Evolution Et Adaptation Fonctionnelle Des Arbres Tropicaux : Le Cas Du Genre Guibourtia Benn

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Evolution Et Adaptation Fonctionnelle Des Arbres Tropicaux : Le Cas Du Genre Guibourtia Benn EVOLUTION ET ADAPTATION FONCTIONNELLE DES ARBRES TROPICAUX : LE CAS DU GENRE GUIBOURTIA BENN. FELICIEN TOSSO COMMUNAUTE FRANÇAISE DE BELGIQUE UNIVERSITE DE LIEGE – GEMBLOUX AGRO-BIO TECH EVOLUTION ET ADAPTATION FONCTIONNELLE DES ARBRES TROPICAUX : LE CAS DU GENRE GUIBOURTIA BENN. Dji-ndé Félicien TOSSO Dissertation originale présentée en vue de l’obtention du grade de docteur en sciences agronomiques et ingénierie biologique Co-Promoteurs : Prof. Jean-Louis Doucet et Dr. Olivier J. Hardy Année 2018 Copyright Cette œuvre est sous licence Creative Commons. Vous êtes libre de reproduire, de modifier, de distribuer et de communiquer cette création au public selon les conditions suivantes : - paternité (BY) : vous devez citer le nom de l'auteur original de la manière indiquée par l'auteur de l'œuvre ou le titulaire des droits qui vous confère cette autorisation (mais pas d'une manière qui suggérerait qu'ils vous soutiennent ou approuvent votre utilisation de l'œuvre) ; - pas d'utilisation commerciale (NC) : vous n'avez pas le droit d'utiliser cette création à des fins commerciales ; - partage des conditions initiales à l'identique (SA) : si vous modifiez, transformez ou adaptez cette création, vous n'avez le droit de distribuer la création qui en résulte que sous un contrat identique à celui-ci. À chaque réutilisation ou distribution de cette création, vous devez faire apparaitre clairement au public les conditions contractuelles de sa mise à disposition. Chacune de ces conditions peut être levée si vous obtenez l'autorisation du titulaire des droits sur cette œuvre. Rien dans ce contrat ne diminue ou ne restreint le droit moral de l’auteur. Dédicace A mes chers parents, Etienne M. Tosso et Jeanne Codjo i ii Résumé Tosso Félicien (2018). Evolution et adaptation fonctionnelle des arbres tropicaux : le cas du genre Guibourtia Benn. Thèse de doctorat. Université de Liège - Gembloux Agro-Bio Tech, Belgique. 273 p. + annexes. Résumé La présente thèse s’intéresse aux mécanismes à l’origine de la diversification des espèces d’arbres tropicaux. Elle utilise le genre Guibourtia Benn. (Fabaceae- Detarioideae) comme modèle biologique afin de comprendre les mécanismes historiques, biologiques et environnementaux, à l'origine de la diversité de ce genre aux niveaux interspécifique et intraspécifique. Plus particulièrement, elle vise à : (1) étudier au niveau interspécifique, le rôle relatif des forces évolutives neutres et de sélection dans la diversification du genre Guibourtia en combinant une phylogénie moléculaire avec la caractérisation des traits et les niches écologiques des espèces ; et (2) au niveau intraspécifique, questionner les causes de la différenciation des populations de trois espèces de Guibourtia. Au niveau interspécifique, la phylogénie datée basée sur le séquençage du génome chloroplastique complet a globalement confirmé la taxonomie actuelle (espèces généralement monophylétiques). Elle a montré une diversification au milieu du Miocène en trois clades qui sont décrits aujourd’hui comme des sous-genres (Guibourtia, Gorskia et Pseudocopaiva). Elle démontre en outre que deux espèces américaines sont issues d’une migration de l’Afrique vers l’Amérique à la fin du Miocène. Il est également apparu que certains traits morphologiques ont été sélectionnés de manière convergente au sein des différents clades du genre Guibourtia en fonction des niches climatiques des espèces. Ce dernier résultat a été consolidé au moyen d’une expérimentation écophysiologique prouvant que la lumière constitue un important facteur de sélection et de différenciation adaptative entre trois espèces (G. ehie, G. coleosperma et G. tessmannii). Au niveau intraspécifique, une étude de phylogéographie de deux espèces a permis de mettre en évidence que les barrières biogéographiques chez G. ehie et les gradients climatiques chez G. coleosperma, auraient contribué à la différenciation génétique des populations. En outre, cette étude montre une forte différenciation entre les populations de G. ehie d’Afrique de l’Ouest et d’Afrique centrale en lien avec quelques traits morphologiques, ce qui préjuge de l’existence d’une nouvelle espèce. Enfin, chez G. tessmannii, espèce aux fruits déhiscents et graines arillées, nous avons identifié les principaux disperseurs dont les calaos (Ceratogymna atrata ) qui pourraient contribuer à une dispersion à longue distance, influençant la structure spatiale de la variation génétique des populations. Cette thèse en utilisant le genre Guibourtia comme modèle d’étude a apporté de nouveaux éléments à la compréhension des mécanismes qui génèrent la diversité au sein des espèces d’arbres. Elle fournit en outre des connaissances originales sur des espèces inscrites en annexe II de la CITES. iii Abstract Tosso Félicien (2018). Evolution and functional adaptation of tropical trees species : the case study of Guibourtia Benn. PhD thesis. University of Liège - Gembloux Agro-Bio Tech, Belgium. 