Megaclavatula polonica (Pusch, 1837) Harzhauser & Landau & Janssen, 2022

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 : 74-75

Megaclavatula polonica (Pusch, 1837) nov. comb.

Figs 23A 1 –A 3 , B 1 –B 3 , C 1 –C 3 , D 1 –D 3, 3T , 5 , 6

* Pleurotoma tuberculosa Bast. Var. polonica —Pusch 1837: 142, pl. 12, fig. 6.

Pleurotoma asperulata Lam. — Hörnes 1854: 341 (partim), pl. 37, figs 4a–b (only) [non Megaclavatula asperulata ( Lamarck, 1822) ].

Pleurotoma tuberculosa Modification bilineata —Quenstedt 1884: pl. 210, fig. 7

Pleurotoma tuberculosa — Quenstedt 1884: 618, pl. 201, figs 8–9.

Clavatula asperulata Lam. — Friedberg 1912: 188, pl. 12, figs 1–2 [non Megaclavatula asperulata ( Lamarck, 1822) ].

Clavatula polonica Pusch — Friedberg 1938: 142.

Clavatula aff. calcarata Grat. —Friedberg: 1912: 197, pl. 12, fig. 5 [non ‘ Clavatula ’ calcarata ( Grateloup, 1832) ].

Clavatula asperulata Lamarck — Strausz 1966: 405 (partim), pl. 15, fig. 17 [non Megaclavatula asperulata ( Lamarck, 1822) ].

Clavatula asperulata tortonica Peyr. — Csepreghy-Meznerics 1969: 95, pl. 6, fig. 11 [non ‘ Clavatula ’ tortonica Peyrot, 1931 ].

Clavatula asperulata tortonica Peyrot — Csepreghy-Meznerics 1972: 32, pl. 15, fig. 24 [non ‘ Clavatula ’ tortonica Peyrot, 1931 ].

Clavatula polonica (Pusch, 1837) — Bałuk 2003: 36, pl. 1, figs 7–8 [non figs 4–6 = Megaclavatula grunerti nov. sp.], pl. 2, figs 6–8 [non figs 3, 5 = Megaclavatula grunerti nov. sp.].

Clavatula amaliae (R. Hoernes & Auinger, 1891) — Mikuž 2009: 30, pl. 10, fig. 132 [non Megaclavatula amaliae ( Hoernes & Auinger, 1891) ].

Clavatula suturalis rosaliae (Hoernes R. & Auinger M. 1890) — Katona et al. 2011: 9, pl. 1, fig. 8 [non Perrona rosaliae ( Hoernes & Auinger, 1891) ]

Clavatula polonica (Pusch, 1837) — Kovács & Vicián 2021: 142, pl. 2, figs 5–7.

Type material. Neotype: The whereabouts of the syntypes, described by Pusch (1837) from the Badenian (middle Miocene) of Korytnica ( Poland) are unknown and the specimens seem to be lost. Therefore, we designate specimen NHMW 2021 /0138/0001, SL: 64.4 mm, MD: 25.9 mm, from Korytnica ( Poland) as neotype, figs 23A 1 –A 3.

Illustrated material. NHMW 2021/0138/0002, SL: 71.8 mm, MD: 29.8 mm, Korytnica ( Poland), figs 23B 1 –B 3, NHMW 2021/0139/0001, SL: 60.6 mm, MD: 24.3 mm, Baden-Sooss ( Austria), figs 23C 1 –C 3, NHMW 1862/0001/0324, SL: 45.3 mm, MD: 18.2 mm, Bad Vöslau ( Austria), illustrated in Hörnes (1854, pl. 37, figs 4a–b), figs 23D 1 –D 3, 3 T.

Additional studied material. NHMW 1831 /0010/0260, 5 spec., Korytnica ( Poland) ; NHMW 1868 /0001/0068, 1 spec., Baden-Sooss ( Austria) ; NHMW 2021 /0132/0003, 6 spec., Baden-Sooss ( Austria) ; NHMW 1889 /0089/0060, 1 spec., Baden-Sooss ( Austria) ; NHMW 2021 /0132/0004, 5 spec., Baden-Sooss ( Austria) ; NHMW 2013 /0300/0414, 11 spec., Baden-Sooss ( Austria) ; NHMW 1855 /0045/0930, 10 spec., Baden-Sooss ( Austria) ; NHMW 2013 /0078/0427, 8 spec., Baden ( Austria) ; NHMW 2013 /0078/0428, 2 spec., Baden ( Austria) ; 5 spec., NHMW 1997 z0178/1130, Bad Vöslau ( Austria) ; NHMW 1846 /0037/0309, 1 spec., Gainfarn ( Austria) ; NHMW 2021 /0126/0008, 1 spec., Gainfarn ( Austria) ; NHMW 2015 /0389/0483, 2 spec., Enzesfeld ( Austria) ; NHMW 1872 /0031/0017, 3 spec., Kirin ( Croatia) ; NHMW 1851 /0010/0039, 5 spec., Mikulov (the Czech Republic) .

