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.     

Implications of Harvest on the Boundaries of Protected Areas for Large Carnivore Viewing Opportunities

The desire to see free ranging large carnivores in their natural habitat is a driver of tourism in protected areas around the globe. However, large carnivores are wide-ranging and subject to human-caused mortality outside protected area boundaries. The impact of harvest (trapping or hunting) on wildlife viewing opportunities has been the subject of intense debate and speculation, but quantitative analyses have been lacking. We examined the effect of legal harvest of wolves (Canis lupus) along the boundaries of two North American National Parks, Denali (DNPP) and Yellowstone (YNP), on wolf viewing opportunities within the parks during peak tourist season. We used data on wolf sightings, pack sizes, den site locations, and harvest adjacent to DNPP from 1997–2013 and YNP from 2008–2013 to evaluate the relationship between harvest and wolf viewing opportunities. Although sightings were largely driven by wolf population size and proximity of den sites to roads, sightings in both parks were significantly reduced by harvest. Sightings in YNP increased by 45% following years with no harvest of a wolf from a pack, and sightings in DNPP were more than twice as likely during a period with a harvest buffer zone than in years without the buffer. These findings show that harvest of wolves adjacent to protected areas can reduce sightings within those areas despite minimal impacts on the size of protected wolf populations. Consumptive use of carnivores adjacent to protected areas may therefore reduce their potential for non-consumptive use, and these tradeoffs should be considered when developing regional wildlife management policies.     

Meta-Analysis of Relationships between Human Offtake,Total Mortality and Population Dynamics of Gray Wolves (Canis lupus)

Following the growth and geographic expansion of wolf (Canis lupus) populations reintroduced to Yellowstone National Park and central Idaho in 1995–1996, Rocky Mountain wolves were removed from the endangered species list in May 2009. Idaho and Montana immediately established hunting seasons with quotas equaling 20% of the regional wolf population. Combining hunting with predator control, 37.1% of Montana and Idaho wolves were killed in the year of delisting. Hunting and predator control are well-established methods to broaden societal acceptance of large carnivores, but it is unprecedented for a species to move so rapidly from protection under the Endangered Species Act to heavy direct harvest, and it is important to use all available data to assess the likely consequences of these changes in policy. For wolves, it is widely argued that human offtake has little effect on total mortality rates, so that a harvest of 28–50% per year can be sustained. Using previously published data from 21 North American wolf populations, we related total annual mortality and population growth to annual human offtake. Contrary to current conventional wisdom, there was a strong association between human offtake and total mortality rates across North American wolf populations. Human offtake was associated with a strongly additive or super-additive increase in total mortality. Population growth declined as human offtake increased, even at low rates of offtake. Finally, wolf populations declined with harvests substantially lower than the thresholds identified in current state and federal policies. These results should help to inform management of Rocky Mountain wolves.     

Habitat selection by wolvesCanis lupus in the uplands and mountains of southern Poland

Using data from the National Wolf Census, carried out in Poland in 2000–2001, and GIS techniques we analysed habitat selection by wolvesCanis lupus Linnaeus, 1758 in uplands and mountains of southern Poland. We compared ten habitat variables and two parameters related to wolf abundance in 52 circular plots (154 km² each) with recorded wolves and 97 randomly selected plots with no signs of wolf presence. Wolf plots were characterized by higher elevation and closer location to the state border than wolf-free plots. Furthermore, wolf plots had higher forest cover, but smaller number of villages and towns and shorter railways and roads than plots without wolves. The best model explaining wolf distribution included forest cover, number of villages, length of roads and railway lines. Compared to northern Poland, the southern part of the country offers worse habitats for wolves due to significantly denser network of settlements and transportation routes.     

