Mediterranean Sepiola aurantiaca Jatta, 1896, versus the NE Atlantic Sepiola pfefferi Grimpe, 1921 (Cephalopoda, Sepiolinae)

2012·0.92 MB·

by  Goud, Jeroen

B76-Goud:Basteria-2010 11/07/2012 19:19 Page 1
Mediterranean Sepiola aurantiaca Jatta, 1896,
versus the NE Atlantic Sepiola pfefferi Grimpe, 1921
(Cephalopoda, Sepiolinae)
Jeroen Goud & Ate de Heij
NCBNaturalis,P.O.Box9517,NL-2300RALeiden,TheNetherlands;[email protected]
phylogenyoftheSepiolinaeintheNorthSea(Groenenberg
OriginaldescriptionsandliteraturereferencesofSepiolaau- etal.,2009).Foroneofthespeciesweusedexplicitlythe
rantiacaandS.pfefferiarecomparedwith156NEAtlantic nameSepiolapfefferiGrimpe,1921,insteadofthemorecom-
specimensfrommuseumcollectionsandrecentcollecting monlyusednameSepiolaaurantiacaJatta,1896.Inthispaper 1
fromtheInternationalBottomTrawlSurvey(IBTS)andthe weanalyzetheuseofthesenamesintheliteraturethusfar
BeamTrawlSurvey(BTS).ItappearsthatSepiolapfefferi andwecomparebothdescriptionswith156additionalspec-
Grimpe,1921,isnotasynonymofS.aurantiacaJatta,1896. imensreportedherefromdifferentlocalitiesintheNorth
SeaandtheNEAtlantic.
Keywords:Cephalopoda,Sepiola,nomenclature,taxonomy,
Mediterranean,Atlantic. Abbreviations:BMN,UniversityMuseumofBergen,Norway;NMSZ,
RoyalScottishMuseum,Edinburgh,Scotland;RMNH,NCBNaturalis,
Leiden,theNetherlands;ZMA,ZoologischMuseumAmsterdam,now
Introduction inNCBNaturalis,Leiden.
IntheNationalMuseumofNaturalHistory,NCBNaturalis Sepiolaspecies occuring in the North Sea
(Leiden),wecouldstudysince2004,throughthekindcoop-
erationofImares,Wageningen(Dutchfisheriesresearchsta- IntheliteraturewehavefoundfourspeciesnamesofSepiola
tion,IJmuiden),manyoftheSepiolinaesamplescollected supposedlyoccurringintheNorthSea:S.atlantica
duringtheInternationalBottomTrawlSurvey(IBTS)and d’Orbigny,1839-1842,S.pfefferiGrimpe,1921,S.aurantiaca
theBeamTrawlSurvey(BTS),undertheauspicesoftheIn- Jatta,1896andS.rondeletiiLeach,1834.Byfarthecommon-
ternationalCouncilfortheExplorationoftheSea(ICES).By estnameisSepiolaatlantica.Asshowninourmostrecent
meansofaninitialDNAsurveyin2006-2007itwasshown paper(DeHeij&Goud,2010)S.atlanticaintheNorthSea
that,asexpected,severaloftheidentificationsprovedtobe consistsoftwocloselyrelatedspecies.Onegenerallyliving
wrong.ThisresultedinawiderDNAstudyandamolecular inmorecoastalshallowconditions,thetrueS.atlantica,and
Basteria76(1-3):1-11(2012)
B76-Goud:Basteria-2010 11/07/2012 19:19 Page 2
thenewlydescribedSepiolatridensDeHeij&Goud,2010,
livingmoreoffshoreatdepthsover50m.InDeHeij&
Baayen(2005)thedistributionofS.atlanticaintheNorthSea
wasdescribed.Infacttheirmaterialstudiedconsistedof
threeSepiolaspecies,mainlyofS.atlanticaandS.tridensbut
alsosomespecimensofathirdspecies,whichcanbere-
ferredtoasS.pfefferiGrimpe,1921orS.aurantiacaJatta,
1896.Bothnameshaveformerlybeenusedinliteratureon
cephalopodsfromtheNorthSea(Russell,1909:454-455,
1922:28-29;Grimpe,1925:19-21;Jaeckel,1958:577-578;
Nesis,1987:128,132).TheoccurrenceofS.rondeletiiinthe
NorthSeaisonlysuggestedbythedistributionmapsin
Roperetal.(1984:68)andinJereb&Roper(2005:167-168).
