PRIMATE FIELD STUDIES Series Editors: Robert W. Sussman, Washington University Natalie Vasey, Portland State University Series Editorial Board: Simon Bearder, Oxford-Brookes University Marina Cords, Columbia University Agustin Fuentes, Notre Dame University Paul Garber, University of Illinois Annie Gautier-Hion, Station Biologique de Paimpont Joanna Lambert, University of Wisconsin Robert D. Martin, Field Museum Deborah Overdorff, University of Texas Jane Phillips-Conroy, Washington University Karen Strier, University of Wisconsin Series Titles: The Spectral Tarsier Sharon L. Gursky, Texas A&M University Strategies of Sex and Survival in Hamadryas Baboons: Through a Female Lens Larissa Swedell, Queens College, The City University of New York The Behavioral Ecology of Callimicos and Tamarins in Northwestern Bolivia Leila M. Porter, Northern Illinois University The Socioecology of Adult Female Patas Monkeys and Vervets Jill D. Pruetz, Iowa State University Apes of the Impenetrable Forest: The Behavioral Ecology of Sympatric Chimpanzees and Gorillas Craig B. Stanford, University of Southern California PRIMATE FIELD STUDIES Many of us who conduct field studies on wild primates have witnessed a decline in the venues available to publish monographic treatments of our work. As researchers we have few choices other than to publish short tech- nical articles on discrete aspects of our work in professional journals. Also in vogue are popular expositions, often written by non-scientists. To counter this trend, we have begun this series. Primate Field Studies is a venue both for publishing the full complement of findings of long-term studies, and for making our work accessible to a wider readership. Interested readers need not wait for atomized parts of long-term studies to be published in widely scattered journals; students need not navigate the technical literature to bring together a body of scholarship better served by being offered as a cohesive whole. We are interested in developing monographs based on single or multi-species studies. If you wish to develop a monograph, we encourage you to contact one of the series editors. About the Editors: Robert W. Sussman (Ph.D. Duke University) is currently Professor of Anthropology and Environmental Science at Washington University, St. Louis, Missouri and past Editor-in-Chief of American Anthropologist, the flagship journal of the American Anthropological Association. His research focuses on the ecology, behavior, evolution and conservation of nonhuman and human primates, and he has worked in Costa Rica, Guyana, Panama, Madagascar and Mauritius. He is the author of numer- ous scientific publications, including Biological Basis of Human Behavior, Prentice Hall (1999), Primate Ecology and Social Structure (two volumes), Pearson Custom Publishing (2003), and The Origin and Nature of Sociality, Aldine de Gruyter (2004). Natalie Vasey (Ph.D. Washington University) is currently assistant profes- sor of anthropology at Portland State University in Portland, Oregon. Her work explores the behavioral ecology, life history adaptations, and evolu- tion of primates, with a focus on the endangered and recently extinct pri- mates of Madagascar. She has presented her research at international venues and published in leading scientific journals. She is dedicated to educating students and the public-at-large about the lifestyles and conser- vation status of our closest relatives in the animal kingdom. The Spectral Tarsier Sharon L. Gursky Texas A&M University First published 2007 by Pearson Education, Inc. Published 2016 by Routledge 2 Park Square, Milton Park, Abingdon, Oxon OX14 4RN 711 Third Avenue, New York, NY 10017, USA Routledge is an imprint of the Taylor & Francis Group, an informa business Copyright © 2007 Taylor & Francis. All rights reserved. All rights reserved. No part of this book may be reprinted or reproduced or utilised in any form or by any electronic, mechanical, or other means, now known or hereafter invented, including photocopying and recording, or in any information storage or retrieval system, without permission in writing from the publishers. Notice: Product or corporate names may be trademarks or registered trademarks, and are used only for identification and explanation without intent to infringe. Credits and acknowledgments borrowed from other sources and reproduced,with permission, in this textbook appear on appropriate page within text. ISBN : 97801318933 20 (pbk) Cover Designer: Kiwi Design Library of Congress Cataloging-in-Publication Data Gursky, Sharon The spectral tarsier/submitted by Sharon Gursky. p. cm — (Primate field series) ISBN 0-13-189332-7 1. Tarsius spectrum. I. Title. II. Series. QL737.P965G87 2007 599.8'3—dc22 2006040599 Dedicated to my Nana, Lucille