Tomellana aueri, 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 : 144-145

Tomellana aueri nov. sp.

Figs 42F 1 –F 3 , G 1 –G 3 , H, 4 Aa, 5, 7

Pleurotoma subcanaliculata Münster —Münster in Goldfuss 1841: 20 (partim), pl. 71, fig. 3. Pleurotoma semimarginata Lam. — Hörnes 1848: 20 [non Tomellana semimarginata ( Lamarck, 1822) ]. Pleurotoma semimarginata Lam. — Hörnes 1854: 347, pl. 38, figs 7–8 [non Tomellana semimarginata ( Lamarck, 1822) ]. Clavatula semimarginata (Lamck.) Varietà C — Bellardi 1877: 291. Pleurotoma (Clavatula) semimarginata Lamk. — Hoernes & Auinger 1891: 359, pl. 47, figs 17a–c [non Tomellana semimarginata ( Lamarck, 1822)]. Clavatula semimarginata (Lmk.) — Boettger 1902: 48 [non Tomellana semimarginata ( Lamarck, 1822) ]. Clavatula semimarginata (Lmk.) — Boettger 1906: 63 [non Tomellana semimarginata ( Lamarck, 1822) ]. Clavatula semimarginata Lmk. — Montanaro 1937: 131, pl. 5, figs 62–66 [non Tomellana semimarginata ( Lamarck, 1822) ]. Perrona semimarginata (Lam.) — Sieber 1947: 111 [non Tomellana semimarginata ( Lamarck, 1822) ]. Perrona semimarginata (Lk.) — Sieber 1953: 188 [non Tomellana semimarginata ( Lamarck, 1822) ]. Perrona (Perrona) semimarginata (Sm.) — Sieber 1956: 245 [non Tomellana semimarginata ( Lamarck, 1822) ]. Perrona (Perrona) semimarginata ( Lamarck 1822) — Steininger 1963: 56 [non Tomellana semimarginata ( Lamarck, 1822) ]. Clavatula (C.) semimarginata Lam. — Stancu & Andreescu 1968: 464 [non Tomellana semimarginata ( Lamarck, 1822) ]. Perrona semimarginata Lamarck — Nicorici 1972: 69 [non Tomellana semimarginata ( Lamarck, 1822) ]. Clavatula (Perrona) semimarginata mediterranea nom. nov. — Csepreghy-Meznerics 1972: 32, pl. 15, fig. 18 [nomen nudum]. Clavatula (Perrona) semimarginata (Lamarck) 1822 — Bohn-Havas 1973: 1063, pl. 6, fig. 9 [non Tomellana semimarginata ( Lamarck, 1822)]. Perrona (Perrona) semimarginata servata ( Sacco, 1890) — Nikolov 1994: 60, pl. 6, figs 11–13, pl. 7, figs 7–8. [non Tomellana ? servata ( Sacco, 1890) ]. Clavatula (Perrona) semimarginata ( Lamarck, 1822) — Mikuž 1998: 81, pl. 6, fig. 4 [non Tomellana semimarginata ( Lamarck, 1822)]. Perrona (Perrona) semimarginata praecursor (Schaffer) — Schultz 1998: 74, pl. 30, fig. 6 [non Tomellana praecursor (Schaffer, 1912)]. Perrona semimarginata ( Lamarck, 1822) —Harzhauser 2002: 120, pl. 11, fig. 7 [non Tomellana semimarginata ( Lamarck, 1822)]. Perrona semimarginata ( Lamarck, 1822) — Mikuž 2009: 34, pl. 11, fig. 154 [non Tomellana semimarginata ( Lamarck, 1822) ]. Perrona semimarginata ( Lamarck, 1822) — Kovács & Vicián 2021: 146, pl. 3, figs 19–20 [non Tomellana semimarginata

( Lamarck, 1822)].

non Clavatula (Perrona) semimarginata Lmk. — Tavani 1939: 19, pl. 2, fig. 4 [is an unidentifiable internal cast].

Type material. Holotype: NHMW 1874 /0033/0009a, Bad Vöslau ( Austria), SL: 100.5 mm, MD: 29.5 mm, Figs 42F 1 –F 3 . Paratypes: NHMW 1874 /0033/0009b, Bad Vöslau ( Austria), SL: 85.5 mm , MD: 26.7 mm, Figs 42G 1 –G 3 ; NHMW 2013 /0078/0436, SL: 20.4 mm , MD: 7.5 mm, Baden ( Austria), figs 42H, 4 Aa .

