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ARTICLES

Huma n Evolution in the Middle Pleistocene:The Role of Homo heide lbe rge nsisG. PHILIP RIGHTMIRE

For paleoanthropologists working in

the Middle Pleistocene, these are inter-

esting times. New discoveries of arti-

fa c t s a n d h u m a n fo ss il s h a v e b e en

reported from western Europ e, so that

it now looks as though this continent

w as populated 800, 000 years ago, if

not earlier. O ne of th e fos s ils, from

Ceprano in Italy, is described as H o m o

erectus. W hether th is ancient s pecies

ever reached Eu rope has been repeat-

edly questioned, but the Ceprano cra-

nium is com plete enough to provide

some har d evidence.

O t h er fi n d s f r om S p a in a r e e ve n

m o r e s p ec t a cu l a r. T h e S im a d e lo s

Hu esos (Pit of Bo nes) in the Sierra

de A tapuerca h as yielded a w ealth of

s keletons that are bes t interpreted as

early N eanderthals , perhaps clos e to

300,000 years in age. Older but unfor-tunately m ore fragm entary rem ains ,

also from Atapu erca, display no Nean-

d e r th a l f e a tu r e s a n d a r e c la i m ed a s

represen tatives of a new sp ecies. Ho mo

antecessorwill require close study.

These European dis coveries focus

fres h attention on the evidence accu-

m ulating from Africa and As ia. H u -

ma n bones are known from th e earlier

Middle Pleistocene of Africa at locali-

t ie s s u c h a s B od o i n E t h io p ia a n d

B r ok en H i ll i n Z a m b ia . T h e c r a n ia

s how anatom ical features that distin-

guish them from Hom o erectus. In the

F ar Ea s t, the people at D ali and other

s it es a r e a ls o m o r e a d va n ce d t h a n

Homo erectus, but their affinities to

groups in the West are un certain.

This Middle Pleistocene record, still

s p a r se b u t i n cr e a si n gl y w e ll d a t ed ,rais es im portan t ques tions . O ne con-

cerns the fate ofHom o erectus in d iffer-

ent r egions of th e Old World. Another

is how m any distinct species should be

recognized am ong the des cendants of

this ancient lineage. It is apparent that

t h e t r a d it io n a l a p p r o a c h o f l u m p i n g

diverse hu ma ns to gether as arch aic

Hom o sapiens w ill n o lo n ge r w o r k.

T h e p i ct u r e i s h i gh l y c o m p le x, a n d

s everal taxa p robably are n eeded to

accom m odate the fos s ils . Evolution-

ary relationships among these popula-

tions must be clarified, but pose some

major problems. I will address only a

s u bs et o f t h es e t o pi cs p e rt a in i ng

m ainly to earlier M iddle P leistocene

hom inids .

HOMO ERECTUSIN PERSPECTIV E

This extinct species has been at the

center of m uch controvers y. A t pre-

s ent, there is no fi rm cons ens us as to

whether it should be defined as a long

lasting, polytypic lineage or as a group

of r elatively specialized populations

g eo gr a p h i ca l ly c o n fin e d t o t h e F a r

East. In my view, Hom o erectus origi-

n a t e d i n Afr i ca a n d t h e n s p r e a d t o

Euras ia. The hypodigm is m a de up of

s pecim ens from J ava, Zhoukoudian,

a n d o t h er s it e s i n Ch i n a , T er n i fin e

(now Tighenif) in Northwest Africa,

O ld u va i G or g e, t h e Tu r k a n a B a si n ,

and Swartkrans in South Africa (Fig.

1). Several fossils recently discovered

in w es tern A s ia and in Europe prob-

ably s hould be counted as w ell. This

species spans an interval of at least 1.5

million years. Indeed , some East Asian

populations m ay have s urvived into

later Pleistocene times.

As docum ented b y Weidenr eich, von

Koenigswald, Le Gros Clark, and oth-

e r s, m e m b e r s o f t h i s t a x on s h a r e a

s uite of characters by w hich they can

be distinguished from recent humans.

S om e of the principal differences re-

late to cra nial capacity, keeling on the

m idline of the vault, parietal length,

occipital proportions , the anatom y of

the cra nial base, facial projection, the

fo r m o f t h e m a n d i b u la r s ym p h y si s,

tooth size, the relative narrowness of

the pelvis, and th e length of the femo-

r a l n e ck . A la r ge n u m b er o f t r ai ts

generally describe Homo erectus an d

diagnose this species relative to living

people.

These as s um ptions have been chal-

lenged by several workers, on highly

diverse grounds. One point of conten-

t io n c o n ce r n s t h e m a t e r ia l f r o m t h e

Tu r k a n a B a si n . I t h a s b e en c la i m e d

that the early Kenyan crania lack spe-

cial features developed by the Asian

p o p u la t io n s . A m i d li n e k ee l o n t h e

vault, an angular torus at the postero-

inferior corner of the parietal bone,

c er t a in c h a r a ct e r s o f t h e b a s e, a n d

overall thickening of the braincase a re

s aid to be abs ent from the s pecim ens

at K oobi F ora but w ell expres s ed in

the rem ains from Trinil, Sangiran, an d

Zhoukoudian. These differences have

prom pted investigators, including An-

drews,1 Groves,2 and Larick and Cio-

chon,3 to recognize two species and to

s ugges t that Homo erectus m u s t b e

geographically res tricted to the F ar

East. Wood 4,5 agrees , on the bas is of

fa c ia l m e a s u r em e n t s, p e r h a p s s o m e

aspects of tempora l bone m orphology,

and dental differences , that the early

African hom inids s hould b e s et apart

G. Philip Rightmire is Professor of Anthro-pology at the State University of New Yorkat Binghamton. In his research, which fo-cuses on the evolution of the genus Homo,he has studied fossils from many of theimportant prehistoric localities in Europe,Africa, and the Far East. He is particularlyinterested in questions about Homo erec-tus, Middle Pleistocene hominids, and theorigins of more modern humans.

Key words: species; phylogeny; Homo erectus;Homo heidelbergensis;Neanderthals

218 Evolutionary Anthropolog y


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from later Homo erectus. W ood now

refers the Turkan a Basin specimens to

Hom o ergaster, w hich, in h is opinion,

is m ore likely than Homo erectus to

have played a role in the evolution of

later people.

