Morphological Variation in 20

Homo neanderthalensis and

Homo sapiens in the Levant
A Biogeographic Model
YOEL RAK

Introduction

It was only the geographic proximity of the Mount Carmel specimens that
spared them from being assigned to many different taxa as "separate form[s] of
humanity." Were it not for these circumstances, McCown and Keith ( 1939) would
not have conceived of accommodating such a great range of variation in one
taxon. However, a clear morphological dichotomy between the hominids from
Skhul and those from Tabun (C-1, the female skeleton, and C-2, the isolated
mandible) emerges from their monograph. The existence of two kinds of homi-
nids in a relatively small geographic area of the Middle East has since been
confirmed through discoveries at several additional sites. Other hominids have
been found, including specimens from Amud (Suzuki and Takai, 1970) and
Shanidar (Trinkaus, 1983), which can be grouped comfortably with the Tabun
specimens, whereas specimens uncovered at Qafzeh (Vandermeersch, 1981) can
be added to the Skhul group. Until quite recently, McCown and Keith's basic
contention-that the Neanderthal-looking Tabun group represented the earlier,
primitive anatomy, and the modern-looking Skhul group represented the later,
derived anatomy-was generally accepted.

YOEL RAK • Department of Anatomy, Sackler Faculty of Medicine, Tel Avtv University, Tel Avtv,
Israel, and Institute of Human Origins, Berkeley, California 94 709.
Speues, Speczes Concepts, and Pnmate Evolutwn, edited by William H. Kimbel and Lawrence B. Martin.
Plenum Press, New York, I 993.

523

W. H. Kimbel et al. (eds.), Species, Species Concepts and Primate Evolution

© Springer Science+Business Media New York 1993
524 BIOGEOGRAPHIC MODEL OF H. NEANDERTHALENSIS AND H SAPIENS

The purpose of this chapter is threefold. The first is to demonstrate that the
Neanderthals differ, on the specific level, from Homo sapzens; in other words, to
show that from the perspective of the traditional morphological standards used
in primate taxonomy, the magnitude of difference in the face and in the pelvis,
justifies such a taxonomic decision. The second goal is to demonstrate that the
phylogenetic relationship between H. sapiens and Neanderthals, which is based
on their morphology and the emerging timetable, cannot be anagenetic. Even
when Neanderthals are recognized as a separate species, the morphological and
temporal factors do not allow us to consider the Neanderthal an ancestral species
to H. sapiens, as did some investigators in the past who endorsed the Nean-
derthal's species status. The third goal of the chapter is to propose an explana-
tion for the morphological variation characterizing the hominids of the
Mousterian period in the Middle East.

Morphological and Temporal Considerations

Although in the past the Qafzeh specimens were claimed by some to be

contemporary with or even older than the Neanderthals (much older than com-
monly thought at the time; Haas, 1972; Tchernov, 1984), there was a general
reluctance to accept this possibility. At the root of this reluctance were the facts
that the chronology was based on biostratigraphic considerations rather than
absolute dates and, perhaps even more significantly, the difficulty that many
investigators have in accepting a cladogenetic scenario for such a late segment of
human evolution. Indeed, this combination of factors led McCown and Keith
(1939) to argue with Garrod about the need to view the Tabun skeleton as
chronologically earlier than the specimens from Skhul; and, many years later, it
also led Trinkaus (1984, 258) to write as follows:

All of the techniques that have been used to date the Qafzeh remains have limitations,
and each IS insufficient to confirm a date. However, the younger date would alleviate
the need to account for how two dtstmct groups of Homo sapzens could have coexisted
for several millennia in the small area of northern Israel, using the same cultural
adaptive complex and yet remainmg biologically distinct. For the purposes of this
dtscussion, I will employ Occam's Razor, use the date for the Qafzeh remains that
creates the fewest problems, and consider them to date to around 40 ky BP, about the same
age as the Skhullayer B remains. [my emphasis!

