Habitat variables associated with wolf (Canis lupus) distribution and abundance in northern Poland

Based on data collected during the National Wolf Census in 2000–01, we analysed the main habitat factors influencing the distribution and abundance of the wolf, Canis lupus, in northern Poland. The study region forms the western border of the continuous Eastern European range of wolves, although attempts at westward dispersal have been observed. Using Geographic Information System techniques, we measured nine habitat variables and three parameters related to wolf occurrence in 134 circular sample plots (radius 7 km, area 154 km² each). We compared 72 plots where wolves were recorded and 62 plots with no signs of wolf presence. Wolf plots were characterized by significantly higher forest cover, less fragmentation of forests, lower density of villages, towns, motorways, and railways than wolf‐free plots. We found a positive correlation between the sum of wolf observations in plots and forest cover. The number of domestic animals killed by wolves was higher in areas with higher indices of wolf abundance and lower forest area. In multiple regression analysis, four independent variables explained 59% of the variation in wolf distribution and abundance in northern Poland: straight‐line distance to continuous range of wolves in Eastern Europe; forest cover; forest fragmentation; and length of major motorways. We conclude that protection of wolves in Poland (since 1998) may not be an adequate conservation measure, especially because of the increasing density of highways and express motorways. Existing forest corridors should be protected and new ones should be restored to ensure long‐term conservation of wolves and allow range expansion into Western Europe.     

Invading white-tailed deer change wolf–caribou dynamics in northeastern Alberta

Human-caused habitat change has been implicated in current woodland caribou (Rangifer tarandus caribou) population declines across North America. Increased early seral habitat associated with industrial footprint can result in an increase in ungulate densities and subsequently those of their predator, wolves (Canis lupus). Higher wolf densities can result in increased encounters between wolves and caribou and consequently higher caribou mortality. We contrasted changes in moose (Alces alces) and deer (Odocoileus spp.) densities and assessed their effects on wolf–caribou dynamics in northeastern Alberta, Canada, pre (1994–1997) versus post (2005–2009) major industrial expansion in the region. Observable white-tailed deer (O. virginianus) increased 17.5-fold but moose remained unchanged. Wolf numbers also increased from approximately 6–11.5/1,000 km². Coincident with these changes, spatial overlap between wolf pack territories and caribou range was high relative to the mid-1990s. The high number of wolf locations in caribou range suggests that forays were not merely exploratory, but rather represented hunting forays and denning locations. Scat analysis indicated that wolf consumption of moose declined substantively during this time period, whereas use of deer increased markedly and deer replaced moose as the primary prey of wolves. Caribou increased 10-fold in the diet of wolves and caribou population trends in the region changed from stable to declining. Wolf use of beaver (Castor canadensis) increased since the mid-1990s. We suggest that recent declines in woodland caribou populations in the southerly extent of their range have occurred because high deer densities resulted in a numeric response by wolves and consequently higher incidental predation on caribou. Our results indicate that management actions to conserve caribou must now include deer in primary prey and wolf reduction programs. © 2010 The Wildlife Society     

Encounter frequencies between GPS-collared wolves (Canis lupus) and moose (Alces alces) in a Scandinavian wolf territory

Over 6,000 GPS fixes from two wolves (Canis lupus) and 30,000 GPS fixes from five moose (Alces alces) in a wolf territory in southern Scandinavia were used to assess the static and dynamic interactions between predator and prey individuals. Our results showed that wolves were closer to some of the moose when inside their home ranges than expected if they had moved independently of each other, and we also found a higher number of close encounters (<500 m) than expected. This suggests that the wolves were actively seeking the individual moose within their territory. Furthermore, the wolves showed a preference for moving on gravel forest roads, which may be used as convenient travel routes when patrolling the territory and seeking areas where the moose are. However, due to the particularly large size of the wolf territory combined with relatively high moose densities, the wolves generally spent a very small proportion of their time inside the home range of each individual moose, and the frequency of encounters between the wolves and any particular moose was very low. We suggest that the high moose:wolf ratio in this large Scandinavian wolf territory compared to that typically occurring in North America, results in a relatively low encounter frequency and a low predation risk for individual moose, as the predation pressure is spread over a high number of prey individuals.     

