REINTRODUCING PLAINS BISON (BOS BISON) TO AMERICAN … · REINTRODUCING PLAINS BISON (BOS BISON) TO AMERICAN PRAIRIE FOUNDATION LANDS IN NORTHCENTRAL MONTANA: 5-YEAR CONSERVATION

Diane Hargreaves / hargreavesphoto.com

REINTRODUCING PLAINS BISON (BOS BISON) TO AMERICAN PRAIRIE

FOUNDATION LANDS IN NORTHCENTRAL MONTANA: 5-YEAR CONSERVATION AND MANAGEMENT PLAN

Kyran Kunkel, Steve Forrest, Curtis Freese

for

Northern Great Plains Ecoregion Program World Wildlife Fund – US

PO Box 7276 Bozeman, MT 59771

REVIEWERS DRAFT

This is a draft document. The authors request that the contents not be cited without permission from the lead author or World Wildlife Fund.

APF BISON REINTRODUCTION PLAN 1

Table of Contents EXECUTIVE SUMMARY..............................................................................................4 INTRODUCTION

Problem..................................................................................................................5 Genetic and population concerns Ecological role of bison and associated ecosystem benefits Bison v. cattle Factors limiting bison restoration

Solutions..................................................................................................................8 Opportunities for Ecological Restoration of Bison in the Northern Great Plains Partnerships for Restoration in the Montana Glaciated Plains

Conservation Goals and Objectives....................................................................10 10-year objectives Premises

METHODS

Study area description.........................................................................................13 Biological feasibility.............................................................................................15

Reasons for bison absence from area Availability of suitable habitat Minimum area required Elimination of threats

Hunting Disease

Genetic management and habitat loss Significance of cattle introgression

Previous reintroductions Source herd evaluation.........................................................................................17

Castle Rock, New Mexico Wind Cave National Park, Wyoming Elk Island National Park, Alberta Henry Mountains, Utah Source herd recommendation

Optimal release strategy modeling.....................................................................20 Parameters

Minimum founding size Minimum viable population size

Population demographic inputs Modeling results Comparison with other populations

Additional justification for release strategy......................................................24 Habitat suitability................................................................................................25

Number of bison area can support Rangeland condition Mid-term capacity

Ecologically effective minimum population size...............................................26 Effect of reintroducing bison..............................................................................26

Special status species Availability of suitable release stock and impacts to donor populations


Socio-economic and legal considerations...........................................................27 Qualifications of restoration team Public planning and participation

TRANSLOCATION

Acquisition methods.............................................................................................28 Disease and genetics screening............................................................................29 Release methods....................................................................................................30

Handling facility Fencing

MANAGEMENT

Disease....................................................................................................................31 Escapes...................................................................................................................31 Mortality................................................................................................................32

Culling Disposition of culled animals

Genetics and population management................................................................33 Bison movement and pasture management........................................................34 Grassland management........................................................................................35 Other wildlife management.................................................................................36 Organizational Risk Management Controls......................................................36

Risk Management Dissolution of the Project

MONITORING AND RESEARCH PROGRAM (EXPERIMENTAL DESIGN)

Justification...........................................................................................................38 Bison Research Needs...........................................................................................38 Methods.................................................................................................................40 Dissemination of Results......................................................................................42

CRITERIA FOR MEASURING SUCCESS..................................................................42 ACKNOWLEDGMENTS................................................................................................44 LITERATURE CITED....................................................................................................45 APPENDICES

Appendix 1. Variables used in the Vortex Modeling........................................56 Appendix 2,3,4. Sample BLM Environmental Assessments............not attached Appendix 5. Bison Care and Use Plan.....................not attached, in development Appendix 6. APF IRS Determination Letter....................................not attached Appendix 7. Bison restoration review team and collaborators.......................60

FIGURES, TABLES

Fig. 1. Site Location Map.......................................................................................13 Fig. 2. Site Detail and Proposed Pastures Map......................................................14


Fig. 3. Bison Population Trend..............................................................................23 Table 1. Representative acreages in ownership.....................................................25

Table 2. Effect of bison restoration on special status species in and near APF project area.................................27

Table 3. Project Milestone Chart (similar for each of 2004, 2005, 2006)....................................................43


EXECUTIVE SUMMARY Perhaps no species is as emblematic of the grasslands of North America as the plains bison (Bos bison1), whose numbers were described as “innumerable” by accounts of European explorers of the Great Plains in the early 19th century. The plains bison is also an icon of the conservation movement, one of the first species that stirred a national effort to intervene on behalf of a species that seemed destined for extinction. Until recently, these efforts were regarded as a conservation success story. A number of herds are maintained on public lands and numbers of bison in production herds are increasing throughout North America. Recent work by Dr. James Derr and his associates at Texas A&M University (Halbert 2003) has found, however, that at least some of this success has come at the cost of the bison genome—with a few important exceptions, a vast majority of the bison in both public and private herds today are contaminated with cattle DNA, a result of crossbreeding in the early days of the last century. In some cases, small herd size exacerbates the erosion of this limited gene pool. The American Prairie Foundation (APF) is a 501(c)(3) Montana not-for-profit land trust with holdings in south Phillips County, Montana, within what is described as the Montana Glaciated Plains, a mixed-grass prairie. The base property controls federal and state leases2 that are currently grazed by cattle, some within the existing boundaries of the Charles M. Russell National Wildlife Refuge. APF’s goal is to manage its deeded and leased holdings for the benefit of wildlife and wildlife habitat. Consistent with that goal, and in the interest of furthering restoration of native grazers to the mixed-grass prairie ecosystem, World Wildlife Fund (WWF) is assisting APF in developing this reintroduction proposal to establish a conservation herd with the express purpose of maintaining the bison genome in perpetuity and of restoring the ecological role of bison over a large area. This herd should be one of several new conservation herds that need to be established in North America. We are prepared to expend the resources and to work with stakeholders and cooperators to ensure the greatest likelihood of success for this project. This document will serve as our reintroduction plan and feasibility study to ensure that we have taken the necessary steps to achieve our objectives, as well as to provide information for our collaborators and partners. This plan is being circulated for review by the IUCN Bison Specialist Group and all of our collaborators (Appendix 5).

1 We follow the convention of van Gelder (1977) for inclusion in the genus Bos, although Bison is traditionally used and also appropriate (Halbert 2003). 2 The current BLM Telegraph Creek Allotment (#5654) and Montana state leases 8401, 7013, 8443, 5303, and 8159 within the boundaries of the CMR Refuge. CMR and APF currently manage grazing on these internal leases through an exchange of use agreement that allows APF grazing on Refuge pastures contiguous to APF property.


INTRODUCTION Problem Genetic and population concerns Once numbering around 30 million (Lott 2002), the plains bison (Bos bison) was a dominant keystone species (Soulé et al. 2003) that ranged over almost all the grasslands of North America, with the Great Plains its stronghold. Today, however, the plains bison is ecologically extinct at any significant scale (Estes et al. 1989; Novaro et al. 2000; Boyd 2003a, 2003b). There are few plains bison populations within the original range that exist under natural conditions, and none that are considered viable by the current benchmark (Boyd 2003a). Free-ranging, disease–free (i.e., free of key problematic diseases such as brucellosis) populations that are potentially influenced by naturally regulating factors in the original range of plains bison account for only 1,289 bison (6.7% of total conservation population). These are in 4 herds: Antelope Island and Henry Mountains, Utah; Prince Albert National Park, Saskatchewan; and Primrose Air Weapons Range, Alberta/Saskatchewan. One of these herds is subject to handling. None of them are >400 animals, well below the minimum of 1,000 animals considered necessary for long-term maintenance of 90% of genetic (allelic) diversity (Gross and Wang 2005). The only 3 federal U.S. herds where bison conservation is a priority objective are Fort Niobrara National Wildlife Refuge (NWR) in Nebraska, the National Bison Range in Montana, and Wichita Mountains NWR in Oklahoma, but all of these herds entail conservation problems such as unnatural culling and confinement to relatively small areas. Therefore, no disease-free viable populations exist under natural conditions. More problematic than ecological extinction because of its irreversibility is the erosion of the wild bison genome. The 96% of North America’s bison that are found in private herds are subject to artificial selection for domestication, with ease of handling and marketable meat production as major goals. Small herd size (usually only tens or hundreds of animals), unnatural culling practices, artificial genetic selection, and lack of a coordinated management strategy for these herds make them questionable repositories for conservation of the plains bison genome. Even more troubling is the existing level of introgression of domestic cattle genes in the bison genome. Nearly every private bison herd tested to date (49 out of 50 tested; J. Derr, Texas A and M University, personal communication) harbors cattle DNA (Halbert 2003). In addition, nearly all significant public herds have been tested and most exhibit cattle gene introgression. Among plains bison, the only public herds known for which there is a good probability of genetic purity (97% probability of non-contamination) are Henry Mountains (Utah), Yellowstone National Park (Wyoming and Montana), Wind Cave National Park (South Dakota) (Halbert 2003) and Elk Island National Park (Alberta; Ward et al. 1999). Efforts are just beginning to comprehensively address genetic issues in


the management of these herds and of other herds with cattle gene introgression that are nevertheless important for conserving genetic diversity in bison. Based on these significant conservation concerns, Boyd’s (2003a) exhaustive review indicated bison warrant consideration for listing as an imperiled species under the relevant Canadian and U.S. laws. Montana lists free-ranging herds as a species of concern with an S2 status: “imperiled because of rarity or because other factors demonstrably making it very vulnerable to extinction throughout its range” (Carlson 2003). Montana classifies wild plains bison as a game animal but there is no season or license issued. A season is currently being examined for bison ranging out of Yellowstone National Park. The U.S. National Park Service is now developing management guidelines to maintain the genetic diversity of its herds (P.Gogan, US Geological Survey, pers. comm.). Canada’s Committee on the Status of Endangered Wildlife in Canada (COSEWIC) in May, 2004 listed plains bison as a threatened species (COSEWIC 2004). The World Conservation Union, formerly known as the International Union for Conservation of Nature and Natural Resources (IUCN) Red Book lists bison as Lower Risk, conservation dependent (1996; IUCN 2002), indicating that bison, as a species, are currently the focus of a conservation program. If current conservation programs were to cease, the species would become vulnerable, endangered or critically endangered within 5 years. No distinction is made between the wood and plains bison subspecies. Given this conservation situation, the IUCN North American Bison Specialist Group (BSG) is currently completing a status assessment and will conduct a review in the next 12 months of the need to red-list the species (the “red list” is IUCN’s threatened and endangered species list for the world). That the plains bison is perhaps more imperiled than previously thought has important management implications for the genome and bison conservation in general. Ecological role of bison and associated ecosystem benefits Increasingly, ecological data indicate that second-order consequences of species loss—the loss of species interactions—may be even more ominous than the first-order loss of species (Soulé et al. 2003). Accordingly, a primary mission of conservation should be to restore interactive species (Soulé et al. 2003). Highly interactive species that are also very abundant have been termed foundation species. Perhaps the most profound loss of foundation species in North America has been the loss of free-ranging wild bison. Although bison exist in a few small public herds and the species is gaining popularity as an alternative breed of domestic livestock, the bison of the Great Plains is today ecologically extinct at any significant scale (Soule et al. 2003; Pyare and Berger 2003). Bison disturbance (grazing, trampling, and wallowing) no longer influences native vegetation and species composition over large scales as it once did. Bison-style grazing no longer creates the mosaic of vegetative structures over large areas that provided habitats for many other species. Grazing, wallowing, and movement of herds were instrumental in shaping the prairie landscape, including influencing the distribution of many prairie birds (Truett et al. 2001). Bison likely played a significant role in


establishment of prairie dogs, the distribution of other large herbivores, and nutrient cycling (Truett et al. 2001). As the principal large converters of grass to animal biomass, they supported large populations of predators and scavengers, such as aboriginal humans, wolves, bears, wolverines, bald eagles, ravens, coyotes, and swift foxes (Mead 1899; Roe 1951:155-158; Bogan 1997). Finally, decomposing bison carcasses no longer create rich patches of nutrients for vegetative growth (Towne 2000). It has been argued that management of domestic livestock can be employed to mimic the effects of bison. The reality, however, is that livestock, and even few bison herds, are managed in this way today (Plumb and Dodd 1993; Hartnett et al. 1997; Frelich et al 2003). There are probably few communities or species in the Great Plains that in one form or another were not affected by the presence of bison. One of the most significant interactive relationships occurred between bison and prairie dogs. Historical accounts and recent studies leave little doubt that bison co-existed with and benefited prairie dogs (Cynomys spp., J. Truett, Turner Endangered Species Fund, unpublished report). Prairie dogs in turn elevated biodiversity by serving as food or providing burrows for numerous other species (Miller et al. 1994; Kotliar et al. 1999; Truett et al. 2001). Birds known to selectively use black-tailed prairie dog (C. ludovicianus) colonies include mountain plovers (Charadrius montanus) (Knowles et al. 1982), ferruginous hawks (Buteo regalis) (Roth and Marzluff 1989), and burrowing owls (Speotyto cunicularis) (Desmond and Savidge 1996; Truett and Savage 1998, Agnew et al. 1986). These 3 species have attracted recent attention as priority species for conservation; the mountain plover and ferruginous hawk have both been proposed for listing under the U.S. Endangered Species Act at various times (Roth and Peterson 1997; US Fish and Wildlife Service 1999) and, along with the burrowing owl, are species at risk or extirpated in Canada. Bison v. cattle While cattle are often proposed as ecological surrogates for bison (cite), differences in the ecological influences of bison versus cattle have been noted. At Konza Prairie and Utah (Knapp et al. 1999, Van Vuren 2001), results indicate that the abundance and richness of annual forbs, and the spatial heterogeneity of biomass and cover, are higher in sites with bison than in sites with cattle. Plumb and Dodd (1993) reported a difference in diet of the 2 species with bison selecting more strongly than cattle for graminoids vs. forbs. Bison spend less time grazing than do cattle. Bison spent significantly more time on other behaviors including social interactions and defense. Bison grazing, especially in combination with burning, has the potential for inducing a much more complex set of local interactions and spatial and temporal patterns than cattle grazing (Steuter 1997). We will address these questions as hypotheses in our restoration monitoring plan below. Factors limiting bison restoration The most fundamental limit to bison recovery is the lack of large conservation areas where bison populations of minimum viable size acting on an ecologically meaningful scale can be established. Large areas of suitable habitat of native or largely native prairie


exist (Forrest et al. 2004). Growth and augmentation of current bison herds are potential methods for bison recovery, but over half of managers of bison conservation herds indicate there is no room for expansion and none indicated room for any significant expansion (Boyd 2003a). We know of no large conservation area where significant expansion of the current range for bison is contemplated. The pressures of a developed landscape, a burgeoning commercial bison industry, and localized issues like disease and absence of predators constrain the current possibilities for effective bison recovery. Identification and evaluation of new recovery sites is needed (Boyd 2003a). We know of no current plans for development of conservation herds of wild bison in the U. S., although a few initiatives in Canada are underway (P. Fargey, Grasslands National Park, pers. comm.; S. Michalsky, Nature Conservancy Canada, pers. comm.). Solutions The current distribution of the uncontaminated bison genome, in fewer than 5 herds, calls for the need to establish additional conservation herds that will contribute to a comprehensive strategy to maintain the genome in an environment of natural selection pressure. Bison herds need to be established on a scale that: 1) reduces the risk of genetic erosion posed by small herd size; 2) allows the full expression of ecological, behavioral and evolutionary relationships; and 3) accounts for incorporation of ecosystem processes such as fire and drought. Areas as large as 3.2 million acres (5,000 sq mi/12,500 sq km) have been suggested as necessary to sustain wild bison on ecologically meaningful scales (Lott 2002), but the issue of “how large,” beyond minimum levels needed to population and genetic viability, remains to be tested. Needless to say, landscapes that have the potential to accommodate large numbers of bison are a prerequisite. Reintroductions are the only way to expand the species’ range, as there are few source populations and natural dispersal has been eliminated through management polices on lands adjacent to these populations. While reintroductions are relatively lengthy, complex, risky, and expensive conservation endeavors (IUCN/Species Survival Commission Re-introduction Specialist Group 1998), we propose that the status of wild bison will remain unchanged or worsen unless proactive management actions like reintroductions are taken. Opportunities for Ecological Restoration of Bison in the Northern Great Plains With less than 1.5% of the Northern Great Plains (NGP) ecoregion land area managed primarily for biodiversity conservation, few areas currently exist where bison conservation, and more broadly grassland conservation at ecologically significant scales, can be accomplished. A 2004 assessment of the Northern Great Plains (NGP) ecoregion by WWF and the Northern Plains Conservation Network (NPCN; Forrest et al. 2004), and more recently by the Commission on Environmental Cooperation (Hoth, pers. comm.) identified a number of biologically significant landscapes in all or portions of the Great Plains. The WWF/NPCN assessment identified 10 terrestrial landscapes in the U.S. and Canadian plains of 2-3 million acres in size as having sufficient intact grasslands in combination with existing land management where opportunities exist to restore large-scale ecological processes and provide habitat for significant populations of native wildlife.


