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A.dEtc and reliabl. mexrurmsns rqun. no.-pomus .ndoca$ mrtends, .lexn glassu{., clem(pr.rcnHy dnt cd) wt.a xnd x..uor bahncs mdwight.I! is ilwys diresing ro Mrh bubbles dis.due ro iopropd or inconplere .oatlng oi the morcpoious F.nions of th. erdocst, but .uch ar the drn,ges inhe4nt in Pat.on.uolog.

PARTIAL BRÎN ENDocAsT VoLUME5Ii is ofrei thc case ihn the avalhble .ranial ponionsm roo ;icomplete b .Iow r rtlirble bhl acon$nction of the hissing prs, xs rhe missing pRft saedwhrt n prsit, Tobias ,nd Hughd Gcc, in prticultr,Tobis, 1991) tun into tAis problem whcn anemptinA

to lird rhe ..do.nni.l tulume olOH 7, rnd HdlMy(1972) had similt poblcms in aftemptins ro 6nd tot,1cndNnnirl voluda fo! thc M,trprnsgr speiEcnsof theMLDl o..ipitalponion,ndihe56 19conialbroe portio'. In thsc sibrtions on. ries ro xs.eftinvhrt pere.hge of rhc tord rclume is rtprcs.ied in! poftion ofrn c.do.!$. In doing thG wnh a wietyoftua, one hop.r ro rivc at a perc..hge 6g!rt thrn fairl) tighdy disribubd. Tobirs's (1991) dt.usionscgrrdi.g the bipri.r bnncl olol-I 7 povide x de-bilcd de{dption ol he this done. (W tunh h ad4P.flkak.llt, trat htu wJal uarA flrl/ tr dahz ;n thiaa, bdt k t?a,t tb aaoihbk dat brr:ld4rtuE dn@wi: ,11 k nardt.ttti, 6tinak:.)

ENDOCAST VOLUME RELIABILITYIn the Volure md M(hod sedion of exch hq inthis volume k give our Gsc$m.ni ofrhe rcLxbilitI ofrgiven endo.st volunc. Tfic reliability is smrtd r.codinE to nethod (letd d$isnation) rnd ou .lduxrion ofthc sim volune (ilhcncd d4kmtion). Th. endoca* volums gikn hdc rft obhined lir on. of lourndhods: (1) difd sxqdisplacem.ni of.irhd x full or! hcmiendocx$ sirh minifri disbfrion rnd plsticenc4ondruction, (2)det trination using , pdi.l.ndo-.a* as dcdibed by Tobir 0967, 1971) rnd Holloqy(1970), (3) *ensiv. plcti.cnc Econsdction rn.o.!ing to harorthe tord .ndoc6t, and (4) volune cdd-hed liom rcgre$ion fornula or c*inrted on rhe b6i!of a fs measuftm.nG, whi.h rre rhen plqg.d intolorqdas *ch thr olferd by MacKnnor *.r. (1956):

rl = J10.5(LtrB + LWl! )l

H{e I is mdlnln lcngth, ,/ is uidrh, , n tngthtiom bftsmr b dr pGtuior limit olihe cr.bellum, ,9n the height lrom tfic scfrq to the de.pe$ prt ot th.tcmponl lob., bd/ n rhc tun spe.itic .o.6cnnqxreaes to paviously publishcd wlus rhxr nxve eiticrhen .on6rned by us or, in etuin cases, not. Inxddition to b\e endo.a* ncthod sores, erdr .ndoca*n soEd nunen.ary ro indi$r our asssfenr of rhcdj$ility oi the given vol!tu. Thc* rlirbillry vJu6m.nluated on a s.le of1 to 3 wheE 1 i.dicxks thchigLci ftlirbilrt 2 dD fic giren rclune is gtneeltalirblc, and 3 thd tl]. giren rclume hs low dirbility

Bx,rtr Enoocesr Voluuss By FoRMULAAr uitb rny physn.l objcct, th.a RE reldionships bcMcn the rclume xnd thc linc{ me,$rnen$ rhat

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HonMy RL 1970. AsE,ropirhsi'€ .ndos* (T,uDslpdmen, 1924): A i.w yoluft dd<oinatio!. s.l.ne

Hollowa),Rl-. 1972. AsE:ropith<itr. odod6, bnin @-ludotr in rhr Hofindide md a modd of honin:d .vo-lurion. rn, Tu,nc R, !d. Th. Futrniond riJ Erctdoeny Bioloey of l}inrc. chilzso, Ardinc/Athsto' Pr$,

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Holroqy RL 1976. Son piobrcms ofhoEirid bsii .ndorj* kostutioq,xoodrr md rcuin idglniatior.In: Tobiis PV, Coppcs Y' cds, Coloqu! \,I of ttr L\coryG$ Is Plus Anctns Honinid

Horlowsy RL. 1973. tlobrcms of b!ai! cDdoQ* inrrpFhiio! &d Arioa hominid 4olurion. li: Jory c, .d,Erly Hodidids of Afaci. Londoo: Duckwrh, pp 379

src of rh. hnre jl]s h cdy hominid .voludon. In:Tobir ?v, €d, Honinid Erclurioni pN, Pesri aDd futuc. N.w Yo& A.R. Liss, pp a7+2.

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Honowy RL ?00o. Bnin. ti: Ddsdd r, nftcE?r r, v.'couwnnsJ,B@ok3As,.ds,EicIlopedirofHumoErcludon md Prhisrorji 2!d id. NsyYo* Gehnd, pp 141

Honomr RL, dc L{Gr Lu)mondn MC. 1932. nnjr.ndoqn *jnndry id pdryids md hooiDidr som. prc-liminu) fitrdinss o! lh. pst.omologl of atblt domimie. An J PhJ,s And@pol 53i01-n0.

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k$rli L. 1911. Fonulr ao! dctlminsrion of rhe sprciryoath. N.gb sku[ ftun drii,] rq$r@cft. Biomdike

Jlnsn HJ 1e76. Diw$ion prP*: Pdeblurion oar2ryuzgs ,{n NY Aqd s.i 230:370 332.

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Kochdkop r"r. 1973. Prteo'ruFlog} wrhinsron,

reM!) M. 1976. Molphorosi..l ftrcbiJ rsydfr.ris ofmo,l.m mq md idDnumn pliEaEs. ,4ln NY A..d &i

IracKnio, rL, K.inedyl4 Dayi6 TV 1916. Thl 4tin.rion oash op{iq &on

'o.nrg.no1o€i. fe$cdctrr.

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scheph cwH. 1950. The bnln sss of rh. rcidy [email protected] Pr,,,rrror'$ skurL. rn, nrcom R, Robiiso' JT,s.hcpcB GwIl .,1s, Sc*fonftii ,peman, ?rJtlrutd Tnisv Ms Mcm 4. Trusr"r Musuq Prbd4 S.

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shardra TR, Maooche sL. 1969. Th. bnjn oa{hiEpoac(P,, tusrrr,r. rn, BouE GH, .4 Th. Chinpria'A'aboy. nlhrvior, .nd Dls*s ofcB.iiifoie uoiE6ity Irdk I,!c$, pp 137-237.

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Iigure 1,10, Bnin

S* 505 boin volune Gdtei thr 550 nl) n hq.rrbxn no*.hinFxnz.es, but ir wodd b. r.xson,!l. roexped thxt th. Stu 505 body sia ws .lso gr.xEi W..m quibble xb.ut these rbelute md rchnve &ain sizs,but lhe frjor point $ens cl.rly provrd: ihe hominidbnin u.dffinr ftorgmizrtion befor n unddw.nrany ndor expansion in size.

