Ebenomitra leucozona ( Andrzejowski, 1830 ) Harzhauser & Landau, 2021

Harzhauser, Mathias & Landau, Bernard, 2021, An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea, Zootaxa 4982 (1), pp. 1-70 : 28-30

Ebenomitra leucozona ( Andrzejowski, 1830) nov. comb.

Figs 4E , 10A 1 –A 2 , B 1 –B 2 , C 1 –C 2 , 11A 1 –A 2

* Mitra leucozona Nobis — Andrzejowski 1830: 98, pl. 4, figs 6a–b.

Mitra striata m.— Eichwald 1830: 221.

Mitra striata — Eichwald 1852: pl. 8, fig. 16.

Mitra leucozona Andrzeiowski —Pusch 1837: 119, pl. 11, figs 6a–b.

[ Mitra ] buccinula n. sp. — Hauer 1837: 417.

Mitra striata — Eichwald 1840: 12.

[ Mitra ] buccinula [Partsch]— Hörnes 1848: 17.

[ Mitra ] parvula [Partsch]— Hörnes 1848: 17.

Mitra buccinula Partsch — d’Orbigny 1852: 54, nr. 922.

Mitra ebenus Lam. — Hörnes 1852b: 109, pl. 10, figs 11–13 [non Lamarck, 1811].

Mitra striata m.—Eichwald: 1853: 202.

[ Mitra ] ebenus Lam. — Auinger 1871: 8 [non Lamarck, 1811].

Volutomitra ebenus Lamk. —Hoernes 1880: 125 [non Lamarck, 1811].

Mitra (Volutomitra) ebenus Lamk. — Hoernes & Auinger 1880: 82 pars, pl. 7, figs 10–12, 14 [non pl. 7, fig. 13 = Pusia (Ebeno-mitra) pseudopyramidella ( Boettger, 1906) , non pl. 9, figs 22–25 = Pusia avellanella ( Boettger, 1906) ] [non Lamarck, 1811].

Mitra leucozona Andz. — Hilber 1882: 2.

Mitra striata Eichw. — Hilber 1882: 2, pl. 1, figs 2–3.

Turricula ebenus Lam. — Friedberg 1911: 18, pl. 1, figs 10–11 [non Lamarck, 1811].

Turricula ebenus Lam. var. leucozona Andrz. — Friedberg 1911: 20, pl. 1, figs 12–13.

Mitra ebenus Lam. — Meznerics 1933: 322 [non Lamarck, 1811].

Vexillum (Uromitra) ebenum (Lk.) — Sieber 1953: 188 [non Lamarck, 1811].

Mitra ebenus Lamarck — Csepreghy-Meznerics 1954: 46, pl. 6, figs 6–7 [non Lamarck, 1811].

Vexillum (Uromitra) ebenum (Lm.) (= leucozona )— Sieber 1956: 244 [non Lamarck, 1811].

V [exillum]. (V [exillum].) ebenus ebenus (Lm.) — Sieber 1958a: 153 [non Lamarck, 1811].

V [exillum]. (V [exillum].) ebenus leucozona (Andrz.) — Sieber 1958a: 153.

Vexillum (V.) ebenus (LM.) — Sieber 1958b: 149 [non Lamarck, 1811].

Vexillum (Vexillum) ebenus var. leucozona ( Andrzejowski 1830) —Kojumdgieva in Kojumdgieva & Strachimirov 1960: 160, pl. 42, figs 9a–b.

Mitra (Pusia) ebenus Lamarck — Strausz 1966: 366, pl. 26, fig. 5, figs 3–5 [non Lamarck, 1811].

Vexillum (Vexillum) ebenus leucozona (Andrzejowsky, 1830) — Hinculov 1968: 146, pl. 36, figs 8a–b, 9a–b.

Vexillum ebenus leucozona (Andrzejowskj) — Bałuk 1970: 119, pl. 12, fig. 7.

Mitra (Pusia) ebenus Lamarck, 1811 — Nicorici & Sagatovici 1973: 173, pl. 27, figs 16–17 [non Lamarck, 1811].

Mitra (Ebenomitra) ebenus ebenus Lamarck, 1811 — Švagrovský 1982: 397, pl. 4, fig. 3 [non Lamarck, 1811].

Vexillum (Vexillum) leucozona ( Andrzejowski, 1830) — Bałuk 1997: 35, pl. 9, figs 1–2.

Vexillum (Vexillum) leucozona ( Andrzejowski, 1830) — Mikuž 2009: 29, pl. 9, figs 126–127.

? Turricula leucozona Andrzejowski — Peyrot 1938: 243.

