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41 LARGE CARNIVORES, MOOSE, AND HUMANS: A CHANGING PARADIGM OF PREDATOR MANAGEMENT IN THE 21 st CENTURY Charles C. Schwartz 1 , Jon E. Swenson 2 , and Sterling D. Miller 3 1 Interagency Grizzly Bear Study Team, U.S. Geological Survey, Biological Resources Division, Northern Rocky Mountain Science Center, Montana State University, Bozeman, MT 59717, USA; 2 Department of Biology and Nature Conservation, Agricultural University of Norway, Box 5014, N- 1432 Ås, Norway; 3 National Wildlife Federation, 240 North Higgins, Suite #2, Missoula, MT 59802, USA ABSTRACT: We compare and contrast the evolution of human attitudes toward large carnivores between Europe and North America. In general, persecution of large carnivores began much earlier in Europe than North America. Likewise, conservation programs directed at restoration and recovery appeared in European history well before they did in North America. Together, the pattern suggests there has been an evolution in how humans perceive large predators. Our early ancestors were physically vulnerable to large carnivores and developed corresponding attitudes of respect, avoidance, and acceptance. As civilization evolved and man developed weapons, the balance shifted. Early civilizations, in particular those with pastoral ways, attempted to eliminate large carnivores as threats to life and property. Brown bears ( Ursus arctos) and wolves ( Canis lupus) were consequently extirpated from much of their range in Europe and in North America south of Canada. Efforts to protect brown bears began in the late 1880s in some European countries and population reintroductions and augmentations are ongoing. They are less controversial than in North America. On the other hand, there are no wolf introductions, as has occurred in North America, and Europeans have a more negative attitude towards wolves. Control of predators to enhance ungulate harvest varies. In Western Europe, landowners own the hunting rights to ungulates. In the formerly communistic Eastern European countries and North America, hunting rights are held in common, although this is changing in some Eastern European countries. Wolf control to increase harvests of moose ( Alces alces) occurs in parts of North America and Russia; bear control for similar reasons only occurs in parts of North America. Surprisingly, bears and wolves are not controlled to increase ungulates where private landowners have the hunting rights in Europe, although wolves were originally exterminated from these areas. Both the inability of scientific research to adequately predict the effect of predator control on ungulate populations and a shift in public attitudes toward large carnivores have resulted in an accelerating number of challenges to predator management in places where it is still espoused. Utilitarian attitudes towards wildlife are declining in Western cultures and people now increasingly recognize the intrinsic value of wildlife, including large predators. In the future, agencies responsible for managing resident wildlife will face increased pressure to balance the needs of the hunting public with the desires of non-hunting publics. We suggest that in the next century we will witness a continued shift in how wildlife agencies manage both moose and large carnivores. More attention will be paid to maintaining and restoring intact ecosystems and less toward sustainable yield of meat. ALCES VOL. 39: 41-63 (2003) Key words: Alces alces, brown bear, Canis lupus, gray wolf, grizzly bear, moose management, predator control, Ursus arctos

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  • ALCES VOL. 39, 2003 SCHWARTZ ET AL. - PREDATOR MANAGEMENT AND MOOSE

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    LARGE CARNIVORES, MOOSE, AND HUMANS: A CHANGINGPARADIGM OF PREDATOR MANAGEMENT IN THE 21st CENTURY

    Charles C. Schwartz1, Jon E. Swenson2, and Sterling D. Miller3

    1Interagency Grizzly Bear Study Team, U.S. Geological Survey, Biological Resources Division,Northern Rocky Mountain Science Center, Montana State University, Bozeman, MT 59717, USA;2Department of Biology and Nature Conservation, Agricultural University of Norway, Box 5014, N-1432 Ås, Norway; 3National Wildlife Federation, 240 North Higgins, Suite #2, Missoula, MT 59802,USA

    ABSTRACT: We compare and contrast the evolution of human attitudes toward large carnivoresbetween Europe and North America. In general, persecution of large carnivores began much earlierin Europe than North America. Likewise, conservation programs directed at restoration andrecovery appeared in European history well before they did in North America. Together, the patternsuggests there has been an evolution in how humans perceive large predators. Our early ancestorswere physically vulnerable to large carnivores and developed corresponding attitudes of respect,avoidance, and acceptance. As civilization evolved and man developed weapons, the balanceshifted. Early civilizations, in particular those with pastoral ways, attempted to eliminate largecarnivores as threats to life and property. Brown bears (Ursus arctos) and wolves (Canis lupus)were consequently extirpated from much of their range in Europe and in North America south ofCanada. Efforts to protect brown bears began in the late 1880s in some European countries andpopulation reintroductions and augmentations are ongoing. They are less controversial than inNorth America. On the other hand, there are no wolf introductions, as has occurred in NorthAmerica, and Europeans have a more negative attitude towards wolves. Control of predators toenhance ungulate harvest varies. In Western Europe, landowners own the hunting rights toungulates. In the formerly communistic Eastern European countries and North America, huntingrights are held in common, although this is changing in some Eastern European countries. Wolfcontrol to increase harvests of moose (Alces alces) occurs in parts of North America and Russia;bear control for similar reasons only occurs in parts of North America. Surprisingly, bears andwolves are not controlled to increase ungulates where private landowners have the hunting rightsin Europe, although wolves were originally exterminated from these areas. Both the inability ofscientific research to adequately predict the effect of predator control on ungulate populations anda shift in public attitudes toward large carnivores have resulted in an accelerating number ofchallenges to predator management in places where it is still espoused. Utilitarian attitudes towardswildlife are declining in Western cultures and people now increasingly recognize the intrinsic valueof wildlife, including large predators. In the future, agencies responsible for managing residentwildlife will face increased pressure to balance the needs of the hunting public with the desires ofnon-hunting publics. We suggest that in the next century we will witness a continued shift in howwildlife agencies manage both moose and large carnivores. More attention will be paid tomaintaining and restoring intact ecosystems and less toward sustainable yield of meat.

    ALCES VOL. 39: 41-63 (2003)

    Key words: Alces alces, brown bear, Canis lupus, gray wolf, grizzly bear, moose management,predator control, Ursus arctos

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    There is ample evidence that both wolves(Canis spp.) and bears (Ursus spp.) kill andeat moose. There is also empirical datasuggesting that this predation can limit moosenumbers under certain conditions (Ballardand Van Ballenberghe 1998). Wolf reduc-tion can result in increased numbers ofmoose under some circumstances (Gasawayet al. 1983). However, there are only a fewempirical studies supporting the principlethat bear reduction programs result in en-hancement of moose numbers (Stewart etal. 1985, Ballard 1992). To our knowledge,no study has addressed the long-term ef-fects of bear control on moose numbers.

    In only a few places in North Americadoes control or reduction in wolf and bearnumbers continue to be strongly advocatedby some citizens groups and agencies re-sponsible for wildlife management. A simi-lar attitude regarding wolf control for live-stock safety also exists in Europe. How-ever, in the past 2 decades, agencies havewitnessed increasing levels of concern to-ward and criticism of predator control pro-grams by a more vocal public, particularlythe environmental community. These groupsquestion both the scientific validity and thephilosophical basis for carnivore control.Concurrently, there is an apparent shift inhow society in general values large preda-tors, particularly bears and wolves (Duda etal. 1998). In recent times, a larger contin-gent of the public and scientific communityfinds inherent intrinsic value in carnivoresand perceives their role as necessary inecosystem function (Miller et al. 2001); theyoppose predator control that favors the moreutilitarian attitude of “game production” asthe objective of wildlife management. Theseattitudinal shifts are prevalent in both NorthAmerica and Europe despite dissimilar legalsystems of game management.

    Here, we review the evolution of preda-tor control in North America and Europe,agency culture and big game management,

    the scientific basis of predator manage-ment, and an apparent shift in social valuesin the past decades away from predatorcontrol and toward large carnivore conser-vation and management.

