BIOLOGICAL OCEANOGRAPHY AN INTRODUCTION Related titles ELDER & WILLIAMS Fluid Physics for Oceanographers and Physicists, 2nd edition OPEN UNIVERSITY The Ocean Basins: Their Structure and Evolution Seawater: Its Composition, Properties and Behaviour, 2nd edition Ocean Circulation Waves, Tides and Shallow-Water Processes Ocean Chemistry and Deep-Sea Sediments Case Studies in Oceanography and Marine Affairs PARSONS, TAKAHASHI & HARGRAVE Biological Oceanographic Processes, 3rd edition PICKARD & EMERY Descriptive Physical Oceanography: An Introduction, 5th edition POND & PICKARD Introductory Dynamical Oceanography, 2nd edition TAIT & DIPPER Elements of Marine Ecology, 4th edition TOMCZAK & GODFREY Regional Oceanography: An Introduction BIOLOGICAL OCEANOGRAPHY AN INTRODUCTION Second Edition CAROL M LALLI and TIMOTHY R PARSONS University of Britisli Columbia, Vancouver, Canada ELSEVIER BUTTERWORTH HEINEMANN AMSTERDAM • BOSTON • HEIDELBERG • LONDON • NEW YORK • OXFORD PARIS • SAN DIEGO • SAN FRANCISCO • SINGAPORE • SYDNEY • TOKYO Elsevier Butterworth-Heinemann Linacre House, Jordan Hill, Oxford 0X2 8DP 30 Corporate Drive, Burlington, MA 01803 First published 1993 Reprinted with corrections 1994, 1995 Second edition 1997 Reprinted 1999, 2000, 2001, 2002, 2004, 2006 Copyright © 1993, 1997, C M Lalli andX R Parsons All rights reserved The right of C M Lalli andT R Parsons to be identified as the author of this work has been asserted in accordance with the Copyright, Designs and Patents Act 1988 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 WIT 4LP Applications for the copyright holder's written permission to reproduce any part of this publication should be addressed to the publisher Permissions may be sought directly from Elsevier's Science & Technology Rights Department in Oxford, UK: phone: (+44) 1865 843830, fax: (+44) 1865 853333, e-mail: permissions@elsevier.co.uk You may also complete your request on-line via the Elsevier homepage (http://www.elsevier.com), by selecting 'Customer Support' and then 'Obtaining Permissions' British Library Cataloguing in Publication Data Lalli, Carol M Biological oceanography: an introduction - 2nd ed Marine biology Marine ecology I Title II Parsons, Timothy R (Timothy Richard), 1932574.9'2 Library of Congress Cataloguing in Publication Data Lalli, Carol M Biologocial oceanography: an introduction / Carol M Lalli and Timothy R Parsons - 2nd ed p cm Includes bibliographical references and index Marine biology Marine ecology Oceanography I Parsons, Timothy Richard, 1932- II Title QH91.L35 96-42139 574.92-dc20 CIP ISBN 7506 3384 For information on all Elsevier Butterworth-Heinemann publications visit our website at www.bh.com Working together to grow libraries in developing countries www.elsevier.com | www.bookaid.org | www.sabre.org ELSEVIER l!,?5?.^ A;.^ Printed and bound in China Sabre Foi CONTENTS ABOUT THIS VOLUME CHAPTER XI INTRODUCTION 1.1 1.2 1.3 1.3.1 1.4 1.5 CHAPTER SPECIAL PROPERTIES AFFECTING LIFE IN THE SEA CLASSIFICATIONS OF MARINE ENVIRONMENTS AND MARINE ORGANISMS BASIC ECOLOGICAL TERMS AND CONCEPTS r- and /C-selection THE HISTORICAL DEVELOPMENT OF BIOLOGICAL OCEANOGRAPHY SUMMARY OF CHAPTER 13 THE ABIOTIC ENVIRONMENT 2.1 2.1.1 2.1.2 2.2 2.2.1 2.2.2 2.3 2.3.1 2.3.2 SOLAR RADIATION Radiation at the sea surface Radiation in the sea TEMPERATURE Sea surface temperatures 16 17 18 21 21 Vertical temperature distribution 24 SALINITY Range and distribution of salinity 25 26 Biological importance of salinity 28 2.4 DENSITY 2.5 PRESSURE 2.6 SURFACE CURRENTS 2.6.1 2.7 CHAPTER 30 33 Biological significance of currents 34 35 SUMMARY OF CHAPTER 36 PHYTOPUNKTON AND PRIMARY PRODUCTION 3.1 3.1.1 3.1.2 SYSTEMATIC TREATMENT Diatoms Dinoflagellates 40 40 42 3.1.3 Other phytoplankton 45 PHOTOSYNTHESIS AND PRIMARY PRODUCTION Methods of measuring biomass and primary productivity 46 48 RADIATION AND PHOTOSYNTHESIS 