Navigating snowscapes: scale-dependent responses of mountain sheep to snowpack properties.

Winters are limiting for many terrestrial animals due to energy deficits brought on by resource scarcity and the increased metabolic costs of thermoregulation and traveling through snow. A better understanding of how animals respond to snow conditions is needed to predict the impacts of climate change on wildlife. We compared the performance of remotely sensed and modeled snow products as predictors of winter movements at multiple spatial and temporal scales using a data set of 20,544 locations from 30 GPS-collared Dall sheep (Ovis dalli dalli) in Lake Clark National Park and Preserve, Alaska, USA from 2005 to 2008. We used daily 500-m MODIS normalized difference snow index (NDSI), and multi-resolution snow depth and density outputs from a snowpack evolution model (SnowModel), as covariates in step selection functions. We predicted that modeled snow depth would perform best across all scales of selection due to more informative spatiotemporal variation and relevance to animal movement. Our results indicated that adding any of the evaluated snow metrics substantially improved model performance and helped characterize winter Dall sheep movements. As expected, SnowModel-simulated snow depth outperformed NDSI at fine-to-moderate scales of selection (step scales < 112 h). At the finest scale, Dall sheep selected for snow depths below mean chest height (<54 cm) when in low-density snows (100 kg/m3 ), which may have facilitated access to ground forage and reduced energy expenditure while traveling. However, sheep selected for higher snow densities (>300 kg/m3 ) at snow depths above chest height, which likely further reduced energy expenditure by limiting hoof penetration in deeper snows. At moderate-to-coarse scales (112-896 h step scales), however, NDSI was the best-performing snow covariate. Thus, the use of publicly available, remotely sensed, snow cover products can substantially improve models of animal movement, particularly in cases where movement distances exceed the MODIS 500-m grid threshold. However, remote sensing products may require substantial data thinning due to cloud cover, potentially limiting its power in cases where complex models are necessary. Snowpack evolution models such as SnowModel offer users increased flexibility at the expense of added complexity, but can provide critical insights into fine-scale responses to rapidly changing snow properties.

Habitat selection by Dall's sheep is influenced by multiple factors including direct and indirect climate effects.

Arctic and boreal environments are changing rapidly, which could decouple behavioral and demographic traits of animals from the resource pulses that have shaped their evolution. Dall's sheep (Ovis dalli dalli) in northwestern regions of the USA and Canada, survive long, severe winters and reproduce during summers with short growing seasons. We sought to understand the vulnerability of Dall's sheep to a changing climate in Lake Clark National Park and Preserve, Alaska, USA. We developed ecological hypotheses about nutritional needs, security from predators, energetic costs of movement, and thermal shelter to describe habitat selection during winter, spring, and summer and evaluated habitat and climate variables that reflected these hypotheses. We used the synoptic model of animal space use to estimate parameters of habitat selection by individual females and calculated likelihoods for ecological hypotheses within seasonal models. Our results showed that seasonal habitat selection was influenced by multiple ecological requirements simultaneously. Across all seasons, sheep selected steep rugged areas near escape terrain for security from predators. During winter and spring, sheep selected habitats with increased forage and security, moderated thermal conditions, and lowered energetic costs of movement. During summer, nutritional needs and security influenced habitat selection. Climate directly influenced habitat selection during the spring lambing period when sheep selected areas with lower snow depths, less snow cover, and higher air temperatures. Indirectly, climate is linked to the expansion of shrub/scrub vegetation, which was significantly avoided in all seasons. Dall's sheep balance resource selection to meet multiple needs across seasons and such behaviors are finely tuned to patterns of phenology and climate. Direct and indirect effects of a changing climate may reduce their ability to balance their needs and lead to continued population declines. However, several management approaches could promote resiliency of alpine habitats that support Dall's sheep populations.