InthereferenceslistedbytheseFAOguidesnodatacon-
cerningtheoccurrenceofS.rondeletiiintheNorthSeacould
befoundandwealsodidn’tfindanyS.rondeletiiinthehun-
dredsofsampleswehavestudiedinthepasttenyears.The
distributionofS.auranticaseemstoberestrictedtotheBay
ofNaples.
Literature studies on usage of the names
2 S.aurantiacaandS.pfefferi
In1896JattadescribedthespeciesSepiolaaurantiacafromthe
GulfofNaplesintheMediterraneanSea(pp.130-133,plate
14:31-46).InFigs1a-bwereproducethemainfiguresgiven
byJatta,1896:plate14figs34,36,38,40&42. Fig.1.SepiolaaurantiacaafterJatta,1896:plate14.1a,ventralview(34),
SomeofthemaincharactersgivenbyJattaare:theskin dorsalview(38);1b,armcomposition(36),hectocotylus(40),tentacle
colouroftheanimals,“niceyelloworange”;theU-shaped club(42).
ventralmantlecurvingwidelyaroundthefunnel(Fig.1a:
34);theshapeofthetentacleclub(smoothwithequally Therightarmalsocarriesinthefirstthirdofitslengthtwo
smallsuckers)(Fig.1b:42);thehectocotylizationofthemale leaf-likeexpansions,whichare,however,muchsmallerthan
arms(Fig.1b:40). thoseoftheleftarm;theremainingpartofthearminun-
Thislastcharacter,themostimportantdiscriminating changed.”[translationfromItalianbyG.Bello].Jattade-
characterofmalecephalopods,isdescribedbyJattaasfol- scribedandfiguredclearlytwoleaf-likeexpansionsaspart
lows:“Hectocotylizationinvolvesallthearms(pl.14fig.36). ofthehectotylizationinbothfirstmalearms.
Thearmsofthefirstpairarejoinedtoeachotherattheir Russell(1909)usedJatta’snameSepiolaaurantiacafor20
baseforashortextent;theleftarmcarriesonitsventralpart specimenscaughtNEofScotland.Hebasedthisidentifica-
twoverylargeleaf-likeirregularlyshapedexpansions, tionontheskincolourandtheU-shapedventralmantle
placedclosetoeachother(pl.14figs36,40).Theremaining aroundthefunnel,buthealsorecognizedtheabsenceofthe
partofthearmisunchanged,butitssuckersarequitelarger leaf-likeexpansionsonthefirstrightmalearmasdescribed
andaremorewidelyspacedthanthoseonthefemalearms. byJatta.Inasubsequentpublication(Russell,1922)he
Basteria76(1-3)
B76-Goud:Basteria-2010 11/07/2012 19:19 Page 3
enlargedsuckersinbotharmsmuchbiggerthanthosein
Jatta’sdrawing(Fig.1b:40).Russellapparentlydidnotrec-
ognizethese20specimensasbelongingtoadifferentspecies
inspiteofthedemonstrateddifferences.
Naef(1912:271)studiedsevenspecimensofSepiolaau-
rantiacainthecollectionofthe‘StazioneZoologicadiNapoli’
andbrieflyredescribedthespeciesreferringtotheoriginal
descriptionofJatta(1896:130-133).Naef(1923:590-593[612-
615inoriginaledition])publishedanadditionaldrawingof
thehectocotylusofS.aurantiaca.Thetwoexpansionsatthe
baseoftherightarmaredrawnveryprominently(seeFig.
3).Hedescribedtheseexpansionsasextendedstalksof
suckerswith,inthecaseofyoungmales,thesuckerssome-
timesstillinplace.Inlaterpublicationsitisalwaysthis
drawingwhichisreproduced(e.g.Muus,1963,sheet94:2;
Nesis,1987:135N;Bello,1995:47)
In1921Grimpe(pp.4-12)describedthecatchofthree
specimensofSepiolaattheeastcoastofEngland,south-east
ofFlamboroughHead.Theseanimals,althoughsimilarto
3
Fig.2.SepiolaaurantiacaafterRussell,1922:figs8-11.8,dorsalside;9,
ventralside;10,hectocotylus;11.tentacleclub.
figuredtheventralanddorsalsideofafemalespecimen
(Fig.2:8,9),atentacleclub(Fig.2:11)andthemale’sfirst
armpair(Fig.2:10).Wecannowpointoutthreecharacters
beingdifferenttothoseofSepiolaaurantiaca.Thetentacle
showsaclubwithaclearseparationinamanusandadacty-
luspartwhereasJatta(1896)(Fig.1b:42)figuredaclub
withoutsucha‘digit’.Russellfiguresthefirstarmpairwith
onlytheleftarmbeinghectocotylizedwhereasJattagavea Fig.3.SepiolaaurantiacaafterNaef,
descriptionandafigureofbotharmsbeinghectocotylized. 1912:fig.1d,hectocotylus.
