The reality of Neandertal symbolic behavior at the Grotte du Renne, Arcy-sur-Cure, France


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The reality of Neandertal symbolic behavior at the Grotte du Renne, Arcy-sur-Cure, France
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PLOS ONE
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Caron, François
d'Errico, Francesco
Del Moral, Pierre
Santos, Frédéric
Zilhão, João
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Neanderthals ( lcsh )
Caves ( lcsh )
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serial ( sobekcm )
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Europe -- France -- Bourgogne-Franche-Comté -- Arcy-sur-Cure

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The question of whether symbolically mediated behavior is exclusive to modern humans or shared with anatomically archaic populations such as the Neandertals is hotly debated. At the Grotte du Renne, Arcy-sur-Cure, France, the Châtelperronian levels contain Neandertal remains and large numbers of personal ornaments, decorated bone tools and colorants, but it has been suggested that this association reflects intrusion of the symbolic artifacts from the overlying Protoaurignacian and/or of the Neandertal remains from the underlying Mousterian. We tested these hypotheses against the horizontal and vertical distributions of the various categories of diagnostic finds and statistically assessed the probability that the Châtelperronian levels are of mixed composition. Our results reject that the associations result from large or small scale, localized or generalized post-depositional displacement, and they imply that incomplete sample decontamination is the parsimonious explanation for the stratigraphic anomalies seen in the radiocarbon dating of the sequence. The symbolic artifacts in the Châtelperronian of the Grotte du Renne are indeed Neandertal material culture.
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TheRealityofNeandertalSymbolicBehavioratthe GrotteduRenne,Arcy-sur-Cure,FranceFranc ¸oisCaron1,Francescod’Errico2,3* ,PierreDelMoral1,Fre ´ de ´ ricSantos2,Joa ˜ oZilha ˜ o41 InstitutNationaldeRechercheenInformatiqueetenAutomatiqueBordeauxSud-Ouest,InstitutdeMathe ´ matiquesdeBordeaux,Universite ´ deBordeaux,Talence, France, 2 Unite ´ MixtedeRecherche5199DelaPre ´ histoirea ` l’Actuel:Culture,EnvironnementetAnthropologie,Universite ´ deBordeaux,Talence,France, 3 Institutefor Archaeology,History,CulturalandReligiousStudies,UniversityofBergen,Bergen,Norway, 4 Seminarid’EstudisiRecerquesPreisto ` riques,UniversitatdeBarcelona/ Institucio ` CatalanadeRecercaiEstudisAvanc ¸ats,Barcelona,SpainAbstractBackground:Thequestionofwhethersymbolicallymediatedbehaviorisexclusivetomodernhumansorsharedwith anatomicallyarchaicpopulationssuchastheNeandertalsishotlydebated.AttheGrotteduRenne,Arcy-sur-Cure,France, theCha ˆ telperronianlevelscontainNeandertalremainsandlargenumbersofpersonalornaments,decoratedbonetoolsand colorants,butithasbeensuggestedthatthisassociationreflectsintrusionofthesymbolicartifactsfromtheoverlying Protoaurignacianand/oroftheNeandertalremainsfromtheunderlyingMousterian.Methodology/PrincipalFindings:Wetestedthesehypothesesagainstthehorizontalandverticaldistributionsofthe variouscategoriesofdiagnosticfindsandstatisticallyassessedtheprobabilitythattheCha ˆ telperronianlevelsareofmixed composition.Ourresultsrejectthattheassociationsresultfromlargeorsmallscale,localizedorgeneralizedpostdepositionaldisplacement,andtheyimplythatincompletesampledecontaminationistheparsimoniousexplanationfor thestratigraphicanomaliesseenintheradiocarbondatingofthesequence.Conclusions/Significance:ThesymbolicartifactsintheCha ˆ telperronianoftheGrotteduRenneareindeedNeandertal materialculture.Citation: CaronF,d’ErricoF,DelMoralP,SantosF,Zilha ˜ oJ(2011)TheRealityofNeandertalSymbolicBehaviorattheGrotteduRenne,Arcy-sur-Cure,France.PLoS ONE6(6):e21545.doi:10.1371/journal.pone.0021545 Editor: DavidFrayer,UniversityofKansas,UnitedStatesofAmerica Received May18,2011; Accepted June1,2011; Published June29,2011 Copyright: 2011Caronetal.Thisisanopen-accessarticledistributedunderthetermsoftheCreativeCommonsAttributionLicense,whichpermits unrestricteduse,distribution,andreproductioninanymedium,providedtheoriginalauthorandsourcearecredited. Funding: ThisworkwassupportedbytheEuropeanResearchCouncil(FP7/2007/2013/ERCGrantTRACSYMBOLSn u 249587).Thefundershadnoroleinstudy design,datacollectionandanalysis,decisiontopublish,orpreparationofthemanuscript. CompetingInterests: Theauthorshavedeclaredthatnocompetinginterestsexist. *E-mail:f.derrico@pacea.u-bordeaux1.frIntroductionFormostofthe20thcentury,personalornaments,systematic pigmentuseandelaboratebonetechnologywereassociatedwith theco-emergenceof‘‘Cro-Magnon’’peopleandtheUpper Paleolithic(beginninginWesternEuropewiththeCha ˆtelperronian)[1].Overthelastthreedecades,anumberoffindings challengedthisview[2–9],namely:(a)inAfrica,ornaments, pigmentsandabstractdecorationoccuramongtheimmediate ancestorsofmodernhumans;(b)anatomicalmodernityemerged earlierinAfrica;(c)burialritual,jewelryandbodypaintingare knownamongMiddlePaleolithicEuropeanNeandertals;(d) wheretheCha ˆtelperronianisfoundwithdiagnosticfossils,these