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University of Florida Thesis Or Dissertation Formatting EVOLUTION AND DEVELOPMENT OF PETALS IN AIZOACEAE (CARYOPHYLLALES) By SAMUEL FRASER BROCKINGTON A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY UNIVERSITY OF FLORIDA 2009 1 © 2009 Samuel Fraser Brockington 2 To my ma and pa 3 ACKNOWLEDGMENTS I acknowledge the assistance of numerous people and organizations that have contributed to this doctoral work. First and foremost, I thank my advisors Douglas and Pamela Soltis for their patient support and example. I thank Walter S. Judd for inspiring my interest in the evolution of angiosperms and for his insight into various aspects of this project. I thank David Oppenheimer and Paris Gray for their expertise and use of their in-situ facilities. I thank Elena M. Kramer for her advice and expertise at various stages of this project. The morphological development work would not have been possible without the help of Paula J Rudall at the Royal Botanic Gardens, Kew and Michael Frohlich at the Natural History Museum, London. I thank my supervisory committee for taking the time to read this dissertation. I thank Mark Whitten for taking so many superb photos and sharing his photographic equipment. Kurt Neubig kindly shared equipment. I thank my two undergraduate helpers Alexandre Roolse and Jeremy Ramdial for putting in some long hours and collecting substantial amounts of data. The research was funded by an Assembling the Tree of Life grant EF-0431266 (NSF) to D. E. Soltis and P. S. Soltis, Floral Genome Project NSF grant PGR-0115684 to D. E. Soltis and P. S. Soltis, and a DDIG grant DEB-0808342 (NSF) to D. E. Soltis and S. F. Brockington, and a Botanical Society of America Karling Award to S. F. Brockington. Finally I thank my friends in Florida and England, and my siblings Dan, Alice and Grace for their endless support and encouragement. Above all I thank my parents for their love and dedication over so many years. 4 TABLE OF CONTENTS page ACKNOWLEDGMENTS.................................................................................................................... 4 LIST OF TABLES................................................................................................................................ 9 LIST OF FIGURES ............................................................................................................................ 11 LIST OF ABBREVIATIONS ............................................................................................................ 13 ABSTRACT ........................................................................................................................................ 14 CHAPTER 1 INTRODUCTION....................................................................................................................... 16 The Angiosperm Perianth as a System for Evolutionary Developmental Biology (Evo-Devo) ....................................................................................................................... 16 An Evo-Devo Framework for the Study of Petal Evolution ............................................. 17 Caryophyllales: A Model Clade for the Study of Perianth Evolution .............................. 18 Dissertation Outline: Developing the Caryophyllales as a System for Evo-Devo Research............................................................................................................................ 18 2 PHYLOGENY OF THE CARYOPHYLLALES SENSU LATO: REVISITING HYPOTHESES ON POLLINATION BIOLOGY AND PERIANTH DIFFERENTIATION IN THE CORE CARYOPHYLLALES ............................................... 20 Introduction ................................................................................................................................. 21 Materials and Methods ................................................................................................................ 26 Taxon Sampling ................................................................................................................... 26 DNA Isolation and Amplification ...................................................................................... 27 Alignment and Phylogenetic Analysis ............................................................................... 28 Character Reconstructions .................................................................................................. 30 Results .......................................................................................................................................... 34 Individual Plastid Single Copy Data Sets .......................................................................... 34 Inverted Repeat Data Set ..................................................................................................... 35 Combined Plastid Genes from the Single Copy Region ................................................... 35 Combined Plastid Single Copy and Nuclear Genes .......................................................... 36 Total Evidence Data Set ...................................................................................................... 36 Discussion .................................................................................................................................... 37 Phylogenetic Analyses ........................................................................................................ 38 Reconstruction of Pollination Mechanism ......................................................................... 42 Reconstruction of Perianth Differentiation ........................................................................ 44 Multiple Origins of Perianth Differentiation ..................................................................... 46 Recruitment of Preceding Bracts ........................................................................................ 47 Petaloid Modification of the Androecium.......................................................................... 50 5 Differentiation of Homologous Perianth Parts .................................................................. 51 Caryophyllales as a System for Floral Evo-Devo.............................................................. 52 3 LABILE EVOLUTION OF PIGMENTATION IN THE CORE CARYOPHYLLALES ..... 85 Introduction ................................................................................................................................. 86 The Unique Origin of Betalains .......................................................................................... 87 The Intercalation of Betalain and Anthocyanin Lineages ................................................. 88 Mutual Exclusion of Anthocyanins and Betalains............................................................. 89 Biosynthetic Inhibition ........................................................................................................ 89 Materials And Methods .............................................................................................................. 91 Taxon Sampling and Data Assembly ................................................................................. 91 Phylogenetic Analyses ........................................................................................................ 91 Character Coding ................................................................................................................. 93 Character-state Reconstruction ........................................................................................... 93 Stochastic Mapping ............................................................................................................. 94 Prior Specification ............................................................................................................... 94 Rates of Transformation and Ancestral State Reconstruction .......................................... 95 Results .......................................................................................................................................... 96 Phylogenetic Relationships ................................................................................................. 96 Character Mapping with Unordered Coding ...................................................................... 97 Character Mapping with Ordered Coding .......................................................................... 98 Discussion .................................................................................................................................... 99 Polyphyletic Molluginaceae ................................................................................................ 99 Early-Diverging Anthocyanic Lineages ........................................................................... 100 Derived Anthocyanic Lineages ......................................................................................... 101 Evidence for Reversals to Anthocyanin Pigmentation .................................................... 102 Effect of Ordered Character Coding - Support for Ehrendorfer (1976. ......................... 103 The Occurrence of Unpigmented Lineages...................................................................... 104 Multiple Origins of Betalain Pigmentation ...................................................................... 105 Conclusion ................................................................................................................................
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