Spatial and temporal variability in summer diet of gray wolves (Canis lupus) in the Greater Yellowstone Ecosystem.

The role of predation by large carnivores in suppressing prey populations and structuring ecosystems is highly debated, calling for a detailed understanding of carnivore diets. Wolves (Canis lupus) roam across three continents and persist throughout widely different ecosystems. Their diet is flexible and may vary spatially as well as seasonally, which requires analysis of diet on different spatial and temporal scales. Few studies have investigated the summer diet of wolves, which is more variable, consists of smaller prey, and requires different methods than studying their winter diet. To better understand the summer diet of wolves, we combined three independently collected wolf scat data sets from three distinctly different portions of the Greater Yellowstone Ecosystem: Yellowstone National Park (2009), Grand Teton National Park (2003 - 2009), and the Absaroka-Beartooth Wilderness (2009 - 2010). These areas represent different ecological conditions and management regimes, which may impact wolf diet. We estimated relative biomass and compared occurrence of different prey species among packs, years, as well as the three regions. In total, we analyzed 1,906 wolf scats and found that neonate cervids, adult elk, and adult deer were the most important prey species in the summer diet of the wolves. We found dietary variation among packs residing in the same area, as well as across years. The occurrence of neonate cervids displayed the most variation, and low occurrence of this prey type often was associated with a more diverse diet. Wolf packs within the national parks had a higher occurrence of medium-sized prey (~ 50 - 70 kg) and lower occurrence of small-sized prey (≤ 20 kg) compared to wolves in the Absaroka-Beartooth Wilderness. These results demonstrate flexibility in summer diet across packs, years, and between regions within the Greater Yellowstone Ecosystem.

Animal migration amid shifting patterns of phenology and predation: lessons from a Yellowstone elk herd.

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.

Linking anti-predator behaviour to prey demography reveals limited risk effects of an actively hunting large carnivore.

Ecological theory predicts that the diffuse risk cues generated by wide-ranging, active predators should induce prey behavioural responses but not major, population- or community-level consequences. We evaluated the non-consumptive effects (NCEs) of an active predator, the grey wolf (Canis lupus), by simultaneously tracking wolves and the behaviour, body fat, and pregnancy of elk (Cervus elaphus), their primary prey in the Greater Yellowstone Ecosystem. When wolves approached within 1 km, elk increased their rates of movement, displacement and vigilance. Even in high-risk areas, however, these encounters occurred only once every 9 days. Ultimately, despite 20-fold variation in the frequency of encounters between wolves and individual elk, the risk of predation was not associated with elk body fat or pregnancy. Our findings suggest that the ecological consequences of actively hunting large carnivores, such as the wolf, are more likely transmitted by consumptive effects on prey survival than NCEs on prey behaviour.

Elk migration patterns and human activity influence wolf habitat use in the Greater Yellowstone Ecosystem.

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.

Sarcoptic mange found in wolves in the Rocky Mountains in western United States.

We documented sarcoptic mange caused by mites (Sarcoptes scabiei) in 22 gray wolves (Canis lupus) in the northern Rocky Mountain states of Montana (n=16) and Wyoming (n=6), from 2002 through 2008. To our knowledge, this is the first report of sarcoptic mange in wolves in Montana or Wyoming in recent times. In addition to confirming sarcoptic mange, we recorded field observations of 40 wolves in Montana and 30 wolves in Wyoming displaying clinical signs of mange (i.e., alopecia, hyperkeratosis, and seborrhea). Therefore, we suspect sarcoptic mange may be more prevalent than we were able to confirm.