Testing Global Positioning System Telemetry to Study Wolf Predation on Deer Fawns

ABSTRACT We conducted a pilot study to test the usefulness of Global Positioning System (GPS) collars for investigating wolf (Canis lupus) predation on white-tailed deer (Odocoileus virginianus) fawns. Using GPS collars with short location-attempt intervals on 5 wolves and 5 deer during summers 2002–2004 in northeastern Minnesota, USA, demonstrated how this approach could provide new insights into wolf hunting behavior of fawns. For example, a wolf traveled ≥1.5–3.0 km and spent 20–22 hours in the immediate vicinity of known fawn kill sites and ≥0.7 km and 8.3 hours at scavenging sites. Wolf travel paths indicated that wolves intentionally traveled into deer summer ranges, traveled ≥0.7–4.2 km in such ranges, and spent <1–22 hours per visit. Each pair of 3 GPS-collared wolf pack members were located together for ≤6% of potential locations. From GPS collar data, we estimated that each deer summer range in a pack territory containing 5 wolves ≥1 year old and hunting individually would be visited by a wolf on average every 3–5 days. This approach holds great potential for investigating summer hunting behavior of wolves in areas where direct observation is impractical or impossible.     

Assessment of Shock Collars as Nonlethal Management for Wolves in Wisconsin

ABSTRACT Lethal control alone has not proven entirely effective in reducing gray wolf (Canis lupus) depredations in chronic problem areas. Opponents of lethal control argue that more emphasis should be placed on integrating nonlethal strategies into current management. However, few evaluations have tested the effectiveness of nonlethal options. We compared behavior patterns in terms of frequency and duration of bait station visits for 5 wolves fitted with shock collars to 5 control animals inhabiting wolf pack territories in northern Wisconsin during summers of 2003 and 2004. Shock collared wolves spent less time and made fewer visits to bait station zones than did control animals. During and after shocking, wolves shifted 0.7 km away from the bait station zone. Although active shocking did restrict wolf access, which could be useful in controlling wolf depredations during a limited time period, conditioning was not clearly demonstrated once shocking ceased. The effect of shock collar design and operation on long-term conditioning and shock-conditioned wolves on pack behavior needs further study. If long-term conditioning is possible, shock collars could be used by wildlife managers as a nonlethal wolf management method in chronic problem areas where lethal control has proven ineffective.     

Evidence of wolf dispersal in anthropogenic habitats of the Polish Carpathian Mountains

In the course of their maturation, most young wolves leave their natal pack and disperse in search for mating partners, improved food availability and new territories. We investigated whether this dispersal is affected by anthropogenic infrastructure in a 5,000 km2 area of the eastern region of the Polish Carpathian Mountains occupied by wolves. A radio-collared male wolf covered 230 km while dispersing through forested hills and densely populated valleys. To test if such dispersal is common in the population we analysed by microsatellite genotyping 39 samples taken from live-trapped wolves or wolves found dead in the study area. Although the obtained genotypes were assigned to different clusters in Bayesian tests, we could not ascribe this structure to landscape features, but rather to shared ancestry of wolf individuals found in distant locations. Moreover, we could not detect a spatial genetic structure in the wolf population, indicating a random occurrence of genotypes within the study area. Observation of the dispersing wolf and the absence of spatial genetic structure imply that wolves are still able to roam the entire area despite high densities of roads and a dense human population. Thus, we concluded that the existing anthropogenic infrastructure does not restrict wolf dispersal in the area and the studied wolves represent a coherent part of the Polish Carpathian wolf population.     

Multistage,Long-Range Natal Dispersal by a Global Positioning System-Collared Scandinavian Wolf

Abstract: We document a new record dispersal for wolves worldwide. The natal straight-line dispersal distance of a Global Positioning System-collared female wolf from the Scandinavian population was 1,092 km from southeast Norway to northeast Finland, with a multistage actual travel distance of >10,000 km. Natural gene flow to the isolated, inbred Scandinavian wolf population may occur if survival of dispersers is improved.     

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.     

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.     

