The Kocabaş hominin (Denizli Basin, Turkey) at the crossroads of Eurasia: New insights from morphometric and cladistic analyses

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C.R.Palevol17(2018)17–32

ContentslistsavailableatScienceDirect

Comptes

Rendus

Palevol

www . s c ie n c e d i r e c t . c o m

Human

Palaeontology

and

Prehistory

The

Kocabas¸

hominin

(Denizli

Basin,

Turkey)

at

the

crossroads

of

Eurasia:

New

insights

from

morphometric

and

cladistic

analyses

L’homininé

de

Kocabas¸

(bassin

de

Denizli,

Turquie)

au

carrefour

de

l’Eurasie

:

nouvelles

données

obtenues

à

partir

d’analyses

morphométriques

et

cladistiques

Amélie

Vialet

a

_,

_Sandrine

_Prat

b

_,

_Patricia

_Wils

c

_,

_Mehmet

_Cihat

_Alc¸

_ic¸

_ek

d

a_UMR_7194,_CNRS,_Muséum_national_d’histoire_naturelle,_UPVD_CERP_de_Tautavel,_1,_rue_{René-Panhard,}₇₅₀₁₃_Paris,_France b_UMR_7194,_CNRS,_Muséum_national_d’Histoire_naturelle,_UPVD,_Musée_de_l’Homme,_17,_place_du_Trocadéro,₇₅₁₁₆_Paris,_France c_UMS_2700,_outils_et_méthodes_de_la_{systématique}_{intégrative,}_Muséum_national_d’Histoire_naturelle,_43,_rue_Buffon,₇₅₀₀₅_Paris,_France d_Department_of_Geology,_Pamukkale_University,₂₀₀₇₀_Denizli,_Turkey

a

r

t

i

c

l

e

i

n

f

o

Articlehistory: Received4June2017

Acceptedafterrevision21November2017 HandledbyYvesCoppens

Keywords: Hominin Turkey Morphometry Cladistics OutofAfrica

a

b

s

t

r

a

c

t

TheKocabas¸skullcap(DenizliBasin),datedbetween1.2and1.6Ma,istheonlyancient homininfossilfromTurkeyandispartofdiscussionsfocusingonthefirstsettlementoutside theAfricancontinent.OurmorphometricstudytendstolinkthisspecimenwiththeAfrican fossils,HomoergasterandearlyHomoerectus,andtodistinguishitfromthespecimensfrom DmanisiandAsianHomoerectus.Theseresultsareconfirmedbyacladisticanalysis,which showsaseparationofKocabas¸fromtheEurasiancladecomprisingtheDmanisihominins andgroupingitwiththeAfricanfossilsdatedtoaround1Ma(KNM-OL45500,Daka-Bouri BouVP2/66,BuiaUA31).AsintheKocabas¸fossil,thedivergenceofthefrontalboneisnot verymarkedontheselatterfossilsandthetemporallinesareseparatedontheparietal bone.TheKocabas¸skullseemstopointtoadifferentevolutionaryhistorythanthatofthe Dmanisifossils,andcouldreflectalater“out-of-Africa”expansion.

Motsclés: Homininé Turquie Morphométrie Cladistique

Expansionhorsd’Afrique

r

é

s

u

m

é

Seulfossiled’homininéancienenTurquie,lacalottecrâniennedeKocabas¸(bassinde Deni-zli),datéeentre1,2et1,6Ma,s’inscritdansladiscussionsurlespremierspeuplements endehorsducontinentafricain.Notreétudemorphométriquetendàlerapprocherdes fossilesafricains,HomoergasteretHomoerectusrécents,etàledistinguerdesspécimens deDmanissietdesHomoerectusasiatiques.Cesrésultatssontconﬁrmésparuneanalyse cladistiquequimontreuneséparationdufossiledeKocabas¸ducladeeurasiatiquequiinclut leshomininésdeDmanissi,etleregroupeaveclesfossilesafricainsdatésautourd’1Ma (KNM-OL45500,Daka-BouriBouVP2/66,BuiaUA31).CesdernierspartagentavecKocabas¸ unedivergencedel’osfrontalpeumarquéeetuneséparationdeslignestemporalessurl’os pariétal.LecrânedeKocabas¸sembletémoignerd’unehistoireévolutivedifférentedecelle desfossilesdeDmanissi,quipourraitcorrespondreàuneexpansionhorsd’Afriqueplus tardive.

E-mailaddress:amelie.vialet@mnhn.fr(A.Vialet).

https://doi.org/10.1016/j.crpv.2017.11.003

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18 A.Vialetetal./C.R.Palevol17(2018)17–32

1. Introduction

Severalhominin fossils are known from Africa after 7Ma, when thefirst earlyHomois recorded at2.8 Ma (Villmoareetal.,2015).Thereisasignificanttimelagof about1MabetweenAfricaandEurasia,wherethefirst humanfossilsaredatedto1.77MaatthesiteofDmanisiin Georgia.Thislatteristheearliestknowndirectevidence ofthe“out-of-Africa”expansion.Based onlithic assem-blagesdiscoveredinAsia,thesefirstwavesofsettlement couldbeconsiderablyolderthan 2Ma (Prat, 2018,this issue).However,thesediscoveriesremainscarce. Inthis context,theKocabas¸skull,fromtheDenizliBasininTurkey, providesimportantevidenceofearlyhumanpresencein theMiddleEast,oran“out-of-Africa”expansionalongthe Levantinecorridor,andpossibledispersaltowardEurope – via the Bosphorus Strait – and toward Asia – across mountainousterritoriessuchastheTaurus,Zagros,and theCaucasus.AlthoughtheKocabas¸skullcapisvery frag-mented,itfillsapaleoanthropologicalgapbetween1.6and 1.2Ma,notonlyinAfricabutalsoinEurasia.Indeed,there isnofossilbetweenOH9fromOlduvai in Tanzaniaand theone-million-year-oldHomoerectus-likehomininsfrom EastAfrica(KNM-OL45500,Daka-BouriBouVP2/66,Buia UA31).In Europe,apartfromthedeciduoustoothfrom theOrceBasindatedto1.4Ma(Toro-Moyanoetal.,2013), theoldesthumanfossilsarefromtheAtapuerca–Simadel Elefantesitedatedto1.2Ma(Carbonelletal.,2008).

