Graduate Theses & Dissertations

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Spatial Dynamics of Wind Pollination in Broadleaf Cattail (Typha latifolia)
Natural populations of flowering plants rarely have perfectly uniform distributions, so trends in pollen dispersal should affect the size of the pollination neighbourhood and influence mating opportunities. Here I used spatial analysis to determine the size of the pollination neighbourhood in a stand of the herbaceous, wind-pollinated plant (Typha latifolia; broad-leaved cattail) by evaluating patterns of pollen production and seed set by individual cattail shoots. I found a positive correlation between pollen production and seed set among near-neighbour shoots (i.e., within 4 m2 patches of the stand; Pearson's r = 0.235, p < 0.05, df = 77) that was not driven by a correlation between these variables within inflorescences (Pearson's r = 0.052, p > 0.45, df = 203). I also detected significant spatial autocorrelations in seed set over short distances (up to ~ 5 m) and a significant cross-correlation between pollen production and seed set over distances of < 1 m indicating that the majority of pollination events involve short distances. Patterns of pollen availability were simulated to explore the shape of the pollen dispersal curve. Simulated pollen availability fit actual patterns of seed set only under assumptions of highly restricted pollen dispersal. Together, these findings indicate that even though Typha latifolia produces copious amounts of pollen, the vast majority of pollen dispersal was highly localized to distances of ~ 1 m. Moreover, although Typha latifolia is self-compatible and has been described as largely selfing, my results are more consistent with the importance of pollen transfer between nearby inflorescences. Therefore, realized selfing rates of Typha latifolia should largely depend on the clonal structure of populations. Author Keywords: clonal structure, correlogram, dispersal curves, pollination, spatial analysis, Typha latifolia
Constraints on phenotypic plasticity in response to predation risk
Inducible defenses are plastic responses by an organism to the perception of predation risk. This dissertation focuses on three experiments designed to test the hypothesis that plastic ability is limited by energetic constraints. Chapter 1 provides a general introduction to phenotypic plasticity research and the theoretical costs and limitations affecting the expression of plastic traits. In Chapter 2, I tested the hypothesis that costs of early plasticity may be manifested by a reduced response to risk in later life stages. I found that amphibian embryos are able to detect and respond to larval predators, but that the energetic cost of those plastic responses are not equivalent among behavioural, growth, and morphological traits, and their expression differs between closely-related species. Chapter 3 explicitly examines the relationship between food resource availability and plasticity in response to perceived predation risk during larval development. Food-restricted tadpoles showed limited responses to predation risk; larvae at food saturation altered behaviour, development, and growth in response to predation risk. Responses to risk varied through time, suggesting ontogeny may affect the deployment of particular defensive traits. Chapter 4 examines the influence of maternal investment into propagule size on the magnitude of the plastic responses to predation risk in resulting offspring. I found that females in better body condition laid larger eggs and that these eggs, in turn, hatched into larvae that showed greater morphological plasticity in response to predation risk. Maternal investment can therefore affect the ability of offspring to mount morphological defenses to predation risk. Last, Chapter 5 provides a synthesis of my research findings, identifying specific factors constraining the plastic responses of prey to perceived predation risk. Overall, I found constraints on plastic responses imposed by the current environment experienced by the organism (resource availability), the prior experience of the organism (predator cues in the embryonic environment), and even the condition of the previous generation (maternal body condition and reproductive investment). Together, these findings both provide new knowledge and create novel research questions regarding constraints limiting phenotypic variation in natural populations. Author Keywords: behaviour, inducible defense, Lithobates pipiens, morphometrics, phenotypic plasticity, predation risk
Ice age fish in a warming world
In the face of climate change, the persistence of cold-adapted species and populations will depend on their capacity for evolutionary adaptation of physiological traits. As a cold-adapted Ice Age relict species, lake trout (Salvelinus namaycush) are ideal for studying potential effects of climate change on coldwater fishes. I studied the thermal acclimation capacity and aerobic metabolism of age 2+ lake trout from four populations across four acclimation temperatures (8ºC, 11ºC, 15ºC, and 19ºC). One population had marginally significant higher active metabolic rate (AMR) and metabolic scope across all temperatures. There was no interpopulation variation for critical thermal maximum (CTM), standard metabolic rate (SMR), or thermal acclimation capacity. Acclimation resulted in a 3ºC increase in thermal tolerance and 3-fold increase in SMR for all populations. At 19ºC, SMR increased and AMR declined, resulting in sharply reduced metabolic scope for all populations. The limited intraspecific variation in thermal physiology suggests that climate change may threaten lake trout at the species rather than population level. Author Keywords: Climate Change, Lake Trout, Metabolic Rate, Salvelinus namaycush, Temperature, Thermal Acclimation
