Habitat loss and fragmentation is a crisis affecting wildlife worldwide. In Tanzania, East Africa, a dramatic and recent (<80 years) expansion in human settlement and agriculture threatens to reduce gene flow among protected areas for many species of large mammals. Wildlife linkages can mitigate population isolation, but linkage designs lacking empirical justification may be controversial and ineffective. Connectivity conservation requires an understanding of how biogeographic factors shaped gene flow prior to habitat loss or fragmentation, however the history of interaction among populations is rarely known. The goal of my study was to provide context for connectivity conservation in central and southern Tanzania by identifying landscape features that have shaped gene flow for three ungulate species with different dispersal capabilities. I investigated historical patterns of connectivity for Maasai giraffe (Giraffa camelopardalis tippelskirchi), impala (Aepyceros melampus), and eland (Tragelaphus oryx) by estimating genetic structure among four to eight protected areas per species. Genetic structure changes very slowly among large populations and thus is likely to reflect historical processes instead of recent anthropogenic influences. I collected noninvasive DNA samples and generated microsatellite genotypes at 8 to 15 loci per species, then estimated genetic diversity metrics (allelic richness, AR, and expected heterozygosity, H[subscript E]) for each population (defined by reserve). I also calculated genetic distance (F[subscript ST] and Nei's unbiased genetic distance, D[subscript hat]) and an estimate of gene flow (Nm) between all population pairs for each species. To elucidate the possible causes of genetic structure between these populations, I tested for isolation by distance and isolation by resistance based on a suite of biogeographic factors hypothesized to affect gene flow. To do this, I created GIS-based resistance surfaces that assigned different costs of movement to landscape features. I created one or more resistance surfaces for each hypothesis of landscape effect. I used circuit theory to estimate the cumulative resistance between each pair of reserves for each weighting scheme, and then performed Mantel tests to calculate the correlation between these resistances and the observed population pairwise genetic distances (D[subscript hat]). I chose the optimal resistance model for each species as the model that was most highly correlated with observed genetic patterns. To verify that the correlation of resistance models with genetic distance was not an artefact of geographic distance, I performed partial Mantel tests to calculate correlation while controlling for the effect of geographic distance. Finally, I compared historical gene flow patterns to the distribution of contemporary human activity to predict areas that are at risk of a loss of connectivity. Indices of genetic diversity were moderate for all three species and comparable to previously reported values for other savannah ungulates. Diversity (both H[subscript E] and A[subscript R]) was highest in eland and lowest in giraffe for these populations, and was not consistently correlated with reserve size as has been reported for other species in East Africa. Although patterns in genetic distance were broadly similar across all three species there were also striking differences in connectivity, highlighting the importance of cross-species comparisons in connectivity conservation. At this scale, resistance models based on slope strongly predicted population structure for all three species; distance to water was also highly correlated with genetic distance in eland. For all three species, the greatest genetic distances occurred between populations separated by the Eastern Arc Mountains, suggesting that the topography of this area has long acted as a barrier to gene flow, but this effect is present in varying degrees for each species. I observed high levels of historical gene flow between centrally located populations (Ruaha National Park and Rungwa Game Reserve) and those in the southwest (Katavi National Park and Rukwa Game Reserve). Although human settlement in this area has been low relative to other areas, the connection between the Katavi/Rukwa and Ruaha ecosystems may be threatened by increased human activity and warrants conservation. High levels of historical gene flow were also seen between reserves in the northeast (Tarangire National Park, Swagaswaga Game Reserve) and the central and southwest populations. These connections appear highly threatened due to current land use practices, and may have already suffered a loss of gene flow. Field surveys in the lands surrounding the northeastern reserves are needed to quantify current levels of connectivity and determine whether corridors could be established to maintain or restore gene flow with other reserves. / Graduation date: 2012