Thegoalof thispaper is,ononehand, to character-izemorphometricallytheKocabas¸fossiland,ontheother hand,totestitslinkwithotherPleistocenehomininsfrom AfricaandAsiausingcladisticanalysis.Themainquestion istoestablishwhethertheTurkishhominin,foundatthe crossroadsofAfricaandAsiabetween1.6and1.2Ma,is indeedclosertoAfricanorAsianspecimens.

2. Background

TheKocabas¸fossilwasdiscoveredin2002byoneofus (M.C.A.)inoneofthequarriesintheDenizliBasin(Alçiçek andAlçiçek,2014).Thesmallskullwasslicedbytheblades usedtocutblocksinthistravertinequarry.Althoughthe discoveryoftheskullwasaccidental,associatedfaunais abundantinthefindhorizon(Alçiçek,2014,Boulbesetal., 2014).ThisfossiliferoushorizonfromwhichtheKocabas¸ fossilcomes from, is identified as theupper travertine (UT)(Lebatardetal.,2014a,2014b),whichwastheonly quarriedtravertineatthetimeofdiscoveryin2002,and theonlyunittohaveyieldedpaleontologicalmaterialup untilnow.

ThestratigraphicandchronologicalcontextoftheUT bearing the homininskull and an abundantUpper Vil-lafranchianfaunawerecomprehensively definedduring thefield missionsin 2011and2012.Thisfieldworkhas beenfocusedonthedepositsfromtheFaberQuarrywhere the succession reaches a depth of over 90m and con-tainspreserved levels comparable tothose from which thefossilswereextractedin2002withintheUT.ThisUT unit is situated stratigraphically betweentwo fluviatile levels. The resultsobtained from 26_Al/10_Be _cosmogenic

nuclidesanalysisontheoverlyingandunderlyingﬂuviatile

conglomeratelevels(Lebatardetal.,2014a,2014b) indi-cate an ageof 1.1 Ma and 1.6 Ma respectively, setting achronologicalbracketbetweenthesetwodatesforthe Kocabas¸faunaandthehumanfossil.The magnetostratig-raphyforthewholesequenceofthedeposits(Khatibetal., 2014)allowsacorrelationofthebaseoftheupperfluvial unit (normal polarity) with the Cobb Mountain excur-sion, dated to1.22Ma.Moreover,the biochronologyof thefaunaArchidiskodonmerionalismeridionalis (Elephanti-dae)andPalaeotragus(Giraffidae)associatedwiththeskull (Boulbesetal.,2014)confirms thischronological frame-workbetween1.2and1.6Ma.

Nolithicartefactsarestrictlyassociatedwiththehuman andanimalfossilsfromtheUToftheDenizliBasin. Archae-ologicalprospectioninthesurroundingterraceshasjust startedrecently(Maddyetal.,2015;Aytekcomm.pers.), leadingtothediscoveryoflithicartefacts,mostlyonthe surface.Moregenerally,fewLowerPalaeolithicsitesare knownin Turkey(Dinçer,2016)andnoneareas oldas theKocabas¸locality.Wehavenoevidenceforthemoment to ascertain whether at 1.2/1.6 Ma, the technological modecouldberelatedtoOldowanorAcheulean techno-complexes.However,theDursunlusite(Güleçetal.,2009) andYarimburgazCave(Howelletal.,1996)haveyielded stoneflakes.Theopen-airsiteofKaletepeDeresi3attested tothepresenceofwell-shapedhandaxesmadeofobsidian (Slimaketal.,2004,2008)datedtothemiddlePleistocene (Tryonetal.,2009).Somehandaxeswerediscoveredinthe lowerlevelsoftheKarainECave(Tas¸kıran,2008,Tas¸kıran, 2018thisissue).Atthissite,theMousterianindustryiswell representedintheupperlevelsinassociationwith20 frag-mentaryNeandertal-likehumanremains(Chevalieretal., 2015)olderthan125ka(Otteetal.,1998).

3. Anthropologicalsettings

The Kocabas¸ skull was attributed to Homo erectus (Kappelmanetal.,2008).Itiscomposedof3fragments sep-aratedatthecranialsutures,whichwerenotcompletely fusedduetotheyoungageofthespecimen(Fig.1).The three cranial fragments werescanned using thePhilips helicalscanneratthePamukkaleteachinghospital,in Deni-zli,onSeptember14th,2009;slicethicknesswas0.80mm, thespacebetweensliceswas0.4mm(ﬁeldofview:20cm, matrix:512×512,power:175mA,intensity:120kV).

Afirstvirtualreconstructionwascarriedoutbasedon theCTdata(Vialetetal.,2011,2012).Thispaperconcerns thesecondvirtualreconstructionwithGeomagicStudio12 softwarebyA.V.andP.W.(technicalplatformASIM,atthe MNHN).Inthisreconstruction,called“Kocabas¸2”,theright partofthefrontalboneismoreaccuratelysituated.Indeed, thereisagapofafewmillimetersbetweentheparietaland frontalsutureswhichpreventsthedirectanatomical con-nectionbetweenthetwobones,aswaspreviouslythought andconsequentlyappliedtothefirstreconstruction.This reconstructionproceededasfollows.First,thetwo pari-etalfragments(rightandleft)wererearticulatedfollowing thecongruenceoftheirsagittalsuture.Suchrearticulation couldbedoneusingdifferentangles.Toensurethatthe rearticulatedboneswerenotoutoftheinitialvolume,we confirmedthattheyaresituatedbetweenthetworeference

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A.Vialetetal./C.R.Palevol17(2018)17–32 19

Fig.1.SuperiorviewofthethreeisolatedfragmentscomposingtheKocabas¸fossil.a:therightpartofthefrontalbone;b:theleftfronto-parietalpart;c: therightparietalfragmentarybone.

Fig.1. VuesupérieuredestroisfragmentsosseuxisolésquicomposentlefossiledeKocabas¸.a:lapartiedroitedel’osfrontal;b:lapartiefronto-pariétale gauche;c:l’ospariétalfragmentairedroit.

Table1

Comparison ofthe measurementsof thetwo reconstitutionsof the Kocabas¸ skull.

Tableau1

ComparaisondesmesuresdesdeuxreconstitutionsducrânedeKocabas¸.

