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Notice: ©1987 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: Pugh, P. R., & Harbison, G. R. (1987). Three new species of prayine siphonophore (Calycophorae, Prayidae) collected by a submersible, with notes on related species. Bulletin of Marine Science, 41(1), 68-91. BULLETINBULLETINOF MARINEOF SCIENCE.MARINE SCIENCE,41(1): 68-91.68-91,1987 41(11987): 68-91 ,1987
THREETHREENEW NEWSPECIESSPECIESOF PRA OFYINEYlNEPRAYINESIPHONOPHORESIPHONOPHORE (CALYCOPHORAE,(CALYCOPHORAE,PRAYIDAE)PRAYIDAE)COLLECTEDCOLLECTEDBY A BY A SUBMERSIBLE,SUBMERSIBLE,WITHWITHNOTESNOTESON RELATEDON RELATEDSPECIESSPECIES
P. R. PughP. R.and PughG.andR. HarbisonG. R. Harbison
ABSTRACTABSTRACT Three newThreespecies newof speciessiphonophore of siphonophorebelongingbelongingto the calycophoran to the calycophoransub-familysub-familyPrayinaePrayinaeare are described.described.The individual The individualspecimensspecimensof these ofwere thesecollected, were collectedin the Bahamas, in the Bahamasregion, by regionthe , by the submersiblesubmersibleDSRV }OHNSON-SEA-LINKJOHNSON-SEA-LrNKDSRV ]OHNSON-SEA-LINKII, usingII,her usingsophisticated her sophisticatedsamplingsamplingdevices. devices.One of One of the newthespecies newhas speciesbeen hasascribed been ascribedto the genus to thePrayola genus PrayolaCarre, 1969.Carre,The 1969.other Thetwoothershowtwo show characterscharactersthat are thatintermediateintennediateare intermediatebetweenbetweenthose found thosein foundexisting in existingprayine genera prayineand generahave and have necessitatednecessitatedthe erectionthe erectionoftwo newoftwogenera,newCraseoa genera, Craseoaand Mistoprayina.and Mlstoprayinato encompass, to encompassthem. them. The morphologicalThe morphologicalcharacterscharactersthat are thatof taxonomicare of taxonomicsignificancesignificancein prayinein siphonophoresprayine siphonophores are reviewed,are reviewedand a reclassification, and a reclassificationofone existing of onespecies, existingRosacea species, RosaceavillqfrancaeCarre,villa/rancaevillafrancaeCarre, 1969,Carre, 1969, is proposed.is proposed.
AlthoughAlthoughmany speciesmany speciesof siphonophoresof siphonophorescan becancollected be collectedadequatelyadequatelyby netby net samplingsamplingsystems,systemsit has, becomeit has becomeapparentapparentin recentin recentyears thatyearstherethataretherea large,are a large, and everandincreasing,ever increasing,numbernumberthat arethatnot.areFor not.instance,For instance,SCUBASCUBAdivingdivingtechniquestechniques have shownhave shownthat manythat ofmanythe apparentlyofthe apparentlyrare siphonophorerare siphonophorespecies,species,describeddescribedin in the lastthecenturylast centurybut notbutseennotsince,seenactually since, actuallyare quiteare common.quite common.This apparentThis apparent paradoxparadoxis explainedis explainedeasily byeasilythe byfact thethat factthethatanimalsthe animalsare extremelyare extremelyfragile fragileand, and, as theyaswouldthey wouldbe tombetotomshreds,to shredsdo not, doappearnot appearin net collections.in net collections. Submersibles,Submersibles,with theirwithsophisticatedtheir sophisticatedsamplingsamplingdevices,devices,have madehaveitmadepossibleit possible to extendto extendthese inthesesituinstudiessitu studiesto greaterto greaterdepths,depths,and withandsignificantwith significanteffect. Oneeffect. One importantimportantfeaturefeatureof theseofinthesesituinsampling situ samplingmethodsmethodsis thatisthethatanimalsthe animalsare colare col lected individually.lected individually.In the caseIn theofcasesiphonophores,ofsiphonophores,whose whosestems canstemsstretchcan stretchout to out to many metersmany metersin length,in length,it is notit isalwaysnot alwayspossiblepossibleto captureto capturethe wholethe wholeanimal.animal. However,However,even ifevenonlyifpartonlyofpartthe animalof the animalis collectedis collectedand evenandif,evensubsequently, if, subsequently, this becomesthis becomesdissociateddissociatedinto itsintocomponentits componentpieces,pieces,one knows one knowsunequivocallyunequivocallythat that all theseallpiecesthese piecesbelongbelongto a singleto aspecimen. single specimen. DuringDuringa seriesaof seriesdivesof(OctoberlNovemberdives (OctoberlNovember1984) in1984)the invicinity the vicinityof the ofBathe Ba hamas,hamas,over 130overindividual130 individualspecimensspecimensofsiphonophoresofsiphonophoreswere collectedwere collectedusing theusing the submersiblesubmersibleDSRVDSRV]OHNSON-SEA-LINKJOHNSON-SPA-LINK]OHNSON-SEA-LINKII. A fullII. Alist fullof listtheseofdivesthese anddivesa andbriefa brief descriptiondescriptionof the ofsamplingthe samplingdevicesdevicesused isusedgivenisin givenPugh inand PughHarbisonand Harbison(1986).(1986). These Thesedevicesdevicesconsistconsistof slurpofguns,slurpwherebyguns, wherebyeven 10+evenm-Iong10+ m-longsiphonophoressiphonophores can becangently besucked gently suckedinto individualinto individualcontainers,containers,and detritusand detritussamplers,samplers,into whichinto which the requiredthe requiredanimalanimalis broughtis broughtby delicateby delicatemaneuveringmaneuveringoftheofthe submersible.ofthe submersible.More More than 30thanspecies30 speciesofsiphonophoresofsiphonophoreswere identifiedwere identifiedfrom thesefromcollectionsthese collectionsand aboutand about half ofhalftheseofweretheseundescribed.were undescribed.AmongstAmongstthelatter the werelatterthree,were belongingthree, belongingto the to the calycophorancalycophoransub-familysub-familyPrayinae,Prayinae,which whichare describedare describedhere. Onehere.species One speciesclearlyclearly is attributableis attributableto the genusto the Prayolagenus PrayolaCarre, Carre,1969, while1969,the whileother the twoothershowtwochar- show char- .acters.actersthat arethatintermediateare intermediatebetweenbetweenthose foundthose foundin existingin existinggenerageneraand haveand have necessitatednecessitatedthe erectionthe erectionof twoofnewtwogenera, new genera,CraseoaCraseoaand Mistoprayina,and Mistoprayina,to en to en compasscompassthem. them. 68 68 PUGH AND HARBISON: NEW SPECIES OF PRAYINE SIPHONOPHORE 69 PUGH AND HARBISON: NEW SPECIES OF PRAYINE SIPHONOPHORE 69
CLASSIFICA nON CLASSIFICAnON Sub-order Calycophorae Leuckart, 1854 Sub-orderFamilyCalycophoraePrayidae K6lliker,Leuckart,18531854 Sub-familyFamily PrayidaePrayinaeK6lliker,Chun, 18531897 Sub-family Prayinae Chun, 1897 Craseoa new genus Craseoa new genus Diagnosis. - Prayine siphonophores with an apposed pair of cylindrical necto- Diagnosis.phores. The- Prayinesomatocystsiphonophoresis simple, withoutwith an eitherapposedan pairascendingof cylindricalor descendingnecto phores.branch. The lateralsomatocystradialiscanalssimple,on withoutthe smalleithernectosacan ascendingare S-shaped.or descendingThe bracts branch.are roundedThe andlateraldividedradialintocanalstwo onalmostthe smallequal nectosaclobes by aaredeepS-shaped.fissure. SixThebractealbracts arecanalsroundedare present.and dividedThe gonophoresinto two almostare equalasymmetriclobes byinashapedeep fissure.with wing-likeSix bractealex- canalspansions.are Thepresent.sub-umbrellaThe gonophorescavity isarerelativelyasymmetricsmall,inoccupyingshape withonlywing-likepart of exthe pansions.lower halfTheof thesub-umbrellagonophore. cavityThe arrangementis relativelyofsmall,the twooccupyingmantle canalsonly partis slightlyof the lowerasymmetric.halfoftheSpecial,gonophore.asexualThenectophoresarrangementare absent.ofthe two mantle canals is slightly asymmetric. Special, asexual nectophores are absent. Type Species. - C. lathetica new species described herein. Type Species. - C. lathetica new species described herein. Etymology. - The generic name Craseoa can be derived from the Greek Kpauur, Etymology.KpaUEWU meaning- Theagenericblending,namereferringCraseoato thecanmixturebe derivedof prayinefrom thecharactersGreek Kpauur,found KpaUEWUin the specimens.meaning aItblending,is also anreferringanagramtooftheRosacea.mixture ofprayine characters found in the specimens. It is also an anagram ofRosacea. Remarks. - The outward appearance of the nectophores of C. lathetica. in that Remarks.they are cylindrical- The outwardand possessappearancerelativelyof thesmallnectophoresnectosacs,of C.is similarlathetica.to inthatthatin theythe speciesare cylindricalof two otherand prayinepossess genera,relativelyRosaceasmall andnectosacs,Desmophyes.is similarHowever,to thatthein theabsencespeciesof ofbothtwoanotherascendingprayineandgenera,descendingRosaceabranchand Desmophyes.to the simpleHowever,somatocyst,the absencetogether ofwithboththeanS-shapedascendingcourseand ofdescendingthe lateralbranchradial canalsto the onsimplethe nectosac,somatocyst,are togetherimportantwithmorphologicalthe S-shapedcharacterscourse of(seethe lateralTable radial1)and canalspreventonthetheinclusionnectosac,ofareC. importantlathetica inmorphologicaleither of thesecharactersgenera. Nonetheless,(see Table 1)theseandcharactersprevent theapplyinclusionalso tooftheC. latheticaspecies ofintheeithergenusofthesePrayola,genera.but Nonetheless,the shape of thesethe nectophore,characters applyand particularlyalso to the species of the genus Prayola, but the shape of the nectophore, and particularly
Figure 1. Craseoa lathetica. Photographs of the living holotype specimen. A. Nectosome. Scale = 5 Figuremm; BI.a cormidialCraseoa lathetica.group. ScalePhotographs= 2 mm. Photosof the livingby R.holotypeW. Gilmer.specimen. A. Nectosome. Scale = 5 mm; B a cormidial group. Scale = 2 mm. Photos by R. W. Gilmer. 70 70 BULLETIN OFBULLETINMARINEOFSCIENCE.MARINEVOLSCIENCE.41. NO.VOLI. 198141,41. NONO.1..1, 1987 c c ----
~~-----~~---'~--~""
Figure 2. NectophoresFigure 2. Nectophoresof CraseoaoflalhelicaCraseoa(JSLlathetica 996 specimen):(JSL 996 specimen):specimen)A. Ventral: AA.and. VentralB. Lateraland B.viewsLateralof views of the N, nectophore. C. Ventral and D. Lateral views of the N, nectophore. Scale = 1 em. the N, nectophore.the N, nectophore.C. VentralC.andVentralD. Lateraland D.viewsLateralof theviewsN 2 nectophore.of the N 2 nectophore.Scale = I em.Scale = I cm. the nectosac,the nectosac,is very differentis very differentand Prayolaand Prayolaspecies arespeciesthe onlyare theprayineonly siphonoprayine siphono phores tophoreshave onlyto havefiveonlycanalsfiveincanalstheir bracts,in theirthebracts,dorsaltheonedorsalbeingoneabsent.beingTheabsent. The stem groupsstemofgroupsC. latheticaof C. latheticadiffer fromdifferall fromotherall prayineother speciesprayineinspeciesthat the in thatgonthe gon ophores ophoreshave a relativelyhave a relativelysmall sub-umbrellasmall sub-umbrellacavity. Allcavity.theseAllcharactersthese characterstogether together necessitatenecessitatethe establishmentthe establishmentofa newofgenus,a newCraseoa;genus, Craseoa;to includetotheincludenew speciesthe newspecies which iswhichdescribedis describedbelow. below.
