IVth International Wildlife Management Congress
Proceedings James W. Cain III and Jason P. Marshal, Editors
9-12 July 2012
Durban, South Africa Proceedings of the IVth International Wildlife Management Congress
Cooperative Wildlife Management Across Borders: Learning in the Face of Change
Editors James W. Cain III Jason P. Marshal
Copyright © 2013 by The Wildlife Society All rights reserved. No part of this book may be reproduced in any form, by photostat, microform, retrieval system, or by any other means, without the prior written permission of the publisher.
Distributed by The Wildife Society 5410 Grosvenor Lane Bethesda, MD 20814 ISBN 0- ISSN 0-
9-12 July 2012 Durban, South Africa Kohl et al. • Future of Bison Conservation
WHAT IS THE FUTURE OF BISON (n = 57) have <1,000 individuals, and 8% (n = 5) CONSERVATION? are managed on areas of >2,000 km2. Based on these data, new questions have been posed regarding the direction of bison restoration at the continental scale. MICHEL T. KOHL, Boone and Crockett Wildlife Taking into account challenges associated with bison Conservation Program, University of Montana, management (e.g., disease transmission, genetic intro- 32 Campus Drive, Missoula, MT 59812 USA gression of domestic cattle, legal designation of bison, availability of restoration sites), we examine whether JEROD A. MERKLE, Département de Biologie, the current model of numerous small, confined bison Université Laval, Pavillon Alexandre-Vachon, populations represents ecological recovery of bison. 1045 Avenue de la Médecine, Québec, QC, G1V We then outline recent conservation initiatives to dem- OA6 Canada onstrate that a decision on the future objectives of bi- son conservation needs to be addressed. PAUL R. KRAUSMAN, Boone and Crockett Wildlife Conservation Program, University of Montana, KEY WORDS: bison, Bison bison, conservation chal- 32 Campus Drive, Missoula, MT 59812, USA lenges, conservation, management, restoration
KYRAN KUNKEL, Wildlife Biology Program, Uni- Proceedings of the IVth International Wildlife versity of Montana, 32 Campus Drive, Missoula, Management Congress: 1-8, 2013 MT 59812, USA Plains bison (Bison bison; hereafter referred to as bi- KEITH AUNE, Wildlife Conservation Society North son) historically ranged across North American from American Program, 301 North Willson Avenue, the Rocky Mountains to the eastern seaboard and the Bozeman, MT 59714, USA plains of Canada to the northern reaches of Mexico (Reynolds et al. 1982, Danz 1997). In total, bison C. CORMACK GATES, Faculty of Environmental resided in 4 Canadian provinces, 42 U.S. states, and 5 Design, University of Calgary, 2500 University Mexican states, an area of >9,000,000 km2, thus en- Drive NW, Calgary, AB T2N 1N4, Canada compassing the largest distribution of any indigenous large herbivore in North America (Figure 1; Gates et al. 2010b). Throughout the Great Plains, bison in- SAM. D. FUHLENDORF, Department of Natural teracted with a host of species including pronghorn Resource Ecology and Management, Oklahoma (Antilocapra americana), elk (Cervus canadensis), State University, 008C Agricultural Hall, Stillwa- deer (Odocoileus spp.), prairie dogs (Cynomys spp.), ter, OK 74078, USA wolves (Canis lupus), grizzly bears (Ursus arctos) and grassland bird species through ecosystem alterations ABSTRACT: The conservation of the plains bison (Coppock et al. 1983, Krueger 1986, Knopf 1996, (Bison bison) is considered one of the greatest conser- Freese et al. 2007). Bison also play critical roles in vation success stories in North America. Although the grassland ecology through the facilitation of vegeta- historic distribution of bison (plains and wood bison tive heterogeneity (Knapp et al. 1999, Fuhlendorf et al. [B.b. athabascae]) was larger than any other indig- 2008). Wallowing activities can lead to standing