Pustulosia submitralis ( Eichwald, 1851 ) Harzhauser & Guzhov & Landau, 2023

Harzhauser, Mathias, Guzhov, Aleksandr & Landau, Bernard, 2023, A revision and nomenclator of the Cainozoic mudwhelks (Mollusca: Caenogastropoda: Batillariidae, Potamididae) of the Paratethys Sea (Europe, Asia), Zootaxa 5272 (1), pp. 1-241 : 158-162

Pustulosia submitralis ( Eichwald, 1851) nov. comb.

Figs 32A–F, 33A–H

* Cerith [ium]. submitrale m.— Eichwald 1851: 88, pl. 7, fig. 16.

[ Cerithium mitrale m.] var. bicostata — Eichwald 1851: 88, pl. 7, fig. 12.

Cerithium bicostatum m— Eichwald 1852: 2, pl. 7, fig. 12.

Cerithium submitrale — Eichwald 1852: 2, pl. 7, figs 16a–c.

Cerith [ium]. submitrale m.— Eichwald 1853: 156.

Cerithium bicostatum m— Eichwald 1853: 158.

Cerithium submitrale Eichw. — Andrae 1855: 296.

Cerithium nodoso-plicatum Ĥrn. — Ĥrnes 1855: 397, pl. 41, figs 19a–c [non 20 a–c = Theodisca biseriata ( Friedberg, 1914) ].

C [erithium]. Eichwaldi R. Hoern. u. Auing. In coll.— Hilber 1881: 184 [nomen nudum].

C [erithium]. Schaueri n. sp. — Hilber 1881: 184 [nomen nudum].

Cerithium eichwaldi R. Hoernes und Auinger nova forma — Hilber 1882: 7, pl. 1, figs 12, 13.

Cerithium Schaueri Hilber nova forma — Hilber 1882: 7, pl. 1, figs 15 [non fig. 14 = Theodisca biseriata or T. theodisca ].

Cerithium Vitense n. f.— Handmann 1883a: 172.

[ Cerithium Vitense ] var. costulata — Handmann 1883a: 172 [non Lamarck, 1822].

Potamides nodosoplicatus M. Hoern. — Friedberg 1914: 280, pl. 17, fig. 14.

Potamides Schaueri Hilb. var. Eichwaldi R. Hoern. i Auing.— Friedberg 1914: 286, pl. 17, figs 22–24.

Pirenella Schaueri Hilb. — Friedberg 1914: 284, pl. 17, fig. 18, text-fig. 62 [non fig. 19 = Theodisca biseriata or T. theodisca ].

Potamides melanopsiformis Auing. (in coll.)— Friedberg 1914: 281, pl. 17, figs 15–17.

Potamides petersi Auing. (in coll.)— Friedberg 1914: 288, pl. 17, figs 20–21.

Potamides (Pirenella) borsodensis n. sp. — Schréter 1929: 385, pl. 9, fig. 2.

Pirenella nodosoplicata (Ĥrnes, M.) — Sieber 1937: 479, pl. 24, fig. B5.

Pirenella Schaueri Hilber — Sieber 1937: 481, pl. 24, A1, 2, 4.

Pirenella picta Defr. var. melanopsiformis Auing. in coll.— Sieber 1937: 486, pl. 24, figs D 1, 2.

Potamides bicostatus Eichwald — Csepreghy-Meznerics 1950: 27, pl. 1, fig. 15.

Potamides petersi Auinger (in coll.)— Csepreghy-Meznerics 1950: 27, pl. 1, fig. 14.

Pirenella schaueri eichwaldi (R. Hoernes und Auinger in coll.) Hilber— Papp 1952: 115, pl. 3, figs 18–21.

Pirenella picta melanopsiformis (Auinger in coll.) Friedberg— Papp 1952: 116, pl. 3, figs 25–31.

Pirenella picta melanopsiformis (Auinger in coll.—Friedberg)— Papp 1953: 224.

Pirenella picta (Defr.) var. melanopsiformis Auing. (in coll.) — Švagrovský 1954: 12, pl. 1, figs 44–49, 52, 53.

Pyrenella [sic] nodosoplicata nodosoplicata (M. Ĥrn.) — Seneš 1955: 94, pl. 8, figs 6–8.

Pyrenella [sic] biseriata biseriata (Friedb.) — Seneš 1955: 95, pl. 8, figs 13–16.

