Graduate Theses & Dissertations

Beyond Habitat
My objective was to understand how individual variation, in conjunction with variation in habitat, can affect individual and population-level variation in animal space use. I used coyotes (Canis latrans) as a model species to investigate the roles of hybridization, an inherited intrinsic factor, and spatial memory, a learned intrinsic factor, on space use. I used a diversity of methods and approaches, including meta-regression, multiple imputation, simulations, resource selection functions, step selection functions, net-squared displacement analysis, and survival analysis. A major contribution was my investigation of the performance of multiple imputation in a meta-regression framework in Chapter 2. My simulations indicated that multiple imputation performs well in estimating missing data within a meta-regression framework in most situations. In Chapter 3, I used published studies of coyote home range size in a meta-regression analysis with multiple imputation to examine the relative roles of hybridization and environmental variables on coyote home range size across North America. I found that hybridization with Canis species was a leading factor driving variation in coyote space use at a continental scale. In Chapter 4, I used telemetry data for 62 coyotes in Newfoundland, Canada, to investigate the influence of cognitive maps on resource use. I found that resident coyotes used spatial memory of the landscape to select or avoid resources at spatial scales beyond their immediate sensory perception relative to transient coyotes, presumably increasing their fitness. Taken together, my dissertation demonstrates that intrinsic factors, such as genetic ancestry and spatial memory, can have substantial influences on how animals use space at both individual and population levels, and at both a local and a continental scales. Author Keywords: canis latrans, hybridization, meta-regression, multiple imputation, Newfoundland, spatial memory
Conservation Genetics of Woodland Caribou in the Central Boreal Forest of Canada
Maintaining functional connectivity among wildlife populations is important to ensure genetic diversity and evolutionary potential of declining populations, particularly when managing species at risk. The Boreal Designatable Unit (DU) of woodland caribou (Rangifer tarandus caribou) in Ontario, Manitoba, and Saskatchewan has declined in southern portions of the range because of increased human activities and has been identified as 'threatened' by the Committee on the Status of Endangered Wildlife in Canada (COSEWIC). In this dissertation, I used ten microsatellite DNA markers primarily from winter-collected fecal samples to delineate genetic structure of boreal caribou in declining portions of the range and increase understanding of the potential influence of the non-threatened Eastern Migratory DU of woodland caribou on genetic differentiation. Eastern migratory caribou are characterized by large home ranges compared to boreal caribou and migrate seasonally into portions of the Boreal DU range. A regional- and local-scale analysis using the spatial Bayesian clustering algorithm in program TESS delineated four regional clusters and 11 local clusters, with the majority of local clusters occurring along the southern periphery of the range. One of those clusters in Ontario corresponded spatially with the seasonal overlap of boreal and eastern migratory caribou and was characterized by substantial admixture, suggesting that the two DUs could be interbreeding. Next, I decoupled the impacts of historical and contemporary processes on genetic structure and found that historical processes were an important factor contributing to genetic differentiation, which may be a result of historical patterns of isolation by distance or different ancestry. Moreover, I found evidence of introgression from a currently unsampled population in northern Ontario, presumably barren-ground caribou (R. t. groenlandicus). Finally, because our analysis suggested recent processes were also responsible for genetic structure, I used a landscape genetics analysis to identify factors affecting contemporary genetic structure. Water bodies, anthropogenic disturbance, and mobility differences between the two DUs were important factors describing caribou genetic differentiation. This study provides insights on where conservation and management of caribou herds should be prioritized in threatened portions of the boreal caribou range and may have implications for future delineation of evolutionarily significant units. Author Keywords: boreal forest, genetic structure, landscape genetics, microsatellite DNA, Rangifer tarandus, woodland caribou
Demography and habitat selection of Newfoundland caribou
