Graduate Theses & Dissertations


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
mechanistic analysis of density dependence in algal population dynamics
Population density regulation is a fundamental principle in ecology, however there remain several unknowns regarding the functional expression of density dependence. One prominent view is that the patterns by which density dependence is expressed are largely fixed across a species, irrespective of environmental conditions. Our study investigated the expression of density dependence in Chlamydomonas reinhartti grown under a gradient of nutrient densities, and hypothesized that the relationship between per capita growth rate (pgr) and population density would vary from concave-up to concave-down as nutrients became less limiting. Contrary to prediction, we found that the relationship between a population's pgr and density became increasingly concave-up as nutrient levels increased. Our results suggest that density dependence is strongly variable depending on exogenous and endogenous processes acting on the population, implying that expression of density regulation depends extensively on local conditions. Population growth suppression may be attributable to environments with high intraspecific competition. Additional work should reveal the mechanisms influencing how the expression of density dependence varies across populations through space and time. Author Keywords: Chlamydomonas reinhartti, density dependence, logistic model, population dynamics, single species growth, theta-logistic equation
Evaluating the Effects of Habitat Loss and Fragmentation on Canada Lynx
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
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
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
Investigating Ecological Niche Differentiation Among Wild Candids Experiencing Hybridization in Eastern North America
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
Phylogeography and Genetic Structuring of Moose (Alces alces) Populations in Ontario, Canada
Moose are an iconic species, known for their large size and impressive antlers. Eight subspecies are classified in circumpolar regions of the planet - four in North America. Two subspecies are similar in shape and size, the north-western moose (Alces alces andersoni) and the eastern moose (Alces alces americana). It was previously believed that these two subspecies meet in northern Ontario. Earlier genetic population studies used a small number of samples from Ontario, primarily in broad studies covering all of North America. A comprehensive genetic study of moose populations in Ontario has not previously been conducted. We examined the genetic diversity and population structure at 10 polymorphic loci using 776 samples from Ontario, as well as outgroups from representative populations – Manitoba/Cape Breton, representing A. a. andersoni, and New Brunswick/Nova Scotia, representing A. a. americana. Results indicated three genetic populations in the province, in north-western Ontario, north-eastern Ontario and south-central Ontario. RST values, compared against both FST and Jost’s D values for phylogenetic analyses, indicated no phylogenetic pattern which suggests no subspeciation present in the province. Population movement patterns in Ontario were studied. Gene flow was estimated using genetic and spatial data. Isolation by distance was only seen within the first distance class of 100 kilometres and then not seen again at further distances, indicating that moose display philopatry. There were very few migrants travelling across the province, with a greater number moving gradually north and west, towards better habitat and food sources. A forensic database in the form of an allele frequency table was created. Three loci showed very low levels of heterozygosity across all three populations. Probability of identity was calculated for the three populations and quantified. Samples with known geographic origins were run against the database to test for sensitivity, with identification of origin occurring at an accuracy level between 87 and 100%. Within Ontario, there are not two different subspecies, as previously believed, but two different populations of the same subspecies meeting in northern Ontario. The genetic data does not support previous research performed in Ontario. The sample sizes in our research also provide a more comprehensive view of the entire province not seen in any previous studies. The comprehensive research enabled the building of a reliable forensic database that can be used for both management and forensic purposes for the entire province. Author Keywords: Alces alces, Genetic Diversity, Moose, Ontario, Phylogeography, Subspecies
Elemental Variation in Daphnia
Environmental variation can affect consumer trait expression and alter ecological and evolutionary dynamics in natural populations. However, although dietary nutrient content can vary by an order of magnitude in natural ecosystems, intra-specific differences in consumer responses to food quality have not been thoroughly investigated. Therefore, the purpose of my dissertation was to examine the influence of dietary nutrition and other environmental factors on consumer phenotypic variation using the freshwater cladoceran Daphnia. I conducted a series of complementary laboratory and field studies where I examined the effects of dietary phosphorus (P) content and additional biological/environmental variables (multi-elemental limitation, genetic variation, and temperature) on daphnid life-history, biochemistry, body elemental composition, and population growth. In general, phenotypic expression within a species varied significantly in response to all experimental variables, but the relative influence of each was highly context dependent. In my first chapter, I