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Mikaël BILI Préparée À L’UMR 1349 « IGEPP » Institut De Génétique, Environnement Et Protection Des Plantes UFR Sciences De La Vie Et De L’Environnement

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Mikaël BILI Préparée À L’UMR 1349 « IGEPP » Institut De Génétique, Environnement Et Protection Des Plantes UFR Sciences De La Vie Et De L’Environnement ANNÉE 2014 THÈSE / UNIVERSITÉ DE RENNES 1 sous le sceau de l’Université Européenne de Bretagne pour le grade de DOCTEUR DE L’UNIVERSITÉ DE RENNES 1 Mention : Biologie Ecole doctorale Vie – Agro – Santé présentée par Mikaël BILI Préparée à l’UMR 1349 « IGEPP » Institut de Génétique, Environnement et Protection des Plantes UFR Sciences de la Vie et de l’Environnement Eléments de Thèse soutenue à Rennes le18 décembre 2014 différenciation de la devant le jury composé de : Didier BOUCHON niche écologique chez Professeur, Université de Poitiers/rapporteur deux coléoptères Emmanuel DESOUHANT Professeur,Université de Lyon 1 / rapporteur parasitoïdes en Geneviève PREVOST Professeur,Université Picardie Jules Verne / compétition : examinateur Cécile LE LANN Maître de Conférences,Université de Rennes 1/ comportement et examinateur communautés Denis POINSOT Maître de Conférences,Université de Rennes 1/ directeur de thèse bactériennes. Anne Marie CORTESERO Professeur, Université de Rennes 1 / co-directrice de thèse Remerciements Après trois années en thèse, comme tout doctorant, il y a un grand nombre de personnes que j'aimerais remercier tant au sein de l'UMR IGEPP que parmi les gens qui m'épaulent (i. e. "me supportent") depuis plus longtemps. Mais tout d’abord je tiens à remercier sincèrement Didier Bouchon, Emmanuel Desouhant, Geneviève Prevost et Cécile Le Lann pour avoir accepté sans hésiter de prendre le temps et de se déplacer à Rennes (avec plus ou moins de difficultés) afin de juger ce travail. Votre intérêt et vos questions m’ont permis de savourer pleinement la défense de cette thèse. Je n'oublierai pas mes encadrants, Denis Poinsot et Anne Marie Cortesero. Je vous remercie de tout cœur pour ces trois années partagées et pour les précieux conseils prodigués toujours avec humour et bienveillance. Vous avez su faire preuve de patience tout au long de cette aventure et en particulier pendant la rédaction de ce manuscrit. Votre soutien et votre attention ont été essentiels pour me permettre de réussir cette étape de mon parcours professionnel et je vous en serai éternellement reconnaissant. Je n’oublierai pas non plus les membres de mes deux comités de thèse : Gwennola Ermel, Marlène Goubault, Yannick Outreman, Sylvain Fournet, Christophe Mougel et « mon tuteur de thèse » David Renault. Merci infiniment d’avoir porté autant d’intérêt à mon travail et d’avoir confronté mes projets à leur faisabilité. Vos conseils ont également été essentiels dans le développement de ce travail. Je voudrais également remercier les personnes avec qui j’ai vécu au quotidien au sein de l’ex « Laboratoire d’écophysiologies des insectes parasitoïdes ». Les chercheurs : Lilianne Krespi, Dennis Webb et Sebastien Dugravot. Vos conseils et votre humour ont toujours été d’un grand soutien. Les techniciennes : Valérie, Chrystelle, Maryline et Sonia. Grâce a vos petites attentions et votre présence, j’ai vraiment apprécié de venir travailler au quotidien. Les doctorants : Céline, toujours à la bourre, Alan, mon « collègue de bureau », Maxime, toujours d’un grand soutien pour les statistiques et c’est la moindre des tes qualités et Fabrice avec qui on peut se « couper de la thèse » en à peine deux jours. Les stagiaires, ceux que j’ai encadré : Lucile, Nicolas, Aurélie, Bénédicte et Florent et aussi les autres pour leur v bonne humeur et leur humour dans des conditions de travail parfois pas très marrantes. Enfin, un merci tout particulier à Jeff pour ses visites qui nous ont fait voyager sans bouger du labo. Je tiens à remercier également l’ensemble des personnes que j’ai eu la chance de rencontrer au sein de l’UMR IGEPP. Citer l’ensemble des personnes de l’unité de recherche que j’ai rencontré au cours de cette thèse serait fastidieux et surtout dangereux. Par peur d’oublier des noms, je me contenterai de tous vous remercier globalement pour votre accueil, les moments que nous avons partagé pendant une présentation, une réunion d’équipe, un repas ou pendant une manip. Que vous soyez chercheurs, personnel technique ou doctorant, ces moments resteront gravés dans ma mémoire. Je tiens tout de même à remercier particulièrement Denis Tagu pour sa bienveillance avec les doctorants intégrés à son institut de recherche et Jean Christophe Simon pour avoir su animer les réunions d’équipe toujours avec beaucoup d’humour même quand elles commençaient à trainer en longueur et pour m’avoir accordé du temps lorsque j’avais des difficultés pour analyser les résultats que j’obtenais. Enfin, j’aimerais remercier mon entourage, ma famille, qui m’a toujours soutenu dans mes choix de poursuivre sans fin mes études (ah bah si c’est fini!), ma belle-famille, pour leur soutien (notamment lors de la défense de la thèse) et mes amis pour m’avoir soutenu même lorsque j’ai eu moins de temps à partager avec eux. Enfin je clôturerai ces remerciement par ma moitié, France, qui m’a