Turritella inaequicingulata Hölzl, 1958

Harzhauser, Mathias & Landau, Bernard, 2019, Turritellidae (Gastropoda) of the Miocene Paratethys Sea with considerations about turritellid genera, Zootaxa 4681 (1), pp. 1-136 : 18-20

Turritella inaequicingulata Hölzl, 1958

Figs 6B , 8A 1 –A 2 , 8B 1 –B 2 , 8C 1 –C 2 , 9A

Turritella terebralis Lam. var. gradata Menke — Schaffer, 1912: 160, pl. 52, figs 17–19 [non Turritella gradata Menke, 1855 ].

Turritella terebralis Lam. var. percingulellata Sacco — Schaffer, 1912: 160, pl. 52, figs 20–22 [non Turritella terebralis percingulellata Sacco, 1895 ].

Turritella (Haustator) vermicularis Brocc. var. lineolatocincta Sacco — Schaffer, 1912: 162, pl. 53, fig. 1 [non Helminthia lineolatocincta ( Sacco, 1895) ].

Turritella terebralis Lam. var. gradata Menke — Rutsch, 1929: 36, pl. 2, fig. 4 [non Turritella gradata Menke, 1855 ].

* Turritella terebralis innaequicingulata [sic] nov. var. — Hölzl, 1958: 182, pl. 17, figs 8–8a.

Turritella terebralis percingulellata Sacco, 1895 — Hölzl, 1958: 183, pl. 17, fig. 9 [non Turritella terebralis percingulellata Sacco, 1895 ].

Turritella ( Turritella ?) terebralis terebralis LM. — Sieber, 1960: 248, pl. 2, fig. 1, pl. 3, fig. 8/14 [non Turritella terebralis Lamarck, 1822 s.s.]. Turritella ( Turritella ?) terebralis eggenburgensis ssp. n. — Sieber, 1960: 249.

Turritella ( Turritella ?) terebralis percingulellata Sacco — Sieber, 1960: 249 [non Turritella terebralis percingulellata Sacco, 1895 ].

Turritella ( Turritella ?) terebralis subgradata Sacco — Sieber, 1960: 250 [non Turritella terebralis subgradata Sacco, 1895 ].

Turritella terebralis Lamarck, 1822 — Báldi, 1964: 142, pl. 3, fig. 9 [non Turritella terebralis Lamarck, 1822 s.s.].

T [urritella]. terebralis inaequicingulata Hölzl, 1958 — Báldi, 1964: 142, pl. 3, fig. 9.

Turritella ( Turritella ?) terebralis Lamarck, 1822 — Steininger et al., 1971: 373, pl. 6, fig. 10 [non Turritella terebralis Lamarck, 1822 s.s.].

Turritella ( Turritella ?) terebralis eggenburgensis Sieber, 1960 — Steininger et al., 1971: 373, pl. 6, fig. 11.

Turritella ( Turritella ?) terebralis percingulata [sic] Sacco, 1895 — Steininger et al., 1971: 373, pl. 6, fig. 8 [non Turritella terebralis percingulellata Sacco, 1895 ].

Turritella ( Turritella ?) terebralis subgradata Sacco, 1895 — Steininger et al., 1971: 374, pl. 6, fig. 12 [non Turritella terebralis subgradata Sacco, 1895 ].

Turritella (Haustator) terebralis Lamarck, 1822 — Ondrejickova, 1972: 102, pl. 17, fig. 6 [non Turritella terebralis Lamarck, 1822 s.s.].

Turritella terebralis subgradata Sacco, 1893 — Fözy & Leél-Össy, 1985a: 121 [non Turritella terebralis subgradata Sacco, 1895 ].

Turritella (Turritella) terebralis terebralis— Stürmer, 1989: 97, pl. 2, fig. 1, pl. 9, fig. 3 [non Turritella terebralis Lamarck, 1822 s.s.].

