Oligodia bicarinata (Eichwald, 1830)

Figs 6P, 7H, 16A 1 –A 2, 16B 1 –B 2, 16C 1 –C 2, 16D, 16E 1 –E

* Turritella bicarinata m. — Eichwald, 1830: 220.

Turritella scalaria— von Buch, 1830: 132.

Turritella scalaria Léop. de Buch — Dubois de Montpéreux, 1831: 36, pl. 2, fig. 18.

Turritella Archimedis Alex. Brong. — Dubois de Montpéreux, 1831: 38, pl. 2, figs 21–22 [non Turritella archimedis Brongniart, 1823].

[ Turritella] subarchimedis d’Orb. — d’Orbigny, 1852: 33, nr. 440 [new name for Turritella archimedis Dubois de Montpéreux, 1831: 38, pl. 2, figs 21–22].

Turritella bicarinata— Eichwald, 1852: plate captions, pl. 10, fig. 23.

Turritella bicarinata mihi — Eichwald, 1853: 280.

Turritella bicarinata Eichw. — Hörnes, 1855: 426, pl. 43, figs 10–11.

[ Turritella bicarinata] subarchimedis d’Orb. — Hörnes, 1855: 427, pl. 43, figs 8–9.

[ Turritella bicarinata] scalaria v. Buch — Hörnes, 1855: 427, pl. 43, fig. 12.

Turritella Oligodia bicarinata Eichw. — Handmann, 1882: 217.

Turritella (Oligodia) arenaria n. f. — Handmann, 1882: 220.

Turritella (Belone) anceps n. f. — Handmann, 1882: 221.

Turritella bicarinata Eichw. — Handmann, 1889: 64, pl. 4, fig. 42.

[ Turritella] A. [rchimediella] bicarinata var. subunocincta Sacc. — Sacco, 1895: 15 [new name for Turritella bicarinata Hörnes, 1855, pl. 43, figs 8–9].

Turritella (Archimediella) conospira n. sp. — Boettger, 1907: 166, no. 567.

Turritella bicarinata Eichw. — Schaffer, 1908: 117, pl. 11, fig. 24.

Turritella bicarinata Eichw. — Friedberg, 1909: 261, pl. 3, figs 9–12.

Turritella bicarinata Eichw. var. scalaria Buch — Friedberg, 1909: 262, pl. 3, fig. 16.

Turritella bicarinata Eichw. — Friedberg, 1914: 332, pl. 19, figs 19–21.

Turritella (Archimediella) conospira Boettger — Zilch, 1934: 216, pl. 6, fig. 9.

Turritella bicarinata Eichw. — Friedberg, 1935: 137, pl. 4, figs 8–12.

T. [urritella] scalaria de Buch — Friedberg, 1935: 137, pl. 4, figs 1–7.

Turritella (Eichwaldiella) bicarinata Eichw. var. — Sieber, 1947: 156.

Turritella (Eichwaldiella) bicarinata (Eichw.) — Sieber, 1953: 186.

Turritella (Archimediella) subarchimedis d’Orb. — Strausz, 1954: 40, pl. 1, fig. 13.

Turritella (Torculoidella) bicarinata Eichwald, 1830 — Moisescu, 1955: 125, pl. 10, figs 9, 12.

Turritella (Archimediella) subarchimedis d’Orb. — Csepreghy-Meznerics, 1956: 431, pl. 2, fig. 19.

Turritella (Eichwaldiella) bicarinata Eichw. — Sieber, 1956: 238.

Turritella (Torculoidella) bicarinata Eichw. — Sieber, 1958b: 147.

Turritella (Torculoidella) scalaria scalaria Buch — Sieber, 1958b: 147.

Turritella (Torculoidella) bicarinata bicarinata Eichw. — Sieber, 1960: 257, pl. 2, figs 7, 19, 21, 23, pl. 3, fig. 8/18.

Turritella (Torculoidella) bicarinata levis ssp. n. — Sieber, 1960: 258, pl. 2, fig. 22.

