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Seasonality and reoccurrence of depredation and wolf control in western North America

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Abstract

Due primarily to wolf (Canis lupus) predation on livestock (depredation), some livestock producers and other interest groups oppose wolf conservation, which is an important objective for large sectors of the public. Predicting depredation occurrence is difficult, yet necessary to prevent it. Better prediction of wolf depredation also would facilitate application of sound depredation management actions. In this paper we analyze temporal trends in wolf depredation occurrence and wolf control, which is employed as a depredation management action. We gathered data from wolf depredation investigations for Alberta, Canada, from 1982–1996 and for Idaho, Montana, and Wyoming, USA, from 1987–2003. We showed that wolf attacks occurred with a seasonal pattern, reflecting the seasonality of livestock calving, grazing practices, and seasonal variation in energetic requirements of wolf packs. Seasonal wolf attacks were auto-correlated with lags of one year, indicating annual reoccurrence. Cross-correlation analyses indicated that limited wolf control was rapidly employed as a short-term response to depredation, and was not designed to decrease wolf depredation at a regional scale or in the long-term. We therefore discovered a reoccurring seasonal-annual pattern for wolf depredation and wolf control in western North America. Ranchers and managers could use our data for focusing investment of resources to prevent wolf depredation increases during high-depredation seasons.

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... Lethal control, as a management tool, can be effective (Conover 2002). However, livestock depredations commonly recur annually after wolves are removed lethally following a depredation (Fritts et al. 1992; Gehring et al. 2003), and does not appear to reduce depredations at a regional scale (Musiani et al. 2005). Nonlethal management tools are regarded by society as more humane than lethal control (Reynolds and Tapper 1996; Reiter et al. 1999). ...
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Human-wildlife conflicts, especially those involving large carnivores, are of global conservation and livelihood concern and require effective and locally-adapted prevention measures. Risk of lion attack on livestock (i.e., depredation) may vary seasonally and may be associated with variation in wild prey abundance or landscape characteristics. To test these competing hypotheses, we used a resource selection approach, and determined whether prey catchability (indicated by geo-spatial variables), or prey availability (indicated by modeled abundance recorded via camera traps) explained spatial and seasonal variation in livestock depredation risk by African lions on Manyara Ranch Conservancy, a multi-use area in northern Tanzania. Seasonal variation in vegetative productivity and proximity to surface water appeared to be strong predictors of livestock depredation risk. Correlates for depredation risk were different between wet and dry seasons. During the dry season, depredation risk was positively correlated with vegetative productivity, whereas depredation risk during the wet season was highest near livestock enclosures (bomas). During both seasons, depredation risk was high closer to surface water. Landscape-driven risk maps were created to identify low risk areas that may be compatible with livestock grazing. Our results on depredation risk by lions are similar to other studies in protected areas and suggest that both prey catchability and prey availability are instrumental in predicting kill sites of lions. To facilitate lion and livestock coexistence in multi-use areas of Africa, we recommend minimizing spatiotemporal overlap between livestock and abundant wild prey by developing alternative livestock water and feeding locations and increasing caution near surface water areas.
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Despite the generally positive trend of European populations, the wolf (Canis lupus) is still today a challenging species to conserve, particularly in the most anthropogenic southern European countries, because of its conflict with humans. In this chapter we summarize the dynamics of wolf distribution in Italy, one of the most densely populated European countries, over the last 50 years. We track changes in the wolf’s diet by comparing its change in Italy with other countries, with the aim of understanding how these changes may have affected the evolution of the human-predator conflict in Italy. In particular, we summarize the results of studies both in Italy and in other European countries to clarify the true impact of wolf predation on both livestock and wild ungulates, which represent the two main causes of predator-human conflict. In order to provide specific insight about the past and the current distribution and feeding habits of the wolf in Italy, and to take stock of the conflict between wolves and humans, we present three case studies. All were carried out over recent decades in northern Italy, i.e. in the area where wolf packs, and particularly their ability to produce dispersing individuals, could affect the future of the entire Italian population. Finally, we consider how to mitigate wolf-human conflict and suggest effective management of wolf populations.
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The cheetah is a prominent example for human–carnivore conflicts and mitigation challenges. Its global population suffered a substantial decline throughout its range. Here, we present an in-depth and new understanding of the socio-spatial organization of the cheetah. We show that cheetahs maintain a network of communication hubs distributed in a regular pattern across the landscape, not contiguous with each other and separated by a surrounding matrix. Cheetahs spend a substantial amount of their time in these hubs, resulting in high local cheetah activity, which represents a high local predation risk for livestock. Implementing this knowledge, farmers were able to reduce livestock losses by 86%.
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Recovery of gray wolves ( Canis lupus ) in the Great Lakes region has been accompanied by an increase in wolf human conflicts. The interface between owners of domestic animals and wolf recovery presents unique challenges for wildlife management. Investigating wolf complaints, explaining wolf ecology, conservation goals, and litigation that has impacted wolf management to people who have had domestic animals killed by wolves are challenges faced by those involved with managing wolf human conflicts. In this chapter, we describe wolf human conflicts and management, focusing on the period 1974 2006, when wolves were protected under the Federal Endangered Species Act (ESA). The patterns of European settlement and wolf persecution were similar in Minnesota, Wisconsin, and Michigan. Minnesota maintained a bounty system for wolves from 1849 to 1965, aerial hunting of wolves persisted until 1956, from 1965 to 1973 wolves could be harvested for fur, and depredation control existed through a state program until May 1974, removing ∼250 wolves per year (Minnesota Department of Natural Resources [MNDNR] 2001 ; United States Fish Wildlife Service [USFWS] 2007). Wisconsin maintained a bounty system for predators, including wolves, from 1839 to 1957. A wolf bounty was the ninth law passed by the first Michigan legislature in 1838. By 1910, wolves were extirpated from Michigan s Lower Peninsula. The bounty continued until 1922. From 1922 to 1935, a state trapper system was in effect. The bounty was reinstated in 1935 and repealed in 1960, after wolves were nearly extirpated from Michigan. In 1915, the United States Congress appropriated funds for a federal wolf control program administered by the United States Department of Agriculture, Bureau of Biological Survey (Young and Goldman 1944). Managers quickly recognized that public acceptance and effective depredation management were necessary for wolf recovery (USFWS 1978a, 1992 ; Peek et al. 1991). During recovery, depredation management was an important component of federal and state wolf management plans (USFWS 1992 ; Michigan Department of Natural Resources [MIDNR] 1997 ; Wisconsin Department of Natural Resources [WDNR] 1999, 2006 ; MNDNR 2001).