Partnerships for Restoration in the Montana Glaciated Plains Among the 10 exceptional areas identified in the WWF/NPCN Northern Great Plains ecoregional assessment is the Montana Glaciated Plains, a mixed-grass prairie landscape of northcentral Montana (Figure 1), which includes extant populations of several at-risk grassland and shrubland-steppe species (black-footed ferrets, prairie dogs, sage grouse, mountain plovers, burrowing owls, ferruginous hawks, among others) as well as some of the largest blocks of intact grasslands in the ecoregion. The ecological importance of the area has also been noted by The Nature Conservancy in its Northern Great Plains Steppe ecoregional plan (TNC 2000) and as one of 8 conservation areas identified statewide in Montana by the Partners for Fish and Wildlife (Undated). In short, the Montana Glaciated Plains not only retains a great amount of historic biological integrity but also contains unfragmented grasslands of sufficient area that it could support a large number of bison. WWF, partnering with the APF, launched the American Prairie Restoration Project (APRP) in 2002 with the long-term goal of creating a large-scale prairie reserve capable of supporting several thousand or more bison in semi-natural conditions. In January 2004 APF acquired the project’s first 20,000-acre ranch adjacent to the C.M. Russell National Wildlife Refuge (CMR) in southern Phillips County, Montana, within the Montana Glaciated Plains. More than 90% of this landscape consists of intact native prairie. The 1-million-acre (0.4-million-ha) CMR, contiguous to the project area, is the largest protected area in the northern Great Plains (Fig. 2). APF plans to utilize private and leased grazing capacity to expand the herd to a minimum size of 400 animals over the short term, and expand capacity as additional land and forage become available over time. To this end, APF will cooperate with the Bureau of Land Management (BLM), the Montana Department of Natural Resources and Conservation (DNRC), and other interested private parties to obtain sufficient leased grazing capacity, as well as acquiring additional capacity as other properties come on the market. The CMR offers a critical buffer to reduce potential conflicts with private ranch lands to the south of the project area as well as complementing APF and WWF’s wildlife management objectives. Our plans call for starting the herd by beginning bison reintroduction by the fall of 2005 or spring of 2006. The new data on threats to the wild bison genome give added impetus to ensuring that this herd is established in the context of a North American bison conservation strategy. We are currently collaborating with the National Park Service and the IUCN Bison Specialist Group to develop criteria for herd selection and composition, to arrange acquisition of animals from appropriate sources, and to design a reintroduction and management plan for this conservation herd.

At the same time, because of WWF’s interest and leadership in conservation in the Northern Great Plains ecoregion, and because of the IUCN Bison Specialists Group’s interest in having WWF participate in the development of a North American bison action plan, the information and plans developed for this bison introduction will be instrumental in guiding bison conservation efforts underway elsewhere in the ecoregion and in shaping the action plan.


Conservation Goals and Objectives Our goal is to establish a landmark herd for conservation of the plains bison in North America, with a focus on both conserving the wild bison genome and re-establishing the bison’s role as a dominant keystone species on a major prairie reserve. This project will enable us to expand and disperse the extant known pure and relatively diverse bison genome within the Great Plains. Currently, Wind Cave National Park [WC] and Nature Conservancy Canada’s Old Man on His Back preserve have herds meeting this description. Establishing a number of these satellite herds will likely be a key component in a North American recovery strategy for the bison. If we are successful in expanding this herd to 2,000 animals in the first decade of the project, it will be the only genetically pure herd in the Great Plains, the historic stronghold for the species, that comfortably exceeds the estimated minimum viable population size of 400 (or 1,000 necessary for retaining 90% of allelic diversity) needed for long-term viability (WC’s population is currently about 375 animals), and one of the 3 or 4 most important conservation herds in North America. 10-year objectives Our 10-year objectives are to:

1. Establish a self-sustaining, naturally regulated, and ecologically effective population of bison that is free of cattle-gene introgression, semi-free ranging and subject to natural selective forces on and around APF lands in north-central Montana;

2. Establish a population that serves as a source of individuals for wild bison restoration throughout the region;

3. Establish a population that enhances the long-term survival of the species genetically, behaviorally, and ecologically and that promotes prairie conservation;

4. Establish a population that contributes to removal of wild bison from the Montana list of species of concern;

5. Establish a bison population capable of sustaining a variety of consumptive and nonconsumptive values and contributing to the cultural, aesthetic, economic, and social well-being regionally and nationally;

6. Collect and disseminate scientific information on reintroduction techniques and the ecological requirements for successful wild bison restoration;

7. Collect and disseminate scientific information on the ecology of bison; and 8. Contribute to restoring and maintaining natural ecological processes and native

biological diversity in north-central Montana.

Premises Following the recommendations of Boyd (2003a), we will emphasize the wildlife conservation value of bison and the differences between wild and domestic bison. We view this project as a wildlife reintroduction and as such will follow all appropriate strategies and protocols recommended for wild species reintroductions, recognizing


limitations to this model imposed by husbandry practices and legal constraints. We recognize in the short term the need to plan and manage bison as semi-domestic and confined, but our long-term goal is for a naturally regulated free-ranging population of wildlife. With respect to our premise, our primary goal will be to restore a population of bison that is ecologically effective (Pyare and Berger 2003) over a large landscape. Such a goal is very much different than managing bison for production, or in small confined areas, or in the traditional livestock management paradigm. As a highly interactive and foundation species, bison played a significant role in maintaining the historic abundance and diversity of plains biota. Bison grazed heavily in some areas and lightly in others, thereby creating a mosaic of vegetation (Hartnett et al. 1996; Knapp et al. 1999), which influenced not only the plant community, but a diverse suite of animals (Cody 1985; Clark et al. 1989). In contrast, western ranges in the United States, though relatively resistant to homogenization, show similar trends in management with the traditionally ideal pasture being uniformly grazed by cattle and free of livestock predators and competitors (Fuhlendorf and Engle 2001;Truett et al. 2001; Freilich et al. 2003). Livestock management, which has provided the general model from which modern bison management evolved, was designed to produce maximum sustained yields of a single species for human consumption, usually the domestic cow. For a number of reasons, this model is not always compatible with biodiversity conservation (Truett et al. 2001; Freilich et al. 2003). The model we propose will deviate from conventional range and livestock management in the following ways:

1. Livestock production goes primarily to feed people. The growth of human numbers and demands for food make it impractical for commercial producers to focus on restoring the historic diversity and abundance of predators, scavengers and decomposers. The annual increase of livestock, including bison, now is harvested for consumption off the range, removing the once-tremendous protein base consumed in place. Where possible, it would be desirable to allow some production to cycle through the system in place by allowing some natural mortality and carcass deposition.

2. Managers commonly attempt to remove species that reduce forage available to livestock. For example, removal of woody plants has been an ongoing focus of range managers in and near the Great Plains for several decades (e.g., Stoddart and Smith 1955; Vallentine 1989) because under some circumstances woody vegetation reduces forage production. Prairie dogs have been controlled because they have been viewed as competitors for forage (Stoddart and Smith 1955:201). Alternative ways of managing prairie dogs (Coppock et al. 1983; Detling 1998) offer fertile areas for biodiversity restoration.

3. Managers usually try to avoid heavy grazing and severe soil disturbance. Because the historic occurrence of these phenomena over large areas reduced the long-term ability


of many landscapes to support cattle, their prevention ranks among the most sacred tenets of good range management. However, some rare native plants (e.g., blowout penstemon, Penstemon haydenii) (Stubbendieck et al. 1997) and animals (e.g., black-footed ferret, mountain plover, and other prairie-dog-ecosystem species) (Koford 1959; Knowles et al. 1982; Knowles 1986) are associated with sites considered overgrazed by conventional standards. Management for these and some other “disturbance” species will require balancing their needs against potentially undesirable side effects of heavy grazing and disturbance, such as encroachment of invasive exotics.

4. Managers strive for uniform cropping of the forage base. Managing grazing animals on western ranges was built on the use of water and salt distribution, fencing, and sometimes herding to achieve spatially uniform utilization of forage (Stoddart and Smith 1955:324, Coughenour 1991). As this review suggests, the concept of uniform utilization ignores habitat needs of some species that rely on lightly and heavily grazed grasslands. Because spatially patchy grazing is normal behavior for grazing animals, capitalizing on this tendency can be a prime management tool for enhancing biodiversity.

In sum, development of habitat mosaics, perhaps on a smaller scale than occurred historically, is the key to present-day conservation of biodiversity. Historical accounts suggest that patch size in the Great Plains grassland mosaic in the early 1800’s often may have been rather large. Prairie dog colonies sometimes extended unbroken for miles (Knowles et al. 2002) and forage sometimes had been depleted by bison over similarly large areas (Hart 2001). Prairie fires probably created a larger-scale mosaic than would have occurred with grazing alone (Vinton et al. 1993), and recurring fires left vast areas devoid of woody plants (Wells 1965).

To “pack” more species into smaller acreages than was the historical norm, patch size can be reduced by applying the same historical patch-makers, but on a smaller scale. Strategic placement of water (which may itself attract many species) and fences in mid- and tall-grass areas can induce bison to create a mosaic by grazing heavily in some areas and lightly in others. In such situations relatively small prairie dog colonies can be maintained in heavily grazed sites without fear of their spreading rapidly into areas where the grass is denser and taller (Snell and Hlavachick 1980; Licht and Sanchez 1993). Controlled application of range fires on a small scale can help manipulate grazing distribution and at the same time preserve windbreaks, shrub thickets, and other patches of woody plants critical to the persistence of many animals. Fire can be used to maintain shrub savannas (McPherson 1995), which may attract a variety of wildlife species (Nelson et al. 1999) not found in shrubless grasslands.

METHODS Study area


Bison will initially be released on the APF Weiderrick property (20,000 ac; 8,000 ha) in southern Phillips County in north-central Montana (47o 40’ – 47o 55’ N, 107o 35’ – 108o 30’ W; Fig 1). The release area is bounded on the south by the CMR along the Missouri River. The Montana Glaciated Plains is a mixed grass prairie with sagebrush flats bordering the southwestern edge of the prairie pothole region. Predominant vegetation includes big sagebrush (Artemesia tridentata), silver sagebrush (Artemesia cana) greasewood (Sarcobatus vermiculatus; total shrub canopy coverage ranges form 0-20%), yellow sweetclover (Melilotus officinalis; total forb canopy coverage ranges form 5-20%), green needlgrass (Stipa virdula), western wheatgrass (Agropyron smithii; total grass canopy coverage ranges from 40-80%) and blue grama (Bouteloua gracilis). Active black-tailed prairie dog colonies contain variable amounts of bare ground interspersed with sparse vegetation that included fringed sagewort (Artemesia frigida), plains prickly pear (Opuntia polycantha), blue grama, needle-and-thread grass (Hesperostipa comata), and Sandberg bluegrass (Poa secunda). Grass forage production ranged from 215 - 751 kg/ha from 1976-1986 (Hamlin and Mackie 1989).

Figure 1. Site Location Map.


Figure 2. Site Detail and Proposed Pastures. Mean annual precipitation is 33 cm, most of which falls from May to July (Hamlin and Mackie 1989). Periods of drought interspersed with periods of high moisture are the rule (Hamlin and Mackie 1989). Mean annual temperature is 6.5 C and ranges from –8.4 in January to 20.8 in July. Snow depths exceeding 0.3 m are rare, though occasionally accumulations up to 0.9 m persist for significant periods. The average frost-free season is 128 days. The topography consists of flats cut by intermittent drainages, including Telegraph Creek, flowing to the Missouri River, and rolling hills. The primary topographic feature is the Missouri River that cuts through the area and creates the associated “breaks.” The breaks near the river are lightly to moderately forested with ponderosa pine (Pinus ponderosa). Scattered and extensive stand of cottonwood (Populus deltoides), willow (Salix spp.), wild rose (Rosa arkansana) and snowberry (Symphoricarpus occidentalis) occur in most drainages. The mean elevation is ~930 m. Soils of the area are primarily derived from the Bearpaw shale formation and are predominately heavy clay loams of the Lismas-Thebo series, with moderate amounts of salt. The soils are relatively impermeable to water and runoff is high. The Missouri River provides permanent water and Telegraph Creek runs intermittently during spring and following heavy rainfall. There are several man-made stockwater


impoundments, which, due to clay soils that impede surface infiltration, often remain full throughout the year. Current land use consists primarily of livestock grazing, with some crop production in bottomlands such as along Telegraph Creek. Ninety percent of the MGP remains in grasslands. Approximately 51% of the MGP is public land (about 41% of Phillips County is public land [US BLM 1992]), primarily BLM and CMR. The area is occupied by numerous species of mammals including pronghorn, elk, coyotes, cougars, bobcats, and prairie dogs. Biological feasibility Reasons for bison absence from area Bison were extirpated locally and throughout the Great Plains through unregulated hunting (McHugh 1958, Isenberg 2000). Disease resulting from import of European cattle possibly also contributed to their demise (Flores 1991; Isenberg 2000), as would habitat loss had bison survived in large numbers into the last century. As recent evidence suggests, forced interbreeding with domestic livestock and inadvertent introduction of those progeny to conservation herds is the only recently recognized threat. Availability of suitable habitat Distribution of prehistoric bison-kill sites dating to the last 10,000 years indicates a relatively high abundance in this area of Montana (Lyman and Wolverton 2002). One of these sites sits on the southern boundary of the APF property. Historical records, including data from the Lewis and Clark expedition, indicate that this was a region of particularly high bison abundance. More bison were encountered along this stretch of the expedition than anywhere else on their journey (Martin and Sutzer 1999, Lyman and Wolverton 2002; Laliberte 2003). Similarly, Maximilian (1843) reported that on 27 July, 1833, two members of his party, “Messrs Mitchell and Cuthbertson … had seen at a distance the Little Rocky Mountain range, like blue clouds. In the green extensive hollow towards the mountains, they saw the whole prairie covered with herds of buffaloes” in what was likely Sun Prairie or the Fourchette Creek drainage. Messiter (1890) reported on bison numbers between the Milk and Missouri Rivers in 1862: "...as we approached the Missouri they were in good-sized bands and toward evening one day, we saw an immense number of them in the distance." And as recently as 1875, Ludlow (1876) reported that on September 15, on the prairie north of Carroll, Montana (15 mi. east of Robinson Bridge on S. bank Missouri R., ca. Hutton Bottoms, flooded by the Ft. Peck dam), that about 3 or 4 miles out he “… saw 2 or 3 herds of buffalo… grazing in sight" and that "buffalo chips were abundant.” However, over a brief span bison were exterminated over most of their range (Hornaday 1889), including the project area. Bison were no longer reported in the project area by 1884, and cattle replaced bison about that time in the region and have been present since. Ranched bison currently occupy this habitat approximately 50 km from our restoration area. The Ft. Belknap Indian Reservation bison herd is approximately 100 km northwest of our area. We will work with these and other neighbors raising bison to minimize likelihood of escape into our herd.


Minimum area requirements To restore a naturally regulated semi-free ranging herd (restricted from moving onto neighboring private lands) of bison that is largely under natural rather than artificial selection will require a very large landscape. In part, we have designed this project to learn more precisely how large an area is required for this to occur, given that the historic record provides little evidence of bison migration and habitat use. Bison on the northern range of Yellowstone, in the Henry Mountains of Utah and the interior of Alaska and Canada are largely naturally regulated and semi-free ranging and under primarily natural selection. Landscapes in these areas range in the hundreds of thousands of hectares. Approximately 600 bison in the Theodore Roosevelt National Park in North Dakota are confined to 18,000 ha (46,000 acres) of river breaks habitat similar to that along the Missouri River in the project area and thrive there (Norland 1985). To reach the goal of a minimally viable population of >400 bison will conservatively require approximately 20,000 ha (50,000 acres). Elimination of threats Hunting Unregulated hunting no longer occurs in the region. Legally, bison managed by APF are, for the present, considered the property of APF, and therefore not subject to regulation as wildlife. Illegal poaching, as with other trespass, will be controlled by patrolling by APF ranch personnel to ensure poaching is not a problem.

