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Systematics, Climate, and Ecology of Fossil and Extant Nyssa (Nyssaceae, Cornales) and Implications of Nyssa Grayensis Sp

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Systematics, Climate, and Ecology of Fossil and Extant Nyssa (Nyssaceae, Cornales) and Implications of Nyssa Grayensis Sp East Tennessee State University Digital Commons @ East Tennessee State University Electronic Theses and Dissertations Student Works 8-2013 Systematics, Climate, and Ecology of Fossil and Extant Nyssa (Nyssaceae, Cornales) and Implications of Nyssa grayensis sp. nov. from the Gray Fossil Site, Northeast Tennessee Nathan R. Noll East Tennessee State University Follow this and additional works at: https://dc.etsu.edu/etd Part of the Biodiversity Commons, Climate Commons, Paleontology Commons, and the Plant Biology Commons Recommended Citation Noll, Nathan R., "Systematics, Climate, and Ecology of Fossil and Extant Nyssa (Nyssaceae, Cornales) and Implications of Nyssa grayensis sp. nov. from the Gray Fossil Site, Northeast Tennessee" (2013). Electronic Theses and Dissertations. Paper 1204. https://dc.etsu.edu/etd/1204 This Thesis - Open Access is brought to you for free and open access by the Student Works at Digital Commons @ East Tennessee State University. It has been accepted for inclusion in Electronic Theses and Dissertations by an authorized administrator of Digital Commons @ East Tennessee State University. For more information, please contact [email protected]. Systematics, Climate, and Ecology of Fossil and Extant Nyssa (Nyssaceae, Cornales) and Implications of Nyssa grayensis sp. nov. from the Gray Fossil Site, Northeast Tennessee ___________________________ A thesis presented to the faculty of the Department of Biological Sciences East Tennessee State University In partial fulfillment of the requirements for the degree Master of Science in Biology ___________________________ by Nathan R. Noll August 2013 ___________________________ Dr. Yu-Sheng (Christopher) Liu, Chair Dr. Tim McDowell Dr. Foster Levy Keywords: Nyssa, Endocarp, Gray Fossil Site, Miocene, Pliocene, Karst ABSTRACT Systematics, Climate, and Ecology of Fossil and Extant Nyssa (Nyssaceae, Cornales) and Implications of Nyssa grayensis sp. nov. from the Gray Fossil Site, Northeast Tennessee by Nathan R. Noll The Late Hemphillian (latest Miocene or earliest Pliocene, 7-4.5 Ma) Gray Fossil Site in northeastern Tennessee is interpreted to represent a lacustrine paleokarst fed by a river or stream. This research focuses on the morphological and systematic relations of Nyssa endocarps (fruit pits) from the fossil site to extinct and extant Nyssa species. A combination of metric and nonmetric traits allows recognition of a new species: Nyssa grayensis sp. nov. This fossil species shares the most similarities with the extant Nyssa ogeche Bartram ex Marshall from southeast North America and the Eocene fossil Nyssa eolignitica Berry from western Tennessee. Affinities with Nyssa ogeche Bartram ex Marshall suggest a warmer winter climate than the present and periodically fluctuating water levels. Fossil vertebrates (Alligator, Heloderma, Hesperotestudo) and plants (Fossil relatives of Ilex vomitoria, Quercus virginiana, Taxodium, Acer leucoderme) with modern counterparts distributed in areas with mild winters support this interpretation. 2 ACKNOWLEDGEMENTS I would like to thank my thesis committee Dr. Liu, Dr. Levy, and Dr. McDowell as well as Dr. Peter Fritsch and Dr. Michael Zavada, for their guidance and support. I would like to thank Dr. Barry Hammel, Daniel Santamaria, Dr.Tao Su, Alan Trently, Dr. Steven Manchester, and the faculty and staff of the Duke Herbarium for assistance in obtaining specimens used in this study. Thank you to Dr. Xiang for providing preliminary results from molecular studies on the extant Nyssa that which assisted my systematic interpreatations. Thank you to Dr. Fred Hossler and Dr. Zavada for your time and assistance with SEM analysis. Thank you to Ms. Alison DePollo for going above and beyond in helping me obtain rare literature that normally is not loaned and for working with other Illiad staff to help me obtain the best possible scans of fossil figures. Thank you to Dr. Mead for providing use of microscopes and other supplies. I would also like to thank the excavators, screen washers, curators, and other faculty and staff at the Gray Fossil Site. Without each role, research on the fossils from the site would not be possible. Funding for this project was supported by the NSF GK-12 Fellowship Program, the ETSU Study Abroad Scholarship, and an NSF CAREER award to Dr. Liu. Finally I would like to thank my family and friends, especially my wife and parents, for the support and patience they have provided during my program. 3 TABLE OF CONTENTS Page ABSTRACT…………………………………………………………………. 2 ACKNOWLEDGEMENTS…………………………………………………. 3 LIST OF TABLES………………………………………………………….... 6 LIST OF FIGURES………………………………………………………….. 