Ptychidia erynella, Harzhauser & Landau, 2019

Harzhauser, Mathias & Landau, Bernard, 2019, Turritellidae (Gastropoda) of the Miocene Paratethys Sea with considerations about turritellid genera, Zootaxa 4681 (1), pp. 1-136 : 67-69

Ptychidia erynella new sp.

Figs 6V , 18E, 18F, 18G, 18H, 18J, 18K, 18M, 18N 1 – N 2

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

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

Turritella (Haustator) cf. eryna Tourn. — Sieber, 1947: 156.

Turritella (Haustator) cf. eryna d’Orb. var. communiformis Voorth. — Sieber, 1949: 108 [non Turritella eryna communiformis van Voorthuysen, 1944 ].

? Turritella (Haustator) turris Basterot, 1825 var. — Hölzl, 1958: 180, pl. 18, fig. 1.

Turritella (Haustator) eryna eryna Orb. — Sieber, 1960: 235, pl. 1, figs 10, 13, pl. 3, fig. 8/2 [non Turritella eryna d’Orbigny, 1852 ].

Turritella (Haustator) eryna turriformis Voorthuysen– Sieber, 1960: 237, pl. 1, fig. 5 [non Turritella eryna turriformis van Voorthuysen, 1944 ].

Turritella (Haustator) eryna communiformis Voorth. — Sieber, 1960: 236, pl. 1, figs 6–7 [non Turritella eryna communiformis van Voorthuysen, 1944 ].

Turritella (Haustator) badensis plana ssp. n. — Sieber, 1960: 240, pl. 1, fig. 32 [non Turritella plana Nelson, 1870 ].

Turritella (Haustator) striatellata Sacco — Sieber, 1960: 242, pl. 1, fig. 19 [non Haustator striatellatus Sacco, 1895 ].

Turritella eryna Orbigny, 1852 — Strausz, 1966: 96, pl. 3, fig. 5.

Turritella (Haustator) eryna eryna Orbigny, 1852 — Tejkal et al., 1967: 195, pl. 10B, figs 1–5 [non Turritella eryna d’Orbigny, 1852 ].

Turritella (Haustator) eryna communiformis Voorthuisen, 1944 — Tejkal et al., 1967: 196, pl. 10B, figs 8–10 [non Turritella eryna communiformis van Voorthuysen, 1944 ].

Turritella (Haustator) eryna communiformis Voorthuisen, 1944 — Steininger et al., 1971: 371 [non Turritella eryna communiformis van Voorthuysen, 1944 ].

Turritella (Haustator) eryna eryna Orbigny, 1852 — Steininger et al., 1973: 397, pl. 3, fig. 8 [non Turritella eryna d’Orbigny, 1852 ].

Turritella (Haustator) eryna communiformis Voorthuisen, 1944 — Steininger et al., 1973: 398 [non Turritella eryna communiformis van Voorthuysen, 1944 ].

Turritella (Haustator) eryna eryna Orbigny, 1852 — Steininger et al., 1978: 311, pl. 1, fig. 8 [non Turritella eryna d’Orbigny, 1852 ].

Turritella (Haustator) eryna? d’Orbigny, 1852 — Mikuž, 1982: 61, pl. 1, figs 4–5 [non Turritella eryna d’Orbigny, 1852 ].

Turritella (Haustator) eryna communiformis Voorthuysen, 1944 — Mikuž, 1982: 66, pl. 1, fig. 9, pl. 2, figs 1a–b [non Turritella eryna communiformis van Voorthuysen, 1944 ].

Turritella (Haustator) eryna turriformis Voorthuysen, 1944 — Mikuž, 1982: 70, pl. 2, fig. 4 [non Turritella eryna turriformis van Voorthuysen, 1944 ].

Turritella (Erynella) eryna eryna d’Orbigny — Stürmer, 1989: 107, pl. 3, figs 4–7, pl. 10, fig. 8 [non Turritella eryna d’Orbigny, 1852 ].

Haustator Miohaustator [sic] badensis planus Sieber — Stürmer, 1989: 123, pl. 4, fig. 1, pl. 10, fig. 4 [non Turritella plana Nelson, 1870 ].

Haustator Miohaustator [sic] striatellatus Sacco — Stürmer, 1989: 126, pl. 4, fig. 7, pl. 10, fig. 10 [non Haustator striatellatus Sacco, 1895 ].

Turritella (Haustator) eryna communiformis Voorthuysen — Schultz, 1998: 56, pl. 21, fig. 1 [non Turritella eryna communiformis van Voorthuysen, 1944 ].

Turritella (Haustator) badensis plana Sieber — Schultz, 1998: 56, pl. 21, fig. 3 [non Turritella plana Nelson, 1870 ].

