Burness, Gary

Due to climate change and managed care, birds may face interacting effects of novel diets and environmental temperatures. I examined how captive zebra finches (Taeniopygia guttata) responded to a 6-month acclimation to either thermoneutral or cool temperatures when provided with saturated or unsaturated fatty acid enriched diets. Body mass was lower on the unsaturated fatty acid diet, but only in cool conditions. Individuals in cool temperatures and saturated fatty acid enriched diets had heavier organs, but pectoralis mass and metabolic rate did not display an effect of diet. While birds on each treatments illustrated similar initial increases in plasma fatty acid unsaturation, composition and concentration of constituent fatty acid varied based on their diet and temperature. Individuals on the unsaturated fatty acid enriched diet and cool treatments had lower survivorship. Collectively, my research will allow researchers to better predict how birds will be impacted by shifting or novel environments.

Author Keywords: chronic acclimation, fatty acids, metabolic rate, Taeniopygia guttata, temperature, unsaturation

Many species of shorebirds depend on stopover sites to rest and refuel during their long-distance migrations. To determine how shorebirds use migratory stopover sites, we tracked three species of shorebirds at two stopover sites in British Columbia, Canada from 2018-2021 during northward and southward migration using automated telemetry. Western Sandpipers (Calidris mauri) stayed longer at the Fraser River Estuary (4-8 days) compared to Tofino (2-6 days). We assessed habitat use of Sanderlings (Calidris alba), Semipalmated Plovers (Charadrius semipalmatus), and Western Sandpipers between beaches and mudflat at the Tofino stopover site. Time spent at the beach and mudflat habitats varied by species, tidal period, time of day, migration period, and human disturbance. This study shows that different stopover sites, and habitats within stopover sites, offer a unique set of characteristics used by birds exhibiting varying migration strategies, highlighting the importance of conserving a diversity of migration stopover locations and habitats.

Author Keywords: habitat use, human disturbance, length of stay, migration, shorebird, stopover site

The influence of stress exposure on the body temperature of vertebrates has been known for nearly two-thousand years. While the proximate mechanisms supporting this phenomenon are well described, the ultimate mechanisms remain enigmatic. In this thesis, I propose a novel hypothesis which states that changes in body surface temperature (henceforth "surface temperature") following stress exposure occur to reduce energetic expenditure toward thermoregulation, thus freeing energy for use in the stress response (hereafter, the "Thermoprotective Hypothesis"). Using a paired experimental design, I first show that black-capped chickadees (Poecile atricapillus, Linnaeus, 1766) exposed to repeated stressors decrease their surface temperatures at low ambient temperatures, and increase their surface temperatures at high ambient temperatures relative to unstressed controls. These changes in surface temperature contribute to a relative reduction in heat loss in the cold, and a relative increase in heat dissipation in the warmth among stress-exposed individuals, thus reducing their energetic demands toward more costly thermoregulatory strategies. Next, I show that stress-induced changes in surface temperature are most pronounced in chickadees that experience naturally-occurring resource-restrictions, suggesting that this response occurs to balance allocation of energy among the stress response and thermoregulation (i.e. a true energetic trade-off ). Third, I show that the magnitudes of chronic, stress-induced changes in surface temperature are highly variable among, and highly consistent within, chickadees, therefore suggesting that this response could hold adaptive significance if such variation among individuals is heritable. Finally, using domestic pigeons (Columba livia domestica, Gmelin, 1789) as a model species, I show that stress-induced changes in surface temperature are highly pronounced at bare tissues with a critical role in thermoregulation for some avian species (the bill), when compared with responses at surrounding bare tissues (the eye region). Together, these findings strongly support the Thermoprotective Hypothesis and suggest that endotherms may trade energetic investment toward thermoregulation with that toward the stress response in resource-limiting environments.

