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Botswana), a Subtropical Flood-Pulsed Wetland

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Botswana), a Subtropical Flood-Pulsed Wetland Biodiversity and Biomass of Algae in the Okavango Delta (Botswana), a Subtropical Flood-Pulsed Wetland Thesis submitted for the degree of Doctor of Philosophy by LUCA MARAZZI University College London Department of Geography University College London December 2014 I, LUCA MARAZZI, confirm that the work presented in this thesis is my own. Where information has been derived from other sources, I confirm that this has been indicated in the thesis. LUCA MARAZZI 2 ABSTRACT In freshwater bodies algae provide key ecosystem services such as food and water purification. This is the first systematic assessment of biodiversity, biomass and distribution patterns of these aquatic primary producers in the Okavango Delta (Botswana), a subtropical flood-pulsed wetland in semiarid Southern Africa. This study delivers the first estimate of algal species and genera richness at the Delta scale; 496 species and 173 genera were observed in 132 samples. A new variety of desmid (Chlorophyta) was discovered, Cosmarium pseudosulcatum var. okavangicum, and species richness estimators suggest that a further few hundred unidentified species likely live in this wetland. Rare species represent 81% of species richness and 30% of total algal biovolume. Species composition is most similar within habitat types, thus varying more significantly at the Delta scale. In seasonally inundated floodplains, algal species / genera richness and diversity are significantly higher than in permanently flooded open water habitats. The annual flood pulse has historically allowed more diverse algal communities to develop and persist in these shallower and warmer environments with higher mean nutrient levels and more substrata and more heterogenous habitats for benthic taxa. These results support the Intermediate Disturbance Hypothesis, Species-Energy Theory and Habitat Heterogeneity Diversity hypotheses. Higher algal biodiversity supports higher algal biomass in the floodplains, where species form three-dimensional communities of attached and periphytic algae requiring more nutrients than phytoplankton assemblages. Multivariate analyses demonstrate that habitat type, flooding frequency and conductivity most importantly influence the relative abundance of algal species, genera and phyla in the Okavango Delta. This study’s findings highlight how the preservation of water level fluctuations and habitat heterogeneity is crucial to maintaining biodiverse and thus resilient food webs in this unique ecosystem which faces increasing anthropogenic threats, such as global warming and upstream water abstraction plans. 3 Acknowledgments I wish to thank Anson W. Mackay, my Principal Supervisor, for giving me the great opportunity of undertaking this Ph.D. and for constantly advising me throughout my research; Vivienne Jones, my Subsidiary Supervisor for her constructive criticism and suggestions and Sophie des Clers and Lars Ramberg who co-supervised me in the early-mid phases of the project for their advice and encouragements. I gratefully acknoweldge my colleague Nqobizitha Siziba who collected numerous samples of algae during the periods when I could not be in Botswana (July and October 2009); Ponde Kauheva, Richard Mazebedi, Thomas Davidson, Ewan Shilland, Ben Goldsmith and the Okavango Research Institute staff for their work during the sampling campaigns and UCL laboratory staff for support on microscope work. My thanks also go to: the Royal Geographical Society, the UCL Geography Department and Graduate School and the UK DEFRA Darwin Initiative for funding fieldtrips to the Delta in April/May 2009 and February/March 2010; Barbara Leoni and Letizia Garibaldi (University of Milano Bicocca) for organizing the introductory course on phytoplankton identification in February 2009; Brian Whitton and David John for organizing the advanced course on phytoplankton identification in July 2011 and for their interest in my research; David Williamson and Chris Carter for providing me with very useful drawings and pictures of desmids from our samples. I also want to thank: Responding To Climate Change, Mouseion Limited, Campbell Harris Tutors, Greater London Tutors and my A-level and GCSE students and their parents as well as UCL staff members for providing me with work experiences to self-fund my Ph.D.; Jan Axmacher and Gina Clarke for advice on diversity data analyses and phytoplankton microscopy. I thank my colleagues Victoria Shepherd and Matt Owen for very useful discussions on my work and other colleague-friends for having shared tips, joys and frustrations during our respective Ph.D. adventures, especially in the good old days in the microscope-room. Last, but definitely not least, I wish to thank all my family - Carlo, Angela, Sara, Alba, Valter, Lisa, Giorgio, Anna - and my fiancée Efe for the great support they have been giving me throughout these years. This Thesis is dedicated to my grandparents Luciano, Giorgio and Olga and my great-aunt Paola who are always in my heart. 4 Table of contents List of figures..................................................................................................................8 List of tables..................................................................................................................12 GLOSSARY OF KEY TERMS AND ABBREVIATIONS.......................................14 CHAPTER 1 – WETLANDS AND ALGAE 1.1. OVERVIEW OF THE THESIS...........................................................................17 1.2. WETLANDS..........................................................................................................18 1.2.1. Wetland environment............................................................................................20 1.2.2. Ecosystem services and economic value................................................................20 1.2.3. Threats and conservation........................................................................................21 1.2.4. Flood-pulsed wetlands.............................................................................................23 1.2.5. Subtropical wetlands...............................................................................................25 1.3. MAIN FEATURES OF ALGAE..........................................................................29 1.3.1. Morphology and definitions................................................................................ 29 1.3.2. Algal cell structure, life and reproduction strategies .........................................31 1.3.3. The role of algae in aquatic food webs...................................................................33 1.3.4. Algae as providers of ecosystem services...............................................................35 1.4. BIOGEOCHEMICAL CYCLES: THE ROLE OF ALGAE............................41 1.5. ENVIRONMENTAL DRIVERS OF ALGAL DISTRIBUTION.....................43 1.5.1. Geography and space...............................................................................................43 1.5.2. Limnology ............................................................................................................44 1.5.3. Seasonality and flood-pulse ................................................................................. 46 1.5.4. Habitat structure .................................................................................................47 1.6. THEORETICAL FRAMEWORK FOR THE THESIS....................................49 1.6.1. Intermediate Disturbance Hypothesis (IDH).........................................................50 1.6.2. Resource Competition Theory (RCT)....................................................................53 1.6.3. Species-Energy Theory (SET).................................................................................55 1.6.4. Habitat Heterogeneity Hypothesis (HHH).............................................................56 CHAPTER 2 – STUDY REGION: THE OKAVANGO DELTA 2.1. THE OKAVANGO RIVER BASIN.....................................................................57 2.1.1. Geography and climate....................................................................................... 58 2.1.2. Socio-economic aspects ....................................................................................... 60 2.1.3. Threats ................................................................................................................ 61 2.1.4. Scenarios ............................................................................................................. 62 2.2. THE OKAVANGO DELTA.................................................................................64 2.2.1. Geology, topography and soil ............................................................................. 65 2.2.2. Climate and hydrology ....................................................................................... 66 2.2.3. Water chemistry and quality..................................................................................69 2.2.4. Hydroecology ......................................................................................................70 2.2.5. Biodiversity .........................................................................................................72 2.2.6. Ecosystem services and socio-economic aspects................................................. 75 2.3. AIM, RESEARCH QUESTIONS AND OBJECTIVES....................................78 CHAPTER 3– STUDY SITE, FIELD AND LABORATORY METHODS 3.1. STUDY SITES AND SAMPLING CAMPAIGNS.............................................84
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