THE INFLUENCE OF COASTAL UPWELLING ON THE BIODIVERSITY OF SANDY BEACHES IN SOUTH AFRICA Pamela Helen Cramb A Thesis Submitted for the Degree of PhD at the University of St Andrews 2015 Full metadata for this item is available in Research@StAndrews:FullText at: http://research-repository.st-andrews.ac.uk/ Please use this identifier to cite or link to this item: http://hdl.handle.net/10023/6553 This item is protected by original copyright This item is licensed under a Creative Commons Licence The influence of coastal upwelling on the biodiversity of sandy beaches in South Africa Pamela Helen Cramb This thesis is submitted in partial fulfilment for the degree of PhD at the University of St Andrews March 2015 1. Candidate’s declarations: I, Pamela Helen Cramb, hereby certify that this thesis, which is approximately 65,000 words in length, has been written by me, and that it is the record of work carried out by me, or principally by myself in collaboration with others as acknowledged, and that it has not been submitted in any previous application for a higher degree. I was admitted as a research student in February, 2010 and as a candidate for the degree of PhD in February, 2011; the higher study for which this is a record was carried out in the University of St Andrews between 2010 and 2014. Date ……………………… signature of candidate ………………………………………………………..………. 2. Supervisor’s declaration: I hereby certify that the candidate has fulfilled the conditions of the Resolution and Regulations appropriate for the degree of PhD in the University of St Andrews and that the candidate is qualified to submit this thesis in application for that degree. Date ……………......... signature of supervisor …………………………………………………………………. 3. Permission for publication: (to be signed by both candidate and supervisor) In submitting this thesis to the University of St Andrews I understand that I am giving permission for it to be made available for use in accordance with the regulations of the University Library for the time being in force, subject to any copyright vested in the work not being affected thereby. I also understand that the title and the abstract will be published, and that a copy of the work may be made and supplied to any bona fide library or research worker, that my thesis will be electronically accessible for personal or research use unless exempt by award of an embargo as requested below, and that the library has the right to migrate my thesis into new electronic forms as required to ensure continued access to the thesis. I have obtained any third-party copyright permissions that may be required in order to allow such access and migration, or have requested the appropriate embargo below. The following is an agreed request by candidate and supervisor regarding the publication of this thesis: PRINTED COPY a) No embargo on print copy ELECTRONIC COPY a) No embargo on electronic copy Date ……………… signature of candidate …………………………………………………………… signature of supervisor …………………………………………………………… i Acknowledgements After 4 ½ years, 97 tons of sand, plenty of blood, sweat and tears, it is a pleasure to finally submit “The Thesis”. There are so many people who have helped with this project that it has often been said my acknowledgements would be as long as my thesis, but I’ll try to keep it brief! First of all, this project would not have been possible without the funding provided by the University of St Andrews, the Andrew Mellon Foundation, the South African Research Chairs Initiative of the Department of Science and Technology, and the National Research Foundation. I am overwhelmingly grateful to each funding body for their generous contribution. I also wish to thank South African National Parks for their co-operation and access to Namaqualand National Park. I am very grateful to Charles Griffiths and Tris Wooldridge, who took the time to verify my species identifications, and I would like to thank Bob Clarke and Paul Somerfield for their statistical advice. I’d like to say a massive thank you to both my supervisors, Dave and Christopher for being there with your wisdom, patience, support and encouragement through all the good and the bad! This project was a steep learning curve in many ways, with plenty of challenges, but you both remained remarkably calm in every crisis and helped “make a plan” as the South Africans say. I really appreciate everything you did for me! Beccy and Vicky, you guys are stars! Project design, pilot studies, troubleshooting, taxonomy, analysis and writing, you have been my support crew in each country – always there with a smile and a laugh to cheer me on. I couldn’t have done it without either of you, thank you! “Pams Adventure Fat Camp” was the best of times and the worst of times. What a beautiful country, but what a