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The Gray Whale: Eschrichtius Robustus PDF

587 Pages·1984·14.079 MB·English
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The Gray Whale Eschrichtius robustus Edited by Mary Lou Jones Steven L Swartz Cetacean Research Associates San Diego, California Stephen Leatherwood Hubbs-Sea World Research Institute San Diego, California with illustrations by Pieter Arend Folkens Oceanic Society San Francisco. California 1984 ACADEMIC PRESS, INC. (Harcourt Brace Jovanovich, Publishers) Orlando San Diego New York London Toronto Montreal Sydney Tokyo COPYRIGHT © 1984, BY ACADEMIC PRESS, INC. ALL RIGHTS RESERVED. NO PART OF THIS PUBLICATION MAY BE REPRODUCED OR TRANSMITTED IN ANY FORM OR BY ANY MEANS, ELECTRONIC OR MECHANICAL, INCLUDING PHOTOCOPY, RECORDING, OR ANY INFORMATION STORAGE AND RETRIEVAL SYSTEM, WITHOUT PERMISSION IN WRITING FROM THE PUBLISHER. ILLUSTRATIONS ON THE COVER, TITLE PAGE, AND PART TITLE PAGES COPYRIGHT © 1984 BY PIETER AREND FOLKENS. ACADEMIC PRESS, INC. Orlando, Florida 32887 United Kingdom Edition published by ACADEMIC PRESS, INC. (LONDON) LTD. 24/28 Oval Road, London NW1 7DX Library of Congress Cataloging in Publication Data Main entry under title: The Gray Whale. Includes index. 1. Pacific gray whale. 2. Whaling--Pacific Coast (North America) II. Jones, Mary Lou. II. Leatherwood, J. Stephen. III. Swartz, Steven L. QL737.C425G73 1984 599.5'1 84-3113 ISBN 0-12-389180-9 (alk. paper) PRINTED IN THE UNITED STATES OF AMERICA 84 85 86 87 9 8 7 6 5 4 3 2 1 This volume is dedicated to Carl L Hubbs, Laura C. Hubbs (top), and Raymond M. Gilmore (bottom), three pioneers of modern gray whale research. Contributors Numbers in parentheses indicate the pages on which the authors' contributions begin. LAWRENCE G. BARNES (3), Section of Vertebrate Paleontology, Natural History Museum of Los Angeles County, Los Angeles, California 90007 ALFRED ANTONOVICH BERZIN (409), Pacific Ocean Science Research Institute of Fisheries and Oceanography, T.I.N.R.O., 690600 Vladivostok, U.S.S.R. SERGEI ALEKSEEVICH BLOKHIN (487), Pacific Ocean Science Research Institute of Fisheries and Oceanography, T.I.N.R.O., 690600 Vladivostok, U.S.S.R. LUYDMILA S. BOGOSLOVSKAYA (465), A.N. Severtsoff Institute of Evolutionary Mor- phology and Animal Ecology, Academy of Sciences of U.S.S.R., 117071 Moscow, U.S.S.R. HOWARD W. BRAHAM (249), National Marine Mammal Laboratory, Northwest and Alaska Fisheries Center, National Marine Fisheries Service, N.O.A.A., Seattle, Washington 98115 PETER J. BRYANT (375), School of Biological Sciences, University of California, Irvine, Irvine, California 92717 MARILYN E. DAHLHEIM1 (511), Department of Zoology, University of British Columbia, Vancouver, British Columbia V6T 2A9 Canada JAMES D. DARLING2 (267), University of Victoria, Victoria, British Columbia V8W 2Y2, Canada and Vancouver Public Aquarium, Vancouver, British Columbia V6B 3X8, Canada JEANNETTE DA SILVA CURIEL (451), College of Veterinary Medicine, University of Saskatchewan, Saskatoon, Saskatchewan S7N 0W0, Canada H. DEAN FISHER (511), Department of Zoology, University of British Columbia, Van- couver, British Columbia V6T 2A9, Canada PIETER AREND FOLKENS, Oceanic Society, Suite E-225, Fort Mason, San Francisco, California 94123 JAMES T. HARVEY (561, 577), Marine Science Center, Oregon State University, New- port, Oregon 97365 DAVID A. HENDERSON (159), Whaling Museum, New Bedford, Massachusetts 02740 Present address: National Marine Mammal Laboratory, Northwest and Alaska Fisheries Center, National Marine Fisheries Service, N.O.AA, Seattle Washington 98115. 2Present address: West Coast Whale Research Foundation, Vancouver, British Columbia V7X 1L3, Canada. xv CONTRIBUTORS DENISE L HERZING (289), Marine Science Center, Oregon State University, Newport, Oregon 97365 MARY LOU JONES3 (309), Moss Landing Marine Laboratory, Moss Landing, California 95035 IGOR I. KRUPNICK (103), Institute of Ethnography, Academy of Sciences of the U.S.S.R., 117036 Moscow, U.S.S.R. CHRISTOPHER M. L7\FFERTY4 (375), School of Biological Sciences, University of California, Irvine, Irvine, California 92717 SUSAN K. U\FFERTY5 (375), School of Biological Sciences, University of California, Irvine, Irvine, California 92717 DONALD K. LJUNGBL^D (543), Naval Ocean Systems Center, San Diego, California 92109 BRUCE R. MATE (289, 561, 577), Marine Science Center, Oregon State University, Newport, Oregon 97365 SAMUEL A. MCLEOD (3), Section of Vertebrate Paleontology, Natural History Museum of Los Angeles County, Los Angeles, California 90007 JAMES G. MEAD (33), Division of Mammals, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560 EDWARD