ECOLOGICAL DYNAMICS ON YELLOWSTONE’S NORTHERN RANGE

Committee on Ungulate Management in Yellowstone National Park

Board on Environmental Studies and Toxicology

Division on Earth and Life Studies

National Research Council

NATIONAL ACADEMY PRESS
Washington, D.C.



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Ecological Dynamics on Yellowstone’s Northern Range ECOLOGICAL DYNAMICS ON YELLOWSTONE’S NORTHERN RANGE Committee on Ungulate Management in Yellowstone National Park Board on Environmental Studies and Toxicology Division on Earth and Life Studies National Research Council NATIONAL ACADEMY PRESS Washington, D.C.

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Ecological Dynamics on Yellowstone’s Northern Range NATIONAL ACADEMY PRESS 2101 Constitution Ave., N.W. Washington, D.C. 20418 NOTICE: The project that is the subject of this report was approved by the Governing Board of the National Research Council, whose members are drawn from the councils of the National Academy of Sciences, the National Academy of Engineering, and the Institute of Medicine. The members of the committee responsible for the report were chosen for their special competences and with regard for appropriate balance. This project was supported by Cooperative Agreement 1443CX2605-98-005, between the National Academy of Sciences and the U.S. Department of Interior National Park Service. Any opinions, findings, conclusions, or recommendations expressed in this publication are those of the author(s) and do not necessarily reflect the view of the organizations or agencies that provided support for this project. Library of Congress Control Number 2002108821 International Standard Book Number 0-309-083451 Additional copies of this report are available from: National Academy Press 2101 Constitution Ave., NW Box 285 Washington, DC 20055 800–624–6242 202–334–3313 (in the Washington metropolitan area) http://www.nap.edu Copyright 2002 by the National Academy of Sciences. All rights reserved. Printed in the United States of America

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Ecological Dynamics on Yellowstone’s Northern Range THE NATIONAL ACADEMIES National Academy of Sciences National Academy of Engineering Institute of Medicine National Research Council The National Academy of Sciences is a private, nonprofit, self-perpetuating society of distinguished scholars engaged in scientific and engineering research, dedicated to the furtherance of science and technology and to their use for the general welfare. Upon the authority of the charter granted to it by the Congress in 1863, the Academy has a mandate that requires it to advise the federal government on scientific and technical matters. Dr. Bruce M.Alberts is president of the National Academy of Sciences. The National Academy of Engineering was established in 1964, under the charter of the National Academy of Sciences, as a parallel organization of outstanding engineers. It is autonomous in its administration and in the selection of its members, sharing with the National Academy of Sciences the responsibility for advising the federal government. The National Academy of Engineering also sponsors engineering programs aimed at meeting national needs, encourages education and research, and recognizes the superior achievements of engineers. Dr. Wm.A.Wulf is president of the National Academy of Engineering. The Institute of Medicine was established in 1970 by the National Academy of Sciences to secure the services of eminent members of appropriate professions in the examination of policy matters pertaining to the health of the public. The Institute acts under the responsibility given to the National Academy of Sciences by its congressional charter to be an adviser to the federal government and, upon its own initiative, to identify issues of medical care, research, and education. Dr. Kenneth I.Shine is president of the Institute of Medicine. The National Research Council was organized by the National Academy of Sciences in 1916 to associate the broad community of science and technology with the Academy’s purposes of furthering knowledge and advising the federal government. Functioning in accordance with general policies determined by the Academy, the Council has become the principal operating agency of both the National Academy of Sciences and the National Academy of Engineering in providing services to the government, the public, and the scientific and engineering communities. The Council is administered jointly by both Academies and the Institute of Medicine. Dr. Bruce M.Alberts and Dr. Wm.A.Wulf are chairman and vice chairman, respectively, of the National Research Council.