Turritella (Turritella) terebralis subgradata Sacco — Stürmer, 1989: 97, pl. 2, fig. 2, pl. 9, fig. 5 [non Turritella terebralis subgradata Sacco, 1895 ].

Turritella ( Turritella ?) terebralis terebralis Lamarck — Schultz, 1998: 56, pl. 21, fig. 8 [non Turritella terebralis Lamarck, 1822 s.s.].

Turritella terebralis Lamarck 1822 — Pfister & Wegmüller, 2007: 92, pl. 8, figs 10–12, pl. 9, figs 4–6 [non Turritella terebralis Lamarck, 1822 s.s.].

Type material. Holotype Nr. 2906, Bayerisches Geologisches Landesamt, Munich, Germany; Kaltenbachgraben , Bavaria, Germany; early Miocene , Eggenburgian.

Studied material. 3 spec. NHMW 1872/0020/0195, Kaltenbachgraben ( Germany); 1 spec. NHMW 2018/0166/0001, 6 spec. NHMW 1860/0050/0268, 1spec. NHMW 1912/0001/0012 (holotype of Turritella terebralis eggenburgensis Sieber 1960 ; illustrated in Schaffer 1912, pl. 52, fig. 15), 1 spec. NHMW 1912/0001/0013 (illustrated in Schaffer 1912, pl. 52, fig. 16), Nondorf ( Austria); 6 spec. NHMW 1860/0050/0254, Mold ( Austria); 10 spec. NHMW 1934/0001/0080, 5 spec. NHMW 2018/0167/0001, (including specimens illustrated by Schaffer 1912, pl. 52, figs 17–19), Maria Dreieichen ( Austria); 1 spec. NHMW 1963/0259/0000, Gauderndorf ( Austria), illustrated in Sieber (1960, pl. 2, fig.1); 6 spec. NHMW 1845/0015, 0048, Loibersdorf ( Austria).

Illustrated material. Figs 8A 1–2 : Nondorf ( Austria): SL: 119.3 mm, MD: 29.1 mm, NHMW 1912/0001/0013; Figs 8B 1–2 : Nondorf ( Austria): SL: 118.6 mm, MD: 35.1 mm, NHMW 1912/0001/0012; Figs 8C 1 – 2 : Nondorf ( Austria): SL: 101.5 mm, MD: 30.6 mm NHMW 1860/0050/0268. Fig. 9A : Mörtersdorf ( Austria): several specimens on sediment, NHMW 2018/0165/0001 (note that specimen A 1 represents Peyrotia desmarestina ).

Revised description. Shell large, solid, moderately slender. Protoconch and earliest teleoconch whorls unknown. Teleoconch comprising at least 16 whorls; early whorls frustate, bicarinate with prominent B and C spiral cords; angulation coinciding with C cord. Flat to weakly convex sutural ramp with 4 – 5 prominent secondary spiral cords. Whorl profile weakly convex below angulation bearing 1 – 2 secondary spiral cords. Whorl profile soon becoming convex with periphery in abapical third of whorl; periphery sometimes angulated even in late teleoconch whorls. Adapical part of whorl weakly convex or rarely nearly flat. C spiral cord usually more prominent throughout growth accompanied by 7 – 9 broad spiral cords of rectangular cross-section, separated by narrower interspaces. Spiral sculpture fading out on last teleoconch whorls. Lateral sinus with steep angle, very wide, deep, simple, regularly concave, LS p distinctly abapical, without sigmoidal part in adapical half. Basal sinus straight to moderately prosocyrt. Aperture thin, subcircular; inner lip narrowly reflected passing into thin glossy, sheet-like parietal callus. No inner lirae present.

Shell measurements and ratios. Complete specimens are rare; largest nearly complete specimen: SL = 119.3 mm, MD = 29.1 mm. n = 5 adult specimens: AA: μ = 19.4° (σ = 1.67°), PA: μ = 15.8° (σ = 1.1°). Lateral sinus (n = 5): LS angle = 26.5° (σ = 3.7°), LS p = 1.4 (σ = 0.21), LS d = 3.2 (σ = 0.36).