A quite different interpretation has

been offered by Wolpoff and cowork-

ers,6 w h o n o w c l a im t h a t t h e n o m e n

Hom o erectus i s u n n e c es sa r y a n d

should be discarded altogether. Here

the question is really whether there is

continuity from earliest P leistocene

t im e s r igh t t o t h e p r es en t ; t h a t i s,

whether just one long lineage, with no

bran ches or extinctions, can be r ecog-

nized. S uch a l ineage w ould include

t h e a n c ie n t Tu r k a n a p o p u la t io n s a s

well as those later resident in Africa

a n d E u r a si a. I f i t c ou ld b e d em o n -

strated that no splitting had occurred,

one m ight argue that any division be-

tween taxa wou ld have to be ar bitrary.

Wolpoff et al.6 and Wolpoff7 (see also

Tobias 8) go a step further and say that

there s im ply is no bas is for keeping

m o r e t h a n o n e s p e ci es , w h ic h m u s t

then be Hom o sapiens.

It does not seem to me that either of

these scenarios can be supported fully.

Certainly there is geographic variation

a m o n g t h e s eve r a l a s se m b la ge s o f

Hom o erectus, but th e fossils from th e

Turkana Bas in, O lduvai G orge, and

other sites in Northwest Africa exhibit

essentially the same set of traits as do

t h os e f r om t h e F a r E a st .9 Discrete

c h a r a c t e r s s a i d t o b e u n i q u e t o t h e

Asian pop ulations are variable in their

expres s ion and, in fact, m os t can be

identifi ed in the earlier Eas t A frican

material.10 Cranial dim ens ions s how

m uch overlap.11,12 Vault thickness, as

m e a su r ed n e ar t h e ju n c ti on o f t h e

frontal and par ietal bones, is about the

s am e in t h e Afr ic an a n d Asi an

samples.13 The faces of KNM-ER 3733

from Koobi Fora and KNM-WT 15000

from N ariokotom e conform in nearly

all res pects to the anatom y of H o m o

erectus as reconstructed from the San-

giran and Zhoukoudian s pecim ens .14

Also, the teeth from the Turkana Basin

a r e c l o s e i n s i z e a n d s h a p e t o t h o s e

from Zhoukoudian.15 Apparen tly there

are not m any traits that can be used to

diagnose Hom o ergaster, and probably

jus t one polytypic s pecies s hould be

recognized. Nevertheless, Hom o erec-

tus, as broadly defi ned, d oes pos s ess

ma ny anatom ical distinctions, extend-

ing not only to the s kull and teeth bu t

to the postcranial skeleton as well. All

of the better-preserved individuals, in-

cluding even the late s urviving on es

fr o m s om e o f t h e F a r E a s te r n s it e s,

can be s et apart from Homo sapiens.

The boundary betw een thes e taxa is

not ar bitrarily defined.

NEW EVIDENC E FROM EURASIA

Al th o u g h t h e r e i s d i sa g r ee m e n t

about taxonomy, most workers would

concede that populations res em bling

Homo erectus dispersed from Africa

into Euras ia w ell before 1.0 m illion

years ago. These m ovemen ts occurred

p r o b a b ly o ve r a l on g p e r io d . T h e

h o m in i ds m a y h a ve m a d e r e pe at ed

s orties , introducing crude chopping

tools a nd s tone fl akes or Acheulean

han daxes into different regions.16

Evidence docum enting the s pread

o f h u m a n s i n t o w e s te r n As ia c o m e s

from several sites, including Dmanisi

in Geor gia an d Ubeidiya in Isr ael. AtDmanisi, a lithic industry and a well-

pres erved m an dible w ith teeth have

been r ecovered.17 The jaw res em bles

t h o se o f Hom o erectus.18 T h e b o n e -

bear ing levels overlie a lava flow da ted

at 1.8 million years, but the age of the

tools and fossils remains problemati-

cal.16 At Ubeidiya, an extensive ma m-

ma lian fauna excavated from lake sedi-

men ts suggests a relatively cool climate

at a date of perhaps 1.4 to 1.0 million

Homo erectus, a s

broadly defined, doespossess ma ny

a natomica l distinctions,

extending not only to the

skull a nd tee th but to the

postcra nia l skeleton a s

well. All of the be tter-

preserved individuals,

including e ven the late

surviving one s from someof the Fa r Ea stern sites,

ca n be set apa rt from

Homo sa piens.

Figure 1. A ma p showing the principa l loca lities that have yielded fossil rema ins of Homo

erectus. Cep rano in Italy is the first site to dem onstrate that the spe cies rea che d Europe,

proba bly before the o nset of the Middle Pleistoc ene.

ARTICLES Evolutiona ry Anthropolog y 219


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years . In addition to the fauna, there

are s tone choppers , s pheroids , picks ,

and bifaces , w hich Tchernov19 com -

pared to the lithic material from u pper

B ed I I a t O ld u va i G or ge . H o w t h e

differences betw een the non-Acheu-

lean and Acheulean as s em blages at

this Jordan Valley site should be inter-preted is unclear, but the tools m os t

prob ably were used by group s ofH o m o

erectus.

A ncient traces of hom inid activity

a r e f o u n d a l s o i n E u r o p e , a t F r e n c h

localities such as Le Vallonet, Isernia

in Italy, and in the Neuwied Basin in

German y. This scattered archeological

evidence, cons is ting m ainly of core-

choppers an d flakes, sometimes found

w ith broken an im al bones taken to b e

fo o d w a s t e, m a y d e m o n s t ra t e a h u -

m a n p r es en c e i n t h e E a r ly P le is -

tocene20 (but also s ee Roebroeks21).U nfortunately, the oldes t European

s ites have not produced m ore than a

few bits of huma n skeleton. A notable

exception is Ceprano in central Italy.

H e r e a fr a gm e n t a r y b u t fa i r ly c o m -

plete brainca se was discovered in 1994.

T h e f o ss il w a s p i ck ed u p i n c la y

depos its , w hich contain no volcanic

ma terial that is directly datable. Potas-

s ium -argon dates have been obtained

fr o m vo lc a n ic s an d s h i gh e r i n t h e

stratigraphic sequence; these are said

by As cenzi and cow orkers22 to indi-

cate an age greater tha n 700,000 years.

Insofar as can be determined from the

description p rovided by its discover-

ers , this hom inid dis plays the heavy

continuous brow, low vault, an gled

occiput, and thick cranial bones that

a r e c h a r a ct e r is ti c o f Hom o erectus.

T h is i s i m p o r t a n t i n fo r m a t i on . Al -

though the Ceprano specimen is dam-

aged in som e key respects, it seems to

confi rm the identity of one group of

p eo p le w h o e n te r ed E u r op e i n t h e

Early Pleistocene.