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Ecological Dynamics on Yellowstone’s Northern Range COMMITTEE ON UNGULATE MANAGEMENT IN YELLOWSTONE NATIONAL PARK Members DAVID R.KLEIN (Chair), University of Alaska Fairbanks, Fairbanks DALE R.MCCULLOUGH (Vice Chair), University of California, Berkeley BARBARA ALLEN-DIAZ, University of California, Berkeley NORMAN CHEVILLE, Iowa State University, Ames RUSSELL W.GRAHAM, Denver Museum of Natural History, Denver, Colorado JOHN E.GROSS, Commonwealth Scientific and Industrial Research Organization, Australia JAMES MACMAHON, Utah State University, Logan NANCY E.MATHEWS, University of Wisconsin, Madison DUNCAN T.PATTEN, Montana State University, Bozeman KATHERINE RALLS, Smithsonian Institution, Washington, D.C. MONICA G.TURNER, University of Wisconsin, Madison ELIZABETH S.WILLIAMS, University of Wyoming, Laramie Project Staff DAVID J.POLICANSKY, Project Director LEE PAULSON, Staff Officer MARGARET WALSH, Postdoctoral Research Associate CAY BUTLER, Editor KELLY CLARK, Editorial Assistant MIRSADA KARALIC-LONCAREVIC, Research Assistant RAMYA CHARI, Project Assistant KATHY IVERSON, Senior Project Assistant Sponsor U.S.DEPARTMENT OF INTERIOR

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Ecological Dynamics on Yellowstone’s Northern Range BOARD ON ENVIRONMENTAL STUDIES AND TOXICOLOGY1 Members GORDON ORIANS (Chair), University of Washington, Seattle JOHN DOULL (Vice Chair), University of Kansas Medical Center, Kansas City DAVID ALLEN, University of Texas, Austin INGRID C.BURKE, Colorado State University, Fort Collins THOMAS BURKE, Johns Hopkins University, Baltimore, Maryland WILLIAM L.CHAMEIDES, Georgia Institute of Technology, Atlanta CHRISTOPHER B.FIELD, Carnegie Institute of Washington, Stanford, California DANIEL S.GREENBAUM, Health Effects Institute, Cambridge, Massachusetts BRUCE D.HAMMOCK, University of California, Davis ROGENE HENDERSON, Lovelace Respiratory Research Institute, Albuquerque, New Mexico CAROL HENRY, American Chemistry Council, Arlington, Virginia ROBERT HUGGETT, Michigan State University, East Lansing JAMES H.JOHNSON, Howard University, Washington, D.C. JAMES F.KITCHELL, University of Wisconsin, Madison DANIEL KREWSKI, University of Ottawa, Ottawa, Ontario JAMES A.MACMAHON, Utah State University, Logan WILLEM F.PASSCHIER, Health Council of the Netherlands, The Hague ANN POWERS, Pace University School of Law, White Plains, New York LOUISE M.RYAN, Harvard University, Boston, Massachusetts KIRK SMITH, University of California, Berkeley LISA SPEER, Natural Resources Defense Council, New York, New York Senior Staff JAMES J.REISA, Director DAVID J.POLICANSKY, Associate Director and Senior Program Director for Applied Ecology RAYMOND A.WASSEL, Senior Program Director for Environmental Sciences and Engineering KULBIR BAKSHI, Program Director for the Committee on Toxicology ROBERTA M.WEDGE, Program Director for Risk Analysis K.JOHN HOLMES, Senior Staff Officer SUSAN N.J.MARTEL, Senior Staff Officer SUZANNE VAN DRUNICK, Senior Staff Officer RUTH E.CROSSGROVE, Managing Editor 1   This study was planned, overseen, and supported by the Board on Environmental Studies and Toxicology.

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Ecological Dynamics on Yellowstone’s Northern Range OTHER REPORTS OF THE BOARD ON ENVIRONMENTAL STUDIES AND TOXICOLOGY The Airliner Cabin Environment and Health of Passengers and Crew (2002) Arsenic in Drinking Water: 2001 Update (2001) Evaluating Vehicle Emissions Inspection and Maintenance Programs (2001) Compensating for Wetland Losses Under the Clean Water Act (2001) A Risk-Management Strategy for PCB-Contaminated Sediments (2001) Toxicological Effects of Methylmercury (2000) Strengthening Science at the U.S. Environmental Protection Agency: Research-Management and Peer-Review Practices (2000) Scientific Frontiers in Developmental Toxicology and Risk Assessment (2000) Copper in Drinking Water (2000) Ecological Indicators for the Nation (2000) Waste Incineration and Public Health (1999) Hormonally Active Agents in the Environment (1999) Research Priorities for Airborne Paniculate Matter (3 reports, 1998–2001) Ozone-Forming Potential of Reformulated Gasoline (1999) Risk-Based Waste Classification in California (1999) Arsenic in Drinking Water (1999) Brucellosis in the Greater Yellowstone Area (1998) The National Research Council’s Committee on Toxicology: The First 50 Years (1997) Carcinogens and Anticarcinogens in the Human Diet (1996) Upstream: Salmon and Society in the Pacific Northwest (1996) Science and the Endangered Species Act (1995) Wetlands: Characteristics and Boundaries (1995) Biologic Markers (5 reports, 1989–1995) Review of EPA’s Environmental Monitoring and Assessment Program (3 reports, 1994–1995) Science and Judgment in Risk Assessment (1994) Pesticides in the Diets of Infants and Children (1993) Protecting Visibility in National Parks and Wilderness Areas (1993) Dolphins and the Tuna Industry (1992) Science and the National Parks (1992) Assessment of the U.S. Outer Continental Shelf Environmental Studies Program, Volumes I–IV (1991–1993) Human Exposure Assessment for Airborne Pollutants (1991) Rethinking the Ozone Problem in Urban and Regional Air Pollution (1991) Decline of the Sea Turtles (1990) Copies of these reports may be ordered from the National Academy Press (800) 624–6242 or (202) 334–3313 www.nap.edu

