Murray, Dennis L

I tracked summer movements and activity of snowshoe hares (Lepus americanus) in southwestern Yukon, Canada, during their population cycle (2015–2022) to assess the primary drivers of movement. Hares exhibited variable movement and activity during the cycle, exhibiting increased home range size and higher daily displacement during low population densities. Males exhibited more dramatic increases in their home ranges (>3-fold), and had greater and more variable movement rates and time spent travelling than females. The ratio of predators to hares was highest at low hare densities when hares were moving most, and seasonal activity of hares seemed unrelated to that of predators. Differences between sexes imply that reproductive activities likely were the main driver of hare movement variation rather than food availability. These findings reinforce that, even in highly variable environments, potential rewards associated with successful mate search and reproduction may outweigh the risks associated with reproductive behaviour.

Author Keywords: behaviour, boreal forest, GPS telemetry, home range, movement ecology, predation risk

Anthropogenic activities such as human activity and livestock grazing are responsible for the global rise in disturbance impacts on wildlife and may underlie regional changes in biodiversity and ecosystem dynamics. Few studies have tried to disentangle the effects of different anthropogenic activities on wildlife behaviour, leaving a major gap in our understanding of conservation and management needs in disturbed areas. Human activity and livestock grazing are increasing in spread and intensity worldwide, thereby imposing pressure on both wildlife and natural areas. In this thesis, I used a camera trapping and occupancy modeling framework to assess whether human presence and livestock grazing had different impacts on site occupancy and activity of 10 wild mammal species, and how responses differed across taxa. Specifically, I predicted that all species would be sensitive to disturbance, but the type and intensity of the response would depend on disturbance type. I detected different responses to each disturbance type across species, but response type (displacement, activity change, crepuscularity) was not associated with species characteristics such as body. Importantly, disturbance intensity had a strong effect on wildlife activity levels, with many species exhibiting marked reductions in activity at high human or livestock disturbance intensity. It remains unclear whether all species' responses are a direct consequence of disturbance versus indirect outcomes of shifts in behaviour of other species in the wildlife community (i.e., disturbance-related changes in prey activity may affect predator activity). Although on the whole disturbance intensity and effect sizes tended to be relatively low in this study, responses were exhibited across all species, implying that disturbance responses may be universal in wild mammals and largely underestimated. Ultimately, my work offers a template for the robust assessment of disturbance impacts on wildlife and provides new avenues for future research to deepen our understanding of wildlife sensitivity to anthropogenic activities.

Author Keywords: activity, anthropogenic disturbance, human activity, livestock, occupancy, wildlife

Climate change has had numerous impacts on species' distributions by shifting suitable habitat to higher latitudes and elevations. These shifts pose new challenges to biodiversity management, in particular translocations, where suitable habitat is considered crucial for the reintroduced population. De-extinction is a new conservation tool, similar to reintroduction, except that the proposed candidates are extinct. However, this novel tool will be faced with similar problems from anthropogenic change, as are typical translocation efforts. Using ecological niche modelling, I measured suitability changes at translocation sites for several Holarctic mammal species under various climate change scenarios, and compared changes between release sites located in the southern, core, and northern regions of the species' historic range. I demonstrate that past translocations located in the southern regions of species' ranges will have a substantial decline in environmental suitability, whereas core and northern sites exhibited the reverse trend. In addition, lower percentages (< 50% in certain scenarios) of southern sites fall above the minimal suitability threshold for current and long-term species occurrence. Furthermore, I demonstrate that three popular de-extinction candidate species have experienced changes in habitat suitability in their historic range, owing to climate change and increased land conversion. Additionally, substantial increase in potentially suitable space is projected beyond the range-limits for all three species, which could raise concerns for native wildlife if de-extinct species are successfully established. In general, this thesis provides insight for how the selection of translocation sites can be more adaptable to continued climate change, and marks perhaps the first rigorous attempt to assess the potential for species de-extinction given contemporary and predicted changes in land use and climate.

Author Keywords: climate change, de-extinction, ecological niche models, MaxEnt, reintroduction, translocation

Population density regulation is a fundamental principle in ecology, however there remain several unknowns regarding the functional expression of density dependence. One prominent view is that the patterns by which density dependence is expressed are largely fixed across a species, irrespective of environmental conditions. Our study investigated the expression of density dependence in Chlamydomonas reinhartti grown under a gradient of nutrient densities, and hypothesized that the relationship between per capita growth rate (pgr) and population density would vary from concave-up to concave-down as nutrients became less limiting. Contrary to prediction, we found that the relationship between a population's pgr and density became increasingly concave-up as nutrient levels increased. Our results suggest that density dependence is strongly variable depending on exogenous and endogenous processes acting on the population, implying that expression of density regulation depends extensively on local conditions. Population growth suppression may be attributable to environments with high intraspecific competition. Additional work should reveal the mechanisms influencing how the expression of density dependence varies across populations through space and time.

