I UNIVERSITÉ DU QUÉBEC À RIMOUSKI DISPARITÉ MORPHOLOGIQUE ET MODULARITÉ DES NAGEOIRES MÉDIANES ET PAIRES CHEZ LES VERTÉBR

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I UNIVERSITÉ DU QUÉBEC À RIMOUSKI DISPARITÉ MORPHOLOGIQUE ET MODULARITÉ DES NAGEOIRES MÉDIANES ET PAIRES CHEZ LES VERTÉBR i UNIVERSITÉ DU QUÉBEC À RIMOUSKI DISPARITÉ MORPHOLOGIQUE ET MODULARITÉ DES NAGEOIRES MÉDIANES ET PAIRES CHEZ LES VERTÉBRÉS BASAUX THÈSE PRÉSENTÉE COMME EXIGENCE PARTIELLE DU DOCTORAT EN BIOLOGIE EXTENTIONNÉ DE L’UNIVERSITÉ DU QUÉBEC À MONTRÉAL PAR OLIVIER LAROUCHE SEPTEMBRE 2017 ii v UNIVERSITÉ DU QUÉBEC À RIMOUSKI Service de la bibliothèque Avertissement La diffusion de ce mémoire ou de cette thèse se fait dans le respect des droits de son auteur, qui a signé le formulaire « Autorisation de reproduire et de diffuser un rapport, un mémoire ou une thèse ». En signant ce formulaire, l’auteur concède à l’Université du Québec à Rimouski une licence non exclusive d’utilisation et de publication de la totalité ou d’une partie importante de son travail de recherche pour des fins pédagogiques et non commerciales. Plus précisément, l’auteur autorise l’Université du Québec à Rimouski à reproduire, diffuser, prêter, distribuer ou vendre des copies de son travail de recherche à des fins non commerciales sur quelque support que ce soit, y compris l’Internet. Cette licence et cette autorisation n’entraînent pas une renonciation de la part de l’auteur à ses droits moraux ni à ses droits de propriété intellectuelle. Sauf entente contraire, l’auteur conserve la liberté de diffuser et de commercialiser ou non ce travail dont il possède un exemplaire. iii REMERCIEMENTS Je tiens à exprimer ma reconnaissance envers tous les membres du Laboratoire de Paléontologie et de Biologie évolutive de l’UQAR qui ont contribué, de près ou de loin, au succès de ce projet. Je souligne particulièrement les contributions de Marion Chevrinais et de Cyrena Riley : leurs commentaires et suggestions ont été d’une aide inestimable lors de la finalisation de l’introduction générale et de la conclusion générale de cette thèse doctorale. Je remercie aussi tout spécialement mes directeurs de recherche, Richard Cloutier et Miriam L. Zelditch, non seulement pour leur encadrement et leur support tout au long de ce projet, mais aussi pour ces innombrables discussions enrichissantes qui m’ont fait évolué en tant que biologiste au cours des cinq dernières années. Finalement je tiens à remercier Dr. R. Craig Albertson, Dr. Piero Calosi et Dr. William Vickery pour avoir accepté de faire partie du commité d’évaluation lors de ma soutenance de thèse doctorale. Leurs commentaires judicieux et leurs suggestions m’ont ouvert de nouvelles pistes de réflexion non seulement en ce qui concerne mon project doctoral, mais aussi pour la suite de ma carrière académique. Ce projet n’aurait pu être mené à terme sans les contributions financières de plusieurs organismes subventionnaires, notamment le Conseil de Recherches en Sciences naturelles et en Génie du Canada, les Fonds de Recherches Nature et Technologies Québec, et les Services aux Étudiants de l’UQAR. iv v AVANT-PROPOS « There once were two watchmakers, named Hora and Tempus, who manufactured very fine watches. Both of them were highly regarded, and the phones in their workshops rang frequently - new customers were constantly calling them (…) The watches the men made consisted of about 1000 parts each. Tempus had so constructed his that if he had one partly assembled and had to put it down - to answer the phone say - it immediately fell to pieces and had to be reassembled from the elements (…) The watches that Hora made were no less complex than those of Tempus. But he had designed them so that he could put together subassemblies of about ten elements each. Ten of these subassemblies, again, could be put together into a larger subassembly ; and a system of ten of the latter subassemblies constituted the whole watch. Hence, when Hora had to put down a partly assembled watch in order to answer the phone, he lost only a small part of his work (…) » (Simon 1962, p. 468) vi vii TABLE DES MATIÈRES REMERCIEMENTS………………………………………………………………….ii AVANT-PROPOS……………………………………………………………………iv TABLE DES MATIÈRES…………………………………………………………...vii LISTE DES FIGURES………………………………………………………………..xi LISTE DES TABLEAUX…………………………………………………………...xiii RÉSUMÉ…………………………………………………………………………….xv ABSTRACT……………………………………………………………………..…xvii INTRODUCTION GÉNÉRALE…………………………………………………...…1 Disparité morphologique……….......................................................................2 Modularité……………………………………………………………………..3 Relation entre modularité et disparité morphologique………………………...6 Les poissons comme organismes modèles………………………………….…8 Les nageoires chez les poissons : définitions opérationnelles……………...…11 Disparité morphologique dans la configuration des nageoires…………….…13 Apparition séquentielle des nageoires…………………………………….