Clavatula ’ mariae ( Hoernes & Auinger, 1891 )

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 : 94-95

‘ Clavatula ’ mariae ( Hoernes & Auinger, 1891)

Figs 28E, F 1 –F 3 , G 1 –G 2 , 4B , 5 , 7

Pleurotoma concatenata Grat. — Hörnes 1854: 344, pl. 37, figs 18–19 [non ‘Clavatula’ concatenata ( Grateloup, 1832) ].

* Pleurotoma (Clavatula) Mariae — Hoernes & Auinger 1891: 341.

Pleurotoma (Clavatula) Dorotheae nov. form. — Hoernes & Auinger 1891: 336, pl. 46, fig. 4 [non ‘ Clavatula ’ dorotheae ( Hoernes & Auinger, 1891) ].

Pleurotoma (Clavatula) Mariae Hörn. et Auing. — Schaffer 1912a: 135, pl. 49, fig. 11.

Pleurotoma (Clavatula) Mariae Hörn. et Auing. var. persculpta Schff. — Schaffer 1912a: 135, pl. 49, figs 12–17.

Pleurotoma (Clavatula) Mariae Hoern. et Auing. — Schaffer 1912b: 327.

Pleurotoma (Clavatula) Mariae Hoern. et Auing. var. persculpta Schff. — Schaffer 1912b: 327.

Pleurotoma (Clavatula) Mariae Hörnes & Auinger — Glaessner 1926: 116.

C [lavatula]. (C [lavatula]) mariae (R. Hörn. et Au.) — Sieber 1958a: 156.

C [lavatula]. (C [lavatula]) mariae persculpta Schff.— Sieber 1958a: 156.

Clavatula (Clavatula) mariae persculpta (Schaffer, 1912) — Steininger et al. 1971: 405.

Clavatula mariae (Hörnes & Auinger) — Mandic & Steininger 2003: 274.

Clavatula (Clavatula) mariae persculpta (Schaffer) — Mandic & Steininger 2003: 274.

Clavatula dorotheae (Hörnes & Auinger, 1891) —Harzhauser 2002: 118, pl. 10, figs 12–13 [non ‘ Clavatula ’ dorotheae ( Hoernes & Auinger, 1891) ].

non Clavatula (Clavatula) aff. mariae persculpta (Schaffer)— Pavlovsky 1960: 214, pl. 2, fig. 3 [unidentifiable fragment].

Type material. Lectotype designated herein: NHMW 1849/0013/0009, Mold ( Austria), SL: 23.7 mm, MD: 8.5 mm, illustrated in Hörnes (1854, pl. 37, figs 18a–b) and Schaffer (1912a, pl. 49, fig. 11), figs 28F 1 –F 3, 4B.

Illustrated material. KM Inv. No. F /1127, Maria Dreieichen ( Austria), SL: 20.1 mm , MD: 8.6 mm, illustrated in Schaffer (1912a, pl. 49, fig. 14), syntype of Pleurotoma (Clavatula) mariae persculpta (Schaffer, 1912) , fig. 28E ; KM Inv. No. F /1128, Maria Dreieichen ( Austria), SL: 20.5 mm , MD: 8.1 mm, illustrated in Schaffer (1912a, pl. 49, fig. 15), syntype of Pleurotoma (Clavatula) mariae persculpta (Schaffer, 1912) , figs 28G 1 –G2 .

Additional studied material. GBA (no number) Maria Dreieichen, illustrated in Schaffer (1912a, pl. 49, figs 16–17), stored in the Geological Survey Vienna ( Austria) (Stojaspal 1975); NHMW 1949 /0005/0040d, SL: 16.8 mm , MD: 7.2 mm, Kleinebersdorf ( Austria), illustrated in Hoernes & Auinger (1891, pl. 46, figs 4a–c) as Clavatula dorotheae .

Revised description. Shell medium-sized, biconic; apical angle ~40°. Protoconch not preserved. Teleoconch of at least nine whorls. Early teleoconch whorls flat-sided with tripartite sculpture. First two teleoconch whorls with tripartite sculpture; three beaded cords, the beads axially aligned forming riblets. Third whorl beads on subsutural cord weaken, mid-whorl concave, central cord disappears, suprasutural cord beaded. Delicate spiral threads cover entire surface including suprasutural tubercles, most prominent at mid-whorl. Suture narrowly incised, running just below suprasutural beads. Last whorl ~60% of total height, fusiform. Subsutural collar well developed, moderately swollen, subsutural ramp of medium width, weakly concave. Shoulder weakly delimited by weak to subobsolete row of tubercles, convex below, moderately constricted at base. Siphonal fasciole swollen, twisted. Sculpture of low spiral cords, broadest cords at mid-whorl and over fasciole, sometimes vaguely tubercular, with irregular secondaries intercalated, all covered by numerous delicate spiral threads. Aperture moderately wide, ovoid. Outer lip not preserved, smooth within. Anal sinus moderately wide and deep, asymmetrically U-shaped, with apex midramp. Siphonal canal moderately long. Columella moderately excavated in upper third, straight below, weakly twisted at fasciole. Columellar and parietal callus thickened, well delimited from base, forming broad callus rim.

