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A Review of the North American Freshwater Snail Genus Pyrgulopsis (Hydrobiidae)

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A Review of the North American Freshwater Snail Genus Pyrgulopsis (Hydrobiidae) A Review of the North American Freshwater Snail Genus Pyrgulopsis (Hydrobiidae) ROBERT HERSHLER SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 554 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 554 A Review of the North American Freshwater Snail Genus Pyrgulopsis (Hydrobiidae) Robert Hershler SMITHSONIAN INSTITUTION PRESS Washington, D.C. 1994 ABSTRACT Hershler, Robert. A Review of the North American Freshwater Snail Genus Pyrgulopsis (Hydrobiidae). Smithsonian Contributions to Zoology, 554, 115 pages, 53 figures, 2 tables, 1994.—The Recent nominal species belonging to the hydrobiid snail genus Pyrgulopsis Call and Pilsbry, 1886, are reviewed. Pyrgulopsis is a large group of 65 Recent species inhabiting diverse inland waters of North America. This genus is distinguished from other members of the subfamily Nymphophilinae by the combination of small size; generally simple, ovate-conic shell; and penis ornamented with relatively few glands. Members of this genus have a globose to elongate-conic shell with near smooth to strongly punctate protoconch, smooth or rarely carinate teleoconch, and simple apertural lip often loosened from the body whorl. The taenioglossate radula features a central radular tooth with well-developed lateral angles, narrow basal process, and, usually, a single pair of basal cusps. The operculum generally is paucispiral with eccentric nucleus. The ventral operculum is notable for its variably thickened attachment scar margin and central callus. The caecal chamber of the stomach is variably developed. Animal pigmentation is variable, sometimes including dark stripes along the mantle. Cephalic tentacle ciliation generally is weak. The gonads usually are of simple lobes, and sometimes overlap the stomach. The prostate gland usually has a small pallial section and the vas deferens usually exits from the gland sub-terminally. The penis is generally large and distally bifid; with a terminal filament bearing the vas deferens, and accessory lobe (sometimes absent). The penis surface is very weakly ciliated and is variably ornamented with dorsal and ventral glands. The albumen gland usually has a small pallial section. The capsule gland is almost always bipartite, and contains a well-developed ventral channel. The genital aperture is either simple or opens to an anterior vestibule. The coiled oviduct usually is simple, and joins the bursal duct just behind, to well anterior to, the pallial wall. Both bursa copulatrix and seminal receptacle usually are present, the former being larger and often at least partly posterior to the albumen gland. The bursal duct is of variable size and shape, and often embedded, sometimes deeply so, within the albumen gland. The seminal receptacle is usually just behind the coiled oviduct and typically has a short duct. Females are oviparous, and lay single, hemispherical egg capsules. A cladistic analysis of relationships among 60 species was conducted using Nymphophilus Taylor as single outgroup. Monophyly of Pyrgulopsis was supported by 15 character-state transformations, including six non-homoplasious synapomorphies. Four large clades were recognized: a group of nine species distributed in Mississippi River drainage and other waters to the east, a group of six species from southern Nevada and southeastern Arizona; a group of 10 species from eastern California, northern Arizona, and Snake River drainage; and a group of 18 species scattered throughout the West. The remaining 17 western taxa included four species occupying basal positions on the cladogram, a poorly resolved group of eight species, and two small clades consisting of two species from