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Xenurobryconin Phylogeny and Putative Pheromone Pumps in Glandulocaudine Fishes (Teleostei: Characidae)

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Xenurobryconin Phylogeny and Putative Pheromone Pumps in Glandulocaudine Fishes (Teleostei: Characidae) Xenurobryconin Phylogeny and Putative Pheromone Pumps in Glandulocaudine Fishes (Teleostei: Characidae) STANLEY H. WEITZMAN and SARA V. FINK m SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 421 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 421 Xenurobryconin Phylogeny and Putative Pheromone Pumps in Glandulocaudine Fishes (Teleostei: Characidae) Stanley H. Weitzman and Sara V. Fink SMITHSONIAN INSTITUTION PRESS City of Washington 1985 ABSTRACT Weitzman, Stanley H. and Sara V. Fink. Xenurobryconin Phytogeny and Putative Pheromone Pumps in Glandulocaudine Fishes (Teleostei: Characidae). Smithsonian Contributions to Zoology, number 421, 121 pages, 81 figures, 4 tables, 1985.—A phylogenetic analysis of the glandulo- caudine genera Argopleura, Iotabrycon, Scopaeocharax (new), Tyttocharax, and Xenurobrycon indicates that they form a monophyletic group, the Xenurobryconini. Their relationships to other glandulocaudine characids and the monophyly of the Glandulocaudinae remain uncertain. Hypotheses concerning the evolution and relationships of the glandulocaudine characids are reviewed and evaluated. The phylogenetic analysis of the Xenurobryconini is based on innovative, mostly sexually dimorphic characteris in males and on reductive features of both sexes. The large number of sexually dimorphic apomorphies we list for some of the taxa is a measure of the complexity of the sexual morphology in xenurobryconins. The reductive characters are mostly reduction and/or loss of parts of the laterosensory canal system and some of the associated bones of the head and shoulder girdle. These reductions are similar to those recorded for other groups of small to minute characids but are independently derived. Usually small adult body length, reductive features, and innovative sexual characters are congruent in xenurobryconins. The described laterosensory canal reductions and certain innovative aspects of the caudal pump mechanisms may be correlated with increased resistance of water to flow encountered in these structures in small fishes of 12 to 25 millimeters in adult standard length. Our phylogenetic hypothesis indicates that reversal of laterosensory reductions to a more primitive state may occur as a correlate of reversal to a somewhat larger body size. The phylogenetic analysis of the xenurobryconin genera indicates that Argopleura, whose member species attain the greatest body length of all species in the Xenurobryconini, is the most primitive morphologically and the sister group of all other xenurobryconins; that Iotabry- con is the sister taxon of a lineage comprising Xenurobrycon, Scopaeocharax, and Tyttocharax; and that Xenurobrycon forms a sister group to Scopaeocharax and Tyttocharax. The distributions of the genera are summarized and a three-area cladogram is presented. Xenurobrycon was found to consist of three species: X. marcopus from several localities in the Rio Paraguay-Rio Parana basin of Paraguay and Brazil; X. pteropus, a new species known only from Fonte Boa on the middle Rio Solimoes, Brazil; and X. heterodon, a new species known from the Rio Pachitea of eastern Peru, and from the Rio Aguarico (Rio Napo drainage) and upper Rio Pastaza, both of eastern Ecuador. These latter rivers are Andean tributaries of the western Amazon system. The phylogenetic relationships among the species of Xenurobrycon were not studied. The anatomy of the putative pheromone pumping mechanisms in the caudal region of glandulocaudines is described and discussed. Two widely different pumping mechanisms were found, a diaphragm pump and a fin-ray pump. The first of these has at least two functionally and anatomically different forms, including a passive pump apparently operated by tail-beating during courtship display and a directly driven muscular diaphragm pump found in one subgroup of miniature xenurobryconins. The hydrodynamics of the pump systems are briefly discussed in relation to their morphology, size differences, and the resistance of water to flow in small tube-like structures. 