A Brief Review of the Status of Plains Bison in North America JOW, Spring 2006, Vol

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Boyd and Gates: A Brief Review of the Status of Plains Bison in North America JOW, Spring 2006, Vol. 45, No. 2 — 15 A Brief Review of the Status of Plains Bison in North America

Delaney P. Boyd and C. Cormack Gates

N 1879, for the first time in the post-glacial history cutting timber, or killing game in YNP (Dary 1989; of the Great Plains, the bison hunt failed. It hap- Danz 1997). I pened first in Canada, and then was repeated two In Canada, official protection of bison began in 1877 years later in Montana (Foster 1992). Once numbering with the passing of An Ordinance for the Protection of in the tens of millions, the plains bison (Bison bison Buffalo (Ogilvie 1979; Gates et al. 2001). This act was bison) was driven to near extinction during a brief frenzy nearly impossible to enforce by the small regiments of of exploitation. Commercial hunting for meat and hides Northwest Mounted Police scattered across the plains, by Native Americans and Euro-Americans was the prox- and was later repealed (MacEwan 1995). In 1883, the imal cause of the decline (Hornaday 1889; Isenberg Ordinance for the Protection of Game was passed, but it 2000). Other contributing factors included subsistence was also ineffective (Ogilvie 1979). The government of hunting, indiscriminate slaughter for sport, transection the Dominion of Canada enacted the Unorganized of the plains by railroads, and political motivations (He- Territories Game Preservation Act in 1894, in part as a witt 1919; Mayer and Roth 1958; Dary 1989; Geist response to reports that the wood bison (B. b. athabas- 1996; Danz 1997). Environmental factors such as re- cae) population in northern Canada had declined to 250 gional drought, introduced bovine diseases, and compe- (Ogilvie 1979). Enforcement of this legislation was tition from domestic livestock and domestic and wild minimal, however, until the Northwest Mounted Police horses also played a role (Flores 1991; Isenberg 2000). was given an enforcement mandate in 1897 (Soper The early conservation of plains bison was made pos- 1941). sible through the independent actions of private citizens Despite protective legislation, prior to 1907 plains combined with protective legislation. Prominent leaders bison in Canada were reduced to a small herd in Rocky included James McKay and Charles Alloway (Manito- Mountains Park (Banff) and a few animals near Win- ba), Charles Goodnight (Texas), Walking Coyote (Mon- nipeg (Ogilvie 1979). In 1907, the Dominion of Canada tana), Frederick Dupree (South Dakota), Charles J. purchased the entire Pablo-Allard herd (approximately “Buffalo” Jones (Kansas), and Michel Pablo and 716 plains bison) from Michel Pablo of Montana. Four Charles Allard (Montana) (Coder 1975; Dary 1989). hundred and ten of these bison were temporarily held at Their efforts to establish breeding herds from the few re- Elk Island National Park (EINP) near Edmonton, Al- maining bison resulted in preservation of foundation berta until most were transported to Wainwright Buffalo stock for much of the subsequent recovery of the sub- Park in east-central Alberta; forty-eight bison were left species. at EINP, forming the nucleus of that national park herd In the United States, numerous bills to protect bison (Coder 1975; Fuller 2002). The Pablo-Allard bison were were introduced by members of Congress between 1871 subsequently used to establish herds in other Canadian and 1876, but no laws were enacted (Ogilvie 1979; Dary national parks. 1989; Danz 1997). Several state and territorial govern- Founded in 1905, the American Bison Society ments were able to enact legislation to protect bison pressed Congress to establish several public bison herds during the last three decades of the 1800s; however, including Wichita Mountains National Wildlife Refuge, these laws were largely ineffective and unenforceable the National Bison Range, Sullys Hill National Game (Danz 1997). In 1872, President Grant created Yellow- Preserve, and Fort Niobrara National Wildlife Refuge stone National Park (YNP) to protect all natural re- (Coder 1975). National parks in both the United States sources, including bison, within its borders. By 1894, and Canada were also established in part to conserve however, poaching had reduced the park bison popula- bison and other large mammal populations (Ogilvie tion from 200 to only 25 animals (Danz 1997). On May 1979). 