Late Neandertals in Southeastern Iberia: Sima de las Palomas del Cabezo Gordo, Murcia, Spain


previous item | next item

Citation

Material Information

Title:
Late Neandertals in Southeastern Iberia: Sima de las Palomas del Cabezo Gordo, Murcia, Spain
Series Title:
Proceedings of the National Academy of Sciences of the United States of America
Creator:
Walker, Michael J.
Gibert, Josep
López, Mariano V.
Lombardi, A. Vincent
Pérez-Pérez, Alejandro
Zapata, Josefina
Ortega, Jon
Higham, Thomas
Pike, Alistair
Schwenninger, Jean-Luc
Zilhão, João
Trinkaus, Erik
Publisher:
National Academy of Sciences
Publication Date:

Subjects

Subjects / Keywords:
Neanderthals ( lcsh )
Paleolithic period ( lcsh )
Shafts (Excavations) ( lcsh )
Genre:
serial ( sobekcm )
Location:
Europe -- Spain -- Murcia -- Torre-Pacheco

Notes

Abstract:
Middle Paleolithic fossil human remains from the Sima de las Palomas in southeastern Iberia (dated to ≤43,000–40,000 calendar years before present) present a suite of derived Neandertal and/or retained ancestral morphological features in the mandibular symphysis, mandibular ramus, dental occlusal morphology, and distal hand phalanx. These traits are combined with variation in the mandibular corpus, discrete dental morphology, tooth root lengths, and anterior dental size that indicate a frequency difference with earlier Iberian and more northern European Neandertals. The Palomas Neandertals therefore confirm the late presence of Neandertals associated with the Iberian persistence of the Middle Paleolithic, but suggest microevolutionary processes and/or population contact with contemporaneous modern humans to the north.
Original Version:
Volume 105, Issue 52
General Note:
6 p.

Record Information

Source Institution:
University of South Florida
Holding Location:
University of South Florida
Rights Management:
All applicable rights reserved by the source institution and holding location.
Resource Identifier:
K26-05545 ( USFLDC DOI )
k26-5545 ( USFLDC Handle )

USFLDC Membership

Aggregations:
University of South Florida
Karst Information Portal

Postcard Information

Format:
serial

Downloads

This item is only available as the following downloads:


Full Text

PAGE 1

LateNeandertalsinSoutheasternIberia:Simadelas PalomasdelCabezoGordo,Murcia,SpainMichaelJ.Walkera,JosepGibertb,MarianoV.Lo ´ peza,A.VincentLombardic,AlejandroPe ´ rez-Pe ´ rezd,JosenaZapataa, JonOrtegaa,ThomasHighame,AlistairPikef,Jean-LucSchwenningere,Joa ˜ oZilha ˜ of,andErikTrinkausg,1aA ´ readeAntropologíaFísica,DepartamentodeZoologíayAntropologíaFísica,FacultaddeBiología,CampusUniversitariodeEspinardo,Universidad de Murcia,30100Murcia,Spain;bInstitutPaleontolo ´gic‘‘Dr.M.Crusafont,’’Carrerdel’EscolaIndustrial23,08201Sabadell,Barcelona,Spain;c2584Blossom Lane,NewCastle,PA16105;dSeccio ´ndeAntropología,DepartamentodeBiologíaAnimal,FacultaddeBiología,UniversitatdeBarcelona,AvenidaDiagonal 645,08028Barcelona,Spain;eResearchLaboratoryforArchaeologyandtheHistoryofArt,UniversityofOxford,OxfordOX13QY,UnitedKingdom;fDepartmentofArchaeologyandAnthropology,UniversityofBristol,43WoodlandRoad,BristolBS81UU,UnitedKingdom;andgDepartmentof Anthropology,WashingtonUniversity,SaintLouisMO63130 ContributedbyErikTrinkaus,November7,2008(sentforreviewOctober17,2008)MiddlePaleolithicfossilhumanremainsfromtheSimadelasPalomas insoutheasternIberia(datedto < 43,000–40,000calendaryears beforepresent)presentasuiteofderivedNeandertaland/orretained ancestralmorphologicalfeaturesinthemandibularsymphysis,mandibularramus,dentalocclusalmorphology,anddistalhandphalanx. Thesetraitsarecombinedwithvariationinthemandibularcorpus, discretedentalmorphology,toothrootlengths,andanteriordental sizethatindicateafrequencydifferencewithearlierIberianandmore northernEuropeanNeandertals.ThePalomasNeandertalstherefore conrmthelatepresenceofNeandertalsassociatedwiththeIberian persistenceoftheMiddlePaleolithic,butsuggestmicroevolutionary processesand/orpopulationcontactwithcontemporaneousmodern humanstothenorth.dentition mandible MiddlePaleolithic postcraniaIthasbecomeapparentthatthetransitionfromaEurope populatedbyNeandertalstooneoccupiedbyearlymodern humansduringmarine(oxygen)isotopestage(MIS)3consisted ofawestwardspreadofmodernhumans,variablyabsorbingat leastsomelocalNeandertalpopulations(1,2).However,the detailedaspectsofthispopulationprocessremainobscureasa resultofthescarcityofwell-datedlateNeandertalsandearly modernhumans,despitetherecentdirectdatingofseveral specimensfrombothsamples(3–9).Ifwearetounderstandthe paleobiogeographyofthisprocess,andhencethepotential patternsofinteractionsbetweenthesetwomorphologically definedhumangroups,thenitisessentialtodocumentthe biologyofboththeearliestmodernhumansandthelatest Neandertals. IthasbeenrecognizedforsometimethattheMiddlePaleolithic,generallypresumedinEuropetohavebeentheproductof Neandertals,persistedsubstantiallylongerinIberiasouthofthe Pyrenees(southofthe‘‘EbroFrontier’’)thanelsewherein Europe,to 34,000( 34ka)calendaryearsbeforepresent(cal BP)( 30karadiocarbonyearsbeforepresent(14CBP)](10; supportinginformation(SI) Appendix ,Fig.1).Eventhoughthere areIberianNeandertalremainsthathavebeenreferredtothis age(11),theirpurportedlateagehasbeenplacedindoubt(12). Atpresent,themostrecent,securelydated,diagnosticNeandertalfossilfromtheregionistheOliveira1middlemanual phalanxat 43,500calBP( 39ka14CBP)(13),aboutthesame ageasthenorthernSpainElSidro ´nremains(7).Itisinthis contextthatwepresentaseriesofNeandertalremainsfromthe upperlevelsoftheSimadelaPalomasinsoutheasternSpain. ResultsContextofthePalomasHumanRemains.TheSimadelasPalomas (Rock-DoveHole)isakarsticshaft(37°47 59 N,0°53 45 W) inthePermo-Triassicmarbleofahill(CabezoGordo)inTorre Pachecotownship,Murcia,reaching312ma.s.l.,overlookingthe MarMenorcoastallagoonoftheMediterraneanSea( SIAppendix ,Figs.1and2).Themainshaftis18mdeep,opening beneathoverhangingrockat 123ma.s.l.( SIAppendix ,Figs.3 and4).Theshaft’sbrecciatedcontentswerelargelyemptiedby 19thcenturyminers,wholeftasedimentcolumndownoneside andscatteredfossiliferousrubbleonthehillside.Systematic collectionofdisturbedremainsandexcavationoftheupper brecciadepositswerebegunbyM.J.W.andthelateJ.Gibert afterdiscoveryofacrushedfacialskeleton(Palomas1)fromthe uppermostbrecciabyJ.C.Blancoin1991(14,15).Fossilhuman remainshavebeencollectedfromtheminerubble(1992–1999) andexcavatedinsitu(1994–present)( SIAppendix ,Table1). Anumberoftheinsituhumanremainscomefromabove, within,andslightlybelowafusiformlensofdark-graysediment (burnt,accordingtoX-raydiffractionandfluorescenceanalyses).Itattainedamaximumthicknessof 20cmintheangleof anL-shapedexcavationareaintheUpperCutting,coveringa thin,oblongmarbleslab( 40 30 10cm)betweenlevels2k and2m.Thelenspeteredout,bothneartheopenshaftand belowtheentrancewhereitcoveredpartofthefootofa downwardandinwardlysteeplysloping(30°–40°)e ´boulisor screeofmarbleblocks 50kg.Amoreconsistentandwidespreadlowerlayerofdark-graysedimentliesbeneathlevel2l, fadingawaybesidetheopenshaft(Fig.1; SIAppendix ,Figs.5 and6). Onlyhumanbonesandteethfromsedimentspostdatingthe e ´boulis(fromlevelsnodeeperthan2l)aredealtwithhere. Althoughtheselatersedimentscontainfewlargerocksandwere laiddownhorizontally,theyoftenhaveacoarsetextureimplying thattheywerewashedfromthehillsideintothecaveanddown thesteepinteriorscreeslope;theslopehaddevelopedneitheran erodedsurfacenoracalcretecrust,incontrasttothethincalcite depositonthehillsidethateventuallysealedoffthee ´boulis. TheselatersedimentscontainedMiddlePaleolithicartifacts, animalbones(someofwhichareburnt),andpollenindicativeof mildclimaticconditions(14,16,17).Becauseofcarbonate precipitation,thee ´boulisisinpartcementedintoirregular brecciatedconglomeratemasses,inwhichthereareMiddle Paleolithicartifacts,faunalremains,andvariablyarticulatedand crushedhumanremains.ThehumanfossilsincludethePalomas 92and96partialskeletons,thePalomas93to95teeth,andthe remainsofatleast2moreindividuals,includingcraniawith mandibles. Authorcontributions:M.J.W.,J.G.,andE.T.designedresearch;M.J.W.,J.G.,M.V.L.,A.V.L., A.P.-P.,J.Zapata,J.O.,T.H.,A.P.,J.-L.S.,J.Zilha ˜o,andE.T.performedresearch;M.J.W., M.V.L.,J.Zapata,J.O.,T.H.,A.P.,J.-L.S.,andE.T.analyzeddata;andM.J.W.,T.H.,A.P.,J.-L.S., J.Zilha ˜o,andE.T.wrotethepaper. Theauthorsdeclarenoconictofinterest.1Towhomcorrespondenceshouldbeaddressed.E-mail:trinkaus@artsci.wustl.edu. Thisarticlecontainssupportinginformationonlineat www.pnas.org/cgi/content/full/ 0811213106/DCSupplemental . ©2008byTheNationalAcademyofSciencesoftheUSAwww.pnas.org cgi doi 10.1073 pnas.0811213106PNASDecember30,2008vol.105no.5220631–20636 ANTHROPOLOGY

