Graduate Theses & Dissertations

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Habitat Characteristics, Density Patterns and Environmental Niches of Indo-Pacific Humpback Dolphins (Sousa chinensis) of the Pearl River Estuary and Eastern Taiwan Strait
The purpose of this thesis is to quantify the habitat characteristics, density patterns and environmental niches of two groups of Indo-Pacific humpback dolphins: Chinese white dolphins (CWD) of the Pearl River estuary (PRE), and Taiwanese white dolphins (TWD, =Taiwanese humpback dolphin, THD) found in the eastern Taiwan Strait (ETS). Much work has already been done on the habitat use of CWDs in parts of the PRE, so the purpose of my first two chapters was to advance knowledge of the TWD to a comparable level. Chapter 2 contains the first published description of the relatively shallow, inshore, estuarine habitat of the TWD. General environmental characteristics and observed group sizes were consistent with other populations of humpback dolphins, and group sizes were not correlated with the environmental variables measured during surveys. Chapter 3 investigated density patterns of TWDs, finding spatiotemporal heterogeneity across the study area. Humpback dolphin densities fluctuated from year to year, but some parts of the study area were consistently used more than others. Environmental characteristics again did not influence dolphin densities, though more dolphins than expected were sighted in waters adjacent to major land reclamations, which may be related to the location of these areas close to major rivers. In Chapter 4, niches of the TWD and CWDs found in the PRE were compared using species distribution models, which indicated significant niche overlap. This may be due to niche conservatism maintaining similar fundamental niches between the two groups since their historical split >10,000 years ago, or a result of the intrinsic biotic factors that influence occurrence data affecting the hypervolume dimensions of each realized niche in similar ways. Geographic predictions indicate that most of the TWD’s range has likely been surveyed, and that there may be connectivity between PRE humpback dolphins and at least one neighbouring putative population due to continuous predicted suitable habitat in waters that remain poorly surveyed. Overall, my thesis demonstrates that density patterns may vary over time, but on a broad temporal scale, these two allopatric groups of Indo-Pacific humpback dolphins have similar habitat requirements in geographically isolated, but environmentally similar locations. Author Keywords: density, habitat, Indo-Pacific humpback dolphin, niche overlap, Sousa chinensis, species distribution model
Habitat use within and among roosts of chimney swifts (Chaetura pelagica)
Chimney swifts are listed as Threatened nationally and in many provinces within Canada due to rapid population declines. I examined large-scale spatial variation in the maximum size of chimney swift roosts at the northern edge of their range to identify where larger roosts occur. I used multi-sourced data collected across Ontario and Quebec between 1998 and 2013. I found that larger roosts were found at more northerly latitudes, and that very large roosts (>1000 birds) only occurred north of 45°. I also investigated fine-scale patterns of chimney swift positioning inside one of the largest roosts in Ontario. Using digitally recorded images, I calculated the angular position of swifts inside the roost relative to ambient and roost temperature. I found that swifts showed a strong preference for clinging to the south facing wall and clustered more when ambient air temperature was warmer. Thus, huddling in swifts provides additional or alternate benefits, other than serving purely to reduce costs of thermoregulation at low ambient temperatures. This research contributes to the understanding of chimney swift roosting ecology and identifies large roosting sites that should be retained for conservation. Author Keywords: chimney swift, communal roosting, conservation, group size, social thermoregulation, species-at-risk
Human Activity and Habitat Characteristics Influence Shorebird Habitat Use and Behaviour at a Vancouver Island Migratory Stopover Site
Pacific Rim National Park Reserve's 16 km of coastal beaches attract many thousands of people and shorebirds every year. To identify locations where shorebirds concentrate and determine the impact of human activity and habitat characteristics on shorebirds, I conducted shorebird and visitor surveys at 20 beach sectors during fall migration in 2011 to 2013 and spring migration in 2012 and 2013. The probability of shorebird presence decreased with increasing number of people at a beach sector. The time that shorebirds spent at a sector increased with increasing sector width. Close proximity to people increased the proportion of time shorebirds spent moving while shorebirds spent more time moving and less time foraging on wider beaches than on narrower ones. My findings suggest that placing restrictions on beach access and fast moving activities (e.g., running) may be necessary to reduce shorebird disturbance at Pacific Rim and similar stopover areas. Author Keywords: habitat use, human disturbance, predation risk, prey availability, shorebird, stopover
Influence of Habitat on Woodland Caribou Site Fidelity