The objective of this thesis is to better understand the demography and habitat selection of Newfoundland caribou. Chapter 1 provides a general introduction of elements of population ecology and behavioural ecology discussed in the thesis. In Chapter 2, I examine the causes of long-term fluctuations among caribou herds. My findings indicate that winter severity and density-dependent degradation of summer range quality offer partial explanations for the observed patterns of population change. In Chapter 3, I investigate the influence of climate, predation and density-dependence on cause-specific neonate survival. I found that when caribou populations are in a period of increase, predation from coyotes and bears is most strongly influenced by the abiotic conditions that precede calving. However, when populations begin to decline, weather conditions during calving also influenced survival. I build on this analysis in Chapter 4 by determining the influence of climate change on the interplay between predation risk and neonate survival. I found that the relative equilibrium between bears and coyotes may not persist in the future as risk from coyotes could increase due to climate change. In Chapter 5, I investigate the relationships in niche overlap between caribou and their predators and how this may influence differential predation risk by affecting encounter rates. For coyotes, seasonal changes in niche overlap mirrored variation in caribou calf risk, but had less association with the rate of encounter with calves. In contrast, changes in niche overlap during the calving season for black bears had little association with these parameters. In Chapter 6, I examine broad-level habitat selection of caribou to study trade-offs between predator avoidance and foraging during the calving season. The results suggest that caribou movements are oriented towards increased access to foraging and the reduction of encounter risk with bears, and to a lesser extent, coyotes. Finally, I synthesize the major findings from this thesis and their relevance to caribou conservation in Chapter 7, to infer that Newfoundland caribou decline is ultimately driven by extrinsic and intrinsic elements related to density-dependence. Reduction in neonate survival emerged from nutritionally-stressed caribou females producing calves with lower survival. Author Keywords: Behavioural ecology, Black bear (Ursus americanus), Coyote (Canis latrans), Population ecology, Predator-prey interactions, Woodland caribou (Rangifer tarandus)
Environmental structure, morphology and spatial ecology of the five-lined skink (Plestiodon fasciatus) at high latitude range limits
Detecting relevant and meaningful patterns from the complex, interconnected network of relationships between organisms and their environment is a primary objective of ecology. Ecological patterns occur across multiple scales of space and time. In this dissertation, I examine aspects of environmental structure that influence a species’ distribution and are expressed in that species’ population dynamics. I compare the morphology of the five-lined skink (Plestiodon fasciatus) near its high latitude range limits with a lower latitude population and evaluate the economics of their behaviour in the context of its reproductive strategy. I tested the conformity of this species to biogeographical rules postulated by MacArthur, Bergmann, and Rensch. Spatial ecology was investigated in the context of the environmental potential for polygamy proposed by Emlen and Oring (1977) The five-lined skink, Plestiodon fasciatus, conformed to these biogeographic rules. Specifically, abiotic factors were the primary limiting factors affecting distribution at the high latitude range limits of the species; body size was larger in high latitude populations; and the degree of sexual size dimorphism was greater at high latitude than at low latitude. Spatial ecology at the individual scale was influenced by sites with suitable thermal conditions which facilitate the polygynandrous mating system documented in P. fasciatus in high latitude populations. My results confirm the importance of microsites with suitable thermal profiles as key habitat for ectothermic vertebrates at high latitudes. The influence of temperature as a limiting abiotic factor is expressed in population density, body size, spatial ecology, and reproductive strategy of P. fasciatus. Conservation and restoration of high latitude populations of ectothermic vertebrates should focus on ensuring thermal requirements of the species of concern are met before other factors are addressed, as temperature is likely the single most important limiting factor at high latitude range limits. Author Keywords: biogeography, lizard, Plestiodon fasciatus, range limits, sexual size dimorphism, spatial ecology
Hybridization Dynamics between Wolves and Coyotes in Central Ontario
Eastern wolves (Canis lycaon) have hybridized extensively with coyotes (C. latrans) and gray wolves (C. lupus) and are listed as a `species of special concern' in Canada. Previous studies have not linked genetic analysis with field data to investigate the mechanisms underlying Canis hybridization. Accordingly, I studied genetics, morphology, mortality, and behavior of wolves, coyotes, and hybrids in and adjacent to Algonquin Provincial Park (APP), Ontario. I documented 3 genetically distinct Canis types within the APP region that also differed morphologically, corresponding to putative gray wolves, eastern wolves, and coyotes. I also documented a substantial number of hybrids (36%) that exhibited intermediate morphology relative to parental types. I found that individuals with greater wolf ancestry occupied areas of higher moose density and fewer roads. Next, I studied intrinsic and extrinsic factors influencing survival and cause-specific mortality of canids in the hybrid zone. I found that survival was poor and harvest mortality was high for eastern wolves in areas adjacent to APP compared