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We used 13 years of historical data to investigate effects of coyote (Canis latrans) removal on depredation of domestic sheep. The 2,168-ha study area maintained >1,000 breeding ewes that produced lambs yearly. Records from 1981 through 1994, which included numbers of sheep, numbers of sheep known killed by coyotes, known numbers of coyotes removed, and annual numbers of trapper hours were summarized and analyzed on a yearly, seasonal, and monthly basis. We used regression analysis and found that annual, seasonal, or monthly depredation losses were not correlated with number of coyotes removed. Both annual number of lambs killed and number of coyotes removed were positively correlated with number of trapper hours. We used a cross-correlation analysis to detect any relation between coyote removal and subsequent depredation losses at all monthly intervals from 0 to 24 months. We found a trend of low negative correlation between depredation losses and number of coyotes removed for lags of 2-12 months, suggesting some reduction of sheep killing due to control efforts. Low correlations within years may be due to inconsistent removal of depredating coyotes while removing primarily young, nondepredating coyotes. Lack of correlation between years may have occurred because past control efforts have not had a lasting reduction on coyote density due to immigration, the compensatory nature of control efforts on coyote mortality, reproductive compensation in the resident coyote population, or all 3 factors.
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Financial compensation for damages caused by wildlife is an alternative to lethal wildlife damage management techniques, but little is known about the use of these programs in North America. We conducted surveys requesting information on wildlife species and type of damage covered by compensation programs, annual cost of programs, and the monitoring and assessment of program success to the wildlife agencies of all states and Canadian provinces. We also requested information on programs providing producers with damage-abatement materials instead of or in addition to financial compensation. All states and provinces responded to our survey. Nineteen states and 7 provinces had compensation programs, and 34 states and 7 provinces provided damage-abatement materials. Most programs were funded by the state, but private and federal organizations also funded some programs. Deer (Odocoileus spp.) were the most common species in compensation programs (in 14 states and provinces) followed by bear (Ursus spp.; in 12), elk (Cervus elaphus; in 10), moose (Alces alces; in 7), waterfowl (in 6), pronghorn antelope (Antilocapra americana; in 6), wolves (Canis spp.; in 5), mountain lions (Puma concolor; in 4), and coyotes (Canis latrans; in 3). Compensation programs involving ungulates included damage to cultivated crops (in all 15 states and provinces), standing hay crops and pastures (in 5), stored hay (in 6), and damage to other property including fencing and irrigation equipment (in 8). Programs for predators involved livestock losses. Programs for bears involved damage to crops, livestock, and beekeeping equipment. In general, compensation programs were established for problems that were recent in origin, exacerbated by governmental actions, or caused by highly valued species. Few states or provinces had formal evaluation procedures for their programs. Given the expense of compensation programs and divided opinions about the programs, we recommend that all states and provinces implement a formal review system.
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Predation by coyotes (Canis latrans) on livestock continues to plague producers in the United States. Agricultural interests are concerned about coyote predation because sheep inventories in the U.S. have declined >85% in the past 60 years, with a 25% decline between 1991 and 1996, This decline in sheep numbers has been attributed to low economic returns among producers, with coyote predation cited as a major causative factor. Generalizations about the magnitude and nature of depredations can be misleading because of the varied nature of sheep operations, including size of operations, differences in management, and environmental circumstances surrounding individual operations. Coyote depredation rates appear to be influenced by sheep management practices, coyote biology and behavior, environmental factors, and depredation management programs, Most nonlethal depredation control techniques fall within the operational purview of the producers. The major controversy regarding depredation management focuses on programs that remove coyotes to prevent or curtail predation on domestic stock, especially on public lands. Differences in the magnitude, nature, and history of problems caused by coyotes, as well as the circumstances in which they occur, dictates a need for a variety of techniques and programs to resolve problems. The resolution of coyote depredation upon livestock remains controversial for producers, resource managers, and the general public. Because various segments of society attach different values to coyotes, resolution of depredations should use management programs that integrate the social, legal, economic, and biological aspects of the animals and the problem. Preferred solutions should involve procedures that solve problems as effectively, efficiently, and economically as possible in the least intrusive and most benign ways. Predation management requires a partnership among producers and wildlife managers to tailor programs to specific damage situations so the most appropriate techniques can be selected. This paper attempts to clarify the issues surrounding depredation management, synthesize past and current research, and provide information to resource managers associated with coyote depredation management. This synthesis integrates current understandings of coyote biology and behavior, the nature of depredations upon sheep producing enterprises, and the merits of various depredation control strategies and techniques.
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Animals commonly choose among habitats that differ both in foraging return and mortality hazard. However, no experimental study has attempted to predict the level of increase in resources, or the decrease in mortality hazard, which will induce a forager to shift from a safer to a more hazardous (but richer) foraging area. Here we present and test a model that specifies the choice of foraging areas ("habitats") that would minimize total mortality risk while allowing collection of some arbitrary net energy gain. We tested the model with juvenile creek chubs (Semotilus atromaculatus) in an experimental field stream in which the foragers could utilize a foodless refuge and choose between two foraging areas that differed in experimentally manipulated resource densities (Tubifex spp. worms in sediments) and mortality hazard (adult creek chubs). For the case tested, the model specified a simple rule: "use the refuge plus the site with the lowest ratio of mortality rate (μ) to gross foraging rat (f)," i.e., "minimize μ./f." Independent prior measurements of mortality hazard (as a function of predator density) and gross foraging rate (as a function of resource density) allowed us to predict the resource level in the more hazardous foraging site that should induce a shift from the safer to the more hazardous site. The chubs' preferences in subsequent choice experiments agreed well with the theoretical predictions. The "minimize μ/f" rule (deaths per unit energy), perhaps in modified form, provides a simple alternative to the "maximize f" (energy per unit time) criterion that applies to long-term rate maximization when predation hazard does not differ among choices.