Author Keywords: heat transfer, infrared thermography, stress physiology, surface temperature, thermoregulation, trade-off

In endotherms, physiological functioning is optimized within a narrow range of tissue temperatures, meaning that the capacity to dissipate body heat is an important parameter for thermoregulation and organismal performance. Yet, experimental research has found mixed support for the importance of heat dissipation capacity as a constraint on reproductive performance. To investigate the effects of heat dissipation capacity on organismal performance, I experimentally manipulated heat dissipation capacity in free-living tree swallows, Tachycineta bicolor, by trimming feathers overlying the brood patch, and monitored parental provisioning performance, body temperature, and offspring growth. I found that individuals with an enhanced capacity to dissipate body heat (i.e., trimmed treatment) provisioned their offspring more frequently, and reared larger offspring that fledged more consistently. Although control birds typically reduced their nestling provisioning rate at the highest ambient temperatures to avoid overheating, at times they became hyperthermic. Additionally, I examined inter-individual variation in body temperature within each treatment, and discovered that body temperature is variable among all individuals. This variability is also consistent over time (i.e., is repeatable), irrespective of treatment. Further, I found that individuals consistently differed in how they adjusted their body temperature across ambient temperature, demonstrating that body temperature is a flexible and repeatable physiological trait. Finally, I used a bacterial endotoxin (lipopolysaccharide) to examine the regulation of body temperature of captive zebra finches (Taeniopygia guttata) during an immune challenge. Exposure to lipopolysaccharide induces sickness behaviours, and results in a fever, hypothermia, or a combination of the two, depending on species and dosage. I asked what the relative role of different regions of the body (bill, eye region, and leg) is in heat dissipation/retention during the sickness-induced body temperature response. I found that immune-challenged individuals modulated their subcutaneous temperature primarily through alterations in peripheral blood flow, particularly in the legs and feet, detectable as a drop in surface temperature. These results demonstrate that the importance of regional differences in regulating body temperature in different contexts. Taken together, my thesis demonstrates that heat dissipation capacity can affect performance and reproductive success in birds.

Author Keywords: body temperature, heat dissipation, tree swallow, zebra finch

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

The distribution of animals is rarely random and is affected by various environmental factors. We examined space-use patterns, habitat selection and stress responses of whip-poor-wills to mining exploration activity.To the best of my knowledge, fine scale patterns such as the habitat composition within known home ranges or territories of eastern whip-poor-wills have not been investigated. Using a population at the northern edge of the distribution in an area surrounding a mining exploration site, we tested whether variations in habitat and anthropogenic disturbances influence the stress physiology of individuals. We found no effect of increased mining activity on the stress physiology of birds but found a significant scale-dependent effect of habitat on their baseline and stress-induced corticosterone levels, and we suggest that these are the result of variations in habitat quality. The importance of other factors associated with those habitat differences (e.g., insect availability, predator abundance, and microhabitat features) warrants further research.

Author Keywords: anthropogenic disturbances, Antrostomus vociferus, corticosterone, eastern whip-poor-will, habitat selection, radio-­telemetry

In the face of climate change, the persistence of cold-adapted species and populations will depend on their capacity for evolutionary adaptation of physiological traits. As a cold-adapted Ice Age relict species, lake trout (Salvelinus namaycush) are ideal for studying potential effects of climate change on coldwater fishes. I studied the thermal acclimation capacity and aerobic metabolism of age 2+ lake trout from four populations across four acclimation temperatures (8ºC, 11ºC, 15ºC, and 19ºC). One population had marginally significant higher active metabolic rate (AMR) and metabolic scope across all temperatures. There was no interpopulation variation for critical thermal maximum (CTM), standard metabolic rate (SMR), or thermal acclimation capacity. Acclimation resulted in a 3ºC increase in thermal tolerance and 3-fold increase in SMR for all populations. At 19ºC, SMR increased and AMR declined, resulting in sharply reduced metabolic scope for all populations. The limited intraspecific variation in thermal physiology suggests that climate change may threaten lake trout at the species rather than population level.

Author Keywords: Climate Change, Lake Trout, Metabolic Rate, Salvelinus namaycush, Temperature, Thermal Acclimation

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