No abstract

Summary1. We examined the eects of food and predators on population limitation in the arctic ground squirrel (Spermophilus parryii plesius Richardson) in the boreal forest of the south-western Yukon. We focused on ground squirrel reproduction and overwinter survival. 2. Squirrel populations were monitored by live-trapping and radio-telemetry from 1993 to the spring of 1996 on four control and four experimental areas (one predator exclosure treatment, two food addition treatments, and one predator exclosure plus food addition treatment). 3. Predator exclusion increased body condition, percentage lactating, percentage weaning litters, litter size, and doubled population density relative to controls, but had no eect on juvenile growth rate, overwinter survival, or juvenile emergence date. 4. Food addition advanced juvenile emergence date and increased adult body condition, percentage lactating, percentage weaning litters, litter size, population density relative to controls (4±7 fold), but had no eect on juvenile growth rate or overwinter survival. 5. Predator exclusion combined with food addition increased adult body condition, percentage lactating, percentage weaning litters, and population density relative to controls (19-fold). 6. We conclude that arctic ground squirrels in the boreal forest are limited by an interaction between food and predation, acting primarily through changes in reproduction, and that their impact on density was multiplicative.

We tested the hypothesis that adult male arctic ground squirrels (Spermophilus parryii plesius) exhibit an adaptive stress response during the mating period that may compromise their survival, whereas males at other times (nonreproductive adult males and juvenile males) have a normal functional stress response. We assessed the physiological responsiveness of the stress axis, of energy mobilization, and of the immune response by subjecting adult breeding males, adult nonbreeding males, and juvenile males to a hormonal challenge and an immunocompetence challenge. At the onset of the breeding season in spring, only 25–30% of the population were males, and of those present during the mating period, half disappeared soon thereafter, and 82% were not replaced by immigrants. Adult breeding males had the highest levels of free cortisol, the lowest maximum corticosteroid‐binding capacity, slight dexamethasone resistance, the lowest hematocrit, the lowest number of white blood cells, the highest number of eosinophils, and the poorest ability to respond to the foreign antigen challenge in comparison with the other two male classes. All of these characteristics were indicative of chronic stress in breeding males that may directly compromise their survival. Juvenile males in mid‐August also showed many, but not all of these characteristics, indicative of a prolonged period of stress, presumably associated with the period of dispersal. Testosterone levels remained high irrespective of age or breeding condition, decreased when dexamethasone was injected, and increased when ACTH was injected. These latter results are unique in mammals. High testosterone levels and their augmentation with stressors may play a key role in maintenance of aggressive behavior. We conclude that breeding male arctic ground squirrels exhibit an adaptive stress response in which they trade off survival for reproduction. We hypothesize that similar stress responses may have evolved in other species with comparable life histories.

Results of the study by Blackburn et al. (2004a) of avifauna on oceanic islands suggest that distance from the mainland and time since European colonization have major influences on species extinctions and that island area is a significant but secondary contributing factor. After augmenting the data of the study on geographical properties for some of the islands they examined, we used a causal analysis approach with structural equation modeling to reexamine their conclusions. In our model geographical properties of islands, such as island area and isolation, were considered constraints on biological factors, such as the number of introduced mammalian predators and existing number of avifauna, that can directly or indirectly influence extinction. Of the variables we tested, island area had the greatest total influence on the threat of extinction due to its direct and indirect effects on the size of island avifauna. Larger islands had both a greater number of threatened bird species and more avifauna, increasing the number of species that could become threatened with extinction. Island isolation also had a significant, positive, and direct effect on threats to island avifauna because islands farther from the mainland had fewer current extant avifauna. Time since European colonization had a significant negative, but relatively weaker, influence on threats compared with the traditional biogeographic factors of island area and distance to the mainland. We also tested the hypothesis that the amount of threat is proportionally lower on islands that have had more extinctions (i.e., there is a "filter effect"). Because the proportion of bird extinctions potentially explained only 2.3% of the variation in the proportion of threatened species on islands, our results did not support this hypothesis. Causal modeling provided a powerful tool for examining threat of extinction patterns of known and hypothesized pathways of influence.

Abstract. We tested the hypothesis that adult male arctic ground squirrels (Spermophilus parryii plesius) exhibit an adaptive stress response during the mating period that may compromise their survival, whereas males at other times (nonreproductive adult males and juvenile males) have a normal functional stress response. We assessed the physiological responsiveness of the stress axis, of energy mobilization, and of the immune response by subjecting adult breeding males, adult nonbreeding males, and juvenile males to a hormonal challenge and an immunocompetence challenge. At the onset of the breeding season in spring, only 25-30% of the population were males, and of those present during the mating period, half disappeared soon thereafter, and 82% were not replaced by immigrants. Adult breeding males had the highest levels of free cortisol, the lowest maximum corticosteroidbinding capacity, slight dexamethasone resistance, the lowest hematocrit, the lowest number of white blood cells, the highest number of eosinophils, and the poorest ability to respond to the foreign antigen challenge in comparison with the other two male classes. All of these characteristics were indicative of chronic stress in breeding males that may directly compromise their survival. Juvenile males in mid-August also showed many, but not all of these characteristics, indicative of a prolonged period of stress, presumably associated with the period of dispersal. Testosterone levels remained high irrespective of age or breeding condition, decreased when dexamethasone was injected, and increased when ACTH was injected. These latter results are unique in mammals. High testosterone levels and their augmentation with stressors may play a key role in maintenance of aggressive behavior. We conclude that breeding male arctic ground squirrels exhibit an adaptive stress response in which they trade off survival for reproduction. We hypothesize that similar stress responses may have evolved in other species with comparable life histories.

