Granulatocincta granulatocincta

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 : 50-52

Granulatocincta granulatocincta (Münster in Goldfuss, 1841) nov. comb.

Figs 18A, B, C 1 –C 2 , D 1 –D 2 , E 1 –E 3 , F 1 –F 3 , G 1 –G 3 , H 1 –H 3, 3V , 5 , 6

Pleurotoma brachyura Partsch — Hauer 1837: 419 [nomen nudum].

Pleurotoma cataphracta Bast. var. β– Pusch, 1837: 144, pl. 12, fig. 15.

* Pleurotoma granulato-cincta Münster —Münster in Goldfuss 1841: 20, pl. 171, fig. 5.

Pleurotoma granulato-cincta Münst. — Hörnes 1848: 20.

Pleur [otoma]. nodifera m.— Eichwald 1852: 184 [non Pleurotoma nodifera Lamarck, 1822 ].

[ Pleurotoma ] granulatocincta — Naumann 1852: pl. 70, fig. 1.

Pleurotoma granulato-cincta Münst. — Hörnes 1854: 344, pl. 37, figs 14–17.

Pleurotoma granulato-cincta Mü. — Schauroth 1865: 231.

Pleurotoma granulato-cincta — Quenstedt 1884: 618, pl. 201, figs 4–6.

Pleurotoma granulato-cincta Münst. — Handmann 1888: 69, pl. 4, fig. 31.

Pleurotoma (Clavatula) granulato-cincta Münst. — Hoernes & Auinger 1891: 353 (partim), pl. 43, 14a–b, pl. 46, figs 16a–b [non pl. 43, figs 11a–b = Granulatocincta schreibersi ( Hörnes, 1854) ].

[ Pleurotoma ] granulato-cincta Münst. — Rosiwal 1894: 84.

Clavatula granulato-cincta Münst. — Friedberg 1912: 200, pl. 12, figs 8–9.

Clavatula granulato-cincta Münst. — Sieber 1947: 157.

Clavatula granulato-cincta (Münst.) — Sieber 1949: 111.

Clavatula granulato-cincta (Münster) — Csepreghy-Meznerics 1950: 59.

Clavatula granulatocincta (Münst.) — Csepreghy-Meznerics 1953: 20, pl. 1, figs 23–24.

Clavatula nógrádensis n. sp. — Csepreghy-Meznerics 1954: 141, pl. 7, fig. 13.

Clavatula granulato-cincta (Münst.) — Sieber 1953: 188.

Clavatula (Clavatula) granulatocincta granulatocincta Münster — Švagrovský 1958: 13, pl. 3, figs 1–3.

C [lavatula]. (C [lavatula]) granulato-cincta (Münst.) — Sieber 1958a: 157.

Clavatula (C.) granulato-cincta (Münst.) — Sieber 1958b: 150.

Clavatula granulatocincta Hörnes, 1856 — Glibert 1960: 39.

Pleurotoma (Clavatula) granulato-cincta Münst. — Florei 1961: 679, pl. 6, fig. 35.

Clavatula granulatocincta Münster (in Goldfuss), 1843— Strausz 1966: 407, pl. 16, fig. 11.

[ Clavatula ] granulatocincta — Powell 1966: 56.

Clavatula (Clavatula) granulatocincta (Münster in Goldfuss, 1843)— Atanacković 1985: 166, pl. 38, figs 12–13.

Clavatula granulatocincta (Münster) — Bałuk & Radwański 1991; 18, pl. 1, figs 2a–c.

Clavatula (Clavatula) granulatocincta (Münster) — Schultz 1998: 74, pl. 30, fig. 3.

Clavatula (Clavatula) granulatocincta (Münster, 1840) — Mikuž 1998: 76, pl. 5, fig. 1.

Clavatula granulatocincta (Münster, 1843) — Bałuk 2003: 34, pl. 6, figs 1–3.

Clavatula nógrádensis Csepreghy-Meznerics, 1954 — Pálfy et al. 2008: 195.

