‘Neanderthal bone flutes’ : simply products of Ice Age spotted hyena scavenging activities on cave bear cubs in European cave bear dens


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‘Neanderthal bone flutes’ : simply products of Ice Age spotted hyena scavenging activities on cave bear cubs in European cave bear dens
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Royal Society open science
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Diedrich, Cajus G.
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Royal Society Publishing
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Cave bear ( lcsh )
Spotted hyena ( lcsh )
Animal remains (Archaeology) ( lcsh )
Glacial epoch ( lcsh )
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serial ( sobekcm )
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Europe -- Slovenia

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Abstract:
Punctured extinct cave bear femora were misidentified in southeastern Europe (Hungary/Slovenia) as ‘Palaeolithic bone flutes’ and the ‘oldest Neanderthal instruments’. These are not instruments, nor human made, but products of the most important cave bear scavengers of Europe, hyenas. Late Middle to Late Pleistocene (Mousterian to Gravettian) Ice Age spotted hyenas of Europe occupied mainly cave entrances as dens (communal/cub raising den types), but went deeper for scavenging into cave bear dens, or used in a few cases branches/diagonal shafts (i.e. prey storage den type). In most of those dens, about 20% of adult to 80% of bear cub remains have large carnivore damage. Hyenas left bones in repeating similar tooth mark and crush damage stages, demonstrating a butchering/bone cracking strategy. The femora of subadult cave bears are intermediate in damage patterns, compared to the adult ones, which were fully crushed to pieces. Hyenas produced round–oval puncture marks in cub femora only by the bone-crushing premolar teeth of both upper and lower jaw. The punctures/tooth impact marks are often present on both sides of the shaft of cave bear cub femora and are simply a result of non-breakage of the slightly calcified shaft compacta. All stages of femur puncturing to crushing are demonstrated herein, especially on a large cave bear population from a German cave bear den.
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rsos.royalsocietypublishing.orgResearch Citethisarticle: DiedrichCG. Neanderthalboneutes:simplyproductsof IceAgespottedhyenascavengingactivitieson cavebearcubsinEuropeancavebeardens. R.Soc.opensci. :. http://dx.doi.org/./rsos. Received:May Accepted:February SubjectCategory: Biology(wholeorganism) SubjectAreas: behaviour/palaeontology Keywords: Neanderthals,pseudo-boneutes,Late Pleistocenecavebeardens,hyenascavenging, toothmarks,femurdestructionstages Authorforcorrespondence: CajusG.Diedrich e-mail: cdiedri@gmx.net Neanderthalboneutes: simplyproductsofIceAge spottedhyenascavenging activitiesoncavebearcubs inEuropeancavebeardens CajusG.Diedrich PaleoLogic,PetraBezruce,Zdice,CzechRepublic CGD, ---.SummaryPuncturedextinctcavebearfemoraweremisidentiedin southeasternEurope(Hungary / Slovenia)as‘Palaeolithicbone utes’andthe‘oldestNeanderthalinstruments’.Theseare notinstruments,norhumanmade,butproductsofthe mostimportantcavebearscavengersofEurope,hyenas.Late MiddletoLatePleistocene(MousteriantoGravettian)IceAge spottedhyenasofEuropeoccupiedmainlycaveentrances asdens(communal / cubraisingdentypes),butwentdeeper forscavengingintocavebeardens,orusedinafewcases branches / diagonalshafts(i.e.preystoragedentype).Inmost ofthosedens,about20%ofadultto80%ofbearcubremains havelargecarnivoredamage.Hyenasleftbonesinrepeating similartoothmarkandcrushdamagestages,demonstrating abutchering / bonecrackingstrategy.Thefemoraofsubadult cavebearsareintermediateindamagepatterns,comparedto theadultones,whichwerefullycrushedtopieces.Hyenas producedround–ovalpuncturemarksincubfemoraonlybythe bone-crushingpremolarteethofbothupperandlowerjaw.The punctures / toothimpactmarksareoftenpresentonbothsides oftheshaftofcavebearcubfemoraandaresimplyaresultof non-breakageoftheslightlycalciedshaftcompacta.Allstagesof femurpuncturingtocrushingaredemonstratedherein,especially onalargecavebearpopulationfromaGermancavebearden..Introduction..FirstboneutedescriptionsTherst‘Neanderthalcavebearboneute’fromtheMiddle Palaeolithicwasbelievedtohavebeendiscoveredinthe 1920sfromPoto ckaZijalkaJamaCave(i.e.PotokCave), TheAuthors.PublishedbytheRoyalSocietyunderthetermsoftheCreativeCommons AttributionLicensehttp://creativecommons.org/licenses/by/./,whichpermitsunrestricted use,providedtheoriginalauthorandsourcearecredited.

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rsos.royalsocietypublishing.org R.Soc.opensci. :................................................ Figure. StudiedandreferredLatePleistocene(MIS…d)EuropeancavesiteswithPalaeolithiccavebearpseudo-boneutes,and comparedcavebeardenswithhyenainuence(hyenapalaeobiogeographyofsites[ ]).Slovenia[ 1 ].Thiswasalargercavebearden(cf.[ 2 , 3 ])andLatePalaeolithicAurignacian(not Neanderthal)usedrocksheltercampsiteattheentrance(cf.[ 1 ]; gure1 ).Inthiscave,cavebearhunts byCro-Magnonhumansseemtobeindicatedonacavebearshoulderbladepathology(largediagonal impacthole,partlyhealeddiagonalhole)thatseemstohavebeenmadebyaprobableMlade cprojectile bonepoint[ 5 ]. Othercavebearcubfemorawithholes(‘boneutes’)werethenreportedfromtheIstállóskö Cave,Hungaria(cf.[ 6 ]).ThiswasasmallercavebearandIceAgespottedhyena( Crocutacrocuta spelaea )carnivoredenwhichoverlapswithanotherAurignaciancampsite,butagain,ithasnoMiddle PalaeolithicNeanderthaloccupationsigns(cf.[ 7 ]). Brodar[ 8 ]reportedcavebearcubfemoraandothercavebearbones‘withholes’asfurtherproofofthe ‘oldestinstrumentsintheworld’fromtheMokriškaJamaCave(orMedvedjaJamaCave = BearCave), Slovenia.Also,thisisalargecavebeardenwhichhadagainanAurignaciancampsiteattheentrance, andagainnoNeanderthaloccupationatall(cf.[ 9 , 10 ]). Allaforementionedfemoraandothercavebearboneswith‘holes’(i.e.ribs,humeriandjaws) werecompiledandstudiedbytheethnologist / musicianOmerzel-Telep[ 11 ],withoutanynatural science,norpalaeozoologybackground,especiallytheimportantecologyofcavebearsandtheir predators/scavengers,non-humantoppredatorsoftheIceAgeandthewidedistributionofcavebear dencavesinEurope(cf.[ 3 , 12 – 21 ];gures 1 and 2 ),wherealwayslargeamountsofdamagedandalso