D. MITCHELL (33), Arctic Biological Station, Ste. Anne de Bellevue, Quebec H9X 3R4, Canada SUE E. MOORE (543), SEACO, Inc., San Diego, California 92106 KAREN R. MORIN (451), Department of Zoology, University of British Columbia, Van- couver, British Columbia V6T 2A9, Canada DEBRA J. MURIE (451), Department of Zoology, University of Guelph, Guelph, Ontario NIG 2Wl, Canada L7MJRIE D. MURISON (451), Department of Zoology, University of Guelph, Guelph, Ontario N1G2W1, Canada MARY NERINI (423), National Marine Mammal Laboratory, Northwest and Alaska Fisheries Center, National Marine Fisheries Service, N.O.A.A., Seattle, Washington 98115 BETH UMJRA O'LEARY (79), Department of Anthropology, University of New Mexico, Albuquerque, New Mexico 87107 HIDEO OMURA (57), Whales Research Institute, Tokyo 136, Japan M. MICHAEL POOLE (389), Division of Biology, Sonoma State University, Rohnert Park, California 94928 RANDALL R. REEVES6 (187), Division of Mammals, National Museum of Natural His- tory, Smithsonian Institution, Washington, D.C. 20560 3Present address: Cetacean Research Associates, San Diego, California 92107. 4Present address: 2157 Pattiz, Long Beach, California 90815. 5Present address: 2157 Pattiz, Long Beach, California 90815. 6Present address: Arctic Biological Station, Ste-Anne-de-Bellevue, Quebec. H9X 3R4 Canada. CONTRIBUTORS XV» STEPHEN B. REILLY (203), Southwest Fisheries Center, National Marine Fisheries Service, N.O.AA, La Jolla, California 92038 DAVID J. RUGH (225), National Marine Mammal Laboratory, Northwest and Alaska Fisheries Center, National Marine Fisheries Service, N.O.A.A., Seattle, Washington 98115 HAZEL SAYERS (121), American Cetacean Society, San Diego, California 92122 JAMES D. SCHEMPP (511), Ship Silencing Support Division, Puget Sound Naval Ship- yard, Bremerton, Washington 98314 STEVEN L SWARTZ7 (309), University of California, Santa Cruz, Santa Cruz, California 95064 ALEXEY V. YABLOKOV (465), N. K. Kol'tsoff Institute of Developmental Biology, Academy of Sciences of the U.S.S.R., 117334 Moscow, U.S.S.R. 7Present address: Cetacean Research Associates, San Diego, California 92107. Preface As student cetologists in the 1970s we learned that scholars had been at work for well over a century trying to piece together the gray whale story. Like many others interested in cetaceans, we often turned to this species to find a story with some sense of depth. The available accounts of other baleen whales were simply less comprehensive. But for the gray whale there existed an extensive body of "facts," mostly the offspring of a marriage of early scientific literature with a recent and rapidly expanding popular litera- ture. Much of what appeared in those earlier publications derived from observations of an exploited whale population during its recovery from a depleted state, and presumed gray whales were increasing in the absence of any density-dependent reduction of their annual reproductive rate. During the 1970s, however, it was generally agreed that gray whales in the eastern Pacific had returned to a stable near preexploitation stock size of between 11,000 and 13,000 animals following their protection from commercial harvest. This created a renewed interest in gray whale population dynamics and reproductive biology as it was hoped that a better understanding of these whales in particular might lead to greater insight into the life processes, reproduction, and growth of large ceta- cean populations in general. Much of what we knew or thought we knew about gray whales in the 1970s derived from the writings of Charles M. Scammon, Roy C. Andrews, Carl L Hubbs, Gordon C. Pike, Raymond M. Gilmore, David A. Henderson, Theodore J. Walker, and others. In 1971 Dale W. Rice and Alan A. Wolman of the National Marine Mammal Laboratory published The Life History and Ecology of the Gray Whale, a book that contributed substantially to the understanding of the species. In 1974 the Marine Fisheries Review released a special issue devoted to the California gray whale, dealing largely with research on a single gray whale yearling, "Gigi." This animal was collected as a calf in Baja California in 1971, was accessible for study during its year in captivity, and was finally radio tagged and released into the ocean in March 1972. With the above as background, investigators in the 1970s and early 1980s studied the accessible gray whale in all parts of its range, and research expanded rapidly. Due in part to legal protection afforded this species in Canada, Mexico, and the United States and in part to increased sensitivity to living whales, field researchers recently have accelerated the development and use of "benign nonconsumptive" techniques