Michael D. Jimenez

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.

Habitat Selection by Recolonizing Wolves in the Northern Rocky Mountains of the United States

Abstract Gray wolf (Canis lupus) populations have persisted and expanded in northwest Montana since 1986, while reintroduction efforts in Idaho and Yellowstone have further bolstered the regional population. However, rigorous analysis of either the availability of wolf habitat in the entire region, or the specific habitat requirements of local wolves, has yet to be conducted. We examined wolf-habitat relationships in the northern Rocky Mountains of the U.S. by relating landscape/habitat features found within wolf pack home ranges (n = 56) to those found in adjacent non-occupied areas (n = 56). Logistic regression revealed that increased forest cover, lower human population density, higher elk density, and lower sheep density were the primary factors related to wolf occupation. Similar factors promoted wolf pack persistence. Further, our analysis indicated that relatively large tracts of suitable habitat remain unoccupied in the Rocky Mountains, suggesting that wolf populations likely will continue to increase in the region. Analysis of the habitat linkage between the 3 main wolf recovery areas indicates that populations in central Idaho and northwest Montana have higher connectivity than either of the 2 recovery areas to the Greater Yellowstone recovery area. Thus, for the northern Rocky Mountains to function as a metapopulation for wolves, it will be necessary that dispersal corridors to the Yellowstone ecosystem be established and conserved.

Survival of Colonizing Wolves in the Northern Rocky Mountains of the United States, 1982-2004

Abstract After roughly a 60-year absence, wolves (Canis lupus) immigrated (1979) and were reintroduced (1995–1996) into the northern Rocky Mountains (NRM), USA, where wolves are protected under the Endangered Species Act. The wolf recovery goal is to restore an equitably distributed metapopulation of ≥30 breeding pairs and 300 wolves in Montana, Idaho, and Wyoming, while minimizing damage to livestock; ultimately, the objective is to establish state-managed conservation programs for wolf populations in NRM. Previously, wolves were eradicated from the NRM because of excessive human killing. We used Andersen–Gill hazard models to assess biological, habitat, and anthropogenic factors contributing to current wolf mortality risk and whether federal protection was adequate to provide acceptably low hazards. We radiocollared 711 wolves in Idaho, Montana, and Wyoming (e.g., NRM region of the United States) from 1982 to 2004 and recorded 363 mortalities. Overall, annual survival rate of wolves in the recovery areas was 0.750 (95% CI  =  0.728–0.772), which is generally considered adequate for wolf population sustainability and thereby allowed the NRM wolf population to increase. Contrary to our prediction, wolf mortality risk was higher in the northwest Montana (NWMT) recovery area, likely due to less abundant public land being secure wolf habitat compared to other recovery areas. In contrast, lower hazards in the Greater Yellowstone Area (GYA) and central Idaho (CID) likely were due to larger core areas that offered stronger wolf protection. We also found that wolves collared for damage management purposes (targeted sample) had substantially lower survival than those collared for monitoring purposes (representative sample) because most mortality was due to human factors (e.g., illegal take, control). This difference in survival underscores the importance of human-caused mortality in this recovering NRM population. Other factors contributing to increased mortality risk were pup and yearling age class, or dispersing status, which was related to younger age cohorts. When we included habitat variables in our analysis, we found that wolves having abundant agricultural and private land as well as livestock in their territory had higher mortality risk. Wolf survival was higher in areas with increased wolf density, implying that secure core habitat, particularly in GYA and CID, is important for wolf protection. We failed to detect changes in wolf hazards according to either gender or season. Maintaining wolves in NWMT will require greater attention to human harvest, conflict resolution, and illegal mortality than in either CID or GYA; however, if human access increases in the future in either of the latter 2 areas hazards to wolves also may increase. Indeed, because overall suitable habitat is more fragmented and the NRM has higher human access than many places where wolves roam freely and are subject to harvest (e.g., Canada and AK), monitoring of wolf vital rates, along with concomitant conservation and management strategies directed at wolves, their habitat, and humans, will be important for ensuring long-term viability of wolves in the region.

