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The Impact of Dehorning on the White Rhinoceros (Ceratotherium Simum) and the Evaluation of Novel Anti-Poaching Tactics

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The Impact of Dehorning on the White Rhinoceros (Ceratotherium Simum) and the Evaluation of Novel Anti-Poaching Tactics The impact of dehorning on the white rhinoceros (Ceratotherium simum) and the evaluation of novel anti-poaching tactics By Samuel G. Penny November 2019 University of Brighton A thesis submitted in partial fulfilment of the requirements of the University of Brighton for the degree of Doctor of Philosophy Abstract Abstract The white rhinoceros (Ceratotherium simum) is experiencing unsustainable poaching losses fuelled by an increased demand for horn. In an attempt to reduce poaching pressure and faced with rising security costs, increasingly, private and state reserves are dehorning their rhino populations as a management approach. Despite the procedure’s wide-scale practice, significant knowledge gaps exist on how dehorning may affect the behaviour or physiology of white rhinos. Given that rhinos require continued protection after dehorning, there is also a need to develop effective, low-cost techniques to aid on- the-ground conservation efforts. This research employed a combination of field studies and laboratory experiments to address this. To determine whether dehorning affected the social behaviour of white rhinos, behavioural observations were conducted on free-ranging horned and dehorned populations in South Africa. Dehorned individuals exhibited similar agonistic and cohesive social behaviours to horned individuals. However, a rise in the rate of agonistic interactions was detected after a repeat dehorning procedure. The results also provide the first evidence of a non-territorial dominance hierarchy among free-ranging white rhinos, with the lowest ranked individuals shifting in social position after dehorning. To investigate whether white rhinos exhibited a long-term physiological response to horn removal, changes in adrenal and gonadal steroid levels were monitored in faecal samples. Long-term hormone profiles were not influenced by the number of times a rhino had been dehorned. Furthermore, there was no detectable difference in corticoid concentrations between a dehorned and horned population. Dehorning did not appear to act as a chronic stressor and thus compares favourably to other conservation techniques, such as translocation. Horn use in a non-social context was examined by camera trapping and behavioural observations. Dehorning did not have a detectable impact on resource access, with no change in digging behaviours observed during geophagy and wallowing, nor on the frequency of horn rubbing behaviours, suggesting limited impact of the procedure on this aspect of the species ecology. i Abstract The movement of rhinos from at-risk areas could provide an effective anti-poaching tactic. Evaluation of the efficacy of novel deterrent-based techniques to move rhinos from poaching hotspots into safer areas was undertaken. White rhinos were exposed to acoustic, olfactory and drone-based stimuli and repeatedly fled from low-altitude drone flights and the siren noises. However, olfactory stimuli were ineffective at inciting avoidance behaviour. The drone’s long transmission range and capability of pursuit make it most suited to manipulating rhino movement as a means to reduce poaching risk. ii Contents Contents Abstract i Contents iii List of Tables vii List of Figures ix List of Acronyms xii Acknowledgments xiii Author’s declaration xiv Chapter 1 – General introduction 1 1.1 Introduction 1 1.2 Rhinoceros population change 1 1.2.1 Asian rhino species 2 1.2.2 African rhino species 4 1.3 Private management of the white rhino 9 1.4 Demand for horn 11 1.5 Current poaching epidemic 13 1.6 Value of the white rhino 16 1.6.1 Economic value 16 1.6.2 Ecosystem value 19 1.7 Anti-poaching strategies 21 1.7.1 Dehorning 21 1.7.2 Reducing consumer demand 33 1.7.3 Reducing illegal supply 34 1.7.4 Modifying poaching risk 36 1.8 Research aims and thesis outline 37 Chapter 2 – General methodology 39 2.1 Introduction 39 2.1.1 Study sites 39 2.2 Study populations 44 2.2.1 Rhino classification 45 2.2.2 Site A population 47 2.2.3 Site B population 48 iii Contents 2.2.4 Dehorning procedure 49 2.3 Data collection and analysis 50 2.3.1 Behavioural observations 50 2.3.2 Faecal sample collection 53 2.3.3 Data analysis 53 2.3.4 Ethics and permissions 54 Chapter 3 – The social effects of dehorning 55 3.1 Introduction 55 3.1.1 Social organisation 55 3.1.2 Agonistic social behaviour 57 3.1.3 Non-territorial dominance 60 3.1.4 Aims 62 3.2 Methods 63 3.2.1 Experimental design 63 