lot of sand! Never before has a bakkie been so tightly packed, or stuck in the sand so often! To all of those that took part at one time or another: Pati, Adam, Margaux, Gavin, Mathilde S, Al, Ele, Gareth, Nick, Vicky J, Tess, Isla, Lucas, Diane, Patrick W, Max, Austin, Precotia, Hlenge, Dave H, Melissa, Sox, Tristan, Sven, Mike, Patrick – Thank you, you lightened the load and taught me the ii value of good field assistants! To those who kept coming back for more: Simone, Linda, Gauthier, Mathilde B, Caroline, Frikkie, Jenny, Tim, Lindsay & Keaton – I am eternally grateful. You guys were my core crew and I REALLY couldn’t have done it without you. The mountain of sand didn’t stop in the field. All the help in the lab was really appreciated. For those not already mentioned: Mpmelelo, Gwen B, Emma, Claire, Jade, Sheldon and the school interns – Thank you! Jaqui and Simon, what can I say? You were there for so much of the field and lab work, always ready and willing, come rain or shine, and often going beyond the call of duty. Good assistant are difficult to find, but I found them in you guys – really, for all your hard work, help and support, I cannot thank you enough! Good admin staff also go a very long way, thank you Gwen, you made my life so much easier - you are amazing at your job, and breathe a breath of fresh air into the lab. It’s been a pleasure to be part of two lovely lab groups, SERG and CRG. I’ve made lots of friends and exchanged a lot of knowledge, eaten a lot of cake and drank a lot of tea! Thank you all for your help, laughter and comfort in times of need – I’m sad to leave! Armand, you have been amazing and I am 100 % sure that I could not have done this thesis without your endless love, support and understanding; not to mention your help in the field, in the lab, and drawing maps! You never failed to calm me down, never complained at my crazy working hours and so often had dinner on the table for me when I’d get home. I couldn’t ask for more, and thank you doesn’t quite seem enough! A huge thank you has to go to my family. You kept me grounded at all times (no it wasn’t a beach holiday) but never failed to give immense support and help whenever it was needed, with a special thanks to Fiona for helping with data entry to save my eyes, and Mum for checking my grammar! Finally, Mum and Dad, you have given me so much love, support and encouragement in every endeavour since the day I was born. There is no thank you big enough for that, but I hope I have made you proud. This is for you! iii Contents Declaration……………………………………………………………………………………………………………………i Acknowledgements…………………………………………………………………..…………………………………ii Contents…………………..…………………………………………………………………………………………………iv List of Tables………………………………………………………………………………………………………………..v List of Figures…………………………………………………………………………………………………..……….viii Abstract……………………………………………………………………………………………………………………….x Chapter 1: General Introduction……………………………………………………………………….......1 Chapter 2: Study sites…………………………………………………………………………………………..20 Chapter 3: The influence of upwelling on macrofaunal assemblages across multiple scales…………………………………………………………………………………….30 Chapter 4: Testing the design - small scale variability………………………………………...141 Chapter 5: Influence of temperature on the filtration rate and oxygen consumption of the sandy beach bivalve Donax serra (Röding)…………211 Chapter 6: General Discussion…………………………………………………………………………...236 References……………………………………………………………………………………………………………….250 Appendix 1: Taxonomic descriptions and catalogue…...............................................CD Appendix 2: Statistical outputs from Chapter 3……………………………………………….…….CD Appendix 3: Statistical outputs from Chapter 4…………………………………………………….CD iv List of Tables Table 2.1 Summary table of site locations ………………………………………………………………… 23 Table 3.1 Summary of physical characteristics ……………………………………………………. 54-56 Table 3.2 Species list; including taxonomic groupings, feeding guilds and developmental modes ………………………………………….………...............……….…..62-63 Table 3.3 Total species richness over the entire sampling period from each geographic area, and the total species richness from each region………………………………….. 64 Table 3.4 Summary of significant results from PERMANOVA and subsequent tests in all regions using presence/absence transformed data…..……………………………………67 Table 3.5 Total numbers of individuals collected from each geographical location over the entire 2 year sampling period, and totals per region……………………….……….69 Table 3.6 Summary of significant PERMANOVA results and subsequent tests from region A, based on 4th root transformed abundance data…………………………73-74 Table 3.7 Summary of significant PERMANOVA results and subsequent tests from region B, based on 4th root transformed abundance data…………………………74-75