EVOLUTIONARY RELATIONSHIPS IN POLYGONACEAE WITH EMPHASIS ON TRIPLARIS BY ADRIANA SANCHEZ A Dissertation Submitted to the Graduate Faculty of WAKE FOREST UNIVESITY GRADUATE SCHOOL OF ARTS AND SCIENCES in Partial Fulfillment of the Requirements for the Degree of DOCTOR OF PHILOSOPHY Biology May 2011 Winston-Salem, North Carolina Approved By: Kathleen A. Kron, Ph.D., Advisor Fabián A. Michelangeli, Ph.D., Chairman Miles R. Silman, Ph.D. William K. Smith, Ph.D. Brian W. Tague, Ph.D. ACKNOWLEDGEMENTS This dissertation would not have been possible without the support, mentoring, and help of many people. I would first like to thank my family and my husband Benoit for their constant support and patience. Sometimes it was difficult to be away for many months at a time, but Ben was an incredible support and I am grateful he went through this journey with me. I thank Wake Forest University for the opportunity to be at this incredible institution for so many years. I also want to thank my advisor, Kathleen A. Kron, for her support through my entire doctorate, because her door was always open when I needed her, and for her advice both professional and personal. I am very grateful for her patience and time (especially when editing my manuscripts) and for teaching me how to be a better scientist. Fabian A. Michelangeli was a constant source of support and advice, and he helped me with all the contact for my field work; without his help I would not have been successful in my trips. Thanks to my committee, Dr. William K. Smith, Miles R. Silman, and Brian W. Tague, for their help when I needed it. I would also like to thank my undergraduate and master‘s advisor, Santiago Madriñán; without his guidance and formation I would not have continued my career as a scientist. I had constant support from my labmates: Catherine M. Bush, Tanja M. Schuster, and Sabrina Setaro. They made my days in the lab more enjoyable and they were a continuous source of knowledge and help. I would also like to acknowledge all the people that helped me collecting in the field trips. At times it was difficult to have a positive attitude with so many stinging ants, but they always kept their spirits high and ii made my field trips unforgettable. Also thanks to the people that hosted me and to the institutions that made my trips successful. Finally I would like to thank my friends and Thor for keeping me sane. Chapter-specific acknowledgements Chapter 2 - We thank all those who helped us obtain plants: A. Ayodele (University of Ibadan, Nigeria); J. Burke and J. Reveal (Cornell University); the Chase Lab (Royal Botanical Gardens Kew); the Donoghue Lab (Yale University); E. Kempton (Rancho Santa Ana); F. Khasanov and H. Shomurodov (Uzbek Academy of Plant Sciences); T. Pennington (Royal Botanical Gardens Edinburgh); J. Solomon (Missouri Botanical Gardens); and A. Lamb-Frye, M. Silman, and the Wake Forest Herbarium (WFU). Financial support was provided by the Wake Forest University Graduate School of Arts and Sciences and by theWake Forest University Department of Biology. Chapter 3 - The authors acknowledge the people who helped them acquire plant material: S. Avendaño (Instituto de Ecología-Jalapa), G. Carnevali (Centro de Investigación Científica de Yucatán), G. Castillo (Instituto de Ecología-Jalapa), R. Duno (Centro de Investigación Científica de Yucatán), E. Kempton (Rancho Santa Ana Botanical Garden), S. Leython (Jardín Botánico de Caracas), J. Pale (Instituto de Ecología-Jalapa), J. Reveal (Cornell University), and S. Romero (Jardín Botánico de Caracas). They also thank staff of the herbaria A, BH, NY, GH, and US for access to material. They are grateful to Jim Reveal and one anonymous reviewer for useful improvements to the manuscript. This work was supported by funding from Wake Forest iii University (A.S.), Cornell University Department of Plant Biology, American Society of Plant Taxonomist, and Botanical Society of America (J.B.). Chapter 4 - This study was funded by Wake Forest University Graduate School of Arts and Sciences, Wake Forest University Department of Biology, the American Society of Plant Taxonomists (ASPT), and the Lewis and Clark Exploration Fund. We thank all the people who helped us obtain plant material: J. Burke (Cornell University), the Chase Lab (Royal Botanical Gardens Kew), T. Pennington (Royal Botanical Gardens Edinburgh), J. Solomon (Missouri Botanical Gardens), the New York Botanical Garden, Santiago Madriñán (Universidad de los Andes, Colombia), and A. Lamb-Frye and M. Silman (Wake Forest University).We also thank Idea Wild for field equipment. Special thanks to James L. Reveal for his comments and help with the manuscript. Chapter 5 - This study was funded by Wake Forest University Graduate School of Arts and Sciences, Wake