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Rosaceae), with Emphasis on the Pleistocene Radiation of the High Andean Genus Polylepis

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Rosaceae), with Emphasis on the Pleistocene Radiation of the High Andean Genus Polylepis ABSTRACT Title of dissertation: A PHYLOGENETIC AND BIOGEOGRAPHIC ANALYSIS OF SANGUISORBEAE (ROSACEAE), WITH EMPHASIS ON THE PLEISTOCENE RADIATION OF THE HIGH ANDEAN GENUS POLYLEPIS. Malin Sofia Kerr, Doctor of Philosophy, 2004 Dissertation directed by: Dr. Charles F. Delwiche Dr. James L. Reveal A phylogenetic and biogeographic analysis of the tribe Sanguisorbeae (Rosaceae) was conducted with emphasis on the radiation of the Andean tree Polylepis. Phylogenetic analyses of coding and non-coding nuclear markers reveal a complex evolutionary history of the tribe including ancient and recent allopolyploid hybridization. Sanguisorba sensu lato is shown to be paraphyletic and split between the allopolyploid hybrid Sanguisorba and the non-hybrid Poterium and Poteridium. A monophyletic origin of the southern hemispheric subtribe Sanguisorbinae is supported, and this clade is given a phylogentic taxon name (Verruchaena). Dating analyses using the penalized likelihood method suggest that this taxon originated in the late Miocene. A biogeographic hypothesis is presented in which Verruchaena originated in the New World with subsequent transoceanic dispersals to southern Africa and Australasia. The paramo genus Polylepis most likely arose from hybridization between two Andean ancestors supporting a “vertical” rather than “horizontal” origin of this taxon. A PHYLOGENETIC AND BIOGEOGRAPHIC ANALYSIS OF SANGUISORBEAE (ROSACEAE), WITH EMPHASIS ON THE PLEISTOCENE RADIATION OF THE HIGH ANDEAN GENUS POLYLEPIS. by Malin Sofia Kerr Dissertation submitted to the Faculty of the Graduate School of the University of Maryland, College Park in partial fulfillment of the requirements for the degree of Doctor of Philosophy 2004 Advisory Committee: Associate Professor Charles F. Delwiche, Chair/Advisor Research Associate Torsten Eriksson Associate Professor Irwin N. Forseth, Jr. Associate Professor Patrick S. Herendeen Professor Charles Mitter Copyright by Malin Sofia Kerr 2004 AKNOWLEDGEMENTS I would like to thank Dr. James L. Reveal, who was my primary advisor for two years before his retirement, for enthusiastically sharing his immense expertise in classical plant taxonomy and nomenclature. The educational foundation provided by Dr. Reveal was perfectly complemented by the guidance and support I received from Dr. Charles F. Delwiche during the following four years, primarily in the field of molecular phylogenetics. I am grateful to Dr. Delwiche for giving me the confidence and freedom to allow my research to develop in new directions. My dissertation research benefited greatly from the daily interactions I had in the Delwiche lab with students and staff. I am particularly grateful to Tsetso Bachvaroff, Mathew Cimino, John Hall, Ken Karol, Jeff Lewandowski, Tanya Marushak and Maria Virginia Sanchez Puerta for support and inspiration. I would also like to thank Dr. Torsten Eriksson for introducing me to my study group, the Sanguisorbeae, and for countless inspiring discussions on Rosaceae evolution and on systematics in general. I am also greatly indebted to several people who generously shared rare plant material with me, in particular Michael Kessler (Universität Göttingen), Christopher Whitehouse (University of Cape Town), and John Clark (George Washington University). In addition I would like to acknowledge Daniel Larrea and Mónica Moraes R. (National Herbarium of Bolivia) for their valuable assistance, making collecting in Bolivia a very pleasant and productive experience. ii The following institutions generously provided herbarium and/or fresh garden material for study: The Royal Botanic Garden Edinburgh, the New York Botanical Garden, the Missouri Botanical Garden, the Gray Herbarium (Harvard University), and the National Herbarium at the Smithsonian. My dissertation research was funded by grants from the National Science Foundation (Doctoral Dissertation Improvement Grant), the American Society of Plant Taxonomists, and the Department of Cell Biology and Molecular Evolution at the University of Maryland. Finally I would like to acknowledge my Swedish family who cheered me on despite the geographic distance, and most of all I want to thank my husband Dr. Peter H. Kerr who shared with me this experience from start to finish as we entered graduate school on the same day and defended our dissertations in the same week. Without Peter I would have never started on this path in the first place and undoubtedly would not have had the endurance to see it through. iii TABLE OF CONTENTS Chapter I General Introduction ..................................................................................... 1 Introduction .............................................................................................................. 1 Morphology, ecology and distribution ..................................................................... 3 Agrimoniinae....................................................................................................... 4 Sanguisorbinae .................................................................................................... 4 Polylepis .............................................................................................................. 6 Previous taxonomic treatments of the Sanguisorbeae and Polylepis ....................... 7 Sanguisorba......................................................................................................... 8 Cliffortia and Acaena .......................................................................................... 9 Polylepis ............................................................................................................ 10 Previous phylogenetic studies of Sanguisorbeae.................................................... 11 Species recognition in Polylepis............................................................................. 13 Objectives and significance.................................................................................... 13 Objectives .......................................................................................................... 13 Significance ....................................................................................................... 14 Chapter II Phylogeny and character evolution in the Sanguisorbeae inferred from chloroplast sequence data ........................................................................................... 16 Introduction ............................................................................................................ 16 Exploratory morphological analysis.................................................................. 17 Materials and methods............................................................................................ 18 Taxon sampling ................................................................................................. 18 Phylogenetic marker.......................................................................................... 24 DNA extraction ................................................................................................. 24 PCR Amplification and Sequencing.................................................................. 25 Phylogenetic methods........................................................................................ 27 Results .................................................................................................................... 28 Phylogenetic nomenclature and the application of taxon names....................... 35 Discussion .............................................................................................................. 37 Sanguisorba and Poterium ................................................................................ 40 Verruchaena ...................................................................................................... 43 Acaena and Margyricarpus/Tetraglochin ......................................................... 44 Polylepis ............................................................................................................ 47 Character evolution ................................................................................................ 48 Floral evolution ................................................................................................. 48 Evolution of habit .............................................................................................. 52 Chapter III Phylogenetic analyses of the Sanguisorbeae based on duplicated polymorphic nuclear sequence data............................................................................ 54 Introduction ............................................................................................................ 54 Polyploidy and gene duplication in Sanguisorbeae........................................... 55 Materials and methods............................................................................................ 57 Taxon sampling ................................................................................................. 57 Phylogenetic markers ........................................................................................ 60 iv DNA extraction, PCR amplification and sequencing........................................ 65 Phylogenetic methods........................................................................................ 68 Results ...................................................................................................................
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