Graduate Theses & Dissertations

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Adaptive Genetic Markers Reveal the Biological Significance and Evolutionary History of Woodland Caribou (Rangifer tarandus caribou) Ecotypes
Migratory and sedentary ecotypes are phenotypic distinctions of woodland caribou. I explored whether I could distinguish between these ecotypes in Manitoba and Ontario using genetic signatures of adaptive differentiation. I anticipated that signatures of selection would indicate genetic structure and permit ecotype assignment of individuals. Cytochrome-b, a functional portion of the mitochondrial genome, was tested for evidence of adaptation using Tajima’s D and by comparing variations in protein physiology. Woodland caribou ecotypes were compared for evidence of contemporary adaptive differentiation in relation to mitochondrial lineages. Trinucleotide repeats were also tested for differential selection between ecotypes and used to assign individuals to genetic clusters. Evidence of adaptive variation in the mitochondrial genome suggests woodland caribou ecotypes of Manitoba and Ontario corresponded with an abundance of functional variation. Woodland caribou ecotypes coincide with genetic clusters, and there is evidence of adaptive differentiation between migratory caribou and certain sedentary populations. Previous studies have not described adaptive variation in caribou using the methods applied in this study. Adaptive differences between caribou ecotypes suggest selection may contribute to the persistence of ecotypes and provides new genetic tools for population assessment. Author Keywords: Adaptation, Cytochrome-B, Ecotype, RANGIFER TARANDUS CARIBOU, Selection, TRINUCLEOTIDE REPEAT
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
Calving site selection and fidelity in a restored elk (Cervus elaphus) herd in Bancroft, Ontario, Canada
ABSTRACT Calving site selection and fidelity in a restored elk (Cervus elaphus) herd in Bancroft Ontario, Canada. Michael R. Allan Parturition site selection by ungulates is believed to be influenced by forage abundance and concealment from predators. In 2011 and 2012, I used vaginal implant transmitters and movements to identify calving sites for 23 GPS collared elk (Cervus elaphus) from a restored herd. I tested the hypothesis that maternal elk used sites with higher forage and denser concealment compared to pre-calving sites at micro and macrohabitat levels. I detected no significant microhabitat differences from direct measurements of vegetation. At the macrohabitat scale, based on proximity of landcover classes, mean distances to hardwood forests was significantly less for calving (153 m) than pre-calving sites (198 m). Site fidelity is hypothesized to offer security in terms of familiarity to an area. I tested the hypothesis that females demonstrated fidelity to their previous year's location during pre-partum, parturition, post partum, breeding and winter periods. Elk were more philopatric during parturition and post partum than during breeding. Compared to winter elk were more philopatric during pre-partum, parturition and post-partum periods. Expressed as distance between consecutive-year calving locations, site fidelity varied with 27% of females exhibiting high (<1 km), 18% moderate and 55% (>2.9 km) low fidelity. I measured nearest-neighbour distances at calving time, exploring the hypothesis that females distance themselves from conspecifics. Elk increased the average distances to collared conspecifics during parturition; however, sample sizes were small. This strategy might influence calving site selection. Rapid movement prior to parturition, low site fidelity and spacing-out of females during parturition appear to be strategies to minimize predator risk and detection. Little evidence of selection for vegetation structure suggests this may not be limiting to these elk. Author Keywords: calving, elk, fidelity, movement, parturition, selection
Canid Predation of Domestic Sheep (Ovis aries) on Ontario Farms
Livestock predation by wild predators is a frequent and complicated issue, often cited as a significant factor in the decline of livestock production and justification for killing predators. Coyotes (Canis latrans) are the primary predators of sheep in Ontario. Some farms appear to be more susceptible to predation than others, despite the use of mitigation techniques. I explored land cover in the vicinity of farms as a potential influence on the level of predation, as coyote abundance and wild prey are correlated with certain habitat types. Using model competition, I show that landscape explains little variation in predation levels over all farms, but can explain 27% of variation in the percent of a flock killed. Total forest edge habitat and distance between forest patches were both positively associated with losses, suggesting a reduction in forest cover surrounding a farm puts the flock at greater risk. In addition, I tested four disruptive deterrents for effectiveness at protecting flocks. A matched-pairs analysis did not show a statistically significant benefit of these non-lethal mitigation tools. Author Keywords: Alternative Prey, Canis latrans, Coyote, Landscape, Predation Deterrents, Sheep