Russell’sdrawingofthefirstarmpairshowssomevery
Goud,J.&DeHeij,A.–Sepiolaaurantiacavs.S.pfefferi
B76-Goud:Basteria-2010 11/07/2012 19:19 Page 4
Fig.4.SepiolapfefferiafterGrimpe,1921:figs3,5.a,hectocotylus;
b,malearmswithcompletesuckercomposition
Jatta’sdescription,wereaccordingtoGrimpenotonlydiffer-
entinthehectocotylus,withonlythefirstleftarmhecto-
cotylized,buttheyalsohadamuchdeeperV-shapedventral Fig.5.SepiolapfefferiafterGrimpe,1921:figs1a,b;holotype.5a,ventral
mantleindentation,lyingmuchcloseraroundthefunnel side;5b,dorsalside.
(Fig.5),whereasJatta(1896)describedawiderU-shaped
ventralmantleindentation(Fig.1a:34).Grimpe(1921:11) imens:1sp.Roscoff(France);4sp.Bergen(Norway);1sp.
alsoindicatesadifferenceinthenumberofsuckerrowson FirthofForth(Scotland)and3sp.NorthSea,butstillcon-
4 thetentacleclubs.Hesaw6clearrowsofsuckersopposite sideredtheclearlydescribeddifferencesasofminorimpor-
toJatta,whosedrawing(Fig.1b:42)shows8rowsofsmall tanceandjustenoughtousethenameS.pfefferifora
suckers.Grimpe(1921:4-12)describedthesethreespeci- subspeciesofS.aurantiaca.
mensasSepiolapfefferi. Inreturn,Grimpe(1925:21),alsoinafootnote,com-
Inhiswellknowncephalopodmonographfrom1923 mentsuponNaef’sopinionbystatingthatthedifferences
Naefoncemoredescribesandfiguresthehectocotylusof betweenanewlydescribedspeciesbyNaefanditsclosest
S.aurantiacaandcommentsuponS.pfefferi.Fromthefoot- relativeareevensmallerthanthedifferencesbetweenhis
noteonpage591(613inoriginaledition)welearnthatNaef S.pfefferiandS.aurantiaca.Asanexampleisgiventhediffer-
studiedthetypespecimensofS.pfefferiGrimpe,1921,and encebetweenSepiettaowenianaPfeffer,1908andSepiettane-
noticedthatinS.pfefferi“[translatedfromtheGerman]the glectaNaef,1916.ApartfromthebiggersizeS.oweniana
funnelindentationisstilldeeper,almostpointed.Thesuck- differs‘only’inhaving32rowsoftinysuckersonthetenta-
ersofthemalearemoreenlargedandthereareapparently cleclubagainst16rowsinS.neglectaandtheirmaturemales
nochangesonthebaseofthedorsalarm.Thetentacleclubs haveaslightlydifferenthectocotylus.(Bello,1998:84).We
arelargerandresemblethetypebecauseofthedistinctsix couldsaythesameaboutSepiolaatlanticad’Orbigny,1841
rowsandtheslightlylargersuckers.”Despitetheseclear andthecloselyrelatedSepiolatridensdeHeij&Goud,2010,
differencesNaefstillconsideredS.pfefferionlyavarietyof twospecieswithsmall,butconsistant,morphologicaldiffer-
S.aurantiaca.AccordingtoNaef,Grimpeneverstudiedthe encesmainlyinthesizeofthetentacleclubsandthemale
knownspecimensandtheirvariation,butsimplybrought hectocotylus.NaefsawthedifferencesbetweenS.aurantiaca
togetherthespecimensreportedasS.aurantiacafromthe andS.pfefferi,butdidnotagreethatthesecouldbeindica-
NorthSeaandtheChannelandrecognizedtheseas tivefortwodifferentspecies.
