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Bertrand MALLET Effet De L'habitat Et Des Pollinisateurs Sur La THÈSE Présentée à l’Université de la Réunion pour obtenir le diplôme de DOCTORAT SPÉCIALITÉ : Biologie des Populations et Écologie ÉCOLE DOCTORALE : Sciences Technologies Santé RÔLE DES FACTEURS ÉCOLOGIQUES DANS LE PROCESSUS DE SPÉCIATION EN MILIEU INSULAIRE Effet de l’habitat et des pollinisateurs sur la diversification du faham (Jumellea spp., Orchidaceae) aux Mascareignes Bertrand MALLET Soutenue publiquement le 7 novembre 2014 devant le jury composé de : Guillaume BESNARD chargé de recherche première classe, CNRS, Toulouse Rapporteur Bertrand SCHATZ, directeur de recherche, CNRS, Montpellier Rapporteur Pascale BESSE, professeure, Université de La Réunion Examinatrice Luc GIGORD, directeur du Conservatoire Botanique National de Mascarin Examinateur Laurence HUMEAU, maître de conférence, Université de La Réunion Examinatrice Thierry PAILLER, professeur, Université de La Réunion Directeur Résumé Les mécanismes à l’origine de la mise en place de l’isolement reproducteur sont relativement peu étudiés dans les radiations adaptatives insulaires. Selon la théorie de la spéciation écologique, l’isolement reproducteur résulte de la sélection divergente agissant sur des populations écologiquement différentes. Par leurs caractéristiques uniques, les îles océaniques constituent des systèmes idéals pour étudier le rôle des facteurs écologiques dans la diversification des lignées endémiques. Le rôle de l’habitat et des pollinisateurs dans la restriction des flux géniques inter et intraspécifique est examiné chez le faham (Jumellea rossii et J. fragrans), une orchidée endémique des Mascareignes. Pour ce faire, les patrons de différenciation phénotypique (traits floraux morphologiques et chimiques), environnementale (habitat, altitude, climat) et génétique (microsatellites nucléaires) ont été confrontés. A l’échelle intraspécifique, les résultats montrent que les flux de gènes sont principalement limités par les différences environnementales entre habitats sans rôle évident des pollinisateurs. A l’échelle interspécifique, outre les variations d’habitat, le changement de pollinisateur semble expliquer l’absence de flux de gènes. Dans le cadre du continuum de spéciation chez le faham, l’isolement reproducteur évoluerait en premier lieu en réponse à l’adaptation aux différents habitats, puis serait renforcé par l’adaptation à différents pollinisateurs. La révision de la taxinomie du clade et les priorités en terme de conservation sont également discutées. Abstract THE ROLE OF ECOLOGY IN ISLAND SPECIATION Effect of habitat and pollinator shifts on the diversification of faham (Jumellea spp., Orchidaceae) in the Mascarene Islands Mechanisms responsible for the establishment of reproductive isolation are little studied in island adaptive radiations. According to the theory of ecological speciation, reproductive isolation results from the ecologically-based divergent selection. By their unique characteristics, oceanic islands are ideal systems to study the role of ecological factors in the diversification of endemic lineages. This study focuses on the role of habitat and pollinators in restricting inter and intraspecific gene flow between populations of faham (Jumellea spp.), an orchid endemic to Mascarene Islands. To do this, patterns of phenotypic differentiation (floral traits), environmental differentiation (habitat, altitude, climate) and genetic differentiation (nuclear microsatellites) were compared. At the intraspecific level, the results show that gene flow restriction is mainly due to the colonization of different habitat types with no obvious role of pollinators. At the interspecific level, in addition to the role of habitat, pollinator shift seems to explain effective complete reproductive isolation. By placing these results into the continuum of speciation, it appears that reproductive isolation evolve initially in response to adaptation to different habitats and, in a second phase, would be enhanced by pollinator-driven divergent selection. Operationally, this study reviews the taxonomy of faham and proposes to define management units and priorities in terms of conservation. Remerciements ‘Autrui, pièce maîtresse de mon univers.’ Michel Tournier, Vendredi ou les limbes du Pacifique. En premier lieu, je tiens à remercier très sincèrement mon directeur de thèse Thierry Pailler et mon encadrante Laurence Humeau de m’avoir permis de découvrir la recherche scientifique en écologie évolutive il y a déjà six ans et d’avoir ensuite accepté de m’encadrer sur une thèse de doctorat. Je remercie Thierry en particulier pour avoir partagé ses nombreuses connaissances sur les milieux naturels de la Réunion et pour avoir su m’emporter dans ses réflexions aussi passionnantes que déstabilisantes. J’ai vraiment apprécié les sorties en forêt que l’on a pu faire ensemble et les discussions scientifiques que l’on a pu avoir. Je remercie Laurence pour son soutien sans faille, pour sa disponibilité inconditionnelle, pour sa rigueur, pour sa gentillesse, pour les nombreuses discussions de travail où elle a toujours accepté mes ‘Non, mais …’, pour m’avoir initié à la biologie moléculaire, à la rédaction d’article scientifique. Bref, pour avoir fait en sorte que je puisse réaliser ce travail de thèse dans les meilleures conditions possibles. Je remercie Florent Martos pour sa précieuse contribution à ce travail de thèse, pour son enthousiasme communicatif et pour avoir permis la collaboration avec l’Afrique du sud. Merci au Conseil Général de la Réunion, à l’Université de la Réunion, au Ministère des Outre-mer, à la Région Réunion et à la Société Botanique de France d’avoir participé au financement de ce projet. Merci au Parc national de la Réunion, à l’Office National des Forêts de la Réunion et au National Parks & Conservation Services qui ont permis la réalisation de ce travail en fournissant les autorisations administratives et les appuis logistiques des campagnes de terrain. Merci à Claudia Baider du Mauritius Herbarium et Steve Johnson de l’Université du Kwazulu-Natal d’avoir accepté de collaborer à ce travail. Merci aux membres du jury – Guillaume Besnard, Bertrand Schatz, Pascale Besse et Luc Gigord – d’avoir accepté d’évaluer ce travail. Je tiens aussi à remercier toute l’équipe de l’UMR PVBMT de m’avoir accueilli dans leur laboratoire. Merci en particulier à Dominique, Johanna, Pascale, Fabien, Lionel, Karine, Denis et Marie-Hélène pour leur aide et leurs encouragements. Je remercie chaleureusement Jacques Fournel, Sébastien Albert, Vincent Florens, Alain Jubault et Patrice Bernet pour la collecte d’échantillons. Merci à Christian Guillermet de m’avoir accompagné lors d’une ‘chasse’ aux pollinisateurs et d’avoir identifié les pollinisateurs à partir des vidéos. Merci aux stagiaires – Sylvain Guichard, Ali M’Dere, Anne-Sarah Padre et Ludivine Sam-Long – pour leur contribution à ce travail. Merci à Alex, Amandine, Audrey, Benoît, Elise, Fred, Gaël, Guillaume, Laurence, Laury, Morgane et Michel pour votre aide sur le terrain. Un grand merci à ceux qui m’ont permis de négocier la dernière ligne droite plus ou moins sereinement en acceptant de réaliser certaines tâches ingrates. Je pense à Stéphanie et Laury pour la relecture du mémoire ainsi qu’à Anne, David et Corto pour la mise en page du document. Un immense merci à mes acolytes réunionnais, normands et épônois pour leur soutien infaillible, leur encouragement et pour tous ces moments qui rendent la vie plus douce. Un énorme merci à ma famille qui m’a toujours soutenu, aidé et encouragé, même lorsque j’ai décidé d’aller m’installer à l’autre bout du monde. J’ai une pensée particulière pour Léonie que je hâte de rencontrer. Merci enfin à ma bretonne préférée de non seulement supporter un normand dans son foyer mais en plus de prendre soin de lui, cette thèse est aussi la tienne. Table des Matières 1 Introduction générale 1 1.1. Cadre théorique: la spéciation écologique .............................................. 3 1.1.1. De Darwin à la définition actuelle de la spéciation écologique .......................... 3 1.1.2. Le continuum de spéciation ............................................................................... 7 1.1.3. Les mécanismes de la spéciation écologique ...................................................... 9 1.1.4. Les facteurs affectant la progression de la spéciation écologique ..................... 11 1.1.5. L’étude de la spéciation écologique ................................................................. 12 1.1.6. L’importance de la spéciation écologique dans la nature ................................. 15 1.1.7. Les mécanismes alternatifs .............................................................................. 16 1.2. La spéciation écologique dans les systèmes insulaires ........................... 19 1.2.1. Intérêt dans l’étude de la spéciation en milieu insulaire .................................. 19 1.2.2. Les exemples connus de spéciation écologique en milieu insulaire ................... 22 1.2.3. Le cas des Mascareignes .................................................................................. 23 1.3. Contexte de l’étude : les orchidées Angraecinae des Mascareignes ........ 28 1.3.1. Biogéographie des Angraecinae aux Mascareignes .......................................... 30 1.3.2. Rôle de l’interaction plante-pollinisateur dans la radiation de la section Hadrangis ........................................................................................................ 31 1.3.3. Le genre Jumellea ...........................................................................................
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