scyphomedusae spawn while still very
small. Chrysaora reared by the author began r-pawning at
the size of 55 mm and spawned almost nightly thereafter.
Fertilization usually takes place within the gastric cavity
or on the surface of the ovary. In Cyanea and Aurelia the
embryos are retained on the oral arms. Aurelia has brood
p<;('kets on the oral arms of the female where em bryos are
r"!ained until the planula is released.
The embryology is quite variable even within the same
speci~s. Segmentation is usually total but gastrulation
depends upon egg size, with ingression in small eggs and
invagination in large ones and by both methods in
medium-sized ova. Usually a planula is produred, whkh
settles and forms a polyp, but very large <.l!;gs tr· ay hypass
both the planula and polyp and form a m.,ciitied ephyra
as they do in Pelagia (BerriIl1949).
Symbiosis is widespread in the Cnidaria and occurs
frequently in the Scyphozoa. There are numerous reports
of juvenile fishes associated with scyphomedusae; Mensueti (1963) discussed these associations. Associations
between juvenile fish and medusae range from the fish
seeking refuge under the umbrella, or between the ten·
tacles, to predation with the larger fish eating bits of the
tentacle or gonad. It is not fully understood how certain
species of fishes, 10'.1-(., Peprilus and Poronotus, are able to
feed on medusae without ill effects, but it is generally
thought that antibodies may be involved and that the
secretion of murus by the fish may prevent the
nematocysts [mm di,charging.
Crustacea are also known to pllrasitize
scyph"medusne: hyperiid amphipods may both !1teal
food from medusae and feed on medusa! tissue. Phillips
et aI. {I 968) discussed the parasitic hahits of the two
species of brachyuran crabs on medusae in the Gulf of
Mexien.
STINGING MEDUSAE
Cllry.'aora is popularly known as the sea nettle because
its sting can be very painful. Cyanea also is a stinger:
Russell (1970) noted that Cyanca was the caust; of a
mysterious death in Conan Doyle's story, "Ad\'en1.ure of
the Lion's Mane." The majoritv of the Cubomedusae are
stingers. as are coronate nll'dusae, some producing a \'ery
severe stillg. Alire/ia, RllClpilema,· Stomolophus.
Phacellophora, and the Stauromedusae ar~ not known to
be severe stingers. Halstead (1965) gave an account of
t he nature of the wounds and medical aspects of stings.
Cnrgo and Schultz (1967) found that when meat tenderizers containing proteolytic enzymes were mixed with
water and applied t, the affected area, the pain was
quickl~' relieved.
GLOSSARY
ancllors (Fig. 81 adhesive organs between arms of some
Stauromeduslle.
calyx flmne!'5haped umbrella of Stnuromedusae.
I.'nidoblar,l c(,ll in which the nematocY3t forms.
t'llidume complement of nematocyst types present.
co"le'llcrull gastric cavity or stomnch.
5
.'.
,I'.""
c~nal furrow (Fig. 2) circular groove which divides
'S'
the exumbrelle. of the coronate
furrow orthe coronate medusae (Fig. 2); bladelike pro·
jections of Cubomedusae to which the tentacles arl;)
attached (Fig, 1).
peduncle (Fig. 8) stalk of polyps, or Stauromedusae.
po(ydisc _.trobi/ntion (Fit!,. 9) several ephyrae produced
from a polyp at the same time.
rlwpalium (Figs. 1, 2) hollow, club-shaped sense organ
l(eneTally located near the umbrella marl(in, usually
with a statocyst and, rarely, an ocellus or ocelli.
scyph,stoma polY!l of scyphozoan medusae excluding
th", Stauromedusae and Cubomedusne.
sensor)' niche open structure partially enclosing rhoplllium in Cubomedusae.
s(mbilo scyphistoma which is forming ephyrae.
str"bilntion process of producing ephyrae.
-'llbL,mbrelia orai or underside of umbrella.
,·..rurilHTI (Fig. 1) sheltlike structure at margin of um1lrella in Cubomedusae ar.d Aurelin.
medlJ~ae.
cross-shaped.
planktonic larva oi Sl'yphozoa except
Gllbornedusae and Staurcrnedusse. Usually with 8
'f
Pairs of arms and 8 rhopaHa.