areofNeandertals,notmodernhumans;(e)theearliestevidence foranatomicalmodernityinEuropepost-datesbymanymillennia theemergenceoftheCha ˆtelperronian,corroboratingthe Neandertalauthorshipofthelatter;(f)nosuddenchangeor accretionisobservedintheearlystagesofsymbolicmaterial culturebutratheradiscontinuouspatternofasynchronous emergence,disappearanceandre-emergenceofitsfeaturesamong both‘modern’and‘archaic’populationsofthetwocontinents.As aresult,mostpaleoanthropologistsnowacknowledgethat ‘‘symbolicthinking’’and‘‘modernbehavior’’arenotspeciesspecificfeaturesofanatomicallymodernhumansandthat Neandertalswerethemakersofasymbolicmaterialculture [4,7–13]. TheCha ˆtelperronianlevels(VIII,IXandX)ofaFrenchcave site,theGrotteduRenne(Arcy-sur-Cure,Burgundy),document theassociationbetweenNeandertalremainsandlargenumbersof personalornaments,bonetools(someofwhicharedecorated),and colorants(Figure1).Althoughexcavated(byA.Leroi-Gourhan) between1949and1963[14–15],thetechniquesusedwere modern(stratigraphicdiggingandareaexposureofoccupation surfaces,spatialplottingofkeyfindsandfeatures,systematic sievingofthedeposits),andthegeologicalrealityofthedescribed succession—confirmedbylimitedexcavationofastratigraphic baulkcarriedoutin1998[16]—isuncontroversial.Evenso,the followingconcernshavebeenvoicedwithrespecttothe homogeneityoftheartifactassemblages[17–20]:(a)noinvestigationofpotentialrefittingofstonetoolsacrosslevelshasbeen carriedout,sotheextenttowhichtheGrotteduRennesequence wasaffectedbypost-depositionaldisturbancecannotbeassessed; (b)theexistenceofoverlyingProtoaurignacian,Aurignacian/ GravettianandGravettianlevels(VII,VIandV,respectively)and thefactthatsomeornamenttypespresentintheCha ˆtelperronian sequence(e.g.,piercedfoxteeth)arecommoninlaterEarlyUpper Paleolithic(EUP)technocomplexesraisethepossibilitythatthe symbolicartifactsfoundintheCha ˆtelperronianareintrusivefrom PLoSONE|www.plosone.org1June2011|Volume6|Issue6|e21545

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above;(c)conversely,asthehabitationstructuresbuiltbythefirst Cha ˆtelperronianoccupantsofthesiteconceivablydisturbedthe underlyingMiddlePaleolithiclevels(XI–XIV),theNeandertal remainsrecoveredinlevelsVIII–Xcouldbeintrusivefrombelow. Onthebasisoftheseconcerns,threealternativestothe stratigraphicintegrityoftheCha ˆtelperronianlevelsoftheGrotte duRenneareconceivable:(a)thepersonalornaments,bonetools andcolorantsareProtoaurignacianorlaterandtheNeandertal remainsareMousterian,sothesite’sCha ˆtelperronianlacked symbolicartifactsandisofunknownauthorship(hereafter Hypothesis1);(b)thepersonalornamentsareProtoaurignacian orlater,sothesite’sCha ˆtelperronian,althoughmadeby Neandertals,lackedsymbolicartifacts,asthecolorantsandbone toolsmayberegardedaspurelyfunctional(hereafterHypothesis 2);and(c)thepersonalornaments,bonetoolsandcolorantsare Cha ˆtelperronian,buttheNeandertalremainsareMousterian,so theCha ˆtelperronianmaywellhavebeenmadebymodern humansinstead(hereafterHypothesis3). Inthefollowing,thedistributionsofornaments,bonetools, colorants,pigmentprocessingtoolsandhumanteethare comparedwiththoseofdiagnosticstonetools:Levalloisflakes (Mousterian);Cha ˆtelperronpointsandconvergentsidescrapers, thelatterasaproxyforthe racloircha ˆtelperronien ,atooltypethat Leroi-Gourhan[21]definedascharacteristicofthesite’s Cha ˆtelperronianbutisnotdiscriminatedassuchinavailable inventories;Dufourbladeletsandtheirunretouchedblanks (Protoaurignacian).Usingthistestedtaphonomicapproach[22], weassesswhetherthefinalconditionrepresentedbytheobserved distributionscanbederivedbypost-depositionaldisturbancefrom anyoftheinitialconditionsimpliedbyHypotheses1–3,which, statistically,areournullhypotheses.Ourresultsrejectthem. Therefore,thealternativeview—theassociationbetweensymbolic artifacts,Cha ˆtelperronianstonetoolsanddiagnosticNeandertal fossilscharacteristicoflevelsVIII–Xisgenuineandreflectsthe behavioroftheGrotteduRenne’sNeandertaloccupants—stands unrefuted.ResultsOfthedifferentpost-depositionalprocessesthatcanaffectan archeologicalsite[23],generalized,largescaledisturbancecanbe rejectedfromfurtherconsiderationinthiscasebecausethe diagnosticartifacts,uponwhichthedifferentlevelswereassigned todifferenttechnocomplexes,wouldhavebecomescrambledand thesequencewouldhavebeenrecognizedfromtheoutsetas stratigraphicallymixed.Asshownbythedistributionsforlevels VII–XIV(Table1),nothingcanbefarthestfromthetruth,as 100%ofallLevalloisflakeswererecoveredinMousterianlevels XI–XIV,99%ofallCha ˆtelperronianpointswererecoveredin Cha ˆtelperronianlevelsVIII–X,and100%ofthebladelets(both Dufourbladeletsandunretouchedblanks)wererecoveredin ProtoaurignacianlevelVII. Conceivably,twoothermechanismscouldhavecreatedthe notionalcontradictionbetweenastratigraphicdistributionoflithic diagnosticsthatmatchesexpectationsin . 99%oftheinstances andtheputativedisplacementofentirecategoriesofitemsfound Figure1.GrotteduRenne,Cha ˆ telperroniansymbolicartifacts. Personalornamentsmadeofperforatedandgroovedteeth(1–6,11),bones (7–8,10)andafossil(9);red(12–14)andblack(15–16)colorantsbearingfacetsproducedbygrinding;boneawls(17–23).1–9courtesyofM. Vanhaeren,12–16courtesyofH.Salomon,17–23modifiedafter[24]. doi:10.1371/journal.pone.0021545.g001 NeandertalSymbolicBehavioratGrotteduRenne PLoSONE|www.plosone.org2June2011|Volume6|Issue6|e21545

