* The Neotropical Fish Family Ctenoluciidae (Teleostei: Ostariophysi: Characiformes): Supra and Intrafamilial Phylogenetic Relationships, with a Revisionary Study RICHARD P. VARI I SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 564 SERIES PUBLICATIONS OF THE SMITHSONIAN INSTITUTION Emphasis upon publication as a means of "diffusing knowledge" was expressed by the first Secretary of the Smithsonian. In his formal plan for the institution, Joseph Henry outlined a program that included the following statement: "It is proposed to publish a series of reports, giving an account of the new discoveries in science, and of the changes made from year to year in all branches of knowledge." This theme of basic research has been adhered to through the years by thousands of titles issued in series publications under the Smithsonian imprint, commencing with Smithsonian Contributions to Knowledge in 1848 and continuing with the following active series: Smithsonian Contributions to Anthropology Smithsonian Contributions to Botany Smithsonian Contributions to the Earth Sciences Smithsonian Contributions to the Marine Sciences Smithsonian Contributions to Paleobiology Smithsonian Contributions to Zoology Smithsonian Folklife Studies Smithsonian Studies in Air and Space Smithsonian Studies in History and Technology In these series, the Institution publishes small papers and full-scale monographs that report the research and collections of its various museums and bureaux or of professional colleagues in the world of science and scholarship. The publications are distributed by mailing lists to libraries, universities, and similar institutions throughout the world. Papers or monographs submitted for series publication are received by the Smithsonian Institution Press, subject to its own review for format and style, only through departments of the various Smithsonian museums or bureaux, where the manuscripts are given substantive review. Press requirements for manuscript and art preparation are outlined on the inside back cover. I. Michael Heyman Secretary Smithsonian Institution SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 564 The Neotropical Fish Family Ctenoluciidae (Teleostei: Ostariophysi: Characiformes): Supra and Intrafamilial Phylogenetic Relationships, with a Revisionary Study Richard P. Vari SMITHSONIAN INSTITUTION PRESS Washington, D.C. 1995 ABSTRACT Vari, Richard P. The Neotropical Fish Family Ctenoluciidae (Teleostei: Ostariophysi: Characiformes): Supra and Intrafamilial Phylogenetic Relationships, with a Revisionary Study. Smithsonian Contributions to Zoology, number 564, 97 pages, 51 figures, 12 tables, 1995.—Osteological and soft anatomical features of the species of the Neotropical characiform family Ctenoluciidae and other characiforms were studied to examine the hypothesis that the family was monophyletic, and to advance an hypothesis of phylogenetic relationships within the family and of the Ctenoluciidae to proximate outgroups. A series of derived features corroborate the hypothesis of monophyly of the Ctenoluciidae and are congruent with the hypothesis that the Ctenoluciidae and Erythrinidae are sister groups. A less-extensive series of derived features indicate that the African family Hepsetidae is the sister group of the clade formed by the Ctenoluciidae and Erythrinidae, with the Neotropical family Lebiasinidae the sister group to the clade formed by those three families. Shared derived features of a variety of body systems define phylogenetic subunits of the Ctenoluciidae and characterize the majority of its species. The ctenoluciid genera Ctenolucius Gill (1861a) and Boulengerella Eigenmann (1903) are defined as monophyletic units. Luciocharax Steindachner (1878) and Belonocharax Fowler (1907) are considered synonyms of Ctenolucius, with Spixostoma Whitley (1951) placed as a synonym of Boulengerella. Two species are recognized in Ctenolucius. Ctenolucius hujeta (Valenciennes in Cuvier and Valenciennes, 1849) ranges from the Lago Maracaibo basin of northwestern Venezuela through the Rfo Magdalena system to the Rio Sinu of northwestern Colombia. Ctenolucius beani (Fowler, 1907) occurs in the Rio Atrato and Rio San Juan basins of northwestern Colombia and the rivers of the Pacific slopes of Panama as far west as Veraguas Province. Luciocharax insculptus Steindachner (1878) is a synonym of C. hujeta, and Luciocharax striatus Boulenger (1911) is a synonym of C. beani. Five species are recognized in Boulengerella. Boulengerella lateristriga (Boulenger, 1895) occurs in the Rio Negro drainage of the Amazon basin in Brazil and Venezuela and uppermost portions of the Rio Orinoco in southern Venezuela. Boulengerella maculata (Valenciennes in Cuvier and Valenciennes, 