Zoology

Arctic breeding shorebirds are often considered alternate or secondary prey to avian and terrestrial predators that primarily feed on cyclic populations of rodents. Predation risk can strongly influence habitat selection; however, there is limited understanding of how nest placement by shorebirds and subsequent risk of nest predation are affected by spatiotemporal patterns involving the indirect effects of alternate prey on predator activity. I examined nest fates of Whimbrel (Numenius phaeopus) and Dunlin (Calidris alpina) at the southernmost edge of their breeding ranges at Polar Bear Provincial Park, Ontario, using field surveys including direct observations, camera monitors and thermal probes. I used resource selection functions (RSFs) to compare used and available habitat features near shorebird nests. I assessed whether shorebird nest predation was affected by spatial patterns in avian predator activity, habitat, and small mammal alternate prey and annual variation in cyclic small mammal abundance. RSFs confirmed strong habitat selection for graminoid fen habitats with avoidance of tundra heath, presumably to avoid terrestrial predators that may use them as travel corridors. Shorebird nest predation was not affected by the spatial variation in avian predator activity. However, Dunlin nest predation risk was affected by spatial variation in small mammal abundance. Further, the effect was strongest in years consistent with peaks in the small mammal population cycle. These findings highlight the important effects of spatiotemporal variation of alternate prey densities on Arctic-Subarctic food webs, providing insights on the complex relationships affecting shorebird nest success.

Author Keywords: apparent competetion, Dunlin, Empirical Bayesian Kriging, Shorebird, Subarctic, Whimbrel

Globally, fledglings of more than 50 seabird species are vulnerable to stranding in human settlements near their colonies as they leave their nests for the first time. This phenomenon has been attributed to attraction toward and disorientation by artificial light at night (ALAN). Grassroots rescue programs find more birds in illuminated locations than in dark ones, but it is unclear to what extent this pattern is driven by seabird versus rescuer behaviour. I studied behavioural responses to ALAN in stranding-prone Atlantic puffin (Fratercula arctica) and Leach's storm-petrel (Hydrobates leucorhous) fledglings and investigated detection probabilities of 3D-printed puffin fledgling decoys by rescuers. I hypothesized that seabird strandings are due to attraction toward ALAN; that post-stranding behaviour varies with taxon and lighting conditions; and that post-stranding behaviour (e.g., concealment) and lighting conditions drive variance in detection probability of stranded seabirds by rescuers. To address my first hypothesis, I alternately illuminated two beaches near a Newfoundland puffin colony and found significantly more puffin fledglings stranded when the beaches were illuminated than when they were dark, supporting the light attraction hypothesis. Stranded puffins similarly moved toward light over darkness in a Y-maze choice experiment and were less active under LED light than in darkness or under high-pressure sodium light, demonstrating a preference toward certain light conditions post-stranding. In contrast, storm-petrels predominantly avoided lit conditions in a Y-maze and were less active in darkness than under light. Further, when provided with a place to hide, more than half of storm-petrels concealed themselves when in light conditions versus none in darkness. To test my third hypothesis, I deployed puffin decoys and found that rescuers detected them more easily in higher light levels, in less concealed locations, and with increased search effort. My results support historical deductions that seabird fledgling strandings are caused by attraction toward artificial light but demonstrate that post-stranding behavioural responses to ALAN depend on species and light type, which may in turn affect detection probability by rescuers.

Author Keywords: artificial light, atlantic puffin, charadriiform, Leach's storm-petrel, light attraction, procellariiform

White-nose syndrome (WNS) is a skin disease of bats caused by the fungus Pseudogymnoascus destructans (Pd) that damages flight membranes during hibernation and can lead to death. The disease causes mortality of multiple bat species in eastern North America and is spreading into western North America. Future impacts of WNS on naïve bat populations are unknown. Variation in host susceptibility occurs among and within species, but mechanisms driving this variation are unclear. Multiple studies have characterized immunological responses to WNS, but skin physiology as a barrier to pathogens is understudied. The unique ability of Pd to actively penetrate the normal, intact skin of its mammalian host makes WNS an interesting study system to understand skin defenses. Aspects of the mammalian skin environment that can influence disease susceptibility include pH, sebaceous lipids, and microbiomes. I found skin mycobiomes of WNS-susceptible species had significantly lower alpha diversity and abundance compared to bat species resistant to Pd infection. Using these data, I predicted that most naïve bat species in western North America will be susceptible to WNS based on the low diversity of their skin mycobiomes. Some fungi isolated from bat wings inhibited Pd growth in vitro, but only under specific salinity and pH conditions, suggesting the microenvironment on wings can influence microbial interactions and potentially WNS-susceptibility. I measured the wing-skin pH of bats in eastern Canada and found that Eptesicus fuscus (WNS-tolerant) had more acidic skin than M. lucifugus (WNS-susceptible). Differences in sebum quantity and composition among and within mammalian species may help explain variation in skin disease susceptibility and the composition of skin microbiomes. This is due to the antimicrobial properties of sebum and the use of sebum as a nutrition source by microbes. Outcomes of this work further our understanding of inter- and intra-specific differences among bat species and individuals in skin mycobiomes and physiology, which may contribute to variation in WNS-susceptibility. Future research should focus on characterizing the physical and chemical landscape of skin as this is essential for understanding mechanisms structuring skin microbial assemblages and skin disease susceptibility in wildlife.

