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Cataglyphis Cursor Université Paris VI - Pierre et Marie Curie Thèse Présentée pour l’obtention du grade de Docteur de l’Université Paris VI Discipline : Écologie et Biologie Évolutive Evolution des stratégies de dispersion et de reproduction chez la fourmi Cataglyphis cursor Par Johanna CLEMENCET Soutenue le 17 Mai 2006 devant le jury composé de : Robert BARBAULT Professeur……………………………………………. Président (Université de Paris VI, France) Jürgen HEINZE Professeur…………………………………………….Rapporteur (Universität Regensburg, Allemagne) Frédérique VIARD Chargée de recherche............................................Rapporteur (Roscoff, France) Alain LENOIR Professeur…………………………………………….Examinateur (Université de Tours, France) Claudie DOUMS Maître de conférences………………………………Directrice de (EPHE, Université de Paris VI, France) Thèse Christian PEETERS Directeur de recherche……………………………..Directeur de (CNRS, Université de Paris VI, France) Thèse Laboratoire Fonctionnement et Évolution des Systèmes Écologiques CNRS ENS UMR 7625 École Doctorale Diversité du Vivant Université Paris VI - Pierre et Marie Curie Thèse Présentée pour l’obtention du grade de Docteur de l’Université Paris VI Discipline : Écologie et Biologie Évolutive Evolution des stratégies de dispersion et de reproduction chez la fourmi Cataglyphis cursor Par Johanna CLEMENCET Soutenue le 17 Mai 2006 devant le jury composé de : Robert BARBAULT Professeur……………………………………………. Président (Université de Paris VI, France) Jürgen HEINZE Professeur…………………………………………….Rapporteur (Universität Regensburg, Allemagne) Frédérique VIARD Chargée de recherche............................................Rapporteur (Roscoff, France) Alain LENOIR Professeur…………………………………………….Examinateur (Université de Tours, France) Claudie DOUMS Maître de conférences………………………………Directrice de (EPHE, Université de Paris VI, France) Thèse Christian PEETERS Directeur de recherche……………………………..Directeur de (CNRS, Université de Paris VI, France) Thèse Laboratoire Fonctionnement et Évolution des Systèmes Écologiques CNRS ENS UMR 7625 École Doctorale Diversité du Vivant 1 RÉSUMÉ Chez les insectes sociaux, les stratégies de dispersion et de reproduction ont un rôle majeur dans l’évolution des actes altruistes puisqu’elles déterminent l’apparentement entre individus au sein et entre les colonies. Le but de ce travail est d’explorer les stratégies de dispersion et de reproduction, a priori exceptionnelles, de la fourmi Cataglyphis cursor. Notre étude de génétique des populations, à l’aide de marqueurs nucléaires (microsatellites) et mitochondriaux (gène COI), supporte l’idée que les colonies se reproduisent par fission chez cette espèce monogyne et souligne l’influence de l’habitat sur la distribution de la variabilité génétique à fine échelle. Une étude socio-génétique menée dans deux populations issues de différents habitats a confirmé que les reines utilisent la parthénogenèse thélytoque pour produire les nouvelles reines, mais s’accouplent avec plusieurs mâles pour produire les ouvrières. Nous montrons, pour la première fois chez les fourmis, qu’à tailles égales, les colonies les plus diverses génétiquement ont en moyenne des ouvrières de plus grande taille. Dans plusieurs populations étudiées, la taille des ouvrières est positivement corrélée avec la taille des colonies. Une augmentation de la taille des ouvrières peut représenter un avantage important chez cette espèce thermophile. Elle permettrait aux colonies de fourrager à des températures difficilement supportables pour les petites ouvrières et les autres espèces. Nous avons montré expérimentalement que la résistance à la dessiccation augmente avec la taille des ouvrières. Toutefois, sur le terrain, les fourrageuses n’étaient pas significativement plus grandes aux heures les plus chaudes de la journée. Dans les colonies orphelines de C. cursor, les ouvrières peuvent également produire des oeufs diploïdes (ouvrières et reines) par parthénogenèse thélytoque. Des observations comportementales ont mis en évidence que le potentiel reproducteur des ouvrières augmente avec leur taille. Ceci souligne que chez C. cursor, les traits phénotypiques des colonies et des ouvrières sont le résultat de pressions de sélections potentiellement divergentes (sélection individuelle vs sélection au niveau de la colonie). Mots-clefs: Insectes sociaux, Cataglyphis cursor (Hymenoptera, Formicidae), dispersion, fondation dépendante, viscosité des populations, parthénogenèse thélytoque, polyandrie, taille de colonie, taille d’ouvrière, fécondité, dessiccation, niveaux de sélection, microsatellites, ADNmt 2 ABSTRACT In social insect, understanding the reproductive and dispersal strategies are of major interest as they determine the relatedness between interacting individuals (within and among colonies), a