Kevin S. White

Habitat selection predicts genetic relatedness in an alpine ungulate

Landscape heterogeneity plays an integral role in shaping ecological and evolutionary processes. Despite links between the two disciplines, ecologists and population geneticists have taken different approaches to evaluating habitat selection, animal movement, and gene flow across the landscape. Ecologists commonly use statistical models such as resource selection functions (RSFs) to identify habitat features disproportionately selected by animals, whereas population genetic approaches model genetic differentiation according to the distribution of habitat variables. We combined ecological and genetic approaches by using RSFs to predict genetic relatedness across a heterogeneous landscape. We constructed sex- and season-specific resistance surfaces based on RSFs estimated using data from 102 GPS (global positioning system) radio-collared mountain goats (Oreamnos americanus) in southeast Alaska, USA. Based on mountain goat ecology, we hypothesized that summer and male surfaces would be the best predictors of relatedness. All individuals were genotyped at 22 microsatellite loci, which we used to estimate genetic relatedness. Summer resistance surfaces derived from RSFs were the best predictors of genetic relatedness, and winter models the poorest. Mountain goats generally selected for areas close to escape terrain and with a high heat load (a metric related to vegetative productivity and snow depth), while avoiding valleys. Male- and female-specific surfaces were similar, except for winter, for which male habitat selection better predicted genetic relatedness. The null models of isolation-by-distance and barrier only outperformed the winter models. This study merges high-resolution individual locations through GPS telemetry and genetic data, that can be used to validate and parameterize landscape genetics models, and further elucidates the relationship between landscape heterogeneity and genetic differentiation.

Benefits of migration in relation to nutritional condition and predation risk in a partially migratory moose population.

The costs and benefits of alternative migratory strategies are often framed in the context of top-down and bottom-up effects on individual fitness. This occurs because migration is considered a costly behavioral strategy that presumably confers explicit benefits to migrants in the form of either decreased predation risk (predation risk avoidance hypothesis) or increased nutrition (forage maturation hypothesis). To test these hypotheses, we studied a partially migratory moose (Alces alces) population and contrasted explicit measures of predation risk (i.e., offspring survival) and nutrition (i.e., accumulation of endogenous energy reserves) between resident and migratory subpopulations. We relied on data collected from migratory and nonmigratory radio-marked moose (n = 67) that inhabited a novel study system located in coastal Alaska between 2004 and 2010. In this area, 30% of the population resides year-round on a coastal foreland area, while 48% migrate to either a small island archipelago or a subalpine ridge system (the remainder exhibited one of six different low-occurrence strategies). Overall, we determined that accumulation of body fat during the growing season did not differ between migratory or resident modalities. However, calf survival was 2.6-2.9 times higher for individuals that migrated (survival, islands = 0.49 +/- 0.16 [mean +/- SE], n = 35; ridge = 0.52 +/- 0.16, n = 33) than those that did not (survival, resident = 0.19 +/- 0.08, n = 57). Our results support the predation risk avoidance hypotheses, and suggest that migration is a behavioral strategy that principally operates to reduce the risk of calf predation and does not confer explicit nutritional benefits. We did not directly detect trade-offs between predation risk and nutrition for migratory individuals. Yet we identify an indirect life history mechanism that may mildly dampen the apparent fitness benefits of migration. The proximate factors accounting for differences in migration-specific neonate survival are likely linked to accessibility of refugial habitats for moose at local and landscape scales, landscape factors that affect hunting efficacy of large carnivores, and interactions with rural human communities. Conservation of ungulate populations can be aided by integrating knowledge about migratory behavior, life history strategies, and factors that alter ungulate vulnerability, particularly those induced by human activity.