As fturc 1,10 shos, *ly honinid bmin volunsm nosdy $ ic dnoughout rhe .rcluriomry life sFxnfon th. Hxdr specinens ihrcugh rhe Plocenc gkileSouth Atican speimens G.s., Truns), .rthoush ibehns pobxbly shsvs r slght non€lloneti. incftasein bnir sia wer the .did Al 1-14 condition. Morspe.imens qil be n.esRry io denonsr,E ihis. Induded in this rppmnt shsis ofbnl. size is th. o.einomplete bmin endo.a* Rvsihble ld 2. trrti, rt.onstucted b 450 ml. The robu* ,ustalopithecines, rside bnnch .ot le?ding or contibur\nlb rJE Hamaline, rc clearly lRrg{ bnined rh.n ten 2. .tianu!cousinq but this intreN rns lik ly an allonetric o.erlatd to r lq.r bodr size. This shsis vB perhrpssome *o nilion )€aB in duntion, during whi.h it islike$ but as }tt nnprwed, that our anestal s,,,lincascus bcsinning to bnnch offro n.nner,4. rlrnana

or ,r. s.r,, fro* lilcl thc lafrcr, sivcn thc tomainingd.nbl lid coiiJ @idcn.c (AsFaw ct !., 1999). Tnmtu* au$olopithecine rcpaened * Swdtkais b'sK 1585 (Hollowqt 1972t) shos { clcr lcfr o&ipiEllsrnmci, and a pu.Leriig in thc po*dor o.cipiEllobe, su8g.sring x ftduction of primry visu codThis .rn hrdly b. rxk n ,s prooE hNft( ?.lh.Frhe .*[e$ ,9 fu lin. hld hn]ns not only lcldjng 6x dnn ic size in.rta. hut .lso showing rhc ps.l-i.l pnrtrns sseided widr h. h,nd.dnes rnd ,ls.ercbnl henisptu;c +e.i:liz{io n . ycr rh. ,4 boiaiforms, su.h xs oH 5, Konso, or KNM-ER 406, donot rppetr 6 show such cletr armddricr. Nor do tI!edier Wst L.le Tukam sp.cim..s unmbiguoDdyindnlk .irher enlrgtd or ffirgbized bnins. W. tcn-r iv.! asrt. rvith r.lker.l. {2000)ihlr

'hel.,y';arBprcfonbl lob. rppes les poinr.d rhrn in edlisrusE:lopnheci.e sp..imens, bur \l lso belicv. rh.tcaurio. is neces.rybecrus rht f.rtu is quire frirblceen lat Prn bdglalth\ nd 2 ?d,,Jra. Thet Brcci!cap regions do not rpper humm like on th. specineGcuftndy rmilxble. This $$ss rhrt wirh i.ftsinArdrpttio.s to nird envisnnens, md r gndu.l de$iccation of aore$s, mtor,l $lectio. frvorcd x &,i.

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Otbtr lo$il honinids, su.h as KNM ERtgt3 and1305, m fonklt puzzti.a, lid pshaps cD bd licy.das idv,.ced rusfr lopithecin$.rrs*mpt.sof rniniadaptive rxdixrion oaedy t4,, lbninids. Whilc it ncomnoo ro vj$ KNM ER 1313 as simpty r f.mrteKNM ER 1470 (Srin$r 1986i Wotpofi 199e), whd thrt rbe d.snc ol dinoiphism oa br,ln hlumc(51G-752 nl) lid brai. endo.in horphology doe6 notsupport ihe sul!]csiion rh* rhey nc th. sne speci.s.KNM ER 1305 n a puticullly diiiculi sp..inen bft(. we \ould h! more wili ig ,o (nrctuin rhe fKulatiw poiibilir rh Ki\i\l-IF t305 rnd KNM-ER1313 aE sdu:lly dimolphic form. oai di$eftnr qnvtara (ondvh@d ausrdopithein.) Iine, at lest tu;thc vies?oi.r .f bmin votunc, ,rtnougn th. edo.nnial norphologics m quiE dilTcrnr (Hotowry, 1973,

la ths. concbsions, dnwn fon tlle p,ts.edolog,cil rnd ortrd crrnial ,.d po$.nniRl evide..e, rccorcd, mo* ofrhe im[Frhnr fto4.nizationd chxn$swcE .omptete by bce.en 1.5 and 2.0 MYA. Ir n.r,h! fh tunher !h!ng.s 6ofi

'n !.trbBl ir mned

-jBrcc?s :nd Wtrnrktsrsion..rnd oAcrc.rcbrJ, aEbclhr G.e Wcrver 2001), and submiicil rcgions such a5rhclimbicslstent ol< place, bur sid. Fom thc ftftbel-Iun, w. cannot dekd ttuse Rith our nrcscnt n thods.

This lerB us rh a s.sidcnblc hiatos in EuoDcin l4T prtr berl* rhe Atrpumr dnd Dm,.isi ei-docs$ ?re noi

'tr d.srihed (sid. fbo stimries of

bnin rclume). It ds.laves us uirh tfic Ncrndetrh.ts(xs rcprcsented xt lecr in We*ein EurF) shdinEeddgcd cmniRl c,px.nis ind no evid.n.. oicerebi:lpnnitirncs, *c.pr in th. ph+ephdi. shape oarhecnni!. Mos enphrticdlr, w *e .o digcrcnes i. rhenoehol%yofrbe pdontal ponion ofthc fDnin bbe,r inding ongrenr uirh tlE morphon.ric an,lFis ofaookein d .r. (1999). While the Nandcnhrts nishrbxve bccn a sepmt sr.cies (in dre sns! oa. morphGpecics rnd .ot

' biological spcies ii rhe modtrn

'Mryrixn' scnse), we do not se how eirhd the b.tBviorrl orundntyinanruDtogrcJ morphotogrJ eM-

Weprefcrto agrd Nerndftluts s sepme fDm mod,si 1444.4rt'r tr oity ih( subspai6. t.vd (HouonnlqSi,hutof our,e rnrnopen ro mort lonnncinge_id.n..onthisnsu..The160i000Ft atdHcfto Honorlr,,i?ar, cmnir (whire ct al., 2003) wix cBntu,ryyield endocmnirl rcnxlns thr ft e$m6:lly th. sm;as in fr.dctn Harn nlnE G.E , size, rsynndi.i lDbr pxtrns, and perh'ps neninge,l v.$.t,

B!tsa!,roRAL DnAM jcswtit. mu.h hI brtn Gtrd isl mrd. of lhe nquitum-,t .utrrJ Jd\:n\!r beginniia r lhr up.r I{c i,thn Nirh 'rue ro,r Jalt,H!.h s btool rndnioBj c?flings, pdictsl rtj jes.ln rvicnd rcLgr s rite, le.g, *c l(jfln, 1q90, Kl.nE+q 2002, md severil p1Pcs in clw,2002reBrd d tuDLdJnsmtr5 r fic mo,r Ike]yeryt?n:dlorrhserdqn(. Dfief ihh norotogi. /b.hrqo

.baq€s in cognitive crprciries b&d on r single or nol-

.pre muhrions or rhe rcquisnion oa n$r.t hodurtr,.e., Du!hnn. *i1., 2001 I trF, rhouah, ihsdipo$ible tAd dmncs i. oft undcrtR.di.g of thc gFnetic eLielb thRi relft ro bfrin dev€lopn.nt ni&illumimr h@ the hunan biain wotkd ar rhis lfuGe. Pieu$ ct a1., 2003). Thc findinss ofrxxbo (20031shouing frgread gc.eti. di{crences in bnin gcnomicmdi.1for1taru thm chinpanzce, derly su8ge*tlEnatofll sl.*ion h.s optrri.d on rhe nflrogendiGbumao brtn cvolution. If one think xbour rhe difr+.nc6 in .ulhnl b.havior ton rh. iime oi srn Edisto the pfts.i with the u$ of.ohp[r6 rnd ou &FFdcnce oo npid *chR.g.s ofinlomation, Sxtilirs andthc likq one should b. r litd. slcptic.l about unddlFing muhrions subseni.s cosnitivc abitries ihxt .oi;nicnd] spnnginto b.ing $me 50,000ltas ,go. We b-li.E thrt w have rmained thc same spaGs, rnd @perhaps the smc specis 150.000 to 200,000 y.!s t-I6E ihat. Ifoie wcE to only thint oftbe no'pdiJEttrcmrins (i.e., sbne tools) of lhc Ausralim Aloridnavc muld $il bc hrvi.s ditiol9, undNmdii; di.ompldity ol ihc; soci.l sruoural slrftms. Wou]dnot ou gr*-g4ndfnheB, in&.d, grndfrrh.6, tmaz.d rhe obvious difertnccs in int.Istozl, n.edal, and spinbd liacgles rhrt fiavc .evotwd" F@rhcnlfetincs ro ouFwithout rnyenpiic'ldemonsd-don ofBen rn (uhiions u.ddlyinF su.h belxvioG?

