8SocialBehaviour

Introduction

Formanyyearsitwasthoughtthatthedomesticcatwasanessentiallysolitarycreature,whichonlytoleratedthecloseproximityofitsconspecificsformatingandwhilerearingoffspring.Ithasoftenbeensaidthatthelionistheonlyfullysocialfelid.However,whileitistruethatthesocialsystemofthelioniscomplex,involvingcommunalhuntingandcooperationwithinbothmaleandfemalegroups,socialinteractionbetweenmembersofotherwisesolitaryfelidspecieshasbeenobserved(Caro,1989).Studiesofthesocialstructureofgroupsofdomesticcatshaveshownthattheyarenotjustartefactsoftheconditionsunderwhichhousecatsarekept;infact,socialstructureismostclearlypresentingroupsthatarebarelytolerantofhumancompany.Lookingobjectivelyatthevastrangeofpopulationdensitiesrecordedfordomesticcats(Fig.8.1),itseemsveryunlikelythatauniformsystemforintraspecificinteractionscouldbeeffective,whenindividualcatscanfindthemselvesspacedatanythingfromanaverageoftentoseveralthousandmetresapart.IncommonwithothermembersoftheCarnivorathatcanadapttoawiderangeofpopulationdensities,thesocialstructurethatpertainstoeachpopulationofdomesticcatsvariesaccordingtotheecologicalcircumstancesinwhichtheyfindthemselves(Macdonald,1983).Essentially,groupsmaybeformedwhentheavailabilityanddispersionoffoodallowstwoormoreindividualstoliveincloseproximity,andonalltheoccasionsthatthishasbeendocumentedmuchofthisfoodhasstemmedfromman’sactivities.Thisinevitablyraisestwoquestions:whethersuchcoalitionscaneveroccurwithoutman’stacitcollaborationand,ifnot,howthenecessarybehaviourpatternsevolved,unlesstheyareaby-productofdomestication.

Fig.8.1.Therelationshipbetweenhomerangesizeandcatdensity,forentiremales,femalesandneuters.Thebrokenlineindicatesthesizeofhomerangeexpectedifeachpartoftheavailablespacewasallocatedtothehomerangeofonemaleandonefemale;pointswellabovethislineindicateoverlapbetweentherangesofmembersofthesamesex,whilepointswellbelowindicatethatnotalltheavailablespaceisused(datafromLibergandSandell,1988,withadditionsfromChipman,1990,andJ.W.S.Bradshaw,unpublished).

SolitaryCats

Thequestionofwhethercatskeptsinglyinhouseholdsaretrulysolitarywillbelefttothefollowingchapter,butitcanbearguedthat,likethedogthatappearstoperceiveitshumanownersaspartofits‘family’,mostcatsdirectspecies-typicalbehaviourtowardstheirhumankeepers.Trulysolitarycatsthathavelittleconspecificorhumancontactformuchoftheyearhavenotbeenstudiedinmuchdetail,notleastbecausetheyaredifficulttolocateandhardertoapproach.TheEuropeanwildcat(Feliss.silvestris),althoughnottheclosestwildancestortothedomesticcat,isthoughttobealmostentirelysolitaryandmaybegeneticallypredisposedtobeso,sinceevenitskittensareverydifficulttodomesticate.LargelysolitarypopulationsofFeliss.catusarealsoknown,forexamplethoseinthebushofsouth-eastAustraliaandonsomeuninhabited

islands.Generally,thesepopulationssupportthemselvesbyhuntingand,becauseitisrareforpreysuitablefordomesticcatstobehighlyabundantinanyonelocationforanylengthoftime,socialgroupsarerare.Undersuchcircumstancescatsarerarelyseeninthecompanyofothercats,exceptformale/femalepairsatoestrus,andfemaleswithjuveniles.Theadultsareusuallyterritorialtosomeextent,althoughthemechanismwherebysuchterritoriesaremaintained,giventhatindividualssorarelyencounteroneanother,isnotclear.Whenfoodismorepatchilydistributed,buteachpatchorgroupofpatchesisstillinsufficienttosupportmorethanonecat,homerangescanoverlapquiteextensively,butsomesystemoftemporalseparationmaythenoperate,sothattwocatsrarelyhuntthesameareaatthesametime.Scentmarkinghasbeenimplicatedinmaintainingthis‘time-sharing’arrangement(seeChapter5,thisvolume).AsdiscussedinChapter7,thehomerangesoffemalesencompasssufficientfoodandshelterfortheirneedsandthoseoftheiroffspringwhiletheyaredependent.Evenwhencatsarewelldispersed,thehomerangesofmalesarelargerthanthoseoffemales(Fig.8.1)–3.5timeslargeronaverage.Malerangesofupto10km2havebeenrecorded,anditislikelythatsomemales,labelledas‘transient’inmoststudies,eitherhaveevenlargerhomerangesthanthisoraremoreorlessnomadic(LibergandSandell,1988).Malerangesappeartobedictatedbytheavailabilityofbreedingfemales,whetherthesearesolitaryorsocial,andthedegreeofcompetitionforthem;thefactorsdeterminingthesizeofindividualmalerangeswillbediscussedfurtherbelow.

Group-livingCats

Bothwide-rangingsurveysandmoredetailedstudiesofsmallareashavedocumentedtheexistenceofcoloniesofdomesticcats.Theconditionsfortheestablishmentofthesecoloniesalmostalwaysinvolvealocalizedconcentrationoffood,arisingdeliberatelyoraccidentallyfromhumanactivities(KerbyandMacdonald,1988).Someoccuraroundrubbishdumps,studiedinlocationsasdiverseasPortsmouthnavaldockyardintheUKandaJapanesefishingvillage.Othersaremoredirectproductsofprovisioning,suchasthesemi-wildpopulationsoftenfoundonindustrialandhospitalsitesorinvariouspublicareasinRome.Farmcatsareprovisionedbothdirectlyandindirectly,bydirecthandoutsoffood,bytheconcentrationofrodentsingrainstoresandsometimesbythetheftoffoodintendedforlivestock.Thesizeofthecoloniesseemstobeveryvariable;inonesurveyof300coloniesonindustrialsitesintheUK,mostcomprisedbetweenoneandtenindividualsbut7%containedover50cats.The

comprisedbetweenoneandtenindividualsbut7%containedover50cats.Thecriticalfactorsdeterminingthesizeofthecoloniesaretheavailabilityoffood,infantmortalityduetofelinepanleucopeniaandotherviruses,anddirectkillingofadultsbyman.

