Neanderthal cranial ontogeny and its implications for late hominid diversity

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Homo neanderthalensis has a unique combination of craniofacial features that are distinct from fossil and extant ‘anatomically modern’ Homo sapiens (modern humans). Morphological evidence, direct isotopic dates1 and fossil mitochondrial DNA from three Neanderthals2,3 indicate that the Neanderthals were a separate evolutionary lineage for at least 500,000 yr. However, it is unknown when and how Neanderthal craniofacial autapomorphies (unique, derived characters) emerged during ontogeny. Here we use computerized fossil reconstruction4 and geometric morphometrics5,6 to show that characteristic differences in cranial and mandibular shape between Neanderthals and modern humans arose very early during development, possibly prenatally, and were maintained throughout postnatal ontogeny. Postnatal differences in cranial ontogeny between the two taxa are characterized primarily by heterochronic modifications of a common spatial pattern of development. Evidence for early ontogenetic divergence together with evolutionary stasis of taxon-specific patterns of ontogeny is consistent with separation of Neanderthals and modern humans at the species level.

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Figure 1: Shape variability in an ontogenetic series of Neanderthals (filled circles; see Methods for specimen labels) and modern humans (open circles/diamonds indicate extant/fossil specimens, respectively) for craniomandibular (a), cranial (b) and mandibular (c) landmark configurations.
Figure 2: Correlations between shape (w1), centroid size S and dental age (in postnatal years) for craniomandibular morphologies of Neanderthals (filled circles) and extant/fossil modern humans (open circles/diamonds, respectively).
Figure 3: Patterns of shape change during postnatal development in Neanderthals and modern humans.
Figure 4: Shape differences between Neanderthal and modern human skulls (ac) and mandibles (df).


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We are grateful to R. D. Martin and P. Stucki for support of our research. C. B. Stringer's continuous help and advice is gratefully acknowledged. We thank C. Howell, J.-J. Jaeger, D. Lieberman, J. Schwartz and C. Pryce for valuable comments on an earlier version of this paper. We appreciate the help of the following curators in providing access to fossil specimens: J.-J. Cleyet-Merle, J.-M. Cordy, V. Kharitonov, M. Marinot, F. Menghin, R. Orban, I. Pap, Y. Rak, C. B. Stringer, B. Vandermeersch. We also thank the radiologists, physicists and technicians engaged in fossil computer tomography scanning: E. Berenyi, G. Bijl, P. Dondelinger, V. Dousset, W. Fuchs, S. Louryan, V. Makarenko, U. Shreter, N. Strickland and C. L. Zollikofer. This work was supported by the Swiss National Science Foundation and a habilitation grant of the Canton of Zurich.

Author information

Correspondence to Christoph P. E. Zollikofer.

Supplementary information

Tables 1 to 3

Table 1: Description of the Neanderthal and the "anatomically modern" Homo sapiens (AMH) sample. (XLS 21 kb)

Table 2: Definitions of the cranial and mandibular landmarks.

Table 3: Statistics of relative warp analyses of the cranio-mandibular, cranial and mandibular landmark configurations.

The animation shows shape transformation of an average Neanderthal skull into an average AMH skull while size is held constant. (MOV 587 kb)

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