Download/List/Vascular.Asp; Accessed Jan
Total Page：16
File Type：pdf, Size：1020Kb
i PHYLOGENY, BIOGEOGRAPHY, AND REPRODUCTIVE BIOLOGY OF THE COSMOPOLITAN FLOWERING PLANT GENUS STELLARIA L. by MATHEW THOMAS SHARPLES B.A., University of Massachusetts, 2008 A dissertation submitted to the Faculty of the Graduate School of the University of Colorado in partial fulfillment of the requirement for the degree of Doctor of Philosophy Department of Ecology and Evolutionary Biology 2019 ii This dissertation entitled: Phylogeny, biogeography, and reproductive biology of the cosmopolitan flowering plant genus Stellaria L. written by Mathew Thomas Sharples has been approved for the Department of Ecology and Evolutionary Biology ________________________________ Dr. Erin A. Tripp ________________________________ Dr. Jeffry Mitton ________________________________ Dr. Mitchell McGlaughlin ________________________________ Dr. Stacey D. Smith ________________________________ Dr. William Bowman Date 4 November 2019 The final copy of this thesis has been examined by the signatories, and we find that both the content and the form meet acceptable presentation standards of scholarly work in the above mentioned discipline. iii Sharples, Mathew Thomas (Ph.D., Ecology and Evolutionary Biology) Phylogeny, biogeography, and reproductive biology of the cosmopolitan flowering plant genus Stellaria L. Dissertation directed by Associate Professor and COLO Herbarium Curator Dr. Erin A. Tripp The flowering plant genus Stellaria L. (Caryophyllaceae; the “starworts”) numbers around 112 species and exhibits a cosmopolitan distribution. To gain familiarity with the Caryophyllaceae and a broad number of temperate plant taxa overall in the field, I conducted a floristic inventory of the South San Juan Wilderness of southern Colorado, documenting 533 species of mostly native vascular plants present there, including five species of Stellaria. Research on Stellaria beyond Colorado was then conducted across five continents based on field and museum work, with the proximate goal of forming a comprehensive phylogenetic hypothesis of evolutionary relationships within this and other flowering plant lineages based on RADseq data. This research revealed the existence of five clades of a core genus Stellaria, offered clarification of taxonomic boundaries of starworts, provided evidence for resurrection of two plant genera related to Stellaria (Adenonema and Mesostemma), and also provided evidence for description of two new plant genera, Nubelaria and Rabelera of Eurasia. Circumscription of core Stellaria allowed for addressing of downstream evolutionary hypotheses. Namely, a well- sampled phylogeny of Stellaria enabled the biogeographical history of starworts to be reconstructed as well as the history of petal evolution. Analyses revealed a Miocene origin of Stellaria likely in northeastern Eurasia with myriad dispersals and radiations across the world subsequently, and also suggested widespread habitat and climate lability in the genus. Data also iv revealed numerous, recurrent losses of petals in unrelated starwort lineages, and a field experiment on the alpine tundra of Niwot Ridge (Boulder County, Colorado) suggested that these losses may be associated with transitions from outcrossing to self-fertilizing pollination systems. v ACKNOWLEDGMENTS This work could not have been completed without the assistance from a great many people and institutions. Curator of COLO Erin Tripp encouraged this work from Day One and therefore holds my utmost and foremost gratitude. The remainder of my Ph.D. committee comprised of professors William Bowman, Jeffry Mitton, Mitchell McGlaughlin, and Stacey Smith (as well as her lab members) provided extensive helpful feedback and other assistance throughout this dissertation, as did Nolan Kane. Stephanie Mayer served as both a teaching and life mentor throughout this dissertation. Early-era lab members Yi-Hsin Erica Tsai, Heather Stone, and Vanessa Díaz, as well as late-era lab members Louise Barton, Matthew Schreiber, Carly Anderson Stewart, Sophia Warsh, Justin Williams, and Yongbin Zhuang are particularly thanked for edifying discussions and other assistance over the years. Tyson Burch also provided early mentorship. A magnificent array of botanists and herbaria hosted and helped me throughout this work, including but not limited to ALTB (Alexander Shmakov, Sergei Smirnov), CANB (Cathy Miller), CAS (Tom Daniel, Rebecca Peters, Debra Trock), CHR (Ines Schönberger), COLO (Ryan Allen, Dina Clark, Tim Hogan), E (Leslie Scott), IBSC (Sunan Huang, Deng Yunfei), K (Marie Briggs), KUN, LE (Galina Konechnaya, Roman Ufimov, Marina Yarichina), MEL (Pina Milne), MO (Jim Solomon), NSK (Andrey Erst, Dmitri Shaulo), NY (Jackie Kallunki, James Lendemer), P (Vanessa Invernon), PE, UC (Kim