taxonID	type	description	language	source
03D787A4FFE2FFE3FF3FFC6AFEA59000.taxon	type_taxon	Type species. Scala elata Thiele, 1925; original designation by Thiele 1928: 92). Present-day, Indian Ocean.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFE2FFE3FF3FFC6AFEA59000.taxon	diagnosis	Diagnosis. “ Shell with broad axial ribs; no intritacalx; protoconch conical, of 3 – 4 whorls, with distinct axial ribs and sometimes interstitial spiral striae, often dark in colour. ” (Kilburn 1985: 295).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFE2FFE3FF3FFC6AFEA59000.taxon	discussion	Discussion. This genus as currently understood might need revision. The type species, Opaliopsis opalina (Dall, 1927), from the western Atlantic, and Opaliopsis hiranoi (Shikama, 1962), from the Indo-West Pacific are turreted shells with low whorls, moderately impressed suture and axial sculpture of weakly raised ribs with weak spiral sculpture. These species have a marked basal angulation with a prominent peribasal cord (e. g., Weil et al. 1999: figs 350, 393; Andrade et al. 2011: fig. 13 D). Other species treated as Opaliopsis by Andrade et al. (2011) and Pimenta et al. (2018) differ in their strong spiral sculpture and strongly raised varices [e. g., ‘ Opaliopsis’ atlantis (Clench & Turner, 1952)]. The Miocene Opaliopsis compacta nov sp. and the Pliocene Opaliopsis savii (Bogi & Cauli, 1998) and Opaliopsis cf. savii (Bogi & Cauli, 1998), described by Landau et al. (2006: pl. 5, figs 1 – 2), have stocky, subcylindrical whorls and are reminiscent of the extant Opaliopsis rabalaisi García, 2005 from the Gulf of Mexico, which however has more prominent varices. Thus, the placement of the Paratethyan species in Opaliopsis is tentative.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFE2FFEDFF3FF978FB2294D9.taxon	description	Figs 1 A – B, 2 H	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFE2FFEDFF3FF978FB2294D9.taxon	materials_examined	Type material. Holotype, NHMW 2024 / 0285 / 0001, SL: 4.8 mm, MD: 2.0 mm, Porzteich at Břeclav (Czech Republic), Figs 1 A 1 – A 2, 2 H. Paratype, NHMW 2024 / 0285 / 0002, SL: 3.9 mm, MD: 1.6 mm, Porzteich at Břeclav (Czech Republic), Figs 1 B 1 – B 2. Type locality. Porzteich at Břeclav (Czech Republic), Vienna Basin. Type stratum. Baden Formation. Age. Middle Miocene, middle Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFE2FFEDFF3FF978FB2294D9.taxon	etymology	Etymology. From compactus (Latin for compact, stout), referring to the shape.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFE2FFEDFF3FF978FB2294D9.taxon	diagnosis	Diagnosis. Very small, moderately slender, faintly cyrtoconoid shell with relatively low subcylindrical whorls, deeply incised suture and prominent axial ribs separated by interspaces of about equal width, overrun by six much weaker spiral cords, aperture subcircular, small, duplex.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFE2FFEDFF3FF978FB2294D9.taxon	description	Description. Shell very small, moderately slender turriculate, faintly cyrtoconoid, apical angle ~ 23 °). Protoconch unknown. Teleoconch of at least seven relatively low whorls. Early teleoconch whorls weakly convex, later whorls becoming subcylindrical. Suture deeply impressed, slightly undulating. Sculpture of prominent, orthocline axial ribs, not aligned axially (13: 13: 14), separated by slightly wider interspaces, overrun by six very weak, somewhat blurred spiral cords, most prominent in axial interspaces, separated by narrower grooves. Last whorl attaining about 35 % of total height, distinct basal angulation delimited by prominent peribasal cord, basal disc with slightly scalloped margin, bearing weak axial ribs and two indistinct spiral cords. Aperture small, subcircular. Outer lip slightly thickened. Columellar and parietal callus forming continuous, narrow rim, detached from base. Peristome duplex. Umbilicus reduced to very narrow chink.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFE2FFEDFF3FF978FB2294D9.taxon	discussion	Discussion. Opaliopsis savii (Bogi & Cauli, 1998), from the Pliocene of Italy, has a higher last whorl, prosocline axial ribs and more prominent spiral cords (see Bogi & Cauli 1998: figs 4 – 5). Opaliopsis cf. savii (Bogi & Cauli, 1998), from the Pliocene of Spain, differs also in its more prominent spiral sculpture but is otherwise very similar to Opaliopsis compacta (see Landau et al. 2006: pl. 5, figs 1 – 2). Opaliopsis compacta sp. nov. is reminiscent of the Mediterranean Punctiscala cerigottana (Sturany, 1896) but lacks a punctate microsculpture typical of that genus (see Weil et al. 1999: fig. 122). Paleoenvironment. Unknown.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFE2FFEDFF3FF978FB2294D9.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienna Basin: Porzteich at Břeclav (Czech Republic) (hoc opus). Genus Papuliscala de Boury, 1911	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFE2FFEDFF3FF978FB2294D9.taxon	type_taxon	Type species. Acirsa praelonga Jeffreys, 1877; original designation. Present-day, eastern Atlantic.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFE2FFEDFF3FF978FB2294D9.taxon	diagnosis	Diagnosis. “ Shell turriculate, teleoconch with several whorls of somewhat rounded outline; with or without carina. Teleoconch sculpture of usually equally strong axial and spiral cords, forming weak to strong nodules on intersections; with development of peripheral basal spiral cord, without umbilicus; basal area with weak to nearly obsolete sculpture. ” (Pimenta et al. 2018: 20).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFE2FFEDFF3FF978FB2294D9.taxon	discussion	Discussion. Miocene species of this genus have sometimes been misplaced in Triphoridae by Boettger (1907), Zilch (1934) and Bałuk (2006). Therefore, Papuliscala was not reported from the Paratethys Sea until now. This is surprising considering its remarkable diversity of six species documented herein. Similarly, the extant diversity of the genus has been strongly underrated until the paper by Hoffman et al. (2020), which more than doubled the number of known species. Currently, 19 extant species are listed by MolluscaBase eds (2024 b). Papuliscala was placed in Nystiellinae Clench & Turner, 1952 by Bouchet & Warén (1986: 494) on the base of an axially ribbed, multispiral protoconch of one Oligocene species. All extant species have paucispiral protoconchs. Papuliscala koeneni (A. W. Janssen, 1967), from the Miocene of the North Sea, also has a multispiral protoconch (see A. W. Janssen, 1967, pl. 14, fig. 2). Herein, we document Papuliscala aquaensis sp. nov. as a further fossil species with a ribbed, multispiral protoconch. This is one of the genera that has lost its planktotrophic mode of development over time so that all extant species have paucispiral protoconchs (see also Landau et al. in press).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFECFFEDFF3FFD43FA2491C9.taxon	description	Figs 3 A – B, 2 A, 4 A	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFECFFEDFF3FFD43FA2491C9.taxon	materials_examined	Type material. Holotype, NHMW 1869 / 0001 / 0209 a, SL: 3.3 mm, MD: 1.1 mm, Baden (Austria), Figs 3 A 1 – A 2, 3 A, 4 A. Paratype, NHMW 1869 / 0001 / 0209 b, SL: 3.2 mm, MD: 0.9 mm, Baden (Austria), Figs 3 B 1 – B 2. Type locality. Baden (Austria), Vienna Basin. Type stratum. Baden Formation. Age. Middle Miocene, middle Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFECFFEDFF3FFD43FA2491C9.taxon	etymology	Etymology. Referring to Aquae, the Roman name for the city Baden in Austria,	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFECFFEDFF3FFD43FA2491C9.taxon	diagnosis	Diagnosis. Very small, glossy, slender conical with turreted multispiral protoconch and biangulate teleoconch whorls with broad, steep subsutural ramp and prominent axial ribs bearing small nodes at sculptural intersections.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFECFFEDFF3FFD43FA2491C9.taxon	description	Description. Shell very small, slender conical (apical angle ~ 16 – 19 °). Protoconch slender, turreted of 4.5 smooth, strongly convex whorls, with low nucleus. Last protoconch whorl widening with rounded mid-whorl angulation and delicate, strongly opisthocline axial riblets. Teleoconch of six whorls. Early teleoconch whorls strongly convex with prominent, convex, orthocline axial ribs. Second and third teleoconch whorls successively developing moderately steep, flattened subsutural ramp with angled shoulder, subcylindrical periphery, strongly constricted below basal angulation. Suture deeply impressed, linear. Sculpture of moderately prominent, convex orthocline axial ribs, not aligned axially, separated by wider interspaces (11: 12 / 13: 12 / 13). Spiral sculpture of two indistinct cords delimiting angulations forming small nodes at intersections with ribs. Shell surface smooth, glossy. Marked basal angulation delimited by prominent peribasal cord and weaker mid-basal cord. Base nearly flat, smooth. Aperture ovate. Outer lip weakly thickened. Columellar parietal callus delicate, adherent. Peristome simple. No umbilicus. Discussions. The protoconch with axial riblets on the last whorl confirms placement in Nystiellinae (Bouchet & Warén 1986: 494). Papuliscala aquaensis sp. nov. is reminiscent of the Pliocene to Recent Mediterranean Papuliscala tavianii Bouchet & Warén, 1986, which differs in its paucispiral protoconch, more prominent axial ribs and well defined spiral cords (see Bouchet & Warén, 1986: fig. 1163, Landau et al. 2006: pl. 5, figs 3 – 4). Papuliscala koeneni (Janssen, 1967), from the Middle Miocene of the North Sea, also has a multispiral protoconch with axial riblets on the last two whorls but is distinguished from the Paratethyan species by its more prominent spiral cords (see Janssen 1967: pl. 14, fig. 1), with a third cord intercalated between the two primaries on the last two whorls. Papuliscala ambulator Lozouet 1999, from the late Oligocene of France, has a multispiral protoconch but differs in its higher number of spiral cords and convex whorl profile (see Lozouet 1999: pl. 10, fig. 10). Paleoenvironment. Occurrences in the Baden Formation of the Vienna Basin suggest middle to outer neritic settings in up to 250 m water depth (Kranner et al. 2021), which agrees with the environmental requirements of extant species (Bouchet & Warén 1986; Hoffman et al. 2020).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFECFFEDFF3FFD43FA2491C9.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienna Basin: Baden (Austria) (hoc opus).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFEFFFEFFF3FFAE3FE37944D.taxon	description	Figs 2 E, 3 C 1 – C 2, 4 B	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFEFFFEFFF3FFAE3FE37944D.taxon	materials_examined	Type material. Holotype, NHMW 1871 / 0010 / 0094 a, SL: 6.3 mm, MD: 2.0 mm, Porzteich at Břeclav (Czech Republic), Figs 2 E, 3 C 1 – C 2, 4 B. Type locality. Porzteich at Břeclav (Czech Republic), Vienna Basin. Type stratum. Baden Formation. Age. Middle Miocene, middle Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFEFFFEFFF3FFAE3FE37944D.taxon	etymology	Etymology. In honor of Leon Hoffman (Senckenberg Research Institute, Wilhelmshaven, Germany), in recognition of his contributions in malaocology.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFEFFFEFFF3FFAE3FE37944D.taxon	diagnosis	Diagnosis. Small, slender turreted shell with protoconch of 2.5 whorls, strongly convex teleoconch whorls, prominent axial ribs especially over subsutural ramp, overrun by six spiral cords forming low rounded nodes on abapical three spiral cords.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFEFFFEFFF3FFAE3FE37944D.taxon	description	Description. Shell small, slender turreted (apical angle ~ 17 °). Protoconch turreted of 2.5 smooth, weakly convex whorls with small nucleus. Teleoconch of more than seven whorls with narrow, moderately steep, weakly convex subsutural ramp, convex periphery placed just above mid-whorl. Suture deeply impressed, linear. Sculpture of prominent, convex, moderately widely spaced, weakly prosocline axial ribs (9: 12:?), separated by slightly wider interspaces. Axial ribs overrun by six broad, convex, somewhat blurred spiral cords, separated by narrower interspaces. Axial ribs most prominent on subsutural ramp, with comparatively weak spiral cords. Abapical three spiral cords forming prominent, low, rounded nodes at intersections. Last whorl poorly preserved; basal sculpture with one, possibly two very weak basal cords.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFEFFFEFFF3FFAE3FE37944D.taxon	discussion	Discussion. Differs from Papuliscala subsoluta (Boettger, 1907) in the higher position of the periphery, the fewer, comparatively blurred spiral cords, and the distinctly less clearly defined rectangular sculpture. The extant Papuliscala elongata (Watson, 1881), from the western and eastern Atlantic, is slenderer and has narrower, sharper axial ribs, and much narrower spiral cords. It also differs in having a protoconch of only two whorls with a big bulbous first whorl (see Brunetti & Soccio 2006: fig. 2 b; Pimenta et al. 2018: fig. 15 E). They also differ in the greater number of more strongly developed cords on the base of P. subsoluta. Paleoenvironment. Unknown. Extant Papuliscala species are found in deep water environments (Bouchet & Warén, 1986).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFEFFFEFFF3FFAE3FE37944D.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienna Basin: Porzteich at Břeclav (Czech Republic) (hoc opus).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFEEFFEFFF3FFD37FC02908A.taxon	description	Figs 2 C, 5 C – D	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFEEFFEFFF3FFD37FC02908A.taxon	materials_examined	Type material. Holotype, NHMW 1871 / 0010 / 0094 b, SL: 4.5 mm, 2.8 MD: mm, Porzteich at Břeclav (Czech Republic), Figs 2 C, 5 C 1 – C 2. Paratype, BkK-G 1206, SL: 3.3 mm, MD: 1.2 mm, Korytnica (Poland), illustrated in Bałuk (2006: pl. 9, fig. 10), Fig. 5 D. Type locality. Porzteich at Břeclav (Czech Republic), Vienna Basin. Type stratum. Baden Formation. Age. Middle Miocene, middle Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFEEFFEFFF3FFD37FC02908A.taxon	etymology	Etymology. In honor of Chris Mays, paleobotanist at the NHMW.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFEEFFEFFF3FFD37FC02908A.taxon	diagnosis	Diagnosis. Very small to small, slender shell with turreted, multispiral protoconch, early teleoconch whorls with rounded shoulder, later whorls convex with deeply incised suture and cancellate sculpture of prominent axial ribs and five spiral cords, adapical cord placed immediately below suture, abapical cord partly obscured by suture.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFEEFFEFFF3FFD37FC02908A.taxon	description	Description. Shell very small to small, slender (apical angle ~ 18 °). Protoconch turreted, 3.5 high, weakly convex whorls with low nucleus (Fig. 5 D). Early teleoconch whorls moderately high with broad, steep, weakly convex subsutural ramp, rounded shoulder, narrow subcylindrical periphery, constricted below. Sculpture of prominent, narrow, weakly prosocline axial ribs separated by wider interspaces (9: 12:?), overrun by weak spiral cord on subsutural ramp and two prominent cords along periphery. Later whorls with weakly convex subsutural ramp and convex periphery. Three prominent spiral cords and weaker cord at adapical suture. Fifth spiral cord at abapical suture partly covered by subsequent whorl, forming cancellate sculpture with small nodes developed at intersections. Suture deeply impressed, linear. Last whorl not preserved.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFEEFFEFFF3FFD37FC02908A.taxon	discussion	Discussion. This species is almost identical to Papuliscala presselierensis Landau, Ceulemans & Van Dingenen, 2018, from the Tortonian of France, in general shape and sculpture but differs in its turreted, multispiral protoconch. It differs from Papuliscala parvicancellata sp. nov. in having fewer spiral cords and the pronounced subsutural ramp. Paleoenvironment. Unknown. Extant Papuliscala species are found in deep water environments (Bouchet & Warén, 1986).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFEEFFEFFF3FFD37FC02908A.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Korytnica Basin: Korytnica (Poland) (Bałuk 2006); Vienna Basin: Porzteich at Břeclav (Czech Republic) (hoc opus).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFEEFFE8FF3FF8F2FD52905D.taxon	description	Figs 2 D, 3 D – E	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFEEFFE8FF3FF8F2FD52905D.taxon	materials_examined	Type material. Holotype, NHMW 1863 / 0015 / 0804, SL: 4.7 mm, MD: 2.5 mm, Niederleis (Austria), Figs 2 D, 3 D 1 – D 2. Paratype, BkK-G 1209, SL: 4.6 mm, MD: 2.0 mm, Korytnica (Poland), illustrated in Bałuk (2006: pl. 9, fig. 13), Fig. 3 E. Type locality. Niederleis (Austria), Vienna Basin. Type stratum. Baden Formation. Age. Middle Miocene, middle Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFEEFFE8FF3FF8F2FD52905D.taxon	etymology	Etymology. After parvus (Latin for small) and cancellata (Latin for cancellate), referring to the delicate, cancellate pattern.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFEEFFE8FF3FF8F2FD52905D.taxon	diagnosis	Diagnosis. Very small to small, slender turreted shell with moderately high convex whorls and prominent finely cancellate sculpture of seven spiral cords with weak nodes at intersection with ribs.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFEEFFE8FF3FF8F2FD52905D.taxon	description	Description. Shell very small to small, slender turreted (apical angle ~ 15 °). Protoconch and early teleoconch whorls unknown. Teleoconch whorls moderately high and convex with narrow, weakly convex subsutural ramp, regularly convex below, periphery just above mid-whorl. Suture impressed, linear. Sculpture of prominent, narrow, weakly prosocline axial ribs (?: 15: 16), separated by wider interspaces, overrun by seven subequal spiral cords, forming finely cancellate sculpture, with small nodes developed at intersections. Base convex with prominent peribasal cord and five spiral cords. Aperture ovate, columella weakly concave. Outer lip not preserved. Columellar and parietal callus delicate.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFEEFFE8FF3FF8F2FD52905D.taxon	discussion	Discussion. This species is based on two fragmentary specimens. Nevertheless, the available material is highly characteristic and allows for a description. It is the most finely cancellate Papuliscala species in the Paratethyan assemblages. The morphologically closest species is Papuliscala plioligustica (Sacco, 1891), from the Pliocene of the Mediterranean Sea. It is almost identical in shape and sculpture, but differs only in the higher number of spiral cords and axial ribs in the Pliocene species, resulting in even finer cancellate sculpture (see Ferrero Mortara et al. 1984: pl. 6, fig. 12). Papuliscala parvicancellata nov. sp differs from most congeners, such as Papuliscala presselierensis Landau, Ceulemans & Van Dingenen, 2018, from the Tortonian of France, in its greater number of spiral cords. The extant Papuliscala luuki Hoffman, Gofas & Freiwald, 2020, from the Great Meteor Seamount, has comparable sculpture but its axial ribs are more delicate and the suture is more deeply incised (Hoffman et al. 2020: figs 26 – 28). Papuliscala gretae Brunetti & Soccio, 2006, from the Pliocene of Italy, is reminiscent of the Paratethyan species, but develops only five spiral cords and has lower whorls (see Brunetti & Soccio 2006: figs 1, 2 a). The extant Papuliscala cerithielloides Bouchet & Warén, 1986 also has similar sculpture, but differs in its more convex whorls and it lacks nodes at the sculptural intersections (see Bouchet & Warén 1986: figs 1164). Papuliscala voorthuyseni (Anderson, 1964) from the Middle Miocene of Dingden (Germany) differs in its spiral cords, which distinctly overrun the more numerous axial ribs (see Anderson 1964: pl. 14, fig. 123). Papuliscala voorthuyseni (Anderson, 1964) (sensu Janssen 1984), from the Hemmoorian of the North Sea, is also reminiscent of P. parvicancellata in its very slender shape and cancellate sculpture but develops only five spiral cords (see Janssen 1984: pl. 50, figs 10 – 11). Paleoenvironment. The Niederleis section, sediment and biota from coastal and lagoonal environments not exceeding 30 m water depth were transported by tempestites into offshore settings (Mandic et al. 2002). Extant Papuliscala species are found in deep water environments (Bouchet & Warén, 1986).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFEEFFE8FF3FF8F2FD52905D.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Korytnica Basin: Korytnica (Poland) (Bałuk 2006); Vienna Basin: Niederleis (Austria) (hoc opus).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFE9FFEBFF3FF9C7FE37976D.taxon	description	Figs 2 B, 3 F 1 – F 2	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFE9FFEBFF3FF9C7FE37976D.taxon	materials_examined	Type material. Holotype, NHMW 1871 / 0010 / 0129, SL: 3.6 mm, MD: 1.2 mm, Porzteich at Břeclav (Czech Republic), Figs 2 B, 3 F 1 – F 2. Type locality. Porzteich at Břeclav (Czech Republic), Vienna Basin. Type stratum. Baden Formation. Age. Middle Miocene, middle Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFE9FFEBFF3FF9C7FE37976D.taxon	etymology	Etymology. Referring to Papuliscala elongata (Watson, 1881), which might be a closely related species.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFE9FFEBFF3FF9C7FE37976D.taxon	diagnosis	Diagnosis. Very small, slender turreted shell with markedly angled teleoconch whorls, moderately steep subsutural ramp with prominent axial ribs and cancellate sculpture below subsutural ramp.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFE9FFEBFF3FF9C7FE37976D.taxon	description	Description. Shell very small, slender turreted (apical angle ~ 18 °). Protoconch and early teleoconch whorls unknown. Teleoconch of about eight whorls. Whorl profile with moderately steep, slightly concave subsutural ramp, marked shoulder angulation, subcylindrical periphery, constricted below. Suture deeply impressed, linear. Sculpture of prominent, narrow, orthocline axial ribs, not aligned axially, separated by wider interspaces (12: 14: 12). Axial ribs overrun by three prominent spiral cords, forming horizontally elongate cancellate sculpture below shoulder, with small nodes developed at sculptural intersections. Nodes along shoulder angulation weakly pointed. Subsutural ramp devoid of spiral sculpture. Finer spiral cord at abapical suture partly covered by subsequent whorl. Last whorl attaining about 35 % of total height, moderately angled basal angulation delimited by narrow but prominent peribasal cord, base flattened bearing three further cords. Aperture ovate. Outer lip not preserved. Columellar and parietal callus indistinct, adherent. No umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFE9FFEBFF3FF9C7FE37976D.taxon	discussion	Discussion. This species is characterized by its sharp shoulder angulation with small, pointed nodes, prominent axial ribs on the subsutural ramp and cancellate sculpture below. Papuliscala elongata (Watson, 1881), from the western and eastern Atlantic, is exceptionally similar concerning the marked shoulder angulation and cancellate sculpture. They are distinguished by the slenderer shape of Papuliscala elongata (see Brunetti & Soccio 2006: fig. 2 b; Pimenta et al. 2018: fig. 15). The base of P. elongata is smooth, or almost so, whereas the base in P. praeelongata sp. nov. has three cords. Paleoenvironment. Unknown. Extant Papuliscala species are found in deep water environments (Bouchet & Warén, 1986).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFE9FFEBFF3FF9C7FE37976D.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienna Basin: Porzteich at Břeclav (Czech Republic) (hoc opus).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFEAFFF4FF3FFED7FEED9785.taxon	description	Figs 2 F, 5 A – B * Cerithiopsis (Metaxia) subsoluta n. sp. — Boettger 1907: 148.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFEAFFF4FF3FFED7FEED9785.taxon	materials_examined	Type material. Holotype, SMF 12 2449, SL: 6.0 mm, MD: 2.0 mm, CoŞteiu de Sus (Romania), illustrated in Zilch (1934: pl. 9, fig. 51). Illustrated material. BkK-G 1207, SL: 7.8 mm, MD: 2.0 mm, Korytnica (Poland), illustrated in Bałuk (2006: pl. 9, fig. 11), Figs 5 A 1 – A 2. NHMW 1871 / 0010 / 0094 c, SL: 6.0 mm, MD: 2.6 mm, Porzteich at Břeclav (Czech Republic), Figs 2 F, 5 B. Revised description. Shell small, very slender turreted of at least nine teleoconch whorls (apical angle ~ 13 °). Protoconch unknown. Early teleoconch whorls strongly convex with broad, weakly convex subsutural ramp. Prominent, moderately widely spaced, weakly prosocline axial ribs. Spiral sculpture of prominent, broad subsutural cord, seven prominent, strongly convex cords below and one weaker cord at abapical suture. Later whorls becoming allometrically higher and less convex, with broad, steep, weakly convex and poorly delimited subsutural ramp, faint angulation, periphery just below mid-whorl. Suture deeply impressed, linear. Sculpture of prominent, widely spaced, weakly prosocline axial ribs (~ 10: ~ 10: ~ 11), separated by wider interspaces, overrun by up to eight broad, strap-like spiral cords, separated by narrower groove-like interspaces, forming low, rounded nodes at intersections. Basal angulation delimited by prominent peribasal cord. Base weakly concave with four prominent spiral cords. Aperture ovate. Outer lip thin, weakly convex. Columellar and parietal callus indistinct. Peristome simple. No umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFEAFFF4FF3FFED7FEED9785.taxon	discussion	Discussion. Papuliscala subsoluta (Boettger, 1907) differs from all its Paratethyan congeners in having broad, strap-like spiral cords separated by narrow grooves, so that the sculpture is not cancellate as seen in most species. The basal cords are also stronger than in any other Paratethyan Papuliscala species. It was erroneously placed in the Triphoridae genus Metaxia Monterosato, 1884 by Boettger (1907) and subsequent authors but the prominent axial ribs exclude a relation with Triphoridae. Paleoenvironment. Unknown. Extant Papuliscala species are found in deep water environments (Bouchet & Warén, 1986).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFEAFFF4FF3FFED7FEED9785.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Korytnica Basin: Korytnica (Poland) (Bałuk 2006); Vienna Basin: Porzteich at Břeclav (Czech Republic) (hoc opus); Făget Basin: CoŞteiu de Sus (Romania) (Boettger 1907).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF5FFF4FF3FFDFFFAA8959D.taxon	type_taxon	Type species. Scalaria borealis Lyell, 1841, by monotypy. Pleistocene, Sweden.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF5FFF4FF3FFDFFFAA8959D.taxon	diagnosis	Diagnosis. “ Shape stocky; whorls anteriorly convex; grooved suture; fine spiral cords; subobsolete axial ribs, fading on the last subangular whorl; base without disc, but nevertheless limited by a distinct periphery. Aperture oval, slightly sloping forwards; peristome rather thin, oblique, not bordered on the outside; columellar margin excavated, slightly reflected outwards and fully covering the umbilical region, ” (translated from Cossmann 1912: 95). “ Species of Acirsa are characterized by a solid shell with broad axial ribs, and the absence of a basal cord and basal disc. Spiral sculpture of incised spiral groves is usually present. ” (Bouchet & Warén 1986: 526).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF5FFF5FF3FFB86FC9392B8.taxon	description	Figs 4 E, 6 A – B, 7 E, I	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF5FFF5FF3FFB86FC9392B8.taxon	materials_examined	Type material. Holotype, NHMW 1873 / 0026 / 0043, SL: 19.3 mm, MD: 7.3 mm, Lăpugiu de Sus (Romania), Figs 4 E, 6 A 1 – A 2. Paratypes, NHMW 1873 / 0026 / 0043 a, SL: 14.4 mm, MD: 5.0 mm, Lăpugiu de Sus (Romania), Fig. 6 B. NHMW 1873 / 0026 / 0043 b, SL: 15.9 mm, MD: 5.6 mm, Lăpugiu de Sus (Romania), Fig. 7 I. Type locality. Lăpugiu de Sus (Romania), Făget Basin. Type stratum. Silt and clay of the Dej Formation. Age. Middle Miocene, Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF5FFF5FF3FFB86FC9392B8.taxon	etymology	Etymology. Referring to the fusiform shape.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF5FFF5FF3FFB86FC9392B8.taxon	diagnosis	Diagnosis. Medium-sized, moderately slender fusiform shell with elevated protoconch with conical outline of five convex whorls with deep suture and strongly teleoconch convex whorls with sigmoidal-opisthocyrt axial ribs overrun by numerous prominent, narrow spiral cords separated by narrow grooves; cords on last whorl subobsolete mid-whorl.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF5FFF5FF3FFB86FC9392B8.taxon	description	Description. Shell medium-sized, moderately slender fusiform, rather thin-shelled (apical angle 23 °). Protoconch multispiral, highly elevated, of five strongly convex whorls with small, pointed nucleus. Last two whorls with faint mid-whorl angulation, slightly constricted below, with some sinuous axial riblets on the last quarter whorl. Teleoconch of seven strongly convex whorls with deeply impressed, linear suture. First teleoconch whorl with about eight narrow spiral cords separated by wider interspaces crossing narrow, widely spaced, orthocline axial ribs. Axial ribs strengthening on later whorls, forming prominent, convex, sigmoidal to opisthocyrt ribs, separated by wider interspaces (20: 17: 20). Spiral cords increasing in number by intercalations of secondary cords (about 15 spiral cords on penultimate whorl), overrunning ribs and forming delicate nodes at intersections. Spiral cords subobsolete mid last whorl. Last whorl attaining about 42 % of total height. Rounded transition into moderately constricted base bearing about 12 prominent spiral cords. Aperture partly missing; ovate, columella weakly concave. Columellar callus forming thin, adherent rim, no parietal callus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF5FFF5FF3FFB86FC9392B8.taxon	discussion	Discussion. Acirsa exvaricula Sacco, 1891, from the Tortonian of S. Agata (Italy), differs in its slenderer shape, less convex whorls and straight, weakly prosocline axial ribs (see Sacco 1891: pl. 2, fig. 102). Acirsa miopedemontana Sacco, 1891, from the Burdigalian of Baldissero (Italy), has comparable outline with strongly convex whorls but lacks prominent axial ribs (Sacco 1891: pl. 2, fig. 103). Paleoenvironment. Unknown.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF5FFF5FF3FFB86FC9392B8.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienna Basin: Bad Vöslau (Austria) (hoc opus); Făget Basin: Lăpugiu de Sus (Romania) (hoc opus).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF4FFF6FF3FFAB5FF609032.taxon	description	Figs 6 C 1 – C 2, 7 F	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF4FFF6FF3FFAB5FF609032.taxon	materials_examined	Type material. Holotype, NHMW 1860 / 0001 / 0289, SL: 14.5 mm, MD: 4.9 mm, Steinebrunn (Austria), Figs 6 E 1 – E 2, 7 F Type locality. Steinebrunn (Austria), Vienna Basin. Type stratum. Silt sand of the Baden Formation. Age. Middle Miocene, Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF4FFF6FF3FFAB5FF609032.taxon	etymology	Etymology. In honor of Efterpi Koskeridou (National and Kapodistrian University of Athens, Greece), in recognition of her contributions to Mediterranean paleontology.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF4FFF6FF3FFAB5FF609032.taxon	diagnosis	Diagnosis. Medium-sized, moderately slender, conical shell with weakly convex whorls, faint subsutural cord and opisthocline axial ribs with delicate spiral threads in interspaces, angled columella.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF4FFF6FF3FFAB5FF609032.taxon	description	Description. Shell medium-sized, moderately slender (apical angle ~ 25 °). Protoconch unknown. Teleoconch of about 10 whorls. Whorl profile weakly convex with steep, weakly concave subsutural ramp, periphery distinctly below mid-whorl. Suture weakly impressed, weakly undulating. Faint subsutural collar. Sculpture of moderately close-set, low, convex, opisthocline axial ribs not aligned axially (10: 13: 14). At least one indistinct, low varix per whorl. Spiral sculpture consisting of numerous delicate spiral threads overrunning axial ribs, most prominent in axial interspaces, fading over axial ribs. Last whorl attaining ~ 40 % of total height, axial ribs fading over convex base, no basal angulation, no peribasal cord, numerous spiral threads over base. Aperture ovate, columella straight, angled at transition to parietal area. Fasciole short, broad, with prominent growth lines. Outer lip not preserved. Columellar callus indistinct. No parietal callus. No umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF4FFF6FF3FFAB5FF609032.taxon	discussion	Discussion. This species is slightly reminiscent of Hemiacirsa oscari (De Boury in Cossmann, 1912). They are distinguished by the fewer, relatively sharper axial ribs in H. oscari. Moreover, the ribs in H. oscari are opisthocyrt whereas they are opisthocline in Acirsa koskeridouae sp. nov. In addition, the whorls of Hemiacirsa oscari are higher and more convex. Acirsa corrugata (Brocchi, 1814), from the Pliocene of Italy, is higher, has more prominent spiral cords and an angled base (see Pinna & Spezia 1978: pl. 59: fig. 3). Paleoenvironment. Shallow marine, inner neritic with sea grass (own data, M. H.).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF4FFF6FF3FFAB5FF609032.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienna Basin: Steinebrunn (Austria) (hoc opus).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF7FFF7FF3FF92DFCD19079.taxon	description	Figs 6 D	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF7FFF7FF3FF92DFCD19079.taxon	materials_examined	Type material. Holotype, M. 63.7792, SL: 14.8 mm, MD: 5.0 mm, Hidas, MÁSZ drill core (Hungary), illustrated in Csepreghy-Meznerics (1950: pl. 2, fig. 6), Figs 6 D. Revised description. Shell medium-sized, slender conical (apical angle ~ 17 °). Protoconch and early teleoconch unknown. Teleoconch of weakly convex whorls, periphery placed mid-whorl. Suture moderately impressed, linear, slightly emphasized by narrow, convex subsutural spiral cord. Sculpture strongly reduced, comprising few broad, indistinct spiral cords. No axial sculpture. Base of last preserved whorl with rounded angulation. Last whorl and aperture unknown.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF7FFF7FF3FF92DFCD19079.taxon	discussion	Discussion. This is a poorly known species, which was based on a single fragmentary and probably worn specimen. A second, more complete specimen was confused by Csepreghy (1950) with the freshwater dwelling Pachychilidae genus Tinnyea Hantken, 1887. Unfortunately, the specimen is lost (pers comm. Alfréd Dulai, 04.11.2024). Tinnyea inornata (Wenz, 1932) has a comparable spire with weak sculpture but has a very large aperture (see Wenz 1932, pl. 1, fig. 3). The extant Mediterranean-northeastern Atlantic Acirsa subdecussata (Cantraine, 1835) is most similar and differs only in its more prominent spiral cords. Given the poor preservation of the holotype, it is difficult to evaluate the value of this difference. Acirsa miopedemontana Sacco, 1891, from the Burdigalian of Baldissero (Italy), is stockier and differs in its more convex whorls and more prominent spiral sculpture (Sacco 1891: pl. 2, fig. 103). Paleoenvironment. Shallow marine, inner neritic.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF7FFF7FF3FF92DFCD19079.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Pannonian Basin: Hidas (Hungary). Genus Acrilloscala Sacco, 1890	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF7FFF7FF3FF92DFCD19079.taxon	type_taxon	Type species. Turbo geniculatus Brocchi, 1814; original designation by Sacco (1890: 321, footnote). Pliocene, Italy.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF7FFF7FF3FF92DFCD19079.taxon	diagnosis	Diagnosis. “ Medium thick shell. Medium size; shape narrow, turriculate; spire long, with regularly conical curve, polygyrate; protoconch smooth, acute, styliform; whorls convex, separated by deep, but continuous sutures; ornamentation composed of fine axial lamellae, not very prominent, slightly inclined, not deviated on the sutures, where they join the corresponding lamella of the previous whorl by a cochleariform inflexion; in their intervals, an excessively fine lattice of spiral striae and growth lines can be distinguished with a magnifying glass. Last whorl not very high, not reaching a quarter of the total height, subangular at the periphery of the base, which is covered by a thin, opaline disc, crossed by lamellae that then converge towards the imperforate center, where there is a very slight bulge with a rudimentary auricle. Circular aperture, with a slightly calloused, almost discontinuous peristome, the inner layer tapering completely to the parietal region; slightly oblique peristome, broadly bordered on the outside. ” (translated from Cossmann 1912: 65). “ Acrilloscala can be used for a group of Pliocene and Recent E. Atlantic species characterized by a poorly defined basal disc over which the axial ribs extend. ” (Bouchet & Warén 1986).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF7FFF7FF3FF92DFCD19079.taxon	discussion	Discussion. The genus is still represented by two species: Acrilloscala fusca (G. B. Sowerby II, 1844) from the tropical eastern Atlantic and the Cape Verde Archipelago (Gofas et al. 1985; Rolán 2005), and Acrilloscala lamyi (de Boury, 1909), from the northeastern Atlantic and the Cape Verde Islands (de Boury 1909 b; Bouchet & Warén 1986; Rolán 2005). Fossil records are also confined to the northeastern Atlantic, Proto-Mediterranean Sea and Central Paratethys Sea (Sacco 1891; Cossmann 1912; hoc opus).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF6FFF0FF3FF9E3FEA995D5.taxon	description	Figs 2 G, 9 A – D	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF6FFF0FF3FF9E3FEA995D5.taxon	materials_examined	Illustrated material. Holotype, NHMW 2024 / 0278 / 0001, SL: 5.6 mm, MD: 1.8 mm, Bad Vöslau (Austria), Figs 9 D 1 – D 2. Paratypes, NHMW 2010 / 0004 / 0526 a, SL: 4.5 mm, MD: 1.6 mm, Bad Vöslau (Austria), Figs 9 A 1 – A 2. NHMW 1997 z 0178 / 0969 a, SL: 5.7 mm, MD: 1.8 mm, Bad Vöslau (Austria), Figs 2 G, 9 B 1 – B 2. NHMW 2013 / 0078 / 0156 a, SL: 6.2 mm, MD: 2.1 mm, Baden (Austria), Figs 9 C 1 – C 2. Type locality. Bad Vöslau (Austria), Vienna Basin. Type stratum. Baden Formation. Age. Middle Miocene, middle Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF6FFF0FF3FF9E3FEA995D5.taxon	etymology	Etymology. Austriacus, referring to Austria.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF6FFF0FF3FF9E3FEA995D5.taxon	diagnosis	Diagnosis. Small, turreted, slender shell with faintly angled, convex whorls, weakly raised, slightly prosocline, moderately widely spaced lamellae and delicate spiral threads, marked basal angulation with broad peribasal cord and weakly concave basal disc.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF6FFF0FF3FF9E3FEA995D5.taxon	description	Description. Shell small, turreted, slender (apical angle ~ 16 – 18 °). Protoconch unknown. Teleoconch of at least nine convex whorls, with weakly concave subsutural ramp, faintly angled below, periphery placed just below mid-whorl. Suture deeply impressed, linear. Sculpture of delicate, thin, weakly raised, slightly prosocline, moderately widely spaced lamellae, only partly axially aligned (15: 16 / 18: 18 / 19). Tips of lamellae weakly alate, attached to preceding whorl. Crests of lamellae weakly rolled over. Delicate spiral threads in axial interspaces. Last whorl attaining about 35 % of total height, basal angulation with broad peribasal cord, basal disc weakly concave with distinct lamellae. Aperture ovate, columella weakly concave. Columellar callus forming narrow adherent rim. Parietal callus delicate. Peristome not preserved.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF6FFF0FF3FF9E3FEA995D5.taxon	discussion	Discussion. Acrilloscala propugnaculum sp. nov. differs from Acrilloscala austriaca sp. nov. in its angled whorls, wider-spaced and lower lamellae and weaker peribasal cord. Acrilloscala terebralis (Michelin, 1831) (= Acrilloscala degrangei De Boury in Cossmann, 1912), from the Serravallian of France, differs in its fewer but broader lamellae (see Ivolas & Peyrot 1900: pl. 2, figs 6 – 8; Cossmann 1912: pl. 3, figs 17 – 19; Cossmann & Peyrot 1922: pl. 4, figs 114 – 115). Scala (Acrilloscala) terebralis of Glibert (1949: pl. 11, fig. 1) is probably not conspecific with A. terebralis and is closer to the Paratethyan species in its more numerous and finer lamellae but is lower spired. Paleoenvironment. Occurrences in the Baden Formation of the Vienna Basin suggest middle to outer neritic settings in up to 250 m water depth (Kranner et al. 2021).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF6FFF0FF3FF9E3FEA995D5.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienna Basin: Baden, Bad Vöslau (Austria) (hoc opus).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF1FFF1FF3FFC4FFE7B9469.taxon	description	Figs 2 I, 8 E 1 – E 2	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF1FFF1FF3FFC4FFE7B9469.taxon	materials_examined	Type material. Holotype, NHMW 1865 / 0001 / 1096, SL: 13.4 mm, MD: 4.4 mm, Grund (Austria), Figs 2 I, 8 E 1 – E 2. Type locality. Grund (Austria), North Alpine-Carpathian Foreland Basin. Type stratum. Grund Formation. Age. Middle Miocene, early Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF1FFF1FF3FFC4FFE7B9469.taxon	etymology	Etymology. Fortior (Latin comparative singular of fortis, stronger), referring to the increasing strength of the varices.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF1FFF1FF3FFC4FFE7B9469.taxon	diagnosis	Diagnosis. Medium-sized, thin, moderately slender shell with strongly convex whorls, rounded angulation mid-whorl, bearing delicate, close-set lamellae on early teleoconch whorls, becoming wider spaced and more prominent on later whorls, convex base without peribasal cord.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF1FFF1FF3FFC4FFE7B9469.taxon	description	Description. Shell medium-sized, thin, moderately slender (apical angle ~ 20 °). Protoconch unknown. Teleoconch of about 11 whorls, with broad, weakly concave subsutural ramp and rounded angulation mid-whorl. Suture moderately impressed, linear. Sculpture on early teleoconch whorls of delicate, weakly raised, only weakly prosocline close-set lamellae, only partly aligned axially. Lamellae becoming wider spaced and more prominent on later whorls (16: 18: 21). Crests of lamellae only weakly rolled over on last whorl. Tips of lamellae faintly alate, attached to preceding whorl. Few more prominent varices on last whorl formed by amalgamation of several lamellae. Interspaces between lamellae with numerous delicate, barely visible spiral threads. Last whorl attaining 38 % of total height, lamellae persisting over base, strongly rolled over, no peribasal cord, base convex delimited by faint basal angulation. Aperture ovate, columella weakly convex. Very short, broad fasciole with distinct growth lines. Outer lip thin, basal lip thin, relatively narrow. Columellar callus forming narrow, adherent rim, parietal callus delicate. Peristome simple. No umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF1FFF1FF3FFC4FFE7B9469.taxon	discussion	Discussion. Acrilloscala terebralis (Michelin, 1831) (= Acrilloscala degrangei De Boury in Cossmann, 1912), from the Serravallian of France, is much slenderer and has higher whorls (see Ivolas & Peyrot 1900: pl. 2, figs 6 – 8; Cossmann 1912: pl. 3, figs 17 – 19; Cossmann & Peyrot 1922: pl. 4, figs 114 – 115). Acrilloscala sp. 1, from the Badenian of the Vienna Basin, is also much slenderer and has more prominent lamellae on early teleoconch whorls. Similarly, Acrilloscala geniculata (Brocchi, 1814), from the Pliocene of the Mediterranean Sea, differs in its slenderer shape and has distinct, relatively widely spaced lamellae on early whorls (see Bouchet & Warén 1986: figs 1222 – 1223). Subuliscala subvaricosa (Cantraine, 1842) as illustrated by Cossmann (1912: pl. 2, figs 16 – 19) is reminiscent of Acrilloscala fortior sp. nov. in general shape but differs in its more deeply incised suture and the strongly thickened peristome. Paleoenvironment. At the locality Grund channels formed in middle to outer neritic environments and were filled, with allochthonous assemblages uniting coastal-mudflat faunas with inner neritic ones (Zuschin et al. 2005; Roetzel 2009). The preservation suggests that the species was part of assemblages from middle to outer neritic environments.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF1FFF1FF3FFC4FFE7B9469.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): North Alpine - Carpathian Foreland Basin: Grund (Austria (hoc opus).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF3FFFCFF3FFF2EFD3B97DD.taxon	description	Figs 2 J, 8 F, 9 A – C	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF3FFFCFF3FFF2EFD3B97DD.taxon	materials_examined	Type material. Holotype, NHMW 1864 / 0001 / 0616, SL: 7.7 mm, MD: 2.4 mm, Grund (Austria), Figs 9 B 1 – B 2. Paratypes, NHMW 2024 / 0279 / 0001, SL: 12.1 mm, MD: 4.6 mm, Grund (Austria), Figs 8 F 1 – F 2. NHMW 2007 / 0086 / 0165 b, SL: 9.3 mm, MD: 3.2 mm, Baden (Austria), Figs 2 J, 9 A 1 – A 2. NHMW 2007 / 0086 / 0165 a, SL: 9.9 mm, MD: 3.3 mm, Baden (Austria), Figs 9 C 1 – C 2. Type locality. Grund (Austria), North Alpine-Carpathian Foreland Basin. Type stratum. Grund Formation. Age. Middle Miocene, early Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF3FFFCFF3FFF2EFD3B97DD.taxon	etymology	Etymology. Propugnaculum (Latin for a rampart; figurative for a defense), noun in apposition.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF3FFFCFF3FFF2EFD3B97DD.taxon	diagnosis	Diagnosis. Medium-sized, turreted, slender shell with moderately impressed suture, multispiral protoconch, weakly angled teleoconch whorls and widely spaced, relatively broad, low, prosocline lamellae and delicate microsculpture of spiral cords in axial interspaces.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF3FFFCFF3FFF2EFD3B97DD.taxon	description	Description. Medium-sized small, turreted, slender (apical angle ~ 15 °). Protoconch turreted of four smooth, weakly convex whorls (fide Bałuk 1975). Teleoconch whorls high, moderately convex, with steep, weakly concave subsutural ramp, angled at shoulder, convex below, periphery placed just below mid-whorl. Suture moderately impressed, linear. Sculpture of delicate, moderately broad, weakly raised, widely spaced, slightly prosocline lamellae, vaguely aligned axially (10: 11 / 12: 12). Tips of lamellae weakly alate, attached to preceding whorl. Faint spiral threads in axial interspaces only visible under strong magnification. Last whorl attaining ~ 35 % of total height, bearing few more prominent varices formed by amalgamation of lamellae, weak basal angulation without clear peribasal cord, lamellae persisting over weakly convex base, strongly rolled over. Fasciole very short, broad, with prominent growth lines. Aperture elongate ovate, columella weakly convex. Outer lip not thickened. Columellar callus forming narrow, attached rim. Parietal callus delicate. Peristome simple. No umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF3FFFCFF3FFF2EFD3B97DD.taxon	discussion	Discussion. Acrilloscala subuloprisca (Sacco, 1891), from the Tortonian of the Proto-Mediterranean Sea, has strongly angled whorls that are more constricted below the periphery. In addition, A. subuloprisca has narrower lamellae (see Ferrero Mortara et al. 1984: pl. 6, fig. 8 b). Acrilloscala terebralis (Michelin, 1831) (= Acrilloscala degrangei De Boury in Cossmann, 1912), from the Serravallian of France, is slenderer and has slightly less angled and higher whorls (see Ivolas & Peyrot 1900: pl. 2, figs 6 – 8; Cossmann 1912: pl. 3, figs 17 – 19; Cossmann & Peyrot 1922: pl. 4, figs 114 – 115). The extant Acrilloscala lamyi (de Boury, 1909) differs in its greater number of lamellae and much weaker shoulder angulation (Bouchet & Warén 1986: figs 1219 – 1221) but is otherwise similar to Acrilloscala propugnaculum sp. nov. in shape and the delicate spiral sculpture. The Pliocene type species Acrilloscala geniculata (Brocchi, 1814) is much larger (SL up to 28.9 mm, own data B. L.), higher, slenderer and has evenly convex whorls (see Bouchet & Warén 1986: figs 1222 – 1223). Paleoenvironment. At the locality Grund channels formed in middle to outer neritic environments and were filled with allochthonous assemblages uniting coastal-mudflat faunas with inner neritic ones (Zuschin et al. 2005; Roetzel 2009). The occurence in the Baden Formation suggests middle to outer neritic environments.