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(Cetartiodactyla : Suidae) Au Plio-Pléistocène En Afrique Orientale THESE Pour l'obtention du grade de DOCTEUR DE L'UNIVERSITÉ DE POITIERS UFR des sciences fondamentales et appliquées Institut de paléoprimatologie, paléontologie humaine : évolution et paléoenvironnements - IPHEP (Diplôme National - Arrêté du 7 août 2006) École doctorale : Sciences pour l'environnement - Gay Lussac Secteur de recherche : Paléontologie des vertébrés Présentée par : Antoine Souron Histoire évolutive du genre Kolpochoerus (Cetartiodactyla : Suidae) au Plio-Pléistocène en Afrique orientale Directeur(s) de Thèse : Jean-Renaud Boisserie, Timothy D. White Soutenue le 11 décembre 2012 devant le jury Jury : Président Michel Brunet Professeur des Universités - Collège de France Rapporteur Denis Geraads Directeur de recherche CNRS - Paris Rapporteur John M. Harris Professor - Natural history museum of Los Angeles Membre Jean-Renaud Boisserie Chargé de recherche CNRS - Université de Poitiers Membre Timothy D. White Professor - University of California, Berkeley Membre Patrick Vignaud Professeur des Universités - Université de Poitiers Membre Stéphane Ducrocq Directeur de recherche CNRS - Université de Poitiers Membre Sevket Sen Directeur de recherche - Muséum national d'histoire naturelle de Paris Pour citer cette thèse : Antoine Souron. Histoire évolutive du genre Kolpochoerus (Cetartiodactyla : Suidae) au Plio-Pléistocène en Afrique orientale [En ligne]. Thèse Paléontologie des vertébrés. Poitiers : Université de Poitiers, 2012. Disponible sur l'Intranet de l'Université de Poitiers <http://theses.univ-poitiers.fr> THESE Pour l’obtention du Grade de DOCTEUR DE L’UNIVERSITE DE POITIERS (Faculté des Sciences Fondamentales et Appliquées) (Diplôme National – Arrêté du 7 août 2006) Ecole Doctorale : Sciences pour l’Environnement Gay Lussac Secteur de Recherche : Terre Solide et Enveloppes Superficielles Discipline : Paléontologie des vertébrés Présentée par Antoine SOURON Histoire évolutive du genre Kolpochoerus (Cetartiodactyla : Suidae) au Plio-Pléistocène en Afrique orientale Sous la direction de Jean-Renaud BOISSERIE, Chargé de Recherche Tim D. WHITE, Professeur Thèse soutenue publiquement le 11 décembre 2012 Devant la Commission d’Examen Membres du Jury Denis GERAADS Directeur de Recherche, CNRS, Paris Rapporteur John M. HARRIS Professeur, Muséum d’Histoire Naturelle de Los Angeles Rapporteur Jean-Renaud BOISSERIE Chargé de Recherche, Université de Poitiers Examinateur Michel BRUNET Professeur, Collège de France, Paris Examinateur Stéphane DUCROCQ Directeur de Recherche, Université de Poitiers Examinateur Sevket SEN Directeur de Recherche, MNHN, Paris Examinateur Patrick VIGNAUD Professeur, Université de Poitiers Examinateur Tim D. WHITE Professeur, Université de Californie, Berkeley Examinateur 1 2 Résumé La sous-famille des Suinae est largement répandue en Afrique au Plio-Pléistocène et a été abondamment utilisée pour corréler biochronologiquement les sites à hominidés en se basant sur l’évolution morphologique rapide des troisièmes molaires dans différentes lignées. À partir d’un échantillon important de suinés africains actuels, les schémas de variation morphologique crânio-mandibulaire et dentaire sont quantifiés par morphométrie géométrique. Ce référentiel actuel sert à estimer la variabilité morphologique dans le registre fossile. La révision de la paléobiodiversité du genre Kolpochoerus (le suiné plio-pléistocène le plus abondant) dans les bassins du Turkana et de l’Awash s’appuie sur l’étude anatomique et morphométrique de matériels publié et inédit découverts dans la basse vallée de l’Omo (formation de Shungura) et dans la vallée moyenne de l’Awash en Éthiopie. Une nouvelle espèce fossile est décrite, les trajectoires évolutives au sein de chaque espèce sont quantifiées, et les interprétations biostratigraphiques sont révisées. Les liens de parenté au sein du genre Kolpochoerus sont décrits grâce à l’analyse cladistique. L’étude de la paléoécologie des suinés africains est basée sur l’anatomie comparée et la biogéochimie des isotopes stables (carbone et oxygène). Les suinés fossiles présentent une gamme de taille importante, ainsi que des régimes alimentaires et des habitats variés. Les nouvelles connaissances acquises sur leur biodiversité, leur phylogénie, et leur écologie permettent de proposer une histoire paléobiogéographique des Suinae en Afrique. Evolutionary history of the genus Kolpochoerus (Cetartiodactyla : Suidae) during the Plio-Pleistocene in eastern Africa. The subfamily Suinae is widespread in Plio-Pleistocene deposits in Africa, and was heavily used as a biochronological tool to correlate hominid-bearing sites based on the rapid morphological evolution of third molars in different lineages. A large sample of extant African suines enables to quantify