The Biology and Management of Lobsters. Volume II: Ecology and Management PDF
Preview The Biology and Management of Lobsters. Volume II: Ecology and Management
Panulirus Cygnus, the West Australian Spiny Lobster. Photo by C. Purday. THE BIOLOGY AND MANAGEMENT OF LOBSTERS Volume II Ecology and Management Edited by J. STANLEY COBB Department of Zoology University of Rhode Island Kingston, Rhode Island BRUCE F. PHILLIPS CSIRO, Division of Fisheries and Oceanography North Beach, Western Australia Australia ACADEMIC PRESS 1980 A Subsidiary of Harcourt Brace Jovanovich, Publishers New York London Toronto Sydney San Francisco COPYRIGHT © 1980, 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. ACADEMIC PRESS, INC. 111 Fifth Avenue, New York, New York 10003 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 Biology and management of lobsters. Includes bibliographies and index. CONTENTS: v. 1. Physiology and behavior.—v. 2. Ecology and management. 1. Lobsters. 2. Lobster fisheries. I. Cobb, J. Stanley. II. Phillips, Bruce F. QL444.M33B56 595.3'841 79-6803 ISBN 0-12-177402-3 (v. 2) PRINTED IN THE UNITED STATES OF AMERICA 80 81 82 83 9 8 7 6 5 4 3 2 1 List of Contributors Numbers in parentheses indicate the pages on which the authors' contributions begin. David B. Bennett (317), Fairways, Burnham-on-Crouch, Essex CMO 8NR, England Β. K. Bowen (243), Department of Fisheries and Wildlife, Perth, Western Australia 6000 James M. Carlberg (333), Department of Biology, San Diego State University, San Diego, California 92182 C. J. Chapman (143), Marine Laboratory, Aberdeen AB9 8DB, Scotland R. A. Cooper (97), Manned Undersea Research and Technology, National Marine Fisheries Service, Woods Hole, Massachusetts 02543 R. L. Dow (265), Department of Marine Resources, Augusta, Maine 04333 Richard F. Ford (3), Center for Marine Studies, San Diego State University, San Diego, California 92182 D. A. Hancock (181), Western Australian Marine Research Laboratories, North Beach, Western Australia 6020 John T. Hughes (333), Massachusetts Division of Marine Fisheries, Lobster Hatchery and Research Station, Vineyard Haven, Massachusetts 02568 P. Kanciruk (59), Environmental Sciences Division, Oak Ridge National Lab oratory, Oak Ridge, Tennessee 37830 G. Marchesseault (219), New England Regional Fisheries, Management Council, Peabody, Massachusetts 01960 G. R. Morgan* (189)> Western Australian Marine Research Laboratories, North Beach, Western Australia 6020 B. F. Phillips (11), CSIRO, Division of Fisheries and Oceanography, North Beach, Western Australia 6020 S. B. Saila (219), Graduate School of Oceanography, University of Rhode Island, Kingston, Rhode Island 02881 *Present Address: Kuwait Institute for Scientific Research, Safat, Kuwait ix χ List of Contributors A. N. Sastry (11), Graduate School of Oceanography, University of Rhode Island, Kingston," Rhode Island 02881 J. R. Uzmann (97), Manned Undersea Research and Technology, National Marine Fisheries Service, Woods Hole, Massachusetts 02543 Jon C. Van Olst* (333), Department of Biology, San Diego State University, San Diego, California 92182 •Present Address: California Mariculture Corporation, La Jolla, California 92037 Preface The animals generally called lobsters fall into several taxonomically distinct groups: the clawed lobsters (Nephropidae), the spiny lobsters (Palinuridae), the slipper lobsters (Scyllaridae), and the coral lobsters (Synaxidae). Despite the taxonomic distinctions, there are many reasons to treat these animals together in a single text. Most notably, they are large, abundant animals that play important roles in the ecosystems in which they are found, and virtually all the abundant species are subject to intense and similarly applied fishing pressure. Much atten tion has been directed toward management of the valuable lobster stocks and to research related to this management. The study of lobster biology has a long history that well illustrates the intertwining of basic and applied research. All of the contributors to this volume are actively involved in attempts to understand lobster biology more fully and to apply that understanding to aiding management decisions. In the first of these two volumes an introductory chapter on the general biology of lobsters was included to provide an overview that we hoped would be helpful as a background for the entire work. The remainder of the first volume deals with lobster physiology and behavior. In this volume lobster ecology, population dynamics, management, and culture are discussed in depth. While the information presented in this book on the clawed lobsters and spiny lobsters is relatively voluminous, it is remarkable that information on the slipper lobster is very sparse, and knowledge of the coral lobster is almost nonexistent. All the authors were specifically asked to include information about the scyl- larids, but almost uniformly their response was that there was not enough infor mation to make any sort