water enous large herbivore in North America, market hunt- (Knapp et al. 1999), which in turn supports numer- ing and competition with the livestock industry reduced ous plant species (Collins and Uno 1983, Polley and the plains bison to ≤1,000, with only 25 free-ranging Wallace 1986) and provides habitat for prairie amphib- animals left in the world by 1902. Through the coop- ians (Bragg 1940, Corn and Peterson 1996). Vegeta- eration of private individuals, non-profit organizations, tion communities are affected by bison through graz- and the federal governments of the U.S. and Canada, ing, physical disturbance, nutrient cycling, and seed bison were saved from extinction and are now scat- dispersal (McHugh 1958, Knapp et al. 1999). These tered throughout much of their historical range, num- activities influence grassland heterogeneity, supporting bering >500,000 individuals. Despite the numerical prairie obligate species in the tall, mixed, and short- recovery of the species, recent questions have surfaced grass prairies (Powell 2006, Fuhlendorf et al. 2008, regarding the true success of this effort as <21,000 Gates et al. 2010a). plains bison (<5% of all bison) are managed within Prior to the arrival of Europeans in North Amer- conservation herds (n = 62). Thirteen percent (n = ica, the estimates of bison numbers ranged from 15 8) of herds are outside of their historical range, 92% to 100 million individuals (Dary 1989, Shaw 1995),
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Figure 1. Original ranges of plains bison (light gray area) and wood bison (dark gray area). Permission granted by Potter et al. 2010. History of bison in North America. Page 7 in Gates et al. 2010a)
however most estimates range from 30 to 60 million (Coder 1975, Danz 1997). In Canada, federal conser- bison (Seton 1929, McHugh 1972, Lott 2002). Fol- vation began in 1907 with the purchase of the privately lowing European settlement, bison numbers declined owned Pablo-Allard herd from Montana, U.S. (Freese rapidly as a result market hunting by European set- et al. 2007). tlers (Hornaday 1887, Isenberg 2000) in addition to These conservation efforts increased the bison competition with domestic livestock (McHugh 1972, population, which doubled between 1888 and 1902 Dary 1989, Danz 1997, Isenberg 2000). As a result, (Coder 1975) and increased steadily to approximately <1,000 bison were in North America by 1890 (Hor- 30,000 by the 1970s (McHugh 1972). Of the 30,000 naday 1887, Seton 1929) and wild, free-ranging bison bison, half resided in conservation herds (Freese et al. were extirpated from Canada (Freese et al. 2007) and 2007), defined as bison populations managed primar- nearly extirpated from the U.S. (Meagher 1973). ily for conservation rather than commercial produc- In the U.S., the loss of large bison herds led to tion (Boyd 2003). In the 1980s, commercial bison the first major conservation movement to preserve a production increased, resulting in >500,000 bison in species on the brink of extinction (Coder 1975). These North America today; yet, <5% of bison reside in efforts were led by private individuals who established conservation herds (Boyd 2003). In fact, the number small herds throughout the Great Plains (Boyd 2003). of individuals in conservation herds has remained rela- Private herds would later form the foundation for most tively stagnant since the 1930s, and today only 20,504 of the public herds (Boyd 2003). The second conser- individuals are located in 62 conservation herds (Boyd vation effort was led by the American Bison Society et al. 2010). In other words, despite the dramatic in- (formed 1905) who influenced the U.S. Congress to es- crease in bison during the past 120 years, the number tablish public conservation herds throughout the U.S. of free-ranging wild bison has lagged far behind.
