S M I T H S O N I A N C O N T R I B U T I O N S TO P A L E O B I O L O G Y ? N U M B E R 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 A B S T R A C T 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 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 \Discordichilus Cossmann, 1918 80 Genus Hystricoceras Jahn, 1894 80 Genus Streptotrochus Perner, 1903 80 Genus Elasmonema Fischer, 1885 80 Suborder MURCHISONIINA COX and Knight, 1960 80 Superfamily JMURCHISONIOIDEA Koken, 1896 Family JHORMOTOMIDAE Wenz, 1938 Genus "\Hormotoma Salter, 1859 Genus -fCoelocaulus Oehlert, 1888 Genus Catazone Perner, 1903 Genus Mesocoelia Perner, 1907 Genus Plethospira Ulrich in Ulrich and Scofield, 1897 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 ^Loxonema 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 -fClathrospira 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 fGosSELETlNiDAE Wenz, 1938 85 vi SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY Subfamily f EURYZONINAE, 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 Platyloron Oehlert, 1888 87 Genus Umbotropsis Perner, 1907 87 Family IPHANEROTREMATIDAE 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 ^Straparollina 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 Phylogenetic Relationships of the Earliest Anisostrophically Coiled Gastropods Peter J. Wagner1 Introduction The renewed interest in gastropod phylogenetics (see Bieler, 1992, for a review) has generally neglected fossil taxa. This is unfortunate because solely neontological studies exclude many interesting gastropod clades and likely underestimate the com? plexity of gastropod evolution. The earliest members of di? verse, long-lived clades also might possess informative combi? nations of plesiomorphies and apomorphies (see Gauthier et al., 1988; Donoghue et al., 1989). Many major taxa (extinct and extant) apparently diverged very early in gastropod history (Knight et al., 1960) without obvious intermediates (Erwin, 1990a); therefore, a phylogenetic analysis of the earliest gastro? pods could contribute much to contemporary ideas about gas? tropod relationships. The last phylogenetic study to concentrate on Early Paleo? zoic gastropods was by Knight (1952). Later workers presented alternative ideas about relationships among particular taxa (e.g., Yochelson, 1967, 1984; Runnegar, 1981; Linsley and Kier, 1984), but none have conducted large-scale phylogenetic analyses. In this paper, I discuss the results of phylogenetic analyses that encompass 295 species of early anisostrophically 'Author's Note: The scientific content of this paper originally appeared as a chapter in the author's 1995 dissertation, "The Generation and Maintenance of Morphologic and Phylogenetic Diversity among Early Gastropods" (Uni? versity of Chicago). This paper was slightly modified from that chapter and was accepted for publication in 1996. Thus, this work is older than studies published by the author since 1997, and readers should consider conclusions in those papers to supercede conclusions in this work. Also, this paper refers to no studies published after 1997 (except in cases where this paper originally referred to works in press or in preparation, or to published abstracts now rep? resenting journal articles). As a result, several studies using similar methods and/or data sets are not mentioned. The author regrets any confusion that might arise because of the misleading publication date. Peter J. Wagner, Department of Geology, Field Museum of Natural History, 1400 South Lake Shore Drive, Chicago, Illinois 60605-2496. E-mail address: pwagner@fmnh.org. coiled gastropods. These results are contrasted with the many previous phylogenetic estimates that gastropod systematists have presented. Although the primary goal of the study is to es? timate relationships among early gastropods, the study ad? dresses (by necessity) some larger phylogenetic issues. These topics include the relationship of early gastropods to other early Paleozoic molluscs, the relationships of the Paragas- tropoda (Linsley and Kier, 1984) to each other and to gastro? pods and other molluscs, and the relationships of the problem? atic bellerophonts to gastropods, other molluscs, and (to a much lesser extent) each other. This study differs from its predecessors in two ways. First, it includes only species that appeared from the Cambrian through the Silurian, whereas studies such as those cited above esti? mated gastropod phylogeny using species that appeared long after major taxa (i.e., orders and suborders) diverged (Yochel? son, 1984). Second, this study is essentially a species-level analysis, whereas previous studies typically used one or two exemplar species to represent each higher taxon. Erwin (1990b) rendered both strategies suspect, as a cladistic analysis of higher taxa that used late Paleozoic exemplar species sug? gested very different relationships than did an analysis of the same higher taxa that used early Paleozoic exemplars. The analysis presented herein avoids both problems by making no assumptions about the definitions or diagnoses of higher taxa. ACKNOWLEDGMENTS A Smithsonian Predoctoral Fellowship during the summer of 1991, sponsored by D.H. Erwin and E.L. Yochelson, funded much of the research presented herein. Additional research funding was provided by an NSF doctoral dissertation im? provement grant, the Geological Society of America, Sigma Xi, the Hinds Fund from the Division of the Biological Sci? ences at the University of Chicago, National Aeronautics and Space Administration (U.S.A.) grant NAGW-1693 to J.J. Sep- koski, Jr., and NSF grant #EAR-84-177011 to D. Jablonski. I SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY thank the following people for allowing me access to the col? lections housed at their institutions (and also for outstanding hospitality): J. Cooper and N.J. Morris at The Natural History Museum, London; R. Owens at the Welsh Museum of National History in Wales; J. Bergstrom and V. Jaanuson at the Swedish Natural History Museum in Stockholm; R.J. Horny at the Nar- odni Museum, Prague; and the staff at the Okresni Museum, Rokycany, Czech Republic. The scanning equipment and gray? scale printers that made many of the figures possible were made available by D. Rowley and D. MacAyeal. I thank the following for extended to near-endless discussions about gas? tropod biology and evolution, and/or phylogenetic theory and methods in general, or for suggesting useful revisions to this manuscript: W. Allmon, J. Alroy, R. Bieler, D.H. Erwin, M. Foote, D. Jablonski, R. Homy, M. LaBarbera, D. Lindberg, R. Linsley, D. Miller, N.J. Morris, P.J. Morris, J.S. Peel, J.A. Schneider, J.J. Sepkoski, Jr., S. Suter, E.L. Yochelson, and the Field Museum systematics discussion group. By no means do I wish to imply that any of the above agree with all (or any) of my results, methods, and/or conclusions. "Archaeogastropods"?A Temporary Definition Workers have classified most early Paleozoic gastropod spe? cies within the order Archaeogastropoda; however, the defini? tions and diagnoses of the taxon has changed drastically in re? cent years, and there is little consensus on the taxon's meaning or utility. Hickman's (1988) monophyletic definition of extant taxa (Pleurotomarioidea + Fissurelloidea + Haliotoidea + Scis- surelloidea + Trochoidea; = the Vetigastropoda of Salvini-Pla- wen and Haszprunar, 1987) is not obviously applicable to early Paleozoic species. Neither is Haszprunar's (1988) paraphyletic definition, which is based on the nervous system. Gastropod systematics is weighed down with excessive names and an un? stable taxonomy (Bieler, 1992), so retaining a definition of the "Archaeogastropoda" that is monophyletic through the Silurian (and hence used in quotes) is useful, if only for purposes of this discussion. For this paper, I define "archaeogastropods" as dex- trally coiled molluscs with an anal emargination (i.e., a sinus) and a peripheral band (see diagnosis below), plus all of their descendants through the Silurian. This definition differs from that of other paleontologists (e.g., Knight et al., 1960) and also from the paleontological definition of the Vetigastropoda given by Tracey et al. (1993) in that "archaeogastropods" include not only putative pleurotomarioids and trochoids, but also the earli? est species assigned to the Apogastropoda2 and other orders. "Archaeogastropods" might be synonymous with a crown- group definition of gastropods (i.e., the last common ancestor of all extant gastropods and all of its descendants), but this analysis cannot demonstrate that possibility. Review of Previous Phylogenetic Hypotheses A standard neontological depiction of gastropod phylogeny suggests that the planispiral Bellerophontina3 were the earliest gastropods and that bellerophontinae were ancestral to anisos- trophic pleurotomarioids (e.g., Barnes, 1987; see Figure lA). Pleurotomarioids later produced other archaeogastropods, apo- gastropods, and the Patellogastropoda (e.g., Fretter and Gra? ham, 1962). The traditional paleontological depiction is simi? lar, essentially differing only by suggesting that a second extinct group of anisostrophic species (i.e., the Macluritoidea and Euomphaloidea) evolved from bellerophontinae (e.g., see Knight, 1952; Knight et al, 1960; Figure IB). Proposed alternatives exist to nearly every relationship shown in Figure lA,B. Some alternatives are only slightly dif? ferent. For example, Koken (1898, 1925; also N.J. Morris and Cleevely, 1981; P.J. Morris, 1991) thought that macluritoids and pleurotomarioids shared a helically coiled ancestor (Figure lc). Other proposals are radically different. A key controversy concerns the affinities of the Bellerophontina. Yochelson (1967, 1984) suggested that bellerophontinae evolved from (rather than giving rise to) pleurotomarioids (Figure ID). Ul? rich and Scofield (1897) considered bellerophontinae derived, but they thought that bellerophontinae and pleurotomarioids evolved independently from limpet gastropods (Figure IE). Haszprunar (1988) also considered limpet gastropods to be the ultimate gastropod ancestors of bellerophontinae, without com? menting on the relationship between bellerophontinae and heli- The Apogastropoda include fossil taxa, such as the Loxonematoidea, that Wenz (1938) and Knight et al. (1960) classified in the Caenogastropoda as well as modern caenogastropods, allogastropods, opisthobranchs, and pulmonates (Tracey et al., 1993). Therefore, I use the Apogastropoda as a replacement for the traditional paleontological definition of the Caenogastropoda. In this discussion and elsewhere, the Bellerophontina refers to a diagnosed taxon, which presumably should represent a monophyletic or paraphyletic gToup. "Bellerophont" denotes a grade of bilaterally symmetrical, planispiral molluscs that might be polyphyletic. I use the former when discussing potential clades and the latter when discussing a morphologic type. FIGURE 1 (opposite).?Summary of previous phylogenetic hypotheses for the Gastropoda. "T" denotes the hypothesized onset of torsion, which is the chief synapomorphy of gastropods. A, Traditional neontological hypothesis. B, Tra? ditional paleontological hypothesis (e.g., Knight, 1952), with bellerophontinae as gastropods that give rise to macluritoids and pleurotomarioids separately, with pleurotomarioids giving rise to murchisonioids. (Murchisonioids later gave rise to apogastropods.) C, Macluritoids and pleurotomarioids sharing a pleurotomarioid-like ancestor (e.g., Koken, 1898, 1925; N.J. Morris and Cleevely, 1981; P.J. Morris, 1991). D, Bellerophontinae as derived pleuroto? marioids (Yochelson, 1967, 1984). E, Bellerophontinae and pleurotomarioids derived separately from early limpet gastropods (Ulrich and Scofield, 1897). F, Bellerophontinae as monoplacophorans with no close relationship to gastro? pods (e.g., Wenz, 1938; Runnegar and Pojeta, 1974; N.J. Morris, 1990). G, Bel? lerophontinae as monoplacophorans and gastropods and macluritoids (includ? ing onychochilids) evolving torsion independently from a pelagiellid ancestor (Runnegar, 1981). H, Bellerophontinae as a collection of monoplacophorans and primitive gastropods (e.g., Homy, 1965; Peel, 1991a). I, Macluritoids as paragastropods and not closely related to gastropods (Linsley and Kier, 1984). NUMBER 88 __ Patellogastropods A T v /^ A Monoplacophorans -A , Bellerophonts ?k Pleurotomarioids - ^ Other Archaeogastropods Caenogastropods + "Higher" Gastropods Subulitoids T - ^ Macluritoids ?V, Euomphaloids <_ B Monoplacophorans i - k Bellerophonts Murchisonioids -X ?W Loxonematoids \ 7^ \J >^ Pleurotomarioids - ^ Trochoids ^ Other Caenogastropods Patellogastropods - - . Euomphaloids ?* Macluritoids C T V v f >k Loxonematoids Monoplacophorans I k Bellerophonts ? \ Raphistomatoids " T r r v " ^ Murchisonioids v " ^ Murchisonioids \ i ^7 >i - * Pleurotomarioids ^ * X > ? Caenogastropods D Monoplacophorans L? Pleurotomarioids ? \ Macluritoids \ Euomphaloids i i i Subulitoids jP Trochoids Bellerophonts Trochoids Patellogastropods ?Patellogastropods -^i'ateiiogasrropoas y ^^ ?Euomphaloids^* Macluritoids n Tv / \ A y C, Monoplacophorans ?? Limpet Urgastropod^^*Raphistomatoids , _ Pleurotomarioids ? ? Trochoids N i -^?Eotomarioids -^Bellerophonts ^ 2* Murchisonioids # Loxonematoids ? ? Subulitoids .Bellerophonts F Monoplacophorans Murchisonioids ? Loxonematoids ? Caenogastropods ? Patellogastropods Pleurotomanoids ^ Other Archaeogastropods Macluritoids ? Euomphaloids > Subulitoids Bellerophonts ^1 G Monoplacophorans -? Onychochilids ? * Macluritoids - ^ Pellagiellids T- Pleurotomarioids > Other Gastropods H Monoplacophoran T v Gastropod V ? ? ? J V ^ . _ Bellerophonts " 7 Bellerophonts ~ T Pleurotomarioids?^ Other Gastropods 1 Monoplacophorans Macluritoids > Other Paragastropods Bellerophontoids > Pleurotomarioids ? Other Gastropods SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY cally coiled gastropods. Conversely, segmented muscle scars in Devonian Cyrtonella Hall, 1879, led Wenz (1938) to conclude that the Bellerophontina were untorted molluscs. Runnegar and Pojeta (1974, 1985; also Runnegar, 1981, 1996; Dzik, 1981; N.J. Morris, 1990) advocated similar views and considered bel? lerophontinae and gastropods to be distant relatives (Figure 1F,G). Finally, some workers considered traditional definitions of the Bellerophontina to include both untorted and torted mol? luscs (e.g., Horny, 1965, 1991, 1992b; Harper and Rollins, 1982; Peel, 1991a,c, 1993; Wahlman, 1992). The last author suggested that bellerophont gastropods evolved from anisos? trophically coiled gastropods, whereas the first three authors considered some bellerophont gastropods ancestral to all other gastropods (Figure lH). The second model is not inconsistent with most traditional models (Figure 1 A,B), because traditional models often do not specify the gross morphology of the imme? diate ancestor of gastropods. The affinities of the Macluritoidea represent another source of contention. Traditionally, workers considered macluritoids to be one of the earliest offshoots within the gastropod clade (e.g., Knight, 1952; N.J. Morris and Cleevely, 1981). Consider? ations about molluscan functional biology led Linsley and Kier (1984) to suggest that macluritoids belonged to the Paragas- tropoda, a group of untorted molluscs (Figure ll). The Hyper- strophina (= Mimospirina of Dzik, 1982, plus the Omphalocir- ridae), a group of molluscs with highly ultra-dextral shells4, are pivotal in this controversy. Workers traditionally assumed that macluritoids evolved from the Onychochilidae (e.g., Knight, 1952; Linsley and Kier, 1984; Runnegar, 1981, 1996), an early family of the Hyperstrophina. Runnegar (1981) also considered onychochilids to include the ancestors of true macluritoids. In this scheme, torsion evolved independently in true gastropods and onychochilids, with untorted pelagiellids being the com? mon ancestor of both torted onychochilids and true gastropods. Functional analyses by P.J. Morris (1991), however, suggested that taxa such as Maclurites Le Sueur were torted, whereas hy- perstrophinae were untorted. Morris and others (e.g., Peel, 1991a,b) considered hyperstrophinae and gastropods (includ? ing Maclurites) as only distant relatives. seum, London, the National Museum of Wales (Cardiff), the Natural History Museum of Sweden (Stockholm), the Narodni Museum (Prague), and the Okresni Museum (Rokycany, Czech Republic). These specimens included material from North America (including Alaska), China, Malaysia, the British Isles, Scandinavia, Estonia, and western and central Europe. I also examined material collected in Kentucky (U.S.A.) by the United States Geological Survey (Wagner, 1990). Published photographs were used when available. The analyses presented herein include 295 species. I ex? cluded most members of the Bellerophontinae, although sev? eral were used as outgroups (see below). I also excluded most species assigned to the Subulitidae, except for the earliest and also some later ones suspected of belonging to other clades (Er? win, 1992). Finally, I also omitted species assigned to the Platyceratidae. In this case, I attempted to include some of the earliest species, but I could not identify their relationships sat? isfactorily (see below). In addition to the deliberately excluded taxa, I also excluded at least 50 "archaeogastropod" species that likely represent valid taxa. Nearly all of those species are known from only one or a few localities, and I was not able to examine enough specimens to code them adequately; however, these species also appear to have been short-lived and apomor- phic. As I could include close relatives (which often were po? tential ancestors of the short-lived species), the exclusion of the short-lived species should not interfere with the stated goal of this paper, i.e., identifying the relationships among major gas? tropod taxa in the early Paleozoic. Notably, this analysis ex? cluded no species known from five or more localities. I examined multiple specimens of most species while coding the character states, which allowed me to evaluate ranges of in- traspecific variation and also to observe ontogenetic variation within species. I granted no special status to type material be? yond determining species assignments. This is important be? cause much of the material that I examined has not been de? scribed, especially from the Early Ordovician. Nearly all of the specimens examined fit within the diagnoses of previously de? scribed species. A question mark precedes a species name throughout the paper if I was unable to examine the type speci? men (or if the type was so poor as to be uninformative). Material SPECIMENS I based the phylogenetic analysis on specimens housed in the type, biologic, and stratigraphic collections at the National Mu? seum of Natural History (Washington, D.C), the Field Mu? seum of Natural History (Chicago), the Natural History Mu- "Ultradextral" shells often are described as "hypertrophic." Hyperstrophy and orthostrophy, however, refer to the orientation of the internal anatomy, whereas "dextral" and "ultra-dextral" refer to whether the shell coils "down" or "up" the coiling axis. Opercula indicate that ultra-dextral species, such as Ma? clurites, had orthostrophic organizations (Yochelson, 1990), so I use the latter set of terms instead. BlOGEOGRAPHY OF ANALYZED SPECIES The biogeographic affinities of the examined species are somewhat complex, largely because of geographic evolution from the latest Cambrian through the Silurian. "Archaeogastro? pods" from the latest Cambrian through the early Arenig (Early Ordovician) appear to have been restricted to the Laurentian fauna (i.e., eastern North America and Scotland). Even after the Early Ordovician, the tropical Laurentian fauna appears to have maintained the highest "archaeogastropod" diversity of the early Paleozoic realms. By the late Early Ordovician (i.e., middle Arenig), "archaeogastropods" also existed in the Baltic, Toquima-Table Head, Celtic, and Gondwanan faunas (see Neuman and Bruton, 1989; Cocks and Fortey, 1990, for general NUMBER 88 descriptions of those faunas). The Toquima-Table Head fauna represents a tropical to equatorial assemblage, whereas the Bal? tic fauna represents a temperate fauna, and the Celtic and Gondwanan faunas represent a temperate to near-polar assem? blage. The Toquima-Table Head and Celtic faunas (i.e., gastro? pod species from western North America, the northern east coast of North America, Wales, and parts of Norway) are some? what problematic. Some workers recognize both faunas as dis? tinct provinces that were unique to the late Early to Middle Or? dovician (i.e., Neuman and Bruton, 1989; Neuman and Harper, 1992). Others consider both faunas to be mixtures of Lauren? tian and Baltic faunas (McKerrow and Cocks, 1986; Cocks and McKerrow, 1993). Gastropods from the Celtic fauna are not well known, but the few described species (e.g., Neuman, 1964) are also known from the Toquima-Table Head faunas (pers. obs.). Although these gastropods do have affinities with both Laurentian and Baltic species, they appear to represent a separate fauna. Ordovician gastropods from the temperate-to- polar Gondwanan realm appear to have been rare, and I exam? ined species only from the Middle Ordovician of western and central Europe. Early to Middle Ordovician gastropods also have been reported from the Gondwanan faunas of South America (e.g., Beresi and Rigby, 1993), but I was not able to examine any specimens. Paleogeographic reconstructions sug? gest that those South American faunas should have been equa? torial, and general published descriptions suggest that the gas? tropods belonged to the tropical Toquima-Table Head fauna. "Archaeogastropod" biogeography simplified in the Late Or? dovician, which witnessed an increasing homogeneity among realms, not only for gastropods, but also for bryozoans (Anstey, 1986), trilobites (Cocks and Fortey, 1990), and brachiopods (Cocks and Ruang, 1988; Cocks and Fortey, 1990). Notably, all of the faunas are thought to have been closer to the equator dur? ing the Late Ordovician than they had been in the Early and Middle Ordovician (Scotese, 1989). The Early Silurian (i.e., Llandovery-Wenlock) shows still greater homogeneity, as "ar? chaeogastropods" appear to have represented a single equato? rial fauna. Some differentiation is noticeable by the Late Sil? urian (Ludlow-Pridoli), with the temperate Gondwanan fauna distinct from the tropical Laurentian and Baltic faunas. Cladistic Characters HOMOLOGY VERSUS ARCHITECTURE It is important to discuss codings and a priori hypotheses of homology when conducting cladistic or phenetic analyses. Un? fortunately, most of the terms used to describe gastropod shells (or portions of those shells) refer to architectural features that might or might not be present depending on the interactions of different character suites (e.g., coiling parameters and aperture shape). For example, terms such as "columella" or "umbilical carina" are not used to label homologous regions of the shell on species with very different gross morphologies. To avoid con? fusion, I avoid these terms and instead use slightly less com? mon terms. For example, the inner margin forms a columella on a "typical" gastropod shell (e.g., Lophospira perangulata (Hall, 1847)) and thus usually is labeled either the columella or the columellar lip (Cox, 1960; e.g., Figure 2A). On species with sufficiently low shell curvature, such as Clathrospira el- liptica (Hisinger, 1829) (Figure 2 B ) or Spiroraphe bohemica Barrande in Perner, 1907 (Figure 2c), the "columellar lip" fails to form a columella unless it is extremely thick. A columella might still be formed by the parietal inductura (i.e., a funicle) on species with the same basic shell geometry (e.g., Siluripho- rus gotlandicus (Lindstrom, 1884)) (Figure 2D). Finally, the homologous region on nearly planispiral taxa, such as Barne- sella llecanospiroides Bridge and Cloud, 1947 (Figure 2E) or Palliseria robusta Wilson, 1924 (Figure 2F), forms the base of the shell rather than a columella (see, e.g., Figure 2F). Given the inconsistent relationship between this feature and the col? umella (and the fact that the columella is an architectural fea? ture rather than a true homology), I refer to that portion of the aperture as the inner margin. A columella at the base of the inner margin typically encir? cles the umbilicus and hence is labeled a basal carina. On nearly planispiral forms, however, the carina is at the periphery of the shell base. Accordingly, I refer to the feature as a basal carina. In addition, the "upper" and "lower" ramps of normally coiled species (e.g., Figure 2 A - D ) are the "right" and "left" ramps of nearly planispiral species (Figure 2E,F). AS the latter terminol? ogy also refers to the post-torsional orientation of internal or? gans, I use right ramp throughout this paper. Batten (1989) la? beled the left ramp the alveozone, a term that I use herein. THE PAUCITY OF SHELL CHARACTERS REVISITED Appendix 1 gives the characters and character states used in this study. Appendix 2 gives the data matrix. I discuss addi? tional data relevant to both appendices below. I used 143 char? acters encompassing 352 character-states for this study (Ap? pendix 1; note that I count continuous characters as only one state). There are four reasons why I used so many characters. First, the species analyzed herein encompass sufficient mor? phological disparity to be classified in multiple orders. One usually can describe any specimen with fewer than 50 charac? ters, but one needs many more characters to describe the entire spectrum of gastropod shell morphologies. The second reason for the high number of characters is that this is a species-level analysis. Supraspecific phylogenetic analyses of gastropods average less than one shell character per operational taxonomic unit (OTU), whereas species-level anal? yses average more than one shell character per OTU (Table 1). This likely is because many shell characters vary within clades as well as among clades, so if a study uses only a few exemplar species to represent higher taxa, then many shell characters be? come uninformative. Thus, many shell characters that are use? ful in phylogenetic analyses of closely related species are not useful in phylogenetic analyses of distantly related ones. SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY RR PB NBC FIGURE 2.?Some basic terms and characters used in the analyses. AC = Carina at base of Alveozone; AL = Alveozone (= Post-Torsional Left Ramp; Batten, 1989); B = Base of Inner Margin; BC = Basal Carina (typically an umbilical carina); IM = Inner Margin; PB = Peripheral Band (almost always located at the sinus apex); RR = Right Ramp; SA = Sinus Apex (for specimens without a peripheral band). A, Lophospiraperangulata (Hall): IM thick, nearly straight, trends nearly parallel to coiling, reflects around coiling axis and forms columella; BC absent; AC present, sharp; left and right ramps symmetrical in shape (concave) and length; PB trilineate (i.e., bearing both peripheral lira and a medial lirum), bisects left and right ramps and oriented approximately 30? adapically from perpendicular to IM. B, Clathrospira elliptica (Hisinger) IM little thicker than rest of shell, curved and nearly parallel to coiling axis; BC and AC absent; AL and RR convex, but AL shorter than RR; PB bilineate (i.e., peripheral lira present), bisecting AL and RR and oriented nearly perpendicular to IM. C, Spiroraphe bohemica Barrande in Perner: IM curved, oriented nearly 30? off parallel to coiling axis; BC and AC absent; AL and RR convex, but RR more convex and longer than AL; PB bilineate, falling partially on RR and oriented approximately 10? abapically from perpendicular to IM. D, Siluriphorus gotlandicus (Lindstrom): IM strongly curved, thicked above base, approximately 15? off perpendicular to coiling axis; BC present, thin and weak; AC present, "squared" and strong; AL and RR convex, with RR very short relative to AL; PB absent, with SA near suture. E, Barnesella llecanospiroides Bridge and Cloud: IM runs nearly perpendicular to coiling axis (forming flat base of shell); BC thick but weak; AC absent; AL and RR flat and long, symmetric in shape and length; PB monolineate, sharp, weak. F, Palliseria robusta Wilson: IM thick, curved and oriented at very high angle (-120?) relative to coiling axis (forming curved base of the aperture); BC dull thickening; AC absent; AL and RR equal in length, but with convex AL and flat or concave RR. The third reason for the large number of characters is that this analysis used many traits that are inapplicable to extant species. Characters describing sinuses, peripheral bands, slits, and differential shell asymmetry are not relevant to studies of most extant taxa, but they are very important to the study of early Paleozoic species (Table 2). For example, only a few types of peripheral bands exist on a handful of extant species; however, most early Paleozoic species possessed peripheral NUMBER 88 TABLE 1.?Operational taxonomic units (OTUs) versus shell characters for some previous phylogenetic analyses of gastropods. Included are the number and taxonomic level of taxa and the number of shell characters that they utilized. Only characters that could be utilized in this study were counted, so some characters, such as shell min? eralogy, were excluded. "No. of OTUs" is the number of taxa that were analyzed, including outgroup taxa. Study Ponder and Lindberg (1996) Haszprunar (1988) Houbrick(1988) Ponder and Lindberg (1997) Davis etal. (1985) Hickman and McLean (1990) Hickman (1996) Hickman (1996) Davis and Pons da Silva (1984) Ponder (1984) Bieler (1988) Reid (1989) Jung(1992) Kool (1993b) Kool (1993a) Houbrick(1984)/ Erwin (1988) Michaux(1989) Wagner (1995a) This study Taxon Gastropoda Gastropoda Cerithioidea Gastropoda Rissoidea Trochoidea Trochoidea Turbinidae Hydrobiidae Iravadiidae Architectonicidae Littorinidae Planorbidae Ocenebrinae Rapaninae Cerithidea Glyptospira Ancillinae Lophospiridae "Archaeogastropods" Taxic level of OTUs "superfamily" "superfamily" superfamily "family" subfamily subfamily/tribe subfamily/tribe subfamily/tribe genus genus genus genus genus genus genus subgenus species species species species No. of OTUs 22 15 15 25 8 29 20 9 8 14 12 36 9 5 24 4 8 32 42 295 No. of shell characters 3 3 15 5 1 26 20 13 4 9 12 2 27 3 4 9 21 32 79 352 TABLE 2.?Importance of "archaeogastropod" shell characters in this analysis versus their importance in other analyses. Columns give the number of character states used in each study for the particular trait. Asterisks denote examples discussed in the text. PB denotes characters describing perhipheral bands, and InAn denotes characters describing apertural inclination. Coiling denotes coiling and/or growth parameters. Study Ponder and Lindberg (1996) Haszprunar (1988) Houbrick(1988) Ponder and Lindberg Davis etal., 1985 (1997) Hickman and McLean (1990) Hickman (1996) Davis and Pons da Si Ponder (1984) Bieler (1988) Reid (1989) Kool (1993a) Kool (1993b) Houbrick(1984) Erwin (1988) Jung (1992) Michaux(1989) Wagner (1995a) This study va(1984) Characters Slit 0 1* 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 1 9 Sinus 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 3 21 PB 0 0 0 0 0 0 0* 0 0 0 0 0 0 0 3 0 0 5 58 Asymmetry 0* 0 0 1 0 0* 0 0 0 0 0 0 0 0 0 0 0 0 15 InAn 0 0 0 0 0 1 1 0 3 0 0 0 0 0 0 0 0 4 5 Coiling 0 2 1 0 1 2 1 1 0 1 1 1 0 0 1 5 6 5 11 Ornament 0 0 1 0 0 4 1 1 4 8 1 0 2 2 11 4 1 4 16 bands, and several different types existed. The transition from symmetric morphologies to asymmetric ones introduces addi? tional character states, as both the left and right sides of several features must be coded independently. Some additional impli? cations of this pattern are discussed in detail below. Finally, this study used finer divisions of shell characters than employed by previous workers. Gastropods can produce very similar shell shapes using different combinations of growth parameters, aperture shapes and orientations, and shell thicknesses. Character complexes often have been treated as single characters, but I divided these into several characters and multiple states. Hickman and McLean's (1990; see also Hick? man, 1996) cladistic analysis of the Trochoidea used tangential and radial apertures (i.e., inclined versus noninclined) as two states of one character. Early Paleozoic gastropods produced inclined apertures in many ways (Table 2; Appendix 1, charac- SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY ters 109-119). Figure 3 illustrates some examples. The aper? tures of some species, such as Siluriphorus gotlandicus, form a plane that is inclined relative to the coiling axis (Figure 3A). For other species, different parts of the aperture are inclined to different degrees. On species such as the Middle Ordovician Clathrospira conica Ulrich and Scofield, 1897, the left5 side of the aperture is much more inclined than the right side (Figure 3 B ) . The opposite condition exists on Pleurorima migrans (Barrande in Perner, 1907), where the right side provides most of the inclination (Figure 3c). The base of the aperture also can deviate from radial by pro? jecting either anteriorly or posteriorly. An example of the former is the Middle Ordovician species Helicotoma tennes- seensis Ulrich and Scofield, 1897 (Figure 3D). The Early Or? dovician species Pararaphistoma qualteriata (Schlotheim, 1820) is an example of the latter (Figure 3E). Note that the pos? terior projection is not the same as a basal excavation (where the base has a sigmoidal or U-shape), which is coded as a sepa? rate character. Posterior projection of the base is especially common in species with very low spire heights (especially planispiral species) and might serve to alter the center of grav- "Left" and "right" here and elsewhere refer to the post-torsional left and right. For species with selenizones, the right corresponds to the area between the selenizone and the coiling axis (i.e., usually above the "upper whorl"). ity within a nearly planispiral shell in a manner that is analo? gous to the standard inclination of the aperture on a normally coiled shell (e.g., Linsley, 1977). Anterior projection of the base occurs in both low-spired and high-spired species and also might serve to enhance apertural inclination. Aperture shape is another trait that previous workers have used as a single character (e.g., Houbrick, 1984; Hickman and McLean, 1990). Different parts of "archaeogastropod" aper? tures (or, more appropriately, different portions of the outer and inner whorl faces) can have different shapes (e.g., convex, flat, concave) on the left and right side of the aperture owing to the decoupling of left-right homologies with increasing asymme? try. Accordingly, I coded the shape of the left and right sides of the aperture separately rather than trying to code a whole shape for the aperture. An additional important point to this breakdown of character complexes concerns the functional biology of gastropod shells. Features such as spire height, apertural inclination, aperture shape, and columellar type strongly affect how easily snails can balance and move their shells (e.g., Linsley, 1977, 1978; Mc- Nair et al., 1981; Signor, 1982). Thus, convergent evolution of similar functional complexes among different clades is a major concern. The characters comprising functional complexes are used herein, however, rather than the complexes themselves. Because different combinations of characters will yield very zzzzzzzzzzi Inclination of whole aperture Inclination of right side Inclination of left side Projection of the base Plane radial to the coiling axis FIGURE 3.?Different ways in which early Paleozoic gastropods produced tangential apertures. A, Inclination of the whole aperture. B, Inclination provided primarily by the left side of the aperture only. Note that the lower part of the aperture has a stronger inclination than the upper part. C, Inclination provided primarily by the right side of the aperture. D, Anterior projection of the aperture base. E, Posterior projection of the aperture base. NUMBER 88 similar character complexes (e.g., gross aperture shapes, spire heights, overall apertural inclinations, and columellas), two distantly related but grossly similar morphologies that func? tional biologists might consider to be the "same" probably will have very different character codings. Similarly, two closely re? lated but grossly different morphologies might have very simi? lar character codings. This does not mean that the coding scheme used herein will identify all functional convergences. Functional convergence between closely related clades or in? volving simplification of the shell (and necessarily reducing applicable characters) will confound any coding scheme. The breakdown of the characters (coupled with the use of strati? graphic data as tests of parsimony estimates), however, should increase the accuracy of the phylogenetic analyses. SINUSES, SLITS, SELENIZONES, AND PERIPHERAL BANDS Among modern gastropods, the sinus (a broad, acute emar- gination culminating at the presumed site of the exhalent cur? rent; Figure 4A) exists only on some vetigastropods (i.e., pleu? rotomarioids and some scissurelloids) and on some basal caenogastropods (e.g., turritellids), whereas the slit (a thin lin? ear cleft, also culminating at the presumed exhalent current; Figure 4 B ) occurs on other vetigastropods and on species in some other clades, such as the Architectonicidae. Neontolo- gists generally have ignored the sinus and have considered the slit to be a synapomorphy of the earliest gastropods (e.g., Hasz? prunar, 1988). Paleontologists noted that the sinus evolved prior to the slit, but most considered the two features to be ho? mologous (e.g., Knight, 1952). Horny (1962), however, sug? gested that the two features represented independent homo- logues, at least in the case of bellerophonts. This study supports Horny's idea. Although much attention has been paid to the slit, the sinus is far more informative phylogenetically, as si? nuses provide multiple characters that are distinguishable among nearly all "archaeogastropod" species (Table 2; see also Appendix 1, characters 1-11). Slits provide far fewer traits (Ta? ble 2; see also Appendix 1, characters 34-36), but as slits are much rarer than sinuses, they tend to be highly informative where they exist (see below). An associated issue herein is the relationship between the slit and the peripheral band (i.e., the "slit-band" of 19th and early 20th century literature). Knight (1934) relabeled the feature the "selenizone" and defined it as a structure generated by a slit. This assumed that the band was simply a distortion produced by a linear cleft in the shell. Several previous workers (e.g., Lindstrom, 1884; Ulrich and Scofield, 1897; Donald, 1902, 1906) had noted that slit-bands predate slits in the fossil record. Although slits were not common before the Devonian and were very rare during the Ordovician, the ubiquitous peripheral band appeared by the Late Cambrian. Knight (1941, 1952) later rec? ognized this and considered the band to be homologous on both slit-bearing and slitless species. Instead of abandoning the pre? vious morphogenetic hypothesis, Knight (1952) abridged it by inferring an unseen notch in the aperture that generated the pe? ripheral band for slitless species. Knight (and subsequent work? ers) used the term "pseudoselenizone" to describe such periph? eral bands. Despite frequent allusions to a notch, this feature has never been observed and its existence has been inferred solely on the assumption that a peripheral band is an artifact of a cleft in the aperture. Knight and others apparently did not consider the possibility that peripheral bands had no morphogenetic relation to slits or notches. One line of evidence suggesting that this is the case is that the pseudoselenizone of species with no such slit (e.g., "Longstaffid" "laquetta" (Lindstrom, 1884) (Figure 4A) typi? cally differs little from selenizones of closely related species with slits (e.g., "Seelya" lloydi (Sowerby in Murchison, 1839) (Figure 4B). A related point is that when the slit is a variable feature on individual specimens, the selenizone is unaffected. The growth lines of some species (e.g., Clathrospira subconica (Hall, 1847) (Figure 4C) suggest that the specimens had incon? sistent slits, i.e., the presence or absence of the slit varied. The growth lines within the peripheral band (i.e., lunulae) corre? spond with the growth lines outside the band, which indicates that the animal did not have a slit (or a "notch") when that part of the shell was secreted. There are fewer lunulae than growth lines, however, and the initial growth did not include shell dep? osition within the selenizone. This suggests that deposition within the peripheral band was halted for a period, resulting in the production (and subsequent lengthening of) a slit. The short slit later is filled, leaving the shell temporarily slitless. Despite the inconsistent nature of the slit, the peripheral band of C. subconica remains the same, which indicates that the slit is not responsible for the structure. Species such as Pararaphistoma qualteriata provide an ex? ample similar to that of Clathrospira Ulrich and Scofield (Fig? ure 4F). Pararaphistoma Vostokova species and their relatives lack slits on their juvenile whorls, and the presence of a slit sometimes is erratic on the adult whorls. The nature of the sele? nizone, however, does not vary over ontogeny or change with the production of a slit. The apex of a sharp (e.g., V-shaped) sinus, such as seen on Clathrospira, Pararaphistoma, and most other early Paleozoic taxa, might act as Knight's notch. Several species with very sharp sinuses (e.g., the Silurian Sinuspira tenera Barrande in Perner, 1907), however, lose the peripheral band over ontogeny without any corresponding ontogenetic changes in sinus mor? phology. If the band were purely an artifact of sinus morphol? ogy, this would not be possible. If a slit generated the peripheral band, then we would expect the position of the two features to coincide on the shell. This usually is true, as the peripheral band usually borders the slit (e.g., Figure 4 B - D ) , especially on species with bilineate bands (i.e., two carinae). Species such as Pararaphistoma qualteri? ata, however, have slits that are wider than the peripheral band and are positioned somewhat differently on the aperture (Fig? ure 4 F ) . Another extreme is shown by Oehlertia scutulata 10 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY A B C D FIGURE 4.?Sinuses, slits, and periph? eral bands. A, Gastropod with a sinus (the curved emargination), a periph? eral band (the paired threads at the apex of the sinus), but no slit. B, Gas? tropod with a sinus, peripheral band, and periodic production of a slit in the middle of the peripheral band, c, D, Gastropods with sinus, peripheral bands, and slits. E, Gastropod with a narrow, shallow sinus, and a broad peripheral band lunulae angling straight into a very narrow slit that is bordered by two sharp threads. F, Gas? tropod with a sharp, single carina for a peripheral band, and a slit that begins above the base of the peripheral band and terminates above the top of the peripheral band. In this case, separate views of the right (upper) and left (lower) ramps are shown, as well as a profile with arrows denoting the posi? tions of the ramps. Note that the slit is not always produced, as some growth lines give way immediately to sigmoi- dal-shaped lunulae. A and C-F taken from Lindstrom (1884); B taken from Ulrich and Scofield (1897). (Lindstrom, 1884), which has a very thin slit (shown by the in? ner pair of sharp lira in Figure 4E) that is much narrower than the peripheral band (shown by the outer pair of sharp lira in Figure 4E). Note that the outer pair appear to be homologous with the peripheral band of other gastropods, whereas the inner pair represent a feature unique to Oehlertia and related taxa. In summary, (1) peripheral bands evolved long before slits did, (2) peripheral bands on early Paleozoic gastropods cannot have been created by slits or notches, and (3) there is no evi? dence that "notches" of any sort ever existed. The definition of a "pseudoselenizone" is rendered logically complex and cer? tainly must be discontinued. Knight's definition of the seleni? zone also is logically complex (as the peripheral bands do not coincide with slits in either ontogeny or phylogeny). Knight (1952) and previous workers did, however, recognize the ho? mology of selenizones and pseudoselenizones. Although a cor- NUMBER 88 11 rected definition of "selenizone" could be used, the term has become too strongly identified as a by-product of a slit rather than an independent anatomical feature. Accordingly, the use of selenizone also should be abandoned, at least in reference to the peripheral band of most Paleozoic gastropods. The term "fasciole" suffers from the same problems. In the absence of a more appropriate term, I will use the cumbersome "peripheral band" to label the band at the apex of the sinus (and presum? ably denoting the location of the anus). In addition to clarifying terminology, the importance of this lengthy discussion is that it indicates the need for a slightly broader coding scheme than is implied by the earlier literature. The morphogenetic scheme of Knight (1952) implies that the peripheral band is an architectural artifact similar to a col? umella, which therefore would require coding only the pres? ence/absence of a notch/slit (e.g., Hickman, 1996). Instead, the presence or absence of a slit represents one character and the presence/absence of a peripheral band represents another char? acter. As noted above, the presence/absence of a sinus is a com? pletely independent third character. SHELL MINERALOGY AND PROTOCONCHS Many workers consider shell mineralogy and protoconch morphology to be phylogenetically informative (e.g., Batten, 1972, 1984; Bandel, 1988, 1991; Ponder, 1990a, 1990b); how? ever, I did not use these characters in this analysis. Shell miner? alogy probably varied widely among "archaeogastropods," but it rarely is possible to identify the exact types of shell mineral? ogy. Taphonomic characteristics reveal which species had at least partially calcific shells, but recrystalization usually ob? scures the relative amount of calcite or its exact nature. Most of the specimens that I examined were silicified. Silicification can reveal the number of mineral layers and their relative thick? nesses, but it leaves no other evidence about those layers (see Carson, 1991). When possible, I do discuss some basic aspects of mineralogy (i.e., aragonitic versus calcific shells), especially if they support or contradict the results presented herein. I omitted protoconchs for a different reason. A distinct proto? conch morphology typifies modern species, but not early Pale? ozoic ones. I examined many extremely well-preserved speci? mens representing a number of different taxa, but the boundaries between protoconch and teleoconch usually were vague at best (some exceptions are discussed below). I also ex? amined microfossils from beds rich in gastropods. Sinus and peripheral band morphologies usually were not observable on microfossils, but the basic profiles of the shells matched those of macrofossils known from the same beds. Dzik (1978) fig? ured protoconchs of two early Paleozoic "archaeogastropod" species, which also possessed adult profiles but lacked clearly defined sinuses and peripheral bands. Many species show distinct ontogenetic changes. These dif? fer from the abrupt transitions between protoconchs and teleo- conchs because the changes are gradual. For example, species classified as Macluritella Kirk, Teiichispira Yochelson and Jones, and Malayaspira Kobayashi have juvenile shells that are similar in overall morphology to Prohelicotoma Flower (Figure 5A). Similarly, species of Pararaphistoma Vostokova and Cli? macoraphistoma Vostokova have juvenile shells similar to that of Lesueurilla Koken (Figure 5B). Therefore, I did not code ju? venile shell types as separate characters. Instead, I coded the types of ontogenetic changes (e.g., counter-clockwise rotation of the aperture and differential expansion of the left side in ma- cluritids, or clockwise rotation of the aperture and increased translation in raphistomatids) as present or absent. I then coded the traits that changed during ontogeny as polymorphic, with all the states produced by an ontogenetic trajectory coded as present. Phylogenetic Analysis CHARACTER DEWEIGHTING 1?BALANCING CONTINUOUS CHARACTERS There are two standard justifications for character weight? ing: accounting for differential homoplasy among characters (e.g., Farris, 1969; Goloboff, 1993) and mitigating the effects of ordered characters (Thiele and Ladiges, 1988; Chappill, 1989; Hauser and Presch, 1991; Skelton and McHenry, 1992). I did not weight or reweight characters because of homoplasy. I did deweight ordered characters, however, to accommodate continuous characters. In all cases these represented continu? ous morphologic features, such as shell growth parameters or apertural inclination. An ordered trait with 10 characters will result in a nine-step difference between species coded as " 1 " and those coded as "10"; however, the maximum difference < N FIGURE 5.?Ontogenetic changes in shell morphology. A, Morphology of adult Prohelicotoma and juvenile Macluritella or Teiichispira. B, Morphology of adult Teiichispira, with negative translation, the left side of the aperture expanded, and the entire aperture rotated counter-clockwise, c, Morphology typical of adult Lesueurilla and juvenile Climacoraphistoma or Pararaphis? toma. D, Morphology of adult Climacoraphistoma, with higher translation and the aperture rotated clockwise. 12 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY between a presence/absence character is only one step (i.e., "0" to "1"). The concern herein is that ordered characters with many states will have an excessively strong effect on a cladis? tic analysis. The "over-weighting" effects of ordered charac? ters noted by Hauser and Presch (1991) is of particular con? cern herein, as malacologists have documented the potential plasticity of shell characters within species (e.g., Kemp and Bertness, 1984; Palmer, 1985; Boulding and Hay, 1993). Un? weighted continuous characters then might separate close rela? tives by several steps due to evolutionarily minor differences. Therefore, I deweighted continuous characters as follows: continuous character weight = - 1 no. of character states - 1 (see Thiele and Ladiges, 1988). One presence/absence synapo? morphy always contributes at least as much (and usually more) to the parsimony analysis as does a continuous synapo? morphy. Deweighted continuous characters are denoted with a "C" in Appendix 1. Farris' (1990) contentions against the deweighting of con? tinuous characters assume that discrete characters are recog? nizable. This assumption is unsound for the continuous char? acters used herein. Techniques used to categorize quantitative characters (e.g., Thorpe, 1984; Archie, 1985; Goldman, 1988) all assume that intraspecific variation is roughly equal for all species, but intraspecific variation can vary among different morphotypes (Schindel, 1990) and at different times in a clade's history (Hughes, 1991). The latter problem appears to have been the case among early gastropods (Wagner, 1995b, 1996). These factors also discouraged the use of simple maxi? mum likelihood methods for quantitative characters (e.g., Felsenstein, 1981, 1988); therefore, I used segment coding (Chappill, 1989), which simply divides continuous characters into equal distributions (e.g., "narrow" < 0.1, "medium" > 0.1 and < 0.2, etc.). This defines no discrete states, so the charac? ters are meaningful only if ordered. Segment coding links ad? jacent characters so that "narrowness" is a synapomorphy of a "narrow" and "very narrow" species relative to a "wide" out? group. This is an inexact method of describing character states, but it makes the fullest use of the available data and thus should be used in analyses such as this one. Several authors have criticized segment coding (e.g., Pimen- tel and Riggins, 1987; Cranston and Humphries, 1988) on the grounds that because one cannot define homologies in a repeat- able manner for continuous characters, such features lack phy? logenetic information. These arguments assume that meaning? ful evolution always proceeds like a cladistic character state optimization, with states clicking on and off without intermedi? ate forms (see Janvier, 1984; Gayon, 1990). This is a conten? tious view of morphologic evolution that is better examined within the contexts of phylogenetic estimates than assumed when conducting phylogenetic analyses. CHARACTER DEWEIGHTING 2?ASYMMETRY AND CHANGING HOMOLOGIES I employed character deweighting for another situation that previous workers have not discussed. Evolutionary events can couple or decouple homologies (e.g., Schaeffer and Lauder, 1986; Wake and Roth, 1989; Atchley and Hall, 1991), result? ing in multiple characters becoming one or one character be? coming several. In such cases, some species possess multiple characters for a morphologic structure whereas other species have only one. Among gastropods, the loss of bilateral sym? metry represents an example of the decoupling of homologies. This has had appreciable effects on the internal anatomy. Among gastropods, left and right homologues often serve dif? ferent functions (or the right organ is absent); these organs are paired in other molluscs. Indeed, trends toward asymmetrical conditions have been so pervasive that parsimony optimiza? tion finds bilaterally symmetrical conditions to be derived within gastropods (e.g., Ponder and Lindberg, 1996). This pat? tern also is reflected in shell morphology. For example, the left and right sides of the sinus and aperture are symmetrical on most early appearing species, but asymmetrical morphologies appeared by the Early Ordovician. This leads to a coding para? dox. The left and right can change independently on asymmet? rical species and, thus, can represent separate character states. If one codes the left and right sides as separate traits, however, then symmetric species have many of the same characters coded twice. Ideally, one would code features that vary in symmetry in the following manner. First, one would distinguish "symmet? ric" versus "asymmetric" as a presence/absence character. There are two mutually exclusive types of asymmetries: greater development of the right side and greater development of the left. Without a priori evidence that one type cannot evolve from the other, one should code these as unordered character states. Among species with the same type of asymmetry, any dif? ference on the left or right side should be coded as one step (or ln th of a step for continuous characters with n+1 states). When describing the difference between asymmetric and sym? metric species, the coding must avoid assuming how asymme? try evolved. An aperture with a more pronounced left side can be produce by enlarging the left side or contracting the right side. Thus, one should code such asymmetric species so that they are equally close to symmetric species with identical left or right sides. For example, consider a species on which the right side of the sinus retreated at 30? whereas the left side re? treated at 50?. That species should be considered equally sim? ilar to species with symmetric sinuses retreating at either 30? or 50?. The 30?:50? species should be coded as one step away from either the 30?:30? or 50?:50? species (i.e., the absence versus the presence of symmetry), with the 20? difference on the left or right side considered to be produced by the onset of asymmetry. NUMBER 88 13 If a symmetrical species differs from both the left and right sides of an asymmetrical species, then one should code species so that there is a difference of one step (symmetry versus asymmetry) plus the minimum number of steps needed to make either the left or right side of the symmetrical species identical to the left or right side of the asymmetrical one. Con? tinuing the example started above, the 30?:50? species would differ from a 40?:40? species by one step (symmetry to asym? metry) plus l/nth steps (either 30? to 40? on the left side or 50? to 40? on the right side, with l/nth being one over the number of character states +1; see the discussion of continuous charac? ters, above). The 30?:50? species would differ from a 20?:20? species by one step (symmetry to asymmetrically deep left side) plus two more steps (i.e., 20? to 40? on the left side). The major difference between species for the other side is attrib? uted to the change in symmetry. This scheme is the most parsi? monious possible because it assumes the minimum differences between species. Step matrices (Swofford and Olsen, 1990; Maddison, 1993) permit the character coding scheme described above. Unfortu? nately, using step matrices slows down computer analyses so much that I could not analyze even small data sets; therefore, I could not use step matrices in this analysis. Instead, I coded the symmetry as "present," "absent (left side greater)" or "absent (right side greater)." I then weighted the left and right sides of potentially asymmetric characters as separate characters. The deweighting means that 30?:30? species and 50?:50? species differ by 2n steps (30? to 50?) instead of 4n (30? to 50? on the right side and 30? to 50? on the left side). The 30?:50? species, however, differs from either species by 1 step (symmetry to asymmetry) plus 2n (the difference on the right or left side). There are two disadvantages to this scheme. First, the number of differences between symmetrical and asymmetrical species is slightly greater than it should be (i.e., l+2n steps instead of 1), as differences in both the left and right sides are tallied. More importantly, this scheme means that the cladistic analysis can imply chimeras. Figure 6A gives an example where parsi? mony will predict an ancestor with symmetrical sinus but dif? ferent left and right sides. In these cases, I had to reoptimize the character states so that no chimeras existed (e.g., Figure 6B). I then kept the shortest trees without chimeras. Until cladistic programs can implement step matrices efficiently, the imper? fect approach used herein represents the best solution for this type of problem. CHARACTER ANALYSES I analyzed the data using PAUP 3.1 (Swofford, 1993). I used heuristic searches, with multiple replications and random se? quence addition of species employed to account for islands of similar trees (see D.R. Maddison, 1991). This does not guaran? tee finding the shortest trees, especially for a matrix of this size; therefore, I reanalyzed smaller portions of the data, with the initial results providing estimates of subclade membership and appropriate outgroups. Ultimately, these analyses were re? duced to four clades, which are referred to below as the "helic- otomatids," "euomphalinae" (minus "helicotomatids"), "eoto- marioids,'' and "murchisoniinae" (minus "eotomarioids"). A disadvantage of this strategy is that the basic clade assign? ments might represent local minima rather than global solu? tions (D.R. Maddison, 1991). This also permitted the analysis of a far greater number of trees within those local minima. The phylogenetic estimate presented is not derived from the most-parsimonious cladograms, but instead from the most- parsimonious trees that stratigraphic data could not reject. Many systematists have asserted that stratigraphic data cannot sym: L: shallow R: shallow > > asym: L: mod R: deep sym: L: shallow R: shallow FIGURE 6.?Effects of asymmetrical characters. Hypothetical cladogram for species on which the sinus becomes asymmetrical. A, The most parsimonious optimization of characters states, given that symmetry/asymmetry is coded as a presence/absence feature, and the left and right sides are coded independently. Note that the node pre? dicts an impossible collection of character states, as the hypothesized ancestor is symmetrical yet features differ? ent left and right sides. B, The shortest acceptable interpretation, in which the hypothetical ancestral morphology is assumed to be identical to the middle morphology. Coding with step matrices (Swofford and Olsen, 1990) would permit parsimony analyses to produce these results; however, these were not computationally feasible in this study. 