Ptychidia vindobonensis (Handmann, 1882)

Figs 6U, 7J, 18A 1 –A 2, 18B 1 –B 2, 18C, 18D, 18O

Turritella Vindobonensis Partsch — Hörnes, 1849: 21 (pars), no. 302 [nomen nudum].

Turritella turris Bast. — Hörnes, 1855: 423 (pars), pl. 43, figs 15–16 [non Turritella turris de Basterot, 1825].

? Turritella Hörnesi Neugeboren — Neugeboren, 1856: 97.

* Turritella (Ptychidia) Vindobonensis Partsch forma typica— Handmann, 1882: 215 [new name for Turritella turris Hörnes, 1855, pl. 43, fig. 15].

[ Turritella (Ptychidia) Vindobonensis Partsch] var. vermicularis— Handmann, 1882: 215 [non Helminthia vermicularis (Brocchi, 1814)].

[ Turritella (Ptychidia) Vindobonensis Partsch] var. quadricarinata— Handmann, 1882: 215 [non Turritella quadricarinata (Brocchi, 1814)].

[ Turritella (Ptychidia) Vindobonensis Partsch] var. tricarinata— Handmann, 1882: 215 [new name for Turritella turris Bast. in Hörnes, 1855, pl. 43, fig. 16; non Turritella tricarinata (Brocchi, 1814)].

[ Turritella (Ptychidia) Vindobonensis Partsch] var. consolida— Handmann, 1882: 215.

[ Turritella (Ptychidia) Vindobonensis Partsch] var. exsuperans— Handmann, 1882: 215.

[ Turritella (Ptychidia) Vindobonensis Partsch] var. complanata— Handmann, 1882: 215.

[ Turritella (Ptychidia) Vindobonensis Partsch] var. Archimedica— Handmann, 1882: 216.

[ Turritella (Ptychidia) Vindobonensis Partsch] var. Semiarchimedis— Handmann, 1882: 216.

Turritella Vindobonensis Partsch — Handmann, 1888: 19, pl. 4, fig. 41.

[ Turritella] T. [urritella] turris cf. var. Badensis Sacc. — Sacco, 1895: 3 [new name for Turritella turris Hörnes, 1856, pl. 43, figs 15–16, non de Basterot, 1825].

Turritella Turris Bast. — Schaffer, 1908: 116, pl. 11, fig. 23.

