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University of Cape Town The copyright of this thesis vests in the author. No quotation from it or information derived from it is to be published without full acknowledgementTown of the source. The thesis is to be used for private study or non- commercial research purposes only. Cape Published by the University ofof Cape Town (UCT) in terms of the non-exclusive license granted to UCT by the author. University EFFECTS OF HABITAT TRANSFORMATION ON TEMPORARY WETLANDS IN THE SOUTH-WESTERN CAPE, SOUTH AFRICA by Matthew S. Bird Thesis submitted for the degree of Doctor of Philosophy in Zoology at the University of Cape Town Town Faculty of ScienceCape University ofof Cape Town February 2012 University DECLARATION This thesis reports original research carried out in the Department of Zoology, University of Cape Town, between 2007 and 2012. It has not been submitted in whole or in part for a degree at any other university. The data presented here are my own. A subset of the data from chapters 2 and 3 was also used by M. Mlambo for a separate and unrelated analysis of diversity patterns of macroinvertebrate assemblages in temporary wetlands (M.Sc dissertation, UCT). Any assistance I have received is fully acknowledged. M. S. Bird Town Several publications are affiliated with the data collected during this thesis. The data from chapters 2 and 3 contributed to the following publications: Bird, M.S. (2010). Aquatic invertebrates as indicatorsCape of human impacts in South African wetlands. WRC Report no. TT 435/09, Waterof Research Commission, Pretoria. 144pp. Bird, M.S. and J.A. Day. (2010). Investigating the use of aquatic macroinvertebrates as indicators of human impairment of seasonal wetlands in the Western Cape Province, South Africa. Pages 323-328 in P. Fraga I Arguimbau, editor. International Conference on Mediterranean Temporary Ponds. Proceedings & Abstracts. Consell Insular de Menorca. Recerca, 14. Maó, Menorca.University Mlambo, M.C., M.S. Bird, C.C. Reed and J.A. Day. (2011). Diversity patterns of temporary wetland macroinvertebrate assemblages in the south-western Cape, South Africa. African Journal of Aquatic Science 36:299-308. II TABLE OF CONTENTS Acknowledgements IV Abstract VI Chapter 1. General introduction 1 Chapter 2. The influence of habitat transformation on environmental conditions 11 in temporary wetlands Chapter 3. The influence of habitat transformation on aquatic invertebrate 56 assemblages in temporary wetlands Town Chapter 4. Effects of alien vegetation invasion on temporary wetlands: 125 a case study of the Kenilworth Racecourse Conservation Area, Cape Town (South Africa) Cape Chapter 5. Synthesis of 185 References 195 Appendices 208 University III ACKNOWLEDGEMENTS I wish to thank a number of people who made this project possible and without whom the fieldwork and sample processing could not have been completed. Big thanks are in order to Musa “the wet hound” Mlambo for fieldwork assistance during 2007 and help with laboratory processing of samples. Jeremy Shelton was my fieldwork assistant for the sampling at Kenilworth during 2008 and 2009 and taught me how to think like a naturalist, as well as the art of the long coffee break! For further help with processing and enumeration of invertebrate samples I would like to thank Candice Daly, Emma Gray, Jon Aronson and Alanna Rebelo. Nutrient analyses were performed by Craig Attwood in the Department of Oceanography (Dr Howard Waldron lab), University of Cape Town (UCT), and I am very grateful for this assistance. Thanks to Liesl Phigeland for help with processing chlorophyll a samples and organising sampling equipment for me; you are one of the warmest and most accommodating people I have ever worked with. My sincere Townthanks to Maya Beukes, the reserve manager at Kenilworth Racecourse Conservation Area, for granting access to the grounds for my fieldwork and providing me with extensive information on this unique fragment of lowland fynbos habitat. Cape Specialist expertise was called upon in theof process of getting good resolution taxonomic identifications of invertebrates and in this regard I wish to thank: Dr Zsolt Gido, University of Debrecan, Hungary, (Ostracoda); Dr Denise Schael, NMMU, (Copepoda and Cladocera); Prof. M. Hamer, UKZN, (Anostraca); Prof. C.C. Appleton, UKZN, (Mollusca); Mr P. Reavell, Stellenbosch University, (Hemiptera and Coleoptera); Prof. M. Coetzee, Wits, (Culicidae); Dr D. Perkins, Harvard University, USA, (Hydraenidae); Mr P. Simaka and Prof. M. Samways, Stellenbosch University,University (Odonata); Prof. C. Griffiths, UCT, (Amphipoda); Assoc. Prof. J. Day, UCT, (Conchostraca); Dr E. Ueckerman, ARC, Pretoria, and Dr R. Gerecke, Tubingen, Germany, (Water mites); and Dr P. Cranston UCD, USA, (who provided a CD for identifying