PAGE 2

ChronologicalAgeofthePalomasHumanRemains.Theageofthe humanfossilsofconcernhereisprimary,becausetheissueisthe morphologyofthelatestNeandertalsinEurope.ThePalomas artifactsareallMiddlePaleolithic,madeonretouchedflakesof flint,quartz,rockcrystal,andmarble(14–16,18).Givencurrent reliabledatesforIberianMiddlePaleolithicassemblages(10, 19),thisindicatesanageof 34kacalBP( 30ka14CBP)for thedeposits. CollagenpreservationispoorinthePalomashumanremains, anddirect14Cdatingofthehumanboneshasnotbeenpossible. However,acombinationofdatingtechniquespermitschronologicalcontroloftheUpperCutting(see SIAppendix ,Figs.7and 8,forthestratigraphicpositionsofthesamples). AburntfaunalbonedirectlyadheringtothePalomas59 mandibleinlevel2fprovidedadateof34,450 60014CBP (OxA-10666)[95%CI:40,950–37,622calBPusingtheCariaco0614Ccurve(20)].Aburntlagomorphbonefromthedeeper level2lprovidedastatisticallyidenticalageof35,030 27014C BP(OxA-15423)(95%CI:40,986–38,850calBP).Giventhe naturesofthedatingspecimensandthemoderatelyhighC:N atomicratiosforthesesamples(albeitanexpectedelevation giventheburning),itislikelythattheseagesfallclosertothe upperlimitsofthe95%confidenceintervalsforthesedates( SI Appendix ,SectionIII).Thisinterpretationofthesedatesfrom theuppersedimentfillissupportedbytheassociatedpalynology (17),whichshowsrelativelytemperateconditions,probably duringGIS9andthereforeslightlybeforetheonsetofthe severelycoldHeinrich4oscillation 40kacalBP(21). Toassessfurthertheagesofthesedepositsandthestratigraphicallyoldere ´boulisdeposits,U-series(LA-ICP-MS)dates wereobtainedfrom3bones( SIAppendix ,SectionIV).Adate of43,800 750calBP(APSLP4)isfromafaunalspecimenfrom level2i;itisslightlyearlierthanthelikely14Ccalibratedranges fromlevels2fand2l.Twomucholderandstatisticallysimilar U-seriesdatescomefromthestratigraphicallydeeper,steeply slopinge ´boulis:oneonaPalomas96metacarpalfromlevel2e of54,000 3,850calBP(APSLP1)andtheotheronafaunal bonefromlevel2lof51,000 1,250calBP(APSLP6).These U-seriesestimatespresumethatthesampleswereclosedsystems,whichcannotbeverified.Theyshouldthereforeberegardedonlyascorroboratingthe14Candpaleoclimatological assessmentsoftheageoftheuppersedimentfillandtheearlier ageofthee ´boulislevel,includingsomemixedmaterialinthe burntdark-graysedimenthorizon. Inaddition,asedimentsample(X2509)fromthetopoflevel 2kdirectlyoverlyingthemarbleslabandhencebelowthe dark-graysedimentlevelinthenortheasterncorneroftheUpper Cuttingwasdatedbyusingopticallystimulatedluminescence (OSL)( SIAppendix ,SectionV).Thesampleprovidedanage estimateof54,700 4,700calBPforthesesediments.This determinationincreasesconfidenceintheU-seriesdatesfor stratigraphicallysimilarspecimensandhenceintheageofthe stratigraphicallyyoungerdepositsdatedby14C. Thesedatingassessmentsthereforecombinetoindicateanage fortheuppersedimentfill 40kacalBPbutpossiblyslightly older(i.e., 43kacalBP).ThesubsampleofthePalomashuman remainsfromthisportionofthesedimentincludes63elements, 54or85.7%ofwhichareisolatedteethortoothfragments.They makethesePalomasNeandertalsthemostrecent,andlargest, sampleofsouthernIberianlateNeandertalscurrentlyknown. TheotherNeandertalscloseinagearetheInitialUpper PaleolithiconesfromSpy(9),andprobablySaint-Ce ´saire, Arcy-Renne,andVindijaG1(22–24).ThesePalomasfossilsare alsoapproximatelythesameageastheearliestmodernhumans inEurope(4,25),albeitattheotherendofEurope.Iftheearliest phasesoftheAurignacianwereindeedmadebymodernhumans (26,27),thenthePalomasremainsshouldoverlapintimewith modernhumansascloseasthenorthernPyrenees(10).ThePalomasHumanRemains.ThePalomasfossilhumanremains thereforeconsistof3samples.Therearetheundatedand isolatedremainsdiscoveredintheminers’rubble.Therearethe partialface,partialskeletons,andisolatedremainsfromthe brecciatede ´boulis.Andtherearethe63isolatedremainsfrom theexcavateddepositsatorabovethelevelsdatedto 40–43ka calBP. Thediagnosticremainsfromthefirst2samplescanallbe attributedtoLatePleistoceneNeandertals.Fortheremainsout ofcontext,therelevantaspectsincludesupraorbitaltoruspresence(Palomas11,12,and62),retreatingmandibularsymphyses andlateralcorpusthickness(Palomas6and23),incisorand molarocclusalmorphologyandincisorrootlength(Palomas24 and50),manualmiddlephalanxbreadth(Palomas65),and femoraldiaphysealshape(Palomas52).Forthebrecciated remains,theisolatedteethappearundiagnostic,andPalomas96 andtheotherassociatedskeletonsarestilllargelyinbreccia.Yet, thePalomas1mandiblehasaretreatingsymphysis,aretromolar space,aprominentcoronoidprocess,andanasymmetrical mandibularnotch.ThePalomas92partialskeletonalignswith theNeandertalsindistalhumeral,proximalulnarandfemoral diaphysealmorphology,aswellasinferredbodyproportions. AlthoughmanyoftheisolatedremainsfromPalomasare Fig.1. SchematicdrawingofthecurrentprolesoftheUpperCutting(see SIAppendix ,Figs.5and6).( A )Thee ´boulisabovenorthwesterncornerofthe excavatedcutting.( B )Levels2m-2obrecciacontaininghumanbones.( C )Projectionofthee ´boulisscreeslope,whichislessperceptibleinthisprolethanithad beeninnowremovedsectionsparalleltoitintheforeground.( D )Uppermostlimit(levels 2h-2i)oflensofburntsedimentmainlyinthenortheasternareaof thecutting.( E )Lowerlimitofburntashysediment(levels2m-2o)inthenorthernandeasternareaofthecutting.20632www.pnas.org cgi doi 10.1073 pnas.0811213106 Walker etal.