Turritella turris var. badensis Sacco — Friedberg, 1909: 255, pl. 2, figs 3–4. Turritella turris var. sexcincta mihi — Friedberg, 1909: 255, pl. 2, fig. 5 [non Turritella sexcincta Goldfuss, 1841]. Turritella turris var. oligocincta mihi — Friedberg, 1909: 255, pl. 2, figs 6–7. Turritella turris var. duplicata mihi — Friedberg, 1909: 255, pl. 2, figs 8–9 [non Zaria duplicata (Linnaeus, 1758)]. Turritella turris var. tricincta mihi — Friedberg, 1909: 255, pl. 2, fig. 10 [non Turritella tricincta Borson, 1821] Turritella Pythagoraica Hilb. var. infirmior mihi. — Friedberg, 1909: 259, pl. 3, fig. 1. Turritella turris Bast. — Friedberg, 1914: 327, text-fig. 70, pl. 19, figs 14–15 [non Turritella turris de Basterot, 1825]. Turritella turris var. sexcincta Friedb. — Friedberg, 1914: 329 [non Turritella sexcincta Goldfuss, 1841]. Turritella turris var. oligocincta Friedb. — Friedberg, 1914: 329, text-fig. 71. Turritella turris var. duplicata Friedb. — Friedberg, 1914: 330 [non Zaria duplicata (Linnaeus, 1758)]. Turritella turris Bast. var. gracilis Friedb. — Friedberg, 1914: 330, pl. 19, fig. 16 [new name for Turritella turris var. tricincta Friedberg, 1909 non Turritella tricincta Borson, 1821], [non Turritella gracilis von Klipstein, 1843, nec Jeffreys, 1867]. Turritella (Haustator) turris Bast. var. — Sieber, 1947: 156. Turritella (Haustator) turris Bast. — Sieber, 1949: 108. Turritella (Haustator) turris badensis (Scc.) — Sieber, 1953: 185. Turritella (Haustator) badensis badensis Sacc. — Strausz, 1954: 13, pl. 1, fig. 15. Turritella (Archimediella) turis [sic] Basterot, 1825 — Moisescu, 1955a: 123, pl. 10, fig. 11 [non Turritella turris de Basterot, 1825]. Turritella (Haustator) badensis Sacco — Csepreghy-Meznerics, 1956: 381, pl. 2, fig. 17. Turritella (Haustator) turris carinatoides Sacco — Csepreghy-Meznerics, 1956: 382, pl. 2, figs 20–21. Turritella (Haustator) badensis sexcincta Friedberg — Csepreghy-Meznerics, 1956: 382, pl. 2, fig. 30 [non Turritella sexcincta Goldfuss, 1841]. Turritella (Haustator) badensis Sacco — Sieber, 1958b: 147. Turritella turris Basterot — Jovanović & Dolić, 1958: pl. 2, fig. 14 [non Turritella turris de Basterot, 1825]. Turritella turris Basterot — Stevanović & Miloševic, 1959: 95, pl. 4, fig. 2 [non Turritella turris de Basterot, 1825]. Turritella turris Basterot — Eremija, 1959: plate captions, pl. 1, fig. 11 [non Turritella turris de Basterot, 1825]. Turritella (Haustator) badensis badensis Sacco — Sieber, 1960: 239, pl. 2, fig. 2, pl. 3, fig. 8/6. Turritella (Haustator) badensis tricarinata (Handm.) — Sieber, 1960: 240, pl. 1, fig. 18 [non Turritella tricarinata King, 1832]. Turritella (Haustator) badensis carinata ssp. n. — Sieber, 1960: 241, pl. 1, fig. 24, pl. 3, fig. 8/7 [non Turritella carinata Lea, 1833, nec Turritella carinata Münster, 1841]. Turritella (Haustator) badensis Sacco — Báldi, 1960: 58, pl. 1, figs 13a–c. Turritella (Archimediella) turris var. badensis Sacco, 1895 — Kojumdgieva in Kojumdgieva & Strachimirov, 1960: 113, pl. 32, fig. 8. Turritella turris Bast. — Florei, 1961: 633, pl. 7, fig. 54. Turritella (Haustator) turris Basterot — Atanacković, 1963: 69, pl. 11, figs 7–7a. Turritella turris Basterot — Grković, 1963: 119, pl. 1, fig. 4 [non Turritella turris de Basterot, 1825]. Turritella (Haustator) turris carinatoides Sacco — Kókay, 1966: 39, pl. 3, fig. 11. Turritella turris badensis Sacco, 1895 — Strausz, 1966: 99, pl. 3, figs 13-17, pl. 78, fig. 4. Archimediella turris Basterot — Zelinskaya et al., 1968: 170, pl. 41, fig. 2 [non Turritella turris de Basterot, 1825]. Archimediella turris var. gracilis (Friedberg, 1909) — Zelinskaya et al., 1968: 170, pl. 41, fig. 3 [non Turritella gracilis von Klipstein, 1843, nec Jeffreys, 1867]. Archimediella turris badensis Sacco, 1895 — Atanacković, 1969: 197, pl. 9, figs 1–2. Turritella turris badensis Sacco, 1895 — Stancu et al., 1971: 125, pl. 8, fig. 1. Turritella (Archimediella) turris Basterot — Eremija, 1971a: 69, pl. 6, fig. 3 [non Turritella turris de Basterot, 1825]. Turritella (Haustator) badensis Sacco, 1895 — Bohn-Havas, 1973: 1040, pl. 3, fig. 11. Turritella (Haustator) turris carinatoides Sacco, 1895 — Bohn-Havas, 1973: 1040, pl. 3, fig. 12. Turritella (Haustator) badensis Sacco, 1895 — Bałuk, 1975: 105, pl. 12, figs 22–27. Turritella (Haustator) badensis badensis Sacco, 1895 — Steininger et al., 1978: 331. Turritella (Haustator) badensis plana Sieber, 1960 — Steininger et al., 1978: 364, pl. 2, fig. 1 [non Turritella plana Nelson, 1870]. Turritella (Haustator) badensis carinata Sieber, 1960 — Steininger et al., 1978: 364, pl. 2, fig. 2 [non Turritella carinata Lea, 1833]. Turritella (Haustator) badensis tricarinata (Handm.) — Steininger et al., 1978: 364, pl. 2, fig. 3 [non Turritella tricarinata King, 1832]. Turritella (Haustator) badensis tricarinata Sieber — Steininger et al., 1978: 364, pl. 2, fig. 4 [non Turritella tricarinata King, 1832]. Turritella (Haustator) badensis badensis Sacco, 1895 — Švagrovský, 1982: 14, pl. 4, fig. 1. Turritella (Haustator) turris turris Basterot, 1825 — Mikuž, 1982: 58, pl. 1, figs 1–3, 6–8 [non Turritella turris de Basterot, 1825]. Turritella (Haustator) badensis carinata Sieber, 1960 — Mikuž, 1982: 80, pl. 5, fig. 3 [non Turritella carinata Lea, 1833]. Turritella (Haustator) badensis cf. tricarinata (Handmann, 1882) — Mikuž, 1982: 82, pl. 6, fig. 1.