Pyrenella [sic] schaueri schaueri (Hilb.) — Seneš 1955: 95, pl. 8, figs 9–12.

Pirenella nodosoplicata Ĥrnes, 1856 — Strausz 1955a: 57, 180, pl. 4, fig. 38, text-fig. 3, 9, 10.

Pirenella nodosoplicata petersi Auing. in Friedberg, 1928— Strausz 1955a: 64, 186, pl. 4, fig. 37, text-fig. 14.

Pirenella nodosoplicata eichwaldi (R. Hoernes Auinger) — Švagrovský 1959: 222, pl. 6, fig. 12.

Pirenella nodosoplicata biquadrata Hilber — Boda 1959: 618, pl. 26, fig. 14 [non Hilber, 1882].

Pirenella picta melanopsiformis (Auinger) — Pavlovsky 1960: 215, pl. 1, fig. 5a.

Pirenella schaueri schaueri (Hilber) — Pavlovsky 1960: 215, pl. 1, fig. 3a.

Pirenella nodosoplicata (Hoernes 1856) —Kojumdgieva in Kojumdgieva & Strachimirov 1960: 108, pl. 31, fig. 16.

Potamides (Pirenella) borsodiensis Schréter 1929 — Boda 1964: 132.

Pirenella biquadrata (Hilber) — Mariani & Papp 1966: 143, pl. 1, figs 6–7 [non Hilber, 1879a].

Cerithium vitense Handmann — Mariani & Papp 1966: 143, pl. 1, figs 8–10.

Potamides (Pirenella) moravicus borsodensis Schréter, 1929 — Strausz 1966: 148, pl. 10, fig. 11.

Pirenella picta melanopsiformis (Auinger) — Spajič 1966: 33, pl. 3, fig. 16.

Potamides (Pirenella) nodosoplicatus Ĥrnes, 1856 — Strausz 1966: 151, pl. 7, fig. 20.

Potamides (Pirenella) nodosoplicatus schaueri Hilber, 1882 — Strausz 1966: 152, text-fig 66.

Potamides (Pirenella) nodosoplicatus petersi Auinger (in Friedberg), 1928— Strausz 1966: 152, pl. 7, fig. 19.

Pirenella eichwaldi (Hoern. et Auing.) — Kókay 1966: 43, pl. 4, fig. 9.

Pirenella eichwaldi elongata (Lomn.) — Kókay 1966: 43, pl. 4, fig. 10.

Pirenella petersi (Auinger in coll.)—Hinculov in Iliescu et al. 1968: 126, pl. 30, figs 3–3b.

Pirenella nodosoplicata nodosoplicata (Ĥrnes, 1856) —Hinculov in Iliescu et al. 1968: 126, pl. 30, figs 4–6.

Pirenella nodosoplicata schaueri ( Hilber, 1882) —Hinculov in Iliescu et al. 1968: 127, pl. 30, figs 8a–b.

Potamides melanopsiformis (Auinger) Friedberg, 1914 — Zelinskaya et al. 1968: 158, pl. 39, figs 7–8.

Potamides schaueri ( Hilber, 1882) — Zelinskaya et al. 1968: 160, pl. 39, figs 21–22.

Potamides (Pirenella) schaueri Hilb. — Bałuk 1970: 117, pl. 10, figs 7–10.

Pirenella nodosoplicata (M. Hoern.) — Švagrovský 1970: 197, pl. 5, figs 4–5.

Pirenella borsodensis Schrét. var.— Kókay 1972: pl. 1, fig. 7.

Pirenella borsodensis Schréter 1929 — Steininger et al. 1973: 405, pl. 4, figs 5a–b.

Pirenella nodosoplicata (Ĥrnes) 1856 — Bohn-Havas 1973: 1045, pl. 4, fig. 7.

Pirenella nodosoplicata (Hoernes) — Ali-Zade 1974: 112, pl. 38, fig. 5 [non Ĥrnes, 1855].

Pirenella schaueri ( Hilber 1882) — Krach 1981: 60, pl. 16, fig. 17.

Pirenella aspera sp. n. — Švagrovský 1982: 22, pl. 6, figs 3–4.

Pirenella schaueri (Hilber) — Ionesi & Nicorici 1994: 61, pl. 4, fig. 25.

