The gray wolf (Canis lupus) was one of the first highly
visible animals to be included on the U. S. Endangered Species
list. The creature now symbolizes endangered species and has become
the cause celebre of numerous animal-interest groups. Probably
because of the affinity of the wolf to the dog (Canis lupus
familiaris) and certainly because the species has historically
been so persecuted (Young and Goldman 1944), a new mythology about
the wolf has evolved; the vile wolf has been replaced by the unjustly
persecuted wolf.
As this wolf deification took place, remnant wolf populations
were relegated to only the most pristine wilderness of North America
and the least developed parts of the rest of the world. Thus,
both laypeople and resource managers widely believed that wolves
preferred wilderness. The animal came to symbolize wilderness,
"for wolves and wilderness are inseparable . . ." (Theberge 1975:152).
However, the wolf survived only in wildernesses mostly because
it was exterminated everywhere else. After the U. S. Endangered
Species Act of 1973 protected the wolf in the 48 contiguous United
States as of 1974 and public attitudes about wolves improved,
wolves began to colonize a wide variety of habitats and to demonstrate
that they did not require wilderness. The wolf has now begun to
recover in the northern U. S. and in several parts of Europe.
The question of the next decade will not be how to save the wolf,
but rather how best to manage the animal. This paper traces the
history of the wolf's status and recovery and explores the dilemma
of its management.
Originally, gray wolves were distributed throughout the northern
hemisphere in every habitat where large ungulates were found.
Saturating most of the region between 20° N latitude (mid-Mexico
and India) and the North Pole, in temperatures from -40°
to +40° C, the wolf inhabited areas as diverse as Israel
and Greenland.
Every kind of northern ungulate, as well as beavers (Castor
canadensis) and arctic hares (Lepus arcticus), can
serve as prey for wolves, and wolves easily switch their prey
from wild to domestic species. Conflict between wolves and humans
over domestic animals probably became an issue soon after ungulates
were domesticated.
As firearms, poisons, and traps were developed, they were used
ruthlessly against wolves with devastating effectiveness (Young
& Goldman 1944). In Eurasia, most wolf populations reached their
lowest point between the 1930s and the 1960s (Pimlott 1973; Delibes
1990; Promberger & Bibikov 1993). In the more-developed regions
of Eurasia, wolves disappeared except in the central Appenine
Mountains of Italy, the Cantabrian mountains of northern Spain,
the Carpathians of Eastern Europe, the northern parts of the former
Soviet Union, and the central plains and mountainous regions of
Asia. Some populations also remained in the deserts of the Middle
East. In North America, wolf numbers were lowest in the late 1950s.
Populations survived primarily in Canada and Alaska (Mech 1970).
In the 48 contiguous United States only the wilderness of northern
Minnesota and nearby Isle Royale National Park in Lake Superior
held wolves.
The environmental revolution ushered in the first endangered species
legislation in the U.S, the Endangered Species Act of 1966. This
act did not protect endangered species but only encouraged federal
agencies to give them special consideration and to promote their
recovery.
At this time, about the only information available on wolves
was anecdotal and hearsay. Historical notes by Young and Goldman
(1944) and Murie's (1944) field study on Mt. McKinley wolves were
practically the only available published information. A few more
studies followed. After the considerable publicity produced by
Durward Allen's seminal investigation of the wolves and moose
of Isle Royale National Park, published in National Geographic
(Allen & Mech 1963) wolf studies proliferated. In 1967, the first
wolf symposium was held by the American Society of Zoologists,
culminating in the publication of the proceedings in the May 1967
issue of American Zoologist. By then the full force of
the environmental movement could be felt. Private wolf organizations
sprang up in many areas, and the wolf quickly gained a popular
constituency in the U.S. and abroad.
In Italy, Luigi Boitani and Eric Zimen pioneered a study of
the wolf in the Abruzzo Mountains east of Rome (Zimen 1981; Boitani
1986). The World Wildlife Fund and the International Union for
the Conservation of Nature and Natural Resources (IUCN), now the
World Conservation Union, took great interest in the wolf, and
the animal was listed in IUCN's Red Data Book of endangered species.
The IUCN Wolf Specialist Group was formed in 1973 (Pimlott 1975).
