ABSTRACT Title: THE EVOLUTION, PHYLOGEOGRAPHY, AND CONSERVATION OF THE GOLDEN LANGUR (Trachypithecus geei) IN BHUTAN Tashi Wangchuk, Doctor of Philosophy, 2005 Directed By: Professor David Inouye, Department of Biology The golden langur (Trachypithecus geei) is an endangered endemic species of primate in Bhutan. Conservation of this species is addressed here through phylogeographic and habitat management tools. I hypothesize that rivers and mountains in Bhutan isolated a population of capped langurs (Trachypithecus pileatus) and that this population later speciated into the morphologically distinct golden langur. Trachypithecus, the genus to which both capped and golden langurs belong, spread north from a paleo-refuge in south China and Semnopithecus (grey langurs) spread east and northward from a refuge in south India. My results show that these two genera both arrived in Bhutan but could not mix since the Sunkosh River and Pelela range form a biogeographic barrier. Likewise, a population of capped langurs isolated from parental populations by rivers speciated into the distinct golden langur. I conducted field surveys covering the entire range of langurs in Bhutan, confirming the distribution and isolating barriers among the three langurs. Grey langurs and golden langurs are isolated from each other by the Sunkosh River and Black Mountain range in west Bhutan. In the east, the Manas river system (Manas- Mangde-Chamkhar) served as a barrier between golden and capped langurs. However, this barrier has been broken in the last 30 years due to the construction of bridges over the Chamkhar river. A hybrid zone was found and the implications are discussed. A cyt b phylogeny showed the grey langur of Bhutan grouping into a distinct clade with other congenerics of Semnopithecus. The south-Indian clade of grey langurs is more ancient, with the Bhutan and Nepal grey langurs having diverged later. The golden and capped langur from Bhutan grouped with Trachypithecus from South East Asia. Finally, I explore conservation of golden langur habitat in Bhutan and estimate available habitat at 3,089 km2 and an estimated population of about 6,000 individuals. I also find that the most viable strategy for conservation of langur habitat is to give ownership of the forests to local people, with monitoring by the Department of Forestry. THE EVOLUTION, PHYLOGEOGRAPHY, AND CONSERVATION OF THE GOLDEN LANGUR (Trachypithecus geei) IN BHUTAN By Tashi Wangchuk Dissertation submitted to the Faculty of the Graduate School of the University of Maryland, College Park, in partial fulfillment of the requirements for the degree of Doctor of Philosophy 2005 Advisory Committee: Dr. David Inouye, Chair Dr. James Dietz Dr. Matthew P. Hare Dr. Charles Delwiche Dr. Jonathan Ballou © Copyright by Tashi Wangchuk 2005 Dedication To the memory of my beloved grandmother Angay Thinley Bidha of Wang-Simu. ii Acknowledgements I am grateful to Professor David Inouye for his support and inspiration throughout the entire process of my graduate work at Maryland. I write this after having successfully defended my dissertation on 31 March 2005. David readily responded to all my requests and needs and guided me with gentle persuasion and unconditional support, a trademark uniquely David’s and a blessing for all his graduate students. I am also grateful to the rest of my committee members, Dr. Matt Hare for guiding me through the phylogenetic work in his lab and for allowing me full access to his lab. Dr. Chuck Delwiche for teaching me molecular systematics and taking time to scrutinize my data. Dr. Jon Ballou for traveling with me to my field sites and reading the 300 page dissertation and thoughtful comments and suggestions on the dissertation. Dr. Jim Dietz for teaching me the basic concepts of Conservation Biology back in 1992 when I was an undergraduate and for placing my dissertation within the framework of Conservation Biology. Dr. Eric Haag provided not only a place for me to stay during the crucial stages of writing the final parts of the dissertation but also a supportive environment and great conversations, meals, and home brewed bear. Thinley Namgyel, compatriot and graduate student in the Conservation Biology and Sustainable Development Program readily participated in all of the festivities. Eric and Thinley provided much needed friendship and distractions from the rigor and worry of preparing for the defense. Dr. Doug Gill provided a place to stay in 2003 during a visit for lab work. Many ecological theories were analyzed. Some we failed to reject. Others we rejected outright - all over a bowl of spicy oxtail stew at his dining table. Dr. Kerry Shaw’s iii seminar on Species and Speciation provided deep insights for someone grappling with species concepts. Lois Reid negotiated the university bureaucracy with ease and always ensured that all my paperwork was in order. Gary Dodge, lab mate and fellow dissertation writer, took time to answer my many questions on statistics despite the pressure of preparing for his own defense two weeks after mine. Colin Rose kindly taught me to use Modeltest. I am also grateful to Peter and Colleen Nicholas for providing a warm and welcoming home for me and my family for over a year in their College Park home in 2000 and 2001, otherwise known also as Bhutan House. They opened their home to all manner of Bhutanese, grateful for a home away from home. The field research would not have been possible without the help of many people in the Department of Forests in Bhutan. Sangay Dorji, Phuntso Thinley, Phurba, Kinley Gyeltsen, Kinga Deki, Kinzang Gyeltsen, Dorji Phuntso, and Kezang Wangchuk provided critical support during the field surveys. The people of Zhemgang and Trongsa willing carried field gear from campsite to campsite, answered numerous questions on land use and taught me a great deal about the land use practices. I am also grateful to the University of Maryland for teaching assistantships, the Bhutan Trust Fund for Environmental Conservation for a research grant, the Royal Government of Bhutan for a free education right up to the Master’s level, and David Hulse for helping with a roundtrip airfare between Bhutan and College Park. Dr. Yoshi Kawamoto and Dr. Mitsuru Aimi of the Primate Research Institute of Kyoto University provided advice and invited me to do some of the lab work in Inuyama. Lastly, my wife Kelzang Lhaden, daughters Thinley Bidha, Megom Tsogyel, and iv Tashi Lham translocated half way across the world without complaint and bravely went to public school in America. Tashi Lham wasn’t quite so sure about the big yellow bus on the first day of her kindergarten in 2001. v Table of Contents Dedication..................................................................................................................... ii Acknowledgements......................................................................................................iii Table of Contents......................................................................................................... vi List of Tables ............................................................................................................... ix List of Figures............................................................................................................... x Dissertation Summary................................................................................................... 1 Chapter 1: Evolution and Phylogeography of the langurs of Bhutan ........................... 8 Introduction: Non-human Primates of Bhutan.......................................................... 8 Ecological and Geophysical Background of Study Area.................................... 15 Ecological Zones and Species Distribution ........................................................ 17 Hypotheses about the phylogeny of the langurs ................................................. 20 Fossil history of the Colobines ........................................................................... 26 Phylogeny and Classification of the Asian Colobines........................................ 29 Methods................................................................................................................... 30 Sample Collection............................................................................................... 33 Mitochondrial DNA............................................................................................ 34 Lab methods for DNA analysis........................................................................... 35 DNA Amplification and Sequencing (Cytochrome b)........................................ 36 Cytochrome b (Cyt b) Data Analyses ................................................................. 37 Divergence Times ............................................................................................... 43 Results..................................................................................................................... 46 Biogeography...................................................................................................... 46 Grey Langur Distribution.................................................................................... 46 Golden Langur Distribution................................................................................ 48 Capped Langur Distribution ............................................................................... 49 Phylogenetics .....................................................................................................