Greater Sage-Grouse: Ecology and Conservation of a Landscape Species and Its Habitats 9780520948686

Table of contents :
CONTENTS
CONTRIBUTORS
PREFACE
FOREWORD
Greater Sage-Grouse and Sagebrush: An Introduction to the Landscape
Part I. Management and Conservation Status
CHAPTER ONE. Historical Development, Principal Federal Legislation, and Current Management of Sagebrush Habitats: IMPLICATIONS FOR CONSERVATION
CHAPTER TWO. The Legal Status of Greater Sage-Grouse: ORGANIZATIONAL STRUCTURE OF PLANNING EFFORTS
PART II. Ecology of Greater Sage-Grouse
CHAPTER THREE. Characteristics and Dynamics of Greater Sage-Grouse Populations
CHAPTER FOUR. Characteristics of Greater Sage-Grouse Habitats: A LANDSCAPE SPECIES AT MICRO- AND MACROSCALES
CHAPTER FIVE. Molecular Insights into the Biology of Greater Sage-Grouse
CHAPTER SIX. Predation on Greater Sage-Grouse: FACTS, PROCESS, AND EFFECTS
CHAPTER SEVEN. Harvest Management for Greater Sage-Grouse: A CHANGING PARADIGM FOR GAME BIRD MANAGEMENT
CHAPTER EIGHT. Parasites and Infectious Diseases of Greater Sage-Grouse
CHAPTER NINE. West Nile Virus Ecology in Sagebrush Habitat and Impacts on Greater Sage-Grouse Populations
Part III. Ecology of Sagebrush
CHAPTER TEN. Characteristics of Sagebrush Habitats and Limitations to Long-Term Conservation
CHAPTER ELEVEN. Pre–Euro-American and Recent Fire in Sagebrush Ecosystems
CHAPTER TWELVE. Ecological Influence and Pathways of Land Use in Sagebrush
CHAPTER THIRTEEN. Influences of the Human Footprint on Sagebrush Landscape Patterns: IMPLICATIONS FOR SAGE-GROUSE CONSERVATION
CHAPTER FOURTEEN. Influences of Free-Roaming Equids on Sagebrush Ecosystems, with a Focus on Greater Sage-Grouse
Part IV. Population Trends and Habitat Relationships
CHAPTER FIFTEEN. Greater Sage-Grouse Population Dynamics and Probability of Persistence
CHAPTER SIXTEEN. Connecting Pattern and Process in Greater Sage-Grouse Populations and Sagebrush Landscapes
CHAPTER SEVENTEEN. Influences of Environmental and Anthropogenic Features on Greater Sage-Grouse Populations, 1997–2007
CHAPTER EIGHTEEN. Factors Associated with Extirpation of Sage-Grouse
Part V. Conservation and Management
CHAPTER NINETEEN. Greater Sage-Grouse as an Umbrella Species for Shrubland Passerine Birds: A MULTISCALE ASSESSMENT
CHAPTER TWENTY. Energy Development and Greater Sage-Grouse
CHAPTER TWENTY-ONE. Energy Development and Conservation Tradeoffs: SYSTEMATIC PLANNING FOR GREATER SAGE-GROUSE IN THEIR EASTERN RANGE
CHAPTER TWENTY-TWO. Response of Greater Sage-Grouse to the Conservation Reserve Program in Washington State
CHAPTER TWENTY-THREE. Restoring and Rehabilitating Sagebrush Habitats
CHAPTER TWENTY-FOUR. Conservation of Greater Sage-Grouse: A SYNTHESIS OF CURRENT TRENDS AND FUTURE MANAGEMENT
LITERATURE CITED
INDEX
Series Titles

Citation preview

GREATER SAGE-GROUSE

STUDIES IN AVIAN BIOLOGY A Publication of the Cooper Ornithological Society HTTP://WWW.UCPRESS.EDU/GO/SAB

Studies in Avian Biology is a series of works by the Cooper Ornithological Society since 1978. Volumes in the series address current topics in ornithology and can be organized as monographs or multi-authored collections of chapters. Authors are invited to contact the series editor to discuss project proposals and guidelines for preparation of manuscripts.

Series Editors Carl D. Marti Brett K. Sandercock, Kansas State University Volume Technical Editor Clait E. Braun, Grouse, Inc. Editorial Board Frank R. Moore, University of Southern Mississippi John T. Rotenberry, University of California at Riverside Steven R. Beissinger, University of California at Berkeley Katie M. Dugger, Oregon State University Amanda D. Rodewald, Ohio State University Jeffrey F. Kelly, University of Oklahoma Science Publisher Charles R. Crumly, University of California Press See complete series list on page 645.

GREATER SAGE-GROUSE Ecology and Conservation of a Landscape Species and Its Habitats Steven T. Knick and John W. Connelly, Editors Studies in Avian Biology No. 38

A PUBLICATION OF THE COOPER ORNITHOLOGICAL SOCIETY

University of California Press Berkeley

Los Angeles

London

University of California Press, one of the most distinguished university presses in the United States, enriches lives around the world by advancing scholarship in the humanities, social sciences, and natural sciences. Its activities are supported by the UC Press Foundation and by philanthropic contributions from individuals and institutions. For more information, visit www.ucpress.edu. Studies in Avian Biology, No. 38 For digital edition of this work, please see the UC Press website. University of California Press Berkeley and Los Angeles, California University of California Press, Ltd. London, England © 2011 by the Cooper Ornithological Society Library of Congress Cataloging-in-Publication Data Greater sage-grouse : ecology and conservation of a landscape species and its habitats / Steven T. Knick and John W. Connelly, editors. p. cm. — (Studies in avian biology ; no. 38) “A Publication of the Cooper Ornithological Society.” Includes bibliographical references and index. ISBN 978-0-520-26711-4 (cloth : alk. paper) 1. Sage grouse—Ecology. 2. Sage grouse—Habitat—Conservation. 3. Sagebrush—Ecology. I. Knick, Steven T. II. Connelly, John W. (John William), 1952– QL696.G27E26 2011 639.9'78636—dc22

2010052102 19 18 17 16 15 14 13 12 11 10 9 8 7 6 5 4 3 2 1

The paper used in this publication meets the minimum requirements of ANSI/NISO Z39.48-1992 (R 1997)(Permanence of Paper). Cover image: Greater Sage-Grouse. Photo by Terry R. Steele.

