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A SYSTEMATIC STUDY OF SELECT SPECIES COMPLEXES OF ELEOCHARIS SUBGENUS LIMNOCHLOA (CYPERACEAE) A Dissertation by DAVID JONATHAN ROSEN Submitted to the Office of Graduate Studies of Texas A&M University in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY December 2006 Major Subject: Rangeland Ecology and Management A SYSTEMATIC STUDY OF SELECT SPECIES COMPLEXES OF ELEOCHARIS SUBGENUS LIMNOCHLOA (CYPERACEAE) A Dissertation by DAVID JONATHAN ROSEN Submitted to the Office of Graduate Studies of Texas A&M University in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY Approved by: Chair of Committee, Stephan L. Hatch Committee Members, J. Richard Carter William E. Fox III James R. Manhart Fred E. Smeins Head of Department, Steven G. Whisenant December 2006 Major Subject: Rangeland Ecology and Management iii ABSTRACT A Systematic Study of Select Species Complexes of Eleocharis Subgenus Limnochloa (Cyperaceae). (December 2006) David Jonathan Rosen, B.S., Texas State University; M.S., Texas A&M University Chair of Advisory Committee: Dr. Stephan L. Hatch A systematic study of two complexes of closely related species within Eleocharis subg. Limnochloa was conducted to better define poorly understood species and to lay the foundation for a worldwide revision of this group. Research utilized scanning electron microscopy (SEM), study of more than 2300 herbarium specimens and types from 35 herbaria, multivariate analysis, and field studies in the southeast United States and Mexico. Examination of achene gross- and micromorphology using SEM indicated a relationship among the species of the Eleocharis mutata complex (comprising E. mutata, E. spiralis, and E. cellulosa), their distinctness from the E. acutangula complex (comprising E. acutangula s.l. and E. obtusetrigona) and support the placement of all taxa studied within subg. Limnochloa. All species examined shared micromorphological characters typical of subg. Limnochloa. A systematic study of the worldwide morphological variation of E. mutata suggests it is represented by a single taxon throughout its distribution and is native to the New World and iv probably introduced in tropical Africa. The taxonomic relationship of Eleocharis mutata and E. spiralis was explored using discriminant analysis, PCA and phyto- geography, supporting the recognition of the two taxa as distinct at the rank of species. A systematic study of E. cellulosa suggests it is represented by a single taxon throughout its distribution, with an apparent Caribbean genotype distinguished by the presence of perianth bristles with retrorse spinules. A systematic study of the worldwide variation of E. acutangula s.l. resulted in its segregation into two infraspecific taxa, E. acutangula subsp. acutangula and E. acutangula subsp. breviseta, and two new South American species, E. neotropica and E. steinbachii. Nomenclatural history and lectotypification of certain taxa studied are discussed, and two basyionyms and two synonyms of accepted taxa are lectotypified. A taxonomic treatment is provided that includes a key, detailed descriptions and complete synonymy of each species, line drawings prepared from select specimens examined, and notes on habitat and distribution. Future research goals and needs are discussed. v DEDICATION This dissertation is dedicated to my grandparents Max and Molly Freedman, my brother Sam Rosen, and my good friend Robert M. Combs. Your passing has left a void in my life that will never be filled. vi ACKNOWLEDGMENTS I would like to offer my sincere thanks to my major professor Dr. Stephan Hatch for providing me the opportunity to earn this degree when others would not, and for his wisdom, direction, encouragement, and patience. I thank my committee members, Dr. Bill Fox, Dr. Jim Manhart, and Dr. Fred Smeins for reviewing my research, and for challenging me to expand the depth and breadth of my knowledge. I offer special thanks to Dr. Richard Carter for agreeing to serve as a special appointment, for being a mentor and friend, for companionship during field work in Texas, Florida, and Mexico, and for critically reviewing this research with patience and enthusiasm. Special thanks are due to Dr. M. Socorro González-Elizondo for suggesting several research problems, for sharing her expertise, and for being a friend, mentor, and generous host during my field work in Mexico. Special thanks are also due to José Marcos Pinedo Reyes for his assistance in the field, and especially for driving us over 3800 km through central and southern Mexico. I’d like to acknowledge numerous colleagues near and far who provided assistance and freely shared their knowledge and offered advice: Stanley Jones, J. Wipff, Fran Menapace, Galen Smith, Tony Reznicek, Eric Roalson, Paul