    EARLY HUMAN–PREDATORRELATIONSHIPS

    Three stages between humans and theirenvironment have been described: hunting,shepherding, and agricultural (Boitani 1995).Human attitudes toward large carnivoreshave been shaped by these relationships.Hunting economies were centered on her-bivores as an important source of food.Large predators were perceived as compe-tition, but not as a threat. Hunters hadrespect for and kinship with predators. Thiswas reflected in attitudes of aboriginal peo-ples. Nomadic shepherds disdained wolvesas threats to their livestock and basis forlivelihood. In contrast, sedentary herdershad more tolerant attitudes toward wolvesbecause they had housing to protect theirlivestock. Farmers, producing crops andlimited livestock, had leeway to be moretolerant. However, agricultural people liv-ing in Latin cultures, characterized by closedvillages, were more tolerant than those liv-ing in Germanic societies, characterized bymore open settlements and solitary farms(Breitenmoser 1998).

    Bears and wolves played important rolesin the legends, beliefs, and lives of prehis-toric peoples. Bears were potentially dan-gerous and fearsome creatures to hunt andkill and had physical similarities to man(Rockwell 1991). Like man, bears wereomnivorous, generalists, intelligent animalswith binocular vision. They livedsympatrically with humans and ate many ofthe same foods; when they stood erect orwere skinned out, they shared physical simi-larities to humans (Shepard 1996). Thesecharacteristics may have contributed tomany similar myths and legends involving

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    creatures that were half human and halfbear as a consequence of mating betweenhumans and bears (Rockwell 1991, Shepardand Sanders 1992). Bears disappeared inthe fall and reappeared in the spring givingrise to beliefs that they had a special 2-wayroute to the afterlife and could commutereadily between worlds; an ability worthy ofgreat respect. All these characteristicsgave bears a special importance in earlycultures closely bound to wild animals.Throughout the shared ranges of bears andhumans, people developed elaborate anddetailed rituals to appease the spirits ofbears they killed (Rockwell 1991, Shepardand Sanders 1992, Edsman 1994) and manyof these rituals continue today in some Na-tive American and Eurasian cultures (Black1998). Wolves played a great mythic-reli-gious role because they possessed manysimilar characteristics with humans; theywere great hunters, members of a pack(tribe or clan), defended territories, andhunted cooperatively (Lopez 1978).

    Large carnivores were eliminated frommuch of their former range both in Europeand North America. The extermination ofwolves in Europe started in the Middle Agesand continued well into the early part of the20th century (Mallinson 1978, Boitani 1995).In Great Britain, wolves were considered athreat to livestock and exterminated by the17th century. They were exterminated fromNorthern Europe by the beginning of the20th century, but survived in lower numbersin Southern and Eastern Europe (Boitani1995). The extermination of brown bearsfollowed a similar pattern; bears were extir-pated from Denmark before the Middle Ages(Jessen 1929) and from Britain during the10th century (Corbet and Harris 1991).Bears survived in relict populations in North-ern and Southern Europe, and in greaterpopulations in Eastern Europe (Swenson etal. 2000).

    The eradication of wolves and bears,

    and the Eurasian lynx (Lynx lynx), a preda-tor on small ungulates, was directly causedby persecution, including bounties, and indi-rectly through habitat destruction and elimi-nation of prey. Bounties on wolves wereinitiated in England in the 1500s, on wolvesand bears in Sweden in 1647, and Norway in1733 (Myrberget 1990, Swenson et al. 1995,Elgmork 1996). The destruction of theforests, due to the expansion of cultivationand overgrazing by domestic livestock, wasan important factor, as was the extermina-tion of native herbivores (Breitenmoser1998). For example, by the year 1200, 40%of Switzerland’s forests had been cleared(Breitenmoser 1998). The Hungarian land-scape was converted from 87% forest andwetland around 900 to less than 11% forestby 1920 (Csányi 1997). Also, the Napoleanicwars in the 1800s resulted in the spread ofmodern firearms. The result was the virtualelimination of the remaining big game spe-cies in much of Europe (Breitenmoser 1998),even though over–hunting had contributedto the extinction of the wild boar (Sus scrofa)in England in the 1500s, the capercaille(Tetrao urogallus) in Britain around 1790,and the complete extinction of the auroch(Bos primigenius) by 1627 (Myrberget1990). The famous Swedish taxonomist,Carl von Linné (Linneaus) probably neversaw a wild moose, and his description of thespecies in 1746 was based on a captiveindividual. In 1789, a Swedish law allowedlandowners unrestricted hunting on theirland, with the result that moose were almosttotally exterminated in Sweden by 1825,when the law was repealed (Bergström etal. 1993). With little wild ungulate prey, thelarge carnivores attacked the abundant do-mestic animals, increasing conflicts andpersecution by people.

    When Europeans colonized the NorthAmerican continent, they brought their oldworld culture and traditions with them, in-cluding a view of wilderness as “something

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    alien to man–an insecure and uncomfort-able environment against which civilizationhad waged an unceasing struggle” (Nash1982:8). To early settlers, wilderness wasvillainous as were the wild animals andIndians living in it. Taming wilderness meantthe extermination of large carnivores, par-ticularly wolves and bears. The prevailingattitude of colonial America was summa-rized in a quote from John Adams in 1756:“The whole continent was one continuingdismal wilderness, the haunt of wolves andbears and more savage men. Now theforests are removed, the land covered withfields of corn, orchards bending with fruitand the magnificent habitations of rationaland civilized people” (Kellert 1996:104).The difference between European colonizersand the American Indians in attitude to wildlands and wild places was eloquently phrasedby Sioux Chief Luther Standing Bear (1932,from Deloria 2001): “We did not think of thegreat open plains, the beautiful rolling hills,and winding streams with tangled growth as‘wild.’ Only to the white man was nature a‘wilderness’ and only to him was the land‘infested’ with ‘wild’ animals and ‘savage’people. To us it was tame. Earth wasbountiful and we were surrounded with theblessings of the Great Mystery.”

    As civilization moved westward, thepioneers viewed predators much like thenomadic shepherds described by Boitani(1995). Since the prevailing form of live-stock husbandry was to allow large herds ofcattle and sheep to graze freely over vastareas, carnivores, particularly wolves andgrizzly bears, were considered an economicthreat. The pervasive attitudes of the timewere captured in 2 salient quotes (fromNRC 1997:135). Historian and trapperStanley Young wrote: “There was sort of anunwritten law of the range that no cow manwould knowingly pass by a carcass of anykind without inserting in it a goodly dose ofstrychnine sulfate, in the hope of killing one

    more wolf” (Young 1946:27). An earlydirector of the U.S. Biological Society, E.A. Goldman, wrote, “Large predatory ani-mals destructive of livestock and game, nolonger have a place in our advancing civili-zation” (Dunlap 1988:51). An early Ameri-can stockman had similar views: “Thedestruction of these grizzlies is absolutelynecessary before the stock business…couldbe maintained on a profitable basis.” (Bai-ley 1931 cited in USFWS 1993).

    With the exception of extreme northernMinnesota, wolves were eliminated fromthe conterminous 48 states by the 1900s(Boitani 1995). Between 1800 and 1975,grizzly bears were eliminated from nearly98% of their historic range (USFWS 1993,Mattson et al. 1995). At the time of theLewis and Clark expedition, grizzly bearsinhabited most of the western United Statesand extended out into the Great Plains(Servheen 1999). They flourished wherePacific salmon (Oncorhynchus spp.) wereabundant far inland into eastern Idaho.Today, they exist as only 5 remnantpopulations south of Canada. Three ofthese populations contain 350 individuals (Servheen1999). One population, in the north Cas-cades along the Pacific coast, is highlyendangered in both the United States andCanada. No bears have been verified onthe United States side of the border inrecent decades.

