Deep Skull from Niah Cave and the Pleistocene Peopling of Southeast Asia


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Deep Skull from Niah Cave and the Pleistocene Peopling of Southeast Asia

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Deep Skull from Niah Cave and the Pleistocene Peopling of Southeast Asia
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Frontiers on Ecology and Evolution
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Curnoe, Darren
Datan, Ipoi
Taçon, Paul S. C.
Ung, Charles Leh Moi
Sauffi, Mohammad S.
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Modern Humans ( local )
Pleistocene ( local )
Southeast Asia ( local )
Australasia ( local )
Niah Cave ( local )
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serial ( sobekcm )

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The Deep Skull from Niah Cave in Sarawak (Malaysia) is the oldest anatomically modern human recovered from island Southeast Asia. For more than 50 years its relevance to tracing the prehistory of the region has been controversial. The most widely held view, originating with Brothwell's 1960 description and analysis, is that the Niah individual is related to Indigenous Australians. Here we undertake a new assessment of the Deep Skull and consider its bearing on this question. In doing so, we provide a new and comprehensive description of the cranium including a reassessment of its ontogenetic age, sex, morphology, and affinities. We conclude that this individual was most likely to have been of advanced age and female, rather than an adolescent male as originally proposed. The morphological evidence strongly suggests that the Deep Skull samples the earliest modern humans to have settled Borneo, most likely originating on mainland East Asia. We also show that the affinities of the specimen are most likely to be with the contemporary indigenous people of Borneo, although, similarities to the population sometimes referred to as Philippine Negritos cannot be excluded. Finally, our research suggests that the widely supported “two-layer” hypothesis for the Pleistocene peopling of East/Southeast Asia is unlikely to apply to the earliest inhabitants of Borneo, in-line with the picture emerging from genetic studies of the contemporary people from the region.
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Frontiers on Ecology and Evolution, Vol. 4 (2016-06-27).

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ORIGINALRESEARCH p ublished:27June2016 doi:10.3389/fevo.2016.00075 FrontiersinEcologyandEvolution|www.frontiersin.org 1 J une2016|Volume4|Article75 Editedby: K .ChristopherBeard, UniversityofKansas,USA Reviewedby: JrgenKriwet, UniversityofVienna,Austria GraemeWilliamBarker, UniversityofCambridge,UK *Correspondence: DarrenCurnoe d.curnoe@unsw.edu.au Specialtysection: Thisarticlewassubmittedto Paleontology, asectionofthejournal FrontiersinEcologyandEvolution Received: 18February2016 Accepted: 08June2016 Published: 27June2016 Citation: CurnoeD,DatanI,TaonPSC,Leh MoiUngCandSauf)-267(MS(2016)Deep SkullfromNiahCaveandthe PleistocenePeoplingofSoutheast Asia.Front.Ecol.Evol.4:75. doi:10.3389/fevo.2016.00075 DeepSkullfromNiahCaveandthe PleistocenePeoplingofSoutheast Asia DarrenCurnoe 1 * ,IpoiDatan 2 ,PaulS.C.Taon 3 ,CharlesLehMoiUng 2 and MohammadS.Sauf)]TJ/T1_4 5.23 Tf88.038 3.258 Td(2 1 Palaeontology,GeobiologyandEarthArchivesResearchCentre,SchoolofBiological,EarthandEnvironmentalSciences, UniversityofNewSouthWales,Sydney,NSW,Australia, 2 SarawakMuseumDepartment,Kuching,Malaysia, 3 Place, EvolutionandRockArtHeritageUnit,SchoolofHumanities,LanguagesandSocialScience,GrifthUniversity,GoldCoast, QLD,Australia TheDeepSkullfromNiahCaveinSarawak(Malaysia)istheoldestanatomicallymodern humanrecoveredfromislandSoutheastAsia.Formorethan50yearsitsrelevanceto tracingtheprehistoryoftheregionhasbeencontroversial.Themostwidelyheldview, originatingwithBrothwell's1960descriptionandanalysis,isthattheNiahindividualis relatedtoIndigenousAustralians.HereweundertakeanewassessmentoftheDeep Skullandconsideritsbearingonthisquestion.Indoingso,weprovideanewand comprehensivedescriptionofthecraniumincludingareassessmentofitsontogenetic age,sex,morphology,andafnities.Weconcludethatthisindividualwasmostlikelyto havebeenofadvancedageandfemale,ratherthananadolescentmaleasoriginally proposed.ThemorphologicalevidencestronglysuggeststhattheDeepSkullsamples theearliestmodernhumanstohavesettledBorneo,mostlikelyoriginatingonmainland EastAsia.Wealsoshowthattheafnitiesofthespecimenaremostlikelytobewith thecontemporaryindigenouspeopleofBorneo,although,similaritiestothepopulation sometimesreferredtoasPhilippineNegritoscannotbeexcluded.Finally,ourresearch suggeststhatthewidelysupported“two-layer”hypothesisforthePleistocenepeopling ofEast/SoutheastAsiaisunlikelytoapplytotheearliestinhabitantsofBorneo,in-linewith thepictureemergingfromgeneticstudiesofthecontemporarypeoplefromtheregion. Keywords:modernhumans,Pleistocene,SoutheastAsia,Australasia,NiahCave INTRODUCTION DiscussionsabouttheinitialsettlementofSoutheastAsiaandAustralasiabyanatomicallymodern humans(AMH)havehistoricallyfocusedonevidencefromasmallnumberofLatePleistocene humanremainsscatteredacrossthisbroadregion( Thorneetal.,1999;Dizonetal.,2002;Dtroit e tal.,2004;Olleyetal.,2006;Barkeretal.,2007,2013;Mijaresetal.,2010;Demeteretal.,2012; Stormetal.,2013).Inmanycases,thegeologicalage,andsometimeseventaxon omicanity,of theseancientremainscontinuestobeuncertain(Thorneetal.,1999;Dizonetal.,2002;Dtroit e tal.,2004;Olleyetal.,2006;Barkeretal.,2007,2013;Mijaresetal.,2010;Demeteretal.,2012; Stormetal.,2013).Thisonlyservestoconfoundthealreadycomplicatedtaskof reconstructing thecolonizationroutesandtimingofthedispersaloftheearliestAMHacrossSoutheastAsia

