Cover of Yellowstone Wolves
Yellowstone Wolves Science and Discovery in the World's First National Park
edited by Douglas W. Smith, Daniel R. Stahler and Daniel R. MacNulty, foreword by Jane Goodall
University of Chicago Press, 2020
Cloth: 978-0-226-72834-6 | Electronic: 978-0-226-72848-3
DOI: 10.7208/chicago/9780226728483.001.0001

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ABOUT THIS BOOKAUTHOR BIOGRAPHYREVIEWSTABLE OF CONTENTS

ABOUT THIS BOOK

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 behaviors, 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, this book 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 behaviors 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.

AUTHOR BIOGRAPHY

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

"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

“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 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

“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"

“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

"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

"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."
— Booklist

"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."
— Nature

"An informative, evocative compendium celebrating the famed predators. Wolf. The mere word evokes images that range from a mystical creature representing the genuine awe of the wild to a demon hell-bent on destroying livestock and livelihoods. The truth, according to Yellowstone Wolves, is one that science and reason—not myth—should discern. The past twenty-five years of studying the animal after a Herculean and successful reintroduction of wolves to Yellowstone provide bounties of both."
— Christopher Lancette, Washington Independent Review of Books

"How did wolves go from feared, hunted and decimated to a protected part of Yellowstone National Park's ecosystem? Yellowstone Wolves tells the fascinating story. . . . It's a comprehensive look at what happened when wolves were driven out of Yellowstone, and how nature is recovering now that they're back. . . . Part scientific tome, part rallying cry against humans' impulse to interfere with the natural world, Yellowstone Wolves is a powerful testament to what happens when 'people unite to give Mother Nature a chance,' as Jane Goodall puts it in her foreword to the book."
— Erin Blakemore, Washington Post

"The most complete book of which I am aware of all aspects of the behavior of majestic grey wolves. . . . The collection of essays, by a who's who of wolf researchers, is encyclopedic. It fully chronicles the epic journey of Yellowstone's wolves and the devoted researchers who have studied them there and in many other locations and is a must-read."
— Marc Bekoff, Psychology Today

"This handsome book may be coffee-table in size, but it is thoroughly scholarly in content. . . . Its coeditors could not represent greater specific expertise in the subject. . . . This splendid book is a major contribution to this vital, continuing debate."
— Geography Realm

“I had high expectations for this book and I wondered how it could ever be possible to cover and summarize the complexity of the project in a single book. Well, this book exceeded all my expectations and I strongly recommend it as one of the most important books on wolf biology ever written, not only for its content but also as superb example of an elegant and effective way of communicating science.”
— Luigi Boitani, Biological Conservation

"The combination of academic content, excellent photography, guest essays, and an online bonus documentary with interviews make this the go-to reference work for anyone wanting to go beyond the headlines on this reintroduction project. . . . By reintroducing wolves, scientists have had the unique opportunity to study the complexities of living, breathing ecosystems. Written by the very people who spent decades in the field doing the research, Yellowstone Wolves is a formidable achievement that presents a wide range of scientific topics in a well-organized, readable, and beautifully illustrated book."
— Leon Vlieger, Inquisitive Biologist

"The book’s contributors, who include many top wolf biologists, have compelling and inspiring stories to tell about conducting science in a special place. . . . A graphic in the afterword summarizes the activities of the Yellowstone Wolf Project from 1995 through 2018. I was impressed by the many scientific publications (85) and conference presentations (45) that resulted from the project, and my jaw dropped when I saw that 1,915 interviews were conducted and that another 4,305 formal presentations and 14,767 informal presentations were made. I suspect that much of the book’s narrative was developed before live audiences. I recommend this collection highly; it is full of important stories told well."
— American Scientist

"I highly recommend this book to wildlife biologists seeking an immersive reading experience, but specifically to those working in applied fields, as there are many lessons in wildlife management to be learned within this volume."
— Killian J. Murphy, Basic and Applied Ecology

"In this book, written by the biologists who made this possible, the reader finds not only historical and biological facts about wolves, but also a well-told story. This, together with a fluent narrative, makes it easily accessible to scientists from other fields."
— Conservation Biology