Scavenging Makes Cougars Susceptible to Snaring at Wolf Bait Stations

Abstract: In western Canada it is illegal to trap or snare cougars (Puma concolor), but cougars are sometimes caught accidentally in snares placed near carrion baits, a technique commonly used by trappers to harvest wolves (Canis lupus). We studied cougar foraging ecology and survival in west-central Alberta to estimate the propensity for cougars to scavenge, their susceptibility to snaring at trapper bait stations, and the implications these have for managing cougar populations. During 2005–2008, we used data from visits to 3,407 Global Positioning System (GPS) location clusters and >400 km of snow tracking of 44 cougars to locate foraging events and calculate scavenging rates. We identified 83 instances of scavenging, and 64% of monitored cougars scavenged at least once. Scavenging rates were higher in winter (0.12 events/week) than in summer (0.04 events/week), reflecting seasonal variation in carrion availability. Individual cougars scavenged at different rates, and winter feeding on carrion occupied up to 50% of total carcass handling time for some cougars. Based on these results we conclude that cougars are facultative scavengers. A propensity to scavenge made cougars susceptible to snaring causing high annual mortality in radiocollared cougars (0.11, 95% CI = 0.03–0.21). Provincial cougar mortality data demonstrate that snaring has increased dramatically as a mortality source in Alberta over the last 2 decades. Mortalities of radiocollared cougars during our study were 100% human caused and the addition of snaring mortality to already high hunting mortality resulted in low annual survival (0.67, 95% CI = 0.53–0.81). Our study is one of the first to identify population-level consequences for nontarget animals killed unintentionally by indiscriminate harvest techniques in a terrestrial ecosystem. Maintaining sustainable cougar harvest where snaring at carrion baits is permitted may require flexible hunting quotas capable of accommodating high cougar snaring mortalities in some years.     

What Drives Wolves: Fear or Hunger? Humans,Diet, Climate and Wolf Activity Patterns

Activity patterns of animals depend on environmental and intrinsic factors. Studies undertaken across the current wolf ( Canis lupus ) range suggested a number of variables that may be correlated with activity patterns of wolves. These factors vary locally and there has been no attempt so far at defining those that ubiquitously impact wolf behaviour. I compared 11 studies (from Alaska to Israel) to assess the influence of (1) public road density, (2) human population density, (3) human-caused mortality, (4) proportion of domestic animals in wolf diet, (5) proportion of forest, (6) latitude and (7) mean annual temperature on nocturnal wolf activity and movements. Nocturnal activity was mainly correlated to the proportion of domestic animals in the diet and the density of public roads, whereas nocturnal movements were mainly correlated to latitude. The importance of latitude indicates that sun periodicity might represent an important signal (`Zeitgeber') for circadian rhythms in wolves. Environmental constraints such as high temperatures during the day and a higher hunting success in crepuscular periods probably limit the ability of wolves to avoid humans by nocturnal behaviour. I therefore suggest that in regions where wolves hunt wild prey, they experience a trade-off between predation risk by humans and increased hunting success during twilight hours.     

Know Thy Enemy: Experience Affects Elk Translocation Success in Risky Landscapes

ABSTRACT To maximize success, reintroduction programs generally select predator-free release areas having high habitat quality. Past studies provide little insight into recovery efforts where multiple, potentially novel, mortality hazards occur. The ability of translocated animals to cope with novel environments can be affected by both pre- and postrelease experiences with habitat and mortality risks. We experimentally released elk (Cervus elaphus) having different background experiences into an area where predators and hunters were prevalent and habitat quality varied. Using a competing risks approach, we predicted the postrelease survival of individuals and their fidelity to release areas as a function of animal source and postrelease encounters with forage resources and areas used by wolves (Canis lupus) or humans. Mortality patterns were consistent with prerelease exposure to mortality risks but not habitat differences among source areas. Wolf predation, poaching, and legal Native hunting were equivalent in magnitude and accounted for the majority of elk mortalities. Familiarity with either wolves or hunters prior to release yielded first-year survival rates 1.9-2.2 times greater than observed for animals naive to both risks. These 2 primary sources of mortality traded off temporally as well as spatially given the proximity of roads, which wolves avoided. The prevalence of forage resources in release areas increased fidelity to release sites but coincided with higher mortality risk during the critical first year, potentially setting an ecological trap for animals naïve to local risks. Translocated individuals largely mediated their respective vulnerabilities over time, showing second-year survival rates equivalent to resident elk. In addition to using source populations that are able to adjust to mortality risks in release areas, spatial and temporal variation in mortality risks might be exploited when planning releases to increase the success of translocations into risky landscapes. (JOURNAL OF WILDLIFE MANAGEMENT 71(2):541–554; 2007)     

