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A Systematic Revision of Passiflora Section Dysosmia (Passifloraceae)

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A Systematic Revision of Passiflora Section Dysosmia (Passifloraceae) A Systematic Revision of Passiflora Section Dysosmia (Passifloraceae) A dissertation presented to the faculty of the College of Arts and Sciences of Ohio University In partial fulfillment of the requirements for the degree of Doctor of Philosophy Harlan T. Svoboda August 2018 © 2018 Harlan T. Svoboda. All Rights Reserved. 2 This dissertation titled A Systematic Revision of Passiflora Section Dysosmia (Passifloraceae) by HARLAN T. SVOBODA has been approved for the Department of Environmental and Plant Biology and the College of Arts and Sciences by Harvey E. Ballard, Jr. Professor of Environmental and Plant Biology Joseph Shields Interim Dean, College of Arts and Sciences 3 ABSTRACT SVOBODA, HARLAN T., Ph.D., August 2018, Environmental and Plant Biology A Systematic Revision of Passiflora Section Dysosmia (Passifloraceae) Director of Dissertation: Harvey E. Ballard, Jr. One of the most taxonomically difficult groups in the passionflower genus (Passiflora, Passifloraceae) is section Dysosmia. For centuries, the taxa belonging to this group have proven difficult to classify or even name. Various authors over the years have attempted to tackle the complex task of describing the diversity in this section and have done so with varying degrees of success; often times shuffling names between species or recognizing taxa at differing ranks. Ellsworth Killip, in his 1938 monograph of the family, introduced a litany of varietal names for one of the most notoriously complicated species, Passiflora foetida, in an attempt to make sense of the staggering degree of morphological variation. His work remains a foundation for the systematics of Dysosmia as well as that of the rest of the genus to this day. The section was most recently revised by John Vanderplank in 2013, who took a more conservative approach by synonymizing many names and renaming others to more accurately reflect our changing taxonomic hypotheses. At the outset of this project 21 species and nine varieties were recognized and accepted in the section. The primary goal of this research was to use modern techniques and provide new evidence to aid in the delimitation of the taxa within section Dysosmia. An exhaustive nomenclatural study was conducted in order to ascertain the extent to which the sectional 4 diversity has already been named. Dozens of names were subsequently typified and corrected in order to bring them into compliance with the current Code of Nomenclature. Intensive morphological studies were performed to investigate the use of macro- and micromorphological features—some of which never before assessed—for the delimitation of taxa. Robust statistical analyses revealed at least three species assemblages or lineages within section Dysosmia that were then the focus of subsequent work, particularly a conspicuous red-fruited complex. Geometric morphometric descriptors were also pursued for the purposes of classification and show promise for objective classification of an otherwise unruly group. Molecular approaches were explored to test the hypotheses of monophyly and relationships within the section but were not successful. Finally, climatic and soil data were employed to investigate a complex of taxa and delimit distinct entities at the rank of species. These data were also informative in describing habitat characteristics of the species. This is the first time that evidence of this kind has been used for taxonomic delimitation in Passiflora. The culmination of this research was a comprehensive synopsis of a portion of the section, comprising all of the red-fruited taxa, in which several new species and combinations were proposed. The current understanding of section Dysosmia brings the number of recognized taxa to 31 species and five putative varieties. 5 DEDICATION This dissertation is dedicated to my partner, Ricky, for his unwavering support and love throughout the adventure of grad school. I also dedicate this to my family who have believed in my every endeavor from the very start. 6 ACKNOWLEDGMENTS First and foremost, I thank my advisor and mentor, Dr. Harvey Ballard, Jr., for his steadfast support throughout every aspect of my development as a researcher, scholar, and educator. I would also like to thank my current committee members, Drs. Morgan Vis, Rebecca Snell, Alycia Stigall, and John MacDougal as well as previous committee members, Drs. Elizabeth Hermsen and Shawn Kuchta. I also thank several Passiflora specialists who have guided and influenced my research, including Drs. Christian Feuillet, Peter Jørgensen, Shawn Krosnick, John MacDougal, and John Vanderplank. I express my gratitude to my current and former labmates: Ben Gahagen, Jen Hastings, Anne Sternberger, Bina Swasta Sitepu, and Bethany Zumwalde for their support. I also appreciate the work of Harold Blazier for growing plants for my research, and to Zach Parinello and Todd Roth for assistance with digitizing and databasing herbarium specimens. I extend my thanks to the Ohio University Graduate Student Senate for funding my research with two Original Work Grants and several Travel Awards to attend scientific meetings, the Department of Environmental and Plant Biology for travel funding, the Ohio University Graduate College for a Student Enhancement Award, and the Ohio Center for Ecology and Evolutionary Studies for a research fellowship. I am indebted to the following herbaria for access to their collections: A, AMD, ASU, B, BAR, BHCB, BM, BOLO, BRIT, C, CAS, CHIC, DES, DUKE, E, ECON, F, FLAS, FTG, G, GENT, GH, GOET, HA, HCIB, HUA, HUEFS, IND, INPA, K, L, LINN, LISC, LSU, MA, MBM, MEXU, MICH, MIN, MISSA, MPU, MO, NY, P, PH, R, RB, RSA, S, SD, SP, TEX-LL, U, UC, UCR, UPCB, UPRRP, US, USF, UTC, VT, W, WAG, and YU. 7 TABLE OF CONTENTS Page Abstract ........................................................................................................................... 3 Dedication ....................................................................................................................... 5 Acknowledgments ........................................................................................................... 6 List of Tables ................................................................................................................ 10 List of Figures ............................................................................................................... 11 Chapter 1: Introduction .................................................................................................. 14 Morphology ............................................................................................................. 16 Current Systematics ................................................................................................. 20 Phylogenetic Relationships ...................................................................................... 25 Geographic Distribution........................................................................................... 26 Chapter 2: Typifications and Nomenclatural Notes in Passiflora Section Dysosmia (Passifloraceae) ............................................................................................................. 28 Abstract ................................................................................................................... 28 Introduction ............................................................................................................. 29 Nomenclatural Treatment ........................................................................................ 30 Typifications ...................................................................................................... 30 Nomenclatural Clarification ............................................................................... 43 Ambiguous Names ............................................................................................. 51 Invalid and Illegitimate Names .......................................................................... 55 Acknowledgements ................................................................................................. 56 Chapter 3: Phenetic and Cladistic Studies Help Clarify Species Assemblages in Passiflora Section Dysosmia (Passifloraceae) ................................................................ 63 Abstract ................................................................................................................... 63 Introduction ............................................................................................................. 64 Methods................................................................................................................... 66 Results and Discussion ............................................................................................ 68 Conclusions ............................................................................................................. 70 Acknowledgements ................................................................................................. 71 Chapter 4: Leaf Geometric Morphometrics .................................................................... 85 Introduction ............................................................................................................. 85 8 Comprehensive Methods for Leaf Geometric Morphometric Analyses ..................... 86 Abstract ............................................................................................................. 87 Background ......................................................................................................
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