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Copyright by Natalia Beatriz Biani 2009 Copyright by Natalia Beatriz Biani 2009 The Dissertation Committee for Natalia Beatriz Biani Certifies that this is the approved version of the following dissertation: Alliances and Struggles in the Miniature Ecosystem of a Socially Flexible Bee Committee: Ulrich G. Mueller, Co-Supervisor Larry E. Gilbert, Co-Supervisor Mike C. Singer Lauren Ancel Meyers John L. Neff Alliances and Struggles in the Miniature Ecosystem of a Socially Flexible Bee by Natalia Beatriz Biani, M.S. Dissertation Presented to the Faculty of the Graduate School of The University of Texas at Austin in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy The University of Texas at Austin August, 2009 Dedication To my parents, for their gift of life. To Evangelina, Marianela and Juan Francisco, the most marvelous siblings. To Juan, my love. To our son, Ignacio, who is loved even before existing. Acknowledgements Doing a PhD. is not a work of months, not even a year… it involves many years, many moments, many feelings, many struggles. And is not the work of a single person, it is a relatively collaborative process. This dissertation work would not have been accomplished without the guidance, support, and encouragement of Bill Wcislo and Ulrich Mueller. I am infinitely grateful to Bill, who in the midst of uncertainty suggested me to “watch those mites” in Megalopta bee nests. Bill is not only a brilliant and passionate scientist, but more importantly, he is a fantastic person. When I met him for the first time, we had a long talk, about graduate school and science, and it was the most peaceful and friendly talk I have ever had with a scientist. I am thankful to you for teaching me how to do science, for being always so supportive, understandable and kind person and for granting me the privilege to get you know you. I am extremely thankful to Ulrich, who timidly accepted me as one of his students, and rescue me from dropping this whole PhD. project. I am immensely grateful to you for your essentially kind and clever advice and your intellectual and moral support and friendship over these years. Thank you for allowing me to work in your lab and for honoring me with the privilege of being a “muellerita”. I thank Larry Gilbert, who opened the doors of UT and who introduced me to the beauty of tropical forests. The other members of my committee, Lauren Meyers, Jack Neff and Mike Singer were plentiful sources of support, advice and passion throughout this learning process. v I am indebted to Juan Carlos di Trani and Fernando Soley for training me on finding and opening Megalopta’s nests. Field work was both possible and enjoyable thanks to the invaluable help, company and friendship of Hermogenes Fernandez-Marin, Adam Smith, Simon Tierney, Therany Gonzales, Marcos Rios, Cecilia Blundo, Gonzalo Rivas, Piotr Lukasik, and Juan Carlos Santos. Greg Gilbert was a wonderful fungi mentor and taught me the delicacies of these unique and ubiquitous organisms. Hermógenes Fernández-Marín and Julia Sarco provided essential logistic help and advice in the Gamboa mycology lab. Hans Klompen and Jon Shik were essential with the identification of the mite species. Andre Rodrígues suggested the best microbiology method to test the cleaning effect of mites. The Smithsonian community in Gamboa is the most stimulating, supporting and fun environment one can ever hope for during field work. I am thankful to the ones I had the privilege to overlap with. I am grateful to the Ryan Lab for accepting me as a “social parasite” during our summers in Gamboa. I am obliged to the Smithsonian helpful and friendly staff for their logistical support and for their help with the collection and exportation permits. I want to give special thanks to Paola Galgani who has not only allowed this research to be possible, but has been a wonderful friend. Oris Acevedo and Belkys Jimenez are amazingly accommodating and caring persons. Oris, I will never forget the afternoon we returned from BCI in a little boat, avoiding transatlantic ships under a heavy tropical rain. The family who I stayed with in Gamboa, were constant source of laugh, support and gifted me with the thrilling experience of driving Emiliana all the way to Panama City while she was in labor. vi The Mueller Lab is the most wonderful, supportive and fun atmosphere ever. I am thankful to all of you: Barrett Klein and Christian Rabeling, for being such enjoyable, funny and delightful friends and office mates, Rachelle Adams for her support during my first years and beyond, Alexander Mikheyev for his help and advice with the AFLP data and for his constant compliments about my baking skills, Ruchira Sen, Sze Huei, Heather Ishak the sweet and loving ladies of the lab, and Mike Cooper for his help during molecular work. Sandy Monahan could not be sweeter