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Paleobiology, Biostratigraphy, and Taphonomy of Neoproterozoic Eukaryotes and Cambrian

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Paleobiology, Biostratigraphy, and Taphonomy of Neoproterozoic Eukaryotes and Cambrian Paleobiology, Biostratigraphy, and Taphonomy of Neoproterozoic Eukaryotes and Cambrian Animals with Carbonaceous Preservation Qing Tang Dissertation submitted to the faculty of the Virginia Polytechnic Institute and State University in partial fulfillment of the requirements for the degree of Doctor of Philosophy In Geosciences Shuhai Xiao Sterling Nesbitt Benjamin C. Gill Brian Romans October 10, 2018 Blacksburg, Virginia Keywords: Paleobiology, Biostratigraphy, Taphonomy, Carbonaceous preservation Copyright 2018 © Q. Tang Paleobiology, Biostratigraphy, and Taphonomy of Neoproterozoic Eukaryotes and Cambrian Animals with Carbonaceous Preservation Qing Tang ABSTRACT Carbonaceous fossil preservation is an important taphonomic window that provides critical perspectives on the evolutionary history of life. However, phylogenetic interpretation of carbonaceous fossils is not straightforward. This is largely because critical biological information is usually lost during fossilization and three-dimensional morphologies are flattened into two- dimensional compressions. Hence, innovative techniques and methods are required in order to better understand the evolutionary significance of these fossils. To achieve this goal, this dissertation is focused on using an array of innovative research techniques to investigate the paleobiology, biostratigraphy, and taphonomy of carbonaceous fossils in critical times of early life evolution, including Neoproterozoic and Cambrian. Chapters 2–5 in this dissertation present original research that helps to decipher hidden biological structures of various carbonaceous fossils using a series of research methods. An improved understanding of these carbonaceous remains will ultimately advance our knowledge regarding the early evolutionary history of life on Earth. Chapter two describes new cellular structures of the carbonaceous compression macrofossil Chuaria using backscattered electron scanning electron microscopy. The data show that Chuaria, which is one of the most common fossils in Neoproterozoic and whose phylogenetic interpretation has been uncertain, is likely a multicellular eukaryote. Chapter three is aimed to resolve a long debate on the depositional age of the Gouhou Formation in the Huaibei region of North China and to constrain the Precambrian-Cambrian (P-C) boundary in this area. Using a low manipulation maceration technique, this study reveals a diverse assemblage of organic-walled microfossils from the lower Gouhou Formation, suggesting that the lower Gouhou Formation is Tonian in age and the P-C boundary may be located within the Gouhou Formation. Chapter four reports a group of problematic carbonaceous compression macrofossils from the Hetang Formation in South China. Taphonomic analysis using optical and electron microscopy tentatively suggests that these carbonaceous macrofossils are probably carapaces of bivalved arthropods. The last chapter describes a group of sponge fossils with carbonaceous preservation from the early Cambrian Hetang Formation in South China. Using an array of electron microscopy techniques, this study reveals that siliceous spicules of the Hetang sponges have large axial filaments and large proportions of organic material, suggesting early sponge in the Precambrian and Cambrian may have had weakly mineralized or entirely organic skeletons. Results from this study helps to reconcile the apparently conflicting molecular clocks, biomarker fossils, and spicular fossils of early sponges. GENERAL AUDIENCE ABSTRACT Carbonaceous fossils can provide important information about the life on Earth in deep time. However, biological interpretation of carbonaceous fossils is not always straightforward, largely because critical biological information is usually lost during fossilization. To address this matter, this dissertation presents original research that helps to decipher hidden biological structures of various carbonaceous fossils using a series of innovative research techniques and methods. Specifically, Chapter two describes new cellular structures of the carbonaceous compression macrofossil Chuaria using backscattered electron scanning electron microscopy. The data show that Chuaria, which is one of the most common fossils in Neoproterozoic and whose biological interpretation has been uncertain, is likely a multicellular eukaryote. Chapter three is aimed to resolve a long debate on the depositional age of the Gouhou Formation in the Huaibei region of North China and to constrain the Precambrian-Cambrian (P-C) boundary in this area. Using a low manipulation maceration technique, this study reveals diverse organic- walled microfossils from the