Reconstitution M10 M9 M43

Kocabas¸1

FromVialetetal.,2012

102 85 116

Kocabas¸2

FromVialetetal.,2014

106 88 118

planes(i.e.thecuttingsectionsmadebythebladesofthe machineshownbytheexposureofdiploictissueatthe cen-tralpartofthefrontalandposteriorpartoftheparietals). Then,usingthemirror-imagingtechnique,weduplicated therightpartofthefrontalbonetoreconstructtheleftone. Finally,theanteriorblock(madebythecompletedfrontal) wasconnectedtotheposteriorblock(madebythe reartic-ulatedparietals)usingasguidelinesthesagittalsutureand thefrontalcrest(Fig.2).

Toconclude,thetworeconstructionsaredifferent.This isthereasonwhythemeasurementstakenforeachoneare notexactlythesame,resultinginaslightlylargerskullfor theKocabas¸2reconstruction(Table1).

Basedonseveralmorphologicalchangesofvarioussize observedontheendocranial surfaceof therightfrontal fragment,Kappelmanandco-workers(Kappelmanetal., 2008)arguedforthepresenceofaLeptomeningitis tubercu-losa(tuberculousmeningitis).However,thevalidityofthis diagnosiswasrefutedandcriticalcommentsweremadeby Robertsandco-workers(Robertsetal.,2009).

Complementarystudiesemployingnew macromorpho-logicalandmicro-CTanalysisontheendocraniallesions areongoinginordertoclarifytheirmorphologicalaspect, especiallybyfocusingonsofttissueimprintsandtheir pos-sibleinnerextension.

4. Material

Castsofthemainfossilhomininskullsfromthelower toupperPleistoceneinAfrica,Asia,andEuropewere con-sidered in this study (Table 2).They are stored at the NationalMuseumofNaturalHistoryinParis.28skullsin totalwereincludedinthe2Dmorphometricanalysisand amongthem,20wereusedforcladisticanalysis(including Sts5astheoutgroupspecimen).

5. Methods

5.1. 2Dmorphometry

Standard metrical measurements from Martin and Saller(1957)wereusedtocharacterizethedevelopment ofthefrontalbone(Table3).Principalcomponentanalysis wasgeneratedbyPast3.1.software(Hammeretal.,2001). 5.2. Cladisticanalysis

To investigate the phylogenetic relationships of the Kocabas¸hominin,wecarriedoutacladisticanalysisusing thecharactersdeﬁnedinTable4.Quotationwasdoneon theoriginalsand castsof thefossilskullsand fromthe

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20 A. Vialet et al. / C. R. Palevol 17 (2018) 17–32

Fig.2.ThevirtualreconstructionofKocabas¸2.Fromlefttoright:the3stepsoftheprocess:1,puttingtogetherthetwoparietals;2,mirroringtherightpartofthefrontaltoreconstructtheoverallbone;3, connectingthefrontalandparietalparts;4,toobtainthemostcompletereconstruction(Kocabas¸2).

Fig.2. LareconstitutionvirtuelledeKocabas¸2.Degaucheàdroite:lestroisétapesduprocessus.1,miseenconnexiondesdeuxpariétaux;2,reconstitutiondel’ensembledufrontal,parimagemiroirdela partiedroite;3,miseenconnexiondespartiesfrontaleetpariétale;4,pourobtenirlareconstitutionlapluscomplète(Kocabas¸2).

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Table2

Listofthehomininfossilsusedinthisstudy.

Tableau2

Listedeshomininésfossilesutilisésdanscetteétude.

Provenience N Name 2D Clad Dating Reference

Africa

SouthAfrica 2 Sterfontein

Sts5 X 2.8–2.4 Schwartzetal.,1994

2.16–2.05 HerriesandShaw,2011

Stw53 X 2.6–2Ma KumanandClarke,2000

Kenya 6 KoobiFora

KNM-ER1470 X X 2.06Ma Joordensetal.,2013

KNM-ER1813 X X 1.78Ma Feibeletal.,2009

KNM-ER3733 X X 1.65Ma McDougalletal.,2012

KNM-ER3883 X X 1.49–1.65Ma Feibeletal.,1989

KNM-ER42700 X 1.55Ma Spooretal.,2007

KNM-OL45500 X 0.97–0.90Ma Pottsetal.,2004

Tanzania 2 Olduvai

OH24 X 1.88Ma Hay,1976

OH9 X X 1.5–1.4Ma SchwartzandTattersall,2003

Ethiopia 1 Daka-Bouri

BouVP2/66 X X 1Ma Asfawetal.,2002

Eritrea 1 Buia

UA31 X 1Ma Abbateetal.,1998

Zambia 1 Kabwe X >125Ka SchwartzandTattersall,2003

Georgia 4 Dmanisi

D2280 X X 1.77Ma Lordkipanidzeetal.,2007

D2282 X

D2700 X X

D3444 X

Turkey 1 DenizliBasin

Asia Kocabas¸2 X X 1.6–1.2Ma Lebatardetal.,2014a,2014b

Indonesia 5 Sangiran

S17 X X 0.8Ma Sémahetal.,2010,Hyodoetal.,2011

Sambungmacan

Sm3 X 0.5Ka Yokoyamaetal.,2008

Ngandong

Ng5,10,12 X 150–27Ka Indriatietal.,2011

China 7 Yunxian

YunxianIIR X 936Ka LumleyandTianyuan,2008

Zhoukoudian

ZKD3,11,12 X X 780±80Ka Shenetal.,2009

Hulucave

Nankin1 X X 641±39Ka Bahainetal.,2017

Hexian 412±25Ka Grünetal.,1998

Dali 270Ka Xiaoetal.,2002

Europe

Greece 1 Petralona X 150–250Ka Grün,1996

France 3 Arago21-47 X 438±31Ka Falguèresetal.,2015

LCAS–LaChappelleauxsaints X MIS3-MIS4 Renduetal.,2014

LF1–LaFerrassie1 X 54±3to40±2Ka Guérinetal.,2015

Total 34 28 19

Table3

Conventionalmeasurementsusedinthisstudy.M9.1isonlyusedforthecladisticanalysis.

Tableau3

Mesuresconventionnellesutiliséesdanscetteétude.M9.1aétéutiliséuniquementdansl’analysecladistique.