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rsos.royalsocietypublishing.org R.Soc.opensci. :................................................ Figure. Cavebearscavengingmodelsinlargercavebeardencaves(hereZoolithenCave,Germany)forallthreetoppredatorsthat hunted,killedandscavengedoncavebearsalloverEuropewithincavesinborealforestpalaeoenvironments.Thebonecrusherof longboneswasonlytheIceAgespottedhyena,whichproducedround / ovalpuncturemarksoncavebearcubbonesbythebonecrushing premolarteeth,i.e.boneuteholes(composedandadaptedfrom[ , , , , ];illustrationsG.Teichmann).puncturedcavebearbonesarepresent,suchasguredwithmanynewexampleshereinforthenorthern GermanWeißeKuhleCaveandothercavebeardens(gures 3 – 7 )...ThelongdiscussedSlovenianpuncturedcavebearcubbonendAnotherjuvenilebearcubfemurwithholesfromDivjeBabeICave,Slovenia,asmallcavebearden (cf.[ 25 ]; gure5 (4)),wherealsoNeanderthalMousterianlayerswerebelievedtobepresent[ 26 ],was

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rsos.royalsocietypublishing.org R.Soc.opensci. :................................................ Figure. Carnivorepunctureholesincavebearskulls,jawsandpostcranialbonescausedbytoppredatorcanineteeth(lions,leopards, hyenasandwolves),butaremainlyproductsatlongbonesandlowerjawsofthepremolarcrackingteethofhyenas(cf. gure ).()Cub skull(smallcavebearform U.spelaeuseremus )fromtheWeißeKuhleCave,Germany,whichwasscavengedstronglyontheleftside. Punctureholesareproducedbycanines(incranium),whereasthebreakageoftheleftmandibleistheresultofhyenapremolarcracking teeth.(a)Dorsal,(b)lateral,(c)detailoflateraltoothmarkholes(producedbycarnivorecanines,bestttingtohyenasorlions)(PAL collection).()Singleprobablycanineimpactofalargecarnivore(lion,hyena)onacubskull(largecavebearform U.ingressus )from theGroßeTeufelsCave,Germany.(a)Dorsal,(b)lateral,(c)detailoftoothmarkhole(GTCPcollection).()Mandible( U.s.eremus )from theWeißeKuhleCaveofacubwithhyenapremolarimpactholes(crackingpurpose).Suchmandibleswerecrushedalwayssimilarwith damagingtheramus,orakesofthelowerdistalmandible.(a)Lateralouterview,(b)lateralinnerview,(c…e)detailsofpunctureholes ofbothsidesand(f)rettingofthejawwithalltoothmarksofbothsidesprojectedinonelevelwhichtinonetoothmarkofthebone crushingteethoftheupperjawofahyena(allPALcollection).declaredtwiceincorrectlyasthe‘oldestinstrument’,a43140BPold‘Neanderthalute’fromlayer8 [ 26 , 27 ]( gure5 (4)).Thiswasalreadycontradictorytotheresultsofthearchaeologicalinventorythat iswellacceptablydeclaredtobesolelyof,again,Cro-MagnonhumanLatePalaeolithicorigin,andnot

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rsos.royalsocietypublishing.org R.Soc.opensci. :................................................ Figure. Carnivorepunctureholesincavebear( U . s .subsp.and U.ingressus )longbones(humerus,radius,tibia)andpelvicandpedal bonesbytoppredator(lions,leopards,hyenasandwolves)canineandmainlypremolarhyenateeth.(…)CubhumerifromtheWeiße KuhleCave,Germany.(…)CubradifromtheWeißeKuhleCave,Germany.(…)CubtibiaefromtheWeißeKuhleCave,Germany.() CubcoxafromtheWeißeKuhleCave,Germany.(…)CubandadultcalcaneifromtheWeißeKuhleCave,Germany(allPALcollection).ofMousterian(cf.[ 28 ]).TheAurignacianlithicmaterialappearsalsotogetherwithcavebearremains [ 25 ].Therefore,thereisnoevidenceforaNeanderthal(Mousterian)contextandthecavebearremains, whichevenoccurinseveralolderandyoungerLatePleistocenelayers(cf.[ 25 ]).Theonlyabsolutedate wasmadesolelyonacavebearbone,the‘boneute’,whoseagewoulddateintotheNeanderthalor ‘cavebearden’timeperiod.Thisreportofa‘cavebearfemurboneute’wasnotthe‘oldest’,neither historically,norbystratigraphy.Thebone’sholesonthedorsalsideappearnottolineup,whereason theventralsideanotherholewasdeclaredasthe‘thumbhole’.Thestudieseventhrilledupto‘exact musicalstudies’[ 29 ].Fink[ 30 , 31 ]declaredthentothetopofthis,withoutnaturalscienticstudies,that

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rsos.royalsocietypublishing.org R.Soc.opensci. :................................................ Figure. PseudoNeanderthalboneutesofdierentagedcavebear( U . s .subsp.and U.ingressus )cubfemora(lessthanyear individualage)fromvariousEuropeanlargecavebeardensites.()FemurfromMokrikaJamaCave,Slovenia(photosadaptedfrom [ ];NMLScollection).()FemurfromKepplerCave,Germany(photosadaptedfrom[ ];SMMcollection).()FemurfromSophiesCave, Germany(photosadaptedfrom[ ];SMMcollection).()FemurfromDivjeBabeCave„theNeanderthalboneuteholotype,Slovenia (photosfromNMLScollection).()FemurfromOaseCave,Romania(IRcollection).()FemurfromHermannsCave,Germany(photos adaptedfrom[ ];RCcollection).theholespacingmatcheda‘diatonicscalesequence,amongthemostwidespreadscalesknown’—which underlines,alsocontradictory,thatthisisnotofhumanorigin.Ethnologist / musicianscreatedthena waveof‘cavebearboneinstruments’basedsolelyon‘holesinbones’(compiledin[ 11 ]),fromallkinds ofcarnivorepuncturedcavebearbones,evenotherthanfemora.Otherauthorsdoubtedthe‘ute’and humanoriginhowever(e.g.[ 32 – 38 ])orwereghtingforpro-arguments(e.g.[ 39 , 40 ]).Atleast,very