    Bears and wolves faired better north ofthe 49th parallel. The chronology of wolfextirpation in southern Canada followed thepattern of agricultural and industrial settle-ment (Hayes and Gunson 1995). Wolveswere extirpated from many areas in theeastern provinces by the early 1900s, andsignificantly reduced in the western prov-inces by the 1930s (Carbyn 1987, Hayesand Gunson 1995). Grizzly bears followeda similar pattern. They were extirpatedfrom part of their historic range in Mani-

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    toba, Saskatchewan, and Alberta, primarilyin the prairies and boreal plains and arescarce in southern Alberta and British Co-lumbia, where human populations are con-centrated (Macey 1979, Banci 1991, Banciet al. 1994, McLellan and Banci 1999).

    Even in Alaska, where wolves andbrown and black bears are still abundant,there were several attempts to eliminatethem. According to Sherwood (1981:24)“Periodically Alaskan civic leaders advo-cated the extermination of the animals[brown bears] and make Ursus the symbolof an alleged colonialism that they claimedwas inspired by conservationist sentimentand directed by bureaucrats in WashingtonD.C., working in concert with vested ab-sentee interests. This colonialism, theybelieved, prevented resident Alaskan en-trepreneurs from exploiting the Territory’snatural resources and prevented residentpoliticians from setting the terms of thatexploitation.”

    Brown bears were perceived as a di-rect threat to the fledgling cattle industry onKodiak Island (Van Daele 2003). Politicalpressure from the industry resulted in regu-lations in 1929 allowing Kodiak cattlemen tokill bears at any time they were considereda menace to livestock or property. Accord-ing to Sherwood (1981:59) one rancher’sadvice to anyone encountering a brownbear was, “shoot them in the guts, in thefoot, any place, but get a bullet into them.”

    Prior to statehood in 1959, wolves inAlaska were targeted in a major predatorcontrol program led by the United StatesGovernment. Wolf control was pervasivewith the intent to increase moose and cari-bou populations. Strychnine and cyanidewere commonly used and later aerial gun-ning was employed as a very effectivetechnique. Following statehood, state man-agement eventually evolved to where thewolf was listed as a game animal in 1963(NRC 1997).

    As civilization expanded and humandensities increased, predators were eithersignificantly reduced or eliminated frommuch of their range in both North Americaand Europe (Woodroffe 2000). As we shallsee, these early attitudes toward large car-nivores have persisted, to varying degrees,around the world.

    EARLY DEVELOPMENT OFWILDLIFE MANAGEMENT

    SYSTEMSThroughout most of the historical record,

    wildlife managers have targeted predatorsand have had a significant impact on theirnumbers, distribution, and more recently,conservation. Because wildlife has valuefor recreation and food, there are signifi-cant economic incentives for individuals,such as landowners, to acquire propertyrights to wildlife. Thus, the system ofwildlife management in a country can con-tribute to the attitudes towards predation bylarge carnivores on big game (Lueck 1995).

    EuropeIn most of Europe, the Kings and their

    chieftains controlled most of the lands andhunting rights during this feudal period, whichwas at its peak around 1000. The oldestgeneral game law, introduced by King Knutof Denmark in 1016, established that no oneowned wild animals, but that the king hadsome hunting privileges. This system de-clined as a result of corruption and theBlack Death, and around 1348-49, huntingprivileges, especially for big game, weretransferred to the large landowners. Thiswas completed in Europe by the 17th and18th centuries. In some countries, such asFinland, Norway, and Switzerland, smallfarmers and others held many hunting rights(Myrberget 1990). The Norwegian Parlia-ment transferred all hunting rights, exceptfor killing large carnivores, to landowners in1899 (Søilen 1995).

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    Landowners often lost exclusive hunt-ing rights following changes in political sys-tems, such as the revolutions in France andRussia and the introduction of communismin Eastern Europe. Interestingly, the deci-sion that hunting rights in the United Stateswould not be held by the landowner, as is thecase in Britain, was made following theAmerican Revolution. After World War II,Europe had essentially 2 systems, with land-owners having the hunting rights in WesternEurope and governments managing huntingin the communistic Eastern Europe. Hunt-ing remained open to all citizens of Portugal,Italy, Greece, and Turkey (Myrberget 1990).Since the fall of communism, countries ofEastern Europe have been in flux aboutwhether to revert to the former system oflandowners owning hunting rights, as Hun-gary has, or to retain state control overwildlife management, independent of landownership, as has Poland, Slovakia, Ukraine,and Romania (Csányi 1997, Salvatori et al.2002).

    North AmericaIn North America, prior to European

    colonization, the American Indian tribesclaimed rights to wildlife by protecting hunt-ing and fishing territories (Carlos and Lewis1995). “The ownership of game amongNative Americans had an uncanny resem-blance to current United States institutions.Indian tribal societies, like state agencies,controlled wildlife stocks by enforcing therights to hunting and fishing territories andrestricting the time and method of harvestby tribal members.” (Lueck 1995:3).

    American game laws are rooted in thepast history of English common law. How-ever, today’s American and English wildlifelaws are markedly different. In the UnitedStates, ownership of wildlife resides withthe people and is administered on theirbehalf by government, primarily the stategovernments. In Great Britain, the law

    places nearly all control in the hands ofprivate landowners. In Canada, wildlife ismanaged by the Provincial Governments onbehalf of the people similar to the UnitedStates, but ownership is vested in the Crownuntil the wildlife is legally killed. At thispoint property rights transfer to the hunter.The states and provinces have retainedcontrol over most wildlife management, butthey have lost some authority to the federalgovernments with international treaties (e.g.,the Migratory Bird Treaty Act and MarineMammal Protection Act) and, in the UnitedStates, with the Lacey Act (controls inter-state transportation of game) and the En-dangered Species Act. But the states andprovinces have vigorously fought to retainthe authority to manage wildlife (Peek 1986).

    CONFLICTS SURROUNDINGPREDATOR CONTROL TO

    IMPROVE HARVEST OF WILDUNGULATES

    In his classic book “Game Manage-ment,” Leopold (1933:3) defined game man-agement as “the art of making land producesustained annual crops of wildlife for rec-reational use.” This definition espoused autilitarian philosophy of game managementthat established the direction of wildlifemanagement for the next half century.Wildlife agencies “managed” gamepopulations for a “sustained yield.” Theirprimary clients were the hunting public, andup until the 1970s virtually all state andprovincial wildlife agencies operated pri-marily under the principle of sustained use.Most universities that trained students inthe field of wildlife management were LandGrant or Agricultural colleges with a focuson production and emphasized sustainableyield concepts.

    Peek (1986:25), discussed 2 groups ofconservationists as defined by Harry et al.(1969): those with a conservation-utilizationemphasis, and those with a conservation-

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    preservation emphasis. Both groups wereconcerned with the perpetuation of naturalresources and therefore could be classed asconservationists. However, people with autilization emphasis were oriented towardthe goal of resource exploitation, such ashunting, with aims of producing sustainedyields by cropping surpluses. “Wise use”was the doctrine of those with conserva-tion-utilization emphasis and their philoso-phy was adopted by most wildlife and natu-ral resource management agencies. Thiswas encouraged by the importance of feespaid by hunters that were vital to manage-ment activities including the salaries of themanagers. Conservationist-preservation-ists, by contrast, were not oriented towards“wise use” but rather espoused an appre-ciative interest in the resource, preferablyin its “natural state” (Harry et al. 1969).State wildlife agencies were slow to ac-knowledge this doctrine in part, perhaps,because there was no mechanism for con-servation-preservationists to regularly sup-port wildlife management efforts with theirfees.

    The conservationist-preservationistmovement in North America greatly in-creased in influence in the early 1970s andbecame a major part of the biopoliticalscene during the 1980s. It was largelyresponsible for broadening activities of wild-life and land management agencies innongame management and has evolved intowhat is termed environmentalism (Peek1986). Conservation-preservationists wereand still are largely responsible for chal-lenging predator control programs.

