Reference:Biol Hull 193:S-I.V(August, 1497) Chrysaom achlyos, a Remarkable New Species of Scyphozoan from the Eastern Pacific JOEL W. MADRATVIIND1,G.LICSAA-RAGNON4GAENRDSHDWAIVNI2D, JAO.SEBPLHOOWM.3BURNETT1, , ^NaturalHistoryMuseumofLosAngelesCounty, LosAngeles, California90007;2CabrilloMarine Aquarium.SanPedro, California90731;^UniversityofMarylandatBaltimore,Departmentof Dermatology, Baltimore,Maryland21201;and^ChesapeakeBiologicalLaboratory, Solomons,Maryland20688 Abstract. Anenormousnewspeciesofscyphozoanjel- cies belonged in the genus Chrysaora but clearly was lyfish, Chrysaoraachlyos, isdescribed from the eastern noneoftheknownspeciesinthatgenus(R.Larson,pers. Pacific.Thedescriptionisbasedprimarilyoncolorpho- comm.; see also Larson, 1990; Larson and Arneson, tographs and video footage of living animals and the 1990). Interestingly,thespecieshadbeenphotographed morphology offour specimens collected in 1989. The beforebutnotdescribed;MartinandKuck(1991)attrib- naturalhistory,lifecycle,andsporadicappearanceofthe utedphotographsappearinginCrowder(1926)andHal- speciesall areunknown. Thespeciesappeared mostre- stead (1965) to this species. The Crowder photograph centlyinlargenumbersin 1989buthasappearedatleast waslabeledonly"blackjellyfish"andwaspublishedina twicepreviously inthiscentury;published photographs popularmagazine.ThephotographsinHalstead(1965), (unlabeled or incorrectly identified) appeared in 1926 oneofwhich wasreprinted in Halstead(1992), deserve and 1965. The species iseasily distinguished by itssize special mention. Both photographs were incorrectly and coloration from other known species in thegenus, identifiedasCyaneacapillala,aspeciesnotatallsimilar all of which are considerably smaller. Morphological tothenewChrysaorainaspectsotherthansize,andwere charactersaredescribed,andlimiteddataonnematocyst attributedtoP.Saunders,whoworkedexclusivelyinthe typesarepresented.Becauseofthesizeofthenewspecies Pacific. Halstead'slabel indicatingthatthephotographs and the known potency ofthe sting ofcongeners, we weretakenoffthecoastofFloridaisthereforeobviously mention briefly the possible consequences of human anerror. Since 1989theanimal hasbeenthesubjectof contact. several popular articles (see synonymy below) and at least one film (Nature's Seasons in the Sea, National Introduction Geographic/Thirteen-WNETandHowardHallProduc- talat6ispyienlunmfvnmelgIediettnsttaaahmMobectlneeerlaiwbteoxldleaeosnoaisusfctawc1sw2kis9nioat0me8tdihae9(hnwte,ntrMaoceeeabasolxid4ernnalpl0tdtole1iliuerrcvanisnatmernmcdtslgareiyeliniioisinnftnbdncnehe.rtulp(idKhiirmTMceuvbekaabhecessfreeodkeiitrreb,rniasblsevdeldnpiiel1eocvanlda9cfiogr9infwdsnemdualp1sfssaas)eoiK.rloucegrsuTfmetix.mcho,shmtekseecLeedw,tryonnilvntpnsdt1yehghhe9rCab,od9dtyaua1zaltpd)tdoribee.hiaferklserloenCithlprasciojauncapndenpirhcteil-a-s--e--- tcd(cobtmirorhfaeCoileenDmnllLMdessleebos,os.eAcslp)taol1Ti.isfMA9tohih9anAtnerno0lghsrter)leeaite.wecslnhsteeeaee.iprsrnaeeeAnfctClAaihmlhiocqmoaemtutauudeenlfasentantslher.stlyadaiwtacWoeu(rlifremneelLesa1tAira9hasnCgir8ereeneM9peeN,)onanam,SowaituaarasnumnanrsrdcbdnieaoendtlnpgoPrhdh,efaHosiodtintoatrsienosnootolfhg,donyiorrs,aioayflnaCnpwotsdalMhiuilyetrotuvidhhfifspse,doipheoureeuCfnanuccnaellieim--wysa-s oftheoralarmsremain. Since the Martin and Kuck paper, concerted efforts Received 1 May1997;accepted2June1997. have been made to properlydescribe the new scypho- NEW PACIFIC SCYPHOZOAN Figure1. ('lirv.