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绪论 虎鲸类型和种群(第2页)

Northernwaters

NorthPacific

ResearchoffthewestcoastofCanadaandtheUnitedStatesinthe1970sand1980sidentifiedthefollowingthreetypes:

Resident(fish-eating)orcas:Thecurveddorsalfinsaretypicalofresidentfemales.

Resident:ThesearethemostmonlysightedofthethreepopulationsinthecoastalwatersofthenortheastPacific.Residentsdietsconsistprimarilyoffishandsometimessquid,andtheyliveinplexandcohesivefamilygroupscalledpods.Femaleresidentscharacteristicallyhaveroundeddorsalfintipsthatterminateinasharpcorner.Theyvisitthesameareasconsistently.BritishColumbiaandWashingtonresidentpopulationsareamongstthemostintensivelystudiedmarinemammalsanywhereintheworld.Residentorcascanbedividedintoatleastthreedistinctmunities;northern,southernandsouthernAlaskan.SouthernAlaskanresidentorcasaredistributedfromsoutheasternAlaskatotheKodiakArchipelagoandnumberover700individuals.Thesewhalesconsistoftwointerbreedingclansdistinguishedbyacousticcallsandwhoserangesoverlap.ThenorthernresidentmunitylivesincoastalandinlandwatersfromsoutheasternAlaskatoVancouverIsland.Itconsistsofthreeclansand16podsandnumberover300orcastotal.ThesouthernresidentmunitygenerallyinhabitstheinlandwatersofsouthernBritishColumbiaandWashington,butcanbefoundintheouterwatersoffVancouverIsland,Washington,OregonandCalifornia.Theyconsistofoneclanandthreepods,andnumberlessthan80individualsandarelistedasendangered.

TransientorBiggs:Thedietsoftheseorcasconsistalmostexclusivelyofmarinemammals.Theyliveinthesameareasasresidents,butthetwoavoideachother.Transientsgenerallytravelinsmallgroups,usuallyoftwotosixanimals,andhavelesspersistentfamilybondsthanresidents.Transientsvocalizeinlessvariableandlessplexdialects.Femaletransientsarecharacterizedbymoretriangularandpointeddorsalfinsthanthoseofresidents.Thegreyorwhiteareaaroundthedorsalfin,knownasthe“saddlepatch“,oftencontainssomeblackcolouringinresidents.However,thesaddlepatchesoftransientsaresolidanduniformlygrey.Transientsroamwidelyalongthecoast;someindividualshavebeensightedinbothsouthernAlaskaandCalifornia.TransientsarealsoreferredtoasBiggsorcainhonourofcetologistMichaelBigg.Thetermhasbeeincreasinglymonandmayeventuallyreplacethetransientlabel.Thetransientecotypeisestimatedtohavediverged700,000yearsago.Thereareatleastthreedifferent“stocks“oftransientsoffNorthAmerica,theAT1stockwhichoccursfromPrinceWilliamSoundtoKenaiFjords,theGulfofAlaskaAleutianIslandsBeringSea(GOAAIBS)stockandthewestcoaststockwhichrangesfromsoutheastAlaskatoCalifornia.AT1isconsideredadepletedstock;itwasaffectedbytheExxonValdezoilspillanddeclinedfrom22individualstoeightbetween1989and2004.TheGOAAIBSstockmaynumberaround500whaleswhilethewestcoasttransientsnumberover320orcaswithover200alongsoutheastAlaska,BritishColumbiaandWashingtonandover100orcasoffCalifornia.Californiatransientsdonotappeartointerminglemuchwiththosefurthernorthandwestcoasttransientsmaybedividedintosub-munities.

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Offshore:AthirdpopulationoforcasinthenortheastPacificwasdiscoveredin1988,whenahumpbackwhaleresearcherobservedtheminopenwater.Astheirnamesuggests,theytravelfarfromshoreandfeedprimarilyonschoolingfish.However,becausetheyhavelarge,scarredandnickeddorsalfinsresemblingthoseofmammal-huntingtransients,itmaybethattheyalsoeatmammalsandsharks.TheyhavemostlybeenencounteredoffthewestcoastofVancouverIslandandnearHaidaGwaii.Offshorestypicallycongregateingroupsof20–75,withoccasionalsightingsoflargergroupsofupto200.Littleisknownabouttheirhabits,buttheyaregeneticallydistinctfromresidentsandtransients.Offshoresappeartobesmallerthantheothers,andfemalesarecharacterizedbydorsalfintipsthatarecontinuouslyrounded.

Separatefish-eatingandmammal-eatingorcamunitiesalsoexistoffthecoastoftheRussianFarEastandHokkaido,Japan.RussianorcasaremonlyseenaroundtheKamchatkaPeninsulaandCommanderIslands.Over2,000individualresident-likeorcasand130transient-likeorcashavebeenidentifiedoffRussia.Atleast195individualorcashavebeencatalogedintheeasterntropicalPacific,rangingfromBajaCaliforniaandtheGulfofCaliforniainthenorthtothenorthwestcoastofSouthAmericainthesouthandwesttowardsHawaii.OrcasappeartoregularlyoccurofftheGalápagosIslands.OrcassightedinHawaiianwatersmaybelongtoagreaterpopulationinthecentralPacific.

NorthAtlanticandadjacent

Orcatail-slappinginVestfjorden,Norway

Atleast15,000whalesareestimatedtoinhabittheNorthAtlantic.IntheNortheastAtlantic,twoorcaecotypeshavebeenproposed.Type1orcasconsistofsevenhaplotypesandincludeherring-eatingorcasofNorwayandIcelandandmackerel-eatingorcasoftheNorthSea,aswellasseal-eatingorcasoffNorway.Type2orcasconsistoftwohaplotypes,andmainlyfeedonbaleenwhales.

