Megaclavatula laevigata ( Eichwald, 1830 ) 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 : 68-70

Megaclavatula laevigata ( Eichwald, 1830) nov. comb.

Figs 21E 1 –E 3 , F 1 –F 3 , 22A 1 –A 3, 3R , 5 , 6

* P [leurotoma]. laevigatum m.— Eichwald 1830: 225.

? P [leurotoma]. aculeatum m.— Eichwald 1830: 225.

Pleurotoma suturalis — Andrzejowski 1833: 441, pl. 13, fig. 5.

? Pleur [otoma]. aculeata m.— Eichwald 1852: 182.

Pleur [otoma]. laevigata m. — Eichwald 1852: 183.

Pleurotoma laevigata — Eichwald 1853, plate captions, pl. 8, figs 3a–b.

? Pleurotoma aculeata — Eichwald 1853: plate captions, pl. 8, fig. 4.

Pleurotoma (Clavatula) Austriaca Mayer-Eymar — Mayer-Eymar 1886: 308, pl. 16, fig. 6.

Pleurotoma tuberculosa Modification unilineata —Quenstedt 1884: 319, pl. 210, fig. 8

Clavatula Susannae —Hoernes 1891: 130 [nomen nudum].

Pleurotoma (Clavatula) Susannae nov. form.— Hoernes & Auinger 1891: 347, pl. 45, figs 7–8.

Clavatula Austriaca M. -E.— Hilber 1892: 1022.

? Pl [eurotoma]. aculeta Eichw. — Mikhailovsky 1903: 123.

Clavatula laevigata Eichw. — Friedberg 1912: 191 (partim), pl. 12, fig. 3, text-figs 45–46.

Clavatula laevigata Eichw. — Friedberg 1938: 142.

Clavatula asperulata (Lam.) — Csepreghy-Meznerics 1953: 9, pl. 1, figs 19–20 [non Megaclavatula asperulata ( Lamarck, 1822) ].

Clavatula (Clavatula) cf. rosaliae R. Hörn. et Auing. — Pavlovsky 1957: 53, pl. 2, figs 2a–b [non Perrona rosaliae ( Hoernes & Auinger, 1891) ].

C [lavatula]. (C [lavatula]) suzannae ( R. Hörn. et Au.)— Sieber 1958a: 157.

Clavatula (Clavatula) styriaca Auinger — Švagrovský 1958: 12, pl. 2, figs 5–7 [non Perrona styriaca ( Hilber, 1879) ].

Clavatula asperulata (Lam) — Eremija 1959: 187, pl. 1, fig. 2 [non Megaclavatula asperulata ( Lamarck, 1822) ].

Clavatula cfr. laevigata (Eichw.) — Báldi 1960: 80, pl. 3, fig. 2.

Clavatula sublaevigata n. sp. — Báldi 1960: 80, pl. 3, figs 1a–c.

Clavatula (Clavatula) laevigata Eichwald — Pavlovsky 1960: 214, pl. 2, figs 4a–b.

Clavatula laevigata Eichwald , sp. 1830— Glibert 1960: 39.

Clavatula (Clavatula) laevigata var. amaliae (Hoernes und Auinger 1891) —Kojumdgieva in Kojumdgieva & Strachimirov 1960: 198, pl. 47, fig. 9 [non Megaclavatula amaliae ( Hoernes & Auinger, 1891) ].

Clavatula asperulata Lamarck, 1822 — Strausz 1966 (partim): 405, pl. 15, figs 20–21 (only) [non Megaclavatula asperulata ( Lamarck, 1822) ].

Clavatula asperulata suturalis Andrezejowski, 1833 — Strausz 1966: 407, pl. 15, fig. 22, pl. 16, figs 1–2.

Clavatula styriaca Auinger (in Hilber)— Strausz 1966: 402, pl. 15, figs 11–14 [non Perrona styriaca ( Hilber, 1879) ].

Clavatula (C.) laevigata Eichw. — Stancu & Andreescu 1968: 464, pl. 6, fig. 67.

Clavatula laevigata ( Eichwald, 1830) — Zelinskaya et al. 1968: 221, pl. 50, figs 18–19.

Clavatula laevigata (Eichwald) — Radwański 1969: 162, pl. 37, figs 13, 15–16.

Clavatula asperulata susannae Hoernes et Auinger — Csepreghy-Meznerics 1969: 94, pl. 6, figs 1–2.

