taxonID	type	description	language	source
A67987E6FFA7676AF2BD010CFD00F93A.taxon	type_taxon	Type species. Bulla tornatilis Linnaeus, 1758; original designation by Montfort (1810: 314). Present-day, Mediterranean Sea.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFA7676AF2BD010CFD00F93A.taxon	discussion	Discussion. Acteon is a problematic group and proper generic placement of Recent taxa requires anatomical data. Therefore, we use Acteon not in a strict sense.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFA76765F2BD002FFD1CF86D.taxon	description	Figs 2 A – E	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFA76765F2BD002FFD1CF86D.taxon	materials_examined	Type material. Holotype, NHMW 1855 / 0045 / 0782 a, SL: 18.3 mm, MD: 8.8 mm, Grund (Austria), illustrated in Hörnes (1856: 506, pl. 46, figs 21 a – b), Figs 2 D 1 – D 2. Paratypes: NHMW 1855 / 0045 / 0782, SL: 8.4 mm, MD: 4.3 mm, Grund (Austria), Figs 2 A 1 – A 2. NHMW 1855 / 0045 / 0782 b, SL: 8.2 mm, MD: 4.3 mm, Grund (Austria), Figs 2 B 1 – B 2. NHMW 1855 / 0045 / 0782 c, SL: 18.5 mm, MD: 9.5 mm, Grund (Austria), Figs 2 C 1 – C 2. NHMW 1855 / 0045 / 0782 b, SL: 18.6 mm, MD: 9.2 mm, Grund (Austria), Figs 2 E 1 – E 2. Additional paratypes. 5 spec., NHMW 1855 / 0045 / 0782 d, Grund (Austria). Type locality. Grund (Austria). Type stratum. Grund Formation. Age. Early Badenian, Langhian, Middle Miocene.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFA76765F2BD002FFD1CF86D.taxon	etymology	Etymology. Ancient Greek anti - (against or opposite) and pinguis referring to Acteon pinguis (d’Orbigny, 1852).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFA76765F2BD002FFD1CF86D.taxon	diagnosis	Diagnosis. Large, moderately broad shell with moderately high spire and subparallel last whorl; sculpture of about 25 relatively uniform, prominent, broad, partly bifid, flattened spiral cords separated by weakly punctate grooves; single strong columellar fold.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFA76765F2BD002FFD1CF86D.taxon	description	Description. Shell large, relatively solid, moderately broad (SL / MD = 1.9 – 2.0), with moderately high spire and broad apex (MD / ApD = 1.3 – 1.4). Protoconch small, depressed of one smooth whorl. Teleoconch of up to five convex whorls separated by narrow, deeply impressed, linear suture. First teleoconch whorl subcylindrical, smooth. Later spire whorls moderately convex, smooth with periphery above abapical suture, apical angle 66 – 81 °. Sculpture of about six broad, flattened spiral cords separated by punctate grooves; spiral cords narrowing adapically, occasionally bifid. Last whorl 78 – 83 % of total height, subcylindrical to weakly convex with weak, rounded shoulder, base moderately constricted, with periphery placed mid-whorl (SL / PH = 1.9 – 2.0). Sculpture of about 25 broad, flattened, spiral cords, separated by weakly punctate grooves. Cords often bifid, raising number of spiral cords. Spiral sculpture not weakening mid-whorl. Aperture 65 – 71 % of total height, moderately wide (MD / MAW = 2.4); outer lip simple, weakly concave mid-whorl, strongly rounded and slightly flared abapically. Columella thickened, slightly erect abapically, with single strong fold. Columellar callus below columellar fold forming broad, detached rim; columellar callus above columellar fold and parietal callus forming thin, delicate sheet.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFA76765F2BD002FFD1CF86D.taxon	discussion	Discussion. The Pliocene to Recent Acteon tornatilis (Linnaeus, 1758) differs in its more numerous and narrower spiral cords, much weaker shoulder, and more convex last whorl (see Cossignani & Ardovini 2011: 36, 368, unnumbered fig.; Gofas et al. 2011: 352, unnumbered fig.; Landau et al. 2023: pl. 21). Acteon pinguis (d’Orbigny, 1852), from the Aquitanian and Burdigalian of France, is very similar but differs in its smaller size (SL up to 13 mm), weaker columellar fold and more prominently punctate spiral grooves (see Peyrot 1932: pl. 11, figs 55 – 57, pl. 14, fig. 12). Some authors (Sieber 1958; Strausz 1966; Csepreghy-Meznerics 1972) considered this Paratethyan species to be conspecific with Acteon inflatus (Borson, 1821). As discussed by Landau et al. (2023: 367), A. inflatus is a poorly known Italian Pliocene species that has rarely been illustrated. However, Acteon antipinguis sp. nov. clearly differs from the holotype of A. inflatus (figured by Pavia 1976 b: pl. 2, fig. 13) in being considerably slenderer and higher spired. Paleoenvironment. At Grund (Austria) fossiliferous channel fills, which formed in middle to outer neritic environments, bearing allochthonous assemblages uniting coastal-mudflat faunas with inner neritic ones occur (Zuschin et al. 2005; Roetzel 2009). Preservation of the fossil shells suggests that the shells belonged to the allochthonous shallow water fauna.