A Monograph of Usnea from Sao Tome and Principe A thesis submitted to the faculty of % San Francisco State University MS In partial fulfillment of E><0L the requirements for the Degree • U 33 Master of Science In Biology: Ecology and Systematic Biology by Miko Rain Abraham Nadel San Francisco, California December 2015 Copyright by Miko Rain Abraham Nadel 2015 CERTIFICATION OF APPROVAL I certify that I have read Monograph of Usnea from Sao Tome and Principe by Miko Rain Abraham Nadel, and that in my opinion this work meets the criteria for approving a thesis submitted in partial fulfillment of the requirement for the degree Master of Science in Biology: Ecology and Systematic Biology at San Francisco State University. DennisE. DesL6rdin, Ph.D. Professor ofjsiology San Francisco State University Jose R. de la Torre, Ph.D. Associate Professor of Biology San Francisco State University Robert C. Drewes, Ph.D. Curator of Herpetology, Emeritus California Academy of Sciences A Monograph of Usnea from Sao Tome and Principe Miko Rain Abraham Nadel San Francisco, California 2015 A systematic monograph of the lichen genus Usnea from the remote, under-explored islands of Sao Tome and Principe (ST&P) in tropical West Africa is presented, treating 11 species representing 2 subgenera, Usnea and Eumitria. Ten of the 11 species are reported as new for ST&P, including U. firmula, U. baileyi, U. pectinata, U. aff. flammea, U. sanguinea, U. picta, U. krogiana and three undetermined, potentially new species, Usnea species A, B and C. Usnea articulata is confirmed for the main island of Sao Tome. Determinations were made using morphological data, chemistry via thin layer chromatography, and a molecular phylogeny based on nDNA Internal Transcribed Spacer (ITS) sequences. Usnea baileyi, U. pectinata, U. articulata and Usnea species C represent species complexes consisting of two or more morphotypes each. Comprehensive descriptions, photographs, and a dichotomous key to the species are provided. correct representation of the content of this thesis De nnjs E. Desjarefm; Chair, Thesis Committee Date ACKNOWLEDGEMENTS I’d like to express my deepest appreciation to the many people who supported me on this journey of exploration and education. Thank you to Dr. Bob Drewes for the incredible opportunity to travel to such an amazing place for my collections and to those in ST&P who provided us with all the support in the field. Thanks also to Dr. Dennis Desjardin for agreeing to work with me despite my strange fascination with lichen, as well as Dr. Jose de la Torre for serving on my committee. Thank you to those who were able to give their time, experience and/or resources in helping me learn and become a better scientist, including: James Shevock, Doug Glavich, Dr. Bruce McCune, Dr. Frank Cipriano, Dr. Yoshihito Ohmura, Dr. Irwin Brodo, and Dr. Felix Schumm. I’m also immensely grateful for my father and the rest of my family, loved ones and friends and especially my grandmother, Ruth “Dolly” Nadel, who taught me to persevere and to believe in myself from a very young age when life was challenging. Thank you to all my colleagues at San Francisco State University for assistance along the way whether teaching me, or allowing me to learn by teaching you, every little bit helped. Lastly, thank you to the groups that granted me funds towards my research; The California Academy of Sciences, The California Lichen Society, The American Bryological and Lichenological Society, and The Mycological Society of San Francisco. v TABLE OF CONTENTS List of Table....................................................................................................................................... vii List of Figures....................................................................................................................................viii List of Appendices.............................................................................................................................. ix Introduction.......................................................................................................................................... 1 Usnea in Sao Tome and Principe......................................................................................... 2 Materials and Method..........................................................................................................................5 Literature Review..................................................................................................................5 Specimen and Exterior Data Collection................................................................................. 5 Morphological Analysis.......................................................................................................... 6 DNA Extraction, Amplification and Sequencing....................................................................7 Sequencing Data Analysis.................................................................................................... 8 Chemical Analysis..................................................................................................................9 Results............................................................................................................................................... 11 Phylogenetic Results............................................................................ 11 Subgenus Eumitria..................................................................................................12 Subgenus Dolichousnea.........................................................................................13 Subgenus Usnea.....................................................................................................13 Thin Layer Chromatography Results....................................................................................15 Taxonomy.................................................................................................................... 17 Key to species.........................................................................................................17 Species descriptions................................................................................................20 Usnea firmula............................................................................................ 20 Usnea baileyi............................................................................................. 22 Usnea pectinata........................................................................................ 24 Usnea aff. flammea................................................................................... 27 Usnea sanguinea...................................................................................... 28 Usnea species A ....................................................................................... 30 Usnea species B ....................................................................................... 32 Usnea articulata........................................................................................ 34 Usnea picta............................................................................................... 37 Usnea species C....................................................................................... 38 Usnea krogiana......................................................................................... 41 Discussion.................................................................................................................................. 43 References........................................................................................................................................66 Appendices........................................................................................................................................76 vii LIST OF PLATES Plate Page 1. Usneafirmula.................................................................................................................45 2. Usnea baileyi.................................................................................................................46 3. Usnea pectinata 1........................................................................................................47 4. Usnea pectinata 2 ........................................................................................................48 5. Usnea aff. flammea....................................................................................................... 49 6. Usnea sanguinea...........................................................................................................50 7. Usnea species A ........................................................................................................ 51 8. Usnea species B ........................................................................................................ 52 9. Usnea articulata.............................................................................................................53 10. Usnea picta....................................................................................................................54 11. Usnea species C............................................................................................................55 12. Usnea krogiana..............................................................................................................56 LIST OF FIGURES Figures Page 1. Usnea phylogeny .........................................................................................................57