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Notice: ©1990 Rosenstiel School of Marine and Atmospheric Science, University of Miami. This manuscript is available at http://www.rsmas.miami.edu/bms and may be cited as: Larson, R. J. (1990). Scyphomedusae and cubomedusae from the eastern Pacific. Bulletin of Marine Science, 47(2), 546-556. BULLETIN BULLETIN OFOFMARINEMARINESCIENCE.SCIENCE.47(2):47(2):546-556,546-556,19901990

from 0 SCYPHOMEDUSAE SCYPHOMEDUSAE ANDANDCUBOMEDUSAECUBOMEDUSAEFROMFROMTHETHE large 0 EASTERNEASTERNPACIFICPACIAC expedi 1968a; R,R,J.J.LarsonLarson medus The ABSTRACTABSTRACT report The purpose study The purpose ofofthisthispaperpaperisistotoreviewreviewthetheliteratureliteratureononthethescyphomedusaescyphomedusaeandandcubome­cubome­ dusae from the eastern and Fl dusae from the eastern Pacific.Pacific.Forty-threeForty-threespeciesspeciesofscyphomedusaeofscyphomedusaeareareknownknownfromfromAlaskaAlaska to Chile. Cubomedusae easten to Chile. Cubomedusae arearerepresentedrepresentedonlyonlybybyCarybdeaCarybdeamarsupialismarsupia/isfromfromthethecoastcoastofof southern California. Ten SOIl southern California. Ten speciesspeciesofofstauromedusaestauromedusae areareknownknownfromfromthisthisarea,area,mostmostoccuroccurinin shallow, subarctic and are ec shallow, subarctic and temperatetemperatewaterswatersofofthetheNorthNorthPacific;Pacific;HaliclystusHaliclystusoctoradiatusoctoradiatusisisthethe most common. Fourteen most common. Fourteen speciesspeciesofofcoronatecoronatemedusaemedusaeandandoneonepolyppolyp(Stephanoscyphus(Stephanoscyphussp.)sp.) some have been reported, most have been reported, most arearemesopelagicmesopelagicandandmanymanyarearewidelywidelydistributed.distributed.ThirteenThirteenspeciesspecies Work ofsemaeostomes ofsemaeostomes areareknownknownfromfromthetheeasterneasternPacific,Pacific,mostmostareareneritic.neritic.However,However,twotwomeso­meso­ of scy bathypelagic , bathypelagic species, DeepstariaDeepstaria enigmaticaenigmaticaandandPoraliaPoralia ru/escens.rufescens, havehavebeenbeencollectedcollectedatat ducti, several locations, several locations, includingincludingsouthernsouthernCalifornia.California.ChrysaorafuscescensChrysaorajuscescensisisthethemostmostabundantabundant HamI semaeostome species semaeostome species inin thethe northeasternnortheastern PacificPacificandandcancan occuroccuratathighhigh biomassbiomasslevelslevelsinin inshore 1988) inshore waters.waters. RhizostomesRhizostomes areare representedrepresented byby onlyonly fivefive species,species, allallsubtropical-tropical.subtropical-tropical. The scyphomedusan repon The scyphomedusan faunafauna alongalong thethe entireentire easterneasternPacificPacificcoastcoastisisbadlybadlyininneedneedofofstudy.study. The ofa number The taxonomy ofa number ofofspeciesspeciesrequiresrequiresclarification.clarification.TheThegeographicgeographicdistributionsdistributionsofof mostmost speciesspecies areare unlrnown.unknown. LittleLittleisis knownknown aboutabouttheirtheirbiologybiologyandandecology.ecology. Me Scyphomedusae, porte, Scyphomedusae, because of theirtheir size,size, upup toto I1 m m andand more,more, andand oftenoften con-con­_ I spicuous Th' spicuous coloration, are familiar toto nearly everyoneeveryone who frequentsfrequents thethe seashore.seashore. They are elsew They are also important predators that can have a largelarge biomass inininshoreinshorewaterswaters t (Pearcy, 1976; l the b (Pearcy, 1976; Moller, 1984; Shenker, 1984; Fancett and Jenkins, 1988). Yet, along the trati

546546 lARSON:lARSON: EASTERNEASTERN PAClFlCPACIFlCSCYPHOMEDUSAESCYPHOMEDUSAE ANDANDCUBOMEDUSAECUBOMEDUSAE 547547 from off southern California and Mexico. Segura-Puertas (1984)(1984) reportedreported onon thethe largelarge collection of medusae (mostly(mostly hydromedusae)hydromedusae) takentaken byby thethe EASTROPACEASTROPAC expeditions inin thethe tropicaltropical easterneastern Pacific.Pacific. Further south,south, KrampKramp (1952;(1952; 1966;1966; 1968a; 1968b), Fagetti (1973),(1973), and Larson (1986)(1986) describeddescribed thethe pelagicpelagic scypho­scypho­ medusae from thethe coast ofofChile. The benthic areare lessless well known. TheThe mostmostextensiveextensivetaxonomictaxonomic report on this group was done by Gwilliam (1956),(1956), but unfortunatelyunfortunately mostmostofofthisthis study was never published. More recently,recently, papers by LarsonLarson (1988)(1988) andand LarsonLarson and Fautin (1989)(1989) described threethree new speciesspecies ofofstauromedusaestauromedusae fromfrom thethe north­north­ eastern Pacific, includingincluding a new genus andand family.family. Some ofthethe more recent studies on thethe scyphomedusaescyphomedusaefromfrom thetheeasterneasternPacificPacific are ecological. Alvarifio (1967; 1976) has described thethe depth distributions ofof some oceanic scyphomedusae fromfrom thethe coastscoasts of CaliforniaCalifornia andand BajaBaja California.California. Work by Pearcy (1972; 1976) and Shenker (1984)(1984) havehave shownshown thatthat thethe biomassbiomass of scyphomedusae, mostly fuscescens,fuscescens, isis extremely high offoffthethe pro­ ductive upwelling coast of Oregon. Studies inin thethe Bering Sea (Harrison,(Harrison, 1984;1984; Hamner and Schneider, 1986) and inin British Columbia (Strand(Strand andand Hamner,Hamner, 1988) also emphasize thethe ecological importanceimportance of thisthis group.group. TheseThese andand otherother reports suggest thatthat scyphomedusae are an importantimportant part ofpelagic foodfood chains.chains.

