Bowman, Jeff

Fishers (Pekania pennanti) were extirpated from many parts of Ontario in the early 20th century, but as of the early 2000s the species had recolonized most of its historical range. While the primary population genetic structure of fishers in central and eastern Ontario has not changed drastically over the past ten years, we did find evidence of increased secondary structure and a reduction in northward movement from southeastern Ontario, a site of recent immigration from the Adirondacks in northern New York. This may be indicative of a reduction in density and thus in density-dependent migration, or it may be a consequence of the population reaching equilibrium following a period of rapid expansion associated with recolonization. We also observed no variation within central and eastern Ontario at 14 of 15 candidate functional loci we screened, suggesting possible directional or stabilizing selection and a lack of adaptive potential.

Author Keywords: fisher, functional genes, Ontario, Pekania pennanti, population genetics, recolonization

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

Currently there are large areas of the North American landscape that are occupied by Canis spp. hybrids of several varieties, leading to the logical question as to the genetic structure and ecological function of Canis populations across the continent, and to what extent hybrids reflect contemporary landscapes. This study illustrated patterns of niche differentiation between parental canid species and their hybrids using individual high quality genetic profile and species distribution models to support the intermediate phenotype hypothesis. In general, hybrids demonstrated an intermediate habitat suitability compared to its parental species, across most environmental variables used. A similar trend was observed in the niche metric analysis, where we found that hybrids exhibit intermediate niche breadth, with eastern coyotes and eastern wolves exhibiting the broader and narrower niche, respectively. Our results demonstrate that the intermediate phenotype hypothesis is supported even at a large scale and when involving highly mobile large mammal species.

Author Keywords: canid, ecological niche modelling, hybridization, intermediate phenotype, microsatellite genotype, niche differentiation

Variation in habitat quality and disturbance levels can strongly influence a species' distribution, leading to spatial variation in population density and influencing population dynamics. It is therefore critical to understand how density can lead to variability in demographic responses for effective conservation and recovery of species. My dissertation illustrates how density and spatial familial networks can be integrated together to gain a better understanding of the influence of density on population dynamics of boreal caribou. First, I created an analytical framework to assess results from empirical studies to inform spatially-explicit capture-recapture sampling design, using both simulated and empirical data from noninvasive genetic sampling of several boreal caribou populations in Alberta, Canada, which varied in range size and estimated population density. Analysis of the empirical data indicated that reduced sampling intensity had a greater impact on density estimates in smaller ranges, and the best sampling designs did not differ with estimated population density but differed between large and small population ranges. Secondly, I used parent-offspring relationships to construct familial networks of boreal caribou in Saskatchewan, Canada to inform recovery efforts. Using network measures, I assessed the contribution of individual caribou to the population with several centrality measures and then determined which measures were best suited to inform on the population demographic structure. I found substantial differences in the centrality of individuals in different local areas, highlighting the importance of analyzing familial networks at different spatial scales. The network revealed that boreal caribou in Saskatchewan form a complex, interconnected familial network. These results identified individuals presenting different fitness levels, short- and long-distance dispersing ability across the range, and can be used in support of population monitoring and recovery efforts. Finally, I used a spatial capture-recapture analytical framework with covariates to estimate spatial density of boreal woodland caribou across the Saskatchewan Boreal Plains, and then reconstructed parent-offspring relationships to create a familial network of caribou and determined whether spatial density influenced sex-specific network centrality, dispersal distance, individual reproductive success, and the pregnancy status of females. I show that caribou densitygreatly varied across the landscape and was primarily affected by landscape composition and fragmentation, and density had sex-specific influences on dispersal distance, reproductive success, and network centrality. The high density areas reflected good-quality caribou habitat, and the decreased dispersal rates and female reproductive output suggest that these remnant patches of habitat may be influencing demographic responses of caribou.