Craseoa Craseoalatheticalatheticanew speciesnew species Figures 1-5Figures 1-5
Material Examined.-Material ExaminedExamined.-Three specimens.- Three specimenscollected incollectedthe regionin theof theregion Bahamasof the byBahamasthe submersibleby the submersible JOHNSON-SEA-LlNKJOHNSON-SEA-LINKJOHNSON-SEA-LINKIIII in 1984.II in 1984. Holotype.-SpecimenHolotype. -Specimencollected atcollecteda depth of607at a depthm usingof607a detritus m usingsamplera detritusduringsamplerJSLduringdive 996.JSL(31dive 996. (31 October 1984;October2so22.SN.19841984;; 2so2225°22.5N.77°S4.9'W)..SN, nOS4.9'W).77°54.9'W).Preserved Preservedin 5% fonnalin.in S%5% Presentedformalin. 10Presentedthe Britishto theMuseumBritish Museum (Natural History)(NaturalandHistory)enteredandas enteredRegd. No.as Regd.1986-6-1-1. No. 1986-6-1-1. Paratypes.-IndividualParatypes.-IndividualParatypes. -Individualspecimens specimenscoUected duringcollectedJSLduringdives 959JSL anddives998.9599S9JSLanddive998.959:JSL 17dive October959:9S9: 17 October 1984; 26"25.6'N.1984;26"25.6'N. 26°2S.6'N7S·44.5/W;, 78°44.5'W;78°44.S'W;Detritus sampler;DetritusDepthsampler;ofcoUection:Depth ofcollection:715 m. JSL71S715dive mm.998:, JSL31dive October998: 998: 31 October 1984; 2so22.0'N,1984; 2so22.025°22.0'N.77°51.6'W;'N, 77°51.6'W;nOSI.6'W;Slurp gun; SlurpDepth gun;ofcollection:Depth ofcollection:659 m; Temperature:6596S9 m; Temperature:IO.3OC. These10.3"C.10.3"C.two These two specimens specimensare housedarein thehousedbiologicalinthe collectionsbiological collectionsofthe InstituteoftheofInstituteOceanographicofOceanographicSciences. Wonnley.Sciences.Sciences, WormleyWormley.. Diagnosis.DiagnosisDiagnosis.- A monotypic.-A- A monotypicgenus andgenusspeciesandofspeciesprayineofsiphonophore,prayine siphonophore,whose prinwhose prin cipal characterscipal charactershave beenhavegivenbeenin giventhe diagnosis in the diagnosisofthe genus.oftheIn genusgenus.this species. In thisthespecies the PUGH AND HARBISON: NEW SPECIES OF PRA YINE SIPHONOPHORE PUGH ANDPUGHHARBISON:AND HARBISON:NEW SPECIESNEW SPECIESOF PRAOFYINEPRASIPHONOPHOREVINE SIPHONOPHORE 71 71
A A
. " ~. "",-,-- . ~
FigureFigure3. Nectophores3. 3. Nectophoresof Craseoaof Craseoalatheticalathetica (JSL 959(JSLspecimen):959 specimen): A. Ventro-IateralA. Ventro-Iateraland B.andLateralB. Lateral views viewsofN, ofofN,nectophore.N I nectophore.C. Ventro-IateralC. Ventro-IateralVentra-lateraland D.andLateralD. Lateral views views ofN~ ofnectopbore.N, nectopbore.neetophore.Scale =ScaleI em.= 1Icm. em. dorsaldorsalbractealbractealcanalcanalarisesarisesfromfromthe distalthe distalend ofendtheofrightthe right(posterior)(posterior)longitudinallongitudinal canal.canal. Description.Description.- NECTOPHORES-NECfOPHORES- NECfOPHORES(Figs.(Figs.IA, 2, 1A,lA,3).2, Each 3).Each specimenspecimenpossessespossessestwo apposedtwo apposed nectophoresnectophores(N. and(NI(NINand2) (Fig.N 22)){Fig.(Fig.IA). The1A).lA). nectophoresThe nectophoresare rounded,are rounded,roundedcylindrical, cylindricaland and bear bearno pronouncedno pronouncedridgesridgesor visibleor visiblepigmentation.pigmentation.pigmentationThe. NtThenectophoresN I1 nectophoresvary varyin in size fromsize from14.5 14.5mm mmin lengthin lengthby 12.5by 1212.5mm.5 mmin widthin width(JSL(JSL996 specimen)996 specimen)to 23.5to 23.5 by 19.5 mm (JSL 959); the N 2 nectophore from 14.5 x 11 mm (JSL 996) to by 19.5 mm (JSL 959); the N 2 nectophore from 14.5 x II mm (JSL 996) to 21.5 21.5x 18xmm18 (JSLmm (JSL959).959). The hydroecium of the N I nectophore is a broad gutter extending along the The hydroeciumThe hydroeciumof theofNtheI nectophoreN nectophoreis a broadis a broadguttergutterextendingextendingalongalongthe the entireentirelengthlengthof theofventralthe ventralsurface,surface,althoughalthoughappearingappearingonly onlyas a asverya veryslightslight indentation at the apical and basal ends (Figs. 2A, 3A). In the N nectophore the indentationindentationat theatapicalthe apicaland basaland basalendsends(Figs.(Figs.2A, 3A).2A, 3A).In theInNthe2 nectophoreN 2 nectophorethe the hydroeciumhydroeciumis narroweris narrowerand enclosesand enclosesthe buddingthe buddingzonezoneoftheofsiphosome.the siphosome.It doesIt does not reachnot reachto thetoapexthe apexof theofnectophorethe nectophoreas theassidesthe sidesuniteunitetogether,together,in a inbroada broad curve,curve,just belowjust belowthat pointthat point(Figs.(Figs.2e, 3C).2e, 2C,3C). 3C)In the. Inlargestthe largestspecimen,specimen,the depththe depthof of the hydroecium decreases rapidly towards the base of the N nectophore as a the hydroeciumthe hydroeciumdecreasesdecreasesrapidlyrapidlytowardstowardsthe basethe baseof theofNthe2 nectophoreN 2 nectophoreas a as a resultresultof a sharpof a sharpincurvingincurvingof itsoflateralits lateralflaps flaps(Fig. (Fig. 3C). 3C). In both types of nectophore (N.(NI and N 2)) the somatocyst lies in the midline, In both types of nectophore (N. and N 2) the2 somatocyst lies in the midline, aboveabovethe centralthe centralpart partof theofdorsalthe dorsalwall wallof theofhydroecium,the hydroecium,and endsand endsat aboutat about ~th the4Aith%thheightthe heightof theofnectophore.the nectophore.It is Itunbranched,is unbranched,withoutwithouttenninalterminalswellings,swellings,swellings , and andhas neitherhas neitheran ascendingan ascending(apically(apicallyinto intothe mesogloea)the mesogloea)nor anordescendinga descending (basally(basallybeyondbeyondthe pointthe pointof originof originof theofpedicularthe pedicularcanal)canal)branchbranch(Figs.(Figs.2B, D;2B, D; 3B, D).3B,3B, D). BULLETI N OF MARINE SCIEN CE, VOL. 41, NO.1, 1987 72 72 BULLETINBULLETINOF MARINEOF MARINESCIENCE.SCIENCE,VOL. VOL.41. NO.1.41, NO,1987I, 1987
A A
FigureFigure4. Stem 4. StemgroupsgroupsofCraseoaofCraseoalalhe/ica:/athetlalhelica:A.ica:TwoA. Twoattachedattachedstemstemgroupsgroupsviewedviewedventrally.ventrally.B. DorsalB. Dorsal viewviewofolderofolderbract.bract.C. Side C. Side(proximal)(proximal)viewviewofyoungofyoungstem stemgroup. group.D. Detail D. Detailofarrangementofarrangementofbractealofbracteal and gonophoraland gonophoralcanalscanalsin the instem the stemattachmentattachmentregion.regionregion.A. and. AA.. andB. fromB. fromJSL 998 JSLspecimen; 998 specimenspecimen;C. and; C.D. and D. fromfromfromJSL JSL996 996specimen.specimenspecimen.Scale. Scale= 0.1= em.0.1 ernIncm.Figure. In Figure3B the 3Bbracteal thethe bractealcanalscanalsare: d: areare:dorsal;: dd:: dorsaldorsal;v: ventral;; vv:: ventralventral;; !.h. andl.h.I.h. andr.h.: r.hr.h.:left.:andleftleft right and righthydroecial; hhydroecial;ydroecialand ; 1.1.andand1.11.1.. r.!.:andleft r.l.:r.1.:and left rightand rightlongitudinal.longitudinal.longitudinal.
