OCEANOGRAPHY and MARINE BIOLOGY AN ANNUAL REVIEW Volume 47 OCEANOGRAPHY and MARINE BIOLOGY AN ANNUAL REVIEW Volume 47 Editors R.N. Gibson Scottish Association for Marine Science The Dunstaffnage Marine Laboratory Oban, Argyll, Scotland [email protected] R.J.A. Atkinson University Marine Biological 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 International Standard Serial Number: 0078-3218 CRC Press Taylor & Francis Group 6000 Broken Sound Parkway NW, Suite 300 Boca Raton, FL 33487-2742 © 2009 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-13: 978-1-4200-9421-3 (Hardcover) This book contains information obtained from authentic and highly regarded sources. Reasonable efforts have been made to publish reliable data and information, but the author and publisher cannot assume responsibility for the valid- ity of all materials or the consequences of their use. The authors and publishers have attempted to trace the copyright holders of all material reproduced in this publication and apologize to copyright holders if permission to publish in this form has not been obtained. If any copyright material has not been acknowledged please write and let us know so we may rectify in any future reprint. 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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 Contents Preface vii Erratum to “Biology of the planktonic stages of benthic octopuses” ix Roger Villanueva & Mark D. Norman Transregional linkages in the north-eastern Atlantic — an ‘end-to-end’ analysis of pelagic ecosystems 1 Clive Fox, Roger Harris, Svein Sundby, Eric Achterberg, J. Icarus Allen, John Allen, Alex Baker, Corina P.D. Brussaard, Paul Buckley, Elizabeth Cook, Stephen R. Dye, Martin Edwards, Liam Fernand, Peter Kershaw, Julian Metcalfe, Svein Østerhus, Ted Potter, Egil Sakshaug, Douglas Speirs, Erling Stenevik, Mike St. John, Frede Thingstad & Ben Wilson Zooplankton diel vertical migration — a review of proximate control 77 Jonathan H. Cohen & Richard B. Forward Jr Cephalopods in the north-eastern Atlantic: species, biogeography, ecology, exploitation and conservation 111 L.C. Hastie, G.J. Pierce, J. Wang, I. Bruno, A. Moreno, U. Piatkowski & J.P. Robin Exploitation and conservation of echinoderms 191 J. Micael, M.J. Alves, A.C. Costa & M.B. Jones Anchovy and sardine in the Adriatic Sea — an ecological review 209 E. Betulla Morello & Enrico Arneri A review of underwater stereo-image measurement for marine biology and ecology applications 257 Mark Shortis, Euan Harvey & Dave Abdo Author Index 293 Systematic Index 295 Subject Index 297 v Preface The 47th volume of this series contains six reviews written by an international array of authors that, as usual, range widely in subject and taxonomic and geographic coverage. The editors welcome sug- gestions 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 time- table for submission, evaluation and acceptance of manuscripts, potential contributors are advised to make contact with 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 CRC Press, especially Marsha Hecht, in ensuring the timely appearance of this volume. vii Erratum to “Biology of the planktonic stages of benthic octopuses” ROger Villanueva1 & Mark D. NOrman2 1Institut de Ciències del Mar (CSIC), Passeig Marítim de la Barceloneta 37-49, E-08003 Barcelona, Spain E-mail: [email protected] 2Sciences, Museum Victoria, GPO Box 666, Melbourne, Vic 3001, Australia E-mail: [email protected] Refers to: Villanueva, R. & Norman, M.D. 2008. Biology of the planktonic stages of benthic octo- puses. Oceanography and Marine Biology: An Annual Review 46, 105–202. The publisher regrets the error introduced after proofreading in the scales of Figures 20 and 25 in the above paper. The corrected figures are reproduced as follows: ix Erratum To “BIOLOGY OF the planktOnic stages OF benthic OctOpuses” A BCD 50µm 10µm 