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30
i l ri rtr.o s,\N D Il^r .Rr r r - s of DN!o.,\sr IN,\r ,fsIs
to lird rhe ..do.nni.l tulume olOH 7, rnd HdlMy
(1972)had similt poblcms in aftemptins ro 6nd tot,1
cndNnnirl voluda fo! thc M,trprnsgr speiEcns
of theMLDl o..ipitalponion,ndihe56 19conial
broeportio'. In thsc sibrtions on. ries ro xs.eftin
vhrt pere.hge of rhc tord rclume is rtprcs.ied in
! poftion ofrn c.do.!$. In doing thG wnh a wiety
oftua, one hop.r ro rivc at a perc..hge 6g!rt thr
n fairl) tighdy disribubd. Tobirs's(1991)dt.usions
cgrrdi.g the bipri.r
bnncl olol-I 7 povide x debilcd de{dption ol he this done. (W tunh h ad4
trat htu wJal uarAflrl/ tr dahz;n thi
P.flkak.llt,
aa, bdt k t?a,t tb aaoihbk dat brr:ld4rtuE dn
@wi: ,11 k nardt.ttti,
6tinak:.)
Figur.11. Dcoo!*Etion otendo.st rctummicdti
m{ior n.rhod, sins w*er disph..mei. a Endoe*
h sobmeigeJi! ! llrge b*k{ frl.d b tlt qLlnow ou.
let, ,nd rh! dcph<d uit$ is .rptu(d i! r bsks of
knownw*ht. Br Thc bek{ holdingdrednphed qrr
iswigh.d, givingihc wcight/volun ofrfi. tnd4d
wcigh it in wtei Th. dif.an.c siould b. the vollnd
A.dEtc and reliabl. mexrurmsns rqun. no.pomus .ndoca$ mrtends, .lexn glassu{., clem
(pr.rcnHy dnt cd) wt.a xnd x..uor bahncs md
wight.I! is ilwys diresing ro Mrh bubblesdis.
due ro iopropd or inconplere .oatlng oi the morc
poiousF.nions of th. erdocst, but .uch ar the drn,
ges inhe4nt in Pat.on.uolog.
PARTIAL BR^IN ENDocAsT VoLUME5
Ii is ofrei thc caseihn the avalhble .ranial ponions
m roo ;icomplete b .Iow r rtlirble bhl acon$nc
tion of the hissing prs, xs rhe missingpRft saed
whrt n prsit, Tobias,nd Hughd Gcc,in prticultr,
Tobis, 1991) tun into tAis problemwhcn anemptinA
ENDOCASTVOLUME RELIABILITY
In the Volure md M(hod sedion of exch hq in
thisvolumek giveour Gsc$m.ni ofrhe rcLxbilitI ofr
givenendo.st volunc. Tfic reliability is smrtd r.cod
inEto nethod (letd d$isnation) rnd ou .lduxrion of
thc sim volune (ilhcncd d4kmtion). Th. endo
ca* volums gikn hdc rft obhined lir on. of lour
ndhods: (1) difd sxqdisplacem.ni of.irhd x full or
! hcmiendocx$sirh minifri disbfrion rnd plsticenc
4ondruction,(2)det trination using, pdi.l.ndo.a* asdcdibed by Tobir 0967, 1971)rnd Holloqy
(1970),(3) *ensiv. plcti.cnc Econsdction rn.o.!
ing to harorthe tord .ndoc6t, and (4) volune cddhed liom rcgre$ionfornula or c*inrted on rhe b6i!
of a fs measuftm.nG,whi.h rre rhen plqg.d into
lorqdas *ch thr olferd by MacKnnor *.r. (1956):
rl = J10.5(LtrB
+ LWl! )l
H{e I is mdlnln lcngth, ,/ is uidrh, , n tngth
tiom bftsmr b dr pGtuior limit olihe cr.bellum, ,9
n the height lrom tfic scfrq to the de.pe$ prt ot th.
tcmponl lob., bd/ n rhc tun spe.itic .o.6cnnq
xreaes to paviously publishcdwlus rhxr nxveeiticr
hen .on6rned by us or, in etuin cases,not. In
xddition to b\e endo.a* ncthod sores, erdr .ndoca*
n soEd nunen.ary ro indi$r our asssfenr of rhc
dj$ility oi the given vol!tu. Thc* rlirbillry vJu6
m.nluated on a s.le of1 to 3 wheE 1 i.dicxks thc
higLci ftlirbilrt 2 dD fic giren rclune is gtneelt
alirblc, and 3 thd tl]. giren rclume hs low dirbility
Bx,rtr Enoocesr Voluuss By FoRMULA
Ar uitb rny physn.l objcct, th.a REreldionshipsbc
Mcn the rclume xnd thc linc{ me,$rnen$ rhat
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36
Il rrn !u s,\N D lI^r r Rr \r .s oF INr oc^hr ,\r ^L\sr s
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Pcfr R. 1905.Bioodiczr sodi$ in m.i.I. vxnrdod ma
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The SourhA&i..n FossilAF Mo, TheAusralopnh*i
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'uruo:ds SO9aS aqr udj uiis Nq :a s
lnr
iJ
PUr DUo ans sqr .Pn:f r'lt'u ruoq Fi Dtro
!unorru Brv r:,hos!p rqq qrh
P$r9rd-Ipar
rnoruoqeqn suo!:Drd.s.'jr sdqbd 1t86t rFJ
,tq 4q pur (5/6t ,rt6t ,qsuqlu /,q hua,4r prc
ad !' hoqs lF,ttr uoD,pro: s !h1n. rruq pud
pnlb!}uud pt8trqu
{buau. @ qrk,por:!,!s
leJouoqlPuotrtrq&ouosDrdur{s,qrP,uqr{q?qdd
roa dnof I@sDm rultr suo€r l4uoij ro prdoro
Dqrlr u llrruulsr F4rD: rurtrgrtu lu! iou s4q
lqra lo Auruap|o,i ro uotsuldri F{q ou
Frrqld
p*oqs {qrqod rrsr usrq.qr:Ftr lu OOfo oorJo
:tur zs rF ursh dnorSF$'u: w qrnsJo{rD?dz)
P'urr rqrre$ runs* ) \',r,9Drqpwhrqr,{ap
€ ftrDUrD.ur Fu,uo'jrqr or4$l
'ro1rsutu
ur ui$ rD4 aqr$a lPqa us!:N oriqro q.e. $h
sd"tsuolrtrjo uoDrumE:p arm} u: eIIr or F9
an p'uh qinoutr rpr ord ro! op .{tdus irui^orsrp
rser!iq\t$
uutula ro )tutu,ru e,qt'
qpuu''/'!,tu
rk,bd t,&,q8,tux
4qupy
'nrq4r'lpv
':1tuqrqt
sNwas sptrgr!-r sr iureu
"./aqtu,Ftt"s
,pup
rrg sV r,!r,/,
,. uo{ hrDusd u,
Vrr{ t
or t uo5 !.uor$rdoFrsnD .qr uol $uor uornl
oD Jrlou sqr uo aurrq ourph arqib rql
rt'F9
,{rnu u:rq i^!q,!ers snu s$ood IEN:' iqr ,rzB
uruiq u' iriDur u? s, pros I$oJ :r| Jo srinn* :rfl
uo! $:urh uD.{ rrJlp r.L{rou:qd pure or.u
rqr rqr rq nFrlttr s, 1r:trqa puv {rE* loql'u oI
or 9 rst rqr u'qrh priras ltqE,tdd rou .poFd
ru! uo5 rurudoF^rp kuolnrdrjo
s,!'r Fnph'p
u eo Drl {rl srdr ruap rNqri.qu:ur
or r1!
idur ethp srr, !r ahoH rsllllo8 pur $zuidurp
rnrF Jo loqr or nlur rD{ffius rra ld,qsuorq
r Fruorplrdntu rsor rosnrrdqq
rd} ur uo{
uonnto r .qr :!!nP Prnrtro eSuz
snu:r1 lalrt prqbtrqns eqrr8n::o bqr ur'ta.$q.
r@d ta'!B'!
D!r{
.qrjo
qtrnu
aor{
rnoqs
&rii'p
rouurtr:
r rJ wd .z-r s.rqBl
stqq
\'(,{liuunr
r
-t
pur !tru i:qv ,'trtiu !,uuq :rI Jo uoDszrurrid
Jo
$posd: r|h (pqqn'pDru, ro) prrrdsarur !r*D
-psa srur Dr|o pip4.nrrund sru! r, 1n!l r$oE
nro€ rqr !tj,.tJ
Nornlong crvsol/4l
\ r1N1c r nns .s \r r u u N:l o r Nr n n tl
286
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S o iE CriE R^ r No rL r
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8
EI H.
a
n
IVILLIONYEARS
Iigure 1,10,Bnin
S* 505 boin volune Gdtei thr 550 nl) n hq.r
rbxn no*.hinFxnz.es, but ir wodd b. r.xson,!l. ro
exped thxt th. Stu 505 body sia ws .lso gr.xEi W.
