NOAA Technical Report NMFS Circular 397

Marine Flora and Fauna ofthe Northeastern United States.Cnidaria: Scyphozoa

Ronald J. Larson

August 1976

U.S. DEPARTMENT OF COMMERCEElliot l. Richardson, Secretary

National Oceanic and Atmospheric AdministrationRob\~rt M. White, Administrator

National Marine Fisheries ServiceRobert W. Schoning, Director

t'ot Sale by the Superintendent of Documents. U.S. GO'o'crnment Printing OfficeWo,hington. D.C. 20~02 - Stock No. 003.017-00387

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12-73) BIBLIOGRAPHIC DATA SHEET NATIONAL OCEANIC AtJD Al'MOSPHERIC ADMINISTRATION

1. NOAA ACCE:;SION NUMBER1

2• 1

3•

RECIPIENT'S ACCESSION NUMBER

NOAA-7610l8l54. TITLE AND SL,'ST:TLE S. REPO~T DATE

Marine Flora and Fauna of the North~astern United States. Au!! 1976Cnidaria: Scyphozoa 6.

7. AUTHOR!SI 8. REPORT NO.

Ronald J. Larson CU.S. National Museum, Washington. DC) NOAA-TR-NMFS Circular ~~79. PERFORMING ORGANIZATION NAME AND ADDRESS 10. PROJECT/TASK NO.

NOM, National Marine Fisheries Service,Seattle, WA, 181 CJ S' 11. CONTRACT/GRANT NO.

Scientific Publications Staff

12, SPONSORING ORGANIZATION NAME AND ADDRESS 13. TYPE OF REPORT AND PERIODCOVERED

Same1'~~h Rent'14.

15. PUBLICATION REFERENCE

NOAA Technical Report NMFS Circular 397, August 1976. 21 p, 28 fig, 24 ref.

16. ABSTRACT

This manual is an introduction to the scyphomedusae found in coastal waters fromMaine to the Chesapeake Bay. It includes a discussion of their identification,

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collection, rearing, preservation, and nematocysts. Also included is an introductionto the natural history of the scyphopolyps and medusa,e, a discussion of stingingscyphomedusae, a glossary of terms, an illustrated synopsis of ephyrae, anillustrating key to the scyphomedusae (including the Stauromedusae), an annotatedsystematic list, a bibliography of major refer~nces, and an index.(Author)

17. KEY WORDS AND DOCUMENT ANALYSIS

17A. 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

6C18. AVAILABILITY STATEMENT 19. SECURITY CLASS 21. NO. OF PAGES

(This report) d-5UNCLASSIFIED20. SECURITY CLASS 22. PRICE

(This report) -:>-~OUNCLASSIFIED ")..

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NOAA TEPHNICAL REPORTS

National Marine Fisheries Service, Circulars

T!le major responsibiJities of the National Marine Fisheries ServiC"e lNMFS) arl' to monitor and aue" the abundance and Reographic distribution of fbitJeryresources. to under:'lland and predict nuctuations in the quantity and di.!ltribution of the~e resourt'es. Iond to establish leveb for optimum use or the resourteS.NMFS is also charged with the dew:lopmen;. and implementation of policies for managing "ationa.! fL!lhing grounds. development Ind en(oN!ement of domestic'f15heries regulations, surveillance of foreign Hshing oU United Slales ("OMta1 waters, and the development and enforcement of intern. tiona: fishery 8gret!:nentsand ~liries. NMFS also M5ists the fishing industry through marketing service and economic analysis programs, and mortK!lg~ insurance and vessel constructionSIJbSldi~s. It collects, analyzes, and publishes statistics on various ~hases of the industry.

The NOAA Technical Report NMFS eiRC !terie! continue! a !eries that has been in exist~nce since 1941. The Circulars are technical publications of general interestintended to aid conservation and management. Publications th~t. review in considerable detail and at a hi,h technical level certain broad areas of rt-..ear('h appear in thi5s~ries. T~chnical p,~rs originatinR in economics studies and from managemeru. investigations appear In the Circular s~ries.

NOAA T«hnical Reports ~MFS CIRf" are a\iailable rree in limited numbers to governm~ntal agendes. both Federal and State. The)' art' also available inexchange for other scientific ...nd technical publil"ations in the marine sciences. Individual copies may be obtained lunless otherwis.e nOlA'd) from 083, Tec:hnic:l1Information Divisioll' Envirllnmental Scien,'""e Information Center, NOAA. Wllshin!{ton. D.C. 20235. Recent Circulars are:

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i!lII!I

IIi\

I

I

315. Synopsis of biologi~al data on the chum salmon. Oncorh1l1ldlu$ kelt!IWal~.uml 119'~. By Richard G. Bakkala. lI.arch 1970. iii + 89 p.. 15 fop .• 51tab!{'s.319, Dureau or Commerl'ial Fi'St;eries Great Lakes Fi.IIlt'lery Laboratory, AnnArbor, Michigan. By Bureau of Co:nmercial Fishuies. March 197(),8 p.• 7 figs,

330. EASTROPAC Atlas: Vol•. )·7. Camlog No. I 49.4:3301Ivo\.l 11 vol•.Availabl~ from fl)e Superintendent of Documents. U.S. Government PrintingOffice. WashinKlon. D.C. 20402.

331. Guidelint'5 for the processing of hot·smo!.:ed chuh. By H. L. SCl1RTan, J.T. Graikoski. and J. A. Emerson. January H~·,U. iv + 23 p., 8 fiJVi .• 2 tablt"s.

:3.12. Pacific hake. 02 articles hy 20 l\utlJond March 1970, iii + 152 p .• 72figs., 47 tables.

333. Recommended practices for vc.:isl'i sanitation and fisll handHng. ByEdJitar W. BC"\lman anti Alfred LarsL·n. March 1970. hI + 27 p.• 6 rigs.

335. Pro~ress I""t'port of the Bureau of Commercial Fisheries Cenler forEstuarine and Me"lhaden Research. Pt'sticide Field Station. Gulf Brp('ze. ,.~ta .•lise" I )'car 1969. By the Laboratory !Ower. AUFrU5t 1970, iii + 33 p., 29 rigs"12 tables.

336. The northern rUT seal. By r.:alph C. Raker. Ford Wilke. and C. H.JWIlMBalun. April 1970. iii + 19 p .. 13 ro"•.

331, ProKram of Div\sion of Economic Re~earch. llure.llU of CommeITialFisheries, fi!lical )'ear 1%9. By Oiv!sion of Economic Research. April 1970. iii+ :?9 p., 12 fiKS., 7 tables.

338. Bureau of Commercial Fisheries Biol:>Kieal Laboratory. Auke 8a)'.AlaskD.. By Bureau or C.,mmercial Fisheries, June 1970, 8 p., 6 riR's.

