A Brief Review of the Status of Plains Bison in North America

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and was later repealed (MacEwan 1995).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

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

Boyd and Gates: ABrief Review of the Status of Plains Bison in North America JOW, Spring 2006, Vol. 45, No. 2 —17
TABLE 1. Plains Bison Conservation Herds in North America (Boyd 2003). Numbers refer to locations on the distribution map in Figure 1.
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.
5Yellowstone 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 HenryMountains 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 ParkCO 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

18 —JOW, Spring 2006, Vol. 45, No. 2 Boyd and Gates: ABrief Review of the Status of Plains Bison in North America
natural dispersal or are being managed actively for
expansion (Boyd 2003).
Genetics
Bison experienced a severe population decline in the
nineteenth century. Since then, they have undergone
artificial hybridization with cattle, been subject to do-
mestication, and have been separated into isolated
populations. All these factors could have affected the
integrity of the bison genome.
Genetic diversity: At the species level, genetic diver-
sity provides the mechanism for evolutionary change
and adaptation (Allendorf and Leary 1986; Meffe and
Carroll 1994; Chambers 1998). Reduction in genetic di-
versity can result in reduced fitness, diminished growth,
increased mortality, and shrinking evolutionary flexibil-
ity of individuals within a population (Ballou and Ralls
1982; Mitton and Grant 1984; Allendorf and Leary
1986; Berger and Cunningham 1994). North American
bison approached extinction in the late 1800s and expe-
rienced a severe demographic bottleneck. Consequently,
extant bison populations may have lower genetic diver-
sity compared to pre-decline populations. Inventories of
genetic diversity held in conservation herds and studies
of herd population dynamics are needed to develop
genetic management plans for North American bison.
Hybridization: The concept of crossing bison with
domestic cattle dates back to Spanish colonizers of the
sixteenth century (Dary 1989). Cross-breeding was at-
tempted in Virginia, the Carolinas, and Pennsylvania
during the 1700s (Ogilvie 1979). In 1888, C. J. “Buf-
falo” Jones coined the term catalo to refer to hybrids
between cattle and bison. Private ranchers involved with
salvaging bison had aspirations of combining the hardi-
ness and winter foraging ability of bison with the meat
production traits of cattle through hybridization (Ogilvie
1979; Dary1989). The Canadian government pursued
experimental production of crossbred animals from
1916-1964 (Ogilvie 1979; Polziehn et al. 1995).
Historical cross-breeding attempts havecreated a
legacy of genetic issues related to the introgression of
cattle DNA into bison herds. Introgression refers to gene
flowbetween populations caused by hybridization fol-
lowed by backbreeding of the hybrid offspring to their
respective parental populations (Rhymer and Simber-
loff1996). The introgressed DNA displaces sections of
the original genome, thereby affecting the genetic in-
tegrity of a species. Many contemporary bison herds are
founded on, and supplemented with, animals from herds
with a history of hybridization. Seven of fifty conser-
vation herds currently show evidence of cattle DNA in-
trogression (Ward et al. 1999; Ward 2000). There is a
high percentage of untested herds (68 percent), creating
alarge information gap in understanding hybridization
prevalence among plains bison conservation herds
(Boyd 2003). Plains bison herds with no evidence of
hybrids include all five U.S. National Park herds, two
of five U.S. National Wildlife Refuge herds, the state-
managed Henry Mountains herd in Utah, and the Elk
Island National Park herd in Canada. These herds ac-
count for approximately 7,984 bison, or 42 percent of
the total estimated plains bison in conservation popula-
tions (Boyd 2003).
Domestication: The commercial bison population in
North America is at least 500,000 and growing. Ranch-
ers continue to enter the bison industry to capitalize on
economic opportunities afforded by bison. The increase
in commercial bison production may reflect recognition
of advantages afforded by the adaptations and ecological
efficiency of bison as an indigenous range animal. Bison
possess several traits that make them preferable to cattle
as a range animal, including greater ability to digest low
quality forage (Hawley et al. 1981), to defend against
predators (Carbyn et al. 1993), and lower incidence of
calving difficulties (Haigh et al. 2001). The primary
goal of many commercial bison ranchers is to increase
profit by maximizing calf production, feed-to-meat con-
version efficiency, and meat quality (Schneider 1998).
This requires nonrandom selection for traits that serve
this purpose, including conformation, docility, reduced
agility, growth performance, and carcass composition.
Selection for these traits reduces genetic variation and
changes the character of the animal over time (Hodgson
1994). The demand for bison meat cannot currently
compete with the much larger scale of the beef produc-
tion industry. Therefore, many bison producers apply
cattle husbandry practices and standards to bison; stan-
dards that may be practical for the bison business, but
will not maintain the bison genome.
Disease
Only one disease, bovine brucellosis, is of concern
for plains bison conservation (Boyd 2003). Bovine bru-
cellosis, also known as Bang’sdisease,is caused by
infection with the bacterium Brucella abortus (Tessaro
1992). Current evidence suggests that brucellosis was
introduced to North America from Europe during the
1500s (Aguirre and Starkey 1994; Meagher and Mayer
1994). The disease is primarily transmitted through oral
contact with aborted fetuses, contaminated placentas,
and uterine discharges (Tessaro 1992). The impacts of
brucellosis on female bison are abortion, inflammation
of the uterus, and retained placenta (Tessaro1992). Male
bison experience inflammation of the seminal vessels,
testicles, and epididymis, and in advanced cases, steril-
ity (Tessaro 1992). Both sexes aresusceptible to bursitis
and arthritis caused by concentrations of the organism in
joints, resulting in lameness, and possibly increased
vulnerability to predation (Tessaro 1992). There is cur-
rently no fully effective vaccine for preventing bovine
brucellosis (Cheville et al. 1998); however, there is
active research testing the efficacy and biosafety of new
vaccines (Olsen et al. 1998; Roffe et al. 1999).
Two of fifty plains bison conservation herds in North