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFA76765F2BD002FFD1CF86D.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene):? Polish-Ukrainian Fore-Carpathian Basin: Wieliczka (Poland) (Reuss 1867); North Alpine Foreland Basin: Grund (Austria) (Hörnes 1856); Pannonian Basin: Várpalota (Hungary) (Strausz 1966); Bükk Mountains: Borsodbóta (Hungary) (Csepreghy-Meznerics 1972); Făget Basin: Coşteiu de Sus (Romania) (Boettger 1902).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFA96766F2BD0275FC28FF03.taxon	description	Figs 3 A – C	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFA96766F2BD0275FC28FF03.taxon	materials_examined	Type material. Holotype, NHMW 2018 / 0248 / 0485, SL: 3.3 mm, MD: 1.6 mm, Coșteiu de Sus (Romania), Figs 3 A 1 – A 2. Paratype, NHMW 2018 / 0248 / 0485 a, SL: 3.8 mm, MD: 1.7 mm, Coșteiu de Sus (Romania), Figs 3 B 1 – B 2. NHMW 2018 / 0248 / 0485 b, SL: 4.1 mm, MD: 1.8 mm, Coșteiu de Sus (Romania), Figs 3 C 1 – C 2. Type locality. Coșteiu de Sus (Romania), Făget Basin. Type stratum. Silt and clay of the Dej Formation. Age. Middle Miocene, middle Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFA96766F2BD0275FC28FF03.taxon	etymology	Etymology. Latin gutta (= drop), referring to the shape; noun in apposition.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFA96766F2BD0275FC28FF03.taxon	diagnosis	Diagnosis. Small, moderately slender ovate, glossy, drop-shaped shell with conical spire; sculpture of numerous, hardly visible, close-set spiral cords on spire whorls and ad- and abapical portions of last whorl leaving smooth mid-portion; columella with prominent fold.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFA96766F2BD0275FC28FF03.taxon	description	Description. Shell small, moderately slender ovate, drop-shaped (SL / MD = 2.0 – 2.4), with moderately high spire and broad apex (MD / ApD = 1.3 – 1.4). Protoconch low, tilted, of one smooth, convex whorl. Teleoconch of up to four whorls separated by narrow, weakly incised, canaliculate, linear suture. Spire whorls nearly straight sided, glossy, with periphery just above abapical suture, apical angle 45 – 48 °. Sculpture of numerous, faint, close-set spiral cords. Last whorl 73 – 80 % of total height, weakly convex, with periphery placed mid-whorl (SL / PH = 2.0). Sculpture of weak, moderately broad, flattened, spiral cords on base, separated by faintly punctate grooves; cords fading mid-whorl leaving smooth, glossy surface. Aperture 60 – 65 % of total height, moderately wide (MD / MAW = 2.6 – 2.8); outer lip thin. Basal lip strongly rounded and somewhat flared abapically. Columella thickened, slightly erect abapically, with a single strong fold. Columellar callus above fold and parietal callus forming delicate, thin sheet.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFA96766F2BD0275FC28FF03.taxon	discussion	Discussion. Acteon orthesi (Benoist, 1889), from the Early Miocene of France, also has a conical spire with indistinct suture, but differs in its much larger size (SL: 13 mm) and higher last whorl (Peyrot 1932: pl. 11, figs 63, 67). A specimen described by Kókay (1966) from Herend-Márkó (Hungary) might represent a stout specimen of Acteon gutta sp. nov. Kókay (1966) suggested these specimens might be conspecific with A. laevigatus (Grateloup, 1828) from the Lower Miocene Aquitanian and Burdigalian of the Aquitaine Basin, France. We have studied specimens of Acteon laevigatus from the Burdigalian of St. - Paul-lès-Dax (France) (B. L. collection) and confirm the separation of both species based on the higher spire and more incised suture of Acteon laevigatus. In addition, it has a broad, low protoconch, which is not tilted. Paleoenvironment. Probably shallow marine, based on the occurrence at Várpalota (own data, M. H.).