Wolves of the World: Perspectives of Behavior, Ecology and Conservation (Noyes Series in Animal Behavior, Ecology, Conservation, and Management) PDF
Preview Wolves of the World: Perspectives of Behavior, Ecology and Conservation (Noyes Series in Animal Behavior, Ecology, Conservation, and Management)
WOLVES OF THE WORLD NOYES SERIES IN ANIMAL BEHAVIOR, ECOLOGY, CONSERVATION AND MANAGEMENT BenjaminB. Beck, Ph.D., ConsultingEditor A series of professional and reference books in ethology devoted to the better understanding ofanimal behavior, ecol ogy, conservation, and management. WOLVES OF THE WORLD: Perspectives of Be havior, Ecology, and Conservation. Edited by Fred H. Harrington andPaul C. Paquet IGUANAS OF THE WORLD: Their Behavior, Ecology, and Conservation. Edited by Gordon M. Burghardt and A. Stanley Rand HORSE BEHAVIOR: The Behavioral Traits and Adaptations ofDomestic and Wild Horses, Includ ing Ponies. By George H. Waring GAZELLES AND THEIR RELATIVES: A Study in Territorial Behavior By Fritz R. Walther, Elizabeth Cary Mungall, and Gerald A. Grau THE MANAGEMENTANDBIOLOGY OFANEX TINCT SPECIES: PERE DAVID'S DEER Edited by Benjamin B. Beck and Christen Wemmer WOLVES OF THE WORLD Perspectives of Behavior, Ecology, and Conservation Edited by FRED H. HARRINGTON Mount Saint Vincent University Halifax, Nova Scotia Canada and PAUL C. PAQUET Portland State University Portland, Oregon U.S.A. NOYES PUBLICATIONS Park Ridge, New Jersey, U.S.A. Copyright ©1982 by Noyes Publications No part ofthis book may be reproduced in any form without permission in writing from the Publisher. Library of CongressCatalog Card Number: 82-3397 ISBN: 0·8155·0905·7 Printed in the United States Published in the United Statesof America by NoyesPublications Mill Road, Park Ridge, New Jersey07656 10Q8 7 6 543 Libraryof CongressCataloging in Publication Data Main entry undertitle: Wolves of the world. Most ofthe papers included were originally presented at the 1979 Portland International Wolf Symposium. Includesbibliographiesand index. 1. Wolves--Congresses. I. Harrington, Fred H. II. Paquet, Paul C. III. Portland International Wolf Symposium (1979 : Or.) QL737.C22W65 599.74'442 82-3397 ISBN 0·8155·0905·7 AACR2 Preface "Only a mountain has lived long enough to listen objectively to the howl of a wolf" -Aldo Leopold, Game Management, 1933 Aldo Leopold was mistaken. We have lived and interacted with wolves from prehistoric to modern times, certainly ample time to comprehend and view them rationally. Exactly how long our ac quaintance has endured is uncertain, but our association probably be gan when our ancestors first immigrated to the northern hemisphere. There, we encountered our ecological counterpart and perhaps most direct competitor, the wolf. Our lives intertwined to such an extent that we selected the wolf (or were selected by it) as our first domes tic companion. Yet despite countless opportunities over untold gen erations, our perceptions of wolves reflected our prejudices more than reality. Our fancied notions about wolves finally began to erode in the 1940s when Adolf Murie, Rudolf Schenkel, Ian McTaggart Cowan and Sigurd Olsen pioneered modern, scientific study into the an imal's ways. During the following decade, insights continued to accu mulate, although investigators were few and the research lacked a consistent direction. Near the end of the 1950s, however, two pro jects were conceived which would soon provide the needed direc tion. Durward Allen recognized that Isle Royale was theperfectwilder ness laboratory-isolated, relatively small, yet teeming with wolves, moose and beaver. For two decades he guided a line ofgraduate stu dents through a series of ground-breaking studies. At the same time, Douglas Pimlott saw the potential of Algonquin Provincial Park. Pimlott's students, like Allen's, broke ground in a number of areas, includingthe introduction ofthe radio-collar in wolfresearch. By 1970, the stage was set for synthesis. David Mech, the first of Allen's students on Isle Royale, gathered together what was then v vi Preface known into The Wolf. The book provided an immediate focus for subsequent research efforts which, until then, had been somewhat diffused. At the same time, Mech brought the radio-collar to north ernMinnesotaand initiateda study which would become a model and standard for future work. The effects were staggering. Currently, field studies using radio-telemetry are being pursued from Alaska south to Minnesota and Montana, in Europe, the Middle East, and perhaps soon in Asia. At present, observations of wolves are out strippingourability to synthesize and place them into perspective. The 1980s should provide us with a more complete and refined picture of wolves, a picture with sufficient complexity to encompass the extensive variation now seen amongst them. This book is part of that process. Most ofthe chapters included were originally presented at the 1979Portland WolfSymposium, Portland,Oregon. That meet ing brought together the majority of wolf researchers from around the world. Our originalhope was that the conference would generate an updated synthesis of wolf ecology and behavior. We now realize that our