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The Invertebrates of Indigenous Forests in Limpopo Province, South Africa: Diversity, Biogeography and Conservation

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The Invertebrates of Indigenous Forests in Limpopo Province, South Africa: Diversity, Biogeography and Conservation THE INVERTEBRATES OF INDIGENOUS FORESTS IN LIMPOPO PROVINCE, SOUTH AFRICA: DIVERSITY, BIOGEOGRAPHY AND CONSERVATION by JOHANNA LYNN HORN Submitted in fulfilment of the academic requirements for the degree of Master of Science in the Discipline Zoology, School of Botany and Zoology Faculty of Science and Agriculture University of KwaZulu-Natal Pietermaritzburg 2004 Horn – Forest Invertebrates ii ABSTRACT In this study I investigated patterns of invertebrate diversity in Limpopo Province indigenous forests, in order to highlight forests and taxa of special conservation significance. Invertebrates from seven target taxa were sampled in 11 patches of indigenous forest in Limpopo Province from February 2001 to January 2002, including six forests in the Soutpansberg and five forests in the northern Drakensberg. Selected forests comprise three distinct vegetation subtypes and the target taxa selected were millipedes, centipedes, earthworms, terrestrial molluscs, spiders, scorpions and amphipods. Invertebrates were sampled by active searching of quadrats and line transects and pitfall traps. A total of 11 969 indigenous target group individuals were sampled, comprising 14 orders, 50 families, 86 genera and 142 species (including at least nine new). There was a significant difference in the total invertebrate species richness and diversity of forest patches but results varied considerably when different target group figures were analyzed. With the exception of spiders, the factors influencing total and individual target group richness in forests could not be determined. Introduced invertebrates comprised a large proportion of the species and individuals sampled, but were not shown to affect indigenous fauna. Invertebrate species assemblages were most similar between forests sharing the same vegetation subtype and between forests in the same mountain region. However, each forest patch had unique species and some even had unique families. Limpopo Province forests support high numbers of endemic invertebrates. A total of 47 endemic invertebrate species were sampled, including six site endemics, eight local endemics, nine regional endemics and 24 national endemics. The numbers and scales of endemism varied by target group. Invertebrate species’ distributions in Limpopo Province forests generally support the biogeographic theories of Pleistocene forest refugia and the Limpopo River valley as a radiation barrier, although some important contradictions were found. Local endemism in Limpopo Province forests is likely the product of historical processes. Although some significant relationships were found between surrogate and true measures, single taxon biodiversity indicators, the higher taxon method, morphospecies and land classes could not accurately predict patterns of target invertebrate species richness in Horn – Forest Invertebrates iii Limpopo Province forests. Results show that formal species identification should be used if accurate richness estimates are desired; the use of surrogates is not supported by this study. Conservation of Limpopo Province forests is vital for the preservation of valuable invertebrate communities. No forest sampled in this study can be considered unimportant. Effective forest conservation and management is dependent upon the protection of forests of varying patch size, careful evaluation and control of utilization and the establishment and maintenance of corridors linking isolated forest patches. Horn – Forest Invertebrates iv PREFACE The experimental work described in this dissertation was carried out in the School of Botany and Zoology, University of KwaZulu-Natal, Pietermaritzburg, from January 2000 to December 2003, under the supervision of Dr Michelle Hamer and co-supervision of Professor Michael Lawes. 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: J. L. Horn (candidate) Signed: Dr Michelle Hamer (supervisor) Signed: Professor Michael Lawes (co-supervisor) Horn – Forest Invertebrates v TABLE OF CONTENTS Page ABSTRACT ii PREFACE iv TABLE OF CONTENTS v LIST OF TABLES ix LIST OF FIGURES xiii ACKNOWLEDGEMENTS xviii CHAPTER 1: GENERAL INTRODUCTION 1 1.1 BIODIVERSITY 1 1.1.1 What is biodiversity? 1 1.1.2 The three levels of biodiversity 3 1.1.3 Estimating global species diversity 4 1.1.4 Invertebrate abundance and diversity 5 1.1.5 Benefits of biodiversity to humans 7 1.1.6 Global species extinction 9 1.1.7 Threats to global biodiversity 10 1.2 FOREST BIODIVERSITY 13 1.2.1 Global forests and biodiversity 13 1.2.2 Forest invertebrate abundance and diversity 14 1.2.3 Why study forest invertebrates? 