    Alaska has carried out a program ofwolf control with the specific goal of in-creasing ungulate populations for huntersduring the same period that conservationand enhancement programs were underwayelsewhere in the United States. The historyof predator control in Alaska provides agood example of the conflicts between the

    wise use and the conservation-preservationgroups. During the decade of the 1980s,controversy grew around the state’s wolfcontrol programs (Stephenson et al. 1995),and environmental groups filed several courtcases against the states in an attempt tostop wolf control. These conflicts reacheda peak in the early 1990s. By 1994, thegovernor of Alaska suspended the state’swolf control program because it was judgedto be an unacceptable treatment of wolves(NRC 1997). The governor called for ascientific review and indicated that he wouldnot reinstate predator control unless it met3 criteria: (1) it was based on solid science;(2) a full cost-benefit analysis showed itmade economic sense for Alaskans; and (3)it had broad public support. Similar con-frontations over wolf control programs havealso occurred in Canada (Hayes and Gunson1995).

    Alaska began efforts to reduce grizzlybears in some areas in 1980 in response toindications that moose numbers in certainareas were being maintained at chronicallylow levels by bear predation on neonates.Moose numbers declined in the 1970s as aconsequence of a series of severe wintersand overharvest by hunters, in addition tosignificant predation on moose calves bybears (Ballard et al. 1981). Unlike effortsto control wolves, grizzly bear control ef-forts in selected areas were done by liber-alization of hunting regulations instead of bytrapping and shooting by state employees.Bear reductions started in 1980 and ulti-mately included elimination of requirementsthat residents buy a special tag to hunt bearsand generous bag limits that allowed hunt-ers to take bears more frequently in tar-geted areas than elsewhere in Alaska.Moose numbers increased during the 1980sand 1990s but calf survival remained low.Available evidence indicated that increasesin moose numbers were unrelated to theincreased bear harvests (Miller and Ballard

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    1992).Efforts are ongoing in targeted areas of

    Alaska to reduce bear numbers in order toprovide more moose for hunters. Unlikeefforts to reduce wolf abundance to accom-plish the same objective, bear reductionefforts have generated less controversy,perhaps because control efforts are donegradually by legal sport hunters instead ofby government agents using trapping andaerial gunning techniques. Also, unlike thegovernment sponsored control efforts in theUnited States south of Canada in the early20th century, bear reduction by hunters hasnot yet resulted in measurable declines inbear density in the portions of Alaska whereit is ongoing, although it has caused changesin population composition (Miller 1997).

    Control of wolves to enhance huntingopportunity is supported by 48% of Alaskanvoters and by 65% of Alaskan hunters(Miller et al. 1998). The most vehementopposition to wolf control efforts comesfrom conservation groups outside of Alaska,but these groups have been effective atstopping or curtailing active wolf reductionprograms. It is possible that the generalpublic may be more amenable to reductionsin bear abundance than to reductions in wolfabundance because, unlike wolves, bearsoccasionally attack humans, resulting in afear of bears in a significant proportion ofthe population. A survey of Alaska votersindicated that 34% had concerns about bearsthat sometimes kept them from going intothe countryside (Miller et al. 1998).

    Similar to the grizzly bear case inAlaska, black bears (Ursus americanus)and mountain lions (Puma concolor) werethought to be reducing recruitment of elk(Cervus elaphus) in the Clearwater areaof central Idaho. In 2000, a research pro-gram was proposed to evaluate the biologyof this relationship through predator reduc-tion efforts accomplished by liberalized hunt-ing regulations as well as control by govern-

    ment officials. Predator advocacy groupsorganized a national campaign against theproposed research similar to that conductedagainst the wolf reduction effort in Alaska.Idaho officials modified their research pro-posal to accomplish targeted reductions usingonly liberalized regulations for hunting ofpredators. Subsequently, objections to theresearch have largely dissipated even thoughthe liberalized hunting regulations have beenexpanded beyond the boundaries of theoriginally proposed research area. Difficul-ties in convincingly documenting trends inblack bear and mountain lion abundance arelikely to confound interpretation of anychanges in elk recruitment or abundancefound in the Idaho study.

    We know of no case of planned reduc-tion in bear densities in Europe to increaseungulate numbers. The European brownbear is much less aggressive towards hu-mans than the North American brown/griz-zly bear (Swenson et al. 1996), which mayalso influence people’s attitudes. The per-secution of wolves in recent times has beenjustified partially by the reduction of preda-tion on ungulates in some areas. Neverthe-less, we are unaware of any case where thiswas the primary justification, as reductionsof livestock losses seem to be the primaryobjective (Promberger and Schröder 1993).We expected that planned reductions ofthese predators would have occurred in thecountries where landowners own the eco-nomic rights to hunting. However, in suchcountries, bears seem to be popular enoughfor hunting in their own right that owners ofthe hunting rights accept this tradeoff. Nocountry with this system has a huntablepopulation of wolves, which might not be acoincidence. However, wolves recentlyhave been established in Norway and Swe-den, where this system exists. In Norway,the state pays compensation for the loss ofhunting income due to wolf predation, but itis a country where conflicts surrounding

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    depredation on free-grazing sheep are veryintense. In Sweden, where there are almostno free-grazing sheep, large forest compa-nies are contributing to the funding of wolfresearch because they want to know ifwolves can help reduce forest damagecaused by high moose densities.

    STUDIES OF PREDATOR–PREYRELATIONSHIPS

    Ballard and Larsen (1987), VanBallenberghe (1987), Boutin (1992), NRC(1997), and Ballard and Van Ballenberghe(1998) all provide thorough reviews of therecent studies of predator-prey dynamics ofmoose in North America; only 1 intensivestudy has been conducted in Europe(Swenson et al. 2001). Wolves, brownbears, and American black bears are theprincipal predators of moose. There isgeneral agreement that predation is a limit-ing factor for moose populations, but thereis controversy regarding the magnitude ofthis limitation and if the evidence supportsthe hypothesis that predation regulatesmoose numbers (Boutin 1992). Many of thepredator-prey studies dealing with carni-vore- (wolves and bears) moose relation-ships were not designed to answer specificquestions about the impacts of predatorcontrol on moose demographics. Addition-ally, the full impact of wolf predation andwolf control on moose demographics hasreceived considerably more attention thansimilar impacts from bears. Few studiesdetail the influences of both bear and wolfpredation on moose in the same system atthe same time (Table 1). Even fewerstudies have been conducted long enough todetermine the long-term impacts of preda-tor control on moose population dynamics.In a review of predator control in Alaska,the NRC (1997) came to 3 conclusions: (1)predator control experiments provide onlynegative evidence for the existence of analternative stable state with relatively high

    numbers of both predators and prey. Only2 studies were monitored long enough toreveal the existence of such a state, and theevidence from those studies was negativeor equivocal. Existing evidence suggeststhat if predator control is to be used as a toolto increase ungulate populations, controlmust be both intensive and relatively fre-quent. There is no factual basis for theassumption that a period of intensive controlfor a few years can result in long-termchanges in ungulate population densities;(2) experiments that resulted in increases inmoose populations were conducted wherewolves were relatively numerous, wherebears were relatively uncommon and werenot preying heavily on ungulate calves, wherehabitat quality was high, and weather wasrelatively benign. The evidence is inconclu-sive, but there is reason to believe that anintensive control effort, during which wolfpopulations are greatly reduced for severalyears and other factors are favorable, canresult in short-term increases in moosepopulations; and (3) control experimentsthat appeared to have had some successused methods, such as aerial shooting, thatare not currently politically acceptable.

    During the 1970s and early 1980s, wolfremoval had strong support among manage-ment agencies in Alaska and the Yukon asa means of increasing moose densities(Gasaway et al. 1983, Ballard and Larsen1987). However, some wolf control experi-ments met with limited success, and telem-etry studies during the same period(Franzmann et al. 1980, Ballard et al. 1981,Franzmann and Schwartz 1986, Boertje etal. 1988, Ballard and Miller 1990) impli-cated bears as a limiting factor in calfsurvival. Consequently, emphasis shiftedtoward both wolf and bear control pro-grams, yet there was little empirical data tosupport bear control as a long-term man-agement tool to increase moose numbers(Table 1). Current science allows manag-

  • PREDATOR MANAGEMENT AND MOOSE - SCHWARTZ ET AL. ALCES VOL. 39, 2003

    50

    Tab

    le 1

    . Pre

    dato

    r red

    uctio

    n ex

    peri

    men

    ts (f

    rom

    NR

    C 1

    997)

    .

