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Species Recognition in FlumanPaleontology

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c r i t r : r ia l i r r d is t ingu ish ing s1 ; r ' c ics in th t hurn i rn l i r ss i l rccord ar t . < lc l i c icn t .N lo rcovcr . in d iscuss inc s l tc r : i t ' s d is t in t : t io r rs . in te r - a r r r l in t r : r -s1 tc t : i csvar iab i l i t v a rc o l i r :n con l i scd . l ' he c lccoup l inq o l rnorpho loe i t : : r l l i om tax icchanqe in thc cvo lu t ionarv J ) r { )cess n t ( ' :u ls tha t thc r ( car ) } ; r ' no a l ;so lu tct : r i t c r ia l i r r rccor {n iz inq sJrcc ics on nror l t l ro loq ica l q rounr ls ; hou,cvcr , ranqcsof morJr l to lo t i ca l var ia t ion in c los t : l v rc la tcc l spc t i cs in the l i v i r rq l lun : rr to rnra l l v ovcr l : rp subs tan t ia l l v o r coml t l c tc l t in l los t charac t ( ' rs i son tcc losc l r . r t ' l a tcc l spcc ics cannot bc d is t ingu is l r t . r l o r r t l r r 'b r rs is o l 'h : r rc l 1 ; : r r ts .H e n c c t h e r t ' i s a t c n d c r r t : t t o u n r l c r e s t i r n a t c l l r c n r r r n l l e r o l ' s l t e c i t s i n l l r ef t rss i l rccor t l , u 'hc rc on lv c l t a r os tc r i loq ica l o r c lcn ta l r l i s t insLr is l r i r rq l i ' a tu rcsc : rn bc uscc l to d isc r im ina tc l )e tu ,ec t r s l t t c ics . I t i s thus in r l )o r lan t lo ( .ns r r r (

t l ra t . rvh t : r t 'd is t in r : t n rorphs r lo cx is ( i r r thc l i r ss i l r t cor< I , t l r cv l r rc no trc lcq : r t t ' c l to the c l ) i l )h ( r r { )n rcno loq i t : r l s t r l r t s o l su l rs l t cc ies u r r less t } re re i st r rn rpc l l i t tu rcason l i r r c l r i inq so . I t i s no l . l ( ' a r n l rv a l l t l r c var ious morphsd is t rner r i sha l r l c in thc \ l idd lc - ro - l , i r te I ) l c is to t :c r rc a rc ger rc ra l l v su l rsumet lw i t h i n t h c s i t t u l t ' s p t r : i c s l l o m a . ; a l i t n . r . S t v t r a l r l i s l i n r : t h r i n r i n i r i s l t c c i t s a r crcprcsr : r r tcc l in thc l i r ss i l rccor r l o l t l r i s l in re pcr io<1.

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Introduction

Ovcr thc pas t scvcra l vcars inc rc2rs ing a t tcn t ion has b t ' cn pa i ( l to thc scurch l i r r pa t t r : rns inthc human {bss i l rccord (c .g . , Cron in e l a l . , l99 l ; R igh tmi rc . l l ) f l l ; l , ) l c l rcc lec & ' l ' a t t r : rs : r l l ,l 9B2) . Thc rc l iab i l i t y o f anv a t tcmpt to rccosn iz ( 'pa t tc rn , l ro lv t ' r , c r , i s r :ons t ra incc i by thcaccuracv with which w(' arc ablc to rccoqnizc sl)ccics in that r( 'corcl; this is truc r,r 'ht ' thur r i l rcsubscr ibcs to thc v icw tha t sp t :c ics a rc ob . jcc t i v ( ' cn t i t i cs in l in i t ' as w( ' l l : r r i r r spu t , , ( ) r ! l l r ( ) l ' ( 't rad i t iona l l y , ho lds tha t thc p rac t ica l p rob lcm is to c l i s t insu ish b t ' t rv t ' cn so l ra ra tc ly ( ' \ ' o lv i r )s

l incaecs w i th in wh i< :h spcc i t ' s a rc cphcmcra l in t imr ' , i { ' no t in spacr t ' . I r r t ' sp t :c t i v r : o1 'on( "sp o s i l i o n o t l t h ( ' n l t t u r ( o l l h c t v r , l u t i ( , n a r \ l ) r ( ) ( { s s . s p , ' r ' i t ' s ( , r : r t i r n \ r ' t r l c l i m t - s u t t t . s s i r r .l i ncagcs o l - spcc ics , a rc thc un i ts w i th wh i r :h onc has to c lca l in t ry ing to u r l ravc l th ( lscqucncc o f cvo lu t ionarv cvcn ts . ' l ' hc dc l inca t ion o f ' thcs t 'un i ts i s thus o f ' r : r i t i< :a l co l l cor l l .My a im in th is shor t L -ssay is no t to cont r i l )u t ( ' to thc c icba tc ovcr c \ 'o lu t ionarv mcchan ism,wh ich w i l l doubt lcss wc l l ou t l i vc th is ccn turv n ' i t l rou t mv hc l l ) . Rathcr i t i s , I i r s t , to [ i )cusat tcn t ion upon thc obscrva t io t r tha t whatcvcr thc cxac l n2r tu r ( 'o f ' cur rc l l t l y popu lar c r i t c r iao f spcc ics rccosn i t ion in pa lcoanthropo losy (and thc l 'a rc ra rc lv i1 'cv r : r a r t i cu la t t :d ) , tojudgc by thc i r rcsu l ts thcy ar t : unrca l i s t i c ; and, s ( ' cond, to po in t ou t som( ' o l ' th ( 'conscqucnccs o l ' th is lb r our in tc rprc ta t ions o f thc la tc r s tag t :s o l 'human c \ 'o lu t ion .

In thc i r scmina l papcr on N{ ioccnc homino ids , S imons & I ) i lbe am (1965) camc about asclosc as any palcoanthropoloeist has yct comc to prccision on thc quostion ol- how oncprocecds to sort asscmblasL's of fossi ls into discrctr: groups: " ln ordcr t t i t :stabl ish a val idspecics i t shouid bc ncccssary to shou, characlcrs in thc avai lablc lbssi l matcrial whichpurport to bc ol- thc samc masnitudo as thosc rvl i ich st:paratc rclatcd l iving spt:ci t :s"(p. l0l ) . But although fcw havc sccn f i t to quarrcl with i t this sasc advi i :c has bt:cn hrinort:da lmost cxc lus ivc ly in thc b rcach, cv r :n by i t s au thors thcmsc lvcs . l ' o r w l i i l c S imons &Pilbeam specif ical ly urged thc cvaluation ol- intt :r-spccics \ ariat iol l , rccr:nt

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pa lcoanthroPo log ica l p rac t ic ( 'hus bcc t t to l i r cus i t t t ( n t i ( ) l l ' n in t r i r - \ l ) ( ' c i cs var ia t ion . ' l ' l ' r c

qucs t ion is , in c f l f c t , most ( ' . J l r tmon lv rcnc l t ' r t ' d : "do t l t t ' l i n r i t s o1 ' r ' : t r ia t ion w( .s ( .c in ( )u l 'i bss i l s t t xcccc l thc l in r i t s o1 ' r 'a r ia t ion \ \ : ( ' scc in sunrp lcs c l lunn l iom t ' x t l r r r t p r in r : r tc s l )cc ics?"Indccc l , thc t ' t : l t r s s i t rcc S imot rs & l ) i lbcanr r . " r 'o tc huv t ' sc t ' r r a s t t 'uc l l f lou ,o f ' thcscs tha tdr tcumt :n t thc rangcs o f ' r ' l r r ia t ion shor ' r 'n l rv nunr t ' rous chr i rac t< ' r ' s , n ros t l r c l t ' r r t l r l . in apc : rnc lhuman skc lc t i i l s : rmp lcs . . \ ln ' ros t in r ' : r r iab l r ' . suc l r s tuc l i t ' s h l rv t ' l r c t 'n a i r l t ' r l l r t cs ta l r l i sh ingnt i rms agzr i t rs t i vh ich thc var ia t ion s t ' r ' r r in , sar ' . l rus t ru lo l t i th s ; r t r r l t l t ' s n tav l t r : t ' r ' l i l uarcc l .H a n d i n h a n d w i t h t h i s c o t t c c r t r l o r v u r i u l i o n h a s q o n t ' r h t ' t r i u r n p l r o l ' t h c l u n r p i n q ( ' t h i cwh ich , in la r tc la i r l v rc jcc t i r re thc 1 r 'po loe ica l th i r rk i r rg t l ra t l r t '< l t ' r ' i l l c r l pa l t 'on to loq l ip t I t 'ca r l i c r v t : i i r s o f ' th is c ( 'n tu r ' \ ' , . lu rs in cc r t l t in c2rs ( ' s l ) r 'o ( ' ( ' t ' r [ t ' r l u i lh un c t lua l lack o l ' r ' t ' l r l i s rn tothc oppos i t t ' cx t rcmc.

