The appearance of anatomically modern humans in Europe and the nature of the transition from the Middle to Upper Palaeolithic are matters of intense debate. Most researchers accept that before the arrival of anatomically modern humans, Neanderthals had adopted several ‘transitional’ technocomplexes. Two of these, the Uluzzian of southern Europe and the Châtelperronian of western Europe, are key to current interpretations regarding the timing of arrival of anatomically modern humans in the region and their potential interaction with Neanderthal populations. They are also central to current debates regarding the cognitive abilities of Neanderthals and the reasons behind their extinction1,2,3,4,5,6. However, the actual fossil evidence associated with these assemblages is scant and fragmentary7,8,9,10, and recent work has questioned the attribution of the Châtelperronian to Neanderthals on the basis of taphonomic mixing and lithic analysis11,12. Here we reanalyse the deciduous molars from the Grotta del Cavallo (southern Italy), associated with the Uluzzian and originally classified as Neanderthal13,14. Using two independent morphometric methods based on microtomographic data, we show that the Cavallo specimens can be attributed to anatomically modern humans. The secure context of the teeth provides crucial evidence that the makers of the Uluzzian technocomplex were therefore not Neanderthals. In addition, new chronometric data for the Uluzzian layers of Grotta del Cavallo obtained from associated shell beads and included within a Bayesian age model show that the teeth must date to 45,000–43,000 calendar years before present. The Cavallo human remains are therefore the oldest known European anatomically modern humans, confirming a rapid dispersal of modern humans across the continent before the Aurignacian and the disappearance of Neanderthals.

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  1. 1.

    , , , & Neanderthal acculturation in Western Europe? A critical review of the evidence and its interpretation. Curr. Anthropol. 39, 1–44 (1998)

  2. 2.

    , & Radiocarbon dating of interstratified Neanderthal and early modern human occupations at the Châtelperronian type-site. Nature 438, 51–56 (2005)

  3. 3.

    , & Confirmation of Neanderthal/modern human interstratification at the Châtelperronian type-site. Proc. Natl Acad. Sci. USA 104, 3657–3662 (2006)

  4. 4.

    et al. Analysis of Aurignacian interstratification at the Châtelperronian-type site and implications for the behavioral modernity of Neandertals. Proc. Natl Acad. Sci. USA 103, 12643–12648 (2006)

  5. 5.

    , & in La lunga storia di Neandertal. Biologia e comportamento (eds & ) (Jaca Book, 2009)

  6. 6.

    , & Uluzzian bone technology and its implications for the origin of behavioural modernity. Quat. Int.. 10.1016/j.quaint.2011.03.039 (in the press)

  7. 7.

    & Découverte de restes humains dans un niveau castelperronien à Saint-Césaire (Charente-Maritime). CR Acad. Sci. Paris 291, 187–189 (1980)

  8. 8.

    , , , & A late Neanderthal associated with Upper Palaeolithic artefacts. Nature 381, 224–226 (1996)

  9. 9.

    , & First Neanderthal remains from Greece: the evidence from Lakonis. J. Hum. Evol. 45, 465–473 (2003)

  10. 10.

    & Dental remains from the Grotte du Renne at Arcy-sur-Cure (Yonne). J. Hum. Evol. 50, 485–508 (2006)

  11. 11.

    & Who were the makers of the Châtelperronian culture? J. Hum. Evol. 59, 586–593 (2010)

  12. 12.

    et al. Chronology of the Grotte du Renne (France) and implications for the context of ornaments and human remains within the Châtelperronian. Proc. Natl Acad. Sci. USA 107, 20234–20239 (2010)

  13. 13.

    & Quatre dents humaines paléolithiques trouvées dans des cavernes de l’Italie Méridionale. L’Anthropologie 71, 249–262 (1967)

  14. 14.

    & Makers of the early Aurignacian of Europe. Am. J. Phys. Anthropol. 113 (Suppl. 31). 61–115 (2000)

  15. 15.