Response of Moose Hunters to Predation following Wolf Return in Sweden

Background

Predation and hunter harvest constitute the main mortality factors affecting the size and dynamics of many exploited populations. The re-colonization by wolves (Canis lupus) of the Scandinavian Peninsula may therefore substantially reduce hunter harvest of moose (Alces alces), the main prey of wolves.

Methodology/Principal findings

We examined possible effects of wolf presence on hunter harvest in areas where we had data before and after wolf establishment (n = 25), and in additional areas that had been continuously exposed to wolf predation during at least ten years (n = 43). There was a general reduction in the total number of moose harvested (n = 31,827) during the ten year study period in all areas irrespective of presence of wolves or not. However, the reduction in hunter harvest was stronger within wolf territories compared to control areas without wolves. The reduction in harvest was larger in small (500-800 km²) compared to large (1,200-1,800 km²) wolf territories. In areas with newly established wolf territories moose management appeared to be adaptive with regard to both managers (hunting quotas) and to hunters (actual harvest). In these areas an instant reduction in moose harvest over-compensated the estimated number of moose killed annually by wolves and the composition of the hunted animals changed towards a lower proportion of adult females.

Conclusions/Significance

We show that the re-colonization of wolves may result in an almost instant functional response by another large predator—humans—that reduced the potential for a direct numerical effect on the density of wolves’ main prey, the moose. Because most of the worlds’ habitat that will be available for future colonization by large predators are likely to be strongly influenced by humans, human behavioural responses may constitute a key trait that govern the impact of large predators on their prey.     

Optimal predator management for mountain sheep conservation depends on the strength of mesopredator release

Large predators often suppress ungulate population growth, but they may also suppress the abundance of smaller predators that prey on neonatal ungulates. Antagonistic interactions among predators may therefore need to be integrated into predator–prey models to effectively manage ungulate–predator systems. We present a modeling framework that examines the net impact of interacting predators on the population growth rate of shared prey, using interactions among wolves Canis lupus, coyotes Canis latrans and Dall sheep Ovis dalli dalli as a case study. Wolf control is currently employed on approximately 16 million ha in Alaska to increase the abundance of ungulates for human harvest. We hypothesized that the positive effects of wolf control on Dall sheep population growth could be counteracted by increased levels of predation by coyotes. Coyotes and Dall sheep adult females (ewes) and lambs were radiocollared in the Alaska Range from 1999–2005 to estimate fecundity, age‐specific survival rates, and causes of mortality in an area without wolf control. We used stage‐structured population models to simulate the net effect of wolf control on Dall sheep population growth (λ). Our models accounted for stage‐specific predation rates by wolves and coyotes, compensatory mortality, and the potential release of coyote populations due to wolf control. Wolves were the main predators of ewes, coyotes were the main predators of lambs, and wolves were the main source of mortality for coyotes. Population models predicted that wolf control could increase sheep λ by 4% per year in the absence of mesopredator release. However, if wolf control released coyote populations, our models predicted that sheep λ could decrease by up to 3% per year. These results highlight the importance of integrating antagonistic interactions among predators into predator–prey models, because the net effect of predator management on shared prey can depend critically on the strength of mesopredator release.     

