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Taxonomic Study of the Clematis Reticulata Species Complex (Ranunculaceae: Subgenus Viorna)

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Taxonomic Study of the Clematis Reticulata Species Complex (Ranunculaceae: Subgenus Viorna) TAXONOMIC STUDY OF THE CLEMATIS RETICULATA SPECIES COMPLEX (RANUNCULACEAE: SUBGENUS VIORNA) By Thomas H. Murphy A Thesis Submitted in Partial Fulfillment of the Requirements for the Degree of Master of Science in Biology Austin Peay State University August 2020 Thesis Committee: Dr. L. Dwayne Estes, Committee Chair Dr. Don Dailey Dr. Rebecca Johansen iii Statement of Permission to Use In presenting this thesis in partial fulfillment of the requirements for the Master of Science in Biology at Austin Peay State University, I agree that the library shall make it available to borrowers under the rules of the library. Brief quotations from this thesis are allowable without special permission, provided that accurate acknowledgement of the source is made. Permissions for extensive quotation or reproduction of this thesis may be granted by my major professor, or in his absence, by the Head of the Interlibrary Services when, in the opinion of either, the proposed use of the material is for scholarly purposes. Any copying or use of the material in this thesis for financial gain shall not be allowed without my written permission _____________________________________________________________ Thomas H. Murphy 07/01/2020 iv I dedicate this research to my family. v ACKNOWLEDGMENTS I would like to acknowledge and thank my advisor and mentor Dr. Dwayne Estes for giving me the opportunity to undertake this study for my graduate research, as well as providing guidance during my time at Austin Peay State University (APSU). I thank Dr. Johansen and Dr. Dailey, my committee members, for providing guidance and helpful comments for my manuscript. I would like to thank Mt. Cuba Center for funding the majority of this study and the broader revisionary systematics research of North American Clematis subgenus Viorna with the Mt. Cuba Center Graduate Fellowship in Eastern North American Botany. I would also like to thank the British Clematis Society for funding additional fieldwork and supplies to complete our research, Lyndy Broder for funding fieldwork, and APSU Center of Excellence for Field Biology for allowing the use of field vehicles. I am grateful to Zach Irick, Mason Brock, Joseph Aroh, Dr. Brian Keener, Kevin England, Jesse Harris, Kate Love, Glen Dale Bryant, and Steve Scarborough for assistance and guidance during fieldwork. I thank Dr. Jared Barnes, Sonnia Hill, and Theo Witsell for sending herbarium specimens and/or living plants to include in our study. I would like to thank Claire Ciafre and Annie Lindsey for generously caring for common garden plants while I was travelling for fieldwork. I thank Dr. Aaron Floden for discussing and providing valuable insight into Clematis taxonomy. I am appreciative of Dr. Kanchi Gandhi for expertise and helpful comments in regard to the Shenzhen Code. I am grateful to Alan Cressler for allowing me to use his photograph from Pine Mountain, Georgia. I also thank the curatorial staff of all herbaria that generously sent specimen loans to APSC, as well as MO for allowing me to visit. I would like to thank my parents for their unwavering support of me. Lastly, I would like to thank my partner, Kate, for always being supportive, encouraging, and understanding while we worked towards our degrees from a distance. vi ABSTRACT Chapter 1 is a literature review of the taxonomic history of Clematis subg. Viorna. Chapter 2 is a morphometric study and taxonomic treatment of the Clematis reticulata species complex. Clematis subgenus Viorna of North America currently consists of 21 species, most of which are distributed in the southeastern United States. Past taxonomic treatments of subg. Viorna have varied greatly, which has led to unclear species limits. Often recognized as a single, polymorphic species with a high degree of ecological amplitude, variation in C. reticulata s.l. species complex is typically attributed to phenotypic plasticity. A combination of herbarium specimen examination and extensive fieldwork led to formation of seven a priori morphological groups, or hypothesized taxa, in the C. reticulata species complex based on previously used and novel morphological characters. A total of 16 morphological characters, quantitative and qualitative, were measured to generate two datasets from herbarium specimens and a common garden experiment which were subjected to the same statistical analyses. To visualize morphological separation in multivariate space, data were subjected to factor analysis of mixed data. To infer if hypothesized taxa were diagnosable, partial least squares discriminant analysis models were used to assess classification accuracy based on a priori groups. Morphological characters were subjected to non-parametric univariate analyses. Of the seven hypothesized taxa, six were supported by evidence from morphology, biogeography, and ecology. Four new species are recognized: C. arenicola sp. nov., C. cumberlandensis sp. nov., C. ouachitensis sp. nov., and C. terminalis sp. nov. Viorna subreticulata is given a new combination in Clematis and the newly defined C. reticulata s.s. is re-circumscribed. An updated dichotomous key, species descriptions, ecological notes, and distribution maps are provided. vii TABLE OF CONTENTS CHAPTER I: A Review of the Systematics and Taxonomy of Clematis subgenus Viorna A. Gray (Ranunculaceae) ......................................................................................................................... 1 Introduction ............................................................................................................................ 1 Nomenclatural History ............................................................................................................ 2 Taxonomic Treatments ........................................................................................................... 3 Palynological Studies .............................................................................................................. 6 Leaf Surface ........................................................................................................................... 6 Seedling Phyllotaxy ................................................................................................................ 7 Cytological Studies ................................................................................................................. 8 Chemotaxonomic Studies........................................................................................................ 9 Molecular Methods ................................................................................................................. 9 Conclusion ............................................................................................................................ 10 Literature Cited ..................................................................................................................... 12 CHAPTER II: A Morphometric Approach to Delimiting Species in the Clematis reticulata Species Complex (Ranunculaceae: Subgenus Viorna) ............................................................... 16 Introduction .......................................................................................................................... 16 Materials and Methods .......................................................................................................... 20 Herbarium specimen sampling .......................................................................................... 20 Common garden sampling................................................................................................. 20 viii Common garden experiment ............................................................................................. 21 Character selection ............................................................................................................ 22 Data subsetting ................................................................................................................. 23 Multivariate analyses ........................................................................................................ 24 Univariate analyses ........................................................................................................... 25 Comparison of herbarium specimens and common garden datasets ................................... 26 Results .................................................................................................................................. 27 Herbarium specimens ........................................................................................................ 27 Common garden experiment ............................................................................................. 28 FAMD .............................................................................................................................. 28 PLS-DA ............................................................................................................................ 30 Univariate analysis of quantitative and ordinal characters.................................................. 31 Nominal characters ........................................................................................................... 34 Comparison of herbarium specimens and common garden datasets ................................... 35 Discussion ............................................................................................................................ 35 Utility of common garden
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