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)
Detection of four at-risk freshwater pearly mussel species (Bivalvia
Environmental DNA (eDNA) detection uses species-specific markers to screen DNA from bulk samples, such as water, to infer species presence. This study involved the development and testing of species-specific markers for four freshwater pearly mussels (Unionidae). The markers were applied to water samples from intensively sampled mussel monitoring sites to compare species detections from eDNA with established sampling method detections. Target species were detected using eDNA at all sites where they had previously been detected by quadrat sampling. This paired design demonstrated that eDNA detection was at least as sensitive as quadrat sampling and that high species specificity can be achieved even when designing against many sympatric unionids. Detection failures can impede species conservation efforts and occupancy estimates; eDNA sampling could improve our knowledge of species distributions and site occupancy through increased sampling sensitivity and coverage. Author Keywords: conservation genetics, cytochrome oxidase subunit I (COI), environmental DNA (eDNA), quantitative PCR (qPCR), species at risk (SAR)
Discriminating grey wolf (Canis lupus) predation events in a multi-prey system in central Saskatchewan
I investigated if spatio-temporal behaviour of grey wolves (Canis lupus) determined via GPS collar locations could be used to discriminate predation events generally, and among prey species, in Prince Albert National Park during winter, 2013-2017. I used characteristics of spatio-temporal GPS clusters to develop a predictive mixed-effect logistic regression model of which spatial clusters of locations were wolf kill sites. The model suffered a 60 % omission error when tested with reserved data due to the prevalence of deer kills with correspondingly low handling time. Next, I found a multivariate difference in the percentage of habitat classes used by wolves in the 2 hours preceding predation events of different prey species, suggesting that wolf habitat use reflects prey selection at a fine-scale. My results highlight the difficulty and future potential for remoting discriminating wolf predation events via GPS collar locations in multi-prey ecosystems. Author Keywords: Canis lupus, GPS clusters, GPS collars, grey wolf, habitat use, predation
ECTOPARASITIC INFLUENCES OF DIPTERA ON THE ACTIVITY AND BEHAVIOUR OF WOODLAND CARIBOU (RANGIFER TARANDUS) IN A MANAGED BOREAL FOREST LANDSCAPE IN NORTHERN ONTARIO
Caribou experience direct and indirect negative effects of harassment from biting flies, influencing behavior and activity on several spatial and temporal scales. I used systematic insect collection surveys during the summers of 2011 and 2012 to examine the spatial and temporal distributions of black flies (Simuliidae), mosquitoes (Culicidae), and deer flies and horse flies (Tabanidae) in a managed boreal forest in northern Ontario. Mosquitoes had a positive association with densely treed habitats, whereas black flies more often occurred in open areas, and tabanids had a strong presence in all habitat types. Habitats in proximity to large bodies of water had fewer biting flies than inland areas. Young stands supported higher abundances of tabanids despite vegetation community type. Next, I tested for seasonal effects of biting fly abundance on caribou activity by modelling the seasonal trend in abundance for each fly family for each year and compared this to an index of daily activity for 20 radio-collared female caribou in 2011 and 10 females in 2012. I modeled this index of caribou activity for each animal in each year and extracted the set of partial correlation coefficients from multiple regressions to test for effects of biting fly abundances on caribou activity. Caribou reduced their daily activity when tabanids were more numerous, and increased activity when mosquitoes were numerous. This divergent response may reflect a difference in the efficacy of moving to reduce harassment, owing to the stronger flight capabilities of tabanids. Author Keywords: Activity, Anthropogenic Disturbance, Behaviour, Insect harassment, Temporal distribution, Woodland Caribou
Effects of Geographic Factors on the Wild Harvest of Large Mammals across North America
While the harvest of mammals is monitored in each jurisdiction across Canada and the USA, there has been no analysis of this wild harvest at a continental scale across North America. The recreational wild harvest of large mammals varies geographically across North America, and I hypothesized that this variation is influenced by both anthropogenic and other environmental factors on the landscape. I tested this hypothesis using annual harvest tallies collected by Conservation Visions Inc. for mammals for each state, provincial, and territorial jurisdiction in Canada and the USA. I built multiple additive models of the harvest, in one harvest year, 2015 – 2016, to test for landscape gradients that explain the variation in harvest levels for seven large mammal species: white-tailed deer (Odocoileus virginianus), black bear (Ursus americanus), bighorn