Representa tives ofHom o erectus also

reached the East Asian tropics before

m oving into m ore tem perate regions.

I t h a s b e e n a s s u m e d t h a t m o v e m e n t

i n t o t h e F a r E a s t b e g a n 1 .0 m i ll io n

years ago or slightly earlier, but radio-

m e t r ic d a t es f ro m m i n e r a l s a m p le s

collected at Modjokerto and Sangiran

now s ugges t that the oldes t Indone-

sian loca lities may be 1.8 to 1.6 m illion

years old.23 If this res ult can be veri-

fied, then it will look as thou gh H o m o

erectus spread quite rap idly across the

Old World. These hom inids flourished

fo r a l on g t im e . At s it e s i n c lu d i n g

Zh o uk ou d ia n a n d Lo n gt a nd on g

(Hexian) in China, the species is known

from d eposits of the later Midd le Pleis-

tocene, while at Ngandong in Java, at

least one group of archaic people may

h a v e s u r v ive d i n t o t h e L a t e P l e is -tocene.24 P opulations s uch as that at

N gandong m ay docum ent the las t ap-

pearan ce of the lineage.

SPECIATION IN AFRICA?

The picture em erging is one ofHomo

erectus as a widespread, polytypic spe-

cies, with groups persisting longer in

s om e regions than in others . The pat-

tern docum ented in China and es pe-

cially in Java contr asts with that in the

West, where Hom o erectus s eem s to

d i s a p p e a r f r o m t h e r e c o r d a t a r e l a -

tively early da te.25,26 Also, it is int erest-

ing that the A sian popu lations appa r-

ently are more specialized in the sense

o f e xh i b it i n g a h i gh e r i n ci d en c e o f

some morphological characters associ-

ated w ith cranial robu s ticity. These

traits are subject to geographic varia-

tion and do not mark a species bound-

ary, but they m ay nevertheless delimit

groups that had d ifferent evolutionary

fates.

There is no reas on to s uppos e that

a ll d e m es o f Hom o erectus evolved

furth er. The evidence is consistent with

e ve n t u a l e x t in c t io n o f s o m e o r a ll

p o p u la t io n s i n t h e F a r E a s t . A l es s

specialized branch of the species may

well have given rise to later h um ans. 9,27

T h is b u d d in g o f a d a u g h te r li n ea g e

from Hom o erectus m u s t h a ve o c-

curred very early in the Middle Pleis-

tocen e, if not b efore. African a nd west -

ern Asia are likely areas in which the

fir st m o r e a d va n ce d h u m a n s o r ig i-

na ted (Fig. 2). An Africa locu s is consis-

tent w ith fi ndings from archeology,e n vi r on m e n t a l r e co n s tr u c t io n , a n d

pattern s of animal dispersal.28

Fossils that shed light on this specia-

tion event have turned up at s everal

localities in Africa. One is Bodo in the

Middle Awash r egion of Ethiop ia. The

B o do c r a n iu m a n d , l a t e r, a b r o ke n

par ietal from a second individual, were

found in conglomerates and san ds con-

taining mammalian bones and Acheu-

lean ar tifacts.29,30 F auna from the s ite

has been com pared to that from Bed

IV at Olduvai Gorge and Olorgesailie

in K enya, and an early M iddle P leis -tocene age is indicated. N ew argon-

a r go n d a t es r e p or t ed b y C la r k a n d

colleagues31 s upport this biochronol-

ogy. The evidence from faun a, arch eol-

ogy, a nd laser-fusion determina tions

p o in t s t o a n a ge o f a b o u t 6 00 ,0 00

years for the Bodo h ominids.

The face and anterior portion of the

braincase are reasonably complete; it

can be es tablis hed that Bodo is l ike

Homo erectus in s om e features . The

ma ssive facial bones, projecting br ow,

low and constricted frontal with mid-

line keeling, par ietal angular torus ,

and thick vault give th e s pecim en an

archaic appearance. In other respects,

t h e c r a n i u m i s m o r e a d va n c ed i n i t s

morphology. Brain size is close to 1,300

cc, which is substantially greater th an

is expected for Hom o erectus. The fron-

tal bone proportions , arched s hape of

t h e s qu a m o u s t em p o r al, a n d s om e

traits of the cran ial base are like those

o f m o r e m o d er n h u m a n s . Al th o u g h

t h e f ac e i s v er y b r o a d a n d h e a vi ly

cons tructed, the s upraorbital tori are

divided into m edial and lateral s eg-

men ts, the mar gin of the nose is verti-

cal rather than forw ard s loping, and

t h e i n ci si ve c a n a l o p e n i n g i n t o t h e

f r o n t o f t h e h a r d p a l a t e s h o w s a d e -

r i ve d c o n d it i on p r e se n t i n r e ce n t

Homo.32

This mix of characters suggests that

the Middle Awash individua ls are in-

term ediatein their m orph ology. How-

ever, s everal of the res em blances to

Hom o erectus are plesiomorphies that

c a n n ot b e c on s id e re d d ia gn o st ic .

The pa ttern doc umented

in China and espe cially

in Ja va contra stswith

tha t in the West, whereHomo erectusseemsto

disa ppe a r from the

rec ord at a relatively

ea rly da te. Also, it is

interesting that the Asianpopulationsapparently

are more specialized.

220 Evolutionary Anthropolog y ARTICLES


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Moreover, it is clear that the cranium

shares other apom orphic features with

m o r e m o d e r n p o p u la t io n s . I t s ee m s

r e a so n a b le t o g r ou p B o do w it h t h e

fam ous fos s il from Broken H ill (K a-

bw e) in Zam bia (F ig. 3), along w ith

s p ec im e n s fr o m E l a n d sfo n t e in i n

South Africa, Lake Ndutu in Tanzania,

and probably Eyasi, also in Tanzania.