Author Keywords: Chlamydomonas reinhartti, density dependence, logistic model, population dynamics, single species growth, theta-logistic equation

Inducible defenses are plastic responses by an organism to the perception of predation risk. This dissertation focuses on three experiments designed to test the hypothesis that plastic ability is limited by energetic constraints. Chapter 1 provides a general introduction to phenotypic plasticity research and the theoretical costs and limitations affecting the expression of plastic traits. In Chapter 2, I tested the hypothesis that costs of early plasticity may be manifested by a reduced response to risk in later life stages. I found that amphibian embryos are able to detect and respond to larval predators, but that the energetic cost of those plastic responses are not equivalent among behavioural, growth, and morphological traits, and their expression differs between closely-related species. Chapter 3 explicitly examines the relationship between food resource availability and plasticity in response to perceived predation risk during larval development. Food-restricted tadpoles showed limited responses to predation risk; larvae at food saturation altered behaviour, development, and growth in response to predation risk. Responses to risk varied through time, suggesting ontogeny may affect the deployment of particular defensive traits. Chapter 4 examines the influence of maternal investment into propagule size on the magnitude of the plastic responses to predation risk in resulting offspring. I found that females in better body condition laid larger eggs and that these eggs, in turn, hatched into larvae that showed greater morphological plasticity in response to predation risk. Maternal investment can therefore affect the ability of offspring to mount morphological defenses to predation risk. Last, Chapter 5 provides a synthesis of my research findings, identifying specific factors constraining the plastic responses of prey to perceived predation risk. Overall, I found constraints on plastic responses imposed by the current environment experienced by the organism (resource availability), the prior experience of the organism (predator cues in the embryonic environment), and even the condition of the previous generation (maternal body condition and reproductive investment). Together, these findings both provide new knowledge and create novel research questions regarding constraints limiting phenotypic variation in natural populations.

Author Keywords: behaviour, inducible defense, Lithobates pipiens, morphometrics, phenotypic plasticity, predation risk

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)

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

Populations have long been delineated by physical barriers that appear to limit reproduction, yet increasingly genetic analysis reveal these delineations to be inaccurate. The eastern and mid-continent populations of sandhill cranes are expanding ranges which is leading to convergence and warrants investigation of the genetic structure between the two populations. Obtaining blood or tissue samples for population genetics analysis can be costly, logistically challenging, and may require permits as well as potential risk to the study species. Non-invasively collected genetic samples overcome these challenges, but present challenges in terms of obtaining high quality DNA for analysis. Therefore, methods that optimize the quality of non-invasive samples are necessary. In the following thesis, I examined factors affecting DNA quality and quantity obtained from shed feathers and examined population differentiation between eastern and mid-continent sandhill cranes. I found shed feathers are robust to environmental factors, but feather size should be prioritized to increase DNA quantity and quality. Further, I found little differentiation between eastern and mid-continent populations with evidence of high migration and isolation-by-distance. Thus, the two populations are not genetically discrete. I recommend future population models incorporate migration between populations to enhance our ability to successfully manage and reach conservation objectives.

Author Keywords: feathers, genetic differentiation, non-invasive DNA, population genetics, population management, sandhill crane (Antigone canadensis)

Wild populations are notoriously difficult to study due to confounding stochastic variables. This thesis tackles two components of investigating wild populations. The first examines the use of niche modeling to quantify macro-scale predator-prey relationships in canid populations across eastern North America, while the second examines range-wide molecular structure in Canada lynx. The goal of the first chapter is to quantify niche characteristics in a Canis hybrid zone of C. lupus, C. lycaon, and C. latrans to better understand the ecological differentiation of these species, and to assess the impacts of incorporating biotic interactions into species distribution models. The goal of the second chapter is to determine if DNA methylation, an epigenetic marker that modifies the structure of DNA, can be used to differentiate populations, and might be a signature of local adaptation. Our results indicated that canids across the hybrid zone in eastern North America exhibit low levels of genetic and ecological differentiation, and that the importance of biotic interactions are largely lost at large spatial scales. We also identified cryptic structure in methylation patterns in Canada lynx populations, which suggest signatures of local adaptation, and indicate the utility of DNA methylation as a marker for investigating adaptive divergence.

Author Keywords: Ecological Epigenetics, Ecological Genetics, SDM

Mountain ranges are often thought to restrict movement of wildlife, yet previous studies evaluating the role of the Rocky Mountains as a dispersal barrier for Canada lynx (Lynx canadensis) have been contradictory. Our study uses neutral microsatellite loci to evaluate the role of the Rocky Mountains as a barrier to gene flow for lynx. Although lynx exhibited low genetic differentiation, we detected a limited effect of the mountains. Furthermore, we inferred the role played by landscape variables in gene flow (genetic differentiation predicted by landscape resistance). Limited gene flow most strongly related to resistance from physical factors (low snow cover and elevation), rather than other topographic and ecological factors (high terrain roughness, low forest cover, low habitat suitability, and geographic distance). Structural connectivity was a relatively poor predictor of functional connectivity. Overall, the Rockies represent an area of reasonably high functional connectivity for lynx, with limited resistance to gene flow.

Author Keywords: Canada lynx, connectivity, gene flow, genetic structure, landscape genetics, Rocky mountains