Allmonia paucicincta ( Sacco, 1895 ) 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 : 87-90

Allmonia paucicincta ( Sacco, 1895) new comb.

Figs 6 Ab, 25A, 25B 1 –B 3, 25C, 25D

Turritella (Proto) cathedralis Brong. — Hörnes, 1855: 419 (partim), pl. 43, fig. 1 [non Allmonia cathedralis ( Brongniart, 1823) ].? [ Proto ] nanus May. — Mayer, 1872: 25 [nomen nudum]. Turritella (Hemitropis) cathedralis Brong. — Handmann, 1882: 212 [non Allmonia cathedralis ( Brongniart, 1823) ].? Turritella (Protoma) Telleri Hilber, 1893: 1025 , pl. 1, fig. 1 [taxon inquirendum]. * P [rotoma]. cathedralis var. paucicincta Sacc. — Sacco, 1895: 32 [new name for Turritella cathedralis in Hörnes, 1855 , pl. 43, fig. 1, non Brongniart, 1823]. [ Protoma cathedralis var.] paucicincta— Sacco, 1896: 80. Protoma cathedralis Brong. var. paucicincta Sacco — Schaffer, 1912: 164, pl. 53, figs 17–21. Protoma cathedralis Brong. var. quadricincta Schaffer, 1912: 165 , pl. 53, figs 15–16. Turritella (Haustator) vermicularis Brocc. var. perlatecincta Sacco — Schaffer, 1912: 162, pl. 52, figs 26–28 [non Turritella (Haustator) vermicularis perlatecincta Sacco, 1895 ]. Turritella cathedralis Brongn. — Toula, 1918: 449, pl. 27, fig. 20 [non Allmonia cathedralis ( Brongniart, 1823) ] Protoma cathedralis (Brong.) var. paucicincta Sacco — Rutsch, 1929: 42, pl. 2, fig. 15. Protoma cathedralis (Brongniart) var. paucicincta Sacco — Mongin, 1952: 49, pl. 1, fig. 14. Protoma proto (Bastérot) var. fabreae new var. — Mongin, 1952: 50, pl. 1, fig. 15. Protoma (Protoma) cathedralis paucicincta Sacco — Csepreghy-Meznerics, 1954: 17, pl. 1, figs 15, 22. P [rotoma]. (P [rotoma].) cathedralis paucicincta Sacco — Sieber, 1958a: 138. Protoma proto (Bast.) — Seneš, 1958: 134, pl. 21, fig. 270 [non Turritella proto de Basterot, 1825 = Allmonia cathedralis ( Brongniart, 1823) ]. Protoma cathedralis var. quadricincta Schff. — Seneš, 1958: 134, pl. 21, figs 276–277. Protoma cathedralis var. bellunensis Venzo — Seneš, 1958: 134, pl. 21, fig. 281 [non Allmonia cathedralis bellunensis ( Venzo, 1937) ]. Protoma cathedralis var. (?) pseudolaevis Sacc. — Seneš, 1958: 135, pl. 21, fig. 273 [non Allmonia cathedralis var. pseudolaevis ( Sacco, 1895) ].

Protoma cathedralis var. (?) concavosimplex Sacc. — Seneš, 1958: 135, pl. 21, figs 274–275 [non Allmonia cathedralis var. concavosimplex ( Sacco, 1895) ].

Protoma cathedralis var. (?) paucicincta Sacc. — Seneš, 1958: 135, pl. 21, figs 271–272.

Protoma cathedralis quadricincta Schaffer, 1912 — Hölzl, 1958: 186, pl. 18, fig. 2.

Protoma cathedralis paucicincta Sacco, 1895 — Báldi, 1964: 142, pl. 3, fig. 10.

Protoma (Protoma) cathedralis paucicincta Sacco — Sieber, 1960: 263, pl. 3, fig. 7.

Protoma (Protoma) cathedralis quadricincta Schaffer — Sieber, 1960: 264, pl. 3, fig. 5.

? Protoma (Protoma) cathedralis quadricincta Schaffer, 1912 — Steininger, 1963: 48, pl. 10, figs 4–5.

Turritella (Protoma) cathedralis var. alterspirata Schaff. — Csepreghy-Meznerics, 1967: 93.

? Protoma cathedralis Brongn. ex quadricincta Schaffer — Вáldi & Radóсz, 1971:145, pl. 2, fig. 4.

Protoma (Protoma) cathedralis paucicincta Sacco — Steininger et al., 1971: 374, pl. 7, fig. 1.

Protoma (Protoma) cathedralis paucicincta Sacco — Steininger et al., 1971: 375, pl. 7, figs 2–3.

