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The Phylogeny, Morphological Evolution and Biogeography Of THE PHYLOGENY, MORPHOLOGICAL EVOLUTION AND BIOGEOGRAPHY OF THE GAULTHERIEAE (ERICACEAE) By CATHERINE MARY BUSH A Dissertation Submitted to the Graduate Faculty of WAKE FOREST UNIVERSITY GRADUATE SCHOOL OF ARTS AND SCIENCES In Partial Fulfillment of the Requirements for the Degree of DOCTOR OF PHILOSOPHY in the Department of Biology May 2010 Winston-Salem, NC Approved by: Kathleen A. Kron, Ph.D., Advisor ____________________________________ Examining Committee: Zack Murrell, Ph.D., Chair ____________________________________ William K. Smith, Ph.D. ____________________________________ Peter D. Weigl, Ph.D. ____________________________________ Clifford Zeyl, Ph. D. ____________________________________ ACKNOWLEDGEMENTS This dissertation would absolutely not have been possible without the support, mentoring and patience of many people. I leaned on my family and fiancé to get through this process, at times wonderful and at times overwhelming. I had their support as I spent months doing field work in six countries. The distance was never easy for either side, but they knew my work made me happy and they were behind me even when it was difficult. I would like to thank my advisor, Kathleen A. Kron, who allowed me to become an independent scientist. Kathy’s door was always open when I needed a push in the right direction and I have learned a tremendous amount from being in her lab. Thanks also to my advisory committee, Dr. Zack Murrell, Dr. William Smith, Dr. Peter D. Weigl and Dr. Clifford Zeyl for their support and guidance. I would like to thank Dr. Gerald L. Smith, my undergraduate advisor who led me with his enthusiasm and kindness on the road to botanical exploration and research. He is a colleague but also a dear friend and I treasure the times we have spent together in the lab or in the field, trying to understand the ever-delightful and elusive spider lilies. I also want to thank the entire biology department faculty at High Point University. My alma mater professors have kept in touch as I progressed in this degree and they are like an extended family to me. I absolutely could not have made it through this degree without the empathy and shared laughter of my labmates past and present – Ann Powell, Emily Gillespie, Adriana Sanchez, Tanja Schuster and Sabrina Setaro. I want to thank Dr. Peter W. Fritsch for taking a chance on me and inviting me on my first major field expedition to China. I am very grateful for the collaboration and ii friendship that grew from that first trip. Peter has been constant source of encouragement and guidance throughout this journey. One of the unexpected gifts of this degree was Lu Lu, my Chinese collaborator and dear friend. She has taught me so much, about both science and life. Finally, I want to thank the generous and kind people who hosted me around the world: Darren Crayn, João Luiz M. Aranha Filho, Angela B. Martins, Ted Oliver, Karen Rea, Rimi Repin, Gisela Sancho, Ross Turner and Steve Wagstaff. Your hospitality will not be forgotten. Chapter-specific acknowledgements Chapter 2 - We thank the anonymous reviewers for their helpful suggestions. We would also like to acknowledge the financial support from Wake Forest University and the National Science Foundation (DEB-0717724, DEB-0717711). Chapter 3 - We thank the anonymous reviewers for their helpful suggestions. We would also like to acknowledge the financial support from Wake Forest University and the National Science Foundation (DEB-0717724; DEB-0717711). Chapter 4 - We acknowledge the financial support from Wake Forest University and the National Science Foundation (DEB-0717724; DEB-0717711). We thank the curators of the Allan Herbarium, the National Herbarium of New South Wales and the Tasmanian Herbarium for access to the databases and collections as well as the New Zealand Department of Conservation and Biodiversity Conservation Branch (Department of iii Primary Industries, Water and Environment; Tasmania) for their helpful advice and for administering our collection permits. We also thank Darren Crayn for help in the field, and two anonymous reviewers for their helpful manuscript suggestions. Chapter 5 - We would like to thank the curators of the following herbaria for loaning us the material needed for this morphological revision and/or for their expert advice regarding type specimens: AK, B, BM, CHR, E, GOET, HO, K, L, LINN, LIV, MELU, MO, MPN, MW, NSW, NZFRI, P, PH, UNSW, UPS, W, WAIK and WELT. We thank Dr. Gerald L. Smith for his knowledgeable critiques of this manuscript. We acknowledge the financial support from Wake Forest University and the National Science Foundation (DEB-0717724; DEB-0717711). iv TABLE OF CONTENTS ACKNOWLEDGEMENTS………………………………………………………….........ii LIST OF TABLES…………………………………………………………………...