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As wolves (Canis lupus) recolonize mixed forest and agriculture areas in the Lake Superior region of the United States, their depredations on livestock are increasing, along with public complaints and compensation payments. We documented 176 complaints about wolves in Wisconsin between 1976 and 2000 and analyzed the regional and temporal patterns for the 87 verified incidents involving the injury or death of 377 domestic animals. Calves were the most frequent target of wolf depredation, but game-farm deer losses demanded higher compensation payments. Sixty-six property owners were affected by wolf depredations over the 25-year period examined. Compensation costs averaged $96.00 per capita of wolf/year. Two thirds of 71 breeding wolf packs were never suspected of causing depredations, but 4 packs were involved in ≥4 incidents. These data were collated to aid in preventing wolf depredation and provide a foundation for policy-making surrounding the impending federal delisting of the wolf.
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The use of aversive conditioning, repellents and deterrents in the management of predator-livestock problems is evaluated based on a comprehensive literature review, contact with leading authorities and visits to areas with similar predation problems. The status of these management tools is reported and their applicability under Scandinavian conditions evaluated. Aversive conditioning usually involves treating baits with an emetic compound (usually lithium chloride), and has shown inconsistent and inconclusive results. Repellents and deterrents include physical, chemical and acoustic stimuli or devices that cause predators to stop an unwanted behaviour or to retreat from an area. Chemical repellents are not particularly effective against coyotes but have been effective for wolverines and bears under some conditions (e.g. with the availability of untreated, alternative prey). Projectile repellents give an immediate, positive result with bears, but their use is limited. Visual and acoustic devices work well, but only for a limited time, as predators quickly habituate to these devices. To summarize, these methods generally show little promise in reducing livestock depredation on a large-scale or long-term basis, especially under the conditions prevailing in Scandinavia.
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The use of domestic animals to protect livestock was reviewed through visits to actual users, discussions with experts and a thorough literature search. Costs and benefits were analysed in terms of reduced livestock losses. The most common guardian animals are dogs, which have been shown to reduce predation (documented mostly for coyote) by 11?100%. Livestock guardian dogs have also been used effectively against bear, wolf and cheetah. Donkeys are also used as guardian animals, and their effectiveness lies in their natural herding behaviour and aggression, especially against canids. The effectiveness of donkeys varies considerably dependent upon the predator species and the temperament of the individual donkey. Llamas are also used as a guardian animal, with approximately the same characteristics as the donkeys, and will defend themselves against most predators. The use of guardian animals appears to be an effective tool for reducing livestock depredation and should be evaluated in areas with high predation losses against the cost of changing production systems.
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To balance conXicting demands for food and safety from predation, feeding animals have two useful tools. First, they can vary the amount of time they devote to harvesting patches that vary in predation risk and feeding rates. Second, they can use vigilance to trade-oV food and safety while feeding from a food patch. I present a model for predicting how an optimal forager should jointly use these two tools. Factors inXuencing the use of these tools include encounter rate with predators, predator lethality in the absence of vigilance, eVectiveness of vigilance in reducing predator lethality, the marginal value of energy to the forager and the forager's survivor's Wtness. Patch-use behaviours inXuenced by these factors include vigilance level, quitting harvest rate and giving-up density (GUD). All three of these patch-use behaviours should increase in response to an increase in encounter rate with predators, predator lethality and the forager's survivor's Wtness, and decrease with an increase in the marginal value of energy. In response to increasing the eVectiveness of vigilance, vigilance should increase and the GUD and quitting harvest rate should decline. The amount of food left by a forager in a depletable food patch, the GUD, provides an empirical link for testing the model's predictions. Giving-up densities should increase with increasing predation risk, and GUDs should increase with declining food-density- speciWc harvest rates. DiVerences in GUDs among food patches attributable to diVerences in quitting harvest rates measure the contribution of time allocation to managing diVerences in predation risk. DiVerences in GUDs attributable to diVerences in food-density-speciWc harvest rates measure the contribution of vigilance to managing predation risk.
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We studied the kill rate by wolves (Canis lupus) after a large-scale wolf removal when populations of wolves, moose (Alces alces), and woodland caribou (Rangifer tarandus caribou) were all increasing. We followed a total of 21 wolf packs for 4 winters, measuring prey selection, kill rates, and ecological factors that could influence killing behavior. Wolf predation was found to be mainly additive on both moose and caribou populations. Kill rates by individual wolves were inversely related to pack size and unrelated to prey density or snow depth. Scavenging by ravens decreased the amount of prey biomass available for wolves to consume, especially for wolves in smaller packs. The kill rate by wolves on moose calves was not related to the number of calves available each winter. Wolves did not show a strong switching response away from moose as the ratio of caribou to moose increased in winter. The predation rate by wolves on moose was best modeled by the number and size of packs wolves were organized into each winter.
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In Alberta, Canada (1982-2001), and in Idaho, Montana, and Wyoming, United States (1987-2001), wolves (Canis lupus) killed various domestic animals, among which the major prey were sheep in the United States (68%, n = 494) and cattle in Canada (95%; n = 1633). Under recovery programs, the wolf population increased in the United States, and depredation events increased proportionately. In both countries, the number of domestic animals killed each year was correlated with the number of wolves killed by government authorities for depredation management. We tested the ability of anti-wolf barriers made of flags hanging from ropes to impede wolf access to food and livestock. In 18 experiments, barriers prevented captive wolves (n = 9) from accessing food for up to 28 hours and allowed daily separation of wolves to administer contraceptive pills to a female wolf. Barriers prevented access by wild wolves to 100-m2 baited sites during two 60-day tests. We also set barriers around three cattle pastures. In Alberta during two 60-day trials on 25-ha pastures, wolves approached barriers on 23 occasions but did not cross them, and no cattle were killed. Wolves killed cattle on neighboring ranches during the trials and before and after the trials on the tested ranches. In Idaho four radio-collared wolves crossed barriers and killed cattle in a 400-ha ranch after 61 days of barrier exposure. Our results suggest that anti-wolf barriers are effective in deterring captive and wild wolves for >1 and ≥60 days, respectively, and that wild wolves switch to alternative livestock when excluded from one herd of livestock. Our depredation data indicate that protecting livestock from wolves reduces the necessity for killing wolves. Barriers could play a role among the limited set of preventive measures available and offer a cost-effective mitigation tool for the problem of livestock depredation on a local scale.
