REVISION OF PASSIFLORA SUBGENUS DECALOBA SUPERSECTION CIECA (PASSIFLORACEAE) By KRISTEN E. PORTER-UTLEY A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY UNIVERSITY OF FLORIDA 2003 Copyright 2003 by Kristen E. Porter-Utley ACKNOWLEDGMENTS This study was supported by grants from the National Science Foundataion (Doctoral Dissertation Improvement Grant, DEB-0104824), the American Society of Plant Taxonomists (Research Grants for Graduate Students), the Passiflora Society International (Graduate Research Award), Sigma Xi (Grant-in-Aid of Research), and the Haitian Resource Development Foundation (Travel Award). Norris Williams also provided funds for DNA sequencing and allowed me unlimited access to his laboratory in the University of Florida Herbarium (FLAS) at the Florida Museum of Natural History. Mark Whitten spent countless hours in the DNA sequencing lab (FLAS) with me where he patiently taught me various molecular techniques and generously provided advice, and for that I am eternally grateful. Pam Soltis also allowed me access to her laboratory and I greatly appreciate all of her help. Windy Zomlefer kindly donated her time to teach me techniques of botanical illustration and her expert advice is greatly appreciated. Tom Emmel generously allowed me to use his greenhouse so that I could cultivate the species in my study group. Peter Jørgensen sent me digital photographs that he took of herbarium specimens at the Muséum National d'Histoire Naturelle, Paris, France (P), the Real Jardín Botánico, Madrid, Spain (MA), and the Universität Göttingen, Göttingen, Germany (GOET) that I would have otherwise not seen. Emil Kugler is a great historian of the Passifloraceae and kindly translated into English several old German publications for me. Axel iii Frank also kindly donated his time to help me with several German to English translations and sent me digital copies of old paintings of several species of supersection Cieca. Special thanks are due to Hugh Popenoe who has been a source of encouragement and support throughout my graduate career. Alexandra Paul accompanied me on two collecting trips and spent several long days helping me search for species of Passiflora. In addition, her wonderful family kindly allowed me to stay in their home during our trip to Haiti. Tracy Commock and Mr. E. Scott of the Institute of Jamaica were very knowledgeable and helpful during my field trip to Jamaica. George Proctor gave me advice on collecting localities and assisted me in finding several of the species in my study group while I was in Jamaica. Andrea Donaldson, of the Natural Resources Conservation Authority, assisted me in obtaining collecting permits in Jamaica. I am especially grateful to German Carnevali for helping me with collecting permits and giving me collecting advice during my trip to México. Demetria Mondragón- Chaparro was my assistant and traveling companion during my field trip to México and is a very dear friend. Her ecological knowledge and experience in traveling around México made for a flawless trip. Demetria's family also opened up their hearts and homes to me while I was México. I would also like to thank those who provided me with greenhouse plants and material for DNA extraction: Richard Abbott, Noel Atherton, Carolee Boyles, Ron Boender, Doug Goldman, David Hearn, Jean Jacques, Jutta Jedzig, Elma Kay, John MacDougal, Mario Posla, Tim Skimina, John Vanderplank, and Max Weigend. I am particularly grateful to Norris Williams and Kent Perkins for providing space at the University iv of Florida Herbarium, Florida Museum of Natural History, Gainesville, Florida, USA (FLAS) for my thousands of loaned herbarium specimens. I would like to thank the following institutions for sending herbarium specimens or providing digital images of herbarium specimens: Arnold Arboretum, Harvard University, Cambridge, Massachusetts, USA (A); University of Aarhus, Aarhus, Denmark (AAU); Botanischer Garten und Botanisches Museum Berlin-Dahlem, Zentraleinrichtung der Freien Universität Berlin, Berlin, Germany (B); The Natural History Museum, London, England, UK (BM); University of Copenhagen, Copenhagen, Denmark (C); California Academy of Sciences, San Francisco, California, USA (CAS); Centro de Investigación Científica de Yucatán, México (CICY); University of Cambridge, Cambridge, England, UK (CGE); Museo Nacional de Costa Rica, San José, Costa Rica (CR); Dudley Herbarium, San Francisco, California, USA (DS); Duke University, Durham, North Carolina, USA (DUKE); Economic Herbarium of Oakes Ames, Harvard University, Cambridge, Massachusetts, USA (ECON); Field Museum of Natural History, Chicago, Illinois, USA (F); Museo di Storia Naturale dell'Università, Firenze, Italy (FI); Conservatoire et Jardin