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rsos.royalsocietypublishing.org R.Soc.opensci. :................................................ Figure. Continuousdocumentationofdestructionstagesofcavebear( U . s .subsp.and U.ingressus )cubfemora:(…)puncture,(…) part-ake,(…)fullbreakage-akes„allwithpunctureholesorhalfpreservedholesaftersplittinginakes„ofdierentagedcave bearcubfemora(lessthanyearindividualage)anddierentspecies( U.s.eremus and U.ingressus )„allfromtheWeißeKuhleCave, Germany(PALcollection).correctly,the‘holes’weremostlydiscussedtobeof‘carnivorechewingdamage’origin(cf.[ 32 , 33 , 37 , 41 ]), whereasX-rayscansdidnotproveany‘drill-scratchesaroundtheholes’oranymarksofstonetoolson thebones,andleftagainthequestionoftheholeoriginopen(cf.[ 42 ]).Theexactcarnivorewasnever estimated,evenbynewerandfullycontroversialstudiesbyTurk etal .[ 24 ]thatlackcarnivoreecology knowledge,especiallyintoothandjawfunctionoftoppredators.Ignoringthetoppredatorbonedamage onIceAgeanimalbones,againthepseudo-boneutewasnotonly‘conrmed’,evenmoreboneute ndswereaddedbythesameSlovenianauthor(cf.[ 43 ]),whomisidentied:(a)thesiteoccupationby Neanderthals,asthoseofAurignacians[ 28 ],(b)thebone,byrotatingitupsidedown(see[ 44 ]),the180rotationofwhichiscorrectedherein( gure5 a ),(c)thegeneralbonetaphonomyofcavebearbones,and (d)carnivorejawfunctions,especiallyhyenas,correctlypresentedherein( gure2 ).

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rsos.royalsocietypublishing.org R.Soc.opensci. :................................................ Figure. ( Captionopposite .)..HyenaandcavebearpopulationsoverEurope:specializedcavebearscavengesInthiscontribution,notonlysolecarnivoredamagecanbedemonstratedonallpreviouslypublished ‘pseudo-boneutes’,whichwerealreadyrevisedinsomecases[ 4 , 16 ]( gure2 ).Herein,evenmoreof suchcavebearboneswithholescanbeaddedwithfocusonlyonthefemora(gures 5 – 7 ),fromGerman andRomaniancavebeardensites(thereforenotlimitedtoSloveniaatall,asmentionedbyTurk etal. [ 24 ];seegures 1 , 5 – 7 and table1 ).Theirproducer,alargecarnivore,andthemainscavenger / bone

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rsos.royalsocietypublishing.org R.Soc.opensci. :................................................Figure. ( Opposite .)Examplesofthedestructionstagesoffemoraofcavebearcubs,subadulttoadultcavebears( U . s .subsp.and U.ingressus ).()Thisfemurofanadultcavebear( U.s.eremus )fromtheGroßeTeufelsCave,Germany(POcollection),isthebestproof forthehyenatoothmarkanddamageorigin,wheretwodiagonaltoothmarks(i.e.diagonalcut)canbereconstructed,andwherelower andupperjawpremolarteethandtheirantagonistictoothmarkimpactholestexactlytothehyenaskulldentition.Ahyenatriedtocut thedistaljoint.(a)Cranialview,(b)detailofthecranialtoothmarkholes,(c)caudalview,(d)detailofthecaudaltoothmarkholes,(e) reconstructionrettingoftheP-teethintothecranialandcaudaltoothpits,demonstratingexactttingandtwooverlappingdiagonal toothmarks(GTCPcollection).()Proximallychewedandpuncturedfemurjointofasubadultcavebear( U.s.spelaeus or U.ingressus ) fromtheWeißeKuhleCave,Germany.Theimpactmarksaretwotypes:(a)fullanddeepintothespongiosa,i.e.toothwithintactcrown tip);(b)roundsurfacebreakagesofcompacta,i.e.toothwithrubbedordamagedtiporslightimpact(PALcollection).()Cutofproximal joint( U.s.eremus )demonstratedatafemurfromtheKepplerCave,Germany,cranial(SMMcollection).()ShaftfromtheOaseCave, Romania,cranial(IRcollection).()Shaftofasubadult(largecavebear U.ingressus )withdistallycrackedparts(allfoundinthecave closetoeachotherwitholdfractures)fromtheWeißeKuhleCave,Germany,cranial(PALcollection).()Selectedfemurfragmentsofcub tosubadultcavebears( U.s.eremus and U . s .subsp.)partlywithspiralbreakage,andtoothmarkimpactmarksonthesurfacesfromthe WeißeKuhleCave,Germany(PALcollection).()Manyselectedfemurfragmentsofsubadulttoadultcavebears( U.s.eremus and U . s . subsp.)partlywithspiralbreakage,andtoothmarkimpactmarksonthesurfacesfromthePerickCaves,Germany(PCHcollection). destructoroftheIceAge,theIceAgespottedhyena Crocutacrocutaspelaea ,willbediscussedastheoval holeproducerherein( gure2 ),basedontheintensiveLatePleistocenecentralEuropeancavebearand toppredatorstudiesinandoutsidecavesofthepastyears(e.g.[ 3 , 12 – 16 , 18 – 22 , 51 , 54 , 55 ]).Thisresultsina differentviewpointofmodernzooarchaeology,multipleanimal/humanuseoflargercavesystemsand cavemodels( gure2 ).TheIceAgetoppredatorresearchinEuropefocusedthesepastyearsonhunting ofcavebearsinlargecavebeardens.There,damageoncavebearbonesisnowwellknownandreported inseveralpublications(e.g.[ 3 , 4 , 16 , 18 – 21 , 51 , 56 ]; gure2 ).Allformerarchaeological,ecologicalfocuscave bear‘boneute’studiesforgotallfourcavebearpredators—steppelions( Pantheraleospelaea ),leopards ( Pantherapardusspelaea ),IceAgespottedhyenas( Crocutacrocutaspelaea )andIceAgewolves( Canislupus spelaeus )—whichareknownnowtobecavebearkillers,andmainconsumersinmountainregions,where mammothsteppemegafaunawereabsent[ 4 , 18 – 21 ].Thesepredatorsspecializedinconsumingmainly (andespeciallyinwintertimesduringcavebearhibernation)cavebearsinborealforestmountain regions,butindifferentwaysandwithdifferentimpactonthecarcassesandbonedestruction(cf. gures 2 and 3 ).However,themain‘bonedestructor’isknowntobetheEuropeanIceAgespotted hyena[ 19 ]( gure2 ),withcavebearbonedamagerstunderstoodattheoverlappinghyenaden(cave entrance)andcavebeardenofthePerickCaves[ 50 – 52 ],withnewerproofatSophie’sCave[ 21 , 22 ],and Hermann’sCave[ 16 ]orZoolithenCave[ 18 ]andhereinbestdemonstratedandnewlyaddedforthe WeißeKuhleCave(gures 3 , 4 , 6 and 7 )..MaterialandmethodsFirst,fromtheliteratureavailable,‘boneutes’werecompiledandreinterpretedhereinwithnew drawings( gure5 and table1 ).Fromtheliterature,newinterpretationsweremadeofthesitesin thearchaeologicalcontent(NeanderthalversusAurignaciansites),andoverlapincarnivoredenuse (hyena / wolfden—alwaysatentranceareas)andidenticationassmalltolargecavebeardens( gure1 and table1 ).Severalcavebeardenswerestudied,aswellaslargerbonecollections( gure1 and table1 ). Tothosenon-Slovenian / Hungarian(whereboneuteswerethoughtonlytobefound)sitesbelongthe RomanianUr¸ silorCaveandOaseCave.GermansitesareHermann’sCave,PerickCaves,especiallya largepopulationofsmall( Ursusspelaeuseremus )andlargecavebears( Ursusingressus )andlargeamount ofmaterialindifferentdestructionstagesfromtheWeißeKuhleCave,butalsosomerelevantbones fromthedensitesKepplerCave,ZoolithenCave,Sophie’sCave,GroßeTeufelsCaveandtheCzech SloupCave. Generalcavebearbonedamagebylargecarnivores(lion,hyena,wolf)ispresentinallofthoselarge cavebeardens.Onaverage80%ofthecub,and20%oftheadultcavebearboneshavelargepredator damage.Thecomparisonfocusedonthepresence / absenceandpositionsofround–ovalpuncturemarks. Hereinthebonedamagestages1–3(1,chewedjoint;2,shaft;3,fragments)arepresentedindetailfor cavebearfemoraofcubs,subadultandadultcavebears.Thismaterialiscomposedandcomparedfrom theaforementionedpublicationstoendthelongdiscussionabout‘MiddlePalaeolithicNeanderthalbone