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The selective removal of carnivores from local areas is sometimes proposed to reduce the number of attacks on livestock. For the lynx, neither the existence of problem individuals nor the efficacy of their selective removal has been demonstrated. In France, from 1989 to 1999, eight lynx and two large carnivores thought to be lynx were legally removed from high conflict areas by trapping (n=7), shooting (n=1) or poisoning with toxic collars on sheep (n=2). The efficacy of the 10 removals was assessed on the farms where a lynx was caught and in the 5-km-radius areas encompassing both these farms and nearby sheep farms. The sex-ratio of captured lynx was seven males:one female. On four farms and in six 5-km-radius areas lynx attacks on sheep reappeared within 40 days after lynx removal, but we observed a significant decrease in the overall number of attacks. In the medium-term (48–365 days), the number of attacks decreased on two farms and in four 5-km radius areas when compared with the number observed in control plots >10 km away from the removal sites. In the long-term, attacks reappeared on the same sites, indicating a “site” effect. In such situations, selective removals may only temporarly reduce the problem of concentrated lynx damage. The only way to obtain a durable effect is to improve shepherding techniques.
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Mountain lion (Puma concolor) depredation incidents on livestock herds were recorded at 15 ranches in southern Brazil from 1993 to 1995. Maximum losses to mountain lions were 78% for goats, 84% for sheep, and 16% for cattle. Cattle mortality arising from causes other than depredation assumed a greater importance in herd productivity. In contrast, attacks on sheep and goats were more frequent than losses to other causes, but could be reduced to acceptable levels when flocks were corralled at night. Most depredation incidents occurred when weather and light conditions were unfavorable to human activity. We explain these patterns and inter-ranch variation in depredation rates on the basis of a risk-avoidance strategy by the mountain lions. Stock losses can be minimized by understanding these patterns and by applying appropriate herd husbandry, thus reducing the urge to persecute this protected species.
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Australian examples of surplus killing by mammalian predators were collated. These included surplus killing of native mammals and birds by foxes (Vulpes vulpes) and stock, native mammals and native birds by dingoes (Canis lupus dingo). We found no examples of surplus killing by feral cats (Felis catus). Incidents collated include historical anecdotes of surplus killing by foxes as they colonised the Australian mainland, recent examples where foxes killed threatened native species at sites despite intensive management to exclude foxes, and recent examples of the killing of native species on formerly fox-free islands to which foxes gained entry. Episodes of surplus killings by foxes, other than predation on captive or closely confined animals, appeared different in kind and frequency to those documented for co-evolved predator–prey systems on the large continental landmasses. They did not appear to be uncommon events associated with synchronised births of prey species, unusual or extreme weather that disadvantaged prey species, or seasonal food caching by a predator. Rather, surplus killing events appeared to reflect ineffective anti-predator defences by prey species when encountering a novel and efficient predator to which they have had no evolutionary exposure. We suggest that surplus killing by foxes may have been a feature of, and major contributor to, the rapid mainland extinction or contraction in range of many native species in Australia. In contrast to foxes, examples of surplus killing by dingoes relate mostly to domestic stock (calves and sheep). The arrival of dingoes to the Australian continent preceded that of foxes by 3500–4000 years, but they appear not to have had the dire impact on native mammals that we attribute to foxes. This may be due to fundamental differences in hunting styles and prey size and to their sparse populations in pre-European Australia. Active persecution of non-commensal dingoes by Aborigines, the lack of free-water, and the absence of European rabbits (Oryctolagus cuniculus) as an alternative food supply would have limited their numbers and their impact on native mammals.
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Phenotype-based selective harvests, including trophy hunting, can have important implications for sustainable wildlife management if they target heritable traits. Here we show that in an evolutionary response to sport hunting of bighorn trophy rams (Ovis canadensis) body weight and horn size have declined significantly over time. We used quantitative genetic analyses, based on a partly genetically reconstructed pedigree from a 30-year study of a wild population in which trophy hunting targeted rams with rapidly growing horns, to explore the evolutionary response to hunter selection on ram weight and horn size. Both traits were highly heritable, and trophy-harvested rams were of significantly higher genetic 'breeding value' for weight and horn size than rams that were not harvested. Rams of high breeding value were also shot at an early age, and thus did not achieve high reproductive success. Declines in mean breeding values for weight and horn size therefore occurred in response to unrestricted trophy hunting, resulting in the production of smaller-horned, lighter rams, and fewer trophies.
Article
Wolves (Canis lupus) were once common throughout North America but were deliberately exterminated in the lower 48 United States, except in northeastern Minnesota, primarily because of depredations on livestock. Wolves remained abundant in areas with few livestock such as most of Canada and Alaska. Sixty years after being nearly exterminated, the gray wolf was listed under the United States Endangered Species Act (Act) in 1974. The combination of natural recovery in NW Montana, and reintroduction in central Idaho and the Greater Yellowstone area (NW Wyoming, eastern Idaho, and SW Montana) has resulted in an expanding wolf population (Bangs et al. 1998). In this paper we discuss our attempts to minimize conflicts between wolves and livestock and to build human tolerance for restoring wolf populations.
