Helminthia vermicularis (Brocchi, 1814)

Figs 6K, 7 G–H, 13D 1 –D 2,13E 1 –E 2, 13F

* Turbo vermicularis nob. — Brocchi, 1814: 372, pl. 6, fig. 13.

Turritella vermicularis Brocch. var. — Hörnes, 1855: 422, pl. 43, figs 17–18.

Turritella Helminthia vermicularis Brocc. forma typica— Handmann, 1882: 214.

[ Turritella (Helminthia) vermicularis Brocc.] var. conica— Handmann, 1882: 214 [non Turritella conica Klipstein, 1843].

[ Turritella (Helminthia) vermicularis Brocc. var.] conica subv. quadricarinata— Handmann, 1882: 214 [non Turritella quadricarinata (Brocchi, 1814)].

[ Turritella (Helminthia) vermicularis Brocc. var.] conica subv. tricarinata— Handmann, 1882: 214 [non Turritella tricarinata King, 1832, nec Turritellinella tricarinata (Brocchi, 1814)].

[ Turritella (Helminthia) vermicularis Brocc.] var. imminuta— Handmann, 1882: 214.

[ Turritella (Helminthia) vermicularis Brocc. var.] imminuta subv. quadricarinata— Handmann, 1882: 214 [non Turritella quadricarinata (Brocchi, 1814)].

[ Turritella (Helminthia) vermicularis Brocc.] var. dilatata— Handmann, 1882: 214.

[ Turritella (Helminthia) vermicularis Brocc.] var. contorta— Handmann, 1882: 214.

[ Turritella (Helminthia) vermicularis Brocc. var.] contorta subv. tricarinata— Handmann, 1882: 215 [non Turritella tricarinata King, 1832, nec Turritella tricarinata (Brocchi, 1814)].

[ Turritella (Helminthia) vermicularis Brocc. var. contorta subv.] quadricarinata— Handmann, 1882: 215 [non Turritella quadricarinata (Brocchi, 1814)].

[ Turritella (Helminthia) vermicularis Brocc. var. contorta subv.] subdivisa— Handmann, 1882: 215.

[ Turritella (Helminthia) vermicularis Brocc.] var. tricarinata— Handmann, 1882: 214 [non Turritella tricarinata King, 1832, nec Turritella tricarinata (Brocchi, 1814)].

Turritella tricincta Bors. — Friedberg, 1909: 22, pl. 14, fig. 15 [non Turritella tricincta Borson, 1821].

Turritella (Haustator) vermicularis var. tricincta Schff. — Schaffer, 1912: 161, pl. 52, figs 23–24 [non Turritella tricincta Borson, 1821] (non fig. 25 = Helminthia doublierii Matheron, 1842).

Turritella tricincta Bors. — Friedberg, 1914: 350, pl. 20, fig. 15 [non Turritella tricincta Borson, 1821].

Turritella (Haustator) aff. vermicularis Br. — Csepreghy-Meznerics, 1956: 382, pl. 2, figs 26–27.

Turritella (Haustator) vermicularis Brocc. — Sieber, 1947: 156.

Turritella (Haustator) vermicularis Brocc. var. tricincta Schff. — Sieber, 1949: 108 [non Turritella tricincta Borson, 1821].

Turritella (Haustator) vermicularis tricincta Schff. — Sieber, 1953: 185 [non Turritella tricincta Borson, 1821].

Turritella (Haustator) vermicularis tricincta Schff. — Sieber, 1958b: 147 [non Turritella tricincta Borson, 1821].

Turritella (Haustator) vermicularis (Brocchi 1814) — Hölzl, 1958: 184, pl. 17, fig. 7.

Turritella (Haustator) vermicularis tricincta Schaffer — Sieber, 1960: 244, pl. 3, fig. 8/8 [non Turritella tricincta Borson, 1821].

Turritella (Haustator) vermicularis (Brocchi) — Răileanu & Negulescu, 1964: 176, pl. 14, fig. 5.

Turritella (Haustator) vermicularis planulata Sacco — Kókay, 1967: 89, pl. 1, figs 3–4.

Turritella tricincta Borson — Bałuk, 1970: 117, pl. 10, fig. 20 [non Turritella tricincta Borson, 1821].

Turritella tricincta Borson — Eremija, 1971a: 69, pl. 6, fig. 2 [non Turritella tricincta Borson, 1821].

Turritella (Haustator) aff. vermicularis Brocchi, 1814 — Bohn-Havas, 1973: 1040, pl. 3, fig. 17.