Discussion. The Paratethyan specimens differ from the largest French shells of Turritella terebralis in their smaller size (corresponding to the height of the last whorl of the Atlantic specimens) and in the much broader and rectangular spiral cords. Even specimens of Turritella terebralis s. s. with prominent sculpture, from Léognan and Saucats ( France), display a much more delicate sculpture on early teleoconch whorls and the spiral cords are more or less trigonal in cross-section ( Figs 8D 1 –D 2 , SL: 128.2 mm, MD: 39.3 mm Léognan ( France), NHMW 1850/0018/0100). Typical specimens of T. terebralis from the Burdigalian of the Torino Hills (NHMW 1862/0035/0463) agree in sculpture with those from France and lack prominent spiral sculpture. Therefore, it seems to be justified to separate the Paratethyan populations from typical Miocene northeastern Atlantic Turritella terebralis Lamarck, 1822 . Schaffer (1912) compared the Paratethyan shells with T. t. percingulellata Sacco, 1895, from the Burdigalian of the Torino Hills. The spiral cords of this Italian morphotype, however, are narrower and Sacco (1895) compared its sculpture with that of Turritella venus d’Orbigny, 1852 . This sculpture type is also seen in Turritella terebralis sulcata Dollfus, Cotter & Gomes, 1903 and Turritella terebralis lusitanica Zbyszewski, 1957 from the early Miocene of Portugal. For the Paratethyan shells, Hölzl (1958) introduced Turritella terebralis innaequicingulata as variation name based on specimens from the Eggenburgian of Bavaria [note that innaequicingulata is a typographical error for inaequicingulata , which was correctly used by Hölzl (1958) several times in the discussion and the plate caption. Therefore, we emend the incorrect original name]. Later, Sieber (1960) introduced Turritella terebralis eggenburgensis as a subjective junior synonym of T. t. inaequicingulata for specimens from the Eggenburg region in Austria. The prominent sculpture of the early spire is somewhat variable concerning width of spiral cords and their interspaces but a separation of these always co-occurring morphotypes as biological subspecies is surely unjustified.

Turritella inaequicingulata is a very characteristic species, which is restricted in the Paratethys to the early and middle Eggenburgian (early Burdigalian). It disappeared already during the late Eggenburgian and is unknown from Ottnangian strata. During the Karpatian (late Burdigalian) it was already fully replaced by Turritella gradata Menke in Hörnes, 1855 .

Paleoenvironment. Turritella inaequicingulata is found in large numbers, forming “turritelline-dominated assemblages” ( Allmon & Knight 1993) in the Loibersdorf Formation in the Eggenburg area ( Fig. 9A ); the depositional environment represents shallow sublittoral settings ( Mandic et al. 2004b).

Distribution in Western and Central Paratethys. Eggenburgian (early Miocene): North Alpine Foreland Basin: numerous localities in the St. Gallen/Belpberg region ( Pfister & Wegmüller 2007); Kaltenbachgraben (Bavaria, Germany) ( Hölzl 1958); Burgschleinitz, Eggenburg, Gauderndorf, Maria Dreieichen, Mold, Loibersdorf, Nondorf ( Steininger et al. 1971); Vaďovce Basin: Vaďovce ( Slovak Republic) ( Steininger et al. 1971); Horná Nitra Basin: Vélka Čausa ( Slovak Republic) ( Steininger et al. 1971); Nógrád–Filakovo Basin: Lipovany, Kalonda ( Slovak Republic) ( Ondrejickova 1972); Pannonian Basin: Budafok-Budapest ( Hungary) ( Báldi 1964; Steininger et al. 1971); Eastern Mátra Mountains: Illona valley ( Hungary) ( Fözy & Leél-Össy 1985a, b).

