Graduate Theses & Dissertations

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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
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)
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
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
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
spatial and temporal distribution of tabanid (Chrysops, Hybomitra and Tabanus) species in the Nakina district of northwestern Ontario
This thesis focused on expanding knowledge of Hybomitra, Chrysops and Tabanus (Diptera: Tabanidae) distributions north of Lake Nipigon, Ontario, in a managed boreal forest. As land use and climate changes accelerate, there is increased pressure to increase knowledge from which to monitor changes. In 2011 and 2012, 8928 individuals representing, 44 species were captured using sweep netting. Major northward range extensions were observed for Chrysops shermani, C. aberrans and Tabanus fairchildi. Smaller range extensions and in-fills were observed for another 15 species. 23 species had exntensions to their previously known seasonal range. C. carbonarius was the only species that showed an extension to both sides of its season. In general, harvested stands had 50% more individuals and 30% greater species richness than younger stands. A possible link between stand age and interspecific competition was identified. Information has been provided to build baseline of species richness, relative abundance and distribution of Tabanid flies. Author Keywords: diptera, distribution, natural history, northern Ontario, species range, tabanid
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)
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
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
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
Regional differences in the whistles of Australasian humpback dolphins (genus Sousa)
Most delphinids produce narrowband frequency-modulated whistles with a high level of plasticity to communicate with conspecifics. It is important to understand geographic variation in whistles as signal variation in other taxa has provided insight into the dispersal capabilities, genetic divergence and isolation among groups, and adaptation to ecological conditions. I investigated whistle variation of Indo-Pacific humpback dolphins (Sousa chinensis chinensis), Taiwanese humpback dolphins (S. c. taiwanensis) and Australian humpback dolphins (S. sahulensis) to test whether differences in whistles support the hypotheses of population structure, regional and species differences in the genus Sousa, which were based on morphological and genetic data. I also investigated important factors that may contribute to local distinctiveness in whistles including behavioural state, group size, and the influence of vessel noise. Multivariate analyses of seven acoustic variables supported the hypotheses of population structure, regional and species differences. Acoustic diversification between groups is likely influenced by behaviour and social contexts of whistles, and environmental noise. The use of sound to identify discrete groups of humpback dolphins may be important in future studies where genetic and morphological studies may not reveal recent differentiation or are difficult to conduct. Author Keywords: Bioacoustics, Cetacean, Geographic variation, Population biology, Sousa, Whistle characteristics
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

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