Wolves: Behavior, Ecology, Conservationby L. David Mech (Editor), Luigi Boitani (Editor), David L. Mech (Editor), Luigi Boitani
Wolves are some of the world's most charismatic and controversial animals, capturing the imaginations of their friends and foes alike. Highly intelligent and adaptable, they hunt and play together in close-knit packs, sometimes roaming over hundreds of square miles in search of food. Once teetering on the brink of extinction across much of the United States and
Wolves are some of the world's most charismatic and controversial animals, capturing the imaginations of their friends and foes alike. Highly intelligent and adaptable, they hunt and play together in close-knit packs, sometimes roaming over hundreds of square miles in search of food. Once teetering on the brink of extinction across much of the United States and Europe, wolves have made a tremendous comeback in recent years, thanks to legal protection, changing human attitudes, and efforts to reintroduce them to suitable habitats in North America.
As wolf populations have rebounded, scientific studies of them have also flourished. But there hasn't been a systematic, comprehensive overview of wolf biology since 1970. In Wolves, many of the world's leading wolf experts provide state-of-the-art coverage of just about everything you could want to know about these fascinating creatures. Individual chapters cover wolf social ecology, behavior, communication, feeding habits and hunting techniques, population dynamics, physiology and pathology, molecular genetics, evolution and taxonomy, interactions with nonhuman animals such as bears and coyotes, reintroduction, interactions with humans, and conservation and recovery efforts. The book discusses both gray and red wolves in detail and includes information about wolves around the world, from the United States and Canada to Italy, Romania, Saudi Arabia, Israel, India, and Mongolia. Wolves is also extensively illustrated with black and white photos, line drawings, maps, and fifty color plates.
Unrivalled in scope and comprehensiveness, Wolves will become the definitive resource on these extraordinary animals for scientists and amateurs alike.
“An excellent compilation of current knowledge, with contributions from all the main players in wolf research. . . . It is designed for a wide readership, and certainly the language and style will appeal to both scientists and lucophiles alike. . . . This is an excellent summary of current knowledge and will remain the standard reference work for a long time to come.”—Stephen Harris, New Scientist
“This is the place to find almost any fact you want about wolves.”—Stephen Mills, BBC Wildlife Magazine
"Everything you wanted to know about wolves!"
Mark S. Boyce
John B. Theberge
Astrid Vik Stronen
"This book is the first comprehensive review of wolf biology since Mech's original work. It provides an excellent compilation of current knowledge, with contributions from all the main players in wolf research over the past few decades who are still alive. It is designed for a wide readership, and certainly the language and style will appeal to both scientists and lucophiles alike. . . . This is an excellent summary of current knowledge and will remain the standard reference work for a long time to come. And by today's standards, it's refreshingly cheap."
"Everything you wanted to know about wolves!"
"There is no single source that compiles more information about wolves, and this compendium will stimulate a new generation of field biologists to design studies to fill the many gaping holes in our understanding. . . . For its price, this is a bargain of a book."
"One of the best books ever published about any wild carnivore species. Its 13 chapters cover a great swathe of topics—social ecology, behaviour, reproduciton, communication, feeding ecology, predator-prey relationships, population dynamics, physiology, molecular genetics, evolution and taxonomy, interactions with non-prey species and with humans, and conservation."
“This book, edited by two outstanding wolf biologists, is the first definitive work on the species since Mech’s monograph in 1970. Its expert contributors deal with all aspects of wolf biology—social life, hunting, prowess, physiology, genetics, worldwide distribution, and relations with humans—and this is the place to find almost any fact you want about wolves.” Stephen Mills, BBC Wildlife Magazine
"Wolves provides an outstanding overview of the biology and conservation of the species, delivering an up-to-date and unparalleled account of grey wolf biology. . . . This is undoubtedly the most important book ever published on grey wolves. A heft volume, beautifully published, with many black and white photographs and quality colour plates, at a reasonable price. . . . A must-have reference book for the carnivore biologist, boreal ecologist, landscape conservationist, behavioural ecologist, student of people-wildlife conflicts, and countless wolf lovers worldwide.”
"[The book] provides one of the most extensive descriptions of the ecology of any mammal species in the world. . . . [It] brings together research conclusions on any conceivable topic related to wolves, very useful either for the interested person or the biologist."
"The book is a highly valuable addition to the bookshelf of anyone who would like to know more about why living with wolves will continue to challenge us and provide perspective on our own ecosystem role."
- University of Chicago Press
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Read an Excerpt
Behavior, Ecology, and Conservation
University of Chicago Press
Copyright © 2003
University of Chicago
All right reserved.
L. David Mech and Luigi Boitani
The wolf is truly a special animal. As the most widely distributed of all
land mammals, the wolf, formally the gray wolf (Canis lupus), is also one
of the most adaptable. It inhabits all the vegetation types of the
Northern Hemisphere and preys on all the large mammals living there. It
also feeds on all the other animals in its environment, scavenges, and can
even eat fruits and berries. Wolves frequent forests and prairies, tundra,
barren ground, mountains, deserts, and swamps. Some wolves even visit
large cities, and, of course, the wolf's domesticated version, the dog,
thrives in urban environments.