In recent years, excavations were reopened at the Kebara Cave in Israel

(Bar-Yosef et al., 1986). As a result, the Neanderthal presence on Mount Carmel
was reconfirmed through diagnostic elements of the skeleton, and the chro-
nology and the stratigraphy of the Middle Paleolithic in Israel were reassessed
with the aid of modern techniques of absolute dating (Valladas et a!., 1987,
1988). In its general appearance and anatomical detail, the mandible of the
skeleton discovered in Kebara in 1983 can easily be identified as that of a "clas-
sical" Neanderthal. Tillier et al. ( 1989) reached this conclusion and thus lumped
this specimen with the other Neanderthal specimens of the Middle East (those
from Amud, Shanidar, and Tabun). On the basis of the mandible's typical ap-
 YOEL RAK 525
pearance, I am confident that the face of this individual, whose cramum ts
missing, was that of a "classical" Neanderthal.
Similarly, the specimen's pelvis exhibits the very long and slender superior
pubic ramus found in other Neanderthals, in contrast to the short, thick ramus
seen in modern humans and the Mousterian hominids from Qafzeh and Skhul
(Rak, 1990; see also Trinkaus, 1976). The pelvis's superb state of preservation
provides new information on Neanderthal pelvic anatomy and permits us to
evaluate the functional significance of these differences. I believe that the ana-
tomical differences between the Neanderthal pelvis and the modern pelvis indi-
cate profound biomechanical differences.
The length of the superior pubis, about 90 mm, is greater by far than what
we would expect, given the size of the inlet. Specimens of modern H. sapiens
having an inlet size similar to that of the Kebara Neanderthal exhibit an average
ramus length of only 70 mm! The ratio of ramus length to inlet width in the
Neanderthal, whose index is 60.3, is much greater than that in modern humans,
where the mean index for males is 55.3 (n = 43; SD = 3.1). The pelvic inlet of the
Neanderthal is only 13% wider (transversely) than the average inlet of modern
males, whereas the superior pubic ramus of the former is 31% longer than that
of the latter. Clearly, it is not the size of the inlet, of which the ramus constitutes a
major part, that dictates the length of the ramus. In an attempt to understand
how to accommodate such a long ramus with a normal-sized inlet, I came to the
conclusion that it is not the ramus length in itself that is significant but, rather,
the distance from the acetabulum to the symphysis pubis. In other words, the long
ramus of the Neanderthal stems from the fact that the acetabulum is located
farther posterolaterally from the symphysis (along the circumference of the inlet
rim) than in the modern male. Indeed, most of the anatomical differences that
distinguish the Neanderthal pubis from that of modern humans stem from that
shift in the position of the acetabulum.
The position of the acetabulum can be evaluated metrically through the use
of an index that relates it to the anteroposterior dimension of the inlet. With the
pelvis held such that the inlet rim is in a horizontal orientation, perpendiculars
are dropped from the anterior edge of the acetabulum and from the anterior
and posterior ends of the inlet at the midline. An index with a small value
denotes a relatively large distance between the coronal plane of the acetabulum
and that of the symphysis pubis, and a relatively small distance between the coronal
plane of the acetabulum and the sacrum. The average index in modern males is
Sl.O (n = 57; SD = 3.5), and in the Kebara Neanderthal, it is 61.5. In other
words, the acetabulum in the modern male is located closer to the coronal plane
of the symphysis and more anterior to the sacrum than in the Neanderthal. It is
my contention (Rak, manuscript in preparation) that the forward migration of
the acetabula in modern humans has made the pelvis into a specialized cushion-
ing device that absorbs the weight of the cyclic dropping of the trunk during
walking (Fig. 1). The discrepancy between the coronal planes of the acetabula
and the sacrum prevents the falling center of mass from landing directly on and
pounding against the femur heads. The forward position of the acetabula and
the concomitantly short superior pubic ramus result in a short lever arm. The
result is that the ramus is subjected to great bending moments. The thickened
modern ramus can be viewed, therefore, as a necessary reinforcement of this
526 BIOGEOGRAPHIC MODEL OF H NEANDERTilALENSIS AND H SAPIENS

Fig. I. A schematic vtew of two pelvic configurations depicted in an exaggerated

manner. Upper. The primitive configuration as in the Neanderthal. Lower: The
derived state, as m the modern human, where a relatively large dtstance is formed
between the acetabula and the venebral column.

