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THÈSE Docteur L'institut Des Sciences Et Industries Du Vivant Et De L N° /__/__/__/__/__/__/__/__/__/__/ THÈSE pour obtenir le grade de Docteur de l’Institut des Sciences et Industries du Vivant et de l’Environnement (Agro Paris Tech) Spécialité : Biologie de l’Evolution et Ecologie présentée et soutenue publiquement par ROY Lise le 11 septembre 2009 11 septembre 2009 ECOLOGIE EVOLUTIVE D’UN GENRE D’ACARIEN HEMATOPHAGE : APPROCHE PHYLOGENETIQUE DES DELIMITATIONS INTERSPECIFIQUES ET CARACTERISATION COMPARATIVE DES POPULATIONS DE CINQ ESPECES DU GENRE DERMANYSSUS (ACARI : MESOSTIGMATA) Directeur de thèse : Claude Marie CHAUVE Codirecteur de thèse : Thierry BURONFOSSE Travail réalisé : Ecole Nationale Vétérinaire de Lyon, Laboratoire de Parasitologie et Maladies parasitaires, F-69280 Marcy-L’Etoile Devant le jury : M. Jacques GUILLOT, PR, Ecole Nationale Vétérinaire de Maisons-Alfort (ENVA).…………...Président M. Mark MARAUN, PD, J.F. Blumenbach Institute of Zoology and Anthropology...…………...Rapporteur Mme Maria NAVAJAS, DR, Institut National de la Recherche Agronomique (INRA)..………... Rapporteur M. Roland ALLEMAND, CR, Centre national de la recherche scientifique (CNRS).……………Examinateur M. Thierry BOURGOIN, PR, Muséum National d’Histoire Naturelle (MNHN)......….... ………….Examinateur M. Thierry BURONFOSSE, MC, Ecole Nationale Vétérinaire de Lyon (ENVL)...……………..… Examinateur Mme Claude Marie CHAUVE, PR, Ecole Nationale Vétérinaire de Lyon (ENVL)…...………….. Examinateur L’Institut des Sciences et Industries du Vivant et de l’Environnement (Agro Paris Tech) est un Grand Etablissement dépendant du Ministère de l’Agriculture et de la Pêche, composé de l’INA PG, de l’ENGREF et de l’ENSIA (décret n° 2006-1592 du 13 décembre 2006) Résumé Les acariens microprédateurs du genre Dermanyssus (espèces du groupe gallinae), inféodés aux oiseaux, représentent un modèle pour l'étude d'association lâche particulièrement intéressant : ces arthropodes aptères font partie intégrante du microécosystème du nid (repas de sang aussi rapide que celui du moustique) et leurs hôtes sont ailés. En outre, D. gallinae est une espèce d'importance économique, ce qui rend possible des comparaisons entre colonisation de milieux anthropisés et sauvages. Au début de l'étude, les espèces du groupe gallinae sont très mal délimitées. Les caractères morphologiques utilisés sont variables au sein de l'espèce, voire de la population, très chevauchants entre espèces. Afin de mieux comprendre les exigences écologiques du développement de D. gallinae et d'appréhender ses voies de dissémination, une investigation comparative basée sur des séquences d’ADN entre espèces du groupe gallinae a été adoptée. Un cheminement d'ordre taxinomique a permis de poser les bases nécessaires. Ensuite, l'exploration de certaines caractéristiques écologiques du groupe gallinae en relation avec sa phylogénie (spécificité d'hôte, flexibilité évolutive) a été menée à bien. Une espèce a été décrite, D. apodis, une lignée de D. gallinae constitue aussi une probable espèce inédite et D. longipes regroupe deux entités. Des différences écologiques marquées entre D. gallinae et les autres espèces semblent résulter non seulement de l'activité humaine, mais aussi de caractéristiques intrinsèques. Aujourd'hui, le rôle des flux commerciaux dans la dispersion de D. gallinae en élevage de pondeuses s'avère primordial, au moins en France, celui des oiseaux sauvages presque nul. Mots-clés Système hôte-parasite, microprédateur, phylogénie, espèce, population, microhabitat, Dermanyssus, mésostigmate, acarien 1 Evolutionary ecology of a hematophagous mite genus: Phylogeny-based approach for interspecific delineations and comparative characterization of populations in five species of the genus Dermanyssus (Acari: Mesostigmata) Abstract Micropredator species of Dermanyssus (Moss'gallinae-group), which parasitize birds, represent an interesting model for the study of loose associations. Thus, these unwinged arthropods do not stay on host (blood meal as quick as mosquitoe's), are part of the nest's microecosystem and their hosts are winged. Moreover, micropredator Dermanyssus include at least one species of economic importance in fowl farms, D. gallinae (the Poultry Red Mite), which enables the comparison between species restricted to wild avifauna and synanthropic species. At the beginning of the study, micropredator species are not clearly delimited. Most of species specific morphological characters are variable within species, in some cases within population, and are overlapping between species. In the aim to investigate the ecological needs for proliferation in D. gallinae and its ways for dispersal, a DNA-based comparative analysis involving this species and its close relatives has been performed. The first section consists of the clearing