Clavatula

Harzhauser, Mathias, Landau, Bernard & Janssen, Ronald, 2022, The Clavatulidae (Gastropoda, Conoidea) of the Miocene Paratethys Sea with considerations on fossil and extant Clavatulidae genera, Zootaxa 5123 (1), pp. 1-172 : 43

bimarginata- clade

Members of this clade have bipartite early teleoconch sculpture with a granulose subsutural cord and suprasutural spiral cord of equal strength with indistinct, bifid beads ( Fig 3F , 9E–F ). The moderately concave interspace bears prominent, comma-shaped axial riblets formed by growth lines. Spiral threads form a somewhat cancellate pattern especially in the abapical half of the whorls. Aside from ‘ Clavatula ’ veronicae ( Hoernes & Auinger, 1891) , from the middle Miocene of the Central Paratethys Sea, we place here also ‘ Clavatula ’ neogradata Glibert, 1954 from the Tortonian of France as described by Landau et al. (2020: 84, pl. 74). The bimarginata- clade differs from other bipartite groups: Clavatula (s.s.) in being more slender fusiform and in having the adsutural cords of equal strength; in Clavatula (s.s.) the subsutural cord is weak, smooth, and the suprasutural cord is prominent with large beads. The ‘ Clavatula ’ juliae group is immediately separated by its broader profile and strongly constricted base and long siphonal canal. The ‘ Clavatula ’ camillae group also has a slender fusiform profile, but differs in having a smooth concave mid-portion without axial riblets.

Figures

FIGURE 3. Early teleoconch whorls of extant and fossil Clavatulidae species. A. Clavatula regia (Röding, 1798), NHMW 1683 6/28, West Africa. B. Clavatula rubrifasciata (Reeve, 1845), NHMW 10203, Gorée (Senegal). C. ‘Clavatula’ diadema (Kiener, 1839), NHMW 1884/0010/0023, Gorée (Senegal). D. ‘Clavatula’ mystica (Reeve, 1843), MNHN-IM-2013-59897, Est de l'ile de Gorée (Senegal). E. ‘Clavatula’ bimarginata (Lamarck, 1822), MNHN-IM-2013-43369, Dakar (Senegal). F. ‘Clavatula’ pyramidata (Kiener, 1839), MNHN-IM-2009-24982, Baie de Pointe-Noire (Republique du Congo). G. Clavatula sorini nov. sp., paratype, NHMW 1874/0025/0046, Lăpugiu de Sus (Romania). H. ‘Clavatula’ romana (Defrance, 1826), NHMW 1999/0098/0004, Modra-Kráľová (Slovakia). I. ‘Clavatula’ interrupta (Brocchi, 1814), NHMW 1861/0033/0420, Buituri (Romania). J. ‘Clavatula’ hirmetzli Kovács & Vicián, 2021, NHMW 1868/0001/0069, Baden-Sooss (Austria). K. ‘Clavatula’ irisae nov. sp., NHMW 2021/0126/0001, holotype, Gainfarn (Austria). L. ‘Clavatula’ sophiae (Hoernes & Auinger, 1891), lectotype, NHMW 2021/0127/0001, Enzesfeld (Austria). M. ‘Clavatula’ veronicae (Hoernes &Auinger, 1891), NHMW 1855/0043/0044b, Lăpugiu de Sus (Romania). N. ‘Clavatula’ santhai Kovács & Vicián, 2021, holotype, HNHM, PAL 2021.20.1, Letkés (Hungary).O. ‘Clavatula’ camillae (Hoernes &Auinger, 1891), lectotype, NHMW 1949/0005/0042, Lăpugiu de Sus (Romania). P. Megaclavatula amaliae (Hoernes & Auinger, 1891), lectotype, NHMW 1949/0005/0049, Lăpugiu de Sus (Romania). Q. Megaclavatula grunerti nov. sp., paratype, NHMW 2021/0127/0003, Enzesfeld (Austria). R. Megaclavatula laevigata (Eichwald, 1830), NHMW 1853/0038/0035a, Korytnica (Poland). S. Megaclavatula pilleri nov. sp., holotype, NHMW 1949/0005/0051a, Baden-Sooss (Austria). T. Megaclavatula polonica (Pusch, 1837), NHMW 1862/0001/0324, Bad Vöslau (Austria).U. Granulatocincta angelae (Hoernes &Auinger, 1891), paralectotype, NHMW 1949/0005/0099, Möllersdorf (Austria). V. Granulatocincta granulatocincta (Münster in Goldfuss, 1841), NHMW 1846/0028/0009, Enzesfeld (Austria). W. Granulatocincta nataliae (Hoernes & Auinger, 1891), NHMW 1855/0035/0021, holotype, Belgrade/Rakovica (Serbia). X. Granulatocincta schreibersi (Hörnes, 1854), paralectotype, NHMW1997z0178/1133, Bad Vöslau (Austria).Y. Granulatocincta callim nov. sp., paratype, NHMW 2021/0128/0001, Bad Sauerbrunn (Austria). Z. Granulatocincta contorta (Švagrovský, 1958), holotype, SNMB Z 3076, Kuzmice (Slovakia), Aa. Granulatocincta theoderichi nov. sp., paratype, NHMW 2016/0026/0320b, Ritzing (Austria). Ab. Olegia doderleini (Hörnes, 1854), paralectotype, NHMW 1858/0015/0147, Wiesen (Austria). Ac. Olegia mandici nov. sp., paratype, NHMW 1872/0031/0008a, Bekića vrelo, Čremušnica at Donji Kirin (Croatia). Ae. Olegia dorotheae (Hoernes & Auinger, 1891), paralectotype, NHMW 1949/0005/0040, Kleinebersdorf (Austria). Scale bar is 5 mm for all specimens except for Granulatocincta contorta, Olegia doderleini and O. mandici.

FIGURE 9. Extant Clavatulidae species. A. Clavatula rubrifasciata (Reeve, 1845), Gonçalo Rosa, Atoll Specimen Shells (Senegal). B. Clavatula rubrifasciata (Reeve, 1845), NHMW 10203, Gorée (Senegal). C. ‘Clavatula’ mystica (Reeve, 1843), MNHN-IM-2013-59897, Est de l'ile de Gorée (Senegal). D. ‘Clavatula’ mystica (Reeve, 1843), Gonçalo Rosa, Atoll Specimen Shells (Senegal). E. ‘Clavatula’ bimarginata (Lamarck, 1822), MNHN-IM-2013-43369, Dakar (Senegal). F. ‘Clavatula’ bimarginata (Lamarck, 1822), NHMW 2042, Gorée (Senegal).G. ‘Clavatula’ diadema (Kiener,1839), NHMW 1884/0010/0023, Gorée (Senegal). Specimens C and E have been used for molecular analysis by Nicolas Puillandre (unpublished data).