Kohl et al. • Future of Bison Conservation
A further examination of these conservation herds CONSERVATION CHALLENGES provokes questions regarding the ecological effective- Domestication ness of bison restoration at a landscape scale. Of the Domestication may permanently alter the bison genetic 62 conservation herds, 13% (n = 8) are located outside pool while producing significant changes to morphol- the historical range of bison and 92% (n = 57) consist ogy, physiology, and behavior as a result of anthropo- of <1,000 individuals, a population size considered to genic selection and the loss of natural selection (Freese be genetically viable over the long term (Gates and El- et al. 2007). Within the commercial herds, cattle hus- lison 2010). Furthermore, only 8 % (n = 5) of herds bandry practices are common resulting in non-random are located on landscapes >2,000 km2 (Gates and Elli- selection of traits leading to docility, reduced agility, son 2010) and 16% do not contain breeding age males. and growth performance while also altering sex ratio Ecologically, wolves are the only effective predator of and age structure (Gates et al. 2010a). The elimination adult bison, yet they are associated with only 10% of of mature males further influences mate competition conservation herds (Gates and Ellison 2010). and natural selection (Gates et al. 2010a). Domesti- As a result, conservationists are questioning cated bison may also pose significant issues to bison whether bison are facing an ecological extinction event conservation if commercial animals establish cattle (Freese et al. 2007). To reverse this trend, the Wild- introgression within conservation herds when inten- life Conservation Society designed a foundation for tionally or accidently mixed with conservation herds bison restoration through the Vermejo Statement (Red- (Boyd et al. 2010). Furthermore, increased commer- ford and Fearn 2007) which states, “Over the next cen- cial herds may lead to the misconception that bison are tury, the ecological recovery of the North American no longer vulnerable to conservation issues because of bison will occur when multiple large herds move freely demographic recovery (Gates and Gogan 2010). across extensive landscapes within all major habitats of their historical range, interacting in ecologically sig- Hybridization nificant ways with the fullest possible set of other na- A concerted effort to create “beefalo” through the tive species, and inspiring, sustaining and connecting cross-breeding of bison and domestic cattle occurred human cultures.” during the late 1800s and early 1900s to create a more Expanding on this work, Sanderson et al. (2008) resilient winter species while maintaining the meat pro- established a scoring matrix to quantifying the conser- duction qualities of cattle (Dary 1989, Oglivie 1979). vation value of these herds. Exceptional contributors The concept dates back to the 16th century (Dary 1989) to ecological restoration included naturally structured and was actively pursued by the Canadian government herds of >5,000 individuals. Herds should consist as late as the early 1960s (Ogilvie 1979). These ef- of genetically pure and disease free animals which are forts have resulted in widespread domestic cattle gene impacted by all natural ecological interactions includ- introgression in the mitochondrial and nuclear DNA ing predation. Lastly, herds located on landscapes of bison (Halbert and Derr 2007). Today, only 8 con- >2,000 km2 are considered excellent contributors to servation herds are free of genetic introgression (Boyd bison recovery (Sanderson et al. 2008). Similarly, et al. 2010) and only the Yellowstone National Park Lott (2002) hypothesized that >13,000 km2 is neces- (YNP) and Wind Cave National Park (WCNP) herds sary for an ecologically functional prairie landscape. have had large enough samples to confidently evaluate More recently, Kohl (2012) examined single foraging introgression levels (Boyd et al. 2010). Of these 2, patch sizes of bison that when multiplied by historical recent work has demonstrated minimal introgression in spatial and temporal scales equates to landscape scales bison in WCNP (K. Kunkel, unpublished data). similar in size to these previous estimates. Further complicating the conservation of the bi- Given the current status of bison (Boyd 2003, son genome are questions related to historical and geo- Gates et al. 2010a) and restoration guidelines (Redford graphic differences. Despite low levels of cattle intro- and Fearn 2007, Sanderson et al. 2008, Gates et al. gression, some public herds may contribute to bison 2010b), we outline the important conservation chal- conservation due to unique historical and geographic lenges facing the ecological restoration of bison today, lineages (Halbert 2003, Halbert and Derr 2007). As and then discuss these challenges in light of the current a result, these lineages are important in the long-term model for bison conservation (i.e., numerous small, conservation of the bison genome regardless of intro- confined bison populations). In conclusion, we outline gression levels and should be preserved (Boyd et al. contemporary steps that are being taken to conserve 2010). Similarly, herds considered free of cattle intro- bison, and provide a comment on how things need to gression should be managed in isolation from hybrid- change to prevent the ecological extinction of bison. ized herds (Boyd et al. 2010).
