Radiocarbon dates from the Grotte du Renne and Saint-Césaire support a Neandertal origin for the Châtelperronian


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Radiocarbon dates from the Grotte du Renne and Saint-Césaire support a Neandertal origin for the Châtelperronian

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Radiocarbon dates from the Grotte du Renne and Saint-Césaire support a Neandertal origin for the Châtelperronian
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Proceedings of the National Academy of Sciences of the United States of America
Creator:
Hublin, Jean-Jacques
Talamo, Sahra
Julien, Michèle
David, Francine
Connet, Nelly
Bodu, Pierre
Vandermeersch, Bernard
Richards, Michael P.
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National Academy of Sciences
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Neanderthals ( lcsh )
Radiocarbon dating ( lcsh )
Paleolithic period ( lcsh )
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serial ( sobekcm )
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Europe -- France -- Occitanie -- Saint-Césaire

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The transition from the Middle Paleolithic (MP) to Upper Paleolithic (UP) is marked by the replacement of late Neandertals by modern humans in Europe between 50,000 and 40,000 y ago. Châtelperronian (CP) artifact assemblages found in central France and northern Spain date to this time period. So far, it is the only such assemblage type that has yielded Neandertal remains directly associated with UP style artifacts. CP assemblages also include body ornaments, otherwise virtually unknown in the Neandertal world. However, it has been argued that instead of the CP being manufactured by Neandertals, site formation processes and layer admixture resulted in the chance association of Neanderthal remains, CP assemblages, and body ornaments. Here, we report a series of accelerator mass spectrometry radiocarbon dates on ultrafiltered bone collagen extracted from 40 well-preserved bone fragments from the late Mousterian, CP, and Protoaurignacian layers at the Grotte du Renne site (at Arcy-sur-Cure, France). Our radiocarbon results are inconsistent with the admixture hypothesis. Further, we report a direct date on the Neandertal CP skeleton from Saint-Césaire (France). This date corroborates the assignment of CP assemblages to the latest Neandertals of western Europe. Importantly, our results establish that the production of body ornaments in the CP postdates the arrival of modern humans in neighboring regions of Europe. This new behavior could therefore have been the result of cultural diffusion from modern to Neandertal groups.
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Volume 109, Issue 46
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6 p.

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RadiocarbondatesfromtheGrotteduRenneand Saint-CésairesupportaNeandertaloriginfor theChâtelperronian Jean-JacquesHublin a,1 ,SahraTalamo a ,MichèleJulien b ,FrancineDavid b ,NellyConnet b,c ,PierreBodu b , BernardVandermeersch d ,andMichaelP.Richards a,e a DepartmentofHumanEvolution,MaxPlanckInstituteforEvolutionaryAnthropology,04103Leipzig,Germany; b UnitéMixtedeRecherche7041duCentre NationaldelaRechercheScienti que,ArchéologiesetSciencesdel ’ Antiquité,92023Nanterre,France; c Institutnationalderecherchesarchéologiques préventives(INRAP),86000Poitiers,France; d UnitéMixtedeRecherche5199duCentreNationaldelaRechercheScienti que — Préhistoireàl ’ Actuel:Culture, EnvironnementetAnthropologie,UniversitéBordeaux1,33405Talencecedex,France;and e DepartmentofAnthropology,UniversityofBritishColumbia, Vancouver,BC,CanadaV6T1Z1 EditedbyRichardG.Klein,StanfordUniversity,Stanford,CA,andapprovedSeptember24,2012(receivedforreviewJuly27,2012) ThetransitionfromtheMiddlePaleolithic(MP)toUpperPaleolithic(UP)ismarkedbythereplacementoflateNeandertalsby modernhumansinEuropebetween50,000and40,000yago. Châtelperronian(CP)artifactassemblagesfoundincentralFrance andnorthernSpaindatetothistimeperiod.Sofar,itistheonly suchassemblagetypethathasyieldedNeandertalremainsdirectly associatedwithUPstyleartifacts.CPassemblagesalsoinclude bodyornaments,otherwisevirtuallyunknownintheNeandertal world.However,ithasbeenarguedthatinsteadoftheCPbeing manufacturedbyNeandertals,siteformationprocessesandlayer admixtureresultedinthechanceassociationofNeanderthal remains,CPassemblages,andbodyornaments.Here,wereport aseriesofacceleratormassspectrometryradiocarbondateson ultra lteredbonecollagenextractedfrom40well-preservedbone fragmentsfromthelateMousterian,CP,andProtoaurignacian layersattheGrotteduRennesite(atArcy-sur-Cure,France).Our radiocarbonresultsareinconsistentwiththeadmixturehypothesis.Further,wereportadirectdateontheNeandertalCPskeleton fromSaint-Césaire(France).Thisdatecorroboratestheassignment ofCPassemblagestothelatestNeandertalsofwesternEurope. Importantly,ourresultsestablishthattheproductionofbody ornamentsintheCPpostdatesthearrivalofmodernhumansin neighboringregionsofEurope.Thisnewbehaviorcouldtherefore havebeentheresultofculturaldiffusionfrommoderntoNeandertalgroups. stimulusdiffusion | culturalmodernity C rucialtounderstandingthereplacementprocessesofNeandertalsbymodernhumansinwesternEurasiaatthebeginningoftheUpperPaleolithic(UP)istheinterpretationofsocalled “ transitionalindustries. ” Inarcheologicalsites,theselithic assemblagesarefoundundervarioustypesofearlyUP(mostly Aurignacian)and/orabovegenuineMiddlePaleolithic(MP) assemblages.TheydisplayvariousproportionsofUPartifacts andtechnologies,butinseveralcases,adirectorigininthelocal MPhasbeenargued(1,2).Although,inEurope,onlyNeandertalremainshavebeenfoundinassociationwithMousterian industriesandonlyanatomicallymodernhumanremainshave beensecurelyidenti edinAurignacianlayers,thebiological identityofthemakersofthetransitionalindustriesremainsunclear(3). ThatNeandertalsmadesomeofthetransitionalindustrieshas attimesbeensuggestedbasedonfragmentarypaleontological evidence(4,5)oronindirectchronologicalarguments(6). However,todate,onlytheChâtelperronian(CP),atransitional assemblagefromcentralandsouthwesternFrance,andnorthern Spain,hasyieldedwell-identi edandrelativelyabundantNeandertalremains,speci callyfromtwoFrenchsites(LaGrotte duRenneandSaint-Césaire)(7 – 9). Despitethisfossilevidence,thequestionofwhetherNeandertalsmanufacturedtheCPremainsthetopicofintensedebate. ThisresultsinpartfromthefactthattheCPistheso-called “ transitional ” assemblagethatdocumentsthebroadestspectrum ofbehavioralfeaturesreminiscentofthesubsequentlocalUP, whichwasundisputedlycreatedbymodernhumans.Thesefeaturesincludetheproductionofbladelets(10),boneartifacts,and