TABLE OF CONTENTS


DOI: 10.7208/chicago/9780226728483.003.0001
[wolf;Yellowstone National Park;history;western settlement;extirpation;ecosystem]
This chapter summarizes the history of wolves in Yellowstone National Park. The history of the park has frequently been written about, but these accounts differ in interpretation and rarely do they specifically discuss wolves. The park is often depicted as pristine, yet impacts due to Eur-American exploration and western settlement began almost a century before 1872 when the park was established. Wolf extirpation began prior to this time and changed many ecological relationships, most prominently predation on elk, well before anyone was able to document these impacts. Other carnivores were being simultaneously eliminated as well, changing the entire ecological community making Yellowstone very ‘unnatural’, contrary to popular perception. This historical omission, or overlooking of what carnivore extirpation did to the ecosystem and the much debated ‘naturalness’ it impacted – a long-held Park Service goal –and the cultural hurdles that had to be overcome to reintroduce wolves, some of which are still present in society, is the subject of this chapter.


DOI: 10.7208/chicago/9780226728483.003.0002
[wolf;Yellowstone National Park;reintroduction;recovery;Endangered Species Act]
This chapter explains the process and the challenges it took to reintroduce wolves to Yellowstone National Park with references to central Idaho. The recent history and status of wolves in the region, their natural immigration to northwest Montana, and how this led to a recovery process under the Endangered Species Act encompassing the entire region of Idaho, Montana, and Wyoming is explained. This includes the biological, political, cultural, and personal challenges of wolf reintroduction to those involved as well as public reaction. Many inside stories and previously unknown elements of the process are revealed with an emphasis on the uniqueness of the opportunity and the likelihood that such an ambitious project is unlikely to be repeated in the near or possibly distant future.


DOI: 10.7208/chicago/9780226728483.003.0003
[wolf;Yellowstone National Park;scientific research]
This guest essay by one of the leading wolf biologists in the world reflects on theunique opportunity for wolf research in Yellowstone National Park. The essay also summarizes key findings and debates in the scientific research of wolves, and offers a personal perspective on the important contribution wolf recovery to the Yellowstone ecosystem has made for conservation and science.


DOI: 10.7208/chicago/9780226728483.003.0004
[aging;body mass;development;generalist;senescence;reproduction;skeletal growth;life span]
The purpose of this chapter is to synthesize fundamentalfacts about the phenotype of wild wolves, and to highlight new phenotypic information revealed by Yellowstone wolves. The species is a quintessential generalist with traits that reflect a tradeoff between the ability to exploit a wide variety of foods – ranging from bison to berries, warm flesh to frozen bones – and the ability to use each one. A generalized skull and dentition together with a modest jaw musculature permit use of many different foods at the expense of maximal use of any one food. Wolves compensate for this tradeoff with a heightened olfactory sense and efficiencies in locomotion and digestion. Data from Yellowstone wolves indicates that this generalist feeding mode is associated with a relatively fast life history, including rapid development (skeletal growth ceases by 6-8 months; 80% of adult body mass acquired by 9 months), early first reproduction (mean age of primiparity = 2.7 years-old), high fecundity (4-5 pups/litter/year), short generation time (4-5 years), and short life span (median survival time = 6 years). Together, these phenotypic details indicate the wolf is a resilient species capable of overcoming substantial environmental adversity if given the opportunity to do so.


DOI: 10.7208/chicago/9780226728483.003.0005
[wolves;Yellowstone;cooperative breeding;social ecology;reproduction;kinship;breeding strategies]
This chapter discusses the social ecology of Yellowstone wolves, focusing on how wolf packs are structured in size, composition, and genetic relatedness and the variety of ways in which they form, operate, and dissolve. Additionally, this chapter discusses mating systems and breeding strategies, and the importance of cooperative breeding to wolf fitness. Factors influencing the social structure of packs are related to annual rates of pup survival, kinship, presence of multiple litters, adult survival, and dispersal. Ultimately, selective forces favoring particular attributes of packs and their members are related to cooperative breeding, territoriality, and food acquisition. Yellowstone research has made some key advances about wolf social ecology. These findings include the importance of group dispersal in a competitive landscape and its consequences for pack assembly, identification of factors driving diverse breeding strategies and their resulting influences on mating systems, and the nuanced effects of pack size and composition on female reproductive success.


