Detecting Responses of Rocky Shore Organisms to Environmental Change Following Wave Energy Extraction
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Detecting responses of rocky shore organisms to environmental change following wave energy extraction Andrew Want Submitted for the degree of Doctor of Philosophy Heriot-Watt University Institute of Life and Earth Sciences April 2017 The copyright in this thesis is owned by the author. Any quotation from the thesis or use of any of the information contained in it must acknowledge this thesis as the source of the quotation or information. Abstract: The highly energetic waters surrounding Orkney have recently attracted attention as a renewable resource to generate electricity. While the importance of wave exposure to littoral assemblages is well known, the ecological consequences of industrial-scale extraction of marine energy have not been directly studied on rocky shores. The aim of this study was to examine the potential consequences of wave energy extraction and other long-term forcing agents, such as climate change, to rocky shore assemblages. Baseline surveys were conducted in areas not previously described in scientific detail to serve as ‘before’ and ‘control’ sites in a BACI-style design. Composition and abundances of biological assemblages were compared with topographic measurement of coastal features expected to modify exposure through dissipation of incoming wave energy. Observed variation in assemblages between sites was explained by differences in exposure and topography, particularly substrate complexity. Rocky shore species were selected for monitoring long-term changes using a paired-species monitoring method, including key structuring organisms on high exposure shores. Monitoring and experimental manipulation identified species which respond to changes in wave energy extraction and temperature. This research will assist in elucidating ecological responses that might occur following removal of wave energy, amid warming seas. i To my Family… ii Acknowledgments: The work contained in these pages would not have been possible without the support of many people. First and foremost are my supervisors: Prof. Jon Side, who has been instrumental in establishing this project and serving as a font of ecological knowledge that has helped guide me in developing this work; Dr. Mike Bell, who has played a critical role in all aspects of this work, from field assistance to advice on statistical approaches, delivered with great patience, and whose precision in communication has helped make me a better writer; and Dr. Jo Porter, whose infectious enthusiasm for marine biology continues to inspire me past challenging obstacles, and who has helped provide me with invaluable opportunities to broaden my research. I would like to thank UKCMER SuperGen for sponsoring this studentship and to the administrative team at the Orkney campus of Heriot-Watt University for their essential support: Kate Pirie, PJ Dewar, Erlend Leslie, Fiona Currie, Cat Morrison, and Eileen McKay. Over the years, many folk have contributed their valuable time to join me in the field even if only to shout ‘wave!’ periodically to ensure my safety. So, thank you to Bob Beharie, Dr. Susana Baston Meira, Dr. Mel Broadhurst, and Matt Coleman, with extra thanks to the latter for assistance with GIS. Best wishes to my office-mates who have supported me over this last period and whom I hope to offer my congratulations on their own PhD completions in the not too distance future: Rob Hiley, Beta Rodrigues, Kate Rydzkowski, Simon Waldman, and Steph Weir. My field work would not have been possible without boat-based deployments, provided by the team from SULA Diving: Kevin Heath, Bertrand Belafonte Taylor, and Dr. Malcolm Thomson. Land-based field work at times required the kind support of local landowners who provided access, sometimes bemusement, an occasional jar of home brew, but mostly, genuine curiosity in this work. Special thanks to Morag iii MacInnes, John McGill, and John Liptrot for your interest in Fucus distichus. Thanks to the team led by Prof. Mike Burrows, with Robin Harvey and Dr. Gail Twigg, for the opportunity to complement Orkney studies with field work on Lewis. Additional field assistance in Orkney was provided by Nuffield students: Tansy Branscombe, Alice Farnie, and Emma Hourston, with supervisory collaboration from my UHI colleague Dr. Ingrid Mainland. Various esteemed colleagues have imparted knowledge, advice, or suggestions which have in some manner informed this document. Thank you for the contribution: Dr. Raeanne Miller, Dr. João Neiva, Dr. Anne Marie Power, Prof. Ester Serrão, and lastly, the legendary phycologist Harry Powell, of whom I had the great pleasure to meet shortly before his death in January 2016. Special thanks to Anne Bignall for the excellent schematic drawings featured in Figure 3.3.1.2, and to Dr. Owen Brimijoin and Katherine Fryer for proof-reading and improvements to the Introduction chapter. Finally, thank you to the examiners of this thesis for their time in considering this work. If I have omitted anyone who has contributed but remains unmentioned, please forgive my oversight. iv v Table of contents: Abstract .............................................................................................................................. i Dedication ......................................................................................................................... ii Acknowledgements .......................................................................................................... iii Declaration Statement ....................................................................................................... v List of abbreviations ....................................................................................................... xiv 1. Introduction ......................................................................................................... 1 1.1 Orkney and marine renewable energy............................................................. 1 1.2 Ecological impacts and the marine renewable energy industry ...................... 7 1.3 Rocky shore ecological studies ..................................................................... 10 -Community classification ...................................................................... 11 1.4 Littoral zonation ............................................................................................ 12 -Fucoid-barnacle-patella complex .......................................................... 15 1.5 Rocky shore community and exposure ......................................................... 17 -Morphological adaptations to wave exposure ....................................... 19 -Consequences of energy extraction ....................................................... 21 1.6 Climate change .............................................................................................. 22 -Climate change on rocky shores ............................................................ 26 -Recent cooling periods........................................................................... 28 1.7 SuperGen Marine Energy Consortium .......................................................... 30 1.8 Road map to the current studies .................................................................... 30 vi 2 Setting the scene: methodological development of long-term littoral studies on West Mainland, Orkney ......................................................................................... 34 2.1 Introduction ................................................................................................... 34 -Littoral studies in Orkney ...................................................................... 34 -Wave exposure on WMO ...................................................................... 36 -Biological responses to environmental changes .................................... 37 -Detecting responses against a background of other environmental changes .................................................................................................... 40 2.2 Rationale and preliminary surveys ................................................................ 41 2.2.1 Selection of indicator species ...................................................... 43 2.2.2 Candidate species ........................................................................ 45 2.2.3 Littoral site selection ................................................................... 48 2.2.4 Selected sites ............................................................................... 49 2.3 Meteorological data ....................................................................................... 50 2.4 Discussion ..................................................................................................... 53 3 Before impact and control assessment of WMO: the relationship between topography, wave exposure, and the littoral community .................................... 56 3.1 Introduction ................................................................................................... 56 -Measuring exposure at ecologically meaningful scales ......................... 56 -Topography and exposure ...................................................................... 59 -Topographic variables............................................................................ 60 3.1.1 Objectives .................................................................................... 65 3.1.2 Hypotheses .................................................................................. 66 3.2