volume 19 • issue 1 • 2011
Yellowstone Bison
Carrying Capacity and Movement Balancing Brucellosis Risk and Wildlife Conservation Historical Wolf Population Numbers ©Dan S t a h ler
Juvenile male bison spar during the rutting season in Hayden Valley. They struggle for dominance and the opportunity to secure their genetic legacy.
A Little Learning Can Be A Provocative Thing
lthough Yellowstone Science is used as a means of Ever since the idea of reintroducing wolves in the conveying the depth and breadth of research that takes Yellowstone area caught on decades ago, people have wanted place in the park, acknowledging what we haven’t fig- to know how many wolves resided here before their wide- Aured out yet is also important. In or between its lines, Yellow spread slaughter began in the nineteenth century. Lee stone Science offers many questions still in search of answers. Whittlesey and Paul Schullery show how letters written by In this issue, we present new information on an old a member of the 1872 Hayden Survey add to the evidence topic—why do bison leave the park? Well, yes, because the on this enigmatic topic, but they also point out the limita- grass is greener there, at least in early spring, but there’s a tions of such evidence. Can we be reasonably certain that the lot more to it than that. Authors P.J. White, John Treanor, number of skins Joseph Savage reported were those of wolves Michael Coughenour, and Rick Wallen raise and attempt to and not coyotes? answer questions such as why bison have left even in years By raising such questions, we hope to provoke readers’ when less than half of the forage in the park had been eaten. curiosity and inspire further efforts to fill the information Seasonal timing is also a critical factor in another manage- gaps. ment challenge addressed in this issue, brucellosis transmis- We hope you enjoy the issue. sion by elk and bison. a periodical devoted to natural and cultural resources NPS
volume 19 • issue 1 • March 2011 Each spring, Yellowstone staff work with Montana Department of Livestock personnel to conduct bison hazing operations along the park’s west boundary. Mary Ann Franke Guest Editor Tami Blackford E ditor F e at u r e s Janine Waller Graphic Designer Associate Editor 8 Carrying Capacity and Movements of Emily Yost Jen Baseden Yellowstone Bison A ssistant Editors Biologists use modelling to study how many bison Yellowstone Artcraft Printers, Inc. can support and explore the reasons for bison migration and Bozeman, Montana Printer dispersal outside the park. P.J. White, Glenn E. Plumb, Michael B. Coughenour, and Rick L. Wallen 15 Balancing Brucellosis Risk Management and Wildlife Conservation Land managers and park scientists examine the complex Yellowstone Science is published periodically. Support for Yellowstone Science is provided scientific and social issues surrounding the how, when, and by the Yellowstone Association, a nonprofit where of brucellosis transmission. educational organization dedicated to serving the park and its visitors. For more information P.J. White, John J. Treanor, and Rick L. Wallen about the association, including membership, or to donate to the production of Yellowstone Science, visit www.yellowstoneassociation.org or 23 Wolves in the Historical Record write: Yellowstone Association, PO Box 117, How many wolves lived in Yellowstone before it became a Yellowstone National Park, WY 82190. The opinions expressed in Yellowstone Science are park? A newly discovered account from 1872 sheds light on the authors’ and may not reflect either National Park Service policy or the views of nineteenth century wolf abundance. the Yellowstone Center for Resources. Lee Whittlesey and Paul Schullery Copyright © 2011, Yellowstone Association for Natural Science, History & Education. For back issues of Yellowstone Science, please see www.greateryellowstonescience.org/ys. D e pa rt m e n t s
Submissions are welcome from all investigators conducting formal research in the Yellowstone area. To submit proposals for articles, to 2 News & Notes subscribe, or to send a letter to the editor, UW-NPS Research Center Agreement • Tested Bison in please write to the following address: Editor, Yellowstone Science, PO Box 168, Gardiner Basin • DOI Bison Conservation Initiative • Yellowstone National Park, WY 82190. You may also email: [email protected]. P.J. White Receives Director’s Award • Winter Elk Count • Cougar Skulls Yellowstone Science is printed on recycled paper with a soy-based ink. 6 Shorts Insights about Surface and Groundwater Flow • Nutrient
Cover photo: Cycling: Transitioning from an Elk- to Bison-dominated Bison and elk grazing on Grassland System • Older Hot Spring Alteration in the Grand Yellowstone’s northern range. Canyon of the Yellowstone River • Migration and Dispersal: NPS photo by Jim Peaco. Key Processes for Conserving National Parks News & Notes N
NPS and UW Renew PS/Pea
Research Center Agreement co 2011 An agreement signed at the AMK Ranch in Grand Teton National Park will continue the longest-standing and one of the most successful part- nerships between a university and the National Park Service. On July 21, 2010, University of Wyoming President Tom Buchanan and Grand Teton Superintendent Mary Gibson Scott signed a 10-year agreement for continued use of the historic Grand Teton Ranch, the site of the University of Wyoming-National Park Service A group of 25 brucellosis seronegative bison being released from the Stephens (UW-NPS) Research Center. Scientists Creek holding facility within Yellowstone National Park. from around the world conduct biolog- ical and physical science research and Tested Bison Allowed in cultural and social studies at the facility Society, and the Wyoming Game and Gardiner Basin in support of resources throughout the Fish Department. In 1953, the UW Greater Yellowstone Ecosystem. joined in operating and sponsoring On January 19, 2011, a group of 25 The ceremony featured remarks the station and its research program at bison bulls and cows that tested from Superintendent Scott, President the Jackson Hole Biological Research negative for exposure to brucellosis Buchanan, University of Wyoming Station near the old Moran town site. (seronegative) were released from the Vice President for Research and The UW-NPS Research Center was Yellowstone National Park Stephens Economic Development Bill Gern, established in 1977 when the headquar- Creek capture facility to a 2,500‑acre and Zoology Professor and UW-NPS ters were moved to the AMK Ranch. grazing area in the adjacent Gallatin Research Center Director Hank At any given time during the summer, National Forest by Interagency Bison Harlow. The UW-NPS Research the AMK Ranch serves as a base for Management Plan (IBMP) members. Center traces its roots to 1946, when 50 to 60 scientists. The AMK Ranch The bison were trailed by riders on the first research station was launched is also used for many seminars, field horseback as they moved north through with the help of the Jackson Hole trips, workshops, and symposia. “We’re the park and the Royal Teton Ranch Preserve, Inc., the New York Zoological honored to be a part of this relation- along the Old Yellowstone Trail and ship—one that onto the national forest, where, based
N has produced on the IBMP, they may remain until PS significant sci- April 15, 2011. Bison are permitted to entific research access the national forest via the pri- and crucial vately-owned Royal Teton Ranch under information a 30-year agreement with the Church that helps Universal and Triumphant signed in inform our 2008. Each of the bison is marked and current and fitted with monitoring devices. future manage- Thirteen of the 25 bison repeatedly ment of park left the authorized grazing area. They resources,” said were captured by the Montana Depart- Scott during ment of Livestock and released back The Berol Lodge at AMK Ranch, location of the University of the signing into the park. One bull from the test Wyoming-National Park Service Research Center. ceremony. group was shot after it entered private
2 Yellowstone Science 19(1) • 2011 N PS/Pea areas. There are also 4,000 woods
co bison free-roaming in Canada. The 2011 Department of the Interior (DOI) manages almost 7,000 bison in national wildlife refuges and national parks. The initiative proposes several actions to address the health and genetic composition of the bison popu- lations on the DOI lands. To promote cooperative conservation in bison management, the DOI will strengthen Interagency staff trail the small group of bison across the Royal Teton Ranch and existing partnerships and build new into the Gallatin National Forest. ones with state agencies, American Indian tribes, landowners, agricultural interests, conservationists and others property and behaved aggressively. The symbols of the American frontier is the interested in bison health and recovery. cooperating agencies will monitor the image of vast herds of bison grazing on Where there is strong local support, movement of the remaining 11 bison to the western plains…Americans today partnerships could permit small bison determine how they use this landscape still find inspiration in bison ranging herds to recreate their natural role in and may agree to allow as many as freely on the landscape, as Yellowstone areas where they are not currently pres- 100 untested bison on the authorized National Park demonstrates.” Kemp- ent. Such arrangements may help sup- grazing area in the future. Other bison thorne further remarked that, while port the restoration or maintenance of management within Yellowstone and the days of millions of free-roaming other native species and habitats, and at the park boundary will continue as bison are gone, the Department policy become important tourist attractions. outlined in the IBMP and park man- must acknowledge the important role An interagency working group agement policies. of bison on the landscape, in tribal cul- will coordinate management and sci- The IBMP, established in 2000, is ture, and in our national heritage. ence needs and activities related to part of an interagency effort to con- There are more than 500,000 plains the DOI’s bison herds and carry out serve a viable, wild bison population bison in North America today, but cooperative efforts with other parties. while preventing the transmission of most are privately owned, have a large The group includes representatives brucellosis from bison to cattle. The extent of cattle genetic introgression, from the Fish and Wildlife Service, the agencies cooperating under the IBMP and are kept in herds of less than 1,000 National Park Service, the Bureau of are the National Park Service, the US that are fenced within relatively small Land Management, the US Geological Forest Service, the USDA Animal and
Plant Health Inspection Service, the N PS/Pea Montana Department of Livestock, the co
Montana Department of Fish, Wildlife 2011 and Parks, the InterTribal Buffalo Council, the Confederated Salish Kootenai Tribes, and the Nez Perce Tribe.
Department of the Interior Bison Conservation Initiative
On October 28, 2008, Secretary of the Interior Dirk Kempthorne announced an initiative for the Department to work with state, tribal, and agricultural interests to strengthen bison conserva- tion efforts. In his announcement, Bison at the Stephens Creek facility are tested to determine the degree to which Kempthorne stated “One of the classic the population is infected (called seroprevalence) with the Brucella abortus bacteria.
19(1) • 2011 Yellowstone Science 3 Survey, and a tribal liaison from the they represent the best of NPS efforts • Preparation of convincing legal Bureau of Indian Affairs, as well as to safeguard some of the most precious responses to a federal lawsuit representatives of states in which bison qualities and natural resources of our against the US Forest Service and are located. The Animal and Plant park landscapes.” Department of the Interior alleging Health Inspection Service of the US As noted in the nomination submit- impairment of Yellowstone bison and Department of Agriculture, which ted by then Yellowstone Superintendent other complaints due to management plays a major role in bison disease Suzanne Lewis, P.J. White has been a actions; and issues, and the US Forest Service and champion of producing objective sci- • Completion of an assessment of Department of Defense, which are ence to support decisions regarding ecological process management in major public land managing agencies, the conservation of natural resources Yellowstone, with recommendations have been invited to participate as well. and the processes that sustain them in for improved approaches and prac- Additional information on the Bison Yellowstone National Park. He excels at tices to mitigate future uncertainties Conservation Initiative can be accessed building relationships between groups such as climate warming, invasive at www.nature.nps.gov/biology. of scientists to conduct mission-critical species, and land use changes. research, making science accessible to Each of these accomplishments White Receives 2010 managers and visitors by creating reflects the culmination of years of Intermountain Region Natural understandable products, and formu- effort and required significant long- Resource Stewardship Award lating and negotiating effective plans to term vision, perseverance, patience, and
N conserve and restore species and pro- diligence. PS cesses across jurisdictions. P.J.’s com- The other 2010 recipients are: mitment to the science-informed man- • Superintendent of the Year for Natural agement of park resources contributed Resource Stewardship—Alexa Roberts, to four significant accomplishments in superintendent, Bent’s Old Fort 2010: National Historic Site • Completion of an adaptive manage- • Natural Resource Research—William ment plan that significantly increased G. Parker, physical scientist, Petrified tolerance for Yellowstone bison Forest National Park migrating to essential winter ranges • Natural Resource Management in a in Montana and reduced the capture Small Park—Chris Ford, chief of Dr. P.J. White. and culling of bison due to concerns Integrated Resources, Grant-Kohrs about disease transmission to cattle; Ranch National Historic Site P.J. White, Branch Chief of Aquatic • Completion of an environmental • Natural Resource Professional— and Wildlife Resources at Yellowstone impact statement analyzing the Gregory Mark Anderson, aquatic National Park, is one of six employees impacts of remote vaccination of bison ecologist, Glen Canyon National in the Intermountain Region (IMR) (i.e., without capture) for brucellosis; Recreation Area to have received Natural Resource
Stewardship Awards for 2010. The N PS/Pea awards, in categories ranging from co
management to maintenance to 1999 research, recognize outstanding con- tributions to the care, protection, and appreciation of the landscapes, wildlife, and other natural attributes of National Park Service sites. White received the Director’s Award for Natural Resource Management. “In their everyday duties, these six colleagues give us daily lessons in how to accomplish better a key part of our National Park Service mission: the protection and preservation of park resources,” IMR Director John Wessels said. “With diligence and creativity, An elk herd searches for winter forage in Lamar Valley.
