Graduate Theses & Dissertations


multi-faceted approach to evaluating the detection probability of an elusive snake (Sistrurus catenatus)
Many rare and elusive species have low detection probabilities, thereby imposing unique challenges to monitoring and conservation. Here, we assess the detection probability of the Eastern Massasauga (Sistrurus catenatus) in contrast to a more common and conspicuous species, the Eastern Gartersnake (Thamnophis sirtalis). We found that patterns of detection probability differed between species, wherein S. catenatus was detected less often and under a more specific set of sampling conditions. Correspondingly, detection trials with S. catenatus found a high non-detection rate, while detection trials with artificial models suggest that regional differences in detection probability are driven by variation in population density and habitat use. Our results suggest that current monitoring efforts are not sufficient, and that S. catenatus is frequently undetected. Accordingly, we highlight the importance of species-specific monitoring protocols when monitoring rare and elusive species, and recommend a multi-faceted approach that estimates detection probability and identifies species-specific challenges to monitoring. Author Keywords: detection probability, elusive species, monitoring programs, non-detection, S. catenatus, snakes
Making eDNA count
Environmental DNA (eDNA) is rapidly becoming an established method for the detection of species in aquatic systems and has been suggested as a promising tool to estimate species abundance. However, the strength of the relationship between eDNA concentrations and taxon abundance (density/biomass) can vary widely between species. I investigated the relationship between eDNA concentration and species abundance using two common and closely-related amphibians in eastern North America, the wood frog (Rana sylvatica) and northern leopard frog (Rana pipiens). I manipulated tadpole density in 80 L mesocosms and documented the relationship between tadpole density, biomass, and eDNA concentration. Species were comparable in biomass but differed in the amount of detectible genetic material produced; density and biomass were the superior abundance metric correlated with eDNA concentration for wood frogs and leopard frogs, respectively. However, increases in eDNA concentration reflected increasing tadpole biomass, therefore biomass is likely a better metric of abundance than density. Overall my findings support that eDNA concentration can be used as an index of species abundance, but that species-specific calibration may be needed before eDNA concentration can be effectively translated to an abundance metric. Future research should refine our understanding of how biotic and abiotic factors influence eDNA production, degradation, and recovery across species, before the method can receive widespread use as a monitoring tool in natural settings. Author Keywords: abundance estimates, environmental DNA, mesocosm, Rana pipiens, Rana sylvatica
How Abiotic and Biotic Factors Can Alter the Competitive Landscape in an Aggressive Species Complex (Genus
Competition is known to impact population dynamics through both indirect and direct interactions, and direct interactions can often lead to injury in one or both parties. As such, response to injury through tissue regeneration can be important for surviving post-competitive interaction. However, the impacts of outside factors like temperature and genome size (e.g. polyploidy) are not well studied, especially in syntopic systems. We addressed this knowledge gap by comparing regeneration rates of diploid Ambystoma laterale and triploid unisexual Ambystoma at two ecologically-relevant temperatures. Environmental factors appeared to have stronger effects on regeneration than ploidy level, but overall mass was impacted more strongly by ploidy level. Interestingly, there was an interaction between temperature and time within unisexuals that was absent when comparing different ploidy levels, implying temperature has a more complex effect on polyploids. This study supports the hypothesis that polyploid organisms are better equipped to respond to shifts in their environments, which can give them a competitive advantage at the northern range limit of this species complex. Author Keywords: Ambystoma, Genome dosage, Hybrid vigor, Polyploidy, Thermal optimum, Tissue regeneration
cascading effects of risk in the wild
Predation risk can elicit a range of responses in prey, but to date little is known about breadth of potential responses that may arise under realistic field conditions and how such responses are linked, leaving a fragmented picture of risk-related consequences on individuals. We increased predation risk in free-ranging snowshoe hares (Lepus americanus) during two consecutive summers by simulating natural chases using a model predator (i.e., domestic dog), and monitored hare stress physiology, energy expenditure, behaviour, condition, and habitat use. We show that higher levels of risk elicited marked changes in physiological stress metrics including sustained high levels of free plasma cortisol which had cascading effects on glucose, and immunology, but not condition. Risk-augmented hares also had lowered daily energy expenditure, spent more time foraging, and decreased rest, vigilance, and travel. It is possible that these alterations allowed risk-exposed hares to increase their condition at the same rate as controls. Additionally, risk-augmented hares selected, had high fidelity to, and were more mobile in structurally dense habitat (i.e., shrubs) which provided them additional cover from predators. They also used more open habitat (i.e., conifer) differently based on locale within the home range, using familiar conifer areas within cores for rest while moving through unfamiliar conifer areas in the periphery. Overall, these findings show that prey can have a multi-faceted, highly plastic response in the face of risk and can mitigate the effects of their stress physiology given the right environmental conditions. Author Keywords: behaviour, condition, daily energy expenditure, predator-prey interactions, snowshoe hare, stress physiology