Coexistence of wolves and humans in a densely populated region (Lower Saxony,Germany)

Since the first sporadic occurrences of grey wolves (Canis lupus) west of the Polish border in 1996, wolves have shown a rapid population recovery in Germany. Wolves are known to avoid people and wolf attacks on humans are very rare worldwide. However, the subjectively perceived threat is considerable, especially as food-conditioned habituation to humans occurs sporadically. Lower Saxony (Germany) has an exceedingly higher human population density than most other regions with territorial wolves; thus, the potential for human–wolf conflicts is higher. Using hunters’ wildlife survey data from 455 municipalities and two years (2014–2015) and data from the official wolf monitoring (557 confirmed wolf presences and 500 background points) collected between 2012–2015, grey wolf habitat selection was modelled using generalized additive models with respect to human population density, road density, forest cover and roe deer density. Moreover, we tested whether habitat use changed in response to human population and road density between 2012/2013 and 2014/2015.Wolves showed a preference for areas of low road density. Human population density was less important as a covariate in the model of the survey data. Areas with higher prey abundance (5–10 roe deer/km²) and areas with >20% forest cover were preferred wolf habitats. Wolves were mostly restricted to areas with the lowest road and human population densities. However, between the two time periods, avoidance of human density decreased significantly.Recolonization of Germany is still in its early stages and it is unclear where this process will halt. To-date authorities mainly concentrate on monitoring measures. However, to avoid conflict, recolonization will require more stringent management of wolf populations and an improved information strategy for rural populations.     

What Cues Do Ungulates Use to Assess Predation Risk in Dense Temperate Forests?

Anti-predator responses by ungulates can be based on habitat features or on the near-imminent threat of predators. In dense forest, cues that ungulates use to assess predation risk likely differ from half-open landscapes, as scent relative to sight is predicted to be more important. We studied, in the Białowieża Primeval Forest (Poland), whether perceived predation risk in red deer (Cervus elaphus) and wild boar (Sus scrofa) is related to habitat visibility or olfactory cues of a predator. We used camera traps in two different set-ups to record undisturbed ungulate behavior and fresh wolf (Canis lupus) scats as olfactory cue. Habitat visibility at fixed locations in deciduous old growth forest affected neither vigilance levels nor visitation rate and cumulative visitation time of both ungulate species. However, red deer showed a more than two-fold increase of vigilance level from 22% of the time present on control plots to 46% on experimental plots containing one wolf scat. Higher vigilance came at the expense of time spent foraging, which decreased from 32% to 12% while exposed to the wolf scat. These behavioral changes were most pronounced during the first week of the experiment but continuous monitoring of the plots suggested that they might last for several weeks. Wild boar did not show behavioral responses indicating higher perceived predation risk. Visitation rate and cumulative visitation time were not affected by the presence of a wolf scat in both ungulate species. The current study showed that perceived predation risk in red deer and wild boar is not related to habitat visibility in a dense forest ecosystem. However, olfactory cues of wolves affected foraging behavior of their preferred prey species red deer. We showed that odor of wolves in an ecologically equivalent dose is sufficient to create fine-scale risk factors for red deer.     

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.     

Wolf Depredation on Domestic Animals in the Polish Carpathian Mountains

Abstract As wolves (Canis lupus) recover in Poland, their depredation on domestic animals is increasing, as have conflicts between wolves and farmers. From 1998 to 2004, I investigated spatial and temporal patterns of 591 verified incidents of wolf depredation in the eastern part of the Polish Carpathian Mountains. The wolf population I surveyed covered an estimated range of 4,993 km². Depredation occurred over 1,595 km² of that area. Sheep accounted for 84.8% of domestic animals killed by wolves. Depredation on sheep and number of sheep farms attacked by wolves increased between 1998 and 2004 (r² = 0.61, P = 0.04 and r² = 0.89, P = 0.02, respectively). The number of wolf attacks on sheep farms in a given year were negatively correlated to red deer (Cervus elaphus) population numbers (R² = 0.69, P = 0.02). The amount of depredation caused by each of the 4 monitored packs was best explained by farm density in their territories (R² = 0.59, P = 0.004). Number of attacks recorded on farms was positively correlated to distance from the farm to the pack's den and rendezvous sites (R² = 0.16, P = 0.04). Of depredation recorded in the 4 pack's territories I surveyed, 77% occurred in 4 farms with no or inadequate protection. I concluded that wolf depredation in the studied area is opportunistic. Wolf predation intensity is a function of decreasing abundance of red deer, the density of sheep farms, and proximity of farms to the summer activity centers of wolf packs, and it is facilitated by poor husbandry practices. These results can aid in preventing wolf depredation and provide a foundation for a wolf management plan.     