PAGE 3

undiagnosticastohumangroupinaLatePleistocenecontext, therearesufficientindicatorstoaligntheseremainswiththe Neandertals.Innocasedoanyoftheremainsexhibituniquely derivedcharacteristicsofmodernhumans(cf.,ref.28). GiventheconcernwithlateNeandertalpaleobiology,the considerationsherearelimitedtotheremainsfoundinsituator abovethe40–43kacalBPlevels.Thecomparativesamples consistofMIS5–3westernEurasianNeandertals,MIS5Middle Paleolithicearlymodernhumans(MPMH),circum-MediterraneanEarlyUpperPaleolithicmodernhumans(EUP)( 33ka calBP),andwesternEurasianMiddleUpperPaleolithicmodern humans(MUP)( 33–24kacalBP).TheMandibles.Fourpartialmandibleswerefoundinsituinthe youngerlevels(Palomas49,59,80,and88)(Fig.2).Palomas59 isaleftcorpuslackingthefullsymphysis,andtheothersare variablycompleteimmaturespecimens. ThepreservedboneofPalomas59indicatesthatithada relativelyverticalsymphysisbutnoprominentdevelopmentof eitheratubersymphyseosorlateraltubercles[probablymentum osseumrank3(29)].TheimmaturePalomas49hasasimilaror moreretreatingsymphysealprofile.Thedentalarcadeisonly intactforPalomas49,andaswithotherveryyoungNeandertals (30),itsbi-dc1externalarcadediameter(35.2mm)isbeyond thoseofsimilarlyagedearlymodernhumans(Fig.3),including thatoftheearlierAurignacianLaQuina-Aval4mandible( 30.0 mm).Palomas59hadaretromolarspace,andPalomas80hasa prominentcoronoidprocessandanasymmetricalmandibular notch,butanopenmandibularforamen.Mostofthesefeatures alignthemprincipallywiththeNeandertalsamongMIS5–3 humans(8). Atthesametime,thementalforaminaoftheadultPalomas 59(atP4M1)andtheinfantPalomas49and88(atdm1)are moderatelymesial(8,31),butthejuvenilePalomas80mandible hasanunusuallymesialmentalforamen,becauseitwasdistinctly mesialoftheP3P4break.Thereislittledifferenceinlateral mandibularcorpusheightbetweenNeandertalsandearlymodernhumans(Kruskal–Wallis P 0.251),butthereisasignificant ( P 0.0002)differenceinbreadth( SIAppendix ,Table4).The otherPalomasmandibles(1,6,and23)arewithotherNeandertalsincorpusbreadth(Fig.4).Palomas59,however,isamong theEUPandMUPmodernhumans;itscorpusbreadthis2.21 standarddeviationsfromtheNeandertalmean(withtheKebara 2highoutliertrimmed,1.97standarddeviationswithKebara2 included).TheDentitions.Theabundanceofisolatedteeth,plusthosein Palomas59and80,makeseveralobservationspossible.Allfour ofthemaxillarycentralincisors(I1s:Palomas34,73,79,and90) exhibitmoderatetomarkedlabialconvexity,andthe3lingually preservedoneshavelargemarginalridgesandlingualtubercles (Fig.5).The2maxillarycanines(C1s:Palomas35and74)have verysmalllingualtubercles,andPalomas35haslittleifany shoveling(Fig.5).Ofthe4P4s(Palomas57,59,78,and87),3 eachexhibitatransversecresteventhoughonlythePalomas59 crestispronounced(Fig.6).Allofthemhaveamesially displacedmetaconid,and3haveextralingualcusps.Onlyoneof theP4s,Palomas59,haslingualasymmetry,butitprobably lackedextralingualcusps.Yet,therearenoconsistentassociationsbetweenthesetraitsacrossthe4PalomasP4s.Ofthe5first andsecondlowermolars(M1sandM2s:Palomas29,80,and84, withoneeachfromPalomas59),allexhibitanteriorfoveabut3 lackmidtrigonidcrests(Fig.7).TheoneM2(Palomas36)hasa skewedprofileandcentrallyplacedcusps. MostofthesedentalocclusaltraitsoccurinboththeNeandertalsandotherPleistocene(andrecent)humansamples,and Fig.2. OcclusalviewoftheimmaturePalomas(SP)49mandibularcorpusand lateralviewsofthePalomas59,80and88mandibles.Palomas59isin norma lateralis ,andPalomas80and88areintheplanesoftheirlateralcorpori.Scale incentimeters. Fig.3. Bivariateplotofexternalbi-deciduouscanine(dc1)dentalarcade breadthversusdevelopmentalageforimmatureLatePleistocenehuman mandibles.Symbols:blackdiamond,Palomas49;graycircles,Neandertals; blacksquares,MPMH;blacktriangle,EUPmodernhuman(LaQuina-Aval4); opentriangles,MUPmodernhumans.Agesarebasedondentalcalcication relativetoextanthumans. Fig.4. Bivariateplotofmandibularcorpusbreadthversusheightatthe mentalforamen,forLatePleistocenematuremandibles.SymbolsasinFig.2; numberedsymbolsareforPalomas1,6,23,and59,therst3ofwhichare geologicallyolderthanPalomas59orundated.Theearlymodernhumans withhighcorpusbreadthsareQafzeh9andSkhul4(MPMH),NazletKhater2 (EUP),andCro-Magnon1(MUP);thehighNeandertaloutlierisKebara2.Walker etal. PNASDecember30,2008vol.105no.5220633ANTHROPOLOGY