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Curnoeetal. DeepSkullandthePeoplingofSoutheastAsia andAustralasiaaswellastheirpossiblerelationshipstorec ent populations.Still,withrecentadvancesindatingmethods,severalnewelddiscoveries,thereexaminationofexistin g butpoorlycharacterizedremainsandgeneticinvestigation sof contemporarypopulations(e.g., Capellietal.,2001;Karafetetal., 2001;Hilletal.,2007;Soaresetal.,2008,2016;Tumonggor etal., 2013;Trejautetal.,2014 )itisbecomingclearthattheearliest AMHsettledEastAsiabyatleast45ka( Mijaresetal.,2010; Demeteretal.,2012;Barkeretal.,2013;HuntandBarker,20 14 ), andprobablyintherangeof80–120ka( Hilletal.,2007;Soares etal.,2008,2016;Baeetal.,2014;Liuetal.,2015;Curnoee tal., 2016 ). SuperimposedontheLatePleistocenehistoryoftheregion aremorerecentprehistoricmigrationsbyagriculturalists such asSino-Tibetan,TaiandAustroasiaticspeakingpeopleintomainlandSoutheastAsiaandAustronesianspeakersacrossoceanicSoutheastAsia( Bellwood,1997 ).Theideathatthese migrationsresultedinthereplacementofmostofthehunter-gatherersoftheregonbyNeolithicpopulationshasbeendebat ed nowfor80years(e.g., vanSteinCallenfels,1936;Hooijer,1950; VonKoenigswald,1952;Brothwell,1960;Coon,1962;Bellwo od, 1997;MatsumuraandHudson,2005 ).Inparticular,ithasbeen widelyacceptedthattheseLatePleistocenetoearlyHolocenehunter-gathererswererelatedtorecentIndigenousAustra lians andNewGuineans,potentiallyevenrepresentingtheearliestAMHtohavesettledtheregion( Matsumuraetal.,2008 ).This model,dubbedthe“two-layer”hypothesis( Jacob,1967 ),on accountofthesubsequentreplacementofthesehunter-gathe rers byimmigrantNeolithicpeople( Bellwood,1997 ),hasenjoyed somewhatofarevivaloflate( Matsumuraetal.,2008;Oxenham andBuckley,2016 ),beingextendedeventoencompassJapan ( Kaifuetal.,2011 ). Notably, KrigbaumandManser(2005) undertookatestof thetwo-layerhypothesisemployingremainsfromNiahCave(excludingtheDeepSkull).Usinga3Dmorphometricapproach,theyexaminedfacialshapeincraniafrompre-NeolithicandNeolithiclayersoftheWestMouth,comparingthemtovarioussamplesfromEastAsia,SoutheastAsia,AustraliaandthePac ic. Theirresearchaimedtoassesswhetherthesetemporallydisti nct samplesfromNiahCavemightrepresentdierentbiologicalpopulations.Theresultshighlightedanabsenceofsignican t dierencesinfacialshapebetweensamples,suggestingbiologi cal andtemporalcontinuitybetweenthepre-NeolithicandNeolit hic peopleatNiahCave,aswellashintingatpossibleanitiesofbothgroupstoSoutheastAsianandevenPolynesiangroups( KrigbaumandManser,2005 ).Owingtopoorpreservationofthe remains,however,thestudyincludedonlyasmallsampleofNia h Cavecrania(n4/7)andfew(mostlybilateral)landmarks(n6 /8)in eachanalysis,makingthesignicanceoftheirndingsdic ultto establishrmly. Formorethan50yearstheDeepSkullfromNiahCave ( Figures1A–K )hasbeenakeyspecimeninthedebate surroundingtheoriginsofAMHinSoutheastAsiaaswellasthetwo-layerhypothesis.Thiscranium,lackingamandible,wasrecoveredin1958atthelevelof106–110inchesinatrialtrenchdubbed“Hell”intheWestMouthofthegreatNiahCave(GuaNiah)systeminSarawak,Malaysia( Harrisson,1967 ). Soonafter,a 14 Cdateoncharcoalsuggestedapossibleage ofc39,600 1000BP(GRO1339)forthispartialcranium. Whilesomeresearchershaveraiseddoubtsaboutitsstratigr aphic context,suggestingitmayhavebeenanintrusiveburial( Bellwood,1997;Wolpo,1999 ),recentresearchincluding detailedstratigraphicinvestigations,directuranium-se riesdating ofcranialboneand 14 Cofcharcoalfromadjacentsedimentshas conrmedtheDeepSkulltobeofLatePleistoceneantiquity—derivingfromtheperiodc45–39kaandmostlylikelyaround37ka( Barkeretal.,2013;HuntandBarker,2014 ).Crucially,this makestheDeepSkulltheearliestsecurelydatedAMHremainsinislandSoutheastAsia. Brothwell(1960) hasprovidedtheonlydetailed,but nonethelessincomplete,descriptionofthespecimen,whichwa s publishedmorethan50yearsago.Hetentativelyreportedthatthecraniumbelongedtoanadolescent(15–17yearsofage),wasofunknownsex,andshowedstrongestresemblancestoTasmanians,speculatingthattheDeepSkulllaywithinanevolutionarylineagetothe“Negritoids.”Atthetime,theNegritoid“race”wasseenasoneofthetwofoundingmodernhumanpopulationstohavesettledSoutheastAsiaandAustralasiaduringthePleistocene,theotheronebeingtheso-called“Australoids”orAustralo-Melanesianpeople(e.g., Hooijer,1950;VonKoenigswald,1952 ).Brothwell's identicationofthecraniumasbeingofTasmaniananitywa s alsopremisedontheassumptionthatTasmaniansandmainlandAustraliansbelongedtodierentpopulations,andwhilethiswasawidelyheldviewatthetime,traceabletotheeighteent h andnineteenthcenturyexplorerslikeCook,LaPérouse,andLabillardière( Mulvaney,1958 ),ithassincebeendiscredited(e.g., Presseretal.,2002 ).Nonetheless,hisinferencesandmethods werethesubjectofstrongcriticismbyevenhiscontemporari es ( Macintosh,1965;Mulvaney,1966 ),asituationoverlookedin recentdiscussionsoftheDeepSkull. Asmallnumberofspecialistssubsequentlyoeredopinions abouttheanitiesoftheremains,andtheyhavebeenusefullysummarizedby Kennedy(1977) and KrigbaumandDatan (2005) .Generally,disagreementoverthepastfourdecadeshas centeredonwhether: TheDeepSkulliswellenoughpreservedtoplayanyroleatallindiscussionsaboutLatePleistocenehumanevolutioninSoutheastAsia(e.g., KammingaandWright,1988 ).Although, Kennedy(1977) hasnotedthatfewworkersinthepasthave studiedtheoriginalremains,andthisislargelytrueevenun til today. Itisfromanadolescent( Brothwell,1960 )oranadult( Birdsell, 1978;KrigbaumandDatan,2005 ). Thesexofthespecimenisdiagnosableatall,owingtoitsyoun g ontogeneticage( Brothwell,1960 ),ormightsampleafemale ( Birdsell,1978 ). TheNiahcraniumbelongstoanAustralo-Melanesianpopulation,asrstproposedby Brothwell(1960) ,oragroup morecloselyrelatedtoEastAsians.Whilemostresearcherswhohavediscussedthespecimenhaveaccepted Brothwell's (1960) diagnosis(e.g., Barkeretal.,2007 ), Birdsell(1978) identiedaNegritocomponenttoitsmorphology, Harrisson FrontiersinEcologyandEvolution|www.frontiersin.org 2 June2016|Volume4|Article75

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Curnoeetal. DeepSkullandthePeoplingofSoutheastAsia FIGURE1|DeepSkullfromNiahCave. Calvariain (A) superiorview, (B) leftlateralview, (C) anteriorview, (D) inferior/endocranialview,and (E) posteriorview; (F) leftisolatedparieto-temporo-occipitalfragmentinlate ralview(greenarrow,mastoidprocess;bluearrow,occipit alsquama);maxilla: (G) inferior/palatalview, (H) anteriorview,and (I) enlargedviewofdentalcrowns, (J) rightlateralview,and (K) superior/internalview. FrontiersinEcologyandEvolution|www.frontiersin.org 3 June2016|Volume4|Article75

