Table of Contents

Study Area Map

A Note on Accompanying Video
Robert K. Landis

Foreword
Jane Goodall

Preface
Douglas W. Smith, Daniel R. Stahler, and Daniel R. MacNulty
Part 1 History and Reintroduction
1 Historical and Ecological Context for Wolf Recovery
Douglas W. Smith, Daniel R. Stahler, Daniel R. MacNulty, and Lee H. Whittlesey

Box 1.1 Wolf History and Surveys in Yellowstone National Park John Weaver
2 How Wolves Returned to Yellowstone Steven H. Fritts, Rebecca J. Watters, Edward E. Bangs, Douglas W. Smith, and Michael K. Phillips
Box 2.1 To Reintroduce or Not to Reintroduce, That Is the Question
Diane Boyd
Guest Essay: Why Are Yellowstone Wolves Important? L. David Mech
Part 2 Behavioral and Population Ecology

3 Essential Biology of the Wolf: Foundations and Advances
Daniel R. MacNulty, Daniel R. Stahler, Tim Coulson, and Douglas W. Smith

4 Ecology of Family Dynamics in Yellowstone Wolf Packs Daniel R. Stahler, Douglas W. Smith, Kira A. Cassidy, Erin E. Stahler, Matthew C. Metz, Rick McIntyre, and Daniel R. MacNulty
Box 4.1 Naming Wolf Packs Daniel R. Stahler
5 Territoriality and Competition between Wolf Packs Kira A. Cassidy, Douglas W. Smith, Daniel R. Stahler, Daniel R. MacNulty, Erin E. Stahler, and Matthew C. Metz
 Box 5.1 Auditory Profile: The Howl of the Wolf 
John B. Theberge and Mary T. Theberge
6 Population Dynamics and Demography Douglas W. Smith, Kira A. Cassidy, Daniel R. Stahler, Daniel R. MacNulty, Quinn Harrison, Ben Balmford, Erin E. Stahler, Ellen E. Brandell, and Tim Coulson
Guest Essay: Yellowstone Wolves Are Important Because They Changed Science Rolf O. Peterson and Trevor S. Peterson
Part 3 Genetics and Disease

7 Yellowstone Wolves at the Frontiers of Genetic Research Daniel R. Stahler, Bridgett M. vonHoldt, Elizabeth Heppenheimer, and Robert K. Wayne
8 The K Locus: Rise of the Black Wolf Rena M. Schweizer, Daniel R. Stahler, Daniel R. MacNulty, Tim Coulson, Phil Hedrick, Rachel Johnston, Kira A. Cassidy, Bridgett M. vonHoldt, and Robert K. Wayne
9 Infectious Diseases in Yellowstone’s Wolves Ellen E. Brandell, Emily S. Almberg, Paul C. Cross, Andrew P. Dobson, Douglas W. Smith, and Peter J. Hudson
Guest Essay: Why Are Yellowstone Wolves Important? A European Perspective Olof Liberg
Part 4 Wolf-Prey Relationships

10 How We Study Wolf-Prey Relationships Douglas W. Smith, Matthew C. Metz, Daniel R. Stahler, and Daniel R. MacNulty
 Box 10.1 Nine-Three-Alpha
Douglas W. Smith
 Box 10.2 The Bone Collectors
Ky Koitzsch and Lisa Koitzsch
11 Limits to Wolf Predatory Performance  Daniel R. MacNulty, Daniel R. Stahler, and Douglas W. Smith
 Box 11.1 Tougher Times for Yellowstone Wolves Reflected in Tooth Wear and Fracture
Blaire Van Valkenburgh
12 What Wolves Eat and Why Matthew C. Metz, Mark Hebblewhite, Douglas W. Smith, Daniel R. Stahler, Daniel R. MacNulty, Aimee Tallian, and John A. Vucetich
Box 12.1 Bison in Wood Buffalo National Park
L. N. Carbyn
13 Wolf Predation on Elk in a Multi-Prey Environment Matthew C. Metz, Douglas W. Smith, Daniel R. Stahler, Daniel R. MacNulty, and Mark Hebblewhite
 Box 13.1 Generalizing Wolf-Prey Dynamics across Systems: Yellowstone, Banff, and Isle Royale
Mark Hebblewhite
 Box 13.2 The Predator’s Perspective: Biomass of Prey
Matthew C. Metz