Correlates of Mortality in an Exploited Wolf Population

Abstract We investigated the influence of habitat use on risk of death from hunting and trapping of 55 radiocollared gray wolves (Canis lupus) from an exploited insular population in Southeast Alaska, USA. We compared mortality rates for resident and nonresident wolves and used Cox proportional hazards regression to relate habitat composition within 100-m circular buffers around radiolocations to risk of death of resident and nonresident wolves. In addition, we included covariates representing distances to roads, logged stands, and lakes and streams in those analyses. We also compiled harvest data from 31 harvest units within the study area to compare densities of roads and distances from human settlements with rates of harvest. During our study 39 wolves died, of which 18 were harvested legally, 16 were killed illegally, and 5 died from natural causes. Legal and illegal harvest accounted for >87% of the mortality of radiocollared resident and nonresident wolves. Mean annual survival was 0.54 (SE = 0.17) for all wolves. Annual survival was 0.65 (SE = 0.17) for resident wolves and 0.34 (SE = 0.17) for nonresidents. Very few (19%) nonresident wolves survived to colonize vacant territories or join existing wolf packs. Roads, muskegs, and distances from lakes and streams were covariates positively associated with death of resident wolves. Clear-cuts were positively associated with risk of death of nonresident wolves. Rate of harvest increased with density of roads; however, road densities >0.9 km/km² had little additional effect on harvest rates. Harvest rates decreased with ocean distances from nearest towns or settlements. Roads clearly increased risk of death for wolves from hunting and trapping and contributed to unsustainable rates of harvest. Wildlife managers should consider effects of roads and other habitat features on harvest of wolves when developing harvest recommendations. They should expect substantial illegal harvest where wolf habitat is accessible to humans. Moreover, high rates of mortality of nonresident wolves exposed to legal and illegal harvest may reduce or delay successful dispersal, potentially affecting linkages between small disjunct wolf populations or population segments. We conclude that a combination of conservative harvest regulations and large roadless reserves likely are the most effective measures for conserving wolves where risks from human-caused mortality are high.     

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.     

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.     

Differential risk effects of wolves on wild versus domestic prey have consequences for conservation

Predators play integral roles in shaping ecosystems through cascading effects to prey and vegetation. Such effects occur when prey species alter their behavior to avoid predators, a phenomenon called the risk effects of predators. Risk effects of wild predators such as wolves are well documented for wild prey, but not for free ranging domestic animals such as cattle despite their importance for ecosystem function and conservation. We compared risk effects of satellite‐collared wolves (n = 16) on habitat selection by global‐positioning‐system‐collared elk (n = 10) and cattle (n = 31). We calculated resource selection functions (RSFs) in periods before, during and after wolf visits in elk home ranges or cattle pastures. The habitat variables tested included: distance to roads and trails, terrain ruggedness, food‐quality and distance to forest. When wolves were present, elk stayed closer to forest cover and selected less for high‐quality‐food habitat. Thus, the risk effects of wolf presence on elk produced a change in the tradeoff between food and cover selection. Cattle responded by avoiding high‐quality‐food habitat and selecting areas closer to roads and trails (where people likely provided security), but these effects manifested only after wolves had left. Artificial selection in cattle may have attenuated natural anti‐predator behaviors. The effects of predators on ecosystems are likely different when mediated through risk effects on domestic compared to wild animals. Furthermore, predator control in response to livestock predation, an important conservation issue, may produce broad ecosystem effects triggered by decrease of an important predator species. Conservation planners should consider these effects where domestic herbivores are dominant species in the ecosystem.     

Predicting occurrence of wolf territories in Scandinavia

We used logistic regression to compare a set of habitat features inside known Scandinavian wolf Canis lupus territories with the same habitat features in areas outside known territories, but still close enough to be available for wolf colonization. In addition, we analysed changes in habitat variables over time within wolf territories. Wolf territories had lower densities of roads, built-up areas and open land than areas outside wolf territories, but there was no difference in the density of the wolves' main prey, elk Alces alces . The logistic regression model classified 79% of Scandinavia outside the reindeer husbandry area as suitable wolf habitat, that is with a probability of wolf occurrence >0.5. The proportion of built-up areas within the wolf territory decreased as the 'borders' of the wolf territory changed over time. Our model had a reasonably high predictive power, with correct classification in 90% (18 of 20) of the observed wolf territories in the study area. Polygons, randomly distributed outside the observed wolf territories, were correctly classified as not being occupied by wolves in 85% of the cases (17 of 20). This allows a more effective use of resources to, for example, prevent wolf depredation on livestock and dogs.     

