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Wolves, Bears, and their Prey in Alaska: Biological and Social Challenges in Wildlife Management 3 Alaska's People, Biomes, and Wildlife Species of Concern INTRODUCTION To evaluate a science-based program, one must understand the contexts in which the program is carried out, including the ecological, economic, and political contexts. This chapter provides an overview of the physical and biological environments in which wolves, bears, and their prey interact. The two following chapters discuss what is known about predator-prey interactions in general (chapter 4) and what has been learned from different attempts to alter predator-prey interactions through wolf and/or bear reductions (chapter 5). The socioeconomic environment is covered in chapter 6. An overview of the climate, vegetation, and soils of Alaska and the major biomes into which Alaska has been divided serves 2 purposes. First, by illustrating the great diversity of Alaskan environments, it demonstrates why management programs must be based on area-specific information. Second, it shows how the vastness of Alaska and the limitations of personnel and financial resources available for biological research make it inevitable that management decisions are based on less-complete information than is desirable and than would be possible if resources were less limited or the area smaller. This survey is followed by a review of the ecology and natural history of wolves, bears, and their primary prey—moose and caribou. The review demonstrates the substantial differences among the species of concern and provides a basis for designing management programs that are tailored to the life-history traits of the managed species and how they respond to changes in their environment.
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Wolves, Bears, and their Prey in Alaska: Biological and Social Challenges in Wildlife Management THE PEOPLE OF ALASKA The first humans in the Western Hemisphere are believed to have come from Asia across the Beringian land bridge into Alaska 12,000–15,000 years ago. The first to arrive were the Paleoindians, who spread throughout North America and South America and from whom most native American cultures derived, including the Haida and Tlingit Indians of the southeastern coast of Alaska (Greenberg 1987). Later migrations of people are believed responsible for the Athabascan Indian cultures that are present throughout the interior and south-central regions of Alaska and in parts of northwestern Canada. The marine-oriented Eskimos of Arctic, western, and southwestern Alaska (represented today by the Inupiat, Yup'ik, and Koniak cultures) arrived much later, apparently by boat across Bering Strait. The Aleut culture of the Aleutian Islands and adjacent Alaska Peninsula has its closest affinity to early Eskimo cultures. Today, the human population of Alaska is about 610,000, with the majority concentrated in and around Anchorage, Fairbanks, Juneau, and smaller southcoastal cities of a few thousand each. The Alaskan population is younger than the rest of the United States (median age 30 years versus 33.4 years for the whole United States), and its rate of population increase in recent years is second only to that of Nevada (Alaska Bureau of Vital Statistics 1995). The nonindigenous residents of Alaska (those who are not Alaska Natives of Eskimo, Indian, or Aleut descent as defined by the Alaska Native Claims Settlement Act of 1971) make up about 84% of the Alaskan population, and about 80% of them live in urban communities. The non-Native residents of Alaska are primarily first-or second-generation immigrants from the other states and reflect the racial and ethnic diversity that characterizes the United States. There are some differences in the racial make-up between Alaska and the United States as a whole. Alaska's population is 4.1% black (12.1% for the entire United States), 16% Native American (0.8%), and 3.2% Hispanic of any race (9%), according to the 1990 US census. Alaska Natives currently make up 16.5% of the state's population and most live in rural communities (Wolfe 1996). There are about 225 rural communities of fewer than 500 residents scattered throughout the state but concentrated in southeastern Alaska. The residents of all but a few of those communities are predominantly Alaska Natives. Human activities have had less effect on the ecosystems of Alaska than elsewhere in the United States. Conversion of land to agricultural use has been minimal, as is the extent of land alteration through mining and petroleum development. The greatest alteration of ecosystems has been through extensive logging of forests in southeastern Alaska. More than 40% of Alaska is managed by federal agencies through the National Park Service, Fish and Wildlife Service, and Bureau of Land Management.
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Wolves, Bears, and their Prey in Alaska: Biological and Social Challenges in Wildlife Management BIOMES: CLIMATE, VEGETATION, SOILS, PERMAFROST Alaska is one-fifth the size of the lower 48 states and occupies 1,477,270 km2. It extends more than 20° in latitude from Pt. Barrow to Amatiguak Island in the Aleutians; it spans 42° in longitude from Portland Canal in southeastern Alaska to Attu Island in the western Aleutians. The topography, climate, and ecosystems of Alaska are characterized by great diversity (Selkregg 1976; Klein and others 1997). Alaska's coastline extends more than 54,700 km, bordering on the North Pacific, Bering Sea, and Arctic Ocean. The average length of the frost-free period varies from 40 days in the Arctic to more than 200 days in parts of southeastern Alaska. The Alaskan interior, because of its relatively warm, very long summer days, has relatively high plant productivity, except where permafrost is present. Permafrost is ground that remains perennially frozen except for a shallow summer-active layer. Annual precipitation ranges from less than 25 cm in the Arctic to 500 cm in parts of the Alexander Archipelago of southeastern Alaska. The low precipitation of interior Alaska would result in desert conditions at lower latitudes, but most of the winter precipitation remains on the land as snow until spring, and low summer evaporation rates and drainage (impeded by permafrost) retain soil moisture throughout the summer in most areas. Permafrost is present throughout most of the Arctic and northwestern Alaska except beneath lakes, rivers, and adjacent riparian zones. South of the Brooks Range in the interior and in southwestern Alaska, permafrost is discontinuous and confined mostly to lowlands, north-facing slopes, and higher elevations. The position of Alaska between the cold Arctic Ocean and the relatively warm North Pacific, its extensive coastline and southern islands, and its high mountain ranges and associated ice fields with intervening and extensive lowlands are responsible for the ecological diversity that characterizes the state. Alaska can be divided into 6 major biogeographic regions or biomes (figure 3.1). Coastal Temperate Rain Forest and Coast Range Mountains The coastal temperate rain forest grows under the moderating maritime influence of relatively warm ocean currents of the North Pacific and Gulf of Alaska. It is a continuation of the temperate coniferous rain forest that extends from northern California along the northwestern coast to northern Kodiak Island. The distribution of large mammals in this region, because of its many islands and the relatively short time since it was heavily glaciated, is more complex than that in the remainder of Alaska. Sitka black-tailed deer (Odocoileus hemionus sitkensis) are common on the islands and adjacent mainland of southeastern Alaska and have been successfully introduced to the islands of Prince William Sound and to Kodiak Island. Moose (Alces alces) occupy the major river valleys that penetrate the Coast Range Mountains—such as the Stikine, Taku, and Chilkat rivers—but do not normally occur in the coniferous forests of the islands. Mountain goats
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Wolves, Bears, and their Prey in Alaska: Biological and Social Challenges in Wildlife Management FIGURE 3.1 Major biogeographic regions of Alaska. (Oreamnos americanus) occur in scattered localities in the mainland mountains and have been introduced to Baranof and Revilla islands. Brown, or grizzly, bears (Ursus arctos) are present on the mainland; Admiralty, Baranof, and Chichagof islands of southeastern Alaska; the islands of Prince William Sound; and Kodiak and Afognak islands. Black bears (Ursus americanus) and wolves (Canis lupus) are present throughout the mainland and on the islands of southeastern Alaska south of Frederick Sound. Densities of bears are high on the islands and mainland, especially in the vicinity of salmon spawning streams. Wolves vary in density in relation to the availability of their primary prey, blacktailed deer. Deer densities fluctuate with the frequency and severity of winters of deep snow. Deer habitat has been greatly modified by forest harvesting. Interior Boreal Forest Much of interior Alaska, which is sheltered by high mountains from the moist maritime air to the south and the cold Arctic air to the north, has a continental climate. Winters are cold and long; summers are warm and short. Seasonal changes are rapid. Altitude strongly influences plant growth, the presence and composition of forests, and the extent of permafrost. Fire, caused mostly by lightning, is a natural feature of the ecology of the interior boreal forest. The pattern of vegetation is a complex juxtaposition of plant communities that vary with fire history, soil temperatures, drainage, and exposure. The present boreal forest in interior Alaska is part of the northern boreal forest that extends from the
