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New fossils from Jebel Irhoud, Morocco and the pan-African origin of Homo sapiens

A Correction to this article was published on 13 June 2018

This article has been updated

Abstract

Fossil evidence points to an African origin of Homo sapiens from a group called either H. heidelbergensis or H. rhodesiensis. However, the exact place and time of emergence of H. sapiens remain obscure because the fossil record is scarce and the chronological age of many key specimens remains uncertain. In particular, it is unclear whether the present day ‘modern’ morphology rapidly emerged approximately 200 thousand years ago (ka) among earlier representatives of H. sapiens1 or evolved gradually over the last 400 thousand years2. Here we report newly discovered human fossils from Jebel Irhoud, Morocco, and interpret the affinities of the hominins from this site with other archaic and recent human groups. We identified a mosaic of features including facial, mandibular and dental morphology that aligns the Jebel Irhoud material with early or recent anatomically modern humans and more primitive neurocranial and endocranial morphology. In combination with an age of 315 ± 34 thousand years (as determined by thermoluminescence dating)3, this evidence makes Jebel Irhoud the oldest and richest African Middle Stone Age hominin site that documents early stages of the H. sapiens clade in which key features of modern morphology were established. Furthermore, it shows that the evolutionary processes behind the emergence of H. sapiens involved the whole African continent.

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Figure 1: Facial reconstruction of Irhoud 10.
Figure 2: Irhoud 11 mandible (lateral and occlusal views).
Figure 3: Comparative shape analysis.

Change history

  • 13 June 2018

    In the originally published version of this Letter, the PC1 and PC2 values in Fig. 3a were incorrect; in addition, the enumeration of Qafzeh samples should have been 'Qafzeh (9, 25)', and Extended Data Fig. 1c and Extended Data Table 2 have been altered accordingly. This has been corrected online.

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Acknowledgements

The research program at Jebel Irhoud is jointly conducted and supported by the Moroccan Institut National des Sciences de l’Archéologie et du Patrimoine and the Department of Human Evolution of the Max Planck Institute for Evolutionary Anthropology. We are grateful to the many curators and colleagues who, over the years, gave us access to recent and fossil hominin specimens for computed tomography scanning or analysis, to E. Trinkaus for providing comparative data and to C. Kiarie, M. Lui, C. Piot, D. Plotzki, A. Buchenau and H. Temming for their technical assistance.

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Contributions

The study was conceived by J.-J.H., A.B.-N. and P.G. Cranial metrical and non-metrical data were compiled and analysed by J.-J.H., A.B.-N., S.E.F., S.N., K.H. and P.G. Mandibular metrical and non-metrical data were compiled and analysed by J.-J.H. and I.B. Dental metrical and non-metrical data were compiled and analysed by S.E.B., M.M.S., A.L.C. and S.B. J.-J.H. and P.G. wrote the manuscript with contributions from all other authors.

Corresponding authors

Correspondence to Jean-Jacques Hublin or Philipp Gunz.

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The authors declare no competing financial interests.

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Reviewer Information Nature thanks R. G. Klein, C. Stringer and the other anonymous reviewer(s) for their contribution to the peer review of this work.

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Extended data figures and tables

Extended Data Figure 1 Mandibular morphology.

a, Symphyseal section of the Irhoud 11 mandible showing the mental angle. b, Mental area of Irhoud 11 before virtual reconstruction (top) and Irhoud 3 (bottom). Both images are surface models generated from micro-computed tomography data. c, Bivariate plot of mandibular corpus breadth versus height at the mental foramen. Irhoud 11 (pink star) falls within the EMH distribution and has one of the largest corpus heights among Middle to Late Pleistocene hominins. Values are in mm. n indicates sample size. Data sources and sample compositions can be found in the Methods. Scale bar, 20 mm.

Extended Data Figure 2 Dental morphology.

a, Shape–space PCA plot of Late Early and Middle Pleistocene archaic Homo, Neanderthals and RMH M1 crown outlines. The deformed mean crown outlines in the four directions of the PCs are drawn at the extremity of each axis. Sample compositions and abbreviations can be found in the Methods. b, EDJ morphology of the M2 and P4. Top left, the PCA analysis of the EDJ shape of the M2 places Irhoud 11 intermediate between H. erectus and RMH (along with other north Africa fossil humans) and distinct from Neanderthals. Surface models illustrate EDJ shape changes along PC1 (bottom left) and PC2 (top right); the former separating H. erectus from RMH, Neanderthals and north African EMH and the latter separating Neanderthals from RMH and north African EMH. Bottom right, a PCA analysis of the EDJ shape of the P4 groups Irhoud 11 with modern and fossil humans.

Extended Data Figure 3 Shape analysis of I2 roots.

A between-group PCA shows a complete separation between Neanderthals and a worldwide sample of recent modern humans based on subtle shape differences. Irhoud 11 (pink star) plots at the fringes of RMH, close to the EMH from Contrebandiers 1 (Tem). Colour-coded Procrustes group mean shapes are plotted in the same orientation as the I2 root surface of Irhoud 11. Although Irhoud 11 is more similar, overall, to Neanderthals in terms of root size, its root shape is clearly modern. The H. erectus specimen KNM-WT 15000 and hypothetical EMH Tabun C2 have incisor root shapes similar to Neanderthals, suggesting that roots that are labially more convex than in RMH represent a conserved primitive condition with limited taxonomical value. Sample compositions and abbreviations can be found in the Methods.

Extended Data Figure 4 Shape analysis of the external vault.

a, PC scores of PC1 and PC2 of external braincase shape in H. erectus, archaic Middle Pleistocene Homo, geographically diverse RMH and Neanderthals. Results are consistent with the analysis of endocranial shape (Fig. 3a). However, several EMH and Upper Palaeolithic specimens fall outside the RMH variation. This is probably owing to the projecting supraorbital tori in these specimens. b, Shape changes associated with PC1 (two standard deviations in either direction) shown as thin-plate spline deformation grids in lateral and oblique view. PC1 captures a contrast between elongated braincases with projecting supraorbital tori (low scores, in black) and a more globular braincase with gracile supraorbital tori (high scores, in red). Sample compositions and abbreviations can be found in the Methods.

Extended Data Figure 5 Facial and endocranial shape differences among Homo groups.

Visualizations of GMM shape analyses in Fig. 3. a, Average endocranial shape differences between H. erectus, recent H. sapiens and Neanderthals. Thin-plate spline grids are exaggerated. b, Visualization of shape changes along PC1 in Fig. 3b in frontal, lateral and superior view; two standard deviations in either direction from the mean shape (grey, negative; black, positive). c, Shape changes along PC2. All recent and fossil modern humans (low scores along PC2) share smaller, orthognathic faces, that differ from the larger, robust and prognathic faces of the Middle Pleistocene humans and Neanderthals (high scores along PC1). Arrow length is colour-coded (short, blue; long, red). As these visualizations are affected by the Procrustes superimposition, we also show grids for the maxilla and the supraorbital area. The arrow points to the plane of the maxillary thin-plate spline (red) in the template configuration.

Extended Data Table 1 List of hominin specimens
Extended Data Table 2 Measurements of the Irhoud 11 mandible after reconstruction
Extended Data Table 3 Dental measurements (upper dentition)
Extended Data Table 4 Dental measurements (lower dentition)
Extended Data Table 5 Morphological dental trait comparison

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Hublin, JJ., Ben-Ncer, A., Bailey, S. et al. New fossils from Jebel Irhoud, Morocco and the pan-African origin of Homo sapiens. Nature 546, 289–292 (2017). https://doi.org/10.1038/nature22336

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