TheThenectosacnectosacis relativelyis relativelysmall,smallsmall,extending, extendingto abouttoto aboutoneonequarterquarterthe thetheheightheightof of the thethenectophores.nectophores.TheTheostialostialopeningopeningis basal,isis basalbasal,but, butthe thetheventralventralpartpartof theofthethenectosacnectosac is moreisis moreextensiveextensivethanthanthanthe thedorsalthe dorsalpartpart(Fig.(Fig.IA).lA).1A).TheTheradialradialcanalscanalsarisearisetogethertogethertogether fromfromthe thethepedicularpedicularcanal.canal.TheThedorsaldorsalandandventralventralcanalscanalsusuallyusuallyrunrunstraightstraightto toto the thethecircularcircularcanalcanalaroundaroundthe thetheostium,ostium,butbutin oneinin onespecimenspecimen(JSL(JSL959) 959)the thedorsalthe dorsal canalcanalof bothof bothnectophoresnectophoresis slightlyisis slightlyS-shapedS-shaped(Fig.(Fig(Fig.3).. The3)3).. Thelaterallateralradialradialcanalscanals in allininnectophores all nectophoresare arebroadlybroadlyS-shapedS-shaped(Figs.(Figs.2, 3). 2, 3). TheThewholewholesurfacesurfaceof theof thethenectophorenectophoreis coveredisis coveredby minuteby minutepapillae.papillae. Siphosome.Siphosome.TheThemaximummaximumnumbernumberof stemof stemgroupsgroupscaughtcaughtin associationinin associatassociationionwithwith the thethenectophoresnectophoreswaswasfour. four.It isItveryisis verylikely likelylikelythatthatthatothersotherswerewerelost lostduringduringcollectioncollection (see(seeIntroduction),Introduction)Introduction),as ,mostas mostrelatedrelatedspeciesspeciesusuallyusuallyhavehavebetweenbetween30 and30 and100100stemstem
-, "" " , .....,..... '.' ". "",. " . -'.~., .«...... •~;-...-t-..+.. :--".... PUGH AANDND HARBISONHARBISON:: NEW SPSPECIESECIES OF PRAVINEPRAYINE SIPHONOPHORE 73 PUGH AND HARBISON: NEW SPECIES OF PRAVINE SIPHONOPHORE 73
Figure 5. Craseoa lathetica: A. Gonophore-lateral viewview.. Scale = 00.1.1 emem.. BB.. Tentillum. Scale = 00.1.1 Figure 5. Craseoa lathetica: A. Gonophore-lateral view. Scale = 0.1 em. B. Tentillum. Scale = 0.1 mm. Both from JSL 998 specimen. rom.mm.BothBothfromfromJSLJSL998998specimen.specimen. groups.groups.AsAsall allthe theexistingexistingpartspartswerewereattachedattachedtogethertogethertogetherononcollection,collection,itititis isisclearclear thatthatthatthesethesethesestemstemgroupsgroupsrepresentrepresentthethetheyoungest,youngest,mostmostrecentlyrecentlydevelopeddevelopedones,ones,asas theytheytheyareareattachedattachedclosestclosesttototothe thethesiphosomalsiphosomalbuddingbuddingzone.zone.In Inlivinglivinglivingspecimensspecimensthethethe onlyonlycolorationcolorationDotednotedwaswasthe thedeep-reddeep-redpigmentationpigmentationofofthethethegastrozooids.gastrozooids.TheThe tentillatentillatentillawerewerewhitewhiteandandopaque,opaque,whilewhilethe theotherotherstemstemcomponentscomponentswereweretransparent.transparent.transparent. BractsBracts(Figs.(Figs.IB, IB,IB,4). 4).TheTheyoungyoungbractsbractsarearerounded,rounded,rounded,hemisphericalhemisphericalstructuresstructuresstructures measuringmeasuring5-65-6mmmminininheightheightandandaboutabout5 5mmmminininwidth.width.TheyTheyarearedivideddividedintointointo twotwotwoalmostalmostequalequallobeslobeslobesbybya adeepdeepincisionincisionincisionwhichwhichliesliesliesin ininthe thetheplaneplaneof,of,andandthroughthroughthrough C). whichwhichwhichpasses,passes,thethethesiphosomalsiphosomalstemstemstem(Fig.(Fig.(Fig.4A, 4A,C).C).TheTheincisionincisionincisionononthe theproximalproximal sidesideside(that(that(thatclosestclosesttototothe thethesiphosomalsiphosomalbuddingbuddingzone)zone)zone)isisisconsiderablyconsiderablydeeperdeeperthanthanthan thatthatthatononthethethedistaldistalside,side,andandextendsextendsalmostalmosttototothethetheapexapexofofthethethebract.bract.However,However,thethethe heightheightofofthethethedistaldistalsidesidesideofofthethethebractbractisisisgreatergreaterthanthanthanthethetheproximalproximalside,side,side,andandconcon sequentlysequentlysequentlythethethestemstemstemrunsrunsrunsobliquelyobliquelydownwardsdownwardsacrossacrossthethethehydroecialhydroecialcavitycavity(Figs.(Figs.(Figs. IA,lA,lA,4A).4A).In Inolderolderbractsbractsthethetheincisionincisionincisionbetweenbetweenthethethetwo twotwolobes lobeslobesis isisless lesslesspronounced,pronounced, andandthethethelobes lobeslobesare areflattenedflattenedflatteneddorso-ventrallydorso-ventrally(Fig.(Fig.(Fig.4B). 4B).TheThewholewholeouteroutersurfacesurfacesurfaceofof a abracta bractisisiscoveredcoveredininina aaregularregularregularpatternpatternofofsmallsmallsmallroundedroundedroundedpapillae,papillae,papillae,whichwhicharearelargerlargerlarger andandmoremoreabundantabundantthanthanthanthosethosethoseononthe thenectophores.nectophores. TheThedorsaldorsalbractealbractealcanalcanalarisesarisesfromfromfromthe thethedistaldistalendendofofthethetherightrightrightlongitudinallongitudinallongitudinal [NOTE: canalcanal(Fig.(Fig.(Fig.4B). 4B).[NOTE:[NOTE:TheTheterminologyterminologyterminologyforforforthethethearrangementarrangementarrangementofofthethethebractealbractealcanalscanals usedusedinininthisthisthispaper,paper,asasin ininFigureFigure4B,4B,is isbasedbasedononthatthatthatusedusedbybyBigelow BigelowandandSearsSears (1937)(1937)(1937)and,and,forforforinstance,instanceinstance,Carre, Carre(1969a;(1969a;(1969a;I969b)].1969b)]1969b)].The. Thedorsal,dorsal,ventralventralventralandandhydroecialhydroecial canalscanalsoftenoftenendendinininsmallsmallswellingssmalt"swellings,swellings, and, andinininoneonecasecase(Fig. (Fig.4A) 4A)4A)the thetherightrightrighthydroecialhydroecial canalcanalcanalwaswasforked.forked.forked.TheTheleftleftlefthydroecialhydroecialcanalcanalisisisslightlyslightlyslightlylongerlongerlongerthanthanthanthethetheright. right.right.TheThe 74 BULLBULLETINETIN OF MARINE SSCIENCE.CIENCE , VOLVOL.. 4141,, NONO.. II,, 1987 74 BULLETIN OF MARINE SCIENCE. VOL 41, NO, I, 1981 longitudinal canals are well developed, and run up the sides of the bract to end longitudinallongitudinalcanalscanalsarearewellwelldeveloped,developed,andandrunrunupupthethesidessidesofofthethebractbracttotoendend at approximately the same height as the lateral incisions. The thin dorsal canal atatapproximatelyapproximatelythethesamesameheightheightasasthethelaterallateralincisions.incisions.TheThethinthindorsaldorsalcanalcanal extends obliquely up to the dorsal surface ofthe bract and ends in a small depres extendsextendsobliquelyobliquelyupuptotothethedorsaldorsalsurfacesurfaceofofthethebractbractandandendsendsinina asmallsmalldepresdepres sion, slightly distal to the midline. Narrow diverticulae ofthe main stem underlie sion,sion,slightlyslightlydistaldistaltotothethemidline.midline.NarrowNarrowdiverticulaediverticulaeofofthethemainmainstemstemunderlieunderlie the proximal parts ofthe ventral and hydroecial canals (Fig. 4D). The main stem, thetheproximalproximalpartspartsofofthetheventralventralandandhydroecialhydroecialcanalscanals(Fig.(Fig.4D).4D).TheThemainmainstem,stem, in the region where the bract and other stem group components attach, appears ininthetheregionregionwherewherethethebractbractandandotherotherstemstemgroupgroupcomponentscomponentsattach,attach,appearsappears considerably expanded, but this may be an illusion caused by the extreme con considerablyconsiderablyexpanded,expanded,butbutthisthismaymaybebeananillusionillusioncausedcausedbybythetheextremeextremeconcon traction of the broken parts on either side. tractiontractionofofthethebrokenbrokenpartspartsononeithereitherside.side. Gonophore (Figs. lB, 5A). The gonophore is asymmetric in shape, being flat GonophoreGonophore(Figs.(Figs.1B,1B,5A).SA).TheThegonophoregonophoreisisasymmetricasymmetricininshape,shape,beingbeingflatflat tened, with wing-like expansions. The sub-umbrella cavity is restricted to the tened,tened,withwithwing-likewing-likeexpansions.expansions.TheThesub-umbrellasub-umbrellacavitycavityisisrestrictedrestrictedtotothethe lower half of the gonophore (Fig. IB) and is much less extensive than in other lowerlowerhalfhalfofofthethegonophoregonophore(Fig.(Fig.IB)IB)andandisismuchmuchlesslessextensiveextensivethanthanininotherother prayine species. Four straight radial canals are present in its lining. The pedicular prayineprayinespecies.species.FourFourstraightstraightradialradialcanalscanalsarearepresentpresentininitsitslining.lining.TheThepedicularpedicular canal is relatively long and at its apex gives rise to two mantle canals, which canalcanalisisrelativelyrelativelylonglongandandatatitsitsapexapexgivesgivesriserisetototwotwomantlemantlecanals,canals,whichwhich underlie the region of attachment of the gonophore to the stem (Fig. 4D). These underlieunderliethetheregionregionofofattachmentattachmentofofthethegonophoregonophoretotothethestemstem(Fig.