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inassociationwithsuchdiagnostics[23].Onesuchmechanismis localized,largescaledisplacementreachingnon-adjacentlevelsof thestratigraphy(e.g.,asaresultofmammalburrowingor subsurfaceanthropogenicintervention);theotherissmallscale, gradualandcumulativedisplacementofindividualobjectsacross theentiresequencethroughgeologicalorpedogeneticprocesses (e.g.,asaresultofcryoturbationorroot/wormactivity).Localized,largescaledisplacementThestratigraphicconsistencyofverticaldistributionsdoesnot counter,perse,thatunrecognizeddisturbanceofaparticulararea ofagivenlevelmayexplainthepresenceinthatstratigraphicunit offindcategoriesotherwiseabsentfromintactareas.Forinstance, inthecaseofCha ˆtelperronianlevelXoftheGrotteduRenne,the ornamentsandothersymbolicitemscouldcorrespondtoa subsurfacecachecreatedbylevelVIIProtoaurignacianpeople thatburrowinganimalssubsequentlymovedfurtherdown,while theNeandertalremainscouldcomefromadiscreteaccumulation (relatedtoe.g.secondaryburial)thatCha ˆtelperronianconstructionactivityoranimalburrowingsubsequentlymovedupfrom MousterianlevelsXI–XII. Theexpectedoutcomeofthedisplacementofsuchfindclusters isapatternwheretheoriginalconcentration,evenifdilutedbythe disturbanceprocess,wouldbepreservedtosomeextent,defininga scatterwithaclearcenterclosetoitsoriginallocationand increasinglysparsetowardtheperiphery.Incontrasttosuchan expectation,thepersonalornamentsinlevelXfeaturea homogeneous,lowdensitydistributionacrosstheentireexcavated surface,withmostfindscomingfromoutsidetheareaofdensest Protoaurignacianoccupation,whiletheboneawls[24]formtwo broadclustersthatcoincidewiththelocationoftheCha ˆtelperronianhabitationfeatures(Figures2,3).TheNeandertalremainsare alsoscatteredand,althoughfoundoutsidethehabitationfeatures, wererecoveredwellinwardofthedripline,counteringthenotion thattheyreflectdumpingatthecaveentranceofMousterian sedimentsremovedbyCha ˆtelperronianconstructionactivity (Figure4).TheapparentconcentrationagainsttheEastwall reflectsthepresenceofthreegroupsofteethfromthreedifferent individuals,interpretedastheinsitudisintegrationofsingle mandibularormaxillarypieces[5]. AnotherhypothesisisthattheNeandertalteethderivedby progradationfromstratigraphicallylowerbuttopographicallyhigher Mousteriandeposits[19].Conceivably,thiscouldhavehappened southwardofrow8andwestwardofrowC,becauseofthemarked slopeofthestrataoutsidetheMousteriandripline.Mostteeth(80%), however,werefoundinwardofthisdripline,wherethestratification oftheCha ˆtelperroniandepositsishorizontal.Mostwerealsofound againsttheeasternwallofthecave—14outofthe22securely provenancedtolevelX(64%)camefromrowsC–Dofthegrid (Figure4).Giventhisspatialdistribution,thepresenceofNeandertal teethinlevelXcannotresultfromprogradationprocessesassociated withthewestwarddipofimmediatelyunderlyingMousterianlevel XI.Suchprocessesaregravity-driven,sotheycouldhavediplaced materialintotheadjacentY-Browslowerdownbutnotintotheexact sameC–Drowshigherup.Ratherthanthe25Neandertalteethin Cha ˆtelperronianlevelXreflectingdownslopeerosionofthe Mousteriandepositstowardsthecenterofthecave,itisthetwo teethinMousterianlevelsXI–XIIthatlikelycorrespondto downwarddisplaceditems,asisthecasewiththefewboneawls andCha ˆtelperronpointsalsofoundinlevelsXI–XII[7,24]. Thedistributionofornaments,bonetools,pigmentsand pigmentprocessingtools[25–29]iscongruentwiththeplacement ofthehabitationfeaturesrecognized(Figures3,5).InProtoaurignacianlevelVII,thereisaconcentrationontheeasternside ofthecave,almostexclusivelyinbandsB–D,wherethefew pigmentsandpigmentprocessingtoolswereassociatedwith hearths(alargespotofredpigmentwasalsonotedtowardthe entrance).InCha ˆtelperronianlevelX,thespatialdistributionof theabundantpigmentschangesslightlybetweensublevelsbut,in agreementwiththelocationofthetoolsforpigmentprocessing, maximumconcentrationssystematicallyoccurinsidethefeatures identifiedinthecentralandnorthwesternpartsofthecave,where mostornamentsandbonetoolswerealsofound.Thesepatterns strengthenthecaseforthestratigraphicintegrityoflevelsVIIand Xandfurthercounterthenotionthatthesymbolicitemsfound thereinweredifferentiallyaffectedbypost-depositionaldisturbance.Generalized,smallscaledisplacementLimitedpost-depositionalmovementacrosstheboundariesof adjacentlevelsisaubiquitousfeatureofcaveandrockshelter stratigraphies.TheGrotteduRenneisunlikelytohavebeen immunetosuchprocesses,andexcavationerrorcanresultinthe misassignmentoffindsfromtheinterfacebetweendifferentunits. Indeed,aswiththetwoawlsintheMousterianthatprobablycome fromCha ˆtelperronianlevelX,foursmallornamentalivory fragmentsfromuppermostCha ˆtelperronianlevelVIIIprobably originatedintheimmediatelyoverlyingProtoaurignacian[7,30]. Dosuchordinarypost-depositionalmechanismssufficeto supportanyoftheHypotheses1–3above?Intuitively,no,because Table1. GrotteduRenne.Stratigraphicdistributionofkeyfinds*.LevelsOrnamentsPigments Worked bone Neandertal teeth Dufour bladelets Cha ˆ telperron points Levallois flakes Unretouched bladelets Convergent sidescraprers VII8397002871028000 VIII8146271029002 IX22861730670028 X29118313925028400105 XI051102904 XII0411001402 XIII000000301 XIV000300002 *Inordertosimplify,andalsobecauseinventoriesofthedifferentfindcategoriesareunavailableforthem,levelsVIandabovearenotconsideredh ere. doi:10.1371/journal.pone.0021545.t001 NeandertalSymbolicBehavioratGrotteduRenne PLoSONE|www.plosone.org3June2011|Volume6|Issue6|e21545

