An apex carnivore's life history mediates a predator cascade.

Apex predators can shape communities via cascading top-down effects, but the degree to which such effects depend on predator life history traits is largely unknown. Within carnivore guilds, complex hierarchies of dominance facilitate coexistence, whereby subordinate species avoid dominant counterparts by partitioning space, time, or both. We investigated whether a major life history trait (hibernation) in an apex carnivore (black bears Ursus americanus) mediated its top-down effects on the spatio-temporal dynamics of three sympatric mesocarnivore species (coyotes Canis latrans, bobcats Lynx rufus, and gray foxes Urocyon cinereoargenteus) across a 15,000 km2 landscape in the western USA. We compared top-down, bottom-up, and environmental effects on these mesocarnivores using an integrated modeling approach. Black bears exerted top-down effects that varied as a function of hibernation and were stronger than bottom-up or environmental impacts. High black bear activity in summer and fall appeared to buffer the most subordinate mesocarnivore (gray foxes) from competition with dominant mesocarnivores (coyotes and bobcats), which were in turn released by black bear hibernation in winter and early spring. The mesocarnivore responses occurred in space (i.e., altered occupancy and site visitation intensity) rather than time (i.e., diel activity patterns unaffected). These results suggest that the spatio-temporal dynamics of mesocarnivores in this system were principally shaped by a spatial predator cascade of interference competition mediated by black bear hibernation. Thus, certain life history traits of apex predators might facilitate coexistence among competing species over broad time scales, with complex implications for lower trophic levels.

Plasticity in elk migration timing is a response to changing environmental conditions.

Migration is an effective behavioral strategy for prolonging access to seasonal resources and may be a resilient strategy for ungulates experiencing changing climatic conditions. In the Greater Yellowstone Ecosystem (GYE), elk are the primary ungulate, with approximately 20,000 individuals migrating to exploit seasonal gradients in forage while also avoiding energetically costly snow conditions. How climate-induced changes in plant phenology and snow accumulation are influencing elk migration timing is unknown. We present the most complete record of elk migration across the GYE, spanning 9 herds and 414 individuals from 2001 to 2017, to evaluate the drivers of migration timing and test for temporal shifts. The timing of elk departure from winter range involved a trade-off between current and anticipated forage conditions, while snow melt governed summer range arrival date. Timing of elk departure from summer range and arrival on winter range were both influenced by snow accumulation and exposure to hunting. At the GYE scale, spring and fall migration timing changed through time, most notably with winter range arrival dates becoming almost 50 days later since 2001. Predicted herd-level changes in migration timing largely agreed with observed GYE-wide changes-except for predicted winter range arrival dates which did not reflect the magnitude of change detected in the elk telemetry data. Snow melt, snow accumulation, and spring green-up dates all changed through time, with different herds experiencing different rates and directions of change. We conclude that elk migration is plastic, is a direct response to environmental cues, and that these environmental cues are not changing in a consistent manner across the GYE. The impacts of changing elk migration timing on predator-prey dynamics, carnivore-livestock conflict, disease ecology, and harvest management across the GYE are likely to be significant and complex.

A Retrospective Assessment of a Failed Collaborative Process in Conservation.

Collaboration provides one tool for managing the complicated and often the contentious natural resource issues. Successful collaborative arrangements involve a mix of actors bringing key attributes to the table: power, capacity, motivation, mandate, and synergy. These attributes, if missing or if one overshadows the rest, can derail the collaborative process and/or the conservation outcomes. We offer a case study of natural gas field development impacts on America's only endemic ungulate-pronghorn (Antilocapra americana)-winter range in the Upper Green River Basin (UGRB), Wyoming, USA. We illustrate how a collaborative process can go awry, given asymmetries between the relative strengths and the associated attributes of actors, and the subsequent extent to which this imbalance created an unfavorable situation for continued collaboration. The case study reveals disagreements on technical data and potential insight on agency capture operating at a local scale. Despite these process challenges, some conservation outcomes resulted from work generated by the collaboration. Our experience underscores the importance of defining a clear purpose for collaborative processes at the outset, articulating specific roles, ensuring transparency among actors, and flexibility for long-term management as possible ways, in which the groups involved in collaborations to manage natural resources can complement each other's strengths and strive for better conservation outcomes.

Identifying impediments to long-distance mammal migrations.

In much of the world, the persistence of long-distance migrations by mammals is threatened by development. Even where human population density is relatively low, there are roads, fencing, and energy development that present barriers to animal movement. If we are to conserve species that rely on long-distance migration, then it is critical that we identify existing migration impediments. To delineate stopover sites associated with anthropogenic development, we applied Brownian bridge movement models to high-frequency locations of pronghorn (Antilocapra americana) in the Greater Yellowstone Ecosystem. We then used resource utilization functions to assess the threats to long-distance migration of pronghorn that were due to fences and highways. Migrating pronghorn avoided dense developments of natural gas fields. Highways with relatively high volumes of traffic and woven-wire sheep fence acted as complete barriers. At crossings with known migration bottlenecks, use of high-quality forage and shrub habitat by pronghorn as they approached the highway was lower than expected based on availability of those resources. In contrast, pronghorn consistently utilized high-quality forage close to the highway at crossings with no known migration bottlenecks. Our findings demonstrate the importance of minimizing development in migration corridors in the future and of mitigating existing pressure on migratory animals by removing barriers, reducing the development footprint, or installing crossing structures.