Kersh), VBGI (Evgeny Boltenkov, Valentina Kalinkina), and VLA. Funding was generously provided by the American Society of Plant Taxonomists, the Botanical Society of America, the Department of Ecology and vi Evolutionary Biology at the University of Colorado (Boulder), a University of Colorado (Boulder) Cynthia H. Schultz Graduate School Small Grant, the University of Colorado Museum of Natural History, and the University of Colorado Undergraduate Research Opportunities Program. Finally, I wish to thank my parents and sister for being fully supportive of my endeavors across the years. vii CONTENTS INTRODUCTION................................................................................................................1 CHAPTER I. VASCULAR FLORA OF THE SOUTH SAN JUAN MOUNTAINS (COLORADO, U.S.A.): A FLORISTIC INVENTORY OF TWO SOUTHERN ROCKY MOUNTAINS SLOPES Introduction.......................................................................................3 Methods.............................................................................................22 Results and Discussion......................................................................24 Checklist of the Vascular Flora of the South San Juan Mountains...37 II. Part I: PHYLOGENETIC RELATIONSHIPS WITHIN AND DELIMITATION OF THE COSMOPOLITAN FLOWERING PLANT GENUS STELLARIA L. (CARYOPHYLLACEAE): CORE STARS AND FALLEN STARS Introduction......................................................................................99 Materials and Methods.....................................................................104 Results..............................................................................................116 Discussion........................................................................................126 Revised Taxonomic Concepts..........................................................141 Part II: TAXONOMIC OBSERVATIONS WITHIN STELLARIA L. (CARYOPHYLLACEAE): INSIGHTS FROM ECOLOGY, GEOGRAPHY, MORPHOLOGY, AND PHYLOGENY SUGGEST WIDESPREAD PARALLELISM IN STARWORTS AND ERODE PREVIOUS INFRAGENERIC CLASSIFICATIONS viii Introduction....................................................................................152 Materials and Methods...................................................................153 Results and Discussion...................................................................157 III. BIOGEOGRAPHIC AND ECOLOGICAL HISTORY OF THE COSMOPOLITAN ANGIOSPERM GENUS STELLARIA L. (CARYOPHYLLACEAE) Introduction....................................................................................183 Materials and Methods...................................................................193 Results............................................................................................201 Discussion......................................................................................216 IV. THE EVOLUTION OF PETAL LOSS IN THE COSMOPOLITAN GENUS STELLARIA L. (CARYOPHYLLACEAE) Introduction....................................................................................233 Materials and Methods...................................................................235 Results............................................................................................242 Discussion......................................................................................251 BIBLIOGRAPHY..............................................................................................................261 APPENDICES AND SUPPLEMENTARY FIGURES.....................................................285 ix TABLES Table 1. Number of species in each family in the South San Juan Mountains....25 2. The eight largest genera in the South San Juan Mountains...................26 3. Number of families and species in each Division in the South San Juan Mountains..........................................................................26 4. Number of species and percentage of the flora in each distribution type in the South San Juan Mountains...................30 5. A generalized classification of distribution types in the South San Juan Mountains..........................................................................30 6. List of core Stellaria plus outgroup accessions sampled, including specimen ages and data yields...................................................105 7. Estimate of number of core Stellaria species inhabiting major geographical regions as well as total global Stellaria................111 8. Known clades of core Stellaria..............................................................158 9. Stellaria species sampled for biogeographical analysis.........................202 10. BioGeoBEARS model comparisons......................................................209 11. Ancestral state reconstruction of