Turritella (Haustator) badensis cf. sexcincta Friedberg, 1909 — Mikuž, 1982: 84, pl. 6, figs 2–6. Turritella (Haustator) badensis cf. gracilis Friedberg, 1914 — Mikuž, 1982: 86, pl. 6, fig. 7. Turritella (Haustator) badensis vajndoliana n. ssp. — Mikuž, 1982: 88, pl. 6, figs 8–9, pl. 7, figs 1a–1b (nomen nudum; unpub- lished PhD dissertation). Turritella (Archimediella) turris Basterot, 1825 — Atanacković, 1985: 93, pl. 21, figs 13–16. Haustator Miohaustator [sic] badensis badensis Sacco — Stürmer, 1989: 120, pl. 4, figs 2–4, pl. 11, figs 2–4. Turritella (Haustator) badensis plana Sieber, 1960 — Mikuž, 1982: 78, pl. 5, fig. 2 [non Turritella plana Nelson, 1870]. Haustator Miohaustator [sic] badensis carinatus Sieber — Stürmer, 1989: 124, pl. 4, fig. 5, pl. 12, fig. 14 [non Turritella carinata Lea, 1833].? Turritella badensis Sacco, 1895 — Iljina, 1993: 35, pl. 3, fig. 12. Turritella (Haustator) badensis Sacco — Jakubowski, 1996: 706, pl. 217, figs 11–12. Turritella (Haustator) badensis tricarinata Handmann — Schultz, 1998: 56, pl. 21, fig. 2 [non Turritella tricarinata King, 1832]. Turritella (Haustator) badensis carinata Sieber — Schultz, 1998: 56, pl. 21, fig. 4 [non Turritella carinata Lea, 1833]. Turritella badensis Sacco — Mikuž, 2003: 308, pl. 10, fig. 22. Turritella (Haustator) turris turris Basterot, 1825 — Mikuž, 2009: 8, pl. 1, fig. 1 [non Turritella turris de Basterot, 1825]. Turritella (Haustator) badensis badensis Sacco, 1895 — Mikuž, 2009: 10, pl. 1, fig. 11. Turritella (Haustator) badensis plana Sieber, 1960 — Mikuž, 2009: 10, pl. 1, fig. 12 [non Turritella plana Nelson, 1870]. Turritella (Haustator) badensis carinata Sieber, 1960 — Mikuž, 2009: 10, pl. 1, fig. 13 [non Turritella carinata Lea 1833]. Turritella (Haustator) badensis cf. tricarinata (Handmann, 1882) — Mikuž, 2009: 11, pl. 1, fig. 14. Turritella (Haustator) badensis cf. sexcincta Friedberg, 1909 — Mikuž, 2009: 11, pl. 1, fig. 15. Turritella (Haustator) badensis cf. gracilis Friedberg, 1914 — Mikuž, 2009: 11, pl. 1, fig. 16. Turritella (Haustator) badensis vajndoliana (n. ssp?) — Mikuž, 2009: 11, pl. 1, figs 17–18. Turritella (Haustator) eryna? d’Orbigny, 1852 — Mikuž, 2009: 9, pl. 1, fig. 4. Turritella (Haustator) eryna communiformis Voorthuysen, 1944 — Mikuž, 2009: 9, pl. 1, figs 5–6. Turritella (Haustator) eryna turriformis Voorthuysen, 1944 — Mikuž 2009: 9, pl. 1, fig. 8. Ptychidia vindobonensis (Handmann, 1882) — Landau et al., 2013: 56, pl. 5, fig. 9, pl. 55, fig. 7, pl. 79, fig. 1. Turritella (Haustator) badensis Sacco, 1895 — Popa et al., 2014: 8, pl. 1, fig. 8.