DOI: 10.7208/chicago/9780226728483.003.0006
[aggression;territoriality;wolves;inter-pack behavior;conspecific strife]
Gray wolves are territorial and conflict between packs accounts for half of all adult wolf mortality. This conspecific strife varies with wolf population density and at high levels can cause a decrease in adult survival. Inter-pack aggression is also affected by the amount of available prey as prey presence and abundance can influence resource competition between packs. In addition, the success or failure of fights between packs can have long-term consequences on pack survival and resource-holding potential. These fights are impacted by pack and individual wolf characteristics where larger packs with certain valuable pack members—those with experience or large body size—are more likely to regularly win during conflicts. This chapter describes nearly 25 years of observations of aggressive inter-pack behavior and explains scientific results related to territoriality. In addition, this chapter discusses how intense inter-pack aggression and intra-pack sociality have shaped the gray wolf’s evolutionary life history related to group-living, defense of young and other resources, cooperation, and the value of aging.


DOI: 10.7208/chicago/9780226728483.003.0007
[wolf;Yellowstone;population dynamics;population growth rate;vital rates;survival]
This chapter reviews what is known about what causes wolf populations to fluctuate, emphasizing that previous research concluded that wolf population size (density) is usually determined by available prey. The underlying goal of this chapter is to confirm this conclusion, but that this long-held belief stymied further research prolonging discovery of the mechanism that adjusts wolf density to prey availability (biomass). Through a detailed discussion of wolf vital rates and explanation of a model constructed for Yellowstone, the authors conclude that adult survival is most likely the most important vital rate driving changes in wolf population growth rate. These findings likely will not resolve the debate, but hopefully invigorate it so a better understanding of what drives wolf population dynamics may someday inform management decisions about this controversial carnivore.


DOI: 10.7208/chicago/9780226728483.003.0008
[wolf;Yellowstone;word cloud;text mining;science;predation;elk]
This guest essay applies word cloudand text mining analytical tools to evaluate and visualize the shift in science publications on wolves through time. Prior to Yellowstone wolf reintroduction, science was primarily focused on elk, grizzly bears, and forest fires. Following wolf recovery, the focus shifted to wolf impacts on elk and predation.


DOI: 10.7208/chicago/9780226728483.003.0009
[genetics;pedigrees;molecular ecology;evolution;connectivity]
This chapter provides an overview of the role genetics has played in the story of Yellowstone wolves and their contribution to an ever-advancing field of molecular research. Yellowstone has established a genetic baseline for wolves using a wide variety of genetic markers, along with establishment of a population pedigree of nearly 400 wolves traced back to the founding individuals. These genetic tools have been used to evaluate the genetic health of Yellowstone wolves and connectivity with wolves across the Northern Rockies. The integration of genetic, pedigrees, demographic, behavioral, and physiological data has helped identify the genetic underpinnings of behavior and physiological traits. Yellowstone wolves have contributed to a detailed map of the North American wolf genome, which has been used to measure mutation rates of individual genes that influence phenotype and adaptation. We now understand the basic factors influencing gene expression and regulation in a wild population that can limit responses to future stressors in the environment. Yellowstone wolves have been at the frontier of genetic research in wild canids and have contributed to a greater understanding of molecular ecology and evolution in general.


DOI: 10.7208/chicago/9780226728483.003.0010
[coat color;melanism;K locus;fitness;wolf;Yellowstone]
This chapter is about the fascinating story of coat color variation in wolves. Specifically, it chronicles the discovery of the origin and persistence of the K locus gene that leads to black pelage in wild wolves. The legacy of black coat color (i.e. melanism)is linked to domestic dogs and human-dog-wolf interaction thousands of years ago. Research in Yellowstone has demonstrated important links to demographic, behavioral, health, and ecological processes. The K locus has become an important scientific case study because it demonstrates that variants that appear under domestication can be viable in the wild and adaptive in natural populations. Additionally, the ability to identify the genetic architecture of coat color variation has helped to determine the mechanisms of selection and maintenance of genetic variation in nature. This story reveals fitness implications and trade-offs involving reproduction, survival, health, aggression, and mate choice in a social carnivore. This story not only highlights the contribution of Yellowstone wolves to science, but also reveals how carefully planned, long-term, and interdisciplinary scientific research can sometimes result in serendipitous discoveries.