    L’Uluzzien: faciès italien du leptolithique archaique. L’Anthropologie 93, 783–812 (1989)

  16. 16.

    in Sourcebook of Paleolithic Transitions (eds & ) (Springer, 2009)

  17. 17.

    & Contemporaneità di paleantropi e fanerantropi sulle coste dell'talia meridionale. Zephyrus 26–27, 7–30 (1976)

  18. 18.

    et al. Comparison of dental measurement systems for taxonomic assignment of first molars. Am. J. Phys. Anthropol. 144, 342–354 (2011)

  19. 19.

    et al. Dental tissue proportions and enamel thickness in Neandertal and modern human molars. J. Hum. Evol. 55, 12–23 (2008)

  20. 20.

    Patterns of molar wear in hunter-gatherers and agriculturists. Am. J. Phys. Anthropol. 63, 39–56 (1984)

  21. 21.

    et al. IntCal09 and Marine09 radiocarbon age calibration curves, 0–50,000 years cal BP. Radiocarbon 51, 1111–1150 (2009)

  22. 22.

    & Middle-Upper Palaeolithic transition in Southern Italy: Uluzzian macromammals from Grotta del Cavallo (Apulia). Quat. Int.. 10.1016/j.quaint.2011.03.028 (in the press)

  23. 23.

    et al. The role of climate in the spread of modern humans into Europe. Quat. Sci. Rev. 30, 273–279 (2011)

  24. 24.

    European Middle and Upper Palaeolithic radiocarbon dates are often older than they look: problems with previous dates and some remedies. Antiquity 85, 235–249 (2011)

  25. 25.

    , , & Revised age of late Neanderthal occupation and the end of the Middle Paleolithic in the northern Caucasus. Proc. Natl Acad. Sci. USA 108, 8611–8616 (2011)

  26. 26.

    in Continuity and Discontinuity in the Peopling of Europe (eds & ) (Springer, 2011)

  27. 27.

    The impossible coincidence: a single species model for the origins of modern human behavior in Europe. Evol. Anthropol. 14, 12–27 (2005)

  28. 28.

    Significance of enamel thickness in hominoid evolution. Nature 314, 260–263 (1985)

  29. 29.

    . R: a language and environment for statistical computing. 〈〉 (2008)

  30. 30.

    & Extensions of the Procrustes method for the optimal superimposition of landmarks. Syst. Zool. 39, 40–59 (1990)

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We thank the Soprintendenza per i Beni Archeologici della Puglia which facilitated the excavation of Grotta del Cavallo over the years. We also thank M. A. Gorgoglione who supported and helped in the collection of samples for 14C dating and encouraged the collaboration with the University of Siena for the study of the archaeological remains. P. Boscato, H. Klempererova, F. Ranaldo and S. Ricci have all helped in aspects of the research and are especially thanked. We are grateful to G. Gruppioni for providing the Italian modern human sample used in this work. We thank M. Francken, B. Trautmann, I. Trautmann, H. Scherf, M. Dockner and R. Ginner for technical assistance. We thank F. L. Bookstein for suggestions on statistics. Access to the fossil specimens was made possible by the Croatian National History Museum, the French Musée National de Préhistoire, the French Muséum National d’Histoire Naturelle, Paleoanthropology, Eberhard Karls Universität Tübingen and the NESPOS Database 2011 (https://www.nespos.org/display/openspace/Home). We acknowledge the Centre de Microtomographie (Université de Poitiers), the Vienna micro-CT Laboratory (University of Vienna), VISCOM AG Hannover, the Paleoanthropology High Resolution Computing Tomography Laboratory (Eberhard Karls Universität Tübingen), the European Synchrotron Radiation Facility beamline ID17, the AST-RX platform (French Muséum National d’Histoire Naturelle) and the Oxford Radiocarbon Accelerator Unit (ORAU). The authors would like to thank T. Higham and R. E. M. Hedges for their input in the radiocarbon dating part of the project, for important comments and proofreading this manuscript. The radiocarbon dating was funded by the Natural Environment Research Council (NERC) NRCF programme. K.D. is part of the Ancient Human Occupation of Britain project, funded by the Leverhulme Trust. This work was supported by the NSF 01-120 Hominid Grant 2007, A.E.R.S. Dental Medicine Organisations GmbH FA547013, the Fondation Fyssen, the DFG INST 37/706-1 FUGG and the NERC Grant (NE/D014077/1).