Article
The level of Canis lupus depredation on livestock in Minnesota during 1975-86 showed a slight upward trend but did not increase significantly. Conflicts were highly seasonal and involved primarily cattle (mainly calves), sheep, and domestic turkeys. Site-specific trapping and removal of wolves in response to depredations was the primary control method. -from Authors
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Data support the hypothesis that Canis lupus depredation on domestic animals is inversely related to winter severity. Wolves take domestic animals inversely to the availability of white-tailed deer Odocoileus virginianus fawns. -from Authors
Article
We investigated wolf (Canis lupus)- and dog-livestock conflicts (1992-1995) and costs of compensation (1991-1995) in the Tuscany region of central Italy. The regional indemnity program cost US $345,000 (± 93,000 SD) annually. Most depredations (95.2%) involved sheep, with a mean (± SD) annual loss of 2,550 ± 730 sheep, or 0.35% of the regional stock. Sheep lost to predators by province were correlated with sheep density within areas containing wolves (r(s)= 0.88, n = 9, P = 0.0015), but marked geographical and temporal fluctuations were reported in compensation costs. Highest levels of conflict were observed in the provinces at the border of the regional wolf range, where livestock was left unattended most of the year and sheep density reached its highest regional levels. Based on 527 reports of approved claims during 1992-1995 from the National Health System, depredations were highly seasonal, increasing steadily from spring to early fall, possibly following trends in sheep availability on pastures and density fluctuations of local wolf packs. An average of 3 sheep (range = 1-18) were killed per attack (n = 483), and 42% of the attacks involved killing of ≤2 sheep. Additionally, 21-113 sheep were killed or attacked in mass slaughters which comprised 2.3% of the depredation events and 19% of the sheep lost. Depredations also resulted in 35% (n = 168) of sheep injured and 33% (n = 158) missing. Most sheep depredations occurred during the night, in pastures interspersed with wood or vegetative cover, and involved free-ranging flocks unattended by either the shepherd or guard dogs. High levels of conflict occurred in localized areas of intensive sheep production; 6% of the affected farms and 8% of the affected municipalities accounted for 32% of the sheep lost to both wolves and dogs at the regional level. Compensation programs alone were not effective in reducing the conflict or in preventing illegal, private efforts to control wolf numbers. Improved husbandry should be encouraged and facilitated through financial incentives and public education.
Article
Grazing systems for the Northern Great Plains based on the exclusive use of natural grassland are no better than continuous grazing. Since the quality of the majority of the ecosystems within the region makes seeded grass pastures feasible, seeded pastures containing highly adapted cultivars such as crested wheatgrass, Russian wildrye and alfalfa can be used in various grazing systems to balance and extend the grazing season. The growth habits and nutrient characteristics of the herbage of the native grasses are of maximum value for a relatively short period during the year. Grazing systems detailed include one in which the requirement per animal unit is reduced from 24.8 to 11.4 acres.
Article
Coyotes (Canis htrans), black bears (Ursus amerkanus), and wolves (Canis lupus) were reported responsible for 35, 31, and 16%, respectively, of confirmed predation losses of cattle in Alberta during 1974-78. Coyotes selected for calves over adults, and adults over yearlings, black bears selected for calves over yearlings, and yearlings over adults, and wolves selected for calves and yearlings over adults. Predation of cattle by coyotes, bears, and wolves peaked during March-June, May-July, and August-September, respectively. Little information is available on predation losses ofcattle, even though the value of losses from predation are comparable for cattle and sheep in the United States (Anon. 1978, Gee 1979). In Alberta, the value of cattle lost to predation from coyotes, black bears, and wolves exceeds that of other species of livestock.1 This paper describes the monthly chronology and age distribution of preda-tion losses of cattle in Alberta, Canada during 1974-78.
Article
Wolf (Canis lupus) depredations on livestock cause considerable conflict and expense in Minnesota. Furthermore, claims are made that such depredations are fostered by the type of animal husbandary practiced. Thus, we tried to detect factors that might predispose farms in Minnesota to wolf depredations. We compared results of interviews with 41 cattle farmers experiencing chronic cattle losses to wolves (chronic farms) with results from 41 nearby "matched" farms with no wolf losses to determine farm characteristics or husbandry practices that differed and that therefore might have affected wolf depredations. We also used a Geographic Information System (GIS) to detect any habitat differences between the 2 types of farms. We found no differences between chronic and matched farms in the 11 farm characteristics and management practices that we surveyed, except that farms with chronic losses were larger, had more cattle, and had herds farther from human dwellings. Habitat types were the same around farms with and without losses. The role of proper carcass disposal as a possible factor predisposing farms to wolf depredations remains unclear.
Article
After gray wolves (Canis lupus) were extirpated over a large portion of their North American range during the early 1900s, researchers reviewed the history of wolf-human encounters and concluded that wild, free-ranging wolves posed little or no threat to human safety. However, documented cases of wolf aggression toward people have recently increased, indicating a need for further examination of wolf-human interactions. I reviewed 80 cases of wolf-human encounters and compared behaviors of wild wolves that interacted with people in different contexts in Alaska and Canada. Only 1 case of unprovoked wolf aggression was documented between 1900 and 1969, but 18 cases of unprovoked wolf aggression toward people occurred between 1969 and 2000, including 3 cases of serious injury to children since 1996. Increases in wolf protection, human activities in wolf habitat, and wolf numbers occurred concurrently with increases in unprovoked aggressive encounters. Aggressive behavior was documented in all regions and among all wolf subspecies of Alaska and Canada. Wolves rarely vocalized during unprovoked aggressive encounters, but wolves that were defending dens consistently displayed loud vocalizations. Behavior of rabid wolves was variable and ranged from stubborn, persistent approaches to prolonged attacks. Habituation contributed to unprovoked wolf aggression toward people in 11 cases; nonhabituated wolves in remote areas displayed unprovoked aggression in 7 cases. Where wolves are protected and frequently encounter people, some level of negative conditioning should be applied to prevent habituated and food-conditioned behaviors in wolves.
Article
Three of the most commonly used mathematical relationships between zooplankton ingestion and food concentration are derived from a queue model, by assuming different feeding strategies of the individual predators or grazers. The model suggests that the abruptness of the transition between maximum search or filtration rate and maximum ingestion rate depends on the size ratio between the animals and their food items. The results show that the stochastic nature of predation and grazing cannot be neglected when the food particles are relatively big.