These localities are Middle Pleistocene

in age. In addition to th e hum an skull-

cap, defl ation s urfaces (bays ) at

E l a n d sfo n t ei n h a ve yi eld e d a l a rg e

fa u n a , t o ge th e r w it h Ac h eu le a n

han daxes. Dating of th is assemblage is

complicated by the fact tha t several of

t h e e x t i n c t m a m m a l s p e c i e s a r e u n -

k n o wn e ls ew h e r e, b u t c o m p a r is o n s

with other African sites imply that th e

b o n e s w e r e a c c u m u l a te d b e t we e n

700,000 an d 400,000 years a go.33 Ani-

mal remains possibly associated with

the Broken H ill cranium s ugges t an

age within this same broad interval. 34

As h a s b e en r e co gn i ze d fo r s o m e

time, the African h ominids a re similar

to other, roughly contemp orary people

know n from Europe. Like that from

B r ok en H i ll, t h e c r a n iu m f ro m P e -

tralona in Greece is particularly well

preserved (Fig. 4). Although there is

doubt about both its original prove-

nience within layers of stalagmite de-

posited on the cave floor and its asso-

c ia tio n w it h a n im a l b on e s, t h is

individual is of Middle Pleistocene an-

tiquity. Some of th e flowstone m ay be

about 200, 000 years old,35 b u t o t h e r

fos s ils from the s ite im ply a greater

age for th e cave contents (s ee Cook

and cow orkers36 for a review). In any

cas e, the P etralona and Broken H ill

crania differ only slightly in orbit size,

frontal proportions , and prom inence

of the torus crossing the occipital bone;

in general, they are remarkably alike.

R es em b l an c es a r e a p p a r en t i n t h e

height, breadth, and m assive construc-

tion of the upper face and cheek, sev-

eral m eas ures of projection in th e fa-c ia l m i d li n e , c o n fi gu r a t i on o f t h e

thickened brows, and many aspects of

vault shape.9 Becaus e the Bodo m ate-

rial is less complete, comparisons be-

tw een it and the G reek cranium m us t

be m ore limited in scope, but here also

there a re similarities.32

M ultivariate s tudies of s kull form

have been car ried out by several work-

ers, who noted resemblances between

t h e Afr ic a n a n d E u r o pe a n s p ec i-

m ens .3739 Van Vark40 has us ed 17 di-

m ens ions of the face and braincas e to

c o n st r u c t a m e a s u r e o f g en e r a li ze ddis tance (D2) , w h ic h s h o ws t h a t t h e

Broken Hill and Petralona specimens

differ from on e anoth er less than is the

case for Upper Paleolithic and recent

hum ans. When a reconstruction of the

partial cranium from Arago Cave in

France is included in this analysis, it

also falls clos e to that from Broken

Hill. There seem to be good phenetic

grounds for lum ping thes e hom inids

together.

O t h er a n c ie n t E u r o p ea n fi n d s a r e

m ore fragm entary. H um an bones and

teeth have been recovered from the

quarry at Bilzings leben in G erm any

and from several earlier Middle Pleis-

tocene sites in Italy, while an occipital

bon e is o n rec ord from Vert esszollos

i n H u n g a r y. T h e r e i s a l so t h e r e a r

p o r t io n o f a b r a in c a se fr o m S w a n s-

combe in En gland. The man dible from

M auer in G erm any an d a t ibia as s oci-

ated w ith A cheulean bifaces at Box-

grove in E ngland are arguably am ong

the oldest hominids from Eu rope. Both

are on the order of 500, 000 years in

age.41 O f c o u r s e t h e r e a r e q u e s t i o n s

about the a ffinities of this mater ial, as

few anatom ical clues are pres erved.

But if the Mauer mandible is grouped

with the other specimen s from E urope

and Africa, then the ent ire assemblage

can be referred to Hom o heidelbergen-

sis.9,42,43

Th is sp ecies w a s n am ed b y

S c h oe t en s a ck i n 1 90 8 t o a c co m m o -

date the jaw found a year earlier in the

bas al s and and gravel com plex of the

G ra fe n r ai n p it n e a r H e id e lb e rg .

Figure 2. A tree illustrating the evolution and geo grap hic d istribution o f Homo in th e Pleis-

tocene. Homo erectusisassumed to ha ve o riginated in Africa and then spread quickly to Asia

and probably to Europe. Homo heidelbergensisisdistribut ed from Africa into Eurasia d uring t heMiddle Pleistoc ene. Whether this spe cies reac hed the Far East is still a question. Europea n

Homo heidelbergensis gave rise to the Neand erthals, while an Africa n b ranc h of Homo

heidelbergensisisanc estral to mod ern huma ns. Four spec iation events(S) are depicte d.



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Schoetensack was impressed with the

prim itive character of his foss il bu t

recognized that i t m us t be hum an, asthe canines are redu ced in size and the

tooth crowns generally display the pro-

portions expected for modern popula-

tions.

Later authors continued to em pha-

s ize the prim itive appearance of the

ma ndible. Howell44 pointed to its mas-

sive construction, multiple mental fo-

r a m i n a , a n d ve r y t h i c k s ym p h y s is ,

w hich lacks any indication of a chin,

a s c h a r a ct e rs s h a r ed b y o t h e r e a r ly

repres entatives of Ho mo . However,

H ow ell w as careful to note that other

features of the specimen , including itsram us breadth, relatively great ante-

rior depth of the corpus, and m oderate

s iz e o f t h e d e n t it io n , d i st in g u is h i t

from both F ar Eas tern Hom o erectus

and the Ternifi ne people. H e argued

that the M auer hom inid m us t be s pe-

cifically distinct from archa ic lineages

in Asia or Northwest Africa. Howell

left open the relations hip of this is o-lated fossil to other European groups,

i n c lu d i n g t h e N e a n d e r t h a l s. W h i le

there are still obvious difficulties with

linking the m andible to individuals

such as tha t from Petralona, for which

no lower jaw has been recovered, the

fo s si l c a n b e lu m p e d w it h e a r li er

Middle Pleistocene hu ma ns in the way

I h a ve o u t li n ed . As d e fin e d o n t h is

basis, Hom o heidelbergensis retains a

num ber of archaic characters and m ay

be the stem from which both Neand er-

thals and m odern people are derived.

ALTERNATIVE VIEWS OF HOMO

HEIDELBERGENSIS

J us t as there is controvers y about

w hether Hom o erectus should be parti-

t io n ed i n to t wo t a xa , s o t h er e a r e

questions about the m ake-up ofH o m o

heidelbergensis. Some authorities hold

that the E uropean and African s peci-m ens s hould be s et apart as repres en-

tatives of distinct lineages. Proponents

o f t h i s v ie w a g r e e t h a t t h e F r en c h ,

G er m a n , a n d G r ee k f os si ls s h a r e a

s eries of features w ith the hom inids

from Bodo and Broken Hill. But they

c la i m t h a t e ve n t h e e a r li es t M id d le

Pleistocene Europeans exhibit apomor-

phic traits that align them only w ith

Neanderthals. 4549

In this reading of the evidence, the

fossils from Ma uer a nd p robably Box-

grove, Arago Cave, P etralona, Bilz-

ingsleb en , Vert esszollos, an d Swa ns-combe document a single line evolving

tow ard the populations at S teinheim

in G erm any and A tapuerca in S pain.