Type material. Holotype NHMW 1853/0010/0030, illustrated in Hörnes (1855, pl. 43, fig. 15); SL: 67.7 mm, MD: 17.2 mm, middle Miocene, Badenian, Enzesfeld (Austria). The lot from which Hörnes (1855) selected the specimen for illustrations also contained specimens of Archimediella abundans (Handmann, 1882) and Archimediella carpathica . Handmann (1882: 215) explicitly referred to the specimen fig. 15 of Hörnes (1855) as ‘forma typica’, without mentioning additional material. Therefore, this specimen is the holotype of Ptychidia vindobonensis .

Studied material. 1 spec. NHMW 1862/00017/0233, Baden (Austria) , illustrated in Hörnes (1855, pl. 43, fig. 16); 7 spec. NHMW 2018/0223/0001, 1 spec. NHMW 1963/0437/0000, Baden-Sooss (Austria) ; holotype of Turritella badensis carinata Sieber 1960; 36 spec. NHMW 1997z0178/1010, 6 spec. NHMW 1851(0006/0017, Bad Vöslau (Austria) ; 6 spec. NHMW 2018/0223/0003, 16 spec. NHMW 2018/0223/0002, Baden (Austria) ; 8 spec. NHMW 1862/0001/0279, Möllersdorf (Austria) ; 19 spec. NHMW 1871/0010/0381, Steinebrunn (Austria) ; 6 spec. NHMW 1868/0015/0758, Niederleis (Austria) ; 6 spec. NHMW 1868/0051/0019, Perchtoldsdorf (Austria) ; 7 spec. NHMW 1853/0010/0030, including specimen illustrated in Hörnes (1855, pl. 43, fig. 15) ; 5 spec. NHMW 1890/0003/0105; 4 spec. 2018/0224/0001; 4 spec. NHMW 2018/0224/0002; 7 spec. NHMW 1868/0001/0170, Enzesfeld (Austria) ; 17 spec. NHMW 1997z0178/1360; 1 spec. NHMW 1937/0002/0305, illustrated in Schaffer (1908, pl. 11, fig. 23); 3 spec. NHMW 1846/0037/0390; 8 spec. NHMW 2018/0225/0001; 1 spec. NHMW 2018/0225/0002; 15 spec. NHMW 1989/0037/0001; 3 spec. NHMW 1997z0178/1506, Gainfarn (Austria) ; 17 spec. NHMW 1861/0012/0118, Hrušovany nad Jevišovkou (Czech Republic) ; 8 spec. NHMW 1860/0001/0287, Hlohovec (= Bischofswart) (Czech Republic) ; 6 spec. NHMW 1847/0037/0100, Mikulov (Czech Republic) ; 12 spec. NHMW 1860/0001/0280, Coşteiu de Sus (Romania) ; 5 spec. NHMW 1870/0032/0159, Lăpugiu de Sus (Romania) .

