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The Roles of Fecal Cases in Neochlamisus Life History

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The Roles of Fecal Cases in Neochlamisus Life History BEHAVIORAL ECOLOGY OF NEOCHLAMISUS LEAF BEETLES: THE ROLE OF FECAL CASES AND ASSOCIATED TRAITS By Christopher G. Brown Dissertation Submitted to the Faculty of the Graduate School of Vanderbilt University in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY in Biological Sciences May, 2009 Nashville, Tennessee Approved: Professor Daniel J. Funk Professor Dave McCauley Professor Patrick Abbot Professor Manuel Leal Professor Jim Fordyce This work is due to my parents, Glen and Sylvia Brown, whose loving care brought me to the top of the hill and to my beloved wife, Jennell Talley, whose patience and (mostly) gentle encouragement pushed me to the other side of it. ii ACKNOWLEDGEMENTS They say it takes a village to raise a child and it probably takes that much or more to instill a doctorate. This work was supported by the financial help of a Student Research Grant from the Animal Behaviour Society, an Exploration Grant from the Explorers Club, and a Dissertation Enhancement Grant from Vanderbilt University, as well as NSF IOB 0616135. I appreciate all the support I received from my major advisor, Daniel J. Funk, who allowed me to work on his beetle babies and yet not on his main passion, their process of speciation. I am also grateful for the other members of my committee, Dave McCauley, Patrick Abbot, Jim Fordyce, and Manuel Leal who not only gave me wings to try new directions, but boundaries that kept me grounded in possibilities. Through them I have learned what it means to be a scientist. Thanks to Roz Johnson, who always had an answer or knew where to find one. I will forever remember the people that I met during this journey. Thanks to my cohort: Dr. Jennifer "Flowergirl" Ellis and Natasha Sherman, we did it together. Thanks to the hall: Dr. David Wills, Dr. and Mrs. Mark Chapman, Vikram Chakre, Jesse Wenzler, Eric Janson, Dan Erickson, Officer Mark Mandel, Ryan Waples, and John "Nutmeg" Gibbons. Thanks to those not on the hall, but who might have wished they were: Jenifer Ferguson, Robin Bairley, Elizabeth Thatcher, and Corey Snelson. And of course, I owe a great debt to the rambling gamblers: Scott P. Egan and Dan Duran. These people taught and retaught me how to laugh, to exercise, how to ignore with conviction, to speak up when it may or may not be necessary, to catch beetles with cheese wrappers, iii to listen to birds, to listen to rap, to eat more healthily, the rules of roller derby, how to care passionately about subjects that no one else in the room has even heard of, about the theory of expanding Earth, to live life to the fullest, how to walk a 10-minute mile, and to appreciate every possibility, especially if there is evidence. Thanks also goes to many friends and family, most of which I knew before I came to Vanderbilt, others I've met since then. I would like to thank my original family: Glen, Sylvia, and Matt Brown and Doris Peterson, my extended family: Mr. And Mrs. Talley, Houston, and Samantha. Thanks to friends from home: the Hutchisons and the Winters, and friends from abroad: the Vanhooses, Lovelesses, Campbells, RJ, and Derek. I also want to remember Gould and Ouida Brown and Clarence "CC" Peterson, who told me repeatedly that education was the most important gift I could ever give myself. But above them all, I thank Jennell Talley, who stood beside me when I couldn’t stand up, who listened to me when I should have shut up, and who loves me no matter what. Without her here, none of this would have happened. Everything I am is for and because of her. She constantly reminds me what it means to be human and that simply being human is not good enough. I love you. iv TABLE OF CONTENTS Page DEDICATION .................................................................................................................... ii ACKNOWLEDGEMENTS ............................................................................................... iii LIST OF TABLES ............................................................................................................... x LIST OF FIGURES ............................................................................................................ xi Chapter I. INTRODUCTION ........................................................................................................1 There's no place like home .......................................................................................1 It's a fecal matter ......................................................................................................7 A special case .........................................................................................................10 In the beginning .....................................................................................................16 Follow the directions..............................................................................................25 What's next .............................................................................................................29 References ..............................................................................................................31 II. ASPECTS OF THE NATURAL HISTORY OF NEOCHLAMISUS (COLEOPTERA: CHRYSOMELIDAE): FECAL CASE-ASSOCIATED LIFE HISTORY AND BEHAVIOR, WITH A METHOD FOR STUDYING INSECT CONSTRUCTIONS .....41 Abstract ..................................................................................................................41 Introduction ............................................................................................................42 Materials and Methods…………………………………………………………...47 Study animals .............................................................................................47 Oviposition and egg case construction ......................................................48 Larval biology and a method for studying case construction ....................48 Pupation and adult emergence ...................................................................51 Additional observations .............................................................................51 Results and Discussion ..........................................................................................53 Oviposition and egg case construction ......................................................53 Larval biology ............................................................................................58 Pupation and adult emergence ...................................................................67 Adult biology and phenology.....................................................................71 Comparisons with other camptosomates ...................................................71 v Acknowledgments..................................................................................................78 References ..............................................................................................................78 III. TESTING THE EFFECTS OF FECAL CASES ON SURVIVAL AND GROWTH RATE OF LARVAL NEOCHLAMISUS PLATANI IN FIELD CONDITIONS ...............84 Abstract ..................................................................................................................84 Introduction ............................................................................................................85 Methods..................................................................................................................90 Test animals .................................................................................................90 Host plants ...................................................................................................91 Predator exclusion treatments ......................................................................91 Experimental setup.......................................................................................92 Data collection and statistics ........................................................................92 Results and Discussion ..........................................................................................94 Effect of treatments on survival ...................................................................94 Rate of development in field conditions ......................................................98 Larval wandering .......................................................................................100 Potential factors contributing to mortality .................................................101 Possible competitors ..................................................................................105 Conclusion .................................................................................................106 Acknowledgements ..............................................................................................108 References ............................................................................................................108 IV. ANTIPREDATORY PROPERTIES OF A BIZARRE ANIMAL ARCHITECTURE: HOW COMPLEX FAECAL CASES THWART ARTHROPOD ATTACK .....................................112 Abstract ................................................................................................................112 Introduction ..........................................................................................................113 Methods................................................................................................................118 Study Animals ..........................................................................................118
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