EVOLUTIONARY HISTORY OF SIMAROUBACEAE (SAPINDALES): SYSTEMATICS, BIOGEOGRAPHY AND DIVERSIFICATION By JOSHUA WILLIAM CLAYTON 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 2008 1 © 2008 Joshua William Clayton 2 To WilsonPhillips-X, for all the good times 3 ACKNOWLEDGMENTS I would like to thank my committee, Doug Soltis, Pam Soltis, Walter Judd, Steve Manchester, and Gustav Paulay. I am very grateful to everyone who provided material for this study: Christopher Quinn at the Sydney Botanic Gardens, Frieda Billiet at the National Botanic Garden of Belgium, Libby MacMillan at the Royal Botanic Gardens, Edinburgh, Andrew Ford at CSIRO, Bruce Wannan of EPA, Queensland, Richard Abbott at FLMNH, Wang Hong at the Shishuangbanna Tropical Botanical Garden, China, Serena Lee at Singapore Botanic Gardens, Kew DNA Bank, MOBOT DNA Bank, and curators of MO, E, AAU, WAG, NY and CAY. Thanks also to Rick Ree (Field Museum, Chicago, USA) and Stephen Smith (Yale University, USA) for help with Lagrange, Alexei Drummond for comments regarding BEAST, Aleksej Hvalj (Komorov Institute, St. Petersburg, Russia) for providing information on Leitneria fossils, and Wayt Thomas, Michael Moore, Scot Kelchner, Susanne Renner and three anonymous reviewers for their helpful comments. Funding was provided by the National Science Foundation (angiosperm AtoL EF-0431266 to DES, PSS et al.; DDIG DEB-0710202 to JWC, DES), Botanical Society of America, and American Society of Plant Taxonomists. Finally, I would like to thank all my friends in the Soltis Lab, Crysta, Andrew, and my family, for their support. 4 TABLE OF CONTENTS page ACKNOWLEDGMENTS ...............................................................................................................4 LIST OF TABLES ...........................................................................................................................8 LIST OF FIGURES .........................................................................................................................9 ABSTRACT ...................................................................................................................................12 CHAPTER 1 INTRODUCTION ..................................................................................................................14 2 OVERVIEW OF SIMAROUBACEAE (SAPINDALES) .....................................................16 Family Description .................................................................................................................16 Vegetative Morphology ..........................................................................................................17 Vegetative Anatomy ...............................................................................................................18 Inflorescence Structure ...........................................................................................................18 Flower Structure .....................................................................................................................19 Embryology ............................................................................................................................20 Pollen Morphology .................................................................................................................20 Karyology ...............................................................................................................................21 Reproductive Biology .............................................................................................................21 Fruits and Seeds ......................................................................................................................21 Dispersal .................................................................................................................................23 Phytochemistry .......................................................................................................................23 Distribution and Habitats ........................................................................................................24 Fossil History ..........................................................................................................................25 Affinities .................................................................................................................................25 Relationships within the Family .............................................................................................26 Uses and Economic Importance .............................................................................................27 Key to the New World Genera ...............................................................................................27 Key to the Old World Genera .................................................................................................28 Descriptions of Genera ...........................................................................................................30 1. Picrasma Blume ...................................................................................................30 2. Castela Turpin ......................................................................................................31 3. Holacantha A.Gray ...............................................................................................31 4. Ailanthus Desf. .....................................................................................................32 5. Leitneria Chapm. ..................................................................................................32 6. Soulamea Lam. .....................................................................................................33 7. Amaroria A.Gray ..................................................................................................33 8. Brucea J.F.Mill .....................................................................................................34 9. Laumoniera Noot. .................................................................................................34 10. Nothospondias Engl. .............................................................................................35 5 11. Picrolemma Hook.f. ..............................................................................................35 12. Quassia L. .............................................................................................................36 13. Samadera Gaertn. .................................................................................................36 14. Eurycoma Jack ......................................................................................................37 15. Gymnostemon Aubrév. & Pellegr. ........................................................................37 16. Perriera Courchet .................................................................................................38 17. Hannoa Planch. .....................................................................................................38 18. Odyendea (Pierre) Engl. .......................................................................................39 19. Iridosma Aubrév. & Pellegr. ................................................................................39 20. Pierreodendron A.Chev. ......................................................................................39 21. Simarouba Aubl. ...................................................................................................40 22. Simaba Aubl. ........................................................................................................41 3 MOLECULAR PHYLOGENY OF SIMAROUBACEAE BASED ON CHLOROPLAST AND NUCLEAR MARKERS ..................................................................42 Introduction .............................................................................................................................42 Methods ..................................................................................................................................45 Taxon Sampling ...............................................................................................................45 DNA Extraction, Amplification and Sequencing ............................................................46 Alignment and Indel Coding ...........................................................................................47 Maximum Parsimony Analyses and Data Congruence ...................................................48 Bayesian Analyses and Partitioning Strategies ...............................................................49 Results .....................................................................................................................................51 Data Congruence .............................................................................................................51 Phylogenetic Analyses and Tree Topology .....................................................................52 Partitioning Strategies .....................................................................................................53 Discussion ...............................................................................................................................54 Data Congruence and Partitioning Strategies ..................................................................54