You are on page 1of 10

218 Evolutionary Anthropology


Human Evolution in the Middle Pleistocene:

The Role of Homo heidelbergensis

For paleoanthropologists working in show anatomical features that distin- species spans an interval of at least 1.5
the Middle Pleistocene, these are inter- guish them from Homo erectus. In the million years. Indeed, some East Asian
esting times. New discoveries of arti- Far East, the people at Dali and other populations may have survived into
facts and human fossils have been sites are also more advanced than later Pleistocene times.
reported from western Europe, so that Homo erectus, but their affinities to As documented by Weidenreich, von
it now looks as though this continent groups in the West are uncertain. Koenigswald, Le Gros Clark, and oth-
was populated 800,000 years ago, if This Middle Pleistocene record, still ers, members of this taxon share a
not earlier. One of the fossils, from sparse but increasingly well dated, suite of characters by which they can
Ceprano in Italy, is described as Homo raises important questions. One con- be distinguished from recent humans.
erectus. Whether this ancient species cerns the fate of Homo erectus in differ- Some of the principal differences re-
ever reached Europe has been repeat- ent regions of the Old World. Another late to cranial capacity, keeling on the
edly questioned, but the Ceprano cra- is how many distinct species should be midline of the vault, parietal length,
nium is complete enough to provide recognized among the descendants of occipital proportions, the anatomy of
some hard evidence. this ancient lineage. It is apparent that the cranial base, facial projection, the
Other finds from Spain are even the traditional approach of lumping form of the mandibular symphysis,
diverse humans together as ‘‘archaic’’ tooth size, the relative narrowness of
more spectacular. The Sima de los
Homo sapiens will no longer work. the pelvis, and the length of the femo-
Huesos (‘‘Pit of Bones’’) in the Sierra
The picture is highly complex, and ral neck. A large number of traits
de Atapuerca has yielded a wealth of
several taxa probably are needed to generally describe Homo erectus and
skeletons that are best interpreted as
accommodate the fossils. Evolution- diagnose this species relative to living
early Neanderthals, perhaps close to
ary relationships among these popula- people.
300,000 years in age. Older but unfor-
tions must be clarified, but pose some These assumptions have been chal-
tunately more fragmentary remains,
major problems. I will address only a lenged by several workers, on highly
also from Atapuerca, display no Nean-
subset of these topics pertaining diverse grounds. One point of conten-
derthal features and are claimed as
mainly to earlier Middle Pleistocene tion concerns the material from the
representatives of a new species. Homo
hominids. Turkana Basin. It has been claimed
antecessor will require close study.
that the early Kenyan crania lack spe-
These European discoveries focus
cial features developed by the Asian
fresh attention on the evidence accu- HOMO ERECTUS IN PERSPECTIVE populations. A midline keel on the
mulating from Africa and Asia. Hu-
This extinct species has been at the vault, an angular torus at the postero-
man bones are known from the earlier
center of much controversy. At pre- inferior corner of the parietal bone,
Middle Pleistocene of Africa at locali-
sent, there is no firm consensus as to certain characters of the base, and
ties such as Bodo in Ethiopia and
whether it should be defined as a long overall thickening of the braincase are
Broken Hill in Zambia. The crania
lasting, polytypic lineage or as a group said to be absent from the specimens
of relatively specialized populations at Koobi Fora but well expressed in
geographically confined to the Far the remains from Trinil, Sangiran, and
G. Philip Rightmire is Professor of Anthro- East. In my view, Homo erectus origi- Zhoukoudian. These differences have
pology at the State University of New York nated in Africa and then spread to prompted investigators, including An-
at Binghamton. In his research, which fo-
cuses on the evolution of the genus Homo, Eurasia. The hypodigm is made up of drews,1 Groves,2 and Larick and Cio-
he has studied fossils from many of the specimens from Java, Zhoukoudian, chon,3 to recognize two species and to
important prehistoric localities in Europe, and other sites in China, Ternifine suggest that Homo erectus must be
Africa, and the Far East. He is particularly
interested in questions about Homo erec- (now Tighenif) in Northwest Africa, geographically restricted to the Far
tus, Middle Pleistocene hominids, and the Olduvai Gorge, the Turkana Basin, East. Wood4,5 agrees, on the basis of
origins of more modern humans.
and Swartkrans in South Africa (Fig. facial measurements, perhaps some
1). Several fossils recently discovered aspects of temporal bone morphology,
Key words: species; phylogeny; Homo erectus; in western Asia and in Europe prob- and dental differences, that the early
Homo heidelbergensis; Neanderthals ably should be counted as well. This African hominids should be set apart
ARTICLES Evolutionary Anthropology 219

to the postcranial skeleton as well. All

of the better-preserved individuals, in-
cluding even the late surviving ones
from some of the Far Eastern sites,
can be set apart from Homo sapiens.
The boundary between these taxa is
not arbitrarily defined.


Although there is disagreement
about taxonomy, most workers would
concede that populations resembling
Homo erectus dispersed from Africa
into Eurasia well before 1.0 million
years ago. These movements occurred
probably over a long period. The
hominids may have made repeated

