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Quotes "The small part of ignorance that we arrange and classify we give the name of knowledge."-A. Bierce "I love fools' experiments. I am always making them."- C. Darwin "Spend it foolishly."- J. S. Karafit "When the center is in the middle, you have a cross section." - G. W. Rothwell "It's not "breaking up", it's a fossil, it's just sitting there."- R. A. Stockey "It takes a long time to pick up after a glacier."- L. E. Thompson "No way of thinking or doing, however ancient, can be trusted without proof." - H. D. Thoreau There should be no monotony In studying your botany; It helps to train And spur the brain— Unless you haven't gotany... - B. Braley Uber alien Gipfeln IstRuh, In alien Wipfeln Spiirest du Kaum einen Hauch; Die Vogelein schweigen im Walde, Warte nur, balde Ruhest du auch. - Goethe University of Alberta Permineralized Late Cretaceous plants from the Eden Main fossil localities, British Columbia, Canada by Steven James Karafit (fy\ A thesis submitted to the Faculty of Graduate Studies and Research in partial fulfillment of the requirements for the degree of Master of Science m Systematics and Evolution Department of Biological Sciences Edmonton, Alberta Fall 2008 Library and Bibliotheque et 1*1 Archives Canada Archives Canada Published Heritage Direction du Branch Patrimoine de I'edition 395 Wellington Street 395, rue Wellington Ottawa ON K1A0N4 Ottawa ON K1A0N4 Canada Canada Your file Votre reference ISBN: 978-0-494-47277-4 Our file Notre reference ISBN: 978-0-494-47277-4 NOTICE: AVIS: The author has granted a non­ L'auteur a accorde une licence non exclusive exclusive license allowing Library permettant a la Bibliotheque et Archives and Archives Canada to reproduce, Canada de reproduire, publier, archiver, publish, archive, preserve, conserve, sauvegarder, conserver, transmettre au public communicate to the public by par telecommunication ou par Plntemet, prefer, telecommunication or on the Internet, distribuer et vendre des theses partout dans loan, distribute and sell theses le monde, a des fins commerciales ou autres, worldwide, for commercial or non­ sur support microforme, papier, electronique commercial purposes, in microform, et/ou autres formats. paper, electronic and/or any other formats. The author retains copyright L'auteur conserve la propriete du droit d'auteur ownership and moral rights in et des droits moraux qui protege cette these. this thesis. Neither the thesis Ni la these ni des extraits substantiels de nor substantial extracts from it celle-ci ne doivent etre imprimes ou autrement may be printed or otherwise reproduits sans son autorisation. reproduced without the author's permission. In compliance with the Canadian Conformement a la loi canadienne Privacy Act some supporting sur la protection de la vie privee, forms may have been removed quelques formulaires secondaires from this thesis. ont ete enleves de cette these. While these forms may be included Bien que ces formulaires in the document page count, aient inclus dans la pagination, their removal does not represent il n'y aura aucun contenu manquant. any loss of content from the thesis. Canada Dedication To my family, friends, and colleges Abstract Permineralized fossil remains of ?dinoflagellates, mosses, ferns, gymnosperms and angiosperms have been found in calcium carbonate concretions from the Eden Main fossil localities, British Columbia, Canada. Fossils come from the Upper Cretaceous Comox Formation of the Nanaimo Group and have been dated as Coniacian in age. A survey of the fossil floral assemblage has revealed morphotaxa of eleven seedless vascular plants from stipes and rhizomes. A new species of Paralygodium, a schizaeaceous fern, is described from fertile pinnules. Seven gymnosperm organs are known including a cycadophyte stem, cupressaceous leaves, twigs and cones, and pinaceous leaves and a pollen cone. Twenty-five angiosperm morphotaxa are also currently recognized based on flowers, inflorescences, fruits, and wood. The presence of several taxa with extant relatives that are currently found in the tropics, and abundant fusain indicates that western North America was warm during the Coniacian, probably experienced wet and dry seasons. Acknowledgements I wish to thank my supervisor Dr. Ruth Stockey (University of Alberta) for her guidance and hospitality over the last few years, none of this would have been possible without her assistance. I also wish to thank my committee, Dr. Randolph Currah and Dr. Jocelyn Hall and Alexander Wolfe (University of Alberta) for their advice and assistance as I've run into questions during my research. I'd also like to thank Dr. Gar Rothwell (Ohio Universaity) for sparking my interest in the field of paleobotany with his inspiring teaching, and for allowing me to work in