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Phylogenetic Relationships of the Earliest Anisostrophically Coiled Gastropods

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Phylogenetic Relationships of the Earliest Anisostrophically Coiled Gastropods SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY • NUMBER 8 Phylogenetic Relationships of the Earliest Anisostrophically Coiled Gastropods Peter J. Wagner SUB Gfittingen 215 090 918 2002 B 3558 Smithsonian Institution Press »\ Washington, D.C. 2002 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 II. 1. "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 m 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 fSlNUOPEIDAE 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 fOPHlLETOiDEA trans, nov. Knight, 1956 70 Genus ^Ophilela Vanuxem, 1842 71 Genus "fLecanospira Butts, 1926 71 Genus Ecculiomphalus Portlock, 1843 71 Genus fAsgardaspira, new genus 71 Genus Lytospira Koken, 1896 71 Superfamily MACLURITOIDEA Fischer, 1885 71 Genus -fMacluritella Kirk, 1927 72 Genus -fTeiichispira 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 fEuOMPHALOlDEA de Koninck, 1844 73 Family fRAPHiSTOMATiDAE 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 -fEccyliopterus Remele 1888 75 Genus ^Lesueurilla Koken, 1898 76 Genus Mestoronema, new genus 76 Genus Pararaphistoma Vostokova, 1955 76 Family fHEUCOTOMlDAE Wenz, 1938 76 NUMBER 88 Genus fLophonema 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 JEUOMPHALIDAE de Koninck, 1881 77 Genus \Boucotspira Rohr, 1980 77 Genus fEuomphalopterus Roemer, 1876 77 Genus Spinicharybdis Rohr and Packard, 1982 78 Genus -fPoleumita Clarke and Ruedemann, 1903 78 Genus Nodonema Linsley, 1968 78 Genus Centrifugus Bronn, 1834 78 Genus Euomphalus Sowerby, 1814 78 Genus Straparollus de Montfort, 1810 79 Genus Micromphalus Knight, 1945 79 Family ANOMPHALIDAE Wenz, 1938 79 Genus ^Trochomphalus Koken, 1925 79 Genus Pycnomphalus Lindstrom, 1884 79 Family PSEUDOPHORIDAE Miller, 1889 79 Genus Pseudophorus Meek, 1873 79 Genus Pseudotectus Perner, 1903 80 Genus fDiscordichilus Cossmann, 1918 80 Genus Hystricoceras Jahn, 1894 80 Genus Streptotrochus Perner, 1903 80 Genus Elasmonema Fischer, 1885 80 Suborder MURCHISONHNA COX and Knight, 1960 80 Superfamily fMuRCHlSONlOlDEA Koken, 1896 81 Family fHORMOTOMIDAE Wenz, 1938 81 Genus -fHormotoma Salter, 1859 81 Genus iCoelocauIus Oehlert, 1888 81 Genus Catazone Perner, 1903 81 Genus Mesocoelia Perner, 1907 81 Genus Plethospira Ulrich in Ulrich and Scofield, 1897 81 Family MURCHISONIIDAE Koken, 1896 82 Genus ^Murchisonia d'Archaic, 1841 82 Genus Morania Horny, 1953 82 Genus Michelia Roemer, 1854 82 Superfamily LOXONEMATOIDEA Koken, 1889 82 Family LOXONEMATIDAE Koken, 1889 82 Genus -fLoxonema Phillips, 1841 82 Genus fOmospira Ulrich and Scofield, 1897 .' 83 Genus Diplozone Perner, 1907 83 Genus Rhabdostropha Donald, 1905 83 Genus Spiroecus Longstaff, 1924 83 Genus Macrochilus Lindstrom, 1884 83 Genus Stylonema Perner, 1907 84 Superfamily EOTOMARIOIDEA Ulrich and Scofield, 1897 84 Family EOTOMARIIDAE Wenz, 1938 84 Genus \Clathrospira Ulrich and Scofield, 1897 84 ?Genus Spirotomaria Koken, 1925 84 Genus \Eotomaria Ulrich and Scofield, 1897 84 Genus Paraliospira Rohr, 1980 84 Genus Liospira Ulrich and Scofield, 1897 85 Family fGosSELETINlDAE Wenz, 1938 85 VI SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY Subfamily fEuRYZONlNAE, new subfamily 85 Genus ^Deaechospira, new genus ;. .. 85 Genus ^Cataschisma Branson, 1909 86 ?Genus Palaeoschisma Donald, 1902 86 Genus \Pleurorima Perner, 1907 86 Genus Euryzone Koken, 1896 86 Genus Latitaenia Koken, 1925 86 Subfamily GOSSELETININAE Wenz, 1938 86 Genus \Stenoloron Oehlert, 1888 86 Genus Platybron Oehlert, 1888 87 Genus Umbotropsis Perner, 1907 87 Family fPHANEROTREMATlDAE Knight, 1956 87 Genus ^Brachytomaria Koken, 1925 87 Genus Phanerotrema Fischer, 1885 87 Genus Ulrichospira Donald, 1905 88 Family LUCIELLIDAE Knight, 1956 88 Genus Conotoma Perner, 1907 88 Genus Prosolarium Perner, 1907 88 Genus Oehlertia Perner, 1907 88 Superfamily LOPHOSPIROIDAE Wenz, 1938 89 Family LOPHOSPIRIDAE Wenz, 1938 89 Genus Ectomaria Koken, 1896 89 Genus Donaldiella Cossmann, 1903 89 Genus Lophospira Whitfield, 1886 89 Genus Proturritella Koken, 1889 89 Genus Eunema Salter, 1859 89 Genus Gyronema Ulrich in Ulrich and Scofield, 1897 89 Genus Ruedemannia Foerste, 1914 89 Genus Loxoplocus Fischer, 1885 90 Genus Longstaffia Cossman, 1908 90 Genus Arjamannia Peel, 1975 90 Genus Trochonemella Okulitch, 1935 90 Family TROCHONEMATIDAE Zittel, 1895 90 Genus Trochonema Salter, 1859 90 Genus Globonema Wenz, 1938 90 Superfamily STRAPAROLLINOIDEA, new superfamily 90 Family STRAPAROLLINIDAE, new family 90 Genus ^Strapamllina Billings, 1865 90 Genus Daidia Salter, 1859 90 Genus Haplospira Koken, 1897 91 Superfamily SUBULITOIDEA Lindstrom, 1884 91 Family SUBULITIDAE Lindstrom, 1884 91 Genus ^Eroicaspira, new genus 91 Genus ^Subulites Emmons, 1842 91 Genus Fusispira Hall, 1872 91 Genus Cyrtospira Ulrich in Ulrich and Scofield, 1897 91 Conclusions 91 Appendix 1: Characters and Character States 93 Appendix 2: Data Matrix 100 Appendix 3: Stratigraphic Data 134 Literature Cited 143.
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