parameter with a major influence on the evolution of altruistic behaviour (division of labour, regulation of reproduction). The aim of this work was to explore the peculiar reproductive and dispersal strategies observed in the ant Cataglyphis cursor. A population genetic study of C. cursor populations strongly supported the idea that colonies reproduce by fission in this monogynous species and emphasized the role of habitat in driving the pattern of local population genetic structure. A socio-genetic study conducted in two populations from different habitat types showed that the peculiar reproductive system, previously demonstrated in a single population, appears to be a general situation. Indeed, in both populations, the queens used thelytokous parthenogenesis to produce new queens but mated with several males (5-12) to produce the workers. We found that the more genetically diverse colonies harboured a greater mean worker size after controlling for colony size. In the two populations studied, worker size was indeed positively correlated with colony size. In this thermophilic species, an increase of worker size could be advantageous to the colony by allowing a foraging temperature that small workers and other species can not tolerate. Effectively, we experimentally found an increase of worker thermal tolerance with size. However, in the field, foragers were not significantly larger than inside nest workers. In orphaned colonies of C. cursor, workers can also produce diploid eggs (gynes and workers) by thelytokous parthenogenesis. Behavioural observations of an orphan colony indicated that size is also linked to the reproductive potential of workers, larger workers being more likely to reproduce than smaller. This underlines that colony and individual phenotypic traits in C. cursor are the result of conflicting (individual vs colony level selection) selective pressures. Key words: Social insect, Cataglyphis cursor (Hymenoptera, Formicidae), sex-biased dispersal, dependant colony foundation, population viscosity, thelytokous parthenogenesis, polyandry, colony size, worker size, worker re productive potential, desiccation, level of selection, microsatellites, mtDNA 3 REMERCIEMENTS En premier lieu, je tiens à remercier le professeur Robert Barbault (président du jury), les rapporteurs Jürgen Heinze et Frédérique Viard, les examinateurs Alain Lenoir et Christian Peeters d’avoir accepté de faire partie du jury de ma thèse. Mille remerciements à ma directrice de thèse Claudie Doums pour la qualité de son encadrement scientifique et humain. Son dynamisme, sa disponibilité, sa détermination, sa confiance, son soutien, ses encouragements, sa patience …. (et ses « figolus » pour attirer les fourmis !) ont largement contribué à la réalisation et à l’aboutissement de cette thèse. Je tiens à remercier chaleureusement les personnes qui ont aidé à la récolte des données sur le terrain ou au laboratoire : - Barbara Viginier pour le travail de développement des marqueurs mitochondriaux. Merci pour m’avoir initiée aux techniques de biologie moléculaire au cours de mon DEA et d’avoir avec enthousiasme participé aux missions de terrain. Merci pour les petits plats après les longues journées de terrain, un véritable régal ! - Laurent Cournault pour avoir toujours répondu présent pour les missions de terrain, dans des conditions parfois difficilement supportables (vive l’écran total !), pour sa détermination, son initiation à l’entomologie et ses discussions sans fin. - Thibaud Monnin d’avoir accepté de participer à la première mission de récolte en « rollers », pour les nombreuses discussions scientifiques et les relectures de manuscrits. - Lionel Brazier et Raphaël Sylvain pour leur participation à la mission de terrain la plus chaude de tous les temps. - Aurélien Odent, Alexandra Mélé, Annabelle Leblond et Aurélie Bessière pour leur travail au laboratoire et leur gentillesse. - Quentin Rome pour sa patience qui a permis d’obtenir une énorme quantité de données comportementales sur la reproduction dans une colonie orpheline de Cataglyphis Cursor. - Pierre Fédérici pour son aide dans le travail de biologie moléculaire et ses nombreux conseils. - Claude Doums pour son accueil chaleureux dans sa maison des Corbières, ses omelettes aux asperges sauvages, ses pronostics météorologiques… 4 Merci à toute l’équipe « fourmis » pour les nombreuses discussions scientifiques, et les relectures de manuscrits : Sébastien Barrate, Emmanuelle Baudry, Aurélie Bocher, Else Fjerdingstad (merci pour tous les tuyaux biblio !), David Laloi, Mathieu Mollet, Léa Zinck, Thibaud Monnin et Christian Peeters. Merci aux chercheurs Alain Lenoir et Xim Cerdá qui m’ont apporté de précieuses informations sur l’espèce Cataglyphis cursor. Merci à l’ensemble des membres des
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