Clavatula asperulata susannae Hoernes et Auinger — Csepreghy-Meznerics 1972: 32, pl. 15, figs 22–23.

Clavatula asperulata (Lamarck) 1822 — Bohn-Havas 1973: 1062, pl. 4, fig. 7 [non Megaclavatula asperulata ( Lamarck, 1822) ].

Clavatula (Clavatula) styriaca ( Hilber, 1879) — Švagrovský 1982: 413, pl. 8, fig. 2 [non Perrona styriaca ( Hilber, 1879) ]. Clavatula laevigata (Eichwald) — Bałuk & Radwański 1996: 286, pl. 3, fig. 13, pl. 5, fig. 2.

Clavatula (Clavatula) cf. styriaca ( Hilber, 1879) — Mikuž 1998: 73, pl. 4, fig. 1 [non Perrona styriaca ( Hilber, 1879) ]. Clavatula (Clavatula) laevigata ( Eichwald, 1830) — Mikuž 1998: 72, pl. 3, figs 1–2.

Clavatula laevigata ( Eichwald, 1830) —Harzhauser 2002: 118, pl. 10, fig. 17, pl. 11, fig. 1.

Clavatula suturalis ( Andrzejowski, 1833) — Bałuk 2003: 39, pl. 8, figs 1–8.

Clavatula laevigata ( Eichwald, 1853) — Bałuk 2003: 35, pl. 7, figs 7–10.

? Clavatula styriaca ( Hilber, 1879) — Bałuk 2003: 38, pl. 6, fig. 4 [non Perrona styriaca ( Hilber, 1879) ].

Clavatula laevigata (Hilber) — Mikuž 2003: 308, pl. 10, fig. 29

Clavatula sublaevigata Báldi, 1960 — Pálfy et al. 2008: 105.

Clavatula laevigata ( Eichwald, 1830) — Mikuž 2009: 31, pl. 10, figs 137–138.

Clavatula suturalis susannae (Hoernes R. & Auinger M. 1890) — Katona et al. 2011: 9, pl. 1, fig. 7.

Clavatula suturalis ( Andrzejowski, 1833) — Kovács & Vicián 2021: 143, pl. 2, figs 12–18.

non Clavatula (Clavatula) laevigata ( Eichwald 1853) —Kojumdgieva in Kojumdgieva & Strachimirov 1960: 197, pl. 47, figs 7–8 [= ‘ Clavatula ’ juliae ( Hoernes & Auinger, 1891) ].

Type material. Syntype: Described by Eichwald (1830) from Szuszkowce ( Ukraine). The specimen might be stored in the Geological Museum of the Geological Department of Kiev University ( Nosowska 2020), but we did not receive any information from that institution. The syntype of Clavatula suturalis , which was illustrated in Andrzejowski (1833, pl. 13, fig. 5) derives from Varivtsi (= Warowce) ( Ukraine). The whereabouts of the collections of Antoni Lukianowicz Andrzejowski (1784–1868) is unknown ( Daszkiewicz & Bauer 2008).

Illustrated material. NHMW 1853/0038/0035a, SL: 74.7 mm, MD: 30.8 mm, Korytnica ( Poland), figs 21F 1 –F 3; NHMW 1853/0038/0035b, SL: 65.8 mm, MD: 25.5 mm, Korytnica ( Poland), figs 21E 1 –E 3, 3 R; NHMW 1846/0037/0308, Baden ( Austria), SL: 76.1 mm, MD: 28.7 mm, syntype of Pleurotoma (Clavatula) Susannae Hoernes & Auinger, 1891 , illustrated in Hoernes & Auinger (1891, pl. 45, fig. 8), figs 22A 1 –A 3; M60.139, SL: 38.4 mm, MD: 15.4 mm, Bíró-völgy, Szokolya ( Hungary), holotype of Clavatula sublaevigata Báldi, 1960 , illustrated in Báldi (1960, pl. 3, figs 1a–c), stored in the Hungarian Natural History Museum, Budapest ( Hungary).

Additional studied material. NHMW 2021/0134/0001, 26 spec., Grund ( Austria); NHMW 1846/0037/0308, 1 spec., Baden ( Austria); NHMW 1853/043/0034, 2 spec., Korytnica ( Poland); NHMW 1997z0178/1446, 3 spec., Gainfarn ( Austria), NHMW 2002z0181/0287, 4 spec., Weitendorf ( Austria); NHMW 2000z0001/0067, 21 spec., Kleinebersdorf ( Austria). S 1415, 32 spec., Grund ( Austria), stored in the ETH Zürich ( Switzerland); syntypes of Pleurotoma (Clavatula) austriaca Mayer-Eymar, 1886 .

