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Format Not Finished PHYLOGENY OF GEOPHAGINE CICHLIDS FROM SOUTH AMERICA (PERCIFORMES: LABROIDEI) A Dissertation by HERNÁN LÓPEZ FERNÁNDEZ 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 2004 Major Subject: Wildlife and Fisheries Sciences PHYLOGENY OF GEOPHAGINE CICHLIDS FROM SOUTH AMERICA (PERCIFORMES: LABROIDEI) A Dissertation by HERNÁN LÓPEZ FERNÁNDEZ Submitted to Texas A&M University in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY Approved as to style and content by: ___________________________ ___________________________ Kirk O. Winemiller Rodney L. Honeycutt (Co-Chair of Committee) (Co-Chair of Committee) __________________________ ___________________________ Melanie L. J. Stiassny John D. McEachran (Member) (Member) ___________________________ ___________________________ James B. Woolley Lee A. Fitzgerald (Member) (Member) ___________________________ Robert D. Brown (Head of Department) August 2004 Major Subject: Wildlife and Fisheries Sciences iii ABSTRACT Phylogeny of Geophagine Cichlids from South America (Perciformes: Labroidei). (August 2004) Hernán López Fernández, B.S., Universidad de Los Andes, Venezuela Co-Chairs of Advisory Committee: Dr. Kirk O. Winemiller Dr. Rodney L. Honeycutt Three new species of cichlid fishes of the genus Geophagus, part of the Neotropical subfamily Geophaginae, are described from the Orinoco and Casiquiare drainages in Venezuela. Phylogenetic relationships among 16 genera and 30 species of Geophaginae are investigated using 136 morphological characters combined with DNA sequences coding for the mitochondrial gene NADH dehydrogenase subunit 4 (ND4) and the nuclear Recombination Activating Gene 2 (RAG2). Data from previous studies are integrated with the new dataset by incorporating published DNA sequences from the mitochondrial genes cytochrome b and 16S and the microsatellite flanking regions Tmo- M27 and Tmo-4C4. Total-evidence analysis revealed that Geophaginae is monophyletic and includes eighteen genera grouped into two major clades. In the first clade, the tribe Acarichthyini (genera Acarichthys and Guianacara) is sister-group to a clade in which Gymnogeophagus, ‘Geophagus’ steindachneri, and Geophagus sensu stricto are sister to ‘Geophagus’ brasiliensis and Mikrogeophagus; all these are in turn sister-group to Biotodoma, Dicrossus and Crenicara. In the second clade, Satanoperca, Apistogramma (including Apistogrammoides), and Taeniacara are sister to Crenicichla and Biotoecus. Monophyly and significantly short branches at the base of the phylogeny indicate that genera within Geophaginae differentiated rapidly within a relatively short period. High morphological, ecological, and behavioral diversity within the subfamily suggest that geophagine divergence may be the result of adaptive radiation. iv In memory of Mom, to Dad, to Diego In memory of “El Chino” Foong To Don Taphorn v ACKNOWLEDGEMENTS I am indebted to a large number of people and institutions without whom this dissertation would remain just an idea. Above all, Kirk Winemiller, Rodney Honeycutt, and Melanie Stiassny have been involved to such extent that I feel this work is as much theirs as it is mine, although all mistakes and opinions are product of my stubbornness. Kirk started it all when he invited me to be his student some seven years ago, and since then, he has been advisor, professor, colleague, and friend. In Texas, and from Venezuela to West Africa, the times spent with Kirk will remain some of the best and more stimulating of my life. Rodney has been gracious enough to frequently exchange mammals for cichlids with an enthusiasm and passion that turns research into an adventure. Working with him has been one of the most fortunate and inspiring turns of this research. Melanie took this project as her own from the beginning and has made sure that everything I ever needed was there for me, from lessons on anatomy and systematics to lessons on life. It has been an honor to work with all of them and to come to see them as colleagues, friends, and role models. John McEachran, Jim Woolley, and Lee Fitzgerald formed the rest of a superb committee. Thoughts from classes and many conversations with all of them are present everywhere in this dissertation, and have shaped many of my thoughts about fish, systematics, and evolution. Working with a group of fishes that spans an entire continent was possible only through the help of many museums and willing friends and colleagues that loaned specimens and/or collected tissues. For the loan of specimens from their collections, I am grateful to Melanie Stiassny (American Museum of Natural History, New York), Donald Taphorn (Museo de Ciencias Naturales de Guanare, Venezuela), Erica Pellegrini Caramaschi and Paulo