Meanwhile radio-tracking was developed in the early 1960s (Cochran
& Lord 1963), a technique especially valuable to wolf research.
Wolves were difficult to study with traditional methods because
they were restricted to wilderness areas, highly elusive, and
low in population density. Kolenosky and Johnston (1967) first
radio-tracked wolves in Ontario. Mech and Frenzel (1971) then
combined that technique with aerial tracking and observation,
and numerous studies using these techniques followed.
The second U. S. Endangered Species Act passed in 1973 and protected
the wolf in the contiguous 48 United States beginning in August
1974. Recovery teams were appointed by the U. S. Fish and Wildlife
Service for three wolf subspecies, the eastern timber wolf, the
northern Rocky Mountain wolf, and the Mexican wolf, as well as
the red wolf (U.S. Fish and Wildlife Service 1975, 1982a, 1982b,
1987). At first many wolves were killed illegally (Mech 1977),
but eventually that number dropped (Fuller 1989), and wolf reservoir
populations in less accessible areas expanded (Fuller et al. 1992)
They first recolonized the more remote areas in their surroundings,
reinforcing the view that they were creatures of the wilderness.
Much of the public misinterpreted the wolf's endangered status
in the 48 contiguous states, thinking it meant no wolves were
left anywhere else in the world. Private groups began to raise
wolves to help restore populations, not realizing that Canada
alone supported 50,000 of them. The wolf's apparent dependence
on the wilderness was quantified in the 1970s and 1980s using
road density as a measure. Roads were the routes by which the
public and the government had been able to reach wolves to kill
them. Thiel (1985) found that recolonizing wolves in Wisconsin
lived only where the road density was 0.6 km per km2, a figure
corroborated for Michigan (Jensen et al. 1986) and Minnesota (Mech
et al. 1988). The wolf then officially became a wilderness animal,
and road densities became the yardstick by which wolf habitat
suitability was measured by agencies and recovery teams.
As more was learned about the wolf, the increasingly urbanized
public continued to favor wolf recovery. Even though illegal taking
of wolves persists in local areas of North America and Europe,
it has not been sufficient to prevent wolf population growth.
In Minnesota, some 75 percent of the public viewed the wolf favorably
(Kellert 1986), a statistic that may be mirrored in much of the
northern hemisphere.
Minnesota's wolf population, now probably about 2000 based on
trend estimates by Fuller et al. (1992), proliferated into neighboring
Wisconsin and Michigan (Thiel 1978; Mech et al. 1995b), where
they currently number over 100 (Mech et al. 1995a). Other Minnesota
wolves eventually spread into the Dakotas (Licht & Fritts 1994).
Canadian wolves were no longer killed when they reached Montana,
and they began to recolonize the Glacier Park National Park area
(Ream & Mattson 1982). One pair even raised pups amongst a herd
of cattle on the prairies of the Rockies' eastern front (Diamond
1994). Montana now supports an estimated 70 wolves, and additional
animals from Canada are entering Idaho and Washington state (Mech
et al. 1995a).
Europe has seen the same trend. In Italy the wolf population
responded to the protection resulting from the research and educational
efforts of Boitani (1986) and increased to 300 individuals that
inhabit even areas around the outskirts of Rome. In Spain wolf
numbers reached 1500-2000 (Blanco et al. 1990), and in Poland,
about 850 (Bobek et al. 1993). Overflow to develop in Finland
(Pulliainen 1993) and eventually a nascent population developed
that straddles Norway and Sweden, currently numbering 20-25 (Promberger
et al. 1993a). Wolves also are spreading from northern Italy into
France and from Poland into eastern Germany (Promberger et al.
1993b).
The much improved public attitude toward wolves, coupled with
publicity and law enforcement have allowed the burgeoning wolf
populations to use areas that had not been wolf habitat for decades,
thus demonstrating the wolf's inherent adaptability. The wolf's
new range includes areas of higher road density (Fuller et al.
