Geometric morphometrics and paleoproteomics enlighten the paleodiversity of Pongo.

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Authors
Paterson, Ryan 
Patramanis, Ioannis 
Barker, Graeme 
Demeter, Fabrice 
Abstract

Pleistocene Pongo teeth show substantial variation in size and morphology, fueling taxonomic debates about the paleodiversity of the genus. We investigated prominent features of the enamel-dentine-junction junction (EDJ)-phylogenetically informative internal structures-of 71 fossil Pongo lower molars from various sites by applying geometric morphometrics and conducted paleoproteomic analyses from enamel proteins to attempt to identify extinct orangutan species. Forty-three orangutan lower molars representing Pongo pygmaeus and Pongo abelii were included for comparison. The shape of the EDJ was analyzed by placing five landmarks on the tip of the main dentine horns, and 142 semilandmarks along the marginal ridges connecting the dentine horns. Paleoproteomic analyses were conducted on 15 teeth of Late Pleistocene Pongo using high-resolution tandem mass spectrometry. The geometric morphometric results show variations in EDJ shape regarding aspects of the height and position of the dentine horns and connecting ridges. Despite the issue of molar position and sample size, modern molars are distinguished from fossil counterparts by their elongated tooth outline and narrowly positioned dentine horns. Proteomic results show that neither a distinction of P. pygmaeus and P. abelii, nor a consistent allocation of fossil specimens to extant species is feasible. Based on the EDJ shape, the (late) Middle to Late Pleistocene Pongo samples from Vietnam share the same morphospace, supporting the previous allocation to P. devosi, although substantial overlap with Chinese fossils could also indicate close affinities with P. weidenreichi. The hypothesis that both species represent one chronospecies cannot be ruled out. Two fossil specimens, one from Tam Hay Marklot (Laos, Late Pleistocene), and another from Sangiran (Java, Early to Middle Pleistocene), along with some specimens within the Punung sample (Java), exhibit affinities with Pongo abelii. The Punung fossils might represent a mix of early Late Pleistocene and later specimens (terminal Pleistocene to Holocene) related to modern Pongo. The taxonomy and phylogeny of the complete Punung sample needs to be further investigated.

Description

Acknowledgements: We express our gratitude to the Werner Reimers Foundation, Bad Homburg (Germany) for making the Gustav Heinrich Ralph von Koenigswald collection available for scientific research as a permanent loan to the Senckenberg Research Institute and Natural History Museum Frankfurt. We thank Alessandro Urciuoli and Jonas Elsborg for their valuable comments on data analyses and interpretation. For access to comparative material, we sincerely thank P. Bayle, A. Mazurier, R. Macchiarelli, and Jean-Jacques Hublin from the Department of Human Evolution of the Max Planck Institute for Evolutionary Anthropology for the access to scans of P. abelii and P. pygmaeus according to the agreement signed with the Directors of the Museum fûr Naturkunde Berlin–Leibniz Institute for Evolution and Biodversity Science (J. Vogel and S. Junker), and the curators of the Phyletic Museum, Jena and the MorphoSource database.

Keywords
Animals, Pongo, Hominidae, Pongo abelii, Proteomics, Tooth, Molar, Pongo pygmaeus, Fossils
Journal Title
PLoS One
Conference Name
Journal ISSN
1932-6203
1932-6203
Volume Title
18
Publisher
Public Library of Science (PLoS)
Sponsorship
HORIZON EUROPE Marie Sklodowska-Curie Actions (861389)
HORIZON EUROPE Marie Sklodowska-Curie Actions (101021361)
CNRS (PICS n°5712)