? Turritella (Haustator) vermicularis Brocchi, 1814 — Steininger et al., 1973: 399, pl. 3, figs 11a–b.

Turritella (? Haustator) tricincta Borson, 1821 — Bałuk, 1975: 108, pl. 12, figs 7–9 [non Turritella tricincta Borson, 1821].

Turritella (Haustator) tricincta Borson, 1821 – Švagrovský, 1981: 126, pl. 39, fig. 1 [non Turritella tricincta Borson, 1821].

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

Archimediella Archimediella [sic] vermicularis trivolvata n. ssp. — Stürmer, 1989: 140, pl. 8, figs 7–9, pl. 12, fig. 3. [new nom. pro tricincta Schaffer, 1912 non Borson, 1821; nomen nudum).

Turritella (Haustator) vermicularis tricincta Schaffer — Schultz, 1998: 56, pl. 21, fig. 5 [non Turritella tricincta Borson, 1821].

Turritella vermicularis (Brocchi, 1814) — Landau et al., 2011: 11, pl. 2, fig. 9 (with additional synonyms).

Helminthia vermicularis (Brocchi, 1814) — Landau et al., 2013: 62, pl. 5, fig. 11 (with additional synonyms).

non Turritella vermicularis Brocchi, 1814 — Strausz, 1966: 94, pl. 3, fig. 3, pl. 78, figs 2–3 [= Helminthia tricincta (Borson, 1821)].

non Turritella (Haustator) vermicularis Brocchi — Tiţă, 2007: 548, fig. 6/f.

Type material. Holotype (according to Rossi Ronchetti 1955) illustrated in Brocchi (1814, pl. 6, fig. 13) and Rossi Ronchetti (1955: 109, fig. 50); Inv. Nr. 176; Museo Civico di Storia Naturale di Milano (Italy); Pliocene, Piacentino (Italy).

Studied material. 1 spec. NHMW 1862/0001/0391 (illustrated as Turritella vermicularis in Hörnes 1855, pl. 43, fig. 18), 12 spec. NHMW 1846/0037/0387, 15 spec. NHMW 1997z0178/1361, 1 spec. 2018/0272/0007, Gainfarn (Austria) ; 1 spec. NHMW 1868/0001/0091, Nussdorf-Vienna (Austria) ; 1 spec. NHMW 1860/0001/0275a (illustrated as Turritella vermicularis in Hörnes, 1855, pl. 43, fig. 17), 6 spec. NHMW 1860/0001/0265, Steinebrunn (Austria) ; 20 spec. NHMW 1865/0001/0821, Grinzing (Austria) ; 1 spec. NHMW 2018/0179/0001, Windpassing (Austria) ; 6 spec. NHMW 1865/0001/0950, 1 spec. NHMW 2018/0180/0001, Pötzleinsdorf-Vienna (Austria) ; 1 spec. NHMW 2018/0181/0001, 1 spec. NHMW 2018/0181/0010, Lăpugiu de Sus (Romania) ; 3 spec. NHMW 1854/0002/0058, Palermo (Italy) .

Illustrated material. Fig. 7G: NHMW 2018/0272/0007, Gainfarn (Austria); Fig. 7H: NHMW 2018/0181/0010, Lăpugiu de Sus (Romania); Figs 13D 1 –D 2: SL: 80.5 mm, MD: 19.8 mm, NHMW 1860/0001/0275a, Steinebrunn (Austria) (illustrated in Hörnes 1855, pl. 43, fig. 17); Figs 13E 1 –E 2: SL: 75.4 mm, MD: 19.6 mm, NHMW 1862/0001/0391, Gainfarn (Austria) (illustrated in Hörnes 1855, pl. 43, fig. 18); Fig. 13F: SL: 64.9 mm, MD: 18.9 mm, NHMW 2018/0181/0001, Lăpugiu de Sus (Romania).

Revised description. Shell medium-sized, robust, moderately slender. Protoconch and first teleoconch whorls unknown. Earliest preserved whorls weakly angulated with prominent B spiral cord and weaker C spiral cord, entire surface covered by delicate spiral threads. B and C spiral cords sometimes very prominent resulting in bicarinate profile. Spiral cords developing into three strongly raised, broad, convex cords during 8 th– 10 th teleoconch whorls (A spiral cord usually slightly weaker), separated by narrower, deep, concave interspaces. A spiral cord separated from adapical suture by broad concavity, C spiral cord separated by narrower concavity from abapical suture. Rarely, d and r spiral cords developed. Entire shell surface, including primary spiral, cords covered by densely spaced spiral threads. Whorl profile of adult shell only weakly convex; suture moderately incised. Lateral sinus with steep angle, ranging around ~24°; deep with distinctly adapical LSp; w LS p eak inflection point close below adsutural suture; second, prominent inflection point in abapical quarter of whorl passing via gentle sinus into base. Basal sinus weakly opisthocyrt. Base nearly flat, bearing three to four broad spiral cords. Aperture subquadrate, higher than wide; inner lip narrow passing into broad, thin, glossy parietal callus. Inner lirae present on late teleoconch whorls, consisting of up to 10 densely spaced palatal lirae and six basal lirae (occasional).

