AUTECOLOGY OF THE SUNDA PANGOLIN (MANIS JAVANICA) IN SINGAPORE LIM T-LON, NORMAN (B.Sc. (Hons.), NUS) A THESIS SUBMITTED FOR THE DEGREE OF MASTER OF SCIENCE DEPARTMENT OF BIOLOGICAL SCIENCES NATIONAL UNIVERSITY OF SINGAPORE 2007 An adult male Manis javanica (MJ17) raiding an arboreal Oceophylla smaradgina nest. By shutting its nostrils and eyes, the Sunda Pangolin is able to protect its vulnerable parts from the powerful bites of this ant speces. The scales and thick skin further reduce the impacts of the ants’ attack. ii ACKNOWLEDGEMENTS My supervisor Professor Peter Ng Kee Lin is a wonderful mentor who provides the perfect combination of support and freedom that every graduate student should have. Despite his busy schedule, he always makes time for his students and provides the appropriate advice needed. His insightful comments and innovative ideas never fail to impress and inspire me throughout my entire time in the University. Lastly, I am most grateful to Prof. Ng for seeing promise in me and accepting me into the family of the Systematics and Ecology Laboratory. I would also like to thank Benjamin Lee for introducing me to the subject of pangolins, and subsequently introducing me to Melvin Gumal. They have guided me along tremendously during the preliminary phase of the project and provided wonderful comments throughout the entire course. The Wildlife Conservation Society (WCS) provided funding to undertake this research. In addition, field biologists from the various WCS offices in Southeast Asia have helped tremendously throughout the project, especially Anthony Lynam who has taken time off to conduct a camera-trapping workshop. The Singapore Zoo provided funding and also assisted in many aspects of the work. I would like to thank Charlene Yeong for the kind patience with all my requests and the wonderful offer of sending other Singapore Zoo staff (e.g. Faizal Hussin, Desmond Ling and Joe Ong) to assist in the fieldwork on Pulau Tekong. The vets (especially Dr. Oh Soon Hock) have never turned down a single request to help examine and treat wild pangolins before releasing them back into the wild. The National Parks Board (NParks) and the Ministry of Defence (MINDEF) granted the permission to conduct the research on both mainland Singapore and Pulau Tekong respectively. It was most privileged for me to have iii obtained kind permission from MINDEF to conduct long-term research on the largely forested island of Pulau Tekong. The staff from both the Public Relations Branch, MINDEF (especially Jude Ng) and CCO Pulau Tekong (especially Major Lim and SSG Hasan) have assisted greatly in the logistics. I am also grateful to the staff of NParks, especially Chew Ping Ting who has provided much assistance when using the GIS system and Benjamin Lee who is always enthusiastic with pangolin “rescue operations”. The research would not have been possible if not for the many people who took the trouble to travel to Pulau Tekong, endured the less-than-comfortable conditions at the base-camp and accompanied me for the many nights of fieldwork. I am most grateful to each and every one of them, and truly enjoyed their company. Of special mention are Chan Kwok Wai, Reuben Clements Gopalasamy, Jeremy Woon, Alan Yeo and Yeo Suay Hwee, who are ever so keen and responsive to last-minute requests. David Lohman provided much assistance for the identification of ants, while Reuben Clements Gopalasamy provided guidance for the statistical analysis. Friends and colleagues from the Systematics and Ecology Laboratory, and the Raffles Museum of Biodiversity Research gave me much support, advice and encouragement along the way. I am also grateful to the friends and collaborators in the neighbouring countries who provided valuable information on the various aspects discussed in this work. Last but not least, I would wish to thank my parents, other members of the family and also my fiancée for tolerating my idiosyncrasies (especially that of sneaking into the forest during the night) all this while. This piece of work (and other research projects) would not have been possible without their continued encouragement and support. iv TABLE OF CONTENTS Acknowledgements………………………………………………………………… iii List of tables, figures and plates…………………………………………………… viii Summary…………………………………………………………………………… xi Chapter 1. Introduction 1.1 Pangolin species and taxonomy …………………………………… 1 1.2 Conservation status of Asian pangolins……………………………. 3 1.3 Literature review on Manis javanica………………………………. 4 1.4 Status of Manis javanica in Singapore…………………………….. 9 1.5 Specific objectives…………………………………………………. 12 2. Materials and methods 2.1 Study site…………………………………………………………… 13 2.2 Capture and radio-telemetry……………………………………….. 14 2.3 Camera-trapping and natal den description………………………... 17 2.4 Prey preference…………………………………………………….. 18 2.5 Data analyses 2.5.1 Sexual dimorphism and sex ratio…………………………... 19 2.5.2 Daily activity patterns……………………………………… 20 2.5.3 Home range………………………………………………… 20 2.5.4 Habitat preference………………………………………….. 