1 Disturbance in the Udzungwas: Responses of Monkeys and Trees to Forest Degradation ANDREW ROBERT MARSHALL Ph.D. Centre for Ecology, Law and Policy Environment Department January 2007 2 Abstract Tropical forests are one of the world’s most threatened biomes. As tropical forests are increasingly destroyed and degraded, there is increasing need for research into the implications for rare species and habitats, and for conservation management. This thesis is a series of four papers investigating species and community responses in the Udzungwa Mountains, an area of international importance for biodiversity. The focus is on both rare species and potential indicators of habitat degradation. The most direct impact of human disturbance is on vegetation structure. However the response of most tropical plant species to disturbance has not been quantified. The Udzungwa area contains many rare and restricted range plant species, and furthermore, is arguably Africa’s most important single site for primate conservation. Primates also have high potential as indicators. In particular, diurnal monkeys are easily observed, are mostly dependent on tropical forests, and may show a diverse range of demographic and behavioural responses to disturbance. Human impacts on the monkeys and trees of the Udzungwas are here assessed at two levels: 1) habitat loss, and 2) habitat degradation. The first analytical chapter (Chapter 2) investigates the relationship between monkey species richness and forest fragment size among 22 sites. Multivariate techniques are used to consider the relationship together with other confounding variables. The results show that there is a log-linear species-area relationship, highlighting the importance of large forests (above 150 km2) for biodiversity conservation. The results also suggest that hunting and isolation have further influenced the species composition, particularly in the smaller fragments. The second analytical chapter (Chapter 3) assesses vegetation responses to disturbance using 120 plots, along six transects in the heavily disturbed lowland forest of Matundu. Multivariate analyses are used to assess variation in community composition, species abundance, stem density and diversity in relation to disturbance, environmental and topographic variables. With the exception of diversity, all measures of vegetation structure and composition are shown to have been affected by disturbance. In particular, rare species diversity was negatively related to disturbance, and also the presence of large animal paths. Therefore both humans and elephants seem to have had a large impact on the Matundu ecosystem. Three common tree species, Funtumia africana, Vangueria volkensii and Parinari excelsa, that have significant negative correlation with disturbance, could be used as indicators of forest health. Several environmental and topographical correlates with vegetation structure and composition are also identified. The third analytical chapter (Chapter 4) is a methodological chapter reviewing techniques for estimating density of clustered animals. This is included in the thesis for use in the monkey-habitat analysis in Chapter 5, and because of the continuing intense debate regarding method selection for primates. The aim of the chapter is to develop a simple guide to method selection. It begins with a 3 summary of the main debate, and an introduction to four alternative methods that are currently employed for surveying primates. The main controversy in the literature surrounds the debate over prioritising for mathematical framework (perpendicular methods) or for minimising correction factors (animal-observer methods). Three of the four methods have shown reasonable accuracy compared to known primate densities, and the fourth has yet to be tested. Perpendicular methods are the most desirable given appropriate field conditions. However problems arising from the five criteria of visibility, habituation of animals, cluster spread, and available resources, may often preclude their use. Finally, chapter 5 looks at the relationship between monkey relative abundance, density and social grouping versus habitat. Data are presented on four monkey species (Udzungwa red colobus Procolobus gordonorum, Angolan black and white colobus Colobus angolensis palliatus, Sykes monkey Cercopithecus mitis subsp. and yellow baboon Papio cynocephalus) counted along the six line transects used for vegetation survey in Chapter 3. In accordance with ecological theory, the four species respond to habitat disturbance as expected from their dietary specialisation, geographic range, and from previous studies on these and closely related species. The Udzungwa endemic red colobus showed the closest relationship with habitat, and may have the most