Turritella (Torculoidella) bicarinata taurevanescens Sacco — Sieber, 1960: 258, pl. 2, fig. 25.

Turritella (Torculoidella) bicarinata subunocincta Sacco — Sieber, 1960: 258, pl. 2, fig. 20.

Turritella (Torculoidella) scalaria praescalaria ssp. n. — Sieber, 1960: 259, pl. 2, figs 24, 26 [non nom. pro Turritella bicarinata Hörnes, 1855, pl. 43, fig. 12].

Turritella bicarinata Eichw. — Eremija, 1959: plate captions, pl. 1, fig. 1 (not mentioned in text).

Turritella (Torculoidella) bicarinata var. subarchimedis (Orbigny, 1852) — Kojumdgieva in Kojumdgieva & Strachimirov, 1960: 115, pl. 32, fig. 15 [non Turritella subarchimedis d’Orbigny, 1852].

Turritella bicarinata Eichwald — Strausz, 1962: 24, pl. 2, fig. 13.

Turritella bicarinata Eichwald, (1930 [sic], 1853 — Strausz, 1966: 86, pl. 2, fig. 13.

Turritella (Torculoidella) bicarinata subarchimedis Orbigny, 1852 — Strausz, 1966: 87, pl. 2, figs 7–8 [non Turritella subarchimedis d’Orbigny, 1852].

Turritella (Eichwaldiella) bicarinata orthezensis Tourn. — Brébion, 1967: 297, text-fig. 2 [non Turritella orthezensis Tournouër in de Bouillé, 1875].

Torculoidella bicarinata (Eichwald) — Zelinskaya et al., 1968: 170, pl. 41, fig. 4.

Turritella (Archimediella) subarchimedis d’Orb. — Csepreghy Meznerics, 1969: 19, pl. 2, fig. 6.

Turritella (Torculoidella) bicarinata Eichw. — Eremija, 1971a: 69, pl. 6, fig. 8.

Turritella (Archimediella) subarchimedis d’Orbigny, 1852 – Bohn-Havas, 1973: 1041, pl. 3, fig. 10, pl. 9, fig. 3.

? Turritella (Archimediella) cf. bicarinata percingulata (Sacco, 1895) — Steininger et al., 1973: 401, pl. 3, fig. 13.

Turritella (Eichwaldiella) bicarinata Eichwald, 1850 — Bałuk, 1975: 109, pl. 12, figs 1–6.

Turritella (Eichwaldiella) bicarinata Eichwald, 1850 — Steininger et al., 1978: 332, pl. 3, figs 1–4.

Turritella (Eichwaldiella) scalaria (v. Buch, 1821) — Steininger et al., 1978: 333, pl. 3, fig. 6.

Turritella (Archimediella) bicarinata Eichwald, 1830 — Švagrovský, 1981: 128, pl. 38, fig. 11.

Tirritella [sic] (Eichwaldiella) bicarinata Eichwald, 1830 — Švagrovský, 1982: 16, pl. 4, fig. 6.

Turritella (Torculoidella) bicarinata bicarinata (Eichwald) — Atanacković, 1985: 94, pl. 22, fig. 9.

? Turritella bicarinata Eichw. (?nov. subsp.) — Popov, 1986: 89, pl. 8, fig. 1.

? Turritella bicarinata Eichwald, 1830 — Iljina, 1993: 35, pl. 3, fig. 11.

Turritella (Eichwaldiella) bicarinata bicarinata Eichwald — Schultz, 1998: 56, pl. 21, fig. 14.

Turritella (Eichwaldiella) scalaria praescalaria Sieber — Schultz, 1998: 56, pl. 21, fig. 15.

Turritella (Eichwaldiella) bicarinata (Eichwald, 1830) — Popa & Ianoliu, 2000: 85, pl. 2, fig. 8.

