Graduate Theses & Dissertations

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evolutionary ecology of Alaska's mountain goats with management implications
The integration of genetic and environmental information can help wildlife managers better understand the factors affecting a species’ population structure and their response to disturbance. This thesis uses genetic techniques to assess the broad and fine scale population structure of mountain goats in Alaska. The first chapter aims to determine the number of genetically distinct subpopulations and model the demographic history of mountain goats in Alaska. The second chapter investigates the population structure and demographic history of mountain goats in Glacier Bay National Park and examines the impact that climate change will have on these mountain goats. My results indicate that Alaska has eight subpopulations which diverged during the Wisconsin glaciation. In Glacier Bay, population structure is reflective of the landscape during colonization, and mountain goat population density and movement corridors are likely to decline due to future climate change. Author Keywords: Alaska, biogeography, gene flow, landscape genetics, mountain goat, population genetic structure
Using genomic and phenotypic data to explore the evolution and ecology of the North American mountain goat
Evaluating the impact of climate change is arguably one of the main goals of conservation biology, which can be addressed in part by studying the demographic history of species in the region of interest. In North America, landscape and species composition during the most recent Pleistocene epoch was primarily influenced by glaciation cycles. Glacial advance and retreat caused species ranges to shift as well, leaving signatures of past population bottlenecks in the genetic code of most species. Genomic tools have shown to be important tools for understanding these demographic events to enhance conservation biology measures in several species. In my thesis I first reviewed the state of ungulate genomics, with a focus on how such data sets can be used in understand demography, adaptation, and inform conservation and management. Importantly, the review introduces key analyses like the pairwise sequentially Markovian coalescent and features like variation in antlers and horns and selection pressures that are used throughout subsequent chapters. Using the North American mountain goat as a model species, I then explored the genomic and phenotypic variation in this alpine specialist mammal. Starting with the generation of the first genome assembly for the mountain goat, I identified genes unique to the mountain goat and modeled demographic history going back millions of years using a pairwise sequentially Markovian coalescent approach. Species’ effective population size generally paralleled climatic trends over the past one hundred thousand years and severely declined to under a thousand individuals during the last glacial maximum. Given the biological importance of horns in mountain goats and the recent scientific interest in genetic basis of headgear, I analyzed over 23,000 horn records from goats harvested in British Columbia, Alaska and Northwest Territories from 1980 to 2017. Overall, variation in horn size over space and time was low; goats harvested further North had shorter horn lengths and smaller horn circumferences in one year old and 4 years and older age classes and 4 years and older age class, respectively. Proximity of roads, which was used as an indicator of artificial selection, had a small effect on horn size, with larger horns being harvested closer to major roads. Finally, I used two range-wide genomic data sets sequenced with a whole genome re-sequencing and reduced representation approaches to provide estimates of genetic diversity, contemporary effective population sizes and population structure. These insights can help inform management and will potentially make an impact in preserving the mountain goat. Author Keywords: genome assembly, horn size, Oreamnos americanus, population demography, reduced representation sequencing, whole genome resequencing
Using environmental DNA (eDNA) metabarcoding to assess aquatic plant communities
Environmental DNA (eDNA) metabarcoding targets sequences with interspecific variation that can be amplified using universal primers allowing simultaneous detection of multiple species from environmental samples. I developed novel primers for three barcodes commonly used to identify plant species, and compared amplification success for aquatic plant DNA against pre-existing primers. Control eDNA samples of 45 plant species showed that species-level identification was highest for novel matK and preexisting ITS2 primers (42% each); remaining primers each identified between 24% and 33% of species. Novel matK, rbcL, and pre-existing ITS2 primers combined identified 88% of aquatic species. The novel matK primers identified the largest number of species from eDNA collected from the Black River, Ontario; 21 aquatic plant species were identified using all primers. This study showed that eDNA metabarcoding allows for simultaneous detection of aquatic plants including invasive species and species-at-risk, thereby providing a biodiversity assessment tool with a variety of applications. Author Keywords: aquatic plants, biodiversity, bioinformatics, environmental DNA (eDNA), high-throughput sequencing, metabarcoding
Using DNA Barcoding to Investigate the Diet and Food Supply of a Declining Aerial Insectivote, the Barn Swallow (Hirundo rustica)