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Wolves, Bears, and their Prey in Alaska: Biological and Social Challenges in Wildlife Management Atlantic coast of Canada across the northern portion of the continent into Alaska. Mammals characteristic of this major biome are similar throughout this vast area. The extensive interior boreal forest region is broken by several mountain complexes that support typical mountain species, such as caribou (Rangifer tarandus) and brown bear. The lowland areas of mixed forest, shrub zones, and wetlands support moose and black bears. Moose also venture into the mountains, especially in summer. Wolves are present throughout the region, varying in density with the distribution and availability of prey, primarily moose and caribou. Mammal populations in this largely intact ecosystem undergo natural fluctuations, primarily in relation to variations in winter snow depth, plant succession after wildfire, and variation in rates of predation by wolves and bears. Interior Mountains (Montane Habitats) This biogeographic region is a complex of mountain ranges characterized by extreme physiographic variability. Elevation, slope steepness, and exposure vary widely locally and between major mountain masses. The distribution of vegetation, although dominated primarily by alpine forms, reflects the terrain variability, and vegetation at lower elevations includes elements from the boreal forest. Climatic variability is pronounced. Because oceanic air masses lose much of their moisture on the seaward sides of mountain ranges as rain or snow, interior sides are characterized by more arid conditions. The progression of summer plant growth is also highly variable, a condition that favors mammals—such as mountain sheep (Ovis dalli), moose, and caribou—that are able to move extensively over the variable terrain to forage. The depth of winter snows usually limits the availability of winter habitat for mountain sheep, moose, and caribou in these mountain areas. Brown bears and wolves are common, especially in areas of high prey densities. Maritime Tundra (Southwestern Alaska and Bering Sea Islands) The maritime tundra that dominates southwestern Alaska and the Bering Sea islands is the product of the cool climate generated by the cold Bering Sea waters. However, there is a gradation from the more-humid and milder conditions prevailing in Bristol Bay and the coastal Alaska peninsula bordering the Aleutian region to the Seward Peninsula, where the adjacent seas are ice-covered for 8 months of the year. Coastal wetlands are extensive throughout the region and dominate the broad expanse of the delta of the Yukon and Kuskokwim rivers (Selkregg 1976). In its climate and vegetation, the region is transitional between the Aleutian and Arctic biogeographic regions. Moose, caribou, brown bears, and wolves have been absent or rare throughout much of this region in the past. In recent years, caribou re-entered the area from the expanding Mulchatna Herd, a re-established herd in the Kilbuck Mountains,
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Wolves, Bears, and their Prey in Alaska: Biological and Social Challenges in Wildlife Management and segments of the Western Arctic Herd, which winters on the western Seward Peninsula and in Kotzebue and Norton Sound drainages. About 30,000 reindeer (domesticated caribou introduced from Scandinavia) in several herds privately owned by Alaska Natives are grazed on the Seward Peninsula and have come into increasing conflict with wintering caribou from the Western Arctic Herd. Moose have become established in recent decades in riparian areas on the Seward Peninsula. Their populations appear to have peaked, with some possible declines, perhaps because the sparse winter habitats along the major rivers have become heavily browsed. Introduced muskoxen live on Nunivak Island, on Nelson Island on the Yukon Delta, and on the Seward Peninsula. Their expanding populations are hunted under permit systems by both subsistence and sport hunters. Brown bears are common in coastal areas of the Alaska Peninsula and present at lower densities on the Seward Peninsula. Aleutian Region The Aleutian Islands and adjacent Alaska Peninsula, an interface between the North Pacific and the Bering Sea, includes the southernmost land area in Alaska. The Aleutians, which extend nearly 1,900 km from the Alaska Peninsula to Attu Island, are renowned for their cool, foggy, and windy weather and temperature consistency. The mean daily temperature of 3.9°C has an annual range of only 9.4°C. Native large terrestrial mammals are absent from virtually all the Aleutian Islands. In the Aleutian biogeographic region, only the southern portion of the Alaska Peninsula and closely adjacent Unimak Island support caribou (currently at a low density), brown bears, and wolves. The bears are moderately abundant and depend heavily on the productive salmon streams in the area. Wolf numbers are low, presumably because of the low prey density. Arctic Tundra The Brooks Range mountains separate the boreal forest and the Arctic tundra biogeographic regions. The Arctic tundra of Alaska, on the higher and drier ground at its southern limit, descends to the broad and wet coastal plain that continues to the Arctic Ocean. Coastal plain tundra is interspersed with thousands of shallow lakes. The Arctic tundra experiences strong northeasterly winds in winter that are generated by the Arctic high-pressure system over the frozen Arctic Ocean. The little snow that falls throughout the long winter is redistributed by winds into drifts wherever there is variation in the terrain. In spite of climatic extremes of the Arctic tundra—which results in a short plant growth season, low mean annual temperature, and cold soils underlain by permafrost—this region is extremely productive of life, much of which (such as nesting birds and caribou) migrates out of the region during winter. The Arctic
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Wolves, Bears, and their Prey in Alaska: Biological and Social Challenges in Wildlife Management tundra supports Alaska's largest caribou herds, the Western Arctic Herd (about 500,000) and Porcupine Herd (about 170,000), as well as the smaller Central Arctic and Teshekpuk herds, each numbering around 20,000. In the latter half of the 20th century, moose moved into riparian habitats along the Collville, Sagavanirktok, Canning, and other larger rivers of the Arctic. Barren-ground brown bears occur at low densities, being more numerous in the southern foothills and western Arctic where they are important predators on young caribou during calving. Wolf densities are relatively low in the Arctic, largely because of the low density of ungulate prey in winter. Their numbers are highest in the northern foothills and adjacent Brooks Range mountains, where mountain sheep, moose, and some wintering caribou can be present. Muskoxen have been reestablished in the Arctic through introductions in the 1960s and 1970s in the eastern and western areas. They are increasing and dispersing into unoccupied habitats. The total numbers in the Arctic tundra appear to be approaching 1,000. ECOLOGY OF LARGE MAMMALS IN NORTHERN ECOSYSTEMS The population dynamics of large mammals in southern Canada and the lower 48 states have been dramatically altered over the past two or three centuries. Not only have most of the natural habitats been converted to agricultural lands, managed forests, or high density human use areas, but dominant species such as wolves, grizzly bears, and millions of migratory bison have largely been eliminated. The population dynamics of large mammals in the pre-Columbian system are unknown. What remains of the system can be managed carefully to provide predictable harvests for hunters and trappers. In northern Canada and Alaska, however, natural habitats have not been substantially altered. Alaskan ecosystems are still much the same as they were when Europeans first arrived in North America and caribou, moose, wolves, and bears are not threatened with extirpation. The large mammal system remains largely intact and is highly volatile. Most caribou migrate over large, and often unpredictable areas, and their numbers fluctuate enormously. For example, in Alaska, the Mulchatna herd increased from about 30,000 to 110,000 individuals between 1984 and 1993. In Quebec and Labrador, the George River herd increased from about 10,000 in the 1950s to approximately 800,000 in the 1990s. Such enormous shifts in one species influence the dynamics of the entire system, and large changes in moose abundance and distribution have also been evident. Biologists have debated how and when factors such as the very slow growth of terrestrial lichens, the highly variable Arctic weather, and predation at different caribou densities, influence caribou and moose populations. What isn't debated is that the northern systems are much more dynamic than what is left of the southern systems. Consequently, we cannot expect to manage northern systems to provide predictable numbers of ungulates and predictable harvests; Alaska is
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Wolves, Bears, and their Prey in Alaska: Biological and Social Challenges in Wildlife Management not Colorado, and caribou are not elk. Alaskans should not expect as constant a supply of game as people expect in the southern portion of the continent. Many factors influence the population dynamics of moose, caribou, wolves, and bears. Among the most important are quantity and quality of food, weather, diseases, parasites, predation, intraspecific strife, and human harvest. These factors can act individually, but often they act in concert. For example, animals weakened by poor nutrition are more likely to succumb to disease or predators than well-fed animals are. Together, these factors determine the ecological carrying capacity of the environment—that is, the number of animals that can be supported on a long term basis by the resources of the environment (see box). Another definition of carrying capacity is the number that will persist if disease organisms and predators are present. Because the goal of predator control and management in Alaska is to increase prey numbers, we use the former definition of carrying capacity in the report. Economic carrying capacity, the density of animals that will allow maximum sustained harvest, is always lower than ecological carrying capacity (Caughley 1976 in Krebs 1994). Carrying capacities fluctuate because abiotic and biotic environmental factors change. Assessing environmental carrying capacity is important because success or failure of management programs often depends on the environmental conditions under which they are carried out. Ecological carrying capacities of Alaskan environments for ungulates are low because arctic, alpine, and subalpine soils are typically poor in nutrients. In combination with short growing seasons, this limits the potential quality and production of forage. Shallow, high-latitude soils underlain by permafrost are fragile and easily damaged and vegetation may recover slowly from overgrazing. Arctic and sub-Arctic plants grow rapidly during the short growing season, and growth rates are higher during warm, sunny summers than during cloudy, cold summers (Klein 1970). Plant species are adapted to different soils, terrain, exposure, and water availability (Maessen and others 1983) and are distributed in a mosaic throughout Alaska (Maessen and others 1983). This mosaic distribution influences the foraging patterns of large herbivores and omnivorous bears. Large herbivores, in turn, can strongly affect plant succession, species composition, and productivity. High densities of large herbivores, such as moose and caribou, can cause both short-term declines in aboveground plant biomass and long-term declines in the quality of plants (Leader-Williams and others 1981). Lower biomass or poorer-quality plants (for example, plants that have been severely browsed) reduce the carrying capacity of the habitat for ungulates (Klein 1968). Plants respond to being fed on by moose or caribou in a variety of ways. Light grazing can stimulate growth, and plants that grow beyond the reach of browsers can complete their annual growth more rapidly (Edenius 1993; Edenius and others 1993; Molvar and others 1993; Robbins and others 1987). Browsing and grazing can also stimulate chemical changes that alter plant palatability (Edenius 1993; Bryant and Kuropat 1980). Thus, plants and herbivores are