DISCUSSION More than 40 species of scyphomedusae and one cubomedusa have beenbeen re­re­ ported from thethe eastern Pacific (Table(Table 1).I). The taxonomic status of some scyphomedusaescyphomedusae fromfrom thethe easterneastern PacificPacific andand elsewhere is confused. Primarily thisthis isis because many species were describeddescribed onon the basis of only a few specimens, sometimes inin mutilated condition,condition, andand illus­illus­ trations were inadequate.inadequate. This has ledled toto confusion,confusion, especially regardingregarding speciesspecies in the genus Chrysaora. Type specimens, even ififtheythey exist, areare very mutilatedmutilated oror damaged and coloration has fadedfaded so thatthat theythey are oflittlelittle help. Although therethere is some scyphomedusan material from the Pacific coast inin museums, moremore spec­spec­ imens are badly needed. Ifpossible theythey should be accompanied bybycolorcolor photos,photos, or notes accurately describing pigmentation. Cubomedusae. -Carybdea rasloni was the only known cubomedusa fromfrom thethe northeastern Pacific. Stiasny (1922) first reportedreported itit fromfrom La Jolla,Jolla, California.California. However, I have examined cubomedusae material from southern California and concluded that Carybdea marsupialis, not C. rasloni,rastoni, isis thethe correct determination (Larson and Arneson, inin press). This isis thethe firstfirst timetime thatthat thisthis speciesspecies hashas beenbeen described from thethe Pacific Ocean. Gladfelter (1973)(1973) described itsits swimmingswimming me­ chanics and noted thatthat it was present occasionally inin thethe kelp beds off Santa Barbara. Pearse and Pearse (1978) published a paper on thethe elaborate lensedlensed eyeseyes of C. marsupiamarsupialis,lis, and neurophysiological studies on control of swimmingswimming were done by Satterlie (1979) and Satterlie and Spencer (1979).(1979). It is puzzling why only one species ofcubomedusa has been reportedreported fromfrom thethe eastern Pacific whereas four or more species occur inin the tropical Atlantic and inin the western Pacific. Perhaps other species, e.g., Carybdea a/alaalala Reynaud and Chiropsa/musChiropsalmus quadrumanus Haeckel, two widely distributed subtropical-tropical species, will be found with further collecting. Slauromedusae. -Stauromedusae have not receivedreceived much attention because theythey are cryptic and occur only in a few habitats. Not until Gwilliam (1956;(1956; 1960) studied their taxonomy and physiology was much known about thisthis group fromfrom Table Suborder Order Family Suborder Order Class Class Family Family Family Family Family Table I. Annotated check list of and Cubozoa from the eastern Pacific Family Atorella Atolla Atorella Atalla Atalla Nausithoe Nausithoeatlantica Nausithoe "Stenoscyphopsis Manania Manania Atalla Haliclystus K.vopoda Craterolophus Nausithoe Manania /{aliclystus C'aryhdea /{aliclystus Vo ~ 1. Taxon 00 Scyphozoae Distribution Cubozoa Stauromedusa

Coronatae References·' Annotated Nausithoidae Depastridae Atollidae Carybdeidae Linuchidae Kyopodiidae chuni gigantea vanhoeffeni w.vvillei

Class Cubozoa Cleistocarpida Eleutherocarpida arcwri vanhoeffeni unguiculata lamberti gwilliami distincta handi marsupialis albatrossi punctata Family Carybdeidaerubra stejnegeri "californiensis" salpinx octoradiatus Caryhdea marsupialis (Linnaeus, Vanhoffen, 1758) S. California 15,35,47 sp. Haeckel, Maas, Bigelow, Lan;on check Class ScyphozoaeVanhoffen, vermiformis" Lan;on, Russell, (Kishinouye, Clark, Lan;on Kolliker,

Order StauromedusaBroch, Bigelow, (Maas, (Schwartz, Kishinouye,

Suborder Eleutherocarpida (Linnaeu5, 1897 Taxon list and 1880 (Lamarck, Family Lucernariidae 1902 1928 1863 of 1988 and lIaliclvstus octoradiatus (Lamarck,1957 1816) 1914 Alaska to C. Calif.

Fautin, 5, ?14, 722, 24

1897) ~ 1902 Scyphozoa 1853 lIalidvstus salpinx1909 Clark, 1863 Vancouver I., S. Alaska 24,37,47 Fautin, 1788) Haliclystus stejnegeri Kishinouye, 1899 1899) N. Alaska 9 § 1758) Haliclystus "californiensis" S. California1899 z 1816) 24 "Stenoscyphopsis verrniforrnis" 1989 o C. California 24 "1 1989

Family Kyopodiidae and ;l> Kyopoda lamberti Larson, 1988 B. Columbia, Calif. 45 ";x3

Cubozoa Z Suborder Cleistocarpida tTl

Family Depastridae 15 sp. Craterolophus Pribilof Islands 47 ~ Manania distincta (Kishinouye, 1899) Alaska to Oregon 46 from R Manania gwilliami Larson and Fautin, 1989 B.C. to Baja Calif. 24,46 cosmopolitan S. S. S. Colombia Chile Oregon, C. Alaska Vancouver S. Panama, Panama, Pribilof Mexico Panama, B. B.C. Vancouver Alaska N. MananiaMexico. handi Larson and Fautin, 1989 Vancouver I. Region 24,46 ~ the Calif., Chile California California Columbia, California ... Order Coronatae Alaska ..... to

eastern z to Family Atollidae to Baja to

Islands 9 B. Colombia S. S. Chile Panama C. Atolla chuni VanhOffen, 1902 Oregon S. Chile !" Chile 44 Columbia I. I., California Chile '0

Atolla gigantea Maas, 1897 Distribution Calif. Panama, S. ChileCalif. 8,44 Region Calif. Pacific S. 8 Atolla vanhoejfeni Russell, 1957 Oregon, B. Columbia 33,44 Atolla w.vvillei Haeckel, 1880 cosmopolitan Alaska 8, II, 12, 16, 17,20,27,29,30,31,33,34,36,44 Family Nausithoidae Atorella arcturi Bigelow, 1928 Colombia 16 Atorella vanhoejfeni Bigelow, 1909 Panama, S. California 11,32,36 Nausithoe albatrossi (Maas, 1897) Panama, Chile 8,22,41 Nausithoe atlantica Broch, 1914 Chile 34 Nausithoe punctata Kolliker, 1853 Mexico to Chile 11,16,26,32,36,41,47 Nausithoe rubra Yanhoffen, 1902 S. Calif., Panama 11,16,27 Family Linuchidae 47 24,46 44 46 45 33,44 34 24,37,47 24,46 24 24 11,16,26,32,36,41,47 11,16,27 16,47 16 11,32,36 15,35,47 8, 9 8,22,41 8, Linuche unguiculata (Schwartz, 1788) Mexico, Colombia5, 16,47 1,?22, ?14, 44 II, 2 16, 12, 24 17,20,27,29,30,31,33,34,36,44

~ ... ".,, .. _».~ ~,...... - ...... _. _•. __ .. ' ...... _.e.'.'__ .... __ .• .._"'

Table I. Continued Referencese' Distribution ".'_.,..~."" "" ...----... ".O-.'_'~_"'''''_''' __ '_'''' __ ~' '''''''''~_'~'''''''''''':''~''-''~'''''''''''''''''''

Table I. Continued I • (1981),39 20 29 K"hinouye Fautin Taxon Distribution Table References-' - - Order Order Family Family Family Family Family Family Family Family Family R.f.renct5: Ilig.low Kramp Aurelia Aurelia Phacellophora Pela!(ia Chrysaora Chrysaora Ca/ostylus Cyanea Pela!?ia Phyllorhiza Poralia Periphy/lopsis Periphylla Catostylus Chrysaora Chrysaorajuscescens Deeps/aria Paraphy/lina Family Periphyllinidae Lychnorhiza (1989),

Paraphy/lina ransoni Russell, 1956 Oregon 33 1. - (I Rhi7.0stomeae (1968a), Smith Continued Cyaneidae Catostylidae Family Periphyllidae(1940),21 899h), Lychnorhizidae Periphyllinidae Periphyllidae Mastigiidae 47

- 1 30,33,3~ aurita limbata colorata n~(escens capi/lata Periphy/la periphy/la (Peron and Lesueur, 1809) cosmopolitannoctiluca 8,11,12,16,20,27, 29, 36,44 - (1982),

l..elson 11 Larson, Periphy/lopsis braueri Vanholfen, 1902 Peru periphy/la sp. ornate/lus plocamia /a!?i melanaster II eni!(ma/ica punc/ata 30 sp. - ransoni meleagris

- ~ braueri (Linnaeus,

Order Semaeostomeae- cam/schatica Big.low

(Haeckel, :>:l Stia.ny (Brandt, Kramp (1810), 40 Russell, unpubl.,

Family Pelagiidae (Linnaeus. ~ Vanholfen, (Forskrtl, Chrysaorajuscescens- Brandt, 1835 Bering Sea to Mexico 2,3,4, 7, 10,38,42,47 :z (Lesson, Purcy (Peron (1909). (Vanholfen, and von Brandt, Russell. (1968h), Vanholfen, Chrysaora melanaster - 2 Brandt, 1835 ~ Bering Sea Brandt, 2,3,12 Taxon Russell, 48 L.

