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Les Faunes Mammaliennes Du Miocène Supérieur Du Tchad : Structure Des Communautés Et Implications Paléoenvironnementales

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Les Faunes Mammaliennes Du Miocène Supérieur Du Tchad : Structure Des Communautés Et Implications Paléoenvironnementales THESE Pour l’obtention du grade de DOCTEUR DE L’UNIVERSITE DE POITIERS U.F.R Sciences Fondamentales et Appliquées (Diplôme National – Arrêté du 25 avril 2002) Ecole Doctorale : Sciences pour l’Environnements Gay Lussac Secteur de Recherche : Terre Solide et Enveloppes Superficielles Discipline : Paléontologie des Vertébrés Présentée par Soizic LE FUR Les faunes mammaliennes du Miocène supérieur du Tchad : structure des communautés et implications paléoenvironnementales Sous la direction de Patrick VIGNAUD, Professeur Emmanuel FARA, Maître de Conférences Thèse soutenue publiquement le 4 décembre 2009 Devant la Commission d’Examen Membres du Jury Z. ALEMSEGED PhD, Chair and Curator, California Academy of Sciences Rapporteur L.J. FLYNN PhD, Assistant Director, Harvard University Rapporteur M. BRUNET Professeur, Université de Poitiers & Collège de France Examinateur/Président E. FARA Maître de Conférences, Université de Bourgogne Examinateur J.-J. JAEGER Professeur, Université de Poitiers Examinateur H.T. MACKAYE Maître de Conférences, Université de N’Djamena Examinateur P. VIGNAUD Professeur, Université de Poitiers Examinateur Remerciements Ces trois années de thèses ont été l’occasion pour moi de côtoyer et d’apprécier un certain nombre de personnes, appartenant au monde de la recherche ou autre. Je tiens à présent à leur exprimer toute ma gratitude pour leur contribution, de près ou de loin, à ce travail de recherche, par leurs conseils avisés mais également par leur soutien inconditionnel. C’est ainsi que je me lance dans l’exercice périlleux des remerciements que je ferai concis, mais je l’espère sans oubli. Je commencerai tout naturellement par remercier mes directeurs de thèse, Patrick Vignaud et Emmanuel Fara, sans qui bien sûr je n’en serais pas là aujourd’hui. La complémentarité de leurs domaines de compétences a été une vraie richesse pour mon apprentissage. Merci pour toute l’aide que vous m’avez apporté, vos conseils précieux, votre soutien moral, et pour votre grande disponibilité, malgré des emplois du temps outrageusement chargés. Je remercie également Michel Brunet et Jean-Jacques Jaeger pour m’avoir accueilli au sein de l’iPHEP. En particulier, Michel Brunet pour avoir contribué à proposer ce sujet de thèse. J’espère finalement avoir répondu à vos attentes et surtout en avoir suscité de nouvelles. Je tiens également à remercier les autres membres du jury, mes rapporteurs, Lawrence Flynn et Zeresenay Alemseged, ainsi que Hassan Taïsso Mackaye, d’avoir bien voulu consacrer de leur temps à évaluer mon travail de recherche. J’ai passé trois années extrêmement enrichissantes au sein de l’iPHEP, sur le plan professionnel mais également sur le plan humain, et c’est à vous tous que je le dois : Ghislaine Florent, Carine Noël, Franck Guy, Cécile Blondel, Mathieu Schuster, Olga Otero, Géraldine Garcia, Jean-Renaud Boisserie, Stéphane Ducrocq, Xavier Valentin, Olivier Chavasseau, Sabine Riffaut, Mouloud Benammi, Louis de Bonis, Vincent Lazzari et Amal Le Beurre. i J’adresse également mes remerciements à Anne Brunellière, pour sa remarquable efficacité dans la recherche d’articles et sa grande gentillesse, ainsi