I t i s no t harc l to sc t 'n ' l t v in t laspcc ics va l ia t ion lns p rov t ' r r so r r r r r t ' l r n ro l t ' l r t ' { r r i l i ng topa lco : tn th ropo loe is ts t l t iu r has i r r tc rs l i t ' r : i l i t ' r ' l L l iu t io r r . l "o r ' . r loc t l i r r t ' o l cornnron s ( ' l l s ( 'thoue l i S in to t ts & I ) i l l r c : r rn 's cx l to r t : r t ion u l ) l )curs to l r r ' , i t o l l i ' r ' s no Pr i r t ' t i ta l q r r i t l c to th t 's o r t i t t g - o u t o l ' f i r s s i l a s s < ' n t l r l a g t ' s . ' l ' l t t ' n ' u s o n l i r t l r i s i s , o l r ' o r r l s r ' . t l n t t l r t ' r t , i s r r o r l i r c c trc la t ions l r ip , in t l t ' t ' c l I lo cor rs is tcn t r t ' l a t io r rs l r ip a t l r l l . l r t ' tn c t ' r r sp t ' r ' i ; r t ion : t r r< lm < i r P h o l < x { i ( r a l c h : t t t g t ' ( t ' . u . \ ' r l r r r , I 1 ) { 3 0 ) . ( ) r r t l r t ' o r r t ' I l r r r r l . s p t ' r ' i a t i o r r n r : r v t l r k t ' p l : r c t ' i r r t } r t 'absc t tcc o l ' t rppr t ' t ' i : t l r l t ' n to rp l to loe ica l c l i r '< ' rqcn t ' r ' . r r l r i l t ' on t l r t ' o t l r t ' r cor rs i r l c l i r l r l t ' l o< i r lc l i { ] i t r t ' n t ia t io t l n r r l \ ' oc ( 'u r , l )u r t i ( 'u l l r r l v w, i th in g t 'oq lap l r i t ' l r l l v n i r l t ' sp r t ' r r r l sp t ' t ' i< ' s . rv i t l ro r r la l t Y r u p t u r ( ' o l t l r t ' r t ' p r o c l L r c t i v r ' ( ' o n t i r ) u i t v t l r : r t l r o t l r l r i r r c l s l r r r i l l i r r r i t s t l t c s p t ' r ' i t ' sp < . r p u l a t i o n . ' l ' h L r s . l v l r i l t ' l i ' u r v o u l t l r l i s a g n ' t ' t l u t t t l r c l t ' l l r s l r t ' t ' u n ( ) r l r ( ) r ' ( ' s i g r r i l i r ' : r r r tt h c o r t ' t i c a l : r t l v l t t t t t t ' i r t s v s t t ' n u t t i c s t l l l n t l r t ' s l r i l i l i o r n t l r c n r o r ' l r l r o l o q i c l t l t o t l r t l r i o l o r g i c l r lco l l ccp t o l sp t ' t i t . s , th is t l r l tng t ' l l Ls op t ' t t t ' r l t l t t ' r loo l to l l tos t o l p l l rc t i t a l p lo l r l c r r rs u l r t ' n i tc l o m o s t o t l t < ' t t t ' t L t a l i t t t t ' r 1 r r t ' t u t i o l t o l t l r c l i r s s i l l t ' t ' o r r l . l r o l r l r r i t t ' l l x r l t l i o r r r t l r c c o r r s i r l t ' r ' r r l r l t 'c l i f l i cu l tv o l ' t r ; tns lu t i t tg r r t : rg r r i t r r r l t ' s o1 ' r ' : r r iu t io r r l io r r r o r rc sp t ' r ' i t ' s o l sp t . t ' i t ' s p l r i l to r r r ro t l rc r .i t i s a l s o t ' r ' i c l c n t t h u t s u t ' l t l r r x o r r o r n i t ' c l t ' c i s i o n s a r t ' r ' r i t i c a l l v l l l c t t t ' < l l r v t l r t ' r ' l r o i < t ' o l r . r , l r i t ' l rl i v i r r e s p t ' c i c s - o r p u i t ' o 1 ' r t ' l a t t ' r l s P t ' t ' i c s i s t o l r t ' t u k t ' r r r L s : r r l r i t t ' r ' o l ' t l r t ' r ' i r r i l l r i l i t vp c r m i s s i l t l t ' i n a I i t s s i l u s s c t r i l l l ; r q c l t ' t ' o q r r i z c t l a s l r s p t ' r ' i t ' s . I ' l r t ' r ' t ' i s n ( ) q u a l a n t ( ' ( ' t l r i L t t l r t '" t t l o s t ' s t l i v i r r e r t ' l a t i v t " ' ( a s i t L r s u l r l l r l r o i l s c l o u , r r t o , w ] r t ' t l r t ' r ' o l r r o t t l r i s i s r l c t c l r r i i r r i r l r l cw i t h l r n t ' c t ' r t a i n l l ) u ' i l l l r t ' t l t c r t r o s t l i P P r o l l r i ; L t c r r r o t l t ' l t o a ; ; P l t ; I i r r i r r s l . r r r r , . . r \ l x ' ( i ( . so c r : u p f i t t t l : r l i n t i t c c l l t ' l i c t r l i : ; t r i l r u t i o n ( o r r t ' o l ' t l r t ' g t ' r ' l r t r r p t ' s . 1 r t ' r ' l r L p s : ' ) u o L r l < l a l n i o s tc c r t : t i t t l v l t c l t l t u r t r t ' l i a l r l t ' g L r i r l c t o t h c t l t ' l i n t ' r r t i o n ( ) 1 ' l r * i r l t ' l r ' - r l i s t l i l > r r t t ' t l r ' < ' l l t t t ' t l l i r s s i lspc t : i t ' s . N Ior t 'ov t ' r . l t t i t l t r tL tc l t Inon ' i rn1 ;or tun t l \ . r r l l r ) ( )s t i r t l i r l i a l r l r n , l l r l i s t ' r ' a l ru r tc r l i su : i th i r t -spe t ies v l i r i i t t io t t , w l r i t ' h r ' r ' i l l l r c l i r r r r r r l to l r q r ( ' i t r ( ' r ' o l i r l ( ' ss ( ' r ' r ' r r t ' r r t i r r u l l l r r l r tonr ic lL lsvs toms. [Ju t l vha t i s i l l l l )o r tan t in t l i s t inqr r i sh in r - l a r r ro r rg sp t ' t ' i t ' s i s b r l r t , t :u r . - . t f t r r ' i r , . r r l r r ia t io t r .\Vhcrc spc t ' i cs l t r t ' t ' l osc lv r t ' l a t t ' c l ( l t r r r l I r t ' nc t ' l ) ros t l i k ( ' l v to l r t ' t o r r l i r sc r l i r r t l r t ' l i r ss i l l t ' t o r '< l ) ,var ia t ion o f ' t l r i s k inc l i s l i k t ' l t to l r< ' r ' r ' s t r i t ' t t ' c l to l { i ' r r t . ln r .a t ' t t ' r . s on l r ' : t l r t ' r ' us t n r i r jo r . i t l ' o lanatomica l ch : rn rc l ( ' r s rv i l l ovcr lap s r r l rs tun t i i r l l v o l to t lL l l r ' .

l ) t ' s l t i t t ' th t ' lack o l ' r t ssoc i l t t io t t l r t ' t r v t '< 'n t l i x i c unc l n ror 'p l ro loq i t ' t ' r ' o l r r t io r r . I rowcr '< ' r ' . uccr ta in an tount < t l ' q t ' t t t ' ra l i z : t t io r r l ionr thc l i v i r rq l i r r r r r i r i s l ross i l r l c r rs i r l r ro r r r l gLr i r l c tos ) ' s tcmat ic p rac t ic ( ' i r t ana l l z i r tq thc l i r ss i l r t ' t ' o r r l . l "o l s l t< ' t ' i a t ion l r r r r ' l r ' . i l ' r ' r ' t ' r ' . i r rvo lvcsq u a n t t l m m o r p h o l o u i c l L l l t ' r L p s . I l ' o r r t ' s u r v c r . s t l r t ' p r i n r a t ( ' s l r . s l r w , l r o l r ' , o r r t ' l i r r r l s t l l t t t h t 'morpho loe ica l c l i f l i ' r cn t :cs l r t ' t r ' r ' r ' cn c los t ' l v r t ' l a tcc l sp t ' t ' i t ' s (s l i r sp t ' t ' i t ' s usu l r l l v c l r rss i l i t ' t lw i th in thc s : rmc g( 'nus or subgcr r r rs ) a rc comnron lv s r r l t l l . unr l rcs t r i t ' t t ' c l to on lv o r rc o r : r l i ' r vcharac tc rs . I i spcc iu lh 'uhcn 'such spcc i ( ' s u r ( ' \ ' ( ' r v c los t ' l v r ( ' l a t ( ' (1 , I i r l i r rs t lL r rcc n l r t ' r '< ' t r r ,ospcc ics : r rc c l t rs t :c t tc i t ' c l l i o r r t l t c sanrc parcu t l r l sp t ' r ' i t ' s , < l i s t i r rqu is l ) ing l i ' i r t r r r t ' s u r '< 'o l i t ' r r

sPF, ( i I l ) s R[ , ( ]o ( ;NIT IO\ t67

l im i ted to so f t t i ssuc charar : l c r i s t i cs u 'h ic l r ck r no t p r ( ' s ( ' rvc in thc l i r ss i l r t ' co r r l . Anc l cvcn inthosc cascs u 'hcr t ' c l i s t inc t ions in cc r t2 r in harc l - t i ssu t ' churac tc rs a rc o l ts t ' r r ' : rb l t ' l r c t 'nvccn

c losc lv r t : la tcc l spcc ics . subs tan t ia l o r conrp lc tc ovcr lap u 'ou lc l l r c cx l . rc t : t cc l in a l l o thcrs .Bccausc o{ ' th is i t i s casv l i r r thc pa l t 'on to loq is t to conf is t ' r 'u r iu t ions o1 ' th t ' t i , r .o k i r rc ls .t - -spcc ia l l v n 'h t :n 1hc t imc ar r i v t ' s l i r r laxononr ic . juc lg< 'n ' r t 'n t on snra l l sanr l t l t ' s o { ' l i r ss i l s . ' l ' hck t : t :nc r o l l c ' s upp l ' c ( : ia t ion o1 ' t l l c in rpor tan t l i i c t o f in t ra -sp t ' t ' i t ' s var ia l r i l i t r ' , { i rn t l r , anc ljus t i f iab l l ' l i xcc l as t l t c k t . r ,s to r rc o1 'modcrn pa lcount l r ropo log ica l svs t ( 'nn t ics . th t 'q r t 'u t t ' rth t : dangcr o l ' c lo ing th is l r t ' conr t ' s . ( l l ca r l r ' . cvcn 'sp t ' c ics i s var iu l r l t ' . anr l th is ( ' ( ' r ) t ra l r t 'a l i t vcan l lo t bc i suorcc l i t t comPul i r tq tu ,o l i r ss i l s , l i r r . ius t r rs s i rn i lu r i t v in u g iv t 'n s t ' t o f ' [ i ' a tu r t ' sdocs no t guar i t l r t c ( ' t lu t l tu 'o l i r t t r t s b< ' lon11 to th t ' san i t ' sPt ' r ' i t ' s . o l rs< 'n c r l r l i l l i ' r cn t ' t ' s n rav l lo timp lv spcc ics c l i s t inc : t ion . Nt ' r ' r ' r thc l< 'ss . ( ' l r c l ) s lx ' ( i ( ' s r ' : r r i t ' s u i t l r i r r l i rn i t s , a r r i l i t i s I ra r r l toar "o ic l th t ' c< tn t : lus io r t t lu r t r r r r r l t ' r ( 'u r rcn t taxor ro r r r i c p rac t i t ' r ' thc r t ' i s a c l i s t i r rc t t c r r r l< 'nc t ' tou n d c r c s t i m a t c t h t ' a l r u r t t [ u r r t ' t ' o l ' s p c c i t ' s i n t h t ' 1 l r ' i r r r a t r ' , u r r r l n o t a l r l v t ] r t ' I r o n r i n i c l . l ' o s s i lrccrlrcl .