Th. erli.* pirses of honinid si*ena d. p*ti.ularly open ro .pcculative enbodc.y. lui when dlis s.id rnd don , n Bmrins rh. sonc tool i.dusrisor ftditio.s rhr can inforn us thc mos rboN h6ninid cognitiE abilities. This docs nor me,. rhxi Ednqard rchaslogi.al conrdb such as the frun cnains, home brses, rhe evid.rc. (or hck olit) for tiGinponrtion owr long dishn.es of stones used ii n.t-ing &ols, de iahing cmsses, or c{n crnnibdiso.Holloway (1967, 1969, 1931) sugg.stld thr so'e rootmrkng rnd lmgu!!. mighrhrt hJd

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FUg u rurPiq uetr sFunru ruor rur:lrP rll{r rrDrddr rrla ?qs FD'u' rur:ulp,G:^ pq rlqr (s:tqqor !tr)

+a'uJo lu"eSta' ri,rualn /d!,n la lbtuanf ru,tud),? !aun,!,z-,! uu1 pn $4 p4,&o,u,op!? 3uo4,14,?, t"-|t,rt1q pr, tbrd tuthqlu) buoHtpu o ru,p,2t ter{tu}rilrlap4e, qus

I3aElS

$'as :rqr r?q rz sr58ns .a lru:si !t {566r i6l6r i5l6r .t96r) iholoH lq pas.5a.s s? u,uq nll

roJ uoDrmtdF {$qod sou :qr s{! F nlrtrtrropu: prloror )u:3 norqntr tur

^to^x uo prseq !s9!rqr trroryro@:rt piEA, ,M

'grtdr urunq to trtqrdrtr Iflrg rou :r.h sFq!trpuNrNqr

^'l P'u:Pur se!?rt r .!crt

FrFrlpur:N JI ur sraDj . D'uudJo !r1,tu. qrra P:u,quo Irrr rDUotr : \ r. lJrioJo 80qr s. iiPrrllrDll'lP sqj sl$o runoroid Plnolsl?tl!trpurxN rrqr onF I'qr puD .(otl d),,qr:tr& iqt

6uzqrrn-\ lvnrNrD rnus \1\\.,ri Nil rNrn ,ll

290 soNr- GrrER,\r . Nor Es

behrvioa ora h.ightcned thresbold torvitfiinlroup rg-gft$ion, p€rnining Bnups to liv. noE dcnscly Rithmoa .oopentiR behrvior as bener potccrion again$both pEd.tor rnd otiq lrcminid goup5. Concomibndy, homonil hrg€t tissu. chrng5 allccEd gbfrhnfts {lonsd dumtiom of srffih and pblons.d &-pend.ng) po$ib! uith sfrc illomctic in.e* inbnin srcnh thrt wouu navc tcci rcflectcd in sLgndyhishcr EQ ihrn in aFjs. Th.n bipcd'ilnn no$ pbtd-blr allowcd for gllatd dn!!s ofconicl'c dicGiq andc+lo tion, lnd th& |roljng sophisiiotion about ob-jcss .nd $atid (latio.ships qs prohlbly insnmen-bl in sdcaion lor ! (htvc d!.tion in ?vC md r(hdvc c{ansion ofpo$dior Friebl lobe. E*.nsionof loastlt rnd menory wodd obviously hrre beenadqntlgcous, fd .pp.! b s b be msc..r in tl1cshominids g.(n rhcn snc*fid bddd prcaront l lobesr.d dpxnd.d hnposl lobes.

Lakr runi.lopnheci.es pnbxbly s.r usi.g toolsbur wer noi slbwing cltr shndrdizrtion oflorm ei-thd in 6eir cnoia ofrmls or then nmnfrce, the.r!*t evidena lor $one rmls gDing b.ck to ,pprdimdely 2.6 IIYA. Il so.i,l communicatory skills uge3€1td thn sdge rhrn wh* ve vitn $ in dd.ichirprnz.s, rhis muld nor likly have been ssociried uirh soci,l aFect rnd contol than with hunting

Ldn art Aom Rrtt Hano {actus ?brn, ,-ftatuiry m"eld.tb af tuEN I ad th| d.uk?no' dhbaua'aP.ni.s;.h.rutndndn b'nt lit ,n brula"nmti6, it hdks RrcMi M?,qia6 afthr t .fnntulntua ad bdh allandtn and

','.ttorni iN66 aJ

ttuin @|uft, uith atkdiry inw ia EQ.TXk sbsc iocludd clabo.ltion rd Ngmcibtion

of ttu changer in so.i.l beidvior hentioied ii $rge 1,More importrnn ir invohd 1 gowing dcpcndcnc. onsociil .oh$ion, [email protected]!ioi, lid sd{ol. ofplenen-hrion in dl eononi. hs!5 i.volving b*in.i.g lr-gl1ge bGed on albitdy s]nbols, however !;ni6v.,rnd ihe hanuFa.bring of sbn. tools to *md.idizedpntens $it[ cLr oSrnive asmirdons beMen.M-ftrni tools rnd hskr. Biped,l lo.onotion ]A f yhudm. The bnin *pmded in sik bot[ absolnielymd rthrively, rtn.ding dor l]ordoml{rgeFdsu.inr.sdio.s in the nodtrn hunm &.crion. Ths..hrqrs l.d ro furL\ei rductio.s in siz dimorphisrb.tue. dr sdes with x posible incrclse i. seon&rysm.l diddpl]isn, pl]siolo3ic,nt, mrtanic,I]. md

b.hamRll) The. ,n tum p.e,lcd ,h. br.n rdserns poshrn dcp{ndu.) ;nd hmins.incrc6iig. t(Mrd prtunhon, ind ,hc i4r{d dudiion.h'ldhood to *"u! mdurJ SduJ krninaormaLng hunhng, rdllcr ig,tu(ngrig,rnd rcpnve stnllis

-e( il1 in r posinve fe.dbrk sls

bcsccn culbril conFldity rnd bnln eiftJg.mbut.ily r. ,ht drn, rhr br . ob{.ri.J .onsmiwca iot trorhcd. As Holexy noEd in this

L!igttrs.bdruorbnm morqmns! &r op.d@sini{ bchrior ofrmqc ncrl} humr ryFoped, *heft lxnguge xnd idl mrkirg m$ lilm

rco t,itorca n ,h. +rsr. rnJ rinsrisr hid pdinponxie ii nxiMining so.ixl.ohsion rtrd.ondd

ing indr*d rnd th.r ms rorc {(ss ji nr&igrnd huniins lq€! samc Th?rc wa' a rlcdion ror irGsld bodr siz., bip.dll lgiljlr aid prdi.riv. xbiliridro' moE suasstul hunri's (o€roioc th. 6r[ food qdin lhich rcmq and cbildici dr idpddnr bndibFtun). The {.irl b.hxvioin cnrngs o in.d in ses6

siscni hudiiig rid lor piot.dod ofr nor $dF hlnbs. |o! f!o,]$ and Fu!g. {ho wE Plwiding smdk

samc ud ycsdiblcs. Thl "iritid kj*" dr "huhr *olurion' n nLly d Ntrd l..ds b slg. 3.

Cnukg Nhbltatkn aJurrrul :Ailt basd tui,, dlaaglqa airy tuitrury t-b,l :r5knt ard d.nhfi,tt b a ryh an n Eo kE f a : n i! | fadna& E b t i a n'bil bei Mb.h@i. l e,flniry and b.ain eah.gnr"i; al:a n,tir-liry r6a.Mk k t.ai,?kri 4!nn4&t ad tdi-, f b, n :p, i o t ad ;n fo. ! nrc t 2 t i t, a {bat, a n d, o,ioldt

stos.3 n $ 'nA

w b.tifla fiarab,inn {Naa-,.dral. It *.ms to us unlilcly thli nrjor.hr8€ i!po$mbl dep.nde.cy6ne, g.$nion, prtuitio., orGpodutiE age oc.ucd du;ng thn sbgei hving b.6for tlll ho* pd .onpl.bd in iJ.lui. ,qoza

'al;d(,.l. dl/?/r?a{a;) bd pft-Ner.dcrthllen tid*, 6$en ar Lx$ in We$tr. Euopm N.rddth,ls ofrh.'tlssic' sod. We :lso believ. tl] it wa r shge du;rgslnch in a$.d Lr.inA Nodd lrNe ben *ftn lyinporhnt ,nd u.dtr frn\' stingenr soci,r conMl,as cultu.l conplexities wft o(udng in noft com-plex md stinDlating mrtd,l rnd soci.l envi6nnen6.Tne mjor neunlogic.r drmgs wrt piobrbLy ni.d

l1tt u:rNrq rlru:rteslP rP \ .qr u 5 Pneq'u'rq plnoqs 9351 )ts to lPos ruTutrd.Pu, 'rFrqnduI rP,umoq s:q!o lsropulqr Jo lSotoqdroupur s.unto^ rqr qroq Jo IPD5 rurPu.drPu .trnotru: pForts IFJ nro puD (H-u) sn Jo euo u$N.q

suqrP.qI sPru .q pFotr r.qr $ro lsroprr :tqssod PUE tPug m8J uroErrd istru3qr lo uoruru iu urs rou }rtl r |uurq P.qu$eP i6I !S qrk Prrmo$ uorrod hFtr FloP tql

'35 !S uo ,rPas !$rg Jo dFiaatrP trq] .urlPun* truDlqrrdoFnsm UDBY qrnos.qr Suouv

