Ecology

Caribou experience direct and indirect negative effects of harassment from biting flies, influencing behavior and activity on several spatial and temporal scales. I used systematic insect collection surveys during the summers of 2011 and 2012 to examine the spatial and temporal distributions of black flies (Simuliidae), mosquitoes (Culicidae), and deer flies and horse flies (Tabanidae) in a managed boreal forest in northern Ontario. Mosquitoes had a positive association with densely treed habitats, whereas black flies more often occurred in open areas, and tabanids had a strong presence in all habitat types. Habitats in proximity to large bodies of water had fewer biting flies than inland areas. Young stands supported higher abundances of tabanids despite vegetation community type. Next, I tested for seasonal effects of biting fly abundance on caribou activity by modelling the seasonal trend in abundance for each fly family for each year and compared this to an index of daily activity for 20 radio-collared female caribou in 2011 and 10 females in 2012. I modeled this index of caribou activity for each animal in each year and extracted the set of partial correlation coefficients from multiple regressions to test for effects of biting fly abundances on caribou activity. Caribou reduced their daily activity when tabanids were more numerous, and increased activity when mosquitoes were numerous. This divergent response may reflect a difference in the efficacy of moving to reduce harassment, owing to the stronger flight capabilities of tabanids.

Author Keywords: Activity, Anthropogenic Disturbance, Behaviour, Insect harassment, Temporal distribution, Woodland Caribou

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

The structure and composition of the landscape can facilitate or impede gene flow, which can have important consequences because genetically isolated groups of individuals may be prone to inbreeding depression and possible extinction. My dissertation examines how landscape structure influences spatial patterns of genetic differentiation and diversity of American marten (Martes americana) and Canada lynx (Lynx canadensis) in Ontario, Canada, and provides methodological advances useful for landscape geneticists. First, I identified the effects of map boundaries on estimates of landscape resistance, and proposed a solution to the bias: a buffer around the map boundary. Second, I assessed the sensitivity of a network-based estimate of genetic distance, conditional genetic distance, to incomplete sampling. I then used these landscape genetic tools in a pairwise, distance-based analysis of 653 martens genotyped at 12 microsatellite loci. I evaluated whether forest management in Ontario has influenced the genetic structure of martens. Although forest management practices had some impact, isolation by distance best described marten gene flow. Our results suggest that managed forests in Ontario are well connected for marten and do not impede marten gene flow. Finally, I used a site-based analysis of 702 lynx genotyped at 14 microsatellite loci to investigate spatial patterns of genetic diversity and differentiation at the trailing (contracting) edge of the lynx distribution in Ontario. I analyzed harvest records and found that the southern edge of lynx range has contracted by >175 km since the 1970s. I also found that neutral genetic diversity decreased towards the trailing edge, whereas genetic differentiation increased. Furthermore, I found strong correlations between gradients of lynx genetic structure and gradients of climate and land cover in Ontario. My findings suggest that increases in winter air temperature, decreases in snow depth, and loss of suitable habitat will result in further loss of genetic diversity in peripheral populations of lynx. Consequently, the adaptive potential of lynx populations on the southern range periphery could decline. In conclusion, my dissertation demonstrates the varying influences that contemporary landscape structure and climate gradients can have on genetic diversity and differentiation of different species.

Author Keywords: Circuitscape, genetic network, landscape genetics, Lynx canadensis, Martes americana, range shift

