Northrup, Joseph M

Animal migration is defined as the seasonal movement from one independent and non-overlapping range to another. Understanding how and why animals migrate is important not only to understand their life history processes but also for informing other important ecological processes such as the spread of wildlife disease and habitat alteration. Animal migrations have been impacted by human activity with instances of complete loss of migrations in human-altered areas. Understanding the drivers of migration can help predict responses to future environmental changes and potentially help conserve these phenomena. Seasonal movements of white-tailed deer (deer; Odocoileus virginianus; Zimmerman, 1780) have been linked to seasonal changes in environmental conditions that impact their ability to find food resources and risk of predation. The human shield hypothesis posits that prey species will select habitat close to people to use predator fear of humans to protect themselves from predation. Using global positioning system (GPS) collars, we examined the onset of deer migrations and assessed how environmental variables including snow, temperature, and plant biomass influenced migration departure dates using time-to-event models. We compared deer locations to data from GPS collared coyotes (Canis latrans; Say, 1823) within the same study area to explore daily space-use differences and examine if deer migrations were food or predation-risk driven using generalized linear mixed effects regression models. We found substantial annual and individual variation in deer migration dates. Snow depth was the strongest and most consistent predictor of deer migration, with individuals departing earlier with greater snow depth. Our regression analyses showed that deer selected for habitats closer to and with greater density of anthropogenic structures than coyotes at all times. After removing the animal locations close to areas with active supplemental feeding, these effects were diminished showing no differences in proximity or density of structures. Overall, we found more support for a food driven migration rather than a predator driven human shield. With the reduction in natural food caused by snow cover, we suggest that supplemental feeding is likely influencing the use of wintering areas by deer. The high proportion of deer migrating to human developed areas with supplemental feeders highlights the need for continued research into the impacts of human activity on animal behaviour.

Author Keywords: coyote, human shield, migration, supplemental feeding, white-tailed deer

Quantifying the impacts of environmental conditions on the abundance of wildlife populations is important for making informed management decisions in the face of increasing environmental threats. Managers require robust tools to estimate abundance and density of wildlife rapidly and with precision. Within the context of studying white-tailed deer, I evaluated the use of camera-traps and a recently developed spatial-mark resight model to estimate deer density and evaluate habitat and land use factors influencing deer density. The study was conducted in central Ontario, Canada on approximately 16 km2 of public land including the protected Peterborough Crown Game Preserve. Telemetry locations from 39 radio-collared deer were used and one hundred camera-traps were deployed for a total of 140 days from January 2022 to May 2022. Using telemetry locations and camera-trap photos I built a two-step spatial-mark resight model to estimate deer density. Deer density varied during the study as a portion of the population migrated to wintering areas outside of the study area. Despite fluctuations in precision, estimates improved towards the end of the study as more data became available and deer space use stabilized. The average deer density during the entire study was 3.0 deer/km2 (95% CI= 0.1, 5.8; SD= 1.7; CV= 55%; N= 238 deer). The lowest mean density was 0.2 deer/km2 (95% CI= 0.1, 0.4; SD= 0.1; CV= 50%; N= 15 deer) from February 26th to March 11th and the highest mean density was 4.8 deer/km2 (95% CI= 3.1, 6.2; SD= 0.8; CV= 17%; N= 378 deer) from May 7th to May 20th. When I incorporated spatial covariates into the model to estimate effects on deer density, higher proportions of mixed forest, deciduous forest, and road and trail density all had negative effects on deer density. While models contained some uncertainty, deer density appeared higher in the portion of the study area protected from licensed hunting. This thesis provides a framework for managers to use camera-traps and the spatial-mark resight model to monitor deer populations and link environmental covariates to spatial variation in density. As environmental threats such as habitat loss and infectious diseases increase in severity, monitoring wildlife population numbers will be vital for informed responses to these threats. The two-step spatial-mark resight model with environmental covariates provides managers with a long-term monitoring tool to evaluate management efforts and population health in forested areas.