Disease Although bison populations have on occasion been reduced due to disease outbreaks, direct mortality from disease is less a threat than herd culling intended to limit the spread of diseases carried by bison to domestic livestock (e.g., brucellosis, see Boyd 2003a). Bison released on the project area will be certified as disease free and will be monitored and managed as described below to ensure they remain disease free.

Habitat loss and genetic diversity The primary objective of the project is to acquire access to sufficient grazing capacity to restore a bison herd of suitable size and genetic composition to maintain itself as self sufficient within the existing carrying capacity. BLM and CMR lands in the project area cannot be tilled under current federal regulations and thus will be maintained in a prairie cover suitable for bison. APF will not till any additional land it owns and will work to maintain and restore native cover to cultivated land it now owns thereby reversing habitat loss that has occurred in the region. It is likely that, once established, the herd would be integrated into metapopulation management for a North American plains bison recovery strategy, thereby contributing to maintenance of the plains bison genome. Significance of cattle DNA introgression For endemic or threatened species, the negative consequences of interspecific hybridization can be both biological and legal. Interspecific hybridization can result in the disintegration of genetic integrity and the loss of native genetic variation or locally adapted gene complexes (genetic swamping; Ward et al 1999). In addition, the presence


of hybrid animals in remaining populations of threatened species may result in legal challenges to their protected status (O’Brien & Mayr, 1991; Hill, 1993; Rhymer & Simberloff, 1996). Consequently, interspecific hybridization has become an increasingly important issue in conservation biology. Since bison and domestic cattle do not naturally hybridize, and there are clear negative fitness consequences in at least the F1 generation, it seems plausible that the introgression of domestic cattle genes into bison germplasm might also be under negative selection. The maintenance of introgressed domestic cattle regions of the DNA for 15 – 20 generations post-hybridization in the federal populations examined by Halbert (2003) suggests that any negative fitness effects, at least in these regions of DNA, is minimal. It is worth noting that the probability a deleterious allele will be purged from a population is correlated with the size of the population. Only alleles under strong negative selection will be able to overcome the effects of genetic drift in small populations. The location of genes and their respective functions within and near the 15 nuclear regions examined by Halbert (2003) are largely unknown; it is therefore not possible at this point to directly investigate the involvement of natural selection on domestic cattle introgression in these regions. Previous reintroductions There have been numerous projects to “reintroduce” bison to locations throughout North America (Boyd 2003). Most of these have been to relatively small areas with no intent to conserve bison genetics or restore bison ecological effectiveness to large landscapes like we propose. Boyd (2003) outlines in detail the history of most of the public bison populations in North America. Source herd evaluation Of the 10 federal herds examined by Halbert (2003), only 3 herds (Yellowstone, Wind Cave and Grand Teton National Parks) were identified as free of introgression of cattle genes. Herds at Yellowstone (YNP), Neil Smith National Wildlife Refuge, and Wind Cave (WC; in that order) had the greatest overall allelic diversity. Yellowstone and Wind Cave herds contained the majority of genetic variation in the herds examined. As such, Halbert (2003) concluded that these populations should be given conservation priority and be maintained in isolation from those populations identified as containing domestic cattle introgression. Since both the YNP and WC populations contain high levels of genetic variation and no evidence of domestic cattle introgression, consideration should be given to starting satellite herds using stock from these populations. The establishment of such satellite herds from WC would be considerably easier than from YNP simply due to the brucellosis-free status of the WC population. The maintenance of satellite herds in this manner will help ensure the future preservation of pure bison germplasm. Castle Rock, New Mexico The only known private herd shown to date to be pure bison with no cattle gene detected is the Turner-owned Castle Rock herd (CR) on the Vermejo Park Ranch (VPR) in northeastern New Mexico (J. Derr, Texas A&M University, unpublished data). Although


the full history is not completely clear, it appears CR animals originated from Yellowstone stock with some genetic influx from bulls from the Maxwell herd in Kansas (one of the 6 founding herds for all bison; D. Hunter, Turner Enterprises, unpublished report). The herd started with 5-8 bison already at the Valle Vidal (nearby unit of Carson National Forest) when Mr. Gourley purchased the VPR in 1945. These few bison were believed to have been moved to VPR with elk that were captured and moved from YNP sometime between the 1920s and 1940s. Thirteen additional bison were purchased from the nearby Philmont Scout Ranch in the 1950s and released with the CR bison. The age structure and composition of the herd from that period was unknown. The Philmont herd was originally founded from bison purchased from YNP in the early 1920s. There are approximately 257 bison in the herd at CR. The herd has 126 cows, 70 bulls, 21 yearlings and approximately 60 calves. The DNA of every adult in the herd has been documented (J. Derr, Texas A&M University, unpublished data). The intense culling of the YNP herd until 1967 removed three/fourths of the herd, leaving only 397 bison and possibly resulting in some loss of genetic diversity (J. Derr, Texas A&M University, pers. comm.). The CR bison were purchased before the large culling events in YNP and the likely loss of some alleles. The only impacts on the genetic pool of the CR herd were due to the process of natural selection, genetic drift, selection for predator (bear, cougar, and coyote) avoidance, and limited hunting by ranch employees. These animals should hold a large proportion of the genes (4/6 foundation herds are represented) that are found in North American bison before European settlement of the West (pre-1850) and possibly a greater diversity of the historic genes than are incorporated in the YNP bison herd. Bison at Castle Rock live on mixed-grass ponderosa-pine forest-edge habitat. The bison have not been supplementally fed, are semi-free-ranging, and are naturally regulated. Wind Cave National Park, South Dakota In 1913, 14 bison (6 bulls and 8 cows) were donated to the Wind Cave National Game Preserve by the New York Zoological Society as part of an effort by the American Bison Association to return bison to the Great Plains. Six additional bison (2 bulls and 4 cows) were brought in from Yellowstone National Park in 1916. The present Wind Cave bison herd originated from these 20 animals (Boyd 2003). The population is approximately 350-450 animals on about 10,000 ha (25,000 acres). In 1986, the WC bison population was released from quarantine by South Dakota and no animals have tested positive for brucellosis antibodies since (Barbara Muenchau, NPS pers. comm. 2004). Vaccination was ended in 1988. In an attempt to maintain a 50:50 sex ratio in the younger age classes, annual roundups are conducted and yearling bison are culled from the WC population. Ten yearling bison of each sex are withheld in the park, producing a herd representative of all age classes (Barbara Muenchau, pers.comm. 2004). Excess bison are typically transferred to various American Indian tribes. Since resuming the live distribution of bison in 1987, 820 (98%) of the bison culled from the herd were distributed live to various Native American Tribes. Ward et al. (1999) examined 37 WC bison and found a single bison mitochondrial haplotype and no


evidence of domestic cattle mitochondrial introgression (375 bison sampled, 0.9999 probability of introgression not >0.01% in the herd; Ward 2000, Halbert 2003. Bison at Wind Cave live on mixed-grass ponderosa-pine forest-edge habitat, are fenced, and coexist with cougars and coyotes. WC has expressed interest in working with us to use our reintroduction site as a satellite herd for managing genetic diversity of WC bison. Gross and Wang (2005) have indicated that managed bison populations of <400 animals are at risk of loss of genetic diversity. One way to retain that diversity would be to start another herd using WC animals. We are interested in helping WC meet this goal. The best information to date indicates, however, that to meet this objective we would only be able to use WC bison to start and supplement the reintroduction herd. We are currently assessing if this may too severely limit our management options and our ability to meet our objectives. Elk Island National Park, Alberta, Canada Elk Island was the temporary home in 1907 for the first shipments of the Pablo Allard herd purchased by the Canadian government from Mr. Pablo of Montana (Coder 1975; Boyd 2003). These bison descended from 4 yearlings captured by Walking Coyote and from 26 geographically diverse bison from Buffalo Jones. All but 48 of these bison were removed late in 1909 and taken to Fort Wainwright, Canada. The remaining 48 experienced exponential growth, and culling began in the 1920s. The population reached 2,479 in 1936. Systematic culling began in 2000 to retain the population on 14,000 ha (35,000 acres) at around 500 animals. The herd has been genetically tested and no cattle introgression has been detected introgression in the herd (Ward 2000;Halbert 2003). Excess bison are available from this population for translocation, and this herd would serve as an excellent source except for current import restrictions in place between Canada and the U.S. According to the Montana Department of Livestock and the Animal and Plant Health Inspection Service of the U.S. Department of Agriculture, it is not possible to import live bison from Canada at this point in time due to fears associated with Bovine Spongiform Encephalopathy (BSE). Henry Mountains, Utah The free-ranging and minimally managed herd on approximately 1,000 mi2 (2,590 km2) of BLM lands in south-central Utah originates from 18 animals from YNP released in 1941 (Boyd 2003). The herd is currently approximately 279 animals managed through hunting (Van Vuren 1982, Van Vuren 2001). Samples from harvested animals indicate it is disease free and has no cattle introgression (Ward 2000). Bison at Henry Mountains live in pinyon pine and juniper habitat and occur with cougars. Source herd recommendation CR bison are semi-free-ranging, naturally regulated, little handled, have no history of brucellosis, and may have the greatest genetic diversity of the potential source herds. The genetic history, however, is clearer with the WC than the CR herd. We will likely have a greater selection for the age and sex composition from the CR herd than from the WC herd. While the Henry Mountain herd has similar features, capture of these animals for translocation will be more difficult. We are continuing to assess optimal strategies for


managing genetic diversity both to meet APF and WC objectives. WC animals and CR bison have been offered and are available in fall 2005. Our primary genetic goal is to obtain a representative amount of the genetic diversity present in the plains bison subspecies. Therefore, the usefulness of obtaining bison from more than one source population will depend on the levels of diversity within the 2 sources and the amount of unique genetic diversity present within the source herds, as can be measured by a genetic distance value. We will assess if it is desirable to obtain animals from both WC and CR herds by determining if the diversity contained within one is different than that in the other (i.e. they each contain a number of unique alleles). If the histories of these populations are different (as would seem to be the case), we would expect to find differences in their gene pools, and therefore obtaining animals from both populations will be important for meeting our goal. Optimal release strategy modeling We used VORTEX (Miller and Lacey 1999) to model the success of various reintroduction strategies to compare against range carrying capacity and release-area size and to assess long-term population viability. The simulation program models deterministic and stochastic processes based on a variety of user-defined variables. We generally followed the model used by Wilson and Zittlau (2004) for wood and plains bison in Elk Island National Park (EINP), Alberta for defining input variables. They describe the rationale for selecting particular input variables. We modeled different reintroduction strategies to maximize population performance (Haig et al. 1990). For the short-term assessments we modeled populations at 2-year intervals over 20 years using 100 simulations for each model run (Appendix 1). For long-term assessments, we modeled populations at 5-year intervals over 100 years using 100 simulations for each model run and a carrying capacity of 5,000 animals. Parameters Minimum founding size The founder population is important because it should represent the natural genetic variability of bison. Lacy (1994, cited in Morrison 2002) suggests that this can be accomplished with a relatively small initial population size. For example, an initial population of 50-100 individuals should represent about 97-99% of the expected heterozygosity in the source population if the chosen animals are unrelated. Additionally, the larger the founder population, the greater the probability of including rare alleles. Alleles with a known frequency of 0.001 in the source population will, theoretically, have a 10% probability of inclusion in a founder population of 50 animals and increases to 19% if 100 animals are released (Lacy 1994, cited in Morrison, 2002). This suggests that genetic testing of the released animals should be undertaken to minimize accumulation of closely related individuals and to have a clear understanding of the founder population genetic status for future management scenarios. The Pink Mountain bison population in British Columbia, founded in 1971 from 48 individuals from EINP, appears to contain a representative amount of the genetic


variation present in its parent population, judging from genetic distance data (Wilson and Strobeck 1999). The population is now comprised of 1,000 animals in 2,000 km2. This number of founders may then be a minimum level to aim for when new bison populations are started (Wilson and Strobeck 1999). Forty-eight animals also founded the EINP population. In order to retain > 95% of the heterozygosity and allelic diversity found in the source population, Leberg (1990) recommended > 25 individuals (1/2 males and 1/2 females) in founding populations. To retain >95% of the heterozygosity and allelic diversity found in the source population, we will reintroduce 8-20 young (< 3 year-olds 50% female:male ratio ) bison in year 1. While a greater percentage of females would result in a greater rate of population increase, more males may allow better retention of genetic diversity because more males may be able to breed when they are all young as opposed to later when a male dominance structure develops. We will supplement an additional 28 (same sex and age characteristics as year 1) bison in year 2 and year 3. Given anticipated mortality rates, this should ensure that we have >50 founders in the population. The female-weighted sex ratio should maintain more of the imported genetic diversity as research indicates that only dominant males reproduce. Thus, the genes of subordinate males are lost after the first generation. Decreased genetic variability resulting from inbreeding depression leads to negative consequences such as lowered fecundity, reduced growth, and greater susceptibility to disease (see review by Lacy 1997). Although the reintroduction of 50 unrelated animals should prevent inbreeding, the herd may also be managed to reduce the impact of inbreeding simply by ensuring the introduction of one effective (i.e., actively breeding) individual into the population every generation. Minimum viable population size Wilson and Zittlau (2004) assessed management strategies for minimizing the potential negative effects of inbreeding. One of the management goals for EINP was to retain 90% genetic diversity over 200 years. As they indicated, this goal is only based on the minimum population size necessary to reduce the risk of inbreeding. Higher levels of diversity may be required for a population to adapt to environmental changes. Furthermore, they reasoned that the diversity found within the bison populations at EINP may not be easily replaced from an outside source. Therefore, a second management goal of maintaining 90% diversity over 500 years was also examined. They believed that the values suggested for meeting these goals should be considered lower limits, and not necessarily values to aim toward. Also, it is important to note that these goals describe the maintenance of 90-95% of current levels of diversity and not absolute levels of genetic diversity. The minimum population size required to retain 90% of their genetic diversity (heterozygosity) over 200 years was 175. For the more conservative goal of retaining 90% diversity for 500 years, 430 plains bison are required. The current managed size of EINP herd will retain 96% diversity in plains bison over 200 years if harvesting strategies target an equal proportion of young males and females every second year. These are


reasonable levels of genetic diversity to retain according to most recommendations (Leberg 1990; Wilson and Zittlau 2004). There are currently no quantitative NPS or US Fish and Wildlife Service management objectives for conserving genetic diversity. Gross (2000) used a goal to achieve a 90% probability of maintaining 90% of the selectively neutral genetic variation for 200 years, following recommendations by Soule et al. (1986). This goal is consistent with U.S. Bureau of Land Management operational guidelines for wild horse management (Coates-Markle 2000). If retention of H0 is the primary aim of management, simulations of Gross and Wang (2005) suggest that a population objective of about 400 animals is likely to achieve a goal of retaining 90% of currently existing H0 if management perfectly followed the simulation model. Since management can seldom be so precise, a larger population is probably needed to meet this goal. A much larger population objective – on the order of 1000 bison - is required to achieve a reasonable assurance of retaining 90% of currently existing alleles (Gross and Wang 2005). However, given the need for bison reintroductions to new areas and for bison to adapt to large current (e.g., exotic diseases) and future (e.g., climate change) alterations in their habitats, as well as the intrinsic value of conserving genetic diversity, we believe the goal should be retention of at least 95% of allelic diversity over 200 years. As summarized by Gross and Wang (2005) “In evolutionary terms, H0 is an index to the overall degree of genetic variance at a locus and it would be expected to reflect the magnitude of short-term responses to artificial or natural selection (James 1971). High allelic diversity will virtually always be correlated with the occurrence of many alleles that have a low frequency in the population. These rare alleles are unlikely to contribute substantially to short-term population responses to selection, but they can be a very important limit to the response to selection over many generations (James 1971; Allendorf 1986). Allelic diversity is thus considered important to the long-term survival of a species, especially where there may be substantial environmental changes, range expansions, or (re)introduction into new sites.” In a more general approach to ensuring evolutionary potential and long-term conservation of populations Franklin and Frankham (1998) and Lynch and Lande (1998) argued that minimum wild population sizes need to be in excess of 5,000. Thus, the rationale for obtaining as large a habitat block as is feasible to support additional animals. Population demographic and behavioral inputs for VORTEX models We generally followed inputs of Wilson and Zitlau (2004) for developing our models. We defined mature individuals as females aged 3 years or older, and males aged 6 years or older, as based on the EINP census data, as well as from the common age of maturity reported for other bison populations (Females: McHugh1972; Fuller 1966; Halloran 1968, Meagher 1973, Haugen 1974, Reynolds et al. 2003, Green and Rothstein 1991; Males: Mahan 1978, Maher and Byers 1987, Wilson et al. 2002). For the short-term projection, we conducted an extra simulation where we defined males as mature at 3 years of age and allowed all males to breed due to the small number of young animals and little competition among males. Senescence in our model occurs at 20 years of age.