his lab as an undergraduate student. To my lab mates past and present, Dr. Genaro Rodrigo Hernandez Castillo, Dr. Stefan Little, Dr. Selena Smith, Dr. Randal Mindell, AJ Harris, Jen Gemmel, Ashley Klymiuk, Kirstin Brink and Brian Rankin. I would also like to acknowledge Dr. Heather Sanders, Marie Davey, Marcie Plishka, and Dr. Thomas Simonsen, for their scholarly conversations and friendship. Also the following people deserve acknowledgement for their help over the last few years, Dr. Dirk Meckert and Rick Ross (Courtenay, BC) for help in the field, Dr. Peter Mustard (Simon Fraser University) for discussion on the geology of Eden Main as well as allowing me to modify figures for this thesis; Dr. Jim Haggart (Geological Survey of Canada) for information on the geology of Eden Main, and George Braybrook (University of Alberta) for his superb SEM assistance. I would also like to thank the following people for helpful discussion, Dr. A.R. Sweet (Geological Survey of Canada) Dr. Harufumi Nishida (Chuo University), Dr. Estella Leopold (University of Washington), Dr. Vera Pspelova (University of Victoria), Dr. Eva Koppelhus (University of Alberta), Dr. Catherine LaFarge (University of Alberta), Nathan Jud (Ohio University), Dr. Steven Manchester (University of Florida), Dr. Elsa Marie Friis (Swedish Museum of Natural History), Dr. Jim Doyle (University of California, Davis), Dr. John T. Mickel (New York Botanical Garden), Dr. Thomas N. Taylor (University of Kansas), Dr. Peter Crane (University of Chicago), Dr. Bruce Tiffhey (University of California, Santa Barbara) and Dr. Alfred Traverse (Pennsylvania State University). Last but not least, I would like to thank my family. My mom, Brenda and dad, Jim Karafit for their endless support and to my brother Joe. Thanks is also due to Uncle John for sparking my interest in fossils by sharing his knowledge about rocks and fossils with me as a child. To Emilie Karafit and Margie Wenning, my grandmothers, for their - support and worry about my well being. This thesis was supported in part by NSERC grant A-6908 to RAS. Table of Contents Chapter 1: Introduction 1 Significance , 2 Materials and Methods 3 The Present Work 5 Chapter 2: Eden Main Flora: Seedless taxa 13 Introduction 13 Dinoflagellates 13 Moss 14 Ferns 16 Conclusions 35 Chapter 3: Eden Main Flora: Paralygodium meckertii 37 Introduction 37 Specific diagnosis ..37 Description 39 Discussion 40 Chapter 4: Eden Main Flora: Seed plants 47 Gymnosperms 47 Angiosperms 62 Conclusions 89 Chapter 5: Conclusions 91 Seedless taxa 92 Gymnosperms 93 Angiosperms 94 Climate, Fire, and Paleolatitude 95 Concluding Remarks 97 Literature Cited 169 List of Tables Chapter 1 Table 1. Eden Main Localities Chapter 2 Table 2. Comparison of Solenostelopteris and Loxomopteris species. Chapter 3 Table 3. Comparison of morphology, spores and age of fertile pinnules of Paralygodium spp. and Anemia poolensis. Chapter 4 Table 4. Comparison of anatomically preserved Cretaceous Pinaceae. List of Text Figures Figure 1. Map of Vancouver Island showing Eden Main localities. 8 Figure 2. Geological map of Eden Main localities. 10 Figure 3. (a) Generalized lithostratigraphy of Nanaimo Group. (b) Generalized stratigraphy of members of Comox Formation. 12 List of Plates Plate 1. Dinoflagellates and bryophytes. 100 Plate 2. Gleicheniaceous rhizome, and frond segments. 102 Plate 3. Branching gleicheniaceous frond. 104 Plate 4. Schizaeaceous and cyatheaceous fronds. 106 Plate 5. Transverse sections of various Eden Main fern rachides. 108 Plate 6. Exarch solenostelic fern rhizome. 110 Plate 7. Transverse sections of amphiphloic, solenostelic fern rhizome. 112 Plate 8. Paralygodium meckertii sp. nov., general morphology. 114 Plate 9. Paralygodium meckertii sp. nov., general morphology and histology. 116 Plate 10. Paralygodium meckertii sp. nov., spores. 118 Plate 11. Paralygodium meckertii sp nov., three dimensional reconstruction. 120 Plate 12. Cycadeoid with resin canals in the cortex and a woody eustele 122 Plate 13. Pinus leaves in transverse section. 124 Plate 14. Pinaceous pollen cone and pollen. 126 Plate 15. SEM images of bisaccate pollen grains from pinaceous cone. 128 Plate 16. Organs of probable taxodiaceous Cupressaceae. 130 Plate 17. Conifer seed, cf. Araucaria. 132 Plate 18. Lauraceous flower and inflorescences. 134 Plate 19. Lauraceous fruit. 136 Plate 20. Transverse sections of angiosperm wood cf. Paraphyllanthoxylon. 138 Plate 21. Radial longitudinal section of angiosperm wood cf. Paraphyllanthoxylon. 140 Plate 22. Tangential longitudinal section of angiosperm wood cf. Paraphyllanthoxylon. 142 Plate 23. Inflorescences and fruits. 144 Plate 24. SEM of platanaceous pollen. 146 Plate 25. Nyssaceae fruits in transverse section. 148 Plate 26. Longitudinal
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