Conventionalmeasurements fromMartinandSallers1957 Deﬁnitions

M9–Minimalfrontalwidth Betweenthetwofrontotemporals(ft) M9.1–Postorbitalbreadth Minimumpostorbitalbreadth

M10–Maximalfrontalwidth Onthecoronalsuture,betweenthetwocoronion(co) M43–Superiorfacialwidth Betweenthetwofronto-malar-temporals(fmt) FrontalbonelengthM29.2chordofthecerebralpartofthefrontalbone Fromsupra-glabellar(sg)tobregma(b)

literatureforKNM-ER42700(Spooretal.,2007),BuiaUA31 (Abbateetal.,1998,Bruneretal.,2016),Daka-BouriBouVP 2/66(Asfawetal.,2002,2008)andKNM-OL45500(Potts etal.,2004).

Eighteencharactersofthefrontalbonewereused;14 aremorphologicalcharactersand4aremetriccharacters. Asthereisnoconsensusonthetaxonomicclassiﬁcationof numeroushomininspecimens,wedeﬁnetheOperational

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Table4

Featuresusedforthecladisticanalysis.

Tableau4

Caractèresutiliséspourl’analysecladistique.

Feature number

Description References

1 Continuityofthepostorbitalgroove

0:absent,1:presentbutincomplete,2:presentcontinuously

Zeitoun,2000(feature4p54)

2 Shapeofthefrontalsuperiorborderinsuperiorview 0:straight,1:convex

3 Heightofthelateralpartofthesupra-orbitaltorus(c)incomparisonwithits centralpart(mid-orbitalposition)(b)

0:c>b,1:c<b,2:c=b

Zeitoun,2000(modiﬁedfromfeature 12p59)

4 Widthofthetemporalband(spacebetweentemporallines)onthefrontal bone

0:narrow,1:large

5 Projectionofthetemporalband(spacebetweentemporallines)onthefrontal bone

0:ﬂat,1:prominentandstronglyprominent

6 Bregmaticeminenceoverlappedontheparietalbone 0:absent,1:present

7 Supratrigonaldepression(ontheantero-medialborderofthetemporalline) 0:absent,1:present

8 Heightofthelateralpartofthesupra-orbitaltorus(metric) 0:weak,1:developed,2:strong

Prat,2000,2004,2005(modiﬁedfrom feature6p165)

9 Developmentofthezygomaticprocessofthefrontalbone(whichformsthe lateralborderoftheorbit)

0:short,1:long

Featureexclusivetothisstudy

10 Proportionofthefrontalscale(M29.2/M9.1)(metric) 0:short,1:long,2:verylong

11 Postorbitalconstriction(M9/M43)(metric) 0:strong,1:developed,2:lessdeveloped

12 Frontalbonedivergence(M9.1/M10)(metric) 0:verydivergent,1:divergent,2:lessdivergent

13 Convexityoftheexternallateralwallofthefrontalbonebelowthetemporal lines

0:vertical,1:convex

14 Sagittalkeelontheanteriorhalfofthesagittalsuture 0:absent,1:present

15 Postcoronaldepression 0:absent,1:present

16 DiscontinuityofthetemporallinesatStephanion 0:absent,1:present

17 Temporallines(superiorandinferior)separatedontheparietalbone 0:separated,1:fused

18 Shapeoftheorbitalroof

0:globular(shortroof),1:conical(long)

TaxonomicUnit(OTU)bythespecimenorgroupof spec-imens rather than by the species (as often used). This approach has been used by Caparros (1997); Zeitoun (2000);Gilbertetal.(2003);Prat(2004,2005);Cameron etal.(2004);MounierandCaparros(2015);Mounieretal. (2016)and Zeitounet al.(2016). Characterpolarity has beendeterminedbyrootingtheoutgroup.The polymor-phismiscodedasmultiplestates(0&1,0&2,1&2,0&1&2) with the polymorphism option of the Paup 4.01 soft-ware(Swofford,1998).Thequantitativecharacterswere codedusingthemethodproposedbyThiele(1993)(aftera logarithmictransformationofdata,theselatterbeing stan-dardizedusingtheformulaxs=((x–min)/(max–min))×n;

n=maximumnumberoforderedstatesallowablebythe algorithmused(32for PAUP).Therehasbeen consider-able debateconcerning methods of coding quantitative characters.ThemethodofThiele(1993)wasusedbecause it allows thecoding of allcharacters in a similar man-ner.Thedatawerecomputedina non-arbitrarywayto avoidanypreconceivedphylogenetichypothesis.Mostof

these characters have been taken from Zeitoun (2000)

and Prat (2000, 2004, 2005). Some of them have been deﬁnedinthisstudy(Table4).Thecharactershavebeen coded on ten OTU and the allocation of the character state for each one is given in Table 5. The OTU are as follows:Australopithecusafricanus(Sts5),whichisthe out-groupintheanalysis;earlyHomo(whichcomprisessome specimenscommonlyallocatedtoHomohabilisandHomo rudolfensis:OH24,KNM-ER1470;KNM-ER1813;Stw53); earlyAfricanHomoerectus/Homoergaster(KNM-ER3733, KNM-ER 3883; OH 9, KNM-ER 42700); Homo georgicus (D2280 andD2282);lateAfricanHomoerectus(BuiaUA 31, Daka-BouriBouVP2/66,KNM-OL45500); Kocabas¸2; IndonesianHomoerectus(Sangiran17);ChineseHomo erec-tus(ZKD3,11,12;Nankin1).Excepttheindividualsfrom this lattergroup,Asian andEuropeanmiddle andupper pleistocenespecimenswerenotconsidered,asour anal-ysis is more focused on the relationshipsbetween the Kocabas¸ fossil and lower and early middle Pleistocene hominins.

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Table5

CharacterstatescodingbyOperationalTaxonomicUnit.

Tableau5

Codagedesétatsdecaractèresparunitétaxinomiqueopérationnelle(OTU).