1849) is widely distributed through the Rio Amazonas, Rio Tocantins, and Rio Orinoco basins. Boulengerella lucius (Cuvier, 1819) occurs in the Rio Amazonas and Rio Orinoco systems. Boulengerella cuvieri (Agassiz in Spix and Agassiz, 1829) is widespread through the Rio Orinoco, Rio Amazonas, and Rio Tocantins basins, the Essequibo River of Guyana, the Oyapock River along the boundary between French Guiana and Brazil, and the coastal rivers of Amapd and Par£ states in Brazil. Boulengerella xyrekes, new species, a relatively rare form, inhabits the Rio Orinoco and Rio Amazonas basins. Xiphostoma taedo Cope (1872) is a synonym of B. maculata. Xiphostoma oseryi Castelnau (1855) and Xiphostoma ocellatum Schomburgk (1841) are considered synonyms of B. cuvieri. Xiphostoma longipinne Steindachner (1876), based on a juvenile from the mouth of the Rio Negro in Brazil, tentatively is considered a synonym of B. cuvieri. Reports of Boulengerella from the Rio Pamafba of northeastern Brazil and in the Rio de La Plata system in Argentina either are based on specimens with questionable locality data or are probable misidentifications. Keys are provided to distinguish Ctenolucius and Boulengerella and for the species in each genus. Lectotypes are designated for Xiphostoma hujeta Valenciennes, Luciocharax insculptus Steindachner, Xiphostoma ocellatum Valenciennes, and Luciocharax striatus Boulenger. The phylogenetic biogeography of the group indicates a vicariance event between the Hepsetidae and the lineage consisting of the Ctenoluciidae and Erythrinidae associated with, or predating, the final separation of Africa and South America about 85 mya. Fossil and distributional data indicate that major cladogenesis within the Ctenoluciidae and Erythrinidae predates the late Miocene. OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: The coral Montastrea cavernosa (Linnaeus). Library of Congress Cataloging-in-Publication Data Vari, Richard P. The neotropical fish family Ctenoluciidae (Teleostei: Ostariophysi: Characiformes) ; supra and intrafamilial phylogenetic relationships, with a revisionary study / Richard P. Vari. p. cm.—(Smithsonian contributions to zoology ; no. 564) Includes bibliographical references (p. ) and index. 1. Ctenoluciidae—South America—Phylogeny. 2. Ctenoluciidae—South America—Classifica- tion. I. Title. II. Series. 597'.52—dc20 94-36338 © The paper used in this publication meets the minimum requirements of the American National Standard for Permanence of Paper for Printed Library Materials Z39.48 1984. Contents Page Introduction 1 Acknowledgments 2 Methods and Materials 2 Terminology 5 Abbreviations 5 Institutional Abbreviations 5 Text and Figure Abbreviations 5 Character Description and Analysis 6 Antorbital 9 First Infraorbital and Supraorbital 10 Fourth Infraorbital 11 Fifth Infraorbital 11 Sixth Infraorbital 12 Nasal 12 Vomer 13 Lateral Ethmoid 13 Frontal 13 Dilatator Groove 13 Epiphyseal Bar 15 Laterosensory Canals in Frontal and Parietal 16 Pterotic 17 Supraoccipital 17 Suprapreopercle 18 Jaws and Associated Soft Tissues 18 Upper Jaw 18 Lower Jaw 20 Dentition 21 Palatine 22 Ectopterygoid Teeth 23 Mesopterygoid Teeth 24 Metapterygoid-Quadrate Fenestra 24 Basihyal 24 Anterior and Posterior Ceratohyals 24 Branchiostegal Rays 25 Interhyal 26 Gill-Rakers 26 Postcleithra 26 Weberian Apparatus 27 Tripus 27 Fourth Pleural Rib 27 Ligament between Tripus and Fourth Pleural Rib 27 Transverse Process of Third Neural Arch 27 Laterosensory Canal System on Body 28 Scales 28 Squamation at Posterodorsal Margin of Head 29 Anal Fin 30 in iv SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY Sexual Dimorphism 30 Ontogenetic Variation 31 Dorsal-Fin Position 31 Adipose Fin 32 Caudal Fin Bony Stays 32 Myology 32 Dilatator Operculi Muscle 32 Adductor Mandibulae Muscle 34 Levator Arcus Palatini Muscle 34 Pigmentation 34 Body and Head Pigmentation 34 Fin Pigmentation 34 Phylogenetic Reconstruction 35 Monophyly of the Lebiasinidae, Hepsetidae, Erythrinidae, and Ctenoluciidae Clade 35 Comments on Monophyly of Lebiasinidae and Subunits 35 Monophyly of the Hepsetidae, Erythrinidae, and Ctenoluciidae Clade 37 Comments on Monophyly of Hepsetidae 37 Monophyly of the Erythrinidae and Ctenoluciidae Clade 37 Comments on Monophyly of Erythrinidae 38 Monophyly of Ctenoluciidae 38 Monophyly of Ctenolucius 39 Monophyly of Boulengerella 40 Intrageneric Relationships in Boulengerella 40 Family CTENOLUCIIDAE Schultz, 1944 41 Key to Ctenolucius Gill and Boulengerella Eigenmann 42 Genus Ctenolucius Gill, 1861 42 Key to Species of Ctenolucius Gill, 1861 43 Ctenolucius hujeta (Valenciennes in Cuvier and Valenciennes, 1849) 43 Ctenolucius beani (Fowler, 1907) 52 Genus Boulengerella Eigenmann, 1903 57 Key