Biology

Researchers have shown increasing interest in biological invasions for the associated ecological and economic impacts as well as for the opportunities they offer to study the mechanisms that induce range expansion in novel environments. I investigated the strategies exhibited by invasive species that facilitate range expansion. Invasive populations exhibit shifts in life-history strategy that may enable appropriate responses to novel biotic and abiotic factors encountered during range expansion. The spatio-temporal scales at which these shifts occur are largely unexplored. Furthermore, it is not known whether the observed dynamic shifts represent a consistent biological response of a given species to range shifts, or whether the shifts are affected by the abiotic characteristics of the new systems. I examined the life-history responses of female round gobies Neogobius melanastomus across fine and coarser spatial scales behind the expansion front and investigated whether invasive populations encountering different environmental conditions (Ontario vs France) exhibited similar life-history shifts. In both study systems, I found an increase in reproductive investment at invasion fronts compared to longer established areas at coarse and fine scales. The results suggest a similar response to range shifts, or a common invasion strategy independent of environmental conditions experienced, and highlight the dynamic nature of an invasive population's life history behind the invasion front.

The second part of my research focused on the development of an appropriate eDNA method for detecting invasive species at early stages of invasion to enable early detection and rapid management response. I developed a simple, inexpensive device for collecting water samples at selected depths for eDNA analysis, including near the substrate where eDNA concentration of benthic species is likely elevated. I also developed a protocol to optimise DNA extraction from water samples that contain elevated concentration of inhibiters, in particular near-bottom samples. Paired testing of eDNA and conventional surveys was used to monitor round goby expansion along its invasion pathway. Round gobies were detected in more sites with eDNA, permitting earlier, more accurate, upstream detection of the expansion front. My study demonstrated the accuracy and the power of using eDNA survey method to locate invasion fronts.

Author Keywords: Age-specific reproductive investment, DNA extraction, Energy allocation, Fecundity, Invasion front, Range expansion

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

Effective population size (Ne) is a foundational concept in conservation biology, in part due to its relationship to the adaptive potential of populations. Although Ne is often estimated for wild populations, it is rarely calibrated against actual population estimates (Nc) other than to produce Ne/Nc ratios. This project used demographic and genetic data for from two intensively-studied populations of lake trout (Salvelinus namaycush) in Ontario's Experimental Lake Area (ELA) as baseline data for evaluating the performance of multiple Ne estimators. Several temporal and single-time (point) genetic methods of estimating Ne were compared against demographic Ne estimates and known population data, as well as variation and consistency within and among Ne estimators. Changes in genetic Ne estimates over time were also compared to changes in demographic structure and fluctuating census estimates, including the effect of an experimentally manipulated population bottleneck on demographic and genetic Ne estimates during population reduction and recovery. Sampling years that included the most pre-, during and post-bottleneck data revealed the lowest estimates using temporal estimators (Ne = 16 to 18) despite pre- and post-bottleneck census estimates of 591 and 565. Estimation of Ne had increasingly tighter confidence intervals as sample sizes approached the actual number of breeding individuals in each population. Performance differences among the tested estimators highlight their potential biases and reliance on different assumptions, illustrating their potential value and caveats for assessing adaptive potential of wild populations.

Author Keywords: Effective Population Size, Experimental Lakes Area, Fish Population Assessment, Lake Trout, Population Demographics, Population Genetics

Leaf litter decomposition represents a major pathway for nutrient cycling and carbon flow in aquatic ecosystems, and macroinvertebrates play an important role in the processing of this material. To assess the causes of variable leaf breakdown and nutrient fluxes, I measured decomposition rates and the nutrient release ratios of decomposing leaf material across a broad latitudinal gradient in Ontario boreal lakes which varied in nutrients, temperature, and pH. I examined the effects of macroinvertebrates using inclusion and exclusion bags. Generally, leaves decomposed faster in nutrient-rich, warmer lakes. Macroinvertebrates increased decomposition rates but their effects were relatively small compared to regional effects of nutrients and temperature. In addition, we found differential effects of nutrients and temperature on nutrient release ratios, which were partially determined by the release and retention of N and P. These results indicate that changes in these important environmental lake variables could alter decomposition dynamics in Ontario lakes, with implications for nutrient cycling and the storage of this important external carbon source.

