3 Biology and Ecology of Cephalopod Species Commercially Exploited in Europe

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3 Biology and ecology of cephalopod species commercially exploited in Europe

3.1 Species accounts

PatriziaȱJereb,ȱUweȱPiatkowski,ȱLouiseȱAllcock,ȱPaolaȱBelcari,ȱManuelȱGarciaȱ Tasende,ȱAngelȱGonzález,ȱAngelȱGuerra,ȱLeeȱC.ȱHastie,ȱEugeniaȱLefkaditou,ȱ AnaȱMoreno,ȱSantiagoȱPascual,ȱJoãoȱPereira,ȱGrahamȱJ.ȱPierce,ȱPilarȱSánchez,ȱ PaoloȱSartor,ȱSoniaȱSeixas,ȱIgnacioȱSobrino,ȱandȱRogerȱVillanuevaȱ

3.1.1 Introduction to the main groups of fished cephalopods in Europe Inȱ nearlyȱ allȱ commercialȱ situationsȱ throughoutȱ Europe,ȱ thereȱ isȱ noȱ speciesȱ determinationȱ inȱ catchesȱ ofȱ cephalopods.ȱ Thisȱ hasȱ beenȱ underlinedȱ inȱ WGCEPHȱ reportsȱoverȱtheȱyearsȱbecauseȱitȱisȱreflectedȱinȱtheȱofficialȱstatisticsȱmadeȱavailableȱtoȱ theȱ scientificȱ community.ȱ Inȱ addition,ȱ cephalopodsȱ areȱ landedȱ byȱ bothȱ commercialȱ andȱ artisanalȱ fleets,ȱ andȱ landingsȱ fromȱ theȱ latterȱ haveȱ beenȱ relativelyȱ poorlyȱ documentedȱ inȱ theȱ past.ȱ Asȱ aȱ resultȱ ofȱ theseȱ problems,ȱ theȱ utilityȱ ofȱ cephalopodȱ landingsȱdataȱforȱassessmentȱofȱstockȱstatusȱhasȱbeenȱcalledȱintoȱquestionȱ(ICES,ȱ2008,ȱ 2009).ȱ Theȱ newȱ métierȬbasedȱ dataȬcollectionȱ systemȱ introducedȱ inȱ 2009ȹmayȱ helpȱ overcomeȱ theseȱ issues,ȱ inȱ thatȱ allȱ speciesȱ caughtȱ inȱ sampledȱ métiersȱ shouldȱ beȱ identifiedȱand,ȱindeed,ȱallȱimportantȱmétiersȱshouldȱbeȱsampledȱ(seeȱSectionȱ5).ȱȱ Officialȱ cephalopodȱ landingȱ statisticsȱ inȱ Europeȱ generallyȱ referȱ toȱ fourȱ groupsȱ ofȱ speciesȱthatȱareȱanalogousȱfromȱcountryȱtoȱcountry,ȱvaryingȱonlyȱtoȱtheȱextentȱthatȱ theirȱcompositionȱreflectsȱspeciesȱdistribution:ȱ(i)ȱlongȬfinnedȱsquid,ȱ(ii)ȱshortȬfinnedȱ squid,ȱ(iii)ȱcuttlefish,ȱandȱ(iv)ȱoctopus.ȱInȱsomeȱcountriesȱ(e.g.ȱSweden),ȱtheȱtwoȱsquidȱ groupsȱareȱlumpedȱtogether.ȱȱ Althoughȱthereȱhasȱbeenȱlittleȱorȱnoȱofficialȱspeciesȱdetermination,ȱtheȱidentityȱofȱtheȱ speciesȱ comprisingȱ eachȱ ofȱ theȱ groupsȱ is,ȱ largely,ȱ knownȱ fromȱ currentȱ orȱ previousȱ scientificȱsampling.ȱInȱsomeȱcases,ȱtheȱproportionsȱofȱtheȱmainȱspeciesȱinȱeachȱofȱtheȱ groupsȱ haveȱ beenȱ determined,ȱ usuallyȱ withinȱ theȱ scopeȱ ofȱ shortȬtermȱ projects.ȱ Proportionsȱofȱ theȱ differentȱ speciesȱ areȱ foundȱ toȱ varyȱ seasonallyȱ andȱ annuallyȱ andȱ mayȱalsoȱdifferȱaccordingȱtoȱtheȱgearȱusedȱ(seeȱBruno,ȱ2008,ȱinȱrelationȱtoȱlandingsȱofȱ IllexȱcoindetiiȱandȱTodaropsisȱeblanaeȱinȱGalicia,ȱSpain).ȱȱ Inȱthisȱsection,ȱshortȱaccountsȱofȱeachȱofȱtheȱmainȱspeciesȱthatȱoccurȱinȱcommercialȱ catchesȱwithinȱEuropeanȱwatersȱareȱpresented,ȱandȱtheȱcountriesȱwhereȱspeciesȱareȱ landedȱ areȱ identified.ȱ Eachȱ ofȱ theseȱ speciesȱ isȱ brieflyȱ described,ȱ andȱ theirȱ mainȱ biologicalȱ andȱ ecologicalȱ charactersȱ areȱ highlighted.ȱ Informationȱ isȱ drawnȱ fromȱ mainstreamȱ scientificȱ publications,ȱ “grey”ȱ literature,ȱ andȱ unpublishedȱ sources,ȱ includingȱoriginalȱresearchȱconductedȱbyȱtheȱauthors.ȱFullȱdescriptionsȱofȱtheȱmainȱ speciesȱwillȱbeȱpresentedȱinȱaȱseparateȱreport.ȱ

3.1.1.1 Long-finned squid Thisȱ groupȱ isȱ composedȱ exclusivelyȱ ofȱ speciesȱ belongingȱ toȱ theȱ familyȱ Loliginidae.ȱ Allȱ ofȱ theȱ speciesȱ exploitedȱ inȱ Europeȱ occurȱ inȱ relativelyȱ shallowȱ waterȱ andȱ areȱ mostlyȱcaughtȱbyȱdemersalȱgear.ȱTheyȱareȱamongȱtheȱmostȱvaluedȱsquidȱspeciesȱandȱ generallyȱcommandȱhigherȱpricesȱinȱtheȱmarketȱthanȱshortȬfinnedȱsquid.ȱ FourȱspeciesȱareȱcaughtȱandȱlandedȱinȱEurope,ȱofȱwhichȱtwoȱareȱrelativelyȱlargeȱandȱ twoȱ areȱ veryȱ small.ȱ Adultsȱ ofȱ theȱ twoȱ largerȱ speciesȱ areȱ similarȱ inȱ sizeȱ andȱ appearanceȱ butȱ canȱ normallyȱ beȱ distinguishedȱ fromȱ eachȱ other,ȱ whereasȱ theȱ twoȱ

10 | Cephalopod biology and fisheries in Europe

smallerȱspeciesȱareȱdifficultȱtoȱdistinguishȱfromȱeachȱotherȱandȱfromȱtheȱjuvenilesȱofȱ theȱ twoȱ largerȱ species.ȱ Theyȱ canȱ occurȱ sympatrically,ȱ andȱ theirȱ identification,ȱ therefore,ȱposesȱproblems.ȱ PostȬrecruitȱ specimensȱ ofȱ Loligoȱ forbesiiȱ canȱ beȱ superficiallyȱ distinguishedȱ fromȱ L.ȱ vulgarisȱprincipallyȱbyȱtheȱlargerȱsizeȱofȱtheȱmedialȱsuckersȱonȱtheȱtentacularȱclubȱinȱL.ȱ vulgaris,ȱwhereasȱinȱL.ȱforbesii,ȱallȱtheȱsuckersȱareȱsubequalȱinȱsizeȱ(Roperȱetȱal.,ȱ1984).ȱ Inȱaddition,ȱtheȱpapillaeȱpresentȱonȱtheȱterminalȱportionȱofȱtheȱhectocotylusȱofȱmatureȱ maleȱ L.ȱ forbesiiȱ areȱ notȱ foundȱ inȱ matureȱ malesȱ ofȱ L.ȱ vulgaris.ȱ Loligoȱ forbesiiȱ hasȱ prominentȱlongitudinalȱstripesȱofȱdarkȱchromatophoresȱonȱtheȱanteriorȱandȱventroȬ lateralȱ surfacesȱ ofȱ theȱ mantle.ȱ Evenȱ thoughȱ largeȱ specimensȱ ofȱ L.ȱ vulgarisȱ canȱ sometimesȱdisplayȱsimilarȱstripes,ȱtheseȱareȱsmallerȱthanȱinȱL.ȱforbesii,ȱlessȱnumerousȱ andȱ differentȱ inȱ appearanceȱ (Jerebȱ andȱ Ragoneseȱ 1986);ȱ thoseȱ ofȱ L.ȱ forbesiiȱ areȱ veryȱ characteristicallyȱ“flameȬlike”:ȱ eachȱstripeȱ isȱ surroundedȱ byȱ whiteȱ tissueȱ(devoidȱ ofȱ chromatophores)ȱ afterȱ whichȱ aȱ lineȱ orȱ borderȱ ofȱ slightlyȱ darkerȱ chromatophoresȱ isȱ visibleȱtoȱdelimitȱtheȱareaȱsurroundingȱtheȱstripe.ȱInȱlife,ȱL.ȱforbesiiȱisȱusuallyȱcolouredȱ withȱmoreȱorangeȱtonesȱandȱL.ȱvulgarisȱwithȱmoreȱpurpleȱtones.ȱ Alloteuthisȱspp.ȱcanȱbeȱseparatedȱfromȱotherȱmembersȱofȱtheȱfamilyȱLoliginidaeȱonȱtheȱ basisȱ ofȱ theirȱ smallȱ sizeȱ asȱ adultsȱ andȱ theȱ factȱ thatȱ theȱ mantleȱ lengthȱ (ML)ȱ isȱ 6Ⱥ–Ⱥ15ȱ timesȱgreaterȱthanȱitsȱwidth,ȱi.e.ȱitȱhasȱaȱnarrowerȱbodyȱthanȱallȱotherȱspecies.ȱAdultȱ maleȱA.ȱsubulataȱandȱA.ȱmediaȱpossessȱtentacularȱclubȱsuckersȱsimilarȱtoȱthoseȱfoundȱ inȱL.ȱvulgaris,ȱbutȱtheȱfinsȱdoȱnotȱextendȱbeyondȱ50Ⱥ%ȱofȱtheȱtotalȱML,ȱexceptȱpossiblyȱ inȱ maleȱ A.ȱ subulata.ȱ Untilȱ recently,ȱ relativeȱ tailȱ lengthȱ wasȱ theȱ characteristicȱ mostȱ commonlyȱ usedȱ toȱ distinguishȱ A.ȱ mediaȱ andȱ A.ȱ subulata.ȱ However,ȱ molecularȱ dataȱ fromȱ Alloteuthisȱ specimensȱ acrossȱ theȱ geographicȱ rangeȱ ofȱ theȱ speciesȱ haveȱ demonstratedȱthisȱcharacteristicȱtoȱbeȱunreliableȱ(Andersonȱetȱal.,ȱ2008).ȱAsȱthisȱraisesȱ doubtsȱ aboutȱ theȱ reliabilityȱ ofȱ previousȱ identifications,ȱ weȱ treatȱ theseȱ twoȱ speciesȱ underȱ Alloteuthisȱ spp.ȱ inȱ thisȱ section,ȱ althoughȱ weȱ continueȱ toȱ referȱ toȱ previousȱ studiesȱusingȱtheȱspeciesȱnameȱasȱdeterminedȱbyȱtheȱauthor(s)ȱofȱthoseȱstudies.ȱ

3.1.1.2 Short-finned squid Thisȱ groupȱ comprisesȱ speciesȱ mainlyȱ belongingȱ toȱ theȱ familyȱ Ommastrephidae.ȱ Occasionallyȱ veryȱ abundantȱ andȱ veryȱ importantȱ inȱ catches,ȱ landings,ȱ andȱ exportsȱ fromȱ Europe,ȱ shortȬfinnedȱ squidȱ areȱ lessȱ valuableȱ inȱ theȱ marketȱ thanȱ longȬfinnedȱ squidȱ andȱ moreȱ proneȱ toȱ extremeȱ variationsȱ inȱ abundance.ȱ Theyȱ allȱ haveȱ pelagicȱ distributionsȱand,ȱalthoughȱsometimesȱcaughtȱdemersally,ȱareȱmainlyȱtakenȱfromȱtheȱ waterȱcolumnȱbyȱtrawlingȱandȱfromȱhighȬrisingȱgillȬȱandȱtrammelnets.ȱ Threeȱ speciesȱ areȱ mostȱ commonlyȱ landedȱ inȱ Europe.ȱ Illexȱ coindetiiȱ andȱ Todaropsisȱ eblanaeȱ mainlyȱ occurȱ inȱ theȱ south,ȱ butȱ bothȱ areȱ occasionallyȱ frequentȱ inȱ theȱ north.ȱ Todarodesȱ sagittatusȱ isȱ moreȱ abundantȱ inȱ theȱ north.ȱ Particularlyȱ inȱ theȱ northernȱ extremes,ȱ Gonatusȱ fabriciiȱ Lichtenstein,ȱ 1818,ȱ aȱ shortȬfinnedȱ squidȱ ofȱ theȱ familyȱ Gonatidae,ȱisȱ taken.ȱ Weȱ haveȱ notȱ includedȱ anȱ accountȱ forȱ thisȱspecies,ȱ orȱforȱotherȱ lessȱ frequentȱ orȱ occasionalȱ species,ȱ butȱ theȱ factȱ thatȱ otherȱ speciesȱ mayȱ beȱ landedȱ meansȱthatȱspecimensȱthatȱdoȱnotȱappearȱtoȱfitȱtheȱdescriptionsȱofȱtheȱmostȱcommonȱ speciesȱrequireȱcarefulȱidentification.ȱȱ ȱ

3.1.1.3 Octopus Landingsȱ inȱ thisȱ categoryȱ consistȱ exclusivelyȱ ofȱ speciesȱ belongingȱ toȱ theȱ familyȱ Octopodidae.ȱCatchesȱandȱlandingsȱareȱdominatedȱinȱweightȱandȱnumberȱbyȱaȱsingleȱ

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species,ȱOctopusȱvulgaris,ȱwhichȱisȱtakenȱinȱgreaterȱnumbersȱinȱtheȱsouthernȱpartȱofȱ theȱICESȱAreaȱandȱfartherȱsouth.ȱLandingsȱofȱtheȱtwoȱEledoneȱspeciesȱinȱtheȱareaȱareȱ increasing.ȱTheȱgenusȱOctopusȱisȱeasyȱtoȱdistinguishȱfromȱtheȱgenusȱEledone,ȱprimarilyȱ byȱtheȱpresenceȱofȱaȱdoubleȱrowȱofȱsuckersȱonȱtheȱarmsȱofȱtheȱformerȱvs.ȱaȱsingleȱrowȱ onȱ theȱ armsȱ ofȱ theȱ latter.ȱ However,ȱ theȱ twoȱ Eledoneȱ speciesȱ mayȱ beȱ difficultȱ toȱ distinguishȱfromȱeachȱotherȱifȱspecimensȱareȱpreservedȱorȱhaveȱbeenȱdeadȱforȱsomeȱ time.ȱ Severalȱ otherȱ Europeanȱ octopusȱ speciesȱ mayȱ beȱ landedȱ asȱ catchȱ fromȱ commercialȱvessels,ȱbutȱtheirȱlowȱabundanceȱmakesȱitȱhardȱtoȱdetermineȱtheȱextentȱtoȱ whichȱtheyȱareȱrepresentativeȱofȱregularȱcatches.ȱ

3.1.1.4 Cuttlefish OfȱtheȱcephalopodȱcategoriesȱlandedȱinȱWesternȱEurope,ȱcuttlefishȱlandingsȱareȱtheȱ highest,ȱmostlyȱfromȱcatchesȱconcentratedȱoverȱtheȱcontinentalȱshelfȱinȱtheȱregionȱofȱ theȱsouthernȱCelticȱSea,ȱtheȱapproachesȱtoȱtheȱEnglishȱChannel,ȱandȱtheȱnortheasternȱ Bayȱ ofȱ Biscay.ȱ Mostȱ landingsȱ areȱ composedȱ ofȱ catchesȱ ofȱ aȱ singleȱ species,ȱ Sepiaȱ officinalis,ȱ butȱ landingsȱ ofȱ atȱ leastȱ twoȱ otherȱ speciesȱ areȱ alsoȱ commonȱ inȱ someȱ Europeanȱcountries.ȱCatchesȱareȱmadeȱdemersallyȱbyȱmeansȱofȱtrawlingȱandȱartisanalȱ trapping.ȱ Theȱ distinctionȱ betweenȱ speciesȱ canȱ poseȱ difficulties,ȱ particularlyȱ inȱ juveniles,ȱ andȱ moreȱ acutelyȱ inȱ someȱ geographicalȱ areasȱ thanȱ inȱ others,ȱ becauseȱ theȱ externalȱ appearanceȱofȱS.ȱofficinalisȱvariesȱacrossȱitsȱdistributionalȱrange.ȱȱ Formerly,ȱS.ȱofficinalisȱincludedȱtwoȱsubspecies,ȱS.ȱo.ȱofficinalisȱandȱS.ȱo.ȱhierredda,ȱtheȱ firstȱdistributedȱinȱtheȱICESȱAreaȱandȱtheȱsecondȱjustȱoutside,ȱtoȱtheȱsouth.ȱAlthoughȱ theyȱareȱveryȱsimilarȱinȱgeneralȱappearance,ȱmorphologicalȱandȱgeneticȱanalysisȱhasȱ demonstratedȱthatȱS.ȱofficinalisȱLinnaeus,ȱ1758ȱandȱS.ȱhierreddaȱRang,ȱ1837,ȱareȱclearlyȱ differentȱspeciesȱofȱtheȱsameȱgenusȱ(Guerraȱetȱal.,ȱ2001).ȱAsȱS.ȱhierreddaȱmayȱalsoȱbeȱ foundȱwithinȱtheȱICESȱArea,ȱitȱisȱusefulȱtoȱknowȱhowȱtoȱtellȱthemȱapart.ȱTheȱmantleȱofȱ S.ȱ hierreddaȱ isȱ narrowerȱ thanȱ inȱ S.ȱ officinalis,ȱ andȱ theȱ unmodifiedȱ armsȱ andȱ theȱ hectocotylizedȱarmȱareȱshorter.ȱTheȱnumberȱofȱtransverseȱrowsȱofȱreducedȱsuckersȱonȱ theȱhectocotylusȱisȱgenerallyȱlargerȱ(8ȱorȱ9ȱtoȱ14)ȱinȱS.ȱhierreddaȱthanȱinȱS.ȱofficinalisȱ(4ȱ orȱ5ȱtoȱ8ȱorȱ9).ȱTheȱstriatedȱzoneȱofȱtheȱcuttleboneȱofȱS.ȱofficinalisȱisȱsomewhatȱsmallerȱ (41Ⱥ%ȱofȱtheȱML)ȱthanȱinȱS.ȱhierreddaȱ(47Ⱥ%).ȱTheȱcuttleboneȱofȱS.ȱofficinalisȱisȱslightlyȱ widerȱ andȱ thickerȱ (38Ⱥ%ȱ andȱ 13Ⱥ%ȱ ofȱ theȱ ML,ȱ respectively)ȱ thanȱ thatȱ ofȱ S.ȱ hierreddaȱ (35Ⱥ%ȱandȱ12Ⱥ%,ȱrespectively).ȱTheȱphragmoconeȱandȱtheȱouterȱconeȱofȱS.ȱofficinalisȱareȱ widerȱthanȱinȱS.ȱhierredda.ȱTheȱcuttleboneȱofȱS.ȱofficinalisȱisȱslightlyȱacuminateȱatȱtheȱ anteriorȱend,ȱwhereasȱitȱisȱveryȱacuminateȱinȱS.ȱhierredda.ȱTheȱspineȱofȱtheȱcuttleboneȱ inȱ S.ȱ officinalisȱ isȱ usuallyȱ coveredȱ byȱ chitin,ȱ especiallyȱ inȱ adults,ȱ whereasȱ thatȱ ofȱ S.ȱ hierreddaȱisȱneverȱcoveredȱbyȱchitin.ȱAsȱS.ȱhierredaȱhasȱonlyȱrecentlyȱbeenȱrecognizedȱ asȱaȱseparateȱspecies,ȱfewȱlandingsȱdataȱhaveȱbeenȱattributedȱtoȱthisȱspecies.ȱAȱspeciesȱ accountȱisȱthereforeȱnotȱincluded.ȱ SepiaȱelegansȱandȱS.ȱorbignyanaȱareȱalsoȱsometimesȱconfusedȱwithȱeachȱotherȱandȱwithȱ S.ȱ officinalisȱ (particularlyȱ youngerȱ individuals).ȱ Sepiaȱ elegans,ȱ however,ȱ isȱ theȱ narrowestȱ ofȱ theȱ three,ȱ whereasȱ S.ȱ orbignyanaȱ hasȱ aȱ moreȱ pronouncedȱ spineȱ atȱ theȱ terminalȱendȱofȱtheȱcuttleboneȱthanȱeitherȱofȱtheȱotherȱtwo.ȱȱ