with other sympatric Canis types outside of APP and eastern wolves within APP. Contrary to previous studies of wolves and coyotes elsewhere, I hypothesized that all Canis types exhibit a high degree of spatial segregation in the Ontario hybrid zone. My hypothesis was supported as home range overlap and shared space use between neighboring Canis packs of all ancestry classes were low. Territoriality among Canis may increase the likelihood of eastern wolves joining coyote and hybrid packs and exacerbate hybridization. Canids outside APP modified their use of roads between night and day strongly at high road densities (selecting roads more at night), whereas they responded weakly at lower road densities (generally no selection). Individuals that survived exhibited a highly significant relationship between the difference in their night and day selection of roads and availability of roads, whereas those that died showed a weaker, non-significant response. My results suggest that canids in the unprotected landscape outside APP must balance trade-offs between exploiting benefits associated with secondary roads while mitigating risk of human-caused mortality. Overall, my results suggest that the distinct eastern wolf population of APP is unlikely to expand numerically and/or geographically under current environmental conditions and management regulations. If expansion of the APP eastern wolf population (numerically and in terms of its geographic distribution) is a conservation priority for Canada and Ontario, additional harvest protection in areas outside of APP may be required. If additional harvest protection is enacted, a detailed study within the new areas of protection would be important to document specific effects on eastern wolf population growth. Author Keywords: Canis, coyotes, eastern wolves, hybridization, resource selection, survival
Reproductive Fitness of Smallmouth Bass (Micropterus dolomieu) Under Heterogeneous Environmental Conditions
Identifying the biotic and abiotic factors that influence individual reproductive fitness under natural conditions is essential for understanding important aspects of a species’ evolutionary biology and ecology, population dynamics, and life-history evolution. Using next generation sequencing technology, I developed five microsatellite multiplex reactions suitable for conducting large scale parentage analysis of smallmouth bass, Micropterus dolomieu, and used molecular pedigree reconstruction techniques to characterize the genetic mating system and mate selection in adult smallmouth bass nesting in Lake Opeongo, Ontario, Canada. I used multivariate spatial autocorrelation analysis to indirectly infer the occurrence and extent of natal philopatry among spawning adults, to assess the strength and direction of sex-bias in natal dispersal patterns, and to evaluate the degree of nest site fidelity and breeding dispersal of spawning adults. I also evaluated how differences in littoral zone water temperature caused by wind-induced seiche events influence the relative reproductive success of spawning adults. Lastly, I provide a synopsis of potential future research aimed at further exploring factors that influence the reproductive fitness of smallmouth bass in Lake Opeongo. This information will contribute to our understanding of the factors regulating smallmouth bass populations, and provide insight into the factors controlling the variance in individual reproductive success and thus recruitment dynamics in this species. Author Keywords: Dispersal, Fitness, Mate selection, Mating systems, Philopatry
Habitat loss and fragmentation can disrupt population connectivity, resulting in small, isolated populations and low genetic variability. Understanding connectivity patterns in space and time is critical in conservation and management planning, especially for wide-ranging species in northern latitudes where habitats are becoming increasingly fragmented. Wolverines (Gulo gulo) share similar life history traits observed in large-sized carnivores, and their low resiliency to disturbances limits wolverine persistence in modified or fragmented landscapes - making them a good indicator species for habitat connectivity. In this thesis, I used neutral microsatellite and mitochondrial DNA markers to investigate genetic connectivity patterns of wolverines for different temporal and spatial scales. Population genetic analyses of individuals from North America suggested wolverines west of James Bay in Canada are structured into two contemporary genetic clusters: an extant cluster at the eastern periphery of Manitoba and Ontario, and a northwestern core cluster. Haplotypic composition, however, suggested longstanding differences between the extant eastern periphery and northwestern core clusters. Phylogeographic analyses across the wolverine's Holarctic distribution supported a postglacial expansion from a glacial refugium near Beringia. Although Approximate Bayesian computations suggested a west-to-east stepping-stone divergence pattern across North America, a mismatch distribution indicated a historic bottleneck event approximately 400 generations ago likely influenced present-day patterns of haplotype distribution. I also used an individual-based genetic distance measure to identify landscape features potentially influencing pairwise genetic distances of wolverines in Manitoba and