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Studies in the Stable Isotope Ecology of Coral- Reef Fish Food-Webs

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Studies in the Stable Isotope Ecology of Coral- Reef Fish Food-Webs Studies in the stable isotope ecology of coral- reef fish food-webs Yiou Zhu School of Natural and Environmental Sciences Newcastle University Submitted to Newcastle University for the degree of Doctor of Philosophy April 2019 i Studies in the stable isotope ecology of coral-reef fish food-webs Yiou Zhu Supervisors: Prof N Polunin, Dr W Reid, Dr S Newman Examiners: Prof N Graham and Prof J Bythell Abstract Coral-reef fish food-webs are complex and few studies have explored the range of production sources or how these support the higher trophic levels of coral-reef food- webs. I collected muscle tissue samples of abundant coral-reef fish at Bahamian and Maldivian sites and used bulk stable isotope (δ13C and δ15N) data in bi-plot to analyse their isotopic niches in relation to their putative trophic guilds; some species (e.g. some corallivorous Chaetodontidae) are evidently confined to their guild, but others seem to be utilizing multiple food sources. I also analysed δ15N (as a proxy for trophic position) to body size relationships among all individuals, individuals of the same species, individuals sharing the same trophic pathway and of the community as a whole. At species, some fishes had relatively flat relationships whereas others varied greatly with size. Individuals relying on the same production source type (e.g. planktivorous Chaetodontidae and Acanthuridae, Maldivian data only) had similar body size-δ15N trends with variation potentially due to at least family-related traits. At community level, there was a positive linear relationship in the Bahamas sites but a parabolic relationship in the Maldives, suggesting trophic function changes at different size classes in different ways in the two locations. I further tested whether the putative trophic guilds precisely portrayed the strict feeding patterns among fishes and whether location affected source partitioning using both bulk (SIA) and compound-specific stable isotope data (CSIA). The results suggested that certain consumers (e.g. corallivores [SIA] and detritivores [SIA and CSIA]) were confined to the trophic guild to which they have typically been assigned, while others show some evidence of relying on multiple sources (e.g. diurnal planktivores [SIA and CSIA]) indicating that it is imprecise to assign them to the single guilds to which they have commonly been assigned. At outer-atoll sites, some i fish fed more on diurnal plankton than those in the inner-atoll suggesting some geographic characteristics could affect the feeding preferences. I have applied stable isotope data to elucidate the importance of body size and geographic location in determining the feeding strategies of fish to better understand how these diverse food-webs work. The study provides pointers to future work such as on benthic-pelagic coupling, roles of sponges, contribution of SIA vs CSIA data and finer-detailed diet composition. ii Acknowledgements I would like to thank those who provided support, resources and funding of this PhD. First, I am grateful for the guidance, teaching and patience of my supervisory team: Prof Nicholas Polunin, Dr William Reid and Dr Steven Newman. They have been involved in the planning, preparing, conducting and writing up of both fieldwork and individual chapters. Prof Nicholas Polunin has helped me set up clear objectives for each project, gain underwater visual census and fish sampling skills in the field, carefully monitor the research progress, patiently and relentlessly improve my scientific writing style, and prepare papers from the work for publication. Dr William Reid has been highly involved in the statistical design of the work from the site selection and sample collection to data analysis, and thesis and paper writing. Dr Steven Newman was involved in the fieldwork design, data analysis, writing and publication of both projects. I thank them all. The Bahamian project was with Cape Eleuthera Institute. I would like to thank Dr Edward Brooks, Dr Annabelle Brooks and Dr Jocelyn Curtis-Quick for providing accommodation, permits and constant support during and after the fieldwork. In addition, I would like to share gratitude for interns at Cape Eleuthera Institute during the fieldwork, who helped with fish surveys (Jordan Atherton) and sample collection (Will Clark, Dom Ruddock and Alex Von Roenn). The Maldivian project was a collaboration with Banyan Tree Marine Labs (Maldives). I would like to thank Dr Steven Newman for supporting me tremendously in the Maldives by providing accommodation, workspace, diving gear and logistics. In addition, I would like to thank Christina Skinner and Banyan Tree Marine Labs interns for helping with surveying and sample collection: Aru, Ali, Shameem, Mode and Nadia. Also, I would like to thank Prof John Bythell for identifying coral species collected in this project. The bulk stable isotope analysis was funded by Newcastle University and analysed by Iso-Analytical Ltd. The compound-specific stable isotope analysis was funded by Newcastle University (sample preparation) and the NERC LSMSF at the University of Bristol (training and IRMS). I would like to thank Alison Kuhl at the School of Chemistry in Bristol for training me in the compound-specific isotope analysis step by iii step and monitoring my progress when I set up the preparation producers at Newcastle University. Also, I appreciate the workspace and nitrogen gas provided by the School of Geoscience at Newcastle University (Prof Martin Jones and David Earley). Finally, I would like to thank my friends for keeping me mentally and physically well throughout (Sheri, Martin, Chris and Glenn), and most importantly, my parents in funding me with my PhD study for these five years. Their support and encouragement kept me going and staying focused, and nothing can beat that. iv Table of Contents Studies in the stable isotope ecology of coral-reef fish food-webs .. i Abstract .................................................................................................. i Acknowledgements ............................................................................. iii Table of Contents ................................................................................. v List of Tables ........................................................................................ x List of Figures .................................................................................... xiii Acronyms and Symbols ................................................................... xvii Chapter 1. Coral-reef fish community trophic structure and source partitioning… ........................................................................................ 1 1.1 Introduction ................................................................................................. 1 1.2 Size structuring of coral-reef fish community .......................................... 2 1.2.1 Size spectra .......................................................................................... 3 1.2.2 Trophic structure ................................................................................. 4 1.3 Diverse production sources and feeding strategies ................................ 5 1.3.1 Benthic production sources ............................................................... 5 1.3.2 Pelagic production sources ................................................................ 8 1.4 Energy pathways......................................................................................... 9 1.4.1 Algivory ................................................................................................ 9 1.4.2 Microphagy ........................................................................................... 9 1.4.3 Detritivory ........................................................................................... 10 1.4.4 Corallivory .......................................................................................... 10 1.4.5 Spongivory ......................................................................................... 10 1.4.6 Planktivory.......................................................................................... 10 1.4.7 Zoobenthivory .................................................................................... 11 1.4.8 Piscivory ............................................................................................. 11 1.5 Feeding strategies of coral-reef fish ....................................................... 12 1.6 Diet analysis methods .............................................................................. 12 1.7 Bio-tracer diet analysis ............................................................................. 15 1.7.1 Analysis approaches ......................................................................... 15 1.7.2 Trophic position estimation .............................................................. 18 1.8 Geography ................................................................................................. 19 1.9 Objectives of the thesis ............................................................................ 19 Chapter 2. Size structuring and trophodynamics of a Bahamian coral-reef fish community* ................................................................ 21 2.1 Introduction ............................................................................................... 21 2.2 Materials and methods ............................................................................. 23 v 2.2.1 Study site ..........................................................................................
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