1992) and much more open, accessible, and populated areas. Breeding
packs now live less than 90 km from Minneapolis and St. Paul,
Minnesota. One wolf was radio-tracked out of the forests in which
it had been raised and into farm fields within 30 km of St. Paul's
center (Wydeven 1994). The animal roamed the farmlands for several
weeks before returning to forest. Other wolves making their way
south of Minneapolis and St. Paul are being killed by cars or
shot when mistaken for coyotes (Canis latrans). Wolves dispersing
into North and South Dakota have been crossing great expanses
of open areas (Licht & Fritts 1994).
In Spain wolves live like coyotes in wheat and sunflower fields
in regions with human densities of up to 200 people per km2 (Vila
et al. 1993). The animals scavenge garbage and livestock remains
and hunt smaller mammals. In Canada, Alaska, Scandinavia, the
Mideast, and much of Asia, wolf numbers are stable or increasing
(Ginsberg & Macdonald 1990).
Given protection, wolves can expand their range rapidly (Fuller
et al. 1992). Average litter sizes reach five to six (Mech 1970).
The territorial packs produce young each year, and maturing individuals
disperse (Fritts and Mech 1981, Gese and Mech 1991) distances
that may exceed 800 km straightline (Fritts 1983). They search
out mates and begin new packs (Rothman and Mech 1979) in new areas
(Ream et al. 1991).
As wolves dispersed from wildernesses, they successfully contended
with more highways, traffic, residences, habitat fragmentation,
and other human disturbances (Mech et al. unpublished data). Some
probably were unable to adapt, especially the first waves. Nevertheless,
wolves that did settle semi-wilderness areas probably became more
habituated to the increased disturbances, and as a population
then adapted more to increasing disturbance.
In Italy, Spain, and Portugal, where much of the wolf's food
is comprised of garbage, wolves have long inhabited the wooded
mountains during the day and made their way into rural villages
to scavenge at night (Zimen & Boitani 1979). In North America,
ungulate population densities are high close to population centers.
Thus, wolves have plentiful natural prey when they move to new,
nonwilderness areas.
As wolves show up in new regions they gather new constituencies
that support their recovery. In Europe the European Wolf Network
dedicated to the recovery of the wolf in central Europe (Promberger
& Schroder 1993) became a branch of the IUCN Wolf Specialist Group
in 1992. Other organizations have formed in North America that
call for the reintroduction of wolves into such places as Arizona,
Colorado, northern New York, and New England.
As wolves move into agricultural areas, conflicts with humans
greatly increase. For example, when Minnesota wolves increased
from 1988 through 1993 by an estimated 15 percent, the number
of wolves killed by the U.S. Department of Agriculture Animal
Damage Control Program increased from 59 to 139, or 223 percent
(Paul 1994). In Spain estimated damage by wolves now exceeds $1
million per year (Vila et al. 1993).
With these conflicts come a distinct danger of public backlash.
Not only will wolves in semi-agricultural areas take increasing
numbers of livestock and incur the wrath of the livestock industry,
which often has strong political support, but they also will kill
pets. In Minnesota wolves killing dogs has caused considerable
public animosity (Fritts & Paul 1989). As the media begin publicizing
such issues, the public gains an exaggerated impression of the
problem. A strong backlash of antiwolf sentiment could result
in management practices that would again restrict wolves to wilderness
areas. Poland has experienced three such cycles of wolf protection
and persecution (Okarma 1992). How can these problems be avoided
and the wolf be restored to as many places as possible? Until
some nonlethal method of controlling wolf populations is discovered,
it appears that lethal control will remain the ultimate means
of curbing wolf damage to livestock and pets.
Several non-lethal methods of preventing livestock losses to
wolves have been tried and abandoned. In Italy and other European
countries, for example, traditional husbandry techniques relied
on guard dogs and shepherds tending small flocks of livestock;
such techniques today are uneconomical. Use of guard dogs alone
has been tried against wolves in Minnesota with only limited success
(Fritts et al. 1992), although the method may be useful in specific
cases. Wolves have also been translocated to other areas, but
many either returned to where they were caught or became problems
elsewhere (Fritts et al. 1984, 1985). Aversive conditioning (Gustavson
& Nicolaus 1987) has not yet proven effective with wild wolves
(Fritts et al. 1992). Currently an electric fence in use in Sweden
seems to hold some promise for protecting livestock from wolves,
but it has not yet been subject to controlled testing (Eles 1986).