4 Yellowstone Science 19(1) • 2011 Y ELL
• Natural Resource Stewardship through Biologists expect the reduction in -134140/
Maintenance—Jackie Messer, facil- the number of wolves and the elimina- co ity manager, Chickasaw National tion of the late season hunt will result lle ct e Recreation Area in some increase in the elk population. d 8/18/1994 As regional winners, the six employ- The Northern Yellowstone ees are eligible for the national 2010 Cooperative Wildlife Working Group NPS Director’s Awards for Natural will continue to monitor trends of the Resources, which will be presented dur- northern Yellowstone elk population ing the biennial George Wright Society and evaluate the relative contribution Skull of a male cougar that died from Conference on Parks, Protected Areas, of various components of mortality. starvation at 6.5 years of age. and Cultural Sites, March 14–18, 2011 in New Orleans. Cougar Skulls Acquired by Heritage and Research Center Cougars were essentially elimi- Winter Elk Count nated from their native habitat in the The museum at Yellowstone National Yellowstone region in the early 1900s The annual aerial survey of the north- Park’s Heritage and Research Center as part of a predator control program. ern Yellowstone elk herd conducted recently received cougar skulls col- The last known cougar was killed in in December 2010 counted 4,635 elk, lected during Phase One of the Cougar 1925. Occasional sightings of cougars 24% less than the 6,070 reported in Project in the park. Researchers studied that might have dispersed from other January 2010. predation and the reproductive success areas were reported in the following Park and university biologists say of cougars in northern Yellowstone years and increased after the 1950s. The increased predation, ongoing drought, and the surrounding area from 1987 project studied this population. hunting pressure, and other factors to 1996. Conclusions from the project Researchers collared 80 of the 88 have contributed to an apparent 70% helped inform the park’s management cougars captured during the project. decline in the herd population in the about cougar numbers and population They concluded the population’s prey last fifteen years. The elk count in dynamics, this carnivore’s food require- was primarily comprised of elk and 1995, when wolf restoration began in ments, and interactions with other mule deer, with elk calves being the Yellowstone National Park, was 16,791. predators. most important prey and adult elk Predation by wolves and grizzly being the least important relative to Y
bears is cited as a major reason for the ELL their availability. Researchers also con- decline. Both prey primarily on elk, -134142/C cluded that resident males (as opposed and predation on newborn elk calves to roaming males) sired nearly all cou- o may limit the population’s ability to lle gar litters. The number of breeding-age ct e recover. d females present within a male’s territory 12/18/1989 Drought during the early 2000s and the male’s age were correlated with appears to have impacted the quality the number of litters the male sired and abundance of forage, possibly low- each year. ering reproduction rates in some elk. Interviews with project lead Kerry The number of permits issued for Murphy appear in early issues of the antler-less Gardiner Late Elk Hunt Yellowstone Science, 2(3) and 2(4), and was reduced from 1,102 in 2005 to 100 can be accessed at www.greateryellow during the 2006–2010 seasons and has stonescience.org/ys. been eliminated for 2011. The Heritage and Research Center Biologists suspect predator num- is home to Yellowstone National Park’s bers may be responding somewhat museum collections and holds more to the decline in the elk population. than 13,000 natural specimens, includ- The number of grizzly bears seen on ing 10,000 herbarium specimens. To the northern range during elk calving access the collections, contact the season has decreased slightly in recent museum staff at (307)344-2662 for an years. Also, the wolf population on The skull of a six month-old male appointment or go to www.nps.gov/yell/ the northern range inside Yellowstone cougar (pictured at two angles) shows historyculture/collections.htm for more National Park has dropped from 94 the two large puncture wounds where information. wolves in 2007 to 39 wolves in 2010. it was killed by adult male cougar.
19(1) • 2011 Yellowstone Science 5 N PS/O bsidi an Cree
ShortsShortsShortsShorts k
th r ough
Surface and Groundwater Flow in Yellowstone fall than rivers outside the caldera. This is because the exten- W i ll
Gardner, W.P., D.D. Susong, D.K. Solomon, and H. Heasler. 2010. Snowmelt sive and young rhyolitic volcanic rocks in Yellowstone form ow hydrograph interpretation: Revealing watershed-scale hydrologic characteris- large, fractured groundwater aquifers. Much of the spring Fla tics of the Yellowstone volcanic plateau. Journal of Hydrology, 383:209–222. ts snowmelt infiltrates these aquifers and is discharged to riv- In this study, Gardner and others gained insights about surface ers throughout the year rather than primarily during spring and groundwater flow in the Yellowstone volcanic plateau by runoff. Geochemical dating of water discharged from cool examining annual hydrographs from US Geological Survey springs in the rhyolitic volcanic rocks indicates groundwa- stream gauges and dating groundwater samples from cool ter ages of less than 50 years. This short residence time for water springs. Analyses of annual streamflow hydrographs the groundwater implies a rapid circulation of groundwater indicate that groundwater inflows—rather than direct runoff within the rhyolitic volcanic aquifers. The results presented from precipitation—are the dominant influence on stream- in this paper are part of a larger study investigating the shal- flow within the Yellowstone caldera and rhyolite volcanic low, groundwater-flow system in Yellowstone and its connec- plateaus. Rivers with headwaters in the Yellowstone caldera tion to the Yellowstone hydrothermal system. and associated rhyolitic plateaus have smaller spring runoff —David D. Susong, Supervisory Hydrologist, peaks, carry less sediment, and have larger base flows in the US Geological Survey
Nutrient Cycling: Transitioning from an Elk to of drought. Though early green-up provides high nutrition Bison Dominated Grassland System for ungulates during the latter stages of gestation and lacta- Based on data and fieldwork from Yellowstone National Park ungulate tion, contributing to increased calf survival, an early decrease monitoring projects. in forage could leave females with marginal fat reserves for Plant production in grassland ecosystems is generally limited pregnancy and survival the following winter. by moisture and nutrient availability, and may decrease or Relatively low elk body condition and pregnancy rates increase depending on how the plants are grazed. Although have been detected during recent years in the eastern por- intensive grazing by ungulates may reduce nutrient cycling tion of the northern grassland. The reduction in wintering and plant production, more moderate grazing can maxi- elk numbers by more than one-half may have contributed mize forage quality and quantity. The high densities of elk to the more than doubling of bison numbers from 1996 to (12–17/km2) on Yellowstone’s northern range during the 2008. Elk and bison have dietary overlap, but elk feed more 1980s and 1990s deposited large quantities of nitrogen, on isolated plants and most migrate to higher elevations after phosphorus, and other nutrients, stimulating plant produc- early spring. Bison, which comprise less than 5% of wolf tion. The reduction of elk numbers to 10–11/km2 as a result kills, tend to create distinct grazing patches and remain on of hunter harvest, predation by a growing wolf population, the northern range through the summer, resulting in differ- severe winterkill in 1997, and drought during 1999 and 2000 ent effects on nutrient cycling. Bison grazing and wallowing resulted in conditions less conducive to plant growth. Studies convert grass-dominated sites to sites of greater plant diver- of the northern range suggest that nutrient dynamics and the sity and spatial heterogeneity. Depending on future bison system as a whole are within long-term variations. Migratory density, the bison’s larger biomass, strong herding tendencies, ungulates concentrate foraging for shorter intervals and and selective consumption of dominant grasses and sedges transfer nutrients via feces and urine from other areas, while could contribute to extensive areas of grazing-tolerant plants predators keep ungulate densities below carrying capacity. that are repeatedly grazed throughout the season, but shift However, there is some indication that the northern across the landscape from year to year, as well as possible range grassland is in a state of flux. The decline in elk density long-term changes in rates of nitrogen cycling and nutrient to approximately 3–5/km2 during 2006 to 2010 reduced for- redistribution. Research on how recent changes in elk and age consumption and nutrient deposition. Also, warmer tem- bison distributions have affected ecosystem processes such as peratures in northern Yellowstone during the past 50 years nutrient cycling is planned to begin in summer 2011 with Dr. have brought an earlier peak in the growing season followed Douglas Frank of Syracuse University. by earlier curing of vegetation and more frequent periods —P. J. White and Rick L. Wallen, Yellowstone National Park
6 Yellowstone Science 19(1) • 2011 Mt. Washburn Older Hot Spring Alteration in the Grand Canyon of the Yellowstone River
R Larson, P.B., A. Phillips, D. John, M. Cosca, C. Pritchard, A. Andersen, J. Manion. EOCENE ABSAROKA VOLCANICS E IV
R 2009. A preliminary study of older hot spring alteration in Sevenmile Hole, E N Grand Canyon of the Yellowstone River, Yellowstone Caldera, Wyoming. O WST Journal of Volcanology and Geothermal Research 188: 225–236. LO L GIN YE MAR ER A ALD Erosion in the Grand Canyon of the Yellowstone River has C exposed a cross-section of more than 300 vertical meters of Canyon IM R rhyolites left by the most recent caldera collapse (640 ka) that Junction N YO have been altered by hydrothermal activity for about 150,000 CA N N years. Research on the canyon walls in the Sevenmile Hole Lower area found minerals (quartz, opal, kaolinite, dickite, alunite, Falls 2km illite, adularia, and pyrite) that formed two alteration zones Upper Falls PLEISTOCENE UPPER BASIN characteristic of shallow hydrothermal systems in volcanic MEMBER OF THE PLATEAU RHYOLITES environments. The advanced argillic zone, located in the first 100 meters below the canyon rim, has an intermediate argillic Sevenmile Hole area (Modified from Christiansen and Blank zone directly beneath it. This transition in clay mineralogy 1975; Prostka et al. 1975; and Christiansen 2001). is also found in active alkali-chloride hydrothermal springs elsewhere in Yellowstone, at temperatures of 150°C to 170°C. across the northern part of the caldera. There it is concealed The Sevenmile Hole area lies at the eastern end of a band of by the younger tuff of Sulphur Creek (480 ka). hydrothermal features that may be aligned with a ring fault —Peter Larson, Washington State University
Migration and Dispersal: Key Processes for Although more than half of the Greater Yellowstone Conserving National Parks ecosystem is on public land, livestock grazing is extensive on Based on data and fieldwork from Yellowstone National Park ungulate the national forests, and valley bottoms and flood plains with monitoring projects. higher plant productivity and more moderate winter condi- The seasonal round-trips undertaken by migrating wildlife tions are primarily on private land. Culling to prevent bison can increase their access to food and mates, and release areas migration and expansion of winter range outside the park from intensive use for part of the year. Dispersal, in which has perpetuated irruptive population dynamics and reduced one or more members of a population make a one-way move- female cohorts. Forcing bison to stay in the park would create ment to another area, can keep population density within artificially high densities that would have negative effects on the food-limited capacity of the environment. In Yellowstone vegetation, soils, and other ungulates. National Park, the presence of species such as the bald eagle Thus, initiatives are needed to maintain and restore eco- and trumpeter swan, for which the park has little suitable logical processes across management jurisdictions and prevent year-round habitat, depends on migration and dispersal from conflicts with species such as bison, bears, and wolves out- more productive areas. Elk, bison, deer, and pronghorn need side the park. Ideally, partnerships of government agencies, to migrate beyond the park’s boundaries to access some of American Indian tribes, non-governmental organizations, their traditional winter range. and private landowners would develop a consensus on objec- However, migration and dispersal have undergone a tives and management actions. However, management of world-wide demise as wildlife has been compressed into wildlife outside the park is the prerogative of the surrounding disconnected areas. The proportion of pronghorn migrat- states, where many officials and citizens remain unconvinced ing from their truncated winter range near the park’s north- that ecological integrity is in their best interests. Over time, ern boundary during summer increased in recent decades, however, ecosystem process management can enhance the possibly in response to extended droughts. Also, the first lives of all of Greater Yellowstone’s residents by sustaining its significant presence of pronghorn in the southern Paradise natural resources and making it possible to address other chal- Valley in many decades apparently occurred when a group lenges such as climate change, forest die-offs, and emerging of Yellowstone pronghorn dispersed approximately 30 kilo- diseases that can be transmitted between wildlife, livestock, meters north of the park in 2000 and grew to 82 animals and humans. by 2009. Immigration and gene flow between pronghorn in —P. J. White, Glenn E. Plumb, and Rick L. Wallen, Yellowstone and the Paradise Valley could improve the long- Yellowstone National Park term viability of both populations.