Author Keywords: camera-trap, chronic wasting disease, landscape ecology, spatial-capture recapture, white-tailed deer, wildlife management

Animal migrations are ubiquitous and one of the most threatened ecological processes globally. Because of the multifaceted nature of migration – seasonal movements between home ranges – it can be difficult to tease apart the underlying mechanisms influencing this behaviour. It is necessary to understand these mechanisms, not only to deepen our fundamental understanding of migration in animals, but also because migrations in many species are vulnerable to environmental change. In Chapter 2, I first systematically identify the broad proximate drivers of migration and offer generalities across vertebrate taxa. I quantitatively reviewed 45 studies and extracted 132 observations of effect sizes for internal and external proximate drivers that influenced migration propensity. Through this meta-analysis, I found that internal and external drivers had a medium and large effect, respectively, on migration propensity. Predator abundance and predation risk had a large effect on migration propensity, as did individual behaviour. Of the studies that examined genetic divergence between migrant and resident populations, 64% found some genetic divergence between groups. In Chapter 3, I explore the genetic basis for migration and identified genes associated with migration direction from pooled genome-wide scans on a population of 233 migrating female mule deer (Odocoileus hemionus) where I identified genomic regions including FITM1, a gene linked to the formation of lipids, and DPPA3, a gene linked to epigenetic modifications of the maternal line. These results are consistent with the underlying genetic basis for a migratory trait which contributes to the additive genetic variance influencing migratory behaviours and can affect the adaptive potential of a species. Finally, in Chapter 4 I used a pedigree-free quantitative genetic approach to estimate heritability and sources of environmental variation in migration distance, timing, and movement rate of the same population of mule deer. I found low heritability for broad patterns of migration timing, and greater variation in heritability for behaviours during migration, with low heritability for distance and duration and high heritability for movement rate along the route. Insights into the genetic and environmental sources of variation for migration are critical both for the eco-evolutionary dynamics of migration behaviour, and for the conservation of species whose migrations may be vulnerable to environmental change. My thesis reveals that broad patterns of migration are driven largely by environmental effects while within these broad patterns, migration behaviour is driven to a measurable degree by genetic variation.

Author Keywords: heritability, migration, Odocoileus hemionus, reduced representation sequencing, whole genome sequencing

Rapidly changing landscapes introduce challenges for wildlife management, particularly for large mammal populations with long generation times and extensive spatial requirements. Understanding how these populations interact with heterogeneous landscapes aids in predicting responses to further environmental change. In this thesis, I profile American black bears using microsatellite loci and pooled whole-genome sequencing. These data characterize gene flow directionality and functional genetic variation to understand patterns of dispersal and local adaptation; processes key to understanding vulnerability to environmental change. I show dispersal is positively density-dependent, male biased, and influenced by food productivity gradients suggestive of source-sink dynamics. Genomic comparison of bears inhabiting different climate and forest zones identified variation in genes related to the cellular response to starvation and cold. My thesis demonstrates source-sink dynamics and local adaption in black bears. Population management must balance dispersal to sustain declining populations against the risk of maladaptation under future scenarios of environmental change.

Author Keywords: American black bear, Dispersal, Functional Genetic Variation, Gene Flow Directionality, Genomics, Local Adaptation

Understanding the complex genomic architecture underlying quantitative traits can provide valuable insight for the conservation and management of wildlife. Despite improvements in sequencing technologies, few empirical studies have identified quantitative trait loci (QTL) via whole genome sequencing in free-ranging mammal populations outside a few well-studied systems. This thesis uses high-depth whole genome pooled re-sequencing to characterize the molecular basis of the natural variation observed in two sexually selected, heritable traits in white-tailed deer (Odocoileus virginianus, WTD). Specifically, sampled individuals representing the phenotypic extremes from an island population of WTD for antler and body size traits. Our results showed a largely homogenous genome between extreme phenotypes for each trait, with many highly differentiated regions throughout the genome, indicative of a quantitative model for polygenic traits. We identified and validated several potential QTL of putatively small-to-moderate effect for each trait, and discuss the potential for real-world application to conservation and management.

Author Keywords: evolution, extreme phenotypes, genetics, genomics, quantitative traits, sexual selection