C' e1Cumbrella aboral or upper surface of umbrella.
-..
gastric cirri or phacelloe (Fig. 1) hollow fingerlike pro·
jections in the coelenteron which aid in digestion.
(appet flaplike structure making up the scalloped marin of the umbrella.
,~'.
,_ odise strobilation (Fig. 10) a single ephyra produced
f. ii'".
r strobila at one time.
~
~:! arm armlike extensions of the corners of the mouth
1' '
which hang from the subumbrella of the Scyphozoa,
to!
,1 in number in the semaeostomes (Fig. :l) and 8 in the
..~
rhizostomes (Fi~. 4).
pl'dalia radial thickenings between lappets and coronal
,'-
",uciform
'. ,..,' phyra (Fig. 5)
i ::
'.,
'4
I"
INTRODUCTION TO THE KEYS TO THE SCYPHOZOA
',.
~ An attempt has been made in constru~ting the keys to
remain unchanged. Because of the delicate nature of
mp,dusae .hey are often injured and parts may be missing
or abnormal. Therefore, in using the key some discretion
must be used for each specimen to determine which
('haracterg are usable.
include those features which 8re most obvious or which
occur in both juvenile and adult specimen.~_ As medusae
!{I'ow, structures such as tentacles usually increase in
number and oral arms become increasingly folded or
complex. while other structures, e.g., rhapaHa, usually
Key to the StBuromedusae
of the Northeast Coast of the United States
With anchors . . . . . . .
:2
Without obvious anchor;;
2
(1)
Anchors larj!r: no shieldlike covering: gonads extend into arm!'
:!
(I)
Anchor" small with shieldlike covering at base: arms shon: gonads do not extend into arms
.. ,
,
3
, , . .
. .. , Thauma/Uscyphus at/anticus
c
·:;;:;:":t:, r-/'~/
.
.
.
b
.. ".
' '·
G
"
'.
'-~,
Fi(ure 11.- Th"umatoacyphWl nl/antic.a.
R. side vie",'
b. \\\ at vif!w
c. nnchor.liidevicw
d. anchor. tront vie'\1:
(j
.
.
d
3 (2) Arms short; anchors bean-shaped; calyx as high as broad
.Haliclystus auricula
(J:
.'.';
.~
.
.:.' ,',".
''':''.
b
C
@d
:,:,t:·::
, ,'.. '
Figure 12.-Hallelyohul auricula.
a. lideview
c. ancbor. Iide view
b. oral view
d. anchor, front view
3 (2) Arms long, anchors trumpet-shaped with short projection at center; calyx broader thail high
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . Halidystu.~ salpinx
b
Figure 13.-HaliclYBtU8 BcJpillX.
8. side view
c. anchor. side view
7
b. oral view
d. anchor. front dew
4 (1) Peduncle and arms long; arms paired; calyx broader than high; largest New England StauroLucernaria quadricornis
medusa
i-.
b
a
Figure 14.-Lucerruvia qruufri<,Dm;'.
a. side view
ba orlll view
4 (l) Peduncle and arms short: arms not paired; calyx higher than brood . . . . . . . Craterolophus canuouulus
b
a
Figure
)1),-
Craterolophu.a CORlIovulus.
8. side "'Iiew
8
b. ofal view
Synopsis of Ephyrae of ChryBaora. Cyanea. Aurelia, and Rhopilema.
(See Russell 1970.)
Newly released ephyrae ca. 2.5 mm in diameter
(Fig. 5)
Morphology of
ephyral arms
Shape of lappets;
pos i tion of rhopalia
lappets slender
pointed tips;
rhopaliar cleft deep
ca. 1/2 arm length
lappets slender
pointed tips;
rhopaliar cleft deep
ca. 1/2 arm length
Chrysaorc.