Turritella eryna d’Orbigny, 1852 — Harzhauser, 2002: 77.

? Turritella eryna eryna d’Orb. — Kókay, 2002: 372, pl. 2, fig. 12, pl. 3, fig. 1 [non Turritella eryna d’Orbigny, 1852 ].

Turritella badensis Sacco, 1895 — Mandic & Harzhauser, 2003: 97, pl. 1, fig. 1 [non Ptychidia vindobonensis ( Handmann, 1882) ].

non Turritella eryna rotundata Schaff. — Kókay, 1972: 48, pl. 1, fig. 2 [probably an Archimediella ].

non Turritella (Haustator) eryna rotundata Schaffer, 1912 — Steininger et al., 1973: 398, pl. 3, figs 7a–c [probably an Archimediella ]. non Turritella (Haustator) eryna turriformis Voorthuysen, 1944 — Švagrovský, 1982: 14, pl. 4, fig. 2.

Types. Holotype NHMW 1963/0522/0000 ( Fig. 18F ), SL: 41.8 mm, 10.4 MD: mm, Guntersdorf ( Austria), illustrated in Sieber (1960, pl. 1, fig. 10) as Turritella (Haustator) eryna eryna d’Orbigny . Paratypes NHMW 1861/0034/0032 ( Fig. 18E ), SL: 55.1 mm, MD: 13.7 mm, Kalladorf ( Austria) ; NHMW 2018/0279/0002 ( Figs 18G 1 –G 2 ), SL: 56.2 mm, MD: 12.2 mm, Guntersdorf ( Austria); NHMW 2019/00277/0003 ( Fig. 18H ), SL: 58.3 mm, MD: 13.9 mm, Guntersdorf ( Austria); NHMW 2018/0277/0002, ( Fig. 18N ), SL: 19.9 mm, MD: 5.1 mm, Grund ( Austria).

Additional illustrated material. Fig. 18J : NHMW 1963/0521/0000, SL: 45.1 mm, MD: 13.4 mm, Grund ( Austria), illustrated in Sieber (1960, pl. 1, fig. 5) as Turritella (Haustator) eryna turriformis van Voorthuysen ; Fig. 18K : NHMW 1963/0520/0000, SL: 44.0 mm, MD: 12.8 mm, Grund ( Austria), illustrated as Turritella (Haustator) eryna communiformis Voorth. in Sieber (1960 , pl. 1, figs 6–7); Figs 18M : NHMW 1963/0543/0000, SL: 43.6 mm, MD: 12.7 mm, Immendorf ( Austria), illustrated as Turritella (Haustator) striatellata Sacco in Sieber (1960 , pl. 1, fig. 19).

Type stratum. Silty sand of the Grund Formation ( Zuschin et al. 2004).

Type locality. Guntersdorf ( Austria) .

Age. Middle Miocene, Badenian.

Etymology. Referring to Turritella eryna d’Orbigny, 1852 , with which the species was frequently confused.

Studied material. 1 spec. NHMW 1861/0034/0032, 3 spec. NHMW 1869/0003/0068, Kalladorf ( Austria); 1 spec. NHMW 2018/0277/0003, 50 spec. NHMW 2018/0277/0004, 1 spec. NHMW 2018/0277/0001, 1 spec. NHMW 2018/0277/0002, 1 spec. NHMW 1963/0521/0000, Grund ( Austria), illustrated in Sieber (1960, pl. 1, fig. 5) as Turritella (Haustator) eryna turriformis van Voorthuysen , 1 spec. NHMW 1963/0520/0000, illustrated in Sieber (1960, pl. 1, figs 6–7) as Turritella (Haustator) eryna communiformis van Voorthuysen , 1 spec. NHMW 1963/0524/0000, illustrated in Sieber (1960, pl. 1, fig. 13) as Turritella (Haustator) eryna eryna d’Orbigny , 2 spec. NHMW 1989/0032/0001, 2 spec. NHMW 1989/0033/0001, Grund ( Austria); 8 spec. NHMW 2018/0279/0001, 1 spec. NHMW 2018/0277/0002, Guntersdorf ( Austria); 1 spec. NHMW 1963/0522/0000, Guntersdorf ( Austria), illustrated in Sieber (1960, pl. 1, fig. 10) as Turritella (Haustator) eryna eryna d’Orbigny ; 20 spec. NHMW 2018/0280/0001, Nodendorf ( Austria); 7 spec. NHMW1861/0050/0056, Karnabrunn ( Austria); 1 spec. NHMW 1860/0050/0114, Laa ( Austria); 7 spec. NHMW 1863/0015/0993, Hrušovany nad Jevišovkou ( Czech Republic); 16 spec. 1845/0031/0053, 1 spec. NHMW 1872/0020/0213, 1 spec. NHMW 1963/0517/0000, Wirtatobel at Bregenz ( Austria), illustrated in Sieber (1960, pl. 1, fig. 26) as Turritella turris Basterot ; 2 spec. NHMW 1872/0020/0196, Kaltenbachgraben ( Germany); Ermingen ( Germany), 1 spec. NHMW1963/0527/0000, Immendorf ( Austria), illustrated as Turritella badensis tricarinata Handmann, 1882 , in Sieber (1960, pl. 1, fig. 18);