.m quibblexb.ut theserbelute md rchnve &ain sizs,
but lhe frjor point $ens cl.rly provrd: ihe hominid
bnin u.dffinr
ftorgmizrtion befor n unddw.nr
anyndor expansionin size.
As fturc 1,10shos, *ly honinid bmin voluns
m nosdy $ ic dnoughout rhe .rcluriomry life sFxn
fon th. Hxdr specinensihrcugh rhe Plocenc gkile
South Atican speimens G.s., Truns), .rthoush ibe
hns pobxbly shsvs r slght non€lloneti. incftase
in bnir sia wer the .did Al 1-14condition. Mor
spe.imensqil be n.esRry io denonsr,E ihis. In
duded in this rppmnt shsis ofbnl. size is th. o.e
inomplete bmin endo.a* Rvsihbleld 2. trrti, rt.on
stucted b 450 ml. The robu* ,ustalopithecines, r
side bnnch .ot le?ding or contibur\nlb rJE Hama
line, rc clearly lRrg{ bnined rh.n ten 2. .tianu!
cousinqbut this intreN rns lik ly an allonetric o.e
rlatd to r lq.r bodr size.This shsis vB perhrps
some*o nilion )€aB in duntion, during whi.h it is
like$ but as}tt nnprwed, that our anestal s,,,lin
cascus bcsinningto bnnch offro n.nner,4. rlrnana
or ,r. s.r,, fro* lilcl thc lafrcr, sivcn thc tomaining
d.nbl lid coiiJ @idcn.c (AsFawct !., 1999).Tn
mtu* au$olopithecine rcpaened * Swdtkais b'
sK 1585(Hollowqt 1972t) shos { clcr lcfr o&ipiEl
lsrnmci, and a pu.Leriig in thc po*dor o.cipiEl
lobe, su8g.sringx ftduction of primry visu cod
This .rn hrdly b. rxk n ,s prooE hNft(
?.lh.F
rhe .*[e$ ,9 fu lin. hld hn]ns not only lcldjng 6
x dnn ic size in.rta. hut .lso showing rhc ps.li.l pnrtrns sseided widr h. h,nd.dnes rnd ,ls
.ercbnl henisptu;c +e.i:liz{io n. ycr rh. ,4 boiai
forms, su.h xs oH 5, Konso, or KNM-ER 406, do
not rppetr 6 showsuchcletr armddricr. Nor do tI!
edier Wst L.le Tukam sp.cim..s unmbiguoDdy
indnlk .irher enlrgtd or ffirgbized bnins. W. tcnr iv.! asrt. rvith r.lker.l. {2000)ihlr
'hel.,y';arB
prcfonbl lob. rppes les poinr.d rhrn
in edlis
rusE:lopnheci.e sp..imens, bur \l lso belicv. rh.t
caurio. is neces.rybecrus rht f.rtu is quirefrirblc
een lat Prn bdglalth\ nd 2 ?d,,Jra. Thet Brcci!
capregionsdo not rpper humm like on th. specineG
cuftndy rmilxble. This $$ss rhrt wirh i.ftsinA
rdrpttio.s to nird envisnnens, md r gndu.l de$
iccation of aore$s,mtor,l $lectio. frvorcd x &,i.
44^4 et \tu )rta,qi4 8u13urj
q ,t r41 zaP r/,P|nr Piruqburdh| Lr
rt,f,pa
Frq InaB t,tt' ou.Hla 'upl 44 r,q| ut ,a,r$g
,tuqpu lprtta'lD i'
tu'nt'!u, 3,
rol
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tqrud
ou PUD
prorr ltl }rssin'J i^qu16 ii'l rl4et
FrqiFur uJrNig suolizq.'jd. P4,D: fur sruPr
luflrd PBid r't8u 9:l
-purq qro! qrc\ tr!,fo*i
! \dD 1'nlg r' s,!ruu^r q ql
-ui urJq rioqi t|u * uiq l s,,s trdu$,(u! $ra
tuFip !. ra rr9! rqurur or ruuodu' s, rt rr
-^?qrq urui'i unfou or tuPDt i:!q .qr ur srslsql
rlqlsqr
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(536I l?MloH.Idur
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rzK u'r,] u' trsBrtrurrr!:don! rurpuur
u" qrN-tn'.un'qor lPritjjs u' $8 ur !s u:.q lErl
tp,j!:puDN :r| or qsndqr
19,1u tJrpur r.!r
iuo os pur'utrturDlt '+t
suop!:a ourdrC DnndErV
'r'luors!?^{s uD,tuDrs & Prrrsrd.J s, (rr,:4?
o\'flrt
a/,I7,o
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+?tr/ H) otuaHiqn
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runto u,u'tro ru:urfiNlu: N.rlnts r nd rrqf r,,
rd,J ,z,H Urpou uog atqD\srn€uls,pu llrru 3zs
,ipo,t r pErt \4rr,t, t? :pnpu :r\ rrq p.r) r,r,,
,&4?
1s:aoq iuftrDr rr r
ur lpoq u,
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Iu 003 PU' 091uarN,q or :unto unrq ur rs$u,
:qr ru]r iIJtFn sr r pnFtrr rnq trzc lpoq q, aotril
ro! oP,idur r { ot}r xg- IN) roj sv lurufq
!r Frqr.r uo P:rq loDQdrPr Fjooeq.q Prtrw^Pr
rotr r roj srnserd uoDtr:tx FJ Sulrrgrr tsDord
rDruollruou r {a rB ur4 ur x
rrlr trEFrrds or snsprq sql p.poq lq uq t!qr{u!
)r3a $!r!Dq lln: rsr,! rvr rxddp pFoa rr 'r.^.
qutruePqsmus:qr PUPsHouo4
aoq
z9
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r$ uror:I lza rPoqJo erruelr Fgsn iPu
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lnru.utuJ oor pu. a{ dor t? r*oC MnPtO
uuj queuDlr Fruunsod xw (ttr*uq4jun
(a002i F ri lntr^) rur lu 09t q0E9
xqr ur 3trqu n'ropur ur.Jq J^4 itrutrnddi .p4ur!rp
lllnl rou +q^\ 'rdorg uasg 'tdroD udj st$lql
c'uru(I r|J I'oto{Jlu
pui luruitrlur
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upnrd,r!r'ual/rtug!.PrEruoPdt uo]Jsru'trrd!
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i3t6t qooq 04
I4idr ,ifurpdtrdrpurpur '(266r
s'q u' ,{qii-I,iq farodr .rt6l u' tudoloH Iq prq
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r'! uo! eu'rqldolurnB
iIrE io tt<] rttp ouaH lo
rso'rr 1!l ,rou 4.rrug.P * rsqr suo€r drr s,Dorfl
pus urld IBrpFd FUo! rrtttr-IDrdD:o rfl s,ipB:ttr
r
qr '0t Iq wN){ qrra r. tuol{ rnrtns azunl
Pru,uo 4onasod rr:ttr 3 soqs rr:oPu] ur Ho rqtr
'uoDitoiuo: Pruo, ron{lr Pr.qr :qr lo suo€r drtr
rnPr trurluu{sD
Du !5nour rou sr
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r $s$sod turunrds
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sp I$tq :qr puB 9r HO pruru'rJj {ru:nF
rqi
rcJ rd:ri: su:rDds saqr lo euou rnF rno iurod oi
ntrNdm sr rI rruauFql Fu€uo eqr Jo uoltpuotr
P.qsutr rqr u.^€ rd:rr or rFrg,P Pug ra uo4h
nsqo u? I Ho roj uD!.d FFld Fitrnd {.t e
pruntr sq (l36t sDrqol q9no9F lr*dde rou .n
sullrd Frorrzur€ror
rlJ 19r pue 'tr !t HO ur
s)
ar ro
.r s) peroNp pqr
aarduour
('z
Ho
D D s!.uD.ds c^nplo rqr Jo $nur
aql
'rnv puD:ddng qroq u sdnof r4,, ,!@IJ ur.pou
roujouolqo :a\rurru:^rru.xoduIFnoPu.urr
r sk rrqr .oN6m rDqtv-Jojno" Artr lD^. s,\
]AP rD$ srqsqqrF,{tn:ttr rr qcno$F (zo0z T rtr
sunoqrC) q,tdt,!@ri.
s splg wdroC $luruc
rqrto rurrrlnoum ru.u.qr sr!