Protoma cathedralis ( Brongniart, 1823) — Ondrejickova, 1972: 106, pl. 17, figs 4–5 [non Allmonia cathedralis ( Brongniart, 1823) ].

Protoma proto (Basterot), 1825 — Bohn-Havas, 1973: 1042, pl. 3, figs 15–16 [non Turritella proto de Basterot, 1825 = Allmonia cathedralis ( Brongniart, 1823) ].

Protoma cathedralis paucicincta Sacco, 1895 — Bohn-Havas, 1973: 1043, pl. 4, fig. 1.

Protoma cathedralis ( Brongniart, 1823) s.l. — Báldi, 1973: 253, pl. 28, fig. 12 [non Allmonia cathedralis ( Brongniart, 1823) ].

Protoma cathedralis Brongniart, 1825 — Báldi & Steininger, 1975: 344, pl. 8, fig. 10 [non Allmonia cathedralis ( Brongniart, 1823) ].

[ Protoma cathedralis ( Brongniart, 1823) ] var. paucicincta Sacco, 1895 — Ferrero Mortara et al., 1984: 237.

Protoma cathedralis ( Brongniart, 1823) — Fözy & Leél-Össy, 1985a: 121, pl. 3, fig. 3 [non Allmonia cathedralis ( Brongniart, 1823) ].

Protoma cathedralis ( Brongniart, 1823) — Fözy & Leél-Össy, 1985b: 184, pl. 3, fig. 1–2 [non Allmonia cathedralis ( Brongniart, 1823) ].

Protoma (Protoma) cathedralis paucicincta Sacco — Schultz, 1998: 56, pl. 21, fig. 17.

Protoma cathedralis paucicincta Sacco, 1895 — Harzhauser, 2002: 76.

Protoma (Protoma) aff. telleri ( Hilber, 1893) — Mikuž, 2009: 15, pl. 2, fig. 34.

Protoma (Protoma) obelisca ( Grateloup, 1832) — Mikuž, 2009: 15, pl. 2, figs 35–37 [non Turritella cathedralis var. obelisca Grateloup, 1846 = Allmonia cathedralis ( Brongniart, 1823) ].

non Protoma cathedralis funiculata Bors. — Steininger, 1973: 402, pl. 3, fig. 14 [probably not an Allmonia ].

non Protoma (Protoma) cathedralis paucicincta Sacco, 1895 — Mikuž, 2009: 15, pl. 2, fig. 33 [ Allmonia carniolica ( Stache, 1858) ].

Type material. Holotype NHMW 1846/0010/0014, illustrated in Hörnes 1855, pl. 43, fig. 1); Eggenburgian (early Miocene) Burgschleinitz ( Austria).

Studied material. 1 spec. NHMW 1846/0010/0014, Burgschleinitz (illustrated in Hörnes 1855, pl. 43, fig. 1, holotype of Turritella (Protoma) cathedralis paucicincta Sacco, 1895 ), 1 spec coll. Anton Breitenberger, 4 spec. NHMW 2018/0128/0001, NHMW 2018/0128/0002, NHMW 2018/0128/0003, NHMW 2018/0128/0004, Burgschleinitz ( Austria), 17 spec. NHMW 1849/0013/0015, NHMW 1860/0050/0323, NHMW 1864/0001/0481, NHMW 1866/0020/00012, Eggenburg ( Austria), 7 spec. NHMW 1851/0006/0020, NHMW 1850/0009/0050, NHMW 1860/0050/3720, NHMW 1866/0040/0038, 1 spec. NHMW 2018/0129/0001, Loibersdorf (illustrat- ed in Schaffer 1912, pl. 53, fig. 19), 2 spec. NHMW 1850/0009/0037, Mörtersdorf ( Austria), 2 spec. NHMW 1989/0031/0001, NHMW 1989/0031/002, Maria Dreieichen ( Austria), 2 spec. KM, Maria Dreieichen (illustrated in Schaffer 1912, pl. 53, figs 15, 16), 3 spec. KM, Maigen (illustrated in Schaffer 1912, pl. 53, figs 17, 18, 21), 4 spec. NHMW 1890/0025/0015, Engelsdorf ( Austria), 1 spec. NHMW 1860/0050/0253 Molt ( Austria), 1 spec. NHMW 2018/0130/0001, Neuruppersdof at Laa ( Austria), 2 spec. NHMW 2018/0131/0001, NHMW 1855/0045/0178, Steinebrunn ( Austria), 4 spec. NHMW 1863/0015/0528, NHMW 1869/0001/0545, Forchtenau ( Austria), 1 spec. NHMW 1863/0015/0000 Rudice ( Czech Republic), 5 spec. NHMW 1861/0040/0117, NHMW1864/0001/5420, Hrušovany ( Czech Republic). 1 spec. UMJG&P 1247 (Inv. No. 200323), Ivanji dol ( Slovenia), illustrated in Hilber (1893, pl. 1, fig. 1), holotype of Turritella (Protoma) telleri Hilber, 1893 .