By modeling reproduction throughout an individual’s lifetime, the complete age distribution can be incorporated into the model. VORTEX assumes that all individuals of reproductive age have an equal probability of breeding. Each year, breeding males are paired with breeding females. For wood bison, Wilson et al. (2002) found that approximately 40% of male bison successfully sired offspring in an average year. In Badlands, 48% of matings were attributed to 11% of males from 9-15 years old (Berger and Cunningham 1994). We set 40% of males as breeding. For female wood and plains bison, the average calving rate from 1991 to 2001 in EINP was determined to be 75% (W. Olson, EINP, pers. comm.). Calving rates reported in other bison populations range from less than 50% to 100% (Halloran 1968, Meagher 1973, Haugen 1974, Lott 1979, Rutberg 1986, Lott and Galland 1987, Gates and Larter 1990). The rate for bison on Turner Ranches in the northern Great Plains ranges from 80-90% (M. Kossler, Turner Enterprises, personal communication). Low rates (<60%) are typically reported under poor forage conditions. Variance in calving rates due to environmental stochasticity was again based on calving rates at EINP and was set at 7.5 for plains bison. The primary sex ratio, calculated from 35 years of population structure data from EINP, was 55% males in both the wood and plains bison populations (W. Olsen, EINP, pers. com. 2003). Typically, sex ratios at birth are around 50%, with a slight bias toward males (Fuller 1966, Halloran 1968, Haugen 1974, Rutberg 1986) The ratio can vary with density and range productivity (Rutberg 1986, Gates and Larter 1990). We set the ratio at 50%. A comparison of census counts between consecutive years provided a rough estimate of annual mortality for EINP. Due to the small sample sizes, standard deviations were high. As the estimates were not well supported, a range of mortality rates (3% to 20%), obtained from published literature, was tested for each age- and sex-class of each herd by Wilson and Zitlau (2004) and we used their final selected values. Mortality rates range from 2% to 45% in calves (Gates and Larter 1990, Berger 1992, Kirkpatrick et al. 1996), 3% to 8% in juveniles (Fuller 1966, Berger 1992), and 1% to 3% in adults (Gates and Larter 1990). Mortality rates for adult bison in young age structure herds on Turner Ranches is approximately 4% (M. Kossler, Turner Enterprises, personal communication). Reports of high mortality rates are associated with herds that experience predation. The EINP populations are not commonly subjected to predation. Although bison populations are in most cases likely density dependent (Lartner et al. 2000), we modeled as density independent because at small populations forage should not be limiting. Modeling results


For the conservative simulation (BISCON) where males first breed at age 6 and then only 40% breed, we have 35 bison at year 2 and 61 in year 3 (Fig. 2). The bison population increased at r = 0.226 (0.331) over 20 years (Fig. 2). Bison reached target population size for the APF property and associated allotments of 165 (based on available AUMs — see discussion below) by year 10. Bison reached threshold for long-term population viability (>500) by year 18 and met genetic heterozygosity goals. For the more liberal simulation (BISLIB60), the bison population reached 39 at year 2 and 71 at year 3. The bison population increased at r = 0.229 (0.321) over 20 years. Bison reached target population size at year 9. Bison reached threshold for long-term population viability (>500) by year 18 and met genetic heterozygosity goals. Bison reached a population of

Figure 3. Bison Population Growth Estimates Based on Founder Inputs. 3,000 animals at year 30 and 5,000 animals at year 35. Comparison with other populations Eighteen bison were reintroduced into the Mackenzie Bison Sanctuary of Northwest Territories Canada in 1963. The population peaked at 2,400 in 1989. Rates of increase ranged from –0.05 – 0.45 depending on density. Bison increased on the National Bison

Bison Population Growth Estimate from Vortex

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Range from 37 in 1909 to 554 in 1922 (Fredin 1984). The rate of increase for any bison population depends on the population’s age structure, sex ratio, latitude, and presumably forage quantity and quality and mortality rates. In relatively unmanaged herds like Yellowstone with severe winters, high densities of predators, and older population structure, productivity rates are low (20 calves/100 cows). Where bison are managed for higher productivity by reducing carrying capacity and altering age and sex ratios, productivity is closer to 50 calves/100 cows. Additional justification for release strategy Our approach to bison reintroduction starts with a small number of animals in year 1 and phases in releases over 3 years for several reasons. Eight animals released in year 1 in a small pasture provide our team with experience handling, managing, and assessing impacts of bison in the area on a reasonable scale. It also ensures limited donor population impact in any 1 year. We are proposing to use young animals because of their greater ease in handling and managing and because of the opportunity to “imprint” younger animals on the release site so that they wander less and will “habituate” and likely learn to use the area in a fashion most appropriate for the area rather than as a result of “lessons” learned elsewhere (D. Campbell, Brokaw bison mgr., pers. comm. 2004; C. Gates, University of Calgary, pers. comm.). We are requesting an even sex ratio as this is likely the sex ratio of wild populations and to ensure slower herd growth as we start the project. In addition to selecting bison based on genetic and disease criteria, it has been hypothesized that social organization may be an important consideration for bison restoration (F. Provenza, Utah State University, unpublished data). As outlined by Provenza (Utah State University, unpublished data), social organization is important for securing protection from predators (Carbyn et al. 1993, Sherman 1977), obtaining social status (Holekamp and Smale 1991, Smale et al. 1993, Brookshier and Fairbanks 2003), improving reproductive success (McComb et al. 2001), learning traditional routes (Tulloch 1978, Festa-Bianchet 1991), and sharing feeding sites (Johnson 1986, Kojola 1989). Social organization is also important because social interactions affect foraging behavior, which may influence how bison use landscapes and how biophysical environments respond to their use (Provenza, Utah State University, unpublished data). For example, in sheep and cattle, social interactions within groups encourage animals to eat a broader array of foods (Scott et al. 1995) and to forage in a greater variety of locations (Howery et al. 1998) as individuals maintain the cohesiveness of the group and respond to ever-changing preferences of individuals within the group (F. Provenza, Utah State University, unpublished data). These outcomes are important ecologically because bison reared as families may graze in ways that impact landscapes, especially riparian areas in ways more aligned with long-term historical impacts (Bob Jackson and Sharon Magee, personal communication, Tall Grass Bison, Iowa). We recognize the potential value of using social groups for reintroductions and will work to that end, but recognize that may not be possible due to donor and handling constraints


in this initial reintroduction. The possible importance of social organization notwithstanding, we believe that the benefits of starting a reintroduction with young animals for the reasons outlined above may outweigh the possible benefits of starting with socially intact and mature groups. Additionally, we believe that the young animals we introduce will develop the necessary social organization on their own in a way matched to this landscape. We will monitor social organization in these animals and its relationship to environmental factors and adjust our reintroduction strategy if it appears that young unrelated animals are faring poorly in this landscape. Habitat Suitability Number of bison area can support Estimating liberally, a 454 kg (1,000-lb) cow bison will eat and trample about 36 air-dry pounds of forage per day (Holecheck 1988; Steuter 1997), or 4,904 kg (13,140 lbs) annually. A 270 kg (600-pound) yearling will use about 18 pounds. Grass forage production in the area ranged from 215- 751 kg/ha (190 – 662 lbs/acre) from 1976-1986 (Hamlin and Mackie 1989). Carrying capacity (animal unit months; AUMS) for cattle in the area has been determined by the BLM. The generally accepted rate for converting carrying capacity for cattle to bison is 1:1 (See, e.g., USDA 2003; US BLM 1997). The current number of AUMs and the permitted use for the Telegraph Creek Allotment by pasture is listed in Table 1. APF proposes utilizing pastures 1-4 and additional Base Property, possibly introducing 8-20 bison into a smaller area (Pasture #3 or #4) of the allotment (Fig. 1) initially. Year-round carrying capacity of this pasture is estimated by BLM to be 103 AUMs. In years 2 and 3 we would incorporate additional pastures, increasing to 5,600 ha (14,000 acres) (Fig. 1) and a target population of 165 bison using approximately 2100 AUMs. Table 1. Telegraph Allotment

Pasture No. Ownership Acres TOTAL AC Ownership AUMs TOTAL AUMS

BLM State Private CMR BLM State Private CMR 1 2642 2642 393 393 2 3455 3455 472 472 3 836 640 1340 2816 129 96 111 336 4 3447 3447 353 353 5 870 1638 360 2868 170 313 47 530

With Base 1166 3380 4546 134 450 584 2668

Rangeland condition The most current assessment is that the range condition of the allotment is 1717 ha (4245 acres) in “good” and 2,944 ha (7275 acres) in “fair” condition (U.S. BLM 1992). The proposed change will result in a net reduction in the number of AUMs and therefore should result in an improvement of range condition from the following standpoints: 1) bison will likely graze intensively in some areas and less intensively in others, at least up


to the point where bison reach carrying capacity of the allotment, resulting in a wider distribution or mosaic of grassland stand conditions, from intensively grazed to lightly grazed, which is believed to mimic natural range condition (Fuhlendorf and Engle 2001); 2) the net reduction in AUMs will likely result in higher retention of residual grass and grass height in lightly grazed areas; and 3) it is likely that bison will graze less intensively than livestock in riparian areas and thus riparian condition will be maintained or improve. Ecologically effective minimum population size Similar to attempts to develop a working definition for keystone species (Mills et al. 1993), it will be difficult to develop a definition of an ecologically effective population. The primary issue is one of scale and magnitude. This is complicated by the fact that we know little about historic structure, ecology, and effects of bison over large landscapes. We believe, however, there are several characteristics that would typify an ecologically effective population. We believe such a population would exhibit landscape and autecology-driven robust demographic and behavioral patterns (Lartner et al. 2000), measurably shape vegetation structure and dynamics at a large scale (Knapp et al. 1999; Fuhlendorf and Engle 2001, Truett et al. 2001), would significantly promote population and behavioral responses in some mammal and avian species (Truett et al. 2001), and would serve as a focus of selection for large predators and scavengers and promote demographic responses in these (Carbyn et al. 1993; Larter et al. 1994). We believe populations on the order of and distributed over a landscape similar to YNP, Wood Buffalo, or the Mackenzie Sanctuary would constitute minimally effective populations. Thus, in order for this reintroduction to reach this goal, APF will have to significantly expand the range available to bison. Effect of bison reintroduction Impacts to special status species We expect the following special status species to benefit directly and indirectly from the change in use to bison. Because we are both reducing the existing stocking levels and introducing an indigenous herbivore, it is anticipated that the impacts to most of the species described here as special status (Table 3) will be either improve or have no impact. Impacts to black-footed ferrets and mountain plovers may not be as beneficial in the early stages since they rely on heavy grazing. Table 2. Effect of bison restoration on special status species in and near APF project area Species Federal State References as status status¹ to benefits


Bison (Bos bison) S2 Black-tailed prairie dog (Cynomys ludovicianus) CT S3 Truett 2003 Mountain Plovers S2 Knowles et al.1982 (Charadrius montanus) ¹ Carlson (2003) Availability of suitable release stock and impacts to donor populations Wind Cave National Park has assessed their population management needs and carrying capacity for the area and determined that surplus animals are available for translocation annually. Turner Enterprises has assessed the CR herd size and management goals determined that surplus animals are available. Socioeconomic and legal considerations We are working with CMR, BLM, Montana DNRC, and the South Phillips County Stewardship Council to develop guidelines related to reintroducing bison on federal grazing leases and allotments existing on CMR, BLM and state lands. Specifically, the BLM has initiated an environmental assessment (EA) that will be completed in early 2005 for the Telegraph Creek allotments to permit change in type of livestock and season of use. Lands within the allotments currently under state (DNRC) and CMR jurisdiction (grazing under an exchange of use for other APF-controlled capacity within the Refuge) will be addressed as part of this EA. Bison grazing is consistent with the management goals of the CMR and the Refuge System (“The mission of the System is to administer a national network of lands and waters for the conservation, management, and where appropriate, restoration of the fish, wildlife, and plant resources and their habitats within the United States…,” National Wildlife Refuge System Improvement Act of 1997, P.L. 105-57-Oct. 9, 1997, Sec. 4) and the Judith Valley/Phillips Resource Management Plan (JVPRMP; U.S. BLM 1992) and BLM Standards for Rangeland Health and Guidelines for Livestock Grazing Management for the Lewistown District (Standard #5…”Management for indigenous vegetation and animals is a priority.”). Bison reintroduction would possibly improve habitat for prairie dogs, thus benefiting federal recovery efforts for the endangered black-footed ferret. We are also working on the terms of an MOU with the South Phillips County Stewardship Council to establish protocols for fencing, herd health, and damages that will meet with general approval of neighboring landowners. We are collaborating with Peter Gogan of the U.S. Geological Survey who is overseeing the U.S. federal government’s modeling program for management of national park bison herds. We envision this new herd as a valuable component to any conservation strategy that is developed from the plan. The IUCN Bison Specialist Group has recognized this project as a top conservation priority for bison.


Qualifications of restoration team (including administrators, biologists, and veterinarian and/or animal caretakers) We have assembled an excellent team of collaborators that have a substantial base of experience designing and conducting reintroduction projects and will be able to apply those skills to this project. Our collaborators have led or participated in successful reintroduction projects of species including swift fox (Kunkel et al. 2003), prairie dog (Truett et al. 2001), black footed ferret (Forrest and Biggins 1988), and bighorn sheep (Krausman et al. 2001). WWF and APF personnel and cooperators will manage and implement this project. Curt Freese, WWF program director, will provide overall project supervision and liaise with the cooperating agencies. Freese, Steve Forrest, Cortland Barnes, and Kyran Kunkel will serve as collaborating project investigators and on-site project leaders. Dave Hunter of Turner Enterprises will serve as our consulting veterinarian and we will engage local veterinarian assistance. Scott Laird is APF field coordinator for the overall restoration project. APF will provide an on-site manager/biologist who will monitor bison on a daily basis. APF will hire additional field staff as needed. We are also working to enlist a science and review team (Appendix 7). Public planning and participation Part of the BLM/state EA process will require public meetings in the area to outline our plans and gather input from neighbors. We are in regular contact with our closest neighbors discussing our plans. We are developing MOUs on bison health and management with the Southern Phillips County Ranchlands Stewardship Group. We will hold 3-4 public meetings in the area in 2004 and 2005. We are also meeting with local resource sportsmen, conservationists, and livestock production groups. TRANSLOCATION Acquisition methods We will work with managers at source populations who have extensive experience handling and moving bison, assuring safe and humane treatment of all animals during the acquisition process. Transport will be contracted with a qualified livestock transporter. Dave Hunter of Turner Enterprises will serve as collaborating veterinarian during loading, transport and upon arrival at the project area, providing all oversight and protocols in cooperation with veterinarians and biologists responsible for source herds. We will develop animal care and use guidelines that will be approved by independent veterinarians and biologists (i.e., Kunkel et. al. 2003; Appendix 7). All bison will be disease tested and PIT tagged upon capture. We will test for BVD I and II, PI3, IBR, Anaplasmosis, Bluetongue and EHD. We will vaccinate with killed vaccine only. We will vaccinate with Triangle 4 with Type II BVD twice, 3 weeks between vaccinations, one dose of Alpha 7 (Clostridium sp.) We will worm with cydectin. A


