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Zingiberalean Fossils from the Late Paleocene of North Dakota, USA and Their Significance to the Origin and Diversification of Zingiberales

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Zingiberalean Fossils from the Late Paleocene of North Dakota, USA and Their Significance to the Origin and Diversification of Zingiberales Zingiberalean fossils from the Late Paleocene of North Dakota, USA and their significance to the origin and diversification of Zingiberales by John C. Benedict A Dissertation Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy Approved April 2012 by the Graduate Supervisory Committee: Kathleen B. Pigg, Chair Martin F. Wojciechowski Melanie L. DeVore Patricia L. Fall ARIZONA STATE UNIVERSITY May 2012 ABSTRACT The Zingiberales, including the gingers (Zingiber), bananas (Musa) and ornamental flowers (Strelitzia, Canna, and Heliconia) are a diverse group of monocots that occupy the tropics and subtropics worldwide. The monophyly of the order is well supported, although relationships between families are not well resolved. A rapid divergence of the Zingiberales has been proposed to explain the poor resolution of paraphyletic families in the order, and direct fossil evidence shows members of both of these lineages of Zingiberaceae and Musaceae were present by the Late Cretaceous. Comparisons of the fossils with extant relatives and their systematic placement have been limited because variation within modern taxa is not completely known. The current study focuses on describing zingiberalean fossil material from North Dakota that includes seeds, leaves, buds, adventitious roots and rhizomes. A survey of extant zingiberalean seeds was conducted, including descriptions of those for which data were previously unknown, in order to resolve the taxonomic placement of the fossil material. Upon careful examination, anatomical characters of the seed coat in fossil and extant seeds provide the basis for a more accurate taxonomic placement of the fossils and a better understanding of character evolution within the order. i DEDICATION To Mitsuko and Shiro, Always there when I need support, or to just to help motivate me to get off my lazy butt and go for a walk. ii ACKNOWLEDGMENTS This study was completed with the gracious efforts of countless individuals that offered their thoughts, guidance, patience and most importantly their time. Without these individuals this study would never have been completed and probably never would have made it through the “good idea” stage. I would like to thank the faculty, staff ASU for their time, suggestions on coursework and offering courses that fueled my passion for plants. ASU students, graduate and undergraduate, that have been great friends, co-workers and pupils for which I have had the pleasure to teach. I would like to acknowledge my dissertation committee, Martin F. Wojciechowski, Melanie L. Devore, Patricia L. Fall for their helpful commentary on this work and well as their constant support throughout my PhD studies. I would like to thank the many people that have extended a helping hand in providing specimens for study, or useful guidance on this subject. Steven Manchester, Barbra Mohr, and Sergio Cevallos-Ferriz for providing comparative material for study. Walter Freidrich for donating much of his Zingiberales material and graciously allowing me to stay at his lovely home during my visit to Aarhus, Denmark. W. John Kress, Ida Lopez and Vinita Gowda of the Smithsonian Institution for providing access to the living and herbarium collections and providing critical comparative material for this study. I especially thank Selena Y. Smith for her help with many faucets of this project, in particular, enlightening me to the realm of synchrotron imaging to iii obtain the locked secrets inside zingiber seeds. I would like to thank my funding sources, Arizona State University Travels Grants to Disseminate this and other works throughout my PhD program and the Michael Cichan Award, the Botanical Society of America, American Society of Plant Taxonomy, as well as the Heliconia Society International Awards for botanical and horticultural research projects. Last but not least I would like to thank Dr. Kathleen B. Pigg, my chair, PhD advisor, and friend. Words cannot express how much I owe you for everything you have done to help cultivate me as a scientist. iv TABLE OF CONTENTS Page LIST OF TABLES.....................................................................................................vii LIST OF FIGURES..................................................................................................viii CHAPTER 1 INTRODUCTION.................................................................................. 1 Introduction to the Zingiberales ......................................................... 1 Relationships within the Zingiberales................................................ 9 The Fossil Record of Zingiberales ................................................... 12 Vegetative Fossil Remains of Zingiberales ..................................... 12 Reproductive Fossil Remains of Zingiberales. ................................ 18 2 MATERIALS AND METHODS ........................................................ 34 Stratigraphy and Geological Setting ................................................ 34 Fossil Localities and Preservation.................................................... 38 The Almont and Beicegel Creek Floras........................................... 39 Fossil Monocotyledonous Remains from Almont and Beicegel Creek ................................................................................................. 42 3 NOMENCLATURE OF ZINGIBERALES FRUITS AND SEEDS . 55 Historical Studies of the Seeds of Zingiberales ............................... 55 Assessing Seed and Fruit Character States in Zingiberales............. 59 4 RESULTS PART I: DESCRIPTIONS OF EXTANT SEEDS........... 67 v CHAPTER Page 5 RESULTS PART II: SYSTEMATICS AND DESCRIPTION............96 Systematics and Description of Kressia dakotensis gen. et sp. nov.96 Systematics and Description of Zingiberopsis hickeyi sp. nov. .... 100 6 DISCUSSION..................................................................................... 110 The Taxonomic Placement of Kressia dakotensis gen. et sp. nov.. ......................................................................................................... 110 Seed Anatomy Variation within Zingiberaceae............................. 116 Comparison of Kressia dakotensis with other fossil Zingiberales ......................................................................................................... 118 Justification of a New Leaf Species Zingiberopsis hickeyi sp. nov. ......................................................................................................... 123 Revisions and New Findings in Spirematospermum wetzleri....... 124 Conclusions..................................................................................... 128 REFERENCES ...................................................................................................... 141 APPENDIX A THE ALMONT AND BEICEGEL CREEK FLORAS............. 160 vi LIST OF TABLES Table Page 1. History of the taxonomic relationships of the Zingiberales................ 27 2. Reports of Cannaceae in the fossil record ........................................... 28 3. Reports of Heliconiaceae in the fossil record...................................... 29 4. Reports of Musaceae in the fossil record ............................................ 30 5. Reports of Zingiberaceae in the fossil record...................................... 31 6. Reports of fossils attributed to Zingiberales without family placement ............................................................................................................. 32 7. Previous reports of fossil Zingiberales that are no longer accepted ... 33 8. Fruit characters in Zingiberales ........................................................... 65 9. Seed characters in Zingiberales ........................................................... 66 vii LIST OF FIGURES Figure Page 1. The current distribution of the families of the order Zingiberales...... 24 2. The relationships of the families in Zingiberales based on the single most parsimonious tree obtained from 36 morphological characters and molecular data from chloroplast rbcL and atpB genes and nuclear 18S sequence data.............................................................................. 25 3. The Williston Basin of Western Interior, North America................... 50 4. The Williston Basin stratigraphic column and physiographic map.... 52 5. Photographs of the Almont and Beicegel Creek Localities................ 54 6. Seed morphology and anatomy of Ensete glucum (Roxb.) Cheesman (Musaceae)......................................................................................... 74 7. Seed morphology and anatomy of Musa sikkimensis Kurz (Musaceae) ............................................................................................................ 75 8. Seed morphology and anatomy of Heliconia bihai (L.) L. (Heliconiaceae) .................................................................................. 77 9. Seed morphology and anatomy of Strelitzia reginae Aiton (Strelitziaceae). .................................................................................. 78 10. Seed and fruit morphology and anatomy of Orchidantha vietnamica (Lowiaceae)........................................................................................ 79 11. Seed morphology and anatomy of Canna paniculata Ruiz & Pav. (Cannaceae)........................................................................................ 81 viii 12. Seed morphology and anatomy of Ctenanthe lanceolata Petersen (Marantaceae).
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