? Vexillum (Vexillum) leucozona Andrzejowski — Glibert 1952: 360, pl. 12, figs 2a–b [non 2c].

? Vexillum leucozona ( Andrzejowski, 1830) — Davoli 2000: 198, pl. 7, figs 8, 23–30.

non Mitra (Vexillum) ebenus leucozona (Andrzejowskj) — Krach 1981: 72, pl. 20, figs 1–5 [= Pusia nov. sp.?].

non Vexillum leucozona ( Andrzejowski, 1830) — Iljina 1993: 101, pl.13, figs 13–14.

Type material. Syntype described and illustrated by Andrzejowski (1830, pl. 4, figs 6a–b) from Szuszkowce (Shushkivtsi) ( Ukraine). The whereabouts of the Andrzejowski collection are unknown ( Daszkiewicz & Bauer, 2008).

Additional material. NHMW 1846 /0037/0110, SL: 21.0 mm, MD: 8.9 mm, Gainfarn ( Austria) , illustrated in Hörnes (1852b, pl. 10, fig. 11), figs 10B 1 –B 2. NHMW 1868 /0001/0401, SL: 16.7 mm, MD: 8.2 mm, Lăpugiu de Sus ( Romania) , illustrated in Hoernes & Auinger (1880, pl. 7, figs 11a–b, figs 10C 1 –C 2. NHMW 1870 /0033/0034, SL: 13.8 mm, MD: 6.3 mm, Lăpugiu de Sus ( Romania) , illustrated in Hoernes & Auinger (1880, pl. 7, figs 12a–b), figs 10A 1 –A 2. NHMW 1876 /0011/0008, SL: 13.5 mm, MD: 6.4 mm, Lăpugiu de Sus ( Romania) , figs 4E; NHMW 1854 /0035/0099, SL: 23.4 mm, MD: 9.8 mm, Lăpugiu de Sus ( Romania) , illustrated in Hoernes & Auinger (1880, pl. 7, figs 10a–b); NHMW 1857 /0014/0008, SL: 14.0 mm, MD: 6.9 mm, Steinebrunn ( Austria) , figs 11A 1 –A 2. NHMW 2020 /0065/0001, SL: 14.5 mm, MD: 6.6 mm, Ternopil ( Ukraine) , illustrated in Hoernes & Auinger (1880, pl. 7, figs 14a–b). NHMW 1857 /0014/0008, 12 specimens ; NHMW 2020 /0066/0001, 3 specimens, Steinebrunn ( Austria) ; NHMW 2020 /0083/0001, 4 specimens, Baden ( Austria) ; NHMW 1872 /0030/0022, 1 specimen , NHMW 1866 /0001/0626, 1 specimen, Baden-Soo ( Austria) ; NHMW 1997 z0178/1106, 20 specimens, Bad Vöslau ( Austria) ; NHMW 2020 /0068/0001, 6 specimens , NHMW 1997 z0178/1429, 17 specimens, Gainfarn ( Austria) ; NHMW 2020 /0067/0001, 1 specimen, Grund ( Austria) ; NHMW 2020 /0070/0001, 36 specimens, Vienna / Pötzleinsdorf ( Austria) ; NHMW 1861 /0001/0235, 5 specimens, Pöls ( Austria) ; NHMW 1863 /0015/0293, 2 specimens, Szob ( Hungary) ; NHMW 1858 /0047/0004, 1 specimen, Sedlec (= Porzteich) ( Czech Republic) ; NHMW 1851 /00010/0016, 2 specimens, Mikulov ( Czech Republic) ; NHMW 1876 /0011/0008, 46 specimen, Lăpugiu de Sus ( Romania) .

Revised description. Shell moderately large, solid, broadly biconic, with moderately high, gradate spire. Protoconch (fig. 4E) paucispiral, with low nucleus and about two moderately convex whorls (height: 550 μm, diameter: 550 μm). Transition to teleoconch not well preserved. Teleoconch of about eight whorls; early spire whorls straight sided, with convex, orthocline axial ribs, separated by slightly narrower interspaces. Ribs and interspaces wider on fourth to sixth teleoconch whorls, ribs broad, poorly delimited on last whorl (eight to ten on last whorl) causing faint convexity of whorls. Suture impressed, undulating. Last whorl short, barrel-shaped to faintly convex, moderately constricted at base. Aperture wide, with shallow, narrow anal canal. Columellar callus narrow, indistinct. Columella with three folds, fourth fold subobsolete; adapical fold most prominent, sharp. Outer lip thin, without denticles; about 10 delicate long lirae inside aperture. No spiral sculpture except for weak spiral threads on base and more prominent spiral cords on fasciole, extending from columellar folds. Siphonal canal short, straight or bent weakly to left, with shallow siphonal notch. Colour pattern frequently preserved, consisting of orange-brown background and white band along abapical suture on spire whorls, continuing above mid-whorl on last whorl; two narrower white bands along base and fasciole.