7, 1894, President Cleveland signed the National Park Protective Act (Lacey Act), ameliorating the longstand- Current Numerical and Geographic Status ing problem with jurisdiction and law enforcement in Following protection from hunting, bison populations YNP. This was the first U.S. federal law to provide spe- increased rapidly, doubling in number between 1888 and cific protection for bison. It carried a two-year jail term 1902; by 1909, the plains bison was considered safe and a $1,000 fine for anyone removing mineral deposits, from extinction (Coder 1975). Initially sparked by 16 — JOW, Spring 2006, Vol. 45, No. 2 Boyd and Gates: A Brief Review of the Status of Plains Bison in North America

FIGURE 1: Distribution of plains bison conservation herds in North America. Numbers correspond with the herd list in Table 1. The shaded area represents original plains bison range. reverence for the animal and nostalgia, motivations for Conservation Issues bison conservation and recovery became increasingly The most important conservation-related issues es- driven by the animal’s commercial value (Yorks and sential to the survival and increase of bison herds are Capels 1998). By 1970, there were 30,000 plains bison habitat, population genetics, disease, and legal issues. in North America; approximately half were in public herds located in national parks, wildlife refuges, and Habitat state wildlife areas, and the other half were privately The plains bison was originally a land-intensive, no- held (Shaw and Meagher 2000). madic species that roamed over great distances on the A recent survey of the numerical status of plains North American landscape. Large-bodied animals are bison estimated that there are over 500,000 bison in especially vulnerable to the effects of habitat fragmen- North America including both commercial and conser- tation because they require a large amount of space vation populations (Boyd 2003). Of these, 95 percent are (Berger and Cunningham 1994). Fragmented popula- under commercial production. Conservation herds enu- tions can be more susceptible to inbreeding pressures, merated in the survey included those managed by mu- loss of genetic diversity, and extinction (Berger and nicipal, state, provincial, and federal governments, and Cunningham 1994). Conservation of plains bison is lim- private organizations having clear conservation objec- ited because most of the original range has experienced tives. There are fifty plains bison conservation herds in change from competing land uses including cultivation, North America (Table 1), 32 percent of which have less cattle ranching, commercial bison ranching, natural re- than fifty bison (Boyd 2003). Thirteen herds have popu- source extraction, and urban expansion (Johnson et al. lations greater than 400 (Boyd 2003), the number cur- 1994). These land uses constrain the potential of pre- rently estimated as the minimum viable population for serving or restoring large tracts of habitat for bison bison (Gates et al. 2001). The number of plains bison in conservation. conservation herds is estimated at 19,200 with 90 per- Current plains bison conservation herds are widely cent in the United States, 10 percent in Canada, and scattered and isolated across the original range of the none in Mexico (Boyd 2003). Only 22 percent of plains subspecies (Figure 1). Thirty-eight percent of conserva- bison conservation herds are increasing in size. Most tion herds reside on ranges smaller than 10 km2; 60 per- herds are within original plains bison range (Figure 1). cent have ranges smaller than 100 km2 (Boyd 2003). Eight of the fifty herds are distinctly outside plains bison There is no range expansion potential for 52 percent of range (Arizona, California, northern British Columbia, plains bison conservation herds. Of herds with expan- and Alaska) (Boyd 2003). sion potential, only eleven are currently expanding by Boyd and Gates: A Brief Review of the Status of Plains Bison in North America JOW, Spring 2006, Vol. 45, No. 2 — 17

Herd Location Jurisdiction Managing Authority Population