or prPPv z9 Ho tr DnuroP.qr sefl.r\ sr '9031d3
'I IN)PUrtr3tU€J ]N) sr!trDssu.u'tr.o3Pluruoq
Dqro Iq p4Eqduotr lqrry s' rqtrld iql O0OSII{\
'I^IN1 iqrnol ruoo|ornN .qr ur Iu 006 ,{tqtnor q
Otrt UgJ\iN)I ur IU zS1uoJ riu.r ruruaJs r$:t
trqI rrqqu.Prru.rruor s?suoqPut$pr!{d:ri'ra
aaaH
$u.rhr,,
{inolr !n ortu'truuuorI|nFiqtd
pur !pit4 auas urx q.q uoBhrp rqr rq t'DDa
uiErtrids0005rI\\ J INI rql uot t335ta NN)
Pur tlra d! ll{N) J. rqds iilrl r$ Ptrt $bIlrPtu
o4arl o luiuntdr $'lJo rdtult
EdJ ro'rr utgr FlPoq
nr rou rqJ trnDc
'fuiflo
rtnt tur f^iurrq i?lq ,i!"rtr rt VI'^t g I ,tF trd
f ,uiunids o6sr!3 ttN) PUrolrr !3 NN}I :'lI
(l Ho) ru o .r iorr or (rz Ho) r0 06s,lllid
doj tuL3dr,!'ridb p,uur !trArquru{ mqs sFsot
$rtJ nlug.{qrtr,l :qr.{q +Eu it rzuqJ,'ir5roC
udJ rr?,, ,4,a * P4ruls,P tPUg F:I
'r^iflorrr uo
rir
P q \rullqBq r$ 3r or Frx{rr u::'l
rut trsrqJ alti,uurFod Pnr shu'r't
qroq Pruu
reqr sPmuoq Ja :u'qruuq lIuoDnl
Pr trrou r*
o^tr {r o lxd ur iP:t rsqr strt!flp Fru:rroruul
ruuodur u:,q aEq rtru trDqrV,{I^I Z lfl5rd rV
'(966r {honoH .$) /]1 rol {oL5t s
$.qr uxqr Ptrtrur^PxraD $rlrddd D:rqo Pn sdqs
uolrtr lrotds 5!!fltrrddN Jo .IqsdE: uolErE,lo
!1JZ
288
lloNr!rD E N D. c . \ $ rs : s o iiD
Otbtr lo$il honinids,su.hasKNM ERtgt3 and
1305,m fonklt puzzti.a, lid pshaps cD bd licy.d
asidv,.cedrusfr lopithecin$.rrs*mpt.sof rnini
adaptiverxdixrion oaedy t4,, lbninids. Whilc it n
comnoo ro vj$ KNM ER 1313as simptyr f.mrte
KNM ER 1470 (Srin$r 1986i Wotpofi 199e), w
hd thrt rbe d.snc ol dinoiphism oa br,ln hlumc
(51G-752nl) lid brai. endo.in horphology doe6not
supportihe sul!]csiion rh* rhey nc th. sne speci.s.
KNM ER 1305 n a puticullly diiiculi sp..inen b
ft(. we \ould h! morewili ig ,o (nrctuin rhefK
ulatiwpoiibilir rh Ki\i\l-IF t305 rnd KNM-ER
1313 aE sdu:lly dimolphic form. oai di$eftnr qnv
tara (ondvh@d ausrdopithein.) Iine,at lest tu;
thc vies?oi.r .f bmin votunc, ,rtnougn th. edo.n
nial norphologics m quiE dilTcrnr (Hotowry, 1973,
la ths. concbsions,dnwn fon tlle p,ts.edo
log,cil rnd ortrd crrnial ,.d po$.nniRl evide..e, rc
corcd, mo* ofrhe im[Frhnr fto4.nizationd chxn$s
wcE .omptete by bce.en 1.5 and 2.0 MYA. Ir n.r,
h! fh tunher!h!ng.s 6ofi !.trbBl ir mned
'n oAcrc.rcbrJ,aE-j
Brcc?s:nd Wtrnrktsrsion..rnd
bclhr G.eWcrver2001),andsubmiicil rcgionssucha5
rhclimbicslstent ol<place,bur sid. Fom thc ftftbelIun, w. cannotdekd ttuse Rith our nrcscntn thods.
This lerB us rh a s.sidcnblc hiatosin EuoDc
in l4T prtr berl* rheAtrpumr dnd Dm,.isi eidocs$ ?re noi
d.srihed (sid. fbo stimries of
bnin rclume). It'trds.laves us uirh tfic Ncrndetrh.ts
(xs rcprcsentedxt lecr in We*ein EurF) shdinE
eddgcd cmniRlc,px.nis ind no evid.n.. oicerebi:l
pnnitirncs, *c.pr in th. ph+ephdi. shapeoarhe
cnni!. Mos enphrticdlr, w *e .o digcrcnes i. rhe
noehol%yofrbe pdontal ponion ofthc fDnin bbe,
r inding ongrenr uirh tlE morphon.ric an,lFis of
aookein d .r. (1999).While the Nandcnhrts nishr
bxve bccn a sepmt sr.cies (in dre sns! oa. mor
phGpecicsrnd .ot biologicalspcies ii rhe modtrn
' do not se how eirhd
'Mryrixn' scnse),we
the b.
tBviorrlorundntyinanruDtogrcJ
morphotogrJeMWeprefcrto agrd Nerndftluts s sepme fDm mod,
si 1444.4rt'r tr oity ih( subspai6.t.vd (Houonn
lqSi,hutof our,e rnrnopen romortlonnncinge_
id.n..onthisnsu..The160i000Ft atdHcftoHono
rlr,,i?ar,
cmnir (whire ct al., 2003)wix cBntu,ry
yield endocmnirlrcnxlns thr ft e$m6:lly th. sm;
asin fr.dctn Harn nlnE G.E , size,rsynndi.i
lD
br pxtrns, and perh'ps neninge,l v.$.t,
B!tsa!,roRAL DnAM jcs
wtit. mu.h hI brtn Gtrd isl mrd. of lhe n
quitum-,t .utrrJ Jd\:n\!r beginniiar lhr u
p.r I{c i,thn Nirh'rue ro,r Jalt,H!.h
s b
tool rndnioBj c?flings, pdictsl rtj jes.ln
rvi
cnd rcLgr s rite, le.g, *c l(jfln, 1q90, Kl.n
E+q 2002,md severilp1Pcsin clw,2002
reBrd d tuDLdJnsmtr5
r fic mo,rIke]yeryt?n:d
lorrhserdqn(.
Dfief ihh norotogi. /b.hrqo
.baq€s in cognitivecrprciriesb&d on r singleor nol.pre muhrions or rhe rcquisnion oa n$r.t hodurt
r,.e.,Du!hnn. *i1., 2001I trF, rhouah,ihsdi
po$ible tAd dmncs i. oft undcrtR.di.g of thc gF
netic eLielb thRi relft ro bfrin dev€lopn.nt ni&
illumimr h@ the hunan biain wotkd ar rhis lfu
Ge. Pieu$ ct a1.,2003).Thc findinssofrxxbo (20031
shouingfrgread gc.eti. di{crencesin bnin gcnomic
mdi.1for1taru thm chinpanzce,derly su8ge*tlE
natofll sl.*ion
h.s optrri.d
on rhe nflrogendiG
bumaobrtn cvolution. If one think xbour rhe difr+
.nc6 in .ulhnl b.havior ton rh. iime oi srn Edis
to the pfts.i with the u$ of.ohp[r6 rnd ou &FF
dcnceoo npid *chR.g.s ofinlomation, Sxtilirs and
thc likq one shouldb. r litd. slcptic.l aboutunddlF
ing muhrions subseni.s cosnitivcabitries ihxt .oi;
nicnd] spnnginto b.ing $me 50,000ltas ,go. We bli.E thrt w havermained thc samespaGs, rnd @
perhapsthe smc specis 150.000to 200,000y.!s tI6E ihat. Ifoie wcE to only thint oftbe no'pdiJEtt
rcmrins (i.e.,sbne tools)of lhc Ausralim Aloridna
vc muld $il bc hrvi.s ditiol9, undNmdii;
di
.ompldity ol ihc; soci.l sruoural slrftms. Wou]d
not ou gr*-g4ndfnheB, in&.d, grndfrrh.6, t
maz.d
rhe obviousdifertnccs in int.Istozl, n.
edal, and spinbd liacgles rhrt fiavc .evotwd" F@
rhcnlfetincs roouFwithout rnyenpiic'ldemonsddon ofBen rn (uhiions u.ddlyinF su.h belxvioG?
Th. erli.* pirses of honinid si*ena d. p*
ti.ularly open ro .pcculativeenbodc.y. lui when dl
is s.id rnd don , n Bmrins rh. sonc tool i.dusris
or ftditio.s rhr can inforn us thc mos rboN h6
ninid cognitiE abilities.This docs nor me,. rhxi E
dnqard rchaslogi.al conrdb suchasthe frun c
nains, home brses,rhe evid.rc. (or hck olit) for tiG
inponrtion owr long dishn.es of stonesusedii n.ting &ols, de iahing cmsses, or c{n crnnibdiso.