14 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY offer tests of parsimony estimates, on the assumption that the effects of incomplete sampling distorts stratigraphic data more than it does parsimony estimates (e.g., Smith, 1994). Simulations studies indicate the opposite, however, at least when sampling levels are comparable to those observed among most shelly invertebrates (Huelsenbeck, 1991a; Wag? ner, 2000). Several methods that integrate stratigraphic data into phylogenetic analyses have been proposed to date (e.g., Fisher, 1991; Huelsenbeck, 1994), but like parsimony itself, these methods are fundamentally ad hoc because they use stratigraphy as an arbitrary parsimony or reweighting crite? rion. A non-ad hoc method uses confidence intervals on strati? graphic ranges to test parsimony assessments of phylogeny (Wagner, 1995a). I saved all trees as short as or shorter than the shortest tree known to be fully consistent with strati? graphic data and then used a computer program that searched for trees that implied no statistically significant stratigraphic gaps. I evaluated the significance of stratigraphic gaps using confidence intervals on stratigraphic ranges (Strauss and Sa? dler, 1989; Marshall, 1990). As a first step, I used Appearance Event Ordination (AEO; Alroy, 1994a) to array the fossilifer- ous horizons that preserve "archaeogastropods," bellero? phontinae, tergomyans, bivalves, and rostroconchs with ara- gonitic shells6. I then used the AEO ordination as a substitute for stratigraphic ranges and calculated confidence intervals using the formula given by Strauss and Sadler (1989) and Marshall (1990) (Appendix 3). The calculation of confidence intervals assumes that hori? zons are distributed randomly throughout stratigraphic ranges (Strauss and Sadler, 1989; Marshall, 1990; but see Marshall, 1994). Differences in ecology, taphonomy, and/or biogeogra- phy will violate this assumption (Wagner, 1995a). Closely re? lated species might be assumed to have similar ecologies and taphonomies owing to phylogenetic autocorrelation; however, different clades and morphotypes likely will have different ecologic distributions and taphonomies, and hence have dif? ferent fossil records. The absence of species from horizons in which no close relatives or morphologically similar snails oc? cur are not meaningful (e.g., Bottjer and Jablonski, 1988) and to include such horizons inflates confidence intervals unrealis- tically. To control for such differences in expected preserva? tion patterns, I described the stratigraphic ranges of species based on only horizons including members of their more in? clusive clades (see Appendix 3). To control for differences in biogeography, I determined stratigraphic ranges and confi- 6Species with calcific shells occur in horizons in which aragonitic shells are absent or rendered unrecognizable. Including these horizons biases the statisti? cal analyses of stratigraphic ranges for calcific species by giving such species more finds (and thus shorter confidence intervals) than is possible for arago? nitic species with identical stratigraphic ranges. Using only horizons that in? cluded noncalcitic "archaeogastropods" offers a taphonomic control and, thus, each species is evaluated based on fossiliferous horizons that could have pre? served any gastropod shell. dence intervals within the individual provinces discussed above (see "Biogeography of Analyzed Species," above). As a result, some very widespread species have stratigraphic ranges in multiple provinces. When sister species existed in different provinces, the stratigraphic ranges could not be contrasted di? rectly. In such cases, stratigraphic correlations based on Har- land et al. (1990) were used to determine whether any strati? graphic gap was significant. This was done loosely, so if parsimony considered a Caradocian species from Baltica to be the sister species of a Llanvirn species from Laurentia, then the relationship was rejected only if the confidence interval's lower bound for the Baltic species was restricted to the Lland- eilo or Caradoc. If that lower bound for the Baltic species ex? tended into the Late Llanvirn, then the estimated relationship was accepted even if the Laurentian species was known from the Early Llanvirn. This accommodates the imprecision of cross-provincial stratigraphic correlations. Appendix 3 gives the stratigraphic ranges of analyzed spe? cies plus the 95 percent confidence intervals on those strati? graphic ranges. Appendix 3 also includes species that could not be included in the cladistic analyses due to inadequate numbers of well-preserved specimens, but that might bridge strati? graphic gaps. I used ACCTRAN character optimization, which favors par? allelisms to reversals (Swofford and Maddison, 1987). The choice of character optimization does not affect the initial re? sults of parsimony analyses. Ancestor-descendant hypotheses eliminate many significant stratigraphic gaps, however, and ACCTRAN makes it less likely that putative ancestors will have apomorphies. Any putative autapomorphies of ancestral species must be considered reversals in descendants, which lengthens a cladogram. Therefore, ACCTRAN ultimately can imply shorter lengths for identical cladistic topologies (Wagner, 1995a). Culling trees that implied statistically significant strati? graphic gaps (i.e., gaps greater than the 95% confidence inter? val extensions on the stratigraphic ranges of relevant species) resulted in the analysis finding very few equally parsimonious alternatives. Typically only a few trees out of several thou? sand would contain no significant stratigraphic inconsisten? cies. Some subclades did produce multiple trees of equal length and no significant inconsistencies. In these cases, I used the following criteria. First, I selected the tree that re? quired the fewest unknown ancestors. This represents a sec? ondary parsimony criterion that other workers have advocated (Alroy, 1995; see also Fisher, 1994; Smith, 1994). Second, I chose trees that had the smallest stratigraphic parsimony debt (Fisher, 1994; Suter, 1994). Any stratigraphic gaps at this point were not statistically significant; however, given the choice of two otherwise equal assessments of a phylogeny, it is logical to select the one that comes closest to predicting the observed pattern in the fossil record (Fisher, 1991; Smith, 1994). Finally, I chose some trees simply because a particular set of synapomorphies led me to prefer that topology over its NUMBER 88 15 rivals. This last criterion is obviously ad hoc; therefore, when discussing these trees below, I describe the rival topologies and detail why I accepted one over the other. Other phylogenetic methods do yield results that have some notable differences from those presented herein. This is partic? ularly true of strict parsimony, as characters or (especially) suites of characters identified by some methods as parallelisms between sister clades are identified as synapomorphies of more inclusive clades by parsimony. The general results de? scribed herein (i.e., the membership of basic clades and the re? lationships among those basic clades) are replicated by other phylogenetic methods, including strict parsimony. Thus, a ma? jor change in phylogenetic methods would be required to ob? tain radically different results using this character data. Sam? pling is another concern, as the ability of parsimony to reconstruct phylogeny decreases with decreasing sample size (Lanyon, 1985; Lecointre et al., 1993). Sampling also de? creases the accuracy of methods incorporating stratigraphic data, albeit to a lesser extent (Wagner, 2000). The sampling density of early gastropods, however, appears to be quite good. Based on the metrics of Foote and Raup (1996), be? tween 50% and 60% of the broadly distributed species appar? ently are included in this analysis (estimates vary according to binning criteria and subclade). These levels are even higher during the crucial early phases of gastropod evolution (i.e., the latest Cambrian and Early Ordovician of Laurentia, where es? timates improve to 70%). This means that we should have sampled many direct and indirect ancestors (see Foote, 1996), especially from the critical intervals during which the major groups were diverging. Such sampling greatly increases the efficacy of phylogenetic methods (Huelsenbeck, 1991a) and greatly reduces the concern that new finds will radically alter the estimates presented herein. I place least confidence in the estimated relationships among groups appearing in the earliest Silurian, as this represents the interval of poorest sampling. Unfortunately, this poor sampling seems to coincide with rapid diversification during the rebound from the end-Ordovician mass extinction. As a result, relation? ships among clades appearing in the Early Silurian form many polytomies, and there are several species and very small clades with very uncertain affinities. The discovery of heretofore un? known Llandovery species undoubtedly will change some of the relationships proposed herein. Many of the cases where this is especially true are emphasized in the text, with the accepted and alternative estimates presented. It should be stressed that these difficulties concern relatively fine-scale relationships, however, and do not effect the estimated relationships among the major groups that were established in the Ordovician. Thus, Silurian sampling is unlikely to affect the basic results of this analysis. As noted above, obtaining radically different results probably will require radically different reinterpretations of shell characters. CAMBRIAN MOLLUSCS AND THE CHOICE OF AN OUTGROUP Smith (1994) suggested that, of the available methods for rooting a cladogram, the outgroup method of polarizing charac? ters makes the fewest a priori assumptions. Choosing an out? group however, makes a major assumption about a group's phylogeny, namely, that the close relatives of the group of in? terest (i.e., the ingroup) are known (Adrain and Chatterton, 1990). This often makes the choice problematic (e.g., Ballard et al., 1992). The different phylogenetic models summarized above and in Figure 1 suggest very different outgroups for the "archaeogastropods"; essentially, nearly every possible rela? tionship among bellerophontinae, macluritinae, and "archaeo? gastropods" has been proposed at one point. Ballard et al. (1992) addressed a similar problem for arthropods by using a wider phylogenetic analysis to establish an appropriate out? group. I adopted a similar strategy by including over 20 differ? ent Cambrian molluscs as potential outgroups. These included species assigned to the Bellerophontina, Onychochilida, Pe- lagiellida, Helcionelloida, and Tergomya. I also included two Late Cambrian members of the Hypseloconidae, which might represent the ancestors of cephalopods (Yochelson et al., 1973; Webers and Yochelson, 1989; but see Teichert, 1988). These species are important because cephalopods are likely gastro? pods' closest relatives among the major extant molluscan classes (Naef, 1911; Wingstrand, 1985). I included an Early Ordovician cyrtonelloid, as the only known Late Cambrian rep? resentative of the group is represented by only a few poorly preserved specimens (McGhee, 1989). Finally, I also included some early Ordovician bellerophonts, owing to the relatively poor preservation of known Cambrian bellerophonts. I coded all outgroup species as if they were gastropods. For example, I treated the circumbasal carinae of onychochilids as a gastropod peripheral band. Thus, the analysis would not sepa? rate onychochilids and pelagiellids from "archaeogastropods" based on a priori interpretations of homology or assumptions about higher taxonomic associations. Density of species sampling strongly affects the precision of phylogenetic analyses (Lecointre et al., 1993). The outgroup analysis represents a very incomplete sample of the known Cambrian molluscs, which in turn represents only a portion of the species that actually existed. One might worry about the potential affect of species that I did not include. To address this, I reran the analyses multiple times after jackknifing the out? group (i.e., deleting one of the species; see Lanyon, 1985). I also employed rudimentary rarefaction (see Raup, 1975; Lecointre et al., 1993), running 50 analyses with random sub? sets of 50% and 75% of the outgroup species each. The rarefied analyses test whether the cladistic relationships among the in? group species are dependent on a few outgroup species. If the relationships within the ingroup vary widely depending upon the outgroup species included (especially at the 75% level), then estimated relationships among ingroup species likely de? pend upon the inclusion of particular outgroup species. This 16 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY suggests that adding more species could easily change esti? mates of relationships among "archaeogastropods" and that more outgroup species are needed before the results should be trusted. Conversely, if the relationships are stable (especially at the 50% level), then the ingroup results obviously are not strongly dependent on particular outgroup species. In this case, adding new Cambrian molluscs is not likely to affect phyloge? netic inferences among "archaeogastropod" species. Results CLADOGRAM STATISTICS AND DESCRIPTIONS I subjected the character-state matrix to two tests for phylo? genetic signal, thegj skewness test (Huelsenbeck, 1991b; Hil- lis and Huelsenbeck, 1992) and the Permutation Compatibility (PC) test (Alroy, 1994b). PAUP estimated the g{ statistic based on 10,000 random trees. For the entire matrix, g\ = -0.234 (p < 0.0001, Sokal and Rohlf, 1981:174). The significant result indicates that the character-state matrix is significantly more structured than a random collection of characters. Phylogenetic autocorrelation (and thus signal) is a primary candidate for that structure, although nonphylogenetic signals also will produce similar structure. Simulation studies suggest that the most par? simonious trees derived from structured matrices are likely to be good approximations of the true phylogeny (albeit, not nec? essarily exactly correct; Huelsenbeck, 1991a). The PC test examines whether a matrix contains greater char? acter compatibility and hierarchical signal than randomly scrambled matrices. Unlike other permutation tests (e.g., the Permutation Tail Test of Faith, 1991; Faith and Cranston, 1991), the PC test does not assume a particular model of tree- building. Also, although phylogenetic autocorrelation (see Raup and Gould, 1974; Felsenstein, 1985) will produce charac? ter correlations, other nonrandom but nonphylogenetic signals (e.g., allometry or functional complexes) also will produce character correlations (Kallersjo et al., 1992). The PC test, how? ever, should be less susceptible to nonphylogenetic signals (Al? roy, 1994b). Finally, the PC test is computationally feasible and could be applied to the whole matrix. Ten thousand permuta? tions found that none of the random permutations of the origi? nal data matrix possessed either the character compatibility or the hierarchical signal of the original data matrix. This indicates that the character state matrix possesses the type of character correlation that is expected if a phylogenetic signal is present. Each figure includes the consistency index (C.I.) and reten? tion index (R.I.) for the species shown, plus the immediate out? group (given at the base of the figure). The C.I. is simply the reciprocal of the average number of steps per derived state (Kluge and Farris, 1969); the R.I. is the difference between hy? pothesized tree length and theoretical minimum tree length di? vided by the difference between the theoretical maximum tree length and the theoretical minimum tree length (Farris, 1989; Archie, 1989). Note that the uninformative characters were ex? cluded when calculating C.I. and R.I. These homoplasy indices (especially C.I.) are meaningful only when compared to other analyses (Archie, 1989; Sanderson and Donoghue, 1989), and I include them for that reason only. Observed C.I.'s did typically fall within the range expected given the number of OTUs (see Sanderson and Donoghue, 1989). Workers discussing individual genera and species often have made suggestions about the phylogenetic relationships of those taxa. Whenever possible, I discuss whether this analysis cor? roborates those ideas. Also, I treat the taxonomic schemes pre? sented in major works (e.g., Knight et al., 1960, in the Treatise on Invertebrate Paleontology) as general phylogenetic schemes. In an evolving system, such as early Paleozoic gas? tropods, higher taxonomic definitions should be either mono? phyletic or cohesively paraphyletic (sensu Estabrook, 1986). Some of the higher taxa defined by Knight et al. (1960) were explicitly polyphyletic (see Tracey et al., 1993). I will treat all other taxonomic schemes, however, as general phylogenetic es? timates (i.e., proposed monophyly or cohesive paraphyly) when contrasting my results with the ideas of previous authors. Figures 7 through 35 present the accepted cladograms. The nodes are numbered uniformly throughout these figures: for example, node 1 is the same in Figures 7 and 8 and node 11 is the same in Figures 9 and 11. Nodes from the outgroup analysis are lettered rather than numbered. Species with open boxes are identical to the nodes to which they are attached; these species are considered to be ancestors in this study. Black boxes denote species with sufficient autapomorphies to be considered not an? cestral to any other taxa included in this analysis. Gray boxes denote equivocal cases (i.e., the taxa lack autapomorphies but there are missing data). The cladogram includes line sketches of each species. Note that these are drawn to a roughly uniform size, not to actual or relative scale. The figure captions give the morphologic changes for each node, as hypothesized by ACCTRAN parsimony (Swofford and Maddison, 1987). I have edited these so that the captions do not simply list the character-state changes, but instead de? scribe the types of morphologic change. Because one type of change can affect two otherwise independent characters (e.g., see the above discussions about the right and left sides of asymmetric morphologies), this presents a more realistic metric of the amount of morphologic change than does a simple count of synapomorphies. Several points about my use of taxonomy in the following sections need to be stressed here. When I use the Linnaean tax? onomy formally (e.g., the Murchisoniina, Murchisonioidea, Murchisoniidae, etc.), I am referring to taxa previously defined by other workers. The one exception is the boxed generic names shown on the cladograms in Figures 7-35, which outline the taxonomic revision discussed below. When I use suprage- neric names in quotes, I am labeling noteworthy clades defined by this analysis. Clade names are entirely informal and are de? signed to describe the phylogeny, obviating the need to follow conventional taxonomic rules. To avoid confusion, all sub? clades have different names than their more inclusive clades. NUMBER 88 17 For example, the "murchisoniinae" clade does not include a "murchisonioid" or "murchisoniid" subclade simply because the names look too similar in print. Also, not every species is assigned to a subclade at every level. This means that the para? phyletic collection of "raphistomatids" that do not belong to the "lesueurillines" or "holopeines" (see below) are not as? signed to any "-ine" subclade. I usually name clades after their least derived members in? stead of following the conventions of priority. I use previously named higher taxa whenever possible, but in many cases, no previously named higher taxa are appropriate. Naming clades after their plesiomorphic members is the only way that mor? phologic grade affects clade names. As a result, the "murchiso? niinae" included many species with morphologies that are very different from that of traditional diagnoses of murchisonioids. Finally, note that "subclade" describes a monophyletic group within the clade that is being discussed immediately. Thus, the "lesueurillines" are a "raphistomatid" subclade if I am discuss? ing relationships among the "raphistomatids," but it is a clade if I am discussing relationships among the "lesueurillines." Ta? ble 3 summarizes the "archaeogastropod" clades and subclades discussed herein. As noted above, this classification is for use in this discussion only and must not be interpreted as a formal taxonomic revision. I also include a numerical ranking of the clades and sub? clades in Table 3 and in the discussions below. This follows the scheme proposed by Hennig (1969) as a replacement for the Linnaean hierarchy. Again, this is not meant as a formal reclas? sification but simply as an aid for the reader. A formal taxonomic revision for the early gastropods based on this analysis and adhering to traditional Linnaean taxonomy is presented at the end of the paper. As noted above, the revised generic taxonomy also is presented in Figures 7-35. OUTGROUP ANALYSES Figure 7 shows the results of the outgroup analysis, with "ar? chaeogastropods" condensed into the Schizopea typica Ulrich and Bridge in Ulrich et al., 1930, plus Dirhachopea normalis Ulrich and Bridge in Ulrich et al., 1930, clade. Jackknifing the outgroups and rarefying them to 75% of the initial total had no effect on the relationships within the ingroup (Figure 8). Rar? efying the outgroup species to 50% of the initial total produced the same ingroup relationships in 46 of 50 analyses and in 205 of the 216 total trees produced by those analyses. The runs that did not result in S. typica being the stem-"archaeogastropod" did not include tropidodiscines, such as Strepsodiscus major Knight, 1948. In all other runs, those species are considered the immediate outgroups of the "archaeogastropods." Thus, the re? sults shown in Figure 8 depend only on our knowing about bel? lerophonts, such as S. major, and do not depend on the inclu? sion of any one species of Cambrian mollusc. This reduces the concern that adding more Cambrian molluscs will affect as? sessments of gastropod relationships. TABLE 3.?"Archaeogastropod" clades and subclades discussed herein. Note that I have added "-itcs" to described subclades within "-ides." "Archaeogastropods" (Figures 7-35, nodes 1-215) I. "Euomphalinae" (Figures 8-19, nodes 2, 11-108) 1.1. "Ophiletoids" (Figure 9, nodes 12-24) 1.1.1. "Ecculiomphalids" (Figure 9, nodes 15-18) 1.1.2. "Lytospirids" (Figure 9, nodes 19-24) 1.2. "Macluritoids" (Figure 10, nodes 25-36) 1.3. "Ceratopeatoids" (Figure 11-19, nodes 37-108) 1.3.1. "Raphistomatids" (Figures 11-14, nodes 42-69) 1.3.1.1. "Lesueurillines" (Figure 12, nodes 44-54) 1.3.1.2. "Scalitines" (Figures 13, 14, nodes 55-69) 1.3.1.3. "Holopeids" (Figure 14, nodes 62-69) 1.3.2. "Heiicotomids" (Figures 11, 15-19, nodes 41, 70-108) 1.3.2.1. "Ophiletinines" (Figure 15, nodes 74-80) 1.3.2.2. "Euomphalopterines" (Figures 16-19, nodes 81-108) 1.3.2.2.1. "Anomphalides" (Figure 16, nodes 83-87) 1.3.2.2.2. "Poleumitides" (Figures 15, 17, 18, nodes 81, 88-96) 1.3.2.2.3. "Pseudophorides" (Figure 19, nodes 97-108) II. "Murchisoniinae" (Figures 8, 20-35, nodes 7, 109-215) 11.1. "Plethospiroids" (Figure 20, nodes 110, 111) 11.2. "Straparollinoids" (Figure 21, nodes 112-118) 11.3. "Hormotomoids" (Figures 22-26, nodes 119-154) 11.3.1. "Subulitids" (Figure 22, nodes 120-123) 11.3.2. "Cyrtostrophids" (Figure 23-26, nodes 124-154) 11.3.2.1. "Goniostrophines" (Figure 24, nodes 131-138) 11.3.2.2. "Omospirines" (Figures 25, 26, nodes 139-154) 11.3.2.2.1. "Loxonematides" (Figure 25, nodes 141-145) 11.3.2.2.2. "Rhabdostrophides" (Figure 26, nodes 146-154) 11.4. "Eotomarioids" (Figures 27-35, nodes 155-215) 11.4.1. "Lophospirids" (Figure 27, nodes 158-165) 11.4.2. "Clathrospirids" (Figures 27-35, nodes 157, 168-215) 11.4.2.1. "Liospirines" (Figures 28, 29, nodes 170-181) 11.4.2.2. "Brachytomariines" (Figures 28, 30-35, nodes 167, 168, 182-215) 11.4.2.2.1. "?Palaeoschismides" (Figures 30, 31, nodes 183, 188) 11.4.2.2.2. "Phanerotrematides" (Figures 30, 32, nodes 185, 186, 190, 191) 11.4.2.2.3. "Luciellides" (Figures 30, 33, nodes 184, 197-201) 11.4.2.2.4. "Planozonides" (Figures 34, 35, nodes 202-215) H.4.2.2.4.1. "Coelozonites" (Figure 34, nodes 202-208) II.4.2.2.4.2. "Gosseletinites" (Figure 35, nodes 209-215) Figure 7 places onychochilids and pelagiellids as distant out? groups of tergomyans (nodes E-J), and considers "archaeogas? tropods" to be a tergomyan subclade (i.e., node L and above). This matches the predictions of Peel (1991a). In addition, these results support the hypothesis that gastropods originated in the late Middle to Late Cambrian (e.g., Linsley and Kier, 1984; Peel, 1991a; Tracey et al., 1993) rather than in the Early Cam? brian (e.g., Knight, 1952; Runnegar and Pojeta, 1974, 1985; Pojeta, 1980). The exact relationships among tergomyans are vague as there are no obvious synapomorphies linking hypselo- conoids, cyrtonelloids, or the clade of bellerophontinae + "ar? chaeogastropods". The sole synapomorphy linking Cyrtolites Conrad, 1838, to early gastropods at node N is the presence of a peripheral band, and most workers would not interpret the ca? rina of Cyrtolites as homologous with the peripheral band of gastropods. If the features are convergent, then it would be 18 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY CI: 0.464 RI: 0.677 TAXON ^ Pelagielloid Helcionelloid F ? ^ Tergomyan TXZl Onychochilid Hypseloconoid Bellerophontoid Euomphaloid Macluritoid Equivocal Helcionella subrugosa Oelandia rugosa Latouchella merino Coreospira rugosa Sinuella minuata Euomphalopsis involuta==^-:--^>^ Macluritella? walcotti/*^-^-^^^ Costipelagiella L^^J zazvorkai Cj?--JP^ j) Pelagiella ' 1 ^ ^ subangulata "Maclurites" thomsoni Scaevogyra swezeyi Kobayashiella circe Matherellina walcotti Matherella saratogensis Kiringella pyramidalis 2j Hypseloconus elongatus Knightoconus antarcticus Cyrtolites sp. Chippewaella patellitheca Strepsodiscus major Strepsodiscus paucivoluta ~ Chalarostrepsis praecursort Eobucania mexicana Peelerophon oehlerti Schizopea typica Dirhachopea normalis (go to Node 4) NUMBER 88 19 FIGURE 7 (opposite).?Outgroup analysis. Nodes with letters denote the out? groups, whereas nodes with numbers denote "archaeogastropods." Planispiral species are shown from a side view, anisostrophic species are shown from an apertural view. In this and subsequent figures the following conditions apply: (1) line drawings are not to scale; (2) numbers (or letters in small capitals) denote nodes, whereas numbers in parentheses denote characters (see Appen? dix 1); (3) abbreviations for synapomorphy linking include the following: ACh = anal channel beneath peripheral band; P = angle between inner margin and peripheral band (90? = peripheral band perpendicular to inner margin; 0? = par? allel); BC = basal carina; BL = paired peripheral lira of peripheral band; CA = coiling axis; E = shell expansion; GL = growth lines; IM = inner margin (= col? umellar lip of "normally" coiled species); K = shell curvature; LR = alveozone; LRC = alveozone carina; ML = single peripheral lira of peripheral band; PB = peripheral band; PI = parietal inductura; RR = right ramp; RRC = right ramp carina; T = shell torque; || = parallel to the coiling axis; 1 = perpendicular to the coiling axis; and (4) branch patterns denote previous higher taxonomic assign? ments (see text). Node A ("Helcionelloids"), tiny univalve shell with no sinus or PB, very strong growth lines, symmetric aperture with moderately broad, short ramps, posterior projection of the aperture, and channeled, curved IM. Node B (Latouchella merino clade), straight IM (95). Node C, very long ramps (55, 56); moderately broad aperture (58, 59); IM channel lost (102). Node D, ACh at top of aperture (48); IMNbase angle ? 60? (94); IM 1 to CA (98); base projected posteriorly -10? (116, 117); extremely low E (121); open coiling (123); anisostrophic shell (?) (125). Node E ("Paragastropods"), asymmetric aperture (contracted right side) with very long, narrowly projected ramps (55-59); IM at -45? to CA (98); PI || to aperture (104); moderate K (123); ani? sostrophic shell (125); loss of septation (128). Node F (Pelagielloids), sigmoi- dal aperture (13); ACh rotated towards top of aperture (48); IM at high angle relative to CA (98); inclined aperture (109); moderate K (123); anisostrophic shell (125). Node G (Onychochiloids), sharp ML (28); P = -10? (48); flat RR (51, 52); asymmetric aperture with longer right side (54-56); IM -15? past 1 to CA (98); aperture inclined backwards (115); anterior projection of aperture (116, 118); high K (123); moderate ultradextral coiling (126). Node H, long LR (56); asymmetric aperture (right side broader) (57-59); IM\base angle = 105? (94); curved IM (95). Node I, extremely strong GL (15); concave RR (52); very low E (121); high ultradextral T (126). Node J, P = -30? (48); very narrow broad right side of aperture (58). Node K {Sinuella minuata + Coreospira rug? osa), moderately long ramps (55, 56); very broad aperture (58, 59); RR swell? ing present (60); moderate E (121); small size (141). Node L (Tergomyans), very narrow aperture (58, 59); curved IM (95); base projected posteriorly -50? (117). Node M (Hypseloconoids), low K (123); large size (141). Node N, PB present (19). Node o (?Gastropods), presence of a sinus (1); dull, lump-like ML (28); extremely long ramps (55, 56); RR swelling present (60). Node P, swollen base of RR and LR (60, 73); low E (121); low K (123); curvature decreases over ontogeny (124). Node Q, narrow sinus (6, 7); BL present (21); slit present (34); weak swelling atop RR (60, 61); straight IM (95). Node R, moderate K (123); isometric curvature (124). Node 1 ("Archaeogastropods"), round ML (28); P = 100? (48); RR swelling dulls over ontogeny (62); moderate swelling at LR base (73, 74); BC present (89); IM 15? off parallel to CA (98); base projected posteriorly -30? (117); anisostrophic shell (125); low T (126). more parsimonious to consider cyrtonelloids and gastropods to have evolved separately from a limpet-like tergomyan similar to Kiringella Rosov, 1975. Horny (1965, 1991) objected to the suggested link between cyrtonelloids and gastropods in part be? cause of the absence of Late Cambrian cyrtonelloids. Since then, Berg-Madsen and Peel (1994) described Telamocornu, a cyrtonelloid from the Late Cambrian. Telamocornu existed in the Avalonian province, whereas early gastropods and their other tergomyan relatives dwelt in the Laurentian province, so there is still a biogeographic gap. Extended species-level analy? ses of Cambrian tergomyans obviously are needed to make any firm estimates of exact tergomyan relationships. Although this should be a priority for future research, the resampling results described above suggest that such research will not greatly af? fect estimates of relationships within the Gastropoda. The limpet-like bellerophont Chippewaella patellitheca Gunderson, 1993, is the least derived member of the tergomyan subclade that includes "archaeogastropods." Synapomorphies uniting this clade include a strongly curved sinus and a dull (i.e., broad but weakly expressed) peripheral band. This is a noteworthy result because C patellitheca has an external mor? phology appropriate for Haszprunar's (1988:407) "urgastro- pod." Chippewaella patellitheca is only known from one Late Cambrian specimen (Gunderson, 1993), so I consider this re? sult tentative. Excluding C. patellitheca produced the same in? group topologies, so even if C. patellitheca is important for un? derstanding the immediate relationships and origins of gastropods, it does not affect assessments of "archaeogastro? pod" relationships. Perhaps the most intriguing result of the outgroup analysis is that the Bellerophontina appear to be diphyletic, including a cohesive paraphylum of species assigned to the Tropidodisci- nae (i.e., Chalarostrepsis Knight, 1948, Eobucania Yochelson, 1968, and Peelerophon Yochelson, 1982) and a secondarily de? rived clade (Figure 8, node 10, which includes Sinuites Koken, 1896, and Owenella Ulrich and Scofield, 1897). Essentially, this results from the hypothesis that lenticular-shaped apertures with deep, hyperbolically curving sinuses, monolineate periph? eral bands, and posterior projection of the aperture (e.g., Strep? sodiscus or Schizopea Butts, 1926) are plesiomorphic, whereas apertures with swollen left and right ramps and compressed midsections, U-shaped, shallow sinuses, and bilineate or absent peripheral bands (e.g., Taeniospira Ulrich and Bridge in Ulrich et al., 1930, Sinuopea, or Sinuites) are derived. This means that the definition of "archaeogastropods" used herein includes many species assigned to the Sinuitidae. Four of the outgroup analyses run at 50% rarefaction considered Sinuites and rela? tives to be the immediate outgroup of coiled gastropods and Sinuopea sweeti Whitfield, 1882, to be the most primitive coiled gastropod; however, as noted above, these runs did not include species such as Strepsodiscus. Characters shared among Strepsodiscus and species such as Schizopea typica re? sult in a phylogeny that considers species with round apertures, U-shaped sinuses, and no peripheral bands to be highly derived rather than primitive. This study does not resolve whether taxa such as Strepsodis? cus or Tropidodiscus Meek and Worthen, 1866, were gastro? pods. Interpretations of Sinuites as untorted center partially on a hypothesized association between of the origin of torsion and anisostrophic coiling (e.g., Ghiselin, 1966; Runnegar, 1981). Regardless of the merits of that hypothesis, its implications are irrelevant for any bellerophonts that evolved from anisostroph? ically coiled species. The muscle scars of some sinuitids are segmented, which has been cited as evidence that sinuitids 20 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY CI: 0.528 RI: 0.758 Figure 7, NodeO TAXON Bellerophontina Euomphalina Pleurotomariina Murchisoniina I M I I H I I I I Schizopea Euom phs . Euconia Schizo. G asconadia Schiz. Sinuopea M urchs . Sinuopea Sinuitid s D I D D D Strepsodiscus major (see Figure 7) Schizopea typica (see Figure 7) Ophileta supraplana (go to Node 11) Prohelicotoma uniangulata (go to Node 25) Jarlopsis D ? I'D ? Euconia etna Rhombella umbilicata Dirhachopea normalis (see Figure 7) Gasconadia, put ilia Dirhachopea subrotunda Sinuopea basiplanata Taeniospirat 1st. clairi Hormotoma SH Isimulatrix (go to Node 109) ??| Taeniospira ? emminencis Sinuope sweeti Owenella antiquata Cloudia buttsi Sinuites sowerbyi NUMBER 88 21 FIGURE 8 (opposite).?Relationships among the earliest "archaeogastro? pods." Genus names in boxes denote proposed generic revisions (see "Sys? tematic Paleontology," below). For abbreviations, see legend to Figure 7. Node P, swollen base of RR and LR (60, 73); low E (121); low K. (123); cur? vature decreases over ontogeny (124). Node 1 "Archaeogastropods," round ML (28); P - 100? (48); RR swelling dulls over ontogeny (62); moderate swelling at LR base (73, 74); BC present (89); IM 15? off parallel to CA (98); base projected posteriorly -30? (117); anisostrophic shell (125); low T (126). Node 2 ("Euomphalinae"), moderately strong, flange-like ML (28, 29); PB curves adapically (42); ACh present (45); p = 70? (48); very long ramps (55, 56); weak LR swelling (71, 74); IM at -45? to CA (98); curved base (120). Node 3 (Rhombella clade), asymmetric sinus (shallower left side, wider right side) (2-7); p = 110? (48); asymmetric aperture (left side shorter and nar? rower) (54-59); IM\base angle = 120? (94); IM 30? off parallel to CA (98); thin PI (103); inclined aperture (109); moderate K (123); moderate T (126). Node 4 {Dirhachopea normalis clade), BL present (21); IM\base angle = 75? (94). Node 5, sinus angle - 50? (3, 4); PB bilineate throughout ontogeny (27, 41); P = 90? (48); RR swelling with isometric shape (62). Node 6, sinus angle = 40? (3, 4); sinus curvature continuous (9, 10); PB width = 20? (20); strong LR swelling (71, 74); IM || to CA (98); radial base (116, 118); septation absent (128). Node 7, PB width = 25? (20); elongated RR (54-56); broad symmetric aperture (58, 59); weak swelling atop RR (60, 61) and base of LR (74); IM thicker than shell (87); IM\base angle = 90? (94); thin PI (103); IM reflected around umbilicus (106); high K (123); very high T increasing over ontogeny (126, 127). Node 8, strong RR swelling (61). Node 9, narrow, U- shaped sinus (6, 7, 9, 10); PB lost (19); weak lunulae (39); high K (123); small size (141). Node 10 (Sinuitids), sinus angle = 30? (3, 4); isostrophic, planispiral shell (125, 126). were untorted (e.g., Runnegar, 1981) and as evidence that sinu? itids were highly derived gastropods (e.g., Haszprunar, 1988; Peel, 1991c; Horny, 1992b, 1995a). This analysis obviously supports the latter scenario. Given the sparse number of sinu- itid species included in this analysis, however, broader analyses of sinuitids and other bellerophonts obviously are necessary. RELATIONSHIPS AMONG EARLY ORDOVICIAN SPECIES Two major subclades of "archaeogastropods" diverged by the Tremadoc (Figure 8). These correspond to nodes 2 and 7 on the cladogram. One clade contains species classified in Plethospira Ulrich in Ulrich and Scofield, 1897, Hormotoma Salter, 1859, and Turritoma Ulrich in Ulrich and Scofield, 1897 (Figures 8 and 20, nodes 7 and 109). Workers have classified the above genera in the Murchisoniina, so I refer to their clade as the "murchisoniinae" hereafter. Synapomorphies include a bilineate peripheral band with two moderately strong, rounded threads, a thickened and reflected inner margin, a relatively shallow, continuously curving sinus, and high translation that increases over ontogeny. The other clade includes species clas? sified in genera, such as Ophileta Vanuxem, 1842, Ceratopea Ulrich, 1911, and Lecanospira Butts, 1926 (Figures 8-11, nodes 2, 11, 37). Previous workers provided no consensus about the superfamilial classifications of these genera: Knight et al. (1960) considered Ophileta to be a pleurotomarioid, Cer? atopea to be an euomphaloid, and Lecanospira to be a macluri- toid; Yochelson (1973, 1984) considered all three to be pleuro? tomarioids, and N.J. Morris and Cleevely (1981; also Ulrich and Scofield, 1897; Koken, 1925; Wenz, 1938) considered all three to be euomphaloids. The phylogeny suggested in Figures 8-19 best matches the phylogenetic model of N.J. Morris and Cleevely, so it seems most appropriate to designate the second clade the "euomphalinae." There also is no evidence suggest? ing that the Mesozoic Pleurotomaria Defrance, 1824, evolved within this clade rather than from the "murchisoniinae"; in fact, as discussed below, the opposite seems more likely. "Eu? omphalinae" synapomorphies include strong, flange-like monolineate peripheral bands that curve abapically, an inner margin that projects away from the coiling axis, curved bases, and circumumbilical thickenings. I. "EUOMPHALINAES" There are several "euomphalinae" subclades, including three that approximate traditional definitions of the Raphistomatidae, Macluritoidea, and Euomphaloidea. The "euomphalinae" also include most of the putative trochoids of the early Paleozoic. As noted above, the "euomphalinae" clade defined herein is very similar to the Euomphaloidea sensu N.J. Morris and Cleevely (1981). Those authors suggested that the Euompha? loidea included two basic clades, the Ophiletidae and the Helic- otomidae (with two other families, the Euomphalidae and the Omphalotrochidae, considered to have evolved from helicoto- mids). This analysis suggests that most of the taxa assigned by N.J. Morris and Cleevely (1981) to the Schizopea group of the Ophiletidae (e.g., Schizopea, Ophileta, Ceratopea, Dirhacho? pea) represent a paraphylum relative to all later gastropods: Ophileta species (e.g., O. supraplana Ulrich and Bridge in Ul? rich et al., 1930, and O. complanata (Miller, 1889)) are plesio? morphic relative to later-appearing "euomphalinae," Dirhacho? pea species are plesiomorphic relative to the "murchisoniinae," and Schizopea is plesiomorphic relative to all other "archaeo? gastropods." 1.1. "Ophiletoids" Most of the genera assigned by N.J. Morris and Cleevely (1981) to two other ophiletid groups (e.g., the Lecanospira and Lytospira groups) represent a single clade of early "euomphali? nae" (Figure 9, nodes 11-24). The earliest member of this clade is the type species of the genus Ophileta, so I refer to the clade as the "ophiletoids." "Ophiletoid" synapomorphies include a very lenticular aperture, the loss of the swelling at the base of the alveozone (i.e., left ramp), and a very strong peripheral band. More-derived species (e.g., the Lecanospira compacta clade) share a very narrow, sharp monolineate peripheral band that is near the top of the whorl, a concave right ramp and slightly ultra-dextral coiling that give specimens a "bowl" shape, and a flat "base" that is perpendicular to the coiling axis. Septation is plesiomorphic to "archaeogastropods," but it is es? pecially prominent in many "ophiletoids." Some derived spe? cies assigned to the genus Lytospira display a long channel 22 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY CI: 0.533 RI: 0.583 Figure 8, I?I Ophileta 1 ?' supraplana (see Figure 8) FAMILY ] Ophiletidae |Helicotomidae IMacluritidae ? Ophileta complanata I?' Lecanospira compacta H Lecanospira nereine I I Barnesella llecanospiroides | 1 Malayaspira rugosa ? Maclurina lannulata I I Malayaspira hintzei I?I Rossospira harrisae < ^ 3 ^ ? Ecculiomphalus bucklandi Barnesella measuresae I I Lytospira yochelsoni I?- Lytospira I?I gerrula Lytospira r ^ ^ ^ T J 'norvesica ?*&&' I Inorvegica Ophiletina I aff. O. sublaxa {sensu Rohr 1988) [ Lytospira angelini | Lytospira subrotunda NUMBER 88 23 FIGURE 9 (opposite).?Relationships among the "ophiletoids." For abbrevia? tions, see legend to Figure 7. Node 11 ("Euomphalinae"), strong BC (91); thin PI (103); very low E (121). Node 12 ("Ophiletoids"), strong ML (29); LR flat? ter than RR (51, 52, 72); long ramps (55, 56); loss of LR swelling (73). Node 13 {Lecanospira compacta clade), PB width = 10? (20); sharp ML (28); weak lunulae (39); PB on top of aperture = 0? (48); extremely narrow aperture (58, 59); weak RR swelling (61); IM 1 to CA (98); low ultradextral T (126). Node 14 {Barnesella llecanospiroides clade), sinus angle = 45? (3, 4); juvenile GL stronger (16); V-shaped lunulae (38); ACh lost (45); asymmetric aperture (LR contracted) (54-56); RR swelling dulls over ontogeny (62). Node 15 ("Eccu- liomphalids"), strong lunulae (39); P = 10? (48); asymmetric ramp shapes (RR more convex) (52, 53, 72); narrow aperture (58, 59); RR becoming shorter and rounder over ontogeny (63); BC lost (89); curved IM (95); IM becoming thicker and rounder over ontogeny (97); IM nearly 1 to CA (98). Node 16, left side of sinus sharper than right (2^1); weakly imbricated GL (17); LR wider than RR (57-59); RRC and LRC present (64, 75). Node 17, moderately broad left side of aperture (59); small size (141). Node 18, basal GL strength same as rest of shell (18); RR swelling acute through ontogeny (62); IM 1 to CA (98); open coiling (123). Node 19 ("Lytospirids"), sinus curvature continuous (9, 10); flat RR (52); projecting, peg-like BC beneath outer margin (90, 91, 93). Node 20, very narrow aperture (58, 59); RR swelling becoming acute over ontogeny (62); open coiling (123); carrier-shell scars (140). Node 21, PB width = 05? (20); PB 1 to aperture (42). Node 22, (Ophiletina cf. O. sublaxa clade), sinus angle -30? on right side (3) with little curve on right side of sinus (9). Node 23, left side of sinus sharper than right (2-4); sinus angle -50? on left side (4); left angle of sinus sharper (6-8); extremely narrow aperture (58, 59); extremely low E (121); open coiling (123). Node 24 {Lytospira angelini clade), no ontogenetic change in GL strength (16); concentric lunulae (38); convex RR (52); symmetric ramp widths (54-56). along the base (see Rohr, 1993). This likely is a site of muscle attachment, which suggests that the base of "ophiletoid" spe? cies is homologous with the inner margin of more typical gas? tropods. Although Rohr (1993) described this channel as unique to Lytospira, similar channels exist on the columellas of species classified in Ceratopea, Pararaphistoma, and Pachys- trophia. Thus, the columellar channel seems to have been a re? curring, polyphyletic character among the "euomphalinae." A noteworthy feature in "ophiletoid" evolution is the parallel development of open-coiling in the "ecculiomphalids" (Figure 9, nodes 15-18) and "lytospirids" (Figure 9, nodes 19-24). This shell form evolved only infrequently after the Early Or? dovician, but it appeared many times among Early Ordovician "euomphalinae." The open-coiled forms illustrated in Figure 9 almost certainly were sessile filter-feeders, as the animals would not have been able to effectively balance their shells (Yochelson, 1971; Linsley, 1977; N.J. Morris and Cleevely, 1981). This also has been considered the likely mode of life for nearly planispiral but close-coiled (i.e., coiled with whorls in contact) gastropods, such as those that occupy the base of the "ophiletoid" cladogram (Linsley, 1978). By deriving these more specialized sessile morphologies from the close-coiled forms more than once, these results support the idea that sessile filter feeding first appeared early in "ophiletoid" evolution. "Lytospirid" species bear external shell scars suggesting that they were carrier shells, i.e., small shells, shell fragments, or pebbles were cemented onto the side of the shell as the animal grew (Rohr, 1993; see Linsley and Yochelson, 1973). The one exception is the slightly problematic species Ophiletina cf. O. sublaxa (sensu Rohr, 1988, not Ulrich and Scofield, 1897), which possesses a broad frill in the same position as the shell scars on Lytospira species. Linsley and Yochelson (1973) sug? gested that attached shells and large shell frills might serve a similar functional role, i.e., serving as a support or a "snow- shoe" on soft substrates (see also Linsley et al., 1978). In addi? tion, early Lytospira species possess a moderately strong, peg? like (i.e., with a parallel surface) carina in nearly the same loca? tion (at the alveozone-base intersection), which is just below the scars. Therefore, it seems plausible that the frill is function? ally homologous with shell carrying and phylogenetically ho? mologous with the carina of Lytospira. There has been little discussion of the phylogenetic relation? ships among these snails. In general, the analysis supports the suggestion that Lecanospira and its relatives are not closely re? lated to macluritoids (Linsley and Kier, 1984; Yochelson, 1984; contra Knight et al., 1960). It also supports the idea that Lecanospira and Barnesella Bridge and Cloud, 1947, are closely related (node 14; Bridge and Cloud, 1947; Knight et al., 1960). Rohr (1994) suggested a close relationship between Rossospira Rohr and Malayaspira, which this analysis also supports; however, this analyses' suggestion that Eccu- liomphalus Portlock evolved from that clade (see node 18) ap? pears to be novel. Finally, there are Silurian and Devonian spe? cies assigned to Lytospira (e.g., Horny, 1992a), but this study suggests that those species belong to the "raphistomatid" clade (see below). Thus, it appears that the "ophiletoids" were en? tirely extinct by the Early Silurian. 1.2. "Macluritoids" This analysis indicates that Prohelicotoma, Macluritella, Tei? ichispira, Monitorella Rohr, 1994, Maclurites, and Palliseria Wilson, 1924, represent a subclade of "euomphalinae" (Figure 10, node 25). "Macluritoid" synapomorphies include a straight, shallow sinus with a weak monolineate peripheral band, exag? gerated basal growth lines, a nearly round aperture, and nearly planispiral coiling that opens in the adult whorls. Many of these characters are apparent in the juvenile whorls of species classified as Macluritella or Teiichispira, which produced char? acters typical of more-derived macluritids later in ontogeny. More-derived species (e.g., nodes 27-36) share distinctive on? togenetic changes involving differential expansion of the base and left side, counter-clockwise rotation of the aperture, and ul? tra-dextral coiling. Still more-derived species (e.g., nodes 31-36) have narrow sinuses and complete filling of juvenile whorls instead of septa. Additional synapomorphies not used in this analysis are as? sociated with the calcareous opercula of "macluritoid" species (Yochelson, 1975). Opercula associated with Teiichispira spe? cies are fibrous, horn-shaped structures, which include knob? like muscle attachments on later species. The opercula of more- derived species (i.e., Maclurites, Monitorella, and possibly 24 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY CI: 0.523 RI: 0.732 Figure 8 Node 2 FAMILY Macluritidae Ophiletidae Helicotomidae Prohelicotoma Q uniangulata (see Figure 8) Macluritella H stantoni Teiichispira I I loceana I?I Teiichispira '?'odenvillensis Monitorella I I auricula Teiichispira I kobayashi "Eccyliopterus I ornatus" Mitrospira | ] longwelli Palliseria robusta I?- Teiichispira I?I sylpha I. ? Maclurites magna Maclurites expansa Maclurites bigsbyi Maclurina logani . Maclurites I I sedgewicki | Maclurina manitobensis NUMBER 88 25 FIGURE 10 (opposite).?Relationships among the "macluritoids." For abbrevia? tions, see legend to Figure 7. Node 25 ("Macluritoids"), sinus angle - 30? (3, 4); sinus curve nearly straight (9, 10); PB width = 10? (20); ML becoming weaker over ontogeny (30); weak lunulae (39); p = 40? (48); globular ramps (52, 72); moderately long ramps (55, 56); IM nearly 1 to CA (98); moderately high E (121). Node 26, sinus angle = 20? (3, 4); weak GL (15); PB width = 05? (20); PB on top of aperture = 0? (48); counter-clockwise rotation of aperture over ontogeny (49); RR becoming more concave over ontogeny (53); IM 1 to CA (98); low ultradextral T (126). Node 27, extremely narrow aperture (58, 59); differential ontogenetic expansion of LR (86, 122). Node 28, loss of RR and LR swellings (60, 73); convex LR (72); base projected posteriorly -10? (117); straight base (120). Node 29 {Monitorella auricula clade), PB width = 10? (20); round ML (28); p = -10? (48); IM\base angle - 75? (94); weakly curved IM (95). Node 30 {Teiichispira kobayashi clade), PB 1 to aperture (42); moderately long RR and LR (55, 56); IM nearly 1 to CA (98); high E (121). Node 31 {Teiichispira sylpha clade), narrow sinus (6, 7); isometric ML strength (30); lunulae same strength as GL (39); ACh lost (45); symmetric ramp shapes (52,53, 72); asymmetric ramp lengths (LR longer than RR) (54-56); asymmet? ric aperture (RR wider than LR) (57-59); thickened IM (87); complete infilling of juvenile whorls (128). Node 32 {Palliseria clade), weak ML (29); flat RR (52); BC lost (89); strongly curved IM with very thick middle (87, 95, 96); IM -30? past 1 to CA (98); isometric E (121, 122); moderate ultradextral T (126). Node 33 {Maclurites clade), very narrow sinus (6, 7); fine, sharp GL (15); PB width = 05? (20); PB _L to aperture (42); no ontogenetic rotation of aperture (49); asymmetric ramp shapes (RR more convex) (52, 53, 72); base projected posteriorly -20? (117); high E (121). Node 34 {Maclurites bigsbyi clade), weak ML (29); flat RR (52); moderately long RR (55); narrow right side of aperture (58); no differential expansion of left side (86, 121, 122); straight IM (95); IM 1 to CA (98); base projected posteriorly -10? (117); ornate base (129). Node 35, very weak ML (29); stronger ornament (131). Node 36, omate LR (129). Palliseria; but see Rohr, 1994) are shield-like rather than horn? like, but retain the knob. The cladogram suggests that the shield-like operculum is derived from the horn-like one. Unfor? tunately, there are too few exact shell-operculum associations to use opercular characters, but more detailed analyses of ma? cluritoid opercula and associated shells could be used to test these results in the future. Although Knight (1952) and Erwin (1990b) previously sug? gested a close relationship between euomphaloids and macluri? toids, this study proposes a very different model of "macluri? toid" evolution than those authors suggested. Knight (1952) and others (e.g., Wangberg-Eriksson, 1979; Runnegar, 1981, 1996; Linsley and Kier, 1984) assumed that Maclurites and re? lated species evolved from onychochilids and, therefore, di? verged from the main gastropod clade during the Early to Mid? dle Cambrian. This analysis, however, suggests that onychochilids and pre-Ordovician species assigned to the Ma- cluritidae (e.g., Macluritella! walcotti Yochelson and Stinch- comb, 1987, and Euomphalopsis involuta Ulrich and Bridge in Ulrich et al., 1930) are related only distantly to the "macluri? toids" and other gastropods (see Figure 7, nodes E, G-J). The results of this study corroborate the predictions of P.J. Morris (1991) and, in part, McLean (1981), Linsley and Kier (1984), and Yochelson (1984), but the traditional models merit further investigation. This is best done by examining whether a more traditional estimate of phylogeny produces a signifi? cantly longer tree than the one accepted herein. The shortest tree linking the two clades assumes that the Middle Ordovi? cian Palliseria and Maclurites are the least derived members of the clade and that the earliest species (e.g., Macluritella and early Teiichispira) are the most derived. Sampling of "maclu? ritoid" species (especially of Palliseria and Maclurites) is dense, so stratigraphic data reject at high levels any tree that considers the morphologic intermediates to be phylogenetic intermediates (see Appendix 3). Accordingly, I used the short? est non-rejectable tree that linked the "macluritoids" with members of the Onychochilidae (which placed "macluritoids" outside the Gastropoda) and compared this tree to the accepted tree (including the outgroups and "macluritoid" species that appeared through the Early Arenig (Early Ordovician)). To test whether a less-parsimonious tree is significantly longer than a more-parsimonious tree, Alroy (pers. comm., 1994) proposed bootstrapping character-states matrices to determine whether random sections of the matrix suggested that one esti? mate of phylogeny is as short as another. Bootstrapping is ap? propriate only when resampling independent units. This as? sumption is violated by the additive coding schemes used herein (see O'Grady et al., 1989); for example, it is impossible to know that some species have a particular type of peripheral band without simultaneously knowing that those species have a peripheral band. Thus, if we sampled character 21 (the pres? ence or absence of peripheral lira on the peripheral band), then we know that we have sampled character 19 (the presence or absence of the peripheral band itself). Similarly, if one ran? domly samples a character describing the left side of some fea? ture, one should know that the species has a right side of that same feature. As an alternative, I bootstrapped the branch lengths on the two different cladograms. For two trees with the same taxa and characters, greater branch lengths indicate some combination of greater homoplasy and poorer sampling of evolutionary transitions (i.e., with poorer sampling, more mor? phologic change happens between sampled species). This means that bootstrapping branch-lengths tests whether one tree requires significantly more homoplasy and/or worse sam? pling than another. None of the 1000 bootstrap analyses of the accepted tree are as long as the alternative tree, nor are any of the 1000 bootstrap analyses of the alternative tree as short as the accepted tree. This apparently is because the putative homologies between the "macluritoids" and onychochilids exist largely in late ap? pearing "macluritoids," not in the early appearing species. Overall, these data suggest that the traditional hypothesis is sig? nificantly less parsimonious than the tree presented herein and also strongly implies that "macluritoids" represent a very re? stricted clade of Ordovician gastropods that are not closely re? lated to other ultra-dextral molluscs of the early Paleozoic. Thus, contrary to Runnegar (1981, 1996), there is neither a morphologic nor a stratigraphic progression linking ony? chochilids and "macluritoids." 26 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY 1.3. "Ceratopeoids" The other major "euomphalinae" subclade (Figures 11-14, nodes 37-69) contains a number of species classified in the genus Ceratopea. This genus has not been used previously to label any higher taxon, but the plesiomorphic nature of Cer? atopea species makes it the most appropriate label for this clade. The earliest "ceratopeoids" (Figure 11) possess deep, strongly curved sinuses, strong abapically hooked peripheral bands (see Figure 11), a sharp, thick basal carina, a crenulated base, and a Lytospira-like channel within the columella. Syna? pomorphies linking later members of the clade include similar ontogenetic changes in aperture orientation, asymmetrical si? nuses, and peripheral bands placed asymmetrically onto the right ramp. "Ceratopeoids" feature two major subclades, the "raphisto? matids" and "helicotomids," and one smaller subclade. This small subclade includes species that have been classified under a variety of generic names, including Bridgeites Flower, 1968a, and Bridgeina Flower, 1968a (Figure 11, nodes 38, 39). The "bridgeitids" are diagnosed by a sharply hooked peripheral band, strongly exaggerated basal and juvenile growth lines, and an extremely lenticular aperture. More-derived species feature nearly planispiral shells with open-coiling in adult stages. This short-lived subclade was relatively diverse during the early Arenig (Early Ordovician) but apparently became extinct shortly thereafter. 