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Replace This with the Actual Title Using All Caps SYSTEMATICS OF ANTIGONON AND TROPICAL ERIOGONOIDEAE: PHYLOGENY, TAXONOMY, AND INVASION BIOLOGY A Dissertation Presented to the Faculty of the Graduate School of Cornell University In Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy by Janelle Marie Burke May 2011 © 2011 Janelle Marie Burke SYSTEMATICS OF ANTIGONON AND TROPICAL ERIOGONOIDEAE: PHYLOGENY, TAXONOMY, AND INVASION BIOLOGY Janelle Marie Burke, Ph. D. Cornell University 2011 The genera of Polygonaceae have historically been segregated into two subfamilies, Eriogonoideae and Polygonoideae, based on a few key morphological characters. Using ITS, morphology and five chloroplast markers, a phylogeny for Eriogonoideae was reconstructed, with an emphasis on sampling of the tropical genera. Results support the placement of nine of twelve woody, tropical genera within Eriogonoideae, where these genera form a paraphyletic assemblage giving rise to Eriogoneae (Eriogonum and allies). My work corroborates previous phylogenetic studies, and suggests a broader circumscription of Eriogonoideae. Also based on these results, I propose the resurrection of a third subfamily, Symmerioideae, in Polygonaceae, and propose two new tribes, Gymnopodieae and Leptogoneae, in Eriogonoideae. Within the subfamily, the genus Antigonon provides a systematic challenge. Although Antigonon is a small, easily-recognized genus, the boundaries of species within it have never been resolved satisfactorily. A taxonomic treatment for the genus is presented, based on morphology and molecular phylogenetic data from two chloroplast markers (psaI-accD, psbA-trnH ) and one nuclear marker (LFY , 2nd intron). Four species are described, and a new subspecies, Antigonon leptopus subsp. coccineum is proposed. Antigonon leptopus is also known as corallita, a pantropical invasive vine particularly problematic on islands. Spread of the invasive populations was investigated using data collected for this systematic project. Label data from herbarium specimens were used to map the spread of invasive A. leptopus over time. These data show the history of corallita introduced as a garden ornamental, with subsequent naturalization and escape. Molecular data from plastid DNA sequence were analyzed in a phylogenetic context. Results suggest introductions outside the native range were not from one seed source, as exemplified by the genetic variation found within one island locality. The known data of biology, spread and control of corallita are reviewed and summarized in an action alert. BIOGRAPHICAL SKETCH Janelle Burke was born in St. Charles, Illinois on March 2, 1982 near the calm, yet moderately polluted waters of the Fox River. She spent the first 18 years of her life in the same town, frolicking through the parks of suburbia. She graduated from St. Charles High School in 2000. She then completed her bachelor of arts at the Johns Hopkins University majoring in behavioral biology in 2004, yet she managed to avoid the pre-med track, and is currently pursuing her Ph.D. in Plant Biology at Cornell University. When not pondering broad evolutionary quandaries, she enjoys eating spicy food and knitting; preferably not at the same time. iii To Dr Mom, for always setting a good example iv ACKNOWLEDGMENTS I am greatly indebted to so many people who assisted me during my dissertation work. I will start with technical support: I would like to thank everyone who helped me collect plants in the field. Gonzalo Castillo, Sergio Avendaño and Jesus Pale at the Instituto do Ecología, Xalapa helped me travel to Veracruz and Oaxaca, as well as offered their warm hospitality. Rodrigo Duno and German Carnevalí accompanied me around the Yucatán, as well as providing access to gastronomic delights. I am very grateful for their help and friendship. Sirli Leython and Severo Romero made so many arrangements and were great hosts in Venezuela. Lucia Vazquez, Patricia Hernandez and James Cohen helped arrange travel in Michoacán and Guerrero, and providing great companionship. Nicole Esteban and Lisa Driver were great help in gathering data on St. Eustatius. Numerous herbaria support staff have assisted me, and facilitated my study of loans and specimens. First and foremost I would like to thank Robert Dirig, Peter Fraissinet and Anna Stalter at the Bailey Hortorium. They were always willing to assist, and have helped tremendously with my herbarium work. Rusty Russell and Deborah Bell have been very accommodating to my frequent visits to the Smithsonian Institution. I would especially like to thank Stan Yankowski for helping me image specimens to use for publication. For financial support, I would like to thank the Department of Plant Biology for fellowships and teaching assistantships, as well as the