Ipnr DqxnJ rJ $rtrr r,utd puistsot rD rDopu: or 'otrqr uo +oa or nuucrp .!oqlro qr$ot rFJo e9n\r u, soqr Sul5.norurJo .doq:qru

^spq urD uoru:o tra pu? lq8noDqr :q o rdu$.

no r srsNrnt SunF.uos rDqr rqr !!ru! rrJ .qI,{pDs nrluryJo Il]rrm s !tr,fqo :sqr prr.tr rqtusrqnsr nqro rr$ rooq s:uoq r'p$prt prprus lpFlduor $,{F!r'rgns u$q aiq

ro! s, iFruriod Br FNr

rsrrrPlt tqrjo IF tu9!r1orronrs ag orra srsvrooNg

'rq (rE r?l]1 !P tuu4qtuJ 4nutlnJrtrpuoa irrgc inojo Fnpdd PU' rurEosu, r91$ ,(tucu.r pu? '&alll.d ,ttuoqqo^r hojo rnri PuiDnpod trqr $oq ! uruq upuD{ r,ll uoslttddtqrJoqtrnu urtdF Ilqrgorl lrr stu"liqqitiu suoD4nurenFs a.J r uo lutluidiP uo!!r u' atl D rou Pur,pro erxl lollu D'i {q$od su dop:p nEuoq

-nto^. Su.IPuw-tuot e s! rourqrq ucun'l rl!1 hrNo'l!n rqhu.r lroP q.crPu: ro lPrus no \sruuru'q

Jo rDrd tu, lrd { rptsuotr * sqnqDr piohr!:qlqld !^4i9a ro 'itd'uun'j 5runP luos iuoDrPlr

plp uoq?.xrdd? iqrqn 'uq"u

loor ruoF't$dd!tu.$ard luoqoudot

lep:drq !8uuoll]r sE rPff 3!oq

-.rdipr prour,lr,l pi'urduorr reqr eturqr lBrsotoi-niu ras rtl tu4rrulr+ 4t sr|']ru:bgns r ipqsIr$ Jo a4runL 3'qr ! i:uoP rou rs qqroPu!rlrruor lrohEqrq:nqr prs Frraolo$ u.Dllp lr]rh*Ddo !i^r)!drqr $r!rd l4:F:jsorDsnuJo suoqEt

fru.r nm! Nll PUt lldrum Jorou fuE luoqu.u

q' u' osP !'q sdrrtsuoDqr Pr.Frtsolnsm u:guEq: ritduE rou spltr uflldnorodo:ot u.Susqr v sunmu ptrBoFnr I (l:qr pp ru) .rrrdo uE: s!.uoduor rsrlrjo :!oN rtsntq ro br:lqo 3!r,6rr

'd'l' uornrd Jo ruru:lgr :r| uolouorot pp:drqr 9r !P'L'uo,j rqr Jo qu:loduotr PiSotoq&ourrlro d' ,rtuiqr Auh.UoJ r|td toolBqr Suqr.us

$nf pu tur uoqnpr prrrdort i'a D:d$ Frrrr rnrjoou, u? ur4 sr[|t 3Bq u'rq uEunq:qrlo uolqon:.qr Iqr lriEd rlrql u' {o'ls or Podu4s r sq .l

isprrr0oq!o$u 5z ruoc Jo rohE,i:'] PUr sur4 r'{r rn$aupw .lrsqo or cn Jo rdo, Jo :!o rj4 rr flnoa SuoIeq tu'{r.u iuq r

'}k ur3 suuq rno ntDrrd

u, Pui .uo{nlod

P'uruoq :'txi:P nr ra lFsottr aorl

Jo ruiu$*e rno ur 1qury :q ol DJ.id :a rnq ]I!qo! ud{ i,tl@ uEr $Dlans q:nu rsr{ qqN+Burr' r' PUr't )uo trql uo rur?trP .I!I rr"u PFo{(uoldun$r no iuE ur :rr:Jrp .pNru8ro ,o rrprosv i96r00.0 D,r00m0 0 sr p+rEs arq *rEqr flft0,!Fu'uog t$$odJo :tulrrd :!I $sB:opua 00IJordursr rj tra lutrsB ruoDrtnrp: mo Irqdun olrnfiFaldr !dq trqr

^sq rnoqs tu

!$tr p'urroPuiJo ducs lEPiu:eld ino r h€ sqrrqr\ 3pruru.q pnpb,pu! 000.000 9u:q o rno sruold.liDuit id sno, 0o9 su q srolrDur' 00o os iruo1!'lrjruudp pr!l rê'j ptnod sp'rpNpur IIBU doq i(rn

t! tqqbptr ,{F r !e lrFs o) snr,{ 00o omJ NdIr Jo rnno: r']r Dro 'uds Puoor.u:8 luu turnpsFos tu,trnfddr oos ro rzu (rM) looqndod Frussl&!koi r pri lxJ OZ .{Es lo llds uo!u:!rs I ruB{ .1 1!4 rtdus t s rt fauq lllleurPoE$ os s.

Pod InsoJ ino rrqr rd$E ollnrgrp,{tnlD@d s,r' rnsoP ioj rs.P IrN! s.Puru usunq :qr ur

rn,{ro spuusnoqrjo3 uJ ,:jds u'qrra.pnsNrn4, rn't Fnps9 rou llqzqdd r:a s:AuErttr Fiodua $qI T:ds or os 5nsou

:qr uqrh $'Esoujo astl$od :qr rno rlnr rou oP #!'q lnonuquor,ttiruJ se eSus .srqr PrEaJ ostD ]

^ !r PUr rSrnAUEI trqo'tuls fliJ Jo $tsrr )lI] or rDnb:

P ss{ uruiq :qr ro.J:'] luoonloDr, r'qrlorFd lddour sE prtx or n.ddB Iusu nqa ssaurh ol prq rolqr Pft 4qrsod !, Pn r&nguq :pNu ru$. FUo!utru :Fur E ir8as

Plqr sllP r r!:q oq { srnftrlar no qrd ar'ssrp IFsrdp 1,!\

Jrytn .{n rnoqrk 1{ntrDsnqd rFrnr/Iuoq Jo $ot ruo px?q asEJ$p rDruonr !r s P:untop rq

^trns $ou ptnod sql {366t ft:qruu:Fr iq prF:Fsns $'pon:d :uD s.Iepor or trqrlorFd lddn rqr roj trzsuBq ontosqE rno u x?rrrP n?os pur Fnpsar Prmtr:o rrlr urla Pasr5:ns rq rqSru rsos lrnoJ v

'8uqBu toor puu 5trurp 5u6 sr qrns r u:udoF^rprrsDr pu? ltun9nq luapp Fnuu ur rutrullour rn:qrD s II sr.loDrFpor (unrms) Fp,uBndur$ pn'$Dunoss lororsosus u tmq paux$or Ip$$ qr to ruamugr pur ]zs Jo srsemu,

tLz

292 l lof rNrn [ûoL,\brs: SurL CrNrR,\ ] , Norrs

et.r. (2000) rnd HanMy {1972h), xnd Hotmy d at.(2000) r8lding irs volune.

Whne ,I ofrhe .onsid.ndons abovc apply to 6othft.ln xnd Erhiopim ender$, we brlicre re hared.{dbed mcr oftl].n, a hrs lobi6(1991). On tn.other hmd, hominids such xs KNM-ER 406, KlMDR ,107, snd KNM ER 732 should be r-sndicd Gndrc cRst), si.e RLHS origin.r ..do.xss wec lcft inN3iDbi md hxve lonA sinc. d.idionkd. Tfic dcscriptions in this volume re bxsed on pla$i. ap1i.a6 ofoftcnpoor qu.lity KNM ER 406 is 6lLd sitl] *rcmethadmrrix md req*es nuch non study tlrolgfi nodqn$.nninA dhni,tues, $ n is very u,JiLely rhn tic madi\ cin .K b. rmor€dwnhout seiioudy dmrgjngthcoriginRl cnnium. Ammiak rclune esdnxe woutl

N.wr frnds, such a &4,2,rtry,/25!A, andothcr fmgnens llon Loneh,i, shodd rield, rl@ugnmod*n scmning techniqu*, r good oppoftuniq tdlind an amnte rclume, pdionrly in lieu ofwhir!{2003) asAsment ofpossible disbnion to rh. holot?.,(NM-WT 4m00. NatorsII acent diKoveiies such *A-rin,Srkladb"ala rd,4tdi?ithauw yit]lj6.rhalumcdc and morphologic,l fertufts thri wil rscrin undcisfrftling ihe e ier phases ofhominid balncvolution. To our cuftnt und4$rnding, the angnenGofticsc cnnia are bo incompl.r b provide such in-fom.tio!, tut thc future alwys holds prcmis.l