10µm 5µm E FGH 50µm 50µm 5µm 5µm Figure 20 Statoliths of Octopus vulgaris paralarvae. Scanning electron micrographic images from antero- lateral (A) and posterior (C) views of hatchling statoliths with their respective crystalline surface structure presented inside the rectangles (B, D). In paralarvae aged 30 days, statolith growth is observed on the posterior side of the statolith (E, F). The crystalline structure of the surface observed inside the lower (G) and upper (H) rectangle of the image F is also indicated. Individuals obtained from rearing experiments described in Villanueva et al. (2004). Original. x Erratum To “BIOLOGY OF the planktOnic stages OF benthic OctOpuses” A B C Figure 25 Denticulation of beaks in Octopus vulgaris paralarvae. Scanning electron micrographic images of (A) oral view of hatchling; (B) 50-day-old specimen in presettlement stage, 7.3 mm mantle length (ML) (fresh); and (C) 60-day-old recently settled individual of 9.3 mm ML (fresh). Note the broken denticles on the lower beaks of posthatching individuals and the rostral tip of the beak in the settled individual, in transition to the typical adult beak form. Individuals obtained from rearing experiments described in Villanueva (1995). Original. xi Erratum To “BIOLOGY OF the planktOnic stages OF benthic OctOpuses” References Villanueva, R. 1995. Experimental rearing and growth of planktonic Octopus vulgaris from hatching to settle- ment. Canadian Journal of Fisheries and Aquatic Sciences 52, 2639–2650. Villanueva, R., Riba, J., Ruiz-Capillas, C., González, A.V. & Baeta, M. 2004. Amino acid composition of early stages of cephalopods and effect of amino acid dietary treatments on Octopus vulgaris paralarvae. Aquaculture 242, 455–478. xii Oceanography and Marine Biology: An Annual Review, 2009, 47, 1-76 © R. N. Gibson, R. J. A. Atkinson, and J. D. M. Gordon, Editors Taylor & Francis TransregiOnal linkages in the NOrth-eastern Atlantic — an ‘end-TO-end’ analYsis OF pelagic ecOSYstems Clive FOX1, ROger Harris2, Svein SundbY11, Eric Achterberg4, J. Icarus Allen2, JOhn Allen4, AleX Baker5, COrina P.D. Brussaard6, Paul BuckleY7, EliZabeth J. COOK1, Stephen R. DYE7, Martin Edwards8, Liam Fernand7, Peter Kershaw7, Julian Metcalfe7, Svein Østerhus3, Ted POtter7, Egil Sakshaug9, DOuglas Speirs10, Erling Stenevik11, Mike ST. JOhn12, Frede Thingstad13 & Ben WilsON1 1Scottish Association for Marine Science, Dunstaffnage Marine Laboratory, Oban, PA37 1QA, UK 2Plymouth Marine Laboratory, Prospect Place, Plymouth, PL1 3DH, UK 3Bjerknes Centre for Climate Research, Allegata 55, N-5007 Bergen, Norway 4National Oceanography Centre, University of Southampton, Waterfront Campus, Southampton, SO14 3ZH, UK 5School of Environmental Sciences, University of East Anglia, Norwich, NR4 7TJ, UK 6NIOZ–Royal Netherlands Institute for Sea Research, P.O. Box 59, 1790 AB Den Burg, Texel, The Netherlands 7CEFAS, Pakefield Road, Lowestoft, NR33 0HT, UK 8SAHFOS, The Laboratory, Citadel Hill, Plymouth, PL1 2PB, UK 9Norwegian University of Science and Technology, Hogskoleringen 1, Gloshaugen, 7491 Trondheim, Norway 10University of Strathclyde, Livingstone Tower, Richmond Street, Glasgow, G1 1XH, UK 11Institute of Marine Research, Nordnesgate 50, P.O. Box 1870 Nordnes, 5817 Bergen, Norway 12Hamburg University, Institute for Hydrobiology and Fisheries Science, Olbersweg 24, 22767 Hamburg, Germany 13University of Bergen, Jahnebakken 5, P.O. Box 7800, N-5020 Bergen, Norway E-mail: [email protected] Abstract This review examines interregional linkages and gives an overview perspective on marine ecosystem functioning in the north-eastern Atlantic. It is based on three of the ‘systems’ considered by the European Network of Excellence for Ocean Ecosystems Analysis (EUR-OCEANS was established in 2004 under the European Framework VI funding programme to promote integra- tion of marine ecological