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1 History of Vitaceae Inferred from Morphology-Based

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1 History of Vitaceae Inferred from Morphology-Based HISTORY OF VITACEAE INFERRED FROM MORPHOLOGY-BASED PHYLOGENY AND THE FOSSIL RECORD OF SEEDS By IJU CHEN 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 Iju Chen 2 To my parents and my sisters, 2-, 3-, 4-ju 3 ACKNOWLEDGMENTS I thank Dr. Steven Manchester for providing the important fossil information, sharing the beautiful images of the fossils, and reviewing the dissertation. I thank Dr. Walter Judd for providing valuable discussion. I thank Dr. Hongshan Wang, Dr. Dario de Franceschi, Dr. Mary Dettmann, and Dr. Peta Hayes for access to the paleobotanical specimens in museum collections, Dr. Kent Perkins for arranging the herbarium loans, Dr. Suhua Shi for arranging the field trip in China, and Dr. Betsy R. Jackes for lending extant Australian vitaceous seeds and arranging the field trip in Australia. This research is partially supported by National Science Foundation Doctoral Dissertation Improvement Grants award number 0608342. 4 TABLE OF CONTENTS page ACKNOWLEDGMENTS ...............................................................................................................4 LIST OF TABLES...........................................................................................................................9 LIST OF FIGURES .......................................................................................................................11 ABSTRACT...................................................................................................................................14 CHAPTER 1 SEED MORPHOLOGY OF VITACEAE..............................................................................16 Introduction.............................................................................................................................16 Materials and Methods ...........................................................................................................17 Results.....................................................................................................................................18 Leea, Cissus, and Cyphostemma .....................................................................................20 Tetrastigma and Rhoicissus.............................................................................................22 Acareosperma and Cayratia............................................................................................24 Ampelocissus, Nothocissus, and Pterisanthes .................................................................26 Vitis, Ampelopsis, Clematicissus, Parthenocissus, and Yua............................................26 Discussion...............................................................................................................................28 2 MORPHOLOGY-BASED PHYLOGENY OF VITACEAE: COMPARING TWO DIFFERENT TREATMENTS FOR CODING CONTINUOUS CHARACTERS................75 Introduction.............................................................................................................................75 Materials and Methods ...........................................................................................................80 Taxon Sampling...............................................................................................................80 Terminology of Morphological Characters.....................................................................80 Character Measurement...................................................................................................82 Character Coding.............................................................................................................82 Phylogenetic Analyses.....................................................................................................84 Results.....................................................................................................................................85 Discussion...............................................................................................................................91 The Influences of Coding Methods .................................................................................91 Relationships Within the Family, Comparisons with the Molecular Data......................92 Morphology of Vitaceae and Character Evolution..........................................................99 Growth habit.............................................................................................................99 Phyllotaxy...............................................................................................................101 Tendrils...................................................................................................................103 Stipules...................................................................................................................105 Leaves.....................................................................................................................106 Hairs .......................................................................................................................111 Sexuality.................................................................................................................112 5 Inflorescence-branch architecture ..........................................................................113 Inflorescences architecture.....................................................................................118 Floral morphology..................................................................................................121 Pollen morphology .................................................................................................125 Fruits.......................................................................................................................125 Seeds.......................................................................................................................126 Concluding Remarks ............................................................................................................127 3 THE BIOGEOGRAPHICAL HISTORY OF VITACEAE INFERRED FROM FOSSIL SEEDS ..................................................................................................................................159 Introduction...........................................................................................................................159 Materials and Methods .........................................................................................................162 Results and Discussion .........................................................................................................165 Classification of Fossil Vitaceaous Seeds .....................................................................165 1) st-Ampelocissus-wide infolds.............................................................................165 2) st-Ampelocissus-rugose......................................................................................165 3) st-Ampelopsis-smooth........................................................................................166 4) st-Ampelopsis-rugose .........................................................................................167 5) st-Ampelopsis-xs ................................................................................................167 6) st-Vitis ................................................................................................................168 7) st-Vitis-Ampelopsis ............................................................................................168 8) st-Vitis rotundifolia ............................................................................................169 9) st-Parthenocissus ...............................................................................................169 10) st-Parthenocissus clarnensis............................................................................170 11) st-Cayratia .......................................................................................................171 12) st-Tetrastigma ..................................................................................................171 13) st-perichalaza ...................................................................................................171 14) uncertain specimens with affinity to Vitaceae .................................................172 Summary of seed type classification......................................................................173 Geographic Distribution of Fossil and Extant Vitaceous Seeds....................................175 Europe ....................................................................................................................176 Siberia.....................................................................................................................177 Asia.........................................................................................................................177 North America........................................................................................................178 Central and South America ....................................................................................180 Africa......................................................................................................................180 Australia
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