Hidden Markov movement models reveal diverse seasonal movement patterns in two North American ungulates.

Animal movement is the mechanism connecting landscapes to fitness, and understanding variation in seasonal animal movements has benefited from the analysis and categorization of animal displacement. However, seasonal movement patterns can defy classification when movements are highly variable. Hidden Markov movement models (HMMs) are a class of latent-state models well-suited to modeling movement data. Here, we used HMMs to assess seasonal patterns of variation in the movement of pronghorn (Antilocapra americana), a species known for variable seasonal movements that challenge analytical approaches, while using a population of mule deer (Odocoileus hemionus), for whom seasonal movements are well-documented, as a comparison. We used population-level HMMs in a Bayesian framework to estimate a seasonal trend in the daily probability of transitioning between a short-distance local movement state and a long-distance movement state. The estimated seasonal patterns of movements in mule deer closely aligned with prior work based on indices of animal displacement: a short period of long-distance movements in the fall season and again in the spring, consistent with migrations to and from seasonal ranges. We found seasonal movement patterns for pronghorn were more variable, as a period of long-distance movements in the fall was followed by a winter period in which pronghorn were much more likely to further initiate and remain in a long-distance movement pattern compared with the movement patterns of mule deer. Overall, pronghorn were simply more likely to be in a long-distance movement pattern throughout the year. Hidden Markov movement models provide inference on seasonal movements similar to other methods, while providing a robust framework to understand movement patterns on shorter timescales and for more challenging movement patterns. Hidden Markov movement models can allow a rigorous assessment of the drivers of changes in movement patterns such as extreme weather events and land development, important for management and conservation.

Wind-energy development alters pronghorn migration at multiple scales.

Migration is a critical behavioral strategy necessary for population persistence and ecosystem functioning, but migration routes have been increasingly disrupted by anthropogenic activities, including energy development. Wind energy is the world's fastest growing source of electricity and represents an important alternative to hydrocarbon extraction, but its effects on migratory species beyond birds and bats are not well understood. We evaluated the effects of wind-energy development on pronghorn migration, including behavior and habitat selection, to assess potential effects on connectivity and other functional benefits including stopovers. We monitored GPS-collared female pronghorn from 2010 to 2012 and 2018 to 2020 in south-central Wyoming, USA, an area with multiple wind-energy facilities in various stages of development and operation. Across all time periods, we collected 286 migration sequences from 117 individuals, including 121 spring migrations, 123 fall migrations, and 42 facultative winter migrations. While individuals continued to migrate through wind-energy facilities, pronghorn made important behavioral adjustments relative to turbines during migration. These included avoiding turbines when selecting stopover sites in spring and winter, selecting areas farther from turbines at a small scale in spring and winter, moving more quickly near turbines in spring (although pronghorn moved more slowly near turbines in the fall), and reducing fidelity to migration routes relative to wind turbines under construction in both spring and fall. For example, an increase in distance to turbine from 0 to 1 km translated to a 33% and 300% increase in the relative probability of selection for stopover sites in spring and winter, respectively. The behavioral adjustments pronghorn made relative to wind turbines could reduce the functional benefits of their migration, such as foraging success or the availability of specific routes, over the long term.

Eyes on the herd: Quantifying ungulate density from satellite, unmanned aerial systems, and GPScollar data.

Novel approaches to quantifying density and distributions could help biologists adaptively manage wildlife populations, particularly if methods are accurate, consistent, cost-effective, rapid, and sensitive to change. Such approaches may also improve research on interactions between density and processes of interest, such as disease transmission across multiple populations. We assess how satellite imagery, unmanned aerial system (UAS) imagery, and Global Positioning System (GPS) collar data vary in characterizing elk density, distribution, and count patterns across times with and without supplemental feeding at the National Elk Refuge (NER) in the US state of Wyoming. We also present the first comparison of satellite imagery data with traditional counts for ungulates in a temperate system. We further evaluate seven different aggregation metrics to identify the most consistent and sensitive metrics for comparing density and distribution across time and populations. All three data sources detected higher densities and aggregation locations of elk during supplemental feeding than non-feeding at the NER. Kernel density estimates (KDEs), KDE polygon areas, and the first quantile of interelk distances detected differences with the highest sensitivity and were most highly correlated across data sources. Both UAS and satellite imagery provide snapshots of density and distribution patterns of most animals in the area at lower cost than GPS collars. While satellite-based counts were lower than traditional counts, aggregation metrics matched those from UAS and GPS data sources when animals appeared in high contrast to the landscape, including brown elk against new snow in open areas. UAS counts of elk were similar to traditional ground-based counts on feed grounds and are the best data source for assessing changes in small spatial extents. Satellite, UAS, or GPS data can provide appropriate data for assessing density and changes in density from adaptive management actions. For the NER, where high elk densities are beneath controlled airspace, GPS collar data will be most useful for evaluating how management actions, including changes in the dates of supplemental feeding, influence elk density and aggregation across large spatial extents. Using consistent and sensitive measures of density may improve research on the drivers and effects of density within and across a wide range of species.