Shell measurements and ratios. SL: 12.0– 21.2 mm, MD: 5.5–9.5 mm; AA: 45–52°, SL/MD: 2.2–2.3, AL/AW: 2.1–2.8, AH/S: 3.4–3.6.

Discussion. Eichwald (1830, 1840, 1852, 1853) described this species as Mitra striata although he was aware that it was identical to M. leucozona Andrzejowski, 1830 (e.g. Eichwald 1840: 12). Andrzejowski’s paper, however, appeared earlier than Eichwald’s paper ( Nosowska 2020) and M. leucozona takes precedence.

Ebenomitra leucozona is a very characteristic species due to its subcylindrical last whorl bearing broad, poorly delimited ribs. Generally, it displays only moderate variability concerning size, slenderness and axial ribbing on the last whorl. Therefore, it is surprising that this species was mixed by Eichwald (1840, 1853), Hörnes (1852b) and Hoernes & Auinger (1880) with smooth and slender shells of other species such as Ebenomitra pseudopyramidella ( Boettger, 1906) and Pusia avellanella ( Boettger, 1906) . The large, barrel-shaped, smooth specimen from Lăpugiu de Sus ( Romania), illustrated in Hoernes & Auinger (1880, pl. 7, figs 10a–b), seems to be an aberrant specimen of E. leucozona based on its identical colour pattern and columellar folds. The specimens from the Badenian of Węglinek i Łychów ( Poland), described by Krach (1981, pl. 20, figs 1–5), are not conspecific with E. leucozona . They are relatively slender with high spire whorls, bear well defined axial ribs and have an almost smooth last whorl. They are slightly reminiscent of Pusia pseudorecticosta ( Boettger, 1906) but are less elongate. These specimens might represent an undescribed species.

The specimens described by Davoli (2000) from the Tortonian of Montegibbio in Italy differ in their broader outline and wider apical angle (e.g. figs 27–30) and are most probably not conspecific with E. leucozona . Similarly, the specimen from the middle Miocene of the Loire Basin ( France) illustrated by Glibert (1952, pl. 12, figs 2a–b) might represent a separate species.

Iljina (1993) mentioned this species also from the Konkian (middle Miocene) of Agara ( Georgia) and the Ustyurt plateau, which would expand the distribution area also into the Eastern Paratethys Sea. The illustrated specimens, however, differ clearly from Ebenomitra leucozona ( Andrzejowski, 1830) in their slender outline, convex spire whorls blunt axial ribs. It represents a yet undescribed Ebenomitra species.

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).

Distribution in Central Paratethys. Korytnica Basin: Korytnica ( Poland) ( Bałuk 1997); Roztocze Hills: Łychów ( Poland) ( Bałuk 1997); Nowy Sącz Basin: Niskowa ( Poland) ( Bałuk 1970); Voronyaky Hills: Podhorce (Pidhirtsi) ( Ukraine) ( Friedberg 1911); Ukrainian Fore-Carpathian Basin: Dryszczów (Nadrichne), Tarnoruda (Tarnoruda), Ternopol (Ternopil), Żabiak, Zaleśce (Salisze), Zborów (Zboriv), Żukowce (Zhukivtsi) ( Ukraine) ( Friedberg 1911); North Alpine Carpathian Foredeep: Grund, Windpassing ( Sieber 1947, 1953); Vienna Basin: Baden, Gainfarn, Enzesfeld, Vienna /Pötzleinsdorf, Steinebrunn ( Austria), Mikulov ( Czech Republic), Borský Mikulás ( Slovakia) ( Hoernes & Auinger 1880; Švagrovský 1982); Eisenstadt-Sopron Basin: Forchtenau ( Austria) ( Sieber 1956); Pannonian Basin: Sámsonháza, Letkés, Illés-Street Budapest ( Hungary); Krka Basin: Stara vas pri Šentjerneju ( Slovenia) ( Mikuž 2009); Mehadia Basin ( Romania) ( Hinculov, 1968); Zarand Basin: Minişul de Sus ( Romania) ( Nicorici & Sagatovici 1973); Făget Basin: Lăpugiu de Sus ( Romania); Dacian Basin: Staropatica, Târnene ( Bulgaria) (Kojumdgieva in Kojumdgieva & Strachimirov 1960).