lower Gouhou Formation, suggesting that the lower Gouhou Formation is Tonian in age and the P-C boundary may be located within the Gouhou Formation. Chapter four reports a group of problematic carbonaceous compression macrofossils from the Hetang Formation in South China. This study, using optical and electron microscopy, suggests that these carbonaceous macrofossils are probably carapaces of bivalved arthropods. The last chapter describes a group of sponge fossils with carbonaceous preservation from the early Cambrian Hetang Formation in South China. Using an array of electron microscopy techniques, this study reveals that the Hetang sponges developed spicules with large proportions of organic material, suggesting early sponge in the Precambrian and Cambrian may have had weakly mineralized or entirely organic skeletons. Therefore, an improved understanding of these carbonaceous remains presented in this dissertation will ultimately advance our knowledge regarding how the early life on Earth evolved through time. DEDICATION I dedicate this work to my mother, father, wife, and brother, who have been tremendously supportive to me and keep me mindful of the past and present, and more importantly, hopeful about the future. iv ACKNOWLDEGEMENTS I would like to thank those who have supported my pursuit of education; contributed to my development as a scientist; and played a role in my personal establishment. Certainly, it is impossible to acknowledge all the help I have received through the years. Still, I would like to take this opportunity to acknowledge and thank, in no particular order, the following exceptional people for their support: • My mother, Lianxiu Chen, who dedicated all her life to her family. • My farther, Cunde Tang, who taught me to be a good person with patience, diligence, and perseverance. • My wife, Yuli Yang. I feel so lucky and thankful to meet her and marry with her on the campus. • My little brother, Li Tang, who I spent memorable childhood with and takes more responsibility than me to take care of our parents ever since I studied abroad. • My fellow graduate students—particularly those in paleontology, including Andrew (Drew 1) Hawkins, Anthony (Drew 2) Muscente, Natalia Bykova, Chris Griffin, Caitlin Colleary, Candice Stefanic, Krista Koeller, Mitchell Riegler, Kiersten Formoso, Dana Korneisel, Devin Hoffman, Brenen Wynd, Morrison Nolan, as well as visiting scholars in the paleo-group Zhou Wang, Wei Wang, Junfeng Guo, Ke Pang, Bin Wan, Qin Ye, Fanfan Kong, and Juliana Okubo—for fun conversations, scientific discussions, constructive feedbacks, and good, peaceful times. • The teachers, professors, and mentors who were instrumental in my education from primary school to college. • My PhD committee: Sterling Nesbitt, Ben Gill, and Brian Romans. • My co-authors and collaborators—Ke Pang, Bin Wan, Xunlai Yuan, Chuanming Zhou, Jie Hu, Guwei Xie, Leimin Yin, Jinlong Wang, Zhe Chen, and Anthony Drew Muscente—who contributed to the work in this dissertation. • The faculty and staff in the Virginia Tech Department of Geosciences, particularly Michelle Stocker, Connie Lowe, and April Newcomer. • Xunlai Yuan, who introduced me to the world of Precambrian paleontology, encouraged my pursuit of PhD abroad, and has continued to support my professional development. • Last but not least, my advisor Shuhai Xiao, who led me to the geobiology world, gave me the opportunity of studying abroad, and has been extremely generous, particularly with his time and knowledge, as an academic adviser, professional mentor, and respected friend. v ATTRIBUTIONS Chapter Two, or “Electron microscopy reveals evidence for simple multicellularity in the Proterozoic Chuaria”, was published in Geology, 2017, volume 45, pages 75–78, by Q. Tang, K. Pang, X. Yuan, and S. Xiao. Q. Tang and S. Xiao conceived the project. Q. Tang, K. Pang, and X. Yuan collected materials. Q. Tang collected all data. Q. Tang and S. Xiao developed the manuscript with contributions from K. Pang and X. Yuan. Chapter three is an updated version of the paper “Organic-walled microfossils from the Tonian Gouhou Formation, Huaibei region, North China Craton, and their biostratigraphic implications”, which was published in Precambrian Research, 2015, volume 266, page 296–318, by Q. Tang, K. Pang, S. Xiao, X. Yuan, and B. Wan. X. Yuan, S. Xiao, and Q. Tang conceived the project, Q. Tang, K. Pang, B. Wan, and X. Yuan collected materials, Q. Tang collected all data, Q. Tang and S. Xiao developed the manuscript with contributions from K. Pang, X. Yuan, and B. Wan. Chapter four has been submitted and is under review in Journal of Paleontology. The initial title of the publication is “A problematic animal fossil from the early Cambrian Hetang Formation, South China”. Co-authors in the manuscript include J. Hu, G. Xie, X. Yuan, B. Wan, C. Zhou, X. Dong, G. Cao, and S. Xiao. S. Xiao
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