S.pfefferi.Naefindicatesthathehasseenthefollowingspec- ThemonographbyGrimpe(1925)onthecephalopodsofthe
Basteria76(1-3)
B76-Goud:Basteria-2010 11/07/2012 19:19 Page 5
S.aurantiaca.Lateron,thesamespecimenswerelistedby
Heppell&Smith(1983:15)andbyYau(1994)underthename
S.aurantiaca.Yau(pers.com.)indicatesdoingthataccording
toNesis(1987).FinallyHayward&Ryland(1990:789)arelist-
ingS.aurantiacaaccompaniedbytheoriginaldrawingofthe
ventralsideofGrimpe’sS.pfefferi.Whatstartedastwodiffer-
enttaxaaccordingtoGrimpe,orsubspeciesaccordingto
Naef,becamefinallysynonymswithoutthestudyofanyad-
ditionalspecimens.
Searching for the type samples
Fig.6.SepiolapfefferiafterGrimpe,1925:fig.5onphotoplate.Ventral Wealreadyanalyzedandlistedthedifferencesinthede-
sideofthreespecimens.Therightspecimencanberecognizedasthe scriptionsofS.aurantiacaandS.pfefferi,butwewouldhave
holotype(comparefig.5);notethedifferenceindepthofindentationin likedtocomparethetypeseriesaswell.Wehaverequested
theventralmantle. thetypesofS.aurantiacafromtheZoologicalStationAnton
Dohrn,Napels(SZN-StazioneZoologicadiNapoli),but
evenafterseveralsearchestheyhavenotbeenabletotrace
NorthSeashowsaphotoplatewiththreespecimensof thetypesinthecollection(pers.com.Dr.Andrea
S.pfefferi(Fig.6).Therightspecimenhasexactlythesame Travaglini);Dr.G.Belloinformedus,thathehasstudiedthe
shapeasthelinedrawinggivenbyGrimpein1921withthe S.aurantiacatypesamplesintheseventiesoflastcentury.
originaldescription(Fig.5)andindicatedastheholotype. Whythesesamplescannotbefoundnowadaysisstillamys- 5
Theindentationoftheventralmantlearoundthefunnelis tery.
V-shaped,butfarlessdeepasindicatedinthelinedrawing. SearchesforthetypesofS.pfefferiweremoreelaborate.
Thespecimenleftonthephoto(Fig.6)showsanevenmore Felleyetal.(2001-2003),indicatethatthetypesprobablyare
shallowU-shapedindentation. intheZMB(ZoologicalMuseumBerlin).DrMatthias
BasedonthiscontroversybetweenNaefandGrimpewe Glaubrechtinformedus,thatthesetypesarecurrentlynotin
seenowthatJaeckel(1958:577-578)andNesis(1987:128,132) theZMBcollection.Grimpematerialmightbeinthe
reportbothS.aurantiacaaswellasS.pfefferiaspartoftheNE NaturkundeMuseumLeipzig.KatrinSchniebsinformedus,
Atlanticcephalopodfauna.Nesisindicates(probablyafter thattherearenocephalopodsamplescollectedordonated
Naef)thatS.pfefferimightnotbeadistinctspecies,butaform byGrimpeintheircollection.Shesuggestedtocheckwith
ofS.aurantiaca.InfactNaefandGrimpewerereferringtothe theUniversityofLeipzigcollection.Thecollectionmanager
samespecimensunderdifferentnames.From1925ondiffer- oftheLeipzigUniversitycollectioninformedus,thatthere
entauthorshavebeenusingboththesetwonamesforspeci- arenoGrimpesamplesofSepiolaintheircollection.
mensoccurringintheNEAtlanticarea,whichobviouslydo Grimpepublishedhismonographonthecephalopodsof
belongtothesamespecies.Grieg(1933:11-12)usedS.auranti- theNorthSeaintheHelgoländerWissenschaftlicheMeere-
acaforspecimensfromthefjordsnearBergen,Norway, suntersuchungen.WehaveinquiredabouttheHelgoländer
whereasGrimpe(1925:19)listedthesamespecimensinhis collectionwithDr.Heinz-DieterFranke.Heinformedus,
monographonthecephalopodsoftheNorthSeaasS.pfefferi. thatthecollectionatHelgolandwaslargelydestroyeddur-
Stephen(1944:254-255)oftheRoyalScottishMuseum(Edin- ingWorldWarIIandthattherearenoSepiolasamplesfrom
burgh)calledtheScottishspecimensS.pfefferi,including Grimperemaining.TheinstituteatHelgolandnowadaysis
thosecollectedbyRussell(1909),whooriginallynamedthese partoftheUniversityofKiel.Dr.U.Piatkowskiinformed
Goud,J.&DeHeij,A.–Sepiolaaurantiacavs.S.pfefferi
B76-Goud:Basteria-2010 11/07/2012 19:19 Page 6
Fig.7.Sepiolapfefferi,ventralside
ofmalewithmantleedgeturned
over.RMNH.MOL.108846.1,
NorthSea,EofScotland,90m
depth;photoJ.Goud.
