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Great Escarpment): Assessing Migration Routes and Endemism

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Great Escarpment): Assessing Migration Routes and Endemism The Phytogeography of the Sneeuberg, Nuweveldberge and Roggeveldberge (Great Escarpment): Assessing Migration Routes and Endemism A thesis submitted in the fulfilment of the requirements for the degree of Doctor of Philosophy of Rhodes University by Vincent Ralph Clark May 2010 Supervisor: Prof. N.P. Barker (Department of Botany, Rhodes University) Co-supervisor: Prof. L. Mucina (Department of Environmental & Aquatic Sciences, Curtin University of Technology) Dedication “For by Him were all things created that are in heaven and that are on earth, visible and invisible…All things were created through Him and for Him. And He is before all things and in Him all thing consist” Colossians 1:16 17 This thesis is dedicated to the Lord Jesus Christ, Who knows unequivocally what actually has gone on on this planet in ages gone by (!) i Abstract The Great Escarpment forms a semi-continuous mountain system 5 000 km long, stretching from Angola in the north-west, south through Namibia, and into western, southern and eastern South Africa, including Lesotho and Swaziland. It is composed of a wide variety of geological suites but is unified in representing the edge of the African plateau and the passive Gondwanan continental margin. The Great Escarpment falls into all major climatic zones on the subcontinent, is a repository of palaeo- and neo-endemics, hosts more than half of southern Africa’s centres of plant endemism, and has a rich suite of endemic fauna. In addition, the Great Escarpment is believed to be both a refugium and corridor for biological diversity. Despite the biological richness of the Great Escarpment, research to date has been fragmented and many sections of the Great Escarpment have not been studied. The aim of this study is to contribute to research on the Great Escarpment by undertaking a detailed floristic study of the southern Great Escarpment (the Sneeuberg, Nuweveldberge and Roggeveldberge). Together these mountains comprise approximately 1 000 km (one fifth) of the Great Escarpment, and occupy a transition zone between the summer rainfall zone in the east and the winter rainfall zone in the west. They are also the sections of Great Escarpment most closely situated to the Cape Floristic Region (CFR) and would thus be involved in hypothesised migration routes for lineages that also occur further north through the Drakensberg Alpine Centre (DAC) to the East African mountain chain. Detailed fieldwork of the southern Great Escarpment was undertaken over a period of four years in all seasons. Approximately 8 000 specimens were collected. Particular emphasis was placed on areas that may represent refugia, i.e. the highest plateaux and peaks, mesic areas and cliff-lines. An overview of each mountain range, together with their endemic plant species and phytogeography, is provided. Approximately ten new species have been discovered during this study, two of which have been ii described to date. Numerous endemics only known from their types have also been rediscovered. The Sneeuberg is defined as a new centre of plant endemism on the Great Escarpment (endemism of 2.3%), and the role of the Boschberg and Groot-Bruintjieshoogde (part of the Sneeuberg) as a nexus for floristic migration routes is discussed. The Nuweveldberge is shown to have low endemism despite a floristic tally similar to the Sneeuberg, while the Roggeveldberge are confirmed to be the most endemic-rich section of the southern Great Escarpment. The field data collected was augmented by available data in taxonomic revisions, and floras for the Sneeuberg, Nuweveldberge and Roggeveldberge were compiled. In order to floristically compare the southern Great Escarpment with other sections of the Great Escarpment and the CFR, a database of some 12 000 taxa was created using available floristic data for the CFR, DAC and Great Winterberg Amatolas, together with the data collated for the Sneeuberg, Nuweveldberge and Roggeveldberge. These data were analysed using phenetic methods and Parsimony Analysis of Endemicity (PAE). The results indicate stronger linkages in the east, particularly between the Sneeuberg and Nuweveldberge, and between the Sneeuberg and the Great Winterberg Amatolas. The relationship of the Roggeveldberge with the rest of the southern Great Escarpment remains ambiguous. In order to refine notions of connectivity and migration routes, 19 well-sampled phylogenies were assessed for sister-taxon disjunctions to explore CFR Great Escarpment connections. Palaeo- connectivity between the CFR and southern Great Escarpment is most strongly supported for the south-eastern (SE) connection, and less so for the north-western (NW) and Matjiesfontein connections. There is support for the current (or recent) use of these three connections from numerous species that occur on both sides of the connections. Results of these analyses indicate that the southern Great Escarpment is a palaeo-corridor, the functioning of which has been broken by the aridification of the Nuweveldberge since the Last Glacial Maximum (LGM). Floristic connectivity is strongest in the east, from the Nuweveldberge to the DAC, and is less so in the west between the Nuweveldberge iii and the Roggeveldberge a finding attributed to the transition from a reliable winter rainfall regime on the Roggeveldberge to an unpredictable moisture regime on the Nuweveldberge. The mountains of the southern Great Escarpment are thus a series of refugia from a previous moister, cooler climate and are a corridor between the eastern and western components of the Great Escarpment. The SE connection is the primary link between the CFR and the eastern Great Escarpment Afromontane region in southern Africa. The implications of this research are that accurate conservation assessments and Red Data listings for many of the previously poorly-known endemics can now be made, and appropriate conservation measures implemented. Climate change remains the primary threat to these endemics and montane taxa in general, while degradation of wetlands is the primary threat to the water catchment service provided by the southern Great Escarpment. Future detailed research on the Great Winterberg Amatolas and Stormberg and a comprehensive flora of the Hantamm Roggeveldberge will further enhance our understanding of the floristics of the southern Great Escarpment, and provide the necessary data for comprehensive GIS-based models of proposed climate change scenarios for local, regional and national conservation planning. iv Table of Contents Dedication .......................................................................................................................................... i Abstract ............................................................................................................................................. ii Table of Contents .............................................................................................................................. v List of Figures .................................................................................................................................. ix List of Tables ................................................................................................................................. xiii List of Appendices........................................................................................................................... xv Acknowledgements ....................................................................................................................... xvii Declaration ..................................................................................................................................... xxi Preamble ....................................................................................................................................... xxii Chapter 1: The Great Escarpment of Southern Africa. ........................................................................ 1 1.1. Introduction ............................................................................................................................ 1 1.2. The geology, geomorphology, origin and physical characteristics of the Great Escarpment...... 6 1.2.1. Origins ............................................................................................................................. 6 1.2.2. Geology ........................................................................................................................... 7 1.2.3. Relief, Hydrology and Intervals ........................................................................................ 7 1.2.4. Climate .......................................................................................................................... 11 1.2.5. Comparisons with other mountain systems in the world .................................................. 12 1.3. The Biodiversity of the Great Escarpment ............................................................................. 16 1.3.1. Angola ........................................................................................................................... 18 1.3.2. Namibia ......................................................................................................................... 19 1.3.3. South Africa, Lesotho and Swaziland ............................................................................. 20 1.3.4. Zimbabwe–Mozambique ................................................................................................ 21 1.4. Research
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