of study. The study of "live" long-lived mammals requires commitment to long-term programs to provide the basis for population trend analysis and meaningful behavioral studies; such long-term commitments have been beyond the scope of many cetacean field research programs. Therefore, even though the results of some programs completed in the past PREFACE decades are exciting, most must be viewed as preliminary and not exhaustive treat- ments. Results of most remain unpublished, and accounts of their progress are scattered in technical and contract volumes difficult for even the most dedicated cetacean biolo- gist to obtain. Late in the 1970s, at numerous scientific meetings sponsored by the International Whaling Commission, World Wildlife Fund, International Union for Conservation of Nature, Sociedad Mexicana para el Estudio de Mamiferos Marinos, National Marine Fisheries Service-National Marine Mammal Laboratory, Marine Mammal Commission, and other agencies, this new information on gray whales was being discussed. It be- came increasingly apparent to the participants that a publication featuring the results of these recent programs was highly desirable. With this goal in mind, we announced our intention to collect and edit a volume of research papers on gray whales, and issued a call for contributions at the IV Biennial Conference on the Biology of Marine Mammals in November of 1981 in San Francisco, California. The response was overwhelming and far exceeded our greatest expectations. We and the colleagues who joined us believe this volume will fill a need for better distribution of information. In this sense we see The Gray Whale as a transitional vehicle, a summary of current information upon which to build. Scientific research almost always raises as many questions as it answers; it is our sincere hope that this volume will stimulate both more work and publication. We are particularly pleased that The Gray Whale is not provincial. It contains contribu- tions from researchers working in Canada, Japan, Mexico, the United States, and the Soviet Union and information from throughout the gray whale's past and present ranges both in the Atlantic and Pacific. Such diversity points up the fact that neither understand- ing nor conservation of this species can be effected without truly sincere international commitment and cooperation. Finally, we wish to extend our thanks to all those individuals, organizations, agencies, translators, and reviewers, anonymous and otherwise, who graciously contributed their time, expertise, and energy to the production of The Gray Whale. Special thanks to Pieter Folkens for his original art for the subject sections, cover, and title page and Janice J. Sherer for her maps. Introduction The gray whale is clearly an important species from the standpoint of public educa- tional and recreational value, aesthetic appeal, economic significance, and scientific interest. Because of its coastal habits the gray whale is the only large whale that can be regularly observed from shore. Its lengthy annual migration is one of the world's most outstanding wildlife spectacles, viewed by more than two million people each year. The gray whale is the sole member of the family Eschrichtiidae and is considered by many to be the most primitive surviving baleen whale. There are two extant populations. One, the western Pacific or Korean population, is critically depleted. The other, eastern Pacific or California population, is a well-known conservation success. Whaling reduced the latter population in the 1850s, following the discovery of the breeding lagoons, and again after the turn of the century with the introduction of floating factories. Even so, this population twice made remarkable recoveries. Since it was afforded protection from commercial whaling in 1946 by the International Convention for the Regulation of Whaling, the population has grown to its current estimated level of 16,000 or more animals and continues to grow. Most whaling historians and biologists believe the preexploitation stock size was between 15,000 and 24,000 animals. Presently, a non- commercial quota is set each year. In the Arctic, Alaskan Eskimos from St. Lawrence Island still take a few gray whales annually, and Soviet-government catcher boats take about 140 to 200 gray whales each year for use by Siberian aborigines. In 1978 the International Whaling Commission (IWC) reclassified the California stock, moving it from protected to sustained-management status. Even with this reclassification and continuing annual harvest, however, the Scientific Committee (IWC) has acknowledged that coastal development and industrial activity pose very