The Effects of Breeder Loss on Wolves

Abstract Managers of recovering wolf (Canis lupus) populations require knowledge regarding the potential impacts caused by the loss of territorial, breeding wolves when devising plans that aim to balance population goals with human concerns. Although ecologists have studied wolves extensively, we lack an understanding of this phenomenon as published records are sparse. Therefore, we pooled data (n = 134 cases) on 148 territorial breeding wolves (75 M and 73 F) from our research and published accounts to assess the impacts of breeder loss on wolf pup survival, reproduction, and territorial social groups. In 58 of 71 cases (84%), ≥1 pup survived, and the number or sex of remaining breeders (including multiple breeders) did not influence pup survival. Pups survived more frequently in groups of ≥6 wolves (90%) compared with smaller groups (68%). Auxiliary nonbreeders benefited pup survival, with pups surviving in 92% of cases where auxiliaries were present and 64% where they were absent. Logistic regression analysis indicated that the number of adult-sized wolves remaining after breeder loss, along with pup age, had the greatest influence on pup survival. Territorial wolves reproduced the following season in 47% of cases, and a greater proportion reproduced where one breeder had to be replaced (56%) versus cases where both breeders had to be replaced (9%). Group size was greater for wolves that reproduced the following season compared with those that did not reproduce. Large recolonizing (>75 wolves) and saturated wolf populations had similar times to breeder replacement and next reproduction, which was about half that for small recolonizing (≤75 wolves) populations. We found inverse relationships between recolonizing population size and time to breeder replacement (r = −0.37) and time to next reproduction (r = −0.36). Time to breeder replacement correlated strongly with time to next reproduction (r = 0.97). Wolf social groups dissolved and abandoned their territories subsequent to breeder loss in 38% of cases. Where groups dissolved, wolves reestablished territories in 53% of cases, and neighboring wolves usurped territories in an additional 21% of cases. Fewer groups dissolved where breeders remained (26%) versus cases where breeders were absent (85%). Group size after breeder loss was smaller where groups dissolved versus cases where groups did not dissolve. To minimize negative impacts, we recommend that managers of recolonizing wolf populations limit lethal control to solitary individuals or territorial pairs where possible, because selective removal of pack members can be difficult. When reproductive packs are to be managed, we recommend that managers only remove wolves from reproductive packs when pups are ≥6 months old and packs contain ≥6 members (including ≥3 ad-sized wolves). Ideally, such packs should be close to neighboring packs and occur within larger (≥75 wolves) recolonizing populations.

People and Wildlife: Managing wolf–human conflict in the northwestern United States

INTRODUCTION The grey wolf ( Canis lupus ) is the most widely distributed large carnivore in the northern hemisphere (Nowak 1995) and has a reputation for killing livestock and competing with human hunters for wild ungulates (Young 1944; Fritts et al . 2003). Wolves rarely threaten human safety, but many people still fear them. In the western USA, widespread extirpation of ungulates by colonizing settlers, wolf depredation on livestock and negative public attitudes towards wolves resulted in extirpation of wolf populations by 1930 (Mech 1970; McIntyre 1995). By 1970, mule deer ( Odocoileus hemionus ), white-tailed deer ( O. virginianus ), elk ( Cervus elaphus ), moose ( Alces alces ) and bighorn sheep ( Ovis canadensis ) populations had been restored throughout the western USA while bison ( Bison bison ) were recovered only in Yellowstone National Park. However, grey wolves were still persecuted. In 1974, grey wolves were protected and managed by the US Fish and Wildlife Service under the federal Endangered Species Act of 1973. In 1986, the first recorded den in the western USA in over 50 years was established in Glacier National Park by wolves that naturally dispersed from Canada (Ream et al . 1989). Restoration of wolves in that region emphasized legal protection and building local public tolerance. Wolves from Canada were reintroduced to central Idaho and Yellowstone National Park in 1995 and 1996 to accelerate restoration (Bangs and Fritts 1996; Fritts et al . 1997). The Northern Rocky Mountains wolf population grew from 10 wolves in 1987 to 663 wolves by 2003 (US Fish and Wildlife Service et al . 2003) (Fig. 21.1, Table 21.1).

Estimation of Successful Breeding Pairs for Wolves in the Northern Rocky Mountains, USA

Abstract Under the Endangered Species Act, documenting recovery and federally mandated population levels of wolves (Canis lupus) in the Northern Rocky Mountains (NRM) requires monitoring wolf packs that successfully recruit young. United States Fish and Wildlife Service regulations define successful breeding pairs as packs estimated to contain an adult male and female, accompanied by ≥2 pups on 31 December of a given year. Monitoring successful breeding pairs will become more difficult following proposed delisting of NRM wolves; alternatives to historically intensive methods, appropriate to the different ecological and regulatory context following delisting, are required. Because pack size is easier to monitor than pack composition, we estimated probability a pack would contain a successful breeding pair based on its size for wolf populations inhabiting 6 areas in the NRM. We also evaluated the extent to which differences in demography of wolves and levels of human-caused mortality among the areas influenced the probability of packs of different sizes would contain successful breeding pairs. Probability curves differed among analysis areas, depending primarily on levels of human-caused mortality, secondarily on annual population growth rate, and little on annual population density. Probabilities that packs contained successful breeding pairs were more uniformly distributed across pack sizes in areas with low levels of human mortality and stable populations. Large packs in areas with high levels of human-caused mortality and high annual growth rates had relatively high probabilities of containing breeding pairs whereas those for small packs were relatively low. Our approach can be used by managers to estimate number of successful breeding pairs in a population where number of packs and their sizes are known. Following delisting of NRM wolves, human-caused mortality is likely to increase, resulting in more small packs with low probabilities of containing breeding pairs. Differing contributions of packs to wolf population growth based on their size suggests monitoring successful breeding pairs will provide more accurate insights into population dynamics of wolves than will monitoring number of packs or individuals only.

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.

Successful rearing of young by wild wolves without mates

Two lone-female wolves ( Canis lupus ) and a lone-male wolf each successfully raised litters of young in northwestern Montana and southeastern British Columbia. This unusual reproductive behavior occurred in a low-density population of wolves colonizing an area containing a relatively dense population of ungulates.