(Fig.4D).4D).TheseThese canals are arranged asymmetrically, with one being slightly longer than the other. canalscanalsarearearrangedarrangedasymmetrically,asymmetrically,withwithoneonebeingbeingslightlyslightlylongerlongerthanthanthetheother.other. None of the gonophores were sexually mature. . NoneNoneofofthethegonophoresgonophoreswereweresexuallysexuallymature.mature. . . Gastrozooid and Tentacle. The gastrozooids have a deep-red coloration in GastrozooidGastrozooidandandTentacle.Tentacle.TheThegaslrozooidsgastrozooidshavehavea adeep-reddeep-redcolorationcolorationinin their basigaster and proboscis or buccal regions, while the stomach region is less theirtheirbasigasterbasigasterandandproboscisproboscisororbuccalbuccalregions,regions,whilewhilethethestomachstomachregionregionisislessless heavily pigmented. The long tentacle bears numerous side branches or tentilla heavilyheavilypigmented.pigmented.TheThelonglongtentacletentaclebearsbearsnumerousnumeroussidesidebranchesbranchesorortentillatentilla (Fig. 5B), which are similarto that ofrelated prayine species (Carre, 1969a; 1969b). (Fig.(Fig.5B),5B),whichwhicharearesimilarsimilartotothatthatofofrelatedrelatedprayineprayinespeciesspecies(Carre,(Carre,1969a;1969a;1969b).1969b). The cnidoband of the tentillum is kidney-shaped, and a single, highly coiled TheThecnidobandcnidobandofofthethetentillumtentillumisiskidney-shaped,kidney-shaped,andanda asingle,single,highlyhighlycoiledcoiled terminal filament stretches out from its distal end. tenninalterminalfilamentfilamentstretchesstretchesoutoutfromfromitsitsdistaldistalend.end. The arrangement of the nematocysts on the tentillum has not been examined TheThearrangementarrangementofofthethenematocystsnematocystsononthethetentillumtentillumhashasnotnotbeenbeenexaminedexamined in detail. However, in the cnidoband there are two rows ofup to 20 or 30 elongated inindetail.detail.However,However,ininthethecnidobandcnidobandtheretherearearetwotworowsrowsofofupuptoto2020oror3030elongatedelongated nematocysts measuring about 105 x 10 ~m. These are probably microbasic mas nematocysts measuring about 105 x 10 ILm. These are probably microbasic mas tigophores as these nematocysts are generally found in related species (Biggs et tigophorestigophoresasasthesethesenematocystsnematocystsarearegenerallygenerallyfoundfoundininrelatedrelatedspeciesspecies(Biggs(Biggsetet al., 1978). Other types ofnematocysts were scarcein the cnidoband, but included aI.,aI.,1978).1978).OtherOthertypestypesofnematocystsofnematocystswerewerescarcescarcein thethecnidoband,cnidoband,butbutincludedincluded a few (?)anisorhizas,(?)anisorhizas, measuring 30 x 5 ~m,~m , and desmonemes (9 ~m in diameter).diameter). a few (?)anisorhizas, measuring 30 x 5 ILm, and desmonemes (9 ILm in diameter). Desmonemes and rhopalonemes (18.5 x 5 ~m) were found in the terminal fila Desmonemes and rhopalonemes (18.5 x 5 ILm) were found in the tenninal fila ment, but it was not possible to determine whether they were arranged in the ment,ment,butbutititwaswasnotnotpossiblepossibletotodetenninedeterminewhetherwhethertheytheywerewerearrangedarrangedininthethe usual prayine pattern (Biggs et al., 1978), due to the highly contracted state ofthe usualusualprayineprayinepatternpattern(Biggs(BiggsetetaI.,aI.,1978),1978),dueduetotothethehighlyhighlycontractedcontractedslatestateofofthethe filaments. filaments.filaments. Distribution. -Known only from the region of the Bahamas, at depths between Distribution.Distribution.-Known- KnownonlyonlyfromfromthetheregionregionofofthetheBahamas,Bahamas,atatdepthsdepthsbetweenbetween 607 and 715 m. 607607andand715715m.m. Etymology. - The name lathetica is derived from the Greek Aa!/JT/nKw(J,'AacPl1TtKWI1 , meaning Etymology. - The name lathetica is derived from the Greek Aa¢TJTLKWU, meaning liable to escape notice. liableliabletotoescapeescapenotice.notice. Remarks. - The nectophores of C. lathetica, by their general shape, can be con Remarks.Remarks.- -TheThenectophoresnectophoresofofC.C.lathetica.lathetica.bybytheirtheirgeneralgeneralshape,shape,cancanbebeconcon fused only with species of the genera Rosacea and Desmophyes. However, they fusedfusedonlyonlywithwithspeciesspeciesofofthethegeneragenera Rosaceaandand Desmophyes.However,However,theythey can be distinguished easily from thoseRosaceaofR. plicata, R.Desmophyes.cymbiformis, and R.jlaccida cancanbebedistinguisheddistinguishedeasilyeasilyfromfromthosethoseofofR.R.plicata,p/icata.R.R.cymbijormis.cymbiformis.andandR.jlaccidaR.jlaccida by the absence ofa descending branch to the somatocyst and the S-shaped, rather bybythetheabsenceabsenceofofa adescendingdescendingbranchbranchtotothethesomatocystsomatocystandandthetheS-shaped,S-shaped,ratherrather than W-shaped, course of the lateral radial canals on the nectosac. Although the thanthanWW-shaped,-shaped,coursecourseofofthethelaterallateralradialradialcanalscanalsononthethenectosac.nectosac.AlthoughAlthoughthethe nectophores of R. villafrancae and D. annectens also do not have descending nectophoresnectophoresofof R. vi//afrancaeandand D. annectensalsoalsododonotnothavehavedescendingdescending branches to theirR. somatocystsvillafrancae, theyD.do annectenshave ascending branches, penetrating dor branchesbranchestototheirtheirsomatocysts,somatocysts,theytheydodohavehaveascendingascendingbranches,branches,penetratingpenetratingdordor sally into the mesogloea, and their lateral radial canals are straight (Table 1). sallysallyintointothethemesogloea,mesogloea,andandtheirtheirlaterallateralradialradialcanalscanalsarearestraightstraight(Table(TableI).1). In the bracts of C. lathetica the origin of the dorsal canal, from the distal end InInthethebractsbractsofofC.C. latheticathetheoriginoriginofofthethedorsaldorsalcanal,canal,fromfromthethedistaldistalendend of the right longitudinallatheticaone, differs markedly from most other prayine species ofofthetherightrightlongitudinallongitudinalone,one,differsdiffersmarkedlymarkedlyfromfrommostmostotherotherprayineprayinespeciesspecies except R. jlaccida, Mistoprayina fragosa (described below), and Praya species. exceptexcept R. jlaccida. Mistoprayina fragosa(described(describedbelow),below),andand Prayaspecies.species. However,R. jlaccida.as discussedMistoprayinalater, it isfragosanot certain whether this characterPrayais of more However,However,asasdiscusseddiscussedlater,later,ititisisnotnotcertaincertainwhetherwhetherthisthischaractercharacterisisofofmoremore than specific importance. This may apply also to the number of mantle canals in thanthanspecificspecificimportance.importance.ThisThismaymayapplyapplyalsoalsototothethenumbernumberofofmantlemantlecanalscanalsinin the gonophore, although such a character clearly distinguishes Praya species, thethegonophore,gonophore,althoughalthoughsuchsucha acharactercharacterclearlyclearlydistinguishesdistinguishesPrayaPrayaspecies,species, PUGH AND HARBISONHARBISON:: NEW SSPECIESPECIES OfOF PRAYINEPRA YINE SIPHONOPHORESJPHONOPHORE 75
which are the only ones to have three such canals.canals. Nonetheless, the slightly asymmetrical arrangement of these canals and the relatively small size of the sub-umbrellasub~umbrella cavity of the gonophore in C. lathetica are distinctive featuresfeatures..
Mistoprayina new genus Diagnosis. - Prayine siphonophores with an apposed pair ofconoid nectophores. The nectosac is extensive occupying the basal two-thirds of the nectophorenectophore,, and hasa wide dorso-basaldorso~basal openingopening.. In one nectophore (NI)(N(N,)I) the lateral radial canals are straight,straight, while in the other (N2) they areslightly curved. A descending branch to the somatocyst is present in both nectophores, while a simple ascending branch, penetrating dorsally into the mesogloea, occurs only in the N 2 nectophore. The young bracts are saddle-shapedsaddle-shaped,, but with age become flattened, with the stem attachment region raised on a mesogloeal process. Six bracteal canals are present. The rounded gonophores possess avery extensive sub-umbrella cavity. The two mantle canals are of different lengths, the longer being distinctly recurved. No special, asexual nectophores are present. Type Species. - MM.. fragosa new species described herein. Etymology. - The name Mistoprayina is used to suggest that the specimens show morphological characters that area mixture ofthose found in other prayine genera. Remarks. - The general shape of the nectophores in Mistoprayina fragosafragosa,, the sole representative ofthis newgenus, in that they are conoid and possess extensive nectosacs, suggests a close affinity with three other prayine genera, namely Prayola, Lilyopsis, and Stephanophyes. However, all four genera easily can be distinguished when consideration is given to the arrangement ofthe somatocyst and the lateral radial canals on the nectosac (Table 1)1).. In addition,addition, the stem groups possess features which, in combination, are ofsuch taxonomic importance as to separate easily the other genera and to necessitate the establishment ofaofa new genus, Misto prayina, to include the species described herein. A more detailed discussion of these points is given at the end of the description of M. fragosafragosa..
