CONTENTS IN FULL

Declaration i

Acknowledgements ii

Abstract v

Contents in brief vi

Contents in full vii

List of figures xi

List of tables xiii

Scientific names xiv

Acronyms and abbreviations xv

Chapter 1 Introduction 1
1.1 Introduction 1
1.1.1 Spatial resource heterogeneity 2
1.1.2 Optimal spatial foraging 2
1.1.3 Animal water dependency 4
1.1.4 Foraging range 8
1.1.5 Spatial impacts 9
1.1.6 Plant response 12
1.1.7 Management 16
1.1.8 Permanency of effect 18
1.2 Modelling 20
1.2.1 Simplification of a complex system 20
1.2.2 Conceptual models 20
1.2.3 The use of conceptual models in rangeland management 21
1.2.4 The general logistic model 21
1.2.5 The use of the logistic model in rangeland management 24
1.2.6 Regression models 24
1.2.7 The prediction of animal densities 25
1.2.8 Looking for gradients from outer space 25
1.2.9 Productivity gradients 26
1.2.10 System models 27
1.2.11 Patterns in forage production and utilisation 27
1.2.12 A grid-based model of a single piosphere 32
1.2.13 Grid-based models of animal densities 33
1.2.14 Modelling the physiology of foraging behaviour 33
1.2.15 Distance to water versus animal energy expenditure 34
1.2.16 Piospheres and the socio-economics of livestock production 35
1.2.17 Miscellaneous models 36
1.2.18 Conclusions and gaps in knowledge 36
1.3 This thesis 38
1.3.1 Problem statement 38

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 1.3.2 Research questions and objectives 40
1.3.3 Thesis structure 42
1.3.4 Experiment locations 43
1.3.5 The model animal - the domestic goat (Capra hircus) 44
Chapter 2 Utilisation of resources in a spatially heterogeneous landscape 45
Abstract 45
2.1 Objectives 45
2.2 Introduction 46
2.2.1 Specific research questions 49
2.3 Methods 50
2.3.1 Experimental paddock 50
2.3.2 Vegetation 50
2.3.3 Plant communities 52
2.3.4 Animals 54
2.3.5 Animal activity 55
2.3.6 Spatial analysis 57
2.3.7 Animal impacts 58
2.4 Results 60
2.4.1 Animal activity 60
2.4.2 Animal locations 62
2.4.3 Spatial impacts 62
2.4.4 Community selection 65
2.4.5 Landscape utilisation 68
2.5 Discussion 73
2.6 Conclusions 78
The roles of animal watering behaviour and travel costs in determining the
Chapter 3 79
distribution of spatial impacts
Abstract 79
3.1 Objectives 79
3.2 Introduction 80
3.3 Specific research questions 81
3.4 Goat water requirements 83
3.5 Goat water intake 85
3.6 Goat water utilisation 87
3.7 Water deficiency in goats 88
3.8 Goat watering frequency 90
3.9 Literature estimates of goat foraging range 91
3.10 Model estimates of goat foraging range 95
3.11 Model 1 - Foraging range from body size relations 95
3.12 Model 2 - Foraging range from predicted energy balance 97
3.13 Model 3 - Foraging range from predictions of spatial foraging behaviour 100
3.13.1 Model description 101
3.13.2 Spatial extensions 104
3.13.3 Scale issues 104
3.13.4 Energy intake 106
3.13.5 Spatial foraging 107

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 3.13.6 Vision 110
3.13.7 Assessment 112
3.13.8 Global and local G* 114
3.13.9 Memory 116
3.13.10 Site selection 119
3.13.11 Constraints on intake 120
3.13.12 Model 3 parameters and output 121
3.14 Model 3 results and discussion 125
3.14.1 Test of Hypothesis 1: Geometry 130
3.14.2 Test of Hypothesis 2: Foraging activity 135
3.14.3 Test of Hypothesis 3: Travel costs 136
3.15 Conclusions 140
Chapter 4 Consequences of a utilisation gradient on individual intake 143
Abstract 143
4.1 Objectives 143
4.2 Introduction 144
4.2.1 The herbivore functional response 145
4.2.2 The browser functional response 147
4.2.3 Specific research questions 150
4.3 Methods 150
4.3.1 Paddock structure 150
4.3.2 Paddock treatment 151
4.3.3 Vegetation survey 151
4.3.4 Bite rate 152
4.3.5 Foraging velocity 153
4.3.6 Analysis 154
4.4 Results 156
4.4.1 Paddock character 156
4.4.2 Foraging velocity 159
4.4.3 Animal feeding response to the utilisation gradient 162
4.4.4 Intake rate 162
4.4.5 Functional response 165
4.5 Discussion 167
4.5.1 A note on spinescence 169
4.5.2 Thresholds in patch browsing 170
4.6 Conclusions 173
Chapter 5 Emergence of spatial pattern and its consequences for dry season survival 174
Abstract 174
5.1 Objectives 174
5.2 Introduction 175
5.2.1 2001: A spatial odyssey 175
5.2.2 Emergent properties 175
5.2.3 The nature of emergence 176
5.2.4 Defining emergence 179
5.2.5 Tests of Emergence 181
5.2.6 Specific research questions 184

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 5.3 Methods 187
5.3.1 Model parameters 187
5.3.2 Model output and analysis 188
5.3.3 Emergence test 1 188
5.3.4 Emergence test 2 189
5.4 Results and discussion 190
5.4.1 Example 1 - Syntactic emergence 192
5.4.2 Example 2 - Semantic emergence 195
5.4.3 Search for mechanisms 200
5.4.4 Summary of results 205
5.5 Conclusions 206
5.5.1 Consequences of the emergence of spatial pattern 206
5.5.2 Resource matching and ecosystem dynamics 207
5.5.3 Consequences for management 209
Chapter 6 Discussion 213
6.1 Synopsis 213
6.2 Synthesis of experimental findings 214
6.2.1 Resource heterogeneity 214
6.2.2 Animal assessment of resource heterogeneity 214
6.2.3 Animal response to resource heterogeneity 215
6.2.4 The organisation of resource heterogeneity into utilisation gradients 215
6.2.5 Animal response to utilisation gradients 216
6.2.6 Population response to utilisation gradients 217
6.2.7 Species response to utilisation gradients 218
6.3 Gaps in knowledge and future research 221
6.3.1 Fieldwork 222
6.3.2 Modelling 223
6.4 Summary of contributions and conclusion 224
Appendices 225
A1 Appendix 1: Species composition of plant communities 225
A2 Appendix 2: Sources of error in animal location measurements 230
A2.1 Accuracy of animal locations 230
A2.2 Independence of animal locations 231
A3 Appendix 3: Goat diet selection from impact scores 232
A4 Appendix 4: Goat diet selection from feeding duration 234
A5 Appendix 5 Poster presented to irc 2001 pretoria, sa. 237
A6 Appendix 6: Semi-arid grazing system model software 240
A6.1 Program structure 240
A6.2 Climate and Soil modules 243
A6.3 Plant module 243
A6.4 Animal module 243
A6.5 Module structure 244
A7 Appendix 7: Use Biomass Rule 257
A8 Appendix 8: Biterate software 258
References 272