3.2.2 Behavioural monitoring 63 3.2.3 Data analyses 68 3.3 Results 75 3.3.1 Agonistic social interaction rate 75 3.3.2 Social dominance hierarchy 80 3.4 Discussion 83 3.4.1 Agonistic social interaction rate 83 3.4.2 Social dominance hierarchy 87 3.4.3 Conclusion 90 Chapter 4 – Physiological effects of dehorning 91 4.1 Introduction 91 4.1.1 Function and mechanism of stress 91 4.1.2 Factors influencing stress 93 4.1.3 Validating rhino endocrine responses 96 4.1.4 Aims 98 4.2 Methods 99 4.2.1 Experimental design 99 4.2.2 Metabolite extraction and detection 100 4.2.3 Data analyses 107 4.3 Results 116 4.3.1 Assay validity 116 iv Contents 4.3.2 Analysis 1. Physiological differences in age and sex 117 4.3.3 Analysis 2. Physiological differences between populations 119 4.3.4 Analysis 3. Acute physiological response 120 4.3.5 Analysis 4. Chronic physiological response 120 4.3.6 Analysis 5. Multiple dehorning procedures – FCMs 122 4.3.7 Analysis 6. Multiple dehorning procedures – FPMs 123 4.3.8 Analysis 7. Multiple dehorning procedures – FAMs 126 4.4 Discussion 127 4.4.1 Physiological differences in age and sex 128 4.4.2 Physiological differences between populations 128 4.4.3 Acute physiological response 129 4.4.4 Chronic physiological response 130 4.4.5 Multiple dehorning procedures – FCMs 131 4.4.6 Multiple dehorning procedures – FPMs 131 4.4.7 Multiple dehorning procedures – FAMs 132 4.4.8 Limitations of the study and future work 133 4.4.9 Conclusion 135 Chapter 5 – Horn use in a non-social context 136 5.1 Introduction 136 5.1.1 Defensive behaviours 136 5.1.2 Resource access 137 5.1.3 Rubbing behaviour 138 5.1.4 Aims 140 5.2 Methods 140 5.2.1 Experimental design 140 5.2.2 Digging 141 5.2.3 Rubbing 141 5.2.4 Data Analysis 143 5.3 Results 144 5.3.1 Horn digging 144 5.3.2 Horn rubbing 147 5.4 Discussion 151 5.4.1 Conclusion 152 Chapter 6 – Eliciting rhinoceros avoidance behaviour using novel deterrents 153 6.1 Introduction 153 v Contents 6.1.1 Poaching risk 153 6.1.2 Mechanism of deterrents 155 6.1.3 Usage of deterrents 156 6.1.4 Aerial deterrents 157 6.1.5 Olfactory deterrents 158 6.1.6 Aims of the study 159 6.2 Methods 160 6.2.1 Experimental design 160 6.2.2 Acoustic deterrents 163 6.2.3 Drone deterrent 165 6.2.4 Olfactory deterrent 167 6.2.5 Data analysis 168 6.3 Results 169 6.3.1 Acoustic deterrent 169 6.3.2 Drone deterrent 170 6.3.3 Olfactory deterrent 173 6.4 Discussion 174 6.4.1 Effectiveness of the acoustic deterrent 175 6.4.2 Effectiveness of the drone deterrent 176 6.4.3 Effectiveness of the olfactory deterrent 179 6.4.4 Conservation implications 180 6.4.5 Conclusion 182 Chapter 7 – General discussion 183 7.1 Introduction 183 7.2 General discussion 183 7.3 Conservation implications 187 7.4 Opportunities for future research 192 7.5 Final statement 194 References 195 Appendix 1 – Other social behaviours 247 Appendix 2 – Social behaviour data 252 Appendix 3 – Inter-calf Intervals in Site A 258 vi List of Tables List of Tables Table 1.1. Major changes in policy affecting the domestic and international trade in rhino horn. 31 Table 2.1. Floristically important taxa of the Central Bushveld Bioregion. 41 Table 2.2. Presence and absence of medium to large bodied (> 10 kg) grazers and browsers at Site A and Site B. 42 Table 2.3. Terms and definitions as defined by the African Rhino Specialist Group (AfRSG). 45 Table 2.4. Population characteristics of rhinos at Site A. 48 Table 3.1. The behavioural states of the focal subjects were classified into one of several exclusive categories. 65 Table 3.2. Social behaviours recorded by all occurrences sampling. 65 Table 3.3. Age-sex class of subjects in the dehorned population (Site A) and horned population (Site B). 69 Table 3.4. Total effective observation length of subjects pre- and post- a dehorning procedure under any group composition and as a stable sub-grouping of six individuals. 72 Table 3.5. Total observation lengths for all observation sessions in which six stable rhinos were together as a stable long-term grouping. 74 Table 3.6. Statistical output from GLMM analysis indicating the effect of horn status on the rate of agonistic social interactions (n = 45, obs = 168). 76 Table 3.7. Statistical output from GLM analysis indicating the effect of sex-age class and horn status on the rate of agonistic social interactions (n = 45, obs. = 45). 76 Table 3.8. Social breakdown of dyadic agonistic interactions between a stable group of six subadult rhinos prior to and after their dehorning. 81 Table 4.1. Assay protocol flow chart for endocrine analyses. 103 Table 4.2. Cross-reactivates listed as a percentage of the actual concentration of cross- reactant in the sample against the level detected in the assay.
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