Site fidelity is the behaviour of individuals to return to the same location; for female woodland caribou it may reflect reproductive success and depend on habitat quality. I investigated the influence of landscape and disturbance conditions on fidelity among three populations in Manitoba and Ontario, Canada. Habitat classifications were based on Forest Resource Inventory (FRI) and Landsat TM landcover maps. A total of 261 sites were ground-truthed to determine mapping accuracy. An amalgamated map incorporating FRI and Landsat TM data was estimated from field measurements to have an overall accuracy of 69.0%. Site fidelity was expressed as the distance between consecutive-year locations of individuals and was investigated during five week-long periods representing calving, early and late post-calving, winter, and breeding. Site fidelity was strongest during the post-calving seasons and weakest during the winter. Habitat had little influence on site fidelity in all seasons, excepting winter, even under highly disturbed conditions, suggesting maintenance of fidelity may be a maladaptive trait. Individual variation proved a strong predictor and cursory mapping indicated that caribou may return to sites visited two or more years earlier. Conservation management and policy should recognize that site fidelity may represent an ecological trap. Author Keywords: calving, disturbance, habitat, movement, Rangifer tarandus caribou, site fidelity
Intra-seasonal Variation in Black Tern Nest-site Selection and Survival
Resources and risk are in constant flux and an organism’s ability to manage change may improve their likelihood of persistence. I examined intra-seasonal variation in nest-site selection and survival of a declining wetland bird, the Black Tern (Chlidonias niger surinamensis). I modelled nest site occupancy and survival of early and late-nesting birds as a function of static and dynamic factors. Early-nesting birds selected nest sites based on the degree and direction of habitat change that occurred over the nesting cycle, while late-nesting birds selected sites based on static conditions near the time of nest-site selection. Nest age had the strongest influence on daily survival rate for both early and late-nesting birds, but the shape of this relationship showed intra-seasonal differences. Additionally, early-season survival improved slightly with increasing vegetation coverage and distance between conspecific nests, while late-season survival increased with clutch size. My results suggest that intra-seasonal variation in nest-site selection and survival is driven by changing habitat conditions and predator behavior. Author Keywords: Black Tern, Chlidonias niger surinamensis, daily survival rate, intra-seasonal variation, nest-site selection
Islands, ungulates, and ice
Central to wildlife conservation and management is the need for refined, spatially explicit knowledge on the diversity and distribution of species and the factors that drive those patterns. This is especially vital as anthropogenic disturbance threatens rapid large-scale change, even in the most remote areas of the planet. My dissertation examines theinfluence of land- and sea-scape heterogeneity on patterns of genetic differentiation, diversity, and broad-scale distributions of island-dwelling ungulates in the Arctic Archipelago. First, I investigated genetic differentiation among island populations of Peary caribou (Rangifer tarandus pearyi) in contrast to continental migratory caribou (Rangifer tarandus) and evaluated whether genetic exchange among Peary caribou island populations was limited by the availability of sea ice – both now and in the future. Differentiation among both groups was best explained by geodesic distance, revealing sea ice as an effective platform for Peary caribou movement and gene flow. With future climate warming, substantial reductions in sea ice extent were forecast which significantly increased resistance to caribou movement, particularly in summer and fall. Second, I assessed genetic population structure and diversity of northern caribou and deciphered how Island Biogeography Theory (IBT) and Central Marginal Hypothesis (CMH) could act in an archipelago where isolation is highly variable due to the dynamics of sea ice. Genetic differentiation among continental and island populations was low to moderate. In keeping with IBT and CMH, island-dwelling caribou displayed lower genetic diversity compared to mainland and mainland migratory herds; the size of islands (or population range) positively influenced genetic diversity, while distance-to-mainland and fall ice-free coastlines negatively influenced genetic diversity. Hierarchical structure analysis revealed multiple units of caribou diversity below the species level. Third, I shifted my focus to the terrestrial landscape and explored the elements governing species-environment relationships. Using species distribution models, I tested the response of caribou and muskoxen to abiotic versus abiotic + biotic predictors, and included distance to heterospecifics as a proxy for competitive interactions. Models that included biotic predictors outperformed models with abiotic predictors alone, and biotic predictors were most important when identifying habitat suitability for both ungulates. Further, areas of high habitat suitability for caribou and muskoxen were largely disjunct, limited in extent, and mainly outside protected areas. Finally, I modelled functional connectivity for two genetically and spatially disjunct groups of island-dwelling caribou. For High Arctic caribou, natural and anthropogenic features impeded gene flow (isolation-by-resistance); for Baffin Island caribou we found panmixia with absence of isolation-by-distance. Overall, my dissertation demonstrates the varying influences of contemporary land- and sea-scape heterogeneity on the distribution, diversity and differentiation of Arctic ungulates and it highlights the vulnerability of island-dwelling caribou to a rapidly changing Arctic environment. Author Keywords: Circuitscape, connectivity, Island Biogeography, landscape genetics, population structure, species distribution models
