Wolf History, Conservation, Ecology and Behavior
Wolf (Canis lupus) Diet in Latvia: Seasonal, Geographical and Sexual Variations
Andersone, Z., Acta Zoologica Lituanica, 13/1 (March 2003)
Diet of wolves (Canis lupus) was studied in Latvia from 1997 to 2001. In total, 302 scats and 107 stomachs of wolves from different parts of Latvia were analysed. Prey species were microscopically identified by their hair structure. In order to avoid bias due to the similar hair structure, cervids -- moose (Alces alces), red deer (Cervus elaphus) and roe deer (Capreolus capreolus) were pooled together. Wild ungulates -- cervids, wild boar (Sus scrofa) and beaver (Castor fiber) -- were found to be the staple food for wolves in Latvia. Cervids were found in 50% of samples (62% of biomass), wild boar were found in 24.7% (21% of biomass), and beavers were found in 13.7% (11.5% of biomass). Beaver was an important prey in summer, its significance being much higher than elsewhere in Europe. Beaver may have been a buffer prey, assisting to maintain high wolf densities during the early 1990s, when the ungulate population decreased. Thus, wild boar was positively selected, especially in winter when its ratio in the diet increased to 33.9% from 20% in summer. It was more a common prey in the east of the country. The ratio of beavers, small rodents and plant food was higher in summer, which resulted in a greater food niche breadth in summer than winter food niche breadth .... However, the percentage of the biomass consumed was dominated by ungulates in both seasons. The role of domestic animals in the wolf diet was insignificant except for winter when they were consumed as carrion (13.1%). Livestock depredation was rather a seasonal and local problem that could have been prevented in most cases should the proper husbandry techniques have been used. Minor sexual differences in the diet of wolves were found: males consumed beavers considerably more often .... Thus, it should be concluded that in Latvia wolves prey generally on wild animals, and conflicts with livestock owners are only local, the main source of conflict being competition for the ungulate prey with hunters.
Winter Habitat and Travel Route Selection by Wolves in the Northern Appenines, Italy
Ciucci, P.; Masi, M.; Boitani, L, Ecography, 26/2 (April 2003), 223 (13pp)
To assess the wolf-habitat relationship on the home range scale, we developed a resource selection probability function (RSPF) through a multiple logistic regression model based on the winter travel routes of a wolf pack in the northern Appenines, Italy (1991-95). Both travel routes (240 km) and habitat variables were mapped at 1:10 000 scale, digitised as Geographical Information System (GIS) layers, and overlaid with a 100 x 100 m pixel grid to census all used and unused resource units....According to the model, travel routes by wolves were not randomly located within the home range but were clearly associated with selected bio-physical factors, including human-related habitat modifications (i.e., roads), which appeared to affect the wolves' resource selection and, ultimately, habitat quality....Although wolves are generally considered habitat generalists, this study shows that patterns of habitat selection are disclosed at finer scales of analysis. In this perspective, resource selection probability functions at finger scales offer different and complementary insight with respect to regional landscape applications, and provide a useful management tool for assessment of habitat quality at the local scale.
Wolf at the Gate: The Anti-Carnivore Alliance and the Symbolic Construction of Community, A
Skogen, K.; Krange, O., Sociologia Ruralis, 43/3 (July 2003) 309 (17pp)
Controversies over the return of large carnivores (e.g. wolves) are often interpreted as clashes between rural traditionalism and urban modernity. Rural communities, however, have never been culturally monolithic, and modernization increases their diversity. However, the popular image is one of rural communities united against vermin and urban romantics. An important reason for this is probably the successful construction of the anti-carnivore front as a last line of defence against destructive forces threatening rural life. Drawing on examples from a study in Østerdalen, Norway, the struggle against wolf protection is discussed as an instance of symbolic construction of community. Images of a threatened community are vital to the self-understanding of the wolf adversaries, but cleavages run through the alliance. Three principal groups may be identified: sheep farmers, landowners who lease hunting, and people with strong ties to traditional land use practices (primarily hunting) and a rural working-class culture. These groups have not always been allies, and conflicts of interest run through the ‘resistance front’. The task here is to identify the social forces that now bring them together, and to explain why the carnivore issue is well suited as a significant component in their symbolic construction of community.