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Ecological Dynamics on Yellowstone’s Northern Range Acknowledgment of Review Participants This report has been reviewed in draft form by individuals chosen for their diverse perspectives and technical expertise, in accordance with procedures approved by the NRC’s Report Review Committee. The purpose of this independent review is to provide candid and critical comments that will assist the institution in making its published report as sound as possible and to ensure that the report meets institutional standards for objectivity, evidence, and responsiveness to the study charge. The review comments and draft manuscript remain confidential to protect the integrity of the deliberative process. We wish to thank the following individuals for their review of this report: Donald Barry, The Wilderness Society Peter A.Bisson, U.S. Forest Service Mark S.Borsity of Alberta Ingrid Burke, Colorado State University Charles C.Capen, Ohio State University Michael B.Coughenour, Colorado State University Amy E.Hessl, West Virginia University Douglas B.Houston, National Park Service (retired) Peggy Johnson, Pennsylvania State University

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Ecological Dynamics on Yellowstone’s Northern Range Walter Klippel, University of Tennessee Frederic Wagner, Utah State University James E.Womack, Texas A&M University Although the reviewers listed above have provided many constructive comments and suggestions, they were not asked to endorse the conclusions or recommendations nor did they see the final draft of the report before its release. The review of this report was overseen by Simon Levin, Princeton University, and Ellis Cowling, North Carolina State University. Appointed by the National Research Council, they were responsible for making certain that an independent examination of this report was carried out in accordance with institutional procedures and that all review comments were carefully considered. Responsibility for the final content of this report rests entirely with the authoring committee and the institution.

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Ecological Dynamics on Yellowstone’s Northern Range Preface National Park policy for management of ungulates within Yellowstone National Park (YNP) has undergone major changes since Yellowstone, the world’s first national park, was established in 1872. These changes, from little emphasis on wildlife at the time of establishment to a major focus on wildlife today, have accompanied an evolving interest in wildlife by the visiting public together with increased understanding of the ecosystem relationships of wildlife by the National Park Service. The ecological relationships of the elk, bison, and other ungulates that inhabit the northern portion of YNP are, however, extremely complex. Acquiring adequate knowledge of this complexity as a basis for park management policy has been complicated by both the dynamic changes that have characterized the system and the recognition that the natural boundaries of this ecosystem extend well outside of YNP, largely to the north into Montana. As a consequence, any management activities that may affect the ungulates within YNP, such as previous reduction hunts and the more recent reintroduction of wolves to the ecosystem, are not independent of land use and associated human activities in the northern range area outside of the park. The national park concept originated in the United States with the National Park Service’s mandate for management of YNP has been to assure protection of the geological, landscape, and biological features for which it was established and to provide opportunities for the public to visit and appreciate its unique natural values. Interests and expectations of visitors to YNP regard-