ChrysaoraMud.n plocamia (Lesson, 1830) Peru, N. Chile I, 18,22 1835)

1964 ~ 1869) Brandt 1758) Lendenfeld. - and Agassiz, :>:l 12 Chrysaora sp. S. California1775) 47 F. Brandt,

and Z 1758) 31 1902 - Stuiver Sammen, 1835 Pelagia colorata Russell, 1964 S. California [830) 7,25, 35,47 1956

1835 ~ (1943), Big.low - (1833). Pelagia noctifuca (ForskAI, 1775) 1967 Mexico to Chile II, 16,20,30,32,34,41,43,47 (') Lesueur, Alvariilo 1902 1888) 5i (1983),41 Family Cyaneidae 1862 (') 1835 22 - 3 (1913), Cyaneapen. capi/lata (Linnaeus, 1758) Bering Sea to Central California 4,12,14,21,23,38,42,47,48 1884 -

(1969). Q Brandt

Family UlmaridaeKI'1Imp comm., ~ 1809)

Aurelia aurita13 (Linnaeus, 1758) cosmopolitan 2,4,23,33,38,42,47 - o ~ - s"gura·Pu.rta.

Aurelia limbata32 (Brandt, 1835) Alaska, ?8. Columbia 2,3, 12, 14,21 (1838).

(1952),23 ttl Bigelow \

Deeps/aria- I'nigmatica Russell, 1967 California, Oregon 28,40 990; g

Alvariilo 34,38,4~44,47 Phace/lophora camtschatica Brandt, 1835 entire coast 4, 12,22,26, ~ - 4

(S ttl

Poralia rufescens(1914). Vanholfen, 1902 S. California, Peru 11,39

• )­ ., -

Aga Z MacGiniti. Shih

Order Rhizostomeae(1984),42 (\972), entire cosmopolitan S. S. S. S. S. California, cosmopolitan o Alaska, Oregon Mexico Ecuador Bering Panama Bering Mexico Bering Peru Peru, .. 14 i, Calif. California California, California Family Catostylidae California et 2 - (1862).

Ca/ostylusal. ornate/lus (Vanholfen, 1888) Ecuador 6 ~ Bigelow 33 N.

coast ~ (1971) Sea Sea Panama Sea 15,47 (1953), Catostylus- tagi (Haeckel, 1869) - ?8.

to ttl to Chile Shenker

P.arcy o 3

Family Lychnorhizidae Chile c: Oregon to to Ecuador - (1920),

Columbia ell for Lychnorhiza sp. Mexico 47 Distribution )­ Clark 24 Peru Central Mexico ttl additional (1972), - Family Mastigiidae(1984), Gwilliam 13 Phy({orhiza punclata(1863).6 von Lendenfeld, 1884 S. California 47 - 34 Stiasny California

Family Rhizostomatidae43 record. - S. Calif. to Ecuador 6,13,15,19,20,30,47 Slomolophus- meleagris L. Agassiz, 1862 (1956), Fag.tti - Alvariilo (1922), •I R.f.rencts: I- lessonVanhoffen (1810), 2 - Bl'1Indt (1833). 3 - Bl'1Indt (1838). 4 - Aga.. iz (1862).3 - Clark (1863). 6 - Vanhall'.n (1888). 7 - Fewkes (1889),8 -M... (1897), 9 - Ki.hinouye (1899a), 10­

