Studies in the Cyperaceae of Georgia: Distribution of Georgia Sedges, Analysis of the Cyperus squarrosus-granitophilus Complex & Two New Species A Thesis submitted to the Graduate School Valdosta State University in partial fulfillment of requirements for the degree of MASTER OF SCIENCE In Biology in the Department of Biology of the College of Arts and Sciences July 2018 Phillip Douglas Lowe BS, Valdosta State University, 2014 © Copyright 2018 Phillip Lowe All Rights Reserved FAIR USE This thesis is protected by the Copyright Laws of the United States (Public Law 94-553, revised in 1976). Consistent with fair use as defined in the Copyright Laws, brief quotations from this material are allowed with proper acknowledgement. Use of the material for financial gain without the author’s expressed written permission is not allowed. DUPLICATION I authorize the Head of Interlibrary Loan or the Head of Archives at the Odum Library at Valdosta State University to arrange for duplication of this thesis for educational or scholarly purposes when so requested by a library user. The duplication shall be at the user’s expense. Signature _______________________________________________ I refuse permission for this thesis to be duplicated in whole or in part. Signature ________________________________________________ ABSTRACT The sedge family (Cyperaceae) is one of the most impactful families to agriculture and a key constituent of graminoid communities in many ecosystems. However, there is no full accounting of all of sedge species vouchered in Georgia. Understanding which sedge taxa occur in the state and where they exist is necessary for both effective pest management and protection of communities and species of interest. I examined holdings at the three largest herbaria in the state to inventory the state's sedge flora. For each taxon, I assessed phenology and the physiographic regions where it occurred. Along with this inventory, I did morphometric analyses of three narrow taxonomic problems. The first analyzed the relationship between the granite outcrop endemic Cyperus granitophilus and its close generalist relative, Cyperus squarrosus. In addition to confirming these species are morphometrically distinct, I discovered a new variety of C. squarrosus which is restricted to granitic uplifts of Texas and Oklahoma. I also investigated two putatively new sedge species from Georgia, one, a Cyperus restricted to exsiccated karst ponds in southwestern Georgia, and, the other, an Eleocharis restricted to high-marshes of Camden County, Georgia. My analysis supports both as distinct species. i TABLE OF CONTENTS I. Chapter I – An Annotated Checklist of the Cyperaceae of Georgia Based on the Holdings of the Three Largest Herbaria in the State ................................................................................................. 1 Introduction ........................................................................................................................ 1 Materials and Methods ....................................................................................................... 2 Results ................................................................................................................................. 4 Discussion ........................................................................................................................... 4 References .......................................................................................................................... 7 Tables and Figures ............................................................................................................ 12 Appendix A: A Checklist of the Cyperaceae of Georgia .................................................... 28 Appendix B: Exsiccatae for Checklist ................................................................................ 69 II. Chapter II – A Morphometric analysis of the Cyperus squarrosus-granitophilus Complex in North America .............................................................................................................................. 133 Introduction .................................................................................................................... 133 Materials and Methods ................................................................................................... 134 Results ............................................................................................................................. 136 Discussion ....................................................................................................................... 136 Taxonomic Treatment and Key ....................................................................................... 139 References ...................................................................................................................... 142 Tables and Figures .......................................................................................................... 144 Appendix A: Vouchers Examined Physically and Digitally .............................................. 165 III. Chapter III – A Morphometric Analysis of a Putative New Cyperus from Karst Ponds in Southern Georgia ........................................................................................................................................ 185 Introduction .................................................................................................................... 185 ii Materials and Methods ................................................................................................... 185 Results ............................................................................................................................. 186 Discussion ....................................................................................................................... 187 Taxonomic Treatment and Key ....................................................................................... 188 References ...................................................................................................................... 190 Tables and Figures .......................................................................................................... 191 Appendix A: Additional Vouchers Examined .................................................................. 205 IV. Chapter IV -- A Morphometric Analysis of a Putative New Eleocharis from the High Marsh of Camden County ............................................................................................................................ 212 Introduction .................................................................................................................... 212 Materials and Methods ................................................................................................... 213 Results ............................................................................................................................. 213 Discussion ....................................................................................................................... 214 Taxonomic Treatment and Key ....................................................................................... 214 References ...................................................................................................................... 217 Tables and Figures .......................................................................................................... 218 Appendix A: Additional Specimens Examined Physically and Digitally ........................... 230 iii LIST OF TABLES Chapter I Table 1. Number of species, infraspecific taxa, and total taxa vouchered at Georgia herbaria GA, GAS, and VSC. .......................................................................................................................... 12 Table 2. Sedges vouchered on 2017 Kyllinga collecting trip. ........................................................ 13 Table 3. Sedges vouchered on 2017 Glascock County collecting trip. ........................................... 14 Table 4. Sedges vouchered on 2017 Montgomery County collecting trip..................................... 15 Table 5. Phenology of sedge genera based on herbarium specimens at GA, GAS, and VSC herbaria label data. ........................................................................................................................ 16 Table 6. Numbers of sedge species restricted to one or two physiographic regions, seasonal phenology, nativity, and weed status, based on vouchers at GA, GAS, and VSC herbaria label data. ............................................................................................................................................... 17 Table 7. Sedge genera in Georgia by number of taxa, physiographic region, seasonal phenology, nativity, and weed status ............................................................................................ 18 Chapter II Table 1. List of characters and abbreviations used in principal component and discriminant analyses ................................................................................................................... 144 Table 2. Results of PCA of 24 measurements of 405 specimens ................................................. 145 Table 3. MANOVA for all 24 characters between all four taxa .................................................... 146 Table 4. Results of discriminate analysis ....................................................................................