    Pred

    ator

    redu

    ctio

    nM

    etho

    d an

    dlo

    catio

    nW

    olve

    sB

    ears

    Res

    ults

    Air

    -ass

    iste

    dE

    ast-c

    entra

    lW

    olf

    popu

    latio

    n re

    duce

    dN

    ot d

    one

    AK

    to 5

    5-80

    % b

    elow

    pre

    -con

    trol

    (GM

    U 20

    A)

    num

    bers

    for 7

    yrs

    (197

    6-82

    ).

    Finl

    ayso

    n,49

    -85%

    of w

    olf p

    opul

    atio

    nN

    ot d

    one

    Yuk

    onre

    mov

    ed fo

    r 6 y

    rs (1

    983-

    89);

    hum

    an h

    arve

    st ra

    te o

    f moo

    sean

    d ca

    ribo

    u re

    duce

    d by

    90%

    .

    Sout

    hwes

    tW

    olf n

    umbe

    rs re

    duce

    d by

    Bea

    r pop

    ulat

    ion

    Yuk

    on40

    -80%

    for 5

    yea

    rsre

    duct

    ion

    estim

    ated

    (Ros

    e Lak

    e; 1

    982-

    87).

    at 7

    -9%

    (198

    2-87

    ).

    Ais

    hihi

    k,A

    ppro

    xim

    atel

    y 76

    % o

    f wol

    fN

    ot d

    one

    Yuk

    onpo

    pula

    tions

    rem

    oved

    ove

    r4

    yrs (

    1993

    -96)

    . M

    oose

    and

    cari

    bou

    hunt

    ing

    curt

    aile

    d.

    Nor

    ther

    n B

    C1,

    000

    wol

    ves r

    emov

    ed in

    Not

    don

    e10

    yea

    rs (1

    978-

    87);

    alm

    ost

    800

    of w

    hich

    wer

    e rem

    oved

    in th

    e la

    st 4

    yrs

    of

    rem

    oval

    .

    Qué

    bec

    Wol

    f po

    pula

    tion

    redu

    ced

    toA

    tota

    l of 8

    1 be

    ars

    rem

    oved

    48-6

    2% o

    ver 4

    yea

    rs (1

    982-

    85).

    over

    a 3

    yr p

    erio

    d in

    adi

    ffer

    ent a

    rea (

    1983

    -198

    5).

    Ave

    rage

    ann

    ual r

    ate

    of in

    crea

    se o

    f moo

    se p

    opul

    atio

    ns w

    as 1

    5% d

    urin

    gw

    olf c

    ontr

    ol, a

    nd 5

    % fo

    r 12

    yrs

    afte

    r the

    end

    of w

    olf c

    ontr

    ol.

    Ave

    rage

    annu

    al ra

    te o

    f inc

    reas

    e of

    car

    ibou

    pop

    ulat

    ions

    was

    16%

    dur

    ing,

    and

    6%

    for 7

    yrs

    aft

    er th

    e en

    d of

    wol

    f con

    trol

    .

    Incr

    ease

    d su

    rviv

    al o

    f adu

    lt ca

    ribo

    u; in

    crea

    sed

    num

    bers

    of c

    alve

    s/10

    0co

    ws

    for b

    oth

    moo

    se a

    nd c

    arib

    ou.

    Ave

    rage

    ann

    ual r

    ate

    of in

    crea

    se fo

    rm

    oose

    and

    car

    ibou

    abo

    ut 1

    6-18

    %.

    Hun

    ting

    succ

    ess

    incr

    ease

    d. S

    even

    year

    s af

    ter w

    olf c

    ontr

    ol e

    nded

    , moo

    se a

    nd c

    arib

    ou n

    umbe

    rs b

    egan

    tode

    clin

    e. N

    o su

    bsta

    ntia

    l inc

    reas

    e in

    moo

    se p

    opul

    atio

    ns o

    r cow

    :cal

    f rat

    ios

    duri

    ng p

    reda

    tor r

    emov

    al.

    No

    subs

    tant

    ial i

    ncre

    ase

    in m

    oose

    pop

    ulat

    ions

    or c

    ow:c

    alf r

    atio

    s du

    ring

    pred

    ator

    rem

    oval

    .

    Incr

    ease

    d nu

    mbe

    rs o

    f car

    ibou

    cal

    ves/

    100

    cow

    s. R

    espo

    nse

    of m

    oose

    high

    ly v

    aria

    ble

    and

    not c

    lear

    ly re

    late

    d to

    wol

    f red

    uctio

    n. C

    ontr

    ol e

    nded

    in 1

    996,

    too

    rece

    ntly

    to a

    sses

    s lo

    ng-t

    erm

    tren

    ds.

    Cal

    f sur

    viva

    l rat

    es a

    nd p

    opul

    atio

    n si

    zes

    appa

    rent

    ly in

    crea

    sed

    for a

    llun

    gula

    tes

    in th

    e ar

    ea.

    Whe

    n w

    olf c

    ontr

    ol e

    nded

    , wol

    f num

    bers

    incr

    ease

    dra

    pidl

    y an

    d ca

    lf s

    urvi

    val d

    ecre

    ased

    to p

    re-c

    ontr

    ol le

    vels

    .

    No

    appa

    rent

    cha

    nge

    in m

    oose

    cal

    f sur

    viva

    l rat

    e in

    eith

    er w

    olf o

    r bea

    rre

    mov

    al ar

    ea.

  • ALCES VOL. 39, 2003 SCHWARTZ ET AL. - PREDATOR MANAGEMENT AND MOOSE

    51

    Eas

    t-cen

    tral

    Wol

    f po

    pula

    tion

    was

    red

    uced

    Not

    don

    eA

    K b

    y 28

    -58%

    for 3

    yrs

    (198

    1-84

    ).(G

    MU

    20E)

    Sout

    h-ce

    ntra

    lE

    xten

    sive

    aer

    ial s

    hoot

    ing

    Pois

    onin

    g pr

    obab

    lyA

    Kan

    d po

    ison

    ing

    redu

    ced

    also

    red

    uced

    bea

    r(G

    MU

    13)

    wol

    f num

    bers

    dra

    mat

    ical

    lynu

    mbe

    rs.

    (194

    8-19

    54).

    Wol

    ves

    wer

    e re

    duce

    d by

    Aft

    er w

    olf c

    ontr

    ol42

    -58%

    for 3

    yrs (

    1976

    -78)

    .en

    ded,

    60%

    of t

    hebe

    ar p

    opul

    atio

    nw

    as tr

    ans-

    loca

    ted

    orre

    duce

    d by

    libe

    raliz

    edhu

    ntin

    g re

    gula

    tions

    .G

    roun

    d-ba

    sed

    Ken

    aiR

    ecre

    atio

    nal

    harv

    ests

    Peni

    nsul

    a,in

    crea

    sed

    for 4

    yrs

    AK

    (197

    6-79

    ).

    Van

    couv

    erW

    olf

    dens

    ity w

    as r

    educ

    edN

    ot d

    one

    Isla

    nd, B

    Cov

    er a

    4-yr

    per

    iod

    (198

    3-86

    )to

    abo

    ut 1

    0% o

    f pre

    -con

    trol

    leve

    ls.

    Sask

    atch

    ewan

    Not

    don

    eU

    nkno

    wn

    prop

    ortio

    n of

    bla

    ckbe

    ar p

    opul

    atio

    n w

    as re

    mov

    edin

    spri

    ng o

    f 198

    3, a

    nd fr

    oman

    othe

    r are

    a in

    spri

    ng o

    f 198

    4.

