ECOLOGY OF NATIVE OIL-PRODUCING PALMS AND THEIR POTENTIAL FOR BIOFUEL PRODUCTION IN SOUTHWESTERN AMAZONIA By JOANNA MARIE TUCKER LIMA 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 2010 1 © 2010 Joanna Marie Tucker Lima 2 To my parents who taught me to appreciate and marvel at the natural world. 3 ACKNOWLEDGMENTS This work is the culmination of a long journey and the encouragement and support of many people throughout my academic career. As this chapter of my life comes to a close, I wish to extend special thanks to my PhD advisor, Karen Kainer, who was a continuous and consistent inspiration and source of encouragement to me. Her expertise and insight enriched my research from beginning to end. I also thank Evandro Ferreira for his contagious love for palms and mix of practical and scientific advice. His students, Janice and Anelena, as well as the research team at the Parque Zoobotanico at UFAC, especially Plinio, Lira, and Edir, were always willing to help with field work and logistics, for which I am very grateful. Anelise Regiane and her chemistry students at UFAC (Thayna, Nubia and Marcia) gave unselfishly of their time to help me process palm fruits and run chemical analysis that I could never have done alone. I am indebted to Francis “Jack” Putz for sharing his curiosity for the natural world and his persistent search for answers to both basic and complex ecological issues that affect our daily lives. His prodding stretched my ideas and encouraged me to look beyond the easy answers. I wish to thank Emilio Bruna and Jane Southworth for their insightful feedback as I developed my dissertation, and for sharing their deep understanding in their fields of expertise. I am also grateful to Meghan Brennan and Christina Staudhammer for their brilliant help with statistical analyses. I am truly grateful for all the financial support I received during my degree from various sources—NSF Integrative Graduate Education Research and Training Program, Tropical Conservation and Development Program (UF), International Palm Society, and the Environmental Protection Agency. I cannot forget to mention the debt I owe to the principle figures during my academic formation, including Emilio Moran, Eduardo 4 Brondizio, Andrew Henderson, and Daniel Zarin. These men inspired me, awakened my zeal for the Amazon, Brazil, and palm trees, and contributed the foundation, both academic and personal, that brought me to this point. Friends and family have given me unconditional support throughout this journey, and I thank them from the bottom of my heart for their patience, wisdom, listening ears, and encouragement. I especially thank my parents who unwaveringly stood by me in the good and the difficult times. Last but not least, I thank my husband, Evandro, without whom this work would have been impossible. I thank him for his reliability and wisdom as he helped with my fieldwork, and for sharing his ideas and intimate understanding of Amazonian forests. I thank him for his patience and willingness to stick it out during the “last year” of finishing-up, and for supporting me unselfishly during that time. Finally, I thank God, for bringing all these wonderful people into my life, for never letting me go, and for providing me the strength, hope, perseverance, intelligence, and faith to complete my dissertation. 5 TABLE OF CONTENTS page ACKNOWLEDGMENTS .................................................................................................. 4 ABSTRACT ................................................................................................................... 12 CHAPTER 1 INTRODUCTION .................................................................................................... 15 2 FLOWER COLOR VARIATION IN ATTALEA PHALERATA (ARECACEAE) ......... 19 Field Observations .................................................................................................. 21 Phenology ........................................................................................................ 21 Flower Color Variation ...................................................................................... 22 Color Polymorphism in Palms ................................................................................. 24 Possible Explanations for Color Polymorphism in Attalea phalerata Flowers ......... 27 Final Considerations ............................................................................................... 28 3 DOES LANDSCAPE CHANGE ALTER REPRODUCTIVE PHENOLOGY, SEX EXPRESSION AND PRODUCTIVITY OF ATTALEA PALMS IN SOUTHWESTERN AMAZONIA? ............................................................................ 30 Introduction ............................................................................................................. 30 Materials and Methods ............................................................................................ 34 Study Area ........................................................................................................ 34 Study Species .................................................................................................. 35 Site Selection ................................................................................................... 37 Data Collection ................................................................................................. 37 Reproductive phenology ............................................................................ 37 Size and illumination measurements ......................................................... 38 Palm dissections ........................................................................................ 38 Statistical Analyses .......................................................................................... 39 Results .................................................................................................................... 40 Comparison of Flower and Fruit Production in Forest and Pasture .................. 40 Flowering and fruiting phenology ............................................................... 40 Flowering and fruiting frequency ................................................................ 43 Associations between Reproduction, Light Availability, and Tree Size ............. 47 Palm Dissections .............................................................................................. 48 Discussion .............................................................................................................. 49 Pasture Palms Out-Produced Forest Palms ..................................................... 49 Phenology ........................................................................................................ 49 Hidden Mechanisms Control Sex Expression ................................................... 50 Sunlight and Plant Size Linked to Sex Expression and Productivity ................. 52 Sex Expression and Male Dominance .............................................................. 53 6 Pollen Availability may Alter Operational Sex Ratios ........................................ 55 Conclusions ............................................................................................................ 56 4 RECOVERY OF ATTALEA PHALERATA MART. EX SPRENG. PALM POPULATIONS AFTER SLASH-AND-BURN AGRICULTURE IN SOUTHWESTERN AMAZONIA .............................................................................. 58 Introduction ............................................................................................................. 58 Study Area .............................................................................................................. 61 Study species ......................................................................................................... 62 Materials and Methods ............................................................................................ 63 Research Design .............................................................................................. 63 Data Collection ................................................................................................. 64 Statistical Analysis ............................................................................................ 65 Results .................................................................................................................... 66 Pre-Burn Palm Populations .............................................................................. 66 Immediate Effects of Slash-and-Burn ............................................................... 68 Recovery .......................................................................................................... 69 Recruitment and survival ........................................................................... 69 Effect of slash-and-burn on A. phalerata populations ................................ 71 Palm demography ...................................................................................... 73 Leaf production and growth ........................................................................ 75 Discussion .............................................................................................................. 77 Palm Colonization or Persistence? ................................................................... 77 Impediments to seedling recruitment ........................................................