Character numbers Kocabas¸ D2700,D2280 OH24, KNM-ER1813, KNM-ER1470, Stw53 KNM-ER37333, KNM-ER3883,OH9, KNM-ER42700

BuiaUA31,Daka BouriBOUVP2/66, KNM-OL45500

Zhoukoudian3,11, 12,Nankin

S17 Sts5

1 2 1 0&2 0&1&2 1&2 2 0 0

2 1 0&1 0&1 1 0&1 0&1 1 0

3 0 0&1 0&1 0&1&2 1 0&1&2 0 0

4 0 0 0&1 0&1 1 0 0 0

5 1 0 0&1 0&1 1 0 0 0

6 0 1 0 0 ? 1 1 0

7 1 0 0&1 0&1 1 0&1 0 0

8 1 0 0 1 ? 1 2 0 9 0 0 0&1 0&1 0 1 0 0 10 1 2 2 1 0 2 1 0 11 2 1 1 1 1 2 2 1 12 2 1 1 1 2 1 2 0 13 1 1 0&1 1 0 1 0 1 14 0 1 0 0&1 0 1 ? 0 15 0 1 0 0 ? 1 1 0 16 0 0 0 1 1 0 0 0 17 0 0&1 1 1 0 0&1 1 1 18 0 1 0 1 1 1 1 ? Table6

Measurementsonthefrontalbonesof28fossilhominins.

Tableau6

Mesuressurlesosfrontauxde28homininésfossiles.

Locality No.specimen Measurements

M9 M10 M43 M29.2

Denizli–Turkey Kocabas¸2 88 106 118 80

Kenya–KNM ER1813 66 88 93 64

ER1470 70 90 106 78

ER3733 83 109 116 77

ER3883 81 108 115 87

Tanzania–Olduvai OH9 84 105 130 84

Georgia–Dmanisi D3444 70 94 105 75

D2280 74 106 114 86

D2282 67 91 105 78

D2700 67 90 97 71

Ethiopia–Daka-Bouri BouVP2/66 89 105 124 75

China–Yunxian YIIReconst. 102 136 130 74

China–Hulucave Nankin 83 101 110 80

China–Zhoukoudian ZKD3 81 105 109 90

ZKD11 84 106 111 90

ZKD12 91 110 119 97

Java–Sangiran Sang17 96 115 119 94

China Hexian 96 116 111 76

France–Arago Arago21Reconst. 109.1 113.1 123 92

Greece Petralona 108 120 130 92 Zambia Kabwe 98 117 133 105 China Dali 103 121 123 101 Java–Sambungmacan Sm3 99 110 112 93 Java–Ngandong Ng5 100 121 116 97 Ng10 102 121 121 99 Ng12 103 117 123 95

France–LaChapelle-aux-Saints LCAS 106 129 118 89

France–LaFerrassie LF1 108 126 120 92

Tobringtolightthehomoplasiesandsynapomorphies, we performed a parsimony analysis following the pro-tocol developed by MiguelCaparros in Caparros(1997)

andMounierandCaparros(2015).Aﬁrstlow-level anal-ysis is conducted with a branch and bound search in ordertoascertainthephylogeneticinformationcontentof

thecharacterbyitsretentionindex.Theretentionindex (RI)iscalculatedbysubtractingtheobservedtree-length fromthemaximumpossibletree-lengthandthen divid-ingthis value bythedifferencebetweenthemaximum andtheminimumlengths(Archie,1989;Farris,1989).All charactersareunorderedandallhaveequalweight.Ina

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24 A.Vialetetal./C.R.Palevol17(2018)17–32 secondstep,18characterswerereanalyzedafter

reweight-ingwiththerescaleconsistencyindex(RC=CI(consistency index)×RI(retentionindex)).Theconsistencyindex(CI)is calculatedastheminimumpossibletree-lengthdividedby theobservedtree-length(Farris,1989; KlugeandFarris, 1969).Ifthereisnohomoplasyinatree,thenitsobserved lengthequalstheminimumtree-lengthandtheCIequals one.Ifhomoplasyispresent,thentheCIislessthanone. WeusetheACCTRAN method(whichmaximizes rever-sals)inordertoresolvetheambiguitiesofthecharacter changesatthehypotheticalancestornodes,asitprovides morecharactersupporttoclades.

Themostparsimonioustreeswereobtainedusingthe heuristicsearchalgorithmwiththebranch-swapping algo-rithm(TBR).WeusePAUPversion4.01(Swofford,1998), andMacClade(v.4.06)inordertoillustratethesupporting charactersatthehypotheticalancestor’snodes.

6. Results

6.1. Anthropologicalstudy–2Dmorphometry MetricsarepresentedinTable6.

ThePCAwascomputedbasedon4variables(M9,M10, M43, M29.2) measured on 28 fossils. On the ﬁrst axis (whichaccountsfor82.5%ofthetotalvariance), individ-ualsarespreadfollowingtheirglobalsize(Fig.3).Kocabas¸ 2isamongtheHomoerectuss.l.fromAfricaandAsia.The DmanisifossilsaresmallerandtheHomoheidelbergensis, lateHomoerectusandNeandertalsarebigger.

Onthesecond axis(11.3%ofthe totalvariance), the lengthof thefrontalboneseparates thefossils. Kocabas¸ 2isdistinctnot onlyfromtheZhoukoudianHomo erec-tusbutalsofromSangiran17,OH9andKNM-ER3883.It isclosertoAfricanfossilssuchasKNM-ER3733and Daka-BouriBouVP2/66(Fig.3)whichshowarelativelyshorter frontalscale.

Thedistributionofthefossilsacrossthethirdaxis(4% ofthetotalvariance)islinkedwiththewidthofthe supra-orbitaltorus(Fig.4).Again,Kocabas¸2isinanintermediate positionbetweentheAsian Homo erectus(Zhoukoudian withagracile supra-orbitaltorus) and theAfrican ones (OH9,Daka-BouriBouVP2/66withastrongsupra-orbital torus).Forthisfeature,theTurkishspecimenisclosertothe DmanisiandtheEastTurkana(KNM-ER3733andER3883) specimens.

Theweakdivergenceof thefrontalbone (proportion betweenminimal-M9andmaximal-M10frontalwidths)is expressedonthefourthaxis(2.1%),onwhichtheKocabas¸2 skullisfarfromtheAsianandAfricanHomoerectus,except OH9(Fig.5).

Theresultsofthisstudyallowus:

• tolinkKocabas¸2withearlyandlateAfricanHomoerectus; • todistinguishitfromtheearlyHomospecimensandfrom

thesmallGeorgianhominins;

• todistinguishitfromtheAsianHomoerectus.

