Oceanography and Marine Biology, An Annual Review, Volume 41: An Annual Review: Volume 41 (Oceanography and Marine Biology) PDF
Preview Oceanography and Marine Biology, An Annual Review, Volume 41: An Annual Review: Volume 41 (Oceanography and Marine Biology)
OCEANOGRAPHY and MARINE BIOLOGY AN ANNUAL REVIEW Volume 41 OCEANOGRAPHY and MARINE BIOLOGY AN ANNUAL REVIEW Volume 41 Editors R.N. Gibson The Dunstaffnage Marine Laboratory Oban, Argyll, Scotland [email protected] R.J.A. Atkinson University Marine Biological Station Millport, Isle of Cumbrae, Scotland [email protected] Founded by Harold Barnes First published 2003 by Taylor & Francis 11 New Fetter Lane, London EC4P 4EE Simultaneously published in the USA and Canada by Taylor & Francis Inc 29 West 35th Street, New York, NY 10001 Taylor & Francis is an imprint of the Taylor & Francis Group This edition published in the Taylor & Francis e-Library, 2004. ©2003 R.N. Gibson and R.J.A. Atkinson 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 andaccurate 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 A catalog record for this book has been requested ISBN 0-203-18057-7 Master e-book ISBN ISBN 0-203-23081-7 (Adobe eReader Format) ISBN 0-415-25463-9 (Print Edition) CONTENTS Preface vii Oxygen minimum zone benthos: adaptation and community response to hypoxia 1 Lisa A. Levin Antarctic marine benthic diversity 47 Andrew Clarke & Nadine M. Johnston Influence of marine allochthonous input on sandy beach communities 115 I. Colombini & L. Chelazzi The effects of sedimentation on rocky coast assemblages 161 Laura Airoldi Exotic molluscs in the Mediterranean basin: current status and perspectives 237 Serge Gofas & Argyro Zenetos Detritus in the epilithic algal matrix and its use by coral reef fishes 279 Shaun K. Wilson, David R. Bellwood, J. Howard Choat & Miles J. Furnas Ecology of whale falls at the deep-sea floor 311 Craig R. Smith & Amy R. Baco The diet of harbour porpoise (Phocoena phocoena) in the northeast Atlantic: areview 355 M.B. Santos & G.J. Pierce Author index 391 Systematic index 419 Subject index 431 v PREFACE The forty-first volume of this series contains eight reviews written by an international array of authors that, as usual, range widely in subject and taxonomic and geographic coverage. The majority of articles were solicited but the editors always welcome suggestions from potential authors for topics they consider could form the basis of appropriate contributions. Because an annual publication schedule necessarily places constraints on the timetable for submission, evaluation and acceptance of manuscripts, potential contributors are advised to make contact with the editors at an early stage of preparation so that the delay between sub- mission and publication is minimised. In line with recent developments in electronic publishing, it is a pleasure to announce that the series will now be available in eBook format. Details can be found at the publisher’s eBookstore at http://www.ebookstore.tandf.co.uk The editors again gratefully acknowledge the willingness and speed with which authors complied with the editors’ suggestions, requests and questions and the efficiency of the copy editor and publishers in ensuring the regular annual appearance of each volume. vii Oceanography and Marine Biology: an Annual Review2003, 41, 1–45 ©R.N. Gibson and R.J.A. Atkinson, Editors Taylor & Francis OXYGEN MINIMUM ZONE BENTHOS: ADAPTATION AND COMMUNITY RESPONSE TO HYPOXIA LISA A. LEVIN Integrative Oceanography Division, Scripps Institution of Oceanography, La Jolla, CA 92093-0218 USA e-mail: [email protected] Abstract Mid-water oxygen minima ((cid:1)0.5mll(cid:2)1 dissolved O) intercept the continental 2 margins along much of the eastern Pacific Ocean, off west Africa and in the Arabian Sea and Bay of Bengal, creating extensive stretches of sea floor exposed to permanent, severe oxygen depletion. These seafloor oxygen minimum zones (OMZs) typically occur at bathyal depths between 200m and 1000m, and are major sites of carbon burial along the continental margins. Despite extreme oxygen depletion, protozoan and metazoan assemblages thrive in these envi- ronments. Metazoan adaptations include small, thin bodies, enhanced respiratory surface area, blood pigments such as haemoglobin, biogenic structure formation for stability in soupy sedi- ments, an increased number of pyruvate oxidoreductases, and the presence of sulphide-oxidising symbionts. The organic-rich sediments of these regions often support mats of large sulphide- oxidising bacteria (Thioploca, Beggiatoa, Thiomargarita), and high-density, low-diversity meta- zoan assemblages. Densities of protistan and metazoan meiofauna are typically elevated in OMZs, probably due to high tolerance of hypoxia, an abundant food supply, and release from predation. Macrofauna and megafauna often exhibit dense aggregations at OMZ edges, but depressed densities and low diversity in the OMZ core, where oxygen concentration is lowest. Taxa most tolerant of severe oxygen depletion ((cid:1)0.2mll(cid:2)1) in seafloor OMZs include calcare- ous foraminiferans, nematodes, and annelids. Agglutinated protozoans, harpacticoid copepods, and calcified invertebrates are typically less tolerant. High dominance and relatively low species richness are exhibited by foraminiferans, metazoan meiofauna, and macrofauna within OMZs. At dissolved oxygen concentrations below 0.15mll(cid:2)1, bioturbation is reduced, the mixed layer is shallow, and chemosynthesis-based nutrition (via heterotrophy and symbiosis) becomes important. OMZs represent a major oceanographic boundary for many species. As they expand and con- tract over geological time, OMZs may influence genetic diversity and play a key role in the evo- lution of species at bathyal depths. These ecosystems may preview the types of adaptations, species, and processes that will prevail with increasing hypoxia over ecological and evolution- ary time. However, many questions remain unanswered concerning controls on faunal standing stocks in OMZs, and the physiological, enzymatic, metabolic, reproductive and molecular adap- tations that permit benthic animals to live in OMZs. As global warming and eutrophication reduce oxygenation of the world ocean, there is a pressing need to understand the functional consequences of oxygen depletion in marine ecosystems. 1
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