Author Keywords: boreal caribou, density, familial networks, population dynamics, rangifer tarandus caribou, spatial capture-recapture

The Pearl River Estuary (PRE) and eastern Taiwan Strait (ETS) populations of Indo-Pacific humpback dolphins (Sousa chinensis) contain ~600 and 100 catalogued individuals, respectively. Population demography is important for conservation actions but few animals have been sexed by conventional methods. Longitudinal analysis of tooth-rake scars on dorsal fins showed scars disappear within 7 months; sexing adults by scarring is likely not impacted by juvenile scarring. Using dorsal fin photographs, sex assigned for 87% of catalogued PRE adults (n=300) and for 93% of ETS adults (n=60), using scars hypothesized from male-male competition, was in concordance with sex assigned by DNA, calf association, and ventral photographs. Scarring was higher in presumed males than females and in PRE females than ETS females. Female:male sex ratios were 3:2 (PRE) and 2:1 (ETS), though this likely results from biases in photo-identification methods. Social analysis with presumed sexes showed strong female-female associations in both populations but stronger female-male and male-male associations in PRE. These results support sex differentiation by scarring, which was a non-invasive approach, and sex assignment for many PRE and ETS individuals.

Author Keywords: Indo-Pacific humpback dolphins, photo-identification, scarring, sex differentiation, sexes, social structure

Current major issues in conservation biology include habitat loss, fragmentation and population over-exploitation. Animals can respond to landscape change through behavioural flexibility, allowing individuals to persist in disturbed landscapes. Individual behaviour has only recently been explicitly included in population models. Carnivores may be sensitive to changing landscapes due to their wide-ranging behaviour, low densities and reproductive rates. Canada lynx (Lynx canadensis) is a primary predator of snowshoe hares (Lepus americanus). Both species range throughout the boreal forests of North America, however lynx are declining in the southern range periphery. In this dissertation, I developed new insights into the effects of habitat loss and fragmentation on lynx. In Chapter 2, I created a habitat suitability model for lynx in Ontario and examined occurrence patterns across 2 regions to determine if habitat selection is flexible when different amounts of habitat are available. Although lynx avoided areas with <30% suitable habitat where suitable land cover is abundant, I found that they have flexible habitat selection patterns where suitable land cover is rare and occurred in low habitat areas. In Chapter 3, I investigated the effects of dispersal plasticity on occupancy patterns using a spatially explicit individual-based model. I showed that flexible dispersers, capable of crossing inhospitable matrix, had higher densities and a lower risk of patch extinction. In contrast, inflexible dispersers (unable to cross inhospitable matrix), were most limited by landscape connectivity, resulting in a high extinction risk in isolated patches. I developed three predictions to be explored with empirical data; (1) dispersal plasticity affects estimates of functional connectivity; (2) variation in dispersal behaviour increases the resilience of patchy populations; and (3) dispersal behaviour promotes non-random distribution of phenotypes. Finally, in Chapter 4, I examined the consequences of anthropogenic harvest on naturally cycling populations. I found that harvest mortality can exacerbate the effects of habitat fragmentation, especially when lynx densities are low. Dynamic harvest regimes maintained lynx densities and cycle dynamics while reducing the risk of population extinction. These results suggest that lynx display some flexibility to changing landscapes and that the metapopulation structure is more resilient to increasing habitat loss and fragmentation than previously understood. Future studies should focus on determining a threshold of connectivity necessary for population persistence and examining the effects of habitat loss on the fecundity of lynx.

Author Keywords: Fluctuating Populations, Habitat Fragmentation, Landscape Ecology, Occupancy Dynamics, Population Ecology, Spatially Explicit Population Models