339. Salmon research at Ice Harbor Dam. By W~.sl •.·y J. Eta'i. April 1970,6p.. 4 figs.

340. Bllreau of Commercial Fisheries Technological LaboraLory, Glouct'ster.Ma"lsachusetls, By Hureau of Commercial F'i'iheries. June 1970. 8 p., 8 figs.

~1. Rt-'port of the Bureau of Commercial Fisheries Biological Laboratory,Beaufort. N.C., for the fiSCRI year end inK June 30. 1968. RJo· the Laboratorystaff. August W70. iii + 24 p., 11 Ci~s .• 16 tllbles.

34.2. Report of the Burt'3u of Commercial Fisheries Biological Laboratory.St. Petersburg Beach. Florida. fiscal year 1969. By the Laboratory staff.August 1970. iii + 22 p.• 20 figs., 8 tables.

343. Report Df the Bureau of Commercial Fisheries Biological Laboratory.Galveston, Totus. fiscal yf!aT 1969. By the Laboratory stafr. August H)iO. ii,+ 39 p.• 28 figs .. 9 tables.

34·1. Bureau of Commercial Fisheries Tmpical Atlantic aiological LaboratoryprQRress in research 1965·69. Miami, Florida. By Ann Wee~s. Oc:tober 1970. iv+ 65 p.. 53 fi~s ..

346. Sportsman's guide to handling, smoking, and preserving Great Lakescoho ~almon. By Shearon Dudley, J. T. Graikoski, H. L. Seagran. and Pall) M.Earl. September 1970. iii + ~8 p.. 15 figs.

347. Synopsis of biolosieal data on Pacific ocean perch, Sebtutodes alutUI.By Richard L. Major and Herbert H. Shippen. De-cember 1970, iii + 38 p.• 31figs.• It table,.

349. LJ!liL' of abstrat'ts lind 'lummarics as communication devices in technicalartidl:s. By F. Drure Sanford. l"cbruary 1971. iii + 11 p., I fig.

3..'10. Resl!srch in fiscal year 1969 at the Bureau of Cummercial Fi5herit~s

Biolol{ical Laborator.r. B..~aufort. N.C. By the Laboratory stafL November 1970.ii + 49 p.. 21 figs,. 17 tables.

3,')1. Burenu of Commercial F;sheries E)"plorl1tory Fishin,li, and Gear R~,earch

fiase, (Jasca~oula, Mississippi, July 1. 1967 to ..Tunc 30. 1969. By Harvey R.Bullis. Jr. and John R. Thompson. November 1970, iv + 29 p., 29 figs .• 1t3hll>.

:lS2. lJpslrl~3m pas~.aKe of anlldromou!i fish through navigation locks and u~e

of tht! strt'am for spawning and nursery habitat, Capp F("ar Rivt-'r, N.C.,n_li2-(lii. II)' Paul R. Nichols and Darl·ell F... Loud~r. October 1970. iv + 12 p..!l fi,,;!'i., .1, tahles,

:~.,'i6. FloatinK laboratory for stud)' of aquatic orhanisms and their environrm'nl. By G('Orge R. Snydl'r. Theodore II. B1ahm, and RiJocrt J, McConnell.Mn)' 1971, iii + It. p., 11 fiKS ..

:~61. Rt·Kinn..1 and other related aSpt'Ct5 of sbellfish ["onsumption - some­prdiminary findings frum tht, 1969 Consume r I>anel Survey. lJ}' MorLon M.Mill.'r and I>arrl'1 t\. Nash. June 1971, iv + 18 p.. 19 figs .. 3 Lahles, 10 apps.

:162. Ht'search \'e!'sels of lht· National Marint' Fi!lhcries Scrvj..t'. By Roh.'rt S.Wnlf. t\ugu.st 1971, iii + 46 p.• 2,') fiKS .• :J tabl£'s. Fo!" sale b)' theSupl'Tintt'ndl'nt of DlX·uments. U.S. Go\,("rnnl{-'nt Printing OffiCl', WashinK!on,n.c. 20·102.

;kj.1, Hi~tory and onelopmcnt or surf dam har\ll'!ltiug gear. By Phillip S.Parker. Ortoht'r 1971, iv + 15 p., 16 fiR'S. For sail! b)· the Supt'rintcnderlr ofnOl.~Umt>nt5. u.s, Go\'('rnment Printing Ofrice, Wnshinfrloll. D.C. 2;:,402.

:165. Procl'ssing Et\STHOPAC STD dnLD and thl' con'ilruction of vfl"'licnlh'mp(>raturc and !':alinit)" sloctions by co~putt'r. By Forrest R. Miller and}\('nnl'th A. Bliss. February 1972, iv + 11 p., 8 figs., 3 appl'ndix figs. Forsale hy tht' ~~uperintenrlent of J)ocument~. U.S. GO\'ernment Printinl{ omct'.\\'u!lhinKton. D.C. 'lO402.

aBO. Key to field identification of IIndromuu.s jU\'f'nil£' snlmonids in thf> PacificNorthwest. By Robt'rt J. M,..Conneli and George R. :"';n)·df'r. January 1972. i\l+ 6 po, 4 fil':s. f~()r salp by lht· SU~t'rintl'ndent of DoclJmcnts. U.S.Gn\,"('rnmt'nt PrintinK OHiC:l', Wa~hin~rton, D.C. 20402.

:167. l':nKlDel'rin~ economic model f('lr fish prmt'in conc{ntration pro{'ess('~. ByK. K. Almenas. L. C. Durilla. R. C. Ernst. J. W. Gelltr)'. M. 3. Hale, and J.M. Mnrcht'llo. O<"lober 1972. iii + \75 p., t' fi~s.• 6 tahh·~. ror saIl" by theSuperintendent of DOC'uments, U.S, GovcrlllTlPnL Printin~ Offil."l'. WashinR\on,D.G. ~0402.

af'JR. COflpl'rative Gulf of Mexico f1stU:Lrinl' invl'ntory lind stud)·. Florida:Phasl' I. area dcscriptio!1. By J. Kneeland ~ ..Nult)·. William N. LindaU, Jro,and Jamt'~ 1-;. SJo·kt's. Novembt>r 1972. \Iii + 126 p., 46 fiRS., 62 tahles. Forsale by the Superintendent of Documents. U.S. Government Printing Office,W..,hinglon. D.C. 20402.

3lJ9. Field guide to thl' anKlefishf!s (Pomacunthi::laet in thl' ..... t'stern Atlantir.By Henry A. Feddern. November 1972, iii + 10 p.. 17 figs .• ror sale b}' theSuperintt'ndcnt or Documents. u..S. Gon·roment PrintinK Office, Wa~hington,D.C. 2o.102.

Continued on inside back cover.