Furthermore, such fences tested for coyotes have generally been
expensive, hi gh-maintenance, and better suited for smaller areas
(Dorrance & Bourne 1980, Nass & Theade 1988).
Compensation for livestock losses is useful for minimizing public
animosity toward wolves, especially when wolf populations are
low and each wolf is important to the population. In Italy, compensation
was important in changing public attitudes toward acceptance of
wolves in agricultural areas. But as wolf populations proliferate,
compensation payments must also increase, sometimes disproportionately.
At some point compensation payments will become politically unpopular
as the public learns it is subsidizing wolves via payments to
farmers for their wolf-killed livestock. Thus many government
agencies are wary of even initiating such payments.
An innovative alternative to public payment for livestock killed
by wolves was instituted by the Defenders of Wildlife in the U.S.
This private, nonprofit organization established a fund to reimburse
ranchers in the western U.S. and even encouraged ranchers to allow
wolves to raise pups on their private land via a payment of $5000
per den (Fischer et al. 1994). The public may well begin demanding
that animal organizations assume these burdens from the government
as the costs increase. In any case, without wolf population control,
people would eventually object to the payments or the damages
caused by wolves.
With natural habitat in so many areas greatly fragmented and wolves
adapting to travel through relatively settled and open areas,
some disjunct wolf populations are developing where wolves can
live without causing livestock damages. For example, about 90
km northwest of Minneapolis and St. Paul, Minnesota, a pack has
lived and bred for at least two years on a wildlife management
area surrounded by agricultural land without killing local livestock.
Similar instances are known in Montana (Diamond 1994) and other
parts of Minnesota (Fritts & Mech 1981, Fritts et al. 1992). This
suggests that management zoning could allow wolves to inhabit
areas where they can feed on natural prey while they are kept
out of agricultural areas.
The approach is to designate zones of potential wolf habitat
and distinguish them from areas that should be kept wolf-free.
Zoning is common in regulating wildlife harvesting and has been
applied on a large scale in wolf recovery plans (U. S. Fish and
Wildlife Service 1975, 1987). If public attitudes continue to
lean toward protectionism, pressure may develop to apply zoning
on local levels such that small sanctuaries are maintained and
control is applied only outside these areas.
The scale of zoning is important. Wolves could be zoned out
of entire states or zoned into only large national parks or nature
preserves. Or they could be allowed to inhabit any area they naturally
colonize as long as their sole prey is wild species. For example,
in a wildlife refuge of only 100 km2 surrounded by farmland including
livestock, wolves could be protected in the refuge but destroyed
immediately outside it. This is similar to the situation in Riding
Mountain National Park, Manitoba, which, although a much larger
area, is an island of wilderness in a sea of agricultural land
(Carbyn 1982).
The main advantage of large-scale zoning is simplification and
efficiency of management. Any wolf in a designated no-wolf state
or outside any large wolf refuge would be subject to legal taking,
while those inside would be protected or managed through regulated
taking. This scenario could allow wolf populations to remain in
the Lake Superior states and much of the mountainous regions of
the western U.S., depending on how large the zones are.
The main disadvantage of large-scale zoning is the need to protect
livestock that would inevitably live inside some of the larger
zones. In Minnesota this would perpetuate the current situation
in which close to 150 wolves are killed by government controllers
annually for about $1225 each. A second disadvantage is that wolves
would probably not be allowed in many areas where they really
could live. This might mean banishing wolves two packs have been
living without causing livestock depredations. Furthermore, in
most of Europe where there are few if any large, remote regions
left, large-scale zoning would be very difficult.
With small-scale zoning, the main disadvantage for management
agencies is complexity. At one extreme even single wolf packs
in areas without livestock would be protected, while immediately
outside wolves could be taken. This could present difficult law
enforcement problems, although such problems are not unlike those
that currently exist for other species in wildlife refuges, national
parks and other protected areas. A small-scale zoning proposal
in Italy (Boitani & Fabbri 1983) was opposed by wolf protectionists
because of the difficulty of law enforcement and the feeling that
wolves would be relegated to areas too small to maintain viable
populations.