Homo erectus, as
broadly defined, does
possess many
anatomical distinctions,
Figure 1. A map showing the principal localities that have yielded fossil remains of Homo extending not only to the
erectus. Ceprano in Italy is the first site to demonstrate that the species reached Europe,
probably before the onset of the Middle Pleistocene. skull and teeth but to the
postcranial skeleton as
from later Homo erectus. Wood now among the several assemblages of well. All of the better-
refers the Turkana Basin specimens to Homo erectus, but the fossils from the preserved individuals,
Homo ergaster, which, in his opinion, Turkana Basin, Olduvai Gorge, and
is more likely than Homo erectus to other sites in Northwest Africa exhibit including even the late
have played a role in the evolution of essentially the same set of traits as do surviving ones from some
later people. those from the Far East.9 Discrete
A quite different interpretation has characters said to be unique to the
of the Far Eastern sites,
been offered by Wolpoff and cowork- Asian populations are variable in their can be set apart from
ers,6 who now claim that the nomen expression and, in fact, most can be
Homo erectus is unnecessary and identified in the earlier East African
Homo sapiens.
should be discarded altogether. Here material.10 Cranial dimensions show
the question is really whether there is much overlap.11,12 Vault thickness, as
continuity from earliest Pleistocene measured near the junction of the sorties, introducing crude chopping
times right to the present; that is, frontal and parietal bones, is about the tools and stone flakes or Acheulean
whether just one long lineage, with no same in the African and Asian
handaxes into different regions.16
branches or extinctions, can be recog- samples.13 The faces of KNM-ER 3733
Evidence documenting the spread
nized. Such a lineage would include from Koobi Fora and KNM-WT 15000
of humans into western Asia comes
the ancient Turkana populations as from Nariokotome conform in nearly
from several sites, including Dmanisi
well as those later resident in Africa all respects to the anatomy of Homo
and Eurasia. If it could be demon- erectus as reconstructed from the San- in Georgia and ‘Ubeidiya in Israel. At
strated that no splitting had occurred, giran and Zhoukoudian specimens.14 Dmanisi, a lithic industry and a well-
one might argue that any division be- Also, the teeth from the Turkana Basin preserved mandible with teeth have
tween taxa would have to be arbitrary. are close in size and shape to those been recovered.17 The jaw resembles
Wolpoff et al.6 and Wolpoff7 (see also from Zhoukoudian.15 Apparently there those of Homo erectus.18 The bone-
Tobias8) go a step further and say that are not many traits that can be used to bearing levels overlie a lava flow dated
there simply is no basis for keeping diagnose Homo ergaster, and probably at 1.8 million years, but the age of the
more than one species, which must just one polytypic species should be tools and fossils remains problemati-
then be Homo sapiens. recognized. Nevertheless, Homo erec- cal.16 At ‘Ubeidiya, an extensive mam-
It does not seem to me that either of tus, as broadly defined, does possess malian fauna excavated from lake sedi-
these scenarios can be supported fully. many anatomical distinctions, extend- ments suggests a relatively cool climate
Certainly there is geographic variation ing not only to the skull and teeth but at a date of perhaps 1.4 to 1.0 million
220 Evolutionary Anthropology ARTICLES

years. In addition to the fauna, there Old World. These hominids flourished This budding of a daughter lineage
are stone choppers, spheroids, picks, for a long time. At sites including from Homo erectus must have oc-
and bifaces, which Tchernov19 com- Zhoukoudian and Longtandong curred very early in the Middle Pleis-
pared to the lithic material from upper (Hexian) in China, the species is known tocene, if not before. African and west-
Bed II at Olduvai Gorge. How the from deposits of the later Middle Pleis- ern Asia are likely areas in which the
differences between the non-Acheu- tocene, while at Ngandong in Java, at first more advanced humans origi-
lean and Acheulean assemblages at least one group of archaic people may nated (Fig. 2). An Africa locus is consis-
this Jordan Valley site should be inter- have survived into the Late Pleis- tent with findings from archeology,
preted is unclear, but the tools most tocene.24 Populations such as that at environmental reconstruction, and
probably were used by groups of Homo Ngandong may document the last ap- patterns of animal dispersal.28
erectus. pearance of the lineage. Fossils that shed light on this specia-
Ancient traces of hominid activity tion event have turned up at several
are found also in Europe, at French SPECIATION IN AFRICA? localities in Africa. One is Bodo in the
localities such as Le Vallonet, Isernia Middle Awash region of Ethiopia. The
in Italy, and in the Neuwied Basin in The picture emerging is one of Homo Bodo cranium and, later, a broken
Germany. This scattered archeological erectus as a widespread, polytypic spe- parietal from a second individual, were
cies, with groups persisting longer in
evidence, consisting mainly of core- found in conglomerates and sands con-
some regions than in others. The pat-
choppers and flakes, sometimes found taining mammalian bones and Acheu-
tern documented in China and espe-
with broken animal bones taken to be lean artifacts.29,30 Fauna from the site
cially in Java contrasts with that in the
food waste, may demonstrate a hu- has been compared to that from Bed
man presence in the Early Pleis- IV at Olduvai Gorge and Olorgesailie
tocene20 (but also see Roebroeks21). in Kenya, and an early Middle Pleis-
Unfortunately, the oldest European tocene age is indicated. New argon-
sites have not produced more than a The pattern documented argon dates reported by Clark and
few bits of human skeleton. A notable in China and especially colleagues31 support this biochronol-
exception is Ceprano in central Italy. ogy. The evidence from fauna, archeol-
Here a fragmentary but fairly com- in Java contrasts with ogy, and laser-fusion determinations
plete braincase was discovered in 1994. that in the West, where points to an age of about 600,000
The fossil was picked up in clay years for the Bodo hominids.
deposits, which contain no volcanic
Homo erectus seems to The face and anterior portion of the
material that is directly datable. Potas- disappear from the braincase are reasonably complete; it
sium-argon dates have been obtained record at a relatively can be established that Bodo is like
from volcanic sands higher in the Homo erectus in some features. The
stratigraphic sequence; these are said early date. Also, it is massive facial bones, projecting brow,
by Ascenzi and coworkers22 to indi- interesting that the Asian low and constricted frontal with mid-
cate an age greater than 700,000 years. line keeling, parietal angular torus,
Insofar as can be determined from the populations apparently and thick vault give the specimen an
description provided by its discover- are more specialized. archaic appearance. In other respects,
ers, this hominid displays the heavy the cranium is more advanced in its
continuous brow, low vault, angled morphology. Brain size is close to 1,300
occiput, and thick cranial bones that cc, which is substantially greater than
are characteristic of Homo erectus. West, where Homo erectus seems to is expected for Homo erectus. The fron-
This is important information. Al- disappear from the record at a rela- tal bone proportions, arched shape of
though the Ceprano specimen is dam- tively early date.25,26 Also, it is interest- the squamous temporal, and some
aged in some key respects, it seems to ing that the Asian populations appar- traits of the cranial base are like those
confirm the identity of one group of ently are more specialized in the sense of more modern humans. Although
people who entered Europe in the of exhibiting a higher incidence of the face is very broad and heavily
Early Pleistocene. some morphological characters associ- constructed, the supraorbital tori are
Representatives of Homo erectus also ated with cranial robusticity. These divided into medial and lateral seg-
reached the East Asian tropics before traits are subject to geographic varia- ments, the margin of the nose is verti-
moving into more temperate regions. tion and do not mark a species bound- cal rather than forward sloping, and
It has been assumed that movement ary, but they may nevertheless delimit the incisive canal opening into the
into the Far East began 1.0 million groups that had different evolutionary front of the hard palate shows a de-
years ago or slightly earlier, but radio- fates. rived condition present in recent
metric dates from mineral samples There is no reason to suppose that Homo.32
collected at Modjokerto and Sangiran all demes of Homo erectus evolved This mix of characters suggests that
now suggest that the oldest Indone- further. The evidence is consistent with the Middle Awash individuals are ‘‘in-
sian localities may be 1.8 to 1.6 million eventual extinction of some or all termediate’’ in their morphology. How-
years old.23 If this result can be veri- populations in the Far East. A less ever, several of the resemblances to
fied, then it will look as though Homo specialized branch of the species may Homo erectus are plesiomorphies that
erectus spread quite rapidly across the well have given rise to later humans.9,27 cannot be considered diagnostic.
ARTICLES Evolutionary Anthropology 221

