University of Alberta Paleobotanical Studies of the Appian Way Fossil Locality by Randal Alexander Mindell © A thesis submitted to the Faculty of Graduate Studies and Research in partial fulfillment of the requirements for the degree of Doctor of Philosophy in 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: 978-0-494-46384-0 Our file Notre reference ISBN: 978-0-494-46384-0 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. 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Canada ABSTRACT The Eocene Appian Way locality of Vancouver Island, British Columbia, Canada, has been surveyed for plant fossils. More than sixty taxa of mosses, ferns, conifers and flowering plants have been identified. This dissertation presents data on five new fossil taxa in three vascular plant families (Gleicheniaceae, Platanaceae and Fagaceae). The plants are anatomically preserved in calcium carbonate concretions, often at different developmental stages. This three-dimensional preservation allows for correlation of internal anatomy to external morphology. A branching fern rhizome and associated stipes are described and placed in the family Gleicheniaceae based on stelar anatomy and leaf trace divergence. The protostelic rhizome is designated as a new species of the extant genus Gleichenia and is the only Cenozoic fossil record of the family in North America. Globose, staminate inflorescences of Platanaceae (sycamores) from the locality have five-parted flowers with in situ pollen and rudimentary carpels. This is the first occurrence of rudimentary bisexuality in the fossil record of the family and the inflorescences are described as a new genus, Gynoplatananthus. Cupules and nuts of the beech family (Fagaceae) are the most common angiosperm fruiting structures in the concretions. The three species described in this thesis represent both Castaneoideae and Fagoideae, the two major subfamilies of Fagaceae. Trigonal, wingless, single-seeded, tricarpellate nuts are placed as a fossil species of the genus Fagus and represent the earliest wingless fruits of subfamily Fagoideae. Two species of spiny, cupulate, ovoid to globose fruits are described and placed in a new genus, Cascadiacarpa. They differ from all extant and fossil species of Fagaceae in being strictly bicarpellate. Their placement in subfamily Castaneoideae is supported by a phylogenetic analysis of morphological characters. The diversity and abundance of Fagaceae at the locality supports a Paleogene radiation of the family. A framework for studying fossil plants from the Appian Way locality has been established and taxa described, thus far, suggest a unique and diverse assemblage that will broaden our understanding of Eocene vegetation. Acknowledgements I would first like to acknowledge you, acknowledgement reader. You are a special breed and the world is better off for your curiosity. I would like to acknowledge all the people who serendipitously or knowingly helped in the work that comprises this dissertation. Dr. W.D Tidwell at Brigham Young University wrote The Field Guide to Common Fossil Plants of Western North America, which proved itself to be a gateway drug into the real annals of botany. John McPhee's The Annals of a Former World described time as if it was the most delicious thing ever concocted. Who wouldn't eat it up? Dr. Jim Haggart at the Geological Survey of Canada in Vancouver provided numerous insights into the geology of the Appian Way locality, as did Drs. Peter Mustard and Rolf Mathewes of Simon Fraser University. Dr. Art Sweet at the Geological Survey of Canada's Calgary office studied a plurality of sediment samples for pollen and this work has proven very useful. The paleo people of Vancouver Island are perhaps a 21st Century equivalent to a legendary Norse—make that Welsh—tribe, only armed with rock hammers in place of swords and fighting with bedrock in place of other humans. In my experience, I came to see Graham and Tina Beard of Qualicum Beach, British Columbia as the leaders of this noble enterprise. Their pure passion for fossils was matched only by their hospitality and graciousness. Were it not for the efforts of the Beards and their people, none of these fossils would ever have been known to exist, let alone available for study. Funding for this research was generously provided by the Department of Biological Sciences at the University of Alberta, who actually paid me to teach. Dr. Ruth Stockey supplemented this out of her own research funding for the first three summers (NSERC A-6908). The Natural Sciences and Engineering Research Council of Canada supported the final two years of this work (NSERC PGS-D), suggesting the very height of a civilization so wealthy that it knowingly funds work that is as interesting as it is inapplicable. Travel funding during my thesis research was provided by Dr. Ruth Stockey, the Faculty of Graduate Studies and Research and the Department of Biological Sciences. The Paleobotany Lab at the University of Alberta was a happening place during my tenure. The many people who populated it helped in countless ways. While many of them now have new titles, I knew them as Brian Rankin, Nick Wiebe, Stefan Little, Selena Smith, Steve Karafit, Simone Mageaux, A. J. MacLean, Lindsay Elliott, Kirsten Brink, Matt Vavrek and Genaro Rodrigo Hernandez-Castillo. All of them participitated in this research, sometimes in the field, sometimes in the lab, sometimes in discussion and many times in all cases. Outside of the lab but within the department, there was always stimulating company in the mycology lab, where Marcie Plishka, Sam Skinner, Connie Fe-Gibas, Winnie Wang, Marie Davie and Melissa Day were typically helpful and convivial. Tara Narwani and Takuya Konishi likewise provided the always-necessary outsider's perspective. Outside the University, Pam Clark and Eric Smith gave me a second home in Edmonton. Dr. Wilf Schofield, Professor Emeritus in the Department of Botany at the University of British Columbia offered a crucial suggestion regarding the Fagus fossils. Drs. Gar Rothwell, Ohio University, Patrick Herendeen, George Washington University, Else Marie Friis, Swedish Museum of Natural History and Steven Manchester, University of Florida all provided helpful feedback on this work. My immediate committee deserves more than a platitude for the meetings (formal and informal), exams and proposals that they oversaw. My supervisor, Dr. Ruth Stockey, provided an exceedingly memorable experience that went beyond paleobotany. Fieldwork, lab work, manuscript work, parenting work and my vernacular were all enlivened by her efforts and encouragement. The guidance and feedback from Drs. Randy Currah and Michael Caldwell is greatly appreciated. Outside of this trio, Drs. Enrico Scarpella and Charles Schweger both served on my candidacy committee, while Dr. Alex Wolfe served on my defense committee. Dr. Paul Manos, Duke University, came all the way up from North Carolina to serve as my external and allowed for rare, precious, uninhibited, nut-centric conversations. Dr. David Cass, while never serving on any of my committees, nevertheless provided helpful input on my work, as did Dr. Jocelyn Hall. Of course these acknowledgements would be incomplete without mention of my family. My mother and father, brothers and sister and in-laws all showed good humor and encouragement with regard to this endeavor. Eileen Gibson, my grandmother remains happy to look at rocks. Evelyn Mindell, my late grandmother, advocated on behalf of and literally made possible the education of all her grandchildren as naturally as she served bagels. My wife Barbara is more engaging than even the most fossiliferous of concretions and cannot be thanked enough. Our daughter Claire showed up late in the