Wolf Depredation on Domestic Dogs in Central Part of European Russia
Bologov, V.V.; Miltner, D., World Wolf Congress 2003
Wolf (Canis lupus) depredation on domestic animals, i.e. livestock and pets, has occurred throughout the world probably ever since and wherever these species have coexisted. The subject has been studied quite intensively since depredation on domestic animals is a major challenge to human tolerance of the wolf and will always influence the relationship between the two where wilderness and agriculture/human settlements meet. Due to importance most studies conducted so far in terms of wolf management and control programs focused on the depredation of livestock. But wolves also kill considerable numbers of pets, especially dogs (Canis familiaris), though the livestock-pet ratio is usually clearly biased towards livestock. In Central European Russia however depredation is more frequent on dogs than on livestock....[W]e collected data about 123 attacks of wolves on dogs, dating from July 1996 to March 2001. Of these, 117 dogs were killed (95.12%) and 6 (4.88%) survived the attack; three survived because the owner drove the wolves off. One dog was attacked twice in one year. It survived the first assault but was killed the second time. Including livestock, casualties of domestic animals total 146 animals. Besides the 123 attacks on dogs (84.25%), there were 14 sheep (9.6%), 6 cows (4.1%), two goats (1.37%) and one cat (0.68%) killed.
Wolf Functional Response and Regulation of Moose in the Yukon
Hayes, R.; Harestad, A., Canadian Journal of Zoology, 78 (January, 2000): 60-66 (7 pp)
We studied kill rates by wolves (Canis lupus) on a rapidly growing moose population in the east-central Yukon. We added these data to the cumulative functional response curve obtained in other North American wolf studies. Our kill rates are higher than those predicted at low moose densities. The kill rate increases rapidly, reaching 2.4 moose per wolf per 100 days at 0.26 moose/km2 and remains constant at this level. No data are available below 0.2 moose/km2 to indicate the shape of the ascending curve. Based on moose distribution and the low prey-switching ability of wolves, we suggest that the functional response curve is of type II. Our wolf predation rate model predicts that moose are held to a low density equilibrium between 0.07 and 0.12/km2, slightly below densities observed in interior Alaska and the Yukon.
Wolf (Canis lupus) Numbers, Diet and Damage to Livestock in Relation to Hunting and Ungulate Abundance in Northeastern Belarus during 1990-2000
Sidorovich, V.E.; Tikhomirova, L.L.; Jedrzejewska, B., Wildlife Biology, 9/2 (June 2003), 103-111
Wolf Canis lupus relationships with wild ungulates, domestic animals and humans were studied in an area of ca 800 km(2) at the head of the Lovat River in northeastern Belarus during 1990-2000. The region was dominated by natural habitats (78%) consisting mainly of forests and bogs, but also lakes and rivers. The abundance of wild ungulates, such as moose Alces alces, wild boar Sus scrofa, and roe deer Capreolus capreolus, as censused by snow tracking and assessed by game wardens, declined 5 to 6-fold between 1990 and 1996, most probably due to uncontrolled exploitation and poaching. During 1997-2000, the numbers of ungulates began to recover. Wolves responded to the shortage of wild ungulates by a strong shift in feeding habits. When wild ungulates were numerous, wolf diet as studied by scat analysis was composed of wild ungulates (80-88% of consumed biomass), with small additions of medium- and small-sized wild animals (7-13%), mainly beaver Castor fiber and hare Lepus sp., and domestic animals (4-6%), mainly cattle. In the years when the recorded numbers of wild ungulates were at their lowest, wolves preyed on domestic animals (38% of biomass consumed), wild ungulates (32%), and medium- and small-sized wild prey (29%). Wolf