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3.1.2 Loligo vulgaris Lamarck, 1798 Commonȱ names.ȱ Europeanȱ squidȱ (English);ȱ encornetȱ (French);ȱ Europäischerȱ Langflossenkalmarȱ (German);ȱ calamaryȱ (̍΅Ώ΅ΐΣΕ΍;ȱ Greek);ȱ calamaroȱ comuneȺ1ȱ (Italian);ȱlulaȬvulgarȱ(Portuguese);ȱcalamarȱcomúnȱ(Spanish).ȱȱ

3.1.2.1 Adult diagnostic features Corneaȱpresent.ȱFinsȱrhomboidal,ȱtheirȱlengthȱupȱtoȱca.ȱ66Ⱥ%ȱofȱdorsalȱML.ȱArmsȱwithȱ twoȱrowsȱofȱsuckers;ȱsuckerȱringsȱwithȱ20ȱteeth,ȱtheȱdistalȱonesȱlargeȱandȱpointedȱandȱ theȱproximalȱonesȱminuteȱorȱabsent.ȱTentacularȱclubsȱwithȱfourȱlongitudinalȱrowsȱofȱ suckersȱ inȱ theȱ “manus”,ȱ theȱ suckersȱ inȱ theȱ centralȱ rowsȱ beingȱ largerȱ thanȱ theȱ marginalȱ ones.ȱ Buccalȱ membraneȱ withȱ 15ȱ tinyȱ suckersȱ onȱ theȱ extremity,ȱ eachȱ withȱ chitinousȱrings.ȱLeftȱventralȱarmȱ(IV)ȱofȱmaleȱhectocotylized.ȱSimpleȱfunnelȬlockingȱ cartilageȱ(Roperȱetȱal.,ȱ1984).ȱ

3.1.2.2 Distribution and habitat Loligoȱ vulgarisȱ occursȱ alongȱ theȱ easternȱ Atlanticȱ fromȱ theȱ Britishȱ Islesȱ (55°N)ȱ toȱ theȱ GulfȱofȱGuineaȱ(20°S;ȱRoperȱetȱal.,ȱ1984),ȱextendingȱoutȱtoȱMadeiraȱ(ClarkeȱandȱLu,ȱ 1995),ȱandȱthroughoutȱtheȱMediterraneanȱSeaȱ(Belcari,ȱ1999a).ȱAdultsȱareȱoccasionallyȱ reportedȱoffȱtheȱnorthwestȱcoastȱofȱScotlandȱatȱ57°Nȱ(Pierceȱetȱal.,ȱ1994a;ȱHastieȱetȱal.,ȱ 2009a),ȱtheȱNorthȱSeaȱ(DeȱHeijȱandȱBaayen,ȱ2005),ȱandȱtheȱKattegatȱandȱwesternȱBalticȱ Seaȱ(Jaeckel,ȱ1937;ȱMuus,ȱ1959;ȱHornborg,ȱ2005).ȱParalarvaeȱareȱabsentȱnorthȱofȱtheȱ EnglishȱChannelȱ(Yau,ȱ1994;ȱCollinsȱetȱal.,ȱ2002).ȱL.ȱvulgarisȱisȱaȱnectobenthicȱspeciesȱ thatȱlivesȱinȱtheȱcircumlittoralȱandȱupperȱbathyalȱzonesȱ(Worms,ȱ1983).ȱ

3.1.2.3 Life history Loligoȱvulgarisȱdemonstratesȱhighȱgeographicȱvariabilityȱofȱreproductiveȱandȱgrowthȱ parameters,ȱ andȱ temperatureȱ isȱ oneȱ ofȱ theȱ mainȱ factorsȱ inducingȱ suchȱ variabilityȱ (Morenoȱetȱal.,ȱ2002).ȱMalesȱattainȱgreaterȱlengthsȱ(640ȹmmȱML)ȱthanȱfemalesȱ(540ȹmmȱ ML),ȱwithȱtheȱmaximumȱvaluesȱrecordedȱonȱtheȱSaharanȱBankȱ(Raya,ȱ2001).ȱTheȱlifeȱ cycleȱmayȱbeȱcompletedȱwithinȱapproximatelyȱoneȱyear,ȱwithȱmaximumȱlifespansȱofȱ 15ȹmonthsȱrecordedȱinȱwesternȱIberiaȱ(Morenoȱetȱal.,ȱ1996;ȱRochaȱandȱGuerra,ȱ1999).ȱ Loligoȱvulgarisȱisȱaȱterminalȱspawner,ȱbutȱoocyteȱmaturationȱandȱeggȬlayingȱoccurȱinȱ separateȱbatchesȱduringȱtheȱspawningȱperiodȱ(RochaȱandȱGuerra,ȱ1996;ȱRochaȱetȱal.,ȱ 2001).ȱFecundityȱhasȱbeenȱestimatedȱatȱbetweenȱ10ȹ150ȱandȱ42ȹ000ȱeggsȱ(Baddyr,ȱ1988;ȱ Coelhoȱetȱal.,ȱ1994;ȱGuerraȱandȱRocha,ȱ1994;ȱLaptikhovski,ȱ2000).ȱMeanȱspawningȱageȱ isȱtenȱmonths,ȱandȱmeanȱageȬatȬmaturityȱisȱnineȱmonths.ȱHatchingȱtimeȱsignificantlyȱ influencesȱmeanȱageȬatȬmaturity,ȱwhichȱisȱhigherȱinȱwinterȱcohortsȱthanȱinȱsummerȱ cohortsȱ(Morenoȱetȱal.,ȱ2005).ȱ Theȱspawningȱseasonȱisȱshorterȱinȱtheȱnorthernȱpartȱofȱtheȱdistributionȱ(Morenoȱetȱal.,ȱ 2002;ȱSifnerȱandȱVrgoc,ȱ2004).ȱElsewhere,ȱspawningȱoccursȱthroughoutȱtheȱyear,ȱwithȱ twoȱmainȱpeaksȱbetweenȱNovemberȱandȱJune,ȱalthoughȱtheseȱoccurȱearlierȱinȱmoreȱ southernȱ latitudesȱ andȱ earlierȱ inȱ theȱ Atlanticȱ thanȱ inȱ theȱ Mediterraneanȱ (Baddyr,ȱ 1988;ȱ Coelhoȱ etȱ al.,ȱ1994;ȱGuerraȱ andȱRocha,ȱ1994;ȱMorenoȱ etȱ al.,ȱ 1994,ȱ2002;ȱRocha,ȱ 1994;ȱ Bettencourtȱ etȱ al.,ȱ 1996;ȱ Villaȱ etȱ al.,ȱ 1997;ȱ Rayaȱ etȱ al.,ȱ 1999;ȱ Belcari,ȱ 1999a).ȱ Spawningȱareasȱareȱpoorlyȱknown,ȱbutȱeggȬmassȱrecoveriesȱindicateȱthatȱspawningȱ occursȱatȱdepthsȱofȱ2Ⱥ–Ⱥ120ȹmȱ(Baddyr,ȱ1988;ȱVillaȱetȱal.,ȱ1997;ȱPereiraȱetȱal.,ȱ1998).ȱ

ȱȱȱȱȱȱȱȱȱȱȱȱȱȱȱȱȱȱȱȱȱȱȱȱȱȱȱȱȱȱȱȱȱȱȱȱȱȱȱȱȱȱȱȱȱȱȱȱȱȱȱȱȱȱȱȱȱȱȱ 1 All Italian common names are taken from Bello and Borri (1990).

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3.1.2.4 Trophic ecology Predators.ȱVariousȱcetaceans,ȱseals,ȱlargeȱpelagicȱfish,ȱandȱsomeȱdemersalȱfishȱhaveȱ beenȱ reportedȱ toȱ preyȱ onȱ L.ȱ vulgarisȱ (e.g.ȱ Gonzálezȱ etȱ al.,ȱ 1994a;ȱ OrsiȬRelini,ȱ 1994;ȱ Morteȱetȱal.,ȱ1997;ȱBörjessonȱetȱal.,ȱ2003;ȱSalman,ȱ2004;ȱPeristerakiȱetȱal.,ȱ2005).ȱȱ Prey.ȱ Paralarvaeȱ feedȱ mainlyȱ onȱ crustaceanȱ larvae,ȱ smallȱ mysids,ȱ andȱ fishȱ larvaeȱ (Boletzky,ȱ 1979a).ȱ Verticalȱ migrationsȱ ofȱ paralarvaeȱ andȱ juvenilesȱ areȱ linkedȱ toȱ feeding,ȱ andȱ juvenilesȱ consumeȱ moreȱ planktonicȱ preyȱ thanȱ benthoȬpelagicȱ preyȱ (Nigmatullin,ȱ 1975;ȱ Worms,ȱ 1983;ȱ Turkȱ etȱ al.,ȱ 1986).ȱ Fishȱ speciesȱ increaseȱ inȱ importanceȱ withȱ increasingȱ squidȱ sizeȱ andȱ areȱ theȱ mostȱ frequentȱ preyȱ forȱ adultsȱ (Rochaȱetȱal.,ȱ1994;ȱCoelhoȱetȱal.,ȱ1997;ȱLefkaditou,ȱ2006).ȱ

3.1.2.5 Fisheries InȱtheȱAtlanticȱandȱMediterranean,ȱL.ȱvulgarisȱisȱmainlyȱaȱbycatchȱinȱtheȱmultispeciesȱ bottomȱ andȱ pelagicȱ trawlȱ fisheries.ȱ Inȱ theȱ Englishȱ Channelȱ andȱ Frenchȱ waters,ȱ landingsȱofȱLoligoȱareȱnormallyȱaȱmixtureȱofȱL.ȱvulgarisȱandȱL.ȱforbesiiȱ(theȱspeciesȱareȱ normallyȱ notȱ separatedȱ inȱ theȱ officialȱ statistics).ȱ Fartherȱ southȱ (e.g.ȱ offȱ Spainȱ andȱ Portugal),ȱL.ȱvulgarisȱisȱtheȱdominantȱlongȬfinnedȱsquidȱinȱtheȱlandingsȱ(seeȱChenȱetȱ al.,ȱ 2006).ȱ Itȱ isȱ aȱ secondaryȱ targetȱ speciesȱ inȱ theȱ Saharanȱ Bankȱ cephalopodȱ trawlȱ fisheryȱ (Rayaȱ etȱ al.,ȱ 1999)ȱ andȱ inȱ theȱ westernȱ Portugueseȱ coastalȱ demersalȱ trawlȱ fisheryȱ (Fonsecaȱ etȱ al.,ȱ 2008).ȱ Thereȱ areȱ alsoȱ smallȬscale,ȱ handȬjigȱ artisanalȱ coastalȱ fisheriesȱatȱdepthsȱofȱ20Ⱥ–Ⱥ350ȹm,ȱespeciallyȱinȱSpainȱandȱPortugalȱ(Guerraȱetȱal.,ȱ1994).ȱ Nearȱ theȱ coasts,ȱ whereȱ theȱ speciesȱ concentratesȱ duringȱ autumnȱ andȱ winterȱ forȱ spawning,ȱsmallȬscaleȱprofessionalȱandȱsportȱfishersȱusuallyȱtargetȱtheȱspeciesȱusingȱ handȬjigs.ȱȱ OnȱtheȱGreek,ȱPortuguese,ȱandȱnorthwestȱSpanishȱcoasts,ȱspawningȱaggregationsȱareȱ alsoȱoccasionallyȱtargetedȱbyȱaȱlimitedȱnumberȱofȱfishingȱvesselsȱusingȱbeachȬseineȱ(aȱ gearȱthatȱcatchesȱhigherȱquantitiesȱthanȱhandȬjigsȱandȱotherȱartisanalȱgears),ȱgillnets,ȱ andȱtrammelnets.ȱInȱGaliciaȱ(northwestȱSpain),ȱaȱseasonalȱ(mainlyȱJulyȺȭȺSeptember)ȱ fisheryȱ targetingȱ juvenileȱ L.ȱ vulgarisȱ takesȱ placeȱ insideȱ theȱ Ríasȱ Bajasȱ inȱ watersȱ ofȱ 3ȺȭȺ40ȹmȱdepthȱusingȱseinenetsȱwithȱaȱbagȱ(“boliche”;ȱGuerraȱetȱal.,ȱ1994;ȱTasendeȱetȱal.,ȱ 2005).ȱ Juvenilesȱ generallyȱ recruitȱ toȱ theȱ fisheryȱ atȱ approximatelyȱ 5ȹmonthsȱ ofȱ ageȱ (Morenoȱ etȱ al.,ȱ 2007).ȱ Nevertheless,ȱ theȱ ageȱ andȱ sizeȱ atȱ recruitmentȱ toȱ theȱ fisheryȱ obviouslyȱ dependsȱ onȱ theȱ fishingȱ gearȱ used.ȱ Thereȱ isȱ noȱ regularȱ assessmentȱ andȱ managementȱ forȱ L.ȱ vulgarisȱ fisheries,ȱ butȱ landingȱ sizeȱ restrictionsȱ areȱ imposedȱ inȱ Portugalȱ(andȱinȱSpain).ȱ TheȱspeciesȱhasȱbeenȱidentifiedȱinȱcommercialȱlandingsȱinȱtheȱUK,ȱFrance,ȱPortugal,ȱ Spain,ȱItaly,ȱandȱGreece.ȱ

3.1.3 Loligo forbesii Steenstrup, 1856 Commonȱ names.ȱ Veinedȱ squidȱ (English);ȱ encornetȱ veinéȱ (French);ȱ Langflossenkalmarȱ (German);ȱ calamaryȱ (̍΅Ώ΅ΐΣΕ΍;ȱ Greek);ȱ calamaroȱ venato,ȱ occhioneȱ(Italian);ȱcalamarȱveteadoȱ(Spanish);ȱlulaȬriscadaȱ(Portuguese).ȱ

3.1.3.1 Adult diagnostic features Corneaȱpresent.ȱRhomboidalȱfinsȱofȱlengthȱca.ȱ75Ⱥ%ȱofȱdorsalȱML,ȱposteriorȱbordersȱ slightlyȱ concave.ȱ Armsȱ withȱ twoȱ rowsȱ ofȱ suckers.ȱ Suckersȱ onȱ theȱ manusȱ ofȱ theȱ tentacularȱ clubȱ areȱ subequalȱ inȱ size;ȱ suckerȱ ringsȱ withȱ 13ȺȭȺ18ȱ sharpȱ conicalȱ teeth.ȱ UnlikeȱL.ȱvulgaris,ȱitȱlacksȱenlargedȱmedialȱsuckersȱonȱtheȱtentacularȱclubȱ(suckersȱinȱ allȱ fourȱ rowsȱ areȱ fairlyȱ similarȱ inȱ size).ȱ Inȱ males,ȱ theȱ leftȱ ventralȱ armȱ (IV)ȱ isȱ hectocotylizedȱ inȱ itsȱ distalȱ thirdȱ byȱ modificationȱ ofȱ suckersȱ intoȱ longȱ papillaeȱ thatȱ

14 | Cephalopod biology and fisheries in Europe

graduallyȱ decreaseȱ inȱ sizeȱ distally.ȱ Simpleȱ funnelȬlockingȱ cartilage.ȱ Diagnosisȱ adaptedȱfromȱRoperȱetȱal.ȱ(1984).ȱ

3.1.3.2 Distribution and habitat Loligoȱforbesiiȱisȱaȱneriticȱandȱlargelyȱdemersalȱspeciesȱoccurringȱinȱcoastalȱwatersȱandȱ theȱcontinentalȱshelfȱfromȱtheȱFaroeȱIslandsȱtoȱ20°NȱinȱtheȱeasternȱAtlantic,ȱincludingȱ theȱ Northȱ Sea,ȱ extendingȱ intoȱ theȱ Mediterraneanȱ andȱ outȱ toȱ Madeira,ȱ theȱ Canaryȱ Islands,ȱandȱtheȱAzores.ȱTheȱspeciesȱisȱrelativelyȱrareȱsouthȱofȱtheȱBayȱofȱBiscayȱandȱisȱ absentȱfromȱtheȱBalticȱSeaȱ(Roperȱetȱal.,ȱ1984).ȱ