\u<truachlyii.i.newspecies,in\iliiphotographtakenbymembersofHowardHallPro- ductionsfilmcrewoftCoronadoNorte. IslasdeLosCoronados.northernBajaCalifornia, Mexico,July 1989.Sideviewofintactanimalinfrontofdiver;belldiameterestimatedat I m.Notedistinctivecolor pattern,lengthoftentacles,andespeciallylengthofcombinedoralarms(estimatedtobe6m),whichtrail tobottomrightofphotograph(terminatingbeyondframeofphotograph).UsedbypermissionofHoward HallProductions. zoan on the basis of nematocyst ultrastructure, allo- specimen was preserved in ethyl alcohol and was not zyme electrophoresis, immunology, and morphology. subjectedtohistologicalexamination. Unfortunately,becauseofthepaucityofspecimensand theirpoorcondition, a full description ofthisinterest- Results siinbglea.nWimealdebsycrmiboedeitrhnersetapnrdiamradrislaypopanrtehnetlbyasiissnooftgprooss-s OrderSemaestomeae morphology,color,and limited informationon nema- F(a'mliirlyysaoPrealaagcihildyaoes, newspecies tocysttypes. Black Jellyfish. Crowder, 1926: 190 (photograph withcaptiononly). MaterialsandMethods Cyanea capil/ala (Linnaeus). Halstead, 1965: plate 43 (misidentincation oftwo P. Saunders photographs, Specimenswerecollected eitherdead ordyingin the incorrectly attributed to coast of Florida). Halstead, surfzonefrom severalareasalongthesouthernCalifor- 1992: plate 86 (one ofthe same photographs that ap- niacoastinlate 1989(seeMaterialexaminedunderRe- pearedinHalstead, 1965). sultsandalso Martin and Kuck, 1991). Photographsof Giant pelagic jellyfish. Hall, 1990: 29, 2 unnum- livinganimals(Figs. 1,2)andvideofootagetakenoffthe beredfigures. coastofCoronadoNorte,IslasdeLosCoronados,north- Purple jellyfish. Cranston, 1993: 2-3 (photograph ern BajaCalifornia, Mexico, in July 1989, were kindly withcaptiononly). senttousbyHoward HallofHowardHall Productions. LargeJellyfish. Whiteman, 1996: 19(photographby AllthreepreservedLACMspecimenswereinitiallyfixed H.Hall). in formalin and latertransferred to 70% ethyl alcohol. Unlabeled figure. Straus and Lisowski, 1997: 509 These specimens, preserved in 1989, were subjected to (photographbyH.Hall). histological examination 7 years later. Tissue samples Chrysaorasp. Larson, 1990:549(Table1). Martin were taken from the edges ofthe bell and lappets, and and Kuck, 1990: 64 (abstract). Larson and Arneson, wereembeddedinparaffinandstainedwithhematoxylin 1990: 130. Kuck and Martin, 1990: 48 (two pho- andeosinaswellasMasson'strichrome.ThesoleCMA tographsbyH.Hall). MartinandKuck, 1991:89,figs. 10 J. W. MARTIN ETAL 24cm. found dead floatingon surface in thesurfzone, VeniceBeach.California,collected25August 1989byJ. MartinandH. Kuck. Description Bell:Hemispherical,fleshy, large;upto25cmdiame- terinpreservedspecimensexamined;reachingestimated 1 m bell diameter in life (based on still photographs, video footage, and eyewitness accounts; see Figs. 1. 