Spatial and Temporal Patterns of Wolf Harvest on Registered Traplines in Alberta,Canada

Abstract: Gray wolf (Canis lupus) populations are recovering in many parts of the world and managers from various jurisdictions will be faced with difficult decisions as wolf populations continue to increase. Wolf management in Alberta, Canada, is achieved mostly through trapping on a registered trapline system. Wolf harvest increased over the last 22 years relative to population increases. Most wolf harvests occurred in the Rocky Mountains and surrounding foothills area and this pattern was consistent over time. On average, trappers only harvested an estimated 9.8% of the provincial population annually despite the lack of bag limits or quotas. Harvests are spatially autocorrelated with peak autocorrelation coinciding with average home-range size for wolves in Alberta. When wolf control actions are deemed necessary, trappers are unlikely to remove a sufficient number of wolves over a large enough area to limit subpopulations under the registered trapline system.     

Habitat suitability,corridors and dispersal barriers for large carnivores in Poland

Carnivores are often particularly sensitive to landscape fragmentation. Ecological corridors may help to connect local populations, ensuring gene flow and retaining viable meta-populations. We aimed to establish habitat suitability models for two large carnivores in Poland, the grey wolf Canis lupus Linnaeus, 1758 and the Eurasian lynx Lynx lynx Linnaeus, 1758, based on ecological niche factor analysis (ENFA). Secondly, we calculated least cost paths (LCPs) based on cost values obtained from ENFA. Thirdly, we determined structures that might act as barriers, thus diminishing the value of the corridor unless appropriate conservation measures are taken. We compared some of the results with actual dispersal data of four lynx in eastern Poland. Results indicate that both species are highly marginalised. Less habitat that is currently available in Poland is suitable for lynx than for wolves. We determined a total of 76 LCPs. Comparison of these theoretical corridors with actual dispersal routes suggests that the traits of calculated LCPs are mostly within the range of those of real routes. We highlight a variety of features that might act as barriers, such as major roads (including planned highways), urbanized areas, and large un-forested areas. We give suggestions where concerted conservation efforts (eg wildlife passages) might be particularly well-directed.     

A predictive spatial model for gray wolf (Canis lupus) denning sites in a human-dominated landscape in western Iran

Predictive models of species denning habitat are useful for understanding distribution of core use areas and identifying areas with potential conflicts. Wolf den site selection in unmanaged areas with dominance of agriculture-lands and human disturbance is not completely understood. We used a GIS multivariate model based on the Mahalanobis distance statistic and 11 digital habitat layers to evaluate gray wolf denning site suitability in Hamedan province (HP), western Iran. Results showed that >74 % of dens occurred in potential denning sites that occupied 14 % (2,785 km²) of the study area. Based on sensitivity analysis with ROC, we found that the distance to rangelands, elevation, and distance to roads, human settlements and streams were the most predictive variables. Our potential distribution model of wolf den sites showed a very good overall performance according to classification accuracy and discrimination capacity. Denning in elevated rugged terrains adjacent to rangelands and away from roads and villages revealed that wolf den distribution in HP is highly influenced primarily by factors associated with human disturbance. The derived predicted distribution map can be used to prioritize areas for conservation, identify areas with potential conflicts, and to implement adaptive management in regions beyond wilderness and protected areas.     

Assessment of anadromous salmon resources in the diet of the Alexander Archipelago wolf using stable isotope analysis