bodyornaments(11,12),allofwhicharevirtuallyunknownin theNeandertalworld.Furthermore,thesebodyornaments,often consideredasanexpressionofadvancedsymbolicbehavior,have onlybeenfoundintheCPoftheGrotteduRenneandQuinçay, bothatthenorthernmostlimitoftheCPgeographicaldistribution( Fig.S1 ). Whenaccepted,theassociationoftheseobjectswithNeandertalshasbeeninterpretedintwoways.Forsome(e.g.,refs.8 and13 – 15),thisnewbehaviordemonstratesaculturalimpacton thelastNeandertalsbycontemporaneousUPmodernhuman populations,alreadypresentfurthereastinEurope.Forothers (16),itrepresentsaNeandertalinventionoftheirownUP-style industryindependentlyofmodernhumans. AthirdgroupofauthorsspeculatesthattheNeandertal/CP associationmayinfactresultfromvarioussiteformationprocessesincludinglayeradmixture(17).FortheGrotteduRenne speci callyithasbeensuggestedthattheNeandertalremains mighthavebeendugoutoftheunderlyingMousterianlayersby CPmodernhumans(17)and/orthatCPbodyornamentsand otherboneartifactsmayhavemoveddownfromtheoverlaying UPlayers(18).AtSaint-Césaire,thestratigraphicintegrityofthe CPdepositshasalsobeenchallenged(17). Radiometricargumentsarerelevanttotheassessmentof possiblemixingofyoungerorolderintrusivematerialintothe CPlayers.TheprecisedatingoftheCPisalsocentraltothe discussionsurroundingitsinterpretation.Inparticular,thenotionthatthedevelopmentoftheCPresultedfromthecultural in uenceofmodernpopulationsimpliesthatCPinnovations postdatethearrivalofmodernhumansinEurope.TheradiometricdatingoftheCPattheGrotteduRenneandinothersites haslongbeenchallengingbecausetheseassemblageslayatthe limitoftheapplicationrangeofthe 14 Cmethod.Inparticular, contaminationproblemshaveimpairedmostofthedatesobtained Authorcontributions:J.-J.H.designedresearch;J.-J.H.,S.T.,M.J.,F.D.,N.C.,andP.B.performedresearch;S.T.,B.V.,andM.P.R.contributednewreagents/analytictools;J.-J.H., S.T.,M.J.,F.D.,N.C.,P.B.,andM.P.R.analyzeddata;andJ.-J.H.andS.T.wrotethepaper. Theauthorsdeclarenocon ictofinterest. ThisarticleisaPNASDirectSubmission. FreelyavailableonlinethroughthePNASopenaccessoption. 1 Towhomcorrespondenceshouldbeaddressed.E-mail:hublin@eva.mpg.de. Thisarticlecontainssupportinginformationonlineat www.pnas.org/lookup/suppl/doi:10. 1073/pnas.1212924109/-/DCSupplemental . www.pnas.org/cgi/doi/10.1073/pnas.1212924109PNAS | November13,2012 | vol.109 | no.46 | 18743 – 18748 ANTHROPOLOGY

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beforerecenttechnicalimprovementsinthepretreatmentof samples,whichhaveincreasedthereliabilityof 14 Cdatesbeyond 40ka 14 CBP(19 – 21). Inthiscontext,aseriesof31acceleratormassspectrometry (AMS)ultra ltereddatesonbones,antlers,boneartifacts,and teethfromkeyarchaeologicallayersoftheGrotteduRenne havebeenpublished(22).Thisstudyshowedanunexpectedly highdegreeofintralayervariationintheradiometricagesand broughtsomesupporttotheadmixturehypothesis. Incontrast,asubsequentreinvestigationofthehorizontaland verticaldistributionofvariouscategoriesofarchaeologicallydiagnosticobjectsattheGrotteduRennedidnotsupportthe hypothesisoflargeorsmallscalepostdepositionalmixing.It insteadsuggestedthatincompletesampledecontaminationisthe mostparsimoniousexplanationfortheobservedradiometric anomalies(11).Inaresponse,theauthorsoftheAMSdatesdid notdisputethisspatialevidencebutalsofoundnogroundforthe claimofcontaminationbyconsolidants(23).Thedebatealso