2. andphotographsin Hall, 1990;Anderson, 1992;Camp- bell, 1992;Cranston, 1993).Surfaceofbellsmooth,lack- ingnematocystwarts(seeRussell, 1970). Marginallappet's:32;allsquarewithroundedcorners. Marginal SCUM' organs: 8; each with two rhopaliar cones, onefromexumbrellaandextendinginward,one betweenandformedbyjunctionofadjacentlappetbor- dersanddirectedupward.Openingofexumbrellarcone stronglytriangularin somespecimens(e.g.. LACM 89- 206.1). Both cones wide and deep; e.g.. in specimen LACM 89-205.1 exumbrellar cone approximately mm mm 7.8 diameterand 5.5-7.0 deep; rhopaliar(lap- mm pet) cone slightly narrower but deeper, 7.2-6.9 in diameter and 8.0-9.0mm deep. Rhopalia interradial and perradial. and set into cones approximately 1 cm frommarginofbell. Tentacles:24;whitetolight pinkin color(from vari- ousstillphotographsandvideofootage),eacharisingbe- tween lappets, in sets of 3 between adjacent rhopalia. Figure2. Chrysaomaclilynx,newspecies,ventralviewofdifferent Eachwithbasalswelling,withonesidemoreprotrusive specimenphotographedonsamedayandinsamelocationasinFigure than other (Fig. 2). Long, extending to perhaps l/2 to I.withdiverinspectingoralarmsandundersideofbell.Notenumber 3/4lengthoforalarms;delicate. and arrangement oftentaclesand spiralingand interlockingoforal Alaniihriuni.Thick,stiff,partlyencircling4interradial arms.UsedbypermissionofHowardHallProductions. subumbrellarostia,throughthecenterofwhichprotrude thickfingerlikeprojectionsbearinggonadsinmaturein- dividuals. 1-3,table 1. Anderson, 1992:26(fourphotographsby Oralarms:4;perradial,extremelylarge,marginsfrilly, (Ha.llHpahllo)t.ogrCaapmhpsbbeyllH..1H9a9ll2):.2. 16,andinsidefrontcover sbiepnliglraianlnidcnogeuaxnnttdeenridcniltnoegcrkltwoocisksoienmgdeiir6nec"mtcioborenklsaoscwrvemiwae"nweumdbanrfnireuormm,iatnbwiolsivtve-e Materialexamined:Holotype,CMAAce.No.89.28.1 animals(Figs. 1,2). (Cabrillo MarineAquarium. San Pedro, California), fe- Radialsepia: 16;twobetweenadjacentrhopalia,each male,belldiameterapproximately22cm,founddeadby terminatingoncenteroflappet. staff members ofthe Cabrillo Marine Aquarium, Los Gonads: 4, interradial. surrounding and protruding AngelesBreakwaterLight,nearLongBeach,California, throughsubumbrellarostiaandborneonendsoffinger- 15July 1989. Paratypes: LACM 89-206.1 (Natural His- likeprojections(seemanubriumabove). tory Museum ofLos Angeles County), probable male, Color: Bell dark purple to black, opaque. Oral arms belldiameterapproximately23cm, found liveapproxi- lighter, morepurplethanblack. Darkred-brownmucus mately 100m offshore ofVenice Beach, California, 20 produced when handled (Martin and Kuck, 1991). Pe- August 1989, collected by B.T. Hogue and D. Golles. rimeter with distinctive light brown to tan spotted or LACM 89-205.1, sex