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
Calving site selection and fidelity in a restored elk (Cervus elaphus) herd in Bancroft, Ontario, Canada
ABSTRACT Calving site selection and fidelity in a restored elk (Cervus elaphus) herd in Bancroft Ontario, Canada. Michael R. Allan Parturition site selection by ungulates is believed to be influenced by forage abundance and concealment from predators. In 2011 and 2012, I used vaginal implant transmitters and movements to identify calving sites for 23 GPS collared elk (Cervus elaphus) from a restored herd. I tested the hypothesis that maternal elk used sites with higher forage and denser concealment compared to pre-calving sites at micro and macrohabitat levels. I detected no significant microhabitat differences from direct measurements of vegetation. At the macrohabitat scale, based on proximity of landcover classes, mean distances to hardwood forests was significantly less for calving (153 m) than pre-calving sites (198 m). Site fidelity is hypothesized to offer security in terms of familiarity to an area. I tested the hypothesis that females demonstrated fidelity to their previous year's location during pre-partum, parturition, post partum, breeding and winter periods. Elk were more philopatric during parturition and post partum than during breeding. Compared to winter elk were more philopatric during pre-partum, parturition and post-partum periods. Expressed as distance between consecutive-year calving locations, site fidelity varied with 27% of females exhibiting high (<1 km), 18% moderate and 55% (>2.9 km) low fidelity. I measured nearest-neighbour distances at calving time, exploring the hypothesis that females distance themselves from conspecifics. Elk increased the average distances to collared conspecifics during parturition; however, sample sizes were small. This strategy might influence calving site selection. Rapid movement prior to parturition, low site fidelity and spacing-out of females during parturition appear to be strategies to minimize predator risk and detection. Little evidence of selection for vegetation structure suggests this may not be limiting to these elk. Author Keywords: calving, elk, fidelity, movement, parturition, selection
Comparative Evaluation of Effective Population Size Genetic Estimation Methods in Wild Brook Trout (Salvelinus fontinalis) Populations
Effective population size (Ne) is a key concept in population genetics, evolutionary biology and conservation biology that describes an important facet of genetic diversity and the capacity of populations to respond to future evolutionary pressures. The importance of Ne in management and conservation of wild populations encouraged the development of numerous genetic estimators which rely on a variety of methods. Despite the number and diversity of available Ne methods, however, tests of estimator performance have largely relied on simulations, with relatively few tests based on empirical data. I used well-studied wild populations of brook trout (Salvelinus fontinalis) in Algonquin Park, Ontario as a model system to assess the comparative performance of multiple Ne estimation methods and programs, comparing the resultant Ne estimates against demographic population size estimates. As a first step, the genetic diversity and ancestry of wild brook trout populations was determined using 14 microsatellite loci. Genetic structure of brook trout populations showed variable contributions from historical supplemental stocking and also identified localized gene pools within and between watersheds, reflecting variable levels of connectivity and gene flow. Once the genetic ancestry and connectivity of populations had been resolved, single sample (point) and two samples (temporal) genetic estimators were used to estimate Ne of populations with pure native ancestry. Values obtained from genetic estimators utilizing both methods were variable within as well as among populations. Single sample (point) estimators were variable within individual populations, but substantially less than was observed among the temporal methods. The ratios of Ne to the estimated demographic population size (N) in small populations were substantially higher than in larger populations. Variation among estimates obtained from the different methods reflects varying assumptions that underlay the estimation algorithms. This research further investigated the effect of sampling effort and number of microsatellite loci used on Ne values obtained using the linkage disequilibrium (LD) estimation method. Ne estimates varied substantially among values generated from subsets of loci and genotyped individuals, highlighting the necessity for proper sampling design for efforts aiming to measure Ne. Despite the variation observed among and within estimation methods, the Ne concept is a valuable for the conservation and management of both exploited and endangered species. Author Keywords: Brook Trout, Effective population size, Genetic Diversity, Genetic Structure
Factors Controlling Peat Chemistry and Vegetation Composition in Sudbury Peatlands after 30 Years of Emission Reductions
