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Systematics of the Neotropical Characiform Genus Steindachnerina Fowler (Pisces: Ostariophysi)

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Systematics of the Neotropical Characiform Genus Steindachnerina Fowler (Pisces: Ostariophysi) * Systematics of the Neotropical Characiform Genus Steindachnerina Fowler (Pisces: Ostariophysi) RICHARD P. VARI I SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 507 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 Astrophysics 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. Robert McC. Adams Secretary Smithsonian Institution SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 507 Systematics of the Neotropical Characiform Genus Steindachnerina Fowler (Pisces: Ostariophysi) Richard P. Vari SMITHSONIAN INSTITUTION PRESS Washington, D.C. 1991 ABSTRACT Van, Richard P. Systematics of the Neotropical Characiform Genus Steindachnerina Fowler (Pisces: Ostariophysi). Smithsonian Contributions to Zoology, number 507, 118 pages, 80 figures, 10 tables, 1991.—The genus Steindachnerina Fowler (1906) of the characiform family Curimatidae is defined as a monophyletic unit. Shared derived features of the gill arches and possibly in dorsal-fin pigmentation diagnose the genus. Derived features of the gill arches, buccopharyngeal complex, infraorbital series, palatine arch, and neurocranium, form of the prepelvic region of the body, details of pigmentation, and meristic features define monophyletic subunits of Steindachnerina or are autapomorphic for species. Steindachnerina has three junior synonyms Curimatorbis, Cruxentina, and Rivasella, all described by Fernandez-Yepez (1948). Twenty-one species are recognized in Steindachnerina: S. amazonica (Steindachner, 1911) of the Rio Tocantins; S. argentea (Gill, 1858) of Trinidad, the Rio Orinoco, and coastal rivers of northern Venezuela; S. atratoensis (Eigenmann, 1912b) of the Rio Atrato; S. bimaculata (Steindachner, 1876) of the Rio Amazonas and Rfo Orinoco; S. binotata (Pearson, 1924) of the Rio Madeira; S. biornata (Braga and Azpelicueta, 1987) of the Rio de La Plata basin and the coastal rivers of Uruguay and southern Brazil; S. brevipinna (Eigenmann and Eigenmann, 1889) of the Rfo Uruguay, Rfo Paraguay, and lower Rfo Parana", and possibly the upper Rio Xingu; S. conspersa (Holmberg, 1891) of the Rfo Paraguay and lower Rfo Parang S. dobula (Gunther, 1868a) of the western Amazon basin; 5. elegans (Steindachner, 1874) of the coastal rivers of eastern Brazil; S.fasciata (Vari and Gery, 1985) of the Rio Madeira basin; 5. gracilis Vari and Vari (1989) of the Rio Tocantins; S. guentheri (Eigenmann and Eigenmann, 1889) distributed from Guyana through the Rfo Orinoco basin, and the Andean peidmont to northern Bolivia; S. hypostoma (Boulenger, 1887a) of the Amazon basin; S. insculpta (Femindez-Yepez, 1948) of the upper Rfo Parana^ S. leucisca (Gunther, 1868) of the Amazon basin; 5. notonota (Miranda-Ribeiro, 1937) of northeastern Brazil; S. planiventris Vari and Vari (1989) of the Amazon basin; S. pupula, new species, endemic to the Rfo Orinoco; S. quasimodoi Vari and Vari (1989) from the Rfo Yavari (Rio Javarf); and S. tuna, new species, from Surinam, French Guiana, and Brazil. A key to the species of Steindachnerina is provided. Nineteen species and subspecies are placed into synonymy in this study. Curimatus leuciscus bolivae described by Eigenmann and Ogle (1908) and Allenina pectinata Fernandez-Yepez (1948) are placed into the synonymy of Steindachnerina leucisca. Curimatus trachystetus Cope (1878), C. bimaculatus sialis Eigenmann and Eigenmann (1889), Prochilodus pterostigma Fowler (1913), Curimatus semiornatus Steindachner (1914), and Curimata melaniris Fowler (1940) are all placed into the synonymy of Steindachnerina bimaculata. Curimata stigmosa Vari (1987) is a synonym of Curimata biornata. Curimatus nitens Holmberg (1891) and Curimatus nigrotaenia Boulenger (1902) are placed into the synonymy of Steindachnerina