Illustrated material. Fig. 25A : Holotype, Burgschleinitz ( Austria): SL: 133.8 mm, MD: 33.1 mm, NHMW 1846/0010/0014; Figs 25B 1 –B 3 : Burgschleinitz ( Austria): SL: 123.8 mm, MD: 33.5 mm, coll. Anton Breitenberger; Fig. 25C : Burgschleinitz ( Austria): SL: 110 mm, MD: 28.5 mm, NHMW 2018/0128/0002; Fig. 25D : Burgschleinitz ( Austria): SL: 116.8 mm, MD: 28.2 mm, 2018/0128/0001.

Revised description. Shell large, robust, moderately slender, of at least 15–18 teleoconch whorls. Protoconch unknown. Whorl profile of first 5–6 teleoconch whorls bicarinate convex, grading into straight sided profile by 12 th to 14 th teleoconch whorl, followed by concave, telescoped whorls. First teleoconch whorls bicarinate convex with B and C spiral cords of approximately equal strength; B cord slightly below mid-whorl. Suture moderately incised. Primary spirals initially sharp becoming convex around 5 th teleoconch whorl; occasionally, secondary u spiral cord present. Bifid spiral cord A appears around 8 th to 9 th teleoconch whorl, being equal in strength to B and C cords until 9 th to 11 th teleoconch whorl. Separation into A 1 and A 2 indicated by weak incision or deep furrow. Later, A 1 and A 2 merge into a prominent, raised subsutural collar. B and C fade out within 13 th to 15 th teleoconch whorls, coinciding with transition from flat sided whorl profiles into moderately concave profiles. Abapical spiral cord d may occur close to abapical suture or may be nearly hidden by spiral cord A of subsequent whorl. Faint spiral threads cover entire shell forming delicate, cancellate pattern at cross sections with growth lines. Lateral sinus with steep angle; wide, moderately deep, simple concave with steeply prosocline adapical part and moderately opisthocyrt abapical part, passing via prominent inflection into basal sinus; adapical LS p. Last whorl with marked basal angulation coinciding with spiral cord d. Base low, markedly concave between angulation and fasciole. Very broad fasciole often with scalloping growth increments. Inner lip forming thick glossy sheet in parietal area, bordering fasciole. Outer lip thin, nearly straight sided passing via flaring convexity into moderately convex basal lip. Basal sinus deeply incised, occasionally forming short, slightly deflected canal.

Shell measurements and ratios. Complete specimens are extremely rare; largest nearly complete specimen: SL = 133.8 mm, MD = 33.1 mm. n = 8 adult specimens: mean MD 28.4 mm (σ = 3.4 mm), AA: μ = 20.0° (σ = 2.33°), PA: μ = 14.8° (σ = 0.90°). Lateral sinus (n = 5): LS angle = 23.4° (σ = 4.3°), LS p = 1.1 (σ = 0.16), LS d = 5.8 (σ = 0.82).

Discussion. Hörnes (1855: 419) already emphasized that the Austrian shells differed from those from France, without separating them taxonomically. A comparison with shells from the early Miocene of Léognan, Bordeaux and Saucats ( France, Aquitaine Basin) in the NHMW collections supports this view. French specimens differ in having slightly higher early teleoconch whorls with a more abapical position of the B spiral cord, resulting in a higher and steeper sutural ramp. Later teleoconch whorls lack the prominent subsutural collar, resulting in a distinctly less telescoped profile. The secondary spiral cords s and t of A. cathedralis may attain nearly the same strength as the primary spiral cords even within the last spire whorls and the reduction of the spiral cords on the last whorls is much weaker. Moreover, the basal sinus of A. paucicincta is deeper. The differences in sculpture and profile were also discussed by Sacco (1895), who introduced paucicincta as variation name for the Austrian shell illustrated by Hörnes (1855, pl. 43, fig. 1). Similarly, Cossmann & Peyrot (1922) stressed these differences and supported the taxonomic separation from ‘ Protoma’ cathedralis . Herein, we treat the Paratethyan specimens as distinct species due to their geographic separation from the Italian and French populations and due to their rather constant morphology. Allmonia paucicincta differs from the coeval Proto-Mediterranean A. funiculata in its strong reduction of the spiral sculpture in late stages of growth and the markedly telescoped and adapically carinate profile. Specimens of Allmonia funiculata in the NHMW collections, differ also in their smaller size, more slender shells and a weakly scalloped fasciole.