real threats to the survival of gray whales. Collectively the chapters in The Gray Whale reflect the intimate relationship between gray whales and the coastal habitat in which they have evolved and now live. It is precisely their coastal habits that place gray whales in direct conflict with humans. They cannot avoid exposure to our pollution, vessel traffic, industrial noise, and activities associated with the development of outer continental shelf resources over virtually their entire range. Offshore oil and natural gas exploration and development by the U.S.S.R., the United States, Canada, and Mexico could negatively affect the gray whales on their nothern feeding grounds, during migration, and in their wintering grounds by disturbing these critical habitats. Industrial activities, fishing, commercial whale watching, and private boating along the entire west coast of North America may obstruct the progress of migrating gray whales. Finally, Mexico has jurisdiction over much of their winter range xxii INTRODUCTION where reproduction is accomplished, and potential disturbances there include local industrial development and foreign recreational pursuits, mainly U.S. tourism. The balance among aesthetics and scientific desires, local economic needs that contribute to environmental disturbance, and whale well-being is both a complicated and sensitive conservation issue. Gray whales can only be assured of survival if their lives, habitats, and food resources are managed according to an admittedly difficult but responsible ecosystem approach. Concern for the future of gray whales has provided an incentive for private and government support of much recent research within many disciplines, a trend we wish to see continue. Mexico's establishment of gray whale sanctuaries in Laguna Guerrero Negro, Laguna Ojo de Liebre, and Laguna San Ignacio, the first ever for cetaceans, is an encouraging step we hope will be matched by sensitive treatment of gray whale needs in other parts of their range. The diversity of interest in gray whales was exemplified by the diversity of manuscripts we received. Contributions came from historians, anthropologists, paleontologists, biologists, and naturalists alike. In addition to chapters presenting new findings on the evolution, phylogeny, biology, demography, and behavior of gray whales, we received excellent chapters dealing with the history of their exploitation by both aborigines and industrialized nations. The papers we received fell into four more or less obvious categories which have become the major sections of this volume. Section I presents two essays dealing with gray whale evolution, fossils, and subfossil remains, systematics, and range in historical times. Barnes and McLeod describe the only known bonafide gray whale fossil and examine the phyletic relationship between gray whales and other mysticetes. Mead and Mitchell provide a detailed review of the gray whale subfossil remains that have been discovered on both sides of the Atlantic Ocean and which confirm that the gray whale was, at sometime, common in the western Atlantic American waters and was taken commercially as recently as American colonial times. Section II documents historical aspects of gray whale exploitation and the economic importance of these whales to humans, beginning with aboriginal societies in Asia and North America, and leading up to the most recent period of commercial exploitation at the turn of the century. Omura describes two populations of gray whales that migrated to the coastal waters of Japan prior to 1900 and suggests that the Seto Inland Sea of southern Japan served as a calving area. Krupnik traces the origins of aboriginal hunting along the Chukotka Peninsula and, based on excavations of ancient settlements, sug- gests that the gray whale, particularly young animals, and not the bowhead whale, was the primary object of much of the aboriginal hunting. O'Leary reviews the relevant ethnographic, historical, and archaeological data on aboriginal whaling and challenges previous assumptions about the importance of gray whale subsistence hunting by the Indian groups of the Pacific Northwest. Sayers reconstructs the rise and fall of the colorful and sometimes profitable shore whaling industry of the California coast, from its beginnings in Monterey in 1854 until its final collapse in 1901. Henderson's detailed account of the lagoon whaling period places this episode in the more general perspec- tive of nineteenth century whaling throughout the Pacific. He describes how Yankee and European whalers pursued gray whales during the summer in the Bering, Okhotsk, and

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