Holloway (1967,1969,1931)sugg.stld thr so'e root
mrkng rnd lmgu!!. mighrhrt hJd
'm'tr.oBn,b€
uid(p'nnings,prfndrlj iadr sbn!
rcot!jhohtd
.l.d dden.e ofshndTdization offd.m 6om clem..6
rssritr
u, uoornpr r or p,t ) Bqrq5rd sn:$u rns
n! rrSnr pur Fuomq usNrq ro!trmru
:qr ur
sisurqJ rsurs /ruoDnro r n ur suoDudspEAsu
ltFquaodJo
?-uor
Fs
e.u
Fns to Lpp :qr plr
.jJurur.q pur Du:purd:p
FrBUNodto spomd prtuot
-od rqr rJ Fllssr r* rE$ sr5]rus tuD{u ru:,]
-gt tou or P]l suoqqdDPD
Ftrruorzu? PUUP:€otor
itq.l
pnqroJo
sllll turdsgo pus
rrs
ltuourq]q
!1'u1J \rt u u:4:q
sdq$orrlr
FrU:uqduol
lu.door rx,u pue lsorr Jo rrs r ru,.u 4qsqoid
sr8!r'l: rirlr r4&,/ta, !'d Iq p
! lide u' r,! uuJ ru!.J.IP Ir EdIrFmstoqn
P'F! ? or Pl r?qr srtu.qtr Frourqrq osp lnqord
r
trfll liliuiJ ur m$! rsod
ru$g'P Pu! {erq prirqu. rlruroDd s rpns $D
suDarqr Pn$s ,trpuotrrs Jo smr.:J rtrur$d. ur
rsl u' u! rnq i,s ipoq pu. qroor osqdourp Fn
b, Jo q,lrnpr ruq'uoruor r {q.qord sh .rqI
'(uoDdtrsip EroJ !196r .,rtuonoH st (zB rpoq puE
u,uttjo ruid,HrFr pu?f,ro plurji .qr $ s. srsurql
qrN u,r lrq :j:?qP.lj r^4sodJo uoar Sumlluo rqr
-r'r) .uoD,rg,dur u.4t6it. prn"r *n rqi
,rrq
.A lpoqpc uDq
.qr.,o stnpirPs l{u.udq$ip
rqr pntr4r rqr sns!
rrfta pu suouo,jtuh|du,
lfu.gr trDru:5!o p:*q
r
s.tusq:p'trs$i!I!
r u, l[qr Surdno$FDos
qor Fs rq&ourp .. Du:doo: rioa ! Jo ru:odolr
rp aqr s:pq:r .ssqd rubr,}rdorAsni ,(pE rql
'\ba' ,/a ,oas tu!
pu rlq r4dt tb/b!,1{lr n8@p ttqr
Ppu'tuts,ru
.rnqn
r*at ur4rr^'"tr4,4)d!1ab./Jr,?e&4
-u, n,rPl Plt rr-El'urq tLrrgp h tuaqDrryBt
+a'uJo lu"eSta' ri,rualn
/d!,n la lbtua
nf ru,tud),? !aun,!,z-,!
uu1 pn
$4 p4,&o
,u,op!?
3uo4,14,?,
pr,
tuthql
t"-|t,rt1q
tbrd
qus
u) buoHtpu o ru,p,2t ter{tu}rilrlap4e,
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
:qr s
sou
{$qod
{! F nlrtrtrropu: prloror
)u:3 norqntr tur
uo prseq!s9!rqr trroryro@:rt piEA, ,M
^to^x
'grtdr
urunq to trtqrdrtrIflrg rou :r.h sFq!trpuNrN
qr
se
^'l P'u:Pur
!?rt r .!crt FrFrlpur:N JI ur sraDj
. D'uud
Jo !r1,tu. qrra P:u,quo Irrr rDUotr: \ r. lJrio
Jo 80qr s. iiPrrllrDll'lP sqj sl$o runoroid Plnol
sl?tl!trpurxN rrqr onF I'qr puD.(otl d),,qr:tr& iqt
6uz
q rrn - \ l vn rN rD
rn u s
piddErpuq Inxr€otoqdrou rou *a spq!.puu.N
tqr 5! $ u Trs Frt op ra
,, :I"s l:ql rsBqp,u
iJtr rrlJo uo!trnnsuotrr r$,! z uo P.$qs, uonnPUotr
uDs rqlE Prrror
3.^ i rt nqr I$ Pu. suDunq
,t iidoto!&or
rn sFqlrpnrN ro rr! p:tu :qr
r^rq (aOOZ) rr unH lFUr.)I
rtql
P+:i'm
F
'qrsos u.unq urPor
Jo uoqvuoqdqr ur {Iuo ruaDdu Sunq uoDEodFrR
q rtrra .Ftrqur $ou r:a rDlll sru.urF lun:u :qr
*arI lqr IsJ !$u r \'rSDntnt roj &Dzdlr prlu
Prurn Pui lur$P t triuI{
ruardu
rcq or sz rD
Jd rqr sPnop
sqr q 1r00atqru pn qnu)rr:p
Pr u PUr(r0oz! F rr EnurqsrN)sr:znduqr u,
ursrp prtuliq aoqs re{l !!dr
ruJrr lq pm€Eur
* :M rt?ijJq puuoqjo rrcud
:isl' pqur!$ ,tjqntosqr rouJ, ruaDdur u, ua:q sBq
rnEI Jo run$t rll1 *qr 5urstrl'ns paruNrd ur:q
$rt uouBlnsd, ,lfu pui !ur!g rtr u.qr rou sr{d
,.snorjo 5tti9sr-t lqlno{l
Asno rduqrFId
0or rou ieJdr ol turuuJtxq.n
ta,zruduqr ru4xr roj xogr ueqr !rer5 r:sJn
pc 5!l:lrd loorn4utu,tu pur SuDUnq,oossdlq
rd .5!rSurrr! 9u!trr .rjosu!
:rlr Jo uolqndrrBu PUr sll!)f Fr
rrqr llP rd::trE ,{turqlr !'r * qtnoqrF t.Dqrq.
2zxt8nLltrss,a.d \rnq!{rpaqyJo
strDedsIur J.I]r
:PFq rou oP .a IIrrS u,:9 rrq rrltu rr .^!urd
!^r sr{ ?u4t snu:t.qr ioj duitr itriSuq llrr n
or rllr:rr rij snrl qA i^Dsrrrn! ,!rns * suorq!
nr aqlnq irtsntuq fl,y'rouuo suoDsDosr s:qr
,rsnorjo suo€r d?r
rqr u, uar pu!.r'qds
*d{
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ufPou 46pur
-u{ rqr u eDrurKr
uD
rsFoPUs :rp p!" r!rl,L/,.IJ
m s'B lufl s:$,1 .uoq pus fu#ulD! ,!q! Ilr
pw
lo.ru:purs:p srrDrtl UroJu' fDpnp
iNEq ltD.tr m sloor :uos ,np,, rtudH \!tM
-u$ 'ulunq
:t/6I s$:!t "3 a) auaH 3n!$ tg!
12002 \mqr:l
Jo sp,u,uo! ttn: r :Err FrDs:a I sh rDrl rrql rr
-n or llntrw? r PUgtu tuBnor:dr n:qr uxqr l3,oo
$rt ou pu. F:o. ${ ou IIrN trfr sf,uruort irrs
''ur$rod $Br€or pur rooutroslr3 !]oq qru
.uoqqrotr rcuodu P,q mq rq8
,oq nuunP zB rfl,q'rr IDEFj P
lqr s6.qr (rooa) qD Er Iq Prn9nrd osP !:^rao!
.$iA\ utr^uP
Nr
^
^Ilualntr
luuol ^Pt{ u rur
Pntr ns il rnq
FUg
Piq uetrsFunru ruor rur:lrP rll{r rrDrddr rrl
a ?qsFD'u' rur:ulp,G:^ pq rlqr (s:tqqor!tr)
\ 1 \ \ . , ri
Nil
rNrn , ll
290
behrvioaora h.ightcned thresboldtorvitfiinlroup rggft$ion, p€rnining Bnups to liv. noE dcnsclyRith
moa .oopentiR behrvior asbener potccrion again$
both pEd.tor rnd otiq lrcminid goup5. Concomi
bndy, homonil hrg€t tissu. chrng5 allccEd gbfrh
nfts {lonsd dumtiom of srffih and pblons.d &in
pend.ng) po$ib! uith sfrc illomctic in.e*
bnin srcnh thrt wouu navctcci rcflectcdin sLgndy
hishcr EQ ihrn in aFjs.Th.n bipcd'ilnn no$ pbtdblr allowcdfor gllatd dn!!s ofconicl'c dicGiq and
c+lo tion, lnd th& |roljng sophisiiotion aboutobjcss .nd $atid (latio.ships qs prohlbly insnmenbl in sdcaion lor ! (htvc d!.tion in ?vC md r
(hdvc c{ansion ofpo$dior Friebl lobe. E*.nsion
of loastlt rnd menory wodd obviouslyhrre been
adqntlgcous, fd .pp.! b s b be msc..r in tl1cs
l lobes
hominids
g.(n rhcnsnc*fid bddd prcaront
r.d dpxnd.d hnposl lobes.