Insular and disjunct distribution of the Arctic wolf in Greenland, 1978�C1998

Little research has been conducted on the spatial and temporal distribution of wolves in the High Arctic, and the processes that influence distributional patterns are not well understood. The present study addressed this information void in north and east Greenland by determining bi-seasonal distribution patterns and core areas based upon 303 sightings of wolves or their tracks during 1978–1998. The data suggested that this wolf population were predominantly distributed in semi-isolated patches in an insular and disjunct distribution. Evidence of wolf occurrence in some areas was so irregular that wolves should be considered absent in most years. Where such areas were clustered over several hundreds of kilometers, they collectively constituted areas of general wolf absence that could be considered gaps in distribution. There was no evidence that this population functioned in a mosaic of closely, interlocking pack territories similar to those reported in lower latitudes. The observed distributional pattern likely reflected the exceptionally impoverished and fragmented polar desert and semi-desert habitat that compelled wolves to adopt differing spatial distribution patterns relative to wolves in temperate areas. The broader scientific significance of this study is that, even in the most remote, vast, and uninhabited regions, it is possible to produce meaningful results on distributional patterns of rare, terrestrial carnivores that can serve as foundation for the next generation of testable hypotheses.     

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.     

Effects of Wolves on Elk and Cattle Behaviors: Implications for Livestock Production and Wolf Conservation

Background

In many areas, livestock are grazed within wolf (Canis lupus) range. Predation and harassment of livestock by wolves creates conflict and is a significant challenge for wolf conservation. Wild prey, such as elk (Cervus elaphus), perform anti-predator behaviors. Artificial selection of cattle (Bos taurus) might have resulted in attenuation or absence of anti-predator responses, or in erratic and inconsistent responses. Regardless, such responses might have implications on stress and fitness.

Methodology/Principal Findings

We compared elk and cattle anti-predator responses to wolves in southwest Alberta, Canada within home ranges and livestock pastures, respectively. We deployed satellite- and GPS-telemetry collars on wolves, elk, and cattle (n = 16, 10 and 78, respectively) and measured seven prey response variables during periods of wolf presence and absence (speed, path sinuosity, time spent head-up, distance to neighboring animals, terrain ruggedness, slope and distance to forest). During independent periods of wolf presence (n = 72), individual elk increased path sinuosity (Z = −2.720, P = 0.007) and used more rugged terrain (Z = −2.856, P = 0.004) and steeper slopes (Z = −3.065, P = 0.002). For cattle, individual as well as group behavioral analyses were feasible and these indicated increased path sinuosity (Z = −2.720, P = 0.007) and decreased distance to neighbors (Z = −2.551, P = 0.011). In addition, cattle groups showed a number of behavioral changes concomitant to wolf visits, with variable direction in changes.

Conclusions/Significance

Our results suggest both elk and cattle modify their behavior in relation to wolf presence, with potential energetic costs. Our study does not allow evaluating the efficacy of anti-predator behaviors, but indicates that artificial selection did not result in their absence in cattle. The costs of wolf predation on livestock are often compensated considering just the market value of the animal killed. However, society might consider refunding some additional costs (e.g., weight loss and reduced reproduction) that might be associated with the changes in cattle behaviors that we documented.     

Long-distance dispersal of a wolf, <Emphasis Type="Italic">Canis lupus</Emphasis>, in northwestern Europe

Several mammal species have recolonized their historical ranges across Europe during the last decades. In November 2012, a wolf-looking canid was found dead in Thy National Park (56° 56′ N, 8° 25′ E) in Jutland, Denmark. DNA from this individual and nine German wolves were genotyped using a genome-wide panel of 22,163 canine single nucleotide polymorphism (SNP) markers and compared to existing profiles based on the same marker panel obtained from northeastern Polish (n?=?13) wolves, domestic dogs (n?=?13) and known wolf-dog hybrids (n?=?4). The Thy canid was confirmed to be a wolf from the German-western Polish population, approximately 800 km to the southeast. Access to the German reference database on DNA profiles based on 13 autosomal microsatellites of German wolves made it possible to pinpoint the exact pack origin of the Thy wolf in Saxony, Germany. This was the first documented observation of a wolf in Denmark in 200 years and another example of long-distance dispersal of a carnivore.     

Can Occupancy–Abundance Models Be Used to Monitor Wolf Abundance?

Estimating the abundance of wild carnivores is of foremost importance for conservation and management. However, given their elusive habits, direct observations of these animals are difficult to obtain, so abundance is more commonly estimated from sign surveys or radio-marked individuals. These methods can be costly and difficult, particularly in large areas with heavy forest cover. As an alternative, recent research has suggested that wolf abundance can be estimated from occupancy–abundance curves derived from “virtual” surveys of simulated wolf track networks. Although potentially more cost-effective, the utility of this approach hinges on its robustness to violations of its assumptions. We assessed the sensitivity of the occupancy–abundance approach to four assumptions: variation in wolf movement rates, changes in pack cohesion, presence of lone wolves, and size of survey units. Our simulations showed that occupancy rates and wolf pack abundances were biased high if track surveys were conducted when wolves made long compared to short movements, wolf packs were moving as multiple hunting units as opposed to a cohesive pack, and lone wolves were moving throughout the surveyed landscape. We also found that larger survey units (400 and 576 km²) were more robust to changes in these factors than smaller survey units (36 and 144 km²). However, occupancy rates derived from large survey units rapidly reached an asymptote at 100% occupancy, suggesting that these large units are inappropriate for areas with moderate to high wolf densities (>15 wolves/1,000 km²). Virtually-derived occupancy–abundance relationships can be a useful method for monitoring wolves and other elusive wildlife if applied within certain constraints, in particular biological knowledge of the surveyed species needs to be incorporated into the design of the occupancy surveys. Further, we suggest that the applicability of this method could be extended by directly incorporating some of its assumptions into the modelling framework.     