Kishinouyo (Ifrom 899b). II - Big.low (1909). 12 - Big.low (1913). 13 - Bigelow (1914). 14 - Bigelow (1920). 13 - Stia.ny (1922). 16 - Bigelow (1928). 17 - Boone (1933), 18 - Stia.ny (1937), 19 - Boone (1938), 23 20 - Ilig.low (1940),21(1973), - Stia.ny and M.. den (1943), 22 - KI'1Imp(1932),23 - MacGinitie (1955). 24 - Gwilliam (1956), 23 - Ru... 11(1964),26 - KI'1Imp (1966).27 - Alvariilo (1967).28 - Ru... 11(1967). 29 - Kl'1Imp (1968a). 30 - Kl'1Imp (1968h). 31 - Alvariilo (1969). 32 - Alvariilo (1972). 33 - Pearcy (1972). 34 - Fag.tti (1973). 33 - Gladf.lter (1973), 36 - A1variilo (1976), 37 - Otto (1976), 38 - Mm. ~ - 16 Brili.h (1981).39 - SmIthand (1982), 40 - Purcy and Stuiv« (1983),41 - Segul'1l·Pu.rta. (1984),42 - Shenker (1984), 43 - Alvariilo and Ambr6. (1986), 44 - Lanon (1986), 43 - Larson (1988), 46 - Larson and \0 Ru.sell - Faulin (1989), 47 - Larson,(1888), unpubl.. 48 - F. Sommen. pen. comm.• 1990; (Se.: Shih etal. (1971) foraddilional records from Briti.h Columbia). 33 Ambr6. Bigelow Columbia). - 47 28,40 47 47 33 11,16,20,30,32,34,41,43.47 15,47 11,39 11 (1964),26 Gladf.lter 7,25, 4, 6 4,12,14,21,23,38,42,47,48 8,11,12,16,20,27,29,30,33,34,36,44 6,13,15,19,20.30,47 2,3,4,7,10,38,42,47 2,4,23,33,38,42,47 2.3,12 2, 1. - 7 (1928), (1986), 3, 12,22,26, 18,22 Fewke. 2 14, 12, 35,47 - (1973), 17 44 KI'1Imp (1889),8 - - Boon. 21 Larson 36 34,38,4~44,47 (1966),27 - (1933), M - A1variilo (1986), References-' ... 18 - (1897), 43 A1variilo (1976), - - Slia.ny Larson - 9 37 (1967),28 (1937), Ki.hinouy. - (1988), OUo 19 (1976), 46 - - Ru.sell (1899a), - Boone Larson 38 - (1967), (1938), Mill. 10­ and \0 VI ~ 550 BULLETIN OF MARINE SCIENCE.SCIENCE. VOL. 47.47.NO.2.NO.2. 19901990 the eastern Pacific. Unfortunately, Gwilliam's taxonomictaxonomic studiesstudies werewere neverneverpub­pub­ lished. Recently, Larson (1988) described a new worm-like stauromedusa,stauromedusa, Kyo­Kyo­ poda lamberti, from British Columbia and California. LarsonLarson andand FautinFautin (1989)(1989) added two additional new species fromfrom thethe northeast PacificPacificbybydescribingdescribingMana­Mana­ nia gwilliami and M. handi. They also added aa newnew distributionaldistributional record,record, M.M. distincta, which was not previously known fromfrom thethe northeast Pacific.Pacific. TwoTwo ofof Gwilliam's new taxa (Haliclystus "californica" and "Stenoscyphopsis vermifor­vermifor­ mis") are yet to be formally described; theythey were reportedreportedoffoffsouthernsouthernandandcentralcentral California, respectively. Studies on the biology ofstauromedusae fromfrom thetheeasterneasternPacificPacificarearefew.few. HymanHyman (1940) described some aspects ofthethe general behaviorbehaviorofofHaliclystusHaliclystus octoradiatusoctoradiatus (as H. "sanjuanensis"). Gwilliam (1960) did an in-depthin-depth studystudy onon thethe neuro­neuro­ muscular physiology of H. octoradiatus and Otto (1976)(1976) examinedexamined thethe earlyearly de­de­ velopment and locomotion of the planula of Haliclystus salpinx.salpinx. Haliclystus octoradiatus (=H.(=H. auricula Rathke) (Larson(Larson andand Fautin,Fautin, 1989)1989) isis the most common stauromedusa along thethe westwest coastcoast ofofNorthNorth AmericaAmerica (Fig.(Fig. I).1). Around Vancouver IslandIsland and Puget SoundSound itit isis foundfound inin aa varietyvariety ofofintertidalintertidal and subtidal habitats and reaches relativelyrelatively high densitiesdensities (unpublished(unpublished obser­obser­ vations). Kozloff (1983) illustratesillustrates thisthis speciesspecies (fig.(fig. 345,345, asas H.H. stejnegeri).stejnegeri). ThisThis isis the only known stauromedusa fromfrom thethe southeastern PacificPacific (Kramp,(Kramp, 1952)1952) butbut further collecting will probably yield other species.species. L.L. BarrBarr (pers.(pers. comm.,comm., 1975)1975) sent me photos of a Craterolophus sp. fromfrom thethe PribilofPribilofIslands.Islands. ThisThis genusgenus isis unreported in the Pacific. Coronatae. - This is a small though widely distributed group.group. MostMostcoronatescoronates areare oceanic or they live along the outer part ofthethecontinentalcontinental shelf;shelf; howeverhoweverLinucheLinuche unguiculata and Nausithoe punctata, both tropicaltropical species,species, areare thethe onlyonly coronatecoronate medusae that are regularlyregularly foundfound inin shallowshallow water.water. TheThe mesopelagicmesopelagic species,species, especially Atolla spp. and Periphylla periphylla, can bebe expectedexpected toto bebe collectedcollected in deep wateranywhere along thethe entire coast. Additionally, oneonewidelywidelydistributeddistributed I species ofAtolla,Atalla, A. parva Russell, probably occursoccurs onon thisthis coastcoast butbut hashas notnot yetyet been reported there. The above-mentioned medusae areare describeddescribed andandillustratedillustrated by Russell (1970) and Larson (1986).(1986). AA coronatecoronatepolyppolypStephanoscyphusStephanoscyphushashasbeenbeen reported off Panama, but isis probably widelywidely distributeddistributed elsewhereelsewhere inin thethe easterneastern Pacific since it is cosmopolitan (Kramp,(Kramp, 1959). Semaeostomeae. - This ordercontains thethemost commoncommonscyphomedusae.scyphomedusae.SeveralSeveral genera reach 25-50 cm and largerlarger (e.g.,(e.g., Aurelia,Aurelia, Chrysaora,Chrysaora, Cyanea,Cyanea, ,Deepstaria, Phacellophora), and the majority are conspicuouslyconspicuouslypigmentedpigmented(Aurelia(Aureliaauritaauritaandand Deepstaria enigmatica being exceptions). MostMost areare neritic;neritic; howeverhowever twotwo speciesspecies (Deepstaria enigmatica and Poralia ruJescens) are meso-bathypelagic. These latter Fi (Deepstaria enigmatica and Poralia ruJescens) are meso-bathypelagic. These latter di medusae have been photographed and observed inin situsitu fromfrom submersiblessubmersibles offoff e~ southern California and are described and illustratedillustratedbybyLarsonLarson(1986).(1986). DeepstariaDeepstaria C enigmatica has a very thin transparenttransparent umbrellaumbrella whichwhich hashas aa networknetwork ofofthinthin di Ii, canals (white in formalin)formalin) runningrunning throughthrough it.it. ItItwaswas firstfirst seenseen byby BarhamBarham (1969)(1969) and was described by Russell (1967).(1967). Poralia ruJescensruJescens hashas aa darkdark brownish-redbrownish-red umbrella with these same pigments occurring inin thetheumbrellarumbrellarmesoglea.mesoglea. ItItisisveryvery fragile and readily fragments during collection withwith nets,nets, whichwhich explainsexplains whywhy itit has not been reported previously. ItIt was photographedphotographed althoughalthough notnotidentifiedidentified byby Hartman and Emery (1956,(1956, pI. 2, fig.fig. d).d). SmithSmith (1982)(1982) foundfound thatthatitit waswasabundantabundant in the Santa Catalina Basin and diddidsomesome ininsitusitustudiesstudiesononitsitsphysiology.physiology. AnotherAnother cosmopolitan, but rare deep-sea semaeostome, StygiomedusaStygiomedusa giganteagigantea (Brown),(Brown), likely will be found along thethe Pacific coast. ItIt hashas anan umbrellaumbrella whichwhich reachesreaches II LARSON: EASTERN PACIFIC SCYPHOMEDUSAE AND CUBOMEDUSAE 551

Figure 1. Scyphomedusae from the eastern Pacific: A) Haliclystus octoradiatus side view, 2.5 em diarn., Oregon; B) Haliclystus salpinx side view, 1.5 em high, Vancouver Island; C) Pelagia colorata exumbrella, 35 em diarn., S: California; D) Chrysaorafuscescens side view,S em diam., Oregon; E) Chrysaora melanaster side view, from Brandt (1838); F) Phacellophora camtchatica exumbrella, 6 em diam., Oregon; G) Chrysaora plocamia portion of exumbrella, from Stiasny (1937) and H) Aurelia limbata.side view and subumbrella from Brandt (1838).