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(a)explainingthe‘‘advanced’’findsmadeintheCha ˆtelperronian levelsasentirelyintrusivefromthelaterEUPinlevelsVIIor abovefacestheproblemthatmostsuchfindscamefromlevelX, notlevelVIII,and(b)althoughmostNeandertalremainsfromthe Cha ˆtelperronianwereinlevelX,thenotionthatadjacent MousterianlevelsXI–XIIseededthefossilsrecoveredinthe immediatelyoverlyingunitfacestheproblemthatonly6%would haveremainedput[7–8,30–31]. Inordertomorerigorouslytestwhethertheoperationofthis mechanismcouldbeheldresponsiblefortheobserveddistributions,wemodeleditasarandomprocesswherebythefinal condition(Table1)derivesfromtheinitialcondition(TableS1)as aresultofsmallscaledisplacementsaffectingtheentirethickness ofthedepositsconcernedhere(seeMaterialsandMethods).In eachcase,theresultsofthegoodnessoffittestreject,fora probabilitythresholdof0.01,thepossibilitythattheobserved distributioncanbetheoutcomeofthedisturbance,viathe modeledmechanism,ofasequencepresentinganyoftheoriginal distributionsimpliedbyHypotheses1–3.UnderHypotheses1and 2,thedistributionoftheitemsputativelyintroducedfromlevelVII orabovewouldhavetofeatureagradualdecreasewithdepth insteadoftheobservedmarkedconcentrationinlevelX(whichis allthemoreremarkableinthecolorants’caseas,byweight,the observedtotalsare0.45kginlevelVIIand14.58kginlevelX; Table2).Likewise,underHypothesis3,theupwardlydisplaced Neandertalremainswouldhavetobeinmuchsmallernumbers thanthoseremaininginsitu.Finally,underanyofthehypotheses, thedisplacementswouldsignificantlyaffectthediagnosticstone tooltypes:underHypotheses1and2itisextremelyunlikelythan nobladeletsaccompaniedthedownwarddisplacementofthe ornamentsandunderHypothesis3itisextremelyunlikelythatno Levalloisflakesaccompaniedtheupwarddisplacementofthe Neandertalremains. Assuminggeneralized,smallscaledisturbance,theworstcase scenarioforthestratigraphicintegrityoftheGrotteduRenneis onewhere,againstthegeologicalevidence,thethreeCha ˆtelperronianlevels(VIII–X)areconflatedintoasingleoccupation,and post-depositionaldisplacementismodeledasoccurringbetween twoadjacentblocksonly—fortheornaments,involvingoverlying ProtoaurignacianlevelVII,and,fortheNeandertalteeth, involvingunderlyingMousterianlevelsXI–XII(seeMaterials andMethods).Evenso,theobserveddistributionofthelithic diagnosticsimpliesthat,forthepredictedinitialdistributions (TableS1)andaprobabilitythresholdof0.01,nomorethan1 ornament(Hypothesis2)or7Neandertalteeth(Hypothesis3) couldhavefoundtheirwayintothatsingleCha ˆtelperronianblock (FigureS3),whereastheobservedtotalsare39and29, respectively.DiscussionSincetheobservedverticalandhorizontaldistributionpatterns cannotbederivedbypost-depositionaldisturbancefromtheinitial setofconditionsimpliedbyHypothesis1–3,weconcludethatthe associationbetweenCha ˆtelperronpoints,personalornaments, bonetools,colorantsandNeandertalremainsfoundattheGrotte duRenneisgenuine.Incontrast,usingBayesianstatisticsapplied tothestratigraphicdistributionofbonetoolsandhuman-modified faunalremainsdirectlydatedbyradiocarbon,arecentassessment ofthesite’sintegrityarguedforsignificantproblemsbecausemore thanonethirdofthedatesturnedouttobeoutliers[20]. Ourresultsshowthatthereasonforthisanomalymustbe soughtelsewhere,asthedistributionofthelithicdiagnosticssets strictlimitstothepotentialdisturbanceoftheotherfindcategories, includingthoseaddressedbytheradiocarbonstudyandnot consideredhere(suchasthefaunalremains).Forinstance,8(out of21)Cha ˆtelperroniansamplescameoutyoungerthanor overlappingwiththe , 36.5ka14CBPlimitsetbychronostratigraphicpatternsforthebeginningoftheProtoaurignacianin Europe[32].But,since0outof287Dufourbladeletsand0outof 2800unretouchedbladeletsintruded,(a)theprobabilitythatthose 8samplescorrespondtointrusiveitemsis , 1e-18and(b)the probabilitythatmorethanonedatedCha ˆtelperroniansampleis displacedfromlevelVIIis , 0.01(FigureS3).Giventhese probabilities,itisclearthattheaccuracyoftheresultsobtained shouldbetreatedasanopenissue,whichinvalidatestheoutlier Figure2.GrotteduRennedistributions. Left:personalornamentsfromCha ˆ telperronianlevelsVIII–Xplottedagainstthemostdenselyoccupied areaofProtoaurignacianlevelVII.Right:NeandertalremainsfromCha ˆ telperronianlevelsVIII–XplottedagainstthehabitationfeaturesinlevelX.The areasingreyarehearths. doi:10.1371/journal.pone.0021545.g002 NeandertalSymbolicBehavioratGrotteduRenne PLoSONE|www.plosone.org4June2011|Volume6|Issue6|e21545