The Alexander Archipelago wolf (Canis lupus ligoni) is unique to southeast Alaska, occurring on islands south of Frederick Sound and along the mainland between Dixon Entrance and Yakutat Bay. Sitka black-tailed deer (Odocoileus hemionus sitkensis) are an important prey species for wolves across the southern part of the region. Spawning salmon (Onchorynchus sp.) are seasonally available but their presence in wolf diets has not previously been quantified. We examined the range of bone collagen δ¹³C and δ¹⁵N values for wolves throughout southeast (n = 163) and interior (n = 50) Alaska and used a dual-isotope mixing model to determine the relative contribution of salmon-derived marine protein in the diet. Southeast Alaska wolves consumed significantly more salmon (mean ± SE: 18.3 ± 1.2%) than did wolves from interior Alaska (9.1 ± 0.6%, P<0.001). Wolves on the southeast Alaska mainland appeared to have higher marine isotopic signatures than island wolves, although this difference was not significant. Variation among individual wolf diets was higher for southeast than for interior Alaska wolves, and variation was highest in coastal mainland wolf diets (P<0.001). Marine resources may augment the diet of southeast Alaska wolves during seasonal or annual fluctuations in the availability of deer, particularly in those areas on the mainland where densities of terrestrial ungulates are relatively low. Received: 1 July 1998 / Accepted: 23 February 1999     

Territory size of wolves Canis lupus: linking local (Białowieża Primeval Forest, Poland) and Holarctic-scale patterns

Factors affecting territory size in wolves Canis lupus were studied at 2 scales, the local population (Bia?owie?a Primeval Forest (BPF), eastern Poland) and the geographic range of species (literature review from 14 localities in the Holarctic). Four packs of wolves were studied by radio‐tracking in BPF from 1994 to 1999. The annual territories of packs (Minimum convex polygons with 95% of locations) averaged 201 km² (SD 63, range 116–310). Core areas of territories (50% MCP) covered from 14 to 78 km² (mean 35). Territory sizes and core areas both were negatively correlated to the encounter rates of ungulates (mean number of ungulates seen per unit time spent in the forest by human observers). Pack size (3–8 wolves) did not influence territory size. Home ranges of individual wolves from the same pack varied with season as well as the age, sex, and reproductive status of the wolf. Review of literature from North America and Europe (42–66^oN), showed that latitude and prey biomass were essential factors shaping the biogeographic variation in wolf territory size. Territories increased with latitude and declined with growing biomass of prey. The analysis showed that latitude acted partly independently of the south–north gradient in prey abundance. At similar standing crop of ungulate biomass (100 kg km^?2), wolf territories would average 140 km² at 40^oN, 370 km² at 50^oN, and 950 km² at 60^oN. Pack size was larger at northern latitudes, but the increase did not keep pace with enlargement of territories. Within‐territory density of wolves declined from 2.5–3 wolves 100 km^?2 at 40–45^oN to 0.7 wolves 100 km^?2 at 60^oN. Our analyses documented similarities regarding the role of prey resources in shaping wolf territoriality at the different scales. Furthermore, a macroecological approach revealed additional factors affecting wolf territory size that were not emergent from knowledge of local population.     

Recolonizing gray wolves increase parasite infection risk in their prey

The recent recolonization of Central Europe by the European gray wolf (Canis lupus) provides an opportunity to study the dynamics of parasite transmission for cases when a definitive host returns after a phase of local extinction. We investigated whether a newly established wolf population increased the prevalence of those parasites in ungulate intermediate hosts representing wolf prey, whether some parasite species are particularly well adapted to wolves, and the potential basis for such adaptations. We recorded Sarcocystis species richness in wolves and Sarcocystis prevalence in ungulates harvested in study sites with and without permanent wolf presence in Germany using microscopy and DNA metabarcoding. Sarcocystis prevalence in red deer (Cervus elaphus) was significantly higher in wolf areas (79.7%) than in control areas (26.3%) but not in roe deer (Capreolus capreolus) (97.2% vs. 90.4%) or wild boar (Sus scrofa) (82.8% vs. 64.9%). Of 11 Sarcocystis species, Sarcocystis taeniata and Sarcocystis grueneri occurred more often in wolves than expected from the Sarcocystis infection patterns of ungulate prey. Both Sarcocystis species showed a higher increase in prevalence in ungulates in wolf areas than other Sarcocystis species, suggesting that they are particularly well adapted to wolves, and are examples of “wolf specialists”. Sarcocystis species richness in wolves was significantly higher in pups than in adults. “Wolf specialists” persisted during wolf maturation. The results of this study demonstrate that (1) predator–prey interactions influence parasite prevalence, if both predator and prey are part of the parasite life cycle, (2) mesopredators do not necessarily replace the apex predator in parasite transmission dynamics for particular parasites of which the apex predator is the definitive host, even if meso‐ and apex predators were from the same taxonomic family (here: Canidae, e.g., red foxes Vulpes vulpes), and (3) age‐dependent immune maturation contributes to the control of protozoan infection in wolves.     