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Curnoeetal. DeepSkullandthePeoplingofSoutheastAsia (1976) suggesteditmightshowanitiestothecontemporary DayakpeoplefromBorneo,and Wu(1987) proposedthat itbelongedtoa“southernMongoloid”groupthatincludedtheAustronesianspeakingpeopleinhabitingtheregiontoday.Subsequently, Wu(1992) opinedthattheDeepSkullwaspart ofawiderSoutheastAsianrace,whichhassometimesbeendubbed“Malays”or“Proto-Malays”( Jacob,1967,1968 ). Itisclear,therefore,thatdespiteitsLatePleistoceneage ,there isstillnoconsensusabouttheimportanceoftheDeepSkulltoaddressingquestionssurroundingtheoriginsofAMHinSoutheastAsia.Thepresentstudysetsouttotesttheaboveissuesbyproviding:(1)anewdescriptionoftheDeepSkullincludinganassessmentofitspreservation,morphologyandpotentialtoprovidemetricdata;(2)areanalysisofitsontoge netic ageandsex;and(3)areconsiderationofitspossibleanitiestoLatePleistocene/EarlyHoloceneandrecentpopulationsfro m East/SoutheastAsiaandAustralasia,withadiscussionoft he implicationsforcurrentdebatesabouttheoriginsofrecentpopulationsfromtheregion.MATERIALSANDMETHODSAlthoughtwopostcranialbonesaresuggestedtorepresentthesameindividualastheDeepSkull( KrigbaumandDatan, 2005 ),ourfocushereisentirelyonthecranialremainsheld bytheSarawakMuseumDepartmentinKuching(Malaysia).AllobservationsandmeasurementsoftheDeepSkullweremadeandcheckedbyDCduring2013and2014.MorphologicalobservationswerealsomadebyDConarangeoforiginalhumanremainsandcastsincludingPleistoceneremainsfromWadjak,Keilor,KowSwamp,theWillandraLakes,ZhoukoudianUpperCaveandLiujiang.Metricaldataforcomparativematerialsweretakenfromtheliterature( VonBonin,1931; SuzukiandHanihara,1982;Cuong,1986;Brown,1989,2015;Storm,1995;WuandPoirier,1995;Howells,1996;MatsumuraandZuraina,1999;Matsumuraetal.,2001,2008;Bulbeck,2005;GreenandCurnoe,2005;MatsumuraandPookajorn,2005;MatsumuraandHudson,2005;CurnoeandThorne,2006;Demeteretal.,2012;Baeetal.,2014;Xiaoetal.,2014;Liuetal.,2015 )andoncastsororiginalremains byDC. Weundertookmultivariateanalysesof18variablescombini ng continuousanddiscrete(presence/absence)data.Alldataw ere log-transformedpriortoanalysis.WeperformedprincipalcoordinatesanalysisandUPGMAclusteranalysisemployingtheGowerdistanceusingthesoftwarePASTVersion3.11( Hammer etal.,2001 ). RESULTSDevelopmentalAgeandSex Brothwell(1960) reportedthepresenceofanuneruptedM 3 fromtheDeepSkull,whichwascentraltohisdiagnosisofthespecimenasprobablyrepresentinganadolescent(15–17yearsold).Unfortunately,thistoothisnotpresentinthecollecti onof theSarawakMuseumDepartmentandsocouldnotbeassessed inourstudy.WhileM 3 emergenceiscommonlyconsideredtobe askeletalmarkerofadulthood( Hillson,1996 ),thefailureofthe M 3 stoeruptiscommoninrecenthumanpopulations,including indigenouspeopleinSoutheastAsia( TurnerandEder,2006 ). Therefore,whilethepresenceoferuptedM 3 scanreasonably betakentoindicateadulthood,absence,onitsown,cannotb e consideredareliablemarkerofontogeneticage. Severalmarkersofdevelopmentalchronologysuggestthat theDeepSkullwasmostlikelytobefromanadult.First,thespheno-occipitalsynchondrosisisobliterated[notedalsoby Brothwell(1960) ].Closureofthisjointincontemporary populationshasbeenwidelyreportedtooccuronaverageataround17–20yearsofage( Bassedetal.,2010;Ekizoglu etal.,2016 ).Ageestimatesdo,however,varyonaccountof sex,geographicpopulationandmethodofassessment,withcombinedsexestimatesofcompleteclosureintherangeofc12–25yearsold( Bassedetal.,2010;Ekizogluetal.,2016 ).Thus, obliterationofthespheno-occipitalsynchondrosisimpliesa nage ofatleastadolescence,butmorelikelyearlyadulthood,int he caseoftheDeepSkull.Someofthevaultsuturescanalsobeassessedforclosureandarecharacterizedbyadvancedfusi on inseverallocations,givingtheimpressionthattheDeepSku llis fromanadult.Forexample, MeindelandLovejoy's(1985) widely deployedcriteriaappliedinternallyatbregmaandmid-corona l (othersutureslikelytobemisleadingowingtotaphonomicdamage)indicateadvancedfusion(scores2–3).Again,onit s own,theseareinsucientgroundstodiagnoseitsontogeneti c age. ThedegreeoftoothwearseenontheDeepSkull'sM 1 and M 2 isoneofthemostrevealingfeaturesintermsofdiagnosing itspersonalage.Thelevelofocclusalwearwouldbelargeforanadolescentevenamonghunter-gatherers( Figure1I ).For example,applying Lovejoy's(1985) standardsforagebasedon occlusalwearprovidesanestimateof40–50years(minimallyPhaseH)fortheNiahCaveindividual'smolars. Finally,thereisevidencefordegenerativechangesparticu larly onthearticularsurfaceofthemandibularfossaandtherightoccipitalcondylepossiblyindicatingosteoarthritis.W hile osteoarthritisofthetemporomandibularjointisgenerallyregardedtobeadegenerativediseaseassociatedwithaging, itis knownclinicallytooccurinchildrenandadolescents( Choand Jung,2012 ). Weconcludethatwhileincompletenessandtaphonomic damageprecludesadetailedassessmentoftheontogeneticag e oftheDeepSkull,theavailableevidencetakentogethersug gests thisindividualwasanadultatthetimeofdeath,andperhapsev en oneofadvancedage,ratherthananadolescent,asproposedby Brothwell(1960) . RegardingthesexoftheDeepSkull,musclemarkingsareweak overtheentirecranium,withveryweakorabsentcranialcre sting. Thevaultshowsstrongbossing,andisstronglyarched/roun ded ineachregion(frontal,parietalsandoccipital)inallplanes . Featuressuchasthezygomaticarchesaredelicatelybuilt. The mastoidprocessesaresmallandweaklyprojecting( Figure1F ). Scoresforthesexingstandardsof BuikstraandUbelaker(1994) ascanbeappliedaregivenin Table1 .Together,thesefeatures indicatetheDeepSkullislikelytobefromafemale. FrontiersinEcologyandEvolution|www.frontiersin.org 4 June2016|Volume4|Article75

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Curnoeetal. DeepSkullandthePeoplingofSoutheastAsia TABLE1|CranialsexingstandardsappliedtotheDeepSkull a . Trait Score Nuchalcrest 1 Mastoidprocess 1 Supraorbitalmargin 1 Supraorbitalridge/glabella 1 a Standardsfrom BuikstraandUbelaker(1994) . DescriptionandComparisonCalvariaIn Table2 ,weprovidearevisedlistoftheremainingbones oftheDeepSkullaspresentlyheldbytheSarawakMuseumDepartment.Thecalvariashowssignsofdistortioninvariou s placesandisfragmentary( Figure1A ).Theentirecraniumhas beencoatedinshellac,andinparts,PlasterofParishasbeenusedtollcracks.Whileweagreethatitmaybeunfossilized (see Barkeretal.,2007 ),thiswouldneedtobeconrmedwithXRF orasimilaranalyticaltechnique.Distortionofthecalott eisat itsworstontherightfronto-orbitalregionandrightposter ior parietalfragment,whichhavebeendisplacedsuperiorlyalong a majorcrack.Thereisalsoalargecrackthroughthemiddleoftheleftparietal,thetwohalvesbeingessentiallyundistor ted,but withtheposteriorhalfbeingmisalignedduringreconstruct ion. Ontherightside,theparietalscomprisetwomajorpartsintheanteroposteriordirection,theposteriorpartbeingmisalign ed alongacommonfracturewiththeanteriorfragment. Whilealloftheseproblemsleadtoanexaggerationof theglobularityofthevault,itisnonethelessevidentthat the specimenisoverallverydelicatelybuilt,withastronglyar ched vault(sagittalandcoronalplanes)andsportswell-developedfrontalandparietaleminences.Theparieto-occipitalcurvat ure isstronglyverticallyinclinedandlacksthesignsofawell developedoccipitalbun.Thelargestcranialfragmentfromth e DeepSkullhasbeenassembledfrom 23piecesandcomprises alargelycompletefrontalboneandmuchoftheleftandrightparietals.Thefrontalincludesasectionoftherightlatera lorbital roofandsupraorbitaltrigone.Thisregionisdistorted,hav ing beendisplacedposteriorlyandmediallyasaresultofpost-bur ial compression. Ontheleftside,asmallsectionofthelateralroofand supraorbitalmarginispresent:thissideofthecalotteisess entially undistorted.Nosignofasupraorbitalridgecanbediscernedlaterallyontherightorleftsides.Thesuperciliaryregionispreservedontheright,butbarelydiscernible:i.e.,aver y small,scarcelyswollen,eminenceisdiscoverableonlythr ough palpation.Ontheleft,thisregionseemsnottobepreserved,beingundiscerniblethroughobservationorpalpation.Whiletheglabellaisincomplete,itssuperiorhalfispresent,butth is regionofthefrontallackstheinferiorpartandnasalroot.I t seemsclear,however,thatglabellawasveryweaklydevelope d, in-linewiththepoorlydenedsuperciliaryridgesandeviden tly thinsupraorbitaltrigones.Theanterior-mostpartofthefro ntal squamabulges(swolleninappearance)foritsanteriormostth ird. Itthencurvesevenlybacktobregma,thevaultbeinghighlyroundedandarched.Ontherightside,aprominentfrontalbos s TABLE2|FragmentsbelongingtotheDeepSkullheldbytheSara wak Museum. Calvariacomprisingfrontal,andleftandrightparietalfr agments. Temporo-occipital-sphenoidfragment:comprisingleft(p etrous)andright temporal(petrous,partofsquamaandmandibularfossa)bon es,basioccipital fragment,andrightfragmentofgreaterwing. Parieto-occipito-temporalfragment(leftside):withpos teroinferiorparietal squama,occipitalplaneandsomeofthenuchalplane,andlef ttemporal squamaandmastoidpartwithmastoidprocess. Isolatedfragmentofprobabletemporalbone. Zygomaticbone(right,largelycomplete). MaxillawithM 1 andM 2 . Frontalfragment:castIA. Frontalfragment:castIB. Temporo-sphenoidfragment. Temporalfragment:mastoidprocess. Temporalfragment:zygomaticprocess. Spheno-frontalfragment. Probablesphenoidfragment(sphenoidfragment#3). Probablesphenoidfragment(sphenoidfragment#4). Occipitalfragment(leftside,fragment#1). Occipitalfragment(leftside,fragment#2). Occipitalfragment(rightside). Temporalfragment:castII. Ethmoidfragment(cribriformplate). Vomerfragment. Cranialfragmentsheldtogetherbymatrix(parietaland?te mporal). Unidentiedfragment,probablybasicranialfragment. Bagof9“doubtful”fragments. isnotable.Nofrontalkeelingcanbediscerned,buttheregi on straddlingthesagittalsutureisslightlyswollenor“heaped up.” Leftandrighttemporalcrestsarepresent,butpoorlydenedalongmostoftheircourse,indicatingweakscarringassoci ated withthetemporalmuscles.Thetemporalfossaearemoderatelyshallowinkeepingwithitsoverallsize:i.e.,thecraniumha s littlepostorbitalconstrictionofthefrontal.Viewedposter iorly, thewallsarehighlydivergentsuperiorlywiththeparietalbos ses stronglyoverhangingtheinferior,orparieto-temporal,reg ions. Theparietalsareshortandstronglyarchedposteriorlyinboth thesagittalandtransverseplanes:i.e.,theyformadistinct and moreorlessverticalplaneintheirposteriorone-thirdratherthanbulgingposteriorly( Figures1A–E ).Thedenticulationsof thelambdoidsuturearediscernibleanddeeplyinvestedonth e rightsidealongthesuture'smedialtwo-thirds.Theoccipit alwas evidentlytall,narrow,andwidelypeaked,butthepositionoflambdacannotbeaccuratelydetermined. Asecondreconstructedvaultfragmentfromtheleftside comprisesaposteroinferiorpieceoftheparietalsquama,partoftheoccipitalplaneandsomeofthenuchalplane,lefttemporalsquamaandmastoidpartwithmastoidprocess( Figure1F ).The restorationisproblematicinpartduetodicultiesensuingfrompost-burialdistortion.Theoccipitalfragmentlacksanexternaloccipitalprotuberanceorcrest/torusandiswithou t clearevidenceofmusclemarkings.Theoccipitalplaneitselfi s rounded,evidentlydeviatinglittlebeyondthelambdoidsu ture FrontiersinEcologyandEvolution|www.frontiersin.org 5 June2016|Volume4|Article75