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF3FFFCFF3FFF2EFD3B97DD.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Korytnica Basin: Korytnica (Poland) (Bałuk 1975); North-Alpine Carpathian Foreland Basin: Grund (Austria (hoc opus); Vienna Basin: Baden (Austria) (hoc opus); Pannonian Basin: Várpalota (Hungary) (Kecskeméti-Körmendy 1962); Făget Basin: CoŞteiu de Sus (Romania) (Cossmann 1912). Genus Adiscoacrilla Sacco, 1890	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF3FFFCFF3FFF2EFD3B97DD.taxon	type_taxon	Type species. Acrilla coppii de Boury, 1890, original designation by Sacco (1890: 321). Pliocene, Italy.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF3FFFCFF3FFF2EFD3B97DD.taxon	diagnosis	Diagnosis. Small, moderately slender to slightly stocky conical shells with evenly convex to faintly shouldered whorls, deeply incised suture, moderately close-set to moderately wide-spaced, thin, moderately raised lamellae, not aligned axially, crests weakly rolled over or straight, adapical tips often slightly alate. Interspaces with delicate spiral cords or threads. More or less distinct basal angulation, often with weak peribasal cord and flattened basal disc bearing distinct, low lamellae or ribs. Aperture ovate with thin peristome.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF3FFFCFF3FFF2EFD3B97DD.taxon	discussion	Discussion. The original diagnosis by Sacco (1890: 321) is very brief and misguiding because Sacco emphasized the absence of a “ real basal disc ”, which is obviously incorrect. This statement, however, led Cossmann (1912: 58) to reject Adiscoacrilla and to synonymize it with Acrilla. Consequently, Adiscoacrilla herthae (Boettger, 1906) was placed by Zilch (1934) and subsequent authors in Acrilla H. Adams, 1860 [type species Scalaria acuminata G. B. Sowerby II, 1844; present-day, Strait of Malacca]. However, Acrilla is very slender with delicate thread-like lamellae (Weil et al. 1999: 206) and is not very similar to the species placed herein in Adiscoacrilla. The genus comprises Adiscoacrilla anticoppii sp. nov., A. herthae, A. laticostata (Bałuk, 1975) and A. walaszczyki sp. nov. from the Langhian and Serravallian of the Central Paratethys Sea, A. silentiosa (Cossmann & Peyrot, 1922), from the Serravallian of the northeastern Atlantic, A. libassii (Seguenza, 1876) and A. interposita (Sacco, 1891), from the Tortonian of the Po Basin, and A. coppii (de Boury, 1890) [= A. lacunocostata (Sacco, 1891), A. depressocoppi (Sacco, 1891)], A. opaliaeformis (Sacco, 1891), A. stefanii (De Boury, 1890) and A. villalvernensis (Sacco, 1891) from the Pliocene of the Mediterranean Sea.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFFDFFFCFF3FFE47FEB890CD.taxon	description	Figs 7 J, 9 D 1 – D 2	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFFDFFFCFF3FFE47FEB890CD.taxon	materials_examined	Type material. Holotype, NHMW 2024 / 0279 / 0004, SL: 18.3 mm, MD: 7.3 mm, Grund (Austria), Figs 7 J, 9 D 1 – D 2. Type locality. Grund (Austria), North Alpine-Carpathian Foreland Basin. Type stratum. Grund Formation. Age. Middle Miocene, early Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFFDFFFCFF3FFE47FEB890CD.taxon	etymology	Etymology. Anti (Ancient Greek for ‘ against’) and coppii, referring to Adiscoacrilla coppii (de Boury, 1890).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFFDFFFCFF3FFE47FEB890CD.taxon	diagnosis	Diagnosis. Medium-sized, moderately slender, turreted shell with strongly convex whorls, delicate, widely spaced, weakly raised lamellae with prominent spiral threads in interspaces, weak basal angulation; lamellae distinctly increase in strength during ontogeny.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFFDFFFCFF3FFE47FEB890CD.taxon	description	Description. Shell medium-sized, moderately slender, turreted (apical angle ~ 22 °). Protoconch and early teleoconch whorls unknown. Teleoconch of strongly convex whorls with indistinct subsutural ramp, periphery placed just below mid-whorl. Suture deeply impressed, linear. Sculpture of numerous, delicate, widely spaced, weakly raised, weakly prosocline lamellae, not aligned axially (?: ~ 20: ~ 25), separated by much wider interspaces. Some lamellae slightly more prominent, occasionally forming narrow varices. Lamellae distinctly increase in strength during ontogeny. Crests of lamellae straight. Interspaces with numerous prominent spiral threads, occasionally with intercalated secondary spirals. Last whorl convex, attaining about 40 % of total height, basal angulation weak without peribasal cord, base weakly convex with delicate spiral threads, lamellae slightly rolled over. Short, broad fasciole with prominent growth lines. Aperture elongate ovate, columella weakly concave. Columellar callus forming narrow rim slightly detached from fasciole, no parietal callus. Umbilicus reduced to narrow chink. Peristome not preserved.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFFDFFFCFF3FFE47FEB890CD.taxon	discussion	Discussion. This species is reminiscent of Adiscoacrilla herthae (Boettger, 1906) but is larger, has more numerous lamellae and lacks a peribasal cord. The whorls are regularly convex, whereas in A. herthae they are distinctly shouldered. Adiscoacrilla coppii (de Boury, 1890), from the Pliocene of Italy, is similar in general shape but differs in its wider apicalangle, more elevated lamellae and marked basal angulation (see de Boury 1890: pl. 4, fig. 1; Landau et al. 2006: pl. 13, figs 5 – 6; Chirli 2009: pl. 18, figs 8 – 15). Paleoenvironment. At the locality Grund channels formed in middle to outer neritic environments and were filled with allochthonous assemblages uniting coastal-mudflat faunas with inner neritic ones (Zuschin et al. 2005; Roetzel 2009). The preservation suggests that the shell might have belonged to the middle and outer neritic assemblages.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFFDFFFCFF3FFE47FEB890CD.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): North Alpine-Carpathian Foredeep: Grund (Austria).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFFDFFFDFF3FF8B7FD6F9333.taxon	description	Figs 7 K, 9 E	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFFDFFFDFF3FF8B7FD6F9333.taxon	materials_examined	Type material. Holotype, SMF 380169 _ 12 - 1454 (XII 1454 a), SL: 5.9 mm, MD: 2.1 mm, CoŞteiu de Sus (Romania), illustrated in Zilch (1934: pl. 10, fig. 78), Figs 7 K, 9 E. Revised description. Shell small, moderately slender turreted (apical angle 22 °). Protoconch and early teleoconch whorls unknown. Teleoconch whorls convex with weakly concave subsutural ramp, angled at shoulder placed just above mid-whorl, convex below. Suture deeply impressed. Sculpture of thin, slightly prosocline, moderately to strongly raised, moderately close-set lamellae, not aligned axially (?: ~ 14: ~ 17); adapical tips of lamellae weakly alate, crests of varices weakly rolled over. Interspaces with delicate spiral threads. Last whorl attaining about 40 % of total height, with marked basal angulation delimited by prominent, slightly scalloped peribasal cord. Basal disc wide, flattened with prominent axial ribs, no spiral sculpture. Aperture ovate, columella weakly concave. Outer lip not preserved. Columellar callus forming thin, indistinct adherent rim.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFFDFFFDFF3FF8B7FD6F9333.taxon	discussion	Discussion. The holotype has only moderately raised lamellae, whereas a specimen from the Vienna Basin, illustrated by Švagrovský (1982), has higher lamellae, which are faintly pointed close to the adapical suture. Nevertheless, we treat both fragmentary specimens provisionally as conspecific. Adiscoacrilla coppii (de Boury, 1890), from the Pliocene of Siena (Italy), is exceptionally similar and differs mainly in its much larger size (SL: 20 mm) (de Boury 1890: pl. 4, fig. 1; Landau et al. 2006: 42). Other specimens of Adiscoacrilla coppii, from the Pliocene of the Toscana, are stockier with a wider last whorl (see Chirli 2009: pl. 18, figs 8 – 15). Adiscoacrilla lacunocostata (Sacco, 1891) and A. depressocoppi (Sacco, 1891), from the Pliocene of Asti, are reminiscent of the Paratethyan species in their shell shape but differ in having fewer lamellae (see Sacco 1891: pl. 2, figs 66, 67). Adiscoacrilla opaliaeformis (Sacco, 1891) differs in its more rounded basal angulation and narrower basal disc (see Sacco 1891: pl. 2, fig. 69). Adiscoacrilla villalvernensis (Sacco, 1891), from the Pliocene of the Mediterranean Sea, differs from A. herthae in its much stockier shape (see Landau et al. 2006: pl. 13, figs 9 – 10, pl. 14, fig 1). Adiscoacrilla silentiosa (Cossmann & Peyrot, 1922), from the Serravallian of France, is also stockier and has weaker lamellae (see Cossmann & Peyrot 1922: pl. 4, figs 64 – 65, 130 – 131). Adiscoacrilla herthae is superficially reminiscent of the much larger Parviscala paratethyca sp. nov. in sculpture but the distinct basal disc excludes placement in Parviscala de Boury, 1887. Paleoenvironment. Shallow marine, inner neritic.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFFDFFFDFF3FF8B7FD6F9333.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienna Basin: Borský Mikuláš (Slovakia) (Švagrovský 1982); Pannonian Basin: Herend (Hungary) (Kókay 1966),? Szokolya (Hungary) (Báldi 1960); Făget Basin: CoŞteiu de Sus (Romania) (Boettger, 1906).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFFCFFFEFF3FFA2DFF5C931D.taxon	description	Figs 7 G, 10 B 1 – B 2	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFFCFFFEFF3FFA2DFF5C931D.taxon	materials_examined	Type material. Holotype, Z. PAL. U. W., BkK-G 35, SL: 5.0 mm, MD: 2.6 mm, Korytnica (Poland), illustrated in Bałuk (1975: pl. 21, fig. 8), Figs 7 G, 10 B 1 – B 2 Revised description. Small, thin-shelled, moderately broad, conical (apical angle 33 – 35 °), gradate, coronate. Protoconch multispiral conical, of about four smooth, slightly convex whorls [fide Bałuk 1975: 177]. Teleoconch of six relatively low, convex whorls with narrow, weakly concave subsutural shelf, marked shoulder angulation, subcylindrical below, with periphery placed mid-whorl. Suture deeply impressed. Sculpture of close-set, moderately raised, prosocline lamellae with short, trigonal spine at shoulder (14 / 16: ~ 24: ~ 33). Crests of lamellae weakly rolled over on later whorls. Interspaces with numerous, delicate spiral threads. Last whorl attaining about 50 % of total height, with weak basal angulation bordering flattish base. Lamellae strongly rolled over on base. Aperture ovate, columella weakly concave. Columellar callus forming narrow rim, detached from base; parietal callus narrow. Peristome slightly thickened. Umbilicus reduced to narrow chink.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFFCFFFEFF3FFA2DFF5C931D.taxon	discussion	Discussion. Adiscoacrilla villalvernensis (Sacco, 1891), from the Pliocene of the Mediterranean Sea, differs in its wider spaced lamellae and weaker shoulder angulation (see Landau et al. 2006: pl. 13, figs 9 – 10, pl. 14, fig 1). Superficially similar, Anguloacrilla kimakowiczi (Boettger, 1896) differs at the same growth stage in its more convex whorls and convex axial ribs. Anguloacrilla karamanensis sp. nov. lacks the distinct subsutural shelf at same growth stage. As well as these differences, Anguloacrilla nov. gen. species have a more strongly developed basal disc. Paleoenvironment. Unknown.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFFCFFFEFF3FFA2DFF5C931D.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Korytnica Basin: Korytnica (Poland) (Bałuk 1975).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFFFFFFFFF3FFA07FD429549.taxon	description	Figs 4 F, 7 L, 9 F 1 – F 2	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFFFFFFFFF3FFA07FD429549.taxon	materials_examined	Type material. Holotype, NHMW 2024 / 0278 / 0002, SL: 3.9 mm, MD: 1.9 mm, Bad Vöslau (Austria), Figs 4 F, 7 L, 9 F 1 – F 2. Type locality. Bad Vöslau (Austria), Vienna Basin. Type stratum. Baden Formation. Age. Middle Miocene, middle Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFFFFFFFFF3FFA07FD429549.taxon	etymology	Etymology. In honor of Ireneusz Walaszczyk (Faculty of Geology, University of Warsaw, Poland) for his help searching for type material published by Wacław Bałuk.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFFFFFFFFF3FFA07FD429549.taxon	diagnosis	Diagnosis. Very small, moderately broad conical, turreted shell with strongly convex whorls, weak shoulder placed high, bearing close-set, erect lamellae, prominent spiral threads in interspaces, strong and sharp basal angulation.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFFFFFFFFF3FFA07FD429549.taxon	description	Description. Very small, thin-shelled, moderately broad, conical (apical angle 35 °). Protoconch conical, of three smooth, slightly convex whorls. Teleoconch of at least six strongly convex whorls with narrow, shallow, weakly convex subsutural ramp, delimited by high-placed faint shoulder angulation, with periphery placed just above mid-whorl. Suture deeply impressed, linear. Sculpture of moderately raised, closely set, weakly prosocline, erect lamellae (18: 22: 24), not aligned axially. Crests of lamellae only weakly rolled. Interspaces with numerous, delicate spiral threads. Last preserved whorl with sharp basal angulation delimited by narrow peribasal cord. Basal disk flat with prominent axial ribs, corresponding to lamellae. No umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFFFFFFFFF3FFA07FD429549.taxon	discussion	Discussion. Adiscoacrilla walaszczyki sp. nov. differs from the otherwise similar Adiscoacrilla laticostata (Bałuk, 1975) in its protoconch of only three instead of four whorls, the more convex whorls and it lacks a coronate shoulder. Adiscoacrilla herthae (Boettger, 1906) is slenderer and has fewer lamellae. Adiscoacrilla villalvernensis (Sacco, 1891), from the Pliocene of the Mediterranean Sea, differs from A. walaszczyki in its wider spaced lamellae (see Landau et al. 2006: pl. 13, figs 9 – 10, pl. 14, fig 1). Adiscoacrilla coppii (de Boury, 1890), from the Mediterraean Pliocene, is stockier with a wider last whorl (see Chirli 2009: pl. 18, figs 8 – 15). This species was confused by Bałuk (1975) also with Adiscoacrilla? orientalis (Friedberg, 1928), which differs in its much more prominent and wider spaced lamellae. Paleoenvironment. Occurrences in the Baden Formation of the Vienna Basin suggest middle to outer neritic settings in up to 250 m water depth (Kranner et al. 2021).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFFFFFFFFF3FFA07FD429549.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Korytnica Basin: Korytnica (Poland) (Bałuk 1975); Vienna Basin: Bad Vöslau (Austria) (hoc opus).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFFEFFFFFF3FFC33FC3993E2.taxon	discussion	Discussion. Kókay (1966) illustrated two spire fragments of a small, slender species with a high, turreted protoconch of four whorls, and high, angled whorls with close-set, prominent lamellae (30), weak spiral threads and distinct basal angulation (SL: 4 mm). This species was confused by Kókay (1966) with Subuliscala lagusensis (de Boury in Cossmann, 1912), from the Burdigalian of France, which differs in its narrower and more numerous lamellae, and more rounded whorls (see Cossmann 1912: pl. 2, figs 14 – 15; Cossmann & Peyrot 1922: pl. 4, figs 49 – 50). We tentatively place this species in Adiscoacrilla. Adiscoacrilla herthae (Boettger, 1906) is superficially similar but broader and has fewer lamellae. We do not describe this species formally due to the poorly preserved material. Paleoenvironment. Shallow marine.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFFEFFFFFF3FFC33FC3993E2.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Pannonian Basin: Herend (Hungary) (Kókay 1966); Făget Basin: CoŞteiu de Sus (Romania) (hoc opus).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFFEFFF8FF3FFA5AFE469395.taxon	description	Fig. 10 A	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFFEFFF8FF3FFA5AFE469395.taxon	materials_examined	Type material. Lectotype (designated herein), specimen illustrated in Friedberg (1928: pl. 38, fig. 24), SL: 6 mm [fide Friedberg 1928: 606; this differs from the scale on plate 38, which suggests a size of about 3 mm], Szuszkowce (Ukraine), Fig. 10 A (re-illustrated from Friedberg 1928). We are not aware of the whereabouts of the material. Friedberg (1928) illustrated two specimens, which are almost certainly not conspecific. The fragmentary specimen illustrated on pl. 38, fig. 23 differs from the lectotype in its slenderer shape, prosocline lamellae and represents another Epitoniidae species. Revised description. Small, moderately broad conical (apical angle 27 °). Protoconch and early teleoconch whorls unknown. Teleoconch whorls evenly convex with periphery placed mid-whorl. Suture moderately impressed. Sculpture of prominent, moderately raised, slightly prosocline lamellae, not aligned axially, separated by wider interspaces (?: ~ 16: ~ 16). At least one varix per whorl. Interspaces with delicate spiral threads. Last whorl attaining about 40 % of total height with weakly angled base delimited by weak peribasal cord. Lamellae weakening over base. Fasciole short, broad, with prominent growth lines. Aperture subcircular, columella nearly straight. Columellar callus forming narrow rim. Parietal callus weak, peristome thickened, duplex, inner layer continuous, narrow. No umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFFEFFF8FF3FFA5AFE469395.taxon	discussion	Discussion. This species is only known from a single, fragmentary specimen. Its placement in Adiscoacrilla is tentative. The weakly angled base excludes placement in the otherwise comparable Sodaliscala de Boury, 1909 [type species Sodaliscala multistriata (Say, 1826); present-day, western Atlantic]. The shell seems to be composed of an outer and inner layer; the inner one whitish. The two layers are most obvious at the labial varix, where they form a ‘ double rim’. We are not sure what the implication of this is, but it is also seen in other epitoniids, but not in other members of the genus Adiscoacrilla. Paleoenvironment. Probably shallow marine.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFFEFFF8FF3FFA5AFE469395.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Polish-Ukrainian Fore-Carpathian Basin: Shushkivtsi (= Szuszkowce) (Ukraine) (Friedberg 1938). Genus Amaea H. Adams & A. Adams, 1853	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFFEFFF8FF3FFA5AFE469395.taxon	type_taxon	Type species. Scalaria magnifica G. B. Sowerby II, 1844; subsequent designation by de Boury (1909 a: 258). Present-day, Japan.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFFEFFF8FF3FFA5AFE469395.taxon	diagnosis	Diagnosis. “ Thin shell, quite large, turriculate, with convex and cancellated whorls; the base of the last whorl is convex and bears a rudimentary disc; but the main characteristic of this acrilliform genus is its oval opening, holostome in front, narrowed behind, and particularly the columellar bulge that is not seen in any other Scalidae; the peristome is weak, except when it coincides with one of the varices. ” (translated from Cossmann 1912: 25).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFFEFFF8FF3FFA5AFE469395.taxon	discussion	Discussion. Some of the discussed species have been placed previously in Scalina Conrad, 1865 by us (Harzhauser et al. 2014). However, Scalina Conrad, 1865 (type species Scala staminea Conrad, 1860, Eocene, Alabama, USA) is currently treated as subjective junior synonym of Amaea by MolluscaBase eds. (2014 d), which is followed herein. DeVries (2007: 231) distinguished Scalina from Amaea based on “ their contrasting basal and lateral sculpture ”. DeVries (2007) described several Scalina species from the Early Miocene to Pliocene of the western Atlantic and eastern Pacific that are morphologically clearly closely related to the Proto-Mediterranean- Paratethyan species discussed herein. Nevertheless, several extant Amaea species, such as Amaea gazeoides Kuroda & Habe in Habe 1961 and Amaea splendida (de Boury, 1913 a) display nearly identical sculpture and support the synonymization of both genera.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF9FFF9FF3FF990FB4593A8.taxon	description	Figs 2 K, 11 A – B	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF9FFF9FF3FF990FB4593A8.taxon	description	A [maea]. (A [crilla].) amoena subreticulata (Orb.) — Sieber 1958: 129 [non Amaea subreticulata (d’Orbigny, 1852)].	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF9FFF9FF3FF990FB4593A8.taxon	materials_examined	Type material. Holotype, NHMW 1858 / 0015 / 0196, SL: 16.5 mm, MD: 6.0 mm, Baden (Austria), illustrated in Hörnes (1856: pl. 46, figs 11 a – b), Figs 2 K, 11 A 1 – A 2. Illustrated material. NHMW 1867 / 0019 / 0256, SL: 13.7 mm, MD: 6.3 mm, CoŞteiu de Sus (Romania), Figs 11 B 1 – B 2. Revised description. Shell medium-sized, moderately slender (apical angle ~ 22 °). Protoconch unknown. Teleoconch of more than nine relatively low whorls. Whorl profile strongly convex. Early teleoconch whorls with straight-sided to weakly concave subsutural ramp, weakly angled periphery placed mid-whorl; later whorls evenly convex. Suture moderately impressed, linear. Sculpture of delicate, slightly raised orthocline to weakly prosocline lamellae, not aligned axially (~ 20: ~ 40: ~ 45). Lamellae slightly alate and bent in adapertural direction. Spiral sculpture on early whorls of two weak spiral cords on subsutural ramp and three primary spirals along periphery. Secondary spiral cords intercalated on later whorls. Spiral sculpture of later whorls only slightly less raised than lamellae, forming fine, regularly cancellate sculpture. Faint spiral threads and delicate growth lines in interspaces. Last whorl attaining about 37 % of total height, with about seven to nine spiral cords, secondary cords typically weaker but may attain equal strength to primaries, basal angulation delimited by moderately prominent, narrow peribasal cord, delicate lamellae and spiral threads on basal disc. Aperture ovate with parietal concavity. Outer lip and peristome not preserved. Columellar callus and parietal callus forming broad, relatively thick rim, slightly detached from base. Umbilicus reduced to very narrow chink.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF9FFF9FF3FF990FB4593A8.taxon	discussion	Discussion. Scalaria (Acrilla) miobronni Sacco, 1891, from the Tortonian of S. Agata (Italy), corresponds in all features with the Paratethyan shells. Therefore, we treat it as subjective junior synonym of Amaea hoernesi (De Gregorio, 1889). A specimen from the Serravallian of Saubrigues (France, illustrated by Cossmann (1912: pl. 3, figs 10 – 11) has a weak shoulder angulation and slightly wider cancellate sculpture and is excluded herein from synonymy. Amaea subreticulata (d’Orbigny, 1852) from the Early Miocene of the North Sea, as illustrated by Glibert (1952: pl. 7, fig. 1), has much weaker lamellae that are not elevated over the spiral cords. Amaea hoernesi was confused by Hörnes (1856) with Amaea amoena (Philippi, 1844), from the Oligocene of the North Sea, which differs in its less incised suture and less convex whorls. The Oligocene species has broad spiral cords, which are separated by narrow interspaces (Janssen 1978: pl. 13, fig. 65). Amaea bronni (Seguenza, 1876), from the Mediterranean Pliocene, differs in its higher lamellae and spiral cords and the weakly coronate shoulder (see Landau et al. 2006: pl. 14, figs 2, 3; Chirli 2009: pl. 17, figs 12 – 17, pl. 18, fig. 1). For a separation from Amaea grunerti sp. nov. see below. Paleoenvironment. Unknown.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF9FFF9FF3FF990FB4593A8.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Korytnica Basin: Korytnica (Poland) (Bałuk 2006); Vienna Basin: Baden (Austria) (Hörnes 1856); Făget Basin: CoŞteiu de Sus (Romania) (Boettger 1902). Proto-Mediterranean Sea. Tortonian (Late Miocene): Po Basin: S. Agata (Italy) (Sacco 1891).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF8FFC5FF3FF993FA329689.taxon	description	Figs 2 L, 11 C – F	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF8FFC5FF3FF993FA329689.taxon	materials_examined	Type material. Holotype, GBA 1875 / 01 / 37, SL: 53.4 mm, MD: 16.3 mm, Ottnang (Austria), illustrated in Hoernes (1875: 362, pl. 10, fig. 9), Figs 2 L, 11 D 1 – D 2. Paratypes, NHMW 2014 / 0379 / 00334, SL: 34 mm, MD: 12.5 mm, Allerding (Austria), illustrated in Harzhauser et al. (2014: pl. 3, fig. 3), Fig 11 C. GBA 1875 / 01 / 37, SL: 33.6 mm, MD: 14.9 mm, Ottnang (Austria), illustrated in Hoernes (1875: 362, pl. 10, fig. 8), Figs 11 E 1 – E 2. NHMW 2014 / 0379 / 00333, SL: 18.8 mm, MD: 8.5 mm, Allerding (Austria), illustrated in Harzhauser et al. (2014: pl. 3, fig. 2), Fig 11 F. Type locality. Ottnang (Austria), North Alpine Foreland Basin. Type stratum. Ottnang Formation. Age. Early Miocene, early Ottnangian (mid-Burdigalian).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF8FFC5FF3FF993FA329689.taxon	etymology	Etymology. In honor of Patrick Grunert (University Köln, Germany) in recognition of his contributions on the stratigraphy of the North-Alpine Foreland Basin.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF8FFC5FF3FF993FA329689.taxon	diagnosis	Diagnosis. Very large, moderately slender shell with convex whorls, faint shoulder angulation and prominent cancellate to axially elongate rectangular sculpture of thin, low lamellae crossing up to six slightly broader primary cords plus one primary cord on subsutural ramp.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF8FFC5FF3FF993FA329689.taxon	description	Description. Shell very large, moderately slender (apical angle ~ 22 °). Protoconch unknown. Teleoconch of more than ten relatively low whorls. Whorls strongly convex with steep, straight sided to weakly convex subsutural ramp, faint shoulder angulation and convex periphery placed just above mid-whorl. Suture deeply impressed, linear. Sculpture of delicate, weakly raised, prosocline lamellae, not aligned axially (~ 16: ~ 45: ~ 55). Broader, band-like, flat varices may develop by amalgamation of several lamellae. Spiral sculpture on early whorls of one spiral cord on subsutural ramp and four to five primary spiral cords below. Sixth spiral cord intercalated at or above abapical suture on later whorls. Spiral sculpture equal in strength and elevation to lamellae, forming sharp, regularly cancellate to weakly axially elongate rectangular sculpture. Secondary spiral cords may be intercalated in some specimens. Last whorl attaining about 35 % of total height, basal angulation rounded to indistinctly angled delimited by narrow peribasal cord of same strength as primary spirals, base weakly convex with thin lamellae and numerous prominent spiral cords. Fasciole very short, broad with prominent growth lines. Aperture ovate. Columella weakly concave. Outer lip thin. Basal lip slightly excavated. Columellar callus with moderate swelling, forming broad, relatively thick rim, attached to base. No umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF8FFC5FF3FF993FA329689.taxon	discussion	Discussion. Already Pantanelli (1886) recognized that the Miocene shells described by Hoernes (1875) were not conspecific with Amaea amoena (Philippi, 1844) and discussed the specimens illustrated by Hoernes (1875) from Ottnang in his description of Scalaria (Cirsotrema) hörnesi from the Langhian of Pantano of Reggio Emilia in Italy. He tentatively suggested that both records were conspecific but emphasized, that this conclusion was based solely on the illustration in Hoernes (1875) and not direct comparison. In our opinion the two records are not conspecific and therefore, the name Amaea hoernesi (Pantanelli, 1886) refers to the species occurring in the Middle Miocene (Langhian) of Italy and cannot be used for the Ottnangian species, as done by Sieber (1956) and Steininger (1973). Sacco (1891: 53) proposed Scalaria (Cirsotrema) mioeuropaeum as name for Scalaria (Cirsotrema) hoernesi Pantanelli, 1886 because he considered it to be a homonym of Scalaria pumicea hornesi De Gregorio, 1889. Both names, however, are available because Article 57.6 of the ICZN (1999) states: “ a one-letter difference between species-group names combined with the same generic name is sufficient to prevent homonymy ”. According to article 32.5 of the ICZN special characters like ä, ö and ü used in species names before 1985 must be corrected to ae, oe and ue. Therefore, Scalaria hornesi De Gregorio, 1889 and S calaria (Cirsotrema) hoernesi Pantanelli, 1886) are both available. Amaea grunerti sp. nov. differs from the Middle Miocene A. hoernesi in its larger size, its less angled basal angulation and its convex base. Amaea hoernesi has three primary spiral cords with secondary cords intercalated, whereas Amaea grunerti has five to six primary spiral cords and usually lacks prominent secondary cords. In addition, A. hoernesi has evenly convex whorls and lamellae, which are raised above the spiral cords, whereas A. grunerti has a faint shoulder angulation and spiral and axial sculpture are almost equal in strength. A specimen from the Ottnangian of Hungary, illustrated by Kovács et al. (2023) as Scalina subreticulata (d’Orbigny, 1852) differs slightly from the Austrian specimens in its broader, convex primary spiral cords and the presence of prominent secondary spiral cords but otherwise agrees in the pattern of primary cords. Therefore, we treat both occurrences tentatively as conspecific.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF8FFC5FF3FF993FA329689.taxon	distribution	Distribution in Central Paratethys. Ottnangian (Early Miocene): North Alpine Foreland Basin: Allerding, Ottnang (Austria) (Hoernes 1875; Harzhauser et al. 2014); Early Miocene North Hungarian Basin: Parádfürdö (Hungary) (Kovács et al. 2023). Genus Anguloacrilla nov. gen.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF8FFC5FF3FF993FA329689.taxon	type_taxon	Type species. Scalaria (Acrilla) kimakowiczi Boettger, 1896. Middle Miocene, Central Paratethys Sea (Romania).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF8FFC5FF3FF993FA329689.taxon	etymology	Etymology. Coinage of angulus (Latin for angle) and Acrilla, referring to the angled shoulder.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF8FFC5FF3FF993FA329689.taxon	diagnosis	Diagnosis. Medium-sized, moderately slender, finely coronate shells with deeply impressed suture, relatively low, convex whorls with narrow, shallow subsutural ramp, numerous close-set lamellae, often forming low spines at shoulder, with numerous spiral threads in interspaces, spirals override the lamellae giving them irregular or granular appearance, marked basal angulation with flat basal disc and prominent peribasal cord; abapical part of the outer lip ends in short spout.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF8FFC5FF3FF993FA329689.taxon	description	Description. Medium-sized, moderately slender, finely coronate shells. Protoconch unknown from type species. Teleoconch whorls relatively low, moderately convex with narrow, shallow subsutural ramp, more or less prominent shoulder angulation, typically reinforced by weakly protruding spines. Periphery placed mid-whorl. Suture deeply impressed. Sculpture of numerous weakly prosocline lamellae and more or less prominent spiral threads in interspaces. Spirals override axials causing delicate granulation on laminar crests. Last whorl attaining about 35 % of total height with very prominent basal angulation, coinciding with prominent peribasal cord. Basal disc flattened with weak laminae and spiral threads. Aperture semicircular, thin. Peristome unknown. Abapical part of the outer lip ends in short spout. Columellar callus forming broad rim, adherent but well delimited from base. No umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFF8FFC5FF3FF993FA329689.taxon	discussion	Discussion. Since Boettger (1896), the species placed herein in Anguloacrilla nov. gen. were treated as Acrilla or Amaea (Acrilla) (e. g., Kojumdgieva in Kojumdgieva & Strachimirov 1960; Landau et al. 2013). The type species of Acrilla is the Recent Scalaria acuminata G. B. Sowerby II, 1844 from the Indo-West Pacific, which is a tall shell with numerous, evenly convex whorls lacking any shoulder angulation and with close-set, narrow, convex, sigmoidal axial ribs instead of lamellae (see Adams 1860: pl. 35, fig. 130; https: // www. gbif. org / occurrence / gallery? taxon _ key = 6126747). According to MolluscaBase eds (2024 c), Acrilla acuminata is the only living species of this genus. Acrilla differs distinctly from Anguloacrilla, which has comparatively low whorls with narrow subsutural ramp, thin lamellae and distinct spiral sculpture. Some Amaea species (type species Scalaria magnifica G. B. Sowerby II, 1844) may also develop rather low, convex whorls, but Amaea lacks shoulder angulation. Among the extant Amaea species, Amaea sericogazea (Masahito, Kuroda & Habe in Kuroda, Habe & Oyama, 1971) is most reminiscent of Anguloacrilla but lacks shoulder angulation and has no protruding lamellae close to the adapical suture (see Garcia 2003: fig. 18). The extant ‘ Gyroscala’ mikeleei García, 2003, from Indonesia, is reminiscent of Anguloacrilla in its slightly coronate shoulder and thin lamellae but lacks a basal angulation and does not develop a peribasal cord (see García 2003: figs 51 – 52) [this species was placed in Gyroscala de Boury, 1887 by García (2003) but differs strongly from the type species Gyroscala commutata (Monterosato, 1877), which is characterized by few, very prominent varices aligned axially].	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFC4FFC5FF3FFEF3FF5C91B1.taxon	description	Figs 7 B, 12 D – F	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFC4FFC5FF3FFEF3FF5C91B1.taxon	materials_examined	Type material. Holotype, SMF 380170 _ 12 - 1443 (XII 1443 a), SL: 25 mm, MD: 8.5 mm, CoŞteiu de Sus (Romania), illustrated in Zilch (1934: pl. 10, fig. 79), Fig. 12 D. Illustrated material. NHMW 1876 / 0011 / 0081, SL: 32.2 mm, MD: 11.2 mm, Lăpugiu de Sus (Romania), Figs 7 B, 12 E 1 – E 2. NHMW 1868 / 0001 / 0540, SL: 29.2 mm, MD: 10.5 mm, Lăpugiu de Sus (Romania), Figs 12 F 1 – F 2. Additional material. 3 spec., NHMW 1855 / 0043 / 0067, Lăpugiu de Sus (Romania). Revised description. Shell very large, moderately slender (apical angle ~ 25 °), finely coronate. Protoconch unknown. Teleoconch of about 13 relatively low whorls. Early teleoconch whorls moderately convex with slight mid-whorl angulation and distinctly impressed suture. Sculpture of prominent, strongly raised, convex axial ribs, no spiral sculpture. Later whorls moderately convex with narrow, shallow subsutural ramp and weakly angled shoulder, periphery placed mid-whorl. Sculpture of numerous, close-set, thin, not very high, only weakly prosocline lamellae not aligned axially (10: 45: 55), forming indistinct, short spines along shoulder. Crests of varices erect. Interspaces between lamellae with numerous, prominent primary and secondary spiral threads, separated by narrower interspaces. Crests of lamellae delicately granulose at intersections with spirals, especially close to abapical suture. Last whorl attaining 35 % of total height, with marked basal angulation, delimited by moderate peribasal cord, flattened basal disc. Lamellae weakening over base. Aperture subcircular with parietal concavity. Columella concave. Outer lip not thickened. Columellar callus forming very broad rim, adherent to but well delimited from base, parietal callus absent. Abapical part of the outer lip ends in short spout. Inner layer of peristome separated from outer layer by weak groove. No umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFC4FFC5FF3FFEF3FF5C91B1.taxon	discussion	Discussion. Anguloacrilla kimakowiczi (Boettger, 1896) is a rare species that was based on a spire fragment. The revision of the type specimen and the newly available material from Romania show that prominent spiral cords characterize adult shells of this species, whereas subadult specimens (like the holotype) lack conspicuous spiral sculpture. Anguloacrilla karamanensis sp. nov. from the Serravallian of the Karaman Basin (Turkey) differs in its much weaker spiral sculpture throughout ontogeny. It remains unclear if another Serravallian occurrence from France, mentioned by Cossmann (1912), represents either of these species. Similarly, the specimen illustrated by Janssen (1967) from the Late Burdigalian / Langhian of the North Sea Basin differs from Anguloacrilla kimakowiczi in its relatively weak and widely spaced lamellae and represents an additional yet undescribed species. Paleoenvironment. Unknown.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFC4FFC5FF3FFEF3FF5C91B1.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Pannonian Basin: Szokolya (Hungary) (Báldi 1960); Făget Basin: CoŞteiu de Sus, Lăpugiu des Sus (Romania) (Boettger 1902); Dacian Basin: Opanec (Bulgaria) (Kojumdgieva in Kojumdgieva & Strachimirov 1960).? Northeastern Atlantic. Serravallian (Middle Miocene): Aquitaine Basin: Saubrigues (France) (Cossmann 1912).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFC7FFC7FF3FF9F4FC0F959D.taxon	description	Figs 7 C – D, 12 A – C	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFC7FFC7FF3FF9F4FC0F959D.taxon	materials_examined	Type material. Holotype, NHMW 2024 / 0279 / 0002, SL: 24.8 mm, MD: 13.8 mm, Grund (Austria), Figs 7 C, 12 B 1 – B 2. Paratypes, NHMW 1859 / 0019 / 0109 a, SL: 21.2 mm, MD: 10.3 mm, Porzteich (Czech Republic), Figs 7 D, 12 A. NHMW 2024 / 0279 / 0003, SL: 21.6 mm, MD: 14.4 mm, Grund (Austria), Figs 12 C 1 – C 2. Type locality. Grund (Austria), North Alpine-Carpathian Foreland Basin. Type stratum. Grund Formation. Age. Middle Miocene, early Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFC7FFC7FF3FF9F4FC0F959D.taxon	etymology	Etymology. In honor of Werner Schwarzhans (Hamburg, Germany), in recognition of his contributions on the knowledge of the Paratethyan fish fauna.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFC7FFC7FF3FF9F4FC0F959D.taxon	diagnosis	Diagnosis. Large, moderately slender, finely coronate shell with narrow, shallow subsutural ramp and angled shoulder, bearing numerous, close-set lamellae forming indistinct spines along shoulder, prominent, close-set spiral cords in axial interspaces, differing from congeners in its fewer lamellae.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFC7FFC7FF3FF9F4FC0F959D.taxon	description	Description. Shell large, moderately slender (apical angle ~ 25 °), finely coronate. Protoconch unknown. Teleoconch of relatively low, moderately convex whorls with narrow, shallow subsutural ramp and angled shoulder. Periphery placed mid-whorl. Sculpture of numerous, moderately close-set, low, only weakly prosocline lamellae not aligned axially (~ 20: ~ 29 / 38: ~ 33 / 38), forming indistinct short spines along shoulder. Interspaces between lamellae with numerous fine spiral threads, separated by interspaces of about equal width. Secondary spiral cords may occasionally occur. Crests of lamellae erect and delicately granulose at intersections with spiral cords especially close to abapical suture. Last whorl with marked basal angulation, delimited by prominent peribasal cord, flat basal disc, lamellae weakening over base, numerous prominent spiral threads. Fasciole very short, broad, with prominent growth lines. Largely covered by columellar callus in some specimens. Aperture subcircular with parietal concavity. Columella concave. Outer lip not preserved. Columellar callus forming very broad rim, adherent to but well delimited from base, parietal callus delicate. Peristome not preserved. Abapical part of the outer lip ends in short spout. No umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFC7FFC7FF3FF9F4FC0F959D.taxon	discussion	Discussion. Anguloacrilla schwarzhansi sp. nov. is morphologically close to Anguloacrilla kimakowiczi (Boettger, 1896) but differs in its fewer lamellae, fewer spiral threads, and the more prominent peribasal cord. Moreover, A. kimakowiczi develops distinct secondary spiral cords. Another difference is the slightly wider and steeper subsutural ramp in A. schwarzhansi. Anguloacrilla lima sp. nov. is distinguished by its more numerous, closer set lamellae and the rounded shoulder. Paleoenvironment. At the locality Grund, channels formed in middle to outer neritic environments and were filled, with allochthonous assemblages uniting coastal-mudflat faunas with inner neritic ones (Zuschin et al. 2005; Roetzel 2009). The preservation suggests that the shells belonged to the inner neritic assemblages.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFC7FFC7FF3FF9F4FC0F959D.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): North Alpine-Carpathian Foreland Basin: Grund (Austria); Vienna Basin: Porzteich at Břeclav (Czech Republic).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFC6FFC0FF3FFB88FB529731.taxon	description	Figs 13 A 1 – A 2	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFC6FFC0FF3FFB88FB529731.taxon	materials_examined	Type material. Holotype, NHMW 1874 / 0058 / 1129, SL: 31.7 mm, MD: 11.2 mm, Akpınar (Turkey). Figs 13 A 1 – A 2. Five paratypes NHMW 1847 / 0058 / 1130. Type locality. Akpınar (Turkey), Karaman Basin. Type stratum. Silt and clay of the Týrtar Formation. Age. Middle Miocene, Serravallian.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFC6FFC0FF3FFB88FB529731.taxon	etymology	Etymology. Referring to the city Karaman in Turkey.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFC6FFC0FF3FFB88FB529731.taxon	diagnosis	Diagnosis. Very large, moderately slender, finely coronate shell with narrow, flat subsutural ramp, distinct shoulder angulation and numerous delicate lamellae, differing from congeners in its very weak spiral sculpture.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFC6FFC0FF3FFB88FB529731.taxon	description	Description. Shell very large, moderately slender (apical angle ~ 22 °), finely coronate. Protoconch unknown. Teleoconch of more than 11 relatively low whorls. Teleoconch whorls moderately convex with narrow, flat subsutural platform and distinctly angled shoulder, periphery placed mid-whorl. Sculpture of very numerous, thin, low, close-set, only weakly prosocline lamellae not aligned axially (~ 15: ~ 40: ~ 50), forming short spines along shoulder. Crests of lamellae erect and delicately granulose at intersections with spiral threads. Interspaces between lamellae with very delicate spiral threads. Last whorl attaining 35 % of total height, very narrow subsutural platform, convex below shoulder, marked basal angulation delimited by weak peribasal cord, flattened basal disc, lamellae rapidly weakening over base. Aperture subcircular. Columella concave. Outer lip not preserved. Columellar callus forming broad rim, adherent to but well delimited from base, parietal callus absent. Abapical part of the outer lip ends in distinct spout. Peristome not preserved. No umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFC6FFC0FF3FFB88FB529731.taxon	discussion	Discussion. This species was confused in Landau et al. (2013) with the Paratethyan Anguloacrilla kimakowiczi (Boettger, 1896), but they differ in the less convex early teleoconch whorls bearing less prominent lamellae, on later whorls the lamellae are much finer and less elevated, and the spiral sculpture is much weaker in Anguloacrilla karamanensis sp. nov. Acrilla lima sp. nov. differs in its much greater number of lamellae and Anguloacrilla schwarzhansi sp. nov. has fewer but more prominent lamellae and prominent spiral cords. The spout-like abapical portion of the aperture, which represents a short siphonal canal and typical for the genus Anguloacrilla nov. gen. is unusual amongst epitoniids and is more strongly developed in A. karamanensis. Paleoenvironment. Shallow marine environments (Landau et al. 2013). Proto-Mediterranean Sea. Karaman Basin: Akboğaz, Akpınar (Turkey) (Landau et al. 2013).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFC1FFC1FF3FFABCFD0692B5.taxon	description	Figs 7 A, 13 B 1 – B 2	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFC1FFC1FF3FFABCFD0692B5.taxon	materials_examined	Type material. Holotype, NHMW 1865 / 0001 / 0244, SL: 28.6 mm, MD: 11.3 mm, Lăpugiu de Sus (Romania), Figs 7 A, 13 B 1 – B 2. Type locality. Lăpugiu de Sus (Romania), Făget Basin. Type stratum. Silt and clay of the Dej Formation. Age. Middle Miocene, Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFC1FFC1FF3FFABCFD0692B5.taxon	etymology	Etymology. After lima (Latin for file), referring to the sculpture; noun in apposition.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFC1FFC1FF3FFABCFD0692B5.taxon	diagnosis	Diagnosis. Large, moderately slender shell with low, convex whorls, very dense sculpture of granulose lamellae and numerous delicate spiral threads in narrow axial interspaces, broad, well-developed abapical apertural spout.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFC1FFC1FF3FFABCFD0692B5.taxon	description	Description. Shell large, moderately slender (apical angle ~ 27 °). Protoconch and early teleoconch whorls unknown. Teleoconch of at least eight relatively low whorls. Early spire whorls convex with distinctly impressed suture. Later whorls moderately convex with very narrow subsutural ramp. Periphery placed mid-whorl. Sculpture of numerous, thin, low, densely spaced, weakly prosocline lamellae, not aligned axially (33: ~ 100: ~ 90). Crests of lamellae erect. Interspaces between lamellae with numerous, prominent spiral threads, separated by interspaces of equal width, resulting in extremely fine cancellate pattern. Last whorl attaining 35 % of total height, marked basal angulation delimited by narrow peribasal cord, flattened basal disc, lamellae indistinct over base. Aperture subcircular with parietal concavity, broad, well-developed abapical spout. Columella weakly convex. Outer lip not preserved. Columellar callus forming broad rim, detached from base, parietal callus absent. Peristome duplex. No umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFC1FFC1FF3FFABCFD0692B5.taxon	discussion	Discussion. This species is superficially reminiscent of Anguloacrilla kimakowiczi (Boettger, 1896) but is not distinctly coronate and differs in its much greater number of lamellae. The close-set lamellae also distinguish Anguloacrilla lima sp. nov. from other Paratethyan congeners. ‘ Acrilla’ leptoglyptamoena Sacco, 1891, from the Langhian or Burdigalian of the Colli Torinesi (Italy), and ‘ Acrilla’ praecurrens Sacco, 1891, from the Rupelian of Carcare (Italy), are reminiscent of Anguloacrilla lima sp. nov. in their densely spaced lamellae but lack any shoulder and their spiral sculpture equals the axial lamellae in strength (see Sacco 1891: pl. 2, figs 60 – 61, Ferrero Mortara et al. 1984: pl. 6, fig. 1). This species resembles Amaea species, but has a narrow, but distinct subsutural ramp and shoulder, absent in Amaea. Paleoenvironment. Unknown.