found that dietary P content and environmental calcium (Ca) concentrations both altered Daphnia body Ca:P ratios and growth rates of individuals and affected intrinsic rates of increase at the population level. However, food quality appeared to have a much larger effect on trait expression, and body Ca:P ratios were highly sensitive to other forms of dietary nutrient limitation. Next, I documented significant quantitative genetic variation and phenotypic plasticity in daphnid P content, growth, and P use efficiency of field collected animals grown across dietary P gradients. Trait expression was also influenced by genotype X diet interactions suggesting that consumer responses to dietary nutrient limitation can be heritable and may be adaptive in different nutrient environments. Finally, I found that temperature appeared to override food quality effects and decouple P metabolism in natural Daphnia populations, but total biomass production was affected by both dietary P content and temperature, depending on the nutrient content of the lake. Overall, my dissertation shows that consumer responses to nutrient limitation can vary significantly within a species and that changes in trait expression may be modified by other environmental variables. These results should be incorporated into existing stoichiometric models and used to investigate the eco-evolutionary consequences of consumer phenotypic variation in response to nutritional stress. Author Keywords: ecological stoichiometry, evolution, life-history, nutrient limitation, nutrient metabolism, zooplankton
third wheel
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
Time-dependent effects of predation risk on stressor reactivity and growth in developing larval anurans (Lithobates pipiens)
The predator vs. prey dynamic is an omnipresent factor in ecological systems that may drive changes in life history patterns in prey animals through behavioural, morphological, and physiological changes. Predation risk can have profound effects on the life history events of an animal, and is influenced by the neuroendocrine stress response. Activation of the hypothalamic-pituitary-adrenal/interrenal axis, and the induction of stress hormones (e.g., corticosterone (CORT)) have been shown to mediate the onset of inducible anti-predator defensive traits including increased tail-depth, and reduced activity. The predator-prey relationship between dragonfly nymphs and tadpoles can be a powerful model system for understanding mechanisms that facilitate changes in the stress response in accordance with altered severity of risk. It has been well demonstrated early in tadpole ontogeny that increased corticosterone (CORT) levels, observed within three weeks of predator exposure, are correlated with increased tail depth morphology. However, the reactivity of the stress response in relation to the growth modulation in developing prey has yet to be fully explored. Accordingly, this thesis assessed the stress and growth response processes in tadpoles that were continuously exposed to perceived predation risk later in ontogeny. Continuous exposure of prey to predation risk for three weeks significantly increased CORT levels, and tail depth. However, tadpoles exposed to six weeks of predation risk acclimated to the presence of the predator, which was observed as a significant reduction of stressor-induced CORT levels. In addition, although increased tail depth has been attributed to predator defense, predator-naïve tadpoles began to display similar tail depth morphology as treated tadpoles at the six week time point. Thus, this thesis suggests that the stress response in lower vertebrate systems (e.g., tadpoles) may operate in a similarly complex manner to that observed in higher vertebrates (e.g., rats), for which severity of risk associated with the stressor aids in defining activity of the stress response. Moreover, the lack of morphological difference between treatments among tadpoles exposed later in ontogeny suggests that the mechanisms for inducing defenses are normal morphological traits in the development of the animal. This thesis paves the way for future research to elucidate the relationship between the neuroendocrine stress response and hormonal pathways involved in growth modulation in the presence of environmental pressures. Author Keywords: Acclimation, Corticosterone, Growth Modulation, Predation Risk, R. pipiens, Tadpole
origin and ecological function of an ion inducing anti-predator behaviour in Lithobates tadpoles
Chemical cues are used commonly by prey to identify predation risk in aquatic environments. Previous work has indicated that negatively-charged ions of m/z 501 are possibly a kairomone that induces anti-predator responses in tadpoles. This thesis found that this ion species: (i) is produced by injured tadpoles; (ii) exhibits increased spectral intensity with higher tadpole biomass; and (iii) is not produced by starved predators. These results refute the hypothesis that the ion is a kairomone, and rather support its role as an alarm cue released from tadpoles. High resolution mass spectrometry (HR-MS) revealed a unique elemental composition for [M-H]-, m/z 501.2886, of C26H45O7S-. Collision induced dissociation (CID) of ion m/z 501 formed product ions of m/z 97 and m/z 80, HSO4- and SO3-, respectively, indicating the presence of sulfate. Green frog (Lithobates clamitans) tadpoles exposed to m/z 501, and an industrial analogue, sodium dodecyl sulphate (NaC12H25O4S), exhibited similar anti-predator responses, thereby suggesting the potential role of organic sulfate as a tadpole behavioural alterant. Author Keywords: Alarm cue, Amphibian, Chemical Ecology, Mass spectrometry, Predator-prey interactions
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


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