I studied the biogeography of predaceous diving beetles (Coleoptera: Dytiscidae) in two remote and understudied regions: the Far North of Ontario, and Akimiski Island, Nunavut. I identified 35 species from northern Ontario, including three first provincial records for Ontario, Acilius athabascae Larson (1975), Hygrotus unguicularis (Crotch 1874), and Nebrioporus depressus (Fabricius 1775). I also documented three significant range extensions and six gap-infills for this region. I collected and identified 16 species from Akimiski Island, Nunavut, which include several first time reports for these species for the Nunavut territory. My collections also extend the known ranges of five species into the Hudson Plains Ecozone. This work provides important baseline information on the distribution of diving beetles for these regions.

Author Keywords: biodiversity, Boreal Shield, decomposition, Dytiscidae, ecological stoichiometry, macroinvertebrates

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

The growth of immature insects that develop on human remains can be used to estimate a post-mortem interval (PMI). PMI estimate confidence is negatively affected by: larval killing and preservation methods altering their size, limited morphological parameters to assess larval growth and therefore age, and few available alternate species development data. I compared live specimens to preserved specimens of the same development stages to assess the effects of killing-preservation techniques on morphology, and I introduce a new method that uses digital photography to examine maggot mouthparts for stage grading of Phormia regina. Digital photographic methods enable live insects to be quantified and improve approximations of physiological age. I then use these digital methods to produce a growth-rate model for a beetle commonly found on human remains, Necrodes surinamensis, providing data for PMI estimates that was previously unavailable.

Author Keywords: Forensic Entomology, Insect development, Morphometrics, Necrodes surinamensis, Phormia regina, Postmortem interval

Polar bears are generally described as solitary, but features of their life cycles and

habitats regularly necessitate interaction. Effective conspecific assessment, including accurate

recognition and discrimination, likely confers benefits, especially to females accompanied by

dependent young. Individuals in the Southern (SH) and Western (WH) Hudson Bay

subpopulations are ideal for studying polar bear social behaviours because of the prolonged

high densities of the ice-free season. First, I looked outside family groups to model their fine

scale sociospatial organization on land. Capture locations were more likely to correspond to

family groups when there were fewer neighbouring bears, when a greater proportion of

neighbours were female, and when the focal individual and neighbours were significantly

related. Second, I looked within the family group to assess offspring recognition. Of 288

offspring in 207 family groups captured in the SH subpopulation from 1999 through 2013, only

one case of adoption (of a singleton) was observed.

Author Keywords: Adoption, Kin Recognition, Logistic Regression, Maternity Analysis, Social Discrimination, Sociospatial

The potential release of nanoparticles into aquatic environments is raising global concerns. As antimicrobials, silver nanoparticles (AgNPs) are among the most prominent form in use. Despite this, their fate, long-term toxicity, and ecological relevance have yet to be investigated largely under natural settings with seasonality and environmental complexity. To better understand the environmental significance, we released AgNPs into Lake 222 at the Experimental Lakes Area over two years. AgNPs remained suspended in the water column and were detected throughout the lake and in the lower food web. Total Ag concentrations ranged from below 0.07 to 18.9 μg L-1 in lake water, and were highly dynamic seasonally both in the epilimnion and hypolimnion depending on the physical, chemical and biological conditions of the lake. Approximately 60% of the measured Ag mass in October was present in the sediment in 2014 and 50% in 2015 demonstrating relatively high sedimentation and removal from the water column. During winter months, Ag was largely absent in the water column under the ice. After ice melt and before summer stratification, Ag concentrations increased in the lake suggesting AgNPs may not be tightly bound to the sediment and are able re-enter the water column during spring mixing events. Despite temporal variation, total Ag was highly synchronous across spatial locations for both years, indicating rapid dispersal upon lake entry. When investigating AgNP sizes using spICPMS, size distributions were similar across spatial locations, with the 40-60 nm size class constituting approximately 60% of all particles identified. Large aggregates (>100 nm) and dissolved Ag were infrequently detected within the lake. Ag accumulated in the lower food web ranging from 0.27-16.82 μg Ag mg C-1 in the bacterioplankton and 0.17-6.45 μg Ag mg C-1 in algae (particulate fraction). Partial least squares models revealed the highest predictors of Ag accumulation were dissolved nutrients including DOC, TDN, TDP in bacterioplankton. Major predictors for particulate Ag included temperature, dissolved oxygen, and sampling date. The diversity of predictors among biological compartments emphasizes the importance of understanding the role of environmental complexity within the lower food web. Despite Ag accumulation we did not detect strong negative effects on the lake food web. An increase in particulate and bacterioplankton chlorophyll-a occurred after addition in contrast to reference lakes, which may indicate a hormetic response to low dose AgNP concentrations. Our findings provide the first whole-lake perspective regarding Ag fate and toxicity, suggesting small scale experiments may overestimate environmental