PAGE 4

notallNeandertalsexhibitallofthem(32).However,theseare alltraitsthatoccurinhighfrequenciesamongtheNeandertals, andNeandertalsinparticularhavehighfrequencies,compared withotherPleistocenesamples,ofthesetraitsoccurringin combination.OfthesePalomasteeth,onlytheI1sandtheM2exhibitthefullsuiteoffeaturesincreasinglyconsideredcharacteristicoftheNeandertals. Rootlengths,especiallyofanteriorteeth,havebeenshownto largelydifferentiateNeandertalsandUpperPaleolithichumans (33);additionaldata( SIAppendix ,Table5)indicateasignificant comparativesampledifferenceinrootlengthsforallbuttheI2. Ofthe13Palomasanteriorteethprovidingrootlengths,4are belowtheNeandertalranges,4are 2fromtheNeandertal means,and77%arebelowtheNeandertalmeans. ThePalomas59M1exhibitssupraradiculartaurodontism,and theM2haspronouncedradicularendotaurodontism,thelatter especiallybeingcharacteristicoftheNeandertals(34).Yet,the Palomas29M2lacksanypulpchamberexpansion. Neandertalsandearlymodernhumanshavesimilarpostcaninedentaldimensions(35),buttheformerhavegreaterI1and I2labiolingualcrowndiameters(Kruskal–Wallis P 10 6for each)butcontrastlessinC1breadths(Kruskal–Wallis P 0.025) ( SIAppendix ,Table6).OneoftheinsituPalomasI1s(Palomas 21)and3ofthe5C1s(Palomas26,54,59)arebelowthe Neandertalrange,andtheremainderofthePalomasanterior mandibularteethareatorbelowtheNeandertalmeans[2-tailed Wilcoxon P 0.047(I1),0.312(I2),0.013(C1)].ThePostcrania.Inthepostcrania,despitemultipleelements( SI Appendix ,Table1),only1diagnosticboneisstratigraphically secureinthemorerecentdeposits,thePalomas28distalhand phalanx.Ithasabroad,roundedapicaltuberositylackingungual spines(Fig.8),anarchaic Homo configuration(36,37).Neandertaldistalphalangealbreadthsaresignificantlyabsolutely broaderthanthoseofalmostallearlymodernhumans(Fig.8; Kruskal–Wallis P 0.00003)( SIAppendix ,Table7),andmost ofthemarebroaderrelativetophalangeallengthdespitethe relativelylongerdistalphalangesoftheNeandertals(38).The distalbreadthofthePalomas28phalanx(9.7mm)fallsabsolutelyandrelativelyamongthosearchaichumans(Fig.8and SI Appendix ,Table7). DiscussionNeandertalAffinities.Theseconsiderationsoftheinsituhuman remainsfromtheupperlevelsoftheSimadelasPalomas confirmthattheyarebestseenaslatesouthwesternEuropean Neandertals.Thereisasuiteoffeatures,includingmandibular symphysealconfiguration,ramalshape,dentalocclusalmorphology,andmanualdistalphalanxshape,thatplacesthemwith archaic Homo andseparatefromearlymodernhumans.Moreover,theI1sandtheP4s,M2,M1sandM2sexhibitapparently derivedNeandertalocclusaltraitsorcombinationsoftraits. Otherretainedplesiomorphousaspectslostamongearlymodern humansorautapomorphoustraitsoftheNeandertals(28)are notpreservedorevidentonlyonundatedorolderPalomas fossils.Theseconsiderationsshouldnonethelessbesufficientto Fig.6. OcclusalviewsofPalomas(SP)mandibularsecondpremolars(P4s). Scaleinmillimeters. Fig.7. OcclusalviewsofPalomas(SP)molars.SP36,maxillaryM2;SP59, mandibularM1andM2;SP80,mandibularM2;SP29,mandibularM2;SP84, mandibularM1.Scaleinmillimeters. Fig.8. Bivariateplotofdistalphalangealbreadthversusarticularlengthfor Palomas28andLatePleistocenesamples.SymbolsasinFig.2.Givenuncertaintiesindigitassignmentforisolatedray2–4distalphalanges,valuesare averagedforthoseindividualpreservingmultipledistalphalangestoprovide anindividualvalue. Fig.5. LingualviewsofPalomas(SP)maxillarycentralincisors(I1s)(SP34,79 and90)andmaxillarycanines(C1s)(SP35and74).Scaleinmillimeters.20634www.pnas.org cgi doi 10.1073 pnas.0811213106 Walker etal.