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFC1FFC1FF3FFABCFD0692B5.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Făget Basin: Lăpugiu de Sus (Romania). Genus Cirsotrema Mörch, 1852	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFC1FFC1FF3FFABCFD0692B5.taxon	type_taxon	Type species. Scalaria varicosa Lamarck, 1822, by monotypy. Present-day, Indo-West Pacific.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFC1FFC1FF3FFABCFD0692B5.taxon	diagnosis	Diagnosis. “ Thick, massive shell. Size often quite large; turriculate, conical shape; slowly increasing, more or less gradate spire; acute, multispiral protoconch; convex, superimposed or subjoined whorls, with very deep, but generally invisible, suture; ornamentation composed of powerful axial lamellae, laminated or even frondose, more or less reflexed, depending on the species, sometimes almost joined and interspersed with large, even more prominent varicose veins; they are often spiny at the back; near the anterior suture, these lamellae form a retrocurrent fold which borders the suture almost continuously, and to which is welded - in the interval between two consecutive lamellae - an anterior deviation of the lamella of the preceding whorl, so that the whole completely masks the sutures; the whorl and lamellae are, moreover, adorned with spiral cords in the interspaces. Last whorl equal to a quarter - or at most a third - of the total height, slightly convex at the base, whose periphery is marked by an almost continuous cord, formed by a retrocurrent “ spike ” or “ buttress ” which is detached from the posterior part of each lamella; sometimes, this buttress is reduced to a simple thickening of the lamella, but it is never completely fused to the preceding lamella; the basal disc, between the discontinuous cordon and the imperforate center, is radiated by the sinuous lamina of the lamellae, which flatten considerably as they widen. Aperture approximately circular, with a very thick, continuous peristome, generally with a posterior auricle, and located in a vertical plane. ” (translated and slightly modified from Cossmann 1912: 50).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFC0FFC3FF3FFAAEFD9D9079.taxon	description	Figs 14 A – C, 15 C – D	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFC0FFC3FF3FFAAEFD9D9079.taxon	description	C [irsotrema]. (C [irsotrema].) crassicostatum (Desh.) — Sieber 1958: 129 [non Cirsotrema crassicostatum (Deshayes, 1850). [Cirsotrema duciei] var. crassicostanomala Sacco, 1891 — Ferrero-Mortara et al. 1984: 47, pl. 5, fig. 15.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFC0FFC3FF3FFAAEFD9D9079.taxon	materials_examined	Type material. Lectotype (designated herein): BS. 030.08.016, SL: 25.5 mm, MD: 15.7 mm, Colli Tortonesi, illustrated in Ferrero-Mortara et al. (1984: pl. 5, fig. 15). The origin of the shell described by Sacco (1891) is unclear. The collection label indicated the Late Miocene locality Tortona but Sacco (1891) doubted this provenance and suggested that the specimen is derived from a “ facies elveziana ”, which would be Burdigalian or Langhian. Illustrated material. NHMW 1872 / 0030 / 0080, SL: 46.8 mm, MD: 21.7 mm, Baden-Sooss (Austria), Figs 14 A 1 – A 2, 15 C. NHMW 1866 / 0040 / 0217, SL: 44.6 mm, MD: 22.8 mm, Baden-Sooss (Austria), illustrated in Hörnes (1856: pl. 46, fig. 7), Figs 14 B 1 – B 2, 15 D. NHMW 2013 / 0300 / 0137, SL: 44.4 mm, MD: 18.8 mm, Baden-Sooss (Austria), Figs 14 C 1 – C 2. Additional material. 10 spec., NHMW 2013 / 0078 / 0147, Baden (Austria); 11 spec., NHMW 1863 / 0015 / 1187, Baden-Sooss (Austria); 8 spec., NHMW 1869 / 0001 / 0111 Baden-Sooss (Austria); 15 spec., NHMW 2013 / 0300 / 0135, Baden-Sooss (Austria); 12 spec., NHMW 2013 / 0300 / 0136, Baden-Sooss (Austria); 9 spec., NHMW 1846 / 0037 / 0429, Möllersdorf (Austria); 2 spec., NHMW 1973 / 1615 / 0334, Bad Vöslau (Austria); 4 spec., NHMW 2010 / 0004 / 0521, Bad Vöslau (Austria); 5 spec., NHMW 199 z 0178 / 0809, Baden (Austria); 1 spec., NHMW 1863 / 0015 / 0799, Niederleis (Austria).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFC0FFC3FF3FFAAEFD9D9079.taxon	description	Description. Shell large, very robust, moderately slender (apical angle ~ 25 °). Protoconch and early teleoconch whorls unknown. Teleoconch of about ten low whorls. Early teleoconch whorls with narrow, moderately high lamellae and prominent spiral cords in interspaces forming cancellate sculpture. Later teleoconch whorls moderately convex with steep, weakly concave subsutural ramp and rounded shoulder, periphery placed just below mid-whorl. Suture moderately impressed, nearly completely obscured by lamellae. Sculpture ofclose-set, weakly prosocline, extremely broad, densely lamellose varices, not aligned axially, separated by deep, narrow interspaces (~ 20: ~ 12: 12 / 13). Crests of varices and interspaces with five to six prominent spiral cords. Spiral cords and interspaces with numerous delicate spiral threads causing delicately granulose crests. Last whorl attaining 45 % of total height, basal angulation delimited by moderately prominent peribasal cord, flattened basal disc. Varices prominent on base with reduces spiral sculpture. Fasciole short, broad with very prominent growth lines. Aperture circular. Columella concave. Outer lip strongly thickened by varix, basal lip broad. Columellar callus forming moderately broad rim, detached from base, parietal callus delicate. Short, poorly developed abapical spout. Peristome duplex. No umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFC0FFC3FF3FFAAEFD9D9079.taxon	discussion	Discussion. Cirsotrema crassicostanomala (Sacco, 1891) is characterized by its change in sculpture from cancellate early whorls to extremely broad, lamellose varices on late teleoconch whorls. In addition, it is comparatively broad and has a relatively weakly incised suture. Hörnes (1856) confused it with Cirsotrema lamellosum (Brocchi, 1814), form the Pliocene of the Mediterranean Sea. Cirsotrema lamellosum has a slightly gradate outline, less convex whorls and a weakly coronate shoulder (see Pinna & Spezia 1978: pl. 58, fig. 2; Landau et al. 2006: pl. 12, figs 5 – 6). Later, Sieber (1958) identified the Paratethyan species as Cirsotrema crassicostatum (Deshayes, 1850) from the early Miocene of France. Although C. crassicostatum was recently discussed by Cilia (2024) and Van der Voort (2024), it is still a poorly understood species. Its understanding was distorted by numerous illustrations of specimens from the Miocene of the North Sea Basin, which in our opinion are not conspecific with C. crassicostatum. Deshayes (1850: 42, pl. 70, figs 1 – 3) depicted a moderately slender species with gradate spire, subcylindrical whorls with weakly convex periphery, distinct subsutural shelf and broad, orthocline, internally lamellose varices (about 16 varices on last whorl). This morphology is quite different from C. crassicostanomala and they are clearly not conspecific. Specimens shown by Cossmann (1912: pl. 3, figs 1 – 3) as Cirsotrema crassicostatum are slenderer, weakly coronate, have wider spaced varices and a very deep suture. A specimen from the Early Miocene of France, illustrated by Cossmann & Peyrot (1922: pl. 4, figs 97 – 98), as C. crassicostatum, is much slenderer and has a narrower subsutural shelf. Specimens from the Langhian of Malta described by Cilia (2024: pl. 2, figs A – N) as Cirsotrema crassicostatum (Deshayes, 1850) differ from the Paratethyan species in the slenderer outline, less convex whorls with their periphery below mid-whorl being slightly angled, and the varices are wider spaced. The Miocene Maltese specimens are quite different from Cirsotrema crassicostatum and should be treated as Cirsotrema obliquatus Cowper Reed in Baldacchino, 1939. Similarly, the specimen from the Hemmoorian of the North Sea illustrated by Janssen (1984: pl. 60, fig. 15) differs from the Paratethyan specimens in its wider spaced and narrower varices and in its deeply incised suture. Other specimens from the Miocene of the North Sea Basin, illustrated by Van der Voort (2024: fig. 1) as Cirsotrema crassicostatum differ from the Paratethyan specimens in their very broad varices and strongly convex whorls with deeply incised suture. The early teleoconch whorls of this North Sea species bear narrow axial ribs and lack the cancellate sculpture of the Paratethyan specimens. Cirsotrema taurina (Sacco, 1891) (including its synonyms Cirsotrema pedemontanum (Sacco, 1891) and Cirsotrema rotundicosta (Sacco, 1891) is reminiscent of the Paratethyan species in its sculpture but differs in its incised suture and slenderer outline (see Sacco 1891: pl. 2 figs 12 – 15). Specimens from the Tortonian of Calabria, identified by Carone & Ardovini (2008) as C. crassicostatum might also represent C. taurina. In summary, we recognize a partly unresolved complex of large robust Cirsotrema species in the Circum-Mediterranean region of which C. crassicostanomala is the Paratethyan representative, C. crassicostatum is found in the early Miocene of France, C. obliquatus in the Miocene Proto-Mediterranean of Malta, C. taurina the early Miocene and possibly Tortonian of Italy, and at least two species occurred in the Miocene North Sea Basin. Paleoenvironment. Occurrences in the Baden Formation of the Vienna Basin suggest middle to outer neritic settings in up to 250 m water depth (Kranner et al. 2021).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFC0FFC3FF3FFAAEFD9D9079.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienna Basin: Baden, Baden-Sooss, Bad Vöslau, Möllersdorf, Niederleis (Austria) (Hörnes 1856). Proto-Mediterranean Sea. Tortonian (Late Miocene) of Colli Tortonesi (Italy) or Burdigalian / Langhian of Colli Torinesi (Italy) (Sacco 1891).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFC2FFCDFF3FF9E2FB43944D.taxon	description	Figs 14 D – E, 15 F * [Scalaria] C [irsotrema]. pumiceum var. dertonense Sacc. — Sacco 1891: 55, pl. 2, fig. 41.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFC2FFCDFF3FF9E2FB43944D.taxon	materials_examined	Type material. Lectotype (designated herein), BS. 030.08.029, SL: 16 mm, MD: 8.7 mm, Stazzano (Italy), Tortonian (Late Miocene), stored in the Museo Regionale di Scienze Naturali Torino (Italy), illustrated in Sacco (1891: pl. 2, fig. 41). Illustrated material. NHMW 1997 / 0178 / 0971 a, SL: 7.8 mm, MD: 4.3 mm, Bad Vöslau (Austria), Figs 14 D 1 – D 2, 15 F. NHMW 2010 / 0004 / 0523 a, SL: 8.3 mm, MD: 4.3 mm, Bad Vöslau (Austria), Figs 14 E, 15 H. Revised description. Shell medium-sized, robust, broad, markedly scalate (apical angle ~ 40 °), weakly coronate. Protoconch incomplete, high conical, last protoconch whorl with weak mid-whorl angulation. Teleoconch of five whorls, with broad, shallow, weakly concave subsutural ramp, strongly angled at shoulder forming periphery, weakly convex and constricted below. Suture deeply impressed, fenestrate. Sculpture of broad, prominent, close-set, prosocline, foliaceous varices, not aligned axially, separated by narrow interspaces (~ 16: ~ 18: 20; partly amalgamated). Varices on later teleoconch whorls often touching or amalgamating, forming spongiose, crisscross surface pattern on varix tops. Later whorls with two very prominent varices per whorl with dense sculpture, weakly coronate and bearing low spine at shoulder, with barely visible interspaces. Last whorl attaining 50 % of total height, with prominent basal angulation delimited by broad, flattened peribasal cord, slightly concave basal disc, varices weakening on base. Fasciole short, broad with very prominent growth lines. Aperture subcircular. Columella concave. Outer lip strongly thickened by varix, basal lip broad. Columellar callus and parietal callus delicate. Peristome duplex. No umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFC2FFCDFF3FF9E2FB43944D.taxon	discussion	Discussion. Cirsotrema dertonense (Sacco, 1891) differs from the Pliocene to Recent Atlanto-Mediterranean Cirsotrema pumiceum (Brocchi, 1814) in its stockier, more markedly scalate shell and the conspicuous internal crisscross pattern of the varices (see Landau et al. 2006: 40 for C. pumiceum). Cirsotrema sallomacense de Boury in Cossmann, 1912, from the Langhian of France, might be a closely related species, which differs in its much broader and higher spire whorls (see Cossmann 1912, pl. 6, fig. 13). Cirsotrema bourgeoisi de Boury in Cossmann, 1912, from the Middle and Late Miocene of France, is very similar and differs from C. dertonense mainly in the more prominent lamellae on the subsutural shelf and the more regular, fenestrate sculpture (see Landau et al. 2018: pl. 29, figs 1 – 2). Paleoenvironment. Occurrences in the Baden Formation of the Vienna Basin suggest middle to outer neritic settings in up to 250 m water depth (Kranner et al. 2021).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFC2FFCDFF3FF9E2FB43944D.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Polish-Ukrainian Fore-Carpathian Basin: Niechobrz (Poland) (Friedberg 1938); Korytnica Basin: Korytnica (Poland) (Bałuk 2006); Vienna Basin: Baden, Bad Vöslau (Austria) (hoc opus); Făget Basin: CoŞteiu de Sus (Romania) (Cossmann 1912). Proto-Mediterranean Sea. Tortonian (Late Miocene): Po Basin: Stazzano (Italy) (Sacco 1891).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFCCFFCDFF3FFD36FF5C910F.taxon	description	Figs 14 F	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFCCFFCDFF3FFD36FF5C910F.taxon	description	cf. Scala (Cirsotrema) fontannesi de Bourry [sic] — Csepreghy-Meznerics 1969 b: 74, pl. 2, fig. 5.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFCCFFCDFF3FFD36FF5C910F.taxon	materials_examined	Type material. Holotype, MD: 10.5 mm, Saubrigues (France), illustrated in Cossmann & Peyrot (1922: pl. 4, fig. 66). Illustrated material. M. 70.412, SL: 26.0 mm, MD: 18 mm, Balaton (Hungary), illustrated in Csepreghy-Meznerics (1969 a; pl. 4, fig. 7), Figs 14 F 1 – F 2.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFCCFFCDFF3FFD36FF5C910F.taxon	description	Description. Fragment of a large solid shell comprising penultimate whorl and last whorl. Whorls convex with weakly convex subsutural ramp and periphery placed mid-whorl. Sculpture of prominent, broad, internally densely lamellose, prosocline varices separated by interspaces of about equal width. Varices obliquely aligned axially, fused across suture (?: 16: 15). Spiral sculpture of one weak cord on subsutural ramp and three very prominent cords along periphery, forming elongate nodes on crests of varices. Sharp basal angulation with weak peribasal cord. Basal disk flat with broad varices. Aperture subcircular. Columellar and parietal callus continuous, forming narrow, adherent rim. Peristome duplex. No umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFCCFFCDFF3FFD36FF5C910F.taxon	discussion	Discussion. Cirsotrema fontannesi Cossmann & Peyrot, 1922 was based on a fragmentary specimen and remains a poorly understood species. The illustration in Cossmann & Peyrot (1922) is indeed reminiscent of the specimen from the Middle Miocene of Hungary, which was identified as the French species by Csepreghy-Meznerics (1969 a, b). We tentatively follow Csepreghy-Meznerics (1969 a, 1969 b) and consider the Paratethyan shell probably conspecific with the French specimen. The available specimen is strikingly similar to Cirsotrema crassicostanomalum (Sacco, 1891) and attains an identical size but has narrower varices, which lack a lamellose internal. Cirsotrema pedemontanum (Sacco 1891) (= Cirsotrema rotundicosta (Sacco, 1891), from the Early or Middle Miocene of Italy, differs in its broader, nearly orthocline varices and less prominent spiral cords (Sacco 1891: pl. 2, figs 14 – 15). Cirsotrema subaspromontanum (Sacco 1891), from the Tortonian (Late Miocene) of Italy, agrees in the prominent spiral cords, but differs in its subcylindrical whorls and the strongly angled base (Sacco 1891: pl. 2, fig. 18).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFCCFFCDFF3FFD36FF5C910F.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Bükk Mountains: Balaton (Hungary) (Csepreghy-Meznerics 1969 a). This species was also mentioned by Kochansky-Devidé (1956) from Zagreb (Croatia) but this record will need verification Northeastern Atlantic. Tortonian (Late Miocene): Aquitaine Basin: Saubrigues (France) (Cossmann & Peyrot 1922).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFCFFFCEFF3FFF2EFBAA908F.taxon	description	Figs 15 G – H, 16 A – F	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFCFFFCEFF3FFF2EFBAA908F.taxon	materials_examined	Type material. Holotype, NHMW 1851 / 0013 / 0086, SL: 12.7 mm, MD: 5.2 mm, Steinabrunn (Austria), illustrated in Hörnes (1856: pl. 46, figs 10 a – b), Figs 15 H, 16 A 1 – A 2. Illustrated material. NHMW 1865 / 0001 / 0585 a, SL: 14.4 mm, MD: 5.5 mm, Steinebrunn (Austria), Figs 16 B 1 – B 2. NHMW 1865 / 0001 / 0585 b, SL: 12.5 mm, MD: 5.0 mm, Steinebrunn (Austria), Figs 16 C 1 – C 2. NHMW 1865 / 0001 / 0585 c, SL: 12.9 mm, MD: 5.8 mm, Steinebrunn (Austria), Figs 16 D 1 – D 2. NHMW 2010 / 0004 / 0523 b, SL: 16.2 mm, MD: 6.3 mm, Bad Vöslau (Austria), Figs 15 G, 16 E 1 – E 2. NHMW 1855 / 0045 / 0205, SL: 17.7 mm, MD: 6.7 mm, Steinebrunn (Austria), Figs 16 F 1 – F 2. Additional material. 1 spec., NHMW 1859 / 0027 / 0189, Gainfarn (Austria); 10 spec., NHMW 1865 / 0001 / 0585, Steinebrunn (Austria): Revised description. Shell medium-sized, robust, moderately slender (apical angle ~ 22 °), weakly coronate. Protoconch unknown. Teleoconch of about eight whorls. Early teleoconch whorls convex, slightly gradate, with narrow, close-set, moderately high varices. Later teleoconch whorls moderately convex, subcylindrical, with narrow, shallow, weakly concave subsutural ramp, distinctly angled shoulder, and periphery placed mid-whorl. Suture deeply impressed, fenestrate, almost fully obscured by varices. Sculpture of numerous, close-set, very broad, internally lamellose, weakly prosocline varices, not aligned axially, separated by narrow interspaces (~ 20: ~ 16 / 18: 18; partly amalgamated). Varices touching each other at level of spiral cords, resulting in punctate to fenestrate surface. Crests of varices with densely packed foliaceous laminae resulting in porous surface. Each whorl with three very prominent, broad, swollen varices with dense internal sculpture. Varices weakly coronate along shoulder on some specimens on last teleoconch whorls. Interspaces barely visible. Last whorl attaining 40 % of total height. Last whorl with basal angulation, coinciding with very prominent peribasal cord, delimiting slightly concave basal disc, varices weakening on base. Fasciole short, broad with very prominent growth lines. Aperture circular. Columella concave. Outer lip strongly thickened by varix, basal lip broad. Columellar callus and parietal callus delicate. Peristome duplex. No umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFCFFFCEFF3FFF2EFBAA908F.taxon	discussion	Discussion. Cirsotrema hornesi (De Gregorio, 1889) is highly reminiscent of the Miocene to Recent Atlanto-Mediterranean Cirsotrema pumiceum (Brocchi, 1814), which differs in its more subcylindrical whorls, the subsutural shelf and angled shoulder, which gives it a slightly gradate appearance. In contrast to Cirsotrema hornesi the base of Cirsotrema pumiceum has a strongly angled base and wide basal disc. Subadult specimens of Cirsotrema hornesi, however, are almost indistinguishable from Cirsotrema pumiceum. Like the complex of large solid Cirsotrema species discussed under Cirsotrema crassicostatum (Deshayes, 1850), the smaller Cirsotrema species similar to C. pumiceum also turn out to represent a species complex. Paleoenvironment. Known from both shallow water and deep water settings.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFCFFFCEFF3FFF2EFBAA908F.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienna Basin: Baden, Bad Vöslau, Steinebrunn, Gainfarn (Austria) (Hörnes 1856; hoc opus). Proto-Mediterranean Sea. Langhian (Middle Miocene): Colli Torinesi: Sciolze (Italy) (Sacco 1891). Serravallian (Middle Miocene): Karaman Basin: (Turkey) (Landau et al. 2013).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFCEFFC8FF3FFA30FD3E9689.taxon	description	Figs 17 C 1 – C 2	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFCEFFC8FF3FFA30FD3E9689.taxon	materials_examined	Type material. Holotype, PAL 2022.54.1, SL: 22.0 mm, MD: 10.5 mm, Parádfürdö, Ilona Valley (Hungary), illustrated in Kovács et al. (2023: figs 58 – 59), Figs 17 C 1 – C 2. Revised description. Shell large, robust, moderately slender (apical angle ~ 23 °). Protoconch and early teleoconch whorls unknown. Teleoconch whorls convex with broad, moderately steep, weakly concave, very poorly delimited subsutural ramp, with periphery just below mid-whorl. Suture moderately impressed, relatively shallow. Sculpture of very numerous, close-set, narrow, prosocline, weakly frondose axial ribs, not strictly aligned axially (23 axial ribs on penultimate whorl). Axial ribs overrun by about five spiral cords of equal strength to ribs, forming fine cancellate pattern. Three very prominent, convex varices per whorl vaguely obliquely aligned axially. Secondary spiral cords intercalated on last whorl numbering about 12. Base slowly contracting with narrow, weak basal cord and flattened disc. Aperture subcircular. Columella concave. Outer lip strongly thickened by varix, basal lip broad. Peristome very broad, duplex. No umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFCEFFC8FF3FFA30FD3E9689.taxon	discussion	Discussion. We are not aware of comparable species in the Circum-Mediterranean region. Paleoenvironment. Agitated, coastal marine (Kovács et al. 2023).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFCEFFC8FF3FFA30FD3E9689.taxon	distribution	Distribution in Central Paratethys. Ottnangian (Early Miocene): Early Miocene North Hungarian Basin: Parádfürdö (Hungary) (Kovács et al. 2023).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFC9FFC8FF3FFEF2FEED93BF.taxon	description	Figs 17 B	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFC9FFC8FF3FFEF2FEED93BF.taxon	materials_examined	Type material. Lectotype (designated by Zilch 1934: 230), SMF 380166 _ 12 - 1447 (XII 1447 a), SL: 9.5 mm, MD: 4.5 mm, CoŞteiu de Sus (Romania), illustrated in Zilch (1934: pl. 10, fig. 77), Figs 17 B. Revised description. Shell small, robust, moderately broad (apical angle ~ 29 °), gradate, faintly coronate. Protoconch and first teleoconch whorls unknown. Early teleoconch whorls convex with convex subsutural ramp, strongly raised laminae and three prominent spiral cords in axial interspaces. Later teleoconch whorls with broad, shallow, weakly concave subsutural ramp and weakly angled shoulder, subcylindrical below, with periphery placed mid-whorl. Suture moderately impressed, fenestrate. Sculpture of moderately close-set, prominent, strongly raised, weakly prosocline, frondose lamellae separated by wider interspaces, partly aligned axially and partly fused to lamellae on preceding whorl (~ 14: ~ 16: 20). Lamellae weakly coronate along shoulder of last two whorls. Axial interspaces with three low, broad spiral cords below shoulder. Last whorl attaining ~ 45 % of total height, with sharp basal angulation delimited by prominent, slightly beaded peribasal cord, basal disc slightly concave, lamellae only slightly weaker on base. Fasciole short, broad, with very prominent growth lines. Aperture circular. Columella concave. Outer lip strongly thickened by varix. Columellar callus forming narrow rim, detached from base, parietal callus delicate. Peristome duplex. Umbilicus reduced to narrow chink.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFC9FFC8FF3FFEF2FEED93BF.taxon	discussion	Discussion. Cirsotrema loerentheyi (Boettger, 1902) is characterized by its cancellate sculpture of frondose lamellae and low, broad spiral cords. Cirsotrema miovaricosum (Sacco, 1891), from the Tortonian of Stazzano (Italy), has comparable sculpture but differs in its broader and orthocline varices and the greater number of narrower spiral cords (see Ferrero Mortara et al. 1984: pl. 5, fig. 16). Cirsotrema couffoni de Boury in Cossmann, 1912, from the Tortonian of the Loire Basin (France), might represent a closely related species that differs mainly in the slenderer shape and steeper subsutural ramp (see Cossmann 1912: pl. 5, fig. 11). Cirsotrema duciei (Wright, 1855), from the Langhian of Malta, has comparable sculpture but is slenderer, has a weaker shoulder and has more numerous, less frondose lamellae (see Wright 1855: pl. 7, fig. 3; Cilia 2024: pl. 3, figs J – M). Paleoenvironment. Unknown.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFC9FFC8FF3FFEF2FEED93BF.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Făget Basin: CoŞteiu de Sus (Romania) (Boettger 1906).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFC9FFC9FF3FF998FD599411.taxon	description	Figs 17 D – E	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFC9FFC9FF3FF998FD599411.taxon	materials_examined	Type material. Holotype, NMBE Nr. B 7648, SL: 24 mm, MD: 11 mm, Hohburggraben, Belpberg, illustrated in Pfister & Wegmüller (2007: pl. 2, figs 5 – 9), Figs 17 D 1 – D 3. Paratype, NMBE Nr. B 7649, SL: 18 mm, MD: 10 mm, Hohburggraben, Belpberg, illustrated in Pfister & Wegmüller (2007: pl. 2, figs 10 – 12), Figs 17 E 1 – E 3. Pictures: Peter Vollenweider. Revised description. Shell large, turreted, moderately slender (apical angle ~ 20 °), weakly coronate. Protoconch and first teleoconch whorls unknown. Early teleoconch whorls very high, subcylindrical to weakly scalate with narrow subsutural ramp and marked shoulder angulation. Suture deeply impressed, largely obscured by varices. Sculpture of prominent, close-set, prosocline to weakly sigmoidal varices with flat crests and narrow interspaces (?: 18 / 20: 21). Several very prominent, convex varices occurring at irregular intervals. Up to ten prominent spiral cords in interspaces, forming small nodes on weaker varices. Protruding varices on last whorl with low spines at shoulder. Last whorl attaining about 40 % of total height, moderately convex with weak basal angulation and very prominent peribasal cord, separating moderately narrow basal disc bearing prominent varices. Aperture subcircular. Columella concave. Fasciole short, broad, with prominent growth lines. Columellar callus and parietal callus forming continuous moderately broad rim, detached from base. Peristome poorly preserved, duplex. No umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFC9FFC9FF3FF998FD599411.taxon	discussion	Discussion. Cirsotrema menkveldae Pfister & Wegmüller, 2007 probably belongs to the C. pumiceum (Brocchi, 1814) species group but differs from other species in the group by the slightly sigmoidal varices and the greater number of protruding varices. Cirsotrema dertonense (Sacco, 1891) is stockier and has a much broader subsutural shelf. Paleoenvironment. Shallow marine, inner neritic.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFC9FFC9FF3FF998FD599411.taxon	distribution	Distribution in Central Paratethys. Eggenburgian / Ottnangian (Early Miocene): North Alpine Foreland Basin: Belpberg (Switzerland) (Pfister & Wegmüller 2007).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFCBFFCAFF3FFF2EFF6093B1.taxon	description	Figs 15 I, 18 A 1 – A 2	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFCBFFCAFF3FFF2EFF6093B1.taxon	materials_examined	Type material. Holotype, NHMW 1863 / 0015 / 0803, SL: 12.5 mm, MD: 5.7 mm, Niederleis (Austria), Figs 15 I, 18 A 1 – A 2 Type locality. Niederleis (Austria), Vienna Basin. Type stratum. Baden Formation. Age. Middle Miocene, middle Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFCBFFCAFF3FFF2EFF6093B1.taxon	etymology	Etymology. Referring to the microperforate appearance of the crests of the varices.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFCBFFCAFF3FFF2EFF6093B1.taxon	diagnosis	Diagnosis. Medium-sized, robust, moderately slender, moderately gradate with faintly coronate last whorls, numerous broad, uniform, close-set varices with microperforated crests, narrower interspaced with numerous spiral threads, sharp basal angulation.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFCBFFCAFF3FFF2EFF6093B1.taxon	description	Description. Shell medium-sized, robust, moderately slender (apical angle ~ 30 °), later whorls faintly coronate. Protoconch and first teleoconch whorls unknown. Teleoconch of about ten whorls. Early teleoconch whorls with narrow, moderately high varices and prominent spiral cords in interspaces, resulting in cancellate pattern. Later teleoconch whorls with broad, shallow, weakly concave subsutural ramp and weakly angled shoulder, subcylindrical below, with periphery placed above mid-whorl. Suture deeply impressed, largely covered by varices. Sculpture of numerous, uniform, densely spaced, broad, weakly prosocline varices, not aligned axially, separated by deep, narrower interspaces (~ 20: ~ 25: 28). Crests of varices with fine foliaceous crisscross pattern, resulting in microperforate surface appearance. Varices weakly coronate along shoulder, slightly concave on subsutural ramp. Interspaces with numerous spiral threads. Last whorl attaining 43 % of total height, with marked basal angulation delimited by very prominent peribasal cord, basal disc slightly concave, varices prominent on base, spiral sculpture reduced. Fasciole short, broad with very prominent growth lines. Aperture circular, columella concave. Outer lip strongly thickened by varix, basal lip broad. Columellar callus forming moderately broad rim, detached from base, parietal callus delicate. Peristome duplex. No umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFCBFFCAFF3FFF2EFF6093B1.taxon	discussion	Discussion. This species is unique within the Paratethyan Epitoniidae due to its elegant sculpture of regularly spaced, close-set and uniform varices. The finely pitted surface appearance of the varix tops we have not seen in any congener. Cirsotrema loerentheyi (Boettger, 1902) differs in it cancellate sculpture and Cirsotrema crassicostanomalum (Sacco, 1891) has broader varices overrun by spiral cords. We are not aware of comparable species from the Miocene of the northeastern Atlantic and Proto-Mediterranean Sea. Cirsotrema duciei (Wright, 1855), from the Langhian of Malta, has a comparable outline and close-set varices of similar width but differs in slenderer shell and spiral cords, which overrun the varices (see Wright 1855: pl. 7, fig. 3; Cilia 2024: pl. 3, figs J – M). Paleoenvironment. At the Niederleis section, sediment and biota from coastal and lagoonal environments not exceeding 30 m water depth were transported by tempestites into offshore settings (Mandic et al. 2002).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFCBFFCAFF3FFF2EFF6093B1.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienna Basin: Niederleis (Austria) (hoc opus).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFCBFFCBFF3FF9AAFEEE9188.taxon	description	Figs 15 J, 18 D – F	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFCBFFCBFF3FF9AAFEEE9188.taxon	materials_examined	Type material. Holotype, NHMW 2020 / 0138 / 0360 a, SL: 6.5 mm, MD: 3.0 mm, Figs 18 D 1 – D 2. Paratypes, NHMW 2020 / 0138 / 0360 b, SL: 13.0 mm, MD: 7.8 mm, Grund (Austria), Figs 18 E 1 – E 2. NHMW 2007 / 0086 / 0166, SL: 9.7 mm, MD: 6.2 mm, Grund (Austria), Figs 15 J, 18 F 1 – F 2. Type locality. Grund (Austria), North Alpine-Carpathian Foreland Basin. Type stratum. Grund Formation. Age. Middle Miocene, early Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFCBFFCBFF3FF9AAFEEE9188.taxon	etymology	Etymology. From nudus (Latin for naked), referring to the comparatively reduced sculpture.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFCBFFCBFF3FF9AAFEEE9188.taxon	diagnosis	Diagnosis. Small, robust, very broad, somewhat scalate shell with marked shoulder, relatively narrow varices, narrowing on slightly concave subsutural ramp, sharp basal angulation. Revised description. Shell small, robust, very broad, somewhat scalate (apical angle ~ 35 – 44 °). Protoconch unknown. Teleoconch of more than five whorls. Teleoconch whorls with broad, moderately shallow, weakly concave subsutural ramp, marked shoulder angulation delimiting periphery placed just above mid-whorl, constricted below. Suture deeply impressed, largely obscured by varices. Sculpture of prominent, close-set, relatively narrow, weakly prosocline, internally lamellose varices not strictly aligned axially (~ 13: ~ 21: 16), narrowing over subsutural ramp. Later whorls with up to three prominent varices per whorl. Interspaces with about four, very weak spiral cords. Last whorl attaining ~ 50 % of total height, with prominent basal angulation delimited by broad peribasal cord, basal disc slightly concave, varices weakening on base. Fasciole short, broad with very prominent growth lines. Aperture subcircular. Columella concave. Outer lip strongly thickened by varix, basal lip broad. Columellar and parietal callus forming narrow, continuous rim, detached from base. Peristome duplex. No umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFCBFFCBFF3FF9AAFEEE9188.taxon	discussion	Discussion. Cirsotrema nudum sp. nov. is reminiscent of Cirsotrema dertonense (Sacco, 1891) in its somewhat scalate outline but differs from that species in its much weaker varices, which lack the complex, internal crisscross structure. Cirsotrema microperforata sp. nov. has broader varices with flat crests and a characteristic microperforate surface. Moreover, C. microperforata lacks protruding varices on the last whorl. Paleoenvironment. At the locality Grund channels formed in middle to outer neritic environments and were filled, with allochthonous assemblages uniting coastal-mudflat faunas with inner neritic ones (Zuschin et al. 2005; Roetzel 2009). The preservation suggests that the shells of Cirsotrema nudum sp. nov. belonged to the inner neritic assemblages.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFCBFFCBFF3FF9AAFEEE9188.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): North Alpine-Carpathian Foreland Basin: Grund (Austria).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD5FFD5FF3FFF2EFBD897F9.taxon	description	Figs 19 A – B	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD5FFD5FF3FFF2EFBD897F9.taxon	materials_examined	Type material. Holotype, BSPG 2007 X 110, SL: 4.4 mm, MD: 2.8 mm, Gurlan (Germany), illustrated in Schneider et al. (2009: fig 5 b), Fig. 19 A. Paratype, BSPG 2007 X 111, SL: 3.0 mm, MD: 2.5 mm, Gurlan (Germany), illustrated in Schneider et al. (2009: fig 5 c), Fig. 19 B. Stored in Bayerische Staatssammlung für Paläontologie und Geologie, Munich (Germany). Type locality. Gurlan (Germany), North Alpine Foreland Basin. Type stratum. Upper Marine Molasse. Age. Early Miocene, early Ottnangian (mid-Burdigalian).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD5FFD5FF3FFF2EFBD897F9.taxon	etymology	Etymology. In honor of Jürgen Pollerspöck (Zoologische Staatssammlung München) in recognition of his contributions on paleoichthyology.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD5FFD5FF3FFF2EFBD897F9.taxon	diagnosis	Diagnosis. Small, moderately slender shell with convex whorls, deeply incised suture, faintly coronate shoulder, very prominent, close-set, moderately broad, prosocline, internally lamellose varices, five prominent spiral cords in interspaces, narrow peribasal cord and wide basal disc.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD5FFD5FF3FFF2EFBD897F9.taxon	description	Description. Small, moderately slender (apical angle ~ 25 °). Protoconch and early teleoconch whorls unknown. Teleoconch whorls convex with deeply impressed suture, narrow subsutural shelf with faintly coronate shoulder. Very prominent, close-set, moderately broad, prosocline, internally lamellose varices separated by deep, slightly wider or narrower interspaces with five prominent spiral cords. Three more prominent varices on last whorl. Last whorl with marked basal angulation delimited by narrow peribasal cord, wide basal disc, varices but narrowing prominent on base. Fasciole short, broad, with strong growth lines. Aperture subcircular. Columella concave. Outer lip and basal lip strongly thickened. Columellar and parietal callus forming narrow, continuous rim, detached from base. Peristome strongly thickened, duplex. No umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD5FFD5FF3FFF2EFBD897F9.taxon	discussion	Discussion. Schneider et al. (2009) illustrated two fragments, identified by these authors as Cirsotrema miovaricosum (Sacco, 1891). However, the holotype of C. miovaricosum, from the Tortonian of S. Agata (Italy), differs in its much narrower varices and the cancellate sculpture (see Ferrero-Mortara et al. 1984: pl. 5, figs 16 a – b). Cirsotrema crassicostatum (Deshayes, 1850) [including Cirsotrema thais de Boury in Cossmann, 1912, as illustrated in Cossmann (1912)] differs in the less convex whorls, broader subsutural shelf and the fewer and less prosocline varices (see Cossmann 1912: pl. 3, figs 1 – 3, pl. 6, fig. 16). Cirsotrema subsequenzai (Sacco, 1891), from the Tortonian of S. Agata (Italy) is reminiscent of Cirsotrema pollerspoecki sp. nov. in whorl profile and sculpture but differs in its broader varices, and it lacks a basal angulation with peribasal cord (see Sacco 1891: pl. 2, fig. 17). Cirsotrema pollerspoecki sp. nov. is reminiscent of the Middle Miocene Cirsotrema hornesi (De Gregorio, 1889) but its varices are not fused and differs in its weaker peribasal basal cord and wider basal disc. Paleoenvironment. Agitated rocky shore and coastal environments (Schneider et al. 2009).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD5FFD5FF3FFF2EFBD897F9.taxon	distribution	Distribution in Central Paratethys. Ottnangian (Early Miocene): North Alpine Foreland Basin: Gurlan (Germany) (Schneider et al. 2009), Reinbach at Schärding (Austria) (Steininger 1973).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD4FFD6FF3FFE62FF5C976D.taxon	description	Figs 15 E, 18 B – C	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD4FFD6FF3FFE62FF5C976D.taxon	materials_examined	Type material. Holotype, Nr. 5022, Museo Civico di Storia Naturale di Milano, SL: 15 mm, MD: 7 mm, Neogene, Piacentino (Italy), illustrated in Pinna & Spezia (1978: pl. 58, fig. 1). Illustrated material. NHMW 2010 / 0004 / 0524, SL: 19.2 mm, MD: 8.7 mm, Bad Vöslau (Austria), Figs 15 E, 18 B 1 – B 2. NHMW 1860 / 0040 / 0302, SL: 27.0 mm, MD: 11.7 mm, Lăpugiu de Sus (Romania), Figs 15 L, 18 C 1 – C 2. Revised description. See Landau et al. (2006: 40).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD4FFD6FF3FFE62FF5C976D.taxon	discussion	Discussion. See Landau et al. (2006: 40). It is worth mentioning that Cirsotrema hornesi (De Gregorio, 1889) and C. pumiceum are found together in the Paratethys and therefore do not form a phylogenetic lineage. They might have exploited slightly different habitats. In the extant Atlantic faunas C. pumiceum was formally considered an amphiatlantic species (Garcia Talavera 1983). However, De Jong & Coomans (1988: 53) separated the western Atlantic species as C. dalli Rehder, 1945 based on the color of the animal (Landau et al. 2006: 41). Paleoenvironment. Lives on sand and detritus bottoms in shallow water, between 8 – 60 m deep. Probably associated with Anemonia sulcata (Poppe & Goto 1991).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD4FFD6FF3FFE62FF5C976D.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Korytnica Basin: Korytnica (Poland) (Bałuk 2006); Vienna Basin: Bad Vöslau, Gainfarn, Niederleis (Austria); Pannonian Basin: Várpalota (Hungary) (Strausz 1966). Făget Basin: Lăpugiu de Sus (Romania) (hoc opus). Proto-Mediterranean Sea. Tortonian (Late Miocene): Po Basin: Tortona (Italy) (de Boury 1890); Pliocene: central Mediterranean (Italy) (De Stefani 1878; Sacco 1891; Pavia 1975; Cavallo & Repetto 1992). Northeastern Atlantic. Pliocene: Guadalquivir Basin (Spain) (González Delgado 1986); Estepona Basin (Landau et al. 2006);	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD4FFD6FF3FFE62FF5C976D.taxon	description	Present-day. Mediterranean Sea, Atlantic (Morocco, Angola, Azores) (Ardovini & Cossignani 2004; Chirli 2009).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD7FFD7FF3FFED6FB3D97DD.taxon	description	Figs 19 C 1 – C 2	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD7FFD7FF3FFED6FB3D97DD.taxon	materials_examined	Type material. Holotype, NHMW 2014 / 0379 / 0032, SL: 33 mm, MD: 13.5 mm, Allerding (Austria), Figs 19 C 1 – C 2. Type locality. Allerding (Austria), North Alpine Foreland Basin. Type stratum. Ottnang Formation Age. Early Miocene, Ottnangian (mid-Burdigalian).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD7FFD7FF3FFED6FB3D97DD.taxon	etymology	Etymology. In honor of Simon Schneider (CASP, Cambridge), in recognition of his contributions to Miocene paleontology.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD7FFD7FF3FFED6FB3D97DD.taxon	diagnosis	Diagnosis. Very large, robust, slender shell with superficial suture, prominent subsutural collar and almost straight sided whorls, sculpture of broad varices with flat crests and five weaker spiral cords in deep, narrow axial interspaces.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD7FFD7FF3FFED6FB3D97DD.taxon	description	Description. Shell very large, robust, slender (apical angle ~ 16 °). Protoconch and early teleoconch whorls unknown. Early teleoconch whorls with broad varices and prominent spiral cords in axial interspaces, resulting in cancellate pattern. Later teleoconch whorls with prominent subsutural collar shallow subsutural concavity, weakly convex below mid-whorl. Suture fully obscured by varices. Sculpture of very broad, close-set, weakly prosocline, band-like flattened varices, not aligned axially, separated by deep, moderately narrow interspaces (?: 13: 14). Adapical tips of varices forming prominent subsutural collar. Deep interspaces with five prominent spiral cords. Last whorl attaining about 42 % of total height, basal angulation delimited by broad, low peribasal cord, basal disc relatively narrow, varices weak on base with reduces spiral sculpture. Fasciole very short, broad with very prominent growth lines. Aperture subcircular. Columella concave. Outer lip slightly thickened by varix, basal lip broad. Columellar callus forming moderately broad rim, parietal callus thin, adherent. Peristome poorly preserved. No umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD7FFD7FF3FFED6FB3D97DD.taxon	discussion	Discussion. This species was erroneously identified by Harzhauser et al. (2014) as Cirsotrema crassicostanomalum (Sacco, 1891), from the Burdigalian of the Colli Torinesi (Italy), due to comparable sculpture. However, Cirsotrema schneideri sp. nov. differs from C. crassicostanomalum in its much slenderer shell, the prominent subsutural collar and the very superficial suture. The same features distinguish it from the Pliocene Cirsotrema lamellosum (Brocchi, 1814) and the Miocene Cirsotrema crassicostatum (Deshayes, 1850). Paleoenvironment. Agitated, rocky shore (Harzhauser et al. 2014).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD7FFD7FF3FFED6FB3D97DD.taxon	distribution	Distribution in Central Paratethys. Ottnangian (Early Miocene): North Alpine Foreland Basin: Allerding (Austria). Genus Clathroscala de Boury, 1890	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD7FFD7FF3FFED6FB3D97DD.taxon	type_taxon	Type species. Turbo cancellatus Brocchi, 1814; original designation by de Boury (1890: 215). Miocene, Italy.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD7FFD7FF3FFED6FB3D97DD.taxon	diagnosis	Diagnosis. “ Shell fairly solid, though not very thick. Medium size; elongated, conical shape; spire turriculate, not gradate, with convex, more or less regularly cancellate whorls, with punctations on the test between ribs; deep sutures, not bordered or canaliculated; axial ribs numerous, filiform, not lamellar, not deviated backwards on the sutures, often a little oblique, almost always interspersed with thicker, more widely flattened varices; these ribs are cut by spiral cords, fairly large and spaced apart. Last whorl equal to about a quarter of the total height, convex or subangular at the periphery of the base, which bears a fairly thick disc, often scalloped in outline between the ribs; this disc is concentrically furrowed and obscurely radiated by the trace of the ribs immersed in it; in the center, there is no trace of umbilicus or bead. Rounded aperture, with fairly thick peristome, devoid of auricle; the inner layer is very thin and continuous, the outer layer discontinuous, thin on the columellar edge, bordered by a varix on the labrum which is a little sinuous and sometimes slightly oblique to the vertical axis. ” (translated from Cossmann 1912: 70).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD7FFD7FF3FFED6FB3D97DD.taxon	discussion	Discussion. This genus is known from a few Pliocene species from the Mediterranean Sea, northwestern Atlantic and the North Sea (e. g., Van Dingenen et al. 2016). The Middle Miocene Paratethyan Clathroscala xeniae (Boettger, 1906) is the oldest member of the genus, followed by Clathroscala bureaui de Boury in Cossmann, 1912 and C. cancellata (Brocchi, 1814) from the Late Miocene of France (Landau et al. 2023).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD6FFD1FF3FFE47FDA99549.taxon	description	Fig 20 A, 21 A	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD6FFD1FF3FFE47FDA99549.taxon	materials_examined	Type material. Lectotype (designated by Zilch 1934: 230), SMF 380171 _ 12 - 1450 a (XII 1450 a), SL: 9.5 mm, MD: 3.5 mm, CoŞteiu de Sus (Romania), illustrated in Zilch (1934: pl. 10, fig. 80), Fig 20 A, 21 A. Revised description. Small, moderately slender conical shell of at least nine teleoconch whorls (apical angle 22 °). Protoconch and first teleoconch whorls unknown. Teleoconch whorls convex with indistinct, weakly concave subsutural ramp, evenly rounded periphery placed just above mid-whorl, separated by deeply impressed suture. Sculpture on early teleoconch whorls of very weak, close-set axial ribs (~ 26: ~ 24: ~ 24). No spiral sculpture. Later whorls with slightly wider spaced, low, convex axial ribs, separated by wider interspaces, with five slightly weaker spiral cords, forming regularly cancellate pattern. Delicate spiral threads and growth lines in interspaces forming finely punctate surface. About one prominent varix per whorl. Last whorl about 35 % of total height, weak basal angulation delimited by narrow peribasal cord, axial ribs terminate at peribasal cord, base flattish bearing weak spiral cords. Aperture ovate, columella weakly concave. Columellar callus forming moderately broad rim, weakly detached from base. Parietal callus delicate. Peristome duplex with thin inner layer; outer lip thickened by varix. Umbilicus reduced to narrow chink.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD6FFD1FF3FFE47FDA99549.taxon	discussion	Discussion. Clathroscala cancellata (Brocchi, 1814), from the Pliocene of the Mediterranean Sea, is much larger (SL: 30 mm) and has more spiral cords (see Pinna & Spezia 1978: pl. 60, fig. 1; Landau et al. 2006: pl. 13, figs 3 – 4; Chirli 2009: pl. 18, figs 2 – 7). Clathroscala woodi (Deshayes, 1864), from the Pliocene of the North Sea, is more delicate and more elongate (see Harmer 1920: pl. 68, fig. 9). The Paratethyan species is very similar to the apical fragment from the Tortonian of France figured by Landau et al. (2023: pl. 11, fig. 2) as C. cancellata. The French specimen has one more spiral cord and a slightly wider apical angle. Clathroscala bureaui de Boury in Cossmann, 1912 has a much narrower apical angle and differs in its taller and turreted shape, its cords are narrower and alternate in strength, and the basal sculpture is more pronounced. Paleoenvironment. Unknown.