FOREWORD

Thj~ isslle of the "Circulars" is part of a subseries entitled "Marine Flora and Fauna uf theNortheastern United States." This subseries will consist of original, illustrated, modern manuals onthe identification, classification. and general biology of the estuarine and ~'Oastal marfine plants andanimals of the Northeastern United States. Mcnuals will be published tlt irregular intervals on asmany 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 HoleRegion," 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 geographiccoverage 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':mwith systematic specialists in the United States dnd abroad. Each manual will be based primarily onrecent and ongoing revisionary systematic ,'eser,rch and a fresh examination of tht plant~ andanimals. Each major taxon, treated in a separst,; manual, will include an introduction, :Ilustratedglossary. uniform originally i'lustrated keys, annotated check list with information when available ondistribution. 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 instancesthe 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 ofestuaries seaward to approximately the 200·m depth.;., the c:mtinental shelf from Maine to Virginia, but may vary somewhat with each major taxon andthe inter~sts of collaborators. Whenever possible representative specimens dealt with in the manualswill 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, themanuals will be revised. grouped, and issued as special volumes. These volumes will thus consist ofcompilations of individual manuals within phyla such as th:l Cnidaria, Arthropoc\a, and Moiiuse.., orof groups of phyla.

CONTENTS

Introduction . .. . .Description of orders of 3cyphozoa .Methods

Collecting . .Rearing ...Pre3crvation

Nematocysts ...Nalural historv

StauromedusaeScyphistomac .Ephyrae ....Scyphomedusae

Stinging medu!;ae .Glossary ....introduction to the keys to tho.! Scyphozoa

Kev tn the Stauromedusae of the northeast coast of the United StatesSy~opsis of ephyrae of Chrysaora, Cyanea. and Rhopi/ema .Key to the pelagic; scvphomedusae of Ihe northeast coasl of the United States

Annotated systematic listSelected bibliography ..Index to scientific namesAcknowledgments ....Coordinating eJitor's comments

The National Marine Fisheries Service (NMFS) does not approve, rec­ommend or endorse any proprietary product or proprietary materiaimentioned in this publication. No reference shall be made to NMFS, orto thb publication furnished by ~MFS, in any advertising or sales pro­motion which would indicate or imply that NlIIFS approves, recommendsor endorses any proprietary product or proprietary material mentionedherein, or which has as its purpose an intent to cause directly or indirectlythe advertised product to be used or purchased because of this NMFSpul>1ication.

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Marine Flora and Fauna of the NortheasternUnited 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 theChesapeake Bay. It includes a discussion oCtheir identifi~ation., collection, rearing, preservation. andnematocysts. 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, abibliography of major references, and finally an index.

INTRODUCTIONThe class Scyphozoa of the phylum Cnidaria or

Coelenterata as it once was called, are commonly knownas "jellyfish," "sea nettlt-s," and "sea wasps."Scyphomedusae, as these jellyfish are termed, are themost conspicuom of the jellylike animals. Some speciesreach 1 m or more in diameter and most are very color­ful. All have stinging organs, hence the names "seanettle" anct "sea wasp." 8cyphomedusae are seasonal;most species are seen during the warmer months. Theyrepresent the sexual stage of most scyphozoan species.The asexual stage is a small benthic (bottom-living)po:yp (scyphopolyp) wh;,:h is perennial. Thescyphopolyp generally buds larval scyphomedusae(ephyrae) during the spring. The medusae are large bymidsummer.

SCYP!:lOmedusae have a muscular saucer-shaped orhemispherical bell or umbrella which propels themedusae through the ','!ilter by contracting and expellingwater behind. Long threadlike tentacles often hang fromthe umbrella and are covered by stinging organs(nematocysts) which are us"d to capture prey. Sur­roul1.~ing the mouth are ribbol'.like, curtainlike, orgelatinous arms which transport prey to the mouth.

8cyphopolyps are minute !lnd rarely seen. They aresaclike with a circle of tentaclr;s around the oral end andare attached to the bottom by a stalklike peduncle.

Seyphozoans usually have a scyphomedusa and ascypht)polyp stage in their life history but one groupremains as a'polyp only. The 8tauromedusae or stalkedmedusae lack a medusa stage. 80me scyphomedusaelack a polyp stage; Pelagia, an oceanic medusa, has eggswhich, when fertilized, transform directly into a juvenilemedusa 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 pestsbecause of their irritating stings, but many areecologically important. Chrysaora, well known as the seanettle, consumes large numbers of ctenophores whichmight otherwise be very detrimental to oyster and clampopulations by feeding on their larvae. Otherscyphomedusae prey on jellyfish which have fewpredators and feed on commercially important invl'r­tebra(~ and fish larvae.

FiVl' ;,,'ders of Scyphozoa are recognized. TheStauromedut·.:. or "stalked jellyfish," are not wellknow,' even ~hough they may be abundant. They are in­active and cryptically pigmented and are difficult to seewhen attached to algae. The Cubomedusae, or sea wasps,are also infrequently seen but arc infamous because oftheir potent sting. The Coronatae, or "crown medusae,"are mostly deepwater (bathypelagic). TheSemaeostomeae, commonly known us sea nettles, are themost fam.iliar order, and most shallow-waterscyphozoans belong to this group. The Rhizostomeae,which lack tentacles, are mostly tropical and are un­common in New England .....aters.

DESCRIPTION OFORDERS OF SCYPHOZOA

Order Stauromedusae-Small benthic scyphozo'lnswhich lack a free-swimming medusa stage. Stauromedu­sae are urn- or funnel-shaped and are attached to thesubstrate by a Dtalklike peduncle. In some forms, themargin is divided into eight arms, the knobbed tentaclesproject from the arm tips; in others the tentacles projectfrom the margin. Modified tentacles, which may act asanchors or sense organs, occur on the margin hetwpA'1 thegroups of tentacles in some species. The pnads ;,repaired, leaflike, or folded structures extending along t.:~

length of the calyx. A wl)rmlike planula larva developsfrom the zygote and creeps ever the substrate, even­tually attaches itself and becomes a juvenilestauromedusa. Stauromedusae occur mainly in cold,

shallow marine waters. Adult Stauromedusae usually oc­cur only during the summer, and the wintering juvenilesmay be difficult to find because of their small size.Tropical and deepwater forms are rare.

Order Cubomedusae (Fig. I)-Small- to medium-sizedmedusae with & somewhat cuboidal or rectangular­shaped transparent umbrella. A velarium, or shelflikeprojection, encircles the inner margin of the umbrella.Tentacles vary from four to many and are on bladelikeprojections of the bell, termed pedalia. Four rhopalia arelocated about midway between the corners of the um-

brella near the margin. The mouth is at the end of aflask-shaped stomach which hangs within the umbrella.The life cycle of the Cubomedusae is unlike that of otherscyphozoans. The cubopolyp has several features in com­mon with hydrozoan polyps, e.g., solid tentacles, steno­tele nematocysts. Cubopolyps metamorphose directlyand totally into a single medusa, other scyphopolypsundergo 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_oralarm-----

3

~;gure l.-Cubomedusa.

Figure 2.-Coronalemedu68.

