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Cape Elements on High-Altitude Corridors and Edaphic Islands

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Cape Elements on High-Altitude Corridors and Edaphic Islands CAPE ELEMENTS ON HIGH-ALTITUDE CORRIDORS AND EDAPHIC ISLANDS by CLlNTON CARBUTT BACHELOR OF SCIENCE (HONOURS) (NATAL) Submitted in fulfilment of the academic requirements for the degree of DOCTOR OF PHILOSOPHY (in the Discipline of Botany) School of Botany and Zoology Faculty of Science and Agriculture University of KwaZulu-Natal Pietermaritzburg SEPTEMBER 2004 11 PREFACE The experimental work described in this thesis was carried out in the School of Botany and Zoology, University of Natal, Pietermaritzburg, from January 1998 to December 2003, under the supervision of Professor Trevor Edwards and co- supervision of Professor Richard Beckett. Fieldwork was conducted in numerous Ezemvelo KZN Wildlife reserves, particularly the uKhahlamba- Drakensberg Park, and was coordinated by Mr. C. R. Scott-Shaw. These studies represent original work by the author and have not otherwise been submitted in any form for any degree or diploma to any University. Where use has been made of the work of others it is duly acknowledged in the text. Signed: ....~...... (Clinton Carbutt) . Pietermaritz~?cem ber 2003) Supervisor· (Prof. T. J. Edwards) .. .U!SJJ;~ Co-supervls r f (Prof. R. P. Beckett) III FRONTISPIECE "As for man, his days are like grass, he flourishes like a flower in the field; the wind blows over it and it is gone, and its place remembers it no more. But from everlasting to everlasting the Lord's love is with those who fear Him, and His righteousness with their children's children -with those who keep His covenant and remember to obey His precepts. The lord has established His throne forever, and His kingdom rules over all" (Psalms 103: 15 -19). In loving memory of Leonora (Leonie) Mary Prozesky, who dearly loved the mountains MLAMBONJA WILDERNESS AREA, NORTHERN KWAZULU-NATAL DRAKENSBERG IV PUBLICATIONS The following publications have arisen from this course of study: CARBUTT, C. & EOWAROS, T. J. 2001. Cape elements on high-altitude corridors and edaphic islands: historical aspects and preliminary phytogeography. Systematics and Geography ofPlants 71: 1033-1061. CARBUTT, C. & EOWAROS, T. J. 2004. The flora of the Drakensberg Alpine Centre. Edinburgh Journal ofBotany 60(3): 581-607. FORMAT All chapters with the exception of Chapters 1 and 5 ('Introduction' and 'Conclusion') have or are being prepared for journal submission. The format of this thesis therefore is that of a number of free-standing chapters. Slight repetition in the introductory sections of each chapter is therefore inevitable. References follow after each chapter. v ACKNOWLEDGEMENTS I would firstly like to acknowledge the help of the Lord Jesus, personal friend and Saviour, to whom this work is credited. In addition, this work would not have been possible without the help of the following individuals and organizations: The National Research Foundation (NRF), the Ernest Oppenheimer and Ernst and Ethel Eriksen Trust Funds, the Wildlife and Environment Society of South Africa (KwaZulu-Natal and Sani Branches) and the University of Natal (Gay Langmuir Bursary Fund), for their generous provision of funding. Prof. Trevor Edwards, my supervisor, for his time, generous advice, encouragement and knowledgeable input. Prof. Richard Beckett, my co-supervisor, for his help and friendly advice. Dr. Steve Johnson for input, advice and memorable field trips (long live the mountains). Prof. Peter Under (Institute for Systematic Botany, University of Zurich) and Prof. Braam Van Wyk (Schweickerdt Herbarium, University of Pretoria) for their VI generous provision of literature, expertise and advice. Members of staff of the University of Natal, in particular: Prof. Owen McGee (retired), Prof. Hans Beckedahl, Prof. Michael Savage, Dr. Ed Granger, Richard Fynn, Craig Morris, Essack Abib and Chris Bester (all of the School of Applied Environmental Sciences) and Prof. Roland Schulze and Mark Horan (both ofthe School of Bioresources, Engineering and Environmental Hydrology), for a variety of input. Ezemvelo KZN Wildlife, in particular Rob Scott-Shaw and those members of staff stationed in the uKhahlamba-Drakensberg Park, Oribi Gorge and Umtamvuna Nature Reserves. KwaZulu-Natal Department of Agriculture, particularly Douglas Chapman (Agrometeorology), and Steve Bainbridge and Alan Manson (Soil Fertility and Analytical Services). The National Botanical Instituteforthe use ofdatafrom the National Herbarium, Pretoria (PRE) Computerized Information System (PRECIS). Gary and Carmen Neethling ('Summit Farm', Oribi Flats) for access to theirfarm to collect soils and to Tony Abbott ('Clearwater Farm', Umtamvuna) and Nick and Mariette de Goede (ex-Garden Castle) for monitoring the pot experiments. VII Dr. Robin