Such a ubiquitous creature must, as a species, be able to tolerate a wide
range of environmental conditions, such as temperatures from -56° to +50°C
(-70° to +120°F). To capture its food in the variety of habitats,
topographies, and climates it frequents, the wolf must be able to run,
climb, lope, and swim, and it performs all these functions well. It can
travel more than 72 km (43 mi)/day, run at 56-64 km (34-38 mi)/hr, and
swim as far as 13 km (8 mi) (P. C. Paquet, personal communication), no
doubt aidedby the webs between its toes.
The wolf leads a feast-or-famine existence, gorging on as much as 10 kg
(22 pounds) of food at a time, but able to fast for months if necessary.
Nevertheless, if all goes well (which for most wolves it does not), wolves
can live 13 years or more in the wild (Mech 1988b) and up to 17 years in
captivity (E. Klinghammer and P. A. Goodman, personal communication).
As might be expected, a widely distributed animal like the wolf varies
physically all around its circumpolar home. The desert-inhabiting variety
of Israel can weigh as little as 13 kg (29 pounds), whereas its northern
tundra cousin can reach over 78 kg (172 pounds). The wolf's color varies
across the entire black-white spectrum, with most wolves tending to be a
mottled gray (Gipson et al. 2002).
Wolves live in packs of up to forty-two, but can survive even as lone
individuals. Although wolf packs are usually territorial, where necessary,
they can migrate hundreds of kilometers between where they raise their
pups and where they take those pups in winter to follow their prey. The
great variation in the wolf's environment, and in the creature's behavior
and ecology as it contends with that environment, makes generalizing
difficult. This problem can lead to false generalizations and
misunderstanding about the animal.
Although wolves have provoked human beings by sharing their domestic and
wild prey without permission, these predators have withstood humankind's
resulting assault on them everywhere except where the ultimate weapon,
poison, has been used. Given half a chance, wolves have responded by
repopulating suitable areas with remarkable success. In the process,
wolves have gained the support of many humans and have become a
conservation challenge-one that is rapidly being met. This animal that
sits on its haunches at the top of the food chain has become a symbol of
the wilderness, an icon to environmentalists, and a poster child for
endangered species recovery efforts.
Because of the strong feelings that both wolf haters and wolf advocates
hold, it has been hard to sell the truth about the wolf-folks of each
viewpoint resist accepting information they believe supports views
opposite their own. Yet we must at least present the information as best
we can. In this respect, we have tried throughout this book to draw valid
generalizations where possible while indicating where lack of information
In the following pages, we and the chapter authors have assembled and
synthesized the considerable amount of information available about this
fascinating animal. This book, however, is not a collection of all works
on the wolf, but rather a compendium of basic information. To have
included all of the worthy published material about the wolf, or even to
have referenced it, would have been too much for writers and readers
We start by discussing in chapter 1 the wolf's basic social ecology; that
is, its pack structure and spacing, natural history, and movements. Why do
wolves live in packs? The answer is not as simple as once thought. What
triggers dispersal? How do wolves defend their territories? These and many
other questions are examined and discussed.
Chapter 2 focuses in more intimately on the wolf and dissects the details
of individual behavior, courtship and reproduction, parental care, pack
social dynamics, competition, aggression, rivalry, leadership, and rank
order. How important and pervasive is the wolf dominance hierarchy for
which wolves are so famous in the popular literature? This chapter
explores this and many other questions in depth.
Focusing even more tightly, chapter 3 discusses wolf communication. Here
wolf howling and other vocalizations, scent marking, body postures, and
other significant signals are covered in detail. The composition of wolf
urine, the influence of hormones on communication, the nature of wolf
hearing, seeing, smelling, and tasting, and the role of wolf senses in the
animal's dealings with its environment constitute much of the chapter.
Once these basics are dealt with, we turn to the behavior that has brought
the wolf so much infamy-its food habits-in chapters 4 and 5. Chapter 4
approaches the subject with an overview of the wolf's predatory
adaptations, its digestive system, general feeding habits, specific foods,
and basic hunting behavior. The wolf is shown to be superbly adapted to
its carnivorous lifestyle, from its forty-two teeth and massive jaw
muscles to its large stomach capacity and thorough digestion.
Picking up where chapter 4 leaves off, chapter 5 examines the challenges
wolves face in trying to procure their prey. With quarry ranging from the
fleet and alert white-tailed deer to the massive moose, muskox, and bison,
the wolf must find, catch, and kill regularly. How does it go about
overcoming all the many antipredatory adaptations these animals have
evolved? The many details of this vast subject constitute a large part of
this chapter. Just as important, however, is the chapter's other main
topic-assessing the influences of wolf predation on prey populations. Lack
of agreement characterizes several aspects of this topic.
Less controversial, but still a dominant aspect of wolf biology, behavior,
and conservation, is the subject of chapter 6: wolf population dynamics.
