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DÉTERMINANTS INDIVIDUELS ET ENVIRONNEMENTAUX DU PARASITISME CHEZ LE TAMIA RAYÉ (TAMIAS STRIATUS) par Chelsey Paquette Mémoire présenté au Département de biologie en vue de l’obtention du grade de maître ès sciences (M.Sc) FACULTÉ DES SCIENCES UNIVERSITÉ DE SHERBROOKE Sherbrooke, Québec, Canada, avril 2020 Le 15 avril 2020 le jury a accepté le mémoire de Madame Chelsey Paquette dans sa version finale Membres du jury Professeur Dany Garant Directeur de recherche Département de Biologie, Université de Sherbrooke Professeur Patrick Bergeron Codirecteur de recherche Département de Biologie, Bishop’s University Professeure Jade Savage Codirectrice de recherche Département de Biologie, Bishop’s University Professeure Fanie Pelletier Évaluatrice interne Département de Biologie, Université de Sherbrooke Professeur Marco Festa-Bianchet Président-rapporteur Département de Biologie, Université de Sherbrooke SOMMAIRE Les parasites peuvent infecter presque tous les organismes vivants et ils peuvent avoir un impact important sur leurs hôtes et sur leur dynamique de population. Cependant, la sévérité de ces impacts peut varier et dépendre de l'hôte, du parasite et de leur environnement commun. Pour mieux comprendre la complexité des interactions hôte-parasite, il est essentiel d’évaluer les déterminants individuels, populationnels et environnementaux du parasitisme. La prévalence de l'infestation peut varier au fil du temps en fonction de caractéristiques populationnelles telles que la densité et la démographie, ainsi que des caractéristiques environnementales telles que les conditions météorologiques et la disponibilité de la nourriture. Cependant, au sein d'une population hôte, le nombre de parasites par individu peut aussi varier en raison de différences individuelles en physiologie, morphologie et/ou comportement. Par conséquent, il est nécessaire d'intégrer les caractéristiques de la population, des individus et de l'environnement dans les études sur les interactions hôte-parasite. Dans ce contexte et en utilisant 12 ans de données provenant d'une population sauvage de tamias rayé (Tamias striatus), j’ai étudié les déterminants du parasitisme de Cuterebra (« bot fly ») au niveau individuel et de la population. Au niveau de la population, j’ai évalué la relation entre la prévalence de l'infestation et les conditions météorologiques, les cycles de paisson (« mast ») du hêtre à grandes feuilles (Fagus grandifolia) et la taille de la population de tamia. Au niveau individuel, j’ai étudié la relation entre l'intensité de l'infestation et les caractéristiques des tamias comme le sexe, l'âge, la masse corporelle et le comportement. La prévalence de l'infestation dans la population était positivement liée à la température printanière, aux précipitations printanières et à la taille de la population de tamias, et négativement liée aux cycles de paisson des hêtres à grandes feuilles. Chez les adultes, l'influence du comportement individuel sur l'intensité de l'infestation variait selon le sexe, ce qui suggère que le comportement spécifique à un sexe peut influencer l'exposition aux parasites. Pour les juvéniles, les résultats montrent i l'importance du moment d'émergence associé à la saison de naissance sur l'exposition aux parasites et sur le nombre de parasites qu'un individu aura. Ces résultats soulignent l'importance des conditions environnementales locales sur les hôtes et les parasites et suggèrent que les caractéristiques individuelles, populationnelles et environnementales qui sont avantageuses pour l'acquisition des ressources et la reproduction peuvent aussi augmenter le niveau de parasitisme. Mots clés : Interaction hôte-parasite, Parasite, Rongeur, Variabilité environnementale, Comportement, Cuterebra bot fly, Tamia rayé ii SUMMARY Parasites can infect almost all living organisms and they may have important impacts on individual hosts, also impacting host population dynamics. However, the severity of these impacts can vary and depend on the host, the parasite and their shared environment. Therefore, to better understand the complexity of host-parasite interactions, we must first evaluate the individual, populational and environmental determinants of parasitism. Populations may vary in infestation prevalence over time due to population characteristics such as density and demography, as well as environmental characteristics such as weather and food availability. However, within a host population, individuals may also vary in the number of parasites they have due to individual differences in physiology, morphology and behavior. Therefore, it is necessary to incorporate population, individual and environmental characteristics in studies on host-parasite interactions. In this context and using 12 years of data from a wild population