focusedonthede nitionofthestratigraphicunitstobeusedin theBayesianmodelingoftheradiocarbondates(24,25). Totestthesealternativeinterpretationsoftheassociationat GrotteduRenneofNeandertalswiththeCPandUPtype artifacts,weproducedaseriesofradiocarbonagesfromthe differentarchaeologicallayersofthesiteusingthesamepretreatmentprotocolsbutadifferentsamplingstrategy.Wealso comparedtheseageswithadirectdateoftheNeandertalhuman remainsfromthesiteofSaint-Césaire( Fig.S2 ). GrotteduRenneStratigraphy TheGrotteduRenneisoneoftherichestknownCPsites.Itwas depositedattheendofasmallkarsticgallery,formingacave opentowardthesouth,overtheCureRiver.Onanextensionof approximately10m,fromthebackofthecavetotheupperlimit ofasteeptalusfallingtowardtheriver,thesitecontainssubhorizontal,strati eddeposits( Fig.S3 ).Intheupperpartofthe talus,thisstratigraphycanstillbefollowed,butitisreducedto asequenceofthinobliquelayers.Thesitewasexcavatedbetween1949and1963byA.Leroi-Gourhanusingadvancedexcavationtechniquesforthistimeperiod.Amongotherinnovations, theteamattemptedtorecognizeandexposeoccupationsurfaces. Thearcheologicalmaterial,andtheobservedstructures,were spatiallyrecordedandthesedimentssystematicallysieved. AttheGrotteduRenne,14stratigraphicunitswereoriginally recognized(26),ofwhich5werethesourceofthedatedmaterial reportedinthispaper.LayerXIistheuppermostMousterian layer.LayersXandIXyieldedmostoftheCPmaterialfromthe siteandalsocontainseveralhearthsofvarioussizes.Thesetwo layersareformedbyanaccumulationoflimestoneslabs.Inone areainthebackofthecave,theyreachathicknessof80cmbut areingeneralmuchthinner,especiallyclosetothetalus.Layer IX,inparticular,isquiteirregularasitdisplaysathicknessof only3cminsomeareas( Fig.S3 ).InlayerX,occupation oors (sublayers)weretentativelyidenti edandnumbered(27). However,theiridenti cationandrecognitionwasnotalways straightforwardandseveralartifactre tsweredetectedbetween thesublayersofX(28)( Fig.S4 ).Twohavealsobeendetected betweenlayersXandIX( Fig.S4 ).IncontrasttolayersX – IX, theuppermostCPlayer(VIII)displaysalowerdensityofartifacts.Itiscomposedofyellowclayishgravel,withouthearths, andhasyieldednumerousbonesofhyenasandcavebears, suggestingonlyintermittenthumanoccupation(26).OverCP layerVIII,layerVIIhasyieldedaProtoaurignacianassemblage, whichistodatethenorthernmostassemblageofthistype identi edinwesternEurope(29). Methods Weselected40bonesamplesfromlayersXI – VIIoftheGrotteduRenne(Table 1),allfromdifferentareasofthehorizontalsectionofthesite.Weespecially targetedthoseareasthatyieldedCPbodyornamentsordiagnosticNeandertalremains( Fig.S5 ).Thirty-onesamplesarefromtheCPdeposits,4are fromtheunderlyingMousterianand5arefromtheProtoaurignacian. Ourprimaryselectioncriterionwasgoodbonepreservation.Thus,wedid notsampleworkedelementsbecausetheyoftenhavelittlewell-preserved compactboneandmayhavebeentreatedwithunrecordedconsolidants, whichcanaltertheradiocarbonage.Wesampledalargenumberofthick, untreatedpiecesofcompactbonefromshafts,mostlyofreindeerandhorse, toassesspossiblemovementsofmaterialbetweenstratigraphiclayersandto comparetheiragestothoseofworkedartifactspublishedforthesite(22). Additionally,35%oftheselectedfragmentsdisplayedanthropogenicmarks (cut-marksorpercussionmarks). Extractionofcollagen,isotopicmeasurements,andgraphitizationwere performedattheDepartmentofHumanEvolution,MaxPlanckInstitutefor