Selection of rendezvous sites and reuse of pup raising areas among wolvesCanis lupus of north-eastern Apennines, Italy

A coarse scale analysis was carried out of factors affecting rendezvous site selection and fidelity to pup raising areas in wolfCanis lupus Linnaeus, 1758 packs inhabiting the north eastern Apennines, Italy. From 1993 to 2004, 44 rendezvous sites were identified, and compared with random sites for variables related to topography, habitat, and human presence. Rendezvous sites were significantly more frequent inside protected areas and at a significantly greater distance from paved roads and villages than random sites. Moreover, they were located on a significantly steeper slope and at a higher absolute elevation than random sites, whereas we did not observe selection of a particular aspect. Deciduous forests were selected, whereas open areas were not used. Canopy cover above 70% was positively selected. Distance from protected areas border and paved roads, and presence of deciduous forest proved the main factors affecting rendezvous site distribution. In four cases we observed the reuse of the same rendezvous sites for two consecutive years. Some packs tended to locate their rendezvous sites in small portions of their territory, thus implying that some areas are more suitable for pup raising.     

Surveying Predicted Rendezvous Sites to Monitor Gray Wolf Populations

ABSTRACT We used rendezvous site locations of wolf (Canis lupus) packs recorded during 1996–2006 to build a predictive model of gray wolf rendezvous site habitat in Idaho, USA. Variables in our best model included green leaf biomass (Normalized Difference Vegetation Index), surface roughness, and profile curvature, indicating that wolves consistently used wet meadow complexes for rendezvous sites. We then used this predictive model to stratify habitat and guide survey efforts designed to document wolf pack distribution and fecundity in 4 study areas in Idaho. We detected all 15 wolf packs (32 wolf pack-yr) and 20 out of 27 (74%) litters of pups by surveying <11% of the total study area. In addition, we were able to obtain detailed observations on wolf packs (e.g., hair and scat samples) once we located their rendezvous sites. Given an expected decrease in the ability of managers to maintain radiocollar contact with all of the wolf packs in the northern Rocky Mountains, rendezvous sites predicted by our model can be the starting point and foundation for targeted sampling and future wolf population monitoring surveys.     

Response of Wolves to Experimental Disturbance at Homesites

ABSTRACT Events during the denning period (parturition to first autumn) often determine the reproductive success of wolves (Canis lupus). Consequently, there is concern about the potential adverse effects of human-caused disturbance at wolf den and rendezvous sites (homesites), but relatively little information on this subject is available. We conducted standardized experimental disturbance treatments at 12 unique wolf homesites in the Northwest Territories, Canada, during summers 2002 and 2003. The treatment consisted of an intruder approaching a homesite once per day for 3 consecutive days and recording behavioral responses, response distance, and response intensity of wolves. We counted pups and estimated their ages prior to the initial treatment at each site. Adult wolves moved pups at 3 of 6 treated homesites in each year. The amount and type of known human activity within a pack's home range did not influence whether adults moved pups in response to the treatment. The response intensity of wolves to the treatment was inversely related to the amount of human activity near a homesite. There was no relationship between the distance at which wolves responded to the intruder and the amount or type of human activity. There was a positive relationship between increasing age of pups and their relocation in response to the treatment. Reproductive success was not influenced by the treatment or by the amount and type of human activity. Treated sites were used by wolves the following year in the same proportion as untreated sites. It appears that pups are most vulnerable early in the year when less mobile; therefore, managers should consider age of pups before human activity at or near wolf homesites occurs.     