Winter predation patterns of wolves in Northwestern Wyoming

Wolf (Canis lupus) diets and potential effects on prey have been a prominent subject of interest to wildlife researchers and managers since reintroduction into Yellowstone National Park, Wyoming, USA, in 1995 and 1996. Post-reintroduction, wolves expanded south and recolonized areas in the southern Yellowstone ecosystem. Elk (Cervus elaphus) in this area are supplementally fed during winter (Dec–Mar) at state-managed feedgrounds, resulting in high-density congregations of elk. From December to March 2000–2007, we determined the winter predation patterns of wolves by examining the remains of 289 wolf kills on 3 state-managed feedgrounds and adjacent winter range near Jackson, Wyoming. During winters 2002–2005, we also monitored the movements of radio-collared elk on feedgrounds to describe the response of elk to the presence of wolf kills. Thirty-seven percent (n = 106) of kills were located on elk feedgrounds where elk composition included 49% calves, 42% adult females, 5% adult males, and 5% unknown. Sixty-three percent (n = 183) of kills were located on winter range adjacent to feedgrounds and prey species consisted of 90% elk (38% calves, 35% adult females, 24% adult males, 2% unknown), 9% moose (Alces alces; 13% calves, 69% adult females, 6% adult males, 1% unknown), 1% mule deer (Odocoileus hemionus; 1 fawn, 1 adult female), and 0.5% adult female bison (Bison bison). Mean age of elk killed on feedgrounds was 4.2 years (range = 0–20) and 4.6 years (range = 0–23) on winter range. Calves were selected more than available in most years with female elk killed less than expected. Adult males were killed more than expected in 2005–2007. Eighty-eight percent (n = 198) of the time elk remained on the feedground even when wolves made a kill. Less commonly, elk left the feedground, gathered in larger herds on adjacent feedgrounds absent of wolves, and returned within a few days (6%, n = 13) or left the feedground for another feedground and did not return for the rest of the winter (6%; n = 14). Elk were less likely to leave feedgrounds in the presence of a wolf kill when there were more elk on that feedground. Elk left feedgrounds with greater topography and tree cover (Alkali and Fish Creek) and gathered on the flat, open feedgrounds (Patrol Cabin) more frequently than they left flat, open feedgrounds for feedgrounds with greater topography and tree cover. Our results indicate wolves in our study area primarily preyed on elk and exhibited a strong preference for elk calves. High-density concentrations of elk on feedgrounds will continue to be an attractant for wolves. Although elk leave feedgrounds for reasons other than wolf presence, any displacement of elk from feedgrounds due to wolves will be temporary. State managers have the ability to alter management strategies (e.g., increasing wolf harvest, phasing out elk feeding, increasing the intensity of elk feeding) in an effort to affect predator-prey relationships. © 2019 The Wildlife Society.     

Conservation of wildlife populations: factoring in incremental disturbance

Progressive anthropogenic disturbance can alter ecosystem organization potentially causing shifts from one stable state to another. This potential for ecosystem shifts must be considered when establishing targets and objectives for conservation. We ask whether a predator–prey system response to incremental anthropogenic disturbance might shift along a disturbance gradient and, if it does, whether any disturbance thresholds are evident for this system. Development of linear corridors in forested areas increases wolf predation effectiveness, while high density of development provides a safe‐haven for their prey. If wolves limit moose population growth, then wolves and moose should respond inversely to land cover disturbance. Using general linear model analysis, we test how the rate of change in moose (Alces alces) density and wolf (Canis lupus) harvest density are influenced by the rate of change in land cover and proportion of land cover disturbed within a 300,000 km² area in the boreal forest of Alberta, Canada. Using logistic regression, we test how the direction of change in moose density is influenced by measures of land cover change. In response to incremental land cover disturbance, moose declines occurred where <43% of land cover was disturbed; in such landscapes, there were high rates of increase in linear disturbance and wolf density increased. By contrast, moose increases occurred where >43% of land cover was disturbed and wolf density declined. Wolves and moose appeared to respond inversely to incremental disturbance with the balance between moose decline and wolf increase shifting at about 43% of land cover disturbed. Conservation decisions require quantification of disturbance rates and their relationships to predator–prey systems because ecosystem responses to anthropogenic disturbance shift across disturbance gradients.