sample of the bison will be radio-collared. We will clean weeds from bison prior to transport. Bison will be loaded onto appropriately-sized and equipped and disinfected transport truck/trailer and shipped immediately to the release site. The cattle trailer shall have had no contact with sheep prior to or during this translocation (including at truck stops, etc.). The driver will be educated on the potential risks should these bison come into contact with sheep on their journey to the release area. This is important in preventing any potential transmission of Malignant Catarrhal Fever, which is highly prevalent in sheep herds and is potentially devastating to bison herds. Disease and genetics screening Two conservation concerns arising from disease in bison are 1) the impact of the disease on the viability of the populations, and 2) the potential threat a diseased herd poses to adjacent commercial livestock and humans. Livestock diseases that may restrict trade or pose a risk to human health are designated as “reportable” under state and federal laws. We will follow all state and federal regulations related to reportable diseases. In general our herd health management philosophy will tend toward minimal management of animals while ensuring minimal risk to our neighbors’ cattle. We favor minimal management for 2 reasons. First, handling of animals increases stress on those animals and is inconsistent with general wildlife management objectives and philosophy. Second, native disease organism are a crucial ecological and evolutionary force in natural systems (Wobeser 2002). We consider brucellosis and tuberculosis (TB) to be exotic diseases and will manage against these for that reason and more importantly because they are reportable diseases and to minimize risk to our neighbors. Our overall disease management philosophy for diseases that are a concern and potential risk to our neighbor will be prevention and control. We will translocate only bison that are from certified disease free states and then we will vaccinate those animals as a further preventive measure and finally we will intensively monitor bison in the field to prevent and control diseases that may impact our neighbors. There are several requirements necessary to bring a bison from another state into Montana to meet state livestock health requirements. All female bison are required to be Official Brucellosis Calfhood Vaccinates (OCV), and must be vaccinated between the ages of 4-12 months. No test is required for adult animals, but we would recommend a negative 30-day test for bulls 12 months and older and vaccinated females 24 months and older. If the bison are from a TB free herd, the herd accreditation number and the date of the entire herd test must be included on the certificate of veterinary inspection (See MT official order 03-01-I). No TB test is currently required for bison entering Montana from South Dakota. A TB test (negative 60-day test) is required for animals originating in New Mexico (Department of Livestock interim guidelines, Dec 14, 2004). The third requirement is individual identification for each bison entering Montana. Each individual needs either an ear tag or a bison registration tattoo. Finally, a health certificate as well as a permit number is necessary for each bison imported into the state. A health certificate as defined in 81-2-702 is a legible record written on an official health certificate from the state of origin or an equivalent form of USDA attesting that the


animals have been visually inspected by a federally accredited veterinarian and found to meet the entry requirements of Montana. No additional disease testing is required once bison are in Montana. We will follow all these requirements and collect blood for complete disease profiles on all animals. Reference Administrative Rules of Montana 32.3.224 and 32.3.403. We will also use blood and hair samples from all bison released for DNA analysis. WC disease background issues Wind Cave is a minimally-managed, free-ranging herd. Because a brucellosis eradication program resulted in elimination of the disease from the herd in 1986, the park does not vaccinate. Direct population reduction through shooting in the field, and capture/disease test/slaughter of seropositive bison, led to the eradication of brucellosis in the Wind Cave herd. Since the release of quarantine in December of 1986 to present, 3099 additional bison have been tested for brucellosis with no positives. In 2001 a state law was passed allowing the transfer of unvaccinated heifers within the State of South Dakota as the state was declared a "Brucellosis Free State". At this time, the State of South Dakota is a Certified (Brucellosis) Free State and an Accredited (TB) Free State (SD Animal Industry Board 2004). We will vaccinate females calves from WC for Bangs as required by the Montana state veterinarian’s office. Calves will be vaccinated during a spring roundup or by immobilizing. Calves will also be vaccinated during fall roundup. CR disease background issues CR bison are brucellosis free. Female bison from CR have all received calfhood brucellosis vaccinate and a waiver is thus not required. Release methods We propose to "soft release" and thereby imprint a small number of bison on a small landscape to minimize the tendency of bison to wander. Hard releases of bison have demonstrated that these bison tend to wander widely. We will use a slow multi-year approach to build the bison population through reproduction and additional reintroduction to reach the minimum founding herd size. We hypothesize that the release area will serve as the core area of the herd range and serve as a base to explore the surrounding area (B. Jackson and Sharon Magee, Tall Grass Bison, Iowa, pers. comm.). Bison will be released into the handling facility and held with hay for 2 days and then allowed to move into a 4-5 acre area of electric fencing and held approximately 30 days so that they can be easily monitored upon release and habituate to their new home. This holding pen will also serve as a quarantine facility. Handling facility We will build a release and handling facility on APF deeded property within and convenient to the release pasture (Pasture #3, Fig. 1). The design will use a combination of pre-constructed commercial bison handling pens and large square straw bales stacked


2 high (10 ft) to construct a 50 ft X 50 ft pen. We will work with experienced bison handlers including Turner and TNC bison managers to design the pen and to conduct the initial handling and moving of bison. The pen will be partly enclosed by electric fencing similar to fencing in the release pasture so bison get exposed to this fence prior to release. Bison will be held in handling facility for 2 days after arrival and then allowed to move into the small fenced area (3-4 acres) where they will be held for approximately 3 weeks prior to moving them to large pasture. Disturbances will be kept to a minimum and hay will be provided. The first pasture is a relatively small area (490 ha or 1,200 acres; Fig 1) where they will be held for 1 year prior to removal of fences to allow them to occupy a large release area. Bison containment We will use fences that optimize containment of bison while at the same time do not restrict wildlife movements. Because no fence does both perfectly, and because fence and weather conditions will change, very infrequently bison will escape. Therefore our bison containment will focus on rapid response to any escapes to ensure that no harm is done to neighbors. We have worked with our neighbors and developed an MOU in this regard outlining that we will respond to escaped bison in a short period of time following escape and will resolve the problem to satisfaction quickly (Appendix ). Fencing Karhu and Anderson (2002) compared 3-wire- and 4-wire-high tensile electric fences to assess their ability to contain livestock but also allow passage of wildlife. Karhu and Anderson (2002) found that 3-wire (42 inches high, 22 inches from ground) electric was just as effective as 4-wire (52 inches high, 22 inches from ground) for containing bison. They reported 100% containment for the 3-wire fence. The higher top wire on a 4-wire fence is not needed to contain bison if they are trained to the electric fence and properly managed. They concluded bison can cross just about any fence but that the key to containment relies more on keeping the animals content by providing adequate space, forage, and water. The 3-wire fences were also the best design for allowing crossings by elk. Elk were more averse (spent more time searching for a place to cross) to crossing the 4-wire fence, but eventual number of crossings (outcomes) by elk, deer, and antelope did not differ between the 3 and 4 wire fences. They recommended using 1-inch fiberglass posts a minimum of fifty feet apart without stays to ensure the high tensile wire fence was flexible enough to allow easier crossing by wildlife. Follow-up work to Karhu and Anderson (2002) focused on additional testing of the 3 wire fence completed by Quitmeyer et al. (2004) provided additional data supporting the containment success of the fence for bison and also indicated low maintenance cost and liability risks for this type of fence. Because we will start bison reintroduction using young animals that will be trained on electric fence, we will start fencing conservatively to ensure adequate wildlife passage while minimizing risk of bison escape. To take advantage of the relatively new barbed wire fence that has been constructed in much of the allotment and to ensure that our neighbors have a fence that is readily repairable from their perspective, we will start by using a modified barbed wire fence for bison. We will fit 2 hot electric wires (solar


charged) to existing barbed wire fences. Wires will be attached to stays (every 3rd post) that extend 12 inches internally from the barbed wire fence (Fig ). The top electric wire will be 39 inches above the ground. The bottom wire will be 22 inches above ground. We will use this design for all exterior and internal fences where the barbed wire fence is in good condition. Where the barbed fencing is in bad condition, we will either replace and modify it or go to the 3 wire electric design depending on how this fits into remainder of fencing. We will place doublewide cattle guards at the 2 road crossings. We will monitor response of other wildlife to the fence, and if bison containment appears to be poor, we will work with our collaborators to develop other options including additional electric wires. Similar to Karhu and Anderson (2002), we will set up remote cameras along the fence to monitor wildlife crossings. We will also monitor crossings visually and based on tracks. We will sign the fence to alert hunters and create several hunter access points by placing steps over the fence. MANAGEMENT Disease We will minimally handle bison to reduce stress and to simulate to the extent possible semi free-ranging behavior and conditions. Over the next 5 years, the primary reason we will handle bison at all is to maintain herd health. Our schedule of testing (below) and vaccinations(see Aquisition Methods)approved by our team of oversight and managing veterinarians (Dave Hunter, TESF; Don Woerner, Montana Bison Association and private veterinarian; and Ann Johnson, Malta private veterinarian). The plan will be reviewed annually for the first 5 years. Because animals 1) will come from arguably brucellosis free herds and certainly brucellosis free states, 2) will meet any vaccination requirements of the state, and 3) will only be managed to control reportable disease, we will not conduct annual vaccinations after the initial release vaccinations. Such management has precedent and success in many herds near cattle including bison in the national park and national wildlife refuges including the National Bison Range in Montana (Boyd 2002). This approach has also been followed with no problems in the “free ranging” Henry Mountain, Utah herd that interacts regularly with cattle. It also has success in private herds including the Turner Flying D herd of approximately 2,000 bison. After 8 years of no vaccinations there is a zero prevalence rate of brucellosis (D. Hunter, Turner Enterprises, personal communication). We will conduct routine screening on a sample of animals annually for all diseases and on any animals if outward sign of disease becomes obvious. We used program EPISAMP (Colorado State University) to determine sampling sizes needed to detect presence of disease with 80% probability and prevalence in the population of 10%. With a population of 30 bison, approximately 13 would need to be sampled. Fourteen bison would need to sampled out of 50 and 15 bison out of 100 would need to be sampled. We will sample those animals by darting and immobilizing (see below) or moving them to the handling facility. Sick animals that appear to be untreatable or that have potentially contagious disease will be destroyed or removed from the county. We will do complete necropsies including disease screens on


all animals that are culled or that we find dead from natural mortality. Bison that move beyond fences will be captured and returned or killed. Escapes We will regularly check and maintain all fences to ensure proper containment. Bison will be monitored often via telemetry and observation to ensure containment. When animals infrequently escape, we will recapture by baiting and herding. To facilitate the recapture of bison, gates will be placed at logical places during fence construction. We have staff trained in wildlife immobilization to drug the animal. Baiting, herding or drugging may be options for recapture depending on the circumstances. If the escape occurs during the summer and the bison have been mingling with cattle, the bison will be recaptured, quarantined, and monitored for disease. A protocol and MOU are being developed with our neighbors for use when bison escape. Our drugging protocol will be to immobilize bison with carfentanil and xylazine using radio instrumented darts fired from the ground or air (Roffe et al. 1998). We will antagonize carfentanil and xylazine with naltrexone and yohimbine. We will kill by gunshot any animals that we cannot recapture. Mortality In general, bison will be naturally regulated with no management of reproduction or mortality. Carcasses of animals dying of natural factors will be allowed to decompose at the death site. While the goal is to establish a “self-regulating” bison population on lands it owns and leases, APF recognizes that it may be necessary to limit herd size for a number of reasons beyond natural mortality (from age, debility, or predation). Culling We anticipate that artificial culling may be needed in the following circumstances:

1) Herd size has exceeded carrying capacity and is significantly negatively effecting other resources or biodiversity targets;

2) Disease management

Our modeling and acquisition plans indicate carrying capacity will greatly exceed bison numbers in the short term and we expect exponential population growth. Over the longer term, without removals, we would expect the bison population to exhibit density dependence and to follow a logistic growth curve with declining birth rates and increasing death rates as density increases (Fredin 1984, Lartner et al 2000). Should removals be necessary to keep bison in the bounds of carrying capacity we will use the following guidelines that ensure retention of genetic diversity and that follow natural patterns for bison. Wilson and Zittlau (2004) assessed culling strategies in EINP for minimizing the potential negative effects of inbreeding. They found that removal of immature animals


would result in the slowest rate of diversity loss, as the genetic material they contain is most likely still represented by adults in the population. This will also act to increase generation length, and as diversity is only lost between generations, the loss of diversity will be further reduced. Wilson and Zittlau (2004) reported that although several factors did not exhibit a significant effect on the loss of genetic diversity, an optimal value or range could be identified for each tested variable. If culling is necessary, we will follow the recommendation of Wilson and Zittlau (2004) and remove a generally even number of male and female calves. Culling will involve, first, non-lethal removal, to the extent that receiving areas are available. In the event that no receiving areas are available, lethal removal will be implemented. If lethal removal is used, a portion of harvested animals allowed to remain in the field. Disposition of animals culled

1) Non-lethal removal: a. Establishment or augmentation of public or qualified nonprofit

conservation herds; b. To public or qualified nonprofit agencies for other than conservation

herds; c. To qualified tribal governments; d. Sale to local landowners for establishment of buffer herds.

2) Lethal removal:

a. Local public hunters; b. Harvest by APF staff.

Genetics and Population Management For optimal conservation strategy, Larter et al. (2000) concluded that bison reintroductions would be facilitated if animals were released into several independent sites since all herds would first increase in their new sites before natural range expansion took place. They indicated that recolonization could be facilitated and achieve a faster spread by several releases rather than waiting for a population to spread by its own density-dependent responses. We will assess rate of expansion of our initial release and relate that to overall carrying capacity and land acquisition to determine if starting additional founding herds in the area seems appropriate. We will also work with BSG and NPS to determine if mixing distant source herds is the proper management technique to ensure the best long-term conservation of the wild bison genome or to meet the genetic diversity objectives of WC (starting a satellite herd). While ecological and conservation consequences of combining animals of varied genetic backgrounds have been widely discussed, the demonstration of effects that stem from lineage mixing remains elusive; however, the opportunity for either inbreeding or outbreeding depression may be high (Berger and Cunningham 1995). Berger and Cunningham (1995) examined 4 potential correlates of fitness in the new sympatric


population: (1) female fecundity; (2) juvenile survival; (3) growth rates; and (4) female age at puberty. Of these, neither female fecundity nor juvenile survival was associated with lineage but growth rates were more rapid and ages at puberty were lower for F1 purebred (inbred) juveniles than for F1 hybrid (outbred) juveniles. Possible consequences of this variation in the F1 generation include (1) higher winter mortality in the slower growing line as well as (2) decreased lifetime production of young; both are life-history parameters that could be interpreted as long-term selection against outbreeding. Berger and Cunningham concluded that these interpretations (1) substantiate a level of variation in life-history parameters stemming from lineage mixing; and (2) suggest that advice regarding prudent conservation strategies must be sought concerning the genetic histories of individuals and populations to be selected for reintroduction. The prevalence of outbreeding and inbreeding tolerances within populations of managed and protected species needs verifiable documentation. We will follow their recommendations and determine these data for source herds and develop appropriate recommendations based on that and will subsequently monitor these factors over the period of restoration. If we determine that additional founding herds will enhance bison conservation and fitness of this herd, we will use a mix of source animals including EINP, WCNP, CR, and Henry Mountains. Bison Movement and Pasture Management Fences will be altered or improved initially according to the grazing management plan developed with the BLM. Initially, we will manage bison grazing and movements using the existing pasture system. Because bison numbers will be maintained well below carrying capacity we do not anticipate negative impacts from bison grazing to overall management of habitat and biodiversity. Should we identify significant short-term impacts of bison grazing that are at odds with overall management of biodiversity we will address these possibly through pasture management or culling. We do not intend to supplementally feed bison to address these concerns. However, if it appears that feed is needed in the first release pasture we may provide it during winter. While we expect that bison response to inadequate feed will be to attempt escape, we should become aware of forage limitations based on bison behavior. However, we will also consider a sampling protocol in the initial release years to look at crude fecal protein as a measure of bison health. Salt and/or trace mineral supplements may have to be provided in some circumstances since pasture boundaries will usually exclude access to these highly localized resources. Year-round water sources are available in each pasture. Pasture management and grazing options that we may consider in the short term (first 3 years) include seasonally grazed summer and winter pastures or fire-induced rotation (Bragg et al. 2001). Following Bragg et al. (2001), an approach to growing season rest-rotation of grazing without cross-fencing is to introduce fire as a means of creating zones of high forage quality that attract intense bison use while largely resting the remainder of the landscape. Selecting the location of burn patches to coincide with patches of high fuel accumulations will produce a landscape mosaic with a dynamic spatial and temporal pattern of seral plant communities. Another low-intensity management strategy is


season-long grazing alternated between fenced summer and winter management units. This strategy calls for bison to be grazed in one unit for the duration of the growing season, followed by grazing during the dormant season in a second unit, which has been deferred from grazing during the growing season (equivalent to consuming low quality "standing hay"). Our long-term objective is to create a bison population that is naturally regulated and free ranging and therefore allowed to move as bison biology, ecology and environmental factors dictate. This will require a larger but yet unknown amount of area in this region. This is unknown because we have no good information on how bison moved in this area historically, nor do we have good information on what drove these movements. In modern times, bison in some areas, including Yellowstone National Park, Wichita Mountains Wildlife Refuge, Oklahoma and Wood Buffalo National Park, do make seasonal movements. Roe (1970) concluded that movements of 19th century bison were unpredictable and should be classed more as nomadic rather than migratory. Forage must have been predictably available at the end of migration routes and this was likely not the situation on the Great Plains. Human-caused fires and hunting pressure would have made this unlikely. Bison inhabiting under-utilized grasslands will probably show little tendency to move great distances except during periods of extreme snow and cold. The best contemporary information on factors influencing bison movements comes from YNP. The number of bison leaving YNP in winter has been hypothesized to be driven by snow depth, snow crusting, forage (quantity, quality, and/or accessibility), bison population size, human trail grooming, herd tradition, or some combination of these factors (Meagher 1998, National Research Council 1998, Farnes et al. 1999, Bjornlie and Garrott 2001). A simple examination of relations between population size, winter severity, bison emigration, and human control operations associated with bison movements outside YNP provided support for all of the hypotheses advanced. A multiple regression model incorporating 3 independent variables (year, maximum bison count, and the average of 3 winter severity indices) and all linear interactions among these variables explained >90% of the variability in numbers of bison removed per year (F7,29 = 43.9, p < 0.001, R2 = 0.93). Unfortunately, not only were all the independent variables significantly related to bison removal, but also were all interactions among variables. This suggests that bison emigration is controlled by a complex relationship between weather (the more severe the winter, the greater the number of bison removed by human actions) and population size (the higher the population count, the greater the number of bison that are removed from the population) that has probably changed over time, but it does not elucidate the exact structure of the relationship. Bjornlie and Garrott (2001) noted that increases in population size would encourage more bison to seek areas with easier foraging conditions outside YNP in winters with deep snow or heavy snow crusting, but because winter conditions cannot be predicted, the number of bison actually exiting YNP cannot be predicted based solely on population size. Predators may also have affected both short-term and seasonal movements of bison (Carbyn et al. 1993).


