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[Oceanography and Marine Biology - an Annual Review] R. N OCEANOGRAPHY and MARINE BIOLOGY AN ANNUAL REVIEW Volume 44 7044_C000.fm Page ii Tuesday, April 25, 2006 1:51 PM OCEANOGRAPHY and MARINE BIOLOGY AN ANNUAL REVIEW Volume 44 Editors R.N. Gibson Scottish Association for Marine Science The Dunstaffnage Marine Laboratory Oban, Argyll, Scotland [email protected] R.J.A. Atkinson University Marine Biology Station Millport University of London Isle of Cumbrae, Scotland [email protected] J.D.M. Gordon Scottish Association for Marine Science The Dunstaffnage Marine Laboratory Oban, Argyll, Scotland [email protected] Founded by Harold Barnes Boca Raton London New York CRC is an imprint of the Taylor & Francis Group, an informa business CRC Press Taylor & Francis Group 6000 Broken Sound Parkway NW, Suite 300 Boca Raton, FL 33487-2742 © 2006 by R.N. Gibson, R.J.A. Atkinson and J.D.M. Gordon CRC Press is an imprint of Taylor & Francis Group, an Informa business No claim to original U.S. Government works Printed in the United States of America on acid-free paper 10 9 8 7 6 5 4 3 2 1 International Standard Book Number-10: 0-8493-7044-2 (Hardcover) International Standard Book Number-13: 978-0-8493-7044-1 (Hardcover) International Standard Serial Number: 0078-3218 This book contains information obtained from authentic and highly regarded sources. Reprinted material is quoted with permission, and sources are indicated. A wide variety of references are listed. Reasonable efforts have been made to publish reliable data and information, but the author and the publisher cannot assume responsibility for the valid- ity of all materials or for the consequences of their use. No part of this book may be reprinted, reproduced, transmitted, or utilized in any form by any electronic, mechanical, or other means, now known or hereafter invented, including photocopying, microfilming, and recording, or in any information storage or retrieval system, without written permission from the publishers. For permission to photocopy or use material electronically from this work, please access www.copyright.com (http:// www.copyright.com/) or contact the Copyright Clearance Center, Inc. (CCC) 222 Rosewood Drive, Danvers, MA 01923, 978-750-8400. CCC is a not-for-profit organization that provides licenses and registration for a variety of users. For organizations that have been granted a photocopy license by the CCC, a separate system of payment has been arranged. Trademark Notice: Product or corporate names may be trademarks or registered trademarks, and are used only for identification and explanation without intent to infringe. Visit the Taylor & Francis Web site at http://www.taylorandfrancis.com and the CRC Press Web site at http://www.crcpress.com 7044_Discl.indd 1 5/3/06 1:42:06 PM 7044_C000.fm Page v Tuesday, April 25, 2006 1:51 PM Contents Preface vii Correction to Volume 43 ix Review of three-dimensional ecological modelling related to the North Sea shelf system. Part II: model validation and data needs 1 Günther Radach & Andreas Moll Role, routes and effects of manganese in crustaceans 61 Susanne P. Baden & Susanne P. Eriksson Macrofaunal burrowing: the medium is the message 85 Kelly M. Dorgan, Peter A. Jumars, Bruce D. Johnson & Bernard P. Boudreau Mediterranean coralligenous assemblages: a synthesis of present knowledge 123 Enric Ballesteros Defensive glandular structures in opisthobranch molluscs — from histology to ecology 197 Heike Wägele, Manuel Ballesteros & Conxita Avila Taxonomy, ecology and behaviour of the cirrate octopods 277 Martin A. Collins & Roger Villanueva The ecology of rafting in the marine environment. III. Biogeographical and evolutionary consequences 323 Martin Thiel & Pilar A. Haye Potential effects of climate change on marine mammals 431 J.A. Learmonth, C.D. MacLeod, M.B. Santos, G.J. Pierce, H.Q.P. Crick & R.A. Robinson Author Index 465 Systematic Index 497 Subject Index 515 7044_C000.fm Page vi Tuesday, April 25, 2006 1:51 PM 7044_C000.fm Page vii Tuesday, April 25, 2006 1:51 PM Preface The forty-fourth 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 editors welcome suggestions from potential authors for topics they consider could form the basis of future 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 contact the editors at an early stage of preparation. Contact details are listed on the title page of this volume. The editors gratefully acknowledge the willingness and speed with which authors complied with the editors’ suggestions, requests and questions, and the efficiency of Taylor & Francis in ensuring the timely appearance of this volume. 