3.1.3.3 Life history Loligoȱ forbesiiȱ isȱ commonlyȱ assumedȱ toȱ haveȱ anȱ annualȱ lifeȱ cycle,ȱ withȱ pulsesȱ ofȱ recruitmentȱ inȱ Aprilȱ andȱ AugustȺȭȺNovemberȱ (LumȬKongȱ etȱ al.,ȱ 1992;ȱ Pierceȱ etȱ al.,ȱ 1994a;ȱCollinsȱetȱal.,ȱ1995,ȱ1997;ȱBelcari,ȱ1999b),ȱalthoughȱsomeȱindividualsȱmayȱreachȱ anȱ ageȱ ofȱ ca.ȱ 18ȹmonthsȱ (Rochaȱ andȱ Guerra,ȱ 1999).ȱ Asȱ inȱ otherȱ loliginidȱ squid,ȱ theȱ growthȱpatternȱcomprisesȱanȱearlyȱrapidȱ“exponential”ȱphaseȱfollowedȱbyȱaȱslowerȱ “logarithmic”ȱphaseȱ(ForsytheȱandȱvanȱHeukelem,ȱ1987;ȱForsytheȱandȱHanlon,ȱ1989).ȱ Malesȱcanȱreachȱsizesȱlargerȱ(610ȹmmȱMLȱonȱtheȱcontinentalȱshelf;ȱ940ȹmmȱinȱAzores)ȱ thanȱ femalesȱ (420ȹmmȱ andȱ 460ȹmm),ȱ althoughȱ someȱ malesȱ matureȱ atȱ muchȱ smallerȱ sizesȱ(fromȱca.ȱ120ȹmm;ȱBoyleȱetȱal.,ȱ1995),ȱapparentlyȱrepresentingȱalternativeȱgrowthȱ andȱmaturationȱstrategies,ȱandȱleadingȱtoȱtheȱpresenceȱofȱ(atȱleast)ȱtwoȱmicrocohortsȱ inȱtheȱfishedȱpopulationȱ(Collinsȱetȱal.,ȱ1999).ȱ Loligoȱforbesiiȱisȱsemelparous,ȱdisplayingȱ“intermittent,ȱterminalȱspawning”,ȱinȱwhichȱ theȱ femalesȱ layȱ eggsȱ inȱ batchesȱ andȱ dieȱ shortlyȱ afterȱ spawningȱ (Rochaȱ etȱ al.,ȱ 2001).ȱ Manyȱstudiesȱdescribeȱanȱextendedȱspawningȱseason,ȱalthoughȱitȱisȱunclearȱhowȱlongȱ anȱ individualȱ continuesȱ toȱ spawnȱ (Guerraȱ andȱ Rocha,ȱ 1994;ȱ Morenoȱ etȱ al.,ȱ 1994;ȱ Collinsȱetȱal.,ȱ1995).ȱFemaleȱL.ȱforbesiiȱproduceȱonlyȱaȱfewȱthousandȱ(upȱtoȱ23ȹ000)ȱeggsȱ inȱtheirȱlifetime,ȱwithȱlargerȱfemalesȱproducingȱmoreȱeggs.ȱȱ Clustersȱ ofȱ eggsȱ areȱ normallyȱ attachedȱ toȱ substrataȱ thatȱ includeȱ algae,ȱ shells,ȱ rockȱ crevices,ȱ nets,ȱ ropes,ȱ creels,ȱ andȱ otherȱ fishingȱ gearȱ thatȱ remainsȱ submergedȱ andȱ undisturbed.ȱ Thereȱ areȱ fewȱ reportsȱ ofȱ eggsȱ fromȱ offshoreȱ waters,ȱ butȱ thisȱ mayȱ beȱ becauseȱ theȱ squidȱ spawnȱ overȱ rockyȱ substratum,ȱ makingȱ spawningȱ areasȱ largelyȱ inaccessibleȱtoȱtrawlingȱ(Holme,ȱ1974;ȱLordanȱandȱCasey,ȱ1999).ȱȱ

3.1.3.4 Trophic ecology Predators.ȱ Variousȱ cetaceanȱ species,ȱ seals,ȱ andȱ largeȱ demersalȱ fishȱ haveȱ beenȱ reportedȱ toȱ preyȱ onȱ L.ȱ forbesiiȱ (e.g.ȱ Pierceȱ andȱ Santos,ȱ 1996;ȱ Santosȱ etȱ al.,ȱ 2004a;ȱ Deȱ Pierrepontȱetȱal.,ȱ2005).ȱȱ Prey.ȱ Aȱ largeȱ numberȱ ofȱ preyȱ species,ȱ includingȱ variousȱ fish,ȱ crustaceans,ȱ polychaetes,ȱ andȱ molluscsȱ haveȱ beenȱ identifiedȱ inȱ L.ȱ forbesiiȱ stomachs.ȱ Asȱ inȱ mostȱ squid,ȱfishȱareȱmoreȱimportantȱinȱadults,ȱandȱcrustaceansȱmoreȱimportantȱinȱjuvenileȱ stagesȱ(e.g.ȱPierceȱetȱal.,ȱ1994d).ȱ

3.1.3.5 Fisheries Loligoȱ forbesiiȱ isȱ theȱ dominantȱ squidȱ speciesȱ fishedȱ northȱ ofȱ theȱ Englishȱ Channel,ȱ mainlyȱasȱbycatchȱfromȱdemersalȱtrawlsȱ(Pierceȱetȱal.,ȱ1994b),ȱalthoughȱvariousȱsmallȱ directedȱ trawlȱ fisheriesȱ existȱ inȱ coastalȱ watersȱ (e.g.ȱ Youngȱ etȱ al.,ȱ 2006a)ȱ andȱ areȱ becomingȱincreasinglyȱimportant.ȱItȱisȱtheȱonlyȱsquidȱspeciesȱofȱeconomicȱimportanceȱ inȱtheȱAzores,ȱwhereȱitȱisȱfishedȱbyȱanȱartisanalȱfleetȱequippedȱwithȱhandlinesȱandȱ homemadeȱ jigsȱ (Martins,ȱ 1982;ȱ Porteiroȱ andȱ Martins,ȱ 1994).ȱ Southȱ ofȱ theȱ Englishȱ

ICES Cooperative Research Report No. 303 | 15

Channel,ȱtheȱspeciesȱisȱcurrentlyȱlessȱcommon,ȱbutȱisȱtakenȱbothȱasȱaȱtrawlȱbycatchȱ andȱbyȱcoastalȱartisanalȱboatsȱusingȱjigsȱ(CunhaȱandȱMoreno,ȱ1994;ȱGuerraȱetȱal.,ȱ1994;ȱ Rochaȱetȱal.,ȱ1994).ȱAnalysisȱofȱmorphometricȱvariationȱandȱtheȱuseȱofȱallozymeȱandȱ microsatelliteȱ markersȱ suggestȱ thatȱ thereȱ isȱ noȱ significantȱ separationȱ ofȱ stocksȱ throughoutȱtheȱrangeȱofȱitsȱdistributionȱonȱmainlandȱcoasts.ȱTheȱAzoresȱpopulationȱ probablyȱ representsȱ aȱ separateȱ subspecies,ȱ andȱ animalsȱ fromȱ offshoreȱ banksȱ (e.g.ȱ Rockall)ȱ demonstrateȱ someȱ differencesȱ fromȱ theȱ coastalȱ populationȱ (Pierceȱ etȱ al.,ȱ 1994c;ȱ Brierleyȱ etȱ al.,ȱ 1995;ȱ Shawȱ etȱ al.,ȱ 1999).ȱ Itȱ isȱ alsoȱ consistentlyȱ presentȱ inȱ theȱ easternȱMediterraneanȱtrawlȱcatchesȱ(Machiasȱetȱal.,ȱ2001).ȱ TheȱspeciesȱhasȱbeenȱidentifiedȱinȱcommercialȱlandingsȱinȱtheȱUK,ȱFrance,ȱPortugal,ȱ Spain,ȱItaly,ȱandȱGreece.ȱ

3.1.4 Alloteuthis spp.

Common names Alloteuthisȱ subulataȱ Lamarck,ȱ 1798.ȱ Europeanȱ commonȱ squidȱ (English);ȱ casseronȱ communȱ (French);ȱ Gepfriemterȱ Langflossenkalmarȱ (German);ȱ calamarettoȱ puntutoȱ (Italian);ȱlulaȬbicudaȬcompridaȱ(Portuguese);ȱcalamarínȱpicudoȱ(Spanish).ȱ AlloteuthisȱmediaȱLinnaeus,ȱ1758.ȱMidsizeȱsquidȱ(English);ȱcasseronȱbambouȱ(French);ȱ calamarettoȱ comuneȱ (Italian);ȱ lulaȬbicudaȬcurtaȱ (Portuguese);ȱ calamarínȱ menorȱ (Spanish).ȱ

3.1.4.1 Adult diagnostic features Elongatedȱ mantle.ȱ Rhomboidalȱ finsȱ attenuateȱ posteriorlyȱ intoȱ aȱ long,ȱ slenderȱ tail,ȱ whichȱcanȱbeȱmoreȱthanȱ50Ⱥ%ȱofȱtheȱMLȱinȱmales.ȱArmsȱwithȱtwoȱrowsȱofȱsuckers.ȱ Tentacleȱclubsȱwithȱfourȱrowsȱofȱsuckers.ȱTheȱcentralȱrowsȱofȱsuckersȱareȱ3Ⱥ–Ⱥ4ȱtimesȱ largerȱ thanȱ theȱ marginalȱ suckersȱ (Yau,ȱ 1994;ȱ Laptikhovskyȱ etȱ al.,ȱ 2002).ȱ Theȱ leftȱ ventralȱ armȱ (IV)ȱ canȱ beȱ hectocotylizedȱ inȱ matureȱ males,ȱ withȱ 6Ⱥ–Ⱥ8ȱ pairsȱ ofȱ normalȱ suckersȱproximallyȱfollowedȱbyȱtwoȱlongitudinalȱrowsȱofȱfineȱpapillaeȱdistallyȱ(Yau,ȱ 1994).ȱMaximumȱMLȱofȱA.ȱsubulataȱisȱ20ȹcmȱ(Rodhouseȱetȱal.,ȱ1988).ȱȱ

3.1.4.2 Distribution and habitat AlloteuthisȱsubulataȱandȱA.ȱmediaȱareȱsympatricȱthroughoutȱmuchȱofȱtheȱrangeȱofȱbothȱ speciesȱ (Roperȱ etȱ al.ȱ 1984),ȱ extendingȱ fromȱ approximatelyȱ 60°Nȱ toȱ 20°Nȱ (Guerra,ȱ 1992).ȱTheyȱareȱfoundȱinȱtheȱNorthȱSeaȱandȱtheȱMediterraneanȱ(Belcari,ȱ1999ȱc,ȱ1999d).ȱ Theyȱ bothȱ occurȱ primarilyȱ inȱ shelfȱ watersȱ andȱ areȱ demersal,ȱ althoughȱ A.ȱ mediaȱ isȱ reportedȱtoȱmigrateȱoffshoreȱinȱwinterȱ(Roperȱetȱal.,ȱ1984)ȱandȱtoȱmakeȱdielȱverticalȱ migrationsȱ(ZuevȱandȱNesis,ȱ1971).ȱȱ

3.1.4.3 Life history Hatchlingsȱofȱbothȱspeciesȱareȱ2Ⱥ–Ⱥ3ȹmmȱlongȱ(ZuevȱandȱNesis,ȱ1971;ȱYau,ȱ1994).ȱInȱA.ȱ subulata,ȱ sexualȱ maturationȱ canȱ beȱ achievedȱ atȱ 4Ⱥ–Ⱥ5ȹcmȱ MLȱ inȱ bothȱ sexes,ȱ althoughȱ matureȱ malesȱ haveȱ aȱ widerȱ rangeȱ ofȱ sizesȱ thanȱ matureȱ femalesȱ andȱ reachȱ 50Ⱥ%ȱ maturityȱ atȱ aȱ slightlyȱ smallerȱ sizeȱ (7Ⱥ–Ⱥ7.5ȹcm,ȱ comparedȱ withȱ 7.5Ⱥ–Ⱥ8ȹcmȱ inȱ females).ȱ AlloteuthisȱmediaȱhasȱanȱMLȱofȱupȱtoȱ12ȹcmȱinȱfemalesȱandȱ9ȹcmȱinȱmales.ȱIndividualsȱ hatchingȱinȱspringȱgrowȱ7Ⱥ–Ⱥ8ȹmmȱmonthȺȬ1ȱduringȱsummerȱandȱreachȱanȱMLȱofȱ7Ⱥ–Ⱥ8ȹcmȱ theȱ followingȱ springȱ (Zuevȱ andȱ Nesis,ȱ 1971).ȱ Bothȱ speciesȱ areȱ thoughtȱ toȱ liveȱ forȱ approximatelyȱ oneȱ yearȱ (Rodhouseȱ etȱ al.,ȱ 1988;ȱ MangoldȬWirz,ȱ 1963;ȱ Alidromiti,ȱ 2007).ȱ

16 | Cephalopod biology and fisheries in Europe

TheȱaverageȱestimatedȱpotentialȱfecundityȱofȱfemaleȱA.ȱsubulataȱisȱ2200Ⱥ–Ⱥ13ȹ500ȱeggsȱ perȱ femaleȱ (meanȱ ~Ⱥ5900ȹ±ȹ970ȱ s.e.;ȱ Hastieȱ etȱ al.,ȱ 2009b).ȱ Inȱ A.ȱ media,ȱ theȱ potentialȱ fecundityȱ hasȱ beenȱ estimatedȱ atȱ 950Ⱥ–Ⱥ1400ȱ eggsȱ forȱ theȱ westernȱ Mediterraneanȱ andȱ 1500Ⱥ–Ⱥ2500ȱforȱtheȱeasternȱMediterranean.ȱOocyteȱmaturationȱoccursȱinȱbatches.ȱȱ Spawningȱ occursȱ throughoutȱ theȱ year,ȱ generallyȱ withȱ largerȱ numbersȱ ofȱ matureȱ squidȱinȱ springȺȭȺsummerȱandȱ smallerȱnumbersȱ inȱautumnȺȭȺwinterȱ (Rodhouseȱ etȱ al.,ȱ 1988;ȱNyegaard,ȱ2001;ȱHastieȱetȱal.,ȱ2009b).ȱAȱpredominanceȱofȱsmallȱsquidȱ(<Ⱥ50ȹmmȱ ML)ȱisȱobservedȱduringȱautumnȱ(Hastieȱetȱal.,ȱ2009b).ȱAdultsȱofȱA.ȱmediaȱmigrateȱtoȱ shallowȱ water,ȱ spawningȱ atȱ depthsȱ ofȱ 10Ⱥ–Ⱥ100ȹmȱ onȱ sand,ȱ seagrassȱ meadows,ȱ etc.ȱ fromȱMarchȱtoȱOctoberȱinȱtheȱwesternȱMediterraneanȱ(MangoldȬWirz,ȱ1963)ȱandȱyearȬ roundȱinȱtheȱcentralȱandȱeasternȱregionȱ(LoȱBianco,ȱ1909;ȱNaef,ȱ1923;ȱLaptikhovskyȱetȱ al.,ȱ 2002;ȱ Lefkaditou,ȱ 2006).ȱ Inȱ theȱ Northȱ Sea,ȱ spawningȱ takesȱ placeȱ inȱ JuneȺȭȺJulyȱ (ZuevȱandȱNesis,ȱ1971).ȱRecruitsȱofȱA.ȱmediaȱareȱpresentȱpracticallyȱallȱyearȬroundȱinȱ shallowerȱwatersȱofȱtheirȱdistributionȱ(ReliniȱandȱOrsiȬRelini,ȱ1984).ȱSeasonalȱpeaksȱofȱ recruitmentȱareȱmoreȱevidentȱinȱareasȱwithȱaȱwiderȱcontinentalȱshelfȱ(MangoldȬWirz,ȱ 1963;ȱPapaconstantinouȱetȱal.,ȱ1994).ȱȱ

3.1.4.4 Trophic ecology Predators.ȱ Alloteuthisȱ subulataȱ hasȱ beenȱ identifiedȱ fromȱ theȱ stomachȱ contentsȱ ofȱ severalȱ speciesȱ ofȱ marineȱ mammalȱ (Santosȱ etȱ al.,ȱ 2001a,ȱ 2001b,ȱ 2004a,ȱ 2005a,ȱ 2005b;ȱ Gonzálezȱetȱal.,ȱ1994a;ȱMeynier,ȱ2004)ȱandȱfromȱhakeȱ(Merlucciusȱmerluccius;ȱDalyȱetȱ al.,ȱ2001),ȱandȱLoligoȱspp.ȱ(Pierceȱetȱal.,ȱ1994d;ȱRochaȱetȱal.,ȱ1994).ȱAlloteuthisȱmediaȱisȱ eatenȱ byȱ variousȱ gadidȱ andȱ elasmobranchȱ fishȱ (Zuevȱ andȱ Nesis,ȱ 1971;ȱ Bello,ȱ 1997),ȱ andȱisȱanȱimportantȱfoodȱforȱdemersalȱfishȱinȱtheȱsouthernȱBayȱofȱBiscayȱ(Velascoȱetȱ al.,ȱ2001).ȱ Prey.ȱ Theȱ mainȱ preyȱ ofȱ A.ȱ subulataȱ inȱ theȱ Irishȱ Seaȱ comprisesȱ clupeidȱ fishȱ andȱ crustaceansȱ(Nyegaard,ȱ2001).ȱTheȱdietȱofȱA.ȱmediaȱconsistsȱofȱlarvaeȱandȱjuvenilesȱofȱ fish,ȱcopepods,ȱandȱeuphausiidsȱ(ZuevȱandȱNesis,ȱ1971).ȱȱ

3.1.4.5 Fisheries BothȱA.ȱmediaȱandȱA.ȱsubulataȱcanȱbeȱcaughtȱandȱmisidentifiedȱasȱjuvenileȱLoligoȱandȱ canȱ occurȱ amongȱ longȬfinnedȱ squidȱ catchesȱ throughoutȱ theirȱ range.ȱ Inȱ theȱ Gulfȱ ofȱ Cadiz,ȱAlloteuthisȱspp.ȱareȱcaughtȱasȱaȱbycatchȱofȱtheȱmultispeciesȱbottomȬtrawlȱfleet,ȱ andȱrecordedȱannualȱlandingsȱvariedȱbetweenȱ55ȱandȱ290ȹtȱduringȱ1996Ⱥ–Ⱥ2006.ȱBothȱ Alloteuthisȱspeciesȱareȱlanded,ȱprobablyȱinȱsimilarȱamountsȱ(I.ȱSobrino,ȱpers.ȱcomm.).ȱ InȱtheȱRíasȱBajasȱofȱGaliciaȱ(northwestȱSpain),ȱAlloteuthisȱisȱaȱbycatchȱofȱtheȱboatȬseineȱ fisheryȱdirectedȱatȱL.ȱvulgarisȱ(Tasendeȱetȱal.,ȱ2005).ȱStudiesȱonȱbottomȬtrawlȱdiscardsȱ haveȱ demonstratedȱ that,ȱ inȱ theȱ northeasternȱ Mediterranean,ȱ A.ȱ mediaȱ isȱ totallyȱ discardedȱ(Machiasȱetȱal.,ȱ2001),ȱwhereasȱinȱtheȱwesternȱMediterranean,ȱitȱisȱnormallyȱ landedȱ(Sartorȱetȱal.,ȱ1998).ȱ TheseȱspeciesȱhaveȱbeenȱidentifiedȱinȱcommercialȱlandingsȱinȱSpainȱandȱareȱregularlyȱ landedȱinȱItalyȱ(Belcari,ȱ1999c,ȱ1999d).ȱȱ