Turritella (Eichwaldiella) bicarinata (Eichwald, 1830) — Harzhauser, 2002: 78, pl. 2, figs 17–19.

Turritella (Eichwaldiella) bicarinata cf. subunocincta Sacco, 1895 — Mikuž, 2009: 12, pl. 2, fig. 22.

Oligodia bicarinata (Eichwald, 1830) — Landau et al., 2013: 61, pl. 5, fig. 10, pl. 55, fig. 8, pl. 79, fig. 2.

Turritella (Eichwaldiella) bicarinata (Eichwald, 1830) — Popa et al., 2014: 9, pl. 1, fig. 9.

Oligodia bicarinata (Eichwald, 1830) — Landau et al., 2018: 193, pl. 16, fig. 1.

non Turritella bicarinata m. (non Eichw.) — Pusch, 1837: 104, pl. 10, figs 3a–b [= Archimediella indigena (Eichwald, 1830)]. non Turritella (Archimediella) bicarinata Eichw. — Cossmann, 1912: 121, pl. 6, figs 29–30 [= Archimediella].

Type material. Edouard Eichwald’s collection (not examined) is stored in the Museum of Paleontology and Stratigraphy at the Faculty of Geology of Saint Petersburg State University, Russia. Eichwald (1830) based his description on numerous syntypes from the Ukrainian localities Szuszkowce (Shushkivtsi), Bilka (Biłka), Zaleśce (Salisze), Poczajów Stary (Staryi Pochaiev), Tarnoruda (Tarnoruda) and Sawadynce.

