ALL APES GREAT AND SMALL VOLUME 1: AFRICAN APES Developments in Primatology: Progress and Prospects Series Editor: Russell H. Tuttle, University of Chicago, Chicago, Illinois This peer-reviewed book series will meld the facts of organic diversity with the continuity of the evolutionary process. The volumes in this series will exemplify the diversity of theoretical perspectives and methodological approaches currently employed by primatolo- gists and physical anthropologists. Specific coverage includes: primate behavior in natural habitats and captive settings; primate ecology and conservation; functional morphology and developmental biology of primates; primate systematics; genetic and phenotypic differ- ences among living primates; and paleoprimatology. All Apes Great and Small Volume 1: African Apes Co-edited by Biruté M. F. Galdikas, Nancy Erickson Briggs, Lori K. Sheeran, Gary L. Shapiro, and Jane Goodall ALL APES GREAT AND SMALL VOLUME 1: AFRICAN APES Co-Edited by Biruté M. F. Galdikas President Orangutan Foundation International Los Angeles, California Nancy Erickson Briggs California State University at Long Beach Long Beach, California Lori K. Sheeran California State University at Fullerton Fullerton, California Gary L. Shapiro Vice-President Orangutan Foundation International Los Angeles, California and Jane Goodall President The Jane Goodall Institute KLUWER ACADEMIC PUBLISHERS NEW YORK, BOSTON, DORDRECHT, LONDON, MOSCOW eBook ISBN: 0-306-4761-1 Print ISBN: 0-306-46757-7 ©2002 Kluwer Academic Publishers New York, Boston, Dordrecht, London, Moscow Print ©2001 Kluwer Academic/Plenum Publishers New York All rights reserved No part of this eBook may be reproduced or transmitted in any form or by any means, electronic, mechanical, recording, or otherwise, without written consent from the Publisher Created in the United States of America Visit Kluwer Online at: http://kluweronline.com and Kluwer's eBookstore at: http://ebooks.kluweronline.com Contents Dedication ix Acknowledgments xi Preface xiii Volume Overview xv SECTION ONE – ISSUES IN APE AND HUMAN EVOLUTION Introduction to Section One 1 1. AFRICAN APES AS TIME MACHINES 5 R. Wrangham and D. Pilbeam 2. PRIMATE DIVERGENCE TIMES 19 A. Janke and U. Arnason 3. THE CEREBELLUM: AN ASSET TO HOMINOID COGNITION 35 C.E. MacLeod, K. Zilles, A. Schleicher, and K.R. Gibson SECTION TWO – BONOBOS, THE “FORGOTTEN APE”? Introduction to Section Two 55 4. THE STATUS OF THE BONOBO (PAN PANISCUS) IN THE DEMOCRATIC REPUBLIC OF CONGO 57 J. Dupain and L. Van Elsacker v vi Contents 5. THE STATUS OF BONOBOS IN THEIR SOUTHERNMOST GEOGRAPHIC RANGE 75 J.A. Myers Thompson 6. CURRENT SITUATION OF BONOBOS IN THE LUO RESERVE, EQUATEUR, DEMOCRATIC REPUBLIC OF CONGO 83 C. Hashimoto and T. Furuichi SECTION THREE – CHIMPANZEES, THE BEST-KNOWN APE Introduction to Section Three 91 7 PAN IN PANDEMONIUM 95 S.T. Boysen and T. Butynski 8. PREDATION OF MAMMALS BY THE CHIMPANZEES OF THE MAHALE MOUNTAINS, TANZANIA 107 K. Hosaka, T. Nishida, M. Hamai, A. Matsumoto-Oda, and S. Uehara 9. REPRESENTATIONAL CAPACITIES IN CHIMPANZEES: NUMERICAL AND SPATIAL R EASONING 131 S.T. Boysen and V.A. Kuhlmeier SECTION FOUR – GORILLAS, THE GREATEST OF THE APES Introduction to Section Four 149 10. THE STATUS OF GORILLAS WORLDWIDE 151 F.G.P. Patterson and M.L. Matevia 11. TWENTY-SEVEN YEARS OF PROJECT KOKO AND MICHAEL 165 F.G.P. Patterson