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Curnoeetal. DeepSkullandthePeoplingofSoutheastAsia above.Inferiorly,itisstronglyundercutbyasteppednuchalplanethatbulges(curvesinferiorly)markedlyattherstst ep;the secondstepbeingmoreorlesshorizontalandapproachingthecranialbaseproper(i.e.,vicinityoftheforamenmagnumregi on). Viewedsuperiorly,thecalotteisbroadlyovoidinoutline,b ut muchwiderbiparietallythanacrossthefrontal( Figures1A,C ). Theremainingvaultgivestheimpressionthatthecompleteskullwouldhavebeenoverallquitetall,especiallyheightt o bregma,ofmoderatelength,andrelativelybroad. Brothwell (1960) collectedseveralcranialmeasurementsonthespecimen, andwehavere-takensomeofthem,andalsomeasuredothersthatwereavailable( Tables3,4 ).Maximumcranial length(GOL)wasestimatedbyrettingthemainoccipitalfragmenttotheremainingcalotte.Itwasreconstructedbyustobemoderateatc175mm,avaluedieringby < 3% from Brothwell's(1960) estimate.Placingthiswithinabroader geographiccontext,theDeepSkullislikelytohavebeenshor t, itsreconstructedvaluesittingoutsideof(below)therang e ofPleistocene/EarlyHoloceneSoutheastAsianandAustral ian samples( Table3 ). Maximumcranialbreadth(XCB)ispositionedhighonthe parietalsandisreconstructedbyustohavebeenwideatc145mm,although,thisvalueislikelytohavebeenaectedtosomeextentbypost-burialdistortion.Still,ourestimated iers from Brothwell's(1960) determinationby < 4%,conrmingits precision.ItsuggeststhattheDeepSkullwouldhavebeenabr oad calvariacomparedtoPleistocene/EarlyHolocenesamplesandmostrecenthumansamples,itsvaluesittingoutsideofthera nges ofrecentTolaiandAustralians( Table3 ). ReconstructedcranialindexoftheDeepSkullisveryhigh atc83%andsuggestsitsshapemayhavebeenbrachycephalic.Onaverage,allofthePleistocene/EarlyHolocenesamplesaremesocephalic(EastandSoutheastAsian)ordolichocephalic(Australian),withtheDeepSkullsittingabovethemaximumvalueforallfossilsampleranges( Table3 ).Moreover,the reconstructedvaluefortheNiahspecimenliesoutsideoftherangesofseveralrecentsamplesincludingTolai,Australian sand Tasmanians( Table3 ). MaximumfrontalbreadthoftheDeepSkullisreconstructed tohavebeenmoderateatc118mm,however,thismeasurementwasnotrecordedby Brothwell(1960) .Itsvaluesitsbelowthe minimumforbothPleistocene/EarlyHoloceneEastAsianandSoutheastAsiansamples( Table4 ).Itis,however,wellwithinthe rangeofPleistocene/EarlyHoloceneAustralians,andsimi larto manyrecentpopulations( Table4 ).Anindexoffrontalshape (XFB/GOL)suggeststheDeepSkull,atc67%,possessedafronta l ofmoderatewidth.TheestimatefortheDeepSkullisoutsideoftherangeofPleistocene/EarlyHoloceneEastAsianandAustraliansamples( Table4 ),butisclosetotheaveragefor Pleistocene/EarlyHoloceneSoutheastAsians,andwellwit hinthe rangeofanumberofrecentsamplesexceptingAustraliansandTasmanians( Table4 ). AsnasionisnotpreservedontheDeepSkullitisnotpossible toprovideanestimateoffrontallength.However,frontalle ngth representedbytheglabella-bregmachord(c109mm)isestima ted tobeidenticaltothePleistoceneChinesepartialcraniumfromHuanglong,andwithintherangeofPleistocene/Early HoloceneAustralians,recentChineseandrecentAustralia n samples( Table4 ). Endocranially,thesuperiorsagittalsinusispoorlydevelope d, butthefrontalcrestisdamagedanditssizecannotbereliablydetermined( Figure1D ).Themiddlemeningealvessel impressionsarenotlarge/deep,butareelaborate,beinghigh ly arborizedasexpectedforanAMHcranium( Figure1D ).There aretwolargeimpressionsofarachnoidgranulationsontheri ght sideposteriortothecoronalsuture—theyhavenoterodedthetabularbonethough—andstraddleapoorlydenedsuperiorsagittalsinus.Inthecaseoftheleftcoronal,swellingcor responds toanendocranialimpressionprobablyleftbyeitheragiantarachnoidgranulation,ormorelikely,ameningealorcerebr al tumor.Theectocranialimpressionofthislikelypathologicalfeaturecouldrepresenteitherectocranialremodelinginre sponse topressurewhenthevaultwasdeveloping,orperhapsresultedfromlesioningandremodelingofthediploeassociatedwithahaemoglobinopathy.Theendocranialtableismissing,exposin g corticalbonewherethefrontalsinuswouldreside,andthereseemstohaveaverysmallorevenabsenceofafrontalsinusintheDeepSkull.VaultThicknessVaultthicknesscanbedeterminedatanumberoflocationsontheDeepSkull,andin Table5 weprovideandcomparedataat twocommonlycomparedlocations(notrecordedbyBrothwell).Thicknessatbregmaisverythinat4.4mm,withthespecimensitingbelowtheminimumofPleistocene/EarlyHoloceneEastAsianandAustralianandChineseNeolithicsamples( Table5 ). Amongrecentpopulations,itsvalueisalsosmall,butsitswithi n theirranges( Table5 ).Thicknessattheparietaleminenceisalso thinat4.5mm,againsittingoutsideofthePleistocene/EarlyHoloceneEastAsianandAustralianranges,butwithintheth inly vaultedJomonandChineseNeolithicsampleranges( Table5 ). Amongrecenthumans,itsvaluelieswithinallsampleranges( Table5 ). IsolatedVaultFragmentsThreeadditionalfragmentsofthetemporalsexist.Theseincludealeftfragmentpreservingtherootofzygomaticprocess,withasmall(narrow)temporalgutter(8.9mmwide),thebasalmarginofthetemporalsquama,anincompletemandibularfossawitharelativelylargearticulareminence , butlackingthepostglenoidprocessandexternalacousticmeatus.Foramenspinosumiscompleteandsitswithinanelevatedspine,andmediallytheforamenovaleispreservedinitslateralsectiononly.Thetemporosphenoidsutureisdiscernible.Themandibularfossaseemstobecompletelyhousedwithinthetemporal.Themostlycomplete,butisolated(detached),rightmastoidprocessispreserved.Ithasasmallvolume,issuperoinferiorlyshort,andwouldhavebarelyprojectedbeyondthecranialbase.Thesupramastoidcrestisveryweak.Ontherightside,thezygomaticprocessofthetemporalispreserved,andthisfragmentseemstomatchtheisolatedtemporo-occipital-sphenoidfragmentdescribedabove.Thezygomaticprocessisratherthinandgracile,butdoescurveoutwardsinawaythatsuggeststhetemporal FrontiersinEcologyandEvolution|www.frontiersin.org 6 June2016|Volume4|Article75