Illustrated material. Fig. 7J, Figs 18A 1 –A 2: NHMW 1853/0010/0030, SL: 68.4 mm, MD: 18.0 mm, Enzesfeld (Austria) (holotype); Figs 18B 1 –B 2: NHMW 1862/0017/0233, SL: 70.4 mm, MD: 16.4 mm, Baden (Austria); Fig. 18C: NHMW1963/0527/0000, SL: 64.9 mm, MD: 17.2 mm, Immendorf (Austria), holotype of Turritella (Haustator) badensis carinata Sieber 1960; Fig. 18D: NHMW 1997z0178/1360, SL: 73.4 mm, MD: 19.1 mm, Gainfarn (Austria) ; Fig. 18O: NHMW 1997z0178/1361, Gainfarn (Austria).

Revised description. Shell medium-sized, moderately slender. Protoconch comprising about 2.5 strongly convex whorls. Transition into teleoconch indicated by appearance of B spiral cord in middle of whorl, accompanied by weaker C spiral cord and initially indistinct A spiral cord. Earliest teleoconch whorls tricarinate, high, moderately convex with incised suture, forming very slender apex. Weak r spiral cord may occur on sutural ramp, occasionally accompanied by two tertiary spiral cords. Successively, one or two secondary cords may appear between primaries; similarly, one to three secondaries may be developed below C primary. Whorl profile of later teleoconch very variable ranging from regularly convex to carinate in abapical third (resulting from prominent B spiral cord). Convexity of sutural ramp also variable from moderately convex to nearly flat. Sculpture of post-neanic whorls typically consisting of five to six (if weak d spiral cord is developed) spiral cords with two less prominent secondaries on sutural ramp; B primary spiral cord (or B and C) being strongest. Strength of primaries may vary individually resulting in morphotypes with spiral cords of subequal strength and distance and regularly convex whorls. Lateral sinus angle steep, shallow with singly, weak adapical inflection point; adapical LS p; basal sinus faintly opisthocyrt. Base moderately convex with densely spaced spiral cords. Aperture U-shaped, columella weakly convex, inner lip reflected forming narrow umbilical chink, passing into broad glossy parietal callus, covering large parts of base. Inner lirae appear on post-neanic whorls, consisting of up to eight palatal lirae and three to four weak basal lirae.

Shell measurements and ratios. (n=10): SL = 66.6 mm (σ = 5.1°), MD = 17.0 mm (σ = 0.7°). AA: 18.5° (σ = 1.8°), PA: 13.0° (σ = 1.1°). Lateral sinus (n = 10): LS angle = 22.0° (σ = 4.4°), LS p = 1.2 (σ = 0.2), LS d = 6.5 (σ = 0.9).

Discussion. As pointed out by Bałuk (1975) and Landau et al. (2013), Ptychidia vindobonensis is an extraordinarily variable species and many forms have been described as variations or subspecies. In some cases, the same morphotype was named two or three times: e.g., specimens with prominent carina on post-neanic whorls have been named consolida Handmann, 1882, carinata Sieber, 1960 and vajndoliana Mikuž, 2009. Shells with nearly flat whorls were described as complanata Handmann, 1882 (note that T. badensis plana Sieber, 1960 is Ptychidia erynella new sp.).

Landau et al. (2013) tried to clarify the complex nomenclature of this species and concluded that Turritella (Ptychidia) vindobonensis Handmann, 1882 is the oldest available name. Nevertheless, Turritella hoernesi Neugeboren, 1856, introduced for shells from Lăpugiu de Sus in Romania, might be an even older available name judging from the description in Neugeboren (1856). We are not aware of the whereabouts of the syntypes of Neugeboren and the species has never been illustrated. Thus, the status of Turritella hoernesi Neugeboren, 1856 cannot be clarified at the moment and we treat it as taxon inquirendum. In any case, it would be a junior homonym of Turritella hoernesi Rolle, 1856 [see discussion of Archimediella hoernesi (Rolle, 1856) above].

