Faculty of Environmental Design
University of Calgary
2500 University Dr. NW
Calgary AB Canada T2N 1N4
Northeast Science Station
Cherskii, Sakha Republic (Yakutia)
Robert O. Stephenson
Alaska Department of Fish & Game
1300 College Road
Fairbanks AB USA 99701
Melissa C. Chapin
Institute of Arctic Biology
University of Alaska-Fairbanks
2102 Goldstream Rd.
Fairbanks AB USA 99709
The following article was originally presented at the International Bison Conference in Edmonton, Alberta in August 2000. The conference covered a wide array of bison topics including production, marketing, genetics, history and much more. This article has been reprinted with the permission of the IBC2000 Chairman.
Various forms of the genus Bison have been important faunal elements in northwestern North America and Siberia for 400,000 years or more. The bison was the most common large herbivore during much of the last 100,000 years. Zooarchaeological, ethnographic, and historical records indicate the general chronology and suggest causes for the decline and local extirpation of bison in North America and adjacent Siberia. The Bering Land Bridge geographically linked these regions during the late Pleistocene. There is some evidence that hunting by humans played a central role in the history of northern bison, first in Siberia where bison were extirpated during the past 2000 years, then in North America where only remnant populations remained at the end of the 19th century. The recovery of wood bison (B. b. athabascae) in their original range is currently the focus of cooperative efforts between Canada, Alaska and Russia. Since 1963, Canada has worked to reestablish healthy wood bison populations. There are currently six wild populations in Canada that are free of tuberculosis and brucellosis. In addition, there are three captive breeding herds representing two maternal lineages (Nyarling and Slave River Lowlands). Further recovery of wood bison in Canada is constrained by human encroachment, competing land uses, and the occurrence of cattle diseases in free-ranging bison populations in one area.
The Alaska Department of Fish & Game has been involved in evaluating and planning the reintroduction of wood bison in the Yukon Flats since 1991. This proposed reintroduction enjoys substantial public and agency support, reflecting interest in restoring a historically important animal and the various potential benefits associated with free-roaming populations of wood bison. The Government of the Republic of Sakha (Yakutia), Russian Federation, has developed an experimental program to evaluate the restoration of large grazing mammals in the Kolyma River lowlands in Siberia. The goal of the project is to evaluate the possibility of restoring a natural steppe ecosystem and by bringing together an assemblage of large mammals many of which were present in northern Siberia during the Holocene era. The project also offers the potential to contribute to future sustainable economic development based on hunting, tourism, and meat production. The Alaskan and Siberian initiatives are particularly important to wood bison conservation in view of the limited availability of habitat for wood bison in Canada, and the risk of infection of some Canadian herds with cattle diseases. They represent additional opportunities to secure the survival of the subspecies in geographically separate populations and to augment Canada's participation in international efforts to conserve species at risk and encourage sustainable development.
Holocene, bison, endangered species, ecological restoration, recovery
Bison have been a prominent faunal element in northwestern North America and northeast Asia for 400,000 years or more. The bison was one of the most common large herbivores during the last 100,000 years. Until recently, the Bering Land Bridge linked these regions. Therefore, it is not surprising that the last types of bison to exist in these regions were similar, small-horned forms represented by the surviving wood bison. Various human societies interacted with bison and other large herbivores in Eurasia for tens of thousands of years and in North America prior to and after the glacial recession some 10-15,000 years ago. It is clear that humans were responsible for the near extinction of bison (B. b. bison) on the Great Plains of North America (Roe 1951) and wood bison from northern Alberta, British Columbia and the southwestern Northwest Territories (Gates et al. 1992). The disappearance of bison and some other large herbivores from northern areas on either side of the Bering Strait during the mid to late Holocene likely involved the combined effects of changes in habitat distribution and hunting by humans (Ward 1997; Stephenson et al., in press). The disappearance of bison and other large grazers may have important consequences for the structure and function of northern ecosystems (Zimov et al. 1995). It also affected human cultures and economies (Cohen 1977).
Efforts to protect, conserve or reestablish wood bison populations in the North have spanned more than a century in Canada and are now the focus of international cooperation between Canada, Alaska and Russia. Modest success has been achieved in Canada, and there remain both opportunities and challenges in restoring the species to additional parts of its original range. We briefly review the taxonomy and history of wood bison, their conservation and recovery, and describe opportunities to reestablish additional populations in Canada, Alaska and Siberia.
