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Information to Users INFORMATION TO USERS While the most advanced technology has been used to photograph and reproduce this manuscript, the quality of the reproduction is heavily dependent upon the quality of the material submitted. For example: # Manuscript pages may have indistinct print. In such cases, the best available copy has been filmed. ® Manuscripts may not always be complete. In such cases, a note will indicate that it is not possible to obtain missing pages. • Copyrighted material may have been removed from the manuscript. In such cases, a note will indicate the deletion. Oversize materials (e.g., maps, drawings, and charts) are photographed by sectioning the original, beginning at the upper lefr-hand comer and continuing from left to right in equal sections with small overlaps. Each oversize page is also filmed as one exposure and is available, for an additional charge, as a standard 35mm slide or as a 17”x 23” black and white photographic print. Most photographs reproduce acceptably on positive microfilm or microfiche but lack the clarity on xerographic copies made from the microfilm. For an additional charge, 35mm slides of 6”x 9” black and white photographic prints are available for any photographs or illustrations that cannot be reproduced satisfactorily by xerography. Farslow, Daniel Leslie THE BEHAVIOR AND ECOLOGY OF THE LONG-TAILED MACAQUE (MACACA FASCICULARIS) ON ANGAUR ISLAND, PALAU, MICRONESIA The Ohio State University Ph.D. 1987 University Microfilms I n ter n ât i O n siSOO N. Zeeb Road, Ann Arbor. Ml 48106 PLEASE NOTE; In all cases this material has been filmed in the best possible way from the available copy. Problems encountered with this document have been identified here with a check mark ■/ 1. Glossy photographs or pages_____ 2. Colored illustrations, paper or print______ 3. Photographs with dark background_____ 4. Illustrations are poor copy______ 5. Pages with black marks, not original copy______ 6. Print shows through as there is text on both sides of pag_______ e 7. Indistinct, broken or small print on several pages 8. Print exceeds margin requirements______ 9. Tightly bound copy with print lost in spine_______ 10. Computer printout pages with indistinct print______ 11. Page(s)___________ lacking when material received, and not available from school or author. 12. Page(s)___________ seem to be missing in numbering only as text follows. 13. Two pages num bered . Text follows. 14. Curling and wrinkled pages______ 15. Dissertation contains pages with print at a slant, filmed a s received_________ 16. Other______________________________________________________________________ University Microfilms International THE BEHAVIOR AND ECOLOGY OF IHE LCWG-TAILED MACAQUE (MACACA FASCICULARIS) ON ANGAUR ISLAND, PALAU, MICRONESIA DISSERTATION Presented in partial fulfillment of the requirements for the degree of Doctor of Philoscçhy in the Graduate Division of The C M o State University By Daniel L. Farslow, B.A. (Hons), M.A., 1977 The Ohio State University 1987 Dissertation Conraittee: improved By Frank E. Poirier Paul W. Sciulli Jean M. Brainard Adviser Department of Anthropology lb Ripu Daman Singh ACKNOWLEDGMENTS Reseéirch of this kind cannot be acccnplished without the trust and help of many people. Dr. F. E. Poirier was constantly supportive in both the research and preparation of this manuscript and was instrumental in ïty obtaining financial assistance through departmental teaching assistantships throughout ny time at Ihe Chio State University. I express sincere appreciation to Dr. Poirier for his enthusiasm and faith in me and the project. My thanks and fond memories go out to the late Dr. D. T. Hughes ^lAo kindly gave his support as Department Chairman and introduced me to many of his good friends in Palau and on Angaur Island. Thanks go to the other members of my advisory committee Dr. Paul W. Sciulli, vto was alv&ys there vhen I needed help, and to Dr. Jean M. Brainard, for all their support and encouragement. Everyone in Palau was so helpful it is impossible to name each person, but specific thanks must go to: Sakie Morris, Katherine Kesolei, Andres and Vittorio Uherbelau, Karen Nero, Serge Birk, Mona, Dennis, Kim and Rosie Batcheller, and last but not least. Bob Etchell, vrtiose generosity, inspiration and friendship set a standard v^ich I could never attain. The Magistrate of Angaur, James Robert graciously allowed me to work on the island and he and his wife Mako lent unending support and friendship, as did the entire Moses family and the Massao's. Nick Takarai and later Balerio Pedro provided housing, and Balerio was always there to guide me through my many times of loneliness and confusion. Special mention must go to Ihomas, Alphonse and Rufino Moses, ny brothers, and to Patrick, Leone