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Yves Desdevises Département De Sciences Biologiques Faculté Des Arts Et Sciences Université De Montréal Et Université De Pe

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Yves Desdevises Département De Sciences Biologiques Faculté Des Arts Et Sciences Université De Montréal Et Université De Pe RECHERCHE DES DÉTÉRMINANTS DE LA SPÉCIFICITÉ PARASITAIRE DANS LE MODÈLE LAMELLODISCUS (DIPLECTANIDAE, MONOGENEA)-SPARIDAE (TELEOSTEI) EN MÉDITERRANÉE Yves Desdevises Département de Sciences Biologiques Faculté des Arts et Sciences Université de Montréal Et Université de Perpignan RECHERCHE DES DÉTERMINANTS DE LA SPÉCIFICITÉ PARASITAIRE DANS LE MODÈLE LAMELLODISCUS (DIPLECTANIDAE, MONOGENEA)-SPARIDAE (TELEOSTEI) EN MÉDITERRANÉE par Yves Desdevises Thèse de doctorat effectuée en co-tutelle au Département de sciences biologiques Faculté des arts et des sciences Université de Montréal ET au Laboratoire de biologie animale Discipline: Biologie Formation Doctorale: Parasitologie et Écologie Parasitaire Université de Perpignan Thèse présentée à la Faculté des études supérieures de l'Université de Montréal en vue de l'obtention du grade de Philosophiae Doctor (Ph.D.) en Sciences Biologiques et à l'École Doctorale de l'Université de Perpignan en vue de l'obtention du grade de Docteur de l'Université de Perpignan en Biologie Octobre 2001 © Yves Desdevises, 2001 ii PAGE D'IDENTIFICATION DU JURY Université de Montréal Faculté des études supérieures et Laboratoire de biologie animale Université de Perpignan Cette thèse intitulée RECHERCHE DES DÉTERMINANTS DE LA SPÉCIFICITÉ PARASITAIRE DANS LE MODÈLE LAMELLODISCUS (DIPLECTANIDAE, MONOGENEA)-SPARIDAE (TELEOSTEI) EN MÉDITERRANÉE présentée et soutenue à l'Université de Perpignan par Yves Desdevises a été évaluée par un jury composé des personnes suivantes Président-rapporteur C. Combes, Professeur, Université de Perpignan, France et membre du jury Directeur de recherche S. Morand, DR CNRS, Université de Perpignan, France Directeur de recherche P. Legendre, Professeur, Université de Montréal, Canada Rapporteur R. D. M. Page, Reader, University of Glasgow, Grande- Bretagne Rapporteur R. Poulin, Reader, University of Otago, Nouvelle-Zélande Examinateur G. Lecointre, Maître de Conférences, Muséum National d'Histoire Naturelle, Paris, France Examinateur J.-P. Féral, DR CNRS, Université Pierre et Marie Curie, Observatoire Océanologique de Banyuls-sur-Mer, France iii RÉSUMÉ L'objectif de cette thèse est de mieux comprendre ce qui contrôle la spécificité parasitaire dans le système hôte-parasite constitué par les poissons de la famille des Sparidae et leurs monogènes (Plathelminthes ectoparasites) spécifiques du genre Lamellodiscus. En d'autres termes, il s'agissait de comprendre les causes amenant une espèce parasite à utiliser une seule ou plusieurs espèces hôtes. On suppose que la spécificité est soumise à des contraintes écologiques et évolutives. L'hypothèse d'une augmentation de la diversité taxonomique avec la spécificité a également été testée. Cela est basé sur l'idée que les espèces généralistes, plus tolérantes à leur environnement, sont supposées être moins sujettes à subir des spéciations après un changement d'hôte, et ainsi être moins diversifiées que les espèces spécialistes. Comme l'histoire évolutive des parasites peut être influencée par celle de leurs hôtes à travers des phénomènes de coévolution, il était nécessaire d'obtenir une phylogénie des hôtes et des parasites. L'étude de la coévolution hôte-parasite dans ce système avait pour but de déterminer si le profil d'association hôte- parasite (et donc la spécificité) est contrôlé par des interactions coévolutives. Des phylogénies ont été élaborées pour les hôtes et les parasites à partir de données moléculaires obtenues par séquençage d'ADN. Cette analyse moléculaire a permis de reconsidérer le statut taxonomique de plusieurs espèces de monogènes : sur la base des séquences obtenues, Lamellodiscus virgula et L. obeliae s'avèrent être une seule espèce (L. virgula), alors que Furnestinia echeneis fait partie du genre Lamellodiscus. Plusieurs méthodes d'étude de la coévolution ont été utilisées dans ce travail. L'une d'elles, ParaFit, a été mise au point pendant la thèse. Toutes les méthodes indiquent que ce système complexe ne semble pratiquement pas être soumis à des phénomènes de cospéciation. Aucun lien entre la diversification taxonomique et la spécificité n'a pu être mis en évidence chez les Lamellodiscus et la famille qui les contient, les Diplectanidae. Par contre, un tel lien a été mis en évidence au niveau des groupes principaux de parasites. Les déterminants écologiques et phylogénétiques de la spécificité ont ensuite été recherchés à l'aide d'analyses statistiques multivariables. Les variables considérées étaient des caractéristiques des hôtes considérées comme des déterminants écologiques potentiels de la spécificité. La phylogénie des parasites a été prise en compte dans