Studied material. Oligodia bicarinata is extremely abundant and the NHMW collection houses hundreds of specimens. Therefore, the following list is just a small part of the available material: Types and illustrated specimens: 1 spec. NHMW 1855/0045/0960, Baden (Austria), illustrated as Turritella bicarinata var. subarchimedis in Hörnes 1855, pl. 43, fig. 8; 1 spec. NHMW 1851/0002/0061, Grund (Austria), illustrated as Turritella bicarinata var. subarchimedis in Hörnes, 1855, pl. 43, fig. 9; 1 spec. NHMW 1859/0045/0014, Kleinebersdorf (Austria), illustrated in Hörnes, 1855, pl. 43, fig. 10a; 1 spec. NHMW 1855/0002/0089, Steinebrunn (Austria), illustrated in Hörnes, 1855, pl. 43, fig. 10; 1spec. NHMW 1846/0037/0378, Mikulov, (Czech Republic), illustrated in Hörnes, 1855, pl. 43, fig. 11; 1 spec. NHMW 1963/0552/0000, Steinebrunn (Austria), paratype of Turritella (Torculoidella) scalaria praescalaria Sieber, 1960, pl. 2, fig. 7. 1 spec. NHMW 1963/0530/0000, Grund (Austria), illustrated in Sieber 1960, pl. 2, fig. 19; 1 spec. NHMW 1963/0549/0000, Vöslau (Austria), illustrated as Turritella (Torculoidella) bicarinata subunocincta in Sieber, 1960, pl. 2, fig. 20; 1 spec. NHMW 1963/0542/0000, Grund (Austria), illustrated in Sieber 1960, pl. 2, fig. 21; 1 spec. NHMW 1963/0506/0000, Kleinebersdorf (Austria), holotype of Turritella (Torculoi- della) bicarinata levis Sieber, 1960, pl. 2, fig. 22; 1 spec. NHMW 1963/0507/0000, Grund (Austria), illustrated in Sieber, 1960, pl. 2, fig. 23; 1 spec. NHMW 1963/0513/0000, Grund (Austria), paratype of Turritella (Torculoidella) scalaria praescalaria Sieber, 1960, pl. 2, fig. 24; 1 spec. NHMW 1963/0510/0000, Grossnondorf (Austria), illustrated as Turritella (Torculoidella) bicarinata taurevanescens in Sieber, 1960, pl. 2, fig. 25; 1 spec. NHMW 1963/0513/0000, Steinebrunn (Austria), paratype of Turritella (Torculoidella) scalaria praescalaria Sieber, 1960, pl. 2, fig. 26; 1 spec. NHMW 1963/0512/0000, Steinebrunn (Austria), illustrated as Turritella (Torculoidella) scalaria in Sieber 1960, pl. 2, fig. 27. Additional material studied: 10 spec. NHMW 1997z0178/0611, Stetten (Austria); 7 spec. NHMW 2000z0001/0054, Karnabrunn (Austria); 12 spec. NHMW 1997z0178/0753, Würnitz-Hetzmannsdorf (Austria); 3 spec. NHMW 1859/0045/0046, Grossrussbach (Austria); numerous spec. NHMW 1851/0002/0061, 1963/0507/0000, 1963/0530/0000, 1963/0542/0000, Grund (Austria); 7 spec. NHMW 1859/0045/0137, Niederleis (Austria); 19 spec. NHMW 1860/0001/0275a; 13 spec. NHMW 2018/0194/0001, Steinebrunn (Austria); numerous spec. NHMW 1855/0045/0509, 1970/1396/1487, 1985/0083/0144 a, 1997z0178/1364, 2011/0161/0108, 2013/0479/0701–2013/0479/0715, 2013/0479/2273–2013/0479/2281, Gainfarn (Austria); numerous spec. NHMW 1859/0045/0014, 1973/1593/0016, 2007z0088/0012, 2015/0389/0159–2015/0389/0162, Enzesfeld (Austria); numerous spec. NHMW 1997z0178/1008, 1963/0510/0000, 1963/0549/0000, 2010/0004/0405–2010/0004/0414, Bad Vöslau (Austria); NHMW 1855/0045/0960, 1985/0083/0120, 2013/0078/0705–2013/0078/0707, Baden (Austria); NHMW 2012/0017/0260, 2013/0300/0095–2013/0300/0098, Baden-Sooss (Austria); 1 spec. NHMW 1868/0008/0051, Pötzleinsdorf-Vienna (Austria); 1 spec. NHMW 1865/0001/0817, Grinzing (Austria); numerous spec. NHMW 1870/0037/0032, 1970/1396/1284, 2013/0297/0011, 2016/0026/0165–2016/0026/0167, Ritzing (Austria); 2 spec. NHMW 1861/0001/0269, Pöls (Austria); 1 spec. NHMW 1858/0015/0168, Rousínov (Czech Republic); 8 spec. NHMW 1859/0019/0103, Sedlec (= Porzteich) (Czech Republic); 5 spec. NHMW 1860/0001/0276a, Hlohovec (= Bischofswart) (Czech Republic); 11 spec. NHMW1860/0001/0275b, Mikulov (Czech Republic); 9 spec. NHMW 1847/0046/0041, Szob (Hungary); 3 spec. NHMW 2018/0195/0001, Nemeșești (Romania); numer- ous spec. NHMW 1971/1422/0004, 1973/1615/0011, 2016/0177/0297–2016/0177/0302, Lăpugiu de Sus (Roma- nia); 8 spec. NHMW 1870/0001/0277, Coşteiu de Sus (Romania); 5 spec. 2018/0193/0001, Białozórka (Ukraine); 1 spec. NHMW 1843/0009/0021, Szuszkowce (Ukraine); 2 spec. NHMW 1859/0040/0030, Zaleśce (Ukraine); 1 spec. NHMW 1853/0038/041, Korytnica (Poland).