patterns of variation in cranio-mandibular and dental morphology using geometric morphometrics. This modern referential is used to estimate the morphological variability in the fossil record. Revision of the paleobiodiversity of the genus Kolpochoerus (the most abundant Plio-Pleistocene suine) in Turkana and Awash basins is based on anatomical and morphometric studies of published and unpublished material discovered in the Lower Omo Valley (Shungura Formation) and in the Middle Awash Valley in Ethiopia. A new fossil species is described, evolutionary trajectories within each species are quantified, and biostratigraphic interpretations are revised. Phylogenetic relationships within the genus Kolpochoerus are described by the cladistic analysis. Paleoecological study of African suines is based on comparative anatomy and stable isotopic biogeochemistry (carbon and oxygen). Fossil suines display a wide range of body size, as well as various diets and habitats. The new results produced regarding their biodiversity, phylogeny, and ecology, enable to reconstruct a paleobiogeographic history of Suinae in Africa. Mots clés : Afrique ; biodiversité ; Éthiopie ; évolution ; Hominidae ; paléobiogéographie ; paléoécologie ; Pléistocène ; Pliocène ; phylogénie ; Suidae. Keywords: Africa ; biodiversity ; Ethiopia ; evolution ; Hominidae ; paleobiogeography ; paleoecology ; Pleistocene ; Pliocene ; phylogeny ; Suidae. Laboratoire : iPHEP (Institut de Paléoprimatologie, Paléontologie Humaine : Évolution et Paléoenvironnements) – UMR CNRS 7262 – Université de Poitiers Bât. B35 – 6, rue Michel Brunet, 86022 POITIERS Cedex 3 4 Remerciements À l’issue de ces trois années de thèse, voici enfin l’occasion de remercier toutes les personnes qui ont partagé de près ou de loin cette expérience enrichissante et inoubliable ! Je tiens dans ces quelques pages à remercier tous les gens qui ont rendu ce travail possible et m’ont aidé et soutenu. Après le mémoire de Master 2, c’est la deuxième fois que j’ai la chance et l’honneur d’être encadré par Jean-Renaud Boisserie dans mon cursus paléontologique. C’est plutôt la troisième fois puisqu’il avait également supervisé mon travail en Éthiopie avant le Master 2. Les mots ne suffisent pas pour lui exprimer ma reconnaissance. Depuis que je le connais, Jean-Renaud m’a apporté beaucoup scientifiquement et personnellement. Il a toujours su me faire profiter de son érudition scientifique et de son sens critique sur les différents aspects de la recherche en paléontologie. Il a toujours été présent quand j’en avais besoin et il a fait preuve d’énormément de patience et de compréhension tout au long de cette thèse. Il n’a jamais ménagé son temps (en particulier ses soirées pendant ces dernières semaines) et son énergie pour répondre à tous mes besoins et mes interrogations. En me confiant ce sujet de thèse sur les suidés et le matériel inédit collecté dans la formation de Shungura par son équipe, l’Omo Group Research Expedition, Jean-Renaud m’a mis en mains un sujet passionnant (je pense, autant pour lui que pour moi) et m’a donné tous les éléments pour conduire cette étude. Je lui en serai toujours reconnaissant. Tout au long de la thèse, il m’a guidé dans les différents aspects méthodologiques et théoriques. De la définition des points homologues sur les crânes de cochons à la façon de faire tourner les logiciels de cladistique en passant par toutes les manières possibles d’interpréter un signal isotopique, j’ai pu profiter de son expérience et de son esprit critique. Je le considère comme un exemple, un maître et également un ami. Je tiens également à exprimer toute ma gratitude à mon second directeur de thèse, Tim White. En me confiant l’étude de l’abondant matériel inédit collecté par le Middle Awash Research Project, il m’a témoigné de sa confiance et je l’en remercie. Pendant ses séjours à Addis Abeba, il a tout mis en œuvre pour que la préparation et l’organisation des collections de suidés soient terminées afin que je puisse les étudier. Malgré un emploi du temps surchargé, il a toujours été disponible et réactif (le mot est faible) à mes moindres demandes et m’a également régulièrement apporté son soutien pour des demandes de bourses. Il m’a fait profiter de ses connaissances encyclopédiques sur les suidés fossiles et plus généralement, 5 l’évolution des faunes africaines. Je le remercie particulièrement de m’avoir ouvert les yeux sur l’importance du contexte géologique et des datations (et leurs limites). Cette thèse n’est que la première pierre d’un travail que j’envisage de poursuivre pendant de longues années. Tim White, en m’acceptant comme post-doctorant à l’Université de Californie, Berkeley, à partir de l’année prochaine, va me permettre de continuer notre collaboration et de l’étendre à d’autres projets menés par les
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