    Tab

    le 1

    . con

    tinu

    ed.

    Pred

    ator

    redu

    ctio

    nM

    etho

    d an

    dlo

    catio

    nW

    olve

    sB

    ears

    Res

    ults

    Wol

    f con

    trol

    had

    no

    mea

    sura

    ble

    effe

    ct o

    n ca

    lf s

    urvi

    val.

    Dur

    ing

    and

    afte

    r thi

    s pr

    edat

    or re

    duct

    ion

    peri

    od, c

    arib

    ou n

    umbe

    rsin

    crea

    sed

    and

    had

    mor

    e th

    an d

    oubl

    ed b

    y th

    e ea

    rly

    1960

    s. T

    his

    coin

    cide

    d w

    ith fa

    vora

    ble

    wea

    ther

    and

    rang

    e co

    nditi

    ons

    and

    low

    harv

    est

    by h

    uman

    s.

    Wol

    f con

    trol

    did

    not

    resu

    lt in

    hig

    h an

    nual

    incr

    ease

    s in

    the

    moo

    sepo

    pula

    tion.

    Cal

    f sur

    viva

    l inc

    reas

    ed a

    fter

    bea

    r rem

    oval

    , but

    bea

    rsre

    turn

    ed to

    the

    area

    aft

    er s

    umm

    er a

    nd m

    oose

    cal

    f sur

    viva

    l ret

    urne

    d to

    leve

    ls b

    efor

    e be

    ar re

    mov

    al.

    No

    chan

    ge in

    cal

    f num

    bers

    cou

    ld b

    eat

    trib

    uted

    to in

    crea

    sed

    bear

    har

    vest

    s.

    Wol

    f re

    duct

    ions

    wer

    e as

    soci

    ated

    with

    incr

    ease

    d m

    oose

    pop

    ulat

    ions

    ,bu

    t the

    hig

    hest

    num

    ber

    of m

    oose

    occ

    urre

    d in

    are

    as th

    at h

    ad b

    urne

    d10

    -25

    year

    s ear

    lier r

    egar

    dles

    s of t

    he e

    xten

    t of w

    olf r

    educ

    tions

    .

    Dee

    r pop

    ulat

    ions

    incr

    ease

    d fo

    llow

    ing

    wol

    f con

    trol

    . H

    unte

    r eff

    ort

    appe

    ared

    to b

    e en

    hanc

    ed b

    y w

    olf c

    ontr

    ol.

    Incr

    ease

    d ca

    lf:c

    ow ra

    tios a

    fter

    bea

    r rem

    oval

    . T

    he fo

    llow

    ing

    year

    ,pr

    opor

    tion

    of y

    earl

    ings

    in th

    e m

    oose

    pop

    ulat

    ion

    was

    hig

    her t

    han

    befo

    re p

    reda

    tor c

    ontr

    ol.

  • PREDATOR MANAGEMENT AND MOOSE - SCHWARTZ ET AL. ALCES VOL. 39, 2003

    52

    ers to predict with reasonable confidencethat predator removal at low moose densi-ties can improve calf survival, but increasedrecruitment and subsequent growth of themoose population may or may not occur.Data are generally unavailable on long-terms effects of predator reduction.

    TIMES CHANGE: FROMPREDATOR EXTERMINATION TO

    PREDATOR CONSERVATIONEurope

    Because of the degradation of forestsand loss of indigenous large herbivores, theattitudes of Europeans towards the environ-ment were changing by the end of the1800s. In Switzerland, laws requiring refor-estation, banning livestock from forests,restricting hunting, and establishing gamesanctuaries were passed in 1876(Breitenmoser 1998). Similar laws, and anincrease in effort to reintroduce ungulatesinto former habitats, occurred throughoutEurope. The first modern law regulatinghunting and protection of game in Norwaywas passed in 1845 (Søilen 1995). TheSwedish Hunter’s Association, the Norwe-gian Association of Hunters and Anglers,and the Swiss League for the Protection ofNature were founded in 1830, 1877, and1909, respectively. During this same timeperiod when forests and ungulate populationswere increasing, human numbers were de-clining in rural areas as people migrated toindustrial cities or emigrated to NorthAmerica. As a consequence, numbers oflivestock also declined (Breitenmoser 1998).Efforts to increase ungulate populations havebeen spectacular, and at present, manyEuropean countries have high densities. Forexample, in the mid-1980s, nearly 250,000moose were harvested annually in the Nor-dic countries, compared with about 72,000in all of North America (Haagenrud et al.1987, Kelsall 1987). Most of these greatincreases occurred in the absence of large

    carnivores. Today, forest damage causedby abundant ungulates is a widespread prob-lem in Europe (Bergström et al. 1993,Breitenmoser 1998) and this over-browsinghas reduced the biodiversity of plants andinvertebrates (Suominen et al. 1999).

    During this period, steps were taken inseveral countries to save the remainingbrown bears. In Sweden, for example,official requests to remove the bounty onbears were made in 1889 and 1891, thesecond by a chapter of the Swedish Hunt-er’s Association. Parliament approved theserequests in 1893 (Lönnberg 1929). In 1905the Royal Swedish Academy of Sciencesissued a statement saying, “it is a matter ofhonor for our country that this interestinganimal be protected from complete exter-mination.” Several measures were taken toprotect bears, including complete protec-tion in national parks, which were first es-tablished in 1910. This effort was success-ful, and hunting was reintroduced in 1943(Swenson et al. 1995). Brown bears re-ceived protection in Poland in 1932 and inItaly in 1939 (1992 in the Abruzzo area).But, the bear received protection muchlater in other countries: 1955 in France,1967 in Spain, and 1972 in Norway (Servheenet al. 1999). Efforts to save and increasebear populations in Europe have been suc-cessful in many areas, and there are nowabout 50,000 brown bears in Europe (ca.14,000 outside of Russia) with increasingand expanding, or at least stable, populationsfound in Northeastern Europe, theCarpathian Region, the Dinaric MountainRange in former Yugoslavia, and Scandina-via (Zedrosser et al. 2001). In many in-stances, bears are returning to countriesthat exterminated them because of suc-cessful conservation efforts in neighboringcountries. In addition, the population hasbeen increasing in most of Russia, with thegreatest increase in the European part(Chestin et al. 1992). However, 8 of the 12

  • ALCES VOL. 39, 2003 SCHWARTZ ET AL. - PREDATOR MANAGEMENT AND MOOSE

    53

    European populations are less than 500bears and are decreasing. Efforts to savethe brown bear have concentrated on pro-tection, and the species is protected or is agame animal in all of Europe (Zedrosser etal. 2001). Additionally, 2 reintroductionshave been attempted, and 2 populationshave been augmented. The first reintroduc-tion attempt in the world was in Poland in1938-44 and was unsuccessful. A secondoccurred in the central Pyrenees of Francein 1996-97. Augmentations have occurredin Austria in 1989-93 and an ongoing projectin Italy that started in 1999.

    Brown bears are an important and prizedbig game animal in many countries. This isprobably why Swedish hunters worked ac-tively for the species’ protection in the1800s. It is also an important game speciesin many Eastern European countries(Salvatori et al. 2002). Zedrosser et al.(2001) concluded that communism in East-ern Europe was not nearly as destructive tobear populations as the political systems inWestern Europe, possibly because bearswere managed for hunting by a few hunters,including foreign hunters with convertiblecurrency, and because gun ownership wasstrictly limited. The extreme consequenceof this was the situation in Romania, wherethe dictator Nicolae Ceausescu allowed thebear population to increase to the highestdensities in Europe to provide hunting op-portunities, such as shooting 24 bears in onehunt, and trophies for himself only (Crisan1994). Romania is the only European coun-try that has decided to reduce its brownbear population, from 8,000 to 6,000 byhunting, in order to reduce loss of human lifeand livestock losses (Servheen et al. 1999,Swenson et al. 2000).