Mistoprayina fragosa new species Figures 6-8 MaterialMateria! ExaminedExamined..- Two specimens collected in the vicinity of the Bahamas by the submersible JOHNSON-SEA-LINK nII during 1984. Ho!otype.-SpecimenHolotype.-SpecimenHolotype.-Specimen collected using a detritus sampler during JSL Dive 990 at a depth of 416 m, where the water temperature was16.2°C l6.2°C (29 October 1984; 25°22.5'N, 77°54.5'W).77°54.5'W) . Preserved in 5% fonnalin.formalin. Presented to the British Museum (Natural HHistory)istory) and entered as RegdRegd.. No. 1986-6-1-21986-6-1-2.. ParatParatype.ype.-JSL Dive 993 specimen.specimen. (30 October 1984; 25°24.0'N, 25°24.0'N, 78"00.8'W;78000.8'W; Detritus sampler;sampler; Depth ofcollection: ca. 305 m. m. Housed in the biological collections ofthe Institute ofOceanographic Sciences, Wonnley.WormleyWormley.. . Diagnosis. -A monotypic genus and species ofprayine siphonophore whose prin ..... cipal characters have been defined in the diagnosis of the genus. In the bracts of , this species the dorsal canal arises fromthe distal end of the right longitudinal canalcanal,, and the hydroecial canals have small side branches. Description. - The surfaces of the nectophores, bracts and gonophores are very tacky, both inlife and after preservation. This stickiness makes itdifficult to examine the specimens without damaging themthem.. Nectophores (Fig. 6). Both specimens possess two dissimilar nectophores. They are coniformconiform with rounded apices,apices, and the nectosacs occupy more than half 76 BULLETINBULLETIN OF MARINE SCIENCE.SCIENCE, VOL.VOL. 41,41.41, NO.NO . II,I., 1987
Figure6. Nectophores ofMistoprayinafragosaMistoprayina fragosa:: A. Reconstruction ofofN,N I nectophore fromJSL 990 specimen.specimen. B.B. Ventro-lateral view of N,N 2 nectophore (JSL 993 specimen).specimen). C. Ventral and D. Ventro lateralviews of N,N2 nectophore (JSL 990 specimen).specimen). Scale = 0.10.1 em. their height.height. The mesogloea in the basal half of the nectophores is very thin, so that after preservation the nectophores became distorted and shrivelled.shrivelled. One of these nectophores,nectophores, which in its preserved slatestate measures about 3.3 mm in height by 2.8 mm in width,width, has been designated the N I1 nectophore. Its simple somatocyst is situated within a ridge ridge ofmesogloea that projects ventrally into the very shallow hydroecium. This somatocystsomatocysthas a short descending branch, but no ascending branch.branch. A short pedicular canal runs dorsally from the somatocyst to the ventral wall ofthe nectosac, where the four radial canals arise directly. The courses ofall of these canals are straight, such that they travel directly to the circular canal around the very large ostium, which opens dorso-basally.dorsa-basally. The narrow hydroecium extends the entire length of the ventral surface.surface. ApicallyApically its lateral margins are produced into two small flapsflaps.. PUGH AND HARBISON:HARBISON: NEWNEWSPECIES SPECI ES OF PRAPRAYINE VINEYINE SIPHONOPHORE 77
A c
B
o ~"" ~ .., , ...... ~/\ . , . " ~:.': ....
Figure 7. 7. Bracts of Mistoprayina!ragosa:Mistoprayinajragosa:Mistoprayinafragosa: A. Lateraland B. Ventral views of largerlarger bract (JSL 993 specimen). C. Lateral, D.D. Ventral and E. E. Dorsal views of younger bracts (JSL(lSL 990 specimen). Scale = 0.5 mm.
Unfortunately, no photographs or drawings of the living specimen were made and, since it would be meaningless to illustrate the shrivelled, preserved necto phore, a reasoned reconstruction is presented here(Fig. 6A). This reconstruction is based on the general features ofthe other, better preserved nectophore and the obvious fact that the nectophore possesses an extensive nectosac. It must be emphasized, however, that all of the characters oftaxonomic importance, which have been described above, can clearly be seen on the preserved specimens. The N 2 nectophore, of both specimens, is in a better state of preservation, probably because the thicker mesogloea in the upper half of the nectophore has prevented excessive shrinkage. The N 2 nectophore from JSL Dive 990 (Fig. 6B, C) measures 4.0 mm in height and 3.5 mm in width, while the JSL 993 nectophore (Fig.(Fig, 60) 6D)is smaller (ca.(ca. 2.8 mm x 2.5 mm respectively). As with the N II necto phore, the nectosac of the N 2 one is extensive,extensive, and is surrounded bya very thin ~.... layer ofofmesogloea.mesogloea.mesogloea. The hydroecium is virtually non-existent as almost the entire ventral surface of the nectophore forms a flattened plate. In the upper half of the >::..# ....;:... nectophore there is a central mesogloeal ridge projecting from the ventral surface within which runs the somatocystsomatocyst(Fig. 6B). This somatocyst has both a short descending branch and an apical ascending branchbranch,, which penetrates dorsallydorsally into 78 BULLETIN OF MARINE MARI NE SCIENCE.SCIENCE, VOL.VOL. 41.41, NO.NO . I,I, 1987 the mesogloea. The descending branch is only vaguely defined,defined, probably having become damaged when the two nectophores became separated. The relatively short pedicular canal runs tothe ventral surface of the nectosac and there gives riseto the four radial canals. The course of the lateral radial canals is slightly curved such that they reach the ostial circular canal towards thebase of the nectophore. Any additional meandering in these canals (Fig. 6B) 613) probably is caused by shrinkage. Siphosome. Judging by the number ofgastrozooids present, it appears that the JSL990 specimen possessed over 30 stem groups,groups, while 15IS remained with the JSL 993 one. No pigmentation of any parts was noted. Bracts (Fig.(Fig. 7). The younger bracts are minute (0.6--0.8(0.6-0.8 mm in length and ca. 0.60.6 mm in height), height) , rounded and roughly hemispherical in shape. At this early early stage, they are deeply divided,divided, in the axial plane, into two almost equal lobes (Fig. 7C). As they enlarge they become flattened (Fig. 7D, E) and eventually take on the form shown in Figure 7A,7A, B,13, with a deep hydroecial cavity dividing the hydroeciallobe. At this later stage they measure about 1.75 mm in length by0.6 mm in height. The region of stem attachment is enclosed within the hydroecial cavity in the younger bracts, but in the older ones it lies on a central mesogloeal process protruding below the main body of the bract (cf. the ventral mesogloeal ridge on the nectophores).nectophores), Six bracteal canals are present.present. The thin dorsal canal arises from the distal end ofthe right longitudinal canal, and extends to the dorsal surface ofthe bract where it ends at the base ofa distinct depression, particularly noticeable in the younger bracts (Fig. 7C). The hydroecial canals are of approx imately equal length.length. They bend away from the midline and asthey pass over the sides of the central mesogloeal process they give rise to small side branches (Fig. 7B).713). Gonophore (Fig. 8A). The almost circular gonophore bears no wing-like ex pansions. It is almost entirely occupied by a sub-umbrella cavity,cavity, which hasfour straight radial canals in itslining. The short pedicular canal gives rise, apically, to two unequal mantle canals, which underlie the stem attachment region. One ofthese canals, which is about twice the length ofofthethe other,other, is distinctly recurved. No mature gonophores were found,found, but asexual, special nectophores were not present. Gastrozooid and Tentacle (Fig.(Fig. 8B,8B813,, q.C). The small gastrozooids have a very characteristic shape (Fig. 8B).8-13).BB). The main stomach regionregion is spherical and opaque,opaque, measuring about 0.6 mm in diameter. Atits apex, apex , the mouth is borne on a slight prominence. Proximally, the gastrozooid is reduced to a narrow tube that is sheathed by the basigaster region,region , from which arises the tentacle. No pedicel could be discerned, although its apparent absence may bean artifact resulting from extreme contraction. Most ofthe tentacles had been lost and very few tentillatentilla could be found.found. How ever, those still present were of the typical prayine form (Fig. 8q,8C), 8C), with long nematocysts (?microbasic mastigophores) present in the proximal half of the cnidoband. No detailed studies have 'been made on the types of nematocyst present. Distribution. -Known- Known only from the Bahamas region at depths of305of305 and 416 m. m. Etymology. - The namefragosa refers tothe extremely delicate and fragile nature of the specimens.specimens. Remarks. -In- In general appearance the nectophores of MistoprayinaMistoprayina fragosa rere semble those found in the species of the prayine genera Prayola.Prayola, Lilyopsis and Stephanophyes. However,However, as noted above,above, all four genera can be distinguished PUGH AND HARBISON: NEW SPECIESSPEqES OF PRA VINE SIPHONOPHORE 79
B
Figure 8. Stem components ofMistoprayinajragosaMistoprayinafragosa (1SL(JSL 990 specimen):specimen): A. Gonophores. B. Gastro zooids. C. Tentillum. Scale = O.S0.5 mm. easily when consideration is given tothe arrangement of the somatocyst and the lateral radial canals on the nectosac (Table 1). For instance, with regard to the somatocyst, in Stephanophyes both ascending and descending branches are pres ent, while in Lilyopsis only the ascending one is to be found. Prayola species possess neither branch, whereas in Mistoprayina both branches ofofthethe somatocyst are found in the N 2 nectophore,nectophore, but only the descending branch is present ininthe N lone.lone. There are certain similarities in the arrangement ofthe lateral radial canals in L. rosea and M.M .fragosa inin that in the N II nectophore they are straight, whereas in the N 2 one they are curved, although to a much greater extent in the former species. However, in L. rosea.rosea,rosea, these canals arise from the dorsal radial canal,canal, at the apex ofthe nectosac,nectosac, while in M. fragosa. fragosa, in common with most other prayine species, all the canals arise together at the junction with the pedicular canal.canal. Carre (1(l969b)(1969b)969b) showed that the N II nectophore of L. rosea is the larval one, retainedretained in 80 BULLETIN OF MARINE SSCIENCE.CIENCE , VOL. 4141., NONO.. II., 1987 the adult stage for a variable length of time. However, this is probably not the case for the N 1 nectophore of M. Jragosafragosa as it possesses a descending branch to the somatocyst and such a feature has not been noted in the larval nectophores of other prayine species. The stem groups ofMistoprayinaMistoprayinafragosafragosa also can be distinguished easily from those of the other genera mentioned above. The bracteal canals of LLilyopsisilyopsis and Stephanophyes are arranged asymmetrically and special, asexual nectophores are present (Table 1).I). The absence ofthe dorsal canal in the bracts ofPrayola species distinguishes them from all other prayinesprayines..
Genus Prayola CarreCarre,, 1969 DiagnosisDiagnosis..- Prayine siphonophores with an apposed pair ofconoid nectophores, whose extensive nectosacs (>halfthe height ofthe nectophore) open dorso-basally. The radial canals on the nectosac are slightly curved, suggesting an open S. The somatocyst possesses neither an ascending nor a descending branch branch.. The bracts have only five bracteal canals, there being no dorsal one. The gonophores possess a hydroecial gutter and two mantle canals of equal length. No special, asexual nectophores are present. Type Species.-P.Species. -P. tottoni Carre, 1969. Remarks. - The diagnosis given by Carre (l969c)(I969c) for his new genus Prayola ob viously was based on the morphological characters of his new species, P. tottoni. ItIt iiss necessary to amend this diagnosisdiagnosis slightly in order toto incorporate the second speciesspecies,, P. urinatrix,urinatrix. described herein. NeverthelessNevertheless,, the characters of greatest taxonomic significance remain the samesame.. These are the conoid shape ofthe necto phoresphores;; the absence ofboth ascending and descending branches to the somatocystsomatocyst;; the open S-shape of the lateral radial canals on the nectosac; the absence of a dorsal canal in the bracts; and the absence of special asexual nectophores. The other characters that Carre (1969c)(I969c) mentionedmentioned,; the shortness of the somatocyst and ofthethe bracteal canalscanals,, are considered here to be only of specific importance.importance.
