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Ecology and Evolution of the Sexual and Asexual Timema Stick Insects

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Ecology and Evolution of the Sexual and Asexual Timema Stick Insects Unicentre CH-1015 Lausanne http://serval.unil.ch RRRRYear : 2018 ECOLOGY AND EVOLUTION OF THE SEXUAL AND ASEXUAL TIMEMA STICK INSECTS Larose Chloé Larose Chloé, 2018, ECOLOGY AND EVOLUTION OF THE SEXUAL AND ASEXUAL TIMEMA STICK INSECTS Originally published at : Thesis, University of Lausanne Posted at the University of Lausanne Open Archive http://serval.unil.ch Document URN : urn:nbn:ch:serval-BIB_A0F0E91546761 Droits d’auteur L'Université de Lausanne attire expressément l'attention des utilisateurs sur le fait que tous les documents publiés dans l'Archive SERVAL sont protégés par le droit d'auteur, conformément à la loi fédérale sur le droit d'auteur et les droits voisins (LDA). A ce titre, il est indispensable d'obtenir le consentement préalable de l'auteur et/ou de l’éditeur avant toute utilisation d'une oeuvre ou d'une partie d'une oeuvre ne relevant pas d'une utilisation à des fins personnelles au sens de la LDA (art. 19, al. 1 lettre a). A défaut, tout contrevenant s'expose aux sanctions prévues par cette loi. Nous déclinons toute responsabilité en la matière. Copyright The University of Lausanne expressly draws the attention of users to the fact that all documents published in the SERVAL Archive are protected by copyright in accordance with federal law on copyright and similar rights (LDA). Accordingly it is indispensable to obtain prior consent from the author and/or publisher before any use of a work or part of a work for purposes other than personal use within the meaning of LDA (art. 19, para. 1 letter a). Failure to do so will expose offenders to the sanctions laid down by this law. We accept no liability in this respect. Département d’écologie et d’évolution ECOLOGY AND EVOLUTION OF THE SEXUAL AND ASEXUAL TIMEMA STICK INSECTS Thèse de doctorat ès sciences de la vie (PhD) présentée à la Faculté de biologie et de médecine de l’Université de Lausanne par Chloé LAROSE Biologiste diplômée du Master de l’Université de Rennes 1, France Jury Prof. Ron Stoop, Président Prof. Tanja Schwander, Directeur de thèse Prof. John Pannell, expert Prof. Hanna Kokko, expert Lausanne 2018 Département d’écologie et d’évolution Ecology and evolution of the sexual and asexual Timema stick insects Thèse de doctorat ès sciences de la vie (PhD) présentée à la Faculté de biologie et de médecine de l’Université de Lausanne par Chloé LAROSE Biologiste diplômée du Master de l’Université de Rennes 1, France Jury Prof. Ron Stoop, president Prof. Tanja Schwander, thesis director Prof. John Pannell, expert Prof. Hanna Kokko, expert Lausanne 2018 REMERCIEMENTS Tanja Schwander, merci d’avoir partagé avec moi ces projets peuplés de Timema par milliers (et parsemés de quelques grillons). De m’avoir accueillie comme première doctorante de ta nouvelle vie de Professeur. Et d’avoir partagé avec moi quelques bribes de ton exigence, ta reflexion, et ton excellence scientifique. John Pannell, Thomas Lenormand et Hanna Kokko, merci d’avoir accepté d’être les trois experts de ma thèse, je ne pouvais pas espérer meilleurs experts pour se pencher sur mon travail, et le critiquer. Sergio Rasmann et Darren Parker, qui, à l’exception de mes précédents superviseurs Solenn Stoeckel, Claude Rispe et Tanja, ont été les premières personnes avec qui j’ai eu la chance de vraiment collaborer et travailler. Merci, parce que maintenant je sais à quel point ‘collaborer’ est agréable ..et essentiel. Je souhaite et suis heureuse de pouvoir enfin mentionner et remercier : Catherine Berney pour de nombreuses petites aides ponctuelles tout au long de ces 5 ans, Amaury Avril pour tes nombreux conseils et explications toujours très utiles, précis, efficaces. Dessislava Petrova pour ton aide avec ces amplifications compliquées de gènes phasmiques, Nicolas Rush et Aline Revel pour m’avoir secondé dans le jardinage de mes plantes californiennes, Marco Zini, un jour tu m’as sorti du pétrin. Frank Chalard et tous tes bricolages avisés, Eléonore Genzoni pour tes soins très attentionnés aux Timema exilés et tout chamboulés après toutes ces heures d’avion. Thanks Armand Yazdani my 1st field assistant. Casper van der Kooi pour ton regard respectueux et avisé sur mes écrits, Léna Kolecek pour ta grande patience face à toutes sortes d’étrangetés administratives intempestives. Karine Légeret pour ton aide immense à la toute fin. Merci Tom Richardson, toujours, sans jamais osciller, tout au long de ces cinq années, planté comme une fougère dans mon dos, j’ai apprécié ton existence dans mes alentours, cher office mate. Merci Valentine Rech De Laval, pour ton amitié, ton soutien, ta gentillesse et sérénité, au détour d’un couloir ou d’un petit café. Merci Maud Liegeois d’avoir débarqué au beau milieu de ce doctorat, comme un petit cheveu sur la soupe, ou une éphémère sur la