PERMISSION TO COPY The Cooper Ornithological Society hereby grants permission to copy chapters (in whole or in part) appearing in Studies in Avian Biology for personal use, or educational use within one’s home institution, without payment, provided that the copied material bears the statement “©2011 The Cooper Ornithological Society” and the full citation, including names of all authors. Authors may post copies of their chapters on their personal or institutional website, except that whole issues of Studies in Avian Biology may not be posted on websites. Any use not specifically granted here, and any use of Studies in Avian Biology articles or portions thereof for advertising, republication, or commercial uses, requires prior consent from the series editor.

CONTENTS

Contributors / vii Preface / xi Steven T. Knick and John W. Connelly Foreword / xiii John C. Freemuth Greater Sage-Grouse and Sagebrush: An Introduction to the Landscape / 1 Steven T. Knick and John W. Connelly

Part I • Management and Conservation Status 1 • HISTORICAL DEVELOPMENT, PRINCIPAL FEDERAL LEGISLATION, AND CURRENT MANAGEMENT OF SAGEBRUSH HABITATS: IMPLICATIONS FOR CONSERVATION / 13

Steven T. Knick 2 • THE LEGAL STATUS OF GREATER SAGE-GROUSE: ORGANIZATIONAL STRUCTURE OF PLANNING EFFORTS / 33

San J. Stiver

Part II • Ecology of Greater Sage-Grouse 3 • CHARACTERISTICS AND DYNAMICS OF GREATER SAGE-GROUSE POPULATIONS / 53

John W. Connelly, Christian A. Hagen, and Michael A. Schroeder 4 • CHARACTERISTICS OF GREATER SAGE-GROUSE HABITATS: A LANDSCAPE SPECIES AT MICROAND MACROSCALES / 69

John W. Connelly, E. Thomas Rinkes, and Clait E. Braun

5 • MOLECULAR INSIGHTS INTO THE BIOLOGY OF GREATER SAGE-GROUSE / 85

Sara J. Oyler-McCance and Thomas W. Quinn 6 • PREDATION ON GREATER SAGE-GROUSE: FACTS, PROCESS, AND EFFECTS / 95

Christian A. Hagen 7 • HARVEST MANAGEMENT FOR GREATER SAGEGROUSE: A CHANGING PARADIGM FOR GAME BIRD MANAGEMENT / 101

Kerry P. Reese and John W. Connelly 8 • PARASITES AND INFECTIOUS DISEASES OF GREATER SAGE-GROUSE / 113

Thomas J. Christiansen and Cynthia M. Tate 9 • WEST NILE VIRUS ECOLOGY IN SAGEBRUSH HABITAT AND IMPACTS ON GREATER SAGEGROUSE POPULATIONS / 127

Brett L. Walker and David E. Naugle

Part III • Ecology of Sagebrush 10 • CHARACTERISTICS OF SAGEBRUSH HABITATS AND LIMITATIONS TO LONG-TERM CONSERVATION / 145

Richard F. Miller, Steven T. Knick, David A. Pyke, Cara W. Meinke, Steven E. Hanser, Michael J. Wisdom, and Ann L. Hild 11 • PRE–EURO-AMERICAN AND RECENT FIRE IN SAGEBRUSH ECOSYSTEMS / 185

William L. Baker

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12 • ECOLOGICAL INFLUENCE AND PATHWAYS OF LAND USE IN SAGEBRUSH / 203

Steven T. Knick, Steven E. Hanser, Richard F. Miller, David A. Pyke, Michael J. Wisdom, Sean P. Finn, E. Thomas Rinkes, and Charles J. Henny 13 • INFLUENCES OF THE HUMAN FOOTPRINT ON SAGEBRUSH LANDSCAPE PATTERNS: IMPLICATIONS FOR SAGE-GROUSE CONSERVATION / 253

Matthias Leu and Steven E. Hanser 14 • INFLUENCES OF FREE-ROAMING EQUIDS ON SAGEBRUSH ECOSYSTEMS, WITH A FOCUS ON GREATER SAGE-GROUSE / 273

Erik A. Beever and Cameron L. Aldridge

Part IV • Population Trends and Habitat Relationships 15 • GREATER SAGE-GROUSE POPULATION DYNAMICS AND PROBABILITY OF PERSISTENCE / 293

Edward O. Garton, John W. Connelly, Jon S. Horne, Christian A. Hagen, Ann Moser, and Michael A. Schroeder 16 • CONNECTING PATTERN AND PROCESS IN GREATER SAGE-GROUSE POPULATIONS AND SAGEBRUSH LANDSCAPES / 383

Steven T. Knick and Steven E. Hanser 17 • INFLUENCES OF ENVIRONMENTAL AND ANTHROPOGENIC FEATURES ON GREATER SAGE-GROUSE POPULATIONS, 1997–2007 / 407

Douglas H. Johnson, Matthew J. Holloran, John W. Connelly, Steven E. Hanser, Courtney L. Amundson, and Steven T. Knick 18 • FACTORS ASSOCIATED WITH EXTIRPATION OF SAGE-GROUSE / 451

Michael J. Wisdom, Cara W. Meinke, Steven T. Knick, and Michael A. Schroeder

Part V • Conservation and Management 19 • GREATER SAGE-GROUSE AS AN UMBRELLA SPECIES FOR SHRUBLAND PASSERINE BIRDS: A MULTISCALE ASSESSMENT / 475

Steven E. Hanser and Steven T. Knick 20 • ENERGY DEVELOPMENT AND GREATER SAGE-GROUSE / 489

David E. Naugle, Kevin E. Doherty, Brett L. Walker, Matthew J. Holloran, and Holly E. Copeland 21 • ENERGY DEVELOPMENT AND CONSERVATION TRADEOFFS: SYSTEMATIC PLANNING FOR GREATER SAGE-GROUSE IN THEIR EASTERN RANGE / 505

Kevin E. Doherty, David E. Naugle, Holly E. Copeland, Amy Pocewicz, and Joseph M. Kiesecker 22 • RESPONSE OF GREATER SAGE-GROUSE TO THE CONSERVATION RESERVE PROGRAM IN WASHINGTON STATE / 517

Michael A. Schroeder and W. Matthew Vander Haegen 23 • RESTORING AND REHABILITATING SAGEBRUSH HABITATS / 531

David A. Pyke 24 • CONSERVATION OF GREATER SAGE-GROUSE: A SYNTHESIS OF CURRENT TRENDS AND FUTURE MANAGEMENT / 549

John W. Connelly, Steven T. Knick, Clait E. Braun, William L. Baker, Erik A. Beever, Thomas J. Christiansen, Kevin E. Doherty, Edward O. Garton, Steven E. Hanser, Douglas H. Johnson, Matthias Leu, Richard F. Miller, David E. Naugle, Sara J. Oyler-McCance, David A. Pyke, Kerry P. Reese, Michael A. Schroeder, San J. Stiver, Brett L. Walker, and Michael J. Wisdom Literature Cited / 565 Index / 625 Series Titles / 645