Thc carlicn ,9,,a spccimcns from Tmzmir (OH7, 12, 13, 16, md 24) aF paniculrly in nced offu.hd study, huilding on Tobirss scholsrly rftrdse(1991). Tllc dngiids tfrr RLH nad. in th. edly1970s hrv. su.ly dGiftgEtcd, and the plR*ic Eplicsi. his mU..tjon .( ofpoor quality. Similuly KNMER 1470 and KNM-ER 1590 should 6. rcst rnd i.-d.pend..dy studicd, p.d.ul'ly uith re8:d to pehli.l asymd.dy pan{ns and thc po$ibili9 oa csebmlhemispheri. spc.i.liation, rs w.ll s the Arcc,t crpftgion. Th. fomd is ofltcy ifrpofr?ne q:rding questiors rbour bral. r.dSanizatioi in thc Hona lirc. hde€d, L\e safe rypls to KNM-ER 1305 and yNMER 1313. Th.* Mo endod* ponions arc pdioLiydi6c'ni to xsign sirh my 6onomi. closur. given iheiru.usu.r norpholos,, drfiou8h @ stmd conEdeni regrding ou volnm. derminations, whcthd thcy mh! .rly ,v ,, or xdunc.d au$nlopith.cines.

The Dnanisi speinens of 1.8 MYA ar cxtEneltinpotunt, both sith rcsped to .c.uoe rclunes rndpo$ible norphologi..l ftrtu$. Wc look fonysrd brerding studi.s on th.t pd.on.trologi..l inform?rion.

Fron ou omhrion ofrhc lrous phobgnphsthese specirens, se @ ccaijn tfi* rhey crn be sendmst, xnd would be *iling to do s.

As fd rs s. m xu!., thc Abpucrca uni.6.dsp,i. (4 bd J) hxv. ynldcd .cune volums, ulthen norpholqic,l .hdcernti.. ar. undd study ttArsu,gst rem. Our xft.mF6 rog.in infoh.rion ahdihese po$ible endossb hrv. nottc.n mswcFd. Cidrbe psspcc d[i theit inr.rna] hblc6 ofbon. ar. pe.iadlypeefl ed, rbe p.l.on.uologi$t lppcriEs snou|llinded be rvheftcd. Ths. rprcscnr ! najor hiitus inourundeBrndinAof possible N.anddthi an@snf rndthe EuroP.an honinids i. g.ndd.

The Steinheim honi.id (Gemanl) n )d anoinchiatus in ihe donai. of p.Loncurolo8ical .tudy.41.thouAh an rccure rclune d.r{minlnon vil pbbrbltrsulifrom nffr srnni.g ddccs bd a*omtiw tcd-niques, n is prelenble ro hrve rn r.tud cndod* oflil-icon nbb{ b p,lprE. lnd.rd, .En x dGro.cd r6idk pFLftd to noihing, which is ou.uftnt sibdiodSimilar connens cm be nxd. foi rhe wu-pc*mdP.talona (Crece) and Cepnno (Ir,ly) sp{in.ns. Nlof th!* rrc inpotunt lor x bdr.r unde6unding ofEurop.an honinid erclution md th.ir vrirbilitti

N.wr Indon.si,n 6nds such xs sn a Grb. ( .l-20031 !rc inpofrnt rdditions to or incrcxsing sropLoflndonsian sp.cim.ns of,!.,,,, raaJ, which ro d.Esfi@ litlc in th. q) ofeithtr votln ric d mo.phalogi.ri iidion, in aolutio..ry pornti3l]y int Gtingft1..t ofthc crclution ofthe genus,9,,,, md OOA (outof Aliict and MRE (nultiftgion l .volurion) con@wdes. Wc E Ery lotun.te to hrE hrd th. opportu.i9 to de{iibc Sm 3, th. Polo}! honi.id ofNYC

S.enl Cbinese honinid mnir (rside tom rntolddZh.uloudimdtcowriesorigin,Il.rnedJ't,-44dF,i,;4J,r, sch a Lr irn, Mrbr, Yudim 2,Nxnjin8, Hsim, Snandons, Dali, mdJi.niusbxn, hlEnar ben endo.r* to our knwledge, a.d r f.w mxy hcpdd.'nrry dieicolt b cndocar giren then n,gm.n-hry md derofred .ondition. HopeirB ths. *ill b.rdd.d to our pdconcurol%ical srmple in ihe frrtuft.

Fin lly k notc tfirt theE m sftr.l inpo mtgaps i. ou $udies ofNRndsthal endocilb. I. pdic-

'na rhe lolih Se.op.*orc hominid cs.i,l ponioG

.ould be .ndo.6t, &LCing €ry inpotunt rclunedi.md morpiolqi..l ddils b thc abundrntm.dovtrsidokr rhes. honinid!. FuntumoE the Middl. ENhNcrnd.trh.ls ofrie Mount Camel and Shanidd crGhrE nor b.en

"udied by us given pDblems of ,Ns

.z3r- !3rior ddns Fduqruv ,(,rd I urv p!84! euwqrrdoFn$ NqorJo srDidm Fftftpqr :!rr

rxq 000z )ruq-Nrr't

i3c pFvplort is -ibrljr1r

isr\r

'91 t! dd utrrcld :,tro ^:N

!d-uo3 pu! spo,lDrjr roanp^3 uElg nauDd tp, i(] rFJ ,!,fuqNury:ut Lorito^,tr o(Pru

urelq luqxJo,6un!pl rqr'136r 'cd $qt itll rholoH

9n-fa dd r$rd ,nrslDn PFro asqr €Nuonryo^3 )qoqo.{s u}urH lo loogpq'H tpr ,J sxd v

'966r '.N /ihotroH

sdDo tprluo$hrra t lxmtu€qJ:ut uoont@sErqulunq Jo norqr rDr$uE I pleol s66r Tu rNio[oH

iruY Pr 3 lsFq :uI !$.to trd Nruu ,anu oaaH ta vtztq Eod aql 536r 'T[ ]rNro oH

9rarr pdoqruv turl arJpur pu) sFpou lrDnsq'36I -ru {B\onoH

t*4zr:.,rrdtu N PN,uas,r@D! q, da14,,r'Jo ui!!d r*rpli uuqlutr:a |136r ,l{itu1o ou

.r0r /3zr( .rodoqruv lirD'c !s,qru^v:uoaq

.$ 6r dd lprd .ru.pxrv )jrol tuN \uo!Dq,ru] FDrNq3AizrsuErsrouoDqo l pu! ruNdoFD(r lpa tq

)i.poq.c uN'[,!N uq€H a'I sFqril ! prhor:uoDEru!5rox p@ ,6,uollr,zr uuq 616r 'ru t*\orpH

rI!3 .p. tr rnol:ul uoDtrld: pn'uo'l rADJv pu! uoa

3/6r .IU daonoHio,IH IDNINJo unsq4l uDD.Uv

r'!-1{ruv 'u!Ip4r

J'rl arus rqrro uornp r q)

odd{ruv s,fitd fu,v i.utv .s !u,,tla.N qz6r '11 rLM[oH

$r dd \erJ uoDqrv/rqlP|v :ornqJ sluudl. i8optl,r'€uolrrd Pu! Fuolrur] iql P! ! qlnl:q uo!

z16r .rN ,ixiofioH

ofr arr19rDspmv,$luuY aaa{s $@!N

-ool 5p, il )ja'toN iJ suld nl Dluud ur uoaqqi

Fniujo sriulrd uo suors rb .uos .696r ',l! {imllor.l.6r-rrar $s r:t )oNqrqxrtdoorjouooqq,

i,l, pu! rotrus r"!Du ud.q ss{)us } prm' !!sl96r .,rr ilhonoH

'fr s,fr 'Fdo4lvlnc )glnrulrjo urfirFN$t rt) Pu ftnrrunuuoi xluod &6r 'M5 srDH

6rt si/r9a ros\tJ lor@qJ d:! tr{c z,Dq

tGrrd pqru xJuoiFrD'dsid sJnlnou trqxrrfxp F:o^ i,ipur np

r ,Lqv T{ es ..I-r unH

$. r rqlFd x r '{r,iror5 b,qrF{!rl) rsruluc

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's!9629:}3Z DI:DS ao'qt! uorj Pnnur\rvbJp,F!r,

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posio! in culy hohinid *oludotr. Atr* R- 273,4:59€

Kcii Rc. 1999. Thc Hund Cds: Hunm Biologiel.ndCulunl Odcis, 2nd !d, Chicl8o, Uniwsiry of chili€o

KiLii RC,ldg{ B. 2001. Th. Dsu

|leklr m, L in R.1973. People oftfi.l,le M.nkind,nd lr B.ginn:ngs. Crd€n Citj NY, incho! PEs.