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IntheMediterraneanSea,orcasareconsidered“visitors“,likelyfromtheNorthAtlantic,andsightingsbeelessfrequentfurthereast.However,asmallyear-roundpopulationexistsintheStraitofGibraltar,whichnumberedaround39in2011.DistinctpopulationsmayalsoexistoffthewestcoastoftropicalAfrica,whichhavegeneralizeddiets.

ThenorthwestAtlanticpopulationisfoundyear-roundaroundLabradorandNewfoundland,whilesomeindividualsseasonallytraveltothewatersoftheeasternCanadianArcticwhentheicehasmelted.SightingsofthesewhaleshavebeendocumentedasfarsouthasCapeCodandLongIsland.ThispopulationispossiblycontinuouswithorcassightedoffGreenland.Orcasaresightedyear-roundintheCaribbeanSea,andanestimated267(asof2020)isdocumentedinthenorthernGulfofMexico.

NorthIndianOcean

Over50individualwhaleshavebeencatalogedinthenorthernIndianOcean,includingtwoindividualsthatweresightedinthePersianGulfin2008andoffSriLankain2015.

Southernwaters

OrcabeachingtocapturesealionatValdesPeninsula

AsmallpopulationoforcasseasonallyvisitsthenorthernpointoftheValdesPeninsulaontheeastcoastofArgentinaandhuntforsealionsandelephantsealsontheshore,temporarystrandingthemselves.OffSouthAfrica,adistinctive“flat-tooth“morphotypeexistsandpreysonsharks.Apairofmaleorcas,PortandStarboard,havebeewellknownforhuntinggreatwhitesandothersharksofftheSouthAfricancoast.

OrcasoccurthroughoutthewatersofAustralia,NewZealandandPapuaNewGuinea.TheyaresightedyearroundinNewZealandwaters,whileoffAustralia,theyareseasonallyconcentratedoffthenorthwest,intheinshorewatersofNingalooReef,andthesouthwest,attheBremerregion.GeneticevidenceshowsthattheorcasofNewZealand,andnorthwestandsouthwestAustraliaformthreedistinctpopulations.NewZealandorcasmainlypreyonsharksandrays.

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Antarctic

Around25,000orcasareestimatedaroundtheAntarctic,andfourtypeshavebeendocumented.Twodwarfspecies,namedOrcinusnanusandOrcinusglacialis,weredescribedduringthe1980sbySovietresearchers,butmostcetaceanresearchersareskepticalabouttheirstatus,andlinkingthesedirectlytothetypesdescribedbelowisdifficult.

Someexamplesofvariationsinorcas

TypeAorAntarcticorcaslooklikea“typical“orca,alarge,black-and-whiteformwithamedium-sizedwhiteeyepatch,livinginopenwaterandfeedingmostlyonminkewhales.

TypeB1orpackiceorcasaresmallerthantypeA.Ithasalargewhiteeyepatch.Mostofthedarkpartsofitsbodyaremediumgreyinsteadofblack,althoughithasadarkgreypatchcalleda“dorsalcape“stretchingbackfromitsforeheadtojustbehinditsdorsalfin.Thewhiteareasarestainedslightlyyellow.Itfeedsmostlyonseals.TypeB1orcaareabundantbetweenAdelaideIslandandthemainlandAntarcticpeninsula.

TypeB2orGerlacheorcasaremorphologicallysimilartoTypeB1,butsmaller.Thisecotypehasbeenrecordedfeedingonpenguinsandseals,andisoftenfoundintheGerlacheStrait.

TypeCorRossSeaorcasarethesmallestecotypeandliveinlargergroupsthantheothers.Itseyepatchisdistinctivelyslantedforwards,ratherthanparalleltothebodyaxis.LiketypeB,itisprimarilywhiteandmediumgrey,withadarkgreydorsalcapeandyellow-tingedpatches.ItsonlyobservedpreyistheAntarcticcod.

TypeDorSub-Antarcticorcaswerefirstidentifiedbasedonphotographsofa1955massstrandinginNewZealandandsixat-seasightingssince2004.Thefirstvideorecordofthistypewasmadein2014betweentheKerguelenandCrozetIslands,andagainin2017offthecoastofCapeHorn,Chile.Itisrecognizablebyitssmallwhiteeyepatch,narrowerandshorterthanusualdorsalfin,bulboushead(similartoapilotwhale),andsmallerteeth.Itsgeographicrangeappearstobecircumglobalinsub-Antarcticwatersbetweenlatitudes40°Sand60°S.Althoughitsdietisnotdetermined,itlikelyincludesfish,asdeterminedbyphotographsaroundlonglinevessels,whereTypeDorcasappearedtobepreyingonPatagoniantoothfish(Dissostichuseleginoides).

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TypesBandCliveclosetotheice,anddiatomsinthesewatersmayberesponsiblefortheyellowishcolouringofbothtypes.MitochondrialDNAsequencessupportthetheorythatthesearerecentlydivergedseparatespecies.Morerecently,pletemitochondrialsequencingindicatesthetypesBandCberecognizedasdistinctspecies,asshouldtheNorthPacifictransients,leavingtheothersassubspeciespendingadditionaldata.AdvancedmethodsthatsequencedtheentiremitochondrialgenomerevealedsystematicdifferencesinDNAbetweendifferentpopulations.A2019studyofTypeDorcasalsofoundthemtobedistinctfromotherpopulationsandpossiblyevenauniquespecies.

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