Author Keywords: bat, fungi, microbiome, mycology, physiology, white-nose syndrome

Line transect sampling and photographic-identification (photo-ID) are common survey techniques for estimating the abundance and distribution of cetaceans. Combining these approaches in the field ('combined LTPI' surveys) and using data from both components has the potential for generating comprehensive ecological knowledge that can be far more valuable than when these techniques and their data are used independently. In this thesis, I evaluated the results and conclusions from these two methods, used singly and in tandem, by investigating the population dynamics of two humpback dolphin (Sousa chinensis spp.) populations: the large and widely distributed Chinese white dolphin (S. c. chinensis) of the Pearl River estuary (PRE), and the small and geographically isolated subspecies of Taiwanese white dolphin (S. c. taiwanensis) in the eastern Taiwan Strait. Data from combined LTPI surveys in Hong Kong waters, at the eastern edge of the PRE, revealed a shift in space use with individuals spending less time in these waters than at the start of surveys. Data from combined LTPI surveys in Taiwan provided further support for a subspecies restricted to the central western waters, and identified a commonly used area at the northern part of their limited range. These two case studies demonstrated an overall efficacy of combined LTPI surveys in ecological studies of cetaceans. However, a multi-criteria analysis revealed that combined LTPI surveys with a line transect focus (e.g., Hong Kong) performed better than a LTPI survey with a photo-ID focus (e.g., Taiwan) when considering ecological aspects of the study populations, labour and data requirements, and ecological output. Even so, the photo-ID focus of Taiwan's monitoring program led to better assessments of individual space use patterns, likely helped by the Taiwanese white dolphin population's smaller size and intensive photographic effort. In both cases, the ecological output of combined LTPI surveys could be improved by expanding the study area or extending the field season or frequency of surveys. Overall, I showed that by following a set of general guidelines, different iterations of the combined LTPI approach (i.e., photo-ID focus or LT focus) can serve as powerful tools for uncovering multi-dimensional ecological information on cetaceans.

Author Keywords: abundance, cetacean, distribution, line transect sampling, multi-criteria analysis, Photo-ID

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

Understanding breeding and migration habits of Whimbrel (Numenius phaeopus) in the Hudson Bay Lowlands is important for the conservation of this population. I monitored Whimbrel at two breeding sites: the Churchill region of Manitoba and Burntpoint, Ontario. Annual average nest initiation timing was highly variable and successful nests were initiated significantly earlier than those that failed. Although nests were initiated significantly earlier at Burntpoint than Churchill, annual nest success quantified in program R MARK was similar across sites. Observed nest success rates were lower than historical records and most failure was due to predation. Annual nest survival varied widely and I used a generalized linear model to relate annual nest survival to annual average weather conditions. I observed weak relationships between annual nest survival and weather conditions in the northbound staging grounds. I tracked post-breeding migratory movements using the MOTUS radio telemetry system and observed consistent use of the mid-Atlantic coast of the United States during migration, especially among birds emerging from Churchill. In Burntpoint, I observed more variability in post-breeding migratory trajectories and significantly earlier post-breeding departure as compared to Churchill. The results of my study suggest differences in breeding and migration habits exist across nearby breeding populations, indicating that there is a need for population-specific conservation approaches for this declining species.

Author Keywords: Migration, Movement Ecology, Nesting Ecology, Nest Success, Shorebird conservation, Whimbrel

Blackfin Cisco (Coregonus nigripinnis), a deepwater cisco species once endemic to the Laurentian Great Lakes, was discovered in Algonquin Provincial Park in four lakes situated within a drainage outflow of glacial Lake Algonquin. Blackfin habitat preference was examined by analyzing which covariates best described their depth distribution using hurdle models in a multi-model approach. Although depth best described their distribution, the nearly isothermal hypolimnion in which Blackfin reside indicated a preference for cold-water habitat. Feeding structure differentiation separated Blackfin from other coregonines, with Blackfin possessing the most numerous (50-66) gill rakers, and, via allometric regression, the longest gill rakers and lower gill arches. Selection for feeding efficiency may be a result of Mysis diluviana affecting planktonic size structure in lakes containing Blackfin Cisco, an effect also discovered in Lake Whitefish (Coregonus clupeaformis). This thesis provides insight into the habitat preferences and feeding ecology of Blackfin and provides a basis for future study.