Fossil evidence indicates there was a single species of bison in northern Eurasia and North America during the middle and late Pleistocene (Guthrie 1990). During the last (Wisconsin) glaciation there were two separate populations of bison in North America (Guthrie 1990; van Zyll de Jong 1993). The steppe bison (B. priscus, McDonald 1981 and Guthrie 1990; or B. b. priscus, Van Zyll de Jong 1986), a relative newcomer from Eurasia, occupied Beringia and was adapted to the rigors of the cold steppe. Bison antiquus, a descendant of an invasion of bison from Eurasia during the preceding Illinoian glaciation, was adapted to a temperate climate and open woodlands, and persisted south of the continental ice sheet. By about 14,000 years ago, Eurasia and Alaska were separated by rising sea levels resulting from climatic warming.
Subsequently, bison on both side of the Bering Isthmus underwent parallel evolution from large horned to small horned forms (van Zyll de Jong 1993). Three bison taxa (wood bison, plains bison, and European bison) continue to exist. The two North American subspecies (B. b. athabascae and B. b. bison) represent the most recent variants on the North American continent, while the European bison (B. bonasus), which survived only in the forests of eastern Europe, is the only extant form of bison on the Eurasian continent. A northern form of small-horned bison, similar to wood bison, became extinct in eastern Siberia by the late Holocene (van Zyll de Jong 1993). The taxonomy of this form is not well defined.
A corridor began to form in the ice sheet separating Beringia from central North America about 13,000 years ago, after which northern and southern forms of bison dispersed from their respective ranges and intermingled (McDonald 1981). Existing North American bison are descendants of these two Pleistocene forms. However, the contribution of the southern antiquus and the northern priscus in the evolution of modern North American bison is not well understood. The southern antiquus was probably more widely distributed and abundant than priscus, and may have played a larger role in the evolution of modern North American bison in the southern part of their range (van Zyll de Jong 1993). The fusion of northern and southern forms produced the early Holocene form, occidentalis, which underwent evolutionary change leading to the two modern North American subspecies by about 5000 years ago.
Adapted to northern woodlands, the non-migratory wood bison evolved in the northwestern portion of the species' range, while the migratory plains bison evolved in the extensive grasslands of central and southern North America. The subspecific status of Holocene bison in eastern Siberia is not understood in detail, but available evidence indicates they were morphologically similar, and ecologically equivalent, to wood bison. A well preserved skull from the Kolyma River lowlands, believed to be of late Pleistocene or early Holocene age, is taxonomically intermediate between occidentalis and B. b. athabascae, demonstrating a close affinity between these forms (van Zyll de Jong 1986, 1993). Bison remains have been recorded at archaeological sites dating to 900 A.D. in southern Siberia near Lake Baikal (Lazarev et al. 1998).
Wood bison are recognized as a subspecies based on morphology and DNA (Wood Bison Recovery Team in press). Geist (1991) suggested the subspecific status is not warranted and that observed differences are environmentally induced. More recent morphological studies of plains bison and wood bison show differences in cranial and skeletal characteristics (van Zyll de Jong 1986), as well as in the anterior slope of the hump, location of the highest point on the hump, angle of the hump, cape variegation and demarcation, upper front leg hair, frontal display hair, ventral neck mane and beard. The traits were not sensitive to geographic location, suggesting that phenotypic differences are genetically controlled (van Zyll de Jong et al. 1995).
There are similarities as well as differences in mitochondrial (mtDNA) and genomic DNA (Bork et al. 1991; Strobeck et al. 1993), and erythrocyte antigens and blood proteins in plains bison and wood bison (Zamora 1983; Peden and Kraay 1979). Wilson and Strobeck (1999) studied variability in 11 microsatellite loci of genomic DNA in 11 North American bison herds. Genetic distances between wood bison and plains bison subpopulations were larger than those within either subspecies (Wilson and Strobeck 1999), indicating that wood bison continue to function as a genetic entity separate from plains bison, despite the introduction of plains bison into wood bison range in the 1920s. Strobeck et al. (1993) compared sequence divergence in a section of D-loop in the mtDNA of a small number of wood and plains bison and found that differences between the two subspecies are approximately the same or less than within plains bison. The rate of sequence divergence in mtDNA is on the order of 1% to 2% per million years (Wilson et al. 1985). These findings are consistent with the view that wood bison and plains bison existed as reproductively isolated populations during the last 5,000 -10,000 years, a relatively short time in evolutionary terms, and probably also reflect introgression of plains bison mtDNA into remnant wood bison populations during the 1920s (Van Zyll de Jong 1995). A parallel "evolution", directed by humans, occurred over a similar period (6000 years) with the radiation of various breeds of domestic cattle from the wild progenitor Bos primigenius. The differences between North American subspecies probably also reflects the extensive periods of separation by ice sheets during the late Pleistocene. The occurrence of dated skeletal remains suggests an eastward expansion of northern small-horned populations from the glacial refugia in Alaska and western Yukon following the last glacial recession (Stephenson et al., in press).