Incomparisontothelatter,Kocabas¸2showsashorter andposteriorlynarrowerfrontalboneandamore promi-nentsupra-orbitaltorus.Withitssmallsize,theTurkish

skullisclearlydistinctfromthemiddleandupper Pleis-tocenehominins.Suchresultsconﬁrmthepreviousstudies based on anatomical features and 3D morphometrics (Vialetetal.,2012,2014).

6.2. Cladisticanalysis

Concerningthephylogeneticanalysisbasedon18 char-acters and 7 specimens or group of specimens in the ingroup,threetreeshavebeenobtainedontheﬁrststep of theanalysis.Thetreelengthis31, consistencyindex (CI)=0.7097; retentionindex (RI)=0.5714; and rescaled consistencyindex(RC)=0.4055.

Aheuristicsearchbasedon18frontalfeatures(Table7), entered with equal weights, identiﬁed 9 morphological charactersthatcontainphylogeneticinformation(RI≥0.5). Weidentify6truesynapomorphies(CI,RI,adRC=1,HI=0); characters #5 Projection of the temporal band onthe frontalbone;#6Bregmaticeminenceoverlappedonthe parietalbone;#7Supratrigonaldepression;#14Sagittal keelontheanteriorhalfofthesagittalsuture;#15 Post-coronaldepression;#17Temporallinesseparatedonthe parietalbone.

Forthesecondstep,weusedabranchandbound algo-rithmtoanalyzethereweightedcharactersbyrespective RCs.Threetreeshavebeenobtained.Thestrictconsensus tree isshown in Fig.6. Thetree lengthis13.66, reten-tionindex(RI)=0.8790;consistencyindex(CI)=0.9051and rescaledconsistencyindex(RC)=0.7956.

AtNodeA,thehypotheticalancestorofthegroup(Buia UA31,Daka-BouriBouVP2/66,OL45500andKocabas¸)is wellsupportedbyonesynapomorphy(#17:superiorand inferiortemporallineschangingfromfusedtoseparated ontheparietalbone,theydonotformastrongridge).This nodeisalsosupportedbyoneinformativehomoplasy(# 12)frontalbonechangingfromdivergenttolessdivergent. Kocabas¸exhibitsthefollowinginformativecharacters (RI≥0.5):

• #5–Projectionofthetemporalbandonthefrontalbone (RI=1): changes from ﬂat to prominent and strongly prominent,whichissharedwiththegroupBuiaUA31, Daka-BouriBouVP2/66,KNM-OL45500;

• # 7–Supratrigonal depression (RI=1): changes from absenttopresent,whichissharedwiththegroupBuia UA31,Daka-BouriBouVP2/66,KNM-OL45500;

• #8–Heightofthelateralpartofthesupra-orbitaltorus (RI=0.5): changes from weak to developed, which is shared with the group (KNM-ER 3733, 3883, 42700, OH9)andthegroup(BuiaUA31,Daka-BouriBouVP2/66, KNM-OL45500)andaconvergencewiththehypothetical ancestorofthegroup(Zhoukoudian3,11,12,Nankin1) andSangiran17;

• # 11–Postorbital constriction (RI=0.5): changes from developed to less developed which is a convergence withthehypotheticalancestorofZhoukoudian3,11,12, Nankin1and Sangiran17. Thedecreaseinpostorbital constrictioncanbeassociatedwithincreasedﬂexionof theanteriorcranialbase(Cameronetal.,2004); • # 12–Frontal bonedivergence(RI=0.5): changesfrom

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A. Vialet et al. / C. R. Palevol 17 (2018) 17–32 25

Fig.3. PCAon4measurementsonthefrontalbone(M9,M10,M43,M29.2)from28hominins.Axis1accountsfor82.5%ofthetotalvarianceandaxis2for11.3%ofthetotalvariance.Blacksquares:earlyHomo andAfricanHomoerectus,bluedots:AsianHomoerectus,purplecrosses:MiddlePleistoceneEuropeanhomininsandNeandertals,redstar:theKocabas¸hominin.

Fig.3. ACPsurquatremesuresdel’osfrontal(M9,M10,M43,M29,2)sur28homininés.L’axe1comptepour82,5%delavariancetotaleetl’axe2pour11,3%delavariancetotale.Carrésnoirs:premiersHomo etHomoerectusafricains;pointsbleus:Homoerectusasiatiques;croixviolettes:homininéseuropéensduPléistocènemoyenetNéandertaliens;étoilerouge:l’homininédeKocabas¸.

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26 A. Vialet et al. / C. R. Palevol 17 (2018) 17–32

Fig.4. PCAon4measurementsonthefrontalbone(M9,M10,M43,M29.2)from28fossils.Axis1accountsfor82.5%ofthetotalvarianceandaxis3for4%.Blacksquares:earlyHomoandAfricanHomoerectus, bluedots:AsianHomoerectus,purplecrosses:MiddlePleistoceneEuropeanhomininsandNeandertals,redstar:theKocabas¸hominin.

Fig.4. ACPsurquatremesuresdel’osfrontal(M9,M10,M43,M29,2)sur28homininés.L’axe1comptepour82,5%delavariancetotaleetl’axe3pour4%.Carrésnoirs:premiersHomoetHomoerectusafricains; pointsbleus:Homoerectusasiatiques;croixviolettes:homininéseuropéensduPléistocènemoyenetNéandertaliens,étoilerouge:l’homininédeKocabas¸.

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A. Vialet et al. / C. R. Palevol 17 (2018) 17–32 27

Fig.5. PCAon4frontalbonemeasurements(M9,M10,M43,M29.2)from28fossils.Axis1accountsfor82.5%ofthetotalvarianceandaxis4for2,1%ofthetotalvariance.Blacksquares:earlyHomoandAfrican Homoerectus,bluedots:AsianHomoerectus,purplecrosses:MiddlePleistoceneEuropeanhomininsandNeandertals,redstar:theKocabas¸hominin.

Fig.5. ACPsurquatremesuresdel’osfrontal(M9,M19,M43,M29,2)sur28homininés.L’axe1comptepour82,5%delavariancetotaleetl’axe4pour2,1%delavariancetotale.Carrésnoirs:premiersHomoet Homoerectusafricains;pointsbleus:Homoerectusasiatiques;croixviolettes:homininéseuropéensduPléistocènemoyenetNéandertaliens;étoilerouge:l’homininédeKocabas¸.