1.3.1. "RAPHISTOMATIDS" One of the major "ceratopeoid" clades includes a number of taxa that workers from the turn of the century (e.g., Koken, 1898, 1925) associated with the Raphistomatidae. Accord? ingly, this clade is labeled the "raphistomatids" (Figures 11-14, nodes 42-68). The "raphistomatid" clade is much broader than the most recent definitions of the family (i.e., Wenz, 1938; Knight et al., 1960), but it is similar to the defini? tions presented herein. Synapomorphies include a very high si? nus angle, pronounced growth lines on juvenile whorls, and a crenulated base. 1.3.1.1. "LESUEURILLINES".?Species classified in the genus Lesueurilla dominate one of the major "raphistomatid" sub? clades (Figure 12, nodes 44-54). That genus name has never been used to label a suprageneric taxon, but it is the most ap? propriate name for this particular clade. "Lesueurilline" syna? pomorphies include very strong, hooked peripheral bands and a distinctive sigma-shaped lunulae (see Figure 4F) . An interest? ing feature of "lesueurilline" evolution is that nearly planispiral coiling is the primitive condition both ontogenetically and phy- logenetically. Juvenile and early forms possess Lesueurilla-like morphologies, whereas adult forms (among derived species) possess more lenticular morphologies (e.g., see Figure 3). The "lesueurillines" also include the earliest clade diagnosed by a slit (the L. prima clade, Figure 12, node 48). The slit of these species is not a distinct feature, but instead it is an extension of the sinus where the left and right halves run parallel to each other near the apex. The morphogenetic development of the slit from the sinus is most obvious on species such as Pararaphis? toma qualteriata (Schlotheim) and P. schmidti (Koken, 1925) on which the slit appears later in ontogeny. This analysis supports Yochelson's (1982, 1984) proposition that Lesueurilla and Climacoraphistoma are close relatives. All of the characters that Yochelson cited as linking the two taxa are synapomorphies of relevant nodes, and there are additional synapomorphies that Yochelson did not list, such as the nar? rower, somewhat asymmetrically shaped sinus and the loss of the columellar channel. Another pertinent implication is that Eccyliopterus Remele, 1888, is closely related to Eccu- liomphalus Portlock, 1843 (Knight et al., 1960). Eccu- liomphalus (or at least the type species of that genus, E. buck- landi Portlock, 1843) apparently is an "ophiletoid" (see discussion above), whereas Eccyliopterus represents a sister group to derived Lesueurilla plus the clade of Climacoraphis? toma + Pararaphistoma. A more tentative suggestion by Yoch? elson and Copeland (1974) is that Ceratopea and Pararaphis? toma are synonymous. This analysis implies that the similarities between Pararaphistoma and Ceratopea are due to reversals during the evolution of Climacoraphistoma and Pararaph is toma. The second "lesueurilline" subclade includes species classi? fied as Raphistoma Hall, 1847, and Scalites Emmons, 1842. Synapomorphies of the clade include increased curvature and differential expansion of the lower portion of the shell, which increased both the whorl expansion and translation rates (sensu Raup, 1966) and provided a more oval aperture shape. Some later species (e.g., R. striata Emmons, 1842, R. peracuta Ulrich and Scofield, 1897, and S. katoi Kobayashi, 1934) possess si? nuses with a distorted right half and a sigmoidal left half (Fig? ure 13, nodes 55-59). "Lesueurillines" were relatively diverse during the late Arenig through the Llandeilo (Middle Ordovician). Although a few species survived into the Caradoc (Middle to Late Or? dovician), there are no known Silurian "lesueurillines." It also FIGURE 11 (opposite).?Relationships among the "ceratopeoids." For abbrevia? tions, see legend to Figure 7. Node 11 ("Euomphalinae"), strong BC (91); thin PI (103); very low E (121). Node 37 ("Ceratopeoids"), long ramps (55, 56); weak RR swelling becoming acute over ontogeny (61, 62); flat LR with no swelling (72, 73); thickened IM (87); moderate K (123). Node 38 {Bridgeites clade), P - 50? (48); IM nearly 1 to CA (98); base projected posteriorly -30? (117); low K. (123); curvature decreases over ontogeny (124); nearly planispi? ral coiling (126). Node 39, ACh lost (45); asymmetric ramp shapes (RR rounder (51-53)), ramp lengths (LR contracted (54-56)), and aperture breadth (RR wider than LR (57-59)); strong RR swelling that dulls over ontogeny (61, 62); IM 1 to CA (98); IM channel lost (102). Node 40 {Ceratopea Vaurentia clade), strong basal GL (18); strong, flange-like ML (28, 29); asymmetric ramp shapes (RR rounder) (51-53); pronounced, channeled BC (90); IM\base angle = 105? (94); base projected posteriorly -30? (117); moderate E (121). Node 41 ("Helicotomids"), fine, sharp GL (15); clockwise rotation of aperture over ontogeny (49); ontogenetic increase in T (127). Node 42 ("Raphistomatids"), juvenile GL stronger (16); counter-clockwise rotation of aperture over ontog? eny (49); elongated RR (54-56); RR swelling dulls over ontogeny (62). NUMBER 88 27 CI: 0.525 RI: 0.451 Figure 8, Node 2 FAMILY Ophiletidae EmiH Euomphalidae Helicotomidae ? T3 Ophileta supraplana (see Figure 8) Bridgeites supraconvexa ? Bridgeites Idisjuncta ? ? Ceratopea llaurentia Ceratopea unguis (go to Node 70) ? Helicotoma medfraensis Palaeomphalus giganteus (go to Node 43) 28 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY CI: 0.484 RI: 0.623 Figure 9 Node 17 FAMILY Ophiletidae Helicotomidae 5? Palaeomphalus r LLJ giganteus (see Figures 11,13) ? ? ? I I Lesueurilla V declivis I I Eccyliopterus regularis M Eccyliopterus ^ Iprinceps I?I Eccyliopterus alatus Eccyliopterus owenanus Lesueurilla prima I?I Eccyliopterus '?' buderbacki LesueuriEa infundibuk Eccyliopterus beloitensis Lesueurilb marginah Lesueurilla scotica Lesueurilb bipatelhre ? Climacoraphistoma i vaginaft _z Cfim acoraphistoma damesi I Pararaphistoma qualteriata \Pararaphistoma schmidn NUMBER 88 29 FIGURE 12 (opposite).?Relationships among the "lesueurillines." For abbrevi? ations, see legend to Figure 7. Node 43 {Palaeomphalus giganteus clade), sinus angle = 50? (3, 4); stronger juvenile GL (16); sigma-shaped lunulae (38); loss of RR swelling (60); curved IM (95). Node 44 ("Lesueurillines"), PB width = 10? (20); p - 30?-40? (48); asymmetric ramp shapes (LR rounder than RR) (51, 52, 72); IM thickness same as rest of shell (87); IM\base angle = 90? (94); IM 60? off parallel to CA (98); nearly planispiral coiling (126). Node 45 {Eccyliopterus regularis clade), sinus angle = 40? (3, 4); very strong ML (29); no ontogenetic rotation of aperture (49); PI thickness same as rest of shell (103); extremely low K (123). Node 46, asymmetric aperture (LR contracted) (54-56). Node 47, PB width * 05? (20); extremely strong ML (29); asymmetric aperture (RR wider than LR) (57-59); thickened IM (87); IM at -45? to CA (98). Node 48 {Lesueurilla prima clade), left side of sinus more obtuse (2^1); left side of sinus narrower (5-7); slit present, leaving weak lunulae (34, 39); PB partially on RR (43); IM channel lost (102). Node 49 {Eccyliopterus louder- backi clade), PB slightly raised relative to whorl (33); RR becoming more con? cave over ontogeny (53); BC lost (89); curved base (120); high E (121); large size (141). Node 50, wide left side of sinus (7); no ontogenetic rotation of aper? ture (49); very high E (121); isometric T (127). Node 51 {Lesueurilla margin- alis clade), IM at -45? to CA (98); isometric T (127). Node 52, fine, sharp GL (15); no ontogenetic change in GL strength (16); ACh lost (45); flat LR (72). Node 53 {Climacoraphistoma clade), high rotation of aperture (50); asymmet? ric ramp shapes (RR more convex) (52, 53, 72); IM at -45? to CA (98). Node 54, basal GL strength same as rest of shell (18); symmetric ramp shapes (51); IM 30? off parallel to CA (98); straight base (120); low T (126). bears noting that the two basic "lesueurilline" subclades gen? erally were restricted to different biogeographic realms from their origins in the middle Arenig (Early Ordovician) through the Llandeilo (Middle Ordovician). The Lesueurilla clade (Figure 12) occurs predominantly in the Baltoscandian faunas of Europe, whereas early members of the Scalites clade (Fig? ure 13) exist predominantly in the early Laurentian fauna of North America. Some geographic overlap between the two clades does occur in Malaysia and western North America during that time, and members of both clades occur in both faunas from the Caradoc through the Ashgill (Middle to Late Ordovician). 1.3.1.2. "HOLOPEINES".?The second "raphistomatid" sub? clade comprises species classified in Raphistomina Salter, 1859, Pachystrophia Perner, 1903, Sinutropis Perner, 1903, and Holopea Hall, 1847 (Figures 13, 14, nodes 60-69). This clade is diagnosed primitively by a completely U-shaped sinus (nodes 60-69), although some derived species have no sinus. Pachystrophia includes the least derived members of a clade diagnosed by the loss of a peripheral band (Figure 14, nodes 62-69). Later species (i.e., Pachystrophia gotlandica (Lind? strom, 1884) and Lytospira subuloides Barrande in Perner, 1903) (Figure 14, nodes 68, 69) include the only open-coiled, nearly planispiral species known from the Silurian. In addi? tion, Silurian species placed by Knight et al. (1960) in the fam? ily Sinuopeidae (e.g., Horiostomella Perner, 1903, and Sellinema Perner, 1903) likely also belong to this clade. The status of the post-Silurian members of that family are not known, but it is much more likely that they are "holopeines" than it is that they are related to the Late Cambrian-earliest Ordovician Sinuopea species. Species assigned to the genus Holopea are a particularly con? spicuous development within the "holopeines" (Figure 14, nodes 64, 65). The earliest Holopea species are nearly identical to contemporaneous Pachystrophia species, save for the ab? sence of the sinus (with the absence of a peripheral band being a synapomorphy that links Pachystrophia and Holopea). The trochiform shell typically associated with the genus is shared among derived Holopea species (node 65). Holopea is an im? portant genus when discussing gastropod phylogeny because its family (the Holopeidae) represents the earliest putative members of the Trochoidea in some classification schemes (e.g., Knight et al., 1960). None of the other genera typically assigned to the Holopeidae, however, seem to have been close relatives of Holopea. Although the genus is reported through the Devonian (Knight et al., 1960), this analysis did not find any post-Ordovician members of the H. insignis clade. This makes it seem unlikely that Holopea represents an early tro? choid. Other phylogenetic scenarios concerning the Trochoidea are discussed below. Some general phylogenetic proposals concerning the genus Pachystrophia are not supported here. Wenz (1938) considered Pachystrophia to be a close relative of "ophiletoid" genera such as Ecculiomphalus and Lytospira. Also, some "hol- opeides" from the Silurian (at least one of which extends into the early Devonian; see Homy 1992a) have been classified in the genus Lytospira. A very different suggestion was made by Knight et al. (1960), who considered Pachystrophia to be a jun? ior synonym of Lesueurilla. This analysis contradicts all of these ideas, suggesting instead that Pachystrophia is distantly related to the "ophiletoids" and that Lesueurilla and Pachystro? phia have closer relatives than one another. 1.3.2. "HELICOTOMIDS" The other major "ceratopeoid" subclade includes many of the traditional euomphaloid genera (sensu Knight et al., 1960), plus most of the early Paleozoic taxa classified in the Tro? choidea (also sensu Knight et al., 1960; Tracey et al., 1993). The earliest species of the clade agree well with the definition of the Helicotomidae (sensu Knight et al., 1960, not Wenz, 1938), so I refer to the clade as the "helicotomids." The clade presented herein (Figures 15-19, nodes 70-108) is very similar to N.J. Morris and Cleevely's (1981) genus-level definition of the Euomphalidae + Helicotomidae clade. "Helicotomid" syna? pomorphies include a sigmoidal aperture shape with the base of the aperture projected in front of the inner margin, a broadly expanded (i.e., nearly round or square instead of lenticular) ap? erture, a shallow, weakly curved sinus, a weak monolineate pe? ripheral band, a weak thread-like basal carina, and strong chan? nels beneath the right and left carina. This analysis also corroborates N.J. Morris and Cleevely's (1981) opinion that derived "euomphalinae," such as Po- leumita Clarke and Ruedemann, evolved from early members of the Helicotomidae. Those authors divided the Euompha- 30 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY CI: 0.538 RI: 0.554 Figure 11 Node 40 FAMILY Raphistomatidae Planitrochidae Helicotomidae ESO Euomphalidae Pararaphistoma [~] lemon i (see Figure 11) Palaeomphalus [ j giganteus (see Figures 11,12) I I Scalites katoi Raphistoma teller ensis Raphistoma peracuta Scalites angulatus Raphistoma striata Helicotoma gubanovi [ ~| Raphistomina lapicida Raphistomina aperta Raphistomina fissurata Raphistomina rugata Pachystrophia I?I devexa (go to Node 62) NUMBER 88 31 FIGURE 13 (opposite).?Relationships among the "scalitines" and "holoepeids." For abbreviations, see legend to Figure 7. Node 42 ("Raphisto? matids"), juvenile GL stronger (16); counter-clockwise rotation of aperture over ontogeny (49); elongated RR (54-56); RR swelling dulls over ontogeny (62). Node 43 {Palaeomphalus giganteus clade), sinus angle = 50? (3, 4); stronger juvenile GL (16); sigma-shaped lunulae (38); loss of RR swelling (60); curved IM (95). Node 55 ("Scalitines"), basal GL strength same as rest of shell (18); PB width = 10? (20); very strong ML (29); symmetric ramp shapes (51); asymmetric ramp lengths (right contracted) (54-56); very thick IM (87); IM\base angle - 75? (94); IM -15? off parallel to CA (98); IM channel lost (102). Node 56, wrinkled right side of sinus (?) (11); ACh lost (45). Node 57, squared ML (28); very short RR (55); asymmetric ramp projection (RR higher) (57-59); whole aperture inclined -20? (109, 110); noncrenulated base (120); high E (121). Node 58, U-shaped left side of sinus (10); BL present (21); PB slightly raised relative to whorl (33); slit present (34); convex RR (52); strong, nonprojecting BC (91). Node 59 {Raphistoma tellerensis clade), U-shaped left side of sinus (10). Node 60 ("Holopeids"), sinus angle = 25? (3, 4); U-shaped sinus (9, 10); no ontogenetic change in GL strength (16); basal GL strength same as rest of shell (18); [J = 70? (48); asymmetric ramp shapes (RR rounder) (51-53); symmetric ramp lengths (54-56); RRC present (64); strong, non- projecting BC (91); IM 30? off parallel to CA (98); low E and low K (121, 123); isometric T (127). Node 61, PB 1 to aperture (42); moderately long ramps (55, 56); extremely narrow aperture (57, 58); IM thickness same as rest of shell (87); nonchanneled, weak BC (90, 91); IM\base angle = 105? (94); IM 15? off parallel to CA (98); curved base (120). loidea into two early families, the Ophiletidae and the Helicot? omidae, which they considered to be sister clades. This analy? sis suggests that N.J. Morris and Cleevely's definition of the Ophiletidae (which is very close to the "euomphalinae" as de? fined herein, but without species from the Middle Ordovician and later) is paraphyletic relative to their Helicotomidae. 1.3.2.1. "OPHILETININES".?Two "helicotomid" subclades evolved by the Middle Ordovician. The first of these includes species assigned to Helicotoma Salter, 1859, Palaeomphalus Koken, 1925, and Ophiletina Ulrich and Scofield, 1897 (Figure 15, nodes 74-80). As I previously labeled the more inclusive clade the "helicotomids," I designate this clade the "ophiletin- ines." "Ophiletinine" synapomorphies include a moderately wide, flange-like peripheral band that curves abapically and overlies a weak channel, a wide, obtuse left carina that overlies a shallow channel, and very low translation. No known species matches the hypothetical common ancestor of the "ophiletin- ines," although it likely was most similar to Oriostoma bro- midensis Rohr and Johns, 1992, of the Early Caradoc. Synapo? morphies uniting the O. bromidensis clade (Figure 15, nodes 75, 76) include a very shallow, very narrow sinus and a strong, sharp basal carina. Ophiletina species (node 76) share a peg? like left carina and a bilineate peripheral band. These species also appear to have had partially calcitic shells, although this was not used in the cladistic analysis. Synapomorphies uniting the Helicotoma tennesseensis clade (Figure 15, nodes 77-80) include a rounded right carina that weakens over ontogeny, an expanded, rounded base, very faint growth lines, an unusually strong, hooked peripheral band, and extreme swelling of the left carina. A disc-like paucispiral operculum is associated with one Helicotoma species (Yochel? son, 1966a). This operculum is very different from the horn- shaped opercula associated with early "helicotomids" and other early "ceratopeoids" (e.g., Ceratopea unguis Yochelson and Bridge, 1957; see Yochelson and Wise, 1972). If the phy? logeny presented herein is reasonably accurate, then a pau? cispiral, disc-like operculum is derived, but it unfortunately is not known how common that operculum was among the "heli- cotomatid" clade. "Ophiletinines" were moderately diverse from the Middle Ordovician through the Late Ordovician, but this study found no Silurian members of the clade. 1.3.2.2. "EUOMPHALOPTERINES".?The second "helicoto? mid" subclade includes a diverse array of Silurian species that typically have been assigned to the Anomphalidae, Elasmone- matidae, Euomphalopteridae, and Pseudophoridae (Figures 16-19, nodes 81-108). The Ordovician precursors of this clade have been classified in the genus Euomphalopterus Roemer, 1876 (e.g., Euomphalopterus cariniferus Koken, 1925). I cate? gorize the clade as the "euomphalopterines," even though this clade is very different from previous definitions of the Eu? omphalopteridae (e.g., see Wenz, 1938, or Knight et al., 1960). "Euomphalopterine" synapomorphies include a very strong left carina, a shallow V-shaped sinus with a very narrow sharp pe? ripheral band, and very broadly projecting ramps. 1.3.2.2.1. "Anomphalides".?Three major "euomphalopter? ine" subclades arose during the latest Ordovician and Early Silurian from an Euomphalopterus cariniferus-hke ancestor (Figures 16-19, nodes 81-108). One of these subclades (Fig? ure 16, nodes 83-87) consists predominately of species as? signed to the Anomphalidae (e.g., Pycnomphalus Lindstrom, 1884, and Grantlandispira Peel, 1984a). Characters diagnos? ing the "anomphalide" clade include a strong but dull carina on a swollen alveozone, a strong lirum on the inner margin that partially fills the umbilicus, and a U-shaped sinus with an apex near the suture. Among more-derived species (e.g., nodes 86, 87), the umbilicus is entirely filled and the periph? eral band is highly reduced or lost. Kase (1989) noted that some of the earliest species assigned to the Omphalotrochidae (e.g., Middle Devonian species assigned to Labrocuspis Kase) have synapomorphies with "anomphalide" species; however, the majority of omphalotrochids, including Devonian species assigned to Pseudomphalotrochus Blodgett, 1992, appear to share synapomorphies with Straparollus de Montfort, 1810, and Euomphalus Sowerby, 1814 (Erwin, in prep.; Wagner, in prep.). 1.3.2.2.2. "Poleumitides".?The second major "euompha? lopterine" subclade contains species assigned to Poleumita, Euomphalopterus, Centrifugus Bronn, 1834, and (possibly) Spinicharybdis Rohr and Packard, 1982 (Figures 17, 18, nodes 88-96). This clade bears little resemblance to the Poleumitidae of Wenz (1938), which links Poleumita with the problematic Oriostoma Munier-Chalmas, 1876 (see also Boucot and Yoch? elson, 1966). Because the Poleumitidae is an established name, I designate this clade as the "poleumitides." "Poleumitides" ev? idently shared a common ancestor in the Early Silurian that 32 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY CI: 0.627 RI: 0.698 Figure 13 Node 60. FAMILY Euomphalidae Holopeidae Ophiletidae \65[ k S" S' ?^^ T3 P <-i ^ O -^*-o \ 3 Cfc a Co a" utropi Oi' ^3 a n s- '^t C?J *-?. ~*i 0 13 3- s* >T3 gs >-l Pachystrophia I I devexa (see Figure 13) Pachystrophia I I contigua ? Holopea insignis Holopea pyrene l?- Holopea I?I rotunda Holopea symmetrica Pachystrophia I I spiralis H Sinutropis lesthetica Euomphalus Pachystrophia 1 gotlandica Lytospira I I triquestra Lytospira C subuloides NUMBER 88 33 FIGURE 14 (opposite).?Relationships among the derived "holopeids." For abbreviations, see legend to Figure 7. Node 62 ("Pachystrophides"), fine, sharp GL (15); PB lost (19); ACh lost (45); p" = 60? (48); no ontogenetic rotation of aperture (49); very narrow aperture (58, 59); RRC lost (64); convex LR (72); BC lost (89); thickened PI (103). Node 63 {Pachystrophia contigua clade), asymmetric aperture (left side broader) (57-59); IM 15? off parallel to CA (98); whole aperture inclined -20? (109, 110); moderate K (123). Node 64 {Holopea insignis clade), sinus lost (1); asymmetric ramp lengths (RR con? tracted) (54-56); broad left side of aperture (59); IM\base angle = 60? (94); moderate T (126). Node 65, p = 40? (48); symmetric ramp shapes (51-53); very short RR (55); moderately wide right side of aperture (58); very convex LR (72); thin PI (103); whole aperture inclined -30? (109, 110); base projected posteriorly -10? (117); straight base (120); moderate E (121); high T (126); ontogenetic increase in T (127); septation absent (128). Node 66 {Sinutropis lesthetica clade), asymmetric sinus shape (left curve stronger) (8-10); P = 30? (48); asymmetric ramp shapes (LR rounder than RR) (51, 52, 72); long ramps (55, 56); asymmetric aperture (RR wider than LR) (57-59); base projected pos? teriorly -20? (117). Node 67, asymmetric ramp lengths (LR longer than RR) (54-56); very convex LR (72). Node 68 {Pachystrophia gotlandica clade), IM channel present (102). Node 69, sinus angle = 20? (3, 4); narrow sinus (6, 7); weak GL (15); P = 20? (48); moderately long ramps (55, 56); moderately wide aperture (58, 59); IM 1 to CA (98); base projected posteriorly -10? (116, 117); open coiling (123); low ultradextral T (126). was much like E. cariniferus but with a frill-like left carina, slightly imbricated growth lines, ontogenetic weakening of the right carina and peripheral band, and strong increases in whorl convexity over ontogeny. A calcitic shell also diagnoses this clade, although again I did not use this as a character state. In the Euomphalopterus subcarinatus clade (Figure 17, nodes 88-92), the frill becomes strongly developed (see also Linsley et al., 1978). This frill is highly crenulated on species such as E. praetextus (Lindstrom, 1884) or replaced with a series of tubes on others such as E. togatus (Lindstrom, 1884). The lat? ter feature is best developed in Spinicharybdis, although it is possible that this represents a parallelism. Rohr and Packard (1982) commented on similarities between Spinicharybdis and Euomphalopterus, but they did not explicitly suggest that the two genera were related. The elongate, widely spaced tubes of S. wilsoni Rohr and Packard, 1982, are more similar to those seen on species of Hystricoceras Jahn than they are to the short, tightly-spaced tubes of E. togatus. Yochelson (1966b) cited this as evidence that the species since classified as Spini? charybdis are related to Hystricoceras. Spinicharybdis wilsoni shares other synapomorphies with E. togatus, however, includ? ing the shape of the ramps, the development of the sinus, and the complete absence of a peripheral band. There are other species of Spinicharybdis that I could not include in this analy? sis because the only known specimens are too incomplete. These species do show a series of tubes that are very similar to the tube-bearing frill of E. togatus, save that they are much longer; however, there is no evidence of frill-bounding lira on any of these species, which exist on E. togatus and its rela? tives. Only the bases and lower whorls are visible on any of the pertinent specimens, so it is not known if the sinuses and pe? ripheral bands of these specimens are like those of Euompha? lopterus or like those of Hystricoceras. Therefore, it is con? ceivable that Spinicharybdis actually is a "pseudophoride" (see below). Finally, Spinicharybdis also shares features with contempo? rary Straparollus (e.g., S. paveyi Foerste, 1924), and the spac? ing of the tubes on S. wilsoni is similar to the spacing of carrier shell scars of 5". paveyi (see further discussion, below). The long tubes of Spinicharybdis might have served the same func? tional purpose as the agglutinated shells on Straparollus spe? cies (e.g., Linsley and Yochelson, 1973). Therefore, another possibility is that the spines of Spinicharybdis represent a mor? phologic novelty that maintained a functional "homology" with carrier-shell ancestors such as S. paveyi. In this case, the more frill-like tubes of other species represent convergence to? ward a Euomphalopterus-hke morphology. As a similar func? tional interpretation applies to the extended frill of some Eu? omphalopterus species (but not to the short tubular frill of species such as E. togatus; e.g., Linsley et al., 1978), however, it seems as or more likely that the spines and frills represent different adaptations on the same homologies that were func? tional parallelism. Euomphalopterus apparently is the sister taxon of a Silurian clade that includes Poleumita and Centrifugus (Figure 18, nodes 93-96). The earliest known species from this clade, P. alata (Lindstrom, 1884), retains a strong lower ramp carina, but later species possess a wide, dull swelling in this region. Other synapomorphies include a flat, very shallow sinus, a thickened and rounded inner margin, well-developed ornament, and near planispiral coiling with very low curvature. The diag? nosis of Poleumita differs from that of Euomphalus primarily in that Poleumita possesses ornamentation. Excluding orna? ment (which is plesiomorphic above node 94), "poleumitide" species, such as E. walmstedti Lindstrom, 1884, lack any obvi? ous autapomorphies relative to Euomphalus. Thus, the "po? leumitides" likely represent the "euomphalinae" in the truest sense. As noted above, Late Silurian species classified as Straparol? lus possess carrier shell scars. This feature is retained on many Devonian species (e.g., Linsley and Yochelson, 1973) and might be a synapomorphy between these species and early omphalotrochids (pers. obs.). 1.3.2.2.3. "Pseudophorides".?The final "euomphalopter- ine" subclade (Figure 19, nodes 97-108) includes species as? signed to Pseudophorus Meek, 1873, Discordichilus Coss- mann, 1918, Hystricoceras, Siluriphorus Cossmann, 1918, and Streptotrochus Perner, 1903. Previous workers classified many of these species in the Pseudophoroidea (e.g., Knight et al., 1960), so I refer to this clade as the "pseudophorides." The clade also includes the Late Ordovician Euomphalopterus lor- dovicius Longstaff, 1924. "Pseudophoride" synapomorphies include a tangential aperture (i.e., the inclination of the entire aperture rather than just portions of the aperture), a thickened inner margin with little projection relative to the coiling axis that fills the umbilicus, and moderately high translation. The clade initially retains the sharp peripheral band and the sharp 34 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY CI: 0.609 RI: 0.726 Figure 9 Node 18 FAMILY Ophiletidae Helicotomidae Euomphalopteridae Oriostomatidae I?i Ceratopea ? unguis (see Figure 11) I I Lophonema peccatonica Polehemia taneyensis Walcottoma frydai Oriostoma bromidensis I?I Ophiletina sublaxa Ophiletina angularis Helicotoma tennesseensis Helicotoma planulata Helicotoma robinsoni Helicotoma blodgetti Palaeomphalus Igradatus I Helicotoma! ' Girvan sp. I Boucotspira aff. B. fimbriata Euomphalopterus I cariniferus (go to Node 81) NUMBER 88 35 FIGURE 15 (opposite).?Relationships among the "helicotomids." For abbrevi? ations, see legend to Figure 7. Node 70 ("Helicotomids"), sinus angle = 50? (3, 4); moderately strong, round ML (28, 29); asymmetric ramp lengths (RR nar? rower than left) (54-56); asymmetric aperture (right side wider) (57-59); IM\base angle = 90? (94); curved IM (95); base projected posteriorly -20? (117); curved base (120). Node 71, sigmoidal aperture (13); basal GL strength same as rest of shell (18); PB width = 10? (20); PB 1 to aperture (42); strong ACh (46); P = 60? (48); symmetric ramp shapes (51, 52, 72); moderately broad (slightly asymmetric) aperture (58, 59); loss of RR swelling (60); sharp carina on RR and LR (64, 75); IM thickness same as rest of shell (87); IM\base angle - 75? (94); IM channel lost (102); base projected anteriorly -10? (116, 118, 119); low E (121); moderate T (126). Node 72 {Polehemia taneyensis clade), ornament throughout left side of aperture (129). Node 73 {Boucotspira aff. B. fimbriata clade), sinus angle = 30? (3, 4); weak GL (15); V-shaped lunulae (38); P = 50? (48); no ontogenetic rotation of aperture (49); left and right widths of aperture symmetrical (57-59); base projected anteriorly -20? (119); isometric T (127). Node 74 ("Ophiletinines"), weak ACh (46); low K (123); low T (126); septation lost (128). Node 75 {Oriostoma bromidensis clade), nar? row sinus (6, 7); strong GL (15); projecting BC (91); base projected anteriorly -10? (119); small size (141). Node 76, BL present (21); concentric lunulae (38); very strong lunulae (39); p = 50? (48); highly asymmetric ramps (LR nearly twice as long as RR) (55, 56); slightly asymmetric aperture (left side contracted slightly) (57-59); squared ridge-like LRC (76); thickened IM (87); BC beneath outer margin (93); IM\base angle = 105? (94); inclined aperture (109). Node 77 {Helicotoma tennesseensis clade), weak RRC (65); LR swell? ing present with weak LRC (73, 77); dull, thickened BC (90). Node 78 {Palae? omphalus Igradatus clade), PB width = 05? (20); BC lost (89); strongly curved IM (95). Node 79 {Helicotoma planulata clade), asymmetric aperture (LR moderately contracted) (57-59); thickened IM (87); ontogenetic increase in T (127); ornate LR (129). Node 80, ACh lost (45); asymmetric ramp lengths (LR strongly contracted) (54-56); moderately strong RRC (65); strong, squared ridge-like LRC (76, 77). right and left carinae that are common to plesiomorphic "eu? omphalopterines." The left carina, however, becomes promi? nent with a square periphery, whereas the right carina and the peripheral band are strongly reduced or lost in the Siluriphorus gotlandicus clade (Figure 19, node 104). Also, the sinus is re? duced to a shallow kink near the suture, and the sigmoidal shape of the aperture becomes extreme. Among very derived members of the Discordichilus clade (Figure 19, node 106), the peg-like left carina is very weak and obtuse, whereas the ramps become strongly rounded. Among other members of the Pseudophorus clade, the peg-like carina becomes a hood-like frill (Figure 19, node 105). A second "pseudophoride" subclade (Figure 19, node 101) includes species assigned to Streptotrochus and Hystricoceras. This clade's synapomorphies include an inner margin that re? flects around the coiling axis and that is thickened at the top and bottom, a thin parietal inductura, and notable increases in both shell expansion and shell torque over ontogeny. The most derived species of this clade possess a projected, strongly chan? neled left carina (e.g., S. lundgreni (Lindstrom, 1884)), which forms a series of closely connected tubes on Hystricoceras. Knight et al. (1960) considered Raphistomina to be the earli? est member of the Pseudophoroidea, based on the assumption that the peripheral band on Raphistomina species is homolo? gous with the strong lower ramp carina or frill on species of Sil? uriphorus and Pseudophorus. Previous workers had interpreted this band as a peripheral band because it lies in the middle of a prominent sinus (e.g., Ulrich and Scofield, 1897; Wenz, 1938). I follow the latter interpretation, which leaves Raphistomina species without any important "pseudophoride" synapomor? phies; therefore, this analysis contradicts the relationships im? plied by the taxonomy of Knight et al. (1960). Knight et al. also considered Trochomphalus Koken to be a pseudophoroid. These results agree better with that idea, but they suggest that Trochomphalus is a member of the "anomphalide" clade. Knight et al. (1960) assigned both Streptotrochus and Dis? cordichilus to the Microdomatoidea. This analysis supports a close relationship between these two genera, but it also implies that they have closer relatives than each other among the Pseudophoroidea. The analysis includes only one other puta? tive microdomatoid, Daidia Wilson, 1951, but species belong? ing to that genus are considered to be "murchisoniinae" and not at all closely related to the "pseudophorides" (see the "strap- arollinoids" below) N.J. Morris and Cleevely (1981) previously linked the Pseudophoroidea to the Euomphaloidea. Morris and Cleevely, however, thought that the Pseudophoroidea diverged from those species very early, i.e., prior to the divergence of taxa such as Ophiletina and Helicotoma. This analysis suggests that "pseudophorides" actually are highly derived "euomphali? nae," evolved within the "helicotomids." N.J. Morris and Cleevely (1981) also suggested that another taxon, the Trocho- nematoidea, are even more closely related to the Euompha? loidea than are the Pseudophoroidea. My analyses, however, suggest that the Trochonematoidea evolved from "lophos- piroids" (Wagner, 1995a; see also Ulrich and Scofield, 1897; Knight et al., 1960). As discussed below, lophospirids appar? ently evolved from the "murchisoniinae," so this study does not support that part of Morris and Cleevely's phylogenetic scheme. II. "MURCHISON1INAES" The "Murchisoniinae" contain several important taxa, in? cluding the earliest putative apogastropods and most of the early Paleozoic taxa assigned to the Pleurotomarioidea (Fig? ures 20-35, nodes 109-215). The relationships among the ma? jor "murchisoniinae" subclades (Figure 20, node 109) ap? proaches a star phylogeny (sensu Felsenstein, 1985), with several taxa derived from the same ancestor. Although system- atists usually interpret such polytomies as unresolved relation? ships, the analysis suggests that the polytomy represents the real relationship among these taxa. The polytomy includes Hormotoma Isimulatrix (Billings, 1865), which is plesiomor? phic relative to all "murchisoniinae" subclades. This suggests that the long-lived and geographically wide ranging H. Isimul- atrix is ancestral to these clades through separate cladogenetic events (see Hoelzer and Melnick, 1994; Wagner and Erwin, 1995). The earliest members of the major subclades all co-oc? cur with H. Isimulatrix, and it often was difficult for me to de- 36 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY FAMILY Euomphalopteridae Euomphalidae Anomphalidae Holopeidae Planitrochidae Equivocal Figure 15 Node 74 CI: 0.697 RI: 0.743 FIGURE 16.?Relationships among the "anompha- lides." For abbreviations, see legend to Figure 7. Node 81 ("Euomphalopterines"), sinus angle = 20? (3, 4); PB width = 05? (20); P = 40? (48); broad symmetric aperture (58, 59); strong, peg-like frill LRC with channel (76, 78); round thread-like BC becoming weaker over ontogeny (90, 92); IM\base angle = 105? (94); PI thickness same as rest of shell (103). Node 82 ("Poleumitides"), sinus angle = 10? (3, 4); frilled LRC (78). Node 83 ("Anomphalides"), sinus angle = 30? (3,4); rounded ML (28); weak ML (29); concentric lunulae (38); convex RR (52); loss of LRC channel (78); BC lost (89); IM\base angle = 75? (94); IM 30? off parallel to CA (98); strong columellar lira present (100). Node 84, RRC lost (64); convex LR (72). Node 85, PB lost (19); ACh lost (45); P = 30? (48); highly asymmetric ramp lengths (LR very long, RR extremely short) (55, 56); very broad aperture (58, 59); callous-like columellar lira (100, 101). Node 86, sinus angle = 10?; strongly curved IM (95). Node 87, LRC lost (75). &' Euomphalopterus \ J cariniferus (see Figure 15) Euomphalopterus I lordovicius (go to Node 97) Euomphalopterus j j subcarinatus (go to Node 88) Poleumita B alata (go to Node 93) Trochomphalus I Idimidiatus Straporillina j?? cf. S. circe (sensu Rohr 1988) Grantlandispira I christei Pycnomphalus \ ( acutus Pycnomphalus \ j obesus Turbocheilus immaturum NUMBER 88 37 CI: 0.695 RI: 0.646 FAMILY Euomphalopteridae Umi Euomphalidae L = J Uncertain Figure 16 Node 81 FIGURE 17.?Relationships among the "poleumiti? des," part 1. For abbreviations, see legend to Figure 7. Node 82, sinus angle = 10? (3,4); frilled LRC (78). Node : {Euomphalopterus subcarinatus clade), weakly imbricated GL (17); PB fades over ontogeny (30); RR becoming much more con? vex over ontogeny (53); asymmetric aperture (right side broader) (57-59); RRC becoming weaker over ontogeny (67); very strong, chan? neled frill (LRC) (77, 79, 82); very low E and high K (121, 123). Node 89 {Euomphalopterus togatus clade), dull, lump-like ML (28); right half of aper? ture very broad (58); dull, thickened BC beneath IM (90, 93). Node 90, ACh lost (45); very convex RR becoming flatter through ontogeny (52, 53); crenulated LRC frill (81). Node 91 {Euomphalopterus praetextus clade), U-shaped sinus (8, 9); fine, sharp, nonimbricated GL (15, 17); PB lost (19); concentric lunulae (38); strongly curved IM (95). Node 92 {Euomphalopterus frenatus clade), right side of aperture extremely broad (58); LRC frill forming tubes (85); high T (126). tn Euomphalopterus | \subcarinatus (see Figure 16) Euomphalopterus H alatus Euomphalopterus | | togatus Euomphalopterus H praetextus Euompha lopterus M undulans Euomphalopterus frenatus Spinicharybdis I wilsoni _ Poleumita m alata (go to Node 93) 38 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY CI: 0.643 RI: 0.637 Figure 16 Node 81 FAMILY Euomphalidae tfrnm Euomphalopteridae Euomphalopterus | subcarinatus (see Figures 16,17 I?j Poleumita alata (see Figures 16,17) Poleumita discors Poleumita granulosa Centrijugus planorbis Poleumita rugosa Poleumita octavia Euomphalus walmstedti Straparollus bohemicus NUMBER 88 39 FIGURE 18 (opposite).?Relationships among the "poleumitides," part 2. For abbreviations, see legend to Figure 7. Node 82, sinus angle = 10? (3, 4); frilled LRC (78). Node 93 {Poleumita alata clade), sinus nearly absent (3, 4); crenu? lated aperture (12); highly sigmoidal aperture (14); IM\base angle = 90? (94) straight IM (95); thickened middle of IM (96); IM 30? off parallel to CA (98) very thick PI in concentrated strip, projecting in front of aperture (103, 104) strongly curved base with slight posterior projection (116, 118, 120); moder? ately dense ornament on LR and RR (129, 130, 133, 134). Node 94 {Poleumita discors clade), periodically flared ML (32); weak ACh (46); p => 10? (48); long LR (56); weak RRC (65); thick contusion LRC (76); thickened IM (87); BC lost (89); IM\base angle = 75? (94); low T (126); large size (141). Node 95 {Poleumita rugosa clade), extremely strong GL (15); strongly imbricated GL (17); convex RR (52); dull, lump-like LRC (66); convex LR (72); moderate E (121); denser ornament on LR (130). Node 96 {Euomphalus marix clade), non- imbricated GL (17); convex LR (72); LRC lost (75); uniform thickness of IM (96); IM at -45? to CA (98); base projected posteriorly -10? (116, 117); straight base (120); loss of ornament (129, 133). termine whether a specimen should be classified as H. Isimula- trix or as another Hormotoma species (e.g., H. confusa Cullison, 1944, or H. dubia Cullison, 1944). Hormotoma con? fusa and H. dubia, however, share synapomorphies with differ? ent "murchisoniinae" subclades, so I treated them as separate species based on the "phylogenetic species" concept (de Queiroz and Donoghue, 1988; Nixon and Wheeler, 1990). II. 1. "Plethospiroids" One early "murchisoniinae" subclade includes species classi? fied in the Plethospiridae (Figure 20, nodes 110, 111). "Plethos- piroid" synapomorphies include a shallow sinus and a thick? ened, siphonate inner margin. This morphologically novel clade appears to include no members younger than the Early Ordovician, even though Knight et al. (1960) assigned several other genera to the Plethospiridae. Seelya ventricosa Ulrich in Ulrich and Scofield, 1897, belongs to the "plethospiroids," but none of the other Ordovician or Silurian species assigned to Seelya Ulrich in Ulrich and Scofield belong to the clade. An? other putative plethospirid, Diplozone Perner, 1907, appears to be related to Loxonema Phillips (see below). Erwin (1992) sug? gested that the Plethospiridae include the sister taxa of the apo- gastropod-like subulitoids. This analysis suggests that the two clades are closely related, but that other "murchisoniinae" likely are more closely related to subulitids. This is discussed further, below. II.2. "Straparollinoids" Hormotoma dubia represents the stem-member of a moder? ately diverse "murchisoniinae" subclade that includes species classified as Daidia Salter, Haplospira Koken, 1897, and Straparollina Billings, 1865 (Figure 21, nodes 112-118). Pre? vious workers assigned these taxa to the Holopeidae and the Microdomatidae, which in turn were classified in the Tro? choidea (Knight et al., 1960; Tracey et al., 1993). As noted above, the present definitions of those families (and the Paleo? zoic trochoids) is highly polyphyletic, and it is not clear which of these species, if any, represent the precursors of true micro- domatids or trochoids. Accordingly, I label the clade the "strap? arollinoids," which is appropriate given that species assigned to Straparollina represent some of the least derived members of the clade. "Straparollinoid" synapomorphies include a dull monolin? eate peripheral band (which is bilineate on the juvenile whorls of early species and becomes monolineate with age) and a nar? row sinus. Slightly more-derived "straparollinoids," such as Lophospira grandis Butts, 1926 (nodes 114-118), share a very narrow, sharp peripheral band, a nonreflected inner margin, a well-developed left carina, and higher shell torque. Even more- derived species (e.g., Straparollinapelagica Billings, 1865, and more-derived species, nodes 115 and above) possess very narrow sinuses and reduced right sides of the aperture. The most derived "straparollinoids" (e.g., Daidia and Haplospira species, nodes 116, 117) have a weak monolineate peripheral band near attenuated sutures, no sinus, and an inner margin that is entirely contiguous with the previous whorl. Previous workers (e.g., Knight et al., 1960; Erwin, 1988; Tracey et al., 1993) classified Daidia in the Microdomatidae, but the next oldest microdomatids first appear in the Devonian (Blodgett and Johnson, 1992; Blodgett, 1993). The statistical significance of this gap is difficult to assess without data for those Devonian species. It seems unlikely that Daidia is closely related to those species. The "pseudophoride" clade in? cludes Silurian species assigned to the Microdomatoidea (al? beit, to the Elasmonematidae). Overall, it appears more likely that true microdomatoids arose in that clade rather than from "straparollinoids." II.3. "Hormotomoids" Hormotoma confusa Cullison, 1944, is the least derived member of a clade that includes most of the species classified in the Murchisonioidea, Subulitoidea, and Loxonematoidea (Figure 8, node 7; Figures 22-26, nodes 119-154). The earliest species in this clade have been classified in the genus Hormo? toma by numerous authors, so I refer to the clade as the "hor? motomoids." Note that the definition of this clade is much broader than that of the Hormotominae sensu Wenz (1938). The chief synapomorphy uniting the clade is an asymmetrical sinus that is much broader and much sharper on the left side than on the right. The "hormotomoids" represent the earliest species in which the left side of the aperture is more pro? nounced than the right side, which is a feature associated with reduction of the right organs. II.3.1. "SUBULITIDS" The Subulitoidea represent one of the most poorly under? stood but intriguing products of early gastropod evolution. The earliest subulitoids appeared in the Arenig (Early Ordovician), 40 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY CI: 0.582 RI: 0.733 Figure 16 Node 81 FAMILY Euomphalopteridae Elasmonematidae Pseudophoridae Euomphalopterus j | lordovicius (see Figure 16) Streptotrochus ? lamellosus Streptotrochus! I I visbeyensis Streptotrochus I I incisus Streptotrochus I lundgreni Hystricoceras astraciformis Streptotrochus I?I aff. S. incisus Siluriphorus I?I gotlandicus Siluriphorus undulans Discordichilus ! dalli . Discordichilus I?I mollis . D iscordichilus I?I kolmodini Pseudotectus comes I?1 Pseudophorus '?' stuxbergi I Pseudophorus profundus NUMBER 88 41 FIGURE 19 (opposite).?Relationships among the "pseudophorides." For abbreviations, see legend to Figure 7. Node 97 ("Pseudophorides"), funicle present (107); base projected anteriorly -30? (119); high K (123). Node 98, asymmetric ramp shapes (RR more convex) (52, 53, 72); strong, sharp LRC (76); BC lost (89); IM -15? off parallel to CA (98); aperture inclined -20? (110); high T (126). Node 99 {Streptotrochus? visbeyensis clade), moder? ately strong LRC (77); variable RRC (weak to absent) (63, 64). Node 100, straight lunulae (38); P = 30?-40? (48); asymmetric aperture (RR wider than LR) (57-59); RRC lost (64). Node 101 {Streptotrochus incisus clade), IM thicker at top and bottom and reflected around umbilicus (96, 106); thin, complete PI (103); ontogenetically increasing E and T (122, 127). Node 102, convex RR and LR (52, 72); moderately asymmetric aperture (right side extremely broad, left side broad) (58, 59); very strong LRC (77). Node 103 {Streptotrochus aff. S. incisus clade), PB lost (19); little ontogenetic change in RR convexity (53); right side of aperture inclined (113). Node 104, highly sigmoidal aperture (14); kinked lunulae (38); acute suture (69, 70); strong, squared LRC (76, 77); PI projecting in front of aperture (104); funicle present (107); base projected anteriorly -50? (119); moderate K (123). Node 105 {Pseudophorus clade), broad symmetric aperture (57-59); very strong, hood-like LRC frill (76, 77, 79, 84); both IM and base thick? ened (88); BC present (89); IM 30? off parallel to CA (98); moderate T (126). Node 106 {Discordichilus clade), convex RR and LR (52, 72); very asymmetric ramp lengths (RR extremely short, LR long) (55, 56); weak, squared LRC (77); whole aperture inclined -50? (109, 110, 113). Node 107 {Discordichilus dalli clade), fine, sharp GL (15); thin, complete PI (103); small size (141). Node 108 {Discordichilus kolmodini clade), moderately asymmetric aperture (right side extremely broad, left side broad) (58, 59); thick, contuse LRC (76). Hormotoma Turritoma Hormotoma Turritoma Plethospira !simulatrix aff. T. acrea confusa Cotter Fm. sp. cannonensis (see Figure 8) (go to Node 112)(go to Node 119)(go to Node 155) ? ? ? ? ? Plethospira cassina Seelya ventricosa ? Hormotoma " Straparollinoids" " Subulitoids" " Eotomarioids" Plethospira CI: 0.636 RI: 0.691 Figure 8 Node 7 FAMILY Murchisoniidae Plethospiridae FIGURE 20.?Relationships among basal "murchisoniinae." For abbreviations, see legend to Figure 7. Node 109, weak GL (15); extremely low E and very high T (121, 126). Node 110 ("Plethospiroids"), sinus angle = 30? (3, 4); narrow sinus (6, 7); PB width = 20? (20); P = 70? (48); slightly asymmetric aperture (58, 59); IM\base angle = 60? (94); slightly twisted siphon (99); nonreflected IM filling umbilicus (106, 108); low E (121). Node 111, loss of RR and LR swellings (60, 73); aperture inclined -20? (109, 110); large size (141). 42 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY oo CH "Q C ??; h ? .2 3^ "^3 C ^ Q CJ *? .2?s "3 o D a/ ^ o a Ha pL !n e a ?5 ?** j "5 ? 1 ? k 0 0 a g ^3 ?o "5? c 2.2 ra pa er ig o^ a K ^3 a S""3 ? *? ?5 a i - h &5 o ph os p gr an di H i J h * < 5^ a so ni id 3 p id ae ex o "o c3 n-i om at i* o c o ? o o CN tn in NUMBER 88 45 U S ^ ? 60 0 2 SPu n n n n D I n 00 O (N IT) IT) d d 46 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY eral band being very low on the whorl. A more-derived sub? clade (Catazone; nodes 129, 130) share a reduced peripheral band, a slit, and an inner margin that is contiguous with the pre? vious whorl rather than reflected. 11.3.2.1. "GONIOSTROPHINES".?One of the major "cyr- tostrophid" subclades includes species classified in Murchiso- nia, Hormotoma, Goniostropha Oehlert, and Sinuspira Perner (Figure 24, nodes 131-138). The last genus is highly derived and not typical of the clade, so I label this clade the "goniostro? phines." Synapomorphies of the clade include an asymmetric whorl shape, with a rounded left side and a flat or concave right side that culminates in an acute, attenuated suture. The clade is composed primarily of Silurian species. One Silurian subclade, the H. attenuata clade (node 138), is diagnosed by the loss of the peripheral lira that are primitive to "murchisoniinae." Spe? cies in another Silurian clade (the H. subplicata clade, nodes 132-137) possess very strong, sharp peripheral band lira, flat ramps, a more symmetrical shallower sinus, an acute suture, and an acute basal portion of the inner margin. The last feature becomes somewhat siphonate in the Donaldiella declivis clade (node 136), and a slit is present just under the suture on those species. The definition of Murchisonia given by both Knight et al. (1960) and Wenz (1938) includes Goniostropha. The type spe? cies of Murchisonia, M. bilineata (Dechen in De la Beche, 1832) is known from the Middle Devonian, so it could not be included here. Murchisonia bilineata possesses Goniostropha synapomorphies, such as a wide peripheral band with strong lira and a reduced sinus, so extended analyses probably will support the shared opinion of Wenz and Knight et al. The latter authors also included Cyrtostropha within Murchisonia, but these results suggest that Cyrtostropha evolved independently from Hormotoma. "Goniostrophines" (= Hormotoma salteri clade) include the only early Paleozoic species known to have had distinct proto? conch morphologies. Unfortunately, the high-spired "cyrtostro? phids" rarely preserve the apex. At least two species in the Hormotoma salteri clade (Sinuspira tenera Barrande in Perner, 1907, and Donaldiella declivis (Barrande in Perner, 1907)) possess large, planispiral protoconchs; however, an interesting implication of this analysis is that this distinctive protoconch morphology arose among some "cyrtostrophids" by the Middle Ordovician. 11.3.2.2. "OMOSPIRINES".?The earliest members of the other major "cyrtostrophid" subclade (Figures 25, 26, nodes 139-154) include species classified in Omospira Ulrich in Ul? rich and Scofield, so I refer to the clade as the "omospirines." "Omospirine" synapomorphies include increased shell expan? sion, reduced reflection and thickness of the inner margin, and a more symmetrical sinus. Omospira itself (node 139) is linked by a weaker peripheral band that is closer to the suture, a U-shaped sinus, and increased left to right asymmetry of the aperture. Most previous authors considered Omospira as an aberrant raphistomatid (e.g., Knight et al., 1960), but this analysis cor? roborates Wenz's (1938) classification of Omospira as a murchisonioid. It also should be noted that the "omospirines" bear little resemblance to the definition of the Omospirinae given by Knight et al. (1960), which included putative Silurian Omospira and post-Silurian genera. This analysis suggests that none of the post-Caradoc (Middle-Late Ordovician) members of this clade retained Omospira-like morphologies. Instead, the other "omospirine" species closely match traditional defini? tions of the Loxonematoidea. Most workers (e.g., Ulrich and Scofield, 1897; Koken, 1898; Wenz, 1938; Knight et al., 1960; Erwin, 1990b) considered loxonematoids to be derived murchisonioids. This analysis corroborates that view, with the caveat that Omospira-\ike "murchisoniinae" species are inter? mediate between Loxonema and traditional murchisonioid spe? cies, such as Hormotoma gracilis (Hall, 1847). Two "omospirine" subclades evolved during the Ordovician. One of these includes Loxonema, so I designate it the "loxone- matides." This clade is discussed below. The sister clade (Fig? ure 26, nodes 146-154) of the "loxonematides" is diagnosed by a very weak peripheral band and weak ornament. Previous workers placed these species in the Loxonematidae (e.g., Donald, 1905; Wenz, 1938; Knight et al., 1960), and this analy? sis suggests that they form a sister clade to the one including Loxonema. The genus Rhabdostropha Donald, 1905, appears to be typical of the group, so I refer to this clade as the "rhab- dostrophides" for lack of a more appropriate name. Synapomorphies of the "loxonematides" (Figure 25, nodes 142-145) include the loss of the peripheral band, a thick induc- tura, and a slightly flaring aperture. In the Loxonema berault- ensis clade (Figure 25, node 145), the sinus becomes deeper and culminates close to the suture. This culminates in Diploz- one crispa (Lindstrom, 1884), which has a siphonate base and a nearly slit-like sinus just under the suture. The "rhabdostrophides" are diagnosed by an inner margin that is contiguous with previous whorls, anterior production of FIGURE 24 (opposite).?Relationships among the "goniostrophines." For abbreviations, see legend to Figure 7. Node 125 {Hormotoma gracilis clade), asymmetric sinus shape (left curve stronger) (8-10); p = 70? (48); moderately asymmetric ramp lengths (right side long, left side short) (55, 56). Node 131 ("Goniostrophines"), dull, wide ML present between BL (27); attenuated suture (69, 70). Node 132 {Hormotoma subplicata clade), sharp BL (22); slightly asymmetric aperture (right side broad, left side moderately broad) (58, 59). Node 133, fine, sharp GL (15); ML lost (27). Node 134 {Goniostropha cava clade), thin, incomplete PI (103). Node 135, left sinus angle = 40? (4); P = 90? (48); long, asymmetric ramps (RR moderately long, LR long) (55, 56); flat LR (72). Node 136 {Murchisonia paradoxa clade), PB width ? 10? (20); slit present (34); nonattenuated acute suture (69, 70); strong, sharp LRC present (75-77). Node 137, very asymmetric ramp lengths (RR short, LR very long) (56, 57); very asymmetric aperture (right side extremely broad, left side nar? row) (58, 59); IM\base angle = 60? (94). Node 138 {Hormotoma attenuata clade), fine, sharp GL (15); PB width = 10? (20); BL lost (21); IM\base angle = 60? (94); moderate size (141). NUMBER 88 47 a k "1 s ,g ?a a to rn o S x. o =C ?S 3? 3 a c a Q o- a g o n e 3 ?S iso n -s: ^J M ur a ?? | Ik to a k 4) e ?Si g a a fc a ?a _> "*2S ?J >^ "Xj a ?5 a s c 53 3= I a >^ 2 jg. 0) in "2 :a c o his CJ M ur aj a T3 4?