' l ' h is o l rs t ' rva t io t r n l l l s ( 'ou l r t ( ' r to ' l ' c rnp l t ' l on 's ( l1 ) t32) sp t ' t ' r r l l r t ion t l r i r t s i r r< ' t ' rn : r r r r r r l r l s

: I r ( l t r lo rc p r t t t t t ' l l ta t t o t l t t ' r v t ' r t t ' l r t ' u t ( ' s to l i r rn r po lv t t 'p ic sp t '< ' i t ' s . t l r c r r r r rn l r t ' r o l n r : rn rnr i i lspcc i ( rs in th< ' { i rss i l r t ' co rc l i s l i kc lv to l r t ' ovcr t ' s t in la t ( ' (1 . ' l ' t ' n rp l t ' to r r ' s : r rqu l l ) ( ' r r t . I rou , t ' r ' t ' r . i sb r ts t ' c [ purc lv o r ) l ) r ( ' sL . l lnc ( l l t ' l a t i v t ' l r rop t ' r rs i t i cs o l ' r l i l ] i ' r t ' r r t 1 ;o l ru la t io r r s t ru ( ' tu r ( ' s l ( )

Pr ( )n io t ( ' r tLp ic l po l tL r l : r t io r r r l i v t ' rg t ' r t t ' t ' , l t cucr ' po lv t r 'p isnr , w , l r i c l r I r t ' a l )J ) ; r r ( 'n t l \ ' ( ' ( lu l i t ( ' s

u , i t h c l c t c c t u l r l t ' n r o r ' y r l r o l o q i t : r l s h i l i s . l l r r t : r s u , ' r ' l r r v t ' s c t ' r r . i t i s r r r r l t ' r r l i s t i t ' t o r n : r k c t l r i st :c1ua1 ion , l i r r r l n ro r l tho loq \ , i t s t ' l l . t t ' r tu in lv thosc lLs l r< 'c ts o l ' i t t l r : r t l ) r ( . : i ( ' r \ ' ( ' i r r th t ' l i r ss i lrccor r l , t t t l l s : t t I i l l i ' r t ' n t s to r \ ' . ( ] r : r r r iosk t ' l c ta l r l i l l i ' r ' r ' r r t : t ' s l rc t l v t ' r ' r r p l in l r t t ' s r r l rsp t ' r ' i t ' s o l t l r csamtr s l t t ' t : i t ' s t t ' t t r l to l r t ' t i r r r ' . i { ' o l rsc lva l r l c l t i r l l . l "o l t ' r l r r r rp l r ' . r l i l l i ' r cn t ' t ' s on l } ) ( ' ( ) r ( l ( ' r ' ( ) lt l t t t sc sc l tan i t i r rq th t ' r 'a t ious su l rspcc ics t l ' l )n tp i t ln tus t l iudr t t tu n ,o r r l r l < t ' r t l r i r r l v l r t ' r r i i ss t ' r l l r va p i i l t ro l t to loq is t $ ' i t l r o r t l r '1 t ' t ' t l t r r r t t l l ro r rcs t ( ) ( ' \ run in ( ' , r ' s1 t t ' r ' i l r l l v i l l r c l r t r l a k t ' r ' r r s t ' r rs t 'o li r r l r i i -popu la t ion r ' : t r i r t t io t t . l ' , r ' r ' t t thc h ie l r l r , l ) ( ) l v tv l ) i ( t ' x ta r r t sp t ' r ' i t ' s I lo t r to . ia / ia i . r vur icsn tu t : l t l< 'ss t l t t rn o l t t ' l vou l t l in l i ' r ' l i onr t l r c v lL r ic tv o l ' l i r ss i l rnor 'p l rs t l l r t I r : t s l r t ' t ' n t ' r ' l r r r r rncr li n t o i t .

' l ' l i c t t ' n t [ t ' r r c v t o u l ) ( l ( ' r ( ' s t i r r n t t ' l o w - l t ' r ' t ' l t r L x i c t l i v t ' r ' s i t v i n t l r t ' l i r s s i l l t ' t ' o r r l i s , i t n i u s t l r c

sa id . in l in i t t ' l t ' p r t ' l i ' r l l r l t ' to t l r t ' oppos i t r ' l i r r r l t . u ' i r l t ' s1 t l t ' : r< l in c l r r l i t ' r ' r l l r vs , o l ' l rap t iz i r re t ' i r t ' l rn t ' l v l i r s s i l s l ) ( ' c i n r ( ' n u ' i t h i t s ( ) \ \ ' n n a n r ( ' . l J u t i t n r : r v r r o r r t ' t l r t ' l t ' s s s r r l r t l r , ' q i v t ' r i s t ' t o r rc l c s t r u t ' t i v t ' n r i s c ' o r t c ' t ' p t i o r t . I t n r : n < 1 u i t r ' p t ' r s r r l s i v t ' l v l r t ' l r r q r r t ' < l t l r : t t i t i s t u r i n r l i o l t r r n Il vh t ' t l t t ' r th t ' sp t ' c i< 's r ' r ' t ' r t ' t ' oqn iz t ' i r r th t ' l i r ss i l l c t 'o l r l l r r t " ' r t ' u l " s1 ; t ' r ' i t ' s o l n ( ) t . ' \ s lonr l t rst l tos t ' u r t i t s rv t ' r t ' t ' og t t i z t ' l r s sp t ' c i i ' s lL r t " ' r r l t r r ru l " , o l r r ronop l r r l t ' t i r ' . r locs i t r t ' rL l l v n ra t t t ' rl v l t t ' th t ' r t l t c r " cons is t o l ' i nc l i v i i l u r r l spc t ' i t ' s o r o1 'uqgrcqutcs o l c los t ' l y r t ' l l r t t ' r l spcc ics? \ \ ' l r t ' r ct l t c rc i s t to r t 'usoruL l r l c q r rosp< ' t ' t o l ' t l i s t ingr r i s l r i r rq - s r rch l )u t i r t i \ ' ( ' sp t ' r ' i t ' s o r r t l r t ' l x i s is o lp r c s c r v t ' c l n t a t t ' r i l t l s t h t ' r t ' t ' l r r t . o l ' t ' o r r r s t ' . l r t ' o n l v o r r t ' r t ' l r s o r r l r l r l t ' l r n s u . t ' r t o t l r i s < 1 r r t ' s t i o n .N < t n c t h c l c s s , ' , v l t t ' r c r t ' s p t ' t ' t l i r r i n t r a - s I ) ( ' c i ( ' s v u r i r r l r i l i t v l t ' r r r l s t o t l r t ' i r r t ' l u s i o r r u ' i t l t i n us i n e l c s p c c i t ' s o l ' l o s s i l s r t ' p r t ' s t ' n t i n r l n r o l t ' t l u r r r o n < ' r t ' u r l i l l i r l t ' n t i l l : r l r l t ' n r o r ' 1 t l r . i 1 i sabso lL r tc lv c ruc ia l t l t : t t rv t ' l r t ' to tu l l v r :on l i r l cn t tha t on lv l r s i r rg l t ' sp t ' r ' i t ' s i s i r r l i r c t i r r , , .o lvc r l .' l ' h i s

i s n o s m a l l m a l t ( ' r ' . l " o r r v l r i l c s p t ' r ' i t ' s a r t ' o l r j c t ' t i v t ' I r i s t o r i c i i l ( ' n t i l i ( ' s t h i r t u , t ' r r r t 'ob l iecc l to con l ion t . morp l iok ru i t 'u l v r r r i t ' t i t ' s l r , i l l i a sp t ' t ' i cs l r r< ' : rck r ron ' l c r lgc r l to l r t ' no n ror t 'than cp iphcno l l l c l l : I . rv l rosc inc l i v ic lua l h is to r ica l i r l t ' n t in hus no t l t cor t ' s t l r l r l i s l rc r l .

S p c c i t ' s , r c p r o d u c t i v t ' l v i s o l a t c d f i ' o n t t l r t ' r t ' s 1 o 1 ' t h c l i v i r r q ' " r , o r l c l , a c r l u i r t ' r h t ' i rd isc rc t t :ncss i i s thc rcsu l t o1 'an i r rcvcrs ib l t 'g t ' r r t ' t i c anc l h is to r i t ' : r l 1 l r 'occss . Su l rsp t ' t ' i t ' s , ont h t t o t h c r h a n d . r v h i l c c l i s t i r r g u i s h a b l c i n s o m t ' n ' r r v l i n c l : r t s ( ) n r ( ' l ) ' , i n l , l u ( i r r t r r r r t t p t

r6B I . TATTE,RSAI ,L

ephcmcra , unboundcd bv rcproduc t ivc bar r i c rs : thc i r un iquc idcn t i t v may a t l cas tpo ten t ia l l v be los t a t an \ t imc in mt ' rs ing n ' i th o thcr su l rspcc ics l topu la t i< ins . C)n lv u , i thspec ia t ion , w i th thc dcvckrpmcnt o { - rcproduc t ivc i so la t ion , d r ics thc idcn t i t v o l ' r vha t bcsanas a subspcc ics bccome dc l ln i t i vc lv cs tab l i shcd. C) f ' coursc th is docs no t ncccssar i l r 'mcantha t subspcc ics , as cn t i t i cs , d t i no t dcmand s tudv and cxp lanat ion ; thcv har ,c o r ig ins andh is to r ics jus t as sp t ' c ics do . But thc la t : t rcn ta ins tha t , to l ta lcon to loq is ts in gcncra l ,

subspcc ics a rc cp ip i i t :nomcna w l i i ch do no t mcr i t thc i i t t cn t ion l ta id to sp t ' c i t s : i t i s spcc ics ,not subspccics, that arc thc units r i f ' t :r 'olut ion, or a1 l( 'ast o{ ' t" , olut ionan' stucl l ' . And indccd,to judgc l rom thc ecncra l l y ins ien i f l can t in tc r tax ic d ivc rs i t r , i r r harc l par ts cxh ib i t t : c l a t thclowcs t taxonomic l cvc ls in thc modcrn pr imat t ' l i i una , thc pursu i t o f ' su l rspcc ics in thc { i rss i lrecord is a t bcs t f iaught u ' i th d i l l i cu l tv , : rnd is morc p rob : rbh f i r t i l c . ' l ' l r i s i s r , r 'hv i t i s c r i t i ca lto avoid rclcgating dist inct morpl iokreic:r l 'u 'ari : ints ol ist ' r 'u ' t 'c l in thc l i rssi l rccorcl to thcstatus of 'subspc<:ics-or thcir inl irrnral c<1uir ' :r lcn1-r,r, , i t l rou1 thc lrcst of 'r t : :rsons l i rr r loingso; io r , as n t ( ' rc subspcc i t ' s , cvcn < ' l c : r r l f i c [cn t i l l a l t l t ' n ro rphs c lo no t u rg( ] l l t l y c [ t 'm iL ldana lys is a t td t : xp lanat ion . Indccd. i t m ig l i t r , vc l l l i t ' a rqucc l th r r t i t l vou lc l bc l t c t t t ' r f< i r t l r cc r lmpr r :hcns ivc l )css o l 'our u t tc l t : rs tand ine o l ' t l i c l run l rn l i r ss i l rccorc l th i t t , i l ' t ' r r wc must , wccr r (w i th in rcason! ) on th t ' s idc o f ' r t ' coqn iz ing too nr iu rv r : r t l ) ( ' r t l l an . as i s thc t t 'nc l t ' r rc_v , toolcw spcc ics un i ts . A f ic r a l l , to hark l rack to un car l i t - r c ( ) rnnr ( 'n t , c \ ' ( 'n l r subsp< ' t : i cs has : rh i s t o r l w o r t l r v o l i n r t ' s t i q a l i o r r .