m in diameter and the oral arms may be 10m in length. The gastrovascular canals in the umbrella of S. gigantea form a characteristic anastomosing pattern (Larson, 1986). The well known genus Aurelia is represented along this coast by two species. Aurelia aurita is often abundant south of Alaska, in inshore waters. Hamner and Jenssen (1974) studied its growth in Tomales Bay, California. The ultrastructure of the was described by Chi a et al. (1984). Strand and Hamner (1988) 552 BULLETIN OF MARINE SCIENCE, VOL. 47,47, NO.2, 19901990 described the predation of Phacellophora camtschatica on A, aurita inin waterswaters ofof Califc British Columbia. comn In Alaskan waters occurs a second Aurelia species, A. limbata.limbata. AlthoughAlthoughStiasnyStiasny Th and Maaden (1943) reported it from off Vancouver Island,Island, thisthis requiresrequires corrob­corrob­ Pela~ oration. This cold-water medusa has a distinctive chocolate-brown bellbell margin.margin. (Tab1 A color photograph ofthis species, takentaken inin thethe Aleutian Islands,Islands, appearsappears onon thethe regul cover ofAudubon magazine (January 1974). Uchida and Nagao (1963)(1963) describeddescribed nortl its early development and metamorphosis. byR The taxonomy, and consequently thethe distribution ofofnearlynearly allall speciesspeciesofofChry­Chry­ Pela, saora, are confused and this includes thethe (four) speciesspecies thatthat occuroccuralongalongthethe westwest shov coast of North America. The major problem isis thethe lacklack ofofdefinitivedefinitive taxonomictaxonomic mag: characters. However, the three species presently knownknown fromfrom thisthiscoastcoast areare easilyeasily vesti separated by their coloration when alive or freshlyfreshly preserved.preserved. ChrysaoraChrysaora melanas­me/anas­ poly ter from the Bering Sea has a brownish pigmented patternpattern onon aa pale,pale, whitishwhitish scyp exumbrella. Sixteen streaks radiateradiate from aa broad centralcentral ringring andand betweenbetween thethe I U streaks, near the margin, are 32 crescents. II have seen photosphotosofofthisthismedusamedusafromfrom a wI Alaska (L. Barr, pers. comm., 1975) which closelyclosely agreeagree withwith thethe illustrationillustration inin tent Brandt (1838). Rarely, this medusa can sometimes driftdrift southsouth atat leastleast asas farfar asas iI jUVt Oregon (unpub. obs.), and it is possible thatthatitit isisoccasionallyoccasionally carriedcarriedevenevenfurtherfurther (18l south by the California Current since itit was reportedreported offoffCaliforniaCalifornia byby AgassizAgassiz bee (1862). I have seen no specimens fromfrom there.there.ItItisis possiblepossiblethatthatsomesomeC.C. melanasterme/anaster and specimens have been confused with another species,species, namelynamely C.C. fuscescens.fuscescens. ThisThis 1 medusa is abundant offCalifornia and Oregon during thethesummersummermonthsmonths(Shenk­(Shenk­ am er, 1984; unpub!.unpubl. obs.). The exexumbrellaumbrella of C. fuscescensfuscescens isis aa darkdark amberamber colorcolor ( which is darkest near thethe margin. A faintfaint radiatingradiatingstar-likestar-likepatternpatternmaymaybebevisiblevisible stu on the exumbrella but itit isis much lessless distinct thanthan thethe darkdark patternpattern onon whitewhite me background of C. melanaster.me/anaster. Also, inin C. fuscescensfuscescens thethe oraloral armsarms spiralspiral screw­screw­ w~ like, whereas in C. melanasterme/anaster theythey are lessless twisted.twisted. un: Chrysaora fuscescens was originally described byby BrandtBrandt (1835).(1835). ThreeThree yearsyears pel later, Brandt (1838), for some unknown reason,reason, referredreferred toto thisthis speciesspecies asas C.C. no helvola.he/vola. Specimens were first collected by Mertens inin thethe AleutianAleutian IslandsIslands andand atat thl Sitka, Alaska. Color illustrationsillustrations by Mertens appearappear inin BrandtBrandt (1838).(1838). ShortlyShortly Gl afterwards, A. Agassiz (1862) reportedreported thisthis speciesspecies (as(as MelanasterMe/anaster mertensiimertensii L.L. Agassiz, and PolybostrichaPo/ybostrichahelvola he/vola Brandt) fromfrom California.California. FewkesFewkes (1889)(1889) foundfound Ie. this species at Santa Cruz, California (identified(identified asas C.C. melanaster).melanaster). KishinouyeKishinouye fn (1899b) found C. fuscescens at Monterey, California andand describeddescribed itit asas aa newnew sp species, C. gilberti.gi/berti. Mayer (1910),(1910), inin his monograph ofofthethe medusaemedusae ofofthetheworld,world, w' was also confused over thethe correct identityidentityoftheseofthese ChrysaoraChrysaora species.species. HeHethoughtthought is that C. helvolahe/vola was a valid species and thatthat C. gilbertigi/berti waswas aa varietyvariety ofofC.C. melan­me/an­ PI aster. Kramp (1961) agreed with Mayer (1910).(1910). AAnumnumberber offieldof field guidesguides havehave S1 used one or more of thesethese names. A third species, C. plocamia, occurs along thethe coastscoasts ofofPeruPeru andand N.N. Chile.Chile. ItIt n has a milky-white umbrella with a dark star-likestar-like patternpatternononthetheexexumbrellaumbrella (Stias­(Stias­ b ny, 1937). In addition, an unidentified speciesspecies ofof ChrysaoraChrysaora waswas reportedreported byby Vanhoffen (1888) from near thethe Straits of Magellan. Ii A fourth species of Chrysaora was seenseen alongalong thethe coastcoast ofofsouthernsouthern CaliforniaCalifornia s· in the summer of 1989. This undescribed speciesspecieshadhadaadarkdarknearlynearlyblackblackumbrella.umbrella. (I Recently, Shenker (1984; 1985) has investigatedinvestigated somesome aspectsaspects ofofthethe ecologyecology ( of C. fuscescens off Oregon. He concluded thatthat itit maymay playplay anan importantimportant rolerole inin ~ the pelagic food web because of itsits high biomass. II havehave seenseen numerousnumerous C.C. ( fuscescens along the northern California and OregonOregon coastscoastsandand II mustmustagreeagree withwith t Shenker concerning thethe extreme abundance ofofthisthis species.species, InIn latelate summersummer thisthis I medusa can be heavily infectedinfected with thethe hyperiidhyperiid HyperocheHyperoche medusarummedusarumandandoffoff LARSON:LARSON:EASTERNEASTERNPACIFICPACIFICSCYPHOMEDUSAESCYPHOMEDUSAEANDANDCUBOMEDUSAECUBOMEDUSAE 553553

California,California,juvenilejuvenile crabscrabs(Cancer(Cancergracilis)gracilis)attachattachtotothethemedusamedusa(J.(1.Cooper,Cooper,pers.pers. comm.,comm., 1976).1976). I TheTheoceanicoceanicgenusgenusPelagiaPelagiaisisrepresentedrepresentedininthetheeasterneasternPacificPacificbybytwotwospecies.species. i PelagiaPelagia noctiluca,noctiluca, aa widelywidelydistributeddistributedspecies,species,isisknownknownfromfromMexicoMexicototoChileChile ! (Table(Table I).1). AA secondsecond species,species, thethe spectacularspectacularpurple-stripedpurple-stripedPelagiaPelagia coloratacolorataisis regularlyregularly washedwashedashoreashorealongalongthethesouthernsouthernCaliforniaCaliforniabeaches,beaches,butbutititcancanoccuroccur northnorthofofSanSan FranciscoFrancisco(J.(J. Rees,Rees, pers.pers. comm.,comm., 1973).1973).PreviousPrevioustotoitsitsdescriptiondescription byby RussellRussell (1964),(1964), ititwaswasreferredreferredtotoasasPelagiaPelagiapanopyrapanopyraPeronPeronandandLesueur,Lesueur,oror PelagiaPelagia cyanellacyanella PeronPeronandandLesueur.Lesueur.BeautifulBeautifulcolorcolorphotographsphotographsofofthisthisspecies,species, showingshowingthethe extremelyextremelylonglongandandpleatedpleatedoraloralarms,arms,werewerepublishedpublishedininSkinSkinDiverDiver magazinemagazine (Oct.(Oct. 1974).1974). Recently,Recently, F.F. SommersSommers(pers.(pers. comm.,comm., 1989)1989)hashasbeenbeenin­in­ vestigatingvestigating thethe developmentdevelopment ofofthisthis speciesspecies andand hashas foundfound thatthatitithashasaa benthicbenthic polyppolyp stage.stage. BecauseBecause thethe onlyonly otherother speciesspecies inin thisthis genus,genus, P.P. noctiluca,noctiluca, lackslacks aa scyphistoma,scyphistoma, P.P. coloratacolorata shouldshould bebe placedplaced inin aa newnewgenus.genus. UnlikeUnlike thethe adultadult formform ofofP.P. colorata,colorata, whichwhichhashascharacteristiccharacteristicpurplepurplestripesstripesonon aa whitewhite umbrella,umbrella,juvenilesjuveniles(up(up totoaboutabout 1010cm)cm)havehaveaa palepalepinkpinkexumbrellaexumbrellaandand tentaclestentacles thatthat areare aa dark-reddishdark-reddish colorcolor (F.(F. Sommers,Sommers, pers.pers. comm.,comm., 1989).1989). ThisThis juvenilejuvenilecolorcolor morph waswas firstfirst reportedreportedfromfrom SantaSantaBarbara,Barbara,CaliforniaCaliforniabybyFewkesFewkes (1889),(1889), as P. panopyra. SinceSince thenthen aa numbernumberofofphotographsphotographsofofthisthis medusamedusahavehave been published (Halstead,(Halstead, 1965,1965, pI.pI. XLVII;XLVII; Gladfelter,Gladfelter, 1973,1973, fig.fig. II;11; GalbraithGalbraith and Boehler, 1974, plts. II,II, III).III). The chemical nature of thethe exumbrellarexumbrellar pigmentpigment ofofP.P. coloratacolorata hashas beenbeen ex­ex­ amined by Fox and Millot (1954)(1954) and Millot andand FoxFox (1954).(1954). Cyanea capcapillataillata is a well known cold water speciesspecies whichwhich hashas beenbeen thoroughlythoroughly studied in the Atlantic (Russell, 1970). However, littlelittle isis knownknown aboutabout thisthis largelarge medusa from the Pacific coast. It is seasonally abundant alongalong thethe coastcoast fromfrom Washington northward. It is less numerous off Oregon (Shenker,(Shenker, 1985; Larson,Larson, unpub. obs.), but does occur as far south as Monterey Bay, California (F.(F. Sommers, pers. comm., 1990). A number of other Cyanea spp. were described fromfrom thethe northern Pacific (C ferruginea Eschscholtz, 1829; C postelsi Brandt, 1838) butbut these are probably synonymous with CC. capillata (Mayer, 1910; Kramp, 1961). Gladfelter (1972) studied the functional morphology of the locomotory system. Phacellophora camtschatica is another large medusa reaching a diameter of at leastleast 50 cm.em. It has an extensive range in the eastern Pacific and apparently occurs fromfrom AlaskaAlaska toto Chile (Larson, 1986). In waters around Vancouver Island this speciesspecies isis common during thethe summer months. It resembles a giant bird's egg whichwhich hashas been crackedcracked and poured onto the water's surface. The thin umbrella isis translucenttranslucent toto milky except near thethe center which is a bright, egg yolk-yellow. PhacellophoraPhacellophora camtschaticacamtschatica has aa number of juniorjunior synonyms (Kramp, 1961). StrandStrandandandHamnerHamner(1988)(1988)recentlyrecentlydescribed thetheforagingforaging behavior ofthis species. OneOne otherother semaeostomesemaeostome genusgenus isis likelylikely toto occur inin thethe eastern Pacific. Des­ monemamonema spp.spp. isis knownknown fromfrom TierraTierra deldel FuegoFuego (Larson,(Larson, 1986) and probably will bebe foundfound furtherfurther northnorth alongalong thethe coastcoast ofofsouthsouth Chile.