toujours encouragé tout au long de ce travail, qui m’a soutenu dans les moments de doutes et qui par ses propres déboires de thèse m’a permis de me considérer chanceux lorsque tout allait bien. vi Résumé Lorsque deux espèces exploitent la même niche écologique, elles entrent en compétition pour l'accès aux ressources. Or, un accès limité aux ressources réduit la fitness des individus. La compétition interspécifique va donc agir comme une pression de sélection qui peut mener à des modifications physiologiques ou comportementales pour partager les ressources, car si elles ne sont pas partagées la compétition entraînera le déplacement ou la disparition d'une des deux espèces.Aleochara bilineata et A. bipustulata sont deux coléoptères staphylins parasitoïdes qui s'attaquent à la mouche du chou Delia radicum. Elles ont des paramètres biologiques différents, notamment au niveau des traits d'histoire de vie (qui semblent avantager A. bipustulata) et du spectre d'hôtes (plus généraliste chez A. bipustulata). Ces deux espèces partagent cependant la même stratégie d'exploitation des hôtes et présentent l'originalité que la femelle pond ses œufs à proximité des hôtes et non à l'intérieur, ce qui les distingue deshyménoptères parasitoïdes qui font l'objet de nombreuses études. La larve Aleocharade premier stade est donc mobile et doit trouver et sélectionner elle même un hôte pour s'y développer. Il y a ainsi des possibilités d'adaptations comportementales à la compétition à la fois pour les adultes et les larves de premier stade. Dans ce projet de thèse, nous avons donc choisi d'explorer la niche écologique de ces deux espèces de façon originale en étudiant les modifications comportementales induites par la présence de compétiteurs à la fois chez les femelles adultes et les larves de premier stade. Nous avons également identifié les communautés bactériennes associées aux deux espèces en compétition mais aussi à leur hôte D. radicum et à un autre compétiteur parasitoïde, l'hyménoptère Trybliographa rapae,dans le but d'étudier ultérieurement les impacts des différents partenaires bactériens sur la niche écologique des deux espèces de coléoptères en compétition. Nos résultats montrent que les femelles de l'espèce spécialiste A. bilineata adaptent leurs comportements aux compétiteurs qu'elles rencontrent et sélectionnent les sites de ponte présentant les meilleures chances de succès parasitaire pour leurs larves. Par ailleurs, les larves de premier stade de l'espèce spécialiste dominent largement la compétition larvaire lorsqu'elles sont en compétition avec les larves de l'espèce généraliste. Enfin, les communautés bactériennes des deux espèces de coléoptères sont plus proches entre elles qu'avec les autres membres du réseau trophique étudiés mais comportent des différences à explorer. Ces résultats sont discutés dans le cadre de l'adaptation des choix comportementaux des individus d'une espèce spécialiste à la présence de compétiteurs généralistes et de la coexistence de ces deux espèces dans le milieu naturel. Mots clés : niche écologique, compétition, modifications comportementales, adaptation, communautés bactériennes, réseau trophique, coexistence, Aleochara, coléoptère parasitoïde. vii viii Abstract When two species live in the same ecological niche, they compete for resources. Sincea limited access to resources reduces fitness, interspecific competition represents a selection pressure that can lead to physiological or behavioral changes to share resources, because not sharing themwill cause the displacement or disappearance of the weaker competitor. Aleochara bilineata and Aleochara bipustulata are two coleopteran parasitoids and attack the same host, the cabbage root fly Delia radicum. These two species have different biological parameters, particularly in their life history traits (which seem to favor A. bipustulata) and host spectrum (A. bipustulatais more generalist). These two species share the same strategy to exploit their host (idiobiont ectoparasitoid). Unlike parasitoid wasps (the object of most studies on parasitoids) coleopteran parasitoid females do not lay their eggs directly inside the host but in locations likely to harbour hosts. Aleochara first instars are mobile and need to find and select a host where they will develop. There is thus the possibility of behavioral adaptations to competition for both for adults and first instars. In this project, we have chosen to explore the ecological niche of these two species in an original way by studying behavioral changes induced by the presence of competitors both in adult females and first instars. We also studied bacterial communities associated to the two competing species but also those of their host D. radicum and of another competitor, the parasitoid wasp Trybliographa rapae, in order to later study the impacts of different bacterial partners in the ecological niche of the two beetle species in competition. Our results show that females of the specialist A. bilineata adapt their behavior to the competitors they face and select oviposition sites with the best probability of parasitism success. Moreover, first instars of A. bilineata dominate the larval competition when competing with larvae of the generalist A.
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