PAGE 5

confirmthatat 40–43kacalBP,insoutheasternIberia,the MiddlePaleolithicpopulationconsistedofNeandertals.MorphologicalVariability.Atthesametime,itisevidentthatthe latePalomasNeandertalsexhibitacomplexmixofNeandertal andmore‘‘modern’’features.Threeofthemandibleshave mentalforaminathataremoderatelymesialforNeandertals,but oneisunusuallyso.DistinctivedentalfeaturessuchaslargeC1lingualtubercles,M1andM2midtrigonidcrest,P4transverse crest,lingualtubercles,mesialmetaconidandlingualasymmetry,longanteriortoothroots,anteriorcrowndimensions, andlowermolartaurodontismarereducedorvariablypresent inthesample.NoneoftheP4shasall4ofthe‘‘Neandertal’’ configurations. ItispossibletofindindividualNeandertalteethormandibles thatexhibitoneormoreofmostthese‘‘non-Neandertal’’ aspects,andbiologicalvariationacrosstheNeandertals(includingtrendsthroughtimeandclinalvariationinspace)hasbeen noted(7,39–42).However,thelevelofvariationinthese featuresinthePalomaslateMiddlePaleolithicsampleisunusual foragroupofNeandertals.Itispossiblethatthesecontrastswith otherNeandertalsrepresent:( i )lateNeandertalgeneticdriftin thedirectionofmodernhumanmorphologythroughisolationby-distanceinthecul-de-sacofsouthernIberia,( ii )anadaptive shifttolocalenvironmentalconstraintsinsomeofthesefeatures, and/or( iii )theproductofgeneflowfromearlymodernhuman populationstotheimmediatenorth. Thefirstexplanationwouldemphasizeregional(perhaps clinal)variationamongtheNeandertals( cf. ,refs.7and40). SecurelydatedearlierMIS4–3Neandertalsfromsouthofthe Pyrenees[e.g.,Banyoles(43,44),CovaNegra(45),Gegant(46), Valdegoba(47),Zafarraya(48),plustheearlierPalomasremains]exhibittheNeandertalpatternincomparableelements (widelateralmandibularcorpori,relativelyposteriormental foramina,largeanteriorteeth,anteriorfoveaeandmidtrigonid crestsonM1–M3,andasymmetricalP4swithmesialmetaconids andlingualtubercles)withminorvariationinmentalforamen positionandacoupleofthedentaltraits.Thesefossilsthusimply thatthePalomasvariationisnotmerelytheresultoflong-term isolationofIberianNeandertalsthroughtheearlierLatePleistocene.Moreover,suchisolationwouldnotnecessarilyexplain thepresenceofautapomorphousmodernhumantraits(e.g.,a narrowmandibularcorpus,reducedC1lingualtubercles,short anteriortoothroots,orsmallanteriordentalcrowns)ina Neandertalpopulation.Thesecondexplanationwouldimply thatsomeofthesemandibularanddentalfeaturesconferan advantageonthesepopulations,althoughvariationinsomeof thefeaturesmaywellbeselectivelyneutral.ThethirdconsiderationwouldrequirethattheearliestAurignacianofthe northernPyreneesandelsewherebetheproductofmodern humans,toprovideageographicallyproximatesourceforgene flow.Itwouldalsoinvokehumanbiologicalcontactacrossthe ‘‘EbroFrontier’’duringatimewhenthereislittleevidencefor culturaldiffusion(10).Inthisscenario,theculturalcontrasts acrossthe‘‘EbroFrontier’’wouldbeduetobehavioralchoices, possiblyecologicallydriven,ratherthanisolationofthesouthern Iberianpopulations. Conclusion ThehumanremainsfromtheSimadelaPalomasinsoutheastern IberiathereforedocumentthepresenceofNeandertals,relativelylateintheMiddlePaleolithic.Theyhelptosubstantiate thattheMiddlePaleolithicoftheregionwastheproductof Neandertals,eventhoughdiagnostichumanremainsassociated withtheverylatestphasesofthistechnocomplexinEurope remainelusive.AtthesametimethatthePalomashumans exhibitasuiteofderivedNeandertalfeaturesandarchaic Homo configurationslongsincelostamongearlymodernhumans,their morphologicalvariationindicatesthattheydeviatefromthe expectedNeandertalrangesofvariation.Thispatternmaybe resultofgeneticdriftinrelativeisolation,directionalchangeor, perhapsmorelikely,populationcontacttothenorth.ACKNOWLEDGMENTS. WethankthelandownersofCabezoGordoandthe managementoftheCabezoGordoS.A.quarryforallowingtheexcavationsto occur,T.RodríguezandJ.L.PoloforprovidingX-raydiffractionanduorescence analyses,andthemanycolleaguesandvolunteersovertheyearsforenthusiasticallyprovidingassistanceandexpertise.ThisworkwassupportedbytheSpanish government(CGL2005/02410/BTE;BOS/2002/02375;PB/98/0405;PB/92/0971),the Murciangovernment(PSH93/52;05584/ARQ/07;CTC/DGC/SPH/063/2001;CCE/ DGC/IPH/SAR0/1998;CCE/DGC/IPH/SAR/1997;CCE/DGC/IPH/SAR/011/1996;CCE/ DGC/IPH/SAR/1995;CCE/DGC/IPH/SAR/1994;PSH/93/52),themunicipalityofTorre Pacheco,andtheEarthwatchInstitute(1994–2001).Analysisofthehuman remainswassupportedbytheSpanishgovernment(CGL2007–60802/BTE) (A.P.