Deep-Sea Fishes Biology, Diversity, Ecology and Fisheries
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Deep-Sea Fishes Biology, Diversity, Ecology and Fisheries The technological advances of the last twenty years have brought huge advances in our understanding of the deep sea and of the species inhabiting this elusive and fascinating environment. Synthesising the very latest research and discoveries, this is a comprehensive and much-needed account of deep-sea fishes. Priede examines all aspects of this incredibly diverse group of animals, reviewing almost 3,500 species and covering deep-sea fish evolution, physiology and ecology as well as charting the history of their discovery from the eighteenth century to the present day. Providing a global account of both pelagic and demersal species, the book ultimately considers the effect of the growing deep-sea fishing industry on sustainability. Copiously illustrated with explanations of the deep-sea environment, drawings of fishes and information on how they adapt to the deep, this is an essential resource for biologists, conservationists, fishery managers and anyone interested in marine evolution and natural history. Imants G. Priede is Professor Emeritus at Oceanlab, a field research station of the Institute of Biological and Environmental Sciences, University of Aberdeen. He has over 40 years of experience studying deep-sea fish, from longlining and trawling on the Royal Research Ship Challenger in the 1970s to participating in and leading expeditions to the Pacific Ocean, Atlantic Ocean, Mediterranean Sea and the Mid-Atlantic Ridge. The founder of Oceanlab, he also pioneered satellite tracking of sharks in the 1980s and is the recipient of the 2011 Beverton Medal of the Fish Society of the British Isles. Deep-Sea Fishes Biology, Diversity, Ecology and Fisheries IMANTS G. PRIEDE Professor Emeritus, Oceanlab, University of Aberdeen, Scotland, UK University Printing House, Cambridge CB2 8BS, United Kingdom One Liberty Plaza, 20th Floor, New York, NY 10006, USA 477 Williamstown Road, Port Melbourne, VIC 3207, Australia 4843/24, 2nd Floor, Ansari Road, Daryaganj, Delhi - 110002, India 79 Anson Road, #06-04/06, Singapore 079906 Cambridge University Press is part of the University of Cambridge. It furthers the University’s mission by disseminating knowledge in the pursuit of education, learning, and research at the highest international levels of excellence. www.cambridge.org Information on this title: www.cambridge.org/9781107083820 DOI: 10.1017/9781316018330 © Imants G. Priede 2017 This publication is in copyright. Subject to statutory exception and to the provisions of relevant collective licensing agreements, no reproduction of any part may take place without the written permission of Cambridge University Press. First published 2017 Printed in the United Kingdom by TJ International Ltd. Padstow Cornwall A catalogue record for this publication is available from the British Library. Library of Congress Cataloging-in-Publication Data Names: Priede, I. G., author. Title: Deep-sea fishes : biology, diversity, ecology and fisheries / Imants G. Priede, professor emeritus, Oceanlab, University of Aberdeen, Scotland, UK. Description: Cambridge, United Kingdom ; New York, NY : Cambridge University Press, 2017. | Includes bibliographical references and index. Identifiers: LCCN 2017006037 | ISBN 9781107083820 (Hardback : alk. paper) Subjects: LCSH: Deep-sea fishes. | Deep-sea biology. | Deep-sea ecology. Classification: LCC QL620 .P75 2017 | DDC 597/.63–dc23 LC record available at https://lccn.loc.gov/2017006037 ISBN 978-1-107-08382-0 Hardback Cambridge University Press has no responsibility for the persistence or accuracy of URLs for external or third-party internet websites referred to in this publication and does not guarantee that any content on such websites is, or will remain, accurate or appropriate. To my wife, Maria. CONTENTS Preface page ix 1.7.2 Methods of Fish Capture 51 1.7.3 Imaging Systems 61 1 Introduction 1 1.7.4 Tagging and Tracking of Fishes 65 1.1 Definition of Deep-Sea Fishes 1 1.2 The Structure of the World’s Ocean 2 Colonisation of the Deep Sea by Fishes 68 Basins 2 2.1 Introduction 68 1.2.1 Ocean Depth Zones 2 2.2 Present-Day Depth Distribution of Fishes 68 1.2.2 Ocean Basin Formation, Slopes, 2.3 The History of Deep-Sea Fishes 72 Plains, Ridges, Islands and 2.4 Origins of the Modern Deep-Sea Fish Fauna 76 Seamounts 6 2.4.1 Agnatha 76 1.2.3 The Bathyal Slopes 2.4.2 Chondrichthyes 76 (200–3000 m Depth) 11 2.4.3 Actinopterygii 79 1.2.4 