Copyright by Martin Enrique Timaná 2005 The Dissertation Committee for Martin Enrique Timaná Certifies that this is the approved version of the following dissertation: SYSTEMATIC STUDIES IN PYCNOPHYLLUM AND PYCNOPHYLLOPSIS (CARYOPHYLLACEAE) OF THE HIGH ANDES Committee: ______________________________ Beryl B. Simpson, Supervisor ______________________________ David Cannatella ______________________________ Robert K. Jansen ______________________________ Tom J. Mabry ______________________________ Jose L. Panero SYSTEMATIC STUDIES IN PYCNOPHYLLUM AND PYCNOPHYLLOPSIS (CARYOPHYLLACEAE) OF THE HIGH ANDES by Martin Enrique Timaná, B.Sc.; M.Sc. Dissertation Presented to the Faculty of the Graduate School of The University of Texas at Austin in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy The University of Texas at Austin May, 2005 Dedication To my son, Leonardo Martin, miracle child and first Texan in the family, For offering me a smile every single day since birth, And to my wife, Aleyda My friend and companion for many years, For waiting this long It was the best of times, it was the worst of times… Charles Dickens, A Tale of Two Cities For sudden, the worst turns the best to the brave… Robert Browing, Prospice Acknowledgements This Dissertation represents the work of several years as a graduate student at the University of Texas. I would like to thank first and above all my graduate advisor, Dr. Beryl B. Simpson, for her infinite patience and support and the intellectual freedom I received from her during all these years. Thank you, Beryl, with all my heart for believing in me. Based on reading my fellow graduate students’ dissertations, it seems to me that this Acknowledgements section is probably the only segment where (unless I’m told otherwise) I have almost all the freedom to express my personal thoughts in my own style, without either being subject to the approval (or disapproval) of members of my dissertation committee or constrained by the stiff and unattractive style of academic prose. And that is a good thing. This incredible journey toward a Ph.D. in Botany started on the afternoon of July 30th, 1984, when my undergraduate advisor, mentor and close friend, Dr. Carlos Reynel, invited me to join him as a student assistant in the “Herbario Forestal” at the Universidad Nacional Agraria, La Molina, in Lima, Peru. Since that day my life has been, in one way or another, linked to the pleasures (and displeasures) of Linnaeus’ Scientia amabilis. Thanks to Carlos, I not only discovered the basics of tropical botany, but became a part of a network of researchers working in the tropics, including the late Alwyn Gentry. Although I wish I could thank everyone involved in my academic formation since that particular day, at least I wanted to include my very first mentor, Carlitos. Since the time I left Peru, 13 years ago, until I moved to Austin to start my doctoral education, several people contributed either in great or small part to the vi development of my ideas. Dr. Alwyn Gentry was one of them. For many years, when still intellectually ‘green’, I had assumed that all botanists had the type of life and line of work that Al had, that is, going to remote, exotic lands (although I never understood why Peru would be exotic, given the fact that I grew up there), collecting non-stop, preparing floras and monographs. I don’t recall ever hearing terms such as “phylogeny” or “autapomorphy” from Al. In any case, his influence also determined in great part the type of botany I wanted to do. In this regard, I was also equally influenced by Dr. Bill R. Anderson, my advisor at the University of Michigan in Ann Arbor. If I had to quote a single sentence that had a tremendous effect in determining my research goals, it was when Bill said to me: “in a hundred years people will be still reading my monographs.” After that there was no turning point. I had to do a monograph. In Austin, in addition to my advisor, Beryl Simpson, I must give an eternal “Thank You” to Drs. Paul and Greta Fryxell. They have been almost like parents to me (and later to my wife, Aleyda). I thank you both for your support in every aspect of my life, not only academic, but also during some of the hardest times at various moments of my days in Texas. Thank you Greta, for all your care during the days of my cancer treatment and for being there, whenever I was in need. Sometimes, not being an English native speaker limits our ability to fully express our thoughts. This is one of those occasions. My “Texan experience” was early affected by the unfortunate fact of having suffered from a rare form of cancer called seminoma (probably so rare I usually have to explain my doctors what this cancer is all about), in 