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Wolves, Bears, and their Prey in Alaska: Biological and Social Challenges in Wildlife Management Carrying Capacity Ecological carrying capacity is the number of organisms that the resources of the environment can sustain in a particular region (Pianka 1978, Sharkey 1970). Populations that exceed their carrying capacity must ultimately decline. Populations below their carrying capacity will tend to increase toward it. Carrying capacity itself can vary from year to year because the availability of resources for a population varies from year to year. Because carrying capacities are themselves dynamic, they are difficult to estimate (Dhondt 1988, Pulliam and Haddad 1994). The notion of carrying capacity is crucial in studying predator-prey systems because the effect of predator removal depends largely on how much below its ecological carrying capacity a prey population is. If prey are far below their carrying capacity and if predators are the cause, predator removal can be effective in enhancing the abundance of prey. In contrast, regardless of how many prey are killed by predators, if prey populations are close to their ecological carrying capacity, removal of predators will not produce a marked increase in prey abundance. involved in a coevolutionary process of defense against and response to predation (for example, Coley and others 1985; Edenius 1993; Rosenthal and Janzen 1979). Fire is a natural part of northern ecosystems and has a major impact on plant community organization and structure, and, hence, carrying capacity for ungulates (Viereck 1973; Zackrisson 1977). After a fire, short-lived and fast-growing deciduous shrubs and trees colonize the area, and long-lived and slow-growing species decline (Scotter 1967). Lichens recover from fire much more slowly than shrubs, but, over a long time period, fire rejuvenates lichen communities and helps to create the mosaic of habitats that is characteristic of Arctic and sub-Arctic environments (Miller 1980; Zackrisson 1977). Fires might have increased with settlement and have been blamed for the decline of some caribou herds (Bergerud 1974; Miller 1980). However, after about 200–300 years without a fire, lichens become senescent and grow more slowly (Klein 1982). Because abundance and distribution of animal populations depends on the availability of suitable habitat, habitat management should be explored as a management tool, especially where studies of habitat and surrogate measures of habitat quality (responses of ungulates such as age at first reproduction, reproductive rates, and growth rates) indicate that habitat quality is depressing the growth of ungulate populations. If the data indicate that a population is limited by habitat (especially food), habitat management programs (such as burning or crushing of vegetation) might be effective. Habitat management is a long-term process and requires long-term commitment. It will not solve short-term ''emergency" situations, but it can improve food availability and it might buffer populations during catastrophic events, such as deep persistent snow. Finally, because habitat management is generally socially acceptable, it offers the potential to decrease or
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Wolves, Bears, and their Prey in Alaska: Biological and Social Challenges in Wildlife Management avoid the use of controversial methods (such as killing predators) to increase ungulate populations. Wolf Ecology Distribution and Density Wolves occur throughout the Northern Hemisphere wherever large ungulates occur, from about 20° N latitude to the polar ice pack. They are present in habitats ranging from deserts to tundra, and although they still occur as far south as Saudi Arabia and India, they are most common in more-northern areas of Alaska, Canada, and Russia. In Alaska, wolves are found over most of their historical range, occupying about 85% of the state's 1.5 x 106 km2 (figure 3.2; Stephenson and others 1995). Wolves are absent from areas that they did not colonize after the last glacial recession, including the Aleutian, Kodiak, Admiralty, Baranof, and Chichagof islands. Wolf densities vary geographically. In Alaska, wolf densities range from about 2 to 20/1,000 km2; total numbers were estimated at 5,900–7,200 during the winter of 1989–90 (Stephenson and others 1995). In the Arctic, densities of wolf populations are often less than 5/1,000 km2, but maximal midwinter wolf densities in southern populations often exceed 40/1,000 km2 (Fuller 1989a). On the basis of data from more than 20 intensive studies that measured total average ungulate biomass (often more than 1 ungulate species) and average wolf populations for a period of several years, variations in wolf density in all of North America seem to be strongly correlated with variations in ungulate biomass (Fuller 1989a, 1995; Messier 1985). The relationship between prey abundance and wolf numbers can vary in areas with migratory versus nonmigratory prey or where prey concentrate seasonally. However, all available data suggest that, unless artificially depressed by humans, wolf numbers are typically limited by ungulate numbers and availability. Food availability is the dominant natural factor that limits wolf abundance. Although the correlation between wolf density and prey abundance is high, the ratio of ungulate biomass to number of wolves is highest for heavily exploited wolf populations (Ballard and others 1987; Peterson and others 1984) or newly protected populations (for example, Fritts and Mech 1981; Wydeven and others 1995) and lowest for unexploited populations (Mech 1986; Oosenbrug and Carbyn 1982) or those where ungulates are heavily harvested (Kolenosky 1972). In Alaska, lightly harvested wolf populations occurred at much higher densities per unit of ungulate availability than heavily harvested populations (Gasaway and others 1992). Changes in wolf density in response to varying prey density have been documented by long-term studies in northeastern Minnesota (Mech 1977, 1986), Isle Royale (Peterson and Page 1988), and southwestern Quebec (Messier and
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Wolves, Bears, and their Prey in Alaska: Biological and Social Challenges in Wildlife Management FIGURE 3.2 Alaska wolf population densities as estimated by Alaska Department of Fish and Game. Management biologists with aerial winter surveys and by contacting area trappers and pilots. Research biologists have radio-collared wolves from several packs in 11 Alaska study areas to verify estimates of wolf densities. Crête 1985). Because the numerical response of an individual wolf population lags behind a change in prey density by up to several years, the ratio of ungulate biomass to wolves often differs among years and areas (Peterson and Page 1983). Wolf densities can also vary with prey type. Population densities per unit of prey biomass are lower in areas where wolves prey mainly on moose than where they prey mainly on deer. The reason appears to be that moose are, on the average, less vulnerable to wolf predation (harder to catch) than are deer. Vulnerability of individual prey species may depend on which other prey species also are found in the area. For example, caribou are more vulnerable to wolf predation when they co-exist with moose, but moose and sheep are less vulnerable when caribou are present (Bergurud 1974, Dale and others 1995, Seip 1992). In Alaska, wolf populations are estimated at different levels of precision, depending on management needs. The least-precise estimates are derived from a combination of information resulting from aerial surveys of wolf tracks, incidental observations, reports from the public, and sealing (mandatory registration)
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Wolves, Bears, and their Prey in Alaska: Biological and Social Challenges in Wildlife Management Habitat Quality and Ungulate Population Dynamics Habitat quality is an important determinant of dynamics of populations of large mammalian herbivores (Caughley 1981; Crête and others 1990; Jeffries and others 1994; Pastor 1988). Knowledge of seasonal and annual variability in forage quality and quantity is important for development of management plans for ungulate populations. In Alaska, habitat carrying capacities have been assessed primarily by such indirect indicators as the length of the lower jaw, marrow fat, and body weight of individual animals, or such demographic data as birth rate, calf survival, or population age structure. Those indirect measures, which assume that the status of the animals reflects the status of their habitats, have proved useful when time, expertise, or funding was not available for direct assessment of the habitats. However, those measures are often slow to respond to habitat changes, can be unduly influenced by short-term weather anomalies, and are influenced by factors other than habitat quality. A few long-term studies involving experimental manipulations and monitoring of habitat changes caused by these and natural disturbances, such as fire, have been undertaken. The collaborative research on moose-habitat relationships by ADFG and the Fish and Wildlife Service (FWS) in the Kenai National Wildlife Refuge (formerly the Kenai National Moose Range) is an excellent example. ADFG, with management responsibility for wildlife, and FWS, with responsibility for habitat, entered into a cooperative agreement that yielded basic information on responses of moose habitat to fire and other manipulations in the Kenai Peninsula and on the physiological, nutritional, and population consequences of habitat changes, constraints, and stocking levels (Oldemeyer and Regelin 1987; Regelin and others 1987; Schwartz and others 1988). Information gained from this long-term effort has been extremely valuable in the management of moose in Alaska and especially on the Kenai Peninsula. Intensive habitat-relationship studies like those carried out on the Kenai need not be duplicated in each area where ADFG plans to intensify management activities, but some level of knowledge of the existing habitat is essential if wise decisions are to be made. Desirable information includes description of vegetation, fire history, proportions of habitat in different successional stages after fire, rough biomass estimates of critical forage types (for example, lichens on caribou winter ranges and primary winter browse species in moose habitats), and distribution and depth of snow cover in the winter. REFERENCES Adams LG, BW Dale, and LD Mech. 1995. Wolf predation on caribou calves in Denali National Park, Alaska. In LN Carbyn, SH Fritts, and DR Seip, Eds. Ecology and Conservation of wolves in a changing world: proceedings of the second North American symposium on wolves. Canadian Circumpolar Inst. Univ. Alberta, Edomonton, 1995. Alaska Bureau of Vital Statistics. 1995. Population trends in Alaska.