qu’Audrey pour sa bonne humeur et sa disponibilité. Je ne saurai oublier les « extérieurs », anciens pictaves et/ou collaborateurs, justifiant ainsi leurs visites (trop peu fréquentes) au laboratoire : Stéphane Peigné, avec qui j’ai pu effectuer mes tous premiers pas dans la recherche ; mais également Fabrice Lihoreau, Gildas Merceron et Doris Barboni. Les discussions que j’ai pu avoir avec chacun, touchant de près ou de loin à mes recherches, ont été très enrichissantes à chaque fois. Ces trois années n’auraient pas été les mêmes sans la présence de mes collègues (et amis) thésards et post-docs : Aurélie, Camille, Pauline, Diogo, Thibaut, Diana, Guillaume, Antoine, Alice, Renaud, Faysal et j’ajouterai à cette liste Sohee et Golpar, presque du labo maintenant… Ainsi que Le P’tit Cabaret qui a su accueillir nos soirées quelque peu arrosées. Je remercie également mes collègues du CIES, entre autres : Marc, Fred, Marie, Julien, Alice, Adeline, Romain, Cyrille, avec lesquels j’ai passé des soirées très « animées » et pu partager des expériences CIES I-NOU-BLI-ABLES ! Je terminerais en remerciant ma famille (toujours présente et chaleureuse, merci pour les A/R Poitiers-Bretagne et pour votre soutien), et mes potes !! Alain, Nat, Marie, Seb, Floue, Arnaud, Fanny, Guillaume, Claire, Emeline, Eric, Mily (psst, thanks pour les cours anti-stress sur la fin), Cédrikus, P-Y, Laure, Oliv, Erel, Elod, Yo, Bertrand et Rachelle. Enfin, merci au capo du 4x4 (que je n’aurai malheureusement pas eu le plaisir de rencontrer) qui a permis de révéler toute la diversité faunique du secteur de TM, véritable point de départ de mon sujet de thèse. ii TABLE DES MATIERES INTRODUCTION 1 PREMIERE PARTIE : JEUX DE DONNEES ET METHODOLOGIE 9 Chapitre 1 : Terminologie de la paléoécologie 11 Chapitre 2 : Les faunes fossiles du Mio-Pliocène d’Afrique 15 I- Assemblages mio-pliocènes du Tchad 15 I.1- Secteur fossilifère de Toros-Menalla (TM) 15 I.1.1- Quel est l’effectif minimal à considérer pour qu’un échantillon devienne représentatif de l’assemblage fossile dont il est issu ? 18 I.1.2- La constitution de répliques permet-elle d’obtenir un échantillon plus représentatif de l’assemblage fossile dont il est issu ? 25 I.2- Secteurs pliocènes du désert du Djourab : Koro Toro, Kollé et Kossom Bougoudi 29 II- Assemblages africains du Miocène supérieur-Pliocène basal 31 III- Construction des jeux de données 37 Chapitre 3 : Base de données modernes 41 I- Définition des environnements africains modernes 41 I.1- Classification des environnements africains modernes 41 I.1.1- Selon le type d’habitats 41 I.1.2- Selon l’agencement des habitats 43 I.2- Quel lien entre facteurs climatiques et formations végétales ? 44 II- Jeux de données 45 Chapitre 4 : Techniques d’analyses 49 I- Codage des données 49 I.1- Structure taxinomique, diversité écologique et taxinomique 49 I.1.1- Structure taxinomique 49 I.1.2- Composition taxinomique 53 I.1.3- Diversité écologique 54 I.1.4- Diversité taxinomique 61 I.2- Transformation et standardisation des données 62 II- Techniques d’analyses multivariées 65 II.1- Techniques de classification 66 II.2- Méthodes d’ordination 69 II.3- Analyses discriminantes 72 III- Indices de similitude 74 IV- Tests statistiques 76 iii DEUXIEME PARTIE : LA FAUNE MAMMALIENNE DE TOROS- MENALLA ET SES IMPLICATIONS PALEOENVIRONNEMENTALES 75 Chapitre 1 : Structuration faunique des assemblages fossiles de TM 81 I- Contexte taphonomique : des assemblages représentatifs de leur communauté(s) d’origine ? 