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CONTRIBUTORS

CAMERON L. ALDRIDGE

THOMAS J. CHRISTIANSEN

Natural Resource Ecology Laboratory Colorado State University; and U.S. Geological Survey 2150 Centre Avenue, Bldg. C Fort Collins, CO 80526-8118 [email protected]

Wyoming Game and Fish Department 351 Astle Avenue Green River, WY 82935 [email protected]

COURTNEY L. AMUNDSON

Department of Fisheries, Wildlife and Conservation Biology University of Minnesota 1980 Folwell Avenue St, Paul, MN 55108 [email protected] WILLIAM L. BAKER

Ecology Program and Department of Geography University of Wyoming Laramie, WY 82071 [email protected] ERIK A. BEEVER

U.S. Geological Survey Forest and Rangeland Ecosystem Science Center 3200 SW Jefferson Way Corvallis, OR 97331 (Current address: U.S. Geological Survey, Northern Rocky Mountains Science Center, 2327 University Way, Suite 2, Bozeman, MT 59715) [email protected]

JOHN W. CONNELLY

Idaho Department of Fish and Game 1345 Barton Road Pocatello, ID 83204 [email protected] HOLLY E. COPELAND

The Nature Conservancy Wyoming State Office Lander, WY 82520 [email protected] KEVIN E. DOHERTY

Wildlife Biology Program University of Montana Missoula, MT 59812 (Current address: U.S. Fish and Wildlife Service 3425 Miriam Avenue Bismarck, ND 58501) [email protected] SEAN P. FINN

U.S. Geological Survey Forest and Rangeland Ecosystem Science Center 970 Lusk Street Boise, ID 83706 sfi[email protected]

CLAIT E. BRAUN

JOHN C. FREEMUTH

Grouse Inc. 5572 North Ventana Vista Road Tucson, AZ 85750 [email protected]

Cecil Andrus Center for Public Policy Boise State University Boise, ID 83725 [email protected] vi i

EDWARD O. GARTON

JOSEPH M. KIESECKER

Department of Fish and Wildlife Resources University of Idaho Moscow, ID 83844 [email protected]

The Nature Conservancy Rocky Mountain Conservation Region Fort Collins, CO 80524 [email protected]

CHRISTIAN A. HAGEN

STEVEN T. KNICK

Oregon Department of Fish and Wildlife 63714 Parrell Road Bend, OR 97702 [email protected]

U.S. Geological Survey Forest and Rangeland Ecosystem Science Center 970 Lusk Street Boise, ID 83706 [email protected]

STEVEN E. HANSER

U.S. Geological Survey Forest and Rangeland Ecosystem Science Center 970 Lusk Street Boise, ID 83706 [email protected] CHARLES J. HENNY

U.S. Geological Survey Forest and Rangeland Ecosystem Science Center 3200 SW Jefferson Way Corvallis, OR 97331 [email protected]

MATTHIAS LEU

U.S. Geological Survey Forest and Rangeland Ecosystem Research Center Snake River Field Station 970 Lusk Street Boise ID, 83706 (Current address: Biology Department, College of William and Mary, P.O. Box 8795, 540, Landrum Drive, Integrated Science Center (ISC) RM-2129, Williamsburg, VA 23185) [email protected] CARA W. MEINKE

ANN L. HILD

Department of Renewable Resources University of Wyoming Laramie, WY 82009 [email protected] MATTHEW J. HOLLORAN

Wyoming Wildlife Consultants LLC Laramie, WY 82072 [email protected] JON S. HORNE

Department of Fish and Wildlife Resources University of Idaho Moscow, ID 83844 [email protected] DOUGLAS H. JOHNSON

U.S. Geological Survey Northern Prairie Wildlife Research Center 204 Hodson Hall 1980 Folwell Avenue St. Paul, MN 55108 [email protected]

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U.S. Geological Survey Forest and Rangeland Ecosystem Science Center 970 Lusk Street Boise, ID 83706 (Current address: Stantec Consulting, 9400 SW Barnes Road, Suite 200, Portland, OR 97225) [email protected] RICHARD F. MILLER

Eastern Oregon Agricultural Research Center 202 Strand Agricultural Research Center Oregon State University Corvallis, OR 97331 [email protected] ANN MOSER

Idaho Department of Fish and Game 600 S. Walnut P.O. Box 25 Boise, ID 83707 [email protected] DAVID E. NAUGLE

Wildlife Biology Program University of Montana Missoula, MT 59812 [email protected]

NO. 38

Knick and Connelly

SARA J. OYLER-MCCANCE

MICHAEL A. SCHROEDER

U. S. Geological Survey Fort Collins Science Center, 2150 Centre Avenue Building C Fort Collins, CO 80526 [email protected]

Washington Department of Fish and Wildlife P.O. Box 1077 Bridgeport, WA 98813 [email protected] SAN J. STIVER

Western Association of Fish and Wildlife Agencies 2184 Richard St. Prescott, AZ 86301 [email protected]

AMY POCEWICZ

The Nature Conservancy Wyoming State Office Lander, WY 82520 [email protected]

CYNTHIA M. TATE DAVID A. PYKE

U.S. Geological Survey Forest and Rangeland Ecosystem Science Center 3200 SW Jefferson Way Corvallis, OR 97331 [email protected]

Wyoming Game and Fish Department Wyoming State Veterinary Laboratory 1174 Snowy Range Road Laramie, WY 82070 [email protected] W. MATTHEW VANDER HAEGEN

THOMAS W. QUINN

Rocky Mountain Center for Conservation Genetics and Systematics Department of Biological Sciences University of Denver Denver, CO 80208 [email protected] KERRY PAUL REESE

Department of Fish and Wildlife Resources University of Idaho Moscow, ID 83844 [email protected]

Washington Department of Fish and Wildlife 600 Capitol Way North Olympia, WA 98501 [email protected] BRETT L. WALKER

Wildlife Biology Program University of Montana Missoula, MT 59812 (Current address: Colorado Division of Wildlife, 711 Independent Ave., Grand Junction, CO 81505) [email protected] MICHAEL J. WISDOM

E. THOMAS RINKES

U.S. Bureau of Land Management 1387 Vinnell Way Boise, ID 83709 [email protected]

U.S.D.A. Forest Service Pacific Northwest Research Station 1401 Gekeler Lane La Grande, OR 97850 [email protected]