A l l th is , o f ' coursc , b r :qs th ( ' p rob l t : rn o f ' c l i s t inqu is l r in r l s r 'p : r ra t t ' n ro lp l rs l vhcn 'c l i f l i ' r t ' nccsarc sl ieht, lvhi lc tht ' I ' t t .r t t t 'otrcorclancc ol ' t :rxi t ' rvi lh morPhologit 'ul churrl l r ' s( '( 'ms t()c l im ina tc anv ob i t :c t i vc so lu l io t t t< . r th t 'p ra r : t i r :a l p rob l t ' rn o l ' spc t ' i t ' s rc r :osn i t io r r . \ \har Iwish to cmph:rsiztt l tcrt ' , horvcvr:r, is thut rvhcrc dist inct ntorPhs t 'an rt ' r tr l i lv lx ' ic lcnti f iccl i twou ld sccm mos l Proc luc t ivc 1o : Issum( ' tha t thcv rcpr ( ' scn t spc t : i cs un l t ' ss t l rc rc i sc r impc l l ing cv ic lcncc to l - l c l i cvc o th t : rw isc . ' l ' o l r rush nror l t l ro log ic : t l c l i v t ' r s i t y unc l t ' r l l i t ' ruqo l 'an a l l -cncompass in r l spcc ics i s s i rnp lv to l r l in t l oncs t ' l l ' to t } rc conrp lcx rca l i t i cs o f 'phylogcnv.

The Human Fossil Record

By 1950, a t a t im( 'whcn th t ' knorvn l romin ic l l i r ss i l n rcor r l was v l rs t l v rnor t ' l i r r i i t c r l t l ra r r i t i sa t p rosc l l t , thc namcs o l 'ov t ' r l r dozon g( ' l l ( ' r l r , : rn t l n r l r r rv n ro lc s l tc t ' i cs , ,uvcr t ' t ' onr rnon lvt lncountc rcc l in thc l css tha t r vo luminous l i t t ' ra t r i r t ' on f i rss i l Horn in ic la t ' . I r r tha t v t 'a r ,howt :vc r , a lo t ts t i v t : rduc anc l h ieh ly in l lucn t ia l p : rpcr l rv l , ln rs t \ ,1avr s t rL r tcc l thc lunrp inubandwagot t r r ) l l i ng u , i th thc ck 'c la ra t io r r tha t : " th t 'ucc( 'P l r inc r ( 'o f ' th t ' r ru rv conc t 'P t o1 'b io log ica l l y d t : l lncd po ly t ,vp ic r s i rcc ics ncccss i t : r t cs thc uprv l r rd r t ' r , i s io r r o l 'a l l o thcrca tcsor ics" (Mayr , 1950, p . I 10) . N . lav r spur t tcc l th t ' i c l t ' a o l 'monot \ ' l ) i ( ' q ( 'n ( ' ra as l r ' : r c l i r rs toan "incquali tv ol 'c:rtcuorics", and at t l )at t inrc ht: t ' r ' t 'n urgut 'cl l i rr i rrclurl ine Auslrahtl t iLlLet:u:in Homo, s inc t : "w i th in th is typ t : lhc rc has bccn spcc ia t io r r n .su l t ins i t I Ion to sap in . i ' .Nonc thc lcss , w i th in l in i t t : l y c lc : r rc r v is ion lhan thosc o [ ' h is succcssors rv l ro ra is t 'd thc"s ing lc spcc ics hvpothcs is " to a t r a r t i c lc o l - f a . i th , N Iavr cxp l i c i l l r , ' r ccogn izcd no t on l r , lh l i t t< rar r i vc a t th is taxonomic : conc lus ion rcqu i rcd "a c lc l ibunr t t ' m in imiz ing o l ' th t ' l r ra in as adccisivt- ' taxonomic cltaractcr", but also thi i t tht: nc{-rcssarv assumption that "al l o1'tht 'scIbrms, including Australopithecus, arc csscntial lr , 'mcmbcrs o1'a singlc l inc o1'clt 'sct 'nt", rvascxc t :ed ing ly f rag i l c , lb r "Add i t iona l l inds mig l i t t :ad ih c l i spro 'u 'c th is . " (pp . l l0 l l l . )Subscqucnt palcoanthropcikisical discovt:r ics h:n'c cmphasizccl l l rc u, isclom o1' NIavr's

SPF)C] IL ,S RE(JO(}NIT ION 1 6 9

ca\reat) but thc thrcc and a half 'dt:cadcs f ir lkrrving his salutorv contr ibution havc rvitnr:sst:d

thc asccndancc o f what has f lna l l v bccomc: a dcs t ruc t i 'u c lv min ima l is t approach to homin id

svs tcmat rcs .' l 'odav,

happil ,v, thc rcal izat ion has bcsun to dau'r 'r that a sieni{ lcant cicsrt:c o{ '

taxonomic varicty cxists in thc rcmotcr stagcs o{ ' hor-ninoid historv, amonq t lr t :

australopiths. Thc gcnus 11orna, hor'r 'cvcr. rvhich l i r l lor 'r ' ing N'Iavr 's cxlrort i i t i<in l tas bcctt

cxpandcd to cmbracc ov('r two mil l ion vr:ars o{ ' t imt' uncl :r largc lrrral ' r i l 'dist inct hominid

morplrs, is commonlv r, icn,ccl as crintaining onl\, thrcc spccit 's: Honn habil i . t , l l . ereclt ts, ancl

H. sapiens. ' l 'hc

f irst of ' thcsc poscs acutc prolr lcnrs o{ ' clcl ini t ion i incl rccognit ion,

cxaccrba tcd bv thc fhc t tha t th t ' on l t ' spcr : in rcns in a mot lc r " r :o l l cc t ion tha t a rc t ru lv

dist inct ivc morpl 'roloeir:al lv arc not irssot ' i ;r tccl nith t l ic nnrnt ' . ' l 'hc st 'cr. ,ncl has. in thc vcarss incc N ' lav r wro tc , cxpandcd bcvonc l i t s vaquc ro lc as th t " 'man o l ' th t 'm ic lc l l t ' l ) l t ' i s tocc t t t " '

to inc ludc a d iv t : rsc : r r r : rv o l - { i rss i l s tha t lu tu rc scho la rs n ' i l l sur t ' l v l i r rc l ampl t ' c : rusc to

subd iv idc_un lcss th t .v can cont r i v t . t t r n l r i r r t : t in thc I i c t io r r o { ' thc , ,s r : rc l t . " to kc t :p th is

d ivc rs i tv dcccnt ly obscurcd . ' l ' hc in tc rprc t l r t io r r o1 ' rnos t hunrar r f i r ss i l s su l rs t :c lucr ) t to a lx )u t0'5 mvr as bclonging somchor,r ' Io I tont.o . tr t l t icn.t is pt 'rh:rps t lr t 'most comllrt 'ht 'nsivt:

smokcscrc r : r ' r o [ -a l l . : rnd cou lc l on lv havc bccn n ' ra in la in t 'd l l t ' a zca lous rc lusa l to cons ic l t ' r

charac tc rs < i t l t c r than bra in s iz r ' . I har . ' c suggcs tur l r ' l sc l r ' l r cn ' ( l ' ) l c ln ' r le r '& ' l ' a t tc rsa l l , l9 t i2 )

tha t thc u rqc to inc luc l t ' l i r rms i i s c l i vc rs r ' : rs | t c t r : r lo ru r , S t t ' i n l r t ' i rn , N t 'a r rc l t ' r t l r ; r l anc l vou r inc l

mc in thc s ine lc spcc ics Horno s tLp iens must l r t ' so t ' io los ic i r l i r r o r i { r in - l l i t ' r t ' i s somcth inq

vaquc lv c l i sqracc lu l a l rou t c l i sc r im ina l inq aq ; r i r rs l o th t ' r la rq t ' - l r ra incr l l ro rn in ic ls - l rn r l I c lo

not rv ish to bca t tha t horsc l i r r t l i c r hcrc : bu l th t ' l i L r : l r t ' n rn ins tha t thc on lv ra t iona l

cxp lanat ion { i r r thc taxononr ic t :o r ra l l in r r toq t ' thc l o l ' thcsc iv ic l t ' l r . ' r l i l l i ' r ing l i r ss i l s ( l i r r agc

wi l l no t c l< i i t . Wolpo l l , l1 )U0. to thc t :on t r : r r r . ' ) i s thc s t ' t t ine o l ' i rn uncons t ' ious "c t ' rc l r ra l

Ru i r i con" , pcr l t : rps : r t somcrv l r t ' r c a rounr l l2 (X) ml . ' l ' l r t '

l ' l i c t l l u r t suc l r : r Rub i t :o t t ,

u t rconsc ious or o t l t c ru ' i s t ' , rvou lc l cxc luck ' : r qoor I rn : r r r l , rncnr l rc rs o l ' t l r t ' l i v i r re sPt :c i t ' s l ionr

Honto . t t tp icn . r has qonc unr t 'mzr rk t ' c l . r \nc l ' "vh i l r ' , i l ' rnv surmis t ' i s cor r t ' r ' t , onc t :nn or r l l

applaucl thc qcncrous l i l r t ' ral s( ' l l t inl( ' l l t that l t 'ar ls lo tht ' inclusiort in I tono tuft iens ol"al l

homin ic ls w l ros t ' l r ra in s izc fa l l s coml i r r ta l r l v w, i th in th t ' mor l t ' rn nu lqc , on( ' i s t ' t l ua l l v

ob l igcd to po in t ou t tha t to conc lud t ' tha t no s l ) ( ' c i ( ' s r l i f l t ' r cnc t ' s cx is t u ' i t l r in t l r i s ; r r rav o l '

homin ids is to con lus t ' th r : t l vo k inc ls o1 'n rorpho loq ic :u l r , : r r i : r t io r r . ' l ' l r t '

