S12-9
Denisovans: How to Recognize Them in the Fossil Record
Clément Zanolli1, Anne-Marie Bacon2, Souliphane Boualaphane3a, Fabrice Demeter4, 5, Philippe Duringer6, Jean-Jacques Hublin7, 8, Renaud Joannes-Boyau9, 10, Ottmar Kullmer11a, 12, Thonglith Luangkhoth3b, Jean-Luc Ponche13, Friedemann Schrenk11b, 12, Laura Shackelford13, 14, Matthew M. Skinner8, 15, Daovee Sihanam3c, Bence Viola16, Kira Westaway17, Song Xing18
1Université de Bordeaux, CNRS, MCC, PACEA, UMR 5199, F-33600, France
2Université Paris Cité, CNRS, BABEL, F-75012, France
3Ministry of Information, Culture and Tourism, Laos
4Lundbeck Foundation GeoGenetics Centre, Globe Institute, University of Copenhagen, Denmark
5Muséum national d'Histoire naturelle, CNRS, Université de Paris, Musée de l'Homme, France
6Ecole et Observatoire des Sciences de la Terre, Institut de Physique du Globe de Strasbourg (IPGS), UMR 7516 CNRS, Université de Strasbourg, France
7Collège de France, France
8Department of Human Evolution, Max Planck Institute for Evolutionary Anthropology, Germany
9GARG, Southern Cross GeoScience, Southern Cross University, Australia
10Centre for Anthropological Research, University of Johannesburg, South Africa
11Division of Paleoanthropology, Senckenberg Research Institute and Natural History Museum, Germany
12Department of Paleobiology and Environment, Institute of Ecology, Evolution, and Diversity, Goethe University Frankfurt, Germany
13Université de Strasbourg, Laboratoire Image, Ville Environnement, UMR 7362 UdS CNRS, France
13Department of Anthropology, University of Illinois at Urbana-Champaign, U.S.A.
14Carle Illinois College of Medicine, University of Illinois at Urbana-Champaign, U.S.A.
15School of Anthropology and Conservation, University of Kent, United Kingdom
16Department of Anthropology, University of Toronto, Canada
17‘Traps’ Luminescence Dating Facility, School of Natural Sciences, Macquarie University, Australia
18Key Laboratory of Vertebrate Evolution and Human Origins, Institute of Vertebrate Paleontology and Paleoanthropology, CAS Center for Excellence in Life and Paleoenvironment, China
Denisovans were first identified in 2010, as a distinct extinct human taxon, based on mtDNA extracted from the distal finger phalanx Denisova 3 from the eponym cave in the Altai Mountains, in Siberia. Later, paleogenetic studies enabled the identification of other Denisovan specimens, such as the isolated teeth of Denisova 2, 4 and 8, and the bone fragments Denisova 11 and 13. Paleogenetics demonstrated that Denisovans and Neanderthals are sister taxa and might thus share morphological affinities. Recently, the Xiahe mandible from Baishiya Cave, on the Tibetan Plateau in China, and the isolated molar TNH2-1 from Cobra Cave, in Laos, were attributed to Denisovans based on a combination of geometric morphometric and paleoproteomic analyses. The currently identified Denisovan remains from Siberia, Tibet and Laos show a mixture of Neanderthal-like features and autapomorphies. The recent description of the Harbin cranium, showing a large endocranial volume associated with a mix of archaic and advanced features, and attributed to Homo longi, sparked a debate regarding its taxonomic attribution. We used here 3D geometric morphometrics (3D GM) to compare the enamel-dentine junction (EDJ) of Homo erectus, Neanderthals, Denisova 4, and modern humans with the Pleistocene specimen CA 673 from China, the holotype of the nomen dubium Hemanthropus peii. Results indicate that CA 673 shares features with Denisova 4 (i.e., internally-tilted dentine horns, well-developed hypocone, reduced central basin, as also frequently found in Neanderthals), indicating that it belongs to a Denisovan. Based on these results and in absence of molecular evidence, we make inferences regarding Asian dental and cranial specimens currently lumped as Middle Pleistocene Homo that exhibit Neanderthal-like traits. We reviewed the available evidence for the specimens from Penghu, Tongzi, Panxian Dadong, Xujiayao, Dali, Maba, Xuchang and Harbin and we propose that they could represent Denisovans.