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Techniques for the Study of Primate Population Ecology Subcommittee on Conservation of Natural Populations Committee on Nonhuman Primates Division of Biological Sciences Assembly of Life Sciences National Research Council \w NATIONAL ACADEMY PRESS r; ry f, nV Washington, D.C. 1981
8 )-0 NOTICE: The project that is the subject of this report was approved by the Governing Board of the National Research Council, whose members are drawn from the Councils of the National Academy of Sciences, the National Academy of Engineering, and the Institute of Medicine. The members of the committee responsible for the report were chosen for their special compe- tences and with regard for appropriate balance. This report has been reviewed by a group other than the authors according to procedures approved by a Report Review Committee consisting of members of the National Academy of Sciences, the National Academy of Engineering, and the Institute of Medicine. The National Research Council was established by the National Academy of Sciences in 1916 to associate the broad community of science and technology with the Academy's purposes of furthering knowledge and of advising the federal government. The Council operates in accor- dance with general policies determined by the Academy under the authority of its congres- sional charter of 1863, which establishes the Academy as a private, nonprofit, self-governing membership corporation. The Council has become the principal operating agency of both the National Academy of Sciences and the National Academy of Engineering in the conduct of their services to the government, the public, and the scientific and engineering communities. It is administered jointly by both Academies and the Institute of Medicine. The National Academy of Engineering and the Institute of Medicine were established in 1964 and 1970, re- spectively, under the charter of the National Academy of Sciences. The primary support for development of this report was provided by Contract N01-RS-9-2117 with the Division of Research Services, National Institutes of Health. Additional support was provided by Grant RC-1W from the American Cancer Society, Inc.; Contract APHIS 53- 3294-9-2 with the U.S. Department of Agriculture; Contract DNA001-79-C-0087-P00001 with the Defense Nuclear Agency; Contract DE-AM02-76CH93000 (Task Agreement DE-AT02- 76CH93011) with the Department of Energy; Contract N01-RR-9-2109 with the Division of Research Resources, Animal Resources Branch, National Institutes of Health; Grant PCM- 7921163 with the National Science Foundation; Contract N00014-80-C-0162 with the Office of Naval Research; and contributions from pharmaceutical companies and other industry. Library of Congress Cataloging In PubUcation Data Main entry under title: Techniques for the study of primate population ecology. Bibliography: p. 1. PrimatesâEcology. 2. Population biologyâTechnique. 3. MammalsâEcology. 4. Mammal populations. 1. National Research Council (U.S.) Committee on Nonhuman Primates. Subcommittee on Conservation of Natural Populations. QL737.P9T43 599.8'045 81-11345 ISBN 0-309-03179-6 AACR2 Available from NATIONAL ACADEMY PRESS 2101 Constitution Avenue, N.W. Washington, D.C. 20418 Printed in the United States of America
COMMITTEE ON NONHUMAN PRIMATES NORMAN H. ALTMAN, Papanicolaou Cancer Research Institute, Miami, Florida (Chairman) BENJAMIN G. BRACKETT, Department of Clinical Studies at New Bolton Center, University of Pennsylvania, School of Veteri- nary Medicine, Kennett Square, Pennsylvania ROBERT w. GOY, Wisconsin Regional Primate Research Center, Madison, Wisconsin BERNADETTE M. MARRIOTT, Division of Comparative Medicine, Johns Hopkins University School of Medicine, Baltimore, Maryland ALBERT E. NEW, National Cancer Institute, National Institutes of Health, Bethesda, Maryland JOHN w. SENNER, Oregon Regional Primate Research Center, Beaverton, Oregon Subcommittee on Conservation of Natural Populations JOHN F. EISENBERG (Chairman), National Zoological Park, Smithsonian Institution, Washington, D.C. WOLFGANG p. j. DITTOS, Smithsonian Institution, Ceylon Primate Survey, Kandy, Sri Lanka THEODORE H. FLEMING, University of Miami, Coral Gables, Florida KENNETH GREEN, Howard University, Washington, D.C. THOMAS STRUHSAKER, New York Zoological Society, The Rocke- feller University, New York, and Makerere University, Kam- pala, Uganda RICHARD w. THORINGTON, JR., U.S. National Museum of Natural History, Smithsonian Institution, Washington, D.C. Staff Officers NANCY A. MUCKENHIRN, 1979-1980 EARL W. GROGAN, 1980-1981 111
Preface The need for basic data on population densities and trends for several species of animals has been increasing steadily as wild lands have been under pressure for the extraction of mineral, tim- ber, and animal resources. The development of land for agricul- tural and industrial purposes has generally progressed without any regional planning and has resulted in the degradation or loss of native habitats, especially mature forests, and the fauna and flora associated with these habitats. In some areas the remaining tracts of wild land are becoming so restricted that management of animal populations has become necessary to safeguard their fu- tures and to bring utilization of some of the species into line with the capacity of the habitat to support them. The assessment of the status of wild primate populations re- quires immediate attention if primates are to be protected and managed as renewable wildlife resources. Little is being done to assemble the needed information systematically. Export restric- tions imposed on primates over the past decade are in part a re- sponse to a concern for conserving wild populations. They are