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD6FFD1FF3FFE47FDA99549.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Bükk Mountains: Borsodbóta (Hungary) (Csepreghy-Meznerics 1969 a, b); Făget Basin: CoŞteiu de Sus (Romania) (Boettger, 1906). Genus Clathrus Agassiz, 1837	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD6FFD1FF3FFE47FDA99549.taxon	type_taxon	Type species. Scalaria acuta J. Sowerby, 1812, subsequent designation by Brown & Neville (2015: 21). Eocene, British Isles	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD6FFD1FF3FFE47FDA99549.taxon	diagnosis	Diagnosis. Medium-sized, moderately stocky conical, gradate, often coronate, with weakly convex to subcylindrical whorls, moderately wide, weakly concave subsutural shelf. Suture deeply impressed. Moderately close-set, prominent, prosocline robust, strongly raised lamellae with rolled over crests. Interspaces with broad, low, convex spiral cords. Lamellae not aligned axially, disconnected along suture. Last whorl with sub-angled base without continuous peribasal cord, lamellae prominent on base, strongly rolled over. Short, broad fasciole with prominent growth lines. Aperture subcircular, columellar callus forming broad rim detached from base, parietal callus weak, peristome thickened, duplex.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD6FFD1FF3FFE47FDA99549.taxon	discussion	Discussion. Clathrus was originally introduced by Oken (1815: 255) but his work was placed on the Official Index by Opinion 417 (ICZN 1956). Later, the name was made available by Agassiz (1837: 35). The complex nomenclatorial history was reviewed by Brown & Neville (2015: 21), who clarified the situation by the subsequent designation of Scalaria acuta J. Sowerby, 1812 from the Late Eocene of the British Isles as type species [for illustrations see Sowerby (1812: pl. 16, lower two figs), Duerr 2004: fig. 9 and Savazzi & Saski (2004: fig 11 A – B, H)]. Sacco (1891) used Clathrus for several Miocene species from Italy and Cossmann (1912: 36) gave a first detailed diagnosis for Clathrus but this was based on an incorrect type species [Scalaria communis Lamarck, 1822, = Scala clathrus (Linnaeus, 1758)]. Therefore, Clathrus sensu Cossmann 1912 is an objective synonym of Scala Mörch, 1852. This erroneous concept was also adopted by Weil et al. (1999). Therefore, the understanding of Clathrus varied greatly in the literature and no diagnosis was available so far for Clathrus sensu Brown & Neville (2015). Duerr (2004) placed Scalaria acuta J. Sowerby, 1812 in Cirsotrema, which would made Cirsotrema Mörch, 1852 a subjective junior synonym of Clathrus Agassiz, 1837. Indeed, Clathrus as understood herein comprises a group of extant species, which are currently placed in Cirsotrema or Boreoscala Kobelt, 1902 by most authors (e. g., Boreoscala ponderosa Garcia, 2003, Elegantiscala fimbriatula Masahito, Kuroda & Habe in Kuroda, Habe & Oyama, 1971, Epitonium (Boreoscala) blainei Clench & R. D. Turner, 1953), Scalaria latecostata Strebel, 1905, Scalaria zelebori Dunker in Dunker & Zelebor, 1866). However, these species differ from Cirsotrema in their simple, non-lamellose lamellae / varices and their lamellae do not fuse laterally as seen in many Cirsotrema species. The lamellae of Clathrus are also not fused across the suture as in Boreoscala and several Cirsotrema species. Therefore, we consider Clathrus a valid genus with the type species Scalaria acuta J. Sowerby, 1812, and not a synonym of Scala Mörch, 1852 with the type species Turbo clathrus Linnaeus, 1758 nor Cirsotrema, type species Scalaria varicosa Lamarck, 1822.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD0FFD2FF3FFC33FD909341.taxon	description	Figs 15 K, 17 A 1 – A 2	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD0FFD2FF3FFC33FD909341.taxon	materials_examined	Type material. Holotype, NHMW 2000 z 0092 / 0008, SL: 21.8 mm, MD: 9.9 mm, Karnabrunn (Austria), Figs 15 K, 17 A 1 – A 2. Type locality. Karnabrunn (Austria), Korneuburg Basin. Type stratum. Korneuburg Formation Age. Early Miocene, Karpatian (late Burdigalian).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD0FFD2FF3FFC33FD909341.taxon	etymology	Etymology. Corrugatus (Latin for wrinkled, corrugated).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD0FFD2FF3FFC33FD909341.taxon	diagnosis	Diagnosis. Large, robust, moderately broad, gradate, weakly coronate shell with very prominent, trigonal varices with slightly rolled over crests and seven prominent but low spiral cords giving surface corrugated appearance.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD0FFD2FF3FFC33FD909341.taxon	description	Description. Shell large, robust, moderately broad (apical angle ~ 30 °), weakly coronate. Protoconch unknown. Early teleoconch whorls convex with, strongly raised varices, separated by slightly wider interspaces. Later teleoconch whorls gradate, nearly subcylindrical, with moderately broad subsutural shelf. Suture deeply impressed. Sculpture of moderately broad, prosocline, widely spaced, weakly trigonal, internally slightly lamellose varices, not aligned axially, separated by slightly wider interspaces (10: 12: 12). Crests of varices weakly rolled over. Interspaces with seven prominent but low, convex spiral cords separated by slightly narrower interspaces, giving surface corrugated appearance. Secondary spiral cords appear especially close to the suture. Last whorl attaining about 45 % of total height, with rounded basal angulation delimited by broad, prominent peribasal cord, relatively narrow basal disc, varices weak on base. Fasciole short, broad, with very prominent growth lines. Aperture subcircular, columella concave. Outer lip not preserved. Columellar callus forming moderately broad, adherent rim, parietal callus thin, attached. No umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD0FFD2FF3FFC33FD909341.taxon	discussion	Discussion. Some Cirsotrema species are reminiscent of Clathrus corrugatus sp. nov. but all differ in their lamellose varices and continuous peribasal cord. [e. g.: Cirsotrema crassicostatum (Deshayes (1850), from the Early Miocene of France, is slenderer, is more weakly coronate, and has more numerous and lamellose varices (see Cossmann 1912: pl. 3, figs 1 – 3). A specimen from the Early Miocene of France, illustrated by Cossmann & Peyrot (1922: pl. 4, figs 97 – 98), as Cirsotrema crassicostatum is much slenderer, has more numerous varices and has a narrower subsutural shelf. Cirsotrema subaspromontanum (Sacco, 1891), from the Tortonian of Italy, differs in its broader varices, which are overrun by spiral cords, the more superficial suture, distinctly subcylindrical whorls and markedly angled base]. Paleoenvironment. Shallow marine, lagoonal in vicinity of mangroves (Harzhauser et al. 2002). Distribution in Central Paratethys. Karpatian (Early Miocene): Korneuburg Basin: Karnabrunn (Austria) (Harzhauser 2002). Genus Claviscala de Boury, 1909	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD0FFD2FF3FFC33FD909341.taxon	type_taxon	Type species. Scalaria richardi Dautzenberg & de Boury, 1897; original designation by de Boury (1909 a: 256). Present-day, Azores.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD0FFD2FF3FFC33FD909341.taxon	diagnosis	Diagnosis. “ Fairly large size; narrow, slender shape; very long, subulate spire; concavo-convex, elevated whorls, separated by sutures bordered or even framed by two small cords; sculpture made up of slightly flexuous axial ribs, which do not extend from one suture to the next or even cease without reaching the sutural cord, but which nevertheless correspond from one whorl to the next, forming a slightly twisted pyramid around the axis; they are crossed by very fine, very regular spiral striae, which persist right up to the cords. Last whorl at most equal to one-fifth or one-sixth of the total height; base slightly excavated towards the periphery, where it is bordered by a projecting cord, while in the center it is slightly convex; the entire surface of the basal disc is decorated with spiral striations, even finer than those on the whorls of the spire, which are not latticed. Narrow opening, with an upwardly indented lip, where the basal cord ends. ” (translated from Cossmann 1912: 92 – 93).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD0FFD2FF3FFC33FD909341.taxon	discussion	Discussion. MolluscaBase eds (2024 d) currently lists twelve species under Claviscala de Boury, 1909. A comparison of these species with the type species suggests that this genus is misinterpreted by most authors. The Atlanto-Mediterranean type species and the Miocene Paratethyan Claviscala norica Harzhauser & Landau, 2014 are characterized by weakly convex whorls with a weakly impressed suture, fold-like axial ribs terminating abapically in an adsutural spiral cord and reaching adapically close to a weaker subsutural cord. Claviscala terebraloides (Kilburn, 1975) from the Indian Ocean, and the IWP-species Claviscala kuroharai Kuroda in Habe, 1961 agree well with the type, except for their slightly more convex whorls. In contrast, the Miocene Claviscala shimajiriensis MacNeil, 1961 and the extant C. dushaneae Brown, 2019, C. midwayensis Habe & Kosuge, 1970, C. nagaii Nakayama, 2000, C. nodulosa Nakayama, 2000, C. pellisanserina García, 2003, C. subulae Nakayama, 2000 and C. vivienneae García, 2003 all differ from the type species in their convex whorls, comparatively more deeply incised suture and in their prominent spiral sculpture. These species should probably be excluded from Claviscala. Another potential fossil Claviscala species was described by Hutton (1885) from the Tertiary of New Zealand as Scalaria marginata. We have not seen this species but the description by Hutton (1885: 330) does not exclude placement in this genus. In the European Neogene we are not aware of any further species, except for the Paratethyan Claviscala norica Harzhauser & Landau, 2014.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD3FFD3FF3FFA3BFB3C90EC.taxon	description	Figs 20 B – D, 21 B	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD3FFD3FF3FFA3BFB3C90EC.taxon	materials_examined	Type material. Holotype, NHMW 2014 / 0379 / 0058, SL: 23.0 mm, MD: 4.8 mm, Allerding (Austria), illustrated in Harzhauser et al. (2014: pl. 4, fig. 14), Fig. 20 B. Paratype, NHMW 2014 / 0379 / 0059, SL: 18.0 mm, MD: 4.8 mm, Allerding (Austria), illustrated in Harzhauser et al. (2014: pl. 4, fig. 15), Figs 20 C, 21 B. Illustrated material. NHMW 2014 / 0379 / 0060, SL: 27.0 mm, MD: 5.0 mm, Allerding (Austria), illustrated in Harzhauser et al. (2014: pl. 4, fig. 16), Fig. 20 D. Revised description. Medium-sized, slender shell (apical angle ~ 15 °). Protoconch unknown. Teleoconch of at least 12 weakly convex to almost straight sided whorls. Suture narrow, weakly impressed. Narrow subsutural cord, steep, poorly delimited, faintly concave subsutural ramp, periphery placed about mid-whorl. Axial sculpture of broad, weakly prosocline, vague axial ribs separated by interspaces of about equal width (?: ~ 16: ~ 16), obliquely aligned axially, weakening over subsutural ramp, most prominent along periphery, terminating in prominent spiral cord at abapical suture. Faint spiral threads. Last whorl and aperture unknown.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD3FFD3FF3FFA3BFB3C90EC.taxon	discussion	Discussion. The available specimens are based on natural casts and silicone moulds. Therefore, the delicate spiral threads, typical for the type species Claviscala richardi (Dautzenberg & de Boury, 1897), are poorly preserved. The extant northern Atlantic and Mediterranean C. richardi is very similar but differs from the Ottnangian shell in its slightly broader and larger shell and more convex whorls (see Bouchet & Warén 1986: figs 1246 – 1247). Paleoenvironment. Coastal marine, rocky shore.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD3FFD3FF3FFA3BFB3C90EC.taxon	distribution	Distribution in Central Paratethys. Ottnangian (Early Miocene): North Alpine Foreland Basin: Allerding (Austria). Genus Cylindriscala de Boury, 1909	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD3FFD3FF3FFA3BFB3C90EC.taxon	type_taxon	Type species. Scalaria fulgens de Boury, 1909 [= Cylindriscala acus (Watson, 1883)], original designation by de Boury (1909 a: 256). Present-day, Azores.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD3FFD3FF3FFA3BFB3C90EC.taxon	diagnosis	Diagnosis. “ […] tall, slender epitoniids with broad, not lamellar axial ribs and well defined basal disc; protoconch multispiral, smooth. ” (Bouchet & Warén 1986: 530).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD3FFD3FF3FFA3BFB3C90EC.taxon	discussion	Discussion. This genus displays a major radiation in the Atlantic with few additional species known from the Indo-West Pacific. All living species are deep water dwellers (Lima & Christoffersen 2012). Cylindriscala has not so far been reported from the Central Paratethys Sea, although it is represented by at least four species. Of these, especially Cylindriscala acus suggests a close relationship with the extant Atlantic fauna.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD2FFDCFF3FF957FA879245.taxon	description	Figs 20 F 1 – F 2, 21 E	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD2FFDCFF3FF957FA879245.taxon	materials_examined	Type material. Holotype, BMNH 1887.2. 9.498 (Natural History Museum London), West of Azores, https: // data. nhm. ac. uk / dataset / collection-specimens / resource / 05 ff 2255 - c 38 a- 40 c 9 - b 657 - 4 ccb 55 ab 2 feb / record / 3499770 Illustrated material. NHMWW 1866 / 0001 / 0717 a, SL: 3.3 mm, MD: 1.3 mm, Möllersdorf (Austria), Figs 20 F 1 – F 2, 21 E	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD2FFDCFF3FF957FA879245.taxon	description	Description (of Paratethyan material). Very small shell, moderately slender, turreted (apical angle ~ 20 °). Protoconch and first teleoconch whorls unknown from Paratethyan material. High conical of 2.5 weakly convex whorls in extant specimens. with prominent spiral cords at apical suture and delicate axial ribs on last whorl (see Bouchet & Warén 1986; fig. 1229). Early teleoconch of strongly and evenly convex whorls with poorly delimited, narrow subsutural ramp, periphery placed mid-whorl. Later whorls with convex to vaguely subcylindrical periphery. Suture deeply impressed. Sculpture of moderately close-set, very prominent, convex, orthocline axial ribs, separated by narrower interspaces, not aligned axially (14: 13: 11). Axial ribs most prominent along periphery, rapidly weakening on subsutural ramp. Interspaces glossy, smooth. Last whorl attaining ~ 30 % of total height, basal angulation delimited by very prominent peribasal cord, axial ribs terminate at peribasal cord, base weakly concave, smooth. Aperture subcircular, columella weakly concave. Outer lip not preserved.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD2FFDCFF3FF957FA879245.taxon	discussion	Discussion. This specimen is strikingly similar to the extant Cylindriscala acus (Watson, 1883). Watson (1883) mentioned 15 and 17 axial ribs for the penultimate and last whorls, which is more than seen in the Paratethyan shell, but at same growth stage the Paratethyan shell is indistinguishable from the extant specimens. Therefore, we tentatively treat both occurrences as conspecific despite the large stratigraphic gap of about 14.5 Ma. Cylindriscala rugata sp. nov. has a comparable shape but has fewer and broader axial ribs. Cylindriscala bellissima sp. nov. is much slenderer and has fewer but even more prominent axial ribs. Paleoenvironment. The species lives in bathyal water depths (Bouchet & Warén 1986). Similarly, Miocene occurrences in the Baden Formation of the Vienna Basin suggest middle to outer neritic settings in up to 250 m water depth (Kranner et al. 2021).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD2FFDCFF3FF957FA879245.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienna Basin: Möllersdorf (Austria) (hoc opus). Northeastern Atlantic. Present-day: Portugal, the Canaries and the Azores (Bouchet & Warén 1986).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFDDFFDDFF3FFBC6FE97976D.taxon	description	Figs 20 E, 21 C	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFDDFFDDFF3FFBC6FE97976D.taxon	materials_examined	Type material. Holotype, SMF 380174 _ 12 - 1455 a (XII 1455 a), SL: 7.7 mm, MD: 2.5 mm, CoŞteiu de Sus (Romania), specimen illustrated in Zilch (1934: pl. 10, fig. 73), Figs 20 E, 21 C. Revised description. Shell small, slender, turreted (apical angle ~ 18 °). Protoconch unknown. Teleoconch of at least eight whorls. Whorl profile strongly convex with weakly concave subsutural ramp, periphery at rounded shoulder placed distinctly above mid-whorl, convex below. Suture deeply impressed, weakly undulating. Sculpture of widely spaced, prominent, relatively narrow, convex, orthocline to weakly prosocline axial ribs, not aligned axially (16: 11: 10). Two prominent varices on last whorl. Axial ribs only slightly weakening over subsutural ramp. Interspaces wide, glossy, bearing three weak spiral cords, overrunning and slightly swollen over axial ribs. Last whorl attaining ~ 33 % of total height, basal angulation without continuous peribasal cord but delimited by row of blunt nodes formed by axial ribs; ribs very weak over slightly concave basal disc. Aperture ovate, columella weakly concave. Outer lip thickened by terminal varix. Columellar and parietal callus forming continuous, narrow rim, detached from base. Peristome duplex. Umbilicus reduced to narrow chink.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFDDFFDDFF3FFBC6FE97976D.taxon	discussion	Discussion. Cylindriscala bellissima sp. nov. is very similar in outline but has broader axial ribs and lacks the weak spiral cords. A specimen from the Hemmoorian of Winterswijk-Miste (the Netherlands) described as Turriscala (Rugatiscala) sp. by Janssen (1984: 175, pl. 50, figs 14 a – b) is highly reminiscent of Cylindriscala amandae (Boettger, 1906) and differs mainly in its more numerous axial ribs. Funiscala? miopusilla (Kautsky, 1925), from the Middle Miocene of the North Sea, has closer-set axial ribs and broader whorls with a slightly less incised suture (see Kautsky 1925: pl. 7, fig. 2). Paleoenvironment. Unknown.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFDDFFDDFF3FFBC6FE97976D.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Făget Basin: CoŞteiu de Sus (Romania) (Zilch 1934).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFDCFFDDFF3FFED7FA2693ED.taxon	description	Figs 4 C, 20 G 1 – G 2, 21 D	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFDCFFDDFF3FFED7FA2693ED.taxon	materials_examined	Type material. Holotype, NHMW 2013 / 0078 / 0155, SL: 5.6 mm, MD: 1.8 mm, Baden (Austria), Figs 4 C, 20 G 1 – G 2, 21 D. Type locality. Baden (Austria), Vienna Basin. Type stratum. Baden Formation. Age. Middle Miocene, middle Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFDCFFDDFF3FFED7FA2693ED.taxon	etymology	Etymology. Superlative of bellus (Latin for beautiful); referring to the aesthetic appearance of the species.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFDCFFDDFF3FFED7FA2693ED.taxon	diagnosis	Diagnosis. Small, slender, turreted, glossy shell with rounded shoulder, constricted below, with deeply impressed suture and very prominent, convex axial ribs, no spiral sculpture, prominent, rounded peribasal cord, aperture ovate with duplex peristome.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFDCFFDDFF3FFED7FA2693ED.taxon	description	Description. Shell small, slender, turreted (apical angle ~ 15 °). Protoconch incomplete, last two whorls turreted, weakly convex, smooth. Teleoconch of seven strongly convex whorls, with weakly convex, shallow subsutural ramp, periphery at rounded shoulder placed distinctly above mid-whorl, constricted below. Suture deeply impressed, weakly undulating. Sculpture of moderately widely spaced, very prominent, convex, orthocline to weakly prosocline axial ribs, not aligned axially (11: 10: 10), most prominent along periphery, only slightly weakening over subsutural ramp. Interspaces of about equal width to ribs, smooth, glossy. Last whorl attaining ~ 33 % of total height, three axial ribs of last whorl forming slightly more prominent varices, marked basal angulation without continuous peribasal cord but with row of blunt nodes formed by axial ribs terminating at basal angulation, base weakly concave, smooth. Aperture ovate, columella weakly concave. Outer lip strongly thickened by terminal varix. Columellar and parietal callus forming continuous, narrow rim, detached from base. Peristome duplex. Umbilicus reduced to narrow chink.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFDCFFDDFF3FFED7FA2693ED.taxon	discussion	Discussion. The extant Cylindriscala acus (Watson, 1883) differs in its greater number of axial ribs and evenly convex whorls (see Bouchet & Warén 1986: fig 1236 – 1237; Weil et al. 1999: fig. 82). Similarly, Cylindriscala aurantina (Bouchet & Warén, 1986), from the Azores, has evenly convex to somewhat subcylindrical whorls and differs in its narrower peribasal cord (see Bouchet & Warén 1986: fig 1242; Weil et al. 1999: fig. 83). Paleoenvironment. Occurrences in the Baden Formation of the Vienna Basin suggest middle to outer neritic settings in up to 250 m water depth (Kranner et al. 2021).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFDCFFDDFF3FFED7FA2693ED.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienna Basin: Baden (Austria) (hoc opus).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFDCFFDEFF3FFA57FCD39299.taxon	description	Figs 20 H 1 – H 2, 21 F	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFDCFFDEFF3FFA57FCD39299.taxon	materials_examined	Type material. Holotype, NHMW 1871 / 0010 / 0128, SL: 3.8 mm, MD: 1.4 mm, Porzteich at Břeclav (Czech Republic), Figs 20 H 1 – H 2, 21 F. Type locality. Porzteich at Břeclav (Czech Republic), Vienna Basin. Type stratum. Baden Formation. Age. Middle Miocene, middle Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFDCFFDEFF3FFA57FCD39299.taxon	etymology	Etymology. Rugatus (Latin for wrinkled), referring to the broad, fold-like axial ribs.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFDCFFDEFF3FFA57FCD39299.taxon	diagnosis	Diagnosis. Very small, slender shell with strongly convex, faintly angled whorls, deeply impressed suture, robust, convex axial ribs crossed by six broad, faint spiral cords, marked basal angulation, prominent peribasal cord.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFDCFFDEFF3FFA57FCD39299.taxon	description	Description. Shell very small, slender (apical angle ~ 18 °). Protoconch and first teleoconch whorls unknown. Teleoconch of at least eight strongly convex whorls with indistinct, convex subsutural shelf and faint shoulder angulation at periphery placed just above mid-whorl. Suture deeply impressed, linear. Sculpture of moderately close-set, robust, moderately raised, weakly prosocline, convex axial ribs, not aligned axially (11: 11: 11), adapical tips of ribs fading at suture. Interspaces between ribs slightly wider, with about six very weak, broad, low spiral cords, weakly overrunning axial ribs. Last whorl attaining about 37 % of total height, with distinct basal angulation delimited by moderately prominent peribasal cord, basal disc flattened, nearly smooth. Aperture ovate, columella weakly concave. Outer lip weakly thickened. Columellar callus forming narrow rim, detached from base, parietal callus delicate. Peristome simple. Umbilicus reduced to narrow chink.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFDCFFDEFF3FFA57FCD39299.taxon	discussion	Discussion. This species is reminiscent of Cylindriscala bellissima sp. nov. in its angled whorls but is broader, has relatively higher whorls and the axial ribs are wider but less elevated. Furthermore, C. bellissima lacks spiral sculpture. The extant Cylindriscala acus (Watson, 1883) is higher and has more numerous and narrower axial ribs (see Bouchet & Warén 1986: fig 1236 – 1237; Weil et al. 1999: fig. 82). Paleoenvironment. Unknown.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFDCFFDEFF3FFA57FCD39299.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienna Basin: Porzteich at Břeclav (Czech Republic) (hoc opus). Genus Discoscala Sacco, 1890	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFDCFFDEFF3FFA57FCD39299.taxon	type_taxon	Type species. Scalaria scaberrima Michelotti, 1840; original designation by Sacco (1890: 320). Miocene, Italy.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFDCFFDEFF3FFA57FCD39299.taxon	diagnosis	Diagnosis. “ Thin shell. Medium size; shape turriculate, conical; whorls very convex, not disjointed, but gradate with ramp; at the anterior part of each whorl, there is a depression that is limited above by a bead invisible under the following whorl; ornamentation composed of axial, straight lamellae, deviated on the posterior ramp, not corresponding from one whorl to the next, crenellated by large spiral cords. Last whorl equal to or greater than a third of the total height, with flattened base covered by a decussate disc, limited at the periphery by a crenellated rim; the lamellae cross it and extend, a little sinuously, as far as the disc; no umbilicus in the center of the disc. Aperture rounded, subangular in front, auricle resembling a slightly calloused beak; peristome barely thickened, almost discontinuous, the inner, varnished layer becoming much thinner in the parietal region. ” (translated from Cossmann 1912: 59).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFDCFFDEFF3FFA57FCD39299.taxon	discussion	Discussion. Discoscala is reminiscent of Cirsotrema, but its axial lamellae do not have a tendency to fuse, its lamellae stop abruptly at the peribasal cord and the peribasal cord delimits a well-developed basal disc that bears fine spiral threads and no axial sculpture except for growth lines.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFDFFFD9FF3FFA83FB12920C.taxon	description	Figs 15 A – B, 22 A – D	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFDFFFD9FF3FFA83FB12920C.taxon	description	D [iscoscala]. scaberrima var. subcanaliculata Sacc. — Sacco 1891: 59, pl. 2, fig. 46. D [iscoscala]. scaberrima var. taurotransiens Sacc. — Sacco 1891: 59, pl. 2, fig. 47	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFDFFFD9FF3FFA83FB12920C.taxon	materials_examined	Type material. Syntype?, BS. 030.09.001, SL: 45.0 mm, MD: 20.1 mm, S. Agata (Italy), Tortonian (Late Miocene), illustrated in Ferrero Mortara et al. (1984: 48, pl. 6, figs 3 a – b). As discussed by Ferrero Mortara et al. (1984: 48) the syntype cannot be identified clearly. Illustrated material. NHMW 1863 / 0015 / 1213, SL: 38.1 mm, MD: 18.8 mm, Möllersdorf (Austria), illustrated in Hörnes (1856, pl. pl. 6, fig. 10), Figs 22 A 1 – A 2. NHMW 2010 / 0004 / 0522 a, SL: 29.4 mm, MD: 19.5 mm, Bad Vöslau (Austria), Figs 15 A, 22 B 1 – B 2. NHMW 1859 / 0027 / 0161 a, SL: 20.9 mm, MD: 11.4 mm, Möllersdorf (Austria), Figs 15 B, 22 C 1 – C 2. NHMW 2010 / 0004 / 0522 b, SL: 22.9 mm, MD: 11.9 mm, Bad Vöslau (Austria), Figs 22 D 1 – D 2. Additional material. 2 spec., NHMW 2013 / 0078 / 0149, Baden (Austria); 2 spec., NHMW 1859 / 0027 / 0161, Möllersdorf (Austria); 1 spec., NHMW 1866 / 0040 / 0338, Lăpugiu de Sus (Romania). Revised description. Shell very large, stocky, moderately broad (apical angle 30 – 33 °). Protoconch and first teleoconch whorls unknown. Teleoconch of about ten low, strongly convex whorls, with shallow, weakly concave subsutural ramp and angled shoulder, periphery placed mid-whorl. Suture deeply impressed on early whorls, moderately impressed on later whorls. Sculpture of numerous, thin, close-set, moderately elevated, orthocline, frondose lamellae, not aligned axially (15: 22 / 29: 30) that do not amalgamate, with erect crests. Lamellar interspaces bearing numerous weak spiral cords, overrun by delicate growth lines. Last whorl attaining 40 % of total height, basal angulation strongly developed, delimited by prominent but narrow peribasal cord, lamellae stop abruptly at cord, basal disc well developed, bearing numerous fine spiral threads crossed by growth lines. Aperture subcircular with parietal concavity. Columella nearly straight. Outer lip not preserved. Columellar callus forming moderately broad rim, detached from base, parietal callus absent. Peristome simple, No umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFDFFFD9FF3FFA83FB12920C.taxon	discussion	Discussion. The assumed syntype from the Tortonian of S. Agata (Italy) differs from the specimen illustrated by Hörnes (1856) in its more pronounced, concave subsutural ramp and lamellose but distinctly less frondose varices, whereas the spire fragment agrees well with the syntype. The available Paratethyan material displays some variability in the number of lamellae and the subsutural ramp may become convex during ontogeny as seen in specimens described as variety taurotransiens by Sacco (1891). Therefore, we consider Discoscala scaberrima (Michelotti, 1840) to be somewhat variable and tentatively treat the Paratethyan specimens as conspecific with it, awaiting more material from the Proto-Mediterranean Sea to better characterize that species. The extant Discoscala edgari (de Boury, 1912) differs in its closer-set lamellae and higher subcylindrical whorls (see Weil et al. 1999: figs 249 – 250). Paleoenvironment. Occurrences in the Baden Formation of the Vienna Basin suggest middle to outer neritic settings in up to 250 m water depth (Kranner et al. 2021).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFDFFFD9FF3FFA83FB12920C.taxon	distribution	Distribution in Central Paratethys. Vienna Basin: Baden, Bad Vöslau, Möllersdorf (Austria) (Hörnes 1856; Schultz 1998); Bükk Mountains: Balaton (Hungary) (Csepreghy-Meznerics 1969 a). Proto-Mediterranean Sea. Langhian (Middle Miocene): Colli Torinesi: Sciolze, Albugnano (Italy) (Sacco 1891); Tortonian (Late Miocene): Po Basin: S. Agata, Stazzano, Montegibbio, Tortona (Italy) (Sacco 1891; Glibert 1962). North Sea. Hemmoorian (Early / Middle Miocene): North Sea Basin: Winterswijk-Miste (the Netherlands) (Janssen 1984). Reinbekian (Middle Miocene): North Sea Basin: Dingden (Germany) (Janssen 1967).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFDFFFD9FF3FFA83FB12920C.taxon	type_taxon	Type species. Scalaria (Funiscala) speyeriana Sacco, 1891 misidentified as Scalaria pusilla Philippi, 1844 in the original designation by de Boury (1891: 205); fixed by Beu (2011: 19) under ICZN Article 70.3. Oligocene, Germany.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFDFFFD9FF3FFA83FB12920C.taxon	diagnosis	Diagnosis. “ Shell rather thick. Size fairly small; shape turriculate, conical; spire long, unstaggered; whorls convex, separated by deep sutures; ornamentation composed of thick, slightly sinuous, rarely subvaricose axial ribs, crossed by spiral cords, unpunctuated. Last whorl about a quarter of the total height, with a thick, excavated disc bordered by a peripheral carina, crenellated by the ribs; its surface is furrowed and obtusely radiated by the ribs it covers; no umbilicus, but a faint bulge that simulates a rudimentary cord. Oval-rounded aperture, located in an almost vertical plane, with a split peristome, the inner layer thin and continuous, the outer layer interrupted in the parietal region; labrum bored by a thick varicose varix. ” (translated from Cossmann 1912: 86).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFDFFFD9FF3FFA83FB12920C.taxon	discussion	Discussion. De Boury (1891: 205) clearly stated Scalaria pusilla Philippi, 1844 as type species but referred to the illustration given in Speyer (1864: pl. 41, fig. 9). In the same year, but slightly later, Sacco (1891) introduced Scalaria (Funiscala) pusilla speyeriana Sacco, 1891 as name for this species along with several additional varieties, such as Funiscala pusilla sandbergeriana (Sacco, 1891) (see Sandberger 1863: pl. 11, fig. 1). De Boury (1911) discussed this problem again and stated: “ It is necessary to change the name of the type, which is characterized by the very well developed transverse cords which are not observed in the true S. pusilla. This has only a few grooves. ” (translated from de Boury 1911: 219). Cossmann (1912: 86) agreed that the type was originally misidentified and suggested that Scalaria pusilla should be placed in Bifidoscala Cossmann, 1888 (type species Scalaria lemoinei de Boury, 1883; Thanetian, France). Consequently, Cossmann (1912) listed ‘ Scalaria Speyeri Sacco’ as type species of Funiscala. As pointed out by Beu (2011), the action by de Boury (2011) conforms to ICZN Article 70.3. (misidentified type species). According to Janssen (1978) Scalaria pusilla Philippi, 1844 and Scalaria (Funiscala) pusilla speyeriana Sacco, 1891 are synonyms anyway. To complicate things, De Boury (1913 b: 72) concluded that the Oligocene species placed by him previously in Funiscala should be placed in Bifidoscala and considered Funiscala a synonym of Bifidoscala. Bifidoscala lemoinei (de Boury, 1883) as illustrated by Cossmann & Pissarro (1907: pl. 7, fig. 52 - 24) is indeed very similar to Funiscala pusilla. Nevertheless, Cossmann (1912: 66) emphasized the presence of an umbilicus in Bifidoscala and defined the genus by “ oblique axial ribs, slightly thickened at the base, invariably divided by a longitudinal groove which gives them a bifid appearance; their intervals are adorned with spiral cords which do not seem to rise up the sides of the ribs. ” (translated from Cossmann 1912: 66). Cossmann (1912) emphasized that the peculiar bifid appearance of the axial ribs was not a matter of preservation but a genuine feature and restricted the genus to a group of species, which “ existed only for a very limited time ” (Cossmann 1912: 67). Therefore, we prefer to keep Funiscala and Bifidoscala separate, awaiting a thorough revision of this enigmatic group. For species that de Boury (1913 b) thought to correspond to his original concept of Funiscala, he introduced Rugatiscala for species without punctate microsculpture [type species Scalaria levesquei de Boury, 1913, original designation by de Boury (1913 b: 72); Eocene, France]. The type species of Rugatiscala is characterized by an almost cancellate sculpture in which the ribs and cords are almost equal in strength (de Boury 1913 b; pl. 8, fig. 6). This morphology differs strongly from that observed in the Miocene species and therefore a close relationship is unlikely.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD8FFDAFF3FFB77FB8B93ED.taxon	description	Figs 21 G, 22 E	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD8FFDAFF3FFB77FB8B93ED.taxon	materials_examined	Type material. Holotype, SMF 380175 _ 12 - 1453 a (XII 1453 a), SL: 3.4 mm, MD: 1.3 mm, CoŞteiu de Sus (Romania), specimen illustrated in Zilch (1934: pl. 10, fig. 74), Figs 21 G, 22 E. Revised description. Very small, moderately slender conical shell (apical angle 23 °). Protoconch turreted of four whorls. Teleoconch of six strongly convex whorls with indistinct, convex subsutural ramp and deeply impressed, linear suture. Sculpture of relatively narrow, weakly prosocline, convex axial ribs separated by wider interspaces, vaguely aligned axially (14: 14: 14). Tips of ribs weakly pointed without touching preceding whorl. One prominent varix on last two whorls. Spiral sculpture of numerous, faint, close-set spiral threads. Last whorl convex, attaining ~ 35 % of total height, basal angulation distinct delimited by prominent, rounded peribasal cord, basal disc flattened, almost smooth. Aperture subcircular with concave columella. Columellar and parietal lip continuous, forming narrow rim. Peristome duplex. Outer lip strongly thickened.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD8FFDAFF3FFB77FB8B93ED.taxon	discussion	Discussion. This species is reminiscent of Funiscala grata (de Boury, 1891), from the Tortonian of Montegibbio (Italy), which differs in its broader axial ribs and more prominent spiral cords (de Boury 1891: pl. 4, fig. 12). Funiscala microscopica (Boettger, 1906) differs in its punctate microsculpture, character of the genus Nodiscala, and weaker basal angulation. Funiscala pusilla (Philippi, 1844), from the Rupelian of Germany, differs in its more superficial suture, broader axial ribs and prominent spiral cords in the narrower interspaces (see Janssen 1978: pl. 12, fig. 57). Paleoenvironment. Unknown.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD8FFDAFF3FFB77FB8B93ED.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Făget Basin: CoŞteiu de Sus (Romania) (Zilch 1934). Genus Gregorioiscala Cossmann, 1912	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD8FFDAFF3FFB77FB8B93ED.taxon	type_taxon	Type species. Scalaria romettensis De Gregorio, 1889; original designation by Cossmann (1912: 80). Miocene, Italy.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD8FFDAFF3FFB77FB8B93ED.taxon	diagnosis	Diagnosis. “ Very thick shell. Medium size; shape turriculate, generally convex; spire long, gradate, acute at the apex, whorls very convex in the middle; excavated posteriorly, separated by linear suture, slightly bordered below; framed by two ramps. Sculpture composed of axial ribs, quite numerous, straight, faded on the posterior ramp and not reaching the anterior suture either, crossed by very fine spiral threads, barely visible when the shell is worn. From the fourth whorl before the last, a very large varicose vein, continuous from one suture to the other, even overflowing the sutures, appears and advances - at each whorl - by an amount equal to its own width. Last whorl less than a third of the total height, ornamented like the rest of the spire, limited at the periphery of the base by a large, obtusely crenellated cord; base narrow, imperforate, only slightly convex, finely striated and vaguely striped by the varices. Circular aperture, with very thick, split peristome, the inner layer fairly narrow and cylindrical, nowhere sloping, except for a small calloused portion on the parietal region; outer layer discontinuous, varicose over the extant of the peristome. ” (translated from Cossmann 1912: 80).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFD8FFDAFF3FFB77FB8B93ED.taxon	discussion	Discussion. We are not aware of illustrations of the type species Scalaria romettensis De Gregorio, 1889, aside from the rather poor drawing of a fragmentary specimen in De Gregorio (1889: pl. 1, fig. 21). Therefore, the understanding of this genus remains ambiguous. We follow Bouchet & Warén (1986: 544) and Landau et al. (2006) who defined Gregorioiscala Cossmann, 1912 as characterized by a strong basal cord, a thickened outer lip and a thick, pitted intritacalx. The varices on subsequent whorls are aligned. Compressiscala Masahito & Habe, 1976 (type species Compressiscala japonica Masahito & Habe, 1976; present-day, Indo-West Pacific) is generally considered as subjective junior synonym (e. g. Kilburn 1985: 277; Bouchet & Warén 1986: 544), although molecular data is missing so far. Currently, MolluscaBase eds. (2024 e) lists 17 extant and two fossil species in this genus. This list, however, might need revision. For example, the extant western Atlantic Gregorioiscala pimentai Lima & Christoffersen, 2014 and Gregorioiscala federicoi Bonfitto, 2020 from the Red Sea are comparatively stout and lack prominent and protruding varices (see Lima & Christoffersen 2014: fig. 1; Bonfitto 2020: fig. 1). Similarly, the generic placement of the Recent Gregorioiscala exfoliata Bouchet & Warén, 1986, from the Bay of Biscay, and Gregorioiscala sarsi (Kobelt, 1903), from Norway, included by Bouchet & Warén (1986) in this genus, will need revision as they are not very similar to the type species. The herein described species, however, is morphologically very close to the extant Gregorioiscala pachya (Locard, 1897) and the Pliocene Gregorioiscala abrupta (De Cristofori & Jan, 1832) and fits within the generic concept of Landau et al. (2006).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFDBFFDBFF3FFA57FE82913D.taxon	description	Figs 21 H, 22 F 1 – F 2	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFDBFFDBFF3FFA57FE82913D.taxon	materials_examined	Type material. Holotype, NHMW 1862 / 0029 / 0082, SL: 9.0 mm, MD: 3.6 mm, Rudice (Czech Republic), Figs 21 H, 22 F 1 – F 2. Type locality. Rudice (Czech Republic), Carpathian Foreland Basin. Type stratum. Unknown. Age. Middle Miocene, middle Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFDBFFDBFF3FFA57FE82913D.taxon	etymology	Etymology. From ascia (Latin for ax), referring to the shape of the last whorl.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFDBFFDBFF3FFA57FE82913D.taxon	diagnosis	Diagnosis. Shell small, solid, moderately slender, turreted, of low, strongly convex whorls with broad, prominent axial ribs and last whorl with two prominent, wing-like varices placed at periphery, terminal varix with short, prominent shoulder spine.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFDBFFDBFF3FFA57FE82913D.taxon	description	Description. Shell small, solid, moderately slender, turreted (apical angle ~ 23 °). Protoconch unknown. Teleoconch of about nine relatively low whorls. Whorl profile of early whorls with moderately steep, weakly concave subsutural ramp, angled periphery placed below mid-whorl. Later teleoconch whorls more convex below suture, with convex periphery placed mid-whorl. Suture deeply impressed, weakly undulating. Sculpture of moderately widely spaced, very prominent, convex, weakly opisthocline axial ribs, about equal in width to their interspaces, partly aligned axially (9: 9: 10), most prominent along periphery, slightly weakening over subsutural ramp. Spiral sculpture of numerous subobsolete spiral cords separated by slightly narrower interspaces. Last whorl broad, attaining ~ 30 % of total height, bearing two prominent, wing-like varices placed at periphery, terminal varix with short, prominent spine, basal angulation delimited by very prominent, broad peribasal cord, axial ribs terminate at cord, base weakly concave, smooth. Aperture subcircular, columella weakly convex. Outer lip strongly thickened by terminal varix. Columellar and parietal callus forming continuous, narrow rim, detached from base. Peristome duplex. Umbilicus reduced to narrow chink. Finely pitted intritacalx typical for genus not preserved.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFDBFFDBFF3FFA57FE82913D.taxon	discussion	Discussion. The Paratethyan species is morphologically very close to the Pliocene Mediterranean Gregorioiscala abrupta (De Cristofori & Jan, 1832) from which it differs in the much larger adult size of the Pliocene species (17 mm; based on Estepona specimens fide Landau et al., 2006: 48), less angled whorls, the presence of a shoulder spine and the comparatively narrower basal disc (see Pinna & Spezia, p. 159, pl. 44, fig. 2; Landau et al. 2006: 48, pl. 14, figs 8 – 9). The extant north-eastern Atlantic Gregorioiscala pachya (Locard, 1897) is also slenderer and lacks prominent varices on the last whorl (Cossmann 1912: pl. 4, figs 1 – 2; Bouchet & Warén 1986: figs 1119 – 1121, 1254 – 1255). Paleoenvironment. Probably middle to outer neritic based on the co-occurring assemblage. (own data M. H.). This interpretation would fit to the habitat of extant species (Bouchet & Warén 1986).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFDBFFDBFF3FFA57FE82913D.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Carpathian Foreland Basin: Rudice (Czech Republic) (hoc opus). Genus Hemiacirsa de Boury, 1890	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFDBFFDBFF3FFA57FE82913D.taxon	type_taxon	Type species. Turbo lanceolatus Brocchi, 1814; original designation by de Boury (1890: 268). Pliocene, Italy.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFDBFFDBFF3FFA57FE82913D.taxon	diagnosis	Diagnosis. “ Thick shell; quite large; very narrow, clavoid shape; spire long, turriculate; whorls numerous, conjoint, almost flat or only slightly convex, separated by linear suture; ornamentation composed of axial ribs, thick, barely prominent, fairly wide apart, crossed by more or less close-set spiral grooves. Last whorl less than one-sixth of total length, subangular at periphery of base, which is mediocrely convex and simply furrowed, ribs replaced by a few irregular radiating folds; center imperforate. Aperture oval, slightly narrowed posteriorly; peristome duplex, inner layer fairly calloused and continuous, outer layer thin and discontinuous; peristome slightly oblique, very broadly bordered on the outside; columellar margin slightly excavated, thickened below, slightly sloping forward on the neck. ” (translated from Cossmann 1912: 97 – 98).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFDBFFDBFF3FFA57FE82913D.taxon	discussion	Discussion. Hemiacirsa was treated as subjective junior synonym of Acirsa Mörch, 1857 by Bouchet & Warén (1986) and MolluscaBase eds (2024 f). Herein, we resurrect Hemiacirsa based on its very slender outline, weakly impressed suture, subcylindrical to weakly convex whorls, and prominent axial ribs on early teleoconch whorls. The type species of Acirsa, A. borealis (Lyell, 1841) differs distinctly from any of the species attributed here to Hemiacirsa in its wider apical angle, more convex whorls that give the shell quite a different profile, and weaker axial sculpture. The genus experienced a major radiation during the Miocene and comprises many species, such as: Hemiacirsa duvergieri (de Boury in Cossmann, 1912 (Burdigalian, France), H. prolanceolata (Sacco, 1891) (Burdigalian, Italy), H. cristulosa (Sacco, 1891), H. convexiuscula (Sacco, 1891), H. taurolanceolata (Sacco, 1891) and H. varicosocylindrica (Sacco, 1891) (Burdigalian or Langhian, Italy), H. tauroprysmatica (Sacco, 1891) (Langhian, Italy), H. miocenica (Seguenza, 1880) (Langhian, Austria; Tortonian, Italy), H. abiaeformis sp. nov., H. drevermanni (de Boury in Cossmann, 1912), H. incusa sp. nov. and H. superelongata sp. nov. (Langhian / Badenian, Austria, Hungary, Romania), H. lanceolata (Brocchi, 1814) (Tortonian to Pliocene, Italy). Hemiacirsa became extinct during the Pleistocene (Cerulli-Irelli 1914) [note that Proto-Mediterranean Miocene species will need revision].	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFA5FFA4FF3FFF2EFDD29152.taxon	description	Figs 23 A – C, 24 A – B	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFA5FFA4FF3FFF2EFDD29152.taxon	materials_examined	Type material. Holotype, NHMW 2024 / 0281 / 0001, SL: 23.4 mm, MD: 4.9 mm, Baden-Sooss (Austria), Figs 23 A 1 – A 2, 24 A. Paratypes, NHMW 2024 / 0281 / 0002, SL: 22.0 mm, MD: 5.3 mm, Baden-Sooss (Austria), Figs 23 C 1 – C 2, 24 B. NHMW 1866 / 0015 / 1189, SL: 20.6 mm, MD: 4.6 mm, Baden (Austria), illustrated in Hörnes (1856: pl. 46, fig. 14), Figs 23 B 1 – B 2. Type locality. Baden-Sooss (Austria), Vienna Basin. Type stratum. Baden Formation. Age. Middle Miocene, middle Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFA5FFA4FF3FFF2EFDD29152.taxon	etymology	Etymology: From Picea abies (spruce), referring to the conifer-like outline.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFA5FFA4FF3FFF2EFDD29152.taxon	diagnosis	Diagnosis. Large, slender conical shell of about 13 weakly convex whorls with periphery close above abapical suture, prominent, moderately close-set axial ribs and prominent spiral cords in axial interspaces, aperture ovate with simple peristome differing from its congeners in its comparatively wide apical angle and prominent axial ribs.