    The situation for the wolf is quite differ-ent, even though some Russian scientistsbegan writing about the importance of thewolf’s place in nature at the end of the 19thand beginning of the 20th centuries (Bibikow

    1990). Protection came much later: 1966 inSweden, 1973 in Norway, 1976 in Italy,1993 in France, and 1995 in Croatia andGreece (Promberger and Schröder 1993,Boitani 2000). It is fully protected in 11 of27 European countries and has no protec-tion at all in 9 countries. The total Europeanpopulation outside of Russia is estimated tobe over 18,000, but only 6 countries havemore than 1,000 wolves, 11 have more than500, and 8 have less than 50 (Boitani 2000).Wolves are increasing in many Europeancountries and, like bears, are expanding intocountries where they were formerly extir-pated. In the Soviet Union, wolves weremanaged by zones, with extermination inintensive agricultural and reindeer (Rangifertarandus) husbandry areas, controlled atlow density in areas with fewer people andagriculture, management as a hunted spe-cies in the largest zone, and complete pro-tection in reserves (Bibikow 1990).

    We know of no efforts to reestablishwolves in Europe by reintroductions. How-ever, in addition to the reintroductions andaugmentations of bear populations, lynx havebeen reintroduced into many parts of Eu-rope, and more are planned (Breitenmoseret al. 2000).

    Both the brown bear and the wolf areprotected and managed according to na-tional legislation. In addition, most Euro-pean countries are signatories of the BernConvention, undoubtedly the most impor-tant agreement protecting large carnivoresin Europe. The Bern Convention was rati-fied on 19 September 1979 in Bern, Switzer-land. Its goal is to preserve wild animalspecies and their natural habitats. Membercountries must pay special attention to en-dangered, and potentially endangered, spe-cies listed in different appendices, eachrepresenting a different stage of endanger-ment. The brown bear and wolf are listedin Appendix II (strictly protected faunaspecies) requiring that actions must be taken

  • PREDATOR MANAGEMENT AND MOOSE - SCHWARTZ ET AL. ALCES VOL. 39, 2003

    54

    to protect them; forbidden are the captureor killing; wilful disturbance, and possessionand trade. In addition, the recolonization ofindigenous species must be promoted ifdoing so will enhance the likelihood of pres-ervation. Member countries can make res-ervations to the Bern Convention regardingmeans or methods of killing, capture, orother exploitation of listed species. Sevencountries have made reservations regard-ing protection of the brown bear (Bulgaria,Czech Republic, Finland, Slovenia, Slovakia,Ukraine, and Turkey) and 10 regarding thewolf (Bulgaria, Czech Republic, Finland,Latvia, Lithuania, Poland, Slovenia, Slovakia,Spain, and Turkey) (Boitani 2000, Swensonet al. 2000).

    In addition, Council Directive 92/43/EEC, Conservation of Natural and WildFauna and Flora (ABL L 206, 22.07.1992),binds member states of the European Union(EU). The main goal of the so-called Flora-Fauna-Habitat Directive is to secure spe-cies diversity by protection of habitats andprotection of wild flora and fauna. Actionsmust be taken by member countries to pre-serve all species and their habitats. Thebrown bear is a priority species of the EU.It is listed in Appendix II (species needingspecially protected areas, except thepopulations in Finland and Sweden) andAppendix IV (strictly protected species;capture, killing, and wilful disturbance notpermitted). Possession, transport, and tradeof Appendix IV species are strictly prohib-ited. The wolf is listed in Appendix II andIV (in both cases except for somepopulations in Spain, Greece, and Finland).Exemptions are given when it can be estab-lished that there is no negative impact tospecies preservation, to prevent seriousdamage to culture and livestock, for publichealth, sanitary, and safety reasons, and forscientific, restocking, and re-colonizationpurposes.

    The European Parliament requests that

    members of the EU consider their resolu-tions, although they are not legally binding.Those relevant to large carnivores are: (1)European Parliament Resolution, 24 Janu-ary 1989 (A2-0377/88, Ser. A), which callsfor immediate steps to favor wolf conserva-tion in all European countries, adopts theInternational Union for Conservation ofNature Wolf Manifesto, and invites theEuropean Commission to expand and pro-vide financial means to support wolf con-servation; (2) European Parliament Resolu-tion, 17 February 1989 (A2-339/88, ABL C69/201, 20.3.1989), which states that theEuropean Commission should promote orcontinue programs to protect the brownbear in the EU. Actions for socio-economicdevelopment should be promoted in returnfor communities with protective measuresfor the brown bear. Systems for beardamage prevention and damage compensa-tion should be developed. A network ofconnected reserves and specially protectedareas should be established (called the“NATURA 2000 Network”); and (3) Euro-pean Parliament Resolution, 22 April 1994(A2-0154/94, ABL C 128/427, 09.05.1994),which states that the European Commissionshould not support and finance developmentthat would have a negative effect on bearpopulations. Protected areas and corridorsfor genetic exchange should be establishedto correct actions that have had negativeimpact on bear populations. Measures toprevent the killing and capture of bears andprotect bear habitat should be undertaken.Financial support for damage compensa-tion, and compensation for economic re-strictions due to bear conservation, shouldbe provided.

    North AmericaWolves and grizzly bears were among

    the first species to be protected under theEndangered Species Act (ESA) in the UnitedStates, signed in 1973. Wolves were listed

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    as endangered in 1974 and grizzly bears asthreatened in 1975. The ESA is probablythe most significant law in any nation de-signed to preserve and maintain biodiversity.The ESA establishes that preservation ofanimal and plant species is a national prior-ity that takes precedence over local inter-ests in wildlife management, over economicinterests, and even over certain rights ofowners of private property (Czech andKrausman 2001). Although the intent of theESA to apply to private lands is clear, thelegal basis for this remains controversialand unresolved to some degree (Sax 2001).Distinct population segments can be listedunder the ESA for species like wolves andbears that are reduced in significant propor-tions of their former range but remain abun-dant elsewhere (e.g., wolves and grizzlybears are not listed in Alaska, only south ofCanada). This national priority for speciesrecovery mandated by the ESA has workedwell in the United States to recover largepredators, like wolves and grizzly bears,which sometimes conflict, or are thought toconflict, with local economic or hunter in-terests. The ESA was amended in 1982, toinclude Section 10(j) designed to reducelandowner opposition to restoration of con-troversial species, like wolves and bears, toportions of their former range from whichthey were extirpated. This is accomplishedby designating such reintroduced populationsas “experimental”. Populations restored as“experimental” are permitted more man-agement flexibility on issues such as takingof nuisance individuals, permitting multipleuses of habitat, and reducing the require-ment for federal review of land manage-ment and use activities (such as logging)that could adversely affect the species inexperimental populations. This “experi-mental” provision was successfully used torestore wolves in Yellowstone National Parkand central Idaho. It was also the key to thereintroduction of red wolves (Canis rufus)

    to the southeast and of Mexican wolves (C.l. baileyei) to the southwest of the UnitedStates. So far, however, Section 10(j) pro-visions have been inadequate to accomplishgrizzly bear restoration in the wildernessareas of central Idaho, which, even with thismanagement flexibility, was opposed bykey politicians in Idaho. This oppositionwas based largely on misguided concernsover the level of physical danger grizzlybears posed to humans; a reprise of miscon-ceptions that existed a century ago.

    A number of programs began in the1990s, directed at long-term conservationand enhancement of large carnivores. Muchof this activity in North America was di-rected at areas where both wolves andbears had either been extirpated by earlycolonization or significantly reduced anddeclared threatened or endangered underthe ESA. Wolves were reintroduced intothe Yellowstone Ecosystem and to centralIdaho in 1995 and have recovered throughimproved management and natural disper-sal in the Midwest (Minnesota, Wisconsin,Michigan). In both places, recovery goalsin terms of population number have beenachieved and proposals are pending todownlist and, ultimately, delist the species.Thanks to the emphasis placed on the spe-cies under the ESA, grizzly bear populationshave also increased in the Yellowstone andNorthern Continental Divide Ecosystemsand proposals to delist the species in andaround Yellowstone National Park are ex-pected as numerical population objectiveshave been achieved.