DOI: 10.7208/chicago/9780226728483.003.0011
[wolves;Yellowstone National Park;disease;parasites;canine distemper;infectious diseases]
Wolves in Yellowstone National Park are hosts to numerous viruses, bacteria, and parasites. These pathogens have been monitored for two decades, and their prevalences and effects are quantified and described within this chapter. Direct observation of wolves, as well as the testing of biological samples, has allowed us to detect which pathogens the wolves were exposed to and when. For instance, we identified canine distemper outbreaks in 1999, 2005, and 2008 that killed most pups born in those years. Because Yellowstone wolves are highly visible, some infections were tracked by symptom expression over time (e.g., sarcoptic mange). Packmates interact much more frequently than individuals from different packs, which has consequences for how pathogens and parasites spread. In general, wolves that live in Yellowstone’s interior or in larger packs tend to survive their infections at higher rates, as do wolves with previous exposure and immunity. Studying infectious diseases in Yellowstone wolves has led to many discoveries about the effects of pathogens and parasites in an apex predator.


DOI: 10.7208/chicago/9780226728483.003.0012
[Yellowstone National Park;wolves;genetics;Scandinavia;ecosystem impacts]
This guest essay provides a perspective on the importance of wolf recovery to Yellowstone National Park from a European biologist studying wolves in Scandinavia. The author draws on four main contributions Yellowstone has made to biology. The first is the opportunity to study ecosystem impacts like trophic cascades. The second is the unique opportunity to observe wolves in an open landscape and better understand behavior. Third, Yellowstone has provided the opportunity to study predator-prey dynamics in great detail in a multi-carnivore, multi-prey landscape. Finally, Yellowstone offers the ability to study conservation genetics in a reintroduced population of carnivores.


DOI: 10.7208/chicago/9780226728483.003.0013
[wolf;Yellowstone National Park;research;field work]
This chapter explains how the wolf-prey research in Yellowstone National Park developed and is conducted. This includes a short history of wolf research and how the methods have changed through time and become more sophisticated, and how the Wolf Project adopted them and integrated their use into studying Yellowstone wolves. Besides a detailed discussion of study methods, the challenges and logic behind what was done and how is explained. The chapter concludes by challenging researchers to use a combination of field work and technology rather than abandoning older methods in favor of ever-improving and more convenient technology.


DOI: 10.7208/chicago/9780226728483.003.0014
[aging;body size;cooperation;group size;hunting behavior;wolves]
Fueling many debates about wolves is the assumption that wolves are outstanding hunters of hoofed mammals (ungulates). This chapter explains why this assumption is false. Numerous studies have shown that wolves more often fail than succeed in their attempts to kill ungulates. Whereas traditional explanations for the low success rate of wolves have mainly focused on the role of prey traits such as age, body size, and physical condition, studies of individually identifiable wolves in Yellowstone National Park have revealed how the wolf’s own traits curb its ability to kill. Although the wolf has the bite force, body size, and cooperative behavior to kill a wide variety of ungulates, it lacks the massive size, retractable claws, supinating muscular forelimbs, and specialized skull configuration that would allow it to be a consistently high-success hunter of any one particular prey species. A wolf’s hunting ability is further limited by old age, small body size, and the “free-riding” behavior of its hunting partners. In sum, the wolf’s own biology enforces strict limits on its capacity to kill ungulates. Recognizing these limits may help resolve debates predicated on exaggerated notions of wolf hunting ability.