Environmental and social factors influencing wolf (Canis lupus) howling behavior

Animals communicate in a variety of ways and calls are used for a number of important behaviors. Temperature, wind, time of day, and human activities can affect animals’ use of calls, particularly over long distances. Effects of group size on the use of calls can be particularly influential in territorial social carnivores. Where gray wolves (Canis lupus) are hunted by humans, for example, howling may make it easier for hunters to locate individuals and ultimately increase mortality. We hypothesized that a suite of factors would affect wolves’ responses to simulated howling. Specifically, we predicted that howling behavior would increase with (a) group size, (b) pup age, and (c) during crepuscular time periods and howling behavior would decrease (a) where wolves were harvested and (b) when it was hot or windy. Contrary to our prediction, larger groups did not respond as quickly to simulated wolf howls as smaller groups did and minimum and maximum daily temperatures were not good predictors of wolf howling response rates. Individuals in small litters of pups may have responded more quickly to howls than those in large litters because they are eager to seek safety from and have socialization with adults returning from foraging bouts. Although harvest did not appear to affect vocal communication by wolves, group size, pup age, time of day, wind, and number of howls emitted greatly affected wolves’ behavior and responses during howling surveys. Howling responses did not change because of harvest; response rates from wolves were nearly identical with (2.2%) and without (2.3%) harvest. The year-round benefits of long-distance vocal communication may outweigh the costs of increased mortality arising from howling during harvest season.     

Wolf predation on moose and roe deer: chase distances and outcome of encounters

We examined chase distances of gray wolves Canis lupus Linnaeus, 1758 hunting moose Alces alces and roe deer Capreolus capreolus, and recorded details of encounters between wolves and prey on the Scandinavian Peninsula, 1997–2003. In total, 252 wolf attacks on moose and 64 attacks on roe deer were registered during 4200 km of snow tracking in 28 wolf territories. Average chase distances were 76 m for moose and 237 m for roe deer, a difference likely due to variation in body size and vigilance between prey species. A model including prey species, outcome of the attack, and snow depth explained 15–19% of the variation found in chase distances, with shorter chase distances associated with greater snow depth and with successful attacks on moose but not on roe deer. Wolf hunting success did not differ between prey species (moose 43%, roe deer 47%) but in 11% of the wolf attacks on moose at least one moose was injured but not killed, whereas no injured roe deer survived. Compared with most North American wolf studies chase distances were shorter, hunting success was greater, and fewer moose made a stand when attacked by wolves in our study. Differences in wolf encounters with moose and roe deer likely result from different anti-predator behaviour and predator-prey history between prey species.     

Legal protection of wolves in Poland: implications for the status of the wolf population

Legal protection of wolves (Canis lupus) in Poland was implemented in 1998 after 23 years of management as a game species. Wolves occurring in Poland were interconnected with larger populations in the Carpathian Mountains and Belarus, Baltic States and Russia, stable in numbers, and were not considered endangered before the change in legal status affording protection from hunting. Parties calling for wolf protection wanted to stop killing of wolves because of their symbolic nature, but did not have particular management goals to achieve. The government did not accompany the change in legal status by management plan, and therefore, the ban on wolf hunting was weakly enforced. A wolf distribution monitoring demonstrated that wolf range had not expanded 9 years after the hunting ban was implemented, and no increase in wolf numbers was observed. This failure to recover may be explained by: (1) a significant (up to 35%) reduction in the wolves’ prey base 6 years before wolf hunting was stopped, (2) weak enforcement of the protection law, resulting in lack of poaching control of wolves, (3) probable increasing fragmentation and isolation of wolf habitat caused by rapid economic growth in Poland. Inconsistent application of current management policy toward wolves resulted in weak enforcement of regulations and promoted negative attitudes toward the species. To improve the status of wolves in Poland, I recommend a flexible wolf management planning framework that involves and addresses attitudes of hunters and sheep herders, includes a framework to promote strong law enforcement, and consistent, fair compensation for livestock killed by wolves.     