crania differ only slightly in orbit size,

frontal proportions, and prominence
of the torus crossing the occipital bone;
in general, they are remarkably alike.
Resemblances are apparent in the
height, breadth, and massive construc-
tion of the upper face and cheek, sev-
eral measures of projection in the fa-
cial midline, configuration of the
thickened brows, and many aspects of
vault shape.9 Because the Bodo mate-
rial is less complete, comparisons be-
tween it and the Greek cranium must
be more limited in scope, but here also
there are similarities.32
Multivariate studies of skull form
have been carried out by several work-
ers, who noted resemblances between
the African and European speci-
mens.37–39 Van Vark40 has used 17 di-
mensions of the face and braincase to
construct a measure of generalized
distance (D2), which shows that the
Broken Hill and Petralona specimens
differ from one another less than is the
case for Upper Paleolithic and recent
humans. When a reconstruction of the
partial cranium from Arago Cave in
France is included in this analysis, it
also falls close to that from Broken
Hill. There seem to be good phenetic
grounds for lumping these hominids
Figure 2. A tree illustrating the evolution and geographic distribution of Homo in the Pleis- Other ancient European finds are
tocene. Homo erectus is assumed to have originated in Africa and then spread quickly to Asia
more fragmentary. Human bones and
and probably to Europe. Homo heidelbergensis is distributed from Africa into Eurasia during the
Middle Pleistocene. Whether this species reached the Far East is still a question. European teeth have been recovered from the
Homo heidelbergensis gave rise to the Neanderthals, while an African branch of Homo quarry at Bilzingsleben in Germany
heidelbergensis is ancestral to modern humans. Four speciation events (S) are depicted. and from several earlier Middle Pleis-
tocene sites in Italy, while an occipital
bone is on record from Vertesszöllös
Moreover, it is clear that the cranium mal remains possibly associated with in Hungary. There is also the rear
shares other apomorphic features with the Broken Hill cranium suggest an portion of a braincase from Swans-
more modern populations. It seems age within this same broad interval.34 combe in England. The mandible from
reasonable to group Bodo with the As has been recognized for some Mauer in Germany and a tibia associ-
famous fossil from Broken Hill (Ka- time, the African hominids are similar ated with Acheulean bifaces at Box-
bwe) in Zambia (Fig. 3), along with to other, roughly contemporary people grove in England are arguably among
specimens from Elandsfontein in known from Europe. Like that from the oldest hominids from Europe. Both
South Africa, Lake Ndutu in Tanzania, Broken Hill, the cranium from Pe- are on the order of 500,000 years in
and probably Eyasi, also in Tanzania. tralona in Greece is particularly well age.41 Of course there are questions
These localities are Middle Pleistocene preserved (Fig. 4). Although there is about the affinities of this material, as
in age. In addition to the human skull- doubt about both its original prove- few anatomical clues are preserved.
cap, deflation surfaces (‘‘bays’’) at nience within layers of stalagmite de- But if the Mauer mandible is grouped
Elandsfontein have yielded a large posited on the cave floor and its asso- with the other specimens from Europe
fauna, together with Acheulean ciation with animal bones, this and Africa, then the entire assemblage
handaxes. Dating of this assemblage is individual is of Middle Pleistocene an- can be referred to Homo heidelbergen-
complicated by the fact that several of tiquity. Some of the flowstone may be sis.9,42,43
the extinct mammal species are un- about 200,000 years old,35 but other This species was named by
known elsewhere, but comparisons fossils from the site imply a greater Schoetensack in 1908 to accommo-
with other African sites imply that the age for the cave contents (see Cook date the jaw found a year earlier in the
bones were accumulated between and coworkers36 for a review). In any basal sand and gravel complex of the
700,000 and 400,000 years ago.33 Ani- case, the Petralona and Broken Hill Grafenrain pit near Heidelberg.
222 Evolutionary Anthropology ARTICLES

Figure 3. Facial and lateral views

of the Broken Hill cranium. 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 exhibits features of the na-
sal region, palate, occiput and
base that distinguish it from Homo

Schoetensack was impressed with the cifically distinct from archaic lineages questions about the make-up of Homo
primitive character of his fossil but in Asia or Northwest Africa. Howell heidelbergensis. Some authorities hold
recognized that it must be human, as left open the relationship of this iso- that the European and African speci-
the canines are reduced in size and the lated fossil to other European groups, mens should be set apart as represen-
tooth crowns generally display the pro- including the Neanderthals. While tatives of distinct lineages. Proponents
portions expected for modern popula- there are still obvious difficulties with of this view agree that the French,
tions. linking the mandible to individuals German, and Greek fossils share a
Later authors continued to empha- such as that from Petralona, for which series of features with the hominids
size the primitive appearance of the no lower jaw has been recovered, the from Bodo and Broken Hill. But they
mandible. Howell44 pointed to its mas- fossil can be lumped with earlier claim that even the earliest Middle
sive construction, multiple mental fo- Middle Pleistocene humans in the way Pleistocene Europeans exhibit apomor-
ramina, and very thick symphysis, I have outlined. As defined on this phic traits that align them only with
which lacks any indication of a chin, basis, Homo heidelbergensis retains a Neanderthals.45–49
as characters shared by other early number of archaic characters and may In this reading of the evidence, the
representatives of Homo. However, be the stem from which both Neander- fossils from Mauer and probably Box-
Howell was careful to note that other thals and modern people are derived. grove, Arago Cave, Petralona, Bilz-
features of the specimen, including its ingsleben, Vertesszöllös, and Swans-
ramus breadth, relatively great ante- combe document a single line evolving
rior depth of the corpus, and moderate toward the populations at Steinheim
size of the dentition, distinguish it HEIDELBERGENSIS in Germany and Atapuerca in Spain.
from both Far Eastern Homo erectus Just as there is controversy about These assemblages can be referred to
and the Ternifine people. He argued whether Homo erectus should be parti- Homo heidelbergensis. However, it is
that the Mauer hominid must be spe- tioned into two taxa, so there are recognized that there is no clear sepa-