Such a fine-grained approach would probably require management
agencies to identify possible wolf areas so that when colonized
they would be recognized as wolf sanctuaries. Geographic information
systems would greatly simplify this task. Furthermore, identification
of such sanctuaries could be incorporated into ecosystem management
plans, biodiversity initiatives, and similar strategies as they
are developed for other reasons.
The main advantage of small-scale zoning would be to allow wolves
to live in enclaves throughout much of Europe and the United States
similar to the way they currently inhabit Wisconsin and Michigan
(Hammill 1993; Wydeven et al. 1994). For several reasons, this
approach would not require the very large-scale land and habitat
protection visualized by the Wildlands Project (Mann and Plummer
1993). Although dispersing wolves would be subject to persecution
while passing through nonprotected areas, those moving primarily
at night or outside of hunting seasons would stand a reasonable
chance of survival. With enough small enclaves of wolves, there
should be large numbers of such dispersers to colonize new areas,
resupply reduced populations, provide sufficient outbreeding,
and thus comprise regional metapopulations. Furthermore, inbreeding
depression, while a problem among some captive wolves (Laikre
& Ryman 1991), probably is not in most wild populations because
of the high natural tu rnover and ensuing selection. Deleterious
alleles should get cleansed from the population quickly.
The Isle Royale wolf population is instructive. Isle Royale
is a 538-km2 national park in Lake Superior some 25 km from Ontario.
It was colonized by wolves about 1949 (Mech 1966), probably by
only two unrelated wolves (Rothman & Mech 1979). Genetic testing
after 40 years indicated a single female founder (Wayne et al.
1991). Nevertheless, the population stabilized at about 23 for
a long period and increased to 50 in 1980, the highest wolf density
on record (Peterson & Page 1988). Although the population then
crashed, raising concerns about inbreeding depression and disease
(Peterson & Krumenaker 1989), the wolves survive. In 1994, eight
1993 offspring survived (Peterson 1994). Thus, with just two founders
and 50 percent loss of genetic variability (Wayne et al. 1991),
this population has survived for 45 years. Had it been on the
mainland, chances are good that some outbreeding would have occurred.
Biologically, wolves could inhabit parts of almost all regions
of the U. S. and many of the European countries. Since protection,
they have been recorded in nine and possibly ten U. S. states.
If biology were the only relevant factor, however, wolves would
never have had to be declared endangered. Throughout the wolf's
former range, it has been persecuted because of its tendency to
prey on livestock and pets. Even though it is currently on the
endangered species list in the U.S., control has been applied
in Minnesota, Wisconsin, and Montana. Thus there is every reason
to believe that wolf control will parallel wolf recovery wherever
it takes place (Mech 1979, Fritts 1993).
The inevitability of wolf control, however, introduces a new,
complex element into the equation governing the wolf's future
in all but the remotest areas of the world: wolf protectionism.
The same cultural attitudes that fostered wolf recovery also encouraged
an extreme degree of wolf protectionism. Those of us professionally
involved with wolf recovery have traditionally been maligned by
antiwolf people (Haubner 1990). Now we are vilified by many wolf
lovers as wolf enemies because of our acknowledgement that wolves
often require control.
Wolves are revered for several reasons. Because they tend to
kill prey that are old, sick or weak (Murie 1944; Mech 1970),
many laypeople mistakenly believe that, without wolves, prey would
automatically die out from disease. Wolves are also hailed as
good models for the human race because of their alleged monogamy
and family allegiances. A book was even titled The Soul of the
Wolf (Fox 1980). Other misconceptions about wolves encourage wolf
protectionism. Because of the book Never Cry Wolf by Farley Mowat
(1963) and the popular movie made from the book, many people believe
wolves live primarily on mice rather than ungulates. Both are
fiction (Banfield 1964, Pimlott 1966), but both purport to be
true and are sold and shown by museums and other unsuspecting
educational organizations. Other misconceptions, half truths,
and outdated views that many protectionists hold include the following:
wolves only prey on livestock when no natural prey is available;
the loss of pack members fost ers disastrous social chaos in the
wolf population; wolves socially limit their own population; because
the wolf is on the U. S. endangered species list, this means that
there are very few left anywhere in the world; and wolves are
so shy of humans that they will move out of areas of high activity
or avoid settling in them, and they will maintain dens and pups
only many kilometers from such activity.