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Disease highly complicated because of this conservation lega- Nine diseases are recognized by the International cy, particularly in cases of threatened species listing Union for the Conservation of Nature (IUCN)- Bison (see below). Specialist Group as diseases of concern for bison con- Listing.—From a global perspective, the IUCN servation; however, only the YNP and Grand Teton lists the bison (wood and plains bison) as “Near National Park/National Elk Refuge (Jackson herd) Threatened” (IUCN 2012). According to federal des- herds (24% of the bison conservation population) are ignation, bison are a “Red-Listed” species in Mexico; significantly impacted by chronic disease issues (Aune however, bison are currently not listed in Canada under and Gates 2010). the Species at Risk Act because of potential economic Brucellosis.—Brucellosis (Brucella abortus) is implications for the Canadian bison industry (Aune and primarily found in bovine species; however, elk play a Wallen 2010). Bison are classified as “Threatened” by transmission role in the Greater Yellowstone Ecosys- the Committee on the Status of Endangered Wildlife in tem (Davis 1990). Primarily transmitted through oral Canada. Similarly, bison are not listed under Endan- contact with aborted fetuses, contaminated placentas, gered Species legislation in the U.S. A primary dif- and uterine discharges (Reynolds et al. 1982, Tessaro ficulty for listing in the U.S. and Canada is complica- 1989), brucellosis leads to first pregnancy abortion in tion caused by the classification of hybridized animals > 90% of infected female bison (Davis 1990, Davis et (Boyd and Gates 2006) and the role of commercial bi- al. 1990). Natural immunity reduces the abortion rate son producers in the numerical status of bison. to 20% and ~ 0% by the second and third pregnancy, Classification.—The classification of bison as respectively (Davis 1990, Davis et al. 1990). Due to wildlife, livestock or both, is jurisdictionally depen- similar symptoms in domestic cattle, infected bison dent. Within their historical range, bison are classified populations are heavily managed to minimize transmis- as wildlife in 4 Canadian provinces, 10 U.S. states, sion from bison to domestic livestock (Keiter 1997), and 1 Mexican state; however, free-ranging popula- despite no confirmed cases of transmission in the wild tions do not exist in all these areas. Where bison do (Bienen 2002, Cheville et al. 1998, Shaw and Mea- exist as “wildlife”, they are typically managed within gher 2000). No highly effective vaccine is available; fence preserves (Aune and Wallen 2010). Outside of however, quarantine protocols and test and slaughter these areas, bison are typically classified and managed protocols may effectively eliminate all animals within as livestock by private ownership, thus are governed an exposed population (Aune and Gates 2010). by animal health and trade regulations. In YNP, the Interagency Bison Management Ownership.—Bison conservation within the pri- Program (IBMP) was adopted as a cooperative, multi- vate sector is largely a secondary objective behind com- agency plan to guide bison and brucellosis management mercial production. However, conservation groups to maintain wild, free-ranging bison, while reducing such as The Nature Conservancy, American Prairie the transmission risk of brucellosis to domestic cattle Reserve, and World Wildlife Fund have established (USDOI and USDA 2000). Management has incorpo- privately owned herds focused on bison conservation. rated multiple strategies including spatial and temporal In addition, North American indigenous peoples are separation of bison and cattle, capture-test-slaughter playing a key role in conservation because of the bi- actions, quarantine and translocation actions, hazing son’s cultural importance and role in restoring cultural of bison back into YNP, and vaccination (USDOI and connections in addition to dietary and economic ben- USDA 2000). In the Jackson herd, intense manage- efits. Despite these efforts, bison managed by these ment actions based on spatial and temporal separation groups face difficult challenges such as management are not as prominent as in YNP, rather management restrictions, insufficient funding, and litigation. aims to reduce elk and bison dependency upon the win- ter feeding grounds while utilizing brucellosis vaccina- Availability of Restoration Sites tion program for both species while simultaneously us- The identification of bison restoration sites has been ing hunting to reduce and later maintain the population particularly problematic because landscapes large at 500 individuals (Aune and Gates 2010). enough to support ecologically interactive bison popu- lations are limited. In particular, human development Legal Status and habitat conversion has expanded into many areas Bison do not typically have equal legal or policy status capable of supporting large bison populations. For ex- when compared to other wildlife species because of ample, private agricultural operations can be found in early conservation practices that classified and man- almost all areas suitable for bison restoration, even on aged bison as a form of livestock (Aune and Wallen public land. Besides direct forage competition between 2010). Today, bison management and conservation is bison and livestock, potential conflicts may include hu-
