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An Integrative Taxonomic Study of Ramps (Allium Tricoccum Aiton) Complex

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An Integrative Taxonomic Study of Ramps (Allium Tricoccum Aiton) Complex An Integrative Taxonomic Study of Ramps (Allium tricoccum Aiton) Complex A thesis presented to the faculty of the College of Arts and Sciences of Ohio University In partial fulfillment of the requirements for the degree Master of Science Bina Swasta Sitepu August 2018 © 2018 Bina Swasta Sitepu. All Rights Reserved. 2 This thesis titled An Integrative Taxonomic Study of Ramps (Allium tricoccum Aiton) Complex by BINA SWASTA SITEPU 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 C. Shields Interim Dean, College of Arts and Sciences 3 ABSTRACT SITEPU, BINA SWASTA, M.S., August 2018, Environmental and Plant Biology An Integrative Taxonomic Study of Ramps (Allium tricoccum) Complex Director of Thesis: Harvey E. Ballard, Jr. The Allium tricoccum complex is native to eastern North America and encompasses broad and confusing morphological variation. Previous studies have led to contrary classifications to account for the diversity of morphologies in the Allium tricoccum complex. Living plants of the complex, leaf tissue samples and soil samples were collected from 28 natural populations in seven states. Plants were cultivated in the Ohio University common garden for biweekly observations of morphological traits; growth patterns and phenology of leaves, flowers and fruits; and weekly photography of plant structures. Morphology, phenology, ecology, and genetic diversity were analyzed to delineate and compare distinct taxa found in the Allium tricoccum complex. Two new characteristics of the species were described for the first time: scape growth direction (orientation) and depth of bulb in the ground. Two major groups in the Allium tricoccum complex, Red Ramps (A. tricoccum sensu stricto) and Green Ramps, were broadly distinguished based on many differences in leaf shape and size; pigmentation of leaf, scape and bulb; number of buds, flowers and fruits; scape growth direction, bulb size and depth in the ground. Three distinctive taxa within the broader Green Ramps group were separated based on differences in morphology, ecology, phenology and geography. The Green Ramps group consisted of A. burdickii (Hanes) G. N. Jones in the Great Lakes and Great Plains regions; a South Green Ramps taxon in the Interior Highlands of Kentucky 4 and Tennessee, similar to A. burdickii but distinct in its tendencies toward narrower leaves, more flowers, shorter perianth and shorter stamens, consistent retention of leaves during scape elongation, and preference for more silty or clayey soils; and a Highland Green Ramps taxon in the Appalachian Mountain region, producing broader distinctly petiolate leaves similar to A. tricoccum, with somewhat intermediate flower and fruit traits between A. burdickii and A. tricoccum, absence of reddish-purple pigmentation, and deep bulb and erect scape characteristic of the other Green Ramps taxa. Microsatellite markers failed to provide genetic differentiation among the four taxa, but did provide separation of populations within each, suggesting that loci adapted from cultivated garlic cultivars may be too variable at the species level for the wild species in the A. tricoccum complex. Corroborative evidence from macromorphology, phenology and ecology, and application of the Unified Species Concept, rejected usage of one or two taxa previously proposed, and instead supported the recognition of four distinct evolutionary species to accommodate diversity in the A. tricoccum complex. 5 DEDICATION Magna Vis Fidelitatis, I dedicate this work for my Lord, owner of the knowledge and my life, I would like to give special thanks for my wife, Agustina Dwi Setyowati, and my family for endless support in my effort to achieve my dream. 6 ACKNOWLEDGMENTS I believe this thesis could be finished as a result of hard work by teamwork, and I am really proud to be part of this team. I express my appreciation to my advisor, Prof. Dr. Harvey E. Ballard, Jr., for his support, advice, and scientific expertise. I would also like to thank my committee members, Dr. Arthur Trese and Dr. Rebecca Snell, for their assistance and valuable input for my research and thesis. Special thanks to Dr. Rebecca Snell for her help in statistics and Species Distribution Modelling. Also, I give my thanks to Dr. Jared DeForest for help in the soil analysis and access to the Soil Analysis room, Prof. Brian C. McCarthy, Ph.D., for granting me access to use his lab and equipment for soil analysis, and Prof. Dr. Morgan L. Vis, for the kindness to provide some lab supplies. Thank you to Harlan Svoboda, Anne Sternberger, and Jennifer Hastings, for friendship and endless support during my research. I thank Ohio University and the Graduate Student Senate (for an Original Work Grant) at Ohio University for providing funding for my research. I would like to thank those who have contributed their various time to my research: Danny Wolf, Colin Kruse, Proma Basu, Yemi Ajayi, Dasmeet Kaur, Tasleem Javaid, and Marion Holmes. Thank you to the USAID-Prestasi Program for the scholarship and research funding. Additional thanks to botanists in the Tennessee and Kentucky Heritage Programs for providing Ramps population information, and Bernheim Arboretum and Research Forest in Kentucky and Eagle Crest Nature Preserve in Indiana for permission and assistance in conducting field work on their lands. Dr. Alan S. Weakley and Tom Govus provided me invaluable discussion and information about 7 Ramps in the southern Appalachian region. I am indebted to everybody who has taken part in this effort to contribute this new information for science and yield a small colored point in the ocean of knowledge. 8 TABLE OF CONTENTS Page Abstract ............................................................................................................................... 3 Dedication ........................................................................................................................... 5 Acknowledgments............................................................................................................... 6 List of Tables .................................................................................................................... 10 List of Figures ................................................................................................................... 11 Chapter 1: Introduction ..................................................................................................... 13 Taxonomic History ..................................................................................................... 17 Types of Data Utilized ................................................................................................ 21 Objectives ................................................................................................................... 24 Broader Impacts .......................................................................................................... 25 Chapter 2: Morphology ..................................................................................................... 26 Methods....................................................................................................................... 26 Results ......................................................................................................................... 34 Discussion ................................................................................................................... 44 Chapter 3: Phenology ........................................................................................................ 50 Methods....................................................................................................................... 50 Results ......................................................................................................................... 52 Chapter 4: Environmental Factors .................................................................................... 62 Methods....................................................................................................................... 62 Results ......................................................................................................................... 63 Discussion ................................................................................................................... 68 Chapter 5: Molecular Analysis ......................................................................................... 72 Methods....................................................................................................................... 72 Results ......................................................................................................................... 73 Discussion ................................................................................................................... 75 9 Chapter 6: Species Distribution Models ........................................................................... 77 Methods....................................................................................................................... 77 Results ......................................................................................................................... 78 Discussion ................................................................................................................... 84 Chapter 7: Systematic Review .........................................................................................
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