Murray, Dennis L

While both resource quality and predator-derived chemical cues can each have profound effects on zooplankton populations and their function in ecosystems, the strength and nature of their interactive effects remain unclear. We conducted laboratory experiments to evaluate how stoichiometric food quality (i.e., algal carbon (C):phosphorus (P) ratios) affects responses of the water flea, Daphnia pulicaria, to predator-derived chemical cues. We compared growth rates, body elemental content, metabolic rates, life history shifts, and survival of differentially P-nourished Daphnia in the presence and absence of chemical cues derived from fish predators. We found effects of predator cues and/or stoichiometric food quality on all measured traits of Daphnia. Exposure to fish cues led to reduced growth and increased metabolic rates, but had little effect on the elemental content of Daphnia. Elevated algal C:P ratios reduced growth and body %P, increased respiration, and increased body %C. Most of the effects of predator cues and algal C:P ratios of Daphnia were non-interactive. In contrast, the declines in daphnid survival and related population growth rates that arose because of poor food quality were amplified in the presence of predator-derived cues. Our results demonstrate that stoichiometric food quality interacts with anti-predator responses of Daphnia, but these effects are trait-dependent and appear connected to animal life-history evolution.

Author Keywords: Daphnia, ecological stoichiometry, indirect predator effects, life history, phosphorus, predator-prey relationships

Salamanders are capable of tissue regeneration throughout all life-stages, which requires the dedifferentiation of mature cells to regrow lost tissues. Dedifferentiation is promoted by degradation of the extracellular matrix by matrix metalloproteases, as well as lysosomal degradation of intracellular and cell-surface proteins that mark cells as part of a mature lineage. Salamanders are also capable of developing cannibalistic phenotypes, plastic traits that are elicited by environmental stressors that result in elevated circulating glucocorticoid (e.g., corticosterone) levels that underlie many fundamental adaptive changes in morphology. Interestingly, the direct effect of corticosterone on regeneration and the cannibalistic phenotype have yet to be examined. In the present thesis, axolotls (Ambystoma mexicanum) were exposed to exogenous corticosterone and 50% of the distal tail tissue was removed. The effects of high corticosterone levels on matrix metalloprotease (MMP-2, MMP-9) and lysosomal acid phosphatase (LAP) activity were assessed; these are two classes of enzymes which are markers of extracellular matrix and intracellular remodeling during regeneration, respectively. We found that elevated corticosterone levels inhibited tissue regeneration, by prolonging the dedifferentiation phase as indicated by increased LAP and reduced MMP-2 and MMP-9 activity. Elevated corticosterone levels also promoted the cannibalistic morphology and this effect was strongest among smaller individuals.

Author Keywords: amphibian, cannibalistic morphology, corticosterone, dedifferentiation, regeneration, stress

Numerous prey taxa employ defensive postures for protection against attack by predators. Defensive postures mitigate predation risk at various stages of the predator-prey sequence, including through crypsis, mimicry, thanatosis, aposematism, and deflection. In terrestrial salamanders, defensive postures may be aposematic, or deflect attacks away from vital body parts and towards the tail, however the extent to which these strategies act exclusively or synergistically remains poorly understood. Herein I demonstrate a novel approach to study the function of salamander defensive postures through experimental manipulation of predator response to antipredator behaviour in a natural field setting. I deployed 1600 clay salamander prey on Pelee Island, Ontario, manipulating prey size (small, large) and posture (resting, defensive) and documented attack rates across three predator types to further assess the effect of prey body size and predator type on antipredator efficacy. My research suggests that irrespective of prey body size, defensive posture does not function through aposematism, but rather acts to deflect predator attacks to the tail, which is commonly noxious and expendable in terrestrial salamanders. An intriguing possibility is that this behaviour facilitates taste-rejection by predators. Overall, my research should further contribute to our understanding of the importance and potential evolutionary significance of defensive posturing in Ambystoma salamanders, and more broadly, on the determinants of prey vulnerability to predation. I also briefly discuss the implications of my results to the conservation of Ambystoma populations on Pelee Island.

Author Keywords: Anti-predator behaviour, Aposematism, Attack deflection, Predator avoidance, Small-mouthed salamander, Taste-rejection

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

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

My objective was to understand how individual variation, in conjunction with variation in habitat, can affect individual and population-level variation in animal space use. I used coyotes (Canis latrans) as a model species to investigate the roles of hybridization, an inherited intrinsic factor, and spatial memory, a learned intrinsic factor, on space use. I used a diversity of methods and approaches, including meta-regression, multiple imputation, simulations, resource selection functions, step selection functions, net-squared displacement analysis, and survival analysis. A major contribution was my investigation of the performance of multiple imputation in a meta-regression framework in Chapter 2. My simulations indicated that multiple imputation performs well in estimating missing data within a meta-regression framework in most situations. In Chapter 3, I used published studies of coyote home range size in a meta-regression analysis with multiple imputation to examine the relative roles of hybridization and environmental variables on coyote home range size across North America. I found that hybridization with Canis species was a leading factor driving variation in coyote space use at a continental scale. In Chapter 4, I used telemetry data for 62 coyotes in Newfoundland, Canada, to investigate the influence of cognitive maps on resource use. I found that resident coyotes used spatial memory of the landscape to select or avoid resources at spatial scales beyond their immediate sensory perception relative to transient coyotes, presumably increasing their fitness. Taken together, my dissertation demonstrates that intrinsic factors, such as genetic ancestry and spatial memory, can have substantial influences on how animals use space at both individual and population levels, and at both a local and a continental scales.

Author Keywords: canis latrans, hybridization, meta-regression, multiple imputation, Newfoundland, spatial memory

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

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

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