Figure 4. Lateral and facial

views of the Petralona cra-
nium. This individual from
Greece resembles that from
Broken Hill in many metric
features relating to facial
proportions and vault shape.
Whether the Petralona face
also displays characters
unique to the Neanderthal
lineage is currently debated.
ARTICLES Evolutionary Anthropology 223

ration of the latter from early Neander- but the topography of the midface tunately damaged as a result of its
thals such as those from Biache in seems to anticipate that of later popu- long interment in compacted cave sedi-
France, Ehringsdorf in Germany, or lations. The infraorbital surface and ments. The frontal bone, interorbital
Saccopastore in Italy. Eventually this the side wall of the nose meet at a pillar, nose, and cheeks show numer-
same lineage produced the ‘‘classic’’ shallow angle, producing a slight con- ous cracks; localized areas of crushing
Neanderthals of the Late Pleistocene. cavity. The cheek region is thus not are also present. The discoverers have
So an alternative classification is pos- ‘‘inflated’’ in the extreme manner of been able to correct some of this dam-
sible in which not only the ‘‘classic’’ Neanderthals, but can be interpreted age in a reconstruction,54,55 but signifi-
populations, but also all the older fos- as intermediate in form. Also in the cant distortion remains. In spite of
sils, are placed in one species termed Sima sample, brows are very thick. these problems, some workers can dis-
Homo neanderthalensis50 (see also Car- Continuity of the supraorbital tori at cern definite resemblances to Neander-
bonell and coworkers51 and Arsuaga glabella is said to be reminiscent of thals. Hublin49 notes that the infraor-
and coworkers52). Neanderthals. At the rear of the cra- bital surface of the maxilla is flattened
According to this ‘‘accretion’’ hypoth- nium, the suprainiac area is large but and the cheek bones are obliquely
esis, distinctive Neanderthal charac- not very depressed. This trait and the oriented. Arsuaga and colleagues52 sug-
ters appear first in the facial skeleton. shape of the occipital torus seem to
gest that the Arago midface is actually
Advocates of the model argue that foreshadow the Neanderthal condi-
more Neanderthal-like than that of
such traces can be identified in the tion.
Sima cranium 5 in the extent to which
Mauer and Arago remains. At later Earlier in the Middle Pleistocene,
the infraorbital plate and the side wall
evolutionary stages, apomorphies ac- of the nose are continuous, and this
cumulate in the occiput and finally in surface is inflated or convex. Also,
the temporal region. It is suggested there is much forward protrusion of
that the ancestors of Neanderthals be- More crucial to the the face at subspinale (in the midline,
came increasingly isolated through
time as a consequence of colder cli-
accretion hypothesis, or just below the nasal opening). The
nose itself is limited inferiorly by a
mate conditions. In the second half of at least the version of it sharp rim, as in Neanderthals.
the Middle Pleistocene, barriers that encompasses the These observations must be tem-
formed by glaciers and associated tun-
earliest fossils, are the pered by the fact that cracking and
dra to the north, ice sheets in the
plastic deformation make it difficult to
mountains to the east, and the Mediter- crania from Arago and assess some key aspects of morphol-
ranean to the south reduced contact
and gene exchange with people out- Petralona. Here the ogy. The wall of the Arago maxilla is
generally flattened or even inflated in
side Europe.49 Isolation in this rela- question is whether there the manner characteristic of Neander-
tively harsh environment led to the
full expression of the morphology that
are signs of ‘‘incipient’’ thals, but there is slight hollowing
distinguishes Neanderthal skulls and Neanderthal laterally, below the orbit. This cannot
postcranial bones from those of other be discounted as due entirely to dam-
morphology, especially age. Also, it is not clear that the zygo-
Much of this scenario seems sound. in the facial region. matic bone is swept back (obliquely
Certainly it is easy enough to track the oriented) as noticeably as it is in later
Neanderthals back in time to Stein- populations. In facial forwardness at
heim or even to Swanscombe. The subspinale, as measured by the zygo-
occipital bones of both specimens dis- Neanderthal roots are more difficult to maxillary angle of Howells,56 the Arago
play signs of a suprainiac fossa (a find. Some authors have pointed to cranium, at 1137, is in the Neanderthal
centrally placed elliptical depression Vertesszöllös or even Bilzingsleben as range, and the Petralona specimen, at
with a pitted floor). Moreover, Swans- documenting evolutionary continuity, 1187, shows almost as much protru-
combe possesses a transverse torus but most of this material is too frag- sion. But the value for Broken Hill is
that is weak near the midline but mentary to provide convincing infor- only 1167. Consequently, a low zygo-
bilaterally projecting. These traits are mation. The jaw from Mauer is com- maxillary angle does not necessarily
diagnostic for the lineage. The Sima plete but, in fact, shows few if any align Arago and Petralona with Nean-
de los Huesos at Atapuerca confirms traits that can be taken as specific dertals rather than with other Middle
that Neanderthal features are present links to later European populations. Pleistocene specimens. The sharp infe-
in an assemblage that may be close to More crucial to the accretion hypoth- rior margin of the Arago nose is in-
300,000 years in age.53 esis, or at least the version of it that deed reminiscent of that in Neander-
As described by Arsuaga and his encompasses the earliest fossils, are thals. However, there is variation in
colleagues,47,52 the Sima skulls present the crania from Arago and Petralona. this feature. Petralona is rather less
a combination of plesiomorphic and Here the question is whether there are like the Neanderthals, while some later
derived characters. The well-preserved signs of ‘‘incipient’’ Neanderthal mor- Europeans, including the Sima people,
face of cranium 5 is quite large in phology, especially in the facial region. have a pattern of cresting on the nasal
relation to the braincase. This, by it- The face of the partial cranium from floor resembling that in the Broken
self, is not a Neanderthal apomorphy, Arago is largely complete but unfor- Hill or Bodo fossils. Finally, it is worth
224 Evolutionary Anthropology ARTICLES