(Fig. 16a, b)
Cyanea
(Fig. 16c, d)
AUr2lia
(Fig. 16e, f)
Rhopilema
(Fig. 16g, h)
a pair ofnematocYbt
clusters on each arm;
radial canals in arms
short, consists of 1 pair
radial canals in arms
long, consists of 2 pairs
Postephyrae
5 mm in diameter
Gastric cirri
lack
usually 4
lappets broael, tips
blunt, rounded;
rhopaliar cleft short
1/2 arm length
radial canal to
rhopalium only
with 4 or more
lappets slender, tips
p;..\nted; rhopaliar
cleft deep ca. 1/2 al'm
length
radial canals in arms
long, consists of 1 pair
with 4 or more
a
b
e
Morphology
oral arms split; lmlg
ribbonlike;
marginal tentacles;
no ring canal
oral arms not split,
wide and flaring;
subumbrellar
tentacles;
no ring canal
oral arms split;
short; marginal
tentacles; ring
canal present
oral arm split several
times; no marginal
or subumbrellar
tentacles
c
d
9
h
Figure 16.-Epllyrae (a, c, e, g); postepbyrae (b, d, f, h).
e, b Clu-ysaora qu./nquecirrha
c, d Cya"ea capUlata
e, r Aurelia aur/ta
(from I).. Calder 1972, unpuhlished; and Calder 1973)
9
g. h Rhopi/cma verrilli
Key to the Pelagic Scyphomedul'ae
of the Northeast Coast of the United States
1 Umbrella, saucer·shaped, dome-shaped, or hemispheIical; with 8 or more tentacles, or no tentacles;
2
with 8 or more rhopalia . . • • . . . . . . . . . . . . . . . . . . . . . . .
1 Umbrella tall rectangular; 4 tentacles; 4 rhopalia; sensory niche with covering scale above and
Order Cubomedusae
Tamaya haplanema
below; stomach supported by 4 mesenteries
0
....
•
0
0
••
0
0
0
0
0
0
0
•••••
"
\.{-
~~
b
Figure 17.- Tamoyahap/onem<J.
a. side view
bo sensory niche, front view
Umbrella divided into two regions by a deep circular groove; stomach darkly pigmented; bathy·
Order Coronatae
_
pelagic, rarely at surface or coastal
2 (1) Umbrella not divided by circular groove, not darkly pigmented; mostly coastc.l or at surface
2
(1)
0
3
0
0
0
0
•
•
0
•
0
•
0
0
•
0
0
0
•
•
3
•
0
.4
(2) Umbrella flat, disc-shaped; tentacles and rhopalia equal in number, usually 22; gonads button·
like
0
0
0
0
•
0
0
0
0
0
•
0
•
0
•••
0
0
•••••
0
0
•
0
•
0
0
••••••
0
0
0
0
0
••
0
0
•
0
Figure 18o-Atolla W)'lJilIei••ide view.
10
0
.Atalla wyvillei
3
(2) Umbrelll' conical or dome-shaped; usually 3 tentacles between r.hopalia; 4 rhopalia; 12 tentacles: gonads horseshoe-shaped
,
PeriphyUa periphyl/a
FIl(1a'e 19.- Perlph:.llo perlJ)hyllo. aide vie...
4
4
(2) With marginal or subumbrella tentacles; 4 oral arms; distinct mouth; saucer-shaped umbrella
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Order Semaeostomeae
(2) Without tentacles on umbrella; 8 tough gelatinous oral arms; mouth smali or absent; umbrella
usually hemispherical . . . . . . . . . . . . . . . . . . . . . . . . . . Order Rhizostomeae
S (4) With numerous narrow radial canals extending from stomach to margin; ring canal near margin; tentacles either on margin. short and numerous or in 16 linear groups on subumbrella
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Family Ulmaridae
5
9
8
Figure 20.-Famlly Ulmaridae, radial and
ring canala in block.
Figure 21.- C)lan{!Q and family Pelagirlae.
stomach pouches in block.
5
(4) With broad stomach pouches ext.ending to margin; without ring canal; telltacles long 01' 8 Ushaped subumbrellar groups
.
.
11
6
6 (5) Oral arms ribbonlike; tenta_~les on margin
Family Pelagidae . . . . .. .