short lever arm (Rak and Rosenberg, manuscript in preparation). As the signifi-
cance of the length and thickness of the superior pubic ramus becomes clear, this
element ceases to be merely an anatomical feature of vague functional value.
Even its smallest fragments are of great importance because of their biomechani-
cal implications.
As we cannot carry out the traditional practice of comparing these pelvises
with specimens from an outgroup,* we are relying on biomechanic assessments
to help us determine the polarity of the two bipedal pelvic configurations-the
primitive configuration as opposed to the derived-and are thus, in essence,
applying Ridley's line of thought exposed in his chapter entitled "The Func-
tional Criterion" ( 1986). Ridley portrays the usefulness of functional criteria not
in categorically determining polarity, but rather in increasing the level of confi-
dence in a given phylogenetic decision. He ( 1986: 136) writes:
The functional criterion will not usually give certamty. It is virtually certain in the case
of vestigal organs, but they are an exception. Usually, the method will not allow us to
conclude anythmg more than that one dtrection of evolution is more probable that the
other. No doubt the further study of any particular case might often make the conclu-
sion more certain, but at any one moment we may have to hve with uncertamty.

*There are no Middle Pleistocene pelvises complete enough for comparison, and although in some
aspects the pelvic anatomy ofAustralopt-thecus is reminiscent of that found in H. neanderthalenszs, I am
reluctant to go so far as to view the Pliocene hominid condition as the ancestral morphotype for the
Kebara and other Neanderthal pelvises.
 YOEL RAK 527
On the basis of the morphological comparison and evaluation of the func-
tional advantage of one form over the other, I believe that the Neanderthal
pelvis still represents the primitive configuration, and the modern pelvis repre-
sents the derived character state. Indeed, in long-distance walking, the modern
human pelvis appears to have the edge over the Neanderthal pelvis.
What I view as the primitive morphology, that of the Neanderthal pelvis,
could easily have evolved through a relatively simple modification-the forward
migration of the acetabula-into the derived form exhibited by the modern
pelvis. However, the morphology of the Qafzeh pelvis, which is identical in every
way to that of the modern pelvis (Rak, 1990), combined with the fossil specimen's
geologic age (Valladas et al., 1987), render unlikely the scenario of the modern
pelvis having evolved from the pelvis of the known Neanderthals. In the evalua-
tion of the relationship between the Qafzeh and the Kebara specimens, chrono-
logical and anatomical considerations impose a cladogenetic geometry.
A cladogenetic picture also emerges when the face of the Neanderthal is
examined and compared with that of modern humans. In this case, however, the
Neanderthal face represents the derived morphology. The Neanderthal face
emerges as unique, deviating both morphologically and architecturally from the
pattern seen in other hominid taxa (and in the primates in general; Rak, 1986).
Howells (197 5) concludes his metric analysis with the comment that the faces of
Homo erectus and JI. sapiens have more in common than either of them has with
the Neanderthal face. Even a cursory examination of hominids, such as KNM-
ER 3733 and SK 847, reveals a morphology more akin to H. sapiens than to the
Neanderthal. From the point of view of phylogenetic analysis, it is much more
economical to link H. sapiens directly to the hominid form represented by KNM-
ER 3733 and SK 847 than to interpose the unique facial and cranial anatomy of
the Neanderthal between them. If we accept this simpler phylogenetic hypoth-
esis, then the discovery of hominids (Zuttiyeh, Skhul, and Qafzeh) that are
contemporary with the Neanderthals and yet have a primitive that is, modern-
looking, face should not come as a surprise and should even be expected.
This idea, of course, is not new and has come up in almost every discussion
of the phylogenetic position of the Neanderthals since their discovery (see re-
views by Bowler, 1986; Mann and Trinkaus, 1974; Spencer and Smith, 198I).
Most notably, it was promoted by Boule (1911-1913), who influenced the views
of many. Among the more modern investigators, Howells was the one who in
1942 came to the conclusion that the 1\'eanderthals play no part in the evolution-
ary history of modern humans and actually constitute a separate species from H.
sapiens. Weidenreich ( 1943, p. 39), arguing against Howells' hypothesis, said that
it "proves that the view of the independency of Neanderthal Man is still advo-
cated, in spite of all that has been written against it in recent years." With his
famous diagram (reproduced here as Fig. 2), Weidenreich attempted to demon-
strate the Neanderthal's intermediate position between H. erectus and H. sapiens.
So eager was he to force the Neanderthal into a unilinear evolutionary scheme
that he failed to distinguish between a morphocline with a defined polarity and
the actual phylogeny itself. The advocates of the two slightly different theories-
the "presapiens" and the "preneanderthal"-maintained, to varying degrees,
that the Neanderthals, specifically the European Neanderthals, constituted a
528 BIOGEOGRAPHIC MODEL OF H NEANDERTHALENSIS AND H. SAPIENS
v
9

