A key event in human evolution is the expansion of modern humans of African origin across Eurasia between 60 and 40 thousand years (kyr) before present (bp), replacing all other forms of hominins1. Owing to the scarcity of human fossils from this period, these ancestors of all present-day non-African modern populations remain largely enigmatic. Here we describe a partial calvaria, recently discovered at Manot Cave (Western Galilee, Israel) and dated to 54.7 ± 5.5 kyr bp (arithmetic mean ± 2 standard deviations) by uranium–thorium dating, that sheds light on this crucial event. The overall shape and discrete morphological features of the Manot 1 calvaria demonstrate that this partial skull is unequivocally modern. It is similar in shape to recent African skulls as well as to European skulls from the Upper Palaeolithic period, but different from most other early anatomically modern humans in the Levant. This suggests that the Manot people could be closely related to the first modern humans who later successfully colonized Europe. Thus, the anatomical features used to support the ‘assimilation model’ in Europe might not have been inherited from European Neanderthals, but rather from earlier Levantine populations. Moreover, at present, Manot 1 is the only modern human specimen to provide evidence that during the Middle to Upper Palaeolithic interface, both modern humans and Neanderthals contemporaneously inhabited the southern Levant, close in time to the likely interbreeding event with Neanderthals2,3.

Access optionsAccess options

Rent or Buy article

Get time limited or full article access on ReadCube.


All prices are NET prices.


  1. 1.

    The earliest modern human colonization of Europe. Proc. Natl Acad. Sci. USA 109, 13471–13472 (2012)

  2. 2.

    , , , & The date of interbreeding between Neandertals and modern humans. PLoS Genet. 8, e1002947 (2012)

  3. 3.

    et al. A draft sequence of the Neandertal genome. Science 328, 710–722 (2010)

  4. 4.

    et al. The Upper Palaeolithic of Manot Cave, Western Galilee, Israel: the 2011–12 excavations. Antiquity 87, (2013)

  5. 5.

    et al. Thermoluminescence date for the Mousterian burial site of Es-Skhul, Mt. Carmel. J. Archaeol. Sci. 20, 169–174 (1993)

  6. 6.

    et al. Thermoluminescence dates for the Neanderthal burial site at Kebara in Israel. Nature 330, 159–160 (1987)

  7. 7.

    , & Hominid cranial remains from Upper Pleistocene deposits at Aduma, Middle Awash, Ethiopia. Am. J. Phys. Anthropol. 123, 1–10 (2004)

  8. 8.

    , & The assimilation model, modern human origins in Europe, and the extinction of Neandertals. Quat. Int. 137, 7–19 (2005)

  9. 9.

    & Is the suprainiac fossa a Neandertal autapomorphy? A complementary external and internal investigation. J. Hum. Evol. 58, 1–22 (2010)

  10. 10.

    in L’Homme de Néandertal 3: l’Anatomie 67–73 (ed. ) (E.R.A.U.L., 1988)

  11. 11.

    , , & On the phylogenetic position of the pre-Neandertal specimen from Reilingen, Germany. J. Hum. Evol. 34, 485–508 (1998)

  12. 12.

    , & Cioclovina (Romania): affinities of an early modern European. J. Hum. Evol. 53, 732–746 (2007)

  13. 13.

    A propos de restes inédits du gisement de La Quina (Charente): un trait méconnu des néandertaliens et des prénéandertaliens. Anthropologie 84, 81–88 (1980)

  14. 14.

    , , & Modern human cranial diversity in the Late Pleistocene of Africa and Eurasia: evidence from Nazlet Khater, Peştera cu Oase, and Hofmeyr. Am. J. Phys. Anthropol. 140, 347–358 (2009)

  15. 15.

    et al. The Later Stone Age calvaria from Iwo Eleru, Nigeria: morphology and chronology. PLoS ONE 6, e24024 (2011)

  16. 16.

    & Virtual Anthropology: A Guide to a New Interdisciplinary Field (Springer, 2011)

  17. 17.

    et al. Early modern human diversity suggests subdivided population structure and a complex out-of-Africa scenario. Proc. Natl Acad. Sci. USA 106, 6094–6098 (2009)

  18. 18.

    & Fossil evidence for the origin of Homo sapiens. Am. J. Phys. Anthropol. 143, 94–121 (2010)

  19. 19.

    et al. The expansion of mtDNA haplogroup L3 within and out of Africa. Mol. Biol. Evol. 29, 915–927 (2012)

  20. 20.

    , , & Mitochondrial genome variation and the origin of modern humans. Nature 408, 708–713 (2000)

  21. 21.

    , , , & Genetic and archaeological perspectives on the initial modern human colonization of southern Asia. Proc. Natl Acad. Sci. USA 110, 10699–10704 (2013)

  22. 22.

    et al. Genomic and cranial phenotype data support multiple modern human dispersals from Africa and a southern route into Asia. Proc. Natl Acad. Sci. USA 111, 7248–7253 (2014)

  23. 23.