7
Cyan,'u capillata
6 (5) Oral arms wide, curtainlike; tentacles in 8 V-shaped groups on subumbrella
Q
Figure 22.- Cyane" ""pUlata.
a. olde"lew
b. oral ,"jew, oral arms removed and
rour groupo or lolnlacleo removed
7 (6) With 8 tentacles; exumbrella with prominent wartlike projections; umbrella nearly hemis. . Pclngia noctilum
pherical in shape, usually lees than 100 mm in diameter; oceanic
Figure 23.- Pelagia noctU...,,,.
a. sici.eview
b. oral view, orsl a.rms remov~
7 (6) With more than 8 tentacles; exumbrella smooth; umbrella saucer-shaped, up to 200 mm in
diameter; estuarine or coastal
Chr)'saora quinquecirrha
a
Figure 24.- Chrysaora quinqueclrrha.
a. side view
b. oral view, oral arms romo"ed
8 (;') Marginal ter.<tacles short and very numilrous; oral arms folded, ribbonlike; gonads in 4 horseshoe-shaped groups; B rhopalia . . . . . . . . . . . . . . . . . . . . . .
. A urel ia aurita
a
Figure 25.-AurclialUU'ila.
a••Idevlew
b. orlll view, oral arms relnoved
3 (7) Tent.acles long, in 16 subumhrellar linear groups; oral arms broad. curtainlike; 16 rhopalia
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Phacellophora camtschatica
r-:~
0;.,.,
.' ~.
.
• 7:'"
r~
1\
lb, ~~
I
\
IClMtz; :1
1 'r
~
,\
,
~
I<
\
\ II••
J()I
\ I
I
:
~I;
I
I
I
b
a
Figure 26.-Phae.lloph.ro camt.hallea.
a. aideview
b. oralvle'w, oralarmssndonehalf ortenlacle. remov<'li
13
9 (4) Oral arms free along most of their length; spindlelike '1ppendages hang from oral arms; umbrella hemispherical or flattened . . . . . .
. . . . . . . . . . . . . . . . . . . .Rhopilema uerrilli
/.' .~'J':'
l·
.
f'
.
•
.: . ',
',-•
.
,
,
.
'J.
.
-
~, "
i
,.
Figure '1:1.- Rhop/lema "errilli, aide view.
\::i:,1
:t.
9 (4) Oral arms fused; no spindlelike appf:ndages; umbrella higher than a hemisphere (globular).
. . . . . . . . . . . . . . . .
Stomcilll!?hus meleagris
.:.c
.
..
{,. .
('
Figure 28.-Slomolopllue meleaaria, aide view.
ANNOTATED SYSTEMATIC LIST
HaLiclystus auricula (Rathke 1806)
Massachusetts northward, Northeast, Atlantic,
North Pacific, lower intertidal and subtidal on algae
and Zostera, at Woods Hole most common on brown
algae Scytosiphon. Juveniles pasily confused with
H. salpinx, but arms are very' short. Coloration:
highly variable, brown, greer" red. Berrill (1962);
Mayer (1910).
Haliclystw; salpinx Clark 1863
Massachusetts northward, lower intertidal and slibtidal on algae and Zostera. May be confused with
juvenile H. auricula but with long arms. Coloration:
variable, green, brown. Berrill (1962, 1963).
Lucernaria quadricornis O. F. Muller 1776
Massachusetts northward, Northeast Atlantic, Arctic, lower intertidal and subtidal mostly on
Laminaria. Largest New England stauromedusa
The following list is arranged according to the
classificat:on of Kramp (1961). Distribution notes are
from published records and from museum specimens.
Most of the species listed are represented by specimens
in the U.S. National Museum, Washington, D.C., and
the Gray Museum, Marine Biological Laboratory, Woods
Hole, Mass. Notes on seasonal occurrence, habitat and
life history, and coloration are included. References to
important papers are cited for both the families and for
each species.
Q..ASS
ORDER
SCYPHOZOA
STAUROMEDUSAE
ELEUTHEROCARPIDAE
References: Berrill (1963); Kramp (1961).
FAMILY
14
At least four other coronate medusae may also occni in
the northwestern Atlantic: Linuche unguiculata
(Schwartz), a specimen at the U.S. National Museum,
collected off Cape Hatteras, lat. 35~N, long.
75°W. Nausithoe punctata Kolliksl', a specimen at the
USNM collected off Georges Bank, lat. 40 o N, long.