Box 13.3 Lessons from Denali National Park: Stability in Predator-Prey Dynamics Is a Pause on the Way to Somewhere Else
Layne Adams
14 Population Dynamics of Northern Yellowstone Elk after Wolf Reintroduction Daniel R. MacNulty, Daniel R. Stahler, Travis Wyman, Joel Ruprecht, Lacy M. Smith, Michel T. Kohl, and Douglas W. Smith
 Box 14.1 Wolves and Elk in the Madison Headwaters
Robert A. Garrott, P. J. White, Claire Gower, Matthew S. Becker, Shana Drimal, Ken L. Hamlin, and Fred G. R. Watson
 Box 14.2 Ecology of Fear
Daniel R. Stahler and Daniel R. MacNulty
Guest Essay: The Value of Yellowstone’s Wolves? The Power of Choice Michael K. Phillips
Part 5 Ecosystem Effects and Species Interactions

15 Indirect Effects of Carnivore Restoration on Vegetation Rolf O. Peterson, Robert L. Beschta, David J. Cooper, N. Thompson Hobbs, Danielle Bilyeu Johnston, Eric J. Larsen, Kristin N. Marshall, Luke E. Painter, William J. Ripple, Joshua R. Rose, Douglas W. Smith, and Evan C. Wolf
 Box 15.1 Long-Term Trends in Beaver, Moose, and Willow Status in the Southern Portion of the Absaroka-Beartooth Wilderness
Daniel B. Tyers
16 Competition and Coexistence among Yellowstone’s Meat Eaters Daniel R. Stahler, Christopher C. Wilmers, Aimee Tallian, Colby B. Anton, Matthew C. Metz, Toni K. Ruth, Douglas W. Smith, Kerry A. Gunther, and Daniel R. MacNulty
Guest Essay: Old Dogs Taught Old Lessons Paul C. Paquet
Part 6 Conservation, Management, and the Human Experience

17 Wolves and Humans in Yellowstone Douglas W. Smith, Daniel R. Stahler, Rick McIntyre, Erin E. Stahler, and Kira A. Cassidy
18 The Wolf Watchers Nathan Varley, Rick McIntyre, and James Halfpenny
 Box 18.1 Bob Landis’s Yellowstone Wolves Documentaries 000

Box 18.2 Seeing Wolves
Robert Hayes
19 Conservation and Management: A Way Forward Douglas W. Smith, P. J. White, Daniel R. Stahler, Rebecca J. Watters, Kira A. Cassidy, Adrian Wydeven, Jim Hammill, and David E. Hallac
Guest Essay: Making Better Sense of Wolves Susan G. Clark   Afterword
Rebecca J. Watters, Douglas W. Smith, Daniel R. Stahler, and Daniel R. MacNulty

Acknowledgments

Appendix: Species Names Used in the Text

Literature Cited

List of Contributors

Author Index

Subject Index

About the Author

Douglas W. Smith has studied wolves for more than forty years. In 1994 he was hired by the National Park Service in Yellowstone National Park as the project biologist to reintroduce wolves, and in 1997 he became the project leader, a position he still holds today. Besides wolves in Yellowstone, he is also responsible for supervising the park's bird, elk, and beaver programs. He is coauthor, most recently, of Wolves on the Hunt: The Behavior of Wolves Hunting Wild Prey, also published by the University of Chicago Press. Daniel R. Stahler is the Yellowstone Wolf Project's lead biologist and the project leader of the Yellowstone Cougar Project. In addition, he helps manage the elk program and is Yellowstone National Park's threatened and endangered species coordinator, working with species like lynx, wolverine, and grizzly bears. Daniel R. MacNulty is associate professor of wildlife ecology in the Department of Wildland Resources at Utah State University and was one of the first volunteers hired by the Yellowstone Wolf Project. He is also coauthor of Wolves on the Hunt.

Reviews

"Yellowstone Wolves summarizes over two decades of hard work, involving dozens of dedicated scientists and advocates, to bring these wolves back to Yellowstone. . . . Their voices are skillfully combined to tell the many-faceted narratives in this marvelous book. . . . The overall success of this long-term effort provides information that will be of inestimable value to other restoration projects, sharing methods that can help wolves and humans coexist in a changing world and an example of what can happen if people unite to give Mother Nature a chance."--Jane Goodall, from the foreword