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Functional Ecology of the Obligate Mutualism Between a Fungus Gardening Ant and Its Symbiotic Fungus Jon Nicholas Seal

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Functional Ecology of the Obligate Mutualism Between a Fungus Gardening Ant and Its Symbiotic Fungus Jon Nicholas Seal Florida State University Libraries Electronic Theses, Treatises and Dissertations The Graduate School 2006 Self-Organization and the Superorganism: Functional Ecology of the Obligate Mutualism Between a Fungus Gardening Ant and Its Symbiotic Fungus Jon Nicholas Seal Follow this and additional works at the FSU Digital Library. For more information, please contact [email protected] THE FLORIDA STATE UNIVERSITY THE COLLEGE OF ARTS AND SCIENCES SELF-ORGANIZATION AND THE SUPERORGANISM: FUNCTIONAL ECOLOGY OF THE OBLIGATE MUTUALISM BETWEEN A FUNGUS GARDENING ANT AND ITS SYMBIOTIC FUNGUS By JON NICHOLAS SEAL A Dissertation submitted to the Department of Biological Science in partial fulfillment of the requirements for the degree of Doctor of Philosophy Degree Awarded: Summer Semester, 2006 The members of the Committee approve the Dissertation of Jon N. Seal defended on July 3, 2006. _______________________ Walter R. Tschinkel Professor Directing Dissertation _______________________ Frank Johnson Outside Committee Member _______________________ Frances C. James Committee Member _________________________ Don R. Levitan Committee Member _______________________ Alice A. Winn Committee Member Approved: _________________________________________ Timothy S. Moerland, Chair, Department of Biological Science The Office of Graduate Studies has verified and approved the above named committee members. ii ACKNOWLEDGEMENTS Through the years of this study, countless people have helped me get where I am today. My advisor Walter Tschinkel merits much credit for generating interest and fascination in ant natural history. I was previously someone interested in the social insects that flew, especially wasps, rather than those tiny wingless ‘wasps’ that lived in concealed nests deep in the ground. More importantly, his sharing of his insights into the fungus gardening ants was invaluable and set the stage of not only this dissertation, but also four other projects, including a collecting trip to Panamá. Not only did he tolerate my various side projects that were seemingly tangential to this dissertation, but also indeed supported them in many ways. As a result, our understanding of the behavior, ecology and evolution of this group of ants has been considerably increased. I certainly have had a good experience in all aspects of this dissertation. I am very grateful that my committee was actively involved in my dissertation, even though at times the wisdom of their comments and actions was far from clear. A much stronger dissertation has certainly resulted. My wife Maria seemed to have an endless supply of patience and support that was absolutely necessary for me to survive several nerve-wracking phases. She bore the brunt of my, at times, insane schedule of field work that required me to spend many daylight hours either collecting colonies or monitoring them in the lab. My parents, sister, brother-in-law all offered their support, even though they were quite mystified at my vocational choice that at times consisted of digging holes in the forest in one of the hottest parts of eastern North America for very little money. My fellow graduate students were indispensable with their help of all sorts. My early years benefited greatly from the nearly daily conversations, i.e., arguments, with my lab-mate Sasha Mikheyev. Our arguments solidified many of the projects I conducted during my tenure at FSU, which of course includes this dissertation. Kevin Haight, Joshua King, Kristina Laskis, Chris Smith, Shonna Storz were invaluable at relieving the tension by being in the lab and available for anything from advice to a simple chat. This project would have taken much iii longer if I did not have the cheerful and patient help of Emily Trudeau and Adrian Smith to help me either in the field or in the lab weighing out hundreds of individual ants and placing them in a gelatin capsule for fat extraction or even worse, sorting dead ant colonies from moist fungus garden. I am fairly certain that Adrian will have to have his eyeglass prescription changed in the near future as a consequence. Steve Rehner of the USDA Biocontrol Laboratory in Beltsville, Maryland generously took the time to conduct the molecular assays that were critical in determining the success of the cultivar switches. Margaret Seavy and Steve Miller of the FSU Department of Biological Science Analytical Laboratory assisted with the use of the spectrophotometer and Tom Miller provided a useful piece of equipment. Dr. William H. Outlaw and Danielle Sherdan provided additional support with the chitin assay. The FSU Statistical Consulting Center also aided with several statistical techniques used in this study. Kier Klepzig and Lary Roton of the USDA Southern Research Station in Pineville, Louisiana were extremely helpful in locating the Atta texana colonies. The USDA Forest Service allowed us to collect colonies in the Apalachicola National Forest. Financially my research at FSU was supported by a FSU Dissertation Research Grant, the Horace Loftin Endowment in Graduate Studies (FSU Department of Biological Science), a Short Term Fellowship from the Smithsonian Tropical Research Institute and a grant from the Theodore Roosevelt Memorial Fund from the American Museum of Natural History. Generous funds from grants from NSF, USDA and the Florida State Program Enhancement Program awarded to my advisor, Walter Tschinkel, allowed me to travel to Louisiana and purchase needed supplies and chemicals when I lacked money of my own. His generosity was much appreciated. iv TABLE OF CONTENTS List of Figures ............................................................................................... vi Abstract ..................................................................................................... vii Chapters 1. Literature Review and Conceptual Framework.............................................. 1 2. Complexity and self-organization in an obligate mutualism.......................... 7 Materials and Methods .............................................................................. 9 Results ............................................................................................... 16 Discussion ............................................................................................... 19 3. The Attine Superorganism ............................................................................. 36 Materials and Methods............................................................................... 38 Results ............................................................................................... 45 Discussion ............................................................................................... 51 4. General Conclusions ...................................................................................... 56 APPENDICES A Preference Trials (2002)....................................................................... 59 B Preference Trials (2003)....................................................................... 60 C Preference Trials (2004 (before switch)............................................... 61 D Preference Trials (2004 (after switch).................................................. 62 E DNA Extraction Method ...................................................................... 63 LITERATURE CITED ....................................................................................... 78 BIOGRAPHICAL SKETCH .............................................................................. 86 v LIST OF FIGURES Figure 1:A highly simplified dendrogram of the fungus gardening tribe Attini . 26 Figure 2: Photograph of laboratory colony setup ............................................... 27 Figure 3: Mean number of substrate items collected during preference determination ................................................................... 28 Figure 4: Percentages of total amount of substrate collected in 2003 ................ 29 Figure 5: Fungus garden volume (± SE) as a function of time ........................... 30 Figure 6: Mean percent change in fungus garden volume .................................. 31 Figure 7: Mean total ant biomass (± SE) for each treatment............................... 32 Figure 8: Mean total amounts of substrate (± SE) accepted and final fungus garden weight for data collected in 2003 ................................................... 33 Figure 9: Mean total and percent chitin (± SE) in each fungus garden by treatment. 34 Figure 10: Mean efficiencies of each substrate (± SE) for the production of ant and fungal biomass ........................................................................ 35 Figure 11: Percentages of total amount of substrate collected in 2004 before colonies were given the A. texana cultivar................................................. 68 Figure 12: Percentages of total amount of substrate collected in 2004 after colonies were given the A. texana cultivar. ............................................ 69 Figure 13: Mean (± SE) total amounts of ant performance measured in Joules 70 Figure 14: Mean total sexual biomass (± SE) measured as dry weight (cg) and Joules. ……………………………………………………… 71 Figure 15: Fungus garden volume (± SE) as a function of cultivar and preference over time .................................................................................. 72 Figure 16: Mean (± SE) fungus garden weight (g) ............................................ 73 Figure 17: Mean (± SE) amount of substrate collected by each colony
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