7 Chapter 1. INTRODUCTION…………………………………………………... 12 2. MATERIAL AND METHODS……….……………………………... 20 3. RESULTS AND DISCUSSION: Nyssa grayensis sp. nov...………….. 29 Systematic Description of Nyssa grayensis sp. nov.. ………… 29 Plate 1……………………………………..……........... 31 Plate 2………………………………………..……....... 32 Plate 3………………………………………..…........... 33 Distinguishing Nyssa from Related Genera…………….……. 34 Inter-Specific Variation and Species Identification….….…..... 42 Comparisons with Extant Species………………...……….….. 43 Comparisons with Extinct Species ………………………...… 63 Key to the Fruits of Extant and Extinct Nyssaceae Genera…………………………………….……….….. 73 Key to the Endocarps of Extant and Extinct Nyssa………...…. 74 Key 1: Endocarps with Vascular Bundles Mostly Sunken in the Grooves……………………………………..…. 74 Key 2: Endocarps with Vascular Bundles Mostly 4 Raised on the Crests of Ridges…….….….…………... 86 Key 3: Endocarps with Few Traces of Vascular Bundles on the Surfaces……………..……………………...….. 87 Climatic and Biogeographic Implications………….………..… 88 Fruit Size and Climate………………..…….………………….. 97 Environmental Implications…………………………………… 107 4. SUMMARY AND CONCLUSIONS……………..…………………... 110 REFERENCES………………………………………………………………... 112 APPENDIX………………………………………………………….................. 123 Table 8: Specimens of Extant Nyssa Species Used in Study….…….…. 123 VITA…………………………………………………………………………... 127 5 LIST OF TABLES Table Page 1a. Nyssa Fossils from the Gray Fossil Site……………………….………… 21 1b. Specimens of Extant Nyssa Species Used in This Study............................... 22 2. Nyssa Fossil Fruit Species And Sources Used to Construct the “Key to Nyssa Fruit Species………………………..….…………….... 25 3. Definitions of Variables Used to Correlate Climate With Endocarp Size………………………………….…................................ 28 4. Comparisons of Taxa Similar to Nyssa Grayensis Sp. Nov………..……..… 69 5. Climate and Latitude Data for the Ranges of Extant Nyssa Species……………………………………………………..…….…….. 97 6. Fruit Measurements of Extant Nyssa Species…………….…………...…….. 99 7. Correlation of Fruit Sizes and Climatic and Latitudinal Variables......…….... 100 8. Data on Specimens of Extant Nyssa Species Used in This Study…..……….. 123 6 LIST OF FIGURES Figure Page 1. Phylogeny and Molecular Clock of the Cornales……….………….….…... 16 2. The Distribribution of Extant Nyssa Species……………....…………………. 17 3. Schematic Drawing of a Nyssa Fruit……………….……..………………….. 24 4. Randomly Oriented Fibers in the Extant Species Nyssa talamancana Hammel and Zamora……………………..……….. 35 5. SEM Image of Randomly Oriented Fibers of Nyssa talamancana Hammel and Zamora in Section View………………………...……....... 35 6. SEM Image of Fibers of a Nyssa sinensis Oliver Endocarp in Section View………………………………………………………….... 36 7. SEM Image of Randomly Oriented Fibers Toward the Central Portion of the Endocarp Wall in Nyssa sylvatica Marshall…..…….…... 36 8. SEM Image of Randomly Oriented Fibers of Nyssa grayensis sp. nov……... 37 9. Close-Up SEM Image of the Randomly Oriented Fibers of Nyssa grayensis sp. nov……………………..…..…………………... 37 10. SEM Image of Randomly Oriented Fibers of Nyssa grayensis sp. nov. From the Central Portion of the Endocarp Wall in Section View….….. 38 11. Illustration of a Davidia Endocarp Showing Valves That Dehisce Down Most of its Length…………………………………...………...... 40 12. Immature Fruit Heads of Camptotheca acuminata Decne………….…...…. 41 13. Mature Fruits of Camptotheca acuminata Decne………….………...……... 42 7 14. Comparison of the Fossil Nyssa Endocarps from Gray with Extant Species…………………………………………..……………….. 43 15. A Nyssa talamancana Hammel and Zamora Endocarp Before and After Dehydration…………………………………….…………...…….. 45 16. SEM Image of a Pair of Vascular Bundles in Nyssa talamancana Hammel and Zamora Beneath the Surface…………….……...….....…... 45 17. A Vascular Bundle Mostly Beneath the Surface of A Nyssa ogeche Bartram ex Marshall Endocarp...……………………………………....... 46 18. A Vascular Bundle Completely Obscured beneath the Surface of a Nyssa ogeche Bartram ex Marshall Endocarp…………….………...…. 47 19. SEM Image of Vascular Bundles Obscured beneath the Surface of Nyssa grayensis sp. nov………..………………………..……………….. 47 20. Vascular Bundles Obscured below the Surface of Nyssa grayensis sp. nov. Under a Light Microscope…………….………………….…...... 48 21. A Nyssa aquatica Linnaeus Endocarp From Richmond, NC…………..…...... 49 22. Valve Shape of Nyssa grayensis sp. nov………………....………………….. 49 23. Valve Shape of Nyssa ogeche Bartram ex Marshall…………....…………..... 50 24. Boxplot of Extant Nyssa Stone or Endocarp Lengths in Comparison to Nyssa grayensis sp. nov……………………….…...………...………… 51 25. A Cross-Section of Nyssa talamancana Hammel and Zamora Showing Crescent-Shaped Locules That Result from the Valves Thickening toward the Midline……………………………………………..…....…… 53 8 26. Cross-Section Showing the Roughly “D” Shaped Locule of Nyssa ogeche Bartram ex Marshall with an Even Wall Thickness...…….......................
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