Potamides (Pirenella) nodosoplicatus (M. Hoernes, 1856) — Iljina 1993: 74, pl. 9, figs 15–16.

Bittium deforme Eichw. — Saulea 1956: 82, pl. 3, figs 20–21 [non Eichwald, 1829].

Pirenella nodosoplicata — Aguilà & García García 1997: 20, pl. 1, fig. 7.

Granulolabium bicinctum ( Brocchi, 1814) — Landau et al. 2013: 45 (pars), pl. 4, figs 3–4 [non Tiarapirenella bicincta ( Brocchi, 1814) ].

Potamides schaueri ( Hilber, 1882) — Harzhauser et al. 2018a: 159, figs 10I–K.

non Cerithium nodosoplicatum Hoern. ?— Sokolov 1899: 37, 80, pl. 4, figs 23–24 [= Theodisca biseriata ( Friedberg, 1914) ]. non Cerithium cf. nodosoplicatum Ĥrn. — Bajarunas 1910: 259 [= Theodisca biseriata ( Friedberg, 1914) ]. non Cerithium cf. nodosoplicatum M. Hoern. —Andrussov 1917: plate caption, pl. 15, fig. 24 [= Theodisca biseriata ( Friedberg,

1914)]. non Cerithium nodosoplicatum (Ĥrn.) Sok. — Ossipov 1932: 68, pl. 4, fig. 27 [= Theodisca biseriata ( Friedberg, 1914) ]. non Cerithium nodosoplicatum M. Ĥrn. — Kolesnikov 1935: 227, pl. 28, figs 22–24 [= Theodisca biseriata ( Friedberg,

1914)]. non Cerithium nodoso-plicatum Ĥrn. — Zhizhchenko 1934: 64, pl. 9, figs 21 [= Theodisca biseriata ( Friedberg, 1914) ]. non Potamides nodosoplicatus M. Ĥrnes — Zhizhchenko 1936: 205, pl. 21, figs 11–17 [= Theodisca biseriata ( Friedberg,

1914)]. non Pirenella Schaueri Hilber var. Eichwaldi R. Ĥrn. U. Auing.— Sieber 1937: 482, pl. 24, figs B 3, 4 [= Tiarapirenella tabulata (Ĥrnes, 1856) ]. non Cerithium nodosoplicatum M. Hoernes — Simionescu & Barbu 1940: 91, pl. 2, fig. 22 [= Theodisca biseriata ( Friedberg,

1914)]. non Pirenella nodosoplicata Hoernes — Jekelius 1944: 78, pl. 20, figs 1–9 [= Theodisca biseriata ( Friedberg, 1914) ]. non Potamides nodosoplicatus M. Ĥrnes — Csepreghy-Meznerics 1950: 27, pl. 1, fig. 12 [= Tiarapirenella bicincta ( Brocchi,

1814)]. non Potamides schaueri Hilber — Csepreghy-Meznerics 1954: 19, pl. 1, fig. 26 [? = Lampanella obliquistoma ( Seguenza,

1880)]. non Pirenella nodosoplicata (M. Hoernes) — Papp 1954: 41, pl. 6, figs 25–26 [= Theodisca biseriata ( Friedberg, 1914) ]. non Pirenella picta cf. melanopsiformis (Auinger in coll.) Friedberg— Papp 1954: 40, pl. 6, figs 13–15 [= Tiarapirenella bicincta

( Brocchi, 1814)]. non Pirenella nodosoplicata (M. Ĥrnes) — Švagrovský 1954: 5, pl. 1, figs 1–13 [= Theodisca biseriata ( Friedberg, 1914) ]. non Pirenella picta melanopsiformis Auing. (in coll.) — Strausz 1955a: 41, text-fig. 5 [= Tiaracerithium pictum (de Basterot,

1825)]. non P [otamides]. nodosoplicata nodosoplicata (Ĥrn.) — Sieber 1958: 136 [= Theodisca biseriata ( Friedberg, 1914) ]. non Pirenella nodosoplicata nodosoplicata (M. Ĥrnes) — Švagrovský 1960: 79, pl. 8, figs 1–3 [= Theodisca biseriata ( Friedberg,

1914)]. non Pirenella nodosoplicata eichwaldi (R. Hoernes et Auinger) — Švagrovský 1960: 81, pl. 8, figs 7–9 [= Theodisca theodisca