EvolutionaryAnthropology,inLeipzig,Germany.Allbonesamplespresented hereweresubjecttothefollowingpretreatmentprocedures(19).Samples were rstcleanedbysandblasting,andthen500mgofbonepowderwas taken.Thesamplesweredecalci edin0.5MHClatroomtemperature. Later,0.1MNaOHwasaddedtoremovehumics,andthetreatmentwas completedwiththeadditionof0.5MHCl.TheresultingsolidwasgelatinizedafterLongin(30,31)atpH3inaheaterblockat75°C.Thegelatinwas lteredinanEeze-Filter(ElkayLaboratoryProducts)toremovesmall( < 8 m) particlesandthenultra lteredwithaSartorius “ Vivaspin15 ” 30-kDa ultra lter.Beforeuse,the lterwascleanedtoremovecarboncontaining humectants.Finally,thesampleswerelyophilized. ThecollagenextractsweremeasuredfortheirC:Nratios,%Cand%N,and carbonandnitrogenstableisotopevaluesbyusingaThermiFinniganFlashEA coupledtoaDeltaVisotoperatiomassspectrometer.Thecriteriafordeterminingwhetherthecollagenwaswell-preservedand,therefore,couldbe radiocarbondatedwasacollagenyieldofabove1%andaC:Nratiobetween2.9and3.6(32 – 35).Afterthisdetermination,thecollagensamples weregraphitized. ThegraphitewassenttotheKlaus-Tschira-Accelerator-Mass-Spectrometry (AMS)facilityoftheCurt-EngelhornCentreinMannheim,Germany,forAMS measurements.Alldateswerecorrectedforaresidualpreparationbackgroundestimatedfrompretreated 14 C-freebonesamples,kindlyprovided bytheOxfordRadiocarbonAcceleratorUnit(ORAU). AtibialfragmentofcompactbonefromtheSaint-CésaireNeanderthal wasobtainedtoperformisotopicandpaleogeneticstudies.Itsisotopic valuesandC:Nratiowere rstdeterminedattheMax-PlanckInstitutefor EvolutionaryAnthropologyandwerefoundtobeinacceptableranges. SubsequentlyasubsampleofthebonewassenttotheORAUforcommercialradiocarbondating,andpretreatmentofthebonewasundertaken thereusingtheirstandardprocedures(21).TheisotopicvaluesandradiocarbondatesaregiveninTable1. UsingthestratigraphicinformationandtheOxCal4.1program,which calibratesthedatesbyusingIntCal09(36,37),webuiltaBayesianmodel forthedistributionofages.Thismodelisbasedonthedistinctionoffour separatephasesattheGrotteduRennesite,whichtakeintoaccountthe archeologicalobservationsandtheartifactre ts.Thesephasesarethe Mousterian(layerXI),theCP(combininglayersXandIX),thelaterCP(layer VIII),andtheProtoaurignacian(layerVII).Theuniformdistributionofthe radiocarbonageswastestedbyusingagreementindices.Thisindexshows howthesimplecalibrateddistributionagreeswiththedistributionafter Bayesianmodelingandisexpectedtobemorethan60%whenthedatesare inagreementwiththestratigraphy(37).Thet-typeoutlierdetectionwith priorprobabilitiessetat0.05withintheBayesianmodel(37)wasalsoused todetectproblematicsamples. Results AttheGrotte-du-Renne,alloftheisotopicvaluesandtheC:N ratioarewellwithintheacceptedranges(32 – 35)andmostofour samplesdisplayhighyieldsofcollagen,rangingbetween2and 7%(Table1).Oneofthe40sampleshadacollagenyieldof slightlylessthan1%.However,itdisplayedanormalC:Nratio, wassentfordating,andprovidedadateingoodagreementwith samplesfromthesamelayer. Obtaineddatesrangebetween40,900and43,230 14 CBP( n = 4) fortheMousterianphase,35,500and40,970 14 CBP( n = 26) fortheCPphase,35,380and37,710 14 CBP( n = 5)forthelater CPphase,and29,930and34,810 14 CBP( n = 5)fortheProtoaurignacianphase.Thecalibratedrangesofboundariesand theircon denceintervalsarelistedin DatasetS1 .CPandlaterCP 18744 | www.pnas.org/cgi/doi/10.1073/pnas.1212924109 Hublinetal.

