See discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/311541611 Systematics and Evolution of Menyanthaceae and the Floating-Leaved Genus Nymphoides Article · January 2010 CITATIONS READS 0 212 1 author: Nicholas P. Tippery University of Wisconsin - Whitewater 92 PUBLICATIONS 645 CITATIONS SEE PROFILE Some of the authors of this publication are also working on these related projects: Aquatic plant systematics View project Molecular taxonomy of Hyrcanian Alnus using nuclear ribosomal ITS and chloroplast trnH-psbA DNA barcode markers View project All content following this page was uploaded by Nicholas P. Tippery on 11 July 2017. The user has requested enhancement of the downloaded file. Systematics and Evolution of Menyanthaceae and the Floating-Leaved Genus Nymphoides Nicholas Peter Tippery, Ph.D. University of Connecticut, 2010 Menyanthaceae (70 species) are a family of aquatic and wetland plants that occur worldwide. This diverse group contains different growth habits (emergent and floating-leaved), reproductive systems (heterostyly, homostyly, gynodioecy and dioecy), floral and seed morphologies, and inflorescence architectures. In this study, I have evaluated the phylogenetic relationships, taxonomy, biogeography, and morphological character evolution for over half of the approximately 50 Nymphoides species, plus all species in related genera within the family. In Chapter 1 I investigated generic relationships across Menyanthaceae and found that the contemporary circumscription of Villarsia included three paraphyletic lineages that graded toward a monophyletic Nymphoides. Biogeographical reconstruction supported an Australian origin for the family and also for all of the major clades, with dispersal events corresponding to the boreal sister taxa Menyanthes and Nephrophyllidium, and the South African Villarsia clade. Chapter 2 is a study of the genus Nymphoides that examined all Australian species of the genus and synthesized for the first time the morphological data for all the species worldwide. Morphological data analysis indicated several relationships, including the grouping of species with similar inflorescence habits. Molecular phylogenetic analyses supported a similar division, but species resolution was thoroughly incongruent on trees derived from nuclear (nrITS) or chloroplast (matK/trnK) data, suggesting widespread hybridization during the diversification of the genus. In Chapter 3 I studied the Nymphoides inflorescence architecture, which comprises three types: expanded (pairs of flowers separated by internodes), condensed (single floating leaves each supporting a cluster of flowers), and a morphology unique to N. peltata (pairs of leaves supporting clusters of flowers). I determined that these quite different inflorescence types likely were derived from a common blueprint, from which they deviate only by their relative elongation of internodes or expansion of bracts into foliage leaves. In Chapter 4 I examined the life history and reproductive potential of N. peltata, a Eurasian native that is naturalized in North America. Populations of N. peltata are able to produce abundant fruits with highly germinable seeds, but the plants are genetically identical throughout their naturalized range, possibly the result of inbreeding while in cultivation. Nicholas Peter Tippery – University of Connecticut, 2010 Systematics and Evolution of Menyanthaceae and the Floating-Leaved Genus Nymphoides Nicholas Peter Tippery B.A., University of Dallas, 2000 A Dissertation Submitted in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy at the University of Connecticut 2010 Copyright by Nicholas Peter Tippery 2010 i APPROVAL PAGE Doctor of Philosophy Dissertation Systematics and Evolution of Menyanthaceae and the Floating-Leaved Genus Nymphoides Presented by Nicholas Peter Tippery, B.A. Major Advisor ___________________________________________________ Donald H. Les Associate Advisor ________________________________________________ Gregory J. Anderson Associate Advisor ________________________________________________ Cynthia S. Jones University of Connecticut 2010 ii Dedicated to my father, Kevin Tippery, who introduced me to the beauty and perpetual wonder of nature, who taught me to persevere through adversity, who encouraged me to be active and enjoy life, and who always supported me in whatever I did. iii ACKNOWLEDGEMENTS I could not have gotten even close to the point of finishing my Ph.D. without the continuing support of so many people. I am grateful foremost to my advisor Don Les, who was my guide through the graduate school experience, who taught me by example and by gentle correction, who provided me with laboratory resources whenever I could not support myself, and