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ABSTRACT a MORPHOLOGICAL INVESTIGATION of DICHANTHELIUM SECTION LANUGINOSA (POACEAE) by Justin R. Thomas Much Debate and Confusi

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ABSTRACT A MORPHOLOGICAL INVESTIGATION OF DICHANTHELIUM SECTION LANUGINOSA (POACEAE) by Justin R. Thomas Much debate and confusion has persisted regarding the morphological ranges and limits of those taxa found within section Lanuginosa of the genus Dichanthelium. Phenetic analysis, principal components analysis and pair-wise t-tests of character states were conducted with 285 specimens using 32 morphological characters. An additional examination of 458 herbarium specimens including all relevant type material resulted in the recognition of 16 species and two subspecies. Many of the entities subsumed within the D. acuminatum complex are found to represent distinct species in terms of morphology. Their geographic and habitat distinctions are also discussed. A review of all 160 names associated with the section was conducted and an accurate nomenclature was applied to the group. Dichanthelium lanuginosum is reinstated as distinct from D. acuminatum and two new nomenclatural combinations (D. thermale and D. thermale subsp. sericeum) are presented. A MORPHOLOGICAL INVESTIGATION OF DICHANTHELIUM SECTION LANUGINOSA (POACEAE) A Thesis Submitted to the Faculty of Miami University in partial fulfillment of the requirements for the degree of Master of Science Department of Botany by Justin Ray Thomas Miami University Oxford, Ohio 2008 Advisor ___________________________________ Dr. Michael A. Vincent Reader ___________________________________ Dr. R. James Hickey Reader ___________________________________ Dr. W. Hardy Eshbaugh © Justin Ray Thomas 2008 TABLE OF CONTENTS List of Tables…………………………………………………………………...... iv List of Figures……………………………………………………………………. v Acknowledgements………………………………………………………………. vi Introduction………………………………………………………………………. 1 Materials and Methods…………………………………………………………… 4 Results…………………………………………………………………………….. 6 Discussion…………………………………………………………………………. 8 Taxonomic Treatment…………………………………………………………….. 21 Literature Cited…………………………………………………………………… 62 Appendix A – Characters Sampled……………………………………………….. 87 Appendix B – Histograms................................…..............………...…………….. 91 Appendix C – Raw Data (Long Ligule Group)…..............………...…………….. 106 Appendix D – Raw Data (Short Ligule Group)………....................…….……….. 112 Appendix E – Specimens Sampled……………………………………………….. 116 iii LIST OF TABLES Table 1. Summary of Past Treatments…………………………………………………. 65 2. Characters Sampled for Analysis……………………………………………... 66 3. Results of T-tests for Long Ligule Group………...............…………………… 67 4. Principal Components Loadings for Long Ligule Group…...............………… 68 5. Results of T-tests for Short Ligule Group……….....................……………….. 69 6. Principal Components Loadings for Short Ligule Group………....................... 70 iv LIST OF FIGURES Figure 1. Phenogram of all OTUs.................................................................................... 71 2. Box plot of ligule lengths.................................................................................. 72 3. Phenogram of Long Ligule Group………................…………………………. 73 4. Scatter plot of Principal Components for Long Ligule Group…...............…. 74 5. Scatter plot of Principal Components for D. implicatum and D. lanuginosum. 75 6. Phenogram of Short Ligule Group……….....................…………………...… 76 7. Scatter plot of Principal Components for Short Ligule Group......................… 77 8. Illustrations of ligule and pubescence types within section Lanuginosa……… 78 9 - 24. Distribution maps.....…………………………..............………………..... 79-86 v ACKNOWLEDGMENTS I would like to thank Dr. Michael Vincent for his patience, sage advice and technical support throughout the life of this project. I am grateful to Dr. R. James Hickey, Dr. Linda Watson and Dr. W. Hardy Eshbaugh for their technical assistance as well as for shedding light on the opportunities and challenges presented in this project. I would also like to thank Doug Ladd of the Nature Conservancy of Missouri for making a kid from Brumley believe he could be a botanist. Ultimately, I cannot express enough thanks to my wife and partner, Dana Thomas, for her tireless support and encouragement. Without her pure smiles and gentle understanding this would not have been possible. vi INTRODUCTION The genus Panicum, sensu lato, is one of the largest genera of New World grasses. Because of its diversity and wide distribution, a taxonomic understanding of this notoriously difficult group is vital to floristic works, monitoring projects, restoration efforts and other areas of conservation concern. In 1910 Hitchcock and Chase published Panicum subgenus Dichanthelium to accommodate those taxa that