us,thattherearenoGrimpesamplesintoday’sUniversityof
6 Kielcollection.
Neotype selection
Fig.8.Sepiolapfefferi,ventralsideoffemalewithsharpV-shapedmantle
SinceallthreesyntypesofSepiolapfefferiGrimpe,1921must indentation.RMNH.MOL.117755,NorthSea,EofScotland,51mdepth;
beconsideredlost,weheredesignateaneotype.Wehavese- photoJ.Goud.
lectedafullygrownmale(RMNH.MOL.108845),whichhas
beencollectedon26.i.2005atadepthof38m,verycloseto
theoriginaltypelocality(53,11671°N1,7827°E)andofwhich specimenasS.aurantiaca.SearchesintheZoologicalRecords
aCO1sequencehasbeendepositedinGenbank:FJ231296. procuredonlyonerecentlypublishedspecimen.Thisspeci-
menwasusedinthemolecularstudiesofNishiguchietal.,
Material studied 1998andaCO1sequencewasdepositedinGenbank
(AF035708)andwasshowntobedifferentfromS.pfefferiby
Thestudiedsamplesarelistedinthetableonthenextpage Groenenbergetal.,2009.Thespecimenwasnotavailablefor
(abbreviations:f,female;m,male;sp.,specimen;y,young). morphologicalcomparison(pers.com.Nishiguchi).
SepiolaaurantiacaJatta,1896 SepiolapfefferiGrimpe,1921
FromtheMediterraneanSeanospecimensarefoundinmu- Differentnaturalhistorycollections:RMNH,Leiden;RSM,
seumcollections.BelloandBoletszky(pers.com.)bothlong- Edinburgh;BMN,Bergen,NorwayandZMA,Amsterdam
timeresearchersonMediterraneancephalopodsand arehousingatotalof156specimens:61males,93females
interestedinSepiolainparticularhaveneveridentifieda and2juvenilesofS.pfefferi.Dorsalmantlelength(DML)of
Basteria76(1-3)
B76-Goud:Basteria-2010 11/07/2012 19:19 Page 7
identification revisor regnr lotcontent area latitude longitude depthm date genbank
S.pfefferi Goud&DeHeij RMNH.MOL.110095 1m RiadeArosa 18 26.vii.1963
S.pfefferi Goud&DeHeij RMNH.MOL.110096 1f RiadeArosa 50-60 12.viii.1964
S.pfefferi Goud&DeHeij RMNH.MOL.113973 4f NorthSea 534,407 0,96167 20 26.i.2005
S.pfefferi Goud&DeHeij RMNH.MOL.113657 1m NorthSea 546,768 257,567 20 9.ii.2005
S.pfefferi Goud&DeHeij RMNH.MOL.110056 1f NorthSea 537,677 0,9181 15.ii.2006
S.pfefferi Goud&DeHeij RMNH.MOL.110302 5f NorthSea 572,972 -13,531 94 7.ii.2008
S.pfefferi Goud&DeHeij RMNH.MOL.110303 2m NorthSea 572,972 -13,531 94 7.ii.2008
S.pfefferi Goud&DeHeij RMNH.MOL.110306 21f NorthSea 566,668 -16,123 58 8.ii.2008
S.pfefferi Goud&DeHeij RMNH.MOL.110307 9m NorthSea 566,668 -16,123 58 8.ii.2008
S.pfefferi Goud&DeHeij RMNH.MOL.117755 2f NorthSea 567,617 -2,236 51 9.ii.2009
S.pfefferi Goud&DeHeij RMNH.MOL.117758 1f NorthSea 563,973 -21,002 54 9.ii.2009
S.pfefferi Goud&DeHeij RMNH.MOL.119755 1m NorthSea 577,492 -13,607 95 31.viii.2009
S.pfefferi Goud&DeHeij RMNH.MOL.113962 2f,2m NorthSea 578,345 -25,868 83 31.i.2009.
