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Evidence for increased hominid diversity in the Early to Middle Pleistocene of Indonesia


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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Fig. 1: Virtual rendering of the Indonesian hominid teeth examined for taxonomic reassessment.
Fig. 2: OFA.
Fig. 3: Enamel thickness cartographies.
Fig. 4: Molar crown-root proportions.
Fig. 5: Geometric morphometric analyses of the EDJ and pulp chamber.
Fig. 6: Geometric morphometric analyses of the EDJ and pulp chamber in non-Homo hominids.

Data availability

The authors declare that all data supporting the findings of this study are available within the paper and its Supplementary information files.


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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.

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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.

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Correspondence to Clément Zanolli.

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Zanolli, C., Kullmer, O., Kelley, J. et al. Evidence for increased hominid diversity in the Early to Middle Pleistocene of Indonesia. Nat Ecol Evol 3, 755–764 (2019).

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