I rue( ' var ia t io t r i r t

n r i rma l b ra in s iz t : { i runc l u rnonq mor lc r r r hum:r r rs cxcmpl i { i t ' s r ' r , i1h in -sp t '< ' i t ' s var i : r t io r t , lu tc l

i s la rgc lv o r cn t i rc lv i r r t : luvant 1o thc i ssuc o f ' tax i t ' c l i v t ' r s i l r ' . I h iLvc rL l rcar lv suggcs tcc i t l i a t

c losc ly - rc la t t :d p r imatc spcc ics l v i l l s l r r . rw ic lun t icu l o r sLr l rs tan t i l r l l v ovcr l r ipp i t t s ra r tg t ' s o f '

var i : r . t ion in a lmost a l l morp l ro log ic l r l l i ra tu r t ' s ; iu r r [ rvh i l c i t cou ld , I supposc , ] rc a rqucd th : r t

both t lr t : lowcr-than-moclcrn avcr:rgt ' brairr sizt ' ol ' "r lrchaic I Irnm sapien.r" ancl th(l

h ighcr - than-modcrn a \ /c ragc bra in s izc o l ' thc c l : rss ic : N t 'a r rc lc r thn ls i i r t ' l i ' a tu rcs tha t cou lc l

bc uscd in dt:marcating both lrotn Hono .\aPi?n.t, ivha.t is mut:h morc important in wt ' iglr ing

thc qucstion o1' how mrinv hominid spccics \\ ' 'c arc l iLcccl rvi th in t lr t : midcl lc at)d latc

P lc is toccnc is thc cons tc l la t ion r i l ' l ca tu r r :s in rvh ich l ro th c lcar lv d i f l c r l iom ca . r :h o thcr and

I iom oursc lvcs .

Flvt:n a rclat ivclv cursor\r survcv o{-thc spt 'ctrum ol ' l iv ing primatcs is sul l l t : icnt to rcvcalthat modcrn Homo salt iens, b,v nou, numbcring somc l irur bi l l ion indivicluals ancl l i rr lonereprcscntcd in v i r tua l l , v cvcrv inhab i tab lc rcs ion o f ' thc car t l r , i s u ra thur h ieh lv po ly tvp ic

specics. Noncthclcss, l ikc cr, 'cry othcr spccics, polytypic or olhcrwisc, I Iomo salt iens rs a

cohes ivc b io log ica l cn t i t v wh ich , wh i lc var iab lc . i s f 'a r l iom in { ln i tc lv so . Rcgrcss ing in t imt : ,

t7 {) I . 1 'A1 'TF,RSAI , I

we l ind tha t back to : ibout 110 35 k l ' r ago rnor ( 'o r l css a l l l i r rnnr r l i r ss i l s ta l l in to th t :morphological ranqc bnrckctcd bv thc I iving sPct. ics. Sonr<' subsPcti t 's <l ' Horto sapiens rn.r-t ,' "vcl l , <i1-courst ' , l t :n'r 'cmcrg('d n' i thin t l t ;r t I)cr i(xl onlv to lr t ' t 'onrr '( 'xt inct. ancl thus i t r , . , ,oulcllrc rcas<rnabl( ' to cxpc(' t to l incl. as inclct 'cl u' t ' r lo. thc ot ' t 'usion:,r l I Iomo .rt f funs l i rssi l thats l igh t lv cxc t rc< ls th is ra t tgc in onc chanrc t t ' r o l uno lh t ' r . I1 'u t ' l ook l i r r thc r l rack , hou ' t ' r ' c r , rvcabrupt lv losc thc ab i l i t v to I l t thc humun { i rss i l s n , r ' l l nc l i r r to t l r i s l i r r rn t 'o l ' r ' u r ia t ion . r . " , i t } r t l r ccxccpt io t r o f a l i ' r v p ic t ' cs t ln t a rc t ' i thc r l i ' uqm<' r r t lL rv o l u r r t ' t ' r t lL i r r l v t l : r t t ' r l . l l u t a l thoue l la lmost a l l hunr : rn f i rss i l s p r io r to th is t in r t 'a r ( ' \ ' ( ' r v r l i l l i ' r c r r t i r r as l r t ' r ' t l i onr anv th ing \ \ , ( ' s ( ' ( 'i r r our vc r r " r ' : t r ia l r l t ' l i v inq spcc i t ' s . th t ' r . : r r t ' r ron t ' thc lcss lcgu l r r l l v t ' l ass i l i t ' r l t s I Ion to . to f t i t : r t . t .\ { i t l r t l r t ' p rov iso . t ( ) l ) ( . sur ' ( ' , thu t 1h t ' r ' : r r t ' "u rc lu r ic " ' . o l o th t ' r 'u , i sc r l i l l i ' r< 'n t l bu t I Ion tosap iens nonc thc lcss .

Rar t ' l r . i r t c l t ' t r r l . hon , t ' r ' t ' r ' . I tav t ' Pu lcor r to log is ts t ' r ' t ' r ' I i runr l i t r rc t t ' ss l r l r to c l i s t inqu is l rl t c t ' "v t ' t ' t r " l t rcha i ( " ' l t t t c l "u t ra to t r t i c l t l l v rno i l t ' r ' n " tv l ) ( ' s o l ' t l r t ' su r rc sp t t ' i t ' s . r rnc l t l r c r t ' s t ' r ' r r rss c a n t . i u s t i f i c a t i o n f i r r s c l u t ' r ' z i n { t l t c s t ' t l i s t i r r c t I r o n r i n i r l n r o r ' P I r s i n t o u s i r r q l c s l t c c i c s . I r r l r n r ,e r o u p o t l t t t r t l u u t H o t t t i r t i r l u c t l r t ' l ) r ( ' s ( ' n c ( ' o l ' s c v c l l r l < ' l t ' l r l l v l c t ' o q r r i z u l r l t ' n r o r ' p l r s i r r t l t cr c c o r t l o f ' t h c r n i d t l l t ' ( o u l ) l ) ( ' r l ) l c i s t o t ' t ' r r c n o L r l r l s r r g q t ' s t ( i r r i l t ' r ' r l , r l < ' r r r o n s t r u t t ' ) t h cin 'uo lv t 'mct t t o l ' scvcra l sp t ' c ics . r \n l n r ; r r r r r ru r l ia r r p l l t ' o r r lo log is t s t ' r ' i r rg n rorpho loe i t ' l r lc l i f l i . r c n c c s ( ) l l t l l ( ' ( ) r ( l ( ' r o 1 ' t l r o s t ' s t ' P i t r l t l i r t u r r r o r l t ' r ' r r l r r r r r l r r r s l i o r r r t l r t ' i r r ; r l i r , t r s l ) r ' { ' ( u r \ r ) r s .i i r t t l t l i c l : r t t t ' r I ronr t 'u t l t o t l t t ' r , w ,oLr l t l l r rv t ' no c l i l l i t ' u l t v i r r r t ' t ogr r i z i r r ! r r r ru r r r l rc r o l ' s r 'p i rn r t t 's p c c t i c s . , ' \ n c l i n t l r i s c l c c i s i o r r t h t ' n ' i s r r o o l r v i o t r s p l i r t t ' l i r s p c c i r r l p l t ' u r l i r r g . t ' r ' t ' r r u l r c r t ' i t i so t l r o !v t l t ' l os t ' s t r t ' l i t t i v t ' s t l ta ( u r t ' i r rvo lv r '<1 . I t i s r ro t i r r t l r c l t ' as t s r r r 'p l i s in r l t ln t v l r r iousi i u t l t t r r s , f i r r t ' x a n r p l t ' S t r i n s t ' r t t u l . ( 1 1 ) 7 1 ) ) . \ \ ' L r ( 1 1 ) t t l 1 r r n r l l { i q l r t r r r i l t ' ( 1 1 ) [ i l J ) , I l r v c l r t ' t ' r rl i l t l c to c l i sc :ov t ' r c l t ' r ' i v t ' t l l i ' i t t r r rcs s l l r rc r l w i t l r t l r t ' i l su ( ( ( ' ss ( ) rs i r r s t r t ' l r I i r ss i l s l rs Ku l ru t ' .Nc lu tu . I )c l r i t lo ruL , r \ r : rqo l rn i l l )u l i . l ' l r i s , I ro r , r , t ' r ' t ' r ' . i s r ro l r lq ru r r t ' r r t l i r r t ' o r rsPt 'c i l i c i t v u l r t ' r rthcv : t l so s l ro ' " r ' t l i l l i ' r t ' t t t ' t ' s o1 'u rnagr r i tu r l t ' t l ra t i r r a r r r o t l r t ' r ' p l i rn l r t t ' l i r r r r i l v * 'oLr l< l l r cac t :c l t t c r l ,uv i t l to t t t t l t ' n r r r r l r s r l cn ia lc l l i r r r l s t ' l l rn r t t ' sp t '< i t ' s . S is t t ' r ' s t l r c l t ' r r r i r r l t t ' : rn rongth t ' s t ' n io r l ihs ; cor rspc t ' i l i cs , r ro .

I t i s n o t t r t t ' i t t t t ' r r t i o n l t t ' r ' t ' t o q o i r r t o r l t ' t l L i l o n t l r c n r o r ' p l r o l o u i r ' : r l . j r r s t i l i < ' l r t i o r r s l i r r 'r t rc t rq r r i z i t t t l scvcra l s ( ' J )unr t ( ' sp< ' t ' i cs i r r t l r t ' pos t - l lon to t r r t lu . t l t r r r r i l r r l i r ss i l l t ' r ' o l r l . I r r< l t ' t ' r l .c v t : r r i l l h i s c o n l r i l l t t t i r t t t r l t ' r ' c I ) ( ) t ( l r s t l i s i r n ( ' s s l r . \ ' r i r t l ) ( ' l ' t l t i r n l r s i r r t ' s t ' l l c l r l ' 1 t o l t . t l r i srvou lc l bc i rnprac t i t ' l r l f i r r t r ' ro in tc r r t ' l l r t t ' t l r ( ' i l i ( ) l l s . In t l r t ' l i l s t p l i rc t ' . r l t ' s l r i t t ' t l r t ' t ' x i s t t ' r rc t 'o la l l ( ' x t ( ' I rs i \ ' ( ' l i t t ' r l t t t r r t ' . r l i s t 'uss io r t o1 ' t l r t ' r l c t : r i l t ' r l r r io r ' y ; l ro loq i t a l t ' r ' i r l t ' n t ' t ' l i r l l L r r ru l t ip l i c i t r< t l ' l t u l r t a t t s p t ' t : i t ' s ( ) \ ' ( ' r t l ) ( ' 1 i a s t l r a l l - r r r i l l i o n v t ' ; r r s i s < l i l l i t r r l t w , i t l r o r r t t ' r t c r r s i v r . s t t r < l v o l t l r t 'a< ' tua l foss i l sp t ' t ' i n r t ' t t s i t rvo lv t '<1 .