Lskcy RE. Lsin R 1992. O!ti6 Rcorwh* Miks Ur Hund. London: Lid., BrcM.

Nnhinuft T, Mirmi A, Suzuki J, M,seM ofth. hrlu i! chinpez@ i ua ?@. NdA.d

Pidbo S. 2003. Th. mosi. tbr t ou s€looq Ntu

Plss TM, cac&! M, hu.h6j, .t.1. 2003. Usiig gcnonjs b id$d& humm bnh .pdiiadoB ir J Pb!

R3dilslly LB. 197a. ]lc fd$il .tdei.e ofrntnop.id bniirbludon. An J Pnys Anthrcpol alis-28.

DC, Yuad MS (2003) Morphol.himp.rcp nryvisusl rir&Eosuiaiion prcdii.d bdjn *-

tulinskyLB.1975.PlimeEbGi!aolution.niS.i63:

sbinBs cB. r c8d.fl..rtd,hIhry ofs,-, rr;ft lr:B, Mrnin L, Ar,rrcs B cdq Major ropics i!ild Huru Ercl[ior C,nbridge Crmbids. Utri

Iob!' PV re37 Th! boiD d lo'? ,?rr' A n.{ ldrlqsuiziioi ii .dbr:l.rlu, on.l Hum Evol ldtal-7

T06 d PV lcol.OlduqCoI$,vokaA,aB Th€ Slndoqss. od Te$ oa a/?'? ,arib cmbndg.:

V.[ur A, Lordkiprnidrc D, RighniE CB d z]. 2002.ns skull of*rlr ,9?', f@m Dnuisi, ccorgia. s.i

WaE ACH.2nor Th. eEb€lluo,!d cosnitiEdon in PlioeE and Plckb(nc honinia3. PhD T1l

wlxrcTD. :00r Erly hominldFDiwEiD qdisdio

w]rir TD, ns6{ B, Decusr D, { .1. 2003. Phei. H,d, t'?r4r ft.n M ddh aq'h, ro\iop L Na

worpofl r'L Lqqq. Pns/dhtupology, 2d .d N4 Y

sarNrwoH rrssocsswnroA 'rvrN\/ucoaNg

296 ElnocRîrrr voLU ES oi r rLr FossrL l lorr^r ! r

NorEs FoR APPENDX 1Th. cndociinial voluncs ror rhis hblc te li*ed by dononic gtuup as oftndy dpi.sed in thc liemtuBdd @ngd rlphrhetic,Iy sirlin bqd gcognphicrcgion6. Our u$ ofihese m i. this Trbt iloes noin.u that * conor sith rI ofihBe &signatiom.

Wher tllc'sourc'indicrE m "RlH" th.t memsrhe c.do.nnid eprcities wd .irher dd.min.d b),RLH or published RLH rs i. HoIMy, RL. 2m0.Bnin. In: E. Delson, L TrneNll, J. vr Couvsnns,md AS Broob lEdn. Encyclop.dia ol Hul@ EvelurionndPEhietory, 2d ldidon.NY: Cr :nd Publ.,Inc. pfs. 141 1,49. lf tbe "Some' is 'OTHER", th.wlmcs etoftd reE st n nof thc r.ent lit.ftM $ found in ou Bibli%mphll md nostl), Gom

Ruq CB, Iflnlius. !.. md Hol,d,y, Tw ta97ns.nd enc.ph.liation in A&ban. Hond137171 17b lsupplm.nhry Drr) Th. SMSM.l voluhss c Eom Baba ,,t 2003. HomcJv;um fmm fie Plei{Nne or}q sci

W. hac ddd.d b povide r choiolo3lole.h.n.t nnDl volum. to pftmir ftgrtsingni.r votuie €:jn* tim.. The mgc ofMYA! 6.of rh. honiiidr e too yi& to r(ept mFrilamiddl.vdua, hJ wc hope lh* fie.e dxi6 wilinoft rc.ult s ns dating dhniques beo@rbk Mn! ng, npafrflltr E:nymdo(triJfor th. Mrbd (Chin2) cnnium. we m'nd rdcitation rlvh.rc in th. lihbE thrt w h1E

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Pmvid.d hftin @ thc initirl b*ic $ni6tics for .I tl]ehominid endodo;al snptes re dscibed in this boolTlcse sfrtisris.rn b{ amnsed dijld.nrg 6utwe.hosthcse rrious combiiltions b(ru$, by ftlying o' oFsnsual Nononi. plr.n ns, w wdc .ble to incnxsesanpl. siz6 wiihin tdr, In addition b nch Rnd $m-&d dryiRrion, w include ninimm lnd muinunDrucs, the coeficGnr ofwirtion (cv), sk Mes, mdLunosis. iaw rs.nc thr al honinid populationi en,do.nni:l capriries N.a bsically ofr nomal c.u$irn.1or, the sdo.g ddrions Eon $cn nornslirymight ,ignsl clues r8ldins sanplins md tdononic

Ior @mplq pop'nrtiom uirh v.ry hrg. or verysrdl CVs (i.e., depding *ongly cnher ky Gomxbout 10%l suss*t .lurion is rdvisbL. Thc CV forI a/zzaa of16.1% su8gc.s a f.nltdinorpfiic smple,or r.ombimtion ofm. Wc beli.re tn. 6mz! ro heth. qsc de.piE ihe r.ant Andirys oaReno d.l. (2003)rcg,rding nodem l/,'o,lile lmrt dinorPhiso ofth.femonl h.ad in r arz,i. On the orhd fiand. tlrCVs fot "d\ Hona dkht e 76.296, and 14.996 forerly,9,z,, bccause these sxnpl.s ontrh n'ntip,c mnthd thb stungsmst difiorphism in..doconi, crPrit} HlW4 y. m rot ddfrins th* cvs dnDmorpholosy, onb. rhd th.t night haE a se i' $nri:rr.s us ro qustion rhe brsis for ttrc hid or tow vducs.MoE smpling uill ofmuse esolE these qne$ions.Sinntty [,. 1@ Cys fot 1. rob,nL|/A. baii ptob^-bly sign.l r .onbination oflow smDlc siz md bi6 in

6otn dncdrr' hd merhodotogi!:l 6su3 whmshcnngii\ompt!F endodiB. s,mJrty rhe drirow cvs ror rhc Nsmdons (soto) and lt ,,t&,n groupr sugg.$ eiths smplng or ftco'sbias,*h.rtsti A. afri..nu Nna H batin gnqCvs thdm bo( in fine sith exTeddions ror.dtninid sm,l dimorphism md poFdldon ruidioLb.li4. th. srm..ppli.s to }I ditu witlrcut ttH tudatEatn, Md H. sorEna.

Mosdy non our findings, bd iicluding d'esrcml other woikc6, on th. ,l1:md.tion oan,rl.apKibe5.!( popon m rmg. .ndo.trinpetr/ lat H. e?t4 a.nddldla'' tnzr s lNerhat tbr nodtrn,9ozr gl;,r oatl]. Uppcr PleisTh. addition of No shaljsh (npim (3 md 6)ieff, le$ ibrn 1275 nn, both psb,lly Gmar6,.bary€ ou birs rcgarding rhe ald p:lemttuopo'.hnhuf lhtr Nemdcfthal hsd bdi ii {gcr L5r

'rM, As the noddn ,qara srmpL is Mqusrio

nosdy male, rhis supporition Equn* hoF

Sim'lrlj lhe,k sia6 and laro's 6g!16 urppoxinat du. ro impdfect srmpling or cl6diof d: ln g.neral. *Jus gr. e' or 16rr rhm +skn l signidcant deprtEs. W! do nar suAse* myrofrbes. sotistics to ddehinc .ither thc con 6dwrcngn. $ ofbs I n $ic6l these 6gus ror srnd kfrGt apped ta b. sugg.sinA oveFdl

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Bnd6erd Dc, Honoway RL. 2002. A,ymm.rly of ih.fbrdl .ndoftrniun ii modni hunairicIPreEriotr of aolil odc4s. Am I Phys Anrhbpdl

Bomrs€ TC. 1937. The 6iobgn,r rid .hbnolosi{ir E*rdon ofcdry homiddi I Huo Ercr 1615717r.