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Wolves, Bears, and their Prey in Alaska: Biological and Social Challenges in Wildlife Management Alt GL. 1989. Reproductive biology of female black bears and early growth and development of cubs in Northeastern Pennsylvania. Ph.D. thesis, WV Univ, Morgantown. 116 Pp. Andreyev VN. 1977. Reindeer pastures in the subarctic territories of the USSR. Pp. 275–313 in W Krause, Ed. Handbook of vegetation science, part XIII. Application of vegetation science to grassland husbandry. Dr. W. Junk BV publishers, The Hague, Netherlands. Aune K and W Kasworm. 1989. Final report. East front grizzly studies. Mont Dep Wildl Parks, Helena. 332 Pp. Aune M Madel, and C Hunt. 1986. Rocky mountain front grizzly bear monitoring and investigation. Mont Dep Wildl Parks, Helena. 175 Pp. Ballard WB, JS Whitman, and CL Gardner. 1987. Ecology of an exploited wolf population in southcentral Alaska. Wildlife Monographs 98:1–54. Ballard WB and SL Miller. 1990. Effects of reducing brown bear density on moose calf survival in south-central Alaska. Alces 26:9–13. Ballard WB, SD Miller, and JS Whitman. 1990. Brown and black bear predation on moose in southcentral Alaska. Alces 26:1–8. Ballard WB, JS Whitman, and DJ Reed. 1991. Population dynamics of moose in southcentral Alaska. Wildlife Monographs 114:1–49. Ballard WB, CL Gardner, and DJ Reed. 1995. Use of line-intercept transects for estimating wolf densities. In LN Carbyn, SH Fritts, and DR Seip, Eds. Ecology and conservation of wolves in a changing world. Canadian Circumpolar Inst., Univ. Alberta, Edmonton. Banci V, DA Demarchi, and WR Archibald. 1994. Evaluation of the population status of grizzly bears in Canada. Int Conf Bear Res and Management 9:129–142. Batzli GO, RG White, and FL Bunnell. 1981. Herbivory: a strategy of tundra consumers. International Biological Programme 25:359–375. Becker E and others. In press. A population estimator based on network sampling of tracks in the snow. J Wildlife Management. Beecham JJ. 1983. Population characteristics of black bears in west central Idaho. J Wildlife Management 47:405–412. Berg WE and DW Kuehn. 1982. Ecology of wolves in northcentral Minnesota. In Pacquet, Eds. Wolves: a worldwide perspective of their behavior, ecology and conservation. Noyes Publications, Park Ridge, NJ. Bergerud AT. 1974. Decline of caribou in North America following settlement. J Wildlife Management 38:757–770. Bergerud AT. 1980. A review of the population dynamics of caribou and wild reindeer in North America. Proc Int Reindeer/Caribou Symp 2:556–581. Bergerud AT. 1983. The natural population control of caribou. In FL Bunnell, DS Eastman, and JM Peek, Eds. Symposium on natural regulation of wildlife populations. Proc No. 14. Forest Management and Range Exp Stn, Univ Idaho, Moscow. Bergerud AT. 1996. Evolving perspectives on caribou population dynamics, have we got it right yet? Rangifer Special Issue 9. Bjorge RR and JR Gunson. 1983. Wolf predation of cattle in the Simonette River Pastures in northwestern Alberta. In LN Carbyn, Ed. Wolves in Canada and Alaska. Canadian Wildlife Service Report Series 45. 135 Pp. Bjorge RR and JR Gunson. 1989. Wolf, Canis lupud, population characteristics and prey relationships near Simonette River, Alberta. Canadian Field-Naturalist 103:327–334 Blanchard BM and RR Knight. 1991. Movements of Yellowstone grizzly bear. Biol Cons 58:41–67. Boertje RD. 1984. Seasonal diets of the Denali caribou herd, Alaska. Arctic 37:161–165. Boertje RD. 1985. Seasonal activity of the Denali Caribou herd, Alaska. Rangifer 5:32–42.