-P.)andWashingtonUniversity(E.T.).1.SmithFH,Jankovic ´I,Karavanic ´I(2005)Theassimilationmodel,modernhumanorigins inEurope,andtheextinctionoftheNeandertals. QuatInt 137:7–19. 2.TrinkausE(2007)EuropeanearlymodernhumansandthefateoftheNeandertals. Proc NatlAcadSciUSA 104:7367–7372. 3.SchmitzRW, etal. (2002)TheNeandertaltypesiterevisited:InterdisciplinaryinvestigationsofskeletalremainsfromtheNeanderValley,Germany. ProcNatlAcadSciUSA 99:13342–13347. 4.TrinkausE, etal. (2003)AnearlymodernhumanfromthePes ¸teracuOase,Romania. ProcNatlAcadSciUSA 100:11231–11236. 5.WildEM, etal. (2005)DirectdatingofEarlyUpperPalaeolithichumanremainsfrom Mladec . Nature 435:332–335. 6.TrinkausE(2005)Earlymodernhumans. AnnuRevAnthropol 34:207–230. 7.RosasA, etal. (2006)PaleobiologyandcomparativemorphologyofalateNeandertal samplefromElSidro ´n,Asturias,Spain. ProcNatlAcadSciUSA 103:19266–19271. 8.SocaruA,Dobos ¸A,TrinkausE(2006)EarlymodernhumansfromthePes ¸teraMuierii, BaiadeFier,Romania. ProcNatlAcadSciUSA 103:17196–17201. 9.SemalP, etal. (2008)NewdataonthelateNeandertals:DirectdatingoftheBelgian Spyfossils. AmJPhysAnthropol ,inpress. 10.Zilha ˜oJ(2006)ChronostratigraphyoftheMiddle-to-UpperPaleolithictransitioninthe Iberianpeninsula. Pyrenae 37:7–84. 11.HublinJJ,BarrosoRuizB,MedinaLaraP,FontugneM,ReyssJL(1995)TheMousterian siteofZafarraya(Andalucia,Spain):DatingandimplicationsonthePaleolithicpeoplingprocessesofwesternEurope. CRAcadSci 321:931–937. 12.MichelV, etal. (2003) ElPleistoceneSuperiordellaCuevadelBoquetedeZafarraya , edBarroso-RuízC(ConsejeríadeCultura,JuntadeAndalucía,Sevilla),pp115–155. 13.TrinkausE,MakiJ,Zilha ˜oJ(2007)MiddlePaleolithichumanremainsfromtheGrutada Oliveira(TorresNovas),Portugal. AmJPhysAnthropol 134:263–273. 14.WalkerMJ, etal. (1998)TwoSESpanishMiddlePalaeolithicsiteswithNeanderthal remains:SimadelaPalomasdelCabezoGordoandCuevaNegradelEstrechodelRío Quípar(MurciaProvince). InternetArchaeol 5.Availableathttp://intarch.ac.uk/ journal/issue5/).AccessedOctober24,2008. 15.WalkerMJ, etal. (1999)ExcavationsatnewsitesofearlymaninMurcia,Simadelas PalomasdelCabezoGordoandCuevaNegradelEstrechodelRíoQuípardela Encarnacio ´n. HumEvol 14:99–123. 16.WalkerMJ(2001) AVeryRemotePeriodIndeed ,edsMillikenS,CookJ(Oxbow, Oxford),pp153–159. 17.Carrio ´nJS, etal. (2003)Glacialrefugiaoftemperate,MediterraneanandIbero-North Africanorainsouth-easternSpain:NewevidencefromcavepollenattwoNeanderthalmansites. GlobEcolBiogeog 12:119–129. 18.WalkerMJ, etal. (2004) SettlementDynamicsintheMiddlePalaeolithicandMiddle StoneAge2 ,edConardNJ(Kern,Tu ¨bingen),pp461–511. 19.Zilha ˜oJ,PettittP(2006)OnthenewdatesforGorham’sCaveandthelatesurvivalof IberianNeanderthals. BeforeFarming 2006(3):95–122. 20.HughenK,SouthonJ,LehmancS,BertrandaC,TurnbullJ(2006)Marine-derived14C calibrationandactivityrecordforthepast50,000yearsupdatedfromtheCariaco Basin. QuatSciRev 25:3216–3227. 21.SepulchreP, etal. (2007)H4abrupteventandlateNeanderthalpresenceinIberia. EarthPlanetSciLett 258:283–292. 22.MercierN, etal. (1991)ThermoluminescencedatingofthelateNeanderthalremains fromSaint-Ce ´saire. Nature 351:737–739. 23.BaileySE,HublinJJ(2006)DentalremainsfromtheGrotteduRenneatArcy-sur-Cure (Yonne). JHumEvol 50:485–508. 24.HighamT,BronkRamseyC,Karavanic ´I,SmithFH,TrinkausE(2006)Reviseddirect radiocarbondatingoftheVindijaG1UpperPaleolithicNeandertals. ProcNatlAcadSci USA 103:553–557. 25.Zilha ˜oJ, etal. (2007) RethinkingtheHumanRevolution ,edsMellarsP,BoyleK, Bar-YosefO,StringerC(McDonaldInstituteofArchaeology,Cambridge,UK),pp 249–262. 26.Henry-GambierD,MaureilleB,WhiteR(2004)Vestigeshumainsdesniveauxde l’AurignacienanciendusitedeBrassempouy(Landes). BullMemSocAnthropolParis 16:49–87.Walker etal. PNASDecember30,2008vol.105no.5220635ANTHROPOLOGY

