Allmonia carniolica ( Stache, 1858 ) 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 : 91-94

Allmonia carniolica ( Stache, 1858) new comb.

Figs 6 Ad, 26A, 26B 1 –B 2, 26C 1 –C 2, 26D, 26E, 26F 1 –F 2 ,26G, 26H

* Turritella carniolica— Stache, 1858: 378.

Turritella (Protoma) Carniolica Stache (in litt.) — Hilber, 1893: 1025, pl. 1, figs 2–4.

Turritella (Protoma) Bartelmaica Hilber (nov.) — Hilber, 1893: 1028, pl. 1, figs 5–6.

Zwischenform zwischen Turritella Carniolica Stache und Bartelmaica Hilb. — Hilber, 1893: 1029, pl. 1, fig. 7.

? Protoma costai D.C.G. n. sp. — Dollfus et al., 1903: 1 [plates published in Pereira da Costa, 1867: pl. 28, figs 16a–16b].

? Protoma proto Basterot — Dollfus et al., 1903: 1 a [plates published in Pereira da Costa, 1867: pl. 28, figs 18a–18b] [non Turritella proto de Basterot, 1825 = Allmonia cathedralis ( Brongniart, 1823) ].

Protoma rotifera Lamarck — Dollfus et al., 1903: 2, pl. 29, figs 1a–1b [non Turritella rotifera Lamarck, 1822 ].

Protoma proto Bast. — Strausz, 1954: 40, pl. 1, figs 17a–17c [non Turritella proto de Basterot, 1825 = Allmonia cathedralis ( Brongniart, 1823) ].

? Proto rotifera Lamarck — Zbyszewski, 1957: 158, pl. 11, figs 111–119 [non Allmonia rotifera ( Lamarck, 1822) ].

? [ Proto rotifera Lamarck] var. costai D. C. G. — Zbyszewski, 1957: 158, pl. 11, fig. 116 [non Allmonia rotifera ( Lamarck, 1822) ].

Protoma (Protoma) carniolica Stache — Sieber, 1960: 278, pl. 2, fig. 5.

Protoma P [rotoma]. bartelmaica Hilber — Sieber, 1960: 280, pl. 3, fig. 8/22.

Protoma proto Basterot, 1825 — Strausz, 1966: 111, pl. 4, figs 4–6 [non Turritella proto de Basterot, 1825 = Allmonia cathedralis ( Brongniart, 1823) ].

Turritella (Protoma) carniolica Stache — Kochansky-Devidé, 1970: 11, fig. 4.

Protoma rotifera Lamk. — Kókay, 1972: 48, pl. 2, fig. 2 [non Allmonia rotifera ( Lamarck, 1822) ].

Protoma rotifera Lamarck, 1822 — Steininger et al., 1973: 403, pl. 4, fig. 1 [non Allmonia rotifera ( Lamarck, 1822) ].

Protoma proto Basterot, 1825 — Steininger et al., 1973: 403, pl. 4, fig. 3 [non Turritella proto de Basterot, 1825 = Allmonia cathedralis ( Brongniart, 1823) ].

? Proto rotifera Lamarck — Fonseca, 1977: plate captions, pl. 4, fig. 17.

Protoma rotifera Lamarck — Steininger, et al. 1978: 403, pl. 4, fig. 1 [non Allmonia rotifera ( Lamarck, 1822) ].

Protoma (Protoma) carniolica carniolica (Stache; Hilber) — Mikuž, 1983: 117, fig. 7.

Protoma (Protoma) carniolica n. ssp. — Mikuž, 1983: 117, fig. 8.

Protoma (Protoma) cathedralis (Brongniart) — Mikuž, 1983: 117, fig. 9 [non Allmonia cathedralis ( Brongniart, 1823) ].

Protoma (Protoma) carniolica carniolica (Stache) — Mikuž, 2003: 308, pl. 10, fig. 23.

Protoma (Protoma) bartelmaica ( Stache, 1858) (in Hilber, 1893) — Mikuž, 2009: 14, pl. 2, fig. 28.

Protoma (Protoma) carniolica carniolica ( Stache, 1858) (in Hilber, 1893) — Mikuž, 2009: 14, pl. 2, fig. 29.

