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Orangutan Locomotion

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Orangutan Locomotion HABITAT VARIATION AND ITS INFLUENCE ON THE LOCOMOTOR ECOLOGY OF WILD ORANGUTANS by KIRSTEN L. MANDUELL A thesis submitted to the University of Birmingham for the degree of DOCTOR OF PHILOSOPHY supervised by Dr. Susannah Thorpe Locomotor Ecology and Biomechanics Lab School of Biosciences College of Life & Environmental Sciences University of Birmingham September 2012 University of Birmingham Research Archive e-theses repository This unpublished thesis/dissertation is copyright of the author and/or third parties. The intellectual property rights of the author or third parties in respect of this work are as defined by The Copyright Designs and Patents Act 1988 or as modified by any successor legislation. Any use made of information contained in this thesis/dissertation must be in accordance with that legislation and must be properly acknowledged. Further distribution or reproduction in any format is prohibited without the permission of the copyright holder. Abstract Orangutans are the largest arboreal primate and have a diverse locomotor repertoire. The principal aim of this thesis was to explore the dynamic between morphology, behaviour and habitat to better understand the influences on orangutan locomotion. Positional behaviour data was collected at two peat-swamp forest sites: Sabangau, Central Kalimantan; and Suaq Balimbing, Aceh. We quantified forest structure and support availability in the dry-lowland forest of Ketambe, Aceh, in addition to the aforementioned peat-swamp forests and found that the three forests were structurally different. We used a remote measuring technique to compare limb morphology between species, and found they were similar suggesting selection for an optimal limb length. We found that habitat had a stronger influence on locomotion than either species or study site. Orangutans in different habitats had similar profiles of preferred supports, with the exception that the Sumatran species (Pongo abelii) had a preference for lianas. Orangutans in Sumatran peat-swamp forest used more compliant supports than recorded in dry-lowland forest. However, pronograde bridging was also used to negotiate the most flexible supports. This thesis has shown that habitat has a strong influence on orangutan locomotor behaviour, which is important since their habitat is becoming increasingly altered through human disturbance. Acknowledgements This research was only possible due to the assistance of a great many people. To any of you accidentally omitted here, sorry. and thanks! This work was carried out under the supervision of Dr Susannah Thorpe, and I thank her wholeheartedly for her advice, encouragement and support throughout, and for giving me this amazing opportunity. This work would not have been possible without the cooperation of numerous Indonesian authorities that granted permissions for my work in Indonesia. I thank RISTEK, the Centre for the International Cooperation in Management of Tropical Peatlands (CIMTROP), UNAS, SOCP, the Indonesian Department of Forestry (PHKA), PBKEL and TNGL. In this regard, I am particularly grateful to Dr Ir Suwido H. Limin, CIMTROP Director, for sponsoring me during my research in Sabangau, and to Pak Tatang Mitra Setia for sponsoring my research in the Leuser Ecosystem. I am eternally grateful to the “OUtrop family”, particularly Dr Mark Harrison for introducing me to the study of wild orangutans as his research assistant in 2005, without which I may never have had opportunity to undertake my own research of these fascinating animals. I am also particularly grateful to Dr Helen Morrogh-Bernard for her encouragement when I first expressed an interest in orangutan research. I am extemely grateful to Ian Singleton, Serge Wich and Maria van Noordwijk for enabling me to conduct research in Suaq and Ketambe. I would also like to thank the numerous friends who supported me and kept me sane during my time in the field: Grace Blackham, Susan Cheyne, Simon Husson, Laura Graham, Laura D’Arcy, Angela Benton-Browne, Rosalie Dench, Andrea Höing, Reychell Chadwick, Adam van Casteren, Andrea Permana and Gail Campbell-Smith. Numerous assistants aided in data collection and provided friendship and support whilst in the field, and I offer my most sincere gratitude to them all. Sabangau: Twenti Nolosa Firsman, Adul, Aman, Ambut, Santiano, Yudhi and Dewa. The kindhearted