Proc. Natl. Acad. Sci.

USA
Vol. 96, pp. 7117–7119, June 1999

Commentary

Hominids and hybrids: The place of Neanderthals in human evolution

Ian Tattersall*† and Jeffrey H. Schwartz‡
*Department of Anthropology, American Museum of Natural History, New York, NY 10024; and ‡Department of Anthropology, University of Pittsburgh,
Pittsburgh, PA 15260

As the first extinct human relatives to have become known to Neanderthal apomorphy (anatomical uniqueness), it is unsur-
science, the Neanderthals have assumed an almost iconic prising that the remarkable recent sequencing of a short stretch
significance in human evolutionary studies: a significance that of mtDNA isolated from the Feldhofer individual revealed this
has, of course, been greatly enhanced by the very substantial specimen to be a distant outlier when compared with all
fossil and behavioral record that has accumulated since the modern human populations (11).
original Feldhofer Cave skullcap and partial skeleton were The Neanderthals were highly successful over a large region
accidentally uncovered, on a pre-Darwinian August day in for a substantial period of time, but this situation changed
1856, by lime miners working in Germany’s Neander Valley dramatically with the arrival in Europe of the first modern
(1–3). Yet even now, 14 long decades later, paleoanthropo- humans, Homo sapiens. Indications are that these ‘‘Cro-
logical attitudes toward the Neanderthals remain profoundly Magnons’’ had begun to arrive both in eastern Europe (12) and
equivocal. Thus, although many students of human evolution in the far northeast of the Iberian Peninsula (13) by '40 kyr
have lately begun to look favorably on the view that these ago; and within little more than 10 kyr, the Neanderthals were
distinctive hominids merit species recognition in their own gone. The mechanism of their eviction has long been debated,
right as Homo neanderthalensis (e.g., refs. 4 and 5), at least as but there are four main possibilities (14). The first and second
many still regard them as no more than a strange variant of our of these, that the Neanderthals were eliminated by the
own species, Homo sapiens (6, 7). This difference represents far moderns in direct conflict or by indirect economic competi-
more than a simple matter of taxonomic hair-splitting. For, as tion, both imply the separate species status of the former, as
members of a distinct species, of a completely individuated does any combination of the two. The alternatives, that the
historical entity, the Neanderthals demand that we analyze and Neanderthals had simply evolved rapidly into moderns or that
understand them on their own terms. In contrast, if we see the genes of the invading moderns simply ‘‘swamped’’ those of
them as mere subspecific variants of ourselves, we are almost the Neanderthals, both imply some form of species continuity.
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obliged to dismiss the Neanderthals as little more than an Claims for evidence of ‘‘transition’’ between Neanderthals
evolutionary epiphenomenon, a minor and ephemeral append- and moderns, based on supposedly ‘‘intermediate’’ fossils
age to the history of Homo sapiens. dating from a short window of time around 40–30 kyr ago (15),
Any new information bearing on this matter is therefore have been refuted by the recognition that the fossils concerned
extremely welcome, and there is no doubt that the claims are either typically Neanderthal or modern (10) and, in one
advanced in this issue of the Proceedings by Duarte et al. (8) will significant case, had been misdated (16). Supporters of the
be closely scrutinized by their colleagues. Briefly, Duarte et al. continuity argument have thus tended lately to the view that
propose that the skeleton of a 4-year-old child, recently the disappearance of Neanderthal morphology was due to
unearthed at the 24,500-year-old (24.5 kyr-old) site of Lagar extensive interbreeding between the Neanderthals and the
Velho in Portugal, represents not merely a casual result of a incoming Cro-Magnons, who invaded in sufficient numbers to
Neanderthal/modern human mating, but rather is the product dominate the hybrid gene pool and thus the resulting pheno-
of several millennia of hybridization among members of the types (7, 17). The problem has been, though, that nobody has
resident Neanderthal population and the invading Homo sa- had any idea what a Neanderthal/modern hybrid might look
piens. Species (especially extinct ones) are often tricky to like in theory, and few have dared to suggest in practice that
identify in practice, and speciation, the process (or more any particular known fossil represents such a hybrid. The
probably, assortment of processes) by which new species come Duarte et al. claim for the Lagar Velho skeleton is the closest
about, is poorly understood. But by anyone’s reckoning, long- anyone has recently come to such a contention, hence the
term hybridization of this kind would indicate that the two intense interest that it seems sure to arouse.
populations belonged to the same species. So, if Duarte et al. The potential significance of the Lagar Velho claim is
are right, the case is closed: Neanderthals were indeed no more enhanced by the burial’s Iberian location, because it seems that
than an odd form of Homo sapiens. But is this claim reasonable it was in this peninsular extension of Europe that the Nean-
on the basis of the evidence presented? To clarify this, some derthals lingered longest. Outside Iberia, the latest Neander-
background follows. thals, and survivals of their ‘‘Mousterian’’ culture, are signif-
‘‘The Neanderthals’’ is the informal designation of a mor- icantly more than 30 kyr old. At the southern Spanish site of
phologically distinctive group of large-brained hominids who Zafarraya, however, the Mousterian may have lasted to '27
inhabited Europe and western Asia between '200 and less kyr ago (18) and is associated with typical Neanderthal remains
than 30 kyr ago (1, 2). They are sharply distinguished from at probably not much more than 30 kyr ago. Even more telling,
modern humans by a wide range of cranial and postcranial isolated but reasonably diagnostic fossil teeth suggest that
characters (1–2, 4, 9–10), although they do share a number of Neanderthals were living at the Portuguese cave sites of
derived bony features with other members of the endemic Salemas, Columbeira, and Figueira Brava at '29–30 kyr ago
European/western Asian hominid clade that diversified in this (19, 20). For whatever reasons, the Neanderthals’ last redoubt
part of the world after '500 kyr ago (10). Subsequent to '150 thus seems to have been in Iberia south and west of the Ebro.
kyr ago, the Neanderthals appear to have been the sole It is relevant here that while in certain other regions, Nean-
surviving species of this clade. Given the strong degree of
The companion to this Commentary begins on page 7604.
†To whom reprint requests should be addressed. e-mail: iant@amnh.
PNAS is available online at www.pnas.org. org.