Rhizostomeae.Rhizostomeae.-- ThisThis groupgroup isis mostlymostly tropicaltropical inin distributiondistribution andand only a few speciesspecies havehave beenbeen reportedreported fromfrom thethe easterneastern Pacific.Pacific. TwoTwo speciesspecies ofof CatostylusCatostylus (C(C ornatellasornatellasandand CC tagi)tagi) havehave beenbeen reportedreported fromfrom thethe tropicaltropical easterneastern PacificPacific (Vanhoffen,(Vanhoffen, 1888;1888; Stiasny,Stiasny, 1922).1922). SpecimensSpecimens ofof CatostylusCatostylus ?ornatellus,?ornatellus, atat thethe ScrippsScripps InstitutionInstitution ofofOceanographyOceanography (SIO)(SIO) museum,museum, werewere collectedcollected fromfrom BajaBaja CaliforniaCalifornia (31°N,(31°N, 1116°W)16°W) andand II examinedexamined specimensspecimens atat thethe SmithsonianSmithsonian Insti­Insti­ tutiontution fromfrom thethe GulfGulfofofPanama.Panama. AlsoAlso inin thethe SIOSIO museummuseum areare specimensspecimens ofofaa possiblypossibly undescribedundescribed rhizostome.rhizostome. LychnorhizaLychnorhiza sp.sp. waswas collectedcollected alongalong thethe westwest coastcoastofofMexicoMexicobetweenbetween 17°NI rNandand 24°N.24°N. 554554 BULLETINBULLETINOFOFMARINEMARINESOENCE,SOENCE,VOL.VOL.47,47,NO.2,NO.2,19901990

StomolophusStomolophus meleagrismeleagris hashasbeenbeen reportedreported betweenbetweenS.S. CaliforniaCaliforniaandandEcuadorEcuador 1 (Table(Table 1).1). ItIt isis sometimessometimes referredreferred toto asas thethe "cabbage"cabbagehead"head"oror"cannon"cannonball"ball" Cald, becausebecause ofofitsitsgloboseglobose umbrellaumbrellaandandshort,short,thick,thick,gelatinousgelatinousoraloralarms.arms.AlongAlongthisthis I coastcoast itit usuallyusually isis aa pTUssianpTUssian bluebluecolor,color,whereaswhereasininthetheAtlanticAtlanticitithashasaamilkymilkyoror Chia milkymilky withwith darkdark chocolatechocolate pigmentationpigmentation onon thethe exumbrella.exumbrella. ItsIts lifelife cyclecycle waswas Oar' describeddescribed byby CalderCalder(1982).(1982). Recently,Recently, aa previouslypreviously unreportedunreported rhizostome,rhizostome, PhyllorhizaPhyllorhiza punctata,punctata, hashas ap­ap­ Cro' pearedpeared inin numbersnumbers inin SanSan DiegoDiego Bay,Bay, CaliforniaCalifornia (Larson(Larson andand A.A. Arneson,Arneson, inin Dev press).press). PhyllorhizaPhyllorhiza punctatapunctata maymay havehave beenbeen transportedtransported fromfrom HawaiiHawaii asas polypspolyps Fagl attachedattached toto shipshiphulls.hulls. ItItreachesreachesaa bellbelldiameterdiameterofofupuptoto5050em.em.TheThebellbellisisbluishbluish Fan with regularlyregularly scatteredscattered whitewhite spotsspots andand thethe longlongoral-armoral-armclubsclubs areare tippedtippedwithwith blue and white.white. PhotosPhotos ofofthisthis medusamedusa appearappear inin ReedReed (1971)(1971) andand DevanyDevanyandand Fe'" Eldridge (1977).(1977). Fox With furtherfurther collectingcollectinginshoreinshorealongalongthetheinsufficientlyinsufficientlystudiedstudiedcoastcoastofofMexico,Mexico, Gal and Central and SouthSouth America,America, additionaladditional rhizostomesrhizostomes maymaybebe found.found. Gla ACKNOWLEDGMENTSACKNOWLEDGMENTS