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Table7

Characterstatescoding.

Tableau7

Codagedesétatsdecaractères.

Morhologicalfeatures Range Minsteps Treesteps Maxsteps CI RI RC HI 1Continuityofthepostorbitalgroove 2 2 3 3 0.667 0.000 0.000 0.333 2 Shapeofthefrontalsuperiorborderinsuperiorview 1 1 1 1 1.000 0/0 0/0 0.000 3 Heightofthelateralpartofthesupra-orbitaltorus(c)in

comparisonwithitscentralpart(mid-orbitalposition)(b)

1 1 1 1 1.000 0/0 0/0 0.000

4Widthofthetemporalband(spacebetweentemporal lines)onthefrontalbone

1 1 1 1 1.000 0/0 0/0 0.000

5Projectionofthetemporalband(spacebetween temporallines)onthefrontalbone

1 1 1 2 1.000 1.000 1.000 0.000

6Bregmaticeminenceoverlappedontheparietalbone 1 1 1 3 1.000 1.000 1.000 0.000 7Supratrigonaldepression(ontheantero-medialborder

ofthetemporalline)

1 1 1 2 1.000 1.000 1.000 0.000

8Heightofthelateralpartofthesupra-orbitaltorus (metric)

2 2 3 4 0.667 0.500 0.333 0.333

9Developmentofthezygomaticprocessofthefrontal bone(whichformsthelateralborderoftheorbit)

1 1 1 1 1.000 0/0 0/0 0.000

10Proportionofthefrontalscale(M29.2/M9.1) 2 2 4 5 0.500 0.333 0.167 0.500 11Postorbitalconstriction(M9/M43) 1 1 2 3 0.500 0.500 0.250 0.500 12 Frontalbonedivergence(M9.1/M10) 2 2 3 4 0.667 0.500 0.333 0.333 13Convexityoftheexternallateralwallofthefrontal

bonebelowthetemporallines

1 1 2 2 0.500 0.000 0.000 0.500

14Sagittalkeelontheanteriorhalfofthesagittalsuture 1 1 1 2 1.000 1.000 1.000 0.000 15Postcoronaldepression 1 1 1 3 1.000 1.000 1.000 0.000 16 DiscontinuityofthetemporallinesatStephanion 1 1 2 2 0.500 0.000 0.000 0.500 17Temporallines(superiorandinferior)separatedonthe

parietalbone

1 1 1 2 1.000 1.000 1.000 0.000

18Shapeoftheorbitalroof 1 1 2 2 0.500 0.000 0.000 0.500

Fig.6.MostParsimoniousTree(MPT).OverallRetentionIndexRI=0.8790;andConsistencyIndexCI=0.9051.

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(BuiaUA31,Daka-BouriBouVP2/66,KNM-OL45500)and aconvergencewithSangiran17;

• # 17–Temporal lines separated on the parietal bone (RI=1):superiorandinferiortemporal linesseparated ontheparietalbone, theydo notformastrongridge, changingfromfusedtoseparated,whichissharedwith thegroup (BuiaUA 31, Daka-BouriBouVP2/66, KNM-OL45500).

Furthermore, theKocabas¸ specimenshows a conical orbitalroof(#18,CI=0,5;RI=0),whichisalsoobservedin thegroup(OH24,KNM-ER1813,1470,Stw53).

7. Discussion

Ourmultivariateanalysis,basedonthemain measure-mentsofthefrontalbone,placestheKocabas¸fossilinthe Homoerectuss.l. group(i.e.ZhoukoudianandNankin 1 fromChina,KNM-ER3733,KNM-ER3883fromEastAfrica andthelateAfricanHomoerectussuchasOH9and Daka-BouriBouVP2/66).Itsoverall sizeisgreaterthanthatof earlyHomoandtheDmanisispecimensandsmallerthan Middle and Upper Pleistocene hominins. However, the TurkishfossildiffersfromtheAsianHomoerectusbythe shortnessofitsfrontalscalewhichisclosetotheAfrican fossils, KNM-ER3733 and Daka-BouriBouVP2/66.In our opinion,thisdifference,dealingwiththeproportionofthe frontalboneanditselongation,isaverysigniﬁcant fea-turelinkedwithendocranialdevelopment.Ourconclusions differfromAytekandHarvati’s,whoarguedfora close-nessbetweentheKocabas¸fossiland,notonlytheChinese Zhoukoudian IIIspecimen, but also European hominins suchasCepranoandArago(AytekandHarvati,2016).It isimportanttorecall thattheiranalysisonlyfocuseson thesupra-orbitaltoruswhichisastronglydimorphicarea. However,itistruethattheKocabas¸andZhoukoudianIII specimensare morphologicallycloseforthisanatomical region and theyare not far from the European fossils, suchastheCepranoand Arago21 fossilswhich havea primitivesupra-orbitaltorus.Regardingthisarea,wemust mentionanotherdivergencefrompreviousstudies.Indeed, weconsiderthatKappelmanetal.(2008)over-estimated the supra-orbital torus breadth (124mm vs 118mm in thisstudy)andheight(min16–max19vsmax13inour own measurements),which ledtooveremphasisonthe closenessoftheTurkishfossilwiththeMiddlePleistocene hominins.However,wemustkeepinmindthatonlythe lateralhalfofthesupra-orbitaltorusontheKocabas¸fossilis preservedwhichmeansthatitisnotpossibletoknowhow developedthemedialpartwas.Forthewidthofthe supra-orbitaltorus,ourresultsshowanintermediatepositionfor theKocabas¸homininbetweenontheonehand,agracile groupcomposedoftheAsianHomoerectusandmostofthe DmanisiandearlyHomospecimens,andontheotherhand, themorerobustOH9andDaka-BouriBouVP2/66skulls.