7044_C000.fm Page viii Tuesday, April 25, 2006 1:51 PM 7044_C000.fm Page ix Tuesday, April 25, 2006 1:51 PM Correction to Volume 43 Carney, R.S. 2005. Zonation of deep biota on continental margins. Oceanography and Marine Biology: An Annual Review 43, 211–278. In reviewing various ideas put forward to explain deep benthic depth zonation (Carney 2005), the TROX, Trophic-Oxygen, model for foraminiferan distributions received special mention. This conceptual model incorporates in an easily understood manner several lines of thought concerning the importance of oxygen and labile carbon influx in controlling surficial and interstitial microhab- itats. The origin of the TROX model, however, is misattributed to Loubere and his associates (page 228). The TROX model was proposed in the context of microhabitats with Adriatic transect data used as an example by Jorissen et al. (1995). An important feature of the TROX model is the explicit link between oxygen and carbon flux, such that the ecological importance of food avail- ability changes with oxygen concentration. Given adequate oxygen, food is the primary factor. When microbial consumption of higher food levels reduces microhabitat oxygen, then low oxygen can become the primary factor. There have been other attempts to create general foraminiferan distribution models that incorporate both oxygen and carbon flux somewhat similar to TROX. An algebraically formal model of gradient distribution employing the concept of ‘r’ and ‘K’ selected species was developed by Sjoerdsma & van der Zwaan (1992) and tested with mixed success on archived distribution data from the Gulf of Mexico. Unfortunately, the geochemical relationship between oxygen and carbon flux was omitted. Bottom oxygen was estimated from water column profiles, and flux estimated only on the basis of depth. The conceptual model proposed by Loubere et al. (1993) was an assemblage model incorporating both production and taphonomy. Like the TROX model, it is an important contribution to the understanding of trophic control of geographic distribution. That mode links the geochemistry of oxygen and carbon flux. Flux was estimated from sedimentary oxygen consumption; samples were analysed from the western Gulf of Mexico. REFERENCES Carney, R.S. 2005. Zonation of deep biota on continental margins. Oceanography and Marine Biology: An Annual Review 43, 211–278. Jorissen, F.J., de Stigter, H.C. & Widmark, J.G.V. 1995. A conceptual model explaining benthic foraminiferal microhabitats. Marine Micropaleontology 26, 3–15. Loubere, P., Gary, A. & Lagoe, M. 1993. Generation of the benthic foraminiferal assemblage — theory and preliminary data. Marine Micropaleontology 20, 165–181. Sjoerdsma, P.G. & van der Zwaan, G.J. 1992. Simulating the effect of changing organic flux and oxygen content on the distribution of benthic Foraminifera. Marine Micropaleontology 19, 103–150. 7044_C000.fm Page x Tuesday, April 25, 2006 1:51 PM OCEANOGRAPHY and MARINE BIOLOGY AN ANNUAL REVIEW Volume 44 7044_C000.fm Page xii Tuesday, April 25, 2006 1:51 PM 7044_book.fm Page 1 Friday, April 14, 2006 1:28 PM Oceanography and Marine Biology: An Annual Review, 2006, 44, 1-60 © R. N. Gibson, R. J. A. Atkinson, and J. D. M. Gordon, Editors Taylor & Francis REVIEW OF THREE-DIMENSIONAL ECOLOGICAL MODELLING RELATED TO THE NORTH SEA SHELF SYSTEM. PART II: MODEL VALIDATION AND DATA NEEDS GÜNTHER RADACH & ANDREAS MOLL Institut für Meereskunde (IfM), Universität Hamburg (ZMK-ZMAW), Bundesstr. 53, D-20146 Hamburg, Germany E-mail: [email protected], [email protected] Abstract The aim of this review is to provide an overview of the status of validation of eleven biogeochemical and ecological models of the greater North Sea (COHERENS, CSM-NZB, DCM- NZB, DYMONNS, ECOHAM, ELISE, ERSEM, FYFY, GHER, NORWECOM, POLCOMS- ERSEM) showing the realism achieved as well as the problems hindering a better degree of validity of the models. Several of the models were able to reproduce observations of the state variables correctly within an order of magnitude, but all models are not capable of reproducing every simulated state variable in the range of observations. None of the models can be called a valid model. Comparison of results from different models with datasets are evaluated according to the different spatial and temporal scales, for which data products were available, namely for regional distributions, annual cycles, long-term developments and events. The higher the trophic level, the greater was the discrepancy with the data. Problems still exist in determining the necessary com- plexity of the ecosystem model. More complexity in the model does not necessarily improve the simulations. Special
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