OCR for page 73
Wolves, Bears, and their Prey in Alaska: Biological and Social Challenges in Wildlife Management Boertje RD, WC Gasaway, DV Grangaard, and DG Kellyhouse. 1988. Predation on moose and caribou by radio-collared grizzly bears in east central Alaska. Canadian J Zoology 66:2492–2492. Boertje RD and RO Stephenson. 1992. Effects of ungulate availability on wolf reproductive potential in Alaska. Canadian J Zoology 70:2441–2443. Boertje RD, DG Kellyhouse, and RD Hayes. 1995. Methods for reducing natural predation on moose: an evaluation. In LN Carbyn, SH Fritts, and DR Seip, Eds. Ecology and conservation of wolves in a changing world. Canadian Circumpolar Inst, Univ. Alberta, Edomonton. Boertje RD and CL Gardner. 1996. Factors limiting the Fortymile caribou herd. Alaska Dep Fish and Game. Fed Aid in Wildlife Restor. Prog Rep. Proj W-24-4. Study 3.38. Juneau. 79 Pp. Brand CJ, MJ Pybus, WB Ballard, and RO Peterson. 1995. Infectious and parasitic diseases of the gray wolf and their potential effects on wolf populations in North America. In LN Carbyn, SH Fritts, and DR Seip, Eds. Ecology and conservation of wolves in a changing world. Canadian Circumpolar Inst. Univ Alberta, Edmonton. Brown DE. 1985. The grizzly in the southwest; documentary of an extinction. Univ of Okla Press, Norman. 274 Pp. Bryant JP and PJ Kuropat. 1980. Selection of winter forage by subarctic browsing vertebrates: the role of plant chemistry. Annual Review of Ecology and Systematics 11:261–286. Burkholder BL. 1959. Movements and behavior of a wolf pack in Alaska. J Wildlife Management 23:1–11. Carney DW. 1985. 1985. Population dynamics and denning ecology of black bears in Shenandoah National Park, Virginia. M.S. thesis, VA Polytech Inst and State Univ, Blacksburg. 84 Pp. Caughley G. 1981. Overpopulation. In Problems in management of locally abundant wild mammals. R Jewell and S Holt, Eds. Academic Press, NY Charnov EL. 1982. The theory of sex allocation. Monographs in population biology. Vol. 18. Princeton, NJ, Princeton University Press, Clark. 1991. Ecology of two black bear (Ursus americanus) populations in the interior highlands of Arkansas. Ph.D. thesis, Univ Ark, Fayetteville. 228 Pp. Coley PD, JP Bryant, and FS Chapin III. 1985. Resource availability and plant anti-herbivore defense. Science 230:895–899. Crête M and H Jolicoeur. 1987. Impact of wolf and black bear removal on cow: calf ratio and moose density in southwestern Québec. Alces 23:61–87. Crête M and others. 1990. Food selection during early location by caribou calving in the tundra in Quebec. Arctic 43:60–65. Dale BW, LG Adams, and RT Bowyer. 1995. Winter wolf predation in a multiple ungulate prey system, Gates of the Arctic National Park, Alaska. In LN Carbyn, SH Fritts, and DR Seip, Eds. Ecology and conservation of wolves in a changing world. Canadian Circumpolar Inst. Univ Alberta, Edmonton. Dauphine TC, Jr. 1976. Biology of the Kaminuriak population of barren-ground caribou. Part 4. Growth, reproduction and energy reserves. Canadian Wildl Serv Rep. Ser No. 38. 69 Pp. Davis JL and P Valkenburg. 1991. A review of caribou population dynamics in Alaska emphasizing limiting factors, theory, and management implications. Pp. 184–209 in Proceedings of the Fourth North American Caribou Workshop. C Butler and SP Mahoney, Eds. Newfoundland and Labrador Wildlife Division, St. John's. Dhondt AA. 1988. Carrying capacity: a confusing concept. Acta Oecologia 9:337–346. Eberhardt LL, BM Blanchard, and RR Knight. 1994. Population trend of the Yellowstone grizzly bear as estimated from reproductive and survival rates. Canadian J Zoology. 72:360–363. Edenius L. 1993. Browsing by moose on scots pine in relation to plant resource availability. Ecology 74:2261–2269. Edenius L, K Danell, and R Bergstrom. 1993. Impact of herbivory and competition on compensatory growth in woody plants winter browsing by moose on Scots Pine. Oikos 66:286–292
OCR for page 74
Wolves, Bears, and their Prey in Alaska: Biological and Social Challenges in Wildlife Management Franzmann AW, CC Schwartz, and RO Peterson. 1980. Moose calf mortality in summer on the Kenai Peninsula. J Wildlife Management 44:764–768. Franzmann AW. 1981. Alces alces. Mammalian Species 154:1–7. Franzmann AW and CC Schwartz. 1986. Black bear predation on moose calves in highly productive versus marginal moose habitats on the Kenai Peninsula, Alaska. Alces 22:139–153. French SP and MG French. 1990. Predatory behavior of grizzly bears feeding on elk calves in Yellowstone National Park, 1986–88. Int Conf Bear Res and Management 8:335–341. Fritts SH and LD Mech. 1981. Dynamics, movements, and feeding ecology of a newly-protected wolf population in northwestern Minnesota. Wildlife Monographs 80. 79 Pp. Fritts SH. 1983. Record dispersal by a wolf from Minnesota. J Mammalogy 64:166–167. Fritts SH, WJ Paul, LD Mech, and DP Scott. 1992. Trends and management of wolf-livestock conflicts in Minnesota. US Department of Interior, US Fish and Wildlife Service Resource Publication 181. 27 Pp. Fritts SH and LN Carbyn. 1995. Population viability, nature reserves, and the outlook for gray wolf conservation in North America. Restoration Ecology 3:26–38. Fuller TK and WJ Snow. 1988. Estimating wolf densities from radiotelemetry data. Wildlife Soc Bull 16:367–370. Fuller TK. 1989a. Population dynamics of wolves in north-central Minnesota. Wildlife Monographs 105:1–41. Fuller TK. 1989b. Denning behavior of wolves in north-central Minnesota. American Midland Naturalist 121:184–188. Fuller TK. 1995. Guidelines for gray wolf management in the Northern Great Lakes Region. International Wolf Center Technical Pub 271, Ely, MN. 19 Pp. Gaare E and T Skogland. 1980. Lichen-reindeer interaction studies in a simple case model. In E Reimers, E Gaare, and S. Skjenneberg, Eds. Proceedings of the Second International Reindeer/Caribou Symposium, Direktoratet for vilt og ferskvannfisk, Trodheim, Norway. Garshelis DL. 1994. Density dependent population regulation of black bears. In M Taylor, Ed. Density dependent population regulation in black, grizzly and polar bears. Int Conf Bear Res and Management. Monogr Series No. 3. 43 Pp. Gasaway WC, RO Stephenson, JL Davis, PEK Shepherd, and OE Burris. 1983. Interrelationships of wolves, prey and man in interior Alaska. Wildlife Monographs 84:1–50. Gasaway WC, RD Boertje, DV Grangaard, DG Kellyhouse, RO Stephenson, and DG Larsen. 1992. The role of predation in limiting moose at low densities in Alaska and Yukon and implications for conservation. Wildlife Monographs 120:1–59. Gese EM, and LD Mech. 1991. Dispersal of wolves (Canis lupus) in northeastern Minnesota, 1969–1989. Canadian J Zoology 69:2946–2955. Greenberg JH. 1987. Language in the Americas. Stanford University Press, Stanford, CA. Gunther KA and RA Renkin. 1990. Grizzly bear predation on elk calves and other fauna of Yellowstone National Park. Int Conf Bear Res and Management 8:329–334. Haber GC. 1996. Biological impacts on wolves of exploitation and control. P. 36 in First Annual Conference of the Wildlife Society, Albuquerque, NM (abstract). Haber GC. 1996. Biological, conservation, and ethical implications of exploiting and controlling wolves. Conservation Biology 10:1068–1081. Hamer D and S Herrero. 1991. Elk, Cervus elaphus, calves as food for grizzly bears, Ursus arctos, in Banff National Park, Alberta, Canada. Canadian Field-Naturalist 105:101–103. Hamilton AN. 1988. Classification of coastal grizzly bear habitat for forestry interpretations and the role of food in habitat use by coastal grizzly bears. M.Sc. thesis, University of British Columbia. Harrington FH, LD Mech, and SH Fritts. 1993. Pack size and wolf pup survival: their relationship under varying ecological conditions. Behavioral Ecology and Sociobiology 13:19–26.