CONTRIBUTORS

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PREFACE

Population declines of Greater Sage-Grouse (Centrocercus urophasianus) coupled with extensive loss of their sagebrush (Artemisia spp.) habitats prompted nine petitions from 1999 to 2005 for protection under the Endangered Species Act. As a result, the United States Fish and Wildlife Service (USFWS) faced the daunting task of determining status and trends for a species and its habitat currently distributed across 11 states and two provinces in western North America. The difficulty of this task was in large part due to complex interactions among invasive plants, altered fire and disturbance regimes, and land use. The Conservation Assessment of Greater Sage-Grouse and Sagebrush Habitats (Connelly et al. 2004) was written for the Western Association of Fish and Wildlife Agencies and was intended to aid the USFWS’s range-wide listing decision issued in 2005. The assessment provided detailed information on Greater Sage-Grouse biology as well as analysis of population and habitat trajectories. It now provides the foundation for this volume. The USFWS determined in March 2010 that Greater Sage-Grouse warranted listing under the Endangered Species Act, but that immediate action was precluded by higher priorities. Coalitions of private, state, federal, and nongovernmental entities have been formed, conservation strategies developed, and management actions to benefit sage-grouse and restore sagebrush have been implemented. Sage-grouse and sagebrush are priority concerns for industry, conservation organizations, and management agencies in the United

States and Canada. Yet thousands of square kilometers of sagebrush steppe burn each year or are invaded by exotic plants, energy development has accelerated and influenced many more square kilometers of sage-grouse habitat, West Nile virus has infected local populations of sage-grouse, human densities in the western United States and their use of wildlands have increased, and sage-grouse populations continue to decline in many parts of their range. In this volume, we have documented the status of Greater Sage-Grouse populations and their sagebrush habitats and identified factors that influence their long-term persistence. We have revised, updated, and reconfigured the original content of the 2004 assessment. Gaps caused by time limits in preparing the assessment have been addressed with new chapters that have greatly expanded the scope and depth of information contained in this volume. Management considerations were not provided in the 2004 assessment to avoid influencing policy direction. These now have been developed in a conservation implications section at the conclusion of each chapter. This volume reflects a broad spectrum of expertise in sagebrush and sage-grouse ecology. Thirtyeight authors represented 20 organizations, including 6 state agencies, 3 federal agencies, 6 universities, and 5 nongovernmental organizations. In all, 63 reviewers represented 39 organizations, including 8 state agencies, 8 federal agencies, 18 universities, and 5 nongovernmental organizations. Only 11 individuals both authored and reviewed chapters.

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We appreciate the support of the Cooper Ornithological Society in providing the forum for this collection. Carl Marti1, series editor of Studies in Avian Biology, ensured the consistency and quality of the chapters. All chapters have been peer reviewed, most by both an expert in sage-grouse or sagebrush ecology and an outside expert in the ecological topic. At least one and often both associate editors reviewed each chapter in addition to the technical editor. Chapters authored by United States Geological Survey (USGS) scientists received an agency policy review, although other chapters and the volume as a whole do not necessarily reflect the views of the USGS, the Western Association of Fish and Wildlife Agencies, or any other state or federal agency. The volume has not been endorsed by any private, state, or federal organization but was developed solely as a scientific peer-reviewed publication by the Cooper Ornithological Society. Any use of trade, product, or firm names is for descriptive purposes only and does not imply endorsement by the United States government. We thank the USGS Forest and Rangeland Ecosystem Science Center (STK) and Idaho Department of Fish and Game (JWC). Kathy Peter and Susan Haseltine of the United States Geological Survey provided financial and logistical support.

Carol Schuler, Kate Kitchell, Sue Phillips, and Ruth Jacobs provided administrative support as well as much-needed encouragement. Elena Velasquez and Martina Zucchini translated abstracts into Spanish. The USGS funded page charges. We dedicate this volume to the agency biologists who had the foresight to form the first Western Sage and Columbian Sharp-Tailed Grouse Technical Committee over 50 years ago for the purpose of developing the cooperation across administrative boundaries necessary to conserve prairie grouse. These individuals include Paul D. Dalke, Robert L. Eng, Clifton W. Greenhalph, Gordon W. Gullion, J. Burton Lauckhart, Levon Lee, A. Starker Leopold, Jessop B. Low, Donald D. MacLean, W. V. Masson, Otto C. Nelson, Robert L. Salter, and Charles F. Yocom. Their vision provides a model of collaboration that must be sustained if we are to conserve sage-grouse. Without their efforts, much of our work would not have been possible. We present the information in this volume in hopes that generations long into the future will be able to experience the sun rising on a vast open landscape, smell the pungent scent of sagebrush, and marvel at the centuries-old rite of Greater Sage-Grouse displaying on a lek. STEVEN T . KNICK JOHN W . CONNELLY

1 Carl Marti died during the production of this volume. We will remember working with him as the series editor and also as a colleague and a friend.

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FOREWORD

Thoughts on the Role of Science in Making Public Policy John C. Freemuth

The Greater Sage-Grouse (Centrocercus urophasianus) has become a species whose listing status under the Endangered Species Act (ESA) is fraught with controversy. The recent past has seen court cases, high-level political intervention, and disputes over what constitutes the best available scientific information. Yet the conflict over the warranted listing of the sage-grouse presents an opportunity to engage the public about what is known about the science of the sage-grouse and its sagebrush habitat, and to promote public deliberation about possible solutions. That opportunity can perhaps raise public confidence in science and make the blatant politicizing of science more difficult. S E T T ING T HE STAGE The role of science and expertise in making public policy may strike many readers as a relatively modern concern. Certainly, recent events surrounding the listing decisions for the sage-grouse have brought that role to the foreground and jump-started a new and intense discussion. But we have talked about that role for a long time. Woodrow Wilson, the only president to hold a Ph.D., noted in 1912, What I fear, therefore, is a government of experts. God forbid that in a democratic country we should resign the task and give the government over to experts. What are we for if we are to be scientifically taken care of by a small group

of gentlemen who are the only men who understand the job? Because if we don’t understand the job, then we are not a free people. We ought to resign our free institutions and go to school to somebody and find out what it is we are about. (Davidson 1956:83, as quoted in Smith 1991:2)

