LINKING COMPOSITION, STRUCTURE AND FUNCTIONS OF BIODIVERSITY: RELATIONSHIPS AMONG EPIPHYTES, INVERTEBRATES AND BIRDS IN THE CANOPY OF CHILEAN TEMPERATE RAINFORESTS By IVAN ANDRES DIAZ 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 2009 1 © 2009 Ivan Andres Diaz 2 To my parents, Nora Romero and Fanor Díaz To the nature, which always shares with me its secrets To the canopy, the place where trees let you see the forest 3 ACKNOWLEDGMENTS I am deeply thankful to my friend and advisor, Dr. Kathryn E. Sieving for her permanent support, lessons, friendship, and good will. She has always extended her supportive hand, with a warm smile and the right word when it was needed. Katie has been the kind of scientist and advisor I would like to be, and she has opened to me a world of possibilities because of her generosity and dedication. She honors the profession of being professor. I am also deeply thankful to Dr. Juan J. Armesto, who is my good friend and advisor, opening to me the door of science, giving me the shoes to walk into the forest and unlocking the gate of frontiers to let me go to the world beyond. Juan has been a very important person in my career, giving me the opportunity and the support to continue in the scientific way, an opportunity that finally allowed me to study in this country and in this university. I thank my Advisory Committee, Drs. Jack Putz, Doug Levey, Emilio Bruna, Meg Lowman and Graeme Cumming, for their permanent support, lessons, encouragement, and the willing to support and advise my work at any time. I want to express all my gratitude to Ms. Katia Velazquez and Mr. Javier de la Calle for their generosity with housing and for allowing me to use their forest, which they love. I am also very thankful to the Corporación Nacional Forestal CONAF (Chile), which let me use facilities in Chiloé National Park and supported my work there. I also want express my gratitude to the Institute of Ecology and Biodiversity for providing lab facilities, and to Departamento de Ecología y Evolución, Universidad Austral de Chile for providing access and facilities at the Fundo San Martín site. Also, I would like to acknowledge many friends at this time, including those who helped me from the early stages of my career. Pablo Espejo taught me about science and reptiles, Juan Carlos Torres shared with me the world of birds; Dina Robles supported me as a young, too young kid in science. Mary Willson was our library of knowledge, offering advice and clarity. 4 Sharon Reid, Olga Barbosa, Claudia Papic, Mauricio Soto, Eduardo Soto, Luis Cavieres, Roberto Nespolo, Cintia Cornelius and all members of Senda Darwin Foundation, have been key in the development of my career. I am very thankful for the friends who worked with me in the field: Maurice Peña, who became a right hand in the field, Camila Tejo, Natalia Carrasco, María Paz Peña, Daniela Manuschevich, Pablo Necochea, Melanie, Tio Franz, Juan Larraín, Fernanda Salinas, Juan Luis Celis and Anita Venegas. I am very thankful to Chiloé National Park rangers Americo Panichini, Francisco Delgado, Maestro Lolo, don Paco, and in particular to Don Mario Huinao who always helped us with the very best of his good will. I am also very thankful to Mariano Millacura for all his support in the field. I am grateful to the Institute of Tropical Studies and Conservation, to Peter Lahanas and to Joe Maer, who showed and taught me the secrets of the tropical forest, and how to know the world of the canopy. I want also to express my deepest gratitude to Gary Machlis and the Canon National Parks Science Scholar Program, which funded me for three years, to the Fulbright program, to Fondecyt Grant 1050225 for funding support, and to the Department of Wildlife Ecology and Conservation, for being a great place to learn about science and life. To my big friends of the Mafia Latina, all Maestros: Santiago Espinoza, Alejandro Paredes, Rafael Reyna, Luis Ramos, Sonia Canavelli, Marcela Machicote, Sheda Morshed, to the friends who organized our little world in the office in the basement Olga Montenegro, Alejandro Paredes, to my old roommates Mutsuo Nakamura and Osvaldo Jordan, to Michaela Speirs. Finally, I want to express my appreciation to my family who has always supported me, in the good and the hard times; and to Wara Marcelo, who gave me the comprehension I wanted, the support I needed, the smile I liked, and the love I loved. 5 TABLE OF CONTENTS page ACKNOWLEDGMENTS ...............................................................................................................4 LIST OF TABLES...........................................................................................................................9 LIST OF FIGURES .......................................................................................................................11 ABSTRACT...................................................................................................................................13 CHAPTER 1 LINKAGES AMONG STRUCTURE, COMPOSITION AND FUNCTIONS OF BIODIVERSITY IN THE FOREST CANOPY.....................................................................15 Theoretical Background..........................................................................................................15 Composition, Structure and Functions in Forest Canopies ....................................................16 The Forest Canopy: An Ecological Frontier ...................................................................16 Temperate Rainforest of Southern Chile.........................................................................17 Links Among Epiphytes, Invertebrates and Birds in the Rain Forest Canopy................19 2 EPIPHYTE SPECIES RICHNESS AND BIOMASS LOADS ON Eucryphia cordifolia (CUNONIACEAE) AND Aextoxicon punctatum (AEXTOXICACEAE), TWO CANOPY EMERGENT TREES IN CHILEAN TEMPERATE RAIN FORESTS...............23 Introduction.............................................................................................................................23 Methods ..................................................................................................................................26 Study Area.......................................................................................................................26 Study Design ...................................................................................................................27 Tree Structure ..................................................................................................................28 Epiphyte Species Richness..............................................................................................28 Epiphyte Biomass............................................................................................................28 Ecological Functions Conducted by Epiphytes...............................................................30 Results.....................................................................................................................................31 Structural Features of Eucryphia cordifolia and Aextoxicon punctatum Trees...............31 Species Richness and Distribution of Epiphytes on E. cordifolia and A. punctatum......32 Epiphytic Biomass...........................................................................................................33 Epiphyte Functions..........................................................................................................35 Discussion...............................................................................................................................36 Comparative Diversity of Epiphytes in Southern Chile Temperate Rainforests.............36 Comparative Diversity of Epiphytes with Other Tropical and Temperate Rainforests ...................................................................................................................38 Floristic Relationships between Chilean Epiphytes and those from the Neotropics and Australia-New Zealand .........................................................................................39 Epiphyte Vertical Distributions.......................................................................................41 6 Epiphyte Biomass............................................................................................................43 Epiphyte Functions..........................................................................................................45 Importance of Large Trees with Epiphytes .....................................................................48 3 EFFECTS OF EPIPHYTE LOADS ON INVERTEBRATE SPECIES RICHNESS AND ABUNDANCE ON THE CANOPY OF Eucryphia cordifolia (CUNONIACEAE), AN EMERGENT TREE IN CHILEAN TEMPERATE RAIN FORESTS ..................................65 Introduction.............................................................................................................................65 Methods ..................................................................................................................................67 Study Area.......................................................................................................................67 Study Design ...................................................................................................................68 Tree Characterization ......................................................................................................68