The objective of this thesis was to assess the influence of anthropogenic sulphur (S) and nitrogen (N) deposition, and harvesting on soil acidity and calcium (Ca2+), magnesium (Mg2+), potassium (K+) and N soil pools in plantation forest soils in Ireland. The response to reductions in anthropogenic S deposition was assessed using temporal trends in soil solution chemistry at two long-term monitoring plots--one on a blanket peat, the other on a peaty podzol. At the peat site, there was little evidence of a response to reductions in throughfall non marine sulphate (nmSO₄2-) and acidity; soil water acidity was determined by organic acids. In addition, temporal variation in soil water did not respond to that in throughfall. In the podzol, reductions in anthropogenic S and H+ deposition led to a significant improvement in soil water chemistry at 75 cm; pH increased and total aluminum (Al_tot) concentrations declined. The impact of harvest scenarios on exchangeable Ca2+, Mg2+ and K+ pools was assessed using input-output budgets at 40 sites (30 spruce, 10 pine). Harvest scenarios were stem-only harvest (SOH), stem plus branch harvest (SBH) and stem, branch and needle harvest (whole-tree harvesting; WTH). Average K+ and Mg2+ budgets were positive under these scenarios. However, exchangeable K+ pools were small and due to uncertainty in K+ budgets, could be depleted within one rotation. Average Ca2+ budgets for spruce were balanced under SOH, but negative under SBH and WTH. Nitrogen deposition was high, between 5 and 19 kg N ha-1 yr-1, but was balanced by N removal in SOH. However, N budgets were under SBH and WTH, indicating that these harvesting methods would lead to depletion of soil N over the long-term. Finally, monitoring of N cycling at a spruce plot indicated that N deposition was contributing to large NO₃- leaching, and as such the site was N saturated. However, N cycling did not fit the criteria of the N saturation hypothesis; instead leaching was directly related to N deposition and supported the model of kinetic N saturation.

Author Keywords: acidic deposition, base cations, input-output budgets, Ireland, nitrogen, whole-tree harvesting

Sexual reproduction in flowering plants depends on investment in reproduction, the mode of pollen transfer, the availabilities of nutrient resources and potential mates, and the spatial scales over which these processes take place. In this thesis, I studied the general reproductive biology of Shepherdia canadensis (L.) Nutt. (Elaeagnaceae) and the suite of pollinators that visit the plants in Ivvavik National Park, Yukon, Canada. Across ten sites, I found that S. canadensis females were larger than males, but males produced more flowers than females at most sites. Males typically occurred at higher frequencies than females with the average male to female sex ratio being 1.19 ± 0.08 (mean ± SE, n = 10 sites). Both shrub size and flower production were significantly influenced by interactions between soil nitrogen and sex. Insect visitors to S. canadensis flowers were primarily ants and flower flies (Syrphidae), but exclusion experiments indicated that visitation by flying insects yielded greater fruit production than visitation by crawling insects. I found that fruit set was limited by the density of males within populations, but only over small distances (4-6 m). This is the first study to demonstrate that female reproductive success of a generalist-pollinated dioecious plant is limited by the density of males over small spatial scales.

Author Keywords: dioecy, pollinators, sex ratio, sexual dimorphism, Shepherdia canadensis

The Serial Discontinuity Concept describes the downstream recovery of key biophysical variables below an impoundment. With the proliferation of hydropower dams to meet increasing societal demands, further refinement and understanding of the Serial Discontinuity Concept is needed to accurately predict downstream impacts and ensure the proper management of rivers. In this study, I examine SDC predictions on physical, chemical and biological recovery in regulated rivers providing evidence from 1) a comprehensive literature review and 2) a formal test using two regulated rivers in Northern Ontario. I specifically address how these changes are reflected in benthic invertebrate abundance, diversity, and community composition. The literature review and case studies support the predicted recovery of temperature, periphyton, substrate, and drift. In addition, the study suggests that two recovery gradients exist in regulated rivers: 1) a longer, thermal gradient taking up to hundreds of kilometres downstream; and 2) a shorter, resource subsidy gradient recovering within 1-4 km downstream of an impoundment. Total benthic invertebrate abundance varies considerably and depends on the degree of flow alteration and resource subsidies from the upstream reservoir. In contrast, benthic diversity is reduced below dams irrespective of dam location and operation with little recovery observed downstream. Contrary to SDC predictions, the longitudinal gradient in regulated rivers is not a compaction of functional changes seen over several stream orders in natural rivers but a response to dam design and reservoir conditions. Stoneflies and dragonflies are particularly sensitive to regulation while filter feeding invertebrates are enhanced. Ward and Stanford's (1983) Serial Discontinuity Concept is still a useful framework for testing hypotheses. Future studies should further expand the SDC through empirical estimation within the context of the landscape to gain a better scientific understanding of regulated river ecology.

