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Phylogenetic Relationships of the Earliest Anisostrophically Coiled Gastropods SERIES PUBLICATIONS of the SMITHSONIAN INSTITUTION

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Phylogenetic Relationships of the Earliest Anisostrophically Coiled Gastropods SERIES PUBLICATIONS of the SMITHSONIAN INSTITUTION HKT *£ir «cT 4»BBb %* Alft Phylogenetic Relationships of the Earliest Anisostrophically Coiled Gastropods 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. Lawrence M. Small Secretary Smithsonian Institution SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY • NUMBER 88 Phylogenetic Relationships of the Earliest Anisostrophically Coiled Gastropods Peter J. Wagner ISSUED JAN 3 0 2002 aHHSONIAN INSTITUTION Smithsonian Institution Press Washington, D.C. 2002 ABSTRACT Wagner, Peter J. Phylogenetic Relationships of the Earliest Anisostrophically Coiled Gastro­ pods. Smithsonian Contributions to Paleobiology, number 88, 152 pages, 37 figures, 3 tables, 2002.—In order to explore the phylogenetic relationships among early gastropods, cladistic analyses were conducted of nearly 300 "archaeogastropod" species known from the latest Cambrian through the Silurian. The study includes an extended outgroup analysis of Cambrian molluscs. The resulting estimates of gastropod phylogeny differ not only from traditional ideas about early gastropod relationships, but also from most alternative notions. Outgroup analyses suggest that gastropods had ancestors among the Tergomya (= Monoplacophora of many work­ ers) of the Middle or Late Cambrian. Putative gastropods from older strata (e.g., the Pelagiel- lida and early Onychochilidae) apparently are not closely related to gastropods. The hypothesized ancestor of gastropods possessed dextral-coiling, septation, a deep sinus, and a peripheral band. An anal slit is commonly described as a synapomorphy of gastropods that many clades subsequently lost; however, this study suggests that the slit is a rare, highly derived, and polyphyletic character among early Paleozoic species, and that the ancestors of most "advanced" clades (e.g., the Apogastropoda) never had slits. This study suggests that two major subclades evolved by the earliest Ordovician. The diag­ noses and definitions of these two subclades best correspond to the traditional diagnoses and definitions of the Euomphalina and Murchisoniina. The Pleurotomarioidea is not a paraphyletic ancestral taxon as typically suggested, but instead it is a polyphyletic assemblage derived multi­ ple times from "euomphalinae" and "murchisoniinae" species. The Bellerophontina is at least diphyletic, as the taxon includes both the ancestors of "archaeogastropods" and a clade of planispiral species that is secondarily derived from "archaeogastropods." Macluritoids sensu stricto represent a restricted subclade of the "euomphalinae"; other supposed macluritoids evolved among different euomphalinae subclades or are not gastropods. Early Paleozoic species previously classified as caenogastropods (i.e., the Loxonematoidea and Subulitoidea) represent separate murchisoniinae subclades, with some putative members of the Subulitoidea derived within the Loxonematoidea. Early Paleozoic species assigned to the Trochoidea also represent several subclades, with most of those clades having evolved from the "euomphalinae." An extensive taxonomic revision is presented, which removes all early Paleozoic taxa from the Pleurotomariina and broadly expands the definitions of the Euomphalina and Murchisoniina. OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded in the Institution's annual report, Annals of the Smithsonian Institution. SERIES COVER DESIGN: The trilobite Phacops rana Green. Library of Congress Cataloging-in-Publication Data Wagner, Peter J. Phylogenetic relationships of the earliest anisostrophically coiled gastropods / Peter J. Wagner. p. cm. — (Smithsonian contributions to paleobiology ; no. 88) Includes bibliographic references. 