and more compassionate. I thank her for her friendship and for being always so helpful and kind. I will never forget the day you invited me to visit the tower! My friends in Austin and in Argentina are essential to me. I want to specially thank my friend Natalia Ruiz Juri, with whom I came to Austin for the first time, and who I lived with during the first mesmerizing and overwhelming years of grad school in a foreign country. My friend Lidia Marte, la “Arania”, with whom I share innumerable moments of joy, fear, nausea, hope, etc about grad school and life, who always gives me the anthropologist perspective and the beauty of her art, and with who I share the indescribable feelings of being a latinoamerican. My friends from the EEB cohort of the fall of 2002 and beyond are fundamental pillars enriching my life with theirs, and with their extravagant and delicious dishes from all over the world: Samraat, Catalina, Barrett, Simone, Christian, Juanita, Deepa, Evan, Sasha. I could not have made more marvelous friends. I thank Rachel Page, Ximena Bernal and Rachelle Adams, with whom I shared the adventures of grad school, field work, friendship and the amazing journey to motherhood. I am grateful to the music and the blessed musicians I love, for they make our life and our sometime tedious work in the lab, enjoyable and for they bring us awareness of vii unique memories. Borges, Galeano, Cortazar, Fontanarrosa, Maitena, Girondo, Gelman, Benedetti, Neruda, Garcia Marquez, Les Luthiers, are essential to rescue me from the hard questioning moments of grad school, and are sublime sources of inspiration. I feel eternal gratitude to my parents, Ines and Ricardo, who gifted me with the miracle of life, who taught me the love for the natural world and who instilled me the value of learning. I am immensely thankful to Evi, Marianela y Juan Francisco with whom I share the life, for their unique singularity, for their love and undoubtable fraternity. I thank their beloved ones, for making them truly happy. I am grateful to my grandmas for being so “grandmas”. I will never be able to thank all of them enough for their continual, essential and unconditional support and immense love; for giving me the most grandiose welcoming every time I went back to Rosario; and for the wonderful miracle of being present, connected and sharing the whole year throughout time and distance. Juan has being my friend, my guide, my extra advisor, my field assistant, my lab assistant, my personal statistical consultant, my fervent supporter, and above all, the love of my life. I could not have done this without you. Ignacio, our son, is the most wonderful being in my life and I thank him for being such a sweet and patient baby. Finally, I thank the beautiful bees and the amazing mites for allowing me to peep at their lives. This work was supported by the Smithsonian Tropical Research Institute (STRI) Short Term Fellowships and especially by the research grants from the Program of Ecology, Evolution and Behavior since without those little fellowships, none of this would have even started. I am grateful for the teaching assistantships that were granted to me during all these years. Being an independent international student, I would have been unable to support myself otherwise. viii Alliances and Struggles in the Miniature Ecosystem of a Socially Flexible Bee Publication No._____________ Natalia Beatriz Biani, PhD. The University of Texas at Austin, 2009 Supervisor: Ulrich G. Mueller Lawrence E. Gilbert Cooperation is pervasive in nature but paradoxically also provides opportunity to cheaters. My dissertation involves the study of both cooperation and conflict in two species of Megalopta bees. Megalopta is a Neotropical genus of halictid bees whose biology is characterized by complex life cycles that can range from solitary to eusocial. These bees nest in dead wood and forage under dim light conditions. Megalopta ’s nests are inhabited by an extensive array of organisms and each nest therefore constitutes a miniature ecosystem providing opportunities for cooperation and conflict, both within and between species. I first delineate the social structure of M. genalis and M. ecuadoria nests in several Panamanian populations and integrate the factors that play a role in the behavioral decisions of females when joining a social group or not. Within a kin-selection framework, I discuss how genetic relatedness plays a role in the formation of social nests. Second, I investigate the conflict between host bees and a congener social parasite, and I elucidate reproductive structures that are relevant for understanding the evolution of ix parasitism. Finally, I describe a cleaning mutualism between Megalopta bees and their mite associates. Bee-mite associations encompass a broad spectrum of interspecific interactions.
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