FLORISTIC .AND BIOSYSTEMATIC INVESTIGATIONS IN PLANT TAXONOMY By LINDA CAROL GATTI CLARK Bachelor of Science University of Central Arkansas Conway, Arkansas 1989 Submitted to the Faculty of the Graduate College of the Oklahoma State Univesity in partial fulfillment of the requirements for the Degree of DOCTOR OF PHILOSOPHY July, 1997 ,qci r;, G \1 bfl FLORISTIC AND BIOSYSTEMATIC INVESTIGATIONS IN PLANT TAXONOMY Thesis Approved: Dean of the Graduate College ii ACKNOWLEDGMENTS With any of life's meaningful endeavors, one does not walk alone. During the course of this dissertation, a community of individuals shared this journey with me. I extend my gratitude to those individuals who have been extremely supportive and instrumental in helping me reach my goals. I thank my friends, fellow graduate students, and faculty in the Departments of Botany, Physics, and Zoology for their support during this adventure. I aiso express my appreciation to the members of my committee, Drs. Dave Engle, Anne Ewing, and Bill Henley. I am grateful for their advice, support, and assistance during the course of this study. For instilling in me a love and appreciation for plants, I thank my undergraduate adviser and friend, Dr. Donald Culwell of the University of Central Arkansas, Conway. I thank several individuals who aided the projects presented here. For their assistance in plant identification I would like to thank Dr. Ronald J. Tyrl, Patricia Folley and Cheryl McCreary. I wish to thank Dr. Paul Buck for his thorough review of the Asteraceae character list. Words cannot express my sincere appreciation and gratitude toward my mentor and friend, Dr. Ronald J. Tyrl. His guidance, friendship, and support have been invaluable in the realization of this dissertation. I thank him for treating me as colleague, allowing me to experience all the rewards that comes with such responsibility. 111 For their unwavering support, words of encouragement, and love, I would like to thank my parents and in-laws, Joseph and Patricia Gatti, Sue Clark, and Bob and Leanna Clark. This work could not have been possible without the loving support of my family. To my husband, Michael M. Clark, who has offered his unequivocal support to see me achieve my goal and to my daughter, Shelby T. Clark, who has made life a little more interesting along the way, I offer my deepest thanks for their encouragement, understanding, and love. iv TABLE OF CONTENTS Chapter Page I. ORGANIZATION OF DISSERTATION . 1 PART I: DELTA CHARACTER LIST FOR THE ASTERACEAE OF OKLAHOMA II. INTRODUCTION . 3 III. LITERATURE REVIEW . 5 Ecogeography of Oklahoma and Need for a Modem Taxonomic Treatment . 5 Floristic Work in Oklahoma . 7 Introduction to the Asteraceae. 12 Morphology - Specialized Terminology. 14 Inflorescences . 14 Involucre . 15 Receptacular Bracts . 15 Florets . 15 Pappus . 18 Achenes. 18 Stamens . 19 Styles . 20 Subtribal and Tribal Classification of the Asteraceae . 21 IV. METHODOLOGY . 26 V . RESULTS . 28 VI. LITERATURE CITED . 33 V PART II: MORPHOMETRIC STUDIES OF PAPPOPHORUM SCHREBER (POACEAE: CHLORIDOIDEAE: PAPPOPHOREAE) VII. MORPHOMETRIC STUDIES OF PAPPOPHORUM SCHREBER (POACEAE: CHLORIDOIDEAE: P APPOPHOREAE . 39 Introduction . 39 Material & Methods . 41 Results & Discussion. 46 Taxonomy of Pappophorum. 60 Key to Species . 62 Species Descriptions . 63 Literature Cited . 75 PART III: SURVEY OF THE VASCULAR FLORA OF THE BOEHLER SEEPS AND SANDHILLS PRESERVE VIII. SURVEY OF THE VASCULAR OF THE BOEHLER SEEPS AND SANDHILLS PRESERVE . 78 Introduction . 78 Boehler Seeps & Sandhills Preserve . 79 Method of Survey . 81 Flora of the Preserve . 82 References . 84 Appendices APPENDIX A- DELTA-CHARS ... ; . 87 APPENDIX B- DELTA-SPECS. 162 APPENDIX C - DELTA-ITEMS . 165 APPENDIX D - DELTA-TONAT. 167 APPENDIX E - Herbarium Specimens of Pappophorum used for NT-SYS analysis. 169 VI APPENDIX F - Data Sheet used to record 151 morphological features of each specimen of Pappophorum. 202 APPENDIX G - Phenogram resulting from NT-SYS analysis of Pappophorum (51 characters by 676 OTUs). 206 APPENDIX H - Vascular plant collections from Boehler Seeps and Sandhills Preserve arranged alphabetically by species. Collections of L.C. Gatti Clark during the 1994 and 1995 growing seasons . 224 APPENDIX I - Vascular plant collections from Boehler . Seeps and Sandhills Preserve arranged alphabetically by species. Collecitons made by The Nature Conservancy personnel . 232 APPENDIX J - Vascular plant collections made by L.C. Gatti Clark and The Nature Conservancy personnel arranged by family . 236 vu LIST OF TABLES Table Page PART I 1. Individuals producing taxonomic treatments of the flora of Oklahoma . 8 2. Tribes and genera of Oklahoma composites following Bremer's (1994) classification. 