produce terminal and axillary inflorescences (floral dimorphism), possess winter rosettes in addition to cauline leaves (foliar dimorphism), and rounded rather than pointed spikelets. This is in contrast to members of the subgenus Panicum which only produce terminal inflorescences, lack winter rosettes and generally have pointed spikelets. In addition to these differences, more contemporary studies have demonstrated further distinctions between subgenus Dichanthelium and subgenus Panicum. Brown (1948), Smith and Brown (1973) and Brown and Smith (1975) demonstrated that subgenus Dichanthelium utilize C3 photosynthetic pathways as opposed to subgenus Panicum which are primarily C4 grasses. Boyle (1945) reported that subgenus Dichanthelium has an extremely low incidence of polyploidy (only three out of approximately 100 species) whereas 70-80 percent of subgenus Panicum species are polyploid. Brown et al. (1957) noted that subgenus Dichanthelium has two layers of tunica cells in the shoot apices whereas subgenus Panicum has a single layer. Given the extent of such differences, Gould (1974) elevated subgenus Dichanthelium to the rank of genus. However, some taxonomists have been hesitant to accept this status change stating an abundance of tropical species that appear to be intermediate between the two genera (Lelong 1986; Webster 1988; Crins 1991; Gleason and Cronquist 1991; Yatskievych 1999). The argument against elevation of Dichanthelium to generic rank was further supported by Morrone and Zuloaga (1991) and Zuloaga et al. (1993a, b) who tested the evidence defending Dichanthelium as a distinct genus. Their results demonstrate intermediates and exceptions regarding foliar and floral dimorphism, ploidy level and C3 versus C4 photosynthesis, especially in Central and South American species. 1 More recent evidence has come from research involving the molecular phylogeny of Panicoideae (Giussani et al. 2001; Aliscioni et al. 2003; Morrone et al. 2008). These studies demonstrate that Panicum s.l. is polyphyletic unless Dichanthelium, among others, is treated as a separate genus. Additionally, many of the presumed intermediate taxa (most notably section Cordovensia) that were used to maintain Dichanthelium as a subgenus of Panicum have been shown to represent distinct lineages (Aliscioni et al. 2003; Morrone et al. 2008). Though neither the molecular nor morphological data offer a completely satisfying solution to the debate, their net effect seems to provide reasonable evidence supporting Dichanthelium as a distinct genus. The evolution of an over- wintering rosette in this group of grasses may have led to the diversification of Dichanthelium into temperate North America (Morrone 1991; Zuloaga et al. 1993b). This foliar dimorphism offers a reasonable synapomorphy for the genus. Dichanthelium is a new world genus of between 72-109 species (Gould and Clark 1978; Freckmann and Lelong 2003; Hitchcock and Chase 1910), of which 34 are native to North America (Freckmann and Lelong 2003). Members of the genus are perennial grasses that have erect to decumbent stems which range from 5-150 cm long, vary greatly in degrees of pubescence, and often have short, wide leaves relative to other grasses. Dichanthelium, as its name implies, experiences two flowering periods per growing season (Freckmann and Lelong 2003). All taxa in the genus produce a terminal paniculate inflorescence in spring followed by mid-summer inflorescences on axillary branches that continue to bloom through autumn. The spikelets of the vernal inflorescences are chasmogamous, while those of the autumnal inflorescences are occasionally cleistogamous (Zuloaga et al. 1993b). The spikelets of Dichanthelium range from 0.8-5.2 mm in length and typically have morphologically dissimilar glumes, one sterile lemma and a single fertile floret (Freckmann and Lelong 2003). In 1910, Hitchcock and Chase arranged the taxa of subgenus Dichanthelium into 17 informal groups based on morphological characters. Many authors since have incorrectly used these names as subgenera, sections or sub-sections, but only recently have any groups been validly published as sections within Dichanthelium (Freckman and 2 Lelong 2002). In so doing, Freckmann and Lelong (2002) reduced the 17 informal groups of Hitchcock and Chase (1910, 1951) to 13 sections (Angustifolia, Clandestina, Dichanthelium, Ensifolia, Lancearia, Lanuginosa, Linearifolia, Macrocarpa, Nudicaulia, Pedicellata, Oligosantha, Sphaerocarpa and Strigosa). Section Lanuginosa has the widest range of variation in the genus. Sections Dichanthelium, Lancearia, Angustifolia and Strigosa have the most morphological similarity to section Lanuginosa. The informal groups Lanuginosa, Spreta and Columbiana of Hitchcock and Chase (1910, 1951) (Table 1) were subsumed into section Lanuginosa by the treatment of Freckmann and Lelong (2002).
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