Smallcoloniesmayconsistofasinglesocialunit,whilelargercoloniesusuallycontainseveralsuchgroups.Mostofthegroupsthathavebeenstudiedhaveturnedouttoconsistoffemales,usuallyrelated,togetherwiththeiroffspring,includingimmaturemales,andunrelatedmaturemales.Spontaneousmovementoffemalesbetweengroupsseemstoberare,probablybecausewhilefemaleswithinagrouparegenerallytolerantofoneanother,theyusuallyattackoutsidersonsight,bothmalesandfemales,andtheseattacksgenerallybecomemoreintensewhenthereareyoungkittensinthegroup.Maturemalesareonlylooselyattachedtoanyonegroupand,aswithsolitarycats,theirhomerangestendtobelargerthanthoseoffemales.

SexualBehaviourofFemales

Apartfromthecontactbetweenmotherandoffspring,describedinChapter4,thisvolume,theonlyessentialcomponentofsocialbehaviourrequiredofasolitaryfemaleisthatleadinguptomating.Insolitaryindividuals,therewillbeastrongterritoriallybasedtendencytoattackanycat,andoneofthemainfunctionsofcourtshipbehaviourmaysimplybetobringthesexestogetherwithoutfightingforlongenoughforcopulationtotakeplace.Evenwithinacolonyofcats,whereallindividualsarefamiliartooneanother,ifamaleshowsmorethanafleetingsexualinterestinananoestrousfemale,thefemalewillmoveawayandifnecessaryspitandstrikeoutwithherclaws.Inpro-oestrusthebehaviourofthefemalechanges,firstsubtlyasatendencytomoveaboutmorethanusual,andthenasanincreaseinobject-rubbing(seeChapter5,thisvolume).Malesthatapproachatthisstagearegreetedwithlesshostilitythanbefore,butprolongedcontactisstillnottolerated.Overthenext24htherubbingincreasesinintensityandpersistentboutsofrollingoccur,accompaniedbypurring,stretchingandrhythmicopeningandclosingoftheclaws.Malesarenowpermittedclosetothefemale,andmaybeallowedtolickher,butatthisstageanyattemptsatmountingresultinaconsiderabledisplayofaggression.

Completesexualreceptivitydoesnotensueuntilthebeginningofoestrus,whichisoftenindicatedbyanabruptchangeinbehaviour.Therollingofpro-oestrusisinterruptedbythefemaleadoptingthelordosisposition,suddenlycrouchingwithherheadclosetothegroundandherhindlegstreadingandpartlyextended.Hertailislaterallydisplaced,uncoveringtheperineum,anditisatthe

extended.Hertailislaterallydisplaced,uncoveringtheperineum,anditisatthemomentthatthisdisplayfirstappearsthatanexperiencedmalewillfirstattempttomount.Graspingherneckinhisjaws,hebeginshiscopulatorythrusts,whilethefemaletreadsbackwardswithherhindlegssothattheperineumisrotatedfurtherbackwardsandupwards,untilthemaleachievesintromission.Atthispointthefemaleusuallyemitsaloud,piercingcry,andwithinafewsecondsjumpsawayfromthemaleandturnsonhim,spittingandscratching.Thefemalethengroomshergenitalregionandbeginstorollvigorously.Severalminuteslatershewilladoptthelordosispositionagain,andthiscycleofeventscanberepeated,withtheintervalbetweencopulationslengthening,overthenext1or2days(Michael,1961).Multiplecopulationsarenormallyneededtotriggerovulation,andwithoutcopulationovulationdoesnotoccur.Ontheonehand,ithasbeensuggestedthatthisinducedovulationisanadaptationtosolitaryliving,preventingthefemalefromovulatingwastefully,beforeshehasbeenabletoattractamale.Ontheotherhand,thewholeprocessofpro-oestrus,whenthefemaleisattractivetomalesbutnotreceptive,andalsotheneedformultiplematings,couldalsobedevicestoenhancecompetitionbetweenseveralmalescourtingonefemale,andmaythereforenotonlybeanadaptationthatenhancesthefemale’sfitness,butalsotolivingathighdensity.

Socialaspectsofmaternalbehaviour

Femalestendtostaywithinasinglesocialgroupformuchoftheirlives;asolitarycatmayoccasionallyjoinanestablishedgroup,andformerlygroup-livingfemalescanbecomesolitary,butmigrationbetweengroupsbyfemalesseemstoberatherrare(LibergandSandell,1988).Devillardetal.(2003)foundthatsomefemalesdispersedat1–2yearsofagefromtheirstudycolonyofintact,high-densitycatsandsuggestthatcatsmayshowintraspecificvariabilityofdispersalpatternsrelatedtochangesinenvironment,density,matingsystemandsocialorganization.Theirgeneraltendencytoremainintheirnatalgroup,combinedwiththeamicabletreatmentoffellowmembersofthegroup,contrastingwithaggressiontowardsoutsiders,impliesthatgroup-livingfemalesdefendacommunalcoreterritory,whichislikelytoincludetheirdenningsitesandtheirmajorsourceoffood.Largecoloniesusuallyconsistofoneormore‘central’groupsofrelatedfemalesthatdefendareasthatcontainthebestresourcesforfeedinganddenning,togetherwithother‘peripheral’femalesthatareeithersolitaryorformsmallergroupsthatoccupyinferiorterritories(Macdonaldetal.,2000).Centralfemalesgenerallyproducemoresurvivingoffspringthanperipheralfemales,suggestingthatcombiningtheireffortsto

defendtheirresourcesgivesthemagenuinereproductiveadvantage.Theotherobviousbenefitfromcooperationbetweenthefemalemembersofa

groupisthecommunaldenningandnursingofkittens.Whileinlargegroupssuchcollaborationstendtobewithinmother–daughterpairs,insmallgroupsalltheadultfemalesmaynurseeachother’soffspringandthelittersareoftenpooledincommunaldens.Femaleswillaidinthebirth,thengroom,nurseandguardeachother’skittens.Poolingoflittersmayalsofacilitatethelearningofafamilyidentityamongthekittens,enablingthemtoformkin-basedalliancesonceadult.Otheradvantagesmayalsoexistforsuchpooledlitters.Inonestudy,communallittersoccupiedtwiceasmanynestsassolitarylittersoverthefirst6weeksafterbirth.Thereasonsforsuchfrequentnestmovesareuncertain,butmayhelptomaintainconcealmentofkittensfrompredatorsandavoidpossibleinfanticide(Feldman,1993).Kittensaregraduallymovedtonestsnearertothefoodsourceastheydeveloptowardsindependenceinbothsolitaryandcommunallitters,sofacilitationofweaningmaybeareasonforrepeatedmoves.Whateverthereason,kittensfromcommunalnestsspendlesstimealoneandarethereforeatlessriskfrompredatorsduringsuchmovementsthankittensfromafemaleraisingherkittensalone.Kittensfrompooledlittersalsoleavethenestearlierthanthoseinsolitarylitters,suggestingthatcommunaldenningmayconfersomedevelopmentaladvantagesonthekittens.

