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A Molecular Phylogeny of the Caribbean Subtribe

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A Molecular Phylogeny of the Caribbean Subtribe RESOLVING GENERIC BOUNDARIES IN RHYTIDOPHYLLUM AND GESNERIA: A MOLECULAR PHYLOGENY OF THE CARIBBEAN SUBTRIBE GESNERIINAE (GESNERIACEAE) by DAVID REECE WATSON JOHN L. CLARK, COMMITTEE CHAIR MARTHA POWELL LAURA REED SIMON JOLY A THESIS Submitted in partial fulfillment of the requirements for the degree of Master of Science in the Department of Biological Sciences in the Graduate School of The University of Alabama TUSCALOOSA, ALABAMA 2015 Copyright David Reece Watson 2015 ALL RIGHTS RESERVED Abstract The Caribbean subtribe Geseneriinae is comprised of 4 genera consisting of approximately 91 species. Most species are in Gesneria and Rhytidophyllum and fewer than five are currently recognized in Bellonia and Pheidoncarpa. Hispaniola is the most diverse region with 38 species followed by Cuba with 35 species. The Gesnerieae is nearly endemic to the Caribbean with only two species located in Venezuela and Colombia. Molecular sequence data from chloroplast (matKR) and nuclear markers (ITS and ETS) were used to generate a phylogenetic tree for evaluating the generic circumscription of Gesneria and Rhytidophyllum. Results from this study strongly support the monophyly of Rhytidophyllum and Gesneria. Phylogenetic results support and update classification for the circumscriptions of species previously recognized as Gesneria purpurascens and G. duchartreoides. Results also support the need for a re-evaluation of taxa currently recognized as Gesneria viridiflora complex. ii Dedication This thesis is dedicated to my grandparents. The wisdom they shared with me growing up helped lay the foundation for me to pursue my goal. It is their guidance and support that helped me to reach this point. It is in their memory that I endeavor to reach my academic and personal goals. iii Acknowledgements I would like to acknowledge all of the members of the faculty of the University of Alabama, colleagues in the lab and friends who have helped guide me in preparing this thesis. I would also like to acknowledge my committee members Dr. Laura Reed, Dr. Martha Powell, and Dr. Simon Joly whose guidance and instruction in performing this research was invaluable. Most notably I would like to acknowledge Dr. John L. Clark. This project would not have been possible without his expertise and belief in me. I would also like to acknowledge all of the members of the Clark Lab who have aided in my research and provided support. Laura Clavijo, Jason Martin, and Jon Grammar all provided invaluable assistance in lab work and support during analysis and editing of this study. This research was made possible by the support of the University of Alabama, the Graduate School, the Department of Biological Sciences, my friends, and family. I appreciate everyone who aided me during this process and the encouragement that helped make the completion of this project and thesis possible. iv Contents Abstract ........................................................................................................................................... ii Dedication ...................................................................................................................................... iii Acknowledgements ........................................................................................................................ iv List of Tables ................................................................................................................................ vii List of Figures .............................................................................................................................. viii Introduction ......................................................................................................................................1 Materials and Methods .....................................................................................................................4 Taxon Sampling ......................................................................................................................4 DNA Extraction, Amplification and Sequencing ...................................................................6 Alignment and Phylogenetic Analyses ...................................................................................7 Results ..............................................................................................................................................9 DNA Sequencing and Alignment ...........................................................................................9 Phylogenetic Analyses ............................................................................................................9 Discussion ......................................................................................................................................13 Comparison to Previous Phylogenies ...................................................................................13 Resolving Generic Boundaries .............................................................................................13 Future Directions ..................................................................................................................15 References ......................................................................................................................................16 v List of Tables 1. Specimens in Analysis .........................................................................................................4 vi List of Figures 1. Maximum Likelihood Consensus Tree ..............................................................................10 2. Bayesian Consensus Tree ..................................................................................................11 3. Strict Consensus Tree Based on Maximum Parsimony Analysis ......................................12 4. Examination of the characters of Gesneria fruticosa ........................................................14 vii Introduction The subtribe Gesneriinae is a monophyletic lineage that has diversified in the Caribbean and is comprised of approximately 65 species of Gesneria, 26 species of Rhytidophyllum, two species of Bellonia, and two species of Pheidonocarpa. The majority of species are found on the islands of Cuba and Hispaniola with each having approximately 30 species (Skog 2012). The only other islands with more than five species are Puerto Rico and Jamaica. All other islands are limited to fewer than five species and most have only one or two (Skog 2012). Only two species are non-Caribbean and they are located along the coasts of Colombia and Venezuela. These two species are: Rhytidophyllum onacaense (Rusby) L.E. Skog (mostly found in Colombia) and Rhytidophyllum cumanense (Hanst.) L.E. Skog,(mostly found in Venezuela). The subtribe Gesneriinae as set forth by Linnaeus lacked sufficient definition for the circumscription of genera and thus many new species were incorrectly placed within the tribe. The group was treated as a single genus by Linnaeus (1753) and Swartz (1788) and then divided into two or more genera Martius (1829), Decaisne (1846), Hanstein (1854) and others, losing the name Gesneria L. in doing so. Most notably Kuntze (1891) resurrected the name Gesneria L. while synonymizing Rhytidophyllum while Fritsch (1893-94) reverted back to the system of Hanstein and Bentham, dividing the tribe back into Gesneria and Rhytidophyllum. Urban focused his work on Caribbean plants and as a result he described more than 60 species of Gesneria and Rhytidophyllum, which is more than any other botanist. Urban (1901) followed the classifications of Bentham (1876) and Fritsch (1894 lending support to the separation of Gesneria and Rhytidophyllum. 1 Recent generic boundaries in the tribe Gesnerieae are confusing because Wiehler (1983) and Skog (1976) had different generic concepts for circumscribing Rhytidophyllum and Gesneria. Wiehler recognized Rhytidophyllum as a synonym of Gesneria and therefore combined all Caribbean members of the family into Gesneria. In contrast, Skog recognized Gesneria and Rhytidophyllum as separate genera by the presence of trichomes on the stamens as well as adnation of the stamens to the corolla tube. In addition to Rhytidophyllum and Gesneria, Skog (1976) described Pheidonocarpa as a new genus. Rhytidophyllum are distinguished from Gesneria by large bullate or areolate leaves, campanulate flowers, isolated stomata raised on domes, undulate cell margins, stamens adnate to the base of the corolla, pubescence on the stamen filaments, and maroon to green flowers (Skog 1976). However, no single morphological character can be used to distinguish all species in one genus from another and several of the defining characteristics are only present on the corolla. Some species of Rhytidophyllum have red tubular flowers and have thus been historically classified as Gesneria but they exhibit adnation of the stamens at the base of the corolla tube and pubescence on the stamens which is more similar to Rhytidophyllum. Wiehler (1983) recognized that no single character could be used to differentiate between the entireties of each genus. However, the confusion over these morphological differences and the presence of species displaying intermediate characteristics led him to argue that the diversity is indicative of environmental adaptations of individual species and does not represent generic delimitation. Wiehler interpreted the adnation of the filaments to the corolla tube as minor differences that were only useful for recognizing
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