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test—inthiscontext,asimpleconclusionofthepremisethatthe radiocarbonresultsareaccurate. Underthislight,theGrotteduRenne’sradiocarbondating historyisveryinstructive.Giventhat,inagreementwith predictionsmadeonthebasisofarcheologicalcriteria[32],the newagesforProtoaurignacianlevelVIIfellinthe , 34.5–36.5ka14CBPrange,theunderlyingCha ˆtelperronianlevelsVIII–X shouldbechronometricallyolder.However,only2outofthe17 results(12%)previouslypublishedfortheCha ˆtelperronianofthe GrotteduRennesatisfiedsuchacondition[20],whichisthecase foratleast13ofthenewsamples(62%).Thisprogresscanonly havecomefromamelioratedmeasurementandpretreatment techniques(e.g.,ultrafiltration)[33],notfromthesite’sstratigraphicintegrityhavingimprovedovertheyears.Inthiscontext, theparsimoniousexplanationforwhyaminorityofthenewresults cameoutyoungerthanexpectedshouldbebasedon(a)thepoor preservationofcollagenatthesite,asreported[20],and(b)the incompletedecontaminationofanumberofsamples. Theimpactthateventraceamountsofcontaminantshaveon sampleswhoseagesareclosetothelimitofapplicabilityof radiocarbonishuge.Forinstance,ifasampleis40,000yearsold, aslittleas0.6%ofacontaminantdatedtothetimeofthe excavationofthefinds,50yearsago,wouldsufficetoproducea measuredageof35,000.TheGrotteduRenne’sfaunalremains andbonetoolswereextensivelypreservedwithgluesand consolidants,andindeedthiswasthesinglesiteamongthose datedbytheOxford-ledprojecttoassessthechronologyofthe Middle-to-UpperPaleolithictransitioninEurope[34]wheresuch kindsofsamples,otherwiserejectedasunsuitable,wereused—in fact,attheGrotteduRenne,theyrepresentasmuchas84%ofthe determinations. Tocircumventtheproblemsofpotentialcontaminationthe radiocarbonstudyusedasolventsequencephaseofpretreatment beforecollagenextractionwhenevervisualinspectionindicated curatorialpreservation.Inonecase,Poly-VinylAcetate(PVA)was identified.This14C-freematerialcannotbethecauseforresults Figure3.GrotteduRenne,distributionofboneawls,ornamentsandpigmentprocessingtools. Top:inProtoaurignacianlevelVII. Bottom:inCha ˆ telperronianlevelX.Theareasingreyarehearths. doi:10.1371/journal.pone.0021545.g003 NeandertalSymbolicBehavioratGrotteduRenne PLoSONE|www.plosone.org5June2011|Volume6|Issue6|e21545

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youngerthanexpectedbutthestudyalsofoundthatitwas impossibletoruleouttracecontaminationfromthisorother sources.Thatsomeremainedindeedisindicatedby(a)theC:N valueof3.7associatedwithoneoftheCha ˆtelperronianresults (OxA-X-2226-7),whichthestudyaccepteddespiteacknowledging thatitindicatedthepresenceofexogenouscarbon,(b)theC:N valueof3.6,alsoabovetheOxAthresholdforacceptance, obtainedforasamplefromlevelXdatedto , 23.1ka14CBP (OxA-X-2222-21),(c)theverticaldistributionofthefewdatesfor unconsolidatedsamples,whicharefullyconsistentwithstratigraphyand,atthe95.7%confidencelevel,withthe , 36.5ka14CBP boundaryfortheProtoaurignacian(Figure6),(d)thefactthat outliersonlyoccurinsamplesfromlevelIXorbelow(alldatesfor uppermostCha ˆtelperronianlevelVIIIandthoseabovecorrelate perfectlywithstratigraphicdepth),and(e)theresultsfor MousterianlevelXII,youngerthanthoseobtainedforwelldated occurrencesoftheCha ˆtelperronianelsewhereinFrance,namely atRoc-de-CombeandtheeponymoussiteoftheGrottedesFe ´es [8,22]. TheselevelXIIresultsplayacriticalroleintheoutlieranalysis underpinningreference[20]’sinterpretationthattheGrottedu Rennesequenceissignificantlydisturbed.Intheframeworkofthe Bayesianapproachfollowed,theresultsforlevelXIIconstraintheage oftheoverlyingunits,forinstancemakingthedateof48,700 6 360014CBP(OxA-X-2279-44)forlevelXappeartoooldforits stratigraphicpositionand,hence,asanintrusionformtheunderlying Mousterian.Infact,whenthat , 48.7ka14CBPresultisstatistically comparedone-to-onewiththoseforthesamelevelacceptedbythe outlieranalysis(forinstance,usingthetoolsintheCalib6.0software, http://calib.qub.ac.uk/calib/),allcomeoutasidentical. Moreimportantly,acceptingtheoutcomeoftheoutlieranalysis shouldhaveinvalidatedtheuseofBayesianmodelingtoreconstruct thechronologyofthesite—ifsignificantmovementacrosslevels occurred,thestratigraphicpositionofagivensamplecannotbe usedtoconstraintheprobabilitydistributionassociatedwithits determinedage.Thisisbecause,inthatcase,therealchronological orderofthesamples,anapriorirequirementofthemodeling,must beconsideredunknown—oncesignificantpost-depositionaldisplacementisposited,stratigraphicprovenancecannotbetakenas anindicatorofrelativeage.Forinstance,inFigure2ofreference [20],thetwolevelXIIsamplesareusedtodefinethelower chronologicalboundaryoftheCha ˆtelperronianbutthisassumes thatthesamplesareinsitu.However,if,astheauthorsclaim,as muchas30%oftheir31determinationsareoutliersandthisimplies post-depositionaldisturbance,thenwecannotrejectthepossibility thatthosesamplesrepresentinsteadintrusionsfromoverlying Cha ˆtelperronianX,asotherwisesuggestedbythepresenceofa Cha ˆtelperronianboneawlinthatlevel[24]. Tobeconsistentwiththeresultsoftheoutlieranalysis,thestart ofthesequenceinreference[20]’sagemodelshouldbegivenby the , 48.7ka14CBPresultforlevelXbecause,ifthelatteris statisticallydistinctfromtheothersfromthesameleveland deemedtobestratigraphicallydisplaced,thenitcanonlyhave beendisplacedfromtheMousterian.Thiscarriestheimplication thatthebeginningoftheMousterianphaseshouldberepresented bythat , 48.7ka14CBPresult,not,asassumedbythemodel,by thetworesultsinthe , 37–38ka14CBPrangeobtainedforlevel XII,which,inturn,significantlydecreasesthenumberofoutliers. Thedistortionimposedonthedatabyreference[20]’s unwarranteduseofBayesianmodelingalsoimpactstheestimated durationofthesequence.Thisisreadilyapparentwhen comparingitsfigures2(modeledcalibrateddates)andS2 (unmodeledcalibrateddates):intheformer,thesuccessionof levelsXIItoVIspanssome5000years,whilethelattersuggests thatitmayinfactcorrespondinsteadtoasmuchas10,000. Giventheabove,theinterpretationoftheanomalouslyyoung resultsfortheGrotteduRenneshouldbeidenticaltothatfor