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University of Southampton Research Repository Eprints Soton University of Southampton Research Repository ePrints Soton Copyright © and Moral Rights for this thesis are retained by the author and/or other copyright owners. A copy can be downloaded for personal non-commercial research or study, without prior permission or charge. This thesis cannot be reproduced or quoted extensively from without first obtaining permission in writing from the copyright holder/s. The content must not be changed in any way or sold commercially in any format or medium without the formal permission of the copyright holders. When referring to this work, full bibliographic details including the author, title, awarding institution and date of the thesis must be given e.g. AUTHOR (year of submission) "Full thesis title", University of Southampton, name of the University School or Department, PhD Thesis, pagination http://eprints.soton.ac.uk UNIVERSITY OF SOUTHAMPTON FACULTY OF ENGINEERING, SCIENCE & MATHEMATICS SCHOOL OF OCEAN & EARTH SCIENCES Marine Communities of North Sea Offshore Platforms, and the Use of Stable Isotopes to Explore Artificial Reef Food Webs by Andrew James Guerin Thesis for the degree of Doctor of Philosophy September 2009 UNIVERSITY OF SOUTHAMPTON ABSTRACT FACULTY OF ENGINEERING, SCIENCE & MATHEMATICS SCHOOL OF OCEAN & EARTH SCIENCES MARINE COMMUNITIES OF NORTH SEA OFFSHORE PLATFORMS, AND THE USE OF STABLE ISOTOPES TO EXPLORE ARTIFICIAL REEF FOOD WEBS by Andrew James Guerin Stable isotope methods offer a powerful means of investigating trophic interactions, allowing assessment of the relative importance of multiple nutrient sources to biological assemblages, as well as estimation of the trophic positions of consumers. Differences in the isotope ratios of consumers between habitats can thus indicate differences in the structures of food webs, or the contributions of different food sources to those food webs. Isotope methods were used to compare the food web of an artificial reef located off the south coast of England with that of a nearby natural reef system, revealing a similarly complex food web, with similar trophic structure, and similar inputs from the available food sources. Isotope methods should be incorporated into more artificial reef studies, where they have been seldom applied. ***** Offshore oil and gas platforms in the North Sea are artificial reefs, hosting substantial assemblages of sessile invertebrates and other associated fauna, and attracting large numbers of fish and motile invertebrates. Structural survey footage provided by the oil and gas industry allowed the investigation of the marine life associated with several of these structures, of varied ages and in various locations in the North Sea. At least thirty‐six taxa of motile invertebrates and fish were observed in association with the structures, most of which were present on all platforms surveyed. While most reef‐associated fish were observed around the base of the larger platforms, many thousands of fish were also observed in the water column around these structures at other depths. A small number of sessile taxa dominated the fouling assemblages, in places achieving total coverage of the available surfaces. Fouling composition changed with depth, but this pattern was not identical on all platforms. Platform age and location both affected the fouling assemblages present, but these two factors did not fully explain all the variation. ii Table of Contents ABSTRACT ................................................................................................................................... ii LIST OF TABLES ............................................................................................................................ ix LIST OF FIGURES ........................................................................................................................ xiii DECLARATION OF AUTHORSHIP ................................................................................................ xvii ACKNOWLEDGEMENTS ............................................................................................................ xviii Chapter 1. Introduction to Artificial Reefs .................................................................................... 1 1.1 What is an artificial reef? .......................................................................................... 1 1.2 Artificial reefs and epibiota ....................................................................................... 1 1.2.1 Factors affecting fouling assemblages on artificial reefs ................................. 2 1.2.1.1 Location .......................................................................................... 2 1.2.1.2 Construction material .................................................................... 2 1.2.1.3 Surface orientation......................................................................... 3 1.2.1.4 Reef profile ..................................................................................... 4 1.2.1.5 Structural complexity ..................................................................... 4 1.2.1.6 Timing of deployment .................................................................... 4 1.2.2 Colonisation of artificial reefs by epibiota ....................................................... 4 1.3 Artificial reefs and fish .............................................................................................. 5 1.3.1 Fish populations at artificial reefs ................................................................... 5 1.3.2 Colonisation of reefs by fish ............................................................................ 6 1.3.3 Potential benefits of artificial reefs to fish ...................................................... 7 1.3.3.1 Food ............................................................................................... 7 1.3.3.2 Shelter ............................................................................................ 7 1.3.3.3 Other possible benefits .................................................................. 8 1.3.4 Factors affecting fish populations at artificial reefs ........................................ 8 1.3.4.1 Reef size ......................................................................................... 9 1.3.4.2 Epibiota .......................................................................................... 9 1.3.4.3 Arrangement of reef units .............................................................. 9 1.4 Effects of artificial reefs on surrounding environment ............................................... 9 1.4.1 Attraction versus Production ........................................................................... 9 1.4.1.1 The importance of the attraction‐production debate ................. 10 1.4.1.2 Is there evidence of production? ................................................. 11 1.4.2 Effects on the surrounding benthos .............................................................. 11 1.4.3 Artificial reefs and biological invasions ......................................................... 12 1.5 Artificial reef applications ....................................................................................... 12 1.5.1 Exploitation reefs (fishing reefs) .................................................................... 12 1.5.2 Reefs for fisheries management .................................................................... 13 1.5.3 Reefs for habitat enhancement and restoration ........................................... 14 1.5.4 Recreational reefs .......................................................................................... 14 1.5.6 Other functions .............................................................................................. 15 1.5.7 de facto artificial reefs ................................................................................... 15 1.6 Structure of thesis .................................................................................................. 16 PART I ................................................................................................................. 17 Chapter 2. Introduction to Part I: Investigating artificial reef food webs ..................................... 18 2.1 Food webs on artificial reefs ................................................................................... 18 2.1.1 Trophic transfer to reef fish ........................................................................... 18 2.1.2 Structure of reef food webs .......................................................................... 19 2.2 Stable isotopes in ecological research ..................................................................... 20 2.2.1 Introduction to isotope approach ................................................................. 20 2.2.2 Advantages of isotope approach ................................................................... 21 2.2.3 Applications ................................................................................................... 21 iii 2.3 Aims and objective of Part I .................................................................................... 22 2.4 The Poole Bay Artificial Reef ................................................................................... 22 Chapter 3. Stable isotope ratios of consumers on natural and artificial reefs .............................. 24 3.1 Introduction ..........................................................................................................
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