Clavatula granulatocincta (Münster, 1840) — Mikuž 2009: 33, pl. 11, fig. 145.

Clavatula granulatocincta (Münster in Goldfuss, 1841)— Kovács & Vicián 2021: 142, pl. 1, figs 30–32.

non Clavatula granulato-cincta (Munster) — Peyrot, 1931: 90, pl. 8, figs 76–77 [= ‘ Clavatula’ limbata Peyrot, 1931].

non Clavatula (Clavatula) cf. granulato-cincta (Münster, 1843) — Steininger 1973: 436.

non Clavatula (Clavatula) granulatocincta (Münster in Goldfuss, 1840)— Bošnjak et al. 2021: 228, figs 3I–J [= Granulatocincta schreibersi ( Hörnes, 1854) ].

Type material. Lectotype designated herein: SNSB-BSPG AS VII 2056, SL: 41.0 mm, MD: 17.5 mm, Enzesfeld ( Austria), illustrated in Münster in Goldfuss (1841: 20, pl. 171, fig. 5), stored in the Paleontological Museum of the Bavarian State Collection for Geology and Paleontology, Munich ( Germany), fig. 18B . Paralectotypes: SNSBBSPG AS VII 2057, SL: 39.8 mm , MD: 18.0 mm, Enzesfeld ( Austria), stored in the Paleontological Museum of the Bavarian State Collection for Geology and Paleontology, Munich ( Germany), fig. 18A; SNSB-BSPG AS VII 2058 -2065, 8 spec., Enzesfeld ( Austria) .

Illustrated material. NHMW 2021/0126/0006, SL: 21.7 mm, MD: 9.9 mm, illustrated in Hörnes (1854, pl. 37, figs 17a–b), Gainfarn ( Austria), figs 18C 1 –C 2, 3 V; M.61.4366, SL: 14.0 mm, MD: 7.1 mm, Buda-hegy, Sámsonháza ( Hungary), holotype of Clavatula nógrádensis Csepreghy-Meznerics, 1954 , illustrated in Csepreghy-Meznerics (1954, pl. 7, fig. 13), stored in the Hungarian Natural History Museum, Budapest ( Hungary), figs 18D 1 –D 2; NHMW 1846/0028/0008, SL: 24.7 mm, MD: 11.1 mm, Enzesfeld ( Austria), figs 18E 1 –E 2; NHMW 1846/0028/0009, SL: 34.7 mm, MD: 16.1 mm, Enzesfeld ( Austria), illustrated in Hörnes (1854, pl. 37, figs 16a–b), figs 18F 1 –F 2.

Teratological specimens: NHMW 1855/0045/0486, Gainfarn ( Austria), SL: 77.8 mm, MD: 30.2 mm, illustrated in Hoernes & Auinger (1891, pl. 43, figs 16a–b), figs 18G 1 –G 2; NHMW 1855/0045/0136, Steinebrunn ( Austria), illustrated in Hoernes & Auinger (1891, pl. 43, figs 14a–b), figs 18H 1 –H 2.