BiNi B,W,rk iA, wirdCv,rcaklrzl,rrr,flJ &,tt elqi. ton a$ lire Ttr*da. Ar I

Brd. FB, tueJ jC, Ribr EF. 197r. SkuI vorumc io inf,6.Merhodolosa nomrl ulua rnd xppl,oriois An I Dn

Buho.wd.n Dq c*dqrMa 2002. Thc miniolum endmluioo ofd. bnii.3Ei, B.hxv Esl60125 15r.

cant?]upo c, Hopki!! wD. 2001. Aromdi. Brms mlitr glai rp*. Ntur 414i505.

Chlk WE Lecor 1933. Thl lndocruiil {6t oa thcsws@mbc shx ho!6. JR Aldrcpor r,s 63:61*z

clrk wE li cbi 1947. obswarioE on rhe ,D.omy of$€ 6$il luhl,tri.rdr. J An* 31:300 333.

cr{k wr r, crcs, coorr DM, zuckdmm s 1936. The€trdomnixr en o. dr .hihprl"&. JR rl hrcpol rns

c@rclly cl. 1e50. trndEl oulhdogy of rh€ pir*cbnin. Spdng6dd, Iliidisi CC Thon6.

coorcy cc, vonin IIw Tobis Pv. 190 Erd(ftoiarflfres oi t 6 l n k ? it tt tu, 4t iaau,.v.ded 6, 2 xnd 3 -Dompu.d brognphy. s.i.ic 2t 7$3 441.

conroy Gc, w€b( c, s.id6 H, Tobixs q Kai! A,Bturtd B 1993a. Edo(roiir qpdg i! o qny horinid artrium iom Stiffontin, SduL\ r16ia. Schn(

coiioy Gc, Kanc,c" scidlc!H,wchsc,Tobias Pv 1993b.Edocn'i.l qp..ity oas* 50t ("Mi. Pl.J), a ldsc Ewhoninid cruiuo rrcm sr*foncin. Aml Phys Adhopor

conby cc, wrbd cw s.idld H, Tobis Pv 1999. rn-do@iir apacig' oacd) honini&. scnna 233:9t

Cor6yCC, F.lkD, CuwrJ,wlb( Gw, scidlsH, tuchlnw 2000a. trldomni,r qprciry in srs 71 lAa,.talrhe.6 d'.?rot br drrccdiEoriood @npred bmogliphri

coqoycc,w€bqGw, s€idld H, Rchck w hrNedd.!qjfr H.jrc M. ,000b. tnd@Enier !4rity ofth. Bodocmium dd(mimd fon ihrc-difrefuiond compm.d b@grrphji r1r I ?hys Alarcpor n3:r r Fn8.

cooi cs5 1962. Thc Oish orRac' Ncw YorL KnopfCoibrlis M 2002. L,riali9 and hc@ Tl, .d, Tr€ Sp<iriotr of Mod.n Hano q?ti6.Oxfdidi OJqd Udvr6itr P($ p 137 152.

ciN TJ, .d ?0o2. Thc spdarioa ofModon nruoxfod: oxford unibitl Pcj

Dur R \.1e2s. nadatbaa gn D6.7n wSo(hA6id. NalrE 115,19t 199.

D{' RA. lolb. Trdns- ind i5 'cnifi(r(

Nr }lt

_ /DM uA. rer. Th" ,"r"ti,-hip.rb,r. sr. md b'i,

t_dn:o.hT1l s:.tus

:Ar,Jl4.ds:,1r,3rsDr RA 1956. Th. rclNioiships of bnjn rie !!dp{rf b humn shtu5. s AnJ Mcd sci 21:23!5.

Dr* R. 1959 Adftntui6rhiLddPhil Th€ tnnidcs PE$.

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Dqcon T. 1997. Th€ symbolic spd;s: rhr .d{rcr io.lansuasc ad rhe bni!. N€w York ww Ndron.

D{i D, HtbLin.l.t, Honry Rr, znsL! R te .rhc ph)rosrico!pcr,or of

'heptr.N*drJ Tsi6oh tuilnscn, Gcmur J Hun Ear 34:435-503.

d. H.inarin J, cldk JD, whir T, Hd w. R.nEwordlcrbrh c, B.y.n. Y, v6, E rqqq. Eivnnxnd behlvior of2.5-millon-]re oH noui homidd.

dd coBo M. 1q92, L. moutrs. .ido.iinnn d.rMrtu!) Etud. tr6 prcporootr d coi@uE d€ r.nd€icn w? duE d!6,nioi biomaique I bu 3)sdfri$aDEA ac luriwsii. d. Montpdli€r II, 33 p, 116 p

Ddmd Aj Chimd l. 1e5o L. pft$on d Hatuphn . C I

ftlsn E, Hs*i K, Riddy D, M?@r Lr, M@bny Xrso}t! GJ,.leob, Mrquq s 2001. Th. sxnbunsffi3 r,,o ,err .dv,;, A @EpnnE molphomdic 6doo,lholosi..l {i.\sis. And R6 262330 397.

dc Lumrey M , PiErl l. 1969. ra arcs hunaiG d.h cft. du Luftt (Nic, ,{rp6-M{inms). Mim sa

Dubok r. 1924. On Ae l)lincip.t dmff€6 or rh. ddimmd bnin, rhr nmdiblr ma rhc idh orrlir@4r?6,{tu. Prcc R Agd Amd.'drn 27:26+173.

Dubok E. 1929. Th! ri$uEson lh. fmnrdtob* oapr&d-rr,ta,,ra. l]ie R A.ad Ansrdam 32j32.

Dubon E. r9$. Th. shrp€ md siz. ofdar',{ nd iD P;1ra lrro?4 ?rc. R Aqd Amn.dd

Duch,inc B, cosnida L, Tebl J. 2001. Ercludm-4, l6}lholos] ud $c bnin. Cu Opin Ndrcbiol n:

tdinserT. 1929. Di. fosden c.hift. !€ib! Atu u ErM

l:lqDqN xrurojF:) Jo trsdDn s*:I]l c'ld lHq rruuo4r Jo sooq) rDuluNit roos

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310

1966. CEniar ansciq rtrd nuo! rumend pioposd. Ar.l Phys AndfoPor 2r:

Hono{ay RL. 1967 Thc crctudon ofthe hufu bnh ion.

erurio, or oEplq bchryior c.i sys 12:3 19.Thr crctudon of rhr piinar bnlnl

$ft asp{s oa qustun*ive ieritiors. Bnin tus 7121

HonowyRr 1969. Sofc qusrioro o!dorriio! in priDns Ii: Pdds lj Nob{L c, di coE-pn4i€ snd dolutioniry aTr$ oft|t ktubnft e nlncrcs {n.m A' tr NY Alad Sci 167:332-3,10.

HolrMy Rr-. 1970r. Ntw edo.ruiit qtus for rhe rs-

Honowy RL. 1970b. Asritornh(irc .idoa* crluos!p.riEc!, 1924): A iew slumc dc€lmhsdon. s!iF(

Holowr RL 1971r. Nw esratorithsii. endoq*, sK1535, GoE Srrutk$, SoulhAtia. AElphrs Adhftpol

Hollony Rr 1e72b. A*nitopnh(io. riddrts, bniDddurioi in th( Homhoide. rdd I Eodd ofhomidd *qru,ioa ri: Turde R, d, Tbe tuodiotrd xia crchdonr.],

chi.rso, Ardinc/athsioi r,&$,

Honmy RL. 1973x. Nq cndoqrni.l vdks ao! rbc tra{Afriru culy hooioi&. Nxtu( 243,97 39.

Ilorlovly RL 1973b. trodocnii.l yotunG of th. 4dyAfthin hooinids xid ihc rctr orth! bni,r in hunu nqaicevobrio!. J Hum rvot 2:11e .45e

Horouy RI. 1974. On $c n*'iig of bftin s;c. Scn,c

Holo$y Rr. 1975r Erly hohidd odNiss: voluD6,mo.photogn rid sisoinsrce. I': TutuidioiJ miahoto$ rtrd sou^n Thr Elglc

Honowly RL. 197rb. Thc RoleofHtrndr rvorutioD orin. Bnii 43d J@q Alihr Ltu onde Ewlurioo ofthc Humo Bnin. N

HdnMy RL. 1e76. Somt prcblcms orhoniiid bdn cndoq$ rc.onshcnon, ixon.ry,.nd nrri Tobiar Pv copP.ns Y, cd,, concong*: L6 Prui Ancieis HooiDids. Ni..: pdiinsr,

Hdnqqy Rr-. 1e73. Ttt nrcnr( of.dd*b ao! 5tudFiis pdmic bEi! ml io.. tni Nobek cR, ed. sdF4ss&os h Piinr6. Nq yorL A(dtni! pE$, pD tsll

Horoway RL. 1973b. l'@btcm of bnf cnd46i ioE,?Fh rion ud Aftico hominid rrctudoo ri Jo[y c, id, ErtHoo;ids ofArric! Londor Du.krcrth, pp 37e-!,r

Horr(My RL. 1979 B!ri! si4, s[omer], md co,Eid_Teiodi T(,!Ed i synlhsir r4 Hznn ME, Jensco c,Dudlk 3c, .di D4do|D.ft r Erctudon of 8nj,sizc B$ivjo,, rnprioiiols. Nev yolti: Aud{lnc p!*

Honou] Rl. 1e30a. wnhitr sp{i* b!ai! bod), ftisirnnh der hd odE

lrioii€ spd.s. rr J Ph]! Atrthopol 53j09-r2r.Honsqy RL. 1930b. Tb! Or.I. 7 (Olduni colg., T.tr*ni.) hominid |r{ist b in cd@4r E$ird An J prrF

Houm) RL. 1e30!. rrdorcsntr'sol.i (Nsudory) o,dofdiir 4oNhdioisi somc pElimiwy ob$4ari6and .odPukom wnh N4dJqdral ind r,,, tugraps. Am I Phys Anrbepot 5l:235-29s.