Bennett (Agricultural Research Council), Dr. Hugo Van Niekerk and Tappie Bredenkamp (Small Grains Institute, Bethlehem), Dr. Elisa Raulings (Australia), and Mrs Anne Rennie (Underberg) for help with various aspects of this project. Stephne Stuartand Christina Potgieter, for laboratory and herbarium assistance respectively. Renny Noble, ourtrustworthy and hard-working storeman, and Doug Boddy, for all his creative handiwork. Boyd Escott, Leeann Hoatson, Mike Grundy and Shane Sparg for their company and digging skills on soil collecting trips. Nicky Grieshaber for proofreading a portion of this thesis. My Dad and Mom, Keith and Jenny, and my sister Lara, thank you for everything, especially you mom for your good example in the garden. Finally, to Hilary and Nathi, my beloved family, I love you both very much! VIII ABSTRACT Common to the temperate floras throughout sub-Saharan Africa is a group of taxa with strong ties to the Cape Floristic Region (CFR) (;::: Cape elements). Their distribution is limited to the eastern escarpment of Africa (e.g. the Drakensberg Alpine Centre - DAC), on nutrient-rich humic soils, as well as on isolated sandstone outcrops of low elevation, on nutrient-poor soils (e.g. the Pondoland Centre - PC), suggesting that intrinsic soil fertility is not the primary determinant of their distribution. The principal aim of this study was to determine which aspect of the edaphic environment of the DAC is most influenced by temperature, that may indirectly render it nutrient-poor and therefore provide suitable niches for Cape elements, as in the PC. A multidisciplinary approach involving aspects of plant biogeography, plant ecology, plant ecophysiology and soil chemistry was therefore adopted. The study regions were the DAC, PC and the KwaZulu-Natal Midlands. The flora of the DAC was resurveyed for this study, and is richer than previously thought: 2818 native taxa, most of which (2520) are angiosperms. The phytogeography of the DAC and PC is discussed, and comparisons are made with the floras of Kwazulu-Natal and the CFR. Their climatic environments, as well as those for the CFR and Sneeuberge, were compared using rainfall and temperature data from a range of sources. These climatic regimes were correlated with the floristic patterns of Cape elements for the high-altitude regions of South Africa and Lesotho. Altitude and rainfall increased, and temperature decreased, as the number of Cape elements increased towards the DAC. This study provided a IX contemporary inventory of the Cape elements of the DAC and PC. A total of 89 genera are recognised as Cape elements, of which 60 (c. 67%) are shared between the two regions. The highest number of Cape elements recorded for the eastern escarpment was the DAC (72 genera), with the highest numberfrom all sites analysed being the PC (77 genera). The most Cape elements are contributed by the Asteraceae, Scrophulariaceae, Iridaceae, Fabaceae, Orchidaceae and Restionaceae, partly due to the success ofannual aerial parts and their geophytic growth forms, which are convergent in these families. Furthercompartmentalisation into life and growth forms shows that most Cape elements of the DAC and PC are either ericoid (and sclerophyllous) or mesic herbs and shrubs. The ecological and ecophysiological aspects of this study involved the use of reciprocal pot experiments established along a gradient of altitude from coastal hinterland to mountain, that investigated the interactions between altitude, temperature and substrate on plant productivity in sites known either to support or to exclude Cape elements. Three soils were used at each site, representative of the DAC, PC and KwaZulu-Natal Midlands. The interactions between 'soil' and 'site' (~ the climatic environment) were quantified using a temperate testtaxon (Diascia) that has a strong Cape-centred distribution. Plant characters relating to morphology and nutrient content, and soil characters relating to fertility, were used as the basis for comparing treatment effects (soil-site interactions). Soil nitrogen availability was assayed using pot experiments with Eragrostis curvula (Schrad.) Nees. Wheat pot 3 experiments revealed no A1 + toxicity in 'Drakensberg' soil. Non-metric x multidimensional scaling (NMDS) and redundancy analysis (RDA) indicated that all soil-site interactions were significant contributors to biomass differences, and that the Cape taxon performed poorly in the nutrient-rich Drakensberg soil at low altitude. Soil samples indicated that Drakensberg soil was the most nutrient-rich, and Pondoland soil the mostnutrient-poor. Although total nitrogen in Drakensberg soil was six times higher than Pondoland soil, both soils mineralised similar low levels of nitrogen at their respective spring temperatures. The result for Drakensberg soil (simulated so as to include the effect of altitude)
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