Urban environments are unique because fragments of natural or semi-natural habitat are embedded within a potentially permeable matrix of human-dominated areas, creating increased landscape and, potentially, habitat heterogeneity. In addition, urban areas can provide diet subsidies for wildlife species in the form of fruiting ornamental plants, trash, and domestic animals. Ecological opportunity in the forms of habitat and food heterogeneity are thought to be important mechanisms in maintaining individual specialization. Identifying which contexts, traits, and mechanisms determine the success or failure of individuals within an urban wildlife population could potentially provide predictions about which populations may succeed in human-dominated landscapes and which may experience local extinction. We used both scat and stable isotope analysis of whiskers to investigate the degree to which coyotes (Canis latrans) utilized anthropogenic subsidies and exhibited individual diet specialization across the urban-rural gradient in southern California. Land use surrounding scat and isotope sample locations was also evaluated to determine the effect of land cover on diet. Human food constituted a significant portion of urban coyote diet (22% of scats, 38% of diet estimated by stable isotope analysis). Domestic cats (Felis catus) and ornamental fruit and seeds were also important items in urban coyote diets. Consumption of anthropogenic items decreased with decreasing urbanization. In suburban areas, seasonality influenced the frequency of occurrence of anthropogenic subsidies with increased consumption in the dry season. The amount of altered open space (areas such as golf courses, cemeteries, and landscaped parks) nearby had a negative effect on the consumption of anthropogenic items in both urban and suburban areas. Contrary to our hypothesis, urban coyotes displayed reduced between-individual variation compared to suburban and rural coyotes. It is possible that the core urban areas of cities are so densely developed and subsidized that wildlife inhabiting these areas actually have reduced ecological opportunity. Suburban animals had the broadest isotopic niches and maintained similar individual specialization to rural coyotes. Wildlife in suburban areas still have access to relatively undisturbed natural areas while being able to take advantage of anthropogenic subsidies in neighboring residential areas. Therefore, areas with intermediate urban development may be associated with increased ecological opportunity and specialization.

We investigated the effect of local environment on the demography and population dynamics of arctic ground squirrels (Spermophilus parryii plesius ) by comparing reproduction, survival, and population trends of squirrels living in low elevation boreal forest and high elevation alpine tundra sites in southwestern Yukon Territory, Canada. Contrary to the trend for most birds and mammals, reproduction was significantly lower at the lower elevation and females living at higher elevation did not delay the age at which they first reproduced. Even though survival in the boreal forest was lower in summer than in the alpine, it was higher over winter so annual adult female survival was similar between sites. Sensitivity analysis of model parameters revealed that in the forest, population growth rate (l) was most sensitive to small changes in adult active season survival whereas for the alpine population, l was most sensitive to changes in juvenile winter survival. In their respective habitats, these parameters also showed high year to year variation and thus contributed greatly to the population trends observed. Even though ground squirrels persisted in the boreal forest, the measured demographic rates indicate the forest was sink habitat (lB/1) and may have relied on nearby grassy meadows for immigrants. In contrast, the alpine habitat maintained a ground squirrel population in the absence of immigration (l0/1).The variation in demographic rates between ground squirrels living at high and low elevation may arise from phenotypic responses of squirrels to different habitat structure. Arctic ground squirrels rely on sight to detect predators from a safe distance, and the boreal forest, with its lower visibility and higher predator density, appears to be suboptimal habitat.

2004. Comparision of discriminant function and classification tree analyses for age classification of marmots. Á/ Oikos 105: 575 Á/587.We evaluated the predictive power of two classification techniques, one parametric Á/ discriminant function analysis (DFA) and the other non-parametric Á/ classification and regression tree analysis (CART), in order to provide a non-subjective quantitative method of determining age class in Vancouver Island marmots (Marmota vancouverensis ) and hoary marmots (Marmota caligata ). For both techniques we used morphological measurements of known-age male and female marmots from two independent population studies to build and test predictive models of age class. Both techniques had high predictive power (69 Á/86%) for both sexes and both species. Overall, the two methods performed identically with 81% correct classification. DFA was marginally better at discriminating among older more challenging age classes compared to CART. However, in our test samples, cases with missing values in any of the discriminant variables were deleted and hence unclassified by DFA, whereas CART used values from closely correlated variables to substitute for the missing values. Therefore, overall, CART performed better (CART 81% vs DFA 76%) because of its ability to classify incomplete cases. Correct classification rates were approximately 10% higher for hoary marmots than for Vancouver Island marmots, a result that could be attributed to different sets of morphological measurements. Zygomatic arch breadth measured in hoary marmots was the most important predictor of age class in both sexes using both classification techniques. We recommend that CART analysis be performed on data-sets with incomplete records and used as a variable screening tool prior to DFA on more complete data-sets.