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rsos.royalsocietypublishing.org R.Soc.opensci. :................................................utesandoldestinstruments’.Toothmarktypes,shapesandespeciallytheirpositionsonbothsidesof theshaftends,orthemiddlepart,wereidentiedastheantagonisticupperandlowerjawtoothmarks ofhyenas. Furthermore,theavailablematerialwasstudiedonthetoothmarkmarginsandholesundera microscope,whichallowsidentifyinginarststepwithoutreectionelectronmicroscopyorX-ray photospossiblydrillorstonetoolscratchmarks.Nosuchcut / drillmarkswerefoundwithinthe hereinguredmaterial.Instead,stonetoolcausedcurvedcutmarkswerefoundonasinglecavebear femur( U.ingressus )fromthelatestLatePleistocene(MIS3cavebearlayers,alsoAurignacianperiod) ofHermann’sCave(cf.detailsin[ 23 ]).Otherbonesurfacedamagesintheformofbiteswereobserved ontwocavebearcubhumeri( U.s.eremus )fromSophie’sCave,butthosewerewellidentiedtohave resultedfromporcupine(rodent)incisiveteeth(cf.detailsin[ 21 ]). Finally,thepositionofthebitemarkholesandtheirorientationinovalpitsarecalculatedatsimilar moreorlesscylindricalinthemiddleoftheshaftformedlongbones:radius,femurandtibia. Thegured‘boneutes’ofDivjeBabeCave1,MokriškaJamaCaveandallotherSlovenian‘pseudoboneutes’arehousedintheNationalMuseuminLjubljana,SloveniaandHungary(NMLS).Oase CavematerialwasstudiedintheInstituteEmilRacovita,Romania(IR).WeißeKuhleCave(Germany) bonesareinthePaleoLogicResearchInstitute,CzechRepublic(PAL),PerickCavematerialiskeptin thecollectionofthePerickCaveclubhouseinHemer,Germany(PCH).Hermann’sCavematerialwas analysedinthecollectionoftheRübelandshowcaves,Germany(RC),KepplerCavematerialisinthe StatdmuseumMenden,Germany(SMM),andGroßeTeufelsCavebonesarekeptintheshowcaveofthe villageofPottenstein(PO)..Results..Bonedamagestages:cubandadultdierencesInarststage,oneofthejoints(damagestage1),andinasecondstep(damagestage2)theotherjoint wascutoffusingthescissor-dentitiononverysmall-sizedfemora( gure5 (1)),whichbecomesmore diagonal(i.e.diagonalcut)withincreasingfemursizes(gures 5 – 7 ).Thelargerthedistalfemurjointhas been,themorediagonalthiswascut.Cuttingoffthejointsisrecognizedinallageclassesofcavebears. Betweendamagestages2(cutting)and3(cracking),therearealreadydifferencesincubtoadultcave bearfemora(gures 6 and 7 ).Puncturemarksofpremolarteethareonlypresentincubfemurbones, owingtoslightphosphaticcalcicationoftheshaft.Insomecases,theseshaftsexpose,onbothsides, punctureholesofeachoftheupperjawP3andantagonisticlowerjawP3–4,sometimespartsofM1,which attributesitonlytothecrushingteethtriangleofhyenas(cf.gures 2 and 3 ).Instage3,subadultcavebear femoraalreadystartedtocrush,whichisdemonstratedfromatleastoneexamplefromHermann’sCave ( gure5 (6)).Thedetailcontinuousstagesofcubfemorapuncturetobreakingstagesaredemonstrated forthersttimehereinintheWeißeKuhleCavematerial( gure6 ),whereasbreakageismuchrarer insubadulttoadultfemora(gures 6 , 7 and 8 ).Withincreasingcalcicationoftheshaftcompacta, spiralbreakageandsometimesbackakingpatternsoccur.Suchbonefragments,herecompiledfor thePerickCaves( gure4 ),donotexposeanypuncturemarksofthepremolars,generally,becausethe bonebreaksarewithinthecrushingtriangle,andnotbyapuncturehole.Biomechanicallytheboneshaft (nearlyconicalcrosssection)cracksbecauseofpressureonthreesides(crushingtrianglepremolars).The mainandstrongestimpulseofthecrushcomesfromthemostpowerfulupperP3.Sometimes,cavebear femorashowsmallerround–ovaltoothmarks,orontheshaftendsonlyhalfofthepuncturemarkison themargin(e.g.Sophie’sCaveandDivjeBabe1Cavebearcubfemora;gures 3 (3)and 4 ).Thereare furtherargumentsforthehyenaoriginonlongboneshaftpuncturesfoundsimilartoradiusandtibia cubshaftswhicharesimilartofemurshaftcrosssection,butstrongerintheircompactainthisanimal agestage( gure4 )...PositionofpuncturesandonbonetypesTheamountofbonematerialisstillnotenoughtopresentclearstatistics.However,withthehereinused WeißeKuhlematerialbeingveryrepresentativeforalargecavebearden,punctureholesarefoundin thecavebearcubhumerus(4 × ),ulna(0 × ),radius(2 × ),femur(13 × )andtibia(5 × ).Thepositionofthe holesismostlyonthehereinstudied19cubfemora,ontheventralside,andnotonthedorsalconvex side(cf.gures 5 and 6 ,ande.g.‘boneuteholotype’ gure5 (4)).Thisareaisthinnerinthecompacta thanthedorsalone.Intotal,holesareventral(7 × ),dorsal(3 × )andinvecasesonbothsides.Thoseare

