Helminthia tricincta (Borson, 1821)

Figs 6I, 13A, 13B 1 – B 2, 13C

* Turritella tricincta nob. — Borson, 1821: 342, pl. 6, fig. 11.

[ Turritella] Riepelii Partsch — Hauer, 1837: 420, nr. 121 [nomen nudum].

[ Turritella] Riepelii Partsch — Hörnes, 1849: 21 [nomen nudum].

[ Turritella] Riepeli— Naumann, 1853: pl. 69, fig. 18.

Turritella Riepeli Partsch — Hörnes, 1855: 421, pl. 43, fig. 2.

Turritella Riepeli Partsch — Neugeboren, 1856: 96 .

Turritella (Eurotropis) Riepeli Partsch — Handmann, 1882: 212 .

? Turritella (Eutrops) Gainfarnensis nov. f. — Handmann, 1882: 218 [nomen nudum].

[ Turritella] Haustator tricinctus (Bors.) — Sacco, 1895: 25, pl. 2, fig. 28).

Turritella (Haustator) tricincta Bors. — Sieber, 1949: 108 .

Turritella (Haustator) tricincta Bors. — Sieber, 1956: 238 .

Turritella (Haustator) tricincta Bors. — Sieber, 1958b: 147 .

Turritella (Haustator) tricincta tricincta Bors. — Sieber, 1960: 245, pl. 3, fig. 8/12.

Turritella (Torculoidella) tricincta Borson 1821 — Kojumdgieva in Kojumdgieva & Strachimirov, 1960: 116, pl. 32, fig. 18.

Turritella vermicularis Brocchi, 1814 — Strausz, 1966: 94, pl. 3, fig. 3, pl. 78, figs 2–3 [non Helminthia vermicularis (Brocchi, 1814)].

Turritella (H.) triplicata Brocc. — Stancu, I. & Andreescu, 1968: 462, pl. 3, fig. 26 [non ‘ Turritella ’ triplicata (Brocchi, 1814)].

Turritella cf. riepeli Partsch. — Csepreghy-Meznerics, 1969: 20, pl. 2, fig. 9

Turritella (Haustator) tricincta Borson, 1821 — Pavia, 1976: 152, pl. 1, fig. 10.

Turritella (Archimediella) pythagoraica Hilber — Fotescu, 1976: 115, fig. 14 [non Archimediella pythagoraica (Hilber, 1882)].

Turritella (Haustator) tricincta tricincta Borson, 1821 — Steininger et al., 1978: 332, pl. 2, fig. 5.

Turritella (Torculoidella) tricincta Borson, 1821 — Atanacković, 1985: 95, pl. 22, figs 11–12.

Archimediella Archimediella [sic] tricincta Borson — Stürmer, 1989: 144, pl. 7, figs 8–9, pl. 12, fig. 5.

Turritella tricincta Borson, 1821 — Iljina, 1993: 35, pl. 3, figs 9–10.

Turritella (Haustator) tricincta tricincta Borson — Schultz, 1998: 56, pl. 21, fig. 7.

Helminthia tricincta (Borson, 1821) — Landau et al., 2013: 63.

Turritella riepeli Partsch — Fözy & Szente, 2014: 311, fig. 6.

non Turritella tricincta— Münster, 1841: 119, pl. 13, fig. 21 [placed in Tricarilda Gründel, 1973 by Gründel (2014)]. non Turritella tricincta Bors. — Friedberg, 1909: 22, pl. 14, fig. 15 [= Helminthia vermicularis (Brocchi, 1814)]. non Turritella (Haustator) vermicularis var. tricincta Schff. — Schaffer, 1912: 161, pl. 52, figs 23–25 [= Ptychidia austrorotun- data new nom.]. non Turritella tricincta Bors. — Friedberg, 1914: 350, pl. 20, fig. 15 [= Helminthia vermicularis (Brocchi, 1814)]. non Turritella (Haustator) tricincta (Borson) — Erünal-Erentöz, 1958: 14, pl. 2, figs 1–2 [= Helminthia vermicularis (Brocchi, 1814)]. non Turritella (Haustator) vermicularis tricincta Schaffer — Sieber, 1960: 244, pl. 3, fig. 8/8 [= Ptychidia austrorotundata new nom.]. non Turritella tricincta Borson — Bałuk, 1970: 117, pl. 10, fig. 20 [= Helminthia vermicularis (Brocchi, 1814)]. non Turritella tricincta Borson — Eremija, 1971a: 69, pl. 6, fig. 2 [= Helminthia vermicularis (Brocchi, 1814)]. non Turritella (? Haustator) tricincta Borson, 1821 — Bałuk, 1975: 108, pl. 12, figs 7–9 [= Helminthia vermicularis (Brocchi, 1814)]. non Turritella (Haustator) tricincta Borson, 1821 – Švagrovský 1981: 126, pl. 39, fig. 1 [= Helminthia vermicularis (Brocchi, 1814)]. non Turritella (Torculoidella) tricincta Borson — Atanacković, 1985: 95, pl. 22, figs 11–12 [= Helminthia vermicularis (Brocchi, 1814)]. non Turritella (Haustator) tricincta Borson — Delgado, 1986: 76, pl. 1, fig. 12–13 [= Helminthia vermicularis (Brocchi, 1814)].