新疆卡拉麦里山有蹄类自然保护区冬季狼的家域

正狼(Canis lupus)是食肉目(Carnivora)、犬科(Canidae)、犬属(Canis)食肉类哺乳动物(盛和林等,1999),是世界上分布最广和研究最多的大型食肉动物(Ripple et al.,2014)。目前,我国狼的分布已由过去的全国性分布缩小至东北、西北和内蒙等地区(Zhang,1999)。在斯洛伐克,研究人员通过无线电项圈获取分布在两个国家公园内两群狼的家域利用情况(Findo and Chovancova,2004),加     

Evidence of long-distance dispersal of a gray wolf from the Chernobyl Exclusion Zone

The Chernobyl Exclusion Zone (CEZ) is a ~?4300 km² area in Belarus and Ukraine that remains heavily contaminated with radiation from the nuclear accident of 1986. Long standing controversy persists on the fate of wildlife within the CEZ following human abandonment of the area. Human residency remains extremely sparse, and the CEZ has become a refuge for some populations of wildlife, including gray wolves (Canis lupus). Using GPS telemetry, we documented the first long-distance movements of a young (1–2 years) male wolf from the CEZ into the surrounding landscape. The wolf traveled 369 km from its home range center over a 21-day period in February 2015. In the 95 days prior to dispersal, the wolf maintained a home range of ~?28 km², with daily displacements rarely exceeding 5 km. With the onset of dispersal, daily displacement increased to a mean of 16.8 km. The dispersal of a young wolf is an important observation because it suggests that the CEZ may serve as a source for some wildlife populations outside of the CEZ, and raises questions about the potential spread of radiation-induced genetic mutations to populations in uncontaminated areas.     

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     

Home range, activity and movements of a wolf pack in central Italy

Home range, habitat use, activity and movement patterns were studied in a pack of wolves in a mountainous region of Abruzzo, central Italy from June 1986 to March 1987. The home range, estimated by the minimum convex polygon from 421 radio locations, measured 197 km² and comprised several infrastructures and areas of human presence, including four garbage dumps and two offal sites. Core areas, calculated by the harmonic mean method, were located toward the centre of the home range where human disturbance and road density were lowest but forest cover was highest. During the time-span of the study, home-range use and movement patterns suggested a marked centrality in spatial behaviour and traditionality in retreat areas year-round, both during pup-rearing season and the following months. In addition, by being essentially nocturnal, resident wolves appeared to adopt tactics of temporal segregation from people to exploit food resources safely in the proximity of human settlements. Overall activity correlated with distance travelled (r = 0.90, P << 0.001), and corresponded to cyclic nocturnal movements from retreat to feeding areas. Wolf movement rate between 20:00 and 04:00 h averaged 2.5 km/h but varied up to about 8 km/h, and daily distance travelled (= 27 km/night; range 17–38 km/night) mostly depended on the location of traditional feeding sites. Home-range configuration, habitat use, activity and movements all appeared highly integrated so as to represent the most functional compromise between avoidance of human inteference and exploitation of the available food resources.     

Hypoxia Adaptations in the Grey Wolf (Canis lupus chanco) from Qinghai-Tibet Plateau

The Tibetan grey wolf (Canis lupus chanco) occupies habitats on the Qinghai-Tibet Plateau, a high altitude (>3000 m) environment where low oxygen tension exerts unique selection pressure on individuals to adapt to hypoxic conditions. To identify genes involved in hypoxia adaptation, we generated complete genome sequences of nine Chinese wolves from high and low altitude populations at an average coverage of 25× coverage. We found that, beginning about 55,000 years ago, the highland Tibetan grey wolf suffered a more substantial population decline than lowland wolves. Positively selected hypoxia-related genes in highland wolves are enriched in the HIF signaling pathway (P = 1.57E-6), ATP binding (P = 5.62E-5), and response to an oxygen-containing compound (P≤5.30E-4). Of these positively selected hypoxia-related genes, three genes (EPAS1, ANGPT1, and RYR2) had at least one specific fixed non-synonymous SNP in highland wolves based on the nine genome data. Our re-sequencing studies on a large panel of individuals showed a frequency difference greater than 58% between highland and lowland wolves for these specific fixed non-synonymous SNPs and a high degree of LD surrounding the three genes, which imply strong selection. Past studies have shown that EPAS1 and ANGPT1 are important in the response to hypoxic stress, and RYR2 is involved in heart function. These three genes also exhibited significant signals of natural selection in high altitude human populations, which suggest similar evolutionary constraints on natural selection in wolves and humans of the Qinghai-Tibet Plateau.     