USE OF SALIVARY CORTISOL TO EVALUATE THE INFLUENCE OF RIDES ON THE STRESS PHYSIOLOGY OF DROMEDARY CAMELS (CAMELUS DROMEDARIUS)
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
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
Home range use, habitat selection, and stress physiology of eastern whip-poor-wills (Antrostomus vociferus) at the northern edge of their range
The distribution of animals is rarely random and is affected by various environmental factors. We examined space-use patterns, habitat selection and stress responses of whip-poor-wills to mining exploration activity.To the best of my knowledge, fine scale patterns such as the habitat composition within known home ranges or territories of eastern whip-poor-wills have not been investigated. Using a population at the northern edge of the distribution in an area surrounding a mining exploration site, we tested whether variations in habitat and anthropogenic disturbances influence the stress physiology of individuals. We found no effect of increased mining activity on the stress physiology of birds but found a significant scale-dependent effect of habitat on their baseline and stress-induced corticosterone levels, and we suggest that these are the result of variations in habitat quality. The importance of other factors associated with those habitat differences (e.g., insect availability, predator abundance, and microhabitat features) warrants further research. Author Keywords: anthropogenic disturbances, Antrostomus vociferus, corticosterone, eastern whip-poor-will, habitat selection, radio-­telemetry
Social thermoregulation and potential for heterothermy
Northern and southern flying squirrels (Glaucomys sabrinus and G. volans, respectively) are experiencing a climate change induced increase in range overlap, resulting in recent hybridization. We investigated the occurrence of heterospecific communal nesting, a potential facilitator of hybridization, and aimed to confirm the presence of torpor, a potential barrier to hybridization, in flying squirrels. In wild-caught captive squirrels, we conducted a paired nest choice experiment and found that heterospecific nesting did occur, but in a lower frequency than conspecific nesting. Ambient temperature did not affect the frequency of grouped nesting. We attempted to induce torpor in flying squirrels in a laboratory through cold exposure while measuring metabolic rate and body temperature. Strong evidence of torpor was not observed, and metabolic rate remained unchanged with season. We conclude that torpor is not a barrier to hybridization in flying squirrels, but resistance to heterospecific nesting may indicate the existence of one. Author Keywords: heterospecific group, hybridization, northern flying squirrel, social thermoregulation, southern flying squirrel, torpor
effects of parasitism on consumer-driven nutrient recycling
Daphnia are keystone consumers in many pelagic ecosystems because of their central role in nutrient cycling. Daphnia are also frequently infected, and the parasites causing these infections may rival their hosts in their ability to regulate ecosystem processes. Therefore, parasitic exploitation of Daphnia may alter nutrient cycling in pelagic systems. This thesis integrates existing knowledge regarding the exploitation of Daphnia magna by 2 endoparasites to predict parasite-induced changes in the nutrient cycling of infected hosts and ecosystems. In chapter 1, I I contextualizing the integration of these themes by reviewing the development of the fields of elemental stoichiometry and parasitology. In chapter 2, we show how the bacterial parasite, Pasteuria ramosa, increased the nitrogen (N) and phosphorus (P) release rates of D. magna fed P-poor diets. We used a mass-balance nutrient release model to show that parasite-induced changes in host nutrient accumulation rates and diet-specific changes in host ingestion rates were responsible for the accelerated nutrient release rates that we observed. In chapter 3, we extended our examination of the nutrient mass balance of infected D. magna to include another parasite, the microsporidian H. tvaerminnensis. We found differences in the effects of these two parasites on host nutrient use as well as support for the hypothesis that parasite-induced changes in Daphnia N release are caused by the effects of infection on Daphnia fecundity. In chapter 4, we examined the relationship between P concentrations and the presence and prevalence of H. tvaerminnensis in rock pools along the Baltic Sea. We found that particulate P concentrations were negatively associated with the prevalence of this parasite, a result that is consistent with the increase in P sequestration of H. tvaerminnensis-infected Daphnia that we observed in chapter 3. I discuss the potential implications of the work presented in chapters 2-4 for other parasite-host systems and ecosystems in chapter 5. Overall, the research presented here suggests that parasite-induced changes in host nutrient use may affect the availability of nutrients in the surrounding environment, and the magnitude of this effect may be linked to parasite-induced reductions in fecundity for many invertebrate hosts. Author Keywords: consumer, ingestion rates, mass-balance, nutrient-recycling, parasitism, phosphorus
Comparing Biological Responses to Contaminants in Darters (Etheostoma spp.) Collected from Rural and Urban Regions of the Grand River Watershed, Ontario