noting that neither the Arago nor the Along with the artifacts, there are hu- the maxilla is not seen in archaic
Petralona cranium exhibits the apo- man bones representing at least six hominids. That hollow, the canine
morphic traits identified recently by different individuals. Given the uncer- fossa, is not well developed in speci-
Schwartz and Tattersall.57 A medial tainties surrounding the age of the mens such as those from Bodo, Arago,
projection from the inner margin of find at Ceprano in Italy, these Spanish or Petralona. As noted earlier, Neander-
the nose and a swelling of the poste- discoveries may be the oldest any- thal faces have quite a different appear-
rior nasal wall itself are present in where in Europe. ance, with the cheek region inflated
Neanderthals but seem to be lacking Although many of the fossils are and swept to the rear. If the identifica-
in earlier Middle Pleistocene hominids. fragmentary, they provide informa- tion of the Ceprano braincase is cor-
Given this level of doubt concerning tion about the teeth, skull, and postcra- roborated, the Gran Dolina people may
specific Neanderthal affinities of the nial skeleton. Preliminary descrip- have coexisted with Homo erectus in
Arago and Petralona crania or the tions have been provided by Carbonell the Mediterranean region. However,
Mauer jaw, perhaps it is premature to and colleagues51 and Bermudez de these people seem to be set apart, not
disassociate these specimens from con- Castro and colleagues.59 One impor- only from Homo erectus, but also from
temporary Africans. In my view, all of tant specimen is part of a lower jaw Homo heidelbergensis and later Euro-
the earlier Middle Pleistocene with the molars still in place. More peans.
hominids share both erectus-like fea- teeth belonging to this hominid have The evidence from midfacial topog-
tures and a suite of derived traits been recovered and it is possible that a raphy, along with other cranial, man-
common to later humans. It is hard to piece of frontal bone should also be dibular, and dental characters, has per-
find any morphological basis for re- assigned to the same individual, an suaded the Atapuerca researchers to
stricting Homo heidelbergensis to Eu- adolescent about 14 years old. Differ- name a new species, Homo anteces-
rope. This taxon may well have evolved sor.59 As described so far, the fossils do
elsewhere. However, these people did not exhibit any of the derived features
reach Europe at an early date. Some- of Neanderthals and so do not fit neatly
time later, as climatic conditions Confirmation that the into the accretion model, which holds
changed and populations became iso- Gran Dolina people are all early Europeans to be Neanderthal
lated by ice barriers, speciation pro- ancestors. Members of the new taxon
duced the first Neanderthals (Fig. 2).
more advanced than
can be interpreted as close relatives to
Just when this event occurred is uncer- Homo erectus comes Middle Pleistocene hominids such as
tain, but the ca. 300,000-year-old Sima
fossils, as well as those from Stein-
from a partial face Mauer and Arago. Moreover, some fea-
tures of its mandibular body are very
heim and Swanscombe, foreshadow discovered in 1995. The similar to those of the Sima popula-
the Neanderthal condition. There are morphology of this tion. This suggests evolutionary conti-
good reasons to postulate deep roots nuity with Homo heidelbergensis, taken
for the Neanderthal lineage, and here specimen, representing to be a strictly European lineage. At
the accretion model must be broadly another young the same time, Homo antecessor may
accurate. be generalized enough in its morphol-
individual, is said to be
ogy to be ancestral to more modern
remarkably modern. humans. The species from Spain thus
MORE FOSSILS FROM SPAIN seems to be a candidate for the stem
New evidence challenging both the from which both Neanderthals and
phylogenetic role of Homo heidelber- Homo sapiens are descended. Here the
gensis and the accretion model has ences from Homo erectus are apparent role played by Middle Pleistocene
been reported recently from Spain. in the expansion of the lower incisor populations in Africa is left unex-
The Sierra de Atapuerca contains crowns, the size relationships of the plained, but Bodo and Broken Hill
many sites in addition to the Sima de premolar teeth, and the relative gracil- apparently are neither antecessor nor
los Huesos. One is a limestone cave ity of the mandibular body. Frontal heidelbergensis and must represent still
deposit exposed by workers cutting a breadth, which can be estimated as a another (unnamed) taxon.
railway trench at the turn of the cen- minimum behind the brows, exceeds Undoubtedly, the exciting new mate-
tury. The collapsed cave of Gran Do- that for all but the largest of the Homo rial from Atapuerca will be subjected
lina is filled with a substantial thick- erectus crania from Asia. to further comparative study. In the
ness of sediments. The most ancient Confirmation that the Gran Dolina meantime, there are questions. Atten-
levels must be of Early Pleistocene people are more advanced than Homo tion has been focused particularly on
age. Paleomagnetic sampling indi- erectus comes from a partial face dis- the facial skeleton. The morphology of
cates that the TD6 layers may lie just covered in 1995. The morphology of the ATD6-69 midface does seem to be
below the Brunhes-Matuyama mag- this specimen, representing another distinctive with respect to other Middle
netic reversal, dated at 780,000 years.58 young individual, is said to be remark- Pleistocene hominids, and a well devel-
Excavations in one of the TD6 strata ably modern. Below the orbit, hollow- oped canine fossa is characteristic of
have produced a collection of stone ing of the bone surface is accentuated more modern humans. A complica-
tools consisting of core-choppers and by forward bending of the side wall of tion is that this Gran Dolina individual
flakes, but no handaxes or cleavers. the nose. Such a degree of ‘‘flexion’’ of is juvenile, maybe only 10 or 11 years
ARTICLES Evolutionary Anthropology 225