3.1.5 Illex coindetii Verany, 1839 Commonȱnames.ȱBroadtailȱshortfinȱsquidȱ(English);ȱfauxȱencornetȱ(French);ȱthrapsaloȱ (̋ΕΣΜ΅ΏΓ;ȱ Greek);ȱ totano,ȱ todaroȱ (Italian);ȱ potaȱ voladoraȱ (Spanish);ȱ potaȬvoadoraȱ (Portuguese).ȱ

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3.1.5.1 Adult diagnostic features Mantleȱwidthȱ15Ⱥ–Ⱥ25Ⱥ%ȱofȱtheȱML.ȱRhomboidalȱfinsȱofȱwidthȱ45Ⱥ–Ⱥ60Ⱥ%ȱofȱdorsalȱML.ȱ Armsȱwithȱtwoȱrowsȱofȱsuckers.ȱTentacularȱclubȱwithȱeightȱtransverseȱrowsȱofȱminuteȱ subequalȱ suckers.ȱ Leftȱ orȱ rightȱ ventralȱ armȱ ofȱ maleȱ hectocotylizedȱ withȱ distalȱ trabeculaeȱ modifiedȱ toȱ papilloseȱ flaps.ȱ Inverseȱ TȬshapedȱ funnelȱ cartilage.ȱ Funnelȱ grooveȱ withȱ anteriorȱ foveolaȱ andȱ withoutȱ sideȱ pockets.ȱ Diagnosisȱ adaptedȱ fromȱ Roperȱetȱal.ȱ(1984,ȱ1998)ȱandȱRoperȱandȱMangoldȱ(1998).ȱ

3.1.5.2 Distribution and habitat Illexȱ coindetiiȱ isȱ foundȱ inȱ theȱ Mediterraneanȱ Sea,ȱ theȱ easternȱ Atlanticȱ fromȱ 60°Nȱ toȱ 17°Sȱandȱ30°W,ȱandȱinȱwesternȱAtlanticȱwatersȱfromȱtheȱGulfȱofȱMexicoȱtoȱVenezuelaȱ (Roperȱetȱal.,ȱ1998;ȱBelcari,ȱ1999e).ȱItȱisȱdemersalȱinȱtheȱmiddleȱandȱlowerȱsublittoralȱ andȱupperȱbathyalȱinȱtemperateȱlatitudes,ȱandȱitȱundertakesȱdielȱverticalȱmigrations.ȱ

3.1.5.3 Life history IllexȱcoindetiiȱisȱaȱmediumȬsizedȱsquidȱ(Sánchez,ȱP.,ȱetȱal.,ȱ1998).ȱTheȱmaximumȱMLȱisȱ generallyȱ largerȱ inȱ femalesȱ thanȱ males.ȱ Theȱ largestȱ sizesȱ areȱ foundȱ inȱ Northeastȱ Atlanticȱ populations,ȱ withȱ malesȱ growingȱ toȱ 370ȹmmȱ andȱ femalesȱ toȱ 320ȹmmȱ (Gonzálezȱ etȱ al.,ȱ 1996;ȱ Sánchez,ȱ P.,ȱ etȱ al.,ȱ 1998).ȱ Theȱ lifeȱ cycleȱ isȱ probablyȱ annual,ȱ althoughȱ lifespanȱ estimatesȱ varyȱ fromȱ sixȱ monthsȱ (basedȱ onȱ statolithȱ increments;ȱ Arkhipkinȱetȱal.,ȱ1998)ȱtoȱ24ȹmonthsȱ(lengthȱfrequencyȱanalyses;ȱMangoldȬWirz,ȱ1963).ȱ TheȱageȱatȱwhichȱI.ȱcoindetiiȱmaturesȱisȱvariable:ȱ100Ⱥ–Ⱥ271ȱdaysȱinȱmalesȱandȱ140Ⱥ–Ⱥ285ȱ daysȱinȱfemalesȱ(GonzálezȱandȱGuerra,ȱ1996;ȱGonzálezȱetȱal.,ȱ1996;ȱArkhipkinȱetȱal.,ȱ 2000).ȱTheȱnumberȱofȱmatureȱeggsȱinȱtheȱovaryȱandȱoviductsȱrangesȱfromȱ50ȹ000ȹtoȱ 200ȹ000.ȱItȱisȱanȱ“intermittentȱspawner”,ȱwithȱfemalesȱspawningȱseveralȱtimesȱduringȱ aȱprolongedȱspawningȱperiodȱlastingȱfromȱaȱfewȱdaysȱtoȱaȱfewȱweeksȱ(Gonzálezȱandȱ Guerra,ȱ 1996).ȱ Spawningȱ mayȱ occurȱ yearȬround,ȱ butȱ seasonalȱ peaksȱ existȱ andȱ varyȱ widelyȱthroughȱtheȱgeographicȱrangeȱ(e.g.ȱSánchez,ȱ1984;ȱTursiȱandȱD’Onghia,ȱ1992;ȱ SánchezȱandȱMartin,ȱ1993;ȱJerebȱandȱRagonese,ȱ1995;ȱBelcari,ȱ1996;ȱArvanitidisȱetȱal.,ȱ 2002;ȱHernándezȬGarcia,ȱ2002;ȱCeriolaȱetȱal.,ȱ2006).ȱȱ

3.1.5.4 Trophic ecology Predators.ȱIllexȱcoindetiiȱhasȱbeenȱfoundȱinȱtheȱstomachsȱofȱvariousȱmarineȱmammals,ȱ includingȱ Risso’sȱ dolphinȱ (Grampusȱ griseus),ȱ bottlenoseȱ dolphinȱ (Tursiopsȱ truncatus)ȱ (Santos,ȱ M.ȱ B.,ȱ etȱ al.,ȱ 1997,ȱ 2007),ȱ longȬfinnedȱ pilotȱ whaleȱ (Globicephalaȱ melas),ȱ andȱ commonȱ dolphinȱ (Delphinusȱ delphis)ȱ (Gonzálezȱ etȱ al.,ȱ 1994a;ȱ Silva,ȱ 1999).ȱ Itȱ hasȱ alsoȱ beenȱrecordedȱfromȱswordfishȱ(Xiphiasȱgladius;ȱBello,ȱ1985),ȱgreaterȱforkbeardȱ(Phycisȱ blennoides;ȱMorteȱ etȱ al.,ȱ2002),ȱandȱ theȱsharpnoseȱ sevengillȱ sharkȱ (Heptranchiasȱ perlo;ȱ HendersonȱandȱWilliams,ȱ2001).ȱ Prey.ȱTheȱdietȱofȱI.ȱcoindetiiȱisȱcomposedȱofȱfish,ȱcrustaceans,ȱandȱcephalopods;ȱitȱisȱ alsoȱcannibalistic.ȱItȱisȱmainlyȱaȱneritic,ȱnektoȬbenthicȱpredatorȱwithȱaȱwideȱspectrumȱ ofȱprey,ȱbutȱadultsȱperformȱsignificantȱverticalȱmigrationȱandȱalsoȱtakeȱpelagicȱpreyȱ (Sánchez,ȱP.,ȱetȱal.,ȱ1998).ȱ

3.1.5.5 Fisheries Illexȱcoindetiiȱisȱtakenȱthroughoutȱtheȱyearȱasȱbycatchȱinȱbottomȱandȱpelagicȱtrawls,ȱ and,ȱ toȱ aȱ lesserȱ extent,ȱ inȱ gillȬȱ andȱ trammelnets,ȱ atȱ depthsȱ ofȱ 100Ⱥ–Ⱥ400ȹmȱ inȱ theȱ Mediterranean,ȱ offȱ Westȱ Africa,ȱ andȱ inȱ theȱ Northeastȱ Atlantic.ȱ Itsȱ fisheryȱ valueȱ isȱ increasing.ȱAlthoughȱseparateȱstatisticsȱareȱgenerallyȱnotȱreported,ȱtheȱannualȱcatchȱ

18 | Cephalopod biology and fisheries in Europe

forȱtheȱlastȱdecadeȱprobablyȱrangedȱfromȱ5000ȹtȱtoȱ12ȹ000ȹt.ȱThereȱisȱthoughtȱtoȱbeȱaȱ singleȱstockȱthroughoutȱtheȱregion.ȱ Theȱ speciesȱ hasȱ beenȱ identifiedȱ inȱ commercialȱ landingsȱ inȱ Ireland,ȱ theȱ UK,ȱ France,ȱ Spain,ȱandȱPortugalȱ(inȱtheȱNortheastȱAtlantic),ȱandȱSpain,ȱItaly,ȱandȱGreeceȱ(inȱtheȱ Mediterranean).ȱ

3.1.6 Todarodes sagittatus Lamarck, 1798 Commonȱ names.ȱ Europeanȱ flyingȱ squidȱ (English);ȱ toutenonȱ communȱ (French);ȱ Pfeilkalmarȱ (German);ȱ katamachiȱ (̍΅Θ΅ΐΣΛ΍;ȱ Greek);ȱ beitusmokkurȱ (Icelandic);ȱ totanoȱviolaȱ(Italian);ȱakkarȱ(Norwegian);ȱpotaȬeuropeiaȱ(Portuguese);ȱpotaȱeuropeaȱ (Spanish).ȱ

3.1.6.1 Adult diagnostic features Tentacleȱ clubȱ suckersȱ onȱ dactylusȱ areȱ arrangedȱ inȱ fourȱ rows,ȱ suckersȱ onȱ elongateȱ carpusȱinȱ10ȺȭȺ12ȱpairs;ȱentireȱclubȱrelativelyȱlong,ȱextendingȱalongȱstalk.ȱArmȱsuckersȱ withȱ enlargedȱ centralȱ tooth,ȱ 7ȺȭȺ9ȱ regularȱ teeth,ȱ andȱ virtuallyȱ noȱ smallȱ alternatingȱ teeth.ȱ Funnelȱ grooveȱ withȱ anteriorȱ foveolaȱ andȱ withoutȱ sideȱ pockets.ȱ Reference:ȱ Roperȱetȱal.ȱ(1984).ȱ

3.1.6.2 Distribution and habitat TodarodesȱsagittatusȱisȱfoundȱthroughoutȱtheȱeasternȱAtlanticȱtoȱca.ȱ40°W,ȱandȱfromȱtheȱ Arcticȱ Oceanȱ toȱ ca.ȱ 13°S.ȱ Theȱ rangeȱ includesȱ theȱ Northȱ Seaȱ andȱ theȱ Mediterraneanȱ (Clarke,ȱ1966;ȱZuevȱetȱal.,ȱ1976;ȱRoperȱetȱal.,ȱ1984).ȱTheyȱoccurȱinȱbothȱtheȱopenȱoceanȱ andȱcoastalȱwaters,ȱandȱfromȱtheȱsurfaceȱtoȱnearȬbottomȱatȱdepthsȱupȱtoȱ2500ȹm.ȱTheyȱ migrateȱ verticallyȱ betweenȱ theȱ surfaceȱ atȱ nightȱ andȱ nearȬbottomȱ watersȱ duringȱ theȱ dayȱ(MangoldȬWirz,ȱ1963;ȱKorzunȱetȱal.,ȱ1979;ȱNesis,ȱ1987).ȱ

3.1.6.3 Life history Theȱ largestȱ reportedȱ maleȱ fromȱ northernȱ Europeȱ hadȱ anȱ MLȱ ofȱ 640ȹmm,ȱ andȱ theȱ largestȱunsexedȱspecimensȱanȱMLȱofȱ750ȹmmȱ(inȱClarke,ȱ1966).ȱMoreȱrecentȱrecordsȱ indicateȱthatȱfemalesȱreachȱaȱmaximumȱMLȱofȱ520ȹmmȱ(malesȱ426ȹmm)ȱoffȱnorthernȱ Europeȱ (Lordanȱ etȱ al.,ȱ 2001a)ȱ andȱ 600ȹmmȱ (malesȱ 385ȹmm)ȱ inȱ theȱ Mediterraneanȱ (Cuccuȱetȱal.,ȱ2005).ȱHowever,ȱtheȱrecordsȱgivenȱbyȱCuccuȱetȱal.ȱ(2005)ȱrepresentȱquiteȱ unusualȱ findingsȱ forȱ Mediterraneanȱ populationsȱ (e.g.ȱ Quetglasȱ etȱ al.,ȱ 1999)ȱ andȱ forȱ Italianȱwatersȱ(e.g.ȱJerebȱandȱRagonese,ȱ1990;ȱBelcari,ȱ1999f).ȱItȱwasȱhypothesizedȱthatȱ suchȱ largeȱ sizesȱ mayȱ resultȱ fromȱ unusuallyȱ oldȱ animals.ȱ AgeȬatȬmaturityȱ hasȱ beenȱ estimatedȱatȱ15ȹmonths,ȱalthoughȱNesisȱ(1987)ȱsuggestedȱthatȱindividualsȱlargerȱthanȱ 50ȹcmȱ areȱ atȱ leastȱ 2ȱ yearsȱ old,ȱ aȱ hypothesisȱ alsoȱ proposedȱ byȱ Lordanȱ etȱ al.ȱ (2001a).ȱ Generallyȱspeaking,ȱhowever,ȱindividualsȱinȱtheȱMediterraneanȱandȱtheȱsouthernȱpartȱ ofȱtheȱdistributionalȱrangeȱmatureȱatȱmuchȱsmallerȱbodyȱsizesȱthanȱthoseȱinȱtheȱNorthȱ Atlanticȱ (HernándezȬGarcíaȱ etȱ al.,ȱ 1998a;ȱ Piatkowskiȱ etȱ al.,ȱ 1998;ȱ Arkhipkinȱ etȱ al.,ȱ 1999).ȱFemalesȱproduceȱ12ȹ000Ⱥ–Ⱥ18ȹ000ȱeggsȱ(LaptikhovskyȱandȱNigmatullin,ȱ1999).ȱ Spawningȱprobablyȱtakesȱplaceȱonȱtheȱcontinentalȱslopeȱinȱlateȱwinterȱorȱearlyȱspringȱ offȱ northernȱ Europe,ȱ inȱ Marchȱ andȱ Aprilȱ offȱ France,ȱ andȱ betweenȱ Septemberȱ andȱ Novemberȱ inȱ theȱ westernȱ Mediterraneanȱ (Piatkowskiȱ etȱ al.,ȱ 1998;ȱ Quetglasȱ etȱ al.,ȱ 1998a;ȱArkhipkinȱetȱal.,ȱ1999;ȱLordanȱetȱal.,ȱ2001a).ȱFemalesȱalwaysȱoutnumberȱmales.ȱ Ripeȱ malesȱ seemȱ toȱ beȱ presentȱ throughoutȱ theȱ year,ȱ whereasȱ matureȱ femalesȱ areȱ presentȱexclusivelyȱduringȱtheȱspawningȱperiod.ȱȱ

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3.1.6.4 Trophic ecology Predators.ȱTodarodesȱ sagittatusȱ hasȱ beenȱ foundȱ inȱ theȱ stomachsȱ ofȱ variousȱcetaceansȱ (Gonzálezȱ etȱ al.,ȱ 1994a;ȱ Clarkeȱ andȱ Pascoe,ȱ 1997;ȱ Santos,ȱ M.ȱ B.,ȱ etȱ al.,ȱ 2001b,ȱ 2007;ȱ Öztürkȱetȱal.,ȱ2007)ȱandȱlargeȱfish,ȱsuchȱasȱswordfishȱ(Xiphiasȱgladius;ȱBello,ȱ1985,ȱ1991,ȱ 1993;ȱGuerraȱetȱal.,ȱ1993;ȱSalman,ȱ2004;ȱPeristerakiȱetȱal.,ȱ2005).ȱ Prey.ȱTheȱdietȱofȱT.ȱsagittatusȱisȱcomposedȱofȱfish,ȱcrustaceans,ȱandȱcephalopods;ȱtheȱ presenceȱofȱcannibalismȱhasȱalsoȱbeenȱnoted.ȱInȱnorthernȱwaters,ȱT.ȱsagittatusȱfeedsȱ primarilyȱ onȱ smallȱ herringȱ (Clupeaȱ harengus)ȱ andȱ codȱ (Gadusȱ morhua;ȱ HernándezȬ García,ȱ1992;ȱPiatkowskiȱetȱal.,ȱ1998;ȱQuetglasȱetȱal.,ȱ1999).ȱ

3.1.6.5 Fisheries Aȱ directedȱ fisheryȱ offȱ Norwayȱ hasȱ takenȱ 10ȹ000ȹtȱ inȱ someȱ yearsȱ (e.g.ȱ Wiborg,ȱ 1972,ȱ 1978,ȱ 1979b,ȱ 1987;ȱ Besteiro,ȱ 1985;ȱ Sundet,ȱ 1985;ȱ Bjørkeȱ andȱ Gjøsæter,ȱ 2004).ȱ Inȱ theȱ Mediterranean,ȱ theȱ mainȱ fisheryȱ isȱ operatedȱ byȱ Italianȱ bottomȬtrawlȱ vessels,ȱ withȱ annualȱ catchesȱ (mainlyȱ bycatches)ȱ ofȱ ca.ȱ 3000ȹt.ȱ Inȱ otherȱ partsȱ ofȱ itsȱ distributionalȱ range,ȱitȱisȱtakenȱasȱbycatchȱinȱtrawlȱfisheriesȱ(e.g.ȱJoy,ȱ1990;ȱJónsson,ȱ1998;ȱLordanȱetȱ al.,ȱ 2001b;ȱ Gonzálezȱ andȱ Sánchez,ȱ 2002;ȱ Lefkaditouȱ etȱ al.,ȱ 2003a).ȱ Theȱ fisheryȱ isȱ concentratedȱ fromȱ Augustȱ toȱ Decemberȱ inȱ ICESȱ Divisionȱ IIa.ȱ Commonȱ fishingȱ methodsȱ includeȱ jiggingȱ offȱ Norwayȱ (Sundet,ȱ 1985)ȱ andȱ inȱ theȱ Mediterraneanȱ (RagoneseȱandȱBianchini,ȱ1990)ȱandȱpurseȬseinesȱaroundȱtheȱCanaryȱIslandsȱ(Clarke,ȱ 1966).ȱNoȱlandingsȱdataȱforȱthisȱspeciesȱaloneȱareȱreported;ȱitȱisȱgroupedȱwithȱotherȱ shortȬfinnedȱsquid.ȱItȱisȱprobableȱthatȱtwoȱseparateȱstocksȱexist,ȱoneȱinȱtheȱnorthernȱ andȱanotherȱinȱtheȱsouthwesternȱpartȱofȱtheȱrange.ȱ Theȱ speciesȱ hasȱ beenȱ identifiedȱ inȱ commercialȱ landingsȱ inȱ Norway,ȱ Ireland,ȱ Spain,ȱ Portugal,ȱItaly,ȱandȱGreece.ȱ