> 6 o Lo x 1) O D- O f-1 r > OO CO 48 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY Hormotoma Omospira Omospira Rhabdostropha Loxonema Loxonema Loxonema Loxonema Diplozone trentonensis alexandra laticincta primitiva murrayana crossmanni sinuosa beraultensis crispa (see Figure 23) (go to Node 146) ? ? ? ? ? Omospira Rhabdostropha Loxonema Diplozone CI = 0.773 RI = 0.781 Figure 23 Node 125 FAMILY Murchisonidae l-^fl Loxonematidae Plethospiridae luiiJ Raphistomatidae FIGURE 25.?Relationships among the "omospirines" and "loxonematides." For abbreviations, see legend to Fig? ure 7. Node 139 ("Omospirines"), left sinus angle -40? (4); nonreflected IM (106); very high T increasing over ontogeny (126, 127). Node 140 {Omospira clade), P = 50? (48); very asymmetric ramp lengths (RR very short, LR long) (55, 56); large size (141). Node 141 {Rhabdostropha primitiva clade), left sinus begins at base of LR (7); nonthickened IM (87). Node 142 ("Loxonematides"), PB lost (19); very asymmetric ramp lengths (RR very short, LR very long) (55, 56); broad (asymmetric) aperture (58, 59); acute suture (69); nonthickened IM (87); arched IM (95); PI thickness same as rest of shell (103); weakly flaring aperture (105); very lowE (121). Node 143, leftside of sinus beginning near base of LR (7); fine, sharp GL (15); IM 15? off parallel to CA (98). Node 144 {Loxonema sinuosa clade), sinus angle = 30? (3, 4); IM fills umbilicus (108). Node 145 {Loxonema beraultensis clade), sinus angle = 40? (3,4); symmetric sinus width (5-7); attenuated suture (69, 70); strongly curved IM (95); IM contigu? ous with previous whorl (108); inclined aperture (109). FIGURE 26 (opposite).?Relationships among the "rhabdostrophides." For abbreviations, see legend to Figure 7. Node 146 ("Rhabdostrophides"), nearly symmetric sinus angles = 20? (3, 4); very weak BL (23); aperture contiguous with previous whorl (108); isometric T (127); sparse, fine ornament present (129-133). Node 147 {Girvania excavata clade), symmetric U-shaped sinus (5-7, 9, 10); PB width = 05? (20); dense ornament (130, 134). Node 148, very shallow sinus angles = 20? (3, 4); fine, sharp GLs (15); loss of RR ornament (133). Node 149 {Holopella regularis clade), sinus angle = 10? (3,4); U-shaped sinus (9, 10). Node 150 {Kjerulfonema clade), sinus nearly absent (3, 4); thick? ened IM (87); extremely low E (121); strong ornamentation (129-137). Node 151 {Macrochilus buliminus clade), sinus nearly absent (3, 4); p = 30? (48); highly asymmetric ramp lengths (RR highly contracted) (55, 56); thickened middle of IM (96); PI absent (103). Node 152, extremely asymmetric ramp lengths (RR extremely contracted) (55, 56); IM || to CA (98). Node 153, sinus lost (1); IM\base angle = 45? (94); straight IM (95); IM reflected around umbili? cus (106); open umbilicus (108). Node 154 {Stylonema clade), symmetric sinus width (5-7); p = 70? (48); slightly asymmetric, very broad aperture (very broad right side, broad left side) (58, 59); RR swelling present (60); swollen base of LR (73); thin, incomplete PI (103); IM contiguous with previous whorl (108); curved base (120); ornament throughout left side of aperture (129). NUMBER 88 49 r-4 \?, in vo ? ? II l-H l-H 50 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY the aperture (e.g., Figure 3D), and well-developed ornamenta? tion. Early species in this clade might retain a very weak bilin? eate peripheral band, but it is not clear whether the feature is a peripheral band or general ornament. Among more-derived species, such as the Holopella regularis clade (Figure 26, nodes 149-154), the sinus is extremely reduced and broadly U- shaped. Stylonema Perner, 1907 (Figure 26, node 154) is some? what more divergent, possessing a curved base, a much rounder and slightly more asymmetric aperture, and marked swelling near the suture. Horny (1955) discussed the phyloge? netic position of Stylonema, but only in general terms relative to Loxonema, so this exact estimate cannot be compared to that assessment. The most noteworthy subdivision within the H. regularis clade is the Macrochilus buliminus subclade (Figure 26, nodes 151- 153). This last subclade possesses a gross mor? phology that converges strongly on that of derived "subulitids" (see Figure 22, node 122). Synapomorphies include the near loss of the sinus (which is completely lost in the most derived species), a inner margin that is differentially thickened in the middle, the loss of the parietal inductura, and an extremely asymmetric aperture with the right side strongly reduced. This analysis supports the commonly held view that the Loxonematoidea evolved from the Murchisonioidea (e.g., Wenz, 1938; Knight et al., 1960). I noted above that these re? sults contradicted the proposal by Knight et al. (1960) that the Subulitoidea evolved from the Loxonematoidea. This study does suggest, however, that a clade of species classified in the Subulitoidea (i.e., the Macrochilus buliminus clade) did evolve from the "loxonematides." This corroborates Erwin's (1992) claim that traditional definitions of the Subulitoidea are polyphyletic. In addition to the general suggestions about loxonematoid re? lationships, authors also have discussed more specific relation? ships of these species. As noted above, the idea of Knight et al. (1960) that Diplozone evolved among the Plethospiridae is re? jected by this study. Their suggestion that both Holopella M'Coy, 1851, and Rhabdostropha are closely related to Lox? onema is upheld, although this analysis indicates that the former genera belong to a different subclade than does Lox? onema. The suggestion by Peel and Yochelson (1976) that the Silurian genus Kjerulfonema Peel and Yochelson is related to Girvania is supported herein. II.4. "Eotomarioids" Most of the taxa classified in the Pleurotomarioidea by Knight et al. (1960) form a major clade that includes species classified in the Eotomariidae, Gosseletinidae, Lophospiridae, Luciellidae, and Phanerotrematidae (Figures 27-35, nodes 155-215). This clade is similar to Ulrich and Scofield's defini? tion of the Eotomarioidea, although their definition does in? clude many "raphistomatids" (see above). "Eotomarioid" syna? pomorphies include a peripheral band with stronger, sharper lira and weak channels beneath those lira, a straight inner mar? gin, flatter ramps, and the loss of swellings on top of the ramps. The earliest species belonging to this clade fit the general de? scription of Turritoma, which corroborates Ulrich and Scofield's (1897:1013) suggestion that eotomarioids evolved from Turritoma. II.4.1. "LOPHOSPIRIDS" "Eotomarioids" include two major subclades. One of these contains the Lophospiridae plus species assigned to the genera Solenospira Ulrich and Scofield, 1897, and Ectomaria Koken, 1896 (Figure 27, nodes 156-165). "Lophospirid" synapomor? phies include flat to slightly concave right and left ramps, a strong lower ramp carina, a deep V-shaped sinus, and sharp threads on a bilineate peripheral band. An early appearing "lo? phospirid" subclade that includes several species classified as Solenospira and Ectomaria (node 160 and above) represents the sister taxon of traditional lophospirids. The primary syna? pomorphy of this subclade is an extremely wide peripheral band with very strong peripheral lira. Most workers considered Solenospira to be redundant with Ectomaria (e.g., Wenz, 1938; Knight et al., 1960), a view that this analysis supports. Those authors also considered Ectomaria to be a close relative of Hormotoma and other traditional murchisoniids, which this analysis contradicts. Node 160 and its descendants represents the most diverse "archaeogastropod" clade of the Ordovician, including species classified as Arjamannia Peel, 1975a, Donaldiella Cossmann, 1903, Eunema Salter, 1859, Globonema Wenz, 1938, Gy- ronema Ulrich in Ulrich and Scofield, 1897, Longstaffia Coss? mann, 1908, Lophospira Whitfield, 1886, Loxoplocus Fischer, 1885, Pagodospira Grabau, 1922, Proturritella Koken, 1889, Ptychozone Perner, 1907, Ruedemannia Foerste, 1914, Schizol- opha Ulrich in Ulrich and Scofield, 1897, Trochonema Salter, 1859, and Trochonemella Okulitch, 1935. The primary synapo? morphies of lophospirids are a trilineate peripheral band with a FIGURE 27 (opposite).?Relationships among the "eotomarioids." For abbrevi? ations, see legend to Figure 7. Node 155 ("Eotomarioids"), sharp, moderately strong BL (22, 23); LR swelling lost (73). Node 156, flat ramps (52, 72). Node 157 ("Clathrospirids"), left side of sinus sharper and narrower than right (2-7); very asymmetric ramp lengths (RR very long, LR short) (55, 56); left and right widths of aperture symmetrical (57-59). Node 158 ("Lophospirids"), weak GL (15); p = 70? (48); asymmetric ramp lengths (RR shorter) (54-57); RRC and LRC present (64, 75); both IM and base thickened (88); isometric T (127). Node 159 {Pagodospira cicelia clade), PB width = 15? (20); ML present (27); ACh present (45); asymmetric ramp shapes (RR more convex) (52, 53, 72). Node 160 {Ectomaria adelina clade), PB width = 30? (20); extremely strong BL (23). Node 161, sharp, thread-like RRC and LRC (66, 76). Node 162 {Murchisonia callahanensis clade), sinus angle = 30? (3, 4); concave ramps (52, 72); RR becoming more concave over ontogeny (53); very low E (121); ontogenetically increasing expansion (122); high K (123); isometric T (127). Node 163 {Ectomaria pagoda clade), moderately strong LRC (77). Node 164 {Ectomaria nieszkowskii clade), symmetric ramp lengths (54-56). Node 165 {Ectomaria prisca clade), RRC lost (64). NUMBER 88 51 6 o o * 2 W J 69 9 o n l-H u 72 4 o il 2 52 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY pronounced channel. The phylogenetics of this clade are dis? cussed elsewhere (Wagner, 1990, 1995a, 1999). II.4.2. "CLATHROSPIRIDS" The second major "eotomarioid" subclade includes the re? mainder of the taxa previously classified in the Pleurotomario- idea (Figures 28-35, nodes 166-215). I designate the clade as "clathrospirids" because most of the earliest species have been classified in the genus Clathrospira. "Clathrospirid" synapo? morphies include an extended right ramp, placement of the pe? ripheral band partially on the right ramp, and an asymmetrical sinus with a wider, shallower right half. Like the "raphistomatids," two early "clathrospirid" sub? clades existed in different biogeographic realms, with one pri? marily North American (Laurentian) and the second primarily Baltic; however, there is substantial biogeographic overlap be? tween the two clades by the end of the Ordovician. II.4.2.1. "LIOSPIRINES".?The Laurentian "clathrospirid" subclade includes species classified as Eotomaria, Liospira Ulrich and Scofield, and Paraliospira Rohr, with the first ge? nus representing a paraphylum relative to the latter two (Figure 29, nodes 172-181). I label this clade as the "liospirines," al? though the clade differs greatly from the definition of the Li- ospirinae sensu Knight et al. (1960). "Liospirine" synapomor? phies include the projection of the inner margin at a high angle relative to the coiling axis (e.g., Figure 3c), the presence of a channel under the peripheral band, and a dull carina on top of the right ramp, which fills the suture. Two "liospirine" sub? clades were moderately diverse during the Caradoc and Ash- gill (Middle-Late Ordovician). One includes true Liospira (Figure 29, nodes 174-177). Liospira synapomorphies include a thickened inner margin, very weak threads on the peripheral band, a shallower, more symmetric sinus, isometric translation (in contrast to the increasing translation shown over the ontog? eny by more plesiomorphic members of the clade), and the fill? ing of the umbilicus by the inner margin and an extended in- ductura. The last features suggest that in life Liospira wore their shells like caps on top of the body rather than carrying them by the aperture. A slit does not diagnose the clade, but it is present on the most derived species. The second "liospirine" clade includes species assigned by Rohr (1980) to Paraliospira (Figure 29, nodes 178-181). Paraliospira synapomorphies include very strong square- shaped peripheral band lira, a very shallow right side of the si? nus, and a nearly planispiral, widely umbilicate form produced by very low curvature and translation. In addition, the base is projected much further in front of the inner margin. More-de? rived species (i.e., the P. mundula clade) have much stronger channels beneath the peripheral band, a channeled carina on top of the right ramp, a dull, channeled left carina, and a more symmetrical sinus. It is noteworthy that this clade includes Li? ospira larvata (Salter, 1859) and Eotomaria supracingulata Ulrich and Scofield, 1897, as this substantiates Ulrich and Scofield's (1897) suggestion that those species are closely re? lated to P. mundula (Ulrich and Scofield, 1897), P. angulata (Ulrich and Scofield, 1897), and their allies (which Ulrich and Scofield tentatively placed in Liospira). Ambiguity about character-state polarity affects the exact clade structure within the L. larvata clade. Paraliospira mun? dula and P. rugata (Ulrich and Scofield, 1897) share a very thick inner margin that is folded back into the umbilicus to form a funicle. This feature obscures the inner margin and makes it impossible to determine if an inner margin lirum is present or absent. To be more exact, if the feature was present, then it would be obliterated by the umbilicus. If the feature is assumed to be absent, the best reconstruction is to link P. angu? lata to the Eotomaria rupestris clade. This reconstruction is supported by the fact that some specimens of P. mundula lack? ing funicles (owing either to intraspecific variation or differ? ences in taphonomy) lack the lirum, suggesting that the feature is actually absent. This analysis supports the idea that Liospira and allies are closely related to Eotomaria, which was the opinion of Ulrich and Scofield (1897) and Wenz (1938). The analysis strongly differs from the views of Knight et al. (1960) in two ways. First, those authors considered the Liospirinae to be a subdivision of the Raphistomatidae. There are strong similarities between some "raphistomatids" (e.g., Pararaphistoma) and Liospira. In fact, the widely distributed and somewhat variable species P. qualteriata appears to have been classified as Liospira by some authors (e.g., Longstaff, 1924). Only the most derived Liospira species, such as L. micula (Hall in Hall and Whitney, 1862), dis? play "lesueurilline" synapomorphies. Linking these species di? rectly (i.e., so that L. micula is considered a plesiomorphic member of Liospira or so that L. micula is removed from the "liospirines" to the "lesueurillines") also invokes statistically significant stratigraphic gaps (see Appendix 3). The second difference between these results and the taxon? omy of Knight et al. (1960) is that those authors included post- Ordovician species in the Liospirinae. Although Liospira com? monly is reported from Silurian strata (e.g., Peel, 1977), this analysis found no Silurian "liospirines." This might be due to the poor preservation of the relevant specimens, as several of the important synapomorphies (e.g., peripheral band type and FIGURE 28 (opposite).?Relationships among basal "clathrospirids." For abbreviations, see legend to Figure 7. Node 166, PB width =15? (20); slightly prominent PB (33); left side of aperture inclined -20? (111, 112); base pro? jected anteriorly -20? (118, 119); moderate E (121); moderate K (123); high T (126). Node 167 {Clathrospira Itrochiformis clade), p ? 80? (48); both IM and base thickened (88). Node 168 {Clathrospira euconica clade), symmetric sinus depth (2), ACh lost (45); asymmetric aperture (RR wider than LR) (57-59); RRC present (64). Node 169 {Clathrospira inflata clade), sinus angle -60? on left side (4); asymmetric aperture (RR much wider than LR) (57-59); moder? ately convex whorls (52, 72). Node 170 {Mourlonia mjoela + Eotomaria con? vexa), narrow PB with moderately strong PL (20, 23); very convex whorls (52, 72); IM 15? off parallel to CA (98); moderate size (141). NUMBER 88 53 < r-CN m ( N r> ON TJ- i n d d U ai 60 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY 3 5 C 3 so la , o ce r o b !?. CX QH s i s 3 1 R -5 u a *~^ s ilu n ill ei 5-* ^ E o 3 W r ^ I > t * "2 'o "i E o o w o S3 hi (/3 * , * * , * \ 2 o ?E ?? . - o ?is o b % o c/j O -2. o- E o u H rV ?3 1 o S 73 a> e: o S .2 8 Si * O JJ u ?a > Z u u fc ?? '5 -^ a. S & ? ?j3 u - = O PL, z ? ? O Q. O & te s | if r c u e 2 I ? u. Ho \c 67 all the vetigastropod-like species of the Silurian belong to that clade. I suggest (tentatively) that "clathrospirids," especially within the "planozonide" subclade, represent the most likely candidates because these derived "clathrospirids" share similar slits, sinuses, peripheral bands, and apertural orientations with modern pleurotomarioids and coiled scissurelloids. Many workers have suggested that the Loxonematoidea are the earli? est apogastropods (e.g., Wenz, 1938; Knight et al., 1960; Fret- ter and Graham, 1962; Houbrick, 1979, but see Houbrick, 1988), and many workers (e.g., Haszprunar, 1988; Tracey et al., 1993) consider loxonematoids to be plesiomorphic apogas? tropods. Other "hormotomoids" make poor candidates for the ancestors of the Apogastropoda. If vetigastropods are derived "clathrospirids" and apogastropods are "loxonematides," then the vetigastropod-apogastropod divergence (roughly Figure 20, node 109) occurred no later than the Tremadoc (Early Or? dovician), i.e., around 490 million years ago (Harland et al., 1990). The origins of patellogastropods are less obvious. D.R. Lind? berg (pers. comm., 1993) has suggested that the clade evolved from coiled, sinus-bearing, septate gastropods. If so, then the "euomphalinae" represent the most likely ancestor. This exten? sion of Lindberg's basic scenario fits the neontological data well, as nearly all recent phylogenetic analyses indicate that pa? tellogastropods are the outgroup to all other gastropods (Lind? berg, 1988; Haszprunar, 1988; Tillier et al., 1992). Other as? pects of this phylogenetic model also fit paleontological data, as some trochoid-like "euomphalinae" (e.g., Pseudophorus) possessed morphologies suggesting single-gilled anatomies, adaptations for clamping onto hard substrates, and septa re? stricted to the youngest whorls. These assessments of patello- gastropod-"euomphalinae" relationships clearly must be con? sidered highly tentative, and other data (e.g., protoconch morphology and shell mineralogy) are necessary to corroborate it. If accurate, however, then patelloids and the vetigastropod + apogastropod clade diverged by the Late Cambrian, i.e., around 510 million years ago (Harland et al., 1990). Linking patellogastropods and the "euomphalinae" contra? dicts Haszprunar's (1988) suggestion that the Euomphaloidea are closer to vetigastropods than to patellogastropods. This analysis suggests that the "euomphalinae" are not closely re? lated to the most appropriate precursors to the vetigastropods. McLean (1981, 1989, 1990) linked the Euomphaloidea with the Neomphalina. As with the Neritoidea, neontological studies have reached little consensus about the relationships of the Neomphalina, so paleontological data likely will be important. Unfortunately, almost nothing is known about the fossil ante? cedents of that taxon, so it is difficult for paleontological data to address McLean's phylogenetic hypothesis at this time. The speculations about the early Paleozoic precursors of pa? tellogastropods, vetigastropods, and apogastropods can only be described as consistent with both paleontological and neon? tological data. It still has important implications for phyloge? netic analyses of the higher taxa, however, because it is evi- 68 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY dent that the most likely antecedents of those taxa should have diverged by the Early Ordovician, i.e., shortly after gastropods evolved. As noted above, the phylogenetic affinities of the Neomphalina, Neritoidea, and the various hydrothermal-vent taxa are poorly understood. The one consistent implication of the conflicting results from different molecular and morpho? logic analyses is that these gastropod problematica diverged before or around the time that vetigastropods and apogastro? pods diverged, which this study suggests was some 490 mil? lion years ago (Harland et al., 1990). Long divergence times lead to "long branches," which confound phylogenetic analy? ses (Felsenstein, 1978; Huelsenbeck and Hillis, 1993). Echin- oderms provide an important analogy, as the five extant major classes within that phylum also diverged during the early Pale? ozoic. Cladistic analyses using 18s rRNA have not been able to resolve the relationships among those classes, and different morphologic data sets produce conflicting results (Smith, 1989). Given that extant orders and suborders of gastropods diverged at about the same time, we should expect different neontological data sets to produce very different hypotheses about relationships among the Gastropoda. Figure 37 presents a summary of the general phylogenetic relationships proposed herein, for comparison with the hy? potheses outlined in Figure 1. It also includes the possible im? plications for neontological relationships that this analysis suggests. Systematic Paleontology In the discussion of the estimated relationships among early gastropods, I repeatedly emphasized the many similarities be? tween the results of this analysis and the previously held views of numerous workers. Despite the general congruence, how? ever, it is clear that gastropod taxonomy requires major revi? sion in order to reflect many aspects of gastropod phylogeny, and also to reflect more accurately the evolutionary dynamics within the clade. It must be emphasized that the taxonomy proposed herein is phylogenetic rather than typologic. Type species are granted no special importance except as convenient labels for genera; i.e., if a clade or paraclade (see definition below) includes a previ? ously designated type species, then that genus labels the clade or paraclade. If a clade or paraclade comprises multiple type species, then I adhered to standard taxonomic priority. Because the taxonomy is phylogenetic and given that the text of this pa? per discusses the characters uniting each clade in some detail, little discussion of diagnostic characters is given here. Also, gastropods typically are described by characters such as spire height, general aperture shape and inclination, columella type, and ornament. As noted in the text, these features are com? pound characters that rarely denote particular homologies and convey only indirect phylogenetic information. Accordingly, I eschew "traditional" descriptions, referring the reader to the character matrices and synapomorphy lists provided in the ap? pendices and figures. I did attempt, however, to adhere to more traditional grade-based criteria to separate derived suprage- neric taxa from their "ancestral" taxa. Another important point is that I have attempted to eliminate monotypic genera. Some monotypic genera, however, are known from poorly sampled intervals (especially the Llan- virn-Llandeilo and the Early Llandovery). Further sampling of those intervals could easily reveal that those species belong to separate clades or paraclades; if and when such species are sampled and included in phylogenetic analyses, then those gen? era should be reinstated. In addition to reflecting general phylogenetic relationships, taxonomy also should reflect evolutionary dynamics (e.g., pat? terns of standing diversity and relative extinction and origina? tion intensities). Contrary to the claims that only monophyletic taxonomy reflects evolution (de Queiroz and Gauthier, 1990, 1992, 1994), monophyletic taxa actually reflect history. When using a historical system to define taxa, later evolution will dis? tort the apparent evolutionary patterns of earlier time intervals by "retroactively" affecting which clusters of species are recog? nized as taxa (Van Valen, 1978; Sepkoski, 1987; Janis, 1992). Wagner (1995c) demonstrated the lack of congruence between the evolutionary patterns implied by phylogeny and monophyl? etic taxonomy among the gastropods described in this paper. Thus, regardless of whether strictly monophyletic taxa might be desirable for other reasons, paraphyletic taxa usually are necessary to reflect evolutionary dynamics accurately (Sepko? ski and Kendrick, 1993; Patterson, 1994; Smith, 1994; Wagner, 1995c; Maley et al., 1997). There is a close correspondence be? tween diversity patterns implied by the phylogeny presented herein and the patterns implied by traditional taxa (Wagner, 1995c). This has important implications for historical biodiver? sity studies that use higher taxa. Higher taxa are not considered entities in themselves in such studies, but rather they are sam? pling proxies for species (Sepkoski, 1987). Using higher taxa as proxies for species assumes that those higher taxa are not polyphyletic. The fact that highly polyphyletic gastropod taxa still accurately reflect species-level patterns suggests that taxo? nomic diversity studies are robust to violations of at least one important assumption. Although many of the taxa are paraphyletic, all supraspecific taxa defined herein include at least one monophyletic clade at all times. Such taxa are labeled "paraclades" below (e.g., Raup and Gould, 1974). Paraclades are denoted with a "f" symbol. Patterson and Rosen (1977) proposed this notation for denoting plesions (i.e., extinct sister clades of extant taxa). Although paraclades and plesions are not synonymous, plesions also have been described as plesiomorphic sister groups (e.g., Wiley, 1979; Eldredge and Cracraft, 1980). Paraclades as de? fined herein include a plesiomorphic sister taxon of a morpho? logically derived clade, with the plesion retaining the name as? sociated with the ancestral morphotype. This emendation of the plesion concept allows both the recognition of ancestral types NUMBER 88 69 (if only in a broad sense) and the accurate depiction of diversity dynamics within the clade. Traditional "evolutionary" taxonomy (e.g., Simpson, 1961) has relied as much on perceived morphologic grade and dis? parity as perceived phylogenetic relationships. Cladistic taxon? omy grants no importance to disparity, on the basis that only sister taxa represent real units in biology. Curiously, it has never been demonstrated that clade membership is a better pre? dictor of extinction and speciation dynamics or any other "real" aspects of evolution than is morphologic or ecologic grade. In the absence of appropriate analyses for distinguish? ing such groups (see, e.g., Purvis et al., 1995), I used tradi? tional criteria for separating derived families and superfamilies from their paraphyletic relatives. This is done in part to main? tain as many traditionally recognized taxa as possible, for an overly radical revision of any taxon probably is useless if the taxonomy is unrecognizable to workers specializing on that taxon. Also, it seems unlikely that malacologists would adhere to a taxonomic scheme that would reduce derived taxa, such as opisthobranchs, to a subfamily or genus within the Loxonema? toidea. Many differences usually accorded generic or even fa? milial-level disparities (e.g., ornamentation, closed-or open- coiling, spire height) are recognized, however, only when they coincide with major clades. In the future, methods of optimiz? ing evolutionary parameters onto estimated phylogenies (see, e.g., Sanderson and Bharathan, 1993; Purvis et al., 1995) but designed to recognize major changes in those parameters might be used to construct quantitative, repeatable evolution? ary taxonomies. The primary taxonomic revision proposed herein is the re? moval of all early Paleozoic gastropods from the Pleurotomar- iina and transferring nearly all of them into either the Murchisoniina or Euomphalina. Pleurotomarioids are a sub? clade of the Vetigastropoda (Haszprunar, 1987), which in turn likely is a subclade of (or possibly equivalent to) one of the major subclades documented in this study. Determining the or? igin of the Pleurotomarioidea and other vetigastropods will be a topic of future research. Class GASTROPODA Cuvier, 1797 Order "ARCHAEOGASTROPODA" Thiele, 1925 REMARKS.?As noted in the text, the "Archaeogastropoda" is not a particularly useful label owing to uncertainties and in? consistencies in the grade or paraphylum it defines. Better un? derstanding of the relationships of modern gastropods to their early Paleozoic ancestors should obviate the need for the taxon. Ultimately, ordinal status probably should be granted to the clades corresponding to the Euomphalina and Murchisoniina as defined below; however, too much speculation is required pres? ently to equate those taxa diagnosed herein with clades com? prising modern taxa. Family fSlNUOPEiDAE Wenz, 1938 REMARKS.?Several Late Cambrian and earliest Ordovician gastropods do not belong to either the murchisoniinae or eu? omphalinae clades. It is simplest to place these taxa within the family Sinuopeidae, although the new definition of that taxon is very different from that of either Wenz (1938) or Knight et al. (1960). Wenz's original subfamilial definition of the taxon was limited to Sinuopea and Taeniospira, which are considered synonymous herein. The definition of Knight et al. included many post-Ordovician species that possessed U-shaped sinuses and no peripheral bands (like Sinuopea and Chepultapecia), but this analysis suggests that those species are members of the Raphistomatidae (see definition below). Although this family is paraphyletic relative to the Euomphalina and Murchisoniina (and possibly the Sinuitidae), sinuopeids represent a cohesive group whose termination in the Early Arenig is marked by the extinction of at least two true clades that retain very primitive morphologies. Genus f Sinuopea Ulrich, 1911 FIGURE 8 1 Chepultapecia Ulrich in Weller and St. Clair, 1928. Taeniospira Ulrich and Bridge in Ulrich et al., 1930. TYPE SPECIES.?Sinuopea sweeti (Whitfield, 1882). FIRST KNOWN APPEARANCE (FKA).?S. sweeti: Oneota Sandstone (Late Trempealeauan). LAST KNOWN APPEARANCE (LKA).?Taeniospira !st. clairi Ulrich and Bridge in Ulrich et al., 1930: Smith Basin Lime? stone (Late Tremadoc (Demingian)). ADDITIONAL ASSIGNED SPECIES.?Sinuopea basiplanata Ul? rich and Bridge in Ulrich et al., 1930; Taeniospira emminencis Ulrich and Bridge in Ulrich et al., 1930. REMARKS.?Sinuopea typically is diagnosed as lacking a pe? ripheral band, although at least one previously recognized spe? cies (S. basiplanata) retains the feature. Peripheral bands typi? cally are highly diagnostic features, but the feature evidently was plastic among the earliest gastropods. As there is no phylo? genetic boundary between Sinuopea and Taeniospira, they are synonymized herein. Taeniospira frequently is described as possessing a slit; however, the feature occurs only on the type specimen of one species and, moreover, appears to represent breakage rather than a true slit. Chepultapecia is somewhat more problematic, as it appears to be monotypic and known only from small, poorly preserved specimens. These specimens seem to possess U-shaped sinuses and lack peripheral bands, which are possible synapomorphies with very derived Sinuo? pea. Interestingly, the most parsimonious tree places them in? termediate between S. sweeti and the bellerophont Owenella. Better specimens for both Chepultapecia and Owenella are needed to explore this possibility more fully. 70 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY Genus ^Schizopea Butts, 1926 FIGURES 7,8 Dirhachopea Ulrich and Bridge in Ulrich et al., 1930. Rhachopea Ulrich and Bridge in Ulrich et al., 1930.?Knight et al., 1960. TYPE SPECIES.?Schizopea washburnis Butts, 1926. FKA.?Dirhachopea normalis Ulrich and Bridge in Ulrich et al., 1930; D. subrotunda Ulrich and Bridge in Ulrich et al., 1930; Schizopea typica (Ulrich and Bridge in Ulrich et al., 1930): Eminence Dolomite (Late Trempealeauan). LKA.?Schizopea washburnis: Roubidoux Formation (Late Tremadoc (Demingian)). REMARKS.?The species classified in the genera listed above vary in peripheral band and coiling features. These fea? tures are more static in later gastropods, but they evidently varied among closely related species during the early phases of gastropod evolution. Genus Euconia Ulrich in Ulrich and Scofield, 1897 FIGURE 8 Roubidouxia Butts, 1926. Rhombella Bridge and Cloud, 1947. Jarlopsis Heller, 1954.?Knight et al., 1960. TYPE SPECIES.?Euconia etna (Billings, 1865). FKA.?Jarlopsis conicus Heller, 1954: Gasconade Dolomite (Early Tremadoc (Gasconadian)). LKA.?Euconia etna: Smithville Formation (Early Arenig (Cassinian)). ADDITIONAL ASSIGNED SPECIES.?Rhombella umbilicata (Ulrich and Bridge in Dake and Bridge, 1932). REMARKS.?Several monotypic genera belonging to one clade are lumped herein. The species are united principally by greatly expanded right ramps and contracted left ramps, leav? ing the monolineate peripheral band very low on the whorl. The orientation and shape of the right ramp coupled with the relatively high translation produces a trochiform gross mor? phology. Genus Gasconadia Ulrich in Weller and St. Clair, 1928 FIGURE 8 TYPE SPECIES.?Gasconadia putilla (Sarderson, 1896). FKA.?Gasconadia putilla: Chepultepec Dolomite (Early Tremadoc (Gasconadian)). LKA.?Gasconadia putilla: Gasconade Dolomite (Early Tremadoc (Gasconadian)). REMARKS.?The highly derived nature of this monotypic ge? nus makes determining its relationships difficult. Although it is simplest to link it to Dirhachopea, it is not much less parsimo? nious to derive it from Sinuopea. Therefore, I retain the genus at the present. ?Genus Calaurops Whitfield, 1886 REMARKS.?This genus is represented by numerous poorly preserved specimens. It might be derived from Late Tremadoc Dirhachopea; if so, then the two should be synonymized. It is also equally likely that Calaurops is derived from Bridgeites or even Macluritella. If so, then it should become the senior syn? onym for either of those two genera. Although the genus name has priority over all possible relatives, it should be considered invalid until better material can place its relationships Suborder EUOMPHALINA de Koninck, 1881 REMARKS.?The Euomphalina herein equals the "euompha? linae" clade defined and diagnosed above. Four superfamilies are recognized, the Ophiletoidea, Macluritoidea, Euompha? loidea, and Platyceratoidea. The first three all evolved in the Tremadoc from species that would be classified in the genus Ophileta, which renders the Ophiletoidea nominally paraphyl? etic relative to the others. The final superfamily, the Platycera? toidea, likely evolved within the Euomphaloidea, but the great morphologic (apparently accompanied with ecologic) disparity renders its exact relationships problematic. This study does not thoroughly examine platyceratoids and detailed systematic work certainly is necessary. In the interim, it is prudent to sepa? rate the clade at a fairly high taxonomic level. Because Knight et al. (1960) placed the Euomphaloidea within the Macluritina, one might argue for labeling the taxon the Macluritina in order to preserve priority. Macluritoids are associated with a particular highly derived morphology, how? ever, and workers insisting on grade-based classifications likely would find "macluritoids" an unacceptable classification for many species within this clade. A similar argument would apply to labeling the clade the Patellogastropoda, if in fact pa- telloids did evolve from members of this clade. If the latter is true, then the Euomphalina (as defined herein) likely is equal to the Eogastropoda of Ponder and Lindberg (1996, 1997). Be? cause "Euomphalina" also is associated with a morphologic grade, "Eogastropoda" would be a preferable label for this clade. Superfamily fOPHlLETOIDEA trans, nov. Knight, 1956 REMARKS.?The Ophiletoidea corresponds to the "ophile? toid" clade plus the species Ophileta supraplana. It is much more similar to the Ophiletidae as defined by Morris and Cleevely (1981) than to the Ophiletinae as defined by Knight et al. (1960). Members of this clade typically are diagnosed by a combination of characters yielding a lenticular aperture, a strong monolineate peripheral band, and an inner margin that typically forms the base of the shell. As with other higher taxa, all of these features are modified in the most derived members of the clade. NUMBER 88 71 Genus ^Ophileta Vanuxem, 1842 FIGURES 8,9 Ozarkispira Walcott, 1924. TYPE SPECIES.?Ophileta complanata Miller, 1889. FKA.?Ophileta supraplana Ulrich and Bridge in Ulrich et al., 1930: Chepultepec Dolomite (Early Tremadoc (Gascona? dian)). LKA.?Ophileta complanata: Nittany Dolomite (Late Trem? adoc (Demingian)). ADDITIONAL ASSIGNED SPECIES.?Ozarkispira leo Walcott, 1924. REMARKS.?The inclusion of species classified in Ozark? ispira (one or more small species that typically are poorly pre? served) maintains Ophileta as a paraclade. Genus f Lecanospira Butts, 1926 FIGURE 9 Barnesella Bridge and Cloud, 1947. TYPE SPECIES?Lecanospira compacta (Salter, 1859). FKA.?Lecanospira compacta: Nittany Dolomite (Late Tremadoc (Demingian)). LKA.?Barnesella aff. B. lecanospiroides Bridge and Cloud, 1947 (sensu Rohr et al., 1995): Skoki Formation (Early Llanvirn). ADDITIONAL ASSIGNED SPECIES.?Barnesella Hecano- spiroides Bridge and Cloud, 1947; Lecanospira nereine (Bill? ings, 1865). REMARKS.?Lecanospira now includes most species classi? fied in Barnesella, which previously was separated at the sub? genus or genus level based on stratigraphic position. Ulti? mately, it might be more appropriate to consider Lecanospira a junior synonym of Ophileta. Genus Ecculiomphalus Portlock, 1843 FIGURE 9 Malayaspira Kobayashi, 1959. Rossospira Rohr, 1994. TYPE SPECIES.?Ecculiomphalus bucklandi Portlock, 1843. FKA.?Malayaspira rugosa Kobayashi, 1959: Setul Forma? tion (Late Arenig). LKA.?Ecculiomphalus bucklandi: Balclatchie Group (Mid? dle Caradoc (Harnagian)). ADDITIONAL ASSIGNED SPECIES.?Ecculiomphalus fredricus Raymond, 1908; Maclurina !annulata (Walcott, 1884); Mala? yaspira hintzei Rohr, 1984; Rossospira harrisae Rohr, 1994. REMARKS.?Ecculiomphalus includes species with broad rectangular or pentagonal aperture shapes and a strong periph? eral carina. Open coiling now diagnoses only derived members of the genus, and the frill-like peripheral band does not diag? nose any members of Eccyliopterus. Genus ^Asgardaspira, new genus FIGURE 9 TYPE SPECIES.?Lytospira yochelsoni Rohr, 1994. FKA.?Barnesella measuresae Rohr, 1994: Antelope Valley (Late Arenig). LKA.?Ophiletina aff. O. sublaxa of Rohr (1988) and Eccu? liomphalus !potteri (Rohr, 1988): Port Clarence Limestone (Ashgill). ADDITIONAL ASSIGNED SPECIES.?Lytospira gerrula Rohr, 1993; L. !norvegica (Koken, 1925). ETYMOLOGY.?After the mythological Norse serpent, re? flecting the open-coiled nature of most species. REMARKS.?This paraclade includes species retaining lentic? ular apertures, sharp peripheral bands, and moderately deep V- shaped sinuses. Primitive members are diagnosed by a strong basal carina, which forms a stubby frill at the base of the outer margin. Carrier-shell scars and a columellar lirum typify both Asgardaspira and Lytospira, but the retention of the sharp pe? ripheral band and sharp sinus separates Asgardaspira from Lytospira. Many of the species previously assigned to Lytospira are reassigned here. In addition, some poorly known Baltic species likely belong here. Genus Lytospira Koken, 1896 FIGURE 9 TYPE SPECIES.?Lytospira angelini (Lindstrom, 1884). FKA.?Lytospira angelini: Lower Gray Orthoceras Lime? stone (Late Arenig (Kundan)). LKA.?Lytospira aff. L. subrotunda (sensu Rohr, 1994): Port Clarence Limestone (Ashgill (Richmondian)). !Lytospira valida Koken, 1925: Lyckholm Formation (Late Ashgill (Porkuni)). ADDITIONAL ASSIGNED SPECIES.?Lytospira subrotunda (Ulrich and Scofield, 1897). REMARKS.?Lytospira is substantially reduced from previ? ous definitions and now includes only open-coiled species with shallow V-shaped to U-shaped sinuses and weak peripheral bands. Also, the aperture profile is round (owing to the rounder ramps and the greater projections of the right and left ramps), whereas that of Asgardaspira is lenticular (see above). Lytospira almost certainly includes several other Baltic species described by Koken (1925) and others; however, the available material was too poor for me to include those species here. See remarks on Pachystrophia below. Superfamily MACLURITOIDEA Fischer, 1885 REMARKS.?The Macluritoidea corresponds to the "macluri? toid" clade. As noted herein and elsewhere (e.g., Linsley and Kier, 1984), many of the species previously placed in this taxon likely were not gastropods. Species within this superfamily typically are diagnosed by a base formed from a posteriorly projecting inner margin, nearly planispiral to visually dextral coiling, a shallow V-shaped sinus, and a sharp thin peripheral 72 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY band located on top of the aperture. Another diagnostic charac? ter is the change in shape of the aperture over ontogeny, with the left side expanding differentially to produce a rhombohe- dral adult aperture from a lenticular juvenile one. Finally, a horn-shaped calcified operculum with a handle-like knob diag? noses Teiichispira and more-derived species. Genus ^Macluritella Kirk, 1927 FIGURES 8,10 IBridgeina Flower, 1968a. Prohelicotoma Flower, 1968b. TYPE SPECIES.?Macluritella stantoni Kirk, 1927. FKA.?Prohelicotoma uniangulata (Hall, 1847): Chepulte- pec Dolomite (Early Tremadoc (Gasconadian)). LKA.?Macluritella stantoni: Rochdale Run Formation (Early Arenig (Jeffersonian)). REMARKS.?Macluritella represents a primitive grade and paraclade of macluritoids. Although it is paraphyletic relative to later macluritoids, it appears to include a monophyletic clade of open-coiled, strongly septate species (e.g., M. stantoni and "Lytospira" gyrocera (Roemer)). Unfortunately, species other than M. stantoni are not well enough known to include in this analysis. If it should prove that those species do not belong to a Macluritella paraclade, then Macluritella should be expanded to include species presently assigned to Teiichispira. Genus ^Teiichispira Yochelson and Jones, 1968 FIGURE 10 Monitorella Rohr, 1994. TYPE SPECIES.?Teiichispira kobayashi Yochelson and Jones, 1968. FKA.?Maclurites !oceana (Billings, 1865): Watts Bight Formation (Late Tremadoc (Demingian)). LKA.?Undescribed species (initially described as "Ec? cyliopterus ornata" by E.O. Ulrich in an unpublished manu? script): Murfreesboro Limestone (Early Caradoc (Ashbian)). ADDITIONAL ASSIGNED SPECIES.?Monitorella auricula Rohr, 1994; Teiichispira odenvillensis Yochelson and Jones, 1968. REMARKS.?Teiichispira is a paraclade composed of species with juvenile whorls similar to that of adult Macluritella, but with apertures becoming more lenticular over ontogeny (with the long axis becoming parallel to the coiling axis). The diagno? sis for Teiichispira originally was based on opercula (Yochelson and Jones, 1968), but the definition provided herein is based en? tirely on shell features. Monitorella Rohr, 1994, a monotypic genus from the Antelope Valley Formation, represents a species with somewhat rounder whorls than other species and does not appear to represent a separate clade. Owing to the inherent diffi? culty of linking opercula to shells, it might be advisable to re? place Teiichispira with Monitorella in the future. Early Carado- cian specimens labeled but not published by E.O. Ulrich appear to belong to this genus, which extends the range of Teiichispira somewhat. This highly derived species from the Murfreesboro Formation is noteworthy for strongly crenulated growth lines. Genus ^Maclurites Le Sueur, 1818 FIGURE 10 TYPE SPECIES.?Maclurites magna Le Sueur, 1818. FKA.?Teiichispira sylpha (Billings, 1865): Aguantha For? mation (Early Arenigian (Cassinian)). LKA.?Maclurites expansa Koken, 1925: Bosnes Formation (Ashgill (Pirgu)). ADDITIONAL ASSIGNED SPECIES.?Maclurites crassus Ul? rich and Scofield, 1897; M. klamathensis Rohr, 1980. REMARKS.?Maclurites is distinguished from Teiichispira by (1) greater expansion of the lower left portion of the aper? ture throughout ontogeny (producing a rhombohedral aperture rather than a lenticular one), (2) the peripheral band being above the "umbilicus" rather than above the center of the aper? ture, (3) a flatter inner margin (yielding a flat base), and (4) complete visually dextral coiling, with (at most) only the base of the aperture coiling in a planispiral plane. The inclusion of T. sylpha renders Maclurites paraphyletic relative to both Maclu? rina and Palliseria (see remarks below). Genus Maclurina Ulrich and Scofield, 1897 FIGURE 10 TYPE SPECIES.?Maclurina manitobensis (Whiteaves, 1890). FKA.?Maclurites bigsbyi (Hall, 1861): Platteville Forma? tion (Early Caradoc (Black Riverian)). LKA.?Maclurina manitobensis: Bighorn Dolomite (Ashgill (Richmondian)). ADDITIONAL ASSIGNED SPECIES.?Maclurina logani Salter, 1859; Maclurites sedgewicki (Longstaff, 1924). REMARKS.?Rohr, Blodgett, and Furnish (1992) separated Maclurites and Maclurina based on the latter taxon possessing a strong revolving ornament. The analyses herein suggest that the ornate species do form a clade, so Maclurina is recognized herein. Species not analyzed herein, such as Maclurites kla- menthensis, leave Maclurites a paraclade. Genus Palliseria Wilson, 1924 FIGURE 10 Mitrospira Kirk, 1929.?Knight etal., 1960. Zhuozishanospira Yu, 1961. TYPE SPECIES.?Palliseria robusta Wilson, 1924. FKA.?Palliseria robusta: Antelope Valley Limestone (Late Arenig (Whiterockian Zone M)). LKA.?Palliseria robusta: Oil Creek Formation (Llanvirn). ADDITIONAL ASSIGNED SPECIES.?Mitrospira longwelli Kirk, 1929; Zhuozishanospira sinensis Yii, 1961. REMARKS.?Palliseria is a highly derived clade of visually sinistral shells. The clade is diagnosed by a rounded inner mar- NUMBER 88 73 gin that forms the lower portion of the outer margin and onto? genetic "increase" in translation accompanied by some de? crease in expansion yielding a somewhat cerithioid-form. These features, combined with expansion of the lower left por? tion of the aperture (present in ancestral Maclurites sylpha), produce counter-clockwise rotation of the aperture over ontog? eny. The monotypic Zhuozishanospira differs only in the pres? ence of ornament and is included within Palliseria in order to maintain monophyly. ?Genus Rousseauspira Rohr and Potter, 1987 TYPE SPECIES.?Rousseauspira teicherti Rohr and Potter, 1987. FKA.?Rousseauspira aff. R. teicherti Rohr and Potter, 1987: Klamath Mountains Limestone (Llandeilo). LKA.?Rousseauspira teicherti: Port Clarence Limestone (Ashgill (Richmondian)). REMARKS.?Rohr and Potter, 1987, recognized this genus based solely on opercula. When these opercula are associated with shells, it is likely that they will diagnose a distinct clade within either Maclurites or Maclurina (or possibly even Tei? ichispira). Until that time, the genus should be considered questionable. Superfamily fEuOMPHALOlDEA de Koninck, 1844 REMARKS.?With the likely exception of the Platycera- toidea, the Euomphaloidea corresponds to the "ceratopeatoid" clade described above. Early members of the family are diag? nosed by a sigma-shaped inner margin, a strong channeled basal carina, an adapically curved monolineate peripheral band, and increasing translation accompanying abapical rota? tion of the aperture over ontogeny. Some or all of these features are modified on more-derived members of the clade. LKA.?Helicotoma medfraensis Rohr and Blodgett, 1988: Telsitina Formation (Early(?) Llanvirn). ADDITIONAL ASSIGNED SPECIES (excluding operculum spe? cies).?Ceratopea buttsi (Yochelson and Bridge, 1957); C. hammsi (Stauffer, 1937); C. Haurentia (Billings, 1865); C. pyg- maea (Stauffer, 1937); Pararaphistoma lemoni Flower, 1968a. REMARKS.?Previous diagnoses of the genus have been based purely on operculum morphology, and the type species, Ceratopea keithi, is diagnosed by an operculum. This con? founds typologic categorizations of the genus. Also, some Cer? atopea opercula that are associated with shells are now trans? ferred to other genera. Ceratopea is paraphyletic relative to the rest of the Euomphaloidea. The species retained within the ge? nus, however, typically are diagnosed by lenticular apertures, strong adapically hooked peripheral bands, strong basal cari? nae, and deep strongly curved sinuses. Genus Bridgeites Flower, 1968a FIGURE 11 TYPE SPECIES.?Bridgeites discoideus Flower, 1968a. FKA.?Bridgeites !disjuncta (Billings, 1865): Oxford For? mation (Late Tremadoc (Demingian)). LKA.?Bridgeites planodorsalis (Cullison, 1944): Smith- ville Formation (Early Arenig (Cassinian)). ADDITIONAL ASSIGNED SPECIES.?Bridgeites supraconvexa (Cullison, 1944). REMARKS.?Early Ordovician species from North America previously classified as Lesueurilla by Cullison (1944) and others belong to this clade. The clade is diagnosed by nearly planispiral coiling, very strong basal carinae, and decreasing curvature over ontogeny that yields open-coiling (sensu Yoch? elson, 1971) in later stages. Family f RAPHISTOMATIDAE Koken, 1896 REMARKS.?This family corresponds to the "scalitine" clade diagnosed and defined above, plus the basal members of the Euomphalidae. Post-Silurian genera typically assigned to the Raphistomatidae (e.g., Arizonella Stoyanow, 1948, Buechelia Schliiter, 1894, Denayella Blodgett and Johnson, 1992, Scali- tina Spriesterbach, 1919, and Wisconsinella Blodgett, 1987) appear to be related to eotomarioid taxa, such as Phaner? otrema, rather than to raphistomatids (Wagner, in prep.). Thus, they are removed from the Raphistomatidae. Genus ^Ceratopea Ulrich, 1911 FIGURES 11-13 Proliospira Flower, 1968a. TYPE SPECIES.?Ceratopea keithi Ulrich, 1911. FKA.?Ceratopea canadensis (Billings, 1865): Fort Ann Limestone (Late Tremadoc (Demingian)). Genus Orospira Butts, 1926 FIGURE 11 TYPE SPECIES.?Orospira bigranosa Butts, 1926. FKA.?Orospira gainesvillensis Cullison, 1944: Rich Foun? tain beds, Jefferson Formation (Early Arenig (Jeffersonian)). LKA.?Orospira bigranosa: Cotter Formation (Early Arenig (Cassinian)). REMARKS.?This is essentially an ornate version of Cerato? pea, including at most only two species. Most of the other "species" (see, e.g., Cullison, 1944) are simply variants of O. bigranosa. Genus Raphistoma Hall, 1847 FIGURE 13 TYPE SPECIES.?Raphistoma striata (Emmons, 1842). FKA.?Helicotoma gubanovi (Rohr, 1994): Antelope Valley Formation (Late Arenig (Whiterockian Zone M)). 74 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY LKA.?Raphistoma striata: Valcour Formation (Late Lland- eilo). ADDITIONAL ASSIGNED SPECIES.?Scalites alaskensis (Rohr in Rohr, Blodgett, and Furnish, 1992). REMARKS.?Although most of the species within the "ra? phistomatids" were once classified as Raphistoma, the current definition includes only a very restricted set of highly derived species. The clade is diagnosed in part by a slit and a dull broad peripheral band. Finally, although the stratigraphic record of first appearances within this group appears to run opposite to the cladogram, it should be noted that there is no Late Arenigian record for the eastern United States (which contains the most primitive members of the clade). Thus, there is no reason to think that R. striata evolved prior to the Llanvirn. Genus ^Scalites Emmons, 1842 FIGURE 13 TYPE SPECIES.?Scalites angulatus Emmons, 1842. FKA.?Palaeomphalus giganteus Kobayashi, 1959: Unkaku Beds (Late Arenig). LKA.?Raphistoma tellerensis (Rohr, 1988): Port Clarence Limestone (Ashgill (Richmondian)). ADDITIONAL ASSIGNED SPECIES.?Raphistoma peracuta (Ulrich and Scofield, 1897); Scalites katoi (Kobayashi, 1934). REMARKS.?Scalites is expanded to include most of the spe? cies previously classified in Raphistoma. The type species, S. angulatus, is not typical of the paraclade and shares several sy? napomorphies with Raphistoma. It would be more consistent with the phylogeny to use Scalites for the Raphistoma genus as defined above, but this would require then creating a new ge? nus for the paraclade. Keeping S. angulatus in the same genus as other Scalites species does not distort diversity dynamics (i.e., obscuring any originations or extinctions), so I chose to keep the traditional genera. Members assigned to the genus are diagnosed by flattened right ramps that (owing to the overall orientation of the aperture) are nearly perpendicular to the coil? ing axis, strong adapically hooked peripheral bands, and strong channeled basal carina. Family HOLOPEIDAE Wenz, 1938 REMARKS.?The new definition of the Holopeidae differs drastically from those presented by Wenz (1938) or Knight et al. (1960). Those definitions appear to be highly polyphyletic, and Holopea represents the only trochoid-like genus within the present definition of the family. This is unfortunate, as it means that the genus is very atypical of the eponymous family. Al? though taxonomic priority rules do not apply strictly to su- prageneric taxa, it seems preferable to maintain as much of the old taxonomic structure as possible. The family is diagnosed by U-shaped sinuses, thus including many genera previously as? signed to the Sinuopeidae, and the loss of the peripheral band in all but the least derived members (e.g., Raphistomina species). Genus Raphistomina Ulrich and Scofield, 1897 FIGURE 13 TYPE SPECIES.?Raphistomina lapicida (Salter, 1859). FKA.?Raphistomina lapicida: Lower Antelope Valley Limestone (Late Arenig (Whiterockian, Zone M)). LKA.?Raphistomina rugata Ulrich and Scofield, 1897: Port Clarence Limestone (Ashgill (Richmondian)). ADDITIONAL ASSIGNED SPECIES.?Liospira americana (Billings, 1865); Raphistomina aperta Wilson, 1921; R.fissu- rata Steele and Sinclair, 1971. REMARKS.?The initial analyses (e.g., Wagner, 1995d) sug? gested that a ghost lineage linked Early Caradocian (and possi? ble Llandeilian) Raphistomina with Llanvirnian Pachystro? phia. (Pre-Llanvirnian reports of Raphistomina had invariably proved to be Ceratopea.) Since then, Rohr (1996) described Raphistomina from the Late Arenig and Early Llanvirn of the western United States. This represents one of the first confir? mations of a ghost taxon estimate known to the author. The somewhat problematic specimens of Liospira americana are assigned here provisionally; it appears to represent a primitive grade of Raphistomina that retains a V-shaped sinus rather than a U-shaped one. The species also lacks any of the important sy? napomorphies of Liospira. Genus ^Pachystrophia Perner, 1903 FIGURES 13,14 TYPE SPECIES.?Pachystrophia devexa (Eichwald, 1859). FKA.?Pachystrophia devexa: Asirian Limestone (Late Llanvirn (Asirian)). LKA.?Devonian species classified as "Lytospira" are known through the Late Eifelian. ADDITIONAL ASSIGNED SPECIES.?Lytospira subuloides Barrande in Perner, 1903; L. triquestra (Lindstrom, 1884); Pachystrophia contigua (Ulrich and Scofield, 1897); P. got- landica (Lindstrom, 1884). REMARKS.?Pachystrophia is used herein to label an inor? nate paraclade of species with broad U-shaped sinuses and no peripheral bands. Post-Ordovician species previously classi? fied as Lytospira are removed to Pachystrophia. Late Silurian and younger species are diagnosed by carrier-shell scars. Open-coiled Pachystrophia can be distinguished from Lytospira by the very broad whorls (produced by the very high angles of right and left ramp projections) with the long axis perpendicular to the coiling axis owing to the orientation of the whorl. Lytospira features a nearly circular aperture. In ad? dition, Pachystrophia has a much broader U-shaped sinus than does Lytospira, and Pachystrophia has no trace of a peripheral band. NUMBER 88 75 Genus Sinutropis Perner, 1903 FIGURE 14 TYPE SPECIES.?Sinutropsis esthetica Barrande in Perner, 1907. FKA.?Pachystrophia spiralis Rohr, 1980: Kangaroo Creek Formation (Caradoc (?Harnagian)). LKA.?Sinutropis esthetica: Kopanina Formation (Late Ludlow (Ludfordian)). ADDITIONAL ASSIGNED SPECIES.?Euomphalus tubus (Lind? strom, 1884); Sinutropsis !esthetica (sensu Rohr, 1988). REMARKS.?Sinutropis labels a clade of ornate species de? rived from Pachystrophia. The correlations for beds containing the earliest known species are uncertain; however, quantitative analysis of molluscan assemblages using Appearance Event Or? dination (Alroy, 1994a) suggests that those Kangaroo Creek Formation beds correspond to the Hamagian (Wagner, in prep.). Genus Umbospira Perner, 1903 Horiostomella Perner, 1903. Sellinema Perner, 1903. Turbomaria Perner, 1903. TYPE SPECIES.?Umbospira nigricans Barrande in Perner, 1907. FKA.?Umbospira yochelsoni Peel, 1977: Ross Brook For? mation (Late Llandovery (Telychian)). LKA.?Several species known from the Kopanina Forma? tion (Late Ludlow (Ludfordian)). ADDITIONAL ASSIGNED SPECIES.?Horiostomella otiosa Barrande in Perner, 1907; Sellinema dive Barrande in Perner, 1907; Turbomaria sepulta Barrande in Perner, 1907. REMARKS.?Umbospira represents an apparent clade of largely Gondwanan species derived from Pachystrophia. The genus is diagnosed by a very shallow and fairly narrow sinus. If Leptozone belongs to this clade, then the range of the genus extends through the Early Devonian (Pragian). The numerous junior synonyms of the genus reflects the taxonomy of Bar? rande and Perner in Perner (1903, 1907), which severely over- splits gastropods at both the species and genus level. My exam? inations suggest that many of Perner's genera comprise one recognizable species, with the additional species names within those genera usually reflecting intraspecific variants. This is partially rectified here. Genus Holopea Hall, 1847 FIGURE 14 Litiopsis Salter, 1866. TYPE SPECIES.?Holopea symmetrica Hall, 1847. FKA.?Holopea insignis Ulrich and Scofield, 1897: Lowville Formation (Early Caradoc (Black Riverian)). LKA.?Holopea has been recorded as late as the Early Ser- pukhovian. Given the featureless nature of the genus, however, its present definition likely is very polyphyletic. Holopea palu- diniformis Hall, 1847, and H. parvula Ulrich and Scofield, 1897, represent the latest species that certainly belong to the ge? nus, being known from the Cobourg Formation (Late Caradoc, latest Shermanian/early Edenian). Holopea vermiculosa Bar? rande in Perner, 1907, from the Bohladec Formation (Late Caradoc) represents a roughly contemporaneous species. None of the species identified as "Holopea" by Lindstrom belong to this genus. ADDITIONAL ASSIGNED SPECIES.?Holopea ampla Billings, 1865; H. pyrene Billings, 1865; H. rotunda Ulrich and Scofield, 1897. REMARKS.?Holopea is a highly derived clade of "raphisto? matids." Unfortunately, the derivations involve the reduction of shell features, which limits the number of diagnostic characters (as opposed to diagnostic absences). Many shells that are fea? tureless owing to poor preservation have been inappropriately assigned to Holopea. There do not appear to be valid members of this genus from the Ashgill or later. Family LESUEURILLIDAE, new family REMARKS.?This family corresponds to the "lesueurilline" clade diagnosed above. The most noteworthy features include a sharp peripheral band, often strongly developed, a deep V- shaped sinus culminating in a short notch (producing distinc? tive lunulae), and distinct ontogenetic changes from a "Raphis- toma"-\ike. early morphology to a "Maclurites"-like (e.g., Lesueurilla) morphology. One derived clade shows the oppo? site pattern, however, by producing "Liospira"-like late mor? phologies (i.e., Pararaphistoma). Genus ^Eccyliopterus Remele, 1888 FIGURE 12 TYPE SPECIES.?Eccyliopterus alatus (Roemer, 1876). FKA.?Lesueurilla declivis Koken, 1925: Asaphuskalk (Early Arenig (Volkov)). LKA.?Eccyliopterus owenanus (Meek and Worthen, 1866): Fusispira bed, Prosser Formation (Middle Caradoc (Kirkfield- ian)). ADDITIONAL ASSIGNED SPECIES.?Eccyliopterus !princeps Koken, 1925; E. regularis Koken, 1925. R E M A R K S . ? T h i s paraclade comprises open-coi led "lesueurillines," with a derived clade featuring a very strong frill-like and adapically curved peripheral band. The con? founding of this genus with Ecculiomphalus is puzzling, as the two share very few features. These differences include a num? ber of characters involving the orientation and shape of differ? ent parts of the aperture (resulting in lenticular apertures for Eccyliopterus species and rhombohedral apertures with the in? ner margin forming a base for Ecculiomphalus species), the peripheral band (which is very strong with a pronounced adapical hook on Eccyliopterus species but weak and nearly 76 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY straight on Ecculiomphalus species), and the sinus (which curves back strongly and culminates in a short notch on Ec? cyliopterus species but is fairly shallow and V-shaped on Ec? culiomphalus species). Genus ^Lesueurilla Koken, 1898 FIGURE 12 TYPE SPECIES.?Lesueurilla infundibula (Koken, 1896). FKA.?Lesueurilla prima Barrande in Perner, 1907: Niveau G (Early Arenig (Klabava)). LKA.?Eccyliopterus beloitensis Ulrich and Scofield, 1897: Platteville Formation (Early Caradoc (Black Riverian)). ADDITIONAL ASSIGNED SPECIES.?Eccyliopterus louder- backi Endo, 1932. REMARKS.?Usage of Lesueurilla is now restricted to a par? aclade of nearly planispiral species with lenticular apertures oriented nearly parallel to the coiling axis and curved inner margins forming rounded bases. Lesueurilla and derivatives represent the earliest clade diagnosed by a slit, which is formed by an extension of the sinus and appears later in ontog? eny on primitive members. Some derived Lesueurilla appear to have slits throughout ontogeny. Genus Mestoronema, new genus FIGURE 12 TYPE SPECIES.?Lesueurilla bipatellare Koken, 1925. FKA.?Lesueurilla marginalis Koken, 1925: Revel Forma? tion (Late Llanvirn (Asirian)). LKA.?Lesueurilla acutangulata Koken, 1925: Wesen- berger Shale (Late Caradoc (Vormsi)). ADDITIONAL ASSIGNED SPECIES.?Lesueurilla scotica Longstaff, 1924; L. spiralis Koken, 1925; L. aff. L. marginalis (in Rohr, 1980). ETYMOLOGY.?After the ruler of the intelligent evil gastro? pods from the world's longest running science fiction serial, Doctor Who. REMARKS.?This new genus recognizes a monophyletic group of close-coiled, generally small lesueurillids with strong, blunt and angled peripheral bands, and short slits that are ap? parently present throughout ontogeny and are produced as ex? tensions of the sinus. Species can be distinguished from Lesueurilla by the strong angular umbilical carina, which sometimes creates as much of a profile as the periphal band. Genus Pararaphistoma Vostokova, 1955 FIGURE 12 Climacoraphistoma Vostokova, 1955.?Knight etal., 1960. TYPE SPECIES.?Pararaphistoma qualteriata (Schlotheim, 1820). FKA.?Climacoraphistoma vaginati (Koken, 1925): Niveau G (Early Arenig (Klabava)). LKA.?Climacoraphistoma spiralis (Koken, 1925): Keila Formation (Middle Caradoc (Keila)). ADDITIONAL ASSIGNED SPECIES.?Climacoraphistoma damesi (Koken, 1925); Pararaphistoma schmidti (Koken, 1925). REMARKS.?This derived genus is diagnosed by the devel? opment of a slit on more adult whorls and an ontogenetic rota? tion of the aperture and an increase in translation that produces a Ceratopea-like adult morphology from a Lesueurilla-hke ju? venile morphology. Family THELICOTOMIDAE Wenz, 1938 REMARKS.?The Helicotomidae paraclade corresponds to the Ordovician members of the "helicotomid" clade; however, its extinction at the end of the Ordovician coincides with the true extinction of at least two and possibly three relatively primitive subclades. Nearly all members of this family and its derivatives are diagnosed by an anteriorly produced aperture with a distinct sigmoidal shape when viewed from the side (owing to the anterior angling of the base of the alveozone). Genus fLophonema Ulrich in Purdue and Miser, 1916 FIGURES 11,15 Polehemia Cullison, 1944. Walcotloma Rohr, 1994. TYPE SPECIES.?Lophonema peccatonica Ulrich in Purdue and Miser, 1916. FKA.?Ceratopea unguis Yochelson and Bridge, 1957: Watts Bight Formation (Late Tremadoc (Demingian)). LKA.?Walcottoma frydai Rohr, 1994: Antelope Valley For? mation (Llanvirn (Whiterockian Zone N)). ADDITIONAL ASSIGNED SPECIES.?Polehemia taneyensis Cullison, 1944. REMARKS.?Lophonema represents the basal paraclade of the helicotomids. Typical features include sharp carina on ei? ther side of the peripheral band and an anteriorly projected ap? erture. The monophyletic portion of the paraclade includes some of the earliest ornate species. Genus f Linsleyella Rohr, 1980 Ellisella Rohr, 1980. [Not Gray, 1858.] Yochelsoniella Rohr and Huddleston, 1982 [replacement name for Ellisella]. TYPE SPECIES.?Linsleyella johnstoni Rohr, 1980. FKA.?Linsleyella greggi (Rohr, 1980): Kangaroo Creek Formation (Llandeilo? Caradoc (Harnagian)). LKA.?Linsleyella johnstoni: Kangaroo Creek Formation (Llandeilo? Caradoc (Harnagian)). ADDITIONAL ASSIGNED SPECIES.?Helicotoma spinosa (Salter, 1859). NUMBER 88 77 REMARKS.?This genus is represented only by poor speci? mens; however, it is diagnosed by strong, spinose ornament and strong nodular lira on either side of the peripheral band. The stratigraphic position of the Kangaroo Creek beds in which two of the species are found is uncertain; however, quantitative analyses suggest an Early Caradocian age. This concurs well with the better constrained Black Riverian age for the basal member of the clade, L. spinosa. Genus ^Helicotoma Salter, 1859 FIGURE 15 TYPE SPECIES.?Helicotoma planulata Salter, 1859. FKA.?Helicotoma tennesseensis Ulrich and Scofield, 1897: Murfreesboro Limestone (Early Caradoc (Ashbian)). LKA.?Helicotoma robinsoni Rohr, 1988, and H. blodgetti Rohr, 1988: Port Clarence Limestone (Ashgill (Richmondian)). ADDITIONAL ASSIGNED SPECIES.?Helicotoma planula- toides Ulrich and Scofield, 1897. REMARKS.?The definition of Helicotoma presented herein is restricted to a paraclade of species with a strong adapically curved peripheral band and a round, anteriorly projected aper? ture with a broad dull carina on the border of the alveozone and base. Genus Palaeomphalus Koken, 1925 FIGURE 15 TYPE SPECIES.?Palaeomphalus gradatus Koken, 1925. FKA.?Helicotoma olsoni Rohr, 1980 (? = Helicotoma griffmora Rohr, 1980): Kangaroo Creek Formation (Caradoc (?Harnagian)). LKA.?Palaeomphalus !gradatus: Borkholm Formation (Late Ashgill (Porkuni)). ADDITIONAL ASSIGNED SPECIES.?Euomphalus obtusangu- lus Koken, 1925. REMARKS.?Palaeomphalus is restored here to label a clade of helicotomids with obtuse weak peripheral bands and highly reduced swellings on the alveozone, but with sharper circum- basal carina than found on Helicotoma species. An apparently undescribed species from the Thraive Limestone (Ashgillian) belongs here. ADDITIONAL ASSIGNED SPECIES.?Ophiletina angularis Ul? rich and Scofield, 1897; Oriostoma bromidensis Rohr and Johns, 1990. REMARKS.?Ophiletina represents a monophyletic genus di? agnosed by the characters discussed in the text. Family fEuOMPHALiDAE de Koninck, 1881 REMARKS.?The Euomphalidae represents a paraclade that includes the Silurian species corresponding to traditional diag? noses of the family (e.g., Knight et al., 1960). Ordovician spe? cies previously assigned to Euomphalopterus (reassigned herein to Boucotspira) represent the only Ordovician members of the family, and render the family paraphyletic relative to the Pseudophoridae and Anomphalidae. Although the Silurian as? semblage within the family is monophyletic, preliminary anal? yses suggest that the Euomphalidae as defined herein include the ancestors of Devonian and later taxa presently assigned to both the Omphalotrochidae and the Omphalocirridae. Genus ^Boucotspira Rohr, 1980 FIGURES 15, 16,19 TYPE SPECIES.?Boucotspira ftmbriata Rohr, 1980. FKA.?Boucotspira aff. B. ftmbriata: Lower Antelope Val? ley Limestone (Late Arenig (Whiterockian Zone M)). LKA.?Streptotrochus incisus (Lindstrom, 1884) Slite Bed (Unit G) (Wenlock (Sheinwoodian)). ADDITIONAL ASSIGNED SPECIES.?Euomphalopterus !carin- iferus Koken, 1925; E. !ordovicius Longstaff, 1924; E. aff. E. ordovicius Longstaff, 1924; Streptotrochus lamellosus (Lind? strom, 1884); Streptotrochus! visbeyensis (Lindstrom, 1884); Trochonemella antelopensis Rohr, 1996. REMARKS.?This paraclade is strongly convergent on the lo- phospirid genera Trochonema and Trochonemella. Trochone? mella antelopensis is either a variant of B. ftmbriata or a closely related slit-bearing species. Boucotspira includes the Late Ordovician species that were previously classified as Eu? omphalopterus. Silurian species retaining strong right and left ramp carina are retained within Boucotspira. The paraclade is diagnosed by these prominent right and left carina, with the left (alveozone) carina usually stronger than the right. Genus Ophiletina Ulrich and Scofield, 1897 FIGURE 15 TYPE SPECIES.?Ophiletina sublaxa Ulrich and Scofield, 1897. FKA.?Ophiletina sublaxa: Murfreesboro Limestone (Early Caradoc (Ashbian)) or Oriostoma bromidensis Rohr and Johns, 1990: Bromide Formation (Early Caradoc (Ashbian)). LKA.?Ophiletina sublaxa: Fusispira Beds, Prosser Forma? tion (Middle Caradoc (Kirkfieldian)). Genus ^Euomphalopterus Roemer, 1876 FIGURES 16-18 Pleuromphalus Perner, 1903.?Knight et al., 1960. Bathmopterus Kirk, 1928.?Knight etal., 1960. ISiskyouspira Rohr, 1980. TYPE SPECIES.?Euomphalopterus alatus (Wahlenberg, 1821). FKA.?Euomphalopterus subcarinatus (Lindstrom, 1884): Solvik Formation (Middle Llandovery (Aeronian)). 