Because of these misconceptions and the power of animal rights
groups, wolf control is resisted by much of the public (cf. Garrott
et al. 1993). This attitude has three major negative implications
for wolf recovery. First, some people revere wolves so much that,
rather than having wolves face control, these people would rather
not restore wolves to areas where they would have to be controlled.
Because wolves will probably have to be controlled almost everywhere
they are restored, this sentiment translates into political pressure
against wolf recovery. Second, the antiwolf public, such as some
livestock owners and organizations, intensify their antiwolf attitudes
in reaction to the extremism of the other side. They also fear
the possibility of road closures and other restrictions on land
use that are often fostered by protectionists using the wolf to
prevent logging, mining, snowmobiling or other human uses of semiwilderness
and wilderness. Third, some wolf advocates resort to ter rorism
(Hayes, personal communication) and deceptive advertisements (Anonymous
1992). This zealotry intimidates public officials, who might otherwise
be predisposed toward wolf recovery, to shun it.
Of course, the prowolf contingent holds a wide spectrum of attitudes.
Thus, some people will accept control against livestock depredations
but oppose control prescribed for increasing game herds. Some
will accept control by government agencies but not by the public.
Many people will accept indirect methods of control such as fencing,
guard dogs, or aversive conditioning. These indirect methods are
more acceptable because they do not involve humans killing wolves
directly. Few proponents of these methods seem to realize, however,
that keeping wolves from prey ultimately reduces the carrying
capacity of wolf range, and thus fosters starvation and increased
deaths from intraspecific strife (Mech 1994). This is particularly
true in countries such as Italy, Spain, Israel, where a high percentage
of the total carrying capacity for wolves is comprised of livestock,
but it applies on a smaller scale to North America as well. As
long as wolf deaths are either indirect (and thus not so o bvious)
or natural, many people accept these deaths who would not tolerate
direct or human-caused deaths.
Direct lethal control is still usually the only practical course
under most conditions. There are several ways to apply this control.
Control by government agency, usually the Department of Agriculture
in the U. S., is the type generally most acceptable to wolf advocates,
but it is by far the most expensive and time-consuming. Control
by landowners or their agents is the one most favored by landowners,
but it is difficult to police, and most landowners lack the time
and expertise for it, except by poisoning. Open taking of wolves
year-around in no-wolf zones similar to the taking of coyotes
in most areas of the U. S., and regulated taking by the public,
could be applied in no-wolf zones or in wolf sanctuaries to hold
the population down such as is done in many suburban areas for
white-tailed deer (Odocoileus virginianus), geese (Anser sp.),
and beavers. A modification of this type of control is public
taking by special permit.
All of the non-government approaches to control are much less
expensive but also less precise to the area or to specific wolves
taken and generally are the most disliked by wolf advocates. A
notable exception is the government control of wolves to increase
herds of big game in areas of Alaska and Canada. A public take
of 1200-1500 wolves per year in Alaska brings little or no protest,
but the state's controlling of 150 wolves to increase big game
herds is protested vehemently (Anonymous 1993). While biologically
this seems illogical, politically such state control allows animal-rights
groups to portray this control as a dastardly government program
that must be stopped.
The wolf's high reproductive potential and its tendency to disperse
hundreds of kilometers insure that there are few places where
wolves could be restored without some form of control being necessary.
But the very people most enthusiastically promoting wolf recovery
are generally those who want no control, so this dilemma makes
public officials reluctant to promote recovery.
Because wolf-taking by landowners or the public is the least
expensive and most acceptable by people who do not regard the
wolf as special, there will be greater local acceptance for wolf
recovery in more areas where such control is allowed. Thus, if
wolf advocates could accept effective control, wolves could live
in far more places.
It appears that the best way to promote wolf recovery is to encourage
public education about wolf management issues so that a significant
proportion of the public would support wolf recovery while tolerating
some form of control. Public education programs must include the
message that any restoration of wolves will ultimately result
in a need to control wolves (Fritts et al. 1994). Of course, there
will always be animal-rights advocates who never will accept any
wolf control. If their views are seen by most of the public as
counterproductive to wolf recovery, however, officials can probably
be persuaded to allow wolves to live in far more of their former
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