northern Arizona; and three widely disjunct species from northern Mexico, Snake River drainage, and southeastern California. The following genera herein are added to the list of junior subjective synonyms of Pyrgulopsis: Savaginius Taylor, 1966a, Apachecoccus Taylor, 1987, and Yaquicoccus Taylor, 1987. Fluminicola avernalis carinifera Pilsbry, 1935, is elevated to full species status herein —this and fifteen other species are newly allocated to Pyrgulopsis. Fontelicella pinetorum Taylor, 1987, is a junior subjective synonym of Pyrgulopsis kolobensis (Taylor, 1987); and Pyrgulopsis wabashensis Hinkley, 1908a, is a junior subjective synonym of Pyrgulopsis scalariformis (Wolf, 1869). A new name, Pyrgulopsis bryantwalkeri, is proposed for Fluminicola nevadensis Walker, 1916 (not Pyrgula nevadensis Stearns, 1883). OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded in the Institution's annual report, Smithsonian Yera. SERIES COVER DESIGN: The coral Montastrea cavernosa (Linneaus). Library of Congress Cataloging-in-Publication Data Hershler, Robert A review of the North American freshwater snail genus Pyrgulopsis (Hydrobiidae) / Robert Hershler. p. cm.—(Smithsonian contributions to zoology ; no. 554) Includes bibliographical references. 1. Pyrgulopsis—North America—Classification. I. Title. II. Series. QL1.S54 no. 554 [QL430.5.H9] 591 s-dc20 [594'.3] 93-47246 ® 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 Materials and Methods 2 Acknowledgments 4 Superfamily RISSOOIDEA Gray, 1847 5 Family HYDROBIIDAE Troschel, 1857 5 Subfamily NYMPHOPHILINAE Taylor, 1966b . 5 Genus Pyrgulopsis Call and Pilsbry, 1886 5 Western American Species 14 Pyrgulopsis aardahlii Hershler, 1989 14 Pyrgulopsis amargosae Hershler, 1989 17 Pyrgulopsis archimedis Berry, 1947 17 Pyrgulopsis arizonae (Taylor, 1987) 18 Pyrgulopsis avernalis (Pilsbry, 1935), new combination 19 Pyrgulopsis bacchus Hershler, 1988 21 Pyrgulopsis bernardina (Taylor, 1987) 21 Pyrgulopsis brandi (Drake, 1953), new combination 22 Pyrgulopsis bruneauensis Hershler, 1990 23 Pyrgulopsis bryantwalkeri, new name 23 Pyrgulopsis californiensis (Gregg and Taylor, 1965) 25 Pyrgulopsis carinifera (Pilsbry, 1935a), new combination 26 Pyrgulopsis cedrosensis (Pilsbry, 1927), new combination 27 Pyrgulopsis chihuahua (Pilsbry, 1928), new combination 28 Pyrgulopsis chupaderae (Taylor, 1987), new combination 30 Pyrgulopsis conica Hershler, 1988 30 Pyrgulopsis cry stalls Hershler and Sada, 1987 32 Pyrgulopsis davisi (Taylor, 1987), new combination 32 Pyrgulopsis deserta (Pilsbry, 1916) 33 Pyrgulopsis erythropoma (Pilsbry, 1899) 35 Pyrgulopsis fairbanksensis Hershler and Sada, 1987 36 Pyrgulopsis gilae (Taylor, 1987), new combination 36 Pyrgulopsis giulianii Hershler and Pratt, 1990 38 Pyrgulopsis glandulosa Hershler, 1988 39 Pyrgulopsis hendersoni (Pilsbry, 1933) 39 Pyrgulopsis idahoensis (Pilsbry, 1933) 41 Pyrgulopsis intermedia (Tryon, 1865) 42 Pyrgulopsis isolata Hershler and Sada, 1987 44 Pyrgulopsis kolobensis (Taylor, 1987), new combination 44 Pyrgulopsis longinqua (Gould, 1855) 46 Pyrgulopsis manantiali (Hershler, 1985) 47 Pyrgulopsis merriami (Pilsbry and Beecher, 1892), new combination 49 Pyrgulopsis metcalfi (Taylor, 1987), new combination 49 Pyrgulopsis micrococcus (Pilsbry, 1893) 50 Pyrgulopsis montezumensis Hershler, 1988 52 Pyrgulopsis morrisoni Hershler, 1988 52 Pyrgulopsis nanus Hershler and Sada, 1987 53 Pyrgulopsis neomexicana (Pilsbry, 1916) 54 in iv SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY Pyrgulopsis nevadensis (Steam, 1883) 55 Pyrgulopsis owensensis Hershler, 1989 57 Pyrgulopsis palomasensis (Pilsbry, 1895), new combination 57 Pyrgulopsis pecosensis (Taylor, 1987), new combination 59 Pyrgulopsis perturbata Hershler, 1989 59 Pyrgulopsis pilsbryana (Baily and Baily, 1952) 60 Pyrgulopsis pisteri Hershler and Sada, 1987 60 Pyrgulopsis robusta (Walker,
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