Effect of sociality and season on gray wolf (Canis lupus) foraging behavior: implications for estimating summer kill rate

Background

Understanding how kill rates vary among seasons is required to understand predation by vertebrate species living in temperate climates. Unfortunately, kill rates are only rarely estimated during summer.

Methodology/Principal Findings

For several wolf packs in Yellowstone National Park, we used pairs of collared wolves living in the same pack and the double-count method to estimate the probability of attendance (PA) for an individual wolf at a carcass. PA quantifies an important aspect of social foraging behavior (i.e., the cohesiveness of foraging). We used PA to estimate summer kill rates for packs containing GPS-collared wolves between 2004 and 2009. Estimated rates of daily prey acquisition (edible biomass per wolf) decreased from 8.4±0.9 kg (mean ± SE) in May to 4.1±0.4 kg in July. Failure to account for PA would have resulted in underestimating kill rate by 32%. PA was 0.72±0.05 for large ungulate prey and 0.46±0.04 for small ungulate prey. To assess seasonal differences in social foraging behavior, we also evaluated PA during winter for VHF-collared wolves between 1997 and 2009. During winter, PA was 0.95±0.01. PA was not influenced by prey size but was influenced by wolf age and pack size.

Conclusions/Significance

Our results demonstrate that seasonal patterns in the foraging behavior of social carnivores have important implications for understanding their social behavior and estimating kill rates. Synthesizing our findings with previous insights suggests that there is important seasonal variation in how and why social carnivores live in groups. Our findings are also important for applications of GPS collars to estimate kill rates. Specifically, because the factors affecting the PA of social carnivores likely differ between seasons, kill rates estimated through GPS collars should account for seasonal differences in social foraging behavior.     

Some Aspects of the Population Ecology of Wolves, Alaska

Information on Alaskan wolf populations was obtained from examinationof bounty records, 4,150 wolf radii and ulnae, 1,262 wolf carcasses,and from observations of wolves inhabiting an area of 20,000square miles where wolves were protected. Pregnant adult female wolves averaged 6.5 fetuses; two-year-oldfemales averaged 5.3 fetuses;female pups were not sexually mature. In Alaska, wolves conceive from late February through earlyApril but most females breed in March. Multiparous females breedearlier than first breeders. Multiparous females produce anaverage of 7.3 ova and 6.5 implanted fetuses. The loss of ovafrom ovulation to implantation is significant. Multiparous femalesproduce more ova than first breeders; the difference is highlysignificant. Mortality of pups rather than the lack of initial productionof pups is believed to be the reason for the observed variationsin the proportion of pups in wolf populations. Wolf packs includemembers in all categories of sex and age during the breedingseason. Size of the pack is an indicator of abundance. Wolves in an area where they were protected increased at anaverage rate of 20–30% per year during an 11-year period.     

Fine-scale genetic structure suggests low levels of short-range gene flow in a wolf population of the Italian Apennines

We investigated local gene flow in a high-density wolf (Canis lupus) population of the Italian Apennines, where no effective barrier to wolf dispersal was present. From 1998 to 2004 we examined wolf carcasses and non-invasively collected samples, focusing on three mountain districts, separated by two valleys, where wolf packs showed high spatial stability. Using nine autosomal microsatellites we successfully genotyped 177 samples, achieving the identification of 74 wolves. Genetic relatedness steeply decreased with increasing distance between sampling areas, thus suggesting that short-distance interpack migration is infrequent in this population. In addition, no individual from a central pack under intensive monitoring was sampled in the range of the surrounding packs over a 4-year period. The limited short-distance gene flow resulted in a cryptic genetic structure, which was revealed by Bayesian analysis. A different genetic cluster was found in each of the three mountain areas, and a small proportion of first-generation immigrants was detected. Overall, the present study suggests that local genetic differentiation in Italian wolves might arise from high spatial stability of packs and can be favoured by a combination of long-range dispersal, the attitude to mate between unrelated individuals and a high young mortality rate.     

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.