50 THE EFFECT OF NUTRIENTS ON GROWTH RATE 53 PHYSICAL CONTROLS OF PRIMARY PRODUCTION Oceanic gyres and rings 58 60 3.2 3.2.1 3.3 3.4 3.5 3.5.1 Continental convergence and divergence 62 3.5.3 Planetary frontal systems 63 3.5.4 63 3.5.5 Shelf-break fronts River-plume fronts 3.5.6 Island mass effect and Langmuir frontal zones 66 3.6 GLOBAL PHYTOPLANKTON PRODUCTIVITY 67 3.7 SUMMARY OF CHAPTER 70 3.5.2 CHAPTER 65 ZOOPLANKTON 4.1 COLLECTION METHODS 74 4.2 HOLOPLANKTON: SYSTEMATICS AND BIOLOGY 76 4.3 MEROPLANKTON 86 4.4 VERTICAL DISTRIBUTION 90 4.5 DIEL VERTICAL MIGRATION 94 4.6 SEASONAL VERTICAL MIGRATIONS 98 4.7 ZOOGEOGRAPHY OF THE HOLOPLANKTON 100 Patchiness 103 4.8 LONG-TERM CHANGES IN ZOOPLANKTON COMMUNITY STRUCTURE 108 4.9 SUMMARY OF CHAPTER 109 4.7.1 CHAPTER ENERGY FLOW AND MINERAL CYCLING 5.1 FOOD CHAINS AND ENERGY TRANSFER 5.2 FOOD WEBS 5.2.1 5.3 5.3.1 5.3.2 5.4 5.5 The microbial loop MEASURING SECONDARY PRODUCTION Field studies Experimental biological oceanography A COMPARISON OF MARINE AND TERRESTRIAL PRODUCTION OF ORGANIC MATERIAL 5.5.1 MINERAL CYCLES Nitrogen 5.5.2 Carbon 5.6 CHAPTER SUMMARY OF CHAPTER 112 117 121 123 123 125 134 136 137 141 143 NEKTON AND FISHERIES OCEANOGRAPHY 6.1 NEKTONIC CRUSTACEA 147 6.2 NEKTONIC CEPHALOPODS 148 6.3 MARINE REPTILES 149 6.4 MARINE MAMMALS 149 VII 6.5 SEABIRDS 6.6 6.6.1 MARIHE FISH Fish migrations 6.7 6.7.1 FISHERIES AND FISHERIES OCEANOGRAPHY World fish catch and fisheries management 6.7.2 Fluctuations in the abundance of fish stocks 153 156 162 6.7.3 Regulation of recruitment and growth in fish 6.7.4 Fishing and the use of near real-time oceanographic data 162 163 164 167 170 6.8 MARICULTURE 172 6.9 SUMMARY OF CHAPTER 174 CHAPTER BENTHOS 7.1 7.1.1 7.2 BENTHIC PLANTS Measurements of benthic primary production 178 180 BENTHIC ANIMALS 180 7.2.1 Systematics and biology 182 7.2.2 Sampling and production measurements 191 7.3 DETERMINANTS OF BENTHIC COMMUNITY STRUCTURE 192 7.4 SUMMARY OF CHAPTER 193 CHAPTER BENTHIC CORAMUNITIES 8.1 8.1.1 8.1.2 INTERTIDAL ENVIRONMENTS Tides Environmental conditions and adaptations of intertidal organisms 196 197 197 8.2 8.2.1 8.2.2 ROCKY INTERTIDAL SHORES Zonation Trophic relations and the role of grazing and predation in determining community structure 198 198 201 8.3 KELP FORESTS 202 8.4 SAND BEACHES 205 Environmental characteristics Species composition 205 206 ESTUARIES 209 CORAL REEFS Distribution and limiting factors 213 213 8.4.1 8.4.2 8.5 8.6 8.6.1 8.6.2 Coral structure 214 8.6.3 Diversity 215 8.6.4 Nutrition and production in reefs 217 8.6.5 8.6.6 Production estimates Formation and growth of reefs 218 219 8.6.7 Zonation patterns on reefs 221 8.7 MANGROVE SWAMPS 222 8.7.1 What are mangroves? 223 8.7.2 Ecological features of mangrove swamps 224 8.7.3 Importance and uses of mangroves 225 DEEP-SEA ECOLOGY 226 8.8.1 8.8 Faunal composition 227 8.8.2 Species diversity 231 8.8.3 8.8.4 Biomass Food sources 232 8.8.5 Rates of biological processes 238 8.8.6 Future prospects 238 8.9 8.9.1 HYDROTHERMAL VENTS AND COLD SEEPS Chemosynthetic production 239 8.9.2 Vent fauna 239 8.9.3 Shallow vents and cold seeps 8.9.4 Unique environmental features of sulphide communities 242 8.10 SUMMARY OF CHAPTER 243 CHAPTER 233 238 241 HUMAN IMPACTS ON MARINE BIOTA 9.1 9.2 9.2.1 FISHERIES IMPACTS MARINE POUUTANTS Petroleum hydrocarbons 9.2.2 Plastics 9.2.3 Pesticides and other biologically active organic compounds 9.2.4 Heavy metals 9.2.5 Sewage 9.2.6 9.2.7 Radioactive wastes Thermal effluents 9.3 INTRODUCTIONS AND TRANSFERS OF MARINE ORGANISMS 9.4 9.4.1 IMPACTS ON SPECIFIC MARINE ENVIRONMENTS Estuaries 9.4.2 Mangrove swamps 9.4.3 Coral reefs 9.5 SUMMARY OF CHAPTER 248 251 252 252 253 254 255 255 256 256 258 258 259 260 263 APPENDIX GEOLOGIC TIME SCALE 266 APPENDIX CONVERSIONS 267 SUGGESTED FURTHER READING 269 ANSWERS AND COMMENTS TO QUESTIONS 270 GLOSSARY 287 ACKNOWLEDGEMENTS 304 INDEX 307