Over millennia the "Great Bison Belt" that extended across Eurasia into Beringia and southward across North America, diminished in size (Guthrie 1980, 1990: 51). Although bison likely disappeared from Siberia by the late Holocene, they were still widely distributed in North America, including the Great Plains, in eastern woodlands, and in northwestern Canada and Alaska (Soper 1941; Dary 1989; Stephenson et al., in press). Bison in northern habitats were not as numerous as bison on the Great Plains. The phenomenon of a large migratory population was unique to the Holocene Great Plains, with its huge expanse of contiguous grasslands (Guthrie 1980, 1982, 1990). Beginning in the 1700s, the availability of horses altered the mobility and hunting strategies of Native groups (Isenberg 2000). Horses eventually became sufficiently abundant and widespread to compete with bison for forage (Flores 1996; Fisher and Roll 1997). A number of studies describe the importance of hunting by humans in the dynamics of late Holocene plains bison populations (Roe 1951; Guthrie 1980; Speth 1983; Flores 1991, 1996; Belue 1996; Dobak 1996; Fisher and Roll 1997; Haynes 1997; Morgan 1997; Martin and Szuter 1999; Isenberg 2000).
Radiometric and archaeological data and historic accounts by Native Elders in Alaska and adjacent Canada indicate that the original range of wood bison in North America (Fig. 1) included northern Alberta, northeastern British Columbia, a small portion of northwestern Saskatchewan, the western Northwest Territories, Yukon, and much of Alaska (Skinner and Kaisen 1947; Harington 1977; van Zyll de Jong 1986; Guthrie 1990; Lotenberg 1996; Stephenson et al., in press). Radiocarbon dates for bison remains spanning much of the past 10,000 years have been recovered from a number of archaeological sites. Radiometric and archaeological data demonstrate that the mid to late Holocene distribution of wood bison included much of eastern Alaska, southern Yukon, the western Northwest Territories, and possibly western Alaska. Radiocarbon data indicate that bison occurred in Alaska at least as recently as 170 ± 30 years ago. Well-preserved bones also indicate their presence along the arctic coast of Yukon and Northwest Territories and on Victoria Island (Le Blanc 1984; Morrison 1997). A bison skull with attached horn cores was discovered in 1987 at a site along the banks of the Old Horton River Channel (Le Blanc 1988), near the extreme northwestern arctic coast of the Northwest Territories. The skull was dated to 420 ± 65 years ago (Harington 1990), and was found in close association with the skulls of two muskoxen at a site that had a long history of human use, primarily for caribou hunting and butchering. Faunal remains of large mammals such as moose and bison are well represented at various mid-Holocene sites in southern Yukon (cf. MacNeish 1964; Workman 1978), but few identifiable large mammal bones have been found at late Holocene sites, probably reflecting changes in cooking and preparation techniques.
Written records document the occurrence of bison in the 18th and 19th centuries in the southern Yukon, western NWT, Alberta and British Columbia (Gates et al. 1992, Lotenberg 1996). Bison apparently disappeared from Alaska and western Yukon before their presence was recorded in written records. However, oral narratives provide insight into the late Holocene distribution, human use, and disappearance of wood bison in Alaska and adjacent Yukon. Historic accounts from Native elders in interior Alaska describe how bison were a source of food and raw materials. Hides were tanned and used to make robes, pillows, and clothing, and sewing thread was made from bison hair. Bison were hunted with bow and arrow, spears, dogs, and with snowshoes to gain advantage in deep snow, and possibly with drives. The accounts indicate that substantial populations of wood bison declined or disappeared from Alaska by the early or mid 1800s, with the occurrence of small numbers in the eastern interior as late as the early 1900's (Stephenson et al., in press). The accounts are in general agreement with oral history obtained from First Nation elders in the Yukon (Lotenberg 1996), which also indicated a decline in numbers during the last 400 years and the eventual extirpation of wood bison by the mid-19th century.
Historic accounts as well as paleontological and archaeological data indicate that humans hunted wood bison until their disappearance from Alaska and Yukon during the last few hundred years. Wood bison had apparently become scarce shortly before early Euroamerican explorers, naturalists, and entrepreneurs entered the region from the east, and before firearms became widely available (Jennings 1968, Holmes and Bacon 1982, Guthrie 1990). This decline coincided with the historical period during which plains bison were extirpated from the woodlands east of the Mississippi River (Dary 1989; Belue 1996).