68°W. Nauphantopsis diomedeae Fewkes, not reported
since the original description, off New England. (Kramp
1961.) Atolla vanhoetfeni Russell, a specimen at the
USNM, collected off Virginia lat. 36°39'1'1. long.
74°39W. (Russell 1970.)
reaching over 50 mm in length. Coloration: highly
variable, brown, green, red. Berrill (1962, 1963).
FAMILY CLEISTOCARPIDAE
References: Herrill (1963); Kramp (1961).
Craterolophus con'Jolvulus (Johnston 1835)
Massachusetts, Northeast Atlantic, intertidal on
Fucus where it attaches near the bladders, inconspicuous b~c6use of its shape and color. Coloration: olive green when attached to Fucu.~. Kramp
(1961).
Thaumatoscypus atlanticus Berrill 1962
Maine, subtidal on Laminaria. This species is unusual in that ocelli are located on the
anchors. Coloration: not known. Harrill (1962,
ORDER SEMAEOSTOMEAE
1963).
FAMILY PELAGIDAE
References: Kramp (1961); Russell (1970).
Chrysaora quinquecirrha (Desor 1848)
Possibly cosmcpolitan, western Atlantic distribution from Cape Cod to the Gulf of Mexito. Abundant during the summer months in estuaries, not
common in the open ocean. Scyphistoma occurs on
oyster shells in upper estuary. Popularly called "sea
nettle" because of its stinghg ability. Coloration:
highly variable, pink, yellow, often with 16 yellowocher or red radiating stripes. Cargo and Schultz
ORDER CUBOMEDUSAE
FAMILY CARYBDEIDAE
References: Bigelow (1938); Kramp (1961).
Tamoya hap/onema·F. Muller 1859
Tropical Atlantic, Gulf of Mexico to Massachusetts.
Probably carried north in the Gulf Steam. Mayer's
(1910) report of this species in Long Island Sound is
the only report of T. haplonema in the New England
coastal waters. Coloration: colorless except tentacles, pink. Bigelow (1938); Phillips and Burke
(1966, 1967).
Pelagia noctiluca (Forskal 1775)
Cosmopolitan open ocean species in warm and temperate oceans. Carried north in the Gulf Stream,
sometimes seen in coastal New J£ngland
waters. LackfJ scyphistoma stage. Coloration:
highly variable, blue, yellow, pink, tentacles
red. Russell (1970).
(1970).
Three other Cubomedusae also occur in the
northwestern Atlantic. Carybdea a/ata Reynaud. a
specimen in the U.S. National Museum (USNM) was
collected off Cepe Hatteras at lat. 35°N, long.
75°W. Carybdea marsupia/is (Linneaus) occurs in the
Gulf Stream near Bermuda but has not been reported on
our coast. Chirapsalmus quadrumanus (Muller) occurs
along U.S. coast south 0f Virginia. Bigelow (1938);
Phillips and Burke (1970).
FAMILY CYAl\iEIDAE
References: Kramp (1961); Russell (1970).
Cyanea capillata (Linnaeus 1758)
Cosmopolitan in colder oceans. Occurs in the Gulf of
Mexico and Florida northward. Most abundant
during winter through spring in the south, summer
to fall in the north (New England waters). Popularly
called "lion's mane" because of the long red-yellow
tentacles. Largest New England medusa reaching a
meter or more in diameter. Coloration: highly
variable, pink, red, yellow, brown. Russell (1970).
ORDER CORONATAE
FAMILY ATOLLIDAE
References: Kramp (1961); Russell (1970).
Atalla wyvil/ei Haeckel 1880
Cosmopolitan in deep oceans. A deep-sea species,
bathypelagic, seldom occuring at the surface or on
the continental shelf. Commonly taken in midwater
samples in deep water usually below 500
m. Coloration: stomach dark red-brown. Russell
FAMILY ULMARIDAE
References: Kramp (1961); Russell (1970).
Aurelia aurita (Linnaeus 1758)
Cosmopolitan species in nearly all oceans except the
Arctic. Popularly known as "moon jelly" because
of the whitish umbrella. Coloration: bell clear;
gonads, tentacles, pink, purple. Russell (1970).