Hydrothermal Vents and 2.4.4 Sarcopterygii 85 Cold Seeps 12 2.5 Ancient and Secondary Deep-Sea Fishes 85 1.2.5 The Abyss (3000–6000 m Depth) 13 1.2.6 The Hadal Zone (>6000 m Depth) 15 3 Adaptations to the Deep Sea 87 1.3 History of the Ocean Basins 16 3.1 Introduction 87 1.4 The Deep-Sea Environment 21 3.2 Energy Metabolism 87 1.4.1 Deep Water Formation, Circulation 3.3 Effects of Pressure 90 and Oxygenation of the Deep 21 3.4 Size and Shape 91 1.4.2 Surface Ocean Circulation and Gyres 27 3.5 Buoyancy 93 1.4.3 Ocean Biomes 27 3.5.1 The Swim Bladder and Gas Gland 93 1.5 Food Supply to the Deep and the 3.5.2 Lipid Buoyancy 100 Distribution of Biomass 30 3.5.3 Weight Reduction 101 1.5.1 Export of Particulate Organic 3.6 Muscle Adaptations 102 Matter from the Surface 30 3.7 Gill Area 103 1.5.2 Food Falls 31 3.8 Heart and Circulatory System 104 1.5.3 Distribution of Biomass in 3.9 The Art of Slow Swimming and the the Deep Sea 31 Optimum Speed Fallacy 105 1.6 History of Discovery of Deep-Sea Fishes 35 3.10 Feeding and Growth 108 1.6.1 From the Beginnings of Ichthyology 3.11 Life History Strategies 114 to the First Deep-Sea Discoveries, 3.12 Excretion 118 1554–1860 35 3.13 Sensory and Communication Systems 119 1.6.2 Great Voyages of Discovery, 3.13.1 Diversity of Brain Morphology 119 1860–1914 38 3.13.2 Vision 122 1.6.3 To the First Manned Descents into 3.13.3 Camouflage in the Deep 125 the Deep, 1914–1945 41 3.13.4 Bioluminescence 127 1.6.4 The Great Twentieth-Century 3.13.5 The Olfactory and Gustatory Scientific Expansion from 1945 42 Systems 133 1.7 Collection and Methods 49 3.13.6 Underwater Sound and the 1.7.1 Accessing the Deep Sea 49 Octavo Lateral System 133 viii Contents 3.14 Time in the Deep Sea 135 4.3.25 Order Tetraodontiformes 299 3.15 Diseases and the Immune System 136 4.3.26 Order Lophiiformes 301 3.16 The Maximum Depth Limit for Fishes 138 4.4 Class Sarcopterygii 314 4.4.1 Order Coelacanthiformes 314 4 Systematic Description of Deep-Sea Fishes 139 5 The Deep-Sea Fish Faunas 317 4.1 Class Myxine 139 5.1 Mesopelagic and Bathypelagic Fish 4.1.1 Order Myxiniformes 139 Fauna 317 4.1.2 Order Petromyzontiformes 141 5.1.1 Arctic Ocean 319 4.2 Class Chondrichthyes 142 5.1.2 Pacific Ocean 319 4.2.1 Order Chimaeriformes 142 5.1.3 Indian Ocean 333 4.2.2 Order Hexanchiformes 144 5.1.4 Atlantic Ocean 334 4.2.3 Order Pristiophoriformes 144 5.1.5 The Antarctic Ocean 335 4.2.4 Order Squatiniformes 145 5.2 Bathyal Demersal Fishes 4.2.5 Order Squaliformes 145 (200–3000 m Depth) 335 4.2.6 Order Lamniformes 149 5.3 Abyssal Demersal Species 4.2.7 Order Carcharhiniformes 150 (3000–6000 m Depth) 347 4.2.8 Order Torpediformes 152 5.4 Fishes of the Hadal Zone 4.2.9 Order Pristiformes 152 (>6000 m Depth) 356 4.2.10 Order Myliobatiformes 152 5.5 Fishes of Hydrothermal Vents and 4.2.11 Order Rajiformes 153 Cold Seeps 358 4.3 Class Actinopterygii 155 5.6 The Circumglobal Deep-Sea Fish Fauna 362 4.3.1 Order Albuliformes 155 4.3.2 Order Notacanthiformes 156 6 Deep-Sea Fisheries and Conservation 363 4.3.3 Order Anguilliformes 158 6.1 Global Trends in Deep Fishing 363 4.3.4 Order Alepocephaliformes 166 6.2 Types of Fishing 364 4.3.5 Order Argentiniformes 171 6.2.1 Baited Traps and Lines 364 4.3.6 Order Stomiiformes 175 6.2.2 Static Nets 364 4.3.7 Order Ateleopodiformes 186 6.2.3 Pelagic Trawling 364 4.3.8 Order Aulopiformes 187 6.2.4 Bottom Trawling 365 4.3.9 Order Myctophiformes 196 6.3 Global Trends in Fish Catch 365 4.3.10 Order Lampriformes 203 6.4 Composition of the World Deep-Sea 4.3.11 Order Polymixiiformes 207 Fish Catch 367 4.3.12 Order Zeiformes 208 6.4.1 The Top Twenty Deep-Sea Species 384 4.3.13 Order Stylephoriformes 210 6.4.2 Other Species 391 4.3.14 Order Gadiformes 210 6.5 Overview of the Fishing Trends 400 4.3.15 Order Ophidiiformes 228 6.6 The Impacts of Deep-Sea Fishing 400 4.3.16 Order Batrachoidiformes 238 6.7 Conclusions 403 4.3.17 Order Gobiesociformes 238 4.3.18 Order Stephanoberyciformes 239 Epilogue 404 4.3.19 Order Beryciformes 241 4.3.20 Order Cetomimiformes 245 Appendix 406 4.3.21 Order Syngnathiformes 248 References 415 4.3.22 Order Scorpaeniformes 249 Index 470 4.3.23 Order Perciformes 264 Colour plates can be found between pages 252 and 4.3.24 Order Pleuronectiformes 294 253. PREFACE My first direct experience of deep-sea fishes was during November 1974 when I joined the Royal Research Ship Challenger for an expedition in the NE Atlantic under the leadership of Malcolm Clarke (Marine Biological Association Laboratory Plymouth, MBA) and John Blaxter (Scottish Marine Biological Association, Oban, SMBA).