19(1) • 2011 Yellowstone Science 7 NPS abortus bacteria before 1917, likely from infected In 1969, managers in Yellowstone stopped removing bison bison removing stopped Yellowstone in managers 1969, In Brucella to 60% to up then, since and, 1994) Meyer and (Meagher cattle of the population has tested positive for anti-bodies indi- cating exposure to this nonnative pathogen (Cheville et al. non- of birth the or abortions induce may Brucellosis 1998). viable calves in bison, cattle, and elk (Rhyan et al. 2009). fluctu- to numbers allowed and size population their limit to limitations resource and predators, weather, to response in ate (Cole 1971). Bison abundance increased rapidly under this (fig. regime 2) management and, since the increasing 1980s, numbers have moved during winter outside the park where federal and tribal, state, by or hunted culled been have some agencies (Fuller et al. 2007a, b). Bison movements beyond boundary led to claims that they over- were the Yellowstone abundant and had degraded the range inside the park (e.g., Kay 1998). These claims, in turn, led to calls for intensive However, However, by the early twentieth 1).
19(1) • 2011
Postcard #157: “The Buffalo Herd”, by Frank Jay Haynes. Date unknown. Haynes began issuing Haynes Date unknown. Haynes. Frank Jay by “The Buffalo Herd”, #157: Postcard in 1921. out of his studio in 1897 and he retired postcards
he successful conservation of Yellowstone National the Park bison from a of low two of dozen about animals in 1902 to more than 5,000
Yellowstone Science Yellowstone T P. J. White, Glenn E. Plumb, Michael B. Coughenour, and Rick L. Wallen and Rick L. Coughenour, B. Michael E. Plumb, Glenn White, J. P. of Yellowstone Bison Yellowstone of Carrying Capacity and Movements Carrying Movements and Capacity animals in 2005 has led to enduring conflicts among vari- ous publics and management agencies regarding issues of perceived overabundance and the risk of brucellosis trans- mission to cattle (Plumb et al. occupied 2009). approximately Bison 20,000 historically square kilometers in headwaters the of the Yellowstone and Madison rivers in northern the portion of what is now referred to as the Greater Yellowstone area (fig. (YNP) sanctu- provided Park National Yellowstone century, ary to the only wild and free-ranging plains bison remain- ing in the United States (Plumb and Sucec 2006). These animals were spatially segregated into two herds ing occupy- the central plateau and northern portions of the park, exposed Also, these bison were 1973). (Meagher respectively 8 management to limit the abundance and distribution of bison We used the best available scientific information to inside YNP, including fencing, fertility control, hunting, and evaluate a central question in this debate, which is whether brucellosis test-and-slaughter programs (US Department of bison move outside the park because their abundance has the Interior and US Department of Agriculture 2000a). Since surpassed levels that can be supported by the forage base in 2000, the risk of brucellosis transmission from Yellowstone the park. We also considered other potential explanations bison to livestock has been managed by the federal gov- for bison movements outside the park during winter and ernment and the state of Montana pursuant to the court- spring, and appraised the implications of perceived over- negotiated Interagency Bison Management Plan, which abundance and brucellosis transmission risk for long-term allows for the culling of bison in park boundary areas if hazing bison conservation. (i.e., forcing bison to move back into designated conservation areas) becomes ineffective at maintaining separation between Have Bison Exceeded their Carrying Capacity? bison and cattle (US Department of the Interior and US Department of Agriculture 2000b). These actions are highly Ecological carrying capacity has been defined as the natural contentious and polarizing, and the repeated culling of large limit of a population set by resources in a particular environ- numbers of bison could potentially affect the age structure, ment (Caughley and Sinclair 1994). It is one of the equilib- reproduction, recruitment, or genetics of Yellowstone bison rium points that populations tend toward as animal den- in unintended ways over the long term (White et al. 2009). sity increases and the amount of food, space, cover, or other
r e v Montana Ri e on st w o ll e ± Idaho Y Wyoming
M a d is o n
R
iv
e r
Yellowstone Lake
Legend
Curent Distribution (IMBP 2007)
1940s Distribution (Meagher 1973)
Pre-Settlement Distribution (approximation based on archived reports and journals of expeditions through the area) 0 15 30 60 Park Boundary Kilometers Figure 1. Pre-settlement, mid-twentieth century, and current distribution of Yellowstone bison (Meagher 1973; Plumb and Sucec 2006; Schullery and Whittlesey 2006; Plumb et al. 2009).
19(1) • 2011 Yellowstone Science 9 NPS lum A Simulated Simulated bison population dynamics agreed well with observed data (Coughenour 2005). When the model was run for eight simulations for the northern and central herds simultaneously over 50 years, with random weather varia- tions and allowing no range expansion by bison outside the park, the northern herd increased to a mean of 2,417 bison (range=1,820–3,530, median=2,670) and the central herd increased to a mean of 3,776 bison (range=2,430–5,630, comparison, median=4,030).the actual For maximum total bison during count summer of Yellowstone 2005 was 1,484 bison in the northern herd and 3,531 bison in the central Thus, neither herd. the central nor the northern bison herd capac- carrying food-limited mean estimated its exceeded has varia- inter-annual extensive are there though park, the in ity tions in carrying capacity due to variations in for- weather, et al. and other competition, factors (Plumb age availability, 2009). Also, the model predicted that the bison population would be under nutritional stress well below food-limited carrying capacity during winters with deep snowpacks that restricted bison access to forage. As a result, there would be due mortality adult increased and mortality calf considerable to starvation (Coughenour 2005). Creek in Hayden Valley during summer aerial surveys. during summer aerial surveys. Valley in Hayden Creek Why Do Bison Move Outside the Park? Outside the Park? Do Bison Move Why migrations seasonal increased, numbers bison Yellowstone As along altitudinal gradients within YNP became the higher-elevation from norm, moving herds both in bison some with summer ranges to lower elevations from autumn through returning to summer ranges 1989; (Meagher winter, in June Bjornlie and Garrott 2001; Bruggeman et al. 2009). These seasonal, round-trip movements were initially detected in central YNP during the 1970s, when bison abundance was low (<500) and the summer range should have ample resources for bison year-round (Meagher 1993, 1998; provided Bruggeman 2006). Thus, Yellowstone bison were partially the approach to began abundance their before well migratory Yellowstone bison observed in the upper reaches of bison observed upper reaches in the Yellowstone
YELL-15805/ G. Baggley Vaccinium Year ugust, 1970–2010. ugust, A 19(1) • 2011 anch in December 1930. anch in December 1930.
R umber of Yellowstone bison observed bison during Yellowstone umber of Central Northern N veys of the central (square) and northerncentral (square) of the veys 1970 1980 1990 2000 2010 0
500
4000 3500 3000 2500 2000 1500 1000 No. of Bison of No. amar Buffalo Yellowstone Science Yellowstone L Three mounted rangers separate bison for slaughter at the slaughter at the mounted rangers separate bison for Three shrubs, and coniferous trees. The model used Geographic Information System (GIS) data for soils, vegetation, topog- and raphy, other variables, and was driven by weather data near and in located sites telemetry snow and climate 29 from generated were maps temperature and Precipitation park. the and accumulation the simulated model snow validated a and melting of snow. resources diminishes. Coughenour (2005) evaluated whether whether evaluated (2005) Coughenour diminishes. resources capac- carrying food-limited a reached had bison Yellowstone ity YNP inside by developing and testing a spatially-explicit ecosystem model that (SAVANNA) integrated data on site water balance, plant biomass production, plant population dynamics, litter decomposition and nitrogen cycling, ungu- energy ungulate distribution, spatial ungulate herbivory, late pred- and predation, dynamics, population ungulate balance, ator population The dynamics model (fig. simulated 3). the central and northern herds of bison, Yellowstone as well as Madison) and (northern herds elk wintering resident two the and summer immigrant elk. Nine plants functional were groups simulated, of including fine- graminoids, and forbs, sagebrush, deciduous coarse-leaved shrubs, Figure 2. 2. Figure aerial sur in July– (triangle) herds 10 11 NPS/Peaco ellowstone Science ellowstone Y
19(1) • 2011 Bison also move out of the park in an attempt to expand to attempt an in park the of out move also Bison Decreased foraging efficiency can induce dispersal areas areas at lower elevations and on thermally warmed ground Bruggeman (Meagher 1989; 2006; Bruggeman et al. 2009). Also, snow melts earlier at lower elevations outside the park (e.g., Gardiner basin, Horse Butte peninsula) resulting in earlier green-up and more energy-efficient foraging oppor- tunities than on the higher-elevation summer ranges in the park (Thein et al. 2009).Thus, the numbers and timing of bison migrating from their summer range to their winter range in or outside the park is positively related build-up on the summer higher-elevation range in the park. to snow The return migration from lower-elevation winter of wave the following ranges June, and mid-May in begins typically growing vegetation from lower to higher elevations, similar pronghorn) elk, deer, sheep, bighorn (e.g., ungulates other to 2010; 2007, al. et White 2005; al. et (Gates ecosystem this in Thein et al. 2009). their range. This dispersal, or movement from one spatial of continuum a essentially is return, without another to unit migration. Increasing bison density decreases the resources available for each animal, which can negatively their influence nutrition, body condition, reproduction, and survival (Sinclair 1975; Caughley 1976). Range expansion can delay these responses to food limitation since new ranges provide additional forage and limitations will become apparent pri- (Messier colonized be longer no can ranges new when marily et al. 1988). Indeed, increases in Yellowstone bison winter range areas from 1983 onwards contributed population to growth sustained in both herds, and ecological carrying et capacity al. increased once new ranges found were (Taper 2000; Coughenour 2005; et al. 2005). Gates movements at population levels well below ecological car- rying capacity and bison the Yellowstone by used areas range large-scale winter the in Increases starvation of animals. detected were in the and 1980s continued as bison numbers increased, eventually including movements to areas outside In the winter, Yellowstone bison spend nearly one-third of one-third bison spend nearly Yellowstone In the winter, time displacing snow. their foraging model (Coughenour 2005). ANNA V A There There is substantial variability in the proportion of bison migrating outside the park each winter due to random to due winter each park the outside migrating bison variations in climate that affect the abundance and acces- bison sibility spend of food (Gates et al. 2005).Yellowstone the majority of their time finding and eating forage during winter, with nearly one-third of that time spent displacing snow to reach forage (Bruggeman 2006; Bruggeman et al. 2009).Thus, snow is the primary factor that reduces forag- ing efficiency and bison prefer patches with minimal snow- pack compared to the surrounding landscape (Bruggeman 2006). As snow depth increases, the available foraging area for Yellowstone bison is reduced to increasingly limited estimated food-limited carrying capacity of their range inside range their of capacity carrying food-limited estimated YNP (Bruggeman et more bison al. began 2009). However, migrating earlier to lower-elevation winter ranges, includ- ing outside the park, as density suggesting increased, migra- tion provided greater access to food supplies and allowed more efficient use of resources year-round (Meagher 1998, et al. 2009).Bruggeman Figure 3. Primary (B) components (A) and spatial structure 3. Figure of the S Even if the risk of brucellosis transmission could be eliminated from Yellowstone bison, it is unlikely these massive animals would be tolerated in most areas outside Yellowstone National Park due to social and political barriers…
the park (Taper et al. 2000; Gates et al. 2005). There were Yellowstone and Madison river watersheds historically oper- also pulses of dispersal from the central herd to the northern ated at a scale larger than YNP and recent migration and dis- range during this period (Coughenour 2005; Fuller et al. persal movements by Yellowstone bison represent an attempt 2007a; Bruggeman et al. 2009). These dispersal movements to operate at this larger scale (Gates et al. 2005; Plumb et al. began when population size increased above 1,500 bison for 2009). These movements are a natural process resulting from the central herd and 550 for the northern herd (Gates et successful conservation and population increases inside the al. 2005), which is well below the mean estimates of food- park. However, for much of the past 100 years as Yellowstone limited carrying capacity for Yellowstone bison (Coughenour bison recovered from near extirpation, they were constrained 2005). These findings suggest there was increased competi- to two herds that migrated to discrete wintering areas and did tion for food supplies, even though less than one-half of the not regularly and extensively venture outside the park. This total forage was eaten. Higher-quality foraging areas for bison led to the widespread belief that Yellowstone bison should in YNP are limited in overall area, patchily-distributed, and remain in YNP, which is reflected in the status and authority likely depleted first (Cheville et al. 1998; Bruggeman 2006). for management afforded to bison in states adjacent to the Also, severe winter snow conditions have prompted dispersal park. Idaho’s Comprehensive Wildlife Conservation Strategy movements of bison to low-elevation meadows beyond their mentions bison as a species of concern that is critically imper- historical winter range, as well as pulses of dispersal from the iled, but the state’s agricultural regulations do not recognize central herd to the range of the northern herd, where there is wild bison and consider them livestock. The State of Montana less snowpack and more energy efficient foraging (Meagher considers the Yellowstone bison population to be wildlife, but 1989; Gates et al. 2005; Fuller et al. 2007; Bruggeman et al. authorizes the Department of Livestock to remove or destroy 2009). Yellowstone bison that enter Montana due to the risk of dis- ease transmission to cattle. The State of Wyoming has desig- Implications for Bison Conservation nated specific areas adjacent to Grand Teton National Park and YNP where bison are considered wildlife, but elsewhere Plains bison evolved in the variable climatic environments of they are considered livestock. Even if the risk of brucellosis the North American central grasslands (McHugh 1972) and transmission could be eliminated from Yellowstone bison, it adapted to this variability through large-scale movements is unlikely these massive animals would be tolerated in most (Moodie and Ray 1976; Hanson 1984). Bison occupying the areas outside YNP due to social and political barriers such N N PS PS
Bison moving west along the roadway in the upper Madison Bison select the most productive habitats within the valleys Valley. Each spring a pulse of migrating bison moves out of they occupy. This group of bison have migrated to a lower the interior ranges of the park to seek low-elevation areas elevation and found an irrigated meadow on private land to where spring green up occurs earlier. graze during the early spring.