2

Figure 3.-Semaeoslome medus••

FilCW'O 4.- RWzostome med.us",

c-.:,',-.

dusae but are occasionally transported inti) temperateregions by warm currents. Cubomedusac ure known assea wasps because of their virulent sting.

Order Coronatae (Fig. 2)-Small- or medium-sizedmedusae with flattened or dome-shaped umbrella whichis divided into two regions by a circular (coronal) furrow.The tentacles occur between marginal lappets. Radialthickenings of the margin, termed pedalia, I'ive rise tothe tentacles, lappets, and rhopalia. The gonads are onfour septa which project into the coelenteron. TheCoronatae are mostly bathypelagic and darkly pig·mented. The coronate polyp, all known species beingplaced in the genus Stephanoscyphus, is covered by achitinous cuticle wpich has numerOlIS circular an­nulations (Kramp 1959).

Order Semaeostomeae (Fig. 3)-Large medusae,usually saucer-shaped. The mouth has four long oralarms. These may be folded once as in Pelagia, or folded anumber of times as in Cyanea. Tentacles occur betweenmarj{inal lappets as in the Coronatae, or on the subum­hrella. Gonads occur on folds of the gastrodermis. The

.j.polyp stage is small and naked or has a thin cuticlearound its base and can encyst under certain conditions/Fig. 7). A typical ephyra (Fig. 5) is produced, usually bypolydisc strobilation (Fig. 9). Semaeostomes Ilrc themost common scyphomedusae in temperate coastalwaters.

Order Rhizostomeae (Fig. 4)-Large medu~ae, usuallywith a hemispherical umbrella. Marginal tentacles areab~ent, but eight oral arms serve both to capture andtransport food to the ge.stric cavity by Nay of a canalsystem. A central mouth is usually absent; instead the11llluth is formed by a large number of bifurcating grooveswhich lie on the ventral and lateral sides of the oral arms.The oral Elrms may have terminal appendages whichhave a high concentration of n£>matocysts at their tip.The glmads e,re foids of the gastrodermis as in thesel11ucostome~;. The polyp stage has a long stalklikepeduncle ',vhich is partially covered with a thin cuticle.J<:phyrae ,Irf, usually produced by monodisc strobilation(Fig. 10). Rhizostomes are mostly tropical, with only afew species extending into temperate waters.

5

gastric

8

rhopolium

cirri

mouth

6

9

7

10

Figure 5.-Ephyra, newly released.

Figure H.- SfepMnoscyphus.

Figure 7.-Semaeostome polyp.

Figure 8.-Stauromedusa.

3

Figure 9.-Semaeostome slrobila.

Figure 10.-Rhi,oslorne slrobila.

METHODS

Collecting

To obtain medusae in the best possible condition theyshadd be dipped from the water surface usinp.: a bucketor other large container. Deepwater species can becaught in plankton nets equipped with large sampleuuckels.

Scyphopolyps can be obtained by dredgin!:, collectingat low tide, or diving. Polyps commonly attach to shells,oysters being a favorite substrate. Polyps will also settleon "lass slides placed in areas where medusae spawn.

Rearing

Scyphozoans can be raised from embryos to maturityin the laboratory (Spangenberg 1965). Polyps can bereared in small dishes and fed newly hatched Artemia.Planktoni<: .. cages are more difficult to raise, requiring alarge 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 asAurelia and rhizostome medusae. Ch'J'saora can be fedctenophores, Artemia, or other medusae. (See Russell1970.)

P.·eservation

Scyphozuans should be fixed in a 20% formaldehydesolution and stored in a 5~c neutralized formaldehydesolutio:!. Alcohol dehydrates specimens, making themdistorted. Ordinary paper disintegrates in Formalin.thus labels should be made from high-rag content andwater-resistant paper.

NEMATOCYSTS

Nematocysts occur in all cnidarians. They areproduced by specialized cells, cnidoblasts. A greatamount of work has been done on the cnidoblast and howthe nematocvst functions (Picken and Skaer 1966). Weill(1934) sho\V~d that the cnidome is taxcl10mically impor­tant; and'eeently Calder (l97l) used nematocysts as ameans of identifying polyps of ChrysaoriJ, Cyanea, andAurelia. (Also see Calder 1974.)

NATURAL HISTORY

Stauromedusae

Stauromedusae, contrary to what their name implies.are really polyplike. Some are permanently attached.Others can move in a somersaulting motion by adheringto the substrat.e with the oral eO!d and releasing the pedaldisc, then reattaching the disc at a new location; butnone have been observed to swim.

They attach to algae, sea grass (Zostera), and othersubstrates in shallow areas which have adequate watercirculation. They may be abundant, but because of theirinconspicuous coloration and inactivity, they are rarelyseen. Most of the diet consists of other benthic animals,

4

with crustaceans being the major part of the food. Berrill(1962) noted that Lucernaria fed on the amphipc>dAmphithoe and the small gastrupod Lacuna. At WoodsHole, Mass., I found mostly harpacticoid copepods andgammarid amphipods, and a few chironomid fly larvae ingastric cavities of Craterolophus. Prey, caught by thetentarles, cause one or all arms to fold over the mouthwhich then expands and engulfs the food.

Stauromedusae, like scyphistomae, occur year-round.Berrill (1962) found that Stauromedusae, along the coastof Maine, spawn during the summer and then die..Juveniles appear during the fall and become sexuallymature by summer.

Apparently Stauromedusae are very sensitive to en·vironmental CO'lditions and have been difficlllt to raise inaquaria. Berri11 (1962) remarked that they are becomingrare in Massachusetts Bay due to pollution.

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 achitinous theca around the peduncle. They are flask­shaped with a single whorl of tentacles and a cruciformmouth. They are often found in shallow water attachedto shells or other substrates. Those of the coronatemedusae are completely covered by a chitinous thecaand are found in deeper water.

Scyphistomae feed on small crustaceans and cal: hereared 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 cystswhich can resist temperature extremes at which medusaeare not found (Cargo and Schultz 1967). Scyphistomaealso undergo strobilation, a process which produces fromone to many ephyrae by transverse constrictions of thepolyp's oral region (Figs. 9, 10).

Scyphistomae are difficult to identify. Calder (1971)used nt:matocvsts to aid in identifying the polyps ofAurelia, Ch,,'s~ora, and Cyanea. They can a1;:0 be rearedin the laboratory until they strobilate &nd then theephyre.e or postephyrae can be identified. Increasedfeeding and/or change in waleI' temperature or additionof thyroxin may induce strobilation.

Ephyrne

Ephyrae (Fig. 5) are small, 1-4 mm in diameter, flat­tened larv,,1 medusae which usually have eight arms. Atthe tip of each arm there is a pair of blunt or pointedlappets, and between the lappets, a rhopalium or senseorgan. The mouth is cruciform and a few gastric cirrimay occur in the coelenteron. Ephyrae feed on planktonand grow rapidly. They soon appear like small scypho·medllse.e, at which time they can usually be identifiedwith certainty. (See synopsis of Ephyrae.)