Prayo/aPrayola urinatrix new species Figures 9-12 MateMaterialrial Examined. -- Two specspecimensimens collected in the Bahamas region by the submersible JOHNSON-SEA-LINKJOHNSON-SEA-LrNK II during 1984. HolotHolotype.Holotype.-Specimenype.-Specimen-Specimen collected by a detritus sampler at a depth of 431 m during JSL Dive 1007. (4 NoNovembervember 19841984;; 25°22.425°22.4'N.25°22.4'N,'N , 77°53.5'W)77°53.5'W).. Preserved in 5% formalin.formalin. Presented to the British Museum (Natural History) and entered as RegdRegd.. NoNo.. 1986-6-1-3. ParatParalype.Paratype.ype.-Specimen collected during JSL DDiveive 999 (l(I NoNovembervember 19841984;; 25°22.925"22.9'N,25"22.9'N.'N, 77°5577°55.I'W;77°55.1.1'W'W;; De tritustritus samplersampler;; Depth of collection: 626 m). Housed in the biological collections of ththee InstituteInstitute of OcOceanographiceanographic SciencSciences,Sciences.es, WormleWormley.y. Diagnosis. -As forfor the diagnosis of the genus Prayola given above. In additionaddition,, thethe somatocyst ofthethe nectophore isis well-developed and situated on a mesogloeal process in thethe midline of the flattenedflattened ventral facet.facet. A gutter-like indentationindentation is present at the apex of thethe nectophore, thethe dorsal wall of which isis extended intointo a distinct collar or frill. In the young bracts the ventral canal is approximately twicetwice as long as thethe hydroecialshydroecials.. The mantle canals on the gonophore are longer thanthan those inin the related speciesspecies,, P. tottoni.tottoni. The mature femalefemale gonophores contain threethree eggs. PUGPUGHH AANDND HARBISONHARBISON:: NEW SPSPECIESECIES OF PRAYIPRAVINEYINENE SIPHOSIPHONOPHORENOPHORE 81
. \ I.
Figure 9. Nectophores of PraPrayolayola urinatrixurinalrix (JSL(1SL 1007 specimen)specimen):: AA.. Lateral and B. Ventral vviewsiews ofNofN,, nectophore. C. Ventral and DD.. Ventro-lateralVentra-lateralVentro-Iateral views ofofN,N, nectophoreneetophore.nectophore.. Scale = 0.1 em. 82 BULLETIN OF MARINE SCIENCE.SCIENCE, VOL.VOL. 41,4141., NO.NO . I,I, 1987
DescriptionDescription..- NECTOPHORESNECfOPHORES (Fig. 9). 9). Nectophores were found only on the JSL
1007 specimen. The two nectophores, N,N. and N 2 , differdiffer slightly in construction but,but, like the previous species, have a general shape reminiscent ofPrayola tottoni and Lilyopsis rosea. The larger (N1)I) nectophore measures about 4.1 mm in height and 22.6.6 mm in width (Fig. 9A, 9A, B).Its apex is not smoothly rounded,rounded, as in most other prayine siphonophores,siphonophores, but is interrupted by a gutter-like indentation. The dorsal edge of this gutter forms a distinct collar or frill around the apex of the nectophore and extends fora short distance down its sides before petering out (Fig. 9A). HoweverHowever,, the lateral and ventral facets ofofthethe nectophore arestill clearly demarcated by a rounded extension ofthe ventral edge ofthis gutter. A thickened bolster of mesogloea, in the shape of a horseshoehorseshoe,, formsfOnTIS the lateral and apical perimeter ofthe ventral facet (Fig. 9B). A central mesogloeal process alsois present in the apical part of the ventral facetfacet,, within which runs the somatocyst. The somatocyst is simple, without eithereither ascending or descending branches, but is considerably longer than that found in P. tottoni. Towards its apex, the somatocyst ofofthethe N 1 nectophore bends to the right (left when viewed ventrally, Fig.9B) away from the midline, and ends in a very small swelling.
The nectosac is extensive, occupying all but the apical quarter of the N 1 nec tophoretophore,, and has a large ostium that opens dorso-basallydorso-basally.. The short pedicular canal runs to the ventral surface ofthe nectosac and immediately gives rise to the four radial canals (Fig. 9A)9A).. The course of both the dorsal and ventral canals is straight, the ventral one joinsjoins the ostial circular canal basally, while the dorsal one passes over the apex of the nectosac to join the circular canal at the same level as its point of origin. The lateral canals are slightly curvedcurved,, running apico dorsally away from the pedicular canal before curving round to run down the sides of the nectosac and joining the circular canal close to the base of the neenec tophore (Fig(Fig.. 9A)99A).A).. The N 2 nectophore is smaller,smaller, measuring 3.3 mm in height and 2.1 mm in width in its preserved and clearly shrunken condition (Fig(Fig.. 9C,D). D). The nectosac is more extensive than that of the N 1I nectophore, and occupies so much of the nectophore that thickenings of the mesogloea occur only apically and ventrally. The apical margin of the nectophore is interrupted by a gutter, which is less pronounced than that in theN 1I nectophore,nectophore, as isthe dorsal frill or collar (Fig. 9C).9C). The dorsal wall of the gutter appears to extend down the sides ofthe necto phorephore,, but this feature is very indistinct. The ventral wall ofthe gutter also formsfOnTIS the lateral margins ofthe ventral facet. HoweverHowever,, unlike the N.N( nectophore, these margins do not reach to the base of the nectophore, but are united by a linkage running across the ventral facet at about a quarter the height of the nectophore (Fig.9C, 9C, D).D). Again, there is a horseshoe-shaped bolster of mesogloea that formsfOnTIS the lateral and apical perimeter of the ventral facet. The somatocyst lies on a central mesogloeal process.process. It is simple,simple, without ascending or descending branches, and does not curve away from the midline, as in the N 1I nectophore. The short pedicular canal runs to the ventral wall of the nectosac. It does not immediately give rise to allfour radial canalscanals,, but only to the vventalental and dorsal ones. The lateral canals branch from the dorsal canal a short distance above this point. The courses of all these canals are similar to those in the N,N. nectophorenectophore,, the laterals being slightly curved and joiningjoining the circular canal close to the base of the nectosac. Siphosome. About 14 stem groups were collected with the incomplete JSL 999 specspecimen,imen, while at least 24 gastrozooids were found with the more complete JSL 1007 one, one , whwhichich also possessed sexually mature gonophoresgonophores.. Bracts (Fig. 10). All but oneone·.of the bracts were young,young, and most were found PPUGHUGH AND HARBISONHARBISON:: NEW SPECIES OOFF PRAYINPRAPRAYINEVINEE SIPHONOPHORESJPHONOPHORE 83
A
IJ B ,I \ , '- '- . ,,
I I IJ / I w /-...... , I I I I I I t ,..------..... I ~~~ --- -- - /J , -,-"~--_---t .... , ,...,o!'/'. .... ,..~ ,/
'--, n.. ... '
• j .' FigureFig\lre 1010.. I}raCI$Bracts of PF(lyo/aPraPr associated with theJSL 999 specimen.specimen. Th~s~Thes~These bracts have a characte.ristjccharacteri!!tjccharacteristic saddlesa,ddle shaped appearanceo:lppearance (Fig. lOA) and measure aboutabo\lt 1.4 mm in length an4and 1I rommm in height.heigllt. A deep incision,incision, in the axial plan.e,plane, divides the bract into two almostalrnost equal lobes. The incisioll.incision is deepest on the lateral margins of the brao.tbract while centrilly,centrally,centrally, in the stem attachment region,region, there is a mesogloeal swelling. In addition,addition, the hydroecial cavity is relatively deep and $ubdivi<:iessubdivic;l.essubdivides the bl ' ,' Figure 11.II. Gonophores of Prayola urinatrix: urinatrix : A.A. Young gonophore-dorsalgonophore-dorsal and lateral vviewsiews (JSL 999 specimen). BB.. MatureMature malemale gonophore (JSL 1007 specimen). C. DeDevelopingveloping andand maturemature femalefemale gonophoresgonophores (JSL 1007specimen). specimen). ScaleScale = 0.1O. I emem.. Large gonophore bells were scarce ininthe more developeddeveloped JSL 1007specimen, specimen, but were common in the other,other, although none of the latter contained well-dewell-de velopedvelopedsexual sexual products.products. It is presumed presumed that largelarge bells are developed developed only once on each stem group and that they are lost in associationassociation with ththee release of theirtheir sexual products. Nevertheless, well-developed male and female gonophores wwereere found in the JSL 1007 specimen,specimen, but the medusoidmedusoid bell associated with these was reduced to a small cup-like structure around their bases (Fig.(Fig. 11B11 B,, C).C), The mature female gonophores contained only three eggs. Each stem group possessed gono phores of one sex only, only, but both sexessexes were present on the stem.stem. No special,special, asexualasexual nectophores werewere found.found. PUGH AND HARBISONHARBISON:: NNEWEW SPSPECIESECIES OF PRA YINEVINE SIPHONOPHORE 85 B A c D FFigureigure 1212.. Gastrozooids and tentillumtentillum of Prayola urinatrixurinatrix:: A. Young gastrozooid,gastrozooid, with B. The pattern of cells on its stomach regregionion (JSL(JSL 999 specimen).specimen). C. Older gastrozooid, and D. Tentillum fromfrom JSLJSL 1007 specimen. ScaleScale:: A and C = 0.5 mmmm,, BandBand D = 0.1 mmmm.. Gastrozooid and Tentacle (Fig. 12). The gastrozooids of thethe JSL 999 spec imenimen were relatively small (Fig(Fig.. l2A)12A) and possessed a characteristic pattern of cells on thethe surface of theirtheir stomach regionsregions (Fig. l2B).12B). HoweverHowever,, thethe largerlarger gastrozooids of thethe other specimen did not show thisthis pattern (Fig. l2C).12C). The younger tentillatentillatentilla were of thethe typicaltypical praprayineyine designdesign,, with a J-shapedJ-shaped cnidobandcnidoband,, except thatthat no large nematocysts could be foundfound on thethe cnidoband. InIn thethe more developed tentillatentilla (Fig. l2D)12D) thethe cnidoband had straightened somewhatsomewhat,, with thethe pedicel insertedinserted basally. A few large nematocysts were presentpresent inin thethe proximal part of thethe cnidobandcnidoband,, (?)microbasic mastigophores, but no detailed studiesstudies of thesethese have been mademade.. Distribution. - Known only fromfrom thethe Bahamas regionregion at depths of431 and 626 m. • f ~ 86 BULLETIN OF MARINE SCSCIENCE.SCIENCE,IENCE, VOLVOL.. 