UNITED STATES 1 Badlands National Park SD Federal U.S. National Parks Service 750 2 Theodore Roosevelt National Park ND Federal U.S. National Parks Service 850 3 Wind Cave National Park SD Federal U.S. National Parks Service 375 4 Grand Teton National Park/Nat. Elk Refuge WY Federal and State U.S. NPS; U.S. FWS; WY 700 Fish and Game Dept. 5 Yellowstone National Park WY/MT Federal and State U.S. NPS, NFS, MT Fish and Parks, 4,000 and MT Dept. of Livestock 6 Fort Niobrara National Wildlife Refuge NE Federal U.S. Fish and Wildlife Service 352 7 National Bison Range MT Federal U.S. Fish and Wildlife Service 400 8 Neal Smith National Wildlife Refuge IA Federal U.S. Fish and Wildlife Service 35 9 Sullys Hill National Game Preserve ND Federal U.S. Fish and Wildlife Service 37 10 Wichita Mountains National Wildlife Refuge OK Federal U.S. Fish and Wildlife Service 565 11 Fermilab National Accelerator IL Federal Department of Energy 32 12 Land Between the Lakes National Rec. Area KY Federal USDA Forest Service 130 13 Chitina AK State Alaska Department of Fish and Game 38 14 Copper River AK State Alaska Department of Fish and Game 108 15 Delta Junction AK State Alaska Department of Fish and Game 360 16 Farewell Lake AK State Alaska Department of Fish and Game 400 17 House Rock State Wildlife Area AZ State Arizona Fish and Game Department 217 18 Raymond State Wildlife Area AZ State Arizona Fish and Game Department 72 19 Antelope Island State Park UT State Department of Natural Resources, 600 Division of Parks and Recreation 20 Blue Mounds State Park MN State Department of Natural Resources, 56 Division of Parks and Recreation 21 Finney Game Refuge KS State Kansas Dept. of Wildlife and Parks 120 22 Maxwell Wildlife Refuge KS State Kansas Dept. of Wildlife and Parks 230 23 Prairie State Park MO State Missouri Dept. of Natural Resources 76 24 Fort Robinson State Park NE State Nebraska Game and Parks 500 25 Wildcat Hills State Recreation Area NE State Nebraska Game and Parks 10 26 Custer State Park SD State South Dakota Game Fish & Parks Dept. 1,100 27 Caprock Canyons State Park TX State Texas Parks and Wildlife Department 40 28 Henry Mountains UT State Utah Division of Wildlife Resources 279 29 Sandhill Wildlife Area WI State Wisconsin Dept. of Natural Resources 15 30 Bear River State Park WY State Wyoming State Parks and Historic Sites 8 31 Hot Springs State Park WY State Wyoming State Parks and Historic Sites 11 32 Konza Prairie Biological Station KS State and K-State University, Division of Biology; 275 Foundation The Nature Conservancy 33 Santa Catalina Island CA Foundation Catalina Island Conservancy 225 34 Cross Ranch Nature Preserve ND Foundation The Nature Conservancy 140 35 Medano-Zapata Ranch CO Foundation The Nature Conservancy 1,500 36 Niobrara Valley Preserve NE Foundation The Nature Conservancy 473 37 Ordway Prairie Preserve SD Foundation The Nature Conservancy 255 38 Tallgrass Prairie Preserve OK Foundation The Nature Conservancy 1,500 39 Clymer Meadow Preserve TX Foundation and The Nature Conservancy; Private rancher 320 Private 40 Smoky Valley Ranch KS Foundation The Nature Conservancy 45 41 Daniels Park CO Municipal Denver Parks and Recreation 26 42 Genesee Park CO Municipal Denver Parks and Recreation 26 Subtotal – United States 17,251 CANADA 43 Camp Wainwright AB Federal Department of National Defence 16 44 Elk Island National Park AB Federal Parks Canada Agency 430 45 Prince Albert National Park SK Federal Parks Canada Agency 310 46 Riding Mountain National Park MB Federal Parks Canada Agency 33 47 Waterton Lakes National Park AB Federal Parks Canada Agency 27 48 Primrose Lake Air Weapons Range AB/SK Federal and Dept. of Nat’l Defence; SK Environment, 100 (Cold Lake) Provincial Fish and Wildlife Branch 49 Pink Mountain BC Provincial British Columbia Department of Water, 1,000 Lands and Air Protection 50 Buffalo Pound Provincial Park SK Provincial Saskatchewan Environment, Parks Branch 33 Subtotal – Canada 1,949 TOTAL – NORTH AMERICA 19,200

TABLE 1. Plains Bison Conservation Herds in North America (Boyd 2003). Numbers refer to locations on the distribution map in Figure 1. 18 — JOW, Spring 2006, Vol. 45, No. 2 Boyd and Gates: A Brief Review of the Status of Plains Bison in North America natural dispersal or are being managed actively for of five U.S. National Wildlife Refuge herds, the state- expansion (Boyd 2003). managed Henry Mountains herd in Utah, and the Elk Island National Park herd in Canada. These herds ac- Genetics count for approximately 7,984 bison, or 42 percent of Bison experienced a severe population decline in the the total estimated plains bison in conservation popula- nineteenth century. Since then, they have undergone tions (Boyd 2003). artificial hybridization with cattle, been subject to do- Domestication: The commercial bison population in mestication, and