OCEANOGRAPHY and MARINE BIOLOGY AN ANNUAL REVIEW Volume 37 OCEANOGRAPHY and MARINE BIOLOGY AN ANNUAL REVIEW Volume 37 Editors A.D.Ansell R.N.Gibson Margaret Barnes The Dunstaffnage Marine Laboratory Oban, Argyll, Scotland Founded by Harold Barnes First published 1999 by Taylor & Francis 11 New Fetter Lane, London EC4P 4EE Simultaneously published in the USA by Taylor & Francis 29 West 35th Street, New York, NY1001 Taylor & Francis is an imprint of the Taylor & Francis Group This edition published in the Taylor & Francis e-Library, 2003. © 1999 A.D.ANSELL, R.N.GIBSON, MARGARET BARNES All rights reserved. No part of this book may be reprinted or reproduced or utilised in any form or by any electronic, mechanical, or other means, now known or hereafter invented, including photocopying and recording, or in any information storage or retrieval system, without permission in writing from the publishers. Every effort has been made to ensure that the advice and information in this book is true and accurate at the time of going to press. However, neither the publisher nor the authors can accept any legal responsibility or liability for any errors or omissions that may be made. In the case of drug administration, any medical procedure or the use of technical equipment mentioned within this book, you are strongly advised to consult the manufacturer’s guidelines. British Library Cataloguing in Publication Data A catalogue record for this book is available from the British Library Library of Congress Cataloging in Publication Data are available ISBN 0-203-48482-7 Master e-book ISBN ISBN 0-203-79306-4 (Adobe eReader Format) ISBN 0-748-40919-X (Print Edition) CONTENTS Preface vii Air-sea gas exchange into the millennium: progress and uncertainties 1 Thomas Frost & Robert C.Upstill-Goddard The role of benthic microalgae in neritic ecosystems 47 Lawrence B.Cahoon Mesoherbivore-macroalgal interactions: feeding ecology of sacoglossan sea slugs (Mollusca, Opisthobranchia) and their effects on their food algae 87 S.I.Williams & D.I.Walker Population structuring, gene dispersal and reproduction in the Actinia equina species group 129 M.C.Perrin, J.P.Thorpe & A.M.Solé-Cava The mortality of intertidal cirripedes 153 Margaret Barnes The biology of siphonariid limpets (Gastropoda: Pulmonata) 245 Alan N.Hodgson Author index 315 Systematic index 337 Subject index 345 v PREFACE This, the thirty-seventh volume in the series, contains the usual broad mix of topics covering physical and biological aspects of marine science. As in the past the contents of the articles range widely from global ocean processes through the ecological importance of particular groups to detailed reviews of the biology of individual genera and species that are of worldwide interest. Reviews such as these provide a great service to the marine science community by summarizing and evaluating the literature but they are time consuming to write. Consequently, the incentive to produce such articles may be less strong at a time when publication rates are considered, rightly or wrongly, to be one measure of scientific achievement. Fortunately, the steady influx of high quality manuscripts indicates the willingness of authors to compile reviews and ensures the continuing success of the series. As always, it is a pleasure for the editors to acknowledge the authors’ patience and co- operation in responding to editorial requests and the efficiency of the publishers in maintaining the regular appearance of these volumes. It is with great regret that we have to report the death of Dr A.D.Ansell during the final stages of the production of this volume. He was a long-standing colleague and friend and will be greatly missed. We dedicate this volume to his memory RNG, MB vii Oceanography and Marine Biology: an Annual Review 1999, 37, 1–45 © A.D.Ansell, R.N.Gibson and Margaret Barnes, Editors Taylor & Francis AIR-SEA GAS EXCHANGE INTO THE MILLENNIUM: PROGRESS AND UNCERTAINTIES THOMAS FROST & ROBERT C.UPSTILL-GODDARD Department of Marine Sciences and Coastal Management, University of Newcastle, Newcastle upon Tyne, NE1 7RU, UK. Abstract The current status of research into air-water gas exchange is reviewed. Some relevant basic concepts are re-examined within the context of current progress towards parameterization of the air-sea gas exchange process using models and data from wind-tunnels, and the interpretational difficulties that still persist with these approaches are identified and discussed. Subsequently, field measurements of air-water gas exchange are reviewed, with key developments highlighted, in particular, important recent advances in the use of micrometeorological and chemical tracer-based techniques of measurement. Results obtained with these methods are summarized and compared with laboratory wind-tunnel-based measurements, in order to identify some outstanding gas exchange issues still to be resolved. In the light of these, the potentially important roles played by a variety of physical, chemical and biological forcings are considered and some likely fruitful avenues for nature research are outlined for this important area of global science. Introduction The exchange of gases between oceans and atmosphere is a fundamentally important process in global biogeochemistry. For many gases of biogeochemical interest the oceans are a net atmospheric source; important examples include DMS (Charlson et al. 1987, Turner et al. 1996), NO (Law & Owens 1990, Nevison et al. 1995, Bange et al. 1996a,b), CH (Crutzen 2 4 1991, Owens et al. 1991, Bange et al. 1994), non-methane hydrocarbons (Plass-Dülmer et al. 1993), CO (Johnson & Bates 1996), COS (Andreae & Ferek 1992, Uher & Andreae 1997), and CS (Kim & Andreae 1992). These gases play important roles in atmospheric 2 chemistry and climate regulation and for all of them, the gas exchange rate itself controls directly the sea-to-air flux. Consequently, knowledge of air-sea gas exchange rates is directly relevant to a range of global issues, including the role of marine-derived aerosols in climate regulation (Schwartz 1988, 1989, Wigley 1989), relative source strengths of natural and anthropogenic acid rain sulphur (Nriagu et al. 1987, Turner et al. 1988), the oceanic source strengths of N O, CH, CO, and COS (Bange et al. 1994, Nevison et al. 1995, Johnson & 2 4 Bates 1996, Uher & Andreae 1997) and the role of the oceans in regulating atmospheric halocarbons (Butler 1994, Lobert et al. 1995, Nightingale et al. 1995). In contrast, for CO, for which the oceans are a net atmospheric sink, the situation is 2 somewhat more complex. The rate of CO uptake by the deep ocean is ultimately dictated 2 by the rate at which CO in surface waters can be exchanged vertically downward across 2 the thermocline; on average a doubling of the air-sea gas exchange rate only promotes an 1
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