vi Preface likely to remain in effect until scientific data become available that will justify changing them. In the early 1970's a committee organized by the Institute of Laboratory Animal Resources, of the Division of Biological Sci- ences, Assembly of Life Sciences, National Research Council, was asked to study the status of wild primate populations. This was the Committee on Nonhuman Primates. Under the leadership of R. W. Thorington, Jr., Chairman, and C. H. Southwick, Vice-Chairman, the committee set in motion a program of surveys, or censuses, to assemble data on primate populations in Colombia, Peru, Guyana, and Bolivia. Funds for supporting survey teams were provided by the Pan American Health Organization, the National Institutes of Health, and the U.S. Army Medical Research and Development Command. When these surveys were compared with studies conducted by other investigators, it was obvious that lack of standardized pro- cedures was causing serious inconsistencies in findings and that a manual would contribute significantly to accuracy and uniformity. The Subcommittee on Conservation of Natural Populations, Committee on Nonhuman Primates, was organized to produce such a manual, and the present manual is the result of the sub- committee's efforts. It is hoped that the techniques and observa- tions presented here will lead to increased understanding of forest ecosystems and will encourage worldwide efforts to develop con- servation and management programs. JOHN F. EISENBERG, Chairman Subcommittee on Conservation of Natural Populations
Acknowledgments In addition to the basic contracts supporting this work, the Na- tional Institutes of Health and other institutions supported prep- aration of certain parts of the report. The institutions that pro- vided such support, the persons who received it, and the subjects dealt with in these persons' contributions to the report are as fol- lows: National Zoological Park, John F. Eisenberg, habitat de- scription; Smithsonian Institution, National Science Foundation, and National Institutes of Health, Richard W. Thorington, Jr., habitat use and collecting and marking techniques; Smithsonian Institution, National Science Foundation, National Institutes of Health, and the Deutsche Forschungs-Gemeinschaft, Wolfgang P. J. Dittus, sexing and aging animals, population analysis, and determinants of population density and growth; National Insti- tutes of Health, Howard University, Johns Hopkins University, and National Zoological Park, Kenneth Green, remote-sensing techniques (Dr. Green also prepared a draft of a censusing man- ual and a background discussion of censusing methods.); and New York Zoological Society, National Science Foundation, and Na- vii
viii Acknowledgments tional Institute of Mental Health, Thomas Struhsaker, censusing methods. The Subcommittee on Conservation of Natural Populations is indebted to several persons who made suggestions concerning preparation of various sections of the manuscript. Special thanks are due Anne Baker-Dittus, T. Butynski, Lysa Lelancl. G. Caughley, J. Ballon, J. Frazier, and J. Russell. Artwork was pre- pared by Sigrid James-Briich and Figure 1-1 by Carrie Thoring- ton.
Contents 1 INTRODUCTION 2 SITE SELECTION Maps and Photographs, 5 Faunal Lists and Floral Inventories, 6 Special Studies of Indicator Species, 7 HABITAT DESCRIPTION AND SPECIMEN COLLECTION 10 Habitat Profile, 11 Characteristics of Habitats, 13 Species Maps, 23 Recording Climatic Data, 25 Collecting and Marking Specimens, 26 Phenological Patterns, 31 Satellite Imagery, 33 ix
Contents CENSUS METHODS FOR ESTIMATING DENSITIES 36 Background for Calculations Derived from Transect Censuses, 36 General Guidelines and Field Procedures, 45 Assessing the Impact of Human Activities, 48 Broad Surveys, 49 Applications of Line Transect Censuses in an African Forest, 50 Nonlinear Density Plot Method, 65 Quadrat Censuses, 68 Specialized Census Methods, 71 Long-term Monitoring of Specific Groups, 74 Summary of Important Census Methods for Diurnal Primates, 78 Extrapolation of Density Estimates from Census Area to Other Areas, 79 TECHNIQUES FOR SEXING AND AGING PRIMATES 81 Sexing, 83 Estimating Relative Age Classes for Censuses, 84 Direct Measurement of Chronological Age, 88 Indirect Measurement of Chronological Age, 90 Identifying Individual Primates by Natural Markings, 110 Identifying Individual Primates by Artificial Marking Techniques, 117 HABITAT USE 128 Time Budgets, 128 Use of Space, 130 Feeding Ecology, 132
Contents xi 7 PRIMATE POPULATION ANALYSIS 135 Data Requirements, 136 Births and Fecundity, 136 Estimating Rates of Population Growth, 145 Estimating Vital Statistics from a Stable Standing Age Distribution, 147 Estimating Vital Statistics from Individual Life-History Records, 163 Estimating Vital Statistics from Standing Age Distributions Sampled at Yearly Intervals, 164 Deriving Vital Statistics from Fecundity and Survivorship Schedules, 165 Dispersal, 169 Suggested Readings, 175 8 DETERMINANTS OF POPULATION DENSITY AND GROWTH 176 Measures of Population Size in Relation to the Environment, 176 Determinants of Population Size, 181 Food Supply as a Major Factor Determining Population Size, 185 Population Regulation by Behavioral and Physiological Mechanisms, 189 9 COMMENTARY 197 REFERENCES 205 APPENDIXES A LANDSAT Data Distribution Centers Operative in 1979, 221 B Sources of Interpretive Service for Satellite Imagery in the United States, 223 C Sample Data Sheets, 226 D Checklist of Essential Equipment, 233