damage to domestic animals (28 head of cattle and 247 dogs killed) and wolf-human interaction (100 observations of wolves in and near villages, including one attack by a rabid wolf on 1 people) were recorded in 14 villages. The rate of wolf predation on domestic animals and their appearances in villages increased exponentially with the declining biomass of wild ungulates and ceased again when wild ungulates began to recover; a one-year time lag in wolf response to changes in ungulate abundance was observed. The numbers of wolves as estimated by snow tracking and assessed by game wardens played a weaker role in shaping wolf-livestock and wolf-human interaction. The wolf population was strongly affected by hunting during the study. Wolves responded numerically with a 1 to 2-year time lag to the varying intensity of harvest by humans. Our study showed the role of the human factor in shaping wolf numbers and wolf-livestock interaction in eastern Europe. The three major components of this relationship were: 1) the manifold decline in wild ungulate abundance, which was most probably caused by uncontrolled exploitation by humans in the years of political transformation and economic regress, made wolves shift to predation on domestic animals; inevitably, wolves were frequently seen in the rural areas; 2) people interpreted the growing rates of wolf damage and appearances near the settlements as an effect of greatly increasing numbers of wolves, and demanded that authorities and hunters fight the 'wolf plague'; 3) hunting impact on wolves increased and led to a marked reduction in wolf numbers and a decline in wolf-human conflicts. This scenario was most probably repeated in many areas of eastern Europe during 1990-2000, which was a decade of political and economical transformation. From a management perspective, we suggested that predation levels and wolf-human conflicts could be reduced not only by increased wolf harvest but also by enhancing the density and diversity of wild ungulates.
Wolf Pack Territory Marking in the Bialowieza Primeval Forest (Poland)
Zub, K. ; Theuerkauf, J. ; Jedrzejewski, W. ; Jedrzejewska, B. ; Schmidt, K. ; Kowalczyk, R., Behaviour, 140/5 (2003)
We analysed data on territory marking with urine, scats, and ground scratching by wolves (Canis lupus) belonging to four packs in the Bialowieza Primeval Forest, Poland. The aims were to determine: (1) seasonal variation in the marking rates, (2) significance of various kinds of marking in territory demarcation, and (3) relationship between spatial distribution of wolves' marking and their use of territory. Continuous radio-tracking and subsequent snow tracking of the collared wolves were the main methods. Deposition rates of scats showed little variation in time and space, whereas rates of urine marking and ground scratching showed large seasonal and spatial variation. Wolf marking rates with urine and ground scratching were highest during the cold season (October-March) and peaked during the mating season, in January and February. Marking intensity did not grow with the number of wolves in a pack, and per capita rates of marking were highest in wolves travelling singly or in pairs. Mean marking rates per km of wolf trail were low in the core areas of territories, and increased when wolves approached the boundaries. However, densities of marks (number of marks per square km) increased in territory centre (due to intense use of core area by the pack), and in peripheral areas, which bounded other territories (due to increased marking activity by wolves when moving along the territory edge). Our findings did not support the 'olfactory bowl' model of wolf territory marking, as marks were not distributed equally along territory boundaries. Instead, marks were concentrated in 'hot spots' more vulnerable to penetration by intruders (territory edge) or more valuable to owners (vicinities of breeding dens).