3.1.7 Todaropsis eblanae Ball, 1841 Commonȱ names.ȱ Lesserȱ flyingȱ squidȱ (English);ȱ toutenonȱ souffleurȱ (French);ȱ thrapsaloȱ(̋ΕΣΜ΅ΏΓ;ȱGreek);ȱtotanoȱtozzoȱ(Italian);ȱpotaȬcosteiraȱ(Portuguese);ȱpotaȱ costeraȱ(Spanish).ȱ

3.1.7.1 Adult diagnostic features Mantleȱwidthȱlargerȱthanȱ33Ⱥ%ȱofȱtheȱtotalȱML.ȱRhomboidalȱfinsȱofȱlengthȱlessȱthanȱ 50Ⱥ%ȱ ofȱ dorsalȱ ML,ȱ widthȱ 75Ⱥ–Ⱥ90Ⱥ%ȱ ofȱ dorsalȱ ML.ȱ Armsȱ withȱ twoȱ rowsȱ ofȱ suckers.ȱ Dactylusȱofȱtentacularȱclubȱwithȱfourȱlongitudinalȱrowsȱofȱsmallȱsuckers.ȱManusȱofȱ clubȱwithȱsixȱtransverseȱrowsȱofȱfourȱsuckers,ȱeachȱwithȱmedianȱpairsȱupȱtoȱfourfoldȱ largerȱ inȱ diameterȱ thanȱ lateralȱ suckers.ȱ Leftȱ andȱ rightȱ ventralȱ armsȱ (IV)ȱ ofȱ matureȱ malesȱ areȱ hectocotylized.ȱ Inverseȱ TȬshapedȱ funnelȬlockingȱ cartilage.ȱ Funnelȱ grooveȱ withoutȱanteriorȱfoveolaȱandȱwithoutȱsideȱpockets.ȱReference:ȱRoperȱetȱal.ȱ(1984).ȱ

3.1.7.2 Distribution and habitat Theȱ geographicalȱ distributionȱ isȱ discontinuous.ȱ Theȱ speciesȱ isȱ knownȱ fromȱ theȱ MediterraneanȱSea,ȱtheȱeasternȱAtlanticȱfromȱ61°Nȱtoȱ40°S,ȱtheȱsouthwesternȱPacific,ȱ andȱtheȱsouthwesternȱIndianȱOceanȱoffȱAustralia.ȱItȱisȱdemersalȱandȱassociatedȱwithȱ sandyȱandȱmuddyȱbottoms,ȱmainlyȱinȱtheȱlowerȱsublittoralȱandȱupperȱbathyalȱoverȱ theȱcontinentalȱshelf,ȱinhabitingȱdepthsȱofȱ20Ⱥ–Ⱥ780ȹmȱ(Adam,ȱ1952;ȱBelcari,ȱ1999g).ȱ

3.1.7.3 Life history Eggsȱofȱmatureȱfemalesȱareȱ0.8Ⱥ–Ⱥ2.5ȹmmȱinȱlengthȱ(MangoldȬWirz,ȱ1963;ȱHastieȱetȱal.,ȱ 1994;ȱ Laptikhovskyȱ andȱ Nigmatullin,ȱ 1999).ȱ Theȱ hatchingȱ seasonȱ extendsȱ fromȱ

20 | Cephalopod biology and fisheries in Europe

OctoberȱtoȱMarchȱinȱBritishȱwatersȱ(Hastieȱetȱal.,ȱ1994;ȱCollinsȱetȱal.,ȱ2002)ȱandȱfromȱ Marchȱ toȱ July,ȱ withȱ aȱ peakȱ inȱ May,ȱ inȱ northwesternȱ Africanȱ watersȱ (Laptikhovskyȱ andȱNigmatullin,ȱ1999).ȱGrowthȱratesȱvaryȱwithȱtheȱseasonȱofȱhatchingȱandȱareȱhigherȱ inȱ laterȬhatchedȱ individuals.ȱ Frenchȱ fisheryȱ dataȱ suggestȱ monthlyȱ growthȱ ratesȱ ofȱ 0.76ȹcmȱ (males)ȱ andȱ 1.22ȹcmȱ (females;ȱ Robinȱ etȱ al.,ȱ 2002).ȱ Maximumȱ MLȱ isȱ 290ȹmmȱ andȹ220ȹmmȱ forȱ femalesȱ andȱ males,ȱ respectivelyȱ (Robinȱ etȱ al.,ȱ 2002).ȱ Lifespanȱ isȱ approximatelyȱ1ȺȭȺ2ȹyearsȱforȱbothȱmalesȱandȱfemales.ȱ SizeȬatȬmaturityȱvariesȱwithȱgeographicȱlocationȱ(Gonzálezȱetȱal.,ȱ1994b;ȱHastieȱetȱal.,ȱ 1994;ȱBelcariȱetȱal.,ȱ1999;ȱZumholzȱandȱPiatkowski,ȱ2005),ȱwithȱanȱMLȱofȱ120Ⱥ–Ⱥ130ȹmmȱ inȱmalesȱandȱ140ȺȭȺ200ȹmmȱMLȱinȱfemales.ȱTotalȱfecundityȱinȱmatureȱfemalesȱvariesȱ fromȱ 4500ȹtoȱ 28ȹ000ȱ eggsȱ inȱ Scottishȱ watersȱ (Hastieȱ etȱ al.,ȱ 1994)ȱ andȱ fromȱ 43ȹ000ȱ toȱ 275ȹ000ȱeggsȱoffȱWestȱAfricaȱ(LaptikhovskyȱandȱNigmatullin,ȱ1999).ȱTodaropsisȱeblanaeȱ isȱanȱintermittentȱterminalȱspawner.ȱTheȱspawningȱseasonȱvariesȱgeographically.ȱȱ

3.1.7.4 Trophic ecology Predators.ȱTheȱmainȱpredatorsȱofȱT.ȱeblanaeȱareȱtoothedȱwhalesȱandȱdolphinsȱ(Clarkeȱ andȱPascoe,ȱ1985;ȱPascoe,ȱ1986;ȱBlancoȱetȱal.,ȱ2001;ȱSantos,ȱM.ȱB.,ȱetȱal.,ȱ2001a,ȱ2001b,ȱ 2007),ȱsharksȱ(ClarkeȱandȱStevens,ȱ1974),ȱandȱseveralȱteleostȱfishȱspeciesȱ(Meyerȱandȱ Smale,ȱ1991;ȱLipinskiȱetȱal.,ȱ1992).ȱ Prey.ȱTheȱdietȱisȱcomposedȱofȱfish,ȱcrustaceans,ȱandȱcephalopodsȱ(Raseroȱetȱal.,ȱ1996;ȱ Lordanȱetȱal.,ȱ1998;ȱZumholz,ȱ2000;ȱFormȱandȱOelschlägel,ȱ2004).ȱ

3.1.7.5 Fisheries Juvenilesȱ recruitȱ toȱ theȱ fisheryȱ atȱ ca.ȱ 3ȹmonthsȱ ofȱ age.ȱ Todaropsisȱ eblanaeȱ isȱ takenȱ throughoutȱ theȱ yearȱ asȱ bycatchȱ inȱ otterȬtrawlȱ fisheriesȱ and,ȱ toȱ aȱ lesserȱ extent,ȱ withȱ gillȬȱ andȱ trammelnets,ȱ longlines,ȱ andȱ jiggingȱ atȱ depthsȱ ofȱ 100Ⱥ–Ⱥ400ȹmȱ inȱ theȱ Mediterranean,ȱ offȱ Westȱ Africa,ȱ andȱ inȱ theȱ Northeastȱ Atlanticȱ (Robinȱ etȱ al.,ȱ 2002).ȱ Mostȱ catchesȱ areȱ madeȱ atȱ aȱ depthȱ ofȱ ca.ȱ 200ȹm.ȱ Asȱ forȱ otherȱ ommastrephids,ȱ inȱ general,ȱnoȱseparateȱlandingsȱdataȱareȱavailableȱforȱthisȱspecies;ȱlandingsȱareȱpooledȱ withȱthoseȱofȱotherȱommastrephidȱsquidȱspecies.ȱTodaropsisȱeblanaeȱisȱlessȱimportantȱ thanȱI.ȱcoindetiiȱinȱFrenchȱshortȬfinnedȱsquidȱlandingsȱ(Robinȱetȱal.,ȱ2002)ȱandȱmoreȱ importantȱ thanȱ I.ȱ coindetiiȱ inȱ Spanishȱ andȱ Portugueseȱ shortȬfinnedȱ squidȱ landings,ȱ althoughȱseasonalȱvariationȱoccursȱ(Bruno,ȱ2008).ȱ TheȱspeciesȱhasȱbeenȱidentifiedȱinȱcommercialȱlandingsȱinȱIreland,ȱUK,ȱFrance,ȱSpain,ȱ andȱ Portugalȱ (inȱ theȱ Northȱ Atlantic),ȱ andȱ Spain,ȱ Italy,ȱ andȱ Greeceȱ (inȱ theȱ Mediterranean).ȱ

3.1.8 Eledone moschata Lamarck, 1798 Commonȱ names.ȱ Muskyȱ octopusȱ (English);ȱ elédoneȱ musquéeȱ (French);ȱ Moschuskrakeȱ(German);ȱmoshiosȱ(̏ΓΗΛ΍ϱΖ;ȱGreek),ȱmoscardinoȱmuschiatoȱ(Italian);ȱ polvoȬmosqueadoȱ(Portuguese);ȱpulpoȱcabezónȱ(Spanish).ȱ

3.1.8.1 Adult diagnostic features Armsȱuniserial.ȱLivingȱanimalȱexudesȱaȱveryȱprominent,ȱmuskȬlikeȱodour,ȱreportedlyȱ fromȱglandsȱinȱtheȱskin.ȱLiveȱcolourȱgreyishȬbrown,ȱwithȱblackishȬbrownȱblotchesȱonȱ dorsum.ȱ Skinȱ smoothȱ toȱ veryȱ finelyȱ granulose.ȱ Noȱ ridgeȱ occursȱ aroundȱ theȱ lateralȱ peripheryȱ ofȱ theȱ mantle.ȱ Webȱ moderatelyȱ deepȹ–ȹ30Ⱥ%ȱ ofȱ armȱ length.ȱ Rightȱ armȱ IIIȱ hectocotylized,ȱlengthȱonlyȱ60ȺȭȺ70Ⱥ%ȱofȱleftȱarmȱIII.ȱLigulaȱshortȱ(3Ⱥ%ȱofȱarmȱlength);ȱ calamusȱ absent.ȱ Distalȱ tipsȱ ofȱ allȱ otherȱ armsȱ ofȱ malesȱ modifiedȱ byȱ subdivisionȱ ofȱ

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universalȱ suckersȱ intoȱ twoȱ parallelȱ rowsȱ ofȱ flattenedȱ laminaeȱ orȱ platelets.ȱ Gillȱ demibranchȱwithȱ11ȺȭȺ12ȱlamellae.ȱReference:ȱRoperȱetȱal.ȱ(1984).ȱ

3.1.8.2 Distribution and habitat Thisȱ speciesȱ isȱ distributedȱ atȱ depthsȱ ofȱ 15Ⱥ–Ⱥ200ȹmȱ inȱ Mediterraneanȱ andȱ Iberianȱ waters,ȱwithȱtheȱgreatestȱabundanceȱdownȱtoȱ100ȹmȱ(Lefkaditouȱetȱal.,ȱ1998a;ȱBelcariȱ andȱSbrana,ȱ1999;ȱBelcariȱetȱal.,ȱ2002a;ȱSilvaȱetȱal.,ȱ2004).ȱItȱisȱrelativelyȱabundantȱonȱ theȱ southernȱ Portugueseȱ coastȱ andȱ inȱ theȱ Gulfȱ ofȱ Cadiz,ȱ whereȱ itȱ extendsȱ toȱ 450ȹmȱ (Guerra,ȱ1984;ȱReisȱetȱal.,ȱ1984).ȱ

3.1.8.3 Life history Hatchlingsȱ ofȱ Eledoneȱ moschataȱ areȱ 25Ⱥ–Ⱥ30ȹmmȱ longȱ (MLȱ 10Ⱥ–Ⱥ12ȹmm),ȱ eventuallyȱ reachingȱaȱmaximumȱlengthȱofȱ150ȹmmȱinȱtheȱGulfȱofȱCadizȱandȱalongȱtheȱPortugueseȱ coastȱ andȱ 190ȹmmȱ inȱ theȱ Mediterraneanȱ (Mangold,ȱ 1983;ȱ Silvaȱ etȱ al.,ȱ 2004;ȱ E.ȱ Lefkaditou,ȱpers.ȱcomm.;ȱS.ȱLourenço,ȱpers.ȱcomm.).ȱFemalesȱreachȱmaturityȱatȱanȱMLȱ ofȱ12ȹcmȱinȱtheȱGulfȱofȱCadizȱandȱ11ȹcmȱinȱTunisianȱwatersȱ(EzzeddineȬNajai,ȱ1997;ȱ Silvaȱ etȱ al.,ȱ 2004).ȱ Malesȱ matureȱ atȱ slightlyȱ smallerȱ sizes.ȱ Estimatesȱ ofȱ fecundityȱ (oocytesȱperȱfemale)ȱincludeȱca.ȱ450ȹ±ȹ150ȱforȱtheȱGulfȱofȱCadizȱ(Silvaȱetȱal.,ȱ2004)ȱandȱ 100Ⱥ–Ⱥ500ȱforȱtheȱnorthwesternȱMediterraneanȱpopulations,ȱwhereȱeggȱlengthȱrangedȱ betweenȱ12ȱandȱ16ȹmmȱ(Mangold,ȱ1983).ȱ InȱtheȱGulfȱofȱCadiz,ȱtheȱspawningȱseasonȱextendsȱfromȱOctoberȱtoȱJulyȱ(Silvaȱetȱal.,ȱ 2004).ȱ Peakȱ spawningȱ occursȱ fromȱ Februaryȱ toȱ May,ȱ withȱ aȱ secondaryȱ peakȱ inȱ October.ȱ Theȱ southernȱ Mediterraneanȱ populationsȱ spawnȱ fromȱ Novemberȱ toȱ Julyȱ (EzzeddineȬNajai,ȱ1997).ȱInȱtheȱnorthwesternȱMediterraneanȱwaters,ȱtheȱreproductiveȱ seasonȱisȱshorter,ȱfromȱJanuaryȱtoȱMayȺ–ȺJuneȱ(Mangold,ȱ1983;ȱBelcariȱetȱal.,ȱ2002a).ȱ TheȱlifeȬcycleȱmodelȱofȱthisȱspeciesȱinȱtheȱnorthwesternȱMediterraneanȱisȱbasedȱonȱtheȱ alternationȱofȱshortȬlivedȱandȱlongȬlivedȱlifeȱcyclesȱ(Mangold,ȱ1983;ȱSilvaȱetȱal.,ȱ2004),ȱ wherebyȱpopulationsȱhatchedȱatȱdifferentȱtimesȱvaryȱinȱgrowthȱrateȱandȱrateȱofȱsexualȱ developmentȱaccordingȱtoȱprevailingȱenvironmentalȱconditions.ȱȱ

3.1.8.4 Trophic ecology Hatchlingsȱ areȱ ableȱ toȱ feedȱ onȱ liveȱ crabsȱ ofȱ theirȱ ownȱ size.ȱ Theirȱ preferenceȱ forȱ crustaceansȱ isȱ clearȱ fromȱ theȱ veryȱ earlyȱ stagesȱ onwardȱ (Boletzky,ȱ 1975).ȱ Captiveȱ studiesȱconfirmȱaȱpreferenceȱforȱcrustaceansȱoverȱmolluscsȱandȱfishȱ(jen,ȱ2007).ȱ

3.1.8.5 Fisheries Thisȱ speciesȱ isȱ caughtȱ mainlyȱ asȱ bycatchȱ ofȱ bottomȬtrawlȱ fisheries;ȱ inȱ theȱ Gulfȱ ofȱ CadizȱitȱisȱoftenȱdiscardedȱbyȱtheȱSpanishȱfleetȱbecauseȱofȱitsȱlowȱcommercialȱvalue.ȱ Inȱ theȱ mainȱ portsȱ ofȱ thisȱ region,ȱ annualȱ landingsȱ ofȱ E.ȱ moschataȱ averagedȱ ca.ȱ 100ȹtȱ duringȱ1996Ⱥ–Ⱥ2004,ȱwithȱaȱmaximumȱfromȱJanuaryȱtoȱApril.ȱInȱtheȱMediterranean,ȱthisȱ speciesȱ isȱ ofȱ considerableȱ economicȱ importance,ȱ especiallyȱ onȱ theȱ southȱ andȱ eastȱ coastsȱandȱinȱtheȱAdriaticȱSeaȱ(BelcariȱandȱSbrana,ȱ1999;ȱBelcariȱetȱal.,ȱ2002a).ȱ TheȱspeciesȱhasȱbeenȱidentifiedȱinȱcommercialȱlandingsȱinȱSpain,ȱItaly,ȱandȱGreece.ȱ

3.1.9 Eledone cirrhosa Lamarck, 1798 Commonȱ names.ȱ Hornedȱ octopus,ȱ lesserȱ octopusȱ (English);ȱ élédoneȱ commune,ȱ poulpe,ȱ poulpeȱ blancȱ (French);ȱ moshiosȱ (̏ΓΗΛ΍ϱΖ;ȱ Greek);ȱ moscardinoȱ biancoȱ (Italian);ȱpolvoȬdoȬaltoȱ(Portuguese);ȱpulpoȱblancoȱ(Spanish).ȱ

22 | Cephalopod biology and fisheries in Europe

3.1.9.1 Adult diagnostic features Suckersȱ uniserial.ȱ OrangeȬredȱ andȱ reddishȬbrownȱ dorsally;ȱ greenishȱ iridescenceȱ ventrally.ȱ Dorsalȱ sideȱ coveredȱ withȱ numerousȱ warts.ȱ Aȱ whitishȱ lineȱ encirclesȱ theȱ lateralȱ peripheryȱ ofȱ theȱ mantle.ȱ Aȱ cirrusȱ isȱ presentȱ nearȱ eachȱ eye.ȱ Rightȱ armȱ IIIȱ ofȱ malesȱisȱhectocotylized,ȱshorterȱ(69Ⱥ–Ⱥ76Ⱥ%)ȱthanȱitsȱopposite,ȱwithȱaȱveryȱshortȱligulaȱ (3Ⱥ–Ⱥ4Ⱥ%ȱ ofȱ theȱ lengthȱ ofȱ theȱ hectocotylus),ȱ calamusȱ absent;ȱ terminalȱ suckersȱ ofȱ theȱ otherȱarmsȱofȱmalesȱmodifiedȱandȱtransverselyȱcompressed.ȱDiagnosisȱadaptedȱfromȱ Roperȱetȱal.ȱ(1984).ȱ