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFA5FFA4FF3FFF2EFDD29152.taxon	description	Description. Large, slender conical, slightly cyrtoconoidal shell (apical angle ~ 15 – 17 °). Protoconch unknown. Teleoconch of about 13 whorls. Early teleoconch whorls strongly convex with prominent, orthocline, convex axial ribs separated by narrower interspaces. Later whorls weakly convex with delicate subsutural cord, weakly concave subsutural ramp and convex periphery placed close above abapical suture. Suture superficially incised, slightly undulating. Moderately close-set, prominent, orthocline to weakly opisthocyrt axial ribs, weakening slightly over subsutural ramp, not strictly aligned axially. Axial interspaces of roughly equal width to ribs bearing numerous spiral cords separated by narrow grooves (8 / 10: 10 / 12: 10 / 11). Spiral cord at abapical suture more prominent. Spiral sculpture weak to subobsolete over ribs. About one slightly broader axial rib per whorl forming indistinct varix. Last whorl attaining ~ 27 % of total height, with distinct, rounded basal angulation delimited by one or two slightly more prominent peribasal cords, followed by up to 12 cords on flattened to weakly convex base. Aperture elongate ovate with weakly concave columella, widely angled adapically. Outer lip thin. Columellar callus forming moderately broad rim, detached from base, parietal callus forming narrow rim. Peristome simple. Umbilicus reduced to very narrow chink.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFA5FFA4FF3FFF2EFDD29152.taxon	discussion	Discussion. Hemiacirsa abiaeformis sp. nov. was confused with Hemiacirsa lanceolata (Brocchi, 1814) by Hörnes (1856), with Hemiacirsa miocenica (Seguenza, 1880) by Sieber (1958) and with Hemiacirsa prolanceolata (Sacco, 1891) by Csepreghy-Meznerics (1969 a). It differs from all these species in its wider apical angle, convex whorl periphery placed close above the abapical suture and the prominent, regular axial ribs. Hemiacirsa prolanceolata (Sacco, 1891), from the Burdigalian or Langhian of the Colli Torinese (Italy), is smaller, has sharper axial ribs and a wider aperture (see Sacco 1891: pl. 2, figs 98 a – b; Ferrero Mortara et al. 1984: pl. 6, fig. 15). Paleoenvironment. Occurrences in the Baden Formation of the Vienna Basin suggest middle to outer neritic settings in up to 250 m water depth (Kranner et al. 2021) but specimens from Szob (Hungary) might rather drive from shallow water environments.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFA5FFA4FF3FFF2EFDD29152.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienna Basin: Baden, Bad Vöslau (Austria) (Hörnes 1856); Pannonian Basin: Szob (Hungary) (Csepreghy-Meznerics 1956); Bükk Mountains: Borsodbóta (Hungary) (Csepreghy-Meznerics 1969 a).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFA4FFA6FF3FF8B0FA8692A7.taxon	description	Figs 24 C, 25 A – D	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFA4FFA6FF3FF8B0FA8692A7.taxon	materials_examined	Type material. Holotype, SMF 380167 _ 12 - 1459 (XII 1459 a), SL: 7.5 mm, MD: 2.0 mm, CoŞteiu de Sus (Romania), specimen illustrated in Zilch (1934: pl. 10, fig. 66), Fig 25 A. Illustrated material. NHMW 2018 / 0248 / 0552 a, SL: 7.7 mm, MD: 2.0 mm, CoŞteiu de Sus (Romania), Figs 24 C, 25 D 1 – D 2. NHMW 2018 / 0248 / 0552 b, SL: 7.3 mm, MD: 2.2 mm, CoŞteiu de Sus (Romania), Figs 25 B 1 – B 2. NHMW 2018 / 0248 / 0552 c, SL: 6.7 mm, MD: 1.7 mm, CoŞteiu de Sus (Romania), Figs 25 C 1 – C 2. Additional material. 10 spec., NHMW 2018 / 0248 / 0552 c, CoŞteiu de Sus (Romania). Revised description. Shell small, slender conical (apical angle ~ 15 – 18 °). Protoconch with cyrtoconoidal outline, elevated blunt nucleus (diameter 280 μm); slender, turreted spire of 4.5 smooth, weakly convex whorls, last whorl with weak angulation just below mid-whorl. Teleoconch of eight whorls, with very steep, weakly concave subsutural ramp, and weakly convex periphery placed distinctly below mid-whorl. Suture weakly impressed, linear or slightly undulating. Sculpture of moderately close-set, convex, orthocline to weakly opisthocline axial ribs, not aligned axially (10 / 12: 10 / 11: 10 / 12), separated equal width to slightly wider interspaces, bearing about eight prominent, broad spiral cords, forming more or less prominent rectangular pattern. Slightly more prominent subsutural cord and prominent spiral cord at abapical suture may occur. Last whorl attaining 35 % of total height, axial ribs terminate abruptly at weakly rounded basal angulation delimited by up to three moderate strength peribasal cords. Base weakly convex with about five weak spiral cords. Aperture elongate ovate, columella weakly convex. Outer lip and basal lip thin. Columellar callus forming narrow rim, weakly detached from base, parietal callus delicate. Peristome simple. Umbilicus reduced to narrow chink.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFA4FFA6FF3FF8B0FA8692A7.taxon	discussion	Discussion. Hemiacirsa drevermanni (de Boury in Cossmann, 1912) exhibits some variability in the number of axial ribs, strength of the spiral cords, and strength of the peribasal cords. Typical specimens are characterized by the elongate rectangular sculpture. Hemiacirsa superelongata sp. nov. develops comparable spiral sculpture but differs in its much slenderer outline and blurred axial ribs. Hemiacirsa miocenica (Seguenza, 1880) is larger and has broader and more numerous spiral cords. Paleoenvironment. Unknown.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFA4FFA6FF3FF8B0FA8692A7.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Făget Basin: CoŞteiu de Sus (Romania) (Boettger 1902); Dacian Basin: Opanec (Bulgaria) (Kojumdgieva in Kojumdgieva & Strachimirov 1960).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFA6FFA0FF3FFF2EFF60959D.taxon	description	Figs 24 D, 25 E – H	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFA6FFA0FF3FFF2EFF60959D.taxon	materials_examined	Type material. Holotype, NHMW 1863 / 0015 / 0803 a, SL: 6.3 mm, MD: 1.8 mm, Niederleis (Austria), Figs 24 D, 25 E 1 – E 2. Paratypes, NHMW 1863 / 0015 / 0803 b, SL: 7.2 mm, MD: 2.1 mm, Niederleis (Austria), Figs 25 F 1 – F 2. NHMW 1863 / 0015 / 0803 c, SL: 5.9 mm, MD: 1.9 mm, Niederleis (Austria), Figs 25 G 1 – G 2. NHMW 1863 / 0015 / 0803 d, SL: 6.5 mm, MD: 1.9 mm, Niederleis (Austria), Fig. 25 H. Additional paratypes. 4 spec., NHMW 1863 / 0015 / 0803 e, Niederleis (Austria). Type locality. Niederleis (Austria), Vienna Basin. Type stratum. Baden Formation. Age. Middle Miocene, middle Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFA6FFA0FF3FFF2EFF60959D.taxon	etymology	Etymology. After incusus, Latin for ‘ hammered out’, referring to the sculpture.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFA6FFA0FF3FFF2EFF60959D.taxon	diagnosis	Diagnosis. Small, moderately slender subulate, glossy shell with slightly cyrtoconoid outline, weakly impressed, undulating suture, weakly convex whorls, sculpture of prominent, somewhat fold-like, opisthocline axial ribs, weak to subobsolete spiral sculpture in interspaces, aperture ovate, simple.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFA6FFA0FF3FFF2EFF60959D.taxon	description	Description. Shell small, moderately slender subulate, glossy with slightly cyrtoconoid spire (apical angle ~ 25 °). Protoconch largely missing; last protoconch whorl convex, smooth. Teleoconch of up to eight weakly convex whorls, with periphery placed below mid-whorl. Suture weakly impressed, undulating. Sculpture of moderately prominent, weakly delimited, fold-like, opisthocline axial ribs, vaguely obliquely arranged axially, separated by wider interspaces (8 / 9: 8 / 9: 8), about one rib per whorl varicose. Spiral sculpture of subobsolete spiral cords, barely visible in interspaces. Last whorl attaining about 35 % of total height, with rounded basal angulation delimited by up to three weak peribasal cords, often strongly reduced, base weakly convex, almost smooth, bearing subobsolete spiral cords. Aperture relatively small, elongate ovate. Columella weakly convex. Outer lip slightly thickened by terminal varix close to thin peristome. Columellar callus forming narrow rim, detached from base. Parietal callus delicate. Umbilicus reduced to narrow chink.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFA6FFA0FF3FFF2EFF60959D.taxon	discussion	Discussion. This species could be mistaken for Hemiacirsa drevermanni (de Boury in Cossmann, 1912) but differs in its cyrtoconoid spire, subobsolete spiral sculpture and the relatively more rounded basal angulation. Hemiacirsa miocenica (Seguenza, 1880) is much larger and differs in its prominent spiral sculpture. We are not aware of any other species from the circum-Mediterranean Miocene with comparable shape and sculpture. Hemiacirsa oscari (De Boury in Cossmann, 1912) differs in its more convex whorls. Paleoenvironment. The Niederleis section, sediment and biota from coastal and lagoonal environments not exceeding 30 m water depth were transported by tempestites into offshore settings (Mandic et al. 2002).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFA6FFA0FF3FFF2EFF60959D.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienna Basin: Niederleis (Austria) (hoc opus).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFA1FFA2FF3FFB86FCD495F1.taxon	description	Figs 24 E – G, 26 A – G * S [calaria] lanceolata Brocchi Var. miocenica — Seguenza 1880: 114.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFA1FFA2FF3FFB86FCD495F1.taxon	materials_examined	Type material. Syntypes, Reggio Calabria (Italy), Tortonian (Late Miocene); the material is lost (pers com. Stefano Dominici, 03. 11. 2024). Sacco (1891) illustrated a specimen from the Tortonian of S. Agata (Italy), BS. 030.17.006, SL: 37.0 mm, MD: 7.5 mm, stored in the Museo Regionale di Scienze Naturali Torino (Italy), illustrated in Sacco (1891: pl. 2, fig. 99), Figs 26 D 1 – D 2. This specimen, however, is no type specimen. Illustrated material. NHMW 1854 / 0035 / 0311, SL: 23.7 mm, MD: 5.1 mm, Lăpugiu de Sus (Romania), Figs 26 A 1 – A 2. NHMW 1863 / 0015 / 0202, SL: 22.1 mm, MD: 4.8 mm, Lăpugiu de Sus (Romania), Figs 26 B 1 – B 2. NHMW 1865 / 0001 / 0243, SL: 23.2 mm, MD: 4.9 mm, Lăpugiu de Sus (Romania), Figs 26 C 1 – C 2. NHMW 1867 / 0019 / 0254, SL: 31.8 mm, MD: 5.4 mm, CoŞteiu de Sus (Romania), Figs 24 E, 26 E 1 – E 2. NHMW 2013 / 0078 / 0151, SL: 21.3 mm, MD: 4.0 mm, Baden-Sooss (Austria), Figs 24 F, 26 F 1 – F 2. NHMW 1846 / 0037 / 0128, SL: 18.6 mm, MD: 3.8 mm, Baden-Sooss (Austria), Figs 24 G. NHMW 1872 / 0030 / 0081, SL: 18.2 mm, MD: 3.3 mm, Baden-Sooss (Austria), Figs 26 G 1 – G 2. Revised description. Shell large, very slender (apical angle ~ 8 – 11 °). Protoconch unknown. Teleoconch of up to 15 whorls. Early teleoconch whorls moderately convex, with steep, weakly convex subsutural ramp, convex periphery placed just below mid-whorl bearing broad, prominent, slightly drop-shaped axial ribs. Later whorls high, subcylindrical, with very steep, weakly concave subsutural ramp and faintly convex periphery placed below mid-whorl. Suture weakly impressed, slightly undulating. Sculpture of moderately close-set, convex, weakly prosocline, irregular axial ribs, not aligned axially, fading over subsutural ramp (10: 10 / 11: 8 / 11), weakening on last whorl. About one slightly broader axial rib per whorl forming indistinct varix. Spiral sculpture appearing around eighth teleoconch whorl consisting of about 14 – 16 broad, flat spiral cords separated by narrower interspaces. More prominent spiral cord at abapical suture, often hidden by subsequent whorl. Spiral cords prominent in axial interspaces, only slightly weaker over ribs. Last whorl weakly convex to subcylindrical, attaining ~ 22 – 26 % of total height, with distinct, rounded basal angulation delimited by one or two more prominent peribasal cords, followed by up to 15 spiral cords over flattened base. Aperture elongate ovate with weakly concave columella, widely angled at adapically. Outer lip and basal lip thin. Columellar callus forming moderately broad rim, detached from base, parietal callus forming delicate, narrow, attached rim. Peristome simple. Umbilicus reduced to very narrow chink.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFA1FFA2FF3FFB86FCD495F1.taxon	discussion	Discussion. The shells from the Vienna Basin differ from those from Romania in the often smaller size (18 – 22 mm versus 24 – 36 mm), the usually slenderer shell and the comparatively blurred axial sculpture, which is often strongly reduced on parts of the last whorls. Provisionally, we treat both occurrences as conspecific. Hemiacirsa superelongata sp. nov. is also very slender but smaller, and has even less convex whorls, more numerous and broader axial ribs and rectangular sculpture on the subsutural ramp. Hemiacirsa taurolanceolata (Sacco, 1891), from the Burdigalian of the Colli Torinesi (Italy), differs in its straight-sided whorls and more prominent, narrower axial ribs (see Sacco 1891: pl. 2, fig. 98 bis). Hemiacirsa prolanceolata (Sacco, 1891), from the Langhian of Sciolze (Italy), is smaller and differs in its opisthocline and sharper axial ribs (see Ferrero Mortara et al. 1984: pl. 6, fig. 15). Hemiacirsa varicosocylindrica (Sacco, 1891) and Hemiacirsa cristulosa (Sacco, 1891), both from the Burdigalian or Langhian of the Colli Torinesi (Italy), differ in their even slenderer shells with more convex whorls and more prominent axial ribs (see Sacco 1904; pl. 23, fig. 28; Ferrero Mortara et al. 1984: pl. 7, fig. 4). Hemiacirsa lanceolata (Brocchi, 1814) differs in its subcylindrical whorls and weaker, often blurred axial ribs (see Pinna & Spezia 1978: pl. 60, fig. 3; Cavallo & Repetto 1992: fig. 162; Chirli 2009: pl. 16, figs 13 – 18). According to Sacco (1891), Hemiacirsa eoprysmatica (Sacco, 1891), from the Tortonian of S. Agata and Stazzano (Italy), differs from H. miocenica (Seguenza, 1880) only in its slightly wider spaced ribs on the last whorl and is probably a subjective junior synonym of H. miocaenica. However, we have not seen the syntypes of Hemiacirsa eoprysmatica, which were not illustrated by Sacco (1891) or Ferrero Mortara et al. (1984). Paleoenvironment. Occurrences in the Baden Formation of the Vienna Basin suggest middle to outer neritic settings in up to 250 m water depth (Kranner et al. 2021).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFA1FFA2FF3FFB86FCD495F1.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienna Basin: Baden-Sooss (Austria) (hoc opus). Făget Basin: CoŞteiu de Sus, Lăpugiu de Sus (Romania) (hoc opus). Proto-Mediterranean Sea. Tortonian (Late Miocene): Calabria: Reggio Calabria (Italy) (Seguenza 1880), Po Basin: S. Agata (Italy) (Sacco 1891). Pliocene records mentioned by Sacco (1891) will need verification: Po Basin: (Castelnuovo d’Asti, Zinola, Savona-Fornaci, R. Torsero (Italy).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFA3FFA2FF3FFC6AFEED91FE.taxon	description	Figs 7 H, 26 H	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFA3FFA2FF3FFC6AFEED91FE.taxon	materials_examined	Type material. Holotype: SMF 380168 _ 12 - 1458 (XII 1458 a), SL: 6.6 mm, MD: 2, 2 mm, CoŞteiu de Sus (Romania), illustrated in Zilch (1934: pl. 10, fig. 65), Figs 7 H, 26 H. Revised description. Small, moderately slender conical shell (apical angle 21 °). Protoconch and early teleoconch whorls unknown. Teleoconch whorls strongly and evenly convex. Suture deeply incised, undulating. Sculpture of very prominent, opisthocline to opisthocyrt axial ribs separated by slightly wider interspaces (12: 11: 12). Spiral sculpture weak, of indistinct spiral cords. Last whorl attaining 40 % of total height, faint basal angulation, base weakly convex, no peribasal cord, base with delicate axial ribs. Aperture ovate, Columella weakly concave. Outer lip not preserved. Basal lip thin. Columellar callus forming narrow rim. Umbilicus reduced to narrow chink.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFA3FFA2FF3FFC6AFEED91FE.taxon	discussion	Discussion. This species is placed in Hemiacirsa with some hesitation. It is squatter, with more convex whorls than most of its congeners. The species is slightly reminiscent of Cylindriscala de Boury, 1909 but lacks a well-defined basal disc. Hemiacirsa oscari is superficially reminiscent of Hemiacirsa drevermanni (de Boury in Cossmann, 1912) and Hemiacirsa abiaeformis sp. nov. in its prominent axial ribs but is less slender, lacks distinct spiral sculpture and has more convex whorls. Paleoenvironment. Unknown.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFA3FFA2FF3FFC6AFEED91FE.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Făget Basin: CoŞteiu de Sus (Romania) (Boettger 1902).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFA2FFACFF3FFF2EFA209549.taxon	materials_examined	Type material. Holotype, NHMW 2024 / 0280 / 0002, SL: 13.8 mm, MD: 2.6 mm, CoŞteiu de Sus (Romania), Figs 23 D 1 – D 2, 24 I. Paratypes, NHMW 2024 / 0280 / 0003, SL: 13.1 mm, MD: 2.8 mm, CoŞteiu de Sus (Romania), Figs 23 E 1 – E 2. NHMW 2024 / 0280 / 0004, SL: 12.4 mm, MD: 2.7 mm, CoŞteiu de Sus (Romania), Figs 23 F 1 – F 2, 24 H. Type locality. CoŞteiu de Sus (Romania), Făget Basin. Type stratum. Silt and clay of the Dej Formation. Age. Middle Miocene, Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFA2FFACFF3FFF2EFA209549.taxon	etymology	Etymology. superelongata, referring to the extremely slender and elongate outline.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFA2FFACFF3FFF2EFA209549.taxon	diagnosis	Diagnosis. Medium-sized, very slender shell with almost flat-sided whorls, superficially incised suture, broad, low, blurred axial ribs, fine spiral cords in interspaces forming vaguely cancellate / rectangular sculpture on subsutural ramp.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFA2FFACFF3FFF2EFA209549.taxon	description	Description. Shell medium-sized, very slender (apical angle ~ 10 – 11 °). Protoconch unknown. Teleoconch of up to 13 very high whorls, with very steep, weakly concave subsutural ramp and faintly convex periphery placed below mid-whorl. Suture superficial, linear. Sculpture of moderately close-set, low, convex, somewhat blurred axial ribs, not aligned axially, fading over subsutural ramp (10: 12: 12); ribs orthocline on early whorls, weakly prosocline on later whorls. Interspaces between ribs with about ten spiral cords, including cords at adapical and abapical suture, forming vaguely cancellate / rectangular sculpture especially on subsutural ramp. Last whorl attaining 28 % of total height, axial ribs fading at basal angulation delimited by with weak peribasal cord, base weakly convex bearing about five spiral cords. Aperture elongate ovate, columella weakly convex. Outer lip slightly thickened, basal lip thin, relatively narrow. Columellar callus forming narrow, attached rim, parietal callus delicate. Peristome simple. No umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFA2FFACFF3FFF2EFA209549.taxon	discussion	Discussion. This species is reminiscent of Hemiacirsa miocenica (Seguenza, 1880) but differs in its much smaller size, even slenderer outline, less convex whorls, wider basal angulation and weaker axial ribs on early teleoconch whorls. Hemiacirsa lanceolata (Brocchi, 1814) differs in its much larger size, subcylindrical whorls, marked basal angulation and comparatively more irregular, fold-like axial ribs. Hemiacirsa drevermanni (de Boury in Cossmann, 1912) is stockier, has more prominent axial ribs and broader, less numerous spiral cords. Hemiacirsa taurolanceolata (Sacco, 1891), from the Burdigalian of the Colli Torinesi (Italy), differs in its straight-sided whorls, more numerous axial ribs and wider aperture (see Sacco 1891: pl. 2, fig. 98 bis). Paleoenvironment. Unknown.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFA2FFACFF3FFF2EFA209549.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Făget Basin: CoŞteiu de Sus (Romania) (hoc opus). Genus Hirtoscala Monterosato, 1890	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFA2FFACFF3FFF2EFA209549.taxon	type_taxon	Type species. Scalaria cantrainei Weinkauff, 1866 = Hirtoscala muricata (Risso, 1826)]. Present-day, Mediterranean Sea.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFA2FFACFF3FFF2EFA209549.taxon	diagnosis	Diagnosis. “ Medium size; shape turriculate, stocky, conical or subpyramidal; spire mediocrely elongated, with convex and disjointed whorls adorned with continuous lamellae, which are thin and reflexed, strongly spiny behind, and fused from one whorl to the other over the sutures which are perforated in the lamellae intervals; no spiral ornamentation. Last whorl about one third of the total height, a little depressed at the base, which is imperforate and over which the reflexed lamellae extend, down to the center where there is only a rudimentary cord. Aperture obliquely oval, slightly narrowed posteriorly, with a peristome that is broadly reflexed and continuous, in an almost vertical plane: the inner, varnished layer, slightly thinned in the parietal region, spreads out everywhere else over the outer, discontinuous layer; labrum provided posteriorly with a pointed spine. ” (translated from Cossmann 1912: 32).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFA2FFACFF3FFF2EFA209549.taxon	discussion	Discussion. Species placed herein in Hirtoscala are reminiscent of species placed in Spiniscala de Boury, 1909 in the paleontological literature. Spiniscala, however, was already introduced by de Boury in Ivolas & Peyrot, 1900 with Scalaria trevelyana Johnston, 1841, as type species by monotypy (present-day, northeastern Atlantic). De Boury (1909 a: 257) introduced Spiniscala again as new genus with the fossil Scalaria frondicula Wood, 1848 (North Sea), as type species. Therefore, Spiniscala de Boury, 1909 is a homonym of Spiniscala de Boury in Ivolas & Peyrot, 1900. To complicate things, Ivolas & Peyrot (1900) and Peyrot (1938) referred to specimens from the Langhian of Pontlevoy (France), which were not illustrated and might not be conspecific with the extant northeastern Spiniscala trevelyana. Spiniscala trevelyana is reminiscent of species placed herein in Hirtoscala Monterosato, 1890 but lack a coronate shoulder. Cossmann (1912), referring to Spiniscala de Boury, 1909, pointed out that the differences between Spiniscala and Hirtoscala are subtle at best. Both coronate and more or less gradate but the axial lamellae were said to be fused across the suture in Hirtoscala but separated from each other in Spiniscala. In addition, Cossmann (1912) considered the axial interspaces to be smooth and glossy in Hirtoscala but dull in Spiniscala. We are not convinced of these differences and tentatively prefer to treat Spiniscala de Boury, 1909 as subjective junior synonym of Hirtoscala. Foveoscala de Boury, 1909 [type species Scalaria reflexa Carpenter, 1856, present-day, eastern Pacific] was treated as synonym of Hirtoscala by Cossmann (1912: 32). Consequently, DuShane (1974: 42) placed Scalaria reflexa Carpenter, 1856 in Hirtoscala (see Palmer 1963: 331, pl. 63, figs 5 – 6 for the holotype of Scalaria reflexa). Later, the type species of Foveoscala was placed in Lamelliscala de Boury, 1909 [type species Scalaria fasciata G. B. Sowerby II, 1844] by Weil et al. (1999: fig. 468). Thus, Foveoscala would become a subjective synonym of Lamelliscala both being published simultaneously by de Boury (1909 a). However, Lamelliscala fasciata has well defined lamellae with strongly rolled over crests (see Weil et al. 1999: fig. 147), whereas Hirtoscala / Foveoscala reflexa has flaring lamellae, which form trigonal spines along the shoulder. In addition, Kilburn (1985: 290) and Weil et al. (1999: 211) emphasized a closed umbilicus as important feature to distinguish Hirtoscala from Lamelliscala.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFADFFACFF3FFC33FD66913D.taxon	description	Figs 27 A 1 – A 2, 28 A	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFADFFACFF3FFC33FD66913D.taxon	materials_examined	Type material. Holotype, HH 42 f, Höbarth Museum, Horn (Austria), SL: 8.0 mm, MD: 3.8 mm, Achberg at Maria Dreieichen (Austria), Figs 27 A 1 – A 2, 28 A. Type locality. Achberg at Maria Dreieichen (Austria), North Alpine Foreland Basin. Type stratum. Loibersdorf Formation. Age. Early Miocene, early Eggenburgian (Burdigalian).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFADFFACFF3FFC33FD66913D.taxon	etymology	Etymology. Ceruchus (Latin for rope), referring to the rope-like appearance of the lamellae; Noun in apposition.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFADFFACFF3FFC33FD66913D.taxon	diagnosis	Diagnosis. Shell small, slender, moderately convex teleoconch whorls with periphery below mid-whorl and widely spaced, very prominent lamellae, fused across suture and obliquely aligned axially, with strongly rolled over crests, fused across suture.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFADFFACFF3FFC33FD66913D.taxon	description	Description. Shell small, solid, slender (apical angle ~ 15 °). Protoconch and first teleoconch whorls unknown. Teleoconch of moderately convex whorls with broad, steep, weakly convex subsutural shelf and convex periphery below mid-whorl. Suture deeply impressed, fenestrate. Sculpture of very prominent, widely spaced, weakly prosocline lamellae fused across suture, obliquely aligned axially (?: 8: 8), separated by wider interspaces. Crests of lamellae strongly rolled over. Interspaces between lamellae smooth. Last whorl attaining about 35 % of total height, without basal angulation or peribasal cord. Lamellae strongly rolled over on base. Fasciole very short, broad, with prominent growth lines. Aperture ovate, columella weakly concave. Columellar callus and parietal callus continuous, columellar callus forming narrow rim, attached to fasciole. Peristome thickened, duplex. No umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFADFFACFF3FFC33FD66913D.taxon	discussion	Discussion. This species co-occurs with Hirtoscala uncinata sp. nov. but differs clearly in its lamellae, which are fused across the suture and lack spines. Hirtoscala miofrondiculoides (Sacco, 1891) is similar but is much larger and differs in its wider shell and more numerous lamellae. Paleoenvironment. Shallow marine, inner neritic.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFADFFACFF3FFC33FD66913D.taxon	distribution	Distribution in Central Paratethys. Eggenburgian (Early Miocene): North Alpine Foreland Basin: Achberg at Maria Dreieichen (Austria) (Steininger et al. 1971).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFACFFAFFF3FFA07FD7F976D.taxon	description	Figs 27 D 1 – E 2, 28 B	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFACFFAFFF3FFA07FD7F976D.taxon	materials_examined	Type material. Holotype, BkK-G 340, SL: 4.2 mm, MD: 2.3 mm, Korytnica (Poland), illustrated in Bałuk (1975: pl. 20, fig. 14), Figs 27 E 1 – E 2. Paratype; NHMW 2010 / 0004 / 0525 a, SL: 6.5 mm, MD: 3.3 mm, Bad Vöslau (Austria), Figs 27 D 1 – D 2, 28 B. Type locality. Korytnica (Poland), Korytnica Basin. Type stratum. Pińczów Formation. Age. Middle Miocene, middle Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFACFFAFFF3FFA07FD7F976D.taxon	etymology	Etymology. After cornucopia, Latin for the ‘ horn of plenty’.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFACFFAFFF3FFA07FD7F976D.taxon	diagnosis	Diagnosis. Small, moderately broad, strongly coronate shell with turreted protoconch, gradate spire with nearly uncoiled whorls, robust, strongly raised, widely spaced lamellae forming prominent spines at shoulder angulation, smooth axial interspaces.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFACFFAFFF3FFA07FD7F976D.taxon	description	Description. Shell small, moderately broad (apical angle ~ 30 – 33 °), gradate coronate. Protoconch turreted of four smooth, weakly convex whorls (diameter 320 – 350 μm). Teleoconch of up to six smooth whorls, with broad subsutural shelf and convex periphery and very deeply impressed suture. Sculpture of robust, strongly raised, widely spaced, only weakly prosocline lamellae, only partly aligned axially (10: 9: 8). Crests of lamellae weakly rolled over, adapical tips forming prominent, pointed spines along shoulder. Interspaces between lamellae smooth, glossy. Last whorl attaining 45 % of total height with broad subsutural shelf, lamellae persisting over base, becoming strongly rolled over, no peribasal cord nor basal angulation, base slightly convex. Aperture subcircular, columella weakly concave. Short, broad fasciole with distinct growth lines. Outer lip thickened by prominent varix, basal lip broad. Columellar callus forming broad rim, detached from base, parietal callus delicate. Peristome duplex but inner lip covering most of peristome. No umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFACFFAFFF3FFA07FD7F976D.taxon	discussion	Discussion. Hirtoscala reflexa (Carpenter, 1856), from the eastern Pacific, is exceptionally similar but has fewer lamellae with more rolled over crests and longer spines, which may form tube-like tips (see Palmer 1963: 331, pl. 63, figs 5 – 6). Hirtoscala mitraeformis (G. B. Sowerby II, 1844), also from the modern eastern Pacific fauna, is also highly reminiscent of Hirtoscala cornucopiae sp. nov. but has more numerous teleoconch whorls and its lamellae are less rolled over (see Weil et al. 1999: fig. 467). Hirtoscala muricatocristata (Sacco, 1891), from the Pliocene of Italy, is slenderer and has much weaker shoulder spines (Sacco 1891: pl. 1, fig. 35). Hirtoscala cornucopiae sp. nov. was confused by Bałuk (1975) with Nitidiscala detracta (De Boury in Cossmann, 1912), which differs in its more sharply axially aligned lamellae, the much weaker spines and the less incised suture. Paleoenvironment. Occurrences in the Baden Formation of the Vienna Basin suggest middle to outer neritic settings in up to 250 m water depth (Kranner et al. 2021).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFACFFAFFF3FFA07FD7F976D.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Korytnica Basin: Korytnica (Poland) (Bałuk 1975); Vienna Basin: Baden (Austria) (hoc opus).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFAEFFA8FF3FFED7FE379481.taxon	materials_examined	Type material. Holotype, NHM 1862 / 0029 / 0081, SL: 8.5 mm, MD: 5.6 mm, Rudice (Czech Republic), Figs 29 B 1 – B 2. Type locality. Rudice (Czech Republic), Carpathian Foreland Basin. Type stratum. unknown. Age. Middle Miocene, middle Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFAEFFA8FF3FFED7FE379481.taxon	etymology	Etymology. Referring to its superficial similarity with some species of the terrestrial gastropod Diplommatina.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFAEFFA8FF3FFED7FE379481.taxon	diagnosis	Diagnosis. Small, very broad, stout conical, gradate, coronate shell with prominent shoulder, close-set, strongly raised, thin lamellae, partly obliquely aligned axially, distinct chink between fasciole and columellar callus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFAEFFA8FF3FFED7FE379481.taxon	description	Description. Shell small, very broad, stout conical, gradate, coronate (apical angle ~ 41 °). Protoconch and early teleoconch whorls unknown. Teleoconch of relatively low whorls, with shallow, weakly convex subsutural ramp, prominent shoulder, periphery placed above mid-whorl. Suture distinctly impressed, weakly undulating. Sculpture of close-set, strongly raised, lamellose, weakly prosocline lamellae, partly obliquely aligned axially (?: 17: 16). Lamellae forming short, trigonal spines along shoulder. Second row of weaker spines at suture, separated from shoulder spines by concavity. Interspaces between lamellae glossy, smooth. Last whorl attaining about 45 % of total height, lamellae persisting over base with strongly rolled over crests, resulting in flattish appearance, distinct basal angulation coinciding with slightly angled lamellae, no peribasal cord, base weakly concave. Short, broad fasciole with prominent growth lines. Aperture subcircular, columella concave. Outer lip thickened by varix, basal lip broad. Columellar callus forming narrow rim separated from fasciole by deep chink; parietal callus delicate. Peristome duplex. No umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFAEFFA8FF3FFED7FE379481.taxon	discussion	Discussion. We place this peculiar species in Hirtoscala based on its slightly coronate lamellae and smooth interspaces. Hirtoscala diplommatinaformis sp. nov. is reminiscent of a squat morph of Hirtoscala muricatoides (Sacco, 1891), from the Mediterranean Pliocene, and Hirtoscala pantanellii (de Boury in Cossmann, 1912), from the Mediterranean Pleistocene (see Cossmann 1912: pl. 2, fig. 26; Ferrero Mortara et al. 1984: pl. 5, fig. 3; Landau et al. pl. 8, figs 1 – 2, pl. 9, figs 1 – 6). However, all three species are somewhat untypical for Hirtoscala due to their weak basal disc. Hirtoscala muricatoides (Sacco, 1891) and Hirtoscala pantanellii (de Boury in Cossmann, 1912) were placed in Gyroscala by Cossmann (1912: 48) but lack the broad peribasal cord and the axially fused lamellae typical of that genus. Hirtoscala diplommatinaformis sp. nov. is superficially reminiscent of Tioria Maxwell, 1928 [type species Cirsotrema (Tioria) youngi Marwick, 1928; Paleocene / Eocene, New Zealand]. Tioria is characterized by its gradate shell with broad subsutural ramp and strongly rolled over crests of the lamellae on the base and a chink between fasciole and columellar callus; it lacks a peribasal cord. However, it differs from Hirtoscala diplommatinaformis sp. nov. in its spiral sculpture and the detached parietal lip. Tioria was described so far only from the Paleogene of New Zealand and was documented only by its type species [see Beu & Maxwell (1990: 96) and Beu & Raine (2009)]. Paleoenvironment. Probably middle to outer neritic based on co-occurring assemblage. (own data M. H.).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFAEFFA8FF3FFED7FE379481.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Carpathian Foreland Basin: Rudice (Czech Republic) (hoc opus).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFA9FFA9FF3FFC8CFB0E9144.taxon	description	Figs 28 C, 30 A – B	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFA9FFA9FF3FFC8CFB0E9144.taxon	materials_examined	Illustrated material. Holotype, NHMW 2022 / 0290 / 0169, Vienna / Ottakring (Austria), SL: 6.2 mm, MD: 2.3 mm, Figs 30 B 1 – B 2. Paratype, NHMW 2024 / 0278 / 0003, SL: 7.4 mm, MD: 2.8 mm, Bad Vöslau (Austria), Figs 28 C, 30 A 1 – A 2. Type locality. Vienna / Ottakring (Austria), Vienna Basin. Type stratum. Baden Formation. Age. Middle Miocene, Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFA9FFA9FF3FFC8CFB0E9144.taxon	etymology	Etymology. Hastatus (Latin for armed with a spear), referring to the slender outline.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFA9FFA9FF3FFC8CFB0E9144.taxon	diagnosis	Diagnosis Small, moderately slender, turriculate, coronate shell with narrow subsutural shelf, moderately close-set, robust, strongly raised, weakly prosocline lamellae with low, trigonal spines at shoulder, smooth interspaces.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFA9FFA9FF3FFC8CFB0E9144.taxon	description	Description. Shell small, moderately slender, turriculate (apical angle ~ 20 – 22 °), coronate. Protoconch and early teleoconch whorls unknown. Teleoconch of at least eight whorls, with narrow subsutural shelf and weakly convex to subcylindrical periphery. Suture deeply impressed. Sculpture of moderately close-set, robust, strongly raised, weakly prosocline lamellae, aligned axially (13: 13: 14), adapical tips forming low, trigonal spines along shoulder. Crests of lamellae slightly rolled over. Interspaces between lamellae smooth, glossy. Last whorl attaining about 41 % of total height, lamellae persisting over base, becoming strongly rolled over, flat and broad, no peribasal cord nor basal angulation but lamellae faintly angled at transition into flattish base. Very short, broad fasciole with prominent growth lines. Aperture subcircular, columella weakly concave. Outer lip thickened by prominent varix. Columellar callus forming broad rim, detached from base, parietal callus delicate. Peristome duplex but inner layer covering outer layer largely. No umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFA9FFA9FF3FFC8CFB0E9144.taxon	discussion	Discussion. Hirtoscala miofrondiculoides (Sacco, 1891) differs in its fewer and broader varices, which are distinctly fused over the suture. The extant eastern Atlantic Spiniscala trevelyana (Johnston, 1841) is reminiscent of the Paratethyan species but differs in its wider shell and has an open umbilicus (Weil et al. 1999: fig. 92). ‘ Epitonium trevelyanum’ from the Serravallian of the Karaman Basin (Turkey), is also a Hirtoscala but lacks a coronate shoulder (see Landau et al. pl. 20, fig. 10). Hirtoscala spinosa (Sacco, 1891), from the Tortonian of S. Agata (Italy) might be closely related but differs in in having fewer lamellae (see Sacco 1891: pl. 1, fig 38). Hirtoscala hastata sp. nov. is exceptionally similar to the extant Hirtoscala tenebrosa (G. B. Sowerby II, 1903) from South Africa, which differs mainly in its slightly broader and more convex whorls and deeper suture (see Kilburn 1985: figs 94 – 95; Weil et al. 1999: fig. 142). The Pliocene Mediterranean Hirtoscala spinifera (Seguenza, 1876) differs in its fewer, much broader lamellae (see Landau et al. 2006: pl. 9, figs 11 – 16, Chirli 2009: pl. 14, figs 1 – 7). Parviscala lacinia sp. nov. and Parviscala paratethyca sp. nov. are superficially similar but differ in the presence of spiral sculpture in the axial interspaces and have fewer lamellae. Moreover, H. hastata has a narrower subsutural ramp with weaker spines compared to Parviscala lacinia sp. nov.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFA9FFA9FF3FFC8CFB0E9144.taxon	description	Paleoenvironment. Occurrences in the Baden Formation of the Vienna Basin suggest middle to outer neritic settings in up to 250 m water depth (Kranner et al. 2021).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFA9FFA9FF3FFC8CFB0E9144.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Korytnica Basin: Jawor at Korytnica (Poland) (Friedberg 1938); Vienna Basin: Bad Vöslau, Vienna / Ottakring (Austria) (hoc opus).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFABFFAAFF3FFF2EFB87903D.taxon	description	Figs 27 C 1 – C 2, 28 D	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFABFFAAFF3FFF2EFB87903D.taxon	materials_examined	Type material. Holotype, NHMW 1851 / 0002 / 0007, SL: 5.8 mm, MD: 2.9 mm, Grund (Austria), Figs 27 C 1 – C 2, 28 D. Type locality. Grund (Austria), North Alpine-Carpathian Foreland Basin. Type stratum. Grund Formation. Age. Middle Miocene, early Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFABFFAAFF3FFF2EFB87903D.taxon	etymology	Etymology. In honor of Katarína Holcová (Charles University, Prague), in recognition of her contributions on Paratethyan oceanography.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFABFFAAFF3FFF2EFB87903D.taxon	diagnosis	Diagnosis. Small, moderately broad, weakly coronate shell with subcylindrical whorls, gradate spire with deeply impressed suture, robust, strongly raised, moderately widely spaced lamellae occasionally forming low spines at shoulder angulation, smooth axial interspaces.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFABFFAAFF3FFF2EFB87903D.taxon	description	Description. Shell small, moderately broad (apical angle ~ 28 °), gradate, weakly coronate. Protoconch unknown. Teleoconch of at least six whorls, with moderately broad subsutural shelf and weakly convex to subcylindrical periphery. Suture deeply impressed. Sculpture of robust, strongly raised, moderately widely spaced, weakly prosocline lamellae, aligned axially (9: 10: 11). Crests of lamellae weakly rolled over, adapical tips occasionally forming low, vaguely trigonal spines along shoulder. Other lamellae with slightly flaring convex margins along shoulder. Interspaces between lamellae smooth, glossy. Last whorl subcylindrical, attaining about 45 % of total height, lamellae very prominent over base, becoming strongly rolled over, no peribasal cord nor basal angulation. Base slightly convex. Aperture subcircular, columella weakly concave. Short, broad fasciole with distinct growth lines. Outer lip thickened by prominent varix with small shoulder spine, basal lip broad. Columellar callus forming broad rim, detached from base, parietal callus delicate. Peristome duplex but inner lip covering most of peristome. No umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFABFFAAFF3FFF2EFB87903D.taxon	discussion	Discussion. The extant Mediterranean Hirtoscala muricata (Risso, 1826) [= H. cantrainei (Weinkauff, 1866)] might be closely related but is distinguished by its wider apical angle, more numerous and rather orthocline varices and the wider base (Weil et al. 1999: fig. 98; Chirli 2009: pl. 12, figs 10 – 17). Hirtoscala spinosa (Sacco, 1891), from the Tortonian of S. Agata (Italy), has a much narrower subsutural shelf and more numerous, less prosocline lamellae (see Sacco 1891: pl. 1, fig. 38). Hirtoscala perlongata (Sacco, 1891), from the Pliocene of Italy, differs in its wider subsutural ramp and more numerous varices (see Ferrero-Mortara et al. 1984: pl. 4, fig. 14). Hirtoscala cornucopiae sp. nov. is comparable in size and general outline but differs in the wider subsutural shelf, more prominent spines and fewer varices. Parviscala lacinia sp. nov. is also reminiscent of this species but differs in the presence of spiral sculpture. Paleoenvironment. At the locality Grund channels formed in middle to outer neritic environments and were filled with allochthonous assemblages uniting coastal-mudflat faunas with inner neritic ones (Zuschin et al. 2005; Roetzel 2009). The preservation suggests that the species was part of assemblages from middle to outer neritic environments.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFABFFAAFF3FFF2EFB87903D.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): North-Alpine Carpathian Foreland Basin: Grund (Austria (hoc opus); Pannonian Basin: Várpalota (Hungary) (Strausz 1954).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFABFFABFF3FF924FEB890CA.taxon	description	Figs 28 E, 30 D – F	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFABFFABFF3FF924FEB890CA.taxon	materials_examined	Type material. Holotype, NHMW 2024 / 0282 / 0001, SL: 9.3 mm, MD: 3.7 mm, Steinebrunn (Austria), illustrated in Hörnes (1856: pl. 46, figs 8 a – d), Figs 28 E, 30 D 1 – D 2. Illustrated material. NHMW 2024 / 0278 / 0004, SL: 9.3 mm, MD: 4.1 mm, Bad Vöslau (Austria), Figs 30 E 1 – E 2. NHMW 2024 / 0278 / 0005, SL: 4.7 mm, MD: 2.2 mm, Bad Vöslau (Austria), Figs, 30 F. Revised description. Shell small, moderately slender (apical angle 24 °), faintly coronate. Protoconch and early teleoconch whorls unknown. Teleoconch of at least eight convex whorls, with periphery placed below mid-whorl. Narrow, weakly concave subsutural shelf. Suture moderately impressed. Sculpture of moderately closely set, robust, strongly raised, weakly prosocline lamellae, obliquely aligned axially (11: 11: 11), adapical tips slightly pointed close to suture. Crests of lamellae strongly rolled over. Interspaces between lamellae smooth. Last whorl attaining about 40 % of total height, lamellae persisting over base, strongly rolled over, no peribasal cord nor basal angulation, base slightly convex. Fasciole very short, broad, with prominent growth lines. Aperture ovate, columella weakly concave. Outer lip thickened by varix, basal lip broad. Columellar callus forming moderately broad rim, slightly detached from base, parietal callus delicate. Peristome duplex but inner layer covers outer layer largely. No umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFABFFABFF3FF924FEB890CA.taxon	description	Synonyms. Clathrus parilis De Boury in Cossmann, 1912, was introduced by Cossmann (1912) from CoŞteiu de Sus (Romania). The holotype, illustrated in Cossmann (1912: pl. 5, fig. 5) does not differ from Hirtoscala miofrondiculoides (Sacco, 1891). No good illustration of this species, however, was available to Cossmann (1912) at that time.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFABFFABFF3FF924FEB890CA.taxon	discussion	Discussion. The holotype was mixed by Hörnes (1856) with specimens described herein as Parviscala lacinia sp. nov. Whilst the latter is quite frequent at Steinebrunn (Austria) with more than 50 specimens, Hirtoscala miofrondiculoides is very rare. They are clearly distinguished by the gradate spire and weak spiral cords of P. lacinia. Hirtoscala hastata sp. nov. differs from H. miofrondiculoides in its subcylindrical whorls and slightly wider subsutural shelf. The Pliocene Mediterranean Hirtoscala proxima (de Boury, 1890) might be a closely related species but is slenderer and has a slightly deeper incised suture (see de Boury 1890, pl. 4, fig. 9; Landau et al. 2006: pl. 11, figs 1 – 2). Scalaria clathrata sensu Eichwald (1852: pl. 10, fig. 20) differs in its strongly convex whorls and its varices are not fused across the suture. The status of this species from Ukraine remains unclear. Scalaria spinosa Bonelli refers to a name used in an unpublished catalogue for the collection in Torino (Italy) written by Franco Andrea Bonelli (1784 – 1830) around the year 1826. The name was made available by Sacco (1891: 26, pl. 1, fig. 38), who illustrated a specimen from the Tortonian of S. Agata (Italy). Hirtoscala spinosa (Sacco, 1891) differs from H. miofrondiculoides in its slightly gradate spire and subcylindrical whorls. Paleoenvironment. The occurrence at Steinebrunn (Austria) suggests shallow marine environments with sea grass (own data M. H.).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFABFFABFF3FF924FEB890CA.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Korytnica Basin: Korytnica (Poland) (Bałuk 1975); Vienna Basin: Baden, Bad Vöslau, Steinebrunn (Austria) (Hörnes 1856); Făget Basin: CoŞteiu de Sus (Romania) (Cossmann 1912). Proto-Mediterranean Sea. Serravallian (Middle Miocene): Karaman Basin: Seyithasan (Turkey) (Landau et al. 2013).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFAAFFB4FF3FF8B2FBC3935A.taxon	description	Figs 28 F, 30 C 1 – C 2	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFAAFFB4FF3FF8B2FBC3935A.taxon	materials_examined	Type material. Holotype, NHMW 1867 / 0019 / 0250, SL: 14.9 mm, MD: 5.0 mm, CoŞteiu de Sus (Romania), Figs 28 F, 30 C 1 – C 2. Type locality. CoŞteiu de Sus (Romania), Făget Basin. Type stratum. Silt and clay of the Dej Formation. Age. Middle Miocene, Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFAAFFB4FF3FF8B2FBC3935A.taxon	etymology	Etymology. After the ancient lighthouse Pharos, referring to the tower-like shape; noun in apposition.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFAAFFB4FF3FF8B2FBC3935A.taxon	diagnosis	Diagnosis. Medium sized, moderately slender, turreted shell with moderately impressed suture, moderately close-set, moderately raised, weakly prosocline lamellae, smooth, glossy interspaces, low last whorl.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFAAFFB4FF3FF8B2FBC3935A.taxon	description	Description. Shell medium sized, moderately slender (apical angle ~ 22 °). Protoconch and early teleoconch whorls unknown. Teleoconch of at least 12 convex whorls, with periphery just above mid-whorl. Suture moderately impressed, linear. Sculpture of moderately close-set, moderately raised, weakly prosocline lamellae, not aligned axially (10: 12: 13), adapical tips faintly alate at suture. Crests of lamellae forming weak subsutural concavity, weakly rolled over. Interspaces between lamellae smooth, glossy. Last whorl attaining about 30 % of total height, with weakly convex subsutural ramp, strongly convex periphery, lamellae strongly rolled over on base, no peribasal cord, base slightly sub-angled, weakly convex. Aperture ovate, columella concave. Columellar callus forming narrow rim, detached from base, parietal callus delicate. Peristome weakly thickened, duplex. Narrow umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFAAFFB4FF3FF8B2FBC3935A.taxon	discussion	Discussion. This species is reminiscent of Hirtoscala miofrondiculoides (Sacco, 1891) but differs in its larger size, slenderer shape, more numerous and lower teleoconch whorls and the less rolled over and therefore seemingly narrower varices. Hirtoscala hastata sp. nov. differs in it the presence of a narrow subsutural shelf and in its coronate shoulder. The Pliocene Mediterranean Hirtoscala proxima (de Boury, 1890) might be a closely related species and is a look-alike of the Paratethys species concerning size, shape and general sculpture but differs in its aligned varices (see de Boury 1890, pl. 4, fig. 9; Ferrero Mortara et al. 1984: pl. 4, fig. 16; Landau et al. 2006: pl. 11, figs 1 – 2). This species was confused by previous authors with the extant Fuscoscala tenuicostata (Michaud, 1829) [= Fuscoscala turtonis (Turton, 1819)], which is larger and differs in its striate axial interspaces and has broader lamellae, which are slightly alate at the adapical suture (see Weil et al. 1999: fig. 97; Landau et al. 2006: pl. 12, fig. 3). Paleoenvironment. Unknown.