of statement. We considered but rejected the notion of including a chapter on the ecology of scyllarid juveniles and adults because, although some information exists, it is very fragmented and does not begin to present a satisfactory picture of scyllarid ecology. Despite the lack of informa tion, the Scyllaridae are a widely dispersed group that forms an important fraction of the larger benthic decapods of the world's oceans. It is axiomatic that sound management practices and appropriate culture tech niques must be based on good knowledge of the biology of the animal. In the 70 years since F. H. Herrick wrote his classic monograph concerning one of the lobster species, The Natural History of the American Lobster, a great deal more has been learned about the biology of all lobsters. Much of the new knowledge has arisen through the use of new tools. Some of the techniques used in the xi xii Preface research reviewed in this volume are illustrative: scuba, submersibles, submarine television, sophisticated bioeconomic models, growth equations appropriate to crustaceans, and advanced technology for culture systems. It is through the use of this new set of research instruments that a firm basis for rational management can be laid. Such management practices are badly needed. Many of the world's lobster fisheries are overfished and overcapitalized, with management strategies based on knowledge a half-century old. However, biological knowledge is not the only prerequisite for sound management. In many countries it appears that the management of heavily fished lobster populations is as much a political as it is a biological and economic problem. The political situation is not dealt with in this book. However, we hope that the biological information combined with reviews of the status of several of the fisheries presented in the following pages will aid in the futher development and refinement of management practices for the many species of lobsters. We believe that, in an overall sense, more is known about the biology of lobsters, particularly Homarus americanus, than any other aquatic invertebrate. Despite this, great holes in our knowledge persist. In this book we have tried not only to compile information but to present an integrated view of basic and applied biology while pointing out areas of exciting current and future research. If lobster research progresses at its current pace, we may soon be able to put together a whole view of the animal from evolutionary history through physiolog ical mechanisms to ecology and behavior. Such a picture will emerge only if both formal and informal communication between workers continues. These volumes are intended to stimulate and foster such communication as well as to provide a baseline of information from which to work. Many friends and colleagues have helped in the preparation of this book, and to them all we give our sincere thanks. In addition to those noted in the Preface to Volume I the following deserve special mention: P. F. Berry, Oceanographic Research Institute, Durban; N. G. Hall, Department of Fisheries and Wildlife, Western Australia; M. W. Johnson, Scripps Institute of Oceanography, San Diego; G. Newman, Department of Industries, Capetown; and D. Reid, CSIRO Mathematics and Statistics, New South Wales. The author of Chapter 4 thanks Β. B. Parrish and J. Mason for comments and the following for permission to quote or reproduce unpublished material: R. J. A. Atkinson, O. Bagge, A. Charuau, H. Eiriksson, A. S. D. Farmer, A. Fernandez-Garcia, Μ. M. Figueiredo, J. P. Hillis, G. G. Howard, J. A. Kitching, R. Jones, A. C. Simpson, D. J. Symonds, H. J. Thomas, P. Warren, Β. I. Dybern, and S. Hay. Both of the editors are deeply grateful for the superb assistance given by P. Barbour, S. Dennerlein, J. Downey, S. Koerner, G. McBride, R. Saunders, and C. Squires. J. S. Cobb B. F. Phillips Contents of Volume I Physiology and Behavior General Biology B. F. Phillips, J. S. Cobb, R. W. George Part I. Physiology Introduction W. Dall Molting and Growth D. E. Aiken Neurobiology B. W. Ache and D. L. MacMillan Reproductive Biology D. E. Aiken and S. L. Waddy Nutrition D.E. Conklin Diseases J. E. Stewart Part II. Behavior Introduction J. S. Cobb Spiny Lobsters: Patterns of Movement W. F. Herrnkind Social Behavior J. Atema and J. S. Cobb Index xiii Introduction RICHARD F. FORD From the information described in Chapters 1-4 and in the closely related section concerning fisheries management, it is clear that our knowledge about the ecology and habitat behavior of lobsters is extensive. Research in these areas of lobster biology has increased markedly over the last 10 years. Both the interest in lobsters and the amount of effort devoted to investigating their ecology and behavior are the result of several factors. Surely one of the most important of these is that lobsters are particularly interesting animals, from the standpoint of both their appearance and what they do. As juveniles