Den shifting by wolves in semi-wild landscapes in the Deccan Plateau, Maharashtra, India

Reproductive success is crucial for the survival and persistence of any species. The Deccan Biogeographic Zone of India is the stronghold of a population of the Indian wolf Canis lupus pallipes . Gaining a better understanding of the den-use pattern of wolves in different areas in this zone is thus vital for their conservation and management. The wolves excavated multiple dens in our study sites and kept shifting their litters among them. Major disturbance factors around denning sites were active stone quarries, traffic, crop harvesting and livestock movement. One wolf pack used 14 dens in four breeding seasons. Discriminant function analysis indicated that den shifting by wolves was not entirely governed by disturbance levels at den sites. Increasing age of pups was one of the main factors associated with den shifting rather than the magnitude of disturbance ( χ ²=34.26, d.f.=12, P <0.001). Tolerance to disturbance around dens during the early stages of pup development was negatively correlated ( r =−0.519, P <0.05) with availability of water and age of pups ( r =−0.613, P <0.01). Excavation of multiple dens by wolves was apparently related to den shifting, which seemingly is a survival strategy of wolves in these semi-wild human-dominated landscapes in the Deccan Biogeographic Zone. Principal components analysis indicated that during the initial stages of pup development, nature of the land, den orientation and distance of the den from roads were important cues in addition to age category of pups for den shifting. The analysis also suggested that factors such as distance of the new den where pups are to be transferred, distance from water source and availability of fox Vulpes bengalensis holes for den use were also important.     

Predicting the Spatial Distribution of Wolf (Canis lupus) Breeding Areas in a Mountainous Region of Central Italy

Wolves (Canis lupus) in Italy represent a relict west European population. They are classified as vulnerable by IUCN, though have increased in number and expanded their range in recent decades. Here we use 17 years of monitoring data (from 1993 to 2010) collected in a mountainous region of central Italy (Arezzo, Tuscany) in an ecological niche-based model (MaxEnt) to characterize breeding sites (i.e. the areas where pups were raised) within home ranges, as detected from play-back responses. From a suite of variables related to topography, habitat and human disturbance we found that elevation and distance to protected areas were most important in explaining the locality of wolf responses. Rendezvous sites (family play-back response sites) typically occurred between 800 and 1200 m a.s.l., inside protected areas, and were usually located along mountain chains distant from human settlements and roads. In these areas human disturbance is low and the densities of ungulates are typically high. Over recent years, rendezvous sites have occurred closer to urban areas as the wolf population has continued to expand, despite the consequent human disturbance. This suggests that undisturbed landscapes may be reaching their carrying capacity for wolves. This, in turn, may lead to the potential for increased human-wolf interactions in future. Applying our model, both within and beyond the species’ current range, we identify sites both within the current range and also further afield, that the species could occupy in future. Our work underlines the importance of the present protected areas network in facilitating the recolonisation by wolves. Our projections of suitability of sites for future establishment as the population continues to expand could inform planning to minimize future wolf-human conflicts.     

Wolf denning behaviour in response to external disturbances and implications for pup survival

We conducted a 15-year study (1999–2014) of wolf denning behavior in Naliboki Forest, Belarus, to determine the extent to which some external factors (rain, mosquitoes, human disturbances) and litter size are negatively correlated with pup survival until autumn.For this purpose we entirely documented denning areas of 30 breeding wolf pairs. Initial denning area includes the natal den site and (ii) secondary den sites between which the parent wolves transported sucessively the litter during the rearing season.The number of den sites per denning area ranged from 1 to 4. The distances in kilometres differed significantly between den sites. The total number of dens (surface dens and burrows) was higher in the natal den site compared with the other den sites. The number of surface dens per den site was higher compared with burrows in den sites 1, 2 and 3 but not den site 4. A total of 168 pups was found over the 15 years. Litter sizes ranged from 1 to 11 (average 5.6 ± 1.9). The rate of pup survival ranged from 0 to 100% and averaged 54.5 ± 29%.We examined the bivariate association between environmental conditions (mosquitoes, rainfall and human presence) and number of surface dens, total number of burrows and number of den sites. Environmental conditions were assessed from spring to mid-summer. Results show that there was a positive relationship between environmental conditions and number of den sites, and also that more difficult environmental conditions (rainy weather, many mosquitoes) decreased pup survival both in absolute and in relative terms. A cluster analysis showed that pup survival was also influenced by human presence. Wolves probably have a more complicated denning behaviour following stressful event which may lead to less effectiveness for feeding or protecting pups. The number of pups surviving was also found to be lower when there were many (8–10) pups because den shifting is more risky in this case. Burrows improve pup survival. Our results tend to suggest that, in our study area, a high mortality rate of wolf pups could be related to external and internal factors of stress.     