similaryconsolidatedbonesampleswhosecollagenwassimilarly poorlypreservedandwhosesamplechemistrysimilarlyfailedto meetallthestandardsofthe(same)datinglaboratory—namely, thetwoNeandertalbonesfromVindijadatedatOxford,which treatedthemasminimumagesonly[35].Buteveniftheaccuracy ofallthenewradiocarbonresultsfortheGrotteduRenneis accepted,itcanbecalculated(giventhe38%putativelydisplaced Cha ˆtelperroniansamplesthisimplies,andassumingthatallfinds areequallysusceptibletodisplacement),foraprobabilitythreshold of0.01,thatatleast8ofthe39ornamentsand5oftheNeandertal teethinCha ˆtelperronianlevelsVIII–Xcannotbeintrusive(Text S1).Obviously,thiscalculationignoresthemodelingboundaries imposedbythedistributionofthediagnosticstonetools,asiftheir culturalandchronologicalattributesweremeaninglessandthe Cha ˆtelperronianandProtoaurignacianlabelsvoidtobeginwith. Thepoint,however,isthateventhisunrealisticscenariofailsto rejectthatatleastapartoftheGrotteduRenne’sNeandertal teethandsymbolicartifactsaretrulyinsitu. Finally,wenotethatrecentre-analysisoftheQuinc ¸ay rockshelter,theothermajorCha ˆtelperroniansitefeaturing ornaments(sixpiercedteethofdeer,foxandwolf),showedthat thereislittlegroundtoquestionitsstratigraphicintegrity[36–37]. Asmallcomponentofretouchedbladeletsisfoundinalllevelsbut inassociationwiththecoreswhencethecorrespondingblanks wereextractedandthosecoresareoftypicalCha ˆtelperronian technique.Thismaterialcannotrepresentintrusionfromoverlying Aurignacianlevels,whicharenon-existentatQuinc ¸ay,wherethe Cha ˆtelperroniansequencewassealedbycollapsedlimestoneslabs severalmeterslongand , 50cmthick.ConclusionOurresultsrejectthenotionthattheassociationofsymbolic artifactswithNeandertalsattheGrotteduRenneresultsfrom largescalelocalizedorsmallscalegeneralizeddisplacementof artifactsandhumanremains.Theyimplythattheparsimonious explanationfortheanomaliesobservedintheradiocarbondating ofthesequenceisincompletedecontaminationofthebone samplesused.Moreover,theGrotteduRenneisnotalone. Similar,andevenearlierevidenceofNeandertalsymbolically mediatedbehaviorshasnowbeenproducedfortheMiddle PaleolithicofIberiaandItaly[9,38],andthereissufficientground topostulatethatcoevalmaterialfromtheFrenchCha ˆtelperronian, theItalianUluzzian[39],andanumberofsitesinCentraland WesternEuropewithlessclear-cutstratigraphicpatterns[30,31]is alsoNeandertal-associated.WhilerecentGrotteduRenne researchdevelopmentshavebeenclaimedtoheraldthebursting oftheNeandertalornament‘‘bubble’’[40],ourresultsandtheir widercontextshowthatsuchnewswere‘‘greatlyexaggerated.’’Figure4.DistributionoftheNeandertalremainsinCha ˆ telperronianlevelsVIII–XoftheGrotteduRenne;after [ 5,19 ] ,modified. In theplan(above),theblackcirclesarediagnosticteeth,thewhitecirclesareteethwhosefeaturesareconsistentwith(butdonotprove)assignment totheNeandertals,thewhitesquareistheimmaturetemporalbone,andtheredlinesindicatethepositionofthestratigraphicprofilesshownmiddle andbottom. doi:10.1371/journal.pone.0021545.g004 NeandertalSymbolicBehavioratGrotteduRenne PLoSONE|www.plosone.org7June2011|Volume6|Issue6|e21545

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TheCha ˆtelperronianlevelsoftheGrotteduRennedostandfor Neandertalsymbolism.MaterialsandMethodsToassesspost-depositionaldisplacementofhumanremainswe usedlooseteeth(giventheirsmallsizeandmobilityandthefact that,exceptforatemporalfragment,nootherhumanskeletal partswererecoveredattheGrotteduRenne),andonlyconsidered thoseunambiguouslyprovenancedtoaspecificgridunit.In ornamentcounts,fragmentsofpossiblythesameobjectwere consideredseparately.Thesecriteriaexplaintheminordifferences withpreviouslypublishedinventories[8,30,31].Weuseda continuoustimemodelpositingthatthedifferentitemscanmove Figure5.SpatialdistributionofcolorantsbyweightinCha ˆ telperronianlevelX(allsublevelsincluded);datafrom [ 27 ] . Topleft,all colors(14,580g);topright,red(10,520g);bottomleft,black(3,961g);bottomright,yellow(99.6g).Bubblesizesreflecttherelativefrequency per gridunitandwerecalculatedbyassigningthefollowingvaluestothefourweightclassesgiveninthedatasource:[0.1–10g]=10g;[10–50g] =50g;[50–100g];=100g;[ . 100g]=150g.Majorconcentrationsareapparentdespitethesmoothingofthedistributionscausedbythe quantificationprocedureandtheycoincidewiththehabitationfeatures.Thegreyareasarehearths. doi:10.1371/journal.pone.0021545.g005 NeandertalSymbolicBehavioratGrotteduRenne PLoSONE|www.plosone.org8June2011|Volume6|Issue6|e21545

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fromoneleveltothenextatagivenrate,whichweassumetobe thesameforboththedirectionofthemovement(upwardor downward)andthecategoryconcernedbecausenosignificant differenceinphysicalproperties(size,weightordensity)existsthat justifiesdiscriminatinginthisregardbetween,say,aDufour bladelet,aNeandertaltoothorapiercedfoxcanine.Fromtopto bottom,andusingavailableradiometricinformation[20],weset theuppertimelimitsfortheformationofthedifferent stratigraphicunitsat40.5(VII),41.5(VIII),43.0(IX),44.5(X), 46.0(XI),47.5(XII),49.0(XIII)and50.5(XIV)thousand calendaryearsago(butfoundtheresultsoftheexercisetobe unchangedeveniftheageofthelower,undatedunitswasmoved backintimebyseveralmillennia).Wethenestimated,foreach hypothesis,therateparameterthatbestfitted,accordingtothe Pearsonchisquarestatistic[41,42],thetransformationbetween