PROCEEDINGS OF THE CENTER FOR TROPICAL FOREST SCIENCE – ARNOLD ARBORETUM INTERNATIONAL FIELD BIOLOGY COURSE 2006 Sinharaja World Heritage Site, Sri Lanka 30 July – 28 August 2006 Edited by Min Sheng Khoo, Cynthia Hong-Wa, and Rhett D. Harrison Center for Tropical Forest Science – Arnold Arboretum Asia Program University of Peradeniya Forest Department Sri Lanka Cover photo: Organizers, resource staff and participants of the sixth CTFS-AA International Field Biology Course 2006 (IFBC-2006) at Sinharaja Forest Bungalow, after the opening ceremony on 31 July 2006. See more photos on page 9. Preface i Preface The CTFS-AA International Field Biology Course is an annual, graduate-level field course in tropical forest biology run by the Center for Tropical Forest Science – Arnold Arboretum Asia Program in collaboration with institutional partners in South and Southeast Asia. The CTFS-AA International Field Biology Course 2006 was held at Sinharaja World Heritage Area, Sri Lanka, from 30 July to 29 August and was hosted by the Forest Department, Sri Lanka, and the University of Peradeniya, Sri Lanka. It was the sixth such course organized by CTFS-AA. Last year’s the course was held at Khao Chong, Thailand, and previous courses have been held in Malaysia. Next year’s course will be held at Xishuangbanna, Yunnan, China. The aim of these courses is to provide high-level training in the biology of forests in South and Southeast Asia. The courses are aimed at upper-level undergraduate and graduate students from the region, who are at the start of their thesis research or professional careers in forest biology. During the course, topics in forest biology are taught by a wide range of experts in tropical forest science. There is a strong emphasis on the development of independent research projects during the course. Students are also exposed to different ecosystem types, as well as forest related industries, through course excursions. The CTFS-AA International Field Biology Course 2006 was attended by 21 students from nine countries (Malaysia, Thailand, Philippines, UK, Taiwan, China, India, Madagascar, and Sri Lanka) and a total of 27 resource staff from a variety of national and international institutions gave lectures and practical instruction. The course in 2006 was implemented by Dr. Rhett Harrison (CTFS-AA), Drs. Savitri and Nimal Gunatilleke (University of Peradeniya, Sri Lanka), Mr. Anura Sathurusinghe (Forest Department, Sri Lanka), and Ms Luan Keng Wang (Raffles Museum, Singapore). Due to their considerable efforts the course proved to be an enormous success. The following report illustrates the hard work of the organizers and the enthusiasm and commitment of the students. We look forward to another successful course in 2007. Stuart J. Davies Director, Center for Tropical Forest Science International Field Biology Course 2006 Center for Tropical Forest Science – Arnold Arboretum Asia Program University of Peradeniya Forest Department Sri Lanka ii Acknowledgements Acknowledgements First, the organizers of the CTFS-AA International Field Biology Course 2006 wish to thank all the resource staff who gave their time to teach on the course. Without the commitment of these researchers, many of whom have now taught on the course each year for the past several years, to the success of the field course at Sinharaja, nothing could have been achieved. In particular we would like to thank Ms. Luan Keng Wang (Raffles Museum, Singapore), who as always assisted greatly in the smooth running of the course. We would also like to acknowledge the considerable help of the University of Peradeniya staff and field assistants, especially the following: Drs. Nirmal Weerasekera and Hashendra Kathriarachchi, Mr. Suranjan Fernando and Mr. Tiran Abewardena, who made the computer facility at Sinharaja possible; the field assistants Mr. T. M. N. Jayatissa, Mr. Anura Tennakoon, Mr. T. M. Ratnayaka, and Mr. R. Sirimanna; the drivers Mr. M. A. Gunadasa and Mr. T. Wickramasinghe; the field station caretaker Mr. Wimal Shantha Kumara; and the kitchen staff Mr. M. G. Jayaratna, Mr. K. G. Piyasena and his assistants in the Kudawa forest camp; Mr. Martin Wijesinghe and his family for their unreserved support before and throughout the workshop. A list of all the resource staff who contributed to the course appears at the end of these proceedings. We would like to thank the Forest Department, Sri Lanka for the enormous support they gave to the course. Especially, The Conservator General of Forests, Mr. M. P. A. U. S. Fernando for whole heartedly supporting this field course at Sinharaja from its inception to the end; Mr. E. J. M. J. K. Ekanayake (Additional District Forest Officer), Mr. D. P. Prasad (Forester) and other staff at the Sinharaja (Kudawa) Forest