78 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY LKA.?Euomphalopterus has been recorded as late as the Emsian. ADDITIONAL ASSIGNED SPECIES.?Euomphalopterus aliger Barrande in Perner, 1903; E. praetextus (Lindstrom, 1884); E. togatus (Lindstrom, 1884); E. undulans (Lindstrom, 1884). REMARKS.?Euomphalopterus represents the basal para? clade of the Euomphalidae. It is diagnosed by a very strong al? veozone carina that often produces a frill, which is lost or highly modified in other euomphalid genera. The exceptionally strong development of this feature in the type species appears to be unique. Genus Spinicharybdis Rohr and Packard, 1982 FIGURE 17 TYPE SPECIES.?Spinicharybdis wilsoni Rohr and Packard, 1982. FKA.?!Euomphalopterus frenatus (Lindstrom, 1884): Lower Visby Formation (Late Llandovery (Telychian)). LKA.?Spinicharybdis wilsoni: Barrow Inlet Formation (Late Ludlow (Ludfordian)). ADDITIONAL ASSIGNED SPECIES.?Spinicharybdis billingsi Rohr and Packard, 1982; S. ehlora (Billings, 1865). REMARKS.?Spinicharybdis is diagnosed by strong tubes at the base of the lower ramp. These apparently were derived from the frill of Euomphalopterus, although some alternative phylogenetic scenarios are discussed above in "Results." Genus ^Poleumita Clarke and Ruedemann, 1903 FIGURES 16-18 ?Kiaeromphalus Peel and Yochelson, 1976. 1 Offleyotrochus Peel, 1979. TYPE SPECIES.?Poleumita discors (Sowerby, 1814). FKA.?Poleumita discors: Lower Visby Formation (Late Llandovery (Telychian)). LKA.?Poleumita discors: Upper Ludlow Formation (Late Ludlow (Ludfordian)). (Poleumita are reported as late as the Emsian, but the veracity of these assignments cannot be evalu? ated herein). ADDITIONAL ASSIGNED SPECIES.?Poleumita alata (Lind? strom, 1884); P. granulosa (Lindstrom, 1884). REMARKS.?Poleumita represents a paraclade containing the ancestors of important euomphalids of the late Paleozoic, such as Euomphalus and Straparollus. A thick, strip-like parietal in- ductura projecting well in front of the aperture diagnoses the genus as well as some descendants (e.g., Nodonema and Cen- trifugus). Poleumita differs from Euomphalus by the retention of strong ornament and from Straparollus by the retention of both ornament and a peripheral band. Kiaeromphalus and Offleyotrochus are tentatively placed within Poleumita. Both genera are known from a single spe? cies, and little of the overall characteristics are known, espe? cially for Offleyotrochus. More and better material are required before separate generic assignments can be justified. Genus Nodonema Linsley, 1968 FIGURE 18 TYPE SPECIES.?Nodonema granulata Linsley, 1968. FKA.?Poleumita rugosa (Lindstrom, 1884): Lower Visby Formation (Late Llandovery (Telychian)). LKA.?Nodonema granulata: Anderdon Limestone (Eifel- ian). ADDITIONAL ASSIGNED SPECIES.?Poleumita octavia (M'Coy, 1851). REMARKS.?Nodonema represents a high-spired variant of Poleumita, with strong nodular ornament. It is very similar to the contemporaneous lophospirid genus Globonema, but it dif? fers in its slightly sigmoidal aperture, thick parietal inductura strip, and nodular ornament. Many post-Silurian species previ? ously assigned to Gyronema might belong here. Genus Centrifugus Bronn, 1834 FIGURE 18 TYPE SPECIES.?Centrifugus planorbis Bronn, 1834. FKA.?Centrifugus planorbis: Upper Hemse Beds (Early Ludlow (Gorstian)). LKA.?Centrifugus planorbis: Lower Ludlow Formation (Early Ludlow (Gorstian)). REMARKS.?This is one of the few monotypic genera main? tained herein. It might be better to lump this genus with Po? leumita (in which case, Centrifugus would represent the senior synonym). Given the morphologic disparity between this spe? cies and its closest relatives, however, I keep them separate for now. Genus Euomphalus Sowerby, 1814 FIGURE 18 TYPE SPECIES.?Euomphalus pentangulatus Sowerby, 1814. FKA.?Euomphalus walmstedti Lindstrom, 1884: Wenlock Limestone (Late Wenlock (Gleedonian)). LKA.?Euomphalus is recorded as late as the Norian (e.g., Yin and Yochelson, 1983). REMARKS.?The definition presented herein is highly abbre? viated, as Euomphalus is one of the most commonly described genera of the Paleozoic. Whether it represents a cohesive phy? logenetic unit requires further investigation. Euomphalus al? most certainly represents a phylogenetic unit distinct from Straparollus (pers. obs.), so there is no reason to consider the former a subgenus of the latter. NUMBER 88 79 Genus Straparollus de Montfort, 1810 FIGURE 18 TYPE SPECIES.?Straparollus dionysii de Montfort, 1810. FKA.?Straparollus paveyi Foerste, 1924 (?= S. bohemicus Barrande in Perner, 1907): Racine Dolomite (Early Ludlow (Gorstian)). LKA.?Straparollus is recorded as late as the Guadalupian. REMARKS.?Like Euomphalus, Straparollus is a commonly described genus; however, it seems highly unlikely that it pres? ently constitutes a cohesive phylogenetic unit (pers. obs.). Genus Micromphalus Knight, 1945 REMARKS.?Tassell (1979) assigned a Late Silurian species to this genus but did not specify the species or formation. This genus might belong to the Pseudophoridae. Family ANOMPHALIDAE Wenz, 1938 REMARKS.?This monophyletic family corresponds to the "anomphalides" described above in "Results." In addition to the genera included herein and elsewhere (e.g., Knight et al., 1960), some genera assigned to the Omphalotrochidae (e.g., Labrocuspis Kase, 1989) might belong here. It appears, how? ever, that true omphalotrochids likely evolved elsewhere within the Euomphalidae (Erwin, in prep.). The family is diag? nosed by a U-shaped sinus located high on the aperture and a very strong lirum on the inner margin, which frequently pro? duces a callus that fills the umbilicus. Genus ^Trochomphalus Koken, 1925 FIGURE 16 Grantlandispira Peel, 1984a. TYPE SPECIES.?Trochomphalus !dimidiatus (Koken, 1896). FKA.?Trochomphalus !dimidiatus: Borkholm Formation (Ashgill (Pirgu)). LKA.?Grantlandispira christei Peel, 1984a: Offley Island Formation (Late Llandovery (Telychian)). ADDITIONAL ASSIGNED SPECIES.?Straparollina aff. S. circe of Rohr (1988). REMARKS.?It is not clear that this genus represents a para? clade, so it might prove better to consider this a junior syn? onym of Pycnomphalus. Conversely, given that it certainly spans through the poorly sampled earliest Silurian, it is possi? ble that additional sister species of T. christei exist, which would justify reinstatement of Grantlandispira. Genus Pycnomphalus Lindstrom, 1884 FIGURE 16 INematotrochus Perner, 1903. IPycnotrochus Perner, 1903. Turbocheilus Perner, 1903. llsfarispira Gubanov et al., 1995. TYPE SPECIES.?Pycnomphalus obesus Lindstrom, 1884. FKA.?Pycnomphalus acutus Lindstrom, 1884: Lower Visby Formation (Late Llandovery (Telychian)). LKA.?Pycnomphalus inflatus Barrande in Perner, 1903 (? = P. obesus) Pridoli Formation (Pridoli). ADDITIONAL ASSIGNED SPECIES.?Wematrochus concu- rens Barrand in Perner, 1907; Turbocheilus immaturum (Bar? rande in Pemer, 1903). REMARKS.?Pycnomphalus clearly is paraphyletic relative to later anomphalids; as such, its last appearance must be con? sidered tentative pending additional systematic work on the anomphalids. The genus is diagnosed by a callus filling (or nearly filling) the umbilicus and by the loss of the peripheral band. All species but the most primitive lack the prominent al? veozone. Pycnotrochus (Kopanina Formation, Late Ludlow) might represent basal members of Anomphalus rather than de? rived Pycnomphalus, but too little material is available to ver? ify this. Isfarispira appears to retain the basic synapomorphies of Pycnomphalus, although it clearly has a derived ontoge? netic change in the basic coiling parameters. Gubanov et al. (1995) suggested that other species might be assignable to Is? farispira; if so, then the genus should be maintained as a sepa? rate entity. Family PSEUDOPHORIDAE Miller, 1889 REMARKS.?The family corresponds to the "pseudophoride" clade diagnosed above in "Results." Most of the middle Paleo? zoic gastropods previously classified in the Trochina belong to this family. As extant trochoids are vetigastropods, it is highly unlikely that pseudophorids and trochoids are closely related. The family is diagnosed by strongly tangential apertures pro? duced by a greatly reduced sinus and anterior production of the base, a strongly sigmoidal aperture, and a strong inclination of the aperture. Genus Pseudophorus Meek, 1873 FIGURE 19 TYPE SPECIES.?Pseudophorus antiquus (Meek, 1872). FKA.?Discordichilus kolmodini (Lindstrom, 1884): Con- chidium Bed, Slite Formation (Early Wenlock (Sheinwood- ian)). LKA.?Yochelson and Saunders (1967) record this species from Toumaisian strata. ADDITIONAL ASSIGNED SPECIES.?Pseudophorus profundus Lindstrom, 1884. REMARKS.?Pseudophorus likely represents a paraclade rel? ative to several post-Silurian taxa. More detailed systematic work is necessary to ascertain whether the paraclade actually extends into the Carboniferous. 80 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY Genus Pseudotectus Perner, 1903 FIGURE 19 Planitrochus Pemer, 1903. Palaeonustus Perner, 1903. TYPE SPECIES.?Pseudotectus comes Barrande in Perner, 1907. FKA.?Discordichilus kolmodini (Lindstrom, 1884): Con- chidium Bed, Slite Formation (Early Wenlock (Sheinwood- ian)). LKA.?Pseudotectus comes: Koneprussy Formation (Pra- gian). Blodgett, Rohr, and Boucot (1988) reported Planitrochus from "Lockhovian-Emsian" aged material from the MacKenzie Mountains. Linsley (1979) recorded Eifelian Planitrochus but did not make reference to particular specimens or formations. ADDITIONAL ASSIGNED SPECIES.?Pseudotectus amicus Barrande in Perner, 1907. REMARKS.?Pseudotectus comprises (in part) two mono? typic genera diagnosed by Perner (1907), Planitrochus and Palaeonustus. It is diagnosed by the loss of the sinus, the loss of the peripheral band and right ramp, and a strong thick alveo? zone carina that produces a weak frill. Genus ^Discordichilus Cossmann, 1918 FIGURE 19 ?Umbotrochus Pemer, 1907. Siluriphorus Cossmann, 1918. TYPE SPECIES.?Discordichilus mollis (Lindstrom, 1884). FKA.?Siluriphorus gotlandicus (Lindstrom, 1884): Lower Visby Formation (Early Wenlock (Sheinwoodian)). LKA.?Dischordichilus mollis: Hemse Marl (Early Ludlow (Gorstian)); !Umbotrochus asperus (Barrande in Perner, 1903): Kopanina Formation (Late Ludlow (Ludfordian)). ADDITIONAL ASSIGNED SPECIES.?Dischordichilus dalli (Lindstrom, 1884); Siluriphorus undulans (Lindstrom, 1884). REMARKS.?This paraclade includes the ancestors of Strep? totrochus. In order to make Discordichilus a paraclade, species previously assigned to Siluriphorus now belong to Discordichi? lus. The paraclade appears to become truly extinct in the Lud? low, but Eifelian species have been assigned to Siluriphorus. Umbotrochus is represented by one poorly preserved species, but it is very similar to D. mollis. If it is related, then the range of the genus is extended slightly. Genus Hystricoceras Jahn, 1894 FIGURE 19 TYPE SPECIES.?Hystricoceras spinosum Jahn, 1894. FKA.?Hystricoceras astraciformis (Lindstrom, 1884): Lower Visby Formation (Early Wenlock (Sheinwoodian)). LKA.?Hystricoceras spinosum: Kopanina Formation (Late Ludlow (Ludfordian)). REMARKS.?This genus is diagnosed by distinctive spines on the alveozone-base periphery. Devonian taxa, such as As- tralites, might have evolved from within Hystricoceras; if so, then it is possible that the extinction of the genus is later than reported herein. Genus Streptotrochus Perner, 1903 FIGURE 19 Perneritrochus Cossmann, 1908. TYPE SPECIES.?Streptotrochus venalis (Barrande in Perner, 1903). FKA.?Streptotrochus incisus (Lindstrom, 1884): Lower Visby Formation (Early Wenlock (Sheinwoodian)). LKA.?Streptotrochus carinatus (Barrande in Perner, 1907): Koneprussy Formation (Pragian). Blodgett, Rohr, and Boucot (1988) reported Perneritrochus from "Lockhovian-Emsian" aged material from the MacKenzie Mountains. ADDITIONAL ASSIGNED SPECIES.?Streptotrochus lundgreni (Lindstrom, 1884). REMARKS.?Streptotrochus is used herein to label a para? phyletic group diagnosed by the characters discussed above in "1.3.2.2.3 Pseudophorides." Perneritrochus differs by the nearly complete elimination of the sinus. It is included within Streptotrochus to maintain monophyly. Genus Elasmonema Fischer, 1885 TYPE SPECIES.?Elasmonema bellatula (Hall, 1861). FKA.?Elasmonema imitator (Hall and Whitfield, 1872): Racine Dolomite (Late Wenlock (Gleedonian)). LKA.?Linsley (1979) reported a Frasnian occurrence for this genus. REMARKS.?Preliminary analyses of Devonian members of this genus place it as a sister taxon to Discordichilus mollis. The Silurian specimens that I have examined corroborate this hypothesis. Suborder MURCHISONIINA Cox and Knight, 1960 REMARKS.?The Murchisoniina now include most of the genera traditionally placed in the Pleurotomariina. The Murchisoniina also include the earliest caenogastropods sensu lato and the ancestors of more-derived gastropods, such as opisthobranchs. It seems unlikely that most malacologists will accept ranking those taxa below the suborder (the preceding examples have been accorded ordinal or class level). Thus, fu? ture work likely will render the murchisoniinae paraphyletic. Although the suborder defined herein is similar to the Pleu? rotomariina as defined by Yochelson (1984; see also Gordon and Yochelson, 1983), retaining the Pleurotomariina would be at odds with recent neontological work (e.g., Haszprunar, 1988; Ponder and Lindberg, 1996, 1997). The definition also differs from Yochelson's Pleurotomariina by excluding bellerophonts. NUMBER 88 11 Should future work corroborate suggestions that Sinuites and/ or other bellerophont gastropods belong to this clade, then they too should be included in the Murchisoniina. Finally, future work might equate this clade with the Orthogastropoda of Pon? der and Lindberg (1996); again, this would be a preferable la? bel than "murchisoniinae" simply because "Orthogastropoda" is not associated with a morphologic grade. Superfamily fMuRCHisONiolDEA Koken, 1896 REMARKS.?As the earliest members of the Murchisoniina have been classified as Hormotoma, the Murchisonioidea is used to label the paraclade within the Murchisoniina. In fact, the paraphyletic portions of the Murchisonioidea all are as? signed to Hormotoma, so it is monophyletic excepting for that genus. The clade is diagnosed by several features yielding an asymmetrical aperture, with the post-torsional left side greatly expanded at the expense of the right side. A weak bilineate pe? ripheral band also typifies members of the superfamily, al? though the feature is lost or highly modified on some species. Family fHORMOTOMiDAE Wenz, 1938 REMARKS.?The family Hormotomidae is resurrected to la? bel the paraclade within the Murchisonioidea. The Murchisoni- idae, Eotomarioidea, Subulitoidea, and Loxonematoidea all are derived from members of this family. The paraclade includes at least one monophyletic group at all times, however, and the post-Ordovician definition (see below) is strictly monophyletic. Genus ^Hormotoma Salter, 1859 FIGURES 8, 20,22-24 Cyrtostropha Donald, 1902. TYPE SPECIES.?Hormotoma gracilis (Hall, 1847). FKA.?Hormotoma !simulatrix (Billings, 1865): Watt's Bight Formation (Late Tremadoc (Demingian)). LKA.?Cyrtostropha coralli (Sowerby in Murchison, 1839): lower Ludlow Formation (Early Ludlow (Gorstian)). ADDITIONAL ASSIGNED SPECIES.?Hormotoma centervillen- sis (Foerste, 1923); H. confusa Cullison, 1944; Turritoma !anna (Billings, 1865). REMARKS.?Hormotoma as used herein is a large paraclade including high-spired species with round whorls and strong bi? lineate peripheral bands. The genus terminates with a true ex? tinction with the demise of the H. centervillensis clade in the Late Silurian. Pre-Silurian species classified as Coelocaulus belong to this clade. Genus ^Coelocaulus Oehlert, 1888 FIGURE 23 TYPE SPECIES.?Coelocaulus davidsoni (Oehlert, 1877). FKA.?Hormotoma insignis Koken, 1925: Wesenberger Shale (Late Caradoc (Vormsi)). LKA.?Blodgett et al. (1990) reported Coelocaulus species from unspecified Emsian localities. ADDITIONAL ASSIGNED SPECIES.?Coelocaulus concinnus Horny, 1952; Hormotoma cingulata (Lindstrom, 1884); H. monoliniformis (Lindstrom, 1884). REMARKS.?Silurian Coelocaulus represent a purely para? phyletic group relative to both Catazone and Mesocoelia. A Devonian monophylum that includes the type of the genus (C. davidsoni) certainly is derived from this paraclade, so the name is used for the larger paraclade. The genus differs from Hormo? toma in the presence of a slit. It differs from slit-bearing murchisoniids (see below) in retaining very round whorls, a bi? lineate peripheral band with weak lira, and a highly asymmetri? cal sinus. Genus Catazone Perner, 1903 FIGURE 23 TYPE SPECIES.?Catazone cunea Barrande in Perner, 1907. FKA.?!Catazone argolis Barrande in Perner, 1907, and other species: Pridoli Formation (Pridoli). LKA.?"Murchisonia" nevadana Blodgett and Johnson, 1992: Robert Mountains kockelianus-zone Limestone (Eife? lian). ADDITIONAL ASSIGNED SPECIES.?Catazone allevata Bar? rande in Perner, 1907. REMARKS.?The definition of Catazone used herein is much broader than that based on the traditional diagnosis of the ge? nus. The weak peripheral band low on the whorl now diagnoses only the most-derived species. The major feature now diagnos? ing the clade is an inflated upper ramp that gives the aperture a nearly square shape. Genus Mesocoelia Perner, 1907 TYPE SPECIES.?Mesocoelia j anus Barrande in Perner, 1907. FKA.?Kopanina Formation (Late Ludlow (Ludfordian)). LKA.?Eifelian (Linsley, 1978). REMARKS.?Unfortunately, the available specimens for this genus all are incomplete; however, it does appear to represent a sister taxon to the Catazone argolis clade. If it does represent a separate monophyletic group, then the basal species in Cata? zone should be transferred here, as Mesocoelia retains a more primitive morphology. Genus Plethospira Ulrich in Ulrich and Scofield, 1897 FIGURE 20 Seelya Ulrich in Ulrich and Scofield, 1897. TYPE SPECIES.?Plethospira cassina (Whitfield, 1886). FKA.?Plethospira cannonensis (Stauffer, 1937): Gorman Falls Formation (Late Tremadoc (Jeffersonian)). LKA.?Plethospira cassina and Seelya ventricosa Ulrich in Ulrich and Scofield, 1897: Fort Cassin Formation (Early Arenig (Cassinian)). 82 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY REMARKS.?It actually is inappropriate to put Plethospira in the Hormotomidae, as the other clades branching at this time are accorded superfamilial status. Plethospira was a more rap? idly terminated dead end (producing as few as three species), so according it higher rank than a genus seems absurd. Also, it maintains the paraclade status of the Hormotomidae at a higher level, ensuring that there is always at least one monophyletic clade within the family at all times. Seelya is synonymized herein; it previously was separated only by the presence of or? namentation. As none of the other genera previously classified in the Plethospiridae are closely related to Plethospira, usage of that family should be discontinued. Family MURCHISONHDAE Koken, 1896 REMARKS.?The Murchisoniidae herein are restricted to a monophyletic group of "hormotomoids" with concave upper whorls and fairly acute sutures. The clade also appears to be di? agnosed by a large, planispirally coiled protoconch. Genus ^Murchisonia d'Archaic, 1841 FIGURES 23,24 Goniostropha Oehlert, 1888. TYPE SPECIES.?Murchisonia bilineata Dechen in De la Beche, 1831. FKA.?Hormotoma salteri (Ulrich and Scofield, 1897): Black River Formation (Early Caradoc (Black Riverian)). LKA.?Late Guadalupian. ADDITIONAL ASSIGNED SPECIES.?D. morinenis Horny, 1953; Goniostropha cava (Lindstrom, 1884); G. sculpta (Bar? rande in Perner, 1907); Hormotoma subplicata (Lindstrom, 1884); Murchisonia laphami Whiteaves, 1895. REMARKS.?Murchisonia as defined herein represents a much narrower range of species than it has in traditional defini? tions. It is still a paraclade, however, as Morania is considered a derivative herein. The chief difference between Murchisonia and Hormotoma is the acute, somewhat channeled suture of Murchisonia, which contrasts with the rounded upper whorl of Hormotoma. A slit diagnoses more-derived members of Murchisonia. Genus Morania Horny, 1953 FIGURE 24 ILeptorima Pemer, 1907. TYPE SPECIES.?Morania v-sinuata Horny, 1953. FKA.?Loxonema! attenuata (Lindstrom, 1884): Bed G, Slite Formation (Early Wenlock (Sheinwoodian)). LKA.?Donaldiella declivis (Barrande in Perner, 1907): Ko? panina Formation (Late Ludlow (Ludfordian)). ADDITIONAL ASSIGNED SPECIES.?Murchisonia paradox Lindstrom, 1884. REMARKS.?The problematic Leptorima likely is synony? mous with Morania. Because Leptorima is known from very poor material and has not been used aside from the original de? scription, it cannot be used reliably and should be disregarded. Genus Michelia Roemer, 1854 FIGURE 24 Sinuspira Pemer, 1907. TYPE SPECIES.?Michelia cylindrica Roemer, 1854. FKA.?Hormotoma attenuata (Lindstrom, 1884): Bed B, Slite Formation (Early Wenlock (Sheinwoodian)). LKA.?The type and other species are known from the Em? sian. ADDITIONAL ASSIGNED SPECIES.?Sinuspira tenera Bar? rande in Perner, 1907. REMARKS.?Michelia and Sinuspira are united herein based on their shared loss of the peripheral band late in ontogeny. The deep, highly asymmetrical sinus clearly links them to murchisoniids rather than to loxonematids. Superfamily LOXONEMATOIDEA Koken, 1889 ? = Basal Apogastropoda + Euthyneura REMARKS.?This represents one of the few clades separated at a higher rank because of morphologic disparity. The super- family probably is synonymous with traditional paleontologi? cal definitions of the Caenogastropoda, which is a suborder. Among more modern taxonomies, it likely is synonymous with the Apogastropoda + Euthyneura of Haszprunar, 1988. Whether this disparity contributes to any real difference rela? tive to the Murchisonioidea (i.e., changes in rates of speciation, extinction, or morphologic change) awaits later testing. Family LOXONEMATIDAE Koken, 1889 REMARKS.?The addition of Devonian species will render this a paraclade, as it is unlikely that the earliest heterostrophs (e.g., Pseudozygopleura) will be retained with other loxonema? toids. Again, whether heterostrophic coiling sufficiently alters evolutionary patterns to merit such recognition will require quantitative demonstration. Genus ^Loxonema Phillips, 1841 FIGURE 25 TYPE SPECIES.?Loxonema sinuosa Phillips, 1841. FKA.?Loxonema murrayana Salter, 1859: Paquette Rapids Black River Formation (Early Caradoc (Black Riverian)). LKA.?Wenz (1938) recorded a Norian occurrence of the genus. ADDITIONAL ASSIGNED SPECIES.?Loxonema brogeri Ko? ken, 1925; L. costulata Barrande in Perner, 1907; L. cross- NUMBER 88 83 manni Phillips, 1841; L.fasciata Lindstrom, 1884; L. strangu- lata Lindstrom, 1884. REMARKS.?As used herein, Loxonema represents a para? clade of "omospirines" with deep U-shaped sinuses but no pe? ripheral bands. The paraclade status is maintained in the Sil? urian through several species that are not illustrated. Future work probably will break the present definition (which extends through the Triassic) into several genera. Genus ^Omospira Ulrich and Scofield, 1897 FIGURES 23, 25 TYPE SPECIES.?Omospira laticincta Ulrich and Scofield, 1897. FKA.?Omospira laticincta and O. alexandra Ulrich and Scofield, 1897: Platteville Formation (Early Caradoc (Black Riverian)). LKA.?Turritoma pinguis (Donald, 1902) (? = Turritoma polita (Barrande in Perner, 1907)): Starfish Bed, Thraive Glen Formation (Ashgill (Rawtheyan)). ADDITIONAL ASSIGNED SPECIES.?Hormotoma trentonensis (Ulrich and Scofield, 1897). REMARKS.?This paraclade is diagnosed by the retention of a weak bilineate peripheral band, a shallow nearly symmetrical sinus, globose whorls, and reduced translation. Genus Diplozone Perner, 1907 FIGURE 25 TYPE SPECIES.?Diplozone innocens Barrande in Perner, 1907. FKA.?Diplozone crispa (Lindstrom, 1884): Upper Hemse Marl (Early Ludlow (Gorstian)). LKA.?Diplozone innocens: Koneprussy Formation (Pra- gian). ADDITIONAL ASSIGNED SPECIES.?Loxonema beraultensis Barrande in Perner, 1907. REMARKS.?This Plethospira-like genus apparently repre? sents a short-lived siphonate experiment within "loxonemati? des." It is diagnosed by an extended, tightly coiled columella lip that yields a weak siphon and by a short, narrow but very sharp (almost slit-like) sinus. Genus Rhabdostropha Donald, 1905 FIGURES 25, 26 IHolopellaU 'Coy, 1851. Kjerulfonema Peel and Yochelson, 1976. TYPE SPECIES.?Rhabdostropha grindrodii (Donald, 1905). FKA.?Rhabdostropha primitiva Longstaff, 1924: Shalloch Mill Formation (Late Caradoc (Onnian)). LKA.?Holopella regularis Lindstrom, 1884: Hemse Marls (Early Ludlow (Gorstian)). ADDITIONAL ASSIGNED SPECIES.?Kjerulfonema cancellata (Sowerby in Murchison, 1839); K. quinquecincta Peel and Yochelson, 1976; Murchisonia sp. (Point Clarence Limestone; Rohr, 1988); Rhabdostropha latisinuata Longstaff, 1924. REMARKS.?Rhabdostropha is used for a clade of "omospir? ines" with shallow sinuses and ornament. Holopella has taxo? nomic priority over Rhabdostropha and certainly has been used more frequently. Unfortunately, the diagnosis of Holopella was based on a steinkern, thus making it impossible to determine its affinities and probably contributing to the very inconsistent ap? plications of the genus name. Usage of Holopella should, therefore, be discontinued. Genus Spiroecus Longstaff, 1924 FIGURE 26 Girvania Longstaff, 1924. TYPE SPECIES.?Spiroecus girvanensis Longstaff, 1924. FKA.?Girvania excavata (Longstaff, 1924): Whitehouse Group (Late Caradoc (Onnian)). LKA.?Spiroecus girvanensis: Girvan Limestone (Early Ashgill (Pusgillian)). REMARKS.?This labels an early but highly derived clade of ornate loxonematoids. A key diagnostic feature, a strong lirum on the left side of the sinus apex, might be a remnant peripheral band. Genus Macrochilus Lindstrom, 1884 FIGURE 26 Auriptygma Pemer, 1903. TYPE SPECIES.?Macrochilus fenestratus Lindstrom, 1884. FKA.?Macrochilus fenestratus: Slite Bed, Unit G (Early Wenlock (Sheinwoodian)). LKA.?"Auriptygma" species: Zlichov Limestone (Emsian). ADDITIONAL ASSIGNED SPECIES.?Auriptygma fortior Bar? rande in Perner, 1903; Macrochilina recticosta (Barrande in Perner, 1907); Macrochilus buliminus Lindstrom, 1884; M. cancellatus Lindstrom, 1884. REMARKS.?Knight et al. (1960) considered Macrochilus to be a junior synonym of Soleniscus, which appears to be a true subulitoid. Some post-Emsian "Soleniscus" species, however, might belong to Macrochilus. Although these species con? verge strongly on the subulitioid form, they retain a broad but shallow sinus, and they display a much weaker siphon than do subulitioids. The siphon of Macrochilus also differs from that of Soleniscus. In Macrochilus the siphon is formed by elon? gating and slightly bending the inner margin. In Soleniscus the siphon is produced by widening the base of the inner margin as it is extended beneath and around the base of the penulti? mate whorl (which yields the distinctive plicate columella of subulitioids). 84 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY Genus Stylonema Perner, 1907 FIGURE 26 TYPE SPECIES.?Stylonemapotens Barrande in Perner, 1907. FKA.?Stylonema potens: Kopanina Formation (Late Lud? low (Ludfordian)). LKA.?!Stylonema opposita Barrande in Perner, 1907: Zlichov Limestone (Emsian). ADDITIONAL ASSIGNED SPECIES.?Stylonema domestica Barrande in Perner, 1907; S. mater Barrande in Perner, 1907. REMARKS.?The sinistral S. opposita might not belong to this genus. Another Emsian species, S. dilatata (Spitz, 1907) is too poorly known to accurately place. Thus, the last appearance of this genus might be in the Pragian (S. domestica). Superfamily EOTOMARIOIDEA Ulrich and Scofield, 1897 (? = Vetigastropoda) REMARKS.?This taxon corresponds to the "eotomarioid" clade. Of the superfamilies defined herein, the Eotomarioidea corresponds most closely to traditional definitions of the Pleu? rotomarioidea. It is very possible that the Eotomarioidea are synonymous (or nearly so) with the Vetigastropoda. The clade is diagnosed primitively by a peripheral band with strong pe? ripheral lira, a slight increase in translation over ontogeny, and an asymmetrical sinus with the right side broader and shal? lower than the left side. Family EOTOMARIIDAE Wenz, 1938 REMARKS.?The Eotomarioidea encompasses the basal para? clade of the "eotomarioids," plus the monophyletic "liospir? ines." Again, it includes at least one monophyletic group of clades throughout its history. The family is diagnosed by the features primitively diagnosing the superfamily. Genus ^Clathrospira Ulrich and Scofield, 1897 FIGURES 20, 27-29 Turritoma Ulrich and Scofield, 1897. TYPE SPECIES.?Clathrospira subconica (Hall, 1847). FKA.?Turritoma acrea (Billings, 1865): Boat Harbor For? mation (Early Arenig (Jeffersonian)). LKA.?Clathrospira conica Ulrich and Scofield, 1897: Meaford Formation (Ashgill (Richmondian)). ADDITIONAL ASSIGNED SPECIES.?Clathrospira !trochifar- mis Butts, 1926; Clathrospira species, Smithville Formation; Turritoma ornate species, Cotter Formation. REMARKS.?The most radical change in the present defini? tion of Clathrospira is the inclusion of Turritoma species. That name, however, does not define any type of phylogenetically cohesive group, and the type species is plesiomorphic relative to typical Clathrospira. It is best considered a primitive high- spired version of Clathrospira. Clathrospira is diagnosed by a very strong bilineate peripheral band that is located very low on the whorl owing to a very short left ramp relative to the right ramp. ?Genus Spirotomaria Koken, 1925 FIGURE 28 TYPE SPECIES.?Spirotomaria rudissima Koken, 1925. FKA.?Spirotomaria rudissima: Vaginatum Limestone (Late Arenig (Kundan)). LKA.?!Clathrospira euconica Butts, 1926: ?Tulip Creek Formation (Late Llandeilo (Chazyan)). ADDITIONAL ASSIGNED SPECIES.?Clathrospira !glindmey- en'Rohr, 1996; Clathrospira !ulrichi (Endo, 1932). REMARKS.?These assignments are made based on the simi? larities between S. !glindmeyeri, S. !ulrichi, and figures of the poorly known type species, S. rudissima. As I have not been able to examine any S. rudissima specimens, so the assign? ments must be considered tentative. The high asymmetry in the left and right ramp lengths, coupled with a high angle between the base of the columella and the alveozone, leave the periph? eral band very low on Spirotomaria species. Genus ^Eotomaria Ulrich and Scofield, 1897 FIGURE 29 IPseudocryptaenia Koken, 1925. TYPE SPECIES.?Eotomaria canalifera Ulrich and Scofield, 1897. FKA.?Eotomaria canalifera: Murfreesboro Limestone (Early Caradoc (Ashbian)). LKA.?Eotomaria notablis (Eichenwald, 1859): Lyckholm Formation (Ashgill (Late Pirgu)). ADDITIONAL ASSIGNED SPECIES.?Eotomaria dryope Ulrich and Scofield, 1897; !E. tumida Koken, 1925. REMARKS.?The definition of Eotomaria presented herein is somewhat reduced relative to most definitions, as it focuses on Baltic species. Most Laurentian species previously placed in Eotomaria now are placed in Paraliospira or Liospira. Eoto? maria is a paraclade relative to both genera, but it suffers true extinction with the termination of the Baltic clade. Pre-Carado- cian specimens assigned to Eotomaria usually belong to Cer? atopea. There possibly are multiple Baltic species within this clade, as E. notablis has not been well defined as a species; however, I was unable to examine enough specimens to ascer? tain this. Also, the poorly known Pseudocryptaenia might be? long to this clade; if so, then it should be considered a junior synonym unless it forms a distinct monophyletic group with Eotomaria. Genus Paraliospira Rohr, 1980 FIGURE 29 TYPE SPECIES.?Paraliospira angulata (Ulrich and Scofield, 1897). NUMBER 88 85 FKA.?Liospira larvata (Salter, 1859): Murfreesboro Lime? stone (Early Caradoc (Ashbian)). LKA.?Paraliospira aff. P. angulata: Port Clarence Lime? stone (Ashgill (Richmondian)). ADDITIONAL ASSIGNED SPECIES.?Eotomaria rupestris Ko? ken, 1925; E. supracingulata (Billings, 1865); Paraliospira mundula (Ulrich and Scofield, 1897); P. rugata (Ulrich and Scofield, 1897). REMARKS.?Rohr's (1980) initial definition of Paraliospira remains largely unchanged, save that it now includes some spe? cies previously assigned to Eotomaria. Relative to Eotomaria, Paraliospira is diagnosed by a stronger peripheral band that is higher on the aperture owing to rotation of the aperture (e.g., the projection of the inner margin) and much stronger anterior projection of the outer margin of the aperture. The poorly known Eocryptaenia might belong here. If so, it technically would be a senior synonym, but as the type material was poorly illustrated and subsequently lost (Knight, 1941), the name probably should be abandoned. Specimens from the Nanook Limestone of Alaska (Ashgill (Richmondian)) identified as "Siskyouspira" by Blodgett, Rohr, Harris, and Rong (1988) either are Paraliospira rupestris or are close relatives of that species. Another species placed in Siskyouspira by those authors, S. majewski (intially classified as Pseudocryptaenia majewski by Rohr and Blodgett (1985)) also appears to belong to the Paraliospira clade. The type spe? cies of Siskyouspira Rohr is known only from deformed speci? mens. These specimens, however, appear to show synapomor? phies with "helicotomoids" rather than with "eotomarioids." Genus Liospira Ulrich and Scofield, 1897 FIGURE 29 TYPE SPECIES.?Liospira micula (Hall and Whitney, 1862). FKA.?Eotomaria labrosa Ulrich and Scofield, 1897: Mur? freesboro Limestone (Early Caradoc (Ashbian)). LKA.?Liospira micula: Port Clarence Limestone (Ashgill (Richmondian)); !L. marklandensis McLearn, 1924: Stone- house Formation (Pridoli). ADDITIONAL ASSIGNED SPECIES.?!Liospira affmis Pitcher, 1939; L. angustata Ulrich and Scofield, 1897; L. decipens Ul? rich and Scofield, 1897; L. progne Ulrich and Scofield, 1897; L. subconcava Ulrich and Scofield, 1897. REMARKS.?The question marks above denote Silurian spe? cies that might not belong to Liospira; however, better material is needed to evaluate this possibility. The genus is best diag? nosed by a very strong columella partly produced by a thick? ened inner margin that is folded backwards, but mainly by a fu? nicle produced by an extension of the parietal inductura. Putative Liospira species lacking these features likely belong to Raphistomina or Pararaphistoma. Notably, the type species of the genus is strongly convergent on Pararaphistoma species, although it can be distinguished by (1) anterior rather than pos? terior projection of the aperture, (2) a slit that is consistent throughout phylogeny and that does not appear to be an exten? sion of the sinus, and (3) a thick columella plus a strong funicle filling the umbilicus. The genus also is diagnosed by a strongly reduced peripheral band. On the most-derived species (e.g., L. micula), the band is almost entirely absent, although a true se? lenizone (i.e., distortion in growth lines produced by a slit) ex? ists on the left side of the remnant peripheral band. Family |GOSSELETINIDAE Wenz, 1938 REMARKS.?The definition of this family is nearly identical to the one provided by Knight et al. (1960), which highlights the tendency for traditional taxonomy to be far more congruent with the phylogeny of post-Ordovician taxa than with the phy? logeny of Ordovician genera. As it includes a genus that is paraphyletic relative to the Phanerotrematidae (see below), it is a paraclade. Notably, this clade is comprised predominantly of Baltic species during the Ordovician and provides much of the post-Ordovician diversity. Subfamily fEURYZONiNAE, new subfamily REMARKS.?The definition of this subfamily is nearly identi? cal to that of the Coelozoninae provided by Knight et al. (1960). Coelozone appears to be a junior synonym of Eu? ryzone, so I am simply updating the subfamily name. Genus ^Deaechospira, new genus FIGURE 28 TYPE SPECIES.?Clathrospira elliptica (Hisinger, 1829). FKA.?Deaechospira elliptica: Huk Formation (Early Llan? virn (Kundan)). LKA.?Clathrospira maritima Koken, 1925: Wesenberger Shale (Late Caradoc (Vormsian)). ADDITIONAL ASSIGNED SPECIES.?Clathrospira inflata Ko? ken, 1925. ETYMOLOGY.?Named for the favorite baseball rule of Dou? glas H. Erwin, the Designated Hitter (DH) rule. Among the fea? tures separating Deaechospira from the ancestral Clathrospira is a rounder profile, a feature that typically separates desig? nated hitters from other baseball players. The naming of a typi? cally European clade after a rule in a typically American game is odd and inappropriate, but so is the rule. REMARKS.?Baltic species previously classified in Clath? rospira form their own clade. The clade differs from Clath? rospira by the narrower peripheral band that features weaker but sharper lira. Deaechospira might include more species than I have recognized, as D. elliptica and D. inflata have been used imprecisely. Compared to Clathrospira, Deaechospira has a peripheral band with only moderately strong threads of little prominence. Also, this band bisects the left and right ramps evenly on Deaechospira, whereas the peripheral band of Cla- thospira is located on the base of the right ramp. Deaechospira also displays greater symmetry between the left and right 86 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY ramps than does Clathrospira, placing the peripheral band nearer the center of the whorl. Finally, Deaechospira displays rounder right and left ramps than does Clathrospira. Genus ^Cataschisma Branson, 1909 FIGURES 28,30,31,34, 35 Globispira Koken, 1925.?Knight et al., 1960. TYPE SPECIES.?Cataschisma exquisita (Lindstrom, 1884). FKA.?Mourlonia mjoela Yochelson, 1963: Cephalopod Shale, Huk Formation (Early Llanvirn (Kundan)). LKA.?Cataschisma exquisita: Mulde Beds (Late Wenlock (Whitwellian)). ADDITIONAL ASSIGNED SPECIES.?Clathrospira convexa Ul? rich and Scofield, 1897; Globispirapillula (Koken, 1925). REMARKS.?This is another paraclade that includes the basal members of the Gosseletinidae plus a small clade diagnosed by strong transverse ornament. The genus is distinguished from both Clathrospira and Deaechospira by much rounder ramps and greatly reduced asymmetry between the left and right ramps, leaving the weaker peripheral band in the middle of the aperture. The peripheral band of Cataschisma also is wider than on Deaechospira. On derived members, the peripheral band is very low owing to the right ramp being much broader than the left ramp. ?Genus Palaeoschisma Donald, 1902 FIGURES 30,31 TYPE SPECIES.?Palaeoschisma girvanensis Donald, 1902. FKA.?!Eotomaria elevata Ulrich in Ulrich and Scofield, 1897: Lexington Limestone (Middle Caradoc (Rocklandian)). LKA.?!Clathrospira thraivensis Longstaff, 1924: Starfish Bed, Thraive Glen Formation (Ashgill (Rawtheyan)). ADDITIONAL ASSIGNED SPECIES.?"Bembexia" globosa Longstaff, 1924. REMARKS.?Much about this genus is tentative. Three spe? cies, IP. globosa, !P. thraivensis, and IP. elevata, clearly form a monophyletic group; however, it is not clear that these spe? cies actually are related to P. girvanensis (the type species). Genus ^Pleurorima Perner, 1907 FIGURE 34 TYPE SPECIES.?Pleurorima migrans Barrande in Perner, 1907. FKA.?Pleurorima wisbeyensis (Lindstrom, 1884): Unit G, Slite Formation (Early Wenlock (Sheinwoodian)). LKA.?See remarks. ADDITIONAL ASSIGNED SPECIES.?Globispira prima Bar? rande in Perner, 1907; Seelya moydartensis Peel, 1977. REMARKS.?The last record of Pleurorima is from the Late Ludlow (Ludfordian) of Central Europe and Canada. Prelimi? nary analyses of Devonian species, however, indicate that spe? cies assigned to Mourlonia and Ptychomphalina clearly belong to the Pleurorima clade. Thus, it is not clear when a monophyl? etic assemblage best assigned to Pleurorima became extinct. The genus is diagnosed by a broad and swollen peripheral band with wide but weak lira bordering a seemingly short slit. Genus Euryzone Koken, 1896 FIGURE 34 ICoelozone Perner, 1907. TYPE SPECIES.?Euryzone delphinuloides (Schlotheim, 1820). FKA.?Euryzone calva Barrande in Perner, 1907, and Co- elozone verna Barrande in Perner, 1907: Kopanina Formation (Late Ludlow (Ludfordian)). LKA.?Euryzone has been reported as late as the Frasnian. ADDITIONAL ASSIGNED SPECIES.?Euryzone connulastus Barrande in Perner, 1907; Pleurorima aptychia Barrande in Pemer, 1907. REMARKS.?Euryzone and Coelozone are provisionally united herein, pending work on Devonian species. Coelozone has been recorded in Devonian rocks, so it is possible that it in? cludes a separate clade. The species identified by Koken (1925) as "Euryzone" belong to Brachytomaria. Genus Latitaenia Koken, 1925 TYPE SPECIES.?Latitaenia rotelloidea (Koken, 1896). FKA.?Latitaenia rotelloidea: Lower Chasmops Shale (Early Caradoc (4a-cc = Idavere)). LKA.?Latitaenia aequicrescens Koken, 1925, and Latitae? nia kirnaensis Koken, 1925: Lyckholm Formation (Ashgill (Pirgu)). REMARKS.?Members of this genus are not featured in this analysis because I have not been able to examine any speci? mens personally. Analyses of figured specimens, however, in? dicate that Latitaenia represents a small clade derived from the Cataschisma paraclade. Subfamily GOSSELETININAE Wenz, 1938 REMARKS.?A monophyletic definition of this subfamily is very similar to the one provided by Knight et al. (1960). Analy? ses of Devonian species, however, indicate that the (nearly) bi? laterally symmetrical Porcellidae belong to this clade. Genus ^Stenoloron Oehlert, 1888 FIGURES 34,35 Spiroraphe Perner, 1907. TYPE SPECIES.?Stenoloron aequilatera Oehlert, 1888. FKA.?Stenoloron shelvensis Pitcher, 1939: Pentamerous Beds (Late Llandovery (Telychian)). NUMBER 88 87 LKA.?Stenoloron minor Blodgett and Johnson, 1992: Lone Mountain Limestone, ensensis-zone (Late Eifelian). ADDITIONAL ASSIGNED SPECIES.?Clathrospira biformis (Lindstrom, 1884); Oriostoma angulifer Lindstrom, 1884; Spiroraphe bohemica Barrande in Perner, 1907; Stenoloron pollens Barrande in Pemer, 1907; S. viennayi Oehlert, 1888. REMARKS.?This paraclade includes the ancestors of Platy- loron (see below) and the Porcellidae. If future work corrobo? rates the estimate linking !S. angulifer and !S. biformis, then those species should be removed into a separate genus. Stenoloron is diagnosed by a curved base with a slight posterior projection and by a very narrow peripheral band that appears to encompass a moderately long slit. Some derived members of Paraliospira (e.g., P. rupestris) have very similar gross mor? phologies to Stenoloron; however, Stenoloron can be distin? guished from those species by the narrow sinus and the curved base of the inner margin. Genus Platyloron Oehlert, 1888 FIGURE 35 TYPE SPECIES.?Platyloron bischoffi (Goldfuss, 1844). FKA.?Oriostoma dispar Lindstrom, 1884: Unit G, Slite Beds (Early Wenlock (Sheinwoodian)). LKA.?Platyloron is known through the Famennian. ADDITIONAL ASSIGNED SPECIES.?Stenoloron voluta (Lind? strom, 1884). REMARKS.?This distinctive genus is diagnosed by a periph? eral band near the top of the whorl, owing to both rotation of the aperture and expansion of the left ramp relative to the right ramp. Silurian species, however, retain Stenoloron-like pro? files. Like Stenoloron, the peripheral band encompasses a slit. Genus Umbotropsis Perner, 1907 FIGURE 35 Eocryptaulina Foerste, 1936. TYPE SPECIES.?Umbotropsis albicans Barrande in Perner, 1907. FKA.?Eocryptaulina helcinia (Lindstrom, 1884): Unit A, Lower Visby Formation (Late Llandovery (Telychian)). LKA.?Umbotropsis albicans: Kopanina Formation (Late Ludlow (Ludfordian)). ADDITIONAL ASSIGNED SPECIES.?Eocryptaulina filitexta (Foerste, 1893). REMARKS.?This genus currently is poorly known but seems to represent a monophyletic group. Umbotropsis is diagnosed by a thin, weak peripheral band encompassing a slit and thick? ened bases of the inner margin, sometimes forming a sharp lira. Family fPHANEROTREMATiDAE Knight, 1956 REMARKS.?This family also is very similar to the definition presented by Knight et al. (1960). There is one major differ? ence, however, as preliminary analyses of Devonian species place post-Ordovician "raphistomatids" Buechelia, Scalitina, Denayella, and Wisconsinella in the Phanerotrematidae. These analyses also place Pleuromphalus in the family, which con? verges strongly on a euomphalinae, such as Lesueurilla. Or? dovician and Silurian species are diagnosed by narrow and shallow sinuses with unusually rugose growth lines, including very strong lunulae. Genus ^Brachytomaria Koken, 1925 FIGURES 30-32 1 Promourlonia Longstaff, 1924. TYPE SPECIES.?Brachytomaria baltica Koken, 1925. FKA.?Euryzone kiari Koken, 1925: Keila Formation (Mid? dle Caradoc (Keila)). LKA.?Bembexia cognata Barrande in Perner, 1903: Kopan? ina Formation (Late Ludlow (Ludfordian)). ADDITIONAL ASSIGNED SPECIES.?Brachytomaria papulosa (Billings, 1852); B. semele (Ulrich and Scofield, 1897); B. stri? ata (Ulrich and Scofield, 1897); B., new species, Gotland (some specimens assigned to Pleurotomaria laquetta by Lind? strom (1884); the type of that species, however, is a lophos- piroid); Lophospira kindlei Foerste, 1924; Promourlonia fur? cata Longstaff, 1924. REMARKS.?This paraclade includes the basal members of all Silurian genera of phanterotrematids. In the Silurian it ap? pears to represent a monophyletic group that includes species with lower translations that previously were classified as Pro? mourlonia. If the phylogenetic assessment presented herein is correct, then Promourlonia has priority over Brachytomaria. The lone possible Promourlonia species included herein differs from carinate Brachtyomaria species (e.g., B. semele) largely in having slightly lower curvature, resulting in an umbilicus. Even if morphologic separation is sometimes grounds for sepa? rating supraspecific taxa, small differences in coiling parame? ters certainly are not sufficient for diagnosing genera. Pro? mourlonia has not been used since the original diagnosis, and the name incorrectly implies a close relationship with Mourlo? nia, whereas Brachytomaria has been used by many workers and is a phylogenetically neutral name. Thus, retaining Brachy? tomaria clearly is ultimately preferable. Members of the genus typically are diagnosed by a very shal? low sinus, a strong peripheral band with prominent peripheral lira, and strong carina on the upper and lower ramps. The very strong growth lines and lunulae of most Brachytomaria species are contiguous through the peripheral band, indicating the ab? sence of a slit (contra Knight, 1945). Some of the more-derived species, however, display evidence of short slits. Genus Phanerotrema Fischer, 1885 FIGURE 32 Pseudoscalites Boucot et al., 1967. [Not Diener, 1926.] TYPE SPECIES.?Phanerotrema labrosa (Hall, 1859). 