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Theropithecus Gelada) on an Intact Afro-Alpine Grassland at Guassa, Ethiopia ______ LONG-TERM RANGING PATTERNS OF WILD GELADA MONKEYS (THEROPITHECUS GELADA) ON AN INTACT AFRO-ALPINE GRASSLAND AT GUASSA, ETHIOPIA ____________________________________ A Thesis Presented to the Faculty of California State University, Fullerton ____________________________________ In Partial Fulfillment of the Requirements for the Degree Master of Arts in Anthropology ____________________________________ By Cha Moua Thesis Committee Approval: Associate Professor Peter J. Fashing, Chair Associate Professor Nga Nguyen, Department of Anthropology Associate Professor Elizabeth G. Pillsworth, Department of Anthropology Fall, 2015 ABSTRACT Long-term studies of animal ranging ecology are critical to understanding how animals utilize their habitat across space and time. Although gelada monkeys (Theropithecus gelada) inhabit an unusual, high altitude habitat that presents unique ecological challenges, no long-term studies of their ranging behavior have been conducted. To close this gap, I investigated the daily path length (DPL), annual home ranges (95%), and annual core areas (50%) of a band of ~220 wild gelada monkeys at Guassa, Ethiopia, from January 2007 to December 2011 (for total of n = 785 full-day follows). I estimated annual home ranges and core area using the fixed kernel reference (FK REF) and smoothed cross-validation (FK SCV) bandwidths, and the minimum convex polygon (MCP) method. Both annual home range (MCP - 2007: 5.9 km2; 2008: 8.6 km2; 2009: 9.2 km2; 2010: 11.5 km2; 2011: 11.6 km2) and core area increased over the 5-year study period. The MCP and FK REF generated broadly consistent, though slightly larger estimates that contained areas in which the geladas were never observed. All three methods omitted one to 19 sleeping sites from the home range depending on the year. Thus, neither the MCP nor fixed kernel estimators were more accurate than the other. Similarly, mean annual DPL (± SE m) increased over the study period (2007: 2,848±57 m; 2008: 3,339±65 m; 2009: 3,272±72 m; 2010: 3,835±80 m; 2011: 4,100±86 m). In general, the geladas showed remarkable variation in daily, monthly, and annual ii DPL. I also investigated the effects of movement across uneven topography on DPL, and I discuss the ecological implications of these findings. I compare the ranging behavior of geladas at Guassa to (a) geladas at other study sites, (b) to Papio (baboon) species, (c) to both terrestrial and arboreal primates, and (d) to grazing ungulates. The extensive inter- annual variability in ranging patterns in this study demonstrates the importance of long- term monitoring for wild nonhuman primates and its implications for conservation policy. iii TABLE OF CONTENTS ABSTRACT ................................................................................................................... ii LIST OF TABLES ......................................................................................................... vi LIST OF FIGURES ....................................................................................................... vii ACKNOWLEDGMENTS ............................................................................................. viii Chapter 1. INTRODUCTION ................................................................................................ 1 Research in Animal Ranging Ecology .................................................................. 1 The Importance of Long-Term Ranging Studies ........................................... 4 Gelada Monkeys as a Model System ............................................................. 6 Gelada Monkeys Study Site, Guassa, Ethiopia .............................................. 8 Objectives of the Study .................................................................................. 9 2. METHODS ........................................................................................................... 11 Study site............................................................................................................... 11 The Qero System and its Future .................................................................... 12 Study Subjects ................................................................................................ 13 Data Collection and Analysis ............................................................................... 14 Daily Ranging Data ....................................................................................... 15 Ranging Analysis: Calculation of Daily Path Lengths .................................. 17 Ranging Analysis: Amending Daily Path Lengths to Account for Changes in Altitude ................................................................................................. 18 Home Range Analysis .......................................................................................... 19 Home Range Estimator: Minimum Convex Polygon .................................... 20 Home Range Estimator: Fixed kernel ............................................................ 23 Autocorrelation: Implications on Home Range Analysis .............................. 27 Statistical Analysis ................................................................................................ 30 3. RESULTS ............................................................................................................. 32 Annual Home Range Estimates: MCP.................................................................. 32 Annual Home Range Estimates: FK REF ...................................................... 33 Annual Home Range Estimates: FK SCV ..................................................... 41 iv Comparison of Annual Home Range Across Methods ......................................... 43 Trends in Annual Home Range ..................................................................... 43 Annual Core Area: Use and Trends ............................................................... 46 Ranging Patterns: Daily, Monthly, and annual trends in DPL ............................. 49 Monthly Mean DPL ....................................................................................... 51 Annual Mean DPL ......................................................................................... 53 4. DISCUSSION ....................................................................................................... 56 Summary of Findings............................................................................................ 56 Evaluation of the MCP Method ..................................................................... 57 Evaluation of the Kernel Estimators .............................................................. 60 Implications and Suggestions for Future Research ........................................ 63 Comparison of Gelada Monkey Ranging Behavior Across Sites ......................... 67 How do the Annual Home Range Estimates of Geladas at Guassa Compare to Those for Geladas at Other Sites?......................................................... 68 How do Geladas Utilize Their Home Range at Guassa and How Does it Compare to That of Geladas at Other Sites? ............................................. 70 How do the DPL of Geladas at Guassa Compare to Those of Geladas at Other Sites? .............................................................................................. 71 Comparison of Gelada Monkey Ranging Behavior Across Taxa......................... 71 Comparison of Gelada Ranging Behavior to Papio Species ......................... 72 Comparison of Gelada Ranging Behavior to Terrestrial Nonhuman Primate Species ...................................................................................................... 77 Comparison of Gelada Ranging Behavior to Arboreal Nonhuman Primate Species ...................................................................................................... 83 Comparison of Gelada Ranging Behavior to Terrestrial Ungulate Species .. 89 Implications of Inhabiting in a Topographically Variable Environment on Calculations of Distance Traveled ..................................................................... 91 Ecological Implications of Movement Across Uneven Topography............. 92 Critiques of the Altitudinal Change Formula ................................................ 96 Conclusions ........................................................................................................... 97 APPENDIX: ADDING ERROR TO USER IDENTIFIED DUPLICATE PAIRS ...... 100 BIBLIOGRAPHY .......................................................................................................... 105 v LIST OF TABLES Table Page 2.1 Results of Autocorrelation Analysis ....................................................................... 30 3.1 Comparison of Annual Home Range Estimates for MCP ...................................... 33 3.2 Core Areas (50%) and Annual Home Ranges (95%) Based on the FK 0.6*REF .................................................................................................................. 34 3.3 Core Areas (50%) and Annual Home Ranges (95%) Based on the FK SCV ......... 41 3.4 Monthly Mean DPL ± SE (m), Number of Full-days, and Range of DPL for Each and all Years. ............................................................................................ 52 4.1 Comparison of Gelada Monkey Ranging Patterns Across Sites ............................. 69 4.2 DPL, Home Range, and Core Area of Papio Species............................................. 75 4.3 DPL, Home Range,
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