Movements and Home Sites of Timber Wolves in Algonquin Park

Ecological studies of timber wolves (Canis lupus) in a forestedenvironment have always been difficult to undertake in the past,particularly during the summer months, because of the lack ofa suitable technique. In 1960 Pimlott devised a technique whichemployed the use of broadcast wolf howls in locating wolvesin the field. This report reviews the success of this techniquein studying the movements of two packs of wolves, and theiruse of home sites in Algonquin Park, Ontario. Wolves responded by howling a total of 476 occasions or approximately13% of the occasions that broadcast howls were given. Humanimitations were more successful than tape-recordings in inducingresponses. Wolves responded at any time of day, but dusk wasthe most favorable period. They also responded more frequentlyin July and August than in May and June. Two types of home sites were found: the den site, occupied duringthe early life of the pups, and the rendezvous site, a placeoccupied by wolves during later development of pups. All ofhe home sites were adjacent to some immediate source of water.The movement of wolves appeared also to be concentrated alongthe water courses. The locations of the home sites and the evidence obtained fromhowling responses, tracks, and scats suggested that the summerrange comprised a minimum area of eight square miles. The samerange was utilized by a pack in 1961 and 1963.     

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.     

The importance of fine-scale breeding site selection patterns under a landscape-sharing approach for wolf conservation

For large carnivores persisting in human-dominated landscapes, conservation planning is often hindered by the large spatial requirements of these species, availability of protected areas, and human land uses. Protected areas are usually too small to support viable populations, and scattered across a human land-use matrix. Therefore, large carnivore conservation should be planned at large spatial scales under a land-sharing approach (allowing the coexistence between large carnivores and people in the same landscape), which means increasing the focus on the human-dominated matrix. Most of the critical factors determining large carnivore persistence (i.e., those related to food availability and vulnerability to humans) interact synergistically in space and time during the breeding season. Here, using as a case study a wolf population in NW Iberia, we studied fine-scale breeding site selection patterns (1 and 9 km²) in relation to human pressure, and the availability of food and refuge. The selection of wolf breeding sites in this human-dominated landscape was not determined by potential availability of prey biomass in the immediate vicinity (1 km²). However, wolves selected breeding sites with high availability of refuge (concealing vegetation), and low human accessibility and activity levels. Paved roads showed the highest proportion of independent contribution to explaining breeding site selection patterns (negative influence), being followed by refuge availability (positive influence) and the remoteness of breeding sites in relation to the surrounding spatial context (positive influence). Refuge availability, even at very small spatial scales taking into account the spatial requirements of wolves, may compensate for moderate levels of human activities in the vicinity of breeding sites. The strength of breeding selection patterns varied along a hierarchical process at different spatial scales. Under a landscape-sharing approach, integrating key processes observed in the human-dominated matrix, such as breeding site selection patterns, into landscape planning is of paramount importance for carnivore conservation. By temporally restricting human use on breeding sites and small portions of surrounding lands (~ 1 km²), and by maintaining several small refuge areas interspersed within the human-dominated matrix, we could favor wolf persistence without reducing land availability for other uses, improving the conditions for coexistence between wolves and humans.     

Efficient,Noninvasive Genetic Sampling for Monitoring Reintroduced Wolves

ABSTRACT Traditional methods of monitoring gray wolves (Canis lupus) are expensive and invasive and require extensive efforts to capture individual animals. Noninvasive genetic sampling (NGS) is an alternative method that can provide data to answer management questions and complement already-existing methods. In a 2-year study, we tested this approach for Idaho gray wolves in areas of known high and low wolf density. To focus sampling efforts across a large study area and increase our chances of detecting reproductive packs, we visited 964 areas with landscape characteristics similar to known wolf rendezvous sites. We collected scat or hair samples from 20% of sites and identified 122 wolves, using 8–9 microsatellite loci. We used the minimum count of wolves to accurately detect known differences in wolf density. Maximum likelihood and Bayesian single-session population estimators performed similarly and accurately estimated the population size, compared with a radiotelemetry population estimate, in both years, and an average of 1.7 captures per individual were necessary for achieving accurate population estimates. Subsampling scenarios revealed that both scat and hair samples were important for achieving accurate population estimates, but visiting 75% and 50% of the sites still gave reasonable estimates and reduced costs. Our research provides managers with an efficient and accurate method for monitoring high-density and low-density wolf populations in remote areas.     

