SPATIAL AND TEMPORAL PATTERNS IN RESOURCE DISPERSION AND THE STRUCTURE OF RANGE USE AND CO-EXISTENCE IN A SOCIAL OMNIVORE CHLOROCEBUS AETHIOPS by ALAN SEAN BARRETT submitted in accordance with the requirements for the degree of DOCTOR OF PHILOSOPHY in the subject ENVIRONMENTAL MANAGEMENT at the UNIVERSITY OF SOUTH AFRICA SUPERVISOR: PROF L R BROWN CO-SUPERVISORS: PROF S P HENZI / PROF L BARRETT NOVEMBER 2009 Declaration I declare that SPATIAL AND TEMPORAL PATTERNS IN RESOURCE DISPERSION AND THE STRUCTURE OF RANGE USE AND CO-EXISTENCE IN A SOCIAL OMNIVORE CHLOROCEBUS AETHIOPS is my own work and that all the sources that I have used or quoted have been indicated and acknowledged by means of complete references. .................................................. .................................................. A.S. Barrett Date [email protected] Summary The movements of two vervet monkey troops were studied to determine whether they optimize their rate of food intake in relation to seasonal energy availability. The effect of variation in habitat structure on the troops’ foraging strategies while utilizing temporally and spatially distributed resources was determined. Troop home range boundaries were delineated, the various plant communities and species utilised by the troops identified and classified, and variations in home range and vegetation structure were reported. The diets of the troops were determined and compared. Effects of coexistence on competition were assessed. Vervet food trees were randomly selected, marked and seasonal phenological data collected. Samples of food items constituting the two troops diets were collected for energy analysis. Using geostatistical interpolation techniques, monthly energy values were extrapolated onto home range grids for the two vervet monkey troops. Grids were stored as database files that were interrogated through GIS simulation models. Using the stochastic processes inherent in Markov chain theory, a series of non-returning random walks were simulated for comparison to original routes taken by the two troops. Results from comparisons of home range energy, day range lengths and areas, shortest route energy to actual route energy, time spent in high energy areas, and energy utilisation from actual and randomly generated routes indicated that the two troops optimize resource energy available to them by adopting flexible foraging strategies. In environments where temporal and spatial variations in habitat structure affect the distribution of resources, it is essential that animals develop optimal foraging strategies to survive. For the two troops investigated, foraging strategies fluctuate between being time minimizers in more heterogeneous environments where resources are abundant, and energy maximisers in homogeneous environments where resources are constrained by low diversity and seasonality. Key Terms Animal movement; Range use; Plant communities; Phytosociology; Phenology; Diet; Caloric content; Energy; Bootstrapping; Vervet monkey; Interpolation; Foraging strategies. Page i Table of Contents Summary .............................................................................................................. i Key Terms ............................................................................................................ i Table of Contents ................................................................................................. ii List of Figures ...................................................................................................... vi List of Tables ..................................................................................................... xiii List of Appendices .............................................................................................. xv Acknowledgements ........................................................................................... xvi CHAPTER 1: INTRODUCTION ........................................................................... 1 CHAPTER 2: LITERATURE REVIEW ................................................................. 6 2.1 Loskop Dam Nature Reserve .................................................................... 6 2.2 Vegetation ................................................................................................. 7 2.3 Phytosociology .......................................................................................... 7 2.4 Phenology and energetic content of food .................................................. 8 2.5 Vervet monkeys ....................................................................................... 10 2.6 Habitat and dietary selectivity .................................................................. 11 2.7 Energy and ranging patterns ................................................................... 12 CHAPTER 3: STUDY SITE, SUBJECTS AND METHODS ............................... 17 3.1 Study Site ................................................................................................ 17 3.1.1 Location and size .............................................................................. 18 3.1.2 Topography ....................................................................................... 19 3.1.3 Geology ............................................................................................. 20 3.1.4 Pedology ........................................................................................... 26 3.1.5 Land types ........................................................................................ 26 3.1.6 Vegetation ......................................................................................... 40 Page ii 3.1.7 Fauna ................................................................................................ 44 3.1.8 Climate .............................................................................................. 44 3.2 Study subjects ......................................................................................... 46 3.3 Methods ................................................................................................... 47 3.3.1 Study site .......................................................................................... 48 3.3.2 Phytosociology .................................................................................. 50 3.3.3 Habitat and dietary selectivity ........................................................... 50 3.3.4 Phenology and energetic content of food .......................................... 51 3.3.5 Energy and ranging patterns ............................................................. 51 3.3.6 Statistics ............................................................................................ 52 CHAPTER 4: PHYTOSOCIOLOGY .................................................................. 53 4.1 Introduction .............................................................................................. 53 4.2 Methods ................................................................................................... 54 4.3 Results ..................................................................................................... 57 4.3.1 Donga home range classification ...................................................... 57 4.3.2 Description of the Donga home range plant communities ................. 61 4.3.3 Picnic home range classification ....................................................... 73 4.3.4 Description of the Picnic home range plant communities .................. 79 4.3.5 Community environmental ordination ................................................ 91 4.3.5.1 Donga home range ........................................................................ 91 4.3.5.2 Picnic home range ......................................................................... 93 4.3.5.3 Donga vs. Picnic ............................................................................ 95 4.3.6 Floristic analysis ................................................................................ 97 4.3.6.1 Donga home range ........................................................................ 98 4.3.6.2 Picnic home range ......................................................................... 99 4.3.6.3 Donga vs. Picnic .......................................................................... 101 4.4 Discussion ............................................................................................. 104 Page iii CHAPTER 5: HABITAT AND DIETARY SELECTIVITY .................................. 107 5.1 Introduction ............................................................................................ 107 5.2 Methods ................................................................................................. 108 5.3 Results ................................................................................................... 109 5.3.1 Plant community electivity ............................................................... 109 5.3.2 Diet .................................................................................................. 111 5.3.3 Troop seasonal food type selection................................................. 113 5.3.4 A comparison of the utilization of food types by the two troops ....... 115 5.3.5 Age-sex class seasonal food type selection .................................... 115 5.3.6 Across troop age-sex class seasonal food type selection ............... 118 5.3.7 Number of foraging individuals vs. variety of food types