Forest University Department of Biology, the Richter Award, the American Society of Plant Taxonomists (ASPT), and the Lewis and Clark Exploration Fund. We thank all the people who helped us obtain plant material: J. Burke (Cornell University), the Chase Lab (Royal Botanical Gardens Kew), T. Pennington (Royal Botanical Gardens Edinburgh), J. Solomon (Missouri Botanical Gardens), the New York Botanical Garden, Santiago Madriñán (Universidad de los Andes, Colombia), and A. Lamb-Frye and M. Silman (Wake Forest University); the people who helped with field work: F. Michelangeli, E. Bellota, E. Cosio, F. Ramírez, M. Gaviria, C. Pedraza, P. Pulgarín, D. Daly, H. Medeiros, F. Obermuller, E. Oliveira, M. Silveira, R. Aguilar, B. iv Hammel, and N. Zamora, and to the institutions that helped us obtain collecting permits: INRENA (Peru), InBio (Costa Rica) and CNPq (Brazil). We would also like to thank P. Ward for the identification of Pseudomyrmex and collection information, and F. Azorsa and W. McKay for identification of other ant genera associated to Triplaris. F. Estela contributed to the GIS mapping and Idea Wild with field equipment. v TABLE OF CONTENTS ACKNOWLEDGEMENTS …………………………………………………………….. ii LIST OF APPENDICES ………………………………………………………………... ix LIST OF TABLES ……………………………………………………………………… xi LIST OF FIGURES …………………………………………………………………… xiii ABSTRACT ……………………………………………………………………………... 1 CHAPTER I An Introduction To Polygonaceae And Triplaris ……………..………………… 4 CHAPTER II A Large Scale Phylogeny Of Polygonaceae And Its Implications For Taxonomic Circumscriptions. Published in International Journal of Plant Sciences (2009) ………………….. 13 ABSTRACT ……………………………………………………………. 14 INTRODUCTION ……………………………………………………... 15 MATERIALS AND METHODS ………………………………………. 19 RESULTS ……………………………………………………………… 23 DISCUSSION ………………………………………………………….. 29 LITERATURE CITED ………………………………………………… 36 CHAPTER III Placing The Woody Tropical Genera Of Polygonaceae: A Hypothesis Of Character Evolution And Phylogeny. Published in American Journal of Botany (2010) ……………………………… 56 ABSTRACT …………………………………………….……………… 57 vi INTRODUCTION ……………………………………………………... 58 MATERIALS AND METHODS ………………………………………. 63 RESULTS ……………………………………………………………… 71 DISCUSSION ………………………………………………………….. 75 LITERATURE CITED ………………………………………………… 86 CHAPTER IV Phylogenetic Relationships Of Triplaris And Ruprechtia: Re-delimitation Of The Recognized Genera And Two New Genera For Tribe Triplarideae (Polygonaceae) Accepted for publication in Systematic Botany ….…………………………… 110 ABSTRACT ………………………………………………………….. 111 INTRODUCTION ……………………………………………………. 112 MATERIALS AND METHODS ……………………………………... 116 RESULTS …………………………………………………………….. 120 DISCUSSION ………………………………………………………… 124 LITERATURE CITED ……………………………………………….. 133 CHAPTER V Mutualism Between Triplaris And Pseudomyrmex: Phylogeny And Geographical Distribution ………………...…………………………………………………. 148 ABSTRACT ………………………………………………………….. 149 INTRODUCTION ……………………………………………………. 150 MATERIALS AND METHODS ……………………………………... 154 RESULTS …………………………………………………………….. 160 DISCUSSION ………………………………………………………… 167 vii LITERATURE CITED ……………………………………………..… 179 CHAPTER VI Conclusions And Future Directions ……...…………………………………… 208 CURRICULUM VITAE ……………………………………………………………… 217 viii LIST OF APPENDICES Appendix 2.1 Voucher information for DNA extractions used in this study. Sequences obtained from Genbank are given with their respective site specific numbers. New sequences generated for this study provide the following information: Taxon, collector(s), collection number (in italics), and Genbank accession numbers. Gene abbreviations are as follows: R = rbcL, M = matK, N = ndhF and I = ITS. NA= not used in this study. Appendix 3.1 Voucher information for DNA extractions used in this study. Sequences obtained from GenBank are given with their respective site specific numbers. New sequences generated for this study provide the following information: taxon, collector(s), collection number (in italics), and GenBank accession numbers. Gene abbreviations are as follows: R = rbcL , M = matK , N = ndhF, P = psbM-ycf6 , A = psaI-accD and I = ITS. NA= not used in this study. Appendix 4.1 Voucher information for DNA extractions used in this study. Sequences obtained from Genbank are given with their respective site specific numbers. The following information is provided for new sequences generated by this study: Taxon, collector(s), collection number (in italics), and Genbank accession numbers. Gene abbreviations are as follows: M = matK, N = ndhF, D= ndhC-trnV, K= rps16-trnK, I = ITS, L = lfy2i. NA= not used in this study. Appendix 5.1 Voucher information for plant DNA extractions used in this study. Sequences obtained from Genbank are given with their respective site specific numbers. New sequences generated