88 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY FKA.?Phanerotrema jugosa Pitcher, 1939: Beechhill Cove Formation (Early Llandovery (Rhuddanian)). LKA.?Phanerotrema is recorded through the Bashkirian. ADDITIONAL ASSIGNED SPECIES.?Phanerotrema australis (Etheridge, 1891); P. dolia (Lindstrom, 1884); P. lindstroemi (Boucot et al., 1967); P. !occidens (Hall, 1860); "Seelya" !vi- tellia (Billings, 1865). REMARKS.?Silurian species recently assigned to the genus/ subgenus Pseudoscalites are now returned to Phanerotrema, as the former name is preoccupied by a Triassic caenogastropod genus. Pseudoscalites illustrates an important misconception about Phanerotrema, i.e., that it is related to Scalites and Ra? phistoma. These analyses, plus preliminary ones that include Devonian species, indicate that the similarities (which are strik? ing in some post-Silurian taxa) are convergences. Phaner? otrema appears to represent the paraclade from which post-Sil? urian taxa previously assigned to the Raphistomatidae arose. The genus is diagnosed by rotation and alteration of the aper? ture, leaving the peripheral band higher on the whorl (and pro? ducing a more step-like profile) than observed on Brachy? tomaria. Genus Ulrichospira Donald, 1905 FIGURE 32 TYPE SPECIES.?Ulrichospira similis Donald, 1905. FKA.?Ulrichospira similis: Pentamerous Beds (Middle Llandovery (Telychian)). Note that the phylogenetic analyses indicate that it must have originated by the Early Llandovery. LKA.?"Seelya" lloydi Sowerby in Murchison, 1839: Ko? panina Formation (Late Ludlow (Ludfordian)). ADDITIONAL ASSIGNED SPECIES.?Murchisonia othemenis (Lindstrom, 1884). REMARKS.?This restricted clade of phanerotrematids is di? agnosed by high translations and a weak peripheral band that encompasses a slit. The peripheral band appears to be on the upper periphery owing to the orientation of the left and right ramps plus some displacement of the peripheral band toward the suture. Family LUCIELLIDAE Knight, 1956 REMARKS.?Preliminary analyses suggest that Devonian genera assigned to the Portlockidae belong to the Luciellidae. The clade is diagnosed by a narrow peripheral band that is low on the whorl owing to high asymmetry between the right and left ramps. Genus Conotoma Perner, 1907 FIGURES 30, 33 TYPE SPECIES.?Conotoma eximia (Goldfuss, 1844). FKA.?Conotoma claustrata (Lindstrom, 1884): Lower Visby Formation (Early Wenlock (Sheinwoodian)). LKA.?Conotoma eximia: Kopanina Formation (Late Lud? low (Ludfordian)). ADDITIONAL ASSIGNED SPECIES.?Conotoma glandiformis (Lindstrom, 1884). REMARKS.?This genus represents a paraclade relative to the rest of the Luciellidae. Species within the genus previously have been classified in Clathrospira and Euconospira. The ge? nus is diagnosed by pronounced increase in translation over on? togeny and a peripheral band on which the right (upper) lirum is slightly stronger than the left (lower) lirum. The peripheral band is very low on the whorl owing to the right ramp being much longer than the left ramp. Genus Prosolarium Perner, 1907 FIGURE 33 Platyconus Perner, 1907. Crenilunula Knight, 1945. TYPE SPECIES.?Prosolarium procerum Barrande in Pemer, 1907. FKA.?Crenilunula hallei (Whiteaves, 1895): Lower Visby Formation (Early Wenlock (Sheinwoodian)). LKA.?Platyconus incumbens Barrande in Perner, 1907: Pridoli Beds (Pridoli). Some Devonian members of the Luciel? lidae likely belong to this genus. ADDITIONAL ASSIGNED SPECIES.?Crenilunula limata (Lindstrom, 1884). REMARKS.?Prosolarium labels a clade that evidently in? cludes the Devonian Luciella de Koninck, 1883. It is possible that work in progress will unite Prosolarium and Luciella. Pro? solarium includes species previously placed in Crenilunula in order to maintain a cohesive phylogenetic unit. Platyconus is represented by a scrappy specimen, which appears to be either P. hallei or a closely related species. The genus has not been used since Perner first proposed it and should be eliminated. Prosolarium is diagnosed by extreme asymmetry between the right and left ramps (leaving the peripheral band very low on the whorl), very strong peripheral lira that create a short frill (which Knight et al. (1960) confounded with the frill-like alve? ozone carina of Euomphalopterus; see Linsley et al., 1978), and an unusual "zipper-like" lunulae pattern within the periph? eral band. Genus Oehlertia Perner, 1907 FIGURES 30,33 IQuadricarina Blodgett and Johnson, 1992. TYPE SPECIES.?Oehlertia senilis (Barrande in Perner, 1903). FKA.?Oehlertia scutulata (Lindstrom, 1884): Lower Visby Formation (Early Wenlock (Sheinwoodian)). LKA.?Oehlertia lenticularis (Goldfuss, 1844): Stringo- cephalen Limestone (Givetian). NUMBER 88 89 ADDITIONAL ASSIGNED SPECIES.?Oehlertia cancellata (Lindstrom, 1884); O. gradata (Lindstrom, 1884); Quadricar- ina glabrobasis Blodgett and Johnson, 1992. REMARKS.?Preliminary analyses of Devonian species place Quadricarina within the Oehlertia clade. The genus is diag? nosed by a thin slit bordered by strong lira that lies well within the peripheral band. The assignment of Oehlertia to the Luciel? lidae is considered tentative, as the genus' phylogenetic posi? tion is uncertain. Superfamily LOPHOSPIROIDEA Wenz, 1938 REMARKS.?Wagner (1999) elevated the taxon to superfam? ily status. It includes taxa formerly assigned to the Superfamily Trochonematoidea, including the genus Trochonema; however, due to the highly polyphyletic nature to the Trochonematoidea and also to the very dissimilar taxon definitions, it is recom? mended that the Trochonematoidea be abandoned. Family LOPHOSPIRIDAE Wenz, 1938 REMARKS.?The phylogeny and systematics of this clade are described elsewhere (Wagner, 1990, 1995a, 1999; Wagner and Erwin, 1995); however, a summary of the clade's taxonomy is presented herein for comparison with other taxa. Genus Ectomaria Koken, 1896 FIGURE 27 Solenospira Ulrich and Scofield, 1897.?Knight et al., 1960. TYPE SPECIES.?Ectomaria nieszkowskii Koken, 1925. FKA.?Ectomaria adelina (Billings, 1865): Boat Harbor Formation (Early Arenigian (Jeffersonian)). LKA.?Ectomaria nieszkowskii: Borkholm Formation (Late Ashgill (Porkunian)). ADDITIONAL ASSIGNED SPECIES.?Ectomaria adventa (Butts, 1926); E. laticarinata Koken, 1925 (= E. cf. E. prisca sp. 1 of Rohr (1988)); E. pagoda (Ulrich and Scofield, 1897); E. cf. E. pagoda of Rohr (1988); E. prisca (Ulrich and Scofield, 1897); E. cf. E. prisca (= E. cf. E. prisca sp. 2 of Rohr (1988)); Ectomaria Shakopee Dolomite species ("Hormotoma cassina" of Stauffer, 1937); Murchisonia callahanensis Rohr, 1980. REMARKS.?Ectomaria is a basal paraclade by merit of one species and is monophyletic thereafter. Ultimately, one sub? clade of relatively low-spired species (E. callahanensis, E. lati? carinata, and the unfigured E. adventa) might be separated into a new genus. If so, then this new genus would range from the Late Llanvirn (Lenoir Limestone) through the Ashgill (Port Clarence Limestone). Genus Donaldiella Cossmann, 1903 FIGURE 27 Pagodospira Grabau, 1922.?Knight etal., 1960. TYPE SPECIES.?Donaldiella ftlosa (Donald, 1902). FKA.?Pagodospira cicelia (Billings, 1865): Durness Lime? stone (Early Arenig (Cassinian)). LKA.?"Hormotoma" trilineata Foerste, 1923: Belfast For? mation (Middle Llandovery (Rhuddanian)). Genus Lophospira Whitfield, 1886 FIGURE 27 TYPE SPECIES.?Lophospira milleri (Hall in Miller, 1877). FKA.?Lophospira perangulata (Hall, 1847): Smithville Formation (Early Arenig (Cassinian)). LKA.?Lophospira milleri: High Mains Formation (Late Ashgill (Hirnantian)). Genus Proturritella Koken, 1889 TYPE SPECIES.?Proturritella gracilis Koken, 1889. FKA.?Proturritella bicarinata Koken, 1925: Eines Forma? tion (Late Arenig (Kundan)). LKA.?Proturritella gracilis: Folkeslundekalk (Late Llan? virn (Lasnamagian)). Genus Eunema Salter, 1859 TYPE SPECIES.?Eunema strigillata Salter, 1859. FKA.?Lophospira centralis (Ulrich and Scofield, 1897): Murfreesboro Limestone (Early Caradoc (Ashbian)). See Wag? ner (1999). LKA.?Lophospira quadrisulcata (Ulrich and Scofield, 1897): Maquoketa Formation (Early Ashgill (Maysvillian)). See Wagner (1999). Genus Gyronema Ulrich in Ulrich and Scofield, 1897 TYPE SPECIES.?Gyronemapulchellum Ulrich and Schofield, 1897. FKA.?Gyronema pulchellum: Platteville Limestone (Early Caradoc (Black Riverian)). LKA.?Gyronema quadrisulcata (Ulrich and Scofield, 1897): Rockland Formation (Middle Caradoc (Rocklandian)). Genus Ruedemannia Foerste, 1914 Coronitla Pemer, 1907. [Not Beneden, 1871.] Ptychozone Pemer, 1907. ISchizolopha Ulrich and Scofield, 1897. TYPE SPECIES.?Ruedemannia lirata (Ulrich in Ulrich and Schofield, 1897). FKA.?Ruedemannia humilis (Ulrich and Scofield, 1897): Lexington Limestone (Middle Caradoc (Shermanian)). LKA.?Ruedemannia is reported as late as the Tournaisian. REMARKS.?This definition includes the types of two genera (Schizolopha and Ptychozone) that should have taxonomic pri- 90 ority; however, both genera are (1) monotypic, (2) known from very few specimens, and (3) very derived relative to other spe? cies in the genus. Also, neither genus has been used, whereas Ruedemannia has been used extensively. Thus, usage of the two genera (which has been nearly nonexistent) should be dis? continued. SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY Genus Trochonema Salter, 1859 TYPE SPECIES.?Trochonema umbilicata (Hall, 1847). FKA.?Trochonemella trochonemoides (Ulrich and Scofield, 1897): Valcour Formation (Late Llandeilo). LKA.?Trochonemapanderi Koken, 1925: Borkholm For? mation (Late Ashgill (Porkuni)). Genus Loxoplocus Fischer, 1885 Kiviasukkaan Peel, 1975b. TYPE SPECIES.?Loxoplocus solutus (Whiteaves, 1884). FKA.?Lophospira sedgewicki Donald, 1905: Girvan Lime? stone (Early Ashgill (Pusgillian)). LKA.?Lophospira gotlandica Ulrich and Scofield, 1897: Kopanina Formation (Late Ludlow (Ludfordian)); REMARKS.?Worthenia likely is derived from this genus, which extends its range past the Silurian. Analyses place the monotypic Kiviasukkaan as the sister taxon of the type species, Loxoplocus solutus, with the ubiquitous L. gotlandicus seem? ingly ancestral to that clade (Wagner, in prep.). Genus Longstaffia Cossman, 1908 TYPE SPECIES.?Longstaffia laquetta (Lindstrom, 1884). FKA.?Longstaffia centervillensis (Foerste, 1923): Beech- hill Cove Formation (Middle Llandovery (Rhuddanian)). LKA.?Longstaffia is reported as late as the Eifelian. Genus Arjamannia Peel, 1975 TYPE SPECIES.?Arjamannia cancellata (M'Coy in Sedg? wick and M'Coy, 1852). FKA.?Lophospira bellicarinata (Longstaff, 1924): Shal- loch Mill Formation (Late Caradoc (Onnian)). LKA.?Arjamannia aulongensis Peel, 1975a: Doctors Brook Formation (Middle Wenlock (Whitwellian)). Genus Trochonemella Okulitch, 1935 TYPE SPECIES.?Trochonemella notablis (Ulrich in Ulrich and Schofield, 1897). FKA.?Trochonemella knoxvillensis (Ulrich and Scofield, 1897): Lenoir Limestone (Late Llanvirn). LKA.?Trochonemella churkini Rohr, 1988, and T. reusingi Rohr and Blodgett, 1985: Port Clarence Limestone (Ashgill (Richmondian)). Family TROCHONEMATIDAE Zittel, 1895 REMARKS.?A greatly revised definition and diagnosis of this taxon was presented in Wagner (1999). Genus Globonema Wenz, 1938 TYPE SPECIES.?Globonema bicarinata (Koken, 1925). FKA.?Trochonema salteri Ulrich and Scofield, 1897: Fu- sispira Beds, Prosser Formation (Middle Caradoc (Rocklan- dian)). LKA(?).?Eunema murricata Lindstrom, 1884: Unit A, Hamra Formation (Late Ludlow (Ludfordian)). Superfamily STRAPAROLLINOIDEA, new superfamily Family STRAPAROLLINIDAE, new family REMARKS.?This new superfamily and family represents an early experiment with trochoid-like shell designs. Its members have been assigned to the Microdomatidae and Holopeidae. It is diagnosed by a very small and a strong but dull monolineate peripheral band. Post-Early Ordovician species are diagnosed by a nearly absent sinus and a contiguous coiling of the whorls that yields no columella. Genus ^Straparollina Billings, 1865 FIGURES 20, 21 TYPE SPECIES.?Straparollinapelagica Billings, 1865. FKA.?Turritoma aff. T. acrea: Fort Ann Limestone (Late Tremadoc (Demingian)). LKA.?Straparollina erigione Billings, 1865, and S. circe Billings, 1865: Rockland Formation (Middle Caradoc (Rock- landian)). ADDITIONAL ASSIGNED SPECIES.?Holopea hennigsmoeni Yochelson, 1963; Hormotoma dubia Cullison, 1944; Lophos? pira grandis Butts, 1926; Plethospira! turgida (Hall, 1847); Raphispira martinensis Rohr, 1996. REMARKS.?Straparollina is used herein to label the basal paraclade, which terminates in the extinction of a Middle Or? dovician clade. Post-Caradocian species assigned to Straparol? lina appear to belong to the Anomphalidae. Genus Daidia Salter, 1859 FIGURE 21 TYPE SPECIES.?Daidia cerithioides Salter, 1859. FKA.?Daidia cerithioides: Paquette Rapids Black River Formation (Early Caradoc (Black Riverian)). NUMBER 88 91 LKA.?Daidia aff. D. cerithioides in Rohr, 1988: Port Clar? ence Limestone (Ashgill (Richmondian)). REMARKS.?This represents a very small, highly derived clade of straparollinoids. Genus Haplospira Koken, 1897 FIGURE 21 TYPE SPECIES.?Haplospira variablis Koken, 1897. FKA.?Haplospira !nereis (Billings, 1865): Paquette Rapids Black River Formation (Early Caradoc (Black Riverian)). LKA.?Haplospira sibeliuxeni Peel, 1977: Stonehouse For? mation (Pridoli). Superfamily SUBULITOIDEA Lindstrom, 1884 Family SUBULITIDAE Lindstrom, 1884 REMARKS.?The systematics of this clade will be treated separately (Erwin, in prep.), so I present only a summary of the Ordovician-Silurian genera here. The basal genus of the clade is described here in some detail as it provides a link between "murchisonioid"-grade gastropods and this highly derived clade. Genus fEroicaspira, new genus FIGURE 22 TYPE SPECIES.?Hormotoma bellicincta Hall, 1847, sensu Ulrich and Scofield, 1897. FKA.?Hormotoma artemesia (Billings, 1865): Roubidoux Formation (Late Tremadoc (Demingian)). LKA.?Hormotoma bellicincta: Hudson River Formation (Ashgill (Richmondian)). ADDITIONAL ASSIGNED SPECIES.?Hormotoma augustina (Billings, 1865); H zelleri Flower, 1968b; Hormotoma Setul Formation species (see Kobayashi, 1959). ETYMOLOGY.?Named for Beethoven's Third Symphony, which possesses some of the innovations diagnosing Roman? tic music while retaining elements of the ancestral Classical form. REMARKS.?This paraclade is monophyletic if the earliest species is omitted. It is diagnosed by the strongly asymmetri? cal sinus of derived Hormotoma as well as a siphon formed from a twisting of the base of the columella. The subclade of post-Early Arenig species also is diagnosed by unusually large sizes. Genus fSubulites Emmons, 1842 FIGURE 22 TYPE SPECIES.?Subulites elongata Emmons, 1842. FKA.?Subulites sp.: El Paso Limestone (Early Arenig (Cassinian)). LKA.?Subulites is recorded as late as the Gedinnian. Genus Fusispira Hall, 1872 FIGURE 22 TYPE SPECIES.?Fusispira ventricosa Hall, 1872. FKA.?Fusispira sp.: Smithville Formation (Early Arenig (Cassinian)). LKA.?Blodgett and Johnson (1992) recorded Fusispira from Tortodus kockelianus-zones of the Roberts Mountain and Lone Mountain Limestones (Late Eifelian). Genus Cyrtospira Ulrich in Ulrich and Scofield, 1897 TYPE SPECIES.?Cyrtospira tortilis Ulrich in Ulrich and Scofield, 1897. FKA.?Cyrtospira raymondi Ulrich and Scofield, 1897: Chazy Limestone (Early Llandeilo). LKA.?Cyrtospira ventricosus Lindstrom, 1884: Kopanina Formation (Late Ludlow (Ludfordian)). Conclusions Phylogenetic analyses of early Paleozoic "archaeogastro? pods" suggest that (1) "archaeogastropods" evolved in the Late Cambrian from bellerophonts with monolineate periph? eral bands and deep sinuses but no slits; (2) "archaeogastro? pods" quickly diverged into two clades, corresponding best with previous definitions of the Euomphalina and Murchisoni? ina; (3) sinuitid gastropods might represent secondarily de? rived bellerophonts, which would be the sister taxon of the "murchisoniinae"; (4) the Pleurotomarioidea do not represent a paraphylum containing the ancestors of more-derived gastro? pods but instead represent a polyphyletic assemblage of "eu? omphalinae" and "murchisoniinae" subclades; (5) trochoid- like and apogastropod-like morphologies evolved nearly as frequently as pleurotomarioid-like morphologies; and (6) the paleontological hypothesis presented is easily reconciled with neontological hypotheses. In many ways, the results presented herein better match the estimates of 19th century workers, which were based on early Paleozoic material, than more recent estimates, which appar? ently were based largely on middle to late Paleozoic or Recent species. This analysis cannot directly assess the relationships among extant gastropods, but it does put parameters on viable hypotheses and sets the stage for more inclusive analyses that could further the value of a rich fossil record for determining relationships among extant gastropods. Appendices 1-3 Appendix 1. Characters and Character States A superscript letter "S" denotes characters weighted by one-half to account for symmetry/ asymmetry. A superscript letter "C" denotes continuous characters, which are weighted 1/ (n-1), where n is the number of states. Other multistate characters are considered unor? dered. 1. Sinus 1. Absent 2. Present 2. Symmetry of sinus depth 1. Angle of right side greater (as on "clathrospirids") 2. Symmetrical 3. Angle of left side greater (as on "hormotomoids") s > c3. Sinus depth on right side (= angle of sinus retreat) 0. >0? 1. -10? 2. -20? 3. -30? 4. -40? 5. -50? 6. -60? 7. -70? s , c4. Sinus depth on left side (= angle of sinus retreat) 0. >0? 1. -10? 2. -20? 3. -30? 4. -40? 5. -50? 6. -60? 7. -70? 5. Sinus width symmetry 1. Right side wider (as on Clathrospira species and de? scendants) 2. Symmetric 3. Left side wider (as on Hormotoma confusa and rela? tives) s , c6. Width of right side of sinus (NOTE: variation in the ramp length affects sinus width without changing the char? acter state.) 1. Just above peripheral band 2. Between peripheral band and top of right ramp 3. At top of right ramp 4. Above top of right ramp s ' c7. Width of left side of sinus 1. Just below peripheral band 2. Between peripheral band and alveozone (= left ramp) 3. At base of alveozone 4. Between alveozone and inner margin 8. Symmetry of sinus shape 1. More acute curve on the left (as on Hormotoma spe? cies) 2. Symmetric 3. More obtuse curve on the left (as on Clathrospira or Pleurorima) s9. Shape of right side of sinus 1. Plateaus before apex (half a "U" as on Raphistoma and Raphistomina) 2. Straight into apex (as on Ectomaria) 3. Continuous curve towards apex (as on Hormotoma) 4. Hyperbolic curve towards apex (as on Climacora? phistoma) s 10. Shape of left side of sinus 1. Plateaus before apex (half a "U" as on as on Raphis? tomina) 2. Straight into apex 3. Continuous curve towards apex (as on Clathrospira) 4. Hyperbolic curve towards apex (as on Hormotoma) 11. Wrinkled right sinus (= a strong kink in the sinus, as on Raphistoma) 1. Absent 2. Present 12. Crenulated aperture (usually represented by undulat? ing growth lines) 1. Absent 2. Present 13. Aperture with reverse sigma shape (when viewed from the side) 1. Absent 2. Present (as on "helicotomids") 14. Extent of sigmoidal aperture 1. Absent 2. Sigmoidal 3. Hypersigmoidal 15. Growth-line prominence 1. Not visible 2. Weak 3. Fine sharp 4. Strong 5. Extremely strong 16. Ontogenetic change in growth-line strength 1. No change over ontogeny 93 94 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY 2. Stronger on juveniles (as on Lesueurilla) 17. Imbricated growth lines 1. Absent 2. Weak 3. Moderate 4. Strong 18. Growth lines on base 1. Same 2. Stronger than on the rest of shell (as on Malayaspira) 19. Peripheral band 1. Absent 2. Present c20. Peripheral-band width 1. 1 = 05? 2. 1 = 1 0 ? 3. 1 = 1 5 ? 4. 1 = 20? 5. 1 = 2 5 ? 6. I = 30? 21. Peripheral lira (i.e., lira bordering the peripheral band; present with "single" lirum (27) only if there is on? togenetic change (41) or if the peripheral band is trilineate) 1. Absent 2. Present 22. Peripheral-lira shape 1. Round (as on Hormotoma) 2. Sharp (as on Ectomaria and Lophospira) 3. Square (i.e., with edges as on Eotomaria and Parali? ospira) 23. Peripheral-lira strength 1. Extremely weak 2. Weak 3. Moderate 4. Strong 5. Extremely strong 24. Peripheral lira forming a frill 1. Absent 2. Present (as on Conotoma and Crenilunula) 25. Frill strength 1. Weak 2. Strong 26. Relative strength of right and left peripheral-band lira 1. Same 2. Right lirum stronger than left lirum (as on Conotoma and Crenilunula) 27. Medial lirum (= single keel at apex of sinus; present with peripheral lira (21) only if there is ontogenetic change (41) or if the peripheral band is trilineate) 1. Absent 2. Present 28. Medial lirum type 1. Rounded (as on Schizopea) 2. Sharp (as on Barnesella and Malayaspira) 3. Flange (as on Lesueurilla) 4. Squared ridge (as on Ophiletina) 5. Dull lump (as on Euomphalopterus) 29. Single peripheral-lira strength 1. Very weak 2. Weak 3. Moderate 4. Strong 5. Very strong 6. Extremely strong 30. Ontogenetic change in single peripheral-lira strength 1. No change over ontogeny 2. Becomes weaker 3. Becomes stronger 31. Single peripheral-lira tubes 1. Absent 2. Present 32. Imbricated single peripheral lira 1. Consistent 2. Flares periodically (as on Poleumita discors and rela? tives) 33. Peripheral-band prominence 1. None 2. Slight 3. Strong 34. Slit 1. Absent 2. Present 35. Continuity of slit 1. Periodic slit (i.e., absent periodically as on some Clathrospira) 2. Continuous slit (as on Pleurorima and relatives) 36. Width of slit relative to peripheral band 1. Subsuming peripheral band (as on Pararaphistoma) 2. Within peripheral band (as on Pleurorima and rela? tives) 3. Thinner than peripheral band (as on Oehlertia) 37. Lira within peripheral band (usually bordering thin slits (36)) 1. Absent 2. Two lira bordering slit (as on Oehlertia) 38. Lunulae shape 1. Concentric 2. Zipper-like (as on Crenilunula) 3. Sigma-shaped (as on Pararaphistoma) 4. V-shaped (as on Eccyliopterus) 5. Kinked (as on onychochilids) 6. Straight (as on Oehlertia) 39. Lunulae strength relative to the growth lines 1. Weaker than growth lines (as on Liospira or Pleuror? ima) 2. Same as growth lines 3. Stronger than growth lines (as on Brachytomaria bal? tica and relatives) NUMBER 88 95 40. Ontogenetic change in peripheral-band prominence 1. None 2. Increasing prominence (as on Straparollina) 3. Decreasing prominence (as on Catazone) 41. Ontogenetic change from a bilineate to a monolineate pe? ripheral band 1. Bilineate throughout ontogeny 2. Becoming monolineate over ontogeny (as on early Straparollina) 42. Peripheral-band attitude 1. Projecting normally to the aperture 2. Peripheral band curves adapically (as on "lesueur- illids" or Helicotoma) 43. Symmetry of peripheral band relative to the aperture 1. Asymmetrical on alveozone (as on "gosseletides") 2. Bisects whorl 3. Asymmetrical on right ramp (as on Lesueurilla and Eccyliopterus) 4. Entirely on right ramp (as on Eotomaria and Para? liospira) 44. Ontogenetic change in peripheral-band symmetry 1. No change 2. Moving from the right ramp to bisecting aperture (as on Paraliospira) 45. Channel beneath peripheral band 1. Absent 2. Present (as on Lophospira perangulata) 46. Channel strength 1. Weak 2. Strong 47. Thickenings on either side of peripheral band (NOTE: these might be homologous with the right and left ramp carina) 1. Absent 2. Present (as on Oehlertia) c48. Position of anal notch (based on plane passing through centroid and sinus apex; if perpendicular to inner margin, (3 = 0?; if parallel to inner margin and ori? ented adapically, (3 = 0?; if parallel to inner margin and oriented abapically, p = 180?). 1. P = 150? (e.g., Prosolarium) 2 . 3 = 140? 3. 0=130? 4. p = 120? 5. p=110? 6. P = 100? 7. P = 90? 8. P = 80? 9. p = 70? A. P = 60? B. p = 50? C. p = 40? D. p = 30? E. P = 20? F. p = 10? (e.g., Centrifugus) 49. Ontogenetic change in P (due to rotation of the aperture clockwise or counter-clockwise) 1. Absent 2. Decreasing P (as on basal "raphistomatoids") 3. Increasing P (as on Paraliospira) 50. Magnitude of ontogenetic rotation 1. Small (<30?) 2. Great (> 30?) 51. Symmetry of ramp shapes 1. Right ramp rounder (as on "holopeides") 2. Symmetric 3. Alveozone rounder (as on "lesueurillids") s52. Right-ramp shape (see also 72) 1. Extremely globular (forming an acute angle) 2. Globular 3. Convex 4. Flat 5. Concave 53. Ontogenetic change in right-ramp shape 1. Ramp becoming more concave 2. Ramp becoming slightly more convex (as on Pachys? trophia) 3. Ramp becoming much more convex (as on Parara? phistoma) 54. Symmetry of ramp lengths 1. Right ramp longer (as on "luciellides") 2. Symmetric 3. Alveozone longer (as on Raphistoma striata and rela? tives) s , c 55. Right-ramp length (= degrees between peripheral band and the top of ramp) 0. <20? 1. -30? 2. -40? 3. -50? 4. -60? 5. -70? 6. -80? 7. -90? 8. -100? s , c56. Alveozone length (= degrees between peripheral band and the base of ramp) 0. <20? 1. -30? 2. -40? 3. -50? 4. -60? 5. -70? 6. -80? 7. -90? 8. -100? 9. -110? 57. Symmetry of ramp projections (see 58, 59) 1. Right higher (as on "helicotomids") 2. Symmetric 96 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY 3. Left higher (as on "holopeides") s ' c58. Right-ramp projection (= angle between peripheral band and top of ramp) 1. -20? 2. -30? 3. -40? 4. -50? 5. -60? 6. -70? 7. -80? 8. -90? 9. -100? s ' c59. Alveozone projection (= angle between peripheral band and base of ramp) 0. -20? 1. -30? 2. -40? 3. -50? 4. -60? 5. -70? 6. -80? 7. -90? 60. Thickening of shell at the top of right ramp 1. Absent 2. Present (as on early "archaeogastropods" and bellero? phonts) 61. Degree of thickening 1. Weak 2. Moderate 3. Strong 4. Very strong c62. Ontogenetic change in right-ramp swelling 1. Always acute (as on Ecculiomphalus) 2. Going from acute to obtuse over ontogeny (as on Ma? layaspira) 3. Always obtuse 63. Ontogenetic change in right ramp 1. None 2. Ramp becoming shorter and rounder (as on Mala? yaspira) 64. Right-ramp carina (RRC in figure captions) 1. Absent 2. Present 65. Right-ramp carina strength (in terms of prominence) 1. Weak 2. Moderate (roughly equal to a weak-to-moderate pe? ripheral band) 3. Strong (roughly equal to a strong peripheral band) 66. Right-ramp carina type 1. Dull thick thread 2. Dull thin thread 3. Round thin-to-moderately wide lira 4. Sharp thin-to-moderately wide thread 67. Ontogenetic change in right-ramp carina strength 1. None 2. Becomes weaker (as on Raphistomina species) 3. Becomes sharper (as on Raphistoma species) 68. Channel beneath right-ramp carina 1. Absent 2. Present (as on "poleumitids") 69. Suture type 1. Right ramp oblique at suture 2. Right ramp acute at suture 70. Degree of acuteness 1. Acute (as on subulitids) 2. Attenuated (as on Hormotoma salteri) 11. Aperture curving back at suture 1. Absent 2. Present (as on Liospira) 72. Alveozone shape (see also 52) 1. Extremely globular (forming an acute angle) 2. Globular 3. Convex 4. Flat 5. Concave 73. Thickening of shell at the base of alveozone 1. Absent 2. Present 74. Degree of thickening 1. Weak 2. Moderate 3. Strong 4. Very strong 75. Alveozone carina 1. Absent 2. Present 76. Alveozone-carina type 0. Dull thick thread 1. Thick swelling of shell 2. Sharp thin-to-moderately wide thread 3. Frill (as on Euomphalopterus species?go to charac? ter 79) 4. Square thread (as on Ophiletina sublaxa) 11. Alveozone-carina strength 1. Weak 2. Moderate (= peripheral band) 3. Strong 4. Very strong 78. Channeled alveozone carina 1. Absent 2. Present 79. Frilled alveozone carina 1. Peg (as on Euomphalopterus cariniferus) 2. Weak frill (as on Poleumita alatum) 3. Extended frill (as on Euomphalopterus alatus) 80. Frill bordered by lira 1. Absent 2. Present (as on Euomphalopterus togatus) 81. Crenulated frill 1. Absent NUMBER 88 97 2. Present (as on Euomphalopterus undulans) 82. Channeled frill 1. Absent 2. Present (as on Euomphalopterus alatus) 83. Projection of frill relative to coiling axis 1. Behind aperture 2. Parallel to aperture 3. In front of aperture 84. Hood-like frill 1. Absent 2. Present (as on Pseudophorus praetextus) 85. Alveozone-carina tubes 1. Absent 2. Present (as on Euomphalopterus togatus) 86. Increasing expansion at base of alveozone over ontogeny (inducing a more "bowF'-like apical umbilicus) 1. Absent 2. Present (as on Teiichispira) 87. Inner-margin (= columellar lip of most species) thickness, relative to rest of shell 1. No different 2. Thicker 3. Much thicker 88. Thickened region(s) of inner margin 1. Only top of inner margin thicker 2. Entire inner margin thicker 3. Inner margin and base thicker 4. Only base of inner margin thicker 89. Basal carina (BC in figure captions) 1. Absent 2. Present 90. Basal-carina type 1. Narrow thickening on the base 2. Dull extension with channel 3. Sharp thread 4. Rounded thread 5. Peg 91. Prominence of basal carina 1. Weak 2. Strong 3. Projecting 92. Ontogenetic change in basal carina 1. None 2. Becomes weaker with age 3. Becomes stronger with age 93. Position of basal carina 1. Beneath inner margin of aperture 2. Middle of whorl 3. Beneath outer margin of aperture c94. Angle between inner margin and base 0. <45? 1. >45? 2. >60? 3. >75? 4. >90? 5. >105? 6. >120? 95. Shape of inner margin on the inner margin of aperture 1. Round 2. Curved 3. Straight 4. Arched inwards 96. Shape of inner margin on the outer margin of aperture 1. More obtuse than the inner margin 2. Same 3. More acute than the inner margin 97. Ontogenetic change in inner-margin shape 1. None 2. Becomes rounder with age c98. Inner-margin attitude relative to coiling axis 1. -0? 2. -15? 3. -30? 4. -45? 5. -60? 6. -75? 7. -90? 8. -105? 9. -120? 99. Siphon 1. Absent 2. Slight twist of inner margin 3. Strong extension of inner margin 100. Inner-margin lira 1. Absent 2. Present 101. Type of inner-margin lira 1. Lira (as on Paraliospira) 2. Callous (as on Pycnomphalus) 102. Inner-margin channel 1. Absent 2. Present 103. Parietal-inductura strength 1. Absent 2. Thin and incomplete 3. Thin 4. Same as shell 5. Thicker than shell 104. Inductura projection 1. Parallels rest of aperture 2. Extended in front of aperture (as on Liospira) 3. A thick, concentrate strip extended in front of aper? ture (as on Poleumita) 4. Fills umbilicus (as on Eotomaria) 105. Flaring aperture 1. Absent 2. Weak (as on Loxonema) 3. Strong (as on Gasconadia) 106. Reflected inner-margin lip 1. Absent 98 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY 122. c123 2. Present (as on basal "hormotomoids") 118. 107. Funicle 1. Absent 2. Present (as on Liospira) c l 19. 108. Inner margin fills umbilicus 1. Umbilicus open or filled by inductura 2. Inner margin fills umbilicus 3. Inner margin contiguous with previous whorl 4. Inner margin folds back into umbilicus 109. Whole aperture inclined (species with left and right halves 120. both inclined (111 and 113), but to different degrees, were scored as unknown (?)) 1. Radial/backwards inclination (see 115) 2. Left and right side inclined to same degree (as on Stra- c 121. parollina) -110. Angle of inclination of whole aperture 1. InAn= 10? 2. InAn = 20? 3. InAn = 30? 4. InAn = 40? 5. InAn = 50? 6. InAn = 60? 111. Inclination of left half of aperture 1. Radial 2. Inclined (right side different or radial; e.g., Clath? rospira) -112. Inclination of left side of aperture 0. InAn =5? 1. InAn= 10? 2. InAn = 20? 3. InAn = 30? 4. InAn = 40? 113. Inclination of right half of aperture 1. Radial 2. Right side inclined (left side different or radial; e.g., 124. Pleurorima) "114. Inclination of right side of aperture 1. InAn= 10? 125 2. InAn = 20? 3. InAn = 30? 4. InAn = 40? 115. Aperture inclined "backwards" c126. 1. Absent/forward inclination 2. Present (as on onychochilids) 116. Posterior projection of aperture base 1. Absent (as on all species with anterior projections; see also 120) 2. Present (as on Schizopea) c117. Magnitude of posterior projection 1. ~ -10? 2. ~ -20? 3. ~-30? 127. 4. ~-40? 5. ~-50? Anterior projection of aperture base 1. Absent 2. Present (as on "helicotomids" and "eotomarioids") Magnitude of anterior projection 1. -10? 2. -20? 3. -30? 4. -40? 5. -50? Shape of inner-margin base 1. Straight 2. Excavated (curved) (as on Spiroraphe) 3. Excavated (crenulated) (as on Lesueurilla) Shell expansion (in radians) 1. E < 0.05 (extremely low) 2. 0.05 0.25 (very high) 7. E > 1.0 (extremely high) Ontogenetic change in expansion 1. Decreasing expansion over ontogeny 2. None 3. Increasing expansion over ontogeny Shell curvature around coiling axis (in radians) 0. K < 0.40 (open coiling) 1. K < 0.5 (extremely low) 2. 0.5 < K < 0.55 (very low) 3. 0.55 < K < 0.65 (low) 4. 0.65 < K < 0.75 (moderate) 5. 0.75 < K < 0.85 (high) 6. 0.85 < K < 0.95 (very high) 7. K > 0.95 (extremely high) Ontogenetic change in curvature 1. No change over ontogeny 2. Decreasing curvature over ontogeny Anisostrophy (applies only to coiling, not left-right asymmetry) 1. Isostrophic 2. Anisostrophic Shell torque 0. High ultradextral 1. Moderate ultradextral 2. Low ultradextral 3. Nearly planispiral 4. Low dextral 5. Moderate dextral 6. High dextral 7. Very high dextral 8. Extremely high dextral Ontogenetic change in shell torque 1. Increasing torque over ontogeny 2. Isometric NUMBER 88 99 3. Decreasing torque over ontogeny 128. Septation 1. Absent 2. Septation present (as on most "euomphalinae") 3. Complete filling of juvenile whorls (as on Palliseria) 129. Ornament on left side of aperture 1. Absent 2. Present throughout the alveozone and base 3. Present on the alveozone only 130. Density of ornament 1. One thread per 20? 2. One thread per 10? 3. One thread per 5? 4. One thread per 1 ? 131. Strength of alveozone ornament 1. Fine threads 2. Weak lira 3. Strong lira 132. Consistency of alveozone ornament 1. Uniform 2. Threads stronger higher on the alveozone 3. Threads stronger lower on the lower ramp 4. Highest threads strong, the rest uniform 133. Ornamentation of right ramp 1. Absent 2. Present throughout 3. Ornament present on upper half of right ramp only 134. Density of ornament on right side 1. One thread per 20? 2. One thread per 10? 3. One thread per 5? 4. One thread per 1? 135. Strength of ornament on right side 1. Fine threads 2. Weak lira 3. Strong lira 136. Pattern of ornament on right side 1. Uniform 2. 2:1:2 (every other thread twice as strong as interme? diate thread) 137. Type of right-ramp ornament 1. Local thickenings on shell 2. Local changes in aperture shape 138. Transverse ornament 1. Absent 2. Present (as on Cataschisma) 139. Peripheral-band ornament 1. Absent 2. Present (as on Crenilunula) 140. Carrier-shell scars 1. No scars 2. Scars (as on Lytospira) 141. Size (= shell volume) 0. Very small (micro-mollusc: < 10 mm3) 1. Small(>10mm3<102mm3) 2. Moderate (> 102 mm3 < 103 mm3) 3. Large(>103mm3<104mm3) 4. Huge(>104mm3<105mm3) 142. Protoconch coiling 1. Like teleoconch 2. Planispiral (as on Sinuspira) 143. Protoconch size 1. Small (< 10-'mm) 2. Large (> 10_1 mm, as on Murchisonia and Loxonema) Appendix 2. Data Matrix Data matrix for the analyzed species. Character numbers correspond to those in Appendix 1, and species numbers correspond to those in Appendix 3. Outgroup species have letters rather than numbers. 100 NUMBER 88 101 Number Species AC. AB. AA. Z. Y X. W. V. u. T. S. R. Q. p. 0 . N. M. L. K. J. I. H. G. F. E. D. C. B. A. 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12. 13. 14. 15. 16. 17. 18. 19. 20. 21. 22. 23. 24. 25. 26. 27. 28. 29. 30. 31. Mollusc Helcionella subrugosa Latouchella merino Pelagiella subangulata Costipelagiella zazvorkai Oelandia rugosa Coreospira rugosa Sinuella minuata Chippewaella patellitheca Strepsodiscus major Chalarostrepsis praecursor Eobucania mexicana Strepsodiscus paucivoluta Modestospira poulseni Peelerophon oehlerti Kiringella pyramidal is Cyrtolites sp. Macluritella? walcotti Euomphalopsis involuta "Maclurites" thomsoni Kobayashiella circe Scaevogyra swezeyi Matherella saratogensis Matherellina walcotti Hypseloconus elongatus Knightoconus antarcticus Sinuites sowerbyi Cloudia buttsi Owenella antiquata Dirhachopea normalis Dirhachopea subrotunda Schizopea typica Sinuopea sweeti Taeniospira emminencis Ceratopea canadensis Gasconadia putilla Jarlopsis conicus Ophileta supraplana Rhombella umbilicata Prohelicotoma uniangulata Sinuopea basiplanata Taeniospira 1st. clairi Bridgeites Idisjuncta Bridgeites planodorsalis Bridgeites supraconvexa Euconia etna Ceratopea 1 laurentia Ceratopea pygmaea Orospira bigranosa Macluritella stantoni Teiichispira odenvillensis Teiichispira loceana Palliseria robusta Mitrospira longwelli Teiichispira kobayashi Teiichispira sylpha Monitorella auricula Maclurites magna "Eccyliopterus ornatus" Maclurites bigsbyi I ? 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 ? n n n n n n 2 2 2 2 2 2 2 2 n n n 2 n n n n n n n 2 2 2 2 2 2 2 2 2 2 1 2 1 2 2 2 2 2 2 1 2 2 2 2 2 2 2 2 2 2 2 2 3 2 3 ? n n n n n n 6 4 6 6 6 6 6 6 n n n 1 n n n n n n n 3 3 2 6 5 6 4 4 6 6 6 6 6 3 4 4 6 6 6 6 6 6 5 2 2 2 1 1 2 1 2 2 1 2 4 ? n n n n n n 6 4 6 6 6 6 6 6 n n n 1 n n n n n n n 3 3 2 6 5 6 4 4 6 6 4 6 4 3 4 4 6 6 6 4 6 6 5 2 2 2 1 1 2 1 2 2 3 2 APPENDIX 2.? 5 ? n n n n n n 2 2 2 2 2 2 2 2 n n n 2 n n n n n n n 2 2 2 2 2 2 2 2 2 2 1 2 1 2 2 2 2 2 2 1 2 2 2 2 2 2 2 2 2 2 2 2 2 2 6 ? n n n n n n 3 2 3 2 2 3 3 2 n n n 1 n n n n n n n 2 2 2 3 3 3 2 3 3 1 2 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 2 2 3 2 3 1 3 1 7 ? n n n n n n 3 2 3 2 2 3 3 2 n n n 1 n n n n n n n 2 2 2 3 3 3 2 3 3 1 2 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 2 2 3 2 3 1 3 1 8 ? n n n n n n 2 2 2 2 2 2 2 2 n n n 2 n n n n n n n 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 3 2 -Continued. 9 ? n n n n n n 4 4 4 4 4 4 3 4 n i i n 3 n n n n n n n 1 1 1 4 4 4 1 3 4 4 4 4 4 2 3 3 4 4 4 4 4 4 4 2 2 2 2 2 2 2 2 2 3 2 1 0 ? n n n n n n 4 4 4 4 4 4 3 4 n n n 3 n n n n n n n 1 1 1 4 4 4 1 3 4 4 4 4 4 2 3 3 4 4 4 4 4 4 4 2 2 2 2 2 2 2 2 2 2 2 1 1 7 n n n n n n n 1 n n n n n n n 2 1 2 . ? n n n : n ; n n n n n ' 1 n n 'J. n I n n n n 2 1 1 5 4 ' ? n n > 1 > 1 n n n n n n n n n n n n n n n . n . n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n I 1 5 6 ? ? 4 ? 5 1 2 1 4 1 4 ? 4 1 4 1 4 1 4 1 4 1 3 1 4 1 4 1 5 1 3 i : 4 1 4 1 2 1 4 1 5 1 4 1 5 1 5 1 4 1 4 1 4 1 4 1 4 1 4 1 4 1 4 1 4 1 4 1 4 1 1 1 4 1 4 1 4 1 4 1 4 I 4 1 4 ? 4 1 4 1 2 1 4 1 3 1 3 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 3 ? 2 1 4 1 1 1 1 7 8 9 ? ? ? ? ? 1 \ 1 1 1 2 1 2 1 2 1 2 1 2 1 1 1 2 > 1 1 ! 1 2 2 2 1 1 1 2 1 1 1 2 1 1 1 2 1 1 1 1 1 1 1 1 1 1 1 2 1 2 1 2 1 1 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 ? 2 1 2 1 2 1 2 2 2 1 2 2 5 1 2 } 1 2 1 1 2 1 1 2 1 1 2 1 1 2 1 1 2 1 1 2 1 1 2 1 2 2 1 2 2 1 2 2 2 0 ? n n n n n n n 2 3 3 2 4 n 3 n 3 2 n 1 n 2 n 2 n n n n n 3 3 3 n 4 3 1 2 3 4 2 4 5 2 3 3 3 3 3 3 1 1 1 2 2 1 2 2 1 2 1 2 1 ? n n n n n n n 1 1 2 2 1 n 2 n 1 1 n 1 n 1 n 1 n n n n n 2 2 n 2 2 2 2 2 2 7 n n n n n n n n n 2 2 n n 2 n n n n n n n n n n n n n n 1 1 n n 1 n 1 n n n n 1 1 n n n n n n n n n n n n n n n n n n 2 3 7 n n n n n n n n n 1 1 n n 4 n n n n n n n n n n n n n n 2 2 n n 3 n 2 n n n n 2 2 n n n n n n n n n n n n n n n n n n 2 4 ? n n n n n n n n n 1 1 n n 1 n n n n n n n n n n n n n n 1 1 n n 1 n n n n n n 1 1 n n n n n n n n n n n n n n n n n n 2 5 7 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n ?i n n n n n n n n n n n n n n n n n n n n n n n n n n n 102 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY APPENDIX 2.?Continued. Number Species 2 2 6 7 3 3 2 3 3 3 3 4 5 6 4 4 4 1 2 3 4 4 4 4 5 6 4 4 7 8 4 5 9 0 AC. Mollusc AB. Helcionella subrugosa AA. Latouchella merino Z. Pelagiella subangulata Y Costipelagiella zazvorkai X. Oelandia rugosa W. Coreospira rugosa V. Sinuella minuata U. Chippewaella patellitheca T. Strepsodiscus major S. Chalarostrepsis praecursor R. Eobucania mexicana Q. Strepsodiscus paucivoluta P. Modestospira poulseni 0 . Peelerophon oehlerti N. Kiringella pyramidalis M. Cyrtolites sp. L. Macluritella? walcotti K. Euomphalopsis involuta J. "Maclurites" thomsoni I. Kobayashiella circe H. Scaevogyra swezeyi G. Matherella saratogensis F. Matherellina walcotti E. Hypseloconus elongatus D. Knightoconus antarcticus C. Sinuites sowerbyi B. Cloudia buttsi A. Owenella antiquata 1. Dirhachopea normalis 2. Dirhachopea subrotunda 3. Schizopea typica 4. Sinuopea sweeti 5. Taeniospira emminencis 6. Ceratopea canadensis 1. Gasconadia putilla 8. Jarlopsis conicus 9. Ophileta supraplana 10. Rhombella umbilicata 11. Prohelicotoma uniangulata 12. Sinuopea basiplanata 13. Taeniospira 1st. clairi 14. Bridgeites Idisjuncta 15. Bridgeites planodorsalis 16. Bridgeites supraconvexa 17. Euconia etna 18. Ceratopea? laurentia 19. Ceratopea pygmaea 20. Orospira bigranosa 21. Macluritella stantoni 22. Teiichispira odenvillensis 23. Teiichispira ?oceana 24. Palliseria robusta 25. Mitrospira longwelli 26. Teiichispira kobayashi 27. Teiichispira sylpha 28. Monitorella auricula 29. Maclurites magna 30. "Eccyliopterus ornatus" 31. Maclurites bigsbyi n n n n n n n n n 2 n 2 n n n 2 n n n 2 n n n n n 2 2 2 n n n n n n " n n n n n n n n n n n n n n n n n 1 n n n n n n n n n n n n n n n n n n 2 1 2 n 1 2 2 2 2 2 2 1 1 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 1 n 1 n n 2 1 1 3 1 3 n n 4 2 2 1 3 3 2 1 3 3 1 1 1 1 1 1 1 1 2 n 2 n n 3 1 3 3 2 3 n n 3 3 3 2 4 4 3 3 3 3 2 2 2 3 3 3 3 2 1 n 1 n n 1 1 1 1 1 2 n n 1 1 1 1 1 1 1 2 2 2 1 1 2 1 2 1 n 1 n n n n 1 1 1 1 1 1 1 1 1 n n n n n n n n n n n n n n n 4 n 5 n 4 1 5 1 5 1 5 1 5 1 5 1 n 1 n 2 n n 2 2 2 2 2 n n 1 1 n 1 n 1 n 1 n n 1 1 NUMBER 88 103 APPENDIX 2.?Continued. Number Species AC. AB. AA. Z. Y. X. w. V. U. T. S. R. Q. P. 0 . N. M. L. K. J. I. H. G. F. E. D. C. B. A. 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12. 13. 14. 15. 16. 17. 18. 19. 20. 21. 22. 23. 24. 25. 26. 27. 28. 29. 30. 31. Mollusc Helcionella subrugosa Latouchella merino Pelagiella subangulata Costipelagiella zazvorkai Oelandia rugosa Coreospira rugosa Sinuella minuata Chippewaella patellitheca Strepsodiscus major Chalarostrepsis praecursor Eobucania mexicana Strepsodiscus paucivoluta Modestospira poulseni Peelerophon oehlerti Kiringella pyramidal is Cyrtolites sp. Macluritella? walcotti Euomphalopsis involuta "Maclurites" thomsoni Kobayashiella circe Scaevogyra swezeyi Matherella saratogensis Matherellina walcotti Hypseloconus elongatus Knightoconus antarcticus Sinuites sowerbyi Cloudia buttsi Owenella antiquata Dirhachopea normalis Dirhachopea subrotunda Schizopea typica Sinuopea sweeti Taeniospira emminencis Ceratopea canadensis Gasconadia putilla Jarlopsis conicus Ophileta supraplana Rhombella umbilicata Prohelicotoma uniangulata Sinuopea basiplanata Taeniospira ?st. clairi Bridgeites ?disjuncta Bridgeites planodorsalis Bridgeites supraconvexa Euconia etna Ceratopea ?laurentia Ceratopea pygmaea Orospira bigranosa Macluritella stantoni Teiichispira odenvillensis Teiichispira ?oceana Palliseria robusta Mitrospira longwelli Teiichispira kobayashi Teiichispira sylpha Monitorella auricula Maclurites magna "Eccyliopterus ornatus" Maclurites bigsbyi 5 1 7 2 2 3 2 2 2 2 2 2 2 2 2 2 2 2 2 1 2 3 3 3 3 3 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 1 2 1 1 1 1 2 3 3 2 3 3 1 3 3 3 3 3 5 2 7 2 1 3 2 2 2 3 2 2 2 2 2 2 2 2 3 2 1 3 4 3 4 4 2 2 2 2 2 2 2 2 2 2 2 3 2 2 2 1 2 2 2 2 2 2 2 2 3 1 2 1 3 3 4 4 4 4 3 3 5 3 7 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 1 1 1 2 2 1 2 1 3 1 3 5 4 7 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 ? 2 2 2 2 2 2 2 2 2 2 2 2 1 1 2 1 2 2 1 1 2 1 1 2 2 2 2 2 2 3 3 2 3 2 3 2 3 5 5 7 2 2 7 7 2 3 4 8 8 8 8 8 7 8 7 7 8 7 7 7 7 7 7 7 7 7 7 7 8 7 8 7 7 6 6 8 6 8 4 5 4&5 6 5 5 8 6 5 5 4 5 5 5 5 4 5 5 5 4 3 5 6 7 2 2 7 7 2 3 4 8 8 8 8 8 7 8 7 7 8 7 7 5 3 6 4 7 7 7 7 7 8 7 8 7 7 6 4 4 6 4 4 5 3 4 5 4 3 6 5 5 4 5 5 6 6 6 6 5 7 4 7 5 7 9 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 7 2 2 3 1 1 1 2 2 2 2 2 2 2 2 2 2 2 1 1 2 1 2 2 2 1 2 1 1 2 2 2 2 2 2 1 1 2 1 2 1 2 1 5 8 7 5 5 2 2 5 7 7 2 2 2 2 1 4 2 2 2 2 4 1 3 3 5 6 3 2 3 4 6 2 3 2 5 4 2 4 2 2 2 2 3 5 3 1 1 3 1 1 3 2 1 1 3 2 1 2 1 2 1 3 5 9 7 5 5 2 2 5 7 7 2 2 2 2 1 4 2 2 2 2 4 1 4 1 0 5 3 2 3 4 6 2 3 2 5 4 2 4 1 2 1 2 3 5 2 1 1 1 1 1 3 2 1 1 2 1 1 1 1 2 1 2 6 0 7 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 1 2 2 2 2 2 2 2 2 2 2 2 2 2 2 1 2 2 1 6 1 7 n n n n n 4 4 1 2 1 1 2 2 1 n n n n n n n n n n n 3 3 3 2 2 2 3 3 1 n 2 2 2 2 2 1 3 1 3 2 1 1 1 1 n 1 n n n n n n 2 n 6 6 2 3 7 ] n 1 n 1 n 1 n 1 n 1 1 1 1 1 3 1 2 1 3 1 3 1 3 1 3 1 3 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 3 1 3 1 3 1 2 1 3 1 2 1 3 1 3 1 3 1 n 1 2 1 2 1 2 1 2 1 3 1 3 1 2 1 3 1 2 1 2 1 3 1 n 1 3 1 2 1 n 1 2 1 n 1 n 1 n 1 n 1 n 1 n 1 3 1 n 1 6 6 4 5 7 7 1 n 1 n 2 2 1 n 1 n 1 " 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 i 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n ? n ? n I n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 2 3 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n I n 1 n 1 n 6 6 7 n n 1 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n 4 n n n n n n n n n n n 6 7 7 n n 1 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n 1 n n n n n n n n n n n 6 6 8 9 7 7 n 1 n 1 1 1 n 1 n 1 n 1 n 1 n ? n 1 n 1 n 1 n 1 n 1 n 1 n ? n ? n 1 n 1 n 1 n 1 n 2 n 2 n 2 n ? n ? n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n ] n 1 n n n n n 1 n n n n n 1 n n n n n n n n n n n 7 7 0 I 7 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 1 1 1 1 1 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n ! n n n n n u n n n n n n n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 7 2 7 2 1 2 2 2 2 3 2 2 2 2 2 2 2 2 3 3 1 1 1 2 2 2 2 2 2 2 2 2 2 2 2 2 2 3 2 2 2 1 2 2 3 3 3 3 3 3 3 I 1 1 2 1 1 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 7 3 7 2 1 2 2 2 2 2 2 2 2 2 2 2 2 2 2 1 2 2 2 2 2 2 2 7 7 4 5 7 ? n 1 n 1 n 1 n 1 n 1 n 1 4 ? 1 1 2 1 1 1 1 1 1 1 3 1 1 1 n ? n 2 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n ? n 1 3 ? 3 ? 3 ? 2 1 2 1 2 1 3 1 3 1 n 1 1 1 2 1 1 1 2 1 1 1 3 1 1 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 1 1 n 1 1 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 104 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY APPENDIX 2.?Continued. Number Species AC. AB. AA. Z. Y. X. w. V. u. T. S. R. Q. p. 0 . N. M. L. K. J. I. H. G. F. E. D. C. B. A. 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12. 13. 14. 15. 16. 17. 18. 19. 20. 21. 22. 23. 24. 25. 26. 27. 28. 29. 30. 31. Mollusc Helcionella subrugosa Latouchella merino Pelagiella subangulata Costipelagiella zazvorkai Oelandia rugosa Coreospira rugosa Sinuella minuata Chippewaella patellitheca Strepsodiscus major Chalarostrepsis praecursor Eobucania mexicana Strepsodiscus paucivoluta Modestospira poulseni Peelerophon oehlerti Kiringella pyram idalis Cyrtolites sp. Macluritella? walcotti Euomphalopsis involuta "Maclurites" thomsoni Kobayashiella circe Scaevogyra swezeyi Matherella saratogensis Matherellina walcotti Hypseloconus elongatus Knightoconus antarcticus Sinuites sowerbyi Cloudia buttsi Owenella antiquata Dirhachopea normalis Dirhachopea subrotunda Schizopea typica Sinuopea sweeti Taeniospira emminencis Ceratopea canadensis Gasconadia putilla Jarlopsis conicus Ophileta supraplana Rhombella umbilicata Prohelicotoma uniangulata Sinuopea basiplanata Taeniospira ?st. clairi Bridgeites ?disjuncta Bridgeites planodorsalis Bridgeites supraconvexa Euconia etna Ceratopea ?laurentia Ceratopea pygmaea Orospira bigranosa Macluritella stantoni Teiichispira odenviltensis Teiichispira ?oceana Palliseria robusta Mitrospira longwelli Teiichispira kobayashi Teiichispira sylpha Monitorella auricula Maclurites magna "Eccyliopterus ornatus" Maclurites bigsbyi 7 6 7 n n n n n n ? n n n n n n n ? 1 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n 7 7 7 n n n n n n 7 n n n n n n n ? 1 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n 7 8 7 n n n n n n ? n n n n n n n ? n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n 7 9 7 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n 8 0 7 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n 8 1 7 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n 8 2 7 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n 8 3 7 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n II n n n n n n n n n n n n n n n n n n n n n 8 4 7 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n 8 5 7 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n 8 6 7 2 2 2 2 2 2 2 1 1 1 8 7 7 2 ? 2 2 2 2 2 1 2 2 2 1 1 1 3 3 1 2 1 1 1 1 8 8 7 n n n n n n ? ? n n n n n n n n n n n 7 n n n n n n n n n n n n n 2 n n n n n n 2 2 2 2 n 2 2 2 n n n 2 3 n n n n n n 8 9 7 7 7 7 7 ? ? 1 1 1 1 1 2 2 2 1 1 2 1 2 2 2 2 1 1 2 2 2 1 2 2 2 2 2 2 1 1 2 2 2 2 1 2 9 0 7 n n n n 7 n n n n n n n n n n n n n 7 7 7 ? 7 n n n n n 1 1 1 n n 1 n 1 1 1 1 n n 1 1 1 n 2 2 3 1 1 1 n n 1 1 1 1 n 1 9 1 7 n n n n 7 n n n n n n n n n n n i i n 7 7 7 ? 7 n n n n n 1 1 1 n n 1 n 1 2 1 1 n n 3 1 3 n 3 3 2 1 1 1 n n 1 1 1 1 n 1 9 2 7 n n n n ? n n n n n n n n n n n n n ? 7 7 ? 7 n n n n n 1 1 1 n n 1 n 1 1 1 1 n n 1 1 1 n 1 1 1 7 1 1 n n 1 1 1 1 n 1 9 3 9 n n n n 7 n n n n n n n n n n n n n ? 7 7 ? ? n n n n n 2 2 2 n n 2 n 2 2 2 2 n n 2 2 2 n 2 2 1 ? 2 2 2 2 2 2 2 2 n 2 9 4 7 3 3 3 5 3 5 3 5 4 4 4 4 4 4 5 3 1 2 4 5 5 5 5 5 5 3 3 3 3 3 4 3 3 4 3 6 4 6 4 3 4 3 5 5 6 5 5 4 4 4 4 3 3 3 3 3 3 3 4 9 5 7 1 3 3 2 1 3 3 2 2 3 3 2 7 3 ? ? 3 1 3 2 2 3 2 2 2 ? 2 2 2 2 2 2 2 3 2 2 3 2 1 2 2 2 3 3 2 3 3 3 2 2 2 3 2 3 9 6 7 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 1 2 2 1 2 1 2 1 2 1 2 1 2 2 1 2 1 2 1 2 1 3 1 2 1 2 1 2 1 2 2 2 2 2 2 2 2 2 2 2 2 2 1 1 2 2 2 2 2 2 ) 9 7 8 1 1 1 1 1 4 4 1 1 1 1 1 1 1 1 1 1 1 1 7 7 8 7 8 8 8 1 2 1 2 2 1 1 4 2 3 4 3 6 1 1 7 6 7 3 4 4 2 7 7 7 9 9 5 8 7 7 6 7 1 ? 0 ? 0 ? 7 2 NUMBER 88 105 APPENDIX 2.?Continued. Number Species AC. Mollusc AB. Helcionella subrugosa AA. Latouchella merino Z. Pelagiella subangulata Y. Costipelagiella zazvorkai X. Oelandia rugosa W. Coreospira rugosa V. Sinuella minuata U. Chippewaella patellitheca T. Strepsodiscus major S. Chalarostrepsis praecursor R. Eobucania mexicana Q. Strepsodiscus paucivoluta P. Modestospira poulseni 0. Peelerophon oehlerti N. Kiringella pyramidalis M. Cyrtolites sp. L. Macluritella? walcotti K. Euomphalopsis involuta J. "Maclurites" thomsoni I. Kobayashiella circe H. Scaevogyra swezeyi G. Mat here!la saratogensis F. Matherellina walcotti E. Hypseloconus elongatus D. Knightoconus antarcticus C. Sinuites sowerbyi B. Cloudia buttsi A. Owenella antiquata 1. Dirhachopea normalis 2. Dirhachopea subrotunda 3. Schizopea typica 4. Sinuopea sweeti 5. Taeniospira emminencis 6. Ceratopea canadensis 7. Gasconadia putilla 8. Jarlopsis conicus 9. Ophileta supraplana 10. Rhombella umbilicata 11. Prohelicotoma uniangulata 12. Sinuopea basiplanata 13. Taeniospira ?st. clairi 14. Bridgeites ?disjuncta 15. Bridgeites planodorsalis 16. Bridgeites supraconvexa 17. Euconia etna 18. Ceratopea? laurentia 19. Ceratopea pygmaea 20. Orospira bigranosa 21. Macluritella stantoni 22. Teiichispira odenvillensis 23. Teiichispira loceana 24. Palliseria robusta 25. Mitrospira longwelli 26. Teiichispira kobayashi 27. Teiichispira sylpha 28. Monitorella auricula 29. Maclurites magna 30. "Eccyliopterus ornatus" 31. Maclurites bigsbyi 1 1 1 0 0 0 1 2 3 ? ? 4 ? 2 4 ? 2 4 ? ? ? ? : ? 7 7 7 ? ? ? 7 7 ? ; ? 7 ? ? 7 7 ? ? ? ? ? 7 7 ? ? ? ? 7 ? 7 ? 7 7 ? 7 ? 7 j ? 7 ? ; ? j 1 ? ? 7 ? 7 ? ? ? 7 7 ? 4 4 ' 4 > 4 4 3 4 4 3 4 7 4 4 > 4 4 4 ? 4 4 4 4 4 4 3 4 4 4 4 4 4 4 >. 3 4 3 ' 3 3 4 4 3 3 > 4 4 3 > 2&3 ! 3 3 4 3 1 3 4 3 3 3 4 3 4 1 1 1 1 1 1 0 0 0 0 0 0 4 5 6 ' ? 7 ? 2 1 1 2 3 1 1 1 1 I 1 1 ? i i 2 1 1 7 i j 2 1 1 1 1 1 1 1 1 1 1 1 1 1 1 7 i i 1 1 1 2 1 1 1 1 1 ? i i ? i i ? 2 1 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 2 1 1 2 2 1 7 1 1 7 i j 1 3 2 1 1 2 7 8 9 ) ? i 1 2 1 2 2 2 1 2 1 2 1 2 1 1 0 ? ? 7 2 3 ? ? ? ? ? ? 7 ? ? ? 9 ? 7 ? 7 ? ? ] ? ? ? 1 ? ] ? i 7 7 7 ? ] ? i ? 7 7 ] 3 1 2 1 ? 2 1 ? ? ? ? 7 ? 2 ? ? 7 ? ? ? 7 7 ? ? ? ? ? ? 1 1 1 2 1 ? ? ? 7 7 ? 7 7 ? ? ? ? ? ? ? ? ? ? ? 7 7 7 7 7 ? ] ? ? ? ? ? ? ? ? 7 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 7 ? 7 7 ? ? 1 1 1 1 3 4 ? ? ? 7 ? 7 7 7 ? ? ? ? ? ? ? 7 ? ? ? ? ? : ? 2 ? ; ? : ? : ? i ? ? ? 7 7 7 7 ? ? ? ? ? ? 7 ? ? 7 ? 7 ? 7 ? ? 7 ? ? 7 7 ? ? ? ? 7 ? ? 1 1 1 1 5 6 ? ? 1 2 2 2 2 2 2 1 2 2 2 2 2 2 2 2 2 2 2 1 1 1 1 1 2 2 1 1 1 2 2 2 1 1 2 2 2 2 2 2 1 1 2 2 2 2 2 2 2 2 2 2 2 2 I 2 2 2 2 2 2 1 1 7 7 2 1 2 2 1 3 n 5 5 5 5 5 5 5 5 5 1 1 1 n 'i n \ n \ n "J. n 'i 5 1 5 1 n 1 n n 3 1 3 1 3 1 n 1 n 2 1 4 2 3 3 n n 3 3 3 3 3 3 2 3 1 3 3 1 3 1 1 1 1 3 9 ? ? n n n n n n n n n n n n n n n n n n 2 1 1 1 1 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n 1 2 0 7 2 3 2 3 3 3 1 3 3 2 2 1 2 1 2 1 ? 6 6 6 6 6 4 2 7 3 3 3 3 2 3 7 5 1 1 3 2 3 2 2 7 7 2 2 2 3 3 3 2 4 3 2 3 1 3 4 2 3 3 3 3 2 4 4 2 5 4 5 3 3 6 4 4 5 5 5 1 2 2 ? 2 2 2 2 2 2 2 2 2 2 2 3 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 3 3 2 2 3 3 3 3 3 2 1 2 3 ? 1 1 4 4 1 3 2 1 3 3 4 2 2 4 1 2 0 1 5 5 5 5 5 3 3 3 5 5 4 4 3 5 4 4 5 4 3 4 3 4 5 3 3 3 5 4 4 4 2 4 3 2 2 3 2 4 3 4 3 1 2 4 7 2 2 2 2 2 2 1 2 5 1 1 1 2 2 1 1 1 1 1 1 1 1 I 1 1 1 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2rt 2 2 2 2 2 2 2 2 2 2 2 2 2 2 106 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY APPENDIX 2.?Continued. Number Species AC. AB. AA. Z. Y. X. W. V. u. T. S. R. Q- P. 0 . N. M. L. K. J. I. H. G. F. E. D. C. B. A. 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12. 13. 14. 15. 16. 17. 18. 19. 20. 21. 22. 23. 24. 25. 26. 27. 28. 29. 30. 31. Mollusc Helcionella subrugosa Latouchella merino Pelagiella subangulata Costipelagiella zazvorkai Oelandia rugosa Coreospira rugosa Sinuella minuata Chippewaella patellitheca Strepsodiscus major Chalarostrepsis praecursor Eobucania mexicana Strepsodiscus paucivoluta Modestospira poulseni Peelerophon oehlerti Kiringella pyramidalis Cyrtolites sp. Macluritella? walcotti Euomphalopsis involuta "Maclurites" thomsoni Kobayashiella circe Scaevogyra swezeyi Ma there I la saratogensis Matherellina walcotti Hypseloconus elongatus Knightoconus antarcticus Sinuites sowerbyi Cloudia butts i Owenella antiquata Dirhachopea normalis Dirhachopea subrotunda Schizopea typica Sinuopea sweeti Taeniospira emminencis Ceratopea canadensis Gasconadia putilla Jarlopsis conicus Ophileta supraplana Rhombella umbilicata Prohelicotoma uniangulata Sinuopea basiplanata Taeniospira ?st. clairi Bridgeites ?disjuncta Bridgeites planodorsalis Bridgeites supraconvexa Euconia etna Ceratopea ?laurentia Ceratopea pygmaea Orospira bigranosa Macluritella stantoni Teiichispira odenvillensis Teiichispira ?oceana Palliseria robusta Mitrospira longwelli Teiichispira kobayashi Teiichispira sylpha Monitorella auricula Maclurites magna "Eccyliopterus ornatus" Maclurites bigsbyi 1 1 1 1 1 2 2 2 2 3 6 7 8 9 0 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 1 0 1 0 0 3 3 3 3 3 4 4 4 6 6 4 6 5 4 5 4 6 7 : 3 3 3 5 4 ' 4 : 4 2 2 3 1 1 2 2 2 2 2 2 7 7 7 2 2 i : l 2 ? 1 7 2 7 ? 2 ? 1 ? 1 2 : 2 1 ? 1 1 1 2 2 1 1 1 1 2 2 2 1 1 2 ? 2 2 2 2 1 > 1 ? 2 2 ? } 2 I 2 2 ; 2 ? ? 3 3 ? 3 2 3 2 3 : n n ! 3 n n n n n n n n n n n n n ! 2 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n I 3 n n n n n n n n n n 5 3 1 3 1 9 n n 1 n n n n n n n n n n n n n 2 n n n n n n n n n n n n n n n n n n n n n II n n n n n n n n 1 n n n n n n n n n n 3 1 1 1 3 3 3 2 3 4 9 9 9 n n i : n n n n n n n n n n n n n i : n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n i : n n n n n n n n n n 1 n n ! 3 n n n n n n n n n n n n n . 2 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n > 3 n n n n n n n n n n n 1 3 5 9 n n 1 n n n n n n n n n n n n n 1 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n 1 n n n n n n n n n n n 1 3 6 9 n n 1 n n n n n n n n n n n n n 1 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n 1 n n n n n n n n n n n 1 3 : 1 1 1 3 4 4 7 8 9 0 1 7 n n 1 n n n n n n n n n n n n n 1 1 n n n n n n n n 1 n n n n n n n n n n n n n n n n n n n n n n 1 n n n n n n n n n n n 7 ? 7 n n n n n n n n 1 1 1 1 1 n n n n 1 1 n n n n 0 0 0 0 0 1 1 2 2 2 ? 2 2 2 2 2 1 2 2 0 2 2 2 3 3 2 1 1 2 2 2 1 2 2 1 3 2 2 2 2 2 3 2 2 2 2 2 2 2 2 2 3 4 1 2 1 2 1 2 1 4 1 2 1 4 1 4 2 7 1 1 1 1 7 7 7 ? 7 7 7 ? ? ? 7 7 7 7 1 7 7 ? ? 7 7 7 7 ? 7 7 7 7 ? ? 7 ? 7 ? 7 ? ? 7 ? ? ? ? 7 ? 7 ? ? ? ? ? ? 7 ? ? ? 1 4 3 ? n n n 1 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n NUMBER 88 107 APPENDIX 2.?Continued. Number Species 32. 33. 34. 35. 36. 37. 38. 39. 40. 41. 42. 43. 44. 45. 46. 47. 48. 49. 50. 51. 52. 53. 54. 55. 56. 57. 58. 59. 60. 61. 62. 63. 64. 65. 66. 67. 68. 69. 70. 71. 72. 73. 74. 75. 76. 77. 78. 79. 80. 81. 82. 83. 84. 85. 86. 87. 88. 89. 90. 91. Maclurina logani Maclurina manitobensis Maclurites sedgewicki Maclurites expansa Ophileta complanata Lecanospira compacta Lecanospira nereine Barnesella ?lecanospiroides Malayaspira hintzei Malayaspira rugosa Barnesella measuresae Lytospira angelini Lytospira yochelsoni Maclurina ?annulata Rossospira harrisae Ecculiomphalus bucklandi Lytospira gerrula Lytospira ?norvegica Ophiletina cf. 0. sublaxa Lytospira subrotunda Pararaphistoma lemoni Climacoraphistoma vaginati Lesueurilla bipatellare Lesueurilla marginalis Lesueurilla prima Palaeomphalus giganteus Climacoraphistoma damesi Eccyliopterus alatus Eccyliopterus ?princeps Eccyliopterus regularis Lesueurilla infundibula Eccyliopterus louderbacki Lesueurilla declivis Pararaphistoma qualteriata Pararaphistoma schmidti Helicotoma gubanovi Scalites katoi Helicotoma medfraensis Lesueurilla scotica Pachystrophia devexa Raphistoma striata Raphistomina lapicida Scalites angulatus Holopea insignis Eccyliopterus beloitensis Holopea rotunda Pachystrophia contigua Pachystrophia spiralis Raphistomina aperta Raphistomina fissurata Eccyliopterus owenanus Holopea ampla Holopea pyrene Holopea symmetrica Raphistoma per acuta Raphistomina rugata Raphistoma tellerensis Sinutropis ?esthetica Pachystrophia gotlandica Lytospira triquestra 1 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 1 2 1 2 2 2 2 2 1 1 1 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 3 2 2 2 2 3 3 3 3 3 2 2 2 1 1 1 1 2 1 2 2 2 1 1 2 1 1 2 2 2 3 2 2 2 2 7 1 7 2 2 2 2 2 7 7 7 2 2 2 2 2 2 3 2 2 2 2 6 6 6 4 4 5 4 4 4 5 4 2 3 3 1 4 6 5 6 5 5 5 5 4 3 4 5 5 5 5 6 4 5 5 4 3 4 2 4 7 4 9 3 3 2 2 3 7 7 ? 4 2 3 3 3 2 4 2 2 2 2 6 6 6 4 6 5 4 4 4 6 6 6 5 5 1 4 6 4 5 3 4 5 4 4 3 4 3 4 5 4 4 4 5 5 3 3 4 2 4 7 3 7 3 3 2 2 3 7 7 ? 4 2 3 3 3 2 5 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 1 2 2 2 1 1 1 1 2 1 2 1 2 1 1 2 1 1 2 2 2 1 2 2 2 2 9 1 7 2 2 2 2 2 7 7 7 2 2 2 2 2 2 6 1 1 1 1 3 3 2 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 7 3 7 3 3 3 3 3 9 9 ? 3 3 3 3 3 2 7 1 1 1 1 3 3 2 3 3 3 3 3 3 3 3 3 3 2 3 3 3 2 2 2 2 3 2 3 2 3 2 3 3 2 2 3 3 3 2 3 3 3 3 9 3 7 3 3 3 3 3 7 7 7 3 3 3 3 3 2 8 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 1 1 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 9 2 7 2 1 2 2 2 9 9 2 2 2 2 1 1 1 9 2 2 2 2 4 4 4 4 4 4 3 3 3 4 4 4 2 2 2 1 4 4 4 4 4 4 4 4 3 4 4 4 4 4 4 1 4 4 4 1 1 1 4 7 3 9 1 1 1 1 4 7 7 7 1 1 4 1 1 1 1 1 1 0 1 2 2 1 1 2 1 1 2 1 1 2 1 1 4 1 1 4 1 1 4 1 1 4 1 1 4 1 1 4 1 1 3 1 1 3 1 1 3 1 1 4 1 1 4 1 1 4 1 1 3 1 1 3 1 1 2 1 1 1 1 1 4 1 1 4 1 1 4 1 1 4 1 1 4 1 1 4 1 1 4 1 1 4 1 1 3 1 1 4 1 1 4 1 1 4 1 1 4 1 1 4 1 1 4 1 1 1 1 1 4 1&2 1 4 1 1 4 1 1 1 1 1 1 2 1 I 1 1 4 1 1 7 7 7 3 1 1 7 7 7 1 1 1 3 1 1 1 1 1 1 1 1 4 1 1 7 7 J 9 9 9 ? 1 1 1 2 1 1 1 1 4 2 1 3 1 1 3 1 1 3 1 1 1 1 3 4 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 1 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 Tt 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 1 1 n 1 n 1 n 1 n 1 n 1 n 1 5 4 2 2 3 3 4 4 4 4 4 4 4 4 4 4 4 3 3 2 3 4 4 3 4 4 4 3 4 4 4 4 4 4 4 4 4 2 4 3 3 4 4 4 3 4 3 3 3 4 3 4 3 3 3 4 5 3 3 3 2 1 6 2 2 : 2 2 1 2 1 2 2 2 2 ? 2 2 1 7 2 1 1 1 2 1 2 1 1 1 2 1 2 1 2 1 2 1 2 1 1 1 2 1 2 1 2 1 2 1 2 1 1 1 7 2 1 2 1 2 2 1 7 8 2 1 1 1 9 2 7 2 2 2 2 2 2 2 1 1 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 9 2 1 9 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 1 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 1 2 2 2 1 2 1 1 1 2 2 2 1 1 1 2 2 2 1 1 1 2 0 1 1 1 1 3 2 2 2 2 2 2 1 2 2 2 2 1 2 1 n 1 2 2 2 2 3 2 1 2 2 2 2 2 2 1 2 2 1 2 n 2 2 2 n 1 n n n 2 2 1 n n n 2 2 2 n n n 2 n 2 n n 1 n n n 1 1 1 n n n 1 1 1 n n n 2 2 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n 1 n n n n 1 n n n n n n n n n n n n n n n n n n n 2 3 n n n n n n n n n n II n n n n n n n n n n n n n n n n n n n n n n n n 2 n n n n 1 n n n n n n n n n n n n n n n n n n n 2 4 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n 1 n n n n 1 n n n n n n n n n n n n n n n n II n " 2 5 n n n n n n n n n n n n n n n n n n n n n " n n n n n n n n n n n n n nrt n n n n n n n n n n n n n n n n n n n n n n n n 108 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY APPENDIX 2.?Continued. Number Species 32. 33. 34. 35. 36. 37. 38. 39. 40. 41. 42. 43. 44. 45. 46. 47. 48. 49. 50. 51. 52. 53. 54. 55. 56. 57. 58. 59. 60. 61. 62. 63. 64. 65. 66. 67. 68. 69. 70. 71. 72. 73. 74. 75. 76. 77. 78. 79. 80. 81. 82. 83. 84. 85. 86. 87. 88. 89. 90. 91. Maclurina logani Maclurina manitobensis Maclurites sedgewicki Maclurites expansa Ophileta complanata Lecanospira compacta Lecanospira nereine Barnesella ?lecanospiroides Malayaspira hintzei Malayaspira rugosa Barnesella measuresae Lytospira angelini Lytospira yochelsoni Maclurina ?annulata Rossospira harrisae Ecculiomphalus bucklandi Lytospira gerrula Lytospira ?norvegica Ophiletina cf. O. sublaxa Lytospira subrotunda Pararaphistoma lemoni Climacoraphistoma vaginati Lesueurilla bipatellare Lesueurilla marginalis Lesueurilla prima Palaeomphalus giganteus Climacoraphistoma damesi Eccyliopterus alatus Eccyliopterus ?princeps Eccyliopterus regularis Lesueurilla infundibula Eccyliopterus louderbacki Lesueurilla declivis Pararaphistoma qualteriata Pararaphistoma schmidti Helicotoma gubanovi Scalites katoi Helicotoma medfraensis Lesueurilla scotica Pachystrophia devexa Raphistoma striata Raphistomina lapicida Scalites angulatus Holopea insignis Eccyliopterus beloitensis Holopea rotunda Pachystrophia contigua Pachystrophia spiralis Raphistomina aperta Raphistomina fissurata Eccyliopterus owenanus Holopea ampla Holopea pyrene Holopea symmetrica Raphistoma peracuta Raphistomina rugata Raphistoma tellerensis Sinutropis ?esthetica Pachystrophia gotlandica Lytospira triquestra 2 6 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n 1 n n n n 1 n n n n n n n n n n n n n n n n n n n 2 7 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 i i 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 n 2 2 2 n 2 n n n 2 2 2 n n n 2 2 2 n n n 2 8 1 1 1 1 3 2 2 2 2 2 2 2 2 2 2 2 2 3 2 n 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 4 3 3 3 n 4 3 4 n 3 n n n 3 3 3 n n n 3 3 3 n n n 2 9 2 1 1 3 4 4 4 4 4 3 4 3 4 3 4 3 4 4 2 n 4 4 5 5 4 4 3 6 5 5 4 4 4 4 4 2 5 2 3 n 5 5 5 n 4 n n n 5 3 6 n n n 5 5 5 n n n 3 0 2 n n 1 n n n 1 1 1 n n n 1 1 1 n n n 3 n n n 1 n n n 1 1 1 n n 1 1 1 1 n n n 3 2 n n n 1 n n n 1 1 1 n n n 1 1 1 n n n 3 3 2 2 2 n 2 n 2 n n n 1 1 1 n n n 1 2 1 n n n 3 4 2 2 2 2 1 2 1 1 1 2 2 1 2 2 2 1 1 2 1 2 1 7 1 2 1 1 1 ? ? 1 1 1 1 1 1 1 1 1 1 3 5 n n n n n n n n n n n n n n n n n n n n n 2 2 2 2 n 2 n n n 2 2 n 2 2 n n n 2 n 2 n n n n n n n n n n n n n n n n n n n 3 6 n n n n n n n n n n n n n n n n n n n n n 1 1 1 1 n 1 n n n I 1 n 1 1 n n n 1 n 1 n n n 1 n n n n n n n n " n n n u n n 3 7 n n n 1 n n n 1 1 1 n n n 1 1 1 n n n 3 8 4 4 4 4 1 4 4 4 4 4 4 4 1 1 3 3 3 3 3 3 3 3 3 3 3 3 3 3 1 3 1 3 1 1 1 n n n n 1 1 1 1 3 1 n n 1 1 1 1 1 1 3 9 2 2 2 2 2 1 1 1 3 3 1 n 1 3 3 2 1 1 1 2 2 1 1 1 1 2 3 2 2 2 1 1 2 1 1 1 1 2 1 2 1 2 n n 1 n 2 2 2 2 2 2 n n 2 2 1 2 2 2 4 0 2 n n n n n n 2 n n n 4 1 n n n n n n n n n n n i i n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n 4 2 1 1 1 1 2 2 2 2 2 2 2 1 2 2 2 2 1 1 1 n 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 n 2 2 2 n 2 n n n 2 1 2 n n n 1 1 2 n n n 4 3 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 n 2 3 3 3 3 2 3 2 3 2 3 3 2 3 3 2 2 2 3 n 2 2 2 n 3 n n n 2 2 2 n n n 2 2 2 n n n 4 4 n n n n n 1 n n n 1 1 1 n n 1 1 1 1 n n n 4 5 1 1 1 1 2 2 ? 1 1 1 1 2 1 1 1 ? 