FOREST RESPONSES TO CLIMATE CHANGE ALONG AN ANDES-TO-AMAZON ELEVATIONAL GRADIENT BY WILLIAM FARFAN RIOS A Dissertation Submitted to the Graduate Faculty of the WAKE FOREST UNIVERSITY GRADUATE SCHOOL OF ARTS AND SCIENCES in Partial Fulfillment of the Requirements for the Degree of DOCTOR OF PHILOSOPHY Biology August 2019 Winston-Salem, North Carolina Approved By: Miles R. Silman, Ph.D., Advisor Nigel C. Pitman, Ph.D., Chair T. Michael Anderson, Ph.D. Kathleen A. Kron, Ph.D. William K. Smith, Ph.D. Clifford W. Zeyl, Ph.D. DEDICATION To my beloved parents Julio Farfan and Visitacion I. Rios, and my brothers Maikol and Henry Farfan ii ACKNOWLEDGEMENTS First and foremost, I would like to sincerely thank my advisor, Miles R. Silman, for his continuous support of my research, for his encouragement, knowledge, patience and for being a great mentor and friend over the past several years. I deeply appreciate his time, enthusiasm, and encouragement in pushing me beyond the limits my own mind to develop my critical thinking. If I had to do it all over again, undoubtedly, I would do it with Miles. Likewise, I thank my committee members, Dr. Kathleen A. Kron, Dr. William K. Smith, Dr. Michael T. Anderson, Dr. Clifford W. Zeyl, and Dr. Nigel C. Pitman for their time, helpful ideas and comments that enriched my research. I have been immensely blessed by a wonderful and incredibly family, their love with unwavering support were essential to finish graduate school and perform my research. I owe so much gratitude to my beloved parents Julio Farfan, Visitacion I. Rios and my brothers Maikol and Henry Farfan. Collaboration is important to do science and I was extremely fortunate to work closely with great scientists that enriched my research on so many levels. My sincere gratitude to professors Kenneth Feeley, Yadvinder Malhi, Sassan Saatchi, Oliver Phillips, Norma Salinas, Patrick Meir, Greg Asner, Timothy Baker, Paul Baker, Sherilyn Fritz, James Pease, Brian Enquist and Alexander Shenkin. Thanks to my old and new Silman lab friends for their support, comments on the different drafts of the dissertation chapters and helpful ideas to improve my research, iii thanks to Josh Rapp, Danny Lough, Karina Garcia, Rachel Hillyer, Noah Yavit, Cassie Freund, Ellen Quinlan, Max Messinger, Stephanie Bilodeau, John Gorelick, Jared Beaver and Jorge Caballero. Likewise, thanks for the support to my Deacon friends Jenny Howard, Emily Tompkins, Nicholas Huffeldt, Yoyi Fernandez, Scott Cory, Deusdedith Rugemalila, Robbie Baldwin, Felipe Estela, and Mary J. Carmichael. Thanks to the Causita’s group members Jorge Ortiz, Ciro Astete, Armando Alfaro, Jorge Curo, Juan Gibaja, Victor Chama, Juan Costa, Vicky Huaman, Javier Silva, Johnny Farfan, Jean Latorre and Saul Zuñiga for their support and encouragement to finish graduate school and complete my research. Thanks to Alex Nina for his thoughtful assistance translating the abstract in Quechua. Finally, thanks to my friends Leydi, Gaile, Gerald, Janell, Richard and Craig for their support during my graduate school life. I express my deep gratitude to Robin Foster, Abel Monteagudo, Paul Mass, Ronald Liesner, Jun Wen, Kenneth Wurdack, Nancy Hensold, Henk van der Werff, Fabian A. Michelangeli, Lucia Kawasaki, Percy Nuñez, Karina Garcia, Felipe Sinca and Charlote M. Taylor for the help in the botanical identifications. Likewise, my recognition and thanks to the directors and curators of the different herbaria, including the Herbario Vargas (CUZ), Herbario del Museo de Historia Natural- Universidad Nacional Mayor de San Marcos (USM), Herbario Nacional de Bolivia (LPB), al Field Museum (F) and the Missouri Botanical Garden Herbarium (MO), Herbario Selva Central - Oxapampa (HOXA), Herbario Augusto Weberbauer (MOL) for access and permission to work with the botanical collections. Thanks to GBIF, JSTOR, TROPICOS, TNRS and to all the institutions that made available the botanical data essential for this research. iv This fieldwork effort was not one-man labor, it was only possible thanks to an extraordinary group of people with high levels of enthusiasm and hard work. First, I thank Alex Nina, Jhonatan Sallo, Karina Garcia, Luis Imunda, Natividad Raurau, Judith Huaman and Alberto Gibaja for their crucial help in getting everything done coordinating the different aspects of the fieldwork and plants identification process. Lastly but no less important, I thank the Peruvian students that shared with me the extreme fieldwork conditions, leaving sweat, blood, and tears in our field campaigns. Without them, I would not have been able to complete this research. My eternal gratitude to Adam J. Ccahuana, Albino Quispe, Alex Nina, Alexander Quispe, Alex A. Caceres, Andrea Palomino, Angela Rozas, Bryan G. Valencia, Carlos A. Salas, Catherine