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OPTIMAL FORAGING on the ROOF of the WORLD: a FIELD STUDY of HIMALAYAN LANGURS a Dissertation Submitted to Kent State University

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OPTIMAL FORAGING on the ROOF of the WORLD: a FIELD STUDY of HIMALAYAN LANGURS a Dissertation Submitted to Kent State University OPTIMAL FORAGING ON THE ROOF OF THE WORLD: A FIELD STUDY OF HIMALAYAN LANGURS A dissertation submitted to Kent State University in partial fulfillment of the requirements for the degree of Doctor of Philosophy by Kenneth A. Sayers May 2008 Dissertation written by Kenneth A. Sayers B.A., Anderson University, 1996 M.A., Kent State University, 1999 Ph.D., Kent State University, 2008 Approved by ____________________________________, Dr. Marilyn A. Norconk Chair, Doctoral Dissertation Committee ____________________________________, Dr. C. Owen Lovejoy Member, Doctoral Dissertation Committee ____________________________________, Dr. Richard S. Meindl Member, Doctoral Dissertation Committee ____________________________________, Dr. Charles R. Menzel Member, Doctoral Dissertation Committee Accepted by ____________________________________, Dr. Robert V. Dorman Director, School of Biomedical Sciences ____________________________________, Dr. John R. D. Stalvey Dean, College of Arts and Sciences ii TABLE OF CONTENTS LIST OF FIGURES ............................................................................................... vi LIST OF TABLES ............................................................................................... viii ACKNOWLEDGEMENTS .....................................................................................x Chapter I. PRIMATES AT THE EXTREMES ..................................................1 Introduction: Primates in marginal habitats ......................................1 Prosimii .............................................................................................2 New World monkeys ........................................................................3 Cercopithecinae.................................................................................3 Colobinae and apes ...........................................................................5 The gray langur: A brief history of a very adaptable colobine .........6 Himalayan gray langurs ....................................................................9 Goals of the project .........................................................................12 II. STUDY SITE AND SUBJECTS ....................................................13 Langtang National Park, Nepal .......................................................13 Vegetation types of main study area ...............................................15 Climate of main study area .............................................................17 Mammalian fauna of main study area .............................................17 Study subjects .................................................................................19 Taxonomic identification of study subjects ....................................21 III. DIET, ACTIVITY PATTERNS, AND RESOURCES ...................24 INTRODUCTION ..........................................................................24 METHODS .....................................................................................25 Activity and feeding data ................................................................25 Classification of dietary items ........................................................27 Phenology .......................................................................................28 Data analysis ...................................................................................30 RESULTS .......................................................................................31 Activity patterns ..............................................................................31 Plant production ..............................................................................32 iii Diet ..................................................................................................35 Seasonal diet ...................................................................................38 Food selection .................................................................................40 Daily path lengths ..........................................................................43 DISCUSSION ................................................................................43 IV. EXTRACTIVE FORAGING AND INTELLIGENCE ..................51 INTRODUCTION ..........................................................................51 METHODS .....................................................................................56 Data collection ................................................................................56 Data analysis ...................................................................................58 RESULTS .......................................................................................59 Classes of extractive foraging in Himalayan langurs .....................59 Correlations: food availability and extractive foraging ..................66 Extractive foraging in other colobines ............................................68 DISCUSSION .................................................................................68 V. OPTIMAL FORAGING: CLASSICAL PREY MODEL ...............77 INTRODUCTION ..........................................................................77 METHODS .....................................................................................83 Behavioral observations ..................................................................83 Nutritional analysis and currencies for the model ..........................84 The model .......................................................................................87 Model predictions and statistics ......................................................90 RESULTS .......................................................................................93 Threshold for dropping items and the zero-one rule .......................93 Preference for profitable food types .............................................112 Increased selectivity at higher encounter rates .............................114 Selectivity independent from abundance of low-ranking food .....116 Comparison of currencies .............................................................118 Conformance of model assumptions and predictions ...................120 DISCUSSION ...............................................................................120 VI. OPTIMAL FORAGING: SOCIAL PREY MODEL ....................131 INTRODUCTION ........................................................................131 METHODS ...................................................................................135 Behavioral observations ................................................................135 Nutritional analysis and currency .................................................136 The model .....................................................................................138 Model predictions and statistics ....................................................138 iv RESULTS .....................................................................................143 Patch use strategy, residence time and travel time .......................143 Social foraging and the expanding specialist strategy ..................145 Number of foragers within patch and gain rate at switch .............148 DISCUSSION ...............................................................................150 VII. CONCLUDING REMARKS ........................................................154 Introduction ...................................................................................154 Diet, activity patterns, and resources ............................................155 Colobine cognition ........................................................................157 Optimal foraging theory ................................................................160 Himalayan langurs, foraging theory, and human evolution ..........162 REFERENCES ....................................................................................................166 v LIST OF FIGURES 1. Topographical map of main study area .............................................................14 2. Woody habitats of main study area ...................................................................16 3. Temperature and precipitation at Ghore Tabela, Nepal ....................................18 4. F troop near Langtang Monastery .....................................................................20 5. Variation in forearm coloration in F troop ........................................................23 6. Phenology of broad-leaved vegetative structures at Langtang .........................33 7. Phenology of reproductive plant parts at Langtang ..........................................34 8. Seasonal daily paths for Himalayan langurs .....................................................44 9. Extracted foods of Himalayan langurs ..............................................................61 10. Percent contribution of food types to extractive foraging ..............................62 11. Juvenile langur digging ...................................................................................63 12. Adult female with underground storage organ ...............................................64 13. MEO profitability threshold for juveniles ......................................................106 14. MEH profitability threshold for juveniles ......................................................106 15. CP profitability threshold for juveniles .........................................................107
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