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^3=Wff3==Gk3=33ffffffffff==kkkpzp3fpzzpzzzp=3=kf3fpfpf=pp33f3ppppGf=pfff\G3Gk====I==pffffffzfzfzfzf33333333pppppppppzffpppzfpzpfffffffpzfzfzfzfpppppppp33333ffp3p3p3p3p3ppppppGGGzfzfzfzfp=p=p=ppppppzfp\p\p\pp3pGzfp=zfzfpppzf3pN@3ff=pfffffDCf=f=,=\\f==ff3\k='kkkkq@@@@@@@@@33ppzf"u ^+22PA#522222222225557XAAAA<7FA2A7KAF?@ABCDEFGHIJKLMNOPQRSTUVWXYZ[\]^_`abc  u!7d7  TR 6&A4 (Portrait)3'6&A4 (Portrait)  6&TUKUS&R&R&R&{&&R  W RR &Q&{&&R  Q & &Q7&&dd7@p`ReviewofUpperPleistocenemammalianassemblages  @`inCroatiancaves# &Q&S  Qa##&R&& &QB# &&{&&RԈ  ,  &Q&g& &&_&& &Q_&&_@ u`byNevenkaKrklec&GordanaJambreia,  #&_&+_## &Q&&&_#Ԉ 8X &I&& &Q@2 e`&&! &&I &&InstituteofPaleontologyandGeologyofQuaternary,CroatianAcademyof     SciencesandArts,10000Zagreb,A.Kovaciaa5,Croatia#&& ##&I& &&#   &&! &&IAbstract#&I&g& &4# h  & &! &&I?&&  DatabasecollectedfromthepublishedliteratureshowsthatthemaincharacterofUpperPleistocenemammalianassemblages @ ` foundinCroatiancavesiscommonpresenceoflargecarnivoretaxaandoverwhelmingabundanceofcavebears.ThepaleoecologicalinterpretationoftheseassemblagesyieldedaconclusionthattheconditionsweremoresevereduringtheEarlyUpperPalaeolithic,andthatseveraltaxacannotbeusedasenvironmentalindicators.Atthesametime,thefrequencyofarchaeologicalassemblagesdominatedbycavebearcanbeusedasameasureofintensityofhumansettlement:beardominatedassemblagesaremostcommonduringtheEarlyUpperPalaeolithicandhumanoccupationintheareawasrelativelyscarce.Unfortunately,faunaldataisgreatlyreducedinvalueduetotheabsenceoftaphonomicinformation,andmostofthePalaeolithicsitesneedtobereanalysed. Q  &Q&  QZusammenfassung   & && &Q?&&  DieinderLiteraturverffentlichtenDatenzeigen,da dasHauptmerkmalderGemeinschaftenderSugetieredesOberen   Pleistozns,dieinderkroatischenHhlengefundenwurden,sehrgro eAnzahlvonCarnivorenunddasberwiegendesHhlenbaren,bzw.seinerFossiliensind.DiepalokologischeInterpretationderGemeinschaftenl tdaraufschlie en,da dieBedingungenwhrendderEUP(deslterenPalolitihikums)verschiedenwaren,sodasdieverschiedenenArtenalsUmweltindikatorennichtverwendetwerdenknnen.GleichzeitigkanndieHufigkeitderpalontologischenGemeinschaften,indenenderHhlenbarvorherrscht,alsMa stabderIntensittdermenschlichenAnsiedlungverwendetwerden:dieBrengemeinschaftensindwhrendderEUPamhufigstenunddiesesGebietwurdevonMenschensehrdnnangesiedelt.LeidersinddieFaunadatenhauptschlichinfolgederfehlendentaphonomischenInformationenvermindertundderGro teilderpalolitischenFundortesolltenocheinmalanalysiertwerden. Q  &Q&  Q1.Introduction  & && &Q?&&  SpecificgeologicalframeworkofCroatianland,whichincludesabout75%ofcarbonaterocks,hasbeenthereasonfordevelopment  ofundergroundkarstmorphology,likecaves,shaftsandpits.Basedonmanyyearsofexploration,morethan3500speleologicalobjectshavebeenregistrated.About80localitieshavebeeninvestigated,withtestingsondesandpreliminaryanalysisoffauna.Excavationandpaleontologicalandchronostratigraphicalelaborationofthesitehasbeencarriedoutatsevenlocalities(Velikapeaina,Vindija,Veternica,Cerovackepeaine,Kupiaipeaina,andaljaandKopacina)(Figure1).Fossilassemblagesassociatedwitharchaeologicalfindingshavebeenass%  ignedtothedifferentfourchronologicalperiods(afterGamble,1986):middlePalaeolithic(MP,cca3500050000),earlyupperPalaeolithic(EUP,cca3500020000),lateupperPalaeolithic(LUP,cca2000010000)andMesolithic(MES,cca100007000)(Table1).s%*?/+p| 0 M `%E`0s RRRo#Ro# Ro#Ro#RR Q  &Q&  Q =1]-= # Q& &Q##  Q#    *h dh,// ,j//#,//",// ,dd ,// ,(dd ,//"+    SITE )Dd )m/sl )Dd )COORDINATES )Dd )CHRONOLOGY )Dd )LEVEL )Dd )DATE )Dd )PERIOD )Dd )REF.No "Dd "VINDIJA  q 275  q 461912N q 164438E   * RISSHOLOCEN  q EFdFd/dG1G3K  n 18500300 q 26600900  * 26970623  33000400 <\ 42400430   114100  n MP,EUP,LUP,MES   * 1  q VELIKAPE`INA  4 T  428     461710N    160222E  4 T  RISS/WRMHOLOCEN  4 T  egi     26450    31168 4 T  33850     MP,EUP,LUP     2     VETERNICA    306    455036N   163234E    RISSHOLOCEN    cij  , L  13660   >4320010080000  , L  MP,EUP    3    CEROVAbKEPE`INE    1404  Yy  441624N Yy  165348E    WRMHOLOCEN  Yy  c  Yy  >40000  Yy  EUP  Yy  4,5  Yy  KUPI`IPE`INA    865  ?  