Protoma (Protoma) carniolica stachei (n. ssp.?) — Mikuž, 2009: 14, pl. 2, fig. 30.

Protoma (Protoma) cf. rotifera ( Lamarck, 1822) — Mikuž, 2009: 155, pl. 15, fig. 31 [non Allmonia rotifera ( Lamarck, 1822) ].

Protoma (Protoma) cathedralis ( Brongniart, 1823) — Mikuž, 2009: 15, pl. 2, fig. 32 [non Allmonia cathedralis ( Brongniart, 1823) ].

Protoma (Protoma) cathedralis paucicincta Sacco, 1895 — Mikuž, 2009: 15, pl. 2, fig. 33 [non Allmonia paucicincta ( Sacco, 1895) ].

Protoma carniolica (Stache) — Kókay, 2013: 148.

This species has been discussed in several unpublished PhD theses, which are occasionally cited:

Turritella (Protoma) carniolica Stache — Ramovš, 1974: 56, fig. 145.

Protoma (Protoma) carniolica Stache — Perše, 1980: 14, pl. 5, figs 2–3.

Protoma (Protoma) bartelmaica (Hilber) — Perše, 1980: 15, pl. 5, fig. 4.

Protoma (Protoma) bartelmaica ( Hilber, 1893) — Mikuž, 1982: 123, pl. 2, figs 1a–1b.

Protoma (Protoma) carniolica carniolica ( Stache, 1858) — Mikuž, 1982: 128, pl. 1, figs 3a–3b, pl. 12, figs 1–8.

Protoma (Protoma) cf. rotifera ( Lamarck, 1822) — Mikuž, 1982: 155, pl. 16, fig. 6, pl. 17, fig. 4 [non Allmonia rotifera ( Lamarck, 1822) ].

Protoma (Protoma) carniolica stachei n. ssp. — Mikuž, 1982: 132, pl. 13, figs 1–5.

Type material. The syntypes of Stache (1858) are stored in the collections of the Geological Survey in Vienna (two lots, 47 spec. GBA 2008/027/0003, 10 spec., GBA 2008/027/0005, 1 spec. GBA 1858/007/0001). From the syntypes, we select the specimen illustrated as Fig. 26A as lectotype ( GBA 1858/007/0001); middle Miocene, Badenian, Sveta Marjeta (St. Margarethen), Slovenia.

Studied material. Specimens from Slovenia: syntypes of Turritella (Protoma) Bartelmaica Hilber, 1893 : UMJG&P 1255, Šentjernej (Sankt Bartlmä), UMJG&P 1256, Dolenja Stara vas (= Altendorf), UMJG&P 1257, Gorenje Vrhpolje (= Oberfeld), UMJG&P 1258, Šmarje (St. Marein) (all stored in Universal museum Joanneum Graz). Specimens described by Hilber (1893) as Turritella (Protoma) Carniolica Stache : UMJG&P 1248, Dolenja Stara vas (= Altendorf), UMJG&P 1249 Šmarje (St. Marein) illustrated as fig. 2 in Hilber (1893), UMJG&P 1250 Šentjernej (Sankt Bartlmä), UMJG&P 1251 Šentjernej (Sankt Bartlmä), illustrated as fig. 4 in Hilber (1893), UMJG&P 1252 Gorenje Vrhpolje (= Oberfeld), UMJG&P 1253 Šentjernej (Sankt Bartlmä), illustrated as fig. 3 in Hilber (1893), UMJG&P 1254 Šmarje (St. Marein), UMJG&P 1260–1261 Gorenje Vrhpolje (= Oberfeld). 1 spec. NHMW 1963/0546/0000, ‘Krain’, (illustrated in Sieber 1960: pl. 2, fig. 5); 8 spec., NHMW 1880/0017/0986, Gorjanci; 14 spec., NHMW 2018/0154/0001, 1 spec., NHMW 2018/0154/0002, 1 spec., NHMW 2018/0154/0003, 1 spec., NHMW 2018/0154/0004, Sveta Marjeta; 7 spec., NHMW 1884/0019/2178, 1 spec., NHMW 2018/0155/0001, 1 spec., NHMW 2018/0155/0002, 1 spec., NHMW 2018/0155/0003, 15 spec. NHMW 2018/0155/0004, Ivanji dol; 1 spec., NHMW 2018/0154/0005, 1 spec. NHMW 2018/0154/0006, Sveta Marjeta. 1 spec. NHMW 1863/0057/0069, Sacavém (Lisbon, Portugal).