and W. Gordon 12. WHO’S IN CHARGE? OBSERVATIONS OF SOCIAL BEHAVIOR IN A CAPTIVE GROUP OF WESTERN LOWLAND GORILLAS 177 T.A. Stein SECTION FIVE – PHYSIOLOGICAL BASES FOR BEHAVIOR AND AGING: GREAT APES AND HUMANS Introduction to Section Five 191 Contents vii 13. THE GREAT APE AGING PROJECT: A RESOURCE FOR COMPARATIVE STUDY OF BEHAVIOR, COGNITION, HEALTH, AND N EUROBIOLOGY 195 J.M. Erwin, M. Bloomsmith, S.T. Boysen, P.R. Hof, R. Holloway, L. Lowenstine, R. McManamon, D.P. Perl, W. Young, and A. Zihlman 14. AN INTERNATIONAL DATABASE FOR THE STUDY OF DIABETES, OBESITY, AND AGING IN GREAT APES AND OTHER NONHUMAN PRIMATES 201 R. Martens, R. Couch, B. Hansen, C. Howard, J. Kemnitz, D.P. Perl, and J.M. Erwin 15. STUDIES OF AGE-RELATED NEURONAL PATHOLOGY IN GREAT APES 207 D.P. Perl, P.R. Hof, E.A. Nimchinsky, and J.M. Erwin 16. METABOLITES OF OVARIAN HORMONES AND BEHAVIORAL CORRELATES IN CAPTIVE FEMALE BONOBOS (PAN PANISCUS) 217 M.H. Jurke, L.R. Hagey, N.M. Czekala, and N.C. Harvey 17. SEXUAL MOTIVATION OF MALE CHIMPANZEES DURING THE FEMALE CYCLE, INCLUDING P RELIMINARY DATA ON AGE EFFECTS 231 R.D. Nadler SECTION SIX – THE BUSHMEAT CRISIS: AFRICAN APES AT RISK Introduction to Section Six 237 18. BUSHMEAT, PRIMATE KINSHIP, AND THE GLOBAL CONSERVATION MOVEMENT 241 A.L. Rose 19. STATUS OF THE PROPOSED LOMAKO FOREST BONOBO RESERVE: ACASE STUDY OF THE BUSHMEAT TRADE 259 J. Dupain and L. Van Elsacker 20. WHAT HAPPENED TO GORILLA GORILLA UELLENSIS? A PRELIMINARY INVESTIGATION 275 K. Ammann and N. Briggs 21. APES, PERSONS, AND BIOETHICS 283 P. Singer and P. Cavalieri viii Contents Index 293 Dedication To the co-editors’ families and friends for their support throughout this challenging project. With appreciation for the loving support of our parents: Pat and Bill Chiles, Otto Erickson, and Filomena Galdikas. To the late Antanas Galdikas, whose love of animals resonated in his children. To Vanne Goodall, the late beloved mother of Jane Goodall. To Lorraine P. Jenkins, the late beloved mother of Dr. Gary and Mrs. Ingrianni Shapiro. Her generosity provides for OFI’s annual L.P. Jenkins Memorial Fellowship. We intend for this volume to promote the dignity and well-being of all Africans, whether ape or human. ix Acknowledgments Many of the papers in this volume were first presented at the Third International Great Apes of the World Conference (GAWC-3) held July 3-6, 1998 in Kuching, Sarawak, Malaysia. We thank the Chief Minister of the state of Sarawak, Datuk Taib, Dr. Laila Taib, and Dr. James Masing for their support, without which GAWC-3 would not have been successful. We also thank the participants, who came from all over the world to attend the first conference on the apes held in Malaysia: Altheide, T.; Ammann, K.; Ancrenaz, M.; Andau, M.; Andayani, N.; Bates, M.; Beck, R.; Beckenbach, A.T.; Bennett, E.; Bernard, H.; Blair, S.; Blom, A.; Bloomsmith, M.; Blouch, R.; Boestani, A.N.; Bosi, E.; Boysen, S.T.; Briggs, N.; Butynski, T.; Cocks, L.; Collier, J.; Couch, R.; Czekala, N.M.; de la Rosa, M.; de