sheep (Ovis canadensis), elk (Cervus canadensis), mule deer (Odocoileus hemionus), pronghorn (Antilocapra americana), and moose (Alces alces). I built these models from a suite of nine putative predictor variables that comprised landcover, human footprint, and evapotranspiration. For all large mammal species except for pronghorn, anthropogenic influence had a positive effect on the wild harvest density, consistent with the idea that the proximity of human populations and roads are important for fostering wild harvest activity by providing hunters access to hunting areas. The harvest of white-tailed deer, elk, and pronghorn were negatively affected by vegetation structure, urbanization, and primary productivity, respectively. Understanding the recreational wild harvest at a broad-spatial scale provides a unique perspective of the North American model of wildlife conservation and spurs future comparative analyses of the wild harvest across spatial scales. Author Keywords: Anthropogenic Influence, Hunting, Large Mammals, Primary Productivity, Vegetation Structure, Wild Harvest
Effects of Hydroelectric Corridors on the Distribution of Female Caribou (Rangifer tarandus) on the Island of Newfoundland
A species of concern is caribou (Rangifer tarandus), a species in decline across most of the circumpolar North, including the island of Newfoundland. Resource exploitation across caribou ranges is projected to accelerate in the coming decades as oil extraction, roads, forest harvesting, and mining encroach upon their habitat. Hydroelectric corridors, in particular, are anticipated to expand significantly. The effects of these linear developments on caribou habitat remain unclear. I capitalized on an existing dataset of nearly 700 radio‐tracked female caribou, 1980‐2011, to determine the long‐term effects of hydroelectric corridors on their seasonal distributions. Using an island-wide landcover map, I tested for preference or avoidance hydroelectric corridors in each of 4 seasons using the Euclidean Distance habitat selection technique at the extent of the population ranges (broad scale) for each decade (1980s, 1990s, 2000s). I also examined the distribution of caribou ≤10 km and ≤20 km from corridors (narrow scale) for five herds. At the broad scale, the response was highly variable. Female caribou were most likely to avoid corridors during the 1980s, but they often exhibited little aversion, even preference for corridors, particularly in the 1990s and 2000s. Hydroelectric corridors, therefore, did not appear to be limiting at this scale. I surmise that these long-term shifts reflect the heightened density-dependent food limitation for Newfoundland caribou. At the narrow scale, avoidance of corridors was common – typically, a 50% reduction in use within 2-5 km of the corridor. Consistent with the broad scale, caribou exhibited the strongest tendency for avoidance in the 1980s compared to subsequent decades. Understanding space-use remains central to the study of caribou ecology. Hydroelectric lines in Newfoundland tended to coincide with other anthropogenic features. Cumulative effects must be considered to understand the full range of effects by human developments on caribou. Author Keywords: Caribou, distribution, habitat, hydroelectric, Newfoundland, Rangifer tarandus
Effects of Local, Landscape, and Temporal Variables on Bobolink Nest Survival in Southern Ontario
Populations of grassland birds, including the Bobolink (Dolichonyx oryzivorus), are experiencing steep declines due to losses of breeding habitat, land use changes, and agricultural practices. Understanding the variables affecting reproductive success can aid conservation of grassland species. I investigated 1) whether artificial nest experiments accurately estimate the impacts of cattle on the daily survival rate of Bobolink nests and 2) which local, landscape, and temporal variables affect daily survival rate of Bobolink nests in Southern Ontario. I replicated an artificial nest experiment performed in 2012 and 2015 to compare the daily survival rate of artificial and natural nests at multiple stocking rates (number of cattle × days × ha-1). I also monitored Bobolink nests and modeled daily survival rate using local variables (e.g., stocking rate, field use, patch area), landscape variables (e.g., percent forest within 2, 5, and 10 km), and temporal variables (e.g., year, date of season). Results indicate that artificial nest experiments using clay shooting targets overestimated the impacts of stocking rate on the daily survival rate of Bobolink nests. With natural nests, region (confounded by year and field use), stocking rate, and date of season were the strongest predictors of daily survival rate; with stocking rate and date of season both having a negative effect. Management should focus on conserving pastures with low stocking rates (< 40 cattle × days × ha-1), late-cut hayfields, fallow fields, and other grasslands to protect breeding grounds for the Bobolink and other declining grassland bird species. Author Keywords: Bobolink, Daily survival rate, landscape variables, local variables, Nest survival, temporal variables

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