Kohl et al. • Future of Bison Conservation
man safety issues and property damage (e.g., fencing, levels of cattle introgression were removed from the crop depredation). As a result, the social difficulties of population in 2011. Genetically pure animals from Elk translocation have increased, particularly when consid- Island National Park were used to augment the popula- ering the restoration of bison to mixed-ownership and tion in 2010 (n = 94) and 2012 (n = 70). The bison mixed-management landscapes. These areas require pasture will double in the coming years and will be detailed restoration plans which provide guidelines for further expanded as land purchases allow. No large dealing with management issues and conflicts. In these predators currently exist in the region; however, griz- areas, coordination among private individuals, local, zly bears have ventured within 200 km of the reserve state, federal, and tribal governments, wildlife agen- (Robbins 2011). Future reserve plans include a land- cies, conservation organizations and other concerned scape of >14,000 km2 that support a full suite of native parties has been difficult but successful and is a neces- wildlife including 10,000 bison and large predators. sity for long-term success. El Uno Ecological Reserve DISCUSSION AND REVIEW OF CURRENT INI- The El Uno Ecological Reserve (EUER), located in TIATIVES Chihuahua, Mexico, is owned and managed by The Throughout the last century it has been easier to start Nature Conservancy. The EUER is a part of the larg- and maintain multiple small herds due to challenges er Janos Biosphere Reserve which consists of a mix listed above, however this trend raises questions about of private owners and ejidos (communal agriculture the efficacy of this model in the future of bison con- lands). Bison were translocated from WCNP in 2009 servation. Have bison been restored ecologically? Are with 38 animals currently occupying approximately 20 conservation initiatives actually meeting the restoration km2 of the EUER. The EUER has been registered as criteria set forth by Freese et al. (2007), Sanderson et a Wildlife Management Area with the Mexican Fed- al. (2008), and Gates et al. (2010b)? The following eral Government which will permit an increase in the review of recent bison initiatives provides insight into bison pasture and herd size. As a result, future man- these questions. agement will maintain a sustainable herd size of 157 animals and a maximum of 210 (Laura Paulson, The Grasslands National Park Nature Conservancy, personnel communication). No Grasslands National Park is located on the U.S. – Can- sustainable populations of large predators are currently ada border in southern Saskatchewan. The park was located in the region. formally established in 1981 by the Canadian federal These examples illustrate the range of recent res- government and is currently managed by Parks Can- toration efforts for bison. From government-based ada. Seventy-one bison were translocated from Elk herds on public land (i.e., Grasslands National Park) Island National Park in 2005 and have since grown to to completely private conservation herds (i.e., EUER), approximately 350 individuals. Disease-free and ge- the conservation of bison comes in multiple forms. netically pure, these animals are maintained within 182 However, when considering the criteria for the eco- km2 fenced region of the park. Current management logical restoration of bison, many programs fall short. plans will maintain the herd at approximately 800 ani- Are these bison herds able to move freely across the mals; however, a range expansion may occur in com- landscape? Are the effort’s goals >5,000 individu- ing years providing availability for a larger population. als? Will these herds face historical predation? These Although large predators are not found in the area, questions seem easy to answer, but is this the model future expansion of wolves into the area may lead to of bison conservation that will actually restore bison, overlap with the park. ecologically, in North America?
American Prairie Reserve CONCLUSION American Prairie Reserve (APR) is a non-profit orga- As bison conservation continues forward, we as man- nization established in 2001 with the goal of acquiring agers and conservationists understand the challenges and managing private land and public grazing leases associated with bison restoration. Additionally, we to establish a fully-functioning prairie-based wildlife have been provided with guidelines for the successful reserve in north-central Montana, USA. Since estab- restoration of bison (Redford and Fearn 2007, Sander- lishment, APR has purchased and leased 498 km2 of son et al. 2008, Gates et al. 2010b). After reviewing prairie, of which 57 km2 currently support bison. Ap- the projects above in light of ecologically restored bi- proximately 325 disease-free animals currently graze son populations, it leads us to speculate on the feasibil- the reserve. All animals have been genetically tested ity and intentions of bison restoration efforts. If we and all animals acquired from WCNP which contained understand the challenges and issues associated with