Author information


  1. Department of Anthropology, University of Vienna, Althanstrasse 14, 1090 Vienna, Austria

    • Stefano Benazzi
    • , Cinzia Fornai
    •  & Gerhard W. Weber
  2. Oxford Radiocarbon Accelerator Unit, Research Laboratory for Archaeology and the History of Art, University of Oxford, Dyson Perrins Building, South Parks Road, Oxford OX1 3QY, UK

    • Katerina Douka
  3. Paleoanthropology, Department of Early Prehistory and Quaternary Ecology, Eberhard Karls Universität Tübingen, Rümelnstrasse 23, Tübingen 72070, Germany

    • Catherine C. Bauer
    •  & Katerina Harvati
  4. Department of Paleoanthropology and Messel Research, Senckenberg Research Institute Frankfurt, Senckenberganlage 25, D-60325 Frankfurt, Germany

    • Ottmar Kullmer
  5. Institute of Archaeology, Academy of Sciences of the Czech Republic, Královopolská 147, 612 00 Brno, Czech Republic

    • Jiří Svoboda
  6. Department of Anthropology, Faculty of Science, Masaryk University, Vinarˇská 5, 603 00 Brno, Czech Republic

    • Jiří Svoboda
  7. Department of Anthropology, Hungarian Natural History Museum, Ludovika tér 2-6, 1083 Budapest, Hungary

    • Ildikó Pap
  8. Department of Biology, University of Pisa, Via S. Maria 53, 56126 Pisa, Italy

    • Francesco Mallegni
  9. UMR 5199 PACEA, Université Bordeaux 1, avenue des Facultés, 33405 Talence, France

    • Priscilla Bayle
  10. Paleoanthropology group, Department of Paleobiology Museo Nacional de Ciencias Naturales (MNCM-CSIC), C/ José Gutiérrez Abascal 2, 28006 Madrid, Spain

    • Michael Coquerelle
  11. UMR 6578 CNRS/Aix Marseille/EFS, Laboratoire d’Anthropologie Bioculturelle, Faculté de Médecine/Secteur Nord, CS80011 Bd Pierre Dramard 13344, Marseille Cedex 15, France

    • Silvana Condemi
  12. Department of Environmental Sciences “G. Sarfatti”, U.R. Prehistoric Ecology, University of Siena, via T. Pendola 62, 53100 Siena, Italy

    • Annamaria Ronchitelli
  13. Senckenberg Center for Human Evolution and Paleoecology, Eberhard Karls Universität Tübingen, Rümelnstrasse 23, Tübingen 72070, Germany

    • Katerina Harvati


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S.B., F.M., K.H. and G.W.W. initiated and organized the project. S.B., C.C.B., P.B., J.S., I.P., K.H. and G.W.W. collected the fossils and modern human sample. S.B. and C.F. carried out the dental measurements. S.B. and M.C. analysed the data. K.D. initiated and performed the radiocarbon dating project. S.B., K.D., C.F., O.K., M.C., S.C., A.R., K.H. and G.W.W. discussed the results. S.B., K.D., C.F., M.C., S.C., A.R., K.H. and G.W.W. wrote and edited the manuscript.

Competing interests

The authors declare no competing financial interests.

Corresponding author

Correspondence to Stefano Benazzi.

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    The file contains Supplementary Methods and Results, Supplementary Figures 1-5 with legends, Supplementary Tables 1-6 and additional references.

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