Phacellophora camtschatica Brandt 1838
Occurs north of Cape Cod in the western Atlantic,
probably the entire Pacific coast of the Americas
and in the Mediterranean and off western
Africa. Rarely taken in New England
waters. Coloration: yellowish. Mayer (1910).
(1970) .
FAMILY PERIPHYLLIDAE
References: Kramp (1961); Russell (1970).
Periphylla periphylla (Peron and Lesueur 1809)
Cosmopolitan bathypelagic in deep oceans. Common in midwater trawl samples from deep water. It
has been taken near the surface off New
England. Coloration: stomach dark redbrown. Russell (1970).
15
guiculata. with a brief reexamination of scyphozoan nematocyst
classification. Chesapeake Sci. 15:170-173.
ORDER RHIZOSTOMEAE
CARGO, D. G., and L. P. SHULTZ.
1966. Not... on the biology of the sea nettle, Chrysaora quinquecirrha, in Chesapeake BhY. Cbesapeake Sci. 7:95·100.
1967. Further observati"ns on the biology of the sea nettle and jelly.
fishes in Chespeake Bay. Chesapeake Sci. 8:209-220.
FRASER, J. H.
1969. Experimental feeding of BOrne medusae snd Chaelognatha.
J. ~-i.h. Res. Board -Can. 26:1743-1762.
HALSTEAD, B. W.
1965. Poi•.onous and venomous marine animals of the WNld. I. Invenebrates. U.S. Gov. Print. Off.. Wash., D.C.. 994 p. (see
Phylum Coelenterata, p. 297·371).
KRAMP. P.L.·
1959. Stephanoscyphus (Scyphozoa). Galathea Rep. 1:173-185.
1961. Synopsis of the medusae of the world. J. Mar. BioI. Aaaoc.
U.K. 40:1-469.
MANSUETI, R.
FAMiLY RHIZOSTOMATIDAE
References: Kramp (1961); Mayer (1910).
Rhopilema vurilli (Fewkes 1887)
Occurs in tlle northern Gulf of Mexico, North
Carolina to Connecticut. An uncommon species,
found along ClJast and b mouths of estuaries. Coloration: bell milky; oral arms
brown. Mayer (1910); Calder (1973).
FAMILY STOMOLOPHIDAE
References: Kramp (1961).
Stomolophus meleagris L. Agassiz 1862
Occurs in the western Atlantic from Brazil to Cape
Hatteras and from Panama to San Diego in the
eastern Pacific. Specimens occasionally carried
north to NE'w England waters by the Gulf Stream.
Popularly known as the "cabbage head jelly5sh"
because of its globular shape. Coloration: brown
band at margin. Mayer (1910).
1963. Symbiotic behavior between small fishes and jellyfishes,
with new data on that between the stromateid. Peprilus alepido~
tus, and the Bcyphomedusa, ChrysQora quinqueci"ha. Copeiu
1963:40·80.
MAYER, A.G.
1910.
Medusue of the world.
m. The Sc,)'phomedusse.
CarllcF;ie
Inst. Wash. PubI. 109:499-735.
PHILLIPS. P. J., snd W. D. BURKE.
1970.
The occurrence of sea wasps (Cubomedusse) in MiFosissippi
Bull. Mar. Sci. 20:853·
Sound and the northern Gulf of Mexico.
SELECTED BIBLIOGRAPHY
859.
PHILLIPS, P. J .• W. D. BURKE, and E. J. KEENER.
1969. Observations on the trophic significance af jellyfishes in Mississippi Sound with Quantitative data on the associative behavior
of smaIl fishes with me<lusae. Trans. Am. Fish. Soc. 98:703·712.
PICKSN, L. E. R., and R. J. SKAER.
1966. A review of researches on nematocysts. In W. J. Rees (editor), The Cnidaria and their ~volution. p. 19-50. Zoo!' Soc. Lond.
RUSSELL, F. S.
1970. Thc medusae of the British Isles. II. Pelagic Scyphozoa.
Cambridge Univ. Presa. 264 p.
SOUTHWARD, A. J.
195F·. Observations on the ciliary currents of the jelly-fish Aurelia
aurita L. J. Mar. BioI. Assoc. U.K. 34:201-216.