1. Gastropoda, Fossil. I. Title. II. Series. QE808.W34 1999 560s-dc21 [564\scl36\.3] 98-52464 © 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 1 "Archaeogastropods"—A Temporary Definition 2 Review of Previous Phylogenetic Hypotheses 2 Material 4 Specimens 4 Biogeography of Analyzed Species 4 Cladistic Characters 5 Homology Versus Architecture 5 The Paucity of Shell Characters Revisited 5 Sinuses, Slits, Selenizones, and Peripheral Bands 9 Shell Mineralogy and Protoconchs 11 Phylogenetic Analysis 11 Character Deweighting 1—Balancing Continuous Characters 11 Character Deweighting 2—Asymmetry and Changing Homologies 12 Character Analyses 13 Cambrian Molluscs and the Choice of an Outgroup 15 Results 16 Cladogram Statistics and Descriptions 16 Outgroup Analyses 17 Relationships among Early Ordovician Species 21 I. "Euomphalinaes" 21 1.1. "Ophiletoids" 21 1.2. "Macluritoids'' 23 1.3. "Ceratopeoids" 26 1.3.1. "Raphistomatids" 26 1.3.1.1. "Lesueurillines" 26 1.3.1.2. "Holopeines" 29 1.3.2. "Helicotomids" 29 1.3.2.1. "Ophiletinines" 31 1.3.2.2. "Euomphalopterines" 31 1.3.2.2.1. "Anomphalides" 31 1.3.2.2.2. "Poleumitides" 31 1.3.2.2.3. "Pseudophorides" 33 II. "Murchisoniinaes" 35 ILL "Plethospiroids" 39 11.2. "Straparollinoids" 39 11.3. "Hormotomoids" 39 11.3.1. "Subulitids" 39 11.3.2. "Cyrtostrophids" 43 11.3.2.1. "Goniostrophines" 46 11.3.2.2. "Omospirines" 46 11.4. "Eotomarioids" 50 11.4.1. "Lophospirids" 50 11.4.2. "Clathrospirids" 52 11.4.2.1. "Liospirines" 52 11.4.2.2. "Brachytomariines" 55 in IV SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY 11.4.2.2.1. "?Palaeoschismides" 55 11.4.2.2.2. "Phanerotrematides" 57 11.4.2.2.3. "Luciellides" 58 11.4.2.2.4. "Planozonides" 58 Problematica 61 Platyceratoidea 61 Neritoidea 63 Oriostomatoidea 63 Discussion 65 Paleozoic Pleurotomarioids: An Oxymoron? 65 "Euomphalinaes": Two Gills or One? 65 Implications for Relationships among Extant Gastropods 66 Systematic Paleontology 68 Class GASTROPODA Cuvier, 1797 69 Order "ARCHAEOGASTROPODA" Thiele, 1925 69 Family tSlNUOPElDAE Wenz, 1938 69 Genus ^Sinuopea Ulrich, 1911 69 Genus ^Schizopea Butts, 1926 70 Genus Euconia Ulrich in Ulrich and Scofield, 1897 70 Genus Gasconadia Ulrich in Weller and St. Clair, 1928 70 ?Genus Calaurops Whitfield, 1886 70 Suborder EUOMPHALINA de Koninck, 1881 70 Superfamily tOPHiLETOiDEA trans, nov. Knight, 1956 70 Genus ]Ophileta Vanuxem, 1842 71 Genus \Lecanospira Butts, 1926 71 Genus Ecculiomphalus Portlock, 1843 71 Genus ^Asgardaspira, new genus 71 Genus Lytospira Koken, 1896 71 Superfamily MACLURITOIDEA Fischer, 1885 71 Genus \Macluritella Kirk, 1927 72 Genus ^Teiichispira Yochelson and Jones, 1968 72 Genus \Maclurites Le Sueur, 1818 72 Genus Maclurina Ulrich and Scofield, 1897 72 Genus Palliseria Wilson, 1924 72 ?Genus Rousseauspira Rohr and Potter, 1987 73 Superfamily fEuOMPHALOiDEA de Koninck, 1844 73 Family fRAPHlSTOMATlDAE Koken, 1896 73 Genus \Ceratopea Ulrich, 1911 73 Genus Bridgeites Flower, 1968a 73 Genus Orospira Butts, 1926 73 Genus Raphistoma Hall, 1847 73 Genus 'fScalites Emmons, 1842 74 Family HOLOPEIDAE Wenz, 1938 74 Genus Raphistomina Ulrich and Scofield, 1897 74 Genus \Pachystrophia Perner, 1903 74 Genus Sinutropis Perner, 1903 75 Genus Umbospira Perner, 1903 75 Genus Holopea Hall, 1847 75 Family LESUEURILLIDAE, new family 75 Genus -\Eccyliopterus Remele 1888 75 Genus ^Lesueurilla Koken, 1898 76 Genus Mestoronema, new genus 76 Genus Pararaphistoma Vostokova, 1955 76 Family fHELiCOTOMlDAE Wenz, 1938 76 NUMBER 88 Genus ^Lophonema Ulrich in Purdue and Miser, 1916 76 Genus Linsleyella Rohr, 1980 76 Genus ^Helicotoma Salter, 1859 77 Genus Palaeomphalus Koken, 1925 77 Genus Ophiletina Ulrich and Scofield, 1897 77 Family fEuOMPHALiDAE de Koninck, 1881 77 Genus ^Boucotspira Rohr, 1980 77 Genus \Euomphalopterus Roemer, 1876 77 Genus Spinicharybdis Rohr and Packard, 1982 78 Genus -\Poleumita Clarke and Ruedemann, 1903 78 Genus Nodonema Linsley, 1968 78 Genus Centrifugus Bronn, 1834 78 Genus Euomphalus Sowerby, 1814
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