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(Tarsius Pumilus) in CENTRAL SULAWESI, INDONESIA ALTITUDINAL EFFECTS ON THE BEHAVIOR AND MORPHOLOGY OF PYGMY TARSIERS (Tarsius pumilus) IN CENTRAL SULAWESI, INDONESIA A Dissertation by NANDA BESS GROW Submitted to the Office of Graduate Studies of Texas A&M University in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY Chair of Committee, Sharon Gursky-Doyen Committee Members, Michael Alvard Jeffrey Winking Jane Packard Head of Department, Cynthia Werner August 2013 Major Subject: Anthropology Copyright 2013 Nanda Bess Grow ABSTRACT Pygmy tarsiers (Tarsius pumilus) of Central Sulawesi, Indonesia are the only species of tarsier known to live exclusively at high altitudes. This study was the first to locate and observe multiple groups of this elusive primate. This research tested the hypothesis that variation in pygmy tarsier behavior and morphology correlates with measurable ecological differences that occur along an altitudinal gradient. As a response to decreased resources at higher altitudes and the associated effects on foraging competition and energy intake, pygmy tarsiers were predicted to exhibit lower population density, smaller group sizes, larger home ranges, and reduced sexually selected traits compared to lowland tarsiers. Six groups containing a total of 22 individuals were observed. Pygmy tarsiers were only found between 2000 and 2300 m, indicating allopatric separation from lowland tarsiers. As expected, the observed pygmy tarsiers lived at a lower density than lowland tarsier species, in association with decreased resources at higher altitudes. The estimated population density of pygmy tarsiers was 92 individuals per 100 ha, with 25 groups per 100 ha. However, contrary to expectation, home range sizes were not significantly larger than lowland tarsier home ranges, and average NPL was smaller than those of lowland tarsiers. The average home range size for the observed pygmy tarsiers was 2.0 ha, and the average nightly path length (NPL) was 365.36 m. Pygmy tarsiers exhibited a nonrandom, clumped distribution near forest edges. While insect abundance and biomass were found to decrease as altitude increased, insect abundance and biomass was higher along anthropogenic edges. Thus, tarsiers within the study area may mitigate the decreased availability of insects at high altitudes ii by remaining close to forest edges, which in turn may be related to smaller than expected home range sizes. Further, estimates of pygmy tarsier abundance may be inflated because of increased insect abundance along anthropogenic edges. Contrary to the prediction for smaller group sizes as a response to feeding competition, the observed pygmy tarsiers lived in relatively large groups with multiple adult males. However, in support of the prediction for energetic constraints on body proportions, the observed pygmy tarsiers did not exhibit sexually selected traits. The pygmy tarsiers exhibited low sexual dimorphism and small relative testes mass, a trend opposite from lowland tarsier species, which may indicate a constraint on the development of those traits. Considered together, these results suggest that the observed pygmy tarsiers have adapted to life in an environment with limited resources. Future studies should explore the possible contributing effects of seasonality and topography. iii DEDICATION This dissertation is dedicated to my mother and father, Drs. Arunee Grow and John Grow, for all their love and support through the years. Mom and Dad: you condemned me to become the third Dr. Grow, and for that I am grateful. iv ACKNOWLEDGEMENTS First and foremost, I would like to express my immense gratitude to my committee chair, Dr. Sharon Gursky-Doyen, for her support in all my research and educational pursuits. I would also like to thank my committee members, Dr. Michael Alvard, Dr. Jeffrey Winking, Dr. Jane Packard, for their guidance and support throughout the course of this research, and to Dr. David Carlson. I also want to extend my gratitude to the various funding sources for this research: the National Science Foundation Doctoral Dissertation Improvement Grant, Primate Conservation, Inc., Conservation International Primate Action Fund, American Society of Primatologists Conservation Small Grant, Explorers Club Exploration Fund Grant, and the Texas A&M University Department of Anthropology. Without the generous contributions from these agencies, this research would not have been possible. Thanks also go to my friends and colleagues and the faculty and staff in the Department of Anthropology for making my time at Texas A&M University a memorable and enjoyable experience. Finally, thank you to my family for their moral support. I would like to give special thanks to my mother and father for their encouragement, and to John Blong for always being there. v NOMENCLATURE A.S.L. Above Sea Level DBH Diameter at breast height HA Hectare M Meters MDE Mid-domain effect NPL Nightly path length (travel distance) TV Testicular Volume vi TABLE OF CONTENTS Page ABSTRACT ...................................................................................................................... ii DEDICATION .................................................................................................................. iv ACKNOWLEDGEMENTS ................................................................................................ v NOMENCLATURE .......................................................................................................... vi TABLE OF CONTENTS ................................................................................................. vii LIST OF FIGURES .......................................................................................................... xi LIST OF TABLES .......................................................................................................... xiii 1. INTRODUCTION AND LITERATURE REVIEW, ......................................................... 1 1.1 Research Objectives ............................................................................................. 1 1.2 Tarsier Classification and Biogeography ............................................................... 3 1.2.1 Tarsier Evolution and Phylogeny ................................................................ 3 1.2.2 Tarsier Diversity and Distribution ................................................................ 7 1.2.3 Sulawesian Tarsier Biogeography .............................................................. 8 1.2.4 History of Tarsius pumilus ........................................................................ 10 1.3 Altitudinal Variation .............................................................................................. 13 1.3.1 High Altitude Ecology ................................................................................ 13 1.3.2 Altitudinal Clines in Species Diversity and Abundance ............................. 14 1.3.2.1 The Mid-Domain Effect ...................................................................... 15 1.3.2.2 Species-Area Effect Hypothesis (Elevational Gradient)..................... 17 1.3.2.3 The Massenerhebung Effect of Elevational Vegetation Zones .......... 17 1.3.2.4 Geographic Isolation and the Rescue Hypothesis ............................. 19 1.3.3 Altitudinal Clines in Range Size ................................................................ 20 1.3.3.1 Range Size and Rapoport’s Rule of Species Distribution .................. 20 1.3.3.2 Body Size Clines ................................................................................ 21 1.4 Body Size Variation ............................................................................................. 21 1.4.1 Why Is Body Size Important? ................................................................... 21 1.4.2 The Relationship Between Body Size and Abundance ............................ 23 1.4.3 The Island Rule ......................................................................................... 24 1.4.4 Bergmann’s and Allen’s Rules .................................................................. 26 1.4.4.1 Predictions of Bergmann’s Rule and Corollaries ............................... 26 1.4.4.2 Support for Bergmann’s Rule and Corollaries ................................... 29 1.4.4.3 Evidence for Bergmann’s Rule and Corollaries Among Primates...... 30 1.4.4.4 Application to Altitudinal Gradients .................................................... 33 vii 1.4.4.5 The Thermoregulation Explanation for Bergmann’s Rule .................. 35 1.4.4.6 Alternative Mechanisms Behind Body Size Clines ............................ 39 1.4.4.7 The Heat Dissipation Hypothesis ....................................................... 39 1.4.4.8 The Dispersal Hypothesis .................................................................. 39 1.4.4.9 The Starvation Resistance (Seasonality) Hypothesis ........................ 40 1.4.4.10 The Resource Availability Hypothesis ............................................. 42 1.4.4.11 The Converse of Bergmann’s Rule .................................................. 43 1.4.5 Taxonomic Level of Clinal Variation ......................................................... 44 1.4.6 Are Body Size Clines Adaptive? ............................................................... 45 1.5 Altitudinal Effects on Primates ............................................................................. 45 1.5.1 Spatial and Temporal Variation in Primate Behavior and Morphology ..... 45 1.5.2 Foraging Behavior
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