23 2.5.5 Prey preference…………………………………………….. 25 2.5.6 Miscellaneous (data from mainland Singapore)…………… 26 3. Results 3.1 Body measurements of Sunda Pangolin on Pulau Tekong………… 27 v 3.2 Home-range, activity cycle and other spatial-temporal statistics….. 28 3.3 Habitat preference………………………………………………….. 31 3.4 Natal den usage by pregnant and postpartum Manis javanica MJ6.. 32 3.5 Prey preference…………………………………………………….. 33 3.6 Predation of Manis javanica by Python reticulatus………………... 35 4. Discussion 4.1 Body measurements of Sunda Pangolin on Pulau Tekong………… 37 4.2 Home-range, activity cycle and other spatial-temporal statistics….. 38 4.3 Prey preference…………………………………………………….. 44 4.4 Habitat preference………………………………………………….. 46 4.5 Reproductive biology of MJ6……………………………………… 49 4.6 Predators of Manis javanica……………………………………….. 51 5. Overall conclusions and recommendations 5.1 Challenges for future pangolin research.…………………………... 54 5.2 Priorities for conservation of pangolin species…………………….. 57 5.3 Status of Manis javanica in Singapore…………………………….. 61 5.4 Conclusion…………………………………………………………. 62 References………………………………………………………………………….. 63 Tables, Figures and Plates…………………………………………………………..73 Appendices I. List of Manis javanica sightings in Singapore for the past two decades……………………………………………. I-1 II. Percentage of the various ant genera collected with cheese bait, sugar bait and the combined values. ………………….. II-1 III. Percentage of foraging time spent on the various ant genera by three adult male Manis javanica..…………………... III-1 vi LIST OF TABLES 1. A summary of information on the eight pangolin species of the Old World. 2. Known uses and supposed benefits of pangolin parts. 3. Body measurements (HB – head-body length; T – tail length, in cm; body mass in kg) and sex of the Manis javanica captured on Pulau Tekong. 4. Speed statistics (m/ hr) of adult male Manis javanica on Pulau Tekong. 5. Home-range estimates (ha) of adult Manis javanica using minimum-convex polygon (MCP), harmonic mean and kernel methods, with the various parameters stated and using non-autocorrelated radio locations. 6. Simplified ranking matrix for Manis javanica based on a) comparing roportional habitat use within minimum convex polygon (MCP) home ranges with proportions of total available habitat types, and b) comparing proportion of radio locations (with 20-m radius buffer) for each animal in each habitat type with the proportions of each habitat types within the animal’s MCP range (see text for more details). “+” implies the habitat type of the r was preferred to the habitat type in the row and “-” implies the reverse; a triple sign represents significant deviation from random at P < 0.05. 7. Jacob’s index for each habitat used by pangolins on Pulau Tekong at second-order and third-order selection. 8. Description, measurements and duration of usage of the three dens used by female Manis javanica MJ6 from September to December 2005. 9. Mean and maximum time (min) spent on feeding by three male Manis javanica. 10. Percentage of time spent feeding on ants and termites by three tagged male Manis javanica. 11. Jacob’s index of the ant subfamily consumed by adult male Manis javanica on Pulau Tekong. 12. Jacob’s index of the ant genera consumed by adult male Manis javanica on Pulau Tekong. 13. Date of the locations of radio-transmitter in Figure 1, distance from previous location and description of location. 14. Reported date of birth and body measurements of neonate Manis spp. vii LIST OF FIGURES 1. Location of all known records of Manis javanica sightings and roadkills in Singapore, from 1985-2007. 2. Map of Pulau Tekong, relative to mainland Singapore. The shaded grey shaded area is the only reservoir on the island and the cross-shaded patches are the built- up areas. The remainder of the island is covered with the vegetation. 3. Radio-telemetry error polygons under extreme conditions of maximum distance from transmitter (i.e. 200 m) and maximum bearing difference from optimal angle (i.e. 60° and 120°). 4. Diel activity pattern of female Manis javanica MJ6 (mean ± S.E.), based on 30 days of camera-trapping data from 8th September to 12th December 2005. 5. Diel activity pattern of each of the four adult male Manis javanica. No activity was recorded between 0800 hrs to 2000 hrs. 6. Diel activity pattern of adult male Manis javanica (n = 4), with S.E. as error bars. No activity was recorded between 0800 hrs to 2000 hrs. 7. Accumulative 100% minimum-convex polygon home-range estimates of individual adult male Manis javanica with increasing number of radio-tracking days. 8. Autocorrelation plot of Schoener’s index with time interval of Manis javanica MJ9. Reasonably-independent radio locations were obtained at approximately 2- hr interval (green vertical line), where there are three consecutive values of Schoener’s index greater than 1 (see text for more details). 9. Position and shape of 100 % minimum convex polygon home-range of the four adult male Manis javanica (MJ9, MJ12, MJ22 and MJ23) on Pulau Tekong.