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The Ecological Functions of Asian Elephants in the Sundaic Rainforest: Herbivory and Seed Dispersal

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The Ecological Functions of Asian Elephants in the Sundaic Rainforest: Herbivory and Seed Dispersal The Ecological Functions of Asian Elephants in the Sundaic Rainforest: Herbivory and Seed Dispersal A thesis submitted to the University of Nottingham Malaysia for the degree of Doctor of Philosophy Lisa Ong 2020 Statement of word count: Number of words contained in the document including reference and appendices: 83451 Abstract Asian elephants (Elephas maximus) are among the last remaining megaherbivores in the Sundaic rainforests. In this thesis, I aimed to improve our understanding of the ecological roles of rainforest-dwelling Asian elephants through their herbivory and seed dispersal behaviours. A comprehensive review in Chapter One revealed that there have been few investigations of their interactions with the community and the habitats they live in. In Chapter Two, I advanced our understanding of Asian elephants in modifying the rainforest structure by quantifying their diet selection and herbivory impacts. The principal foods of Asian elephants in rainforests were palms (39% of total trunkfuls consumed) and trees (30%). They preferred monocots, particularly palms (preference ratio=4.6; below 1=avoidance) that are not abundant in the forest, and most monocots sustained high damage due to elephant foraging (above 40% in foraged patches). Their feeding behaviour was associated with a reduced palm density in the Royal Belum rainforest where elephants roam, as compared to a forest lacking elephants in Krau Wildlife Reserve. Although large tree saplings (around 21 cm diameter) were vulnerable to elephant damage, they had high recovery rates, and elephant herbivory is likely not the sole factor causing lower tree sapling density in Belum. Asian elephants are important seed dispersers. In Chapter Three, I recorded rare interactions of Asian elephants and a large-fruited rainforest tree, Irvingia malayana (Irvingiaceae) through the use of camera-traps. The removal rate of fruits by elephants was low, yet seed dispersal by elephants allowed Irvingia malayana seeds to escape from high levels of seed predation by wild boars. In Chapter Four, using network analysis, I identify Asian elephants as dominant functional generalists promoting seed dispersal diversity within the community. This is the first highly diverse seed dispersal network in the Sundaic region, built using an inter-disciplinary approach, which combines ethnobotany knowledge, feeding signs on fruits, camera-trapping, and published literature. Elephants had high species strength and were important to both the dispersal of plants central to the network and the dispersal of large-fruited species. Along with other important seed dispersers such as gibbons (the most efficient seed dispersers), binturong, civets, and the Malayan sun bear, they promoted network nestedness and functional redundancy. Simulation of defaunation of important, highly-connected seed dispersers caused adverse co-extinctions of seed dispersal interactions. Urgent protection of important and vulnerable rainforest species is needed in the Sundaic region. This thesis confirms the importance of Asian elephants as selective feeders for palms and tree saplings. It describes their interactions with tropical Sundaic fruits and confirms Asian elephants as functionally dominant seed dispersers within the seed dispersal community. i Acknowledgment I was privileged to work in one of the world’s most remarkable rainforest, the Belum Temenggor Forest Complex, to study the last remaining megaherbivores, the Asian elephants, and to extend the study to nature’s web of life with the local indigenous people, the Orang Asli. This research has received generous funding support from Yayasan Sime Darby (grant M0005.54.04), and the University of Nottingham Malaysia. I thank Puan Dr. Hajjah Yatela Zainal Abidin for the continuous support she and her team had provided to MEME. I thank the Department of Wildlife and National Parks (DWNP), the Perak State Parks Corporation, the Forestry Department of Perak, and the Department of Orang Asli Development (JAKOA) for the approval of permits. I am grateful to Nasharudding Osman and Mohamad Khairul Adha bin Mat Amin from the Kuala Gandah National Elephant Conservation Center (NECC), for making the elephant feeding observations possible in Krau Wildlife Reserve. My supervisor, Prof. Ahimsa Campos-Arceiz provided this special opportunity and allowed me the freedom to explore my ideas. This thesis would have been impossible without his enthusiasm for seed dispersal and megafauna, and faith in his Ph.D. student. The dynamics he created for his lab, the Management and Ecology of Malaysian Elephants, MEME, has introduced diversity (people and science) into my research environment. My co-supervisor, Dr. Kim McConkey has inspired me with her unrelenting patience for young researchers and obsession with nature. She has stimulated my thinking in the forests and strengthened my knowledge in natural history and seed dispersal science. I am incalculably grateful for both