…viii LIST OF FIGURES……………………………………………………………………....ix LIST OF APPENDICES……………………………………………………………......xiii ABSTRACT……………………………………………………………………………….1 CHAPTER I An Introduction to the Gaultherieae………………………………………………4 CHAPTER II Phylogeny of Gaultherieae (Ericaceae: Vaccinioideae) based on DNA sequence data from matK, ndhF and nrITS. Published in International Journal of Plant Sciences (2009)……………………19 ABSTRACT……………………………………………………………...20 INTRODUCTION……………………………………………………….21 MATERIALS AND METHODS………………………………………...26 RESULTS………………………………………………………………..31 DISCUSSION……………………………………………………………36 LITERATURE CITED…………………………………………………..41 CHAPTER III The Phylogeny of Leucothoë s.l. (Ericaceae: Vaccinioideae) based on morphological and molecular (ndhF, matK, and nrITS) data. Accepted for publication in Systematic Botany (2010)…………………………..54 ABSTRACT……………………………………………………………...55 v INTRODUCTION………………………………………………………56 MATERIALS AND METHODS………………………………………..60 RESULTS………………………………………………………………..63 DISCUSSION……………………………………………………………66 LITERATURE CITED…………………………………………………..70 CHAPTER IV The phylogeny, biogeography and morphological evolution of Gaultheria (Ericaceae) from Australia and New Zealand. Published in Australian Systematic Botany (2009)………………………………79 ABSTRACT……………………………………………………………...80 INTRODUCTION……………………………………………………….81 MATERIALS AND METHODS………………………………………...85 RESULTS………………………………………………………………..92 DISCUSSION……………………………………………………………96 LITERATURE CITED…………………………………………………102 CHAPTER V The Flowering Plant Genus Gaultheria (Ericaceae) in Australia and New Zealand………………………………………………………………………….124 ABSTRACT…………………………………………………………….125 INTRODUCTION……………………………………………………...126 MATERIALS AND METHODS……………………………………….130 RESULTS ……………………………………………………………...132 DISCUSSION…………………………………………………………..134 vi TAXONOMIC TREATMENT………………………………………....139 LITERATURE CITED…………………………………………………165 CHAPTER VI The phylogeny and morphology of the Brazilian Gaultheria (Ericaceae) species. Submitted for publication in Systematic Botany………………………………..192 ABSTRACT…………………………………………………………….193 INTRODUCTION……………………………………………………...194 MATERIALS AND METHODS……………………………………….199 RESULTS………………………………………………………………202 DISCUSSION…………………………………………………………..204 LITERATURE CITED…………………………………………………207 CHAPTER VII Conclusions……………………………………………………………………..215 CURRICULUM VITAE……………..…………………………………………………223 vii LIST OF TABLES Table 4.1. Taxa sampled in this study of the Gaultheria from Australia and New Zealand………………………………………………………………………….107 Table 4.2. Statistics for individual and combined data maximum parsimony analyses in the study of the Gaultheria from Australia and New Zealand……………….…110 Table 5.1 Total number of herbarium sheets examined per species…………………...166 Table 5.2 Characters measured/observed for morphometric study……………………168 Table 6.1 Taxa sampled in this study of the Gaultheria from Brazil………………….209 Table 6.2 Statistics for individual and combined data maximum parsimony analyses in the study of the Gaultheria from Brazil………………………………………...211 viii LIST OF FIGURES Figure 2.1. Strict consensus of the six most parsimonious trees (L = 1,021, CI = 0.58, RI = 0.79) from the total combined data parsimony analysis (matK, ndhF, and nrITS; Gaultherieae phylogeny)…………………………………………………44 Figure 2.2. Single consensus tree from a Bayesian analysis of the total combined data set (matK, ndhF, and nrITS; Gaultherieae phylogeny)……………………………...46 Figure 2.3. Calyx morphology traced onto the single consensus tree from a Bayesian analysis of the total combined data set (matK, ndhF, and nrITS; Gaultherieae phylogeny)……………………………………………………………………….48 Figure 2.4. Fruit morphology traced onto the single consensus tree from a Bayesian analysis of the total combined data set (matK, ndhF, and nrITS; Gaultherieae phylogeny)……………………………………………………………………….50 Figure 2.5. Inflorescence type traced onto the single consensus tree from a Bayesian analysis of the total combined data set (matK, ndhF, and nrITS; Gaultherieae phylogeny)……………………………………………………………………….52 Figure 3.1. Strict consensus of the two most parsimonious trees (L = 155, CI = 0.56, RI = 0.53) based on morphological/non-molecular data (Leucothoë phylogeny)………………………………………………………………….........75 Figure 3.2. The one most parsimonious tree (L = 689, CI = 0.62, RI = 0.67) from the total combined data parsimony analysis (matK, ndhF, nrITS, morphology; Leucothoë phylogeny)……………………………………………………………77 Figure 4.1. Strict consensus of the five most parsimonious trees (length = 733, CI = 0.54, RI = 0.59) from a parsimony analysis of ix the total combined data (matK, ndhF, nrITS, lfy, waxy; Gaultheria from Australia and New Zealand)………………………………………………………………112 Figure 4.2. The single tree from a maximum likelihood analysis of the total combined dataset (matK, ndhF, nrITS, lfy, waxy; Gaultheria
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