8 b

0
0 2 3 4 5 6 7 8 9 10 11 12 13

Modem Man Sinanthropus Pithecanthropus

Neanderthalians Homo soioensis Anthropoids

Fig. 2. Weidenreich's famous diagram showmg "the gradual expansiOn of the bramcase in the course
of human evolution" (1943, p. 46). Wetdenreich offered thts diagram as support of his idea of
unilinear evolution, in which Neanderthals precede H. sap~ens phylogenetically. Apparently, he faded
to distinguish between a morphoclme and the actual phylogeny.

side branch of the main stem leadzng to modern humans (Howell, 1952; Vallois,
1954; Boule and Vallois, 1957).
The truth of the matter is that even among the more enthusiastic advocates
of a unilinear model of human evolution (Weidenreich, 1947, 1949; Weinert,
1953), certain cladogenetic aspects are to be found that undoubtedly stem from
the vague anatomical definition of the Neanderthal taxon itself. One can readily
understand how Hrdlicka's definition of the Neanderthal-a definition still ac-
cepted by some scholars-might be an endless source of taxonomic and phy-
logenetic confusion. He writes that "the only workable definition of Neanderthal
man and period seems to be, for the time being, the man and period of the
Mousterian culture" (1927, p. 251). In the 1970s, a period in which the tax-
onomic pendulum seemed to rest on the side of the unilinear model, F. C.
Howell (1973, p. 123) still asked, in the heading of a chapter, "just who was the
Neanderthal?" In this choice of a title, he signaled his discontent with, and the
problematics of, viewing the Neanderthal's unique morphology as a direct pred-
ecessor of modern human morphology.
Once again, the reluctance to remove the Neanderthal from our ancestry
and to accept the cladogenetic model seems to be what leads Trinka us ( 1983,
1984, 1987) and investigators who endorse his view (Smith and Paquette, 1989)
to see the Neanderthal facial configuration as resulting primarily from two
distinct trends: the posterior withdrawal of the zygomatic bones to their position
in modern humans and the persistence of the prognathic configuration of H.
erectus in the rest of the face. The Neanderthal, according to this approach, can
 YOEL RAK 529
still serve as an intermediate stage between H. erectus and modern humans.
Without entering into the details of their argument, I would like to stress that the
retreat of the zygomatic bones has never been convincingly demonstrated. For
example, the position of the zygomatic process of the maxilla relative to the
dental arcade probably does not indicate a retreat of the zygomatic bone, as
claimed by Trinka us ( 1987), but rather stems from the more sagittal orientation
of both the zygomatic bone and the zygomatic process of the maxilla, and the
attachment of these elements, at that orientation, to a wide maxillary body (see
also Rak, 1986, 1991 ). Furthermore, it is not at all clear how this supposed retreat
could have resulted in the particular morphology manifested in certain Nean-
derthal facial elements that can only be viewed as derived because they differ
from corresponding elements that H. sapiens and H. erectus have in common. For
example, let us note the immense width of the pyriform aperture, the flaring
nasal apophysis, and the horizontally oriented nasal bones; all of these features
combine to produce the dramatic prominence of the nasal bridge and the typical
midfacial prognathism, on the one hand, and the distinct subdivision of the
infraorbital region, on the other (Rak, 1986).
Although many attempts have been made to interpret the uniqueness of the
Neanderthal face (see discussion in Rak, 1986), the question of the functional
significance is of little relevance here. A comparison with the faces of other
hominids and, indeed, other primates, is what reveals this uniqueness.
It is the combination of a derived face with a primitive pelvis that enables us
to diagnose Neanderthal as a separate species from H. sapiens, which is charac-
terized by the reverse-a primitive face and a derived pelvis (Fig. 3). These
opposite combinations and the emerging chronological picture point to the con-
temporaneity of two distinct species and compel us to recognize that two lineages
were evolving separately. It is still the traditional taxonomic considerations-the
magnitude of morphological differences between the Neanderthal and H. sa-
puns-that constitute the basis for proposing species status for the Neanderthal.
Both the morphology and the basic architectural plan of the face, as well as
fundamental differences in the pelvis, have led me to consider the Neanderthals
as a distinct species-Homo neanderthalensis King. In recent years, other investi-
gators have reached the same conclusion (see, for example, Stringer and Gn1n,
1991; Tattersall, 1986; Howell, 1991). On the basis of anatomical considerations
and paleontological criteria, along with the revised geologic age, we cannot but
falsify the hypothesis that H. neanderthalensis represents a stage in an evolution-
ary sequence leading to H. sapzens.
Some of the Middle Eastern Neanderthal specimens (most notably Tabun
C-1, Shanidar 5, and Kebara) are "classical" in appearance and resemble those of
Western Europe. However, other specimens (Tabun C-2, Shanidar 2 and 4, and
Amud I) do not adhere to this classical image. Although a broad consensus
maintains that the latter are Neanderthals, their face and mandible do exhibit a
more modern-looking morphology, and hence, they are considered by some
investigators as closer to modern H. sapiens than the former group is. However, I
believe that these specimens (Tabun C-2, Shanidar 2 and 4, and Amud I) con-
stitute the more przmitive segment of the spectrum characterizing the species;
and since the face of modern H. sapzens (like that of H. erectus) shows, in essence,
the primitive configuration, the more generalized morphology of these spec-
530 BIOGEOGRAPHIC MODEL OF H. NEANDERTHALENSI S AND H SAPIENS