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rsos.royalsocietypublishing.org R.Soc.opensci. :................................................Table. Studiedandfromliteraturecompiledcavebear,hyena,wolfdensiteswithpseudo-boneutes(i.e.puncturedcavebearcubfemora),andoverlapofLat ePalaeolithicAurignaciancampsitesatthecaveentrances,orcave bearhuntsignsdeepincaves.Pseudo-boneutesarenotinMiddlePalaeolithicarchaeological,butofLatePalaeolithicandcavebeardencontextw ithlargecarnivoreinuence. localityageanimal/humanuselargecarnivoresreferences.......................................... ........................................... .......................................... ......................................... ........................................... ........................................... ..................................... ..................................... .....................................DivjeBabeCave(Sl)MIS…d,includingAurignaciansmallcavebearden/Aurignaciancampsite P.l.spelaea,P.p.spelaeus,C.lupus cf. spelaeus [ , , … ].......................................... ........................................... .......................................... ......................................... ........................................... ........................................... ..................................... ..................................... .....................................HermannsCave(D)MIS…d,includingAurignacianlargecavebearden( U.spelaeus subsp.),short-termhyenadenat entrance,Aurignaciancavebearhuntingsitedeepincave P.l.spelaea,P.p.spelaeus,C.c.spelaea,C.lupus cf. spelaeus [ , ].......................................... ........................................... .......................................... ......................................... ........................................... ........................................... ..................................... ..................................... .....................................IstalloskoCave(Hu)?MIS…dcavebearden?[ , ].......................................... ........................................... .......................................... ......................................... ........................................... ........................................... ..................................... ..................................... .....................................KepplerCave(D)MIS…dlargecavebearden( U.spelaeus subsp.),short-termhyenaden, wolfdenatentrance P.l.spelaea,C.c.spelaea,C.lupus cf. spelaeus [ , ].......................................... ........................................... .......................................... ......................................... ........................................... ........................................... ..................................... ..................................... .....................................LukvarkaCave(Sl)?MIS…dcavebearden?[ ].......................................... ........................................... .......................................... ......................................... ........................................... ........................................... ..................................... ..................................... .....................................MoriskaJavaCave(Sl)?MIS…dcavebearden?[ , ].......................................... ........................................... .......................................... ......................................... ........................................... ........................................... ..................................... ..................................... .....................................PerickCaves(D)MIS…dlargehyenadenatentrance(cubraising,communaldentype), largecavebearden( U.spelaeus subsp./ U.ingressus ) P.l.spelaea,C.c.spelaea,C.lupus cf. spelaeus, Canis [ , , … ].......................................... ........................................... .......................................... ......................................... ........................................... ........................................... ..................................... ..................................... .....................................PotokaCave(Sl)MIS…d,includingAurignacianlargecavebearden/Aurignaciancampsite,cavebearhuntingsite P.I.spelaea,C.I. cf. spelaeus [ … , , ].......................................... ........................................... .......................................... ......................................... ........................................... ........................................... ..................................... ..................................... .....................................SloupCave(CZ)MIS…dlargecavebearden/hyenadenatsidebranch P.l.spelaea,C.c.spelaea,C.lupus cf. spelaeus [ ].......................................... ........................................... .......................................... ......................................... ........................................... ........................................... ..................................... ..................................... .....................................SophiesCave(D)MIS…d(?e)largecavebearden/short-termhyenadenatentrance P.l.spelaea,C.c.spelaea,C.lupus cf. spelaeus [ , , , ].......................................... ........................................... .......................................... ......................................... ........................................... ........................................... ..................................... ..................................... .....................................GroßeTeufelsCave(D)MIS…dlargecavebearden/short-termwolfand?hyenadenatentrance P.l.spelaea,C.c.spelaea,C.lupus cf. spelaeus [ , ],herein.......................................... ........................................... .......................................... ......................................... ........................................... ........................................... ..................................... ..................................... .....................................OaseCave(Ro)MIS…dlargecavebearden/Aurignacianskullndsite C.c.spelaea,C.lupus subsp.[ ].......................................... ........................................... .......................................... ......................................... ........................................... ........................................... ..................................... ..................................... .....................................UrilorCave(Ro)MIS…d(?e)largecavebearden,short-termhyenadenatentrance P.l.spelaea,C.c.spelaea,C.lupus cf. spelaeus [ , ].......................................... ........................................... .......................................... ......................................... ........................................... ........................................... ..................................... ..................................... .....................................WeißeKuhleCave(D)MIS…dlargecavebearden( U.spelaeuseremus / U.ingressus ),?short-term wolfdenatentrance P.l.spelaea,C.lupus cf. spelaeus,P.p.spelaeus , Ursusarctos subsp. [ ],herein.......................................... ........................................... .......................................... ......................................... ........................................... ........................................... ..................................... ..................................... .....................................ZoolithenCave(D)MIS…d(?e)largecavebearden( U.spelaeus subsp./ U.ingressus ),hyenaden, wolfdenatentrancechamberarea P.l.spelaea,P.p.spelaeus,C.c.spelaea,C.lupus cf. spelaeus [ , , ].......................................... ........................................... .......................................... ......................................... ........................................... ........................................... ..................................... ..................................... .....................................