and Kasiano. To them and to all the citizens of Angaur I owe ray deepest appreciation and thanks. I wish I could have done more to earn their friendship, Angaur will remain dear to iry heart forever. When the research is complete the work has just begun. Ify thanks to Karen Itoxell v*o labored long and hard under sometimes adverse conditions to expertly type this manuscript. Her patience and tolerance throughout the many revisions are greatly appreciated. A special mention must go to Or. Kerry Stroup %Ao not only suffered through the editing of each and every word of this thesis, but also had to endure ny incessant expression of self-doubt throughout the process. Thank you Kerry for your special skills and understanding. And finally, there is not enough appreciaticm to give to Nancy Kirwin, *Aio joined me in the field and shared the joy and hardship of researcdi. Without her caring energy I would have been much less able to cope with the rigors of science. To Nancy, ny unending thanks. August 2, 1945 ................... B o m - Windsor, Ontario, 1975 ......... ..................B.A., (Honors), University of Windsor, Windsor, Ontario, Canada 1976-1982.........................Graduate Associate, Department of Anthrcçology, Ihe Ohio State University, Columbus, CSiio 1977 ............................ M.A., Ihe Chio State University, Columbus, Chio 1982-1984.........................Lecturer, Erindale College, Ihe University of Toronto Mississauga, Ontario, Canada 1984-Present..................... Graduate Associate, Department of Anthropology, Ihe Ohio State University, Columbus, CSiio PUBLICATimS Poirier, F. E. and D. L. Farslow 1984 Status of the crab-eating macaque on Angaur Island, Palau, Micronesia. lUCN/SSC Primate Specialist Grotç) Newsletter 4:42-43. Sigrom, B. A. and D. L. Farslow 1986 Ihe hindlimb of primates. In: Comparative Primate Biology, Volume I: Systematics, Evolution and Anatoiy, D. R. Swindler, ed.. New York: A. R. Liss. FIELDS OF STUDY Major Field: Physical Anthrc^logy Minor Fields: Primate Behavior and Ecology Human Paleoanthrtçology TABLE OF CONTENTS Page .............................. ...... .iii VITA.......................................................... V LIST OF TABLES........................ ix LIST OF FIGURES..............................................xi CHAPIER I. INTRODUCTION........................................... 1 Introduction............................................1 Conservation............................................2 Socioecology and Island Biology..........................4 Aims................................................... 7 Macaques, Background and Distribution................... 8 The Dong-Tailed Macaque (M. fascicularis) .............. 10 Distribution............................................. 11 Research Background .................................... 16 II. THE STUDY AREA - ANGAUR ISLAND. ...................... 21 Introduction............................................. 21 Geography and Geology................................... 26 Vegetation...............................................32 Climate.................................................38 Human Bihabitants ..................................... 45 The Angaur Macaca fascicularis............................49 Other Fauna............................................. 51 Research Strategy....................................... 54 III. DEMOGRAPHY.............................................. 57 Introduction.............................................57 Methods.................................................58 Population and Distribution.......................... .62 Reproduction.............................................67 Morbidity...............................................68 Mortality...............................................73 Discussion...............................................78 IV. SOCIAL ORGANIZAnCM................................... 86 Introduction............................................86 Methods............................................... 88 Gnxp Ocn%x)sition .............................. 91 Group Dynamics.........................................96 Discuss ic^...............................................101 V. SOCIAL BEHAVIOR......................................... 108 Introduction............................ 108 Methods.................................................109 Ocnmunication Matrix..................................... 116 Grooming.................................................130 Dominance............................................... 135 Play.................................................... 139 Sex-Consort Relationships................................143 Vigilance............... 145 Discuss icxi............................................... 148 VI.
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