7117
 7118 Commentary: Tattersall and Schwartz Proc. Natl. Acad. Sci. USA 96 (1999)

FIG. 2. Mandible of 3–4 year old Neanderthal from Roc de Marsal,

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France (Museé National de Prehistoire, Les Eyzies). (Upper) Oblique

view of essentially featureless front of mandible, which is vertical yet
also broad and somewhat arcuate from side to side. (Lower) Inferior
view illustrating how uniformly thinner the symphyseal region is
buccolingually compared to the bone farther back along the corpus.
Not to scale.

would be implausible even in principle. The expected distri-

bution of traits in hybrids that Duarte et al. discuss is that to
FIG. 1. Mandible of recent 2–3 year old Homo sapiens (American be found in F1 or F2 hybrids, not 200 generations down the line.
Museum of Natural History, uncatalogued) illustrating the essential Still, the claim of mixed ancestry for the Lagar Velho child
features of the symphyseal region. (Upper) Anterior view showing the ultimately rests on the morphology of the specimen. How does
central keel, which broadens at the mental tuberosity, fans out
this hold up? Duarte et al. begin with the skull, represented by
inferiorly, and terminates bilaterally in blunt “corners.” (Lower)
Inferior view showing the marked thickness of the mandible in the a temporal fragment and a partial lower jaw. In describing the
symphyseal region compared to the corpus behind it and the contri- temporal bone, they remark that the mastoid and juxtamastoid
bution of the marginal attributes of the mental tuberosity to mandib- eminences project basally to an approximately equal extent—a
ular shape. Not to scale. characteristic they claim to be intermediate between Nean-
derthals and moderns. However, this is a variable feature
derthals of the 36–33 kyr period appear to have acquired some among Neanderthals (10), and, more importantly, the individ-
of the Cro-Magnons’ behavioral attributes by acculturation ual was 4 years old and thus was developmentally only at the
(21, 22), there is no evidence for this beyond the ‘‘Ebro line,’’ point at which the mastoid process begins to expand signifi-
where—as in most other places—abrupt cultural replacement cantly downward among modern humans (23). Almost cer-
appears to have been the rule. tainly, as an adult this individual would have shown the typical
Nonetheless, if a claim is to be made that any 24.5-kyr-old modern conformation of the region, with a projecting mastoid
fossil individual represents a Neanderthal/modern hybrid, the process. There appears to have been no horizontal suture
obvious place to try it is in Iberia, and especially in Portugal, running beyond the parietal notch, as is found even in young
where the time gap between this individual and the last Neanderthals. And, regrettably, no information is provided on
plausible occurrence of Neanderthals may be as little as 2–3 the internal aspect of the petrosal, which would be highly
millennia. In the larger scheme of things this is hardly an eon, informative.
but it is still probably around 200 generations—a long time in Although damaged, the mandible conforms anteriorly pre-
genetics. This is presumably why Duarte et al. refrain from cisely to the highly characteristic ‘‘inverted-T’’ conformation
identifying the Lagar Velho child as a 50:50 Neanderthal/ we have recently described for Homo sapiens (24), and the
modern hybrid; but by claiming that it is instead the product relative thickness of the mandible across the symphysis in
of a population that had been hybridizing for many centuries, inferior view is also typical for our species (24, 25), as are the
they pose problems for themselves in analyzing it in terms of mental fossae (compare Figs. 1 and 2). The angulation of the
Neanderthal vs. Cro-Magnon traits. After so many genera- subincisal region is variable among both Neanderthals and
tions, genetic introgression would necessarily have proceeded moderns and does not argue here for Neanderthal influence.
so far that dichotomous characterization of phenotypic traits As described, the mandible lacks any sign of the internal
 Commentary: Tattersall and Schwartz Proc. Natl. Acad. Sci. USA 96 (1999) 7119

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