All the factors potentially influencing bison movement in the restoration area will be assessed during our restoration effort (see below). We will use this information to design our land leasing and acquisition efforts and bison release and pasturing plans. Grassland management We will collect baseline information on many environmental variables related to bison impacts (see below), including range condition. We will produce a set of criteria for what we anticipate will be desired future conditions for range-related management of biodiversity. We will develop an adaptive management approach whereby our understanding of bison ecology in the region drives what these criteria are and feeds into rangeland and bison management. We are beginning to assess riparian conditions and needs for restoration. Part of that effort may include the need for exclosures to keep bison out of some riparian areas to aid short-term restoration and assess bison impacts. We will work with CMR to assess the fire history of the region and develop a fire management plan. The lack of trees suitable for fire-scar dating often makes an estimate of pre-settlement fire frequency difficult. Wright and Bailey (1980) suggested a 5- to 10-year fire return-interval for mixed-prairie grasslands. Ponderosa pine (Pinus ponderosa) fire scars in north-central Nebraska indicate an average fire-return interval of about 7.5 years for the pre-1850 period (Bragg et al. 2001). A review of explorer accounts along the upper Missouri River during the 1801-1805 period (Higgins 1986) suggests that at least some areas may have burned annually. In the short term, we will follow the paradigm and principles described by Truett et al. (2001) and Fuhlendorf and Engle (2001) and strive in the long-term for a naturally functioning system with little need for human intervention. Other wildlife management Because bison and prairie dog restoration appear inextricably linked (Truett et al. 2001), we will work to coordinate management and restoration of these 2 species as much as possible, using grazing by each to benefit the other (Truett and Phillips unpublished report). Fencing will be designed to accommodate ungulate movement, and fences will be monitored to assess any impacts to ungulate passage. Research on other aspects of wildlife response will be ongoing, dependent on funding. Work already funded includes construction of several riparian exclosures to be built in 2005, construction of at least 2 upland exclosures, and ongoing monitoring of bird and mammal populations locally. Organizational Risk Management Controls Risk Management APF is an IRS 501(c)(3)-qualified Montana not for profit organization (Appendix 2, IRS determination letter). APF’s primary mission is to acquire and manage lands to benefit biodiversity conservation. To this end, an endowment has been established to provide an income stream to cover core management costs, with the remainder of restoration costs


raised through grants and donations. While the duration of any project of this type is never assured, APF is managing risk of project failure in the following ways: 1) Ensuring success of APF’s mission APF has defined critical key components of its program, which include:

• A commitment to raise sufficient funds to assure the successful execution of the initial (Phase I) land acquisition program;

• Partnering with World Wildlife Fund which provides the technical expertise for

restoration science and planning, as well as financial support for land acquisition and management;.

• Finding and winning-over reliable funding sources to maintain sufficient operation funds to run the business and implement additional phases of the project;

• Establishing the reputation as “buyer-of-choice” with sellers and potential sellers

in the project area; • Securing acceptance and support of this project in the minds of the majority of

project-area residents and the general public; • Establishing and maintaining productive and positive relationships with key

government agencies associated with the project. 2) Maintaining the character of lands in perpetuity APF’s Board has agreed with 3rd party donors to place easements on all APF acquisitions within 3 years or in response to the following triggers:

1) prior to dissolution or corporate termination; 2) loss of charitable status; 3) change in mission set forth in APF’s articles and bylaws; 4) violation of operating standards for land management; 5) transfer or conveyance of Property.

The easement terms would limit activities that would significantly change the character of the property (such as sodbusting). Dissolution of the project In the event that APF is unable to fulfill its mission and is required to transfer or assign the land it owns and dispose of its assets, the following priority list will apply to bison remaining under APF control:


1) APF will provide the donor the opportunity to recover any stock donated or any progeny of stock that can be directly traced to parentage of the donor stock;

2) APF will offer the stock to a federal agency for establishment of a conservation herd on Park, Refuge, or other federal land;

3) APF will donate stock to any public or qualified nonprofit entity wishing to establish or augment a conservation herd;

4) APF will donate stock to any public or qualified nonprofit entity for purposes other than establishing a conservation herd;

5) APF will sell stock to any qualified tribal organization; 6) APF will sell stock and donate the receipts to a qualified nonprofit organization.

MONITORING AND RESEARCH PROGRAM (EXPERIMENTAL DESIGN) Justification The IUCN guidelines for reintroduction programs (IUCN 1998) indicate that post-release monitoring is considered vital and mandatory for any reintroduction. Accordingly, we will monitor this restoration project to maximize knowledge gained concerning wild bison restoration throughout North America. Notwithstanding our limited ability to design into our study plan rigorous replications and control of variables, this reintroduction will be implemented with a premium on gaining knowledge. For example, we will strive to provide clarity concerning the mechanisms of population growth and ecological impacts of large free-ranging populations of bison (Sarrazin and Barboult 1996). We will examine factors related to bison population growth including survival and reproduction (i.e. Kunkel and Pletscher 1999). We will monitor bison use of the landscape to assess potential impacts to biodiversity. We will work with collaborators at Utah State University (F. Provenza, Utah State University, personal communication) to test hypotheses related to social groups and bison movements. We will also compare bison demography, movement and ecosystem impacts to cattle in nearby areas (Van Vuren 2001). We will also conduct genetic monitoring of the herd to assess how quickly the genetic diversity present in the recently founded population is being lost through the process of genetic drift, differential male reproductive success, and offtake. We have also followed IUCN recommendations and established a bison management review team for this project (Appendix 7) and will work to enlist collaborators to develop detailed study designs for components of the monitoring project, as well as coordinating with other ongoing reintroductions in the region. We anticipate possibly enlisting review and collaboration from the Konza LTER researchers or TNC and university researchers to focus on rangeland impacts (Knapp et. al 1999, Fuhlendorf and Engle 2001). Because this restoration effort will necessarily proceed in a series of steps from small to large scale, we will design our monitoring program to assess impacts of bison on the system at each of these scales and compare impacts among the scales (initial, short term, mid term, long term; Turner et al. 1995). As we have indicated, the most significant effects are not likely to occur until bison function over a very large scale. Because this is


a long-term project and bison are a long-lived species and many impacts will likely not become evident for some time, all of our objectives will be carried out for >10 years. Bison research needs This project also provides the opportunity to address many basic research needs on bison. We know relatively little about bison ecology in large landscapes. There has been no direct assessment to date of bison cause-specific mortality rates. Predation rates have only been estimated by monitoring wolf kill rates over short periods of time (Carbyn et al. 1993). Relatively little work has examined ecosystem influences of cattle versus bison and this and most work on bison have been conducted at relatively small scales (Knapp et al. 1999, Fuhlendorf and Engle 2001, Van Vuren 2001). Research will be designed to address the following: Objectives: 1. Estimate bison reproductive rates annually; 2. Estimate bison survival rates annually; 3. Determine bison movement patterns, home range size, and resource selection annually; 4. Determine impacts of bison on range species productivity and diversity; 5. Determine impacts of bison on rangeland heterogeneity; 6. Determine impacts of bison on small mammal, bird, and large mammal density, especially prairie dogs and mountain plovers; 7. Determine impacts of bison carcasses on scavenger abundance. 8. Monitoring breeding, reproductive rates, and offtake rates and compare to genetic composition of herd over time. Hypotheses/predictions: 1. Survival rates of bison will be higher over early phases and high relative to other wild herds with a greater density of large carnivores; 2. Reproductive rates of bison will be higher over the early phases and high relative to other wild herds with a greater density of large carnivores; 3. Senescence will be primary mortality factor for bison in early phases and predation will be the primary in later phases; 4. Bison numbers will increase exponentially in early phases and logistically in later phases with forage availability being the primary limiting factor; 5. Bison home range size (standardized to scale) will be smaller in early vs. late phases; 6. Bison range expansion will occur through a series of pulses; 7. Relative movement distances (standardized to scale) of bison will be lower in early vs. late phases and therefore differential breeding by dominant males will not significantly reduce genetic diversity in the early phase; 8. Bison will exhibit greater and more complex social structure in late vs. early phases; 9. Rangeland heterogeneity of biomass and cover will be greater in late vs. early phases and greater than in comparable areas grazed by cattle; 10. Relative to cattle and availability, bison will select areas of open grassland and greater topographic relief for grazing over wooded areas and riparian areas;


11. Bison movements will be greater and behavior will be more aggregative than cattle at local scale and will be characterized as seasonal at later phases; 12. Bison carcasses will significantly enhance local species richness and abundance; 13. The abundance and richness of annual forbs will be greater in late vs. early phases and greater than comparison areas with cattle.

Methods We will employ a variety of techniques to obtain information on habitat use and behavior. In addition to basic observational monitoring (calving reproduction rates, mortality observation), we propose to fit a sample of bison with GPS radio collars that will allow us to monitor bison movements. Global positioning collars will allow us to obtain very accurate (within ~30 meters) and intensive (> 12 locations/day) bison locations. GPS locations will be uploaded to a GIS (ARC/INFO and ArcView, Environmental Systems Research Institute, Redlands, CA). Spatial and habitat attributes will be generated for bison locations from GIS map layers (30 m minimum mapping unit). Attributes will include land cover type, patch size, percent canopy (shrub, tree and grass), slope, and distance to permanent water. GIS will be used to classify topographic ruggedness and range site (soil type). We will create home ranges for bison using the Animal Movement extension in ArcView and examine ecological correlates of home range size and location. Landscape variables at bison telemetry relocations, control sites and random sites in the study area will be compared to assess resource selection (Kunkel and Pletscher 2000, Boyce et al. 2003). When a mortality is detected, we will investigate the site to locate the carcass and determine cause of death (Kunkel and Pletscher 1999). Survival rates will be determined using MICROMORT (Heisey and Fuller 1985). Key ecological variables including height, biomass, quality, and diversity of grasses and forbs, productivity, and nutrient recycling at bison grazing patches and lawns will be measured and compared over time (Knapp et al. 1999, Fuhlendorf and Engle 2001). Permanent plots and exclosures to monitor long-term trends of range condition will be established. We will follow existing (BLM, USFWS 1985) habitat evaluation procedures and range monitoring protocols to monitor range condition and trends for wildlife indicator species (USFWS 1985; Appendix 2). We will collect data on breeding bird and small mammal diversity in areas with and without bison (Ralph et al. 1993; Slater 1994). We will especially focus on potential impacts to species of management concern ( e.g., mountain plovers and sage grouse) and will collaborate with ongoing work on these species in the region. We will monitor abundance of potential bison predators including coyote and cougars. We will use scat deposition rates to monitor coyotes (Kost 1997) and track transects to monitor cougars (Beier et al. 1996). We will monitor number and time scavengers spend at bison carcasses (Wilmers et al. 2003). We will monitor interactions between bison and other ungulates including pronghorn and elk. We will monitor and identify bison group behavior and associations following protocols established by Provenza (Utah State University).


Dissemination of Results We will produce reports on a regular basis addressing project goals and objectives. These will include a report outlining protocols for bison conservation though reintroductions and peer reviewed scientific publications addressing our monitoring and research objectives. CRITERIA FOR MEASURING SUCCESS Initial success (1-2 years) will be reached when we achieve breeding and production of bison calves in the release area (Kleinman et al. 1991). Short-term criteria (3-5 years) for success will include a population that is 1) self-sustaining, with demographic rates similar to other wild self-sustaining populations (Lartner et al. 2000; see above); 2) partially naturally regulated, with no external inputs into the system by humans; and 3) ecologically effective at a small scale—structuring the vegetation, increasing associated species diversity and abundance, and providing biomass for predators, scavengers, and decomposers. Increasing success will mean expanding these parameters over a larger and larger area over time. We will consider mid-term success (5-10 years) when demographic rates approach self-sustaining levels yielding extinction probabilities over 100 years of <0.20, thus securing a new herd conserving the genetic diversity of wild bison. Long-term success (>10 years) will be achieved when bison are naturally regulated and ecologically effective over a large scale and evolving in a pattern consistent with their long term evolutionary history or that of other wild ungulates. Long-term success will be reached when bison populations expand and connect with other populations of genetically pure conservation herds in the region, are considered free ranging and wild, and contribute to removal of wild bison from the Montana list of species of concern. We may adjust these criteria as we learn more about bison ecology in the restoration area. We will assess factors determining reasons for not meeting any criteria for success. We will use adaptive management to modify management strategies to alleviate problems where needed.

Table 3. Project Milestone Chart (similar for each of 2004, 2005, 2006) Project

Months Calendar

Month(s) Milestone/Task

1-3 April - July Locate source herd and composition of animals to be reintroduced and finalize reintroduction plans

1 –5 April – September

Conduct pre-release surveys of key biodiversity components and prepare release site

6-7 October-November

Translocate bison

7-11 October – April Monitor bison movements, behavior, and survival 10 January Finalize BLM/state change in use EA and expand pastures 2-9 May-December Complete additional land acquisitions and agreements to allow


additional reintroductions or removal of some WR fences 13-19 April - October Monitor response of key biodiversity components to bison

presence 14-21 May –

December Monitor bison movements, behavior, reproduction and survival

22-34 January – December

Finalize plans with BLM allow bison to freely range on some allotments


Complete additional land acquisitions and agreements to allow additional reintroductions or removal of more WR and other private lands fences


Monitor bison movements, behavior, reproduction and survival

26-34 May – December

Analyze data on restoration success and write reports


ACKNOWLEDGMENTS We thank the Earth Friends Wildlife Foundation, J.M. Kaplan Fund, Mary A. and John M. McCarthy Foundation, and WWF’s Species Action Fund for generous financial support of this work. We thank Keith Aune, Pat Fargey, Dave Hunter, Greg Wilson, James Derr, and Peter Gogan, … for reviewing and offering constructive suggestions on a draft of this plan. We also deeply appreciate the assistance, information and advice provided by the following people at various stages in the development of this plan: Cortland Barnes, Scott Laird, Sean Gerrity, ….


Literature Cited Agnew, W., D.W. Uresk, and R.M. Hansen. 1986. Flora and fauna associated with prairie dog colonies and adjacent ungrazed mixed-grass prairie in western South Dakota. Journal of Range Management 39:135-39. Beier P, Cunningham SC. 1996. Power of track surveys to detect changes in cougar populations. Wildlife Society Bulletin 24:540-6.