Description. Shell medium-sized, slender, of about 18 teleoconch whorls. Protoconch unknown. Early teleoconch whorls high, angulated at mid-whorl with deep suture. Onset of primaries not preserved; angulation coinciding with B spiral cord adjoined by weaker C spiral cord and faint A spiral cord (not always visible) starting at least on 3 rd to 4 th teleoconch whorls. Height of whorls successively decreasing¸ angulation shifting slightly below mid-whorl on neanic whorls. B spiral cord most prominent, ranging from keel-like to convex. C spiral cord shifting down, close to suture, resulting in broad, convex interspace between B and C spiral cords. Sutural ramp flat to weakly convex with distinct A spiral cord, often accompanied by r cord and one or two tertiary spiral cords. Secondary spiral cord between B and C spiral cords, rarely equaling primaries; no additional secondary cords below C spiral cord aside from peribasal cord of variable strength. Lateral sinus simple, steep, maximum depth below mid-whorl, adapical part moderately convex without inflection point, abapical part weakly convex, nearly straight. Aperture subquadratic, higher than broad; outer lip thin, inner lip narrowly reflected firming narrow umbilical chink. Parietal callus glossy, thin, indistinct, covering only small part of base. Basal sinus nearly straight. Base moderately convex with about eight narrow, usually poorly developed, secondary spiral cords. No inner lirae detected in Austrian and French specimens.

Shell measurements and ratios. n = 10 near complete specimens: SL: μ = 53.9 mm (σ = 3.2 mm), MD: μ = 12.7 mm (σ = 0.7 mm), AA: μ = 17.9° (σ = 0.5°), PA: μ = 12.6° (σ = 1.0°). Lateral sinus (n = 8): LS angle = 28.6° (σ = 2.4°), LS p = 1.2 (σ = 0.1), LS d = 4.7 (σ = 1.0).

Discussion. Paratethyan occurrences of this species have been identified as Turritella eryna d’Orbigny, 1952 , which was recorded by Cossmann & Peyrot (1922) from the early Miocene of the Aquitaine Basin. Lozouet et al. (2001), however, considered Turritella eryna d’Orbigny, 1852 and Turritella subtriplicata d’Orbigny, 1852 as synonyms of Turritella turris de Basterot, 1825 . No middle Miocene occurrences have been reported by Cossmann & Peyrot (1922) and Lozouet et al. (2001) for these species. Specimens from Léognan and Saucats in France, stored in the NHMW collection and identified as Turritella turris , are indeed similar to Paratethyan specimens in overall shape, but differ in their strongly keeled early neanic whorls. Unlike P. vindobonensis , the initial part of the protoconch of T. turris is depressed and a distinct A spiral cord appears on the first teleoconch whorl [see Fig. 18I : NHMW 1872/0020/0048, SL: 40.8 mm, MD: 12.0 mm, Léognan ( France)].

Sieber (1960) identified some specimens as Turritella eryna communiformis van Voorthuysen, 1944 and T. e. turriformis van Voorthuysen, 1944, which were originally described from the Hemmoorian (late Burdigalian to early Langhian) to Langenfeldian (Serravallian and early Tortonian) of the North Sea (van Voorthuysen 1944). The neanic whorls of the specimens illustrated by van Voorthuysen (1944) are less angulated and differ from the Paratethyan species in their prominent spiral cords on the sutural ramp and the generally broader apical angle, regularly convex whorls and more uniform spiral sculpture. Ptychidia erynella is highly reminiscent of P. vindobonensis in sculpture and whorl profile. In both species ‘ carinata ’ and ‘ plana ’ morphs have been described. Ptychidia vindobonensis , however, is generally larger (about 10 mm at same growth stage) and less slender. Inner lirae are frequent in P. vindobonensis but virtually absent in all investigated P. erynella . Moreover, P. erynella seems to be restricted to the North Alpine-Carpathian Foreland Basin, where it appears in enormous numbers. It is absent from middle Badenian faunas in the Vienna Basin, where P. vindobonensis predominates in numbers.