have been separated into isolated North America is at least 500,000 and growing. Ranch- populations. All these factors could have affected the ers continue to enter the bison industry to capitalize on integrity of the bison genome. economic opportunities afforded by bison. The increase Genetic diversity: At the species level, genetic diver- in commercial bison production may reflect recognition sity provides the mechanism for evolutionary change of advantages afforded by the adaptations and ecological and adaptation (Allendorf and Leary 1986; Meffe and efficiency of bison as an indigenous range animal. Bison Carroll 1994; Chambers 1998). Reduction in genetic di- possess several traits that make them preferable to cattle versity can result in reduced fitness, diminished growth, as a range animal, including greater ability to digest low increased mortality, and shrinking evolutionary flexibil- quality forage (Hawley et al. 1981), to defend against ity of individuals within a population (Ballou and Ralls predators (Carbyn et al. 1993), and lower incidence of 1982; Mitton and Grant 1984; Allendorf and Leary calving difficulties (Haigh et al. 2001). The primary 1986; Berger and Cunningham 1994). North American goal of many commercial bison ranchers is to increase bison approached extinction in the late 1800s and expe- profit by maximizing calf production, feed-to-meat con- rienced a severe demographic bottleneck. Consequently, version efficiency, and meat quality (Schneider 1998). extant bison populations may have lower genetic diver- This requires nonrandom selection for traits that serve sity compared to pre-decline populations. Inventories of this purpose, including conformation, docility, reduced genetic diversity held in conservation herds and studies agility, growth performance, and carcass composition. of herd population dynamics are needed to develop Selection for these traits reduces genetic variation and genetic management plans for North American bison. changes the character of the animal over time (Hodgson Hybridization: The concept of crossing bison with 1994). The demand for bison meat cannot currently domestic cattle dates back to Spanish colonizers of the compete with the much larger scale of the beef produc- sixteenth century (Dary 1989). Cross-breeding was at- tion industry. Therefore, many bison producers apply tempted in Virginia, the Carolinas, and Pennsylvania cattle husbandry practices and standards to bison; stan- during the 1700s (Ogilvie 1979). In 1888, C. J. “Buf- dards that may be practical for the bison business, but falo” Jones coined the term catalo to refer to hybrids will not maintain the bison genome. between cattle and bison. Private ranchers involved with salvaging bison had aspirations of combining the hardi- Disease ness and winter foraging ability of bison with the meat Only one disease, bovine brucellosis, is of concern production traits of cattle through hybridization (Ogilvie for plains bison conservation (Boyd 2003). Bovine bru- 1979; Dary 1989). The Canadian government pursued cellosis, also known as Bang’s disease, is caused by experimental production of crossbred animals from infection with the bacterium Brucella abortus (Tessaro 1916-1964 (Ogilvie 1979; Polziehn et al. 1995). 1992). Current evidence suggests that brucellosis was Historical cross-breeding attempts have created a introduced to North America from Europe during the legacy of genetic issues related to the introgression of 1500s (Aguirre and Starkey 1994; Meagher and Mayer cattle DNA into bison herds. Introgression refers to gene 1994). The disease is primarily transmitted through oral flow between populations caused by hybridization fol- contact with aborted fetuses, contaminated placentas, lowed by backbreeding of the hybrid offspring to their and uterine discharges (Tessaro 1992). The impacts of respective parental populations (Rhymer and Simber- brucellosis on female bison are abortion, inflammation loff 1996). The introgressed DNA displaces sections of of the uterus, and retained placenta (Tessaro 1992). Male the original genome, thereby affecting the genetic in- bison experience inflammation of the seminal vessels, tegrity of a species. Many contemporary bison herds are testicles, and epididymis, and in advanced cases, steril- founded on, and supplemented with, animals from herds ity (Tessaro 1992). Both sexes are susceptible to bursitis with a history of hybridization. Seven of fifty conser- and arthritis