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University of Florida Thesis Or Dissertation Formatting SIDEROXYLON SECTION FRIGORICOLA (SAPOTACEAE): A CLADE ENDEMIC TO TEMPERATE NORTH AMERICA By PAUL THOMAS COROGIN 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 2015 © 2015 Paul Thomas Corogin To Ananda Devi, my wife and soulmate, whose presence beside me gives me strength ACKNOWLEDGMENTS I thank the following herbaria for loans of specimens: A, AUA, FLAS, FNPS, FSU, FTG, GA, GH, LSU, MISS, OKL, TENN, UNA, US, USCH, and USF. I thank the following people without whose help this study could not have succeeded: Kent Perkins of the University of Florida Herbarium (FLAS) for his generous assistance with processing specimen loans; Brett Jestrow of Fairchild Tropical Garden (FTG) and Jennifer Stafford of the National Park Service herbarium (FNPS) for facilitating my examination of south Florida herbarium specimens; Jimi Sadle, Mark Whitten, Bob Simons, Wendy Poag, Lisa Kruse, and Dee Mincey, for assisting with field work; Karen Kelley, electron microscopy manager, and Kim Backer-Kelley, lab technician, of the ICBR Electron Microscopy Core lab at the University of Florida, for their kindness and expertise, and Kurt Neubig for his guidance and insights with the molecular work. And I thank my supervisory committee: Drs. Norris Williams, Pamela Soltis, Nigel Smith, and my advisor, Dr. Walter S. Judd, who gives so generously of his time to inspire in his students something of his passion for scholarship and botanical field expertise. I have indeed studied and learned from one of the best there is. 4 TABLE OF CONTENTS page ACKNOWLEDGMENTS................................................................................................. 4 LIST OF TABLES........................................................................................................... 7 LIST OF FIGURES ........................................................................................................ 8 ABSTRACT.................................................................................................................. 11 CHAPTER 1 INTRODUCTION ................................................................................................... 13 The Family Sapotaceae ......................................................................................... 13 The Genus Sideroxylon ......................................................................................... 16 Biogeography of Sapotaceae in Southeastern North America ............................... 18 The Aims of This Study.......................................................................................... 22 2 GENERIC AND SECTIONAL DELIMITATIONS WITHIN SIDEROXYLON ............ 24 Relationships of North American and Tropical Species of Sideroxylon Based On ITS Sequences ................................................................................................... 26 Materials and Methods .......................................................................................... 26 Results and discussion .......................................................................................... 29 3 PHYLOGENETIC RELATIONSHIPS WITHIN SIDEROXYLON SECTION FRIGORICOLA BASED UPON MORPHOLOGY AND cpDNA SEQUENCES ....... 35 Species and Subspecies Concepts ....................................................................... 36 Materials and Methods .......................................................................................... 37 General Morphology ........................................................................................ 37 Morphological Cladistic Analysis ..................................................................... 38 Chloroplast DNA Analysis ............................................................................... 39 Results and Discussion ......................................................................................... 41 Morphological Cladistic Analysis ..................................................................... 41 Chloroplast DNA Analysis ............................................................................... 44 Combined ITS + cpDNA Datasets ................................................................... 45 Comparison of Phylogenies ............................................................................ 46 Assessment of Phylogenetic Species .............................................................. 48 4 ECOLOGY AND BIOGEOGRAPHY ...................................................................... 62 Historical Biogeography of the Genus Sideroxylon ................................................ 62 The North American Coastal Plain: A Hotspot of Endemic Biodiversity.................. 63 Sideroxylon section Frigoricola: A Group of NACP Endemics ................................ 65 5 Habitats and Distribution ................................................................................. 66 Eco-Geographical Differentiation of the Species ............................................. 76 5 TAXONOMY .......................................................................................................... 79 Key to Species of Sideroxylon sect. Frigoricola ..................................................... 89 Key to the Subspecies of Sideroxylon lanuginosum............................................. 143 Key to the Subspecies of Sideroxylon reclinatum ................................................ 179 6 CONCLUSION .................................................................................................... 244 LIST OF REFERENCES ............................................................................................ 247 BIOGRAPHICAL SKETCH ......................................................................................... 264 6 LIST OF TABLES Table page 2-1 List of taxa, vouchers, and GenBank accession numbers.................................. 31 3-1 Characters used in the morphology-based phylogenetic analysis...................... 50 3-2 Character values used in cladistic analysis ....................................................... 52 3-3 Character totals for DNA regions and combined datasets ................................. 53 3-4 Putative autapomorphies for species of Sideroxylon section Frigoricola. ........... 54 7 LIST OF FIGURES Figure page 2-1 Strict consensus of 12 most parsimonious trees from analysis of ITS sequences ......................................................................................................... 32 2-2 Representative cladogram: one of 12 most parsimonious trees generated in analysis of ITS sequences. ................................................................................ 33 2-3 Best tree generated by maximum likelihood analysis of ITS sequences ............ 34 3-1 Single most parsimonious tree from morphological analysis .............................. 55 3-2 Strict consensus of 20 most parsimonious trees from MP analysis of sequences from five cpDNA regions ................................................................. 56 3-3 Representative cladogram: one of 20 most parsimonious trees generated in MP analysis of sequences from five cpDNA regions.......................................... 57 3-4 Best tree generated by maximum likelihood analysis of sequences from five cpDNA regions .................................................................................................. 58 3-5 Strict consensus of five most parsimonious trees from MP analysis of combined dataset of ITS plus five cpDNA regions ............................................. 59 3-6 Representative cladogram: one of five most parsimonious trees generated in MP analysis of combined dataset of ITS plus five cpDNA regions ..................... 60 3-7 Best tree generated by maximum likelihood analysis of combined dataset of ITS plus five cpDNA regions .............................................................................. 61 4-1 Geographical ranges of the species of Sideroxylon section Frigoricola ............. 78 5-1 Distribution of Sideroxylon sect. Frigoricola. .................................................... 214 5-2 SEM images of abaxial leaf surface of four individuals of Sideroxylon lycioides .......................................................................................................... 215 5-3 Sideroxylon lycioides ....................................................................................... 216 5-4 Distribution of Sideroxylon lycioides ................................................................ 217 5-5 SEM images of abaxial leaf surface of two individuals of Sideroxylon thornei . 218 5-6 Sideroxylon thornei.......................................................................................... 219 5-7 Distribution of Sideroxylon thornei ................................................................... 220 8 5-8 SEM images of abaxial leaf surface of Sideroxylon tenax ............................... 221 5-9 Sideroxylon tenax ............................................................................................ 222 5-10 Sideroxylon tenax, morphological examples typical of species of Sideroxylon sect. Frigoricola ............................................................................................... 223 5-11 Sideroxylon tenax, flower, typical of species of Sideroxylon sect. Frigoricola .. 224 5-12 Distribution of Sideroxylon tenax ....................................................................
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