S.pfefferi Goud&DeHeij RMNH.MOL.113964 1m NorthSea 588,737 -38,178 100 1.ii.2009
S.pfefferi Goud&DeHeij RMNH.MOL.117319 1f NorthSea 538,107 0,82733 37 26.viii.2009
S.pfefferi Goud&DeHeij RMNH.MOL.117322 1f NorthSea 511,833 186,183 39 2.ix.2009
S.pfefferi Goud&DeHeij RMNH.MOL.113758 3f,4m EnglishChanal 503,398 -0,6965 52 21.i.2009
S.pfefferi Goud&DeHeij RMNH.MOL.113758.1 1f EnglishChanal 503,398 -0,6965 52 21.i.2009
S.pfefferi Goud&DeHeij RMNH.MOL.113943 1f,1f CelticSea 514,158 -76,084 86 27.ii.2009
S.pfefferi Goud&DeHeij RMNH.MOL.113944 1m CelticSea 513,418 -74,508 88 27.ii.2009
S.pfefferi Goud&DeHeij RMNH.MOL.113945 1f,1m CelticSea 51,144 -81,529 106 28.ii.2009
S.pfefferi Goud&DeHeij RMNH.MOL.113946 1f,3m CelticSea 513,523 -81,406 92 28.ii.2009
S.pfefferi Goud&DeHeij RMNH.MOL.121490 1m IrishSea 533,943 -54,109 86-89 4.iii.2009
S.pfefferi Goud&DeHeij RMNH.MOL.121491 1m IrishSea 533,332 -52,564 89-92 4.iii.2009 7
S.pfefferi Goud&DeHeij RMNH.MOL.121492 1m IrishSea 535,245 -5,589 69-71 5.iii.2009
S.pfefferi Goud&DeHeij RMNH.MOL.121493 1f IrishSea 541,771 -45,387 78-95 6.iii.2009
S.pfefferi Goud&DeHeij RMNH.MOL.121494 6m,5f IrishSea 533,118 -41,527 48-53 11.iii.2009
S.pfefferi Goud&DeHeij RMNH.MOL.108846.1 1m NorthSea 5,774,243 -13,547 90 26.viii.2005 FJ231292
S.pfefferi Goud&DeHeij RMNH.MOL.108846.2 1y NorthSea 5,774,243 -13,547 90 26.viii.2005 FJ231293
S.pfefferi Goud&DeHeij RMNH.MOL.105642 1f NorthSea 5,344,067 0,96167 20 26.i.2005 FJ231294
S.pfefferi Goud&DeHeij RMNH.MOL.110056 1f NorthSea 534,611 0,9181 38 15.ii.2006 FJ231295
S.pfefferi Goud&DeHeij RMNH.MOL.108845 1m NorthSea 5,311,671 17,827 38 26.i.2005 FJ231296
S.pfefferi Goud&DeHeij RMNH.MOL.110050 1f NorthSea 56,75 -0,5 68 6.ix.2000 +
S.pfefferi Goud&DeHeij RMNH.MOL.110055 1m NorthSea 56,75 -0,5 68 6.ix.2000 +
S.aurantiaca Yau,1991 NMSZ:1958045.78b 2m NorthSea 57°06'N 02°03’W 24-37 4.iv.1931
S.pfefferi Stephen,1944 NMSZ:1958045.89 1m,1f NorthSea 60°22'N 01°31’W 75 8.ix.1920
S.pfefferi Stephen,1944 NMSZ:1958045.90 1m NorthSea 56°05'N 02°32’W 0-40 26.iii.1925
S.pfefferi Stephen,1944 NMSZ:1958045.91 1m,1f NorthSea 58°09'N 03°27’W 45 14.xi.1926
S.pfefferi Stephen,1944 NMSZ:1958045.94 1m NorthSea 56°06'N 02°31.5W 41 20.xii.1928
S.pfefferi Stephen,1944 NMSZ:1958045.98a 1m,2f NorthSea 56°09'N 02°43’W 40 6.xi.1906
S.aurantiaca Yau,1991 NMSZ:1979038.41301b 1m NorthSea 57°42'N 02°30’W 42 6.xii.1978
S.aurantiaca RSM,1980 NMSZ:1980074.41301 1m NorthSea 49°28’N 02°27’W lowwater 24.ix.1980
S.aurantiaca McKay,1982 NMSZ:1982043.41319 5m,8f NorthSea - - - 20.xi.1982
S.aurantiaca Naef/Grieg,1933 BMN:36233 1m,2f Norway Bergens skjærgaard 17m
S.aurantiaca BMN:36258 1y EofShetlands 110m 15.vii.1904
S.aurantiaca Naef/Grieg,1933 BMN:86305 1f Norway Bergen
S.pfefferi ZMA.Moll.162435 1f Norway Stavanger 8ord v.1929
Goud,J.&DeHeij,A.–Sepiolaaurantiacavs.S.pfefferi
B76-Goud:Basteria-2010 11/07/2012 19:19 Page 8
8
Fig.9.Sepiolapfefferi,ventralsideoffemales(left)andmales(right).AllfromsampleNMZS1982043-41319,NorthSea,EofScotland;photoJ.Goud.
