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Wolf History, Conservation, Ecology and Behavior
Temporary Wolf Management Plan for Croatia
Huber, D. Beyond 2000: Realities of Global Wolf Restoration (2000)

After alarmingly low wolf population indices in Croatia in the early 1990s, a campaign to protect the species succeeded by May 1995 when the animal was protected: (1) any killing or disturbance was prohibited (a fine of nearly USD 6000 was established), and (2) any proven damage done by wolves would be compensated by the state. The first provision never worked, as the killings even increased compared to the years before protection: 24 bodies retrieved by the author's lab, at least twice as many actually killed after protection as before. No one was ever fined. Secondly, the state paid over USD $200,000 for damage claims that showed a dramatic increase: 9 in 1995, 176 in 1996, 516 in 1997 and 669 through November 1998. The need for a wolf management plan for Croatia became urgent. Newly acquired data showed that the wolf is present throughout Zone A, an area of 17,270 km2 (6,746 mi2) which was divided into the Zone A1 where wild prey predominates as wolf food and Zone A2 where wolves feed predominantly on livestock. The area of occasional wolf presence is 6,840 km2 (2,672 mi2), and the area presently without wolves is 29,212 km2 (11,411 mi2). Despite high mortality, the estimated 50 wolves in the early 1990s might have increased to over 100 individuals. Possible reasons are wolf reproductive success during the Croatian war due to the abundance of abandoned livestock and due to immigration from outside Croatia. Although the wolf population of this size should still be considered threatened and legally protected, the noted trend permits more flexible management. Due to the lack of knowledge about Croatian wolves, the high illegal kill, and the increasing trend of wolf damages, a temporary wolf management plan was instituted for the period of 1 January 1999 through 31 December 2000. Long and short-term measures were initiated. As a short-term measure for Zone A1, wold reduction will be considered in the case of high losses of trophy wildlife and the scientifically proven stable local wolf population. For Zone A2, an area of 3,177 km2 (1,241 mi2) was defined where 75% of wolf depredations have been claimed. In this area the possibility of permitting local, limited wolf killing to protect livestock is anticipated; in the same period, wolf damage compensations would not be paid. Part of the strategy is to prepare conditions, by 2001, for paying subsidies instead of damage compensations for livestock raising in wolf range.
Threatened Ethiopian Wolves Persist in Small Isolated Afroalpine Enclaves
Marino, J., Oryx, 37/1, January 2003, 62-71

The Ethiopian wolf Canis simensis is endemic to the Afroalpine highlands of Ethiopia. Half of the world population, estimated at c. 500 individuals, lives in the Bale Mountains of southern Ethiopia. Little is known, however, about the presence of wolves and suitable habitat in other Afroalpine ranges. Assessing the distribution, abundance and threats to all extant populations is a conservation priority for this Critically Endangered canid. With these objectives in mind, surveys were conducted between 1997 and 2000 in the little known regions of Arsi, Wollo, Gondar and Shoa. Suitable habitat and resident wolves were found in all regions. Outside Bale the existence of six other isolated populations, including two previously undescribed, was confirmed. All were small, estimated at no more than 50 individuals, and some with fewer than 25 individuals. Two population extinctions were documented, and habitat loss to agriculture largely explained local extinctions in small habitat patches. While Bale remains crucial for the long-term persistence of this species, the finding of several small and threatened populations highlights the need for in situ conservation actions to be expanded to other regions of the Ethiopian highlands.
A Ten-Year History of the Demography and Productivity of an Arctic Wolf Pack
Mech, L.D., Arctic, 48/4, December 1995, 329-332

A pack of two to eight adult wolves (Canis lupus arctos) and their pups were observed during ten summers (1985-95) on Ellesmere Island, Northwest Territories, Canada. The author habituated the wolf pack to his presence in the first summer and reinforced the habituation each summer thereafter. The first alpha female produced four to six pups each year between 1986 and 1989. However, her daughter, who succeeded her as the alpha female, produced only one to three pups each year between 1990 and 1992 and in 1994, and apparently did not whelp in 1993 or in 1995. The tenure of the first alpha male was at least two years, and his successor was alpha male for the remaining eight years of the study. The wolf pack was characterized by highly variable annual productivity. The second alpha male-and-female breeding pair likely was an older brother and a younger sister. Early survival of wolf pups was high and constant, with all pups surviving through August of their first year. The pack's demography was consistent with what is known for wolf packs in other regions of North America, but its productivity was more typical of arctic packs.
Translocation of Wolves Associated with Livestock Depredation in a Recovering Population
Beyer, Jr., D.E.; Hammill, J.H.; Johnson, B.J.; Lonsway, D.H.; Roell, B.J., World Wolf Congress 2003.

Gray wolves (Canis lupus) have become reestablished in the Upper Peninsula of Michigan, through immigration of animals from the neighboring states of Minnesota and Wisconsin and the Canadian province of Ontario. The wolf population has increased from 3 animals in 1989 to 278 animals in 2002. Wolves in Michigan are currently listed as a federally endangered species. In 1998, the first case of wolf depredation on livestock was verified. The only management response to wolf depredation allowed under the federal endangered status is live-capture and relocation of nuisance animals. To date, we have verified 16 cases of wolf depredation on livestock. We have trapped and relocated 24 wolves at 5 farms that experienced wolf depredation. Translocated wolves were released with radiocollars an average of 124 km from the trapping location. None of the translocated wolves remained in the vicinity of their release site. Three wolves returned to their territories and 4 wolves settled into territories in new areas. Translocated wolves often moved long distances in short periods of time. Radio contact was lost with 8 wolves after an average of 58 days, although 3 of these animals were later recovered. Four wolves died before settling into a territory. The fate of 6 wolves trapped and translocated in 2002 is still being assessed. Regardless of their movements or fate, none of the translocated wolves were associated with livestock depredation after they had been moved. Survival of translocated wolves did not differ from wolves that were not moved.
The Trophic Ecology of Wolves and Their Predatory Role in Ungulate Communities of Forest Ecosystems in Europe
Okarma, H., Acta Theriologica, 40/4, December 1995, 335-386