Illustrated material. Fig. 26A : Sveta Marjeta ( Slovenia): SL: 86.7 mm, MD: 19.5 mm, GBA 2008/ 027/0003; Figs 26B 1 –B 2 : Ivanji dol ( Slovenia): SL: 57.4 mm, MD: 14.5 mm, NHMW 2018 / 0155/0001; Figs 26C 1 –C 2 : Sveta Marjeta ( Slovenia): SL: 53.8 mm, MD: 13.9 mm, NHMW 2018 / 0154/0003; Fig. 26D : Sveta Marjeta ( Slovenia): SL: 63.1 mm, MD: 18.2 mm, NHMW 2018 / 0154/0005; Fig. 26F 1 –F 2 : Šentjernej ( Slovenia): SL: 46.0 mm, MD: 23.5 mm, UMJG&P 1251; Fig. 26G : Ivanji dol ( Slovenia): SL: 31.4 mm, MD: 21.4 mm, NHMW 2018 / 0155/0003; Fig. 26H : Ivanji dol ( Slovenia): SL: 35.2 mm, MD: 14.5 mm, NHMW 2018 / 0155/0002; Fig. 26E : Sacavém (Lisbon, Portugal): SL: 89.9 mm, MD: 23.5 mm, NHMW 1863 /0057/0069.

Revised description. Shell large, slender. Protoconch and first teleoconch whorls unknown. Earliest preserved teleoconch whorls bicarinate, frustate with B spiral cord in mid-whorl; flat, broad sutural ramp, slight concavity below C spiral cord. Entire whorl covered by faint spiral threads. Sutural ramp gradually passing into broad subsutural collar; B and C cords developing into prominent, rounded ribs on early spire; afterwards B cord usually disappears completely, whereas C cord continues as delicate spiral cord or faint angulation. Whorl profile changing quickly from bicarinate to straight sided and strongly telescoped with prominent concavity on late teleoconch whorls. Suture insignificant. Prominent, densely-spaced growth lines especially on adult whorls. Lateral sinus with steep angle; wide, deep, simple concave, rarely with faint inflection point close to adapical suture; adapical LS p. Broad concavity below basal angulation of last whorl, delimiting very broad, scalloped fasciole, indicating broad, incised basal sinus. Aperture largely destroyed in all available specimens.

Shell measurements and ratios. No complete specimens are available; largest specimen without aperture: SL = 89.1 mm, MD = 20.5 mm. SL/MD ratio of 8 largest fragmentary specimens = 4.5, suggesting maximum SL of ~ 104 mm based on specimen Fig. 1M (representing last whorl with fasciole); AA (n=11): μ = 16.3° (σ = 1.57°), PA: μ = 11.2° (σ = 1.08°). Lateral sinus (n = 7): LS angle = 21.6° (σ = 3.9°), LS p = 1.1 (σ = 0.17), LS d = 4.3 (σ = 0.36).

Paleoecology. See discussion on genus.

Discussion. Stache (1858) gave a very short diagnosis of the species, comparing it with Protoma cathedralis . He therefore made the name available and is the author of Turritella carniolica , although Hilber (1893) gave a much more comprehensive description. The species is abundant in the Krško Basin in Slovenia and Stache (1858: 378) mentioned that the shells can be collected ‘ by the bushel ’. This rich material reflects a considerable variability concerning spiral sculpture. Especially the strength of the B and C spiral cords and their point of disappearance are variable. In some specimens, a spiral cord may appear along the abapical suture, fading out again on the last whorls. Similarly, the secondary spiral cords may vary in strength. The adapical spiral may appear bifurcate due to the occasional presence of a narrow spiral groove. All these features vary even within single specimens during ontogeny. Hilber (1893) separated Turritella (Protoma) bartelmaica from T. (P.) carniolica based on the less slender profile, the occasional presence of a sixth spiral thread at the abapical suture and the weaker keel. Later, Mikuž (1982, 2009) separated an additional subspecies as P. carniolica stachei [this name became available by the publication of Mikuž (2009) because Mikuž (1982) is an unpublished PhD thesis]. Other morphotypes have been variously identified as P. rotifera , P. cathedralis , and P. cathedralis paucicincta ( Mikuž 2009) . All these taxa are found co-occurring at the same localities within a limited geographic area and even Hilber (1893) recognized intermediate specimens. Therefore, we consider Allmonia carniolica to be a morphologically variable species and reject the taxonomic separation of various morphotypes.