Additional paratypes. NHMW 1846 /0037/0330, Baden ( Austria) , illustrated in Hörnes (1854, pl. 38, figs 8a– b); NHMW 1853 /0016/0021, Baden ( Austria), (1854, pl. 38, figs 7a–b) ; Baden ( Austria); NHMW 1846 /0037/0331, 20 spec., Baden ( Austria) , NHMW 1949 /0005/0108, Baden ( Austria) , illustrated in Hoernes & Auinger (1891, pl. 47, figs 17a–b); NHMW 1997 z0178/0870, 12 spec., Baden ( Austria) ; SMF 351878 /1, Baden ( Austria) ; NHMW 2013 /0300/0426, 19 spec., Baden-Sooss ( Austria) ; NHMW 1866 /0040/0204, 14 spec., Baden-Sooss ( Austria) ; NHMW 2013 /0300/0571, 23 spec., Baden-Sooss ( Austria) ; SMF 351879 /1 Baden-Sooss ( Austria) ; NHMW 2013 /0078/0435, 59 spec., Baden ( Austria) ; NHMW 1874 /0024/0046, 3 spec., Bad Vöslau ( Austria) ; NHMW 2012 /0229/0220, 1 spec. , NHMW 2012 /0229/0221, 2 spec., Traiskirchen ( Austria) ; NHMW 1863 /0015/0498, 2 spec., Forchtenau ( Austria) ; NHMW 2021 /0149/0001, 1 spec., Neuruppersdorf ( Austria) ; NHMW2000 z0001/0071, Karnabrunn ( Austria) , illustrated in Harzhauser (2002, pl. 11, fig. 7); NHMW 1867 /0019/0139, 2 spec., Coşteiu de Sus ( Romania) ; SMF 351876 /1, 351877/2 Coşteiu de Sus ( Romania) .

Type locality. Bad Vöslau ( Austria), Vienna Basin.

Type stratum. Silt and clay of the Baden Formation.

Age. Middle Miocene, middle Badenian (Langhian).

Etymology. In honor of Gerald Auer (University Graz) in recognition of his contributions on Neogene micropaleontology and stratigraphy.

Diagnosis. Very large fusiform shell with slender conical spire. Protoconch multispiral of 2.5 smooth whorls. Teleoconch whorls with very broad and indistinct, rounded collar and broad concave mid-portion. Shell surface smooth except for faint spiral threads. Siphonal canal long, narrow.

Description. Shell very large fusiform of up to 106 mm in height, with tall, slender conical spire; apical angle 25–27°. Protoconch high conical of 2.5 smooth whorls. Teleoconch of up to 13 concave whorls with periphery at abapical suture, separated by narrowly impressed, superficial, linear suture. First four teleoconch whorls weakly concave with weak subsutural spiral cord and suprasutural spiral row of slightly opisthocline beads. On fifth whorl subsutural cord strengthens to form rounded collar. Broad concave mid-portion. Suprasutural cord loses beads. On late spire whorls subsutural collar and suprasutural cord become very broad and indistinct, separated by narrow concave mid-portion, resulting in weakly biconvex whorls with narrow concave mid-portion. Surface smooth except for faint spiral threads. Last whorl about 58% of total height. Broad, weakly inflated, poorly delimited subsutural collar. Weakly concave subsutural ramp. Shoulder and base delimited by a suggestion of an angulation. Mid-whorl inflated. Base strongly constricted. Fasciole weakly developed bearing strong growth increments. Spiral sculpture reduced to very faint cords over fasciole. Aperture moderately wide, pyriform. Outer lip not thickened, smooth within. Anal sinus deep, moderately wide, slightly asymmetrically U-shaped, with apex mid-ramp. Siphonal canal long, narrow, straight, shallowly notched and recurved at tip. Columella smooth, moderately excavated in upper third, straight below, very slightly twisted at fasciole. Columellar and parietal callus thickened, weakly delimited, forming relatively broad callus rim.

Discussion. Tomellana aueri nov. sp. is reminiscent of Tomellana ? servata ( Sacco, 1890) from the Tortonian of Stazzano, S. Agata and Montegibbio ( Italy) ( Bellardi 1877; Montanaro 1937; Caprotti 2011) in size and general shape. The syntype of the Italian species is stored in the Museo Regionale di Scienze Naturali Torino ( Italy), BS.011.07.070, SL: 78 mm, MD: 25 mm, S. Agata ( Italy), Tortonian (late Miocene), illustrated in Sacco (1904, pl. 13, figs 11–12). A comparison of both species (by R.J.) revealed clear differences: Tomellana ? servata has a paucispiral protoconch and lacks beads on early teleoconch whorls. The placement of the Italian species in Tomellana will need verification.