SPANGENBERG. D. B.
1965. Cultivation of the life stages of Aurelia aurita under cu"·
troIled conditions. J. E~p. Zool. 159:303-318.
WEILL, R.
1934. Contribution a retude des cr daires et de leurs nemalocyste•.
Trav. Stn. Zoo. Wimerem 11:351-701.
\VERNER, B.
BERRILL, M.
1962. The biolollY of three New England Stauromeduaae. with a
dcscripti~n of a ncw spccics. Can. J. Zool. 40:1249-1262.
1963. Compamtive functional morphology of the Stauromedusae.
Can. J. Zool. 41:741-702.
BERRILL, N. J.
1949. Devc(opm<atal analyaia of Scyphomeduaae. Bio', Rev.
(Camh.) 24:393-410.
BIGELOW. H. B.
19'26. Plankton of the off.hore waters of the Gulf of Maine. Boll.
U.S. Bur. Fiah. 40:1-509 (aee p. 340·380).
1938. Plankton of the Bermuda Oceanographic Expeditions, VIII.
Medusae taken during the yeara 1929 and 1930. Zoologica (N.Y.I
23:99·189.
CALDER. D. R.
1971. Nematocysta of polyps of Aurelia, C"rysanra, and Cyanea,
and their utility in idehtification. Trans. Am. Microsc. Soc.
90:269-274.
1973. Laboratory observotions on the life history of Rhopilemo
verrilli (Scyphozoa: Rhizostomeae). Mar. BioI. merl.) 2\:109·
1973.
1974.
New investigations on systematics end evolution of the class
Scyphozoa and the phylum Cnidaria. Publ. Seto M,.r. BioI. Lab.
114.
Nematocysts of the coronate scyphomedusa. L;uuche un·
20:35·60.
16
SYSTEMATIC INDEX
Atolla
vanhoeffeni
wyvillei ..
Atollidae . . .
Aurelia aurita
Caa7a~~e~ .
..
marsupialis
·
Carybdeidae . . . . . . . . .
Chiropsalmus quadrumanus
Chrysaora quinquecirrha ..
Cleistoc8rpldae
Coronatae
Craterolophus COTlvovu!lts
Cubomedusae . . . . . .
Cyanea capillata
Cyaneidae . . . .
Eleutherocarpidae
Halicl;lstus
aurlcuUl
.
.
7, 14
salpinx
.
Linuche unguiculata
Lucemaria quadri.:omis
Nausithoe punctata . . .
Nauphantopsis diomedeae
Pelagia nocti/uca .. . . .
· . . . 15
· .10, 15
· . . . 15
9, 13, 15
Pelagidae
1[;
· .15
8, 14
· .15
· .15
.
Periphylla periphylla
·
15
·
15
·
15
9, 12, 15
· . . . 15
· .10, 15
· . B,15
· .10, 15
9, 12, 15
.. 15
Periphyllidae
Rhizostomatidae
.
Rhizostomeae . . . . .
Rhopilema verrilli . . .
Semaeostomeae . . . .
Stautomedusae
.
.
. . 14
Stomolophus meleagris
Tamayo haplanema . . . . .
Thaumatoscyphus atlanticus
7, 14
Ulmaridae
17
12
11, 12, 15
11,15
·
15
· .13, 15
.
Phacellophora camtschatica
Stomolophidae
·
·
·
.
· . . . 16
· .11, 16
9, 14, 16
.11. 15
. 6,14
. . . 16
.14, 16
.10, 15
... 6, 15
. . . 11, 15
ACKNOWLEDGMENTS
Preparation of the "Marine Flora and Fauna of the Northeastern United States" is being coordinated by the following
Board:
Coordinating Editor: Melbourne R. Carriker, Marine Studieg
Center, College of Marine Studies, University of Delaware,
Lewea, DE 19958.
Editorial Advisers:
Marie B. Abbott. Marine Biolr:gkal Laboratory. Woods
Hole. Mass.
Arthur G. Humes, Bosto" L'ni"ersity Marine Program,
Marine Biological Laboratory, Woods Hole, Mass.