Shell measurements and ratios. Largest nearly complete specimen: SL = 81.5 mm, MD = 20.0 mm; n = 6 near complete specimens: SL: μ = 81.0 mm (σ = 6.6 mm), MD: μ = 20.1 mm (σ = 2.4 mm), AA: μ = 19.2° (σ = 1.7°), PA: μ = 14.0° (σ = 0.5°). Lateral sinus (n = 5): LS angle = 23.7° (σ = 1.1°), LS p = 0.76 (σ = 0.05), LS d = 4.65 (σ = 0.23).

Discussion. Helminthia vermicularis has been frequently confused with H. tricincta (see Landau et al. 2013), which occur together at many Paratethyan localities. Both species develop very similar lateral sinus morphologies, and both are characterized by the development of three prominent spiral cords and densely spaced spiral threads, covering also the primary ribs. Nevertheless, they are distinguished easily by the larger size and weakly incised suture of H. tricincta and the much more raised spiral cords of H. vermicularis .

Paleoenvironment. This species is very common in shallow water settings, partly indicating sea grass meadows (e.g. Gainfarn; Zuschin et al. 2007).

Distribution. Central Paratethys. As discussed by Landau et al. (2013) this species was frequently confused with Helminthia tricincta (Borson, 1821) in the Paratethys literature, which makes it difficult to evaluate its dis- tribution. Early Miocene Paratethyan occurrences are dubious: e.g.: Ottnangian (early Miocene):? North-Alpine Foreland Basin: Mettbach (Austria) (Sieber 1960), Pannonian Basin:?Várpalota (Hungary) (Kókay 1967). Badenian (middle Miocene): North-Alpine Foreland Basin: Grund, Guntersdorf, Windpassing, Immendorf, Braunsdorf, (Austria) (Sieber 1947, 1949, 1960); Vienna Basin: Gainfarn, Steinebrunn, Grinzing, Vienna (Ottakring, Heiligenstadt, Nussdorf, Grinzing, Pötzleinsdorf), Baden (Austria) (Sieber 1953, 1958b, 1960), Devínska Nová Ves (Slovak Republic) (Švagrovský 1981), Mikulov, Kienberk (Czech Republic) (Švagrovský 1981); Mattersburg Basin: Forchtenau, Marz (Sieber 1960); Carpathian Foreland Basin: Niskowej, Korytnica (Poland) (Bałuk 1970, 1975); Mecsek Mountains (Hungary) (Bohn-Havas 1973); southern Pannonian Basin: Hrvaćanim Reljevac (Bosnia and Herzegovina) (Atanacković 1985); Prnjavor Basin (Bosnia and Herzegovina) (Eremija 1971a). Eastern Paratethys. Records from the Tarkhanian (early middle Miocene) of the Nakhichevan Autonomous Republic will need verification (Popov 1986). Proto-Mediterranean Sea and northeastern Atlantic. Burdigalian (early Miocene): Torino Hills (Italy) (Sacco 1895); late Burdigalian-Langhian: Kasaba Basin (Turkey) (İslamoğlu 2004); Serravallian (middle Miocene): Karaman Basin (Turkey) (Landau et al. 2013); Tortonian (late Miocene): Guadalquivir Basin: Cantillana (southern Spain) (Wenz 1942); Po Basin (Italy) (Sacco 1895), Tunisia (Stchépinsky 1938). Early Pliocene: Guadalquivir Basin, Spain (Landau et al. 2011), Morocco (Brébion 1979); Estepona Basin (Spain) (Landau et al. 2004), northeastern Spain, (Martinell 1979), Roussillon Basin (France) (Fontannes 1879); central Mediterranean, Italy (Bernasconi 1990), Tunisia (Fekih 1975). Early-late Pliocene: Mondego Basin (Portugal) (Zbyszewski 1957), Morocco (Lecointre 1952); central Mediterranean, Italy (Malatesta 1974; Cavallo & Repetto 1992). Early Pleistocene: central Mediterranean, Italy (Landau et al. 2013).