The Clavatulidae (Gastropoda, Conoidea) of the Miocene Paratethys Sea with considerations on fossil and extant Clavatulidae genera Author Harzhauser, Mathias 0000-0002-4471-6655 mathias.harzhauser@nhm-wien.ac.at Author Landau, Bernard 0000-0002-7768-8494 bernardmlandau@gmail.com Author Janssen, Ronald 0000-0002-4471-6655 mathias.harzhauser@nhm-wien.ac.at text Zootaxa 2022 2022-03-30 5123 1 1 172 http://dx.doi.org/10.11646/zootaxa.5123.1.1 journal article 10.11646/zootaxa.5123.1.1 1175-5326 6399827 036F6B4D-CDCC-4CD7-A914-9A1D8C7A097A Genus Granulatocincta nov. gen. Type species. Pleurotoma granulato-cincta Münster in Goldfuss , 1841; middle Miocene , Langhian (Badenian); Central Paratethys Sea , Austria . Etymology. Referring to the name of the type species, meaning ‘granulose cord’. Diagnosis. Small to large, solid, broad fusiform to buccinoid shells with tripartite early teleoconch sculpture of subsutural and central cords bearing small tubercles, and bifid to trifid suprasutural cord. Entire shell covered by close-set, finely beaded spiral cords or with flattened spiral cords cut by axial growth lines into subquadratic tubercles, siphonal canal short. Description. Small to large, solid, broad fusiform to buccinoid with conical to weakly coronate spire. Protoconch paucispiral. Teleoconch of up to ten whorls. Early teleoconch whorls flat-sided, with tripartite sculpture; finely beaded subsutural cord, beaded mid-whorl cord, and axially elongate, opisthocline beads on suprasutural cord cut by one or two narrow spiral grooves making beads bifid or trifid. Later teleoconch whorls flat-sided or weakly concave; subsutural collar weakly swollen; suprasutural cord weak, largely covered by subsequent whorl. Mid-portion entirely covered by narrow, finely beaded spiral cords. Adsutural cords usually coarsely, beaded ( G. callim , G. contorta , G. sotterii ) although beads overrun by secondary and tertiary spiral cords in some species ( G. pelliscrocodili ). Last whorl 55–70% of total height; subsutural collar beaded or finely coronate. Shoulder obtusely angled by finely tubercular shoulder cord, convex below. Siphonal fasciole short, rounded, twisted. Aperture wide, ovate; outer lip not thickened by varix, smooth or lirate within. Anal sinus wide, moderately deep, asymmetrically U-shaped; apex coinciding with middle row of beads. Siphonal canal short to moderately short, shallowly notched at tip. Columella weakly excavated in upper third, weakly twisted below, smooth. Columellar and parietal callus thickened, sharply delimited, forming relatively broad callus rim, often with pseudumbilical chink. Stratigraphic and geographic range. Central Paratethys Sea: middle Miocene (Badenian to Sarmatian, Serravallian); Austria , Slovakia , the Czech Republic , Hungary , Romania Bulgaria ; Proto-Mediterranean Sea: middle to late Miocene (Serravallian to Tortonian), Colli Torinesi ( Italy ), Karaman Basin ( Turkey ); Pliocene ( Italy , southern Spain ); northeastern Atlantic: middle Miocene (Langhian, Serravallian); France , Pliocene ( Spain ); late Miocene (Tortonian): France . Included species. Central Paratethys Sea: Pleurotoma ( Clavatula ) angelae Hoernes & Auinger, 1891 , Granulatocincta callim nov. sp. , Clavatula contorta Švagrovský, 1958 , Pleurotoma granulato-cincta Münster in Goldfuss, 1841, Pleurotoma ( Clavatula ) nataliae Hoernes & Auinger, 1891 , Pleurotoma schreibersi Hörnes, 1854 , Granulatocincta theoderichi nov. sp. Proto-Mediterranean Sea: middle and late Miocene: Clavatula calcarai Bellardi, 1877 , Clavatula curionii Bellardi, 1877 , Clavatula pellegrinii Simonelli, 1896 , Clavatula pelliscrocodili nov. sp. , Pleurotoma sotterii Michelotti, 1847 , Clavatula turbinata Bellardi, 1877 , Clavatula turgidula Bellardi, 1877 , Clavatula vigolensis Bellardi, 1877 (see Simonelli 1896 ; Bellardi 1877 ; Venzo & Pelosio 1963 ; Ferrero Mortara et al. 1981 for illustrations) [note that some of these species may be mere morphotypes of Granulatocincta sotterii , which is the oldest available name), Clavatula rugata Bellardi, 1877 ; Pliocene: Murex rusticus Brocchi, 1814 . Northeastern Atlantic: late Miocene: Pleurotoma capgrandi Tournouër, 1873 , Pleurotoma obtruta ( Millet, 1865 ) , Pliocene: Clavatula delgadoi Vera-Peláez & Lozano-Francisco, 2001 (see Landau et al . 