Revised description. Shell large, solid, moderately broad fusiform with high, coronate spire; apical angle ~40– 45°. Protoconch not preserved. Teleoconch of at least ten whorls, concave mid-whorl. Early teleoconch whorls with tripartite spiral sculpture consisting of weak, vaguely beaded subsutural cord, even weaker, beaded mid-whorl cord, and prominently beaded suprasutural cord. Sculpture changing around fifth to sixth teleoconch whorl; subsutural cord develops into subsutural collar with wide spaced, spiny, trigonal tubercles; mid-whorl cord disappears. Beads on suprasutural cord turning into indistinct tubercles partially covered by subsequent whorl on mid-spire whorls, completely obscured on penultimate and last whorls. Spiral sculpture of delicate spiral threads on early teleoconch whorls, becoming weak or subobsolete on last whorl. Suture narrowly incised, weakly undulating. Last whorl ~65% of total height, poorly delimited but broad, spinous subsutural collar, distinctly concave subsutural ramp, angled at shoulder and peribasal cords. Base strongly constricted, concave. Siphonal fasciole long, almost straight, with prominent growth increments. Two primary spiral cords at mid-whorl bearing small tubercles, adapical delimiting shoulder, abapical delimiting base, with secondary spirals intercalated. Weak irregular cords over base and siphonal fasciole. Prominent, close-spaced axial growth lines give surface slightly scabrose appearance. Aperture moderately wide, pyriform. Outer lip not thickened, smooth within. Anal sinus moderately wide and deep, asymmetrically Vshaped, with apex below subsutural collar in deepest part of subsutural concavity. Siphonal canal moderately long, straight to slightly deflected, shallowly notched at tip. Columella weakly excavated, slightly twisted at fasciole, smooth. Columellar and parietal callus moderately thickened, adherent, forming broad callus rim.

Discussion. Already Eichwald(1852) stated that Pleurotoma suturalis Andrzejowski, 1833 was a junior synonym of his P. laevigata . Nevertheless, a few subsequent authors used suturalis again as a valid name for Paratethyan Clavatulidae (e.g., Strausz 1966). Bałuk (2003), however, distinguished Clavatula laevigata and Clavatula suturalis , based on the frequent presence of a certain trace-fossil described by Radwański & Bałuk (1997) as Clavatulicola evaephilus, which was considered by these authors to be associated only with C. laevigata . In addition, Bałuk (2003) stated that C. laevigata was smaller and more slender compared to C. suturalis . Tucker (2004) and Kovács & Vicián (2021), in contrast, considered them to be synonyms. Tentatively, we follow this position, but stress the lack of plentiful material. In contrast, we follow Bałuk (2003) in considering Megaclavatula susannae ( Hoernes & Auinger, 1891) as subjective junior synonym of M. laevigata . The only difference between M. laevigata and M. susannae is the largely reduced spiral sculpture of the latter.

The name Pleurotoma suturalis was already used by Bronn (1831: 48) but is a nomen nudum. Moreover, although Bronn (1831) listed P. suturalis as new, he also refers to Defrancia suturalis Millet, 1827 , suggesting that his name was not intended to be new.

This species might have been described by Eichwald (1830) also as Pleurotoma aculeatum Eichwald, 1830 from Szuszkowce ( Ukraine). Eichwald (1852) considered Pleurotoma tuberculosa polonica Pusch, 1837 to be a synonym of his P. aculeatum . Bałuk (2003) listed P. aculeatum also as synonym of Megaclavatula polonica , overlooking that Eichwald’s name would have priority over M. polonica . Pleurotoma aculeatum as described and illustrated by Eichwald (1852, 1853), however, is characterized by a raised central spiral cord on early spire whorls, which persists to the fifth to sixth spiral cord. Such spiral cord is not developed in M. polonica , but may be present in M. laevigata ( Eichwald, 1830) (see Bałuk 2003: pl. 8, figs 6, 7). Moreover, the number and morphology of the tubercles of M. aculeata is close to M. laevigata , but differ from the few but spinose tubercles in M. polonica . Therefore, we tentatively consider Pleurotoma aculeatum Eichwald, 1830 and Pleurotoma laevigata Eichwald, 1830 to be subjective synonyms. As first revisers, we chose Megaclavatula laevigata as name for this species.