Wood bison were nearly eliminated from their remaining range in Canada during the late 1800s, coinciding with the rapid decline of plains bison between 1840 and 1900 (Raup 1933). With the advent of the fur trade extensive hunting for meat and pemmican played a major role in the decline of wood bison in Canada (Gates et al. 1992). The most rapid decline occurred after 1860 (Soper 1941). In 1888 the total population was estimated at between 500 and 600 animals (Schultz 1888). By 1891, 300 wood bison remained in the wilderness between Great Slave Lake and the Peace-Athabasca Delta (Ogilvie 1893). This population reached an estimated low of approximately 250 during 1896-1900 (Soper 1941). After 1900, wood bison were occasionally observed throughout their range in Canada, but numbers were small except in the Slave River lowlands and in the area later designated as Wood Buffalo National Park (Gates et al. 1992). One of the last bison in northeastern British Columbia was shot in 1906 (MacGregor 1952). Lotenberg (1996) cites Clarke (c. 1945: 28) who stated that Indians killed a bull at Lower Post, British Columbia in 1939. Wood bison were extirpated from most of their original range in North America by the late 19th century, persisting only in the area south of Great Slave Lake, where the Canadian Government afforded some protection.
Bison persisted in northern Eurasia into the middle or late Holocene but apparently disappeared earlier than in Alaska and adjacent areas in Canada (van Zyll de Jong 1993; Rusanov 1975; Flerov 1979; Archipov 1989; Lazarev et al. 1998). Numerous bison remains have been found in Holocene deposits, but have not been dated. Van Zyll de Jong (1986, 1993) concluded that bison on either side of the Bering Strait underwent parallel evolution from large to small horned forms. A skull from the Kolyma River lowlands falling within the range of North American Holocene bison probably dated to at most 5,000-6,000 years ago (Sher 1971). This specimen suggests that "bison marooned on either side of the Bering Strait underwent parallel evolution, from large-horned to small-horned forms," and that bison became extinct in Siberia after the thermal optimum (ca. 9000-2500 years ago) (van Zyll de Jong 1993). Flerov (1979) also concluded that B. bison became extinct in Siberia about 6000 years ago, after the thermal optimum.
Skeletal remains and petroglyphs indicate that a small-horned form of bison persisted in eastern Siberia and coexisted with humans for an extended period during the Holocene. Bison remains have been found in association with Neolithic and post-Neolithic archaeological sites at several locations in southern Yakutia near Olekminsk and the confluence of the Olekma and Lena Rivers (Lazarev et al. 1998). Petroglyphs that appear to portray bison, estimated to be 2,000 years old, occur in the Olekminsk region (Archipov 1989). This evidence has led some investigators to suggest that bison were extirpated in Siberia approximately 2,000 years ago (Rusanov 1975). More recent studies suggest that bison existed in conjunction with human settlements over a wide area, possibly until a later date. Bison remains have been found at archaeological sites on the upper Kolyma River and on Kamchatka (Mochanov and Savvinova 1980; Dikov 1979). Recent archaeological studies describe the occurrence of remains of a reduced (small-horned) form of Pleistocene bison in association with cultural artifacts at settlements adjacent to the west and east shores of Lake Baikal, dating as late as the 8th or 9th century A.D. (1,100-1,200 years BP) (Lazarev et al. 1998; Vereschagin and Baryshnikov 1985; Ermolova 1978). The available evidence suggests that descendants of steppe bison survived in Siberia well into the Holocene and were similar to wood bison in size, form and function.
The reasons for the disappearance of bison from eastern Siberia (western Beringia) are not known with certainty. The Pleistocene overkill hypothesis suggests that human hunting caused or contributed to the extinction of various large mammals during the late Pleistocene and early Holocene (Hopkins 1967; Jelinek 1967; Martin and Klein 1984; Martin and Wright 1989, Ward 1997). Mammoths survived on Wrangel Island until 3700 BP (Vartanyan et al. 1993), a time that coincided with the arrival of humans (Lister 1993). Early humans apparently relied on sharpened bones and antlers as hunting weapons. However, by 10000-15000 years ago, Beringia was occupied by people who hunted with stone microblades inserted in wooden or bone shafts (West 1981; Guthrie 1990). They extensively hunted steppe megafauna (Haynes 1982). The spread of microblade hunting technology coincides with a period of significant change in steppe megafauna and with the transition from steppe to tundra vegetation in much of Beringia (Zimov et al. 1995). Perhaps some Pleistocene-grazing mammals could have survived in the absence of human hunting.
Climatic reconstructions using general circulation models suggest the region's climate is presently warmer than during the Pleistocene, but that precipitation and storm patterns may be similar (Manabe and Broccoli 1985; Rind 1987; Zimov et al. 1995). Variations in macroclimate across Beringia (Manabe and Broccoli 1985; Kutzbach and Guetter 1986; Rind 1987) and in microclimate relative to slope, aspect, and elevation (Barnosky et al. 1987; Edwards and Armbruster 1989) were probably greater than the magnitude of change in these parameters from the Pleistocene to the present (Manabe and Broccoli 1985; Kutzbach and Guetter 1986; Rind 1987), suggesting that climatic changes alone do not explain the disappearance of Pleistocene megafauna. In contrast, evidence from paleontology, archaeology, and related disciplines suggests that exploitation by humans affected the abundance and distribution of bison prior to and during the last several millennia.
