Two fossilized human crania (Apidima 1 and Apidima 2) from Apidima Cave, southern Greece, were discovered in the late 1970s but have remained enigmatic owing to their incomplete nature, taphonomic distortion and lack of archaeological context and chronology. Here we virtually reconstruct both crania, provide detailed comparative descriptions and analyses, and date them using U-series radiometric methods. Apidima 2 dates to more than 170 thousand years ago and has a Neanderthal-like morphological pattern. By contrast, Apidima 1 dates to more than 210 thousand years ago and presents a mixture of modern human and primitive features. These results suggest that two late Middle Pleistocene human groups were present at this site—an early Homo sapiens population, followed by a Neanderthal population. Our findings support multiple dispersals of early modern humans out of Africa, and highlight the complex demographic processes that characterized Pleistocene human evolution and modern human presence in southeast Europe.
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The data that support the findings of this study are available from the corresponding authors upon reasonable request.
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This research was supported by the European Research Council (ERC CoG no. 724703) and the German Research Foundation (DFG FOR 2237). We thank all curators and their institutions for access to original specimens or casts used in this study; T. White, B. Asfaw, M. López-Soza, V. Tourloukis, D. Giusti, G. Konidaris, C. Fardelas and O. Stolis for their input and assistance; A. Balzeau (Muséum National d’Histoire Naturelle; MNHN), E. Delson (New York Consortium in Evolutionary Primatology; NYCEP), L. Leakey (africanfossils.org) for providing access to three-dimensional models of specimens used in our figures. C.S.’s research is supported by the Calleva Foundation and the Human Origins Research Fund. We are grateful to S. Benazzi, E. Delson and I. Hershkovitz for their comments and suggestions.
The authors declare no competing interests.
Publisher’s note: Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.
Peer review information Nature thanks Stefano Benazzi, Eric Delson and Israel Hershkovitz for their contribution to the peer review of this work.
Extended data figures and tables
a, The interior of Apidima Cave A, with the ‘skull breccia’ (red box) before its removal from the cave, shown wedged between the cave walls and near the ceiling. A cross-section of the Apidima 1 cranium can be seen in the bottom left corner of the red box. Note the bedded appearance of the breccia remnant (black dashed line) consisting of different clast sizes and distribution similar to those seen in the talus cone outside the cave in c. b, Cast of the ‘skull breccia’ in the early stages of preparation and cleaning. Apidima 1 is seen on the left, Apidima 2 on the right. c, View of the Apidima site from the sea. Images courtesy and copyright of the Museum of Anthropology, Medical School, National Kapodistrian University of Athens.
a, Posterior view. b, Superior view. c, Inferior view. Scale bar, 5 cm.
a–c, Images are the computed surface of the original fossil (a), all segmented fragments (b) and reconstruction 1 from an anterior-superior view (c); segmented fragments are shown in colour and mirrored fragments in grey.
Top row, reconstruction 1 (made by C.R., mirroring criterion). Second row, reconstruction (made by C.R., smoothness criterion). Third row, reconstruction 3 (made by A.B., mirroring criterion). Bottom row, reconstruction 4 (made by A.B., smoothness criterion). Scale bar, 3 cm.
a–d, Images are the computed surface of the original fossil (a), the cropped scan volume (b), the duplicated and mirrored scan volume (c) and the complete reconstruction (d).
Extended Data Fig. 6 Lateral and posterior views of the parietal region of the four manual reconstructions of Apidima 2.
Images as shown in Extended Data Fig. 4. a, Reconstruction 1 (made by C.R., mirroring criterion). b, Reconstruction 2 (made by C.R., smoothness criterion). c, Reconstruction 3 (made by A.B., mirroring criterion). d, Reconstruction 4 (made by A.B., smoothness criterion).
a, b, Posterior and lateral views of the Apidima 1 reconstruction. c, d, Posterior and lateral views of Apidima 2 reconstruction 1. e, f, Posterior and lateral views of the three-dimensional model of La Chapelle-aux-Saints (Neanderthal). g, h, Posterior and lateral views of the three-dimensional model of La Ferrassie 1 (Neanderthal). e–h, Images courtesy of A. Balzeau (MHNH). i, j, Posterior and lateral views of Elandsfontein (MPA). Images courtesy of C.S. k, l, Posterior and lateral views of the three-dimensional model of Sima de los Huesos Cranium 5, cast (MPE). Images courtesy of E. Delson (NYCEP). m, n, Posterior and lateral views of Skhul 5 (H. sapiens). Images courtesy of C.S. o, p, Posterior and lateral views of the three-dimensional model of Eliye Springs (MPA). Images reproduced with permission from https://africanfossils.org/. Scale bar, 5 cm.
Values calculated from the dataset used in analysis 4 (midsagittal posterior cranial profile), based on the axis between the mean Neanderthal and mean modern African shape. Apidima 1 and the remaining fossil samples are projected onto this axis. Violins extend from the minimum to the maximum value; boxes show the 25–75% quartiles and lines indicate the median. Samples as in Fig. 3b, symbols as in Fig. 2; modern Africans, green dots (n = 15).
Extended Data Fig. 9 Apidima 1 reconstruction superimposed manually with Apidima 2 reconstruction 1.
Aidima 1 is shown in yellow; Apidima 2 is shown in rainbow. a, Lateral view. b, Posterior view. c, Ventral view.