SYSTEMATICS AND EVOLUTION OF COMMIPHORA JACQ. (BURSERACEAE) IN MADAGASCAR by Morgan Robert Gostel A Dissertation Submitted to the Graduate Faculty of George Mason University in Partial Fulfillment of The Requirements for the Degree of Doctor of Philosophy Environmental Science and Public Policy Committee: _________________________________________ Dr. Andrea Weeks, Dissertation Director _________________________________________ Dr. Cody Edwards, Committee Member _________________________________________ Dr. James Lawrey, Committee Member _________________________________________ Dr. Donald Seto, Committee Member _________________________________________ Dr. Albert Torzilli, Graduate Program Director _________________________________________ Dr. Robert Jonas, Department Chairperson __________________________________ Dr. Donna Fox, Associate Dean, Student Affairs & Special Programs, College of Science _________________________________________ Dr. Peggy Agouris, Dean, College of Science Date: __________________________________ Summer Semester 2015 George Mason University Fairfax, VA Systematics and Evolution of Commiphora Jacq. (Burseraceae) in Madagascar A dissertation submitted in partial fulfillment of the requirements for the Doctor of Philosophy at George Mason University By Morgan Robert Gostel Master of Science Virginia Commonwealth University, 2010 Bachelor of Science Virginia Commonwealth University, 2008 Director: Andrea Weeks, Associate Professor Department of Biology Summer Semester 2015 George Mason University Fairfax, VA Acknowledgements I owe my deepest gratitude to so many for their support, both emotional and financial, and the ability to begin and complete this dissertation. The genealogy of this work extends quite far and begins of course, as all living things, with my family for their encouragement of curiosity, learning, and appreciation for the beauty in nature. I was first inspired to pursue a career in biological research by my friend and mentor, Dr. Peter deFur, whose levity of spirit, compassion, and earnestness is a reminder of what is most important in life. My interest in botany began as an undergraduate student at Virginia Commonwealth University in the laboratory of Dr. Gregory M. Plunkett, who first offered me the opportunity to pursue graduate research. My Masters research gave way to the opportunity to work with Dr. Andrea Weeks, who has been my lifeline, a rock, and the most supportive advisor I could ever imagine. My deepest thanks to Dr. Weeks for taking me on as a Ph.D. student and giving me the opportunity to study Commiphora, for the many late night and weekend responses to emails and edits, and for not giving up on me or my work and encouraging me to do the same. At George Mason University (GMU) I would like to thank the rest of my committee, Drs. James Lawrey, Cody Edwards, and Don Seto for their support and guidance on my project and recommendations along the way. I am especially grateful to Dr. Edwards for the opportunity to work as a graduate research assistant in the Office of the Provost during the last year of my study, and thus expanding my interest in education. I owe much thanks to the Department of Environmental Science and Policy Executive Committee, especially Dr. Albert Torzilli, Sharon Bloomquist, Roslyn Cress, Joanne Zimmerman, and Susan Cheselka, for administrative support, financial support through teaching assistantships and two research fellowships that greatly improved my dissertation research, and ongoing assistance throughout the five years of my study at GMU. I also extend my thanks Dr. Reid Schwebach for offering guidance and assistance toward my pedagogical development and making me a better teacher. I thank members of my wider professional network, who have supported me throughout the years, sometimes simply by ‘being there’ and other times for reminding me of my part in the ‘bigger picture’. I especially thank the Botanical Society of America and Bill Dahl for inviting me to become involved in the BSA. Immense thanks to the many organizations that have helped to fund this research, including the National Science Foundation, the American Society of Plant Taxonomists, and the GMU Executive Committee. I am grateful for the help of colleagues throughout the course of this work, especially Pete Phillipson and Jeannie Raharimampionona, whose expertise and familiarity with the Malagasy Commiphora was of incomparable aide to this project. I hope to spend many more hours in the herbarium with you both! Many thanks to Dr. Pete ii Lowry II, Richard Razakamalala, and Charles Rakotovao at the Missouri Botanical Garden and Jackie Andriantiana and Hanta Razafindraibe at PBZT in Madagascar, without whom my collections in Madagascar would not have been possible. I could not ask for better company in the field and I hope to see you all soon! I give thanks to my friends, old and new, near and far, who have helped make this possible. This work is the culmination of a series of life events, good, bad, and everything in between. You’ve been there with me all along, but thanks especially to JD, Nate, Critter and Stephy. Thank you for being there and giving me support when I needed it, I love you! Thanks to my family, Gostel and Dal Forno, for their love and support over the years. John, Mary, Laura, and Robbie, you have each been rockstars! Thanks for being here for Manu and me, you have helped make Fairfax much more of a home for us. Obrigrado, Silvani, Luis, Anna, e Lara para todo seu amor e apoio, amo vocês! Most importantly I thank my wife, Manu, who brings happiness, love, and meaning to every day in my life. All of you have played a role in one of the most important chapters in my life so far, thank you for reminding me that I will never stop learning and for giving me the chance to continue doing so. iii Table of Contents Page List of Tables .................................................................................................................... vi List of Figures .................................................................................................................. vii List of Appendices ............................................................................................................ ix Abstract .............................................................................................................................. x Chapter 1: Systematics of the myrrh genus, Commiphora, globally and in Madagascar, 1797–present ............................................................................................... 1 Introduction ....................................................................................................................... 1 References .......................................................................................................................... 5 Chapter 2: Phylogenetic reconstruction of the myrrh genus, Commiphora (Burseraceae), reveals multiple radiations in Madagascar and clarifies infrageneric relationships....................................................................................................................... 7 Abstract .............................................................................................................................. 7 Introduction ....................................................................................................................... 8 Commiphora in Madagascar .......................................................................................... 11 Objectives and Hypotheses............................................................................................. 12 Materials and Methods ................................................................................................... 12 Taxonomic Sampling ..................................................................................................... 12 DNA Extraction and Molecular Sampling .................................................................... 14 Phylogenetic Inference .................................................................................................. 15 Divergence Dating Analysis .......................................................................................... 18 Results .............................................................................................................................. 19 Sequence Data ............................................................................................................... 19 Discussion......................................................................................................................... 22 Monophyly of Commiphora and its Taxonomic Sections ............................................. 22 Species Evolution of Commiphora in Madagascar ....................................................... 23 Dating the Divergence of Commiphora......................................................................... 24 Well-supported Lineages within Commiphora .............................................................. 25 Conclusions ...................................................................................................................... 30 References ........................................................................................................................ 31 Chapter 3: Development of novel EPIC markers from EST databases and evaluating their phylogenetic utility in Commiphora (Burseraceae) .......................... 48 Abstract ...........................................................................................................................