Illustrated material. Fig. 7H: NHMW 2018/0272/0008, Gainfarn (Austria); Figs 16A 1 –A 2: NHMW 1851/0002/0061, SL: 60.0 mm, MD: 16.4 mm, Grund (Austria), illustrated in Hörnes 1855, pl. 43, fig. 9; Figs 16B 1 –B 2: NHMW 1846/0037/0378, SL: 51.5 mm, MD: 13.8 mm, Mikulov, (Czech Republic), illustrated in Hörnes 1855, pl. 43, fig. 11; Figs 16C 1 –C 2: NHMW 1963/0513/0000, SL: 43.4 mm, MD: 13.4 mm, Steinebrunn (Austria), illustrated in Sieber 1960, pl. 2, fig. 26; Fig. 16D: NHMW 1860/0001/0275a, SL: 50.2 mm, MD: 14.8 mm, Steinebrunn (Austria); Figs 16E 1 –E: NHMW 2018/0193/0001, SL: 38.2 mm, MD: 11.2 mm, Bilosirka, Ternopil Oblast, Ukraine.

Description. This species was described in detail by Landau et al. (2013).

Shell measurements and ratios. (n=15): SL = 52.5 mm (σ = 4.2°), MD = 15.4 mm (σ = 1.5°). AA: 25.0° (σ = 3.4°), PA: 14.6° (σ = 1.0°). Lateral sinus (n = 7): LS angle = 16.7° (σ = 3.9°), LS p = 0.8 (σ = 0.06), LS d = 5.1 (σ = 0.80).

Discussion. This species was discussed in detail by Landau et al. (2013); in addition, we observe up to eight inner lirae on the palatal wall and several weak basal lirae on most specimens. The lateral sinus of this species was described and illustrated by Marwick (1957), as ‘shallow, oblique’. This is contrasted by our data, which reveal a moderately deep lateral sinus with weak inflection point below the adapical suture and a much more pronounced inflection point at the d spiral cord, with prosocyrt part, which passes into a shallowly sigmoidal basal sinus. It is possible that Marwick (1957) only had subadult or poorly preserved specimens at hand.

This is the most variable turritellid species in the Paratethyan record. Especially the strength and width of B and d spiral cords, the presence or absence of spiral sculpture on the sutural ramp and the overall profile (slender or stocky) display a striking variability. The morphologic plasticity is expressed by a multitude of variation and subspecies names. The most conspicuous morphotype is the screw-like scalaria -morph, which is characterized by strongly detached (but not fully uncoiled) teleoconch whorls, with deeply incised sutural ramp resulting in a very deeply incised suture. Its B and D spiral cords are extraordinarily raised and united in a flat sided or weakly concave spiral band. Friedberg (1935) discussed transitional morphs and own material (NHMW) documents that bicarinata and scalaria - morphs may co-occur at the same localities with transitional morphs (partly corresponding to Turritella praescalaria Sieber, 1960). Friedberg (1935) considered Turritella scalaria to be a late Badenian descendant of T. bicarinata, which developed in the Carpathian Foreland Basin after the Badenian Salinity Crisis (~13.8 Ma, de Leeuw et al. 2010). This view is incorrect, as the scalaria -morph appears in the Vienna Basin in deposits predating the Badenian Salinity Crisis (e.g. Gainfarn, Steinebrunn). Nevertheless, we are not aware of any early Miocene and early Badenian scalaria -morphs and the first specimens are collected from middle Badenian (upper Langhian) deposits. Moreover, this morphotype seems to be restricted to the Paratethys and was not detected from the Proto-Mediterranean.

Turritella anceps Handmann, 1822 is a junior synonym of O. bicarinata . This taxon was introduced by Handmann (1822) with a Latin description and a short discussion on its relationship with other taxa. Therefore, Turritella anceps Woods in Bosworth, 1922 is a junior homonym of Turritella anceps Handmann, 1822 . The Paleogene specimens from Peru, illustrated by Woods in Bosworth (1922) as T. anceps are surely not turritellids and we refrain from proposing a replacement name without providing a systematic revision.

Paleoenvironment. Ubiquitous in all shallow water deposits of the Karpatian and Badenian of the Central Paratethys but rare in offshore settings. It appears in seagrass assemblages (e.g. Gainfarn, Zuschin et al. 2007) and lagoonal environments (e.g. Korneuburg Basin, Zuschin et al. 2014). Stable isotope data suggest a preference for normal marine conditions without influx of brackish water (Latal et al. 2006).

