ISHOKU DOUGEN – THE MEDICINAL USE OF PLANTS AND CLAYS BY WILD JAPANESE MACAQUES (MACACA FUSCATA YAKUI) by CHRISTOPHER DAGG (Under the Direction of Carolyn Ehardt) ABSTRACT There is increasing evidence from behavioral and chemical ecology that non-nutritional resources have a significant effect on the health of wild animal populations, both through deliberate self-medication (zoopharmacognosy) and dietary prophylaxis. This dissertation reports the first systematic investigation of anti-parasitic self-medication in Yakushima Japanese macaques (Macaca fuscata yakui). Adult female macaques of NA2 group on Yakushima were studied for one full seasonal cycle, their health measured through fecal parasitological analysis, and their foraging behavior recorded through focal observation. A set of candidate medicinal plants and clay were selected from literature and preliminary observations, and interactions with these recorded in continuous detail. Statistical analyses were designed to reveal patterns in parasite load, parasitological stimuli for self-medicative behavior and the functional efficacy of those behaviors. Ethnographic interviews were conducted with several practitioners of traditional medicine in Japan, and their knowledge compared with the macaque results. The macaques were found to be infected with five species of gastro-intestinal parasite, but appear to tolerate the infections with few symptoms. The coarse grass Miscanthus sinensis was found to be associated with severe Streptopharagus pigmentatus infection, and may act to physically expel this nematode taxon. Geophagy (clay eating) is practiced year-round, but while it may account for the low incidence of diarrhea, it was not found to be directly associated with parasite infection. Instead it appears to function in the detoxification of invertebrate and plant foods. Records of juvenile observation of their parents suggest that these two behaviors may be a cultural tradition in the population. The ingestion of several other plants seems to incorporate a range of pharmacologically active phytochemicals into the macaque diet. While some are rare and deserve further investigation, others are utilized as food items and any health benefit accrued incidentally to this. Interview data reveal that the traditional pharmacopeia of Japan differs significantly from that available to the macaques, and is proposed to be due to ecological differences between Yakushima and the centers of traditional medicine in Japan. The significance of these results is discussed with regard to conservation medicine, and theories of the emergence of human medicine. INDEX WORDS: Zoopharmacognosy, macaque, Macaca fuscata, Yakushima, geophagy, parasitology, Japan, ethnobotany, conservation medicine, chemical ecology ISHOKU DOUGEN – THE MEDICINAL USE OF PLANTS AND CLAYS BY WILD JAPANESE MACAQUES (MACACA FUSCATA YAKUI) By CHRISTOPHER DAGG B.Sc., University of Edinburgh, Scotland, 1997 A Dissertation Submitted to the Graduate Faculty of The University of Georgia in Partial Fulfillment of the Requirements for the Degree DOCTOR OF PHILOSOPHY ATHENS, GEORGIA 2009 © 2009 Christopher Dagg All Rights Reserved ISHOKU DOUGEN – THE MEDICINAL USE OF PLANTS AND CLAYS BY WILD JAPANESE MACAQUES (MACACA FUSCATA YAKUI) By CHRISTOPHER DAGG Major Professor: Carolyn Ehardt Advisory Committee: René Bobe Susan Tanner Electronic Version Approved: Maureen Grasso Dean of the Graduate School The University of Georgia May 2009 DEDICATION Mook! iv ACKNOWLEDGEMENTS I owe the completion of this dissertation to Angela Hong, without whom I may have never returned. Her support, understanding, love and timely application of internet telephony kept me sane though many a long period in the field, and when faced with the seemingly insurmountable challenges of data analysis and writing. Perhaps the most long-overdue professional acknowledgement is deserved by Micheal Huffman of Kyoto University, for the encouragement, guidance and constant enthusiasm he provided over the past decade. From my first interest in zoopharmacognosy as an undergraduate to this day, he has consistently gone out of his way to support the process of enquiry, my entry to graduate school, the logistics of research and residence in Japan, and the particular issues of writing up. This is all the more remarkable in that he has never been an official part of my advisory committee, and all his help has truly been above and beyond the call of duty. I am privileged to know him as both colleague and friend. Heartfelt thanks are also deserved by Laurent Tarnaud, Alexander Hernandez and Alisa Chalmers of Kyoto University for their invaluable support and companionship throughout and following my time in Japan, Shin-ichiro Aiba of Kagoshima University for the kind donation of his climatological data ,Professor Hasegawa of Oita University for sharing his parasitological expertise, Sachiko Hayakawa for the kind donation of the fecal sample featured in chapter 3, and the Yakushima Forest Environment Conservation Center, Kagoshima Prefecture and Kirishima-Yaku National Park for permission to conduct research in the conservation area. I am also sincerely grateful for the cooperation and time of all the traditional pharmacists, herbalists and wholesalers that kindly donated their time to be interviewed as part of my preliminary research in 2003 and during my ethnomedicinal research of 2007, for the work of my translator, Ueda, without whom I would have been forced to impose my terrible Japanese on unsuspecting informants. v I would like to extend my gratitude to all the researchers and students at the Kyoto University Yakushima Field Research Station for their tolerant support and company throughout this field session: Professor Suguira, Professor Yamagiwa, Professor Hanya, Mari Terakawa, Koda Ryosuke, Naoki and Yoshimi Agetsuma, Miki Matsubara. I must also acknowledge the fortunate acquaintance and kindness of Yuko Obata, who found me both my houses on Yakushima. Finally, and certainly not least, I would like to express my gratitude to the professors who have served on my Advisory Committee: my Major Professor, Carolyn Ehardt, without whom I would not have come to UGA, for her endless patience and support, Brent Berlin for being tolerant of my crazy ideas, Irwin Bernstein for his clarity and rigor of mind, Susan Tanner and Rene Bobe for stepping up to the crease, and also to Alex Brewis who regrettably moved away. Sincere thanks also for the invaluable support of Charlotte Blume, Margie Floyd and LaBau Bryan for keeping me right through the maze of academic and immigration beaurocracy. This study was funded in part by National Science Foundation, Physical Anthropology Dissertation Improvement Award #BCS-0525156 (2006-7 field season) and preliminary research by NSF Ethnographic Research Training Grant #BCS-0091764 (summer 2003). vi TABLE OF CONTENTS Page ACKNOWLEDGEMENTS .......................................................................................................................... v LIST OF TABLES ........................................................................................................................................ x LIST OF FIGURES ..................................................................................................................................... xi CHAPTERS 1. INTRODUCTION AND LITERATURE REVIEW .......................................................................... 1 Introduction .............................................................................................................................. 1 Literature Review .................................................................................................................... 9 Conclusions............................................................................................................................ 33 Literature Cited ...................................................................................................................... 34 2: THE PARASITE ECOLOGY OF JAPANESE MACAQUES (MACACA FUSCATA YAKUI) ON YAKUSHIMA ............................................................................................................. 48 Abstract .................................................................................................................................. 49 Introduction ............................................................................................................................ 50 Methods ................................................................................................................................. 53 Results ................................................................................................................................... 58 Discussion .............................................................................................................................. 67 Conclusions............................................................................................................................ 74 Literature Cited ...................................................................................................................... 75 3: THE INGESTION OF MISCANTHUS SINENSIS AS AN ANTIPARASITIC ADAPTATION IN YAKUSHIMA JAPANESE MACAQUES (MACACA FUSCATA YAKUI) [WITH SATCHIKO HAYAKAWA] ................................................................................. 81 Abstract .................................................................................................................................