Lakr runi.lopnheci.es pnbxbly s.r usi.g tools
bur wer noi slbwing cltr shndrdizrtion oflorm eithd in 6eir cnoia ofrmls or then nmnfrce,
the
.r!*t
evidena lor $one rmls gDingb.ck to ,pprd
imdely 2.6 IIYA. Il so.i,l communicatoryskills uge
thn sdge rhrn wh* ve vitn $ in dd.i
3€1td
chirprnz.s, rhis muld nor likly havebeenssoci
ried uirh soci,l aFect rnd contol than with hunting
Ldn art Aom Rrtt Hano {actus ?brn, ,-fta
tuiry m"eld.tb aftuENI ad th| d.uk?no' dhb
b'nt
lit ,n brul
aua'aP.ni.s;.h.rutndndn
a"nmti6, it hdks RrcMi M?,qia6 afthrt .fnntul
iN66 aJ
ntua ad bdh allandtn and
','.ttorni
inw
ia
ttuin @|uft, uith atkdiry
EQ.
TXk sbsc iocludd clabo.ltion rd Ngmcibtion
of ttu changerin so.i.l beidvior hentioied ii $rge 1,
More importrnn ir invohd 1 gowing dcpcndcnc.on
sociil .oh$ion, .@p.r!ioi, lid sd{ol. ofplenenhrion in dl eononi. hs!5 i.volving b*in.i.g lrgl1ge bGed on albitdy s]nbols, however!;ni6v.,
rnd ihe hanuFa.bring of sbn. tools to *md.idized
pntens $it[ cLr oSrnive asmirdons beMen.Mftrni tools rnd hskr. Biped,l lo.onotion ]A f y
hudm. The bnin *pmded in sik bot[ absolniely
md 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 dimorphisr
b.tue. dr sdes with x posible incrclsei. seon&ry
sm.l diddpl]isn, pl]siolo3ic,nt, mrtanic,I]. md
s o Nr -G r r E R, \ rNo
. rE s
b.hamRll) The. ,n tump.e,lcd ,h. br.n rd
serns poshrn dcp{ndu.);nd hmins.incrc6i
ig. t(Mrd prtunhon, ind ,hc i4r{d
dudiion
.h'ldhoodto *"u! mdurJ SduJ krninaor
maLng hunhng,rdllcr ig,tu(ngrig,rnd rcp
nve stnllis
il1 in r posinvefe.dbrk sls
-e(conFldity rnd bnln eiftJg.m
bcsccn culbril
but.ily r. ,ht drn, rhr br . ob{.ri.J .onsmi
wca iot trorhcd. As Holexy noEd in this
L!igttrs.bdruorbnm morqmns! &r op.d
@sini{ bchrior ofrmqc ncrl} humr ryF
oped,*heft lxnguge xnd idl mrkirg m$ lilm
rco t,itorca n ,h. +rsr. rnJ rinsrisr hid pd
inponxie ii nxiMiningso.ixl.ohsion
rtrd.ondd
ing indr*d rnd th.r ms rorc {(ss ji nr&ig
rnd huniinslq€! samc Th?rcwa' a rlcdion rorir
Gsld bodrsiz.,bip.dll lgiljlr aid prdi.riv. xbilirid
ro' moEsuasstulhunri's (o€roiocth. 6r[ foodqd
in lhich rcmq
and cbildici
tun). The {.irl
b.hxvioin cnrngs o
dr idpddnr
bndibF
in.d in ses6
siscni hudiiig rid lor piot.dod ofr nor $dF hln
bs. |o! f!o,]$ andFu!g. {ho wE Plwiding smdk
ycsdiblcs. Thl "iritid kj*" dr "huhr *
samcud
olurion' n nLly d Ntrdl..ds b slg. 3.
Cnukg Nhbltatkn aJurrrul :Ailt basd tui,,
d
laaglqa airy tuitrury t-b,l :r5knt ard d.nhfi,t
t b aryh an n EokE f a: n i! |fadna& Eb t i an'bil beiM
b.h@i. l e,flniry andb.ain eah.gnr"i; al:an,tirliry r6a.Mk
k t.ai,?kri 4!nn4&t
ad tdi,f b, n :p, iot ad ;n fo. ! nrct2 t i t, a{bat, and, o,ioldt
stos.3 n $
w b.tifla fiarab,inn {Naa,.dral. It *.ms 'nA
to us unlilcly thli nrjor.hr8€
i!
po$mbl dep.nde.cy6ne, g.$nion, prtuitio., orG
podutiE ageoc.ucd du;ng thn sbgei hving b.6
for 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;rg
slnch in a$.d Lr.inA Nodd lrNe ben *ftn
ly
inporhnt ,nd u.dtr frn\' stingenr soci,r conMl,
as cultu.l conplexities wft o(udng in noft complex md stinDlating mrtd,l rnd soci.l envi6nnen6.
Tne mjor neunlogic.r drmgs wrt piobrbLyni.d
l1tt u:rNrq rlru:rteslP rP \ .qr u 5 Pneq'u'
rq plnoqs
lPos ruTutrd.Pu, 'rFrqnd
9351
)ts
to
uI rP,umoq s:q!o
lsropulqr
Jo lSotoqdrou
pur s.unto^ rqr qroq Jo IPD5 rurPu.drPu .trnotr
u: pForts IFJ nro puD(H-u) snJo euo u$N.q
suqrP.qI sPru .q pFotrr.qr $
ro lsroprr :tqssod
tPug m8J uroErrd istru
PUE
3qr 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 pui
stsot rD rDopu: or 'otrqr uo +oa or nuucrp .!oql
ro qr$ot rFJo e9n\r u, soqr Sul5.norurJo .doq:qr
u
urD uoru:o tra pu? lq8noDqr :q o rdu$.
no^spq
r srsNrnt SunF.uos rDqr rqr !!ru!
rrJ .qI
,{pDs nrluryJo Il]rrm s !tr,fqo :sqr prr.tr rqtu
srqnsr nqro rr$ rooqs:uoq r
'p$prt prprus lpFlduor $,{F!r'rgns u$q aiq
ro! s,iFruriod Br FNr rsrrrPlt tqrjo IF tu9!r1
orronrs ag orra
srsvrooNg
'rq (rE r?l]1!P tuu4qtuJ 4nut
inojo Fnpdd PU' rurEosu, r91
lnJrtrpuoa irrgc
$ ,(tucu.r pu? '&alll.d ,ttuoqqo^r hojo rnri Pui
Dnpod trqr$oq ! uruq upuD{ r,ll uoslttddtqrJo
suoD4nu
qtrnu urtdF Ilqrgorl lrr stu"liqqitiu
renFs a.J r uo lutluidiP uo!!r u' atl D rou Pur
,proerxl 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 lPrusno \sruuru'q
Jo rDrd tu, lrd { rptsuotr * sqnqDr piohr!:q
5runP luos iuoDrPlr
lqld !^4i9a ro
'itd'uun'j
plp uoq?.xrdd?iqrqn
loor ruoF't$dd!t
u.$ard luoqoudot lep:drq
'uq"u!8uuoll]r sErPff 3!oq
-.rdipr prour,lr,l pi'urduorr reqr eturqr lBrsotoi
-niu ras rtl tu4rrulr+ 4t sr|']ru:bgns r ipqs
Ir$ 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
lldrum
Jorou
fru.r nm! Nll PUt
fuE luoqu.u
q' u' osP!'q sdrrtsuoDqr
u:guEq: r
Pr.Frtsolnsm
itduE rou spltr uflldnorodo:ot
u.Susqr v sun
mu ptrBoFnr I (l:qr pp ru) .rrrdo uE: s!.u
oduor rsrlrjo :!oN rtsntq ro br:lqo 3!r,6rr
:r| uolouorot pp:drq
'd'l' uornrd Jo ruru:lgr
r 9r !P'L'uo,j rqr Jo qu:loduotr PiSotoq&ou
rrlro d' ,rtuiqr Auh.UoJ r|td toolBqr Suqr.us
$nf pu tur uoqnpr prrrdort i'a D:d$ Frrrr rnr
joou, u? ur4 sr[|t 3Bqu'rq uEunq:qrlo uolqon:
r sq .l
.qr
lriEd rlrql u' {o'ls or
Iqr
Podu4s
isprrr0oq
!o$u 5z ruoc Jo rohE,i:'] PUr sur4 r'{r rn$au
pw .lrsqo or cnJo rdo, Jo :!o rj4 rr flnoa SuoI
ur3 suuq rno ntDrrd
eq tu'{r.u iuq r
'}k :'txi:P nr ra lFsottr aorl
u, Pui .uo{nlod P'uruoq
Jo ruiu$*e rno ur 1qury :q ol DJ.id :a rnq ]I!q
o! ud{ i,tl@ uEr $Dlans q:nu rsr{ qqN+Bur
r' r'
)uo trql uo rur?trP .I!I rr"u PFo{
PUr't
(uoldun$r no iuE ur :rr:Jrp
.pNru8ro ,o rrpro
sv i96r00.0D,r00m0 0 srp+rEs arq *rEqr flft0
,!Fu'uog t$$odJo :tulrrd
:!I $sB:opua
00I
rj tra lutrsB ruoDrtnrp: mo Irqdun ol
Jordursr
!dq trqr
rnoqstu
rnfiFaldr
^sq
!$tr p'urroPuiJo ducs lEPiu:eld ino r h€ sq
rrqr\ 3pruru.q pnpb,pu! 000.0009u:q o rno sruol
d.liDuit id sno, 0o9su q srolrDur' 00o os iruo
1!'lrjruudp pr!l r^e'j ptnod sp'rpNpur IIBU doq i(rn
t! tqqbptr ,{F r !e lrFs o) snr,{ 00o omJ Nd
Ir Jo rnno: r']r Dro 'uds Puoor.u:8 luu turnp
sFostu,trnfddr oosro rzu (rM) looqndod Frussl
&!koi r pri lxJ OZ.{Eslo llds uo!u:!rs I ruB
{ .1
rtdus t s
lllleurPoE$ os s.