12 Yellowstone Science 19(1) • 2011 as human safety concerns (e.g., motor- varies on a decadal scale between 2,500 major modeling and field studies of graz- ists), conflicts with private landowners and 4,500 animals should satisfy the ing ecosystems and assessments of ungu- late carrying capacities in Yellowstone and (e.g., property damage), depredation collective long-term interests of stake- Rocky Mountain national parks, and the of agricultural crops, competition with holders, as a balance between the park’s Pryor Mountain Wild Horse Range. livestock grazing, lack of local public forage base, conservation of the genetic C support, and lack of funds for state integrity of the bison population, pro- ou r t e management (Boyd 2003). Since the tection of their migratory tendencies, sy
of evolution of a substantially larger bison brucellosis risk management, and other G. Pl conservation area outside of YNP is the societal constraints (Plumb et al. 2009). umb prerogative of the states in the Greater Yellowstone area, the prevailing social Acknowledgments carrying capacity of Yellowstone bison is perhaps most limiting (Plumb et al. Portions of this article were taken with kind permission from Elsevier: Plumb, G.E., P.J. 2009). White, M.B. Coughenour, and R.L. Wallen. With the exception of Yellowstone 2009. Carrying capacity, migration, and bison, the plains bison is considered eco- dispersal in Yellowstone bison. Biological logically extinct across North America Conservation, 14:2377–2387 ©2009, Elsevier. (Freese et al. 2007). Thus, conservation The SAVANNA model for the Yellowstone ecosystem was funded by the US Geological of the migratory and nomadic tenden- Survey Greater Yellowstone Bison Research Glenn E. Plumb spent 12 years in cies of Yellowstone bison, as well as Initiative at the Northern Rocky Mountain Yellowstone National Park before stepping their genetic integrity and ecological Science Center, Bozeman, Montana. into his current role as the chief wildlife biologist for the NPS Biological Resource role, is paramount for the perpetuation N PS/Bla Management Division in Fort Collins, of the species (Sanderson et al. 2008).
ckfo Colorado. Plumb has served as Chair of the Approximately 1,000–2,000 Yellow US Animal Health Association committee on r stone bison likely are needed in each d Brucellosis, a member of the DOI Strategic of the central and northern herds to Science Working Group for the Deepwater Horizon oil spill, and a consulting brucello- retain enough genetic diversity to enable sis scientist with the US State Department adaptation to a changing environment Bureau of International Security and Non- through natural selection, drift, and proliferation/Cooperative Threat Reduction. mutation (Gross et al. 2006; Freese et al. N PS 2007). Also, many thousands of bison are likely necessary to fully express their ecological role through the creation of landscape heterozygosity, nutrient redistribution, competition with other ungulates, prey for carnivores, habitat P.J. White is the Branch Chief of Aquatic creation for grassland birds and other and Wildlife Resources at the Yellowstone Center for Resources in Yellowstone species, provision of carcasses for scaven- National Park. He has worked in Yellow gers, stimulation of primary production, stone since 2002. White received his BS and opened access to vegetation through from Cornell University, MS from the snow cover (Freese et al. 2007; Sanderson University of Minnesota, and PhD in wildlife et al. 2008). However, it appears that ecology from the University of Wisconsin. In 2010 he received the NPS Intermountain the interactive effects of severe winters Region’s Directors Award for Natural with population levels greater than Resource Management. 4,700 bison could induce large-scale Michael B. Coughenour is a Senior movements of bison to lower-elevation Research Scientist at Colorado State winter range outside YNP (Geremia et University. He received his PhD from Colorado State University, specializing in Rick L. Wallen has been the lead al. 2010). Such large movements jeopar- systems ecology and nutrient cycling in Wildlife Biologist for the Bison Program dize brucellosis risk management objec- a southern Montana grassland. He later at Yellowstone National Park, since 2002. tives by overwhelming land managers’ studied the Serengeti grazing ecosystem He received his MS from Montana State abilities to maintain separation between of Tanzania, using simulation modeling and University–Bozeman. Prior to his work in Yellowstone, Wallen was involved in ungu- bison and livestock. Thus, we propose experimental studies to determine how the ecosystem supports the world’s largest late management programs in Grand Teton, that a Yellowstone bison population that ungulate herds. He has carried out several Redwood, and Bryce Canyon national parks.
19(1) • 2011 Yellowstone Science 13 Literature Cited Bjornlie, D.D., and R.A. Garrott. 2001. Effects outside protected areas. National Park Ser- Sanderson, E.W., K.H. Redford, B. Weber, of winter road grooming on bison in Yel- vice, Yellowstone National Park, Wyoming. K. Aune, D. Baldes, J. Berger, D. Carter, lowstone National Park. Journal of Wildlife Gross, J.E., G. Wang, N.D. Halbert, P.A. C. Curtin, J. Derr, S. Dobrott, E. Fearn, Management 65:423–435. Gogan, J.N. Derr, and J.W. Templeton. 2006. C. Fleener, S. Forrest, C. Gerlach, C.C. Boyd, D.P. 2003. Conservation of North Effects of population control strategies on Gates, J.E. Gross, P. Gogan, S. Grassel, J.A. American bison: Status and recommenda- retention of genetic diversity in National Hilty, M. Jensen, K. Kunkel, D. Lammers, R. tions. Dissertation, University of Calgary, Park Service bison (Bison bison) herds. US List, K. Minkowski, T. Olson, C. Pague, P.B. Calgary, Alberta. Geological Survey, Biological Resources Robertson, and B. Stephenson. 2008. The Bruggeman, J.E. 2006. Spatio-temporal dynam- Division, Department of Biology, Montana ecological future of the North American ics of the central bison herd in Yellowstone State University, Bozeman, Montana. bison: Conceiving long-term, large-scale National Park. Dissertation, Montana State Hanson, J.R. 1984. Bison ecology in the North- conservation of wildlife. Conservation Biology University, Bozeman, Montana. ern Plains and a reconstruction of bison 22:252–266. Bruggeman, J.E., P.J. White, R.A. Garrott, and patterns for the North Dakota region. Plains Schullery, P., and L.H. Whittlesey. 2006. F.G.R. Watson. 2009. Partial migration in Anthropologist 29:93–113. Greater Yellowstone bison distribution and central Yellowstone bison. In The ecology of Kay, C.E. 1998. Are ecosystems structured abundance in the early historical period. In large mammals in central Yellowstone: Sixteen from the top-down or bottom-up: A new Greater Yellowstone public lands: Proceedings years of integrated field studies. R.A. Garrott, look at an old debate. Wildlife Society Bulletin of the eighth biennial scientific conference on P.J. White, and F.G.R. Watson, editors, 26:484–498. the Greater Yellowstone ecosystem. A.W. Biel, 217–235. Elsevier, San Diego, California. Kilpatrick, A.M., C.M. Gillin, and P. Daszak. editor, 135–140. National Park Service, Yel- Caughley, G. 1976. Wildlife management 2009. Wildlife–livestock conflict: The risk of lowstone National Park, Wyoming. and the dynamics of ungulate populations. pathogen transmission from bison to cattle Sinclair, A.R.E. 1975. The resource limitation In T.H. Croaker, editor, Applied Biology outside Yellowstone National Park. Journal of trophic levels in tropical grassland ecosys- 1:183–246. Academic Press, New York, of Applied Ecology 46:476–485. tems. Journal of Animal Ecology 44:497–520. New York. McHugh, T. 1972. The time of the buffalo. Taper, M.L., M. Meagher, and C.L. Jerde. 2000. Caughley, G.C., and A.R.E. Sinclair. 1994. Alfred A. Knopf, New York, New York. The phenology of space: Spatial aspects of Wildlife ecology and management. Blackwell Meagher, M. 1973. The bison of Yellowstone bison density dependence in Yellowstone Science, Cambridge, Massachusetts. National Park. National Park Service, Wash- National Park. US Geological Service, Cheville, N.F., D.R. McCullough, and L. R. ington, D.C., Government Printing Office, Biological Resources Division, Bozeman, Paulson. 1998. Brucellosis in the Greater Science Monographs 1. Montana. Yellowstone area. National Academy Press, Meagher, M. 1989. Range expansion by bison of Thein, T.R., F. G.R. Watson, S.S. Cornish, T.N. Washington, D.C. Yellowstone National Park. Journal of Mam- Anderson, W.B. Newman, and R.E. Lock- Cole, G.F. 1971. An ecological rationale for the malogy 70:670–675. wood. 2009. Vegetation dynamics of Yellow- natural or artificial regulation of ungulates Meagher, M. 1993. Winter recreation-induced stone’s grazing system. In The ecology of large in parks. Transactions of the North American changes in bison numbers and distribution mammals in central Yellowstone: Sixteen years Wildlife Conference 36:417–425. in Yellowstone National Park. National of integrated field studies. R.A. Garrott, P.J. Coughenour, M.B. 2005. Spatial-dynamic Park Service, Yellowstone National Park, White, and F.G.R. Watson, editors, 113–133. modeling of bison carrying capacity in the Wyoming. Elsevier, San Diego, California. Greater Yellowstone ecosystem: A synthesis Meagher, M. 1998. Recent changes in Yel- US Department of the Interior, National Park of bison movements, population dynamics, lowstone bison numbers and distribution. In Service and US Department of Agriculture, and interactions with vegetation. Natural International symposium on bison ecology and Forest Service, Animal and Plant Health In- Resource Ecology Laboratory, Colorado management in North America. L. Irby and spection Service. 2000a. Final Environmental State University, Fort Collins, Colorado. J. Knight, editors, 107–112. Montana State Impact Statement for the Interagency Bison
14 Yellowstone Science 19(1) • 2011 Balancing Brucellosis Risk Management and Wildlife Conservation P. J. White, John J. Treanor, and Rick L. Wallen N PS
Each fall Yellowstone National Park’s Bison Ecology and Management Office monitors a group of about 30 bison cows to track pregnancy and calf rearing. They use temporary immobilization (as seen above) to assess body condition and brucello- sis status. The team also follows their migration using radio telemetry.
