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2.2 Marine Phytoplankton 27 2.3 Marine Zooplankton 38 2.4 Other Topics 44 Elements of Marine Ecology ThisPageIntentionallyLeftBlank Elements of Marine Ecology Fourth Edition R.V. Tait F. A. Dipper Butterworth-Heinemann Linacre House, Jordan Hill, Oxford OX2 8DP 225 Wildwood Avenue, Woburn, MA 01801-2041 A division of Reed Educational and Professional Publishing Ltd A member of the Reed Elsevier plc group OXFORD BOSTON JOHANNESBURG MELBOURNE NEW DELHI SINGAPORE First published 1968 Reprinted 1970 Second edition 1972 Reprinted 1975, 1977, 1978 Third edition 1981 Reprinted 1983, 1988, 1992 Fourth edition 1998 ᭧ R. V. Tait and F. A. Dipper 1998 All rights reserved. No part of this publication may be reproduced in any material form (including photocopying or storing in any medium by electronic means and whether or not transiently or incidentally to some other use of this publication) without the written permission of the copyright holder except in accordance with the provisions of the Copyright, Designs and Patents Act 1988 or under the terms of a licence issued by the Copyright Licensing Agency Ltd, 90 Tottenham Court Road, London, England W1P 9HE. Applications for the copyright holder’s written permission to reproduce any part of this publication should be addressed to the publishers British Library Cataloguing in Publication Data Tait,R.V.(RonaldVictor) Elements of marine ecology. – 4th ed. 1 Marine ecology I Title II Dipper, Frances 577.7 ISBN 0 7506 2088 9 Typeset by Keyword Publishing Printed and bound in Great Britain Contents Preface ix 1 The oceans 1 1.1 Introduction 1 1.2 Extent and depth of the oceans 2 1.3 Ocean currents 9 1.4 Biological features of the marine environment 19 1.5 Elementary classification of the marine environment 20 2 Marine plankton 25 2.1 Definitions 26 2.2 Marine phytoplankton 27 2.3 Marine zooplankton 38 2.4 Other topics 44 3 Measuring and sampling 46 3.1 Oceanographic data 46 3.2 Biological sampling 61 3.3 Underwater observations 80 4 The seawater habitat – physical and chemical conditions 96 4.1 Introduction 96 4.2 Temperature 97 4.3 The composition of seawater 109 4.4 Specific gravity and pressure 124 4.5 Viscosity 135 4.6 Illumination 137 4.7 Currents 156 5 Organic production in the sea 166 5.1 Primary production 166 5.2 Measurements of organic production 174 5.3 Some factors regulating production 178 5.4 Ocean seasons 193 v vi Contents 5.5 Some mathematical models 202 5.6 Geographical differences of fertility 205 5.7 Turbidity currents 210 6 The sea bottom 213 6.1 The substrate 213 6.2 Benthic communities 220 6.3 Classification of communities 226 6.4 The food supply 232 7 Energetics of a marine ecosystem 244 7.1 Primary production 245 7.2 The grazing chain 248 7.3 The detritus chain 250 7.4 The energy balance sheet 253 7.5 Conclusion 253 8 The seashore 256 8.1 Tides 256 8.2 Waves 260 8.3 The evolution of coastlines 267 8.4 Some problems of shore life 270 8.5 Food sources 272 8.6 Zonation 273 8.7 Fitting the shore environment 275 8.8 Rocky shores 283 8.9 Sandy shores 292 8.10 Muddy shores 296 8.11 Estuaries 297 8.12 Tropical shores and coral reefs 303 9 Sea fisheries 313 9.1 Fishing methods 313 9.2 The biology of some European food fishes 326 9.3 The overfishing problem 350 9.4 Fishery research 358 9.5 The regulation of fisheries 368 9.6 The seas in relation to human food supplies 376 10 Human impact on the marine environment 395 10.1 Marine pollution 395 10.2 Climate change and global warming 419 Contents vii 10.3 Management and marine nature reserves 424 10.4 Effects of fisheries and acquaculture 429 Appendix 1 Topics for further study and class discussion or written work 436 Appendix 2 Some laboratory exercises 439 Appendix 3 Some field course exercises 441 Abundance scale for rocky shore intertidal organisms 444 A field course book list 447 Appendix 4 Addresses of organizations mentioned in the text 451 Index 453 ThisPageIntentionallyLeftBlank Preface It is now sixteen years since the third edition of this student textbook was published, and during this time advances in the field of marine biology have been many and varied. Continuing demand for the book has therefore led to this fourth, completely revised and updated edition. Developments in technology have been particularly rapid, especially in the fields of diving, precision instruments, satellite communications and computers, and these advances have greatly improved and expanded our ability to explore the oceans and their marine life. At the same time, the impact of man on the marine environment is ever increasing and cannot be ignored in any study of marine ecology. Therefore the chapters on ‘Measuring and sampling’ and ‘Sea fisheries’ have been extensively revised, and a new chapter on the ‘Human impact on the marine environment’ has been added. The aim of the latter is to introduce the student to the various ways in which our activities are changing the ecology