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togetherrepresentaperiodtwiceasshortastheProtoaurignacian. ThelaterCPalonelastedonlyapproximately500y,whichmay alsoexplainoverlapsindatesbetweenourtwoCPphases. OxCal ndsanexcellentagreementindex(A_overall = 113.5%, whenthepriorprobabilityoftwooutliersdiscussedbelowissetto 100)betweenthefullsetofdatesandthestratigraphicinformation(Fig.1).Theresultsoftheoutlierdetectionmethod areshowninFig.1(aswellas DatasetS2 ).Onlytwosamples, EVA-29andEVA-56,althoughwe llwithintheacceptedrange forcollagenyieldandC:Nratio,displayasigni cantposterior probabilityindicativeofoutliers.Theyarebothcut-markedand chronologicallycorrespondtothelimitbetweenCPandlaterCP. Theiroccurrencemayresultfromadecontaminationproblemfor theyoungerofthetwo,fromhumanorcarnivoreactivityduringthe laterCPphase,orsimplyfromexcavationerrors. ThedirectdateobtainedfortheSaint-Césaireskeleton( Fig. S6 )canbeacceptedwithsomecaution,becauseitscollagenyield is0.8%.However,theC:Nratioisfullywithintheaccepted range.Itsageofbetween41,950and40,660calBPwithaprobabilityof86%(1 )correspondstothetransitionfromCPtolater CPattheGrotteduRenneandisconsistentwiththeCPassignmentofthespecimen. Table1.RadiocarbondatesoftheGrotteduRennelayersandSaint-CésaireNeandertal Sampleno. Samples(withspatialand stratigraphicorigin)%Coll. 13 C 15 N%C%NC:N 14 CageB.P.+/ 1 Err GrotteduRenne Proto-Aurignacian EVA-79Arcy58RVIIC7b3.7 20.66.227.6103.229,930208 EVA-81*Arcy58VIIC9(178)4.4 20.58.422.88.13.333,850311 EVA-92Arcy58RVIIbC10(348)5.3 20.67.624.28.73.331,610131 EVA-93Arcy58RVIIC11(452)4.6 21.33.428.610.73.133,010182 EVA-95Arcy58RVIID10(254)6.2 20.76.328.410.63.134,810210 LaterCP EVA-52ArcyVIIIB15(24)1.8 205.544.514.43.635,980432 EVA-53ArcyVIII59.C15(38)0.9 19.13.838.914.23.236,230435 EVA-54ArcyVIII59.Y10(50)2.2 18.53.435.413.13.235,380390 EVA-55*ArcyVIII59.C15(36)1.2 19.33.241153.236,630452 EVA-56*ArcyVIII58.C8(92)1.4 20.4737.413.83.237,710533 CP EVA-44ArcyIXaB15(20)1.6 20.76.135.2133.239,280351 EVA-46ArcyIXa58.B112.7 215.64817.63.239,930361 EVA-47ArcyIXaB14(106)4 21.34.739.414.63.239,750360 EVA-33*ArcyIXbY13(43)4.4 18.73.639.114.53.140,970424 EVA-34*ArcyIXbB13(110)4.5 194.639.314.53.240,520389 EVA-35ArcyIXbB12(134)3.4 18.63.139.514.53.239,240341 EVA-36ArcyIXbC12(70)3.5 19.33.345.616.73.237,740307 EVA-37ArcyIXbB12(21)4.3 18.53.74215.43.239,450340 EVA-38ArcyXaA11(3412)1.5 20.85.738.614.33.236,540248 EVA-40ArcyXaA12(42)3.4 19.14.139.114.53.137,510275 EVA-41*ArcyXaA12(75)2.5 19.17.841.115.13.238,730333 EVA-42*ArcyXaB12(36)4 19.23.550.218.53.238,070311 EVA-43ArcyXa59.C9(68)4.1 18.93.137.813.93.239,020352 EVA-23ArcyXb161.C7(464)2.6 21.15.942.515.83.136,840335 EVA-24ArcyXb161.D11(60)5.2 19.18.940.2153.138,400317 EVA-25ArcyXb161.D11(87)2.2 19.63.641.315.23.236,210250 EVA-26*ArcyXb161.Z11(180)4 18.73.641.315.23.239,390334 EVA-27ArcyXb161.D11(93)4.4 18.73.541.915.43.240,230395 EVA-28ArcyXb161.C12(274)4.5 18.72.539.214.53.240,930393 EVA-29*ArcyXb261.Z11(213)1.7 20.95.640.414.83.235,500216 EVA-30*ArcyXb261.B9(327)2.2 215.340.314.93.237,980284 EVA-31ArcyXb261.A11(2641)2.5 19.85.540.114.73.239,290334 EVA-32ArcyXb261.C9(336)5.1 215.140.6153.236,820257 EVA-48ArcyXb262.C9(102)3.5 18.6343.215.43.339,070332 EVA-49ArcyXb262.C9(103)4.4 18.93.941.915.53.140,830778 EVA-51ArcyXb262.Z10(2)3.8 18.67.739.814.63.239,960702 Mousterian EVA-77*Arcy56RXIA126.2 206.929.610.93.242,120805 EVA-83*Arcy59RXIB12(25)5.9 20.49.129.610.93.241,980821 EVA-84Arcy60RXIC15(133)6.4 20.39.325.79.53.243,270929 EVA-85*Arcy60RXIC15(428)5.1 20.78.825.89.53.240,900719 Saint-Césaire OxA-18099SP28;Neanderthaltibia0.77 19.311.727.39.63.336,200750 C:Nratios,%C,%N,andamountofcollagenextracted(%Coll)refertothe > 30-kDafraction. 13 Cvaluesarereportedrelativetothe vPDBstandardand 15 NvaluesarereportedrelativetotheAIRstandard. *Boneswithanthropogenicmodi cations. Hublinetal. PNAS | November13,2012 | vol.109 | no.46 | 18745 ANTHROPOLOGY