1999. After the corresponding surgery I underwent various months of chemotherapy. And it was during this time, that I witnessed and benefited from some of the most vii beautiful expressions of support, camaraderie, and compassion toward the needy one the American people are capable of. Here I want to thank first to Xochitl Munn, who spent a full week taking care of me after my open chest surgery, Beryl Simpson, for organizing a whole team of volunteers ready to drive me back and forth to the oncology clinic for my daily session of chemotherapy, Paul and Greta Fryxell, for their care and love, and various of my fellow graduate students, including Jennifer Tate, Andrea Weeks, Katie Hansen, Megan Helfgott, Heidi Meudt, among others for making sure I had from fresh meals to movies to watch while confined to my apartment. Since I was still single and living alone back then, I don’t think I could have made it without your help. Thank you to you all. For one of those strange circumstances that certainly confirm the concept of probabilities, the day I was diagnosed with cancer I also received news of having been awarded the first Latin American Research Fellowship, offered by the Royal Botanic Gardens, Kew, to spend four months in Europe. What are the odds? I was fortunate that my treatment was completed a few weeks before I was due in London. As somebody who is in love with the Victorian tradition of natural history, this fellowship represented my own “gentleman’s tour” as experienced by many 19th century botanists, including Asa Gray and Joseph Hooker, to name a couple. It was the journey I had planned to do since I was maybe 14 years old. While this is not the place to go into the details of this experience, it also allowed me to meet my good friend and fellow Argentinian botanist, Dr. Jorge Chiapella. Even though we overlapped in London for only a week and only shared the bill of some cheap English breakfast (used for dinner) and some dark Irish beer in a smoky London pub six years ago, Jorge has offered viii me since then his friendship, words of advice, and support either from Vienna or Cordoba. It is indeed such a relief to find out that there is somebody as pessimistic about the future as oneself. Thank you Jorge,and I’m sorry I never shared your enthusiasm for Borges, grasses, and your vinchucas. Graduate school is also supposed to be about graduate life. And although the life of a herbarium taxonomist is almost monastic and in constant isolation, over the years I had the joy of sharing some time with great graduate fellows, who not only made my life more interesting by giving me another glimpse of the American people, but more important, because they were there, when I needed their help and support, from simple things such as a sequencing nightmares, to more relevant aspects of life. Thus I want to thank Heidi Meudt, for sharing her interest in southern biogeography, Megan Helfgott, who introduced me to the intimidating world of DNA systematics, Jennifer Tate, for being funny, and being my travel companion across the South American Andes, Kirsten Triplet, for the long conversations about English culture, Shakespeare, books, and everything that had nothing to do with botany, Katie Hansen, for sharing her experiences as a parent. And of course, there is Leah Larkin. My good friend Leah is truly a rare species (in the good sense); over the years I (and others) witnessed her metamorphosis from a leather-bound road biker to a loving mother and wife, but her essence has remained the same – a person with a deep sense of loyalty, friendship, and work ethic; she is also a genius in multiple aspects of modern phylogenetic systematics and without her help in even the most basic aspects, I would have never completed this work. Thank you Leah for your support, but above all, thank you for your friendship. ix Much of the training of a traditional taxonomist comes from what he or she can learn from senior botanists. In that aspect, I have been fortunate to have access to some great “traditional” botanists here in Austin; in addition to my advisor, Beryl Simpson, I received much help from Dr. Paul Fryxell and Dr. Tom Wendt. Both have always showed their willingness to help me with my constant taxonomic and nomenclatural questions and doubts. From the most basic ones, such as: “what if I annotate a type specimen but years later I change my mind?” - Tom’s answer: “we won’t tell your biographers about it,” to more fundamental questions regarding species concepts, the Code, and lectotypification issues. I also want to thank Tom Wendt for all the logistic support involved in obtaining specimen loans and for allowing me to, quietly, expand my domains on the second floor of the herbarium.