Some combination of climatic and environmental change and human hunting probably led to the extinction of the Pleistocene grazers and a decline in associated grassland vegetation (Martin and Klein 1984; Guthrie 1990; Zimov et al. 1995; Ward 1997). Based on the historical record, there is no way to determine whether a climate-induced change in vegetation, combined with hunting, caused extinction of the megafauna, or the loss of the megafauna as a result of human hunting led to the change in vegetation. However, recent research provides support for the concept that megafauna played an important role in maintaining grassland vegetation during the Pleistocene (Zimov et al. 1995), just as African grazers currently play a key role in maintaining the savannahs of Africa (Owen-Smith 1988).
The relationships between bison, human populations and other environmental factors have been diverse, with no single defining pattern. Nevertheless, it is clear that during the last millennium bison populations were dramatically reduced in Eurasia and much of North America in areas where the amount and distribution of suitable late Holocene habitat were more limited than on the Great Plains. Wood bison were extirpated from most of their original range in northern Canada, and rapidly approached extinction following over-hunting during the 19th century. European bison also declined during the Holocene, with less than 100 wisent (B. bonasus) persisting in the forests of eastern Europe in the early 1900s (Gstalter and Lazier 1996). Habitat reduction and overhunting were key factors causing their near extinction. Bison persisted in northern Eurasia into the middle or late Holocene but apparently disappeared earlier than in Alaska or adjacent parts of Canada (van Zyll de Jong 1993; Rusanov 1975; Flerov 1979; Archipov 1989; Lazarev et al. 1998). Plains bison persisted in a large region in North America despite being hunted extensively before the introduction of firearms. Annual long-range migration was likely a key factor accounting for the relative abundance of plains bison, similar to some African ungulates (Fryxell et al. 1988).
Conservation and recovery in Canada
History of the recovery program
Bison conservation efforts began in Canada in 1877 with the passing of the Buffalo Protection Act (Hewitt 1921). This measure was largely ineffective until it was strengthened in 1893 when the Dominion Government passed additional statutes (Soper 1941). Enforcement was minimal until it was assigned to the Northwest Mounted Police in 1897. The first police outpost in wood bison range was established at Fort Fitzgerald on the Slave River in 1907, when formal patrols of the region began. In 1911, six Buffalo Rangers were appointed to patrol the remaining range of the wood bison. The wood bison population increased slowly to approximately 500 by 1914 (Banfield and Novakowski 1960).
Wood Buffalo National Park (WBNP) was established in 1922 by an Order-In-Council under the Forest Reserves and Parks Act in an attempt to save the wood bison from extinction and to protect its habitat (Soper 1941; Lothian 1979). The total number of wood bison on the southern range at that time was estimated at between 1500 and 2000 (Siebert 1925; Soper 1941). The conservation of wood bison and plains bison became linked in the 1920's, when more than 6,000 surplus plains bison from Wainright Buffalo Park in east central Alberta, were shipped north to Wood Buffalo Park (Graham 1924; Lothian 1981). During 1925-1928, plains bison were released at several sites along the west bank of the Slave River, south and north of Hay Camp, in range that was occupied by wood bison (Soper 1941). The introduction of plains bison into wood bison range occurred despite challenges from the American Society of Mammalogists (Howell 1925) and individual biologists (Harper 1925; Saunders 1925). The plains bison carried Mycobacterium bovis and Brucella abortus, the causative agents of bovine tuberculosis and brucellosis. Bison herds in and around WBNP remain infected to this day (Tessaro et al. 1989). Despite some ensuing hybridization, WBNP bison remain genetically and morphologically distinguishable from plains bison (van Zyll de Jong et al. 1995; Wilson and Strobeck 1999) and warrant conservation as separate forms (Wood Bison Recovery Team, in press).
Three projects have salvaged wood bison from the WBNP area to establish captive breeding herds or wild populations that are free of bovine tuberculosis and brucellosis. During winter 1963, 77 wood bison were captured at Needle Lake and Nyarling River in northwestern WBNP to establish a captive-breeding herd near Fort Smith, NWT. After being tested for diseases, 19 bison were transported to a holding corral near Fort Smith. In June 1963, a second outbreak of anthrax among free-ranging bison north of the holding facility led to the transfer the captive herd to an uninhabited area on the west side of Great Slave Lake. In August, four adult males, 10 adult females and four calves were moved to the Mackenzie Bison Sanctuary near Fort Providence to establish a free-ranging herd (Gates and Larter 1990).
