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And Chemistry-Based Species Circumspections in Lichenized Brigham Young University BYU ScholarsArchive All Theses and Dissertations 2010-07-02 Assessing Traditional Morphology- and Chemistry-Based Species Circumspections in Lichenized Ascomycetes: Character Evolution and Molecular Species Delimitation in Common Western North American Lichens Steven Leavitt Brigham Young University - Provo Follow this and additional works at: https://scholarsarchive.byu.edu/etd Part of the Biology Commons BYU ScholarsArchive Citation Leavitt, Steven, "Assessing Traditional Morphology- and Chemistry-Based Species Circumspections in Lichenized Ascomycetes: Character Evolution and Molecular Species Delimitation in Common Western North American Lichens" (2010). All Theses and Dissertations. 2191. https://scholarsarchive.byu.edu/etd/2191 This Dissertation is brought to you for free and open access by BYU ScholarsArchive. It has been accepted for inclusion in All Theses and Dissertations by an authorized administrator of BYU ScholarsArchive. For more information, please contact [email protected], [email protected]. Assessing traditional morphology- and chemistry-based species circumspections in lichenized ascomycetes: character evolution and species delimitation in common western North American lichens Steven D. Leavitt A dissertation submitted to the faculty of Brigham Young University in partial fulfillment of the requirements for the degree of Doctor of Philosophy Larry L. St. Clair, Chair Byron J. Adams Leigh A. Johnson Roger Rosentreter Jack W. Sites, Jr. Department of Biology Brigham Young University August 2010 Copyright © 2010 Steven D. Leavitt All Rights Reserved ABSTRACT Assessing traditional morphology- and chemistry-based species circumspections in lichenized ascomycetes: character evolution and species delimitation in common western North American lichens Steven D. Leavitt Department of Biology Doctor of Philosophy Accurate species delimitation has critical implications for ecological and conservation studies; and for understanding factors driving diversification. However, a growing body of evidence indicates that morphology-based species circumspection in lichenized ascomycetes often fails to accurately represent the number of fungal species. The use of molecular data in lichen systematics provides an important alternative to traditional morphological characters for identifying natural groups and assessing evolutionary histories in challenging lichen taxa. In this work, I examined two common lichen-forming genera in western North America, Rhizoplaca and Xanthoparmelia, as models for investigating character evolution, species delimitation in morphologically and chemically diverse species, and identification of lineages in the early stages of divergence. Phylogenetic hypotheses were reconstructed to assess character evolution using sequence data from four nuclear ribosomal markers and fragments from two nuclear loci. I applied a multifaceted approach to delimit species in Rhizoplaca and Xanthoparmelia b y assembling multiple lines of evidence using DNA sequence data, and genealogical and population genetic analyses. I have found that traditionally circumscribed species are not supported by molecular data. For example, in Rhizoplaca previously unrecognized lineages were identified within what has thus far been considered a single species. In contrast, morphologically and chemically distinct species within Xanthoparmelia were not supported by molecular data. Distinct medullary chemistries, growth forms, and the production of vegetative diaspores appear to have evolved independently multiple times in Xanthoparmelia. This work clearly indicates that morphological and chemical characters do not always accurately reflect lichen species diversity within even the best known and studied genera. My study of the Rhizoplaca melanophthalma species complex demonstrates that the genus Rhizoplaca, as presently circumscribed, is more diverse in western North American than previously thought. I present these analyses as a working example of species delimitation in morphologically cryptic lichenized fungi. In Xanthoparmelia diagnostic morphological and chemical characters have evolved in a highly homoplasious manner. In contrast to other studies documenting previously undiscovered fungal lineages masked within lichen species circumscribed by traditional morphological and chemical characters, my work suggests that species diversity has been overestimated in the lichen genus Xanthoparmelia. Keywords: character evolution, convergence, lichens, morphology, Parmeliaceae, Rhizoplaca, secondary metabolites, speciation, species concepts, species delimitation, vagrant lichens, Xanthoparmelia ACKNOWLEDGMENTS I wish to thank my graduate committee for their guidance throughout this work. I recognize and sincerely appreciate