    , , & Modern human ancestry at the peripheries: a test of the replacement theory. Science 291, 293–297 (2001)

  24. 24.

    et al. The early Upper Paleolithic human skeleton from the Abrigo do Lagar Velho (Portugal) and modern human emergence in Iberia. Proc. Natl Acad. Sci. USA 96, 7604–7609 (1999)

  25. 25.

    et al. Genetic history of an archaic hominin group from Denisova Cave in Siberia. Nature 468, 1053–1060 (2010)

  26. 26.

    et al. Genome sequence of a 45,000-year-old modern human from western Siberia. Nature 514, 445–450 (2014)

  27. 27.

    et al. U-series and ESR analyses of bones and teeth relating to the human burials from Skhul. J. Hum. Evol. 49, 316–334 (2005)

  28. 28.

    & Tabun revisited: revised ESR chronology and new ESR and U-series analyses of dental material from Tabun C1. J. Hum. Evol. 39, 601–612 (2000)

  29. 29.

    , , , & Sea-land oxygen isotopic relationships from planktonic foraminifera and speleothems in the Eastern Mediterranean region and their implication for paleorainfall during interglacial intervals. Geochim. Cosmochim. Acta 67, 3181–3199 (2003)

  30. 30.

    et al. The timing and spatiotemporal patterning of Neanderthal disappearance. Nature 512, 306–309 (2014)

  31. 31.

    , & Where are inion and endinion? Variations of the exo- and endocranial morphology of the occipital bone during hominin evolution. J. Hum. Evol. 61, 488–502 (2011)

  32. 32.

    et al. Ohalo II H2: A 19,000-year-old skeleton from a water-logged site at the Sea of Galilee, Israel. Am. J. Phys. Anthropol. 96, 215–234 (1995)

  33. 33.

    The People in the Land of Israel from the Epipaleolithic to the Present Times: A Study Based on their Skeleton Remains. PhD thesis, Tel Aviv Univ. (1973)

  34. 34.

    & The Amud Man and his Cave Site 123–206 (Univ. Tokyo, 1970)

  35. 35.

    How and why humans grow thin skulls: experimental evidence for systemic cortical robusticity. Am. J. Phys. Anthropol. 101, 217–236 (1996)

Download references


The excavation at Manot Cave was initiated and supported throughout the years by the late D. David, founder of the ‘Dan David Prize’, and his son A. David. The ongoing research is financially supported by the Dan David Foundation, the Israel Antiquities Authority, Case Western Reserve University, the Leakey Foundation, the Irene Levi Sala CARE Archaeological Foundation, the Keren Kayemet L’Israel and the Israel Science Foundation. Radiocarbon dating research has been supported by the National Science Foundation, the Exilarch’s Foundation and the Max Planck Society–Weizman Institute Joint Center for Integrative Archaeology and Anthropology. Geological research was supported by The Bertha and Louis Weinstein Research Fund. We thank other research members of the Manot team: J.-M. Tejero and S. Weiner. We thank A. Behar and L. Barda for their drawings, C. Amit for the photography of the skull, and V. Essman and Y. Shmidov for surveying and drafting the cave. We thank I. Mureinik for editorial assistance. Special thanks are due to the following students and scholars: L. Weissbrod, D. Stein, H. Cohen, B. Medlej, M. Feldman, O. Hay, T. Abulafia, L. Davis, N. Schneller-Pels, D. Yegorov, M. Ullman and G. Hertzelinger. Thanks are also due to the Maale Yossef Regional Council, the residents of modern Manot and N. Reuven. We are also grateful to the late S. Dorfman, U. Dahari, D. Barshad, E. Stern and J. Goldberg. We thank I. Gilead and O. Bar-Yosef for reading and commenting on a previous version of this paper. We are grateful to C. Stringer for comments and suggestions.

Author information

Author notes

    • Israel Hershkovitz
    •  & Ofer Marder

    These authors contributed equally to this work.

    • Viviane Slon

    Present address: Department of Evolutionary Genetics, Max-Planck-Institute for Evolutionary Anthropology, Deutscher Platz 6, D-04103, Leipzig, Germany.


  1. The Dan David Laboratory for the Search and Study of Modern Humans, Sackler Faculty of Medicine, Tel Aviv University, PO Box 39040, Tel Aviv 6997801, Israel

    • Israel Hershkovitz
    •  & Viviane Slon
  2. The Steinhardt Museum of Natural History and National Research Center, Tel Aviv University, PO Box 39040, Tel Aviv 6997801, Israel

    • Israel Hershkovitz
    • , Daniella Bar-Yosef Mayer
    •  & Hila May
  3. Archaeology Division, Ben-Gurion University of the Negev, PO Box 653, Beer-Sheva 8410501, Israel

    • Ofer Marder
  4. Geological Survey of Israel, 30 Malkhe Israel Street, Jerusalem 95501, Israel