chironomids). To the members of the Freshwater Research Unit of the Zoology Department, you have been like a family to me over the years and I have thoroughly enjoyed working with each and every one of you: Musa Mlambo, Jeremy Shelton, Sean Marr, Fynn Corry, Alistair Fyfe, Heather Malan, Justine Ewart-Smith, Mao Amis, Helen Dallas, Dean Ollis, Cecile Reed, Vere RossGillespie. For comments on the draft of my thesis I wish to thank Assoc. Prof. Coleen Moloney and Dr Deena Pillay (Zoology Department, UCT). This work was funded by grants from the Water Research Commission (WRC) and the National Research Foundation IV (NRF), to whom I extend every thanks for giving me the opportunity to learn how to do freshwater ecological research. This thesis was affiliated with the WRC Wetland Health and Importance Programme, within which my project entitled Aquatic invertebrates as indicators of human impacts in South African wetlands was administered by Dr Heather Malan whose cracking whip made sure I got deliverables finished at least “reasonably” on time. Heather, thanks so much for your support, guidance and good company over the years and for providing me with this opportunity to do research. In connection with the WRC project, I also wish to thank Dr Liz Day who provided invaluable guidance and technical support on the topic. Liz, you have been my mentor from as early as my Honours year, when you first inspired me to study wetland ecology. Hopefully all your help over the years will pay off soon! Special thanks are due to my supervisor Assoc. Prof. Jenny Day for her support and encouragement over the years. Jenny, you have always been a warm, easy-going person and a mentor for me. It has been a pleasure studying under your guidance. Town To my parents, for your love and support through thick and thin. You have always encouraged me to follow my own ambitions and make my own decisions in life and for that I am deeply grateful. To Claire, for your love and friendship during the years of this thesis and for generally putting up with my PhD worries and Capeobsessions. To Jem, for your support and belief in me even at times when I have struggledof to believe in myself. Without the support and encouragement of those mentioned above I could not have finished this thesis. University V Name: Matthew S. Bird Thesis title: Effects of habitat transformation on temporary wetlands in the south-western Cape, South Africa Date: February 2012 ABSTRACT The widespread loss of temporary wetlands to human activities has been well established, but virtually no information exists on how the human transformation of landscapes has altered the ecological character of those wetlands that remain. This thesis investigates environmental conditions and invertebrate assemblages in temporary wetlands in relation to the extent of habitat transformation in the adjacent landscape, using a broad (across multiple landscapes) and a fine (within-landscape) scale of analysis. For the broad-scale studies, I hypothesized firstly that habitat transformation around wetlands is associated with changes in physico-chemical conditions and biotope characteristics in wetlands, and secondly, that invertebrate assemblages are resilient to these human-induced environmental changes because they are well adapted to naturally disturbed environments. Ninety isolated depression wetlands situated on coastal plains of the south-western Cape were sampled for environmental variables, macroinvertebrates and microcrustaceans. Linear regression models were used to relate environmental and invertebrate components to the surrounding extent of habitat transformation. Wetland physico-chemistry was significantly related to the extent of habitat transformation around wetlands,Town but there was no evidence for an effect of habitat transformation on the biotope characteristics or hydro-morphometry of wetlands. Relationships between invertebrate assemblage composition and gradients of habitat transformation were weak, but provided some indication that habitat transformation exerted an influence on invertebrate assemblages. Natural variations in environmental conditions,Cape particularly physico-chemistry, had a strong influence on aquatic invertebrates. of It was expected that at a fine scale of analysis, where natural influence is considerably reduced, the effects of habitat transformation on temporary wetlands would be better elucidated than at the broad scale. Using repeated sampling of a set of 12 wetlands within a single landscape (~1km2), the hypothesis was tested that the loss of indigenous vegetation around wetlands (in this case due to alien vegetation invasion) is associated with changes in wetland physico-chemistry and, given the strong influence of physico-chemistry observed at the broad scale, this
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