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
Assessing Connectivity of Protected Area Networks and the Role of Private Lands in the United States
Forestalling biodiversity loss through the establishment of protected areas is a universally accepted conservation strategy, yet despite established guidelines for protected area coverage and placement, much of the world is currently failing to meet its commitments to conservation planning and landscape protection. Calls for the United States to protect more land usually focus on the need for strategic selection of land parcels to bolster protected area coverage and network functionality, but to date there lacks focused research on either the role of private protected areas in conservation planning or the factors affecting individual protected area selection and importance. We determined gaps in conservation planning in the contiguous United States by analyzing the connectivity of protected area networks by state, and assessing the importance of private protected areas in improving linkages in protected area connectivity. We found that all states had low coverage from protected areas (average <8.4% of total land mass), and especially private protected areas (average <1.1% of total land mass), and that the overall contribution of such areas to protected area network connectivity also was low. Terrain ruggedness was identified as the main factor affecting the current location of protected areas, and that protected area spatial layout is a primary influence on landscape connectivity. We conclude that establishment of private protected areas could offer a viable conservation tool for increasing protected area coverage and connectivity, but that current efforts are inadequate to either adequately link existing protected areas or to meet established land protection guidelines. Author Keywords: Aichi Target 11, conservation planning, graph theory, network theory, private conservation, protected areas
Enduring Attack
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
Complex niche determinants in terrestrial salamanders
I assessed how organisms having multiple biotic attributes may have conflicting niche determinants, and whether the realized niche reflects single or multiple attributes. All-female salamanders engage in two biotic states: hybridism and reproductive parasitism. Hybrids should occupy areas transitional to those used by parental species, whereas parasites that engage in competition with hosts should occupy habitats moderately suitable for hosts. Using niche models, I predicted realized niches for unisexual Ambystoma via a hybrid model (environmental predictors) and a parasite model (host suitability predictors). The hybrid model predicted that the unisexual niche would indeed be transitional between parental Ambystoma spp. The parasite model demonstrated unisexual salamanders occupied habitats moderately suitable for hosts, though model validation did not fully corroborate its predictive power. The hybrid model was more descriptive of unisexual occurrence than the parasite model. When species have competing ecological roles a primary biotic attribute may largely derive the realized niche. Author Keywords: Ambystoma, hybrid, niche, parasite, range, unisexual
Corticosterone Promotes Development of Cannibalistic Morphology and Inhibits Tissue Regeneration in Axolotls (Ambystoma mexicanum)
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
New Interpretations from Old Data
Range contractions and expansions are important ecological concepts for species management decisions. These decisions relate not only to rare and endangered species but to common and invasive species as well. The development of the broad spatiotemporal extent models that are helpful in examining range fluctuations can be challenging given the lack of data expansive enough to cover the time periods and geographic extents needed to fit the models. Archival records such as museum databases and harvest data can provide the spatiotemporal extent needed but present statistical challenges given they represent presence-only location information. In this thesis, I used maximum entropy and Bayesian hierarchical occupancy algorithms fitted with archival presence-only records to develop spatiotemporal models covering broad spatial and temporal extents for snowshoe hare and Canada lynx. These two algorithm types are well suited for presence-only data records and can be adapted to include biological and physical processes, thus improving the ecological realism of the models. Using these modelling methods, I found the extent of occurrence (EOO) and area of occupancy (AOO) varied greatly over time and space for both snowshoe hare and Canada lynx, suggesting that management decisions for these species should include consideration of these variations. While the presence-only data were appropriate for model development and understanding changing values in EOO and AOO, it sometimes lacked the locational accuracy and precision needed to create fine scale ecological analyses, thus resulting in somewhat coarse but potentially relevant conclusions. Author Keywords: Area of occupancy, Bayesian hierarchical models, Canada lynx, Extent of occurrence, Presence-only data, Snowshoe hare
Time to adapt
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
Stoichiometric food quality affects responses of Daphnia to predator-derived chemical cues
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


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