Synopsis of the Morphology and Taxonomy of Carex Section
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SYNOPSIS OF THE MORPHOLOGY AND TAXONOMY OF CAREX SECTION GLAUCESCENTES IN NORTH AMERICA A Thesis by DIANE COSTON MCLAUGHLIN Submitted to the Office of Graduate Studies of Texas A&M University in partial fulfillment of the requirements for the degree of MASTER OF SCIENCE August 2004 Major Subject: Rangeland Ecology and Management SYNOPSIS OF THE MORPHOLOGY AND TAXONOMY OF CAREX SECTION GLAUCESCENTES IN NORTH AMERICA A Thesis by DIANE COSTON MCLAUGHLIN Submitted to Texas A&M University in partial fulfillment of the requirements for the degree of MASTER OF SCIENCE Approved as to style and content by: _____________________________ _____________________________ Stephan L. Hatch Fred E. Smeins (Chair of Committee) (Member) _____________________________ _____________________________ Charles T. Hallmark Stanley D. Jones (Member) (Member) _____________________________ Steven G. Whisenant (Head of Department) August 2004 Major Subject: Rangeland Ecology and Management iii ABSTRACT The Morphology and Taxonomy of Carex Section Glaucescentes in North America. (August 2004) Diane Coston McLaughlin, B.S., Texas A&M University Chair of Advisory Committee: Dr. Stephan L. Hatch Field studies were used to characterize habitat and evaluate morphological characters of Carex glaucescens, C. joorii and C. verrucosa. Morphometric analysis of herbarium specimens along with field studies, Environmental Scanning Electron Microscopy (ESEM), pollen viability and phenology show C. glaucescens, C. joorii and C. verrucosa to be taxonomically distinguishable at the species level. The taxonomy of Carex section Glaucescentes is presented in an artificial dichotomous key to the taxa and in species descriptions. iv DEDICATION To the beginnings of every Botanist. v ACKNOWLEDGEMENTS I would like to thank Dr. Stephan L. Hatch for his research guidance and financial support throughout the course of this research. I would also like to thank my committee members for their contributions and time. Many special thanks to my family and friends for their support and help (Darrel McLaughlin, James Coston Sr., James and Catherine Coston Jr., Lisa Coston, Dale A. Kruse and David Rosen). Financial support for ESEM and collection trips was funded by the Gould Research Award. I would like to thank the curators and staff at the following institutions for providing loans: L.H. Bailey Hortorium (BH), Herbarium of Charles T. Bryson (CTB), Duke University (DUKE), University of Florida (FLAS), Florida State University (FSU), University of Michigan (MICH), North Carolina State University (NCSC), University of North Carolina (NCU), University of Louisiana at Monroe (NLU), New York Botanical Garden (NY), The Academy of Natural Sciences Herbarium (PH), Southern Forest Research Program (SFRP), Southern Weed Science Research Center (SWSL), S. M. Tracy Herbarium (TAES), University of Texas – Austin (TEX/LL), University of Alabama (UNA) and U.S. National Herbarium (US). I would like to thank several people whose correspondence made a valuable contribution to this work: Stanley D. Jones, Anton A. Reznicek; David Rosen, Dale A. Kruse, James Macklin; Thomas A. Zanoni, Jay Wipff, Kanchi Gandhi and Ernie Schuyler. vi TABLE OF CONTENTS Page ABSTRACT ...................................................................................................................iii DEDICATION ............................................................................................................... iv ACKNOWLEDGEMENTS .............................................................................................. v TABLE OF CONTENTS................................................................................................ vi LIST OF TABLES ........................................................................................................viii LIST OF FIGURES ........................................................................................................ x INTRODUCTION ........................................................................................................... 1 METHODS..................................................................................................................... 6 Field Work ............................................................................................................... 12 Cytology................................................................................................................... 12 Chromosome Numbers .................................................................................... 12 Pollen Stainability............................................................................................. 20 Achene and Perigynia Micromorphology.................................................................. 21 Data Analysis........................................................................................................... 21 Univariate Analysis........................................................................................... 21 Multivariate Analysis of Borrowed Specimens .................................................. 23 Multivariate Analysis of Collected Populations Including Suspected Hybrids....24 Multivariate Analysis with Carex joorii Holotype Specimen...............................24 Taxa Distributions.................................................................................................... 24 Phenology................................................................................................................ 24 Species Descriptions ............................................................................................... 25 RESULTS AND DISCUSSION..................................................................................... 29 Cytology................................................................................................................... 29 Chromosome Numbers .................................................................................... 29 Pollen Stainability............................................................................................. 29 Achene and Perigynia Micromorphology.................................................................. 30 Data Analysis........................................................................................................... 33 Univariate Analysis........................................................................................... 33 Multivariate Analysis of Borrowed Specimens .................................................. 33 Multivariate Analysis of Collected Populations Including Suspected Hybrids....49 Multivariate Analysis with Carex joorii Holotype Specimen...............................55 Taxa Distributions.................................................................................................... 55 Phenology................................................................................................................ 62 vii Page Taxonomy................................................................................................................ 64 Type Specimens .............................................................................................. 64 Artificial Dichotomous Key to Carex Section Glaucescentes in North America .66 Species Descriptions........................................................................................ 68 CONCLUSIONS .......................................................................................................... 75 REFERENCES ............................................................................................................ 77 APPENDIX A ............................................................................................................... 80 APPENDIX B ............................................................................................................... 84 APPENDIX C............................................................................................................... 94 APPENDIX D............................................................................................................... 98 APPENDIX E ............................................................................................................. 107 APPENDIX F ............................................................................................................. 116 APPENDIX G............................................................................................................. 120 VITA .......................................................................................................................... 123 viii LIST OF TABLES TABLE Page 1. Herbaria lending specimens and specimen images to TAES for use in this study………………………………………………………………………. 7 2. Characters used in the analysis. Character states are: continuous quantitative (CQ), two-state quantitative (TS), two-state qualitative (TQ) and multiple-state qualitative (MS)………………………………………….. 9 3. Specimens used in the numerical analysis and scanning electron microscopy…………………………………………………………………….. 13 4. Personal collections of Carex section Glaucescentes. Specimens used in PCA and those that are possible hybrid taxa are indicated…………… 16 5. Representative soil descriptions of Carex section Glaucescentes from personal collections…………………………………………………………... 18 6. Pollen stainability. Percent viability is recorded for specimens; pollen staining a dark color are recorded as viable……………………….. 22 7. Carex glaucescens univariate statistics from quantitative data used in PCA’s