OCR for page 75
Wolves, Bears, and their Prey in Alaska: Biological and Social Challenges in Wildlife Management Hart JS, O Heroux, WH Cottle, and CA Mills. 1961. The influence of climate on metabolic and thermal responses of infant caribou. Canadian J Zoology 39:845–856. Hayes RD, A Baer, and DL Larsen. 1991. Population dynamics and prey relationships of an exploited and recovering wolf population in the southern Yukon. Yukon Fish and Wildlife Br Final Rep TR-91-1. Whitehorse. 67 Pp. Hayes RD. 1995. Numerical and functional response of wolves, and regulation of moose in the Yukon. M.Sc. thesis, Simon Fraser University, Burnaby, British Columbia. Heard DC and GW Calef. 1986. Population dynamics of the Kaminuriak northwest territories Canada caribou herd 1968–1985. Rangifer. Special Issue 1:159–166. Heard DC, TM Williams, and DA Melton. 1996. The relationship between food intake and predation risk in migratory caribou and implications to caribou and wolf population dynamics. Rangifer. Special Issue 9:37–44. Helle T, J Aspi, K Lempa, and E Taskinen. 1992. Strategies to avoid biting flies by reindeer: field experiments with silhouette traps. Annales Zoologici Fennici 29:69–74. Hicks MV. 1994. Federal aid in wildlife restoration survey-inventory management report. Alaska Department of Fish and Game, Div Wildlife Cons, Juneau, AK. Holleman DF and JR Luick. 1977. Lichen species preference by reindeer. Canadian J Zoology 55:14–26. Hovey FW and BN McLellan. 1996. Estimating population growth of grizzly bears from the Flathead River drainage using computer simulations of reproduction and survival rates. Can J Zool 74:1409–1416. Huntington S and Rearden 1993. Shadows on the Koyukuk. Alaska Northwest Books. Anchorage, AK Jeffries R and others. 1994. Vertebrate herbivores and northern plant communities: reciprocal influences and responses. Oikos 71:193–206. Jonkel CJ and IM Cowan. 1971. The black bear in the spruce-fir forest. Wildlife Monographs 27:1–57. Keith LB. 1974. Some features of population dynamics in mammals. Trans Int Congr Game Bio 11:17–58. Keith LB. 1983. Population dynamics of wolves. Pp. 66–77 in LN Carbyn, Ed. Wolves in Canada and Alaska, Canadian Wildlife Service Report Series 45. 135 Pp. Kingsley MCS, JA Nagy, and HV Reynolds. 1988. Growth in length and weight of northern brown bears—Differences between sexes and populations. Canadian J Zoology 66:981–986. Klein DR. 1968. The introduction, increase and crash of reindeer on St. Matthew Island. J Wildlife Management 32:350–367. Klein DR. 1970. Interactions of Rangifer-Tarandus reindeer and caribou with its habitat in Alaska. Riistatteteellisia Julkaisuja. 289–293. Klein DR. 1982. Fire, lichens, and caribou. J Range Management 35:390–395. Klein DR. 1992. Comparative ecological and behavioral adaptations of Ovibos-Moschatus and Rangifer Tarandus. Rangifer 12:13–25. Klein DR, DF Murray, RH Armstrong, and BA Anderson. 1997. Alaska. In MJ Mac, PA Opler, and PD Doran, Eds. National status and trends report. US National Biological Service, Washington, DC. Kolenosky GB. 1972. Wolf predation on wintering deer in east-central Ontario. J Wildlife Management 36:357–369. Krebs CJ. 1994. Ecology, the experimental analysis of distribution and abundance, 4th ed. Harper Collins College Publishers, NY, NY. 651 Pp. Larsen DG, DA Gauthier, and RL Markel. 1989. Causes and rate of moose mortality in the southwest Yukon. J Wildlife Management 53:548–557.
OCR for page 76
Wolves, Bears, and their Prey in Alaska: Biological and Social Challenges in Wildlife Management Leader-Williams N, TA Scott, and RM Pratt. 1981. Forage selection by introduced reindeer on South Georgia, South Atlantic Ocean and its consequences for the flora. J Applied Ecology 18:83–106. LeCount AL. 1982. Characteristics of a central Arizona black bear population. J Wildlife Management 46:861–868. Lehman NE, P Clarkson, LD Mech, TJ Meier, and RK Wayne. 1992. A study of the genetic relationships within and among wolf packs using DNA fingerprinting and mitochondrial DNA. Behavioral Ecology and Sociobiology 30:83–94. Lent PC. 1980. Synoptic snowmelt patterns in Arctic Alaska in relation to caribou habitat use. Pp. 71–77 in E Reimers, E Gaare, and S. Skjenneberg, Eds. Proceedings of the Second International Reindeer/Caribou Symposium, Direktoratet for vilt og ferskvannfisk, Trondheim, Norway. Licht DS and SH Fritts. 1994. Gray wolf (Canis lupus) occurrences in the Dakotas. American Midland Naturalist 132:74–81. Lindzey FG, KR Barber, RD Peters, and EC Meslow. 1986. Responses of a black bear population to a changing environment. Int Conf Bear Res and Management 6:57–63. Lindzey FG and EC Meslow. 1977. Population characteristics of black bears on an island in Washington. J Wildlife Management 41:408–412. Maessen O, B Freedman, J Svovoda, and MLN Nam. 1983. Resource allocation in high arctic vascular plants of differing growth form. Canadian J Botany 61:1680–1691. Martinka CJ. 1974. Population characteristics of grizzly bears in Glacier National Park, Montana. J. Mammal. 55:21–29. McEwan EH. 1968. Growth and development of the barren-ground caribou. II. Postnatal growth rates. Can J Zool 46:1023–1029. McLellan B. 1989. Dynamics of a grizzly bear population during a period of industrial resource extraction. III. Natality and rate of increase. Can J Zool 67:1865–1868. McLellan B. 1994. Density-dependent population regulation of brown bears. Pp. 15–24 In M Taylor, Ed. Density-dependent population regulation on black, brown and polar bears. Int Conf Bears Res and Management. Monogr Series No. 3. 43 Pp. McLellan BN and FW Hovey. 1994. The diet of grizzly bears in the Flathead river drainage of southeastern British Columbia. Can J Zool 73:704–712. McLellan BN. In press. Maintaining viability of brown bears along the southern fringe of their distribution. Int Conf Bear Res and Management. Mech LD. 1977. Wolf pack buffer zones as pre reservoirs. Science 198:320–321. Mech LD. 1986. Wolf numbers and population trend in the central Superior National Forest, 1967–1985. US Dept Agric for Serv Res Pap No. NC-270. 6 Pp. Mech LD and ME Nelson. 1989. Polygyny in a wild wolf pack. J Mammalogy 70:675–676. Mech LD. 1994. Biological impacts on wolves of exploitation and control: a response. In First Annual Conference of the Wildlife Society, Albuquerque, NM (abstract). Mech LD, SH Fritts, and D Wagner. 1994. Minnesota wolf dispersal to Wisconsin and Michigan. US National Biological Survey, Minneapolis, MN. Typewritten manuscript 6 Pp. Mech LD, TJ Meier, JW Burch, and LG Adams. 1995. Patterns of prey selection by wolves in Denali National Park, Alaska. In LN Carbyn, SH Fritts, and DR Seip, Eds. Ecology and conservation of wolves in a changing world. Canadian Circumpolar Institute, Occasional Papers 35, Edmonton, Alberta. Meier T, J Burch, LD Mech, and L Adams. 1995. Pack structure and genetic relatedness among wolf packs in a naturally regulated population. Pp. 293–302 In LN Carbyn, SH Fritts, and DR Seip, Eds. Ecology and conservation of wolves in a changing world. Canadian Circumpolar Institute, Occasional Papers 35, Edmonton, Alberta. Messier F and M Crête. 1985. Moose-wolf dynamics and the natural regulation of moose populations. Oecologia 65:503–512.
