Spatial Modelling, Phytogeography and Conservation in the Eastern Arc Mountains of Tanzania and Kenya Philip John Platts PhD THE UNIVERSITY OF YORK York Institute for Tropical Ecosystem Dynamics Environment Department March 2012 This thesis is dedicated to my father Philip Graham Platts 27th November 1937 – 11th February 2012 v Thesis Abstract Forests in the Eastern Arc Mountains are amongst the oldest and most biodiverse on Earth. They are a global priority for conservation and provide ecosystem services to millions of people. This thesis explores how spatial modelling can provide direction for conservation and botanical survey, and contribute to understanding of phytogeographical relationships. The ecoregion is rigorously defined by terrain complexity, vegetation distribution and established geoclimatic divisions, providing a coherent platform upon which to collate and monitor biological and socioeconomic information. Accordingly, 570 vascular plant taxa (species, subspecies and varieties) are found to be strictly endemic. The human population exceeds two million, with median density more than double the Tanzania average. Population pressure (accrued across the landscape) is shown to be greatest adjacent to the most floristically unique forests. Current knowledge on species distributions is subject to sampling bias, but could be systematically improved by iterative application of the bioclimatic models presented here, combined with targeted fieldwork. Tree data account for 80% of botanical records, but only 18% of endemic plant species; since conservation priorities differ by plant growth form, future fieldwork should aim to redress the balance. Concentrations of rare species correlate most strongly with moisture availability, whilst overall richness is better predicted by temperature gradients. Climate change impacts are projected to be highly variable, both across space and between species. Concordant with the theory that past climatic stability facilitated the accumulation of rare species, contemporary climates at sites of known endemic richness are least likely to be lost from dispersal-limiting mountain blocs during the 21st century. Faced with rapid population growth and the uncertainty of climate change, priorities for governance are to facilitate sustainable forest use and to maintain/restore habitat connectivity wherever possible. Overall, the thesis demonstrates that decision makers concerned with biodiversity conservation, particularly in mountain and coastal regions, should be wary of inferring local patterns of change from broad-scale models. The current study is a step toward spatially refined understanding of conservation priorities in the Eastern Arc Mountains, whilst novel methodologies have wider application in the fields of species distribution modelling and mountain analysis. vi Contents Thesis Abstract Contents List of Tables List of Figures Acknowledgements Author’s Declaration 1. General Introduction 1 Overview 3 Forest degradation and loss 4 Conservation prioritisation 5 Species distribution modelling 6 Describing the response 8 Prediction 10 Thesis aims and objectives 11 Study region 13 Geology and climate 13 Biological importance 14 Delineation and classification 15 Forest use and protection 16 Data collection 18 Outline of analytical chapters 19 Chapter 2 – Mountain limits 19 Chapter 3 – Predicting tree distributions 20 Chapter 4 – Distribution models and conservation priority 20 Chapter 5 – Climate change 20 References 22 2. Mountains Limits 27 Article title, authors and abstract 29 Introduction 20 A qualitative definition for the Eastern Arc 31 Mountain limits 32 A global mountain typology 32 vii Methods 33 Regional refinements to the global typology 33 Elevational zonation 33 Terrain parameters 34 Matching to mountain vegetation 34 Bounding the chosen regional typology 35 Amalgamating features 35 Mountain selection based on relative relief 36 Results 36 Sensitivity to local elevation range 36 Boundary placement 39 Protected areas 42 Human populations 44 Discussion 45 Acknowledgements 49 Author contributions 49 Appendix 2A: Description of spatial data 50 Appendix 2B: Mountain classes in the regional typology 51 Appendix 2C: Derivation of the population surface 52 References 54 3. Predicting Tree Distributions 57 Article title, authors and abstract 59 Introduction 60 Model selection 61 Data bias 62 Envelope uncertainty 63 Methods 63 Study region 63 Tree data 65 Environmental predictor variables 65 Statistics for evaluation and calibration 68 Modelling experiments 69 Selecting predictors 69 Weighting absences 70 Spatial autocovariates 73 viii Results 73 Baseline models 73 Weighted models 74 Spatial models 76 Predictors and envelope uncertainty 78 Discussion 79 Conclusions 83 Acknowledgements 84 Author contributions 84 Appendix 3A: Tree species modelled 85 Appendix 3B: Stratified cross-validation 86 Appendix 3C: Neighbourhood size for spatial autocovariates 87 Appendix 3D: Species-specific results 88 References 97 4. Distribution Models and Conservation Priority 101 Article title, authors and abstract 103 Introduction 104 Methods 106 Study region 106 Plant inventory data 107 Environmental data 107 Model calibration 110 Background data 110 Predictor selection 110 Spatial autocorrelation 111 Testing and validation 111 Richness estimates 112 Results 112 Model performance 112 Sampling bias 114 Richness 115 Confirmed at bloc level 115 Predictive estimates 116 Growth form 116 Discussion 121 ix Conclusions 125 Acknowledgements 125 Author contributions 126 Appendix 4A: Details of plant data 127 Appendix 4B: Occurrence thresholds and sensitivity to prevalence 128 Appendix 4C: Analysis of model performance 129 Appendix 4D: Patch occupancy for endemic/threatened taxa 132 Appendix 4E: Drivers of richness and endemism 133 References 135 5. Climate Change 139 Article title, authors and abstract 141 Introduction 142 Methods 145 Study region 145 Plant data 145 Climate data 146 Climates lost and gained 147 Endemism vs. altitude 148 Univariate response 148 Multivariate response 149 Results 149 Spatial variation in change 149 Climates lost and gained 151 Endemism vs. altitude 153 Univariate response 153 Multivariate response 154 Discussion 155 Acknowledgements 162 Author contributions 163 Appendix 5A: Improving normality for statistical regression 164 Appendix 5B: 21st century climate anomalies 167 Appendix 5C: Climates lost and gained 171 Appendix 5D: Multivariate species models 174 References 182 x 6. Summary Discussion 185 Mountain limits 187 Endemism criteria 187 Human populations 189 Methods for species distribution modelling 191 Sample selection 191 Data weights 193 Variable selection 193 Environmental relationships 194 Socioeconomic relationships 197 Directions for fieldwork 199 Plants in context 201 References 204 Thesis Appendix 209 Appendix I: Co-authored paper abstracts 209 Appendix II: Population projections for Tanzania 217 Appendix III: Hydrological modelling 220 References 224 Abbreviations 225
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