The Nature of Savannas That Replace Dry Forests in Chiquitania, Bolivia
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THE NATURE OF SAVANNAS THAT REPLACE DRY FORESTS IN CHIQUITANIA, BOLIVIA By JOSEPH WILLIAM VELDMAN 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 Joseph William Veldman 2 To Aaron, Ava, and Helen 3 ACKNOWLEDGMENTS First and foremost I thank F.E.“Jack” Putz for his tremendous investments in my development as a scientist. I will be forever grateful to Hope College Professors Greg Murray, Kathy-Winnett Murray, William Mungall, and Mark Northuis who encouraged me to attend graduate school. I also thank my committee members, Emilio Bruna, Michelle Mack, and George Tanner for their suggestions and guidance in both the developmental and final stages of my dissertation. Thanks also to Botany faculty Kaoru Kitajima, Stuart McDaniel, Chris Baraloto, and Jack Ewel for their support. Departmental staff Kathy Jones, Kim Williams, Denise Caswell, Karen Patterson, and Tangelyn Mitchell helped me navigate the University of Florida system and kept research funds flowing to me in Bolivia. My parents Jim and Lorraine Veldman, my sister Katie, and dear friend Nathan Geisler gave me the love and support without which this research would mean little. Lin Cassidy and Camila Pizano are my Gainesville family and will forever be my neighbors even if living on different continents. I spent many weekends running at San Felasco with John Engels, Christine Archer, Alfredo Rios, Santiago Espinosa, Franklin Paniagua, Amy Duchelle, Gaby Stocks, Wendy-Lin Bartels, and Peter Derby. Since returning to Florida I have been blessed with new friends Jared Westbrook, Lucas Majure, Meribeth Latvis, Nick Miles and Alejandro Berrio, as well as friends from Bolivia now at the University of Florida: Marlene Soriano and Martijn Slot. I have been privileged to share offices with a number of graduate students in the Departments of Botany and Biology. My first years at the University of Florida were spent at McCarty Hall in the office of Fire and Chemical Ecology with Morgan Varner and Matt Palumbo. On the 3rd floor of McCarty I also had the camaraderie of Geoff Blate, Bill Grauel, Claudia Romero, Skya Murphy, Ana Eleuterio, Paulo Brando, Camila Pizano, Julia Reiskind and 4 Ann Wagner. In Bartram Hall I am fortunate to work with Vincent Mejibe, Drew Silver, Yuan Zhou, Alexander Shenkin, Xavier Haro, and Mathew McConnachie. During time I spent in Ann Arbor, University of Michigan Professor John Vandermeer generously gave me space in his lab; I sincerely appreciate that David Allen, Jahi Chappell, Heidi Liere, Zach Miller, Doug Jackson, Shalene Jha, Aley Joseph, and Senay Yitbarek shared their office, friendship, and ideas. I am also grateful to University of Michigan Professor Robyn Burnham for sharing her excitement for ecology and inviting me to the LOTS discussion group that includes David Marvin and Becca Sonday. This research was possible because of the hospitality of the Bolivian people - I will forever feel that Bolivia is my second home. I owe a huge debt of gratitude to Bonifacio Mostacedo who encouraged me to pursue research in Bolivia. In Bolivia the people of the Instituto Boliviano de Investigación Forestal (IBIF) gave me friendship, guidance, and support: Bonifacio Mostacedo, Marielos Peña-Claros, Zulma Villegas, Marisol Toledo, Lourens Poorter, Geovana Carreño, Juan Carlos Licona, Alfredo Alarcon, Carlos Pinto, Emma Nuñez, Laly Dominguez, Bety Flores, Israel Ibañez, Miguel Ángel Chaves, Mayra Maldonado, Jose Luis Santivañez, Vincent Vroomans, Saskia Grootemaat, Ari Martinez, Skya Murphy, and Margo Stoddard. I additionally wish to acknowledge the friendship of Elibeth Paredo, Remedios Calle, Eva Maria Rivero, Gabriela Gutierrez, and Javier Chambi. I spent much of my time in Bolivia with Lars Markesteijn, fellow doctoral student and biologist, who shared his friendship and ideas about life and ecology. Field work would not have been impossible without the help of field assistants including: Juan Alvarez, Juan Carlos Alvarez, Israel Melgar, the late Juan Pesoa, Juan Pasavare, and Jose Iraipi. In particular, I am honored to have worked with Don Juan Alvarez of Concepcion who 5 accompanied me tirelessly for months of field work. Throughout Chiquitania, ranchers and rural villagers gave me a place to camp, food to eat, permission to study their land, and directions. The Colombara family deserves special thanks for the use of Estancia Cacarachi and for providing horses to ride to field sites. Thanks to Don Juan of San Francisco, the residents of San Jose, Don Negro of Concepcion and many others. The portions of this research conducted in selectively logged forest were facilitated by the managers and employees of INPA Parket LTD; in particular I thank Paul Rozenboom, William Pariona, and Daniel Velasquez. Don Wilfredo and Don Raul kept my pick-up truck functioning between trips to the field. I was fortunate to use the plant collection at the Herbario del Oriente Boliviano (Museo de Historia Natural Noel Kempff Mercado) and thank botanists Fabiana Mamani, Daniel Soto, Moises Mendoza, and Daniel Villarroel. Nelson Joaquin of the Centro de Investigación Agricola Tropical (CIAT-Santa Cruz) helped me learn about tropical forage grasses. Financial support for this research came from the Garden Club of America Award in Tropical Botany, a National Science Foundation Doctoral Dissertation Improvement Grant, and a University of Florida Alumni Doctoral Fellowship. 6 TABLE OF CONTENTS page ACKNOWLEDGMENTS ...............................................................................................................4 LIST OF TABLES ...........................................................................................................................9 LIST OF FIGURES .......................................................................................................................10 ABSTRACT ...................................................................................................................................12 CHAPTER 1 TROPICAL DRY FOREST REPLACEMENT BY SAVANNAS ON THE SOUTHERN EDGE OF THE AMAZON BASIN .................................................................14 Introduction .............................................................................................................................15 Materials and Methods ...........................................................................................................19 Study Area .......................................................................................................................19 Satellite Image Analysis ..................................................................................................19 Savanna Floristic, Fuels, and Soil Analyses ....................................................................21 Results.....................................................................................................................................24 Satellite Image Change Analysis .....................................................................................24 Savanna Floristics ............................................................................................................25 Fuel Loads and Fire History ............................................................................................29 Soil Fertility and Grass Species Composition .................................................................29 Discussion ...............................................................................................................................30 Conclusion ..............................................................................................................................34 2 SELECTIVE LOGGING AND FIRE AS DRIVERS OF ALIEN GRASS INVASION IN A BOLIVIAN TROPICAL DRY FOREST ......................................................................45 Introduction .............................................................................................................................46 Methods ..................................................................................................................................48 Study Site .........................................................................................................................48 Grass Species Distributions .............................................................................................49 Alien Grass Responses to Fire .........................................................................................50 Data Analysis ...................................................................................................................52 Results.....................................................................................................................................53 Grass Species Distributions .............................................................................................53 Alien Grass Responses to Fire .........................................................................................55 Discussion ...............................................................................................................................56 Grass Species Distributions .............................................................................................56 Alien Grass Responses to Fire .........................................................................................59 Conclusions.............................................................................................................................60