Author Keywords: allometric regression, blackfin cisco, habitat, hurdle models, lake whitefish, mysis

Across vertebrate taxa, the hypothalamic-pituitary-adrenal axis (or the stress axis) is highly conserved, and is central to vertebrate survival because it allows appropriate responses to psychological stressors. Habitat shapes successful physiological and ecological strategies, and to appreciate how individual species respond to stressors in their environment, it is essential to have a thorough knowledge of the basic stress physiology of each species. In this dissertation, I studied the functioning and evolution of the stress physiology of New World flying squirrels. I showed that baseline, circulating cortisol levels in northern (Glaucomys sabrinus) and southern (G. volans) flying squirrels are some of the highest ever reported for mammals, indicating that their stress axes operate at a higher set point than most other species. I also assessed other aspects of their acute stress response, including free fatty acid and blood glucose levels, and indices of immune function, and showed that the flying squirrels' physiological reaction to stressors may differ from that of other mammals. Using immunoblotting, I found that corticosteroid-binding globulin (CBG) expression levels in flying squirrels appeared to be higher than previously reported using alternative methods. I also concluded however, that these levels did not appear to be high enough to provide their tissues with the protective CBG-bound buffer from their high circulating cortisol concentrations experienced by the majority of vertebrates. Thus, this arm of cortisol regulation within the flying squirrel stress axes may be weak or non-existent. Following this, I focused on southern flying squirrels and showed evidence that the second arm of cortisol regulation — the negative feedback mechanism at the level of the brain — functions effectively, but that this species is glucocorticoid resistant. Their tissue receptors appear to have a reduced affinity for cortisol, and this affinity may change seasonally to allow for the onset of other biological processes required for survival and reproduction. Due to their distinctive stress physiology, northern and southern flying squirrels may provide comparative physiologists with model systems for further probing of the function and evolution of the stress axis among vertebrates.

Author Keywords: corticosteroid-binding globulin, flying squirrel, Glaucomys, glucocorticoids, physiological ecology, stress physiology

This study examines foraging strategies during the Middle Woodland Period's Sandbanks Phase (A.D. 700–1000) on Boyd Island, Pigeon Lake, Ontario. The faunal remains analyzed in this study were recovered from a site associated with the procurement of aquatic and terrestrial taxa. Detailed taphonomic analyses have revealed that the Boyd Island faunal remains were affected by weathering and human transport decisions. White-tailed deer was the most frequently acquired prey at Boyd Island, followed by black bear. Using the central place forager prey choice model as a framework, the analysis of diet breadth and carcass transport patterns suggests that most animal resources were acquired from both aquatic and terrestrial habitats, at moderate distances from the site. Incomplete carcasses of large game appear to have been transported away from the site, where they were subsequently processed for provisioning or consumption. Comparisons with other Sandbanks faunal assemblages and those dating to later periods indicate significant differences in terms of taxonomic composition, while continuing to emphasize the use of fish. It is suggested that the Middle Woodland foragers adopted subsistence strategies focusing on the exploitation of local habitats in which productivity may have been enhanced through niche construction associated with the low-level food production activities.

Author Keywords: animal resource exploitation, archaeozoology, foraging theory, Middle Woodland, niche construction theory, southcentral Ontario

Population cycles are regular fluctuations in population densities, however, in recent years many cycles have begun to disappear. With Canada lynx this dampening has also been seen with decreasing latitude corresponding to an increase in prey diversity. My study investigates the role of alternate prey on the stability of the lynx-hare cycle by first comparing the functional responses of two sympatric but ecologically distinct predators on a primary and alternate prey. I then populated a three species predator-prey model to investigate the role of alternate prey on population stability. My results showed that alternate prey can promote stability, though they are unlikely to "stop the cycle". Furthermore, stability offered by alternate prey is contingent on its ability to increase intraspecific competition. My study highlights that population cycles are not governed by a single factor and that future research needs to be cognizant of interactions between alternate prey and intraspecific competition.

Author Keywords: alternate prey, Canis latrans, functional response, Lepus americanus, Lynx canadensis, Tamiasciurus hudsonicus