These assemblages can be referred to

Hom o heidelbergensis. However, it is

recognized that there is no clear sepa-

Figure 3. Facial and lateral views

of the Broken Hill c ranium. Bro-

ken Hill is one of the most com-

plete Middle Pleistocene speci-

mens, here attributed to Homo

heidelbergensis. Along with in-

creased brain volume, the cra-

nium e xhibits features of the na-

sal region, palate, occiput and

base that distinguish it from Homo

erectus.

Figure 4. Late ral and fac ial

views of the Petralona cra-

nium. This individua l from

Greece resemblesthat from

Broken Hill in ma ny metric

features relat ing to facial

proportionsand va ult shap e.

Whether the Petralona face

a l so d i sp l a y s c h a r a c t e rs

unique to the Neanderthal

lineag e iscurrently deb ated .



6/10

ration of the latter from early Neander-

t h a ls s u ch a s t h o se fr o m B ia c h e i n

F rance, Ehrings dorf in G erm any, or

Saccopastore in Italy. Eventually this

same lineage produ ced the classic

Neanderthals of the Late Pleistocene.

So an alternative classification is p os-

sible in which not only t he classicpopulations, but also all the older fos-

sils, are placed in one species termed

Hom o neanderthalensis50 (see also Car-

bonell and cow orkers51 and Arsuaga

and coworkers52).

Accord ing to th is accr etion hypo th -

es is , distinctive N eanderthal charac-

ters appear first in the facial skeleton.

Ad vo c a te s o f t h e m o d e l a r g u e t h a t

s u ch t r a ce s c a n b e i d e n t ifi ed i n t h e

M a u e r a n d Ar a go r e m a in s . At l at e r

evolutionary stages, apomorphies ac-

cum ulate in the occiput and fi nally in

the tem poral region. I t is s ugges tedthat the ancestors of Neanderthals be-

c a m e i n c re a si n gl y i so la t e d t h r o u gh

tim e as a cons equence of colder cli-

mate conditions. In the second half of

t h e M id d le P le is to ce n e, b a r r ie r s

formed by glaciers and associated tun -

d r a t o t h e n o r th , i c e s h ee ts i n t h e

moun tains to the east, and the Mediter-

r a n e a n t o t h e s o u t h r e d u c e d c o n t a c t

and gene exchange w ith people out-

s ide Europe.49 Is olation in this rela-

t ive ly h a r s h e n vi r on m e n t l ed t o t h e

full expression of the m orphology that

distinguishes Neanderthal skulls and

postcranial bones from those of other

populations.

Much of this scenario seems sound.

Certainly it is easy enough to track th e

N e a n d e r t h a l s b a c k i n t i m e t o S t e i n -

h e im o r e ve n t o S w a n sc o m b e. T h e

occipital bones of both specimens d is-

p la y s ig n s o f a s u p r a in i a c f o ss a ( a

centrally placed elliptical dep ression

with a pitted floor). Moreover, Swans-

c o m b e p o ss es se s a t r a n sve r se t o r u s

t h a t i s w ea k n e ar t h e m id li n e b u t

bilaterally projecting. These tra its are

diagnos tic for the lineage. The S im a

de los H ues os at A tapuerca confi rm s

that N eanderthal features are pres ent

in an assemblage that may be close to

300,000 years in age.53

As d e sc r ib e d b y Ar s u a ga a n d h i s

colleagues,47,52 the Sima skulls present

a com bination of ples iom orphic and

derived char acter s. The well-preser ved

fa c e o f c r a n iu m 5 i s q u i t e l a r ge i n

relation to the braincas e. This , by it-

self, is not a Neanderthal apomorphy,

b u t t h e t op o gr a p hy o f t h e m i d fa c e

seems to anticipate that of later popu-

lations . The infraorbital s urface and

t h e s i d e w a l l o f t h e n o s e m e e t a t a

shallow angle, producing a slight con-

cavity. The cheek region is thus not

inflated in the extreme ma nner of

N eanderthals , but can be interpretedas interm ediate in form . A ls o in the

S im a s a m p le , b r o w s a r e ve r y t h i c k.

Continuity of the s upraorbital tori at

glabella is s aid to be rem inis cent of

N eanderthals . A t the rear of the cra-

nium , the s uprainiac area is large but

not very depressed. This trait and the

s hape of the occipital torus s eem to

fo r es h a d ow t h e N ea n d e r th a l c o n di -

tion.

Earlier in the M iddle P leistocene,

Neanderth al roots are m ore difficult to

fi n d . S om e a u t h o r s h a v e p o i n te d t o

Vert esszollos or even Bilzingsleb en as

documenting evolutionary continuity,

but m os t of this m aterial is too frag-

mentary to provide convincing infor-

m ation. The jaw from M auer is com -

p le t e b u t , i n f ac t , s h o ws fe w i f a n y

t r a it s t h a t c a n b e t a k e n a s s p e c ifi c

links to later European populations .

M ore crucial to the accretion hypoth-

es is , or at leas t the vers ion of i t that

encom pas s es the earlies t fos s ils, are

the crania from Arago and P etralona.

Here the question is whether there are

signs of incipien t Nean dert ha l m or-

ph ology, especially in the facial region.

The face of the pa rtial cranium from

A rago is largely com plete but unfor-

t u n a t el y d a m a g ed a s a r e su l t o f i ts

long interment in compacted cave sedi-

m ents . The frontal bone, interorbital

pillar, nose, and cheeks show numer-

ous cra cks; localized a reas of crush ing

are a lso present. The discoverers h ave

been able to correct some of this dam -

age in a reconstruction,54,55 but signifi-c a n t d i st o r ti on r e m a in s . I n s p it e o f

these problems, some workers can dis-

cern definite resemblances to Nean der-

thals. Hublin 49 notes that the infraor-

bital surface of the ma xilla is flattened

a n d t h e c h ee k b o ne s a r e o b li qu e ly

oriented. Arsuaga and colleagues 52 sug-

gest that the Arago midface is actua lly

m o r e N ea n d e r th a l -li ke t h a n t h a t o f

Sima cran ium 5 in the extent to which

the infraorbital plate and the side wall

o f t h e n o s e a r e c o n ti n u ou s , a n d t h i s

s urface is inflated or convex. Also,

there is m uch forw ard protrus ion of

the face at subspinale (in the midline,

j u st b e lo w t h e n a s a l o p e n i n g) . T h e

nos e itself is l im ited inferiorly by a

sharp rim , as in Neandertha ls.