81 I.1- Processus taphonomiques identifiés à TM 81 I.2- Biais taphonomiques associés aux assemblages de TM 91 II- Toros-Menalla : un ensemble faunique homogène ? 94 II.1- Structures écologiques des assemblages de TM 94 II.2- Structures taxinomiques des assemblages de TM 97 II.3- Compositions taxinomiques des assemblages de TM 98 II.4- Diversités taxinomiques des assemblages de TM 98 III- Toros-Menalla : deux structures fauniques distinctes 116 III.1- Réel signal écologique ou biais taphonomique ? 116 III.2- Comment s’organisent ces deux ensembles dans le temps et l’espace ? 121 IV- Conclusion 123 Chapitre 2 : Le(s) paléoenvironnement(s) de TM 127 I- Comment la structure faunique permet-elle de discriminer les environnements africains modernes ? 127 I.1- Conséquence de l’exclusion des petites espèces dans la résolution du modèle nul 127 I.2- Identification des habitats présents dans un environnement 129 I.2.1- Apports des diversités écologique et taxinomique 129 I.2.1.1- Densité du couvert arboré 129 I.2.1.2- Présence d’habitats édaphiques 140 I.2.2- Apports de la structure taxinomique 142 I.2.3- Apports de la composition taxinomique 145 I.3- Structuration de l’environnement 147 I.3.1- Diversité d’habitats 147 I.3.2- Agencement des habitats dans l’espace 149 II- Quel signal paléoenvironnemental la faune fossile de Toros-Menalla a-t-elle enregistré ? 150 II.1- Singularité des assemblages fossiles par rapport aux faunes modernes : quelle(s) origine(s) ? 152 II.1.1- Des environnements sans équivalent moderne ? 152 II.1.2- Un biais méthodologique ? 153 II.1.3- Un biais taphonomique ? 154 II.2- Quel environnement pour les faunes de TM ? 160 II.2.1- Densité du couvert végétal 160 II.2.2- Structure de l’environnement 160 III- Conclusion 170 iv TROISIEME PARTIE : TOROS-MENALLA EN AFRIQUE AU MIOCENE SUPERIEUR-PLIOCENE BASAL : SIMILARITES FAUNIQUES ET PALEOENVIRONNEMENTALES AVEC LES AUTRES GISEMENTS FOSSILIFERES 173 I- Diversité des paléoenvironnements en Afrique au Miocène supérieur-Pliocène basal 175 I.1- Identification des biais associés aux assemblages fossiles africains 175 I.1.1- Une sous-représentation des petites espèces ? 175 I.1.2- Un mélange de communautés ? 175 I.1.3- Profil écologique des assemblages fossiles 177 I.2- Densité du couvert végétal 184 I.3- Structure de l’environnement 193 I.4- Bilan : quels étaient les environnements présents en Afrique au Miocène supérieur-Pliocène basal ? 195 II- Provinces mammaliennes représentées en Afrique au Miocène supérieur- Pliocène basal 199 II.1- Quelles sont les limites de telles inférences ? 199 II.2- Quelles étaient les provinces mammaliennes représentées en Afrique au Miocène supérieur-Pliocène basal ? 201 III- Particularités environnementales et taxinomiques des sites à hominidés anciens 212 III.1- Quel habitat les hominidés anciens occupaient-ils ? 212 III.2- Quels sont les taxons clés associés aux hominidés anciens ? 213 CONCLUSIONS ET PERSPECTIVES 217 Références Bibliographiques 225 Annexes 251 v INTRODUCTION 1 CONTEXTE Les mécanismes de l’évolution sont principalement régis par l’interaction entre les caractéristiques intrinsèques des taxons (propriétés héritées telles que la structure de population, le système de fertilisation, le spectre écologique, etc.) et les facteurs environnementaux, biotiques (interaction avec les autres êtres vivants, végétaux ou animaux) et abiotiques (température, précipitations, etc.) (Todd, 1999; Vrba, 1999).
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