A second roundup of wood bison was held in northern WBNP in 1965. Twenty-one of 47 animals captured were successfully transferred to Elk Island National Park (EINP) in central Alberta. Two additional calves were transferred to EINP between 1966 and 1968. The translocated stock carried bovine tuberculosis and brucellosis. However, a rigorous management protocol involving isolation and quarantine of neonates eradicated these diseases by 1971. The EINP breeding herd has provided disease-free founding stock for five free-ranging populations in Canada, two captive breeding herds, and several zoo and park herds. Wood bison have also been provided to two Canadian First Nations to establish commercial ranches in cases where there was a potential conservation benefit involving release of bison to the wild. In the absence of opportunities to use surplus stock for conservation projects, EINP sold wood bison at auction for the first time in 1998.
A third salvage and recovery project was initiated in 1996. Over a three-year period, 62 newborn calves were captured from the Hook Lake herd in the Slave River Lowlands (SRL). They were maintained in enclosures near Fort Resolution and treated with antibiotics for exposure to tuberculosis and brucellosis (Gates et al. 1997). In 2000 the herd consisted of 58 bison captured from the wild and 32 born in captivity. The animals have remained negative for tuberculosis and brucellosis after nine biannual whole herd tests (Gates et al. 1999; J. Nishi pers. comm.). The population is managed under quarantine as an experimental captive breeding herd.
The establishment of a wild population near Fort Providence in 1963 and a captive population at EINP in 1965 represented the first major steps toward the recovery of wood bison in Canada. In 1973, representatives of the Canadian Wildlife Service, Parks Canada and Territorial and Provincial wildlife management agencies in western Canada began meeting annually to coordinate recovery efforts for wood bison. A Wood Bison Recovery Team was formally established in 1988 under the Recovery of Nationally Endangered Wildlife initiative (RENEW). The Recovery Team fosters collaboration between wildlife management agencies within Canada (Manitoba, Alberta, British Columbia, Yukon, Northwest Territories, Parks Canada, and the Canadian Wildlife Service) and with other countries (Alaska and the Russian Federation). Wood bison are classified as "threatened" in Canada's legal classification of endangered wildlife, having been downlisted from "endangered " in 1988. As a range state, Alaska has been a participating member of the Recovery team since 1991. In addition, the Recovery Team has cooperated with the Russian Federation regarding its interest in acquiring wood bison for an ecological experiment and possible reintroduction in eastern Siberia.
Recovery projects in Canada
In 1999/2000 there were approximately 2800 wood bison in six free-ranging disease-free herds, 708 in four captive breeding herds, and about 2900 in diseased, free-ranging herds in and around Wood Buffalo National Park (Fig. 1).
The Mackenzie population was the first disease-free herd to be established in the wild. It was founded in 1963 with the transfer of 18 bison from Wood Buffalo National Park to the Fort Providence area west of Great Slave Lake. The herd currently numbers about 2000 animals and is the largest of six disease-free wild populations. The other free-ranging disease-free herds were also founded with stock from Elk Island National Park.
The Nahanni herd (Northwest Territories) was established in 1980, when 28 bison were released. It was supplemented with 12 bison in 1989 and 60 in 1998. The Nahanni herd is estimated to number 170. A population was successfully reestablished in the southern Yukon with releases totaling 180 bison between 1988 and 1991. The population currently includes about 500 wood bison.
A free-ranging herd of wood bison was established in the northern Interlake Region of Manitoba with a total of 22 wood bison being released near Chitek Lake, north of the Waterhen Wood Bison Ranch, during 1991-93. Nine additional animals were released in January 1993. The herd increased from 31 animals in July 1995 to approximately 50 by December 1996. The population was estimated to be approximately 70 during winter 1999-2000. The area has the potential to support 400-500 wood bison. The Province of Manitoba established a protected area for wood bison and their habitat in the Chitek Lake region.
A program to reestablish wood bison in northwestern Alberta was initiated in 1981 in cooperation with the Dene Tha First Nation. A free-ranging herd was established in 1993 when 49 bison escaped prematurely from a fenced area. The population was estimated as 130 in winter 1999/2000. Wood bison from the Hay-Zama herd occasionally wander into the Hay River drainage in northeastern British Columbia. The Government of Alberta established a 36,000-km2 management area in the northwestern corner of the province, within which wood bison are protected. This area has the potential to support at least 400 wood bison. However, the herd will be maintained at between 250 and 400 animals until the WBNP disease issue is resolved. Bison that occur between the designated bison management zone and WBNP are not protected from hunting, making the area a buffer zone that reduces the potential for disease transmission. Bison are designated as "livestock" in the Alberta Wildlife Act, except within the bison management area.