caused by concentrations of the organism in vation herds currently show evidence of cattle DNA in- joints, resulting in lameness, and possibly increased trogression (Ward et al. 1999; Ward 2000). There is a vulnerability to predation (Tessaro 1992). There is cur- high percentage of untested herds (68 percent), creating rently no fully effective vaccine for preventing bovine a large information gap in understanding hybridization brucellosis (Cheville et al. 1998); however, there is prevalence among plains bison conservation herds active research testing the efficacy and biosafety of new (Boyd 2003). Plains bison herds with no evidence of vaccines (Olsen et al. 1998; Roffe et al. 1999). hybrids include all five U.S. National Park herds, two Two of fifty plains bison conservation herds in North Boyd and Gates: A Brief Review of the Status of Plains Bison in North America JOW, Spring 2006, Vol. 45, No. 2 — 19

plains bison range account for only 1,289 plains bison, or 6.7 percent of the total conservation population (Boyd 2003). Con- servation issues related to genetic diversity and hybridization with domestic cattle further support consideration of the plains bison for listing. Potential complications could accompany the process of listing plains bison. First, the presence of cattle DNA in some conservation herds may pre- clude listing under some legislation, such as the United States Endangered Species Act. Hy- brids are exempt from the ESA when propagated in captivity and when they result from one listed parent and one nonlisted parent (O’Brien and Mayr 1991; U.S. Fish and Wildlife Service 2003). Plains bison with hybridization histories could therefore be exempt from the ESA. Sec- ond, if all plains bison are considered, then the growing FIGURE 2: Great Plains States Counties that lost population between 1990-2000. commercial population pre- cludes any arguments for listing America are chronically infected with brucellosis: YNP based on numerical status. Third, legislation supporting and the Jackson herd in Grand Teton National Park/ listings may prohibit commercial and captive propaga- National Elk Refuge. These populations account for tion of a listed species; a situation that the current 4,700 bison, or 24 percent of the North American plains momentum of the bison industry would not allow. A bison conservation population (Boyd 2003). Manage- legal distinction between wild and domesticated popu- ment of these herds is affected by the presence of bru- lations would be required to support protection of the cellosis because of the potential risk the disease poses to wild form (Boyd 2003). the livestock industry. Transmission of brucellosis from Legal recognition of the wild form is impeded by the bison to cattle has been demonstrated in captive studies; classification of bison as livestock by many state and however, there are no confirmed cases of transmission in provincial governments. In the absence of protection by the wild (Cheville et al. 1998; Shaw and Meagher 2000; wildlife legislation, free-ranging plains bison could Bienen 2002). Nevertheless, the potential exists and has potentially be sequestered into private herds or hunted created a contentious bison management issue in the without regulation. The only legal protection afforded to region. free-ranging bison in this situation would be associated with the legal status of their habitat (e.g., a national Legal Issues park). Classification as non-wildlife could have implica- Plains bison are currently not recognized at the sub- tions for the success of attempts to reintroduce wild specific level on any national or international list for bison herds. Existing free-ranging plains bison herds in species at risk of extinction. A recent survey of plains Alaska, Arizona, Montana, Utah, Wyoming, British bison conservation status reveals trends demonstrating Columbia, and Saskatchewan are managed as wildlife that plains bison warrant consideration for listing (Boyd under state or provincial legislation. Legislative revi- 2003). Although the North American plains bison popu- sions would be required to provide for reintroductions of lation is over 500,000 and is growing, more than 95 per- free-ranging plains bison in other jurisdictions. cent of herds are under commercial production. Of the estimated 19,200 plains bison in conservation herds, Recovery Initiatives only 8,337 are free-ranging (Boyd 2003). Further, free- The ultimate goal of bison conservation is to facilitate