Scyphomedusae

By being planktonic, scyphomedusae may find food,

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 insuringtllat uninhabited substrates are utilized if conditiom arefavorable 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 notI ..lderstood. Chrysaora ephyrae from the ChesapeakeBay are released in early June. By July these medusaearc mature, and most have died by mid-September.Aurelia and Cyanf!a may live somewhat longer.

The occurrence of some scyphomedusae is related towater temperature. Cyanea in the Gulf of Maine are firstseen in April or May and are mature by early summer(Bigelow 1926). Cyanea is found in the Chesapeake Bayearly 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 slightlylater at Woods Hole, Mass. Because of this dependenceon high temperatures, it apparently cannot strobilate incolder water and is not found north of Cape Cod. Aurelia,whose range overlaps' the two previous species. occursduring the wP.rmer months in both the Gulf of Maine andin the Chesapeake Bay, but appears earlier furthersouth. In the Gulf of Mexico and off Florida. both Aureliaand Chrysaora can occur at almost any time. Rhopilemaand Stomolophus also may occur nearly year-round.

Medusae feed on a variety of prey. These includectenophOl'p's, planktonic microcrustacea (i.e., copepods,amphipods, and larvae), larval and small fishes, pelagicpolychaetes, siphonophores, and even other medusae.Fraser (1969) calculated that a Cyanea less than one-halfmeter in diameter would eat over 1.5 million copepods ofthe genus CalantL< in its lifetime. One can only guess atthe amount of food eaten by a 2·m Cyanea. Chrysaorafeeds mostly on ctenophores and other medusae. Studieswhich I have made in Puerto Rico (unpublished) indiclltethat a Chrvsaora medusa would have to consume about15 kg of ~tenophore Mnemiopsis in order to attain adiameter of 150 mm. In Aurelia, food is trapped on theumbrella and by the short tentacles, and is transferred,not dire.::tly to the lips, as in most medusae, hut to a foodgroove formed by the small velarium. Cilia move the foodwhich is mixed with mucus to the adradial canals whereit is then transferred to the lips (Southward 1955).

Rhizostomes are filter feeders and many are activeswimmers. In forms such as Rhopilema, water is forcedaround the oral arms by the contracting urn brella duringswimming. In Stomolophus, water is forced through thekeellike appendages. Minute tentacles, which line thesestructures, filter the suspended food from the water. Thefood is then carried to the stomach by the cilia-linedcanals. Stomolophus collected at Beaufort, N.C .. fedmostly on bivalve veligers and harpacticoid copepods(author, unpublished).

The gonads of scyphozoans are always near the site ofdigestion. In semaeostomes and rhizostomes the gonadsare infoldings of the sl1bumbrella which also bear thegastric cirri. Some scyphomedusae spawn while still very

5

small. Chrysaora reared by the author began r-pawning atthe size of 55 mm and spawned almost nightly thereafter.Fertilization usually takes place within the gastric cavityor on the surface of the ovary. In Cyanea and Aurelia theembryos are retained on the oral arms. Aurelia has broodp<;('kets on the oral arms of the female where em bryos arer"!ained until the planula is released.

The embryology is quite variable even within the samespeci~s. Segmentation is usually total but gastrulationdepends upon egg size, with ingression in small eggs andinvagination in large ones and by both methods inmedium-sized ova. Usually a planula is produred, whkhsettles and forms a polyp, but very large <.l!;gs tr· ay hypassboth the planula and polyp and form a m.,ciitied ephyraas they do in Pelagia (BerriIl1949).

Symbiosis is widespread in the Cnidaria and occursfrequently in the Scyphozoa. There are numerous reportsof juvenile fishes associated with scyphomedusae; Men­sueti (1963) discussed these associations. Associationsbetween juvenile fish and medusae range from the fishseeking refuge under the umbrella, or between the ten·tacles, to predation with the larger fish eating bits of thetentacle or gonad. It is not fully understood how certainspecies of fishes, 10'.1-(., Peprilus and Poronotus, are able tofeed on medusae without ill effects, but it is generallythought that antibodies may be involved and that thesecretion of murus by the fish may prevent thenematocysts [mm di,charging.

Crustacea are also known to pllrasitizescyph"medusne: hyperiid amphipods may both !1tealfood from medusae and feed on medusa! tissue. Phillipset aI. {I 968) discussed the parasitic hahits of the twospecies of brachyuran crabs on medusae in the Gulf ofMexien.

STINGING MEDUSAE

Cllry.'aora is popularly known as the sea nettle becauseits sting can be very painful. Cyanea also is a stinger:Russell (1970) noted that Cyanca was the caust; of amysterious death in Conan Doyle's story, "Ad\'en1.ure ofthe Lion's Mane." The majoritv of the Cubomedusae arestingers. as are coronate nll'dusae, some producing a \'erysevere stillg. Alire/ia, RllClpilema,· Stomolophus.Phacellophora, and the Stauromedusae ar~ not known tobe severe stingers. Halstead (1965) gave an account oft he nature of the wounds and medical aspects of stings.Cnrgo and Schultz (1967) found that when meat ten­derizers containing proteolytic enzymes were mixed withwater and applied t, the affected area, the pain wasquickl~' relieved.

GLOSSARY

ancllors (Fig. 81 adhesive organs between arms of someStauromeduslle.

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.

.'.,I'.""c~nal furrow (Fig. 2) circular groove which divides'S' the exumbrelle. of the coronate medlJ~ae.

,'- ",uciform cross-shaped.'. ,..,' phyra (Fig. 5) planktonic larva oi Sl'yphozoa excepti :: Gllbornedusae and Staurcrnedusse. Usually with 8

'f '4Pairs 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 mar-

'., I"in 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 mouth1' ' 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

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, usuallywith a statocyst and, rarely, an ocellus or ocelli.

scyph,stoma polY!l of scyphozoan medusae excludingth", Stauromedusae and Cubomedusne.

sensor)' niche open structure partially enclosing rho-plllium 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 um­

1lrella in Cubomedusae ar.d Aurelin.

remain unchanged. Because of the delicate nature ofmp,dusae .hey are often injured and parts may be missingor abnormal. Therefore, in using the key some discretionmust be used for each specimen to determine which('haracterg are usable.

INTRODUCTION TO THE KEYS TO THE SCYPHOZOA

',.~ An attempt has been made in constru~ting the keys toinclude those features which 8re most obvious or whichoccur in both juvenile and adult specimen.~_As medusae!{I'ow, structures such as tentacles usually increase innumber and oral arms become increasingly folded orcomplex. while other structures, e.g., rhapaHa, usually

Key to the StBuromedusaeof the Northeast Coast of the United States

With anchors . . . . . . .

