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Trophic Ecology of Frugivorous Fishes in Floodplain Forests Of TROPHIC ECOLOGY OF FRUGIVOROUS FISHES IN FLOODPLAIN FORESTS OF THE COLOMBIAN AMAZON A Dissertation by SANDRA BIBIANA CORREA VALENCIA Submitted to the Office of Graduate Studies of Texas A&M University in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY August 2012 Major Subject: Wildlife and Fisheries Sciences Trophic Ecology of Frugivorous Fishes in Floodplain Forests of the Colombian Amazon Copyright August 2012 Sandra Bibiana Correa Valencia TROPHIC ECOLOGY OF FRUGIVOROUS FISHES IN FLOODPLAIN FORESTS OF THE COLOMBIAN AMAZON A Dissertation by SANDRA BIBIANA CORREA VALENCIA Submitted to the Office of Graduate Studies of Texas A&M University in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY Approved by: Chair of Committee, Kirk Winemiller Committee Members, Spence Behmer Stephen Davis Derbert Gatlin Thomas Olszewski Head of Department, John Carey (Iterim) August 2012 Major Subject: Wildlife and Fisheries Sciences iii ABSTRACT Trophic Ecology of Frugivorous Fishes in Floodplain Forests of the Colombian Amazon. (August 2012) Sandra Bibiana Correa Valencia, B.S., Universidad del Valle; M.S., University of Florida Chair of Advisory Committee: Dr. Kirk Winemiller Diverse fish species consume fruits and seeds in the Neotropics, in particular in the lowland reaches of large rivers, such as the Amazon, Orinoco, and Paraná in South America. Floodplains of the Amazon River and its lowland tributaries are characterized by marked hydrological seasonality and diverse assemblages of frugivorous fishes, including closely related and morphologically similar species of several characiform families. Here, I investigated whether or not these fishes are capable of detecting fluctuations in food availability and if they are, how they adjust their feeding strategies. I tested predictions of optimal foraging, limiting similarity and resource partitioning theories with regard to expansion or compression of niche breadth and reduction in trophic niche overlap among species in relation with fluctuations in the availability of alternative food resources. I monitored fruiting phenology patterns to assess food availability and conducted intensive fishing during the high-, falling-, and low-water seasons in an oligotrophic river and an adjacent oxbow lake in the Colombian Amazon. I combined analysis of stomach contents and stable isotope ratios to evaluate dietary patterns, niche breadth, and niche overlap. Diets of six characiform fish species (Brycon falcatus, B. melanopterus, Myloplus asterias, M. rubripinnis, and M. torquatus) changed in a manner that indicated responses to iv fluctuations in food availability. Feeding strategies during the peak of the flood pulse were consistent with predictions of optimal foraging theory. During times of high fruit abundance, fish preferentially consumed items to which their phenotype is best adapted, maximizing net energy gain and enhancing fitness. As the flood pulse subsided and the availability of forest food resources was reduced in aquatic habitats, there was not a consistent pattern of diet breadth expansion or compression, even though diet shifts occurred, suggesting interspecific differences in foraging efficiencies. Analyses of diets and isotopic ratios revealed a general pattern of increased dietary segregation as the water level receded. Although there never was complete niche segregation among these fishes, these dietary changes effectively reduced interspecific niche overlap. Implications of these results and contribution of allochthonous food resources to diversity maintenance of floodplain fishes are discussed. v DEDICATION To my parents Rosalba Valencia and Alvaro Correa, and to my sister Dr. Maritza Correa Valencia, whose unconditional support has helped me achieve my dreams, even though they cannot explain why I choose to chase after fish in the depths of the Amazon forest for a career. To Dr. Michael Goulding, whose dedication to the conservation of Amazon flooded forests has been an inspiration for my career choice. vi ACKNOWLEDGEMENTS I especially want to thank my advisor Dr. Kirk Winemiller for giving me the great opportunity to be a part of his Aquatic Ecology Laboratory, supporting me throughout the years, and continuously stressing the importance of interpreting patterns in nature in light of classical and current theories. I am very grateful to the other members of my committee: Drs. Spence Behmer, Stephen Davis, Derbert Gatlin, and Thomas Olszewski; for ideas, suggestions, constructive criticism, and support. I would also like to thank the Evans Library and the Writing Center at Texas A&M University (TAMU) for their services throughout my research and in the completion of my dissertation. I am extremely thankful