Boyd and Gates: ABrief Review of the Status of Plains Bison in North America JOW, Spring 2006, Vol. 45, No. 2 —19
America are chronically infected with brucellosis: YNP
and the Jackson herd in Grand Teton National Park/
National Elk Refuge. These populations account for
4,700 bison, or 24 percent of the North American plains
bison conservation population (Boyd 2003). Manage-
ment of these herds is affected by the presence of bru-
cellosis because of the potential risk the disease poses to
the livestock industry. Transmission of brucellosis from
bison to cattle has been demonstrated in captive studies;
however,there are no confirmed cases of transmission in
the wild (Cheville et al. 1998; Shaw and Meagher 2000;
Bienen 2002). Nevertheless, the potential exists and has
created a contentious bison management issue in the
region.
Legal Issues
Plains bison are currently not recognized at the sub-
specific level on any national or international list for
species at risk of extinction. A recent survey of plains
bison conservation status reveals trends demonstrating
that plains bison warrant consideration for listing (Boyd
2003). Although the North American plains bison popu-
lation is over 500,000 and is growing, more than 95 per-
cent of herds are under commercial production. Of the
estimated 19,200 plains bison in conservation herds,
only 8,337 are free-ranging (Boyd 2003). Further, free-
ranging, brucellosis-free populations within original
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 gene-
tic diversity and hybridization
with domestic cattle further sup-
port consideration of the plains
bison for listing.
Potential complications could
accompany the process of list-
ing plains bison. First, the pres-
ence of cattle DNA in some
conservation herds may pre-
clude listing under some legis-
lation, 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 hybrid-
ization histories could therefore
be exempt from the ESA. Sec-
ond, if all plains bison are
considered, then the growing
commercial population pre-
cludes any arguments for listing
based on numerical status. Third, legislation supporting
listings may prohibit commercial and captive propaga-
tion of a listed species; a situation that the current
momentum of the bison industry would not allow. A
legal distinction between wild and domesticated popu-
lations would be required to support protection of the
wild form(Boyd 2003).
Legal recognition of the wild form is impeded by the
classification of bison as livestock by many state and
provincial governments. In the absence of protection by
wildlife legislation, free-ranging plains bison could
potentiallybe sequestered into private herds or hunted
without regulation. The only legal protection afforded to
free-ranging bison in this situation would be associated
with the legal status of their habitat (e.g., a national
park). Classification as non-wildlifecould have implica-
tions for the success of attempts to reintroduce wild
bison herds. Existing free-ranging plains bison herds in
Alaska, Arizona, Montana, Utah, Wyoming, British
Columbia, and Saskatchewan are managed as wildlife
under state or provincial legislation. Legislative revi-
sions would be required to provide for reintroductions of
free-ranging plains bison in other jurisdictions.
RecoveryInitiatives
The ultimate goal of bison conservation is to facilitate
recovery of the species and ensure long-term survival of
FIGURE 2: Great Plains States Counties that lost population between 1990-2000.

20 —JOW, Spring 2006, Vol. 45, No. 2 Boyd and Gates: ABrief Review of the Status of Plains Bison in North America
bison as a wild species (Boyd 2003). Bison recovery
should include the maintenance and reintroduction of
free-ranging or minimally-managed captive herds in
areas within the taxon’s original range. To maximize
conservation value, these herds should occupy large
geographic areas, and should be of sufficient size and
demographic composition to maintain population via-
bility. The herds should be subject to forces of natural
selection, and effective genetic, disease, and range man-
agement. They should also be protected under law and
free of the previous causes of extirpation.
In Canada, the national parks played a pivotal role in
rescuing the bison from extinction through interventions
such as purchasing the Pablo-Allard herd and providing
sanctuary for the struggling species (Ogilvie 1979).
Today, 50 percent of conservation herds in Canada are
found in national parks. Protected public lands offer the
best sanctuary for plains bison in Canada at this time.
There are also emerging opportunities for bison restora-
tion on additional public lands and lands owned or man-
aged by conservation organizations. In Canada, plains
bison reintroductions are being considered for Banff
National Park, Waterton Lakes National Park, Grass-
lands National Park, and the Nature Conservancy of
Canada’s Old-Man-On-His-Back Conservation Area
(Boyd 2003).
Plans to reintroduce plains bison populations in the
United States are limited. The USDA Forest Service
recently conducted an assessment of its management of
national grasslands in Montana, North Dakota, Nebras-
ka, South Dakota, and Wyoming; it dismissed a pro-
posed alternative to restore free-ranging bison (USDA
Forest Service 2001). Two appeals to this decision were
being considered in 2002 by the Chief of the Forest
Service (J. Kessler, Biodiversity Conservation Alliance,
personal communication 2002). There are, however, two
landscape-level grassland restoration projects being
planned that involve reintroducing bison: The Montana
Big Open and The Buffalo Commons’Million Acre Pro-
ject. Although ambitious, these large-scale grassland
restoration concepts consider landscape-level processes,
and focus on multi-stakeholder networks to facilitate
positiveeconomic and environmental rejuvenation of
the prairie region, and reestablish free-ranging bison on
the North American landscape.
ACKNOWLEDGEMENT
This paper is based in part on a conservation status survey for
bison in North America in preparation by the World Conservation
Union/Species Survival Commission’s Bison Specialist Group
(BSG). The authors would like to acknowledge and thank the mem-
bers of the BSG for their support and feedback throughout the devel-
opment of the status survey.
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Delaney P. Boyd graduated from the University of Calgary’s
Faculty of Environmental Design Environmental Science pro-
gram 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 experi-
ence 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 con-
servation 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.