old. It is always tricky to compare are heavily damaged. One has been reminiscent of Homo erectus. In many
children to adults, for little is known crushed nearly flat. In the other the other respects, the Dali braincase is
of the growth patterns in archaic face is reasonably well preserved, al- more like that of later humans. Even
people. Almost certainly, the shape of though the vault has been deformed when crushing of the maxilla is ac-
the maxilla changes as the sinuses and the base is filled with small cracks. counted for, the face must be relatively
expand and the teeth are fully erupted. There are some resemblances to Homo short. Also, the margin of the nose is
In another (adult) specimen from the erectus. The brow is thickened and the vertically oriented and the incisive ca-
site, less hollowing of the cheek is vault is long and low, with an angled nal is placed anteriorly on the hard
present. So one must ask whether this occiput. The cranial base is generally palate, as in Middle Pleistocene Afri-
feature is an appropriate component similar to that in the Zhoukoudian cans and Europeans. As with one of
of the diagnosis of a new species. It specimens and does not seem to ex- the Yunxian faces, the wall of the cheek is
seems to me that one can make a good hibit the flexion apparent in later popu- hollowed to produce a canine fossa.
case for attributing the Spanish fossils lations. At the same time, the Yunxian If it is accepted that these Chinese
to Homo heidelbergensis, where the crania share many features with more individuals (including Yunxian?) are
hypodigm for this taxon is defined advanced humans. The braincase is not Homo erectus, then sorting them
broadly to include European and Afri- large and not very constricted behind to Homo heidelbergensis is one alterna-
can remains. In this view, Homo heidel- the orbits, and the squamous tempo- tive that must be explored. This is
bergensis, like Homo erectus, was a ral is arched. Some traits of the nose suggested in Fig. 2. Depending on the
wide-ranging species rather than just and palate may also be derived relative age of the Yunxian material, the en-
a short segment of a lineage sand- trance of Homo heidelbergensis into
wiched in between the Gran Dolina eastern Asia might have occurred ear-
hominids and later Neanderthals. lier than depicted. The taxon would
Particularly in China,
then have persisted alongside Homo
many localities erectus for a substantial time. A ques-
CONTINUING QUESTIONS document the presence tion arises as to the fate of these
Several questions about the evolu- Chinese populations. The scenario of
of humans more Fig. 2 shows eventual extinction, but a
tionary role of Homo heidelbergensis
have still to be touched on. In my advanced than Homo case for continuity with recent hu-
reading of the record, this species is mans must be considered. In fact, this
erectus, certainly after point is still difficult to resolve from
ancestral not only to Homo neandertha-
lensis but also to modern humans. 300,000 years ago and the paleontological record. Compara-
Figure 2 suggests speciation to Homo perhaps much earlier. tive molecular studies keep open the
sapiens in Africa or western Asia some- possibility that there was some contri-
time in the later Middle Pleistocene. Whether the skeletons bution from archaic Asians to the mod-
However, there are more fossils from should be lumped with ern gene pool.65
the Far East that complicate this pic- Some workers elect instead to sepa-
ture. Particularly in China, many lo-
Homo heidelbergensis is rate the Chinese hominids from Middle
calities document the presence of hu- one issue; how they are Pleistocene populations in the West.
mans more advanced than Homo This preference is based largely on
erectus, certainly after 300,000 years
related to recent Asian observations of the midface, which is
ago and perhaps much earlier. Whether populations is another. said to show modern features at a
the skeletons should be lumped with relatively early date. These workers
Homo heidelbergensis is one issue; how emphasize the development of a ca-
they are related to recent Asian popu- nine fossa, along with lateral promi-
lations is another. Both are fraught to Homo erectus. The midface of the nence of the cheek. If these differences
with controversy. second individual is described as espe- are taken to preclude an identification
Two important crania have been dis- cially ‘‘sapiens-like’’ in that a canine as Homo heidelbergensis, then the fos-
covered in terrace deposits of the Han fossa is present and the infraorbital sils may have to be allocated to a new
River at Yunxian in western Hubei.60 region is set at an angle to the flaring taxon. However, as noted earlier, hol-
The finds were made in a clay layer cheek. Given this mix of characters, Li lowing of the infraorbital surface can
and both hominids were encased in and Etler60 and Etler61 choose to place be documented for faces outside of
hard calcareous matrix. The same level the skulls with Homo erectus, but China. Furthermore, the new finds
has produced mammalian fossils and Zhang63 identifies them with later hu- from Gran Dolina suggest that this
some stone cores and flakes. The fauna mans. feature may appear in Europe at the
suggests a Middle Pleistocene age. Pa- Other fossils are known from later beginning of the Middle Pleistocene.
leomagnetic work coupled with other Middle Pleistocene localities in China. Such evidence will make it harder to
approaches now indicates that the The Dali and Jinniushan specimens argue for isolation of the major Old
Yunxian assemblage may be as much are often described as archaic or pre- World geographic provinces. The
as 600,000 or even 800,000 years modern Homo sapiens.64 The Dali cra- spread of some populations of Homo
old.61,62 nium is quite complete. Its massive heidelbergensis into the Far East can-
Unfortunately, the crania themselves brow, keeled frontal, and low vault are not be ruled out.
226 Evolutionary Anthropology ARTICLES