rt
1!4
fauq
s,
Pod InsoJino rrqr rd$E ollnrgrp,{tnlD@d
s.Puru usunq :qr ur
r' rnsoP ioj rs.P IrN!
rn,{ro spuusnoqrjo
3 uJ ,:jds u'qrra.pnsNrn4, rn't Fnps9 rou llqz
qdd r:a s:AuErttr
Fiodua $qI T:ds or os5nsou
:qr uqrh $'Esoujo astl$od :qr rno rlnr rou oP#
!'q lnonuquor,ttiruJ se eSus .srqr PrEaJ ostD]
!r PUr rSrnAUEItrqo'tuls fliJ Jo $tsrr )lI] or rDnb:^
P ss{ uruiq :qr ro.J:'] luoonloDr, r'qrlorFd lddo
ur sEprtx or n.ddB Iusu nqa ssaurh ol prq ro
lqr Pft 4qrsod !, Pn r&nguq :pNu ru$. FUo!
utru :Fur E ir8as Plqr sllP r r!
:q oq { srnftrlar no qrd ar'ssrp IFsrdp 1,!\
Jo $ot r
Jrytn .{n rnoqrk 1{ntrDsnqdrFrnr/Iuoq
uo px?q asEJ$p rDruonr !r s P:untop rq
^trns
$ou ptnod sql {366t ft:qruu:Fr iq prF:Fsns $
'pon:d :uD s.Ieporor trqrlorFd lddn rqr roj trzs
uBq ontosqErno 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^rp
rrsDr pu? ltun9nq luapp Fnuu ur rutrullo
ur rn:qrD s II sr.loDrFpor (unrms) Fp,u
Bndur$ pn'$Dunoss lororsosus u tmq pau
x$or Ip$$ qr to ruamugr pur ]zs Jo srsemu,
tLz
292
l l o frN rn [^ uoL,\br s: Sur L Cr Nr R,\],Nor r s
et.r. (2000)rnd HanMy {1972h),xndHotmy d at.
(2000)r8lding irs volune.
Whne ,I ofrhe .onsid.ndons abovcapplyto 6oth
ft.ln
xnd Erhiopim ender$, we brlicre re hare
d.{dbed mcr oftl].n, a hrs lobi6(1991). On tn.
other hmd, hominids suchxs KNM-ER 406, KlM
DR ,107,snd KNM ER 732 shouldbe r-sndicd Gnd
rc cRst),si.e RLHS origin.r ..do.xss wec lcft in
N3iDbi md hxvelonAsinc. d.idionkd. Tfic dcscrip
tions in thisvolumere bxsedon pla$i. ap1i.a6ofoftcn
poorqu.lity KNM ER 406is6lLd sitl] *rcmethad
mrrix md req*es nuch non study tlrolgfi nodqn
$.nninA dhni,tues, $ n is very u,JiLelyrhn tic ma
di\ cin .K b. rmor€dwnhout seiioudydmrgjngthc
originRlcnnium. Ammiak rclune esdnxe woutl
N.wr frnds, such a &4,2,rtry,/25!A,
and
othcr fmgnens llon Loneh,i, shodd rield, rl@ugn
mod*n scmning techniqu*, r good oppoftuniq td
lind an amnte rclume, pdionrly in lieu ofwhir!
{2003)asAsment ofpossibledisbnion to rh. holot?.,
(NM-WT 4m00. NatorsII acent diKoveiiessuch*
A-rin,Srkladb"ala
rd,4tdi?ithauw
yit]lj6.rh
alumcdc and morphologic,l fertufts thri wil rscr
in undcisfrftling ihe e ier phasesofhominid baln
cvolution.To our cuftnt und4$rnding, the angnenG
ofticsc cnnia arebo incompl.r b providesuchinfom.tio!, tut thc future alwys holdsprcmis.l
Thc carlicn ,9,,a spccimcnsfrom Tmzmir (OH
7, 12, 13, 16, md 24) aF paniculrly in nced of
fu.hd study,huilding on Tobirss scholsrly rftrdse
(1991).Tllc dngiids tfrr RLH nad. in th. edly
1970shrv. su.ly dGiftgEtcd, and the plR*ic Eplics
i. his mU..tjon .( ofpoor quality. Similuly KNM
ER 1470andKNM-ER 1590should6. rcst rnd i.d.pend..dy studicd,p.d.ul'ly uith re8:d to pehl
i.l asymd.dy pan{ns and thc po$ibili9 oa csebml
hemispheri. spc.i.liation, rs w.ll s the Arcc,t crp
ftgion. Th. fomd is ofltcy ifrpofr?ne q:rding ques
tiors rbour bral. r.dSanizatioi in thc Hona lirc. h
de€d,L\e safe rypls to KNM-ER 1305and yNM
ER 1313.Th.* Mo endod* ponions arc pdioLiy
di6c'ni to xsign sirh my 6onomi. closur.giveniheir
u.usu.r norpholos,, drfiou8h @ stmd conEdenire
grding ou volnm. derminations, whcthd thcy m
h! .rly ,v ,, or xdunc.d au$nlopith.cines.
The Dnanisi speinens of 1.8MYA ar cxtEnelt
inpotunt, both sith rcsped to .c.uoe rclunes rnd
po$ible norphologi..l ftrtu$. Wc look fonysrd b
rerding studi.s on th.t pd.on.trologi..l inform?rion.
Fron ou omhrion ofrhc lrous phobgnphs
thesespecirens,se @ ccaijn tfi* rheycrn be s
endmst, xnd would be *iling to do s.
As fd rs s. m xu!., thc Abpucrca uni.6.d
sp,i. (4 bd J) hxv. ynldcd .cune volums, ul
then norpholqic,l .hdcernti..
ar. undd studytt
Arsu,gst rem. Our xft.mF6 rog.in infoh.rion ahd
ihesepo$ible endossb hrv. nottc.n mswcFd. Cid
rbe psspcc d[i theit inr.rna] hblc6 ofbon. ar. pe.
iadlypeefl ed,rbep.l.on.uologi$t lppcriEs snou|ll
inded be rvheftcd.Ths. rprcscnr ! najor hiitus in
ourundeBrndinAof possibleN.anddthi an@snfrnd
the EuroP.anhoninids i. g.ndd.