The structure and composition of the landscape can facilitate or impede gene flow, which can have important consequences because genetically isolated groups of individuals may be prone to inbreeding depression and possible extinction. My dissertation examines how landscape structure influences spatial patterns of genetic differentiation and diversity of American marten (Martes americana) and Canada lynx (Lynx canadensis) in Ontario, Canada, and provides methodological advances useful for landscape geneticists. First, I identified the effects of map boundaries on estimates of landscape resistance, and proposed a solution to the bias: a buffer around the map boundary. Second, I assessed the sensitivity of a network-based estimate of genetic distance, conditional genetic distance, to incomplete sampling. I then used these landscape genetic tools in a pairwise, distance-based analysis of 653 martens genotyped at 12 microsatellite loci. I evaluated whether forest management in Ontario has influenced the genetic structure of martens. Although forest management practices had some impact, isolation by distance best described marten gene flow. Our results suggest that managed forests in Ontario are well connected for marten and do not impede marten gene flow. Finally, I used a site-based analysis of 702 lynx genotyped at 14 microsatellite loci to investigate spatial patterns of genetic diversity and differentiation at the trailing (contracting) edge of the lynx distribution in Ontario. I analyzed harvest records and found that the southern edge of lynx range has contracted by >175 km since the 1970s. I also found that neutral genetic diversity decreased towards the trailing edge, whereas genetic differentiation increased. Furthermore, I found strong correlations between gradients of lynx genetic structure and gradients of climate and land cover in Ontario. My findings suggest that increases in winter air temperature, decreases in snow depth, and loss of suitable habitat will result in further loss of genetic diversity in peripheral populations of lynx. Consequently, the adaptive potential of lynx populations on the southern range periphery could decline. In conclusion, my dissertation demonstrates the varying influences that contemporary landscape structure and climate gradients can have on genetic diversity and differentiation of different species.

Author Keywords: Circuitscape, genetic network, landscape genetics, Lynx canadensis, Martes americana, range shift

Many facilities attempt to alleviate the risk of chronic stress in captivity by providing environmental enrichment shown to minimize behavioural disorders and stress in several species. One potential form of enrichment used in zoos is training animals to perform rides for guests, however, the effect of this activity on the welfare of individual animals has never been examined. I validated the use of saliva for assessing stress in dromedary camels (Camelus dromedarius) an animal commonly used for rides. I then measured variation in salivary cortisol in four male camels during animal rides for guests at the Toronto Zoo. The camels were sampled during the ride season (from June to August) using four treatments: 1) in their pasture, 2) at the ride area not performing rides, 3) performing a low number of rides (n=50/day) and 4) performing a high number of rides (n=150/day). Furthermore, samples were taken before and after the ride season for comparison. There was a significant difference between the post-ride season treatment and the three treatments involving guest presence during the ride season (ride area, low rides, high rides. This indicates that performing rides is not a stressful experience based on the stress metrics I used, and suggests that rides may be a form of enrichment for dromedary camels.

Author Keywords: ACTH challenge, animal welfare, camels, environmental enrichment, salivary cortisol, stress

I used a GIS raster layer of an area in the Churchill, Manitoba region to investigate the effect of breeding habitat on demography and density of Whimbrel from 2010 through 2013. Program MARK was used to quantify adult and daily nest survival. Apparent annual survival of 0.73 ± 0.06 SE (95% CI = 0.60-0.83) did not significantly differ between sexes or habitats and was lower than expected based on longevity records and estimates for other large-bodied shorebirds. Nest success, corrected for exposure days, was highly variable, ranging from a low of 3% (95% CI = 0-12%) in 2011 to a high of 71% (95% CI = 54-83%) in 2013. The highest rate of nest survival occurred in the spring with the warmest mean temperature. I developed a generalized linear model (GLM) with a negative-binomial distribution from random plots that were surveyed for abundance to extrapolate a local breeding population size of 410 ± 230 SE and density of 3.2 birds per square km ± 1.8 SE. The result of my study suggests that other aspects of habitat not captured by the land cover categories may be more important to population dynamics.

Author Keywords: abundance, apparent survival, curlew, land cover map, nest-site fidelity, nest success

Since their successful reintroduction, the eastern wild turkey (Meleagris gallopavo silvestris) has expanded its range north. Due to different and potentially more severe limiting factors, management approaches generalized from studies within the historical range may not be appropriate to apply to northern populations. To better understand northern wild turkey ecology, GPS and VHF transmitters were used to track habitat selection and survival of female turkeys at the species northern range edge in Ontario, Canada. These northern turkeys exhibited larger seasonal home range sizes relative to those in their historical range, and selected deciduous forest and pasture and fields within the study area. Supplemental food was also selected by turkeys when choosing autumn and winter ranges. The northern turkeys also suffered a low annual survival rate, and high mortality from predation. These findings underscore the challenges of maintaining turkey populations in northern environments, and will help inform management strategies.

Author Keywords: Eastern Wild Turkey, Euclidean distance analysis, Habitat selection, Meleagris gallopavo silvestris, Northern range edge, Survival