Discussion. ‘Clavatula’ mariae ( Hoernes & Auinger, 1891) is reminiscent of Olegia , but the tripartite early teleoconch whorls excluds a placement in that genus. ‘Clavatula’ mariae displays some variability concerning the strength of the beads on the suprasutural cord and of the sculpture below the ramp on the last whorl. Similarly, the depth of the central concavity is slightly variable. Schaffer (1912a) based the separation of his subspecies persculpta on this variability, which in our opinion is unjustified. Some specimens have the entire last whorl covered in fine threads overrunning the primary spiral sculpture ( Fig. 29G ), whilst others do not ( Fig. 29F ). This species was originally identified by Hörnes (1854) as ‘ Clavatula ’ concatenata ( Grateloup, 1832) from the Chattian and Aquitanian of France ( Lauranceau 2015). Indeed, ‘ Clavatula ’ mariae resembles the French species in outline and general shape, but differs in the much more prominent beads. Moreover, the beads of ‘C. ’ concatenata are intersected by spiral grooves (see Peyrot 1931, pl. 8, figs 63, 84, 85; Lauranceau 2015, pl. 24, figs E–F; Lozouet 2017, pl. 33, figs 18–20).

Paleoenvironment. Inner neritic lagoonal environments close to coastal mudflats and sandy bottoms with seagrass (own data for Maria Dreieichen and Mold).

Distribution in Central Paratethys. Eggenburgian (early Miocene): North Alpine-Carpathian Foreland Basin: Maria Dreieichen, Mold, Gauderndorf ( Austria) ( Mandic & Steininger 2003), Karpatian (early Miocene): Korneuburg Basin: Kleinebersdorf ( Austria) ( Hoernes & Auinger 1891).

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 4A. ‘Clavatula’ justinae (Hoernes & Auinger, 1891), NHMW ex1949/0005/0110b, Buituri (Romania). B. ‘Clavatula’ mariae (Hoernes & Auinger, 1891), lectotype, NHMW 1849/0013/0009, Mold (Austria). C. ‘Clavatula’ olgae (Hoernes & Auinger, 1891), lectotype, NHMW 1862/0001/0552, Buituri (Romania). D. ‘Clavatula’ orientoromana Báldi, 1960, holotype, M.60.145, Szokolya (Hungary). E. ‘Clavatula’ juliae (Hoernes & Auinger 1891), paralectotype, NHMW 1949/0005/0043c, Lăpugiu de Sus (Romania). F. ‘Clavatula’ apolloniae (Hoernes & Auinger, 1891), NHMW 1868/0001/0474a, Lăpugiu de Sus (Romania). G. Neoperrona harzhauseri (Kovács & Vicián, 2021), NHMW 2019/0151/019a, Letkés (Hungary). H. Neoperrona zoltanorum nov. sp., holotype, NHMW 2019/0151/0108, Letkés (Hungary). I. Scaevatula sidoniae (Hoernes & Auinger, 1891), NHMW 1855/0043/0045, holotype, Lăpugiu de Sus (Romania). J. Pusionella palatina Strausz, 1954, NHMW 2021/0159/0003, Várpalota (Hungary). K. Pusionella hofmanni nov. sp., NHMW 1867/0019/0136, holotype, Coşteiu de Sus (Romania). L. Perrona barbarae (Hoernes & Auinger, 1891) NHMW 1949/0005/0103a, Rückersdorf (Austria). M. Perrona descendens (Hilber, 1879), NHMW 1846/0049/0017, St. Florian (Austria). N. Perrona emmae (Hoernes & Auinger 1891), NHMW 2021/0127/0005, Enzesfeld (Austria), NHMW 1970/1396/1449, Baden (Austria). O. Perrona ernae (Šuklje, 1929), NHMW 2021/0156/0001, Weinsteig (Austria). P. Perrona floriana (Hilber, 1879), holotype, NHMW 1858/0020/0012, Guglitz at St. Florian (Austria). Q. Perrona ilonae nov. nom., M.61.4367, Szentkút, Mátraverebély (Hungary). R. Perrona lydiae (Hoernes & Auinger, 1891), lectotype, NHMW 1949/0005/0112, Gainfarn (Austria). S. Perrona oliviae (Hoernes & Auinger 1891), paralectotype, NHMW 1873/0026/0027, Lăpugiu de Sus (Romania). T. Perrona rosaliae (Hoernes & Auinger 1891), lectotype, NHMW 1999z0098/0002, Vienna/Pötzleinsdorf (Austria). U. Perrona sabinae (Hoernes & Auinger 1891), paralectotype, NHMW 2021/0154/0001, Lăpugiu de Sus (Romania). V. Perrona styriaca (Hilber, 1879), lectotype, NHMW 1846/0037/0309, Pöls (Austria). W. Perrona ursulae (Hoernes & Auinger, 1891), paralectotype, NHMW 1846/0001/1294, Buituri (Romania). X. Perrona vindobonensis (Quenstedt, 1884), paralectotype, NHMW 1997z0178/1467, Gainfarn (Austria), Y. Perrona wanzenboecki nov. nom., NHMW 2021/0158/0001, neotype, St. Veit an der Triesting (Austria). Z. Tomellana dulaii nov. sp., NHMW 2021/0148/0002, holotype, Nemeşeşti (Romania). Aa. Tomellana aueri nov. sp., NHMW 2013/0078/0436, Baden (Austria). Ab. Striopusionella szekelyhidiae (Kovács & Vicián, 2021), holotype, PAL 2021.19.1, Letkés (Hungary). Ac. Scaevatula sidoniae (Hoernes & Auinger, 1891). NHMW 1855/0043/0045, holotype, Lăpugiu de Sus (Romania). Scale bar is 5 mm for all specimens except for Perrona barbarae and protoconch of Scaevatula sidoniae.