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{ < r c u s t r r t t l r t ' c o n l i n u i t i t ' s l r t ' t u t ' t ' r t . s ; r r , " l r l t ' h a i c l l . s r t l i n t . i ' l r r r r l t t ' r r r P o l l r l l r c o r r t i g L r o u sspt rc i t ' s -Hornr t t r t t lu . t , t l t t ' N t ' : rnc l t ' r ' t l l t l s . o r " l r r r i r t ( )n r i ( l r l l r r r ro r l t ' r r r I Ion to . tu f t i t , t t l " - i rn ( lh t tnc t ' , p rcsunt i t l r l r " . l rL rg t ' l v o r r p r in r i t i v t ' r ' l l r r : rc t t ' r s . N los t o l ' t l r t ' l i t c l lL t r r l t ' . i r r r l t ' r ' r l . t l roLrg l rc r : rn rmt ' r l u ' i t l t co t r t l ta t ' i so r rs , i s t ' x t raor t l inu l i l t r r r r i r r l i r r rn l r t i v t ' i l o r r t 'w ' i sh t ' s to k r rou ' i l t l r t 'charac tc r s l i i t t ' s l i t ' i r tq t :onrpux ' r l a r t 'p r i rn i t i v t 'o l r l t ' r i v t ' c l . o l l ro* . , r ' i r l t ' l r . t l r t ' r ' l r rc s l l r rc r l .Sccond lv . oxc( 'p t \ ' ( ' r v rcc ( 'n t l v in th t ' r 'as t 'o1 ' t l r< ' \ i t ' unr l t ' r ' t l uL ls . l i ' u , r r t t ( ' rn l ) t : i I rav t ' l r t ' t ' nmado to c lc f in t ' c l i s t i r rc t i v t ' l a t t ' r ' honr i r r i t l g rou l )s lLs r r ro rphs . l t ' t r r lo r r t ' : r s sp t ' t ' i r . s . no t on lv i rsa r c s u l t < t 1 ' t h c s c a r t ' l i l i l r c o n t i n u i t v l i r r t l r l s , , . , , r r , l ) r r ' \ u n t ( \ , i r r t o r r s t ' < l t r c r r c t ' o 1 ' t l r c i rt :p iphcnomcno loq ica l s ta tus r rs i r r t raspcc i l i c vur iun ts . ( )nc s l rou l r l r ro t t ' . hou ' t ' r ' t ' r , t l l 2 r t \ \ i t l lthc adopt ion b l ' somc pa l t 'oun thropo log is ts o l ' u n rorc o r l t ' ss t ' l u r l i s t i c approac l r to t l r t 'r ccons t ruc t t ion 0 f 'p l r r . lOgt ' l l \ , , th ( ' rc havc | t ' ccn t lY l r< ' t 'n sor r r t '$ t ' l co r r r r 'Pos i t iV t ' r r r tx t ' s . ' l ' l rusHub l in ( l97 t l ) l i s rcd a s t '1 o f 'aponrorph i t ' s o1 ' th t ' sku l l r ( 'u r u rnor )q Ncr r r r r l c r rha ls , u r r r l l )a r ' &

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th t : namc Hono tup ien . r rn igh t no t l l t ' s t l i t ' r cs t 'ncc l l i r r l i r . rn rans o1 'n roc lc rn as l ) ( ' c t . " t ' x t r t :n rc "

as hc I i ' l t thc suegcs t ion n 'ou lc l appc l r r . In a r r r ' ( ' \ ' ( ' n t . f i ) r th r ' pur 'pos t ' s o l ' th t ' p rcs t ' r r t

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h igh ; par ie t : r l a r t 'h long anr l h iqh , nur ro 'nv in l i ' r ' i o r l r u r r r l l r lo l r r l su l rc l io r l r ' , l i r r r r t r t l l ro r t t '

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t h t ' t ' x t n i o r r l i n a r v l r t ' h i r v i o n r l l L r l v l u r c t ' n l ( ' n l s ( s t ' r ' . I i r r c x u r n l r l t ' , \ \ ' l r i t t ' . l 1 ) 8 2 , u r r r l K l c i r r .

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i n n o v a t i o n .

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l r r rs a r r ro r tq l l t t ' N t ' l r t r r l t ' r t l l t l

a p r r r n r r r p h i c s i r l t ' n t i l i t ' r l l r r H r r l r l i r r ( 1 1 ) 7 t J ) , S t r i r r g c l t ' l u l . ( 1 1 ) U l ) , : r r r r l V r r r t r l t ' r n r t ' r ' r s t l t( 1 9 t 3 5 ) l r r t ' t h r ' l i r l l o u ' i n g : l i r n r l r r l o i r l l l a t t t ' n i n g l r r r t [ " l r u r r n i n q " o l t l r t ' o t ' t ' i p r r t ; t l r t '

n r i c l - s : r q i t t r r l c l i v i s i o r r o l ' t h c o c t ' i p i t a l t o r u s . n ' i t l r l i r r n r : r t i o r t o l r r s r r l r r : t i r t i a c l i r s s l r : t l r t '

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rc t roa t i r )q zvqonur t i c p ro l i l t ' r r r r r I r t ' t ronr r l l t r spr tc t ' s ; n : rs l r l : rp t ' r t r rn ' t ' : tp : r< ' io r rs , l v i t l r r r : t s r t l

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nr ieh t bc ac lc [ t ' c l p ronounct ' c l n r ( l ia l ( ' r l r va tu r t ' . l l l r t t t ' n i r rg o l ' t l r t ' l i r s t r ' : r rpo l t t t ' tu t ' l t t ' p i t l

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lonqs t l i r rd inu c l i spu t ( ' o \ ' ( ' r n 'h t ' t l r t ' r t l r t ' e rouy . r o l 'hor r r in i r l loss i l s to t t tu i r r ing t l r t '

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morp l t ( r t ' coqr ' ) i z t ' c l in i t s hav ing l rccn nanr t ' c l , I ron ' t ' r '< ' r ' i r l rpy r ropr i rL tc l r ' ) tha t i s t ' l t ' u r l v

t72 I . TATTF,RSAI - I

dcmarcatcd at thc spccics lcvcl lrom Httmo ereclus,lrom I lomoneanderthalen.r is,ancl lrom Homosap iens . S t r ingcr ( l9 t t5 ) has summar izcd scvcra l o f i t s c l i s t ineu ish ine charac tc rs in a Vcnndiagram which cmphasizcs t irc rtccd frrr furthcr analvsis o1-thc arpomorphics ol-thc eroup.One might point out, hor'vcl 'cr ' , that t :xccpt in thc t irrm of- i ts suprarirbital torus, thishominid specics appcars dist inct also l iom that bcst rcprt:scntcd l- lr , t l ic Stcinhcim andposs ib ly Swanscombc sp t :c imcns (as St r ingcr , 198: r . has a lso i rnp l i cd in c< immt :n t ins oncharac tc rs o { - t } i c rcar o f ' t l i c c ran ium) . Both Hub l in ( l9 t j2 ) : rnc l \ r : rndcrmccrsch ( l9 t3 t )havc p t i in tcc l to thc p rcscncc in thc S tc inhc im sku l l o l 'u charac tc r in lc rp r< : tab lc as asupra in iac lbssa , and havc suggcs tcd tha t S tc inhc in r rcprcs( 'n ts a l t ro to -Ncanc l t : r tha l ;S t r ingcr ( l9 t i5 ) has . horvcr . ' c r , rc jcc tcd th is in t t . rp rc ta t ion , r , r ,h i l c sur rs t ' s t ins nonc thc l t ' sstha t th is fo rm d t :par ts l iom thc Nc: r .nc lc r t l ra ls la rgc l r , i t r p lcs ion tor l thous char : tc t tc r s ta tcs .In tc rcs t ing l ,v , most spccu la t ion on Stc inh t ' im 's l inks rv i th th t ' \ t .anc lc r tha ls has { i rcus t :c l onthc rcar o f thc sku l l . rv l t i l t ' thosc rv l to havc d is r :ovcrcc l "Nt 'unc lc r th i r l - l i k t " ' I i ' a tu rcs in lhcPctralona-Arago group I i i ivc pointccl most convin<:inql1, to chlrrat: tcrs o{ ' t l r t ' lacc (sccVandcrmccrsch, I 9f l5)

' I ' h is h ieh lv i t t compl t r t t ' l i s t ing o f ' chanrc tc rs , sp t ' r : i rncns ur rc l n rorphs is in rcnc lcc l n i t : rc lv

to bc i l l us t ra t i vc , l tnd c lcar l t ' a g rca t c icu l o { ' r ' r ' o rk r t 'n ru ins to l r t ' dor r t ' i r r sor t inq ou t t l r cnumbcr o l 'd is t inc t morphs r ( ' l ) rcscr ) t ( 'd i r r thc qcosrap l r i cu l l v scant ' rcc l l ronr in ic l f i r ss i lrccord o l - thc N ' I idd lc : rn t l La tc l ) l c is toccnc . Hou ' r ' r ' t ' r ' . i t r lo t ' s r l t ' r r rons tn l l ( ' t l ra t a t l c : ts t th r r . r 'o r Ibur (anc l c lu i tc p ro l t : t l r l v morc i l ' thc cn t i rc s l ) ( ' ( ' t runr o l n i r r tc l i l r l r , r ' t ' r t ' c 'ons id t ' r t ' c [ )s ( rpara t ( r hont in ic l g roups c l in l r t ' c l i s t in { ru ish t ' c l amonr { t l v " 'Honto :a l iuL . i ' l i r ss i l s o f ' th is t i rnc .Morc<x ' t : r , i t s t rong lv sur {g ( ' s ts 1hat , rv l r t 'n n r t 'asurcc l l ) \ , t l l ( ' s ta l l ( l l L r '< ls t l r : t t : rpp lv to o th t ' rp r imatcs thc morpho loq ica l r l i l l c rcn< ' r ' s l i t ' t r , r ' t ' c r r t l r t ' sc g ror rPs r rn rP l r ' . jus t i l \ . sPcr : i t ' sd is t inc t ions . I l ' n rv opcra t ion : r l c r i t t ' r i : r l i r r spc t ics r r ' ( { ' tn i l i ( , l l i r l ) l ) ( l r r son ic lv } l r t i r r l i r rn ia lth is i s ncccssar i l v thc casc , in co l l s ( ' ( lu ( 'n ( ' ( 'o l ' t l r c l : rc .k o1 ' r 'o r r t ' l l r t ion l r t 'nvccr r sPt ' r ' i a t io r r anc ldcgrc t : o1-n io rpho loe ic : r l chanqc; bu t us I har , t 'u l r t ' l t c l r , l r rgu t ' r l . r r ro r l l l ro loq icu l r l i l l i ' r t ' r rccs o lth t t dcgrc t r c l i sc :uss t ' c l invar i l r l r l v inc l i t :a tc : lmonq l i v i r rq I i r r lns tha t spcc i : r t ion has oc t ' r r r r< 'c l .