Pusar, and in loving memory of my Popop, Louis Pusar This page intentionally left blank Contents List of Figures xi List of Tables xvii Acknowledgments xxi 1 Taxonomic History 1 2 Field Site and Data Collection Methods 7 Field Site 8 Geological History of Sulawesi 8 Present-day Sulawesi 8 Tangkoko Dua Saudara Nature Reserve, North Sulawesi 9 Physical Setting 9 Climate 9 Flora 9 Fauna 11 Human Population 14 Tourism 14 Methods 15 Locating Tarsier Groups 17 Mistnetting 19 Radio Tracking 20 Justification for Radio Telemetry 21 Bird-Banding 22 Habituation 23 Data Collection Methods 23 Focal Follows and Locational Data 23 Physical Measurements and Condition 25 Insect Sampling and Identification 25 Demographic Data Collection 26 vii viii CONTENTS Vegetation Analyses 32 Rainfall and Temperature 32 Moon Phase 32 Predation Experiments 33 Infant Predation Experiments 33 Adult Predator Experiments 33 Ad libitum 35 Data Analysis 35 Statistical Autocorrelation Between Data Points 35 Chapter Summary 36 3 Ecological and Social Factors Selecting for Gregariousness: Group Size and Composition, Moonphase, and Predators 37 Group Size and Composition 38 Gregariousness and Group Cohesiveness 48 Predators/Predation 56 Infants and Predators 64 Mother’s Response 67 Moonlight 72 Chapter Summary 81 4 Ecological and Social Factors Selecting for Gregariousness: Territoriality, Diet, Site Fidelity, and Dispersal Patterns 83 Territoriality 84 Description of Territorial Behavior 85 Mate Defense 90 Resource Defense 91 Diet 95 Sleeping Trees 104 Site Fidelity 107 Dispersal Patterns 108 Chapter Summary 110 5 Infant Care and the Cost of Infant Transport 111 Gestation Length 112 Litter Size 113 CONTENTS ix Birth Seasonality 113 Lactation Length and Frequency 114 Infant Transport 118 Mode of Transport: Oral vs. Fur 118 Transport Frequency 119 Parent–Offspring Conflict 122 Infant Parking 125 Infant Falling from Parked Locations 128 Infant Alarm/Distress Calls 129 Infant Environmental Exploration and Developmental Changes 130 Infant Grooming 133 Infant Mortality 135 Mother–Infant Relationships in Galagos and Lorises 138 Modeling Maternal Time Budgets: Why Park? 139 Time Budget of Adult Female Spectral Tarsiers 141 Adult Female Body Weight 142 Infant Birth Weight and Infant Growth 142 Estimating the Energetic Efficiency of Lactation (E) 146 Calculating Time Spent Foraging (Ft) for Adult Female Spectral Tarsiers 147 The Cost of Infant Transport 149 Discussion and Conclusions 151 6 The Cost of Nonmaternal Caretakers: Paternal and Subadult Infant Care 153 Introduction to Male Care 154 Form and Quantity of Paternal Care 156 Energetics of Male Care 162 Male Activity Budgets and Differences with Females 162 Male Care: Discussion and Conclusions 164 Subadult Spectral Tarsiers 165 Allomaternal Care in Primates: Hypotheses 165 Mutualism 165 Reciprocity 167 Kin Selection 167 Benefits of Philopatry 168 Allomaternal Care in Prosimian Primates 168 x CONTENTS Allomaternal Care by Subadult Female Spectral Tarsiers 171 Infant Transport 171 Food Sharing 171 Play 173 Grooming 173 Distance 175 Alarm Calls 177 A Time Budget Model for Subadult Infant Care 178 Activity Budgets of Subadults 178 Subadult Body Weight 179 Time Allocation Model 179 Subadult Care: Discussion and Conclusions 181 Why do Spectral Tarsiers Provide Allocare? 181 Could Subadults Continually Transport the Infants? 182 Chapter Summary 183 7 Conservation Status of Spectral Tarsiers 185 Introduction 185 Population Density 186 Population Viability Analysis 190 Captive Conservation 190 Habitat Destruction 191 8 Summary and Conclusions 199 References Cited 205 Index 225 List of Figures Figure 1–1 Map Illustrating the Geographic Location of the Various Tarsier Species in Indonesia, Malaysia, and the Philippines 4 Figure 2–1 Map Illustrating Position of Tangkoko Dua Saudara Nature Reserve on the Northeastern-Most Tip of the Island of Sulawesi, Indonesia 10 Figure 2–2 The Rainfall (mm) Collected Each Month Between 1994 and 2004 11 Figure 3–1a The Variation in Group Sizes During 1994–1995 Survey 38 Figure 3–1b The Variation in Group Sizes During 1999 Survey 39 Figure 3–1c The Variation in Group Sizes During 2003–2004 Survey 39 Figure 3–2 The Relative Limb Proportions of Male and Female Spectral Tarsiers Expressed as Mean Indices 45 Figure 3–3 The Frequency Distribution of the Observed Distance Between Adults for Each Simultaneous Locational Data Point 50 Figure 3–4 The Frequency that Two Adult Spectral Tarsiers Encountered One Another According to the Hour of the Night 51 Figure 3–5a The Expected Encounter Rates for Group M300, if the Group Members’ Movements Were Random and Independent, Z, Were Compared with the Observed Encounter Rates for Each Inter-individual Distance 52 xi xii LIST OF FIGURES Figure 3–5b The Expected