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFAAFFB4FF3FF8B2FBC3935A.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Southern Pannonian Basin: Miljevići (Bosnia and Herzegovina) (Atanacković 1985). Dacian Basin: Dobrusha, Opanec (Bulgaria) (Kojumdgieva in Kojumdgieva & Strachimirov 1960); Făget Basin: CoŞteiu de Sus (Romania) (hoc opus).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB5FFB5FF3FFAC2FD669469.taxon	description	Figs 27 B, 28 G	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB5FFB5FF3FFAC2FD669469.taxon	materials_examined	Type material. Holotype, HH 42 g, Höbarth Museum, Horn (Austria), SL: 6.2 mm, MD: 3.2 mm, Achberg at Maria Dreieichen (Austria), Figs 27 B 1 – B 2, 28 G. Type locality. Achberg at Maria Dreieichen (Austria), North Alpine Foreland Basin. Type stratum. Loibersdorf Formation. Age. Early Miocene, early Eggenburgian (Burdigalian).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB5FFB5FF3FFAC2FD669469.taxon	etymology	Etymology. In honor of Fritz F. Steininger, (Eggenburg, Austria), in recognition of his tremendous impact on Paratethys research.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB5FFB5FF3FFAC2FD669469.taxon	diagnosis	Diagnosis. Small, solid, moderately slender shell with moderately convex whorls and widely spaced, prominent lamellae with strongly rolled over crests and weak spines along rounded shoulder.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB5FFB5FF3FFAC2FD669469.taxon	description	Description. Shell small, solid, moderately slender (apical angle ~ 20 °), weakly coronate. Protoconch and first teleoconch whorls unknown. Teleoconch of moderately convex whorls with rounded shoulder, periphery placed just above mid-whorl. Suture deeply impressed, fenestrate. Sculpture of very prominent, widely spaced, weakly prosocline lamellae, separated by wider interspaces, obliquely aligned axially (?: 8: 8), forming short, indistinct spines along shoulder. Crests of lamellae fragile, strongly rolled over. Interspaces between lamellae smooth. Last whorl attaining about 40 % of total height, without basal angulation or peribasal cord, lamellae strongly rolled over on base. Fasciole very short, broad, with prominent growth lines. Aperture ovate, columella weakly concave. Columellar callus and parietal callus continuous, columellar callus forming narrow rim, attached to fasciole. Peristome thickened, duplex. No umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB5FFB5FF3FFAC2FD669469.taxon	discussion	Discussion. Hirtoscala steiningeri sp. nov. co-occurs with Hirtoscala ceruchus sp. nov. and Hirtoscala uncinata sp. nov., which all fall in the same size class. Hirtoscala uncinata differs clearly in its strongly coronate shoulder whereas H. ceruchus differs in its uniformly broad lamellae, which lack spines and the low position of the periphery. The Pliocene Mediterranean Hirtoscala spinifera (Seguenza, 1876) might be a closely related species but differs in its much larger size and broader shell (see Landau et al. 2006: pl. 9, figs 11 – 16, Chirli 2009: pl. 14, figs 1 – 7).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB5FFB5FF3FFAC2FD669469.taxon	distribution	Distribution in Central Paratethys. Eggenburgian (Early Miocene): North Alpine Foreland Basin: Achberg at Maria Dreieichen (Austria) (Steininger et al. 1971).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB4FFB6FF3FFDD3FD6696A5.taxon	description	Figs 28 H, 31 A – F	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB4FFB6FF3FFDD3FD6696A5.taxon	materials_examined	Type material. Holotype, HH 42, Höbarth Museum, Horn (Austria), SL: 6.9 mm, MD: 3.4 mm, Achberg at Maria Dreieichen (Austria), Figs 28 H, 31 D 1 – D 2. Paratypes, HH 42 a, Höbarth Museum, Horn (Austria), SL: 8.0 mm, MD: 4.6 mm, Achberg at Maria Dreieichen (Austria), Figs 31 A 1 – A 2. HH 42 b, Höbarth Museum, Horn (Austria), SL: 8.1 mm, MD: 3.2 mm, Achberg at Maria Dreieichen (Austria), Figs 31 B 1 – B 2. HH 42 c, Höbarth Museum, Horn (Austria), SL: 8.0 mm, MD: 3.8 mm, Achberg at Maria Dreieichen (Austria), Figs 31 C 1 – C 2. HH 42 d, Höbarth Museum, Horn (Austria), SL: 7.3 mm, MD: 3.5 mm, Achberg at Maria Dreieichen (Austria), Fig. 31 E. HH 42 e, Höbarth Museum, Horn (Austria), SL: 8.6 mm, MD: 4.0 mm, Achberg at Maria Dreieichen (Austria), Fig. 31 F. Type locality. Achberg at Maria Dreieichen (Austria), North Alpine Foreland Basin. Type stratum. Loibersdorf Formation. Age. Early Miocene, early Eggenburgian (Burdigalian).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB4FFB6FF3FFDD3FD6696A5.taxon	etymology	Etymology. Uncinatus (Latin for hook-like), referring to the axial sculpture.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB4FFB6FF3FFDD3FD6696A5.taxon	diagnosis	Diagnosis. Small, solid, moderately slender, gradate and strongly coronate shell with very prominent, broad, strongly rolled over lamellae forming long, solid spines pointed in adapical direction.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB4FFB6FF3FFDD3FD6696A5.taxon	description	Description. Shell small, solid, moderately slender (apical angle ~ 20 – 25 °), gradate, strongly coronate. Protoconch and first teleoconch whorls unknown. Teleoconch of weakly convex, subcylindrical whorls with broad, concave subsutural shelf and angled shoulder, with periphery placed mid-whorl. Suture deeply impressed, largely obscured by lamellae. Sculpture of very prominent, moderately close-set, only weakly prosocline lamellae, separated by slightly wider interspaces, not aligned axially (?: ~ 10: ~ 11 / 14), forming very prominent, adapically pointed spines along shoulder. Crests of lamellae strongly rolled over. Interspaces between lamellae smooth. Last whorl attaining about 45 % of total height, without basal angulation nor peribasal cord, lamellae strongly rolled over on base. Fasciole very short, broad, with prominent growth lines. Aperture ovate, columella weakly concave. Columellar and parietal callus continuous, columellar callus forming narrow rim, attached to fasciole. Peristome thickened, duplex. No umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB4FFB6FF3FFDD3FD6696A5.taxon	discussion	Discussion. Hirtoscala uncinata sp. nov. is characterized by its extraordinarily prominent lamellae with long, solid spines. This species was confused by Steininger et al. (1973) with the coeval Hirtoscala merignacensis (de Boury in Cossmann, 1912) from the Burdigalian of the Aquitaine Basin in France, which differs in its broader conical outline and fewer lamellae (see Cossmann 1912: pl. 1, figs 30 – 31; Cossmann & Peyrot 1922: pl. 3, figs 39 – 42). Hirtoscala benoisti de Boury, 1912, from the Burdigalian of the Aquitaine Basin, has even fewer lamellae than H. merignacensis and differs from the Paratethyan species in its slightly inflated last whorl (Cossmann 1912: pl. 1, figs 33 – 34; Cossmann & Peyrot 1922: pl. 3, figs 69 – 70). Hirtoscala hastata sp. nov. has more numerous lamellae and less prominent spines. The extant Hirtoscala venosum (G. B. Sowerby II, 1844), from the western Atlantic is comparable in its strongly coronate lamellae and gradate spire but differs in its broader shell (Weil et al. 1999: fig. 28). Paleoenvironment. Shallow marine, inner neritic.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB4FFB6FF3FFDD3FD6696A5.taxon	distribution	Distribution in Central Paratethys. Eggenburgian (Early Miocene): North Alpine Foreland Basin: Achberg at Maria Dreieichen (Austria) (Steininger et al. 1971).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB6FFB7FF3FFCFBFA249101.taxon	description	Figs 28 I, 32 A	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB6FFB7FF3FFCFBFA249101.taxon	materials_examined	Type material. Holotype, NHMW 2007 / 0086 / 0165 c, SL: 8.6 mm, MD: 3.9 mm, Baden (Austria), Figs 28 I, 32 A 1 – A 2. Type locality. Baden (Austria), Vienna Basin. Type stratum. Baden Formation. Age. Middle Miocene, middle Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB6FFB7FF3FFCFBFA249101.taxon	etymology	Etymology. Occidentalis (Latin for western).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB6FFB7FF3FFCFBFA249101.taxon	diagnosis	Diagnosis. Small, moderately slender conical shell with strongly convex, slightly angled whorls, deeply impressed suture and somewhat inflated last whorl, with numerous delicate, slightly raised, sigmoidal lamellae with smooth interspaces, no peribasal cord.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB6FFB7FF3FFCFBFA249101.taxon	description	Description. Shell small, moderately slender conical (apical angle ~ 26 °). Protoconch and first teleoconch whorls unknown. Early teleoconch whorls convex, with moderately steep, weakly concave subsutural ramp, slightly angled below, periphery placed just below abapical suture. Later whorls strongly increasing in width with more prominent angulation mid-whorl. Suture deeply impressed, linear. Sculpture of delicate, slightly raised, close-set, slightly sigmoidal lamellae, not aligned axially (~ 25: ~ 38: ~ 34), occasionally amalgamating into weak varices, tips of lamellae slightly bent in apertural direction, faintly alate, touching preceding whorl. Crests of lamellae slightly rolled over. Interspaces smooth, increasing slightly in width on later whorls. Last whorl attaining ~ 42 % of total height, weak basal angulation without peribasal cord, lamellae persisting over weakly convex base, strongly rolled over. Aperture ovate, columella weakly convex. Outer lip not thickened. Columellar callus forming narrow, attached rim. Parietal callus delicate. Peristome simple. No umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB6FFB7FF3FFCFBFA249101.taxon	discussion	Discussion. This species could be confused with Subuliscala breitenbergeri sp. nov., which is similar in shape and axial sculpture, but differs in the presence of spiral cords in the latter. The extant Hyaloscala amathusium (Melvill & Standen, 1903), from the Indian Ocean, has a broader conical shell with fewer lamellae (see Weil et al. 1999: fig. 202). The Atlanto-Mediterranean Hyaloscala clathratulum (Kanmacher, 1798) has more prominent lamellae and less convex whorls without angulation. Paleoenvironment. Occurrences in the Baden Formation of the Vienna Basin suggest middle to outer neritic settings in up to 250 m water depth (Kranner et al. 2021).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB6FFB7FF3FFCFBFA249101.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienna Basin: Baden (Austria) (hoc opus).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB1FFB1FF3FFA59FF60959D.taxon	description	Figs 28 J, 32 B 1 – B 2	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB1FFB1FF3FFA59FF60959D.taxon	materials_examined	Type material. Holotype, NHMW 2024 / 0278 / 0006, SL: 5.2 mm, MD: 2.3 mm, Bad Vöslau (Austria), Figs 28 J, 32 B 1 – B 2. Type locality. Bad Vöslau (Austria), Vienna Basin. Type stratum. Baden Formation. Age. Middle Miocene, middle Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB1FFB1FF3FFA59FF60959D.taxon	etymology	Etymology. In honor of Bret Raines (Victorville, California, USA), in recognition of his contributions in Epitoniidae systematics.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB1FFB1FF3FFA59FF60959D.taxon	diagnosis	Diagnosis. Small, moderately broad, stocky conical, weakly gradate, faintly coronate, with narrow subsutural ramp, weak shoulder angulation, moderately close-set, moderately raised lamellae, obliquely aligned axially, sub-angled base, subcircular aperture.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB1FFB1FF3FFA59FF60959D.taxon	description	Description. Shell small, translucent, moderately broad, stocky conical (apical angle ~ 32 °), weakly gradate, faintly coronate. Protoconch and early teleoconch whorls unknown. Teleoconch of at least seven whorls. Early whorls with slight mid-whorl angulation, later whorls with narrow, weakly concave subsutural ramp and convex periphery placed mid-whorl. Suture deeply impressed. Sculpture of moderately close-set, moderately raised, only weakly prosocline lamellae, obliquely aligned axially (12: 16: 16), forming low, trigonal spines along shoulder, fused across suture. Crests of lamellae only weakly rolled over. Interspaces between lamellae smooth, glossy. Last whorl attaining 45 % of total height, faintly coronate, lamellae persisting over base, strongly rolled over, base sub-angled, no peribasal cord, flattened. Aperture subcircular, columella weakly concave. Very short, broad fasciole with distinct growth lines. Columellar callus forming narrow rim, attached to fasciole, parietal callus delicate. Peristome simple, slightly flaring but not thickened. No umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB1FFB1FF3FFA59FF60959D.taxon	discussion	Discussion. The extant Hyaloscala opposita (de Boury, 1921), from South Africa, is exceptionally similar and differs only in its slightly weaker shoulder angulation and more convex base. (for holotype see https: // science. mnhn. fr / institution / mnhn / collection / im / item / 2000 - 4366). This African species was placed in Sodaliscala by de Boury (1921), but Sodaliscala has spiral cords in the interspaces between the lamellae (Kilburn 1985: 312). Consequently, the species was transferred to Hyaloscala by Kilburn (1985: 293) and Weil et al. (1999: 54). The Atlanto-Mediterranean Hyaloscala clathratula Kanmacher, 1798 has a much weaker shoulder angulation (see MolluscaBase eds 2924 i). Hyaloscala miotrinacria (Sacco, 1891), from the Late Miocene of Tetti Borelli (Italy), differs in its greater number of lamellae and broader early teleoconch whorls (see Sacco 1891: pl. 1, fig. 42; Ferrero Mortara et al. 1984: pl. 3, fig. 13). Hyaloscala micans Cossmann & Peyrot, 1922, from the Burdigalian of France, is comparably stout with broader spire whorls and larger last whorl (see Cossmann & Peyrot 1922: pl. 4, figs 86 – 87). Hyaloscala exilis de Boury, 1912, from the Burdigalian of France, (Cossmann & Peyrot 1922: pl. 4, fig, 25), is much slenderer and has more numerous, delicate lamellae. Paleoenvironment. Occurrences in the Baden Formation of the Vienna Basin suggest middle to outer neritic settings in up to 250 m water depth (Kranner et al. 2021).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB1FFB1FF3FFA59FF60959D.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienna Basin: Bad Vöslau (Austria) (hoc opus).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB0FFB2FF3FFB87FF609469.taxon	description	Figs 28 L, 32 D – F	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB0FFB2FF3FFB87FF609469.taxon	materials_examined	Type material. Holotype, NHMW 2024 / 0278 / 0007, SL: 6.8 mm, MD: 3.2 mm, Bad Vöslau (Austria), Figs 28 L, 32 D 1 – D 2. Paratypes, NHMW 2024 / 0278 / 0008, SL: 4.8 mm, MD: 2.4 mm, Bad Vöslau (Austria), Figs 32 E 1 – E 2. NHMW 2024 / 0278 / 0009, SL: 4.8 mm, MD: 2.7 mm, Bad Vöslau (Austria), Fig. 32 F. Type locality. Bad Vöslau (Austria), Vienna Basin. Type stratum. Baden Formation. Age. Middle Miocene, middle Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB0FFB2FF3FFB87FF609469.taxon	etymology	Etymology. Rusticus (Latin for rustic), referring to the crude sculpture.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB0FFB2FF3FFB87FF609469.taxon	diagnosis	Diagnosis. Small, moderately broad conical shell with strongly convex whorls, slightly inflated last whorl and prominent axial sculpture of broadly rolled over lamellae, obliquely aligned axially, fused across suture, sub-angled base, ovate aperture.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB0FFB2FF3FFB87FF609469.taxon	description	Description. Shell small, moderately broad conical (apical angle 25 – 29 °). Protoconch and early teleoconch whorls unknown. Teleoconch of at least six strongly convex whorls with weakly concave subsutural ramp, rounded shoulder, periphery placed mid-whorl. Suture deeply impressed, fenestrate. Sculpture of prominent, moderately raised, slightly prosocline lamellae, obliquely aligned axially, separated by wider interspaces (?: ~ 14 / 17: ~ 14 / 17), adapical tips faintly alate, fused across suture. Crests of lamellae strongly rolled over, resulting in broad varix-like appearance of lamellae. Interspaces smooth. Last whorl slightly inflated, attaining about 45 % of total height with sub-angled base, lamellae strongly rolled over on base, very broad. Fasciole short, broad, with prominent growth lines. Aperture ovate, columella nearly straight. Columellar callus forming narrow rim, attached to fasciole and base. Parietal callus weak. Basal lip slightly widened. Peristome moderately thickened, duplex; inner layer covering outer layer largely. No umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB0FFB2FF3FFB87FF609469.taxon	discussion	Discussion. Hyaloscala rainesi nov sp. is smaller than Hyaloscala rustica sp. nov. and differs in its faintly coronate shoulder. Hyaloscala occidentalis sp. nov. has a comparable outline but more numerous and weaker lamellae. The extant Hyaloscala arcanum (DuShane, 1979), from California, is relatively stouter and has fewer lamellae (see DuShane 1979: fig. 1; Weil et al. 1999: fig. 457). Hyaloscala melior (Melvill & Standen, 1903), from the Persian Gulf, has weaker and more widely spaced lamellae (see Melvill & Standen 1893: pl. 7, fig. 9; Weil et al. 1999: fig. 410). Some species of Sodaliscala, like the extant Sodaliscala multicostata (Sowerby, 1844) from the Indo-West Pacific, are quite similar but have distinct spiral cords (Kilburn 1985; figs 129 – 130; Weil et al. 1999: fig. 331). Interspaces in Hyaloscala species are usually smooth, and if they have spiral sculpture, it is much finer or fainter than in Sodaliscala species. Paleoenvironment. Occurrences in the Baden Formation of the Vienna Basin suggest middle to outer neritic settings in up to 250 m water depth (Kranner et al. 2021).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB0FFB2FF3FFB87FF609469.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienna Basin: Bad Vöslau (Austria) (hoc opus).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB2FFBEFF3FFB87FB8C9689.taxon	description	Figs 4 D, 34 E – F, 35 A	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB2FFBEFF3FFB87FB8C9689.taxon	materials_examined	Type material. Holotype, NHMW 1997 z 0178 / 0972 a, SL: 7.4 mm, MD: 3.3 mm, Bad Vöslau (Austria), Figs 4 D, 34 E 1 – E 2, 35 A. Paratype, NHMW 1874 / 0058 / 1126, SL: 3.8 mm, MD: 1.8 mm, Seyithasan (Turkey), illustrated in Landau et al. 2913: pl. 20, fig. 9), Figs 34 F 1 – F 2. Type locality. Bad Vöslau (Austria), Vienna Basin. Type stratum. Baden Formation. Age. Middle Miocene, middle Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB2FFBEFF3FFB87FB8C9689.taxon	etymology	Etymology. Superlative of costata (Latin for ribbed), revering to the high number of axial ribs.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB2FFBEFF3FFB87FB8C9689.taxon	diagnosis	Diagnosis. Small, moderately broad conical shell with turreted protoconch and moderately convex teleoconch whorls with many, close-set, broad, flat lamellae with narrow interspaces and three broad flattened varices per whorl.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB2FFBEFF3FFB87FB8C9689.taxon	description	Description. Shell small, moderately broad conical (apical angle ~ 30 °). Protoconch turreted of 4.5 weakly convex, smooth whorls. Teleoconch of four whorls with steep, weakly concave subsutural ramp, moderately convex below, periphery distinctly below mid-whorl. Suture moderately impressed. Sculpture of numerous, densely spaced, broad, low, weakly prosocline, internally lamellose lamellae, not aligned axially, separated by narrower interspaces (19: ~ 33: ~ 38). Crests of lamellae strongly rolled over with indistinct pimples especially close to the adapical suture. Three varices per whorl more prominent, broad and flattened. Interspaces between varices smooth. Last whorl slightly allometrically widening, attaining 48 % of total height, base convex, no peribasal cord, lamellae persisting over base, strongly rolled over. Fasciole short, broad, with distinct growth lines. Aperture ovate. Columella nearly straight. Columellar callus forming narrow rim attached to fasciole, parietal callus delicate. Peristome strongly thickened, duplex. Basal lip slightly widening. No umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB2FFBEFF3FFB87FB8C9689.taxon	discussion	Discussion. Kilburn (1985) mentioned spiral threads in his diagnosis of Librariscala, which are not seen in the Paratethyan species. Therefore, our generic placement might be disputed. However, the peculiar sculpture (= “ tubular ” varices of Kilburn 1985) of Librariscala costatissima sp. nov. is characteristic for the genus. García (2016) described “ pimple-like projections ” on the varices of his Librariscala boutetorum from French Polynesia (see García, 2016: fig 1 E). These delicate projections are also seen in Librariscala costatissima, which differs from Librariscala boutetorum in its conical outline and less convex whorls (see García 2016: fig. 1). Similarly, Librariscala parvonatrix (Kilburn, 1985), from South Africa, is distinguished by its more strongly convex whorls (Kilburn 1985: figs 110 – 112, Weil et al. 1999: fig. 151). This species was identified as Epitonium cf. pulchellum (Bivona e Bernardi, 1832) by Landau et al. (2013), who discussed the rapidly expanding whorls, non-shouldered convex whorls and very close-set lamellae as differences from the Pliocene to Recent Atlanto-Mediterranean species. We agree with these authors and separate the two, recognizing the Turkish specimen as conspecific with L. costatissima. The teleoconch whorls of Hyaloscala pulchella are regularly increasing in width, whereas the last whorl is slightly allometrically widening in L. costatissima, which differs in its broad crests of the lamellae, whereas Hyaloscala pulchella forms thin lamellae (see Weil et al. 1999: fig. 101; note that Chirli 2009: pl. 13, figs 11 – 17 is probably not this species). The presence of three large varices on the last whorls also distinguishes the Miocene species. The protoconch of L. costatissima is high conical of five whorls but slender, turreted as opposed to four whorls in H. pulchella (see Landau et al. 2006: pl. 11, fig. 3). Paleoenvironment. The occurrence in the Baden Formation suggest middle to outer neritic environments (Kranner et al. 2021), whereas the specimen from Seyithasan (Turkey) derives from a shallow water assemblage.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB2FFBEFF3FFB87FB8C9689.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienna Basin: Bad Vöslau (Austria) (hoc opus). Proto-Mediterranean Sea. Serravallian (Middle Miocene): Karaman Basin: Seyithasan (Turkey (Landau et al. 2013).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB2FFBEFF3FFB87FB8C9689.taxon	type_taxon	Type species. Mazescala thrasys Iredale, 1936; original designation by Iredale (1936: 296). Present-day, Australia.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB2FFBEFF3FFB87FB8C9689.taxon	diagnosis	Diagnosis. “ Scaloid shells with whorls tightly coiled and not umbilicated, lamellae erect and numerous, apical whorls thin and glassy, whorls numerous subshouldered. ” (Iredale 1936: 296).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB2FFBEFF3FFB87FB8C9689.taxon	discussion	Discussion. Although Mazescala is treated as synonym of Epitonium Röding, 1798 by MolluscaBase eds (2024 g) we follow Iredale (1936) and Weil et al. (1999), who accepted Mazescala at genus or subgenus rank. Weil et al. (1999: 212) used this genus for species resembling Hyaloscala de Boury, 1890 [type species Turbo clathratulus Kanmacher, 1798; present-day, eastern Atlantic] but which have numerous convex whorls, a strongly impressed suture and thin, strongly raised lamellae. Hirtoscala Monterosato, 1890 [type species Scalaria cantrainei Weinkauff, 1866; present-day, Mediterranean Sea] differs in its more solid, gradate whorls and more prominent lamellae fused from one whorl to the other over the sutures. In our opinion, Mazescala comprises a group of quite distinctive, small to medium-sized, thin-shelled species, which are characterized by convex whorls with more or less distinct shoulder angulation, deeply impressed suture and numerous thin lamellae often forming short spines along the shoulder. The shells lack a peribasal cord but usually have a weak basal angulation with flattish base. The lamellae persist on the base where they have strongly rolled over crests. The very thin lamellae are frequently damaged resulting in a somewhat ‘ tattered’ appearance of some specimens. The genus is documented by several extant species from Australia [Mazescala bellicosa (Hedley, 1907), M. thrasys Iredale, 1936, M. heloris Iredale, 1936] and Japan [Mazescala koyamai (Nakayama, 1995), M. sakuraii (Kuroda & Habe, 1961), M. sawamurai (Azuma, 1960)]. In most epitoniid genera the presence / absence of spirals is a generic character. In this genus the interspaces can be smooth or have very faint spirals. Most species are found in deeper marine environments. The Pliocene to Pleistocene Mediterranean Mazescala salicensis (Seguenza 1876) and the herein described species from the Middle Miocene of the Paratethys are the only fossil occurrences known to us.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFBFFFBEFF3FFEF3FE3D90CD.taxon	description	Figs 33 A, 34 A – B 2	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFBFFFBEFF3FFEF3FE3D90CD.taxon	materials_examined	Type material. Holotype, NHMW 1865 / 0001 / 0841, SL: 5.5 mm, MD: 2.1 mm, Vienna / Grinzing (Austria), Figs 34 B 1 – B 2. Paratype, NHMW 1866 / 0001 / 0717, SL: 3.5 mm, MD: 1.6 mm, Möllersdorf (Austria), Figs 33 A, 34 A 1 – A 2. Type locality. Vienna / Grinzing (Austria), Vienna Basin. Type stratum. Baden Formation. Age. Middle Miocene, Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFBFFFBEFF3FFEF3FE3D90CD.taxon	etymology	Etymology. Referring to the slightly alate varices.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFBFFFBEFF3FFEF3FE3D90CD.taxon	diagnosis	Diagnosis. Small, moderately slender shell with concave subsutural ramp and angled periphery placed above mid-whorl, deeply impressed suture, numerous, close-set, thin, strongly raised, weakly prosocline lamellae, obliquely aligned axially, broad low, trigonal spines along shoulder.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFBFFFBEFF3FFEF3FE3D90CD.taxon	description	Description. Shell small, moderately slender (apical angle ~ 25 – 27 °), faintly coronate. Protoconch unknown. Teleoconch whorls with weakly concave subsutural ramp and rounded angulation at periphery placed just above mid-whorl. Suture deeply impressed, linear. Early teleoconch whorls with weak, thin axial ribs, becoming successively more prominent. Later whorls with numerous, close-set, thin, strongly raised, weakly prosocline lamellae, obliquely aligned axially (~ 18: ~ 20: ~ 20); crests of lamellae faintly rolled over. Broad low, trigonal spines along shoulder of some lamellae. Interspaces smooth, glossy. Last whorl with relatively broad, weakly concave subsutural ramp and slightly angled periphery, attaining about 45 % of total height, lamellae persisting on base, strongly rolled over, weak basal angulation, no distinct peribasal cord, moderately broad, flat to weakly concave base. Aperture ovate, columella weakly concave. Outer lip thickened by varix with shoulder-spine. Columellar callus forming narrow rim. No umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFBFFFBEFF3FFEF3FE3D90CD.taxon	discussion	Discussion. Mazescala alata sp. nov. is highly reminiscent of Mazescala kostejana (Boettger, 1902) in size and outline but differs in its much higher lamellae, whereas M. kostejana has much lower, slightly frondose lamellae. Mazescala kostejana develops small, delicate, subtrigonal spines along the shoulder whereas M. alata has much broader and larger wing-like spines. In addition, it lacks spiral sculpture and has a wide base. Mazescala alata is a look-alike of the extant Australian Mazescala bellicosa (Hedley, 1907), which is nearly identical in size (SL: 7.5 mm), sculpture and general outline and differs only in its fewer lamellae (see Hedley 1907: pl. 67, fig. 18; Iredale 1936: pl. 22, fig. 4). Mazescala thrasys Iredale, 1936, from Austria, is larger at same number of whorls (SL: 15.5 mm) and slenderer (see Iredale 1936: pl. 22, fig. 2). The Japanese Mazescala sakuraii (Kuroda & Habe, 1961) has many more lamellae (Weil et al. 1999: fig. 377) and Mazescala sawamurai (Azuma, 1960) is slenderer and has weaker angled whorls (Weil et al. 1999: fig. 378). The Plio-Pleistocene Mazescala salicensis (Seguenza 1876) is much larger with more numerous whorls and frondose lamellae (see Landau et al. 2006: pl. 11, fig. 4). Paleoenvironment. Occurrences in the Baden Formation of the Vienna Basin suggest middle to outer neritic settings in up to 250 m water depth (Kranner et al. 2021). Mazescala salicensis (Seguenza 1876) is also found in deeper water deposits, indicating at least 200 m water depth (own data B. L.).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFBFFFBEFF3FFEF3FE3D90CD.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienna Basin: Vienna / Grinzing, Möllersdorf (Austria) (hoc opus).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFBFFFB8FF3FF8B7FAEE9229.taxon	description	Figs 4 G, 33 B – C, 36 A – C	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFBFFFB8FF3FF8B7FAEE9229.taxon	materials_examined	Type material. Holotype, SMF 380173 _ 12 - 1446 a (XII 1446 a), specimen illustrated in Zilch (1934: pl. 10, fig. 76), SL: 4.0 mm, MD: 1.2 mm, CoŞteiu de Sus (Romania), illustrated in Zilch (1934: pl. 10, fig. 80), Fig 36 A. Illustrated material. NHMW 1858 / 0047 / 0048, SL: 5.2 mm, MD: 2.5 mm, Porzteich at Břeclav (Czech Republic), Figs 33 B, 36 C 1 – C 2. NHMW 1859 / 0019 / 0108, SL: 3.2 mm, MD: 1.4 mm, Porzteich at Břeclav (Czech Republic), Figs 4 G, 33 C, 36 B 1 – B 2. Revised description. Shell small, moderately slender (apical angle ~ 25 – 28 °), faintly coronate. Protoconch turreted of three weakly convex whorls. Teleoconch whorls with weakly concave subsutural ramp and rounded angulation at periphery placed just above mid-whorl. Suture deeply impressed, linear. Early teleoconch whorls with weak, indistinct axial ribs, becoming successively more prominent. Later whorls with numerous, close-set, thin, weakly raised, lamellose to foliaceous, weakly prosocline lamellae, not clearly aligned axially (~ 18: ~ 16 / 20: ~ 20), crests of lamellae erect. Small, low, vaguely trigonal spines along shoulder. Interspaces glossy with very faint spiral cords, most prominent close to abapical suture and a transition to base. Last whorl with relatively broad, attaining about 45 % of total height, with weakly concave subsutural ramp, slightly angled periphery, lamellae persisting over base, strongly rolled over, weak basal angulation, no distinct peribasal cord, moderately narrow, flat to weakly concave base. Aperture subcircular, columella weakly concave. Outer lip thickened by varix. Columellar callus forming narrow rim, detached from base, parietal callus delicate. Umbilicus reduced to narrow chink in subadult specimens; closed in fully grown specimens.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFBFFFB8FF3FF8B7FAEE9229.taxon	discussion	Discussion. Mazescala kostejana (Boettger, 1902) is characterized by its angled whorls and the ‘ tattered’ appearance of the lamellae. It is reminiscent of the recent Australian Mazescala bellicosa (Hedley, 1907), which is similar in size (SL: 7.5 mm), sculpture and general outline but differs in its subcylindrical last whorl and the broader base (see Hedley 1907: pl. 67, fig. 18; Iredale 1936: pl. 22, fig. 4). Mazescala thrasys Iredale, 1936, from Australia, is larger at same growth stage (SL. 15.5 mm), has more teleoconch whorls, is slenderer and has a narrower subsutural ramp on the last whorl (see Iredale 1936: pl. 22, fig. 2). The Australian Mazescala heloris Iredale, 1936 is larger (SL. 13.5 mm) and has a broad subsutural shelf, resulting in a gradate outline (see Iredale 1936: pl. 22, fig. 3). Mazescala koyamai (Nakayama, 1995), from Japan, has also somewhat ‘ tattered’ lamellae but has a weaker shoulder angulation and Mazescala sakuraii (Kuroda & Habe, 1961), from Japan, differs in its narrower and shallower subsutural ramp and in its more numerous varices (Weil et al. 1999: fig. 377). The Japanese Mazescala sawamurai (Azuma, 1960) has weaker angled whorls and a more convex subsutural ramp (Weil et al. 1999: fig. 378). The Pliocene to Pleistocene Mediterranean Mazescala salicensis (Seguenza 1876) is larger (SL: 14.0 mm), has more prosocline lamellae and is slenderer (see Landau et al. 2006: pl. 11, fig. 4). Paleoenvironment. Probably middle to outer neritic environments.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFBFFFB8FF3FF8B7FAEE9229.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Korytnica Basin: Korytnica (Poland) (Bałuk 1975); Vienna Basin: Porzteich at Břeclav (Czech Republic) (hoc opus), Borský Mikuláš, Devínska Nová Ves (Slovakia) (Švagrovský 1981, 1982); Pannonian Basin: Szokolya (Hungary) (Báldi 1960); Bükk Mountains: Balaton (Hungary) (Csepreghy-Meznerics 1969 a); Făget Basin: CoŞteiu de Sus (Romania) (Boettger 1902).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB9FFB8FF3FFB13FA249101.taxon	description	Figs 4 H, 33 D, 34 C	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB9FFB8FF3FFB13FA249101.taxon	materials_examined	Illustrated material. NHMW 2024 / 0278 / 0010, SL: 4.2 mm, MD: 1.8 mm, Bad Vöslau (Austria), Figs 4 H, 33 D, 34 C 1 – C 2.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB9FFB8FF3FFB13FA249101.taxon	description	Description. Shell very small, moderately slender (apical angle ~ 22 °), weakly coronate. Protoconch turreted, of four smooth, weakly convex whorls. Teleoconch of seven strongly convex whorls, with weakly concave subsutural ramp, angled shoulder, with periphery placed just above mid-whorl. Suture deeply impressed, weakly undulating. Sculpture of moderately widely spaced, moderately raised, only weakly prosocline, thin but robust lamellae, partly obliquely aligned axially (11: 13: 13). Crest of lamellae only weakly rolled over. Low, broad, trigonal spines at lamellae along shoulder. Interspaces between lamellae smooth, glossy. Last whorl attaining about 38 % of total height, lamellae persisting on base, becoming strongly rolled over, no peribasal cord. Aperture not preserved.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB9FFB8FF3FFB13FA249101.taxon	discussion	Discussion. This species differs from Mazescala kostejana (Boettger, 1902) in its fewer lamellae, presence of faint spiral sculpture in the lamellar interspaces, and its protoconch consist of four instead of three whorls. Mazescala alata sp. nov. is broader and differs in its greater number of lamellae. The Pliocene to Pleistocene Mediterranean Mazescala salicensis (Seguenza 1876) is larger (SL: 14.0 mm) and has more numerous and more prosocline lamellae (see Landau et al. 2006: pl. 11, fig. 4). Paleoenvironment. Occurrences in the Baden Formation of the Vienna Basin suggest middle to outer neritic settings in up to 250 m water depth (Kranner et al. 2021).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB9FFB8FF3FFB13FA249101.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienna Basin: Baden (Austria) (hoc opus).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB8FFBAFF3FFF2EFBB694D9.taxon	description	Figs 33 E, 34 D	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB8FFBAFF3FFF2EFBB694D9.taxon	materials_examined	Illustrated material. NHMW 2010 / 0004 / 0526 b, SL: 4.2 mm, MD: 1.7 mm, Bad Vöslau (Austria), Figs 33 E, 34 D.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB8FFBAFF3FFF2EFBB694D9.taxon	description	Description. Shell very small, moderately slender, turreted (apical angle ~ 23 °), faintly coronate. Protoconch and early teleoconch whorls unknown. Teleoconch of at least seven strongly convex whorls, with narrow, weakly concave subsutural ramp and convex periphery placed mid-whorl. Suture deeply impressed. Sculpture of numerous, close-set, thin, moderately raised, nearly orthocline lamellae, not clearly axially aligned (?: 26: 28), with crests only weakly rolled over. Interspaces smooth. Last whorl not preserved.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB8FFBAFF3FFF2EFBB694D9.taxon	discussion	Discussion. Differs from Mazescala kostejana (Boettger, 1902) and M. alata sp. nov. in its much greater number of lamellae, weaker angulation and less incised suture. Paleoenvironment. Occurrences in the Baden Formation of the Vienna Basin suggest middle to outer neritic settings in up to 250 m water depth (Kranner et al. 2021).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB8FFBAFF3FFF2EFBB694D9.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienna Basin: Bad Vöslau (Austria) (hoc opus). Genus Nitidiscala de Boury, 1909	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB8FFBAFF3FFF2EFBB694D9.taxon	type_taxon	Type species. Scalaria unifasciata G. B. Sowerby II, 1844; original designation by de Boury (1909 a: 257). Present-day, Caribbean Sea.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB8FFBAFF3FFF2EFBB694D9.taxon	diagnosis	Diagnosis. Small, moderately stocky to moderately slender conical, occasionally coronate. Protoconch high conical. Teleoconch whorls convex, occasionally with weak shoulder. Deeply impressed suture. Sculpture of moderately widely spaced, very prominent, robust, slightly flaring lamellae, with crests often becoming successively more rolled over during ontogeny. Often with slightly alate tips at adapical suture. Lamellae often obliquely aligned axially and fused over suture. Shell surface smooth, glossy. Last whorl relatively large with sub-angled, weakly convex base, no peribasal band or basal disc, lamellae very prominent and strongly rolled over on base. Fasciole short, broad, with prominent growth lines. Aperture ovate. Columella weakly concave. Columellar callus forming narrow rim attached to fasciole, parietal callus delicate. Peristome strongly thickened, glossy, duplex with inner layer covering outer layer largely. No umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFB8FFBAFF3FFF2EFBB694D9.taxon	discussion	Discussion. We treat Nobiliscala de Boury 1917 [type species Scalaria foliacea G. B. Sowerby I, 1823; original diagnosis by de Boury (1917: 60); Pliocene, British Islands] as subjective junior synonym of Nitidiscala de Boury, 1909 because we find no feature to separate both genera. Kilburn (1985) restricted Nitidiscala to species with prominent lamellae, such as Nitidiscala hexagonum (Sowerby, 1844) and N. unifasciata (G. B. Sowerby II, 1844) (see Weil et al. 1999: figs 41, 477). Strong (1930), DuShane (1974) and Collin (2000) listed several species in Nitidiscala reflecting a rather wide range of morphologies. Molecular data will be needed to better understand this group. Herein, we place Scala (Clathrus) oligocostata Bałuk, 1975 in Nitidiscala due to its resemblance to the eastern Pacific Nitidiscala statuminata (G. B. Sowerby II, 1844) [= Epitonium strongianum Lowe, 1932] [see DuShane 1974: figs 76 – 77; Weil et al. 1999: fig. 493]. In addition, we accommodate a group of Atlanto-Mediterranean and Paratethyan Miocene to Recent species in Nitidiscala: Nitidiscala remota Cossmann & Peyrot, 1922, from the Burdigalian of the northeastern Atlantic (see Cossmann & Peyrot, 1922: pl. 4, figs 7 – 8) and N. baluki sp. nov., N. bonfittoi sp. nov., N. browni sp. nov. and N. detracta (de Boury in Cossmann, 1912) from the Middle Miocene of the Central Paratethys Sea; Nitidiscala foliacea (G. B. Sowerby I, 1823) (see Chirli 2009: pl. 11, figs 7 – 12, Landau et al. 2006: pl. 11, fig. 8), N. pultoastensis (Sacco, 1891) (see Ferrero Mortara et al. 1984: pl. 5, fig. 9) and N. septemcostata (Conti, 1871) (see Landau et al. 2006: pl. 11, figs 5 – 7) from the Pliocene of the Mediterranean Sea and the extant N. jolyi (Monterosato, 1878) [= N. senegalensis (von Maltzan, 1885)] from the Mediterranean Sea and west Africa (Weil et al. 1999: fig. 103) [note that Landau et al. (2006: 35) considered Scalaria jolyi Monterosato, 1878 to be a subjective junior synonym of Scalaria septemcostata Conti, 1871]. Nitidiscala might appear already during the Middle Eocene in the Paris Basin with species placed by de Boury (1887 a, 1887 b) in Gyroscala, such as ‘ Scalaria’ contabulata Deshayes, 1861 (Deshayes 1861: pl. 11, fig. 11). Several Early Miocene taxa, placed in Gyroscala by Cossmann & Peyrot (1922) are also closer to Nitidiscala but differ in the presence of a peribasal cord (e. g., Gyroscala neovasconiensis Cossmann & Peyrot, 1922 (Cossmann & Peyrot 1922: pl. 4 fig. 67), Gyroscala despecta Cossmann & Peyrot, 1922 (Cossmann & Peyrot 1922: pl. 4 figs 77 – 78), Gyroscala exspectata (de Boury, 1913). Note that Lozouet et al. (2001) synonymized these species with Scalaria subscalaris d’Orbigny, 1852 and we follow these authors. A revision of the Eocene and Miocene species will be necessary to decide on their generic placement. Similarly, ‘ Gyroscala’ lycocephala (Kensley & Pether, 1986), from the Quaternary of South Africa is almost certainly misplaced in that genus (see Kensley & Pether 1986: figs 16 A – B), documenting the problematic situation of the group. The above listed species are partly reminiscent of Gyroscala de Boury, 1887 a [type species Scalaria commutata Monterosato, 1877; present-day, Mediterranean Sea], which has comparable shell shapes and distinctly aligned lamellae and a thin but prominent peribasal cord (Kilburn 1985: 261). The Paratethyan species lack a peribasal cord and this absence of a peribasal cord reminds also of Connexiscala de Boury, 1909 [type species Scalaria connexa G. B. Sowerby II, 1844; present-day, Philippines]. Connexiscala is similar to Gyroscala but differs in its very narrow lamellae which appear fused across the suture (e. g., Connexiscala continens (Melvill & Standen, 1903), from the Gulf of Oman, C. cultellicosta (de Boury, 1913), from Borneo, and C. smriglioi (Bonfitto, 2010) from the Gulf of Aden). The mode of formation of the comparatively solid lamellae and the general outline with clearly impressed suture of the Paratethyan species is quite different from those of Connexiscala.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFBBFF84FF3FFD43FC3F976D.taxon	description	Figs 4 I, 33 H, 37 A 1 – A 2	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFBBFF84FF3FFD43FC3F976D.taxon	materials_examined	Type material. Holotype, NHM 1876 / 0011 / 0083, SL: 3.1 mm, MD: 1.7 mm, Lăpugiu de Sus (Romania), Figs 4 I, 33 H, 37 A 1 – A 2. Type locality. Lăpugiu de Sus (Romania), Făget Basin. Type stratum. Silt and clay of the Dej Formation. Age. Middle Miocene, Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFBBFF84FF3FFD43FC3F976D.taxon	etymology	Etymology. In honor of Wacław Bałuk (1935 – 2022), (University Warsaw, Poland), in recognition of his contributions on Paratethyan paleontology.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFBBFF84FF3FFD43FC3F976D.taxon	diagnosis	Diagnosis. Very small, moderately broad, stocky shell with low spire, globose last whorl and thin, strongly raised lamellae obliquely aligned axially, without being fused at deeply impressed suture.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFBBFF84FF3FFD43FC3F976D.taxon	description	Description. Shell very small, moderately broad, stocky with relatively low spire and globose last whorl (apical angle 46 °). Protoconch high conical of 3.5 smooth, moderately convex whorls with elevated blunt nucleus. Teleoconch of three strongly convex whorls. Sculpture of thin, strongly raised lamellae, obliquely aligned axially without being fully fused, indented at deeply impressed suture (10: 12: 13), crests erect. No spiral sculpture. Last whorl attaining 60 % of total height, no basal angulation nor peribasal cord but lamellae slightly angled at transition to base, lamellae grading into prominent, relatively broad ribs terminating at columellar callus, base weakly convex. Fasciole very short, very broad with prominent growth lines. Aperture ovate, columella weakly concave. Outer lip strongly thickened by varix. Columellar callus forming broad rim, attached to base, parietal callus delicate. Peristome duplex. No umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFBBFF84FF3FFD43FC3F976D.taxon	discussion	Discussion. This species was confused by Bałuk (2006) with ‘ Epitonium ’ exspectatum (de Boury, 1913) [= ‘ Epitonium ’ subscalare (d’Orbigny, 1852)]. The type specimen of ‘ Epitonium ’ exspectatum, from the Aquitanian of Saint Avit (France), differs from Nitidiscala baluki sp. nov. in its much larger size (SL: 6 mm), wider last whorl with broader subsutural ramp, and the more prominent lamellae (see de Boury 1913 a: pl. 8, fig. 14). ‘ Epitonium ’ subscalare (d’Orbigny, 1852, as understood by Lozouet et al. (2001), from the Aquitanian and Burdigalian of Dax and Saucats (France), is also much larger (SL: 14 mm) and differs in its higher spire, the comparatively thinner and more orthocline lamellae and the peribasal cord and is therefore not congeneric (see Cossmann & Peyrot 1922: pl. 4, figs 39 – 40; Lozouet et al. 2001: pl. 19, fig. 5). ‘ Gyroscala’ series de Boury, 1913 from the Burdigalian of Saint- Jean-de-Marsacq (France), is reminiscent of N. baluki and falls within the same size-range, but differs in its thinner lamellae, which are more fused at the suture and it has a delicate peribasal cord (see de Boury 1913 a: pl. 8, fig. 16). Nitidiscala baluki differs from Nitidiscala connexicosta sp. nov. in its globose last whorl and even thinner varices, which are erect. The extant Connexiscala cultellicosta de Boury, 1913 is superficially similar but is stockier and differs in its more numerous lamellae (see de Boury 1913 a: pl. 8, fig. 17).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FFBBFF84FF3FFD43FC3F976D.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene):? Korytnica Basin: Korytnica (Poland) (Bałuk 2006); Făget Basin: Lăpugiu de Sus (Romania) (hoc opus).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF85FF84FF3FFED7FDFE91EE.taxon	description	Figs 33 I, 37 C 1 – C 2	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF85FF84FF3FFED7FDFE91EE.taxon	materials_examined	Type material. Holotype, NHMW 2024 / 0283 / 0002, SL: 3.4 mm, MD: 1.6 mm, Lăpugiu de Sus (Romania), Figs 33 I, 37 C 1 – C 2. Paratype, BkK-G 1219, SL: 7.2 mm, MD: 3.4 mm, Korytnica (Poland), illustrated in Bałuk (2006: 202, pl. 10, fig. 4). Type locality. Lăpugiu de Sus (Romania), Făget Basin. Type stratum. Silt and clay of the Dej Formation. Age. Middle Miocene, Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF85FF84FF3FFED7FDFE91EE.taxon	etymology	Etymology. In honor of Antonio Bonfitto (University of Bologna, Italy), in recognition of its contributions on Epitoniidae systematics.