T h es e o b s e r va t io n s m u s t b e t e m -

p e r e d b y t h e f a c t t h a t c r a c k i n g a n d

plastic deformation make it difficult to

as s es s s om e key as pects of m orphol-

ogy. The wall of the Arago maxilla is

generally flattened or even inflated in

the manner characteristic of Neander-

t h a ls , b u t t h e r e i s s li gh t h o llo w in g

laterally, below th e or bit. This cann otbe dis counted a s due entirely to dam -

age. Also, it is n ot clear that the zygo-

m atic bone is s w ept back (obliquely

oriented) as n oticeably as it is in later

populations . In facial forw ardness at

s ubs pinale, as m eas ured by the zygo-

ma xillary an gle of Howells,56 the Arago

cranium , at 113, is in the Neanderth al

range, and the Petralona specimen, at

11 8, s h o w s a l m o s t a s m u c h p r o t r u -

sion. But the value for Broken Hill is

only 116. Consequently, a low zygo-

m axillary angle does not neces s arily

align Arago and Petralona with Nean-

dertals rather than w ith other M iddle

Pleistocene specimens. The sharp infe-

rior m argin of the A rago nos e is in-

deed rem inis cent of that in N eander-

thals. H ow ever, there is variation in

this feature. P etralona is rather les s

like the Neanderthals, while some later

Eur opeans, including the Sima people,

have a pattern of cresting on the nasal

fl oo r r e se m b li n g t h a t i n t h e B r ok en

Hill or Bodo fossils. Finally, it is wor th

More c rucial to the

a ccretion hypothesis, or

at lea st the version of it

that encompa sses the

ea rliest fossils, a re the

crania from Arago and

Petralona. Here the

question is whether there

a re signsof incipient

Neanderthalmorphology, especially

in the fa cial region.



7/10

noting that neither the Arago nor th e

P etralona cranium exhibits the apo-

m orphic traits identifi ed recently by

S chw artz and Tattersall.57 A m e d i a l

projection from the inner m argin of

the nos e and a s w elling of the pos te-

r i or n a s a l w a ll i t se lf a r e p r e se n t i n

N eanderthals but s eem to be lackingin earlier Middle Pleistocene hom inids.

Given this level of doubt concerning

s pecific N eanderthal affinities of the

Ar a go a n d P et r alo n a c ra n ia o r t h e

Mauer jaw, perhaps it is premature to

disassociate these specimen s from con -

temporary Africans. In my view, all of

t h e e ar li er M id d le P le is to ce ne

hom inids s hare both erectus-like fea-

t u r es a n d a s u it e o f d e r ive d t r ait s

com m on to later hum ans . I t is hard to

fi nd any m orphological bas is for re-

stricting Hom o heidelbergensis t o E u -

rope. This ta xon m ay well have evolvedelsewhere. H owever, th ese people did

reach Europe at an early date. S om e-

t i m e l a te r, a s c li m a t ic c o n d it io n s

changed and populations becam e iso-

lated by ice barriers , s peciation pro-

duced the fi rst N eanderthals (F ig. 2).

Just when this event occurr ed is uncer-

tain , but th e ca. 300,000-year-old Sim a

fos s ils, a s w ell a s thos e from S tein-

heim and S w ans com be, foreshadow

the N eanderthal condition. There are

good reas ons to pos tulate deep roots

for the N eanderthal l ineage, and here

the accretion m odel m us t be broadlyaccurate.

MORE FOSSILS FROM SPAIN

New evidence challenging both the

phylogenetic role of Hom o heidelber-

gensis a n d t h e a c cr et io n m o d el h a s

b e en r e p or t ed r e ce n t ly fr o m S p a in .

T h e S ie r r a d e At a p u e r ca c o n t ai n s

m any s ites in a ddition to the S im a de

los H ues os . O ne is a l im es tone cave

deposit exposed by workers cutting a

railw ay trench at the turn of the cen-

tury. The collapsed cave of Gran Do-

lina is filled with a substantial thick-

nes s of s edim ents . The m os t ancient

levels m us t be of E arly P leis tocene

a g e . P a l e o m a g n et i c s a m p l in g i n d i -

cates that the TD6 layers may lie just

b e lo w t h e B r u n h es -M a t u ya m a m a g -

netic reversal, dated a t 780,000 years. 58

Excavations in one of the TD6 s trata

have produced a collection of s tone

tools consisting of core-choppers and

fl akes , but no handaxes or cleavers .

Along with the artifacts, there are hu-

m a n b o n e s r e p r es en t in g a t l ea s t s i x

different individuals. Given the uncer-

t a in t ie s s u r r o u n d in g t h e a g e o f t h e

find at Ceprano in Italy, these Span ish

d i sc o ve r ie s m a y b e t h e o ld e st a n y -

where in Europe.

Al th o u g h m a n y o f t h e fo ss ils a r efragm entary, they provide inform a-

tion about the teeth, skull, and postcra-

n i a l s k el et o n . P r e li m i n a r y d e s cr i p -

tions have been pr ovided by Carbon ell

and colleagues51 a n d B er m u d ez d e

Castro and colleagues.59 O ne im por-

tant s pecim en is part of a low er jaw

w ith the m olars s til l in place. M ore

teeth belonging to this hom inid have

been recovered and it is possible that a

piece of frontal bone s hould als o be

as s igned to the s am e individual, an

adolescent about 14 years old. Differ-

ences from Hom o erectus are apparent

in the expans ion of the lower incis or

crow ns , the s ize relations hips of the

prem olar teeth, and the relative gracil-

i ty o f t h e m a n d i b u la r b o d y. F r o n ta l

breadth, w hich can be es tim ated as a

m inim um behind the brow s , exceeds

that for all but th e largest of the H o m o

erectus crania from Asia.

Confi rm ation that the G ran D olina

people are m ore advanced tha n H o m o

erectus com es from a partial face dis -

covered in 1995. The m orphology of

this s pecim en, repres enting another

young individual, is said to be r emar k-

ably modern . Below the orbit, hollow-

ing of the bone surface is accentuated

by forward bend ing of the side wall of

the nose. Su ch a degree of flexion of

t h e m a xilla i s n o t s ee n i n a r ch a ic

h o m i n i d s. T h a t h o ll ow, t h e c a n in e

fossa, is not well developed in speci-

men s such as th ose from Bodo, Arago,

or Petr alona. As noted earlier, Neand er-

tha l faces have quite a different appea r-

ance, w ith the cheek region infl ated

and swept to the r ear. If the identifica-tion of the Ceprano braincas e is cor-

roborated, the Gran Dolina people may

have coexisted with Hom o erectus in

the M editerranean region. H ow ever,

these people seem to be set apart, not

only from Hom o erectus, but a lso from

Hom o heidelbergensis and later Euro-

peans .

