Oceanography and Marine Biology, An Annual Review, Volume 41: An Annual Review: Volume 41 (Oceanography and Marine Biology)

444 Pages·2003·5.58 MB·English

by  R. N. Gibson

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.
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