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Discussion AttheGrotteduRenne,ourradiocarbonresultsarenotably consistentwiththestratigraphicdivisions.Mostimportantly,of 31samplesdocumentingtheCPandlaterCPdepositsnota singleoneyieldedadateintherangesobservedfortheoverlying ProtoaurignacianorunderlyingMousterianlayers.Considering Fig.1. CalibratedagesandboundariescalculatedbyusingOxCal4.1(37)andIntCal09(36).TheGrotteduRenneagesareinblackandarecomparedwith theSaint-Césairehumanbonedateinred.Asteriskindicatesanthropogenicallymodi edbones.Theresultsarelinkedwiththe(NGRIP) 18Oclimaterecord. 18746 | www.pnas.org/cgi/doi/10.1073/pnas.1212924109 Hublinetal.

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thenumberofanalyzedsamplesandtheirspatialdistribution, itseemsthereforeunlikelythatmajorcontaminationoftheCP andlaterCPcouldhaveoccurredintheportionofthesitewith horizontallayers. Theseobservationsareatoddswiththepreviousradiometric measurementsonthesamesequence(22).Inthatstudy,nine outliersweredetectedinaseriesof31dates,andintheCPlayers alone,eightsamplesyieldeddatesoutsidetheknownchronologicalrangeoftheCP.Inourview,thishighdegreeofintralayer variationintheradiometricagesresultsnotfromlayeradmixture butratherfromthesampleselectioncriteria.Inparticular,the emphasisputondirectlydatinganthropogenicallymodi ed bones,includingbonetoolsandbodyornaments,mayattimes havebiasedthesamplingtowardpoorlypreservedbones.After aninitialscreeningofthe59collectedsamples,theprevious studydiscarded19samplesbecauseofinsuf cientcollagen quality(22)(nonewerediscardedinourstudy).Still,asawhole, theremainingsamplesdisplayedlowcollagenyields,insome casesbetween0.3and0.5%(Fig.2).Inthissituation,contaminationisahugeissue.Signi cantly,noneoftheCPpendants,and onlythreeCPawlscouldbedated,oneofthemwithayounger agethantheyoungestGravettiandocumentedinthesite. Consideringthecomparablesizesofsomeoftheartifacts datedinthepreviousstudy(22)andofoursamples,itseems improbablethatthedivergingresultsaretheconsequenceof aselectivemigrationthroughthestratigraphyofworkedbones andartifacts.Oursampleswithanthropogenicmodi cationsprovidedagesinperfectagreementwiththoseofbonefragments withoutmodi cations.Additionally,thepossibleselectivemigrationofbodyornamentsfromtheProtoaurignacianlayersinto thelowermostCPlayer(X),wheremostoftheornamentswere discovered,isinconsistentwiththeabsenceofanyundisputable ProtoaurignacianstoneartifactsofsimilarsizeintheseCPlayers. AnupwardselectivemigrationofNeandertalremainstoward theCPlayersisequallydif culttoconceive.AlthoughtheCP andlaterCPdepositsyielded29isolatedteethandonetemporal bonedisplayingclearNeandertalfeatures(7,8),theunderlying Mousterianlayer(XI)isarcheologicallypoor(theareadirectly undertheexcavatedCPdepositsyieldedonlythreehumanteeth fromanexcavationareaof30m 2 ).Furthermore,twoteeth,most likelyNeandertal(7),comefromthelaterCPdepositsimmediatelyunderlyingtheProtoaurignacian. Anotherissuethatmayexplainsomeofthemismatches previouslyobservedbetweenstratigraphyandradiometric