Urban and agricultural activities may introduce chemical stressors, including contaminants of emerging concern (CECs) and current use pesticides (CUPs) into riverine systems. The objective of this study was to determine if fish collected from sites in a river show biomarkers of exposure to these classes of contaminants, and if the biomarker patterns vary in fish collected from urbanized and agricultural sites. The watershed selected for this study was the Grand River in southern Ontario, which transitions from areas dominated by agricultural land use in the north to highly urbanized locations in the southern part of the watershed. Rainbow darters (Etheostoma caerluem) and fantail darters (Etheostoma flabellare) were collected from the Grand River in June, 2014 for biomarker analysis from two urbanized sites and three agricultural sites (n=20 per site). Over the same period of time, Polar Organic Chemical Integrative Samplers (POCIS) were deployed for 2 weeks at each site to monitor for the presence of CUPs and CECs. The amounts of the target compounds accumulated on POCIS, determined using LC-MS/MS were used to estimate the time weighted average concentrations of the contaminants at each site. Data on the liver somatic index for darters indicate site-specific differences in this condition factor (p<0.05). Significant differences in the concentrations of thiobarbituric acid reactive substances (TBARS) in gill tissue (p<0.05) indicate differences in oxidative stress in fish collected from the various sites. Measured concentrations of ethoxyresorufin-O-deethylase (EROD) in liver tissue were significantly different between sites (p<0.05), indicating differences in CYP1A metabolic activity. Finally, acetylcholinesterase (AChE) activity in brain tissue was significantly different between fish from rural and urban sites (p<0.05). The analysis of these biomarkers indicates that fish may be experiencing different levels of biological stress related to different land uses. These data may be useful in developing mitigation strategies to reduce impacts on fish and other aquatic organisms in the watershed. Author Keywords: AChE, Biomarker, Darter, EROD, POCIS, TBARS
Pathogen vs. Predator
Stressors are often an inescapable part of an organism’s life. While the effects of many stressors have been well studied individually, potential interactions between stressors exist that may result in greater than additive negative effects. Stressors may be linked by conflicting demands on energy budgets, interfering with important physiological pathways, or necessitating incompatible adaptive responses. Using Ranavirus (FV3) and larval dragonfly predators (Anax spp.) in a 2x2 factorial experiment on green frog (Lithobates clamitans) tadpoles, I investigate the interactions in behaviour, morphology, and metabolism when both stressors were applied in concert. I demonstrate that activity and feeding are reduced additively by both stressors, and tadpoles increase distance between conspecifics in FV3-exposed tanks, but only in the absence of predators. I also note decreases in mass, and a non-significant marginal increase in metabolic rate of tadpoles exposed to FV3. Interestingly, I provide evidence that FV3 can compromise morphometric responses through antagonistic interactions with perceived predation risk exposure, which may result in significantly elevated mortality even when either stressor is present in sub-lethal quantities. Thus, I conclude that sub-lethal exposure to stressors can nonetheless have substantial impacts on organisms and a more integrative approach to examining the impacts of stressors on individual physiology and fitness is necessary. Author Keywords: Behaviour, Interaction, Morphology, Predation Risk, Ranavirus, Tadpoles
Bank Swallow (Riparia riparia) Breeding in Aggregate Pits and Natural Habitats
I examined Bank Swallow (Riparia riparia) colony persistence and occupancy, in lakeshore, river and man-made aggregate pit habitat. Habitat persistence was highest on the lakeshore and lowest in aggregate pits, likely due to annual removal and relocation of aggregate resources. Bank Swallow colonies in aggregate pit sites were more likely to persist if a colony was larger or if burrows were located higher on the nesting face. I also compared nest productivity and health factors of Bank Swallows in lakeshore and aggregate pit habitats. While clutch size was the same in both habitat types, the number of fledglings from successfully hatched nests was significantly higher in aggregate pit sites than from lakeshore sites. Mass of fledgling Bank Swallows did not differ significantly between habitat types, however mass of adults from aggregate pits decreased significantly over the nesting season. Parasite loads on fledgling Bank Swallows were significantly lower in aggregate pits than in lakeshore sites. According to these indicators, aggregate pits appear to provide equivalent or higher quality habitat for Bank Swallows than the natural lakeshore sites, making them adequate and potentially key for this species’ recovery. Aggregate pit operators can manage for swallows by (1) creating longer, taller faces to attract birds and decrease predation, and (2) supplementing their habitat with water sources to encourage food availability. Author Keywords: Aerial insectivore, aggregate pits, Bank Swallow, colony persistence, ectoparasites, substitute habitat
Stress Axis Function and Regulation in New World Flying Squirrels
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

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