2 2 2 1 2 2 I 2 2 2 1 2 1 2 2 2 2 2 1 1 2 2 1 1 1 2 1 1 2 1 1 1 2 2 2 1 1 1 2 1 1 1 1 1 4 4 4 6 n n n n 1 1 n n n n n 1 n n n n 1 1 1 n 1 1 n 1 1 1 1 1 1 n 1 1 n 1 1 1 1 1 1 1 n n 1 1 n n n 1 n n 1 n n n 1 1 1 n n n 1 n n n n n 7 8 9 1 9 I 9 9 5 7 7 7 8 8 7 7 7 2 6 6 7 6 7 C 5 7 9 9 7 5 8 8 7 5 6 7 7 8 7 B B 7 8 7 B 6 A 7 6 B 9 C 5 7 8 9 7 D [ 8 I 8 I B I 7 1 C 1 7 4 9 2 2 3 3 3 2 2 3 1 ? 1 3 1 2 3 3 2 2 2 3 3 2 5 0 n n n n n n n n n n n n n n n n n n n n n 2 1 1 1 1 2 n n n 1 n 1 2 2 n n n 1 n 1 n n n n n n n n n n n n n n n n n n n NUMBER 88 109 APPENDIX 2.?Continued. Number Species 32. 33. 34. 35. 36. 37. 38. 39. 40. 41. 42. 43. 44. 45. 46. 47. 48. 49. 50. 51. 52. 53. 54. 55. 56. 57. 58. 59. 60. 61. 62. 63. 64. 65. 66. 67. 68. 69. 70. 71. 72. 73. 74. 75. 76. 77. 78. 79. 80. 81. 82. 83. 84. 85. 86. 87. 88. 89. 90. 91. Maclurina logani Maclurina manitobensis Maclurites sedgewicki Maclurites expansa Ophileta complanata Lecanospira compacta Lecanospira nereine Barnesella ?lecanospiroides Malayaspira hintzei Malayaspira rugosa Barnesella measuresae Lytospira angelini Lvtospira yochelson i Maclurina ?annulata Rossospira harrisae Ecculiomphalus bucklandi Lytospira gerrula Lytospira ?norvegica Ophiletina cf. 0. sublaxa Lytospira subrotunda Pararaphistoma lemoni Climacoraphistoma vaginati Lesueurilla bipatellare Lesueurilla marginalis Lesueurilla prima Palaeomphalus giganteus Climacoraphistoma damesi Eccyliopterus alatus Eccyliopterus ?princeps Eccyliopterus regularis Lesueurilla infundibula Eccyliopterus louderbacki Lesueurilla declivis Pararaphistoma qualteriata Pararaphistoma schmidti Helicotoma gubanovi Scalites katoi Helicotoma medfraensis Lesueurilla scotica Pachystrophia devexa Raphistoma striata Raphistomina lapicida Scalites angulatus Holopea insignis Eccyliopterus beloitensis Holopea rotunda Pachystrophia contigua Pachystrophia spiralis Raphistomina aperta Rap his torn ina fis sur at a Eccyliopterus owenanus Holopea ampla Holopea pyrene Holopea symmetrica Raphistoma peracuta Raphistomina rugata Raphistoma tellerensis Sinutropis ?esthetica Pachystrophia gotlandica Lytospira triquestra 5 1 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 2 3 1 3 3 3 3 1 3 3 3 3 3 3 3 2 2 2 2 3 3 1 2 1 2 1 3 2 1 3 1 1 3 1 1 2 2 2 3 3 3 3 5 2 3 3 3 4 3 4 4 4 2 2 3 2 3 2 2 3 3 3 3 2 2 3 2 3 3 2&3 3 3 4 2 4 2 3 3 3 2 3 2 4 2 2 2 3 2 2 1 2 2 2 2 3 2 2 1 3 3 3 2 2 2 5 3 3 3 3 3 2 2 2 2 3 3 2 1 2 3 3 2 2 9 2 2 2 3 2 2 2 1 3 2 9 2 1 1 2 3 3 2 2 2 1 2 2 3 2 2 1 2 2 2 2 2 2 2 2 2 2 1 2 2 2 2 5 4 3 3 3 3 2 2 2 2 2 1 2 2 2 2 1 1 1 1 2 1 2 2 1 1 3 3 1 1 2 3 2 3 3 2 3 2 3 2 2 1 3 3 3 3 2 3 2 2 2 5 5 3 4 4 5 5 5 5 5 5 5 5 5 5 5 5 5 5 6 5 5 5 5 6 6 6 6 6 6 6 6 5 6 6 6 6 2 2 4 6 4 2 4 2 3 6 2 4 4 4 4 6 3 3 0 4 4 3 5 5 4 5 6 7 7 7 7 5 5 5 3 4 4 3 5 4 4 4 4 4 4 3 5 5 5 6 6 6 6 4 6 5 6 3 6 6 5 4 5 5 4 5 4 5 4 5 4 6 4 4 5 4 4 6 4 4 4 5 4 4 5 5 4 5 7 1 1 1 2 2 2 2 2 3 2 2 2 2 3 3 3 2 2 2 1 2 2 2 2 2 2 2 1 2 2 2 2 2 2 3 1 1 2 2 2 1 2 1 3 2 3 3 2 2 2 1 3 3 3 2 2 2 1 1 1 5 8 3 3 4 2 2 1 1 1 3 3 1 2 2 2 3 3 1 1 4 3 1 1 1 1 1 2 2 1 1 1 2 1 1 1 1 4 3 3 1 2 3 1 4 3 1 4 3 4 1 1 1 3 3 4 1 1 2 3 3 4 5 9 2 2 2 2 2 1 1 I 4 3 1 2 2 2 4 4 1 1 4 3 1 1 1 1 1 2 2 1 1 1 2 1 1 1 2 1 1 3 1 2 1 1 1 6 1 5 4 4 1 1 1 5 6 5 1 1 2 2 2 3 6 0 1 1 1 1 2 2 1 2 2 2 2 2 2 2 2 2 2 2 1 2 2 2 6 1 n n n n 2 n 3 3 2 3 n n n n n n n n 1 n n n n n n n n 3 n n n n n n n n n n n n n n n n n n n n n n 6 t 2 3 n 1 n 1 n 1 n 1 3 1 3 1 n 1 2 1 2 2 2 2 2 1 3 1 3 1 2 2 1 2 1 2 3 1 n 1 n 1 3 1 2 1 n ] n 1 n 1 n 1 n 1 n 1 n i n n n n n n n n n 2 n n n n n n n n n n n n n n n n n n n n n n 6 6 4 5 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 2 2 1 n 1 n 1 n 1 n 2 2 2 2 1 n 1 n 1 n 2 1 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 2 1 2 1 1 n 1 n 1 n 1 n 1 n 1 n 2 1 2 1 1 n 1 n 1 n 1 n 1 2 2 1 2 1 1 1 n I 1 n 1 1 n 1 1 n 6 6 n n n n n n n n 4 n n n n 2 4 n n n 4 n n n n n n n n n n n n n n n n n n n n n 1&3 1 n n n n n n 1 1 n n n n 4 1 n n n n 6 7 n n n n n n n n 1 n n n n 2 1 n n n 1 n n n n n n n n n n n n n n n n n n n n n 3 2 n n n n n n 2 2 n n n n 2 1 n n n n 6 t 8 5 n n i n 1 n 1 n ] n 1 n 1 n 1 1 1 n 1 n 1 n 1 n ? 1 1 1 1 n 1 n ? n ? 1 1 n ? n 1 n 1 n ] n 1 n 1 n 1 n 1 n 1 n ' n 1 n n n n n 1 n n n n i i 1 1 n n n n ; n n 1 1 n n n n 1 1 n n n n 7 1 0 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n n n 1 n 1 n n n n n n n I 1 n n n n 1 n i n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 7 1 2 3 2 1 2 I 2 1 2 1 3 1 3 1 3 1 3 1 3 1 3 1 3 1 2 1 3 1 3 1 3 1 3 1 2 1 2 1 3 1 2 1 3 1 3 1 3 I 2 1 2 1 3 1 2 1 2 1 3 1 2 1 2 1 2 1 2 1 3 1 3 1 2 1 3 1 2 1 3 1 2 1 2 1 3 1 3 2 2 1 2 1 3 1 3 1 2 1 2 1 2 1 1 1 3 1 3 1 2 1 2 1 1 1 2 7 7 4 5 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 2 n 1 n 1 n 1 n 1 n 2 n 2 n 2 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 ti 1 n J n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 1 n 1 1 n 1 I n 1 1 n 1 1 n 1 1 n 1 1 n 1 110 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY Number Species 32. 33. 34. 35. 36. 37. 38. 39. 40. 41. 42. 43. 44. 45. 46. 47. 48. 49. 50. 51. 52. 53. 54. 55. 56. 57. 58. 59. 60. 61. 62. 63. 64. 65. 66. 67. 68. 69. 70. 71. 72. 73. 74. 75. 76. 77. 78. 79. 80. 81. 82. 83. 84. 85. 86. 87. 88. 89. 90. 91. Maclurina logani Maclurina manitobensis Maclurites sedgewicki Maclurites expansa Ophileta complanata Lecanospira compacta Lecanospira nereine Barnesella ?lecanospiroides Malayaspira hintzei Malayaspira rugosa Barnesella measuresae Lytospira angelini Lytospira yochelsoni Maclurina ?annulata Rossospira harrisae Ecculiomphalus bucklandi Lytospira gerrula Lytospira ?norvegica Ophiletina cf. O. sublaxa Lytospira subrotunda Pararaphistoma lemoni Climacoraphistoma vaginati Lesueurilla bipatellare Lesueurilla marginalis Lesueurilla prima Palaeomphalus giganteus Climacoraphistoma damesi Eccyliopterus alatus Eccyliopterus ?princeps Eccyliopterus regularis Lesueurilla infundibula Eccyliopterus louderbacki Lesueurilla declivis Pararaphistoma qualteriata Pararaphistoma schmidti Helicotoma gubanovi Scalites katoi Helicotoma medfraensis Lesueurilla scotica Pachystrophia devexa Raphistoma striata Raphistomina lapicida Scalites angulatus Holopea insignis Eccyliopterus beloitensis Holopea rotunda Pachystrophia contigua Pachystrophia spiralis Raphistomina aperta Raphistomina fissurata Eccyliopterus owenanus Holopea ampla Holopea pyrene Holopea symmetrica Raphistoma peracuta Raphistomina rugata Raphistoma tellerensis Sinutropis ?esthetica Pachystrophia gotlandica Lytospira triquestra 1 6 n n n n n n n n 2 n n n n 2 2 2 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n 7 7 n n n n n n n n 2 n n n n 2 2 2 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n 7 8 n n n n n n n n 1 n n n n 1 1 1 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n 7 9 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n APPENDIX 2.? 8 0 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n 8 1 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n ?? n i i n n n n n 8 2 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n 8 3 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n -Continued. 8 4 n n n n n n n n n n n n n n n n n n n n n .. n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n 8 8 5 6 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 2 1 2 n 2 1 2 n ? n ? 1 1 1 ? 1 1 n ? n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n I n 1 n 1 n 1 n 1 n 1 n 1 n 1 ii 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 r> 1 n 1 n 1 n 1 8 7 9 2 2 2 3 3 2 2 2 2 3 1 1 1 8 8 n n n n n n 9 n n n n n n n n n n n n n n n n n n 2 n 2 n n n n n n n 2 2 n n n 2 n 2 n n n n n n n 2 n n n 2 3 3 n n n 8 9 2 2 2 2 2 2 7 2 1 1 2 1 2 1 1 1 1 1 2 1 2 2 2 2 2 2 2 2 2 2 1 1 2 2 2 2 2 2 2 1 2 2 7 2 1 1 1 1 2 2 2 1 2 1 2 2 2 1 I 1 9 0 7 1 n n 5 n 5 n n n n n 5 n 2 2 3 3 2 2 2 2 2 2 n n 2 2 2 2 2 1 2 n 2 1 7 1 n n n n 2 1 2 n 1 n 2 1 2 n n n 9 2 2 7 2 n n 3 n 3 n n n n n 3 n 1 3 3 3 3 3 1 3 2 3 n n 3 2 1 2 3 1 1 n 2 1 7 1 n n n n 2 1 3 n 1 n 2 1 3 n n n 9 2 1 1 1 1 1 1 7 1 n n 1 n 1 n n n n n 3 n 1 1 1 1 1 2 1 1 1 1 n n 2 1 1 1 2 1 1 n 2 2 7 1 n n n n 2 2 1 n 1 n 2 1 1 n n n 9 3 2 2 2 2 2 2 7 2 n n 3 n 3 n n n n n n n 2 2 2 2 2 2 2 2 2 2 n n 2 2 2 2 2 2 2 n 2 2 7 2 n n n n 2 2 2 n 2 n 2 2 2 n n n 9 4 4 4 4 3 4 4 4 3 4 4 3 4 4 4 4 5 4 5 5 4 5 4 3 3 4 5 3 4 5 4 4 4 4 4 5 1 1 4 4 3 1 5 1 1 4 2 3 3 3 5 4 2 1 2 3 5 1 3 3 3 9 5 3 3 2 3 3 3 3 3 2 2 3 3 3 1&2 2 2 3 2 2 2 3 2 2 2 2 2 3 3 2 3 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 3 2 2 2 2 2 2 2 2 2 1 9 9 9 9 0 6 7 8 9 0 2 1 2 1 2 1 2 1 2 1 2 1 ? 1 2 1 2 1 2 : 2 1 2 1 2 1 2 2 2 : 2 : 2 1 3 1 2 1 2 1 1 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 3 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 7 1 7 1 7 1 7 1 4 1 7 1 7 1 7 1 6 1 6 1 7 1 8 1 7 I 6 1 7 1 7 1 7 1 4 1 6 1 4 1 4 1 4 1 4 1 4 1 5 1 4 1 4 1 4 1 6 1 5 1 6 1 5 1 5 1 3 1 3 1 2 1 2 1 3 1 4 3 2 3 2 3 5 2 2 2 3 1 2 1 4 1 2 1 3 1 2 1 2 1 2 1 2 1 5 1 2 1 7 2 2 NUMBER 88 111 APPENDIX 2.?Continued. Number Species 32. Maclurina logani 33. Maclurina manitobensis 34. Maclurites sedgewicki 35. Maclurites expansa 36. Ophileta complanata 37. Lecanospira compacta 38. Lecanospira nereine 39. Barnesella ?lecanospiroides 40. Malayaspira hintzei 41. Malayaspira rugosa 42. Barnesella measuresae 43. Lytospira angelini 44. Lytospira yochelsoni 45. Maclurina ?annulata 46. Rossospira harrisae 47. Ecculiomphalus bucklandi 48. Lytospira gerrula 49. Lytospira ?norvegica 50. Ophiletina cf. 0 . sublaxa 51. Lytospira subrotunda 52. Pararaphistoma lemoni 53. Climacoraphistoma vaginati 54. Lesueurilla bipatellare 55. Lesueurilla marginalis 56. Lesueurilla prima 57. Palaeomphalus giganteus 58. Climacoraphistoma damesi 59. Eccyliopterus alatus 60. Eccyliopterus ?princeps 61. Eccyliopterus regularis 62. Lesueurilla infundibula 63. Eccyliopterus louderbacki 64. Lesueurilla declivis 65. Pararaphistoma qualteriata 66. Pararaphistoma schmidti 67. Helicotoma gubanovi 68. Scalites katoi 69. Helicotoma medfraensis 70. Lesueurilla scotica 71. Pachystrophia devexa 72. Raphistoma striata 73. Raphistomina lapicida 74. Scalites angulatus 75. Holopea ins ignis 76. Eccyliopterus beloitensis 77. Holopea rotunda 78. Pachystrophia contigua 79. Pachystrophia spiralis 80. Raphistomina aperta 81. Raphistomina fissurata 82. Eccyliopterus owenanus 83. Holopea amp la 84. Holopea pyrene 85. Holopea symmetrica 86. Raphistoma per acuta 87. Raphistomina rugata 88. Raphistoma tellerensis 89. Sinutropis ?esthetica 90. Pachystrophia gotlandica 91. Lytospira triquestra 1 0 1 9 7 7 7 ? 7 ? ? 7 9 7 ? ? ? ? ? ? 1 ? 9 7 9 9 9 9 7 9 9 1 ? 7 ? ? ? ? ? ? 7 9 7 ? ? 7 ? 7 7 7 ? 9 7 7 7 7 ? 7 7 7 7 9 7 1 0 2 2 1 2 2 2 2 2 2 2 1 1 1 1 0 0 0 c 3 4 5 6 4 1 1 1 4 1 1 1 ? ? 1 1 3 1 1 1 3 1 1 1 4 1 1 1 ? ? 1 1 4 1 1 1 4 1 1 1 3 1 1 1 4 1 1 1 4 1 1 1 4 1 1 1 4 1 1 1 4 1 1 1 4 1 1 1 4 1 1 1 4 1 1 1 4 1 1 1 4 1 1 3 1 1 3 1 1 3 1 1 ? ? i 3 1 1 3 1 1 4 1 1 4 1 1 4 1 1 4 1 1 3 1 1 3 1 1 3 1 1 3 1 1 3 1 1 3 1 1 4 1 1 3 1 1 3 1 1 4 1 1 1 1 1 2 1 1 4 1 1 4 1 1 3 1 1 3 1 1 4 1 1 4 1 1 3 1 1 3 1 1 4 1 1 4 1 1 4 1 1 3 1 1 2 1 1 1&2 1 1 3 1 1 4 1 1 4 1 1 4 1 1 1 1 1 0 0 0 7 8 9 1 1 2 1 2 2 1 2 2 1 1 2 1 1 1 2 1 1 1 2 1 1 1 2 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 2 1 1 1 2 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 1 1 1 2 1 1 1 1 1 1 1 1 0 1 ? 1 ? 1 ? i 7 i ? ] 7 i 7 ] 7 i ? i ? 1 7 i 7 i 7 i 7 j 7 i 7 ] ? 1 ? 1 ? 1 ? j ? 1 ? 1 ? i ? 1 ? j ? 1 7 ? 1 ? ] 7 j ? 1 ? ] ? ? 1 ? 2 1 1 ? ? ? 2 7 2 2 7 3 2 3 9 ? ? 2 2 3 ? ? 1 9 2 7 1 1 1 1 1 1 2 3 4 ? 1 ? ? J 7 ? 1 ? ? 1 7 ? J 7 ? 1 ? ? 1 7 ? 1 ? ? 1 ? ? 1 7 ? j ? ? 1 "? ? 1 ? ? 1 7 7 1 7 7 1 7 ? J 7 7 1 7 ? 1 ? 7 J 7 ? 1 ? ? J 7 ? 1 7 ? i 7 ? 1 7 ? 1 ? 7 J 7 ? 1 1 ? 1 "3 ? 1 ? i ? i 7 i 1 ? 1 1 ? 1 [ ? 1 [ ? i 1 ? 1 1 ? 1 1 ? 1 1 ? 1 1 1 1 1 5 6 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 1 1 2 1 2 1 2 1 1 1 1 1 1 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 1 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 ? ' 1 2 > 1 ? > 1 2 ' 1 2 > 1 2 > 1 2 > 1 2 ? 1 2 ? 1 2 ? 1 2 ? 1 2 ? 1 2 ? 1 2 ? 1 2 ? 1 2 ? 1 2 ? 1 2 ? 1 2 ? 1 2 1 1 1 1 7 J 1 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 3 1 2 1 3 1 n 2 1 1 2 1 3 1 n 2 n 2 n 2 4 1 3 1 2 1 3 1 3 1 3 1 3 1 4 1 3 1 n 2 3 1 3 1 3 1 3 1 2 2 2 1 3 3 3 n 3 n 3 3 1 3 2 3 3 3 3 3 1 3 2 1 2 2 1 1 1 9 n n n n n n n n n n n n 2 n n n 2 3 2 n n n n n n n n n 3 n n n n n n n n n n n n n ? n n n i n 1 n 1 n | n [ n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 2 0 1 1 1 1 2 2 ? 2 2 2 2 2 2 2 2 2 2 2 2 1 3 3 3 3 3 3 3 3 3 3 2 2 3 1 1 1 3 1 3 2 1 2 1 2 2 1 2 2 2 2 3 2 2 1 3 2 3 2 2 2 1 2 1 5 5 5 5 2 2 2 3 4 2 3 2 1 4 2 2 1 1 3 2 4 4 4 4 4 4 4 3 1 3 5 6 3 4 4 5 4 4 4 3 6 3 5 5 6 4 5 6 3 3 3 5 5 3 3 4 4 5 5 5 1 2 2 2 2 2 3 2 2 2 2 2 3 2 2 2 2 2 2 2 9 2 2 2 2 2 2 2 2 2 2 9 2 2 2 2 2 2 2 2 2 2 2 2 2 2 1 2 2 2 2 2 2 2 2 1 2 2 2 2 2 2 2 1 2 3 3 3 3 3 3 3 3 3 3 3 3 1 0 4 1 1 0 0 3 1 3 3 4 4 3 4 4 1 0 1 4 4 3&4 4 4 5 5 3 4 3 5 4 5 3 4 4 4 3 3 5 2 4 3 4 3 3 5 3 3 1 1 2 4 2 2 2 2 2 2 9 2 1 1 2 1 1 2 1 2 1 1 1 1 1 1 1 1 1 1 1 1 9 1 1 1 1 1 1 1 1 1 1 2 5 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 7 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 112 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY APPENDIX 2.?Continued. Number Species 32. 33. 34. 35. 36. 37. 38. 39. 40. 41. 42. 43. 44. 45. 46. 47. 48. 49. 50. 51. 52. 53. 54. 55. 56. 57. 58. 59. 60. 61. 62. 63. 64. 65. 66. 67. 68. 69. 70. 71. 72. 73. 74. 75. 76. 77. 78. 79. 80. 81. 82. 83. 84. 85. 86. 87. 88. 89. 90. 91. Maclurina logani Maclurina manitobensis Maclurites sedgewicki Maclurites expansa Ophileta complanata Lecanospira compacta Lecanospira nereine Barnesella ?lecanospiroides Malayaspira hintzei Malayaspira rugosa Barnesella measuresae Lytospira angelini Lytospira yochelsoni Maclurina ?annulata Rossospira harrisae Ecculiomphalus bucklandi Lytospira gerrula Lytospira ?norvegica Ophiletina cf. O. sublaxa Lytospira subrotunda Pararaphistoma lemoni Climacoraphistoma vaginati Lesueurilla bipatellare Lesueurilla marginalis Lesueurilla prima Palaeomphalus giganteus Climacoraphistoma damesi Eccyliopterus alatus Eccyliopterus ?princeps Eccyliopterus regularis Lesueurilla infundibula Eccyliopterus louderbacki Lesueurilla declivis Pararaphistoma qualteriata Pararaphistoma schmidti Helicotoma gubanovi Scalites katoi Helicotoma medfraensis Lesueurilla scotica Pachystrophia devexa Raphistoma striata Raphistomina lapicida Scalites angulatus Holopea insignis Eccyliopterus beloitensis Holopea rotunda Pachystrophia contigua Pachystrophia spiralis Raphistomina aperta Raph is torn ina fiss urata Eccyliopterus owenanus Holopea ampla Holopea pyrene Holopea symmetrica Raphistoma peracuta Raphistomina rugata Raphistoma tellerensis Sinutropis ?esthetica Pachystrophia gotlandica Lytospira triquestra 1 2 6 2 2 2 2 4 2 2 2 3 3 3 2 2 2 3 3 2 2 3 2 4 3 3 3 3 4 3 3 3 3 3 3 3 4 4 5 5 4 3 4 5 4 5 5 3 6 4 4 5 5 3 5 5 6 4 4 5 4 4 2 1 2 7 7 7 7 ? 2 2 3 2 2 7 2 2 1 2 2 2 2 2 3 2 2 7 2 2 1 1 2 2 8 9 3 3 3 2 3 2 3 1 2 1 2 1 2 1 ? i ? 1 2 1 ? 1 2 1 2 1 2 1 2 1 2 1 ? i 2 1 7 i 2 1 2 1 2 1 ? i 7 i 2 1 2 1 2 1 2 1 2 1 2 1 2 1 7 i 2 1 2 1 2 1 ? 2 ? 1 2 1 2 1 2 1 1 1 2 1 ? 1 2 1 ? i 1 1 ? 1 2 2 2 1 2 1 ? i ? 1 2 1 1 1 7 i 1 1 2 2 2 2 2 3 2 1 1 3 0 3 3 3 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n 2 n n n n n n n n n n n 4 n n n n n n n n 3 4 1 n 1 3 1 3 3 3 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n 3 n n n n n n n n n n n 1 n n n n n n n n 1 1 2 n 1 1 3 : 2 : I l i l i n n n n n n n n n 1 n n n 1 n n n n n n n n 1 n n n n n n n n n n n n 1 n n n n n n n n n n n i ; n n n n n n n n i : 1 . 1 n 1 3 4 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n > 1 n n n n n n n n > 3 > 4 n n 1 3 5 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n 1 n n n n n n n n 1 1 n n 1 3 6 n n n n n n n n n n II n n n n n n n n n n i i n n n n n n n n n n n n n n n n n n n n n n n n n 1 n n n n n n n n 1 1 n n 1 1 1 3 3 : 7 8 S n 1 n 1 n 1 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 1 n 1 n 1 n 1 n 1 n 1 1 n 1 1 n 1 n 1 n 1 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 ii 1 n 1 n 1 n 1 n 1 n 1 i n 1 n 1 n 1 n 1 n 1 n 1 n 1 1 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 1 1 1 I n 1 n 1 1 1 4 4 0 1 1 4 1 4 1 4 1 4 1 2 1 2 1 2 1 2 1 1 1 2 1 2 2 2 2 3 1 2 1 1 1 3 2 3 2 2 1 1 2 3 1 2 1 2 1 1 1 1 1 2 1 2 1 2 1 2 1 2 1 3 1 3 1 3 1 2 1 3 1 2 1 2 1 3 1 2 1 2 l 1 2 1 3 1 2 1 3 1 2 1 3 1 2 1 2 1 2 1 2 1 1 2 1 1 2 1 2 1 1 2 1 1 1 1 1 2 1 1 1 1 1 3 n 1 2 ti 1 2 n 1 3 1 4 2 ? ? ? 7 ? ? ? 7 ? ? ? ? ? ? 7 7 7 ? ? ? 7 ? 7 7 7 7 ? 7 ? ? ? 7 ? ? ? 7 7 7 ? 7 7 7 7 ? 7 7 7 7 1 4 3 n n n n n n n n n n n n n n n n n n n n n n n n n n n 3 n n n n n 3 n n n n n n n n n n n n n n n n n n n n n n n n n 3 NUMBER 88 113 APPENDIX 2.?Continued. Number Species 92. 93. 94. 95. 96. 97. 98. 99. 100. 101. 102. 103. 104. 105. 106. 107. 108. 109. 110. 111. 112. 113. 114. 115. 116. 117. 118. 119. 120. 121. 122. 123. 124. 125. 126. 127. 128. 129. 130. 131. 132. 133. 134. 135. 136. 137. 138. 139. 140. 141. 142. 143. 144. 145. 146. 147. 148. 149. 150. 151. Euomphalus tubus Lytospira subuloides Ceratopea unguis Boucotspira aff. B. ftmbriata Lophonema peccatonica Polehemia taneyensis Walcottoma frydai Helicotoma planulata Helicotoma tennesseensis Ophiletina sublaxa Ophiletina angularis Oriostoma bromidensis Euomphalopterus ?ordovicius Euomphalopterus aff. E. ordovicius Euomphalopterus cariniferus Palaeomphalus ?gradatus Trochomphalus ?dimidiatus Helicotoma blodgetti Helicotoma robinsoni Helicotoma? Girvan sp. Straporillina cf. S. circe Euomphalopterus alatus Euomphalopterus frenatus Euomphalopterus praetextus Euomphalopterus subcarinatus Euomphalopterus togatus Euomphalopterus undulans Grantlandispira christei Poleumita discors Pycnomphalus acutus Pycnomphalus obesus Discordichilus dalli Discordichilus mollis Discordichilus kolmodini Poleumita alata Poleumita octavia Poleumita rugosa Pseudophorus profundus Pseudophorus stuxbergi Siluriphorus gotlandicus Siluriphorus undulans Streptotrochus incisus Streptotrochus aff. S. incisus Streptotrochus lamellosa Streptotrochus lundgreni Streptotrochus? visbeyensis Hystricoceras astraciformis Poleumita granulosa Euomphalus walmstedti Centrifugus planorbis Spinicharybdis wilsoni Turbocheilus immaturum Pseudotectus comes Straparollus bohemicus Hormotoma artemesia Hormotoma confusa Hormotoma ?dubia Hormotoma ?simulatrix Ectomaria adelina "Hormotoma" "cassina" 1 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 1 2 2 1 2 2 2 2 2 1 1 1 2 2 2 1 7 7 7 1 1 1 1 2 1 2 2 2 1 2 1 1 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 n n n 2 2 2 9 9 7 7 ? 7 7 7 2 7 2 2 1 7 1 7 7 3 3 2 2 2 2 3 2 2 5 3 5 5 5 3 3 3 3 2 2 2 2 3 3 3 3 4 3 1 7 1 1 9 1 3 0 1 1 n n n 0 0 0 9 9 9 9 9 9 9 ? 1 9 0 0 5 9 0 9 9 4 4 4 4 4 4 4 2 2 5 3 5 5 5 3 3 3 3 2 2 2 2 3 3 3 3 4 3 1 7 1 1 9 1 3 0 1 1 n n n 0 0 0 9 9 9 9 9 9 9 7 1 9 0 0 3 9 0 9 9 6 6 4 4 4 4 5 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 3 2 2 2 n n n 2 2 2 9 9 9 7 ? ? 7 7 2 7 2 2 2 7 2 7 7 3 3 2 2 2 2 6 3 2 3 3 3 3 3 3 3 2 2 1 3 3 3 3 3 3 3 3 3 3 3 3 3 3 2 2 3 3 3 n n n 3 3 3 7 7 7 7 7 7 7 7 1 7 3 3 3 7 3 9 7 3 3 2 3 3 3 7 3 2 3 3 3 3 3 3 3 2 2 1 3 3 3 3 3 3 3 3 3 3 3 3 3 3 2 3 3 3 3 n n n 3 3 3 9 9 9 9 9 9 9 ? 1 9 3 3 1 9 3 9 7 4 3 2 3 3 3 8 2 1 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 n n n 2 2 2 7 9 7 7 2 2 7 2 2 7 2 2 2 9 2 7 7 1 2 2 2 2 2 9 1 1 4 3 4 3 3 3 3 3 3 2 2 2 2 3 1 3 3 3 2 1 2 1 2 2 1 1 2 1 1 n n n 2 2 2 7 9 9 7 7 7 7 7 2 7 2 2 4 7 1 9 9 3 3 3 3 3 3 1 0 1 3 4 3 4 3 3 3 3 3 3 2 2 2 2 3 1 3 3 3 2 1 2 1 2 2 1 1 2 1 1 n n n 2 2 2 7 7 9 9 7 ? 9 9 2 7 2 2 4 7 1 7 9 4 3 3 3 3 3 n n n 1 1 1 7 9 9 7 7 7 7 ? 1 7 1 1 1 9 1 9 9 1 1 1 1 1 1 1 2 2 2 2 2 9 2 2 2 2 7 1 3 1 1 1 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 7 1 4 n n n 2 2 2 2 2 1 2 2 2 2 2 2 2 2 1 1 2 2 n n n n n n 1 1 5 6 7 i 3 2 3 2 3 3 3 2 2 4 4 4 1 3 1 3 1 2 1 2 1 3 1 2 1 2 1 3 1 2 1 4 1 4 1 3 1 4 1 4 1 3 1 3 1 4 1 3 1 3 1 3 1 3 1 4 1 4 1 5 1 5 1 4 1 4 1 4 1 5 2 2 4 2 4 2 4 4 4 4 3 4 4 3 2 2 2 2 2 1 1 i ; I l I I I I I I I i I I I I I I I I I I I I I I I I I I 2 1 2 1 1 1 2 1 2 1 1 1 1 1 2 1 1 1 1 1 1 1 1 1 I 1 1 1 4 1 4 1 i : 1 1 1 1 1 1 1 1 1 1 1 1 i ; 2 1 2 2 I 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 I 9 1 1 ! 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 1&2 2 1 2 2 1 1 2 1 1 1 1 1 2 2 2 1 1 1 1 2 1 2 2 2 > 2 2 2 2 ? 1 I 1 1 1 1 2 1 2 1 2 1 2 1 2 1 2 2 0 n n 3 2 2 2 2 3 3 3 3 2 1 1 1 1 1 3 3 1 1 1 1 i i 1 1 n n 1 n n n n n 1 1 1 n n n n 1 n 1 1 1 1 2 1 1 n n n n 5 5 3 5 6 4 2 1 n n n n n 1 n n n n n 1 1 1 n n n n 1 n 2 n n n n 2 2 2 2 2 2 2 2 n n n n n n n n n 1 1 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n 1 n n n n 1 1 1 1 2 2 2 3 n n n n n n n n n 2 2 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n 1 n n n n 2 2 2 2 5 3 2 4 n n n n n n n n n 1 1 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n 1 n n n n 2 5 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n 114 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY APPENDIX 2.?Continued. Number Species 92. 93. 94. 95. 96. 97. 98. 99. 100. 101. 102. 103. 104. 105. 106. 107. 108. 109. 110. 111. 112. 113. 114. 115. 116. 117. 118. 119. 120. 121. 122. 123. 124. 125. 126. 127. 128. 129. 130. 131. 132. 133. 134. 135. 136. 137. 138. 139. 140. 141. 142. 143. 144. 145. 146. 147. 148. 149. 150. 151. Euomphalus tubus Lytospira subuloides Ceratopea unguis Boucotspira aff. B. ftmbriata Lophonema peccatonica Polehemia taneyensis Walcottoma frydai Helicotoma planulata Helicotoma tennesseensis Ophiletina sublaxa Ophiletina angularis Oriostoma bromidensis Euomphalopterus ?ordovicius Euomphalopterus aff. E. ordovicius Euomphalopterus cariniferus Palaeomphalus ?gradatus Trochomphalus ?dimidiatus Helicotoma blodgetti Helicotoma robinsoni Helicotoma? Girvan sp. Straporillina cf. S. circe Euomphalopterus alatus Euomphalopterus frenatus Euomphalopterus praetextus Euomphalopterus subcarinatus Euomphalopterus togatus Euomphalopterus undulans Grantlandispira christei Poleumita discors Pycnomphalus acutus Pycnomphalus obesus Discordichilus dalli Discordichilus mollis Discordichilus kolmodini Poleumita alata Poleumita octavia Poleumita rugosa Pseudophorus profundus Pseudophorus stuxbergi Siluriphorus gotlandicus Siluriphorus undulans Streptotrochus incisus Streptotrochus aff. S. incisus Streptotrochus lamellosa Streptotrochus lundgreni Streptotrochus? visbeyensis Hystricoceras astraciformis Poleumita granulosa Euomphalus walmstedti Centrifugus planorbis Spinicharybdis wilsoni Turbocheilus immaturum Pseudotectus comes Straparollus bohemicus Hormotoma artemesia Hormotoma confusa Hormotoma ?dubia Hormotoma ?simulatrix Ectomaria adelina "Hormotoma" "cassina" 2 6 n n n n n n n n n n M n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n 2 7 n n 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 n 2 2 n n 2 n n n n n 2 2 2 n n n 1 2 n 2 2 2 2 2 2 1 n n n n 1 1 2 1 1 1 2 8 n n 1 2 1 1 2 3 3 3 3 2 2 2 2 1 2 3 3 3 1 2 5 n 2 5 n n 2 n n n n n 2 2 2 n n n n 5 n 2 5 2 5 2 2 n n n n n n n 1 n n n 2 9 n n 3 3 3 3 3 4 4 3 3 4 4 4 3 3 2 4 4 4 1 1&3 1 n 3 3 n n 3 n n n n n 3 3 3 n n n n 1 n 4 1 3 1 4 2 n n n n n n n 2 n n n 3 0 n n 2 2 n 2 2 n n 1 n n n n n 1 1 1 n n n n 2 n 1 2 1 2 1 1 n n n n n n n n n n n 3 1 n n n n n 1 n n n n n 1 1 1 n n n n n n n n n n n 1 n n n 3 2 n n n n n 2 n n n n n 1 2 2 n n n 1 1 n 1 1 1 1 2 1 n n n n n n n 1 n n n 3 3 3 3 4 5 n n i : n n n 1 n n n n n 1 1 1 n n n n n '. n n n n 2 1 3 1 1 1 n n n n n n n n n n > 2 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n I 2 n n n n n n n n n n 3 3 3 6 7 8 n n 1 n n 1 n n n n n n n n 1 n n n n n n n n n n n n n n n n n i n n r n i n n n n n n n n n n n n n n n n n n 1 n n l n n n n n n n n 1 4 1 I 1 4 4 1 1 4 4 4 4 4 1 n 4 4 1 1&4 4 1 4 4 1 l 1 4 l 1 l 1 5 5 5 4 4 4 5 5 5 5 6 6 4 6 4 6 4 4 1 n l 1 5 5 1 1 1 1 1 1 3 9 2 2 2 2 1 1 1 2 2 3 3 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 n 2 2 2 2 2 2 2 2 2 2 2 1 n 2 2 2 2 2 2 2 2 2 4 0 n n n n n 1 n II n n n 1 1 1 n n n 1 1 n 1 n 1 1 1 1 1 n n n n 1 1 2 1 1 1 4 1 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n 1 n n n n 1 1 2 1 1 1 4 2 n n 2 1 1 1 1 2 2 2 2 1 1 1 1 1 1 2 2 2 1 1 1 n 1 1 n n 1 n n n n n 1 1 1 n n n 1 1 n 1 n 1 1 1 1 1 n n n n 4 3 n n 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 n 2 2 n n 2 n n n n n 2 2 2 n n n 2 2 n 2 n 2 2 2 2 2 n n n n 2 2 2 2 2 2 4 4 n n n n n 1 n n n n n 1 1 1 n n n 1 1 n 1 n 1 1 1 1 1 n n n n 4 5 1 1 2 2 2 2 2 2 2 2 2 2 2 2 2 2 ? 1 1 2 2 2 1 1 2 2 1 1 2 1 1 1 1 1 2 2 2 1 1 1 1 2 1 2 I 2 1 2 2 1 ? 1 1 ? 1 1 2 1 1 1 4 4 4 4 5 6 7 8 9 0 n n 1 2 2 2 2 1 1 1 1 1 1 1 1 1 1 1 2 1 2 1 2 1 1 1 n 1 1 n 1 1 1 1 2 1 n n 2 1 1 1 n n 1 1 n n n n n 2 1 1 1 1 1 n n n n 1 n 2 n 2 n 1 1 n n n n n n n 1 n n n A 7 5 : 7 6 : 6 : 8 1 8 1 8 7 1 7 1 A 1 6 1 6 1 8 1 8 1 9 1 9 1 6 1 7 1 9 1 9 1 A 1 9 1 8 1 C 1 9 1 A 1 C 1 A 1 A 1 C 1 C 1 D 1 9 1 C 1 C B 1 9 C C C C 8 B A B C B F 9 A I C I A I 8 1 8 1 7 1 7 1 9 1 8 n n 1 n 1 1 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n 1 n 1 n 1 n [ n 1 n 1 n NUMBER 88 115 APPENDIX 2.?Continued. Number Species 92. 93. 94. 95. 96. 97. 98. 99. 100. 101. 102. 103. 104. 105. 106. 107. 108. 109. 110. 111. 112. 113. 114. 115. 116. 117. 118. 119. 120. 121. 122. 123. 124. 125. 126. 127. 128. 129. 130. 131. 132. 133. 134. 135. 136. 137. 138. 139. 140. 141. 142. 143. 144. 145. 146. 147. 148. 149. 150. 151. Euomphalus tubus Lytospira subuloides Ceratopea unguis Boucotspira aff. B. ftmbriata Lophonema peccatonica Polehemia taneyensis Walcottoma frydai Helicotoma planulata Helicotoma tennesseensis Ophiletina sublaxa Ophiletina angular is Oriostoma bromidensis Euomphalopterus ?ordovicius Euomphalopterus aff. E. ordovicius Euomphalopterus cariniferus Palaeomphalus ?gradatus Trochomphalus ?dimidiatus Helicotoma blodgetti Helicotoma robinsoni Helicotoma? Girvan sp. Straporillina cf. S. circe Euomphalopterus alatus Euomphalopterus frenatus Euomphalopterus praetextus Euomphalopterus subcarinatus Euomphalopterus togatus Euomphalopterus undulans Grantlandispira christei Poleumita discors Pycnomphalus acutus Pycnomphalus obesus Discordichilus dalli Discordichilus mollis Discordichilus kolmodini Poleumita alata Poleumita octavia Poleumita rugosa Pseudophorus profundus Pseudophorus stuxbergi Siluriphorus gotlandicus Siluriphorus undulans Streptotrochus incisus Streptotrochus aff. S. incisus Streptotrochus lamellosa Streptotrochus lundgreni Streptotrochus? visbeyensis Hystricoceras astraciformis Poleumita granulosa Euomphalus walmstedti Centrifugus planorbis Spinicharybdis wilsoni Turbocheilus immaturum Pseudotectus comes Straparollus bohemicus Hormotoma artemesia Hormotoma confusa Hormotoma ?dubia Hormotoma ?simulatrix Ectomaria adelina "Hormotoma" "cassina" 5 1 3 3 1 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 7 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 3 2 3 2 2 2 5 2 2 3 2?3 3 3 3 3 3 3 3 3 3 3 3 3 3 2 3 3 3 2 1 1 1 3 3 1 2 3 2 2 2 2 2 3 2 2 9 3 3 3 3 3 3 2 3 2 3 3 3 1 2 2 2 2 2 3 2 3 3 5 3 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 3 7 1 3 3 2 2 2 2 2 2 2 2 2 2 2 9 3 2 2 3 2 3 3 3 3 2 2 2 1 2 2 2 2 2 2 2 2 2 5 4 3 2 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 1 1 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 2 3 3 3 3 3 3 3 3 3 3 1 1 2 1 3 3 5 5 2 4 273 2 3 3 2 3 3 4 4 3 1 1 2 3 1 4 3 3 1 3 1 3 3 2 1 0 2 0 0 1 1 1 2 2 2 0 2 2 2 2 2 3 2 2 2 3 2 0 1 0 1 2 5 5 5 5 4 5 5 6 7 4 3 3 4 4 5 4 4 8 8 4 2 2 4 5 4 2 2 4 4 5 4 4 4 4 4 5 5 5 5 5 5 7 4 4 5 6 4 4 4 4 4 3 4 3 5 5 5 9 4 5 6 4 4 4 5 4 5 5 5 7 2 2 2 2 2 2 2 2 1 1 2 2 2 1 1 1 1 1 2 2 2 2 1 1 1 2 2 2 2 2 1 1 1 1 2 1 2 1 1 2 1 1 2 1 2 1 2 2 2 1 1 5 8 4 5 1-3 4 4 4 5 4 4 4 4 3&4 5 5 5 4 5 3 4 4 5 5 7 5 5 6 6 6 5 6 6 6 6 8 5 5 5 5 5 7 7 6 6 6 7 5 7 5 5 7 7 6 8 5 5 4&f 5 5 4 4 5 6 9 C 3 1 4 1 2 2 4 1 3 1 3 1 3 1 2 1 4 1 3 1 3 1 3 1 5 1 5 1 5 1 4 1 5 1 2 1 2 1 4 1 5 1 5 1 4 ' 5 1 4 1 5 4 1 6 5 6 6 5 4 6 : 5 5 5 5 5 5 5 5 5 6 5 5 5 6 5 1 4 6 6 5 5 5 5 5 2 3 6 1 n n 2 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n ! n n n n n n n n n n n n [ n [ n [ n 1 n 1 n I n 1 n 1 n 1 n 2 2 2 1 2 1 2 1 1 n 1 n 6 6 2 3 n 1 n 1 3 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n n n n 1 n n 1 n n n n n n n n n n n n n n n n n n n n n 3 n 3 3 n n 6 4 1 1 1 2 2 2 2 2 2 2 2 2 2 2 2 1 2 2 2 2 1 2 1 2 2 2 1 1 2 1 1 1 1 1 2 2 2 1 1 1 1 1 1 1 1 2 1 1 1 2 1 1 1 2 1 2 1 1 1 1 1 1 1 1 1 2 1 1 1 1 1 1 1 1 1 2 1 2 6 5 n n n 3 3 3 1 1 1 3 3 3 3 3 3 n 2 2 2 1 n 1&2 n 1 3 2 n n 1 n n n n n 3 1 1 n n n n n n 3 n 3 n 1 1 n n n n 1 n n n n 1 1 6 6 n n n 4 4 4 2 3 3 4 4 3 4 4 4 n 4 3 3 3 n 4 n 1 4 4 n n 4 n n n n n 4 1 1 n n n n n n 4 n 4 n 4 4 n n n n 1 n n n n 1 1 6 7 n n n 1 1 1 1 2 2 1 1 2 1 1 1 n 1 2 2 2 n 2 n 1 2 2 n n 1 n n n n n 1 1 1 n n n n n n 1 n 1 n 1 1 n n n n 1 n n n n 1 1 6 6 8 9 n 1 n 7 n 1 2 1 2 1 2 1 n 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 n 1 1 1 2 1 2 1 2 1 n 1 2 1 n 1 2 1 2 1 2 1 n n 1 2 1 n n n n 2 n 2 2 1 2 7 7 0 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 1 2 . 1 2 n 1 n 1 n 1 n 2 1 1 n 2 1 1 n 2 1 1 n ^ n n 2 n 2 n 2 2 n n n n 2 n n n n 1 1 I 1 1 n n I n I n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 2 1 1 n 1 n 1 n 1 n 1 n 1 n 1 n 7 2 1 2 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 2 1 1 3 3 3 1 2 3 2 2 2 2 2 3 2 2 4 3 3 3 3 3 3 2 3 2 4 1 2 1 3 1 1 1 1 2 2 1 2 1 1 1 2 1 3 1 2 1 3 1 3 7 7 3 4 1 n 1 n 1 n 1 n 1 n 1 n 1 n 2 1 2 1 1 n 1 n 1 n 1 n 1 " 1 n 2 1 1 n 2 1 2 1 2 1 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n ? n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 2 2 2 1 2 1 2 1 1 n I n 7 5 1 1 1 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 1 2 2 2 2 1 2 2 2 2 2 2 2 2 2 2 2 2 1 2 2 1 2 1 1 1 1 1 2 2 116 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY Number Species 92. 93. 94. 95. 96. 97. 98. 99. 100. 101. 102. 103. 104. 105. 106. 107. 108. 109. 110. 111. 112. 113. 114. 115. 116. 117. 118. 119. 120. 121. 122. 123. 124. 125. 126. 127. 128. 129. 130. 131. 132. 133. 134. 135. 136. 137. 138. 139. 140. 141. 142. 143. 144. 145. 146. 147. 148. 149. 150. 151. Euomphalus tubus Lytospira subuloides Ceratopea unguis Boucotspira aff. B. ftmbriata Lophonema peccatonica Polehemia taneyensis Walcottoma frydai Helicotoma planulata Helicotoma tennesseensis Ophiletina sublaxa Ophiletina angularis Oriostoma bromidensis Euomphalopterus ?ordovicius Euomphalopterus aff. E. ordovicius Euomphalopterus cariniferus Palaeomphalus ?gradatus Trochomphalus ?dimidiatus Helicotoma blodgetti Helicotoma robinsoni Helicotoma? Girvan sp. Straporillina cf. 5. circe Euomphalopterus alatus Euomphalopterus frenatus Euomphalopterus praetextus Euomphalopterus subcarinatus Euomphalopterus togatus Euomphalopterus undulans Grantlandispira christei Poleumita discors Pycnomphalus acutus Pycnomphalus obesus Discordichilus dalli Discordichilus mollis Discordichilus kolmodini Poleumita alata Poleumita octavia Poleumita rugosa Pseudophorus profundus Pseudophorus stuxbergi Siluriphorus gotlandicus Siluriphorus undulans Streptotrochus incisus Streptotrochus aff. S. incisus Streptotrochus lamellosa Streptotrochus lundgreni Streptotrochus? visbeyensis Hystricoceras astraciformis Poleumita granulosa Euomphalus walmstedti Centrifugus planorbis Spinicharybdis wilsoni Turbocheilus immaturum Pseudotectus comes Straparollus bohemicus Hormotoma artemesia Hormotoma confusa Hormotoma ?dubia Hormotoma ?simulatrix Ectomaria adelina "Hormotoma" "cassina' 1 6 n n n 2 2 2 2 1 1 4 4 1 3 3 3 1 2 4 4 1 1 3 3 3 3 3 3 2 1 3 n 1 1 1 3 n 1 3 3 4 4 2 2 2 2 2 2 1 n 1 3 n 1 n n n n n 1 1 7 7 n n n 2 2 2 2 1 1 3 3 2 3 3 3 1 3 3 3 2 1 4 4 4 4 4 4 3 3 3 n 2 1 1 3 n 3 4 4 3 3 2 2 3 4 2 4 3 n 3 4 n 1 n n n n n 1 1 7 8 n n n 2 2 2 2 2 2 2 2 2 2 2 2 2 1 2 2 2 n 2 2 2 2 2 2 1 2 1 n 2 2 2 2 n 2 2 2 2 1 2 2 2 2 2 2 2 n 2 2 n 2 n n n n n 1 1 7 9 n n n n n n n n n 1 1 n 1 1 1 n n n n n n 2 2 2 2 2 2 n n 1 n n n n 1 n n 2 2 1 n n n 1 n n n n n n 2 n n n n n n n n n APPENDIX 2 ? 8 0 n n n n n n n n n 1 1 n 1 1 1 n n n i i n n 1 2 2 1 2 1 n n 1 n n n n 1 n n 1 1 1 n n n 1 n n n n n n 1 n n n n n n n n n 8 I n n n n n n n n n 1 1 n 1 1 1 n n n n n n 1 2 2 1 1 2 n n 1 n n n n 1 n n 1 1 1 n n n 1 n n n n n n n n n n n n n n n n 8 2 n n n n n n n n n 1 1 n 1 1 1 n n n n n n 2 2 2 2 2 2 n n 1 n n n n 1 n n I 1 1 n n n 1 n n n n n n 2 n n n n n n n n n 8 3 n n n n n n n n n 2 2 n 3 3 3 n n n n n n 3 3 3 3 3 3 n n 3 n n n n 3 n n 3 3 3 n n n 3 n n 1 n n n 3 n n n n n n n n n -Continued 8 4 n n n n n n n n n 1 1 n 1 1 1 n n n n n n 1 1 1 1 1 1 n n 1 n n n n 1 n n 2 2 1 n n n 1 n n n n n n 1 n n n n n n n n n 8 8 8 5 6 7 n n n n n 2 2 n n n n 1 n n 2 n n 1 2 n n n n n n n 1 1 I 1 2 1 1 1 1 2 1 2 2 1 3 3 1 1 2 2 2 1 2 1 1 1 1 1 1 3 2 3 3 2 2 2 1 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 1 2 2 2 2 2 2 2 2 2 8 8 n n 2 n n n n 3 n 2 2 n 2 2 n n 2 3 3 n 2 n n n n n n 2 2 2 2 2 1 1 n 2 2 3 3 2 2 2 2 2 2 2 2 2 2 2 n 2 1 2 2 2 2 2 3 3 8 9 1 1 2 2 2 2 2 2 2 2 2 2 2 2 2 1 1 2 2 1 1 2 1 2 2 2 2 2 2 2 9 0 n n 3 3 3 3 3 1 1 3 3 3 4 4 4 n n 1 1 n n 1&4 n 1 4 1 n n n n n n n n 4 n n 3 3 3 n n n n n n n n n n n n n n n n n n n n 9 1 n n 1&3 1 1 1 1 1 1 3 3 3 1 1 1 n n 1 1 n n 1 n 1 1 1 n n n n n n n n 1 n n 1 1 1 n n i i n n n n n n n n n n n n n n n n n 9 2 n n 2 2 3 2 2 2 n n 1 1 n n 2 n 2 2 2 n n n n n n n n 2 n n 2 2 2 n n n n n n n n n n n n n n n n n n n n 9 3 n n 1 2 2 2 2 2 2 3 3 2 2 2 2 n n 2 2 n n 2 n 1 2 1 n n n n n n n n 2 n n 1 1 1 n n n n n n n n n n n n n n n n n n n n 9 4 3 3 4 3 3 3 3 3 3 5 5 3 5 5 5 3 3 3 3 3 3 4 4 4 5 5 3 3 3 3 3 5 6 6 4 3 3 5 5 5 4 5 5 4 5 5 5 3 3 3 5 3 6 3 3 4 4 4 3 4 9 5 2 2 2 2 2 2 2 2 2 2 2 2 1 1 2 1 2 2 2 1 2 2 2 1 2 2 1 2 3 1 1 2 2 3 3 3 3 2 2 2 2 2 2 2 2 2 2 3 3 3 2 1 2 1 3 2 2 2 3 3 1 9 9 9 9 0 6 7 8 9 0 2 2 2 1 2 1 2 1 2 2 2 1 2 1 2 1 2 1 1 1 1 1 2 1 2 1 1 1 2 1 2 1 2 1 1 1 2 1 2 1 2 1 2 1 2 1 2 1 1 1 1 1 1 1 1 1 3 1 3 1 1 1 1 1 1 1 1 1 2 1 2 2 2 3 2 2 3 2 3 1 2 2 2 1 1 2 2 2 2 2 2 2 3 7 4 4 1 4 1 4 1 3 1 4 1 4 1 4 1 4 1 3 1 4 1 4 1 4 1 4 1 3 1 3 1 5 1 4 1 3 1 3&4 1 3 1 3 1 4 1 2 1 3 1 3 1 3 1 3 1 3 1 2 1 2 1 1 1 3 1 1 1 3 1 3 1 3 1 2 1 2 2 2 2 2 2 2 3 4 3 4 3 1 2 1 4 2 2 2 2 2 NUMBER 88 117 APPENDIX 2.?Continued. Number Species 92. 93. 94. 95. 96. 97. 98. 99. 100. 101. 102. 103. 104. 105. 106. 107. 108. 109. 110. 111. 112. 113. 114. 115. 116. 117. 118. 119. 120. 121. 122. 123. 124. 125. 126. 127. 128. 129. 130. 131. 132. 133. 134. 135. 136. 137. 138. 139. 140. 141. 142. 143. 144. 145. 146. 147. 148. 149. 150. 151. Euomphalus tubus Lytospira subuloides Ceratopea unguis Boucotspira aff. B. ftmbriata Lophonema peccatonica Polehemia taneyensis Walcottoma frydai Helicotoma planulata Helicotoma tennesseensis Ophiletina sublaxa Ophiletina angularis Oriostoma bromidensis Euomphalopterus ?ordovicius Euomphalopterus aff. E. ordovicius Euomphalopterus cariniferus Palaeomphalus ?gradatus Trochomphalus ?dimidiatus Helicotoma blodgetti Helicotoma robinsoni Helicotoma? Girvan sp. Straporillina cf. S. circe Euomphalopterus alatus Euomphalopterus frenatus Euomphalopterus praetextus Euomphalopterus subcarinatus Euomphalopterus togatus Euomphalopterus undulans Grantlandispira christei Poleumita discors Pycnomphalus acutus Pycnomphalus obesus Discordichilus dalli Discordichilus mollis Discordichilus kolmodini Poleumita alata Poleumita octavia Poleumita rugosa Pseudophorus profundus Pseudophorus stuxbergi Siluriphorus gotlandicus Siluriphorus undulans Streptotrochus incisus Streptotrochus aff. S. incisus Streptotrochus lamellosa Streptotrochus lundgreni Streptotrochus? visbeyensis Hystricoceras astraciformis Poleumita granulosa Euomphalus walmstedti Centrifugus planorbis Spinicharybdis wilsoni Turbocheilus immaturum Pseudotectus comes Straparollus bohemicus Hormotoma artemesia Hormotoma confusa Hormotoma ?dubia Hormotoma ?simulatrix Ectomaria adelina "Hormotoma" "cassina" 1 1 0 C 1 2 ? 1 ? 7 ? 2 ? i 7 ] 7 | 7 ; 7 7 7 7 7 ] 7 1 ? i ? ] 7 1 ? ? ? i ? ? 7 ? 7 7 2 ? 2 2 7 7 ? ? 7 7 7 7 7 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 1 0 3 4 4 4 3 3 3 3 3 3 3 3 4 3 3 4 3 4 3 3 3 4 4 3 4 4 4 3 3 5 2 2 3 3 2 5 5 5 2 2 2 2 3 2 2 3 2 3 5 5 I 5 3 1 2 1 2 I 5 1 3 1 3 1 3 1 3 1 3 1 3 1 1 0 C 4 f 7 3 2 2 1 2 2 3 3 3 2 2 2 2 3 3 3 1 7 2 2 1 0 6 I 2 1 2 1 2 1 2 1 2 1 2 1 2 1 1 0 C 7 i 2 2 2 1 2 1 2 2 2 2 1 0 9 2 2 2 2 2 2 1 1 1 2 2 2 2 2 2 2 2 2 2 I 2 1 1 1 1 0 1 2 1 7 1 7 1 7 1 ? 1 7 1 ? 1 ? | ? j 2 1 2 1 ? i 7 i 7 i 7 i 7 i 7 ] ? i ? i 7 j 7 i 7 1 7 1 7 i 7 i 7 i 7 7 1 7 j 7 i 7 1 4 1 5 1 5 1 7 7 ] 7 3 3 2 2 3 2 2 4 3 3 7 7 ? ? 9 5 9 ? 9 9 7 ? 9 1 1 1 1 2 3 t. 1 1 7 1 ? 1 7 1 ? i 7 1 7 1 7 1 7 1 7 i ?; ? 1 7 [ 7 i r, ? 1 7 1 ? i i 7 i 7 1 7 1 7 j 7 1 7 1 ? i 7 1 7 1 7 j 7 1 7 1 7 i 7 1 7 1 7 1 7 i 7 2 : ? 2 : 7 2 . 7 2 7 1 7 2 1 ? 1 1 7 1 1 7 1 1 7 1 1 7 1 1 ? 1 1 ? 1 1 1 1 1 1 1 1 5 6 7 1 1 n 1 2 1 1 2 2 1 1 n 1 1 n 1 1 n 1 1 n 1 1 n 1 1 n 1 1 2 1 1 2 1 1 n 1 1 n 1 1 n 1 1 n 1 1 n 1 1 n 1 1 2 1 1 n 1 1 n 1 1 n 1 1 n 1 1 n 1 1 n 1 1 n 1 1 n 1 1 n 1 1 n 1 2 2 1 1 n 1 1 n 1 1 n ? 1 1 n ' 1 1 n ' 1 2 2 ' 1 1 n ' 1 2 2 ? 1 1 n > 1 1 n 5 1 1 n $ 1 1 n > 1 1 n 1 1 1 n ? 1 1 n > 1 1 n ? 1 1 n ? 1 1 n ? 1 2 1 7 1 2 1 7 1 2 2 7 1 1 n 7 1 1 n 7 1 1 n 7 1 2 1 7 1 1 n ? 1 1 n 7 1 1 n 7 1 1 n 7 1 1 n ? 1 1 n 1 1 8 2 1 1 2 2 2 2 2 2 2 2 2 2 2 2 2 7 2 2 2 2 2 2 2 2 2 2 2 1 2 2 2 2 2 1 1 1 2 2 2 2 2 2 2 2 2 2 1 1 1 2 2 2 1 1 1 1 1 1 1 1 1 9 2 n n 2 1 1 1 2 2 1 1 1 3 3 2 1 n 2 2 2 2 2 2 2 2 2 3 2 n 2 2 5 3 5 n n n 5 5 5 5 3 2 3 2 3 5 n n n 1 2 3 n n n n n n n 1 2 0 1 2 2 1 2 2 2 1 1 1 1 1 2 2 2 1 7 1 1 1 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 1 2 2 2 2 1 2 2 2 2 1 2 1 4 3 4 3 3 3 3 4 4 4 4 3 3 3 3 4 3 4 4 4 3 2 2 2 2 2 2 3 3 3 3 2 3 3 3 4 4 2 2?3 1 1 2 2 2 2 3 1 3 3 3 2 3 3 3 1 2 2 2 1 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 3 2 3 3 2 3 2 2 2 2 2 1 2 2 2 2 2 2 2 1 1 2 2 3 4 4 1 1 1 4 1 4 1 5 1 5 1 5 1 3 1 3 1 3 2 3 1 7 1 5 1 5 1 4 1 3 1 4 1 3 1 3 1 4 1 4 1 5 1 5 1 5 1 5 1 5 1 4 1 4 1 4 1 4 1 4 1 5 1 4 1 4 1 4 1 5 1 4 1 4 1 4 1 4 1 4 1 5 1 5 1 5 1 5 1 5 1 6 1 4 1 4 1 3 1 5 1 4 1 4 1 4 1 6 1 6 1 6 1 6 1 6 1 6 1 1 2 5 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 118 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY APPENDIX 2.?Continued. Number Species 92. 93. 94. 95. 96. 97. 98. 99. 100. 101. 102. 103. 104. 105. 106. 107. 108. 109. 110. 111. 112. 113. 114. 115. 116. 117. 118. 119. 120. 121. 122. 123. 124. 125. 126. 127. 128. 129. 130. 131. 132. 133. 134. 135. 136. 137. 138. 139 140. 141. 142. 143. 144. 145. 146. 147. 148. 149. 150. 151. Euomphalus tubus Lytospira subuloides Ceratopea unguis Boucotspira aff. B. ftmbriata Lophonema peccatonica Polehemia taneyensis Walcottoma frydai Helicotoma planulata Helicotoma tennesseensis Ophiletina sublaxa Ophiletina angularis Oriostoma bromidensis Euomphalopterus ?ordovicius Euomphalopterus aff. E. ordovicius Euomphalopterus cariniferus Palaeomphalus ?gradatus Trochomphalus ?dimidiatus Helicotoma blodgetti Helicotoma robinsoni Helicotoma? Girvan sp. Straporillina cf. S. circe Euomphalopterus alatus Euomphalopterus frenatus Euomphalopterus praetextus Euomphalopterus subcarinatus Euomphalopterus togatus Euomphalopterus undulans Grantlandispira christei Poleumita discors Pycnomphalus acutus Pycnomphalus obesus Discordichilus dalli Discordichilus mollis Discordichilus kolmodini Poleumita alata Poleumita octavia Poleumita rugosa Pseudophorus profundus Pseudophorus stuxbergi Siluriphorus gotlandicus Siluriphorus undulans Streptotrochus incisus Streptotrochus aff. S. incisus Streptotrochus lamellosa Streptotrochus lundgreni Streptotrochus? visbeyensis Hystricoceras astraciformis Poleumita granulosa Euomphalus walmstedti Centrifugus planorbis Spinicharybdis wilsoni Turbocheilus immaturum Pseudotectus comes Straparollus bohemicus Hormotoma artemesia Hormotoma confusa Hormotoma ?dubia Hormotoma ?simulalrix Ectomaria adelina "Hormotoma" "cassina" 1 2 6 5 2 4 5 5 5 5 4 4 4 4 7 5 5 5 4 4 4 4 5 4 5 6 5 5 5 5 5 4 5 5 6 6 6 5 6 4 5 5 6 6 6 6 6 6 6 6 4 4 3 6 5 6 5 8 7 7 7 7 7 1 2 7 1 1 2 1 2 2 1 2 1 1 1 1 1 1 1 1 1 2 2 1 1 1 1 2 1 1 1 1 3 1 1 1 1 1 3 1 3 1 1 1 1 2 1 1 2 1 2 3 1 1 1 1 1 1 2 2 2 2 1 1 1 2 8 2 2 2 ? ? ? ? 1 1 ? 7 1 7 7 2 7 7 1 ? ? ? 2 2 2 2 2 2 ? 2 2 2 7 7 ? 2 2 2 2 7 2 2 7 ? ? 2 ? 2 2 2 2 ? ? ? 2 1 2 9 2 1 1 1 1 2 2 3 3 3 2 2 2 2 2 2 3 1 3 0 4 n n n n 1 1 3 n n n n n n n n n 1 3 n n n n n n n n 3 2 n n i i n n 2 3 3 n n i i n n n n n n n 2 n 2 n n n n n n n n n n 1 3 1 1 n n n n 3 3 3 n n n n n n n n n 1 3 n n n n n n n n 1 3 n n n n n 3 3 3 n n n n n n n n n n 1 n 3 n n n n n n n n n n 1 3 2 1 n n n n 2 2 2 n n n n n n n n n 2 2 n n n n n n n n 1 1 n n n n n 1 1 1 n n n n n n n n n n 1 n 1 n n n n n n n n n n 1 3 3 2 7 2 2 2 2 2 2 1 3 4 4 n n n n n n n n n n n n n n n n n n n n n n n n n n n 1 n n n n n 1 3 2 n n n n n n n n n n 1 n 2 n n n n n n n n n n 1 3 5 1 n n n n n n n n n n n n n n n n n n n n n n n n n n n 2 n n n n n 2 2 1 n n n n n n n n n n 2 n 2 n n n n n n n n n n 1 3 6 1 n n n n n n n n n n n n n n n n n n n n n n n n n n n 1 n n n n n 1 1 1 n n n n n n n n n n 1 n 1 n n n n n n n n n n 1 1 3 3 7 8 1 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 7 n ? n 7 n 1 n 7 n 7 n ? 1 1 n 7 n ? n 1 n 1 n 1 1 1 1 1 1 1 n 7 n 7 n ? n ? n ? n ? n ? n 7 n 7 n ? 1 1 n 7 1 1 n 1 n ? n 1 n ? n 1 n 1 n 1 n 1 n 1 n 1 1 1 1 3 4 4 9 0 1 n n n n 1 1 1 1 n 1 1 1 1 n n n n n 1 1 1 1 1 1 n n n n n n n n n n n n 1 ? 3 2 2 2 ? 2 2 1 1 1 2 2 2 2 ? 2 2 2 2 3 2 2 2 2 2 3 3 2 2 1 1 2 2 2 3 2 2 2 2 1 1 2 2 1 2 3 3 3 2 2 2 2 2 I 2 I 2 2 1 1 1 1 4 2 ? ? ? ? ? ? ? ? ? ? ? 7 7 ? ? 7 7 ? 7 ? 7 ? ? 7 ? ? ? ? ? ? ? 7 7 7 7 ? ? ? ? ? 7 7 7 7 ? ? 7 ? 7 ? 7 7 7 ? 7 ? ? ? ? ? 1 4 3 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n NUMBER 88 119 APPENDIX 2.?Continued. Number Species 152. 153. 154. 155. 156. 157. 158. 159. 160. 161. 162. 163. 164. 165. 166. 167. 168. 169. 170. 171. 172. 173. 174. 175. 176. 177. 178. 179. 180. 181. 182. 183. 184. 185. 186. 187. 188. 189. 190. 191. 192. 193. 194. 195. 196. 197. 198. 199. 200. 201. 202. 203. 204. 205. 206. 207. 208. 209. 210. 211. Fusispira Smithville Fm. sp. Hormotoma augustina Hormotoma zetleri Lophospira perangulata Subulitid El Paso Fm. sp. Pagodospira cicelia Plethospira cannonensis Plethospira cassina Seelya ventricosa Lophospira grandis Straparollina pelagica Plethospira? turgida Turritoma acrea Turritoma Cotter Fm. omate sp. Turritoma cf. T acrea Hormotoma Setul Fm. sp. Turritoma ?anna Murchisonia callahanensis Ectomaria prisca Hormotoma gracilis Daidia cerithioides Ectomaria pagoda Haplospira ?nereis Hormotoma bellicincta Hormotoma salteri Hormotoma trentonensis Loxonema murrayana Omospira alexandra Omospira laticincta Straparollina circe Straparollina erigione Girvania excavata Murchisonia Pt. Clarence Fm. sp. Rhabdostropha primitiva Spiroecus girvanensis Daidia aff. D. cerithioides Ectomaria cf. E. pagoda Ectomaria cf. E. prisca Ectomaria laticarinata Ectomaria nieszkowskii Hormotoma insignis Holopella regularis Hormotoma centervillensis Hormotoma cingulata Kjerulfonema cancellata Kjerulfonema quinquecincta Cyrtostropha coralli Goniostropha cava Hormotoma subplicata Hormotoma monoliniformis Hormotoma attenuata Loxonema? attenuata Macrochilus fenestratus Rhabdostropha grindrodii Loxonema crossmanni Loxonema sinuosa Auriptygma fortior Catazone allevata Catazone argolis Catazone cunea 1 1 2 2 2 1 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 1 2 1 2 2 2 2 2 2 2 2 2 2 2 2 1 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 1 2 2 2 2 7 3 3 2 7 2 2 2 2 2 2 2 3 2 2 3 3 2 2 3 7 2 7 3 3 2 2 2 2 2 2 2 2 2 2 7 2 2 2 2 3 2 3 3 2 2 3 3 3 3 3 3 2 2 2 2 7 3 3 3 3 ? 4 3 4 7 4 3 3 3 3 1 4 4 4 4 2&3 3 3 4 3 7 4 7 4 3 4 4 2 2 1 1 2 1 2 2 7 2 4 3 4 3 1 4 3 0 0 4 2 4 3 3 3 0 1 4 3 7 2 2 2 4 2 7 6 4 2 4 3 3 3 3 1 4 5 4 4 7 5 3 4 7 7 4 7 6 7 4 4 2 2 1 1 2 1 2 2 7 2 4 3 4 5 1 7 7 0 0 7 5 4&5 5 5 5 0 1 4 3 7 7 7 7 5 ? 3 3 2 ? 2 2 2 2 2 2 2 1 2 2 3 3 2 2 3 7 2 7 3 3 3 3 3 3 2 2 2 3 3 2 7 2 2 2 2 1 3 3 1 3 3 3 1 2 3 3 3 3 3 3 3 7 1 1 1 6 ? 3 3 3 7 3 2 2 2 2 1 1 3 3 3 3 3 3 3 3 7 3 7 3 3 3 3 3 3 2 2 3 3 3 3 7 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 7 3 2 3 7 ? 4 4 3 7 3 2 2 2 2 1 1 3 3 3 4 4 3 3 4 7 3 7 4 4 4 3 4 3 2 2 3 3 3 3 9 3 3 3 3 4 3 4 4 3 3 4 4 4 4 4 4 3 3 3 3 7 4 4 4 8 ? 1 1 2 9 2 2 2 2 2 2 2 2 2 2 1 2 2 2 1 7 2 7 1 1 2 2 2 2 2 2 2 2 2 2 7 2 2 2 2 1 2 1 1 2 2 1 2 2 1 1 1 2 2 2 2 7 1 1 1 9 ? 3 3 3 7 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 7 3 7 3 3 3 3 3 1 3 3 1 1 3 1 7 3 3 3 3 3 1 3 3 1 1 3 3 3 3 3 3 1 1 3 3 7 3 3 3 1 1 o i ; 3 1 4 1 4 1 3 1 3 1 3 1 3 1 3 1 3 1 1 3 1 3 1 1 3 1 1 3 1 1 3 1 1 3 1 1 4 1 1 3 1 1 3 1 1 3 1 1 4 1 1 7 7 1 3 1 1 7 7 ] 4 1 1 4 1 1 3 1 1 3 1 1 3 1 1 1 1 1 3 1 1 3 1 1 1 1 1 1 3 1 1 1 7 7 3 1 3 1 3 1 3 1 4 1 1 1 4 1 4 1 1 1 1 1 4 1 3 1 3 1 4 1 4 1 4 1 1 1 1 1 3 1 3 1 7 7 4 1 4 1 4 1 1 1 3 4 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n I n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n i 1 n [ 1 n 1 1 n 1 1 n 1 1 n 1 1 n 1 1 n 1 1 5 6 ' 1 1 1 2 1 1 2 1 1 2 i : 2 1 1 2 1 1 2 1 1 2 1 1 2 1 1 2 1 1 2 1 1 4 1 1 3 1 1 3 1 1 2 1 1 2 1 1 2 1 1 2 1 1 2 1 1 2 1 1 4 1 1 2 1 1 3 1 1 2 1 1 2 1 1 2 1 1 2 1 1 2 1 1 2 1 1 3 1 1 3 1 1 2 1 1 3 1 1 2 1 1 2 1 1 4 1 1 2 1 1 2 1 1 2 1 1 2 1 1 3 1 1 3 1 4 1 2 1 3 1 3 1 4 1 3 1 2 1 3 1 3 1 3 1 3 1 3 1 3 1 3 1 3 1 2 1 3 1 2 1 1 1 8 9 1 1 1 2 1 2 1 2 1 1 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 1 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 1 1 2 1 2 1 1 1 1 1 1 2 I 1 2 1 2 1 1 2 1 1 2 1 1 2 1 1 2 1 1 2 1 1 2 2 0 n 4 5 3 n 3 4 4 4 3 2 3 5&6 5 5 4 4 6 6 4 1 6 1 4 4 4 n 4 4 n n n n n 1 1 6 6 6 6 4 n 4 4 n n 4 4 5 3 2 1 n n n n n 3 3 3 2 1 n 2 2 2 n 2 2 2 2 2 1 2 2 2 2 2 2 2 2 2 1 2 1 2 2 2 n 2 2 n n n n n 1 1 2 2 2 2 2 n 2 2 n n 2 2 2 2 1 2 n n n n n 2 2 2 2 2 n 1 1 2 n 2 1 1 1 1 n 1 2 2 1 1 1 2 2 1 n 2 n 1 1 1 n 1 1 n n n n n n n 2 2 2 2 1 n 1 1 n n 1 2 2 1 n 2 n n n n n 1 2 1 2 3 n 2 2 3 n 3 2 2 2 2 n 2 3 3 2 2 2 5 5 2 n 5 n 2 2 2 n 2 2 n n n n n n n 5 5 5 5 2 n 3 3 n n 3 4 2 3 n 1 n n n n n 3 3 3 2 4 n n n n n n n n n n n n n 1 1 1 1 1 n 1 1 n n 1 1 1 1 n 1 n n n n n 1 1 1 2 5 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n 120 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY APPENDIX 2.?Continued. Number Species 152. 153. 154. 155. 156. 157. 158. 159. 160. 161. 162. 163. 164. 165. 166. 167. 168. 169. 170. 171. 172. 173. 174. 175. 176. 177. 178. 179. 180. 181. 182. 183. 184. 185. 186. 187. 188. 189. 190. 191. 192. 193. 194. 195. 196. 197. 198. 199. 200. 201. 202. 203. 204. 205. 206. 207. 208. 209. 210. 211. Fusispira Smithville Fm. sp. Hormotoma augustina Hormotoma zelleri Lophospira perangulata Subulitid El Paso Fm. sp. Pagodospira cicelia Plethospira cannonensis Plethospira cassina Seelya ventricosa Lophospira grandis Straparollina pelagica Plethospira? turgida Turritoma acrea Turritoma Cotter Fm. ornate sp. Turritoma cf. T. acrea Hormotoma Setul Fm. sp. Turritoma ?anna Murchisonia callahanensis Ectomaria prisca Hormotoma gracilis Daidia cerithioides Ectomaria pagoda Haplospira ?nereis Hormotoma bellicincta Hormotoma salteri Hormotoma trentonensis Loxonema murrayana Omospira alexandra Omospira laticincta Straparollina circe Straparollina erigione Girvania excavata Murchisonia Pt. Clarence Fm. sp. Rhabdostropha primitiva Spiroecus girvanensis Daidia aff. D. cerithioides Ectomaria cf. E. pagoda Ectomaria cf. E. prisca Ectomaria laticarinata Ectomaria nieszkowskii Hormotoma insignis Holopella regularis Hormotoma centervillensis Hormotoma cingulata Kjerulfonema cancel lata Kjerulfonema quinquecincta Cyrtostropha cor alii Goniostropha cava Hormotoma subplicata Hormotoma monoliniformis Hormotoma attenuata Loxonema? attenuata Macrochilus fenestratus Rhabdostropha grindrodii Loxonema crossmanni Loxonema sinuosa A uriptygma fortior Catazone allevata Catazone argolis Catazone cunea 2 6 n n n n n n n n n n n 1 n 1 1 1 1 1 n 1 1 n n 1 1 1 1 1 n n n n n n n n n 2 7 n 1 1 2 n 2 1 1 1 I 2 2 1 1 2 1 1 1 1 1 2 1 2 1 2 1 n 1 1 n n n n n 2 2 1 1 1 1 1 n 1 1 n n 1 1 2 1 2 1 n n n n n 1 1 1 2 8 n n n 2 n 2 n n n 1 1 1 n n 1 n n n n n 1 n 1 n 1 n n n n n n n n n 1 2 n n n n n n n n n n n n 1 n 1 n n n n n n n n n 2 9 n n n 3 n 3 n n n 2 2 2 n n 2 n n n n n 2 n 2 n 1 n n n n n n n n n 2 2 n n n n n n n n n n n n 1 n 2 n n n n n n n n n 3 0 n n n n n n n n n n n n n n n n n n n n 1 n 1 n 2 n n n n n n n n n 1 1 n n n n n n n n n n n n 1 n 1 n n n n n n n n n 3 1 n n n 1 n 1 n n n 1 1 1 n n 1 n n n n n 1 n 1 n 1 n n n n n n n n n 1 1 n n n n n n n n n n n n 1 n 1 n n n n n n n n n 3 2 n n n 1 n 1 n n n 1 n 1 n n 1 n n n n n 1 n 1 n 1 n n n n n n n n n n 1 n n n n n n n n n n n n 1 n 1 n n n n n n n n n 3 3 n 2 2 1 n 1 1 1 1 3 3 3 1 1 2 2 1 1 1 1 1 1 1 1 1 1 n 1 1 n n n n n 2 1 1 1 1 1 1 n 1 1 n n 1 1 1 1 1 n n n n n n 1 1 1 3 3 4 5 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 2 2 1 n 1 n 1 n 1 n 1 n 1 n 1 n 2 2 1 n 1 n 1 n 1 n 1 n 2 2 2 2 2 2 3 6 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n 2 n n n n n n n 2 n n n n n 2 2 2 3 7 n n n n n n n n n n n n n n 1 1 1 3 3 8 9 n n 1 2 1 2 1 2 n n 1 2 1 2 1 2 1 2 1 2 4 2 1 2 1 2 1 2 1 2 n n 1 1 1 2 1 2 1 1 1 2 1 2 n 2 1 2 1 1 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 1 1 2 1 1 1 1 1 2 1 2 1 1 1 1 1 1 1 1 I 1 1 1 1 2 1 2 1 2 1 2 n n 1 1 1 2 1 2 4 0 n 1 2 1 n 1 1 1 1 2 2 2 2 n n n n n n n 1 n n n 1 1 1 4 1 n 1 1 1 n 1 1 1 1 2 n 2 1 1 2 1 1 1 1 1 n 1 n 1 1 1 n 1 1 n n 1 n n 1 n 1 1 1 1 1 n 1 1 n n 1 1 1 1 1 1 n 1 n n n 1 1 1 4 2 n n n n n n n n n 1 n n n 1 1 1 4 3 n 2 2 2 n 2 2 2 2 2 2 2 3 2&3 2 2 2 2 2 2 2 2 2 2 2 2 n 2 2 2 2 2 n n 2 2 2 2 2 2 2 n 2 2 n n 2 2 2 2 2 2 n 2 n n n 2 2 2 4 4 n n 2 n n n n n 1 n n n 1 1 1 4 5 ? 2 1 2 7 2 1 1 7 2 2 2 2 2 2 4 4 4 4 5 6 7 8 9 0 n 1 1 n 1 1 n 1 1 n n n 1 1 1 1 n n n n 1 1 n 1 n n n 1 1 n 1 1 n n n n n n n n n n n n 1 1 n n n n n n n n n n n n n n n n n n n n n n n n E 1 9 1 8 1 A 1 E 1 9 1 9 1 8 1 9 1 A 1 C 1 7 1 7 1 7 1 7 1 9 1 8 1 9 1 9 1 9 1 A 1 9 1 A 1 A 1 9 1 B 1 B 1 A 1 B 1 9 1 A 1 B 1 B 1 B 1 A 1 A 7 1 9 9 9 6 9 9 7 9 B 9 8 9 9 A B E B 1 A 1 A 1 C 1 2 1 4 1 4 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n 1 n 1 n 1 n 1 n NUMBER 88 121 APPENDIX 2.?Continued. Number Species 152. 153. 154. 155. 156. 157. 158. 159. 160. 161. 162. 163. 164. 165. 166. 167. 168. 169. 170. 171. 172. 173. 174. 175. 176. 177. 178. 179. 180. 181. 182. 183. 184. 185. 186. 187. 188. 189. 190. 191. 192. 193. 194. 195. 196. 197. 198. 199. 200. 201. 202. 203. 204. 205. 206. 207. 208. 209. 210. 211. Fusispira Smithville Fm. sp. Hormotoma augustina Hormotoma zelleri Lophospira perangulata Subulitid El Paso Fm. sp. Pagodospira cicelia Plethospira cannonensis Plethospira cassina Seelya ventricosa Lophospira grandis Straparollina pelagica Plethospira? turgida Turritoma acrea Turritoma Cotter Fm. omate sp. Turritoma cf. T. acrea Hormotoma Setul Fm. sp. Turritoma ?anna Murchisonia callahanensis Ectomaria prisca Hormotoma gracilis Daidia cerithioides Ectomaria pagoda Haplospira ?nereis Hormotoma bellicincta Hormotoma salteri Hormotoma trentonensis Loxonema murrayana Omospira alexandra Omospira laticincta Straparollina circe Straparollina erigione Girvania excavata Murchisonia Pt. Clarence Fm. sp. Rhabdostropha primitiva Spiroecus girvanensis Daidia aff. D. cerithioides Ectomaria cf. E. pagoda Ectomaria cf. E. prisca Ectomaria laticarinata Ectomaria nieszkowskii Hormotoma insignis Holopella regularis Hormotoma centervillensis Hormotoma cingulata Kjerulfonema cancellata Kjerulfonema quinquecincta Cyrtostropha coralli Goniostropha cava Hormotoma subplicata Hormotoma monoliniformis Hormotoma attenuata Loxonema? attenuata Macrochilus fenestratus Rhabdostropha grindrodii Loxonema crossmanni Loxonema sinuosa Auriptygma fortior Catazone allevata Catazone argolis Catazone cunea 5 1 3 3 3 2 3 2 2 2 2 3 3 3 2 2 3 3 2 2 2 2 3 2 3 2 3 2 2 2 2 3 3 2 2 2 3 3 2 2 2 2 3 2 2 3 2 2 2 2 3 2 3 3 2 2 2 2 2 3 1 3 5 2 ? 2 2 3 7 3 2 2 2 3 3 2 3 2 2 2 2 4 3 2 4 3 3 2 3 2 2 2 2 2 2 2 2 2 4 4 3 3 4 3 2 2 2 2 2 2 2 3 3 2 4 4 2 2 2 2 2 3 2 3 5 3 ? 2 2 3 9 3 2 2 2 2 2 2 2 2 2 2 2 1 2 2 2 2 7 2 2 2 2 2 2 2 2 2 2 2 1 2 2 2 1 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 7 2 2 2 5 4 3 1 2 3 3 1 2 2 1 2 3 3 1 1 2 1 2 3 3 3 3 3 3 2 3 2 3 3 3 3 3 3 3 3 3 3 2 2 3 2 1 3 3 1 3 3 3 3 3 2 3 3 3 3 3 3 3 1 1 1 5 5 ? 5 5 3 7 4 5 5 5 2&3 0 6 7 5 4 5 5 4 3 3 1 3 1 3 3&4 3 2 1 1 0 0 2 2 3 2 1 4 4 4 4 5 2 3 5 2 2 3 2 3 4 3 1 0 1 2 2 0 6 7 6 5 6 6 5 5 5 6 5 5 6 5 5 6 5 2 4&5 4 4 4&5 5 5 4&5 7 5 6 4 5 3 6 5 5 6 6 4 5 4 3 7 3 4 5 4 4 6 4 3 6 6 4 4 4 4 6 6 6 4 6 6 6 2 2 2 5 7 7 1 1 1 9 1 1 1 1 1 1 3 2 1 2 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 2 1 1 2 1 1 2 1 1 1 1 2 2 1 1 1 1 1 1 1 2 2 2 5 8 ? 5 6 4 7 4 6 5 5 6 6 4 5 5 5 5 4 5 4 4&5 8 4 6 5 6 4 6 7 6 6 6 7 8 4 7 8 4 4 5 4 5 7 5 6 7 7 5 5 5 4 6 8 7 8 7 7 8 5 4 5 5 9 5 2&3 5 2 5 2 4 4 4 3 4 5 5 4 5 2&3 3 3 3 3 2 3 3 2&3 2&3 2&3 4 1&2 3 4 4 3 4 4 5 2 3 4 2 2 5 4 4 5 4 4 4 4 5 4 1&2 3 1 4 4 4 1 5 4 5 6 0 7 2 2 1 7 1 2 1 1 1 1 2 1 2 2 2 6 1 n 1 2 n n n 1 n n n n 3 n 1 1 1 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n 6 6 2 3 n 1 3 1 3 1 n 1 n 1 n 1 3 1 n 1 n 1 n 1 n 1 3 1 n 1 3 1 3 1 3 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n ! n n n n n n n n n n n n n n n n n n n n n n n n n n n n 6 4 7 1 1 2 ? 2 2 2 2 2 2 2 6 5 n n n 1 n 1 n n n n n n n n n n n 2 n n 3 1 n n n n n n n n n n n n 2 3 3 n n n n n 2 n n n 3 n n n n n n n n n n n n n 6 6 n n n 1 n 1 n n n n n n n n n n n 4 n n 3 2 n n n n n n n n n n n n 3 3 4 n n n n n 3 n n n 4 n n n n n n n n n n n n n 6 7 n n n 1 n 1 n n n n n n n n n n n 2 n n 1 1 n n n n n n n n n n n n 1 1 1 n n n n n 1 n n n 2 n n n n n n n n n n n n n 6 8 n n n 1 n 1 n n n n n n n n n n n 1 n n 1 1 n n n n n n n n n n n n 2 1 1 n n n n n 1 n n n n n n n n n n n n n n n n n 6 9 2 1 2 1 2 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 2 1 2 2 2 1 1 2 2 2 2 2 1 1 1 1 1 2 1 1 2 2 1 2 2 1 2 2 2 2 2 2 2 2 2 2 7 7 0 1 1 1 n 1 1 1 n 1 1 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 1 1 n 1 n 1 n 1 1 1 n 1 1 I 1 1 1 1 n 1 n 1 1 1 1 1 1 1 1 1 1 1 n 1 n 1 n n n 1 1 n n 1 2 n 2 2 n 2 7 7 2 3 1 2 2 2 1 2 3 1 1 2 3 1 2 2 2 1 2 1 2 1 2 1 2 2 3 1 2 1 2 2 2 2 2 1 4 1 3 1 2 1 2 1 3 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 3 1 2 1 3 1 3 1 4 1 3 1 1 1 2 1 2 1 1 1 2 1 1 2 1 I 2 1 1 2 1 1 3 1 1 2 1 1 3 1 1 2 1 1 2 1 2 1 2 1 2 2 2 1 3 1 3 1 3 7 4 2 1 2 n 2 n 1 n n n n 3 n n 1 1 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n 7 5 1 1 1 2 1 2 1 1 1 2 2 1 1 1 1 1 1 2 2 1 2 2 1 1 1 1 1 1 1 1 1 1 1 1 1 2 2 2 2 2 2 2 2 122 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY APPENDIX 2.?Continued. Number Species 0 1 3 4 5 6 8 9 9 9 1 2 9 9 9 3 4 5 9 9 9 9 0 6 7 8 9 0 152. Fusispira Smithville Fm. sp. 153. Hormotoma augustina 154. Hormotoma zelleri 155. Lophospira perangulata 156. Subulitid El Paso Fm. sp. 157. Pagodospira cicelia 158. Plethospira cannonensis 159. Plethospira cassina 160. Seelya ventricosa 161. Lophospira grandis 162. Straparollina pelagica 163. Plethospira? turgida 164. Turritoma acrea 165. Turritoma Cotter Fm. ornate sp. 166. Turritoma cf. T. acrea 167. Hormotoma Setul Fm. sp. 168. Turritoma? anna 169. Murchisonia callahanensis 170. Ectomaria prisca 171. Hormotoma gracilis 172. Daidia cerithioides 173. Ectomaria pagoda 174. Haplospira ?nereis 175. Hormotoma bellicincta 176. Hormotoma salteri 177. Hormotoma trentonensis 178. Loxonema murrayana 179. Omospira alexandra 180. Omospira laticincta 181. Straparollina circe 182. Straparollina erigione 183. Girvania excavata 184. Murchisonia Pt. Clarence Fm. sp. 185. Rhabdostropha primitiva 186. Spiroecus girvanensis 187. Daidia aff. Z). cerithioides 188. Ectomaria cf. ?. pagoda 189. Ectomaria cf. ?. prisca 190. Ectomaria laticarinata 191. Ectomaria nieszkowskii 192. Hormotoma ins ignis 193. Holopella regularis 194. Hormotoma centervillensis 195. Hormotoma cingulata 196. Kjerulfonema cancellata 197. Kjerulfonema quinquecincta 198. Cyrtostropha coralli 199. Goniostropha cava 200. Hormotoma subplicata 201. Hormotoma monoliniformis 202. Hormotoma attenuata 203. Loxonema? attenuata 204. Macrochilus fenestratus 205. Rhabdostropha grindrodii 206. Loxonema crossmanni 207. Loxonema sinuosa 208. Auriptygma fortior 209. Catazone allevata 210. Catazone argolis 211. Catazone cunea 1 1 1 1 1 2 2 n 2 1 2 n 2 n n n n 1 1 1 1 1 1 2 1 2 2 2 3 2 1 2 2 2 1 n 2 n n n n 2 n 2 n n n n 2 n n n n n n 2 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n 1 1 NUMBER 88 123 APPENDIX 2.?Continued. Number Species 152. 153. 154. 155. 156. 157. 158. 159. 160. 161. 162. 163. 164. 165. 166. 167. 168. 169. 170. 171. 172. 173. 174. 175. 176. 177. 178. 179. 180. 181. 182. 183. 184. 185. 186. 187. 188. 189. 190. 191. 192. 193. 194. 195. 196. 197. 198. 199. 200. 201. 202. 203. 204. 205. 206. 207. 208. 209. 210. 211. Fusispira Smithville Fm. sp. Hormotoma augustina Hormotoma zelleri Lophospira perangulata Subulitid El Paso Fm. sp. Pagodospira cicelia Plethospira cannonensis Plethospira cassina Seelya ventricosa Lophospira grandis Straparollina pelagica Plethospira? turgida Turritoma acrea Turritoma Cotter Fm. ornate sp. Turritoma cf. T. acrea Hormotoma Setul Fm. sp. Turritoma ?anna Murchisonia callahanensis Ectomaria prisca Hormotoma gracilis Daidia cerithioides Ectomaria pagoda Haplospira ?nereis Hormotoma bellicincta Hormotoma salteri Hormotoma trentonensis Loxonema murrayana Omospira alexandra Omospira laticincta Straparollina circe Straparollina erigione Girvania excavata Murchisonia Pt. Clarence Fm. sp. Rhabdostropha primitiva Spiroecus girvanensis Daidia aff. D. cerithioides Ectomaria cf. E. pagoda Ectomaria cf. E. prisca Ectomaria laticarinata Ectomaria nieszkowskii Hormotoma insignis Holopella regularis Hormotoma centervillensis Hormotoma cingulala Kjerulfonema cancellata Kjerulfonema quinquecincta Cyrtostropha coralli Goniostropha cava Hormotoma subplicata Hormotoma monoliniformis Hormotoma attenuata Loxonema? attenuata Macrochilus fenestratus Rhabdostropha grindrodii Loxonema crossmanni Loxonema sinuosa A uriptygma fortior Catazone allevata Catazone argolis Catazone cunea 1 1 1 0 0 0 1 2 3 ? ? ? 7 7 7 7 ? 7 7 ? ? ? ? ? ? ? 7 ? ? ? ? 7 7 7 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 7 ? ? ? ? 7 ? ? ? ? ? ? ? ? ? ? ? 7 ? ? 2 3 3 3 2 3 3 3 3 3 3 4 3 3 3 2 3 3 3 3 1 3 1 3 3 3 4 3 3 1 1 1 ? 1 1 1 4 3 3 2 3 4 3 2 4 3 3 2 3 1 3 3 1 1 4 4 1 2 2 2 1 0 4 ? ? 7 ? 7 7 7 7 7 ? ? 1 1 0 5 1 2 2 2 2 2 2 1 I 1 1 1 1 0 0 0 6 1 2 7 2 1 2 1 1 1 1 1 2 2 2 2 2 2 2 2 2 1 2 1 2 2 1 1 1 1 1 1 1 1 1 1 1 2 2 2 2 2 1 2 2 1 1 2 2 2 2 2 2 1 1 1 1 2 1 1 1 7 8 2 2 2 3 3 3 3 3 3 3 1 1 1 1 1 3 1 1 3 3 1 1 1 1 1 2 3 3 1 2 1 3 3 3 1 0 9 ? 2 2 2 2 2 2 2 2 1 1 0 ? ? ? 7 9 ? ? 2 2 1 1 1 ? ? ? 7 ? 7 ? ? 1 ? 2 ? 7 ? 7 ? ? 1 1 7 ? ? 1 ? 1 ? ? ? ? 7 1 7 ? ? 7 7 ? 7 7 ? 7 ? 1 ? 7 ? 7 ? ? ? 1 1 1 1 1 2 I 7 ? ? ? ? ? ? ? 7 7 7 ? 7 ? ? ? ? 7 ? ? ? ? ? ? ? 7 7 ? ] ? 7 7 1 7 1 7 1 7 1 7 1 7 1 7 1 7 7 1 7 ] ? j ? 7 ] ? 7 ? ? ? ? ? ? 7 ? ? 7 7 ? ? ? ? 1 1 1 1 1 1 3 4 5 7 1 ? i 7 1 ? j ? i 7 1 ? i ? i 7 1 7 1 7 1 7 1 ? i 7 1 7 i 7 1 ? i ? i 7 i 7 1 7 1 7 1 7 1 7 1 7 j ? i 7 1 7 i 7 1 7 ] 7 1 7 i 7 1 ? i 7 i 7 1 7 1 7 i ? i 7 j ? j ? i 7 1 ? i 7 1 7 i 7 i 1 1 1 1 6 7 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 2 n . 