Article
Pastoralists and their livestock share much of the habitat of the snow leopard (Uncia uncia) across south and central Asia. The levels of livestock predation by the snow leopard and other carnivores are high, and retaliatory killing by the herders is a direct threat to carnivore populations. Depletion of wild prey by poaching and competition from livestock also poses an indirect threat to the region's carnivores. Conservationists working in these underdeveloped areas that face serious economic damage from livestock losses have turned to incentive programs to motivate local communities to protect carnivores. We describe a pilot incentive program in India that aims to offset losses due to livestock predation and to enhance wild prey density by creating livestock-free areas on common land. We also describe how income generation from handicrafts in Mongolia is helping curtail poaching and retaliatory killing of snow leopards. However, initiatives to offset the costs of living with carnivores and to make conservation beneficial to affected people have thus far been small, isolated, and heavily subsidized. Making these initiatives more comprehensive, expanding their coverage, and internalizing their costs are future challenges for the conservation of large carnivores such as the snow leopard.
Article
Results of wolf (Canis lupus) control to reduce predation of cattle in northwestern Alberta are reported. Numbers of wolves declined from about 40 prior to control to 3 following the strychnine poisoning of 26 wolves during 2 winters, 1979-80 and 1980-81. Additional losses of wolf pack members occurred from natural mortality and dispersal following the removal of the majority of their packmates. In 3 of 6 instances where packs took baits, entire packs of 2, 4, and 6 wolves were killed. Ingress of wolves occurred within 1-2 years. Total mortality of cattle declined from a mean of 64 (3.4%) during 4 years prior to control to 36 (2.0%) during 2 years following control. Selectivity of strychnine poisoning was reasonably good although more emphasis on preventive management is recommended.
Article
Abstract We examined interactions between wolves (Canis lupus) and domestic calves within a grazing allotment in central Idaho to evaluate the role of wolf predation in calf survival and movements. During the 1999 and 2000 grazing seasons, we radio-marked 231 calves per year, representing 33% of the calf population, on the Diamond Moose Association (DMA) grazing allotment, and monitored their survival and movements relative to wolf distribution. Overall, calf survival was high ( 95%), with relatively few mortalities (n=13) among the marked population. Non-predation calf mortality (pneumonia, unknown natural causes, and fire) and wolf-caused calf mortality represented 61% and 31% of deaths, respectively, while coyote predation accounted for the remaining (7%) mortality. Calves selected by wolves were younger than the surviving cohort by an average of 26 days (P< 0.05). Calf movement patterns and group size did not vary relative to the level of spatial overlap with wolves, however, vulnerability to predation appeared to be correlated with spatial proximity of calves to wolf home,ranges and rendezvous sites. These results suggest that in our study area the overall impact of wolves on calf survival and behavior was modest, and that ranchers could further minimize wolf predation by altering calving periods to favor older calves and minimize spatial overlap between grazing cattle and areas of intense wolf activity. 3
Article
Wolf management can be controversial, reflecting a wide range of public attitudes. We analyzed wolf management case histories representing a spectrum of approaches in Canada and the United States. During the early 20th century, wolves were considered undesirable. They were subject to persecution and were extirpated from large areas of their original range. With increased environmental awareness in the 1970s, attitudes toward wolves began to change. Wolf conservation became a focus of public interest, providing conditions that favored regional wolf recovery. However, in regions where livestock production or big-game hunting is valued, wolves have continued to be controlled by management authorities or through the actions of individual citizens. With US wolf populations recovering in the conterminous states, a rule was approved to delist the species from endangered to threatened status under the Endangered Species Act. Notwithstanding the intent of legal instruments, history has demonstrated that societal values ultimately determine the survival of species such as the wolf.
Article
Carnivore conservation depends on the sociopolitical landscape as much as the biological landscape. Changing political attitudes and views of nature have shifted the goals of carnivore management from those based on fear and narrow economic interests to those based on a better understanding of ecosystem function and adaptive management. In parallel, aesthetic and scientific arguments against lethal control techniques are encouraging the development of nonlethal approaches to carnivore management. We anticipate greater success in modifying the manner and frequency with which the activities of humans and domestic animals intersect with those of carnivores. Success should permit carnivore populations to persist for decades despite human population growth and modification of habitat.Resumen: La conservación de carnívoros depende tanto del paisaje sociopolítico como del paisaje biológico. Cambios en las actitudes políticas y percepciones de la naturaleza han cambiado las metas de manejo de carnívoros de aquéllas basadas en el miedo y las intereses económicos estrechos a metas basadas en un mejor entendimiento del funcionamiento del ecosistema y en el manejo adaptativo. A su vez, los argumentos estéticos y científicos en contra de las técnicas de control letal están fomentando el desarrollo de planteamientos no letales en la gestión de carnívoros. Anticipamos un mayor éxito en la modificación del modo y la frecuencia en que las actividades de humanos y animales domésticos intersectan con las de carnívoros. El éxito debe permitir que las poblaciones de carnívoros persistan por décadas a pesar del crecimiento de la población humana y la modificación de hábitats.