Wilson was not bashing experts but asking us to think about their appropriate role in a democracy. Today, the role of science in public land management is the question before us. It remains part of the context of decisions over whether to list the sage-grouse as an endangered species. I argue that that role can help us learn what we are about regarding our federal lands, while continuing to make decisions grounded in democratic practice. I had the privilege of serving on, then chairing, the Science Advisory Board of the Bureau of Land Management (BLM) during the late 1990s. The board’s purpose was to work with the BLM and its director on improving the communication of the bureau’s science and research needs to other agencies, scientists, the U.S. Congress, and the public; helping get scientific findings to BLM field office staff; and coping with resource management issues. The board lasted until the early years of the Bush administration when its Federal Advisory Committee Act charter was not extended. One of the most interesting and important projects of the board and BLM was writing the bureau’s science strategy. That strategy was to

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delineate the role of science in BLM decision making and public land management. The strategy noted: Science is useful for evaluating alternatives and estimating outcomes. However, it is not the sole factor in making decisions because the state of natural resource science is often insufficient to give definitive cause-effect predictions. Unknowns and uncertainties will always be associated with predictions of decision outcomes. Science may reduce but can never completely eliminate the uncertainty regarding future events. However, the use of the best-available science—along with a consideration of political, social, and economic information—will result in the best-informed decisions. (United States Bureau of Land Management 2000:3–5)

In this document, science was considered a necessary but insufficient condition for making BLM management decisions. It was also my experience, both on the board and as someone who pays attention to the academic and actual worlds of federal land policy, that science can occupy other places in discussions over its role in public-land policy decisions. Indeed, it was likely that most of those involved with the writing of the statement above, as well as many other observers, were aware that there was quite a bit of debate and confusion over the role of science in natural resource and public lands decision making. T WO IS S UE S I N THE SCIEN CE-POL ICY R E LA T IONS H IP Two regularly occurring issues influence any attempt to untangle the science-policy relationship. The first issue concerns viewing science as a process versus science as truth. Science as process refers to the use of the scientific method to produce knowledge, which is then subjected to peer review and published in professional journals. All chapters of this volume of Studies in Avian Biology were subjected to peer review, revised, and, in some cases, reviewed again, certainly meeting the process test. The science-as-truth claim refers to the use of science as the proper way to solve policy conflicts over other ways of deciding. To use the BLM quote above as an example, the science-as-truth claim would thus disallow political, social, and economic concerns. Authors of this document

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did not make any such claim about science, nor were they even given the task of considering such concerns, but it nonetheless remains an important issue in the science-policy dialogue.

Science as Process An extension of the process discussion is a question of particular concern to those who work on public-land management conflicts such as that of the sage-grouse. Is the knowledge produced helpful to land managers and other decision makers? This is a difference between applied and basic science. During my tenure with the BLM board, agency scientists often discussed being evaluated and rewarded for publication in journals, even though the information needed by land managers was of a different sort—more immediate and thus less able to go through long peer-review processes—but it had to be credible or it simply would not stand up in court or anywhere else. This volume is a good example. It has gone through a long and exhaustive review process both to ensure science credibility and to stand up in court. Beyond the immediate purpose of providing information for determining the listing status of Greater Sage-Grouse, how will it get used by managers and other decision makers to influence public-land policy? It was also not clear whether land managers did a very good job at explaining to scientists both within and outside an agency (like BLM) how they made decisions. To put it differently, how did they use science to make a decision while still considering political, social, and economic information, especially under conflicting multiple-use laws? Having this kind of discussion would go far in allowing people to see the alternate perspectives in the science–public policy arena. Indeed, agency-based meetings or conferences should be designed to do exactly that. Many suspect that more may be going on with the process question than how managers use science. Scientists always need to be transparent about how their own values did or did not inform their research choices. In 1994, a fascinating debate appeared in the journal Conservation Biology over whether conservation biologists should link arms with activists in efforts to reform grazing practices (Brussard et al. 1994). Other noted conservation biologists were concerned that conservation biologists might damage their credibility NO. 38

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by openly advocating and supporting political positions. They suggested asking instead how livestock grazing can be managed to have the fewest impacts on biodiversity and ecosystem integrity (Brussard et al. 1994). They worried about the flaw of deductive reasoning which leads to a research question that states: “range management must be dramatically reformed.” . . . Our work as scientists involves recognizing patterns based on data and only then formulating a general rule. More importantly, how can we hope to advance society’s mission to preserve biological diversity if our audience of policymakers assumes that we intend to prove a presumed conclusion instead of attempting to falsify well-framed null hypotheses? (Brussard et al. 1994:920)

In this example, we see well-respected scientists having a polite discussion about the use of scientific information and how it informs policy. What the authors caution against is an approach that confuses scientific assertions with value judgments (grazing reduces biodiversity) and assumptions (grazing has a negative influence on ecosystem integrity). One begins with a supposedly neutral hypothesis arising from a presumed conclusion that grazing is bad or grazing must be reformed. It might then be a relatively easy to find the science in support of the hypothesis. The more appropriate hypothesis in this argument ought to be along the lines that grazing has no effect on biological diversity. If it is falsified, then people can begin to speak of reform. Of course, as Brussard et al. (1994) also note, sometimes the effects of grazing could be positive. We also know that all of these hypotheses are very specific to place and perhaps not as useful in such a complex issue as the sage-grouse status. The authors remind us that good science starts with data, looks for patterns, then forms general rules that are subject to further testing through hypothesis development. A second process-related issue concerns the influence of political appointees on the production and dissemination of science to be used in decision making. If science is indeed necessary (and the requirements and legal interpretations of many laws seem to make that a noncontroversial statement), then it can become strategic to influence what that science might be asserting. In May 2008, the United States Government Accountability

Office issued a report on United States Fish and Wildlife Service reviews of ESA decisions (including Greater Sage-Grouse and Gunnison SageGrouse [Centrocercus minimus]) during the George W. Bush administration’s tenure and concluded that political appointees indeed had intervened. Not surprisingly, there was debate over that role and whether or not there was undue pressure on scientists to alter documents and conclusions. As one former official, Craig Manson, put it, “the fact was the assistant secretary’s office took a very active role in the ESA program, and that’s perfectly proper for the assistant secretary’s office to do so” (Manson 2008:A-7). Manson is correct. Political appointees do take an active role in these sorts of decisions, and they should. They represent the president and the president’s views. They are accountable to the president, and they are accountable to Congress and to all Americans. In the best of circumstances, they have to navigate a labyrinth of laws, values, and conflicting expectations. They are not given the task of simply following best science, because best science is not a trump. However, how they take an active role is important. As one reviewer of this introduction noted, changing data, altering reports, and attempting to intimidate scientists is unacceptable and destroys the integrity of the process. We could decide as a society that best science ought to be the trump. We would have to do that through legislation, and in doing that, we open up a number of other issues regarding science and democracy. Such a solution might be thought extreme by many, though perhaps not by many of my friends working on the science needed for federal land-management decisions. More realistically, what is needed for this particular problem, based on the actions of political appointees in this case, is an administrative framework, a series of guidelines that make it difficult for the excesses reported by the Government Accountability Office to occur. Such a framework ought to provide an open and defensible accounting of why certain scientific information was edited, replaced, not allowed, and so on. After all, this Studies in Avian Biology volume, used by the United States Fish and Wildlife Service in its listing decision, must by law be made easily accessible to the public. There should not be a mysterious administrative process that only partially reveals and certainly does not explain why a decision was made. This is

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similar to the earlier call for scientists to be clear about their values and to disentangle those values from their science. Decision makers ought to be clear, too.