IVth International Wildlife Management Congress bison restoration, shouldn’t conservation target the servation guidelines 2010. International Union for restoration of the species in landscapes large enough Conservation of Nature, Gland, Switzerland. to support ecologically restored bison herds, not small Bragg, A. N. 1940. Observations on the ecology and bison herds maintained behind wire? If so, then we as natural history of Anura I: habits, habitat, and managers and conservationists, along with the general breeding of Bufo congantus Say. American Natu- public, and all interested and affected parties must join ralist 74: 424-438. in the discussion on how we proceed. Cheville, N. F., D. R. McCullough, and L. R. Paul- If we as society desire an ecologically restored son. 1998. Brucellosis in the Greater Yellowstone bison population, we must then initiate a shift in bison Area. National Academy Press, Washington D. conservation which takes us away from small, isolated C., USA. conservation herds and toward ecological recovery. Coder, G. D. 1975. The national movement to pre- Naturally, there will be social, political, and biological serve the American buffalo in the United States ramifications to deal with; however, these issues are and Canada between 1880 and 1920. Dissertation. not new to wildlife conservation. To proceed, we must Ohio State University, Columbus, USA. shift our focus toward expanding existing herds that Collins, S. L., and G. E. Uno. 1983. The effect of have the potential for a large ecological footprint. In early spring burning on vegetation in buffalo wal- regions currently lacking bison, we must focus our ef- los. Bulletin of the Torrey Botanical Club 110: forts on the translocation of bison to areas that can meet 474-481. the aforementioned conservation criteria. Never again Coppock, D. L., J. E. Ellis, J. K. Detling, M. I. Dyer. will bison roam North American by the millions, but it 1983. Plant-herbivore interactions in a North is ecologically, culturally, and historically important to American mixed-grass prairie. II. Responses of bi- continue the conservation of this North American icon. son to modification of vegetation by prairie dogs. Thus, it is time we shift our model of bison conserva- Oecologia 56:10-15. tion toward ecologically functional populations rather Corn, S. P., and C. R. Peterson. 1996. Prairie legacies than livestock and fenced wildlife. – amphibians and reptiles. Pages 125-234 in F. B. Samson and F. L. Knopf, editors. Prairie Conser- vation. Island Press, Washington D.C. USA. LITERATURE CITED Danz, H. P. 1997. Of bison and man: from the annals Aune, K. E. and C. C. Gates. 2010. Reportable or of a bison yesterday to a refreshing outcome from notifiable diseases. Pages 27-38 in C. C. Gates, human involvement with American’s most valiant C. H. Freese, P. J. P. Gogan, and M. Kotzman, of beasts. University of Colorado Press, Niwot, editors. American Bison: Status survey and con- USA. servation guidelines 2010. International Union for Dary, D. A. 1989. The buffalo book: the full saga of Conservation of Nature, Gland, Switzerland. the American Animal. Swallow Press, Chicago, Il- Aune, K. E., and R. L. Wallen. 2010. Legal status, linois, USA. policy issues and listings. Pages 63-84 in C. C. Gates, C. H. Freese, P. J. P. Gogan, and M. Davis, D. S. 1990. Brucellosis in wildlife. Pages 321- Kotzman, editors. American Bison: Status survey 334 in K. Nielsen and J. R. Duncan, editors. Ani- and conservation guidelines 2010. International mal brucellosis. CRC Press, Boca Raton, Florida, Union for Conservation of Nature, Gland, Swit- USA. zerland. Davis, D. S., J. W. Templeton, T. A. Ficht, J. D. Bienen, L. 2002 Informed decisions: conservation cor- Williams, J. D. Kopec, and L. G. Adams. 1990. ridors and the spread of disease. Conservation in Brucella abortus in captive bison. I. serology, bac- Practice 3:10-17. teriology, pathogenesis and transmission to cattle. Boyd, D. P. 2003. Conservation of North American Journal of Wildlife Diseases 26:360-371. bison: status and recommendations. Thesis, Uni- Freese, C. H., K. E. Aune, D. P. Boyd, J. N. Derr, versity of Calgary, Alberta, Canada. S. C. Forrest, C. C. Gates, P. J. P. Gogan, S. Boyd, D. P. and C. C. Gates. 2006. A brief review of M. Grassel, N. D. Halbert, K. Kunkel, and K. H. the status of plains bison in North America. Jour- Redford. 2007. Second chance for the plains bison. nal of the West 45:15-21. Biological Conservation 136:175-184. Boyd, D. P., G. A. Wilson, J. N. Derr, and N. D. Hal- Fuhlendorf, S. D., D. M. Engle, J. Kerby, and R. bert. 2010. Genetics. Pages 19-26 in C. C. Gates, Hamilton. 2008. Pyric herbivory: rewilding land- C. H. Freese, P. J. P. Gogan, and M. Kotzman, scapes through the recoupling of fire and grazing. editors. American Bison: Status survey and con- Conservation Biology 23:588–598.