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Curnoeetal. DeepSkullandthePeoplingofSoutheastAsia TABLE3|Comparisonofoverallcraniallength,breadthandsh ape(numbersinbold,whereDeepSkullliesoutsideofsamplera nge). Maximumcraniallength(GOL)(mm)Maximumcranialbreadth(XCB) (mm)Cranialindex(XCB/GOL)(%) nAvgSD Range nAvgSD Range nAvgSD Range DeepSkull(c175) (c145) (c83) PLEISTOCENE/EARLYHOLOCENE PooledEast/SoutheastAsian1218411.5166–204131377.2126 –15112754.2 70–81 EastAsian818112.1166–20481386.1131–1488774.1 70–81 SoutheastAsian41908.3 181–200 51369.4126–1514722.5 70–76 Australian441949.0 176–220 71389.8126–1517693.0 65–74 RECENTPOPULATIONS Eskimo1081856.5173–2061081334.7 120–142 108723.0 64–79 Buriat1091777.7158–1941091526.8138–167109863.777–94Ainu861857.6168–201861404.9130–15686762.3 70–82 NorthernJapan871798.5157–195871395.9124–15287783.47 1–88 SouthernJapan911777.1162–192911364.9125–15291773.57 0–89 Anyang421814.3171–190421395.4127–15142773.569–84Hainan831746.6160–189831374.8125–14983793.670–88Atayal471746.9161–190471344.7 126–142 47773.170–84 Philippines501776.9162–192501405.6126–15250794.067– 85 Aeta471687.3146–182471416.1118–15147842.978–90Dayak821765.4166–189801386.0123–15280784.270–87Java2031727.5155–2002031425.5128–163203834.271–95AndamanIslands751655.3153–175791354.8122–14675822.87 4–89 Tolai1101796.9164–1961101294.4 120–139 110722.8 65–80 Australian1011867.5169–2011011305.4 117–144 101702.5 63–77 Tasmanian871826.9164–203871366.1118–14987752.7 68–80 musclesmayhavebeenquitewelldeveloped(somewhataringarches). Thebasioccipitalclivusissmall,beinganteroposteriorlys hort andmediolaterallynarrow.Thepharyngealtubercleisbarelydiscernible.Thisfragmentpreservestheanteriormarginof the foramenmagnumwhich,judgingbyitscurvature,wouldhavebeennarrow.Thesphenoidalsinusisnotpreserved.Therightpetrousisstronglyangledsagittally,whiletheleftpetrous retains thebrokenbaseofasmallstyloidprocess.Thestylomastoidforamenislocatedlateralandslightlyposteriortotheproce ss. Thepetrouscrestisverytall,sharpandlaterallyextensive ,passing tomeetthebaseoftheacousticpart.Itissuperoinferiorlyta lland anteroposteriorlyverynarrow.Theopeningofthecarotidcan al iswellpreservedwhilethecochlearaquaductislarge.Inter nally, theregionaroundandincludingtheinternalacousticmeatu s andpertouspyramidarewellpreserved.Thearcuateeminenceisverydistinct.Ontherightside,thebaseofthemastoidpro cess ispreserved(mastoidpr.brokenaway);withthedigastricfos sa preservedanteriorly.Theexternalacousticmeatushasbrok en awayfromthestyloidprocesslaterally.Thecarotidcanalis clear. TheooroftherightmandibularfossaoftheDeepSkull isadeeprectangle,beingboundedonthreesides.Ithasbeenmodiedprobablyasresultofosteoarthritis,anobservatio n supportedbyapparentpittingonitsroof.Theprincipaldimensions(notrecordedbyBrothwell)ofthemandibularfossaare:chordlength16.3/13.1mm(r/l),breadth35mm(l) , anddepthc5.0/4.0mm(r/l).Fewcomparablemeasurementsareavailableintheliterature,however,thisfeaturehasb een studiedinPleistocene/EarlyHoloceneAustraliansandtheyd ier signicantlyfromthemorphologyseenintheDeepSkull:themandibularfossaoftheNiahspecimenisstrikinglyshort(Australian:n7,Avg.29.6mm,SD2.8mm,range26–33mm)andbroad(Australian:n10,Avg.29.9mm,SD6.3mm,range19.8–38.7mm),butwithadepthsimilartothattypicallyseenamongPleistocene/EarlyHoloceneAustralians(Australian:n 11, Avg.4.8mm,SD1.5mm,range1.5–7.0mm).Thepre-glenoidplanerisessteeplyfromthearticulareminenceatanangleofc 45 degrees. Thearticulareminenceisitselflargeandwelldemarcated fromthesurroundingbone.Thepostglenoidprocessisdamaged(fragmentedandmissingitsmedialhalf),butwasevidentlys mall. Endocranially,theinternalacousticmeatusiswellpreserve d,asis theentiremedialwallofthepetrouspart.Thearcuateeminence is verydistinct,beinglargerthantheleftside.Thepetrosqua mosal surfaceisalsopreserved.Ashallowsigmoidsinusisevident . Therightsphenoidfragmentcomprisespartofthegreater wingthroughtothecarotidgroovemedially.Foramenovaleispresent.Partofthesphenoidsinusisvisible.Endocranial ly, thebonesurfaceismostlybrokenawayabovethemastoidpartexposinglargeaircellsdownintothemastoidprocess.Theinternaloccipitalprotuberanceispreservedandretainsthel eft transversesinus,whichisshallow.Thereisalumpofheavil y FrontiersinEcologyandEvolution|www.frontiersin.org 7 June2016|Volume4|Article75

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Curnoeetal. DeepSkullandthePeoplingofSoutheastAsia TABLE4|Comparisonoffrontalsizeandshape(numbersinbold,w hereDeepSkullliesoutsideofsamplerange). Maximumfrontalbreadth(XFB)(mm)Frontalshapeindex1(XFB/GO L)(%)Frontallength(GBL)(mm) nAvgSD Range nAvgSD Range nAvgSD Range DeepSkull(c118) (c67) c109 PLEISTOCENE/EARLYHOLOCENE PooledEast/SoutheastAsian51217.0112–12912737.059–81– ––– EastAsian31262.0 125–129 8774.170–811109–– SoutheastAsian2113.5– 112–115 4666.359–73–––– Australian51117.0105–1215583.8 53–61 421167.297–134 RECENTPOPULATIONS Eskimo1081114.3100–122108602.655–67––––Buriat1091245.9112–145109703.363–78––––Ainu861184.7105–12786642.159–69––––NorthernJapan871145.6102–12787643.057–73––––SouthernJapan911134.7102–12291642.958–73––––Chinese––––––––651075.790–120Anyang421155.0104–12742643.257–70––––Hainan831135.2101–12783653.358–74––––Atayal471124.5101–12247642.260–69––––Philippines501154.7104–12550653.357–72––––Dayak71138.6100–1257626.354–6871069.197–119AndamanIslands701085.197–12070662.561–72––––Tolai1101084.097–119106612.554–68––––Australian961084.396–119101582.3 53–64 1001064.294–119 Tasmanian951105.495–12287612.753–67–––– TABLE5|Vaultthicknesscompared(numbersinbold,whereDeep Skullliesoutsideofsamplerange). Bregma(mm) ParietalEminence(mm) nAvgSD Range nAvgSD Range DeepSkull 4.4 4.5 PLEISTOCENE/HOLOCENE PooledEast/SoutheastAsian89.72.7 6.0–12.5 –––– EastAsian710.13.0 6.0–12.5 37.31.0 6.5–8.5 SoutheastAsian18.0––––––Australian3510.22.3 6.1–17.2 307.82.1 5.3–14.0 Jomon448.41.63.7–13.1476.41.52.0–10.8ChineseNeolithic1767.61.6 4.7–13.8 1895.81.33.4–9.5 RECENTPOPULATIONS Japanese1526.21.22.5–9.91525.11.21.5–8.6Chinese737.01.44.0–10.6626.41.44.0–10.0MurrayValley1247.71.54.0–12.61106.41.43.2–10.8 gluedbonecomprisingmultiplefragmentsofcancellousbonesittingabove,withintheoccipitalsulcus,whichisclearlyn ot initsanatomicallycorrectlocation.Thecerebralfossaea re small.Thetemporalsquamafragmentsportstwoimpressionsofbloodvessels(middlemeningealgrooves),whicharemodera tely incised. Threefurtheroccipitalfragmentsremain:therstisfrom theleftsideandcontainstheoccipitalcondyleand,unusual ly, abifurcatedhypoglossalcanal(thickandmediolaterally continuousbonyseptumbifurcatingthecanal).Thecondylei s 26mminanteroposteriorlength,and12mmatitsmaximummediolateralwidth.Thus,itislongandslender.Thesecondfragmentcomprisestheposterior,leftlateral,marginoftheforamenmagnumandnuchalplanewithinternaloccipitalcrest . Itshouldmatchtheleftoccipitalfragmentwithcondyle,butnoobviousarticulationisobservable,withinterveningbo ne missing.Estimatedmaximumwidthoftheforamenmagnumisc20mm.Thethirdfragmentisfromtherightsideandcomprises FrontiersinEcologyandEvolution|www.frontiersin.org 8 June2016|Volume4|Article75