The Steinheimhoni.id (Gemanl) n )d anoinc
hiatus in ihe donai. of p.Loncurolo8ical .tudy.41.
thouAhan rccure rclune d.r{minlnon vil pbbrblt
rsulifrom nffr srnni.g ddccs bd a*omtiw tcdniques,n is prelenble ro hrve rn r.tud cndod* oflilicon nbb{ b p,lprE. lnd.rd, .En x dGro.cd r6id
k pFLftd to noihing, which is ou.uftnt sibdiod
Similar connens cm be nxd. foi rhe wu-pc*md
P.talona (Crece) and Cepnno (Ir,ly) sp{in.ns. Nl
of th!* rrc inpotunt lor x bdr.r unde6unding of
Europ.an honinid erclution md th.ir vrirbilitti
N.wr Indon.si,n 6nds suchxs sn a Grb. ( .l20031!rc inpofrnt rdditions to or incrcxsingsropL
oflndonsian sp.cim.ns of,!.,,,, raaJ, which ro d.E
sfi@ litlc in th. q) ofeithtr votln ric d mo.pha
logi.ri iidion, in aolutio..ry pornti3l]y int Gting
ft1..t ofthc crclution ofthe genus,9,,,, md OOA (out
of Aliict and MRE (nultiftgion l .volurion) con@
wdes. Wc E Ery lotun.te to hrE hrd th. oppor
tu.i9 to de{iibc Sm 3, th. Polo}! honi.id ofNYC
S.enl Cbinesehoninid mnir (rside tom rnt
olddZh.uloudimdtcowriesorigin,Il.rnedJ't,44dF,i,;4J,r,
sch a Lr irn, Mrbr, Yudim 2,
Nxnjin8, Hsim, Snandons,Dali, mdJi.niusbxn, hlE
nar ben endo.r* to our knwledge, a.d r f.w mxyhc
pdd.'nrry dieicolt b cndocar giren then n,gm.nhry md derofred .ondition. HopeirB ths. *ill b.
rdd.d to our pdconcurol%icalsrmplein ihe frrtuft.
Fin lly k notc tfirt theE m sftr.l inpo mt
gapsi. ou $udiesofNRndsthal endocilb. I. pdicrhe lolih Se.op.*orc hominid cs.i,l ponioG
'na
.ould be .ndo.6t, &LCing€ry inpotunt rclunedi.
md morpiolqi..l ddils b thc abundrntm.dovtrsid
okr rhes. honinid!. FuntumoE the Middl. ENh
Ncrnd.trh.ls ofrie Mount Camel and Shanidd crG
hrE nor b.en
by us given pDblems of ,Ns
"udied
.z3r!3rior ddnsFduqruv ,(,rd I urv p!84
! euwqrrdoFn$ NqorJo srDidm Fftftpqr :!rr
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'91 -ibrljr1r
t! dd utrrcld :,tro ^:N !di(]
uo3 pu! spo,lDrjrroanp^3 uElg nauDd tp,
rFJ
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urelqluqxJo,6un!pl rqr'136r 'cd $qt itll rholoH
9n-fa ddr$rd ,nrslDn PFro asqr €N
uonryo^3)qoqo.{su}urH
tpr ,J sxd v
lo
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'966r '.N /ihotroH
'fr s,fr 'Fdo4lvlnc
)glnrulrjo urfir
FN$t rt) Pu ftnrrunuuoi xluod &6r 'M5 srDH
6rt si/r9a ros\tJ lor@qJ d:! tr{c z,Dq
tGrrd pqru xJuoiFrD'dsid
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IDNINJo
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r ,LqvT{ es ..I-r unH
$. r rqlFd x r '{r,iror5 b,qrF{!rl)
id nproldwot
sdDo tprluo$hrra t lxmtu€qJ:utuoont@sErq
ulunq Jo norqr rDr$uE I pleol s66r Tu rNio[oH
iruY Pr 3 lsFq :uI !$.to trd N
ruu ,anu oaaHta vtztq Eod aql 536r'T[ ]rNro oH
9rarrpdoqruv turl arJpur pu) sFpou lrDns
q'36I -ru {B\onoH
t*4zr:.,rrdtu
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uuq 616r 'ru t*\orpH
rI!3 .p. rnol:ul uoDtrld: pn'uo'l rADJv pu! uoa
tr
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rI v ' '3 6 I ' ( {I r l
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-i!dr4uorords 1002 lrqooJ,,ts
ssrd rmiun Por*o rt ojro r4,(/'i
'a9r Lrr dd5 d,{l's:^ npeJxo ipej
xo rBdu4@r urpot^rjo uoDDDdsiqt.p, fJNrc
uDD.Uv
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a.N qz6r '11 rLM[oH
$r dd\erJ uoDqrv/rqlP|v :ornqJ sluudl.
ptl,r'€uolrrd Pu! Fuolrur] iql P! ! qlnl:q
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'9r) d
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.r iPtenvldaprnv.6b6r.5
r\rvbJp,F!r,
rdns s uosdu$ ,q^luut,o
ror:^o,l .J,!!M 1I$ssy
sarNanrrr{
z16r .rN ,ixiofioH
ofr arr19rDspmv,$luuY aaa{s $@!N
il
iJ suld nl Dluud
)ja'toN
ur uoaqqi
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uo suors rb .uos .696r ',l! {imllor.l
Fniujo sriulrd
.6r-rrar$s r:t )oNqrqxrtdoorjouooqq,
i,l, pu! rotrus
ud.q ss{)us } prm' !!s
r"!Du
.,rr ilhonoH
l96r
t(z
!€rr'u ,a,'J FtruaruD unPou
strDrdsru!]I|rp rFra ss'!
fu} ,sr,l roJ$.4i!t$iP
:uo ro! io fqrrqa \rur!^p lruo{nto tr FrFlpw
rNJ.lu'pue$ipun no u, rurcdFro^d ruruodu
ur r,t or,iplq * IiqJ !4ru ftruru!5aJ,rqr pE
sr r.N l x?r 1D l r os
2<)4
H u rrrD IN Doc^sr s: sor L Cr Nr R^L N.r L$
DC, YuadMS (2003)Morphol
.himp.rcp
nryvisusl rir&
tulinskyLB.1975.PlimeEbGi!aolution.niS.i63:
Eosuiaiion
sbinBscB.r c8d.fl..rtd,hIhry
ofs,-, rr;ft lr:
prcdii.d bdjn *-
posio! in culy hohinid *oludotr. Atr* R- 273,4:59€
Kcii Rc. 1999.Thc Hund Cds: Hunm Biologiel.nd
Culunl 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 Rc
orwh* Miks Ur Hund. London:Lid., BrcM.
Nnhinuft T, Mirmi A, SuzukiJ,M,
seM ofth. hrlu i! chinpez@ i ua ?@. NdA.d
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.{ ldrl
qsuiziioi
ii .dbr:l.rlu,
on.l Hum Evol ldtal-7
T06d PV lcol.OlduqCoI$,vokaA,aB Th€S
lndoqss. od Te$ oaa/?'? ,arib cmbndg.:
V.[ur A, LordkiprnidrcD, RighniE CB d z]. 2002.
ns skullof*rlr ,9?', f@mDnuisi, ccorgia.s.i
WaE ACH.2nor Th. eEb€lluo,!d cosnitiE
don in PlioeE and Plckb(nc
honinia3. PhD T1l
Pidbo S. 2003.Th. mosi. tbr t ou s€looq Ntu
wlxrcTD. :00r Erly hominldFDiwEiDqdisdio
Plss TM, cac&! M, hu.h6j, .t.1. 2003.Usiig gc
nonjs b id$d& hummbnh .pdiiadoB ir J Pb!
w]rir TD, ns6{ B, Decusr D, { .1. 2003.P
R3dilsllyLB. 197a.]lc fd$il .tdei.e ofrntnop.id bnii
rbludon. An J PnysAnthrcpolalis-28.
worpofl r'L Lqqq.Pns/dhtupology, 2d .d N4 Y
hei.
H,d, t'?r4r ft.n M ddh aq'h, ro\iop L Na
sarNrwoHrrssoc
sswnroA'rvrN\/ucoaNg
296
E l n o cR ^ i rrr voLU ESoi r r Lr Fossr Lllor r ^r !r
NorEs FoRAPPENDX1
Ruq CB, Iflnlius. !.. md Hol,d,y,Tw ta97
Th. cndociinial voluncs ror rhis hblc te li*ed by d
ononic gtuup as oftndy dpi.sed in thc liemtuB
dd @ngd rlphrhetic,Iy sirlin bqd gcognphic
rcgion6.Our u$ ofihese m i. this Trbt iloes noi
n.u that * conor sith rI ofihBe &signatiom.
Wher tllc'sourc'indicrE m "RlH" th.t mems
rhe 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@ Eve
lurionndPEhietory,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.ft
M $ found in ou Bibli%mphll md nostl), Gom
ns.nd
enc.ph.liation in A&ban.
Hond
13717117blsupplm.nhryDrr) Th. SM
SM.l voluhss c Eom Baba,,t
2003.Hom
cJv;um fmm fie Plei{Nne or}q
W. hac ddd.d
sci
b povide r choiolo3lol
e.h.n.t nnDl volum.to pftmir ftgrtsing
ni.r votuie €:jn* tim.. Themgc ofMYA! 6.
of rh. honiiidr e tooyi& to r(ept mFrila
middl.vdua, hJ wc hopelh* fie.e dxi6 wili
noft rc.ult s ns dating dhniques beo@
rbk Mn! ng, npafrflltr E:nymdo(triJ
for th. Mrbd (Chin2) cnnium. we m'nd rd
citation rlvh.rc in th. lihbE
thrt w h1E
l6z
nll
lo snl\nlo]\
HTII
H',ll
I (I']N
Hl!