undetermined, bell diameter ap- reticulated pattern extending upward toapproximately p1r9o8x9imbayteLlyAC2M5cImc,htfhoyuonlodgyaliSveec,ticoonl,lejcutsetdnoornth25ofAtuhge a'/l<,ltsop'e/4chiemiegnhstsoefebnelolr(cFoilgl.ec1t).edNeoxhviabriitaetdiosnaemnecocuonltorerpeadt;- Marina del Rey jetty, Venice, California. LACM 89- tern. Allcolorfadedtoatranslucentlightbrown in pre- 26.1, sex undetermined, bell diameter approximately servedindividuals. NhW PACIFIC SCVPHOZOAN I I H a Figure3. Chrysanrauclilyiix.newspecies,selectedcnidocytesfrombiopsyofbellperimeter,(a)Section olbellperimetercontainingisorhiza(largearrow)andseverallargeeuryteles,twoolwhichareindicated (medium-sizedarrows),(b)Adjacentsectionofbellperimetercontainingseverallargeeuryteles,oneof whichisindicated(medium-sizedarrow),andsmallereuryteles(smallarrow).Magnificationofbothpho- tographsis400x. Nematocysts:Fourtypes in onesample from edgeof Quentin, Baja California, Mexico). Subsequent sight- bell(Fig.3).CnidomidenticaltothatofC.quinquecirrha ings,includingthosemadebytheHowardHallfilmcrew from Chesapeake Bay (Burnett el a/., 1968; Sutton and offCoronado Norte, Islas de Los Coronados, Mexico, Burnett, 1969).Alargeisorhizaof15 X22^mdiameter weretothenorth,andoften involvedanimalsthatwere (5%,n= 15),asmallerroundisorhizawithafewcoiled moribund (Martin and Kuck, 1991: fig. IB). The latest threads(rare;sizevariationnotmeasured),a 14 X 7^m sightingswere in late August toearly September, 1989, eurytele (14%, = 30), and a clusterofsmaller, 4 x andwerefromLaJolla,California,toVeniceBeachand 7/urn nematocysts(8%, n = 15) resemblingthe larger Santa Monica. California, all to the north of Isla San euryteleswere identified (Fig. 3a, b)(see Discussion for Martin and Coronado Norte, Islas de Los Coronados. commentsonsizevariation). Thus, the species appears to have arrived in southern Lifecycle:Unknown. Californiawatersviaasouthernroute,comingupalong Distribution:IslaSanQuentin, BajaCalifornia, Mex- theoutercoastoftheBajaCaliforniapeninsula. ico, north to Santa Monica Beach, California (Martin Ofthe fivecurrently recognized speciesofChrysaora andRuck, 1991). (Arai, 1997), the new species is most similarto C. plo- Etymology:AnartificialconstructbasedontheGreek camia(see Table I), a speciesabout which verylittle is achlys, meaning mist, darkness, and obscurity (Brown, known.However,C. achlyosc&nbeimmediatelydistin- 1956), referring both to the dark coloration and infre- guished by its size and color pattern, which in C. plo- quentappearanceofthisremarkablespecies. camiaconsistsofdark radiatingstripesupon atranslu- cent bell background. Additionally, a single sentence Discussion translated from Haeckel's(1880) description ofC. plo- camiaservestoestablishamajordifferencebetweenthe Behavioral notes, includingsizeoftheshoals, known twospecies: "Tentaclesasshortasthe umbrelladiame- orsuspectedsymbionts,associates, parasites,and possi- ter."InC.achy/osthetentaclesareperhaps3or4times ble predators, are found in Martin and Kuck (1991). longerthanthebelldiameter(Figs. 