OCR for page 77
Wolves, Bears, and their Prey in Alaska: Biological and Social Challenges in Wildlife Management Messier F. 1985. Social organization, spatial distribution, and population density of wolves in relation to moose density. Canadian J Zoology 63:1068–1077. Miller DR. 1980. Wildfire effects of barren-ground caribou wintering on the taiga of northcentral Canada: a reassessment. Pp. 84–98 in Reimers, E, Gaare E and S. Skjenneberg, Eds. Proceedings of the Second International Reindeer/Caribou Symposium, Direktoratet for vilt og ferskvannfisk, Trondheim, Norway. Miller F. 1990. Peary caribou status report. Canadian Wildlife Service. West and North Reg Rep. 64 Pp. Miller SD, EF Becker, and WB Ballard. 1987. Black and brown bear density estimates using modified capture-recapture techniques in Alaska. Int Conf Bear Research and Management 7:23–35. Miller SD and WB Ballard. 1992. Analysis of an effort to increase moose calf survivorship by increasing hunting of brown bears in southcentral Alaska. Wildlife Society Bulletin 20:445–454. Miller SD. 1995. Impacts of heavy hunting pressure on the density and demographics of brown bear populations in southcentral Alaska. Alaska Dep Fish and Game. Fed Aid in Wildlife Res Grant W-24-3, Study 4.26. Fairbanks. AK. 28 Pp. Miller SD, GC White, RA Sellers, HV Reynolds, JW Schoen, K Titus, VG Barnes Jr., RB Smith, RR Nelson, WB Ballard, and CC Schwartz. 1997. Brown and black bear density estimation in Alaska using radio telemetry and replicated mark-resight techniques. Wildlife Monographs 113:1–55. Miquelle DG, JM Peek, and V Van Ballenberghe. 1992. Sexual segration in Alaskan moose. Wildlife Monographs 122:1–43. Molvar EM, RT Bowyer, and V Van Ballenberghe. 1993. Moose herbivory, browse quality, and nutrient cycling in an Alaskan treeline community. Oecologia 94:472–479. Murie AO. 1994. The wolves of Mount McKinley. Fauna of the national parks of the US. Fauna Ser No. 5. US Gov Print Off, Washington, DC. 238 Pp. Nagy JA, RH Russel, AM Pearson, MC Kingsley, and CB Larsen. 1983a. A study of grizzly bears on the barren-grounds of Tuktoyaktuk Peninsula and Richards Island, Northwest Territories, 1974–1978. Can Wildlife Ser Rep. 136 Pp. Nagy JA, RH Russel, AM Pearson, MC Kingsley, and BC Goski. 1983b. Ecological studies of the grizzly bear in the Arctic mountains, northern Yukon territory, 1972 to 1975. Can Wildlife Ser Rep. 104 Pp. Nelson ME and LD Mech. 1986. Relationship between snow depth and gray wolf predation on white-tailed deer. J Wildlife Management 50:90–91. Oldemeyer JL and WL Regelin. 1987. Forest succession, habitat management, and moose in the Kenai National Wildlife Refuge. Viltrevy (Swedish Wildlife Journal) Supplement 1:163–180. Osborne TO and others. 1996. Extent, cause, and timing of moose calf mortality in western interior Alaska. Alces 27:24–30. Oosenbrug S and LN Carbyn. 1982. Winter predation on bison and activity pattern of a pack of wolves in Wood Buffalo National Park. Pp 43–53 in FM Harrington and PC Paquet, Eds. Wolves: a worldwide perspective of their behavior, ecology, and conservation. Noyes Publications, Park Ridge, NJ. Packard JM and LD Mech. 1980. Population regulation in wolves. Pp. 135–150 in MN Cohens, RS Malpass, and HG Klein, Eds. Biosocial mechanisms of population regulation. New Haven: Yale Univ Press. Packard JM, LD Mech, and US Seal. 1983. Social influences on reproduction in wolves. Pp. 78–85 in LN Carbyn, Ed. Wolves in Canada and Alaska: their status, biology, and management Canadian Widlife Serv. Rep Ser No. 45. Page RE. 1985. Early calf mortality in northwestern British Columbia. M.Sc. thesis, Univ of Victoria, Victoria, BC 129 Pp.
OCR for page 78
Wolves, Bears, and their Prey in Alaska: Biological and Social Challenges in Wildlife Management Parker KL and CT Robbins. 1984. Thermoregulation in mule deer (Odocoileus hemionus hemionus) and elk (Cervus elaphus nelsoni). Canadian J Zoology 62:1409–1422. Pastor J. 1988. Moose, microbes, and the boreal forest. Bioscience 38:770–777. Pearson AM. 1975. The northern interior grizzly bear. Can. Wildlife. Serv. Rep. Ser. No. 34. 86 Pp. Peek JM. 1974. Initial response of moose to a forest fire in northeastern Minnesota. Am Midland Naturalist 91:435–438. Pegau RE. 1970. Succession in two exclosures near Unalakleet, Alaska. Canadian Field-Naturalist 84:12–25 Pegau RE. 1975. Analysis of the Nelchina caribou range. Biol Pap Univ Alaska special report 1. Peterson RO and RE Page. 1983. Wolf-moose fluctuation in Isle Royale National Park, Michigan USA. Annales Zoologici Fennica 74:251–253. Peterson RO, JD Woolington, and TN Bailey. 1984. Wolves of the Kenai Peninusla. Alaska Wildlife Monographs 88. 52 Pp. Peterson RO and RE Page. 1988. The rise and fall of Isle Royale wolves, 1975–1986. J. Mammalogy 69:89–99. Peterson RO. 1977. Wolf ecology and prey relationships on Isle Royale. National Park Service Scientific Monograph 11. 210Pp. Pianka ER. 1978. Evolutionary Ecology. 2nd Ed. Harper & Row, New York. Pimlott DH, JA Shannon, and GB Kolenosky. 1969. The ecology of the timber wolf in Algonquin Provincial Park. Ontario Dept Lands and For Res Rep (Wildl) No. 87. 92 Pp. Pritchard GT and CT Robbins. 1990. Digestive efficiencies of grizzly and black bears. Can J Zool 68:1645–1651. Pruitt WO. 1981. Application of the varrio snow index to over wintering North American barren-ground caribou (Rangifer tarandus arcticus). Canadian Field-Naturalist 95:363–365. Pulliam HR and NM Haddad. 1994. Human population growth and the carrying capacity concept. Bull. of the Ecological Society of America 75:141–157. Rausch RA. 1969. A summary of wolf studies in southcentral Alaska, 1957–1968. Trans North Am Wildlife and Nat Resour Conf 34:117–131. Regelin WL, CC Schwartz, and AW Franzmann. 1987. Effects of forest succession on the nutritional dynamics of moose forage. Viltrevy (Swedish Wildlife Journal). Supplement. 1:247–264. Reimers E. 1980. Activity pattern; the major determinant for growth fattening in Rangifer. Pp. 466–474 in E Reimers, E Gaare, and S Skjenneberg, Eds. Proceedings of the Second International Reindeer/Caribou Symposium, Direktoratet for vilt og ferskvannfisk, Trondheim, Norway. Reiner DC. 1996. Estimating bear habitat quality in southeastern British Columbia by measuring changes in bear body condition. M.S. thesis, Univ Montana, Missoula, MT. 81 Pp. Renecker LA and RJ Hudson. 1986. Seasonal energy expenditures and thermoregulatory responses of moose (Alces alces). Canadian J Zoology 64:322–327 Reynolds HV and TA Boudreau. 1990. Effects of harvest rates on grizzly bear population dynamics in the northcentral range. Alaska Dep Fish and Game. Div Wildlife Cons. Fed Aid in Wildlife Restoration Research progress report Project W-23-3, study 4.19. Reynolds HV. 1994. Effects of harvest on grizzly bear population dynamics in the northcentral Alaska Range . Alaska Dep Fish and Game. Fed Aid in Wildlife Res Grant W-24-3, Study 4.26. Fairbanks, AK. 28 Pp. Robbins CT, S Mole, AE Hagerman, and TA Hanley. 1987. Role of tannins in defending plants against ruminants reduction in dry matter digestion. Ecology 68:1606–1615. Rogers LL. 1987. Effects of food supply and kinship on social behavior, movements, and population growth of black bears in Northeastern Minnesota. Wildlife Monographs 97:1–72. Rosenthal GA and DH Janzen, Eds. 1979. Herbivores: their interaction with secondary plant metabolites. Academic Press, New York, NY.