Allmonia carniolica has been confused with A. cathedralis ( Brongniart, 1823) , A. paucicincta ( Sacco, 1895) and A. rotifera ( Lamarck, 1822) . A separation from A. cathedralis and A. paucicincta is obvious based on the smaller size, the slender shell, and deeper lateral sinus of A. carniolica . Moreover, the deep concavity of late teleoconch whorls does not occur in A. cathedralis and A. paucicincta . This prominent concavity of the whorl profile, however, agrees well with A. rotifera . A separation of A. rotifera is based on the early teleoconch whorls, which pass from convex to straight-sided at a growth stage in which A. carniolica has bicarinate, frustate whorls. Moreover, A. rotifera develops a sharp keel close to the adapical suture, forming from a well-defined A-spiral cord, which results in a screw-like profile. Such a sharp keel is never developed by A. carniolica .

Although Allmonia carniolica is described so far only from a limited area in the Paratethys, it seems to have had a much wider distribution. Several specimens from the Burdigalian to Tortonian of Portugal, described by Pereira da Costa (1867), Dollfus et al. (1903), and Zbyszewski (1957) are highly reminiscent of Allmonia carniolica , but differ slightly in their broader shells. A specimen from the Tortonian of Sacavém in Lisbon, in the NHMW collection, however, agrees fully with Paratethyan specimens also in its slender profile ( Fig. 26E ); similarly, the Burdigalian specimen illustrated by Fonseca (1977) from the Tagus Basin can hardly be separated from the Paratethyan specimens. A biogeographic link of the Allmonia-carniolica -bearing Paratethyan assemblages with the Portuguese faunas is also indicated by the co-occurrence of the otherwise rare stromboid Pereiraea gervaisi Vezian, 1856 ( Bartol et al. 2014).

Distribution. Central Paratethys. Karpatian (early Miocene): Várpalota and Bántapuszta ( Hungary) ( Kókay 1972; Steininger et al. 1973) [note that Kókay (1972) and Steininger et al. (1973) considered these occurrences as Ottnangian, but Mandic (2003) documented a younger (Karpatian) age for the strata]. Badenian (middle Miocene): Krško Basin: Ivanji dol, Dolenja Brezovica, Šentjernej (= Sankt Bartlmä), Gorenje Vrhpolje (= Oberfeld), Dolenja Stara vas (= Altendorf), Šmarje (St. Marein), Škocjan, Selo, Sveta Marjeta ( Slovenia) ( Stache 1858; Hilber 1893; Mikuž 2009). Styrian Basin: Klockerberg at Wetzelsdorf ( Sieber 1960). Pannonian Basin: Pécsszabolcs, Várpalota ( Bohn-Havas 1973; Kókay 2013). Eastern Atlantic. Along the Portuguese Atlantic coast, the species seems to be present from the Burdigalian to the Tortonian (e.g. Tagus Basin, Sacavém in Lisbon) ( Fonseca 1977; herein).

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 26. A–H. Allmonia carniolica (Stache, 1858). A. Sveta Marjeta (Slovenia), GBA 2008/027/0003. B1–B2. Ivanji dol (Slovenia), NHMW 2018/0155/0001. C1–C2. Sveta Marjeta (Slovenia), NHMW 2018/0154/0003. D. Sveta Marjeta (Slovenia), NHMW 2018/0154/0005. E. Sacavém (Lisbon, Portugal), NHMW 1863/0057/0069. F1–F2. Šentjernej (Slovenia), UMJG&P 1251, G. Ivanji dol (Slovenia), NHMW 2018/0155/0003. H. Ivanji dol (Slovenia), NHMW 2018/0155/000. I1–I2. Allmonia rotifera (Lamarck, 1822), Catalonia, NHMW 9836.