Berger, J. 1992. Facilitation of reproductive synchrony by gestation adjustment in gregarious mammals: a new hypothesis. Ecology 73:323-329.

Berger, J., and C. Cunningham. 1994. Bison: mating and conservation in small populations. Columbia University Press, New York, NY. 330 pp.

Berger, J. and C. Cunningham. 1995. Multiple bottlenecks, allopatric lineages, and badlands bison Bos bison: consequences of lineage mixing. Biological Conservation 71:13-23. Bjornlie. D. D. and R.A. Garrott. 2001. The effects of groomed roads in the behavior and distribution of Bison bison in Yellowstone National Park. Journal of Wildlife Management. 65:560-572. Bogan, M. A. 1997. Historical changes in the landscape and vertebrate diversity of north central Nebraska. Pp. 105-130 in F. L. Knopf and F. B. Sampson, eds., Ecology and Conservation of Great Plains Vertebrates. Springer, New York. Boyd, D. 2003a Conservation of North American Bison: Status and Recommendations. Unpubl. Thesis, Univ. Calgary, Alberta. 222 pp. Boyd, D. 2003b. A Conservation Status Survey for Plains Bison in Canada. Year End Report. Prepared for Banff National Park. Facility of Environmental Design, University of Calgary. Boyce, M. S., J. S. Mao, E. H. Merrill, D. Fortin, M. G. Turner, J. Fryxell, and P. Turchin. 2003. Scale and heterogeneity in habitat selection by elk in Yellowstone National Park. Ecoscience 10:421-431. Bragg, T. K., B Hamilton and A. Steuter. 2001. Guidelines for bison management. The Nature Conservancy, Arlington Viginia. Brookshier, J.S. and W.S. Fairbanks. 2003. The nature and consequences of mother-daughter associations in naturally and forcibly weaned bison. Can. J. Zool. 81:414-423. Carbyn, L. N, S. M. Oosenbrug, and D. W. Anions. 1993. Wolves, bison, and the dynamics related to the Peace--Athabasca Delta in Canada's Wood Buffalo National Park. Canadian Circumpolar Institute Research Series Report 4.


Carlson, J. 2003. Montana Animal Species of Concern. Montana Natural Heritage Program and Montana Fish, Wildlife and parks, Helena, MT., 14pp. http://nhp.nris.state.mt.us/animal/reports/ASOCL_2003.pdf Clark, B. K., D. W. Kaufman, E. J. Finck, and G. A. Kaufman. 1989. Small mammals in tallgrass prairie: Patterns associated with grazing and burning. Prairie Naturalist 21:177-84. Coder, G. D., 1975 The national movement to preserve the American buffalo in the United States and Canada between 1880 and 1920. Ph.D. Dissertation (History). The Ohio State University, Columbus. Cody, M. L. 1985. Habitat selection in grassland and open-country birds. In Habitat Selection in Birds, ed. M. L. Cody, 191-226. San Diego, CA: Academic Press. Coppock, D. L., J. E. Ellis, J. K. Detling, and M. I. Dyer. 1983. Plant-herbivore interactions in a North American mixed-grass prairie. II. Responses of bison to modification of vegetation by prairie dogs. Oecologia 56:10-15. COSEWIC 2004. COSEWIC assessment and status report on the plains bison Bison bison bison in Canada. Committee on the Status of Endangered Wildlife in Canada. Ottawa, ON. vi + 71 pp. (www.sararegistry.gc.ca/status/status_e.cfm Coughenour, M. B. 1991. Spatial components of plant-herbivore interactions in pastoral, ranching, and native ungulate ecosystems. Journal of Range Management 44:530-42. Desmond, M. J., and J. A. Savidge. 1996. Factors influencing burrowing owl (Speotyto cunicularia) nest densities and numbers in western Nebraska. American Midland Naturalist 136:143-48. Detling, J. K. 1998. Mammalian herbivores: ecosystem-level effects in two grassland national parks. Wildlife Society Bulletin 26:438-48. Estes, J.A., D. O. Duggins, and G. B. Rathbun. 1989. The ecology of extinctions in kelp forest communities. Conservation Biology 3: 252-264. Farnes, P., C. Heydon, and K. Hansen. 1999. Snowpack distribution across Yellowstone National Park. Final Report, Cooperative Research Agreement Number CA 1268-1-9017. Yellowstone Center for Resources, Yellowstone National Park, Wyoming. Festa-Bianchet, M. 1991. The social system of bighorn sheep: grouping patterns, kinship and female dominance rank. Anim. Behav. 42:71-82.


Flores, D. 1991. Bison ecology and bison diplomacy: The southern plains from 1800 to 1850. The Journal of American History 78(2):465-85. Forrest, S.C., and D. Biggins, 1988. Black-Footed Ferret Recovery Plan (Revision) U.S. Fish and Wildlife Service. Forrest, S.C., H. Strand, W.H. Haskins, C. Freese, J. Proctor and E. Dinerstein. 2004. Ocean of Grass: A conservation Assessment for the Northern Great Plains. Northern Plains Conservation Network and Northern Great Plains Ecoregion, WWF-US, Bozeman, MT. http://www.worldwildlifefund.org/wildplaces/negp/pubs/ocean_of_grass.cfm

Franklin, I.R. and R. Frankham. 1998. How large must populations be to retain evolutionary potential? Animal Conservation 1: 69-73. Fredin, R.A. 1984. Levels of maximum net productivity of large terrestrial mammals. Pages 381-387 in W. F. Perrin, R. L. Brownell, Jr,. and D. P. DeMaster, editors. Report of the International Whaling Commission Special Issue 6. Cambridge U.K.

Freilich, J. E., J. M. Emlen, J. J. Duda, C. Freeman, and P. J. Cafro. 2003. Ecological effects of ranching: a six-point critique. Bioscience 53:759-765.

Fuhlendorf, S. D., and D. M. Engle. 2001. Restoring heterogeneity on rangelands: ecosystem management based on evolutionary grazing patterns. BioScience 51: 625-632. Fuller, W.A. 1966. The biology and management of the bison of Wood Buffalo National Park. Canadian Wildlife Service Wildlife Management Bulletin Series 1:1-52. Gates, C.C., and N.C. Larter. 1990. Growth and dispersal of an erupting large herbivore population in northern Canada: The MacKenzie wood bison (Bison bison athabascae). Arctic 43:231-238. Green, W.C.H., and A. Rothstein 1991. Trade-offs between growth and reproduction in female bison. Oecologia 86:521-527. Gross,J.E.2000.AdynamicsimulationmodelforevaluatingtheeffectsofremovalandcontraceptionongeneticvariationanddemographyofPryorMountainwildhorses.BiologicalConservation96:319-330. Gross, J. E. and G. Wang. 2005. Effects of Population Control Strategies on Retention of Genetic Diversity in National Park Service Bison (Bison bison) Herds. National Park Service Final Report, Bozeman, Montana. Haig, S. M., J. O. Ballou, and S. R. Derrickson. 1990. Management options for preserving genetic diversity: reintroduction of Guam rails into the wild. Conservation Biology 4:290-300.


Halbert, N.D. 2003. The utilization of genetic markers to resolve modern management issues in historic bison populations: implications for species conservation. Ph.D. Dissertation, Texas A&M Univ., College Station, TX, Halloran, A.F. 1968. Bison (Bovidae) productivity on the Wichita Mountains Wildlife Refuge, Oklahoma. Southwestern Naturalist 13:23-26. Hamlin K. L. and R. J. Mackie 1989. Mule deer in the Missouri River Breaks, Montana: a study of population dynamics in a fluctuating environment. Final Report, Montana Fish Wildlife and Parks. Hart, R.H. 2001. Where the buffalo roamed — or did they? Great Plains Research 11:83-102. Hartnett, D.C., A.A. Steuter and K.R. Hickman. 1997. Comparative Ecology of Native and Introduced Ungulates. Ecology and Conservation of Great Plains Vertebrates. Springer-Verlag New York, Inc.

Haugen, A.O. 1974. Reproduction in the plains bison. Iowa State Journal of Research 49:1-8.

Heisey, D. M. and T. K. Fuller. 1985. Evaluation of survival and cause-specific mortality rates using telemetry data. Journal of Wildlife Management 49:668-674. Hill, K. D. 1993. The endangered species act: what do we mean by species? Environ. Affairs 20: 239–264.

Higgins, K.F. 1986. Interpretation and compendium of historical fire accounts in the northern great plains. USDI Fish and Wildl. Serv. Res. Pub. 161. Holecheck, J. L. 1988. An approach for setting the stocking rate. Rangelands 10:10-14. Holekamp, K.E. and L. Smale. 1991. Dominance acquisition during mammalian social development: the ‘inheritance’ of maternal rank. Am. Zool. 31:306-317. Hornaday, W.T. 1889. The extermination of the American bison, with a sketch of its discovery and life history: Report of the National Museum, Washington, DC: U.S. Government Printing Office. Howery, L.D., F.D. Provenza, R.E. Banner and C.B. Scott. 1998. Social and environmental factors influence cattle distribution on rangeland. Appl. Anim. Behav. Sci., 55:231-244. Isenberg, A.C. 2000. The destruction of the bison. Cambridge University Press. IUCN/SSC Re-introduction Specialist Group. 1998. IUCN guidelines for re-introductions. IUCN, Gland, Switzerland and Cambridge, UK. 10 pp.


James,J.W.1971.Thefoundereffectandresponsetoartificialselection.GeneticalResearch12:249-266. Januszewski MC, Olsen SC, Mclean RG, Clark L, and Rhyan JC. 2001. Biosafety of Brucella abortus strain RB51 in non-target species. Journal of Wildlife Diseases. J. Wildlife Dis. 37: 532-537. Johnson, C.N. 1986. philopatry, reproductive success of females, and maternal investment in the red-necked wallaby. Behav. Ecol. Sociobiol. 19:143-150. Jolly and Messier 2000. Karhu, R. and S. Anderson. 2002. Evaluation of high tensile electric fence design on big game movements and livestock containment. Wyoming Cooperative Fish and Wildlife Research Unit Final Report. Kirkpatrick, J.F., J.C. McCarthy, D.F. Gudermuth, S.E. Shideler, and B.L. Lasley. 1996. An assessment of the reproductive biology of Yellowstone bison (Bison bison) subpopulations using noncapture methods. Canadian Journal of Zoology 74:8-14. Kleinman, D. G. et al. 1991. Costs of reintroduction and criteria for success: accounting

accountability in the golden lion tamarin conservation program. Pages 125-142 in J. H. W. Gipps, editor. Beyond Captive Breeding : Reintroducing Endangered Species to the Wild. Oxford University Press, Oxford, United Kingdom.

Knapp, A.K., J.M. Blair, J.M. Briggs, S.L. Collins, D.C. Hartnett, L.C. Johnson, and E.G. Towne. 1999. The keystone role of bison in North American tallgrass prairie. BioScience 49:39-50. Knowles, C. J. 1986. Some relationships of black-tailed prairie dogs to livestock grazing. Great Basin Naturalist 46:198-203. Knowles, C. J., C.J. Stoner, and S. P. Gieb. 1982. Selective use of black-tailed prairie dog towns by mountain plovers. Condor 84:71-74. Knowles, C.J., J. Proctor, and S.C. Forrest. 2002. Black-tailed prairie dog abundance and distribution in the Great Plains based on historic and contemporary information. Great Plains Research 12:219-54. Koford, C. B. 1959. Prairie dogs, whitefaces, and blue grama. Wildlife Monographs 3:1-78. Kojola, I. 1989. Mother’s dominance status and differential investment in reindeer calves. Anim. Behav. 38:177-185.


Kost, C. D. 1997. Estimation of predator densities in western South Dakota. M.S. Thesis, South Dakota State University, Brookings, South Dakota, USA. Kotliar, N.B., B.W. Baker, A.D. Whicker, and G. Plumb. 1999. A critical review of assumptions about the prairie dog as a keystone species. Environmental Management 24:177-192. Krausman, P., K.E Kunkel, M. Phillips. 2001. Private restoration of bighorn sheep. Page 231-242 in D. Maehr, R. Noss, and J. Larkin, editors. Restoration of Large Mammals. Island Press, California, USA. Kunkel, K.E., K. Honness, M.K. Phillips, and L. N. Carbyn. 2003. Plan for restoring swift foxes to west-central South Dakota. Pages 189-198 in Ecology and Management of Swift Fox in a Changing World. L.Carbyn and M. Sovada, editors. Canadian Plains Research Center, Regina, Saskatchewan, Canada. Kunkel, K. E. and D. H. Pletscher. 1999. Species-specific population dynamics of cervids in a multipredator ecosystem. Journal of Wildlife Management 63:1082-1093. Kunkel, K.E. and D.H. Pletscher. 2000. Habitat factors affecting vulnerability of moose to predation by wolves in southeastern British Columbia. Canadian Journal of Zoology 78:150-157. Laliberte, A. 2003. Human Influences on Historical and Current Wildlife Distributions from Lewis and Clark to Today. Ph.D. Dissertation. Oregon State University. Lacy, R.C. 1994. Managing Genetic Diversity in Captive Population of Animals. Pages 63-89 in M.L.Bowles and C.J. Whelan (Editors). Restoration of Endangered Species: Conceptual Issues, Planning, and Implementation. Cambridge University Press, New York, NY. Lacy, R.C. 1997. Importance of Genetic Variation on the Viability of Mammalian Populations. Journal of Mammalogy 78(2):320-335. Larter, N. C., A. R. E. Sinclair, and C. C. Gates 1994. The response of predators to an erupting bison, Bison bison athabascae population. Canadien Field-Nauralist 108:318–327. Larter, N. C., A. R. E. Sinclair, T. Ellsworth, J. Nishi and C. C. Gates. 2000. Dynamics of reintroduction in an indigenous large ungulate: the wood bison of northern Canada. Animal Conservation 3:299-309 Leberg, P. L. 1990. Genetic considerations in the design of introduction programs. Transactions of the North American Wildlife and Natural Resources Conference 55:609-619.


Licht, D. S., and K. D. Sanchez. 1993. Association of black-tailed prairie dog colonies with cattle point attractants in the northern Great Plains. Great Basin Naturalist 53:385-89. Lott, D.F. 1979. Dominance relations and breeding rate in mature male American bison. Zietschrifte fur Tierpsychologie 49: 418-432.

Lott, D.F. 2002. American Bison: A Natural History. University of California Press, London, England. 229 pp.

Lott, D.F., and J.C. Galland. 1987. Body mass as a factor influencing dominance status in American bison cows. Journal of Mammalogy 68:683-685.

Ludlow, W. 1876. Report on a Reconnaissance from Carroll, Montana Terr. on the Upper Missouri to the Yellowstone National Park and Return, Made in the Summer of 1875. War Dept., Washington, D.C. (MSU library, Bozeman, MT, Renne Coll.). Lynch, M. and R. Lande. 1998. The critical effective size for a genetically secure population. Animal Conservation 1:70-72. Lyman, R. L. and S. Wolverton. 2002. The late prehistoric-early historic game sink in the northwestern United States. Conservation Biology 16:73-85. Mahan, B.R. 1978. Aspects of American bison (Bison bison) social behavior at Fort Niobrara National Wildlife Refuge, Valentine, Nebraska, with special reference to calves. M.S. Thesis, University of Nebraska, Lincoln, Nebraska. Maher, C.R., and J.A. Byers. 1987. Age-related changes in reproductive effort of male bison. Behavioral Ecology and Sociobiology 21:91-96. Martin, P. S. and C. R. Szuter. 1999. War zones and game sinks in the Lewis and Clark west. Conservation Biology 13:36-45. Maximillian, Prince of Weid (Alexander Phillip von). 1843. In Early Western Travels 1748-1846, Maximillian, Prince of Wied’s travels in the interior of North America, 1832-34, Vol. 2, Thwaites, R.G. ed. AMS Press, Inc., New York. McComb, K., C. Moss, S.M. Durant, L. Baker and S. Sayialel. 2001. Matriarchs as repositories of social knowledge in African elephants. Science 292:491-494. McHugh, T. 1972. The Time of the Buffalo. New York, NY: A. A. Knopf.

McPherson, G. R. 1995. The role of fire in the desert grasslands. Pp. 130-151 in M. P. McClaren and T. R. VanDevender, eds., The Desert Grassland. Tucson: University of Arizona Press.