Barn Swallow (Hirundo rustica) populations have declined in North America over the past 40 years and they are listed as Threatened in Ontario, Canada. Changes in the food supply have been hypothesized as a potential cause of this population decline. I used DNA barcoding to investigate the diet and food supply of Barn Swallows and to determine if the food supply affects their reproductive performance. In two breeding seasons, I monitored nests, collected fecal samples, and monitored prey availability by collecting insects from the habitat surrounding breeding sites using Malaise traps. I used DNA barcoding to identify insect specimens collected from the habitat and to identify prey items from Barn Swallow nestling fecal samples. I found that Barn Swallow nestlings were fed a very broad range of prey items but were fed larger prey items more frequently. Prey availability was not related to the timing of reproduction, the number of nests at a breeding site, or the reproductive output of individual nests. This study provides information on the diet composition of Barn Swallows in North America and suggests that food limitation during the breeding season may not be a major factor in their population decline. Author Keywords: aerial insectivore, diet, DNA barcoding, Hirundo rustica, metabarcoding, reproductive success
Understanding the establishment of Typha spp. in North America using population genetics and common garden studies
There are three cattail (Typha) taxa in Canada: T. latifolia (native), T. angustifolia (introduced), and their hybrid T. x glauca. The latter is invasive in regions around the Laurentian Great Lakes, and I investigated the potential role that commercial suppliers may be playing in the introduction of non-native Typha by comparing genotypes of North American, European, and commercially available plants. I found that Ontario garden centres are importing both hybrids and non-native lineages of T. angustifolia into Canada, but was unable to identify the provenance of T. latifolia. I also investigated the possibility that the hybrid cattail leaf litter shade and leachate influences germination and early growth of the parental species of the hybrids. Using three common garden experiments, I found that T. x glauca leaf litter suppresses germination rates of the three taxa. In the early seedling growth experiment, plant performance varied by taxa, and for the competition experiment there were no intra- or interspecific competition or treatment effects on the performance of plants. Overall, my research identified a potential mechanism allowing T. x glauca to dominate wetlands, and also shows that non-native lineages are being introduced into Canada through commercial trade Author Keywords: Competition, Germination, Non-native lineages, Plant nurseries, Seedling Growth, Typha spp.
Time to adapt
To better understand species’ resilience to climate change and implement solutions, we must conserve environments that maintain standing adaptive genetic variation and the potential generation of new beneficial alleles. Coding trinucleotide repeats (cTNRs) providing high-pace adaptive capabilities via high rates of mutation are ideal targets for mitigating the decline of species at risk by characterizing adaptively significant populations. Ultimately, adaptive genetic information will inform the protection of biological diversity below the species level (i.e., “Evolutionarily Significant Units” or “ESUs”). This dissertation investigates cTNRs within candidate genes to determine their prevalence and influence under selection in North American mammals. First, I evaluated the potential for somatic mosaicism in Canada lynx (Lynx canadensis), and found that tissue-specific mosaicism does not confound cTNR genotyping success in lynx. Second, I assessed a selection of clock gene cTNRs across characterized mammals and found that these repeats are abundant and highly variable in length and purity. I also identified preliminary signatures of selection in 3 clock gene cTNRs in 3 pairs of congeneric North American mammal species, highlighting the importance of cTNRs for understanding the evolution and adaptation of wild populations. I further evaluated the influence of selection on the NR1D1 cTNR within Canada lynx sampled across Canada using environmental correlation, where I estimated the variation in NR1D1 cTNR alleles explained by environmental and spatial variables after removing the effects of neutral population structure. Although most variation was explained by neutral structure, environment and spatial patterns in eastern lynx populations significantly explained some of the variation in NR1D1 alleles. To examine the role of island populations in the generation and distribution of adaptive genetic variation, I used 14 neutral microsatellites and a dinucleotide repeat within a gene linked to mammalian body size, IGF-1, and found that both genetic drift and natural selection influence the observed genetic diversity of insular lynx. Finally, I estimated the divergence dates of peripheral lynx populations and made recommendations towards the conservation of Canada lynx; high levels of genetic differentiation coupled with post-glacial colonization histories and patterns of divergence at cTNR loci suggest at least 4 ESUs for Canada lynx across their range. Author Keywords: adaptation, Canada lynx, candidate genes, coding trinucleotide repeats, evolution, natural selection
Testing for Interspecific Hybridization and a Latitudinal Cline Within the Clock Gene Per1 of the Deer Mouse (Peromyscus maniculatus) and the White-Footed Mouse (Peromyscus leucopus)
