Perrona eleonorae ( Hoernes & Auinger, 1891 ) 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 : 116-117

Perrona eleonorae ( Hoernes & Auinger, 1891) nov. comb.

Figs 34E 1 –E 3 , F 1 –F 2 , 35E 1 –E 3 , F 1 –F 2 , 5 , 7

Pleurotoma tuberculata Münster —Münster in Goldfuss 1841: 20, pl. 171, fig. 6c (only) [non Pleurotoma tuberculata Pusch,

1837]. Pleurotoma asperulata Lam. — Hörnes 1854 (partim): 341, pl. 37, figs 5a–b [non Megaclavatula asperulata ( Lamarck, 1822) ]. Clavatula Eleonorae —Hoernes 1891: 130 [nomen nudum]. * Pleurotoma (Clavatula) Eleonorae nobis— Hoernes & Auinger 1891: 349, pl. 45, figs 1–3. Clavatula Eleonorae R. Hoernes i Auinger— Friedberg 1912: 198, pl. 12, fig. 6. Pleurotoma (Clavatula) asperulata Lamarck — Schaffer 1908: 112, pl. 11, fig. 16 [non Megaclavatula asperulata ( Lamarck, 1822)].? Clavatula asperulata Lamarck — Csepreghy-Meznerics 1953: 9, pl. 1, figs 19–20 [non Megaclavatula asperulata ( Lamarck, 1822)]. Clavatula (Clavatula) asperulata eleonorae R. Hoernes et Auinger — Švagrovský 1958: 10, pl. 2, figs 1–3. C [lavatula]. (C [lavatula]) eleonorae (R. Hörn. et Au.) — Sieber 1958a: 157. Clavatula eleonorae Hoernes et Auinger , sp. 1891— Glibert 1960: 38. Clavatula asperulata Lamarck, 1822 — Strausz 1966 (partim): 405, pl. 15, figs 18–19 (only) [non Megaclavatula asperulata ( Lamarck, 1822)].? Clavatula asperulata Lamarck — Strausz 1966: 405, pl. 15, figs 20–21 [non Megaclavatula asperulata ( Lamarck, 1822) ]. Clavatula (Clavatula) eleonorae (Hoernes R. & Auinger) — Mikuž 1998: 71, pl. 2, fig. 3.? Clavatula eleonorae ( Hoernes & Auinger, 1891) — Kovács & Vicián 2021: 142, pl. 1, figs 28–29.

non Clavatula eleonorae ( Hoernes & Auinger, 1891) — Bałuk 2003: 33, pl. 3, figs 6–9 [cf. Granulatocincta schreibersi ( Hörnes, 1854)].

Type material. Lectotype designated herein: NHMW 1949 /0005/0052, Baden-Sooss ( Austria), SL: 52.1 mm, MD: 23.8 mm, illustrated in Hoernes & Auinger (1891, pl. 45, figs 1a–b), figs 34E 1 –E 3 . Paralectotype: NHMW 1846 /0037/0309a, Enzesfeld ( Austria) , SL: 51.5 mm, MD: 21.8 mm, illustrated in Hoernes & Auinger (1891, pl. 45, figs 2), figs 34F 1 –F 2; NHMW 1846 /0037/0309b, Enzesfeld ( Austria) , SL: 41.1 mm, MD: 16.8 mm, illustrated in Hoernes & Auinger (1891, pl. 45, figs 3), figs 35E 1 –E 3; NHMW 1847 /0037/0310, Enzesfeld ( Austria) , SL: 42.9 mm, MD: 19.2 mm, illustrated in Hörnes (1856, pl. 37, fig. 5), figs 35F 1 –F 2; NHMW 1970 /1396/1449a, Baden ( Austria) , SL: 39.3 mm, MD: 14.1 mm. NHMW 1855 /0045/0484, 14 spec., Gainfarn ( Austria) .

Additional studied material. NHMW 1937 /0002/0298, illustrated in Schaffer (1908, pl. 11, fig. 16) ; NHMW 2010 /0004/1112/ 78 spec., Bad Vöslau ( Austria) ; NHMW 2010 /0004/1114, 15 spec., Bad Vöslau ( Austria) , NHMW 1997 z0178/1131, 3 spec., Bad Vöslau ( Austria) ; NHMW 2013 /0300/0413, 1 spec., Baden-Sooss ( Austria) ; NHMW 2021 /0132/0005, 11 spec., Baden-Sooss ( Austria) ; NHMW 1970 /1396/1452, 5 spec., Baden ( Austria) ; NHMW 1847 /0046/0034, 4 spec., Szob ( Hungary) .