CONCLUSIONS ACKNOWLEDGMENTS 16 Bar-Yosef O (1995) The role of climate in the

interpretation of human movements and cultural
In their well-known 1975 volume My studies of Pleistocene hominids transformations in western Asia. In Vrba ES,
Denton GH, Partridge TC, Burckle LH (eds),
After the Australopithecines,66 Karl have been conducted with the assis- Paleoclimate and Evolution with Emphasis on
Butzer and Glynn Isaac noted many tance of many individuals and institu- Human Origins, pp 507–523. New Haven: Yale
uncertainties surrounding human evo- tions, and I am grateful for this help. University Press.
lution in the Middle Pleistocene. Al- The governments of China, Ethiopia, 17 Gabunia L, Vekua A (1994) A Plio-Pleistocene
hominid from Dmanisi, East Georgia, Caucasus.
most a quarter of a century later, the Indonesia, Kenya, and Tanzania Nature 373:509–512.
‘‘muddle in the middle’’ is still evident, granted me clearance to examine fos- 18 Bräuer G, Schultz M (1996) The morphologi-
especially in respect to systematics sils in these countries. The L.S.B. cal affinities of the Plio-Pleistocene mandible
from Dmanisi, Georgia. J Hum Evol 30:445–481.
and classification of the hominids. Per- Leakey Foundation and the Boise Fund
19 Tchernov E (1987) The age of the ‘Ubeidiya
haps the most vexing questions con- supported much of the research on Formation, an Early Pleistocene hominid site in
cern fossils of earlier Middle Pleis- which this paper is based. R.G. Klein, the Jordan Valley, Israel. Israel J Earth Sci 36:3–
tocene antiquity. Specimens from R. Quam, and C.B. Stringer kindly 30.
20 Villa P (1991) Middle Pleistocene prehistory in
Africa and Eurasia have most fre- commented on a version of the manu- southwestern Europe: The state of our knowledge
quently been described as ‘‘archaic’’ script, as did several anonymous re- and ignorance. J Anthropol Res 47:193–217.
representatives of our own species, viewers. 21 Roebroeks W (1994) Updating the earliest
but this situation is unsatisfactory for occupation of Europe. Curr Anthropol 35:301–
several reasons. Fossils such as those REFERENCES 22 Ascenzi A, Biddittu I, Cassoli PF, Segre AG,
found at Bodo, Broken Hill, Arago, Segre-Naldini E (1996) A calvarium of late Homo
1 Andrews P (1984) An alternative interpretation erectus from Ceprano, Italy. J Hum Evol 31:409–
Petralona, and Dali retain many primi- of characters used to define Homo erectus. Cour 423.
tive erectus-like characters, and this Forsch-inst Senckenberg 69:167–175.
23 Swisher CC, Curtis GH, Jacob T, Getty AG,
anatomy sets them apart from recent 2 Groves CP (1989) A Theory of Human and Suprijo A, Widiasmoro (1994) Age of the earliest
humans. Simply lumping diverse an- Primate Evolution. Oxford: Oxford University known hominids in Java, Indonesia. Science 263:
Press. 1118–1121.
cient groups with living populations 3 Larick R, Ciochon RL (1996) The African emer- 24 Swisher CC, Rink WJ, Anton SC, Schwarcz
obscures these differences. gence and early Asian dispersals of the genus HP, Curtis GH, Suprijo A, Widiasmoro (1996)
There is increasing acceptance of Homo. Am Sci 84:538–551. Latest Homo erectus of Java: Potential contempo-
the suggestion that distinct lineages 4 Wood B (1991) Koobi Fora Research Project, Vol. raneity with Homo sapiens in southeast Asia.
4. Hominid Cranial Remains. Oxford: Clarendon. Science 274:1870–1874.
may have evolved during this period. 5 Wood B (1994) Taxonomy and evolutionary 25 Groves CP (1994) The origin of modern hu-
One possibility is that fossils from relationships of Homo erectus. Cour Forsch-inst mans. Interdisciplinary Sci Rev 19:23–34.
Africa and Europe can be sorted to- Senckenberg 171:159–165. 26 Dean D, Delson E (1995) Homo at the gates of
gether to a single taxon, appropriately 6 Wolpoff MW, Thorne A, Jelinek J, Zhang Y Europe. Science 373:472–473.
(1994) The case for sinking Homo erectus: 100 27 Harrison T (1993) Cladistic concepts and the
called Homo heidelbergensis. This spe- years of Pithecanthropus is enough! Cour Forsch- species problem in hominoid evolution. In Kim-
cies may have originated in Africa. If inst Senckenberg 171:341–361. bel WH, Martin LB (eds), Species, Species Con-
the Gran Dolina fossils are also Homo 7 Wolpoff MW (1996) Human Evolution. New cepts and Primate Evolution, pp 345–371. New
York: McGraw-Hill. York: Plenum Press.
heidelbergensis, then these people ap-
8 Tobias PV (1995) The place of Homo erectus in 28 Foley R, Lahr MM (1997) Mode 3 technologies
parently reached Europe at an early nature with a critique of the cladistic approach. and the evolution of modern humans. Cambridge
date. In this region, populations iso- In Bower JRF, Sartono S (eds), Human Evolution Archaeol J 7:3–36.
in its Ecological Context, Vol. 1. Palaeoanthropol- 29 Kalb JE, Wood CB, Smart C, Oswald EB,
lated by glacial conditions perhaps ogy: Evolution and Ecology of Homo Erectus, pp Mabrate A, Tebedge S, Whitehead P (1980) Pre-
were eventually ancestral to the Nean- 31–41. Leiden: Pithecanthropus Centennial Foun- liminary geology and palaeoecology of the Bodo
derthals. In other parts of the species dation. d’Ar hominid site, Afar, Ethiopia. Palaeogeogr
range, including Africa, there are indica- 9 Rightmire GP (1990) The Evolution of Homo Palaeoclimatol Palaeoecol 30:107–120.
Erectus. Comparative Anatomical Studies of an 30 Clark JD, Asfaw B, Assefa G, Harris JWK,
tions that later Middle Pleistocene groups Extinct Human Species. Cambridge: Cambridge Kurashina H, Walter RC, White TD, Williams MA
were evolving in the direction of Homo University Press. (1984) Paleoanthropological discoveries in the
sapiens. Homo heidelbergensis is thus the 10 Bräuer G, Mbua E (1992) Homo erectus fea- Middle Awash valley, Ethiopia. Nature 307:423–428.
tures used in cladistics and their variability in 31 Clark JD, de Heinzelin J, Schick KD, Hart WK,
stem from which both Neanderthals Asian and African hominids. J Hum Evol 22:79– White TD, WoldeGabriel G, Walter RC, Suwa G,
and modern humans are derived. 108. Asfaw B, Vrba E, H.-Selassie Y (1994) African
A problem is whether the same taxon 11 Kramer A (1993) Human taxonomic diversity Homo erectus: Old radiometric ages and young
in the Pleistocene: Does Homo erectus represent Oldowan assemblages in the Middle Awash val-
can be identified in the East. Fossils multiple hominid species? Am J Phys Anthropol ley, Ethiopia. Science 264:1907–1910.
such as those found at Dali and Jinni- 91:161–171. 32 Rightmire GP (1996) The human cranium
ushan in China are more advanced 12 Bräuer G (1994) How different are Asian and from Bodo, Ethiopia: Evidence for speciation in
than Homo erectus and exhibit some African Homo erectus? Cour Forsch-inst Sencken- the Middle Pleistocene? J Hum Evol 31:21–39.
berg 171:301–318. 33 Klein RG, Cruz-Uribe K (1991) The bovids
of the same derived characters as do 13 Walker A (1993) Perspectives on the Narioko- from Elandsfontein, South Africa, and their impli-
the specimens from Africa and Eu- tome discovery. In Walker A, Leakey R (eds), The cations for the age, palaeoenvironment and ori-
rope. But it can be argued that the Nariokotome Homo Erectus Skeleton, pp 411–430. gins of the site. Afr Archaeol Rev 9:21–79.
Cambridge: Harvard University Press. 34 Klein RG (1994) Southern Africa before the
Chinese hominids are distinctive in
14 Rightmire GP (1998) Evidence from facial Iron Age. In Corruccini RS, Ciochon RL (eds),
aspects of their facial morphology. morphology for similarity of Asian and African Integrative Paths to the Past: Paleoanthropological
Some workers will prefer either to representatives of Homo erectus. Am J Phys An- Advances in Honor of F. Clark Howell, pp 471–519.
thropol 106:61–85. Englewood Cliffs: Prentice Hall.
place them in a new species or lump
15 Brown B (1994) Comparative dental anatomy 35 Grün R (1996) A re-analysis of electron-spin
them as early Homo sapiens. This ques- of African Homo erectus. Cour Forsch-inst Senck- resonance dating results associated with the Pe-
tion remains to be resolved. enberg 171:175–184. tralona hominid. J Hum Evol 30:227–241.
ARTICLES Evolutionary Anthropology 227