he increasing emergence or resurgence of infec- Human infection by B. abortus typically involves inges- tious diseases that move between livestock, wildlife, tion of the bacteria or exposure through the eyes, open and humans has raised interest in disease ecology wounds, or by direct contact with skin. Infected bison and andT wildlife health. Wildlife and diseases do not recognize elk pose a health risk for people that handle animal carcasses jurisdictional or political boundaries, and wildlife con- without proper protective equipment. Intensive efforts to servation can become challenging when risks to human eradicate B. abortus in livestock during the twentieth century health arise. Bovine brucellosis is a contagious bacterial dis- decreased infections nationally from 6,500 cases in 1940 to ease caused by Brucella abortus that may induce abortions 70 cases in 1994. There were five confirmed cases reported or the birth of non-viable calves in livestock and wildlife to the Wyoming Department of Health from 1995 to 2005, (Rhyan et al. 2009). The bacterium was likely introduced and 17 confirmed cases reported to the Idaho Department of from European livestock to Yellowstone bison and elk Health and Welfare from 1980 to 2003, though none of these before 1930 (Meagher and Meyer 1994). Infection of bison, cases were attributed to wildlife (Snow 2005). However, cattle, or elk typically occurs through contact with infec- there have been two confirmed cases of hunters contracting tious reproductive tissues shed when a pregnancy is termi- brucellosis from elk in Montana (Zanto 2005). Infection by nated during an abortion or live birth (Rhyan et al. 2009). B. abortus is rarely fatal in humans, but can cause severe,
19(1) • 2011 Yellowstone Science 15 recurring, fever-like symptoms. Humans cannot Most livestock and natural resources pass the disease to animals or other humans. personnel believe the primary sources for Most livestock and natural resources per- sonnel believe the primary sources for brucello- brucellosis transmission to other herds of elk sis transmission to other herds of elk and cattle and cattle are the supplemental feeding of are the supplemental feeding of elk in Wyoming and Yellowstone bison (Bienen and Tabor elk in Wyoming and Yellowstone bison. 2006). Thus, brucellosis management focuses on elk in the southern Greater Yellowstone area and bison in the northern portion. Elk often mingle with bison within the social group where the birth occurred. From cattle, and all recent brucellosis transmissions to cattle have 2004 to 2007, at least one bison interacted with potentially been attributed to elk (Beja-Pereira et al. 2009). Conversely, infectious material in 30% of observed births. Thus, infectious resource agencies prevent bison from mingling with cattle live births could be an important transmission event for bru- through active management practices such as hazing, hunt- cellosis among Yellowstone bison (Rhyan et al. 2009). ing, and culling (US Department of the Interior and US Yellowstone elk also exhibit a high degree of birth Department of Agriculture 2000), which may explain the synchrony, with the majority calving between mid-May lack of transmissions from bison to cattle. We review the state and mid-June. In Yellowstone, the potential for brucellosis of the knowledge regarding brucellosis transmission risk and transmission between elk is likely lower than between bison its control in bison and elk in the Greater Yellowstone area. because females segregate themselves while giving birth and also clean the birth site. Thus, birth sites are dispersed and Brucellosis Transmission Risk the likelihood of other elk encountering infected birth tissues is low. However, transmission risk may be higher during the The risk of brucellosis transmission to livestock or wildlife is potential abortion period from February through April when proportional to the amount of infectious material shed onto many elk aggregate in larger groups on lower-elevation winter the landscape where it can be contacted. Environmental fac- ranges. Spontaneous abortions under these conditions could tors such as deep snow and the presence of predators may influ- expose many susceptible elk to infectious material. During ence brucellosis transmission by increasing stress and concen- the past decade, elk abundance has increased substantially trating animals, which increases exposure to shed infectious in some areas, with large groups congregating on winter tissues. However, expelled tissues are quickly removed by ranges (Cross et al. 2010). Coincident with these changes in scavengers, and the behavior of elk and bison during calving abundance and group sizes, brucellosis seroprevalence (i.e., may limit the risk presence of antibodies to Brucella circulating in blood) has R
. S of transmission to increased 7%–20% on some non-feed ground areas (Cody cho cattle. The birth- and Buffalo Valley, Wyoming). Thus, elk populations far en h
ar i n g p e r i o d f o r from both bison and feed grounds may be becoming viable d Yellowstone bison reservoirs for perpetuating higher levels of brucellosis due is synchronous, to increased densities and group sizes on their winter ranges with 80% of births (Cross et al. 2010). occurring from late The high seroprevalence (40%–60%) of brucellosis in April to late May Yellowstone bison suggests they are a likely candidate for (Jones et al. 2010). transmission to elk, recent data suggests such transmission Birthing females is rare. The peak bison calving period occurs approximately meticulously clean one month earlier than for elk and, overall, there is little and leave birth sites overlap in distribution during the period when B. abortus within two hours is expected to be shed from each species. In areas where elk after calving, there- do mingle with bison during winter and spring, such as the by lowering the risk Madison headwaters area in Yellowstone, elk have much of brucellosis trans- lower seroprevalence rates for brucellosis (3%) than do mission to cattle Yellowstone bison or elk associated with feeding programs and bison that later in Wyoming. Proffitt et al. (2010b) found that brucellosis A bison cow and newborn calf rest encounter the birth transmission risk from bison to elk was quite low in the near the Firehole River. Eighty percent site. Although the Madison headwaters area, despite a high degree of spatial of bison births occur between late potential for expo- overlap when B. abortus is typically shed. Predation risk asso- April and late May. sure is higher for ciated with wolves increased elk and bison spatial overlap
16 Yellowstone Science 19(1) • 2011 N PS/Pea 2006). Wyoming has 22 state feed grounds and one federal co feed ground (National Elk Refuge, US Fish and Wildlife Service) where feeding reduces elk foraging on cattle ranches, but consequently sustains higher numbers of elk than the remaining winter habitat could otherwise support (Cross et al. 2007). Additionally, supplemental feeding creates large elk aggregations that facilitate brucellosis transmission (Bienen and Tabor 2006; Cross et al. 2007). Wildlife and livestock managers remain concerned that reduced feeding of elk would lead to increased brucellosis transmission to cattle as a result of increased mingling of elk and cattle (Cross et al. 2007). In the interim, strategies that reduce the length of the feeding season and the duration of elk aggregation during the peak transmission period (February through May) may N PS/Pea decrease brucellosis seroprevalence in elk on feed grounds in
co the southern Greater Yellowstone area (Cross et al. 2007). Another factor that may influence the maintenance of brucellosis in elk in the Greater Yellowstone area is the increasing human population and land use practices that sus- tain it. Ranching and the development of rural homes have fragmented valley bottom and flood plain habitats crucial for elk migration and use during winter. The conversion of traditional winter habitat for human use has contributed to an increase of elk on cattle feed lines and refuges that results in large elk aggregations (Haggerty and Travis 2006; Cross et al. 2010). Studies of migratory elk near Yellowstone National Most elk births occur between mid-May and mid-June Park suggest they tend to select areas during winter with a (above, June 9, 2010). Female elk thoroughly clean the lower probability of wolf occupancy, lower road density, birthing site and then leave soon after when the calf (here, and higher forage abundance (Proffitt et al. 2010a). Many age 23 minutes) is ready to begin walking. of these areas occur on lower-elevation private ranchlands and portions of public grazing allotments. Effective brucel- losis control measures should recognize that elk aggregating temporarily, but these behavioral responses by elk did not on feed grounds, natural winter ranges, or refuges influence have important disease implications. Also, DNA genotyp- brucellosis transmission, especially near the calving period in ing indicates a relatively high genetic divergence between late winter and early spring. Thus, wildlife agencies need to B. abortus isolates from bison and elk, which suggests that explore strategies for dispersing large aggregations of elk in B. abortus is not extensively exchanged between bison and elk late winter and spring, such as gaining enhanced cooperation (Beja-Pereira et al. 2009). It appears that brucellosis in the from landowners to increase access for hunters, providing Greater Yellowstone area is a disease sustained by multiple increased tolerance and protection of large predators such as hosts and control measures aimed at managing the risk of wolves that may disperse elk, and assisting landowners with transmission to cattle will require addressing both wildlife infrastructure to isolate cattle and their feed from wildlife. reservoirs and the factors maintaining infection. Bison—After intensively managing bison numbers for 60 Brucellosis Control years through husbandry and regular culling, the National Park Service instituted a moratorium on culling in the park Elk—The best available scientific information suggests in 1969 and allowed bison numbers to fluctuate in response that elk-to-elk transmission is primarily responsible for the to weather, predators, and resource limitations. Abundance observed levels of B. abortus exposure within elk popula- increased rapidly and large-scale bison migrations out of the tions and the risk of their transmitting brucellosis to cattle. park during winter began in the late 1980s. These migrations There is general agreement that as sustainable alternatives led to a series of conflicts with stock growers and the state for maintaining elk numbers are developed, supplemental of Montana, largely because of the possibility of brucellosis feeding should be phased out, which may lead to decreased transmission to cattle. As a result, in 2000 the federal govern- brucellosis seroprevalence over time (Bienen and Tabor ment and the state of Montana agreed to a court-negotiated
19(1) • 2011 Yellowstone Science 17 Montana ± Idaho Wyoming
? ?
? ? ?
? Private Land with Cattle Pre-Settlement Bison Distribution IBMP Intensive Management Zones Major Bison Use Areas Yellowstone National Park Roads Rivers Lakes 05 10 20 30 40 50 Miles Figure 1. Bison management zones and major use areas in and near Yellowstone National Park.
Interagency Bison Management Plan (IBMP) that estab- 2011). However, this intensive management is expensive, lished guidelines for (1) cooperatively managing the risk of logistically taxing, and controversial due to culls of more brucellosis transmission from bison to cattle, and (2) preserv- than 1,000 bison which have occurred in some winters ing the bison population and allowing some bison to occupy when large numbers of bison have migrated outside the winter ranges on public lands in Montana (fig. 1). park. The IBMP is designed to adaptively progress through a To reduce the risk of brucellosis transmission to cat- series of management steps that initially tolerate only bison tle, recurrent, small (less than 100) to large (about 1,700) testing negative for brucellosis exposure on winter ranges numbers of bison have been shipped to domestic slaughter outside Yellowstone National Park, but will eventually facilities following capture near park boundaries since the tolerate limited numbers of untested bison on key winter late 1980s. Despite these actions, brucellosis seroprevalence ranges adjacent to the park when cattle are not present (US in Yellowstone bison has not decreased (Kilpatrick et al. Department of the Interior et al. 2000, 2008). The IBMP 2009). This non-random culling strategy serves more as a uses intensive management (e.g., hazing, hunting, and culls) population reduction program than to reduce brucellosis of bison migrating outside the park to maintain separation (Bienen and Tabor 2006). Intensifying this strategy to a between bison and cattle. The agencies have successfully level that would be effective at reducing brucellosis trans- maintained spatial and temporal separation between bison mission would be extremely expensive, unacceptable to the and cattle with no transmission of brucellosis (White et al. public, and questionable as a management practice given
18 Yellowstone Science 19(1) • 2011 Service policy allows for the use of reproductive intervention Another factor that may influence in wildlife if these techniques are appropriate for achieving the maintenance of brucellosis management goals. Thus, if an effective, reliable, and safe in elk in the Greater Yellowstone contraceptive was developed, contraception of seropositive bison or elk might be considered for decreasing brucellosis area is the increasing human transmission; especially when combined with the vaccina- population and land use practices tion of seronegative animals (Ebinger et al. 2010). However, fertility control products may also cause negative long-term that sustain it. effects, such as permanent sterility, altered reproductive or social behaviors, and changes in the age and sex structure of a population. Thus, it is uncertain whether available fertility the National Park Service policy to maintain ecosystem control products can effectively decrease brucellosis infection integrity (Bienen and Tabor 2006; White et al. 2011). in free-ranging bison and elk over a reasonable time frame A more effective and acceptable approach to brucel- without unacceptable side effects. losis reduction might involve management actions on Until effective brucellosis reducing methods are devel- selected bison in combination with brucellosis vaccination. oped, the best approach to control the risk of brucellosis Approaches that target pre-reproductive females for vacci- transmission from bison to cattle is to maintain spatial and nation, while removing reproductively active, seropositive temporal separation. Management agencies should continue females may be effective at reducing disease transmission to allow bison migration to essential winter range areas in (Ebinger et al. 2010). Consistent vaccination of female bison and adjacent to Yellowstone National Park, but actively pre- has the potential to reduce brucellosis prevalence by increas- vent dispersal and range expansion to outlying private lands ing herd immunity, especially if vaccine technology and until there is tolerance for bison in these areas (Plumb et methods for remote vaccine delivery to free-ranging wildlife al. 2009). Bison abundance and distribution on lands adja- are improved (Treanor et al. 2010). However, a vaccine with cent to Yellowstone can be adjusted based on evaluations of low or medium efficacy is unlikely to succeed in controlling available habitat, new conservation easements or land man- brucellosis in the long term without the eventual inclusion agement strategies, reduced brucellosis prevalence in bison, of test and slaughter or fertility control (Ebinger et al. 2010; and new information or technology that reduces the risk of Treanor et al. 2010). Thus, the development of more effec- disease transmission (US Department of the Interior et al. tive vaccines, delivery methods, and diagnostic techniques is 2008). However, the comprehensive wildlife conservation urgently needed. Until substantial improvement is made in strategies of Idaho, Montana, and Wyoming express little these areas, proposed vaccination efforts will, at best, only support for resident, free-ranging wild bison (Plumb et al. result in a relatively slow decline in brucellosis seroprevalence 2009). over decades (Treanor et al. 2010). To increase tolerance for bison, Kilpatrick et al. (2009) Contraception has also been suggested as a method to recommended establishing a local brucellosis infection status reduce brucellosis transmission because the disease is known zone for cattle in the Greater Yellowstone area and testing all to only be transmitted by pregnant bison or elk. National Park cattle within this area for brucellosis (with a “split status” for N PS/D. S t a h ler
Predator species influence the migration patterns and population density of native ungulates. These factors then affect transmission and prevalence rates of disease among herds.