Linking large scale monitoring and spatially explicit capture–recapture models to identify factors shaping large carnivore densities
Understanding the spatial ecology of large carnivores in increasingly complex, multi-use landscapes is critical for effective conservation and management. Complementary to this need are robust monitoring and statistical techniques to understand the effect of bottom-up and top-down processes on wildlife population densities. However, for wide-ranging species, such knowledge is often hindered by difficulties in conducting studies over large spatial extents to fully capture the range of processes influencing populations. This thesis addresses research gaps in the above themes in the context of the American black bear (Ursus americanus) in the multi-use landscape of Ontario, Canada. First, I assess the performance of a widely adopted statistical modelling technique – spatially explicit capture-recapture (SECR) – for estimating densities of large carnivores (Chapter 2). Using simulations, I demonstrate that while SECR models are generally robust to unmodeled spatial and sex-based variation in populations, ignoring high levels of this variation can lead to bias with consequences for management and conservation. In Chapter 3, I investigate fine-scale drivers of black bear population density within study areas and forest regions by applying SECR models to a large-scale, multi-year black bear spatial capture-recapture dataset. To identify more generalizable patterns, in Chapter 4 I then assess patterns of black bear density across the province and within forest regions as a function of coarse landscape-level factors using the same datasets and assess the trade-offs between three different modeling techniques. Environmental variables were important drivers of black bear density across the province, while anthropogenic variables were more important in structuring finer-scale space use within study areas. Within forest regions these variables acted as both bottom-up and top-down processes that were consistent with ecological influences on black bear foods and intensity of human influences on the species’ avoidance of developed habitats. Collectively, this thesis highlights the opportunities and challenges of working across multiple scales and over expansive landscapes within a SECR framework. Specifically, the multi-scale approach of this thesis allows for robust inference of the mechanisms structuring fine and broad scale patterns in black bear densities and offers insight to the relative influence of top-down and bottom-up forces in driving these patterns. Taken together, this thesis provides an approach for monitoring large carnivore population dynamics that can be leveraged for the species conservation and management in increasingly human-modified landscapes. Author Keywords: animal abundance, black bear, capture-recapture, density estimation, statistical ecology, wildlife management
Mixed methods approaches in marine mammal science
This thesis explored the contribution of mixed methods approaches to marine mammal science through the use of concurrent and sequential designs to study distribution and feeding ecology of bowhead whales (Balaena mysticetus) in the Arctic region of Nunavik, Quebec, Canada. The study combines Inuit knowledge (IK), collected through semi-directed interviews with Inuit harvesters, and analyses of stable isotopes and trace elements (SI/TE) in baleen plates. A systematic literature review found that mixed methods are increasingly used in marine mammal ecology studies in remote locations, yet are still relatively rare and face a number of challenges. Both IK and SI/TE, indicated that bowhead whales have a seasonal pattern in their distribution and feeding ecology. They are most commonly present in productive nearshore areas in summertime, feeding in areas of great prey diversity, and moving to offshore areas in winter to fast. Mixed methods approaches used in this case study enabled the collection of complementary knowledge about bowhead whale ecology important for local management in a changing climate. This study also shows the value of mixed methods approaches for future marine mammal studies in Nunavik and elsewhere. Author Keywords: Arctic, bowhead whale, distribution, feeding ecology, mixed methods, traditional ecological knowledge
Population Genetics and Gut Microbiome Composition Reveal Subdivisions and Space Use in a Generalist and Specialist Ungulate
Natural populations are often difficult and costly to study, due to the plethora of confounding processes and variables present. This is of particular importance when dealing with managed species. Ungulates, for example, act as both consumers and prey sources; they also provide economic benefit through harvest, and as such, are of high ecological and economic value. I addressed conservation and management concerns by quantifying subdivision in wild populations and combined movement with non-invasive sampling to provide novel insight on the physiological drivers of space use in multiple species. This thesis explored biological patterns in ungulates using two distinct approaches: the first used molecular genetics to quantify gene flow, while the second examined the relationship between movement and the gut microbiome using high-throughput sequencing and GPS tracking. The goal of the first chapter was to quantify gene flow and assess the population structure of mountain goats (Oreamnos americanus) in northern British Columbia (BC) to inform management. I used microsatellites to generate genotype data and used a landscape genetics framework to evaluate the possible drivers behind genetic differentiation. The same analyses were performed at both a broad and fine scale, assessing genetic differentiation between populations