FIGURE 5. Anal sinus morphology of Paratethyan Clavatulidae.

FIGURE 7. Apertures of Paratethyan Clavatulidae (O. = Olegia, ‘C.’ = ‘Clavatula’, N. = Neoperrona, P. = Perrona, Pu. = Pusionella, Sc. = Scaevatula, S. = Striopusionella, T. = Tomellana).

FIGURE 28 A1–A3. ‘Clavatula’ apolloniae (Hoernes & Auinger, 1891), holotype, NHMW 1866/0040/0310, Lăpugiu de Sus (Romania). B1–B3. ‘Clavatula’ apolloniae (Hoernes & Auinger, 1891), NHMW 1868/0001/0474a, Lăpugiu de Sus (Romania). C1–C3. ‘Clavatula’ apolloniae (Hoernes & Auinger, 1891), NHMW 1868/0001/0474b, Lăpugiu de Sus (Romania). D1–D3. ‘Clavatula’ apolloniae (Hoernes & Auinger, 1891), NHMW 1868/0001/0474c, Lăpugiu de Sus (Romania). E. ‘Clavatula’ mariae (Hoernes & Auinger, 1891), KM Inv. No. F/1127, Maria Dreieichen (Austria). F1–F3. ‘Clavatula’ mariae (Hoernes & Auinger, 1891), lectotype, NHMW 1849/0013/0009, Mold (Austria). G1–G2. ‘Clavatula’ mariae (Hoernes & Auinger, 1891), KM Inv. No. F/1128, Maria Dreieichen (Austria).

FIGURE 29 A. ‘Clavatula’ inedita Bellardi, 1877, Letkés (Hungary), private collection of Zoltán Kovács and Zoltán Vicián. B. ‘Clavatula’ rusticula (Mayer-Eymar, 1886), lectotype, S 1615, Grund (Austria). C. ‘Clavatula’ orientoromana Báldi, 1960, holotype, M.60.145, Szokolya (Hungary). D1–D2. ‘Clavatula’ orientoromana Báldi, 1960, paratype, M60.146, Szokolya (Hungary). E1–E2. ‘Clavatula’ justinae (Hoernes & Auinger, 1891), paralectotypes, NHMW 1849/0005/0110b, Buituri (Romania).F1–F3. ‘Clavatula’ justinae (Hoernes &Auinger, 1891), paralectotype, NHMW 1949/0005/0110a, Buituri (Romania). G. ‘Clavatula’ justinae (Hoernes & Auinger, 1891), lectotype, NHMW 1849/0005/0110c, Buituri (Romania).