The evidence from m idfacial topog-

raphy, along with other cranial, man-

dibular, and d ental chara cters, has per-

s uaded the A tapuerca res earchers to

n a m e a n e w s p e ci es , Hom o anteces-

sor.59 As describ ed so far, the fossils donot exhibit any of the d erived features

of Neanderthals and so do not fit neatly

into the accretion model, which holds

all early Europeans to be Neanderthal

ancestors. Members of the new taxon

can b e interpreted as close relatives to

M iddle P leistocene hom inids s uch as

Mau er an d Arago. Moreover, some fea-

tures of i ts m an dibular body are very

s im ilar to thos e of the S im a popula-

tion. This suggests evolutionary con ti-

nuity with Hom o heidelbergensis, taken

to be a s tr ictly European lineage. A t

the s am e tim e, Hom o antecessor m ay

be generalized enough in its morphol-

o g y t o b e a n c e s t r a l t o m o r e m o d e r n

hum ans . The s pecies from S pain thus

s eem s to be a candidate for the s tem

fr o m w h ic h b o t h N ea n d e r th a l s a n d

Hom o sapiens are descended. Here the

r o le p l a ye d b y M i d d le P l ei st o c en e

p o p u la t i on s i n Afr i ca i s l ef t u n e x-

p la i n ed , b u t B o do a n d B r ok en H i ll

apparently are n either antecessor n or

heidelbergensis and m ust represent still

another (unnam ed) taxon.

Undoubtedly, the exciting new mate-

rial from Atapuerca will be subjected

to further com parative s tudy. In the

meantime, there are questions. Atten-

tion has been focused particularly on

the facial skeleton. The m orphology of

the ATD6-69 m idface does seem to b e

distinctive with respect to oth er Middle

Pleistocene hom inids, and a well devel-

oped canine fossa is characteristic of

m o r e m o d e r n h u m a n s . A c o m p li ca -

tion is that th is Gran Dolina ind ividual

is juvenile, maybe on ly 10 or 11 years

Confirma tion tha t the

Gra n Dolina people a re

more advanced thanHomo erectuscomes

from a pa rtia l fa ce

discove red in 1995. The

morphology of this

specimen, representing

a nother youngindividual, is sa id to be

rem a rka bly modern.



8/10

o ld . I t i s a l w a ys t r ic ky t o c o m p a r e

children to adults, for little is known

o f t h e g ro wt h p a tt er n s in a r ch a ic

people. Almost certa inly, th e sh ape of

t h e m a x il la c h a n ge s a s t h e s in u s es

expand and the teeth are fully erupted.

In another (adult) s pecim en from the

s it e , l e ss h o llo w in g o f t h e c h ee k i spresent. So one must ask whether this

feature is an appropriate com ponent

of the diagnos is of a new s pecies . I t

seems to me that one can m ake a good

case for attributing t he Spa nish fossils

to Hom o heidelbergensis, w h e re t h e

h y po d ig m f or t h is t a xo n i s d e fi n ed

broadly to include Europ ean an d Afri-

can r ema ins. In this view, Hom o heidel-

bergensis, like Hom o erectus, w a s a

wide-ranging species rather than just

a s h or t s eg m e n t o f a li n ea g e s a n d -

w iched in betw een the G ran D olina

hominids and later Neanderthals.

CONTINUING QUESTIONS

S everal ques tions about the evolu-

tionary role of Hom o heidelbergensis

h a ve s ti ll t o b e t ou c h ed o n . I n m y

reading of the record, this s pecies is

ancestral n ot only to Hom o neandertha-

lensis b u t a ls o t o m o d er n h u m a n s.

Figure 2 suggests speciation to H o m o

sapiens in Africa or western Asia some-

tim e in the later M iddle P leis tocene.

However, there are more fossils from

the F ar Eas t that com plicate this pic-

ture. P articularly in China, m any lo-

calities docum ent the p res ence of h u-

m a n s m o r e a d v an c ed t h a n Hom o

erectus, certainly after 300,000 years

ago and perh aps mu ch earlier. Wheth er

the s keletons s hould be lum ped w ith

Hom o heidelbergensis is one issue; how

they are related to recent Asian popu-

l at io n s i s a n o t h er. B o th a r e f ra u g h t

with con troversy.

Two importa nt cran ia have been dis-

covered in terra ce deposits of the Ha n

River at Yunxian in western Hub ei.60

T h e fi n d s w e r e m a d e i n a c la y l a ye r

a n d b o t h h o m i n i d s w e r e e n c a s e d i n

hard calcareous matrix. The same level

has produced m am m alian foss ils and

some stone cores and flakes. The faun a

suggests a Middle Pleistocene age. Pa-

leomagnetic work coupled with other

a p p ro a ch e s n o w i nd ic a te s t h a t t h e

Yunxian a ssemblage may be as m uch

a s 6 00 ,0 00 o r e ve n 8 00 ,0 00 y ea r s

old.61,62

Unfortun ately, the crania th emselves

a r e h e a v i l y d a m a g e d . O n e h a s b e e n

c r u s h e d n e a r l y fl a t . I n t h e o t h e r t h e

face is reasonably well preserved, al-

though the vault has been deform ed

and the base is filled with small cracks.

There are some resemblances to H o m o

erectus. The brow is thickened and t he

vault is long and low, with an angledocciput. The cranial base is generally

s im i la r t o t h a t i n t h e Z h o u k o u di a n

s p ec im e n s a n d d o e s n o t s e e m t o e x -

hibit the flexion apparent in later popu-

lations. At th e sam e tim e, the Yunxian

crania s hare m an y features w ith m ore

a d va n c ed h u m a n s . T h e b r a in c a se i s

large and not very constricted behind

the orbits , and the s quam ous tem po-

ral is arched. S om e traits of the nos e

and palate may also be derived relative

to Hom o erectus. The m idface of the

second individual is described as espe-

cially sapiens-like in that a canine

fos s a is pres ent a nd the infraorbital

region is s et at an angle to the fl aring

cheek. Given this mix of characters, Li

and Etler 60 and Etler 61 choose to place

t h e s ku l ls w it h Hom o erectus, bu t

Zhang63 identifies them with later hu-

m a n s .

O ther fos sils a re know n from later

Middle Pleistocene localities in China.

