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The Role of Frugivores in the Evolution of Fruit in Ficus (Moraceae)

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The Role of Frugivores in the Evolution of Fruit in Ficus (Moraceae) THE ROLE OF FRUGIVORES IN THE EVOLUTION OF FRUIT IN FICUS (MORACEAE) By SILVIA BEATRIZ LOMASCOLO A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY UNIVERSITY OF FLORIDA 2008 1 © 2008 Silvia Beatriz Lomáscolo 2 To my husband and son, who are the compass that keeps me focused on the true North of my life. To my parents, for just giving, from the very beginning. 3 ACKNOWLEDGMENTS I first want to thank Doug Levey, my advisor, for his mentoring, patience, respect for my choices in life and, especially, for believing in me even at times when I doubted of my own capacity to get to the end. Doug was the best adviser I could have asked for my Ph.D. He became a role model as a scientist, teacher, and parent. Every time a piece of equipment broke while I was in the field, Doug rushed to send a replacement piece half a world away. Moreover, the equipment always came accompanied by many chocolate bars, an important “pat in the back” for a woman in despair in the field. The other members of my committee, Rebecca Kimball, Ben Bolker, Mike Miyamoto, and Emilio Bruna, offered great insight from different perspectives. Drs. David Dilcher and Colin Chapman were once part of my committee. They had to be replaced for regulatory and logistic reasons, but were very helpful at the beginning stages of my dissertation. Special thanks go to Rebecca and Ben, who were extremely helpful with data analysis and always willing to offer their time to brainstorm for new ideas on how to tackle my (uncooperative) data. Without Rebecca, the phylogenetic analysis would have been extremely difficult to tame. Without Ben, I would still be wandering aimlessly through hallways, books, and internet forums. Rebecca and Ben are true assets to the Department of Zoology. My time in Papua New Guinea (PNG) was one of the richest, hardest, and most intense periods of my life. Things would have been much harder and a lot less enjoyable without the help of Andy Mack, Ross Sinclair, Debra Wright, and the rest of the Wildlife Conservation Society PNG crew. They unselfishly helped in every aspect of my field work, offering resources and (at times much needed) emotional support. Moreover, they became long-lasting friends. Mónica Alberto and Laurent Bertin hosted me for weeks at a time while I anxiously waited for equipment to arrive, or for rides to the field. I am immensely thankful for their endless 4 generosity, and for offering me a place to stay, when all else failed. Madang wouldn’t have felt like such a familiar place without them. The management at Jais Aben Resort was also generous to offer an extremely good deal to a low-budgeted graduate student. The staff were always extremely nice and made me feel “at home”. Big thanks go to Gisele Maisonneuve too, who shared her house with us a few times, facilitating my work with the coastal fig species around Madang. Unknowingly, she provided a safer environment for the first month of development of our child. Volatile compound work would not have been possible without the help of Peter Teal, Hans Alborn, and Barbara Dueben, who is no longer with us. They are great scientists and amazing people who “adopted” me and helped me without asking anything in return. We need more people like them in science (and everywhere, really). Mark Whitten of the Florida Museum of Natural History also helped me in the fledgling stages of the volatile work. He helped me get started and he led me to Peter Teal, two events that made the volatile compound work a reality. Many people in the Department of Zoology helped in many aspects of my dissertation. Brian McNabb with his contagious curiosity and love for PNG helped me financially and logistically with my first trip to explore that amazing land. He was right: PNG is one of those places on Earth immensely worth seeing. Thanks to many professors, including Ed Braun, Craig Osenberg, Marta Wayne, Larry McEdward, Carmine Lanciani, Richard Kiltie, Colette St. Mary, and others for interesting discussions in their offices or during courses that in one way or another helped to shape this dissertation as it is today. Karen Pallone was always vigilant in helping me over administrative hurdles; I don’t know what Zoology graduate students would do without such an extremely efficient person in our Department. Pete Ryschkewitsch helped me prepare 5 for my fieldwork in many ways and always made my trips to the basement a fun break from my computer. Ken Albergotti and Jared Wright are the most efficient and knowledgeable computer people we’ve had since I came to the Department. I thank them for their many trips to the 6th floor to save me from my computer illiteracy. Thanks also go to Kent Vliet, Kenetha Johnson, Tangelyn Mitchell, Lelia Long, and Jimie Norton from Biological Sciences, who helped support me as a Teaching Assistant. The many hours spent on the 6th floor led me to meet great people who have come and gone through the Department, but who will remain an important part of my life. Special thanks go to Manjula Tiwari, Andreas Heyland, Cristina Dockx, Cedric Worman, Toshi Okuyama, Martin and Veronika Schaefer, and Charlotte Skov, who are long-lasting friends, and who I’ll take home with me in my fond memories. The collaborations I developed throughout my Ph.D. ended up being an important part of this dissertation. I thank George Weiblen for introducing me to Ohu and for his constant help with PNG figs. I want to especially acknowledge Martin Schaefer and Pablo Speranza for fun and productive collaborations. I hope there are many more. Thanks also go to Eric Dyreson for walking me through EPCA, a statistical analysis that seemed written to fill a gap in my dissertation, and for patiently answering all my questions (some of them, more than once). Tamaki Yuri pointed me to EPCA and helped me weave through Mesquite and get the analysis to work. I also want to thank Nina Rønsted for generously providing sequences for all the fig species I needed, before she had even published her work. Thanks go to my undergraduate volunteers, who spent long hours entering and juggling data. I really hope they gained as much as I did from the experience. I thank Eliza Chiang and Derek Cox for sticking with me until the very end. Jamie Morris, Sara Ouly, and Sabrina Simms 6 also helped at different stages. Kore Tau and Francesca Dem, from PNG, helped me by taking full charge of parts of my project, even after I left PNG. They also helped me get to know the mysterious Melanesian culture a bit better. Thanks go to the people in Ohu, especially to my field assistants Amos, Frank, and Ismael, without whom I couldn’t have gathered half of my data. To Brus Isua, for sharing his immense knowledge on the figs of Madang, and to the whole Isua family for hosting me for so long. To Lucy and the many women of Ohu who have showed me how strong a woman can really be. My parents and my sister are an inherent part of this work because they have always been role models for me. They constantly show me with their example how to be a good professional, a good daughter, a good sibling and, now, a good parent. I still have a lot to learn from them, now from a bit closer. Above all, I want to thank my husband for making enjoyable the (at times arid and rough) journey towards finishing this dissertation. He helped with everything that I have thanked other people for: fieldwork, equipment, ideas, interesting discussions about my work, and, especially, emotional support. He taught me the value of persistence and of self-teaching. He was a pillar that helped me stay on my feet in the toughest moments, and a true partner in this enterprise. He also gave me my child, which is what I’m most thankful for in life. 7 TABLE OF CONTENTS page ACKNOWLEDGMENTS ...............................................................................................................4 LIST OF TABLES.........................................................................................................................11 LIST OF FIGURES .......................................................................................................................12 ABSTRACT...................................................................................................................................14 CHAPTER 1 GENERAL INTRODUCTION ..............................................................................................16 2 CORRELATED EVOLUTION OF FIG SIZE AND COLOR SUPPORTS THE DISPERSAL SYNDROME HYPOTHESIS ..........................................................................20 Introduction.............................................................................................................................20 Methods ..................................................................................................................................23 Study System...................................................................................................................23 General Approach............................................................................................................23 Fruit Trait Data................................................................................................................24 Phylogenetic Data and Analyses .....................................................................................26 Sequences.................................................................................................................26
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