Burness, Gary

Many facilities attempt to alleviate the risk of chronic stress in captivity by providing environmental enrichment shown to minimize behavioural disorders and stress in several species. One potential form of enrichment used in zoos is training animals to perform rides for guests, however, the effect of this activity on the welfare of individual animals has never been examined. I validated the use of saliva for assessing stress in dromedary camels (Camelus dromedarius) an animal commonly used for rides. I then measured variation in salivary cortisol in four male camels during animal rides for guests at the Toronto Zoo. The camels were sampled during the ride season (from June to August) using four treatments: 1) in their pasture, 2) at the ride area not performing rides, 3) performing a low number of rides (n=50/day) and 4) performing a high number of rides (n=150/day). Furthermore, samples were taken before and after the ride season for comparison. There was a significant difference between the post-ride season treatment and the three treatments involving guest presence during the ride season (ride area, low rides, high rides. This indicates that performing rides is not a stressful experience based on the stress metrics I used, and suggests that rides may be a form of enrichment for dromedary camels.

Author Keywords: ACTH challenge, animal welfare, camels, environmental enrichment, salivary cortisol, stress

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

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

Glucocorticoids (e.g., corticosterone (CORT)) are hypothesized to mediate decisions regarding reproductive investment during breeding, but the directionality of the relationship is not clear. The CORT-fitness hypothesis posits that high levels of CORT arise from challenging environmental conditions in which an individual will conserve resources for future reproduction or self-maintenance, and thus result in lower reproductive success (a negative relationship). In contrast, the CORT-adaptation hypothesis suggests that, during energetically demanding periods, CORT will mediate physiological or behavioural changes that result in increased reproductive investment and success (a positive relationship). Inconsistencies arise due to the various species and life-history stages studied, and the complex interactions between fitness and glucocorticoids. Using an experimental approach, I investigated the relationship between CORT and reproductive success by manipulating baseline CORT levels in female tree swallows Tachycineta bicolor prior to laying using silastic implants. Implants failed to raise CORT levels of females during either incubation or the nestling stage, and maternal treatment had no effect on indices of fitness at either stage. Using a correlative approach, partial support for the CORT-adaptation hypothesis was found: There was a positive relationship between CORT and hatching success. This only occurred when CORT was measured during incubation, when baseline CORT levels may stimulate increased reproductive effort and success. In contrast, during the nestling stage baseline CORT levels were not related to reproductive investment or success. Maternal CORT levels during incubation also did not influence nestling phenotype, although nestling stress CORT levels were higher in individuals that survived to fledging. In conclusion, CORT mediates reproductive effort and success during some breeding stages, but it is still unclear why this is the case and whether this same pattern will prevail in other contexts.

Author Keywords: corticosterone, HPA axis, maternal effects, reproductive success, tree swallow

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

Due to their effective antibacterial and antifungal properties, silver nanoparticles (AgNPs) have quickly become the most commonly used nanomaterial, with applications in industry, medicine and consumer products. This increased use of AgNPs over the past decade will inevitably result in an elevated release of nanoparticles into the environment, highlighting the importance of assessing the environmental impacts of these nanomaterials on aquatic ecosystems. Although numerous laboratory studies have already reported on the negative effects of AgNPs to freshwater organisms, only a handful of studies have investigated the impacts of environmentally relevant levels of AgNPs on whole communities under natural conditions. This thesis examines the effects of chronic AgNP exposure on natural freshwater littoral microcrustacean, benthic macroinvertebrate and pelagic zooplankton communities. To assess the responses of these communities to AgNPs, I focused on a solely field-based approach, combining a six-week mesocosm study with a three-year whole lake experiment at the IISD – Experimental Lakes Area (Ontario, Canada). Our mesocosm study tested the effects of AgNP concentration (low, medium and high dose), surface coating (citrate- and polyvinylpyrrolidone [PVP]-coated AgNPs), and type of exposure (chronic and pulsed addition) on benthic macroinvertebrates in fine and stony sediments. Relative abundances of metal-tolerant Chironomidae in fine sediments were highest in high dose PVP-AgNP treatments; however, no negative effects of AgNP exposure were seen on biodiversity metrics or overall community structure throughout the study. I observed similar results within the whole lake study that incorporated a long-term addition of low levels of AgNPs to an experimental lake. Mixed-effects models and multivariate methods revealed a decline in all species of the littoral microcrustacean family Chydoridae in the final year of the study within our experimental lake, suggesting that this taxon may be sensitive to AgNP exposure; however, these effects were fairly subtle and were not reflected in the overall composition of littoral communities. No other negative effects of AgNPs were observed on the pelagic zooplankton or benthic macroinvertebrate communities. My results demonstrate that environmentally relevant levels of AgNPs have little impact on natural freshwater microcrustacean and benthic macroinvertebrate communities. Instead, biodiversity metrics and community structure are primarily influenced by seasonal dynamics and nutrient concentrations across both lakes. This thesis highlights the importance of incorporating environmental conditions and the natural variability of communities when examining the potential risks posed by the release of AgNPs into the environment, as simplistic laboratory bioassays may not provide an adequate assessment of the long-term impacts of AgNPs on freshwater systems.