1,2). These authors also established that the first sightings Although nematocyst types and sizes are included were made along the Baja California peninsula in the here,westressthatthereisgreatvariabilitywithnemato- summer of 1989 (7 July, 1989, Isla San Martin, San cystmeasurementsin preservedcnidariansamples,and 12 J. W. MARTIN /;V I/ ;_ NEW PACIFIC SOPHOZOAN 13 thatoursampleswerepreservedfor8yearspriortohis- Anderson,R.1992. Highseas'drifter.Mtt lint 101(12):26-24. tological examination. Comparisonbyoneofus(JWB) Aral,M.N.1997. Il-'nnelimialHiologv lScyi>ho:oa Chapmanami of nematocyst measurements of recent and long-pre- BrowHanl.l,R.Lo1n9d56o.n.3Co1n6iip>po.*iiionolSeiennfiel\'ord* SmithsonianInsti- served specimens ofChrysaora quinquecirrha and also tutionPress.Washington,DC.882pp. thehydrozoanPliysaliaindicategreatvariationin mea- Burnett,J.\\'..J.H.Stone, L.M.Pierce, I).G.Cargo,E.C. l.ajnu, suredsizeduetofixation;sizesgiven hereforC. achylos andJ.8.Sutton.1968. Aphysicalandchemicalstudyofseanettle nematocystsprobablydo not reflect sizein life, andare nematocystsandtheirio\m../ Invest Dermatol.51:330-336. inneedofverificationwithlivingspecimens. CampBbaeyllA,quEa.r1i9u9m2.Fou.n1d(aitninolne,inMoinhtcerIIe'<y,/<C/A<>.///1;<6pJpe.llyfish Monterey The only person stung by C. achlyos (J. Martin) re- Cones, H.N.J. 1969. Strohilaiion ofChrvsaoraqiiinquedrrhapol- calledalocalmildburningsensationthatwaslessintense ypsinthelaboratory. I'lrginuiJ ,S'c; 20: 16-18. thanstingshereceivedfromlivespecimensoftheChes- Cranston, B. 1993. Inieniaiional ll'ildli/e 23(4): 2-3 (photograph afpreoamkeC.Baacyhlynoetstlwee,reC.arqeusiunlqtuoecfihrarnhdal.inHgowtheevebra,relsytilnigvs- Croww2d0i2te.hr,ca\pVt.io1n92o6n.ly:Tnhoealcicfeoomfpatnheyimnogont-ejxet)l.ly.Hal.Oeogr.50: 187- ingordead medusaeaftercollecting,andwereprobably Haeckel, E. 1880. Da* System ilcr Medusen Ersler Theil eiuter caused by free nematocysts floating in the collecting MonographiederMednsen G. Fischer, vormalsF. Mauke.Jena. bucket,asnotentaclesremainedonanyofthecollected 672pp. asntipefelicyciamvnetenncslo.imniCochuarlsycsgauesoneruwasaiossfgtjehenallteyrfoaiflslha.yvPrieecrgthaimradpwesdhtoahsedaemvomesoltdoepsireg-d- HIallallS1,s1a1t1neI*Ia.dtL,uu1ni9B/.s90Ol\.he\in.sn1piT9omh6,ne5i.CK*Ae.lMPpa4hr0lyi-'lpnopuree.msiA.CnonHeuolawelna*tredr<>a/tHtaah.lelPpa\.\nnd2rl9dB7.l-a5vko3el5.PuIibnlIPinosvihesironlnge.-- massiveurticaria,lightheadedness,andnauseaformany hrates. U.S.Government PrintingOffice. Washington. DC. 494: minuteswhenstungbyahostofChesapeakeBaynettles plate43. (('. quinquecirrha)fallingon him froma broken fishing llalCsotelaodr,iBi.la\*V.Da1r99w2i.nPrDeasnsg,ePrroinnc*etAotn/n,aNliJe.A2n6i4mpapl.*ulthe World..1 nethoistedoverhishead(Hartman elul., 1980, patient llartman,K.R.,G.J.Calton,and.1.W.Burnett.1980. Useofradio- A). Ifsuchan injurycouldoccurfromcontactwith me- allergosorbenttestforthestudyofcoelenteratetoxin-specificim- dusaeofsmallernettles(albeit manyofthem), it might munoglobinE.Inl Areli Itlergy.!