who generously included me on his research to help me become well published. I thank also my advisory commitee, who helped to round out my education with their phenomenal experience in their respective areas of expertise. Greg Anderson, whose friendliness knows no equal, was always encouraging and supportive, and he was a valuable resource for helping me think through pollination biology and reproductive system evolutionary concerns. Cindi Jones, who knows all the inner workings of plant anatomy and morphology, was very generous with her time and resources in allowing me to add the morphological component to my dissertation, and her enthusiasm continues to inspire me. I had a number of professional acquatintances who helped to further my development as a scientist. I could not thank Surrey Jacobs enough for sharing two months of his life with me and instructing me in everything from Australian customs to good plant pressing technique. Surrey took my poorly conceived plan to collect Nymphoides across Australia and turned it into a workable project that became rousingly successful. Surrey and Betty Jacobs made Australia a second home for me, and I am forever grateful to have met and shared time with them. Surrey‘s friends and colleagues also helped greatly to make our trip more successful and enjoyable. I thank especially John and Marion Clarkson for opening their home to us, and I was privileged also to meet Helen Aston, Janice and Roger Carolin, Geoff Sainty, Lance Smith, and Karen Wilson while in Australia. Walter Pagels was a steady source of plant material, knowlege and inquiry about Menyanthaceae, and I continue to try to repay him for helping to get me started. I relied heavily on the information gained from herbarium loans, for which I thank the curators of iv the CONN herbarium, Andrew Doran and Bob Capers, and the directors and staff of the many herbaria that kindly sent loans for my research. The graduate students of the UConn EEB Department were my constant support, both emotionally and academically. I enjoyed having made so many kind friends here, and I hope to carry them with me through the rest of my life. I could scarcely name them all, and I deeply regret that I cannot list everyone here, but some people who were particularly helpful include Lori Benoit, Jessica Budke, Jane Carlson, Karolina Fučíková, Laura Forrest, Geert Goemans, Chris Martine, Hilary McManus, Michael Moody, Brigid O‘Donnell, Rachel Prunier, Krissa Skogen, Frank Smith, Kathryn Theiss, Dan Vanderpool, Juan Carlos Villarreal, Amy Weiss, and Norm Wickett. Without these and so many other inspirational people, I would have abandoned hope long ago. In addition to fellow EEB graduate students, I had many friends help me through their personal support, many of whom only tangentially knew what sort of research I was pursuing. These include Dan and Mary Bernier, Russ Bird, Kate and Matt Gilmore, Jeff Kotz, Kate O‘Sullivan, Nicole and Pete Palumbo, Marissa Prosser, Anna and Scott Russell, Emily Simmons, and Liz Werle. I thank them for keeping me grounded and helping me to have fun in the midst of my most serious times. My family has always been there to support me, and they helped me to become the kind of person who could successfully finish a Ph.D. My mom Kitty has always been the rock of our family, keeping us on the straight and narrow, motivating us to be better people, and always presenting a positive outlook on life. My sister Janie is always optimistic and supportive, and there is nothing she wouldn‘t do to help out a friend. My brother Chris helped to raise me by his example, and he was the constant companion of my youth whom I could always rely on to laugh at my jokes. In our adult years, he has continued to support me, and I treasure the time I have been able to spend with him, his wife Allison, and their lovely children. v During the majority of my time at UConn, I have been privileged to have the love and companionship of my girlfriend Megan Johnston. She has helped me to see the end goal through the whole long process of graduate school, and she has made my experience vastly more enjoyable. Megan has supported my role as a graduate student far and above what should be expected of any partner, and at times she was an even stronger motivator than I was for my own work. Her patient support was really one of the major reasons why I was able to finish. vi TABLE OF CONTENTS Chapter 1: Systematics of Menyanthaceae Abstract................................................................................................................................1 Introduction..........................................................................................................................1 Materials and