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Ecological Dynamics on Yellowstone’s Northern Range ing wildlife, although changing over time, have continued to influence wildlife management policy within the park. An understanding of the ecological relationships of wildlife within the park is also an important component of national park management. A major challenge for park management is to meet expectations of the visiting public regarding wildlife; with the cumulative increase in understanding of the ecology of park wildlife. Within this context of ecosystem change and complexity, changing human perspectives and interests in wildlife, and markedly differing land designations and uses inside and outside of YNP, it is not surprising that controversy arose over National Park Service policy toward ungulates within the park. The National Research Council’s Committee on Ungulate Management in Yellowstone National Park, charged to review information of the population ecology about ungulates on the northern range of the Greater Yellowstone ecosystem and associated practices for their management, was cognizant of the diversity of perspectives, concerns, and opinions that the public has expressed about management policy for ungulates in YNP. The committee appreciated the opportunity to hear from a wide range of people interested in the issues the committee was asked to address during open forums in Gardiner, Montana, and in Mammoth, Wyoming. The oral and written submissions, reports, and publications provided to the committee by members of the public; the National Park Service; the U.S. Forest Service; Montana Fish, Wildlife and Parks; the Biological Resources Division of the U.S. Geological Survey; and other agency representatives; were particularly valuable in our efforts to comprehensively review all relevant information. We thank the following people who made presentations to the committee: John Dennis of the Washington office of NPS; Michael Finley, John Varley, Paul Schullery, Ann Johnson, and Wayne Brewster of YNP; Steve Torbit of the National Wildlife Federation; rancher Brian Severin; Mike Harris of Senator Conrad Burns’s office; Frederic H. Wagner of Utah State University; Bob Beschta of Oregon State University; Richard Keigley, Peter Gogan, and Kim Keating of the U.S. Geological Survey; Tom Lemke and Kurt Alt of Montana Fish, Wildlife, and Parks; Clifford Montagne of Montana State University; Timothy Clark of Yale University; Rex Gates of Brigham Young University; and members of the Northern Yellowstone Cooperative Wildlife Working Group. We hope the efforts of the committee, culminating in this report, will enhance understanding of the complexity of the ecological relationships of the ungulates of the Yellowstone northern range and will strengthen the scientific

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Ecological Dynamics on Yellowstone’s Northern Range basis for effectively managing them within the context of the Greater Yellowstone ecosystem. All members of the committee were generous in their commitment of time through participation in the meetings, discussions, and writing of this consensus report. We appreciate the assistance provided by National Research Council staff Lee Paulson, David Policansky, Chris Elfring, Margaret Walsh, Kathy Iverson, Stephanie Parker, Mirsada Karalic-Loncarevic, Kelly Clark, Ramya Chari, and Jennifer Saunders, who supported our efforts to meet, discuss, and prepare the final report. Of particular importance to the completion of the report are the efforts of Lee Paulson, during the initial stage of committee meetings and preliminary writing; Gordon Orians, who as liaison from the Board on Environmental Studies and Toxicology, provided oversight, guidance, and critical review; and David Policansky, who undertook the difficult and final editorial task of melding and integrating the written contributions of committee members. David R.Klein, Ph.D. Chair, Committee on Ungulate Management in Yellowstone National Park

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Ecological Dynamics on Yellowstone’s Northern Range Contents     SUMMARY   1 1   INTRODUCTION   15     Background and Charge to the Committee,   15     Natural Regulation,   20     Historical Context,   24     Climate Versus Elk As Causality of Changes in Aspen and Willow,   30     Organization of This Report,   31 2   HISTORICAL PERSPECTIVE: YELLOWSTONE’S CHANGING ENVIRONMENTS   32     Scales of Change,   32     Paleobiology of the Greater Yellowstone Area,   33     Ungulate Use Over Time As Estimates of Population Densities,   35     Physical Changes from Prehistory to the Present,   35 3   PRESENT CONDITIONS: VEGETATION   43     Upland Shrublands and Grasslands of the Northern Range,   43     Forest Types Associated with the Northern Range,   50     Aspen Communities of the Northern Range,   55     Riparian Communities of the Northern Range,   67

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Ecological Dynamics on Yellowstone’s Northern Range     Wetland Vegetation of the Northern Range,   79 4   PRESENT CONDITIONS: ANIMALS   83     Population Dynamics of Northern Range Ungulates,   83     Major Predators in the Yellowstone Ecosystem,   109 5   CONCLUSIONS AND RECOMMENDATIONS   120     Nature Is Dynamic,   120     Management for Ecosystem State or Ecosystem Process?   122     Large-Scale Interactions and Patterns,   125     Weather, Wolves, and Aspen in Yellowstone,   126     Indicators of Unacceptable Change,   127     Understanding the Consequences of Alternative Management Approaches,   128     Conclusions,   132     Recommendations,   134     Epilogue,   137     REFERENCES   138 APPENDIX A:   UNDERSTANDING THE PAST   168 APPENDIX B:   BIOGRAPHICAL INFORMATION ON THE COMMITTEE ON UNGULATE MANAGEMENT IN YELLOWSTONE NATIONAL PARK   177

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Ecological Dynamics on Yellowstone’s Northern Range Ecological Dynamics on Yellowstone’s Northern Range

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