Buckup (Museu Nacional do Rio de Janeiro, Brazil), Patrice Pruvost and Yves Fermon (Museum National d’Histoire Naturelle, Paris), Patrick Campbell (The Natural History Museum, London), John McEachran (Texas Cooperative Wildlife Collection, College Station), Frank Pezold (Northeastern Louisiana University, Monroe), and Roberto Reis (Museu de Zoologia do PUCRS, Porto Alegre, Brazil). For vi providing tissue samples for DNA sequencing, I am indebted to Nate Lovejoy and Stuart Willis (University of Manitoba, Canada), Izeni Farias (Universidade do Amazonas, Brazil), Angela Ambrosio and Eloísa Revaldaves (Universidade Estadual de Maringá, Brazil), and Yves Fermon (Museum National d’Histoire Naturelle, Paris). Collections in the field were possible thanks to the help of Kirk Winemiller, Albrey Arrington, Stuart Willis, Craig Layman, Tom Turner, José Vicente Montoya, David Hoeinghaus, Carmen Montaña, Leslie Kelso-Winemiller, Jennifer Arrington, and Hugo López. Fishing permits in Venezuela were granted by the Servicio Autónomo para los Recursos Pesqueros y Acuícolas of the Venezuelan Ministerio del Ambiente. Funding for the field portion of this research came mostly through grants from the National Geographic Society and the National Science Foundation to Kirk Winemiller. DNA sequencing and other laboratory research was funded partially by a Jordan Endowment Fund from the American Cichlid Association, a faculty mini-grant from Texas A&M University to Kirk Winemiller, and grants from the National Science Foundation to Rodney Honeycutt. Research on cichlid morphology was partially possible thanks to a Collection Study Grant from the American Museum of Natural History and two Axelrod Fellowships from the Department of Ichthyology at the American Museum. I have been fortunate to interact with many friends and colleagues that made this experience much more than just graduate school. In the Winemiller lab, years of morning coffee with Albrey Arrington were an intellectual treat and built a strong friendship. Stuart Willis was partner of many an adventure and remains an unconditional friend and colleague. I benefited enormously from the time spent with David Jepsen, José Vicente Montoya, David Hoeinghaus, Craig Layman, Senol Akin, Jenny Birnbaum, and Steve Zeug. In the Honeycutt lab, I could not have done anything if April Harlin and Diane Rowe had not been willing to drop everything to help a clumsy beginner to grasp the basics of PCR and DNA sequencing. Joe Gillespie offered invaluable help with the alignment of mitochondrial ribosomal sequences. Discussions with April Harlin, Anthony Cognato, Diane Rowe, Iván Castro, Stuart Willis, Joe vii Gillespie, Isabel Landim, Brian Langerhans, Cliff Rhuel, Bob Schelly, Izeni Farias, José Vicente Montoya, Thom DeWitt, Larry Frabota, Colleen Ingram, the Systematics Discussion Group, and the Ecology of Adaptive Radiations seminar at Texas A&M have benefited my work and shaped many ideas. Over the years in College Station I met some truly brilliant people and some have become very close friends. April Harlin and Anthony Cognato, Dawn Sherry, Iván Castro, Kirk and Leslie Winemiller, Don, Lyn and Anne Willis, Al and Pat Gillogly, and Duane and Judy Schlitter have treated me like family. For their friendship of years, I am especially grateful to José Vicente Montoya, Alexandre Garcia, Scott Brandes, Dave Jepsen, Tammy McGuire, Rob Powell, Gage Dayton, Rebecca Belcher, Lance and Eve Fontaine, Magui Mieres, Stacey Allison, and Damion Marx. Finally, my family and friends from Venezuela have been always close and supportive during this long process. I am deeply grateful to my father, Hugo, and my brother Diego, who saw the beginning of all this, and have believed in me even when I doubted. Don Taphorn started my formal education as an ichthyologist, and has always made sure that I do not stray from the path. María Alexandra Rujano, Luis Gabaldón, Cecilia Rodríguez, Saberio Pérez, Josmary Muñoz, Tatiana Páez, Alfonso Llobet, Mariana Escovar, Ximena Daza, and Jesús Molinari have managed to be close, despite the long distance. viii TABLE OF CONTENTS Page ABSTRACT .................................................................................................. iii DEDICATION .............................................................................................. iv ACKNOWLEDGEMENTS .......................................................................... v TABLE OF CONTENTS .............................................................................. viii LIST OF FIGURES....................................................................................... x LIST OF TABLES .......................................................................................
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