However,therearealsodisadvantagestocommunalliving.Severalcontagiousdiseasesareideallysuitedtotransmissionwithinacommunaldenandbetweencatssharingconcentratedfoodsources.OntheoceanicMarionIsland,thefelinepanleucopeniaviruswasintroduceddeliberatelyasameasuretocontroltheferalcatpopulation,andnotonlywasthereafourfolddecreaseinthepopulation,buttheproportionofcatslivingingroupsdeclinedalso.Onceavirusinfectsafarmcatcolony,itmayspreadrapidlytoallmembers,andnotjustwithincoregroupsthatsharenestsandrestingplaces(Macdonaldetal.,2000):transmissionbetweencoloniesmaysimplybelowbecausemigrationofcatsfromonefarmcolonytoanotherisrare,asisclosecontactbetweencatsindifferentcolonies.

MaleBehaviour

Thereisverylittleevidenceforcooperativebehaviourbetweenintactmalecats;forexample,Dards(1983)neverobservedanyamicablebehaviourbetweenanyofthematuremalesinaPortsmouthdockyard.Whentwounfamiliarmalesmeet

forthefirsttimetheymayinitiallysniffeachother,butthisquicklygiveswaytoaggression,includingthearched-backposture(Fig.5.1),growlingandyowling.Ifonemalegoesontothedefensive,indicatingtheother’ssuperiority,itwilltendtocrouch,hissandstrikeoutwithitsfore-claws.Afterrepeatedencounters,overtaggressionisreducedbuttheoriginalwinnerwilltendtosprayurineandrubobjectsmorefrequentlythantheloser(deBoer,1977b).Affiliativebehaviourbetweenintactmalesappearstoberestrictedtojuvenilesfromthesamefamilygroup,whichmayassociatetogetheruntiltheybecomesexuallymatureanddisperse(Macdonaldetal.,2000).

Territorialityandthematingsystem

Thesizeandorganizationofhomerangesandthematingsystemofmaledomesticcatstendtovaryaccordingtotheenvironmentalconditionsanddensityoftheparticularcatpopulation.Thehomerangesofmalesare,onaverage,about3.5timeslargerthanthoseoffemaleslivingunderthesameecologicalconditions,butthisapparentlysimplerelationshipconcealsagreatdealofvariability.Inruralareaswherecatsarelivingatlowdensity(100–300cats/km2;Libergetal.,2000),malesmaintainexclusivehomerangeswhichmayoverlapthoseofseveralfemales.Theymaydefendaccesstofemalesinoestrusduringthematingseasontherebysecuringmost,ifnotall,ofthematings.Thissystemisclassifiedaspolygynous(PontierandNatoli,1996;Sayetal.,1999),wherebyeachmalecopulateswithseveralfemalesateachoestrousperiod.Thefemalesmayinteractatahighratewithsuchamalewhenheispresent,andmayappeartobetryingtoprolongthemale’spresenceinthatpartofhishomerangethatencompassestheircorearea(Macdonaldetal.,1987).Outsidethebreedingseason,malerangesinsuchlow-densitypopulationstendnottooverlapagreatdeal,butinthebreedingseasontheirrangesexpandandoverlapconsiderably,astheytrytogainaccesstoasmanyreceptivefemalesaspossible.Thismaybeexaggeratedwhentheinterventionofmanreducesthenumberofmatingopportunitiesthroughtheneuteringoffemaleswithinamale’snormalhomerange.Yearlingmalesinthisenvironmentoftenstayclosetotheirmotherormaternalgroup,butastheygrowtheycomeunderincreasingattackfromoldermales.Intheirsecondorthirdyears,theyusuallydisperseawayfromtheirmother’shomerange.Individualmalesthatdonotemigrateappearnottobecomesexuallymature.Atthisstagetheymaybecomestronglysolitary,avoidingcontactwithallothercats,butmosteventuallycometochallengethesuccessfulbreedingmalesforaccesstofemales.Malesinruralpopulationsmaynotstartreproducinguntilaroundtheageof3years(Sayetal.,1999).

Wherecatdensitiesarehigh(upto3000cats/km2;Liberg,2000),suchasinurbanpopulations,territoriesareindistinctandoverlapping,encompassinglargemulti-male–multi-femalegroupsofcats(Natolietal.,2000).Here,eventhosemalesholdingterritoriesmaybeunabletomonopolizeallofthereceptivefemales,leadingtoapromiscuousmatingsysteminwhichbothmalesandfemalesmatewithseveralpartnersateachoestrousperiod(NatoliandDeVito,1991;Sayetal.,2002).Aggregationsofmalesaroundreceptivefemalesarecommonplacewherecatpopulationdensitiesarehigh,andtheirmostsurprisingfeatureisthatunderthesecircumstancesthemalesareoftenlessaggressivetowardseachotherthanwhenthereisnoavailablefemaleinthevicinity.Natolietal.(2007)pointoutthatthecostassociatedwiththeexclusionofallothermaleswouldbetoohighfortheresidentmale–whilehewasfightingwithonecompetitor,othersmightexploitthesituationandmatesuccessfullywiththefemale.Soheoptsinsteadforsharingthematingopportunitiesandthereforethepaternityoflitterswithothermales(Sayetal.,1999,2001).Malesindensepopulationsbeginreproducingassoonastheyreachsexualmaturity(Sayetal.,1999)andmaybelesslikelytodisperse.Devillardetal.(2003)observednoevidenceofmaledispersalatanyageover8yearsinthecolonytheystudied.Possiblythereducedinter-maleaggressionseeninhigh-densitypopulationsenablesjuvenilemalestoremain.

GeneticanalysisofpaternityinDevillardetal.’s(2003)studyshowedthathighlyinbredmatingoccurredinthatcolony,presumablydueinsomeparttothenon-dispersalofmales.Theredidnotappeartobeanydecreaseinlittersize,survivalprobabilitiesorbodyweightofoffspringasaconsequence,suggestingthatperhapsthereareonlylowcostsassociatedwithinbreedingindomesticcats(Devillardetal.,2003).Wherecoloniesarenotisolated,theremaybemoreoutbreedingthaninbreeding(Yamane,1998).

Reproductivesuccessinmales

StudiesofthematingsystemhaveadvancedconsiderablysinceMenotti-RaymondandO’Brien(1995)developedatechniqueusingDNAtodeterminethepaternityofkittens.Researchersarenowabletoaccuratelymeasuremalereproductivesuccessunderdifferentsocialandenvironmentalcircumstances.

Sayetal.(1999)analysedthepaternityofkittensbornintwocontrastinghabitats(ruralversusurban)withdifferentdensitiesofcats(234/km2versus2091/km2).Theyfoundahighrateofmultiplepaternityintheurbanpopulation,with70–83%oflittershavingmorethanonefather,whereasthiswasmuch

lowerintheruralpopulation,withonly0–22%oflittershavingmorethanonefather(Fig.8.2).Onemightexpecttheruralmalestoobtainexclusivesiringofkittens;however‘satellite’malesaresometimesabletocopulatewithfemalesintheabsenceoftheresidentmale(Sayetal.,2002).

Maleshavingaterritoryoverlappingthatoffemales,therefore,havevariedreproductivesuccessdependingonthelocalpopulationdensity.Inurbanareas,suchmalesdonothavecompletecontrolofreceptivefemalesorfullpaternityofsinglelitters.However,theydoachievethehighestreproductivesuccess:Natolietal.(2007)foundthattheresidentmalesiredthehighestpercentageofkittensbutmonopolizedonlyonewholelitteroutofnine,andco-siredthehighestnumberofotherlitters.

Maleswiththelargesthomerangesincludethemostfemalehomerangesandhavethehighestreproductivesuccess(SayandPontier,2004).Theymayalsosuccessfullyreproducewithfemaleswhosehomerangesdonotoverlaptheirs–inSayandPontier’s(2004)study,28%ofkittensweresiredbymaleswhoserecordedhomerangesdidnotoverlapwiththekittens’mothers’homeranges.Thisindicatesthatmalesmaymakequickexcursionsoutsidetheirrangestofindnewmatingopportunities.Thus,althoughrangingmorewidelydecreasestheprobabilityofsiringallthekittensinanysinglelitter(Sayetal.,2001),apparentlyusingalargeareaincreasesfertilizationsuccessbyallowingamalemorematingopportunities(Sandell,1989).Ingroup-livingsituations,therefore,amale’sabilitytomaintainalargehomerangemaybeoneofthemainkeystoimprovedmatingsuccessand,interestingly,hasbeenfoundtobeindependentofhisagonisticbehaviour,whichisconventionallyinterpretedasanindicatorof‘socialrank’(SayandPontier,2004).

Fig.8.2.Distributionofthenumberoffathersperlitterinaruralandanurbanpopulation(redrawnfromSayetal.,1999).

Bodyweighthasalsobeenshowntoaffectreproductivesuccess.Yamane(1998),studyingagroup-livingpopulationonasmallisland,identifiedtwodifferentreproductivetacticsaccordingtobodyweightofthemale.Heaviermalescourtedfemalesbothoftheirowngroupandofothergroups.Lighter-weightmalescourtedfemalesonlyoftheirowngroup.Observationsoftheextra-groupcourtingattemptssuggestedthismaybeafairlyunsuccessfultacticbytheheavymalesastheircourtshiprankwasreducedinsuchgroups.DNAevidence,however,revealedthatoverhalfthekittensfromgroupsoffemaleswerefatheredby‘extra-group’males.Yamanesuggeststhatthediscrepancybetweenobservedcopulationandactualpaternitymayindicatethatsomefemalechoicemaybeoperatingonthematingsysteminthatparticulargroupofcats.

Thequestionofwhetherfemalesexertchoiceoverwhichmalessiretheirkittenshasproducedequivocalresults.Natolietal.(2000)foundthat,althoughfemalescopulatedwithseveralmalesduringasingleoestrousperiod,therewasnoevidenceofmatechoicetobefound.Ishidaetal.(2001),however,foundthattheirfemalesdidnotacceptallmountingorcopulationattemptsandinparticularappearedtoavoidinbreedingwithclosekin(1/4orhigherdegreesofrelatedness).Moredistantrelativeswerenotrefused.

Oneotherimportantvariableaffectingthereproductivesuccessofmalesisthedegreeofoestrussynchronizationamongstfemales.Whenoestrusisasynchronousamorecompetitivemalecanattainhigherreproductivesuccessthanlower-rankingmalesthroughpriorityofaccesstofemales.Thisallowshimtocopulatewithmorefemalesortocopulatewiththesamefemaleseveraltimestoensurepaternity(Sayetal.,2001;SayandPontier,2004).Whenoestrusissynchronized,asisoftenthecaseindenseurbanpopulations,asinglemalewillnotbeabletomonopolizeallthefemalesandothermaleswillbeabletomatewiththemtoo.Sayetal.(2001)foundthatvarianceinmalereproductivesuccesswasfourtimesgreaterinthoseyearswhenfemalesbredasynchronously,withdominantmalessiringthehighestproportionofoffspring.

Infanticide

Whenonecoalitionofmalelionssucceedsinoustinganothergroupofmalesfromagroupoffemales,theyusuallykillallthecubsinthepride,therebybringingthelionessesintooestrusmorequicklythaniftheyhadcompletedlactationthroughnaturalweaning.Examplesofinfanticidebymaledomesticcatshavebeenreported(e.g.Macdonaldetal.,1987;PontierandNatoli,1999).Inthelatterstudy,sixcasesofinfanticideweredirectlyobservedinruralpopulationsofcats.Itisunclearjusthowcommonthisphenomenonis,althoughitmaybeonefactorcausingtheaggressionshownbynursingfemalestowardsstrangemales.Ithasnotbeendescribedforhigh-densitypopulations,andSayetal.(2001)suggestthatsynchronicityofoestrusinfemalesindenselypopulatedbreedinggroupsmayhelpreducetheriskofinfanticidethroughthemixingofpaternitythatensues(seesectiononreproductivesuccessinmales).Sincethebirthintervalinthedomesticcatisonly4months,comparedwith19monthsinthelion,theadvantagetobegainedfrominfanticideinbringingafemaleintobreedingconditionmaynotbegreat(Natoli,1990).Intemperateclimatesthesecond(autumn)litterrarelyproducessurvivingoffspring,butitispossiblethatitschancesofsuccessareincreasedbybringingthetimeofconceptionforwardbyevenafewweeks.Infanticidemightalsobeasuccessfulstrategywhencatdensityislowandadultfemales,foodorsuitablenestsitesareatapremium.Anon-breedingmalecouldenhancethechancesofhisownfutureoffspringsurvivingbyattemptingtokillalltheoffspringofthecurrentresidentmale,sothatwhenthatmalelosthisposition,andhetookover,hisownkittensmightbeborninthebestnestsites(asselectedbythesurvivingfemales)andbefedsufficientlywelltoreachadulthood.However,itisstillunclearwhetherinfanticideisacommonstrategyamongferalcats,oranaberrationonly

practisedbyaveryfew.ThisisdiscussedinmoredetailbyPontierandNatoli(1999).

SocialCommunication

Wehaveseenthat,wherethereisasufficientconcentrationoffood,catsformmoreorlessstablegroups,thebasisofwhichisusuallythecooperativerearingofkittensbyrelatedfemales.Theroleofmalesinthesegroupsisstillpoorlyunderstood,andmayvarywithpopulationdensity.Themembershipofthesegroupsisgenerallystableoutsideaswellasduringthebreedingseason,andasforothersocialCarnivora,thesocialstructureismaintainedbya‘glue’ofinteractivebehaviourpatterns.Inotherspeciesthesepatternshavebeeninterpretedasindicatingsomekindofhierarchicalorganization,andanindividual’spositioninthehierarchyisusuallyestimatedbythebehaviourpatternsitexhibitstowardsothermembersofitsgroup.Someofthemostrevealingofthesepatternsarethoseindicatingsubmission,whereoneindividualsignalsthatitwishesto‘backdown’fromanencounter.However,nosuchpatternhasbeenidentifiedinthecat,whichtendstowardoffaggressiveapproacheswithdefensive,ratherthansubmissive,behaviour.Whereasconsistentpatternsofaggressiveinteractionhavesometimesbeenidentifiedbetweenmales,andinterpretedasa‘dominancehierarchy’,thecollaborationbetweenfemalesismorecooperativethanhierarchical,becauseitiscommonforallthefemalesinagrouptobreedsimultaneously.Certainbehaviourpatternsappeartostrengthenthebondsbetweenindividuals,andtobuildupagroupidentity.Thesesocialbehavioursincludescentmarking,mutualgrooming(allogrooming)andmutualrubbing(allorubbing),andthesewillbediscussedinturn.Toputtheseintoasocialcontext,theflowofinteractionswithinsocialgroupswillbedescribedfirst.

Inanumberofpublishedstudies,notonlyhasthebehaviourofparticularindividualsbeenrecorded,butalsothepartnerstheychosetointeractwith,whetherthoseindividualswereconsideredseparately,ascloserelativesorasrepresentativesofaparticularage/sexclass.Incoloniesthatobtainsomeoftheirfoodbyhunting,thecatstendtovisittheircoreareaindependently,neitheravoidingoneanothernortendingtoalwaysbepresentwithaparticularpartner.However,whentheyarepresenttheytendtosingleoutparticularpartnersforinteraction.Forexample,Wolfe(seeCurtisetal.,2003)studyingtwocoloniesfoundthatsomeadultcatdyadswerewithin1mofeachothermoreoftenthan

wouldbeexpectedbychance–thesehereferredtoaspreferredassociates.Proximitywasnotdependentonlocation,socatswerenotsimplyaggregatingatpreferredresourcesatthesametime.Somecatsappearedactivelytoavoidoneanothertoo–inacolonyofintactcatstheseweremainlymale–malepairs,whileinaneuteredcolonytherewasnoeffectofgender(Wolfe,2001;seealsoCrowell-Davisetal.,2004).Macdonaldetal.(2000)discussinmoredetailthepreferencescatshadforproximitytobothkinandparticularage/sexclassesforthecatsinthreestudycolonies.Moredetailedexaminationofthequalityofsocialbehaviourinsuchcoloniesconfirmsthattheinteractionsarehighlystructured,andthecoloniesarefarmorethansimpleaggregationsaroundfoodsources.

Tailup

Catsusuallyprecedeamicableinteractionsbyraisingtheirtailtothevertical.Inferalcats,tail-upprecedesbothsittingdownwithanothercatandrubbingonanothercat(seeFig.8.3).Petcatsapproachatail-raisedsilhouetteofacatfasterthanonewithitstaildown,confirmingthattheverticaltailsignalsanintentiontointeractamicably(BradshawandCameron-Beaumont,2000).

Allogrooming

Catsspendagreatdealoftimegrooming,andthereisnoevidencetosuggestthatasolitarycatisanylesscleanthanacatthatisgroomedbyothers.Thefunctionofallogrooming,whichismostoftendirectedtotheheadandneckareaoftherecipient(VandenBos,1998),isthereforelikelytobeprimarilyasocialone,exceptinthecaseofyoungkittensthataregroomedbytheirmotherbeforetheybecomecompetentatgroomingthemselves.Inonestudyofallogroominginanindoorcolonyofneuteredadults(VandenBos,1998)theflowofallogroomingboutswithinpairsofcatswasasymmetric,withcatsshowingmoreaggressiongroomingratherthantheotherwayround.Groomingwasalsoassociatedwithaggressivebehaviourin35%ofinteractions(withgroomersshowingaggressivebehaviourmoreoftenthangroomees).Othercolonystudiesalsoindicatethatthehigherthedensityofcatsthelessaggressionandthemoreallogroomingoccurs,suggestingthatallogroomingmaybeawayofredirectingpotentialaggressionandreducingtensionbetweencatslivingtogether.AlthoughVandenBoscouldnotdetectanyeffectofdegreeofrelatednessonfrequencyordurationofgrooming,Curtisetal.(2003)studyingadifferentneuteredcolonyof

catsfoundkinpreferenceinbothallogroomingandproximity.

Fig.8.3.Aclassificationofsocialbehaviourpatternsperformedbyneuteredcats(i.e.excludingsexualbehaviour).Patternsthatareverylikelytobeperformedbythesamecatduringasingleinteractionwithanothercatinitsownsocialgrouparejoinedbysolidlines.Thepatternsfallintoclusters,mostofwhichhaveself-evidentfunctions.Amicableinteractions(top)oftenbeginwiththetail-upsignal(centre)eitherwhenthecatisstationary(tailup)ormoving(TR):thesefallintotwotypes,oneconsistingofallorubbing(topleft)andtheothermutualsniffingandgrooming(topright).Atthebottomisadefensivecluster,andaboveitaclusterofaggressivepatterns,linkedtogetherbychase/cuff.Ofthepatternsnotstronglylinkedtoanyofthesegroups,miaowmaybeanattemptbyonecattoalertanotherthatitisfollowingwiththeintentionofinitiatingaboutofrubbing;pawmaybeanattemptatinitiatinganinteractionwhichtriggersaggressionintheothercatandisfollowedbyrollandthendefensivebehaviour;follow/sniffrear(onlyweaklylinkedtootherpatterns,shownbybrokenlines)maybefollowedbyamicablesittingtogetherorbyaggressivebehaviour,presumablydependinguponthereactionofthecatbeingsniffed.(DatacollectedbySarah

BrownandanalysedbyCharlotteCameron-Beaumont,derivedfrom2044interactivesequencesbetween42neuteredcatsinthreepermanentgroups.Solidlinesrepresentpositive2×2associations;P<0.001bychi-square.)

Allorubbing

Thereisincreasingevidencethatoneofthekeybehaviourpatternsthatcementsanexistingcatgrouptogetherismutualrubbing,inwhichpairsofcatsrubtheirforeheads,cheeks,flanksandsometimestailstogether.Theremaybetwocommunicativeaspectstothisbehaviour,onebeingthetactilesignalsexchangedandtheotherthepotentialmixingofthetwocats’individualscents.Sofarneitheraspecthasbeeninvestigatedseparately,sothatforthepresentthefunctionofthisdisplayhastobededucedfromthebehaviouroftheanimalsthatinitiateit,andthebehaviourpatternsthatprecedeandfollowitinthecourseofasocialinteraction.

Severalstudieshaveshownthatrubbingflowsmarkedlyasymmetricallybetweenmembersofacolonyofcats,withfemalesinitiatingrubbingmorethanmales,andyounganimalsinitiatingmorerubsthanolderones(Fig.8.4).Forkittens,rubbingmayindicatethestrengthoftheirrelationshiptowardsthelactatingfemalesthatsucklethem,becauseacloserelationshiphasbeenfoundbetweenthenumberoftimesthateachkittenrubsonaparticularfemaleandthenumberofnursingboutsitreceivesfromthatfemale.Itisunclearwhichparticipantinitiatesinteraction–doeseachfemalepermitakittentosuckleinproportiontothenumberofrubsshehasreceived,ordoeseachkittenrubindirectresponsetobeingnursed?Whicheverwayroundthisis,rubbingdoesseemtohavespecialsignificanceasfarasnursingisconcerned;forexample,thelevelsofgroomingandnursingthatfemalesgivetoindividualkittensinapooledlitterarenotcloselyrelated.

Fig.8.4.Theflowofallorubbingbetweentheage/sexclassesoffarmcatsinabreedingcolonyatBarleyparkFarm,Oxfordshire,UK.Theproportionsofinteractionsinvolvingrubbingareindicatedbythewidthofthearrows(betweenage/sexclasses)andtheradiiofthecircles(withinage/sexclasses).Smallcirclesareshowninsidesquares,forclarity.Theproportionsarenotcorrectedforthenumbersofindividualcatsineachage/sexclass;averagenumbersareshowninbrackets.AM,adultmales(13);AF,adultfemales(33);JF,juvenilefemales(9);JM,juvenilemales(6);KI,kittensofbothsexes(12)(fromunpublisheddatacollectedbyWarnerPassanisiandDavidMacdonald).

Thesignificanceofallorubbingbetweenadultscanbeexploredfurtherbyexaminingthebehaviouralcontextinwhichitoccurs.Twostudiesofneuteredferalcoloniesshowedthatanindividualthatisabouttorubalmostalwaysraisesitstailasitapproachestheothercat(Brown,1993;Fig.8.5).Theformoftherubbingitselfdependeduponwhetherthereceivingcatalsoraiseditstail.Ifitdid,bothcatsusuallyrubbedsimultaneously;ifnot,therecipientcatsometimesonlyrubbedaftertheinitiatorhad,ornotatall.Femalesandmalesmayuserubbingdifferently:inaneuteredcolonyof(presumably)relatedcats,CafazzoandNatoli(2009)foundthatwhiletailupandrubbingweregenerallydisplayedbyfemalestowardsmales,sniffingnosewasmoreoftendisplayedbymalestowardsfemales.However,BarryandCrowell-Davis(1999)didnotobserveanyallorubbingin20neuteredpairsoffemaleindoorcats,possiblybecausethe

majorityofthesepairsconsistedofunrelatedindividuals.

Fig.8.5.Typicalsequencesofbehaviourthatcontainheadorforeheadrubbingbetweencats.Cat1approachesCat2withitstailraised(TA).IfCat2doesnotraiseitsowntail,Cat1rubsitshead(RH)onCat2,whichmayreciprocatebeforeCat1rubsitsflank(RF)onCat2.IfCat2doesraiseitstail(TU)orapproachesCat1withitstailraised,theysimultaneouslyrubheadsorforeheads(HF)together,beforerubbingflankstogether.Widthsofarrowsareproportionaltofrequencies(exceptforthoseemanatingfromRF).Alltransitionsderivedbyfirst-orderMarkovchainanalysis,excludingthosewithprobabilitiesworsethan0.001bychi-square(Brown,1993).

Anthropomorphically,rubbingseemstobehighlyaffectionatebutitappearstobeusedpredominantlywhentherelationshipissomewhatone-sided,andmaybethenearestthecathastoabehaviourpatternthatisusedbetweenindividualstoreduceconflictthroughreinforcementofaffiliativebonds.However,itisunliketheappeasementbehavioursshownbyothersocialcarnivorestoavoidescalatingaggressionduringinteractions,sincerubbingisrarelyseenininteractionsinvolvinganykindofovertaggression.Exceptionstothishavebeenrecorded,aswhenaparticulartomcatwasoccasionallymildlyaggressivetowardsafemalethatpersistentlyattemptedtorubagainsthim(Macdonaldetal.,1987),buteveninthiscasetherewasnoevidencetosuggestthatthefemale

rubbedtoappeasethetom.

Scentmarking

Socialodoursfeatureprominentlyinthelivesofmanymammalianspecies.Scentscanbespecifictoaparticularindividual,arefairlystablewithtimeandoffertheconsiderableadvantagethattheycanbedepositedintheenvironmentandlaterdetectedanddecodedbyaconspecificintheabsenceoftheemitter.Thescentsknowntobeusedasmarksbycatsincludethosecarriedinandbytheurineandfaeces,andthoseoriginatinginskinglandsonthehead.Urinescentmarksareknowntoconveyindividual-andgroup-specificinformation(describedinmoredetailinChapter5,thisvolume),althoughthewaythisinformationisusedinsocialinteractionsisuncertain.Malecatsfrequentlysprayurinewhenconsortingwithanoestrousfemale,anditispossiblethattherateofsprayingisanindicatorofmatingsuccess.Anypossibleroleofthisscentmarkingintheselectionofsexualpartnerseitherbythemaleorthefemaleremainsunclear.

SocialStructure

Thecommunicativerepertoireofthedomesticcat,althoughnotascomplexasthatofthemostsocialcarnivores–thewolf,forexample–neverthelesssuggeststheexistenceofasocialsystemintowhichallsocialinteractionsshouldfit.Thereissignificantdisagreementastohowthatsystemshouldbeconceptualized.Theearlieststudiesofcatsociality,performedonlaboratorycolonies,usedthedominanceor‘peckorder’concepttoderivehierarchies,basedonpriorityofaccesstoresourcessuchasfood(e.g.Winslow,1938).However,whenbehaviouralecologistsbegantostudyfree-livingcatcolonies,theycouldfindlittleevidencefordominancehierarchies,apartfromoldermalespreventingyoungermalesfrombreeding:catsocietyappearedtorevolvearoundcoalitionsbetweenfemales,whichwereusuallyegalitarian(Macdonaldetal.,1987).Bothoftheseapproacheshavebeenusedintheinterpretationofsocialinteractionswithinmulti-cathouseholds:someauthors(e.g.Crowell-Davisetal.,2004;CafazzoandNatoli,2009;Fig8.6a)haveespousedthehierarchy,whileothers(e.g.Rochlitz,2005b)haveexpresseddoubtsastowhetheritisusefultoapplythisapproachtopetcats.Theusualalternativeapproachinvolvesfocusingonthewaythateachcatusesthespaceavailable(BernsteinandStrack,

1996;BradshawandLovett,2003;Fig.8.6b),basedontheideathatcatsarefundamentallyterritorialanimals.

Fig.8.6.Contrastingapproachestothecharacterizationofsocialstructureinmulti-cathouseholds.(a)Adominancehierarchybasedonagonisticinteractions(fromKnowlesetal.,2004);eachboxrepresentsadifferentcat(N=19outof27catspresent)andthenumbersindicatedominancerank,suffixedbylettersindicatingcatsofequivalentrank.(b)Favouredrestinglocationsfor13cats(fromBernsteinandStrack,1996);mostcatshadseveraloftheseandmanyweresharedbetweentwoormoreindividuals.

Dominancecanbealooselydefinedconcept(Drews,1993)andalsoonethatmaymeanmoretothehumanobserverthantotheanimalsthemselves(Appleby,1993).Inanygroupofanimals,itisusuallypossibletomeasureconsistentasymmetriesintheexchangeofcompetitivebehaviourbetweenpairsofanimals.Combiningthesedyadicinteractionsintoanoverallstructurecanrevealapparenthierarchies–forexample,amoreorlesslinearhierarchyinwhichoneindividualisdominantoverallothersinthegroupand,atthelowerend,oneormorearesubordinatetoallothers.Robustmathematicaltechniquesnowexistfortheconstructionofsuchhierarchies(e.g.Bangetal.,2010),butitisoften

unclearwhethertheseconstructsreflecteverythingthatisofimportancetotheanimalsthemselves,orwhethertheyareanartificialconstructcreatedbythehumanobserver.Firstofall,the‘hierarchy’canchangemarkedlydependingonwhichbehaviourpattern(s)areusedtoconstructthedominancerelationships(e.g.Natolietal.,2001),andthecontextinwhichinteractionsareobserved.Secondly,thefactthatahierarchymaybeapparenttothehumanobserverdoesnotmeanthatitisalsoapparenttothecatsthemselves,orthatpositioninthehierarchyissomethingtowhicheachcataspires.Inotherwords,somegroupsofcatsmaybehaveasiftheyinhabitahierarchy,butareinfactsimplyreactingtothecatsaroundthemintheirownindividualway,orasaconsequenceoflearningfrompreviousencounterstheoptimalmannerinwhichtointeractwithothers.However,thecatsthemselvesareunlikelytobeaimingtoachieveaspecies-specificstructure,adoptinganyparticular‘role’withinthatstructure,ordeterminingbehavioursbasedonchangingtheirrolewithinthestructure.Indeed,becausetoday’sdomesticcatsareonlyafewthousandgenerationsremovedfromtheirsolitaryterritorialancestor,andhuntingremainsasolitaryactivity,itisdifficulttoseehowaspecies-specificsocialstructurecouldhaveevolved,noritsvalueindeterminingsocialbehaviour.

Theapparenthierarchiesthatcanberecordedfromcatskeptinhigh-densitycoloniesmaysimplyreflectstressbroughtonbyovercrowding:ithasbeensuggestedthatthe‘lowest-ranking’catsinthesegroupswouldnormallydisperse(DurrandSmith,1997).Inferalbreedingcolonies,cooperationbetweenfemaleshasbeendescribedas‘centripetal’andnot‘hierarchical’(Macdonaldetal.,1987),butfunctionalhierarchiescanoftenbederivedamongmales,inwhicholdermalesharassyoungermales,especiallyjuveniles,therebypreventingthemfrommating.Adultmalecatsrarely,ifever,cooperate,andthishierarchyappearstobeasimple‘peckingorder’inwhichyoungermaleslearntoavoidolder,moreexperiencedmaleswithwhichtheycannot(yet)effectivelycompete.Although‘dominant’residentmalesachievemorematingsthan‘subordinate’residentmales,doubthasbeencastonwhetheroldermalesactuallyhavemorereproductivesuccess(SayandPontier,2004):femalesshowstrongpreferencesfornon-kinmales(Ishidaetal.,2001),resultinginmany–evenamajorityof–kittensbeingfatheredbymalesfromoutsidethecolony(Yamane,1998).Thusthemostsuccessfulstrategyforamalecatwouldappeartobetovisitanyneighbouringcoloniesasoftenaspossible.Onsuchvisits,evenmalesthatappeardominantintheirowncolonybehavelikesubordinates,suggestingthat‘dominance’isnotanindividualcharacteristicbutaconsequenceofthelearntinteractionswithinfamiliargroups.Inmulti-cathouseholds,evenneuteredcats

canbehaveina‘bullying’way,thusaffectingthemovementoractivityoftheothercats(seeChapter11,thisvolume).However,thebehaviourofthemajorityofcatsinsuchagglomerationsappearstoberegulatedbymutualavoidance,time-sharingand,forsomepairsof(especiallyrelated)individuals,activeaffiliation(BernsteinandStrack,1996),possiblyderivedfromnormalfemale–femalebehaviour.

Mostofthebehaviourpatternsthathavebeenputforwardasindicatorsofdominanceandsubmissionarepartofnormaloffensiveanddefensivebehaviour,asoccursinothercontexts,forexamplebetweenunfamiliarindividuals.Insomebutbynomeansallcolonies,oldermalesmountyoungermales,butitisnotclearwhetherthisisdominance-assertionormerelyanoutletforsexualtensionintheproximityofunreceptivefemales(Yamane,1999).Feldman(1994b)hasreportedjuvenilemalesinonecolonyperformingtherollingbehaviournormallycharacteristicofoestrousfemales,apparentlyasasubmissivebehaviourtowardsoldermales,butthisdoesnotseemtooccuruniversally.Macdonaldetal.(1987)speculatedthatrubbingbehaviourmightbeanindicatorofsubordinatestatusinlargecolonies,butthishasyettobeconfirmedbysubsequentobservations.Overall,itappearsthattheritualized(i.e.evolved,species-typical)signalsdescribedinotherspeciesasdeterminingdominancerelationshipsaremissingfromthedomesticcat’srepertoire.Thisisconsistentwiththeideathattheyhavenospecies-typicalsocialstructureotherthanwhateveremergesasaconsequenceofeachcatlearningaboutthelikelyresponsesofothersunderdifferentcircumstances.Inthisconception,Macdonaldetal.’s‘centripetal’systemcanbethoughtofasarisingfromthepersistenceintoadulthoodofnormalaffiliativerelationshipsbetweenmothersandtheirfemaleoffspring.

TheFunctionsofDomesticCatSociality

Therehasbeenagooddealofspeculationovertheadvantagesthatcatsmightgainfromlivingingroups,buttodatenotagreatdealofevidencehasbeenfoundforanyofthealternatives.Giventhewiderangeofgroupsizesandthehighlyartificialsurroundingsinwhichsomeofthelargestgroupsfindthemselves,itisprobablynotsurprisingthatafunctionalexplanationthatappearstoapplytoagroupinonetypeofsituationdoesnotapplyinanother.Certainlyagreatdealofcareisneededwhenextrapolatingfromthesocialityofdomesticcatstothatofwildfelids.SocialstructuresinwildspeciesofCarnivora

havebeenrefinedbymillionsofyearsofnaturalselection;notonlyisFeliscatusonlyafewthousandgenerationsold,butthecircumstancesunderwhichithasevolvedhavenotbeenconsistent,asthenichessuppliedbyman’sactivitieshavecontinuallychanged.

Torecap,cooperationappearstorevolvearoundtheactivitiesoffemalesattemptingtopooltheirabilitytodefendresources,andtherebyobtainmaximumbenefitfortheiroffspringfromtheconcentratedfoodsourcethathasallowedthecolonytobecomeestablished.Whenallthefemaleswithinagrouparecloselyrelated,andtheyusuallyare,thiscanbeextendedtoincludetheirrelatives’offspring.ThebestevidenceforthiscomesfromthemuchpoorerbreedingsuccessofperipheralfemalescomparedwithcentralfemalesatHorspathFarm(KerbyandMacdonald,1988).Atthatfarmthiscrucialspatialstatuswasdeterminedbykinship.Thetwocentrallineageshadmuchhigherbreedingsuccessthantheperipherallineages(eventhoughsomecentralindividualswouldoccasionallybreedinperipheralsites),whichconsistedoffourside-branchesoftheoriginalcentrallineagesandtwolineagesbasedonimmigrants.Afterseveralgenerations,theseperipherallineageswouldprobablydieout,oremigrate;meanwhile,ifthecentrallineagebredsuccessfully,itwouldtendtofragmentandpushsomeofitsfemalestoperipheralpositions.Inpractice,thisprocessislikelytobedisturbedbyman’sactivities(culling,thetakingofkittensforpets,changesintheamountoffoodandshelteravailable),andsoisunlikelytobereadilyapparentinallcolonies.

TheOriginsofSocialityintheDomesticCat

Sincetherearenopublishedaccountsofthebehaviourofgroup-livingF.silvestrisapartfromthedomesticcat,wecanonlyspeculateastohowthesocialityofwhichthisspeciesisevidentlycapablearoseinthefirstplace.Virtuallyallthesocialgroupsthathavebeenstudiedhavereliedonconcentrationsoffoodsuppliedbyman.Thisraisesthepossibilitythatsocialityinthedomesticcathasarisensecondarily,asaby-productofdomestication;F.s.lybicaisessentiallysolitaryandterritorial.IfweassumethatF.s.lybicaoriginallyexploitedanthropogenicenvironmentsasacommensal,andthatthosenichescontainedreliablesourcesofpreythatcouldfeedmorethanonequeenandlitter,thenindividualanimalscapableofformingcoalitionswithkinmightout-competeindividualsthatcouldnot.Thiswouldpavethewayfortherapidevolutionofcooperativebehaviour.Subsequently,asthecatwasdomesticated,

manmighthavedeliberatelychosenindividualsthattoleratedthecloseproximityofothercats,becauseonehighlyterritorialcatwouldnothaveachievedthedesiredeffect.Lateron,thosecatsthatalsodisplayedaffiliativebehaviourtowardspeoplemighthavebeenselectedfromtheoriginal,conspecific-tolerant,population.Theaffiliativebehavioursthatadultcatsdirecttowardseachotherandtowardspeoplemayhavebeenderivedfromthoseshownbykittenstowardstheirmothers,carriedintotheadultstatebyaprocessofprogressiveartificialneotenization.