Department of Neurobiology for a teaching assistantship. I would also like to thank the Hal Moore Funds through the Dept. of Plant Biology, the American Society of Plant Taxonomists and the Botanical Society of America for research grants. My collaborators with the Kron Lab at Wake Forest have helped me not only with data collection, but also with formulating new ideas and making progress in Polygonaceae systematics. Dr. Adriana Sanchez has been a great person to help move my research forward. Dr. Sang-Hun Oh is a wonderful person to have in the lab. He has provided invaluable assistance in v molecular techniques and advice. And of course, I would like to thank all my friends who have given me emotional and/or scientific support and provided many, many laughs and meals over the years. To my original housemates, Hanh Nguyen and Catherine Martel, with whom I have enjoyed spending so much time. All of the Bailey Hortorium graduate students were phenomenal office mates and cohorts. In particular I would like to thank my lab mate Cindy Skema and my “eternal TA” Mariana Yazbek for their friendship and mentorship. Fronny Plume, Jim Cohen, Shannon Straub, Marcela Martinez-Millan, Caroline Kellogg, Mary Futey, Simon Gunner, Gwynne Lim, Tee Havanada, Dan Ilut, and Jeremy Coate: you are great friends and office mates. I consider myself fortunate to have had an outstanding committee. Dr. Jerrold Davis was always available to listen to my questions, and offered thoughtful feedback. I have especially enjoyed his lectures, sense of humor, and academic advice. Dr. James Reveal is a tremendous resource for all things Polygonaceae, and was willing to review any document I sent his way. Dr. Antonio DiTommaso has helped by encouraging me to publish and present in a field that is not my primary focus. I appreciate his enthusiasm and timely feedback. Dr. Melissa Luckow, my major professor, has been a true mentor and friend. From the time I was a young graduate student looking for a thesis project and a free trip to Venezuela, she has always been available for advice and support. I have learned from her rigid scholarship and I will always appreciate her academic guidance. I would like to thank my family, who has always ready and willing to visit me in Ithaca, even during bad weather. My father, James Burke, was a great help in helping generate distribution maps. And lastly, I want to thank Ahmad Slaibi, who has provided unwavering support and encouragement. ﺑﺤﺒﻚ ﻛﺘﻴﺮ. ﺍﻧﺖ ﻋﻴﻮﻧﻲ. vi TABLE OF CONTENTS Biographical sketch .................................................................................................................. iii Acknowledgements ....................................................................................................................v List of Tables ............................................................................................................................ ix List of Figures ............................................................................................................................x 1. Introduction .................................................................................................................................1 2. Placing the woody tropical genera of Polygonaceae: A hypothesis of character evolution and phylogeny .................................................................................................................................7 3. Revised subfamily classification for Polygonaceae, with a tribal classification for Eriogonoideae .........................................................................................................................58 4. Monograph of Antigonon ..........................................................................................................82 5. Corallita (Antigonon leptopus): Intentional introduction of a plant with documented invasive capability ..............................................................................................................................165 6. Integrating systematics and invasion biology: A case study of the pantropical perennial vine Antigonon leptopus (corallita) ..............................................................................................187 7. Conclusions .............................................................................................................................211 vii LIST OF TABLES 2.1 Traditional tribal circumscription of Coccolobeae and Triplarideae ......................................10 2.2 Table depicting characters used by various workers to either segregate woody tropical genera from the rest of Polygonaceae, or to distinguish tribes from each other ....................12 2.3 Statistics for gene regions used in the phylogenetic analysis.
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