( Rolle, 1856)]. non Pirenella nodosoplicata M. Ĥrnes — Florei 1961: 683, pl. 7, fig. 49 [= Theodisca biseriata ( Friedberg, 1914) ]. non Pirenella nodosoplicata eichwaldi (R. Hoern. et Auing.) — Švagrovský 1964: 84, pl. 16, fig. 6 Theodisca theodisca ( Rolle,

1856)]. non Pirenella nodosoplicata (M. Hoernes) — Spajič 1966: 35, pl. 3, figs 5–6 [= Theodisca biseriata ( Friedberg, 1914) ]. non Potamides (Pirenella) pictus melanopsiformis Auinger (in Friedberg), 1928— Strausz 1966: 146, pl. 8, fig. 20 [= [=

Tiaracerithium pictum (de Basterot, 1825) ]. non Potamides nodosoplicatus (M. Ĥrnes, 1856) — Zelinskaya et al. 1968: 159, pl. 39, figs 15–16 [= Theodisca biseriata

( Friedberg, 1914)]. non Pirenella nodosoplicata nodosoplicata (Ĥrnes, 1856) — Kojumdgieva 1969: 91, pl. 32. figs 9–11 [= Theodisca biseriata

( Friedberg, 1914)]. non Pirenella nodosoplicata (M. Hörnes) — Plǎmǎdealǎ 1970: 118, pl. 8, figs 1–11 [= Theodisca biseriata ( Friedberg, 1914) ]. non Pirenella nodosoplicata nodosoplicata (M. Ĥrnes, 1856) — Švagrovský 1971: 337, pl. 57, figs 8–15 [= Theodisca biseriata

( Friedberg, 1914)]. non Pirenella nodosoplicata (M. Hoernes) — Papp 1974: 344, pl. 7, figs 19–22 [= Theodisca biseriata ( Friedberg, 1914) ]. non Pirenella nodosoplicata (Hoernes) — Ali-Zade 1974: 112, pl. 38, fig. 5 [= Theodisca biseriata ( Friedberg, 1914) ]. non Pirenella nodosoplicata (Hoernes) — Ionesi & Nicorici 1994: 62, pl. 5, fig. 17 [= Cerithiidae ]. non Pirenella nodosoplicata (Hoernes) — Munteanu & Munteanu 1997: 157, pl. 19, fig. 24 [= Theodisca biseriata ( Friedberg,

1914)]. non Pirenella nodosoplicata nodosoplicata (Ĥrnes) — Schultz 1998: 130, pl. 59, figs 8a–d [= Theodisca biseriata ( Friedberg,

1914)]. non Granulolabium nodosoplicatum (Ĥrnes, 1856) — Harzhauser & Kowalke 2002: 66, pl. 13, fig. 6 [non figs 7, 11 =

Tiaracerithium pictum (de Basterot, 1825) ]. non Potamides nodosoplicatus (Hoern.) — Ionesi 2006: 199, pl. 5, fig. 12 [= Theodisca biseriata ( Friedberg, 1914) ]. non Potamides nodosoplicatus ( Ĥrnes, 1855) — Landau et al. 2013: 54, pl. 4, figs 13–14 [= Theodisca theodisca ( Rolle,

1856)]. non Potamides nodosoplicatus ( Ĥrnes, 1855) — Filipescu et al. 2014: 78, fig. 10/5 [= Theodisca biseriata ( Friedberg, 1914) ].

Type material. Lectotype designated herein: PM SPSU 3 /318a (Paleontological Museum, Saint-Petersburg State University), SL: 16.5 mm, MD: 5.1 mm, illustrated in Eichwald (1851: pl. 7, fig. 16), Figs 33A 1 –A 2.

Type locality. Zalistsi ( Ukraine) .

Stratigraphy. Middle Miocene, Badenian.