R ish tmi r r : ( l9 f i5 ) has s imi la r lv m: rc lc th t 'po in1 tha t . i l sP t ' c i t ' s u r ( ' ro l ) ( ' r ' i t ' r , v t ' c l us rcu l i r rt imc as wt : l l : rs i t t s lxLcr ' , i l i s r r t ccssrLr ' \ t ( , I ( ' co ran iz r ' lour spcc ics , I !onut n r t . lus , I Ion tohe ide lbergens i : , Homo neander thu le r t . t i . r unc l Hunut . ru f t i t ' tL .s . i r r t l r t ' rn ic lc l l t ' t ( ) la t ( ' l ) l c i s toc t 'n t 'human loss i l r t ' co rc l . Hc { i runc i th is schcnr t ' \ \ "an t ing . l ro r ,v t ' r '< ' r ' , in t l ru t n lu r ) \ ' ( ) r ' l r l l l l r t c rl ) l c is toccnc homin ic ls f r r rn r . , \ l i ' i ca anc l ( lh ina cLr r ro t c l t 'a r l r ' l i t i n to ( )n ( 'o l l rno th t ' ro l ' t l r< .s t 'spcc i t rs , a r rd p rc { i r r t :d Bonc l t "s ( l l i t l l ) gnrup inq o1 'a l l r rs su l rspcc i t ' s t l ' l l tn tosup iens . [Ju t soto ctrrrcludc is otr<:t ' uq:r in to us( ' t l l ( ' clcvicc o1'an al l-r 'nr l)r ircing l lonto .tr t l t i t t t . .s to olrst:u1'r , 'a r iab i l i t v tha t : rc tua l l y cx is ts , zurc l rcc lu i rcs a rn lvs is . I1 ' r \ l i . i can l rnc l ( lh in t ' s t ' l ro r r r in ic ls o lth is pcr iod sho lv : rpomorph ics s t :par i l t inq t l i ( ' l n 1 i 'om th t ' r ' ss t 'n t ia l l v l ' i u rop t . r rn s r : l r t ' rn t 'onwhich th is c l i v is ion in to spcc ics i s bascc l . i t i s t : c r ta i r r l v uppropr i : r t t ' a t l c : rs t to < 'nc lu in 'whc th t : r vc t morc spcc ics mieh t l rc in . " 'o lv t :c [ . ( ] r ' r ta in lv t l r r 'Po in t i s a l rcac lv conccc lcc l th : r tmort: morphs arc prcscl)t than r:an bc cnrbraccrl l rr , thc l , iuropcan sc. lrcnrr ' , ancl thcscmorphs rcqu i rc ana l l ' s i s and cxp lanat ion o1 ' thc k ind thcr ' r r r ( ' n ios t I i kc lv to l rc c lcn icc l i l ' thcvarc d ismisscd as r ( 'p rcscn ta t i vcs o l ' "Honto sap ien . i ' i n somt 'unc l t ' f incc l l l r lg t ' r ' s t 'ns t ' .

Rccosn iz ins a mul t ip l i c i t v o f -h r , rmin ic l spcc ics in thc n r ic [ - to - la tc l ) l t ' i s to t : t 'n t ' u , i l l , o lcoursc , eo aga ins t th t : g ra in l i r r manv pa lcoanthnrpo log is ts . , , \ l i c r a l l . th t : r t ' i s a l r t ' uu ty inl incar s imp l ic i t y tha t t l t c acs thc tcs amol l {J us , in par t i c ru la r . rvou lc l } )o ro luc tan t 1o sc( lvan ish . Noncthc lcss , th is svs tcmat ic conc lus ion is hard lv surpr is ine r ,vhcn onc cons ic lu rs thcenormous c l imat ic and sca- l t : r ' c l { luc tua t ions tha t markc( l th t : pc r ioc i . \ \ r i thou t unv c loubt ,

S P E ( ] I E S R E C O G N I T I O N t i 3

and most cspccialh' in tcmperatc lat i tudr:s, thcsc f luctuations lrasmcntt:d and rccoalcsccdhominid populat ions, thcrcbv pror. idine idcal condit ions lbr spcciat ion ancl c<impcti t ion.' fhe

potcntial that cxistt :d during thc middlc and latc Plcistoccnc lbr thc local divcrst:ncc ol-populat ions is o'cn rccosnizcd in modcls such as that ad<iptcd bv Wolpoff el a/. ( 1984), rvhoso on to posit iongstandins continuit ics r,vi thin rcsional human populat ions. Ho'"vcvt--r, i fonc acccpts that di l i 'crcnt cnvironmcnts in di l l -crcnt rcgions can procluct: ntorpholoeicaldiscontinuit ics bctrt ' t :cn populat ions, tht:n onc nrust also acccpt th: i t cnvironmcnt:r l changcin si lu can do thc samc. l) t :rccir,ccl long-tt :rm rnorpholoqical trcncls in local humanpopulat ions ovc-r this pcriocl t l icn bcr:omt'mort 'cl i l i l<:ult to clt ' { i 'nd, t-r ' t :n r,r 'hcn lavishlytr ickcd out n' i th gcnctic sophistr l ' . \ \ Ihatcvcr rcasonablc niocicl lvc acccPt ol ' thc Pl i t tcrn ol-cvo lu t ion in thc la tc r par t o l ' thc homin ic l l i r ss i l rc t :o rc l . i t i s c l t 'a r tha t r )o t a l l o1 ' thc honr in ic lvar ian ts o l - thc mid- to - la tc l ) l c is toccnc c2u) h : rvc bccn an<: t ' s t r : r l to our owl l sp t :c ics , anc l tha tthc oriein of- Iktmtt . ;apiens is to bc sougll t in a morJrh rvhost ' or 'r ' rr oriqin rvas l inkcd to :rspcci l ic gcographic rcgiolt . ( ] i r .cn thc ururtomical rarrqc ulrcaclv krrolvrr :rmong suclrmorphs i t i s 'u ' i r tua lh 'cc r ta i t t t l i i L t th is popu l : r t ion rc l t rcsor t t ' c l u c l i s t in t : t hont i r r i c l s l )cc i ( ' swh ich cocx is tcd , a t l cas t tcn tpora l l v , r , r ' i th o thcrs .

' l ' hc d iscuss ion o1 ' th t 'g r ( ' : r l c r spcc t rum o1-nr ic lc l l< ' to la lc l ) l c is toccnc hum:rn f i rss i l s , f io rn

s i tcs wor lc i ' "v ic lc . inc ' " ' i t : tb lv sp i l l s ov t ' r ' i n to t l i c p ro l t l cn ' r o l ' c lc f in i r rg Honut en t lus , n rccn t lya b l v d i s c u s s c c l b v A n d r t ' u ' s ( 1 9 t 3 ' 1 ) . R i e h t m i r t ' ( c . e . l 1 ) U 4 . l ! ) { } 5 ) , S t r i n e c r ( 1 l ) t } , t l ) , : r n r lWoocl (19t34). Str ingcr. i t t p:rr1it 'ular, makt 's thc point that , , \ [ ' r ' icarr:rnrl , , \si ;rn Honto erectu.tcan l l tr assigtr( 'd to a "qntclc" clr l l<'cl I Ionn erct lu.r, l tul nol 1o 2r sl)cci( 's o1't l rat rr;rmr'. Horvcvt 'r ' ,l ikc i t rrr ntf i , Honto eretlu-r is : I sp( 'ci( 's narnt ' : "qnrclt 's" urt ' not rccol lnizt 'cl in thlIn tc rna t ion : t l ( lodc o f 'Zoo loq i ( 'a l Nomcnc la lu r t ' . : r r r r l nc i th t ' r , in an t ,o l r j cc t i v t ' s ( 'nsc1 l r r ' ( 'th t :v {bunc l i t r t ta tu rc , Wolpo l i ' r l a1 . ( l ! , } t3 , t ) to thc r :on t r r rn ,no l rv i t } rs t : rn r l ing .

' l ' h t ' "q r : r t l c " .

indccd, i s onc o [ ' thc n tos l c lcs t ruc t i r , t ' t :a t l r l c ls tha t 1 ta l lo : r1 t I161t9 loqy [ : rs 1 r ' r ' r sc t ' r r { j t t9in l l i c t upon i t sc l f : a mt ' : t t t i t tg l t ' ss anc l unc lc l in t ' c l conc t '1 t1 , u l ) l )a r ( ' r l t l y l can ing h t 'av i l v onbra in s iz r : , tha t < :a t t bc uscd to cn tomb a l l k in t ts o l rnorp l ro loq ica l k rosc cnc ls a r rc l thusc l im ina tc thc n t ' t ' d to cx : rminc thcm. As k rnq as i , v t 'aPPlv s tanc la rc l svs t ( 'n la t i ( . t t . rm ino logr . ,to our lbss i l r t ' co rd , l v t ' s l rou ld bc c : l c : r r a l ro r r l thc mcan inr { o { t l r c tc rn rs w( 'us( ' , : rn r l w t 'shou ld bc warv o l - in t roc luc ing o t l t c rs t l la t s ( ' r \ ' ( ' < . r r r l t t r . r o l r l i sca t t ' .

' l ' hc au t l ro rs namt .c l

abovc havc c lcgant lv s l ro lvn tha t r :huruc l t ' r s t rad i t ion2r l l l , r rs t ' c l to r l t ' l i nc L Ionut t rc t lu : inc l r rdca mishmash o l 'p r im i t i r ' ' c r rn t l dc r i r "cd chur : l c t t ' r s t l r t cs . anr l tha t 1h t ' A l i i can : rn t l As i rL r rIb rms: r rc no t c l i s t i r rqu ishcc l l r r , thc san ic ayromorph i t ' s . N Ior t 'ovcr . r \nc l rc rvs n ' rakcs thcimpor tan t po in t tha t compr i r i sor ts o l 'Honut t re tLu . t t t i l h o t l r t ' r l ronr in ic l sp t ' c i< 's havc l i c t ' r rla rec lv w i tb I lonn . tu l in .s a lon< ' . ( )n lv u 'h< 'n spcc i t ' s h l r '< ' l r< ' r 'n ac l t ' c lua tc lv c lc l ln r :c lmorpho loe ica l l y can a l ) l ) l ( , l l r iu t ( r ' r )n r l ) i r r i i s r )n , j l t c n r : rc lc , anc l thc c l i s t r i l ru t ion o f ' chanrc tc rs ta tcs across spcc ics bc us t :c l to gc t ro f i r t ( 'ph l , loecr rc t i c h r 'Po th t ' scs . LuntP ine c l i vc ' rscmorpho loe ics in to "g r : rd r :s " k i l l s th is p roccss l r t ' f i r r< ' i t c : l l s tu r t .

