In 2020, it will have been twenty-five years since one of the greatest wildlife conservation and restoration achievements of the twentieth century took place: the reintroduction of wolves to the world's first national park, Yellowstone. Eradicated after the park was established, then absent for seventy years, these iconic carnivores returned to Yellowstone in 1995 when the US government reversed its century-old policy of extermination and – despite some political and cultural opposition – began the reintroduction of forty-one wild wolves from Canada and northwest Montana. In the intervening decades, scientists have studied their myriad behaviours, from predation to mating to wolf-pup play, building a one-of-a-kind field study that has both allowed us to witness how the arrival of top predators can change an entire ecosystem and provided a critical window into impacts on prey, pack composition, and much else.
Here, for the first time in a single book, is the incredible story of the wolves' return to Yellowstone National Park as told by the very people responsible for their reintroduction, study, and management. Anchored in what we have learned from Yellowstone, highlighting the unique blend of research techniques that have given us this knowledge, and addressing the major issues that wolves still face today, Yellowstone Wolves is as wide-ranging and awe-inspiring as the Yellowstone restoration effort itself. We learn about individual wolves, population dynamics, wolf-prey relationships, genetics, disease, management and policy, newly studied behaviours and interactions with other species, and the rippling ecosystem effects wolves have had on Yellowstone's wild and rare landscape. Perhaps most importantly of all, the book also offers solutions to ongoing controversies and debates.
Featuring a foreword by Jane Goodall, beautiful images, a companion online documentary by celebrated filmmaker Bob Landis, and contributions from more than seventy wolf and wildlife conservation luminaries from Yellowstone and around the world, Yellowstone Wolves is a gripping, accessible celebration of the extraordinary Yellowstone Wolf Project – and of the park through which these majestic and important creatures once again roam.
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
Appendix: Species Names Used in the Text
List of Contributors
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 co-author most recently of Decade of the Wolf: Returning the Wild to Yellowstone.
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.
"A gathering of scientific essays on the natural history of the reintroduced wolf population in Yellowstone National Park. Wolves were long a key predator in the Greater Yellowstone Ecosystem before they were extirpated. In 1995, they were reintroduced in what the editors characterize as 'a change of heart.' That change of heart is 'the single most important fact of wolf recovery everywhere.' [...] A welcome contribution to the conservation-biology literature on wolves."
– Kirkus 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
"Smith and his colleagues have graced us with the most comprehensive and elegant book ever written about the Yellowstone wolves. They take us deep into the human dimensions of wolf recovery while also giving us exceptional insights about social behavior not only for wolves, but for animal societies in general."
– Mark W. Moffett, author of The Human Swarm: How Our Societies Arise, Thrive, and Fall
"I can't praise this book enough. From the foreword by Jane Goodall to David Mech at the end, to all the now nearly lifetime work of researchers dedicated to this grand ecological experiment, important conservation effort, and inspirational project, Yellowstone Wolves is a model of success that will not only endure, but grow and sprout hope and inspiration globally. Bravo!"
– Bernd Heinrich, author of Mind of the Raven, The Homing Instinct, and White Feathers
"Wolves are surrounded by conflict and controversy, the substance of which has been dimmed by a veil of scientific uncertainty. Yellowstone Wolves is a readable and authoritative account about this iconic species and magical place, written by the scientists, managers, and conservationists who did the work. Read this book and learn what these on-the-ground professionals really know and think about wolves in the Yellowstone ecosystem."
– James A. Estes, author of Serendipity: An Ecologist's Quest to Understand Nature
"I consider the restoration of the gray wolf into the northwestern United States to be the single most important event in the past fifty years of wildlife conservation and management. This volume will serve as a major reference for years as it covers the key social/ecological/behavioral/management issues related to the wolf in Yellowstone National Park. Virtually all of the principal individuals who have been involved in wolf investigations and management across the continent have contributed."
– James M. Peek, professor emeritus, Department of Fish and Wildlife Sciences, University of Idaho
"Very informative. Every reader of this book will learn something new and interesting about wolves, their ecology, or how they compare to other species in the animal world. It will be a great resource, sought out as the de facto authority on wolf knowledge and discovery for Yellowstone most certainly, but also on how the knowledge gained lends itself to understanding wolves elsewhere. Compelling."
– Dean Cluff, Department of Environment and Natural Resources, Government of the Northwest Territories, Canada
"Yellowstone Wolves is a Herculean attempt to synthesize research on Yellowstone National Park subsequent to wolf recovery. It is likely to be touted as the definitive work on the subject for years. This is a necessary, wonderful book."
– Mark Boyce, Department of Biological Sciences, University of Alberta, Canada
"Reintroducing the wolf to Yellowstone is arguably the world's greatest wildlife experiment. The wolves' progress has been documented meticulously by a team of hiking, driving and flying biologists and passionate volunteers – so much so that no wolf study comes close to yielding its abundance of information. The research generated has been distilled into a new book, Yellowstone Wolves, assembled by three of the biologists who studied the wolves' return."
– New York Times
"The huge volume of information presented in this book, plus illustrations, diagrams, and an extensive bibliography, underpin its significance to ecological research as do the many contributors; several guest essays add useful overviews. It is a fitting tribute to the 100 years plus of research pre and post wolf reintroduction, and a sound basis for future studies."
– The Biologist
"An intimate look at these wild canines and their ways. Humans have always felt more about wolves than we've actually known about them. Their emotions seem similar to our own, but they inspire dark fairy tales and anger among some livestock owners. By 1960, the wild canines were nearly extinct in the lower forty-eight, but in 1995, scientists plucked forty-one wolves from Canada and Montana and released them into the wilds of Yellowstone National Park. Since then they've thrived, splitting into myriad packs and offering researchers an unparalleled opportunity to witness their secrets. Now, twenty-five years of intensive research has been distilled into a fascinating new book, Yellowstone Wolves."
– American Way
"Rich with photographs, charts, and maps, this volume provides a wealth of information for those studying the project, ecology, rewilding, and wolves in general, and it also brings the complex workings of the natural world of Yellowstone into clear focus [...] A comprehensive resource well-suited for natural science collections."
"This year marks the twenty-fifth anniversary of one of the largest and most successful conservation efforts in modern history: the reintroduction of wolves into Yellowstone National Park. This gorgeous book explains why wolves were eradicated from the park when it opened nearly seventy years ago, and how their reintroduction fundamentally changed the park's ecosystems for the better. Edited by the people responsible for studying and managing the wolf reintroduction project – and with a forward by Jane Goodall – Yellowstone Wolves is a must-read for anyone interested in wildlife or conservation. The text is accompanied by stunning, full-color photographs and an online documentary by Bob Landis, perhaps best known for his work with National Geographic."
– Amy Brady, LitHub
"Twenty-five years ago, the authors reintroduced wolves to Yellowstone National Park in Wyoming – the first deliberate return of an apex carnivore to a big ecosystem. Here, they relate what they've learnt of the animals' predation, mating, play, genetics, disease and more, and their impact on other species and the landscape. Also detailed are the fraught history, politics and implications of rewilding. Glorious pictures bear witness to fragile gains. US President Donald Trump's silver-anniversary gift? Rolling back protections on the wolves."