Science as Truth The second issue, science as truth, involves the value one places on science and the knowledge it produces. A recent article by Lockwood (2008) on the mythic cowboy sets the stage for this perspective: From the first-hand perspective of a scientist—an insect ecologist, in particular—I can assure you that science cannot provide the answers to the deep questions concerning what we ought to do with western people and lands. Science is necessary, but it is not sufficient. And here’s why. In the Myth of Scientism, the hero is an absolutely objective, rational discoverer of unchanging physical truths (like Isaac Newton and the apple incident, which of course never happened—but we know that myths don’t collapse under the weight of facts). The scientist reveals how the world really is, not some fanciful tale replete with human desires. The problem, of course, is that according to this myth, science is value-free. So the hero can offer no moral lessons regarding our treatment of one another or the land.

If it claims to answer value questions, science, from this second perspective, moves from a way of knowing grounded in a process to a purveyor of truths. It has become a form of higher law. As Gregg Cawley and I have pointed out (Cawley and Freemuth 2007), one of the leading writers in public administration suggests that “the imperative of higher law is always conceived as derived from what is most valid, most powerful, most highly honored. Historically this has most frequently been God. But in the late nineteenth and early twentieth century America it has often been SCIENCE” (original emphasis; Waldo 1984:157). The science-as-truth claim suggests that it can answer questions not simply about what might or could be done but about what should be done. Is there a path that leads away from the problems associated with these two issues? Perhaps, but it requires patience and dialogue. As Albert Teich, director of science and policy programs for the American Association for the Advancement of Science, recently wrote:

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The dominant mode of scientists engaging with the lay public in the past has been “public understanding of science”—an approach based on the belief that if people only knew more about science, they would see the world the way scientists do. This is a paternalistic posture, one that is based on a failure to recognize the legitimacy of people’s values when they conflict with those of the scientist. The constituencies whose values are reflected in some of the political positions of the administration with which many scientists disagree are not necessarily uneducated or irrational. Rather, they have a different perspective on these issues, one that places a different value on the costs and risks versus the benefits of stem cell research, reductions in carbon dioxide emissions, or the preservation of an obscure species of aquatic plant that may block construction of a dam. They need to be engaged in genuine two-way conversation—a dialogue—rather than dismissed out of hand, or “educated.” (Teich 2008:22)

Before scientists get upset at Teich’s remarks, they might consider that some of the scientists who capture the public imagination are those who effectively communicate the excitement and challenges of modern science, such as the late Carl Sagan. A PO S S IBL E PAT H We should experiment with new forms of decision making and public discourse. Let us create a sage-grouse science and public policy deliberative forum, where scientists, the public, and interested decision makers might assemble to talk through the issues surrounding a possible listing of the sage-grouse. Scientists known for their skills in communication could present their best science in an accessible and understandable way, while letting us know how their values and assumptions guided and informed their work. This would mean distilling the diverse offerings that make up this volume, but that can be done, and scientists should be rewarded for doing it. Scientists could also debate and question each other so the public could see science at work. The public could ask questions and talk about the scientific information they think would be useful, but they should be prepared to put their own assumptions and values on the table as well, such as whether they have already decided that they are for or against

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listing. Conserving sage-grouse and sagebrush are very good goals, but some of the measures required to do that may not be accepted by an expanding human population in the western United States that has other priorities. We need to know if that is true. We should hear as well the perspective of various sage-grouse working groups throughout the West who provide a useful way to link questions of science to questions of public and agency concerns and values. Managers and decision makers ought to come—indeed, ought to be compelled to come—and talk about the factors that govern their listing decisions and the role that science plays or

does not play in those decisions. Together, perhaps all parties could develop the sort of guidelines that should be used by managers and decision makers in reviewing the scientific information needed for an effective listing decision. This process would take much work, but it might be more sustainable, and certainly would be more democratic, than our current one. This volume is a hugely important part of this process. The work that has gone into it is extraordinary. But we all need to participate in developing a better way to link it to the other processes that will lead to a decision about the future of the sage-grouse and its habitats.

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GREATER SAGE-GROUSE AND SAGEBRUSH

An Introduction to the Landscape Steven T. Knick and John W. Connelly

The Greater Sage-Grouse (Centrocercus urophasianus) is often called an icon of the West because the species has become the symbol for conserving sagebrush (Artemisia spp.) ecosystems, one of the most difficult environmental challenges in North America. Sage-grouse have undergone long-term population declines and now are absent from almost half of their estimated distribution prior to Euro-American settlement (Schroeder et al. 2004) (Fig. I.1). Overall, population trends have been more stable in recent years, although sage-grouse numbers are still declining in some regions (Connelly et al. 2004). Proximate reasons for population declines differ across the sage-grouse distribution, but ultimately, the underlying cause is loss of suitable sagebrush habitat (Connelly and Braun 1997, Leonard et al. 2000, Aldridge et al. 2008). Some form and quantity of sagebrush within the landscape are necessary to meet seasonal requirements for food, cover, and nesting of sage-grouse (Patterson 1952, Connelly et al. 2000c). Thus, conserving and managing Greater Sage-Grouse is as much about the ecology of the bird as it is about understanding the dynamics of sagebrush ecosystems (Connelly et al. 2000c, Crawford et al. 2004). Concluding that loss and degradation of sagebrush-dominated landscapes cause sage-grouse population declines is deceptively simple, much

like the ecosystems themselves. Many species of conservation concern have either restricted ranges or a small number of factors contributing to their plight. In comparison, sage-grouse currently occupy 670,000 km2 spanning parts of 11 western states and two Canadian provinces (Schroeder et al. 2004). This broad distribution encompasses highly diverse environments and an extensive array of ecological stressors. At least 11 species of sagebrush occur within the sage-grouse range, each differing in their specific plant community structure, productivity, resilience, and resistance to disturbance (West and Young 2000, Miller and Eddleman 2001). The distribution and influence of multiple land uses also vary widely across the sage-grouse distribution, as summarized in Knick et al. (this volume, chapter 12). Conversion to croplands has eliminated or fragmented sagebrush in areas having deep fertile soils or irrigation potential. Sagebrush remaining in these areas has been reduced to agricultural edges or to relatively unproductive environments. Oil and gas resources are being developed primarily in the eastern portion of the sage-grouse range (Knick et al. 2003; Naugle et al., this volume, chapter 20), but exploration and development of wind and geothermal energy is increasing rapidly in many regions. Livestock grazing occurs throughout the sage-grouse range but has a

Knick, S. T. and J. W. Connelly. 2011. Greater Sage-Grouse and sagebrush: an introduction to the landscape. Pp. 1–9 in S. T. Knick and J. W. Connelly (editors). Greater Sage-Grouse: ecology and conservation of a landscape species and its habitats. Studies in Avian Biology (vol. 38), University of California Press, Berkeley, CA.