Additional studied material. NHMW 1853 /0010/0028, Enzesfeld ( Austria) , illustrated in Hörnes (1854, pl. 37, figs 14a–b); NHMW 1849 /0005/0035 , NHMW 1855 /0045/0139, 17 spec., Enzesfeld ( Austria) ; NHMW A1381 , 14 spec., Enzesfeld ( Austria) ; NHMW A425 , 12 spec., Enzesfeld ( Austria) ; NHMW 1846 /0037/0318, 5 spec., Enzesfeld ( Austria) ; NHMW 1853 /0010/0028, 14 spec., Enzesfeld ( Austria) ; NHMW 1846 /0028/0008, 18 spec., Enzesfeld ( Austria) ; NHMW 1847 /0037/0307, 3 spec., Enzesfeld ( Austria) ; NHMW 1997 z0178/1450, 11 spec., Gainfarn ( Austria) ; NHMW 1997 z0178/1451, 25 spec., Gainfarn ( Austria) ; NHMW 1997 z0178/1455, 3 spec., Gainfarn ( Austria) ; NHMW 1868 /0001/0167, Vienna /Pötzleinsdorf ( Austria) ; NHMW 1997 z0178/1134, 14 spec., Bad Vöslau ( Austria) ; NHMW 1851 /0002/0155, 4 spec., Bad Vöslau ( Austria) ; NHMW 1859 /0027/0149, 3 spec., Möllersdorf ( Austria) ; NHMW 1870 /0054/0420, 3 spec., Steinebrunn ( Austria) ; NHMW A904 , 3 spec., Immendorf ( Austria) ; NHMW 1863 /0015/0497, 1 spec., Forchtenau ( Austria) ; NHMW 1851 /0010/0040, 11 spec., Mikulov (the Czech Republic) ; NHMW 1870 /0054/0114, 4 spec., Mikulov (the Czech Republic) ; NHMW 1863 /0015/0315, 11 spec, Szob ( Hungary) ; NHMW 1867 /0019/0141, 4 spec., Coşteiu de Sus ( Romania) .

Revised description. Shell moderately large to large, solid, broad buccinoid, with weakly gradate and coronate spire; apical angle ~38–40°. Protoconch not preserved. 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 initially by two narrow spiral grooves making it trifid. Abapically further spiral grooves subdivide subsutural cord. On sixth spire whorl sculpture changes abruptly, beads on subsutural cord disappear, the cord becomes swollen forming collar, with small, pointed tubercles developed on the most adapical portion of the collar. Mid-whorl cord remains narrow and strongly beaded; suprasutural cord beads weaken leaving row of poorly defined trifid tubercles; abapical tubercle of trifid group usually largely covered by next whorl. Last whorl 60% of total height. Subsutural collar narrow, finely coronate, with 16–18 small pointed tubercles placed at mid-collar. Subsutural ramp poorly delimited, weakly concave, with strengthened secondary beaded cord running at mid subsutural ramp. Shoulder obtusely and weakly angled by tubercular shoulder cord, moderately constricted at base. Siphonal fasciole short, rounded, twisted; sculpture of four primary spiral cords with whorl profile flat to weakly concave between major cords. Adapical shoulder cord also delimiting periphery bearing numerous small rounded tubercles. Mid-whorl, peribasal and perifasciolar cords weaker with smaller tubercles; whole surface covered in relatively coarsely beaded secondary spirals only slightly weaker than the primaries. Aperture wide, ovate. Outer lip not thickened by varix, with up to twelve lirae deep within; anal sinus wide, moderately deep, asymmetrically U-shaped, with apex at strengthened cord running at mid-ramp. Siphonal canal short, weakly deflected to the left, moderately deeply notched at tip. Columella strongly excavated in upper third, straight below, weakly twisted, smooth. Columellar and parietal callus thickened, sharply delimited, forming relatively broad callus rim.

Discussion. Granulatocincta granulatocincta (Münster in Goldfuss, 1841) is among the most common Paratethyan Clavatulidae species and is characterized by its dense sculpture of beads and granules. This species develops many aberrant specimens and figs 18G 1 –G 2 represent a teratological specimen of unusually large size. A specimen with aberrant sculpture is shown in figs 18H 1 –H 2 and a dwarf specimen is represented by figs 18D 1 –D 2. This last specimen ( Figs 18D 1 –D 2 ) was described by Csepreghy-Meznerics (1954) as Clavatula nogradensis . Its sculpture, however, is identical with that of G. granulatocincta and a separation is unjustified. Bałuk (2003) treated one specimen of G. angelae ( Hoernes & Auinger, 1891, pl. 46, fig. 19) as G. granulatocincta . However, there is no reason to separate this specimen from the other syntypes of G. angelae (see discussion and figures of G. angelae herein). Despite some resemblance in sculpture, both species are clearly separated by the more slender outline of G. angelae , the strongly coronate spire with larger spinous tubercles, the more constricted base, and the more uniformly finer granulose sculpture.