of the taxonomy and species delineations. In the second section, ecological and intrinsic data (host specificity, flexibility) are compared between species of the gallinae-group. One species has been described (D. apodis), one lineage within D. gallinae seems to represent a cryptic species and D. longipes currently groups two different entities. Important ecological differences between D. gallinae and other species seem to result not only from human activities, but also from intrinsic characteristics. Currently, the role of trade flows in D. gallinae’s spread in layer farms appears to be essential, at least in France, as opposed to the role of wild birds (nearly nul). Key words Host-parasite system, micropredator, phylogeny, species, population, microhabitat, Dermanyssus, Mesostigmata, Acari 2 Remerciements Je tiens à remercier du fond du cœur mes deux directeurs de thèse, à qui je dois, c’est certain, beaucoup plus qu’il n’est commun de devoir à ses directeurs de thèse. Merci à Claude Chauve, pour le soutien et la confiance qu’elle m’a admirablement prodigués tout au long de cette thèse. Je ne sais comment exprimer ma reconnaissance et mon respect pour son courage et sa loyauté remarquables. Merci à Thierry Buronfosse, pour le soutien, la confiance et la patience dont il m’a fait bénéficier au cours de cette thèse ! Nos continuelles oppositions, contradictions et confrontations ont sûrement été la clé du présent travail. Plus qu'un encadrant, c’est un collègue, voire un ami, que je crois avoir trouvé en lui, qui m'a soutenue et aidée au-delà de toute espérance. Et il a réussi à former une « prof. de Lettres » à la biologie moléculaire ! J'aimerais que cette thèse puisse constituer un remerciement suffisant à l’appui si solide, si bénéfique et sans cesse renouvelé dont ils ont tous deux fait preuve. J'adresse mes vifs remerciements à Maria Navajas et Mark Maraun pour avoir accepté d’endosser l’important rôle de rapporteur. Je remercie cordialement Roland Allemand, Thierry Bourgoin et Jacques Guillot pour avoir accepté de prendre part au jury de cette thèse. Je remercie aussi les deux premiers pour leur suivi dans le cadre du comité de pilotage. Merci à l’ensemble de mes collègues de travail, passés ou présents, en particulier mes collègues de laboratoire qui travaillent ou ont travaillé sur Dermanyssus : Claire Valiente Moro, Sophie Desloire, Marie-Thérèse Poirel. Une pensée émue à Marie-Claude Reynaud. Merci à mes valeureux stagiaires, qui ont subi mon obsession du genre Dermanyssus au long de stages de durée variable, ont, pour la plupart, fourni un travail particulièrement satisfaisant et ont ainsi pleinement participé au travail de recherche que voici : Robin Bellon, Sandrine Bonnet, Jean Filippi, Mathieu Galès, Mehdi Gharbi, Nina Guichard, Guillaume Lallemand, Sophie Merlin, Sabrina El Ouartiti, Anthony Piron, Marie Rigaux. Merci à l’ensemble des personnes qui ont contribué à l’obtention des indispensables échantillons biologiques : Olivier et Nathalie Chovet, le Centre de Soins aux Oiseaux Sauvages du Lyonnais, notamment Pascal Tavernier, Mathilde, l’Ecopôle du Forez, notamment François Boléat, Claire Brucy, Pascal, …., la Réserve du Romelëare, le Centre de Recherche sur la Biologie et les Populations d’Oiseaux, notamment Gérard Gory, Olivier Caparros, Laurent Brucy, Laurent Carrier, Nicolas Vincent-Martin, Yves Beauvallet, Olivier Dehorter, Franck Salmon, Rolf Wahl, J. Girard-Claudon,… En particulier, merci à Olivier Caparros pour m’avoir accueillie au cours de ses passionnantes séances de baguage d’oiseaux sauvages, et m’avoir instruite quant à la biologie et les traits d’histoire de vie de ces animaux fascinants. 3 Merci à l’Association Ornithologique Rhodanienne, en particulier M. André Marchal et M. Marc Serre, pour m’avoir permis d’obtenir des échantillons provenant d’élevages de petits oiseaux de compagnie et/ou de compétition. J’adresse également mes sincères remerciements à M. Jean- Jacques Sage, président du Club Français du Montauban et M. S. Calabro, président de l’Amicale Ornitholoqique Becs Crochus Centre Est. Merci à Christine Basset pour avoir pris le temps de m’accompagner et m’avoir ainsi si souvent permis d’entrer dans tous ces élevages de volailles en Bresse, pour sa sympathie et sa bonne volonté permanentes. Je remercie également le Dr Patrick Chabrol, Jérome Arnal, … , pour leur aide aux prélèvements dans les élevages de pondeuses. Merci à Sophie Lubac pour son aide dans la collecte d’acariens provenant d’élevages de volailles et les informations concernant la filière pondeuse. Son implication, son soutien et son dynamisme ont fortement contribué à une bonne partie de l’étude. Merci au PEP Avicole (Pôle d’Expérimentation et de Progrès, Région Rhône-Alpes) pour l’appui financier sans lequel l’étude, entamée au cours de
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