19(1) • 2011 Yellowstone Science 19 Chronic Chronic brucellosis infection does not adversely affect the Conservation Implications bison and elk both of conservation the for step first essential An is to develop acceptable brucellosis control methods that bal- ance risk management with wildlife conservation. Over time, need and costs the reduce could herein discussed strategies the for brucellosis risk management activities, while maintaining the potential for elk low risk for However, the cattle industry. to serve as a brucellosis maintenance host in the absence of bison and away from feed grounds control. complicates Brucellosis is brucellosis present in elk populations throughout Greater area, Yellowstone where they number approximately 50,000. Eradication of brucellosis from these without populations technology is current with practical or possible not likely resorting to ethically and politically unacceptable techniques such as mass test and slaughter or depopulation (US Animal in brucellosis of management Thus, 2006). Association Health elk populations might best be achieved by curtailing practices that unnaturally increase elk densities and group sizes during the potential abortion period. 2009). al. et (Geremia bison Yellowstone of viability long-term Thus, the Yellowstone bison population will likely continue to and grow attempt to expand its range unless hunting, cull- ing, and/or relocations are used to remove several Thus 2009).al. et (Hobbs population the from year per bison hundred bison far, Yellowstone have only been transported to domes- tic slaughter or research facilities due infection with to brucellosis. These removals the have differentially potential for affected the central and northern breeding herds, altered gen- years, some in females of groups reduced created structure, der and temporarily dampened productivity (White et al. 2011). 1,000 than (more culls large-scale over continued If are bison) time, these effects could diminish the ecological role of the largest remaining free-ranging plains bison population in the processes ecological the diminish would turn, in which, world within the park (White et al. 2011). Thus, there is a need to increase tolerance for bison on key winter ranges in Montana population. the of culls large-scale of frequency the reduce and The shipment of “surplus” Yellowstone bison to quarantine conservation other or tribes Indian American by operated sites would facilities slaughter domestic to than rather organizations be a transformational moment in the American conservation of of culture the plains preserve help would action This bison. Indians while promoting bison conservation by establishing the from populations existing augmenting or populations wild of wild plains bison. source only existing pure
NPS 19(1) • 2011
[Bison] removals have differentially affected the central and northern breeding breeding northern and the central affected differentially have removals [Bison] years, in some of females groups reduced created structure, gender altered herds, productivity… dampened and temporarily Kilpatrick et al. (2009) also recommended the cessation the recommended also (2009) al. et Kilpatrick Yellowstone Science Yellowstone Some studies recommend the cessation of cattle grazing in Some studies recommend the park in winter and improved bison leave where areas strategies in collaboration with private landowners. fencing of cattle grazing in areas where bison leave the park in win- ter and compensating ranchers for lost earnings and wages. success- have agencies government and groups Conservation fully used, and are still pursuing, this strategy with willing landowners (US Department of the Interior et al. 2008). However, further efforts are needed to develop identify Montana, in additional bison for areas conservation and habitat fencing strategies in collaboration with private landowners that raise susceptible cattle, and identify opportunities for to Montana in habitat bison of creation or enhancement the sustain bison from April to early and June discourage bison movements onto private lands with cattle (US Department et al. 2008).of the Interior the remaining portions of Idaho, Montana, and Wyoming). the remaining portionsWyoming). and of Montana, Idaho, In 2010, the Animal and Plant Health Inspection Service published an status interim lower a to rule area or State that Free Class any eliminated of reclassification the automatic if two or more herds are found to have brucellosis within a and Animal Agriculture, of Department (US period two-year detec- rule, this Under 2010). Service Inspection Health Plant tions of brucellosis in domestic livestock within the Greater case-by-case a on with dealt are area surveillance Yellowstone basis, which should eliminate many economic barriers cre- ated by the brucellosis class status system. As long outbreaks are as investigated the and contained, state brucellosis- brucellosiswas fact, In change. not does cattle for status free Idaho, in herds cattle and bison domestic several in detected any without 2010, and 2009 during Wyoming and Montana, changes in state status. 20 P.J. White is the Branch Chief of Aquatic Literature Cited with livestock during the brucellosis trans- and Wildlife Resources at the Yellowstone mission risk period. Journal of Applied Ecology Beja-Pereira, A., B. Bricker, S. Chen, C. DOI: 10.1111/j.1365-2664.2010.01928.x. Center for Resources in Yellowstone Almendra, P.J. White, and G. Luikart. 2009. Proffitt, K.M., P.J. White, and R.A. Garrott. National Park. He has worked in Yellow DNA genotyping suggests recent brucel- 2010b. Spatio-temporal overlap between stone since 2002. White received his BS losis outbreaks in the Greater Yellowstone Yellowstone bison and elk – implications for from Cornell University, MS from the area originated from elk. Journal of Wildlife wolf restoration and other factors for brucel- University of Minnesota, and PhD in wildlife Diseases 45:1174–1177. losis transmission risk. Journal of Applied ecology from the University of Wisconsin. Bienen, L., and G. Tabor. 2006. Applying an Ecology 47:281–289. In 2010 he received the NPS Intermountain ecosystem approach to brucellosis control: Rhyan, J.C., K. Aune, T. Roffe, D. Ewalt, S. Can an old conflict between wildlife and Region’s Directors Award for Natural Hennager, T. Gidlewski, S. Olsen, and R. agriculture be successfully managed? Frontiers Clarke. 2009. Pathogenesis and epidemiology Resource Management. in Ecology and the Environment 4:319–327. of brucellosis in Yellowstone bison: serologic Cross, P.C., E.K. Cole, A.P. Dobson, W.H.
C and culture results from adult females and ou Edwards, K.L. Hamlin, G. Luikart, A.D. their progeny. Journal of Wildlife Diseases r t
e Middleton, B.M. Scurlock, and P.J. White.
sy 45:729–739.
of 2009. Probable causes of increasing brucel- Snow, J. 2005. November 3, 2005 electronic J. Trean losis in free-ranging elk of the Greater mail message to J. MacDonald, Greystone En- Yellowstone Ecosystem. Ecological Applications vironmental Consultants, Greenwood Village, o 20:278–288. r Colorado regarding the number of human Cross, P.C., W.H. Edwards, B.M. Scurlock, E.J. cases of brucellosis in Wyoming. State Public Maichak, and J.D. Rogerson. 2007. Effects of Health Veterinarian, Wyoming Department management and climate on elk brucellosis in of Public Health, Cheyenne, Wyoming. the Greater Yellowstone ecosystem. Ecologi- Treanor, J.J. , J.S. Johnson, R.L. Wallen, S. Cilles, cal Applications 17:957–964. P.H. Crowley, J.J. Cox, D.S. Maehr, P.J. White, Ebinger, M.R., P. Cross, R. Wallen, P.J. White, and G.E. Plumb. 2010. Vaccination strate- and J. Treanor. 2010. Modeling brucellosis gies for managing brucellosis in Yellowstone management in bison: A heuristic model of bison. Vaccine 28S:F64-F72. management actions on disease dynamics. US Animal Health Association. 2006. Enhanc- Yellowstone Center for Resources, Mam- ing brucellosis vaccines, vaccine delivery, and moth, Wyoming. surveillance diagnostics for elk and bison in Geremia, C., P.J. White, R.A. Garrott, R. Wal- the Greater Yellowstone area: A techni- len, K.E. Aune, J. Treanor, and J.A. Fuller. cal report from a working symposium held 2009. Demography of central Yellowstone August 16–18, 2005 at the University of bison: effects of climate, density and disease. Wyoming. T. Kreeger, and G. Plumb, editors. Pages 255–279 in R.A. Garrott, P.J. White, The University of Wyoming Haub School and John Treanor is a wildlife biologist for the and F.G.R. Watson, editors. The ecology of Ruckelshaus Institute of Environment and large mammals in central Yellowstone: Sixteen Yellowstone Wildlife Health Program at Natural Resources, Laramie, Wyoming. years of integrated field studies. Elsevier, San US Department of Agriculture, Animal and Yellowstone National Park. John coordi- Diego, California. Plant Health Inspection Service. 2010. nates a diversity of wildlife health projects Haggerty, J.H., and W.R. Travis. 2006. Out of Brucellosis class free states and certified involving bison, bats, rodents, birds, and administrative control: Absentee owners, brucellosis-free herds; revisions to testing amphibians. John is also a doctoral student resident elk, and the shifting nature of wildlife and certification requirements. Federal Regis- at the University of Kentucky studying bru- management in southwestern Montana. ter 75:81090. cellosis infection in Yellowstone bison and Geoforum 37:816–830. US Department of the Interior, National Park potential management applications. Hobbs, N.T., R. Wallen, J. Treanor, C. Geremia, Service and United States Department of and P.J. White. 2009. A stochastic population Agriculture, Forest Service, Animal and Plant model of the Yellowstone bison population. Health Inspection Service. 2000. Record of Colorado State University, Fort Collins, decision for final environmental impact state- Colorado. ment and bison management plan for the Jones, J. D., J.J. Treanor, R.L. Wallen, and P.J. State of Montana and Yellowstone National White. 2010. Timing of parturition events Park. Washington, D.C. in Yellowstone bison: Implications for bison US Department of the Interior, National Park conservation and brucellosis transmission Service and US Department of Agriculture, risk to cattle. Wildlife Biology 16:1–7. Forest Service, Animal and Plant Health Kilpatrick, A.M., C.M. Gillin, and P. Daszak. Inspection Service, and the State of Montana, 2009. Wildlife-livestock conflict: The risk of Department of Fish, Wildlife, and Parks, pathogen transmission from bison to cattle Department of Livestock. 2008. Adap- outside Yellowstone National Park. Journal of tive adjustments to the interagency bison Applied Ecology 46:476–485. management plan. National Park Service, Meagher, M., and M.E. Meyer. 1994. On the Yellowstone National Park, Wyoming. origin of brucellosis in bison of Yellowstone White, P.J., R.L. Wallen, C. Geremia, J.J. Rick L. Wallen (right) has been the lead National Park: A review. Conservation Biology Treanor, and D.W. Blanton. 2011. Manage- Wildlife Biologist for the Bison Program 8:645–653. ment of Yellowstone bison and brucellosis Plumb, G.E., P.J. White, M.B. Coughenour, at Yellowstone National Park since 2002. transmission risk: Implications for conserva- and R.L. Wallen. 2009. Carrying capacity, tion and restoration. Biological Conservation, He received his MS from Montana State migration, and dispersal in Yellowstone bison. DOI:10.1016/j.biocon.2011.01.003. University–Bozeman. Prior to his work in Biological Conservation 142:2377–2387. Zanto, S. 2005. Montana Public Health and Yellowstone, Wallen was involved in ungu- Proffitt, K.M., J.A. Gude, K.L. Hamlin, R.A. Safety Division, Public Health Laboratory, late management programs in Grand Teton, Garrott, J.A. Cunningham, and J.L. Grigg. personal communication with L. Bambrey, Redwood, and Bryce Canyon national parks. 2010a. Elk distribution and spatial overlap Greystone, August 3, 2005.
