The Tortoise Manouria Emys Emys: Behaviour and Habitat in the Wild
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The tortoise Manouria emys emys: behaviour and habitat in the wild M. Sc. Thesis by Klaus Høybye-Mortensen Supervisor: June 2004 Associate professor, Ph.D. Institute of Biology, Ole Næsbye Larsen University of Southern Denmark Klaus Høybye-Mortensen, University of Southern Denmark, June 2004 ____________________________________________________ Dedicated to the memory of Dr. Annelisa M. Kilbourn The tortoise Manouria emys emys: behaviour and habitat in the wild __________________________________________________ Acknowledgements The author wishes to thank the following organisations and companies for their financial support: Chelonian Research Foundation (Linnaeus Fund), DANCED-SLUSE (TFS-Grant), Internatiotionaliseringspuljen (University of Southern Denmark), HKC-Data, Spejder Sport, Nokia Denmark, MT Contractors, Centrum Skilte, Kosmos Grafisk, Institute of Biology University of Southern Denmark and Institute of Tropical Biology and Conservation, Universiti Malaysia Sabah. Without their support, this project would not have been realised. Thanks to Ole Næsbye Larsen for agreeing to be my supervisor. His patience and help during the whole process has been invaluable to me. Thanks to Frank B. Mortensen for designing and manufacturing the radio telemetry system and assisting in other technological matters. My deepest thanks goes to a long list of people who helped me during my two stays in Sabah: Dr. Datin Prof. Maryati Mohamed for accepting to be my local supervisor, Dr. Annelisa Kilbourn for being a good friend and field colleague, Dr. Edwin J. Bosi for introducing me to this little known tortoise species, Leni Tupang Thomas and the rangers (Makarius, Nimrod, Tinju, Vincent, Francis, Sarinus and Samson) from SOS Rhino for introducing me to jungle life and their help in finding study animals. Thanks to David Gakin, David Antonius, Herman Stawin, Justin, Rayner, Justin and Zul from the Wildlife Department at Tabin Station and their families for taking good care of me. A special thanks to Herman Stawin for inviting me to watch the World Cup 2002 at his house. Thanks to Mike Cadman and the staff at Trekkers Lodge for their care during my stays in Kota Kinabalu. In addition, I would like to thank my fellow students Alan and Jakob for their encouragements and good spirits throughout my university life. Special thanks to Anja for her love and support throughout the project, even though it meant that I had to leave her behind for several months. Thanks to Chuck Schaffer for providing me with “rare” literature pertaining to Manouria emys and his visit in Tabin Wildlife Reserve. Thanks to Betina, Kevin and Chuck for reviewing the thesis and numerous suggestions for improvement. Klaus Høybye-Mortensen, University of Southern Denmark, June 2004 ____________________________________________________ Table of contents i. Landskildpadden Manouria emys emys: naturlig adfærd og habitat ...............................i ii. Abstract ........................................................................................................................... iii iii. Abbreviations ...................................................................................................................iv 1 Introduction.......................................................................................................................1 1.1 The tortoise Manouria emys emys: a biological review ............................................... 1 1.1.1 Distribution......................................................................................................... 2 1.1.2 Morphology ........................................................................................................ 2 1.1.3 Sexual dimorphism and behaviour....................................................................... 3 1.1.4 Nesting ............................................................................................................... 5 1.2 Conservation status..................................................................................................... 6 1.3 Objectives of the present study ................................................................................... 7 2 Methods .............................................................................................................................9 2.1 Research time frame ................................................................................................... 9 2.2 Study area: General description .................................................................................. 9 2.3 Topography .............................................................................................................. 12 2.4 Climate..................................................................................................................... 12 2.5 Flora......................................................................................................................... 14 2.6 Fauna........................................................................................................................ 14 2.7 Recording techniques................................................................................................ 16 2.7.1 Radio telemetry................................................................................................. 16 2.7.2 GPS and GIS..................................................................................................... 18 2.7.3 Home range ...................................................................................................... 19 2.7.4 Daily movements .............................................................................................. 21 2.7.5 Activity............................................................................................................. 21 2.7.6 Diet................................................................................................................... 23 2.7.7 Behaviour ......................................................................................................... 24 2.7.8 Temperature...................................................................................................... 25 2.7.9 Habitat .............................................................................................................. 25 2.7.10 Morphometry.................................................................................................... 26 2.7.11 Statistics ........................................................................................................... 28 3 Results..............................................................................................................................29 3.1 Study animals ........................................................................................................... 29 3.1.1 First sighting ..................................................................................................... 29 3.1.2 Morphometry.................................................................................................... 30 3.2 Diet .......................................................................................................................... 31 3.3 Home Range............................................................................................................. 34 3.4 Activity .................................................................................................................... 43 3.5 Habitat...................................................................................................................... 49 3.5.1 Forest type ........................................................................................................ 49 3.5.2 Temperature and microclimate.......................................................................... 50 3.5.3 Water................................................................................................................ 53 3.6 Nesting behaviour - a case story................................................................................ 55 3.7 Mating- a case story.................................................................................................. 58 4 Discussion ........................................................................................................................60 The tortoise Manouria emys emys: behaviour and habitat in the wild __________________________________________________ 4.1 Study animals and morphometry............................................................................... 60 4.2 Habitat...................................................................................................................... 62 4.3 Temperature preferences........................................................................................... 63 4.4 Diet .......................................................................................................................... 66 4.5 Activity .................................................................................................................... 70 4.6 Home range .............................................................................................................. 72 4.7 Nesting ..................................................................................................................... 75 4.8 Mating ...................................................................................................................... 78 5 Conclusion .......................................................................................................................79