Article
During this century, Canis lupus in Alberta recovered from low numbers during the 1900s to 1920s (few ungulates, bounty, and poisonings) and the 1950s (rabies control). Wolves increased in abundance during the late 1960s and the 1970s when protection was the primary wolf management goal. An average midwinter estimate of 4200 wolves during 1975-1985 was calculated. Wolf management strategies designed in 1983 to enhance ungulate populations and reduce livestock depredations included encouraging wolf trapping and hunting and reducing regional wolf populations. Management of wolf predation on ungulates was ineffective. Although predation seems to be an important factor contributing to low ungulate populations, wolf control was not conducted because of apprehension caused by public controversy. Experimental manipulation of predator and ungulate dynamics is recommended. -from Author
Article
Summary 1. Human-elephant conflict (HEC) in Africa occurs wherever these two species co- incide, and poses serious challenges to wildlife managers, local communities and elephants alike. Mitigation requires a detailed understanding of underlying patterns and processes. Although temporal patterns of HEC are relatively predictable, spatial variation has shown few universal trends, making it difficult to predict where conflict will take place. While this may be due to unpredictability in male elephant foraging beha- viour (the male behaviour hypothesis) it may also be due to variations in the data resolution of earlier studies. 2. This study tested the male behaviour and data resolution hypotheses using HEC data from a 1000-km 2 unprotected elephant range adjacent to the Masai Mara National Reserve in Kenya. HEC incidents were divided into crop raiding and human deaths or injuries. Crop raiding was further subdivided into incidents involving only male ele- phants or family groups. A relatively fine-resolution, systematic, grid-based method was used to assign the locations of conflict incidents, and spatial relations with under- lying variables were explored using correlation analysis and logistic regression. 3. Crop raiding was clustered into distinct conflict zones. Both occurrence and intensity could be predicted on the basis of the area under cultivation and, for male elephant groups, proximity to major settlements. Conversely, incidents of elephant-induced human injury and death were less predictable but were correlated with proximity to roads. 4. A grid-based geographical information system (GIS) with a 25-km 2 resolution utilizing cost-effective data sources, combined with simple statistical tools, was capable of identifying spatial predictors of HEC. At finer resolutions spatial autocorrelation compromised the analyses. 5. Synthesis and applications . These results suggest that spatial correlates of HEC can be identified, regardless of the sex of the elephants involved. Moreover, the method described here is fully transferable to other sites for comparative analysis of HEC. Using these results to map vulnerability will enable the development and deployment of appropriate conflict mitigation strategies, such as guarding, early warning systems, barriers and deterrents. The utility of such methods and their strategic deployment should be assessed alongside alternative land-use and livelihood strategies that limit cultivation within the elephant range.
Article
Summary 1. Most studies of animal movements and habitat selection do not recognize empiric- ally that different components of the environment are important to animals at different scales. Often, availability of habitats is defined at one or more arbitrary spatio-temporal scales, but use of those habitats is constrained to one scale. Identification of scalar movement is the first step in developing models to explain why animals select or move to certain parts of their range. We used a non-linear curve-fitting model of movement rates to identify discontinuities in the scales of movement by woodland caribou Rangifer tarandus caribou collared with global positioning system (GPS) collars. 2. We differentiated intrapatch from interpatch movements, but were unable to distinguish interpatch from migratory-type movements for most combinations of indi- vidual caribou by season. Model fit was stronger for winter than summer movements. We suggest that increased patch heterogeneity during the winter resulted in interseason variation in movements and corresponding model fit. 3. Responses by caribou to the environment were scale-dependent. When we applied logistic regressions, land-cover type, energetic costs of movement, and predation risk differentiated the two scales of movement. Intrapatch movements had a lower cost of movement, were associated with cover types where foraging behaviours probably occurred, and were closer to areas of higher predator risk than interpatch movements. 4. Application of the non-linear model will aid in developing mechanism-based approaches to studying resource selection and animal behaviour.
Article
The gray wolf once inhabited a wide variety of habitats throughout most of the northern hemisphere north of 20-degrees-N latitude. Because the animal preyed on livestock and competed with humans for wild prey, it was extirpated from much of its range outside of wilderness areas. Environmental awareness in the late 1960s brought for the wolf legal protection, increased research, and favorable media coverage. The species has increased in both Europe and North America, is beginning to reoccupy semiwilderness and agricultural land, and is causing increased damage to livestock. Because of the wolf's high reproductive rate and long dispersal tendencies, the animal can recolonize many more areas. In most such areas control will be necessary, but the same public sentiments that promoted wolf recovery reject control. If wolf advocates could accept control by the public rather than by the government, wolves could live in far more places. Insistence on government control discourages some officials and government agencies from promoting recovery. The use of large- or small-scale zoning for wolf management may help resolve the issue. Public education is probably the most effective way to minimize the problem and maximize wolf recovery, but the effort must begin immediately.
Article
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.
Article
Gray wolves (Canis lupus) in parts of the United States and Europe live in networks of disjunct populations, many of which are close to human settlement. Because wolf management goals include sustaining disjunct populations, it is important to ask what types of areas and protections are needed for population survival. To predict the effects of different levels of human-caused mortality, we created a simulation model for a disjunct wolf population living in a semi-wild landscape with abundant, well distributed prey. The landscape included a maximum of 16 territories divided into core and peripheral range. The mortality rate in the core range was 20%, whereas the mortality rate in peripheral range (40%) was higher because of human-caused deaths. We examined the relationship between the proportions of core and peripheral range and the 50-year occupancy of that range by wolf packs, given different assumptions about pup and dispersal mortality and immigration. Simulations showed that occupancy increased as the number of core sites increased, but curve location depended on parameter assumptions. With pup and dispersal mortality rates consistent with those for disease-free and legally protected populations, wolves saturated a 16-territory cluster with as few as two core sites, regardless of immigration rate. When populations had high pup or dispersal mortality, as few as two immigrants per year helped maintain high (>80%) site occupancy in clusters with four or more core sites. Small numbers of immigrants were also important for sustaining colonizing populations and buffering the negative effects of increased environmental variation. The simulations supported the claim that wolves can survive in disjunct populations provided that wolves can move between populations, human persecution is not excessive, and prey is abundant.
Article
Conservation biology requires the development of practical tools and techniques to minimize conflicts arising from human modification of ecosystems. We applied behavioral theory of primary and secondary repellents to predator management by using aversive stimulus devices (electronic training collars) and disruptive stimulus devices (behavior-contingent audio and visual repellents) in a multipredator (Canis lupus, Haliaeetus leucocephalus, Ursus spp. ) study in the United States. We examined fladry and a newly developed disruptive stimulus device contingent upon behavior on six wolf territories in Wisconsin, (US.A.) and determined that the disruptive stimulus device gave the greatest degree of protection from predation. We also compared the efficacy of a primary repellent (disruptive stimulus device) versus a secondary repellent (electronic training collars) to keep captive wolves from consuming a food source. Disruptive stimulus devices effectively prevented captive wolves from consuming the food resource, but did not produce an aversion to that food resource. With training collars, logistical and behavioral variability limited our ability to condition wolves. Our studies highlight the complexity of application of nonlethal techniques in real-world situations.