Without obvious anchor;;

:2

2 (1) Anchors larj!r: no shieldlike covering: gonads extend into arm!' 3

:! (I) Anchor" small with shieldlike covering at base: arms shon: gonads do not extend into arms.. , , , , . . . .. , Thauma/Uscyphus at/anticus

c

dG'''''·'." .

'-~, .

b·:;;:;:":t:, r-/'~/ ... . .. ".

Fi(ure 11.- Th"umatoacyphWl nl/antic.a.R. side vie",' b. \\\ at vif!wc. nnchor.liidevicw d. anchor. tront vie'\1:

(j

3 (2) Arms short; anchors bean-shaped; calyx as high as broad .Haliclystus auricula

(J:.'; ~ .'. .C.:.' ,',".

''':''.

b

@:,:,t:·:: d, ,'..'

Figure 12.-Hallelyohul auricula.a. lideviewc. ancbor. Iide view

b. oral viewd. 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 viewc. anchor. side view

7

b. oral viewd. anchor. front dew

4 (1) Peduncle and arms long; arms paired; calyx broader than high; largest New England Stauro-medusa Lucernaria quadricornis

i-.

ab

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

a bFigure )1),- Craterolophu.a CORlIovulus.

8. side "'Iiew b. ofal view

8

Synopsis ofEphyrae of ChryBaora. Cyanea. Aurelia, and Rhopilema.(See Russell 1970.)

Newly released ephyrae ca. 2.5 mm in diameter(Fig. 5)

Chrysaorc.

(Fig. 16a, b)

Cyanea

(Fig. 16c, d)

AUr2lia

(Fig. 16e, f)

Rhopilema

(Fig. 16g, h)

Shape of lappets;pos ition of rhopalia

lappets slenderpointed tips;rhopaliar cleft deepca. 1/2 arm length

lappets slenderpointed tips;rhopaliar cleft deepca. 1/2 arm length

lappets broael, tipsblunt, rounded;rhopaliar cleft short1/2 arm length

lappets slender, tipsp;..\nted; rhopaliarcleft deep ca. 1/2 al'mlength

Morphology ofephyral arms

a pair ofnematocYbtclusters on each arm;radial canals in armsshort, consists of 1 pair

radial canals in armslong, consists of 2 pairs

radial canal torhopalium only

radial canals in armslong, consists of 1 pair

Gastric cirri

lack

usually 4

with 4 or more

with 4 or more

Postephyrae5 mm in diameter

Morphology

oral arms split; lmlgribbonlike;marginal tentacles;no ring canal

oral arms not split,wide and flaring;subumbrellartentacles;no ring canal

oral arms split;short; marginaltentacles; ringcanal present

oral arm split severaltimes; no marginalor subumbrellartentacles

a b c d

e 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'aeof the Northeast Coast of the United States

1 Umbrella, saucer·shaped, dome-shaped, or hemispheIical; with 8 or more tentacles, or no tentacles;with 8 or more rhopalia ..••....................... 0 • 0 0 •• 0 0 0 0 0 0 0 ••••• " 2

1 Umbrella tall rectangular; 4 tentacles; 4 rhopalia; sensory niche with covering scale above andbelow; stomach supported by 4 mesenteries Order Cubomedusae Tamaya haplanema

Figure 17.- Tamoyahap/onem<J.a. side view

.... \.{-

~~

b

bo sensory niche, front view

2 (1) Umbrella divided into two regions by a deep circular groove; stomach darkly pigmented; bathy·pelagic, rarely at surface or coastal 0 0 0 0 0 • • 0 • 0 • 0 0 • 0 0 0 • • • Order Coronatae _ 3

2 (1) Umbrella not divided by circular groove, not darkly pigmented; mostly coastc.l or at surface 0 .4

3 (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 0 .Atalla wyvillei

Figure 18o-Atolla W)'lJilIei••ide view.

10

3 (2) Umbrelll' conical or dome-shaped; usually 3 tentacles between r.hopalia; 4 rhopalia; 12 ten-tacles: gonads horseshoe-shaped , PeriphyUa periphyl/a

FIl(1a'e 19.-Perlph:.llo perlJ)hyllo. aide vie...

4 (2) With marginal or subumbrella tentacles; 4 oral arms; distinct mouth; saucer-shaped umbrella. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Order Semaeostomeae 5

4 (2) Without tentacles on umbrella; 8 tough gelatinous oral arms; mouth smali or absent; umbrellausually hemispherical . . . . . . . . . . . . . . . . . . . . . . . . . . Order Rhizostomeae 9

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 8

Figure 20.-Famlly Ulmaridae, radial andring 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 U-shaped subumbrellar groups . .

11

6

6 (5) Oral arms ribbonlike; tenta_~les on margin Family Pelagidae . . . . .. . 7

6 (5) Oral arms wide, curtainlike; tentacles in 8 V-shaped groups on subumbrella Cyan,'u capillata

Figure 22.- Cyane" ""pUlata.a. olde"lew

Q

b. oral ,"jew, oral arms removed androur groupo or lolnlacleo removed

7 (6) With 8 tentacles; exumbrella with prominent wartlike projections; umbrella nearly hemis-pherical in shape, usually lees than 100 mm in diameter; oceanic . . Pclngia noctilum

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 indiameter; 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 horse-shoe-shaped groups; B rhopalia . . . . . . . . . . . . . . . . . . . . . . . A urelia 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-:~ .' ~. • 7:'".0;.,.,r~ 1\

I lb, ~~~1 IClMtz :1

\'r ;~ I<

,\ ,

\ \ II•• J()I\ I ~I;

I : II I

aFigure 26.-Phae.lloph.ro camt.hallea.

a. aideview b. oralvle'w, oralarmssndone-halfortenlacle. remov<'li

13

b

9 (4) Oral arms free along most of their length; spindlelike '1ppendages hang from oral arms; um-brella hemispherical or flattened . . . . . . . . . . . . . . . . . . . . . . . . . .Rhopilema uerrilli

/.~.' 'J':'. ',-

f' . •• , , 'J.

. . - .l·· .: ... ',~, "

i \::i:,1

,. :t.

Figure '1:1.- Rhop/lema "errilli, aide view.

9 (4) Oral arms fused; no spindlelike appf:ndages; umbrella higher than a hemisphere (globular).. . . . . . . . . . . . . . . . Stomcilll!?hus meleagris

{.

.:.c .

. .,.

('

ANNOTATED SYSTEMATIC LIST

The following list is arranged according to theclassificat:on of Kramp (1961). Distribution notes arefrom published records and from museum specimens.Most of the species listed are represented by specimensin the U.S. National Museum, Washington, D.C., andthe Gray Museum, Marine Biological Laboratory, WoodsHole, Mass. Notes on seasonal occurrence, habitat andlife history, and coloration are included. References toimportant papers are cited for both the families and foreach species.

Q..ASS SCYPHOZOAORDER STAUROMEDUSAE

FAMILY ELEUTHEROCARPIDAEReferences: Berrill (1963); Kramp (1961).