Vries, H.; Didier, S.; Djojasmoro, R.; Dulek, L.; Dupain, J.; Ely, J.; Erwin, J.; Foitova, I.; Galdikas, B.M.F.; Ghaffar, N.; Gilbertson, W.; Gilmour, L.; Griffiths, M.; Hagey, L.R.; Hansen, B.; Harvey, N.C.; Hashim, N.R.; Hashimoto, C.; Hill, K.; Hof, P.; Holloway, R.; Hoole, W.S.; Howard, C.; Jamart, A.; Janecek, J.; Janke, A., Jerabkova, Z.; Jurke, M.; Jurke, S.; Kanthaswamy, S.; Karesh, W.; Kemnitz, J.; Kilbourn, A.; Komiji, J.; Lackman-Acrenaz, I.; Lardeux- Gilloux, I.; Lee, D.R.; Leiman, A.; Lowenstine, L.; MacKinnon, J.; MacKinnon, K., Martelli, P., Martens, R.; McManamon, R.; Melnick, D.J.; Minesi, C.; Mitrasetia, T.; Moore, B.; Mootnick, A.; Morales, J.C.; Mosman, K.; Muir, C.; Mutalib, A.; Nadler, R.; Nessel, J.; Nimchinsky, E.; Norcup, S.; Ollanski, M.; Onuma, M.; Paredes, G.; Patterson, F.; Perl, D.; Poullet, S.; Redmond, I.; Robertson, Y.; Rose, A.L.; Shapiro, G.; Sheeran, L.K.; Sinaga, D.W.; Singer, P.; Singleton, I.; Skrivankova, J.; Stein, T.; Stevens-Wood, B.; Stone, W.H.; Supriatna, J.; Swartz, K.B.; Tambing, E.; Tanaka, M.; Thompson, J.; Tisen, O.B.; Todd, A.; Tutin, C.; Utami, S.S.; Vancata, V.; Vancatova, M.; Van Elsacker, L.; van Hoof, J.A.R.A.M.; van Schaik, C.; xi xii Acknowledgments Verheyen, R.; Wisdom, L.; Wolfe, N; Young, W.; Zihlman, A.; and Zlamalova, H. We thank the Sarawak Development Institute (SDI), particularly Lelia Sim Ah Hua, Zabariah Matali, Tang Hung Huong, and Daniel Chew, for co- organizing GAWC-3 with the Orangutan Foundation International. The SDI staff was professional, competent, and efficient during the planning stages of and at the conference itself. We also thank the staff and management of the Crowne Plaza Riverside Hotel, particularly General Manager Mohd. Khalid Buang, for hosting the event and providing excellent service to all concerned. We thank the following agencies and organizations for sponsoring GAWC-3: Ministry of Tourism, Sarawak; Sarawak Tourism Board; Malaysia Airlines System; Malaysia Tourism Promotion Board; Orangutan Foundation International; Orangutan Foundation UK; Safeskin Corp. SDN BHD; Datuk Amar (Dr.) Leonard Linggi Jugah; Eric Raymond Foundation; Mr. Ralph Arbus; Sarawak Shell BHD; Sedgewick County Zoo; Zoo Atlanta; and Nature’s Hammock. The following agencies and individuals also provided valuable support in making the conference possible: National Park and Wildlife Department Sarawak; Police Department Sarawak; Immigration Department Sarawak; Chief Minister’s Department; Kuching City North Council; Aloysious Dris; Mohd. Tuah Jais; Robert Basiuk; Sharifa Danial; K.H. Chea; Tan Hyn Teong; Ahmad Fuaad Dahlan; Ismail Hj.Talib; Abu Omar; Tony Champion; Rusty Staff; Marilyn Staff; Karin Harvey; Simone Archambeau; Michael Hawkins; and Philip Yong. Thanks also goes to the following OFI members who contributed to the conference’s success: Rodney Briggs, Inggriani Hartanto Shapiro, Ashley Leiman, Pamela Breitbeil, Gloria Anderson, Filomena Galdikas, Aldona Galdikas, John Pearson, Tony Rose, Lisa Mather, Wendy Hoole, Dandy O’Shea, Chantel Bryan, and Dave Duncan. We are grateful for the institutional support provided by Simon Fraser University (BMFG), California State University-Fullerton (LKS), and California State University-Long Beach (NB) and for our departmental colleagues’ encouragement. Dr. Douglas Eernisse (Department of Biology, California State University-Fullerton) provided helpful commentary on chapter two. We appreciate the patience and assistance of the Kluwer staff, particularly Andrea Macaluso, Deborah Doherty, Susanne St. Claire, and Mary McCarra. We express our gratitude to Marie Marley for her tireless work formatting chapters, arranging layouts, and entering our editorial changes.