Planning and Implementing a Reintroduction of Wolves to Yellowstone National Park and Central Idaho

The Northern Rocky Mountain Wolf Recovery plan proposed reintroduction of Canis lupus (gray wolf) to Yellowstone National Park and central Idaho as part of a wolf restoration plan for the northern Rocky Mountains of the United States. Strong opposition from some factions within the region forestalled the action for two decades. An environmental impact statement, conducted in 1992–1994 with extensive public input, culminated in a proposal to reintroduce wolves designated as “non-essential—experimental” under Section 10 (j) of the federal Endangered Species Act. This approach, approved by the Secretary of the Interior in 1994, provided for wolf restoration while allowing management flexibility to deal with concerns of the local public. A reintroduction plan was developed in the summer and fall of 1994. Acquiring, holding, transporting, and releasing suitable wolves for reintroduction presented a myriad of technical and logistical challenges that required effective planning and coordination by all participants. In January 1995, 29 wolves were captured in Alberta and transported to Yellowstone National Park (14) and central Idaho (15). Idaho wolves were freed immediately upon arrival; Yellowstone wolves (three family groups) were held in acclimation pens in the park until late March. Most Idaho wolves traveled extensively within the area intended for them, averaging 82 km net distance away from release sites after 5 months (range = 30–220 km), and three male-female pairs formed by July. After 5 months in the wild, at least 13 of 15 Idaho-released wolves were alive within the intended area, as were 13 of 14 Yellowstone wolves; one wolf was known to have been illegally killed in each area. No livestock were killed. Wolves released into Yellowstone Park continued to live as packs, stayed closer to their release sites (x = 22 km at end of June), and settled into home ranges; two packs produced a total of nine pups. The progress of the reintroduction program in its first year far exceeded expectations. Reintroductions of about 15 wolves to each area for 2–4 more years are scheduled, but the project may be shortened because of early successes. Future reintroduction planners can expect sociocultural issues to pervade the effort, but they can be optimistic that, from a biological standpoint, reintroduction of wolves has strong potential as a restoration technique.     

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.     

Distribution, status and conservation of Indian gray wolf (Canis lupus pallipes) in Karnataka, India

The Indian gray wolf Canis lupus pallipes is the major large carnivore in the plains of Karnataka, India. We carried out a study on its distribution and status from November 2001 to July 2004. We estimated 555 wolves occupying about 123 330 km² of the state. In the past 40 years, wolves have disappeared from the southern plateau from an area of about 31 801 km². Their distribution is now largely restricted to the north-eastern dry plains. The wolf has also disappeared in recent years from some 'protected areas' such as Melkote Temple Wildlife Sanctuary, and their present population is largely found in 'non-protected' areas. Blackbucks are the only natural prey of wolves in Karnataka, but their density in most parts of the state is extremely low. The major prey species is domestic livestock, especially sheep. The available 'remote area' (forests or rocky terrains) in the wolf-occupied regions determined the status of the wolf. Killing of adult wolves and pups was common throughout the range of the wolf. However, such killings were made largely by local sheepherders with small sheep holdings and not by nomadic shepherds who maintained large sheep herds. The forests in the north-eastern parts of the state exist in small patches every few kilometers. Because each wolf pack ranges over large distances and is by and large a commensal species, we propose that the management of these small forest patches, considering them as components of a larger landscape, is the only effective conservation practice for the wolf. Although existing locally in low densities, because of a large ranging area of a single pack, the seemingly isolated wolf packs can become parts of a large metapopulation, providing a sustainable population.     