19(1) • 2011 Yellowstone Science 21 Y
The Lamar Buffalo Ranch ELL Cori Knudten -5517/ R . R . Robi n so s the vast herds of bison disappeared from the American n A West in the nineteenth century, they became a symbol of the vanishing frontier. By 1885 the bison herd in Yellow- stone National Park was the largest existing bison popula- tion south of Canada (Bartlett 1985). In 1901 park officials attempted to increase bison numbers by starting a captive herd with bison purchased from ranch- ers. An enclosure was constructed near Fort Yellowstone at Corralled bison, circa 1912–1920. Winter feeding of bison at the Mammoth Hot Springs, and 15 cows and 3 bulls were trans- Lamar Buffalo Ranch was discontinued in 1952. ported to the park (Haines 1977). By 1907 the herd was too large for the Mammoth enclosure and was moved to a site on Rose Creek in the Lamar Valley, which became known In 1933 a group of National Park Service biologists as the Lamar Buffalo Ranch. At Lamar, park employees used compiled Fauna of the National Parks of the United States: A cattle ranching techniques to monitor and manage the bison Preliminary Survey of Faunal Relations in National Parks. The herd. Bison roamed freely in the summer, but in the winter report was based on a multi-year survey of wildlife issues they were corralled and fed hay that was raised at the ranch in national parks across the country. It was intended to on 600 acres of plowed, seeded, and irrigated land. The herd be the “first comprehensive statement of natural resource grew from fewer than 50 animals in 1902 to around 950 by management policies” in the National Park Service. The 1929 (Haines 1977; Sellars 1997). biologists argued for letting natural processes dominate The Lamar Buffalo Ranch was a popular attraction with when possible and for restoring parks and their wildlife tourists because they could easily see the bison in the cor- populations to resemble their condition prior to the arrival rals. In the 1920s, under Superintendent Horace Albright, of Europeans. The report stated explicitly that every wildlife large round-ups and stampedes were held for the amuse- species should be allowed to “carry on its struggle for ment of visitors. Although the program started over con- existence unaided” unless a species was threatened with cerns that bison might become extinct in the park, managing extinction (Sellars 1997). the herd as a tourist attraction quickly became an important This publication eventually contributed to a major shift in part of the ranch’s operation. Other wild bison still roamed the operation of the Lamar Buffalo Ranch. As new attitudes the park, but because they ranged in more remote areas they took hold in the National Park Service and Yellowstone, the were subject to little or no interference by park managers function of the Lamar Buffalo Ranch changed with them. (Haines 1977, Sellars 1997). Newton B. Drury, who became Director of the National By the late 1920s, some worried that the population of the Park Service in 1940, stated that Yellowstone was not a Lamar Valley herd was becoming too large. In order to re- “zoological park or game farm” and that the bison should duce the population, the park shipped bison to preserves and be allowed to become “self-sustaining” (Sellars 1997). The zoos around the country and slaughtered “surplus” bison. last winter feeding took place in 1952, and the operations at the ranch were discontinued.
Y The historic buildings at the Lamar Buffalo Ranch cur- ELL
-27615 rently serve as instruction and lodging space for the non- profit Yellowstone Association Institute.
Literature Cited Bartlett, R.A. 1985. Yellowstone: A Wilderness Besieged. University of Ari- zona Press, Tuscon, Arizona. Haines, A.L. 1977. The Yellowstone Story: A History of Our First National Park. Vol. 2. Rev. ed. University Press of Colorado, Niwot, Colorado. Sellars, R.W. 1997. Preserving Nature in the National Parks: A History. Yale University Press, New Haven, Connecticut. Wright, G.M., J.S. Dixon, and B.H. Thompson. 1933. Fauna of the National Parks of the United States: A preliminary survey of faunal relations in National Parks. US Government Printing Office, Washington, DC. The “show herd” in the Mammoth Hot Springs pen, 1924.
22 Yellowstone Science 19(1) • 2011 How Many Wolves Were in the Yellowstone Area in the 1870s? A Revealing Account from 1872 Lee Whittlesey and Paul Schullery
This photo, taken by William Henry Jackson on July 17 or 18, 1871, while he was with the Hayden survey, reveals the substantial scale of the Bottlers’ hide-hunting operations by that year.
NPS
rior to the 1995 restoration of wolves to Our initial publication (Schullery and Whittlesey 1992) Yellowstone National Park (YNP), we researched the analyzed eight times as many accounts as any previous review historical record of large- and medium-sized wild- of the historical record of GYE wildlife, but it was clear to Plife species in the Greater Yellowstone Ecosystem (GYE). us that many hundreds of additional unanalyzed accounts In our initial review of 168 distinct accounts of visits to awaited attention. Besides the customary trapper and pros- the GYE from 1796 through 1881, we found numerous pector diaries, tourist accounts, official survey party reports, general statements indicating that wolves were present or administrative documents, and other readily accessible common and more specific statements reporting individ- material, Yellowstone was the subject of countless articles in ual sightings, howls, or other substantive evidence of wolf newspapers, sporting periodicals, other narrowly distributed presence (Schullery and Whittlesey 1992, 1995). These periodicals, and other sources that are still being discovered. sources of evidence were buttressed by the official admin- Since that initial publication, we have searched for istrative record, especially Yellowstone Superintendent additional information and, with the assistance of additional Philetus Norris’s statement in his annual report for 1880: investigators, compiled a much larger volume of material and a database that further confirm Norris’s assertion that wolves The large, ferocious gray or buffalo wolf, the sneak were widespread and abundant in the park prior to their extir- ing, snarling coyote, and a species apparently between pation by Euro-Americans (Schullery et al. forthcoming). the two, of a dark-brown or black color, were once However, until recently we had not found any documents exceedingly numerous in all portions of the Park, but that allowed us to resolve early Yellowstone-area wolf num- the value of their hides and their easy slaughter with bers any further than the general impressions we published strychnine-poisoned carcasses of animals have nearly in 1992. We have now located one such document, but it is led to their extermination (Norris 1881). necessary first to provide the context for its interpretation.
19(1) • 2011 Yellowstone Science 23 Coming to Terms with Historical Wolf Numbers It is certain, however, that wolves were proportionately abundant Determining historical wildlife numbers in a large geograph- ical area has vexed generations of researchers. The abundance with the bison they preyed and of wolves throughout much of the American West was a mat- scavenged upon in the American ter of historical record, but except for the most general of esti- mates it has been impossible to calculate that abundance with West at the beginning of the any precision. It is almost as difficult to calculate the number nineteenth century. of wolves killed during the slaughter of western wildlife that became widespread starting in the 1860s. Record keeping was often casual, species were sometimes confused (Norris’s before the arrival of Europeans. (Seton 1953). More recently, description of the wild canids in the park is one of many Edward Curnow, whose “The History of the Eradication of examples of this taxonomic uncertainty), and the conditions the Wolf in Montana” (1969) served for some time as a pri- under which the killing took place were not conducive to mary scholarly source text, estimated that in the 1860s there precise accounting or the durability of records (Hampton were “several hundred thousand wolves” in the region that 1997). would become Montana. He said that the number of wolves It is certain, however, that wolves were proportionately killed during the 1860s and 1870s was unknown, but “a con- abundant with the bison they preyed and scavenged upon servative estimate would be over 100,000 per year between in the American West at the beginning of the nineteenth 1870 and 1877.’’ He illustrated the scale of the trade in wolf century. Raymond Burroughs (1961) compiled numerous hides by quoting from an 1873 article in The Daily Herald reports of wolves from the journals of Lewis and Clark, a few of Helena, which reported that a group of “wolfers” using representative examples of which follow, all recorded in what five or six teams of horse-drawn wagons accumulated “about is now Montana. In June 1805, near the mouth of the Marias 10,000 wolf skins among them” in one winter. River, Lewis said that the “innumerable herds of Buffalo were It is important to distinguish between the killing of seen attended by their shepherds the wolves.” In July 1806, wolves by early professional “wolfers” in Montana and YNP, Lewis reported that there were “vast assemblages of wolves” and the later killing of wolves for a state-sponsored bounty. near the Sun River. In late July 1806, on the Yellowstone In the 1870s and early 1880s, wolfers killed wolves for, as River near present Billings, Clark said that “for me to men- Norris said, “the value of their hides.” There was a boom- tion or give an estimate of the different species of wild ani- ing market for wolf hides, just as there was for the hides mals on this river particularly Buffalow, Elk, Antelopes and of other wildlife species (Zmyj 1996; Hampton 1997). After Wolves would be incredible. I shall therefore be silent on the 1883, when Montana’s territorial legislature passed the ter- subject further.” ritory’s “first workable bounty law” (Curnow 1969), killing Historians and naturalists have made occasional attempts was still done for money, but the primary purpose was to to estimate the number of wolves inhabiting all or parts of protect livestock. the American West prior to the arrival of white settlers. The Thanks to the more consistent paperwork required by naturalist Ernest Thompson Seton estimated about 2 mil- the bounty laws, we have a somewhat better grasp of the lion wolves in all of North America and northern Mexico number of wolves killed during the bounty period. From 1883 to 1918, Montana recorded bounty claims on more
N than 80,000 wolves; between 1895 and 1917, Wyoming PS Yellowstone National Park’s recorded almost 30,000 (Hampton 1997). However, it is second superintendent, thought that these official tallies may overstate the number Philetus W. Norris, regularly of wolves killed because bounty hunters were creative and chronicled the park’s wildlife energetic in defrauding authorities (Zmyj 1996). Still, the in his annual reports and historical records, including the one presented below for the other writings. His notes Yellowstone area, are sobering evidence of a massive slaughter provide the first official of Montana and Wyoming wolves after their numbers had record of hide-hunting and already been significantly reduced by the hide-hunters. related killing of wildlife in Subsequent estimates of the Montana wolf population the park. Norris was one of and the number of wolves killed in the 1870s were lower several commentators who than Curnow’s. Dave Walter (1986) thought that the annual spoke highly of the Bottlers kill of Montana wolves in the 1870s was on the order of despite their wholesale 55,000. Bruce Hampton (1997) regarded Walter’s number as slaughter of park animals. still too large, pointing out many problems with early tallies,
24 Yellowstone Science 19(1) • 2011 .03 (1869) No ana t n o WCS-049-OMF-M N
. No p / Ma s e chiv r A nal tio a N
A section of Lieutenant Gustavus Doane’s “Map of the Route of the Yellowstone Expedition,” based on his 1870 visit to the area as commander of the military escort of the Washburn- Langford-Doane expedition, showed “Bottler’s Ranche” on the Yellowstone River near the mouth of Emigrant Creek. From this base of operations, the Bottlers conducted hide- and meat- hunting operations in the region, killing thousands of animals in the present park area. Courtesy of the National Archives.
including the possible combining of wolf and coyote hides of 380,000 wolves in the western United States and Mexico in some totals. prior to the near destruction of both species south of Canada Recently, scientific methods unavailable to research- in the late 1800s and early 1900s. ers only a generation ago have been applied to the ques- Considering the recovery of Yellowstone wolves in this tion of the western wolf population. Leonard et al. (2005) context, it is worth noting that Leonard et al. concluded that extracted mitochondrial DNA from 34 wolf specimens in “modern wolves are a depauperate subset of the historic popula- the collection of the National Museum of Natural History tion,” whose large size “provides a striking contrast to restora- (Smithsonian Institution) in Washington, D.C., that were tion goals in the western US,” which are limited to a total of killed before 1916, some as early as 1856. Most of these wolves 400 wolves for C. l. nubilis and C. l. baileyi. Taking a posi- were killed within the historical ranges of Canis lupus nubilus, tion that makes current wolf recovery goals in the northern the wolf of the Great Plains and Interior West, and C. l. bai American Rockies seem modest in the extreme, Leonard et al. leyi, the Mexican wolf, but a few were from as far north and recommended a much more ambitious restoration effort in east as Labrador. Based on the time required for the gray wolf the West that “would restore wolves to past population sizes to achieve the degree of genetic diversity displayed by this and enable them to significantly influence the dynamics of the set of specimens, Leonard et al. arrived at a “rough estimate” Rocky Mountain ecosystem.”
19(1) • 2011 Yellowstone Science 25 US Ge o l ogic al S u r v e y P hotog ra phic
Lib rar y WHJ-1257
The entire roster of the combined parties of the Hayden Survey, photographed by W. H. Jackson about August 15, 1872, in the Lower Geyser Basin. An enlargement of the lower right corner of the photograph (inset, right) provides a look at #21, Joseph Savage, the author of a historic and previously unknown account of wolf-hide trade in the Yellowstone area.