/)/>/.lininiinol.61:389-393. bcaelrseyamsopntaobmlsetionaanhtiucmiapantetheaqtuaclolnytaocrtemdortehesteevnetraeclcelsinoi-f kKraaknniianpeu,im/aPi.,eaLGY.o.e1t91l69e16.7..BulSlyDneMovpaesrliospllmoifenlntthSelaotmfeadlu*sasciaiyiepihi*oo/fznot1ah3ne(.Iwo)D:ra2le9dl-.y3l3/o.mMelarra theconsiderably largerC. achlyos. Caution istherefore BiolAssoc Lf*40:1-464. advised in approaching orcollectingthisspecies when- Kuck, II.G., and J.W. Martin. 1990. Mysterious medusa. Terra everitnextappears. (NaturalHistoryMuseumofLosAngelesCountri29(1):48-49. Larson, R.J. 1990. Scyphomedusae and Cubomedusae from the easternPacific,hull Mar Sci -47:546-556. Acknowledgments Larson,R.J.,andA.C.Arneson.1990. Twomedusaenewtothecoast Wethank Paul Cornelius forprovidingthe morpho- oufsaCaalnifdorPnhiyal:loCralrnyzhadpeiainmcairasiialpvioalnisLe(nLdmennefaeeluds..11878548,).aarehiuzboosmteodm-e logical frameworkwefollowed forthedescription; Ron scyphomedusa.BullSmith Calif,.hadSci.89(3):130-136. LHaalrlsofnorfaolrlhoewlipnginusdteoduucsienhgisthoericgoirnarlecpthgoetnougsr;apHhos;waMrid- MartsBipaenjc,aiJeC.salWoi.ff,oSarcnnyidpahIoiIn.zoG1a.98Kf9uo.cuknS.dmi1nt9eh9a0r.CsahloirIfn.evAfeerrtaoedmb.rsaSoteuett.a.hsesArobncsitCaratalecistfsoofronalinaAentw-o chelle HallofHoward Hall Productions, Karen Koppel nualMeeting,May11-12.IWO.Abstract64. of Scott Foresman-Addison Wesley Publishers, and Martin,J.W.,andH.G.Kuck. 1991. Faunalassociatesofanunde- HowardYouthofZooGoermagazineforhelpwithliter- scribedspeciesofChrysaora(Cnidaria,Scyphozoa)intheSouthern aatnudre;thHeelsetanfefRaunbdinvsotleuinntefeorrsthoefhtihsetolCoagbyriplrleoparMaatriionne; NatuCwraaelti.efro1rs9pn9ei0ca.ieBsigSihneta,t*hwmeiiat*rheain.noBttuehlseloS.neSmauintNuhastuCiaaollln/ao,lcc.GluerearodegnrScaeepish9io0fc(o3V)ti:hde8re9ow-1aP0rr1em-. Aquariumfortheirencouragement.Thisstudywassup- sentation,incollaborationwithThirteen-WNETandHowardHall ported in part by the Friends ofthe Cabrillo Marine Productions. Aquarium;the DepartmentofBiology,CaliforniaState Pages, F.,J.-M.Gili,andJ. Bouillon. 1992. Medusae(Hydrozoa. sPUtnriiotvugetrreasmiUt;nyda,enNrdogrtrthahedruNiaadttgieeo;nRtaehlseeSHacoricewnhacreidnFoHBuuinogldhoaegtisicoaMlnedvSiiccaiaeRlnEcIenUs- RusstSleiaclcg)yli.,pcShFcSoi.cz.ySoM.pa.ha1or9Cz7uo05b.a6ozw(oisTtauhh)pepaolMfSeetumdhpeeupnslBtaeenIme)ge:uunl1et-lt6htai4e>.CBturhrierteFiinsrhtstI(sslIoe'su.othlIen'amoseltioeirnmneHAIyItdlraPone--- (Research ExperiencesforUndergraduates)supplement medusae CambridgeUniversityPress,London.284pp. toDEB9320397toJ.W. Martin. StraFuossr,esEm.,ana-ndAdMd.isLoinsoWwessklie.y1P9u9b7l..,MBeionlloogyP.arTkh,eCAll.'ch10o1j6Lipfpe..Scott Sutton,J.S.,andJ.\V. Burnett. 1969. A lightandelectron micro- LiteratureCited scopicstudyofnematoeytesofChrysaorat/uini/iicc/rrha.J. Vltras- iniel.Re*.28:214-234. Agassiz, A.,and A.G. Mayer. 1898. On Daelylomelra. Bull. A/i/.v. Whiteman,L.1996. Lights,camera,wildlife!ZooGocr(Friendsofthe Camp./.mil.HarvardUniv.32:1-11,+ 13plates. NationalZoo),Sept-Oct.1996:15-19.