I wish toto thankthank M. Arai, A. Alvariiio, C.C. Arneson,Arneson, L.L. Barr,Barr, J.J. Cooper,Cooper, P.P. Corbin,Corbin,Y.Y. SchnorrSchnorrvonvon Carolsfeld, D. Fautin, C. Mills, J.J. Rees, F.F. Sommers,Sommers, G.G. Snyder,Snyder, andandJ.J. WordWordforforkindlykindlysendingsendingmeme G'I' valuable medusan material, usefuluseful photographs, oror importantimportant distributionaldistributional data.data. C.C. MilJsMills andand anan anonymous reviewerreviewermade highlyhighlyhelpfulhelpfulcommentscommentsononthethemanuscript.manuscript.K.K.LarsonLarsongenerouslygenerouslyprovidedprovided two photos forfor thetheplate. This isis contributioncontributionnumber 725725ofofHarborHarborBranchBranchOceanographicOceanographicInstitution.Institution. Ha LITERATURE CITED LITERATURE CITED H~ Agassiz, A. 1862 (in[in L. Agassiz, 1862).1862]. Pages 1-380 inin ContributionsContributions toto thethe naturalnatural historyhistory ofofthethe United States of America, Vol. 4. Little, Brown, Boston. Alvariiio, A. 1967. Bathymetric distribution of Chaetognatha, Siphonophorae,Siphonophorae, Medusae,Medusae, andand Oe­Oe­ nophorae off San Diego, California. Pac. Sci. 21: 285-474.285-474. H: --. 1969. Zoogeographia del Mar de Cortes; Quetognatos, Sifonoforos && Medusas.Medusas. An.An. lost.Inst. BioI.Bio\. Univ. Na\.Nal. Auton. Mexico 40. Ser.SeT. Cien. de Mar yy Limnol. I:I: II-54.II-54. H --.-_. 19.72. Pages 223-247 in Zooplancton del Caribe, Golfo dede Mexico yy regionesregionesadjacentesadjacentesdeldel H Pacifico. Memorias IV Congreso Nacional de Oceanografia (Mexico).(Mexico). --.-_. 1976. Los indicadores planctonicos: distribucion batimetrica dede alguanasalguanas medusas.medusas. PagesPages K 161-190 in Memorias del II Simposio Latinoamericano sobre Oceanographia Biologica.Biologica. Uni­Uni­ versidad de Oriente, Venezuela. --and M. J. L. A. Ambros. 1986. EI zooplancton del Pacifico equatoriano. Iov.Inv. Mar. CicimarCicimar K 3: 69-110. Barham, E. G. 1969. A window in the sea. Oceans I: 5~0. Bigelow, H. B. 1909. Reports on the scientific resuJtsresults ofthe expedition to thethe eastern tropicaltropical Pacific, inin charge of Alexander Agassiz, by the U.S. Fish Commission steamer 'Albatross:'Albatross: fromfrom October 1904, toto March 1905, Lieut. Commander L. M. Garrett, U.S.N., commanding. XVI. The medusae. Mem. Mus Compo Zool.Zoo\. Harv. 37: 1-243. --. 1913. Medusae and Siphonophrae collected by the U.S. Fisheries steamer 'Albatross''Albatross' inin thethe north-westnorth-west Pacific, 1906. Proc. U.S. Nat. Mus. 44: 1-119.I-I 19. --. 1914. Note on thethe medusan genus Stomolophus from San Diego. Univ. Calif. Pub!.Publ. Zoo!.Zool. 13: 239-241.239-24 I. --.--. 1920.1920. Pages +.23 inin Medusae and ctenophores from the Canadian Arctic Expedition, 1913­ 18.18. Rep. Can. Arct. Exped. 8, panpart H. Medusae and Oenophora. King's Printer, Ottawa. --.--. 1928.1928. Scyphomedusae of thethe Arcturus Expedition. Zoologica 8: 495-524. --.--. 1940.1940. Medusae ofthethe Templeton Crocker and Eastern Pacific zacaZaca Expeditions, 1936-1938. ZoologicaZoologica 25:25: 281-321.281-321. Boone,Boone, L. 1933.1933. Scientific resultsresultsofthethecruises ofthe yachts 'Eagle''Eagle' and 'Ara,' 1921-1928. Coelentera­ ta,ta, EchinodermataEchinodermata andand Mollusca. Bull. Vanderbilt Oceanogr. (Mar.) Mus. 4: 1-217. --.--. 1938.1938. TheThe marinemarine algae.algae. coelenteratacoelenterata ... of the world cruises of the yachts 'Ara: 1928­ 1929,1929, andand 'Alva,''Alva,' 1931-1932,1931-1932, 'Alva''Alva' Mediterranean cruise 1933, and 'Alva''Alva' South American cruise.cruise. Bull.BuJl. VanderbiltVanderbilt Oceanogr.Oceanogr. (Mar.)(Mar.) Mus. 7: 27-76. Brandt,Brandt, J.J. F.F. 1835.1835. ProdromusProdromus descriptionisdescriptionis animaliumanimalium ab Mertensio in orbis terrarum circumna­ vigationevigatione observatorum,observatorum, Fasc.Fasc. I.I. Rec. Act. Acad. Imp.Imp. Sci. St.-Petersburg, 1834.1834.7575 pp. LARSON:LARSON:EASTERNEASTERNPACIFICPACIFICSCYPHOMEDUSAESCYPHOMEDUSAEANDANDCUBOMEDUSAECUBOMEDUSAE 555555

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Maas, O. 1897. Report on an exploration off the west coast of Mexico, Central and South America, and off the Galapagos Islands, XXI. Die Medusen. Mem, Mus. Compo Zoo!. Harv. 23: 1-92. MacGinitie, G. E. 1955. Distribution and ecology of the marine invertebrates of Point Barrow, Alaska. Smithson. Misc. Coil. 128: 1-201. Mayer, A. G. 1910. Medusae of the world. III. The Scyphomedusae. Carnegie Inst. Wash. Pub!. 109: 499-735. Millot, N. and D. L. Fox. 1954. Pigmentation of the Jellyfish, Pelagia noctiluca. Nature 173: 169- 171. Mills, C. E. 1981. Seasonal occurrence of planktonic medusae and ctenophores in the San Juan Archipelago (N.E. Pacific). Wasmann J. BioI. 39: 6-29. Moller, H. 1984. Reduction of a larval herring population by a jellyfish predator. Science 224: 621- 622. Otto, J. 1976. Early development and planula movement in Haliclystus (Scyphozoa: Stauromedusae). Pages 319-330 in G. O. Mackie, ed. Coelenterate ecology and behavior. Plenum Press, New York. Pearcy, W. G. 1972. Distribution and ecology of oceanic animals off Oregon. Pages 351-377 in A. T. Pruter and D. L. Alverson, eds. The Columbia River Estuary and adjacent ocean waters. Univ. Washington Press, Seattle. ---. 1976. Seasonal and inshore-offshore variations in the standing stocks of micronekton and macrozooplankton off Oregon. Fish. Bull. U.S. 74: 70-80. --- and M. Stuiver. 1983. Vertical transport of carbon-I 4 into deep-sea food webs. Deep-sea Res. 30: 427-440. Pearse, 1. S. and V. B. Pearse. 1978. Vision in cubomedusan . Science 199: 458. Reed, S. A. 1971. Some common coelenterates in Kaneohe Bay, Oahu, Hawaii. Pages 37-51 in H. M. Lenhoff, L. Muscatine and L. V. Davis, eds. Experimental coelenterate biology. Univ. Hawaii Press, Honolulu. Russell, F. S. 1964. On scyphomedusae of the genus Pelagia. J. Mar. Bio!. Ass. U.K. 44: 133-136. --. 1967. On a remarkable new scyphomedusan. J. Mar. BioI. Ass. U.K. 47: 469-473. ---. 1970. The medusae of the British Isles. II. Pelagic Scyphozoa. Cambridge Univ. Press, London. 284 pp. Satterlie, R. A. 1979. Central control of swimming in the cubomedusan jellyfish Carybdea rastonii. J. Compo Physiol. 133: 357-367. -- and A. N. Spencer. 1979. Swimming control in a cubomedusanjellyfish. Nature 281: 141- 142. Segura-Puertas, L. 1984. Morphology, systematics and zoogeography of medusae (Cnidaria: Hydro- zoa and Scyphozoa) from the eastern tropical Pacific. Inst. Cienc. del Mar y Limnol. Univ. Nal. Auton. Mexico. Pub!. Esp. 8: 1-320, Shenker, J. M. 1984. Scyphomedusae in surface waters near the Oregon Coast, May-August, 1981. Est. Coast. Shelf Sci. 19: 619-632. ---. 1985. Carbon content of the neritic scyphomedusa Chrysaorafuscescens. J. Plank. Res. 7: 169-173. Shih, C.-T., A. J. G. Figueira and E. H. Grainger. 1971. A synopsis of Canadian marine . Fish. Res. Bd. Can. Bull. 176. 264 pp. Smith, K. L. 1982. Zooplankton of a bathyl benthic boundary layer: in situ rates of oxygen con- sumption and ammonium excretion. Limno!. Oceanogr. 27: 461-471. Stiasny, G. 1922. Die Scyphomedusen-Sammlung von Dr. Th. Mortensen nebst anderen Medusen aus dem Zoologischen Museum der Universitiit in Kopenhagen. Pap. from Dr. Th. Mortensen's Pac. Exped. 1914-1916. XIII. Vid. Medd. Dansk Naturhist. Foren. Kbh. 73: 513-555. --. 1937. Uber Chrysaora plocamia (Lesson) Haeckel von Toxopilla (Nord-Chile). Zoo!. Anz. 119: 18-284. --- and H. van der Maaden. 1943. Uber Scyphomedusaen aus dem Ochotskischen und Kam- tschatka Meer nebst einer Kritik der Genera Cyanea und Desmonema. Zool. Jb. 76: 228-266. Uchida, T. and Z. Nagao. 1963. The metamorphosis of the scyphomedusa, Aurelia limbata (Brandt). Annot. Zool. 1. 36: 83-91. Vanhoffen, E. 1888. Untersuchungen iiber semaeostome und rhizostome Medusen. Bibliotheca Zoologica I: I-52. Werner, B. 1975. Bau und lebensgeschichte des Polypen von Tripedalia cystophora (Cubozoa, class. nov., Carybdeidae) und seine Bedeutung fUr die Evolution der Cnidaria. Helgo. Wiss. Meeresun- ters. 27: 461-504.