ces analyses à l'aide de méthodes iv comparatives, comme la méthode des contrastes indépendants. La spécificité apparaît être fortement contrainte par la phylogénie, ce qui suggére l'existence de déterminants génétiquement transmissibles. Les analyses révèlent également que les parasites spécialistes ont tendance à utiliser les hôtes les plus grands. Cela est interprété comme une spécialisation sur une ressource prédictible. Mots-clés: Spécificité, Monogène, Poisson, Coévolution, Phylogénie moléculaire, Lamellodiscus, Sparidae, Analyse comparative, Diversification taxonomique v ABSTRACT The objective of this thesis is to obtain a better understanding of the factors controlling host specificity in the host-parasite system formed by fish from the family Sparidae and their specific monogeneans parasites from the genus Lamellodiscus (Platyhelminthes). In other words, the goal was to understand the factors determining the number of hosts used by a parasite species. We will assume that specificity is under ecological and evolutionary constraints. The hypothesis of an increase of taxonomic diversity with specificity was also tested. It is based on the idea that generalist species are more tolerant to their environment, and therefore less subject to speciation after a host change event, and are then less diversified than specialist species. Since the evolutionary history of parasites can be influenced by the history of their hosts via coevolutionary interactions, it was necessary to obtain phylogenies for the hosts and parasites. The aim of the study of host-parasite coevolution in this system was to assess if the pattern of host- parasite association (and consequently, specificity) was determined by coevolutionary interactions. Phylogenies were obtained for hosts and parasites from the analysis of DNA sequences. This analysis, carried out at the molecular level, led us to reconsider the taxonomic status of several monogenean species. On the basis of the DNA sequences obtained, Lamellodiscus virgula and L. obeliae appear to form a single species (L. virgula), while Furnestinia echeneis is transferred to the genus Lamellodiscus. Several analytical methods were used to study host-parasite coevolution in this system. Among them, ParaFit was designed during this thesis. All methods agreed that this host-parasite system does not exhibit a general cospeciation pattern. No link between taxonomic diversity and specificity has been found in Lamellodiscus, nor in their family, the Diplectanidae. However, such a link was found when the main groups of parasites were considered. Ecological and phylogenetic determinants of specificity were investigated via multivariate statistical methods. The variables included in the analyses were potential host-related ecological determinants of specificity. The parasite phylogeny was taken into account through comparative methods, including the independent contrasts method. Specificity appears to be strongly constrained by the phylogeny, suggesting the existence of genetically transmitted determinants. The analyses also revealed that Lamellodiscus monogeneans tend vi to specialize on larger hosts. This is interpreted as a specialization on a predictable resource. Key-Words: Specificity, Monogenean, Fish, Coevolution, Molecular Phylogeny, Lamellodiscus, Sparidae, Comparative Analysis, Taxonomic Diversification vii TABLE DES MATIÈRES PAGE D'IDENTIFICATION DU JURY ........................................................ II RÉSUMÉ.................................................................................III ABSTRACT................................................................................V TABLE DES MATIÈRES ..................................................................... I LISTE DES FIGURES.......................................................................IX LISTE DES TABLEAUX.....................................................................XI REMERCIEMENTS ...................................................................... XIII I. INTRODUCTION .......................................................................1 A. PROBLÉMATIQUE ......................................................................2 1. Contexte général....................................................................2 2. Cas des relations hôte-parasite......................................................4 a. La spécificité....................................................................................... 5 i. Définition.......................................................................................................................... 5 ii. Déterminants potentiels....................................................................................................... 7 b. La coévolution hôte-parasite.................................................................... 10 3. Le cas des monogènes............................................................
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