Wolf Predation and the Exotic Disease Debate in Wood Buffalo National Park
Carbyn, Lu, World Wolf Congress 2003
Significant declines in bison have occurred within Wood Buffalo National Park. Numbers declined from about 11,000 in 1971 to about 2,300 in 1998. Prior to 1971, herds had increased from an estimated low of about 500 around the turn of the century to about 15,000 in the mid 1950s. Exotic bovine diseases were introduced to the park from 1925-1928. This paper evaluates the ramifications of wolf predation and disease in bison declines. The predation hypothesis involves the notion that predators alone have caused the decline. A competing hypothesis is that diseases reduce bison calf production and adult survival, thus shifting bison numbers from food competition equilibrium to low density predator regulated equilibrium. Field studies from 1985 to 1999 indicated low cow:calf ratios at about 30 calves per 100 cows. I have concluded that in the presence of both exotic bovine diseases (tuberculosis and brucellosis) and wolf predation, bison numbers are kept at low levels. In the absence of wolves, it is likely that, even in the presence of diseases, bison numbers will increase. My conclusion is that of the two factors, wolf predation has a greater influence on overall numbers than exotic diseases.
Wolf Predation Damages to Livestock, The Golan, Israel
Gilady, P., Beyond 2000: Realities of Global Wolf Restoration (2000)
The Golan Heights, approximately 1,000 km2 (390 sq. mi.), are located in the northern part of Israel, east of the Sea of Galilee. The livestock population there includes about 5,000 ewes and 12,000 cows on 350 km2 (140 sq. mi.) Adult male wolves (Canis lupus pallipes) in Israel weigh about 26-30 kg. (55-70 lb.), while wolves of the Golan population are slightly larger, similar to European wolves. The wolves in Israel are in danger of extinction, and their population size is "guesstimated" to be 150-200 individuals, including 100 in the Golan. Wolves have been protected in Israel since 1954, but are culled due to livestock predation....In recent years, genetic, demographic and behavioral research has been conducted on the Golan's wolf population. According to ranchers' estimates, wolf predation damage to livestock during 1998-99 was approximately $280,000. Predation affected about 6% of all newborn livestock. Identification of the predator was done via inspection of the remains, tracks, teeth marks, biting location and the description of the event by the rancher. It was not always possible to determine the cause, whether it was predation, or the predator's species. The following methods are used to try to minimize wolf depredations on livestock: (1) Two breeds of guard doggs are raised in Israel, the Great Pyrenees and the Maremma Sheep Dog, and recently the Turkish Akbash has been tested. In general, dog presence with the herd may reduce predation. (2) In spite of guard dog presence, it is still difficult to eliminate predation, so some ranchers fence some pastures. We recommend electric fencing of birthing enclosures up to 200 hectares (500 acres) in size. A kilometer (0.6 mi.) of electric fencing costs $800. (3) Controlled hunting of specific wolves is conducted only by wildlife rangers or hunters with special permits, and only during attempted predation. (4) Marking birthing enclosures every two or three days with dogs urinating or defecating along enclosure perimeter. (5) Foot traps are used to capture wolves but only where wolf damage is observed. (6) When dens are located within paddocks, wolves are removed and transferred to local zoos or universities. (7) A governmental compensation fund provides partial compensation for predatory damages and financial assistance to purchase an electric fence and guard dogs. These measures attempt to find the Golden Path that minimizes conflicts between nature and humans.
Wolf Predation in A Multiple-Ungulate System in Northern British Columbia
Bergerud, A.; Elliot, J, Canadian Journal of Zoology, 76 (August, 1998): 1551-1569 (19 pp)
Caribou (Rangifer tarandus), elk (Cervus canadensis), moose (Alces alces), and Stone's sheep (Ovis dallistonei) were either decreasing or stable in numbers in two areas in northeastern British Columbia in 1981-1982, prior to reductions in wolf (Canis lupus) numbers. Following the reduction of wolf numbers, recruitment improved 2-5 times for all four species, and all populations increased, based on either hunting statistics, census results, and (or) recruitments greater than 24 offspring at 9 months of age per 100 females. Recruitment of offspring at 9 months of age, when regressed against wolf numbers, declined with decelerating slopes for all four species. This inverse functional response is hypothesized to result from the preparturient spacing of females to reduce predation risk, and in this regard moose seem the least secure and sheep the most effectively spaced. For the four species, mean recruitment at 9 months of age that balanced adult mortality and provided a finite rate of increase of 1.00 was 24.16 +or- 0.91 offspring/100 females (n = 11, coefficient of variation = 12.5%). The predicted recruitment rate for all four species in the absence of wolves was 53-57 offspring/100 females. But the birth rate of moose was much higher than those of the other species, indicating greater loss to other factors of which bear predation may be the greatest. Following wolf reductions of 60-86% of entire travelling packs, the wolves quickly recolonized the removal zones, with rates of increase ranging from 1.5 to 5.6.