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Curnoeetal. DeepSkullandthePeoplingofSoutheastAsia theanteriormarginoftheforamenmagnumtothemediansagittalplane(includingbasion)andanincompleteoccipitalcondyle.Thehypoglossalcanalisasingleandrelativelysma ll foramen.ZygomaticArightzygomaticfragmentexistsanditcomprisesthefronta l process,partialtemporalprocessandmostofthebody.Theinferiorrimisbrokenawayanteriorlysothatneitherthemassetericattachmentnorthetuberclearepreserved.Thisfragmentdoesnotarticulatewiththecalvariainanyobviou sway. Itmaytontothemaxilla,butagainanypossiblecontactsareunclear.Overall,thezygomaticissuperoinferiorlyshort, butitis surprisinglyrobustwithathick(anteroposteriorlylong)la teral orbitalpillar(frontalprocess),combinedwithwhatmusthav e beenathintemporalprocess.Thesuperoinferiorheightofthezygomatic,fromzygomatico-frontalsutureverticallydow ntothe inferiormargin,is32mm.MaxillaThemaxillaryfragmentcomprisesmuchofthebody;withleftfrontalprocess,medialinferiorrimoftheleftorbitandmuc h oftheinferiororbitalrimrightside;rightzygomaticproce ss; inferiorpiriform(nasal)apertureandmuchoftheoorofthecavity;alveolarprocess;roofofthepalatewithpalatineproces ses andincompleteperpendicularplatesofethmoid;foraminalikel y preservedbutobscuredbyshallac;andrightI 1 -P 2 alveoli,right M 1 -M 2 crowns,andonthelefttheI 1 -P 1 andpartialP 2 alveoli ( Figures1G,I ). Theanteriormarginofthezygomaticprocessofthemaxilla arisesfromaboveP 2 /M 1 .Thelowerandmid-faceareatwhile thecaninefossaistraceonly.Minimumcheekheight(WMH)intheDeepSkullisestimatedtobeshortatc20.0mm(dimensionnotrecordedbyBrothwell).Unfortunately,onlydataemployin g Martin's48(d)forEast/SoutheastAsianandancientAustra lians areavailableintheliteraturelimitingourcomparisons.Stil l, WMHisshortbyrecenthumanstandards,beingidenticaltoaveragevaluesfortheAustralianandTasmaniansamples,andwellwithintherangeofmostrecenthumansamples( Table6 ). Theleftlateralnasalmargin/zygomaticprocesshasnotbeen ttedcorrectly.Ithasbeenrotatedtoofarmediallyandsho uld bepositionedinamoreuprightposition(vertically).Onlytheanteriormarginsoftheorbitaloorsarepreserved.Theentryintothenasalcavityisagentleslope.Abroad,atandbidanteriornasalspinesitsjustoutsideof(anteriorto)thenasalaperture.Itisatandrelativelyindistinctfromthenasalborder.Ontherightside,therearetwocreststhatbeginatthecorneroftheaperture,onecoursinganteromediallytothenasalspine(spinalcrest),moreorlesshorizontallyanddemarcatingtheentrancetothecavityasasmallsharpcrest;theothercoursinginferomediallyontoth e nasoalveolarclivustotheinferiorpartoftheanteriornasa l spine(turbinalcrest).Ashallowandnarrowsillsitsbetwee n them. TABLE6|Comparisonofmaxillarydimensions(numbersinbold,wh ereDeepSkullliesoutsideofsamplerange). CheekHeight(WMH)(mm) NasalBreadth(NLB)(mm) nAvgSD Range nAvgSD Range DeepSkull c20 29 PLEISTOCENE/EARLYHOLOCENE PooledEast/SoutheastAsian––––11282.024–32EastAsian––––5273.024–32SoutheastAsian––––6281.026–30Australian––––36292.618–32 RECENTPOPULATIONS Eskimo108252.419–31108241.620–29Buriat109282.5 23–35 109282.122–34 Ainu 86232.419–3186272.123–32 NorthernJapan87232.518–3087261.821–32SouthernJapan91232.816–3091261.721–31Anyang42272.1 21–30 42282.123–33 Hainan83252.019–2983272.121–31Atayal47212.316–2747262.023–31Philippines50232.315–2850281.626–32Aeta ––––75272.121–31 Dayak––––77252.913–34Java ––––203271.723–32 AndamanIslands70191.816–2370241.621–29Tolai110222.414–28110271.923–32Australian101202.316–27101271.823–34Tasmanian87202.016–2587282.124–34 FrontiersinEcologyandEvolution|www.frontiersin.org 9 June2016|Volume4|Article75

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Curnoeetal. DeepSkullandthePeoplingofSoutheastAsia TABLE7|Molarcrownbuccolingualdiameterscompared. M 1 (mm) M 2 (mm) nAvgSD Range nAvgSD Range DeepSkull c12.3 12.4 PLEISTOCENE/EARLYHOLOCENE EastAsian1811.40.710.2–12.8911.41.010.2–13.4SoutheastAsian1712.40.810.0–14.91712.70.610.8–14.5HoloceneMalaysian1012.40.411.1–13.81112.60.411.3–13 .9 Australian2913.40.712.1–15.13013.50.812.0–15.3 RECENTPOPULATIONS Amami-OkinawaIslands2411.90.510.4–13.41912.10.610.3– 13.9 Chinese25111.30.69.3–12.824811.40.79.1–13.8Philippines1712.10.510.6–13.61811.80.510.3–13.3Luzon“Negritos”2012.00.610.2–13.81811.00.69.2–12.8AndamanIslands2512.10.510.6–13.62412.00.89.6–14.4Laotians2911.60.510.1–13.12711.60.79.5–13.7Vietnamese3411.70.610.2–13.22911.70.79.6–13.8Malay4612.00.610.2–13.84411.80.89.4–14.2Dayak3611.90.79.8–14.03511.90.99.2–14.6LesserSunda/Java1012.60.410.9–13.41011.90.610.1–13.7Sumatra2311.90.610.1–13.72011.70.69.9–13.5Australian4712.80.710.7–14.94813.00.910.3–15.7 ThepiriformapertureoftheDeepSkullisbroadat29mm ( < 3%dierenttoBrothwell'sestimate)( Table6 ).Thisvaluesits withintherangeofallcomparativesamplesandisidenticaltothePleistocene/EarlyHoloceneAustralianaverage( Table6 ). Thenasoalveolarheight(nasospinale-prosthion)is13.3mmi n theDeepSkull,whichisshortcomparedwithsamplesofPleistocene/EarlyHoloceneAustralians(n32,Avg.20.2mm ,SD 3.1mm,range13–27mm),andtheLatePleistoceneChineseZhoukoudianUpperCave101(18mm)andLiujiang(20mm)crania.However,itdoesliewithintherangeofNeolithicCh inese (n178,Avg.20.0mm,SD2.7mm,range12–28mm)andrecentChinese(n63,Avg.19.3mm,SD2.5mm,range12–26mm)samples(dimensionnotrecordedbyBrothwell). Theanteriormarginofthedentalarcadeisevenlyarched andprojectsonlyslightlyforwardofthefacewiththeanteri or toothrootssittingforwardofalineprojectedinferiorlyfro m theanteriornasalmargin(i.e.,lowalveolarprognathism)( Figures1G,H,J,K ).Palatelengthandbreadthcannotbe accuratelyestimated,butpalateheightisreconstructedto be c11mmattheleveloftheM 1 s(nottakenbyBrothwell). Thus,thepalateoftheDeepSkullisshallowcomparedwithGuaGunung(16mm)andMohKiew(17mm),butiswithintherangeofPleistocene/EarlyHoloceneAustralians(n10, Avg. 15mm,SD5.8mm,Rangec7–23mm)andsimilartocraniafromZhoukoudianUpperCave(9–13mm)andLiujiang(9.5mm).ItsvalueisalsosimilartosampleaverageforrecentJapanese(n7 1, Avg.11.4mm,SD2.7mm)andAustralians(n73,Avg.11.5mm,SD2.1mm).TheDeepSkullalsosportsapalatinetorus. DentitionTheocclusalsurfaceoftheM 1 crownisheavilyworn;well intothedentinelingually;whilebuccallyitshowswearintothedentineexposedas“eyes”intheenamelovercuspapices( Figures1G–I ).TheM 2 lacksinterstitialweardistallysuggesting themissingM 3 shadnoterupted(seeabove).Itshowsadvanced wearonitscrown,especiallythebuccalhalfwhereithaswornwellintothedentine,seenasamesiodistallylonggroo ve. Applying Smith(1984) schemeresultsinestimatesofvery advancedwearforbothteeth:Stages5/6(seealso,above). Table7 listsandcomparesbuccolingual(BL)dimensions forthemaxillarymolars.TheDeepSkull'sM 1 crownis moderateinsize(c12.3mm;identicaltoBrothwell'sestimat e), beingclosesttotheaverageforPleistocene/EarlyHoloceneSoutheastAsianandHoloceneMalaysiancrowns,smallertha n Australians,butwellwithintherangeofallothersamples( Table7 ).Amongrecenthumansamples,theDeepSkullM 1 crownismostsimilartosouthernSoutheastAsiansamplesandtheAndamanIslands,butalsowellwithintherangeofallothersamples( Table7 ).Thus,itshowsnoclearanity withanyofthesamplesgivenin Table7 .TheDeepSkull M 2 crownBL(12.4mm;4.5%dierenttoBrothwell'sestimate) isagainmoderateinsize,beingclosesttotheaverageforPleistocene/EarlyHoloceneSoutheastAsianandHolocene(Peninsula)Malaysiansamples,butsmallerthanAustralian s ( Table7 ).ItsvaluesitswithintherangeofallPleistocene/early Holoceneandrecentsamplesandagainshowsnoclearanities( Table7 ). FrontiersinEcologyandEvolution|www.frontiersin.org 10 June2016|Volume4|Article75