Illustrated material. NHMW (no number), SL: 15.2 mm, MD: 5-9 mm, Bernhardsthal ( Austria), Middle Miocene, Badenian, illustrated in Harzhauser et al. (2018a: fig. 10I), Figs 33B 1 –B 2. RGM 794074, SL: 16.1 mm, MD: 6.1 mm, Tilkikaya ( Turkey), Middle Miocene, Serravallian, Figs 33C 1 –C 2. NHMW 1859/0045/094, SL: 13.2 mm, MD: 4.7 mm, Oles'ko (Lviv Region, Ukraine), Middle Miocene, Badenian, Figs 33D 1 –D 2. NHMW 1862/0018/0320, SL: 18.2 mm, MD: 6.2 mm, Hidas ( Hungary), Middle Miocene, Badenian, Figs 33E 1 –E 2. NHMW 1859/0045/094, SL: 15.6 mm, MD: 5.6 mm, Oles'ko (Lviv Region, Ukraine), Middle Miocene, Badenian, Figs 33F 1 –F 2. NHMW 1859/0042/0026, SL: 17.9 mm, MD: 6.4 mm, Hidas ( Hungary), Middle Miocene, Badenian, Figs 33G 1 –G 2. NHMW 1859/0042/0026, SL: 17.6 mm, MD: 6.2 mm, Hidas ( Hungary) Middle Miocene, Badenian, Figs 33H 1 –H 2. ZNG PAN A-I-50/683.1, SL: 21.5 mm, MD: 8.0 mm, lectotype (designated herein) of Potamides melanopsiformis Friedberg, 1914 , Małoszów ( Poland), Middle Miocene, Badenian, illustrated in Friedberg (1914, pl. 17, fig. 15), Figs 32A 1 –A 2. ZNG PAN A-I-50/683.2, SL: 16.1 mm, MD: 6.4 mm, paralectotype of Potamides melanopsiformis Friedberg, 1914 , Małoszów ( Poland), Middle Miocene, Badenian, illustrated in Friedberg (1914, pl. 17, fig. 17), Figs 32B 1 –B 2. ZNG PAN A-I-50/683.3, SL: 17.5 mm, MD: 7.5 mm, paralectotype of Potamides melanopsiformis Friedberg, 1914 , Małoszów ( Poland), Middle Miocene, Badenian, illustrated in Friedberg (1914, pl. 17, fig. 16), Figs 32C 1 –C 2. ZNG PAN A-I-50/705.1, SL: 12.0 mm, MD: 4.3 mm, Niskowa ( Poland), Middle Miocene, Badenian, lectotype (designated herein) of Potamides petersi Friedberg, 1914 , illustrated in Friedberg (1914: pl. 17, fig. 20) ( Figs 32D 1 –D 2). ZNG PAN A-I-50/705.2, SL: 23.0 mm, MD: 7.3 mm, Niskowa ( Poland), Middle Miocene, Badenian, paralectotype of Potamides petersi Friedberg, 1914 , illustrated in Friedberg (1914: pl. 17, fig. 21) ( Figs 32E 1 –E 2).

Additional material. Karpatian (Early Miocene): 47 spec., NHMW 1859/0045/0069, Niederkreuzstetten ( Austria) ; 48 spec., NHMW 1861/0050/0090, Niederkreuzstetten ( Austria) ; 15 spec., NHMW 1859/0045/0069, Niederkreuzstetten ( Austria) ; 3 spec., NHMW 1863/0015/0000, Helfens ( Austria) ; 1 spec., NHMW 1859/0045/0258, Weinsteig ( Austria) .

Badenian (Middle Miocene): 1. spec., NHMW 1999z0004/0017, Ritzing ( Austria), illustrated in Sieber (1937: pl. 24, fig. A1) as Pirenella schaueri Hilber. 1. spec., NHMW 1999z0004/0010, Ritzing ( Austria) , illustrated in Sieber (1937: pl. 24, fig.A2) as Pirenella schaueri Hilber. 1. spec., NHMW 1999z0004/0012, St. Veit an der Triesting ( Austria) , illustrated in Sieber (1937: pl. 24, fig.A4) as Pirenella schaueri Hilber. 1 spec., NHMW 1999z0004/0008, St. Veit an der Triesting ( Austria) , illustrated in Sieber (1937: pl. 24, fig. B3) as Pirenella schaueri eichwaldi Friedberg , 28 spec., NHMW1857/0014/0025, Steinebrunn ( Austria) ; 4 spec., NHMW 1866/0001/0902, Niederleis ( Austria) ; 1 spec., NHMW 1862/0001/0476, Vienna /P̂tzleinsdorf ( Austria) ; 3 spec., NHMW 1859/0045/0069, Enzersdorf ( Austria) ; 10 spec., NHMW 1855/0002/0090, Sedlec (Voitelsbrunn) ( Czech Republic) ; 8 spec., NHMW 1863/0015/0987, Hrušovany nad Jevišovkou ( Czech Republic) ; 6 spec., NHMW 1855/0002/0089, Mikulov ( Czech Republic) ; 126 spec., NHMW 1865/0025/0023, Hidas ( Hungary) ; 1 spec., NHMW 1862/0018/0009, Hidas ( Hungary) ; 61 spec., NHMW 1859/0042/0027, Hidas ( Hungary) ; 11 spec., NHMW 1870/0014/0004, Golubytsya ( Ukraine) ; 85 spec., NHMW 1870/0014/0004, Golubytsya ( Ukraine) ; 73 spec., NHMW 1859/0045/094, Oles'ko (Lviv Region, Ukraine) .