Paleoenvironment. The common occurrence at Gainfarn and Enzesfeld suggests a preference for inner neritic environments with sea grass meadows ( Zuschin et al. 2007).

Distribution in Central Paratethys. Badenian (middle Miocene): North Alpine-Carpathian Foreland Basin: Windpassing, Guntersdorf, Immendorf ( Austria) ( Sieber 1947, 1949), Korytnica Basin: Korytnica ( Poland) ( Bałuk 2003); Vienna Basin: Bad Vöslau, Gainfarn, Enzesfeld, Vienna /Pötzleinsdorf, Steinebrunn, Möllersdorf, ( Austria); Mikulov (the Czech Republic) ( Hoernes & Auinger 1891; Rosiwal 1894; Sieber 1953, Sieber 1958b); Eisenstadt-Sopron Basin: Forchtenau ( Austria) (hoc opus); Pannonian Basin: Letkés, Szob, Sámsonháza ( Hungary) ( Hoernes & Auinger 1891; Csepreghy-Meznerics 1954; Kovács & Vicián 2021); Jazovac ( Bosnia and Herzegovina) ( Atanacković 1985); Zorlenţul Mare Basin ( Romania) ( Florei 1961); Krka Basin: Gorenja Stara vas, Šentjernej ( Slovenia) ( Mikuž 1998, 2009); Făget Basin: Coşteiu de Sus ( Romania) (hoc opus).

Proto-Mediterranean Sea: Glibert (1960: 39) mentions an occurrence from the Tortonian of Montegibbio ( Italy) but this record will need confirmation.

Figures

FIGURE 1. A. Map of central and south–eastern Europe, representing the area that was partly covered by the Central Paratethys Sea. The white insert corresponds to the area that is restored in the palaeogeographic map below. B. Paleogeography of the Middle Miocene Paratethys Sea showing the most important sedimentary basins (NAFB: North Alpine Foreland Basin, E.-Sopr B: Eisenstadt–Sopron Basin).

FIGURE 2. Descriptive terms used in descriptions and measurements to shell morphology, whorl profiles and anal sinus. SL: shell length, MD: maximum diameter, AA: apical angle, LWH: last whorl height, AH: aperture height.AL: aperture length.AW: aperture width.

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 5. Anal sinus morphology of Paratethyan Clavatulidae.

FIGURE 6. Apertures of Paratethyan Clavatulidae (C. = Clavatula, ‘C.’ = ‘Clavatula’, G. = Granulatocincta, M. = Megaclavaltula, O. = Olegia).

FIGURE 18 A. Granulatocincta granulatocincta (Münster in Goldfuss, 1841), paralectotype, SNSB-BSPG AS VII 2057, Enzesfeld (Austria). B. Granulatocincta granulatocincta (Münster in Goldfuss, 1841), lectotype, SNSB-BSPG AS VII 2056, Enzesfeld (Austria). C1–C2. Granulatocincta granulatocincta (Münster in Goldfuss, 1841), NHMW 2021/0126/0006, Gainfarn (Austria). D1–D2. Granulatocincta granulatocincta (Münster in Goldfuss, 1841), 3T; M.61.4366, Buda-hegy, Sámsonháza (Hungary). E1–E3. Granulatocincta granulatocincta (Münster in Goldfuss, 1841), NHMW 1846/0028/0008, Enzesfeld (Austria). F1–F3. Granulatocincta granulatocincta (Münster in Goldfuss, 1841), NHMW 1846/0028/0009, Enzesfeld (Austria). G1–G3. Granulatocincta granulatocincta (Münster in Goldfuss, 1841), NHMW 1855/0045/0486, Gainfarn (Austria). H1–H3. Granulatocincta granulatocincta (Münster in Goldfuss, 1841), NHMW 1855/0045/0136, Steinebrunn (Austria).