Since the first discovery of Pithecanthropus (Homo) erectus by E. Dubois at Trinil in 1891, over 200 hominid dentognathic remains have been collected from the Early to Middle Pleistocene deposits of Java, Indonesia, forming the largest palaeoanthropological collection in South East Asia. Most of these fossils are currently attributed to H. erectus. However, because of the substantial morphological and metric variation in the Indonesian assemblage, some robust specimens, such as the partial mandibles Sangiran 5 and Sangiran 6a, were formerly variably allocated to other taxa (Meganthropus palaeojavanicus, Pithecanthropus dubius, Pongo sp.). To resolve the taxonomic uncertainty surrounding these and other contentious Indonesian hominid specimens, we used occlusal fingerprint analysis (OFA) to reconstruct their chewing kinematics; we also used various morphometric approaches based on microtomography to examine the internal dental structures. Our results confirm the presence of Meganthropus as a Pleistocene Indonesian hominid distinct from Pongo, Gigantopithecus and Homo, and further reveal that Dubois’s H. erectus paratype molars from 1891 are not hominin (human lineage), but instead are more likely to belong to Meganthropus.

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We thank the Pusat Penelitian Arkeologi of Jakarta and the Balai Pelestarian Situs Manusia Purba of Sangiran, Java, and the Muséum national d’histoire naturelle. We thank the many curators and colleagues who granted access to fossil and recent hominid materials for scanning. We are grateful to D. Grimaud-Hervé, C. Hertler, F. Sémah and H. Widianto for their support. We thank J. Braga for sharing the microtomographic scans of South African fossil specimens. For scientific discussion, we thank P. Bayle, S. Benazzi, L. Bondioli, J. Braga, M.C. Dean, F. Détroit, Y. Hou, L. Mancini, B. Maureille, A. Mazurier, L. Puymerail, L. Rook, C. Tuniz and B. Wood. We would like to express our gratitude to C. Hemm, L. Hauser, M. Janocha and L. Strzelczyk for their help with the surface scanning and OFA analysis. Scanning of the Vietnamese specimens was funded by the Projet International de Coopération Scientifique-Centre national de la recherche scientifique (CNRS) grant to A.M.B (PICS 2011-2013 n°5712). Research was supported by the CNRS.

Author information


  1. Laboratoire PACEA, UMR 5199 CNRS, Université de Bordeaux, Pessac, France

    • Clément Zanolli
  2. Laboratoire AMIS, UMR 5288 CNRS, Université Toulouse III Paul Sabatier, Toulouse, France

    • Clément Zanolli
    •  & Jean Dumoncel
  3. Department of Palaeoanthropology, Senckenberg Research Institute and Natural History Museum Frankfurt, Frankfurt am Main, Germany

    • Ottmar Kullmer
    •  & Friedemann Schrenk
  4. Department of Paleobiology and Environment, Institute of Ecology, Evolution, and Diversity, Goethe University, Frankfurt, Germany

    • Ottmar Kullmer
    •  & Friedemann Schrenk
  5. Institute of Human Origins and School of Human Evolution and Social Change, Arizona State University, AZ, Tempe, USA

    • Jay Kelley
  6. Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, Washington, DC, USA

    • Jay Kelley
  7. Department of Human Evolutionary Biology, Harvard University, MA, Cambridge, USA

    • Jay Kelley
  8. Laboratoire BABEL, FRE 2029 CNRS, Université Paris Descartes, Faculté de chirurgie dentaire, Montrouge, France

    • Anne-Marie Bacon
  9. Lundbeck Foundation GeoGenetics Center, Copenhagen, Denmark

    • Fabrice Demeter
  10. UMR 7206 CNRS, Muséum national d’histoire naturelle, Paris, France

    • Fabrice Demeter
  11. Department of Anatomy and Developmental Biology, Monash University, Victoria, Melbourne, Australia

    • Luca Fiorenza
  12. Earth Sciences, University of New England, New South Wales, Armidale, Australia

    • Luca Fiorenza
  13. Department of Anthropology and Department of Anatomical Sciences, Stony Brook University, NY, Stony Brook, USA

    • Frederick E. Grine
  14. Department of Human Evolution, Max Planck Institute for Evolutionary Anthropology, Leipzig, Germany

    • Jean-Jacques Hublin
    •  & Matthew M. Skinner
  15. Anthropological and Palaeoenvironmental Department, The Institute of Archaeology, Hanoi, Vietnam

    • Anh Tuan Nguyen
    •  & Thi Mai Huong Nguyen
  16. Key Laboratory of Vertebrate Evolution and Human Origins, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Beijing, China

    • Lei Pan
  17. State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing, China

    • Lei Pan
  18. Heinz Maier-Leibnitz Center (FRM-II), Technische Universität München, Garching, Germany

    • Burkhard Schillinger
  19. School of Anthropology and Conservation, University of Kent, Canterbury, UK

    • Matthew M. Skinner
  20. Department of Paleoanthropology, Yunnan Institute of Cultural Relics and Archaeology, Kunming, China

    • Xueping Ji
  21. School of Resource Environment and Earth Science, Yunnan University, Kunming, China

    • Xueping Ji
  22. UMR 7194 CNRS, Muséum national d’histoire naturelle, Paris, France

    • Roberto Macchiarelli
  23. Unité de Formation Géosciences, Université de Poitiers, Poitiers, France

    • Roberto Macchiarelli


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The study was initiated by C.Z. during his PhD research project under the supervision of R.M. Microtomography-based data were collected and elaborated by C.Z., A.M.B., F.D., J.K., O.K., L.F., F.S., A.T.N., T.M.H.N., B.S., J.-J.H., M.M.S., X.J. and R.M. Quantitative data were compiled and analysed by C.Z., J.D., O.K., L.F., L.P., M.M.S., F.E.G. and R.M. C.Z, R.M, O.K. and J.K. wrote the manuscript with contributions from all the other authors.

Competing interests

The authors declare no competing interests.

Corresponding author

Correspondence to Clément Zanolli.

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