potential as an indicator of forest habitat quality of all taxa in the Udzungwas. Monkey community composition is also suggested as a good indicator of habitat quality. The results suggest that those areas in the Udzungwa lowlands that contain high densities and large groups of red and black and white colobus monkeys contain the best quality forest in terms of vegetation structure and composition. Those containing mainly Sykes monkeys are the poorest quality. Overall the thesis shows that monkey and tree communities in the Udzungwa Mountains have been negatively impacted by human activities. However, the results also emphasise the importance of disturbed forests for future conservation of biodiversity. Given that most tropical forests have been disturbed to some extent, management of these areas is of paramount importance, as discussed in Chapter 6. The indicator species and communities identified here may be useful for this, both for identifying priority conservation areas and for monitoring forest recovery. 4 Contents Abstract 2 Contents 4 List of Tables 8 List of Figures 10 Preface 12 Acknowledgements 13 Author’s Declaration 14 Chapter 1 – Introduction to Thesis 16 Tropical Forest Disturbance and Loss 17 Selection of Study Site 18 Eastern Arc Mountains 18 Udzungwa Mountains 19 Matundu Forest 21 Selection of Study Taxa 21 Thesis Aims 24 Chapter Outlines and Objectives 24 Chapter 2 - The Species-Area Relationship in a Threatened Monkey Community: 24 Controlling for Confounding Variables Chapter 3 - Tree Species and Community Gradients for Assessing Habitat 25 Disturbance Chapter 4 – Line-Transect Methods for Estimating Density of Clustered Animals: A 25 Review and Guide with Lessons from the Primates Chapter 5 – Monkeys as Indicators in Lowland Udzungwa 26 References 26 Chapter 2 - The Species-Area Relationship in a Threatened Monkey Community: 32 Controlling for Confounding Variables Abstract 33 Introduction 33 Aim and Objectives 35 Methods 36 Study Areas and Species 36 Data Collection 37 Statistical Analyses 38 Results 39 Discussion 42 5 Confounding Variables for the Species-Area Relationship 42 Status of Rare Monkeys in the Udzungwas 44 The Species-Area Relationship and Landscape Management 45 Acknowledgements 46 References 46 Chapter 3 - Tree Species and Community Gradients for Assessing Habitat 51 Disturbance Abstract 52 Introduction 52 Aim and Objectives 54 Methods 55 Study Area 55 Vegetation Data 56 Predictor Variables 56 Statistical Analyses 57 Results 59 Habitat Description 59 Community Composition 61 Density and Diversity 62 Rare Species 64 Discussion 66 Disturbance and Gradients 66 Conservation of Matundu Forest 68 Acknowledgements 69 References 69 Appendix 75 Chapter 4 - Line-Transect Methods for Estimating Density of Clustered Animals: A 80 Review and Guide with Lessons from the Primates Abstract 81 Introduction 81 Rationale for Line-Transect Distance Sampling 82 Points of Contention 83 Aims and Objectives 84 Assumptions and Limitations 85 Cluster Parameters 88 Cluster Size 88 Cluster Centre / Spread 89 6 “Accepted” Methods 90 Perpendicular Distance To Cluster Methods 90 a) Perpendicular Modelling of Cluster Centres (Traditional Detection Function 90 Modelling) b) Perpendicular Modelling of Centre of Measurable Individuals 92 c) Strip Transects (Truncated Distance Method or Kelker Method) 93 Perpendicular Distance to First Individual Methods 96 Non-Perpendicular Distance Method 96 d) Animal-Observer Distance Method (King’s Method) 96 Comparison of Methods 98 General Discussion 99 Method Selection 99 Alternative Methods 102 Acknowledgements 103 References 104 Chapter 5 - Monkeys as Indicators in Lowland Udzungwa 108 Abstract 109 Introduction 109 Aim and Objectives 111 Methods 112 Study Area 112 Transect Walks and Density Estimation 113 Social Group Size 115 Habitat Variables 116 Statistical Analyses 118 Results 119 Coarse-Scale Distribution 119 Fine-Scale Distribution 120 Social Grouping 122 Density and Population 123 Discussion 125 Responses to Forest Disturbance 125 1) Specialists 125 2) Generalists 127 Monkeys as Indicators 128 Conservation of Matundu Forest 130 Acknowledgements 130 References 131 7 Appendices 137 Chapter 6 – Summary Discussion 138 Summary of Relationships to Habitat Disturbance and Loss 139 Indicators 140 Environmental Relationships 142 Future Directions for Udzungwa 145 Connectivity 145 Temporal Trends 145 Increasing Human Population Pressure 146 Collaboration 146 References 147 8 List of Tables Chapter 1 No Tables Chapter 2 Table 1. Description of forest fragments used for investigating the species-area relationship in the Udzungwa Mountains. Table 2. Monkey species present in the study forests. Table 3. GLM analyses and Spearman correlation of species richness versus log forest