    • Avner Ayalon
    • , Miryam Bar-Matthews
    •  & Gal Yasur
  5. Max Planck Society-Weizmann Institute Center for Integrative Archaeology and Anthropology, D-REAMS Radiocarbon Laboratory, Weizmann Institute of Science, Rehovot 76100, Israel

    • Elisabetta Boaretto
    • , Valentina Caracuta
    •  & Bridget Alex
  6. Department of Anthropology and Human Evolutionary Biology, Harvard University, 11 Divinity Avenue, Cambridge, Massachusetts 02138, USA

    • Bridget Alex
  7. Department of Geography, The Hebrew University of Jerusalem, Jerusalem 91905, Israel

    • Amos Frumkin
  8. Institute of Archaeology, The Hebrew University of Jerusalem, Mount Scopus, Jerusalem 91905, Israel

    • Mae Goder-Goldberger
  9. Department of Human Evolution, Max-Planck-Institute for Evolutionary Anthropology, Deutscher Platz 6, D-04103, Leipzig, Germany

    • Philipp Gunz
  10. Department of Anthropology, Columbia University, New York 10027, USA

    • Ralph L. Holloway
  11. Department of Anatomy, Case Western Reserve University, Cleveland, Ohio 44106, USA

    • Bruce Latimer
  12. Department of Orthodontics, Case Western Reserve University School of Dental Medicine, 10900 Euclid Avenue, Cleveland, Ohio 44106, USA

    • Bruce Latimer
    •  & Mark G. Hans
  13. 8 Dan Street, Modi’in 7173161, Israel

    • Ron Lavi
  14. Institute of Earth Sciences, The Hebrew University of Jerusalem, Edmond J. Safra Campus, Givat Ram, Jerusalem 91904, Israel

    • Alan Matthews
  15. Department of Archaeology, Simon Fraser University, 8888 University Drive, Burnaby, British Columbia V5A 1S6, Canada

    • Francesco Berna
  16. Zinman Institute of Archaeology, University of Haifa, Haifa 3498838, Israel

    • Guy Bar-Oz
    •  & Reuven Yeshurun
  17. Department of Anatomy and Anthropology, Sackler Faculty of Medicine, Tel Aviv University, PO Box 39040, Tel Aviv 6997801, Israel

    • Hila May
  18. Department of Anthropology, University of Vienna, Althanstrasse 12–14, A-1090 Vienna, Austria

    • Gerhard W. Weber
  19. The Core Facility for Micro-Computed Tomography, University of Vienna, Althanstrasse 12–14, A-1090, Vienna, Austria

    • Gerhard W. Weber
  20. Israel Antiquities Authority, PO Box 586, Jerusalem 91004, Israel

    • Omry Barzilai


  1. Search for Israel Hershkovitz in:

  2. Search for Ofer Marder in:

  3. Search for Avner Ayalon in:

  4. Search for Miryam Bar-Matthews in:

  5. Search for Gal Yasur in:

  6. Search for Elisabetta Boaretto in:

  7. Search for Valentina Caracuta in:

  8. Search for Bridget Alex in:

  9. Search for Amos Frumkin in:

  10. Search for Mae Goder-Goldberger in:

  11. Search for Philipp Gunz in:

  12. Search for Ralph L. Holloway in:

  13. Search for Bruce Latimer in:

  14. Search for Ron Lavi in:

  15. Search for Alan Matthews in:

  16. Search for Viviane Slon in:

  17. Search for Daniella Bar-Yosef Mayer in:

  18. Search for Francesco Berna in:

  19. Search for Guy Bar-Oz in:

  20. Search for Reuven Yeshurun in:

  21. Search for Hila May in:

  22. Search for Mark G. Hans in:

  23. Search for Gerhard W. Weber in:

  24. Search for Omry Barzilai in:


I.H., O.B. and O.M. are directing the Manot cave research project. I.H., P.G., B.L., V.S., G.W.W., H.M., M.G.H. and R.L.H. performed the various aspects of the anthropological study of the Manot 1 calvaria. O.B., O.M., R.L. and M.G.G. conducted the archaeological studies at the cave. A.A., M.B.-M., G.Y. and A.M. conducted the U–Th dating of the calcitic crust on the Manot 1 calvaria and of speleothems in the cave. A.F. and F.B. conducted the geological study of the cave. E.B., V.C. and B.A. performed the radiocarbon dating and charcoal analysis. G.B.-O., R.Y. and D.B.-Y.M. conducted the study of the faunal remains.

Competing interests

The authors declare no competing financial interests.

Corresponding author

Correspondence to Israel Hershkovitz.

Extended data

Supplementary information

PDF files

  1. 1.

    Supplementary Information

    This file contains Supplementary Text and Supplementary References – see Contents for details.

Excel files

  1. 1.

    Supplementary Table 1

    This file contains detailed age determination results of flowstones, stalagmites and calcitic crusts.

About this article

Publication history






Further reading


By submitting a comment you agree to abide by our Terms and Community Guidelines. If you find something abusive or that does not comply with our terms or guidelines please flag it as inappropriate.