HollNay RL r93t{. The c @dt ofitr. ono ieDitL333y 6 honiiid lpcimn An J PhF Anth!@l i4l0e-

Hollowtry RL r93lh CulnLr,sr,mbots. rnd humh bninqdurior A srlrh6is. Diah( Andhpol 5 237-]03.

Holqqy RL. ]931.. liiplorils $€ do6xt $r6cc oahonioidbninod4ft !byis$Ptohqu$is. Phiros?rns RSocLod rl Brl sd 292i55-166.

Honoway RL 1931d Voluncric snd sydh&r dc€ui,nnoir on (ant hominid tddo!6k spr,I Id ll, Dj.bdrbDud r. end it€ silc //o4 d{r,r spnors on Nruddrrl baii sizc. ]lo l Phys Antb@pd

e'dd.sb eiisiEd AmJ PhJl Anthlopol s5J03 521.Horrowry RL 193ra r.rumsn pxtto olosilz]did.n.c rd-

cant b h,gug. bchayiol r'run Neuiobiot 2i05-n1.Honduy RL 1e3rb Htrfro bnin evoluiioi: x kMh forunis, mod.l5, ind rn$*n. Cd.l ,{lrhopot 3:21j

HollNiyRr-. 1933c. C.rtbiarbnin cDdcd pxrdof ,ratotapi t b.tu 4n rt u n honin:n. N Nn\K 303 1 42o- 422.

Hollour RL. 1933d. rr,u dd'lboii qodoans: volunetsi. fd EolPhobgn.l ob*ry*io!$jdecbrn sydndi6. Lt ,rrrk d ta plaft dc lhomn.dr TauhKl r*mi res honiddrs ao$ihs: coisi5 rnr.osri onxr dc Pir6ohloci. H u d,irc, Nic. 16-21 Ocbbrc1932, rP 355 366. CNRS, P{n.

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IIC

Jdnf H' 2002. On thory ii onp@rirc pryhotogy. rn:sEnbds RJ, Kaufmo J, €&,Th..voludon oainrcni$Nc.l'ranwlhj Nl: L ldbxoh A$4nft. pp ?51-233

Johrn$n Dc, L4€ror Cq Xinbd .tr'Ir, whiE TD, wrdsc, Btrsh ME, radm{ BM, copp.ns Y 1932 M{photos, ofrh€ PrioEie p.riixr hoEioa skctdon (A.L. 233-1)rrc,n rht Hadd formanon, Ethiopix. Ad J pbs rlrhop

Jungc6 \l1 1e33. Ns eiihn$ ohlopirh.iin6. ln, cr;e rE, .d, Evoa thc ,Robus, Asrdopithsinc. Nry york Aidiit dr

Ksppds CUA. 1929. Th. fisurcs of rhe cofter bb€! ofPnkfl b/.tb.rytu Dlhan @6ped u.h thcr ofNcxidtftnir mcr, r?z' ada .nd .hinpilrze. p@c R Aqd

Kaflr€s cuA, Boudo KH. 1939 Coopssoi oaitu cn-dqnnilr a* of rh. p,4a4rrnl6 {{d skuu found byDubois md rcd Kooisvrd! pn 4-17,16 sknl Ko!Ncdcd Abd qn Weris 42i30 40.

kiih A. 1931. N€w DiscNns Retxrirs b ihc Aniiquir,orM.i. Nrw Ydik WrV Nonon

Ki@bd wI.I. 1934 Vliiriio! i! lhc partn of u rl k-ior sirss nd hooinid phylog.n), AiJ phF rlnthlonol

Kmbcr wH,loh.,so, Dc, coppcs y 1932. plioei. ho-minid fuiir rcoaiE eom the Hrdu Foinarioq Ethiorir.AmJ Phys Anrhbpol t7,a53 500

ncirRc 199e. T]t Humn Ce( HulrE Biotogisl.ndcuhui oiigis,2d d. chicaso urnkisig ofchic,so

xlcin RC, trdsft B.2002. Thc Drs ofHuor Cutiur.

Kchdkd? U. 1973. Parcoe@rogr,. wshinglon, Dcl

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Lcke) RXF L.!ii R. 1992. Olisirs Rcefidd.d: tnsqrh oawhn M+!s ui r.rum.n. Bo

LlMry M. 1976. Mqphorosicar erbnt qmm.ri6 ofEodln no, ad nonhumn pdDrs. An! N.Y Asd s.i

L4ii c. 1916. Frri.J od.intuoriqr chlncrc^ ii rk humubniD.AmjPhysAdh$pol22i345 330.

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MkKionon rL, K.niedyjA, Dris Tiioi orskun apeiry arcm rc. c€notogir n*Eoo6

Mdcozj v 1942. Uiftrr f.niry$ rcsri ,ooiii Fe i, ner sinuropo, e ncrr. scinmie. Ri{Anhpol3.:

Mriqu@ s, Moys4y K, s$ls cJ, Jeob I sitE$ a2001. A n$ ro$il hooilid ..tvsi. tod lnddcil-smbungnrcm :r. Ad* Rk 262:s36&

ti@ itr D {olosislconM. N.w Yo& Ar{ilu Mu.um ofNrnmi Hnio+

Mrin RD 1931. tu[drc bnin sizc rd bsi me6bd[.art in rrc*ri,r Elcb*ci Nrhie 293:57 n0.

M in RD. 1ee0. PliEaE onsi's {id *olrioni a ph/eoro Hslr?ined

McHeiJr rn\{. 1930. Ns s.im*es oabody {cigh iocrly hominids rnd rheir sisffarcc !o .ncahr0addnxnd d.sido ia io robu{" ru$x opirhccins. lo, c;o.FE, ed, Evoluriodary Hisbry of th. 'Robun' A$eJop-ithdns. Ns York Atdiie de cturrs, pp 133-143.

M.Hetuy HM. 1992. Body si7. {id Foponions iD etlyhooinids. An J PhIs Andrrcpor 37:107+31.

Nihimu rl Mik{mi,t suzlri.l, Mnszan T. ?003. D}s.ni of $c bryN in chimprn?.r inaxn6. r,ior Nid Asd

osaw I xemiF T, sxkr s, H6orew, H 1970. slmobafttions 0! thc cndocniii sn ofSu?ukj H, Tskt n edibn. Thc roud Mo end His C*si'e'TolgqKeigakulppl0T-4]J0'

o,o lvl, Kubick s, Abqns(hey cD. 19brd s,nci. New Yo* Thioc.

Pebo s 2003. Th. nosi. rh.i is o

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R.u$ TM 2000. Tzring th. m4surc oadr6igi @Epq!niE ,rn.riEs ro rhc Eodcl ad@l plndigm ii oni..lndrcscicu. BEio B€hs lvot 55:237 299

P4$TM 200r. Thc Dnc@,r ofc€rcbnl Diw6itr AnUowl.on. S.ienri6. turcrudor camb dg.: Cxmbdg.

PrcU$ TM, Q! H, KIS,JH 1'99.Dhd os,i?rior ot hufrm piinaq,;s!r oaq. Pmc NdA.id sd usA 96a1601-11606.

P!.u$ TM, Ciarcs M. Lauchs L rlomic b ideniit lNf.n born spsilljatios. Am I Phys

P}tnft WP 1923. Rlod$ho nen: d.*ipdoi of thc 5kulod dbn hu,nrn fndns Fom Brcketr r.rirl rn: P}tdft

Ts.Jrrrtzv En luv uoDqom rtll'ull rol {o!Fdq a:dr Fl5 u, .tr5opuorl Da lJDa ur A4qimrl

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