Peatlands are prevalent in the Sudbury, Ontario region. Compared with the well documented devastation to the terrestrial and aquatic ecosystems in this region, relatively little work has been conducted on the peatlands. The objective of this research was to assess factors controlling peat and plant chemistry, and vegetation composition in 18 peatlands in Sudbury after over 30 years of emission reductions. Peatland chemistry and the degree of humification varies considerably, but sites closer to the main smelter had more humified peat and the surface horizons were enriched in copper (Cu) and nickel (Ni). Copper and Ni concentrations in peat were significantly correlated with Cu and Ni in the plant tissue of leatherleaf, although the increased foliar metal content did not obviously impact secondary chemistry stress indicators. The pH and mineral content of peat were the strongest determining factors for species richness, diversity and community composition. The bryophyte communities appear to be acid and metal tolerant, although Sphagnum mosses are showing limited recovery. Author Keywords: anthropogenic emissions, bryophytes, community comspoition, heavy metals, peatlands, wetland vegetation
Demography and habitat selection of Newfoundland caribou
The objective of this thesis is to better understand the demography and habitat selection of Newfoundland caribou. Chapter 1 provides a general introduction of elements of population ecology and behavioural ecology discussed in the thesis. In Chapter 2, I examine the causes of long-term fluctuations among caribou herds. My findings indicate that winter severity and density-dependent degradation of summer range quality offer partial explanations for the observed patterns of population change. In Chapter 3, I investigate the influence of climate, predation and density-dependence on cause-specific neonate survival. I found that when caribou populations are in a period of increase, predation from coyotes and bears is most strongly influenced by the abiotic conditions that precede calving. However, when populations begin to decline, weather conditions during calving also influenced survival. I build on this analysis in Chapter 4 by determining the influence of climate change on the interplay between predation risk and neonate survival. I found that the relative equilibrium between bears and coyotes may not persist in the future as risk from coyotes could increase due to climate change. In Chapter 5, I investigate the relationships in niche overlap between caribou and their predators and how this may influence differential predation risk by affecting encounter rates. For coyotes, seasonal changes in niche overlap mirrored variation in caribou calf risk, but had less association with the rate of encounter with calves. In contrast, changes in niche overlap during the calving season for black bears had little association with these parameters. In Chapter 6, I examine broad-level habitat selection of caribou to study trade-offs between predator avoidance and foraging during the calving season. The results suggest that caribou movements are oriented towards increased access to foraging and the reduction of encounter risk with bears, and to a lesser extent, coyotes. Finally, I synthesize the major findings from this thesis and their relevance to caribou conservation in Chapter 7, to infer that Newfoundland caribou decline is ultimately driven by extrinsic and intrinsic elements related to density-dependence. Reduction in neonate survival emerged from nutritionally-stressed caribou females producing calves with lower survival. Author Keywords: Behavioural ecology, Black bear (Ursus americanus), Coyote (Canis latrans), Population ecology, Predator-prey interactions, Woodland caribou (Rangifer tarandus)
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
Hybridization Dynamics between Wolves and Coyotes in Central Ontario
Eastern wolves (Canis lycaon) have hybridized extensively with coyotes (C. latrans) and gray wolves (C. lupus) and are listed as a `species of special concern' in Canada. Previous studies have not linked genetic analysis with field data to investigate the mechanisms underlying Canis hybridization. Accordingly, I studied genetics, morphology, mortality, and behavior of wolves, coyotes, and hybrids in and adjacent to Algonquin Provincial Park (APP), Ontario. I documented 3 genetically distinct Canis types within the APP region that also differed morphologically, corresponding to putative gray wolves, eastern wolves, and coyotes. I also documented a substantial number of hybrids (36%) that exhibited intermediate morphology relative to parental types. I found that individuals with greater wolf ancestry occupied areas of higher moose density and fewer roads. Next, I studied intrinsic and extrinsic factors influencing survival and cause-specific mortality of canids in the hybrid zone. I found that survival was poor and harvest mortality was high for eastern wolves in areas adjacent to APP compared with other sympatric Canis types outside of APP and eastern wolves within APP. Contrary to previous studies of wolves and coyotes elsewhere, I hypothesized that all Canis types exhibit a high degree of spatial segregation in the Ontario hybrid zone. My hypothesis was supported as home range overlap and shared space use between neighboring Canis packs of all ancestry classes were low. Territoriality among Canis may increase the