theinitialandfinalconditions.Thisbestfittedrateparameter providesthebesttrade-offregardingobjectsthathavenotmoved (e.g.Dufourorunretouchedbladelets)andobjectsthathave moved(e.g.Neandertalteethorornaments)underthedifferent hypotheses.Wethenconductedagoodnessoffittesttoassessthe likelihoodthatthefinaldistributionwasgeneratedfromthe hypothesizedinitialdistributions(TableS1)viathemodel.Using thebestfittedrateparameterforeachhypothesis,weobtained expectedfinaldistributionsforeachfindcategoryandprobabilities thatanygivenitemmovedfromanygivenleveltoanother(Tables S2,S3).Thesedistributionswerethencomparedwiththeobserved finaldistributions(Table1)toassessthegoodnessoffitofthe modelassociatedwitheachhypothesis.Thisgoodnessoffitis measuredbythePearsonchisquarestatistic,whichprovidesa positivemeasureofdiscrepancybetweentheexpectedandtrue finaldistributionsofobjects.Havingconcludedthatsmallscale, generalizeddisplacementfailedtoaccountforsignificantmovementinsomefindcategoriesandnoneinothers,wethen calculated,usingtheBayestheorem[43],theprobabilitythat itemsfromthekeyfindcategories(Neandertalteethandpersonal ornaments)hadmovedintotheCha ˆtelperronianfromthe overlyingProtoaurignacianortheunderlyingMousterianunder theconstraintsposedbythedistributionofdiagnosticstonetools. Atechnicaldescriptionofourstatisticalapproachisprovidedin TextS1. Table2. Stratigraphicdistributionofcolorants(byweight,in grams)atGrotteduRenne;datafrom[27].LevelsCulturalAttributionRedBlackYellowTotal% VGravettian1179.414.513.91207.86.0 VIAurignacian/Gravettian196.51.5147.2345.21.7 VIIProtoaurignacian404.429.517.3451.22.2 VIIICha ˆ telperronian966.2231.728.21226.16.1 IXCha ˆ telperronian1749.3476.417.22242.911.1 XCha ˆ telperronian10,520.2396199.614,580.872.4 XIMousterian48.5171277.50.4 TOTAL15,064.54731.6335.420,131.5100.0 doi:10.1371/journal.pone.0021545.t002 Figure6.Plotofage(meanvalueand2 s confidenceinterval)againststratigraphicprovenienceofthenewradiocarbondatesfor theGrotteduRenne;datafrom [ 20 ] . LevelsVIII–XareCha ˆ telperronian. doi:10.1371/journal.pone.0021545.g006 NeandertalSymbolicBehavioratGrotteduRenne PLoSONE|www.plosone.org9June2011|Volume6|Issue6|e21545

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SupportingInformationTextS1Hypothesistesting,continuoustimemodeland probabilitymodels. (PDF)FigureS1Realizationfromthecontinuoustimemodel, foranobjectstartinginlevelXI. (TIFF)FigureS2a: S infunctionof l forHypothesis1;b: S infunction of l forHypothesis2,c: S infunctionof l forHypothesis3. (TIF)FigureS3a:probabilitythatkornamentsoverasetof47(the totalnumberofornamentsfromlevelsVIIandVIII–X)are intrusive,giventhenumberofintrudedDufourbladeletsand unretouchedbladelets;b:probabilitythatkNeandertalteethover asetof31(thetotalnumberofNeandertalteethfromlevelsXI– XIIandVIII–X)havemovedfromlevelsXI–XIItolevelsVIII– X,giventhenumberofLevalloisflakesthathavemoved;c: probabilitythatksamplesoverasetof26(thetotalnumberof datedsamplesfromlevelsVIIandVIII–X)areintrusive,giventhe numberofintrudedDufourbladeletsandunretouchedbladelets. (TIF)TableS1Predictedinitialdistributionoffindsforthe differenttestedhypotheses. (DOC)TableS2Expectedfinalvalues E (rounded)associated tothebestfitted l forthedifferenttestedhypotheses. (DOC)TableS3Values p associatedtothebestfitted l forthe differenthypotheses. (DOC)AcknowledgmentsWethankHe ´le `neSalomon,forallowingustouseunpublishedimagesof GrotteduRennecolorants,andMiche `leJulienandMarianVanhaerenfor givinguspermissiontouseimagesofpersonalornamentsandforproviding updatedinformationontheirspatialdistribution.AuthorContributionsConceivedanddesignedtheexperiments:FCFDPDFSJZ.Performedthe experiments:FC.Analyzedthedata:FCFDPDJZ.Wrotethepaper:FC FDJZ.References1.Leroi-GourhanA(1964)LesreligionsdelaPre ´histoire.Paris:Presses UniversitairesdeFrance.164p. 2.DeGiorgioM,JakobssonM,RosenbergNA(2009)Explainingworldwide patternsofhumangeneticvariationusingacoalescent-basedserialfounder modelofmigrationoutwardfromAfrica.ProcNatlAcadSciUSA106: 16057–16062. 3.RightmireP(2009)MiddleandlaterPleistocenehomininsinAfricaand SouthwestAsia.ProcNatlAcadSciUSA106:1604–1605. 4.d’ErricoF,StringerC(2011)Evolution,RevolutionorSaltationscenarioforthe emergenceofmoderncultures?PhilosophicalTransactionsoftheRoyalSociety B:BiologicalSciences366,1567:1060–1069. 5.BaileySE,HublinJ-J(2006)DentalremainsfromtheGrotteduRenneatArcysur-Cure(Yonne).JHumEvol50:485–508. 6.JacobsZ,RobertsR,GalbraithR,DeaconH,Gru ¨nR,etal.(2008)Agesforthe MiddleStoneAgeofSouthernAfrica:ImplicationsforHumanBehaviorand Dispersal.Science322:733–735. 7.d’ErricoF,Zilha ˜oJ,JulienM,BaffierD,PelegrinJ,etal.(1998)Neanderthal acculturationinwesternEurope?Acriticalreviewoftheevidenceandits interpretation.CurrAnthropol39:S1–S44. 8.Zilha ˜oJ(2006)Neandertalsandmodernsmixed,anditmatters.EvolAnthropol 15:183–195. 9.Zilha ˜oJ,AngelucciD,Badal-GarciaE,d’ErricoF,DanielF,etal.(2010) SymbolicuseofmarineshellsandmineralpigmentsbyIberianNeandertals. ProcNatlAcadSciUSA107:1023–1028. 10.ConardN(2008)AcriticalviewoftheevidenceforasouthernAfricanoriginof behavioralmodernity.SouthAfArchaeolSoc10:175–179. 11.HoversE(2009)TheMiddle-to-UpperPaleolithicTransition:WhatNews?In: CampsM,ChauhanP,eds.SourcebookofPaleolithicTransitions:Methods, theories,andinterpretations.NewYork:Springer.pp455–462. 