16 1.3 SOUTH AFRICAN FORESTS 18 1.3.1 Description of South African forests 18 1.3.2 South African forest classification 21 1.3.3 South African Afromontane forests 22 1.4 PURPOSE AND OBJECTIVES 24 CHAPTER 2: STUDY SITES, TAXA AND METHODS 27 2.1 INTRODUCTION 27 2.1.1 Northern Mistbelt Forests of Limpopo Province 27 2.1.2 Species vs. morphospecies in biodiversity studies 29 2.1.3 Focal groups vs. target groups 31 2.1.4 Invertebrate collection techniques 32 2.2 STUDY SITES 34 2.3 STUDY TAXA 38 2.3.1 Millipedes (Class: Diplopoda) 38 2.3.2 Centipedes (Class: Chilopoda) 40 2.3.3 Scorpions (Order: Scorpionida) 41 2.3.4 Web building and ground wandering spiders (Order: Araneae) 42 2.3.5 Terrestrial molluscs (Class: Gastropoda) 44 2.3.6 Earthworms (Class: Oligochaeta) 46 2.3.7 Terrestrial amphipods (Family: Talitridae) 47 2.3.8 Additional taxa 48 2.4 METHODS 48 Horn – Forest Invertebrates vi 2.4.1 Quadrat sampling 48 2.4.2 Line transect sampling 50 2.4.3 Pitfall trap sampling 51 2.4.4 Other sampling techniques 52 2.4.5 Specimen identification 53 CHAPTER 3: INVERTEBRATE SPECIES DIVERSITY OF LIMPOPO PROVINCE FORESTS 55 3.1 INTRODUCTION 55 3.1.1 Biodiversity surrogates 55 3.1.2 Measuring alpha diversity 56 3.1.3 Introduced species 59 3.2 METHODS 60 3.2.1 Calculations of richness, diversity and evenness 61 3.2.2 Comparisons of richness, diversity and evenness 61 3.2.3 Absolute species richness, diversity and evenness 61 3.2.4 Descriptive models - multiple regression analysis 62 3.2.5 Introduced species 63 3.3 RESULTS 63 3.3.1 Total numbers of species and individuals 63 3.3.2 Regional comparisons of richness, diversity and evenness 67 3.3.3 Comparisons of forest richness, diversity and evenness 67 3.3.4 Comparisons of target group richness, diversity and evenness 71 3.3.5 Absolute species richness, diversity and evenness 75 3.3.6 Descriptive models - multiple regression analysis 75 3.3.7 Introduced species 77 3.4 DISCUSSION 80 3.4.1 Species composition 80 3.4.2 Regional richness, diversity and evenness 81 3.4.3 Forest richness, diversity and evenness 82 3.4.4 Target group richness, diversity and evenness 83 3.4.5 Introduced species 89 3.5 CONCLUSION 91 CHAPTER 4: INVERTEBRATE COMMUNITIES, DISTRIBUTION AND BIOGEOGRAPHY 93 4.1 INTRODUCTION 93 4.1.1 Measuring beta diversity 94 4.1.2 Species endemism 95 4.1.3 Limpopo Province forest history in brief 96 4.2 METHODS 98 4.2.1 Site similarity 98 4.2.2 Cluster analysis 98 4.2.3 Species endemism analysis 99 Horn – Forest Invertebrates vii 4.2.4 Comparison with Zimbabwean and KwaZulu-Natal forests 101 4.3 RESULTS 103 4.3.1 Site similarity 103 4.3.2 Cluster analysis 109 4.3.3 Species endemism analysis 109 4.3.4 Comparison with Zimbabwean and KwaZulu-Natal forests 123 4.4 DISCUSSION 127 4.4.1 Site similarity 127 4.4.2 Endemism 128 4.4.3 Forest history and invertebrate distribution 133 4.5 CONCLUSION 135 CHAPTER 5: ASSESSMENT OF BIODIVERSITY SURROGATES 137 5.1 INTRODUCTION 137 5.1.1 Biodiversity indicators and invertebrate studies 139 5.1.2 Land classes as surrogates for species' distributions 143 5.2 METHODS 145 5.2.1 Individual taxa as indicators of species richness 146 5.2.2 Higher taxa as indicators of invertebrate species richness 146 5.2.3 Morphospecies as indicators of formal (true) invertebrate species richness 147 5.2.4 Vegetation classification as a surrogate for invertebrate species' distributions 148 5.3 RESULTS 148 5.3.1 Individual taxa as indicators of species richness 148 5.3.2 Higher taxa as indicators of invertebrate species richness 154 5.3.3 Morphospecies as indicators of formal (true) invertebrate species richness 159 5.3.4 Vegetation classification as a surrogate for invertebrate species' distributions 165 5.4 DISCUSSION 165 5.4.1 Individual taxa as indicators of species richness 165 5.4.2 Higher taxa as indicators of invertebrate species richness 169 5.4.3 Morphospecies as indicators of invertebrate species richness 171 5.4.4 Vegetation classification as a surrogate for invertebrate species' distributions 173 5.5 CONCLUSION 175 CHAPTER 6: SUMMARY AND CONSERVATION RECOMMENDATIONS 176 6.1 INTRODUCTION 176 6.1.1 Uses of and threats to South African forests 176 6.1.2 South African forest conservation 178 6.1.3 Limpopo Province forest conservation at present 180 Horn – Forest Invertebrates viii 6.2 LIMPOPO PROVINCE FOREST INVERTEBRATES 181 6.2.1 Invertebrate species diversity - major findings 181 6.2.2 Invertebrate communities, distribution and biogeography- major findings 182 6.2.3 Assessment of biodiversity surrogates - major findings 183 6.2.4 Additional information 184 6.3 CONSERVATION OF LIMPOPO PROVINCE FORESTS 188 6.3.1 Why conserve Limpopo Province forests? 188 6.3.2 Which forests should be conserved in Limpopo Province? 188 6.3.3 How should we conserve forests in Limpopo Province? 191 6.4 CONCLUSION 192 REFERENCES 194 APPENDICES 222 Horn – Forest Invertebrates ix LIST OF TABLES Page Table 1.1: Afromontane forest by province in South Africa. The total area of the Afromontane forest in each province, the proportion of the province area that Afromontane vegetation occupies and the contribution to the total area of Afromontane forest in South Africa are given (Low & Rebelo 1996).
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