in all of northern BC and in a case management study area northeast of Smithers BC. The results indicated panmixia among mountain goats regardless of scale, suggesting distance and landscape resistance were minimally inhibiting gene flow. Therefore, management at local scales can continue with little need for genetically informed boundaries, but regulations should be tailored to specific regions incorporating data on local access and harvest pressure. My second chapter aimed to determine the extent to which the gut microbiome drives space-use patterns in a specialist (mountain goat) and generalist (white-tailed deer, Odocoileus virginianus) ungulate. Using fecal samples, we generated genomic data using 16S rRNA high-throughput sequencing to evaluate gut diversity and gut microbiome characteristics. Additionally, individuals were fitted with GPS collars so that we could gain insight into movement patterns. Gut microbiome metrics were stronger predictors of space use and movement patterns with respect to home range size, whereas they were weaker predictors of habitat use. Notably, factors of both the gut microbiome and age of a given species were correlated with changes in space use and habitat use. Ultimately, this research linked high-throughput sequencing and GPS data to better understand ecological processes in wild ungulates. Author Keywords: gene flow, genomics, gut microbiome, home range, population genetic structure, ungulates
Prey abundance and habitat during the breeding season for Piping Plovers in the Ontario Great Lakes region
Similar to other shorebird trends around the world, the Piping Plover population (Charadrius melodus circumcinctus) is projected to decline if concerted conservation efforts are relaxed. To date, there is insufficient understanding of the connection between habitat type, prey abundance, and chick behaviour of the Piping Plover breeding population in Ontario. The aim of my thesis was to gain knowledge about prey abundance at recent and historic breeding locations, understanding how habitat influences prey abundance and chick behaviour across the Piping Plover breeding range in Ontario’s Great Lakes. The objective of my first study was to understand prey abundance across the breeding region Great Lakes of Ontario from 2018-2019, including occupied and unoccupied sites, and to quantify variation among habitats and periods of reproduction. To evaluate resources, I sampled 17 locations to compare prey abundance using invertebrate traps (n= 3,507). Sampling took place over the reproductive periods of nest initiation, post-hatch, and fledging and in four habitat types of shoreline, wrack, berm, and back dune. Occupied breeding sites had higher prey abundances, and different assemblages of invertebrate prey than unoccupied sites. Additionally, breeding sites had higher prey abundance during nest initiation and supported higher amounts of prey in shoreline and wrack habitat. The objective of my second study was to understand how habitat types influence chick behaviour. To evaluate behaviour-habitat trends, instantaneous chick observations were recorded at the four nest sites from the post-hatch to fledging stages. In total there were 23 fledged chicks that we observed across the two years. Chicks in this study spent 60.9% of their time foraging, 11.9% of the time displaying alert behaviour, 21.4% of their time resting or being brooded, and 5.9% of their time preening. Chicks spent a large proportion of time foraging in the shoreline, resting in the back dune, and alert in berm habitat. The frequency of these alert, defensive behaviours differed among sites, with Sauble Beach chicks spending more time in defensive behaviours compared to other sites. I concluded that in both nesting and brood-rearing periods, habitat is selected non-randomly by adult and young Piping Plovers to maximize access to invertebrate prey for growth and survival. Access by chicks to the most productive habitats should be considered in local management decisions. Author Keywords: chick behaviour, endangered, Great Lakes Region, habitat, Piping Plovers, prey abundance
Reintroducing species in the 21st century
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
Risk of Mortality for the Semipalmated Plover (Charadrius semipalmatus) Throughout Its Life Cycle
Three long-term mark and recapture/resight data sets of individually marked Semipalmated Plovers (Charadrius semipalmatus) were analyzed using Cormack-Jolly- Seber models. Data came from two breeding populations (Churchill, Manitoba, Canada, n=982, and Egg Island, Alaska, USA, n=84) and one overwintering population (Cumberland Island, Georgia, USA, n=62). For Alaska and Georgia, time-invariant models were best-supported, giving annual survival estimates of 0.67 (95%C.I.: 0.58- 0.76) and 0.59 (95%C.I.: 0.49-0.67) respectively. Data from Manitoba supported a timedependent model: survival estimates varied from 1.00 to 0.36, with lowest estimates from recent years, supporting observations of local population decline. Seasonal survival analysis of the Georgia population indicated lower mortality during winter (monthly Φoverwinter: 0.959, 95%CI: 0.871-0.988; for 6 month period Φoverwinter: 0.780 (0.440-0.929)) than during combined breeding and migratory periods (monthly ΦBreeding+Migration: 0.879 (0.825-0.918); for 8 month ΦBreeding+Migration: 0356 (0.215-0.504)). I recommend, based on high resight rates, continued monitoring of survival of wintering populations, to determine potential range-wide population declines. Keywords: survival, longevity, mortality, shorebird, overwinter, breeding, migration, life cycle Author Keywords: life cycle, longevity, mortality, non-breeding, shorebird, survival

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