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Curnoeetal. DeepSkullandthePeoplingofSoutheastAsia FIGURE2|Resultsofmultivariateanalysisof18continuousa nd discretevariables.(A) Objectplotfromprincipalcoordinatesanalysis (minimumspanningtreeindicated)and (B) Dendrogramresultingfrom UPGMAclusteranalysis(bootstrapscoresfrom100,000repl icates). MultivariateAnalysisIn Figure2 ,wepresenttheresultsofmultivariateanalyses using18variablesandthreePleistocenesamples.Theanalyses usedallofthevariablesin Table8 ,exceptthatvariables2 and4,andvariables6and8,werecombined.Alldataforcontinuousvariablesweretakenfrom Tables3 – 7 (seealso, MaterialsandMethods).Theresultsofprincipalcoordinatesanalysis( Figure2A )highlightedadistinctionbetweentheDeep Skullandallothersamples,although,theminimumspanningtreeshowedittobeclosesttoPleistoceneEastAsians;althoug h, notespeciallyclosetothem.ThePleistocene/EarlyHoloceneAustralianandPleistoceneSoutheastAsiansampleswerealsoshowntobepheneticallyquiteclose.AdendrogramderivingfromUPGMAclusteranalysisconrmedtheresultsofprincipalcoordinatesanalysis,withtheDeepSkullbranchingseparate lyto aclustercontainingEastAsian,SoutheastAsian,andAustr alian samples( Figure2B ).Furthermore,theSoutheastAsianand AustraliansamplesformedaclusterseparatetoPleistoceneEastAsians,withthisclusterreceivingstrongbootstrapsu pport (90%).DISCUSSIONAnumberofresearchershavedismissedtheDeepSkullasprovidinglittleinformationofrelevancetoreconstructing the originsofAMHinSoutheastAsiaonaccountofitspoorpreservation(e.g., KammingaandWright,1988 ).Wedisagree, andwhileitisincomplete,anditscalvariapossessespost-burialdistortion,carefulexaminationofthespecimenreve alsthe presenceofanumberofwell-preservedmorphologicalfeaturesrelevanttoassessingitanities.Theimportanceofthespeci men stemsfromthefactthattheDeepSkullisthemostcompleteLat e PleistocenespecimenrecoveredfromtheNiahCavescomplexandtheoldestAMHtoberecoveredfromislandSoutheastAsia( Barkeretal.,2013;HuntandBarker,2014 ).Therefore, tooverlookitonaccountofitsincompletepreservation,thecasewithsomanyancientremainsglobally,wouldbeimprudent. OurstudiesoftheDeepSkullsuggestthatitmostlikely representsanadultofadvancedage,andisprobablyalsofemale.Thendingsareinbroadagreementwiththepublishedabstractof Birdsell(1978) ,whichalsosuggestedthattheNiah individualwasanadult,butslightlyyoungerataround20–30yearsofageatdeath.Thisisimportantbecause Brothwell's (1960) diagnosisoftheNiahcraniumasrepresentingapossible adolescentprecludedhimandothersfromundertakingamorecomprehensiveassessmentofitsanities.Moreover,theadul t statusofapartialfemurandtibiarecoveredbytheHarrisson s alongwiththeDeepSkull( Harrisson,1967 )addsweighttoour diagnosis( KrigbaumandDatan,2005 ). In Table8 welistandcompare20morphologicalfeatures preservedontheDeepSkullwiththetypicalconditionseenamongstLatePleistocene/EarlyHoloceneremainsfromSoutheastAsia,EastAsiaandAustralia.Giventheoverallfragmentarystateofthespecimen,thisisasurprisinglycomprehensivecatalog.Ofthe20statesshown,theNiahcraniumsharesonlythreewithAustralians,vewithSouthe ast Asians(oneunknown)andsixwithcontinentalEastAsians(oneunknown).Thus,whencomparedwiththesesamplesit FrontiersinEcologyandEvolution|www.frontiersin.org 11 June2016|Volume4|Article75

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Curnoeetal. DeepSkullandthePeoplingofSoutheastAsia TABLE8|ComparisonofsalientfeaturesoftheDeepSkullwithmo dalstatesinPleistocene/EarlyHolocenesamples a . DeepSkull AustralianSoutheastAsianEastAsian 1.Vaultoverallshort b ––– 2.Maximumcranialbreadthlocatedhighonparietals ––– 3.Frontalsagittalridgeabsent +–+ 4.Excessivewidthacrosstheparietals,overhangingtempo rals ––– 5.Vaultrelativelyshort&wide(?brachycephalic) b ––– 6.Frontalabsolutelynarrow b +–– 7.Frontalnarrowrelativetocraniallength b –+– 8.Limitedpost-orbitalconstrictionoffrontal ––– 9.Poorlydevelopedglabella ––– 10.Superciliaryarchespoorlydeveloped ––– 11.Supraorbitaltrigonespoorlydeveloped –+– 12.Strongfrontalandparietalbossing ––– 13.Weakorabsentcresting,especiallyweaksupramastoidc restsandweak/absentoccipitalcrests––– 14.Thinvaultbones(particularlyat bregma ) ––– 15.Absenceofwell-developedoccipitalbun ––+ 16.Shortmandibularfossa –?? 17.Nasoalveolarclivusshortandverticallyoriented(low prognathism) –++ 18.Broadpiriformaperture +++ 19.Flatmid-face(nasalregion) ––+ 20.Moderatemolarsize(buccolingualdimension) –++ a Key:“–”absent,“+”present,and“?”missing/notpreserved. b Observationbasedinpartonreconstructedmeasurement(s)oftheDeep Skull. hasfewfeaturesincommonwiththeLatePleistocene/EarlyHoloceneinhabitantsofthisbroadregion.Theresultsofou r multivariateanalysesemploying18ofthesetraitsconrmth e distinctivenessoftheDeepSkull.Theyalsoshowtheindivi dual tobepheneticallyslightlyclosertoPleistoceneEastAsian sthan toSoutheastAsiansorAustralians. OneissuethatarisesfromthediagnosisoftheDeep Skullasfemaleisthesexuallydimorphicnatureoftheexpressionofmanyofitsmorphologicaltraits.Thisisnot,ofcourse,exclusivetotheDeepSkull,butitmightexplaintosomedegreetherelativegracilityofthespecimen.ArelateddicultyincomparingancientmaterialfromacrossAustraliaandSoutheast/EastAsiaisthatmuchofitseemstobemale,sosexcannotbeadequatelyaccountedforinanyevaluationofitanities.However,byincludingsex-balancedsamplesfromrecentpopulationsinmorphologicalandunivariatecomparisonstheseconcernswereamelioratedtoacertainextentinourstudy.Indoingso,wefurthergainedtheoverallimpressionthattheDeepSkullshowsstrongestanitiestorecentEastAsian(sometimesSouthea st Asian)populationsratherthantoNewGuinean,AustralianorTasmaniansamples.AmongallofthemeasurementswehavecomparedhereonlytwoofthemarecharacterizedbystrongerresemblancestoAustraliansand/orTasmaniansthantoothe r groups. ThisisfurtheremphasizedwhencomparingtheDeep SkullwitharecentIbancraniumandPleistocenesouthernChineseandAustralianremains( Figure3 ).Noteworthyare itsstronglyarchedvaultwithprominentbossing,maximumcranialbreadthlocatedhighontheparietals,absenceofa frontalsagittalridge,excessivewidthacrosstheparietal s, whichoverhangthetemporals,avaultthatisrelativelyshortandwide,withlimitedpost-orbitalconstriction,aweakglabella,poorlydevelopedsuperciliaryridgesandsupraorbita l trigones,weakorabsentcrestingespeciallysupramastoidcre sts andweak/absentoccipitalcrests,absenceofawell-developedoccipitalbun,nasoalveolarclivusthatisshortandvertical ly orientedindicatinglowalveolarprognathism,andaatmid-face. Suchaconclusionisfurtherconrmedbycomparisonsof theDeepSkull'smorphologywiththedetailedcharacterizat ion ofIndigenousAustraliancraniaprovidedby Larnachand Macintosh(1966,1970) . InlightofthegracilityoftheDeepSkullanditsanities assuggestedbyourcomparisons,theNiahindividualshouldprobablybestbeconsideredasbelongingtoapopulationbroadl y relatedtoEastAsiansthatinhabitedBorneoduringtheLatePleistocene.Theevidenceathandsuggeststherearetwomos t plausiblealternativeswithrespecttoitsanities: (1)TheNiahindividualsamplesapopulationrelated(?ancestr al) tosomeofthecontemporaryindigenouspeopleofBorneo(asproposedby Harrisson,1976 ). (2)TheDeepSkullrepresentsaNegritogroup(seealso Birdsell, 1978 ),andalthoughthistermhasconsiderableambiguity historicallyintheliterature( Endicott,2013 ),weapplyitin arestrictedsensetorefertoagracileandsmallstaturedpopulationprobablyrelatedtothepeopleinhabitingthePhilippinestodayandapparentlyalsoduringthePleistocene( Détroitetal.,2013 ). FrontiersinEcologyandEvolution|www.frontiersin.org 12 June2016|Volume4|Article75