In most shells, the B spiral cord coincides with a more or less distinct angulation below mid-whorl. The variability of sculpture results mainly from the strength and presence of the secondary and tertiary spiral cords. In some morphotypes, all spiral cords attain similar strengths and were treated as subspecies communiformis van Voorthuysen, 1944 in the literature, whereas specimens with prominent primaries were identified as subspecies turriformis van Voorthuysen, 1944. Both morphotypes co-occur in the rich lower Badenian localities Grund, Immendorf and Guntersdorf in the North Alpine-Carpathian Foreland Basin, including transitional morphs. Some of these morphotypes are close to the specimens from the Ottnangian (early Miocene) of Ermingen ( Germany), where this turritellid forms a mass-occurrence known as ‘Erminger Turritellenplatte’ ( Baier 2008; Höltke 2009). The preservation of these specimens is rather poor, and the species may be closer to P.? turris than to P. erynella .

(see Fig. 18L : NHMW 1845/00031/0053, SL: 45.9, MD: 13.7 mm, Ermingen ( Germany). Sieber (1960) and Stürmer (1989) identified a pathological specimen from Immendorf ( Austria) as Turritella (Haustator) striatellata Sacco, 1895 ( Fig. 18M ). The early teleoconch whorls of this specimen do not differ from other specimens of P. erynella from the same locality and the subsequent loss of spiral sculpture is caused by severe shell damage at an early stage of growth.

Paleoenvironment. The Paratethyan occurrences derive from inner neritic, coastal and lagoonal settings (e.g. Korneuburg Basin and North Alpine Foreland Basin, Harzhauser et al. 2002; Zuschin et al. 2004). At many localities, it occurs in huge numbers forming “turritelline-dominated assemblages” sensu Allmon & Knight (1993).

Distribution. Western and Central Paratethys. Karpatian (early Miocene): North Alpine -Carpathian Foreland Basin: Platt, Laa/Thaya ( Austria), Slup ( Czech Republic) ( Sieber 1960; Tejkal et al. 1967); Korneuburg Basin: Weinsteig, Stetten, Karnabrunn, Niederkreuzstetten ( Austria) ( Harzhauser 2002); Badenian (middle Miocene): North Alpine-Carpathian Foreland Basin: Gaindorf, Kalladorf, Nodendorf, Niederleis, Braunsdorf, Grund, Immendorf, Guntersdorf, Windpassing ( Austria) ( Sieber 1960; Mandic & Harzhauser 2003); Vienna Basin: Hrušovany nad Jevišovkou ( Czech Republic) (herein); Pannonian Basin: Bántapuszta, Várpalota ( Hungary) ( Steininger et al. 1973).

Figures

FIGURE 1. Paleogeography of the Paratethys during the early Miocene (Burdigalian, Eggenburgian, Sakaraulian) and the middle Miocene (Langhian, Badenian, Tarkhanian) (modified after Popov et al. 2004 and Neubauer et al. 2015).

FIGURE 2. Miocene chronostratigraphy after Gradstein et al. (2012) with regional Paratethys stages after Popov et al. (2004) and Piller et al. (2007).

FIGURE 6. Growth line traces of Turritellidae species based on own data; bold: type species of genera; black: Paratethyan species; asterisk: non-Paratethyan species (unknown basal sinuses are left blank).

FIGURE18.A–D.Ptychidia vindobonensis (Handmann,1882).A1–A2. Holotype, Enzesfeld(Austria), NHMW1853/0010/0030. B1–B2. Baden (Austria), NHMW 1862/0017/0233. C. Immendorf (Austria), holotype of Turritella (Haustator) badensis carinata Sieber, 1960, NHMW1963/0527/0000, D. Gainfarn (Austria), NHMW 1997z0178/1360. E–H. Ptychidia erynella new sp. E. Paratype, Kalladorf (Austria), NHMW 1861/0034/0032. F. Holotype, Guntersdorf (Austria), NHMW 1963/0522/0000. G. Paratype, Guntersdorf (Austria), NHMW 2018/0279/0002. H. Paratype, Guntersdorf (Austria), NHMW 2019/00277/0003. I. Ptychidia turris (de Basterot, 1825), NHMW 1872/0020/0048, Léognan (France). J–K. Ptychidia erynella new sp., Grund (Austria). J. NHMW 1963/0521/0000. K. 1963/0520/0000. L1–L2. Ptychidia sp., Ermingen (Germany), NHMW 1845/00031/0053. M–N. Ptychidia erynella new sp. M. pathologic specimen, Immendorf (Austria), NHMW 1963/0543/0000. N1–N2. Paratype, Grund (Austria), NHMW 2018/0277/0002. 18O. Ptychidia vindobonensis (Handmann, 1882), Gainfarn (Austria), NHMW 1997z0178/1361. P. Ptychidia cf. eryna (d’Orbigny, 1852), Biot (France), NHMW 1897/0032/0047.