Bravo, Claudio Lipa, Chaska Chavez, Cristian E. Solis, Cristian Alvarez Galvez, Cristhian Alvarez C., Darcy F. Galiano, David Lopez, Dayana Suni, Dino Tapia, Edith R. Clemente, Erickson G. Urquiaga, Felipe Sinca, Félix F. Farfan, Flor M. Perez, Flor M. Zamora, Fredy Guizado, Ghylmar J. Tinoco, Giuliana M. Palomino, Gladys Castillo, Guido Vilcahuaman, Guisela J. Zans, Irving C. Costas, Israel Cuba, Janet Mamani, Javier E. Silva, , Jesus Castaneda, Jesus M. Banon, , Jhonatan Sallo, Jhoel Delgado, Jhon G. Quispe, Jhon S. Cansaya, Jimmy R. Chambi, Jimy S. Mesicano, Joel Mendoza, Jonyer H. Zapata, Jose A. Quintano, Jose L. Mancilla, Jose Sanchez, Juan A. Gibaja, Juan J. Calvo, Judit Huaman, Julina Pelaes, , Karina Garcia, Karina Luna, Katherine Quispe, Katia Quispe, Kilmenia Luna, Leonit Tueros, Leydi V. Auccacusi, Leyni Caballero, Lucely Vilca, Luis Imunda, Luis Mancilla, Luz M. Cabrera, Manuel J. Marca, Marco A. Maldonado, Maria A. Cuentas, Maria I. Manta, Mario V. Sanchez, Marlene Mamani, Miguel Salas, Milenka X. Montoya, Mireya N. Raurau, Nohemi Lizarraga, Omar Tacusi, Omayra V. Colque, Paul v E. Santos, Paul Iturbe, Paul Ricardo, Percy O. Chambi, Reymerth Peña, Richard Tito, Rosa M. Castro, Rosalbina Butron, Rudi S. Cruz, Ruly Pillco, Shirley J. Serrano, Tatiana E. Boza, Vicky Huaman, Walter Huaraca, Washinton Luis, Wilson Huaman, Yan C. Nina, Yaneth Quispe, Yeferson Davalos, Yessica Halanocca, Yezen Sanchez, Ylenia Moron, Yovana Yllanes, and Yorka Gutierrez. This study is a product of the Andes Biodiversity and Ecosystem Research Group (ABERG; http://www.andesconservation.org/) with contributions for lowland plot data from John Terborgh and affiliated networks RAINFOR (http://www.rainfor.org/), GEM (http://gem.tropicalforests.ox.ac.uk/), and the ForestPlots.net data management utility for permanent plots. SERFOR, SERNANP, and personnel of Manu National Park - Peru provided assistance with logistics and permission to work in the buffer zone and protected areas in Peru. Pantiacolla Tours and the Amazon Conservation Association provided logistical support. Funding came from Wake Forest University through the Vecellio Grant, Elton C. Cocke Travel Award, Alumni Student Travel Award, and Richter Scholarship Award. Funding also came from the Gordon and Betty Moore Foundation’s Andes to Amazon initiative and the US National Science Foundation (NSF) DEB 0743666, NSF Frontiers in Earth Systems Dynamics (FESD) 1338694, and NSF Long-Term Research in Environmental Biology (LTREB) 1754647. The research was also supported by the National Aeronautics and Space Administration (NASA) Terrestrial Ecology Program grant # NNH08ZDA001N-TE/ 08-TE08-0037. Support for RAINFOR and ForestPlots.net plot monitoring in Peru has come from a European Research Council (ERC) Advanced Grant (T‐FORCES, “Tropical Forests in the Changing Earth System”, vi 291585), Natural Environment Research Council grants (including NE/F005806/), NE/D005590/1, and NE/N012542/1), and the Gordon and Betty Moore Foundation. vii TABLE OF CONTENTS ACKNOWLEDGEMENTS....………..…………..………………………………...….iii LIST OF TABLES...…………………………………………………………………...xi LIST OF FIGURES.….……………………………………………………...……….xiii ABSTRACT….……………………….……………………………………………...xvii CHAPTER I Introduction……..…..…………………………………………………………..1 CHAPTER II Landscape-scale wood density variation across an Andes-to-Amazon elevational gradient ABSTRACT………………….………………………………….………..26 INTRODUCTION……………………………….…………………..........28 METHODS…….…...……………………………………………….........32 RESULTS….………...…………………………………………………...38 DISCUSSION………….…………………………………....…………....42 LITERATURE CITED.…………….……………………….……………55 SUPPORTING INFORMATION.…………………..………….…..…….84 CHAPTER III Movement or mortality? Pervasive but slow upslope tree migration in the Amazon to the Andes revealed through 38 years of forest monitoring ABSTRACT…….……………………………………………….………129 viii INTRODUCTION…………………………………………………........131 METHODS………………………………………………………….......135 RESULTS……………………………………………………………….143 DISCUSSION….……………………………………………………......148 LITERATURE CITED.……….……………………………..………….158 SUPPORTING INFORMATION …..…………………….…………….180 CHAPTER IV Long-term stand and carbon dynamics along the Amazon-to-Andes elevation gradient ABSTRACT…………….……………………………………….………211 INTRODUCTION…………………………………………………........213 METHODS………………………………………………………….......217 RESULTS……………………………………………………………….224 DISCUSSION……………….…………………………………………..230 LITERATURE CITED……….…………………………………………238 SUPPORTING INFORMATION ………………………........................263 CHAPTER V An annotated checklist of trees and their relatives in tropical montane forests of southeast Peru: the importance of continued botanical collecting ABSTRACT………………..……………………………….…..….……277 INTRODUCTION…………………………………………………........279 METHODS………………………………………………………...........281