Predation by ungulate populations in Europe, with special reference to the multi-species system of Bialowieza Primeval Forest (Poland/Belarus) was assessed on the basis [of results] of original research and literature. In historical times (post-glacial period), the geographical range of the wolf and most ungulate species in Europe decreased considerably. Community richness of ungulates and potential prey for wolves, decreased over most of the continent from 5-6 species to 2-3 species. The wolf is typically an opportunistic predator with a highly diverse diet; however, cervids are its preferred prey. Red deer Cervus elaphus are positively selected from ungulate communities in all localities; moose Alces alces are the major prey only where middle-sized species are scarce. Capreolus capreolus [roe deer] are locally preyed on intensively, especially where they have a high density, co-exist mainly with moose or wild boar Sus scrofa, and red deer is scarce or absent. Wild boar are generally avoided, except in a few locations; and European bison Bison bonasus are not preyed upon by wolves. Wolf predation contributes substantially to the total natural mortality of ungulates in Europe: 42.5% for red deer, 34.5% for moose, 25.7% for roe deer, and only 16% for wild boar. Food niche breadth of wolves in Europe, calculated only for the ungulates considered in this study, increases with the number of ungulate species in the community. There is also a significant relationship between ungulate community breadth and food niche breadth of wolves. Food niche breadth for wolves, however, does not achieve very high values even in the richest ungulate communities. Wolves easily adapt to locally abundant food of anthropogenic origin (livestock, garbage). The level of predation on livestock may be a result of different husbandry practices (e.g., use of livestock guarding dogs) rather than of differences in availability of wild and domestic prey. Available data from Europe suggests that wolves likely limit density of red deer and moose in some areas. Roe deer density can be decreased locally by wolves but is limited mainly by lynx. Wild boar density is more influenced by mast crops of Quercus spp. and Fagus silvatica (and to a lesser extent by snow depth) than by wolf predation.
Trophic Facilitation by Introduced Top Predators: Grey Wolf Subsidies to Scavengers in Yellowstone National Park
Wilmers, C.C.; Crabtree, R.L.; Smith, D.W.; Murphy, K.M.; Getz, W.M., Journal of Animal Ecology, 72/6, November 2003, 909-916

The reintroduction of grey wolves Canis lupus (L.) to Yellowstone National Park provides a natural experiment in which to study the effects of a keystone predator on ecosystem function. 2. Grey wolves often provision scavengers with carrion by partially consuming their prey. 3. In order to examine how grey wolf foraging behaviour influences the availability of carrion to scavengers, we observed consumption of 57 wolf-killed elk Cervus elaphus (L.) and determined the percentage of edible biomass eaten by wolves from each carcass. 4. We found that the percentage of a carcass consumed by wolves increases as snow depth decreases and the ratio of wolf pack size to prey size and distance to the road increases. In addition, wolf packs of intermediate size provide the most carrion to scavengers. 5. Applying linear regression models to the years prior to reintroduction, we calculate carrion biomass availability had wolves been present, and contrast this to a previously published index of carrion availability. Our results demonstrate that wolves increase the time period over which carrion is available, and change the variability in scavenge from a late winter pulse dependent primarily on abiotic environmental conditions to one that is relatively constant across the winter and primarily dependent on wolf demographics. Wolves also decrease the year-to-year and month-to-month variation in carrion availability. 6. By transferring the availability of carrion from the highly productive late winter, to the less productive early winter and from highly productive years to less productive ones, wolves provide a temporal subsidy to scavengers.
Two Centuries of the Scandinavian Wolf Population: Patterns of Genetic Variability and Migration During an Era of Dramatic Decline
Flagstad, Ø.; Walker, C.W.; Vila, C.; Sundqvist, A.-K,; Fernholm, B.; Hufthammer, A.K.; Wiig, Ø.; Koyola, I.; Ellegren, H., Molecular Ecology, 12/4 (April 2003), 869 (12pp)

The grey wolf (Canis lupus) was numerous on the Scandinavian peninsula in the early 19th century. However, as a result of intense persecution, the population declined dramatically and was virtually extinct from the peninsula by the 1960s. We examined historical patterns of genetic variability throughout the period of decline, from 1829 to 1979. Contemporary Finnish wolves, considered to be representative of a large eastern wolf population, were used for comparison. Mitochondrial DNA (mtDNA) variability among historical Scandinavian wolves was significantly lower than in Finland while Y chromosome variability was comparable between the two populations. This may suggest that long-distance migration from the east has been male-biased. Importantly though, as the historical population was significantly differentiated from contemporary Finnish wolves, the overall immigration rate to the Scandinavian peninsula appears to have been low. Levels of variability at autosomal microsatellite loci were high by the early 1800s but declined considerably towards the mid-20th century. At this time, approximately 40% of the allelic diversity and 30% of the heterozygosity had been lost. After 1940, however, there is evidence of several immigration events, coinciding with episodes of marked population increase in Russian Karelia and subsequent westwards migration.