and adults they occupy a variety of tropical, temperate, and boreal marine habitats in both shallow and deep water. In many cases they are important carnivores and prey species in the associated benthic communities. They have unusual morphological and behavioral adaptations, modes of life, and population characteristics, both as planktonic larvae and during their benthic existence. Most species of lobsters inhabiting benthic areas in shallow water are readily accessible for study in nature, in part because of their large size, their relatively high densities, and their tendency to remain in or near established shelters. Most of these species also are tolerant of environmental conditions and handling in laboratory aquarium systems. These characteristics make them excellent subjects for field and laboratory studies. Another equally important factor is that species of the families Nephropidae (clawed lobsters) and Palinuridae (spiny lobsters) support some of the most valuable fisheries in the world. Because of this, the population ecology and related characteristics of the species involved in these fisheries have been inves tigated extensively in some cases, using methods of the fisheries biologist. The catch per unit effort for many lobster fisheries has declined markedly, while the demand for both nephropid and palinurid lobsters and the monetary value of the catch continue to increase. These trends have intensified interest in 3 THE BIOLOGY AND MANAGEMENT OF LOBSTERS, VOL. Π Copyright © 1980 by Academic Press, Inc. All rights of reproduction in any form reserved. ISBN 0-12-177402-3 4 R. F. Ford learning more about the habitat requirements, population ecology, and fisheries-oriented behavior of the species involved, so that effects of the fishery can be evaluated and predicted as a basis for effective management. These trends also have intensified interest in developing fisheries for other species and stocks of lobsters, as for example of Panulirus penicillatus in Micronesia (MacDonald, 1971; George, 1972; Ford and Ebert, 1979). Efforts in such fisheries-oriented studies have provided much of the information now available concerning the environmental relationships and behavior of nephropid and palinurid lobsters during their benthic stages. One major disadvantage of this emphasis on studying exploited lobster species is that we know considerably less about the remaining ones and, in the case of many species, almost nothing at all. This is particularly true in general of the family Scyllaridae (slipper lobsters), the family Synaxidae (coral lobsters), deep water species of the Palinuridae, and many other palinurids which are not of commercial importance or support only minor fisheries. Such gaps in our knowl edge are unfortunate, because many of these forms differ from the well-studied species in their larval life histories and benthic habitat requirements, and presum ably also in their population characteristics and behavior. For example, it is significant that there is essentially no discussion of ecological and behavioral characteristics of the Scyllaridae in this volume, other than that concerning larval forms in Chapter 1. This simply reflects the fact that very little is known and still less has been published about these characteristics of slipper lobsters. Similarly, what we know about the ecological characteristics of palinurid lobsters is based on detailed studies of relatively few species, all of which inhabit shallow water areas. Thus, while our knowledge of the ecology of lobsters is substantial, it is not nearly as comprehensive as it should be. A second major problem resulting from our reliance on studies of exploited lobster populations is emphasized in Chapters 2-4 and in Chapter 7 of Vol. I. For practical reasons, fisheries biologists studying lobsters have tended to obtain ecological and behavioral data that have direct bearing on fishery problems and that can be obtained primarily by sampling from the fishery itself. These fishery data concerning distribution and abundance, size structure of the population, mortality, growth, reproduction, movements, and other fishery-related behavior are valuable not only to the fisheries biologist, but to all those studying lobster populations. Yet, by itself this information does not provide an adequate descrip tion of the basic ecological and behavioral characteristics of the lobster species considered. A related problem is that standard trapping and trawling methods used in many lobster fisheries tend to be highly selective against certain life history stages in the population, because of their size and behavior and the characteristics of the benthic habitats sampled. In addition, these fishery sam pling methods usually do not provide particularly accurate quantitative estimates of distribution and abundance of the kind needed in detailed studies of habitat requirements and population ecology.
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