4141.41,, NONO.. Il.I,, 1987 Etymology. - The name urinatrurinatrix.ix, meaning a female (sea) diverdiver,, is used in rec ognition of the invaluable role that the RR.V..V. SEA DIVER, the tender ship for the JOHNSON-SEA-LINK]OHNSON-SEA-LINK II, and her crew played during the OctoberOctober-November-November 1984 cruise. RemarksRemarks..- The species ofthe genus Prayola are immediately distinguishable from all other prayine siphonophores by the fact that a dorsal canal is absent in their bracts (Table 1). In addition, the conoid shape ofthe nectophores, together with thethe absence of both an ascending and descending branch to the somatocyst are distinctive characters. The only other prayine to show the latter character is Craseoa lathetica,lathetica. but the general features 'of the nectophores and bracts set this species apartapart,, as discussed above. The characters that distinguish Prayola urinatrix from P. tottoni are thatthat in the formerformer:: a) the apex of the nectophore isis·interrupted by a gutter andand·therethere is a characteristic mesogloeal frill, particularly in the N II nectophore; b) the nectosac is more extensive although the course of the lateral radial canals in both species is very similar; c) the somatocyst isis relatively long, whereas inPinP.. tottoni it is very short; and d) the lateral radial canals of the N 2 nectophorearisefromnectophore arise from the dorsal canal slightly above the point ofinsertion ofthe pedicular canal, whereas in both nectophores of P. tottoni all four canals arise together. The general structure ofthe stem groups is very similar in both Prayola species. The young bracts are deeply incisedincised laterally and only five bracteal canals are present. The ventral and hydroecial canalsinPcanalsinP.. tottoni are ofalmost equal lengthlength,, while in P. urinatriurinatrixx the ventral is almost twicetwice the length ofthe hydroecials. The young gonophores also are very similarsimilar,, although the mantle canals in PiurinatrixP. urinatrix are longer than thosethose in P. tottonitottoni.. Carre (1969c) makes no mention of reduced gonophores in P. tottonitottoni,tottoni., but he did note that from four toto six eggs were present inin the female gonophores ofhis speciesspecies,, whereas we found only three eggs in those of P. urinatrurinatrix.ix. DISCUSSION Biggs et al. (1978) briefly discussed how thethe nomenclature of the sub-family Prayinae had become very confused and complicated iinn the pastpast,, particularly particularly..with regard to thethe genus Rosacea and one of its speciesspecies,, R. plicata. This species was desdescribedcribed by Quoy and Gaimard (1827)(1827),, but for thethe remainderremainderofthethe 19th century no other specimens were described under thatthat name, although the taxonomytaxonomy is very confused. HoweverHowever,, some authors have considered thatthat other species, par ticularlyticularly Desmophyes annectens Haeckel, 1888, might be the same as thatthat ofofQuoyQuoy and GaimardGaimard.. This may be true but itit isis now impossibleimpossible toto verify as the original specimen of R. plicata is no longer inin existence (Carre, pers. commcomm.)..). Bigelow (1911 a) appears to have been thethe firstfirst toto re-usere-use thethe name RR.. plicata whilst de scribing some material collected inin thethe Bay ofBiscay. Later thatthat year, he (Bigelow(Bigelow,, 1911b)1911 b) figured other specimens fromfrom thethe Eastern Pacific, although he ddidid appear to have some reservations about theirtheir exact identity.identity. Nonetheless, it is apparentapparent,, as detadetailediled by Totton (1965)(1965),, thatthat thethe spspecimensecimens described by BBigelowigelow as RR.. plicata are not thethe same as that described originally by Quoy and Gaimard (1827). Totton decided thatthat since the generic name Rosacea sensu Bigelow had been inin common usage for over half a century itit should be retainedretained andand,, hence, he designated R. plicata sensu Bigelow as its typetype species. We follow thisthis decision and in order toto stabilize thethe nomenclature we designate twotwo nectophores (N II and N 2)2),, fromfrom thethe material thatthat BBigelowigelow (1911(1911a)(1911 a) examinedexamined,, as the neotypeneotype.. These nectophores are housed inin thethe British Museum (Natural(Natural History)History),, and are entered "'. .'.". Table I. A comparison of the morphological characters of related prayine species .~ .s £ £ " '" §o ~~ jl'! -c::'6'~ s ", .~ . ~ "" ~ ~s ~~ ~~ ~ .g 6-~~ ~~ .s., ~.g '-E ",,,, ~~ ~ ~ !1..!! ~] ~t ~ ~ E S"''5.!! .~ ~ l~ ""t ",,,, a.g SE ~~ ",2 ~ '§ h ~ .. ~ .5 ~~ ~ <1:'" ~.g, ~~ Q::-5 t'E ll('il, as Regd. NoNo.. 1939.6.10.1. They were found in aajarjar (Regd. No. 1939.61939.6.10.1-5).10.1-5) which contained three NN,I and three N 2 nectophores of Rosacea plicata sensu Bigelow, plusone larval nectophore ofthe same species. Ofcourse, since none of the nectophores are attached to each otherother,, it is impossible tobe certain that the two nectophores removed belonged originally to the same specimen. However,However, on the basis of size, the NtN N,I and N 2 nectophores can be paired off,off, and the best preserved of these pairs has been selected as the neotype. The specimens were collected during the cruise of H.M.S.H.M.S. RESEARCH. The labelon the jar states that the depth ofcollection was 300-0fm which, judging by the records for R. plicata given by Bigelow (19lla)(191Ia) (1911 a) would indicate that they came from haul 36k. This haul was made on 25-vii-190025-vu-1900 at 47°03'N, 47°03'N, 7°55'W7°55'W.. However, Bigelow (19lla)(191Ia)(1911a) stated that six superior (NI) and two inferior (N(N2)2) nectophores were found in this haul, which is not in accord with the extant material. Perhaps the specimens came from haul 361 (350-0fm), in which Bigelow (191la)(1911a)(191Ia) found three superior and three inferior nectophores; but the museum records are insufficient to resolve this point. Bigelow's (1911(1911a)a) material ofRosacea plicata is ofgreat interest as it is apparent that two of the N 1I nectophores possess a very short apical extension of the so matocyst into a pad of mesogloea that projects into the hydroecium. It was this feature that convinced Bigelow that his material could be identified with that described by Quoy and Gaimard (1827) as R. plicata. NeverthelessNevertheless,, this very short extension in no way resembles the pronounced ascending branch of the somatocyst found in other species, e.g., e.g., Desmophyes annectens and R. villafrancae.villafrancae, and we do not consider itto represent such a structure. HoweverHowever,, it does bear a close resemblance to that found in theN N.1I nectophore ofanother Rosacea species, R.jlaccidaR .jlaccida(Table 1). We have examined other material ofR.R . plicata sensu Bigelow,Bigelow, from recent R.R.S.R.R.S. DISCOVERY collections made in the Bay ofBBiscayiscay (pugh, 1984), and find that this short apical extension ofthe somatocyst is present in some, but not all,all, of the N.N I1 nectophores. Thus,Thus, we confirm that this feature is a true, but inconsistent, morphological characteristic ofthis species. We have examined also many specimens ofR. cymbijormis.cymbiformis, collected by SCUBA divers, and can find no trace of such an apical extension to the somatocyst in any of the nectophores. In recent years four new species ofprayine siphonophores have been described. Two of these were placed in the genus Rosacea,Rosacea, namely R. villafrancae Carre, 1969 and R.R . jlaccida Biggs et al.,a1., al., 1978, whilethe third, PrayolaPrayola toltonitottoni Carre,Carre, 1969, formed the type species of a new genus. The fourth species,species , Nectocarmen antonioi Alvariiio,Aivariiio,Alvarifio, 1983,1983, we regard as an incomplete specimen of Prayadubia. The two minute nectophores,neetophores, which AivariiioAlvariiioAlvarifio (1983) found on her specimen, are quite out of proportion with the rest of the material and would appear to be replacements, at very early stages of development,development, for the pair of full sized nec nee totophoresphores that have become detached. In his discussion ofPP.. toltoni,tottoni, Carre (1969c) produced keyskeys forthe identification of the nectophores and stem groups of all extant prayine genera. However,However, with the present description of three new species, it is apparent that the generic char acters used by Carre (1969c) are no longer wholly satisfactory and that other characters,characters, as detailed in Table 1,I, are also oftaxonomic significance. We consider ..... the following characters to be of particular importance. For the nectophores: a) the general structure of the bell; bell; b) the structure of the somatocyst,somatocyst, including the presence or absence of ascending and/or descending branches;branches; and c) the general structure ofthe nectosac and the courses ofits canals. For the stem groups: a) the general structure of the bract and gonophore; b) the number and arrangement of the bracteal canals; and c) the number and arrangement of the mantle canals on the gonophore. Individually, some of these characters are only ofspecific impor- PUGH AND HARBISONHARBISON:: NEW SPESPECIESCIES OF PRAVINEPRAVINEYINE SIPHONOPHORE 89 A ;~,:::: ~;~>o..,: _1/ .s \,·1 n n E FigureFigure \13.'3. AA.. Bract (dorsal view) ofofDesmophyesDesmophyesanneanannectensnee/ensctens (JSL(JSL 969969 specimen),specimen),with B thevariabilityvariability inin thethe sizesize ofofthethe central vesiclevesicle asas illustratedillustrated byby thethe patternpattern ofofcanalscanals inin otherother bractsbracts fromfrom thethe samesame specimen.specimen.