Paleoenvironment. Most occurrences suggest inner neritic environments; e.g., Zuschin et al. (2007) for Gainfarn and Enzesfeld ( Austria), Švagrovský (1981) for Borský Mikuláš ( Slovakia), Kovács & Vicián (2013) for Letkés ( Hungary).

Distribution in Central Paratethys. Karpatian (early Miocene): Korneuburg Basin: Kleinebersdorf, Gaweinstal ( Austria) (Harzhauser 2002, 2003); Badenian (middle Miocene): Korytnica Basin: Korytnica ( Poland) ( Bałuk 2003); Ukrainian Fore-Carpathian Basin: Varivtsi (= Warowce) ( Ukraine) ( Andrzejowski 1833); North-Alpine Foreland Basin: Grund ( Austria) (hoc opus); Vienna Basin: Baden, Bad Vöslau, Enzesfeld, Forchtenau, Gainfarn, Vienna /Pötzleinsdorf, Steinebrunn ( Austria), Borský Mikuláš ( Slovakia), Mikulov (the Czech Republic) ( Hörnes 1854; Sieber 1953; Švagrovský 1982); Oberpullendorf Basin: Ritzing ( Austria) ( Hörnes 1854); Pannonian Basin: Pécsszabolcs, Hird, Szob, Letkés, Szokolya, Várpalota (Faller Street) ( Hungary) ( Báldi 1960; Strausz 1966; BohnHavas 1973; Katona et al. 2011; Kovács & Vicián 2021); Bükk Mountains: Borsodbóta ( Hungary) (CsepreghyMeznerics 1972); Zapresic-Brijeg at Samobor ( Croatia) ( Pavlovsky 1960); Glina Basin: Glina ( Croatia) ( Eremija 1959; Pavlovsky 1957; Pavlovsky 1960); Salgótarján Basin: Nógrádszakál ( Hungary) ( Bogsch 1936); Caransebeş Basin: Rugi, Delineşti ( Romania) ( Stancu & Andreescu 1968): Krka Basin: Gorenja Stara vas, Dolenja Brezovica, Ivanji dol ( Slovenia) ( Mikuž 1998).

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 21 A1–A3. Megaclavatula evae (Hoernes & Auinger, 1891), paralectotype, NHMW 1949/0005/0047b, Guntersdorf (Austria). B1–B3. Megaclavatula evae (Hoernes & Auinger, 1891), lectotype, NHMW 1949/0005/0047a, Guntersdorf (Austria). C1–C3. Megaclavatula grunerti nov. sp., holotype, NHMW A1382, Enzesfeld (Austria), D1–D3. Megaclavatula grunerti nov. sp., paratype, NHMW 2021/0127/0003, Enzesfeld (Austria). E1–E3. Megaclavatula laevigata (Eichwald, 1830), NHMW 1853/0038/0035b, Korytnica (Poland). F1–F3. Megaclavatula laevigata (Eichwald, 1830), NHMW 1853/0038/0035a, Korytnica (Poland).

FIGURE 22 A1–A3. Megaclavatula laevigata (Eichwald, 1830), NHMW 1846/0037/0308, Baden (Austria). B1–B3. Megaclavatula subsculpta (Schaffer, 1912), lectotype, KM Inv. No. F/1125, Maria Dreieichen (Austria). C1–C2. Megaclavatula nemethi (Kovács & Vicián, 2021), holotype, HNHM, PAL 2021.21.1, Letkés (Hungary). D. Megaclavatula nemethi (Kovács & Vicián, 2021), paratype, NHMW 2021/0002/0003, Letkés (Hungary). E1–E2. Megaclavatula? szontaghyi (Strausz, 1926), holotype, M.116, Bán, Baranya County (Hungary). F. Megaclavatula neudorfensis (Schaffer, 1898), holotype, NHMW 2021/0137/0001, Devínska Nová Ves (Slovakia). G1–G3. Megaclavatula pilleri nov. sp., holotype, NHMW 1949/0005/0051a, Baden-Sooss (Austria). H1–H3. Megaclavatula pilleri nov. sp., paratype, NHMW 1949/0005/0051b, Baden-Sooss (Austria).