14

Figure 28.-Slomolopllue meleaaria, aide view.

HaLiclystus auricula (Rathke 1806)Massachusetts northward, Northeast, Atlantic,North Pacific, lower intertidal and subtidal on algaeand Zostera, at Woods Hole most common on brownalgae Scytosiphon. Juveniles pasily confused withH. salpinx, but arms are very' short. Coloration:highly variable, brown, greer" red. Berrill (1962);Mayer (1910).

Haliclystw; salpinx Clark 1863Massachusetts northward, lower intertidal and slib­tidal on algae and Zostera. May be confused withjuvenile H. auricula but with long arms. Coloration:variable, green, brown. Berrill (1962, 1963).

Lucernaria quadricornis O. F. Muller 1776Massachusetts northward, Northeast Atlantic, Arc­tic, lower intertidal and subtidal mostly onLaminaria. Largest New England stauromedusa

reaching over 50 mm in length. Coloration: highlyvariable, brown, green, red. Berrill (1962, 1963).

FAMILY CLEISTOCARPIDAEReferences: Herrill (1963); Kramp (1961).Craterolophus con'Jolvulus (Johnston 1835)Massachusetts, Northeast Atlantic, intertidal onFucus 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 1962Maine, subtidal on Laminaria. This species is un­usual in that ocelli are located on theanchors. Coloration: not known. Harrill (1962,1963).

ORDER CUBOMEDUSAE

FAMILY CARYBDEIDAEReferences: Bigelow (1938); Kramp (1961).

Tamoya hap/onema·F. Muller 1859Tropical Atlantic, Gulf of Mexico to Massachusetts.Probably carried north in the Gulf Steam. Mayer's(1910) report of this species in Long Island Sound isthe only report of T. haplonema in the New Englandcoastal waters. Coloration: colorless except ten­tacles, pink. Bigelow (1938); Phillips and Burke(1970).

Three other Cubomedusae also occur in thenorthwestern Atlantic. Carybdea a/ata Reynaud. aspecimen in the U.S. National Museum (USNM) wascollected off Cepe Hatteras at lat. 35°N, long.75°W. Carybdea marsupia/is (Linneaus) occurs in theGulf Stream near Bermuda but has not been reported onour coast. Chirapsalmus quadrumanus (Muller) occursalong U.S. coast south 0f Virginia. Bigelow (1938);Phillips and Burke (1970).

ORDER CORONATAE

FAMILY ATOLLIDAEReferences: Kramp (1961); Russell (1970).

Atalla wyvil/ei Haeckel 1880Cosmopolitan in deep oceans. A deep-sea species,bathypelagic, seldom occuring at the surface or onthe continental shelf. Commonly taken in midwatersamples in deep water usually below 500m. Coloration: stomach dark red-brown. Russell(1970) .

FAMILY PERIPHYLLIDAEReferences: Kramp (1961); Russell (1970).

Periphylla periphylla (Peron and Lesueur 1809)Cosmopolitan bathypelagic in deep oceans. Com­mon in midwater trawl samples from deep water. Ithas been taken near the surface off NewEngland. Coloration: stomach dark red­brown. Russell (1970).

15

At least four other coronate medusae may also occni inthe 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 theUSNM collected off Georges Bank, lat. 40oN, long.68°W. Nauphantopsis diomedeae Fewkes, not reportedsince the original description, off New England. (Kramp1961.) Atolla vanhoetfeni Russell, a specimen at theUSNM, collected off Virginia lat. 36°39'1'1. long.74°39W. (Russell 1970.)

ORDER SEMAEOSTOMEAE

FAMILY PELAGIDAEReferences: 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, notcommon in the open ocean. Scyphistoma occurs onoyster shells in upper estuary. Popularly called "seanettle" because of its stinghg ability. Coloration:highly variable, pink, yellow, often with 16 yellow­ocher or red radiating stripes. Cargo and Schultz(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£nglandwaters. LackfJ scyphistoma stage. Coloration:highly variable, blue, yellow, pink, tentaclesred. Russell (1970).

FAMILY CYAl\iEIDAEReferences: Kramp (1961); Russell (1970).

Cyanea capillata (Linnaeus 1758)Cosmopolitan in colder oceans. Occurs in the Gulf ofMexico and Florida northward. Most abundantduring winter through spring in the south, summerto fall in the north (New England waters). Popularlycalled "lion's mane" because of the long red-yellowtentacles. Largest New England medusa reaching ameter or more in diameter. Coloration: highlyvariable, pink, red, yellow, brown. Russell (1970).

FAMILY ULMARIDAEReferences: Kramp (1961); Russell (1970).

Aurelia aurita (Linnaeus 1758)Cosmopolitan species in nearly all oceans except theArctic. Popularly known as "moon jelly" becauseof the whitish umbrella. Coloration: bell clear;gonads, tentacles, pink, purple. Russell (1970).

Phacellophora camtschatica Brandt 1838Occurs north of Cape Cod in the western Atlantic,probably the entire Pacific coast of the Americasand in the Mediterranean and off westernAfrica. Rarely taken in New Englandwaters. Coloration: yellowish. Mayer (1910).

ORDER RHIZOSTOMEAE

FAMiLY RHIZOSTOMATIDAEReferences: Kramp (1961); Mayer (1910).

Rhopilema vurilli (Fewkes 1887)Occurs in tlle northern Gulf of Mexico, NorthCarolina to Connecticut. An uncommon species,found along ClJast and b mouths of es­tuaries. Coloration: bell milky; oral armsbrown. Mayer (1910); Calder (1973).

FAMILY STOMOLOPHIDAEReferences: Kramp (1961).

Stomolophus meleagris L. Agassiz 1862Occurs in the western Atlantic from Brazil to CapeHatteras and from Panama to San Diego in theeastern Pacific. Specimens occasionally carriednorth to NE'w England waters by the Gulf Stream.Popularly known as the "cabbage head jelly5sh"because of its globular shape. Coloration: brownband at margin. Mayer (1910).

SELECTED BIBLIOGRAPHY

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.I23:99·189.

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 Rhopilemoverrilli (Scyphozoa: Rhizostomeae). Mar. BioI. merl.) 2\:109·114.

1974. Nematocysts of the coronate scyphomedusa. L;uuche un·

16

guiculata. with a brief reexamination of scyphozoan nematocystclassification. Chesapeake Sci. 15:170-173.

CARGO, D. G., and L. P. SHULTZ.1966. Not... on the biology of the sea nettle, Chrysaora quinque­

cirrha, 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. In­

venebrates. U.S. Gov. Print. Off.. Wash., D.C.. 994 p. (seePhylum 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.

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. Copeiu1963:40·80.

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 MiFosissippiSound and the northern Gulf of Mexico. Bull. Mar. Sci. 20:853·859.