PAGE 6

27.BaileySE,HublinJJ(2005)WhomadetheEarlyAurignacian?Areconsiderationofthe Brassempouydentalremains. BullMemSocAnthropolParis 17:115–121. 28.TrinkausE(2006)ModernhumanversusNeandertalevolutionarydistinctiveness. Curr Anthropol 47:597–620. 29.DobsonSD,TrinkausE(2002)Cross-sectionalgeometryandmorphologyofthemandibularsymphysisinMiddleandLatePleistocene Homo.JHumEvol 43:67–87. 30.MallegniF,TrinkausE(1997)AreconsiderationoftheArchi1Neandertalmandible. J HumEvol 33:651–668. 31.CoqueugniotH(1999)Lecra ˆned’ Homosapiens enEurasie:Croissanceetvariation depuis100,000ans. BritArchaeolRepS 822:1–197. 32.BaileySE(2006)Beyondshovel-shapedincisors:Neandertaldentalmorphologyina comparativecontext. PeriodBiol 108:253–267. 33.BaileySE(2005) CurrentTrendsinDentalMorphologyResearch ,edZadzinskaE (UniversityofLodzPress,Lodz),pp201–210. 34.KallayJ(1970) Krapina1899 1969 ,edMalezM(JugoslavenskeakademijeznanostiI umjetnosti,Zagreb),pp165–176. 35.TrinkausE(2004)in Miscala ´neaenHomenajeaEmilianoAguirre ,edRubioS(Museo Arqueolo ´gicoRegional,Alcala ´deHenares),pp393–398. 36.TrinkausE(1983) TheShanidarNeandertals (Academic,NewYork). 37.RosasA(1985)FalangeshumanasdelaSimadeslosHuesos(CuevaMayor),Sierrade Atapuerca(Burgos).Estudioanatomicoycomparativo. ActasIVCongEspan ˜ol AntropolBiol 4:557–566. 38.VillemeurI(1994) LaMaindesNe ´andertaliens (CentreNationaldelaRecherche Scientique,Paris). 39.SmithFH(1984) TheOriginsofModernHumans ,edsSmithFH,SpencerF(Liss,New York),pp137–209. 40.TrinkausE(1991)LeshommesfossilesdelaGrottedeShanidar,Irak:E ´ volutionet continuite ´parmileshommesarchaïquestardifsduProche-Orient. L’Anthropol 95:535– 572. 41.TrinkausE,ChurchillSE,RuffCB,VandermeerschB(1999)Longboneshaftrobusticity andbodyproportionsoftheSaint-Ce ´saire1Cha ˆtelperronianNeandertal. JArchaeol Sci 26:753–773. 42.Caramelli etal. (2006)AhighlydivergentmtDNAsequenceinaNeandertalindividual fromItaly. CurrBiol 16:630–632. 43.MarotoJ(ed)(1993)LamandíbuladeBanyolesenelcontextdelsfo `ssilshumansdel Pleistoce `. CtrInvestArqueolGirona 13:1–194. 44.Gru ¨nR, etal. (2006)ESRandU-seriesanalysesofenamelanddentinefragmentsofthe Banyolesmandible. JHumEvol 50:347–358. 45.ArsuagaJL, etal. (2007)NewNeandertalremainsfromCovaNegra(Valencia,Spain). JHumEvol 52:31–58. 46.DauraJ, etal. (2005)ANeandertalmandiblefromtheCovadelGegant(Sitges, Barcelona,Spain). JHumEvol 49:56–70. 47.QuamRM, etal. (2001)HumanremainsfromValdegobaCave(Hue ´rmeces,Burgos, Spain). JHumEvol 41:385–435. 48.Barroso-RuízC,LumleyMAde,CaparrosM,VerduL(2003) ElPleistoceneSuperiordel laCuevadelBoquetedeZafarraya ,edBarroso-RuízC(ConsejeríadeCultura,Juntade Andalucía,Sevilla),pp327–387.20636www.pnas.org cgi doi 10.1073 pnas.0811213106 Walker etal.


printinsert_linkshareget_appmore_horiz

Download Options

close
No images are available for this item.
Cite this item close

APA

Cras ut cursus ante, a fringilla nunc. Mauris lorem nunc, cursus sit amet enim ac, vehicula vestibulum mi. Mauris viverra nisl vel enim faucibus porta. Praesent sit amet ornare diam, non finibus nulla.

MLA

Cras efficitur magna et sapien varius, luctus ullamcorper dolor convallis. Orci varius natoque penatibus et magnis dis parturient montes, nascetur ridiculus mus. Fusce sit amet justo ut erat laoreet congue sed a ante.

CHICAGO

Phasellus ornare in augue eu imperdiet. Donec malesuada sapien ante, at vehicula orci tempor molestie. Proin vitae urna elit. Pellentesque vitae nisi et diam euismod malesuada aliquet non erat.

WIKIPEDIA

Nunc fringilla dolor ut dictum placerat. Proin ac neque rutrum, consectetur ligula id, laoreet ligula. Nulla lorem massa, consectetur vitae consequat in, lobortis at dolor. Nunc sed leo odio.