Finally,Kocabas¸sharesaweakfrontaldivergencewith theone-million-year-oldHomoerectus-likehomininsfrom East Africa (OH9, Daka-Bouri BouVP2/66). This charac-ter distinguishesthem from earlier hominins including theDmanisifossils.Theseresultsarecorroboratedbythe cladisticanalysiswhichconsidersmorphologicalaswellas

metricalfeatures.Ouranalysisclearlyseparatestwoclades: anAsianone (includingtheDmanisispecimens)and an Africanone(includingtheKocabas¸fossil).Itenablesusto consideranevolutionaryhistoryfortheKocabas¸fossil, ﬁll-ingagapinthisfossilrecord,whichisdistinctfromthatof theDmanisigroupandlinkedwiththeAfricanHomoerectus knownbetween1.65Ma(KNM-ER3733),1.5Ma(OH9)and around1Ma(KNM-OL45,500,Daka-BouriBouVP2/66,Buia UA31).TheTurkishspecimenistheﬁrstrepresentativeof suchaclusteroutsideAfrica.Anothergroupisrepresented bytheAsianHomoerectus(ZhoukoudianandNankin1), whichareinasamecladewiththeDmanisifossils show-inganexpansiontowardtheEast,rootedat,atleast,1.77 Ma.However,alessmarkedpostorbitalconstriction(#11) isacommonfeaturebetweenKocabas¸andtheancestor oftheAsianHomoerectusgroupshowingthattheyhave reachedasimilardevelopmentalstage.Inthesameway, thefrontalbone divergence(#12) islesspronouncedin Sangiran17andthegroupincludingKocabas¸,BuiaUA31, Daka-BouriBouVP2/66,andKNM-OL45500whichisa con-vergenttraitduetoendocranialdevelopment.Butamong thesefossils,Kocabas¸showsaconvexityofthefrontalwall belowthetemporallines(#13)whichisverticalinthe oth-ers.Insummary,eventhoughtheAsianHomoerectus,the one-million-oldAfricanspecimensandtheTurkishfossil belongtothesamestageofcranialdevelopment,Kocabas¸ seemsdifferenttosomeextent.Moreover,inthisspecimen, theorbitalroof forms,attheback,a verticalwall(#18) whichsuggeststheorbitalcavitytobeshortand globu-lar,insteadoflongandconicalasobservedingeneralon fossilhomininsandHomosapiens.Thispattern,observed onKNM-ER1813andonStw53asfarasthisareais pre-served,maybelinkedwiththeoverallsmallsizeofthese individualsand/oritsanatomicalage.

An expansion of the Kocabas¸ group representatives towardthewest couldbeexpectedbased ontheir geo-graphical location. Key specimens to test this question arethefossilsfromtheAtapuerca-SimadelElefantesite datedto1.2Ma(Carbonelletal.,2008;Bermúdezde Cas-troetal.,2011)andOrcedatedto1.4Ma(Toro-Moyano etal.,2013).Unfortunately,theTurkish(frontalbone)and Spanishfossils(mandible,handphalanx,fragmentoffemur forSima delElefante, deciduoustoothforOrce)arenot strictly comparable in terms of skeletal representation. Moresuitablearethefrontalandtheparietalbones discov-eredintheAurorastratum(TD6.2)intheAtapuerca-Gran Dolinasite,datedto0.9Ma(Morenoetal.,2015).Indeed, ATD6-15consistsofalargeportionofthefrontalbone (bet-terpreservedonitsrightside)butshowingpost-mortem distortion (Arsuaga et al., 1999). Asfar as they can be compared,theTurkishandSpanishfossilsshowsome sim-ilarities.Onboth,wecanobservethesameconvexityof theorbitalroof(character#18)asaglobular/shortroof(vs conical/longone).Aswehaveseenpreviously,thisfeature appearsinterestingtohighlightthepeculiarityofKocabas¸ incomparisonwiththeotherspecimenseveniftheRIis low(<0.5).MaybethissimilaritybetweentheTurkishand Spanishhomininsisduetotheyoungageofboth individ-uals,astheformerisconsideredayoungadult(Vialetetal., 2012)andthelatterclosetopuberty(Arsuagaetal.,1999). Theparietal bone(ATD6-100/168) is alsofroma young

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30 A.Vialetetal./C.R.Palevol17(2018)17–32 individualbetween8and10yearsoldwhenHomo

sapi-ensisusedasthereference(Bruneretal.,2017).Thatisthe reasonwhythecomparisonislimitedduetoconsiderations ofgrowthpatternsand thestateofpreservation (i.e.in Kocabas¸,onlytheanteriorhalfoftheparietalispreserved).

8. Conclusion

Theresultsoftheanthropologicalanalysishighlightthat thereconstructedKocabas¸partialskull(Kocabas¸2)shows aconformationofitsfrontalbonewhichisdistinctfrom theearly Homopattern and intermediate betweenthat ofAsianHomoerectusandAfricanHomoerectus. Cladisti-callyspeaking,theTurkishfossilisclearlyseparatedfrom theDmanisifossilsandtheAsianHomoerectus,whichare groupedinthesameclade.ItislinkednotonlytotheAfrican Homoerectus bracketedbetween 1.49and 1.65Ma but alsowiththeone-million-year-oldAfricanhomininssuch astheKNM-OL45500,Daka-BouriBouVP2/66andBuiaUA 31skulls.Duetothelackofconsensustonamethelatter specimensand,generallyspeaking,todeﬁneHomo erec-tus,wemaintaintheattributionoftheKocabas¸hominin tothisspeciesinthebroadestsense,whilewecansuggest thattheTurkishfossilsharesanevolutionaryhistorymore withtheAfricangroupsthantheAsianones.Fillinga pale-oanthropologicalgapoutsideAfrica,between1.6and1.2 Ma,itmayprovideevidenceofhumandispersalthrough south-western Europe and the entire Mediterranean area.

Acknowledgements

TheauthorswanttowarmelythankPr.YvesCoppens forhisinvitationinthisspecialissueofC.R.Palevol dedi-catedtothe“Homininsandtools.ExpansionsfromAfrica towardsEurasia”.Mostofthecontributionsofthis issue havebeneﬁtedfromthesupportoftheAgence Universi-tairedelaFrancophonieinthecontextofthesessionB52 oftheXVIIeCongrèsdel’UnionInternationaledesSciences PréhistoriquesetProtohistoriques-UISPP (2014,Burgos, Spain).Thisresearchisanemanationofthe“FirstHomoin Turkey”,ascientiﬁcprojectledbyA.Vialetandsupported bytheCNRS(PICS2016-2018).WearegratefultoLouise ByrneandPhilGlaubermanforlinguisticassistance.MCA issupportedtotheGEB˙IPgrant(TheOutstandingYoung ScientistAward)givenbytheTurkishAcademyofSciences (TÜBA).

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