In th is n t t t c I l tavc a t t ( 'mptcc l to ra isu ryucs t ions ra thcr than 1o so lvc thcnr . : r r rc l a l io .u ,e a l lto d raw a t tcn t io t t to t l t c l : rc t tha t l rv rnask inq thc s i { rn i l i canc t . o1 ' n tu r :h obscn,cc lmorphok te ica l 'u 'a r ia t ion , inac lcc lua tc conc( 'p ts o l ' spcc ics c lc l inca t ion havc in thc pas timpcc lcd advanc t :mct i t s in our u r tdcrs tand ing o1 ' l ro rn in ic l ph t ' logcnr ' . , \ l though i t appcarsd i l l i cu l t o r cvc t r imposs ib lc to a r r i vc a t an ob . jcc t i vc , o r un i r , t - rsa lh ' app l i cab lc , so lu t ion tothc problcm o1'spccit :s rccognit ion in thc f i issi l rct 'orcl, cognizant:t- of ' thc gcncral p:rt t t :rns ofbetwccn-spccics morpholosical r. 'ar ial ion in thc l ivinq biota r,vi l l hclp us lr t : t t t ' r t() asscss t l iatrecord . That wc do so is c ruc ia l , lo r rvc r ,v i l l makc l i t t l c usc lu l p roqr ( ' ss in human

1r-1 I . TATI ' t . ]RS, \LL

evo lu t ionar ,v s tud ics un t i l u 'c ac lop t morc rca l i s t i c n 'o rk ing c r i t c r ia o1 s l r t t i t s r ' ( r ' ( )en i t i r )n .and d t ' l l nc t l tosc sp t ' c i t ' \ \ \ r ' r c ( ( )qn izc i r r t t ' r rns o f ' c lc r i vcc l , o r l t t l t ' as t o f c l i s t inc t .morphokre ics . As I h l i vc no t t ' c l , th is i s so cvcn r rnc lc r a q rac lua l i s t n i (x l ( ' l o l cvo lu t ion . l i r rhowcvcr < inc v ic rvs thc cvo lu t ionurv p roc( 'ss . Pa lcoanthroPo loqy is a t 'onrP l r l r r t i r c t , ' i , n t c i r rl vh ich ra t iona l rc t :oqn i t ion is l r ( ' c ( ' ssar \ ,o f ' t l ) ( 'L ln i ts t ( ) l r t ' t ' onrpa l t ' c l . ( )n lv nhcn u 'c havcr c a l i s t i c a l l v c [ c l i r r < ' d t h c s p t t i c s \ \ ( . r c c ( ' l ] t r v c u , i l l l r t ' a ] r l t ' t o l r r o u c h t h t ' 1 l r ' o l r l c n i o l t l r t '

r c la t ionsh ips bc tu ' r ' cn such spcc ics . r ' r ' hc thcr l in t 'a r o l r 'o l la t t ' ru l . unc l t l r c t t i t t tu r t t p ro t : t ' cc l

towarc ls : rn undt ' rs lanc l ing o1 ' l : r rgcr -s t 'a lc pa t t ( ' rns in h r rn r : rn t ' r ' o lu t io r r .

Summary

' l ' h r : scar r :h {o r p : r t t t ' n rs i r r thc l ru rn l r r r { i rss i l n ' r 'o rc l } r rs l rc t 'n hanr l i t ' uppcr I l rv t l r< '

inac lc r lua t :y o l ' thc c r i t t ' r i l r us ( ' ( l to l c t 'oqn izc syr t ' t ' i t ' s i r r th i t t r t ' r ' o l r l . ' l ' h t ' l ' l i c t t l r : r t t : rx ic lL r tc lmorpho l rx { ic l r l changc r r t '< 'c l no t l r t ' : r ssoc iu t t ' t l r l i r cc t l r n i ( 'uns t l l r t t l r t ' r ' t ' r ' an l r t ' r ro o l r j c t ' t i vcn l ( rans o f ' s< t r t i t tq l i r ss i l s i r r to sy rc t ' i cs . I r r q t 'n< ' r : r l , l rou ' r ' r ' t ' r ' , c losc l r l t ' l l r t t ' t l p l in l r tc sp t ' t : i t ' sshorv on l t ' tn i r ro r n ror 'p l ro loq icu l r l i l ] i ' r ' t ' r r t ' t ' s I io rn o r rc l r r ro t l rc r ' . sor r rc t i rncs i r r p l r l t s o l ' t l r ca t ra tomv tha t c [o no t l ) r ( ' s ( ' r \ ' ( ' i r r thc f i rss i l r t ' r ' o rc l . ' l ' l r t ' r ' t ' i s t l rus lL t t ' r r r l t ' r rcv to unr lc r t ' s t i rnu t< 'sp t :c ics r l i vc rs i tv i r r t l r t ' l i r ss i l rccorc l . unc l w 'h i l c lu rnp ing n l i rv i i l ) l ) ( ' l r l sop l r i s t i ca t t ' t l l r c t ' t rusci t i s a r "o rv t ' r l l l a l l t i - t \ ' l ) ( ) l (x l i c : r l , too nr i rch o l iL goor l t l r i r rq h r rs i r r t l r t ' t ' n r l l r t ' r ' o rn t ' l r l i ; r l r i l i t v too u r i l t t c r p r t ' t u t i o t t o l ' t l t t ' s u l r s t l r n t i r t l r n o r ' p l r o l o g i c l r l r l i v t ' r ' s i t t t l l r t t ' x i s t s i n t l r t ' l r r r n r i r r r l i r s s i lr t t c o r c l . S u l i s p t ' c i { i t ' t ' a t t ' g o t i t ' s : r n ' r ' r ' q : r r t l c r l l r s r ' p i p l r t ' n ( ) n l ( ' n a t l r i r t < l o r r o t . j t r s t i l r ' t h t ': r t t c l l t i o l t u c c o r r l t ' r l t o s p t ' < ' i t ' s , s o i t i s c l i t i c a l t l n t . n v l r c r t ' r l i s t i r r c t n r o r p l r s t ' x i s t i r r t l r r ' l o s s i lrccorc l t l t t ' r ' : t r t ' n ( ) t r ( ' l ( ' g1r t ( ' ( l to su l rspcc i l i c s tu t r rs rv i t l ro r r t goor l rc i rsor r . S t ' r ' r ' r ' i r l r r ro r 'p l rs u . r ' t 'c l i s t i n e u i s h a l r l c i r r t l r r ' l a t c r p r r l t o l ' t l r t ' l r u r n r r n l o s s i l r c t o r r l t l u r t l r l t ' r l i s t i n c t l i o t l r l i ' o r n t ' : r c l ttr t l rcr :r t tr l f rrrnr t l r t ' l iv ing spt 'cics I lono :uft iot.s. ' l ' l r t 'st ' int ' l rrr l t I lortro rrcttrulrr lhulnr.t i .r , I lorttr the ide lb t ' rgen.s i .s . anc l p ro l i : t l r l v a lso I Ion to . t le i r t l t t i t r r r ,a . r r . r . l ' l r t ' r ' t ' r ' oqr r i t io r r o l ' r r ro r 'p l rs i r r th t ' l i r t t ' r 'par t o1 ' t l t t ' hunrar t I i r ss i l r '< 'cor r l l l r s l rccn lx rs t ' r l l l r rgc l r ' ( )n t l ) ( ' l r ss t ' r r r l r lug t ' o l ' l oss i l s l i ' onrl ' i u r o p c , : r t r c l r n : r r r 1 ' l i r s s i l s l i o r r r ' \ s i ; r l n r l , , \ l i i t l t t l o r r o t l l t t l r t ' l ' , t r l o p t ' u r r s c l r c r n t ' . l ) r o l r l r l r l vo t h c r s p t ' c : i t ' s r v i l l 1 ; r o v t ' i t l t ' r r t i l i i L l r l t ' l i o t t t o t l t t ' r ' : r l t ' l r s o l ' t l r t ' u . o r l r l u l r t ' r r t l r c a l i p l o l r l i r r t t 'c o r r t p l r r i s o t r s a n ' r n u r l t ' . ' l ' l r t ' t ' x i s l t ' r r c t ' o l ; r r n u l t i l r l i < ' i t r o l ' l r o r r r i r r i r l s p r ' < i c s i r r t l r < ' n r i r l r l l c t ola tc l ) l c is to t ' r ' r t c u ror r r r r l th t 'u 'o r l< l rvou lc l no t l r t ' s r r r 'p r i s inq i r r v i t ' .nv o l ' t l r t ' g rcu t t l i n r : r t i c l rn r l( tus ta t i c os t ' i l l a t ions . t lo t r l r t l t ' ss i r t vo lv i l tq th t ' l i t ' r ;u t 'n t l i l s -n rcn t lL t ion l r r r r I r ' t '< 'o : r l csccn t ' t ' o lI tumat t popu l lL t ions , th l r t r r l r rkc r I t l r c p t ' r io r l .

Acknowledgements

' l ' h is : r r t i c : l c r ' vas in rprovcr I l rv th r ' ( ' ( )n ln l ( ' l l l s o l l ' ] r ' i c l ) t ' l son . N i l t ' s l l l r l l t ' r lg t ' . i tn r l th r t ' t '

: tnonvmoLls r t ' r ' i t ' r v t ' r s . I to r t t 'o l 'w ' l ro rn n t ' ccss l r r i l v ag l t ' t ' s u i t l r l r r rv t l r i r rg i t cor r t r r i r rs . I l l r r ri n c l c b t c t i t o t h c R i < ' h a r r l L o u r r s l r t ' r r , l i o u n r l a l i o n l i r r i t s ! l ( ' 1 1 ( ' r ' ( ) u s s u l ) l ) ( ) r t . ' l ' h i s i s

C l r in t r ibu t ion r ro . l t J o l ' th t ' l ,ounsbcrv l , r r l ronr to rv o l l J io l rx t i c l r l ' \ r r l l r roPo logr , . r \n r t ' r i car rM u s c u m o f ' N a t u l r l I l i s t o n .

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