ranging, brucellosis-free populations within original recovery of the species and ensure long-term survival of 20 — JOW, Spring 2006, Vol. 45, No. 2 Boyd and Gates: A Brief Review of the Status of Plains Bison in North America bison as a wild species (Boyd 2003). Bison recovery MA: Sinauer, 1986), 57-76. Ballou, J., and K. Ralls, “Inbreeding and Juvenile Mortality in Small should include the maintenance and reintroduction of Populations of Ungulates: A Detailed Analysis,” Biological Con- free-ranging or minimally-managed captive herds in servation, 24 (1982): 239-272. areas within the taxon’s original range. To maximize Berger, J., and C. Cunningham, Bison: Mating and Conservation in Small Populations. (New York, NY: Columbia University, 1994). conservation value, these herds should occupy large Bienen, L., “Informed Decisions: Conservation Corridors and the geographic areas, and should be of sufficient size and Spread of Infectious Disease,” Conservation in Practice 3, 2 demographic composition to maintain population via- (2002): 10-17. Boyd, D. P., Conservation of North American Bison: Status and Rec- bility. The herds should be subject to forces of natural ommendations. (Calgary, Alberta: Master’s dissertation, Universi- selection, and effective genetic, disease, and range man- ty of Calgary, 2003). agement. They should also be protected under law and Carbyn, L. N., S. M. Oosenbrug, and D. W. Anions, “Wolves, Bison and the Dynamics Related to the Peace-Athabasca Delta in Cana- free of the previous causes of extirpation. da’s Wood Buffalo National Park,” Circumpolar Research Series, In Canada, the national parks played a pivotal role in 4 (1993): Can. Circumpolar Institute. rescuing the bison from extinction through interventions Chambers, K. E., “Using Genetic Data in the Management of Bison Herds,” in International Symposium on Bison Ecology and Man- such as purchasing the Pablo-Allard herd and providing agement in North America, June 4-7, 1997, edited by L. R. Irby sanctuary for the struggling species (Ogilvie 1979). and J. E. Knight (Bozeman, MT: Montana State University, 1998), Today, 50 percent of conservation herds in Canada are 151-157. Cheville, N. F., D. R. McCullough, and L. R. Paulson, Brucellosis in found in national parks. Protected public lands offer the the Greater Yellowstone Area. (Washington, DC: National Acade- best sanctuary for plains bison in Canada at this time. my of Sciences, 1998). There are also emerging opportunities for bison restora- Coder. G. D., The National Movement to Preserve the American Buf- falo in the United States and Canada between 1880 and 1920. tion on additional public lands and lands owned or man- (Columbus, OH: Ph.D. dissertation, Ohio State University, 1975). aged by conservation organizations. In Canada, plains Danz, H. P., Of Bison and Man (Niwot, CO: University Press of Col- bison reintroductions are being considered for Banff orado, 1997). Dary, D. A., The Buffalo Book, 2nd ed. (Athens, OH: Swallow /Ohio National Park, Waterton Lakes National Park, Grass- University, 1989). lands National Park, and the Nature Conservancy of Flores, D., “Bison Ecology and Bison Diplomacy: The Southern Canada’s Old-Man-On-His-Back Conservation Area Plains from 1800 to 1850” The Journal of American History, 78, 2 (1991): 465-485. (Boyd 2003). Foster, J. E., “The Metis and the End of the Plains Buffalo in Alber- Plans to reintroduce plains bison populations in the ta,” in Buffalo, edited by J. E. Foster, D. Harrison, and I. S. Ma- United States are limited. The USDA Forest Service cLaren (Edmonton, Alberta: University of Alberta, 1992), 61-77. Fuller, W. A., “Canada and the “Buffalo,” Bison bison: A Tale of Two recently conducted an assessment of its management of Herds,” Canadian Field Naturalist, 116, 1 (2002): 141-159. national grasslands in Montana, North Dakota, Nebras- Gates, C. C., R. O. Stephenson, H. W. Reynolds, C. G. van Zyll de ka, South Dakota, and Wyoming; it dismissed a pro- Jong, H. Schwantje, M. Hoefs, J. Nishi, N. Cool, J. Chisholm, A. James, and B. Koonz, National Recovery Plan for the Wood Bison posed alternative to restore free-ranging bison (USDA (Bison bison athabascae), National Recovery Plan No. 21. (Ot- Forest Service 2001). Two appeals to this decision were tawa, Ontario: Recovery of Nationally Endangered Wildlife, being considered in 2002 by the Chief of the Forest 2001). Geist, V., Buffalo Nation: History and Legend of the North American Service (J. Kessler, Biodiversity Conservation Alliance, Bison (Saskatoon, SK: Fifth House, 1996). personal communication 2002). There are, however, two Haigh, J., J. Berezowski, and R. Munger, “Reproduction and Herd landscape-level grassland restoration projects being Health,” in Bison Are Back — 2000. Proceedings of the Second In- ternational Bison Conference, August 2-4, 2000, edited by B. D. planned that involve reintroducing bison: The Montana Rutley (Leduc, Alberta: Bison Centre of Excellence, 2001), 154- Big Open and The Buffalo Commons’ Million Acre Pro- 173. ject. Although ambitious, these large-scale grassland Hawley, A. W. L., D. G. Peden, and W. R. Stricklin, “Bison and Here- ford Steer Digestion of Sedge Hay,” Canadian Journal of Animal restoration concepts consider landscape-level processes, Science, 61, 1 (1981): 165-174. and focus on multi-stakeholder networks to facilitate Hewitt, G. C., “The Coming Back of the Bison,” Natural History, 19, positive economic and environmental rejuvenation of 6 (1919): 553-565. Hodgson, B., “Buffalo: Back Home on the Range,” National Geo- the prairie region, and reestablish free-ranging bison on graphic, (November 1994): 67-87. the North American landscape. Hornaday, W. T., The Extermination of the American Bison, with a Sketch of Its Discovery and Life History: Annual Report (1887). ACKNOWLEDGEMENT (Washington, DC: Smithsonian Institution, 1889). This paper is based in part on a conservation status survey for Isenberg, A. C., The Destruction of the Bison: An Environmental His- bison in North America in preparation by the World Conservation tory, 1750-1920. (New York, NY: Cambridge University, 2000). Union/Species Survival Commission’s Bison Specialist Group Johnson, D. H., S. D. Haseltine, and L. M. Cowardin, “Wildlife Habi- (BSG). The authors would like to acknowledge and thank the mem- tat Management on the Northern Prairie Landscape,” Landscape bers of the BSG for their support and feedback throughout the devel- and Urban Planning, 28 (1994): 5-21. opment of the status survey. MacEwan, G., Buffalo: Sacred and Sacrificed (Edmonton, Alberta: Alberta Sport, Recreation, Parks and Wildlife Foundation, 1995). REFERENCES AND BIBLIOGRAPHY Mayer, F. H., and C. B. Roth, The Buffalo Harvest. (Union City, TN: Aguirre, A. A., and E. E. Starkey, “Wildlife Disease in U.S. National Pioneer Press, 1958, 2nd printing, 1995). Parks: Historical and Coevolutionary Perspectives,” Conservation Meagher, M., and M. E. Mayer, “On the Origin of Brucellosis in Biology 8, 3 (1994): 654-661. Bison of Yellowstone National Park: A Review,” Conservation Bi- Allendorf, F. W., and R. F. Leary, “Heterozygosity and Fitness in Nat- ology 8, 3 (1994): 645-653. ural Populations of Animals,” in Conservation Biology: The Sci- Meffe, G. K., and C. R. 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Delaney P. Boyd graduated from the University of Calgary’s Faculty of Environmental Design Environmental Science program with an M.E.Des. Her dissertation research involved de- veloping a conservation status survey on North American bison for the World Conservation Union/Species Survival Commis- sion’s (IUCN/BSG) Bison Specialist Group (BSG). She is an Officer of the IUCN/SSC Bison Specialist Group and manages the BSG website (www.notitia.com/bison). She also holds a B.Sc. in Ecology from the University of Calgary.

C. Cormack Gates holds a Ph.D. in Animal Science from the University of Alberta (1980). He is the Director of the Envi- ronmental Science Program in the Faculty of Environmental Design, University of Calgary. Dr. Gates has extensive experience in applied conservation science in Northern and Western Canada. He has facilitated work by a number of academic in- stitutions and government agencies on a diversity of applied conservation problems. Prior to taking a position at the Uni- versity of Calgary he spent many years working on the conservation of Canada’s bison and on research in support of management of northern terrestrial ecosystems. He is the prin- cipal author of Canada’s 2001 National Recovery Plan for Wood Bison and is the co-author of the Alberta species at risk status report for this animal. He chairs the IUCN Species Sur- vival Commission — Bison Specialist Group (North America), and is co-chair of Canada’s National Wood Bison Recovery Team. His research focuses on applied ecology and management of wildlife, species at risk of extinction, landscape ecology, and integrated resource and land use planning.