Resting in risky environments: the importance of cover for wolves to cope with exposure risk in human-dominated landscapes

Centuries of persecution have influenced the behaviour of large carnivores. For those populations persisting in human-dominated landscapes, complete spatial segregation from humans is not always possible, as they are in close contact with people even when they are resting. The selection of resting sites is expected to be critical for large carnivore persistence in human-dominated landscapes, where resting sites must offer protection to counteract exposure risk. Using wolves (Canis lupus) as a model species, we hypothesised that selection of resting sites by large carnivores in human-dominated landscapes will be not only influenced by human activities, but also strongly determined by cover providing concealment. We studied the fine-scale attributes of 546 wolf resting sites and confronted them to 571 random points in NW Iberia. Half of resting sites (50.8 %) were found in forests (mainly forest plantations, 73.1 %), 43.4 % in scrublands, and only 5.8 % in croplands. Compared to random points, wolves located their resting sites far away from paved and large unpaved roads and from settlements, whereas they significantly selected areas with high availability of horizontal (refuge) and canopy cover. The importance of refuge was remarkably high, with its independent contribution alone being more important than the contribution of all the variables related to human pressure (distances) pooled (51.1 vs 42.8 %, respectively). The strength of refuge selection allowed wolves even to rest relatively close to manmade structures, such as roads and settlements (sometimes less than 200 m). Maintaining high-quality refuge areas becomes an important element to favour the persistence of large carnivores in human-dominated landscapes as well as human-carnivore coexistence, which can easily be integrated in landscape planning.     

Spatial variability in wolf diet and prey selection in Galicia (NW Spain)

We studied wolf (Canis lupus) diet for three different landscapes in the north-western Iberian Peninsula, differing in land uses and availability of food for wolves. We examined 2740 scats, collected over a period of 4 years, in order to describe wolf diet, its geographic variation, and trophic preferences. The most consumed species were wild pony, roe deer and cattle. We observed differences in wolf diet among the three study sites, related to the availability and accessibility of food resources in each habitat. For the two study sites in northern and central Galicia, wolves showed similar diet, with high occurrence of wild pony (37 vs. 34%) and cattle (20 vs. 23%), but differing in the consumption of wild ungulates (16 vs. 8%) and carrion (7 vs. 14%). Wolf diet in eastern Galicia’s mountain ranges was entirely different, due to the higher consumption of wild ungulates (70%). Wolves showed clear prey selection patterns. Between wild ponies and livestock, wolves positively selected ponies. Among wild ungulates, wolves positively selected roe deer. Wild pony and roe deer are key species for wolf feeding in Galicia. In the Galician wild pony range, ponies are the main food for wolves. Given that the availability of wild ponies may contribute to the decrease in wolf predation on cattle, it is essential to develop innovative administrative decisions in such areas to preserve this traditional equid population. In the same way, the population of roe deer should be strengthened in the livestock areas outside the range of wild pony.     

Additional considerations for gray wolf management after their removal from Endangered Species Act protections

Mech (2010) provided a review of options involving regulated, public hunting of gray wolves (Canis lupus) when states regain control of wolf management. We agree with his general conclusion that the use of lethal management should focus more in areas of conflict and less in wilderness areas, especially near protected places like national parks. Here, we expand on Mech's work and provide additional considerations that could be incorporated into state management plans to make them more acceptable to an increasingly diverse group of interested stakeholders, including: 1) the use of human dimensions research to understand the conditions under which stakeholders find lethal management acceptable, and to evaluate the acceptability of agency efforts to increase tolerance for wolves; 2) employing preventative measures to protect livestock and pets, especially in cases where wolf packs are highly visible to the public; and 3) selective use of sport hunting in areas where wolf impacts are deemed unacceptable. © 2011 The Wildlife Society.     

Non-invasive acoustic detection of wolves

Monitoring wolves (Canis lupus) is a difficult and often expensive task due to high mobility, pack dynamic, shyness and nocturnal activity of this species. Wolves communicate acoustically through howling, within pack and with packs of the neighbourhood. A wolf howl is a low-frequency vocalization that can be transmitted over long distances and thus it can be used for monitoring. Elicited howling survey is a current method to monitor wolves in different areas all over the world. Elicited howling, however, may be invasive to residential wolf packs and could create possible negative reactions from the human population. Here we show that it is possible to detect wolves by recording spontaneous howling events. We measured the sound pressure level of wolf howls by captive individuals and we further found that elicited howling may be recorded and clearly identified up to a distance of 3 km. We finally conducted a non-invasive acoustic detection of wolves in a free-ranging population. The use of passive sound recorders may provide a powerful non-invasive tool for future wolf monitoring and could help to establish sustainable management plans for this species.     