Testing the risk of predation hypothesis: the influence of recolonizing wolves on habitat use by moose

Considered as absent throughout Scandinavia for >100 years, wolves (Canis lupus) have recently naturally recolonized south-central Sweden. This recolonization has provided an opportunity to study behavioral responses of moose (Alces alces) to wolves. We used satellite telemetry locations from collared moose and wolves to determine whether moose habitat use was affected by predation risk based on wolf use distributions. Moose habitat use was influenced by reproductive status and time of day and showed a different selection pattern between winter and summer, but there was weak evidence that moose habitat use depended on predation risk. The seemingly weak response may have several underlying explanations that are not mutually exclusive from the long term absence of non-human predation pressure: intensive harvest by humans during the last century is more important than wolf predation as an influence on moose behavior; moose have not adapted to recolonizing wolves; and responses may include other behavioral adaptations or occur at finer temporal and spatial levels than investigated.     

Wolf pack rendezvous site selection in Greece is mainly affected by anthropogenic landscape features

In wolves, most offspring mortality occurs within the first 6–8 months of their life. As wolf pups pass this entire period at either the den or rendezvous sites, their selection by wolf packs may affect pup survival and recruitment. Rendezvous sites are important for pup survival as they are used during summer and early autumn, when intense human activity may increase pup mortality. Adult wolves and pups can be killed by livestock guarding dogs during summer and intentionally or accidentally during large game hunting in autumn. This study describes factors related to rendezvous site selection in order to enhance their protection and management. We studied the rendezvous site selection of 30 wolf packs in central and northern Greece between 1998 and 2010, after locating 35 sites using the simulated howling survey method and telemetry. We considered a series of environmental and anthropogenic predictors of wolf rendezvous site selection at two spatial scales. At the landscape-population scale, wolves selected rendezvous sites below 1,200 m asl, with large inter-site distance (mean, 12.9 km), and avoided partially forested or open habitats, indicating preference for covered, spaced areas with seasonally stable resources. At the home range scale, wolves selected rendezvous sites away from forest roads and villages, close to water sources, and in areas with low forest fragmentation, indicating avoidance of human presence and disturbance. In the summer of 2011, we used an ensuing resource selection model (RSF, AUC?=?0.818) to successfully locate seven new rendezvous sites outside our previous survey area, verifying the utility of prediction maps (all new sites were at areas with 0.8–1 model probability). Rendezvous prediction maps can be used to reduce field effort when monitoring wolf populations, assess livestock predation risk, design protected areas, and reduce human disturbance on reproductive wolf packs.     

Road avoidance and its energetic consequences for reptiles

Roads are one of the most widespread human‐caused habitat modifications that can increase wildlife mortality rates and alter behavior. Roads can act as barriers with variable permeability to movement and can increase distances wildlife travel to access habitats. Movement is energetically costly, and avoidance of roads could therefore impact an animal's energy budget. We tested whether reptiles avoid roads or road crossings and explored whether the energetic consequences of road avoidance decreased individual fitness. Using telemetry data from Blanding's turtles (Emydoidea blandingii; 11,658 locations of 286 turtles from 15 sites) and eastern massasaugas (Sistrurus catenatus; 1,868 locations of 49 snakes from 3 sites), we compared frequency of observed road crossings and use of road‐adjacent habitat by reptiles to expected frequencies based on simulated correlated random walks. Turtles and snakes did not avoid habitats near roads, but both species avoided road crossings. Compared with simulations, turtles made fewer crossings of paved roads with low speed limits and more crossings of paved roads with high speed limits. Snakes made fewer crossings of all road types than expected based on simulated paths. Turtles traveled longer daily distances when their home range contained roads, but the predicted energetic cost was negligible: substantially less than the cost of producing one egg. Snakes with roads in their home range did not travel further per day than snakes without roads in their home range. We found that turtles and snakes avoided crossing roads, but road avoidance is unlikely to impact fitness through energetic expenditures. Therefore, mortality from vehicle strikes remains the most significant impact of roads on reptile populations.     