(In(In thesetheseventralventralviewsviews thethedorsaldorsalcanalcanal isis obscured.)obscured.)C.C. Gonophore ofofDesmophyesDesmophyesannectensanneetensannectens (JSL(JSL(ISL 969969 specimen).specimen). DD.. Gonophore (lateral(lateral view)view)andand EE.. Bract (ventral(ventral view)view)ofofRosacea cymbiformiscymbiformis.cymbijormis.. FF.. Bract and GG.. Gonophore (lateral(lateral vviews)iews) ofofRosaRosaceacea plicataplica/a.plicata.. ScaleScale == 00.2.2 ernern.cm.. 90 BULLETIN OF MARINE SCIENCESCIENCE.SCIENCE,. VOVOL.L. 4141,41.. NNO.O. I.I.I, 1987 tance butbut,, when in combination with othersothers,, they can be used to establish separate genera. The presence ofan asexualasexual,, special nectophore inin each cormidiumcormidium,, as in the genera Lilyopsis and Stephanophyes,Stephanophyes. also is an importantimportant generic character. Although such a structure has been described in the stem groups of Desmophyes annectens we believe that it does not exist in this species, as is shown below. ItItisis apparent from Table 1I that the prayine genera can be split into twotwo subgroups on the basis of thethe general shape of thethe nectophore. There are those genera that have cylindrical nectophores with nectosacs occupying less thanthan half the heightheight,, and others which have conoid nectophores with extensive nectosacs.nectasacs. The presence or absence of a descending or ascending branch toto thethe somatocystsomatocyst thenthen separates thethe individual genera in each subgrouping. Thus: Cylindrical Conoid Both branches present: Praya Stephanophytes Ascending branch only: Desmophyes Lilyopsis Descending branch only: Rosacea Neither branch present: Craseoa Prayola Both present inin N 22,, but only one inin NNt:I: Prayoides Mistroprayina As mentioned above, we do not consider thethe very small apical extension ofthethe somatocyst inin thethe NNtII nectophores of Rosacea plicataplicata and R. flaccidaflaccida toto be rep resentativeresentative of a truetrue ascending branch, which runs dorsally intointo thethe mesogloeamesogloea,, and so the genus Rosacea remainsremains distinct fromfrom all thethe other prayine genera. Nevertheless, therethere isis one species, RR.. villafrancae,villafrancae. thatthatdoes not fitfit intointothisthisgeneral scheme as its nectophores do not have a descending branch toto thethe somatocyst. InIn factfact,, by the presence of an ascending branch toto thethe somatocyst and ofstraight radial canals on the nectosac, thisthis species appears toto be more closely allied toto Desmophyes annectens,annectens. although therethere are certain differences in thethe stem groupsgroups,, e.g., in the arrangement oftheofthe bracteal canalscanals.. InIn D. annectens thethe dorsal bracteal canaljoinscanaljoinsthethelongitudinallongitudinalcanal centrally, by way ofa variably sizedsizedbasal swellingswelling (Fig.(Fig. 13A13A,l3A,, '8)'8),B),, while inR. villafrancae thisthiscanal originates fromfrom therightrighthydroecial (see(see Carre, 1969a)1969a).. However, itit should be noted thatthat inin twotwo other species, R. plicata and R. cymbiformis,cymbiformis. which are obviously closely relatedrelated toto each other, therethere isis a comparable situation. InIn R. cymbiformis thethe dorsal bracteal canal arises fromfrom thethe rightright longitudinallongitudinal (Fig.(Fig. 13E),l3E), proximal (central) toto thethe rightright hydroecial, although not inin thethe midline, while in R.R. plicataplicata itit arises fromfrom thethe rightright hydroecialhydroecial (Fig.(Fig. 13F).l3F). Thus itit would seem thatthat thethe arrangementarrangement ofthethe bracteal canals isis only ofspecific, not genericgeneric,, importance.importance. Nonetheless thethe bracts ofthethe twotwo latterlatterspeciesspecies areare very similarsimilar inin basic construction, asas areare theirtheir gonophores (Fig(Fig.. 13D,l3D, G). We conclude thatthat thethe general similaritiessimilarities between thethe nectophores and bracts of DD.. annectens and R. villafrancae suggestsuggest aa closeclose affinityaffinity between thesethese twotwo species andand accordingly we transfertransfer thethe latterlatter intointo thethe genus Desmophyes asas D. villafrancaevillafrancae (Carre,(Carre, 1969). However, we are aware thatthat thisthis arrangementarrangement isis not wholly satisfactory since therethere areare certaincertain differences between themthem,, e.g.,e.g., thethe number of mantlemantle canalscanals inin thethe gonophore. Another obstacle toto thisthis transferencetransference couldcould bbee thethe factfact thatthatDD.. villafrancae does not possesspossess special, asexualasexual nectophores on the siphosomesiphosome while theythey have been described forfor D. annectensannectens.annectens,. The presencepresence of specialspecial nectophores isis certainlycertainly aa valid generic character, butbut we believe thatthat DD.. annannectensectens does not possess suchsuch structuresstructures.. We havehave examinedexamined severalseveral specspec imensimens of D.D,D. annectens caughtcaught by SCUBA divers and byby thethe submersible PUGH AND HARBISONHARBISON:: NEW SPECIESOF PRA PRAYINEVINE SIPHONOPHORE 91 ]OHNSON-SEA-LINKJOHNSON-SEA-LINK II and have been unable to find any sign ofspecial nectophores. Large bells are present in the stem groups, but all ofthese possess either developing sexual products or a small manubrial rudiment. Thus they are all true gonophores (Fig.13C). It does not seem possible that an even larger asexual nectophore also could be attached to the stem and the capture of intact specimens confirms this. History has shown that the observation of a bell without a manubrial process does not necessarily mean that it is an asexual nectophore. Bigelow (1911a)(191Ia) in his description of Rosacea plicata sensu Bigelow, suggested that asexual necto phores were present. However, Bigelow and Sears (1937) noted, for the same species, that the spadix ofthe gonophores could be detached without leaving any trace and concluded that the supposed asexual nectophores were old gonophores that had remained attached to the stem. We believe the same situation applies to D. annectens and,and, thus, that asexual nectophores are notdeveloped. Nonetheless it is apparent that the gonophores ofD. annectens possess two symmetrical mantle canals (Fig. 13C), while those ofD. villa/rancaevillafrancae possess only one. Although Haeckel (1888) described D. annectens as having upto six nectophores, arranged biserially, none ofthethe specimens we have seen have had more than two apposed nectophores. ACKNOWLEDGMENTS Thecruisecruise during which the specimens were collected was funded by the NationalNational Science Foundation under Grant No. OCE-8400243. We thank all those involved in this cruise for their assistance.assistance. This paper represents DSMC (Direct Studies of Mesopelagic Communities) Contribution No. 10. We are grateful to Mrs. C. Darter fordrawing up the figures. Contribution No. 607 of the Harbor Branch Oceanographic Institution. LITERATURE CITED Alvariiio,Alva rifio, A. 1983. Nectocarmen antonioantonioi,i, a new Prayinae,Prayinae, Calycophorae,Calycophorae, Siphonophora fromCal ifornia. Proc. Proc. BioI.Soc. Wash. 96: 339-348. BBigelow,igelow, H.B. 1911a.191 la. Biscayan plankton collected during a cruise ofH.Mof H.M.S..S. 'Research,''Research,' 19001900.. Part XIIIXIII.. The Siphonophorae.Siphonophorae. Trans.Trans. Linn.Linn. SocSoc.. Lond.Lond . (Zool.) 1010:: 337-358337-358.. --. 1911b.191 lb. The Siphonophorae.Siphonophorae. Mem.Mem. MusMus.. CompoCompoZool. Harv.Harv. 38:38: 173-402173-402.. --and M. Sears. 1937. Siphonophorae.Siphonophorae. Rep.Rep. Dan.Dan. Oceanogr.Oceanogr. Exped. Mediterr. (II BioI.) BioI.) 2:2: 1- 144.144. Biggs,Biggs, DD.. c.C.,, PP.. RR.. Pugh and C. Carre. 19781978.. RosaceajlaccidaRosacea j/accida n.sp.,n.sp ., a new species ofofsiphonophoresiphonophore (Calycophorae Prayinae)Prayinae) from the North Atlantic OceanOcean.. Beaufortia 2727:: 207-218207-218.207-218.. Carre, C. 1969a. Rosaceavilla/rancae villafrancae sp.sp. n.un nouveau siphonophore calycophore Prayinae de la mer Mediterranee,Mediterranee. Beaufortia 16: 109-117109-117.109-1 17.. --. 1969b.1969b. Sur Ie genre LilyopsisLilyopsis Chun 18851885,, avec une redescription de l'espeee l'espece Lilyopsisrosea Chun 1885 (Siphonophore,(Siphonophore, Prayinae) et une description de sa phase calyconula. Cah. BioI. MarMar.. 1010:10:71-81.: 71-81. ----.. 1969c.1969<;. Prayola tot/oni tottonitol/oni gen.gen. sp. nn.,., nouveau genre et nouvelle espece de siphonophore caly cophore Prayinae de la mer MediterraneeMediterranee., Vie Milieu 2020:: 31-42. Haeckel,Haeckel , E.'EE..' 1888.1888. Report on the Siphonophorae collected by HH.M.S..M.S. Challenger during the years 1873-18761873-1876.. RepRep.. Sci. Res. HH.M.S..M.S. Challenger (Zool.) 28: 1-3801-380.. PughPugh,, PP.. RR.. 19841984.. The dieldieI migrations and distributions within a mesopelagic community in the Northeast Atlantic. 7.7. Siphonophores. Prog. Oceanog. 13: 461-489. ----andand G.G. RR.. HarbisonHarbison.. 1986. New observations on a rare physonect ssiphonophore,iphonophore, LLychnagalmaychnagalma utriutriculariacularia (Claus, 1879)1879).. J. Mar. BioI. Ass. Ass. U.K.66: 695-710. Quoy,Quoy, JJ.. R.R. C. and JJ.. P. GaimardGaimard.. 1827. ObservatObservationsions zoologiques faites a bord de "1'Astrolabe,""I'Astrolab"I'Astrolabe,"e,' en Mai 1826, 1826, dans Ie detroit de Gibraltar. Ann. Sci. Nat. 1010:: 55-21,172-193.5-21,-21, 172-193172-L93.. Totton, AA.. K. 1965. A synopsis of the Siphonophora.Siphonophora. British Museum (Natural History), LondonLondon.. 230 pp. pp. DATE ACCEPTEDACCEPTED:: October 30, 1986. ADDRESSES:AoDRFSSES: (P(P.R.P.).R.P.) Institute ofo/OceanographicOceanographic Sciences.SciencesSciences,, Wormley, Godalming.GodalmingGodalming,, Surrey,Surrey, GU8 5UB, 5UB, England;(G.R.H.) (G.R.H.) Harbor BranchOceanographic InstitutionInstitution,, 5600Old Dixie HighwayHighway,, Fort PiercePierce,Pierce., Florida 3345033450..