    In much of North America, there ap-pears to be widespread recognition of thevalue of restoring healthy populations ofpredators like wolves and grizzly bears (Dudaet al. 1998, 2001). However, these attitudesare not universal and some states, notablyIdaho and Alaska, retain strong sentimentsagainst predators in favor of aggressivepredator management designed to reduce

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    or eliminate depredations on livestock andthe wild ungulates favored by hunting inter-ests. These attitudes mirror the anti-preda-tor attitudes that resulted in the near extir-pation of predators in the previous century.Such local opposition from political leadershas, temporarily, blocked efforts to restoregrizzly bears to the Bitterroot Ecosystem incentral Idaho. The 14,800-km2 reintroduc-tion area is designated wilderness and rep-resents the best and largest place to restorea significant new population of grizzly bearsin North America. The reintroduction pro-posal provided an unprecedented level oflocal participation and experimental desig-nation of the restored grizzly population(Fischer and Roy 1998, USFWS 2000,Schoen and Miller 2002). It received wide-spread national and local support from thepublic (Duda et al. 1998, Roy 2001) andfrom all professional wildlife managementgroups who commented. We believe it islikely that public support will ultimately re-sult in grizzly bears being restored to thishabitat. Restoration and recovery for con-troversial and environmentally sensitive spe-cies like grizzly bears requires a collabora-tive approach to build popular support(Servheen 1998). However, as the IdahoBitterroot example demonstrates, even suchapproaches are sometimes insufficient toachieve success because of local negativeattitudes about bears.

    CHANGING SOCIAL VALUESWildlife management evolved originally

    as a means to assure continuation of huntingopportunities. Frequently, attitudes of wild-life managers as well as of hunters reflectthese origins and result in policies, such aspredator control, that conflict with the con-cerns and preferences of the general publicover the conservation of predators. Thechanges in official attitudes towards largecarnivores, from policies of extermination,to those of conservation and enhancement,

    reflect changes in the attitudes of the gen-eral populace (Duda et al. 1998, 2001).Obviously, conflict will arise if the manag-ing authorities are seen to be out-of-stepwith the prevailing public attitudes. Oneexample of this was the liberalization ofblack bear hunting regulations, includingspring hunting, hunting with hounds, andbaiting, in Colorado. Citizens objected tothese regulations based on concerns theywere inhumane, but an intransigent bureauc-racy failed to respond. The result was anoverwhelmingly approved citizens’ initia-tive that set aside these regulations (Beckand Gill 1995, Beck et al. 1995). Similarinitiatives passed in Oregon (Boulay et al.1999) and, for mountain lions, in California.In Alaska, voters approved a citizens’ initia-tive that overturned a regulation that al-lowed persons to take wolves by landingaircraft and shooting them. In British Co-lumbia, opponents of hunting grizzly bearssucceeded in getting a moratorium on griz-zly bear hunting implemented in 2001. Themoratorium on hunting has since been liftedbut it is clear that citizens threaten thecontinuation of grizzly bear hunting withconservation-preservationist attitudes to-ward bears.

    Citizens’ initiatives are not practiced inEuropean countries, but there is an obvioustrend towards more protection of large car-nivores. Also, the killing of a few wolves byState employees shooting them from a heli-copter in Norway in winter 2000-2001 re-sulted in enormous European media cover-age and negative public reactions from otherparts of Europe. Some in Europe are ques-tioning the wisdom of the high densities ofungulates, which cause traffic accidents,forest damage, and reductions in forestbiodiversity.

    These examples, and those providedearlier about opposition to predator controlprograms in Alaska, illustrate aspects of atrend that sociologists have been observing;

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    that utilitarian attitudes towards wildlife aredeclining in Western cultures (Decker et al.1992, 2001). A recent metaanalysis(Williams et al. 2002) of 38 studies aboutpeople’s attitudes towards wolves showedthat people were generally positive to wolves(61%), that age, residence, occupation, edu-cation, and income influenced one’s atti-tude, and that about one-fourth of the peo-ple are neutral. The analysis also found thatwolves are less popular in Europe than inNorth America, as is also suggested by ourreview, based on levels and dates of protec-tion. In Europe, this may be influenced bya historical fear of wolves as a carrier ofrabies that dates back to the Middle Ages(Bibikow 1990), or a fear of the wolf as apotential killer of humans, which has morehistorical support from Europe than NorthAmerica (Linnell et al. 2002). Also, atti-tudes towards wolves vary within Europe(Boitani 1995), even in adjacent areas, asNorwegians have more negative attitudestowards wolves than Swedes (Bjerke et al.2001).

    Wolf biologists working in Scandinaviahave reported the impression of an increasedsupport for wolves, although they could notsubstantiate it (Wabakken et al. 2001).However, the metaanalysis, which coveredstudies from the period 1972 – 2000, did notfind any trend in support for wolves.Williams et al. (2002) predicted increasingsupport for wolves over time due to increas-ing education and urbanization, but stressedthat positive attitudes towards wolves in thegeneral public are often weak and have thepotential to shift rapidly if linked to otherstronger attitudes.

    SUMMARYHuman attitudes toward large carni-

    vores have been shaped by centuries ofcoexistence. These attitudes have changedmarkedly as human civilizations maturedand industrialized. Early peoples lacked the

    appropriate weapons to effectively controllarge predators. These cultures adaptedways to live with carnivores. As manacquired modern weapons, large predatorsand native ungulates were exterminated.We witnessed this first in Europe wheremodern civilization first developed. How-ever a second wave of extermination fol-lowed with the European colonization ofNorth America. Old World values weretransported to the New World. In NorthAmerica, attitudes of predator control werealso adopted as part of the evolving profes-sion of wildlife management, where largecarnivores were still abundant in northernenvironments. Carnivores were perceivedas competitive and a threat to ungulatesharvested by hunters.

    Through history, human values towardlarge carnivores seem to be inversely pro-portional to carnivore abundance. Societytends to value them more when they be-come rare or endangered. Because preda-tors were largely eliminated or reduced toremnant population in Europe, social atti-tudes away from extermination and towardsprotection evolved more quickly. Theseattitudinal shifts are reflected in therecolonization of carnivores back into his-toric habitats. Many citizens and scientistsview this reoccupation as a valuable contri-bution to society and ecological processes.A similar transition has come more slowlyto North America and even today there is anapparent inability by some members of thepublic and some wildlife managers to en-gender the changing social dimension thatvalues predators in a broader context ofecosystem function, rather than an impedi-ment to ungulate management for the hunt-ing public.

    Agencies in North America responsiblefor the management of ungulates and carni-vores are currently faced with conflictingvalues and differing demands. If we canuse information about the wolf as a guide, it

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    is possible to make some general conclu-sions. On the one hand, old values prevailwith a continued emphasis on predator con-trol. However, there appears to be a gradualshift toward carnivore conservation, espe-cially among the more highly educated, ur-ban, youth (Williams et al. 2002). Extrapo-lation of these results into the future suggestthat there will likely be a gradual shift awayfrom negative attitudes toward more posi-tive attitudes as the older population is re-placed (Williams et al. 2002). Game man-agement agencies will need to shift towarda more modern construct that recognizesthe intrinsic value of wild ecosystems andthe wildlife they contain, including largepredators. Today’s biologists and moosemanagers face a difficult challenge of bal-ancing biological principles with a diversearray of social and economic values often inconflict with principles of optimum or maxi-mum sustained yield harvest.

    ACKNOWLEDGEMENTSWe thank K. West and R. Jachowski

    for reviews of the manuscript. H. Reynoldsfor providing literature detailing the Alaskaexperience, and organizers of the Norwayconference for kindly inviting the seniorauthor to present this topic.

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