Revised description. Shell large, solid, broad fusiform with high, strongly coronate spire; apical angle ~33– 38°. Protoconch not preserved. Teleoconch of at least ten whorls. Early teleoconch whorls flat-sided, with tripartite spiral sculpture consisting of three beaded cords, subsutural cord narrowest, suprasutural cord widest, the beads made bifid by a spiral groove. Sculpture changing around fifth teleoconch whorl; fine spiral cords cross tubercles on sub- and suprasutural cords that weaken abapically, mid-whorl cord disappears, mid-whorl profile concave, bearing comma-shaped axial riblets. Distinct change in sculpture again on seventh whorl; appearance of broad subsutural collar with long, trigonal, spinous tubercles (10–16 on last whorl), somewhat laterally compressed and upturned, placed a short distance below the suture. Suprasutural row of tubercles almost completely covered by subsutural collar of subsequent whorl. Suture narrowly incised, weakly undulating. Last whorl ~60% of total height; broad, poorly delimited, strongly spinous subsutural collar. Distinctly concave subsutural ramp delimited by narrow, weakly angled shoulder cord. Double peribasal cord set a short distance below so that shoulder and peribasal cords all placed at mid-whorl, all three cords bearing small tubercles with a few weak irregular cords between primaries mid-whorl and over base and siphonal fasciole. Base strongly constricted, concave. Siphonal fasciole moderately short, rounded, twisted, often with prominent growth increments. Prominent, close-spaced axial growth lines give surface slightly scabrose appearance. Aperture moderately wide, ovate. Outer lip not thickened, smooth within. Anal sinus wide and moderately deep, asymmetrically V-shaped, with apex below subsutural collar in deepest part of subsutural concavity. Siphonal canal moderately long, straight to slightly recurved, shallowly notched at tip. Columella strongly to moderately excavated in upper third, straight below, slightly twisted at fasciole, smooth. Columellar and parietal callus moderately thickened, adherent, forming broad callus rim.

Discussion. This species is characterized by its comparatively stout outline and abrupt change of sculpture from a conical spire with beaded spiral cords to a strongly coronate shell with spinose tubercles. Therefore, it is surprising that Bałuk (2003) mixed this species with Megaclavatula grunerti nov. sp., which has a reduced sculpture and a slender fusiform outline. Csepreghy-Meznerics (1969, 1972) identified the Paratethyan specimens from Hungary as ‘ Clavatula ’ asperulata tortonica ( Peyrot, 1931) , which was originally described from the Langhian of Saubrigues in France.Although the French species develops a comparable sculpture of prominent, spiny tubercles, it differs clearly in its slender shape and the presence of spiral sculpture (see Peyrot 1931, pl. 5, figs 19, 24, 30–31). For comparison with Pleurotoma aculeatum Eichwald, 1830 see Megaclavatula laevigata ( Eichwald, 1830) . ‘ Clavatula ’ ernesti ( Toula, 1901) , from the Serravallian of Turkey, is a morphologically very close species with comparably spiny sculpture. The Turkish species, however, differs from M. polonica in its strictly flat-sided whorls, the prominent spiral cords and the higher spire (see Toula 1901: 261, pl. 8, fig. 11; Landau et al. 2013: 283, pl. 47, figs 1–3). Moreover, ‘ Clavatula ’ ernesti has a completely different early teleoconch with axial riblets crossed by spiral cords ( Landau et al. 2013, pl. 73, figs 6–7).

Paleoenvironment. The frequent occurrence in the Baden Formation in the Vienna Basin suggests middle to outer neritic environments with up to 210 m water depth ( Kranner et al. 2021). Occurences at Várpalota and Letkés ( Hungary), in contrast, point to inner neritic environments ( Harzhauser et al. 2002; Kovács & Vicián 2013).

Distribution in Central Paratethys. Badenian (middle Miocene): Korytnica Basin: Korytnica ( Poland) ( Bałuk 2003); Vienna Basin: Bad Vöslau, Baden-Sooss, Gainfarn, Enzesfeld ( Austria) (hoc opus), Mikulov (the Czech Republic) (hoc opus); Pannonian Basin: Letkés, Várpalota, Várpalota (Faller Street) ( Hungary) ( Strausz 1966; Katona et al. 2011; Kovács & Vicián 2021), Kirin ( Croatia) (hoc opus); Bükk Mountains: Borsodbóta ( Hungary) ( Csepreghy-Meznerics 1969); Krka Basin: Dolenja Brezovica ( Slovenia) ( Mikuž 2009).

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 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 23 A1–A3. Megaclavatula polonica (Pusch, 1837), neotype, NHMW 2021/0138/0001, Korytnica (Poland). B1–B3. Megaclavatula polonica (Pusch, 1837), NHMW 2021/0138/0002, Korytnica (Poland).C1–C3. Megaclavatula polonica (Pusch, 1837),NHMW2021/0139/0001,Baden-Sooss (Austria).D1–D3.Megaclavatula polonica(Pusch,1837),NHMW1862/0001/0324, Bad Vöslau (Austria). E1–E3. Megaclavatula pilleri nov. sp., paratype, NHMW 1949/0005/0051c, Baden-Sooss (Austria). F1–F3. Megaclavatula pilleri nov. sp., paratype, NHMW 1949/0005/0051d, Baden-Sooss (Austria).