of eastern chipmunks (Tamias striatus) relying on pulsed food resources, I investigated the determinants of Cuterebra bot fly parasitism at both the population and individual level. First, I assessed the relationship between infestation prevalence and weather conditions, mast cycles of American beech (Fagus grandifolia) trees and chipmunk population size. Then, I investigated the relationship between infestation intensity and chipmunk characteristics such as sex, age, body mass and behavior. Infestation prevalence in the population was positively related to spring temperature, spring precipitation and chipmunk population size, and negatively related to mast cycles of American beech trees. For adults, the influence of individual behavior on infestation intensity varied with sex, suggesting that sex-specific behaviors may influence parasite exposure. For juveniles, the results highlight the importance of parasite exposure and emergence timing on the number of parasites an individual will have. These results emphasize the importance of local environmental conditions on both hosts and parasites, and suggest that the individual, populational and iii environmental characteristics which are advantageous for resource acquisition, reproduction and population growth may also come at the cost of increasing parasitism. Key words: Host-parasite interaction, Parasite, Rodent, Environmental variability, Behavior, Cuterebra bot fly, Eastern chipmunk iv ACKNOWLEDGEMENTS I would like to thank my three research supervisors, Dany Garant, Patrick Bergeron and Jade Savage. I was extremely fortunate to have three tremendous researchers take me under their wings and guide me along my short but sweet journey as a master’s student. Thank you, Dany, for your wisdom, efficiency in everything you do, and for answering all of my questions, even those that were hypothetical. Patrick, for your ability to always provide great advice and for inspiring me as a young biologist. And Jade, for all your support and for fueling my inner entomologist; I was never the only one in the room that was overly fascinated with bot flies. I would also like to thank Denis Réale for his contribution to the Chipmunk Project and his comments on my manuscript, and Marco Festa-Bianchet and Fanie Pelletier, for their feedback on my project and thesis. This project would not have happened without the numerous graduate students and field assistants who have collected data for the Chipmunk Project since its initiation in 2004. After having spent 13 months in the scenic forests of Mansonville, I would like to give a special thank you to Elouana, for teaching me everything there is to know about the field (and chipmunks) and Katherine, for being by my side throughout it all. Thank you also to Edward (L094) for being nothing like the Revenant (J088). To my lab mates, thank you for your support, feedback and wide ranging/interesting conversations. More specifically, thank you Philippine and Caro for being incredible mentors. I would like to thank all of my family and friends for providing me with endless support and encouragement throughout all of my studies, especially mom, Lyssa and Allen. Finally, to my father, thank you for always letting a ray of sunshine peak through the forest canopy when things were not going as planned. This project was supported financially by the Natural Sciences and Engineering Research Council of Canada and Fonds de Recherche du Québec – Nature et Technologies. I would also like to thank the Quebec Centre for Biodiversity Science for providing me with financial support to attend an international conference. v TABLE OF CONTENTS SOMMAIRE ............................................................................................................................ i SUMMARY ......................................................................................................................... iii ACKNOWLEDGEMENTS ........................................................................................................ v LIST OF TABLES ...................................................................................................................... x LIST OF FIGURES .................................................................................................................. xii CHAPTER 1 GENERAL INTRODUCTION ............................................................................. 1 1.1 Determinants of parasitism in wild vertebrate populations ............................................... 2 1.1.1 Population characteristics ........................................................................................... 3 1.1.2 Individual characteristics ...........................................................................................
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