….16 Apparition des nageoires médianes…………………………………..17 Duplications de la nageoire dorsale………………………………….18 Apparition des nageoires paires……………………………………...20 Modularité des nageoires………………………………………………….…24 Hypothèses de modularité des nageoires médianes.………………….24 Hypothèses de modularité des nageoires paires.……………………..27 La problématique…………………………………………………………….30 Objectifs de recherche et méthodologie utilisée…....………………………...32 CHAPITRE I MODULES DE NAGEOIRES : UNE PERSPECTIVE ÉVOLUTIVE SUR LA DISPARITÉ DES APPENDICES CHEZ LES VERTÉBRÉS BASAUX 1.1 Résumé………………………………………………………………39 viii 1.2 Abstract………………………………………………………………42 1.3 Introduction…………………………………………………………..43 1.4 Methods……………………………………………………………...47 1.4.1 Fin presence/absence dataset…………...…………………....47 1.4.2 Fish supertree...……………………………………………...50 1.4.3 Mapping of the fin characters on the supertree……………....53 1.4.4 Covariation in the number of fins present………………….....53 1.5 Results………………………………………………………………..55 1.5.1 Fish supertree...……………………………………………...55 1.5.2 Mapping the evolutionary history of fish appendages………..57 1.5.3 Covariation in the presence/absence of fish appendages…….62 1.6 Discussion……………………………………………………………70 1.6.1 Disparity in fin configurations……………………………….71 1.6.2 Evolutionary history of fish appendages……………………..75 1.6.3 Evidence for fin evolutionary modules……………………….83 1.7 Conclusions..…………………………………………………............87 CHAPITRE II TÊTE, CORPS ET NAGEOIRES : PATRONS D’INTÉGRATION MORPHOLOGIQUE ET DE MODULARITÉ CHEZ LES POISSONS 2.1 Résumé………………………………………………………………89 2.2 Abstract………………………………………………………………92 2.3 Introduction…………………………………………………………..93 2.4 Methods……………………………………………………………...96 2.4.1 Sample………………………………………………………..96 2.4.2 Geometric Morphometrics…………………………………...97 2.4.3 Standardizing for Preservational Artifacts and for Size…..….98 2.4.4 Testing Hypotheses of Modularity………………………...….99 2.5 Results………………………………………………………………106 2.5.1 Distance-Matrix Method……………………………………111 ix 2.5.2 Minimum Deviance Method………………………………...112 2.5.3 Escoufier’s RVs………………………………………….….112 2.5.4 A Posteriori Results.…………………………………….…..113 2.6 Discussion…………………………………….…………………….115 CHAPITRE III TÊTES, NAGEOIRES ET QUEUES : UNE ANALYSE COMPARATIVE DES PATRONS D’INTÉGRATION MORPHOLOGIQUE ET DE MODULARITÉ ENTRE LES ÉCHELLES MICRO- ET MACROÉVOLUTIVES 3.1 Résumé……………………………………………………………..122 3.2 Abstract……………………………………………………………..123 3.3 Introduction…………………………………………………………124 3.4 Methods…………………………………………………………….126 3.4.1 Sample……………………………………………………....126 3.4.2 Geometric morphometrics………………………………….129 3.4.3 Taking into account phylogeny…………………………..….132 3.4.4 Quantifying allometry………………………………………133 3.4.5 Testing hypotheses of modularity………………..………….133 3.4.6 Comparing evolutionary rates and disparity among partitions……………………………...…………………………..139 3.4.7 Software…………………………………………………….139 3.5 Results………………………………………………………………140 3.5.1 Testing evolutionary models…………………………….…..140 3.5.2 Covariance ratios………………………………….………..142 3.5.3 Graphical modeling………………………………………...144 3.5.4 Minimum deviance method………………………………….145 3.5.5 Evolutionary rate ratios…………………………………….146 3.5.6 Comparing micro- and macroevolutionary patterns of integration…………………………………………………..147 3.6 Discussion…………………………………………………………..149 3.6.1 Support for evolutionary fin modules……………………….150 x 3.6.2 Modularity, morphological disparity and rates of morphological evolution……………………………………….........151 3.6.3 Correspondence between micro- and macroevolutionary patterns of integration………………..…..........................................153 3.7 Conclusion………………………………………………………….156 CONCLUSION GÉNÉRALE………………………………………………….…...159 Contexte, originalité de l’étude et rappel des objectifs…………….………..159 Principaux résultats…………………………………………………………160 Portée de l’étude…………………………………………………….………166 Limitations de l’étude……………………………………………….………167 Perspectives de recherche dans le domaine concerné………………….……170 Recherches futures………………………………………………………….172 ANNEXE A : HOW TO DEFINE A FIN?.................................................................175 ANNEXE B : A HISTORICAL ACCOUNT OF APPENDAGE DIVERSITY AND HOMOLOGY AMONG BASAL CHORDATES……………………………….….179 ANNEXE C : LIST OF SOURCE TREES FOR THE SUPERTREE ANALYSIS AND ADDITIONAL RESULTS……………………………………………………..…..217 ANNEXE D : PRESENCE/ABSENCE DATASET………………………………..227 ANNEXE E : ADDITIONAL RESULTS OF MULTIPLE CORRESPONDENCE ANALYSES………………………………………………………………………..239 ANNEXE F : ADDITIONAL RESULTS FOR THE DANIO RERIO AND CHROSOMUS EOS DATASETS INCORPORATING ADDITIONAL MODULARITY HYPOTHESES…………………………………………………..243 LEXIQUE…………………………………………………………………………..247 BIBLIOGRAPHIE...………………………………………………………………..249 xi LISTE DES FIGURES Figure 1 : Synthèse des relations phylogénétiques entre les groupes de poissons existants et fossiles illustrant l’importante disparité morphologique qui caractérise les poissons dans leur ensemble……………………….……...9 Figure 2
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