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF85FF84FF3FFED7FDFE91EE.taxon	diagnosis	Diagnosis. Very small, moderately broad, stocky conical with convex whorls, deeply impressed suture, relatively large last whorl and about 8 or 9 thin, strongly raised, slightly alate lamellae, partly obliquely axially aligned and fused across suture, ovate aperture with strongly thickened peristome.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF85FF84FF3FFED7FDFE91EE.taxon	description	Description. Shell very small, moderately broad, stocky conical (apical angle ~ 35 °). Protoconch high conical with raised small nucleus, of 4.5 smooth, weakly convex whorls, last protoconch whorl with slight mid-whorl angulation (diamater 210 μm). Teleoconch of up to six convex whorls (Bałuk 2006), periphery just below mid-whorl. Suture deeply impressed. Sculpture of moderately close-set, strongly raised, thin but robust, only weakly prosocline lamellae, partly obliquely aligned axially, crossing suture where raised adapical tip align with lamella of preceding whorl (7 / 8: 8 / 9: 8). Crests of lamellae becoming successively more rolled over during ontogeny, partly with alate tips at adapical suture. Interspaces between varices smooth, glossy. Last whorl large, attaining about ~ 46 – 50 % of total height, convex, lamellae persisting on base, strongly rolled over, no basal angulation, no peribasal cord, base slightly convex. Aperture ovate, columella weakly concave. Fasciole short, broad, with prominent growth lines. Columellar callus forming narrow rim, attached to fasciole, parietal callus delicate. Peristome strongly thickened, glossy, duplex with inner layer largely covering outer layer. No umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF85FF84FF3FFED7FDFE91EE.taxon	discussion	Discussion. Nitidiscala bonfittoi sp. nov. differs from Nitidiscala browni sp. nov. in its evenly convex whorls, which lack a shoulder and lack the small trigonal spines developed on the lamellae at the shoulder. The Pliocene Mediterranean Nitidiscala pultoastensis (Sacco, 1891) is very similar but differs in its more prosocline and slightly closer-set lamellae (see Sacco 1891: pl. 1, fig. 30; Ferrero Mortara et al. 1984: pl. 5, fig. 9). The Pliocene Atlanto-Mediterranean Nitidiscala foliacea (G. B. Sowerby I, 1823) is slightly stockier and has more numerous lamellae (see Wood 1848: pl. 8, fig. 17; Chirli 2009: pl. 11, figs 7 – 12) and N. frondicula (Wood, 1842) of Chirli 2009: pl. 11, figs 13 – 18) is distinguished by its wider subsutural ramp with weakly coronate shoulder and N. frondicula (Wood, 1842) in Landau et al. (2006: pl. 22, fig. 8) is slenderer with higher lamellose varices. Nitidiscala baluki sp. nov. is comparatively stouter. The Pliocene to Recent Mediterranean Nitidiscala septemcostata (Conti, 1871) has only seven lamellae (see Landau et al. 2006: pl. 11, figs 5 – 7). Paleoenvironment. Unknown.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF85FF84FF3FFED7FDFE91EE.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Korytnica Basin: Korytnica (Poland) (Bałuk 2006); Vienna Basin: Borský Mikuláš (Slovakia) (Švagrovský 1982); Făget Basin: CoŞteiu de Sus, Lăpugiu de Sus (Romania) (Boettger 1902; hoc opus).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF84FF85FF3FFF2EFEA99299.taxon	description	Figs 4 K, 33 J, 37 D 1 – D 2	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF84FF85FF3FFF2EFEA99299.taxon	materials_examined	Type material. Holotype, NHMW 2024 / 0283 / 0002 d, SL: 2.2 mm, MD: 1.0 mm, Lăpugiu de Sus (Romania), Figs 4 K, 33 J, 37 D 1 – D 2. Type locality. Lăpugiu de Sus (Romania), Făget Basin. Type stratum. Dej Formation. Age. Middle Miocene, Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF84FF85FF3FFF2EFEA99299.taxon	etymology	Etymology. In honor of Leonard G. Brown (Connecticut, USA), in recognition of its contributions on Epitoniidae systematics.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF84FF85FF3FFF2EFEA99299.taxon	diagnosis	Diagnosis. Very small, moderately broad, stocky conical, slightly gradate, weakly coronate, deeply impressed suture, relatively large last whorl with about ten thin, strongly raised lamellae partly obliquely aligned axially, fused across suture, with low trigonal spines developed at shoulder, ovate aperture with strongly thickened peristome.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF84FF85FF3FFF2EFEA99299.taxon	description	Description. Shell very small, moderately broad, stocky conical (apical angle ~ 35 °), slightly gradate, weakly coronate. Protoconch high conical, of 3.5 smooth, weakly convex whorls. Teleoconch of up to five whorls. Early teleoconch whorl convex with periphery placed above mid-whorl. Later whorls with broad, weakly concave subsutural ramp, shoulder angulation weakly convex to subcylindrical periphery. Suture deeply impressed. Sculpture of moderately close-set, strongly raised, weakly prosocline, thin but robust lamellae, partly obliquely aligned axially (13: 12: 10), raised adapical tip fused across suture. Lamellae forming low trigonal spines on last whorl, crests of lamellae moderately rolled over. Interspaces between lamellae smooth, glossy. Last whorl large, convex, attaining about ~ 52 % of total height, lamellae persisting over base, strongly rolled over, base sub-angled, no peribasal cord, base narrow, flattened. Aperture ovate, columella weakly concave. Fasciole short, broad, with prominent growth lines. Columellar callus forming narrow rim, attached to fasciole, parietal callus delicate. Peristome strongly thickened, glossy, duplex with inner layer covering outer layer largely. No umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF84FF85FF3FFF2EFEA99299.taxon	discussion	Discussion. Nitidiscala frondicula (Wood, 1842) of Chirli 2009: pl. 11, figs 13 – 18) is larger and differs in its wider subsutural ramp. Nitidiscala bonfittoi sp. nov. from the Middle Miocene of the Paratethys and the Pliocene Mediterranean Nitidiscala pultoastensis (Sacco, 1891) lack a shoulder (see Sacco 1891: pl. 1, fig. 30; Ferrero Mortara et al. 1984: pl. 5, fig. 9). Paleoenvironment. Unknown.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF84FF85FF3FFF2EFEA99299.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Făget Basin: Lăpugiu de Sus (Romania) (hoc opus).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF84FF86FF3FFA82FD2997F9.taxon	description	Figs 4 J, 33 K, 37 B 1 – B 2	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF84FF86FF3FFA82FD2997F9.taxon	materials_examined	Type material. Holotype, NHMW 2024 / 0284 / 0002, SL: 3.4 mm, MD: 1.9 mm, Hrušovany nad Jevišovkou (Czech Republic), Figs 4 J, 33 K, 37 B 1 – B 2. Type locality. Hrušovany nad Jevišovkou (Czech Republic), Carpathian Foreland Basin. Type stratum. Middle Badenian silt and sand. Age. Middle Miocene, middle Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF84FF86FF3FFA82FD2997F9.taxon	etymology	Etymology. Referring to the axial ribs, which are obliquely aligned axially along the teleoconch.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF84FF86FF3FFA82FD2997F9.taxon	diagnosis	Diagnosis. Very small, moderately broad, stocky shell with strongly convex whorls, deeply impressed suture and strongly raised, widely spaced, weakly prosocline lamellae, obliquely aligned axially, indented along suture.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF84FF86FF3FFA82FD2997F9.taxon	description	Description. Shell very small, moderately broad, stocky (apical angle ~ 33 °). Protoconch and early teleoconch whorls unknown. Teleoconch whorls strongly convex with periphery placed mid-whorl. Suture deeply impressed, weakly undulating. Sculpture of moderately close-set, thin, strongly raised, weakly prosocline lamellae, obliquely aligned axially (?: 10: 11), crests only slightly rolled over. Interspaces smooth, glossy. Last whorl slightly widening, roughly attaining half of the total height (apex is missing), moderately convex with nearly subcylindrical periphery, no peribasal cord but lamellae slightly angled at transition to base, lamellae grading into prominent, relatively broad ribs terminating at columellar callus, base weakly convex. Fasciole very short, broad with prominent growth lines. Aperture ovate, columella weakly concave. Outer lip strongly thickened by varix. Columellar callus forming broad rim, attached to base, parietal callus delicate. Peristome duplex. No umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF84FF86FF3FFA82FD2997F9.taxon	discussion	Discussion. ‘ Epitonium’ subscalare (d’Orbigny, 1852), from the Early Miocene of France, has a higher spire and thinner orthocline varices (see Cossmann & Peyrot 1922: pl. 4, figs 39 – 40; Lozouet et al. 2001: pl. 19, fig. 5). Nitidiscala baluki sp. nov. is stockier and has a larger, more inflated last whorl. Nitidiscala baluki is smaller than N. connexicosta and has slightly fewer lamellae. Paleoenvironment. Unknown.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF84FF86FF3FFA82FD2997F9.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Carpathian Foreland Basin: Hrušovany nad Jevišovkou (Czech Republic) (hoc opus).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF87FF86FF3FFE63FACF9173.taxon	description	Figs 33 L, 37 E – H	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF87FF86FF3FFE63FACF9173.taxon	materials_examined	Type material. Holotype, SMF 380172 _ 12 - 1448 a (XII 1448 a), illustrated in Zilch (1934: pl. 10, fig. 80), SL: 6.8 mm, MD: 3.5 mm, CoŞteiu de Sus (Romania), illustrated in Zilch (1934: pl. 10, fig. 80), Fig. 37 E. Illustrated material. NHMW 1865 / 0035 / 0082, SL: 5.1 mm, MD: 3.2 mm, Szob (Hungary), Figs 37 G 1 – G 2. NHMW 1997 z 0178 / 0973, SL: 4.8 mm, MD: 2.8 mm, Bad Vöslau (Austria), Figs 33 L, 37 F 1 – F 2. NHMW 1997 z 0178 / 0973 a, SL: 4.2 mm, MD: 2.6 mm, Bad Vöslau (Austria), Fig. 37 H. Revised description. Shell small, moderately broad (apical angle ~ 31 °), weakly coronate. Protoconch and early teleoconch whorls unknown. Teleoconch of at least seven strongly convex whorls with periphery placed just above mid-whorl. Suture very deeply impressed. Sculpture of widely spaced, strongly raised, thin, prosocline lamellae, obliquely aligned axially (7: 9: 8), lamellae forming short, vaguely trigonal spines along shoulder. Crests of lamellae erect. Interspaces wide, smooth, glossy. Last whorl slightly widening, attaining about 40 % of total height, with shallow subsutural ramp, strongly convex below, lamellae strongly rolled over on base, persisting to columellar callus, no peribasal cord but lamellae form angled nodes at transition to convex base. Very short, broad fasciole. Aperture subcircular, columella weakly convex. Outer lip thickened by varix. Columellar and parietal callus forming broad rim, detached from base and fasciole. Umbilicus narrow, open on subadult specimens (holotype), fully covered by columellar callus on adult specimens.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF87FF86FF3FFE63FACF9173.taxon	discussion	Discussion. This species was placed in Nobiliscala by de Boury (1917: 60). It was confused by Boettger (1902) with the extant western Atlantic Hirtoscala foliaceicosta (d’Orbigny, 1842) (as Scalaria muricata G. B. Sowerby II, 1844) (see Weil et al. 1999: fig. 35). Both species are indeed very similar, but the lamellae of H. foliaceicosta are erect over the base and not rolled over as in Nitidiscala detracta (De Boury in Cossmann, 1912). A specimen illustrated as this species by Csepreghy-Meznerics (1956) from Szob (Hungary) is slenderer and has a narrower subsutural ramp. We have not seen this specimen and include it here with some hesitation. However, another specimen of N. detracta from Szob is illustrated herein, proving its occurrence at that locality. The specimen illustrated by (Bałuk 1975) from Korytnica (Poland) differs in its much broader subsutural ramp and prominent shoulder spines. That specimen is identified herein as Hirtoscala cornucopiae sp. nov. Paleoenvironment. Unknown.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF87FF86FF3FFE63FACF9173.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienna Basin: Bad Vöslau (Austria) (hoc opus); Pannonian Basin: Szob (Hungary) (hoc opus); Făget Basin: CoŞteiu de Sus (Romania) (Boettger 1902).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF87FF80FF3FF8ECFDEA96A5.taxon	description	Figs 33 F – G, 36 D – F	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF87FF80FF3FF8ECFDEA96A5.taxon	materials_examined	Illustrated material. NHMW 2024 / 0283 / 0001, SL: 3.9 mm, MD: 2.0 mm, Lăpugiu de Sus (Romania), Figs 33 F, 36 D 1 – D 2. NHMW 1863 / 0015 / 1006, SL: 7.4 mm, MD: 3.5 mm, Rousínov (Czech Republic), Figs 33 G, 36 F. NHMW 2024 / 0284 / 0001, SL: 4.5 mm, MD: 2.2 mm, Hrušovany nad Jevišovkou (Czech Republic), Figs 36 E 1 – E 2. Revised description. Shell small, moderately broad (apical angle ~ 28 °). Protoconch and early teleoconch whorls unknown. Teleoconch whorls strongly convex with periphery placed just below mid-whorl. Suture deeply impressed, weakly undulating. Sculpture of widely spaced, strongly raised, slightly flaring, robust, prosocline lamellae, obliquely aligned axially (?: 7: 7), crests slightly rolled over. Interspaces wide, smooth, glossy. Last whorl relatively high, moderately convex, slightly angled at mid-whorl, lamellae persisting and strongly rolled over on base, no peribasal cord, base slightly convex. Aperture depressed ovate, columella weakly concave, nearly straight. Outer lip slightly thickened by varix. Columellar callus forming broad rim, detached from base; parietal callus weak. Peristome duplex but outer layer largely covered by inner layer.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF87FF80FF3FF8ECFDEA96A5.taxon	discussion	Discussion. There are only few comparable species known from the Miocene of the Circum-Mediterranean Region. Nitidiscala recidiva (de Boury in Cossmann & Peyrot, 1922), from the Early Miocene of France, differs in its even wider interspaces between the lamellae and lacks a weak mid-whorl angulation of the last whorl (Cossmann & Peyrot 1922: pl. 3, figs 14 – 15). Nitidiscala benoisti (de Boury in Cossmann & Peyrot, 1922), from the Early Miocene of France, has weaker lamellae on the spire whorls and has a larger last whorl (see Cossmann & Peyrot 1922: pl. 3, figs 69 – 70). Although the French species were placed in the genus Foveoscala de Boury, 1909 [= Hirtoscala Monterosato, 1890] by Cossmann & Peyrot (1922), this allocation is rejected herein based on the rolled over crests of the lamellae and the lack of flaring varices with trigonal shoulder spines (see discussion under Hirtoscala). ‘ Clathrus’ mutinofoliaceus Sacco, 1891, from the Burdigalian or Langhian of the Colli Torinesi (Italy), is stockier and differs in its more convex whorls with deeply impressed suture and fewer, orthocline lamellae (see Sacco 1891: pl. 1, fig. 29). ‘ Clathrus’ miopaucicostulata Sacco, 1891, from the Tortonian of Montegibbio (Italy) is based on a fragment which seems to have a peribasal cord, as typical for Gyroscala, and its lamellae are less flaring and less prosocline (see Sacco 1891: pl. 1, fig. 28). The status of these two Italian species will need revision and we hesitate to place them in Nitidiscala. The Paratethyan species differs from the extant Nitidiscala statuminata (G. B. Sowerby II, 1844) only in its less flaring varices, which are not as fused along the suture. Some species of Laeviscala de Boury, 1909 [type species Scalaria subauriculata Souverbie in Souverbie & Montrouzier, 1866; present-day, New Caledonia] are reminiscent of Nitidiscala oligocostata but Laeviscala is characterized by its cancellate microsculpture in the lamellar interspaces (Bonfitto 2018: 120). Paleoenvironment. Unknown.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF87FF80FF3FF8ECFDEA96A5.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Korytnica Basin: Korytnica (Poland) (Bałuk 2006); North Alpine-Carpathian Foreland Basin: Hrušovany nad Jevišovkou, Rousínov (Czech Republic) (hoc opus). Genus Nodiscala de Boury, 1890	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF87FF80FF3FF8ECFDEA96A5.taxon	type_taxon	Type species. Scalaria bicarinata Sowerby, G. B. II., 1844); original designation by de Boury (1890: 168). Present-day, Indo-West Pacific.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF87FF80FF3FF8ECFDEA96A5.taxon	diagnosis	Diagnosis. “ Thick shell. Size small; shape narrow, somewhat pupoidal; spire elongate, with subconoidal curvature; protoconch smooth, polygyrate; whorls convex, with sutures often bordered above: ornamentation composed of axial ribs, subnodular anteriorly, often interrupted below the sutural bead, and fine rows of spiral punctations, sometimes transforming into deeper grooves, at the anterior part of each whorl. Last whorl little more than a third of the total height, with a convex, imperforate base, often with a very obsolete, spirally punctate disc. Aperture small and circular; peristome split, inner layer thin and continuous; outer layer broadly bordered and discontinuous; labrum varicose, slightly oblique. ” (translated from Cossmann 1912: 84).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF87FF80FF3FF8ECFDEA96A5.taxon	discussion	Discussion. Nodiscala is treated as subjective junior synonym of Opalia H. Adams & A. Adams, 1853 by MolluscaBase eds (2024 h) but Opalia australis (Lamarck, 1822), the type species of Opalia, differs significantly from Nodiscala species in its conical shape, non-crenulate suture and prominent axial ribs, which emerge from the prominent peribasal cord and the lack of spiral sculpture. Therefore, we keep both genera separated (for additional discussions see Landau et al. 2006: 49).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF81FF80FF3FFE9FFDE493A5.taxon	description	Figs 35 B, 38 A – D	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF81FF80FF3FFE9FFDE493A5.taxon	materials_examined	Type material. Holotype, SMF 380181 _ 12 - 1460 a (XII 1460 a), SL: 2.5 mm, MD: 1.0 mm, CoŞteiu de Sus (Romania), specimen illustrated in Zilch (1934: pl. 10, fig. 67), Fig. 38 A. Illustrated material. NHMW 1997 z 0178 / 0970, SL: 5.0 mm, MD: 2.1 mm, Bad Vöslau (Austria), Figs 38 D 1 – D 2. NHMW 1864 / 0001 / 0102 a, SL: 3.8 mm, MD: 1.6 mm, Baden (Austria), Figs 35 B, 38 B 1 – B 2. NHMW 1864 / 0001 / 0102 b, SL: 2.8 mm, MD: 1.3 mm, Baden (Austria), Figs 38 C 1 – C 2. Revised description. Shell small, moderately slender (apical angle ~ 21 – 25 °). Protoconch incomplete but multispiral, high conical, of smooth, weakly convex whorls. Teleoconch of six strongly convex whorls. Whorl profile with moderately steep, weakly concave subsutural ramp, angled periphery placed just below or above mid-whorl. Suture distinctly impressed, weakly undulating. Sculpture of moderately close-set, prominent, convex, orthocline to weakly prosocline low axial ribs, not aligned axially (8: 13: 13). Axial ribs weakening over subsutural ramp, most prominent at periphery. One more prominent, prosocline varix per whorl may appear. Spiral sculpture of numerous spiral cords separated by slightly narrower, deeply punctate interspaces. Last whorl attaining ~ 42 % of total height, moderate basal angulation with subobsolete peribasal cord, base weakly convex to nearly flat with prominent spiral cords. Aperture ovate, columella weakly concave. Outer lip strongly thickened by terminal varix. Columellar and parietal callus forming continuous, narrow rim, detached from base. Peristome duplex. Umbilicus reduced to narrow chink.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF81FF80FF3FFE9FFDE493A5.taxon	discussion	Discussion. Nodiscala macilenta (De Boury in Cossmann, 1912) differs from Nodiscala scacchii (Hörnes, 1856) in its broader conical outline and broader, more prominent axial ribs that lack the vague nodes developed at the shoulder in N. scacchii. The extant Atlanto-Mediterranean Nodiscala coronata (Philippi & Scacchi in Philippi, 1840) [= Nodiscala hellenica (Forbes, 1844)] may develop shells with angled whorls, which are reminiscent of N. macilenta but N. coronata is slenderer, with less convex whorls, the suture is more superficial and strongly crenulate (see Landau et al. 2006: pl. 15, figs 2 – 3). Paleoenvironment. Occurrences in the Baden Formation of the Vienna Basin suggest middle to outer neritic settings in up to 250 m water depth (Kranner et al. 2021).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF81FF80FF3FFE9FFDE493A5.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienna Basin: Baden, Bad Vöslau (Austria) (hoc opus); Southern Pannonian Basin: Miljevići (Bosna and Herzegovina) (Atanacković 1969); Făget Basin: CoŞteiu de Sus (Romania) (Zilch 1934).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF81FF82FF3FF99FFE97959D.taxon	description	Figs 35 C, 38 E – F	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF81FF82FF3FF99FFE97959D.taxon	materials_examined	Type material. Lectotype (designated by Zilch 1934: 229), SMF 380176 _ 12.1457 a (XII 1457 a), SL: 3.5 mm, MD: 1.3 mm, CoŞteiu de Sus (Romania), specimen illustrated in Zilch (1934: pl. 10, fig. 75), Figs 38 E. Illustrated material. NHMW 1862 / 0007 / 0287, SL: 3.6 mm, MD: 1.5 mm, Möllersdorf (Austria), Figs 35 C, 38 F 1 – F 2. Revised description. Very small, moderately slender conical shell (apical angle 25 °). Protoconch turreted of four glossy whorls, last whorl with weak mid-whorl angulation, apically cyrtoconoidal (diameter 300 μm). Teleoconch of four strongly convex whorls with indistinct, weakly convex subsutural ramp and deeply impressed, slightly undulating suture. Sculpture of relatively narrow, weakly prosocline, convex axial ribs separated by wider interspaces, not strictly aligned axially (12: 13: 13). Tips of axial ribs prominent at suture usually separated from preceding whorl. Spiral sculpture of about five primary spiral cords and numerous close-set spiral threads separated by narrow, punctate grooves. Last whorl convex, attaining ~ 45 % of total height, basal angulation moderate with weak peribasal cord delimiting relatively narrow basal disc bearing weak spiral cords. Aperture subcircular with concave columella. Columellar and parietal lip continuous, forming narrow, weakly detached rim. Basal lip truncated. Peristome duplex. Outer lip strongly thickened with radial internal sculpture.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF81FF82FF3FF99FFE97959D.taxon	discussion	Discussion. This species was placed in Funiscala by previous authors but the punctate microsculpture excludes this placement. It is reminiscent of Nodiscala macilenta (De Boury in Cossmann, 1912) is size, shape and sculpture but differs in its strongly convex whorls, better defined axial ribs and coarser spiral cords. Cossmann (1912: 195) proposed Funiscala perpusilla as replacement name for Pliciscala (Funiscala) microscopica Boettger, 1906 and stated “ non Cossmann ” without referring to a certain species or paper. Zilch (1934) accepted this new name. Indeed, Cossmann (1902: 26) had described Scala (Bouryiscala) microscopica from the Bartonian from Bois-Gouët (Loire-Atlantique) [= Graphis (Bouryiscala) microscopica (Cossmann, 1902)] but not a Pliciscala microscopica. Therefore, the replacement name is unnecessary. ‘ Funiscala’ imperfecta de Boury, 1891, from the Tortonian of Montegibbio (Italy), differs in its slightly less convex whorls and much stronger spiral cords (see de Boury 1891: 206, pl. 4, fig. 13) and it does not have a perforated surface and therefore is not congeneric with Nodiscala microscopica (Boettger, 1906). Paleoenvironment. Unknown.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF81FF82FF3FF99FFE97959D.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Făget Basin: CoŞteiu de Sus (Romania) (Zilch 1934).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF83FF8CFF3FFB87FC779715.taxon	description	Figs 35 D – E, 38 G – N	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF83FF8CFF3FFB87FC779715.taxon	materials_examined	Type material. Lectotype (designated herein), NHMW 1865 / 0001 / 0584, SL: 6.5 mm, MD: 2.1 mm, Steinebrunn (Austria), illustrated in Hörnes (1856, pl. 46, fig. 12), Figs 38 G. Illustrated material. SMF 380170 _ 12 - 1445 (XII 1445 a), SL: 8.1 mm, MD: 2.8 mm, CoŞteiu de Sus (Romania), holotype of Nodiscala rugatina De Boury in Cossmann, 1912, illustrated in Zilch (1934: pl. 10, fig. 72), Fig. 38 H. NHMW 1855 / 0014 / 0207, SL: 8.9 mm, MD: 3.2 mm, Steinebrunn (Austria), Figs 38 I 1 – I 2. SMF 380177 _ 12 - 1444 a (XII 1444 a), SL: 4.7 mm, MD: 2.0 mm, CoŞteiu de Sus (Romania), holotype of Pliciscala (Nodiscala) bimonilifera Boettger, 1902, illustrated in Zilch (1934: pl. 10, fig. 70), Fig. 38 J. NHMW 1868 / 0001 / 0541, SL: 7.2 mm, MD: 2.7 mm, Lăpugiu de Sus (Romania), Figs 35 D, 38 K. NHMW 1870 / 0019 / 0251, SL: 5.1 mm, MD: 1.8 mm, CoŞteiu de Sus (Romania), Figs 35 E, 38 L 1 – L 2. SMF 380178 _ 12 - 1461 a (XII 1461 a), SL: 4.0 mm, MD: 1.2 mm, CoŞteiu de Sus (Romania), holotype of Nodiscala angulipunctata De Boury in Cossmann, 1912, illustrated in Zilch (1934: pl. 10, fig. 71), Fig. 38 M. NHMW 1860 / 0011 / 0305, SL: 4.2 mm, MD: 1.6 mm, Lăpugiu de Sus (Romania), Fig. 38 N. Revised description. Shell small, moderately slender (apical angle ~ 16 – 22 °). Protoconch incomplete, high conical, of smooth, moderately convex whorls. Teleoconch of up to six whorls. Whorl profile with indistinct, narrow subsutural collar, steep, weakly concave subsutural ramp and weakly angled periphery placed just below mid-whorl. Suture moderately impressed, weakly undulating, occasionally crenulate. Sculpture of moderately close-set, convex, orthocline to weakly prosocline axial ribs, not aligned axially (~ 11: ~ 10 / 11: ~ 10 / 11). Axial ribs weakening over subsutural ramp, most prominent along rounded angulation. One or two more prominent varices may occur per whorl (angulipunctata - morph). Occasionally, axial ribs may form rounded nodes at shoulder and along basal angulations (bimonilifera - morph). Spiral sculpture of numerous spiral cords separated by slightly narrower, deeply punctate interspaces. Last whorl attaining ~ 45 % of total height, axial ribs rapidly weakening over convex base, basal angulation weak to distinct, often with subobsolete peribasal cord, numerous spiral cords on base. Aperture ovate, columella weakly concave. Outer lip strongly thickened by terminal varix, with radial sculpture. Columellar and parietal callus forming continuous, narrow rim, detached from base. Peristome duplex. Umbilicus reduced to narrow chink.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF83FF8CFF3FFB87FC779715.taxon	description	Paratethyan synonyms. We follow Landau et al. (2006) and consider Pliciscala (Nodiscala) bimonilifera Boettger, 1902, Nodiscala angulipunctata de Boury in Cossmann, 1912 and Nodiscala rugatina de Boury in Cossmann, 1912 as subjective junior synonyms of Nodiscala scacchii (Hörnes, 1856). Pliciscala (Nodiscala) bimonilifera is characterized by its prominent nodes along the shoulder of the last whorl and a second row of weaker rows along the basal angulation and Nodiscala angulipunctata is based on a specimen with very prominent varices.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF83FF8CFF3FFB87FC779715.taxon	discussion	Discussion. This is a very variable and widespread species, which was discussed in detail by Landau et al. (2006: 50). Paleoenvironment. Shallow marine, inner neritic; the occurrence at Steinebrunn (Austria) suggests seagrass environments (own data M. H.).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF83FF8CFF3FFB87FC779715.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Korytnica Basin: Korytnica (Poland) (Bałuk 2006); Vienna Basin: Steinebrunn (Austria) (Hörnes 1856); Pannonian Basin: Hidas, Szob, Várpalota (Hungary) (Csepreghy-Meznerics 1950, 1956; Strausz 1966); Mecsek Mountains (Hungary) (Strausz 1966); Southern Pannonian Basin: Miljevići (Bosnia and Herzegovina) (Atanacković 1985); Făget Basin: CoŞteiu de Sus (Romania) (Zilch 1934). Mediterranean Sea. Pliocene: Estepona Basin: Estepona (Spain) (Landau et al. 2006); Central Mediterranean: Italy (Brunetti 2000). Northeastern Atlantic. Aquitanian (Early Miocene): Aquitaine Basin: Saucats (France) (Lozouet et al. 2011). Pliocene: Loire Basin: Pigeon Blanc (France) (Van Dingenen et al. 2016).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF83FF8CFF3FFB87FC779715.taxon	description	North Sea. Early Miocene: North Sea Basin: Belgium (Glibert 1949); Middle Miocene: North Sea Basin: The Netherlands (Nordsieck 1972; Janssen 1984), Germany (Wienrich 2001).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF8DFF8DFF3FFE0FFB889715.taxon	description	Figs 35 F, 38 O	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF8DFF8DFF3FFE0FFB889715.taxon	materials_examined	Type material. Lectotype (designated by Zilch 1934: 228), SMF 380182 _ 12 - 1456 a (XII 1456 a), SL: 5.1 mm, MD: 1.6 mm, CoŞteiu de Sus (Romania), specimen illustrated in Zilch (1934: pl. 10, fig. 69), Figs 35 F, 38 O. Revised description. Small, moderately slender, somewhat uncoiled shell (apical angle ~ 22 °). Protoconch unknown. Early teleoconch whorls with straight sided subsutural ramp, mid-whorl angulation, constricted below. Later whorls with second angulation close above abapical suture, with periphery placed at shoulder angulation slightly above mid-whorl. Whorls straight-sided below, funnel shaped down to lower angulation. Suture extremely deeply incised below base of whorls resulting in loosely coiled appearance. Sculpture of prominent, strongly raised, broad, convex axial ribs separated by slightly wider interspaces (~ 12: 12: 12). One large, strongly protruding varix may occur per whorl. Spiral sculpture of prominent spiral cords, overriding axial ribs. Punctate microsculpture very weak. Last whorl attaining ~ 40 % of total height, strongly constricted at base with distinct basal angulation without clear peribasal cord. Very prominent, raised, cord-like fasciole. Aperture subcircular; columella moderately concave. Outer lip with small expansion abapically. Peristome duplex with continuous glossy inner layer, forming narrow rim. Outer layer strongly thickened with radial sculpture. Umbilicus forming narrow, relatively deep chink.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF8DFF8DFF3FFE0FFB889715.taxon	discussion	Discussion. The only available specimen is characterized by its slightly decoiled shell. This feature, however, may be pathological. Comparable shells are known for example from the Atlanto-Mediterranean Opalia coronata (Philippi & Scacchi in Philippi, 1840), which usually has ‘ normally’ coiled whorls in which the base of the preceding whorl is not exposed along the suture. It is unlikely that Nodiscala transsylvanica is an aberrant specimen of Nodiscala scacchii (Hörnes, 1856), as it differs from that species in its very prominent axial ribs the spiral cords, which overrun the axial ribs. Paleoenvironment. Unknown.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF8DFF8DFF3FFE0FFB889715.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Făget Basin: CoŞteiu de Sus (Romania) (Boettger 1906). Genus Parviscala de Boury, 1887	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF8DFF8DFF3FFE0FFB889715.taxon	type_taxon	Type species. Scalaria algeriana Weinkauff, 1866; original designation by de Boury (1887 a: 11). Present-day, Mediterranean Sea.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF8DFF8DFF3FFE0FFB889715.taxon	diagnosis	Diagnosis. “ Size small; shape stocky, turbinate, conical; spire mediocrely elongated, somewhat gradate, with smooth, pointed protoconch; whorls very convex, subdivided, although the sutures do not appear perforated; axial lamellae very prominent, auriculate behind, fused from one whorl to the other in the depth of the sutures; spiral ornamentation composed of fairly tight grooves. Last whorl almost equal to half the total height, with convex, imperforate base, almost completely devoid of cords and auricle. Aperture rounded, fairly large, with split peristome, inner layer continuous, outer layer foliaceous and interrupted in the parietal region; labrum slightly oblique, with a spiny projection behind. ” (translated from Cossmann 1912: 35).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF8DFF8DFF3FFE0FFB889715.taxon	discussion	Discussion. Parviscala comprises small species with closed umbilicus, pointed lamellae and spiral cords in the axial interspaces (Weil et al. 1999: 213). Boreoscala Kobelt, 1902 [type species Scalaria greenlandica Perry, 1811; present-day, Greenland] also has spiral cords but differs in its much larger size, more robust shell, its broad lamellae with strongly rolled over crests and it has a peribasal cord (or row of nodes). Hirtoscala Monterosato, 1890 [type species Scalaria cantrainei Weinkauff, 1866 = Hirtoscala muricata (Risso, 1826); present-day, Mediterranean Sea] is morphologically similar but lacks spiral sculpture. The Middle Miocene Paratethyan species probably belong to the lineage leading to the extant Mediterranean Parviscala vittata (Jeffreys, 1884).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF8CFF8EFF3FF894FA249245.taxon	description	Figs 4 L, 39 C – E, 40 C	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF8CFF8EFF3FF894FA249245.taxon	materials_examined	Type material. Holotype, NHMW 2024 / 0278 / 0011, SL: 5.7 mm, MD: 2.2 mm, Bad Vöslau (Austria), Figs 4 L, 39 E 1 – E 2. Paratypes, NHMW 2024 / 0278 / 0012, SL: 6.3 mm, MD: 2.5 mm, Bad Vöslau (Austria), Figs 39 C 1 – C 2, 40 C. NHMW 2024 / 0278 / 0013, SL: 5.2 mm, MD: 2.2 mm, Bad Vöslau (Austria), Figs 39 D 1 – D 2. Type locality. Bad Vöslau (Austria), Vienna Basin. Type stratum. Baden Formation. Age. Middle Miocene, middle Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF8CFF8EFF3FF894FA249245.taxon	etymology	Etymology. Abditus (Latin for hidden) referring to the fact that the specimens were ‘ hidden’ in a lot of Parviscala lacinia sp. nov.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF8CFF8EFF3FF894FA249245.taxon	diagnosis	Diagnosis. Small, moderately slender turreted shell with turreted multispiral protoconch, convex whorls with rounded shoulder placed just above mid-whorl and moderately close-set, moderately raised, robust, weakly prosocline lamellae, obliquely aligned axially, occasionally with low indistinct trigonal spine along shoulder, weak spiral threads in axial interspaces.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF8CFF8EFF3FF894FA249245.taxon	description	Description. Shell small, moderately slender turreted (apical angle ~ 23 – 27 °), faintly coronate. Protoconch turreted, of four smooth, weakly convex whorls, blunt elevated nucleus. Teleoconch of at least seven strongly convex whorls, with weakly concave subsutural ramp, rounded shoulder, periphery placed just above mid-whorl. Suture deeply impressed, linear. Sculpture of moderately close-set, moderately raised, robust, weakly prosocline lamellae, obliquely aligned axially (12: 12: 12), fused across suture. Crest of lamellae moderately rolled over. Some lamellae with weak, low, trigonal spines at shoulder. Interspaces between lamellae glossy with weak spiral threads. Last whorl convex, attaining about 40 % of total height, lamellae persisting over base, strongly rolled over, no peribasal cord, base sub-angled, slightly convex. Aperture ovate, columella weakly concave. Outer lip not preserved. No umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF8CFF8EFF3FF894FA249245.taxon	discussion	Discussion. This species is reminiscent of Parviscala badensis sp. nov., which differs in its angled shoulder, more prominent spines, less fused lamellae, even weaker spiral sculpture and a pointed protoconch. Parviscala paratethyca sp. nov. has less convex whorls and is distinctly coronate. The extant Mediterranean Parviscala vaillanti (Jousseaume, 1912) is similar but differs in its strongly prosocline strongly raised lamellae and deeply impressed suture (see Bonfitto et al. 2021: fig. 1). Paleoenvironment. Occurrences in the Baden Formation of the Vienna Basin suggest middle to outer neritic settings in up to 250 m water depth (Kranner et al. 2021).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF8CFF8EFF3FF894FA249245.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienna Basin: Baden (Austria) (hoc opus).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF8EFF8FFF3FFF2EFA249245.taxon	description	Figs 4 M, 39 B, 40 A	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF8EFF8FFF3FFF2EFA249245.taxon	materials_examined	Type material. Holotype, NHMW 2024 / 0278 / 0014, SL: 5.5 mm, MD: 2.0 mm, Bad Vöslau (Austria), Figs 4 M, 39 B 1 – B 2, 40 A. Type locality. Baden (Austria), Vienna Basin. Type stratum. Baden Formation. Age. Middle Miocene, middle Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF8EFF8FFF3FFF2EFA249245.taxon	etymology	Etymology. After the city Baden in Lower Austria.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF8EFF8FFF3FFF2EFA249245.taxon	diagnosis	Diagnosis. Small, moderately slender, weakly coronate with turreted protoconch, strongly convex teleoconch whorls with rounded shoulder angulation, thin robust lamellae, partly aligned axially, sometimes pointed at shoulder, delicate spiral cords in interspaces, sub-angled base.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF8EFF8FFF3FFF2EFA249245.taxon	description	Description. Shell small, moderately slender turreted (apical angle ~ 25 °), weakly coronate. Protoconch turreted, of four smooth, weakly convex whorls, small elevated nucleus. Teleoconch of seven strongly convex whorls, with weakly concave subsutural ramp, rounded shoulder, periphery placed above mid-whorl. Suture deeply impressed, weakly undulating. Sculpture of moderately close-set, moderately raised, thin but robust, weakly prosocline lamellae, partly aligned axially (10: 10: 10). Crest of lamellae only weakly rolled over. Some lamellae with low, trigonal spines at shoulder. Interspaces between lamellae glossy with delicate cords. Last whorl large, convex, attaining about 38 % of total height, lamellae persisting over base, strongly rolled over, no peribasal cord, base sub-angled, slightly convex. Aperture ovate, columella weakly concave. Outer lip not preserved. Columellar callus forming broad rim, detached from base, parietal callus delicate. No umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF8EFF8FFF3FFF2EFA249245.taxon	discussion	Discussion. Parviscala paratethyca sp. nov. is superficially reminiscent of Parviscala badensis sp. nov. but has a less incised suture, less convex whorls and its spines are closer to the adapical suture. Paleoenvironment. Occurrences in the Baden Formation of the Vienna Basin suggest middle to outer neritic settings in up to 250 m water depth (Kranner et al. 2021).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF8EFF8FFF3FFF2EFA249245.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienna Basin: Baden (Austria) (hoc opus).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF8EFF89FF3FFB3EFADE9309.taxon	description	Figs 4 N – O, 40 B, 41 A – H	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF8EFF89FF3FFB3EFADE9309.taxon	materials_examined	Type material. Holotype, NHMW 2018 / 0248 / 0551 a, SL: 4.9 mm, MD: 2.0 mm, CoŞteiu de Sus (Romania), Figs 41 A 1 – A 2. Paratypes, NHMW 2018 / 0248 / 0551 b, SL: 3, 1 mm, MD: 1.7 mm, CoŞteiu de Sus (Romania), Figs 4 N, 41 C 1 – C 2. NHMW 2018 / 0248 / 0551 c, SL: 5.2 mm, MD: 2.1 mm, CoŞteiu de Sus (Romania), Figs 41 D 1 – D 2. NHMW 1855 / 0045 / 0206 a, SL: 6.1 mm, MD: 2.6 mm, Steinebrunn (Austria), Figs 39 B, 41 E 1 – E 2. NHMW 1855 / 0045 / 0206 b, SL: 6.8 mm, MD: 2.9 mm, Steinebrunn (Austria), Figs 40 B 1 – B 2. NHMW 1887 / 0018 / 0017, SL: 7.8 mm, MD: 4.1 mm, Steinebrunn (Austria). NHMW 2024 / 0278 / 0016, SL: 4.4 mm, MD: 1.7 mm, Bad Vöslau (Austria), Figs 41 F 1 – F 2. NHMW 2024 / 0278 / 0017, SL: 3.1 mm, MD: 1.4 mm, Bad Vöslau (Austria), Fig. 41 G. NHMW 2024 / 0278 / 0018, SL: 3.7 mm, MD: 1.6 mm, Bad Vöslau (Austria), Figs 41 H 1 – H 2. NHMW 1887 / 0018 / 0017, SL: 7.8 mm, MD: 4.1 mm, Steinabrunn (Austria), Fig. 4 O. Additional material. 52 spec., NHMW 1864 / 0001 / 0694, Steinebrunn (Austria). Type locality. CoŞteiu de Sus (Romania), Făget Basin. Type stratum. Silt and clay of the Dej Formation. Age. Middle Miocene, Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF8EFF89FF3FFB3EFADE9309.taxon	etymology	Etymology. After lacinia (Latin for corner, end, tip, point), referring to the small shoulder spines; noun in apposition.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF8EFF89FF3FFB3EFADE9309.taxon	diagnosis	Diagnosis. Small, moderately broad, turriculate, gradate, coronate with convex to subcylindrical whorls, narrow subsutural shelf, robust, strongly raised, weakly prosocline lamellae forming short spines along shoulder, delicate spiral sculpture.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF8EFF89FF3FFB3EFADE9309.taxon	description	Description. Shell small, moderately broad, gradate (apical angle ~ 27 – 29 °), coronate. Protoconch turreted of 4.5 smooth, weakly convex whorls, last whorl with marked mid-whorl angulation, nucleus moderately elevated dome. Teleoconch of five whorls, with narrow subsutural shelf and weakly convex to subcylindrical periphery. Suture deeply impressed. Sculpture of moderately close-set, robust, strongly raised, weakly prosocline lamellae, only partly obliquely aligned axially (9 / 11: 13: 13), adapical tips forming robust, trigonal, strongly pointed spines along shoulder. Crests of lamellae slightly rolled over on spire whorls, strongly rolled over on last whorl. Interspaces between lamellae with broad, low spiral cords. Last whorl attaining about 40 % of total height, lamellae persisting over base, strongly rolled over, flattish, no peribasal cord, base slightly concave. Aperture ovate to subcircular, columella weakly concave. Outer lip thickened by prominent varix. Columellar callus forming broad rim, detached from base, parietal callus delicate. Peristome duplex with inner layer covering outer layer largely. No umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF8EFF89FF3FFB3EFADE9309.taxon	discussion	Discussion. Parviscala lacinia sp. nov. is highly reminiscent of the extant Mediterranean Parviscala vittata (Jeffreys, 1884), which is only slightly larger (SL: 7 mm). It differs from the Paratethyan species in its slenderer shell and more convex whorls (see Weil et al. 1999: fig. 108; Bouchet & Warén 1986: figs 1193 – 1194). The protoconch of Parviscala vittata agrees with that of P. lacinia in its high conical outline with slightly angled last protoconch whorl (see Bouchet & Warén 1986: fig. 1174; https: // www. gruppomalacologicoscalaria. org / atlas / epitonium-vittatumjeffreys- 1884 /). The extant Atlanto-Mediterranean Parviscala algeriana (Weinkauff, 1866), the type species of Parviscala, has a much broader shell, more lamellae and weaker spines (see Bouchet & Warén 1986: figs 1188 – 1190). The extant eastern Pacific Parviscala deifica (Melvill & Standen, 1903) and Parviscala eusculpta (Sowerby, 1903) are both larger and differ in the greater number of lamellae (P. eusculpta) or the wider apical angle (P. deifica) (see Weil et al. 1999: figs 316, 317; Brown & Raines 2004: fig. 1). Parviscala thyraea (Kilburn, 1985), from South Africa, differs from P. lacinia in its weak shoulder angulation and much weaker spines (see Kilburn 1985: fig. 139; Weil et al. 1999: fig. 169). Parviscala climacotum (Kilburn, 1985) and P. columba (Kilburn, 1985), both extant species from South Africa, differ from the Paratethyan species in their broader shells and fewer lamellae with weaker spines (see Kilburn 1985: figs 133, 136; Weil et al. fig. 163). Paleoenvironment. The common occurrence at Steinebrunn (Austria) suggests shallow marine environments with sea grass (own data M. H.).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF8EFF89FF3FFB3EFADE9309.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Polish-Ukrainian Fore-Carpathian Basin: Shushkivtsi (= Szuszkowce) (Ukraine) (Friedberg 1928); Vienna Basin: Steinebrunn (Austria) (hoc opus); Pannonian Basin: Hidas, Szokolya, Várpalota, (Hungary) (Strausz 1966); Southern Pannonian Basin: Jazovac, Hrvaćani (Bosnia and Herzegovina) (Atanacković 1985); Făget Basin: CoŞteiu de Sus (Romania) (hoc opus).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF88FF8AFF3FFA73FB549073.taxon	description	Figs 39 A, 40 D	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF88FF8AFF3FFA73FB549073.taxon	materials_examined	Type material. Holotype, NHMW 1865 / 0001 / 0583, SL: 12.0 mm, MD: 4.0 mm, Steinebrunn (Austria), Figs 39 A 1 – A 2, 40 D. Type locality. Steinebrunn (Romania), Vienna Basin. Type stratum. Baden Formation. Age. Middle Miocene, middle Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF88FF8AFF3FFA73FB549073.taxon	etymology	Etymology. Referring to the occurrence in the Paratethys Sea.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF88FF8AFF3FFA73FB549073.taxon	diagnosis	Diagnosis. Medium, - sized, moderately slender, turriculate, coronate with convex whorls, strongly raised, weakly prosocline, rolled over lamellae forming short spines close to adapical suture, delicate spiral sculpture in interspaces.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF88FF8AFF3FFA73FB549073.taxon	description	Description. Shell medium-sized, moderately slender, turriculate (apical angle ~ 22 °), coronate. Protoconch unknown. Teleoconch of at least eight convex whorls, periphery placed just below mid-whorl. Suture deeply impressed. Sculpture of moderately close-set, robust, strongly raised, weakly prosocline lamellae, only partly obliquely aligned axially (11: 12: 12), adapical tips forming small, trigonal spines close to adapical suture. Crests of lamellae rolled over. Interspaces between lamellae with low spiral cords. Last whorl attaining about 38 % of total height, lamellae persisting on base, strongly rolled over, flattish, no peribasal cord or basal angulation, base slightly convex. Aperture ovate, columella concave. Outer lip not preserved. Columellar callus forming broad rim, detached from base, parietal callus delicate. No umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF88FF8AFF3FFA73FB549073.taxon	discussion	Discussion. Parviscala paratethyca sp. nov. differs from Parviscala lacinia sp. nov., with which it co-occurs at Steinebrunn (Austria), in its much larger size and greater number of teleoconch whorls that are more regularly convex and lack a subsutural shelf, and the much weaker spines. The Pliocene Mediterranean Parviscala vidali (de Boury, 1911) might be a closely related species, which differs only in its slightly wider apical angle and more convex whorls (Chirli 2009: pl. 15, figs 12 – 17, pl. 16, figs 1 – 4). Among the recent species of Parviscala, the Mediterranean Parviscala vittata (Jeffreys, 1884), is most similar to Parviscala paratethyca sp. nov. It attains the same size but differs in the greater number of whorls, the more strongly convex early teleoconch whorls that bear numerous, close-set lamellae (see Weil et al. 1999: fig. 108; Raven et al. 2022: fig. G). All other extant species placed in Parviscala by Kilburn (1985) and Weil et al. (1999) differ significantly in their larger size, wider apical angle and / or gradate spire. Paleoenvironment. The common occurrence at Steinebrunn (Austria) suggests shallow marine environments with sea grass (own data M. H.).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF88FF8AFF3FFA73FB549073.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienna Basin: Steinebrunn (Austria) (hoc opus). Genus Punctiscala de Boury, 1890	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF88FF8AFF3FFA73FB549073.taxon	type_taxon	Type species. Scalaria plicosa R. A. Philippi, 1844; original designation. Pliocene, Italy.