likelihood of eastern wolves joining coyote and hybrid packs and exacerbate hybridization. Canids outside APP modified their use of roads between night and day strongly at high road densities (selecting roads more at night), whereas they responded weakly at lower road densities (generally no selection). Individuals that survived exhibited a highly significant relationship between the difference in their night and day selection of roads and availability of roads, whereas those that died showed a weaker, non-significant response. My results suggest that canids in the unprotected landscape outside APP must balance trade-offs between exploiting benefits associated with secondary roads while mitigating risk of human-caused mortality. Overall, my results suggest that the distinct eastern wolf population of APP is unlikely to expand numerically and/or geographically under current environmental conditions and management regulations. If expansion of the APP eastern wolf population (numerically and in terms of its geographic distribution) is a conservation priority for Canada and Ontario, additional harvest protection in areas outside of APP may be required. If additional harvest protection is enacted, a detailed study within the new areas of protection would be important to document specific effects on eastern wolf population growth. Author Keywords: Canis, coyotes, eastern wolves, hybridization, resource selection, survival
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
Dynamics and Mechanisms of Community Assembly in a Mined Carolinian Peatland
Theoretical work on community recovery, development, stability, and resistance to species invasions has outpaced experimental field research. There is also a need for better integration between ecological theory and the practice of ecological restoration. This thesis investigates the dynamics of community assembly following peat mining and subsequent restoration efforts at Canada's most southerly raised bog. It examines mechanisms underlying plant community changes and tests predictions arising from the Dynamic Environmental Filter Model (DEFM) and the Fluctuating Resource Hypothesis (FRH). Abiotic, biotic and dispersal filters were modified to test a conceptual model of assembly for Wainfleet Bog. Hydrology was manipulated at the plot scale across multiple nutrient gradients, and at the whole bog scale using peat dams. Trends in time series of hydrological variables were related to restoration actions and uncontrolled variables including precipitation, evapotranspiration and arrival of beaver. Impacts of a changing hydrology on the developing plant community were compared with those from cutting the invasive Betula pendula. Transplanting experiments were used to examine species interactions within primary and secondary successional communities. Seedlings of B. pendula and the native Betula papyrifera were planted together across a peat volumetric water content (VWC) gradient. Impacts of beaver dams were greater than those of peat dams and their relative importance was greatest during periods of drought. Cutting of B.pendula had little effect on the secondary successional plant community developing parallel to blocked drains. Phosphorus was the main limiting nutrient with optimum levels varying substantially between species. Primary colonisers formed a highly stable, novel plant community. Stability was due to direct and indirect facilitative interactions between all species. Reduction in frost heaving was the major mechanism behind this facilitation. Interactions within the secondary successional community were mostly competitive, driven by light and space availability. However, restricted dispersal rather than competition limited further species recruitment. Predictions based on the DEFM were partially correct. A splitting of this model's biotic filter into competition and facilitation components is proposed. There was little support for the FRH based on nutrient levels and VWC. B. pendula had higher germination and growth rates, tolerance to a wider range of peat VWCs and a greater resistance to deer browsing than native birch. Peat mining, combined with restoration actions and the arrival of beaver has moved much of the bog back to an earlier successional stage circa 350+ years BP. Evidence points to B. pendula being a "back-seat driver" in the ecosystem recovery process. Indirect facilitation of a native by an exotic congener, mediated through herbivory, has not been described previously. Shifts in relative contributions of facilitation, competition and dispersal limitations to community assembly may be useful process-oriented measures for gauging progress in restoration. Author Keywords: Betula pendula, community assembly, competition, facilitation, peatland, restoration
regional comparison of the structure and function of benthic macroinvertebrate communities within Precambrian Shield and St. Lawrence lowland lakes in south-central Ontario