FACE PELVIS

H.n. H.s. H.n. H.s.

I~ I
Fig. 3. The proposed phylogenetic relationship between H sapzen.1 and Neanderthal. The diagram
on the left is based on the face and shows the Neanderthal as the derived form. The diagram on the
nght is based on the pelvis and shows the Neanderthal as the primitive form and H. sap~ens as the
derived. Only the true pelvis is depiCted.

imens is phylogenetically uninformative. They do not provide evidence of in-

terbreeding, and they certainly do not represent a late or even terminal stage in
the H. neanderthalensis evolutionary sequence-an intermediate link between the
"classical" Neanderthal and modern H. sapzens. Their somewhat modern-looking
face does not suggest that the anatomy appeared late, despite the fact that it
resembles the anatomy of the only surviving species, the latest in the phylogeny.
The face of these individuals is simply more primitive, with no chronological
implications necessarily attached. Similarly, and for the same reasons, we can
reasonably expect to find that in the parallel lineage of H. sapiens, part of the
morphological spectrum of derived pelvises will include specimens with a some-
what more primitive appearance, closer to that of H. neanderthalensis. The pres-
ence of a sapzens-like morphology in the spectrum of Neanderthal faces (like the
predicted, more generalized pelvis in part of the H. sapiens clade) is not evidence
of a chronologically late stage; on the contrary, this morphology is simply a
reflection of phylogenetic heritage.
 YOEL RAK 531
A Biogeographic Model