Wood bison were reintroduced into northeastern British Columbia in March 1995. Forty-nine were transferred from EINP to Aline Lake in the Nordquist Flats area in the Upper Liard River Valley. The site is approximately 80 km from the southern part of the Nahanni herd's range along the Beaver River. Thirty-nine bison were counted prior to calving in 2000. The Nahanni and Nordquist herds are eventually expected to coalesce. The combined herd will be managed to reach a population objective of at least 400 bison.
Issues and challenges
The existence of bovine tuberculosis and brucellosis in herds in and near WBNP remains one of the most significant threats to recovery of healthy populations of wood bison in parts of northern Canada (Gates et al. 1992). The risk of infection of the Mackenzie and Hay-Zama herds is currently being studied. This assessment should provide insight into ways to mitigate disease risk for wild and domestic herds of bison in the region surrounding WBNP (Mitchell and Gates 1999). Wood bison were protected in a special wildlife management area in northwestern Alberta in 1993. Recovery outside of this area is limited not only by the presence of herds infected by tuberculosis and brucellosis in and near WBNP, but by policies and legislation that exclude bison from management as wildlife in the remainder of the province.
The expansion of resource extraction activities, forestry, and associated human access in the boreal forest in northern Alberta and British Columbia presents challenges for wood bison recovery in these areas. Although habitat loss and degradation and increased hunting pressure may result from these activities, the potential also exists for positive effects. For example, Syncrude Canada Ltd. (northeastern Alberta) established a captive-breeding herd on reconstructed grasslands. British Columbia prepared a management plan for wood bison that outlines actions to restore them to areas in the extreme northeastern section of the province (Harper and Gates 1999; Harper et al. 2000). Forage species planted along exploration right of ways provides habitat for wood bison in parts of this region.
The British Columbia plan identifies the presence of game-farmed and free-roaming plains bison within the historic range of the wood bison as the greatest obstacle to successful implementation of the plan. Bison ranching is growing rapidly in northern British Columbia and Alberta, both in terms of the number of captive bison and the number of ranches. The existence of free-ranging wood bison near bison ranches would result in conflict with commercial operations. Wildlife managers and the bison ranching industry should address this issue jointly.
In view of the limited availability of habitat for wood bison in Canada, and the risk of infection of some Canadian herds with cattle diseases, opportunities for reestablishing herds in unoccupied suitable habitat is a priority. Such areas may exist in northeastern British Columbia, in central to northern Yukon, and along the lower Mackenzie Valley in the Northwest Territories.
Another challenge facing wood bison recovery is the conservation of genetic resources. The current national captive breeding herd at Elk Island National Park, although free of tuberculosis and brucellosis, is genetically less diverse than other wood bison populations, including the parent population in Wood Buffalo National Park (Wilson and Strobeck 1999). Recent studies of the Hook Lake captive breeding herd indicate that a significant part of the genetic variability of the parent population in the Slave River Lowlands is represented in the captive herd, making it the most genetically diverse captive wood bison herd (J. Nishi, pers. comm.). Incorporating this genetic diversity in the national wood bison recovery program is desirable. However, precautions are necessary to avoid translocation of tuberculosis and brucellosis.
Proposed reintroduction to Alaska
The Alaska Department of Fish and Game (ADF&G) first considered the possibility of reintroducing wood bison to the Yukon Flats in 1991. The concept was discussed with local residents and landowners. Conservation authorities in Canada responded positively to the idea of supporting a reintroduction to Alaska. As described earlier, several Athabascan elders have provided oral history describing the presence of bison, and their being hunted, prior to their disappearance during the last few hundred years.
A preliminary habitat assessment of the Yukon Flats in 1992 (Gates 1992) indicated there was substantial habitat for wood bison on the Yukon Flats in northeastern Alaska. ADF&G completed a Feasibility Assessment in 1994 (Alaska Department of Fish and Game 1994) and a Habitat Assessment in 1995 (Berger et al. 1995). These studies concluded that a reintroduction is feasible and that the Yukon Flats would provide high quality habitat for a substantial population of wood bison. The area supports extensive meadow systems with plant communities that are nearly identical to those occurring in the best wood bison range in Canada. The project has been discussed in numerous formal and informal meetings with local communities and village councils, the Alaska Board of Game, Fish and Game Advisory Committees, conservation and environmental groups, and various civic organizations. The response has been largely supportive among both local and non-local residents. Alaskan and Canadian support for the project stems from interest in restoring this historically important animal and contributing to wood bison conservation, as well as the variety of other benefits associated with wood bison. These include considerations related to Athabascan culture and history, esthetics, biological and habitat diversity, restoration of ecological processes, and the desire of local communities to enhance local economies based on renewable resources.
