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SYSTEMATIC and EVOLUTIONARY STUDIES in the DICHANTHELIUM ACUMINATUM (POACEAE:PANICEAE) COMPLEX a Dissertation by RICKY LEE HAMME

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SYSTEMATIC and EVOLUTIONARY STUDIES in the DICHANTHELIUM ACUMINATUM (POACEAE:PANICEAE) COMPLEX a Dissertation by RICKY LEE HAMME SYSTEMATIC AND EVOLUTIONARY STUDIES IN THE DICHANTHELIUM ACUMINATUM (POACEAE:PANICEAE) COMPLEX A Dissertation by RICKY LEE HAMMER 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 May 2010 Major Subject: Botany SYSTEMATIC AND EVOLUTIONARY STUDIES IN THE DICHANTHELIUM ACUMINATUM (POACEAE:PANICEAE) COMPLEX A Dissertation by RICKY LEE HAMMER 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: Co-chairs of Committee, James Manhart Stephan Hatch Committee Members, Alan Pepper Hugh Wilson Head of Department, U. J. McMahan May 2010 Major Subject: Botany iii ABSTRACT Systematic and Evolutionary Studies in the Dichanthelium acuminatum (Poaceae:Paniceae) Complex. (May 2010) Ricky Lee Hammer, B.B.A., The University of Texas-Permian Basin; M.S., Texas A&M University Co-Chairs of Advisory Committee: Dr. James Manhart Dr. Stephan Hatch Taxonomic boundaries and systematic relationships in the grass subspecific complex Dichanthelium acuminatum were investigated with both morphological and molecular methods. Circumscription of subspecific taxa comprising the complex has been difficult due to a continuum of morphological character variation among taxa and possibly due to infraspecific and interspecific hybridization. Qualitative and quantitative morphological character data was collected from herbarium specimens and field-collected specimens and analyzed using multivariate statistical techniques. Representative specimens were selected for molecular phylogenetic analysis of DNA sequences from the GBSSI (waxy) nuclear gene. Subspecific boundaries as circumscribed in the most recent taxonomic treatment (10 subspecies) were tested from: 1) a morphological perspective with results of the multivariate statistical analysis to determine if the study specimens formed natural groupings that corresponded to the recent treatment; and, 2) with molecular phylogenetic analysis to estimate the evolutionarily significant lineages present and to determine if such lineages supported the natural groupings revealed from the iv multivariate morphological analysis. A separate investigation was conducted using a molecular technique to screen for putative hybrid specimens from DNA obtained from field-collected specimens. Multivariate statistical analysis of the morphological data provided support for four of the 10 taxa tested and additional support for two taxa considered as a single unit. Further research is needed to determine the appropriate status of the remaining six taxa of the ten taxa tested. Molecular phylogenetic analysis provided support for recognizing four evolutionarily significant units and provided parallel support for four of the five taxa recognized from the morphological analysis. The hybridization investigation identified two putative hybrid specimens, which were confirmed as hybrids with GBSSI sequence data and also with multivariate statistical analysis of morphological data to provide provisional evidence for the role of hybridization in producing specimens with intermediate morphological phenotypes. A taxonomic treatment and dichotomous key was produced for the 10 subspecific taxa of the Dichanthelium acuminatum complex. v DEDICATION I would like to dedicate this work to the memory of my late parents, Jack Leroy Hammer and Leola Francis Williamson Hammer. Countless fishing, camping, and backpacking trips with my father as I was growing up in west Texas provided the genesis and spark for my deep interest in the natural history of Texas and farther afield. That early immersion in natural history has matured and merged into a focus on evolutionary history, an interest which continues to pay rich intellectual rewards. Such gifts often start from the smallest of seeds. Thanks to my mother for encouraging and nurturing the development of a child who would eventually aspire to a career in science and teaching. We are all, in some ways big and small, the products of our parents. I also dedicate this disstertation, which represents many hours and years of work, to my beloved wife, Tiffany Kristen Hammer. Without her constant support and tolerance of many late nights in the laboratory and many Saturdays and Sundays spent in research and writing I would not have been able to complete my doctorate. I will always value her constant love and support. I also dedicate this work to my beloved son, Nathan, who came into the world at the midpoint of my doctoral journey. Time away from both my wife and son has been the only detrimental aspect of my research. I would also like to thank my sister, Charleen Ikeler Jones, for her love and encouragement over the years. vi ACKNOWLEDGEMENTS I would like to thank my committee co-chairs, Dr. James Manhart (Biology) and Dr. Stephan Hatch (Ecosystem Science and Management), for their guidance and support throughout the course of this research. I have benefited greatly from their knowledge and experience. My other committee members, Dr. Alan Pepper and Dr. Hugh Wilson, were also instrumental in the success of my doctoral research at Texas A&M. I thank all of my committee members for the friendship they have extended beyond professional advisement. Significant financial support was provided by several grants from the Fank W. Gould Research Award in Plant Systematics. Dr. Gould left a substantial legacy concerning the study of the genus Dichanthelium at Texas A&M including many fine herbarium specimens from which a novice could begin the difficult task of ―learning‖ how to identify the various members of Dichanthelium. Dale (72‘) and Phoebe Watts kindly provided a grant to assist in the early stages of the molecular work and their contribution is greatly appreciated. Recognition is also due the Department of Biology for providing assistance in the form of teaching assistantships over the years. I would also like to thank Dr. Terry Thomas of the Laboratory for Functional Genomics in the Department of Biology for use of the lab‘s Linux cluster for computational support for the molecular phylogenetic portion of my research. Thanks also go to Dr. Robert Freckmann, retired from the vii University of Wisconsin-Stevens Point, for constructive comments on Dichanthelium classification and for supplying various herbarium specimens. Thanks go to several entitities that allowed me to conduct field research in support of dissertation: Texas Parks & Wildlife Department (various parks and WMAs), Marysee Prairie Preserve (Hardin County, TX), and Texas Nature Conservancy (Roy Larsen Sandylands Preserve, Hardin County, TX). viii TABLE OF CONTENTS Page ABSTRACT .............................................................................................................. iii DEDICATION .......................................................................................................... v ACKNOWLEDGEMENTS ...................................................................................... vi TABLE OF CONTENTS .......................................................................................... viii CHAPTER I INTRODUCTION… ............................................................................ 1 II MORPHOLOGICAL ANALYSIS ...................................................... 6 Introduction ........................................................................................... 6 Materials and Methods .......................................................................... 6 Results ..................................................................................................... 9 Discussion ............................................................................................... 17 III MOLECULAR PHYLOGENETIC ANALYSIS ................................. 39 Introduction ............................................................................................ 39 Materials and Methods .......................................................................... 41 Results ..................................................................................................... 44 Discussion ............................................................................................... 45 IV HYBRIDIZATION IN DICHANTHELIUM ........................................ 63 Introduction ............................................................................................ 63 Materials and Methods .......................................................................... 65 Results ..................................................................................................... 68 Discussion ............................................................................................... 69 Conclusion .............................................................................................. 72 V TAXONOMIC TREATMENT AND SUMMARY ............................. 73 Introduction ............................................................................................ 73 Taxonomic Treatment and Summary.................................................. 74 ix Page LITERATURE CITED ............................................................................................. 91 APPENDIX A: FIGURES ........................................................................................ 98 APPENDIX B: TABLES .......................................................................................... 140 APPENDIX C: COMPUTATIONAL
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