The model that I propose offers an explanation for the great variat.ion
characterizing the Mousterian hominids in the Middle East: how the caves of the
period could have been inhabited by hominids exhibiting such a wide mor-
phological range, or, in other words, how such a relatively restricted geographic
region could have accommodated H. sapiens, H. neanderthalensis, and those popu-
lations that purportedly fill in the morphological gap between the two.
A generalized appearance can stem not only from an early geologic age.
Sexual dimorphism and young ontogenetic age are well-known agents that in-
troduce generalized morphology and thus, also, variation into the spectrum.
Another cause of variation is geographic distance, which has been recognized as
"a powerful mechanism for achieving genetic divergence" (Mayr, 1963, p. 517;
Wright, 1943). The gradual change of a given trait in contemporaneous popula-
tions produces a morphocline that begins at a generalized source and exhibits
differentiation across a broad geographic area, as the distance introduces new
ecological niches. A specific case of such distribution is a long chain of clines that
curves back on itself. The chain runs along a restricted geographic route (such as
the shoreline of an island or the margins of a valley), and the ends overlap in
sympatric populations, producing the well-known "chain of races," or rassen-
kreis. *
What Mayr (1963) calls "speciation by distance" has been demonstrated
frequently. It is expressed in an increase in sterility over geographical distance.
The far ends of the sequence do not interbreed, and they are intersterile. Never-
theless, Goldschmidt ( 1940), for example, insisted that because of genetic con-
tinuity, the ends should not be considered distinct species. Dobzhansky (1964, p.
270), on the other hand, recognized the problem of the systematist who "is
confronted with the impossibility of drawing the 'line' anywhere between the two
[species]" but yet claims that since the far ends are genetically isolated, they
should be considered separate species.
One model that may be able to account for hominid diversity in the Middle
East is based, therefore, on the phenomenon of geographic differentiation along
a cline. If we consider the face, this model places the primitive end of the
morphocline in Africa in the form of an early H. sapiens and the derived end
deep within Europe in the form of H. neanderthalens1s. The model predicts that
the face of specimens from the Middle East, which is situated between these two
regions, will be more generalized, that is, less differentiated, than that of the
more western Neanderthals. However, even such a restricted site as the northern
part of Israel presents a more complicated picture of morphological variation.
This proposed zoogeographic model is horizontal and thus lacks the depth
necessary to reflect the time span of the Mousterian period, during which the
territorial range of H. neanderthalensis habitation might well have f1uctuated. The
caves of Israel, situated on the periphery of the Neanderthal range, thus alter-
natively "sampled" whichever of the populations prevailed at a given time:

*Mayr and Ashlock (1991, p. 427) draw attention to the misuse of the German word rassenkms. They
maintain that the term translates as "polytypic species" rather than "circle of races."
532 BIOGEOGRAPHIC MODEL OF H. NEANDERTHALENSIS AND H. SAPIENS

Homo
neanderthalens1s

Homo
!iapiens

--- .. ---- .. .:·~--- ..

--- ::.r•··-.

-····- ..

Ftg. 4. Fluctuations m the terntory of the Neanderthal (black) and of early H. saptens (white) shown
on a map of circa Mediterranea. The Middle East Moustenan stratigraphy is indicated by the
cylinder.

Expansion of the Neanderthal territory brought a H. neanderthalensis presence

into the caves of the region, and shrinkage of the territory, with the correspond-
ing expansion of the H. sapuns territory, led to the presence of H. sapiens in the
caves. The model actually predicts that the varying size of the Neanderthal
territory will be manifested in a corresponding degree of anatomical differentia-
tion in a given cave. In other words, a particulary extensive expansion of the
Neanderthal territory would have brought a much more classical-looking Nean-
derthal into the Galilee caves than a smaller expansion would have. Hence,
oscillation in the size of the territory can be expected to have introduced differ-
ent anatomical forms that interdigitate stratigraphically, even within a single
cave (Fig. 4).
In essence, the effect of the dimension of time (on a greater scale than the
 YOEL RAK 533
periods described above) on a vertical evolutionary lineage does not differ from
the effect of geographic distance on a horizontal morphocline. According to
Mayr ( 1942, 1949), populations are thus expected to show increasing differentia-
tion along an evolving lineage. As in the case of a geographic morphocline,
researchers such as Mayr (1942), Simpson (1943), and Campbell (1963) consider
the opposite ends of a segment in the lineage, or chronocline, as two distinct
species [termed by Simpson (1961) as chronospeCles or successwnal speczes, and by
Mayr (1942) as allochronic species], whereas others refuse to recognize the two
ends as distinct species, no matter how long the sequence is nor how many
differences accumulate from one end of it to the other (very much analogous to
the way Goldschmidt views the geographic rassenkreis). According to the latter
view, speoatton can only result from a cladogenetic event (Eldredge and
Cracraft, 1980; Hull, 1978; Szalay and Bock, 1991; Wiley, 1981).

