Pusia avellanella Harzhauser & Landau 2021, nov. comb.

Pusia avellanella (Boettger, 1906) nov. comb.

Figs 4H, 13A 1 –A 2, B 1 –B 2, C 1 –C 2, D 1 –D 2, E 1 –E 2, F 1 –F 2

Mitra ebenus Lam. — Hörnes 1852b: 109 (pars) [non Lamarck, 1811].

Mitra (Volutomitra) ebenus Lamk. — Hoernes & Auinger 1880: 82 pars, pl. 7, figs 22–25 [non Lamarck, 1811].

[Mitra] (Costellaria) miocaenica DD n. sp. — Dollfus & Dautzenberg 1886: 102, nr. 189 [nov. nom. pro Mitra ebenus Hörnes & Auinger pl. 9, figs 22–25], [nomen nudum; non Michelotti, 1847].

* Mitra (Uromitra) avellana Bell. var. avellanella n. var. — Boettger 1906: 9.

Mitra (Uromitra) avellana Bell. var. pseudavellana n. var. — Boettger 1906: 9.

Turricula intermittens R. Hoern. i Auinger var. dryszczoviensis mihi— Friedberg 1911: 27, pl. 1, fig. 19.

Vexillum (Uromitra) avellana pseudavellana (Boettger) — Zilch 1934: 259, pl. 17, fig. 20.

Vexillum (Uromitra) avellana avellana (Boettger) — Zilch 1934: 259, pl. 17, fig. 29.

Turricula intermittens R. Hoern. u. Auing- Friedberg, 1938: 132.

Turricula ebenus Lam. var. laevis Eichw. — Sieber 1947: 157 [non Voluta laevis Eichwald, 1829].

V [exillum]. (V [exillum].) ebenus striatum (Eichw.)— Sieber 1958a: 153 [non Mitra striata Eichwald, 1830].

Vexillum (V.) ebenus striatum (Eichw.)— Sieber 1958b: 149 [non Mitra striata Eichwald, 1830].

Vexillum (Vexillum) ebenus var. striata (Eichwald 1830) —Kojumdgieva in Kojumdgieva & Strachimirov 1960: 160, pl. 42, fig. 10 [non Mitra striata Eichwald, 1830].

Vexillum (Vexillum) cognatum (Bellardi, 1887) — Bałuk 1997: 34, pl. 9, figs 3–5 [non Pusia (Ebenomitra) cognata (Bellardi, 1887)].

Vexillum (Vexillum) pseudavellana (Boettger, 1906) — Bałuk 1997: 36, pl. 9, fig. 7.

Type material. Lectotype: SMF 360343 (= XII 2209 a), Senckenberg Museum, Frankfurt/Main, Germany, SL: 4.75 mm, MD: 2.1 mm, Coşteiu de Sus (Romania), designated and illustrated in Zilch (1934, pl. 17, fig. 19), figs 13A 1 –A 2. Paralectotypes: SMF XII 2209 b–c, Senckenberg Museum, Frankfurt / Main, Germany, Coşteiu de Sus (Romania).

Additional material. SMF 360344 (= SMF XII 2210 a), SL: 6.5 mm, MD: 2.8 mm, Senckenberg Museum, Frankfurt/Main, Germany, Coşteiu de Sus (Romania), lectotype of Mitra (Uromitra) avellana pseudavellana Boettger, 1906, figs 13D 1 –D 2; NHMW 2020/0072/0001, SL: 10.1 mm, MD: 3.1 mm, Gainfarn (Austria), figs 13C 1 –C 2,4H; NHMW 2020/0072/0002, SL: 9.9 mm, MD: 4.0 mm, Gainfarn (Austria), figs 13B 1 –B 2; NHMW 2020/0072/0003, SL: 9.2 mm, MD: 3.9 mm, Gainfarn (Austria), NHMW 2020/0072/0004, SL: 9.2 mm, MD: 3.) mm, Gainfarn (Austria), illustrated in Hoernes & Auinger (1880, pl. 9, figs 23a–c), syntype of Mitra (Costellaria) miocaenica Dollfus & Dautzenberg 1886, figs 13E 1 –E 2; NHMW 2020/0073/0001, SL: 11.2 mm, MD: 3.9 mm, Jerutek at Lysice (Czech Republic), illustrated in Hoernes & Auinger (1880, pl. 9, figs 24a–c), syntype of Mitra (Costellaria) miocaenica Dollfus & Dautzenberg 1886; NHMW 1847/0037/0048, SL: 14.1 mm, MD: 5.4 mm, Steinebrunn (Austria), illustrated in Hoernes & Auinger (1880, pl. 9, figs 22a–c), syntype of Mitra (Costellaria) miocaenica Dollfus & Dautzenberg 1886, figs 13F 1 –F 2: NHMW 1859/0037/0006, SL: 11.9 mm, MD: 4.3 mm, Lăpugiu de Sus (Romania), illustrated in Hoernes & Auinger (1880, pl. 9, figs 25a–c), syntype of Mitra (Costellaria) miocaenica Dollfus & Dautzenberg 1886; NHMW 1865/0015/0018, 8 specimens, Jerutek at Lysice (Czech Republic), NHMW 2020/0074/0001, 19 specimens, Steinebrunn (Austria), NHMW 1846/0037/0115, 24 specimens, Gainfarn (Austria), NHMW 2020/0072/0005, 50 specimens, Gainfarn (Austria), NHMW 2020/0075/0001, 3 specimens, Lăpugiu de Sus (Romania). ZNG PAN A-I-50/72, syntype of Turricula intermittens R. Hoern. i Auinger var. dryszczoviensis Friedberg, 1911, illustrated in Friedberg (1911, pl. 1, fig. 19), Dryszczów (Ukraine), stored in the Geological Museum of the Institute of Geological Sciences, Polish Academy of Sciences). ZNG PAN A-I-50/73.1–73.12, 12 specimens, syntypes of Turricula intermittens R. Hoern. i Auinger var. dryszczoviensis Friedberg, 1911, Dryszczów (Ukraine), stored in the Geological Museum of the Institute of Geological Sciences, Polish Academy of Sciences).