of their rich contributions and the time they spent reviewing the long bouts of writing before this final thesis. I was fortunate to collaborate with Prof. John Terborgh and Dr. Lisa Davenport, who enriched my journey with their presence. John Terborgh has been a source of great wisdom. He has stimulated an intellectual atmosphere with his sharpness in piecing up the puzzles of the natural system and had inspired me with his vivid writings. I appreciated Lisa Davenport’s calm and reassuring presence during our trips. Her interests for birds and the amazing migratory route she showed of black skimmers crossing the Peruvian Andes had set a little spark in me for birds. I appreciate the feedback from my examiners. Dr. Markus Eichhorn was the internal examiner for my first- and second-year review, and a good lecturer who answered my questions in ecology during my undergraduate years. As a conservation student, it is inevitable to have self-doubts while carrying out our research. During my final year review with the head of school, Environmental and Geographical Science, Prof. Christopher Gibbins, I felt encouraged by his ii words that a Ph.D. student’s responsibility is to produce good science. Both examiners had provided valuable advice for my progress. Equally important to the successful completion of this thesis, was the constructive and positive feedback provided by both internal (Chris Gibbins) and external examiners. I have benefitted from the penetrating mind and experience of Dr. Luis Santamaria during the viva. My sincere appreciation to Vivienne Loke Pei Wen, Loo Yen Yi, Tan Wei Harn, and Nurul Ain binti Amirrudin for being steadfast and curious thinkers and providing key support as research assistants. Their commitment was important to the project. I felt blessed to have Hii Ning and Adeline Hii as close friends who had worked conscientiously, shared ideas, and explored the forests together with me. Param bin Pura, Ridzuan bin Angah, Cherang Mohamad Tauhid bin Tunil, Husin a/l Sudin, and Khairil bin Othman had been generous with their ecological knowledge and skills, welcomed me to the forest, and ensured the safety of all members in good and bad weathers. Alicia Solana-Mena, field manager of MEME, introduced me around Belum-Temenggor and Gerik town and provided key logistics support. Her familiarity and good relations with the indigenous communities around Belum-Temenggor were helpful in liaising with the right participants for the surveys. Praveena Chackrapani, our team manager has been the pillar for administration and led us to productive directions behind the scenes. I appreciate discussions with other colleagues in MEME on topics that extend beyond elephants, including Dr. Lim Teck Wyn, Dr. Antonio de la Torre, Ange Tan, Dr. Nagulendran Kangayatkarasu, Dr. Wong Ee Phin, Nur Aida Ab Ghani, Vanitha Ponnusamy, Dr. Jamie Wadey, Or Oi Ching, Sinchita Sinha, Dr. Vivek Thuppil and Nurul Azuwa. I thank volunteers and visitors who provided help, took moments off their work, or joined for short field trips for the research project ― including Lily Ong, Mike Meridith, Stephane Arul Mariampilai, Lim Teck Wyn, Sze Yanghong, Violet Loke Pei Xing, Todd Bechtold, Sanjay Ramarao, Bryan Wee, Hii Ning, Chong Siu Ho, Ong Lenna, Ong Lihoon, Gracia Chong Kai Xing, Mohammad Hafiz bin Khairul Wizam, Loh Lih Yann, Bhaskar Reddy, Weinstock Justin, Panazzolo Jessie, Angel Sanchez, Francesc Llimona, and Ange Tan. I deeply appreciate the assistance of all mahouts from the Kuala Gandah National Elephant Conservation Center (NECC); some endured the pain of a wasp attack during our observations: Ali a/l Gadong, Fadhlul Hadi bin Abdul Samad, Khairul Affendi, Muhammad Izamin, Lukman bin Muhammad, Syed NorAzlin, Sulaiman a/l Mar, Rizuan bin Muhammad, and Mohamad Sabri bin Aseli. It was such a delight to have the elephants Cherry, Indah, Sanum, Siti and Timur joined us for the feeding observations. iii This research owes its results to the Orang Asli of Belum-Temenggor, especially our field assistants, and the participants who had contributed to the local ecological knowledge (LEK). I warmly thank Roy bin Hani, Ismail bin Jijih, Mamak bin Pinang, and Hasan a/l Padan from Kampung (Kg.) Tiang, Ismail bin Alok, Eim bin Janing, Hamid bin Pura, Apok bin Alok, Jaafar bin Alok, and Abain bin Goying from Kg. Semerloh, Lanas bin Tirai, Jali bin Ranting, Rajut anak Tirai, Tok Batin Radi Anak Langsat, and Hamid bin Kelilik from Kg. Sungai Kijar, Hasim bin Charol, Ajim bin Along, Sham bin Hobel, and Long bin Titik from Kg. Pulau Tijuh, Beracut bin Lebak, Arrifyn bin Dris, Deney bin Abain, and Johar bin Beracut from Kg. Desa Damai, and Selerwer bin Ngengong, Dalong bin Piloi, Selan bin Melensing, and Saken bin Ubel from Kg. Banun. I am filled with gratitude and was truly humbled by their indigenous knowledge. Most importantly, I am grateful to my beloved parents Ong Pia and Tan Bee Choo for enduring my frequent absences from home and allowing their daughter into the jungles. I am blessed to have my family members Kian Hong, Caiping, Lily, Chunxin, Lenna, Paul, Lihoon, and Kelvin, my nieces and nephews, see the value of my work and supported one another while I pursued this research.
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