botaniques de la Ville de Genève, Genève, Switzerland (G); Gray Herbarium, Harvard University, Cambridge, Massachusetts, USA (GH); Universidad de Antioquia, Antioquia, Medellín, Colombia (HUA); Jardín Botánico Nacional Dr. Rafael M. Moscoso, Santo Domingo, Dominican Republic (JBSD); Jepson Herbarium, University of California, Berkeley, California, USA (JEPS); Royal Botanic Garden, Kew, England, UK (K); Lundell Herbarium, University of Texas at Austin, Austin, Texas, USA (LL); Botanische Staatssammlung v München, München, Germany (M); Real Jardín Botánico, Madrid, Spain (MA); Universidad Nacional Autónoma de México, México City, Distrito Federal, México (MEXU); University of Minnesota, St. Paul, Minnesota, USA (MIN); Missouri Botanical Garden, St. Louis, Missouri, USA (MO); Ludwig-Maximilians- Universität, München, Germany (MSB); United States National Arboretum, USDA/ARS, Washington, District of Columbia, USA (NA); Tulane University, New Orleans, Louisiana, USA (NO); New York Botanical Garden, New York, New York, USA (NY); University of Oxford, Oxford, England, UK (OXF); Academy of Natural Sciences, Philadelphia, Pennsylvania, USA (PH); National Museum in Prague, Praha, Czech Republic (PR); Universidade Federal do Rio de Janeiro, Rio de Janeiro, Rio de Janeiro, Brazil (R); Swedish Museum of Natural History, Stockholm, Sweden (S); Universidad Nacional Autónoma de Honduras, Tegucigalpa, Honduras (TEFH); University of Texas at Austin, Austin, Texas, USA (TEX); University of the West Indies, Saint Augustine, Trinidad, Trinidad, and Tobago (TRIN); Nationaal Herbarium Nederland, Utrecht University Branch, Utrecht, Netherlands (U); University of California, Berkeley, California, USA (UC); Smithsonian Institution, Washington, District of Columbia, USA (US); Fundación Instituto Botánico de Venezuela Dr. Tobías Lasser, Caracas, Venezuela (VEN); University of Wisconsin, Madison, Wisconsin, USA (WIS); Instituto de Ecología, Xalapa, Veracruz, México (XAL). Special thanks are due, again, to all of my committee members, Walter Judd, John MacDougal, Hugh Popenoe, Pam Soltis, and Norris Williams. They have all been very supportive of my work and have kindly donated their time and vi resources to this project. Walter Judd has been a wonderful advisor throughout the course of this study. His door was always open and he constantly provided me with much needed support and encouragement. He is truly a wonderful teacher and I am very fortunate to have had such a great mentor during my dissertation work. I would especially like to express my gratitude to John MacDougal. John has played a critical role in this project from its inception. He introduced me to Passiflora, gave me free access to his abundant files pertaining to the Passifloraceae, constantly sent me information that has enhanced the quality of my work, spent countless hours on the telephone giving me advice, and he and his family provided me with a place to stay and support during visits to the Missouri Botanical Garden. I would like to thank my family for their love and support. My husband, Luke, has been a constant source of love, patience, and encouragement. My parents have continually believed in me and helped me in innumerable ways. My sister and her family have always been willing to open up their hearts and home to me. My brother, Bill, and John May have always been around whenever I needed reassurance. Jim and Dolores Utley have also given me a helping hand whenever they could. I am truly fortunate to have such a caring family. vii TABLE OF CONTENTS page ACKNOWLEDGMENTS .......................................................................................iii LIST OF TABLES ................................................................................................xii LIST OF FIGURES .............................................................................................xiv ABSTRACT ........................................................................................................xxi CHAPTER 1 INTRODUCTION.............................................................................................. 1 2 TAXONOMIC HISTORY................................................................................... 5 3 MATERIALS AND METHODS........................................................................ 16 Morphological Data Set................................................................................... 16 Molecular Data Set ......................................................................................... 23 Phylogenetic Search Strategies...................................................................... 25 Species Concepts ..........................................................................................