Wolf Predation on Blackbuck (Antilope cervicapra) in the Great Indian Bustard Sanctuary, Nannaj, Maharashtra, India
Kumar, S.; Rahmani, A.R., Beyond 2000: Realities of Global Wolf Restoration (2000)
Aspects of predation on blackbuck (Antilope cervicapra), a primary prey of the Indian grey wolf (Canis lupus pallipes) were studied in the Great Indian Bustard Sanctuary from 1991 to 1994. For each kill, data were collected on a) sex and age, b) habitat around kill site, c) biomass left unconsumed, d) distance of kill from the den(s) and nearest vegetative cover. The wolves largely preyed on old and injured blackbuck and had a strong selection for males. On an average, wolves made a kill every 3.65 days during winter and 2.1 days during summer. The consumption rate was found to be 1 kg/wolf/day and it was not correlated with pack size. The wolves depended largely on blackbuck for food during their non-breeding period and on livestock during denning and breeding period. The maximum number of kills were located within 4 m of distance from vegetative cover. The distribution of kills differed significantly between habitats and the maximum number of kills were found in grasslands (37%), followed by scrubland (22.7%), plantations (21.3%) and grazing land (18.7%). The wolves killed blackbuck irrespective of the location of dens. Only two instances of food caching by wolves were recorded during the study period. The wolves removed annually about 4% of the total blackbuck biomass available to them in the Sanctuary. Blackbuck used predator avoidance strategies such as encirclement of herds by large males with longest horns to scare wolves, ground stumping and grunting, flashing tail while running, and galloping as high as possible when the flight distance between the predator and the prey was short.
Wolf Recolonization of Agricultural Areas in Spain
Blanco, J.C.; Cortes, Y., Beyond 2000: Realities of Global Wolf Restoration (2000)
Spain's wolves have been recovering since the seventies, and over the last twenty years have colonised flat, densely populated agricultural areas with hardly any wild ungulates, which used to be considered unsuitable for them. Over the last ten years, the building of many fenced motorways has increased concern about the future of the wolf in these areas. Between 1997 and 1999 we radiocollared 11 wolves in farming areas in central and western provinces of Valladolid and Zamora. These wolves live in packs, some comprising as many as ten. They usually breed in island patches of woodland of between 15-35 km2, about 30 km apart and surrounded by arable land; some packs, however, breed among extensive cereal fields with no tree cover whatsoever. Territories adjoin or overlap, making it possible to rule out the idea that the populations are fragmented. Wolves use overhead asphalted bridges built for traffic, the only available structures, to cross the fenced, four-lane motorways. Three resident wolves crossed a motorway on 4.4%-8.7% of tracking days, while the figure for two floaters was 22.2%-23.0%. Domestic animals (mainly sheep scavenged in carrion pits) made up the greater part of the wolves' diet (77% of biomass), while rabbits were the most common wild prey (11% of biomass). Damage to livestock is rare because the flocks of sheep are tended by shepherds throughout the day and kept indoors overnight. Despite being a game species, the annual mortality rate was only 13%, density (2.5-3 wolves/100 km2 in 1999) has risen considerably over the last ten years and the distribution area has increased. The study shows that, given a moderate level of persecution by people, wolves can thrive in very humanised farming areas that are almost devoid of wild prey.