Revised description. Shell moderately large, solid, broad bucciniform, with gradate to conical, weakly to moderately coronate spire; apical angle ~40–45°. Protoconch not preserved. Teleoconch of up to ten whorls. Early teleoconch whorls with weak, smooth adsutural cords and comma-shaped riblets on broad, concave mid-portion. Profile of later spire whorls subcylindrical with mid-portion concave. On sixth spire whorl sculpture changes. Prominent subsutural collar bearing wide-spaced, short, open spines, broad concave mid-portion and row of indistinct broad suprasutural tubercles. Delicate spiral threads cover entire surface. Suture narrowly impressed, undulating. Last whorl 60% of total height, subcylindrical; with subsutural collar wide, but poorly delimited, bearing 12–14 short spines. Subsutural ramp broad, weakly concave, delimited by weakly tubercular, narrow shoulder cord. Peribasal and perifasciolar cords of similar strength, bearing small tubercles. Whorl profile between major cords straight to weakly concave. Siphonal fasciole strongly swollen, twisted, with prominent growth increments, often separated from columellar callus by pseudumbilical chink. Fine spiral cords cover entire whorl surface. Aperture wide, ovate. Outer lip not thickened, smooth within. anal sinus wide, moderately deep, asymmetrically U-shaped, with apex below subsutural cord; siphonal canal moderately short, wide, twisted, deeply notched at tip. Columella weakly excavated, smooth. Columellar and parietal callus strongly thickened, forming broad callus rim.

Discussion. Perrona eleonorae is morphologically close to Perrona laciniata Bellardi, 1877 , from the Tortonian of Stazzano ( Italy) (see Bellardi 1877: 178, pl. 6, fig. 4) including its varieties P. l. percaudata Sacco, 1891 and P. l. subspinulata Sacco, 1891 (see Sacco 1904: 48, pl. 12, figs 68, 69) (note that we have not seen early teleoconch whorls of P. lacinata to verify its placement in Perrona ). Both species are distinguished by the higher spire and higher base of P. laciniata and its much weaker sculpture of the subsutural cord, which lacks tubercles. Nevertheless, both species might form a phylogenetic lineage, ranging from the Langhian to the Tortonian (see also Montanaro 1937, pl. 5, figs. 53–55).

The specimen illustrated by Kovács & Vicián (2021) from Letkés ( Hungary) differs from typical P. eleonorae by its more concave whorl profile, the rapidly contracting base, and the absence of spiral cords on the spire whorls. It might represent an atypical morph.

Paleoenvironment. Specimens from Enzesfeld ( Austria), Mikulov (the Czech Republic) and Šentjernej ( Slovenia) derive from inner neritic environments, partly with sea grass (own data).

Distribution in Central Paratethys. Badenian (middle Miocene): Vienna Basin: Baden-Sooss, Bad Vöslau, Enzesfeld ( Austria) ( Hoernes & Auinger 1891), Mikulov (the Czech Republic) ( Švagrovský 1958); Pannonian Basin: Szob ( Hungary) ( Strausz 1966); Krka Basin: Šentjernej ( 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 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 7. Apertures of Paratethyan Clavatulidae (O. = Olegia, ‘C.’ = ‘Clavatula’, N. = Neoperrona, P. = Perrona, Pu. = Pusionella, Sc. = Scaevatula, S. = Striopusionella, T. = Tomellana).

FIGURE 34 A1–A3. Perrona vindobonensis (Quenstedt, 1884), NHMW1949/0005/0106a, Gainfarn (Austria). B1–B3. Perrona vindobonensis (Quenstedt, 1884), NHMW 1949/0005/0106c, Gainfarn (Austria). C1–C3. Perrona vindobonensis (Quenstedt, 1884), lectotype, NHMW 1846/0037/0334a, Enzesfeld or Gainfarn (Austria). D1–D3. Perrona vindobonensis (Quenstedt, 1884), paralectotype, NHMW 1846/0037/0334b, Enzesfeld or Gainfarn (Austria). E1–E3. Perrona eleonorae (Hoernes & Auinger, 1891), lectotype, NHMW 1949/0005/0052, Baden-Sooss (Austria). F1–F2. Perrona eleonorae (Hoernes & Auinger, 1891), paralectotype, NHMW 1846/0037/0309, Enzesfeld (Austria).

FIGURE 35 A1–A2. Perrona lydiae (Hoernes & Auinger, 1891), lectotype, NHMW 1949/0005/0112a, Gainfarn (Austria). B1–B3. Perrona lydiae (Hoernes & Auinger, 1891), NHMW 1846/0037/0333, Enzesfeld (Austria). C1–C3. Perrona lydiae (Hoernes & Auinger, 1891), NHMW 1851/0006/0014a, Bad Vöslau (Austria). D1–D3. Perrona lydiae (Hoernes & Auinger, 1891), NHMW 1851/0006/0014b, Bad Vöslau (Austria). E1–E3. Perrona eleonorae (Hoernes & Auinger, 1891), paralectotype, NHMW 1846/0037/0309b, Enzesfeld (Austria). F1–F2. Perrona eleonorae (Hoernes & Auinger, 1891), paralectotype, NHMW 1847/0037/0310, Enzesfeld (Austria). G. Perrona herculea (Mayer-Eymar, 1886), S 1523, lectotype, Grund (Austria), ETH Zürich (Switzerland).