Type material. Holotype illustrated in Borson (1821, pl. 6, fig. 11) and Pavia (1976, pl. 1, fig. 10), stored in the Museo Regionale di Scienze Naturali, Torino (BS. 061.04.046) (Merlino & Campanino 2002). Neogene of the Piedmont region in Italy; no detailed locality information is given by Borson (1821).

Studied material. 1 spec. NHMW 1846/0037/0382 (illustrated as Turritella riepeli in Hörnes 1855, pl. 43,fig. 2), 12 spec. NHMW 1997z0178/0758, Steinebrunn (Austria) ; 2 spec. NHMW 1997z0178/1011, Bad Vöslau (Austria) ; 2 spec. NHMW 1868/0001/0071, 5 spec. 2013/0300/0101, Baden Sooss (Austria) ; 2 spec. NHMW 2013/0078/0718, 5 spec. NHMW2007z0078/0021, Baden (Austria) ; 12 spec. NHMW1997z0178/1362, 7spec. NHMW 1846/0037/0382, 1 spec. NHMW 2013/0479/0697, 2 spec. NHMW 2013/0479/0698, 2 spec. NHMW 2013/0479/2268, 6 spec. NHMW 2007z0071/0021, 1 spec. NHMW 2013/0479/2269, 1 spec. NHMW 1970/1396/1287, 1 spec. NHMW 2013/0479/1289, 2 spec. NHMW 1970/1396/1290, 1 spec. NHMW 2011/0161/0107, 2 spec. NHMW 2005z0095/ 0078a, 1 spec. NHMW 1970/1396/1289, Gainfarn (Austria) ; 7 spec. NHMW 1869/0001/0259, Möllersdorf (Austria) ; 3 spec. NHMW 1865/0001/0815, Vienna-Grinzing (Austria) ; 7 spec. NHMW 1846/0037/0382, Mikulov (Czech Republic) ; 10 spec. NHMW 1870/0033/0155, 4 spec. NHMW 1865/0001/0236, Lăpugiu de Sus (Romania) . 1 spec. NHMW 1858/0047/0035, Sedlec (= Porzteich) (Czech Republic) .

Illustrated material. Fig. 13A: NHMW 1870/0033/0155, SL: 105.1 mm, MD: 28.1 mm, Lăpugiu de Sus (Romania); Figs 13B 1 –B 2: NHMW 1846/0037/0382 (illustrated as Turritella riepeli in Hörnes, 1855, pl. 43, fig. 2), SL: 92.3 mm, MD: 24.9 mm, Steinebrunn (Austria); Fig. 13C: NHMW 1858/0047/0035, SL: 36.2, MD: 12.1 mm, Sedlec (= Porzteich) (Czech Republic).

Revised description. Shell medium-sized to large, robust, moderately slender. Protoconch and first teleoconch whorls unknown. Early teleoconch whorls high, moderately convex; B spiral cord at about mid-whorl most prominent; A and C spiral cords slightly weaker; soon accompanied by r, s, t and u secondary spiral cords. Whorl profile becoming faintly angulated at mid-whorl around 8–10 th teleoconch whorl, B spiral cord coinciding with angulation; A spiral cord becoming obsolete; B and C spiral cords developing into broad cords. Whorl profile becoming flatsided with weakly incised suture after about 10 th teleoconch whorl. A, B and C spiral cords developing into broad, low, convex cords separated by interspaces of about same width. A spiral cord separated from adapical suture by shallow concavity; C spiral cord demarcated by abapical suture. Entire surface by delicate spiral threads (around 70 on last whorl). Base almost flat, bearing three to four widely spaced, low spiral cords and numerous spiral threads. Lateral sinus angle steep, ranging around ~23°, moderately deep with maximum in adapical third of whorl; inflection point close below adsutural suture; second, prominent inflection point in abapical quarter of whorl passing via prominent sinus into base. Basal sinus shallow, opisthocyrt. Aperture subquadrate; inner lip grading into thin, broad glossy parietal callus. No inner lirae developed