Figures

FIGURE 1. A. Map of central and southeastern Europe, representing the area that was partly covered by the Central Paratethys Sea (for detailed maps with the localities mentioned in the text see Kroh (2005) and Nosowska (2020). The white insert corresponds to the area that is restored in the palaeogeographic map below. B. Paleogeography of the Middle Miocene Paratethys Sea (modified from Harzhauser et al. 2007) showing the most important sedimentary basins (NAFB: North Alpine Foreland Basin, E–Sopr B: Eisenstadt–Sopron Basin).

FIGURE 2. Measurements to shell morphology and whorl profiles. SL: shell length, MD: maximum diameter, AA: apical angle, LWH: last whorl height, AH: aperture height. AL: aperture length. AW: aperture width. S: length of siphonal canal.

FIGURE 4. Protoconchs of Paratethyan Costellariidae. A. Antithala filipescui nov. sp., Lăpugiu de Sus (Romania), NHMW 1876/0011/0012). B. Antithala sturi (Hoernes & Auinger, 1880), Lăpugiu de Sus (Romania), NHMW 2020/0058/0001. C. Bellardithala kovaci nov. sp., Steinebrunn (Austria), NHMW 2020/0062/0001. D. Bellardithala partschi (Hörnes, 1852), Steinebrunn (Austria), NHMW 2020/0060/0003. E. Ebenomitra leucozona (Andrzejowski, 1830), Lăpugiu de Sus (Romania), NHMW 1876/0011/0008. F. Ebenomitra pseudopyramidella (Boettger, 1906), Steinebrunn (Austria), NHMW 1857/0014/0009. G. Fedosovia fuchsi (Hoernes & Auinger, 1880), lectotype, Lăpugiu de Sus (Romania), NHMW 1857/0024/0012. H. Pusia avellanella (Boettger, 1906), Gainfarn (Austria), NHMW 2020/0072/0001. I. Pusia? confunda nov. sp., holotype, Coşteiu de Sus (Romania), SMF 360341. J. Pusia pseudorecticosta (Boettger, 1906), Pusia pseudorecticosta (Boettger, 1906), Steinebrunn (Austria), NHMW 2020/0076/0005. K. Pusia schafferi (Meznerics, 1933), Steinebrunn (Austria), NHMW 2020/0078/0002. L. Pusia vexans (Boettger, 1902), Coşteiu de Sus (Romania), NHMW 2020/0081/0001. M. Vexillum intermittens (Hoernes & Auinger, 1880), paralectotype, Baden or Bad Vöslau (Austria), NHMW 2020/0087/0003. N. Vexillum michelottii (Hörnes, 1852), lectotype, Baden (Austria), NHMW 1851/0013/0001. O. Tosapusia pseudocupressina Bałuk, 1997, Lăpugiu de Sus (Romania), NHMW 1870/0033/0037.

FIGURE 10 A1–A2. Ebenomitra leucozona (Andrzejowski, 1830), Lăpugiu de Sus (Romania), NHMW 1870/0033/0034. B1– B2. Ebenomitra leucozona (Andrzejowski, 1830), Gainfarn (Austria), NHMW 1846/0037/0110. C1–C2. Ebenomitra leucozona (Andrzejowski, 1830), Lăpugiu de Sus (Romania), NHMW 1868/0001/0401. D1–D2. Ebenomitra pseudopyramidella (Boettger, 1906), Ternopil (Ukraine), NHMW 2020/0071/0003. E1–E2. Ebenomitra pseudopyramidella (Boettger, 1906), holotype, Ternopil (Ukraine), NHMW 2020/0071/0001. F1–F2. Ebenomitra pseudopyramidella (Boettger, 1906), Ternopil (Ukraine), NHMW 2020/0071/0002. G1–G2. Ebenomitra januszkiewiczi (Friedberg, 1928), lectotype, Zboriv (Zborów) (Ukraine), PAN A-I-50/35. H1–H2. Ebenomitra januszkiewiczi (Friedberg, 1928), paralectotype, Zboriv (Zborów) (Ukraine), ZNG PAN A-I-50/36.1-36.6. I1–I2. Ebenomitra januszkiewiczi (Friedberg, 1928), paralectotype, Zboriv (Zborów) (Ukraine), ZNG PAN A-I-50/36.1-36.6. J. Ebenomitra januszkiewiczi (Friedberg, 1928), paralectotype, Zboriv (Zborów) (Ukraine). ZNG PAN A-I-50/36.1-36.6.

FIGURE 11 A1–A2. Ebenomitra leucozona (Andrzejowski, 1830), Steinebrunn (Austria), NHMW 1857/0014/0008. B1–B2. Ebenomitra pseudopyramidella (Boettger, 1906), Ternopil (Ukraine), NHMW 2020/0071/0004.