MODELLING THE RESPONSE OF LIZARDS TO THERMAL LANDSCAPE Teng Fei Examining committee: Prof. dr. V.G. Jetten University of Twente Prof. dr. A. Veldkamp University of Twente Prof. dr. W. Bouten University of Amsterdam Prof. dr. R.R. Giménez University of Malaga ITC dissertation number 231 ITC, P.O. Box 6, 7500 AA Enschede, The Netherlands ISBN 978-90-6164-359-3 Cover designed by Benno Masselink Printed by ITC Printing Department Copyright © 2013 by Teng Fei MODELLING THE RESPONSE OF LIZARDS TO THERMAL LANDSCAPE DISSERTATION to obtain the degree of doctor at the University of Twente, on the authority of the rector magnificus, prof.dr. H. Brinksma, on account of the decision of the graduation committee, to be publicly defended on Wednesday July 10, 2013 at 14:45 hrs by Teng Fei born on 2 August 1981 in Shanghai, China This thesis is approved by Prof. dr. Andrew K. Skidmore, Promotor Dr. Albertus G. Toxopeus, Assistant promotor Dr. Tiejun Wang, Assistant promotor ACKNOWLEDGEMENTS This PhD thesis is the outcome of years of study at the department of natural resources of the Faculty of Geo-information Science and Earth Observation, University of Twente. This PhD journey for me was a highly valuable period of professional and human growth in a friendly international atmosphere. My first thoughts go to my promoter, Prof. Andrew K. Skidmore, without whom this thesis would not have existed. He was and continues to be an ideal mentor. He was always there to put me back on my feet and provided direction with his wisdom. I will always remember the first day I was in his office, he encouraged me to think more about the ‘why questions’ at the beginning of a scientific research project. This suggestion echoed in my ears as the years passed and as time and hard work slowly turned me from an engineer into a scientific worker. It is my good fortune and honour that his wisdom infuses every part of my research: from proposal preparation to experimental design, from trouble shooting to article writing. I will also cherish the fantastic memories of the annual PhD student BBQs hosted by the friendly Skidmore family. I am greatly indebted to my co-supervisors, Dr. Albertus G. Toxopeus, Dr. Valentijn Venus and Dr. Tiejun Wang, for their strong, valuable, and patient advice on this journey. I am grateful for their guidance on mathematical modelling and reptilian physiology, for their strong support on the design and implementation of the animal experiments, and for helping guide my research through areas of thick fog. To the many researchers who gave me valuable comments and meaningful discussion on this journey, I want to express my gratitude and respect to these people, at ITC of Twente: Martin Schlerf, David Rossiter, Jan de Leeuw; at Wageningen University: Prof. Herbert Prins, Jingsong Li, Yuan Zeng and, at Wuhan University: Guofeng Wu, Wenxiu Gao, Yaolin Liu, Yanfang Liu, Qinyun Du and Lin Li. Thanks to Dierenpark De Oliemeulen my cooperation partner on site where an animal experiment was carried out. Special thanks go to Mr. Sjef van Overdijk who assisted setting up the experiment and maintained the servers placed in the i zoo. I was much impressed by his deep knowledge of biology and his enthusiasm for animals. To my friends, I cannot stop smiling when thinking of you. Thanks go to my colleagues in ITC, in particular Mohammad Abouali, Wei Ouyang, Paco Ferri, Roshanak Darvishzadeh, Sabrina Carvalho, Anas Fauzi, Atkilt Girma, Aidin Niamir, Claudia Pittiglio, Ullah Saleem, Ben Wielstra, Ha Nguyen, Tyas Basuki, Moses Cho, Jelle Ferwerda, Mhosisi Masocha, Nicky Knox, Filiz Bektas Balcik, Ajay Bhakta Mathema, and to the warm Chinese group at ITC, especially Xiang Zhang, Yali Si, Xi Zhao, Pu Hao, Fangfang Chen, Xia Li and Dongpo Deng. They gave me lots of invaluable suggestions not only on my research but also on every aspect of my PhD life – from the selection of the presentation template to the selection of beer. My deep gratitude goes to Ms. Eva Skidmore whose excellent English editing helped me to publish my first paper. I also appreciate Esther Hondebrink who offered her generous help on the translation of the abstract of this thesis into the Dutch language. Many thanks go to Mr. Stephen C. McClure and Dr. Stephan Angsuesser who helped proofreading several thesis chapters. Special thanks go to my friends Mr. Ding Zheng and Ms. Lichun Wang in Enschede, the Netherlands. They treated me like if I were their own son. I do, however, expect for one more badminton game. Most of all I would like to thank all my family who, as always, has given me endless love and support throughout my PhD journey. In particular I would like to thank my wife Meng Bian and my son Alan for their love and understanding. I thank my mother-in-law who spent a tremendous amount of time caring for my son. Without your support, I could do nothing. I am sincerely grateful to the review committee for their supportive and rich comments on this PhD thesis. Their careful comments gave me strong guidance on how to improve the work and provided extremely valuable insights to guide the future progress of my research. ii TABLE OF CONTENTS Acknowledgements ............................................................................. i Chapter 1 General Introduction..................................................... 1 1.1 Background .................................................................................................................. 2 1.2 Research Problem Statement ............................................................................... 6 1.3 Research Objectives and Questions ................................................................... 9 1.4 The Target Species .................................................................................................... 9 1.5 Outline of the Thesis ............................................................................................. 11 Chapter 2 A Body Temperature Model for Lizards as Estimated from the Thermal Environment......................................................... 15 2.1 Introduction ............................................................................................................. 17 2.2 Model Description .................................................................................................. 19 2.2.1 Heat Exchange Terms ........................................................................................ 19 2.2.2 Body Temperature Simulation ....................................................................... 22 2.3 Model Evaluation.................................................................................................... 23 2.3.1 Experimental Design ........................................................................................... 23 2.3.2 Calibration of the Body Temperature Model........................................... 25 2.3.3 Validation of the Body Temperature Model ............................................ 26 2.3.4 Sensitivity Analysis of the Body Temperature Model .......................... 27 2.4 Results ........................................................................................................................ 27 2.4.1 Absorbance of the Lizard’s Skin..................................................................... 27 2.4.2 Calibration of the Body Temperature Model........................................... 28 2.4.3 Validation of the Body Temperature Model ............................................ 29 2.4.4 Sensitivity Tests of the Body Temperature Model ................................. 30 2.5 Discussion ................................................................................................................. 33 2.6 Conclusion ................................................................................................................. 36 Chapter 3 Predicting Micro Thermal Habitat of Lizards in a Dynamic Thermal Environment ....................................................................... 37 3.1 Introduction ............................................................................................................. 39 3.2 Method ........................................................................................................................ 41 3.2.1 Target Animal and Experimental Setup .................................................... 41 3.2.2 Data Collection ...................................................................................................... 43 3.2.3 Model Building ....................................................................................................... 44 3.2.4 Model Validation .................................................................................................. 47 3.3 Results and Discussion ........................................................................................ 48 3.3.1 Model Parameterization ................................................................................... 48 3.3.2 Thermal Habitat Use .......................................................................................... 49 3.4 Conclusion ................................................................................................................. 53 iii Chapter 4 Model the Natural Terrain Elevation of Lizard Micro habitat ................................................................................... 55 4.1 Introduction ............................................................................................................. 57 4.2 Methods .....................................................................................................................