responses.

Author Keywords: Ecotoxicity, Fate, Lower food web, Silver Nanoparticles, Whole-lake addition

According to the Invasional Meltdown Hypothesis (IMH), invasive species may interact in their introduced range and facilitate future invasions. This study investigated the possibility that Alliaria petiolata, an invasive allelopathic herbaceous plant in Ontario, is facilitating invasions by additional alien species. Two allelopathic focal species were chosen for this study: the native Solidago canadensis and the invasive A. petiolata. Field surveys in southern Ontario that quantified plant biodiversity in plots that included one or both focal species revealed no support for the IMH, although fewer species co-existed with A. petiolata than with S. canadensis. A year-long recruitment experiment in Peterborough, Ontario, also produced results inconsistent with the IMH, although did provide some evidence that A. petiolata limited recruitment of other species. These results collectively show negative impacts on regional biodiversity by A. petiolata, even in the absence of an invasional meltdown.

Author Keywords: allelopathy, Alliaria petiolata, co-occurrence surveys, invasional meltdown hypothesis, invasive species, Solidago canadensis

Interspecific hybridization is an important evolutionary process which can contribute to the invasiveness of species complexes. In this dissertation I used the hybridizing species complex of cattails (Typha spp., Typhaceae) to explore some of the processes that could contribute to hybridization rates. Cattails in northeastern North America comprise the native T. latifolia, the non-native T. angustifolia, and their fertile hybrid, T. × glauca. First, I examined whether these taxa segregate by water depth as habitat segregation may be associated with lower incidence of hybridization. I found that these taxa occupy similar water depths and therefore that habitat segregation by water depth does not promote mating isolation among these taxa. I then compared pollen dispersal patterns between progenitor species as pollen dispersal can also influence rates of hybrid formation. Each progenitor exhibits localized pollen dispersal, and the maternal parent of first generation hybrids captures more conspecific than heterospecific pollen; both of which should lead to reduced hybrid formation. I then conducted controlled crosses using all three Typha taxa to quantify hybrid fertility and to parameterize a fertility model to predict how mating compatibilities should affect the composition of cattail stands. I found that highly asymmetric formation of hybrids and backcrosses and reduced hybrid fertility should favour the maintenance of T. latifolia under certain conditions. Finally, I used a population genetics approach to characterize genetic diversity and structure of Typha in northeastern North America to determine the extent to which broad-scale processes such as gene flow influence site-level processes. I concluded that hybrids are most often created within sites or introduced in small numbers rather than exhibiting broad-scale dispersal. This suggests that local processes are more important drivers of hybrid success than landscape-scale processes which would be expected to limit the spread of the hybrid. Though my findings indicate some barriers to hybridization in these Typha taxa, hybrid cattail dominates much of northeastern North America. My results therefore show that incomplete barriers to hybridization may not be sufficient to prevent the continued dominance of hybrids and that active management of invasive hybrids may be required to limit their spread.

Author Keywords: fertility model, genetic structure, Hybridization, invasive species, niche segregation, pollen dispersal