Author Keywords: benthic invertebrates, dams, longitudinal, recovery, River Continuum Concept, Serial Discontinuity Concept

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

Climate change has had numerous impacts on species' distributions by shifting suitable habitat to higher latitudes and elevations. These shifts pose new challenges to biodiversity management, in particular translocations, where suitable habitat is considered crucial for the reintroduced population. De-extinction is a new conservation tool, similar to reintroduction, except that the proposed candidates are extinct. However, this novel tool will be faced with similar problems from anthropogenic change, as are typical translocation efforts. Using ecological niche modelling, I measured suitability changes at translocation sites for several Holarctic mammal species under various climate change scenarios, and compared changes between release sites located in the southern, core, and northern regions of the species' historic range. I demonstrate that past translocations located in the southern regions of species' ranges will have a substantial decline in environmental suitability, whereas core and northern sites exhibited the reverse trend. In addition, lower percentages (< 50% in certain scenarios) of southern sites fall above the minimal suitability threshold for current and long-term species occurrence. Furthermore, I demonstrate that three popular de-extinction candidate species have experienced changes in habitat suitability in their historic range, owing to climate change and increased land conversion. Additionally, substantial increase in potentially suitable space is projected beyond the range-limits for all three species, which could raise concerns for native wildlife if de-extinct species are successfully established. In general, this thesis provides insight for how the selection of translocation sites can be more adaptable to continued climate change, and marks perhaps the first rigorous attempt to assess the potential for species de-extinction given contemporary and predicted changes in land use and climate.

Author Keywords: climate change, de-extinction, ecological niche models, MaxEnt, reintroduction, translocation

In this study, I investigate the effects of nitrogen (N) and phosphorus (P) on the nutritional stoichiometry and growth of filamentous green algae of the family Zygnemataceae in situ and ex situ. I found a mean of Carbon (C):N:P ratio of 1308:66:1 for populations growing in the Kawartha Lakes of southern Ontario during the summer of 2012. FGA stoichiometry was variable, with much of the variation in algal P related to sediment P (p < 0.005, R2 = 0.58). Despite large variability in their cellular nutrient stoichiometry, laboratory analysis revealed that Mougeotia growth rates remained relatively consistent around 0.28 day-1. In addition, Mougeotia was found to be weakly homeostatic with respect to TDN:TDP supply (1/HNP = 0.32). These results suggest that FGA stoichiometry and growth rates are affected by sediment and water N and P. However, they will likely continue to grow slowly throughout the summer despite variable nutrient supply.

Author Keywords: Chlorophyll concentration, Filamentous algae, Growth rate, Homeostatic regulation, Nutritional stoichiometry

Consequences of habitat loss and fragmentation include smaller effective population sizes and decreased genetic diversity, factors that can undermine the long-term viability of large carnivores that were historically continuously distributed. I evaluated the historical and contemporary genetic structure and diversity of American black bears (Ursus americanus) in Ontario, where bear habitat is largely contiguous, except for southern regions that experience strong anthropogenic pressures. My objectives were to understand gene flow patterns in a natural system still largely reflective of pre-European settlement to provide context for the extent of genetic diversity loss in southern populations fragmented by anthropogenic influences. Phylogeographic analyses suggested that Ontario black bears belong to a widespread "continental" genetic group that further divides into 2 subgroups, likely reflecting separate recolonization routes around the Great Lakes following the Last Glacial Maximum. Population genetic analyses based on individual genotypes showed that Ontario black bears are structured into 3 contemporary genetic clusters. Two clusters, located in the Northwest (NW) and Southeast (SE), are geographically vast and genetically diverse. The third cluster is less diverse, and spatially restricted to the Bruce Peninsula (BP). Microsatellite analyses revealed that the NW and SE clusters are weakly differentiated from each other relative to mitochondrial DNA findings, suggesting male-biased dispersal and isolation by distance across the province. I also conducted simulations to assess competing hypotheses that could explain the reduced genetic diversity on the BP, which supported a combination of low migration and recent demographic bottlenecks. I showed that management actions to increase genetic variation in BP black bears could include restoring landscape connectivity between BP and SE; however, the irreversible human footprint in the area makes regular translocations from SE individuals a more practical alternative. Overall, my work suggests that: 1) historical genetic processes in Ontario black bears were likely predominated by isolation by distance, 2) large mammalian carnivores such as black bears can become isolated and experience reduced diversity in only a few generations, and 3) maintaining connectivity in regions under increased anthropogenic pressures could prevent populations from becoming small and geographically and genetically isolated, and should be a priority for conserving healthy populations.

Author Keywords: American black bear, carnivore, conservation genetics, Ontario, phylogeography, population genetics