Revised description. Shell medium sized, slender conical to weakly cyrtoconoid, of 11 teleoconch whorls; apical angle ~31–35°, later decreasing to ~16–20°. Protoconch unknown. Early teleoconch whorls with two faint spiral cords at mid-whorl and along angulated periphery just above abapical suture. Mid-whorl cord becoming beaded on fourth teleoconch whorl. Third spiral cord appears at adapical suture on fourth teleoconch whorl becoming rapidly prominently beaded. Adapical spiral cord and mid-whorl spiral cord form two spiral rows of prominent, rounded beads on subsequent whorls. Abapical spiral cord usually largely covered by subsequent whorls but may form third row of beads in some specimens. Delicate spiral threads in interspaces between primaries in some specimens. Suture narrowly incised, undulating. Last whorl weakly convex to subcylindrical with moderately constricted base bearing about five broad spiral cords. Aperture subcircular-ovate; outer lip not thickened, strongly convex, occasionally granulae or short lirae developed within; siphonal canal short, moderately wide, slightly deflected to the left; anal canal narrowly incised. Columella excavated in adapical half, smooth. Columellar callus forming broad rim, sharply delimited from base.

Synonyms. Although Eichwald (1851, 1853) provided rather good illustrations of this species, subsequent authors have largely ignored it and established several new names for the same species. Of these, Cerithium eichwaldi Hilber, 1882 , Cerithium schaueri Hilber 1882 , Potamides petersi Friedberg, 1914 all were described from the type region of G. submitrale in the Polish-Ukrainian Fore-Carpathian Basin. Cerithium schaueri , as described by Hilber (1882: pl. 1, fig. 15) and Friedberg (1914: pl. 7, fig. 19), does not differ from the type of Pustulosia submitralis . Potamides petersi is characterized by large subquadratic beads but fully agrees with Pustulosia submitralis otherwise. Cerithium eichwaldi differs from the type by its prominent middle spiral cord. Potamides (Pirenella) borsodensis Schréter, 1929 , in contrast, represents a stout morph with a short spire. Cerithium nodosoplicatum Ĥrnes, 1855 is based on specimens from the Badenian of the Vienna Basin but cannot be distinguished from Pustulosia submitralis . According to the brief description by Handmann (1883b), Cerithium vitense might represent another subjective junior synonym (see vitense . Cerithium . Handmann, 1883a for discussion). Cerithium vitense costulata Handmann, 1883 is characterized by axially arranged beads, which are typical for Pustulosia submitralis . Similarly, Potamides melanopsiformis Friedberg, 1914 represents an aberrant morph with prominent beads at the adapical suture but somewhat ‘ bicinctum -like’ sculpture below.

Discussion. Pustulosia submitralis differs from Granulolabium plicatum distinctly in its bipartite sculpture of the early teleoconch whorls and the sculpture of wide-spaced, rounded beads. Some specimens with reduced mid-whorl beads are very close to Tiarapirenella bicincta . Both species, however, are distinctly distinguished by the early spire whorls, which bear spiral cords without beads in Tiarapirenella bicincta . Granulolabium moravicum is superficially similar but differs in its prominent secondary spiral cords and more gradate spire.

Distribution. Pustulosia submitralis appeared during the Early Miocene, when it is known from Central Paratethyan basins such as the North Alpine-Carpathian Foreland Basin and the Pannonian Basin. It persisted into the Serravallian equivalent of the Badenian (e.g., Olesko is of late Badenian age, see Anistratenko & Anistratenko 2007). Sarmatian records are rare and not always unambiguous. Maybe this species became extinct during the Badenian–Sarmatian Extinction Event ( Harzhauser & Piller 2007). In the Proto-Mediterranean Sea, this species is documented from the Langhian to the Serravallian.