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Figure I.1. Current and pre–Euro-American settlement distribution of sage-grouse in North America (Schroeder et al. 2004) within the Sage-Grouse Conservation Area (SGCA). Gunnison Sage-Grouse ranges also are included in southeastern Utah and southwestern Colorado. The SGCA was delineated by buffering the potential pre–Euro-American settlement distribution by 50 km.

more diffuse influence on soils and vegetation in contrast to land uses that remove or fragment habitat. Urbanization and human densities are increasing in the western United States as people choose to live near wilderness and recreation areas (Brown et al. 2005, Hansen et al. 2005). New corridors proposed for energy transmission would affect another 2% of the current sagebrush distribution. Less than 5% of the sage-grouse range presently is >2.5 km from a mapped road. Recreation, including off-road vehicles, is rapidly increasing on public lands. Leu and Hanser (this volume, chapter 13) describe how the collective effect of this human footprint influences the landscape structure of sagebrush-dominated habitats for sage-grouse. The Greater Sage-Grouse is a landscape species; their large annual ranges can encompass >2,700 km2 (Dalke et al. 1963, Schroeder et al. 1999, Leonard et al. 2000). Movements from lek sites used for breeding to nesting locations can exceed 25 km (Holloran and Anderson 2005), and seasonal ranges can be >80 km apart (Connelly et al. 1988). Sage-grouse use a variety of landscapes within their annual

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range that, at broad scales, range from expanses dominated by sagebrush to heterogeneous mosaics of grass and shrublands (Connelly et al. 2004). Large ranges and complex habitat selection challenge conservation because management focused on one area, usually centered around leks or habitat configuration and often emphasizing nesting and brood rearing, may not have the intended benefit when other seasonal ranges or habitat components may be at least as critical (Connelly et al. 2000c, Doherty et al. 2008). The large ranges also can encompass lands under multiple ownerships and uses. Knick (this volume, chapter 1) reviews the historical and legislative background that shaped the development of sagebrush landscapes and resulted in differing land characteristics among the mosaic of owners. Stiver (this volume, chapter 2) describes the efforts among public and private entities having diverse perspectives engaging in the collaborative use of sagebrush lands. These conservation challenges are not unique to Greater Sage-Grouse and sagebrush. Fourteen of 18 grouse species receive national status of concern

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in at least one country, primarily because of habitat degradation and loss (Storch 2000). Globally, temperate grasslands, savannahs, and shrublands are the least protected biomes and have experienced extensive conversion to agriculture or invasion by exotic plant species (Brooks et al. 2004b). Extensive loss of grasslands and shrublands in the Western Hemisphere has led to population declines for many species of birds obligate to these systems (Peterjohn and Sauer 1999, Vickery et al. 1999, Brennan and Kuvlesky 2005, Askins et al. 2007). This volume presents a multifaceted view of the ecology of Greater Sage-Grouse and sagebrush from wildlife biologists, landscape ecologists, and shrubland biologists. Authors and reviewers represented a broad expertise in research and management from across the sagegrouse range and were drawn from agency, academic, and private sectors. Thus, the syntheses of published literature combined with new analyses for this volume provide a foundation to develop effective approaches to conservation. History and our current use of the vast landscapes dominated by sagebrush can tell us much about land use, priorities, values, and resource management. The future will tell others about the effectiveness of conservation actions we implement today. DIS T RI B UT I O N O F GR E A T E R S A GE-GROU SE Sage-Grouse Conservation Area The Sage-Grouse Conservation Area (SGCA) used throughout this volume to define the spatial extent of analyses was delineated from the estimated presettlement distribution of sage-grouse (Connelly et al. 2004, Schroeder et al. 2004). We added a 50-km buffer to the pre-settlement distribution to include adjacent factors, such as urban centers or agricultural regions, which may have contributed to previously extirpated populations (Aldridge et al. 2008) or to current trends (Fig. I.1). The SGCA includes parts of 14 U.S. states and three Canadian provinces encompassing 2,063,000 km2. We first described this region as the boundary for conducting the range-wide conservation assessment (Connelly et al. 2004) and have carried it forward to this volume. No legal basis or agency endorsement exists for this region or our designation of the area. Sagebrush is the dominant land cover on 500,000 km2 within the SGCA (Fig. I.2). Of the

sagebrush species, three subspecies of big sagebrush (Wyoming big sagebrush [Artemisia tridentata ssp. wyomingensis], basin big sagebrush [A. t. ssp. tridentata], and mountain big sagebrush [A. t. ssp. vaseyana]), two low forms (little sagebrush [A. arbuscula] and black sagebrush [A. nova]), and silver sagebrush (A. cana) are the most important to Greater Sage-Grouse (Connelly et al. 2000c). Connelly et al. (this volume, chapter 4) provide a summary of habitat requirements by Greater SageGrouse in a synthesis of published literature. Miller et al. (this volume, chapter 10) describe the characteristics of these primary sagebrush communities within the sage-grouse range and the challenges that exotic plant species, altered fire regimes, and climate change present to long-term conservation. Pyke (this volume, chapter 23) discusses the difficulties in restoring and rehabilitating these arid-land systems.