Mountain Goat Survival in Coastal Alaska: Effects of Age, Sex, and Climate

ABSTRACT Ecological theory predicts that individual survival should vary between sex and age categories due to differences in allocation of nutritional resources for growth and reproductive activities. During periods of environmental stress, such relationships may be exacerbated, and affect sex and age classes differently. We evaluated support for hypotheses about the relative roles of sex, age, and winter and summer climate on the probability of mountain goat (Oreamnos americanus) survival in coastal Alaska. Specifically, we used known-fates analyses (Program MARK) to model the effects of age, sex, and climatic variation on survival using data collected from 279 radio-marked mountain goats (118 M, 161 F) in 9 separate study areas during 1977–2008. Models including age, sex, winter snowfall, and average daily summer temperature (during Jul–Aug) best explained variation in survival probability of mountain goats. Specifically, our findings revealed that old animals (9+ yr) have lower survival than younger animals. In addition, males tended to have lower survival than females, though differences only existed among prime-aged adult (5–8 yr) and old (9+ yr) age classes. Winter climate exerted the strongest effects on mountain goat survival; summer climate, however, was significant and principally influenced survival during the following winter via indirect effects. Furthermore, old animals were more sensitive to the effects of winter conditions than young or prime-aged animals. These findings detail how climate interacts with sex and age characteristics to affect mountain goat survival. Critically, we provide baseline survival rate statistics across various age, sex, and climate scenarios. These data will assist conservation and management of mountain goats by enabling detailed, model-based demographic forecasting of human and/or climate-based population impacts.

Effects of Snow on Sitka Black‐Tailed Deer Browse Availability and Nutritional Carrying Capacity in Southeastern Alaska

Abstract Snow affects the nutritional ecology of northern ungulates during winter through burial of important winter forages. We used nonlinear regression analyses to model snow-burial dynamics of blueberry (Vaccinium spp.) browse biomass, a key winter food item of Sitka black-tailed deer (Odocoileus hemionus sitchensis) in southeastern Alaska, USA. During November 2003–March 2004 we collected data from 546 individually marked twigs located on 100 plants of differing sizes and architectures across a range of snow depths. In general, browse biomass became buried and unavailable to deer at snow depths substantially lower than prewinter twig heights. Plant architecture and plant height were related to the probability of a twig being buried. Probability of twig burial was higher on plants with lateral than on those with erect architectures. Twig height also affected the probability of burial by snow but the relationship was complex. For twigs located on erect plants, probability of burial was greatest for twigs near the bottom and top of the plant due to ground-up burial and bending of flexible apex stems, respectively. We used estimated nonlinear equations to model blueberry browse availability in a simulated upland old-growth habitat patch subject to a range of snow depths. We then compared subsequent estimates of deer winter nutritional carrying capacity for this habitat patch to findings derived using an alternative, simple linear (ground-up) model of winter-browse burial by snow. Comparisons indicated that ground-up models of browse burial overestimated browse availability and nutritional carrying capacity for most snow depths. Our findings demonstrate the importance of applying detailed snow-burial models when characterizing nutritional landscape of northern ungulates during winter.

Deciphering translocations from relicts in Baranof Island mountain goats: is an endemic genetic lineage at risk?

Human-mediated movement of wildlife is a common practice in North America. Some translocations have occurred where local populations were thought to be extinct or simply not present. In Alaska’s Alexander Archipelago, mountain goats (Oreamnos americanus) were not considered indigenous and were introduced to Baranof Island in 1923. However, a range-wide survey using microsatellites and mitochondrial DNA revealed a distinct genetic subpopulation endemic to the island. In this study, we attempted to clarify the evolutionary history of the mountain goats on Baranof Island by examining sequence variation in the Y chromosome. We first screened five regions of the Y chromosome in a subset of mountain goats from across their native range. We detected a single polymorphic site in the SRY promoter, and subsequently sequenced this gene in 100 mountain goats. A unique Y chromosome polymorphism was restricted to Baranof Island and an area near Haines, Alaska, and not detected in the presumed source population. An island-to-mainland dispersal scenario from a cryptic refugial population during the retreat of the Cordilleran ice-sheet would account for this distribution. Overall, these data support the hypothesis that a glacial relict population of mountain goats was present on the island prior to introduction. Based on a combination of mitochondrial, microsatellite, and Y chromosome data, we recommend recognizing Baranof Island mountain goats as an evolutionary significant unit.