hope was premature. As more data have accumulated, the less secure our previous generalizations about wolves seem. More data are required, datafrom diverse areas undervaried environmental conditions. We must avoid hurrying our observations toward prema ture conclusions that so easily, and erroneously, imply "this isTHE WOLF." At present, there is an unspoken urgency as wolf researchers pursuetheir quarry. So often, intriguing observationsareleftdangling as wolves, packs, or even populations succumb to civilization's on slaught. Our studies are no longer esoteric exercises in biology and psychology. What we learn may, or rather must, be used eventually to ensure that wolves continue to exist. Thus, the papers in thisvol ume sample areas which we feel are necessary ifwe are to secure this planetfor the wolves. The first two sections survey the areas ofecology and behaviorin the wild. Because an animal's behavior is dependent on its ecology, we have made no effort to segregatethese two areas. Rather, we have made our division between "disturbed" and "undisturbed"habitats. Studiesofwolves in North Americaprovide aview ofwolvesasclassic predators of big game in relatively undisturbed wilderness habitat. Here we can patiently attempt to disentangle the complex web of interrelations that characterize "natural" communities. As we learn more about the workings of these systems, we can make wiser de cisions on management necessitated by our society's appetite for minerals, lumber and energy. Studies of wolves in Eurasia can then provide an important and necessary contrast, and perhaps a glimpse into the future prospects of wolves in areas still pristine by 1980 standards. Eurasian wolves have managed to survive in habitats long altered by us. Ifwe canunderstand how they have managed to adapt to the drastic alterationswe have made in their environment, we may have an idea ofhow greata shockwolves canwithstand, and whatwe can do to lessenthat shock. Preface vii The third section provides a sampling of behavior studies con ducted in captivity. Captive studies have often been maligned be cause they inevitably must confine a wide-ranging animal to a mere fraction of what it might normally traverse. Butwide-rangingwolves are also elusive. Captive studiesallow us to focus onthose rarely seen aspects ofwolflife and permit usdeeper insights into other areasjust glimpsed in the field. Conservation is the subject of the fourth section. With wolfpop ulations variously labeled as "threatened", "endangered" and "ex tinct", and further inroads being made into wolf habitat day by day, it may soon not be enough to merely set aside refuges for the remaining populations. As several papers inthis section pointout, we must use ourknowledge ofwolf behaviorand ecology to balance our demandsfor the wolf's habitatwith our goals for wolfpreservation. We conclude thebookwith two perspectives onwolves, and wolf study,from observersoutside the typicalbiologist/psychologistframe of mind. As philosophers of science have pointed out for years, our observations as scientists are biased by our cultural, sociological and scientific backgrounds. A recognition and understanding of other viewpoints can only serve to open our eyes wider and allow us to "see" phenomenato which we might otherwise have been blind. Although we have presented the major areas ofpresent-day wolf research, our collection is by no means exhaustive. Some areas have been underrepresented and others, unfortunately, are absent. The areas which are included, however, should provide an overview indi catingwhere the study ofwolves is headed duringthe 1980s. Halifax, Nova Scotia Fred H. Harrington Portland, Oregon Paul C. Paquet April1982 Contributors Warren B. Ballard John C. Fentress Alaska DepartmentofFish & DepartmentofPsychology Game Delhousie University Glennallen, Alaska Halifax, Nova Scotia, Canada William E. Berg PatrickM. Ghezzi Minnesota Departmentof Utah State University Natural Resources Logan, Utah Grand Rapids, Minnesota Fred H. Harrington DmitriI. Bibikov Psychology Department A.N. Severstov Institute of Mt. St. Vincent University Animal Evolution Halifax, Nova Scotia, Canada Morphologyand Ecology DaryllM. Hebert USSR AcademyofScience B.C. FishandWildlife Branch Moscow, USSR Nanaimo, British Columbia, Canada AndersBjarvall Robert E. Henshaw National Environmental N.Y. DepartmentofEnvironmental Protection Board Conservation 801na, Sweden Albany, New York Luigi Boitani Richard A. Hook Istituto di Zoologia, DepartmentofBiology Universita di Roma Northern Michigan University Rome, Italy Marquette, Michigan Susan Bragdon ErikIsakson Williams College National Environmental Protection Williamstown, Massachusetts Board LudwigN. Carbyn Solna, Sweden Canadian Wildlife Service DavidJames Edmonton, Alberta, Canada Alaska DepartmentofFish &Game Carl D. Cheney Fairbanks, Alaska Institute ofAnimal Behavior DougJanz Utah State University B.C. FishandWildlife Branch Logan, Utah Nanaimo, British Columbia, Canada Rick Davies PaulJoslin B.C. FishandWildlife Branch Chicago Zoological Society Nanaimo, British Columbia, Brookfield, lllinois Canada ix
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