Distribution. Central Paratethys. Karpatian (early Miocene): Korneuburg Basin: Stetten, Karnabrunn, Kleinebersdorf, Würnitz-Hetzmannsdorf, Grossrussbach, Niederkreuzstetten (Austria) (Sieber 1960; Harzhauser 2002; herein); Badenian (middle Miocene): North Alpine Foreland Basin: Grund, Immendorf, Windpassing, Niederleis, Grossnondorf (Austria) (Sieber 1947; 1960; herein), Rousínov, Sedlec (Czech Republic) (herein); Carpathian Foreland Basin: Bialozurka, Biłka, Sawadynce, Szuszkowce, Tarnoruda, Zalisce, Poczajów Stary (Ukraine) (Eichwald 1830; Friedberg 1909, 1914); Korytnica, Przeciszów, Małoszów, Gartatowice, Pińczów, Szczaworyż, Benczyn, Bogucice, Zgłobice, Grudna Dolna (Poland) (Bałuk 1975); Badenian (middle Miocene): Vienna Basin: Steinebrunn, Gainfarn, Enzesfeld, Bad Vöslau, Baden, Baden-Sooss, St. Veit, Traiskirchen, Pfaffstätten, Pötzleinsdorf-Vienna, Grinzing-Vienna (Austria) (Hörnes 1855; Sieber 1953, 1960; herein), Hlohovec, Mikulov (Czech Republic), Borský Mikuláš, Devínska Nová Ves (Slovakia) (Švagrovský 1981, 1982); Oberpullendorf Basin: Ritzing (Austria) (Janoschek 1931); Styrian Basin: Pöls, Weitendorf, Wetzelsdorf (Strausz 1946; herein); Krško Basin: Ivanji dol (Slovenia) (Mikuž 2009); Pannonian Basin: Szob, Márkó, Bánd, Herend, Várpalota, Hidas (Strausz 1966; Bohn- Havas 1973; herein); Transylvanian Basin: Nemeşeşti, Lăpugiu de Sus, Coşteiu de Sus (Romania) (Popa et al. 2014; herein); southern Pannonian Basin: Glina (Croatia) (Eremija 1959), Hrvaćanim Reljevac (Bosnia and Herzegovina) (Atanacković 1985); Prnjavor Basin (Bosnia and Herzegovina) (Eremija 1971a); Dacian Basin: Târnene, Trifonovo (Bulgaria) (Kojumdgieva in Kojumdgieva & Strachimirov 1960). An Ottnangian or Karpatian occurrence from Hungary, mentioned by Steininger et al. (1973), is doubtful due to the very poor preservation. Eastern Paratethys. Tschokrakian and Konkian (middle Miocene): Azerbaijan, Ukraine, S-Russia (Popov 1986; Iljina 1993). Proto- Mediterranean Sea and northeastern Atlantic. Burdigalian (early Miocene): Torino Hills: Val Sanfà, Valle Ceppi (Italy) (Zunino & Pavia 2009), Kasaba Basin (Turkey) (İslamoğlu 2004). Langhian (middle Miocene): Loire Basin: Pontlevoy, Manthelan, Bossée, La Chapelle-Blanche, Saint-Maure, Sainte-Catherine de Fierbois, Sepmes, Paulmy, Ferrière-Larçon, Charnizay (France) (Glibert 1949); Torino Hills: Monte de Cappuccini (Italy) (Zunino & Pavia 2009), Kasaba Basin (Turkey) (İslamoğlu 2004). Serravallian (middle Miocene): Karaman Basin: Lale, Akkboğaz, Akpýnar, Seyithasan, Tarlaören (Turkey) (Landau et al. 2013). Tortonian (late Miocene): Po Basin: Stazzano (Italy) (Sacco 1895); Antalya Basin (Turkey) (İslamoğlu & Taner 2003); NW France (Landau et al. 2018)