3.1.9.2 Distribution and habitat EledoneȱcirrhosaȱisȱaȱcommonȱspeciesȱinȱtheȱNortheastȱAtlantic,ȱextendingȱfromȱ67°Nȱ toȱ theȱ northwestȱ Africanȱ coastsȱ (Guerra,ȱ 1992)ȱ andȱ throughoutȱ theȱ Mediterraneanȱ Sea.ȱItȱappearsȱoccasionallyȱinȱtheȱKattegatȱ(westȱofȱSweden;ȱHornborg,ȱ2005)ȱandȱinȱ theȱMarmaraȱSeaȱ(Unsalȱetȱal.,ȱ1999).ȱItȱisȱaȱtypicalȱsoftȬbottomȱeurybathicȱspeciesȱthatȱ livesȱtoȱaȱdepthȱofȱ770ȹmȱ(Massy,ȱ1928),ȱalthoughȱitȱisȱmainlyȱdistributedȱbetweenȱ50ȱ andȱ300ȹmȱ(BelcariȱandȱSartor,ȱ1999a;ȱBelcariȱetȱal.,ȱ2002a).ȱ

3.1.9.3 Life history Femalesȱreachȱaȱmaximumȱsizeȱofȱ190ȹmmȱMLȱcomparedȱwithȱ135ȹmmȱMLȱinȱmales.ȱ TheyȱattainȱmaturityȱatȱsmallerȱsizesȱinȱtheȱMediterraneanȱthanȱinȱtheȱAtlantic,ȱandȱ malesȱ matureȱ atȱ smallerȱ sizesȱ thanȱ femalesȱ (Belcariȱ andȱ Sartor,ȱ 1999a;ȱ Belcariȱ etȱ al.ȱ 2002a;ȱA.ȱMoreno,ȱpers.ȱcomm.).ȱ Aȱcombinationȱofȱaȱ1Ȭȱandȱ2Ȭyearȱlifeȱcycleȱ(fastȬgrowing,ȱearlyȬmaturingȱanimalsȱvs.ȱ slowerȬgrowing,ȱ lateȬmaturingȱ animals)ȱ isȱ proposedȱ forȱ theȱ Northȱ Seaȱ (Boyleȱ andȱ Knobloch,ȱ1982;ȱBoyle,ȱ1983;ȱBoyleȱetȱal.,ȱ1988).ȱInȱtheȱMediterranean,ȱtheȱlifespanȱofȱ mostȱindividualsȱisȱestimatedȱtoȱbeȱapproximatelyȱ2ȱyearsȱ(e.g.ȱMoriyasu,ȱ1981,ȱ1983;ȱ Belcariȱetȱal.,ȱ1990,ȱ2002a;ȱSánchezȱetȱal.,ȱ2004),ȱwithȱlessȱthanȱ10Ⱥ%ȱreachingȱmaturityȱatȱ ca.ȱ3ȹyearsȱoldȱ(LefkaditouȱandȱPapaconstantinou,ȱ1995;ȱCuccuȱetȱal.,ȱ2003;ȱOrsiȬReliniȱ etȱ al.,ȱ 2006).ȱ Note,ȱ however,ȱ thatȱ theseȱ conclusionsȱ areȱ basedȱ onȱ lengthȱ frequencyȱ analysis,ȱ theȱ validityȱ ofȱ whichȱ hasȱ beenȱ calledȱ intoȱ questionȱ (E.ȱ Lefkaditou,ȱ pers.ȱ comm.)ȱ Fecundityȱ estimatesȱ varyȱ widely:ȱ ~Ⱥ9000ȱ eggsȱ inȱ theȱ Northȱ Sea,ȱ ~Ⱥ5500ȱ eggsȱ inȱ theȱ CatalanȱSeaȱ(Boyleȱetȱal.,ȱ1988),ȱandȱ~Ⱥ2000ȱeggsȱinȱtheȱTyrrhenianȱSeaȱ(Rossetti,ȱ1998).ȱ E.ȱ cirrhosaȱ spawnsȱ inȱ summerȺȭȺautumnȱ inȱ theȱ Northȱ Seaȱ (Boyle,ȱ 1983;ȱ Boyleȱ andȱ Knobloch,ȱ1983)ȱandȱinȱspringȺȭȺsummerȱfartherȱsouthȱ(A.ȱMoreno,ȱpers.ȱcomm.).ȱInȱ theȱ Mediterranean,ȱ theȱ spawningȱ seasonȱ isȱ earlierȱ inȱ theȱ westernȱ basinȱ (springȺȭȺsummer)ȱ thanȱ inȱ theȱ easternȱ basinȱ (summerȺȭȺautumn;ȱ Belcariȱ andȱ Sartor,ȱ 1999a;ȱLefkaditouȱetȱal.,ȱ2000).ȱ

3.1.9.4 Trophic ecology Predators.ȱWhales,ȱseals,ȱandȱfishȱareȱconsideredȱtoȱbeȱtheȱmostȱimportantȱpredatorsȱ ofȱ E.ȱ cirrhosaȱ (e.g.ȱ Pierceȱ etȱ al.,ȱ 1991;ȱ Tollitȱ andȱ Thompson,ȱ 1996;ȱ Santosȱ etȱ al.,ȱ 1999;ȱ Brownȱetȱal.,ȱ2001;ȱDalyȱetȱal.,ȱ2001;ȱVelascoȱetȱal.,ȱ2001).ȱȱ Prey.ȱEledoneȱcirrhosaȱisȱaȱcarnivorousȱspeciesȱandȱactiveȱpredator.ȱTheȱdietȱisȱmainlyȱ composedȱ ofȱ decapodȱ crustaceans,ȱ mostlyȱ alpheidsȱ andȱ brachyuridsȱ (Boyleȱ andȱ Knobloch,ȱ1981;ȱSánchez,ȱ1981;ȱAuteriȱetȱal.,ȱ1988).ȱTheȱspeciesȱalsoȱfeedsȱonȱmolluscsȱ andȱcephalopodȱeggs,ȱandȱcannibalismȱhasȱbeenȱobservedȱ(Moriyasu,ȱ1981;ȱGuerra,ȱ 1992).ȱ

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3.1.9.5 Fisheries EledoneȱcirrhosaȱhasȱgreatȱcommercialȱvalueȱinȱtheȱwesternȱMediterraneanȱandȱusuallyȱ lessȱthanȱ10Ⱥ%ȱofȱcatchesȱisȱdiscardedȱ(MangoldȱandȱBoletzky,ȱ1987;ȱSartorȱetȱal.,ȱ1998;ȱ Reliniȱetȱal.,ȱ1998).ȱAcrossȱEurope,ȱlandingsȱcomeȱalmostȱentirelyȱfromȱbottomȬtrawlȱ fisheries,ȱandȱE.ȱcirrhosaȱappearsȱtogetherȱwithȱE.ȱmoschataȱandȱOctopusȱvulgarisȱinȱtheȱ catchȱ statistics.ȱ Itȱ isȱ marketedȱ inȱ twoȱ distinctȱ commercialȱ categoriesȱ inȱ theȱ Mediterranean,ȱ whereȱ smallȱ specimensȱ (<Ⱥ50ȹmmȱ ML)ȱ haveȱ higherȱ economicȱ valueȱ (Belcariȱetȱal.,ȱ1998;ȱBelcariȱandȱSartor,ȱ1999).ȱInȱsomeȱregions,ȱtheseȱsmallȱoctopusȱareȱ aȱtargetȱofȱtheȱmultispeciesȱtrawlȱfisheryȱinȱspringȱandȱsummerȱthatȱcoincidesȱwithȱ theȱrecruitmentȱperiodȱofȱtheȱspeciesȱ(ReliniȱandȱOrsiȬRelini,ȱ1984;ȱBelcariȱetȱal.,ȱ1998,ȱ 2002a;ȱBelcariȱandȱSartor,ȱ1999;ȱSánchezȱetȱal.,ȱ2004).ȱRecently,ȱthisȱfisheryȱwasȱbannedȱ inȱ theȱ Catalanȱ Seaȱ (P.ȱ Sánchez,ȱ pers.ȱ comm.).ȱ Eledoneȱ cirrhosaȱ isȱ theȱ onlyȱ Eledoneȱ speciesȱpresentȱinȱnorthernȱSpanishȱwaters,ȱwhereȱitȱisȱlandedȱbyȱtheȱbottomȬtrawlȱ fisheryȱ(ca.ȱ700ȹtȱyearȺƺ1ȱonȱaverage,ȱ1998Ⱥ–Ⱥ2006)ȱasȱbycatch,ȱalthoughȱitsȱcommercialȱ valueȱisȱveryȱlow.ȱItȱisȱoccasionallyȱlandedȱinȱsmallȱquantitiesȱinȱScotland,ȱbutȱthereȱ appearsȱtoȱbeȱlittleȱcommercialȱinterestȱinȱtheȱspeciesȱ(G.ȱPierce,ȱpers.ȱcomm.).ȱ TheȱspeciesȱhasȱbeenȱidentifiedȱinȱcommercialȱlandingsȱinȱPortugal,ȱSpain,ȱItaly,ȱandȱ Greece.ȱ

3.1.10 Octopus vulgaris Cuvier, 1797 Commonȱ names.ȱ Commonȱ octopusȱ (English);ȱ pieuvre,ȱ poulpeȱ (French);ȱ chtapodiȱ (ΛΘ΅ΔϱΈ΍;ȱGreek);ȱGemeinerȱKrakeȱ(German);ȱpolpoȱcomuneȱ(Italian);ȱpulpoȱcomúnȱ (Spanish);ȱolagarroȱ(Spanish,ȱBasque);ȱpopȱroquerȱ(Spanish,ȱCatalan);ȱpolboȱ(Spanish,ȱ Galician);ȱpolvoȬvulgarȱ(Portuguese).ȱ

3.1.10.1 Adult diagnostic features Suckersȱbiserial.ȱMLȱtoȱ40ȹcm;ȱtotalȱlengthȱtoȱ140ȹcm.ȱArmsȱrobustȱatȱbase,ȱlateralȱarmsȱ longest,ȱ dorsalȱ armsȱ shortest.ȱ Suckersȱ 15Ⱥ–Ⱥ17Ⱥȱ ofȱ armsȱ IIȱ andȱ IIIȱ enlargedȱ inȱ adults,ȱ especiallyȱ males.ȱ Rightȱ armȱ IIIȱ ofȱ matureȱ malesȱ hectocotylized;ȱ ligulaȱ shortȱ andȱ spoonȬshaped.ȱGillȱdemibranchȱwithȱ7ȺȭȺ11ȱlamellae,ȱincludingȱterminalȱlamellae.ȱFourȱ papillaeȱ inȱ theȱ dorsalȱ partȱ ofȱ theȱ mantleȱ (oneȱ situatedȱ inȱ theȱ anteriorȱ part,ȱ anotherȱ posteriorȱandȱtwoȱlaterals).ȱReticulatedȱskinȱwithȱfourȱwhitishȱspots,ȱtwoȱbetweenȱtheȱ eyesȱandȱtwoȱbelowȱtheȱfirstȱdorsalȱpapilla.ȱDiagnosisȱadaptedȱfromȱGuerraȱ(1992).ȱ

3.1.10.2 Distribution and habitat Octopusȱ vulgarisȱ isȱ especiallyȱ abundantȱ inȱ theȱ Mediterraneanȱ Seaȱ andȱ theȱ easternȱ Atlanticȱ (Belcariȱ andȱ Sartor,ȱ 1999b).ȱ Molecularȱ geneticȱ workȱ hasȱ demonstratedȱ thatȱ theȱdistributionȱofȱO.ȱvulgarisȱinȱtheȱAtlanticȱextendsȱtoȱsouthernȱBrazilȱ(Söllerȱetȱal.,ȱ 2000)ȱinȱtheȱwest,ȱtoȱLanzaroteȱandȱSenegalȱinȱtheȱeast,ȱandȱasȱfarȱsouthȱasȱTristanȱdeȱ Cunhaȱ andȱ Falseȱ Bay,ȱ Southȱ Africaȱ (Warnkeȱ etȱ al.,ȱ 2004).ȱ Samplesȱ fromȱ Japanȱ andȱ TaiwanȱinȱtheȱPacificȱalsoȱappearȱtoȱbeȱconspecificȱwithȱO.ȱvulgaris.ȱAȱbenthicȱspecies,ȱ O.ȱvulgarisȱinhabitsȱtheȱcoastlineȱtoȱtheȱouterȱedgeȱofȱtheȱcontinentalȱshelfȱ(200ȹm).ȱItȱ undertakesȱlimitedȱseasonalȱmigrations.ȱ

3.1.10.3 Life history Octopusȱvulgarisȱhasȱaȱlifeȱcycleȱofȱ12Ⱥ–Ⱥ14ȹmonthsȱ(Domainȱetȱal.,ȱ2002;ȱIglesiasȱetȱal.,ȱ 2004)ȱandȱterminalȱspawningȱwithȱeggȱcareȱbyȱtheȱfemale.ȱTheȱincubationȱperiodȱofȱ theȱ eggsȱ isȱ 22Ⱥ–Ⱥ25ȹdaysȱ atȱ 25Ⱥ°Cȱ (Mangold,ȱ 1997),ȱ butȱ longerȱ atȱ lowerȱ temperaturesȱ (36ȹdaysȱ atȱ 23Ⱥ°C,ȱ 60ȹdaysȱ atȱ 21Ⱥ°C,ȱ 80ȹdaysȱ atȱ 17Ⱥ°C,ȱ andȱ 120ȹdaysȱ atȱ 13Ⱥ°C;ȱ Mangoldȱ andȱBoletzky,ȱ1973;ȱCaverivièreȱetȱal.,ȱ1999;ȱMartins,ȱ2003).ȱParalarvaeȱareȱplanktonicȱ

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forȱ1Ⱥ–Ⱥ3ȹmonths,ȱdependingȱonȱtheȱeffectȱofȱtemperatureȱonȱgrowthȱrate,ȱandȱadoptȱ theȱbenthicȱlifeȱmodeȱofȱtheȱadultsȱatȱca.ȱ7.5ȹmmȱMLȱ(Villanueva,ȱ1995).ȱTheȱmortalityȱ onȱtheȱparalarvaeȱphaseȱisȱthoughtȱtoȱbeȱveryȱhighȱandȱdependentȱonȱenvironmentalȱ conditions,ȱwithȱhighlyȱvariableȱrecruitmentȱsuccessȱ(Faure,ȱ2002).ȱ Growthȱ isȱ veryȱ rapid,ȱ andȱ largeȱ individualȱ variationȱ inȱ growthȱ ratesȱ hasȱ beenȱ observedȱ bothȱ inȱ cultureȱexperimentsȱ(Iglesiasȱ etȱ al.,ȱ 2004)ȱ andȱ inȱ wildȱ populationsȱ (Domainȱ etȱ al.,ȱ2000).ȱ Juvenilesȱ canȱ reachȱ 0.5Ⱥ–Ⱥ0.6ȹkgȱ withinȱsixȱmonthsȱ ofȱ hatching,ȱ andȱ 1.4Ⱥ–Ⱥ1.8ȹkgȱ withinȱ eightȱ months,ȱ atȱ aȱ meanȱ temperatureȱ ofȱ 18Ⱥ°Cȱ (Iglesiasȱ etȱ al.,ȱ 2004).ȱMaximumȱsizeȱisȱ400ȹmmȱML.ȱȱ Theȱpotentialȱfecundityȱofȱmatureȱfemalesȱrangesȱfromȱ100ȹ000ȱtoȱ500ȹ000ȱoocytes.ȱTheȱ eggsȱ areȱ small,ȱ approximatelyȱ 2.5ȹmmȱ longȱ (Mangold,ȱ 1997).ȱ Theȱ spawningȱ seasonȱ extendsȱthroughoutȱtheȱyear,ȱwithȱtwoȱpeaksȱinȱAtlanticȱpopulationsȱwithinȱtheȱmainȱ upwellingȱregions:ȱwinterȱandȱsummerȱinȱwesternȱPortugueseȱwatersȱ(Moreno,ȱ2008),ȱ andȱ springȱ andȱ autumnȱ inȱ Moroccanȱ watersȱ (Farajȱ andȱ Bez,ȱ 2007)ȱ andȱ theȱ Canaryȱ Islandsȱ (HernándezȬGarcíaȱ etȱ al.,ȱ 2002).ȱ Aȱ singleȱ spawningȱ peakȱ occursȱ inȱ lateȱ winter/springȱinȱnorthwestȱSpainȱ(FernándezȬRuedaȱandȱGarcíaȬFlórez,ȱ2007;ȱOteroȱetȱ al.,ȱ2007)ȱandȱinȱsummerȱinȱtheȱnorthernȱGulfȱofȱCadizȱ(RodríguezȬRúaȱetȱal.,ȱ2005;ȱ Moreno,ȱ 2008)ȱ andȱ theȱ Mediterraneanȱ (Sánchezȱ andȱ Obarti,ȱ 1993;ȱ Mangold,ȱ 1997;ȱ Belcariȱetȱal.,ȱ2002b).ȱȱ

3.1.10.4 Trophic ecology Predators.ȱFish,ȱmarineȱmammals,ȱbirds,ȱandȱotherȱcephalopodsȱpreyȱonȱO.ȱvulgarisȱ (Hanlonȱ andȱ Messenger,ȱ 1998).ȱ Marineȱ mammalsȱ includeȱ commonȱ dolphinȱ (Delphinusȱ delphis;ȱ López,ȱ 2002;ȱ Santosȱ etȱ al.,ȱ 2004b),ȱ bottlenoseȱ dolphinȱ (Tursiopsȱ truncatus;ȱ Blancoȱ etȱ al.,ȱ 2001),ȱ Risso’sȱ dolphinȱ (Grampusȱ griseus),ȱ longȬfinnedȱ pilotȱ whaleȱ (Globicephalaȱ melas;ȱ López,ȱ 2002),ȱ andȱ Mediterraneanȱ monkȱ sealȱ (Monachusȱ monachus).ȱFishȱpredatorsȱofȱadultsȱandȱjuvenilesȱincludeȱcongerȱeelȱ(Congerȱconger)ȱ andȱ Mediterraneanȱ morayȱ eelȱ (Muraenaȱ helena),ȱ whereasȱ Mediterraneanȱ duskyȱ grouperȱ (Epinephelusȱ marginatus),ȱ serranidȱ fishȱ (Serranusȱ sp.)ȱ andȱ theȱ sandȱ smeltȱ (Atherinaȱpresbyter)ȱpreyȱonȱhatchlingsȱ(VillanuevaȱandȱNorman,ȱ2008).ȱ Prey.ȱParalarvaeȱfeedȱmainlyȱonȱdecapodȱcrustaceanȱlarvaeȱ(VillanuevaȱandȱNorman,ȱ 2008).ȱ Dietȱ ofȱ juvenilesȱ andȱ adultsȱ mayȱ includeȱ crustaceans,ȱ teleostȱ fish,ȱ otherȱ cephalopods,ȱandȱpolychaetes.ȱOnȱtheȱMediterraneanȱcoastȱofȱSpainȱinȱCataluña,ȱ80Ⱥ%ȱ ofȱtheȱdietȱcomprisesȱcrustaceansȱ(Guerra,ȱ1978;ȱSánchezȱandȱObarti,ȱ1993;ȱQuetglasȱetȱ al.,ȱ 1998b),ȱ butȱ inȱ theȱ Algarveȱ (Portugal),ȱ aȱ similarȱ proportionȱ comprisesȱ bivalvesȱ (Rosaȱetȱal.,ȱ2004).ȱȱ