DOI: 10.7208/chicago/9780226728483.003.0015
[wolf;prey selection;elk;bison;Yellowstone;diet;prey vulnerability]
This chapter discusses seasonal ungulate use by wolves throughout Yellowstone National Park (YNP), and also in more detail for northern YNP, over a 22-year period (1995 – 2017). Wolves throughout YNP primarily fed on elk during summer, following the return of migratory elk. However, wolf diet was more varied across YNP during winter. Specific to northern YNP, wolves primarily used elk throughout the year. Wolves frequently killed young elk during summer and early winter, and then adult elk the rest of the year. Deer and bison were occasionally important prey for wolves. For example, wolves increasingly used bison, primarily through scavenging, over time during winter. This increased use of bison was concurrent to a general increase in northern YNP bison, and decline in elkabundance. Wolves continued to strongly select elk, however, when killing their prey. These patterns in wolf diet were driven by the relative availability and prey vulnerability.


DOI: 10.7208/chicago/9780226728483.003.0016
[wolf;Yellowstone National Park;predation;functional response;numeric response;kill rate;predation rate]
This chapter discusses how wolf abundance and predation in the multi-prey system of northern Yellowstone National Park (YNP) have changed over a 22-year period (1995 – 2017). Wolves in northern YNP have primarily preyed on elk, and wolf abundance has displayed a positive numeric response to elk abundance. The functional response (i.e., kill rate in relation to prey abundance) indicated that elk abundance positively affected the rate that wolves killed elk during winter, although the precise shape of the functional response was difficult to discern. Increased use of alternative prey (e.g., bison) by wolves also negatively affected wolf kill rate on elk. Finally, the combined effects of wolf kill rate, wolf abundance, and elk abundance indicated that the percent of the northern Yellowstone elk population killed by wolves (i.e., wolf predation rate) has varied considerably across time, but may also be density-dependent, thereby favoring wolf predation as a stabilizing force.


DOI: 10.7208/chicago/9780226728483.003.0017
[northern Yellowstone;predation;hunter offtake;wolf;population dynamics;cougars;bears]
Changes in the size of the northern Yellowstone elk herd have been a source of controversy throughout the history of Yellowstone National Park. This chapter reviews the influence of wolves on the dynamics of the northern elk herd in the years after the 1995-1997 wolf reintroduction. A population model indicates that the size of the northern herd decreased from about 19,000 elk in 1994 to about 4000 elk in 2013. Several lines of evidence indicate that wolves neither triggered this decline nor acted as its primary driver once it started. This includes the fact that the decline started the year before the initial release of wolves in March 1995. In addition, the annual proportion of the northern herd killed by hunters outside the park was about 2-8 times greater than that killed by wolves for seven consecutive years after the initial wolf release. Subsequent decreases in hunter offtake increased the relative influence of wolves; however, other factors also contributed to the elk population decline including drought and predation by bears and cougars. Increasing herd size since 2013 provides further evidence that wolves have had a generally moderate influence on the dynamics of the northern Yellowstone elk herd.


DOI: 10.7208/chicago/9780226728483.003.0018
[Yellowstone;wolf;reintroduction]
This guest essay written by the first project leader of the Yellowstone Wolf Project reflects on the a key aspect of wolf reintroduction: the power of choice. The essay describes a variety of ways people relate to nature and how the decision to bring back a controversial species like the wolf is a catalyst to saving nature as a whole.


DOI: 10.7208/chicago/9780226728483.003.0019
[trophic cascades;food webs;Yellowstone;browsing intensity;ecosystem change;northern range]
Trophic cascades are indirect species interactions that originate with a predator and spread down-ward through food webs.In some areas of Yellowstone, such indirect effects on vegetation seem dramatic, while in others, little effect is observed. This chapter explains the science underlying the observed variation in plant recovery among species and locations across Yellowstone’s northern range. Assessmentsofnorthernrangevegetationoverthe last two decades indicate the emergence of three generalpatternsofecosystem change:(1)youngplants of deciduous woody species have grown taller in some areas; (2) changes in plant height have varied both temporally and spatially because of several factors, including variation in browsing intensity and in groundwater availability; and (3) in some areas, reductions in elk browsing have been counteracted by increases in bison use, causing continued or renewed suppression of woody plants.