Mead, J. R. 1899. Some natural history notes of 1859. Transactions of the Kansas Academy of Science 16:280-81. Meagher, M.M. 1973. The bison of Yellowstone National Park. Scientific Monograph Series #1. U.S. Government Printing Office, Washington, D.C. 161 pp. Meagher, M. 1998. Recent changes in Yellowstone bison numbers and distribution. Pp. 107-112 in Irby, L. and J. Knight, eds. International Symposium on Bison Ecology and Management in North America. Montana State University, Bozeman, Montana. Messiter, C.A. 1890. Sport and adventure among the North American Indians. R.H. Porter, London. Miller, B., G. Ceballos, and R.P. Reading. 1994. The prairie dog and biotic diversity. Conservation Biology 8:677-681 Miller, P. S. and R. C. Lacy. 1999. VORTEX: A stochastic simulation of the extinction process. Version 8 User’s manual. Apple Valley, MN: Conservation Breeding Specialist Group (SSC/IUCN). Mills, L.C., M.E. Soulé, and D.F. Doak. 1993. The Keystone-species Concept in Ecology and Conservation. BioScience 43:219-24. Morrison, M.L. 2002. Wildlife Restoration: Techniques for Habitat Analysis and Animal Monitoring. Society for Ecological Restoration. Island Press, Washington DC. 209 pages. National Research Council. 2002. Ecological Dynamics on Yellowstone's Northern Range. National Academy Press, Washington, D.C. Nelson, T., J. Holecheck, and R. Valdez. 1999. Wildlife plant community preference in the Chihuahuan Desert. Rangelands 21:9-11. Norland, J.E., L.R. Irby, and C.B. Marlow. 1985. Determination of optimum bison stocking rate in Theodore Roosevelt National Park, North Dakota. Journal of Environmental Management 21:225-239. Novaro, A. J., M. C. Funes, and S. Walker. 2000. Ecological extinction of native prey of a carnivore assemblage in Argentine Patagonia. Biological Conservation 92:25-33. O’Brien, S. J. & Mayr, E. 1991. Bureaucratic mischief: recognizing endangered species and subspecies. Science 251:1187–1188. Olsen SC, Jensen AE, Palmer MV, Stevens MG. 1998. Evaluation of serologic responses, lymphocyte proliferative responses, and clearance from lymphatic organs after vaccination of bison with Brucella abortus strain RB51 Am. J. Vet. Res. 59:410-415.


Olsen SC, Kreeger TJ, Schultz W. 2002 Immune responses of bison to ballistic or hand vaccination with Brucella abortus strain RB51. J. Wildlife Dis.38: 738-745. Olsen SC, Jensen AE, Stoffregen WC, and Palmer MV. 2003. Efficacy of calfhood vaccination with Brucella abortus strain RB51 in protecting bison against brucellosis. Res. Vet. Sci 74: 17-22. Plumb, G.E., and J.L. Dodd. 1993. Foraging ecology of bison and cattle on a mixed prairie: Implications for natural area management. Ecological Applications 3:631-43. Pyare, S. and J. Berger. 2003. Beyond demography and delisting: ecological recovery for Yellowstones’s grizzlies and wolves. Biological Conservation 113-63-72. Quitmeyer, C. J., J. A. Bopp, R. M. Stephens, R. Karhu, and S. Anderson. 2004. High tensile electric fence: phase 2 – liability issues, maintenance costs, and containment of bison. Wyoming Cooperative Wildlife Research Unit Final Report. Ralph, J. C., G. R. Geupel, P. Pyle, T. E. Martin, and D. F. DeSante. 1993. Handbook of field methods for monitoring landbirds. Reynolds, H.W., C. C. Gates, and R.D. Glaholt. 2003. Bison. Wild Animals of North America. Biology. Management. Economics. The John Hopkins University Press, Baltimore. Roe, F.G. 1951. The North American Buffalo: A critical study of the species in its wild state. Univ. of Toronto Press. Roffe T. J., D. Hunter, and K. Aune. 1998. Field experience with etorphine and carfentinal immobilization of free-rangin bison in Yellowstone National Park. Page 194 in L.R. Irby and J.E. Knight (Editors). International Symposium on Bison Ecology and Management in North America, Montana State University, Bozeman, MT. Roffe TJ, Olsen SC, Gidlewski T, Jensen AE, Palmer MV, Huber R. 1999. Biosafety of parenteral Brucella abortus strain RB51 vaccine in bison calves. J. Wildlife Management. 63:950-955. Roth, S. D., Jr., and J. M. Marzluff. 1989. Nest placement and productivity of ferruginous hawks in western Kansas. Transactions of the Kansas Academy of Science 92(3-4):13248.

Roth, D., and R. Peterson. 1997. A neotropical migratory bird prioritization for national forests and grasslands. In Conserving Biodiversity on Native Rangelands: Symposium Proceedings, eds. D. W. Uresk, G. L. Schenbeck, and J. T. O’Rourke, 3-7. Rapid City, SD: US Forest Service Rocky Mountain Forest and Range Experiment Station.


Rhymer, J. M. & Simberloff, D. (1996). Extinction by hybridization and introgression. Ann. Rev. Ecol. Syst. 27:83–109. Rutberg, A.T. 1986. Notes and Comments: Lactation and fetal sex ratios in American bison. American Naturalist 12:89-94. Sarrazin, F. and R. Barbault. 1996. Reintroduction: challenges and lessons for basic ecology. Trends in Ecology and Evolution 11:474-478. Scott, C.B., F.D. Provenza, and R.E. Banner. 1995. Dietary habits and social interactions affect choice of feeding location by sheep. Applied Animal Behavior Science. 45:225 -237. Sherman, P.W. 1977. Nepotism and the evolution of alarm calls. Science 197:1246-1253. Slater, P. J. 1994. Factors affecting the efficiency of the area search method of censusing birds in open forests and woodlands. Emu 94:9-16. Snell, G. P., and B. D. Hlavachick. 1980. Control of prairie dogs--the easy way. Rangelands 2:239-40. Smale, L., L.G. Frank and K.E. Holekamp. 1993. Ontogeny of dominance in free-living spotted hyaenas: juvenile relations with adult females and immigrant males. Anim. Behav. 46:467-477. Soule, M. E., J. A. Estes, J. Berger, and C. Matinez Del Rio. 2003. Ecological effectiveness: conservation goals for interactive species. Conservation Biology 17:1238-1250. Steuter, A. A. 1997. Bison. Pages 339-347 in The tallgrass restoration handbook. S. Packard and C. F. Mutel editors. Island Press, Washington D. C. Stoddart, L. A., and A. D. Smith. 1955. Range Management. New York, NY: McGraw-Hill. Stubbendieck, J., J. A. Lamphere, and J. B. Fitzgerald. 1997. The blowout penstemon, an endangered species. Lincoln: Nebraska Game and Parks Commission. The Nature Conservancy. Northern Great Plains Ecoregional Planning Team. 2000. Ecoregional Planning in the Northern Great Plains Steppe. The Nature Conservancy. 181 pp. http://www.conserveonline.org/scd;internal&action=buildframes.action Towne, G.E. 2000. Prairie Vegetation and Soil Nutrient Responses to Ungulate Carcasses. Oecologia 122:232-239. Truett, J.C. 2003. Migrations of grasslands communities and grazing philosophies in the Great Plains: a review and implications for management. Great Plains Research 13: 3-26.


Truett, J.C., and T. Savage. 1998. Reintroducing prairie dogs into desert grasslands. Restoration and Management Notes 16:189-95. Truett, J.C., M. Phillips, K. Kunkel and R. Miller. 2001. Managing Bison to Restore Biodiversity. Great Plains Research 11: 123-44. Tulloch, D.G. 1978. The water buffalo, Bubalus bubalis, in Australia: grouping and home range. Aust. Wildl. Res. 5:327-354. Turner, M, G., R. H. Gardner and R. V. O'Neill. 1995. Ecological Dynamics at Broad Scales: Ecosystems and Landscapes. BioScience Suppl. p. 529-535.

U.S. Bureau of Land Management. 1992. Judith Valley Phillips Resource Management Plan. U.S. Dept. of Interior, Bureau of Land Management, Montana State Office, Billings.

U.S. Bureau of Land Management. 1997. Roe Allotment Change in Class of Livestock from Cattle to Bison, Environmental Assessment No. MT-076-97-04, Dillon Resource Area, Dillon, MT. U.S. Bureau of Land Management. 2001. Sullivan Electric Fence Environmental Assessment, WY-030-EA1-180, Rawlins Field Office, Rawlins, WY. http://www.wy.blm.gov/nepa/docs/SullivanEleFenceEA.pdf U.S. Department of Agriculture. 2003. National Range and Pasture Handbook. Natural Resources Conservation Service, Grazing Lands Technology Institute. ftp://ftp-fc.sc.egov.usda.gov/GLTI/technical/publications/nrph/nrph-cover.pdf U.S. Fish and Wildlife Service. 1985. Final environmental impact assessment for the Charles M. Russell National Wildlife Refuge. U.S. Fish and Wildlife Service, Denver Colorado. U.S. Fish and Wildlife Service. 1999. Endangered and threatened wildlife and plants: Proposed threatened status for the mountain plover. Federal Register 64(30):7587-7601. Vallentine, J. F. 1989. Range Development and Improvements. San Diego, CA: Academic Press. van Gelder, R.G. 1977. Mammalian hybrids and generic limits. Am. Mus. Novit. 2635:1-25. Van Vuren, D.H. 1982. Comparative ecology of bison and cattle in the Henry Mountains, Utah (Summer diets, preferred forages, distributions, ranges). In: Proceedings of the Wildlife-Livestock Relationships Symposium: held at Coeur d'Alene, Idaho, April 20-22, 1981 / sponsored by Department of Wildlife Resources, University of Idaho; James M.


Peek, P.D. Dalke, editors. v, 614 p. Moscow, Idaho: Forest, Wildlife & Range Experiment Station, University of Idaho, 1982. p. 449-457. Van Vuren, D. H. 2001. Spatial relations of American bison (Bison bison) and domestic cattle in a montane environment. Animal Biodiversity and Conservation 24(1): 117-124. 2001 Van Vuren, D. and M.P. Bray 1986. Population dynamics of bison in the Henry Mountains, Utah. Journal of Mammalogy 67(3): 503-511. 1986. Vinton, M.A., and S.L. Collins. 1997. Landscape gradients and habitat structure in native grasslands of the central Great Plains. In Ecology and Conservation of Great Plains Vertebrates, eds. F. L. Knopf and F. B. Sampson, 3-19. New York, NY: Springer. Ward, T. J. 2000. An evaluation of the outcome of interspecific hybridization events coincident with a dramatic demographic decline in North American bison. PhD Dissertation, Texas A and M University, College Station. Ward, T. J., J. P. Bielawski, S K. Davis, J. W. Templeton and J. N. Derr. 1999. Identification of domestic cattle hybrids in wild cattle and bison species: a general approach using mtDNA markers and the parametric bootstrap. Animal Conservation 2:51–57. Wells, P. V. 1965. Scarp woodlands, transported grassland soils, and concept of grassland climate in the Great Plains region. Science 148:216-49. Wilmers, C. C., R. L. Crabtree, D. W. Smith, K. M. Murphy, and W. M. Getz. 2003. Trophic facilitation by introduced top predators: Grey wolf subsidies to scavengers in Yellowstone National Park. Journal of Animal Ecology 72:909-916. Wilson, G.A., and C. Strobeck. 1999. Genetic variation within and relatedness among wood and plains bison populations. Genome 42:483-496. Wilson, G. A.; W. Olson, and C. Strobeck. 2002. Reproductive success in wood bison (Bison bison athabascae) established using molecular techniques. Canadian Journal of Zoology 80:1537-1548. Wilson G. A. and K. Zittlau. 2004. Management Strategies for Minimizing the Loss of Genetic Diversity in Wood and Plains Bison Populations at Elk Island National Park. Parks Canada Agency Species at Risk Support National Office, 25 Eddy Street Gatineau, Quebec Canada K1A 0M5 Email: [email protected]. 58 pp.

Wobeser, G. 2002. Disease management strategies for wildlife. Review of the Science and Technology Office of International Epizoology 21:159-178.

Wright, H.A. and A.W. Bailey. 1980. Fire ecology and prescribed burning in the Great Plains - a research review. USDA For. Serv., Gen. Tech. Rep. INT-77.


Appendix 1. Input Values for VORTEX Simulations.

Variable Number of Iterations 100 Number of Years 100 Extinction Definition Only 1 sex remains Species Description Inbreeding Depression Lethal Equivalents 3.14, 5.00 % due to recessive lethals 50 EV concordance of reproduction and

survival Yes

# of types of catastrophes 0 Reproductive System Polygamous Age of first offspring for females 3 Age of first offspring for males 6 Max age of reproduction 20 Max number of progeny per year 1 Sex ratio at birth (% males) 50

Density dependent reproduction? No Reproductive Rates % Adult females breeding 75 EV in % breeding 7.5 Distribution of number of offspring per

female per year 100% probability

of 1 offspring

Mortality Mortality of Females as % Mortality from Age 0 to 1 15.0 SD in 0 to 1 Mortality due to EV 4.0 Mortality from Age 1 to 2 5.0 SD in 1 to 2 Mortality due to EV 6.0 Mortality from Age 2 to 3 5.0 SD in 2 to 3 Mortality due to EV 3.0 Annual Mortality After Age 3 5.0 SD in Mortality After Age 3 3.0 Mortality of Males as % Mortality from Age 0 to 1 20.0 SD in 0 to 1 Mortality due to EV 18.0 Mortality from Age 1 to 2 15.0 SD in 1 to 2 Mortality due to EV 5.0 Mortality from Age 2 to 3 5.0 SD in 2 to 3 Mortality due to EV 2.0 Mortality from Age 3 to 4 5.0 SD in 3 to 4 Mortality due to EV 2.0 Mortality from Age 4 to 5 5.0 SD in 4 to 5 Mortality due to EV 2.0


Variable Mortality from Age 5 to 6 5.0 SD in 5 to 6 Mortality due to EV 2.0 Annual Mortality After Age 6 5.0 SD in Mortality After Age 6 2.0 % Males successfully siring offspring 40 Initial population size Female Ages Females Age 1 0 Females Age 2 2 Females Age 3 2 Females Age 4 0 Females Age 5 0 Females Age 6 0 Females Age 7 0 Females Age 8 0 Females Age 9 0 Females Age 10 0 Females Age 11 0 Females Age 12 0 Females Age 13 0 Females Age 14 0 Females Age 15 0 Females Age 16 0 Females Age 17 0 Females Age 18 0 Females Age 19 0 Females Age 20 0 Male Ages Males Age 1 0 Males Age 2 2 Males Age 3 2 Males Age 4 0 Males Age 5 0 Males Age 6 0 Males Age 7 0 Males Age 8 0 Males Age 9 0 Males Age 10 0 Males Age 11 0 Males Age 12 0 Males Age 13 0 Males Age 14 0 Males Age 15 0 Males Age 16 0 Males Age 17 0 Males Age 18 0 Males Age 19 0 Males Age 20 0


Variable Carrying capacity K 5,000 SD in K due to EV 0 Trend in K N Supplementation Pop supplemented? Yes First year of supplement 2 Last year of supplement 3 Interval between supplement 1 # Females supplemented Age 2 supplement 7

Age 3 supplement 7 # Males supplemented Age 2 supplement 7

Age 3 supplement 7


Appendices 2, 3, 4 --- not attached Appendix 5 – not attached; in development Appendix 6. –not attached


Appedix 7. Bison Review Team Table. 1 Bison Project Collaborators Review Team Keith Aune - MFWP Cormack Gates - University of Calgary, BSG Mike Hedrick – CMR Dennis Linghor – BLM Al Steuter – TNC Mark Kossler – TEI Greg Wilson – University of California, Berkely Bob Jackson J. Derr – Texas A & M Tom Roffe – USGS Don Woerner – Montana Bison Association Veterinarian Ann Johnson – Malta Veterinarian Monitoring and Research 1. Forrest/Freese, Knapp et al.?, Fuhlendorf?, and Kunkel will lead work on rangeland impacts. 2. Forrest and Matchett will lead the bison and prairie dog interaction work. 3. Kunkel, Gogan, Matchett, Aune, and Carbyn will lead bison demographics, movement, resource selection, and large mammal interaction work. 4. Freese, Dinerstein, and Matchett will lead bison and bird interaction work. 5. Kunkel and Provenza will lead bison social interaction work 6. Derr and Wilson will lead conservation genetics work. 7. Scott Laird will lead Containment and Neighbor/Community Relations & Education

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