Benthic macroinvertebrtes (BMI) are functionally important in aquatic ecosystems; as such, knowledge of their community structure and function is critical for understanding these systems. BMI were sampled from ten lakes in each of two regions of south-central Ontario to investigate which chemical and physical variables could be shaping their community structure and function. Ten Precambrian Shield lakes in the Muskoka-Haliburton region, and ten St Lawrence lowland lakes in the Kawartha lakes region were sampled. These lakes are geologically and chemically distinct, creating natural chemical and physical gradients within and between both regions. Community function was assessed using stable isotope analysis to elucidate carbon transfer dynamics (δ13C) and food web interactions (δ15N). It was predicted that the BMI from Shield lakes would have a δ13C signature indicative of allochthonous carbon subsidies, whereas the lowland lake BMI signatures would reflect autochthonous production. Additionally, it was predicted that the food web length (measured in δ15N units) would be different in Shield and lowland lakes. Both of these predictions were supported; however, the data indicate that δ13C signatures are more likely influenced by catchment geology (represented by bicarbonate concentration) than the extent of allochthony. The best predictor of food web length was found to be region. To assess BMI community structure, taxonomic richness, %EPT (% Ephemeroptera, Plecoptera, Trichoptera; a water quality index), and distribution of functional feeding groups were examined. Based on chemistry it was expected that the Shield lakes would be more speciose, and of greater water quality (relatively lower nutrient levels). These predictions were rejected; since there were no significant regional differences in taxonomic richness or biologically inferred water quality (%EPT). However, sediment size was found to best explain the variability in both metrics, with greater richness and %EPT found at sites with medium and small substrates than those with large substrates. Significant regional differences were found in the distribution of functional feeding groups. Most notably, there were significantly greater proportions of scrapers and shredders in the lowland and Shield lakes, respectively. Based on the feeding mechanisms of these invertebrates it can be inferred that allochthonous subsidies are likely of greater importance to Shield lake BMI communities than those of the lowland lakes; supporting the carbon transfer prediction. These findings provide insight about the structure and function of BMI communities from two dominant lake types in Ontario, and could be useful when determining how future chemical and physical changes will impact these communities. Author Keywords: benthic macroinvertebrates, community function, community structure, Precambrian Shield, stable isotopes, St. Lawrence lowlands
EVALUATION OF HAYFIELD MANAGEMENT STRATEGIES AND BOBOLINK TERRITORIAL HABITAT IN SOUTHERN ONTARIO
I implemented three hayfield management regimens in southern Ontario (a typical schedule at the farmer`s discretion, a delayed first harvest after July 14, and an early first harvest before June 1 with 65 days before second harvest), and evaluated the costs/benefits to farmers regarding hay quality and feasibility, and to Bobolinks (Dolichonyx oryzivorus) regarding reproductive activity and phenology. Typical management resulted in little to no Bobolink reproductive success, and early harvested sites were not (re)colonized. On delayed harvest sites Bobolinks experienced high reproductive success, but hay quality fell below ideal protein levels for most cattle before harvest. I also examined the habitat features Bobolinks use as the basis for establishing territories and associations between Bobolink territory size and habitat quality. I compared vegetation structure, patch size, and prey abundance between small and large territories. Small territories typically occurred on smaller fields with more preferred vegetation characteristics and greater prey abundance. Author Keywords: agro-ecosystem, Bobolink, Dolichonyx oryzivorus, grassland birds, hayfield management
Widespread changes in growth, diet and depth distribution of lake whitefish (Coregonus clupeaformis) in the Great Lakes are linked to invasive dreissenid mussels
Recent declines in growth and condition of Great Lakes' lake whitefish (Coregonus clupeaformis) have been linked to ecosystem-wide changes stemming from the invasion of dreissenid mussels. To test the influence of invasive mussels on this commercially important coregonid species, we collected archived scale samples from ten Great Lake locations and analyzed long-term changes in growth rates, delta 13C and delta 15N stable isotope ratios before and after mussel establishment. There was a decrease in pre-maturation growth after establishment in all four locations where we examined back-calculated growths. In six of the seven locations with dreissenid populations, a significant increase in delta 13C and a significant decrease in delta 15N was found. In dreissenid-absent locations of Lake Superior, we did not see changes in growth or isotope ratios indicative of a major regime shift. Observed shifts in isotopic signatures provide evidence for an increased reliance on nearshore food sources and shallower depth distribution as a result of dreissenids, which likely contributed to lowered growth of lake whitefish. Author Keywords: Diporeia, Dreissenids, food web, Great Lakes, invasive species, lake whitefish

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