Ibu Yanti, who over the years has become my “Indonesian mother” and together with Mas Lis helped keep me fed and healthy. I am also grateful to Siska at CIMTROP who kept all my admin requirements in order. Ketambe: my assistant Adam, Pak Nusuar, Isa, Agusti and the SOCP field assistants: Mdin, Rabusin, Sumurudin, Roma and Kamarudin and the cooks, Silas, Dina and Baby. Suaq Balimbing: Pak Samsuar, Pak Azril, Pak Ishak and his wife, Herman Denias, my assistant Syharul and pendampingan: Agam, Amar, Fauzi and Adai; the SOCP field assistants: Mahmudin, Fikar, Izumi, Armas and Toni who taught me a lot about the forest of Suaq, were great fun to be around, and always pulled me out of the mud! I am also grateful to Pak Rustam and Pak Syafi’i who looked after me and kept me fed and watered. I would also like to thank the people at the SOCP office in Medan for their help and friendship, Santi DeLavita Lubis, Julie Simanjuntak, Ines, Novi, Melda, Angke and Yasmine. I do miss our girly karaoke nights! I made many friends during my time in Indonesia, too many to mention individually, but I am grateful to you all, and will always have fond memories of your beautiful country. My thanks go to all of the people at Birmingham. Jackie Chappell, my internal assessor, for her useful feedback on reports, the members of the Locomotor Ecology and Biomechanics Lab, Sam Coward, Abigail Phillips, Julia Myatt and Emma Tecwyn for many useful discussions. I would also like to thank the other friends I have made at Birmingham over the years, Simone, Katy, Kaat, Jolyon, Emma and Camille. I am particularly grateful for my friendship with Zoe Demery, who has been a constant source of support. Thank you for your kindness, your computer software expertise, and for being such a good friend. I thank all the orangutans, it was a privilege to spend time with you in your natural environment, one which I will always remember. Finally, I am grateful to my family and friends for their patience, love and support. Especially to my husband Julius, who always believed in me and never stopped encouraging me to follow my dreams. Thank you to the University of Birmingham and the NERC for funding this project. Contents Contents i List of Figures iv List of Tables vi 1 General introduction 1 1.1 Ecomorphology . 1 1.2 Orangutan Biology . 2 Taxonomy . 2 Distribution . 3 Morphology . 6 Species Variation . 9 Diet and Foraging Strategy . 12 1.3 Orangutan Habitat . 13 Forest Productivity . 13 Peat-Swamp Forests . 14 Dry lowland Forest . 16 1.4 Orangutan Evolution (Palaeontology and Functional Anatomy) 17 1.5 Positional Behaviour . 23 Primate Positional Behaviour . 23 Orangutan Positional Behaviour . 27 1.6 Study Sites . 32 Sabangau . 32 Suaq Balimbing . 34 Ketambe . 36 1.7 Aims and Objectives . 37 1.8 Thesis Overview . 38 2 Locomotion in Sabangau 41 2.1 Introduction . 43 2.2 Methods . 47 Field Study . 47 Statistical Analysis . 51 2.3 Results . 56 Descriptive Data . 56 Variable Associations . 58 Analysis of Contingency Tables . 61 2.4 Discussion . 66 2.5 Conclusions . 76 3 Support Use in Sabangau 79 3.1 Introduction . 81 3.2 Methods . 85 Field Study . 85 Statistical Analysis . 88 3.3 Results . 90 Descriptive Statistics . 90 Model Interpretation . 91 3.4 Discussion . 98 3.5 Conclusions . 107 4 Forest Structure and Support Availability 111 4.1 Introduction . 113 4.2 Methods . 118 Study Sites . 118 Habitat Survey . 119 Travel Trees . 123 Statistical Analysis . 124 4.3 Results . 126 Tree and Liana Density . 126 Canopy Variables and Support Availability . 126 Support Use . 129 Assessment of Preference . 131 Travel Trees . 132 Locomotion . 135 4.4 Discussion . 137 4.5 Conclusions . 148 5 Comparative Limb Morphology 151 5.1 Introduction . 153 5.2 Methods . 159 Limb Measurements . 163 Photograph Analysis . 164 Statistical Analysis . 166 5.3 Results . 166 Nyaro Menteng . 166 Validation . 168 ii Inter-specific Differences . 172 5.4 Discussion . 172 5.5 Conclusions . 179 6 Habitat Influences Locomotion 181 6.1 Introduction . 183 6.2 Methods . 188 Field Study . 188 Statistical Analysis . 189 Model Selection . 191 6.3 Results . 192 6.4 Discussion . 201 6.5 Conclusions . 207 7 Orangutans in peat swamp use more compliant supports 209 7.1 Introduction . 211 7.2 Methods . 216 Field Study . 216 Statistical Analysis . 217 7.3 Results . 219 7.4 Discussion . 226 7.5 Conclusions . 232 8 General Discussion 233 8.1 Introduction . 233 8.2 Summary of Main Results . ..
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