The Mesozoic mathildoid Turritella complanata Brösamlen, 1909 is a junior homonym of Turritella complanata Handmann, 1882 but was recognized as subjective junior synonym of Gordenella fahrenkohli (Rouillier, 1848) by Gründel & Nützel (2013). Therefore, a replacement name for the Mesozoic taxon is unnecessary.

Paleoenvironment. Ptychidia vindobonensis is a ubiquitous species, which occurs in shallow marine sublittoral, inner neritic settings and in offshore clays of middle to outer neritic environments.

Distribution. Central Paratethys. Badenian (middle Miocene): North Alpine Foreland Basin: Grund, Immendorf, Windpassing (Austria) (Sieber 1947, 1949, 1960); Carpathian Foreland Basin: Korytnica, Biskupice, Małoszów, Gartatowice, Pińczów, Szczaworyż, Staszów, Szydłów, Rybnica, Skoczów, Andrychów, Benczyn, Żegocina, Bogucice, Błonie, Niechobrz (Poland) (Bałuk 1975), Hołubica (Ukraine) (Friedberg 1909, 1914); Vienna Basin: Vienna (Grinzing, Ottakring, Pötzleinsdorf), Gainfarn, Enzesfeld, Bad Vöslau, Baden, Baden-Sooss, Steinebrunn (Austria), Mikulov, Hrušovany nad Jevišovkou, Sedlec (Czech Republic) (Sieber 1953, 1958b, 1960), Borský Mikuláš (Slovakia) (Švagrovský 1982); Eisenstadt-Sopron Basin: Forchtenau (Austria); Oberpullendorf Basin: Ritzing (Austria) (Sieber 1960); Styrian Basin: Wetzelsdorf (Austria) (Sieber 1960); Pannonian Basin: Szob, Szokolya, Pécsvárad, Püspöklak, Herend, Budapest-Káposztásmegyer (Hungary), (Báldi 1960; Kókay 1966; Strausz 1966); Mecsek Mountains (Hungary) (Strausz 1966); Bükk Mountains (Hungary) (Csepreghy Meznerics 1969); Krško Basin: Vajndol, Gorenja Stara Vas Dolenja, Brezovica, Šentjernej (Slovenia) (Mikuž 2009); southern Pannonian Basin: Hrvaćani-Drenik, Glina, Tešnja, Miljevići, Jurkovica (Bosnia and Hercegovina) (Eremija 1959; Grković 1963; Atanacković 1985), Aranđelovac, Tamnava River (Serbia) (Jovanović & Dolić 1958; Stevanović & Miloševic 1959); Transylvanian Basin: Coşteiu de Sus, Buitur, Lăpugiu de Sus, Nemeşeşti (Moisescu 1955a; Popa et al. 2014; herein); southern Carpathians: Dubova (Romania) (Stancu et al. 1971); Dacian Basin: Staropatica, Opanec, Orehovica (Bulgaria) (Kojumdgieva & Strachimirov 1960). Eastern Paratethys. Some records of Ptychidia turris mentioned from the Tarkhanian of the Caucasus region by Popov (1986) might belong here. A specimen described by Iljina (1993) from the Konkian of the Caucasus might be an additional record but is too incomplete for a definite identification. Proto-Mediterranean Sea and northeastern Atlantic. Serravallian (middle Miocene): Karaman Basin: Lale, Akkboğaz, Akpýnar, Tilkikaya, Tavþanlý, Seyithasan, Tarlaören (Turkey) (Landau et al. 2013);? Tortonian (late Miocene): Guadalquivir Basin: Trujillo (southern Spain) (Wenz 1942).