Department Facility for acceding to every request we made and giving their very best to make the course a success. For the course excursions, we received a great deal of support from various people. The staff at Minneriya Wildlife Sanctuary, especially Mr. P. M. Dharmatilleke (Assistant Director) and his staff at the Giritale Wildlife Training Centre, and Mr. M. H. Chirasena (Park Warden), who addressed the students, and his staff at the Minneriya National Park, who kindly hosted us at their respective facilities. Dr. Wolfgang Dittus and his staff at Polonnaruwa worked extremely hard to give the students both an excellent introduction to primate social ecology and a wonderfully enjoyable visit to the ancient capital (the cricket was a great hit!). As a result of our last minute change of schedule, Dr. S. Wijesundara (Director) was not available to host us at the Royal Botanic Gardens, Peradeniya and we are very grateful for the excellent tour given instead by Mr. D. M. U. B. Dhanasekara (Deputy Director) to these beautiful gardens. Dr. Hashendra Kathriarachchi excelled herself giving the course participants both an enormously enjoyable and informative hike around Horton Plains, rain and all. An idea of how much the excursions were appreciated can be gauged from the enthusiastic reports the students wrote about them. The organizers would like to thank all the above for hosting the course. Financial support for the field course came from the Center for Tropical Forest Science – Arnold Arboretum Asia Program of the Smithsonian Tropical Research Institute and the Arnold Arboretum of Harvard University. Tunghai University very generously provided additional funds through a grant to Dr. I Fang Sun, and funded the flights of the Taiwanese participants. The Eco-lodge project through a USAID grant provided additional funding to cover the costs of the participants from their project. In addition to hosting the course the Forest Department, Sri Lanka and the University of Peradeniya covered the costs of their employees and facilitated the course in various other ways for which we are very grateful. Thanks to all. The organizers, Rhett Harrison (Center for Tropical Forest Science –Arnold Arboretum) Savi Gunatilleke (University of Peradeniya) Nimal Gunatilleke (University of Peradeniya) Anura Sathurusinghe (Forest Department, Sri Lanka) International Field Biology Course 2006 Center for Tropical Forest Science – Arnold Arboretum Asia Program University of Peradeniya Forest Department Sri Lanka Contents iii Contents i Preface ii Acknowledgements iii Contents 1 Field Course Program ABSTRACTS 5 Sinharaja World Heritage Site I. A. U. N. Gunatilleke 6 An Introduction to CTFS-AA Stuart Davies 6 Field Practical on Tree Identification Jim LaFrankie 7 The CTFS-AA Trees of Tropical Asia Database Jim LaFrankie 7 Forests of Sri Lanka C. V. S. Gunatilleke 8 Plant Diversity across a Habitat Gradient I. A. U. N. and C. V. S. Gunatilleke 10 Insects and Insect Sampling David Lohman 10 Herbivory David Lohman 10 Ants Nihara Gunawardene 11 Termites (Order Isoptera) Sushila I. Vitarana 16 Bees Rhett D. Harrison 18 Birds S. Kotagama 18 Herpetological Studies K. Manamendra-Arachchi 19 Vertebrate Sampling Methods C. P. Ratnayke 19 Twenty Questions Rhett D. Harrison 21 Royal Botanic Gardens, Peradeniya Siril Wijesunbara 21 Elephant Ecology R. Sukumar 22 Primate Behaviour and Ecology W. P. J. Dittus 24 Niche Partitioning: The Comparative Ecology and Behavior of Three Species of Sympatric Primates at Polonnaruwa W. P. J. Dittus International Field Biology Course 2006 Center for Tropical Forest Science – Arnold Arboretum Asia Program University of Peradeniya Forest Department Sri Lanka iv Contents 24 Raising Sons and Daughters in Macaque Society W. P. J. Dittus 25 Horton Plains Rohan Pethiyagoda and C. V. S. Gunatilleke 27 An Introduction to R Campbell O. Webb 27 Statistics Richard T. Corlett 27 Tropical Forests Compared Richard T. Corlett 28 Frugivory and Seed Dispersal Richard T. Corlett 29 Conservation Biology - Effects of Small Population Size Priya Davidar 30 Invasive Species Channa Bambaradeniya 30 Plant Diversity in Forests: Negative Density Dependence David Burslem 33 Phylogenetic Methods Campbell O. Webb 33 Evolutionary Philosophy Shawn Lum 34 Molecular Ecology Shawn Lum 34 Fig Biology: An Intricate Interaction Rhett D. Harrison 35 Pollination Rhett D. Harrison 37 Seedling Ecology Mark Ashton 37 Tropical Forest Restoration Mark Ashton 38 Tropical Forest Silviculture / Regeneration Mark Ashton 38 Independent Student Projects Rhett D. Harrison EXCURSION REPORTS 40 Perahera Festival at Kandy, 7 August 2006 Harvey John D. Garcia, Cynthia Hong-Wa, and Simon Jiun-Nan Huang 41 Visit to Minneriya National Park, 8 August 2006 Nurfazliza bt. Kamarulbahrin