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Open Research Online The Open University’s repository of research publications and other research outputs Large-scale forcing of coastal communities. Thesis How to cite: Shelmerdine, Richard (2007). Large-scale forcing of coastal communities. PhD thesis The Open University. For guidance on citations see FAQs. c 2007 Richard Shelmerdine https://creativecommons.org/licenses/by-nc-nd/4.0/ Version: Version of Record Link(s) to article on publisher’s website: http://dx.doi.org/doi:10.21954/ou.ro.0000fb2d Copyright and Moral Rights for the articles on this site are retained by the individual authors and/or other copyright owners. For more information on Open Research Online’s data policy on reuse of materials please consult the policies page. oro.open.ac.uk V y v FV s. i r-:. ! 4 ..../ i j Large-scale forcing of coastal communities Richard Shelmerdine M.Res, B.Sc (Hons) A thesis submitted to Open University in fulfilment of the requirement of the Degree of Doctor of Philosophy Marine Ecology UHI Millennium Institute Scottish Association for Marine Science OaU ^ wv\ : <^c(lcpr ZefP, 0 ^ c w t m j d 17 Ap-l 'XI f \ NATURAL nnnUH| SCOTTISH ENVIRONMENT ’• Millennium ASSOCIATION RESEARCH COUNCIL Institute for MARINE i e J SCIENCE i ProQuest Number: 13917225 All rights reserved INFORMATION TO ALL USERS The quality of this reproduction is dependent upon the quality of the copy submitted. In the unlikely event that the author did not send a com plete manuscript and there are missing pages, these will be noted. Also, if material had to be removed, a note will indicate the deletion. uest ProQuest 13917225 Published by ProQuest LLC(2019). Copyright of the Dissertation is held by the Author. All rights reserved. This work is protected against unauthorized copying under Title 17, United States C ode Microform Edition © ProQuest LLC. ProQuest LLC. 789 East Eisenhower Parkway P.O. Box 1346 Ann Arbor, Ml 48106- 1346 I hereby declare that this thesis is my own work. Where I have used the work of other persons or quoted the work of other persons, the sources of the other work of information have been detailed explicitly in the presentation. Richard Shelmerdine Abstract Two contrasting areas of pelagic primary productivity (PPP) were noted in western Scotland, the Clyde (high PPP) and the west coast (low PPP). It was hypothesised that increased PPP would have a direct bottom-up influence on intertidal community structure with the Clyde expected to be dominated by filter feeders with potentially greater larval recruitment, increased density, faster growth and larger maximum sizes. The study was divided into five sections examining community structure, growth rates, predation and grazing pressures, effects of wave exposure, and stable isotope analysis. Individual species tended to vary between sites within lochs rather than between the two regions. Growth rates of the predator Nucella were found to follow peaks in Semibalanus size, rather than Mytilus, with increased growth on the west coast. An increased density of the grazer, Littorina , at a site had an increased effect on their growth rate although dense localised patches within sites were observed where growth rates were lowest. Mussel size classes were found to have different growth rates, most probably due to differing factors such as predation, food availability, and reproduction. Predation and grazing effects differed between regions. Barnacle cover was shown to be affected by both Nucella and Littorina although the latter may have been an indirect affect due to the biofilm cover which was greater in the Clyde. Predation rates of mussels were found to be greater on the west coast which was most probably due to a change in diet from barnacles to mussels. Mussel shell length and biomass declined with increasing wave exposure throughout western Scotland with the potential for factors varying on small scales to be more important in structuring mussel populations. This was evident when testing for differences in stable isotopes of mussels which suggested site specific variation due to increased freshwater input. The results of this study showed that small scale, local factors are as important, if not more, as regional differences in structuring communities. PPP is important, but only for a subset of the community. Acknowledgements Although many months were spent in front of a computer screen carrying out what seemed like an endless number of various analyses which culminated in this written manuscript, none of this would have been possible if it were not for the many people that accompanied me, taking time out of their own busy schedules, on my various field trips to collect my data. In particular Linda Robb, Robin Harvey, and Jenny Beaumont who all voluntarily helped me on more than one occasion. I still wonder if you were all gluttons for punishment or interested in my work. I like to think it was the latter and thank you most sincerely for your help. Dr Rona McGill at SUERC was always willing to help out with the mussel stable isotope analysis, not just in helping me understand the protocols and running my samples but for helping in the interpretation of the results. Many thanks. Thanks to the Ormsary Estate, Loch Caolisport, for granting me site access. To all my fellow PhD students but in particular Adam, Craig, Jenny, and Susan, it was immensely helpful to be able to bounce ideas off one another and also to know that I was not alone during the past couple of years. Thanks to my thesis panel (Angela Hatton, Craig Brown, Maeve Kelly and the adopted Thom Nickell) and my third party monitor, Jack Matthews. Your ideas and suggestions were always helpful and much appreciated. My parents, friends, and of course Karen, who were always there when I needed them even if I wasn’t stressed but should have been! To those that I have forgotten, apologies but thank you. Of course, I owe a countless amount of thanks to both of my supervisors, Mike Burrows and Dave Hughes, who were incredibly patient with me and always had time to help out with my field work on countless occasions, talk over ideas, go through aspects of the work that I was having problems understanding, and reading over what I had written. I could not have asked for better supervisors. Contents Chapter 1 General Introduction ....................................................................................... 13 1.1 Oceanographic Processes..................................................................................... 15 1.1.1 Previous studies examining the effects of large-scale oceanographic conditions on small-scale intertidal community structure.........................16 1.1.2 Oceanographic processes in western Scotland ............................................17 1.1.3 Clyde system and its oceanographic processes ...........................................20 1.1.4 Scottish west coast and its oceanographic processes ................................. 20 1.2 Phytoplankton ...................................................................................................... 21 1.3 Aims of this study ................................................................................................23 Chapter 2 Colonisation and community structure of intertidal rocky shores in western Scotland ..........................................................................................................27 2.1 Introduction ...........................................................................................................28 2.1.1 Impact of disturbances on species succession and community structure 28 2.1.2 Settlement and recruitment processes of the intertidal ............................... 30 2.1.3 Intertidal barnacle species of western Scotland...........................................32 2.1.4 Aims and hypotheses .................................................................................... 36 2.2 Materials and Methods .........................................................................................38 2.2.1 Preliminary surveys of regional and local variation in community structure ........................................................................................................38 2.2.2 Design and establishment of colonisation and monitoring areas ..............38 2.2.3 Image analyses prior to estimation of abundances and length measurements .............................................................................................. 42 2.2.4 Determination of the abundance of major species ..................................... 43 2.2.5 Density, length, and growth of intertidal barnacle species using image analysis .........................................................................................................43 2.3 Results ...................................................................................................................45 2.3.1 Preliminary surveys of regional and local variation in community structure ........................................................................................................45 2.3.2 Long term variation in abundance of major intertidal species ..................48 2.3.3 Density variation in Semibalanus balanoides............................................ 68 2.3.4 Semibalanus balanoides size........................................................................71 2.3.5 Density variation in Chthamalus montagui.................................................79 2.3.6 Chthamalus montagui size..........................................................................