H'm
mrtS htN)
H']II
H',tt
l l l l \n3od\3
294
E N D o cr^N r^ LVoLUUr sor r r E
APPENDIX 1 Pd
RLH
R]-H
RIII
RLH
RLH
RLH
RLH
RLH
RLH
PIJI
RLH
6 62
H'III
(P?'!r@J)
'
ud I xtcN€ddv
s(r N rNo II lr ssod 1nr r o siLi\r lo ,\ lvr Nr n r o o Nq
300
D N D o c^ rN r^ LV oLUsEso. r xE
I0f
NIJ!!roa
O3 pw $unlo F.Iuropur:tFav:ll
s a rNr r o H l rss o J 3 n r ro s x rn lo l\
Dd r )XONSddV
M\ v ! . odx g
vxvJ ,\s sshrnro^ rvrNvucoaNg
co sasl'rvNv rvcrrsrJVJs
XIC N EddV
30.1
S r.\rrsrrL \
o . E No o c riN
^ N.\Lys r5
Pmvid.d hftin @ thc initirl b*ic $ni6tics for .I tl]e
hominid endodo;al snptes re dscibed in this bool
Tlcse sfrtisris.rn b{ amnsed dijld.nrg 6utwe.hos
thcserrious combiiltions b(ru$, by ftlying o' oF
snsual Nononi. plr.n ns, w wdc .ble to incnxse
sanpl. siz6 wiihin tdr, In addition b nch Rnd$m&d dryiRrion, w include ninimm lnd muinun
Drucs,the coeficGnr ofwirtion (cv), sk Mes, md
Lunosis.iaw rs.nc thr al honinid populationi en,
do.nni:l capriries N.a bsically ofr nomal c.u$irn
.1or, the sdo.g ddrions Eon $cn nornsliry
might ,ignsl cluesr8ldins sanplins md tdononic
6otndncdrr' hd merhodotogi!:l
6su3 whm
shcnngii\ompt!F endodiB.s,mJrty rhedri
rowcvs ror rhcNsmdons(soto)andlt ,,t
&,n groupr sugg.$ eiths smplng or ftco's
bias,*h.rtsti
A. afri..nu NnaH batin gnq
Cvs thdm bo( in finesith exTeddionsror.dt
ninid sm,l dimorphismmd poFdldon ruidioL
b.li4. th. srm..ppli.s to }I ditu witlrcut tt
H tudatEatn,Md H. sorEna.
Mosdynon our findings,bd iicluding d'e
srcml otherwoikc6, on th. ,l1:md.tion oa
n,rl.apKibe5.!( popon m rmg. .ndo.trin
petr/ lat H. e?t4
Ior @mplq pop'nrtiom uirh v.ry hrg. or very
srdl CVs (i.e., depding *ongly cnher ky Gom
xbout 10%l suss*t .lurion is rdvisbL. Thc CV for
I a/zzaa of16.1%su8gc.sa f.nltdinorpfiic smple,
or r.ombimtion ofm. Wc beli.re tn. 6mz! ro he
th. qsc de.piE ihe r.ant Andirys oaRenod.l. (2003)
rcg,rding nodem l/,'o,lile lmrt dinorPhiso ofth.
femonl h.ad in r arz,i.
On the orhd fiand. tlr
CVs fot "d\ Hona dkht e 76.296,and 14.996for
erly,9,z,, bccause
thesesxnpl.s ontrh n'ntip,c m
nthd thb stungsmst difiorphism in..doconi, cr
y. m rot ddfrins th* cvs dnD
Prit} HlW4
morpholosy,onb. rhd th.t night haE a se i' $nri:
rr.s usro qustion rhebrsisfor ttrc hid or towvducs.
MoE smpling uill ofmuse esolE theseqne$ions.
Sinntty [,. 1@ Cys fot 1. rob,nL|/A. baii ptob^bly sign.l r .onbination oflow smDlc siz md bi6 in
a.nddldla''
tnzr s lNe
rhattbr nodtrn,9ozr gl;,r oatl]. UppcrPleis
Th. addition of No shaljsh (npim (3 md 6)
ieff, le$ ibrn 1275nn, both psb,lly Gmar6,
.bary€ ou birs rcgardingrheald p:lemttuopo
'.hnhuf
lhtr Nemdcfthal hsd bdi ii {gcr L5r
As the noddn ,qara srmpL is Mqusrio
'rM,
nosdy male, rhis supporition Equn* hoF
Sim'lrlj lhe,k sia6 andlaro's 6g!16 u
rppoxinat du. ro impdfect srmpling or cl6di
of d: ln g.neral.*Jus gr. e' or 16rr rhm +
skn l signidcant
deprtEs. W! donarsuAse*myr
ofrbes.sotisticsto ddehinc .ither thccon 6d
wrcngn.$ ofbs I n $ic6l these6gus rors
rnd kfrGt apped ta b. sugg.sinA oveFdl
s0t
r Y rJ r { s r Nn lo ! \ lr rNv n s o o N!
:EI,rq$unlo
(n'to?
,tu4Pru.H
\&,liapr/ tr4rq ,pn[!l)
,',r,?]'P{
a8'1p' P/'
rxrdlrv $Pnpun
euD.qrrdoFnsiv [v
(rr"/ sPqru4
,@!t6 y,1t8 H
I,n,&at ,tu?j'?tu b,otp w)
(903r'rr3r $pit:ud
y
y
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,
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ro s ns rly Ny 1! c rrs lry rs
10€
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drqt do.
a|hootr@nqIIqdroiq!'@?,,!P',yl8cu4,E4J
r ulru:unqurs Jo )s.op!! iooz s anbrr r 'SI\I
tru',{.v sr{'c t ruqsl)l ,1u liaotroB ac pP9pNorq
puruSjnr $$.tup
q!!q :8'zdF'l eNqlrn?z eP
udvus<r ur{ !' lpuouq$uC
606r ) truNuPos
u;p!c trola,N u.t\lrsoj t96r nHlatP]! r l:tma
.961
6zr:zarodorqrLyxluEs n! :rFdiqC q :p
rrnos
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ip !Fqd,u:t rr6r 'I
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'rr$oj
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'5rs-61!:6r pddqruvt (r+oc)
lucs n! :0s?r1a N,lrp qr$oj
'uu
rz.-lrz:e o)uv eN) r'! rLY rtr,{Fw rtrFuordroo
Iq ?&,r urpoq pu! rqJ! u' etgdd lguur pr!o{ lur
rduo3 666rF! HDsusso.ttrj
Tsl |!r {D) trFiis puuoLt h,t)o qrll Pn $nrds
rus iqr
ltip! ur ,t,'i uc@dq@ s, itrFui,r'
Jo
r|oi.
rr'a#ais
jo nqs {neFp! s! !O 116r 'd )j}rg
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uorrrls
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urD
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ru{ru ! uoDs4rqd?ru3
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1.onlll) Puqr Fqrl nPirbq euuoq $P u:r@roP
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6s z:eer r{mN duor I $tuiqr
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pu! nuuq&! roola'qrFr,tc
iou:u'dop
sixrdf r00e'p) \i[iz ,vq],.r'c ivrrttrDrqrs \ lunuv
ldmN duoC i lrFqus !:rqrtou'
4, ir pxEru uorir s,i:oiq 666
53u n iH srqtqor{'n pt4fl v D
P0 rnrnql
aaJrc sunJVuaJIT
J 08
Bnd6erd Dc, HonowayRL. 2002.A,ymm.rly of ih.
fbrdl .ndoftrniun ii modni huna
iricIPreEriotrof aolil odc4s. Am I PhysAnrhbpdl
Bomrs€ TC. 1937.The 6iobgn,r rid .hbnolosi{ir E*
rdon ofcdry homiddi I Huo Ercr 1615717r.
BiNi B,W,rkiA, wirdCv,rcakl
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1966. CEniar ansciq rtrd nuo! rum
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Hono{ay RL. 1967 Thc crctudon ofthe hufu
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bnh ion.
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Thr crctudon of rhr piinar bnlnl
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En luv uoDqomrtll'ull rol {o!F
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tr!
314
sEnh GE. 190a.Tn. moehology
of L\.
daft
(Icbtnr hcoisPh€G in nrn md
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q
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wxlkr A, lf}ly Rr, Hdn JM, BtrrE FH. 1936.2J
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i'!flh?nt%
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Ndrc
wcidc@ich n 1943Th€ shn of&",tbrt?u prin rtir A
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rnon df dbdl{
@nw Thc (rher{ forium$ r tun
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