3.1.10.5 Fisheries Octopusȱvulgarisȱisȱtakenȱthroughoutȱtheȱyearȱasȱaȱtargetȱspeciesȱinȱbottomȱandȱpelagicȱ trawlsȱandȱviaȱhandȬjigs,ȱpots,ȱtrammelnets,ȱandȱtrapsȱinȱsmallȬscaleȱcoastalȱfisheriesȱ atȱdepthsȱofȱ20ȺȭȺ200ȹmȱinȱtheȱMediterraneanȱ(Belcariȱetȱal.,ȱ2002b),ȱoffȱWestȱAfrica,ȱandȱ inȱ theȱ Northeastȱ Atlantic.ȱ Althoughȱ someȱ countriesȱ reportȱ landingsȱ ofȱ thisȱ speciesȱ separately,ȱothersȱgroupȱallȱlandingsȱofȱOctopodidaeȱtogether.ȱInȱGaliciaȱ(northwestȱ Spain),ȱwhereȱannualȱlandingsȱforȱ1998Ⱥ–Ⱥ2006ȱwereȱca.ȱ1500ȹt,ȱtheȱspeciesȱisȱtargetedȱ byȱtheȱartisanalȱfleetȱusingȱtrapsȱ(calledȱ“nasaȱdeȱpolbo”),ȱwhichȱareȱfairlyȱspecificȱtoȱ octopusȱ (whichȱ formsȱ 80Ⱥ–Ⱥ90Ⱥ%ȱ ofȱ theȱ totalȱ catchesȱ inȱ thisȱ gear);ȱitȱ isȱ alsoȱ caughtȱ inȱ trapsȱ setȱ forȱ otherȱ species,ȱ inȱ trammelnetsȱ (“trasmallos”ȱ andȱ “miños”),ȱ andȱ onȱ hookedȱlinesȱ(“raña”;ȱBañónȱetȱal.,ȱ2007).ȱInȱtheȱGulfȱofȱCadiz,ȱbothȱbottomȱtrawlersȱ andȱartisanalȱvesselsȱtakeȱthisȱspecies,ȱandȱannualȱlandingsȱhaveȱfluctuatedȱwidely,ȱ fromȱ500ȹtȱtoȱmoreȱthanȱ2000ȹtȱinȱ1994Ⱥ–Ⱥ2006.ȱ

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TheȱspeciesȱhasȱbeenȱidentifiedȱinȱcommercialȱlandingsȱinȱSpain,ȱPortugal,ȱItaly,ȱandȱ Greece.ȱ

3.1.11 Sepia officinalis Linnaeus, 1758 Commonȱnames.ȱCommonȱcuttlefishȱ(English);ȱseicheȱcommuneȱ(French);ȱGemeinerȱ Tintenfischȱ(German);ȱsoupiaȱ(̕ΓΙΔ΍Σ;ȱGreek);ȱseppiaȱcomuneȱ(Italian);ȱsepiaȱcomúnȱ (Spanish);ȱchocoȬvulgarȱ(Portuguese).ȱȱ

3.1.11.1 Adult diagnostic features Upȱ toȱ 450ȹmmȱ ML.ȱ Mantleȱ oval.ȱ Tentacularȱ clubȱ withȱ fiveȱ orȱ sixȱ suckersȱ inȱ eachȱ transverseȱrow,ȱtheȱmedialȱoneȱmoderatelyȱenlarged;ȱswimmingȱkeelȱnotȱextendingȱ proximallyȱbeyondȱtheȱbaseȱofȱtheȱclub.ȱHectocotylusȱonȱleftȱarmȱIVȱofȱmales:ȱafterȱ theȱ5Ⱥ–7ȱproximalȱsuckerȱrows,ȱthereȱareȱfromȱ4Ⱥ–Ⱥ5ȱtoȱ8Ⱥ–Ⱥ9ȱ(medial)ȱhorizontalȱrowsȱofȱ reducedȱsuckers.ȱCuttleboneȱposteriorlyȱroundedȱwithȱparallelȱsides.ȱAnteriorȱstriaeȱ areȱinvertedȱUȬshape,ȱorȱshallowȱMȬshape.ȱColourȱlightȱbrown.ȱHeadȱwithȱscatteredȱ whiteȱ spotsȱ andȱ withȱ darkȱ pigmentȱ aroundȱ eyeȱ orbits.ȱ Armsȱ IȺ–ȺIIIȱ haveȱ aȱ broad,ȱ longitudinalȱbrownishȱbandȱmedially,ȱextendingȱontoȱhead.ȱDorsalȱmantleȱhasȱboldȱ transverseȱ zebraȬstripeȱ patternȱ duringȱ theȱ breedingȱseason;ȱ pairedȱ dorsalȱ eyeȱspotsȱ absent.ȱFinsȱwithȱnarrowȱwhiteȱbandȱalongȱouterȱmarginȱandȱwithȱsmallȱwhiteȱspots,ȱ becomingȱ largerȱ towardȱ junctionȱ ofȱ mantleȱ andȱ fins.ȱ Matureȱ malesȱ withȱ armsȱ IVȱ emboldenedȱbyȱwhiteȱandȱblackȱzebraȱbandsȱandȱwhiteȱarmȱspots.ȱDiagnosisȱadaptedȱ fromȱRoperȱetȱal.ȱ(1984),ȱGuerraȱ(1992),ȱandȱReidȱetȱal.ȱ(2005).ȱ

3.1.11.2 Distribution and habitat SepiaȱofficinalisȱextendsȱthroughȱtheȱeasternȱAtlanticȱandȱMediterraneanȱSeaȱ(Belcari,ȱ 1999i).ȱ Inȱ theȱ easternȱ Northȱ Atlantic,ȱ itȱ extendsȱ fromȱ theȱ Shetlandȱ Islandsȱ andȱ southernȱNorwayȱ(notȱpresentȱinȱtheȱBalticȱSea,ȱexceptȱforȱoccasionalȱincursionsȱwithȱ theȱ northeasternmostȱ Atlanticȱ waters),ȱ southȱ throughȱ theȱ Mediterraneanȱ Seaȱ (includingȱ Aegeanȱ Sea,ȱ Seaȱ ofȱ Marmara,ȱ andȱ Levantineȱ Sea),ȱ toȱ northwestȱ Africa,ȱ withȱ theȱ southernȱ boundaryȱ coincidingȱ approximatelyȱ withȱ theȱ borderȱ betweenȱ Mauritaniaȱ andȱ Senegalȱ (16°N;ȱ Reidȱ etȱ al.,ȱ 2005).ȱ Seasonalȱ migrationsȱ occurȱ fromȱ deeper,ȱoffshoreȱoverwinteringȱgroundsȱtoȱshallower,ȱcoastalȱspawningȱandȱnurseryȱ grounds.ȱȱ

3.1.11.3 Life history Hatchlingȱ MLȱ rangesȱ fromȱ 6ȱ toȱ 9ȹmmȱ (Boletzky,ȱ 1983).ȱ Growthȱ ratesȱ varyȱ directlyȱ withȱtemperatureȱandȱinverselyȱwithȱsize.ȱInȱtheȱEnglishȱChannel,ȱstatolithȱanalysisȱ indicatesȱ thatȱ theȱ growthȱ patternȱ isȱ exponentialȱ inȱ hatchlingsȱ andȱ earlyȱ stages,ȱ whereasȱlogisticȱcurvesȱareȱaȱbetterȱfitȱforȱlargerȱsizeȱclassesȱ(7–106ȹmmȱML;ȱChallierȱ etȱal.,ȱ2005a).ȱȱ Commonȱ cuttlefishȱ attainȱ sexualȱ maturityȱ atȱ aȱ wideȱ rangeȱ ofȱ sizes.ȱ Inȱ theȱ Mediterraneanȱ Sea,ȱ matureȱ malesȱ ofȱ 6Ⱥ–Ⱥ8ȹcmȱ MLȱ wereȱ observed.ȱ Malesȱ overȱ 10ȹcmȱ ML,ȱhowever,ȱmayȱstillȱbeȱimmature.ȱAȱsimilarȱsituationȱexistsȱinȱfemales.ȱLengthȱatȱ firstȱmaturityȱisȱca.ȱ13ȹcmȱMLȱinȱfemales.ȱFemalesȱproduceȱ150Ⱥ–Ⱥ4000ȱeggs,ȱdependingȱ onȱtheirȱsize.ȱTheȱmainȱspawningȱseasonȱcoversȱspringȱandȱsummer,ȱbutȱthereȱisȱalsoȱ winterȱspawning,ȱwhichȱisȱespeciallyȱpronouncedȱonȱtheȱAtlanticȱcoast.ȱTheȱlengthȱofȱ timeȱspentȱunderȱoptimalȱconditionsȱinȱtheȱearlyȱjuvenileȱphaseȱ(inshoreȱspringȱandȱ summerȱ conditions)ȱ determinesȱ whetherȱ anȱ individualȱ becomesȱ sexuallyȱ matureȱ duringȱtheȱfirstȱwinterȱand,ȱhence,ȱdeterminesȱtheȱlifespan,ȱwhichȱmayȱvaryȱfromȱca.ȱ 14ȱ toȱ 24ȹmonths.ȱ Theȱ speciesȱ hasȱ anȱ intermittentȱ terminalȱ spawningȱ strategyȱ

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(Boletzky,ȱ 1983;ȱ Guerraȱ andȱ Castro,ȱ 1988;ȱ Boucaudȱ andȱ Daguzan,ȱ 1990;ȱ BoucaudȬ Camouȱetȱal.,ȱ1991;ȱGauvritȱetȱal.,ȱ1997;ȱBelcari,ȱ1999h;ȱDunn,ȱ1999;ȱRochaȱetȱal.,ȱ2001).ȱ

3.1.11.4 Trophic ecology Predators.ȱ Sepiaȱ officinalisȱ hasȱ beenȱ foundȱ inȱ theȱ stomachsȱ ofȱ marineȱ mammals,ȱ includingȱ Risso’sȱ dolphinȱ (Grampusȱ griseus;ȱ Clarkeȱ andȱ Pascoe,ȱ 1985),ȱ theȱ Mediterraneanȱ monkȱ sealȱ (Monachusȱ monachus;ȱ Salmanȱ etȱ al.,ȱ 2001),ȱ andȱ (inȱ Southȱ Africa)ȱ furȱ sealsȱ (Arctocephalusȱ pusillus;ȱ Castleyȱ etȱ al.,ȱ 1991).ȱ Itȱ isȱ alsoȱ eatenȱ byȱ elasmobranchs,ȱincludingȱtheȱblueȱsharkȱ(Prionaceȱglauca;ȱClarkeȱandȱStevens,ȱ1974)ȱ andȱsmoothȱhoundȱ(Mustelusȱmustelus;ȱMorteȱetȱal.,ȱ1997).ȱȱ Prey.ȱ Theȱ dietȱ mainlyȱ consistsȱ ofȱ smallȱ crabs,ȱshrimps,ȱ demersalȱfish,ȱ cephalopods,ȱ andȱpolychaetes.ȱAȱsignificantȱchangeȱfromȱcrustaceansȱtoȱfishȱoccursȱwithȱgrowth.ȱ Cannibalismȱisȱrelativelyȱcommonȱatȱallȱsizes.ȱȱ

3.1.11.5 Fisheries Sepiaȱofficinalisȱisȱanȱimportantȱspeciesȱforȱtheȱcommercialȱfisheriesȱofȱmanyȱcountries.ȱ TheȱmainȱproducerȱcountriesȱinȱtheȱlastȱdecadeȱwereȱFranceȱinȱtheȱNortheastȱAtlantic,ȱ followedȱ byȱ Italyȱ andȱ Tunisiaȱ inȱ theȱ Mediterraneanȱ Seaȱ (Belcariȱ etȱ al.,ȱ 2002c).ȱ Inȱ northernȱfisheriesȱ(EnglishȱChannel,ȱBayȱofȱBiscay),ȱcuttlefishȱareȱprimarilyȱcaughtȱbyȱ otterȱ andȱ beamȱ trawlers,ȱeitherȱ asȱaȱ targetȱ speciesȱ orȱ asȱ bycatchȱwhenȱ theȱ targetȱ isȱ demersalȱ finfish.ȱ Inȱ southernȱ fisheries,ȱ artisanalȱ gearsȱ dominate,ȱ andȱ cuttlefishȱ areȱ caughtȱbyȱgillnets,ȱtrammelnets,ȱandȱaȱgreatȱvarietyȱofȱhighlyȱselectiveȱgears,ȱsuchȱasȱ traps,ȱ lures,ȱ jigs,ȱ andȱ spears.ȱ Generally,ȱ trawlersȱ operateȱ inȱ inshoreȱ andȱ offshoreȱ fishingȱ grounds,ȱ takingȱ bothȱ juvenileȱandȱ adultȱ specimens,ȱ whereasȱartisanalȱgearsȱ (e.g.ȱ traps)ȱ catchȱ spawningȱ animalsȱ mainlyȱ inshore.ȱ Inȱ Galicia,ȱ cuttlefishȱ isȱ mainlyȱ caughtȱwithȱtrammelnetsȱfromȱJanuaryȱtoȱMayȱ(duringȱtheȱmainȱreproductiveȱperiod)ȱ onȱinshoreȱfishingȱgroundsȱ(Arnáizȱetȱal.,ȱ2001).ȱInȱtheȱinnerȱareaȱofȱtheȱrías,ȱcuttlefishȱ isȱ alsoȱ caughtȱ withȱ fykenetsȱ (“butróns”)ȱ andȱ trapsȱ (“nasaȱ deȱ choco”).ȱ Managementȱ optionsȱofȱinterestȱforȱthisȱresourceȱincludeȱeffortȱregulationsȱ(licencesȱareȱissuedȱinȱ inshoreȱartisanalȱfisheries)ȱandȱminimumȱlandingȱsize.ȱCatchesȱofȱthisȱspeciesȱareȱnotȱ recordedȱ separatelyȱ byȱ allȱ countries,ȱ oftenȱ beingȱ groupedȱ withȱ otherȱ cuttlefish,ȱ althoughȱ FAOȱ recordsȱ separateȱ catchȱ statisticsȱ forȱ thisȱ speciesȱ alongȱ theȱ Spanishȱ Atlanticȱ coast,ȱ andȱ separateȱ statisticsȱ areȱ probablyȱ alsoȱ availableȱ forȱ theȱ Frenchȱ Atlanticȱcoast.ȱ Theȱ speciesȱ hasȱ beenȱ identifiedȱ inȱ commercialȱ landingsȱ inȱ theȱ UK,ȱ France,ȱ Spain,ȱ Portugal,ȱItaly,ȱandȱGreece.ȱSepiaȱelegansȱBlainville,ȱ1827ȱ

3.1.12 Sepia elegans Blainville, 1827 Commonȱ names.ȱ Elegantȱ cuttlefishȱ (English);ȱ seicheȱ eleganteȱ (French);ȱ seppiaȱ eleganteȱ(Italian);ȱchocoȬeleganteȱ(Portuguese);ȱchoquitoȱsinȱpuntaȱ(Spanish).ȱȱ

3.1.12.1 Adult diagnostic features Upȱtoȱ89ȹmmȱMLȱ(Adam,ȱ1952).ȱMantleȱoblong,ȱlengthȱmoreȱthanȱtwiceȱwidth.ȱClubȱ short,ȱ oval;ȱ suckerȬbearingȱ surfaceȱ flattened,ȱ withȱ 6Ⱥ–Ⱥ8ȱ suckersȱ inȱ transverseȱ rows;ȱ suckersȱdifferȱmarkedlyȱinȱsize:ȱ3Ⱥ–Ⱥ4ȱgreatlyȱenlargedȱsuckersȱtowardȱposteriorȱendȱofȱ clubȱ andȱ severalȱ dorsalȱ suckersȱ enlarged,ȱ butȱ notȱ asȱ largeȱ asȱ medialȱ suckers.ȱ Hectocotylusȱpresentȱonȱleftȱventralȱarmȱofȱmales:ȱ1Ⱥ–Ⱥ2ȱrowsȱofȱnormalȱsizeȱsuckersȱ proximally,ȱ9Ⱥ–Ⱥ11ȱrowsȱofȱreducedȱminuteȱsuckersȱmedially,ȱthenȱnormalȱsizeȱsuckersȱ toȱ armȱ tip;ȱ suckersȱ inȱ twoȱ dorsalȱ andȱ twoȱ ventralȱ seriesȱ displacedȱ laterally.ȱ Cuttleboneȱ outlineȱ oblong.ȱ Anteriorȱ striaeȱ areȱ invertedȱ UȬshape.ȱ Spineȱ absent,ȱ butȱ

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ridgeȱandȱsmallȱlateralȱwingsȱpresent.ȱDorsoȬposteriorȱendȱofȱcuttleboneȱwithȱshort,ȱ rugose,ȱ calcareousȱ keel.ȱ Colourȱ reddishȱ brown.ȱ Dorsalȱ mantleȱ pale,ȱ pepperedȱ withȱ scatteredȱpurpleȬblackȱchromatophores.ȱFinsȱandȱventralȱmantleȱpale.ȱReference:ȱReidȱ etȱal.ȱ(2005).ȱ

3.1.12.2 Distribution and habitat SepiaȱelegansȱextendsȱthroughȱtheȱeasternȱAtlanticȱfromȱwesternȱScotlandȱinȱtheȱnorthȱ toȱ Namibiaȱ (21qS)ȱ inȱ theȱ southȱ andȱ isȱ presentȱ throughoutȱ theȱ Mediterraneanȱ Seaȱ (JerebȱandȱRoper,ȱ2005).ȱItȱisȱaȱsublittoralȱspecies,ȱlivingȱonȱsandyȱandȱsandyȺ–Ⱥmuddyȱ bottomsȱatȱdepthsȱtoȱca.ȱ450ȹmȱ(JerebȱandȱRagonese,ȱ1991).ȱTheȱspeciesȱisȱtolerantȱofȱ fluctuationsȱinȱsalinityȱandȱhasȱbeenȱfoundȱinȱbrackishȱwatersȱ(salinityȱ18Ⱥ–Ⱥ25)ȱinȱtheȱ Seaȱ ofȱ Marmaraȱ (Unsalȱ etȱ al.,ȱ 1999)ȱ andȱ theȱ estuarineȱ watersȱ ofȱ theȱ Riaȱ deȱ Vigoȱ (Guerra,ȱ1984).ȱ