Revised description. Shell moderately large, moderately broad to moderately slender fusiform, with high spire. Protoconch (fig. 4H) high conical, mammillate, consisting of 3.5 convex whorls (height: 800 μm, diameter: 650 μm). Protoconch axis tilted from shell axis in many specimens. Teleoconch of eight weakly convex to subcylindrical whorls, with periphery at abapical suture, separated by narrowly impressed suture. Axial sculpture of 15–18 closeset orthocline, convex ribs, separated by slightly narrower interspaces; axials very variable, typically weakening on early spire whorls or rarely persisting over entire spire. Last whorl subcylindrical or weakly convex, with rounded shoulder placed close below adapical suture, rapidly contracting, bearing reduced axial sculpture, often more distinct along basal convexity fading out quickly towards shoulder and abapically on basal concavity. Aperture moderately narrow, ovoid with indistinct, narrow anal sinus, faint parietal denticle. Columella narrow, sharply delimited. Columella with four folds, weakening rapidly abapically. Outer lip thin, with about ten prominent, long lirae within. No spiral sculpture except for few spiral cords on weak fasciole, partly forming continuations of columellar folds. Siphonal canal short, straight or slightly bent to left, with very shallow siphonal notch.

Shell measurements and ratios. SL: 4.8–14.1 mm, MD: 2.1–5.4 mm; AA: 40–45°, A/S: 0.8–0.9.

Discussion. This species was confused by Hörnes (1852b) and Hoernes & Auinger (1880) with the extant Pusia (Ebenomitra) ebenus (Lamarck, 1811), which differs in its much larger size and much broader axial ribs. Consequently, Dollfus & Dautzenberg (1886) introduced Mitra miocaenica (written miocaenica) as new name for the specimens illustrated by Hoernes & Auinger (1880, pl. 9, figs 22–25) in a list of Miocene molluscs from the Touraine (France), without any description or comment. Aside from being a nomen nudum, the name Mitra miocaenica, was already preoccupied for an early Miocene species from Italy by Michelotti (1847: 310). According to ICZN Article 58.1 “ species-group names established for different nominal taxa that differ in spelling only in any of the following respects and that are of the same derivation and meaning are deemed to be homonyms when the nominal taxa they denote are included in the same genus or collective group: use of ae, oe or e.” Therefore, Mitra miocaenica Dollfus & Dautzenberg, 1886 is not available. Boettger (1902) compared this species with Pusia avellana (Bellardi, 1887b) from the early Miocene of Italy and separated it as variety avellanella. This ‘variety’ was based on juvenile specimens (see lectotype), whereas Boettger (1906) described fully grown specimens as variety pseudavellana. As first revisers we chose Pusia avellanella as the valid name for this species.

Pusia avellana (Bellardi, 1887) differs from the Paratethyan species by its larger size and stouter outline (see Ferrero-Mortara et al., 1981 pl. 51, figs 10a–b). Bałuk (1997) identified the Paratethyan species as Vexillum cognatum, which was described by Bellardi (1887b) from the Tortonian of Stazzano (Italy). The holotype, illustrated in Ferrero-Mortara et al. (1981 pl. 51, figs 7a–b), differs from Pusia avellanella in its more prominent axial ribs, which persist over the entire last whorl. Turricula intermittens dryszczoviensis Friedberg, 1911 is based on slender morphotypes of Pusia avellanella. A relation with Pusia intermittens (Hoernes & Auinger, 1880) is unlikely due to the less fusiform shell with shorter spire and shorter siphonal canal.

Pusia avellanella is quite variable concerning size and intensity of its axial sculpture, but specimens with persisting axial sculpture are exceptional.

Palaeoenvironment. This species is most abundant at localities suggesting silty-sandy inner neritic environments partly with sea grass meadows (e.g. Gainfarn, Steinebrunn) (Zuschin et al. 2007) but is also found in pelitic deposits, which might correspond to deeper environments.

Distribution in Central Paratethys. Badenian (middle Miocene): Korytnica Basin: Korytnica (Poland) (Bałuk 1997); Ukrainian Fore-Carpathian Basin: Dryszczów (Nadrichne) (Ukraine) (Friedberg 1911); North-Alpine-Carpathian Foredeep: Jerutek at Lysice (Czech Republic) (Hoernes & Auinger 1880); Vienna Basin: Gainfarn, Steinebrunn (Austria) (Sieber 1958b); Făget Basin: Lăpugiu de Sus (Romania) (Hoernes & Auinger 1880).