Many mammals are social. The most basic social behaviour is when the actions of one conspecific are directed toward another, what we call the dyadic interaction. Both intrinsic and extrinsic factors may affect an individuals propensity to interact with other members of a population. I used a social cervid, elk (Cervus elaphus), as a model species to test the importance of intrinsic and extrinsic factors of sociality on dyadic interactions. Dyadic interactions not only form the basis for social structure and information transfer within a population, but are also routes of pathogen transmission. My objective in this thesis was thus twofold: to improve our understanding of sociobiology, but also to gain insight into how sociality may underlie the transmission of communicable wildlife disease. I used a hierarchical, autecological approach from DNA, through individual, dyad, group, subpopulation, and ultimately population to explore the effects of intrinsic factors (e.g., sex and pairwise genetic relatedness) and extrinsic factors (e.g., season, conspecific density, habitat, and elk group size) on sociality. Elk in Riding Mountain National Park (RMNP), Manitoba, Canada, are exposed to the causal agent of bovine tuberculosis (Mycobacterium bovis; TB); however, spatial variation in apparent disease prevalence suggests that TB can only persist in one subpopulation within the Park. Using the natural RMNP system and a captive elk herd that I manipulated, I explored factors that influence interaction rates and durations (as a proxy for pathogen transmission) among elk. Sexual segregation in elk results in seasonal and sex-based differences in interaction rate and duration; with interactions peaking in autumn-winter for both sexes. Female-female dyads interact more frequently than male-male dyads. However, male-male dyads interact for longer durations than do female-female dyads. Interaction rate and duration did not covary with pairwise relatedness. Conspecific density also had sex-specific results for interaction rate and duration. Whereas male-male dyadic interaction rates increase with density, female-female dyads increase until they reach a threshold and subsequently reduce their interaction rates at high density. I observed density dependence in interaction rates in experimental trials and from field data. Furthermore, social networks revealed that social familiarity (i.e., heterogeneity of interactions) can be both frequency- and- density dependent depending on the strength of the relationship (i.e., number of repeat interactions). Density also affected the likelihood that an interaction would occur; however, this was modified by vegetation association used by elk. My results reveal several ecological and evolutionary implications for information transfer and pathogen transmission. In particular, I show that seasonal inter-sex routes of transfer may exist and that transfer is likely to be density-dependent. Finally, I conclude that such transfer is modified by available resources.

Many mammals are social. The most basic social behaviour is when the actions of one conspecific are directed toward another, what we call the dyadic interaction. Both intrinsic and extrinsic factors may affect an individuals propensity to interact with other members of a population. I used a social cervid, elk (Cervus elaphus), as a model species to test the importance of intrinsic and extrinsic factors of sociality on dyadic interactions. Dyadic interactions not only form the basis for social structure and information transfer within a population, but are also routes of pathogen transmission. My objective in this thesis was thus twofold: to improve our understanding of sociobiology, but also to gain insight into how sociality may underlie the transmission of communicable wildlife disease. I used a hierarchical, autecological approach from DNA, through individual, dyad, group, subpopulation, and ultimately population to explore the effects of intrinsic factors (e.g., sex and pairwise genetic relatedness) and extrinsic factors (e.g., season, conspecific density, habitat, and elk group size) on sociality. Elk in Riding Mountain National Park (RMNP), Manitoba, Canada, are exposed to the causal agent of bovine tuberculosis (Mycobacterium bovis; TB); however, spatial variation in apparent disease prevalence suggests that TB can only persist in one subpopulation within the Park. Using the natural RMNP system and a captive elk herd that I manipulated, I explored factors that influence interaction rates and durations (as a proxy for pathogen transmission) among elk. Sexual segregation in elk results in seasonal and sex-based differences in interaction rate and duration; with interactions peaking in autumn-winter for both sexes. Female-female dyads interact more frequently than male-male dyads. However, male-male dyads interact for longer durations than do female-female dyads. Interaction rate and duration did not covary with pairwise relatedness. Conspecific density also had sex-specific results for interaction rate and duration. Whereas male-male dyadic interaction rates increase with density, female-female dyads increase until they reach a threshold and subsequently reduce their interaction rates at high density. I observed density dependence in interaction rates in experimental trials and from field data. Furthermore, social networks revealed that social familiarity (i.e., heterogeneity of interactions) can be both frequency- and- density dependent depending on the strength of the relationship (i.e., number of repeat interactions). Density also affected the likelihood that an interaction would occur; however, this was modified by vegetation association used by elk. My results reveal several ecological and evolutionary implications for information transfer and pathogen transmission. In particular, I show that seasonal inter-sex routes of transfer may exist and that transfer is likely to be density-dependent. Finally, I conclude that such transfer is modified by available resources.