2020 ). ? North Sea: early/middle Miocene: Clavatula boreointerrupta Kautsky, 1925 . Discussion. The species placed in Granulatocincta have been treated so far as Clavatula by all authors since Hoernes & Auinger (1891) . Separation from Clavatula , as defined herein, is based on the early teleoconch sculpture which is trifid; bifid in Clavatula , and the dense, granulose spiral sculpture covering the entire whorl, which is unknown in any extant Clavatulidae . In Granulatocincta the whorl profile below the shoulder on the last whorl is squatter and more rounded compared to Clavatula due to the comparatively wider periphery and shorter siphonal canal. The tripartite early teleoconch sculpture is reminiscent of that of ‘Clavatula’ mystica Reeve, 1843 but ‘ C .’ mystica lacks the bifid and trifid structure of the suprasutural cord (see Boyer & Hernández 2004: 79 , pl. figs 5, 15). A close relation is also unlikely given the different shell shapes. ‘ C .’ mystica is biconic with a rapidly contracting base and has a marked shoulder, whereas Granulatocincta has a broad evenly rounded base. Moreover, Granulatocincta is characterized by prominent granulose spiral cords, whereas ‘ C .’ mystica has weaker, wider spaced and smooth cords (see Boyer & Hernández 2004 for a detailed description of ‘ C .’ mystica ). ‘Clavatula’ filograna Odhner, 1923 is superficially similar and develops a tripartite early teleoconch sculpture, but its spiral cords bear comparatively wide spaced tubercles at intersections with weak axials instead of densely spaced rows of beads, and its anal sinus is narrower. Pleurotoma capgrandi Tournouër, 1873 , from the middle Miocene of France , seems to represent Granulatocincta ( Peyrot1931 , pl.5, figs 34,37; Glibert 1954 , pl.3,fig.6)based on from the middle Miocene of the northeasternAtlantic, discussed by Peyrot (1931) as closely related with G. granulatocincta are not congeneric with Granulatocincta . ‘ Clavatula ’ occidentalis Peyrot, 1931 has a conical spire without tubercles or spines and its spiral cords lack beads ( Peyrot 1931 , pl. 5, figs 10, 18). Similarly, ‘ Clavatula ’ limbata Peyrot, 1931 has smooth spiral cords ( Peyrot 1931 , pl. 8, figs 76, 77). G. obtruta ( Millet, 1865 ) , from the Tortonian of France , is superficially similar but has a biconic outline, much weaker spiral cords and vague, sinuous axial riblets (see Landau et al. 2020 , pl. 75, figs 1–7). Granulatocincta callim and G. contorta . from the Badenian and Sarmatian of the Central Paratethys Sea, and G. sotterii , G. turgidula and G. curionii , from the Tortonian of Italy form a morphologically closely related group and might represent a monophyletic group (see Bellardi 1877 , pl. 5, figs 17–18, Ferrero Mortara et al. 1981 , pl. 14, figs 2a–b, Montanaro 1937 , pl. 5, figs 2–18, Ruggieri & Davoli 1984 , pl. 4, figs 1, 6, 13–14, 18–19, Davoli 1990 , pl. 8, figs 2–8; Dominici et al. 2020 , fig. 11O). Granulatocincta pelliscrocodili nov. sp. , from the Serravallian of Turkey , is somewhat aberrant due to its reduced beads, which are multiply intersected by spiral sculpture. In the Pliocene the group was represented by G. rustica ( Brocchi, 1814 ) , which was widespread in the Mediterranean and disappeared at the end of the Pliocene ( Scarponi & Della Bella 2004 ; Landau & Harzhauser 2022 ), and G. delgadoi , which is restricted to the Atlantic lower Pliocene of the Guadalquivir Basin ( Vera-Peláez & Lozano-Francisco 2001 ; Landau et al . 2011 ), southwestern Spain . Paleoenvironment. The type species is found in inner neritic paleoenvironments. G. angelae , however, might have dwelled also in deeper habitats. Sarmatian occurrences suggest adaptations to hypersaline environments ( Piller & Harzhauser 2005 ), whereas Serravallian occurrences in Turkey derive from brackish assemblages ( Landau et al. 2013 ).