of the oceans, with potentially far-reaching effects. Additional material on coral reefs, hydrothermal vents, phenomena such as ‘El Nin˜ o’ and ocean processes, including the important ‘microbial loop’ of the food web, have been incorporated into existing chapters. Although the book has been extensively revised, this edition retains its original aim of presenting marine ecology as a coherent science and its scope derives from the original, broad definition of ecology as the study of organisms in relation to their surroundings. The text has been compiled as introductory reading for students undertaking courses in marine biology. It provides information and ideas over the general field of marine ecology with reading lists and references from which additional material can be sought. With the busy student in mind, the text has been re-arranged into clearly labelled and numbered sections. The field course book list in the appendices has been extensively revised and updated as have the references and student texts listed at the end of each chapter. This new edition has been prepared by myself Dr Frances Dipper, with the guidance, help and unbounded enthusiasm of Ronald Tait. It was therefore with great sadness that I received the news of his untimely death last year. He will be sorely missed by his family and friends and by all those past students whom he has introduced to the fascination of the marine environment. It has been a privilege and a pleasure to work on this new edition of his book. Our thanks are due to our many colleagues who have provided us with helpful comments, material and encouragement throughout the preparation of this edition. Frances Dipper ix ThisPageIntentionallyLeftBlank 1 The oceans 1.1 Introduction Ecology is the study of relationships between organisms and their surroundings. This study is fundamental to an understanding of biology because organisms cannot live as isolated units. The activities which comprise their lives are dependent upon, and closely controlled by, their external circumstances, by the physical and chemical conditions in which they live and the populations of other organisms with which they interact. In addition, the activities of organisms have effects on their surroundings, altering them in various ways. Organisms therefore exist only as parts of a complex entity made up of interacting inorganic and biotic elements, to which we apply the term ecosystem. All life on earth constitutes a single ecosystem divisible into innumerable parts. This book is concerned with the greatest of these divisions, the marine ecosystem, occupying a larger volume of the biosphere than any other. The marine ecosystem can be further subdivided into many component ecosystems in different parts of the sea. Living processes involve energy exchanges. Energy for life is drawn primarily from solar radiation, transformed into the chemical energy of organic compounds by the photosynthetic processes of plants; thence transferred through the ecosystem by movements of materials within and between organisms, mainly through the agencies of feeding, growth, reproduction and decomposition. An ecosystem is therefore essentially a working, changing and evolving sequence of operations, powered by solar energy. In the long term, the intake of energy to the system is balanced by energy loss as heat. Most scientists believe that life began within the solar-powered environment of the ocean ecosystem. With the discovery in the 1970s of complex animal communities powered by geothermal energy at deep-sea vents (see Section 6.4.4), some scientists now argue that these vents may have allowed the first primitive organisms, such as bacteria, to evolve. The aim of marine ecological studies is to understand marine ecosystems as working processes. At present our knowledge is incomplete and only speculative analyses can be made. This book provides general information about how marine organisms are influenced by, and have effects on, their environment, and describes some of the methods of investigation which may eventually provide the necessary information for a better understanding of marine ecosystems. A good starting- 1 2 Elements of Marine Ecology pointistodescribebrieflysomeofthemajorphysicalfeaturesoftheoceans.There are several excellent introductory oceanography texts to which the student should referformoredetailedinformation.Thesearegiveninthereferencesattheend of Chapter 1. 1.2 Extent and depth of the oceans Seawater covers approximately 71 per cent of the earth’s surface, an area of about 361 million square kilometres (139 million square miles) comprising the major ocean areas shown in Figure 1.1. In the deepest parts the bottom lies more than 10 000 m from the surface, and the average depth is about 3700 m. Although marine organisms are unevenly distributed, they occur throughout this vast extent of water and have been brought up from the deepest places.
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