L’objectif principal de cette thèse est d’identifier les facteurs urbains et les caractéristiques biologiques de Reticulitermes flavipes qui ont permis à cette espèce invasive de coloniser et de persister en milieu urbain. Plus précisément, les études développées ont visé à mieux comprendre l’origine des colonies introduites à Paris, de déterminer comment celles-ci étaient arrivées en ville et, comment, malgré les moyens de lutte mis en oeuvre au cours de ces dernières décennies elles ont persisté dans le paysage parisien. Une première étude nous a permis de voir que les colonies présentes à Paris sont capables de persister et de réinfester des zones qui ont été précédemment traitées. Ces réinfestations sont possibles grâce aux modes de reproduction (néoténie) et de dispersion (par bouturage) de cette espèce qui lui permettent de survivre localement et de recoloniser des zones où les colonies avaient été partiellement éradiquées et ce, même après 15 ans. Dans une seconde étude, nous avons pu montrer deux facteurs majeurs pouvant expliquer la distribution et la propagation de R. flavipes à Paris : la structure complexe des colonies observées et la combinaison spécifique des variables de l’environnement parisien. Enfin, dans la troisième étude, nous avons décrit la dynamique d’expansion de cette espèce à des échelles nationales et régionales. Cette étude révèle des patterns de distribution variés, reflétant les caractéristiques propres des populations invasives de cette espèce en France. Dans ce travail, nous avons discuté de l’implication des caractéristiques biologiques et paysagères sur le succès invasif de cette espèce. Au vue des données obtenues au cours de cette thèse, nous avons également précisé les scénarios concernant l’histoire de l’invasion de ce termite en France, puis nous avons présenté quelques outils et préconisations qui pourraient permettre, selon nous, d’améliorer la lutte contre cet insecte nuisible en ville. / The main objective of this thesis is to identify the urban factors and biological characteristics of Reticulitermes flavipes which have allowed this invasive species to colonize and persist in urban habitat. More specifically, the developed studies aimed at better understanding the origin of introduced colonies in Paris as well as determining how they persisted in the Parisian landscape, despite implementing pest control during the last decades. A first study allowed us to observe that Parisian colonies were able to persist and reinfest areas which were previously treated. These reinfestations are possible thanks to the mode of reproduction (neoteny) and the way of dispersal (by budding) of this species, which allows it to locally survive and recolonize areas where colonies had been partially eradicated, even fifteen years later. In a second study, we were able to highlight two main factors which could explain the distribution and propagation of R. flavipes in Paris: the observed complex colony structure and the specific combination of the Parisian environmental variables. Finally, in a third study, we were able to identify its dynamic of expansion at national and regional scales which showed varied distribution patterns, reflecting the peculiar characteristics of these invasive species populations in France. In this piece of work, we analyzed the implications of the biological and landscape characteristics on the successful invasion of this species. In the view of the data obtained in this thesis, we also suggested some scenarios on the invasion history of this termite species in France and we provided tools and recommendations which, according to us, could allow the improvement of pest management of this insect in cities.

The genetic diversity and genetic structure of populations, and the processes shaping gene flow within and between populations, are influenced by the landscapes they occur within. Within terrestrial landscapes, rivers and their riparian habitat are among the most dynamic, diverse and complex of landscapes and their linear structure appears as an interlinking feature across large landscapes. This thesis took a landscape genetics approach to examine the influence of river landscape features on Alnus glutinosa populations, a widespread keystone tree species of European riparian ecosystems. By accounting for the differing dispersal mechanisms of A. glutinosa (wind and water), landscape effects on seed- and pollen-mediated gene flow, genetic diversity, demographic and genetic structure were identified at different spatial scales of a large UK river catchment. Widespread gene flow within and between A. glutinosa populations was identified with no apparent limitation of wind-mediated pollen dispersal. Hydrochorous dispersal of seed between populations was evident, and found to increase genetic connectivity between riparian populations; however an isolation by distance effect was identified between populations located further apart from each other. No pattern of genetic diversity was found, with high levels of genetic diversity identified at all spatial and temporal scales. At the river-catchment scale no genetic clustering was observed, either within or between the six rivers studied. Demographic structuring within A. glutinosa populations was evident, and correlated with distance from the main river channel. Interactions between seed dispersal, hydrological disturbance, colonisation, and historical influences are discussed in relation to fine-scale spatial genetic structure between A. glutinosa sapling and adult generations. Central to the landscape genetics approach taken in this thesis was the incorporation of key A. glutinosa life history attributes. By incorporating gene flow analyses, species ecology and landscape features, the research presented here furthers our understanding of riverine landscape influences on their riparian populations at different spatial scales and can be used to inform management principles.