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rsos.royalsocietypublishing.org R.Soc.opensci. :................................................ Figure. StagesofcavebearfemurdestructionbyIceAgespottedhyena.Oncubfemora,whicharenotwellcalciedandelasticspongiousinthecompacta,hyenasproducedinmanycasesonlyholeswiththeirpremolarbonecrushingteeth(mainlyP)dueto unsuccessfulbonecrushing(femurfromOaseCave,Romania).Subadultcavebearfemorainitiallyaked(femurfromHermannsCave, Germany).Adultfemorahavenopuncturemarks,becausethosedirectlyakedintopieces.themostimportanttounderstandtheirhyenatoothmarkimpactorigin,becausethosecanbeattributed welltotheupperandlowerjawantagonisticbonecrushingpremolarteeth(gures 5 (1–6), 6 (10)and 7 (1)). Anotherargumentcomesfromtheovalholes,ifattributedtothebonecrushingpremolarhyenateeth theirelongationaxesareinmostcasesparalleltotheboneshaft,butonlyinholeswithintheshaft.Atthe shaftendsthosevarymore,andresultfromdiagonalcuts,mainly.Onemoreargumentfortheholesto beofhyenaorigin(orcarnivoretoothmarkingeneral)arethetoothmarkholemargins.Therearenever signsof‘drillmarksonthemarginsofthecompacta’,andinmanycasestherearebreakagesaroundthe hole(i.e.impactcircles,cf.gures 4 – 7 ).Thenalproofofholesinfemurshaftscomesfromthecrushed andakedspecimens( gure6 (8–14)),whicharedocumentedhereinforthersttimewithinacavebear den.Intwocases,oldbreakagesaredemonstratedonrettedfemurshaftsofcubs( gure6 (10–11)),and onesubadult / adultshaft( gure7 (5));inallcasesthefragmentshaveevendifferentcoloursandwere embeddedaftercrushingindifferentsedimenttypes / layers..Discussion..Dating:AurignaciansversusNeanderthalsAll‘cavebearcubfemoraboneute’sitesfailedtodateintothe‘Neanderthaltimes’becauseallarenotof Neanderthal(MiddlePalaeolithic)human,butareinsteadfrommodernhumanAurignacianoccupations in‘archaeologicallayers’atentrancesofcavebeardens,cf.[ 1 , 7 , 9 ],ordeeperincavesduetocavebear hunt[ 23 ].There,thecavebearlayersthemselves,whichgenerallyspanfromtheMIS3–5d = 25,000– 113,000BP,overlap / intercalatewiththeCro-Magnontimes,mainlyAurignacian,partlyGravettian, culturallayers[ 5 , 23 , 57 ].Cavebearbonesandarchaeologicallayersarethereforenotexactlyisochronous inseveralcases(evenmixedduetopossiblybioturbationbycavebearsbuildingtheirnests,orburrowing porcupinesordiggingIceAgespottedhyenas;cf.[ 13 , 20 , 21 ]),and‘cavebearboneutes’wouldhave been,ifsuch,frommodernhumanlayers,inallcases.