PHILLIPS, P. J .• W. D. BURKE, and E. J. KEENER.1969. Observations on the trophic significance af jellyfishes in Mis­

sissippi Sound with Quantitative data on the associative behaviorof 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 (edi­

tor), 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.

1973. New investigations on systematics end evolution of the classScyphozoa and the phylum Cnidaria. Publ. Seto M,.r. BioI. Lab.20:35·60.

SYSTEMATIC INDEX

Atollavanhoeffeniwyvillei ..

Atollidae ...Aurelia aurita

Caa7a~~e~ . ..marsupialis

Carybdeidae . . . . . . . . .Chiropsalmus quadrumanusChrysaora quinquecirrha ..Cleistoc8rpldae .Coronatae .Craterolophus COTlvovu!ltsCubomedusae . . . . . .Cyanea capillataCyaneidae ....EleutherocarpidaeHalicl;lstus

aurlcuUl

· ... 15· .10, 15· ... 159, 13, 15

· 1[;· 15· 15· 159, 12, 15· ... 15· .10, 15· . B,15· .10, 159, 12, 15

.. 15

. . 14

7, 14

17

salpinx .Linuche unguiculataLucemaria quadri.:omisNausithoe punctata . . .Nauphantopsis diomedeaePelagia nocti/uca .. . . .Pelagidae .Periphylla periphyllaPeriphyllidae .Phacellophora camtschaticaRhizostomatidae .Rhizostomeae .....Rhopilema verrilli . . .Semaeostomeae . . . .Stautomedusae .Stomolophidae .Stomolophus meleagrisTamayo haplanema . . . . .Thaumatoscyphus atlanticusUlmaridae .

7, 14· .158, 14· .15· .15

· 12· 11, 12, 15· 11,15

· 15· .13, 15· ... 16· .11, 169, 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 followingBoard:

Coordinating Editor: Melbourne R. Carriker, Marine StudiegCenter, 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, Har­

vard University, Cambridge, Mass.Roland L. Wigley, National Marine Fisheries Service Bio·

logical Laboratory, NOAA, Woods Hole, Mass.Robert T. Wilce, Department ofBotany, 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, reviewsmanuscripts, and advises the Scientific Editor of the NationalMarine Fisheries Service.

Thanks go to Marie B. Abbott, Curator of the Gray Museum,Marine Biological Laboratory, for the loan of specimens and forassiatance during a short Visit there; S:laUf, Ruetzler, Curator ofthe 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, andCharles 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 ofMarine Sciences, Gloucester Point, and now at the SouthCarolina 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 dataand figures; Cadet Hand, Kay Petersen, Roland Wigley, andRuth Turner for critically reading the manuscript; and typistsA. 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 formore precise and complete identification of coastal organismsthan has been available. It is ma.,datory, wherever possible,that organisms be identified accurlltely to species. Accurascientific 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 oforganisms that heve been inadequately studied.

Ronald J. Larson started his study of medusae during thesummer of 1969 when he worked on the natural history and sys­tematics of Hydromedusae from the Oregon coast. In 1972 hebegan working at the U.S. National Museum where use of theextensive collections and excellent library facilities furthered hiseducation and initiated an interest in (~Ie systematics of the

Scyphozoa. Currently he is involved with graduate studies atthe University of Puerto Rico, Mayaguez. where he is inves­tigating the functional morphology of scyphomedusae andaspects of growth and reproduction.

Preparation of this manual was supported in part by a grantfrom the Environmental Protection Agency to the EditorialBoard of the "Marine Flora and Fauna of the NortheasternUnited States." Work on the "Marine Flora and Fauna of theNortheastern United States" by the Coordinating Editor is sup­ported by the College of Marine Studies, University ofDelaware.

Manuals are available for purchase from the Superintendentof Documents, U.S. Government Printing Office, Washington,D.C. 20402. The manuals so far published in the series are listedbelow:

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

Co u. S. GOV~RNHtNT PQINTING Of:~ICE: 1976-697.707/108 "EGlON 10

370. Collecting and proce5sing data on fish t"IiCKS and larvae in the· CalilcrniaCurrent re~on. By David Kramer, Mary J. Kalin. Eliubeth G. Stevens.Jamt.'s R. Thrailkill. and James It Zweifel. November 1912. iv + 38 p.• 38figs .• 2: tablt.'s. FQr salt.· by the Superintendt'nt of D()("umenl.'l. U.s. Governme-nt Prinlinl( Office. WashinK\on. D.C. 20402.

371. Ocean fL-lht"ry mllnaKl"mt"nt; Discussions and rest"a~h. By Adam A.SokolMki ,editor). 111 papeJ"!il, :?4 au' hors. I April 1973. "'i + 173 p.. 38 riKS.•:r.! table~. 7 appendix tables.

372 "'ishery publicntion!o. calendar yur 1971: l.i~t~ and indue!. By ThomuA. Manar. Odober 19T1. i ... + 2-1 p .• 1 fig. For !!oalti' uy the Superintendent ofDocuments. U.S. Governmt"nt PrintinJ,r Orrin'. WashinKlon, n.c. 20402.

374. Marine.' flora and (aunH of the.' northrnlt'rn Unitt'd States. Annelida:OHgochaeu. By David G. Cook and Ralph O. Brinkhunt. May 1973. Hi + 23p., 82 fi~s. For ~ale by the.' Superintt>ndt'nt of Dnc·umenb. U.S. Go\'rrnmentPri:"ltinlr\' Oflict.·. Washin..,.on, D.C. 2O~02.

315. ~('w I·olyrharla from Bt'3ufort. with a ke-r to lSll spe::es ffforded fromNorth Carolina. By John H, Day. July 1973. xiii + I.tO p., IB r:,i(!I .• 1 tahle.For ~Ic b)' tht· Superintt'ndent of llocumpnts. U.S. r;o..It.'r~In';.·:1l PrintingOffir('. Wa....hin~on. 11.C. :'>Q40:?

:.176. Bottom·water tt.·mpl'raturt·s on tht, continental ~ht'lf. NOVA :;,('oti~ to~t·w Jt.·r"i.t'!·. Oy John R. Colton. Jr. ;snd Ruth It. Stoddard. JU:It,' !iH:t iii +5:') p.• 15 fiKS., 12 a.ppendix tables. For !itnle b)' the Supt'rintendr~1 ofOuc·umt.·nt.... U.S, Gon·rnmt.'nt Printinj{ Office. WD.ShinKf,on. D.C. 20402.

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/e;

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tlf thl' mariru.' frin~t.'. By I::dwin T. Maul. SI'ptt'mbt'r 1973. iii + 60 p., lOSfigs. 1o"or sale by the Supuintt'ndpnt of Documflnts. U.S. Government PrintillKOffict·. Wol~hinh'1on. I).C. ~>()~02.

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;l.'ii, Marini' nura and fauna of thl' nurtht>uu·rn L~nitt'd Stah·!rI. rru~la{'t'A;

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