T h e D al i a n d J in n i u sh a n s p ec im e n s

are often des cribed as archaic or pre-

m odern Hom o sapiens.64 The Dali cra-

nium is quite com plete. I ts m as s ive

brow, keeled frontal, and low vault are

reminiscent ofHom o erectus. I n m a n y

other res pects , the D ali braincas e is

m ore like that of later hum ans . Even

w h e n c r u sh i n g o f t h e m a x il la i s a c -

counted for, the face must be relatively

short. Also, the margin of the nose is

vertically oriented and the incisive ca-

n a l i s p l a c e d a n t e r i o r l y o n t h e h a r dpalate, as in Middle Pleistocene Afri-

c a n s a n d E u r o p ea n s . As w it h o n e o f

the Yunxian faces,th e wallof the cheek is

hollowed to prod uce a can ine fossa.

If i t is accepted that thes e Chines e

individuals (including Yunxian?) are

n ot Hom o erectus, then s orting them

to Hom o heidelbergensis is one alterna-

t ive t h a t m u s t b e e xp lo r ed . T h i s i s

suggested in Fig. 2. Depending on the

age of the Yunxian m aterial, the en-

t r a n c e o f Hom o heidelbergensis into

eastern Asia might have occurred ear-

lier th an depicted. The taxon w ouldthen have pers is ted alongs ide H o m o

erectus for a substantial time. A ques-

t io n a r is es a s t o t h e fa t e o f t h es e

Chinese populations. The scenario of

Fig. 2 shows eventua l extinction, bu t a

c a se fo r c o n ti n u it y w i t h r e ce n t h u -

ma ns mu st be considered. In fact, this

point is still difficult to resolve from

the paleontological record. Compara-

tive m olecular s tudies keep open the

possibility that there wa s some contri-

but ion from arch aic Asians to the mod-

ern gene pool.65

Some workers elect instead to sepa-

rate the Chinese hominids from Middle

P leistocene populations in the Wes t.

This preference is ba s ed largely on

observations of the midface, which is

s ai d t o s h ow m o d er n fe a tu r es a t a

relatively early date. These w orkers

em phas ize the developm ent of a ca-

nine fos s a, along w ith lateral prom i-

nence of the cheek. If these differences

are taken to pr eclude an identification

as Hom o heidelbergensis, then the fos-

sils may have to be allocated to a new

taxon. H owever, as noted earlier, hol-

lowing of the infraorbital surface can

b e d o cu m e n t e d fo r f ac es o u t si d e o f

Ch i n a . F u r t h e r m o r e, t h e n e w fi n d s

fr o m G r a n D oli n a s u gg es t t h a t t h i s

fe a t u r e m a y a p p e a r i n E u r o p e a t t h e

beginning of the Middle Pleistocene.

S uch evidence w ill m ake it harder to

argue for is olation of the m ajor O ld

Wo r ld g eo g ra p h i c p r o vi n ce s . T h e

s pread of s om e populations of H o m o

heidelbergensis i n to t h e F a r E a s t c a n -

not be ruled out.

Pa rticula rly in China ,ma ny loca lities

document the presence

of humans more

advanced than Homo

erectus, certainly after

300,000 yea rs a go a nd

perha ps much ea rlier.

Whether the skeletons

should be lumped withHomo heidelbergensisis

one issue; how they a re

related to rece nt Asia n

populationsisanother.



9/10

CONCLUSIONS

In their w ell-know n 1975 volum e

After the Australopithecines,66 K a r l

B u t ze r a n d G lyn n I sa a c n o t ed m a n y

uncertainties surrounding human evo-

lution in the Middle Pleistocene. Al-

m os t a quarter of a century later, the

mu ddle in the mid dleis still eviden t,

es pecially in res pect to s ys tem atics

and classification of the hominids. Per-

haps the m os t vexing ques tions con-

cern fos s ils of earlier M iddle P leis-

t o ce n e a n t i qu i t y. S p e ci m e n s f ro m

Af ri ca a n d E u r a s ia h a ve m o s t fr e -

quently been described as archa ic

representatives of ou r ow n s pecies,

but this situation is unsatisfactory for

several reasons. Fossils such as those

found at Bodo, Broken H ill , Arago,

Petralona, and Dali retain m any primi-

tive erectus-like chara cters, a nd this

anatom y s ets them apart from recent

hum ans . S im ply lum ping divers e an-

cient groups w ith living popu lations

obscures these differences.

There is increasing acceptance of

the s ugges tion that dis tinct l ineages

may have evolved during this period.

O ne pos sibility is that fos s ils from

A frica and Europe can be s orted to-

gether to a single taxon, approp riately

called Hom o heidelbergensis. This spe-

cies may have originated in Africa. If

the Gra n Dolina fossils are also H o m o

heidelbergensis, then thes e people ap-

p a r e n tly r e a c h e d E u r o p e a t a n e a r ly

date. In this region, populations is o-

l at e d b y g la c ia l c o n d it i on s p e r h a ps

were eventua lly ancestral to the Nean -

derthals. In other parts of the species

range, including Africa, there are indica-

tions that later Middle Pleistocene groups

were evolving in the direction of Homo

sapiens. Hom o heidelbergensis is thus the

s tem from w hich both N eanderthals

and m odern hum ans are derived.

Aproblem is whether the sam e taxon

can be identifi ed in the Eas t. F os s ils

such as those found at Dali and Jinni-

u s h a n i n Ch i n a a r e m o r e a d v a n ce d

than Hom o erectus and exhibit s om e

of the s am e derived characters as do

t h e s p ec im e n s fr o m Afr i ca a n d E u -

r o p e . B u t i t c a n b e a r g u e d t h a t t h e

Chines e h om inids are distinctive in

as pects of their facial m orphology.

S o m e w o r ke r s w il l p r e fe r e it h e r t o

place them in a new s pecies or lum p

them as early Homo sapiens. This ques-

tion rem ains to be resolved.

ACKNOWLEDGMENTS

My studies of Pleistocene hominids

have been conducted w ith the as s is -

tance of ma ny individuals and institu-

tions, and I am grateful for this help.

The governm ents of China, Ethiopia,

I n d on e si a, K en ya , a n d Ta n z an i a

granted me clearance to examine fos-

s il s i n t h e se c o u n t r ie s. T h e L .S .B .

Leakey Foundation and the Boise Fund

s u p p or t e d m u c h o f t h e r e se a r ch o n

which th is paper is based. R.G. Klein,

R . Q u a m , a n d C.B . S t r i n ge r k in d l y

com m ented on a version of the m anu-

s cript, as did s everal anonym ous re-

viewers.

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