36 Cook J, Stringer CB, Currant AP, Schwarcz d’Homo Sapiens, pp 291–327. Paris: Presses Uni- Parmi les Hominidés Fossiles, pp 154–177. Nice:
HP, Wintle AG (1982) A review of the chronology versitaires de France. CNRS.
of the European Middle Pleistocene hominid 47 Arsuaga JL, Martı́nez I, Gracia A, Carretero 56 Howells WW (1973) Cranial variation in man.
record. Yrbk Phys Anthropol 25:19–65. JM, Carbonell E (1993) Three new human skulls A study by multivariate analysis of patterns of
37 Stringer CB (1974) A multivariate study of the from the Sima de los Huesos Middle Pleistocene difference among recent human populations. Pa-
Petralona skull. J Hum Evol 3:397–404. site in Sierra de Atapuerca, Spain. Nature 362: pers of the Peabody Museum 67:1–259.
38 Stringer CB (1983) Some further notes on the 534–537. 57 Schwartz JH, Tattersall I (1996) Significance
morphology and dating of the Petralona homi- 48 Condemi S (1996) Does the human fossil of some previously unrecognized apomorphies in
nid. J Hum Evol 12:731–742. specimen from Reilingen (Germany) belong to the nasal region of Homo neanderthalensis. Proc
39 Bräuer G (1984) A craniological approach to the Homo erectus or to the Neanderthal lineage? Natl Acad Sci 93:10852–10854.
the origin of anatomically modern Homo sapiens Anthropologie 34:69–78. 58 Pares JM, Peres-Gonzalez A (1995) Paleomag-
in Africa and implications for the appearance of 49 Hublin J-J (1996) The first Europeans. Archae- netic age for hominid fossils at Atapuerca archaeo-
modern Europeans. In Smith FH, Spencer F ology 49:36–44. logical site, Spain. Science 269:830–832.
(eds), The Origins of Modern Humans: A World 50 Stringer CB (1995) The evolution and distribu- 59 Bermúdez de Castro JM, Arsuaga JL, Carbon-
Survey of the Fossil Evidence, pp 327–410. New tion of later Pleistocene human populations. In ell E, Rosas A, Martı́nez I, Mosquera M (1997) A
York: Alan R. Liss. hominid from the Lower Pleistocene of Atapu-
Vrba ES, Denton GH, Partridge TC, Burckle LH
40 Van Vark GN (1995) The study of hominid (eds), Paleoclimate and Evolution with Emphasis erca, Spain: Possible ancestor to Neandertals and
skeletal remains by means of statistical methods. on Human Origins, pp 524–531. New Haven: Yale modern humans. Science 276:1392–1395.
In Boaz NT, Wolfe GD (eds), Biological Anthropol- University Press. 60 Li T, Etler D (1992) New Middle Pleistocene
ogy: The State of the Science, pp 71–90. Corvallis: hominid crania from Yunxian in China. Nature
51 Carbonell E, Bermúdez de Castro JM, Arsuaga
Oregon State University Press. 357:404–407.
JL, Diez JC, Rosas A, Cuenca-Bescos G, Sala R,
41 Roberts MB, Stringer CB, Parfitt SA (1994) A Mosquera M, Rodriguez XP (1995) Lower Pleis- 61 Etler D (1996) The fossil evidence for human
hominid tibia from Middle Pleistocene sediments tocene hominids and artifacts from Atapuerca- evolution in Asia. Ann Rev Anthropol 25:275–301.
at Boxgrove, UK. Nature 369:311–313. TD6 (Spain). Science 269:826–830. 62 Chen T, Yang Q, Hu Y, Bao W, Li T (1997) ESR
42 Stringer CB (1993) New views on modern 52 Arsuaga J-L, Martı́nez I, Gracia A, Lorenzo C dating of tooth enamel from Yunxian Homo
human origins. In Rasmussen DT (ed), The Origin (1997) The Sima de los Huesos crania (Sierra de erectus site, China. Q Sci Rev 16:455–458.
and Evolution of Humans and Humanness, pp Atapuerca, Spain). A comparative study. J Hum 63 Zhang Y (1995) Fossil human crania from
75–94. Boston: Jones and Bartlett. Evol 33:219–281. Yunxian: Morphological comparison with Homo
43 Groves CP, Lahr MM (1994) A bush not a 53 Bischoff JL, Fitzpatrick JA, Leon L, Arsuaga erectus crania from Zhoukoudian. Acta Anthro-
ladder: Speciation and replacement in human JL, Falgueres C, Bahain JJ, Bullen T (1997) pol Sinica 14:1–7.
evolution. Perspect Hum Biol 4:1–11. Geology and preliminary dating of the hominid- 64 Wu XZ, Poirier FE (1995) Human Evolution in
44 Howell FC (1960) European and northwest bearing sedimentary fill of the Sima de los Hue- China. A Metric Description of the Fossils and a
African Middle Pleistocene hominids. Curr An- sos chamber, Cueva Mayor of the Sierra de Atapu- Review of the Sites. New York: Oxford University
thropol 1:195–232. erca, Burgos, Spain. J Hum Evol 33:129–154. Press.
45 Vandermeersch B (1985) The origin of the 54 Spitery J (1982) La face de l’homme de Tau- 65 Harding RM, Fullerton SM, Griffiths RC, Bond
Neandertals. In Delson E (ed), Ancestors: The tavel. In de Lumley M-A (ed), L’Homo Erectus et la J, Cox MJ, Schneider JA, Moulin DS, Clegg JB
Hard Evidence, pp 306–309. New York: Alan R. Place de l’Homme de Tautavel Parmi les Hominidés (1997) Archaic African and Asian lineages in the
Liss. Fossiles, pp 110–136. Nice: CNRS. genetic ancestry of modern humans. Am J Hum
46 Hublin J-J, Tillier A-M (1991) L’Homo sapiens 55 De Lumley M-A, Spitery J (1982) Le maxillaire Genet 60:772–789.
en Europe occidentale: Gradualisme et rupture. de l’homme de Tautavel. In de Lumley M-A (ed), 66 Butzer KW, Isaac GL (1975) After the Australo-
In Hublin J-J, Tillier A-M (eds), Aux Origines L’Homo Erectus et la Place de l’Homme de Tautavel pithecines. The Hague: Mouton.

Related Interests