Wesley N. Tiffney, Department of Biology, Boston Univer·
sity, Boston, Ma58,
Ruth D. Turner, Museum of Comparative Zoology, Harvard University, Cambridge, Mass.
Roland L. Wigley, National Marine Fisheries Service Bio·
logical Laboratory, NOAA, Woods Hole, Mass.
Robert T. Wilce, Department of Botany, University ofMas·
sachusetts, Amherst, Mass.
The Board established the format for the "Marine Flora and
Fauna of the Northeastern United States." invites systematists
to collaborate in i.he preparation of manuals, reviews
manuscripts, and advises the Scientific Editor of the National
Marine Fisheries Service.
Thanks go to Marie B. Abbott, Curator of the Gray Museum,
Marine Biological Laboratory, for the loan of specimens and for
assiatance during a short Visit there; S:laUf, Ruetzler, Curator of
the Division of Echinoderms, U,S. National Museum, Washing·
ton. D.C.. for use of collections. dark room, and other facilities:
Maureen E. Downey, also of the Division of Echinoderms, and
Charles E. Cutress of the University of Puerto Rico, Mayaguez,
for encouragement, helpful suggestions, and assistance in proof·
reading; Dale R. Calder, formerly of the Virginia Institute of
Marine Sciences, Gloucester Point, and now at the South
Carolina WiidHfe and Marine Resources Department, Charles·
ton, for critically reading the manuscript, for permission to ex·
amine the scypbozoan polyps maintained at the Virginia Institute of Marine Sciences, and for the loan of unpublished data
and figures; Cadet Hand, Kay Petersen, Roland Wigley, and
Ruth Turner for critically reading the manuscript; and typists
A. Stonework. S. J. Larson, S. Khoury, K. S. Stemler, andJ. S.
Ballantine.
COORDINATING EDITOR'S COMMENTS
Publication of the "Marine FlO!a and Fauna of the Northeastern United States" is most tim2ly in view of the growing universal emphasis on environmental work and the urgent need for
more precise and complete identification of coastal organisms
than has been available. It is ma.,datory, wherever possible,
that organisms be identified accurlltely to species. Accura
scientific names unlock the great quantities of biological information stored in libraries, obviate duplication of research al.
ready done, and often make poasible prediction of attributes of
organisms that heve been inadequately studied.
Ronald J. Larson started his study of medusae during the
summer of 1969 when he worked on the natural history and systematics of Hydromedusae from the Oregon coast. In 1972 he
began working at the U.S. National Museum where use of the
extensive collections and excellent library facilities furthered his
education and initiated an interest in (~Ie systematics of the
Scyphozoa. Currently he is involved with graduate studies at
the University of Puerto Rico, Mayaguez. where he is investigating the functional morphology of scyphomedusae and
aspects of growth and reproduction.
Preparation of this manual was supported in part by a grant
from the Environmental Protection Agency to the Editorial
Board of the "Marine Flora and Fauna of the Northeastern
United States." Work on the "Marine Flora and Fauna of the
Northeastern United States" by the Coordinating Editor is supported by the College of Marine Studies, University of
Delaware.
Manuals are available for purchase from the Superintendent
of Documents, U.S. Government Printing Office, Washington,
D.C. 20402. The manuals so far published in the series are listed
below:
COOK, DAVID G., and RALPH O. BRlNKHURST. Marine flora and fauna of the Northeastern United States. Annelida:
OIigochaeta.
BORROR, ARTHUR C. Marine flora and fauna of the northeastern United States. Protozoa: Ciliophora.
MOUL, EDWIN T. Marine flora and fauna of the northeastern United States. Higher plants of the marine fringe.
McCLOSKEY, LAWRENCE R. Marine flora and fauna of the northeastern United States. Pycnogonida.
MANNING, RAYMOND B. Marine flora and fauna of the northeastern United States. Crustacea: Stomatopoda.
WILLIAMS, AUSTIN B. Marine flora and fauna of the northeastern United States. Crustacea: Decapoda.
POLLACK, LELAND W. Marine flora and fauna of the northeastern United States. Tardigrada.
LARSON, RONALD J. Marine flora and fauna of the ncrtheastern United States. Cnidaria: Scyphozoa.
18
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