La biologie de la conservation est un domaine de recherche en pleine expansion en raison de la perte accélérée de la biodiversité à l’échelle mondiale. Pour mieux comprendre les processus et les menaces au maintien des populations de petite taille et les effets des facteurs anthropiques sur la biodiversité, la génétique est fréquemment utilisée en conservation. Des analyses génétiques peuvent, par exemple, nous informer sur les tendances à long terme, la diversité des populations et les stratégies de reproduction d’une espèce. La tortue des bois (Glyptemys insculpta) est une espèce endémique à l’Amérique du Nord qui est en danger d’extinction selon l’Union internationale pour la conservation de la nature. Dans le cadre de ma thèse, j’avais comme objectif de caractériser la diversité génétique de cette espèce menacée au Canada. À cet effet, j’ai analysé la génétique des populations de tortues des bois à plusieurs échelles spatiales et temporelles, afin de mieux cerner les processus ayant un impact sur la diversité des populations. Dans un premier temps, les relations de parentalité ont été reconstruites au sein d’une population de tortues des bois pour estimer la fréquence de paternité multiple et de paternité répétée. Les résultats de mes travaux suggèrent que l’emmagasinement de sperme chez la femelle et la reproduction multiple avec les mêmes partenaires pour plus d’une saison de reproduction pourraient expliquer ces phénomènes. Ces stratégies de reproduction pourraient dans ce cas être induites par la faible densité de la population à l’étude, ou encore par la fidélité au site d’hibernation où la majorité des évènements de copulation ont lieu. Par la suite, je me suis intéressée à la diversité génétique des populations de tortues de bois. J’ai voulu comprendre les effets de la configuration spatiale des éléments du paysage et les évènements de dispersion géographique sur la diversité des populations. À l’aide d’une approche de génétique du paysage, mes analyses montrent que la division des populations par bassins versants explique une large fraction de la diversité génétique interpopulations. Ces résultats confirment également que les bassins versants représentent des unités de gestion propices à la protection des populations de tortues des bois. Finalement, des analyses de réseaux ont été utilisées pour mieux cerner la dynamique de flux génique entre les populations de la rive nord et de la rive sud du fleuve Saint-Laurent. Plus spécifiquement, la rive nord se caractérise par un réseau robuste de populations isolées, alors que les populations de la rive sud présentent plutôt une structure de métapopulation. En utilisant les réseaux construits à partir de données génétiques, des scénarios hypothétiques furent comparés pour explorer la sélection de populations à l’aide du logiciel BRIDES. Les résultats de ces analyses ont permis de cibler l’importance de certaines populations de tortues des bois pour la connectivité du réseau. L’importance de ces populations n’aurait pu être prédite par les résultats de la diversité et de la différenciation génétique, les indices de centralité et les analyses d’élimination de nœuds. Grâce à la génétique, cette thèse apporte de nouvelles connaissances sur la tortue des bois, les stratégies de reproduction des différents sexes, le flux génique, la connectivité et l’influence du réseau hydrographique sur la diversité des populations. Ces résultats nous permettent d’avoir une meilleure compréhension des processus affectant la diversité génétique de cette espèce afin de mieux la protéger. Toutes les analyses réalisées pour cette thèse sont directement applicables à l’ensemble des autres espèces longévives avec des générations chevauchantes. / Conservation biology is a rapidly expanding field of research due to the accelerating loss of global biodiversity. To better understand the processes and threats to the persistence of small populations and the effects of anthropogenic factors on biodiversity, genetic approaches are frequently used in conservation. Genetic analyzes can, for example, inform us about long-term trends, population diversity and reproductive strategies of a species. The wood turtle (Glyptemys insculpta) is a species endemic to North America that is endangered according to the International Union for the Conservation of Nature. As part of my thesis, my objective was to characterize the genetic diversity of this threatened species in Canada. In order to better understand the impact of reproductive strategy and landscape structure on population diversity, I analyzed the genetics of wood turtle populations at several spatial and temporal scales. First, parentage relationships were reconstructed in a population of wood turtles to estimate the frequency of multiple and repeated paternity. The results of my work suggest that sperm storage in females and multiple reproduction with the same partners for more than one breeding season could explain these phenomena. These reproduction strategies could in this case be induced by the low density of the study population, or by fidelity to the overwintering site where the majority of copulation events take place. Subsequently, I assessed the genetic diversity of wood turtle populations. I wanted to understand the effects of the landscape configuration and geographic dispersion events on the diversity of populations. Using a landscape genetics approach, my analyzes show that the division of populations by watershed explains a large fraction of the genetic diversity between populations. These results also confirm that watersheds represent management units conducive to the protection of wood turtle populations. Finally, network analysis was used to better understand the gene flow dynamics among populations located on the north and south shores of the St. Lawrence River. More specifically, the north shore is characterized by a robust network of isolated populations, whereas the populations on the south shore present more of a metapopulation structure. Using population graphs, hypothetical scenarios were compared to explore the node selection process using the BRIDES algorithm. The results of these analyzes made it possible to point out specific populations of wood turtles, considering their importance for network connectivity. This could have not been predicted by using genetic diversity and distinctiveness estimates, node-based metrics, and node removal analysis for these populations. Thanks to genetics, this thesis brings new knowledge on the wood turtle, the reproductive strategies of both sexes, the gene flow, the connectivity and the influence of the hydrographic network on population diversity. These results allow us to have a better understanding of the processes affecting the genetic diversity of this species in order to better protect it. All analyses performed for this thesis are directly applicable to other long-lived species with overlapping generations.