Hilber (1993) considered the Austrian specimens described by Hörnes (1855) also to be different from the French A. cathedralis and introduced Turritella (Protoma) telleri as new name, which thus would have priority over Protoma paucicincta Sacco 1895 . Unfortunately, the holotype is a fragment from the Badenian of Slovenia showing four prominent spiral cords, of which the adapical two are somewhat amalgamated ( Fig. 25E : SL: 44.2 mm, MD: 21.0 mm, Ivanji dol ( Slovenia) UMJG&P 1247). This spiral sculpture differs from that of typical A. paucicincta . Moreover, it lacks the typical subsutural collar. It might represent an aberrant and reworked A. paucicincta but due to the fragmentary nature of the single available specimen, the status of Allmonia telleri remains unclear and we treat it as taxon inquirendum.

Schaffer (1912) introduced Protoma cathedralis quadricincta for spire fragments with four spiral cords, resulting from a strong separation of the A 1 and A 2 spiral cords. The specimens were collected along with typical specimens of Protoma cathedralis paucicincta from the same localities and layers. Even Schaffer (1912) observed transitional morphs. Consequently, we treat this taxon as subjective junior synonym of Allmonia paucicincta .

Paleoenvironment. See discussion of genus.

Distribution. Western and Central Paratethys.Egerian (Chattian/Aquitanian): Pilis-W. Cserhát Region: Kováčov ( Slovakia) ( Seneš 1958); Transdanubian Mountains: Máriahalom ( Hungary) ( Báldi & Steininger 1975); Pómaz, Diósjenö ( Hungary) ( Báldi 1973; Báldi & Steininger 1975). Eggenburgian (early Miocene): North Alpine Foreland Basin: Bern, St. Gallen ( Switzerland) ( Pfister & Wegmüller 2007); Kaltenbachgraben, Thalberggraben (Bavaria, Germany) ( Hölzl 1958); Eggenburg, Kühnring, Maigen, Gauderndorf, Molt, Mörtersdorf, Loibersdorf, Nondorf, Maria Dreieichen, Engelsdorf, Burgschleinitz ( Austria) ( Sieber 1960; Steininger et al. 1971); Nógrád–Filakovo Basin: Lipovany ( Slovak Republic) ( Ondrejickova 1972); Váh Valley: Podylava ( Slovakia) ( Ctyroky 1959). Pannonian Basin: Budafok-Budapest ( Hungary) ( Báldi 1964);? Ottnangian: Nógrád–Filakovo Basin: Ipolytarnóc (Csepreghy- Meznerics 1967); Eastern Mátra Mountains: Illona Valley ( Hungary) ( Fözy & Leél-Össy 1985a, b); Burdigalian (early Miocene): Rhône Basin: Beaumadalier (Provence) ( Mongin 1952).

This species was variously recorded also from Karpatian and Badenian deposits: North Alpine Foreland Basin: Neuruppersdof at Laa ( Austria) ( Sieber 1960); Vienna Basin: Steinebrunn ( Austria), Hrušovany, Rudice ( Czech Republic); Eisenstadt Sopron Basin: Forchtenau ( Austria); Styrian Basin: Klockerberg at Wetzelsdorf ( Austria); Pannonian Basin: Pécsszabolcs ( Bohn-Havas 1973); Cserhát Mountains: Bán, Mátraszölös, Mátraverebély ( Hungary) (Csepreghy Meznerics 1969; Bohn-Havas 1973); Krško Basin: Gorenje Vrhpolje, Dolenja Brezovica ( Slovenia) ( Mikuž 2009). All these specimens are worn and/or fragmented and show traces of heavy bioersosion in many cases. Therefore, we interpret these shells as reworked specimens from older strata but cannot exclude that this species might indeed have persisted into the middle Miocene. This scenario, however, is rather unrealistic in respect to the large stratigraphic gap between both occurrences.

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 3. Measurements for shell morphology and whorl profiles (adopted from Allmon 1996) (SL = shell length, MD = maximum diameter, AA = apical angle, PA = pleural angle).

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 25. A–D. Allmonia paucicincta (Sacco, 1895), Burgschleinitz (Austria). A. Holotype, NHMW 1846/0010/0014. B1–B3. Collection Anton Breitenberger. C. NHMW 2018/0128/0002. D. NHMW 2018/0128/000. E. Turritella (Protoma) telleri Hilber, 1993, holotype, Ivanji dol (Slovenia), UMJG&P 1247. F–H. Allmonia cathedralis (Brongniart, 1823), Léognan (France). F1–F2. NHMW 1858/0018/0146. G. NHMW 1858/0018/0146. H. NHMW 1858/0018/0146. I–J. Allmonia funiculata (Borson, 1821), Torino Hills (Italy). I. NHMW 1862/0049/0026. J. NHMW 1854/0016/0209.