273 p. + appendixes. Abstract This thesis studies the diversification processes of tropical tree species. It uses the genus Guibourtia Benn. as a biological model (Leguminosae-Detarioideae) in order to understand historical, biological and environmental mechanisms that are at the origin of the diversity of the genus. In particular, this PhD aims to: (1) understand at the interspecific level the relative role of neutral and selection forces in the diversification of Guibourtia species by combining molecular phylogeny with the characterization of functional traits and ecological niches; and (2) at the intraspecific level, question the causes of population differentiation of three Guibourtia species. At the interspecific level, the molecular phylogeny dating based on whole plastid genome sequencing was congruent with the current taxonomy (species were generally monophyletic). It showed a diversification in the middle of the Miocene in three clades corresponding with the subgenera (Guibourtia, Gorskia and Pseudocopaiva). It also demonstrated that two Neotropical species resulted from a migration from Africa to South America during the late Miocene. Considering climate niches, it was also demonstrated that some morphological traits have been selected in a convergent way within the different clades. This result was confirmed by an ecophysiological experiment, which highlighted that light was an important factor of selection and adaptive differentiation between three species (G. ehie, G. coleosperma and G. tessmannii). At the intraspecific level, a phylogeographic study of two Guibourtia species revealed that the biogeographic barriers in G. ehie and the climatic gradients in G. coleosperma, would have contributed to the genetic differentiation of populations. In addition, this study questions the delimitation of G. ehie since its populations showed a strong genetic differentiation between West and Central Africa. This differentiation was associated with diagnostic morphological features. Finally, in G. tessmannii, a species with dehiscent fruits and arillate seeds, we identified the main dispersal species, including hornbills (Ceratogymna atrata), which may have contributed to long-distance gene flow, influencing the spatial structure of genetic variation among populations. This thesis, using the Guibourtia genus as a biological model brings new elements helping in the understanding of mechanisms generating diversity within tropical tree species. It also provides important information on species listed on Appendix II of CITES. iv Remerciements Remerciements Jérôme Touzalin dira : “Il n’y a pas de hasard… il n’y a que des rendez-vous qu’on ne sait pas lire“. Ce travail de doctorat est le pur produit d’une rencontre. Celle avec l’enseignant de cours de Gestion des écosystèmes tropicaux qui m’a offert sans hésiter l’opportunité de faire mon doctorat. Sans lui, sans le Professeur Jean-Louis Doucet, cette présente thèse ne sera tout simplement pas possible. Jean-Louis, au-delà du cadre académique, c’est ici l’occasion de t’exprimer toute ma gratitude pour l’encadrement de grande qualité, la rigueur scientifique, l’esprit d’ouverture, les conseils, l’attention et l’amitié. Derrière ton implication personnelle dans ce travail se cache également celle de Bérengère, Robin et Florine. Avec mes faibles mots, je vous dis Merci. Tu m’as connecté au Dr. Olivier J. Hardy, co-promoteur de cette thèse qui a joué un rôle central dans l’aboutissement de ce travail. Olivier, je tiens à te remercier pour l’héritage génétique qui restera à jamais gravé dans mon ADN. J’aimerais te dire combien j’ai apprécié ta grande disponibilité tout au long de ce travail. J’ai beaucoup appris de ta simplicité et de ton contact, tant humainement que scientifiquement. Je voudrais exprimer ma gratitude au Dr. Kasso Daïnou pour son coaching de proximité, ses multiples encouragements, son éclairage et conseils qui ont enrichi de façon considérable mes reflexions tout au long de cette thèse. J’en suis profondément reconnaissant. Je souhaiterais aussi addresser mes mots de remerciement aux membres du comité de thèse qui ont suivi le présent travail. En plus des trois personnes mentionnées plus haut, je pense ici au : Prof. Marc Aubinet, Dr. Adeline Fayolle, Dr. Alain Hambuckers et Dr. Christine Moureaux. Vous avez profondément amélioré la qualité de ce travail par vos critiques, discussions, suggestions et contributions. Sans vous, cette thèse ne serait pas ce qu'elle est. Je remercie tout particulièrement le Fonds pour la Formation à la Recherche dans l'Industrie et dans l'Agriculture - Fonds
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