bothsexesrangesfrom10-15mm.Thegeographicdistribu- dactylusatthetoparetypical.Russell(1922),probablynot
tionrangesfromnorthernSpain(42°N)toeastoftheShet- awareofthedescriptionofS.pfefferibyGrimpein1921,fig-
lands(60°30’N)andfromtheCelticSea(8°15’W)tosouthern uredunderthenameS.aurantiacaspecimensfromtheNorth
Norway(5°E).Thedepthrangesfromlowwatertidemark Sea,offthecoastofScotland.Hisfigure11(Fig.2)showsa
to110m. tentacleclubwithaclearseparationbetweenmanusand
Results dactylusaswehaveseeninallourS.pfefferisamples.
Theindentationoftheventralmantlevariesmorethan
TheexactrelationshipbetweentheMediterraneanS.auranti- Grimpe(1921:fig.5)showedinhisfigures.Inparticularthe
acaandtheAtlanticS.pfeffericannotbesettledsincenotype linedrawingofthetypeshowsaverydeepindentation,al-
ofS.aurantiacacouldbetracednorcouldanyMediterranean thoughhis1925publication,onphotoplate(Fig.5)showsa
specimen,identifiedassuch,befound. photoofthesamespecimen(hereFig.6)withafarlessdeep
MorphologicalcomparisonshowedthatalltheAtlantic indentation.Inourspecimenstheindentationvariesfrom
specimens,whichwestudied,areconspecificwiththede- VtoU-shapedandfromdeepertomoreshallow(Fig.9).
scriptionandtypefiguresofS.pfefferiGrimpe,1921.Espe- Remarkableisthattheindentationinfemales(Fig.9left)is
ciallythetentacleclubs(Fig.10),characterizedbythebroad morepronouncedthanthatinmales(Fig.9right).
manuswithlargersuckersontheinsideandthesmall Theindentationoftheventralmantleisverycharacteris-
Basteria76(1-3)
B76-Goud:Basteria-2010 11/07/2012 19:19 Page 9
Fig.10.Sepiolapfefferi,tentacle
club,aftercriticalpointdryingand
coatingphotographedinSEM;
scalebar=1mm.RMNH.MOL.
117322;photoJ.Goud.
ticforthespecies,althoughnotalwayseasilyrecognizable.
ThefirstspecimenscollectedbyImaresintheNorthSea,
thatwesaw,werefrozenafterthecatchbeforeidentification. Fig.11.Sepiolapfefferi,malearms,toptwoarmsarethefirstarmpair,
AsshowninFigure7wecouldhardlyspeakaboutanin- withhectocotylusasleftarm(rightsidefortheviewer);photoJ.Goud.
dentation,itjustseemsthattherearetwowhiteknobson 9
theedgeoftheventralmantle.Nowweknowthatthisisan
artifactduetothefreezingtechnique.Specimensprepared Asaconsequenceofourobservationsitisclearthatthe
inethanol70%showanormallyrelaxedmantlewithclear namesS.aurantiacaJattaandS.pfefferiGrimpecannotbere-
indentation(Fig.8). gardedassynonyms.AllAtlanticspecimensthatwehave
Wegiveaphoto(Fig.11)andadrawing(Fig.12)oftwo seen,shouldbegiventhenameS.pfefferi.Manyoftheiden-
firstarmpairsfromfullyadultmalesofS.pfefferi.Theleft tificationguides(Naef,1923;Jaeckel,1958;Muus,1959,1963;
arm(rightonthefigures)ishectocotylizedandtherightarm Roperetal.,1984;Jereb&Roper,2005)refertothesespeci-
clearlynot.ThisisincontrastwiththesituationinS.auranti- mensaseitherS.aurantiacaorS.pfefferi.Someresearchpa-
aca(Fig.1b:40andFig.3)wherebothJattaandNaefdraw persandafewidentificationguideshowever(Grieg,1933;
twolargepapillaeatthebaseofthefirstrightarm.Asec- Hayward&Ryland,1990;Yau,1994;Collinsetal.,2002;Ha-
ondarysexdifferencecanbeseenintheverylargesuckers stieetal.,2009)calltheAtlanticspecieserroneouslyS.auran-
halfwaythefirst,thesecondandfourtharmpair.About3to tiaca.Weherelistthereferences,whichinouropinion
5couplesofsuckersaremuchlargerthantheothers.Inthe shouldhavebeenconsideredS.aurantiacaforMediterranean
drawingsopJatta(Fig.1b:36)andNaef(Fig.3)weseeonly specimensorS.pfefferiforAtlanticspecimens.