Ecosystem Scale Declines in Elk Recruitment and Population Growth with Wolf Colonization: A Before-After-Control-Impact Approach

The reintroduction of wolves (Canis lupus) to Yellowstone provided the unusual opportunity for a quasi-experimental test of the effects of wolf predation on their primary prey (elk – Cervus elaphus) in a system where top-down, bottom-up, and abiotic forces on prey population dynamics were closely and consistently monitored before and after reintroduction. Here, we examined data from 33 years for 12 elk population segments spread across southwestern Montana and northwestern Wyoming in a large scale before-after-control-impact analysis of the effects of wolves on elk recruitment and population dynamics. Recruitment, as measured by the midwinter juvenile∶female ratio, was a strong determinant of elk dynamics, and declined by 35% in elk herds colonized by wolves as annual population growth shifted from increasing to decreasing. Negative effects of population density and winter severity on recruitment, long recognized as important for elk dynamics, were detected in uncolonized elk herds and in wolf-colonized elk herds prior to wolf colonization, but not after wolf colonization. Growing season precipitation and harvest had no detectable effect on recruitment in either wolf treatment or colonization period, although harvest rates of juveniles∶females declined by 37% in wolf-colonized herds. Even if it is assumed that mortality due to predation is completely additive, liberal estimates of wolf predation rates on juvenile elk could explain no more than 52% of the total decline in juvenile∶female ratios in wolf-colonized herds, after accounting for the effects of other limiting factors. Collectively, these long-term, large-scale patterns align well with prior studies that have reported substantial decrease in elk numbers immediately after wolf recolonization, relatively weak additive effects of direct wolf predation on elk survival, and decreased reproduction and recruitment with exposure to predation risk from wolves.     

Dispersal Behavior and the Connectivity Between Wolf Populations in Northern Europe

ABSTRACT The isolated gray wolf (Canis lupus) population of the Scandinavian Peninsular is suffering from inbreeding depression. We studied dispersal of 35 wolves fitted with very high frequency (20) or Global Positioning System—global system for mobile (15) radiocollars in the neighboring Finnish wolf population. The growing wolf population in Finland has high numbers of dispersing individuals that could potentially disperse into the Scandinavian population. About half (53%) of the dispersing wolves moved total distances that could have reached the Scandinavian population if they had been straight-line moves, but because of the irregular pattern of movements, we detected no wolves successfully dispersing to the Scandinavian population. Dispersal to the Scandinavian population was also limited by high mortality of wolves in reindeer (Rangifer tarandus) management areas and by dispersal to Bothnian Bay at times of the year when ice was not present. We suggest that when a small wolf population is separated from source populations by distance, barriers, and human exploitation, wildlife managers could promote the population's viability by limiting harvest in the peripheral areas or by introducing wolves from the source population.     

Summer movements,predation and habitat use of wolves in human modified boreal forests

Grey wolves (Canis lupus), formerly extirpated in Finland, have recolonized a boreal forest environment that has been significantly altered by humans, becoming a patchwork of managed forests and clearcuts crisscrossed by roads, power lines, and railways. Little is known about how the wolves utilize this impacted ecosystem, especially during the pup-rearing summer months. We tracked two wolves instrumented with GPS collars transmitting at 30-min intervals during two summers in eastern Finland, visiting all locations in the field, identifying prey items and classifying movement behaviors. We analyzed preference and avoidance of habitat types, linear elements and habitat edges, and tested the generality of our results against lower resolution summer movements of 23 other collared wolves. Wolves tended to show a strong preference for transitional woodlands (mostly harvested clearcuts) and mixed forests over coniferous forests and to use forest roads and low use linear elements to facilitate movement. The high density of primary roads in one wolf’s territory led to more constrained use of the home territory compared to the wolf with fewer roads, suggesting avoidance of humans; however, there did not appear to be large differences on the hunting success or the success of pup rearing for the two packs. In total, 90 kills were identified, almost entirely moose (Alces alces) and reindeer (Rangifer tarandus sspp.) calves of which a large proportion were killed in transitional woodlands. Generally, wolves displayed a high level of adaptability, successfully exploiting direct and indirect human-derived modifications to the boreal forest environment.     