The Wood Bison Recovery Team places a high priority on the reintroduction of wood bison to Alaska (Wood Bison Recovery Team in press). Reintroducing wood bison to Alaska would be consistent with the international agreement "Framework for Cooperation Between Environment Canada and the U.S. Department of the Interior in the Protection and Recovery of Wild Species at Risk," which was signed in April 1997.
Issues and challenges
The major challenge to the reintroduction of wood bison to Alaska appears to be related to the fact that much of the information regarding the history of wood bison has emerged fairly recently. Although there is substantial support for the project, information about the history of wood bison in Alaska will help resolve lingering concerns about their status as a native animal.
Other issues that are being addressed include mechanisms for local involvement in implementing the reintroduction, public involvement in long term management of bison, harvest regimes that could provide for an eventual harvest allocation that would accommodate local interests while not excluding other user groups, establishing the population size at which limited harvest would begin, and mechanisms for cooperative management.
Proposed reintroduction in Siberia
History of the project
The proposal to test the feasibility of establishing a productive grazing ecosystem in the taiga is the product of collaboration between Russian, American and Canadian scientists. It would involve reestablishing an assemblage of large grazing herbivores, including wood bison and indigenous Siberian horses that would coexist with resident moose and caribou populations in a fenced area in northeastern Siberia. It is possible the Siberian tiger may eventually be introduced to fill the ecological role of the extinct Pleistocene lion. The large ungulates proposed for the park have been successfully reintroduced in other regions of the Holarctic. For example, bison and muskoxen have been successfully re-established in the Yukon, Northwest Territories and Alaska. Fifty Siberian horses were released at the project site in 1998.
The area designated for the project is located near Cherskii, on the Kolyma River, 100 km south of the Arctic Ocean. It is a 160-km2 scientific reserve, owned and administered by a non-profit corporation, the Pleistocene Park Association, consisting of six ecologists from the Northeast Science Station in Cherskii and the Grassland Institute in Yakutsk. The reserve is surrounded by a 600-km2 buffer zone that will be added to the park once several species have been reestablished.
The process of restoring some of the species that were extirpated, at least in part as a result of human activities, has already begun in parts of Siberia. In the last few decades sable, caribou, muskox and horses have been successfully restored to parts of their former range in eastern Siberia. Reestablishing bison in the region could be another step in an on-going effort to restore wildlife and ecosystems in the region, and is analogous to historic efforts to restore and conserve wildlife in North America. The Russian Federation and the Sakha Republic (Yakutia) support the project because of its potential to restore and protect the environment and because it may eventually provide socioeconomic benefits for the people of the north. Support for the project stated in the Canadian Wood Bison Recovery Plan (Wood Bison Recovery Team in press) and the Government of Canada has indicated its support. Protocol agreements have been prepared on animal health certification and donation of wood bison from Canada's captive breeding herd.
Issues and challenges
The Pleistocene Park project is an experiment designed to evaluate the effects of herbivory and other relationships between ungulates and the northern environment (Zimov et al. 1995). Although available evidence indicates that the most recent form of bison to occupy Siberia was morphologically similar to wood bison, additional work is required to clarify taxonomic and phylogenetic relationships between bison in North America and Siberia. Dr. A. Cooper with the ancient DNA lab at the University of Oxford has undertaken this study. The Wood Bison Recovery Team (in press) has recommended that the release of wood bison to the wild in Siberia be conditional on finding that the Siberian Holocene form is clearly related to the North American Holocene form.
Wood bison have achieved modest recovery in Canada and are no longer threatened by imminent extinction. Further recovery is limited by disease, habitat loss, and the distribution of commercial bison ranches. Reestablishing wood bison in additional areas in Canada, including the northern Yukon and the Lower Mackenzie Valley, in Alaska, and possibly in Siberia will enhance the global security of the subspecies. Reintroductions will contribute to ecological restoration in these regions and create opportunities for bison to play a role in the culture and economies of northern peoples as they did in the past.
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Fig. 1. Distribution of captive and free-roaming wood bison herds in Canada, and locations of a proposed reintroduction to Alaska and a proposed translocation to Siberia. Numbers refer to the following herds and locations. Approximate herd size is shown in parentheses.
1 Mackenzie (1908) 9 Nyarling (260)
2 Nahanni (160) 10 Peace Athabasca Delta (900)
3 Yukon (500) 11 Elk Island National Park (350)
4 Nordquist (50) 12 Syncrude Canada Ltd. (150)
5 Hay Zama (130) 13 Hook Lake recovery project (65)
6 Chitek Lake (70) 14 Yukon Flats (proposed)
7 Etthithun Lake (43) 15 Pleistocene Park (proposed)
8 SRL (600)
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