451212N ?  141022E    WRM?  ?    ?    ?    ?  Ѐ6  ?  ANDALJA   72   445257N  135348E  ~ WRMHOLOCEN   B/gB/sB/dB/CC/sC/d   1083050  12320100 ~ 10500100 7 13070230  13130230 Ii 20750400   EUP,LUP,MES   Ѐ7   KOPAbINA  / 280  / 4322N / 1632E    WRMHOLOCEN  / C1AD1C1    11850220 / 12935250    LUP  / Ѐ7 /  ,# Tabele1:UpperPleistocenelocalitiesinCroatia#,%"# (H#  Q  &Q&  Q2.UpperPleistocenemammalianassemblages# Q& &Q"##  Q"# Mm&   Collectedfaunaisgroupedaccordingtothewayofnourishment(Carnivora,HerbivoraandOmnivora)andthefrequencyoftheirappearenceisshownforeachgroup.Cavebearwereexcludedbecausetheyareeitherpredominantatsomelocalitiesorcompletelyabsentatother.Figure2.showsthatatthelocalitiesVindija,VelikapeainaandCerovackepeainethefrequencyofcavebearisover90%;atVeternicaandKupiaipeainaisover75%;atandaljaonly3%whileinKopacinathebearswerenotfound.Analogically,herbivoreandomnivorearemorefrequentinandaljaandKopacina,whilecarnivoraremainsdonotexceed20%oftotalfossilremainsatanylocality.Thereasonsforsuchdistributionoffossilremainsshouldbeeconsideredinseveralaspects.%  c%1!P ,  S2 c RRRo#Ro#K"k1RR K"k1RR K"k1RR     Fromthegeographicalaspects,localitiescanbegroupedintothreetypes.Vindija,VelikapeainaandVeternicawouldbelongtohilltypehabitat;CerovackeandKupiaipeainaaremountainhabitat;andandaljaandKopacinabelongtocoastaltypeofhabitat.  ThehilltypehabitatwasinhabitedbearbythewholeupperPleistocene.Accordingtothemodeloforiginforcaveassemblagewithpredominatingbearremainsonalargecarnivora,thesehabitatscorrespondtothecriteriaforcavelairs.InarchaeologicalassemblagescavebeardominationcanshowthesignificanceofPalaeolithichumanpresenceinthereareas.We,thereforeassumethatVindija,VeternicaandVelikapeainawereusedbyPalaeolithichuntersinperiodsofbearsabsence.  Inmountaintypehabitatscavebearpopulationdiffersfromthetypicalrepresentatives.Althoughtheyare50000yearsold,dentalanalysisdemonstratesimilaritywithmiddlePleistocenebearsfromVindija.Thisdifferencecanbeexplainedbygeographicalpositionwithdifferentpaleoclimaticandpaleovegetationalconditions.Lowerwintertemperaturesandmoresnowprovokedgrowthofdifferenttypesofvegetation(pineforestswithoakforestsrefuges)asbearnutrimentwhichprobablyaffectedtheevolutionofbearsteeth,andresultedwithuncompletedmolarisationofpremolars.Humanpresencehasbeendocumentedwithonetibiaandwithfewstoneartifacts.Thesecaveswereonlyhunterstations,notapermanentresidenceofupperPleistocenehunters(Malez,1975).  Aminimalfrequencyofcavebearsatcoastaltypelocalitiesmeansthatcaveswerenotusedaslairs.Therefore,omnivoresandherbivorescouldfindsheltersinsuchcaves,aswellasPalaeolithichumans.Themostfrequentanimalsweredeers,does,bullsand + K  horses.  Ithastobepointedoutthatmanyspecies,especiallychamoisandibexwhichwerefoundinallstrataatallsitesandofdifferentperiods,arenotgoodclimaticandecologicalindicators,sotheirpresenceinstratadoesnotnecessarilyindicatebadclimaticconditions.  NumerousprovesofcontinuouscolonizationbyPalaeolithichuntershavebeenfound.Besidethehumanboneremains,numerousboneandstoneartifactswerefound,niceformedfireplacesandbrokenPleistoceneanimalbones.Humanpresenceleadtotheconclusion thathumanswereresponsibleforanimalremainsinhabitats,butwecouldnotbesurehowgreatistheirimpactonthecompositionoffoundassemblages.  PaleontologicalinvestigationofthoselocalitiesinCroatiashowbigdiversityinpresencefrequencyofmammal.ThereasoncouldbeindifferentagesofupperPleistocenelayersanddifferentclimaticandvegetationalconditions,butananswertosuchaseriousquestioncannotbebasedonlyonthedescribedtaxa.Itisnecessarytoapplytaphonomicalanalysisandtoextendresearchondynamicalprocesseswhichparticipateintheoriginofpaleontologicalandarchaeologicalassemblages.SuchmethodsmustbeeappliedtoalllocalitiesinCroatia. 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