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rsos.royalsocietypublishing.org R.Soc.opensci. :..................................................Cavebearsubspeciesandpseudo-boneuteoriginsThepseudo-boneutesallcomefromlayersoftheMIS3–5d(hereinaddeduptoMIS6)andarefrom smallerearlycavebearformsof Ursusspelaeus subsp.(i.e. U.s.eremus,U.s.spelaeussensu taxonomy ofStiller etal .[ 58 ]);interestinglythough,alpineLatePleistocenecavebearforms( U.s.ladinicus )do notshowsuchholesinfemora(i.e.indicatorofabsenceofhyenasinalpineregions,andproofofholes madeonlybyhyenaswhicharefoundonlyinmiddlehighelevatedmountainregions[ 19 ]).Asisnow wellknown,AurignacianhumanslivedinEuropetogetherwiththelastandlargestcavebearspecies U.ingressus [ 16 , 18 , 21 , 23 , 58 , 59 ].AtafewcavesitesinEurope,huntingofcavebearswithpropulsory spearsisdocumentedfortheAurignacians–Gravettians[ 5 , 22 , 23 , 57 ].Therefore,the‘pseudo-boneutes’ originatefrombothsmaller Ursusspelaeus subsp.( eremus or spelaeus )andthelarge U.ingressus ,andfrom mountainousregions,whereIceAgespottedhyenaswerearoundalloverEurope(cf.mapin[ 19 ])...StonetooldrillexperimentsDrilledholeswereproducedexperimentallyforareconstructionofa‘cavebearcubboneute’(cf. [ 60 ])andareverydifferentalsoontheholemarginsandforms.Allhereinguredcubfemorahave, differentfromdrill-holes,distinctcharacters(gures 5 – 7 ):(a)theholesarenotfullyround,insteadovalshaped,andbesidethehole(seealso[ 24 ])abreakage-archindicatesan‘impact’,ratherthandrilling(cf. alsomodernhyenaimpactmarkpicturesin[ 61 ]),(b)themarginsareconvexincross-shape,andnot steep-straightaswithdrills,(c)thecornersaresmoothanddonothavedrill / cutmarksigns,atall,and (d)inmostcases(gures 5 – 7 ),theantagonisticpunctures / toothmarks(lower / upperjawdentitiont) arepresent.Suchantagonistictoothmarksarefoundoftenatdifferentmedium-sizedhyenapreybones includingtheirownspeciesfemoraorevenNeanderthalfemora[ 19 , 20 ],alsodocumentedinthemodern actualisticspottedhyenaboneaccumulationrecord[ 61 – 63 ].Finally,alsoX-raysofthe‘boneute’hole marginsdidnotverifyany‘drilling’noranystonetoolworkonthebone(cf.[ 64 ])...MicetoothmarksandholesSmallparallelrectangularscratchesonthepseudo-boneuteofDivjeBabeCave1weremisinterpreted ascutmarks(cf.[ 24 ]).Alsooneholeofthepseudo-boneuteofIstállósköCaveisclearlyproduced,ora toothmarkholeextended,bymousechewing(cf.photoin[ 6 ])...CaninetoothmarkimpactholetheoriesThesearenotpresentontheboneshafts,asfangteethofhyenas(oranyothercarnivore)arenever usedtocracklongbones(e.g.[ 61 ]).Thedentitionisveryheterodontinthosespecializedmammals(cf. gure2 ).Theincorrectbiomechanicalillustrationofthehyenateethandjawfunctionleadstoincorrect interpretationsofhyenasaspossibleproducers(cf.[ 24 ]),whichattributedpossible‘holes’to‘canines’, whichwascontradictorytoseveralargumentsbyTurk etal. [ 24 ],becausethedistalendswerealready saidtohave‘carnivoredamage’[ 44 ].Thetomography(cf.[ 65 ])restudyoftheboneexcludedIceAge hyenas,arguingwith‘abnormalbitingorchewingbehaviourusingtheircanineteeth’(cf.[ 66 ]).Indeed, neitherhyenas,modernnorextinct,noranycarnivoresmentionedusecaninesfor‘bonecrushing’(e.g. [ 22 , 61 , 67 ]; gure2 ).Also,possiblymostherbivoroussmall U.s.eremus andfullherbivorous U.ingressus (e.g.[ 59 ])cavebearsdidnotusethecaninesfor‘bonecrushing’,whichwerealsoproposedtobethe ‘holeproducersincavebearboneshafts’informertimes,alsoforgettingthetoppredatorsofcavebears inthe‘cavebearbook’(cf.[ 68 ]).Someactualisticproofforthenon-useofthecaninescanalsobefound inmodernbrownbear U.a.arctos preytaphonomystudies,wherethosedidnotcrushanylongbones onungulatecarcasses,whereasthosemightpuncturesofterspongiosapartswiththeircanines[ 69 ]. Microscopicanalysis(cf.[ 27 ])declaredthefemurofayoungcavebear(‘boneute’)probablytobe piercedbythe‘bearsthemselves’alsouncriticallyrepeatedincorrectlybyothers(cf.[ 33 , 70 , 71 ]).These old‘cavebearcannibalisticmodels’werealreadyrevised,withmanyargumentsnottobeexistent, includingthetoppredatorsasbonedamagers(includinghumanbones)inEurope(e.g.[ 4 , 16 , 19 – 21 , 72 ]). IncludingalsothenewstudiesoftheomnivorousbrownbearsofArilla etal. [ 69 ],thoseseemtoexclude evencannibalismwithin U.a.arctos. AlsothegureofbonecrushingbyTurk etal. [ 24 ]wasincorrectly presented,usingonlyonelowerjawpremolar,althoughthebonecrushingtriangleconsistsofthree teeth( gure2 ).UsingallthesestrangepresentationsbyTurk etal. [ 24 ],thisindeedwouldbeabnormal forhyenastotrytocrushlongboneswiththeircanineteeth.Whichhyena(lionandwolf)teethare