the investment of time and assistance generously given throughout the course of this research. Byron Adams provided invaluable help and feedback on early versions that vastly improved my writing. Leigh Johnson has been especially generous with funding, work space and equipment, technical advice, invaluable feedback, and support throughout the entire project; and I sincerely feel that due to his kindness I’ve been able to successfully complete this dissertation. Roger Rosentreter has been incredibly helpful with his vast knowledge of vagrant lichens, and his enthusiasm for my research always came at the most opportune times. Jack Sites also provided invaluable feedback and insights on early versions, expertise in understanding species delimitation, and important conceptual help. Larry St. Clair provided incredible opportunities, liberty, generosity, kindness, support, friendship, and a great example. He is everything that a major advisor should be. I am truly indebted to many great people who made significant contributions to the success of this dissertation research. I express heartfelt thanks to colleagues, friends, and family who collected or contributed specimens for this project: Anna Bennett, Curtis Björk, Stuart Crawford, Bernard de Vries, Mike DeVito, Bob Egan, Ted Esslinger, Roy Fuller, Lawrence Glacy, Teegan Hardle, Steve Hardle, Brenda Hardle, J. Hertz, Derek Howell, Donna Howell, Chris Howell, Mike Felix, Trevor Goward, Melinda Greenwood, Jason Hollinger, Katy Knight, Adele Leavitt, Daniel Leavitt, Dean Leavitt, Don Leavitt, Griffin Leavitt, Hailey Leavitt, Jackson Leavitt, James Leavitt, Wayne Leavitt, Garrat Lind, Bruce McCune, Jenifer Munsha, Mark Robinson, Roger Rosentreter, Gajendra Shrestha, the Starkeys, Larry St. Clair, and Tim Wheeler. I thank Trevor Goward for his support, enthusiasm, discussion, and friendship; Jesse Brienholt for timely help with data analyses; Eric Green and Gajendra Shrestha for valuable discussion; and Christopher Jones, LauraDawn Leavitt, and Peter Ririe for help in the lab. My family has been provided stability, love, and encouragement. Specifically, my parents, Don and Adele Leavitt and Chris and Donna Howell, have encouraged, promoted, and facilitated the completion of this research. The Fullers, Howells, Starkeys, and Leavitts have all been to kind enough to participate in lichen catching forays. My brother, Dean Leavitt, provided consistent support and guidance throughout this work. I acknowledge the incredible contributions of my wife, Hailey Leavitt. She certainly has sacrificed more than anyone else throughout the course of this work. She has been patient, loving, supportive, and always made things work out even when it seemed impossible. My two children, Jack and Griffin Leavitt, provided incredible love and acceptance. I deeply treasure the memories of the love and support from my family that was so apparent during the course of this work. These studies were supported by the California Lichen Society, The Ruth L. Glacy Foundation, Walmart, Imke Schmit at the University of Minnesota, Brigham Young University graduate mentoring, graduate research fellowship awards, and the Brigham Young University Office of Research and Creative Activities. TABLE OF CONTENTS LIST OF TABLES ..................................................................................................................... viii LIST OF FIGURES ..................................................................................................................... ix LIST OF SUPPLEMENTARY DATA....................................................................................... ix CHAPTER ONE: Complex patterns of speciation in cosmopolitan “rock posy” lichens - an integrative approach to discovering and delimiting fungal species in the lichen- forming Rhizoplaca melanophthalma species-complex (Lecanoraceae, Ascomycota) ... 1 Abstract ........................................................................................................................... 2 Introduction ..................................................................................................................... 3 Materials and Methods .................................................................................................... 7 Results …………………………………………………………………………… ..... .17 Discussion ..................................................................................................................... 23 Conclusions ................................................................................................................... 30 Acknowledgements ....................................................................................................... 31 Literature Cited ............................................................................................................
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