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.     

Estimation of pack density in grey wolf (<Emphasis Type="Italic">Canis lupus</Emphasis>) by applying spatially explicit capture-recapture models to camera trap data supported by genetic monitoring

Background

Density estimation is a key issue in wildlife management but is particularly challenging and labour-intensive for elusive species. Recently developed approaches based on remotely collected data and capture-recapture models, though representing a valid alternative to more traditional methods, have found little application to species with limited morphological variation. We implemented a camera trap capture-recapture study to survey wolf packs in a 560-km² area of Central Italy. Individual recognition of focal animals (alpha) in the packs was possible by relying on morphological and behavioural traits and was validated by non-invasive genotyping and inter-observer agreement tests. Two types (Bayesian and likelihood-based) of spatially explicit capture-recapture (SCR) models were fitted on wolf pack capture histories, thus obtaining an estimation of pack density in the area.

Results

In two sessions of camera trapping surveys (2014 and 2015), we detected a maximum of 12 wolf packs. A Bayesian model implementing a half-normal detection function without a trap-specific response provided the most robust result, corresponding to a density of 1.21?±?0.27 packs/100 km² in 2015. Average pack size varied from 3.40 (summer 2014, excluding pups and lone-transient wolves) to 4.17 (late winter-spring 2015, excluding lone-transient wolves).

Conclusions

We applied for the first time a camera-based SCR approach in wolves, providing the first robust estimate of wolf pack density for an area of Italy. We showed that this method is applicable to wolves under the following conditions: i) the existence of sufficient phenotypic/behavioural variation and the recognition of focal individuals (i.e. alpha, verified by non-invasive genotyping); ii) the investigated area is sufficiently large to include a minimum number of packs (ideally 10); iii) a pilot study is carried out to pursue an adequate sampling design and to train operators on individual wolf recognition. We believe that replicating this approach in other areas can allow for an assessment of density variation across the wolf range and would provide a reliable reference parameter for ecological studies.

     

Inter-specific territoriality in a Canis hybrid zone: spatial segregation between wolves, coyotes, and hybrids

Gray wolves (Canis lupus) and coyotes (Canis latrans) generally exhibit intraspecific territoriality manifesting in spatial segregation between adjacent packs. However, previous studies have found a high degree of interspecific spatial overlap between sympatric wolves and coyotes. Eastern wolves (Canis lycaon) are the most common wolf in and around Algonquin Provincial Park (APP), Ontario, Canada and hybridize with sympatric gray wolves and coyotes. We hypothesized that all Canis types (wolves, coyotes, and hybrids) exhibit a high degree of spatial segregation due to greater genetic, morphologic, and ecological similarities between wolves and coyotes in this hybrid system compared with western North American ecosystems. We used global positioning system telemetry and probabilistic measures of spatial overlap to investigate spatial segregation between adjacent Canis packs. Our hypothesis was supported as: (1) the probability of locating wolves, coyotes, and hybrids within home ranges ( $\bar{x}$  = 0.05) or core areas ( $\bar{x}$  < 0.01) of adjacent packs was low; and (2) the amount of shared space use was negligible. Spatial segregation did not vary substantially in relation to genotypes of adjacent packs or local environmental conditions (i.e., harvest regulations or road densities). We provide the first telemetry-based demonstration of spatial segregation between wolves and coyotes, highlighting the novel relationships between Canis types in the Ontario hybrid zone relative to areas where wolves and coyotes are reproductively isolated. Territoriality among Canis may increase the likelihood of eastern wolves joining coyote and hybrid packs, facilitate hybridization, and could play a role in limiting expansion of the genetically distinct APP eastern wolf population.