A Greater Yellowstone Wolf Harvest best of terms with park man- The slaughter of YNP wildlife in the 1870s and early 1880s agers; when Superintendent has been documented repeatedly (Hampton 1971; Reiger Norris’s operating funds were 1975; Haines 1977; Schullery and Whittlesey 1992; Schullery lost in a local bank failure in 1997). Among the published contemporary reports was 1878, the Bottlers provided Superintendent Norris’s annual report for 1877, his first year him with money to meet his in office. Norris estimated that an annual average of 1,000 payroll (Haines 1977). elk, as well as “hundreds if not thousands” of each of the Among the evidence of the Bottlers’ hunting is an other species of large ungulates, had been killed and taken account by Norris of an 1875 visit to the Yellowstone valley, from the park since 1870 (Norris 1878). when he was guided by “my old comrade Frederick Bottler” Though hunting was legal in the park until 1883, (Norris no date). According to Norris, “the Bottler Bros. the act creating the park in 1872 had placed limits on it. assure me that they alone packed over 2,000 elk skins from Visitors were allowed to hunt for their own subsistence and the forks of the Yellowstone [the lower Lamar Valley area], for sport but not for profit. The act directed the Secretary besides vast numbers of other pelts, and other hunters at of the Interior to provide against the wanton destruction of least as many more, in the spring of 1875.” William Henry the fish and game found within said park, and against their Jackson’s 1871 photograph of the Bottler Ranch provides rare capture or destruction for the purposes of merchandise or visual evidence of their work, showing a large shed overflow- profit (Schullery 1996). Until the arrival of the US Cavalry ing with hides (and possibly unbutchered carcasses). An 1872 to protect the park in 1886, little or no notice was paid to this photograph of Bottler with Hayden Survey members and five prohibition against commercial hunting. elk freshly shot near the Grand Canyon of the Yellowstone is The slaughter of thousands of the park’s large mammals additional evidence of Bottler’s hunting success. for their hides and for the regional meat trade was regarded The magnitude of the killing was rarely reported as spe- as a great scandal among conservationists in the 1870s and cifically as the number of elk hides that Bottler described to early 1880s, but it was carried on with remarkable aplomb Norris in 1875. An important exception to the mostly vague by local entrepreneurs. The Bottler family, who established accounts has just come to light, identified by Whittlesey in their ranch in today’s Paradise Valley in the late 1860s, were a series of letters in the Western Home Journal, a newspa- hard-working farmers and ranchers in addition to profes- per published in Lawrence, Kansas, in 1872. As many small sional hunters. In the years just before and after 1872, the local newspapers have been scanned and made available Bottlers routinely hosted parties on their way to and from online in the last decade, hundreds of previously uncol- the park, often serving as guides to important groups. They lected Yellowstone items from the late nineteenth and early were among the best known and most trusted guides for early twentieth centuries have been added to the files of the park’s survey parties, and were apparently forthright in describing Research Library. This item is one of those. their hide-hunting activities. Despite their energetic hide- A series of five letters by Joseph Savage, a member of hunting in and near the park, the Bottlers were often on the Ferdinand Hayden’s US Geological Survey, was published
26 Yellowstone Science 19(1) • 2011 The Savage account of the Bottler Bottler was placed well south of the main commerce centers collection of wolf “peltry” is thus of the Montana Territory hide trade. Hides were routinely gathered in Bozeman and along the Yellowstone River for uniquely important among transport down that river, or in Fort Benton for transport known sources for its ability to down the Missouri River. We infer from this that the people with whom Bottler traded for wolf hides were most likely in clarify our understanding of the his neighborhood, because if they were farther north they distribution and abundance of could have more conveniently dealt with the big shippers in Bozeman or Fort Benton. For this reason, we assume that Yellowstone-area wolves. most or all of the skins Bottler obtained came from animals in Paradise Valley and adjoining areas. As for the wolves that the Bottlers killed, it seems most in three issues of the Western Home Journal. Like some other likely that these were also from the region surrounding the early official travelers who published less formal accounts of ranch where they lived. Their hunting grounds during 1871 their experiences and adventures, Savage wrote letters to what and 1872 may or may not have included the present park seems to have been his home-town newspaper. In his first let- area, but would almost certainly have been confined to the ter, written from “Fort Ellis, Montana Territory, Camp of the northern portion of the Greater Yellowstone ecosystem. US Geological Survey” on July 18, 1872, and published on The Savage account of the Bottler collection of wolf August 1, Savage mentions the “younger Bottler,” Frederick, “peltry” is thus uniquely important among known sources and the “elder Bottler,” Phillip. “The younger Bottler brother for its ability to clarify our understanding of the distribu- is employed as a hunter for our party, and the ‘aching void’ tion and abundance of Yellowstone-area wolves. That the within us, which even the mountains cannot fill, is now sup- Bottlers could accumulate several hundred wolf hides in the plied with tender, juicy elk steak, the fruit of two days’ hunt- space of one year reinforces the reliability of reports of preda- ing.” Only Frederick was listed on the official roll of members tor and prey abundance left by many other early observers. of the 1872 Hayden Survey (Haines 1977), but apparently However, the question necessarily arises whether these 550 Phillip was at Fort Ellis at the time and told Savage about pelts all came from wolves or if some were from coyotes. his hide-hunting: In an assessment of the Savage account, wolf historian Rick McIntyre concluded: The elder Bottler gave us the number of his peltries, killed and traded for during the past year, which he sold There is the issue that many people during that period did in the New York market. It was this: three hundred and not properly distinguish between the two species. I found one elk skins, two hundred and fifty deer and antelope in some of the historic documents I looked at that the skins, and five hundred and fifty-five wolf skins; bringing words coyote and wolf were used interchangeably. In some in market two thousand nine hundred and ninety-one cases, the same animal would be called a wolf in the first dollars and thirty-two cents—freight being three hundred and ninety-seven dollars. (Savage 1872) N PS Interpretations of the Savage Account
As with so many tidbits of wildlife information gleaned from early accounts, this paragraph is both revealing and tantaliz- ing. A few generalizations seem warranted. “The past year” mentioned by Savage as the time during which Bottler gath- ered the 555 wolf hides presumably refers to the period from July 1871 to July 1872. Considering that by 1872 the Bottlers were obviously very active in killing elk and other large mam- mals, they had regular opportunities to poison carcasses and collect the resulting dead wolves. Phillip Bottler apparently did not kill all these animals himself. Savage said that Bottler “killed and traded for” the “The Successful Hunter” by W. H. Jackson shows Frederick hides. The range of Bottler’s killing and trading is therefore of Bottler and other Hayden Survey party members on August interest. Operating from the present Emigrant area along the 6 or 7, 1872, near the Grand Canyon of the Yellowstone Yellowstone River approximately 30 miles north of the park, where Bottler has just killed five bull elk.
19(1) • 2011 Yellowstone Science 27 N
PS Associated University Press, Boulder, CO. part of a sentence and a coyote later in Hampton, B. 1997. The Great American Wolf. the same sentence. Henry Holt, New York. Hampton, H.D. 1971. How the US Cavalry Saved But the Bottlers would have been Our National Parks. Indiana University Press, more experienced in buying and sell Bloomington, Indiana. ing hides than most others in that Leonard, J.A., C. Vila, and R.K. Wayne. 2005. Legacy lost: genetic variability and popula- period and their income depended on tion size of extirpated US grey wolves (Canis being somewhat accurate in tabulat lupus). Molecular Ecology 14:9–17. ing hide types and numbers. As the McIntyre, R., ed. 1995. War Against the Wolf: America’s Campaign to Exterminate the Wolf. elder Bottler told Savage that they Voyageur Press, Stillwater, Minnesota. sold 555 wolf skins, I would have to Norris, P.W. 1878. Report upon the Yellowstone regard that statement as being reason National Park to the Secretary of the Interior. Paul Schullery is a former editor of US Government Printing Office, Washington, ably accurate. The buyers in New York Yellowstone Science and the author or edi- D.C.. would certainly have given them a tor of many books about the American Norris, P.W. 1881. Annual report of the Superin- West, including ten about Yellowstone. tendent of the Yellowstone National Park to the lower price if there were a significant Secretary of the Interior for the year 1880. US For his work as a writer and historian, he number of smaller sized coyote pelts in Government Printing Office, Washington, received an honorary doctorate of letters the shipment. A wolf pelt would have D.C.. from Montana State University and the Norris, P.W. no date. Meanderings of a moun- brought a much higher price than a Wallace Stegner Award from the University taineer, or, The journals and musings (or coyote hide (McIntyre, personal com- of Colorado Center of the American West. storys) of a rambler over prairie (or moun- munication, January 13, 2011). He is currently the Scholar-in-Residence tain) and plain. Ms. prepared from newspaper at Montana State University Library in clippings (1870–75) and handwritten addi- Bozeman, Montana. tions, annotated by the author about 1885. We agree with McIntyre’s assess- Huntington Library, San Marino, Calif.
N Reiger, J. 1975. American Sportsmen and the
ment. Established and locally respected PS Origins of Conservation. Winchester Press, shippers like the Bottlers were not New York. in a position to perpetrate a whole- Savage, J. 1872. From the Yellowstone Expedi- sale fraud on dealers downstream. tion. Western Home Journal (Lawrence, Kansas), August 1. However, as is always the case when Schullery, P. 1997. Searching for Yellowstone: Ecol- evaluating anecdotal material of this ogy and Wonder in the Last Wilderness. Hough- ton Mifflin Company, Boston, Massachusetts. sort, we are left with other unresolved Schullery, P, editor. 1996. The Yellowstone Wolf: questions. From how far afield were A Guide and Sourcebook. High Plains Publishing the hides gathered by the Bottlers? Company, Worland, Wyoming. Schullery, P., and L. Whittlesey. 1992. The Were any, or many, of the wolves killed Documentary Record of Wolves and Related in the present park area? For how Wildlife Species in the Yellowstone National many years did the Bottlers sustain Park Area Prior to 1882.” In J.D. Varley, J.D., and W.G. Brewster, editors. Wolves their trade in wolf hides on this scale? for Yellowstone? A Report to the United States Whatever answers we may eventually Congress, Volume IV, Research and Analysis. Lee Whittlesey is the historian for find to these and other questions, the Pages 1.3–1-174. National Park Service, Yel- Yellowstone National Park. He is the lowstone National Park. Savage account does demonstrate what author, co-author, or editor of ten books Schullery, P., and L. Whittlesey. 1995. Summary many other accounts have shown, that and more than 30 journal articles related of the Documentary Record of Wolves and when Yellowstone National Park was to Yellowstone and the northern Rockies Other Wildlife Species in the Yellowstone National Park Area Prior to 1882. In Ecology established, the area supported large including the recent Storytelling in Yellowstone. Lee holds a master’s degree in history and Conservation of Wolves in a Changing World. L.N. Carbyn, S.H. Fritts, and D.R. numbers of wolves as part of a robust from Montana State University and a JD in community of wildlife species. Seip, editors, 63–76. Canadian Circumpolar Law from the University of Oklahoma. In Institute, University of Alberta, Edmonton, 2001, Idaho State University awarded him Alberta. an Honorary Doctorate of Science and Schullery, P., L. Whittlesey, S. Bone, G. Plumb, Acknowledgements Humane Letters. and A. Rodman, forthcoming. Greater Yel- In Yellowstone National Park we would lowstone Mammals, 1796–1881: An Analysis like to thank Rick McIntyre, Doug Smith, of Historical Observations. Literature Cited Seton, E.T. 1953. Lives of game animals. Volume Janine Waller, and P.J. White for helpful read- Burroughs, R.D., editor, 1961. The Natural 1. Part 1, Cats, Wolves, and Foxes. Charles T. ings and suggestions for important sources History of the Lewis and Clark Expedition. Branford Company, Boston, Massachusetts. of information. For assistance with the Michigan State University Press, East Lansing, Walter, D. 1986. Wolf Wars. Montana Magazine Doane 1870 map, we thank Ann Rodman Michigan. 75, January–February 1986, 21–26. of Yellowstone, Alethea Steingisser of the Curnow, E.E. 1969. The History of the Eradica- Zmyj, P.M. 1996. ‘A Fight to the Finish’: The University of Oregon, and Head Special tion of the Wolf in Montana. M.A. Thesis, Extermination of the Gray Wolf in Wyoming, University of Montana, Missoula, Montana. 1890–1930. Montana the Magazine of Western Collections Librarian Kim Allen Scott at the Haines, A. 1977. The Yellowstone Story. Colorado Montana State University Library, Bozeman. History 46(1), Spring 1996, 14–25.
28 Yellowstone Science 19(1) • 2011 From the Archives Y ELL -2791-3
In this photo from 1964, a group of bison are being hazed into the Crystal Creek pen by helicopter.
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