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Curnoeetal. DeepSkullandthePeoplingofSoutheastAsia FIGURE3|DeepSkullcomparedwitharecentIban,southernChi nesePleistocene(Liujiang)andAustralianPleistocene(W LH3andWLH50)crania: leftcolumn,leftlateralview;middlecolumn,superiorview;and rightcolumn,posteriorview(NB:foreaseofcomparison,allcr aniaarescaledtothe sizeoftheDeepSkull). FrontiersinEcologyandEvolution|www.frontiersin.org 13 June2016|Volume4|Article75

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Curnoeetal. DeepSkullandthePeoplingofSoutheastAsia FIGURE4|ProposedmigrationsroutesfortheLatePleistocene peoplingofSoutheastAsiabyanatomicallymodernhumans(AMH) . (1)earliestAMH arrivefromAfricaviaSouthAsiaintosouthernChinabyatle ast80ka(largewhitearrows);(2)settlementoftheEastAsi anlandmasseventuallyleadstodispersalsinto thenorthernoceanicregionofsoutheastAsiathroughthePh ilippinesandsouthintoBorneo,reachingasfarsouthasSul awesiandeastasMaluku(yellowarrows);(3) anotherdispersalfromthemainlandviatheMalaysianpenin sulaestablishesthepopulationsofsouthernarchipelagoS outheastAsia(lightorangearrows),Australia (redarrows)andNewGuinea(darkorangearrows);and(4)wit hinSoutheastAsiaitselftheEhaplogroupseeminglyaroseb yaround30kaandwassubsequently dispersedbypeopleacrossmuchoftheregionincludingnort hintoTaiwanandeastintoNewGuineaafter 15ka(greenboxwitharrows).(NB:stardenotesNiah Cave;mapsourcedfrom GoogleEarth v7.1.51557). Bothoftheseconclusionsreceivesupportfromthepartialfemu r recoveredalongwiththeDeepSkull( Harrisson,1967 ),withits shortreconstructedlength(c370mm)anddiminutiveestima ted stature(c4.5feetorc137cm: KrigbaumandDatan,2005 ). Reconstructedstaturesitswellwithintherangeofcontempor ary ruralfemaleIban( StricklandandUlijaszek,1994 ),whilefemur lengthandpresumablystaturearewithintherangeofPhilippin e Negritos( Stock,2013 ).However,asnoevidencehasbeenfound fortheNegritooccupationofBorneo,theformerofhypothesisseemsamoreplausibleexplanationforitsanitiesatpresent. FrontiersinEcologyandEvolution|www.frontiersin.org 14 June2016|Volume4|Article75

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Curnoeetal. DeepSkullandthePeoplingofSoutheastAsia In Figure4 wesummarizethepossiblemigrationroutesfor theLatePleistocenepeoplingofSoutheastAsiabyAMHassuggestedbythemorphologicalevidenceandrecentgeneticstudiesoftheindigenouspeopleoftheregion:(1)theearliestAMHarrivedfromAfricathroughSouthAsiaandmigratedintosouthernChinabyatleast80ka(largewhitearrows);(2 ) settlementoftheEastAsianlandmasseventuallyledtodispe rsals intothenorthernoceanicregionofSoutheastAsiathrought he PhilippinesandsouthintoBorneo,perhapsreachingasfarasSulawesiandMaluku(yellowarrows);(3)anotherdispersalf rom mainlandSoutheastAsiaviatheMalaysianpeninsulaestablis hed thepopulationsofsouthernislandSoutheastAsia(lightoran ge arrows),Australia(redarrows)andNewGuinea(darkorangearrows);and(4)withinSoutheastAsiaitselftheEhaplogroupseeminglyarosebyaround30kaandsubsequentlydispersedacrossmuchoftheregionincludingeastintoNewGuineaandnorthintoTaiwanafter 15ka. WenotealsothatgeneticstudiesofrecentSoutheastAsian populationsincludingIndigenousgroupsfromtheAndamanIslands,WestMalaysiaandthePhilippines(Negritos)havedemonstratedtheircloserelationshipwithmainlandEastA sians ( Jinametal.,2012;Aghakhanianetal.,2015 ).Moreover,while theAustronesianspeakingIndigenouspeopleofBorneotodayaresuggestedtohavedescendedfromagriculturaliststhat settled theareafromTaiwanonly4–3kyr( Bellwood,1997;Xuetal., 2012 ),thishasbeenchallengedbyawiderangeofgeneticstudies coveringmtDNA,Y-chromosomeandothergenomicmarkers( Capellietal.,2001;Karafetetal.,2001;Hilletal.,2007;So ares etal.,2008,2016;Tumonggoretal.,2013;Trejautetal.,20 14 ). Instead,itseemsmorelikelythatAustronesianspeakersfro m TaiwanandislandSoutheastAsiashareacommonorigingoingbacktotheLatePleistocenewithonlyalimitedsignalofthe“Out-of-Taiwan”expansionduringtheNeolithicperiod.Itse ems reasonabletoopine,therefore,thattheAustronesianlangu ages themselvesdispersedthroughislandSoutheastAsiathroughsmall-scalemigrationandlanguageshiftbyaNeolithicore ven forager-sherpeopleratherthanlarge-scalemigrationinvo lving populationreplacement( Soaresetal.,2016 ). WeproposethattheDeepSkullrepresentstheearliest representativesofmigration“2”inourmodel( Figure4 ).The geneticmarkersofmainlandEastAsiansarealsoseenineasternIndonesia,albeitincombinationwithsomeMelanesi an haplogroups( Monaetal.,2009 ),whichprobablydispersedinto theregionsomewhatlater.Theestimatedgeologicalageoft he DeepSkullofc37ka( Barkeretal.,2013;HuntandBarker,2014 ) alsopredatesmolecularclockestimatesforthetimetheNegr itos divergedfromotherEastAsianpopulationsaround30–10ka( Jinametal.,2012 )aswellastheestimateddivergencetimefor haplogroupEof 29.7ka( Soaresetal.,2016 ).Thislendssome supportalsotoourconclusionthattheDeepSkullmaybemorecloselyrelatedtotheIndigenouspeopleofBorneoratherthan the NegritosofthePhilippines. Allofthisfurthersuggeststhatthemorphologicallyrobustl y builtLatePleistocenepeopleofSoutheastandEastAsia,widelyconsideredtorepresentapopulationrelatedtoAustral oMelanesians(e.g., Bellwood,1997;MatsumuraandHudson, 2005;Matsumuraetal.,2008;OxenhamandBuckley,2016 ), wereprobablymorerestrictedintheirgeographicdistributio n thanhasbeenproposeduntilnow.Ourstudysuggeststhetwo-layerhypothesisisunlikelytoapplytotheearliestinhabitant s ofBorneo,in-linewithgeneticresearch,andasproposedrecentlyalsoforPleistocenehumanremainsdescribedfromt he Philippines( Détroitetal.,2013 ). Ourworkalsohighlightssomemorphologicalsimilarities betweenPleistocenesouthernoceanicSoutheastAsiansandPleistocene/EarlyHoloceneAustralians,consistentwithourmigration“3”scenario( Figure4 )andsomegeneticresearch ( Monaetal.,2009 ).Inthisinstance,however,thetwolayerhypothesiswouldhaveinvolvedacomplexmixtureofpopulationsoriginallyfrommainlandEastAsiaviaBorneoandadjacentregions(migration“2”: Figure4 ),peoplepossessing haplogroupE(migration“4”: Figure4 )andpossiblypeople migratingbackfromNewGuinea,ratherthanlaterarrivingAustronesianspeakingagriculturalists.Itmightbetterbecharacterized,therefore,asacomplexmulti-layerrathert han simpletwo-layermodel,in-linewithbothfossilmorphologica l evidenceandgeneticdatafromcontemporaryindigenouspeopleoftheregion.AUTHORCONTRIBUTIONSAllauthorsconceivedoftheresearch.DCcollectedandanal yzed thedata.DCwrotethemanuscript,withcriticalinputsfromal l otherauthors.ACKNOWLEDGMENTSThisresearchwasfundedprimarilybytheAustralianResearchCouncilundergrantsFT120100168(awardedtoDC)andLP120200144(awardedtoPTandDC). 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