Selection of den sites and chronology of denning by black bears in the eastern Sierra Nevada and western Great Basin.

Use of dens during winter is an important strategy for American black bears (Ursus americanus) for both energy conservation and reproduction; and occupancy of suitable den sites has implications for reproductive fitness. Denning strategies may change as a result of changing climatic conditions and habitat loss. Black bears occupy arid environments in the eastern Sierra Nevada and the western ranges of the Great Basin Ecosystem. Our objectives were to identify: (1) which physical characteristics of habitat influenced selection of den sites at multiple spatial scales and (2) which environmental factors influenced timing of entrance and exit of dens by females and males. We evaluated selection of den sites by black bears at three spatial scales (300, 1000, and 4000 m) from 2011 to 2022. Terrain ruggedness was important for selection of den sites at all spatial scales. Within a 300-m buffer from the den, bears selected den sites with rugged terrain, lower horizontal visibility, and greater canopy cover, resulting in more concealment and protection than that of the surrounding environment. Within 1000- and 4000-m buffers around each den, bears selected den sites with rugged terrain, northern aspects, and steep slopes. At the 4000-m scale, we observed interactions between sex with slope and distance to roads; females selected den sites on steeper slopes and closer to roads than did males. Females remained in the dens longer than males by entering earlier in the autumn and exiting later in the spring. Male bears exited their dens earlier with increasing consecutive days above freezing temperatures, but that relationship was weak for females. Knowing what characteristics are important for selection of den sites, and influence timing of denning, will be important for understanding how shifting climatic patterns will affect bears, particularly in arid environments that may be prone to wider fluctuations in climatic drivers of denning in the future.

Inferential consequences of modeling rather than measuring snow accumulation in studies of animal ecology.

It is increasingly common for studies of animal ecology to use model-based predictions of environmental variables as explanatory or predictor variables, even though model prediction uncertainty is typically unknown. To demonstrate the potential for misleading inferences when model predictions with error are used in place of direct measurements, we compared snow water equivalent (SWE) and snow depth as predicted by the Snow Data Assimilation System (SNODAS) to field measurements of SWE and snow depth. We examined locations on elk (Cervus canadensis) winter ranges in western Wyoming, because modeled data such as SNODAS output are often used for inferences on elk ecology. Overall, SNODAS predictions tended to overestimate field measurements, prediction uncertainty was high, and the difference between SNODAS predictions and field measurements was greater in snow shadows for both snow variables compared to non-snow shadow areas. We used a simple simulation of snow effects on the probability of an elk being killed by a predator to show that, if SNODAS prediction uncertainty was ignored, we might have mistakenly concluded that SWE was not an important factor in where elk were killed in predatory attacks during the winter. In this simulation, we were interested in the effects of snow at finer scales (< 1 km2) than the resolution of SNODAS. If bias were to decrease when SNODAS predictions are averaged over coarser scales, SNODAS would be applicable to population-level ecology studies. In our study, however, averaging predictions over moderate to broad spatial scales (9-2200 km2) did not reduce the differences between SNODAS predictions and field measurements. This study highlights the need to carefully evaluate two issues when using model output as an explanatory variable in subsequent analysis: (1) the model's resolution relative to the scale of the ecological question of interest and (2) the implications of prediction uncertainty on inferences when using model predictions as explanatory or predictor variables.

Identification of source-sink dynamics in mountain lions of the Great Basin.

Natural and anthropogenic boundaries have been shown to affect population dynamics and population structure for many species with movement patterns at the landscape level. Understanding population boundaries and movement rates in the field for species that are cryptic and occur at low densities is often extremely difficult and logistically prohibitive; however genetic techniques may offer insights that have previously been unattainable. We analysed thirteen microsatellite loci for 739 mountain lions (Puma concolor) using muscle tissue samples from individuals in the Great Basin throughout Nevada and the Sierra Nevada mountain range to test the hypothesis that heterogeneous hunting pressure results in source-sink dynamics at the landscape scale. We used a combination of non-spatial and spatial model-based Bayesian clustering methods to identify genetic populations. We then used a recently developed Bayesian multilocus genotyping method to estimate asymmetrical rates of contemporary movement between those subpopulations and to identify source and sink populations. We identified two populations at the highest level of genetic structuring with a total of five subpopulations in the Great Basin of Nevada and the Sierra Nevada range. Our results suggest that source-sink dynamics occur at landscape scales for wide-ranging species, such as mountain lions, and that source populations may be those that are under relatively less hunting pressure and that occupy refugia.