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF88FF8AFF3FFA73FB549073.taxon	diagnosis	Diagnosis. “ Solid shell. Small size; form and protoconch of Pliciscala; ornamentation composed of axial ribs, more or less thick, and spiral cords, a little depressed, in the intervals of which there are - in addition - fine rows of punctuations. Last whorl roughly equal to a quarter of the total height, angular at the periphery of the imperforate base, which is covered by a flattened disk, bordered by a peripheral carina and obtusely furrowed; this disk is often rudimentary, and bears no bead in the center. Circular aperture with split peristome, inner layer thin, outer layer reflective and varicose, oblique labrum. ” (translated from Cossmann 1912: 83).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF88FF8AFF3FFA73FB549073.taxon	discussion	Discussion. Punctiscala de Boury, 1890 is characterized by shells with prominent axial ribs and a broad, well-defined basal disc over which the axial ribs do not extend and a thickened outer lip (Landau et al. 2006: 54). In contrast to Gregorioiscala, its axial ribs are not aligned along the shell, and it rarely forms a varix (Bouchet & Warén 1986, Weil et al. 1999; e. g. Gregorioiscala sarsi (Kobelt, 1904) may form an occasional varix; pers. comm. Leon Hoffmann, 20. 03. 2025). This genus has not been documented from the Paratethys Sea so far. Both genera have pitted spiral microsculpture. Scalaria disjuncta Bronn, 1831 from the Upper Miocene and Pliocene of the Proto- / Mediterranean was placed in the genus Punctiscala by Sacco (1891), a position followed by subsequent authors (e. g., Landau et al., 2006; Brunetti & Cresti, 2018), but that species is much larger than most Punctiscala species and lacks punctate microsculpture, and should probably be placed in Clathroscala de Boury, 1890.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF8BFF94FF3FF9ECFE739411.taxon	description	Figs 35 G – H, 42 A – D	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF8BFF94FF3FF9ECFE739411.taxon	materials_examined	Type material. Holotype, SMF 380180 _ 12 - 1452 a (XII 1452 a), SL: 7.9 mm, MD: 3.0 mm, CoŞteiu de Sus (Romania), specimen illustrated in Zilch (1934: pl. 10, fig. 68), Fig. 42 A. Illustrated material. NHMW 2013 / 0300 / 0141, SL: 13.0 mm, MD: 4.7 mm, Baden-Sooss (Austria), Figs 35 H, 42 D. NHMW 2013 / 0078 / 0157 a, SL: 12.4 mm, MD: 4.2 mm, Baden (Austria), Figs 35 G, 42 B 1 – B 2. NHMW 2013 / 0078 / 0157 b, SL: 14.8 mm, MD: 3.9 mm, Baden (Austria), Figs 42 C 1 – C 2. Revised description. Shell medium-sized, moderately slender, turreted (apical angle ~ 16 – 20 °). Protoconch turreted of about five, moderately convex whorls (diameter 390 μm). Late whorls high, weakly convex, with numerous delicate axial riblets. Teleoconch of about ten convex whorls. Whorl profile with moderately steep, weakly concave subsutural ramp and convex periphery placed mid-whorl. Suture moderately impressed, linear. Sculpture of prominent, convex, moderately close-set, weakly prosocline axial ribs not aligned axially (~ 20: 18 / 20: 16 / 20), weakening slightly over subsutural ramp. Occasionally with one or two weak, more strongly prosocline varices per whorl. Spiral sculpture of about seven primary spiral cords and several weaker secondary cords separated by slightly narrower, weakly punctate interspaces. Typically one or rarely two rows between spiral cords. Punctate microsculpture most prominent on subsutural ramp but occurring more or less distinctly over entire whorl. Spiral cords most prominent in interspaces, weaker over ribs. Last whorl attaining ~ 33 % of total height, basal angulation coinciding with weak to moderately prominent peribasal cord, base weakly convex to nearly flat bearing weak axial ribs and weak spiral cords. Aperture subcircular to ovate, columella weakly concave. Outer lip thickened by terminal varix without forming broad peristome. Columellar callus and parietal callus forming continuous, narrow, attached rim. Peristome duplex but relatively thin with glossy inner layer. No umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF8BFF94FF3FF9ECFE739411.taxon	discussion	Discussion. The generic placement of this species was controversial. It was placed in various combinations in Pliciscala de Boury, 1887 [type species Scalaria gouldi Deshayes, 1861; Eocene, France], Punctiscala de Boury, 1890 and Opalia [type species Scalaria australis Lamarck, 1822; present-day, Australia]. Pliciscala gouldi, from the Eocene of the Paris Basin, has prominent axial ribs but strongly reduced spiral sculpture. Its peristome is duplex and the base is convex (Deshayes 1861: pl. 11, figs 15 – 16). Similarly, Pliciscala obliqua Garvie, 2021, from the Eocene of Texas, lacks spiral cords and lacks a basal angulation (Garvie 2021, fig. 1 A – B). Punctiscala brandenburgi (Boettger, 1906) differs in its distinct spiral sculpture, the basal angulation and the weakly thickened peristome. Therefore, a close relation with Pliciscala is unlikely. Opalia is a problematic genus in which morphologically quite different species are currently placed. The type species Opalia australis is conical with strongly raised, convex axial ribs, lacks spiral sculpture and has a very prominent peribasal cord from which the axial ribs develop. Its peristome is not very thickened. The same morphology is seen in Opalia wroblewskyi (Mörch, 1875), Opalia montereyensis (Dall, 1907), Opalia gereti (de Boury, 1913 a) and Opalia abbotti Clench & Turner, 1952. Several other species currently placed in Opalia by MolluscaBase (2024 i), Weil et al. (1999), Garcia (2004) and Lima et al. (2012) differ in their pupoid shape and / or crenulated suture and might better be placed in Nodiscala de Boury, 1890, Dentiscala de Boury, 1887 or similar genera. It is beyond the scope of this paper to revise these extant species but in any case, Punctiscala brandenburgi is very different from Opalia sensu stricto and we reject its placement in Opalia. Paleoenvironment. Occurrences in the Baden Formation of the Vienna Basin suggest middle to outer neritic settings in up to 250 m water depth (Kranner et al. 2021).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF8BFF94FF3FF9ECFE739411.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Korytnica Basin: Korytnica (Poland) (Bałuk 1975); Vienna Basin: Baden, Baden-Sooss (Austria) (hoc opus); Novohrad-Nógrád Basin: Nógrádszakál (Hungary) (Bogsch 1936); Bükk Mountains: Balaton (Hungary) (Csepreghy-Meznerics 1969 a); Făget Basin: CoŞteiu de Sus (Romania) (Boettger 1906).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF95FF95FF3FFD0BFCB69101.taxon	description	Figs 40 F, 43 A 1 – A 2	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF95FF95FF3FFD0BFCB69101.taxon	materials_examined	Type material. Holotype, NHMW 1871 / 0010 / 0126, SL: 9.2 mm, MD: 3.1 mm, Porzteich at Břeclav (Czech Republic), Figs 40 F, 43 A 1 – A 2. Type locality. Porzteich at Břeclav (Czech Republic), Vienna Basin. Type stratum. Baden Formation. Age. Middle Miocene, middle Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF95FF95FF3FFD0BFCB69101.taxon	etymology	Etymology. Referring to Moravia, a historical region in the east of the Czech Republic.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF95FF95FF3FFD0BFCB69101.taxon	diagnosis	Diagnosis. Small, slender shell with subcylindrical whorls, deeply incised suture, strongly raised, prominent axial ribs overrun by six much weaker spiral cords, aperture subcircular, small, duplex.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF95FF95FF3FFD0BFCB69101.taxon	description	Description. Shell small, slender (apical angle ~ 17 °). Protoconch unknown. Teleoconch of about nine whorls. Whorl profile weakly convex to subcylindrical, vaguely biangular along shoulder and close above abapical suture. Suture deeply impressed, slightly undulating. Sculpture of prominent, strongly raised, narrow, orthocline axial ribs, not aligned axially (9: 11: 12), separated by wider interspaces, overrun by up to six broad, low spiral cords, separated by wider interspaces. Spiral cords much weaker than ribs, forming rectangular sculpture. Last whorl attaining about 35 % of total height, basal angulation delimited by prominent peribasal cord, basal disc with slightly scalloped margin, bearing about five prominent cords. Aperture small, subcircular. Outer lip strongly thickened. Columellar and parietal callus forming continuous, narrow rim. Peristome duplex. No umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF95FF95FF3FFD0BFCB69101.taxon	discussion	Discussion. Punctiscala moravica sp. nov. is reminiscent of Punctiscala plicosa (Philippi, 1844), from the Late Miocene and Pliocene of the Mediterranean Sea but differs in its smaller size, subcylindrical whorls, less convex axial ribs and thicker peristome (see Landau et al. 2006: pl. 17, figs 1 – 5). The extant Mediterranean Punctiscala cerigottana (Sturany, 1896) may develop a comparable outline but has much more prominent, strongly raised, convex axial ribs and much finer spiral sculpture. Paleoenvironment. Unknown.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF95FF95FF3FFD0BFCB69101.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienna Basin: Porzteich at Břeclav (Czech Republic) (hoc opus). Genus Scala Mörch, 1852	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF95FF95FF3FFD0BFCB69101.taxon	type_taxon	Type species. Turbo clathrus Linnaeus, 1758; by tautonymy of replaced name [Clathrus]; present-day, Mediterranean Sea.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF95FF95FF3FFD0BFCB69101.taxon	diagnosis	Diagnosis. Small to medium-sized, solid, slender to moderately slender conical shell with convex whorls, deeply impressed suture. Sculpture of very prominent, widely spaced lamellae with successively more rolled over crests during ontogeny, lamellae distinctly obliquely axially aligned, fused across suture resulting in deeply fenestrate suture. Axial interspaces glossy, smooth or with faint spiral threads. Last whorl convex, no peribasal cord, lamellae strongly rolled over on base. Short, broad fasciole. Aperture subcircular, peristome thickened by terminal varix, no umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF95FF95FF3FFD0BFCB69101.taxon	discussion	Discussion. Scala was first used by Klein (1753), which is a pre-Linnean work without nomenclatorial status. Later, Mörch (1852: 48) made Scala available as a substitute name for “ Clathrus Ocken [sic] ” and listed [Scalaria (Scala)] clathrus Linnaeus as species, which thus become the type species (see also discussion in Bouchet & Warén 1986: 499). Clathrus was originally introduced by Oken (1815: 255) but his work was placed on the Official Index by Opinion 417 (ICZN 1956). Later, the name was made available by Agassiz (1837: 35). The complex nomenclatorial history was reviewed by Brown & Neville (2015: 21), who clarified the situation by the subsequent designation of Scalaria acuta J. Sowerby, 1812 from the Eocene of the British Isles as type species [for illustrations see Sowerby (1812: pl. 16, lower two figs), Duerr 2004: fig. 9 and Savazzi & Saski (2004: fig 11 A – B, H)]. Sacco (1891) used Clathrus for several Miocene species from Italy. Cossmann (1912: 36) gave a first detailed diagnosis for Clathrus but this was based on an incorrect type species [Scalaria communis Lamarck, 1822, = Epitonium clathrus (Linnaeus, 1758)]. Therefore, Clathrus sensu Cossmann 1912 is an objective synonym of Scala Mörch, 1852. This erroneous concept was also adopted by Weil et al. (1999). Therefore, the understanding of Clathrus varied greatly in the literature and no diagnosis was available so far for Clathrus sensu Brown & Neville (2015). This diagnosis is provided here: Medium-sized, moderately stocky conical, gradate, coronate, with weakly convex to subcylindrical whorls, moderately wide, weakly concave subsutural shelf. Suture deeply impressed. Moderately close-set, prominent, prosocline robust, strongly raised lamellae with rolled over crests; interspaces with broad, low, convex spiral cords. Lamellae not aligned across teleoconch disconnected along suture. Last whorl with sub-angled base without persisting peribasal cord. Lamellae prominent on base, strongly rolled over. Short, broad fasciole with prominent growth lines. Aperture subcircular, columellar callus forming broad rim detached from base, parietal callus weak, peristome thickened, duplex.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF97FF96FF3FFF2EFD5B91FA.taxon	description	Figs 43 B 1 – B 2	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF97FF96FF3FFF2EFD5B91FA.taxon	materials_examined	Type material. Holotype, BkK-G 1218, SL: 6.4 mm, MD: 2.5 mm, Korytnica (Poland), illustrated in Bałuk (2006: pl. 10, fig. 3), Figs 43 B 1 – B 2. Type locality. Korytnica (Poland), Korytnica Basin. Type stratum. Pińczów Formation. Age. Middle Miocene, middle Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF97FF96FF3FFF2EFD5B91FA.taxon	etymology	Etymology. Referring to Poland.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF97FF96FF3FFF2EFD5B91FA.taxon	diagnosis	Diagnosis. Shell small, moderately slender conical with strongly convex whorls and strongly raised lamellae obliquely aligned axially, fused across suture, resulting in deeply fenestrate suture.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF97FF96FF3FFF2EFD5B91FA.taxon	description	Description. Small, moderately slender conical shell (apical angle 22 °). Protoconch and first teleoconch whorls unknown. Teleoconch whorls strongly convex with periphery placed mid-whorl. Suture deeply impressed, fenestrate. Sculpture of prominent, strongly raised lamellae, obliquely aligned axially, fused but indented across suture (?: 8: 8). Crests of lamellae slightly rolled over. Interspaces with faint spiral threads. Last whorl attaining about 40 ° of total height, no basal angulation nor peribasal cord, base convex, lamellae strongly rolled over. Aperture not preserved.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF97FF96FF3FFF2EFD5B91FA.taxon	discussion	Discussion. Scala polonica sp. nov. is highly reminiscent of the extant Scala clathrus (Linnaeus, 1758) from which it differs in its smaller size and more indented lamellae across the suture (see Weil et al. 1999: fig. 110; Landau et al. 2006: pl. 7, figs 1 – 2; Raven & Suárez 2020: figs 5 A – C). Bałuk (2006) identified this species with Fuscoscala kunstleri De Boury in Cossmann, 1912, which was originally described from the Burdigalian of Saucats (France). The two are indeed similar, but Fuscoscala kunstleri differs from the Paratethyan species in its broad and steep subsutural ramp, the slightly angled periphery below mid-whorl and its lamellae are not axially aligned (De Boury in Cossmann 1912: pl. 2, figs 7 – 8; Cossmann & Peyrot 1922: pl. 3, fig. 80). Paleoenvironment. Unknown.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF97FF96FF3FFF2EFD5B91FA.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Korytnica Basin: Korytnica (Poland) (Bałuk 2006). Genus Sthenorytis Conrad, 1863	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF97FF96FF3FFF2EFD5B91FA.taxon	type_taxon	Type species. Scalaria expansa Conrad, 1842; subsequent designation by de Boury (1890: 190). Miocene, Maryland, USA.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF97FF96FF3FFF2EFD5B91FA.taxon	diagnosis	Diagnosis. “ Thick, massive shell. Size fairly large; shape turbinate, conical; spire short, not gradate, with disjointed, very convex, low whorls; axial lamellae strong and prominent, oblique, reflexed, striated, bent backwards where they form a uneven festoon before welding - over the sutures - to the lamellae of the previous whorl; in the intervals, the sutures are visibly perforated and bordered above by a spiral cord which detaches from the left side of each lamella; in addition, in most species, the surface of the whorls is adorned with broad, obtuse spiral cords which rise up on the left side of each lamella, and which are separated by obsolete, narrower grooves. Last whorl approximately equal to two-thirds of total height, rounded at the base, which is loosely bounded by a discontinuous cord between the lamellae; the surface of the base is almost completely covered by the reflexed lamellae, up to the center where they fuse together entirely, leaving no umbilical left. Peristome subcircular, inclined in an oblique plane at 25 ° to the vertical axis; inner layer rather calloused, not distinctly auriculated anteriorly; outer layer reflected outward over the entire extent of the labrum, with a broad posterior expansion that folds back strongly before fusing with the lamella of the previous whorl. ” (translated from Cossmann 1912: 44 – 45).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF97FF96FF3FFF2EFD5B91FA.taxon	discussion	Discussion. The name of this genus was frequently misspelled. Sthenorhytis in Dall (1889) and Stenorhytis in Cossmann (1912) are incorrect subsequent spellings.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF96FF90FF3FFF2EFD9D944D.taxon	description	Figs 44 C	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF96FF90FF3FFF2EFD9D944D.taxon	materials_examined	Type material. Holotype, BSPG 2007 X 40, SL: 31.5 mm, MD: 23.0 mm, Gurlan (Germany), illustrated in Schneider et al. (2009: figs 5 d – e), Fig. 44 C. Stored in Bayerische Staatssammlung für Paläontologie und Geologie, Munich (Germany). Type locality. Gurlan (Germany), North Alpine Foreland Basin. Type stratum. Upper Marine Molasse. Age. Early Miocene, early Ottnangian (mid-Burdigalian).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF96FF90FF3FFF2EFD9D944D.taxon	etymology	Etymology. In honor of Simon Schneider (CASP, Cambridge), who reported this species for the first time.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF96FF90FF3FFF2EFD9D944D.taxon	diagnosis	Diagnosis. Very large, robust, very broad, stocky, gradate, weakly coronate shell with subcylindrical whorls and prominent, uniform, strongly rolled over lamellae separated by slightly wider interspaces and numerous prominent spiral threads.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF96FF90FF3FFF2EFD9D944D.taxon	description	Description. Shell very large, robust, very broad, stocky, gradate (apical angle ~ 65 °), weakly coronate. Protoconch and early teleoconch whorls unknown. Teleoconch whorls broad, subcylindrical with broad subsutural shelf and angled shoulder, separated by deeply impressed suture. Sculpture of prominent, strongly raised, uniform, weakly prosocline, internally lamellose lamellae forming indistinct, short spines along shoulder. Crests of lamellae strongly rolled over. Interspaces between lamellae broader than lamellae, with numerous narrow, prominent spiral threads. Last whorl attaining about 60 % of total height, slightly convex, basal angulation weak, relatively narrow base with crowded, nearly amalgamated lamellae. Fasciole short, broad with prominent growth lines. Aperture subcircular, Columella concave. Columellar and parietal callus form continuous narrow rim detached from base. Peristome strongly thickened, duplex. Umbilicus reduced to narrow chink.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF96FF90FF3FFF2EFD9D944D.taxon	discussion	Discussion. This species was erroneously identified by Schneider et al. (2009) as Sthenorytis proglobosa (Sacco, 1891), originally described from the Burdigalian and Langhian of the Colli Torinesi (Italy). The two are indeed similar but Sthenorytis simoni sp. nov. differs from the Italian species in its subcylindrical whorls and broad subsutural shelf. In addition, S. proglobosa lacks a basal angulation (Sacco 1891: pl. 1, figs 62 – 67; Ferrero-Mortara et al. 1984: pl. 5, figs 1 a – b). Sthenorytis trochiformis (Brocchi, 1814), from the Pliocene of the Mediterranean Sea, differs in its convex whorls, large, globose last whorl and lamellose varices (see Manganelli et al. 2011: figs 2 – 5). Paleoenvironment. Agitated rocky shore and coastal environments (Schneider et al. 2009).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF96FF90FF3FFF2EFD9D944D.taxon	distribution	Distribution in Central Paratethys. Ottnangian (Early Miocene): North Alpine Foreland Basin: Gurlan (Germany) (Schneider et al. 2009).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF91FF91FF3FFD37FC9494D9.taxon	description	Figs 44 D – E	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF91FF91FF3FFD37FC9494D9.taxon	materials_examined	Type material. Lectotype (designated herein), BS. 030.07.005, SL: 24 mm, MD: 21 mm, Stazzano (Italy), illustrated in Sacco (1891: pl. 1, figs 53 a – b). The holotype of Sthenorytis acutispina (Sacco, 1891) was not listed by Ferrero Mortara et al. (1984) and might be lost. Illustrated material. M. 62.6660, SL: 23 mm, MD: 21 mm, Sámsonháza (Hungary), illustrated in Csepreghy-Meznerics (1954: pl. 2, fig. 16), Fig. 44 D 1 – D 2. M. 61.4046, SL: 24 mm, MD: 23 mm, Sámsonháza (Hungary), illustrated in Csepreghy-Meznerics (1954: pl. 2, fig. 20), Figs 44 E 1 – E 2. Revised description. Medium-sized, solid, very broad conical, gradate, weakly coronate shell (apical angle 65 – 70 °). Protoconch and first teleoconch whorls unknown. Early teleoconch whorls gradate with broad subsutural shelf, marked but slightly rounded shoulder, subcylindrical periphery. Last whorl large, inflated, relatively low with broad, shallow subsutural ramp, prominent shoulder and convex periphery. Sculpture of very broad, strongly rolled over lamellae with smooth tops separated by wide interspaces (10 – 12). Small rounded spines on lamellae along shoulder. Spiral sculpture of weak primary cords with single delicate secondary intercalated. No basal angulation, peribasal cord weak, only visible in axial interspaces. Base weakly convex, nearly flat with prominent lamellae. Fasciole very broad, short with prominent growth lines. Aperture subcircular, columella concave. Outer lip strongly thickened. Peristome duplex. No umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF91FF91FF3FFD37FC9494D9.taxon	discussion	Discussion. This species was first described as Scalaria retusa var. spinosa by Bellardi & Michelotti (1840). De Gregorio (1889: 10) introduced Scalaria retusa var. retuspina as new name for it without further explanation. Most probably, de Gregorio (1889) considered the name Scalaria spinosa to be preoccupied by Scalaria spinosa Bonelli as also discussed by Sacco (1891: 35). However, Scalaria spinosa Bonelli, refers to a name used in an unpublished catalogue for the collection in Torino (Italy) written by Franco Andrea Bonelli (1784 – 1830) around the year 1826. The name was made available by Sacco (1891: 26), for a species placed by him in Hirtoscala (as subgenus). Consequently, Scalaria spinosa Bellardi & Michelotti, 1840 has priority and Scalaria retuspina de Gregorio, 1889 is an unnecessary replacement name. As Scalaria spinosa Sacco, 1891 is surely not a Sthenorytis, a replacement name for this species is not required. Sthenorytis acutispina (Sacco, 1891), from the Tortonian of Stazzano (Italy), is probably a subjective junior synonym of Sthenorytis spinosa, based on a subadult or small specimen. The fragmentary specimens from Hungary fully agree with this acutispina - morph. Therefore, we tentatively treat it as conspecific. Sthenorytis paucicostata Sacco, 1891, from the Langhian of Monte dei Cappuccini (Italy), might also be a subjective junior synonym of Sthenorytis spinosa, from which it differs mainly in the more numerous and closer-set lamellae on the spire whorls. However, we have not seen the type specimens.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF91FF91FF3FFD37FC9494D9.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Pannonian Basin: Mátraverebély, Sámsonháza (Hungary) (Csepreghy-Meznerics 1954); Cserhát Mountains: Szilvásvárad (Hungary) (Strausz 1966); Mecsek Mountains: Háromaház (Hungary) (Strausz 1966).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF90FF91FF3FFD43FF5C9186.taxon	description	Figs 44 A 1 – A 2	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF90FF91FF3FFD43FF5C9186.taxon	materials_examined	Type material. Lectotype (designated herein): BS. 030.08.004 (stored in the Museo Regionale di Scienze Naturali Torino, Italy), Superga (Italy), illustrated in Sacco (1891: pl. 2, fig. 5). Illustrated material. Private collection, Anton Breitenberger (Bad Vöslau, Austria), SL: 29.8 mm, MD: 19.9 mm, Pernersdorf (Austria), North-Alpine Foreland Basin; Figs 44 A 1 – A 2.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF90FF91FF3FFD43FF5C9186.taxon	description	Description. Shell very large, stocky, robust (apical angle ~ 52 °), weakly coronate. Protoconch unknown. Early teleoconch whorls convex with close-set lamellae, faint spiral cords. Later teleoconch whorls with narrow, weakly concave subsutural ramp, weakly angled shoulder, moderately convex periphery placed just below mid-whorl. Suture deeply impressed. Sculpture of moderately close-set, prominent, strongly raised, weakly prosocline lamellae separated by wider interspaces, not aligned axially (19: 19: 13), not fused to lamellae on preceding whorl, weakly coronate along shoulder forming short trigonal spines. Crests of lamellae weakly rolled over. Two more prominent varices on last whorl with strongly rolled over crests. Axial interspaces with numerous delicate spiral cords. Last whorl attaining ~ 65 % of total height, base sub-angled delimited by weak peribasal cord running up lamellae, base weakly convex, lamellae only slightly weaker over base, only slightly rolled over. Fasciole short, broad, with very prominent growth lines, separated from peristome by narrow chink. Aperture circular. Columella concave. Outer lip strongly thickened by varix. Columellar and parietal callus forming continuous, narrow rim, detached from base. Peristome very broad, duplex. No umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF90FF91FF3FFD43FF5C9186.taxon	discussion	Discussion. This species was described by Sacco (1891: 43) as Cirsotrema? sthenorytoides and is the type species of Pseudosthenorytis Sacco, 1891 [Peudostenorhytis is an incorrect subsequent spelling by Cossmann (1912: 51)]. Later, Pseudosthenorytis was treated as synonym of Cirsotrema Mörch, 1852 by Wenz (1938) and as synonym of Sthenorytis Conrad, 1863 by de Boury in Cossmann (1912: 51), Eames (1952: 50) and Clench & Turner (1950: 224). We follow de Boury in Cossmann (1912). Sthenorytis sthenorytoides (Sacco, 1891) is slightly less depressed than many Sthenorytis species and has less prosocline axial ribs. Nevertheless, these features do not justify separation at genus level. Paleoenvironment. At Pernersdorf channels formed in middle to outer neritic environments and were filled, with allochthonous assemblages uniting coastal-mudflat faunas with inner neritic ones (own data M. H.). The preservation suggests that the species was part of assemblages from inner neritic environments.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF90FF91FF3FFD43FF5C9186.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): North Alpine - Carpathian Foreland Basin: Pernersdorf (Austria (hoc opus). Proto-Mediterranean Sea. Burdigalian (Early Miocene): Colli Torinesi: Superga, Albugnao (Italy) (Sacco 1891).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF93FF9CFF3FF921FB829481.taxon	description	Figs 40 G – H, 45 A – F	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF93FF9CFF3FF921FB829481.taxon	materials_examined	Type material. Holotype, NHMW 1864 / 0001 / 0615 a, SL: 4.7 mm, MD: 1.9 mm, Grund (Austria), Figs 40 G, 45 A 1 – A 2. Paratypes, NHMW 1851 / 0002 / 0070, SL: 7.2 mm, MD: 2.8 mm, Grund (Austria), Figs 40 H, 45 B 1 – B 2. NHMW 1864 / 0001 / 0615 b, SL: 6.5 mm, MD: 2.7 mm, Grund (Austria), Figs 45 C 1 – C 2. NHMW 2007 / 0086 / 0165 d, SL: 8.3 mm, MD: 3.4 mm, Baden (Austria), Figs 45 D 1 – D 2. NHMW 2007 / 0086 / 0165 e, SL: 8.5 mm, MD: 3.9 mm, Grund (Austria), Figs 45 E 1 – E 2. NHMW 2024 / 0279 / 0005, SL: 6.5 mm, MD: 3.5 mm, Grund (Austria), Figs 45 F 1 – F 2. Type locality. Grund (Austria), North Alpine-Carpathian Foreland Basin. Type stratum. Grund Formation. Age. Middle Miocene, early Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF93FF9CFF3FF921FB829481.taxon	etymology	Etymology. In honor of Anton Breitenberger (Bad Vöslau, Austria), who always supports us by providing access to his collection.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF93FF9CFF3FF921FB829481.taxon	diagnosis	Diagnosis. Small, moderately broad, turreted shell with large, slightly inflated last whorl and sculpture of numerous delicate, thin, weakly raised lamellae and numerous delicate spiral threads.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF93FF9CFF3FF921FB829481.taxon	description	Description. Shell small, moderately broad, elongate conical or weakly coeloconoid, turreted (apical angle ~ 24 – 26 °). Protoconch unknown. Teleoconch of at least eight strongly convex whorls with weakly concave subsutural ramp, faintly angled periphery placed mid-whorl. Suture deeply impressed, linear. Sculpture of numerous, delicate, weakly raised, weakly prosocline lamellae, separated by wider interspaces, slightly more prominent close to adapical suture (~ 25 / 30: ~ 25 / 35: ~ 30 / 40). Interspaces with numerous delicate spiral threads. Last whorl large, slightly inflated, attaining about 45 % of total height, with rounded shoulder angulation placed just above mid-whorl, weak basal angulation without peribasal cord, base weakly convex with distinct lamellae and numerous delicate spiral threads. Short, broad fasciole with prominent growth lines. Aperture ovate, columella weakly concave. Outer lip and basal lip thin. Columellar callus forming broad, thin, attached rim. No parietal callus. No umbilicus. Peristome simple.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF93FF9CFF3FF921FB829481.taxon	discussion	Discussion. This species was confused by Kecskeméti-Körmendy (1962) with the eastern Pacific Aspersiscala minuticosta (de Boury 1912), which differs in its deep and wide umbilicus (see de Boury 1912: pl. 7, fig. 1). Subuliscala banoni (Tournouër, 1874) and Subuliscala lagusensis (de Boury in Cossmann, 1912), from the Burdigalian of France, both differ from the Paratethyan species in their much slenderer shells (see Tournouër, 1874: pl. 9, fig. 4, Cossmann 1912: pl. 2, figs 12 – 13, 14 – 15; Cossmann & Peyrot 1922: pl. 4, figs 22 – 23, 35, 41 – 42, 49 – 50). Adiscoacrilla interposita (Sacco, 1891), from the Tortonian of the Po Basin, differs in its broader, conical shell and more prominent varices (see Ferrero Mortara et al. 1984: pl. 6, fig. 10). Subuliscala breitenbergeri sp. nov. is most similar to Subuliscala lagusensis (de Boury in Cossmann, 1912), from the Burdigalian of France, but differs in its weaker varices and slightly more coeloconoid spire (see Cossmann 1912: pl. 2, figs 14 – 15; Cossmann & Peyrot 1922: pl. 4, figs 49 – 50). Paleoenvironment. At the locality Grund channels formed in middle to outer neritic environments and were filled, with allochthonous assemblages uniting coastal-mudflat faunas with inner neritic ones (Zuschin et al. 2005; Roetzel 2009). The preservation suggests that the shells might have belonged to the middle and outer neritic assemblages.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF93FF9CFF3FF921FB829481.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene):? Korytnica Basin: Korytnica (Poland) (Bałuk 2006); North Alpine-Carpathian Foreland Basin: Grund (Austria) (hoc opus); Vienna Basin: Baden (Austria) (hoc opus); Pannonian Basin: Várpalota (Hungary) (Kecskeméti-Körmendy 1962).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF9DFF9EFF3FFCFBFEB89715.taxon	description	Figs 40 I, 46 B – C	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF9DFF9EFF3FFCFBFEB89715.taxon	materials_examined	Type material. Holotype, NHMW 2024 / 0279 / 0006, SL: 12.5 mm, MD: 4.9 mm, Grund (Austria), Figs 40 I, 46 B 1 – B 2. Paratype, NHMW 2024 / 0279 / 0007, SL: 10.5 mm, MD: 5.2 mm, Grund (Austria), Figs 46 C 1 – C 2. Type locality. Grund (Austria), North Alpine-Carpathian Foreland Basin. Type stratum. Grund Formation. Age. Middle Miocene, early Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF9DFF9EFF3FFCFBFEB89715.taxon	etymology	Etymology. Referring to the similarity with Subuliscala banoni (Tournouër, 1874)	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF9DFF9EFF3FFCFBFEB89715.taxon	diagnosis	Diagnosis. Medium-sized, moderately slender, turreted shell of evenly convex whorls, deeply impressed suture, numerous, very delicate, weakly raised, slightly sigmoidal lamellae with numerous delicate spiral threads in interspaces, basal angulation subobsolete.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF9DFF9EFF3FFCFBFEB89715.taxon	description	Description. Shell medium-sized, moderately slender, turreted (apical angle ~ 20 °). Protoconch and early teleoconch whorls unknown. Teleoconch of strongly and evenly convex whorls with indistinct subsutural ramp, periphery placed mid-whorl. Suture deeply impressed, linear. Sculpture of numerous, delicate, weakly raised, weakly sigmoidal lamellae, not aligned axially (?: ~ 30: ~ 35), slightly bent in apertural direction at adapical suture, separated by wider interspaces. Tips of varices slightly more prominent close to adapical suture, faintly alate, touching preceding whorl. Lamellae not strictly evenly spaced, occasionally reduced or slightly more prominent by amalgamation without forming distinct varices. Interspaces with numerous delicate spiral threads. Last whorl convex, attaining about 40 % of total height, basal angulation subobsolete, without peribasal cord, base weakly convex with distinct lamellae and numerous delicate spiral threads. Short, broad fasciole with prominent growth lines. Aperture elongate ovate, columella weakly concave. Outer lip and basal lip thin. Columellar callus forming moderately broad, thin rim attached to fasciole; no parietal callus. No umbilicus. Peristome simple.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF9DFF9EFF3FFCFBFEB89715.taxon	discussion	Discussion. Subuliscala pseudobanoni sp. nov. is reminiscent of Subuliscala banoni (Tournouër, 1874), from the Burdigalian of France, but differs in its more convex whorls, wider apical angle, and weaker lamellae (see Tournouër, 1874: pl. 9, fig. 4, Cossmann 1912: pl. 2, figs 12 – 13; Cossmann & Peyrot 1922: pl. 4, figs 22 – 23, 35, 41 – 42). Subuliscala lagusensis (de Boury in Cossmann, 1912), from the Burdigalian of France, differs in its angled whorls and regularly spaced and uniform lamellae (see Cossmann 1912: pl. 2, figs 14 – 15; Cossmann & Peyrot 1922: pl. 4, figs 49 – 50). Subuliscala pseudobanoni co-occurs with Subuliscala breitenbergeri sp. nov. but S. breitenbergeri has a narrower spire, a larger last whorl, and slightly stronger lamellae, although they are still relatively weak. Adiscoacrilla? orientalis (Friedberg, 1914) differs in its much more prominent varices and thickened peristome (see Friedberg 1928: pl. 38, fig. 24). Paleoenvironment. At the locality Grund channels formed in middle to outer neritic environments and were filled, with allochthonous assemblages uniting coastal-mudflat faunas with inner neritic ones (Zuschin et al. 2005; Roetzel 2009). The preservation suggests that the shells might have belonged to the middle and outer neritic assemblages.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF9DFF9EFF3FFCFBFEB89715.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): North Alpine-Carpathian Foredeep: Grund (Austria).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF9FFF98FF3FF88BFCC29245.taxon	description	Figs 46 D – H	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF9FFF98FF3FF88BFCC29245.taxon	description	[Plicicscala] T [urriscala]. torulosa subv. supervaricosa (De Greg.) — Sacco 1891: 77. [Plicicscala] T [urriscala]. torulosa var. viennincola (De Greg.) — Sacco 1891: 77. [Plicicscala] T [urriscala]. torulosa var. Rovasendae Sacc. — Sacco 1891: 77, pl. 2, fig. 81. [Plicicscala] T [urriscala]. torulosa var. apuniculata Sacc. — Sacco 1891: 77, pl. 2, fig. 82. [Plicicscala] T [urriscala]. torulosa var. atava Sacc. — Sacco 1891: 78, pl. 2, fig. 83. [Plicicscala] T [urriscala]. torulosa var. cingulata Sacc. — Sacco 1891: 78, pl. 2, fig. 83 bis. [Plicicscala] T [urriscala]. torulosa var. perconica Sacc. — Sacco 1891: 78, pl. 2, fig. 84. [Plicicscala] T [urriscala]. torulosa var. convexiuscula Sacc. — Sacco 1891: 78, pl. 2, fig. 85. [Plicicscala] T [urriscala]. torulosa var. acutinodosa Sacc. — Sacco 1891: 79, pl. 2, fig. 86. [Plicicscala] T [urriscala]. torulosa (Br.) var. supervaricosa Sacco. — Sacco 1904: 106, pl. 23, fig. 25.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF9FFF98FF3FF88BFCC29245.taxon	materials_examined	Type material. Lectotype MCSNM, nr. 4975, Museo Civico di Storia Naturale di Milano, Piacentino (Italy), Neogene. This specimen was erroneously listed as Holotype by Rossi-Ronchetti (1955: 141) but Pinna & Spezia (1978: 165) listed an additional syntype Nr. 5021 (erroneously as paratype). Therefore, the specimen illustrated by Brocchi (1814) represents a lectotype (by erroneous definition as holotype by Rossi-Ronchetti 1955) and the other specimen is a paralectotypes. Illustrated material. NHMW 1851 / 0013 / 0008, SL: 23.4 mm, MD: 9.0 mm, Bad Vöslau (Austria), holotype of Scalaria torulosa viennincola De Gregorio 1889, illustrated in Hörnes (1856: pl. 46, figs 13 a – b), Figs 46 E 1 – E 2. NHMW 1874 / 0024 / 0028 a, SL: 19.6 mm, MD: 7.0 mm, Baden-Sooss (Austria), Figs 46 D 1 – D 2. NHMW 1874 / 0024 / 0028 b, SL: 18.0 mm, MD: 6.1 mm, Baden-Sooss (Austria), Figs 46 F 1 – F 2. NHMW 1853 / 0016 / 0012, SL: 22.9 mm, MD: 8.0 mm, Bad Vöslau (Austria), Figs 46 G 1 – G 2. NHMW 1864 / 0001 / 0101, SL: 21.7 mm, MD: 7.8 mm, Baden (Austria), Figs 46 H 1 – H 2. Revised description. Shell large, slender conical (apical angle ~ 18 °). Protoconch unknown. Teleoconch of 12 whorls. Teleoconch whorls weakly convex, with narrow subsutural collar, steep, weakly concave subsutural ramp, periphery below mid-whorl to close above abapical suture. Suture weakly impressed, slightly undulating. Sculpture of moderately widely spaced, broad, convex, orthocline, slightly drop-shaped axial ribs, not aligned axially, fading over subsutural ramp (11: 10: 10). Last teleoconch whorls with two more prominent varices, causing somewhat more convex appearance. Spiral sculpture of numerous narrow spiral cords separated by narrower interspaces often with delicate secondary intercalated. Spiral cords overrunning axial ribs. Slightly more prominent spiral cord at abapical suture. Last whorl attaining ~ 30 % of total height, axial ribs terminate at basal angulation delimited by more prominent, wavy peribasal cord, followed by up to three weaker spiral cords and second more prominent wavey mid-basal cord, base weakly concave with several weak spiral cords. Fasciole very short to obsolete. Aperture relatively small, subcircular, prosocline; Columella concave. Outer lip strongly thickened by terminal varix, truncated abapically. Columellar and parietal callus forming narrow rim, detached from base. Peristome duplex. Umbilicus reduced to narrow chink.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF9FFF98FF3FF88BFCC29245.taxon	description	Paratethyan synonyms. Scalaria torulosa viennincola De Gregorio, 1889 was introduced by De Gregorio (1889) as name for Scalaria torulosa in Hörnes (1856: pl. 46, figs 13 a – b). We do not see any features which would allow a separation of the Paratethyan specimens and therefore, we treat Scalaria viennincola as subjective junior synonym of Turbo torulosa Brocchi, 1814.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF9FFF98FF3FF88BFCC29245.taxon	discussion	Discussion. Turriscala torulosa is a highly characteristic species due to its very solid shell, conical outline, broad varices on the last teleoconch whorls, and truncated basal lip. Sacco (1891) distinguished several subspecies based on differences in sculpture and outline, which we treat as subjective synonyms of Turriscala torulosa (Brocchi, 1814). This species is known from the Early Miocene to Pliocene in the Mediterranean Sea. Turriscala borealis Kautsky, 1925, from the Hemmoorian of the North Sea Basin, is a closely related species, but differs in its smaller size, comparatively straighter-sided whorls and fewer axial ribs on the last whorl (Kautsky 1925: pl. 6, fig. 36; Janssen 1967: pl. 12, fig. 7; Janssen 1984: pl. 50, fig 6). Turriscala cf. borealis Kautsky, 1925, from the Late Miocene of Germany, is also smaller, has broader whorls, more axial ribs on early whorls and has more prominent spiral sculpture (Janssen & Stein in press: pl. 25, figs 12 ‒ 13). Paleoenvironment. Occurrences in the Baden Formation of the Vienna Basin suggest middle to outer neritic settings in up to 250 m water depth (Kranner et al. 2021).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF9FFF98FF3FF88BFCC29245.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienna Basin: Baden, Bad Vöslau, Baden Sooss (Austria) (Hörnes 1856, hoc opus); Pannonian Basin: Buják (Hungary) (Strausz 1966); Mecsek Mountains (Hungary) (Strausz 1966); Dacian Basin: Opanec, Urovene (Bulgaria) (Kojumdgieva in Kojumdgieva & Strachimirov 1960). Proto-Mediterranean Sea. Burdigalian (Early Miocene): Colli Torinesi: Baldissero (Italy) (Sacco 1891). Langhian (Middle Miocene): Colli Torinesi: Albugnano (Italy) (Sacco 1891). Tortonian (Late Miocene): Po Basin: S. Agata, Stazzano, (Italy) (De Boury 1890; Sacco 1891, 1904; Pelosio 1966). Early Pliocene: Estepona (Spain), Italy (Pelosio 1966; Cavallo & Repetto 1992; Landau et al. 2006).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF99FF98FF3FFB77FD599057.taxon	materials_examined	Material. NMBE Nr. D 2874, SL: 12 mm, Aarwald at Belpberg, illustrated in Pfister & Wegmüller (2007: pl. 2, figs 13 – 14).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF99FF98FF3FFB77FD599057.taxon	discussion	Discussion. Very poorly preserved internal casts with remnants of the shells, which are unidentifiable even at generic level.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF99FF98FF3FFB77FD599057.taxon	distribution	Distribution in Central Paratethys. Eggenburgian / Ottnangian (Early Miocene): North Alpine Foreland Basin: Belpberg (Switzerland) (Pfister & Wegmüller 2007).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF99FF98FF3FF9C0FE0291DE.taxon	materials_examined	Material. SMNS 68634, SL: 4.8 mm, illustrated in Höltke (2009: fig. 37).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF99FF98FF3FF9C0FE0291DE.taxon	discussion	Discussion. A fragment of a relatively low last whorl with moderately closely set, strongly prosocline lamellae, and angled base. The specimen is unidentifiable but does not represent one of the species described herein.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF99FF98FF3FF9C0FE0291DE.taxon	distribution	Distribution in Central Paratethys. Ottnangian (Early Miocene): North Alpine Foreland Basin: Ursendorf (Germany) (Höltke 2009).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF98FF99FF3FFF2EFEF99464.taxon	materials_examined	Type material. Holotype, SL: 25 mm, MD: 12 mm, St. Egida (Croatia?), illustrated Cossmann (1912: pl. 5, fig. 13).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF98FF99FF3FFF2EFEF99464.taxon	discussion	Discussion. This species is documented by a single fragmentary specimen, showing a large, moderately slender conical shell with weakly impressed suture and blunt, cancellate sculpture of low, broad varices and six spiral cords of similar width. Cossmann (1912) gave St. Egida as locality and assumed Croatia as origin with question mark. We are not aware of any Miocene locality of this name in Croatia and therefore, locality data and age of this species remain unclear.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF98FF99FF3FFDDFFC0E934B.taxon	materials_examined	Type material. Holotype, specimen from Shushkivtsi (= Szuszkowce) (Ukraine) illustrated in Dubois de Montpéreux (1831: 43, pl. 2, figs 36 – 37). The collection is lost.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF98FF99FF3FFDDFFC0E934B.taxon	discussion	Discussion. The illustration of this dubious species is very schematic and shows a small specimen with a weakly cyrtoconoid spire and unusually low whorls with deeply impressed suture, moderately widely spaced, low, thin, nearly orthocline lamellae and smooth interspaces. The aperture is unusually large with a thin peristome. The description of Dubois de Montpéreux (1831) does not add any details. The illustration might depict a very poorly drawn Hirtoscala miofrondiculoides (Sacco, 1891). In that case, Scalaria duboisiana d’Orbigny, 1852 would have priority over Scalaria (Hirtoscala) miofrondiculoides (Sacco, 1891) but provisionally, we prefer to keep both taxa separate. Hirtoscala pseudoscalaris (Brocchi, 1814), from the Neogene of Italy, with which Dubois de Montpéreux (1831) identified the Ukrainian shell, differs in its higher whorls and less numerous lamellae (see lectotype in Pinna & Spezia 1978: pl. 53, fig. 3). Paleoenvironment. Probably shallow marine, inner neritic.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF98FF99FF3FFDDFFC0E934B.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Polish-Ukrainian Fore-Carpathian Basin: Shushkivtsi (= Szuszkowce) (Ukraine) (Dubois de Montpéreux, 1831).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF98FF9AFF3FFA35FD989715.taxon	materials_examined	Type material. Syntype, SL: 5 mm, MD: 1.5 mm, Zboriv (Ukraine), illustrated in Friedberg (1914: pl. 20, fig. 18).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF98FF9AFF3FFA35FD989715.taxon	discussion	Discussion. Friedberg (1914) illustrated a fragmentary specimen with weakly convex whorls, moderately incised suture and weak, orthocline axial lamellae and relatively narrowly ovate aperture. Although he formally established a new species, his intention was rather to find a new name for the species, which was erroneously identified by Dubois de Montpéreux (1831) as Scalaria pseudoscalaris. For this very specimen, d’Orbigny (1852) had already introduced a new name Scalaria duboisiana [see Hirtoscala duboisianum (d’Orbigny, 1852)]. In 1928, Friedberg noted that the specimen illustrated by him in 1914 was unidentifiable and illustrated another specimen, which he thought to be closer to the one depicted by Dubois de Montpéreux (1831). This new specimen in Friedberg (1928: pl. 38, fig. 22) is surely not conspecific with the specimen from (1914) and is also quite different from the illustration in Dubois de Montpéreux (1831) but represents Parviscala lacinia sp. nov. Therefore, the status of ‘ Scala ’ duboisi Friedberg, 1914 remains dubious. Paleoenvironment. Shallow marine, inner neritic.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF98FF9AFF3FFA35FD989715.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Polish-Ukrainian Fore-Carpathian Basin: Zboriv (Ukraine) (Friedberg 1938).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF9BFF9AFF3FFE0EFF479411.taxon	description	Epitoniids are very rare in the Miocene deposits of the eastern Paratethys, probably because of unfavourable ecological conditions. Here we list only the literature-based occurrences without critical review of the original material.	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF9BFF9AFF3FFD0BFA879276.taxon	discussion	Discussion. Ossipov (1932) illustrated two small, slender specimens (SL: 9.2 mm) with strongly convex whorls, deeply impressed suture and about 13 / 14 prominent, low lamellae obliquely axially aligned, with smooth and glossy interspaces. These features might justify a placement in Fuscoscala Sacco, 1890 [type species Scalaria tenuicostata Michaud, 1829; present-day, Mediterranean Sea].	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
03D787A4FF9BFF9AFF3FFD0BFA879276.taxon	distribution	Distribution in Eastern Paratethys. Konkian (Middle Miocene): Kerch Peninsula (Ukraine / Russia).	en	Harzhauser, Mathias, Landau, Bernard M. (2025): The Epitoniidae (Gastropoda, Epitonioidea) of the Miocene Central Paratethys Sea-an overwhelming diversity. Zootaxa 5630 (1): 1-142, DOI: 10.11646/zootaxa.5630.1.1, URL: https://doi.org/10.11646/zootaxa.5630.1.1