Mountain goat resource selection in relation to mining-related disturbance

Industrial development can have important direct and indirect effects on wildlife populations. Resource selection function (RSF) modeling provides a powerful tool for assessing the effects of industrial development on spatial use patterns of wildlife. Among North American large mammal species, mountain goats Oreamnos americanus are particularly sensitive to human disturbance. In this study mountain goat seasonal resource selection patterns were examined using GPS radio collar (n = 79 individuals) and remote sensing data in a GIS framework across a 491 km2 regional mountain range in southeast Alaska, 2005–2015. The resulting global RSF model was then applied across a limited spatial extent centered on an industrial mining site in order to assess whether mining activity altered expected spatial use patterns at different distances from the mine. Using a quasi treatment—control experimental framework we examined the occurrence of spatially explicit mine disturbance thresholds. In general, resource selection modeling indicated that mountain goats selected for steep, rugged terrain in close proximity to cliffs in areas with high solar exposure; and they selected for lower elevations in winter than in summer. Mountain goat selection for rugged terrain and proximity to cliffs was stronger in winter than summer. RSF model applications indicated that mountain goat use of predicted habitat in close proximity to the mine was lower than expected at distances up to 1800 m in winter and 1000 m in summer. Because lower elevation winter habitat is closer to mining activity than summer habitat, a greater percentage (42%) of winter habitat within the analysis area was affected by mining activity. The resulting net loss of functional winter habitat carrying capacity is likely to have long-term negative implications for the local mountain goat population. In places where mining is proposed, development should avoid areas within 1800 m of mountain goat winter habitat. In places where mining is already occurring within disturbance thresholds (such as this study), long-term monitoring and more detailed field studies should be conducted to more fully understand population-level consequences of disturbance and identify practicable mitigation measures that have a high probability of success.

Parturition site selection in moose (Alces alces): evidence for social structure

Female natal philopatry has often been implicated as an important factor in moose (Alces alces) home range formation, with many populations showing behavioral evidence of sympatric home ranges among related individuals. However, previous genetic studies have failed to detect genetic subpopulation structure, leading to questions as to whether philopatry is a significant factor contributing to intra- and inter-population genetic structure. Here, we examine calving location data from radiomarked individuals (n = 110) collected in 2 separate populations in Berners Bay and Gustavus, Alaska, to examine the extent to which genetic structure originating from philopatry is evident at fine spatial scales. When populations were combined, their overall relatedness correlogram was significant (P < 0.001), with mean r = 0.079 between 0 km and 1.5 km (P = 0.079). Additionally, 13% of the population shows significantly positive relatedness to their 4 nearest neighbors, with an overall average r = 0.19 of those focal individuals to their neighbors. We suggest that habitat structure, especially linear habitats (i.e., river valleys), or habitat bounded with barriers to dispersal, may be a factor in promoting the development of this structure. This study presents the 1st known evidence for fine-scale social genetic structure in moose and natal philopatry to calving locations in moose. In the context of natal philopatry within cervids, these findings identify several cervid-wide commonalities.

A Survey of Bacterial Respiratory Pathogens in Native and Introduced Mountain Goats (Oreamnos americanus)

Abstract In contrast to broad range expansion through translocations, many mountain goat (Oreamnos americanus) populations have shown signs of decline. Recent documentation of pneumonia in mountain goats highlights their susceptibility to bacterial pathogens typically associated with bighorn sheep (Ovis canadensis) epizootics. Respiratory pathogen communities of mountain goats are poorly characterized yet have important implications for management and conservation of both species. We characterized resident pathogen communities across a range of mountain goat populations as an initial step to inform management efforts. Between 2010 and 2017, we sampled 98 individuals within three regions of the Greater Yellowstone Area (GYA), with a smaller sampling effort in southeast Alaska, US. Within the GYA, we detected Mycoplasma ovipneumoniae in two regions and we found at least two Pasteurellaceae species in animals from all regions. Mannheimia haemolytica was the only pathogen that we detected in southeast Alaska. Given the difficult sampling conditions, limited sample size, and imperfect detection, our failure to detect specific pathogens should be interpreted with caution. Nonetheless, respiratory pathogens within the GYA may be an important, yet underappreciated, cause of mountain goat mortality. Moreover, because of the strong niche overlap of bighorn sheep and mountain goats, interspecific transmission is an important concern for managers restoring or introducing mountain ungulates within sympatric ranges.

Mating effort and space use of an alpine ungulate during the rut

In ungulates, the rut generally leads to increased intra and interpopulation movements for males. Because movements induce energetic costs and missed feeding opportunities, they could be an indication of male mating effort. We studied space use of 44 male mountain goats (Oreamnos americanus) from three neighboring subpopulations in southeast Alaska, during the rut from 2005 to 2008. Using mixed models and an information theoretic approach with AIC, we analyzed the relationships between individual traits of males and their space use. We found no indication of breeding migration between subpopulations. Distances between individual seasonal ranges were not related to any individual trait. Daily movements, home range sizes, and total distance traveled during the rut did not vary with mass or age of individuals. As such, effects of individual traits on male space use during the rut appear weak and observed space use patterns do not support any of the main mating effort hypotheses.