Occurrences in the Miocene of the Proto-Mediterranean Sea may partly be obscured by misidentifications of Tiaracerithium pictum . Such a record was described by Landau et al. (2013) from the Serravallian of Turkey.

Central Paratethys. Karpatian (Early Miocene): North Alpine-Carpathian Foreland Basin: Laa ( Austria); Korneuburg Basin: Gebmanns, Helfens, Niederkreuzstetten, Ŗckersdorf, Weinsteig ( Austria); Pannonian Basin: Pereces, Várpalota ( Hungary) ( Strausz 1966; Kókay 1972; Steininger et al. 1973). Badenian (Middle Miocene): Voronyaky Hills: Golubytsya, Pidgirtsi, Yaseniv, Oles'ko (Lviv Region, ( Ukraine) ( Hilber 1882; Friedberg 1914); Roztocze Hills: Łychów ( Poland) ( Krach 1981); Pre-Carpathian area: Dzhuriv (Dżurów) (Ivano-Frankivsk Region, Ukraine); Polish-Ukrainian Fore-Carpathian Basin: Pidvolochys'k (= Zadniszówka), Zboriv, Zalistsi, Zboriv, ravine Zhabyak near Dzvynyacha, Zhukivtsi (Ternopil Region) ( Ukraine) ( Eichwald 1853; Friedberg 1914); Nowy Sącz Basin: Niskowa ( Poland) ( Friedberg 1914; Bałuk 1970); North Alpine-Carpathian Foredeep: Grund ( Austria) ( Ĥrnes 1855); Hrušovany nad Jevišovkou, Sedlec ( Czech Republic) (own data); Vienna Basin: Bernhardsthal, Berndorf (St. Veit an der Triesting), Niederleis, Steinebrunn ( Austria) ( Harzhauser et al. 2018a; Sieber 1937; Mariani & Papp 1966); Trstené pri Hornáde ( Švagrovský 1970), Borský Mikuláš ( Slovakia) ( Švagrovský, 1982), Mikulov, Sedlec ( Czech Republic) ( Ĥrnes, 1855; Sieber 1937); Oberpullendorf Basin: Ritzing ( Austria) ( Papp 1952); Danube Basin: Kuzmice ( Slovakia) ( Švagrovský 1960); Pannonian Basin: Samobor ( Croatia) ( Pavlovsky 1960), Herend–Márkó, Hidas, Pécsszabolcs, Várpalota, Sámsonháza ( Hungary) ( Csepreghy-Meznerics 1950, 1954; Strausz 1966); Southern Carpathians: Crivineni-Patârlagele ( Romania) ( Ionesi & Nicorici 1994); Mecsek Mountains ( Hungary) ( Strausz 1966); Mehadia Basin: Valea Bela Reca, Valea Globului, Valea Satului ( Romania) (Hinculov in Iliescu et al. 1968); Dacian Basin: Dobrusha ( Bulgaria) (Kojumdgieva in Kojumdgieva & Strachimirov 1960). Boziorul, Odăile ( Romania) ( Saulea 1956).

Eastern Paratethys. Occurrences from the early Chokrakian, listed by Iljina (1993), are based on misidentifications (own observations A.G.). Karaganian (Middle Miocene): Aksengir Mountain [very rounded] (Mangyshlak, Kazakhsan). Konkian: Ciscaucasia: ravine Podvesnaya (near Tul'skii, Adygea), Privol'nyi (Stavropol Krai, Russia); Transcaucasia: Haçaparaq (Nakhchivan Republic, Azerbaijan) ( Ali-Zade 1974). Transcaspia: Aksengir Mountain (Mangyshlak, Kazakhstan) ( Iljina 1993). Sarmatian, Volhynian: Ciscaucasia: Privol'nyi, Razdol'nyi (Stavropol Krai, Russia) (own data).

Proto-Mediterranean Sea. Langhian (Middle Miocene): Vallès-Penedès Basin: Bellaterra ( Spain) ( Aguilà & García García, 1997); Serravallian: Karaman Basin: Tilkikaya ( Turkey) ( Landau et al. 2013).