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 Weitzman, Stanley H. Xenurobryconin phytogeny and putative pheromone pumps in glandulocaudine fishes (Teleos- tei : Characidae). (Smithsonian contributions to zoology ; no. 421) Bibliography: p. Supt. of Docs. no. SI 1.27:421 1. Characidae—Evolution. 2. Characidae—Classification. 3. Characidae—Anatomy. 4. Pheromones. 5. Fishes—Evolution. 6. Fishes—Classification. 7. Fishes—Anatomy. L Fink, Sara. V. II. Title. III. Title: Putative pheromone pumps. IV. Title: Glandulocau- dine fishes. V. Series. QL1.S54 no. 421 591 s [597'.52] 85-600079 [QL638.C5] Contents Page Introduction 1 Glandulocaudine Nomenclature and History 1 Xenurobryconin Nomenclature and History 3 Pheromone Caudal Pump Hypothesis 7 Methods 8 Acknowledgments 11 Phylogenetic Hypotheses and Classification of Xenurobryconini .... 12 XENUROBRYCONINI Myers and Bohlke 13 Genus Argopleura Eigenmann, 1913 26 Subgroup A: lotabrycon, Xenurobrycon, Scopaeocharax, and Tyttocharax 30 Genus lotabrycon Roberts, 1973 42 Subgroup B: Xenurobrycon, Scopaeocharax, and Tyttocharax 44 Genus Xenurobrycon Myers and P. de Miranda-Ribeiro, 1945 ... 52 Subgroup C: Scopaeocharax and Tyttocharax 55 Genus Scopaeocharax, new genus 56 Genus Tyttocharax Fowler, 1913 57 Discussion 61 Xenurobryconin Distribution 70 Key to Generic Groups and Genera 72 The Species of Xenurobrycon 74 Artificial Key to the Species of Xenurobrycon 74 Xenurobrycon macropus Myers and P. de Miranda-Ribeiro 74 Xenurobrycon pteropus, new species 85 Xenurobrycon heterodon, new species 88 Putative Pheromone Pumping Mechanisms in Glandulocaudine Fishes 94 Historical Summary 94 Caudal Pump Morphologies and Putative Function 96 The Diaphragm Scale Pump 96 The Fin-Ray Pump 98 Summary 99 Resumo 100 Appendix 1: Glandulocaudine Specimens Examined 102 Appendix 2: Identification, Taxonomic, and Nomenclatural Notes on the Specimens Examined 109 Appendix 3: A Critique of the Monophyly of the Glandulocaudinae, by Stanley H. Weitzman, Sara V. Fink, and Naercio A. Menezes . 112 Addendum 117 Literature Cited 118 iii Xenurobryconin Phylogeny and Putative Pheromone Pumps in Glandulocaudine Fishes (Teleostei: Characidae) Stanley H. Weitzman and Sara V. Fink Introduction GLANDULOCAUDINE NOMENCLATURE AND hypertrophied caudal tissue is simply assumed to HISTORY be glandular. The presence of glandular tissue has been documented by Atkins and Fink (1979) The Glanduocaudinae is a group of American for Corynopoma riisei Gill (1858:426 [66 of re- characid fish genera first recognized by Eigen- print]). mann (1914:34). Subsequently additional genera The subfamily currently consists of 17 genera were added, subtracted, or discussed by Myers and somewhat more than 40 species. The ge- (in Eigenmann and Myers, 1929:467), Tortonese nera that have been assigned to the Glandulo- (1941-1942:77), Myers and P. de Miranda-Ri- caudinae are: Corynopoma Gill (1858:425[65 of beiro (1945:6), Bohlke (1954:266, 1958a:42), reprint]), Pseudocorynopoma Perugia (1891:646), Gery (1963:11), Nelson (1964a: 126), Gery Diapoma Cope (1894:67), Mimagoniates Regan (1966:233), Roberts (1973:489), Fowler (1907:402), Glandulocauda Eigenmann (1911b:- (1975b:332), Gery (1977:351), and Mahnert and 168), Hysteronotus Eigenmann (1911b:- Gery (1984:510). 171), Gephyrocharax Eigenmann (1912:- Glandulocaudines are usually considered as 23), Pterobrycon Eigenmann (1913:3), Argopleura tetragonopterine-related
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