OCR for page 79
Wolves, Bears, and their Prey in Alaska: Biological and Social Challenges in Wildlife Management Rothman RJ and LD Mech. 1979. Scent-marking in lone wolves and newly formed pairs. Animal Behavior. 27:750–760. Schlegel M. 1983. Factors affecting calf survival in the Lochsa elk herd. Job Completion Report, Federal aid project W-1600-R, Study 1, Job 3. Idaho Dept of Fish and Game, Boise, ID. Schoen JW and LR Beier. 1990. Brown bear habitat preferences and brown bear logging and mining relationships in southeast Alaska. Alaska Department of Fish and Game, Fed Aid in Wildlife Restor Final Report. Project W22:1–6 and W 23:1–3. Juneau, AK 90 Pp. Schwartz CC, WL Regelin, and AW Franzmann. 1988. Estimates of digestibility of birch, willow and aspen in moose. J Wildlife Management 52:33–37. Schwartz CC and AW Franzmann. 1991. Interrelationship of black bears to moose and forest succession in the northern coniferous forest. Wildlife Monographs 113:1–58. Scotter GW. 1967. Effects of fire on barren-ground caribou and their forest habitat in northern Canada. Trans North Amer Wildlife Nat Res Conf 32:246–259. Scotter GW. 1995. Influence of harassment by wolves, Canis lupus, on barren-ground caribou Rangifer tarandus groenlandicus, movements near the Burnside River, Northwest Territories. Canadian Field-Naturalist 109:452–453. Seal US, LD Mech, and V VanBallenberghe. 1975. Blood analyses of wolf pups and their ecological and metabolic interpretation. J Mammalogy 56:64–75. Seber GAF. 1982. The estimation of animal abundance and related parameters, 2nd ed. Griffin, London. 654 Pp. Seip DR. 1992. Factors limiting woodland caribou populations and their interrelationships with wolves and moose in southeastern British Columbia. Canadian J Zoology 70:1494–1503. Seip DR and K Brown. 1996. Introduction to the population ecology of North American caribou. Rangifer 9:11–12. Selkregg LL. 1976. 1974–1976 Alaska Regional Profiles: Vol. I Southcentral Alaska, 225 Pp.; Vol. II Arctic, 218 Pp.; Vol. III Southwest, 313 Pp.; Vol. VI Southeast, 233 Pp.; Vol. V Northwest, 265 Pp.; Vol. VI Yukon, 346 Pp. Arctic Information and Data Center, Anchorage, AK. Sellers RA. 1994. Dynamics of a hunted brown bear population at Black Lake, Alaska. 1993 Annual Progress Report. Alaska Department of Fish and Game. 61 Pp. Sellers RA, SD Miller, TS Smith, and R Potts. 1993. Population dynamics and habitat partitioning of a naturally regulated brown bear population on the coast of Katmai National Park: 1993 Annual progress report. Alaska Dept of Fish and Game, Div of Wildlife Conservation, Juneau. Servheen C 1990. The status and conservation of the bears of the world. Eighth Int Conf Bear Res and Management 9:4–16. Servheen C, FW Hovey, BN McLellan, R Mace, W Wakkinen, W Kasworm, D Carney, T Manley, K Kendall, and R Wielgus. 1994. Report of the northern ecosystems researches on grizzly bear population trends in the north and south forks of the Flathead, and the Blackfeet Indian Reservation of the NCDE; the Cabinet-Yank ecosystem; and the Selkirk ecosystem; and future data needs to improve trend estimates. US Fish and Wildlife Service. Missoula, MT. Sharkey MJ. 1970. The carrying capacity of natural and improved land in different climatic zones. Mammalia 34:564–572. Skogland T. 1985. The effect of density-dependent resource limitations on the demography of wild reindeer. J Animal Ecology 54:359–374. Skoog R. 1968. Ecology of the caribou (Rangifer tarandus granti) in Alaska. Ph.D. thesis, Univ CA, Berkeley. Smith TR and MR Pelton. 1990. Home ranges and movements of black bears in a bottomland hardwood forest in Arkansas. Pp. 213–218 in LM Darling and WR Archibald, Eds. Bears: their biology and management. International Association for Bear Research and Management, Coeur d'Alene.
OCR for page 80
Wolves, Bears, and their Prey in Alaska: Biological and Social Challenges in Wildlife Management Spalding DJ. 1990. The early history of moose (Alces alces) distribution and relative abundance in British Columbia, Victoria BC. Royal British Columbia Museum. Stenlund MH. 1955. A field study of the timber wolf, Canis lupus, on the superior National Forest. MN. MN Dept Conservation Tech Bull 4:1–55. Stephenson RO. 1975. Wolf Report. Alaska Sept of Fish and Game, Juneau. Stephenson RO, WB Ballard, CA Smith, and K Richardson. 1995. Wolf biology and management in Alaska 1981–92. Pp. 43–54 in LN Carbyn, SH Fritts, and DR Seip, Eds. Ecology and conservation of wolves in a changing world. Canadian Circumpolar Institute, Occasional Papers 35, Edmonton, Alberta. Storer TI and LP Tevis. 1955. California grizzly. University of California Press, Berkeley. Stringham SF. 1980. Possible impacts of hunting on the grizzly/brown bear, a threatened species. Int Conf Bear Res and Management 4:337–349. Stringham SF. 1990. Grizzly bear reproductive rate relative to body size. Int Conf Bear Res and Management 8:433–443. Telfer ES and JP Kelsall. 1984. Adaptation of some large North American mammals for survival in snow. Ecology 65:1828–1834. Titus K and LR Beier. 1994. Population and habitat ecology of brown bears on Admiralty and Chichagof Islands, Alaska Dept of Fish and Game, Div of Wildlife Cons, Grant W-24-2 study 4.22, Juneau, AK. US Department of Agriculture. 1994. Wolf depredation on livestock in Minnesota annual update of statistics. USDA, Animal and Plant Health Inspection Service, Animal Damage Control, Grand Rapids, MN. Valkenburg P and JL Davis. 1986. Calving distribution of Alaska's Steese-Fortymile herd: a case of infidelity? Rangifer (special issue) 1:315–325. Valkenburg P, JL Davis, JM Ver Hoef, RD Boertje, ME McNay, RM Egan, DJ Reed, CL Gardner and RW Tobey. 1996. Population decline in the Delta caribou herd with reference to other Alaskan herds. Rangifer (special issue) 9:53–62. Van Ballenberghe V and LD Mech. 1975. Weights, growth, and survival of timber wolf pups in Minnesota. J Mammalogy 56:44–63. Van Ballenberghe V, AW Erickson, and D Byman. 1975. Ecology of the timber wolf in northeastern Minnesota. Wildlife Monographs 43. Van Ballenberghe V and DG Miquelle. 1993. Mating in moose: timing, behavior, and male access patterns. Canadian J. Zoology 71:1687–1690. Van Ballenberghe V. 1983. Extraterritorial movement and dispersal of wolves in southcentral Alaska . J Mammalogy 64:168–171. Viereck LA. 1973. Wildfire in the taiga of Alaska. Quat Res (NY) 3:465 –495. Wayne RK, DA Gilbert, N Lehman, K Hansen, A Eisenhower, D Girman, LD Mech, PJP Gogan, US Seal, and RJ Krumenaker. 1991. Conservation genetics of the endangered Isle Royale gray wolf. Conservation Biology 5:41–51. Wayne RK, N Lehman, and TK Fuller. 1995. Conservation genetics of the gray wolf. Pp. 399–408 in LN Carbyn, SH Fritts, and DR Seip, Eds. Ecology and conservation of wolves in a changing world. Canadian Circumpolar Institute, Occasional Papers 35, Edmonton, Alberta. White GC. 1983. Evaluation of radio tagging, marking, and sighting estimators of population size using Monte Carlo simulations. Pp. 91–103 in JD Lebreton and PM North, Eds. Marked individuals in the study of bird populations. Biskhauser Verlog Basel, Switzerland. White RG, BR Thompson, T Skogland, SJ Person, DE Russel, D Holleman, and JR Luick. 1975. Ecology of caribou at Prudhoe Bay, Alaska. Biol Pap Univ Alaska, Special Rep 2. Pp 151–201. White RG and J Trudell. 1980. Habitat preference and forage consumption by reindeer (Rangifer tarandus tarandus) and caribou (Rangifer tarandus granti) near Atkasook, Alaska. Arctic Alpine Research 12:511–530.
OCR for page 81
Wolves, Bears, and their Prey in Alaska: Biological and Social Challenges in Wildlife Management White RG and FA Milan. 1981. Reindeer herds of the Kuskowin, Alaska. Proc Alaska Sci Conf 32:1–10. Wielgus RB. 1993. Causes and consequences of sexual habitat segregation in grizzly bears. Ph.D. thesis, University of British Columbia. Wielgus RB and FL Bunnell. 1994. Sexual segregation and female bear avoidance of males. J Wildlife Management 58:405–413. Wolfe RJ. 1996. Subsistence food harvests in rural Alaska, and food safety issues. Paper presented to the Institute of Medicine. National Academy of Sciences Committee on Environmental Justice, Spokane, WA, August 13. Wydeven AP, RN Schultz, and RP Thiel. 1995. Monitoring of a recovering gray population in Wisconsin 1979–1991. Pp. 147–156 in LN Carbyn, SH Fritts, and DR Seip, Eds. Ecology and conservation of wolves in a changing world. Canadian Circumpolar Institute, Edmonton. Alberta. Wydeven AP. 1993. Wolves in Wisconsin: reconolization under way. International Wolf 3:18–19. Zackrisson O. 1977. Influence of forest fires on the North Swedish boreal forest. Oikos 29:22–32.
Representative terms from entire chapter: