The Wolves of Denali

L. David Mech, Layne G. Adams, Thomas J. Meier,
John W. Burch, and Bruce W. Dale

Cover Photo by Leo Keeler, 1998

Chapter 1

Nuuk, Reykjavik, Nome, Barrow, Murmansk--all are frozen outposts toward the northern end of the earth that evoke images of snowbound, windswept winters and fleeting, urgent summers. Much farther south on the earth rise huge mountain peaks like K2, Annapurna, and Everest. All higher than 20,000 feet (6,000 meters), these massifs of ice, snow, and rock are so huge they make their own frigid weather.

The effects of both northern latitude and sheer elevation combine around central Alaska's own massif, Mount McKinley (plate 2). There it can snow any time of the year, yet perpetual summer daylight promotes a flush of flowers, as well as a myriad of mosquitoes. At 64' N latitude and 150' W longitude, in the shadow of Mount McKinley ("the Mountain"), lies a land where wildlife and their habitat persist in as natural a state as any found on earth. Throughout this great wilderness live the wolves of Denali National Park and Preserve, a 6.1-million-acre (24,000-square-kilometer) area southwest of Fairbanks and north of Anchorage, where human influence is minimal.

Only by realizing the extreme geophysical setting, the combination of latitude and topography, can one begin to understand the wolves and other wildlife of Denali. Latitude determines the sun's angle of incidence and the pattern of daylight hours. These basic physical factors in turn help set the weather and climate, the plant communities, and the wildlife that depend on them.

The widely divergent patterns of daylight in the Denali area seem alien and unfamiliar. A Fairbanks, Alaska, teacher once asked a student to all in the blanks for "The sun rises in the---(what direction?) and sets in the---(what direction?)." The student, not about to fall for a trick question, responded "summer or winter?" That far north, the sun rises in the northeast in summer, hangs high all day, and briefly dips below the horizon in the northwest at "night." In the dead of winter, however, the sun creeps above the horizon from the south for only a few hours in midday.

In autumn, the days rapidly shorten, and the sun hits the earth at a lower and lower angle. It heats the ground and atmosphere only a little, even during the day (figure 1.1). At night, much heat is lost through radiation and conduction. Moisture in soils freezes so deep that in many areas it never thaws, forming permafrost.

As winter tightens its grip, lakes and rivers freeze. In distant coastal areas, the ocean acts like a giant hot-water bottle, its thermal mass buffering winter heat losses. Moisture-laden air, blowing from oceanic areas, crosses into the colder interior, dropping its water load as snow. Ultimately, the Arctic Ocean, Chukchi Sea, and the northern reaches of the Bering Sea are capped with ice, removing this source of moisture for interior Alaska. With little moisture now available to the north and west, clear winter skies, frigid temperatures, and shortening days ̃nish the job, locking the Denali region into the cold and dark of winter.

"The Mountain"
This combination of high latitude and continental climate is not unique to interior Alaska but reaches across vast expanses of Canada and Siberia as well. Setting the Denali region apart, however, is the Mount McKinley massif with a base of over 1,000 square miles (2,500 square kilometers) of granite and ice projecting abruptly up to 20,320 feet (6,100 meters) above sea level from a piedmont plateau at 2,000 feet (600 meters) above sea level. Many peaks in the park range from 5,000 to 7,000 feet (1,500Đ2,100 meters) high and are characterized by alpine tundra vegetation, rock and scree slopes, and small headwater glaciers. However, the peaks of Mount McKinley and its sister Mount Foraker (17,400 feet, or 5,220 meters) soar thousands of feet higher. Composites of rock, snow, and ice, they are devoid of soil and vegetation, and of any life save the few hardy humans who clamber up their sides in valiant, death-defying attempts to meet some personal challenge. The McKinley massif was born of the collision of gargantuan tectonic plates. Gigantic glaciers gradually cascaded from the high peaks and eventually shaped the surrounding region. Movement of the glaciers crushed underlying rock into a ̃ne silt loam, or loess, that, transported by wind and water, filled in large basins creating the Tanana Flats far to the north. Receding glaciers left U-shaped valleys, and eons of meltwater runoff carved jagged canyons through hills and rocks. Few large lakes lie in the study area, although thousands of ponds dot the landscape.

Our study area lies primarily north of Mount McKinley. Its only road is the gravel "park road" that begins near park headquarters at the east end of the park. It curves and stretches some 90 miles (144 kilometers) to Wonder Lake and Kantishna due north of the Mountain in the center of the park.

Denali Weather
The huge mountains of the Alaska Range, including Mount McKinley, set the stage for an array of life more diverse, abundant, and visible than found on the ßats of the interior just to the north or on the snow-buried slopes of the south side. It is often said that The Mountain makes its own weather, and that is true. Southerly and westerly flows of moist air masses from the Pacific Ocean and Bering Sea predominate much of the year. These warm fronts dump large quantities of snow on the south side due to adiabatic cooling, where moist ocean air masses cool as they lift over the steep terrain. On the north side and in the mountain passes, high winds, due to the gradients between low-pressure systems in the Gulf of Alaska and high pressure in the Arctic, scour snow from the ridges and hills, leaving relatively shallow snowpacks and exposed vegetation. These breaks in the snow provide wintering habitat for caribou, Dall sheep, and other wildlife.

Nevertheless, the 80-inch (200-centimeter) average annual snowfall leaves a knee- to waist-deep snowpack in areas not wind-scoured that persists from late September to early May (table 1.1). Although 80 inches is a great deal of snow and strongly influences the area, it amounts to only 15 inches (38 centimeters) of water. Denali's rugged mountains and foothills can receive dustings of snow in any month. By October, even the lowlands are blanketed with snow and ice, which remains into May (figure 1.2). These conditions result in 8 months of insignĩcant photosynthesis; 8 months of snowpack; icy, overflowing rivers; and frozen ground. Temperatures range from 90 deg. F (32 deg. C) in summer to -53 deg. F (-47 deg. C) in winter, highly dependent on elevation.

Not only are conditions harsh, but like the Toklat-colored grizzly bear, they are unpredictable. In mid-September 1992, 2 to 4 feet (60-120 cm) of heavy, wet snow fell in the Tanana and upper Kuskokwim regions. The early snowstorm caused hardwoods, still leafed, to bend under the weight of the wet snow, in some cases until the tops touched the ground. Cold weather then froze the trees, and in many places they remain forever in that condition. All this followed a late spring; consequently the summer of 1992 was the shortest on record, with ramifications we will discuss later. These variations on the main theme of very long winters in Denali play a pivotal role in the functioning of the ecosystem. One of the most significant aspects of the variations involves the total annual snowfall. During our study, that ̃gure varied fourfold (table 1.1), a natural "experiment" that yielded great dividends to our investigations.

Soils and Vegetation Soils in Denali, where present, are poorly drained because of the silt and permafrost. Standing water often becomes acidic due to tannic acids from slowly decaying organic materials. Plant communities on the flats of the northwest edge of the park and preserve consist of black spruce (Picea) and tamarack (Larix) bogs and wet tussock tundra with sedge (Carex), moss (Sphagnum), and heath (Ericaceae) understories. Cotton grass (Eriophorum) is a common component of wet tussock tundra and is seasonally important to caribou.

More-productive high ground and riparian areas hold white spruce (Picea), birch (Betula), aspen (Populus), and alder (Alnus) with understories of berries and willow (Salix) (figure 1.3). Foothills below roughly 3,500 feet (1,050 meters) support woodlands of white spruce, aspen, and birch. Willows and alder at the tree line give way to alpine tundra and bare ground at higher elevations. The alpine tundra is dominated by Dryas mats and prostrate willows. Lichens, particularly Cetraria and Cladonia, are locally abundant in many vegetative communities, especially woodlands and alpine tundra. Rivers in Denali are heavily laden with glacial silt. These rivers rise during the heat of the day as snowfields and glaciers melt under the summer sun. Clear-water streams and tundra ponds host a variety of aquatic and riparian plants important to moose and other wildlife. In winter, many water bodies freeze to the bottom. Rivers freeze to the substrate, and ßowing groundwater seeks new routes adjacent and atop the ice, creating treacherous overflow and patches of "aufeis" that persist well into summer. The action of ice and water erosion along river courses creates a continually renewing ecotone of pioneering willows and other colonizing plants, which are important forage for moose, caribou, and numerous small animals.

Wildfires caused by lightning strikes are common in the northwest flats of the park, and in a dry year, thousands of acres will burn (plate 3). Willows, forbs, birches, and aspen frequently recolonize after fire, creating mosaics of productive patches of ungulate forage. Wildlife Populations Denali was made a national park largely because of its wildlife resources. The park hosts herds of Dall sheep, caribou, and moose, along with their primary predators, wolves and grizzly bears, as well as wolverines, foxes, coyotes, black bears, many smaller mammals, and a wide variety of raptors, waterfowl, ptarmigan, and other birds (Sheldon 1930).

Dall Sheep
Dall sheep, the species that has figured so prominently in the history of the park, are stout, medium-size ungulates adapted to utilizing steep and rocky terrain (plate 4). In Denali, they are the smallest ungulate available as wolf prey. Adult rams sport large horns that curl in a tight spiral, while ewes carry smaller sickle-shaped horns. Rams weigh around 200 pounds (90 kilograms) but can exceed 250 pounds (110 kilograms). Adult ewes are much smaller, weighing only about 120 pounds (55 kilograms) on average (Bunnell and Olsen 1976; Nichols 1978).

Dall sheep are currently common throughout the Alaska Range at densities similar to those found in Denali. We will never know whether sheep were more abundant in Denali than adjacent areas when the park was designated because so few records exist from outside the park.

Denali's sheep were hunted by native Alaskans, big-game hunters, miners, and market hunters (Walker 1992). Concerns for overharvest of this species helped lead to the creation of the park. Sheep populations have ßuctuated markedly in Denali, with highs in the late 1920s of at least 5,000, and perhaps many more (Murie 1944). Their numbers declined severely to 1,000 to 1,500 after the big-snow winters of the early 1930s. Sheep numbered about 2,000 in the park just before our study (Singer et al. 1981; Taylor et al. 1987). Dall sheep habitat consists of scree and alpine-tundra-dominated mountain slopes. The animals are rarely found far from steep, rocky slopes, which provide refuge from their predators, primarily wolves, grizzly bears, golden eagles, coyotes, and wolverines (Murie 1944). Sheep venture away from escape terrain in search of forbs, grasses, and willows, but if caught there, are easily killed.

In winter, sheep frequent windblown slopes and ridges where favored forages like fescue grasses are more accessible. In some parts of Alaska, sheep winter range overlaps summer range, while elsewhere seasonal migrations of 6 to 12 miles (10Đ20 kilometers) have been documented (Ayres 1986). In Denali, sheep sometimes move between the taller Alaska Range peaks and the lower "Outer Range" ridges to the north. However, they are present in both areas regardless of the season.

Denali's sheep breed in November, and ewes give birth to a single lamb in mid-May (Geist 1971).

Caribou
Caribou numbers have also fluctuated dramatically in Denali, as indicated earlier. The staple meat animal for the region (Rawson 1994), caribou were much more numerous before 1945, likely exceeding 20,000 animals (Murie 1935; Singer 1986). During the late 1940s to the early 1960s, herd estimates ranged from 8,000 to 10,000 caribou (Adams et al. 1989). In the 1960s, several caribou herds throughout Alaska and Canada declined dramatically. The Denali herd was no exception, reaching a low of around 1,000 caribou by 1975 (Troyer 1977, 1978). Since the mid-1970s, the Denali herd has fluctuated between 1,000 and 3,200 animals.

It is important to note that early caribou estimates were confounded by the limitations on the methods available. Many of these estimates were made during seasonal migrations or were based on gross numbers seen by travelers. The highly variable distribution of caribou made it extremely difficult to even determine to which herd a group of caribou may have belonged. With the increasing use of aerial surveys and radiotelemetry in the past couple of decades, the accuracy of herd-size estimates has greatly improved. About 30 caribou herds inhabit Alaska, ranging in size from hundreds to hundreds of thousands of animals. They total about a million (Davis and Valkenburg 1991). Most are of the "Alaskan" ecotype (Davis and Valkenburg 1991). This is an arbitrary class of caribou herds that ̃t behaviorally between the large migratory barren-ground herds of the Arctic and the dispersed low-density woodland populations of the boreal forest.

Alaskan ecotype caribou commonly inhabit mountainous terrain, migrate en masse seasonally (but on a lesser scale than barren-ground herds), and share ranges with grizzly bears and wolves, as well as with other ungulate prey. Although they commonly frequent spruce forests, these caribou usually live above the tree line and on the open tundra. Denali caribou, as well as other Alaskan ecotype herds, are larger than their barren-ground counterparts, and adult bulls carry large ornate antlers in fall (plate 5). Mature bulls average 490 pounds (220 kilograms) but can weigh up to 590 pounds (270 kilograms) (L. Adams, unpublished data). Cows are about half as big, weighing 255 pounds (115 kilograms) on average, and range from 185 pounds (85 kilograms) to 325 pounds (145 kilograms) (L. Adams, unpublished data).

Denali's caribou undertake migrations of about 50 miles (80 kilometers) between summer and winter ranges, rather short compared to the hundreds of miles covered seasonally by barren-ground herds. Their movements cover a wide elevation range from below 2,000 feet (610 meters) in winter to above 6,000 feet (1,830 meters) in summer in search of the lichens, shrubs (particularly willows), graminoids, forbs, and mushrooms on which they subsist (Boertje 1984).

In interior Alaska, caribou breed in late September and early October while migrating from mountain summer ranges to tundra and woodland plains for the winter. Then, in late April and early May, caribou cows return to their calving grounds as the calving season begins.

As we will show in chapter 7, caribou calving is highly synchronous and takes place in mid-May. Most females bear their single calves high in the mountains near glaciers and snow̃elds, concentrated in a relatively small area. Others will calve scattered in the mountains or spruce woodlands. Although quite precocious, walking and feeding a few hours after birth, caribou calves are extremely vulnerable to predators during their first 2 weeks of life.

Cows try to avoid detection by calving in areas hard for predators to search, such as mottled snowfields high in the mountains. If the cow detects a predator, she tries to move away undetected. If detected, the cow will not usually try to defend the calf against a wolf or bear but, rather, will make a run for it. Young calves that are small and slow quickly fall behind and are easily caught.

Later, caribou frequent areas near high mountain snow̃elds and glaciers to escape the mosquitoes, warble flies, and botflies of June. On a calm, warm day in late June, nearly the entire herd can be found huddled and still on snow̃elds and glaciers, seeking relief. Finally, the weather becomes cool and rainy in late July and early August, and the caribou move about freely and feed voraciously on the lush new vegetation.

Moose
Moose (plate 6) occur at moderate to low densities typical of the interior, approximately 6 moose per 10 square miles (2 per 10 square kilometers) (Meier et al. 1991). They were considered plentiful in Denali before about 1920, when market hunters greatly reduced them (Murie 1944). By the late 1930s, however, they had regained "satisfactory" numbers (Murie 1944), which then remained relatively stable through the next several decades (Haber 1977) but may have declined in the late 1970s and early 1980s. Since then, parkwide estimates of 1,400 to 1,900 moose have been made using various types of aerial censuses (Meier et al. 1991).

Moose are the largest wolf prey in Denali, weighing two or three times as much as caribou. Alaskan cow moose average about 880 pounds (400 kilograms), whereas adult bulls average about 120 pounds (55 kilograms) heavier (Franzmann et al. 1978). However, bulls greater than 1,200 pounds (545 kilograms) have been reported (Franzmann et al. 1978; Gasaway and Coady 1974).

Moose, an important food for wolves, are widely distributed in Denali and are locally abundant wherever willow are common (Miquelle et al. 1992). These animals eat a large variety of willows, especially Salix alexensis, S. planifolia, S. arbusculoides, and S. glauca (Risenhoover 1989). Moose also feed on birch and aspen seedlings and blowdowns, and occasionally on the bark of aspens. Although similar in appearance to the preferred shrubs, the abundant stands of dwarf birch, resin birch, and alder are avoided by moose. In summer, moose feed also on aquatic vegetation.

Moose in interior Alaska, including Denali, are an overlapping mixture of migrating and locally resident animals (Gasaway et al. 1983; Ballard et al. 1987). Some travel to foothills for winter and to swamps and lowlands for summer, while others remain in one area year-round, typically in foothills at the tree line, riparian areas, and swamps. Moose are well adapted to snow and cold; however, snow depths greater than about 20 inches (53 centimeters) decrease their foraging ef̃ciency and increase energetic costs of movement, resulting in greater vulnerability to predators (Stephenson 1995). Aside from population surveys (Meier 1987; Dalle-Molle 1987; Meier et al. 1991), only a small segment of resident moose have been studied in Denali (Miquelle et al. 1992).

Smaller Prey
Other prey for wolves include beavers (figure 1.4), which are common wherever stands of deciduous trees or shrubs border streams, rivers, deep ponds, and lakes; and hares, which can be abundant during population highs. Marmots, ground squirrels, numerous microtine rodents, and other small mammals and birds are also occasional food items (Murie 1944). Packs whose territories include salmon rivers feed heavily on salmon at times, especially in autumn and early winter.

Bears and Other Competitors
Grizzly bears (plate 7) in upland habitats and black bears in woodlands compete with wolves for prey in Denali (Murie 1944). Bears of both species in interior Alaska do not have as much access to protein-rich salmon as coastal bears and are much smaller. Adult grizzlies average about 550 pounds (115 kilograms) and 450 pounds (205 kilograms) for females and males, respectively, in spring (J. Keay, unpublished data); black bears average about 40% lighter (J. Hechtel, unpublished data). Both grizzlies and black bears emerge from dens around May and search for winter kills, newborn ungulate calves, and accessible vegetation. They breed during summer. By August, berries, and in some Denali rivers, salmon, become important food resources. Both bear species occasionally prey upon adult ungulates (Dean et al. 1986; Boertje et al. 1988; L. Adams, unpublished). By autumn, they have gained about 30% over their spring weights.

In October, Denali bears enter dens, where from one to three cubs are born in February. Grizzly bear cubs stay with their mother until they are 2 to 3 years of age (Reynolds and Hechtel 1983), while black bear cubs are generally weaned at 1 to 2 years of age (Hechtel 1991). Grizzlies in Denali den in holes typically located on snow-covered hillsides at 3,000 to 4,000 feet (900Đ1,200 meters) (Darling 1987). Black bears den in all types of habitats in holes, brush piles, or simply under a blanket of snow (Smith 1994).

Bears have been censused only in portions of Denali (Dean 1987; J. Keay, unpublished), but density data for those parts suggest a minimum population of 300 grizzlies and conceivably as many as 450, plus an unknown number of black bears.

Wolves and bears tend to avoid each other, but altercations between the two species are not unusual, especially at kills and around dens. Murie (1944) described several wolf-bear incidents in Denali, and our observations tend to parallel his.

During one interaction, the East Fork wolves (see chapter 3) attacked a sow grizzly and 3 yearlings. On July 23, 1989, at about 8:40 a.m., 12 members of the East Fork pack were paralleling the park road, when they headed up a willow drainage in which the 4 bears were rummaging around. Neither the bears nor the wolves seemed aware of each other until the wolves came to within 50 feet (15 meters) of the bears.

The 3 yearlings immediately rushed up the drainage, with the sow right behind. As the wolves caught up, the sow turned to fight them off. The yearlings split, however, and the sow could not defend them all. Two of the yearlings continued on. Five or 6 wolves harassed the sow, while the rest chased the yearlings. The wolves quickly caught up and killed them both. The remaining wolves then rushed over to join in the feast. In this case, the attack was one of sheer predation rather than the usual skirmish between competitors. Also competing with wolves to various degrees are wolverines, lynx, coyotes, red foxes, and golden eagles. None of these compete with wolves to anywhere near the extent that bears do, however.
Most often wolves interact with their competitors around kill remains because kills are magnets for all the meat-eaters. Wolves usually chase away their smaller competitors, sometimes catching and killing them (Mech 1970). The same is true when they encounter a competitor by chance.

Tom Meier recalled an encounter one of our wolf packs had with a wolverine that impressively demonstrates how tough this particular competitor is: In January 1987, pilot Jim Cline and I were radio-tracking the East Fork pack when we spied seven wolves running up a creek bed near the Teklanika River. The wolves overtook and attacked a fleeing wolverine, forming a ring around the animal, lifting it off the ground and shaking it. Making a low pass, we saw that the wolverine was on its back with one wolf continuing the attack. On the next pass, some of the wolves were rolling on the ground, and the others were resting. Several ravens had also arrived. However, we could not find the wolverine.

For 10 minutes we searched. The seven wolves eventually arose and moved on up the creek. Finally, after another 20 minutes, we spotted the wolverine running rapidly down the creek the way it and the wolves had come. The creature appeared unhurt, and no blood was visible at the attack site.

I visited the scene on the ground the next day. Approaching on the wolves' exit trail, I saw drops of blood in their tracks. At the attack site were a few small tufts of wolverine hair and a few drops of blood. It appeared that the wolverine had escaped under a shelf of ice until the wolves left. I saw no blood in the wolverine's exit trail, and I believe it escaped unharmed.

Wolves
Wolves are not unusually abundant in Denali, but the population is both vigorous and viable. The animals have been legally protected in the original park area since 1952 (Harbo and Dean 1983). However, only fragmentary information about their numbers is available from 1952 through 1965. From 1966 through 1974, Gordon Haber (1977) estimated that five wolf packs lived primarily in the old (pre-1980) park, although he was able to define the approximate territories of only two of them. His maximum winter estimates for each of the five packs from 1966 through 1974 total 76 wolves living primarily in the 3,294 square miles (8,432 square kilometers) of the old park, or about 9.0 wolves per 1,000 square kilometers.

Our estimates of late-winter wolf density varied from 3.1 wolves per 1,000 square kilometers in 1987 to 7.8 wolves per 1,000 square kilometers in 1991 (figure 1.5). Roughly a third of the park and preserve is high-altitude rock and ice, not suitable habitat for wolves or their prey, so these numbers translate to an estimated 50 to 125 wolves over the 6,312 square miles (16,160 square kilometers) of available wolf habitat in the park and preserve. In early winter, after pup production and before much dispersal, wolf density is at its highest. Our highest estimate of wolf density at that time was 9.8 wolves per 1,000 square kilometers, or a parkwide estimate of 158 wolves.

Between 1988 and 1993, pups made up 35% to 44% of all wolves observed in fall counts. Combined with parkwide estimates of fall wolf numbers and estimated over-summer mortality of 9% (see chapter 8), these figures yield annual estimates of 65 to 79 pups produced in the park.

Humans in Denali
Before European contact, native Alaskans hunted and trapped throughout the region, and wolves were a part of their harvest. Wolves were respected as hunters and competitors, and were an integral part of the native spiritual culture. In the Denali region, wolves were probably taken primarily by deadfall traps, although pit traps, torsion traps, and puncturing shards frozen in bait may also have been used (Scarff 1972; Rawson 1994). The hides of wolves made warm clothing that enabled survival in the cold winters. The Athabascan Indians that hunted in the Denali region lived primarily along the major waterways north of the park where salmon were available (Brown 1991). They made hunting trips to the foothills during spring through autumn. Some clans may have spent entire summers in the foothills, pursuing caribou and sheep and gathering berries. With European contact came firearms, steel traps, and a market for fur. The hides of wolves and other furbearers became important components of the local economy. Explorations of the Denali area by Alfred Brooks, James Wickersham, and Frederick Cook in 1902 and 1903 resulted in writings about the abundant game herds. These in turn attracted big-game hunters and museum collectors like Charles Sheldon and Belmore Browne.

At the same time, growing gold camps along the Yukon and Tanana drainages created a great demand for game meat. Market hunters frequented the north slope of the Alaska Range, including Denali, to provide caribou, moose, and sheep for mining camps in Fairbanks, Kantishna, and elsewhere (Loftus 1967; Rawson 1994).

Park Establishment and the First Wolf Controversy
Charles Sheldon's concerns over the future of Dall Sheep and other game led him to promote the region as a national park. Through the clout of the Boone and Crockett Club and the Camp FireClub of America, Mount McKinley was designated a national park in 1917. Mount McKinley Park, it turns out, quickly spawned the world's first wolf controversy, a dispute that lasted more than three decades and split the ranks of park personnel, sportsmen, and even scientists.

The controversy began in California. Zoologists Joseph Grinnell and Tracy Storer (1916) published one of the first pleas for protecting predators in general. Grinnell's students spread the philosophy, and in 1924, the American Society of Mammalogists held a symposium on mammalian predators. Charles Adams, one of Grinnell's students, then declared that national parks should be "our main sanctuaries for predaceous animals" (Adams 1925, 90). The New York Zoological Society and the Boone and Crockett Club passed resolutions in 1929 supporting predator protection in national parks (Rawson 1994). Sanctuaries were needed because by this time official government predator control was in full swing, and the wolf was a main target. Bounties and special predator controllers with the U.S. Biological Survey wreaked havoc in an attempt 'to eradicate the wolves and coyotes of Alaska' and elsewhere (Rawson 1994, 92).

On the other side of the issue, Grinnell spoke to a conference of National Park Service (NPS) superintendents about the value of predators. His audience was sympathetic, and NPS director Horace Albright (1931, 185Đ86) proclaimed that "predatory animals are to be considered an integral part of the wildlife protected within national parks, and no widespread campaigns of destruction are to be countenanced." This was just after the government had exterminated the last wolves in Yellowstone.

The plot thickened, however, when prominent members of the Camp Fire Club of America began a campaign to promote extermination of wolves in Mount McKinley Park. Because the Camp Fire Club had been instrumental in establishing the park, the clubƠs inßuence was strong. Its view was that wolves would exterminate the Dall sheep and spill out to all other areas of Alaska.

Murie's Studies and Wolf Protection
Thus began a seesawing of official attitude and action in Denali. From 1930 to 1934, rangers killed 23 wolves; in 1935, acting park director Arno Cammerer forbade rangers from killing wolves; the next year, assistant director Arthur Demaray approved killing wolves "for food habits studies," and rangers shot 14 from 1936 to 1938 (Rawson 1994). Damned if they killed wolves, and damned if they did not, the NPS ended up recommending a scientific study. The scientist was to be Adolph Murie.

Murie's charge was to determine whether the park's sheep faced extermination by the wolves and whether wolf control was necessary. He spent 15 months in the ̃eld in 1940 and 1941 and learned much new information, as discussed earlier (figure 1.6). He concluded that while wolves were the primary factor limiting the sheep, the sheep and wolves were currently in proper balance. Furthermore, management should be directed at maintaining that balance, and wolf control should be applied only when absolutely necessary (Murie 1944).

Murie's book was widely acclaimed as objective and highly informative. However, it was roundly criticized by the anti-wolf forces, who, while becoming increasingly marginalized, still held great power in Alaska. Murie was sent back to the park in August 1945 for a month and found that the sheep population had declined from about 2,700 in 1941 to about 500. He recommended killing 10 to 15 wolves in sheep range and continuing wolf control until the sheep increased.

The Camp Fire Club decided to seek legislation to force wolf control, and in late 1945 a bill was introduced into the U.S. House of Representatives. The NPS, however, successfully resisted the legislation.

Nevertheless, NPS rangers did begin controlling wolves. From 1946 through 1948, some 18 wolves were taken (Rawson 1994). Murie agreed that wolf control should continue until the sheep population reached 2,500, and he personally began snaring wolves. Eleven more wolves were killed from 1949 to 1952. By 1953, sheep numbers had reached 1,500. With the sheep herd increasing, the older members of the Camp Fire Club dying off, and wolves being controlled by federal efforts outside the park, the anti-wolf pressure on the NPS began dissipating. At the same time, a signal request was made of the park that evolved into a historic decision.