Wolf productivity, density, survival, mortality, population change, and
population regulation are all discussed and analyzed. Relationships
between food supply (prey biomass) and several demographic and ecological
factors are explored, and the pervasive effect of food supply becomes
After all the above extensive and external topics are covered, chapter 7
turns to look inside the wolf. "The Internal Wolf: Physiology, Pathology,
and Pharmacology" first examines hormonal aspects of wolf reproduction,
then development, basal metabolic rate, and nutrition. Then it details
internal threats to the wolf's health, including parasites and diseases of
all sorts. Because wolf research and health management of captive wolves
are becoming increasingly important, the chapter also includes a
discussion of drugs used for anesthesia, capture, and treatment of wolves.
Also delving deeply into the wolf is chapter 8. Featuring the relatively
new but highly enlightening techniques of molecular genetics, this chapter
presents some novel perspectives on wolves. Through modern biochemical
analyses, wolf DNA can be parsed, and various inferences can be drawn from
the results. Some of these conclusions challenge existing ideas: was the
dog domesticated from the wolf about 10,000-15,000 years ago, as has long
been thought? Or much longer ago, as molecular analyses suggest?
While chapter 8 uses new technology to get at questions of wolf taxonomy
and evolution, chapter 9 uses long tried and tested methodology to examine
many of the same issues. Fossil evidence and skull measurements are used
to trace the long path of evolution the wolf's ancestors followed, the
relationships among wolves of different geographic regions, and the
possible hybridization of closely related forms. Nicely complementing
chapter 8, this chapter provides one more important perspective on these
subjects. We hope that interested readers will examine chapters 8 and 9
closely and judge for themselves where the weight of evidence falls in any
of the areas where their conclusions differ. The conclusions of both
chapters depend greatly on inference, and in many cases the data base for
the inferences is necessarily more meager than science would like.
Chapter 10 deals with the many interactions wolves have with prominent
animals in their environment other than their regular prey. Species such
as bears, tigers, and cougars not only compete with wolves for prey, but
also sometimes kill wolves or are killed by them. Other smaller creatures,
such as coyotes, foxes, ravens, and eagles, scavenge from wolf kills and
can consume large amounts of the wolf's hard-earned bounty. Chapter 10
outlines these interactions that form such an important part of the wolf's
Chapter 11 differs from the others in that it does not cover a specific
aspect of the wolf, but rather is devoted to describing the recovery of
the red wolf (Canis rufus). We devote a whole chapter to this subject,
while covering other wolf reintroductions as parts of other chapters,
because the red wolf is a special case. The red wolf is the only
long-recognized species of wolf other than the gray wolf, and scientists
disagree on its taxonomic identity. Is the red wolf really a separate
species, or might it be a subspecies of the gray wolf? Some scientists
claim that the red wolf is a hybrid between the gray wolf and the coyote,
while others dispute that conclusion. All workers agree, however, that the
red wolf is endangered. Thus, after several years of raising remnant
members of its population in captivity, the United States Fish and
Wildlife Service reintroduced the red wolf into part of its former range.
Chapter 11 follows the progress of that historic endeavor and summarizes
the lessons learned from it.
That topic leads nicely into the larger subject of chapter 12, the wolf's
interactions with human beings. It is that very subject that makes this
book possible and necessary. The chapter tracks human attitudes toward
wolves through history and many various cultures. It documents the effects
of wolves on human activities and vice versa, including wolf attacks on
human beings and human extermination of entire wolf populations.
Concluding the book after the thought-provoking chapter 12 is our chapter
on conservation of the wolf. Although human beings were responsible for
the demise of the wolf throughout much of western Europe and most of the
contiguous United States, the human species suddenly realized its mistake
and began trying to conserve the wolf and restore it to parts of its
former range. Chapter 13 details these efforts and projects an optimistic
future for the wolf-a fitting conclusion to this book.
Down through the ages, the wolf has never had a neutral relationship with
humanity. It has either been hated, despised, and persecuted or revered,
respected, and protected. It has been, and continues to be, a subject of
myth and legend, folklore and fairy tale. This book is meant to temper
those misrepresentations by presenting a scientific view of the animal-one
that we think is even more interesting, and certainly more accurate.
Excerpted from Wolves
Copyright © 2003
by University of Chicago.
Excerpted by permission.
All rights reserved. No part of this excerpt may be reproduced or reprinted without permission in writing from the publisher.
Excerpts are provided by Dial-A-Book Inc. solely for the personal use of visitors to this web site.
Meet the Author
L. David Mech is a senior research scientist with the Biological Resources Division, U.S. Geological Survey and adjunct professor in the Department of Fisheries, Wildlife, and Conservation Biology and the Department of Ecology and Behavioral Biology at the University of Minnesota. He is author of The Wolf: The Ecology and Behavior of an Endangered Species, The Way of the Wolf, and The Arctic Wolf, among other books, and is coauthor of The Wolves of Denali. Luigi Boitani is a professor of vertebrate zoology and animal ecology at the University of Rome. He is author of Dalla parte del lupo, coauthor of Simon and Schuster's Guide to Mammals, and coeditor of Research Techniques in Animal Ecology.
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