3.1.12.3 Life history Afterȱ hatching,ȱ juvenilesȱ immediatelyȱ adoptȱ aȱ benthicȱ lifestyle.ȱ Growthȱ wasȱ calculatedȱatȱ2.8ȹmmȱmonthȺƺ1ȱforȱmalesȱandȱ3.0ȹmmȱmonthȺȬ1ȱforȱfemalesȱinȱtheȱSicilianȱ Channelȱ(RagoneseȱandȱJereb,ȱ1991),ȱaȱlittleȱfasterȱthanȱthatȱestimatedȱinȱtheȱwesternȱ MediterraneanȱbyȱMangoldȬWirzȱ(1963)ȱandȱinȱtheȱRiaȱdeȱVigoȱbyȱGuerraȱ(1984;ȱ2Ⱥ– 2.5ȹmmȱ monthȺȬ1).ȱ Lifespanȱ rangesȱ betweenȱ 12ȱ andȱ 19ȹmonths.ȱ Theȱ smallestȱ matureȱ malesȱhaveȱbeenȱreportedȱatȱ20ȹmm,ȱandȱfemalesȱatȱ30ȹmmȱMLȱ(GuerraȱandȱCastro,ȱ 1989;ȱ Volpiȱ etȱ al.,ȱ 1990).ȱ However,ȱ mostȱ individualsȱ attainȱ maturityȱ atȱ largerȱ sizesȱ (MangoldȬWirz,ȱ1963;ȱVolpiȱetȱal.,ȱ1990;ȱJerebȱandȱRagonese,ȱ1991),ȱatȱapproximatelyȱ1ȱ yearȱofȱage.ȱMalesȱmayȱcarryȱca.ȱ95ȱspermatophoresȱandȱfemalesȱca.ȱ250ȱeggs.ȱ InȱtheȱMediterraneanȱSea,ȱasȱwellȱasȱinȱtheȱAtlantic,ȱmatureȱmalesȱandȱfemalesȱareȱ presentȱ throughoutȱ theȱ year,ȱ whichȱ suggestsȱ aȱ continuousȱ spawningȱ periodȱ (e.g.ȱ MangoldȬWirz,ȱ 1963;ȱ Roperȱ etȱ al.,ȱ 1984;ȱ Guerra,ȱ 1992;ȱ Belcari,ȱ 1999i).ȱ Theȱ eggsȱ (maximumȱ diameterȱ 5ȹmm;ȱ Guerra,ȱ 1984)ȱ areȱ attachedȱ toȱ alcyonariansȱ (seaȱ fans),ȱ shells,ȱ etc.,ȱ onȱ muddyȱ bottomsȱ or,ȱ lessȱ frequently,ȱ onȱ coralȱ formationsȱ (MangoldȬ Wirz,ȱ 1963).ȱ Continuousȱ recruitmentȱ alsoȱ occurs,ȱ withȱ peaksȱ observedȱ inȱ severalȱ Mediterraneanȱareasȱ(Volpiȱetȱal.,ȱ1990;ȱJerebȱandȱRagonese,ȱ1991;ȱWurtzȱetȱal.,ȱ1991;ȱ D’Onghiaȱetȱal.,ȱ1992;ȱCasaliȱetȱal.,ȱ1998).ȱ

3.1.12.4 Trophic ecology Predators.ȱ Sepiaȱ elegansȱ hasȱ beenȱ identifiedȱ fromȱ stomachȱ contentsȱ ofȱ veryȱ fewȱ predators,ȱbutȱhasȱbeenȱreportedȱfromȱtheȱdolphinfishȱ(Coryphaenaȱhippurus;ȱMassutiȱ etȱal.,ȱ1998)ȱandȱtheȱbottlenoseȱdolphinȱ(Tursiopsȱtruncates;ȱBlancoȱetȱal.,ȱ2001).ȱ Prey.ȱThisȱspeciesȱfeedsȱmainlyȱonȱsmallȱcrustaceans,ȱfish,ȱandȱpolychaetesȱ(Reidȱetȱ al.,ȱ 2005).ȱ Detailedȱ studiesȱ onȱ feedingȱ (e.g.ȱ Guerra,ȱ 1985;ȱ Castroȱ andȱ Guerra,ȱ 1990)ȱ suggestȱ thatȱ thereȱ isȱ noȱ changeȱ inȱ dietȱ withȱ growthȱ and/orȱ maturityȱ andȱ thatȱ theȱ varietyȱofȱpreyȱdoesȱnotȱdecreaseȱwithȱincreasingȱsize.ȱȱ

3.1.12.5 Fisheries SepiaȱelegansȱisȱtakenȱmainlyȱasȱaȱbycatchȱinȱMediterraneanȱandȱWestȱAfricanȱtrawlȱ fisheriesȱ (Roperȱ etȱ al.,ȱ 1984;ȱ Belcari,ȱ 1999i;ȱ Reidȱ etȱ al.,ȱ 2005).ȱ Separateȱ statisticsȱ areȱ generallyȱnotȱreportedȱforȱS.ȱelegans,ȱwhichȱrepresentsȱaȱveryȱsignificantȱpercentageȱofȱ theȱ catchesȱ inȱ someȱ areasȱ ofȱ itsȱ distributionalȱ rangeȱ (Reidȱ etȱ al.,ȱ 2005).ȱ Inȱ theȱ MediterraneanȱSea,ȱitȱisȱmarketedȱalongȱwithȱS.ȱorbignyanaȱandȱsmallȱS.ȱofficinalis,ȱandȱ constitutesȱaȱvaluableȱresourceȱlocally.ȱInȱtheȱSicilianȱChannel,ȱanȱexploitationȱrateȱofȱ 0.73ȱwasȱestimatedȱforȱtheȱspeciesȱ(RagoneseȱandȱJereb,ȱ1991),ȱwhichȱsuggestsȱveryȱ

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intensiveȱfishingȱpressureȱonȱthisȱresource.ȱItȱisȱseparatelyȱrecordedȱinȱlandingsȱfromȱ theȱ Gulfȱ ofȱ Cadiz,ȱ whereȱ itȱ typicallyȱ formsȱ ca.ȱ 7Ⱥ%ȱ ofȱ cuttlefishȱ landingsȱ (theȱ remainderȱbeingȱS.ȱofficinalis).ȱThisȱspeciesȱisȱmarketedȱfreshȱandȱfrozen.ȱ TheȱspeciesȱhasȱbeenȱidentifiedȱinȱcommercialȱlandingsȱinȱItalyȱandȱSpain.ȱ

3.1.13 Sepia orbignyana Férrusac in Orbigny, 1826 Commonȱ names.ȱ Pinkȱ cuttlefishȱ (English);ȱ seicheȱ roseéȱ (French);ȱ seppiaȱ pizzutaȱ (Italian);ȱchocoȬdeȬcaudaȱ(Portuguese);ȱchoquitoȱconȱpuntaȱ(Spanish).ȱ

3.1.13.1 Adult diagnostic features Upȱ toȱ 120ȹmmȱ MLȱ (MangoldȬWirz,ȱ 1963).ȱ Mantleȱ oval.ȱ Clubȱ short,ȱ oval,ȱ withȱ 5Ⱥ–Ⱥ6ȱ suckersȱ inȱ transverseȱ rows.ȱ Suckersȱ differȱ markedlyȱ inȱ size:ȱ threeȱ largeȱ suckersȱ mediallyȱwithȱoneȱslightlyȱsmallerȱsuckerȱonȱeachȱsideȱofȱthese.ȱHectocotylusȱpresentȱ onȱleftȱventralȱarmȱ(IV):ȱ1Ⱥ–Ⱥ2ȱrowsȱofȱnormalȱsizeȱsuckersȱproximally,ȱgreatlyȱreducedȱ suckersȱmedially,ȱthenȱnormalȱsizeȱsuckersȱdistallyȱtoȱarmȱtip.ȱSuckersȱofȱhectocotylusȱ inȱ twoȱ dorsalȱ andȱ twoȱ ventralȱ seriesȱ displacedȱlaterally,ȱ withȱ aȱ gapȱ betweenȱ them.ȱ Cuttleboneȱ outlineȱ oblong.ȱ Anteriorȱ striaeȱ shallowȱ MȬshape,ȱ orȱ wavy.ȱ Spineȱ long,ȱ pointedȱ (prominent),ȱ straight,ȱ directedȱ dorsally,ȱ withȱ ventralȱ keel.ȱ Colourȱ reddishȱ brown.ȱReference:ȱReidȱetȱal.ȱ(2005).ȱ

3.1.13.2 Distribution and habitat Sepiaȱ orbignyanaȱ occursȱ inȱ theȱ easternȱ Atlanticȱ fromȱ theȱ Irishȱ Seaȱ inȱ theȱ northȱ toȱ southernȱAngolaȱ(17°S)ȱandȱisȱpresentȱthroughoutȱtheȱMediterraneanȱSeaȱ(Jerebȱandȱ Roper,ȱ2005).ȱItȱisȱaȱdemersalȱspeciesȱthatȱprefersȱsandyȱandȱsandyȺ–Ⱥmuddyȱbottoms.ȱ Itsȱ greatestȱ abundanceȱ isȱ atȱ depthsȱ ofȱ 50Ⱥ–Ⱥ250ȹmȱ (MangoldȬWirz,ȱ 1963;ȱ Casaliȱ etȱ al.,ȱ 1998).ȱRecordsȱfromȱtheȱSeaȱofȱMarmaraȱindicateȱthatȱitȱcanȱoccurȱinȱbrackishȱwatersȱ (Unsalȱetȱal.,ȱ1999).ȱ

3.1.13.3 Life history Newlyȱhatchedȱanimalsȱobservedȱinȱtheȱlaboratoryȱ(Boletzky,ȱ1988)ȱmeasuredȱ6ȹmmȱ ML.ȱFemalesȱattainȱlargerȱsizesȱandȱgrowȱfasterȱthanȱmalesȱ(Bello,ȱ2001).ȱGrowthȱwasȱ calculatedȱatȱ2.9ȹmmȱmonthȺƺ1ȱforȱmalesȱandȱ3.0ȹmmȱmonthȺƺ1ȱforȱfemalesȱinȱtheȱSicilianȱ Channelȱ(RagoneseȱandȱJereb,ȱ1991).ȱLifespanȱisȱconsideredȱtoȱvaryȱbetweenȱ12ȱandȱ 18ȹmonthsȱ(MangoldȬWirz,ȱ1963).ȱEstimatesȱfromȱlengthȱfrequencyȱanalyses,ȱalthoughȱ widelyȱ consideredȱ toȱ beȱ unreliableȱ forȱ cephalopods,ȱ indicateȱ aȱ longerȱ lifespanȱ ofȱ 2ȱ yearsȱ forȱ malesȱ andȱ 3ȱ yearsȱ forȱ femalesȱ (Ragoneseȱ andȱ Jereb,ȱ 1991).ȱ Malesȱ reachȱ maturityȱ fromȱ 35ȹmmȱ ML,ȱ andȱ femalesȱ fromȱ 65ȹmmȱ MLȱ (Belcariȱ andȱ Sartor,ȱ 1993).ȱ Matureȱmales,ȱagedȱ6ȱorȱ7ȹmonths,ȱcarryȱca.ȱ100ȱspermatophores;ȱfemalesȱagedȱ9ȱorȱ 10ȹmonthsȱcarryȱca.ȱ400ȱeggsȱ(Roperȱetȱal.,ȱ1984).ȱ Inȱ Mediterraneanȱ waters,ȱ spawningȱ isȱ probablyȱ continuous.ȱ Theȱ eggs,ȱ laidȱ individually,ȱwhitishȱorȱgreyishȱinȱcolourȱandȱslightlyȱmoreȱelongateȱthanȱthoseȱofȱS.ȱ officinalisȱ(MangoldȬWirz,ȱ1963),ȱareȱdepositedȱinȱvariousȱspongesȱtoȱformȱclustersȱofȱ 30ȺȭȺ40ȱ(Reidȱetȱal.,ȱ2005).ȱ

3.1.13.4 Trophic ecology Weȱ foundȱ noȱ reportsȱ ofȱ thisȱ speciesȱ fromȱ predatorȱ stomachȱ contents.ȱ S.ȱ orbignyanaȱ feedsȱmainlyȱonȱcrustaceansȱandȱfishȱ(Auteriȱetȱal.,ȱ1988).ȱȱ

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3.1.13.5 Fisheries SepiaȱorbignyanaȱisȱtakenȱmainlyȱasȱbycatchȱthroughoutȱtheȱMediterraneanȱandȱinȱtheȱ WestȱAfricanȱtrawlȱfisheries,ȱalthoughȱtargetedȱfisheriesȱalsoȱexistȱ(e.g.ȱRoperȱetȱal.,ȱ 1984;ȱMangoldȱandȱBoletzky,ȱ1987;ȱBelcari,ȱ1999j;ȱReidȱetȱal.,ȱ2005).ȱSeparateȱstatisticsȱ areȱ notȱ reported,ȱ butȱ S.ȱ orbignyanaȱ representsȱ aȱ veryȱ significantȱ percentageȱ ofȱ theȱ catchesȱinȱsomeȱareas.ȱInȱtheȱMediterraneanȱSea,ȱitȱisȱoftenȱmarketedȱalongȱwithȱtheȱ congenerȱS.ȱelegansȱandȱsmallȱS.ȱofficinalisȱandȱconstitutesȱaȱvaluableȱresourceȱlocallyȱ (Belcari,ȱ 1999j).ȱ Studiesȱ carriedȱ outȱ inȱ theȱ Sicilianȱ Channelȱ demonstratedȱ anȱ exploitationȱrateȱofȱ0.60ȱforȱthisȱspeciesȱ(RagoneseȱandȱJereb,ȱ1991),ȱwhichȱsuggestsȱ intensiveȱfishingȱpressureȱonȱtheȱresource,ȱasȱalreadyȱhypothesizedȱbyȱotherȱauthorsȱ (e.g.ȱBoletzky,ȱ1983).ȱItȱisȱmarketedȱfreshȱandȱfrozen.ȱ TheȱspeciesȱhasȱbeenȱidentifiedȱinȱcommercialȱlandingsȱinȱItaly.ȱȱ

3.2 Application of DNA-based markers in European cephalopod fisheries and biology

PaulȱShawȱ

3.2.1 Introduction Inȱ theȱ pastȱ 30ȱ years,ȱ DNAȬbasedȱ markers,ȱ suchȱ asȱ allozymes,ȱ mitochondrialȱ DNAȱ (mtDNA),ȱandȱmicrosatellites,ȱhaveȱmadeȱaȱsubstantialȱcontributionȱtoȱmanyȱfieldsȱofȱ wholeȱ organismȱ biology.ȱ Inȱ particular,ȱ populationȱ biologyȱ (populationȱ structuring,ȱ individualȱ interactions),ȱ phylogenetics,ȱ andȱ systematicsȱ haveȱ benefitedȱ fromȱ theȱ applicationȱ ofȱ dependable,ȱ geneticallyȱ determined,ȱ inheritedȱ markersȱ thatȱ areȱ notȱ influencedȱ byȱ theȱ environmentȱ andȱ individualȱ habits.ȱ Theȱ valueȱ ofȱ DNAȬbasedȱ markersȱ hasȱ beenȱ demonstratedȱ byȱ theirȱ widespreadȱ useȱ withȱ finfishȱ andȱ theirȱ fisheriesȱ(seeȱCarvalhoȱandȱPitcher,ȱ1995).ȱ Despiteȱtheȱobviousȱbenefitsȱofȱtheseȱmarkers,ȱtheirȱinputȱtoȱcephalopodȱbiologyȱhasȱ beenȱ limitedȱ untilȱ theȱ lastȱ tenȱ years.ȱ Substantialȱ earlyȱ successȱ wasȱ achievedȱ usingȱ allozymesȱ toȱ addressȱ issuesȱ ofȱ populationȱ structuringȱ (e.g.ȱ Brierleyȱ etȱ al.,ȱ 1995),ȱ speciesȱ identificationȱ andȱ systematicsȱ (e.g.ȱ Augustynȱ andȱ Grant,ȱ 1988),ȱ andȱ particularlyȱinȱidentifyingȱcrypticȱspeciesȱ(e.g.ȱYeatmanȱandȱBenzie,ȱ1994).ȱTheȱrecentȱ developmentȱ ofȱ microsatelliteȱ DNAȱ markersȱ forȱ cephalopodȱ speciesȱ andȱ theirȱ applicationȱ toȱ theȱ studyȱ ofȱ populationȱ structureȱ (e.g.ȱ Shawȱ etȱal.,ȱ 1999)ȱandȱ matingȱ strategiesȱ(e.g.ȱShawȱandȱBoyle,ȱ1997)ȹ–ȹandȱtheȱincreasingȱuseȱofȱDNAȱsequencingȱforȱ phylogenetics/systematicsȱpurposesȱ(e.g.ȱStrugnellȱetȱal.,ȱ2005)ȹ–ȹhasȱdemonstratedȱtheȱ potentialȱforȱgreaterȱcontributionsȱtoȱcephalopodȱbiologyȱonȱaȱnumberȱofȱlevels.ȱȱ

3.2.2 Population genetics of commercially important cephalopods in European waters

3.2.2.1 Loligo forbesii Allozymeȱ(Brierleyȱetȱal.,ȱ1995),ȱmicrosatelliteȱ(Shawȱetȱal.,ȱ1999),ȱandȱmitochondrialȱ DNAȱdataȱ(Normanȱetȱal.,ȱ1994)ȱallȱindicateȱthatȱthisȱspeciesȱisȱcomprisedȱofȱaȱsingleȱ geneticȱstockȱthroughoutȱitsȱrangeȱalongȱtheȱAtlanticȱcoastȱofȱEuropeȱ(northwesternȱ Scotlandȱ toȱ southernȱ Portugal),ȱ butȱ thatȱ aȱ significantlyȱ differentiatedȱ populationȱ existsȱ inȱ theȱ Azoresȱ archipelago.ȱ Shawȱ etȱ al.ȱ (1999)ȱ alsoȱ indicatedȱ theȱ possibilityȱ ofȱ geneticallyȱ differentiatedȱ populationsȱ onȱ offȬshelfȱ banksȱ northwestȱ ofȱ Scotlandȱ (particularlyȱ onȱ Rockallȱ Bank).ȱ Thisȱ geneticȱ patternȱ ofȱ largeȬscaleȱ homogeneityȱ isȱ consistentȱwithȱwhatȱisȱknownȱofȱtheȱlifeȬhistoryȱcharacteristicsȱofȱthisȱspecies,ȱwhichȱ isȱhighlyȱmobileȱandȱmigratoryȱasȱjuvenilesȱandȱadults.ȱTheȱgeneticȱdifferentiationȱofȱ