The recent northward expansion of the white-footed mouse (Peromyscus leucopus) in response to climatic changes provides a natural experiment to explore potential adaptive genetic variation within the clock gene Per1 in Peromyscus undergoing latitudinal shifts, as well as, the possibility of hybridization and introgression related to novel secondary contact with its sister species the deer mouse (Peromyscus maniculatus). Because clock genes influence the timing of behaviors critical for survival, variations in genotype may reflect an organism’s ability to persist in different environments. Hybridization followed by introgression may increase the adaptive potential of a species by quickly generating adaptive variation through novel genetic recombination or by the transfer of species-specific alleles that have evolved in response to certain environments. In chapter 2, I used microsatellite and mtDNA markers to test for hybridization and introgression between P. maniculatus and P. leucopus and found that interbreeding is occurring at a low frequency (<1%). In chapter 3, I tested for a latitudinal cline in a polyglycine repeat located within the Per1 gene of Peromyscus and discovered a putative cline in the Per1-142 and Per1-157 allele of P. leucopus and P. maniculatus, respectively. Chapter 4, further expands upon these findings, limitations, and the lack of evidence supporting introgression at the Per1 locus. Despite this lack of evidence, it is possible that novel hybridization has or could lead to adaptive introgression of other genes, allowing for the exchange of adaptive alleles or traits that could be advantageous for range expansion and adaption to future environmental changes. Author Keywords: Clock genes, Hybridization, Latitudinal gradient, Per1, Peromyscus, Range Expansion
Social discrimination by female polar bears (Ursus maritimus) when accompanied by dependent offspring during the ice-free season in southern and western Hudson Bay and James Bay
Polar bears are generally described as solitary, but features of their life cycles and habitats regularly necessitate interaction. Effective conspecific assessment, including accurate recognition and discrimination, likely confers benefits, especially to females accompanied by dependent young. Individuals in the Southern (SH) and Western (WH) Hudson Bay subpopulations are ideal for studying polar bear social behaviours because of the prolonged high densities of the ice-free season. First, I looked outside family groups to model their fine scale sociospatial organization on land. Capture locations were more likely to correspond to family groups when there were fewer neighbouring bears, when a greater proportion of neighbours were female, and when the focal individual and neighbours were significantly related. Second, I looked within the family group to assess offspring recognition. Of 288 offspring in 207 family groups captured in the SH subpopulation from 1999 through 2013, only one case of adoption (of a singleton) was observed. Author Keywords: Adoption, Kin Recognition, Logistic Regression, Maternity Analysis, Social Discrimination, Sociospatial
Sex-Specific Graphs
Sex-specific genetic structure is a commonly observed pattern among vertebrate species. Facing differential selective pressures, individuals may adopt sex-specific life historical traits that ultimately shape genetic variation among populations. Although differential dispersal dynamics are commonly detected in the literature, few studies have investigated the potential effect of sex-specific functional connectivity on genetic structure. The recent uses of Graph Theory in landscape genetics have demonstrated network capacities to describe complex system behaviors where network topology intuitively represents genetic interaction among sub-units. By implementing a sex-specific network approach, our results suggest that Sex-Specific Graphs (SSG) are sensitive to differential male and female dispersal dynamics of a fisher (Martes pennanti) metapopulation in southern Ontario. Our analyses based on SSG topologies supported the hypothesis of male-biased dispersal. Furthermore, we demonstrated that the effect of the landscape, identified at the population-level, could be partitioned among sex-specific strata. We found that female connectivity was negatively affected by snow depth, while being neutral for males. Our findings underlined the potential of conducting sex-specific analysis by identifying landscape elements that promotes or impedes functional connectivity of wildlife populations, which sometimes remains cryptic when studied at the population level. We propose that SSG approach would be applicable to other vagile species where differential sex-specific processes are expected to occur. Author Keywords: genetic structure, Landscape Genetics, Martes pennanti, Population Graph, sex-biased dispersal, Sex-Specific Graphs
Selection on functional genes across a flying squirrel (genus Glaucomys) hybrid zone
While hybridization between distinct taxa can have undesirable implications, it can also result in increased genetic variability and potentially, the exchange of adaptive genes or traits. Adaptive variation acquired through introgressive hybridization may be particularly advantageous for species facing rapid environmental change. I investigated a novel, climate change-induced hybrid zone between two flying squirrel species: the southern (Glaucomys volans) and northern (G. sabrinus) flying squirrel. I was interested in the occurrence of hybridization and introgression, the type of selective pressures maintaining the hybrid zone and the potential for adaptive introgression. I found relatively low hybridization and introgression frequencies (1.7% and 2.9% of the population, respectively) and no evidence of selection on hybrids or backcrosses in particular environments. I conclude that the data are more consistent with a hybrid zone maintained by endogenous (environment-independent) selection. I tested for adaptive introgression using two functional genes: IGF-1 and CLOCK. I documented intermediate functional allele frequencies in backcrosses compared to parental populations, suggesting the alleles do not confer fitness advantages in backcrosses. Despite lack of evidence for current adaptive introgression, genetic admixture between G. volans and G. sabrinus may provide adaptive potential should these species face more rapid or drastic environmental change in the future. Author Keywords: adaptive introgression, flying squirrel, Glaucomys sabrinus, Glaucomys volans, hybridization, introgression
SPATIAL AND TEMPORAL GENETIC STRUCTURE OF WOLVERINE POPULATIONS
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
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

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