DOI: 10.7208/chicago/9780226728483.003.0020
[Yellowstone;scavengers;food web;competition;bears;cougars]
This chapter synthesizes the wolf ’s relationships with other large carnivores and scavengers in Yellowstone. Specifically, it describes how the scavenger community, which includes ravens, eagles, magpies, coyotes, foxes, and bears, responded to the return of wolves from a behavioral and a food acquisition perspective. Second, given that wolf restoration was simultaneous with the natural recovery of grizzly bears and cougars, the nature of competition among these large carnivores is described, which includes elements of both direct and indirect competition. and behavioral interactions. Evaluating the role of interactions between Yellowstone’s large carnivores and scavengers in food web dynamics provides a fresh perspective on the structure and function of natural communities. Arguably, the ecosystem effects of wolf recovery in the presence of other meat eaters reveal one of the most significant, but frequently underappreciated, aspects of Yellowstone’s food web dynamics.


DOI: 10.7208/chicago/9780226728483.003.0021
[wolf;Yellowstone National Park;research;wolf management;conservation]
This guest essay by a prominent Canadian wolf biologistdiscusses how the ecological research that followed wolf reintroduction to Yellowstone National Park has revealed new discoveries about wolves. From revealing nuanced cultural patterns in wolf social dynamics, to clarifying the influence of wolves on ecological communities, Yellowstone research provides a fresh perspective on wolves living in a natural system largely unaffected by humans. The essay offers a philosophical discussion about the ethics of wolf management and the relationship humans have with wolves, conservation, and nature


DOI: 10.7208/chicago/9780226728483.003.0022
[wolf;Yellowstone National Park;management;human disturbance;habituation]
This chapter explains the park philosophy and regulations that guide wolf-human interactions which are designed to protect wolves, yet provide for visitor enjoyment. Yellowstone National Park objectives for wolf management, then are as follows:(1)protectdenand rendezvoussitesfromhuman disturbance, (2) prevent fearless wolf behavior—commonly called habituation, (3) educate park visitors on proper and safe behavior in the presence of wolves, including protective measures in the rare eventofawolf approach,and (4) regulatehumanviewing so as not to disturb “natural” wolf behavior. The park’s first step toward these goals is education and regulation.


DOI: 10.7208/chicago/9780226728483.003.0023
[wolf;Yellowstone;economic benefits;wildlife conservation;visitor enjoyment]
The success of wolf recovery to Yellowstone, coupled with the popularity of the wolves themselves, is a conservation achievement. Not only have these accomplishments led to increased public awareness of wildlife conservation issues and ecological relationships, but they have also informed wildlife restoration efforts elsewhere. This chapter describes the visitor enjoyment phenomena that revolves around watching wild wolves in Yellowstone. From this world-renowned opportunity to view wolves and their interaction with other species, the chapter describes the education and outreach facilitated by wolf observations throughout the park, the economic benefits, and even describes some of the challenges that comes with the public’s passion for Yellowstone wolves.


DOI: 10.7208/chicago/9780226728483.003.0024
[wolf restoration;wildlife management;large carnivores;human attitudes;wolf hunters]
The purpose of this chapter is to examine and recommend a holistic approach to wolf management across jurisdictional boundaries. This is a difficult issue, as management authority resides with multiple agencies with differing mandates and constituencies.Wolves are just one of the more recent examples of this issue. Management of large carnivores has advanced from an era in which they were actively limited to low numbers to one in which they are increasingly tolerated and valued as part of wild ecosystems. Thus, there is increasing pressure on wildlife agencies to balance the desires of wolf hunters and wolf non-hunters. Wolf restoration occurred, in part, because human attitudes changed and people supported it, and the prospect of hunting was a factor in this acceptance.Balancing this dynamic is a key part of wildlife management moving forward in the 21st century and wolves are at the center of this societal issue.


DOI: 10.7208/chicago/9780226728483.003.0025
[Yellowstone;wolves;problem solving]
This closing essay reflects on the challenges humans have in making meaning of the wolf in the greater Yellowstone ecosystem. Applying principals of social science, the essay discusses the different social, political, personal, and commercial constructs humans apply to their views of the wolf. The essay closes with a call for better problem solving and attention to the complexity of our consciousness and how we can better live with wolves.

Book published by University of Chicago Press, 2020. Page maintained by BiblioVault.