CHIMPANZEE (PAN TROGLODYTES ELLIOTI) ECOLOGY IN A NIGERIAN MONTANE FOREST A thesis submitted in partial fulfilment of the requirements for the Degree of Doctor of Philosophy in Ecology in the University of Canterbury by P. E. Dutton University of Canterbury 2012 Acknowledgements Firstly, I would like to thank Dr. Hazel Chapman as supervisor and the director of the Nigerian Montane Forest Project (NMFP) as without her support this research would not have been possible. Furthermore, I would like to thank the NMFP staff for their dedication towards this research, without them my time at Ngel Nyaki would have been very difficult. I would also like to thank Dr. Elena Moltchanova for providing statistical support for this research. I would like to thank Primate Conservation Inc. (PCI) for its financial support as well as the North of England Zoological Society (NEZS), Nexen Nigeria and A. G. Leventis Foundation for their contributions towards the project. Lastly, I would like to thank Annelies Vranken for her tolerance and understanding during my university studies. TABLE OF CONTENTS TABLE OF CONTENTS .................................................................................................... III LIST OF FIGURES ......................................................................................................... VIII LIST OF TABLES ............................................................................................................ XIV ABSTRACT ....................................................................................................................... XVI Chapter 1: Introduction to Pan troglodytes ellioti and Ngel Nyaki Forest Reserve ......... 1 1.1 INTRODUTION ................................................................................................... 1 1.1.1 Pan troglodytes............................................................................................. 1 1.1.2 Pan troglodytes ellioti genetic history.......................................................... 1 1.1.3 History of Pan troglodytes ellioti phylogeny and conservation ................... 3 1.1.4 Distribution and densities of Pan troglodytes ellioti .................................... 3 1.1.5 Previous ecological research on Pan troglodytes ellioti .............................. 4 1.2 AIMS OF THIS STUDY ....................................................................................... 6 1.3 GOALS OF THIS STUDY ................................................................................... 6 1.4 STUDY SITE ........................................................................................................ 7 1.4.1 Description of the region and study area ...................................................... 7 1.4.2 Climate ....................................................................................................... 10 1.4.3 Fauna and Flora .......................................................................................... 14 1.4.4 Human impact ............................................................................................ 14 1.4.4.1 Snares for bush meat .......................................................................... 14 1.4.4.2 Cutting of vegetation for food and construction ................................. 15 1.4.4.3 Land use around the forest ................................................................. 16 1.4.4.4 Grazing pressures ............................................................................... 17 1.4.4.5 Burning regimes ................................................................................. 18 1.4.4.6 Trails ................................................................................................... 19 1.4.5 Direct impact on chimpanzees ................................................................... 19 1.5 REFERENCES .................................................................................................... 20 APPENDIX 1.1 ...................................................................................................................... 25 Chapter 2: Density of Pan troglodytes ellioti in Ngel Nyaki Forest Reserve ................... 26 2.1 ABSTRACT ........................................................................................................ 26 2.2 INTRODUCTION ............................................................................................... 26 2.3 METHODS .......................................................................................................... 27 2.3.1 Study site .................................................................................................... 27 2.3.2 Pilot Study .................................................................................................. 28 2.3.3 Data collection ............................................................................................ 29 2.3.4 Sampling procedure .................................................................................... 32 2.3.5 Sampling designs ........................................................................................ 33 2.3.5.1 Standing Crop Nest Counts ................................................................ 33 2.3.5.2 Marked nest counts ............................................................................. 34 2.3.5.3 Distance sampling .............................................................................. 34 2.3.5.4 Observation ........................................................................................ 35 2.3.5.5 Nest Decay .......................................................................................... 35 2.4 RESULTS ............................................................................................................ 37 2.4.1 Density estimates from Standing Crop Nest Counts (SCNC) .................... 37 2.4.2 Density estimates for Marked Nest Counts (MNC) ................................... 38 iii 2.4.3 Distance sampling ...................................................................................... 39 2.4.4 Direct observation ...................................................................................... 41 07-April 2010; 12:10-1:14 pm; N 07 08.911’ E 11 04.637’ .......................... 41 01-May 2010; 2:50-2:57 pm ........................................................................... 41 07-May 2010; 9:25-11:10 am; N 07 05.005’ E 11 02.915’. .......................... 42 16-March 2011; N 07 05.213’ E 11 02.746’ .................................................. 42 10-October 2009-20-May 2011 ...................................................................... 42 2.5 DISCUSSION and CONCLUSIONS .................................................................. 43 2.5.1 Standing Crop Nest Count .......................................................................... 43 2.5.2 Marked Nest Count .................................................................................... 43 2.5.3 Distance sampling ...................................................................................... 44 2.5.4 Nest discovery ............................................................................................ 44 2.5.5 Nest density ................................................................................................ 45 2.6 REFERENCES .................................................................................................... 47 Chapter 3: Nesting ecology .................................................................................................. 52 3.1 ABSTRACT ........................................................................................................ 52 3.2 INTRODUCTION ............................................................................................... 52 3.3 METHODS .......................................................................................................... 53 3.3.1 Locating nests ............................................................................................. 53 3.3.2 Measuring variables ................................................................................... 53 3.3.3 Analysis ...................................................................................................... 54 3.3.3.1 Descriptive nest variables .................................................................. 54 3.3.3.2 Nesting areas ...................................................................................... 54 3.3.3.3 Transect influence ............................................................................... 55 3.3.3.4 Nest variable associations .................................................................. 55 3.4 RESULTS ............................................................................................................ 58 3.4.1 Descriptive nest variables ........................................................................... 58 3.4.2 Nesting areas .............................................................................................. 61 3.4.3 Transect influence ...................................................................................... 65 3.4.4 Nest variable associations .......................................................................... 69 3.5 DISCUSSION ..................................................................................................... 72 3.5.1 Descriptive nest variables ........................................................................... 72 3.5.1.1