Migration is a striking behavioral strategy by which many animals enhance resource acquisition while reducing predation risk. Historically, the demographic benefits of such movements made migration common, but in many taxa the phenomenon is considered globally threatened. Here we describe a long-term decline in the productivity of elk (Cervus elaphus) that migrate through intact wilderness areas to protected summer ranges inside Yellowstone National Park, USA. We attribute this decline to a long-term reduction in the demographic benefits that ungulates typically gain from migration. Among migratory elk, we observed a 21-year, 70% reduction in recruitment and a 4-year, 19% depression in their pregnancy rate largely caused by infrequent reproduction of females that were young or lactating. In contrast, among resident elk, we have recently observed increasing recruitment and a high rate of pregnancy. Landscape-level changes in habitat quality and predation appear to be responsible for the declining productivity of Yellowstone migrants. From 1989 to 2009, migratory elk experienced an increasing rate and shorter duration of green-up coincident with warmer spring-summer temperatures and reduced spring precipitation, also consistent with observations of an unusually severe drought in the region. Migrants are also now exposed to four times as many grizzly bears (Ursus arctos) and wolves (Canis lupus) as resident elk. Both of these restored predators consume migratory elk calves at high rates in the Yellowstone wilderness but are maintained at low densities via lethal management and human disturbance in the year-round habitats of resident elk. Our findings suggest that large-carnivore recovery and drought, operating simultaneously along an elevation gradient, have disproportionately influenced the demography of migratory elk. Many migratory animals travel large geographic distances between their seasonal ranges. Changes in land use and climate that disparately influence such seasonal ranges may alter the ecological basis of migratory behavior, representing an important challenge for, and a powerful lens into, the ecology and conservation of migratory taxa.

Identifying the ecological dynamics underlying human-wildlife conflicts is important for the management and conservation of wildlife populations. In landscapes still occupied by large carnivores, many ungulate prey species migrate seasonally, yet little empirical research has explored the relationship between carnivore distribution and ungulate migration strategy. In this study, we evaluate the influence of elk (Cervus elaphus) distribution and other landscape features on wolf (Canis lupus) habitat use in an area of chronic wolf-livestock conflict in the Greater Yellowstone Ecosystem, USA. Using three years of fine-scale wolf (n = 14) and elk (n = 81) movement data, we compared the seasonal habitat use of wolves in an area dominated by migratory elk with that of wolves in an adjacent area dominated by resident elk. Most migratory elk vacate the associated winter wolf territories each summer via a 40-60 km migration, whereas resident elk remain accessible to wolves year-round. We used a generalized linear model to compare the relative probability of wolf use as a function of GIS-based habitat covariates in the migratory and resident elk areas. Although wolves in both areas used elk-rich habitat all year, elk density in summer had a weaker influence on the habitat use of wolves in the migratory elk area than the resident elk area. Wolves employed a number of alternative strategies to cope with the departure of migratory elk. Wolves in the two areas also differed in their disposition toward roads. In winter, wolves in the migratory elk area used habitat close to roads, while wolves in the resident elk area avoided roads. In summer, wolves in the migratory elk area were indifferent to roads, while wolves in resident elk areas strongly avoided roads, presumably due to the location of dens and summering elk combined with different traffic levels. Study results can help wildlife managers to anticipate the movements and establishment of wolf packs as they expand into areas with migratory or resident prey populations, varying levels of human activity, and front-country rangelands with potential for conflicts with livestock.

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As an outcome of natural selection, animals are probably adapted to select territories economically by maximizing benefits and minimizing costs of territory ownership. Theory and empirical precedent indicate that a primary benefit of many territories is exclusive access to food resources, and primary costs of defending and using space are associated with competition, travel and mortality risk. A recently developed mechanistic model for economical territory selection provided numerous empirically testable predictions. We tested these predictions using location data from grey wolves ( Canis lupus ) in Montana, USA. As predicted, territories were smaller in areas with greater densities of prey, competitors and low-use roads, and for groups of greater size. Territory size increased before decreasing curvilinearly with greater terrain ruggedness and harvest mortalities. Our study provides evidence for the economical selection of territories as a causal mechanism underlying ecological patterns observed in a cooperative carnivore. Results demonstrate how a wide range of environmental and social conditions will influence economical behaviour and resulting space use. We expect similar responses would be observed in numerous territorial species. A mechanistic approach enables understanding how and why animals select particular territories. This knowledge can be used to enhance conservation efforts and more successfully predict effects of conservation actions.

Little research has evaluated how the migration and distribution of native prey influence patterns of livestock depredation by large carnivores. Previous research suggests that the presence of native prey can increase depredation rates by attracting predators (prey tracking hypothesis). Alternatively, the absence of native prey may facilitate predation on livestock (prey scarcity hypothesis). In this study, we evaluated support for these competing hypotheses through analysis of 4 years of cattle (Bos taurus L., 1758) depredation data (n = 39 kills), 2 years of summer and fall wolf (Canis lupus L., 1758) predation and tracking data (n = 4 wolves), and 3 years of elk (Cervus elaphus L., 1758) movement data (n = 70 elk). We used logistic regression to compare the relative influence of landscape features and elk distribution on the risk of livestock depredation in areas with migratory and resident elk. Cattle depredations occurred in habitats with increased encounter rates between wolves and livestock. In resident elk areas, depredation sites were associated with elk distribution and open roads. In migratory elk areas, depredation sites were associated with wolf dens, streams, and open habitat. Patterns of carnivore–livestock conflicts are complex, and using ungulate distribution data can predict and minimize such instances.

Wolf (Canis lupus) depredations of livestock are a ubiquitous source of conflict in every country where wolves and livestock overlap. We studied the spatial and temporal variation of wolf depredations of livestock in Montana during 2005-2015, including evaluations of targeted control efforts and public harvest as potential means to reduce depredations. During this time we collected spatial data for all confirmed wolflivestock depredations, tallied the annual number of depredation events within hunting districts, and collected data for variables potentially predictive of depredation events. We decomposed variation in depredation data into 2 distinct components: the binary presence or absence of depredation events in each district-year, and the count of depredation events in district-years with !1 event. We found that presenceabsence of depredations increased with wolf presence and wolf density, increased with livestock density, were highest at intermediate proportionate areas of agricultural land, and were a recurrent phenomenon such that districts with depredations the previous year were more likely to continue having them. Targeted removal, but not public harvest, significantly reduced the recurrent presence of depredations. The number of conflicts in district-years with !1 depredation event was positively correlated with wolf density, cattle density, intermediate proportionate areas of forested land, and the number of events during the previous year. Public harvest reduced the counts of depredation events in areas where conflict reoccurred, though with a modest predicted effect size of 0.22 fewer depredations/district-year, or 5.7 fewer depredation events statewide/year (8% of the annual average). Minimizing livestock losses is a top priority for wolf management. These results shed light on the broad-scale patterns behind chronic problems and the effectiveness of wolf management practices in addressing them. Ó

Population sizes and movement patterns of ungulate grazers and their predators have fluctuated dramatically over the past few centuries, largely owing to overharvesting, land-use change and historic management. We used d C and d15 N values measured from bone collagen of historic and recent gray wolves and their potential primary prey from Yellowstone National Park to gain insight into the trophic dynamics and nutrient conditions of historic and modern grasslands. The diet of reintroduced wolves closely parallels that of the historic population. We suggest that a significant shift in faunal d15 N values over the past century reflects impacts of anthropogenic environmental changes on grassland ecosystems, including grazer-mediated shifts in grassland nitrogen cycle processes.

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