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Earth, Wind, and Fire: Resource Allocation and Dispersal Strategies of Rhynchospora knieskernii (Cyperaceae) in a Disturbance-Dependent Ecosystem A Thesis Submitted to the Faculty of Drexel University by Marilyn Carolyn Sobel in partial fulfillment of the requirements for the degree of Doctor of Philosophy June 2015 ii Dedication To my wonderful and supportive family Elizabeth Amanda Sobel Benjamin Carmen Sobel Marcus J Sobel iii Acknowledgements The pursuit of a doctoral degree is a long task that involves the help and support of many people. As a returning student burdened with a full-time job, I benefited greatly from a wide network of people that made it possible for me to complete my thesis. I would like to thank my advisor, Dr. Walter F. Bien for sharing his vast knowledge of the New Jersey Pine Barrens as well as many other areas of ecology. I am so grateful for his support, kindness, friendship, and endless supply of jokes. From the first day of his Pine Barrens Ecology class to the day of my defense Dr. Bien worked to enhance, develop, and polish my scientific understanding and knowledge of ecology. Dr. Bien was always willing to take time to explain concepts and procedures, share books and articles, and teach me a wide range of techniques. He was understanding about my time constraints and willing to work around my schedule. He truly made me a scientist. I also received expert help from my wonderful committee. I am especially grateful to Dr. Dennis M. Gray. He taught me many techniques for analyzing nitrogen and phosphorus, assisted directly in the most technical areas of analysis, made several fruitful suggestions for my research that took it in interesting directions, and generously shared his knowledge and ideas in many discussions of my results. My committee chair, Dr. Michael O’Connor, patiently explained several statistical concepts and was also a great teacher of biophysical ecology, modeling, physiological ecology, and biostatistics. I used information from his classes every single day while writing my thesis. Dr. Gerry Moore shared his botanical expertise about Rhynchospora knieskernii as well as the many associated graminoids. His experience in the field and knowledge of plant structure and taxonomy were invaluable. Dr. James Spotila kindly shared both knowledge and equipment. I could not have conducted the water potential studies without him. iv It would not have been possible for me to write my chapter on community assembly without the help of several New Jersey Pine Barrens botanists. I would like to thank Terry Schmidt, Mark Szutarsky, Donna McBride, and Tom Besselman for patiently sorting through hundreds of plants to ensure all the associated species were correctly identified. Their good humor and expertise turned a tedious task into a special experience. Our laboratory of mavericks provided friendship, support, laughter, and strength when the going got tough. From day one Dane Ward made me feel truly a part of the lab. Thanks to him I got to play king of the mountain, pine cone softball, and broom hockey. We had long discussions about science and conservation and I benefited from his insight and his leadership. Ryan Rebozo was always willing to share data and statistical techniques; he was also quite nice about having a much more attractive study plant. Alina Freire-Fierro shared her extensive botanical knowledge, her specialized taxonomic books, and her sunny personality. Kevin Smith was always helpful with questions about photography and was very supportive during seminars and presentations. Our lab would fall apart without Emily Ostrow, undergraduate extraordinaire. Many thanks for her dedication and willingness to do all kinds of grunt work that kept the lab on track. I am lucky enough to have a very kind and understanding supervisor without whom I could not have taken on this task. I thank Mary Ellen Taggart-Ford of the Steinbright Career Development Center at Drexel University with all my heart for her flexibility, support, and understanding. She even took an interest in snakes, frogs, and my special plant. I am also very appreciative of the support of my colleagues Lauren McHale, Angela Brennan, Nancy LeClair, Catherine Rooney, and Jeanine Rastatter. Mr. Peter Franks, Director of the SCDC, was an early facilitator of my dreams. Many volunteers helped me in the laboratory and field, including Gabrielle Farrell, Jodi Baumgarten, Stuart Berg, Donna Bridger, Christopher Ball, Christopher Funk, and Holly Hagy. I would also like to thank Dr. Roger Marino of Drexel University for his help with the hydrochory v experiments using the flume tank, Robert Cartica of the New Jersey Department of Environmental Protection for assistance with Rhynchospora knieskernii site locations, and Jeremy Markuson of the United States Fish and Wildlife Service for assistance in receiving my permit. Also warm thanks to Major Richard DeFeo and the personnel at Warren Grove Range for logistical support. Last, but certainly not least, I have to thank my wonderful family. My son Ben Sobel and daughter Lizzy Sobel were excited about my doctoral work from the very beginning. They were only 12 and 10 when I started and could not have known what they were going to have to put up with, but they were always gracious and acted as if they did not mind how often I was away in the field. They also helped me check my data, weighed plants, and assisted with my research in many other ways. My husband Marc Sobel put up with having no car most weekends, answered endless questions about statistics, and took care of Ben and Lizzy when I was in the field. I also want to thank my mother-in-law, Florence Sobel for her interest, support, and good wishes. I am sorry my own parents are not alive to enjoy this day, but I am grateful for their focus on the importance of education. I owe a great deal to my mother Jane Carter for demonstrating how happiness is something you create through optimism, hard work, and personal effort. Her example has been a life-long inspiration for me and helped to make this thesis possible. My research received financial support from the 177th Fighter Wing of the New Jersey Air National Guard, the Laboratory of Pinelands Research at Drexel University, and the Philadelphia Botanical Club (Bayard Long Award). vi TABLE OF CONTENTS LIST OF TABLES .............................................................................................................. x LIST OF FIGURES .......................................................................................................... xii ABSTRACT ..................................................................................................................... xix INTRODUCTION .............................................................................................................. 1 1.1. The Genera Rhynchospora .......................................................................................... 1 1.1.1 Study Species ........................................................................................................... 2 1.1.2 Study Sites ................................................................................................................ 4 1.2 Dissertation Structure ................................................................................................... 5 1.2.1 Chapter 2: Earth, Wind, and Fire: Rhynchospora knieskernii Allocation Strategies under Differing Conditions of Resource Availability ...................... 5 1.2.2 Chapter 3: Interactions of Rhynchospora knieskernii with Associated Species and Impacts on Community Assembly .................................................................. 6 1.2.3 Chapter 4: Dispersal Strategies and Germination Cues in a Fire- Dependent Ecosystem ......................................................................................................... 7 1.2.4 Chapter 5: Conservation Implications and Future Directions .................................. 8 Literature Cited ................................................................................................................. 10 2. EARTH, WIND AND FIRE: RHYNCHOSPORA KNIESKERNII ALLOCATION STRATEGIES UNDER DIFFERING CONDITIONS OF REOSOURCE AVAILABILITY ..................................................................................... 15 Abstract ............................................................................................................................. 15 2.1 Introduction ............................................................................................................... 16 2.1.1 Nitrogen Cycling ..................................................................................................... 17 2.1.2 Phosphorus Cycling ................................................................................................ 18 2.1.3 Prescribed Burning and Fire Effects ....................................................................... 19 2.1.4 Rare Plants .............................................................................................................. 20 2.1.5 Resource Allocation ................................................................................................ 23 2..2 Methods .................................................................................................................... 25 2.2.1 Soil Sampling .......................................................................................................... 25 vii 2.2.2 Plant Measurements ...............................................................................................
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