Conclusions

No matter which view (Goldschmidt's vs. Dobzhansky's) one accepts in

reaching a taxonomic decision, both the geographic and the temporal effects
must be taken into account when evaluating the fossil record. The presence in
the Middle East of a highly specialized Neanderthal such as Shanidar 5, for
example, can result from its late geologic age or from an extensive southward
geographic expansion of the H. neanderthalen.m territory. The biogeographic
model suggests that the combination of a late specimen and great territorial
expansion can theoretically result in a Neanderthal with an even more classical,
i.e., derived, appearance in a given cave. On the other hand, the presence in the
Amud cave of a more generalized (modern-looking) H. neanderthalensis spec-
imen, such as Amud I (Suzuki, 1970), can stem from the specimen's great chron-
ological age or from the cave's location on the periphery of the modestly ex-
panded Neanderthal territory. The combination ofthese two factors may, again,
introduce even more generalized specimens. Similarly, morphological dif-
ferences are expected to occur in the H. sapiens lineage. The difference between
the hominids from Zuttiyeh and Qafzeh, two caves that are geographically close,
can be attributed to the time difference between them.
It is, therefore, the specific geographic location of the Middle East that
introduces such morphological complexity into the caves. The model suggests
that at a given point in time, the picture of facial morphology is much less
complex at the extremities of the geographic range of the populations; and, as
we reach the ends of this spectrum, the situation is, indeed, simpler. In western
Europe, only Neanderthals were present almost to the time of their disap-
pearance, whereas in North Africa (for example, Djebel Irhoud and Haua-
fteah), early H. sapiens was the sole inhabitant during the same period. It was
apparently due to the fact that the hominids were unable to cross the Straits of
Gibraltar that the complete ring species was not achieved.
If it is true that no particular degree of morphological change is associated
with the genetic event of speciation (Lambert and Paterson, 1982), it follows that
most morphological changes succeed the actual speciation event itself. This is
534 BIOGEOGRAPHIC MODH. OF H NEANDERTHALENSIS AND H. SAPIENS
apparently the basis for the paleontological phenomenon of underestimating
the number of species on the basis of osteological characters: There may be more
speciation events than the morphological changes that record them (Tattersall,
1986). From this point of view of the fossil record, this implies that a particular
specimen need only reveal its species identity incipiently in order to be consid-
ered part of the taxon. Hence, even incipient Neanderthal morphology is suffi-
cient evidence to merit a specimen's inclusion in H. neanderthalenszs.* It follows,
therefore, that it is less excusable to attribute specimens with such morphology to
H. sapiens than to exclude more generalized specimens-those with few appar-
ent H. neanderthalenszs characters-from the hypodigm of H. neanderthalensis,
although phylogenetically the latter actually belong to this species.
In regard to the combination of features mentioned above (a derived pelvis
and primitive face in H. sapiens, and the opposite in H. neanderthalensis; see Fig.
3) the cladogenetic phylogeny predicts that those specimens on the H. nean-
derthalensis clade that show a more primitive face will still have a typically H.
neanderthalensis, i.e., primitive, pelvis. On the other hand, an anagenetic model,
in which such specimens are viewed as late, that is, "advanced" Neanderthals, on
the verge of becoming H. sapiens, predicts that they will exhibit a more derived
pelvis. It is difficult to reach solid conclusions based on the very fragmentary
evidence available. The Amud I specimen, which is often used as an example of
an "advanced" Neanderthal on the basis of cranial morphology, has little left of
its pelvis (En do and Kimura, 1970). Nevertheless, this fragment, as well as the
Shanidar 4 specimen (Trinkaus, 1983), show hallmarks of the diagnostic H.
neanderthalensis, or primitive, pelvis.
The sense of eternal stability conveyed by a cave and its fossil-bearing depos-
its should not mislead us into thinking that the physical and biotic realms were
likewise invariant across this time span. Although the Mousterian period was
short hom an evolutionary perspective, it was long enough to be characterized
by periodic fluctuations in niche size and shape. It is very likely, given the
geographic position of the Middle East, that such a process was responsible for
the distributional pattern of H. neanderthalenszs and H. sapiens during the upper
Pleistocene of this region.

ACKNOWLEDGMENTS

I would like to thank Dr. William Kimbel, Dr. Eric Delson, and two anony-
mous reviewers for their meticulous comments, which were of great help. Figs. 2
and 4 were skillfully drawn by Douglas Beckner. Finally, I thank the L. S. B.
Leakey Foundation for supporting fieldwork in Israel, especially the Kebara and
Amud projects.

*Referring again to the biogeographic model, we can suppose, therefore, that a taxonomist who is
attempting to draw the geographic boundary between H. neanderthalensts and H. saptens at a given
point in time will tend to place it wtthin the geographic range of H. neanderthalensis rather than at
the true (i.e., genetic) boundary between the two species.
 YOEL RAK 535
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