13 3. A Comparison of Asteraceae Tribes in Oklahoma: Bentham (1873) vs. Bremer (1994) .....· . 24 PART II 1. Characters emphasized in most recent treatments of Pappophorum (Pensiero 1986, Reeder & Toolin1989, Gatti Clark 1997). • . ......-. 42 2. Morphological characters and character states or numerical units used in NT-SYS analysis of species ofPappophorum ..... 44 viii LIST OF FIGURES Figure Page PART II 1. Simplification of phenogram (Appendix G) showing 7 groups of Pappophorum . 48 2. Portions of phenogram (Appendix G) with OTUs labeled showing clustering of P. vaginatum (VA), P. caespitosum (CA), and P. subbulbosum (SU, UN) . 57 3. Univariate analysis of P. vaginatum (diamond), P. 58 caespitosum, (square) and P. subbulbosum (cross) ........ IX CHAPTER I Organization of Dissertation Plant Taxonomy or Systematics is a multifaceted discipline founded on the understanding and interaction of the many conceptual levels of plant biology. This dissertation summarizes three studies that were selected to expose the investigator to several facets of the discipline. Each part is an independent venture and represents a different type of taxonomic work. In Part I, a DELTA character list is presented. It was produced for the Flora of Oklahoma Project and is to be used for the generation of descriptions of heads for the genera and species of the Asteraceae within Oklahoma. Part II is a morphometric (or numerical-taxonomic) investigation of the species of Pappophorum (Poaceae). Part III describes the results of a floristic survey of the Boehler Seeps and Sandhills Preserve in southeastern Oklahoma. To enhance readability and to facilitate submission for publication, each section or part is complete in itself and written in the format traditionally used in taxonomic journals or floras. It is anticipated that Part I will be incorporated into the Flora of Oklahoma, Part II will be submitted to Systematic Botany, and Part III will be submitted to the Proceedings of the Oklahoma Academy of Sciences. 1 PART I DELTA CHARACTER LIST FOR THE . ASTERACEAE OF OKLAHOMA 2 CHAPTER II Introduction "The family Compositae or Asteraceae has attracted, fascinated and even repelled botanists for over two centuries." V.H. Heywood Oklahoma has tremendous climatic and geological diversity. Twenty­ six geomorphic provinces and 19 land resource areas have been described (Johnson et al., 1979; National Resources Conservation Service, 1992). The state's environmental diversity is reflected in its floristic diversity. Plants characteristic of the Gulf Coastal Plain, Ouachita Uplands, Ozark Plateau, Great Plains, southwestern deserts, and Rocky Mountain foothills can be found within the state. Oklahoma ranks second among all states in the number of vascular plant families with 172 that encompass some 850 genera and 2,600 species (Tyrl et al., 1994). A significant component of this flora is the Asteraceae or composite family. Representing approximately 12-13% of the genera and species in the state, it comprises some 104 genera and 350 species (Waterfall, 1969; Taylor and Taylor, 1994). All taxa exhibit the family's characteristic head inflorescence with its distinctive morphological features. A specialized terminology has evolved to describe the features of the composite head and must be dealt with by the authors of taxonomic manuals such as the forthcoming Flora of Oklahoma. State taxonomists writing this 3 treatise have adopted the use of DELTA (DEscriptive Language for TAxonomy, Dallwitz and Paine, 1986), which is an automated description/identification program that provides a standardized format for taxonomic descriptions by using a coded list of characters. Families such as the Asteraceae require the generation of specific character lists which cover their specialized structures and terminology. The work summarized here was conducted in conjunction with the Flora of Oklahoma Project. The objectives were: 1) to create a DELTA-based character list for the features of the heads of Oklahoma composites, and 2) generate descriptions of the heads of representative genera and species of the family. The products of this research effort -- character list and taxon descriptions -- were submitted to the editorial board of the project for subsequent use by contributing taxonomists. 4 CHAPTER III Literature Review Ecogeography of Oklahoma and Need for a Modern Taxonomic Treatment With respect to climate, geology, geography and topography, Oklahoma is quite diverse. Mean annual temperatures across the state range from 1f. in the south to 14°C at the western end of the Panhandle (Ruffner, 1980 average length of the growing season varies from 168 to 225 dav:c 5e rainfall amounts vary as well, from 142 cm per year in the- ~)'.>""uu~ast to around 38 cm per year at the western end of the Panhandle.