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Resting metabolic rate (RMR) is the minimum energy expenditure necessary for survival. RMR varies widely both among and within species and a central question in evolutionary physiology concerns the functional basis for this variation. Juvenile North American red squirrels were used to investigate fitness consequences of variation in RMR by considering how expenditure relates to differences in food availability and to overwinter survival. Additionally, this thesis examines whether red squirrels exhibit phenotypic plasticity in RMR in response to varying levels of food availability. Results indicate that heavier juveniles with relatively low RMRs were more likely to survive overwinter. Moreover, these juveniles were capable of allocating more energy towards mechanical work and possessed larger food stores. Food supplemented yearlings exhibited higher RMRs than unsupplemented controls at the onset of the breeding season, while no difference in RMR was detected following termination of supplementation. / Wildlife Ecology

Classically, testosterone (T) was considered the principal regulator of aggression. However, recent studies in birds have found aggression and T uncoupled during the non-breeding season. Circulating testosterone comes with costs such as immunosuppression and energy expenditure. Instead, the pro-hormone, dehydroepiandrosterone (DHEA), is circulated and activated within the brain via conversion into oestradiol (E2), avoiding the costs associated with T. At present the site of DHEA synthesis is unknown. My thesis investigated the existence of an analogous pathway in non-breeding red squirrels (Tamiasciurus hudsonicus) with two studies: (a) a field study investigating the effects of E2 on aggression and (b) a laboratory study that attempted to determine the site of DHEA synthesis. I conclude that E2 regulates non-breeding aggression in red squirrels and that the adrenals are not the site of DHEA synthesis. My results suggest the existence of a mammalian analogue to the regulatory pathway found in birds.

Classically, testosterone (T) was considered the principal regulator of aggression. However, recent studies in birds have found aggression and T uncoupled during the non-breeding season. Circulating testosterone comes with costs such as immunosuppression and energy expenditure. Instead, the pro-hormone, dehydroepiandrosterone (DHEA), is circulated and activated within the brain via conversion into oestradiol (E2), avoiding the costs associated with T. At present the site of DHEA synthesis is unknown. My thesis investigated the existence of an analogous pathway in non-breeding red squirrels (Tamiasciurus hudsonicus) with two studies: (a) a field study investigating the effects of E2 on aggression and (b) a laboratory study that attempted to determine the site of DHEA synthesis. I conclude that E2 regulates non-breeding aggression in red squirrels and that the adrenals are not the site of DHEA synthesis. My results suggest the existence of a mammalian analogue to the regulatory pathway found in birds.

Northern Goshawk breeding habitat was conceptualized and analyzed as a system composed of different sized components organized within a hierarchy. The organization of the hierarchy was asymmetric with smaller components nested within and subordinate to larger components. Components considered in the breeding habitat hierarchy of goshawks were nest tree, nest site, nest territory, foraging area, and landscape. Choice at any level within the hierarchy was contrasted with the next level in the hierarchy. Thus it was found that goshawks chose nest trees with a largerdbh (79.0 ± 4.4 cm) than the nest site average (43 ± 2.7 cm) (p=0.000036); the averagedbh of the nest site (43.2± 3.1 cm) was significantly larger than the territory (33.2± 1.9cm) (p=0.018). The nest territory when contrasted with random unoccupied locations in the landscape demonstrated a composition of a central site of trees with largedbh and fewer stems when compared with the surrounding forest. I hypothesize that this pattern reduces thepredation risk at the nest site. I investigated the nest site relative to the assumed foraging area (radius of 3000m; approximately 3000ha). Goshawks selected nest sites which were significantly further from young clear cuts (>10 ha). The average distance from a nest site to a recentclearcut greater than 10 ha was 1,350 m of the nest while for unoccupied sites the average distance was 770 m (p = 0.011). Finally, at the landscape level, areas of approximately 10,000 ha, goshawks on Vancouver Island were more abundant, had more nest sites, and had higher productivity per active nest in old growth forests (>120 years) than in either second growth (majority less than 80 years) or fragmented landscapes. The majority of goshawk prey, as determined though pellet analysis was red squirrel (69%). Red squirrels were equally abundant in second growth and old growth yet less abundant in fragmented landscapes. Despite similar abundance of their main prey item goshawks were not as abundant in second growth forests as in old growth forests. I conclude that the structure of second growth and fragmented landscape reduces the availability of key prey species and therefor the suitability of these landscape fpr goshawks is lower. / Graduate

Previous research on the relationship between spatial memory and adult hippocampal neurogenesis has been controversial. In the present study, neurogenesis was compared between two natural populations of the same species that differ in their reliance on spatial memory to cache and retrieve stored food. Western red squirrels store food in a single site whereas eastern red squirrels store food in multiple sites. Neurogenesis was assessed using endogenous markers of the number of proliferating cells (Ki-67) and the number of immature neurons (DCX), and neuronal recruitment was determined by measuring the area of the dentate gyrus of the hippocampus. The number of proliferating cells, immature neurons and neuronal recruitment were enhanced in the eastern compared to the western red squirrels, reflecting the food storing strategies used by the squirrels. This suggests that there is a positive correlation between adult hippocampal neurogenesis and spatial learning and memory.

Previous research on the relationship between spatial memory and adult hippocampal neurogenesis has been controversial. In the present study, neurogenesis was compared between two natural populations of the same species that differ in their reliance on spatial memory to cache and retrieve stored food. Western red squirrels store food in a single site whereas eastern red squirrels store food in multiple sites. Neurogenesis was assessed using endogenous markers of the number of proliferating cells (Ki-67) and the number of immature neurons (DCX), and neuronal recruitment was determined by measuring the area of the dentate gyrus of the hippocampus. The number of proliferating cells, immature neurons and neuronal recruitment were enhanced in the eastern compared to the western red squirrels, reflecting the food storing strategies used by the squirrels. This suggests that there is a positive correlation between adult hippocampal neurogenesis and spatial learning and memory.

A trait must genetically correlate with fitness in order to evolve, however, theory suggests that strong directional selection should erode additive genetic variance (Va) in fitness and limit future evolutionary potential. Sexual antagonism and temporal fluctuations in selection are mechanisms that could maintain Va in fitness. Maternal genetic effects could be an additional source of adaptive genetic variation. I used ‘animal models’ to examine a long-term population of red squirrels to determine 1) if either sexual antagonism or temporal fluctuations in selection were maintaining direct Va in fitness or 2) if maternal genetic effects were a source of indirect Va in fitness. While there were environmental trade-offs on juvenile survival, neither sexual antagonism nor temporal fluctuations in selection maintained Va in fitness. Maternal genetic effects on fitness were significant and provide the Va in fitness needed for rapid microevolution. This is the first instance of maternal genetic effects demonstrated as the only genetic variance available for microevolution. / Northern Scientific Training Program, the Arctic Institute of North America, American Society of Mammologists, Queen Elizabeth II Graduate Scholarship in Science and Technology, NSERC Discovery (to Andrew McAdam), NSF (to Andrew McAdam)