Ontario. Road density and mean spring snow cover were positively associated with genetic distances. Road density was associated with female genetic distance, while spring snow cover variance was associated with male genetic distance. My findings suggest that northward expanding anthropogenic disturbances have the potential to affect genetic connectivity. Overall, my findings suggest that (1) peripheral populations can harbour genetic variants not observed in core populations - increasing species genetic diversity; (2) historic bottlenecks can alter the genetic signature of glacial refugia, resulting in a disjunct distribution of unique genetic variants among contemporary populations; (3) increased temporal resolution of the individual-based genetic distance measure can help identify landscape features associated with genetic connectivity within a population, which may disrupt landscape connectivity. Author Keywords: conservation genetics, Holarctic species, landscape genetics, peripheral population, phylogeography, wolverine
White-Tailed Fear
The primary method used to maintain white-tailed deer (Odocoileus virginianus) populations at densities that are ecologically, economically, socially, and culturally sustainable is hunter harvest. This method considers only the removal of animals from the population (the direct effect) and does not conventionally consider the costs imposed on deer as they adopt hunter avoidance strategies (the risk effect). The impact of risk effects on prey can exceed that of direct effects and there is interest in applying this concept to wildlife management. Deer are potential candidates as they have demonstrated behavioural responses to hunters. I explored the potential of such a management practice by quantifying how human decisions around hunting create a landscape of fear for deer and how deer alter their space use and behaviour in response. I used a social survey to explore the attitudes of rural landowners in southern and eastern Ontario towards deer and deer hunting to understand why landowners limited hunting on their property. I used GPS tracking devices to quantify habitat selection by hunters and hunting dogs (Canis familiaris) to better understand the distribution of hunting effort across the landscape. I used GPS collars to quantify the habitat selection of deer as they responded to this hunting pressure. I used trail cameras to quantify a fine-scale behavioural response, vigilance, by deer in areas with and without hunting. Human actions created a highly heterogeneous landscape of fear for deer. Landowner decisions excluded hunters from over half of the rural and exurban landscape in southern and eastern Ontario, a pattern predicted by landowner hunting participation and not landcover composition. Hunter decisions on whether to hunt with or without dogs resulted in dramatically different distributions of hunting effort across the landscape. Deer showed a high degree of behavioural plasticity and, rather than adopting uniform hunter avoidance strategies, tailored their response to the local conditions. The incorporation of risk effects into white-tailed deer management is feasible and could be done by capitalizing on a better understanding of deer behaviour to improve current management practices or by designing targeted hunting practices to elicit a landscape of fear with specific management objectives. Author Keywords: Brownian bridge movement models, hunting, landscape of fear, resource utilization functions, risk effects, white-tailed deer
cascading effects of risk in the wild
Predation risk can elicit a range of responses in prey, but to date little is known about breadth of potential responses that may arise under realistic field conditions and how such responses are linked, leaving a fragmented picture of risk-related consequences on individuals. We increased predation risk in free-ranging snowshoe hares (Lepus americanus) during two consecutive summers by simulating natural chases using a model predator (i.e., domestic dog), and monitored hare stress physiology, energy expenditure, behaviour, condition, and habitat use. We show that higher levels of risk elicited marked changes in physiological stress metrics including sustained high levels of free plasma cortisol which had cascading effects on glucose, and immunology, but not condition. Risk-augmented hares also had lowered daily energy expenditure, spent more time foraging, and decreased rest, vigilance, and travel. It is possible that these alterations allowed risk-exposed hares to increase their condition at the same rate as controls. Additionally, risk-augmented hares selected, had high fidelity to, and were more mobile in structurally dense habitat (i.e., shrubs) which provided them additional cover from predators. They also used more open habitat (i.e., conifer) differently based on locale within the home range, using familiar conifer areas within cores for rest while moving through unfamiliar conifer areas in the periphery. Overall, these findings show that prey can have a multi-faceted, highly plastic response in the face of risk and can mitigate the effects of their stress physiology given the right environmental conditions. Author Keywords: behaviour, condition, daily energy expenditure, predator-prey interactions, snowshoe hare, stress physiology

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