somelargersuckersaspartofthehectocotylusarm;allother
armshaveregularlysmallersuckers.ThefiguresofGrimpe SepiolaaurantiacaJatta,1896(Figs1-3)
(1921)(Fig.4a,b)illustratewellthesituationinourAtlantic SepiolaaurantiacaJatta,1896:130-134,figs31-46.Naef,1912:85,fig.1d;
samples(43males)whichisshowninFigs11and12. 1923:612-615,figs344a,357-358.Nesis,1987:128,132,fig.30D,N.Bello,
ThefirstarmpairfiguredbyRussell(1922)appearstobeju- 1995:47,50,figs9♂B,9♀B;1998:84,fig.3.Nishiguchi,Ruby&McFall-
venile(Fig.2:10)andnotwellpreserved. Ngai,1998:tab.1,fig.1.Jereb&Ropereds.,2005:180.
Goud,J.&DeHeij,A.–Sepiolaaurantiacavs.S.pfefferi
B76-Goud:Basteria-2010 11/07/2012 19:19 Page 10
SepiolapfefferiGrimpe,1921(Figs4-12)
Sepiolaaurantiaca;Russell,1909:454-455;1922:28-29,figs8-11.Grieg,
1933:11-12.Jaeckel,1958:577-578.Heppell&Smith,1983:15.Hay-
ward&Ryland,1990:788,fig.13.18.Yau,1994:4.Collins,Yau,
Boyle,&Piatkowski,2002:243,tabs2-5.Hastieetal.,2009:140-141.
NotJatta,1896.
SepiolapfefferiGrimpe,1921:4-12,figs1-5;1925:19-21,photo5.Stephen,
1944:254-255.Nesis,1987:128,figs30A-C.Jereb&Ropereds.,2005:
180.Chambers,2008:322.Groenenberg,Goud,DeHeij&Gittenber-
ger,2009:361-369,tabs1-2,figs1-4.
Sepiola(Hemisepiola)aurantiaca;Muus,1959:135,fig.74;1963:sheet94:4,
pl.IIfigs10a,b.
Sepiola(Hemisepiola)pfefferi;Muus,1959:136,fig.75;1963:sheet94:4,pl.
IIfig.11.
Acknowledgements
AcknowledgedfortheirhelpwithpreparationsareKeesvan
derBerg(Naturalis)andBertie-JoanvanHeuven(National
Herbarium).
WearehappytosaythatapartfromImares(Remment
10 terHofstede,IngeborgdeBooisandHenkHeessen,IJ-
muiden,Netherlands),fivemoreinstitutionssupportedus
bysendingvaluablespecimensanddata:IFREMER(Yves
Vérin,BoulognesurMer,JocelyneMartinandPascalLaffar-
gue,Nantes,France),FRSML(CraigDavisandRobert
Wartret,Aberdeen,Scotland),IMR-SE(BarbaraBland,Ly-
sekil,Sweden),MarineInst.Rinville(DavidStokesand
RobertBunn,Oranmore,Ireland)andAFBI(PeterMcCorris-
ton,Belfast,NorthernIreland).
Wealsothankthefollowingcuratorsandresearchersfor
theirhelpwithtracingthetypesandloansofmaterial:Royal
ScottishMuseum,Edinburgh,SankuryPye,ZoologicalCol-
lection;StazioneZoologicaAntonDohrn,Dr.Andrea
Travaglini;NationalMuseumofIreland,Dublin,NigelT.
MonaghanandDrJuliaSigwart;BiologischeAnstaltHel-
Fig.12.Sepiolapfefferi,male,firstarmpairwithhectocotylus;NMZS goland,Prof.Dr.Heinz-DieterFranke;Museumfür
1982043-41319;drawingBasBlankevoort,NCBNaturalis. Naturkunde,Berlin,DrMatthiasGlaubrecht;BergenMu-
seum,NaturalHistoryCollections,UniversityofBergen,Dr
ManuelA.E.Malaquias;StateCollectionsofNaturalHistory
Dresden,MuseumofZoology,DrKatrinSchniebs;Leibniz-
InstitutfürMeereswissenschaften,Kiel,Dr.U.Piatkowski.
Basteria76(1-3)