Distribution of Turritella terebralis Lamarck, 1822 s. l.

Numerous subspecies names have been proposed for various morphotypes of Turritella terebralis ; e.g. grata, aquensis, aturensis and fragilis for French specimens by Cossmann & Peyrot (1922), subgradata, turritissima, subagibbosa and percingulellata for Italian specimens by Sacco, 1895, sulcata and lusitanica for Portuguese specimens by Dollfus et al. (1903) and Zbyszewski (1957). We have not seen the types of these ‘species’ and cannot evaluate if Turritella terebralis s. l. consist of distinct (sub)species with paleogeographic and/or stratigraphic significance (as proposed herein at least for the Paratethyan populations). Turritella terebralis s. s. is most probably restricted to the Aquitanian and Burdigalian of the northeastern Atlantic. Turritella terebralis s. l., however, has a much wider geographic distribution:

Proto-Mediterranean Sea and northeastern Atlantic. Aquitanian and Burdigalian (early Miocene): Aquitaine Basin: Léognan, Saucats, Mérignac, Saint-Paul-lès-Dax ( France) ( Cossmann & Peyrot 1922; Lozouet et al. 2001); Lisbon region: e.g. Val-de-Chellas, Cacilhas, Pragal, Carnide, Forno do Tijolo) ( Dollfus et al. 1903; Zbyszewski 1954, 1957); Burdigalian (early Miocene): Torino Hills: Baldissero, San Raffaele ( Italy) Sacco 1895); Mesohellenic Basin: Odria, Argos Orestikon ( Greece) (herein); Antalya Basin: Kaþ region ( Turkey) ( İslamoğlu 2004); Mut Basin: Burunköy ( Turkey) ( Mandic et al. 2004a); Gebel Gharra (SW Suez, Egypt) (herein); Qom Basin: Qom ( Iran) (herein); Langhian: Aquitaine Basin: Saubrigues ( France) ( Cossmann & Peyrot 1922). Turritella terebralis s.l. is a typical early Miocene species; specimens described from middle Miocene deposits might represent reworked shells (e.g. Peyrot 1938).

non in tropical eastern Atlantic. Böhm (1913) recorded Turritella terebralis sulcata Dollfus, Cotter & Gomes, 1903 and T. t. applanata Böhm, 1913 from the early? Miocene of Angola. Although the lateral sinus of the West African species agrees with that of Turritella terebralis s. l., it differs in its deeply incised suture and high last whorl and is overall rather slender compared to T. terebralis . Later, Dartevelle et al. (1954) and Brébion (1983) reported Turritella terebralis again from the Burdigalian of Angola. The illustrated specimen of Dartevelle et al. (1954) does not allow a clear identification, but it most probably also represents T. applanata . Turritella applanata Böhm seems to be more closely related to the extant West African T. ungulina ( Linnaeus, 1758) and T. ligar Deshayes, 1843 than to the Miocene T. terebralis .

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).

FIGURE 8. A–C. Turritella inaequicingulata Hölzl, 1958, Nondorf (Austria); A1–A2. NHMW 1912/0001/0013. B1–B2. NHMW 1912/0001/0012. C1–C2. NHMW 1860/0050/0268. D1–D2. Turritella terebralis Lamarck, 1822, Léognan (France), NHMW 1850/0018/0100). E–F. Turritella gradata Menke in Hörnes, 1855. E. lectotype, Weinsteig (Austria), NHMW 1846/0037/0397. F1–2. Stetten-Teiritzberg (Austria), NHMW 2018/0169/0001.

FIGURE 9. A. Turritella inaequicingulata Hölzl 1958, NHMW 2018/0165/0001, Mörtersdorf (Austria) with a specimen of Peyrotia desmarestina (de Basterot, 1825), indicated by an arrow. B. Turritella gradata Menke in Hörnes, 1855, Weinsteig (Austria), NHMW 2018/0165/0002.