12.LangleyM,ClarksonC,UlmS(2008)BehaviouralComplexityinEurasian NeanderthalPopulations.CambridgeArchaeologicalJournal18:289–307. 13.NowellA(2010)DefiningBehavioralModernityintheContextofNeandertal andAnatomicallyModernHumanPopulations.AnnuRevAnthropol39: 437–452. 14.Leroi-GourhanArl,Leroi-GourhanA(1964)Chronologiedesgrottesd’Arcy-surCure(Yonne).I:Climatsduquaternairere ´cent.II:IndustriesduPale ´olithique supe ´rieur.GalliaPre ´histoire7:36–64. 15.SchmiderB(2002)L’organisationdel’habitataurignacien.In:SchmiderB,ed. L’AurignaciendelaGrotteduRenne.Paris:CNRSe ´ditions.pp15–24. 16.DavidF,ConnetN,GirardM,LhommeV,MiskovskyJ-C,etal.(2001)Le Cha ˆtelperroniendelagrotteduRennea `Arcy-sur-Cure(Yonne).Donne ´es se ´dimentologiquesetchronostratigraphiques.BulletindelaSocie ´te ´Pre ´historiqueFranc ¸aise98:207–230. 17.WhiteR(2001)PersonalornamentsfromtheGrotteduRenneatArcy-surCure.AthenaRev2:41–46. 18.Bar-YosefO(2006)Neanderthalsandmodernhumans:Adifferentinterpretation.In:ConardNJ,ed.WhenNeanderthalsandModernHumansMet. Tu ¨bingen,Germany:Kerns-Verlag.pp467–482. 19.Bar-YosefO,BordesJG(2010)WhowerethemakersoftheCha ˆtelperronian culture?JHumEvol59:586–593. 20.HighamT,JacobiR,JulienM,DavidF,BasellL,etal.(2010)Chronologyofthe GrotteduRenne(France)andimplicationsforthecontextofornamentsand humanremainswithintheCha ˆtelperronian.ProcNatlAcadSciUSA107: 20234–20239. 21.Leroi-GourhanA(1968)Lepetitracloircha ˆtelperronien.InLaPre ´histoire,Proble `mesetTendances.InMazie `resM,ed.Paris:CNRSe ´ditions.pp275–282. 22.Zilha ˜oJ,d’ErricoF,BordesJ-G,LenobleA,TexierJ-P,etal.(2006)Analysisof AurignacianinterstratificationattheCha ˆtelperronian-typesiteandimplications forthebehavioralmodernityofNeandertals.ProcNatlAcadSciUSA103: 12643–12648. 23.BrantinghamPJ,ToddA,SurovellTA,Waguespack,NA(2007)Modelingpostdepositionalmixingofarchaeologicaldeposits.JAnthropolArchaeology26: 517–540. 24.d’ErricoF,JulienM,LioliosD,VanhaerenM,BaffierD(2003)Manyawlsin ourargument.BonetoolmanufactureanduseintheCha ˆtelperronianand AurignacianlevelsoftheGrotteduRenneatArcy-sur-Cure.In:Zilha ˜oJ, d’ErricoF,eds.TheChronologyoftheAurignacianandoftheTransitional Technocomplexes.Dating,Stratigraphies,CulturalImplicationsLisbon,Portugal:InstitutoPortugue ˆsdeArqueologia.pp247–270. 25.JulienM,BaffierD,LioliosD(2002)L’outillageenmatie`resduresanimales.In: SchmiderB,ed.L’AurignaciendelaGrotteduRenne.Paris:CNRSe ´ditions.pp 217–250. 26.TaborinY(2002)Lesobjetsdeparure.InSchmiderB,ed.L’Aurignaciendela GrotteduRenne.Paris:CNRSe ´ditions.pp253–256. 27.CouraudC(1991)Lespigmentsdesgrottesd’Arcy-sur-Cure(Yonne).Gallia Pre ´histoire33:17–52. 28.SalomonH(2009)Lesmatie `rescolorantesaude ´butduPale ´olithiquesupe ´rieur: sources,transformationsetfonctions.PhDdissertation,UniversityofBordeaux. 29.deBeauneS(2002)L’outillagesurgalet.In:SchmiderB,ed.L’Aurignaciendela GrotteduRenne.Paris:CNRSe ´ditions.pp199–213. 30.Zilha ˜oJ(2007)Theemergenceofornamentsandart:anarcheological perspectiveontheoriginsof‘BehavioralModernity’.JArchRes15:1–54. 31.Zilha ˜oJ(2011)Aliensfromoutertime?Whythe‘‘HumanRevolution’’iswrong andwheredowegofromhere?InCondemiS,WenigerG-C,eds.Continuity andDiscontinuityinthePeoplingofEurope.NewYork:Springer.pp331–366. 32.Zilha ˜oJ,d’ErricoF(2003)AnAurignacian‘‘GardenofEden’’insouthern Germany?AnalternativeinterpretationoftheGeissenklo ¨sterleandacritiqueof theKulturpumpemodel.Pale ´o15:69–86. 33.BronkRamseyC,HighamT,BowlesA,HedgesR(2004)Improvementstothe pretreatmentofboneatOxford.Radiocarbon51:337–3360. 34.HighamT(2011)EuropeanMiddleandUpperPalaeolithicradiocarbondates areoftenolderthantheylook:problemswithpreviousdatesandsomeremedies. Antiquity85:235–249. 35.HighamT,etal.(2006)ReviseddirectradiocarbondatingoftheVindijaG1 UpperPaleolithicNeandertals.ProcNatlAcadSciUSA103:553–557. 36.GrangerJM,Le ´ve ˆqueF(1997)Parurecastelperronienneetaurignacienne:e ´tude detroisse ´riesine ´ditesdedentsperce ´esetcomparaisons.C.R.Acad.Sci.Paris 325:537–543. 37.RousselM,SoressiM(2010)LaGrandeRochedelaPle ´matriea `Quinc ¸ay (Vienne).L’e ´volutionduCha ˆtelperronienrevisite ´e.In:Buisson-CatilJ,NeandertalSymbolicBehavioratGrotteduRenne PLoSONE|www.plosone.org10June2011|Volume6|Issue6|e21545

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PrimaultJ,eds.Pre ´histoireentreVienneetCharente-Hommesetsocie ´te ´sdu Pale ´olithique.Villefranche-de-Rouergue:AssociationdesPublicationsChauvinoises.pp203–219. 38.PeresaniM,FioreI,GalaM,RomandiniM,TagliacozzoA(2010)LateNeandertals andtheintentionalremovaloffeathers asevidencedfrombirdbonetaphonomyat FumaneCave44kyB.P.,Italy.ProcNatlAcadSciUSA108:3888–3893. 39.RielSalvatoreJ(2009)Whatisatransitionalindustry?TheUluzzianofsouthern Italyasacasestudy.In:CampsM,ChauhanP,eds.SourcebookofPaleolithic Transitions.NewYork:Springer.pp377–396. 40.MellarsP(2010)Neanderthalsymbolismandornamentmanufacture:the burstingofabubble?ProcNatlAcadSciUSA23:20147–20148. 41.AgrestiA(2002)CategoricalDataAnalysis.NewYork:Wiley-Interscience. 42.WassermanL(2003)AllofStatistics:AConciseCourseinStatisticalInference. NewYork:Springer. 43.BernardoJM,SmithAFM(2000)BayesianTheory.NewYork:WileyandSons.NeandertalSymbolicBehavioratGrotteduRenne PLoSONE|www.plosone.org11June2011|Volume6|Issue6|e21545


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