Freezing in a warming climate: Marked declines of a subnivean hibernator after a snow drought.

Recent snow droughts associated with unusually warm winters are predicted to increase in frequency and affect species dependent upon snowpack for winter survival. Changes in populations of some cold-adapted species have been attributed to heat stress or indirect effects on habitat from unusually warm summers, but little is known about the importance of winter weather to population dynamics and how responses to snow drought vary among sympatric species. We evaluated changes in abundance of hoary marmots (Marmota caligata) over a period that included a year of record-low snowpack to identify mechanisms associated with weather and snowpack. To consider interspecies comparisons, our analysis used the same a priori model set as a concurrent study that evaluated responses of American pikas (Ochotona princeps) to weather and snowpack in the same study area of North Cascades National Park, Washington, USA. We hypothesized that marmot abundance reflected mechanisms related to heat stress, cold stress, cold exposure without an insulating snowpack, snowpack duration, atmospheric moisture, growing-season precipitation, or select combinations of these mechanisms. Changes in marmot abundances included a 74% decline from 2007 to 2016 and were best explained by an interaction of chronic dryness with exposure to acute cold without snowpack in winter. Physiological stress during hibernation from exposure to cold, dry air appeared to be the most likely mechanism of change in marmot abundance. Alternative mechanisms associated with changes to winter weather, including early emergence from hibernation or altered vegetation dynamics, had less support. A post hoc assessment of vegetative phenology and productivity did not support vegetation dynamics as a primary driver of marmot abundance across years. Although marmot and pika abundances were explained by strikingly similar models over periods of many years, details of the mechanisms involved likely differ between species because pika abundances increased in areas where marmots declined. Such differences may lead to diverging geographic distributions of these species as global change continues.

Ecological consequences of anomalies in atmospheric moisture and snowpack.

Although increased frequency of extreme-weather events is one of the most secure predictions associated with contemporary climate change, effects of such events on distribution and abundance of climate-sensitive species remain poorly understood. Montane ecosystems may be especially sensitive to extreme weather because of complex abiotic and biotic interactions that propagate from climate-driven reductions in snowpack. Snowpack not only protects subnivean biotas from extreme cold, but also influences forage availability through timing of melt-off and water availability. We related relative abundances of an alpine mammal, the American pika (Ochotona princeps), to measures of weather and snowpack dynamics over an 8-yr period that included before and after a year of record-low snowpack in Washington, USA. We sought to (1) quantify any change in pika abundance associated with the snowpack anomaly and (2) identify aspects of weather and snowpack that influenced abundance of pikas. Pikas showed a 1-yr lag response to the snowpack anomaly and exhibited marked declines in abundance at elevations below 1,400 m simultaneous with increased abundances at higher elevations. Atmospheric moisture, indexed by vapor pressure deficit (VPD), was especially important, evidenced by strong support for the top-ranked model that included the interaction of VPD with snowpack duration. Notably, our novel application of VPD from gridded climate data for analyses of animal abundances shows strong potential for improving species distribution models because VPD represents an important aspect of weather that influences the physiology and habitat of biota. Pikas were apparently affected by cold stress without snowpack at mid elevations, whereas changes to forage associated with snowpack and VPD were influential at high and low elevations. Our results reveal context dependency in pika responses to weather and illustrate how snow drought can lead to rapid change in the abundance of subnivean animals.