rivière. Merci Raphael Braunschweig, soutien, duo de choc, équipe de galère, ami intense et fou. Je veux remercier tout particulièrement le soutien très précieux, et teinté d’amitié, que tu m‘as apporté, Nathalie Caloz, acolyte inattendue et essentielle, durant ces années, tu as été parfaite. Je remercie, très fort, mon ami Simon Power, merci pour tout, mais avant toute chose pour cette amitié que nous avons réussi à faire germer, puis fleurir ! Encore tellement de bons moments à venir…monsieur le docteur lord professeur ami génial et for ever. Un merci immense à Loren, pour tout. Tout. Et vraiment tout. Sans oublier les plous, les plis et les plas. Merci mes amis ici, Danielle, Lucie, Amaranta, Jonathan (d’ailleurs il faut que je te raconte…..eh eh), Prof Elsa et aussi à ton Mark avec un k. Et très particulièrement, merci Sacha et Yoann, rencontres belles. Sans vous j’étais foutue. Merci mes amies là-bas, Sonia, Maitena, Marie, Poline, Graciela. Que j’aime. Merci à Bella, ma maman, unique, surtout pour avoir supporté cette étape avec moi, de très loin, mais toujours en réussissant à tolérer mes choix, et à ne pas disparaitre. Alors ….à la passion, à la vie, à la folie, à l’enthousiasme. A la découverte, à la nature. A la créativité, à la poésie, à l’art. A l’intelligence. Au surpassement. Aux choix et décisions que l’on arrive enfin à prendre. Et bien sûr, à la beauté…qui parsème cet univers. Chloé. ‘’ Distance is a relative notion You think you’ve filled the void you reached But you finally realize The more you dig the less you sink The less you stop the more you start You focus on the goal But once you’re close enough If only everything was slightly different.. You feel like getting closer to the edge But you don’t know on which side you stand The beginning might just be the end A point of view could never be shared ’’ I.D A B S T R A C T ‘Reproduction’ is one of the key characteristic of life. Despite this, our knowledge of the evolution of reproductive systems is still incomplete. In particular, the reasons for why the vast majority of eukaryotes use sex, and thus take a complicated and costly detour to reproduction, when straightforward routes, such asexuality, are available, remains a central and largely unanswered question in evolutionary biology. The aim of my thesis is to contribute to the understanding of this evolutionary mystery, and for that I use stick insects of the genus Timema as a study system. This small group of herbivorous insects, endemic to Western United States is ideal for studying and comparing sexual and asexual reproduction as seven asexual lineages have been identified in this group, each with a sexual sister species, allowing us to make multiple independent comparisons between sexual and asexual lineages. The perhaps most broadly accepted theoretical argument is that sex allows selection to work efficiently, which would ultimately favor the adaptive potential of populations. My objective during this thesis was to test two theories directly related to this, but working each time in two successive steps: i) I started by clarifying the ecological and evolutionary aspects and mecanisms concerned by these theories in Timema focusing only on sexual species and thus independently of the reproductive mode, ii) and I then empirically tested these theories. Specifically, I first investigated whether sexuals are able to exploit more ecological niches than asexuals, which would give them an advantage in fluctuating or heterogeneous environments. From this first investigation, I overall found that sexual species are systematically using a larger portion of their environment than their asexual relatives, but I did not find this pattern regarding their intrinsic and physiologic abilities to use their environment. The reduced portion used by asexuals is thus likely a consequence of external and biotic interactions that affect asexuals more strongly than sexuals. I secondly aimed to empirically test if sex confer an advantage when the allele combinations that are favored by selection vary over time, as it is the case in context of coevolution with parasites. My work suggests that parasites are indeed contributing to the maintenance of sex in Timema. In the last part of the thesis, I finally present some preliminary results regarding new Timema populations that I discovered by chance, that feature unusual reproductive strategies with a mixture of sexual, asexual ad facultatively asexual individuals. These populations will be very profitable for future research concerning the evolution of reproduction in Timema. Overall this thesis work contributes to a better understanding of several aspects of the ecology and evolution of Timema stick insects in partiular, and more generally contribute to give novel insights in the understanding of the maintenance of sex in the living world. 9 R E S U M É F R A N Ç A I S L’une des caractéristiques essentielles d’un être vivant est sa capacité à se reproduire.
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