FACTORS AFFECTING THE SIZE AND DISTRIBUTION OF LARGE HERBIVORES IN KAFUE NATIONAL PARK, ZAMBIA by Wigganson Matandiko A dissertation submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy in Fish and Wildlife Biology MONTANA STATE UNIVERSITY Bozeman, Montana May 2016 ©COPYRIGHT by Wigganson Matandiko 2016 All Rights Reserved ii ACKNOWLEDGEMENT I wish to thank my graduate committee, Professor Scott Creel (Chair), Dr. Steve Cherry (Mathematics & Statistics), Dr. Matthew Becker, (CEO – Zambia Canivore Program), Dr. Dave A. Christianson (University of Arizona) for their constructive criticism that has led to the work and results described in this dissertation. I especially thank Professor Creel for his patient guidance. I thank Dr. Matthew Becker and Rachel McRobb who planted the idea of attending graduate School in the United States of America. Dr. Becker worked his way to present a strong case for Fulbright scholarship application. I thank University of Zambia, Schools of Veterinary Medicine and Agriculture Sciences for opening doors to their laboratory facilities. I thank Dr. Paul Schuette who is the backbone of the study design in my research. I thank my colleagues, Elias Rosenblatt, Egil Droge, Jassiel M’soka for their available help. I also wish to thank David McGuire for his assistance in the Creel laboratory. I thank Drs. Mwamba Sichande, Kambwiri Banda, Innocent N’gombwa and Jackson Katampi for participating in my research activities. I want to thank the church family at the Bozeman Adventist Church under the able leadership of Pastor Jim Jenkins and my host family of Mr. and Mrs. Kelly Coffin who took me in as one of their own. The following also helped in data collection and / or organization: Evans Nsende, Rachel Sullivan, Annie Sikanwe, Hannah Rasker, Carolyn Sanguinetti, Brandon Nickerson, Pearson Musonda, Angelina Kelly, Paige Mascol, Aubrey Labarre, Brianna Willey, Samantha Smith, Katelyn Belleville, Wigganson (Jr) Matandiko, Kelvin Yande, Albert Mwangana, Eric Przybyszewski, Teddy Mukula, and others not mentioned. iii TABLE OF CONTENTS 1. FACTORS AFFECTING THE SIZE AND DISTRIBUTION OF LARGE HERBIVORE HERDS IN KAFUE NATIONAL PARK, ZAMBIA: ORGANIZATION OF THE RESEARCH AND THESIS. .......................... 1 2. FACTORS AFFECTING LARGE HERBIVORE DISTRIBUTION AND DENSITY IN KAFUE NATIONAL PARK, ZAMBIA ....................................... 5 2.1. Introduction .............................................................................................................. 5 A. Abiotic ............................................................................................................. 7 B. Biotic ............................................................................................................... 8 C. Anthropogenic Effects..................................................................................... 9 2.2. Materials and Methods ............................................................................................. 9 2.2.1. Study Area ................................................................................................. 9 2.2.2. Study Design ............................................................................................ 11 2.2.2.1. Sampling Design ........................................................................... 11 2.2.3 Covariate Measurement ............................................................................ 13 2.2.3.1 Abiotic Factors ............................................................................... 13 2.2.3.1. a. Soil Chemistry and Texture .............................................. 13 2.2.3.1.b. Water Availability ............................................................. 14 2.2.3.2 Bottom-Up Processes ..................................................................... 15 2.2.3.2.a. Vegetation Communities and Edge Density ...................... 15 2.2.3.2.b. Vegetation Characteristics ................................................. 15 2.2.3.2.c. Anthropogenic Effects ....................................................... 15 2.2.3.3. Top-down Processes: Predation Risk ............................................ 16 2.2.4. Animal Counts ......................................................................................... 17 2.3. Distance Sampling Analysis .................................................................................. 18 2.3.1. Model Set and Model Selection ............................................................... 18 2.3.2. Analysis of the Complete Model Set for Two Years’ Data and a Partial Model Set for Three Years’ Data ........................................................... 21 2.4. Results .................................................................................................................... 21 2.4.1. Sample Size .............................................................................................. 21 2.4.2. Impala ...................................................................................................... 27 2.4.3. Puku ......................................................................................................... 28 2.4.4. Warthog.................................................................................................... 30 2.5. Discussion .............................................................................................................. 32 2.5.1. Impala ...................................................................................................... 33 2.5.2. Puku ......................................................................................................... 36 2.5.3. Warthog.................................................................................................... 38 2.6. Conclusion ............................................................................................................. 42 iv TABLE OF CONTENTS-CONTINUED 3. TOP-DOWN, BOTTOM-UP AND ANTHROPOGENIC EFFECTS ON LARGE HERBIVORE GROUP SIZES IN KAFUE NATIONAL PARK, ZAMBIA .................................................................................... 44 3.1. Introduction ............................................................................................................ 44 3.2. Methods.................................................................................................................. 47 3.2.1. Study Area................................................................................................. 47 3.2.2. Sampling Design and Observations of Group Size ............................................................................... 48 3.2.3. Bottom-up Limiting Factors .................................................................... 51 3.2.4. Anthropogenic Limiting Factors .............................................................. 53 3.2.5. Top-down Limiting Factors: Predation Risk from Lions ................................................................... 53 3.3. Statistical Methods ................................................................................................. 54 3.4. Results .................................................................................................................... 56 3.4.1. Impala ...................................................................................................... 56 3.4.2. Puku ......................................................................................................... 59 3.4.3. Warthog.................................................................................................... 61 3.5. Discussion .............................................................................................................. 64 4. COMPARATIVE ANALYSIS OF ECOLOGICAL AND ANTHROPOGENIC VARIABLES AFFECTING DISTRIBUTION AND SIZE OF LARGE HERBIVORES IN KAFUE NATIONAL PARK, ZAMBIA. ................................................................................... 67 4.1. Introduction ............................................................................................................ 67 4.2. Comparison of Effects on Herd Distribution and Herd Sizes ................................................................................. 69 4.4. Conclusion ............................................................................................................. 77 REFERENCES ................................................................................................................. 78 APPENDIX A: Dedication ................................................................................................90 v LIST OF TABLES Table Page 2.1. Number of observations and range of group sizes for herbivore species analyzed using distance sampling. ....................................................... 22 2.2 (a) and (b). Distance sampling models with results from model selection using Akaike’s information criteria ................................................... 23 2.3 (a) and (b). Coefficient estimates of impala, puku and warthog herd density from highest ranking models. ....................................................... 25 3.1. Effects on impala herd size from stepwise positive Poisson GLM. ................. 58 3.2 Effects on puku herd size from stepwise positive Poisson GLM. ..................... 60 3.3. Effects on warthog group size from stepwise positive Poisson GLM. ............. 62 4.1. Summary of effects on the density of distribution of herds (HD) and herd size (HS) for impala, puku