Due to the different sculpture of early teleoconch whorls, a relation with Tomellana semimarginata ( Lamarck, 1822) . Tomellana semimarginata ( Lamarck, 1822) , described from the early Miocene of the northeastern Atlantic, differs in its bulbous, paucispiral protoconch and long axial riblets on the early teleoconch whorls. The similarity between both species is superficial also if adult specimens are compared: Tomellana semimarginata has a relatively shorter spire, a much higher base with a longer siphonal canal and is smaller (SL: 73 mm) (see Peyrot 1931, pl. 6, figs 14, 15, 23; Lozouet 2017, pl. 33, fig. 6).

The oldest available name for this species is Pleurotoma subcanaliculata Münster in Goldfuss, 1841, who based his description on ‘juvenile’ shells from Bordeaux ( France) and ‘fully grown’ ones from Enzesfeld ( Austria) (Münster in Goldfuss 1841: 21). The specimens from Bordeaux are most probably identical to Tomellana semimarginata ( Lamarck, 1822) , whereas the illustrated specimen seems to represent Tomellana aueri nov. sp. The locality Enzesfeld, however, is problematic because this species is unknown from this shallow water fauna. The type specimens of Pleurotoma subcanaliculata seem to be lost and could not be traced in the collections in Bonn and Munich ( Germany). Therefore, we consider it a nomen dubium. To clarify this confused situation, as first revisers we restrict Pleurotoma subcanaliculata to the occurrences of Bordeaux and assume that this name is a junior synonym of Pleurotoma semimarginata Lamarck, 1822 .

Paleoenvironment. The species is most common in the pelitic Baden Formation suggesting a preference for middle to outer neritic environments with up to 210 m water depth ( Kranner et al. 2021). It is rarely found in shallow water deposits, where it might have been washed ashore.

Distribution in Central Paratethys. Karpatian (early Miocene): Korneuburg Basin: Karnabrunn, Stetten, Neuruppersdorf ( Austria) (Harzhauser 2002, 2003); Badenian (middle Miocene): North Alpine-Carpathian Foreland Basin: Grund, Windpassing ( Austria) ( Hoernes & Auinger 1891; Sieber 1947); Vienna Basin: Baden, Bad Vöslau, Möllersdorf, Pötzleinsdorf ( Austria) ( Hoernes & Auinger 1891), Danube Basin: Modra-Kráľová ( Slovakia) ( Hoernes & Auinger 1891, Sieber 1953); Eisenstadt-Sopron Basin: Forchtenau ( Austria) ( Sieber 1956); Pannonian Basin: Pécsszabolcs, Pécsvárad, Tekeres, Letkés, Hont, Szokolya, Nógrádszakál, Balassagyarmat, Bóta ( Hungary) ( Bohn-Havas 1973, Kovács & Vicián 2021); Bükk Mountains: Borsodbóta ( Hungary) ( Csepreghy-Meznerics 1972); Krka Basin: Gorenja Stara vas ( Slovenia) ( Mikuž 2009); Caransebeş Basin: Rugi, Delineşti ( Romania) ( Stancu & Andreescu 1968); Șimleu Basin: Tusa ( Romania) ( Nicorici 1972); Făget Basin: Coşteiu de Sus ( Romania) ( Boettger 1906); Dacian Basin: Jasen, Opanec ( Bulgaria) ( Nikolov 1994).

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 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 42 A1–A3. Scaevatula sidoniae (Hoernes & Auinger, 1891). NHMW 1855/0043/0045, holotype, Lăpugiu de Sus (Romania). B1–B2. Striopusionella szekelyhidiae (Kovács & Vicián, 2021), holotype, PAL 2021.19.1, Letkés (Hungary). C1–C2. Tomellana dulaii nov. sp., NHMW 2021/0148/0002, holotype, Nemeşeşti (Romania). D1–D2. Tomellana praecursor (Schaffer, 1912), lectotype, KM Inv. No. F/1129, Maigen (Austria). E. Tomellana praecursor (Schaffer, 1912), paralectotype, KM Inv. No. F/1130, Maigen (Austria). F. ‘Clavatula’? czarnockii (Bałuk, 2003), holotype. Catalogue No. 28.II. 179, Korytnica (Poland). F1–F3. Tomellana aueri nov. sp., holotype, NHMW 1874/0033/0009a, Bad Vöslau (Austria). G1–G3. Tomellana aueri nov. sp., paratype, NHMW 1874/0033/0009b, Bad Vöslau (Austria). H. Tomellana aueri nov. sp., paratype, NHMW 2013/0078/0436, Baden (Austria).