brevipinna. Curimatus nasus Steindachner (1882), Curimata hypostoma hastata Allen (in Eigenmann and Allen, 1942), Curimata niceforoi Fowler (1943a), and Prochilodus stigmaturus (Fowler, 1911), are all considered synonyms of Steindachnerina dobula. Steindachnerina guentheri has four junior synonyms: Curimatus morawhannae Eigenmann (1912a), Curimatus issororoensis Eigenmann (1912a), Curimatus metae Eigenmann (1922), and Curimata robustula Allen (in Eigenmann and Allen, 1942). Curimatus elegans bahiensis Eigenmann and Eigenmann (1889) is placed into the synonymy of Steindachnerina elegans. Thephylogenetic biogeography of Steindachnerina indicates numerous cases of large-scale, secondary dispersal during the evolution of the genus. The basal speciation events within Steindachnerina predate the final uplift of the Andes. A major part of the speciation within the genus predates the formation of the Amazon basin and apparently occurred outside of the Rio Amazonas system or in peripheral portions of the basin. The previous occurrence of a Steindachnerina species within the Rfo Magdalena system of Colombia and its subsequent extinction is hypothesized based on phylogenetic and distributional data. 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 Calaloging-in Publication Data Vari, Richard P. Systematics of the neotropical characiform genus Steindachnerina Fowler (Pisces: Ostariophysi) / Richard P. Vari. p. cm. — (Smithsonian contributions to zoology ; no. 507) Includes bibliographical references and index. 1. Steindachncrina—Classification. I. Title. II. Series, QL1.S54 no. 507 [OL638.C891 591s—dc20 [597.52] 90-26412 Contents Page Introduction 1 Systematic Procedures 2 Methods and Materials 3 Osteological Material Examined 3 Abbreviations 4 Institutional Abbreviations 4 Text and Figure Abbreviations 4 Acknowledgments 5 Character Description and Analysis 5 Buccopharyngeal Complex 6 Roof of the Oral Cavity 6 Posterior Lobulate Processes 8 Anterior Posteromedian Flap 8 Lobulate Processes of First and Second Gill Arches 8 Branchial Arches 9 Dorsal Portions of Branchial Arches 9 First Infrapharyngobranchial 9 First Epibranchial 10 Second Infrapharyngobranchial 10 Fifth Upper Pharyngeal Tooth-plate 10 Ventral Portions of Branchial Arches 11 Third Hypobranchial 11 Basihyal and Basihyal Tooth-plate 12 Palatine Arch 13 Mesopterygoid 13 Metapterygoid 15 Infraorbitals 16 Third and Fourth Infraorbitals 16 Fifth Infraorbital 17 Cranial Fontanels 17 Pigmentation 18 Position of Anus 20 Vertebrae 21 Squamation 22 Body Form 23 Synapomorphy List and Phylogenetic Reconstruction 23 Monophyly of Steindachnerina 23 Intrageneric Relationships within Steindachnerina 24 Convergent Characters 26 Genus Steindachnerina Fowler, 1906 27 Key to the Species of Steindachnerina Fowler 29 Steindachnerina leucisca (Gunther, 1868) 32 Steindachnerina binotata (Pearson, 1924) 36 Steindachnerina argentea (Gill, 1858) 37 Steindachnerina conspersa (Holmberg, 1891) 42 Steindachnerina bimaculata (Steindachner, 1876) 46 Steindachnerina biornata (Braga and Azpelicueta, 1987) 51 Steindachnerina hypostoma (Boulenger, 1887) 53 Steindachnerina gracilis Vari and Vari, 1989 55 Steindachnerina quasimodoi Vari and Vari, 1989 57 Steindachnerina planiventris Vari and Vari, 1989 59 Steindachnerina atratoensis (Eigenmann, 1912) 61 Steindachnerina fasciata (Vari and Geiy, 1985) 63 Steindachnerina dobula (Gunther, 1868) 66 Steindachnerina insculpta (Fernandez-Yepez, 1948) 71 Steindachnerina runa, new species 75 Steindachnerina pupula, new species 77 Steindachnerina amazonica (Steindachner, 1911) 80 Steindachnerina guentheri (Eigenmann and Eigenmann, 1889) 83 Steindachnerina notonota (Miranda-Ribeiro, 1937) 89 Steindachnerina elegans (Steindachner, 1874) 92 Steindachnerina brevipinna (Eigenmann and Eigenmann, 1889) 97 Phylogenetic Biogeography 102 Appendix 107 Character Matrix for Steindachnerina 107 Character Summary 108 Resumo 109 Literature Cited Ill Index 116
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