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ABSTRACT LAU, on LEE ANNIE. Evolution Of ABSTRACT LAU, ON LEE ANNIE. Evolution of Tree Architecture in the Brazilian Cerrado. (Under the direction of Dr. William Hoffmann). The tropical savanna-forest boundary is commonly characterized by an abrupt transition in vegetation structure and in tree species composition. It has been hypothesized that differences in architecture between savanna and forest trees have an important role in determining the contrasting structural differences between savanna and forest ecosystems. Because of the importance the vegetation structure in determining the ecosystem properties of these systems, I performed a comparative study of tree architecture to examine differences in plant structure of savanna and forest species. To eliminate the potential bias from phylogenetic relatedness, I used congeneric species pairs containing trees of both habitat types that occur sympatrically in savannas of the Brazilian cerrado habitat at IBGE Ecological Reserve (RECOR). I found that relative to savanna species, forest species have larger crown volumes with more apical meristems and greater height for a given stem diameter. Other traits that influence patterns of light interception also differed, with savanna species exhibiting more convoluted leaf blades and shorter petioles. There was evidence that allometry and other traits are convergent in savanna and forest tree species across lineages, providing strong support for adaptive functions of these traits. Furthermore, the larger canopies of forest species imply that they play a role in reduced light in the understory and the exclusion of grasses, which potentially facilitates further expansion of forest tree species in the absence of fire. Evolution of Tree Architecture in the Brazilian Cerrado. by On Lee Lau A thesis submitted to the Graduate Faculty of North Carolina State University In partial fulfillment of the Requirements for the degree of Master of Science Botany Raleigh, North Carolina 2009 APPROVED BY: _______________________________ ______________________________ Dr. William A. Hoffmann Dr. Qiu-Yun (Jenny) Xiang Advisory Committee Chair Advisory Committee Member ________________________________ ________________________________ Dr. John S. King Dr. Robert R Dunn Advisory Committee Member Advisory Committee Member DEDICATION For my family and friends. ii BIOGRAPHY On Lee Lau was born in Hong Kong and immigrated to San Francisco with her family to spend her formative years learning what science was about as a volunteer and intern at the Calfornia Academy of Sciences, and gaining an appreciation for the outdoors on trail projects with the Student Conservation Association. She studied the ecophysiology of larkspur species ( Delphinium) in an undergraduate Biology thesis at Reed College with Dr. Keith Karoly and measured three-spine sticklebacks ( Gasterosteus aculeatus ) as a lab manager and field technician at University of Texas at Austin with Dr. Dan Bolnick before pursuing a graduate degree. She has done various types of scientific and conservation work in Alaska, California, Costa Rica, Hawaii, Oregon, Texas, Trinidad and Tobago, Vancouver Island BC, and now North Carolina and Brazil. She hopes the future will bring fulfilling work and opportunities to explore more ecosystems. iii ACKNOWLEDGMENTS The author thanks Travis Ingram for comments that significantly improved previous versions of this thesis and for R coding assistance. Also thanks to the super lab group for technical support and insightful discussions: Dr. Sybil Gotsch, Dr. Erika Geiger, Susan Jaconis, Renee Marchin, Alice Wines, Kristen Kostelnik, Wade Wall, Dr. Tom Wentworth, Dr. Marc Johnson and in Brasilia, Marina Calvahlo and Dr. Augusto Franco. And thanks to the esteemed master’s committee, the plant biology office staff, and the thesis editor for providing invaluable support towards graduation. The IBGE Ecological Reserve (RECOR) in Brasilia, DF, Brazil provided great infrastructure and lodging in the Cerrado that made this work possible. Funding for this work was supported by an NSF grant to Dr. William Hoffmann, who cannot be thanked enough for making field work in Brazil possible and being an extremely supportive graduate adviser. iv TABLE OF CONTENTS LIST OF TABLES ............................................................................................................vii LIST OF FIGURES......................................................................................................... viii CHAPTER I. Introduction to the Brazilian Cerrado ........................................................ 1 Evolution of Tree Architecture in High vs Low Light Environments .......................... 4 Effects of Tree Architecture on Ecosystem Properties in the Cerrado ......................... 5 Literature Cited .......................................................................................................... 7 CHAPTER II. Evolution of Tree Architecture and Allometric Relationships in the Brazilian Cerrado ................................................................................................. 13 Introduction ............................................................................................................. 13 Methods ................................................................................................................... 17 Study Site........................................................................................................ 17 Species Selection ............................................................................................ 18 Trait Measurements ......................................................................................... 19 Data Analysis .................................................................................................. 20 Results ..................................................................................................................... 22 Discussion ............................................................................................................... 27 Conclusion ...................................................................................................... 29 Literature Cited ........................................................................................................ 30 CHAPTER III. Differences in Leaf Traits of Savanna-Forest Congeneric Species in the Brazilian Cerrado .................................................................................................. 35 Introduction ............................................................................................................. 35 Methods ................................................................................................................... 38 Study Site........................................................................................................ 38 Species Selection ............................................................................................ 38 Trait measurements and calculations ............................................................... 39 Data Analysis .................................................................................................. 40 v Results ..................................................................................................................... 41 Discussion ............................................................................................................... 47 Literature Cited ........................................................................................................ 50 APPENDIX ........................................................................................................................ 54 vi LIST OF TABLES CHAPTER II. Evolution of Tree Architecture and Allometric Relationships in the Brazilian Cerrado Table 1. List of Species in the Architectural Analysis .............................................. 19 CHAPTER III. Differences in Leaf Traits of Savanna-Forest Congeneric Species in the Brazilian Cerrado Table 1. Means of Leaf Lengths and Petiole Lengths .............................................. 46 Table 2. Variance components of Genus and Species for Multiple Traits ................. 47 APPENDIX Table 1. List of Species in the Leaf Trait Analysis ................................................... 55 vii LIST OF FIGURES CHAPTER I. Introduction to the Brazilian Cerrado Figure 1. Cerrado habitats .......................................................................................... 2 CHAPTER II. Evolution of Tree Architecture and Allometric Relationships in the Brazilian Cerrado Figure 1. Allometric relationships in 15 congeneric species pairs ............................. 24 Figure 2. Least squared means of crown traits in congeners .................................... 25 Figure 3. Correlation of leaf blade area to number of branch tips ............................. 26 CHAPTER III. Differences in Leaf Traits of Savanna-Forest Congeneric Species in the Brazilian Cerrado Figure 1. Means of leaf angle traits in congeners ..................................................... 44 Figure 2. Means for leaf waviness in congeners ...................................................... 45 viii CHAPTER I Introduction to the Brazilian Cerrado The Cerrado region of Brazil is characterized by diverse xeromorphic vegetation that varies in structure from open grasslands (campo limpo) to dense woodlands (cerradao) and riparian gallery forests (Eiten 1972, Oliveira-Filho and Ratter 2002; Figure 1). The Cerrado occupies an
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