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rsos.royalsocietypublishing.org R.Soc.opensci. :................................................responsibleforwhatkindoftoothmarksandbonedamageoncavebearbonefemoraandotherfew selectedcranialandpostcranialmaterialisreguredcomposedofseveralstudies[ 18 – 22 ]( gure2 )...HyenasatcavebeardensinEuropeItisnowonderthenthatfurtherincorrectnessaboutcavebearbonetaphonomyatDivjeBabeCave1was published(cf.[ 73 ]),becauseall‘fragmented’bonesweresimplydeclaredasdueto‘sedimentpressure’. Indeed,somearenaturallyweather-cracked.Astudiedulnaofacavebearatthesiteisoneofthebest examplesofbonecrushingbyhyenapremolarteeth.Eventhepuncturemarksintheuppershaftarea arevisible,demonstratingthescavenging / bonecrackingactivitiesalsointheDivjeBabeCave1,similar tothatfoundinGermancaves(cf.[ 4 , 16 , 18 , 23 ];cf.gures 5 – 7 ).Hisnalargumentsthat‘hyenasare absent’atthissite(cf.alsocarnivorefaunain[ 25 ])arenone,becauseasignoredinintensivecavebear dencavesitetaphonomystudiesofEurope,themodelsofpresenceandabsenceofanylargepredator arewellknown[ 4 , 13 , 18 , 19 , 23 ].Hyenasandothercarnivoresarerarelyfoundatthe‘scavengingsites’, includingcavesandcavebeardens,becausetheyareonlyfoundtherewhentheyoccupiedthecave entrancesas(a)cubraising,(b)communalor(c)preydepotdens(cf.denitionsanddiscussionsin [ 4 , 14 , 18 – 20 , 20 , 21 , 54 , 74 ]..ConclusionThe‘cavebearcubfemorawithholes’are,inallcases,neitherinstrumentsnorhumanmadeatall. Allcavebearpseudo-boneutesarenotdatedtoNeanderthalMiddlePalaeolithicMousterianlayers, butinstead,ifpossibletodate,toLatePalaeolithic,Aurignacian / Gravettianlayers.There,wherethey aredatedabsolutely(DivjeBabeCave1)arewithoutarchaeologicalcontextatall,andsimplyofcave beardenuseduringtheMIS3–5d.Atthesetimes,differentcavebearsubspecies Ursusspelaeus subsp. eremus (smallestcavebear)and spelaeus (i.e.Neanderthaltimes)and U.ingressus (largestcavebear, i.e.Aurignacian / Gravettiantimes)usedcavesalloverEuropeforcubraisingandhibernation.Allthe largecarnivorepuncturedcavebearcubfemora(andotherpuncturedbones)appearalwaysinsmall tolargecavebeardencave / caveentrancecontexts.Thissometimesoverlapswithhyenadensand humancampsitesatcaveentrancesonly,wherecavebearden,carnivoredenandhumanremainsare evenmixedup(partlyseparatedinlayers),alloverEuropeduetocompetitionforandseasonaluseof caveentrances / rockshelters.Thecavebearboneswithround–oval,largerpuncturemarkscanbewell attributedsolelytothemaincavebearscavengerofEurope—theIceAgespottedhyena Crocutacrocuta spelaea .ThismainLatePleistocenebonedestructorinEuropeisknownrecentlywithmorethan150den sites(95%arecavesites)alloverEurope.Atcavebeardenshyenasleft,byperiodicscavenging,upto 20%ofdamagedbones,whereasalsolions(cavebearkillers),leopardsandwolvesplayedalargerrolein thecavebearhunting / scavenging,evendeepincaves.Thoseindeedalsoleft,insomecases,round–oval, largerpuncturesincavebearbones,butwiththeircaninesonlyinsoftspongiosa(pelvis,vertebrae),and neverinanyboneshaftcompacta.Neithercarnivoresnorcavebears(herbivorous)usedtheircanine teethtocrushlongbones,oranyotherbones.Therefore,allothertoppredators—excepthyenas—can beexcluded,atleastfortheround–ovalpuncturesincavebearlongboneshafts.Onlyhyenashave developedacarcassdestructionandbutcherystrategy,alsoforcavebears.Thisstrategyisdemonstrated, hereinindetail,oncavebearfemoradestruction(especiallymaterialfromWeißeKuhleCave,Germany), whichispresentedinthreestagesandfordifferentagedindividuals—cubs(lessthan1year),subadults (1–2years)andadults.Cubbonesare‘soft’andthin-walledintheboneshaftcompacta,whichincreases insubadultstoadults.Thisexplainswhypuncturemarksarefoundonlyincub(less1year)femora,and partlyinsubadults,whereastheyareabsentcompletelyinadults,becausehyenascrackedthosebones intopieceswiththepremolartriangleteeth(i.e.bonecrushingteeth)foraccesstothebonemarrowand easierswallowingofthosepiecesforthebonecollagenuse.Hyenasleft,therefore,‘pseudo-boneutes’ duringtheLateMiddletoLatePleistocenealloverEuropeincavebeardens,andondifferentcavebear species / subspecies.Thisisknownduetolackofbreakageonmostofthecavebearcubfemora,which generallyshowadditionaldiagonalzigzagmargins(fromchewingjointsbyscissorteethofhyenas)or havetriangularorsmallerscratchtoothmarks.Thisevenallowsreconstruction,insomecasesindetail, thetoothmarkoftheupperandlowerjawteethofhyenas—thelasttoothmarkofthepremolarbone crushingtriangleofthepowerfuljawsofthelasthyenasofEurope.Finally,someakesandrettedcub femora,bothwithtoothmarkholes,provethebonecrackingactivitiesatcavesites.

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rsos.royalsocietypublishing.org R.Soc.opensci. :................................................Acknowledgements.IthankforthestudysupportofmaterialfromSophie’sCaveMrsS.Dess.Boneworkwasallowedin theGroßeTeufelsCavebyS.FrühbeißerandwassupportedbyT.Büttner.MaterialstudiesoftheZoolithenCavewere allowedbyPDDrO.Hampe(MuseumfürNaturkundederHumboldt-UniversitätBerlin),whereascaveexploration andboneworkinthecavewasmainlysupervisedbyM.Conrad(FränkischeHöhlen-undKarstForscher).Especially E.Schuhose,G.VolmerandB.Baumbach(RübeländerHöhlenforschergruppe)andC.Hensel(cavemanagement) supportedthecaveexplorationandboneworkinHerman’sCave.J.Toernig-StruckgaveaccesstotheKepplerCave materialoftheStadtmuseumMenden.H.-W.Weber(ArbeitsgemeinschaftHöhleundKarstHemere.V.)helpedinthe PerickCaveexplorationandbonedumpwork.TheOaseCavematerialwasstudiedintheSpeleologicalInstituteEmil Racovita.S.Stevensdidthespell-checkoftherstdraft,whereastheillustrationsarefromG.Teichmann.Finally,I thankDrJ.R.Ardèvolandanunknownreviewerforthereviewoftherstdraft.Fundingstatement.TheresearchwassponsoredbythePrivateResearchInstitutePaleoLogic( www.paleologic.eu ),which runsthe‘EuropeanIceAgespottedhyenaproject’.References.BrodarS,BrodarM.Potokazijalka: visokoalpskapostajaaurignacienskihlovcev (Potokazijalka„einehochalpine Aurignacienjäger-Station).SlovenianAkademije ZnanostiinUmetnostiClassisIDela ,…. .PacherM.DieHöhlenbärenresteausder SammlungGroßausderUschowaHöhle (PotokaZijalka,Slowenien). 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rsos.royalsocietypublishing.org R.Soc.opensci. :................................................Babespecimenandthediatonicscale. Stud.Music Archaeol. ,…. .HorusitzkyFZ.Lesûtespaléolithiques:Divje BabeI,Stállóskö,Lokveetc.pointdevuedes expertsetdescontestataires.Critiquede lappréciationarchéologiqueduspécimenno. deDivjeBabeIetargumentspourladéfensedes specimenspb/etpbduMNMdeBudapest. ArheolokiVestnik ,…. .HoldermannC-S,SerangeliJ.Die NeanderthalerotevonDivje-Babe:Eine RevolutioninderMusikgeschichte?Musica instrumentalis. Z.Organologie , ,…. .DErricoF.JustaBoneoraute? Stud.Music Archaeol. ,…. .TunizC,BernardiniF,TurkI,DimkaroskiL,ManciniL, DreossiD.DidNeanderthalsplaymusic?X-ray computedmicrotomographyoftheDivjeBabe ute. Archaeometry ,…. .TurkM,DimakaroskiL.Neanderthalutefrom DivjeBabeI:oldandnewndings. 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