Shell measurements and ratios. Largest nearly complete specimen: SL = 105.9 mm, MD = 27.1 mm. n = 7 near complete specimens: SL: μ = 97.2 mm (σ = 5.6 mm), MD: μ = 27.1 mm (σ = 4.1 mm), AA: μ = 20.0° (σ = 1.2°), PA: μ = 14.9° (σ = 1.4°). Lateral sinus (n = 7): LS angle = 22.6° (σ = 5.0°), LS p = 0.8 (σ = 0.1), LS d = 5.6 (σ = 0.8).

Discussion. This species has usually been treated as Turritella riepeli Partsch in the Paratethys literature. The name goes back to collection labels written by Paul Maria Partsch (1791–1856) in the k.k. Mineralogisches Hof-Cabinet (now Natural History Museum Vienna). This ‘ in schedis ’ name was subsequently used as nomen nudum by Hauer (1837) and Hörnes (1849). The name was made available unintentionally by Naumann (1852) who provided an illustration of this species. Later, Hörnes (1855) gave a detailed description and discussed the similarity with Turritella tricincta Borson, 1821 . He stated differences in the apical angle and thus separated the two taxa. These differences, however, are pure artefacts due to the poor and sketchy illustration of Borson (1821). A comparison of the Italian holotype of Turritella tricincta with the specimens from Austria clearly shows that they are conspecific. As discussed by Landau et al. (2013: 63) there is a great confusion in the Paratethys literature about the status of Turritella tricincta Borson, 1820 versus Turritella vermicularis Brocchi, 1814 . This was partly caused by the introduction of the new variation name Turritella (Haustator) vermicularis var. tricincta by Schaffer (1912), for the specimens from the Vienna Basin described as T. vermicularis by Hörnes (1855). In later papers, such as Friedberg (1909, 1914) and Bałuk (1970), typical T. vermicularis specimens where described as Turritella tricincta Borson, 1821 (= Turritella riepeli Hörnes, 1855), but the two species occur syntopically at many Paratethyan localities and can be separated easily. Based on shells from the Vienna Basin, the middle Miocene species H. vermicularis differs from H. tricincta in the smaller size, the more slender outline, the deeper sutures and especially in the early whorls, which bear two prominent spiral ribs, whereas the much higher early whorls of H. tricincta have numerous fine spiral threads (see Fig. 13).

Paleoenvironment. The occurrences in offshore clays (Lăpugiu de Sus, Baden, Bad Vöslau, Möllersdorf, Forchtenau) would suggest a preference for middle to outer neritic environments. This contrasts with frequent occurrences in shallow water settings with sea grass meadows (e.g. Gainfarn; Zuschin et al. 2007). Maybe the species had a broad habitat tolerance, or the robust shells have been transported into offshore settings.

Distribution. Central Paratethys. This species was frequently confused with Helminthia vermicularis (Brocchi, 1814) in the Paratethys literature, which makes it difficult to evaluate its distribution. Badenian (middle Miocene): North-Alpine Foreland Basin: Grund (Austria); Vienna Basin: Niederleis, Baden, Grinzing, Bad Vöslau, Steinebrunn, Gainfarn, Möllersdorf (Austria), Mikulov (Czech Republic) (Hörnes 1855, Sieber 1949; 1956; 1960; herein); Mattersburg Basin: Forchtenau, Mattersburg (Austria) (Sieber 1958b, 1960); Pannonian Basin: Illés-Street, Budapest, Szob (Hungary) (Strausz 1966); Transylvanian Basin: Lăpugiu de Sus (Romania) (Neugeboren 1856; Fözy & Szente 2014); Caransebesulu Basin: Rugi, Delineşti (Stancu & Andreescu 1968); Dacian Basin: Staropatica, Opanec (Bulgaria) (Kojumdgieva in Kojumdgieva & Strachimirov 1960). Eastern Paratethys. Tarkhanian (latest early Miocene to early middle Miocene): Tomakivka (Ukraine), Stavropol region (S-Russia) (Iljina 1993). Proto- Mediterranean Sea. Tortonian (late Miocene): Po Basin: Tortona, Codevilla nel Vogherese (?) (Italy) (Sacco 1895); Pliocene: Po Basin: Astigiana (Italy) (Sacco 1895); Pliocene: Majatico at Parma (Italy) (Cocconi 1873).