Recolonizing carnivores: Is cougar predation behaviorally mediated by bears?

Conservation and management efforts have resulted in population increases and range expansions for some apex predators, potentially changing trophic cascades and foraging behavior. Changes in sympatric carnivore and dominant scavenger populations provide opportunities to assess how carnivores affect one another. Cougars (Puma concolor) were the apex predator in the Great Basin of Nevada, USA, for over 80 years. Black bears (Ursus americanus) have recently recolonized the area and are known to heavily scavenge on cougar kills. To evaluate the impacts of sympatric, recolonizing bears on cougar foraging behavior in the Great Basin, we investigated kill sites of 31 cougars between 2009 and 2017 across a range of bear densities. We modeled the variation in feeding bout duration (number of nights spent feeding on a prey item) and the proportion of primary prey, mule deer (Odocoileus hemionus), in cougar diets using mixed-effects models. We found that feeding bout duration was driven primarily by the size of the prey item being consumed, local bear density, and the presence of dependent kittens. The proportion of mule deer in cougar diet across all study areas declined over time, was lower for male cougars, increased with the presence of dependent kittens, and increased with higher bear densities. In sites with feral horses (Equus ferus), a novel large prey, cougar consumption of feral horses increased over time. Our results suggest that higher bear densities over time may reduce cougar feeding bout durations and influence the prey selection trade-off for cougars when alternative, but more dangerous, large prey are available. Shifts in foraging behavior in multicarnivore systems can have cascading effects on prey selection. This study highlights the importance of measuring the impacts of sympatric apex predators and dominant scavengers on a shared resource base, providing a foundation for monitoring dynamic multipredator/scavenger systems.

Sexual predators, energy development, and conservation in greater Yellowstone.

In the United States, as elsewhere, a growing debate pits national energy policy and homeland security against biological conservation. In rural communities the extraction of fossil fuels is often encouraged because of the employment opportunities it offers, although the concomitant itinerant workforce is often associated with increased wildlife poaching. We explored possible positive and negative factors associated with energy extraction in the Greater Yellowstone Ecosystem (GYE), an area known for its national parks, intact biological diversity, and some of the New World's longest terrestrial migrations. Specifically, we asked whether counties with different economies-recreation (ski), agrarian (ranching or farming), and energy extractive (petroleum)-differed in healthcare (gauged by the abundance of hospital beds) and in the frequency of sexual predators. The absolute and relative frequency of registered sex offenders grew approximately two to three times faster in areas reliant on energy extraction. Healthcare among counties did not differ. The strong conflation of community dishevel, as reflected by in-migrant sexual predators, and ecological decay in Greater Yellowstone is consistent with patterns seen in similar systems from Ecuador to northern Canada, where social and environmental disarray exist around energy boomtowns. In our case, that groups (albeit with different aims) mobilized campaigns to help maintain the quality of rural livelihoods by protecting open space is a positive sign that conservation can matter, especially in the face of rampant and poorly executed energy extraction projects. Our findings further suggest that the public and industry need stronger regulatory action to instill greater vigilance when and where social factors and land conversion impact biological systems.

Using community photography to investigate phenology: A case study of coat molt in the mountain goat (Oreamnos americanus) with missing data.

Participatory approaches, such as community photography, can engage the public in questions of societal and scientific interest while helping advance understanding of ecological patterns and processes. We combined data extracted from community-sourced, spatially explicit photographs with research findings from 2018 fieldwork in the Yukon, Canada, to evaluate winter coat molt patterns and phenology in mountain goats (Oreamnos americanus), a cold-adapted, alpine mammal. Leveraging the community science portals iNaturalist and CitSci, in less than a year we amassed a database of almost seven hundred unique photographs spanning some 4,500 km between latitudes 37.6°N and 61.1°N from 0 to 4,333 m elevation. Using statistical methods accounting for incomplete data, a common issue in community science datasets, we identified the effects of intrinsic (sex and presence of offspring) and broad environmental (latitude and elevation) factors on molt onset and rate and compared our findings with published data. Shedding occurred over a 3-month period between 29 May and 6 September. Effects of sex and offspring on the timing of molt were consistent between the community-sourced and our Yukon data and with findings on wild mountain goats at a long-term research site in west-central Alberta, Canada. Males molted first, followed by females without offspring (4.4 days later in the coarse-grained, geographically wide community science sample; 29.2 days later in our fine-grained Yukon sample) and lastly females with new kids (6.2; 21.2 days later, respectively). Shedding was later at higher elevations and faster at northern latitudes. Our findings establish a basis for employing community photography to examine broad-scale questions about the timing of ecological events, as well as sex differences in response to possible climate drivers. In addition, community photography can help inspire public participation in environmental and outdoor activities specifically with reference to iconic wildlife.