resultsrelatestothedistinctionofstratigraphicunits.The discoveryofsomere tsbetweenCPlayersandsublayersled ustogrouptheCPintoonlytwochronostratigraphicunits. Accordingtoourresults,theseunitsrepresent3,300and500y ofdeposit,respectively.Consideringdocumentedhumanand animalactivitiesinthesiteduringthesetwoperiods(38), consideringthechallengingsituationofrecognizingoccupation oorsduringexcavationonlargesurfacesofacavedeposit,and consideringthelimitsofradiocarbonprecisioninthistime period,furtherre nementsinthechronostratigraphypresented hereareunlikely. ThedirectdateobtainedfortheSaint-Césaireskeletondoes notfalsifythenullhypothesisthattheNeandertalsfoundinthe CPdepositsoftheGrotteduRennewereactuallythemakersof thisassemblage.OtherCPsiteshaveyieldedarangeofdatesof whichmostpredatetheuseofAMSandadvancedpretreatment proceduresinradiocarbondatingandarethereforehighly problematic(39).Todate,theGrotteduRennesequenceisthe onlyonedocumentedbyalargenumberofAMSradiocarbon datesobtainedonultra lteredsamples.Thesedatesarequite similartotheCPdatesrecentlyobtainedwiththesametechniqueatLesCottés(France),alsointhenorthernpartoftheCP extent(40). TheProtoaurignacianoftheGrotteduRenneandLesCottés ismuchyoungerthattheoldestoccurrenceofthisassemblagein northernItaly(41)andsouthernFrance(42),suggestingalater arrivaloftheirlikelymodernmakersincentralFrance.Thetime rangeoftheProtoaurignacianisalsowiderthanfoundforthis periodinthesouthernsitesandencompassestheEarlyAurignacianradiocarbondatesfromLesCottés.Ifcontaminationis notanissuethenlayerVIImightbeapalimpsestofProto-and EarlyAurignacian.TheexactageoftheEarlyAurignacianin centralandnorthernFranceremains,todate,poorlyknown. However,thelithicassemblageoflayerVIIprimarilydisplays aProtoaurignacianpattern(29). Finally,accordingtoourresults,theCPNeandertalsofthe GrotteduRenne,Saint-Césaire,andLesCottésclearlypostdate theearliestlikelymodernhumansremainsdocumentedin westernEurope(43)andlargelyoverlapintimewiththeearly AurignacianintheSwabianarea(44)andinsouthwestern France(42).Thisevidenceisfullycompatiblewithamodelof stimulusdiffusion(45),accountingfortheemergenceofbehavioralnoveltiesamongtheCPNeandertals(8,10). Conclusion Onradiometricgrounds,we ndnoevidencetosupportthe previousargumentsformajormovementsofmaterialoccurring betweenarcheologicallayersattheGrotteduRenneinthe sectionofthesitebetweenthecliffandtalus,theareawhich yieldedthebulkofthearchaeologicalmaterial.Thisresult con rmspreviousstudiesonthespatialdistributionofthepaleontological(46)andarcheologicalmaterial(11). Asawhole,theCPandlaterCPlayersofthissiterepresent arathershorttimeperiod.Thisrapiddeposition,thedif cultyin separatingsublayersatthetimeoftheexcavation,somelimited movementofmaterialbetweenCPsublayers,andthelimited resolutionofradiocarbondatesmake nerchronological distinctionsdif cult. Althoughmajorprogresshasbeenmadeinthereliabilityand calibrationof 14 Cdatesbeyond40,000CalBP(47),acomparison ofourresultsandthosepreviouslyobtainedforthissitealso suggeststhatbonepreservationremainsacrucialissuewhen radiocarbondatingisappliedtothistypeofmaterial.Thisissue representsamajortechnologicalchallengeforthedatingof smallartifactsandveryfragmentaryhumanremains. Fig.2. Distributionofthecollagenyieldsinthesamplesusedinthecurrent study(inblack,EVAsamples)andinpreviousradiometricmeasurementsof thesamesequence(22)(inred,OxAsamples). Hublinetal. PNAS | November13,2012 | vol.109 | no.46 | 18747 ANTHROPOLOGY

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