Peterson, Rob Naczi, Mark Strong, and Charles Bryson. I am grateful to Dale Kruse for requesting loans for my research, and the curatorial and library staff of all of the herbaria mentioned for loan of specimens for study, assistance with questions about the disposition of certain specimens, types, and literature, and assistance with nomenclatural vii questions. I am especially grateful to Mark Spencer, Vicki Noble, and Peter Stafford (BM), Hajo Esser (M), Ib Friis (C), Jill Marsden (K), Kanchi Gandhi (GH), and Gina Murrell (CGE) for their friendly responses to my endless queries. I am extremely grateful to my friend Neva Mikulicz for preparing the excellent illustrations, to Alain Touwaide for translating the Latin diagnoses, to Mike Pendleton at the Microscopy Imaging Center for assistance, to Susan Knock for use of her lab, and to Larry Brown for requesting loans and types for me to study at SBSC. Above all, I thank my wife Kimberly and daughter Molly. Like Papa used to say, I make the living, and you make life worth living! I gratefully acknowledge financial support for research related expenses, supplies and travel from the Frank W. Gould Award for Graduate Student Research in Plant Systematics, Springfield Research Award, McCord Scholarship, and Mills Scholarship. viii TABLE OF CONTENTS Page ABSTRACT ………………………………….……………………………….. iii DEDICATION ………………………………………………………………… v ACKNOWLEDGMENTS ……………………………………………………. vi TABLE OF CONTENTS …………………………………………………….. viii LIST OF FIGURES ..……………………….………………………………... x LIST OF TABLES ……………………………………………………………. xiv CHAPTER I INTRODUCTION …………………………………………………. 1 Research Objectives ………………………………………….. 14 II ACHENE MORPHOLOGY OF SELECTED SPECIES OF ELEOCHARIS SUBG. LIMNOCHLOA ………………………… 16 Introduction ……………..……………………………………… 16 Materials and Methods ...……………………………………… 18 Results and Discussion ……………………………………….. 23 Conclusion ……………………………………………………… 41 III MORPHOMETRIC NUMERICAL STUDY OF WORLDWIDE VARIABILITY OF ELEOCHARIS MUTATA ………………….. 44 Introduction .…………………………….……………………… 44 Materials and Methods ………………..……………………… 47 Results ………………………………………………………..... 52 Discussion ……………………………………………………… 58 Conclusion ……………………………………………………… 59 ix CHAPTER Page IV MORPHOMETRIC NUMERICAL STUDY OF THE TAXONOMIC RELATIONSHIP OF ELEOCHARIS MUTATA AND ELEOCHARIS SPIRALIS ….………..…………………… 64 Introduction ……………..……………………………………… 64 Materials and Methods …..……………….…………………… 67 Results ………………………………………………………….. 71 Discussion and Conclusion ….…..…………………………… 80 V MORPHOMETRIC NUMERICAL STUDY OF POTENTIAL INFRASPECIFIC SEGREGATES OF ELEOCHARIS CELLULOSA ………………………………………..…………... 82 Introduction …………………..………………………………… 82 Materials and Methods ..……………………………………… 84 Results …………………………..….………………………….. 86 Discussion and Conclusion ….….….………………………… 92 VI MORPHOMETRIC NUMERICAL STUDY OF WORLDWIDE VARIABILITY OF ELEOCHARIS ACUTANGULA …………… 96 Introduction ..…………………………………………………… 96 Materials and Methods ..…………….………………………… 99 Results …………………………..…….……………………….. 103 Discussion and Conclusion …..……….……………………… 114 VII NOMENCLATURE ………………………………………………. 122 Introduction …………………………….………………………. 122 Nomenclatural History of Eleocharis Subg. Limnochloa ….. 124 Nomenclatural History and Lectotypification in the Eleocharis mutata Complex ………………………………….. 125 Nomenclatural History and Lectotypification in the Eleocharis acutangula Complex …………………………….. 128 VIII TAXONOMIC TREATMENT ……………………………………. 131 Description of Eleocharis Subg. Limnochloa with Key and Species Accounts for Taxa Treated ………………………… 131 x CHAPTER Page IX SUMMARY AND SUGGESTED FUTURE RESEARCH ……… 167 REFERENCES ………………………………………………………………. 171 APPENDIX I ………………………………………………………………….. 188 APPENDIX II …………………………………………………………………. 192 APPENDIX III ………………………………………………………………… 198 APPENDIX IV ………………………………………………………………... 200 APPENDIX V ………………………………………………………………… 208 APPENDIX VI ……………………………………….……………………….. 210 VITA ………………………………………………….……………………….. 212 xi LIST OF FIGURES FIGURE Page 1 SEM micrographs of achenes of representative specimens of Eleocharis mutata……………………………………………. 24 2 SEM micrograph of achenes of representative specimens of Eleocharis mutata and E. cellulosa ……..…………………… 25 3 SEM micrographs of achenes of representative specimens of Eleocharis spiralis ..…………………………………………. 27 4 SEM micrographs of achenes of representative New World specimens of Eleocharis acutangula ..……………………….. 30 5 SEM micrographs of achenes of representative Old World specimens of Eleocharis acutangula ..……………………….. 31 6 SEM micrographs of achenes of representative specimens of Eleocharis acutangula, E. steinbachii, and E. neotropica 32 7 SEM micrographs of achenes of representative specimens of Eleocharis obtusetrigona ..……………………….…………. 34 8 SEM micrographs of achene micromorphology
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