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Gates, C. C. and K. Ellison. 2010. Numerical and geo- Prairie conservation: preserving North America’s graphic status. Pages 55-62 in C. C. Gates, C. H. most endangered ecosystem. Island Press, Wash- Freese, P. J. P. Gogan, and M. Kotzman, editors. ington D.C., USA. American Bison: Status survey and conservation Kohl, M. T. 2012. Bison conservation in the northern guidelines 2010. International Union for Conserva- Great Plains. Thesis, University of Montana, Mis- tion of Nature, Gland, Switzerland. soula, USA. Gates, C. C. and P. J. P. Gogan. 2010. Introduc- Krueger, K. 1986. Feeding relationships among bison, tion: the context. Pages 1-4 in C. C. Gates, C. H. pronghorn, and prairie dogs: an experimental anal- Freese, P. J. P. Gogan, and M. Kotzman, editors. ysis. Ecological Society of America 67:760-770. American Bison: Status survey and conservation Lott, D. F. 2002. American bison: a natural histo- guidelines 2010. International Union for Conserva- ry. University of California Press, Los Angeles, tion of Nature, Gland, Switzerland. USA. Gates, C. C., C. H. Freese, P. J. P. Gogan, and M. McHugh, T. 1958. Social behavior of the American Kotzman. 2010a. American bison: status survey buffalo (Bison bison bison). Zoologica 43:1-40. and conservation guidelines 2010. International McHugh, T. 1972. The time of the buffalo. University Union for Conservation of Nature,, Gland, Swit- of Nebraska Press, Lincoln, USA. zerland. Meagher, M. M. 1973. The bison of Yellowstone Gates, C. C., R. O. Stephenson, P. J. P. Gogan, C. National Park. National Park Service Scientific H. Freese, and K. Kunkel. 2010b. Guidelines for Monograph Series 1, Mammoth, Wyoming, USA. ecological restoration of bison. Pages 103-112 in Ogilvie, S. C. 1979. The Park Buffalo. Calgary-Banff C. C. Gates, C. H. Freese, P. J. P. Gogan, and M. Chapter, National and Provincial Parks Associa- Kotzman, editors. American Bison: Status survey tion of Canada, Calgary, Alberta, Canada. and conservation guidelines 2010. International Polley, H. W. and L. L. Wallace. 1986. The relation- Union for Conservation of Nature, Gland, Swit- ship of plant species heterogeneity to soil variation zerland. in buffalo wallows. American Midland Naturalist Halbert, N. D. 2003. The utilization of genetic mark- 112: 178-186. ers to resolve modern management issues in his- Powell, F. L. A. 2006. Effects of prescribed burns toric bison populations: implications for species and bison (Bos bison) grazing on breeding bird conservation. Dissertation, Texas A&M Univer- abundances in tallgrass prairie. The Auk 123: 183- sity, College Station, USA. 197. Halbert, N. D., and J. Derr. 2007. A comprehensive Redford, K. H., and E. Fearn. 2007. Ecological future evaluation of cattle introgression into U.S. federal of bison in North America: a report from a multi- bison herds. Journal of Heredity 98:1-12. stakeholder. Wildlife Conservation Society Work- Hornaday, W. T. 1887. The extermination of the ing Paper No. 30, Bronx, New York, USA. American bison with a sketch of its discovery and Reynolds, H. W., R. D. Glaholt, and A. W. L. Haw- life history. U.S. National Museum, Washington, ley. 1982. Bison. Pages 972-1007 in J.A. Chap- D.C. USA. man and G.A. Feldhamer, editors. Wild mammals International Union for Conservation of Nature of North America: biology, management, and eco- [IUCN]. 2012. IUCN red list of threatened spe- nomics. The Johns Hopkins University Press, Bal- cies.
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Seton, E. T. 1929. Lives of game animals, 4 volumes. U.S. Department of the Interior and U.S. Department Doubleday, Doran & Co., Garden City, New of Agriculture (USDOI and USDA). 2000. Re- York, USA. cord of the decision for final environmental impact Shaw, J. H. 1995. How many bison originally populat- statement and bison management plan for the state ed western rangelands? Rangelands 17:148-150. of Montana and Yellowstone National Park. U.S. Shaw, J., and M. Meagher. 2000. Bison. Pages 447- Government Printing Office, Washington D.C., 466 in S. Desmarais and P. R. Krausman, editors. USA. Ecology and management of large mammals in North America. Prentice Hall, Upper Saddle Riv- er, New Jersey, USA. Tessaro, S. V. 1989. Review of the diseases, para- sites and miscellaneous pathological conditions of North American bison. Canadian Veterinary Jour- nal 30:416-422.