Sage-Grouse Management Zones The Western Association of Fish and Wildlife Agencies defined seven Sage-Grouse Management Zones for assessing population and habitat trends independent of administrative and jurisdictional boundaries (Fig. I.3; Stiver et al. 2006). Management zones originally were delineated from floristic provinces, within which similar environmental factors influence vegetation communities (West 1983b, Miller and Eddleman 2001). Boundaries of management zones subsequently have been redefined, particularly in Montana and Wyoming, to better reflect potential linkages among populations and to include known leks outside the original zones (S.J. Stiver, pers. comm.). H IE RARC H IC AL O RG AN IZAT IO N O F S AG E BRU S H S Y S T E M S This volume emphasizes regional and range-wide themes that describe sagebrush systems. Authors synthesized results from site-specific studies into broader patterns. Other analyses aggregated localor small-scale events, such as individual fires or counts of sage-grouse at a lek, to provide regional or range-wide patterns of sagebrush communities, disturbance regimes, land use, and sagegrouse behavior and population dynamics. This broad-scale perspective was emphasized because monitoring is most effective when conducted and interpreted at population or range-wide scales (Connelly et al. 2003b, Reese and Bowyer 2007).

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Figure I.2. Distribution of sagebrush land cover within the Sage-Grouse Conservation Area based on land cover mapped in the early 2000s in western North America. The map represents the percent of the landscape dominated by sagebrush land cover (Miller et al., this volume, chapter 10), not site-specific percentages of ground cover. As such, the map delineates a general representation of sagebrush distribution.

Figure I.3. Sage-Grouse Management Zones within the Sage-Grouse Conservation Area (Stiver et al. 2006).

Figure I.4. Hierarchical organization of sagebrush and sage-grouse ecosystems.

Conservation planning and actions also have the greatest benefit for Greater Sage-Grouse when local actions are coordinated at regional or rangewide scales (Hemstrom et al. 2002; Wisdom et al. 2002c, 2005a). However, the validity of this paradigm rests on understanding the organizational structure of sagebrush systems. The model of sagebrush systems as a hierarchical organization arranged along spatial and temporal scales is one of the unifying concepts underlying the information presented in this volume (Fig. I.4). This model presents ecological systems as an integrated assemblage of patterns and processes at smaller scales enclosed within successive levels at larger scales (Allen and Starr 1982, O’Neill et al. 1986, Urban et al. 1987). Levels that are adjacent in this hierarchy are most likely to share similar characteristics (Kotliar and Wiens 1990). Events in widely separated levels may be uncorrelated, and extrapolation of results across scales may lead to incorrect conclusions if organizational structure is not considered (Wiens 1981, 1989b). Sagebrush systems have been studied most often in relatively small spatial and short temporal scales (Brown et al. 2002). Plant communities at sites functioned through time but were not considered as interacting parts of a larger landscape (West 2003b). Modeling sagebrush as a multiscale hierarchy can help illuminate landscape dynamics and long-term changes. Landscapes are mosaics of smaller patches shaped by disturbance patterns that have characteristic spatial and temporal scales

(Delcourt et al. 1983, Levin 1992). Small-scale disturbances are most frequent, but rapid recovery periods maintain relatively stable patterns at larger scales (Urban et al. 1987). Unbalanced dynamics of disturbance relative to recovery at smaller scales can change patterns observed at larger scales. Miller et al. (this volume, chapter 10) describe fire characteristics for each of the Sage-Grouse Management Zones based on fire polygons mapped since the 1980s. Baker (this volume, chapter 11), using a landscape approach to estimate pre–EuroAmerican and current fire rotations, discusses how changes in size and frequency of fires relative to recovery have changed in sagebrush systems. Both chapters conclude that recovery periods are insufficient relative to the characteristics of the new disturbance regime. Consequently, sagebrush landscapes are being converted to exotic grasslands at scales sufficiently large to influence future disturbance dynamics and sage-grouse distributions. A hierarchical perspective may also better describe habitat selection by sage-grouse, as presented by Connelly et al. (this volume, chapter 4). Conceptual models of hierarchical selection predict that coarse features at large spatial scales are important for initial location but finer features in the environment become primary factors in selecting individual sites (Johnson 1980, Kristan 2006). Site-specific characteristics are the most commonly studied components of habitat selection by sage-grouse (Connelly et al. 2000c, Hagen et al. 2007). Most research

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reviewed by Connelly et al. (this volume, chapter 4) was focused on fine-scale features, such as percent canopy cover and height of sagebrush selected by sage-grouse for nesting or brood rearing. Yet largescale (2–50 km2) features are important influences on seasonal use areas, population trends, and persistence (Holloran and Anderson 2005, Doherty et al. 2008). The presence of relatively large (thousands of km2) expanses of sagebrush-dominated shrub steppe at broad spatial scales is necessary to support sage-grouse populations (Connelly et al. 2000c, Aldridge et al. 2008). However, the appropriate patch size or landscape configuration and spatial scale(s) are uncertain. Habitat features and spatial scale of selection by sage-grouse are explored by Hanser and Knick (this volume, chapter 19), Connelly et al. (this volume, chapter 4), and Johnson et al. (this volume, chapter 17). Anthropogenic land use has altered landscapes used by Greater Sage-Grouse in most parts of their range (Knick et al. 2003, Connelly et al. 2004). However, land uses differ in the multiscale signature they impose (Knick and Rotenberry 1997). The Columbia Basin consists primarily of a matrix dominated by agriculture at large scales and interspersed sagebrush patches at small scales. In contrast, oil and gas development in the Wyoming Basin creates fragmentation in landscapes dominated by sagebrush at larger scales. Sage-grouse use each region, although the commingled effect of fragmentation and habitat loss at different scales has influenced each of these populations (Schroeder et al. 2000, Doherty et al. 2008). The diversity of areas used by sage-grouse across their range suggests that we have not identified either the appropriate scales or the relative importance of habitat arrangements across multiple scales. Consequently, our ability to fully understand effects of land uses and alterations often is limited to correlation rather than cause and effect. The hierarchical scaling of sage-grouse as individuals, populations, and metapopulations (Fig. I.4) provides perspective for understanding different sources of mortality. Predation (Hagen, this volume, chapter 6), harvest (Reese and Connelly, this volume, chapter 7), and disease (Christiansen and Tate, this volume, chapter 8) are significant to individuals or local groups but are not significant factors influencing population trends. Similarly, West Nile virus (Walker and Naugle, this volume, chapter 9) has the potential to significantly decrease sage-grouse numbers or eliminate

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relatively small peripheral populations, but the effect on range-wide trends is less clear. The range-wide distribution of sage-grouse, although encompassing a broad spatial extent, is composed primarily of small, mostly independent units. Forty-one populations within the Greater Sage-Grouse range previously were identified using geographic or physical barriers to movements to guide delineation; five spatially large populations were further divided into 24 subpopulations (Connelly et al. 2004). Dispersal distances to new breeding areas by juvenile sage-grouse from the leks where their mother was marked are not well documented but suggest that most distances are