Walt Disney Studios wished to produce a "True Life Adventure" film about wolves in Denali (Rawson 1994). To do so, undisturbed wolves would be needed, so the studio requested a temporary cessation of wolf control. NPS director Conrad Wirth agreed, and Disney produced the film White Wilderness. It portrayed the wolf sympathetically to a wide audience. Public protest against wolf protection was by that time minimal, so the NPS never rescinded their ruling. As a result, wolf protection in Denali has continued uninterrupted since 1952 (Harbo and Dean 1983), and the NPS enjoys considerable public support for that stand.

Wolf-Taking Restrictions
Even today, however, protection of Denali's wolves is not without controversy. Today's laws, while protecting wolves in the original 2.1 million acres (8,400 square kilometers) of the park, provide for a limited harvest of wolves in the park expansion and preserve around the park (figure 1.7).

In 1980, the federal Alaska National Interest Lands Conservation Act (ANILCA) greatly expanded existing national parks and refuges and created expansive new parks and refuges as well. Mount McKinley National Park, renamed Denali National Park and Preserve, swelled from about 3,294 to 9,468 square miles (8,432 to 24,238 square kilometers). However, ANILCA provided for the customary and traditional use of wildlife on federal lands created by the act, even in national parks. The act recognized subsistence lifestyles and the importance of wildlife and other natural resources to the livelihood of local residents. Thus, ANILCA allows the taking of wolves and other wildlife in the 1980 additions to the park by qualified rural residents (figure 1.7). Rural residents qualify if they reside in rural villages that historically used the park lands or if they can demonstrate that they or their families traditionally used the areas. The regulations are liberal and nearly identical in seasons, methods, and means of taking to areas outside the new park expansions. In addition to rural subsistence, sport hunting and trapping are permitted in the preserve (figure 1.7).

Hunting of wolves is allowed from mid- August through April. During this time an individual may take up to 10 wolves. Trapping is legal from November through March, and there is no limit. While these seasons may seem extreme for a national park, they result in very little wolf harvest (about 2% of the park and preserve wolf population annually) (see chapter 2). This is because few residents qualify and actually reside where they have access to the new park. For the same reasons, the harvest of other wildlife in the park is extremely limited. The Denali Caribou Herd is closed to hunting under both state and federal regulations. The taking of wolves and other animals remains prohibited in the original "wilderness" portion of the park.

The Denali Ecosystem
For Denali National Park and Preserve, truly the whole is more valuable than the sum of its parts. Plant and animal populations, virtually all trophic levels, occur in natural and pristine assemblages. Human influences are minimal and have been for decades. Although some small areas have been altered by humans, they seem trivial when compared to the magnificent scale of the entire area.

Denali encompasses a broad ecotonal belt ranging from high and severe mountains of rock and ice, through the mountain and river valley complexes, to the spruce woodlands. This assemblage of habitats supports a diverse array of wildlife communities. Wildfire, a naturally occurring disturbance important for maintaining productive and diverse habitats, has been allowed to run its course since the early 1980s. Another, often overlooked factor distinguishing the Denali ecosystem from other natural areas is that Denali is not an island. Rather, it is part of continuous habitats and populations throughout Alaska and into Canada. Moose, wolves, sheep--virtually all plants and animals--are members of larger, continuous populations that are also relatively undisturbed. This means that population ßuctuations, gene flow, and persistence of populations are not affected by the insular qualities of most natural areas caused by the massive alteration of adjacent habitats and wildlife communities. The presence of metapopulations sharing ecosystems external to the park provides an important added benefit: comparisons can be drawn between systems to help further understanding of the influence of humans on wildlife resources. For all these reasons-species richness, protected nature, surrounding buffer areas, and the long-term integrity of the ecosystem-Denali National Park and Preserve was an ideal location for a long-term study of wolves and their prey.