Article
Most large carnivore species are in global decline. Conflict with local people, particularly over depredation on livestock, is a major cause of this decline, affecting both nominally protected populations and those outside protected areas. For this reason, techniques that can resolve conflicts between large carnivores and livestock farmers may make important contributions to conservation. We monitored rates of livestock depredation by lions ( Panthera leo), leopards (Panthera pardus), cheetahs (Acinonyx jubatus), and spotted hyenas (Crocuta crocuta), and retributive killing of these species by farmers in livestock-producing areas of Laikipia District, Kenya. Farmers killed more lions, leopards, and spotted hyenas where these predators killed more livestock. Livestock husbandry had a clear effect on rates of depredation and hence on the numbers of predators killed. Cattle, sheep, and goats experienced the lowest predation rates when attentively herded by day and enclosed in traditional corrals (bomas) by night. Construction of the boma, the presence of watchdogs, and high levels of human activity around the boma were all associated with lower losses to predators. Although most of this work was carried out on commercial ranches, local Maasai and Samburu pastoralists have practiced nearly identical forms of husbandry for generations. Our study shows that traditional, low-tech husbandry approaches can make an important contribution to the conservation of large carnivores.
Article
As wolf (Canis lupus) populations recover in Wisconsin (U. S. A.), their depredations on livestock, pets, and hunting dogs have increased. We used a mail-back survey to assess the tolerance of 535 rural citizens of wolves and their preferences regarding the management of "problem" wolves. Specifically, we tested whether people who had lost domestic animals to wolves or other predators were less tolerant of wolves than neighboring residents who had not and whether compensation payments improved tolerance of wolves. We assessed tolerance via proxy measures related to an individual's preferred wolf population size for Wisconsin and the likelihood she or he would shoot a wolf. We also measured individuals' approval of lethal control and other wolf-management tactics under five conflict scenarios. Multivariate analysis revealed that the strongest predictor of tolerance was social group. Bear (Ursus americanus) hunters were concerned about losing valuable hounds to wolves and were more likely to approve of lethal control and reducing the wolf population than were livestock producers, who were more concerned than general residents. To a lesser degree, education level, experience of loss, and gender were also significant. Livestock producers and bear hunters who had been compensated for their losses to wolves were not more tolerant than their counterparts who alleged a loss but received no compensation. Yet all respondents approved of compensation payments as a management strategy. Our results indicate that deep-rooted social identity and occupation are more powerful predictors of tolerance of wolves than individual encounters with these large carnivores.
Article
Wolves are some of the world's most charismatic and controversial animals, capturing the imaginations of their friends and foes alike. Highly intelligent and adaptable, they hunt and play together in close-knit packs, sometimes roaming over hundreds of square miles in search of food. Once teetering on the brink of extinction across much of the United States and Europe, wolves have made a tremendous comeback in recent years, thanks to legal protection, changing human attitudes, and efforts to reintroduce them to suitable habitats in North America. As wolf populations have rebounded, scientific studies of them have also flourished. But there hasn't been a systematic, comprehensive overview of wolf biology since 1970. In Wolves, many of the world's leading wolf experts provide state-of-the-art coverage of just about everything you could want to know about these fascinating creatures. Individual chapters cover wolf social ecology, behavior, communication, feeding habits and hunting techniques, population dynamics, physiology and pathology, molecular genetics, evolution and taxonomy, interactions with nonhuman animals such as bears and coyotes, reintroduction, interactions with humans, and conservation and recovery efforts. The book discusses both gray and red wolves in detail and includes information about wolves around the world, from the United States and Canada to Italy, Romania, Saudi Arabia, Israel, India, and Mongolia. Wolves is also extensively illustrated with black and white photos, line drawings, maps, and fifty color plates. Unrivalled in scope and comprehensiveness, Wolves will become the definitive resource on these extraordinary animals for scientists and amateurs alike. “An excellent compilation of current knowledge, with contributions from all the main players in wolf research. . . . It is designed for a wide readership, and certainly the language and style will appeal to both scientists and lucophiles alike. . . . This is an excellent summary of current knowledge and will remain the standard reference work for a long time to come.”—Stephen Harris, New Scientist “This is the place to find almost any fact you want about wolves.”—Stephen Mills, BBC Wildlife Magazine
Article
Biologists, breeders and trainers, and champion sled dog racers, Raymond and Lorna Coppinger have more than four decades of experience with literally thousands of dogs. Offering a scientifically informed perspective on canines and their relations with humans, the Coppingers take a close look at eight different types of dogs—household, village, livestock guarding, herding, sled-pulling, pointing, retrieving, and hound. They argue that dogs did not evolve directly from wolves, nor were they trained by early humans; instead they domesticated themselves to exploit a new ecological niche: Mesolithic village dumps. Tracing the evolution of today's breeds from these village dogs, the Coppingers show how characteristic shapes and behaviors—from pointing and baying to the sleek shapes of running dogs—arise from both genetic heritage and the environments in which pups are raised. For both dogs and humans to get the most out of each other, we need to understand and adapt to the biological needs and dispositions of our canine companions, just as they have to ours.
Managing wolf-human conflict in the northwestern United States. Pages. 340-356 People and wildlife: coexistence or conflict?
  • E E Bangs
  • J A Fontaine
  • M D Jimenez
  • T J Meier
  • E H Bradley
  • C C Niemeyer
  • D W Smith
  • C M Mack
  • V Asher
  • J K Oakleaf
Bangs, E. E., J. A. Fontaine, M. D. Jimenez, T. J. Meier, E. H. Bradley, C. C. Niemeyer, D. W. Smith, C. M. Mack, V. Asher, and J. K. Oakleaf. 2004. Managing wolf-human conflict in the northwestern United States. Pages. 340-356. in R. Woodroffe, S. Thirgood, and A. Rabinowitz, editors. People and wildlife: coexistence or conflict? Cambridge University Press, Cambridge, United Kingdom: in press.