DATEACCEPTED: June 26, 1989.

AnDRESS: Harbor Branch Oceanographic Institution, Inc., 5600 Old Dixie Highway. Fort Pierce, Florida 34946. 556 BULLETINOFMARINESCIENCE,VOL.47, NO.2, 1990

Maas, O. 1897. Report on an exploration off the west coast of Mexico, Central and South America, and off the Galapagos Islands, XXI. Die Medusen. Mem, Mus. Compo Zoo!. Harv. 23: 1-92. MacGinitie, G. E. 1955. Distribution and ecology of the marine invertebrates of Point Barrow, Alaska. Smithson. Misc. Coil. 128: 1-201. Mayer, A. G. 1910. Medusae of the world. III. The Scyphomedusae. Carnegie Inst. Wash. Pub!. 109: 499-735. Millot, N. and D. L. Fox. 1954. Pigmentation of the Jellyfish, Pelagia noctiluca. Nature 173: 169- 171. Mills, C. E. 1981. Seasonal occurrence of planktonic medusae and ctenophores in the San Juan Archipelago (N.E. Pacific). Wasmann J. BioI. 39: 6-29. Moller, H. 1984. Reduction of a larval herring population by a jellyfish predator. Science 224: 621- 622. Otto, J. 1976. Early development and planula movement in Haliclystus (Scyphozoa: Stauromedusae). Pages 319-330 in G. O. Mackie, ed. Coelenterate ecology and behavior. Plenum Press, New York. Pearcy, W. G. 1972. Distribution and ecology of oceanic animals off Oregon. Pages 351-377 in A. T. Pruter and D. L. Alverson, eds. The Columbia River Estuary and adjacent ocean waters. Univ. Washington Press, Seattle. ---. 1976. Seasonal and inshore-offshore variations in the standing stocks of micronekton and macrozooplankton off Oregon. Fish. Bull. U.S. 74: 70-80. --- and M. Stuiver. 1983. Vertical transport of carbon-I 4 into deep-sea food webs. Deep-sea Res. 30: 427-440. Pearse, 1. S. and V. B. Pearse. 1978. Vision in cubomedusan jellyfishes. Science 199: 458. Reed, S. A. 1971. Some common coelenterates in Kaneohe Bay, Oahu, Hawaii. Pages 37-51 in H. M. Lenhoff, L. Muscatine and L. V. Davis, eds. Experimental coelenterate biology. Univ. Hawaii Press, Honolulu. Russell, F. S. 1964. On scyphomedusae of the genus Pelagia. J. Mar. Bio!. Ass. U.K. 44: 133-136. --. 1967. On a remarkable new scyphomedusan. J. Mar. BioI. Ass. U.K. 47: 469-473. ---. 1970. The medusae of the British Isles. II. Pelagic Scyphozoa. Cambridge Univ. Press, London. 284 pp. Satterlie, R. A. 1979. Central control of swimming in the cubomedusan jellyfish Carybdea rastonii. J. Compo Physiol. 133: 357-367. -- and A. N. Spencer. 1979. Swimming control in a cubomedusanjellyfish. Nature 281: 141- 142. Segura-Puertas, L. 1984. Morphology, systematics and zoogeography of medusae (Cnidaria: Hydro- zoa and Scyphozoa) from the eastern tropical Pacific. Inst. Cienc. del Mar y Limnol. Univ. Nal. Auton. Mexico. Pub!. Esp. 8: 1-320, Shenker, J. M. 1984. Scyphomedusae in surface waters near the Oregon Coast, May-August, 1981. Est. Coast. Shelf Sci. 19: 619-632. ---. 1985. Carbon content of the neritic scyphomedusa Chrysaorafuscescens. J. Plank. Res. 7: 169-173. Shih, C.-T., A. J. G. Figueira and E. H. Grainger. 1971. A synopsis of Canadian marine zooplankton. Fish. Res. Bd. Can. Bull. 176. 264 pp. Smith, K. L. 1982. Zooplankton of a bathyl benthic boundary layer: in situ rates of oxygen con- sumption and ammonium excretion. Limno!. Oceanogr. 27: 461-471. Stiasny, G. 1922. Die Scyphomedusen-Sammlung von Dr. Th. Mortensen nebst anderen Medusen aus dem Zoologischen Museum der Universitiit in Kopenhagen. Pap. from Dr. Th. Mortensen's Pac. Exped. 1914-1916. XIII. Vid. Medd. Dansk Naturhist. Foren. Kbh. 73: 513-555. --. 1937. Uber Chrysaora plocamia (Lesson) Haeckel von Toxopilla (Nord-Chile). Zoo!. Anz. 119: 18-284. --- and H. van der Maaden. 1943. Uber Scyphomedusaen aus dem Ochotskischen und Kam- tschatka Meer nebst einer Kritik der Genera Cyanea und Desmonema. Zool. Jb. 76: 228-266. Uchida, T. and Z. Nagao. 1963. The metamorphosis of the scyphomedusa, Aurelia limbata (Brandt). Annot. Zool. 1. 36: 83-91. Vanhoffen, E. 1888. Untersuchungen iiber semaeostome und rhizostome Medusen. Bibliotheca Zoologica I: I-52. Werner, B. 1975. Bau und lebensgeschichte des Polypen von Tripedalia cystophora (Cubozoa, class. nov., Carybdeidae) und seine Bedeutung fUr die Evolution der Cnidaria. Helgo. Wiss. Meeresun- ters. 27: 461-504.

DATEACCEPTED: June 26, 1989.

AnDRESS: Harbor Branch Oceanographic Institution, Inc., 5600 Old Dixie Highway. Fort Pierce, Florida 34946.