to volunteer field assistants who helped me sample fishes and plants, and collect data in the Colombian Amazon: Jed Macdonald, Gaby Tamez, Jhon Patarroyo, Lyda Cruz, Ivan Lopez, and Laura Bravo. I would also like to thank the indigenous field assistants: Diomedes Silva, Jeriel Ahue, Ciro, Edgar Tanimuka, Benjamin Mejia, Luis Tanimuka, Alipio Carevilla Tanimuka, Fabio Yucuna, and Angel Yucuna for their help. Without their hard work and commitment, I would not have been able to accomplish all of what is presented in this manuscript. Thanks to the indigenous communities of the Resguardo Ticoya (Puerto Nariňo) and Resguardo Yaigojé-Apaporis (Puerto Ňumi and Bocas de Taraira) for granting access to their territories and welcoming me with open arms. To conduct research in these areas, permits were granted by the Instituto Colombiano de Agricultura (ICA). Many individuals and institutions in Colombia provided crucial logistic support: Santiago Duque (Universidad Nacional de Colombia, Sede Leticia), Fernando Trujillo vii (Fundación Omacha), Sarita Kendall (Fundación Natütama), Erwin Palacios and Francis Palacios (Conservación Internacional Colombia), Pablo Stevenson (Universidad de los Andes), Francisco Villa (Universidad del Tolima), and Claudia Medina (Instituto Humboldt). Many thanks to Dairon Cárdenas and Nicolas Castaño (Herbario Amazónico Colombiano, Instituto Sinchi), and Pablo Stevenson and Ivonne Vargas (Universidad de los Andes) for helping me identify numerous and diverse plant and fruit samples collected during this project. Thanks to Alejandro Torres for art-work design of educational materials. Funding for this dissertation research was provided by the Wildlife Conservation Society’s Research Fellowship Program. Funds from the College of Agriculture and Life Sciences at TAMU (Excellence Assistantship Award) and the International Federation of University Women (Recognition Award) were used to help conduct this research. Idea Wild generously donated equipment during different phases of this work. The Department of Wildlife and Fisheries Sciences at TAMU continuously funded me with teaching assistantships and travel grants throughout my dissertation. The section of Ecology and Evolutionary Biology at TAMU also provided travel grants to participate in conferences and to conduct museum work in Colombia. I am also indebted to the Tom Slick Senior Graduate Fellowship at TAMU for providing funding during the data analysis. I would like to thank former and current students in the Kirk Winemiller Aquatic Ecology Laboratory and in the Department of Wildlife and Fisheries Sciences at TAMU and my Colombian friends (Adriana Mendez, Marta Ariza, David Toledo, and Sofia Agudelo) for their insightful conversations, moral support, and long-lasting friendships. Lastly, I would like to thank John Baker for his support, friendship, love, and survival throughout my dissertation. viii TABLE OF CONTENTS Page ABSTRACT ............................................................................................................... iii DEDICATION ........................................................................................................... v ACKNOWLEDGEMENTS ....................................................................................... vi TABLE OF CONTENTS ........................................................................................... viii LIST OF FIGURES ................................................................................................... x LIST OF TABLES ..................................................................................................... xii CHAPTER I INTRODUCTION: AQUATIC FRUGIVORES, THE CONSUMPTION OF FRUITS AMONG NEOTROPICAL FISHES . 1 II CHANGES IN FEEDING STRATEGIES IN RESPONSE TO FLUCTUATING FOOD RESOURCES AMONG AMAZONIAN FLOODPLAIN-FOREST FISHES ....................................................... 6 Introduction ..................................................................................... 6 Materials and methods .................................................................... 9 Study site .................................................................................... 9 Sample and data collection ........................................................ 13 Data analysis .............................................................................. 19 Results ............................................................................................. 25 Dietary composition from stomach contents analysis ............... 25 Food availability and food consumption.................................... 31 Dietary similarity based on stomach contents ........................... 43 Dietary similarity based on stable isotopes ...............................
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