2 n : 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 1 n 1 n 1 n 1 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 1 1 1 8 9 1 n 1 n ! n n n n n n n > 2 I 2 n n n n n n n n n n n n n n n n 1 n n 2 1 2 1 n n n n n n n n n n n n n n n n n n n n n n n n n n n n 1 n 1 1 2 2 0 1 1 2 1 1 1 1 1 1 1 1 1 1 3 3 3 4 2 2 2 2 2 2 2 2 2 2 1 2 1 1 1 2 1 2 1 2 1 2 1 2 1 2 1 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 3 2 2 2 2 2 2 2 2 2 2 2 2 2 1 2 2 2 2 1 1 3 2 2 2 2 2 2 2 2 2 2 2 2 2 2 1 2 2 2 2 2 2 1 2 3 7 6 6 6 7 6 5 6 6 5 5 6 6 6 6 6 1 6 1 5 1 6 1 6 1 7 1 6 1 7 1 6 1 6 1 6 1 6 1 6 1 6 1 2 1 5 1 6 1 6 1 6 1 6 1 7 1 6 1 6 1 5 1 6 1 6 1 6 1 6 1 6 1 6 6 6 6 6 6 6 6 6 6 6 6 6 5 6 6 1 1 2 2 * 5 1 2 1 2 1 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 1 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 I 2 I 2 I 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 124 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY APPENDIX 2.?Continued. Number Species 152. 153. 154. 155. 156. 157. 158. 159. 160. 161. 162. 163. 164. 165. 166. 167. 168. 169. 170. 171. 172. 173. 174. 175. 176. 177. 178. 179. 180. 181. 182. 183. 184. 185. 186. 187. 188. 189. 190. 191. 192. 193. 194. 195. 196. 197. 198. 199. 200. 201. 202. 203. 204. 205. 206. 207. 208. 209. 210. 211. Fusispira Smithville Fm. sp. Hormotoma augustina Hormotoma zelleri Lophospira perangulata Subulitid El Paso Fm. sp. Pagodospira cicelia Plethospira cannonensis Plethospira cassina Seelya ventricosa Lophospira grandis Straparollina pelagica Plethospira? turgida Turritoma acrea Turritoma Cotter Fm. ornate sp. Turritoma cf. T. acrea Hormotoma Setul Fm. sp. Turritoma ?anna Murchisonia callahanensis Ectomaria prisca Hormotoma gracilis Daidia cerithioides Ectomaria pagoda Haplospira ?nereis Hormotoma bellicincta Hormotoma salteri Hormotoma trentonensis Loxonema murrayana Omospira alexandra Omospira laticincta Straparollina circe Straparollina erigione Girvania excavata Murchisonia Pt. Clarence Fm. sp. Rhabdostropha primitiva Spiroecus girvanensis Daidia aff. D. cerithioides Ectomaria cf. E. pagoda Ectomaria cf. E. prisca Ectomaria laticarinata Ectomaria nieszkowskii Hormotoma insignis Holopella regularis Hormotoma centervillensis Hormotoma cingulata Kjerulfonema cancellata Kjerulfonema quinquecincta Cyrtostropha coralli Goniostropha cava Hormotoma subplicata Hormotoma monoliniformis Hormotoma attenuata Loxonema? attenuata Macrochilus fenestratus Rhabdostropha grindrodii Loxonema crossmanni Loxonema sinuosa Auriptygma fortior Catazone allevata Catazone argolis Catazone cunea 1 2 6 7 7 8 6 8 7 7 7 6 6 6 7 7 7 7 7 8 7 8 8 7 7 5 7 8 7 7 7 7 4 5 7 7 7 7 7 7 7 6 7 7 7 8 7 7 7 8 8 8 7 8 8 8 7 7 7 7 8 8 8 1 1 1 2 2 2 7 2 1 2 1 2 * 1 2 2 2 2 2 2 2 2 2 ' 1 ' 1 3 1 1 1 1 1 2 1 2 2 2 2 2 3 2 2 1 2 1 2 1 I 9 7 1 2 2 2 2 2 2 2 2 I 1 3 0 n n n n n n n n 2 n n n n 3 n n n n n n n n n n n n n n n n n 3 n 2 3 n n n n n n n n n 1 1 n n n n n n 4 1 n n n n 3 n 1 3 1 n n n n n n n n 2 n n n n 2 n n n n n n n n n n n n n n n n n 1 n 1 1 n n n n n n n n n 3 3 n n n n n n 2 2 n n n n 1 n 1 3 2 n n n n n n n n 1 n n n n 1 n n n n n n n n n n n n n n n n n 1 n 1 1 n n n n n n n n n 1 1 n n n n n 11 1 1 n n n n 1 n 1 3 3 2 2 2 2 2 2 2 2 1 1 3 4 n n n n n n n n 2 n n n n 3 n n n n n n n n n n n n n n n n n 1 1 1 1 n n n n n n n n n 2 2 n n n n n n n 2 n n n n 3 n 1 3 5 n n n n n n n n 2 n n n n 2 n n n n n n n n n n n n n n n n n 1 1 1 1 n n n n n n n n n 3 3 n n n n n n n 2 n n n n 1 n 1 3 6 n n n n n n n n 1 n n n n 1 n n n n n n n n n n n n n n n n n 1 1 1 1 n n n n n n n n n 1 1 n n n n n n n 1 n n n n 1 n 1 3 7 n n n n n n n n 1 n n n n 1 n n n n n n n n n n n n n n n n n 1 1 1 1 1 n n n n n n n n n 1 1 n n n n n n n 1 n n n n 1 n 1 1 3 3 i 9 n n n n 2 1 1 1 t 4 3 1 1 3 3 2 1 2 1 2 3 3 2 2 2 2 1 1 3 1 2 1 1 2 1 ? 3 1 2 2 3 3 ? ? 2 2 2 2 2 1 1 1 2 1 2 2 2 2 2 2 1 1 2 2 2 2 ? 2 2 1 2 2 2 1 4 2 ? ? 7 ? ? ? ? ? ? ? ? ? ? 7 ? ? 7 7 7 ? ? ? ? ? ? ? ? ? ? 7 ? 7 ? ? ? ? ? ? 7 7 2 ? 7 ? ? ? 7 7 7 7 7 7 7 7 7 ? 7 ? 7 1 4 3 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n 2 n n n n n n n n n n n n n n 2 n n n n NUMBER 88 125 APPENDIX 2.?Continued. Number Species 1 2 2 2 2 2 2 1 1 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 3 3 3 2 3 7 9 2 3 3 2 2 3 2 3 2 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 2 3 3 3 3 2 2 2 3 3 3 3 3 3 2 2 1 1 2 2 2 3 3 3 4 3 4 3 4 3&4 7 7 0 3 5 1 1 4 4 4 3 4 4 4 3 4 4 4 4 4 4 4 4 4 4 4 2 3&4 3 4 3 1 2 3 4 2 2 2 2 4 4 2 4 1 2 2 2 3 3 3 2 2 2 1 4 4 5 4 4 4 6 9 9 0 5 7 1 1 5 5 6 5 6 6 5 4 5 5 5 5 5 5 5 6 5 5 5 5 4 5 5 5 1 5 5 5 4 2 2 2 2 4 4 4 4 4 2 2 2 2 3 2 4 5 4 5 2 3 3 3 2 2 9 9 3 2 3 2 2 2 2 2 1 3 2 2 2 2 2 2 1 2 2 2 2 1 3 3 1 1 2 2 2 1 1 2 2 1 1 1 6 2 3 3 3 3 2 7 9 3 3 3 3 3 3 1 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 2 1 2 2 3 3 2 3 2 2 3 3 2 1 3 1 1 1 2 3 2 2 1 1 7 2 4 4 3 3 3 9 9 3 3 4 3 3 3 1 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 1 3 3 3 3 2 3 2 2 3 3 3 2 3 2 2 1 1 8 2 1 2 2 2 1 9 9 2 1 1 2 2 2 2 2 2 2 2 2 2 2 2 1 1 1 1 1 1 2 1 2 2 2 2 2 1 2 2 2 1 2 2 2 2 2 2 2 1 2 2 2 2 7 2 2 2 2 2 2 9 3 3 3 3 3 3 7 7 1 3 3 1 1 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 1 1 1 0 1 2 3 1 1 4 1 1 3 1 1 3 1 1 3 1 1 4 1 1 7 7 1 7 7 1 1 1 1 4 1 1 4 1 1 1 1 1 1 1 1 3 1 1 3 1 1 3 1 1 3 1 1 3 1 1 3 1 1 3 1 1 3 1 1 3 1 1 3 1 1 3 1 1 3 1 1 3 1 1 4 1 1 4 1 1 4 1 1 3 1 1 3 1 1 3 1 3 1 3 1 3 1 3 1 4 I 3 1 3 1 3 1 4 1 3 1 3 1 3 1 3 1 3 1 3 1 3 1 4 1 3 1 3 1 3 1 3 1 3 1 3 1 3 1 3 1 3 1 3 1 3 1 1 1 3 4 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 1 n 1 1 n 2 1 n 2 1 n I 1 n [ 1 n 1 1 n 1 1 n 1 1 n 1 1 n 1 1 n 1 1 5 6 3 1 4 1 3 1 4 1 3 1 3 1 3 1 3 1 4 1 3 1 3 1 3 1 3 1 3 1 4 1 3 1 3 1 3 1 3 1 3 1 3 1 3 1 3 1 3 1 3 1 3 1 4 1 4 1 4 1 2 1 2 1 2 1 3 1 3 1 2 1 2 1 4 1 5 1 4 1 4 2 4 1 4 1 5 1 4 1 5 1 3 1 3 1 3 1 3 1 4 1 3 1 4 1 4 1 3 1 3 1 3 1 3 1 3 1 3 1 4 1 1 1 1 7 8 9 1 1 1 1 1 2 1 1 2 1 1 2 1 1 1 1 1 2 1 1 1 1 1 1 1 1 1 1 1 2 1 1 2 1 1 1 1 1 1 1 1 2 1 1 2 1 1 2 1 1 2 1 1 2 1 1 2 1 1 2 1 1 2 1 1 2 1 1 2 1 1 2 1 1 2 1 1 2 1 1 2 1 1 2 1 1 2 1 1 2 1 1 2 1 1 2 1 1 2 1 1 2 1 1 2 1 1 2 1 1 2 1 1 2 1 1 2 3 1 2 1 1 2 1 1 2 1 1 2 1 1 2 1 1 2 1 1 2 1 1 2 1 1 2 1 1 2 1 1 2 1 1 2 1 1 2 1 1 2 1 1 2 1 1 2 1 1 2 1 1 2 1 1 2 1 1 2 1 1 2 2 0 n 2 2 4 n 4 n n n 2 1 n n 3 4&5 3 3 3 5 3 4 3 3 3 4 2 3 4 3 2 2 2 4 3 2 2 3 4 4 4 3 4 4 4 4 4 3 3 2 2 3&4 2 2 1 1 3 3 2 3 3 2 1 n 2 2 2 n 2 n n n 2 1 n n 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 n 2 2 2 n 2 n n n 2 n n n 2 2 2 2 2 2 2 1 3 3 3 3 2 3 3 3 2 2 2 2 2 2 2 3 2 3 3 3 2 2 2 2 1 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 3 n 2 3&4 4 n 2 n n n 1 n n n 3 4 3 3 3 3 4 2 4 4 4 4 3 4 4 5 1 3 1 4 3 1 1 4 5 5 5 3 4 5 5 5 2 3 3 2 5 3 5 5 3 3 3 3 1 5 5 2 4 n 1 1 1 n 1 n n n 1 n n n 2 2 2 5 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n 2 n n n n n n n n 1 2 212. Diplozone crispa 213. Donaldiella declivis 214. Donaldiella morinensis 215. Goniostropha sculpta 216. Loxonema beraultensis 217. Coelocaulus concinnus 218. Macrochilus buliminus 219. Macrochilus cancellatus 220. Macrochilia recticosta 221. Murchisonia paradoxa 222. Sinuspira tenera 223. Stylonema mater 224. Stylonema potens 225. Clathrospira Smithville Fm. sp. 226. Clathrospira ?glindmeyeri 227. Clathrospira elliptica 228. Clathrospira euconica 229. Clathrospira inflata 230. Mourlonia mjoela 231. Clathrospira ?trochiformis 232. Clathrospira convexa 233. Clathrospira conica 234. Clathrospira subconica 235. Eotomaria canalifera 236. Eotomaria dryope 237. Eotomaria labrosa 238. Liospira larvata 239. Paraliospira mundula 240. Eotomaria supracingulata 241. Liospira angustata 242. Liospira decipens 243. Liospira subconcava 244. Euryzone kiari 245. Eotomaria elevata 246. Liospira micula 247'. L iospira progne 248. Paraliospira angulata 249. Brachytomaria baltica 250. Paraliospira aff. /? angulata 251. Paraliospira rugata 252. Eotomaria notablis 253. Lophospira kindlei 254. Brachytomaria papillosa 255. Brachytomaria semele 256. Brachytomaria striata 257. Cataschisma exquisita 258. Clathrospira thraivensis 259. "Bembexia" globosa 260. Eotomaria rupestris 261. Crenilunula Iimata 262. Clathrospira biformis 263. Phanerotrema jugosa 264. Phanerotrema lindstroemi 265. Oriostoma angulifer 266. Stenoloron shelvensis 267. "Seelya" lloydi 268. Ulrichospira similis 269. Eocryptaulina helcinia 270. Conotoma claustrata 271. Crenilunula hallei 126 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY Number Species 212. 213. 214. 215. 216. 217. 218. 219. 220. 221. 222. 223. 224. 225. 226. 227. 228. 229. 230. 231. 232. 233. 234. 235. 236. 237. 238. 239. 240. 241. 242. 243. 244. 245. 246. 247. 248. 249. 250. 251. 252. 253. 254. 255. 256. 257. 258. 259. 260. 261. 262. 263. 264. 265. 266. 267. 268. 269. 270. 271. Diplozone crispa Donaldiella declivis Donaldiella morinensis Goniostropha sculpta Loxonema beraultensis Coelocaulus concinnus Macrochilus buliminus Macrochilus cancellatus Macrochilina recticosta Murchisonia paradoxa Sinuspira tenera Stylonema mater Stylonema potens Clathrospira Smithville Fm. sp. Clathrospira ?glindmeyeri Clathrospira elliptica Clathrospira euconica Clathrospira inflata Mourlonia mjoela Clathrospira Itrochiformis Clathrospira convexa Clathrospira conica Clathrospira subconica Eotomaria canalifera Eotomaria dryope Eotomaria labrosa Liospira larvata Paraliospira mundula Eotomaria supracingulata Liospira angustata Liospira decipens Liospira subconcava Euryzone kiari Eotomaria elevata Liospira micula Liospira progne Paraliospira angulata Brachytomaria baltica Paraliospira aff. P. angulata Paraliospira rugata Eotomaria notablis Lophospira kindlei Brachytomaria papillosa Brachytomaria semele Brachytomaria striata Cataschisma exquisita Clathrospira thraivensis "Bembexid" globosa Eotomaria rupestris Crenilunula limata Clathrospira biformis Phanerotrema jugosa Phanerotrema lindstroemi Oriostoma angulifer Stenoloron shelvensis "Seelya" lloydi Ulrichospira similis Eocryptaulina helcinia Conotoma claustrata Crenilunula hallei 2 6 n 1 1 1 n 1 n n n n n 2 2 2 2 7 n 1 1 1 n 1 n n n 1 2 n n 2 2 8 n n n n n n n n n n 1 n n n n 2 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n 2 9 n n n n n n n n n n 1 n n n n 1 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n APPENDIX 2 ? 3 0 n n n n n n n n n n 1 n n n n 1 n n n n n n n n n n n n n n n n n n n n n n n n ? I n n n n n n n n n n n n n n n n n n n 3 1 n n n n n n n n n n 1 n n n n 1 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n 3 2 n n n n n n n n n n 1 n n n n 1 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n 3 3 1 1 1 3 n 1 n n n n 1 n n 2 3 1 2 1 1 2 1 2 2 2 2 2 2 2 2 2 2 2 2 1 2 2 2 2 2 2 2 2 2 2 2 1 1 2 1 2 2 2 2 1 1 2 2 1 1 2 -Continued. 3 4 2 2 2 2 2 7 2 2 2 2 2 1 7 2 2 2 2 ? 2 1 2 3 5 n 2 n 2 n 2 n n n 2 n n n n n n n n n n n n 1 n n n n n n n n n n n 2 2 n n n n n n n n n 2 n n 2 1 n n 2 2 2 2 n 2 n 1 3 3 6 7 n 1 2 1 n 1 2 1 n n 2 1 n n n n n n 2 1 n n n n n n n 1 n 1 n 1 n 1 n 1 n 1 n I n 1 n 1 2 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 2 1 2 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 2 1 n 1 n 1 2 1 2 1 n 1 n 1 1 1 2 1 2 1 2 1 n 1 2 1 n 1 2 1 3 3 4 4 4 4 4 8 9 0 1 2 1 1 1 1 1 2 3 1 1 1 2 3 4 1 2 1 1 2 1 1 2 1 1 2 1 1 2 n n n n n 1 2 1 1 2 1 1 2 n n n n n 1 2 n n n n n 1 2 n n n n n 1 1 1 1 1 1 n 2 n r n 2 n r 1 2 1 1 1 2 1 1 1 2 1 1 1 2 1 1 1 2 1 1 1 2 1 1 1 2 1 1 1 2 1 1 1 2 1 1 1 2 1 1 2 1 1 1 2 1 1 2 1 1 2 1 1 1 2 1 1 1 2 1 n n 1 1 2 1 1 2 1 i 1 2 1 1 2 1 1 1 1 1 1 1 1 2 1 1 3 1 1 2 1 1 2 1 1 2 1 I 2 1 1 3 1 1 3 1 1 3 1 1 1 1 1 2 1 1 2 1 1 2 1 2 3 1 1 2 1 1 3 1 1 3 1 1 1 1 1 1 1 1 2 1 1 2 1 1 1 1 1 3 1 2 3 1 1 2 1 1 2 I i n n n i n n n 1 3 1 1 2 1 1 3 1 1 3 1 1 3 1 1 3 1 1 3 1 1 2 1 1 3 1 1 2 1 1 4 1 1 4 1 1 4 1 1 4 1 1 4 1 1 4 1 1 3 1 1 3 1 i 1 3 1 1 2 1 1 4 1 1 3 1 1 3 1 1 4 1 1 2 1 1 4 1 1 4 1 1 4 1 1 2 1 1 2 1 1 2 1 1 2 1 1 2 1 1 4 1 1 4 1 I 4 1 1 1 1 1 4 1 1 1 3 1 1 1 3 1 1 1 4 1 1 1 4 1 1 1 2 1 1 1 2 1 1 1 4 1 1 1 1 1 1 2 1 1 4 5 2 2 2 2 2 2 2 2 2 2 2 2 2 2 1 2 2 2 2 2 4 4 4 4 5 6 7 8 9 0 n n n n n n n n n n n n n 1 1 n 1 1 n 1 1 n 1 1 n n n 1 1 1 1 1 1 1 2 1 2 1 2 1 n n n 1 n n n 1 1 2 n 2 2 1 n n n n n n n 2 n n n n 1 1 n n n n n B 9 7 1 7 1 8 1 7 1 D 1 D 1 C 1 8 1 8 1 8 1 8 1 7 1 8 1 7 1 7 1 7 1 8 1 8 1 6 1 7 1 8 1 4 1 7 1 4 1 6 1 7 1 6 1 4 1 4 1 4 1 9 1 A 1 5 1 4 1 9 1 8 1 6 : 9 7 9 B 9 B 4 9 A A . 2 1 8 1 8 1 8 1 8 1 5 1 B 1 B 1 A 1 6 1 2 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n \ n n 1 n 1 n 1 n n n 1 n 1 n 1 n I n 1 n NUMBER 88 127 APPENDIX 2.?Continued. Number Species 5 1 3 2 2 2 2 3 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 3 2 3 2 3 3 3 2 3 3 3 2 2 2 2 2 2 2 2 2 2 3 3 2 1 3 1 1 3 3 2 2 2 1 1 5 2 3 3 3 3 2 2 2 2 2 2 2 2 2 3 3?4 2&3 3 2&3 3 3 1 3 3 3 3 3 3 3 3 3 3 4 2 4 3 3 3 2 2 3 2 2 2 2 2 1 4 3 2 2 3 2 2 3 2 2 3 2 2 2 5 3 2 2 1 273 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 1 2 2 2 2 2 2 2 2 1 2 1 1 2 2 2 2 2 2 2 2 2 2 2 2 1 2 2 2 2 2 2 2 5 4 3 3 1 3 3 1 3 3 3 1 2 3 3 1 1 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 1 3 1 1 1 2 1 1 1 2 3 2 3 1 3 3 1 1 3 3 3 1 2 3 3 1 1 1 5 5 1 3 4 4 3 5 0 0 1 5 3 2 2 6 7 5 3 5 5 6 6 5 5 4 4 4 4 4 4 4 6 6 5 2 6 4 4 5 5 4 4 4 3 4 3 7 2 2 4 8 1 3 3 5 4 3 3 3 7 8 5 6 4 6 5 5 4 2&3 6 6 4 4 3 5 5 2 4 2 3 2 2 3 2 3 3 4 3 4 3 2 3 2 5 5 4 4 5 2 2 2 2 2 4 4 4 4 4 2 4 4 2 2 2 4 4 2 4 4 4 2 2 1 5 7 2 2 2 2 2 2 1 1 1 3 3 3 2 2 3 1 3 3 1 1 1 1 1 3 1 1 3 3 1 1 1 2 2 1 1 2 2 3 5 8 6 7 5 6 7 5 7 7 8 6 6 6 6 4 5 4 5 4 5 4 3 4 4 4 5 4 3 3 3 2 2 2 5 6 1 2 3 5 3 3 3 5 6 5 6 4 5 5 3 2 6 6 5 4&5 4 6 6 3 2 2 5 5 6 9 0 1 4 3 4 4 4 4&5 1 1 3 5 3 n n n n n n n n n n n 5 2 n 5 2 n 4 2 2 2 2 1 4 3 4 4 4 2 3 4 4 4 1 1 1 4 2 1 2 4 4 4 4 1 4 4 4 4 5 3 4 4 3 4 4 4 4 4 3 3 5 2 3 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n 6 2 n n n n n n n n n n n n n n n n n n n n 1 n n n n n n n n n n n n n n 1 n n n n n n n n n n n n n n n n n n n n n n n n n n 5 6 3 4 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 1 2 2 2 2 2 2 2 1 2 2 2 1 2 2 2 1 2 1 2 2 2 1 1 1 1 1 6 5 n n n n n n n n n n n n n n 1 n 1 n n n n n n 2 1 2 2 3 2 1 1 1 1 3 1 1 2 n 2 3 1 2 2 2 1 n 1 3 1 n 3 2 1 2 2 1 1 n n n 6 6 n n n n n n n n n n n n n n 1 n 3 n n n n n n 1 3 1 1 3 1 1 1 1 3 4 1 1 3 n 3 3 3 4 4 4 2 n 4 3 3 n 3 2 2 3 1 2 4 n n n 6 7 n n n n n n n n n n n n n n 1 n 1 n n n n n n 2 2 2 2 1 2 n n n 6 8 n n n n n n n n n n n n n n 1 n 1 n n n n n n 2 n n 2 2 n n n n 6 9 2 2 2 2 2 1 2 2 2 2 2 2 2 2 2 7 0 7 7 1 2 2 2 2 1 2 n 2 n 1 1 1 1 2 1 1 1 1 1 n n n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n ! 1 1 n n n n 1 n 1 n n n n n n n 1 n n n n 1 1 n n n n n n n 3 3 3 2 1 2 2 2 2 1 2 1 2 2 2 2 3 1 3 1 2&3 1 2&3 1 2&3 1 2&3 1 3 1 1 1 3 1 2&3 1 3 1 3 1 2 1 3 1 3 1 3 1 2 1 2 1 3 1 2 1 4 1 3 1 2 1 3 1 3 1 2 2 2 2 3 2 3 1 1 3 I 3 I 2 I 3 1 3 1 3 1 3 1 3 1 1 1 2 1 3 1 2 1 3 1 3 7 7 3 4 n 1 n n n n n n n n n n 1 1 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n I n I n I n I n 1 n 1 n 1 n 1 " 1 n 1 n 1 n 1 TI 1 n 7 5 1 2 2 2 2 1 2 2 2 2 2 2 2 1 2 1 1 1 1 1 1 1&2 1 2 2 1 1 1 212. Diplozone crispa 213. Donaldiella declivis 214. Donaldiella morinensis 215. Goniostropha sculpta 216. Loxonema beraultensis 217. Coelocaulus concinnus 218. Macrochilus buliminus 219. Macrochilus cancellatus 220. Macrochilina recticosta 221. Murchisonia paradoxa 222. Sinuspira tenera 223. Stylonema mater 224. Sty Ion ema potens 225. Clathrospira Smithville Fm. sp. 226. Clathrospira ?glindmeyeri 227. Clathrospira elliptica 228. Clathrospira euconica 229. Clathrospira inflata 230. Mourlonia mjoela 231. Clathrospira ?lrochiformis 232. Clathrospira convexa 233. Clathrospira conica 234. Clathrospira subconica 235. Eotomaria canalifera 236. Eotomaria dry ope 237. Eotomaria labrosa 238. Liospira larvata 239. Paraliospira mundula 240. Eotomaria supracingulata 241. Liospira angustata 242. Liospira decipens 243. Liospira subconcava 244. Euryzone kiari 245. Eotomaria elevata 246. Liospira micula 247. Z, iospira progne 248. Paraliospira angulata 249. Brachytomaria baltica 250. Paraliospira aff. /? angulata 251. Paraliospira rugata 252. Eotomaria notablis 253. Lophospira kindlei 254. Brachytomaria papillosa 255. Brachytomaria semele 256. Brachytomaria striata 257. Cataschisma exquisita 258. Clathrospira thraivensis 259. "Bembexia" globosa 260. Eotomaria rupestris 261. Crenilunula limata 262. Clathrospira biformis 263. Phanerotrema jugosa 264. Phanerotrema lindstroemi 265. Oriostoma angulifer 266. Stenoloron shelvensis 267. "Seelya" lloydi 268. Ulrichospira similis 269. Eocryptaulina helcinia 270. Conotoma claustrata 271. Crenilunula hallei 128 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY Number Species 212. 213. 214. 215. 216. 217. 218. 219. 220. 221. 222. 223. 224. 225. 226. 227. 228. 229. 230. 231. 232. 233. 234. 235. 236. 237. 238. 239. 240. 241. 242. 243. 244. 245. 246. 247. 248. 249. 250. 251. 252. 253. 254. 255. 256. 257. 258. 259. 260. 261. 262. 263. 264. 265. 266. 267. 268. 269. 270. 271. Diplozone crispa Donaldiella declivis Donaldiella morinensis Goniostropha sculpta Loxonema beraultensis Coelocaulus concinnus Macrochilus buliminus Macrochilus cancellatus Macrochilina recticosta Murchisonia paradoxa Sinuspira tenera Stylonema mater Stylonema potens Clathrospira Smithville Fm. sp. Clathrospira ?glindmeyeri Clathrospira elliptica Clathrospira euconica Clathrospira inflata Mourlonia mjoela Clathrospira ?trochiformis Clathrospira convexa Clathrospira conica Clathrospira subconica Eotomaria canalifera Eotomaria dryope Eotomaria labrosa Liospira larvata Paraliospira mundula Eotomaria supracingulata Liospira angustata Liospira decipens Liospira subconcava Euryzone kiari Eotomaria elevata Liospira micula Liospira progne Paraliospira angulata Brachytomaria baltica Paraliospira aff. P. angulata Paraliospira rugata Eotomaria notablis Lophospira kindlei Brachytomaria papillosa Brachytomaria semele Brachytomaria striata Cataschisma exquisita Clathrospira thraivensis "Bembexia" globosa Eotomaria rupestris Crenilunula limata Clathrospira biformis Phanerotrema jugosa Phanerotrema lindstroemi Oriostoma angulifer Stenoloron shelvensis "Seelya" lloydi Ulrichospira similis Eocryptaulina helcinia Conotoma claustrata Crenilunula hallei 7 6 n 2 n n n n n n n 2 n n n n 1 n n n n n n n n n 1 n n 1 n n n n n 2 n i i 1 n 1 1 1 2 2 2 1 n 2 n n n n n n 2 n 2 2 n n n 7 7 n 2 n n n n n n n 3 n n n n 1 n n n n n n n n n 1 n n 1 n n n n n 1 n n 1 n 1 1 1 2 2 2 1 n 1 n n n n n n 2 n 1 1 n n n 7 8 n 2 n n n n n n n 1 n n n n 1 n n n n n n n n n 1 n n 2 n n n n n 1 n n 2 n 2 2 2 1 1 1 1 n 1 n n n n n n n n 1 1 n n n 7 9 n n n n n n i i n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n APPENDIX 2.? 8 0 n n n n n n n n n n n n n n n n n n n n n n n .. n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n 8 1 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n 8 2 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n 8 3 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n -Continued 8 4 n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n n 8 i 5 ( n n 1 n 1 n n n 1 n n n n 1 n n 1 n n 1 I n n n 1 n n n 1 1 1 1 n 1 1 n 1 n n n n n n n 1 n n n n n n n 1 n n n n n n n n 1 1 n n n 8 7 1 2 2 2 ? 2 1 1 9 2 2 ? ? 2 1 2 2 2 2 2 2 2 2 2 2 3 2 3 2 1 3 3 2 2 2 2 2 3 2 3 2 2 3 2 3 1 2 2 2 2 2 1 3 1 3 1 1 1 2 1 2 1 1 1 2 1 2 8 5 8 9 n 2 1 2 1 2 1 7 2 1 n I n n 2 1 2 1 ? j ? 1 2 1 n 1 2 1 3 1 2 1 2 1 3 1 2 1 3 1 3 1 3 1 2 1 2 1 3 1 3 2 3 1 n 3 1 2 2 1 3 2 3 3 : 3 3 : 3 ' 3 3 3 3 3 n 3 3 2 : 2 3 . 3 3 n n 2 2 n 2 2 9 0 n n n n n n n n n n n n n n n n n n n n n n n n n n n 3 n n n n n n n n . 3 n ! 3 ' ? n n n n n n n n > 3 ? I 4 n 1 n 2 4 n 1 n 1 n 1 n 1 n 1 n 9 1 n n n n n n n n n n n n n n n n n n n n n n n n n n n 2 n n n n n n n n 2 n 2 ? n n n n n n n n 2 n 1 n n 2 n n n n n n 9 2 n n n n n n n n n n n n n n n n n n n n n n n n n n n 1 n n n n n n n n 1 n 1 7 n n n n n n n n 1 n 1 n n 1 n n n n n n 9 3 n n n n n n n n n n n n n n n n n n n n n n n n n n n 1 n n n n n n n n 1 n 1 7 n n n n n n n n 1 n 1 n n 2 n n n n n n 9 4 3 2 3 4 3 3 1 I 2 3 2 3 3 4 3 4 4 4 4 4 3 4 4 4 4 4 5 5 5 5 5 5 4 2 5 5 5 3 5 5 4 4 3 4 3 3 3 4 3 5 4 3 3 4 4 3 3 4 5 5 9 5 1 2 2 3 1 2 3 3 1 3 3 2 2 3 3 3 3 3 3 3 2 3 3 3 3 2 2 2 2 2 2 2 2 3 2 2 2 3 2 2 2 3 3 3 3 3 2 2 2 2 2 3 2 1 1 3 3 2 2 2 9 9 6 1 ? 1 2 1 2 1 1 1 2 1 2 1 1 1 1 1 1 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 2 1 1 1 3 1 3 1 3 1 1 1 1 1 1 1 2 1 2 1 1 1 1 1 3 1 2 3 3 2 2 2 2 2 2 2 2 2 2 1 1 1 2 2 2 2 2 2 2 9 9 8 9 2 1 1 I 1 1 1 1 2 1 1 1 1 1 1 1 2 1 1 1 1 1 2 1 2 1 1 1 2 1 1 1 2 1 1 1 1 1 1 1 2 1 2 1 1 1 4 1 2 1 4 1 4 1 3 1 4 1 4 I 4 1 4 1 1 1 1 1 3 1 4 1 2 1 1 1 4 1 2 1 2 1 1 1 1 1 2 2 1 1 2 1 4 I 3 1 3 1 3 1 2 1 3 1 1 1 1 1 2 1 1 1 3 1 0 0 ? ? ? 1 2 NUMBER 88 129 APPENDIX 2.?Continued. Number Species 212. 213. 214. 215. 216. 217. 218. 219. 220. 221. 222. 223. 224. 225. 226. 227. 228. 229. 230. 231. 232. 233. 234. 235. 236. 237. 238. 239. 240. 241. 242. 243. 244. 245. 246. 247. 248. 249. 250. 251. 252. 253. 254. 255. 256. 257. 258. 259. 260. 261. 262. 263. 264. 265. 266. 267. 268. 269. 270. 271. Diplozone crispa Donaldiella declivis Donaldiella morinensis Goniostropha sculpta Loxonema beraultensis Coelocaulus concinnus Macrochilus buliminus Macrochilus cancellatus Macrochilina recticosta Murchisonia paradoxa Sinuspira tenera Stylonema mater Stylonema potens Clathrospira Smithville Fm. sp. Clathrospira ?glindmeyeri Clathrospira elliptica Clathrospira euconica Clathrospira inflata Mourlonia mjoela Clathrospira ?trochiformis Clathrospira convexa Clathrospira conica Clathrospira subconica Eotomaria canalifera Eotomaria dryope Eotomaria labrosa Liospira larvata Paraliospira mundula Eotomaria supracingulata Liospira angustata Liospira decipens Liospira subconcava Euryzone kiari Eotomaria elevata Liospira micula Liospira progne Paraliospira angulata Brachytomaria baltica Paraliospira aff. P. angulata Paraliospira rugata Eotomaria notablis Lophospira kindlei Brachytomaria papillosa Brachytomaria semele Brachytomaria striata Cataschisma exquisita Clathrospira thraivensis "Bembexia" globosa Eotomaria rupestris Crenilunula limata Clathrospira biformis Phanerotrema jugosa Phanerotrema lindstroemi Oriostoma angulifer Stenoloron shelvensis "Seelya" lloydi Ulrichospira similis Eocryptaulina helcinia Conotoma claustrata Crenilunula hallei 1 1 0 C 1 2 9 ] 7 1 ? | ? ] ? ] 7 1 7 1 7 1 7 7 ? ? 7 7 ? ? ? ? ? ? ? ? ? ? ? 7 ? ? ? ? ? 1 0 3 1 3 2 1 4 4 1 1 1 3 3 2 2 2 3 2 3 2 2 2 ? 3 3 3 3 3 3 4 1 3 3 3 3 1 3 3 3 3 4 3 3 3 3 3 1 3 1 1 3 3 1 2 1 3 I 4 I 4 4 1 4 1 3 1 3 1 2 1 2 1 2 1 0 4 7 7 2 ? 7 7 7 ? 2 ? 7 1 0 5 2 7 2 1 2 1 1 1 2 ? 1 9 9 7 1 1 0 C 6 1 1 1 7 1 2 1 2 1 1 1 2 1 2 1 2 1 1 1 7 1 2 1 2 1 2 1 2 1 2 2 ? 1 0 8 3 2 2 2 3 1 1 1 3 2 2 3 3 4 4 4 . 4 I 2 ? ? 4 1 2 ? ? 1 0 9 2 2 7 7 7 9 7 7 7 1 1 0 1 9 9 9 9 7 9 9 9 9 9 9 9 9 9 9 9 7 9 ? 9 ? 7 7 7 7 7 9 7 7 7 7 ? 7 7 7 7 9 7 7 7 ? 7 7 ? 7 ? 7 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 1 2 2 2 2 2 2 2 1 2 2 2 2 2 2 1 2 2 2 2 2 2 2 2 2 2 1 1 2 2 2 1 1 2 7 7 7 7 ? ? ? 7 ? 7 7 ? 7 2^3 2 3 1 3 2 2 2 2 2 2 1 2 3 3 3 7 1 1 2 2 2 1 3 7 1 3 1 2 ? ? 9 3 2 2 1 2 2 2 2 9 7 2 2 3 2 2 2 2 1 2 2 1 1 2 1 1 1 1 1 2 1 1 1 1 4 f 7 1 7 1 7 1 7 i 7 1 7 1 7 ] 7 ] 7 1 7 1 7 ] 7 i 7 j 7 1 7 1 7 ] 7 ] 7 i 7 1 7 i 2 1 7 1 7 1 7 1 7 1 ? i 7 i 7 1 7 1 7 ] 7 i 7 i 1 1 1 1 7 1 7 i 7 ] 7 ] 7 7 1 ? 7 ? 7 1 1 1 7 3 1 7 7 ? ? 9 7 9 9 3 1 1 1 1 6 7 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 TI 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 ri 1 n 1 n 1 n 1 n 2 1 1 n 1 n 1 n 1 n 1 n 1 n 2 1 1 n 2 1 1 n 2 1 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 2 1 1 n 1 1 n 1 1 n 1 1 n 1 2 1 1 2 1 1 1 n 1 7 n 1 1 n 1 1 n 1 1 n 8 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 1 2 2 2 2 2 2 1 1 1 2 1 2 1 1 1 1 2 2 1 2 2 1 1 2 2 1 9 2 2 2 1 1 1 2 9 C n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 2 n 2 2 1 2 1 2 1 1 1 2 1 2 1 2 1 1 1 2 1 2 1 2 1 1 1 3 1 4 1 5 1 5 1 n 1 3 1 5 1 1 1 1 1 5 2 5 1 n ! n n 5 1 n 1 n n n n 1 1 n 3 2 n n 2 2 n n 1 1 2 I 2 1 2 1 I 1 2 1 2 2 2 1 1 2 1 3 3 3 3 3 3 3 4 3 3 3 3 4 3 3 3 4 4 4 3 4 4 4 3 2 3 3 3 3 3 3 3 I 4 I 3 1 4 1 3 1 3 1 2 1 4 1 4 1 3 1 3 1 2 1 2 1 3 1 2 1 3 1 2 2 2 2 1 2 2 2 2 2 1 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 1 1 2 2 3 4 6 1 6 1 6 1 6 1 6 1 6 1 6 1 6 1 6 1 6 1 6 1 6 1 7 1 4 1 4 1 4 1 4 1 4 1 5 1 5 1 5 1 5 1 5 1 5 1 4 1 5 1 4 1 4 1 4 1 4 1 4 1 4 1 5 1 5 1 4 1 4 1 4 1 5 1 4 1 4 1 4 1 5 1 5 1 5 1 5 1 5 1 5 1 5 1 4 1 4 1 5 1 5 1 4 1 ? 1 4 1 5 1 5 1 4 1 5 1 4 1 1 2 5 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 130 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY Number Species 212. 213. 214. 215. 216. 217. 218. 219. 220. 221. 222. 223. 224. 225. 226. 227. 228. 229. 230. 231. 232. 233. 234. 235. 236. 237. 238. 239. 240. 241. 242. 243. 244. 245. 246. 247. 248. 249. 250. 251. 252. 253. 254. 255. 256. 257. 258. 259. 260. 261. 262. 263. 264. 265. 266. 267. 268. 269. 270. 271. Diplozone crispa Donaldiella declivis Donaldiella morinensis Goniostropha sculpta Loxonema beraultensis Coelocaulus concinnus Macrochilus buliminus Macrochilus cancellatus Macrochilina recticosta Murchisonia paradoxa Sinuspira tenera Stylonema mater Stylonema potens Clathrospira Smithville Fm. sp. Clathrospira ?glindmeyeri Clathrospira elliptica Clathrospira euconica Clathrospira inflata Mourlonia mjoela Clathrospira ?trochiformis Clathrospira convexa Clathrospira conica Clathrospira subconica Eotomaria canalifera Eotomaria dryope Eotomaria labrosa Liospira larvata Paraliospira mundula Eotomaria supracingulata Liospira angustata Liospira decipens Liospira subconcava Euryzone kiari Eotomaria elevata Liospira micula Liospira progne Paraliospira angulata Brachytomaria baltica Paraliospira aff. P. angulata Paraliospira rugata Eotomaria notablis Lophospira kindlei Brachytomaria papillosa Brachytomaria semele Brachytomaria striata Cataschisma exquisita Clathrospira thraivensis "Bembexia" globosa Eotomaria rupestris Crenilunula I imat a Clathrospira biformis Phanerotrema jugosa Phanerotrema lindstroemi Oriostoma angulifer Stenoloron shelvensis "Seelya" lloydi Ulrichospira similis Eocryptaulina helcinia Conotoma claustrata Crenilunula hallei 1 2 6 7 8 8 8 7 7 7 7 7 8 8 7 7 6 6 6 6 6 6 6 6 6 6 5 5 5 4 4 4 5 5 5 6 6 5 5 4 6 4 4 5 6 6 6 6 6 6 6 5 5 ? 6 5 ? 5 6 6 5 6 6 APPENDIX 2 1 l l 2 2 2 7 8 9 2 ' 2 2 ? 2 2 2 2 2 2 2 2 3 1 1 1 3 3 3 2 2 1 2 1 2 1 1 2 2 2 2 2 1 2 2 1 ' 2 2 2 2 2 2 2 2 2 > j 2 2 2 2 2 2 2 3 1 2 1 1 2 2 ?Continued 1 3 0 n n n n n n n 4 n n n 3 3 n n n n n n n n n 3 n n n n n n n n n n n 3 n n n n n n n n n n 4 n n n 3 3 3 3 1 n 2 n n 3 2 1 3 1 n n n n n n n 2 n n n 1 1 n n n n n n n n n 1 n n n n n n n n n n n 1 n n n n n n n n n n 1 n n n 3 1 2 2 3 n 2 n n 1 3 1 3 2 n n n n n n n 1 n n n 1 1 n n n n n n n n n 1 n n n n n n n n n n n 1 n n n n n n n n n n 1 n n n 2 1 1 1 2 n 1 n n 1 1 1 3 3 2 2 2 2 2 2 2 2 2 1 3 4 n n n n n n n 4 n n n 3 3 n n 2 n n n n n n 4 n n n n n n n n n n n 4 n n n n n n n n n n 4 n n n 3 n n n n n 2 n n 3 2 1 3 5 n n n n n n n 1 n n n 1 1 n n 1 n n n n n n 1 n n n n n n n n n n n 1 n n n n n n n n n n 1 n n n 3 n n n n n 2 n n 1 3 1 3 6 n n n n n n n 1 n n n 1 1 n n 1 n n n n n n 1 n n n n n n n n n n n 1 n n n n n n n n n n 1 n n n 1 n n n n n 1 n n 1 1 1 1 3 1 1 1 1 3 4 4 7 8 9 0 1 n n n n n n n 1 1 n n n 1 1 1 1 n n 1 1 1 n n n n n n 1 1 n n n n n n n n n n n 1 1 n n n n n n n n n n i : n n n 1 n n n n n 1 n n 1 1 n n 1 n 1 n n 2 1 n 1 2 1 2 1 2 2 2 2 3 2 2 ? 2 2 2 2 3 3 3 3 3 3 3 2 2 2 2 2 2 1 1 3 2 1 1 2 1 1 2 1 2 3 1 2 2 2 2 2 2 1 2 1 ? 2 I 2 I 4 1 2 1 2 1 ? 1 7 1 2 1 2 1 3 1 4 2 ? 2 ? ? ? 7 7 7 7 7 2 7 ? 7 ? 7 7 7 ? ? 7 7 ? 9 ? 7 7 ? ? 7 ? 7 7 7 ? ? 7 7 7 7 7 7 7 7 7 7 ? ? 1 ? ? ? ? ? 1 4 3 n 2 n n n n n n n n 2 n n n n n n n n n n n n n n n n n n n n n n n i i n n n n n n n n n n n n n n n n n n n 1 n n n n n NUMBER 88 131 Number Species 272. 273. 274. 275. 276. 277. 278. 279. 280. 281. 282. 283. 284. 285. 286. 287. 288. 289. 290. 291. 292. 293. 294. 295. Oehlertia gradata Oehlertia scutulata Pleurorima wisbeyensis Promourlonia aff. P. furcata "Longstaffia" "laquetta" Phanerotrema ?occidens Stenoloron aequilatera Oriostoma dispar Murchisonia othemensis "Seelya" ?vitellia Coelozone verna Conotoma glandiformis Euryzone connulastus Globispira prima Oehlertia cancellata Prosolarium procerum Pleurorima migrans Pleurorima aptychia Phanerotrema dolia Spiroraphe bohemica Stenoloron pollens Stenoloron voluta Umbotropis albicans Seelya moydartensis 1 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 3 3 3 3 3 2 1 1 2 2 3 3 3 2 3 3 3 3 2 1 1 1 2 3 3 1 1 2 2 2 2 5 3 2 2 2 1 2 1 1 1 2 2 3 5 4 3 2 2 4 5 5 4 4 4 2 3 2 2 2 4 3 3 3 4&5 4 3 3 3 3 2 2 2 3 APPENDIX 2 - 5 1 1 1 2 2 2 1 3 2 2 1 2 1 1 1 1 1 1 2 1 1 3 2 1 6 1 1 2 2 2 1 2 1 2 1 1 1 1 2 1 1 1 2 1 2 2 2 1 1 7 1 1 2 2 2 1 1 3 2 1 2 1 2 2 1 1 2 2 1 3 1 3 2 2 8 2 2 2 2 2 2 2 3 2 2 2 2 2 2 2 2 2 2 2 2 2 3 2 2 -Continued. 9 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 1 1 0 1 3 1 3 1 3 1 3 1 3 1 3 1 3 1 1 1 3 1 3 1 3 I 3 1 3 1 3 1 3 1 3 1 3 1 3 1 1 1 1 2 3 4 1 1 n 1 I n 1 1 n 1 1 n [ 1 n 2 1 n 1 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 1 n 3 1 2 1 n 3 1 3 1 1 1 3 1 3 1 1 n 1 n 1 n 1 n 1 n 1 5 4 3 3 4 4 4 4 3 3 4 4 3 4 3 4 4 3 3 1 4 1 4 1 4 1 3 1 2 1 3 1 1 1 5 7 1 3 2 2 3 I 2 1 2 1 2 1 1 1 8 9 1 2 1 2 1 2 1 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 0 4 4 3 2 2 2 1 1 3 2 5 2 5 3 3 2 5 5 5 1 1 1 2 5 2 1 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 1 2 2 2 2 2 2 2 1 2 1 1 2 2 1 1 2 2 2 2 2 1 2 3 3 3 2 3 3 5 4 3 3 3 3 4 : 3 2 3 5 2 2 1 3 1 3 1 4 1 3 1 3 1 1 1 2 1 2 2 4 5 1 n 1 n n n n n n n n n n . 1 n n n 2 n n n n n n n n APPENDIX 2.?Continued. Number Species 272. 273. 274. 275. 276. 277. 278. 279. 280. 281. 282. 283. 284. 285. 286. 287. 288. 289. 290. 291. 292. 293. 294. 295. Oehlertia gradata Oehlertia scutulata Pleurorima wisbeyensis Promourlonia aff. P. furcata "Longstaffia" "laquetta" Phanerotrema loccidens Stenoloron aequilatera Oriostoma dispar Murchisonia othemensis "Seelya" ?vitellia Coelozone verna Conotoma glandiformis Euryzone connulastus Globispira prima Oehlertia cancellata Prosolarium procerum Pleurorima migrans Pleurorima aptychia Phanerotrema dolia Spiroraphe bohemica Stenoloron pollens Stenoloron voluta Umbotropis albicans Seelya moydartensis 2 6 2 2 2 7 2 2 2 2 2 8 n n 5 n n n n n n n n n n 5 n n 5 n n n n n n 5 2 9 n n 1 n n n n n n n n n n 1 n n 1 n n n n n n 1 3 0 n n 1 n n n n n n n n n n 1 n n 1 n n n n n n 1 3 1 n n 1 n n n n n n n n n n 1 n n 1 n n n n n n 1 3 2 .. n 1 n n n n n n n n n n 1 n n 1 n n n n n n 1 3 3 2 2 1 2 2 2 2 1 2 n 1 1 1 1 1 2 1 1 1 2 2 1 2 1 3 4 2 2 2 1 1 7 2 2 2 1 2 1 2 2 2 2 2 2 ? 2 2 2 2 2 3 5 2 2 2 n n n 2 2 2 n 2 n 2 2 2 1 2 2 n 2 2 2 2 2 3 6 3 3 2 n n n 2 2 2 n 2 n 2 2 3 2 2 2 n 2 2 2 2 2 3 7 2 2 2 3 8 4 4 n 4 n 3 4 9 0 2 1 2 1 1 1 2 1 2 1 3 1 2 1 n 1 2 1 2 1 2 1 2 1 1 1 1 1 2 1 3 1 1 1 1 1 3 1 2 1 1 1 n 1 1 1 1 1 4 4 4 4 1 2 3 4 1 1 2 1 1 1 2 1 1 1 2 1 1 1 2 1 1 1 2 1 1 1 3 1 1 1 4 1 2 1 4 1 1 1 2 1 1 1 3 1 1 1 3 1 1 1 1 1 1 1 3 1 1 1 2 1 1 1 2 1 1 1 1 1 1 1 2 1 1 1 2 1 1 1 3 n 1 2 4 1 1 1 4 1 2 1 4 1 1 1 4 1 1 1 2 1 4 5 2 2 2 2 2 1 1 4 6 n n n n n n 1 1 n n n n n n n n n n n 1 1 1 n n 4 7 2 2 4 4 8 S 8 1 9 1 4 1 A 1 A 1 9 1 5 1 9 1 9 1 A 1 5 1 5 1 5 1 6 8 1 4 5 A 7 6 8 9 4 5 0 n n n n n n n n n n n n n i i n n n n n n n 1 n I n 1 n 132 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY Number Species 272. 273. 274. 275. 276. 277. 278. 279. 280. 281. 282. 283. 284. 285. 286. 287. 288. 289. 290. 291. 292. 293. 294. 295. Oehlertia gradata Oehlertia scutulata Pleurorima wisbeyensis Promourlonia aff. P. furcata "Longstaffia" "laquetta" Phanerotrema ?occidens Stenoloron aequilatera Oriostoma dispar Murchisonia othemensis "Seelya" Ivitellia Coelozone verna Conotoma glandiformis Euryzone connulastus Globispira prima Oehlertia cancellata Prosolarium procerum Pleurorima migrans Pleurorima aptychia Phanerotrema dolia Spiroraphe bohemica Stenoloron pollens Stenoloron voluta Umbotropis albicans Seelya moydartensis 5 1 2 2 1 3 3 1 1 2 2 1 1 1 1 1 2 1 1 1 1 1 3 3 2 1 5 2 3 2 2 2 2 2 1 2 3 2 1 2 1 2 3 2 2 2 2 1 1 2 2 2 5 3 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 5 4 1 1 1 3 3 3 1 2 3 3 1 1 1 1 1 1 1 1 3 1 2 2 1 1 APPENDIX 2.? 5 5 8 8 7 2 2 3 7 6 5 3 7 7 7 6 4&5 8 7 7 4 6 4 4 5 7 5 6 1 1 2 3 3 4 3 6 4 6 2 2 2 2 2 1 2 2 5 2 4 4 2 2 5 7 3 3 2 1 1 1 3 2 1 1 2 2 3 2 3 3 3 3 1 3 3 1 1 3 5 8 4 5 3 6 6 5 2 3 5 5 2 3 2 4 5 2 3 3 4&5 4 4 4 4 3 -Continued. 5 9 6 6 4 5 5 4 1 3 3 3 2 3 4 4 6 2 5 4 4 1 4 4 2 5 6 0 1 1 2 1 1 1 1 1 1 1 2 1 2 2 1 1 2 2 1 1 1 1 1 2 6 1 n n 1 n n n n n n n 3 n 3 3 n n 1 3 n n n n n 1 6 < 2 : n n n n n n n n n n n n n n n n n n n n n n n n 5 6 3 4 ? ? 1 2 2 2 1 1 1 1 1 1 1 1 ? 1 1 1 2 1 2 2 1 1 6 5 n n n 3 3 2 n n n n n n n n n n n n 3 n 2 2 n n 6 6 n n n 4 4 2 n n n n n n n n n n n n 2 n 1 3 n n 6 7 n n n 2 1 1 n n n n n n n n n n n n 1 n 1 1 n n 6 8 n n n 1 1 1 n n n n n n n n n n n n 1 n 1 1 n n 6 7 9 0 1 n 1 TI 1 n 1 n 1 n 1 1 1 n 1 n 1 n 1 n 1 n 1 n 1 n I n 1 n 1 n 1 n 1 n 2 1 1 ! n 1 n 1 n n [ n 7 7 2 3 2 1 2 2 2 3 2 2 3 2 3 2 2 2 3 3 2 2 3 2 1 1 2 2 7 7 3 4 n n n n n n n n n n n n n n n n n n n n n n n n 7 5 2 2 APPENDIX 2.?Continued. Number Species 272. 273. 274. 275. 276. 277. 278. 279. 280. 281. 282. 283. 284. 285. 286. 287. 288. 289. 290. 291. 292. 293. 294. 295. Oehlertia gradata Oehlertia scutulata Pleurorima wisbeyensis Promourlonia aff. P. furcata "Longstaffia" "laquetta" Phanerotrema ?occidens Stenoloron aequilatera Oriostoma dispar Murchisonia othemensis "Seelya" ?vitellia Coelozone verna Conotoma glandiformis Euryzone connulastus Globispira prima Oehlertia cancellata Prosolarium procerum Pleurorima migrans Pleurorima aptychia Phanerotrema dolia Spiroraphe bohemica Stenoloron pollens Stenoloron voluta Umbotropis albicans Seelya moydartensis 7 6 n n n 2 2 n n n n n n n n n n n n n n n n n n n 7 7 n n n 3 3 n n n n n n n n n n n n n n n n n n n 7 8 n n n n n n n n n n n n n n n n n n n n n n n n 7 9 n n n n n n n n n n n n n n n n n n n n n n n n 8 0 n n n n n n n n n n n n n n n n n n n n n n n n 8 1 n n n n n n n n n n n n n n n n n n n n n n n n 8 2 n n n n n n n n n n n n n n n n n n n n n n n n 8 3 n n n n n n n n n n n n n n n n n n n n n n n n 8 4 n n n n n n n n n n n n n n n n n n n n n n n n 8 8 8 5 6 7 n 1 n 1 1 n n n n ' n n n n n n n n n n n n n n n 2 2 1 2 2 3 1 ? ' 2 2 2 2 2 1 2 2 2 2 1 1 1 7 1 2 8 8 3 3 n 3 3 3 n 7 2 3 1 2 1 n 2 2 2 2 2 n n 7 n 2 8 9 2 2 2 9 0 2 2 n n n n n n n n n n n n n n n n n n n n n n 9 1 1 1 n n n n n n n n n n n n n n n n n n n n n n 9 2 1 1 n n n n n n n n n n n n n n n n n n n n n n 9 3 1 1 n n n n n n n n n n n n n n n n n n n n n n 9 4 3 3 3 4 3 3 5 4 3 3 5 3 5 3 3 6 3 3 3 5 5 4 5 3 9 5 2 2 2 2 3 3 2 2 3 2 2 3 2 2 2 2 2 2 3 2 1 2 2 2 1 9 9 9 9 0 6 7 8 9 0 1 1 2 2 2 1 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 1 1 1 1 1 1 2 1 1 2 1 1 1 1 1 1 1 1 3 1 1 4 1 1 1 1 1 1 1 1 2 1 1 1 1 1 2 1 1 2 1 1 1 1 1 4 1 1 2 1 1 2 1 1 2 1 1 3 1 1 3 1 1 3 1 1 2 1 1 2 1 1 NUMBER 88 133 APPENDIX 2.?Continued. Number Species 272. 273. 274. 275. 276. 277. 278. 279. 280. 281. 282. 283. 284. 285. 286. 287. 288. 289. 290. 291. 292. 293. 294. 295. Oehlertia gradata Oehlertia scutulata Pleurorima wisbeyensis Promourlonia aff. P. furcata "Longstaffia" "laquetta" Phanerotrema ?occidens Stenoloron aequilatera Oriostoma dispar Murchisonia othemensis "Seelya" ?vitellia Coelozone verna Conotoma glandiformis Euryzone connulastus Globispira prima Oehlertia cancellata Prosolarium procerum Pleurorima migrans Pleurorima aptychia Phanerotrema dolia Spiroraphe bohemica Stenoloron pollens Stenoloron voluta Umbotropis albicans Seelya moydartensis 1 1 1 0 0 0 1 2 3 7 ? ? 7 7 ? ? ? 7 7 ? 7 ? ? 7 7 ? ? ? ? ? ? 7 ? 1 1 1 3 3 4 4 4 3 3 4 1 4 3 2 2 4 4 4 2 2 4 4 4 1 1 1 1 0 0 0 0 4 5 6 7 7 1 1 1 7 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 7 1 1 1 1 1 1 1 1 1 ? ? 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 7 2 1 7 1 1 1 2 1 1 1 1 1 1 1 7 1 1 1 1 1 1 1 1 1 0 0 8 9 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 7 i 1 1 * 1 1 ' I 1 ' 1 1 ' 1 1 ' 1 1 ' 1 ? 1 1 1 ) 1 ? 2 > 2 > 2 ? 2 ? 2 ? 2 ? 2 ? 1 ) 1 } 2 t 2 > 1 1 2 ? 2 > 2 ' 2 1 1 7 2 1 1 ' 1 1 1 1 ' 1 1 ' 1 1 1 1 ' 1 1 ' ' 2 > 2 ' 2 ' 2 > 1 > 2 ? 2 1 1 2 2 2 3 3 2 2 0 7 7 2 4 7 3 3 3 4 3 3 3 0 0 7 1 3 1 1 3 2 2 2 2 2 1 1 1 1 2 1 2 1 2 1 2 2 1 1 1 1 1 1 2 1 1 4 2 2 2 2 1 ? ? ? ? 1 ? I ? 2 7 3 1 7 ? ? ? 7 1 7 1 1 1 1 1 1 5 6 1 1 1 1 1 1 2 2 ? 1 1 1 1 1 1 1 1 1 1 2 2 2 1 1 1 1 7 n n n n n n 1 2 n n n n n n n n n n n 1 1 1 n n 1 1 8 2 2 2 2 2 2 1 2 7 2 2 2 2 2 2 2 2 2 1 1 1 2 2 2 1 1 9 n 1 2 2 1 1 n 1 n 1 3 2 2 2 1 2 2 2 n n n 2 2 2 1 2 0 2 2 1 2 1 3 3 5 4 4 3 3 3 2 4 5 1 5 5 1 3 5 5 4 3 3 3 3 5 1 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 1 2 2 2 2 1 2 3 6 5 5 4&5 . 5 5 4 3 5 5 3 6 3 5 1 6 4 5 1 4 1 5 1 4 1 4 1 4 1 3 1 5 1 1 1 2 2 * 5 1 2 1 2 I 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 APPENDIX 2.?Continued. Number Species 272. 273. 274. 275. 276. 277. 278. 279. 280. 281. 282. 283. 284. 285. 286. 287. 288. 289. 290. 291. 292. 293. 294. 295. Oehlertia gradata Oehlertia scutulata Pleurorima wisbeyensis Promourlonia aff. P. furcata "Longstaffia" "laquetta" Phanerotrema ?occidens Stenoloron aequilatera Oriostoma dispar Murchisonia othemensis "Seelya" ?vitellia Coelozone verna Conotoma glandiformis Euryzone connulastus Globispira prima Oehlertia cancellata Prosolarium procerum Pleurorima migrans Pleurorima aptychia Phanerotrema dolia Spiroraphe bohemica Stenoloron pollens Stenoloron voluta Umbotropis albicans Seelya moydartensis 1 2 6 6 6 6 5 6 6 5 3&4 6 6 4 6 5 6 7 6 6 6 7 5 5 5 4 6 1 1 1 2 2 2 7 8 9 2 2 1 3 2 2 1 3 1 2 1 2 1 1 ? 2 1 1 1 1 1 1 1 1 2 2 2 1 1 2 1 1 1 1 1 2 1 2 2 2 2 2 2 1 1 2 1 2 1 3 0 4 4 4 n n 3 n n n n n 3 n 4 4 3 4 4 2 n n 1 n 2 1 3 1 1 1 1 n n 2 n n n n n 1 n 1 2 3 1 1 3 n n 1 n 3 1 3 2 1 1 1 n n 1 n n n n n 1 n 1 1 1 1 1 1 n n 1 n 1 1 3 3 2 2 2 2 1 2 2 2 2 2 2 1 3 4 4 4 4 n n n n n n n n 3 n 4 4 n 4 4 n n n n n 4 1 3 5 1 1 1 n n n n n n n n 1 n 1 2 3 1 1 n n n n n 1 1 3 6 1 1 1 n n n n n n n n 1 n n n n n n 1 1 1 3 : 7 i 1 1 1 1 1 n n n 1 n n n n n 1 n n n n n n 1 1 1 1 3 4 4 1 9 0 1 2 1 2 1 2 1 n 2 1 2 2 I 2 2 2 1 n n 1 1 1 2 2 2 2 3 3 2 3 2 ? 3 3 2 2 2 1 3 3 3 1 3 1 2 I 2 I 1 I 3 1 4 2 ? 7 7 ? 7 7 7 7 ? 7 7 7 ? ? ? ? 7 7 ? 1 1 7 7 ? 1 4 3 n n n n n n n n n i i n n n n n n n n n 1 1 n n n Appendix 3. Stratigraphic Data Species are arranged within clades or paraclades of similar species (e.g., "Hormotomoids" include all high-spired "murchisoniinae"). Stratigraphic "time" scales for each clade or paraclade precede the data giving first appearance (FKA), last appearance (LKA), lower and upper 95% confidence intervals (LB and UB), and number of sampled horizons (H). Biogeographic time scales reflect the numbers of sampling opportunities within the four main Ordovician/Silurian provinces (see Wagner, 1995a, for details). Generic names re? flect the names used prior to the revisions suggested by this study. Early "Archaeogastropods' "Time" Scales Stage/Province Laurentia Toquima- Table Head Baltica Gondwana Dolgellian Early Tremadoc Late Tremadoc Early Arenig Middle Arenig 1-10 11^18 49-64 65-111 Finds and Ranges Species 1. Dirhachopea normalis 2. Dirhachopea subrotunda 3. Schizopea typica 4. Sinuopea sweeti 5. Taeniospira emminencis 6. Ceratopea canadensis 7. Gasconadia putilla 8. Jarlopsis conicus 9. Ophileta supraplana 10. Rhombella umbilicata 11. Prohelicotoma uniangulata 12. Sinuopea basiplanata 13. Taeniospira ?st. clairi 14. Bridgeites ?disjuncta 15. Bridgeites planodorsalis 16. Bridgeites supraconvexa 17. Euconia etna 18. Ceratopea ?laurentina 19. Ceratopeapygmaea 20. Orospira bigranosa Province Laur Laur Laur Laur Laur Laur Laur Laur Laur Laur Laur Laur Laur Laur Laur Laur Laur Laur Laur Laur H 10 8 13 5 8 9 21 16 19 5 7 8 3 4 17 8 8 4 22 10 FKA 11 41 49 49 49 49 65 65 65 LB -4.0 -5.8 -13.4 -33.1 -26.2 12.0 1.0 -3.8 0.0 -66.9 -30.5 -9.4 -133.9 14.5 32.6 20.4 6.2 44.3 56.1 52.0 LKA 10 10 40 40 40 90 60 63 60 63 60 40 40 63 111 90 111 73 111 90 UB 15.0 16.8 54.4 84.1 67.2 119.0 70.0 77.8 71.0 140.9 101.5 60.4 184.9 97.5 127.4 118.6 153.8 93.7 120.2 103.0 134 NUMBER 88 135 1. "EUOMPHALINAES": 1.1. "Ophiletoids" and 1.2. "Macluritoids" "Time" Scales Stage/Province Dolgellian Early Tremadoc Late Tremadoc Early Arenig Middle Arenig Late Arenig Llanvirn Llandeilo Early Caradoc Middle Caradoc Late Caradoc Ashgill Laurentia - 1-21 22-36 37-53 - 54 55-69 70-77 78-90 91-114 115-122 123-125 Toquima- Table Head - - - - - 54-79 80-85 96 100 - - - Baltica - - - - - 55-60 61 62-63 64-65 - 66 67-72 Gondwana - - - - - - - - - - - - Finds and Ranges Species 9. Ophileta supraplana 11. Prohelicotoma uniangulata 21. Macluritella stantoni 22. Teiichispira odenvillensis 23. Teiichispira ?oceana 24. Palliseria robusta 25. Mitrospira longwelli 26. Teiichispira kobayashi 27. Teiichispira sylpha 28. Monitorella auricula 29. Maclurites magna 30. "Eccyliopterus ornatus" 31. Maclurites bigsbyi 32. Maclurina logani 33. Maclurina manitobensis 34. Maclurites sedgewicki 35. Maclurites expansa Maclurites expansa 36. Ophileta complanata 37. Lecanospira compacta 38. Lecanospira nereine 39. Barnesella ?lecanospiroides Barnesella ?lecanospiroides 40. Malayaspira hintzei 41. Malayaspira rugosa 42. Barnesella measuresae 43. Lytospira angelini 44. Lytospira yochelsoni 45. Maclurina ?annulata 46. Rossospira harrisae 47. Ecculiomphalus bucklandi Ecculiomphalus bucklandi 48. Lytospira gerrula 49. Lytospira ?norvegica 50. Ophiletina cf. O. sublaxa Ophiletina cf. O. sublaxa 51. Lytospira subrotunda Province Laur Laur Laur Laur Laur Laur ToqTab ToqTab ToqTab ToqTab Laur Laur Laur Laur Laur ToqTab Laur Bait Laur Laur Laur Laur ToqTab ToqTab ToqTab ToqTab Bait ToqTab ToqTab ToqTab ToqTab Laur ToqTab Bait ToqTab Laur Laur H 19 7 2 8 7 4 5 11 10 3 25 6 6 24 15 3 1 1 8 7 4 2 2 8 10 7 9 2 5 1 3 4 1 3 2 1 2 FKA 1 1 22 40 33 46 60 55 46 60 55 82 82 78 82 90 123 67 1 22 22 42 55 55 54 60 55 60 60 60 90 55 90 62 97 123 102 LB -6.0 -25.6 -295.1 30.5 15.6 51.7 21.8 39.6 26.0 -65.6 47.9 72.5 72.5 71.7 68.7 56.2 undef undef -26.9 12.9 -3.3 -73.3 -89.1 26.4 36.5 38.4 48.0 -1122.0 21.8 undef 36.9 2.1 undef 37.9 -18.3 undef -589.9 LKA 32 32 32 53 53 54 85 89 85 85 96 90 90 114 125 96 125 72 41 32 32 45 59 96 88 85 66 100 85 85 100 77 96 66 100 125 125 UB 39.0 58.6 349.1 62.5 70.4 56.3 123.2 104.4 105.0 210.6 103.2 99.5 99.5 120.3 138.2 129.8 undef undef 68.9 41.1 57.3 160.3 203.2 124.6 105.5 106.6 73.0 1282.0 123.2 undef 153.1 129.9 undef 90.2 215.3 undef 816.9 136 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY I. "EUOMPHALINAES": 1.3.1 "Raphistomatids" "Time" Scales Stage/Province Dolgellian Early Tremadoc Late Tremadoc Early Arenig Middle Arenig Late Arenig Llanvirn Llandeilo Early Caradoc Middle Caradoc Late Caradoc Ashgill Early Llandovery Late Llandovery Early Wenlock Late Wenlock Early Ludlow Late Ludlow Pridoli Laurentia - - - 1 - - 2-19 20-30 31-84 85-103 104-117 118-121 122-123 124-125 126-132 133-135 - - - Toquima- Table Head - - - - 2-5 6-21 22-34 25-40 - 4 1 ^ 2 41-42 41-42 - - - - - - Baltica - - - - 2 3-26 2 7 ^ 6 47-48 49-66 67-73 74-94 95-97 - - - - - - - Gondwana - - - - 2 3-21 22 23-34 35^10 - - - - - - - 136 137 138 Finds and Ranges Species 52. Pararaphistoma lemoni Pararaphistoma lemoni 53. Climacoraphistoma vaginati Climacoraphistoma vaginati 54. Lesueurilla bipatellare 55. Lesueurilla marginalis Lesueurilla marginalis 56. Lesueurilla prima 57. Palaeomphalus giganteus 58. Climacoraphistoma damesi 59. Eccyliopterus alatus 60. Eccyliopterus ?princeps 61. Eccyliopterus regularis 62. Lesueurilla infundibula 63. Eccyliopterus louderbacki 64. Lesueurilla declivis Lesueurilla declivis 65. Pararaphistoma qualteriata Pararaphistoma qualteriata 66. Pararaphistoma schmidti Pararaphistoma schmidti 67. Helicotoma gubanovi 68. Scalites katoi 69. Helicotoma medfraensis 70. Lesueurilla scotica 71. Pachystrophia devexa 72. Raphistoma striata 73. Raphistomina lapicida Raphistomina lapicida 74. Scalites angulatus 75. Holopea ins ignis 76. Eccyliopterus beloitensis Province Laur ToqTab Gond Bait Bait Bait ToqTab Gond ToqTab Bait Bait Bait Bait Bait ToqTab Bait ToqTab Bait ToqTab Bait ToqTab ToqTab ToqTab ToqTab ToqTab Bait Laur ToqTab Laur Laur Laur Laur H 1 8 4 10 8 36 1 4 11 1 6 18 20 9 2 5 2 15 6 8 1 1 14 1 2 22 29 2 7 5 14 5 FKA 1 2 3 5 2 2 36 2 6 5 5 24 5 5 3 5 3 5 3 5 36 6 6 22 22 27 2 6 11 2 31 31 LB undef -20.4 -6.2 -17.0 -30.0 -8.5 undef -16.4 -6.8 undef -41.6 12.0 -4.2 -6.0 -544.8 -12.6 -544.8 -22.0 -37.3 -24.9 undef undef -3.1 undef -352.8 13.5 -1.8 -455.3 -57.1 -40.6 2.3 -48.4 LKA 1 34 6 48 48 96 40 9 34 16 48 73 48 23 21 16 21 94 40 48 40 21 40 34 34 97 30 21 92 30 117 84 UB undef 56.4 15.2 70.0 80.0 106.5 undef 27.4 46.8 undef 94.6 85.0 57.2 34.0 568.8 33.6 568.8 121.0 80.3 77.9 undef undef 49.1 undef 408.8 111.1 33.8 482.3 160.1 72.6 145.7 163.4 NUMBER 88 137 Species 77. Holopea rotunda 78. Pachystrophia contigua 79. Pachystrophia spiralis 80. Raphistomina aperta 81. Raphistominafissurata 82. Eccyliopterus owenanus 83. Holopea ampla 84. Holopea pyrene 85. Holopea symmetrica 86. Raphistoma peracuta 87. Raphistomina rugata 88. Raphistoma tellerensis 89. Sinutropis ?esthetica 90. Pachystrophia gotlandica 91. Lytospira triquestra 92. Euomphalus tubus 93. Lytospira subuloides Province Laur Laur ToqTab Laur Laur Laur Laur Laur Laur Laur Laur Laur Laur Laur Laur Laur Gond H 4 1 5 41 1 13 8 4 5 5 2 2 3 8 6 1 3 FKA 65 31 6 33 65 93 32 85 67 85 93 118 118 122 125 127 136 LB -22.4 undef -31.1 26.0 undef 84.7 -3.9 43.6 29.4 46.4 -743.1 2.7 98.7 114.5 105.9 undef 121.5 LKA 102 32 40 102 84 115 84 102 94 117 121 121 121 132 142 132 138 UB 189.4 undef 77.1 109.0 undef 123.3 112.1 143.4 127.6 130.6 957.1 236.3 140.3 139.5 161.1 undef 142.5 I. "Euomphalinaes": "Helicotomids" "Time" Scales Stage/Province Dolgellian Early Tremadoc Late Tremadoc Early Arenig Middle Arenig Late Arenig Llanvirn Llandeilo Early Caradoc Middle Caradoc Late Caradoc Ashgill Early Llandovery Late Llandovery Early Wenlock Late Wenlock Early Ludlow Late Ludlow Pridoli Laurentia - - 1 1-44 - - - - 45-69 70-83 84-86 87-96 97-103 104-110 111-142 143-158 159-180 181-189 - Toquima- Table Head - - - - - 45-48 45-48 - 49-59 49-59 49-59 - - - - - - - Baltica - - - - - - - 49-50 - - - 51-60 - - - - - - - Gondwana - - - - - - - - - - 159 160-171 172 Finds and Ranges Species 19. Ceratopeapygmaea 94. Ceratopea unguis 95. Boucotspira aff. B. ftmbriata 96. Lophonema peccatonica 97. Polehemia taneyensis 98. Walcottomafrydai 99. Helicotomaplanulata 100. Helicotoma tennesseensis 101. Ophiletina sublaxa 102. Ophiletina angularis 103. Oriostoma bromidensis Province Laur Laur ToqTab Laur Laur ToqTab Laur Laur Laur Laur Laur H 22 16 8 12 15 2 26 11 6 1 1 FKA 2 1 45 2 2 45 45 45 45 70 45 LB -6.2 -11.3 34.8 -14.8 -10.9 -70.3 36.7 34.0 4.7 undef undef LKA 44 44 59 44 44 48 96 69 82 79 46 UB 52.4 56.3 69.2 60.8 56.9 163.3 104.4 80.0 122.3 undef undef 138 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY Species 104. Euomphalopterus ?ordovicius Euomphalopterus ?ordovicius 105. Euomphalopterus aff. E. ordovicius 106. Euomphalopterus cariniferus 107. Palaeomphalus?gradatus 108. Trochomphalus ?dimidiatus 109. Helicotoma blodgetti 110. Helicotoma robinsoni 111. Helicotoma? Girvan sp. 112. Straparollina cf. S. circe 113. Euomphalopterus alatus Euomphalopterus alatus 114. Euomphalopterus frenatus 115. Euomphalopterus praetextus 116. Euomphalopterus subcarinatus 117. Euomphalopterus togatus 118. Euomphalopterus undulans 119. Grantlandispira christei 120. Poleumita discors 121. Pycnomphalus acutus 122. Pycnomphalus obesus Pycnomphalus obesus 123. ?) iscordich ilus dalli 124. Discordichilus mollis 125. Discordichilus kolmodini 126. Poleumita alata 127. Poleumita octavia 128. Poleumita rugosa 129. Pseudophorus profundus 130. Pseudophorus stuxbergi 131. Siluriphorus gotlandicus 132. Siluriphorus undulans 133. Streptotrochus incisus 134. Streptotrochus aff. 5. incisus 135. Streptotrochus lamellosus 136. Streptotrochus lundgreni 137. Streptotrochus? visbeyensis 138. Hystricoceras astraciformis 139. Poleumita granulosa 140. Euomphalus walmstedti 141. Centrifugus planorbis 142. Spinicharybdis wilsoni 143. Turbocheilus immaturum 144. Pseudotectus comes 145. Straparollus bohemicus Province ToqTab Laur ToqTab Bait Bait Bait Laur Laur Laur Laur Laur Gond Laur Laur Laur Laur Laur Laur Laur Laur Laur Gond Laur Laur Laur Laur Laur Laur Laur Laur Laur Laur Laur Laur Laur Laur Laur Laur Laur Laur Laur Laur Gond Gond Gond H 1 2 1 5 7 3 2 1 1 3 35 5 1 1 11 3 2 1 34 9 6 3 2 5 2 3 7 24 1 4 10 4 6 4 2 3 4 2 5 3 11 1 4 1 3 FKA 49 87 49 49 46 46 87 87 87 87 101 159 101 101 97 101 101 97 101 101 101 159 107 127 127 104 128 107 132 107 104 132 107 107 112 112 111 143 143 143 159 181 158 159 159 LB undef -201.3 undef -145.0 33.6 -26.5 -201.3 undef undef 38.7 91.4 122.3 undef undef 57.0 -285.4 14.5 undef 91.2 54.6 16.2 101.0 -1392.2 47.6 -334.3 -161.7 81.5 94.4 undef -10.3 84.5 72.2 68.8 24.2 -781.7 -37.7 37.4 -289.5 126.8 -40.5 149.3 undef 128.1 -244.6 101.0 LKA 59 96 59 180 60 60 96 96 96 96 187 183 103 103 187 180 103 103 189 180 180 170 158 180 142 158 183 180 142 157 142 157 142 142 142 142 142 157 153 180 180 189 170 170 170 UB undef 384.3 undef 374.0 72.5 132.5 384.3 undef undef 144.3 196.6 219.8 undef undef 227.0 566.4 189.5 undef 198.8 226.4 264.8 228.0 1657.2 259.4 603.3 423.7 229.5 192.9 undef 274.3 161.5 216.8 180.2 224.8 1035.7 291.7 215.6 589.5 169.2 363.5 189.7 undef 199.9 575.6 228.0 NUMBER 88 139 II. MURCHISONIINAES": II.3. "Hormotomoids" "Time" Scales Stage/Province Dolgellian Early Tremadoc Late Tremadoc Early Arenig Middle Arenig Late Arenig Llanvirn Llandeilo Early Caradoc Middle Caradoc Late Caradoc Ashgill Llandovery Late Llandovery Early Wenlock Late Wenlock Early Ludlow Late Ludlow Pridoli Laurentia - 1-3 4-12 13-87 - 88-89 90 91 92-134 135-216 217-295 296-308 309-324 325-330 331-338 339-344 345-360 361-364 365-367 Toquima- Table Head - - - - - 88-110 111-122 123-126 127-128 127-128 127-128 - - - - - - - - Baltica - - - - - - - - - - 288-290 291-300 - - - - - - - Gondwana - - - - - - - - - - - - - - - 345 346-357 358-359 Finds and Ranges Species 13. Taeniospira ?st. clairi 146. Hormotoma artemesia 147. Hormotoma confusa 148. Hormotoma?dubia 149. Hormotoma ?simulatrix 150. Ectomaria adelina 151. "Hormotoma" "cassina" 152. Fusispira Smithville Fm. sp. 153. Hormotoma augustina Hormotoma augustina 154. Hormotoma zelleri 155. Lophospira perangulata 156. Subulitid El Paso Fm. sp. 157. Pagodospira cicelia Pagodospira cicelia 158. Plethospira cannonensis 159. Plethospira cassina 160. Seelya ventricosa 161. Lophospira grandis 162. Straparo I Una pelagica 163. Plethospira? turgida 164. Turritoma acrea 165. Turritoma Cotter Fm. ornate sp. 166. Turritoma cf. T. acrea 167. Hormotoma Setul Fm. sp. 168. Turritoma?anna 169. Murchisonia callahanensis 170. Ectomaria prisca 171. Hormotoma gracilis Hormotoma gracilis 172. Daidia cerithioides 173. Ectomaria pagoda Province Laur Laur Laur Laur Laur Laur Laur Laur Laur ToqTab Laur Laur Laur Laur ToqTab Laur Laur Laur Laur Laur Laur Laur Laur Laur ToqTab ToqTab ToqTab Laur Laur Bait Laur Laur H 3 15 8 3 37 6 8 5 7 20 3 85 2 7 7 6 4 1 23 7 5 1 2 3 6 7 4 11 80 3 4 11 FKA 1 4 4 4 4 13 36 36 30 91 20 36 36 36 91 16 20 20 16 13 20 13 13 36 91 91 88 92 91 288 93 93 LB -13.5 -21.2 -53.1 -121.6 -5.2 -107.8 0.6 -40.4 -18.1 84.3 -28.3 25.1 -1463.2 -7.2 61.1 -60.3 -136.4 undef -37.5 -51.7 -80.0 undef -188.8 -215.2 57.1 64.4 -6.3 -3.5 80.1 -923.3 -102.5 -2.0 LKA 3 87 87 29 87 126 87 87 87 122 29 308 87 87 126 87 87 29 312 90 87 15 19 87 122 122 128 308 308 300 177 308 UB 17.5 112.2 144.1 154.6 96.2 246.8 122.4 163.4 135.1 128.7 77.3 318.1 1586.2 130.2 155.9 163.3 243.4 undef 368.8 154.7 187.0 undef 220.8 338.2 155.9 148.6 222.3 403.5 319.3 1314.3 372.5 403.0 140 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY Species 174. Haplospira ?nereis 175. Hormotoma bellicincta 176. Hormotoma salteri 177. Hormotoma trentonensis 178. Loxonema murryana 179. Omospira alexandra 180. Omospira laticincta 181. Straparollina circe 182. Straparollina erigione 183. Girvania excavata 184. Murchisonia Pt. Clarence Fm. sp. 185. Rhabdostropha primitiva 186. Spiroecus gir\'anensis 187. Daidia aff. D. cerithioides 188. Ectomaria cf. ?. pagoda 189. Ectomaria cf. ?. prisca 190. Ectomaria laticarinata 191. Ectomaria nieszkowskii 192. Hormotoma ins ignis 193. Holopella regularis 194. Hormotoma centervillensis 195. Hormotoma cingulata Hormotoma cingulata 196. Kjerulfonema cancellata 197. Kjerulfonema quinquecincta 198. Cyrtos tropha cor alii 199. Goniostropha cava 200. Hormotoma subplicata 201. Hormotoma monoliniformis 202. Hormotoma attenuata 203. Loxonema? attenuata 204. Macrochilus fenestratus 205. Rhabdostropha grindrodii 206. Loxonema crossmanni 207. Loxonema sinuosa 208. Auriptygmafortior 209. Catazone allevata 210. Catazone argolis 211. Catazone cunea 212. Diplozone crispa 213. Donaldiella declivis 214. Donaldiella morinensis 215. Goniostropha sculpta 216. Loxonema beraultensis 217. Coelocaulus concinnus 218. Macroch ilus bulim in us 219. Macrochilus cancel la t us Macrochilus cancellatus 220. Macrochilina recticosta 221. Murchisoniaparadoxa 222. Sinuspira tenera 223. Stylonema mater 224. Stylonema potens Province Laur Laur Laur Laur Laur Laur Laur Laur Laur Laur Laur Laur Laur Laur Laur Laur Laur Bait Bait Laur Laur Laur Gond Laur Laur Laur Laur Laur Laur Laur Laur Laur Laur Laur Laur Gond Gond Gond Gond Laur Gond Laur Gond Gond Gond Laur Laur Gond Gond Laur Gond Gond Gond H 8 57 36 93 6 9 1 3 2 1 2 4 2 1 2 3 1 3 8 4 2 6 I 8 2 2 7 2 2 10 1 1 2 1 4 4 4 2 2 3 6 2 2 3 1 2 3 1 3 1 4 1 3 FKA 93 93 93 93 93 93 93 93 135 288 288 288 288 298 298 298 298 291 288 300 309 313 300 313 311 328 327 325 327 325 330 330 328 328 339 346 346 346 346 345 346 345 346 346 346 345 345 346 346 345 346 346 346 LB 35.2 75.7 69.2 85.2 2.9 43.7 undef -317.6 89.4 undef -317.4 239.7 -317.4 undef 244.9 -19.1 undef 242.7 281.2 159.7 251.3 262.1 undef 294.4 -63.8 -623.4 312.1 36.7 -19.0 307.0 undef undef -623.4 undef 272.3 270.1 270.1 -74.8 -74.8 267.7 286.2 -116.3 -74.8 237.2 undef -116.3 267.7 undef 232.4 undef 270.1 undef 237.2 LKA 177 308 308 287 177 177 134 177 177 295 308 308 308 308 308 308 308 300 297 360 310 360 312 323 323 360 344 334 338 360 338 338 360 332 367 357 357 357 357 360 312 357 357 357 358 360 360 357 359 360 357 357 359 UB 234.8 325.3 331.8 295.0 267.1 226.3 undef 587.6 222.6 undef 913.4 356.3 913.4 undef 361.1 625.1 undef 348.3 303.8 500.3 367.7 410.9 undef 362.6 697.8 1311.4 358.9 622.3 684.0 378.0 undef undef 1311.4 undef 433.7 341.9 341.9 686.8 686.8 437.3 325.8 821.3 686.8 374.8 undef 821.3 437.3 undef 380.6 undef 341.9 undef 374.8 NUMBER 88 141 II. "MURCHISONIINAES": II.4. "Eotomarioids" "Time" Scales Stage/Province Dolgellian Early Tremadoc Late Tremadoc Early Arenig Middle Arenig Late Arenig Llanvim Llandeilo Early Caradoc Middle Caradoc Late Caradoc Ashgill Early Llandovery Late Llandovery Early Wenlock Late Wenlock Early Ludlow Late Ludlow Pridoli Laurentia - - - 1-2 3-4 5 6 7 8-61 62-126 127-174 175-192 193-212 213-219 220-241 242-248 249-263 264-272 273-275 Toquima- Table Head - - - - - 5-8 9-14 15-21 21-28 21-28 21-28 - - - - - - - - Baltica - - 8-17 18-33 34 35-59 60-66 67-71 72-83 - - - - - - - Gondwana - _ - - - - - _ - - - - - 249-256 257-273 274 Finds and Ranges Species 225. Clathrospira Smithville Fm. sp. 226. Clathrospira ?glindmeyeri 227'. Clathrospira elliptica 228. Clathrospira euconica 229. Clathrospira inflata 230. Mourlonia mjoela 231. Clathrospira ?trochiformis 232. Clathrospira convexa 233. Clathrospira conica 234. Clathrospira subconica 235. Eotomaria canalifera 236. Eotomaria dryope 237. Eotomaria labrosa 238. Liospira larvata 239. Paraliospira mundula 240. Eotomaria supracingulata 241. Liospira angustata 242. Liospira decipens 243. Liospira subconcava 244. Euryzone kiari 245. Eotomaria elevata 246. Liospira micula 247. Liospira progne 248. Paraliospira angulata 249. Brachytomaria baltica 250. Paraliospira aff. P. angulata 251. Paraliospira rugata 252. Eotomaria notablis 253. Lophospira kindlei Lophospira kindlei 254. Brachytomaria papillosa Brachytomaria papillosa 255. Brachytomaria semele Province Laur ToqTab Bait Laur Bait Bait Laur Laur Laur Laur Laur Laur Laur Laur Laur Laur Laur Laur Laur Bait Laur Laur Laur Laur Bait Laur Laur Bait Laur Bait Laur Bait Bait H 8 3 45 1 24 9 3 4 17 30 4 15 1 11 7 11 4 12 19 7 1 35 76 6 3 2 3 16 1 1 2 1 4 FKA 1 5 5 5 12 11 7 20 8 8 8 8 8 10 8 20 20 8 10 60 62 102 25 25 69 177 175 35 177 82 177 72 72 LB -58.2 0.0 -1.0 undef 4.9 -24.4 -89.6 -377.9 -40.1 -3.1 -111.6 -22.6 undef -34.0 -83.3 -19.6 -187.0 -31.8 -12.0 41.7 undef 92.0 16.0 -153.1 6.2 -284.3 146.0 15.9 undef undef -284.3 undef -169.5 LKA 87 15 71 6 53 71 26 192 192 92 59 109 9 109 117 109 109 109 109 81 126 192 174 192 81 192 180 81 192 83 192 81 176 UB 146.2 0.0 77.0 undef 60.2 106.4 122.6 589.9 240.1 103.1 178.6 139.6 undef 153.0 208.3 148.6 316.0 148.8 131.0 99.3 undef 202.0 183.4 370.1 143.8 653.3 209.0 101.1 undef undef 653.3 undef 417.5 142 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY Species Brachytomaria semele 256. Brachytomaria striata 257. Cataschisma exquisita 258. Clathrospira thraivensis 259. "Bembexia" globosa 260. Eotomaria rupestris 261. Crenilunula Iimata 262. Clathrospira biformis 263. Phanerotrema jugosa 264. Phanerotrema lindstroemi 265. Oriostoma angulifer 266. Stenoloron shelvensis 267'. "Seelya" lloydi 268. Ulrichospira similis 269. Eocryptaulina helcinia 270. Conotoma claustrata 271. Cre/i ilunula hallei Crenilunula hallei 272. Oehlertia gradata 273. Oehlertia scutulata 274. Pleurorima wisbeyensis 275. Promourlonia aft. P. furcata 276. "Longstaffia" "laquetta" 211. Phanerotrema ?occidens Phanerotrema ?occidens 278. Stenoloron aequilatera 279. Oriostoma dispar 280. Murchisonia othemensis 281. "Seelya" ?vitellia 282. Coelozone verna 283. Conotoma glandiformis 284. Euryzone connulastus 285. Globispira prima 286. Oehlertia cancellata 287. Prosolarium procerum Prosolarium procerum 288. Pleurorima migrans 289. Pleurorima aptychia 290. Phanerotrema dolia 291. Spiroraphe bohemica 292. Stenoloron pollens 293. Stenoloron voluta 294. Umbotropsis albicans 295. Seelya moydartensis Province Laur Laur Laur Laur Laur Bait Laur Laur Laur Laur Laur Laur Gond Laur Laur Laur Laur Gond Laur Laur Laur Laur Laur Laur Gond Laur Laur Laur Laur Gond Laur Gond Gond Laur Gond Laur Gond Gond Laur Gond Gond Laur Gond Laur H 2 1 3 2 1 3 19 3 5 6 4 9 17 2 2 5 11 7 6 4 1 5 1 6 2 9 1 1 2 3 2 4 3 3 1 1 16 1 1 3 2 1 2 4 FKA 177 177 177 177 177 72 210 210 193 210 210 199 210 199 210 213 213 257 213 216 229 213 213 227 227 219 229 227 242 257 249 257 250 249 257 273 250 257 264 257 257 249 257 264 LB 61.7 undef -151.4 -284.3 undef 14.0 198.1 41.0 168.0 176.1 163.0 174.1 193.1 -118.1 -799.1 170.4 190.6 242.1 158.9 156.2 undef 124.8 undef 187.8 -205.5 201.6 undef undef -392.3 170.1 -183.5 215.6 129.3 176.6 undef undef 243.0 undef undef 170.1 -261.9 undef -261.9 243.3 LKA 180 192 244 192 192 83 263 244 209 263 241 241 274 209 244 241 263 274 263 241 241 272 215 263 241 248 241 241 263 274 263 274 274 263 274 275 274 274 272 274 274 263 274 272 UB 295.3 undef 572.4 653.3 undef 141.0 274.9 413.1 234.0 274.9 288.0 265.9 290.9 526.1 1253.1 283.6 285.4 288.9 317.1 300.8 undef 360.2 undef 302.2 673.5 265.4 undef undef 897.3 360.9 695.5 315.4 394.8 335.5 undef undef 281.0 undef undef 360.9 792.9 undef 792.9 292.7 Literature Cited Adrain, J.M., and B.D.E. 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