Evaluating how hunters see and react to telemetry collars on white-tailed deer

Fates of individuals outfitted with radiotransmitters commonly are used for estimating survival rates in populations of large animals that are hunted. Despite precautions, this practice may be subject to complex biases associated with hunter reaction to presence of radiotransmitters. To assess this potential bias we conducted an experiment using artificial deer (i.e., decoys) to measure hunters' abilities to see deer and determine if deer seen were wearing radiocollars. We used logistic regression to quantify probabilities that seeing deer and subsequently seeing radiocollars might be influenced by distance, percent visual obstruction, body orientation, hunter experience, and antler characteristics of deer. Additionally, we evaluated how experience and antler characteristics of deer might influence a hunter's decision to harvest a radiocollared deer. We found that 25.8% of the potentially observable collared deer (n = 663) were subsequently observed by hunters. Odds of observing deer and radiocollars increased 95% and 230%, respectively, for each additional log(yr) of hunting experience. Willingness to harvest radiocollared deer increased 89% for each additional log(yr) of hunting experience and 144% for large-antlered deer relative to antlerless deer. When hunting is an important source of mortality, analysts need to understand how potential biases associated with observing deer are associated with hunters' reactions to and subsequent decisions to harvest radiocollared animals. Our study suggested that presence of radiocollars may influence a deer's potential risk of being harvested and in turn bias telemetry-based estimates of survival, given that hunting mortality is the largest component of total mortality in hunted deer populations. Collar-based telemetry is used nearly universally by wildlife managers and researchers throughout North America and elsewhere to estimate and monitor the survival of big game populations that are managed through hunting. Our findings demonstrate that these estimates are likely subject to complex and systematic biases that managers should consider when evaluating future population-level effects of managed hunting. © 2011 The Wildlife Society     

Wolf population dynamics in the U.S. Northern Rocky Mountains are affected by recruitment and human-caused mortality

Reliable analyses can help wildlife managers make good decisions, which are particularly critical for controversial decisions such as wolf (Canis lupus) harvest. Creel and Rotella (2010) recently predicted substantial population declines in Montana wolf populations due to harvest, in contrast to predictions made by Montana Fish, Wildlife and Parks (MFWP). We replicated their analyses considering only those years in which field monitoring was consistent, and we considered the effect of annual variation in recruitment on wolf population growth. Rather than assuming constant rates, we used model selection methods to evaluate and incorporate models of factors driving recruitment and human-caused mortality rates in wolf populations in the Northern Rocky Mountains. Using data from 27 area-years of intensive wolf monitoring, we show that variation in both recruitment and human-caused mortality affect annual wolf population growth rates and that human-caused mortality rates have increased with the sizes of wolf populations. We document that recruitment rates have decreased over time, and we speculate that rates have decreased with increasing population sizes and/or that the ability of current field resources to document recruitment rates has recently become less successful as the number of wolves in the region has increased. Estimates of positive wolf population growth in Montana from our top models are consistent with field observations and estimates previously made by MFWP for 2008–2010, whereas the predictions for declining wolf populations of Creel and Rotella (2010) are not. Familiarity with limitations of raw data, obtained first-hand or through consultation with scientists who collected the data, helps generate more reliable inferences and conclusions in analyses of publicly available datasets. Additionally, development of efficient monitoring methods for wolves is a pressing need, so that analyses such as ours will be possible in future years when fewer resources will be available for monitoring. © 2011 The Wildlife Society.