Author Keywords: Benthic macroinvertebrates, IISD - Experimental Lakes Area, Littoral microcrustaceans, Silver nanoparticles, Whole lake experiment, Zooplankton

Understanding breeding and migration habits of Whimbrel (Numenius phaeopus) in the Hudson Bay Lowlands is important for the conservation of this population. I monitored Whimbrel at two breeding sites: the Churchill region of Manitoba and Burntpoint, Ontario. Annual average nest initiation timing was highly variable and successful nests were initiated significantly earlier than those that failed. Although nests were initiated significantly earlier at Burntpoint than Churchill, annual nest success quantified in program R MARK was similar across sites. Observed nest success rates were lower than historical records and most failure was due to predation. Annual nest survival varied widely and I used a generalized linear model to relate annual nest survival to annual average weather conditions. I observed weak relationships between annual nest survival and weather conditions in the northbound staging grounds. I tracked post-breeding migratory movements using the MOTUS radio telemetry system and observed consistent use of the mid-Atlantic coast of the United States during migration, especially among birds emerging from Churchill. In Burntpoint, I observed more variability in post-breeding migratory trajectories and significantly earlier post-breeding departure as compared to Churchill. The results of my study suggest differences in breeding and migration habits exist across nearby breeding populations, indicating that there is a need for population-specific conservation approaches for this declining species.

Author Keywords: Migration, Movement Ecology, Nesting Ecology, Nest Success, Shorebird conservation, Whimbrel

Contemporary evolution has the potential to help limit the biological impact of rapidly changing climates, however it remains unclear whether wild populations can respond quickly enough for such adaptations to be effective. In this thesis, I used the introduction of native North American Pumpkinseed (Lepomis gibbosus) into the milder climate of Europe over 140 years ago, as a 'natural' experiment to test for contemporary evolution to a change in climate in wild populations. In 2008, four outdoor pond colonies were established in central Ontario using adult Pumpkinseed from two native Canadian populations, and two non-native populations from northeastern Spain. By raising native and non-native Pumpkinseed within a common environment, this design minimized the impact of phenotypic plasticity on differential trait expression, and allowed me to interpret differences in the phenotype among pond-reared Pumpkinseed as evidence of genetic differences among populations. I demonstrated that Canadian and Spanish Pumpkinseed have similar thermal physiology except when acclimated to seasonally warm temperatures; trait differences are consistent with Spanish Pumpkinseed being better adapted to a warmer climate. Populations also had similar overwintering ecology, however some differences, such as higher survival under starvation conditions and greater energetic benefits associated with winter feeding, indicated that Canadian populations are better adapted to harsh winter conditions typical of the native range. Finally, I determined that the relatively fast life history expressed in wild European Pumpkinseed is largely driven by plastic responses to the local environment; however, the higher reproductive investment by European populations has a genetic basis. Most climate change research considers taxa that are expected to be negatively impacted by warming: my research demonstrates that even warm-tolerant taxa that are unlikely to experience strong climatic selective forces can respond to a warming environment through evolutionary changes. The potential for adaptive contemporary evolution in warm-tolerant taxa should be taken into account when predicting future ecosystem effects of climate change, and when planning management strategies for species introduced into novel climates.

Author Keywords: climate change, contemporary evolution, fish, non-native species, thermal biology, winter ecology

Recent declines in North American shorebird populations could be linked to habitat loss on the non-breeding grounds. Sea-level rise and increased frequency of coastal storms are causing significant erosion of barrier islands, thereby threatening shorebirds who rely on shoreline habitats for foraging. I conducted shorebird surveys on Bulls Island, South Carolina in the winters of 2018 and 2019 and examined habitat selection and foraging behaviour in Dunlin (Calidris alpina), Sanderling (Calidris alba), Semipalmated Plovers (Charadrius semipalmatus), and Piping Plovers (Charadrius melodus). Area, tidal stage, and invertebrate prey availability were important determinants of shorebird abundance, behaviour, and distribution. My study highlights the importance of Bulls Island's habitat heterogeneity to supporting a diverse community of non-breeding shorebirds. Considering both the high rate of erosion and the increased frequency of disturbance along the shoreline of the island, intertidal habitats should be monitored to predict negative effects of changes in habitat composition and area on non-breeding shorebirds.

Author Keywords: foraging behaviour, habitat loss, habitat selection, invertebrate prey, non-breeding, shorebirds

I examined Bank Swallow (Riparia riparia) colony persistence and occupancy, in lakeshore, river and man-made aggregate pit habitat. Habitat persistence was highest on the lakeshore and lowest in aggregate pits, likely due to annual removal and relocation of aggregate resources. Bank Swallow colonies in aggregate pit sites were more likely to persist if a colony was larger or if burrows were located higher on the nesting face. I also compared nest productivity and health factors of Bank Swallows in lakeshore and aggregate pit habitats. While clutch size was the same in both habitat types, the number of fledglings from successfully hatched nests was significantly higher in aggregate pit sites than from lakeshore sites. Mass of fledgling Bank Swallows did not differ significantly between habitat types, however mass of adults from aggregate pits decreased significantly over the nesting season. Parasite loads on fledgling Bank Swallows were significantly lower in aggregate pits than in lakeshore sites. According to these indicators, aggregate pits appear to provide equivalent or higher quality habitat for Bank Swallows than the natural lakeshore sites, making them adequate and potentially key for this species' recovery. Aggregate pit operators can manage for swallows by (1) creating longer, taller faces to attract birds and decrease predation, and (2) supplementing their habitat with water sources to encourage food availability.

Author Keywords: Aerial insectivore, aggregate pits, Bank Swallow, colony persistence, ectoparasites, substitute habitat