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFA96766F2BD0275FC28FF03.taxon	distribution	Distribution in Central Paratethys. Pannonian Basin: Várpalota (Hungary) (Strausz 1966),? Herend-Márkó (Hungary) (Kókay 1966); Făget Basin: Coşteiu de Sus (Romania) (hoc opus).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFAB6761F2BD0636FDA5FDEB.taxon	description	Figs 4 A – C	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFAB6761F2BD0636FDA5FDEB.taxon	materials_examined	Type material. Holotype, NHMW 2016 / 0177 / 1298, SL: 15.6 mm, MD: 7.8 mm, Lăpugiu de Sus (Romania), Figs 4 A 1 – A 2. Paratypes: NHMW 2016 / 0177 / 1298 a, SL: 14.0 mm, MD: 6.7 mm, Lăpugiu de Sus (Romania), Figs 4 B 1 – B 2. NHMW 2016 / 0177 / 1298 b, SL: 14.1 mm, MD: 6.8 mm, Lăpugiu de Sus (Romania), Fig. 4 C. Type locality. Lăpugiu de Sus (Romania), Făget Basin. Type stratum. Silt and clay of the Dej Formation. Age. Middle Miocene, middle Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFAB6761F2BD0636FDA5FDEB.taxon	etymology	Etymology. Referring to Romania.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFAB6761F2BD0636FDA5FDEB.taxon	diagnosis	Diagnosis. Large, solid, moderately slender shell with moderately high spire and ovate last whorl; sculpture of numerous, subequal, flattened, often bifid spiral cords separated by narrow, distinctly punctate grooves with numerous, close-set pits, often touching each other.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFAB6761F2BD0636FDA5FDEB.taxon	description	Description. Shell large, solid, moderately slender (SL / MD = 2.0 – 2.1), with moderately high spire and broad apex (MD / ApD = 1.3). Protoconch low, depressed of one smooth whorl. Teleoconch of up to six whorls separated by narrow, deeply impressed, linear suture. First teleoconch whorls subcylindrical. Later spire whorls moderately convex with periphery at abapical suture, apical angle 58 – 66 °. Sculpture of up to eight moderately broad, flattened spiral cords separated by punctate grooves; spiral cords occasionally bifid. Last whorl 74 – 79 % of total height, moderately convex, ovate, with periphery placed just above mid-whorl (SL / PH = 1.9 – 2.0). Base moderately constricted. Sculpture of last whorl of about 35 moderately broad, subequal, flattened, spiral cords, separated by narrow, distinctly punctate grooves with numerous, close-set pits. Pits almost touching, separated by delicate or interrupted axials. Cords often bifid, raising number of spiral cords. Spiral sculpture not weakening mid-whorl. Aperture 62 – 68 % of total height, moderately narrow (MD / MAW = 2.9 – 3.0); outer lip simple, weakly flattened mid-whorl, strongly rounded and slightly flared abapically. Columella thickened, slightly erect abapically, with a single strong fold. Columellar callus below columellar fold forming broad, detached rim; columellar callus above columellar fold and parietal callus forming thin, delicate sheet.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFAB6761F2BD0636FDA5FDEB.taxon	discussion	Discussion. Acteon romanicus sp. nov. is a very distinctive species due to its conical spire and ovate last whorl with crowded spiral cords and finely punctate spiral grooves. ‘ Acteon’ benoisti (Peyrot, 1932), from the Burdigalian of France, differs in its slenderer shape and higher spire (see Peyrot 1932: pl. 14, figs 5 – 7). Acteon antipinguis sp. nov. differs in its higher spire and lower number of spiral cords (~ 35 versus ~ 25). Rictaxis achatinus (Sacco, 1897), as defined by the lectotype illustrated in Ferrero Mortara et al. (1984: pl. 52, fig. 1), is slenderer, has a weaker columellar fold and a higher spire. Sacco (1897: pl. 3, figs 43 – 46) illustrated several large specimens from the Pliocene of Asti as ‘ Actaeonidea achatina ’, which are probably not conspecific with Rictaxis achatinus. These shells differ from Acteon romanicus sp. nov. in their subcylindrical last whorl and lower spire. ‘ Acteon ’ brevispira (Sacco, 1897), from the Tortonian of Italy, has a much shorter spire (see Sacco 1897: p. 3, figs 48 – 49) and ‘ Acteon ’ pseudopinguis (Sacco, 1897), from the Middle Miocene of the Colli Torinesi (Italy), differs in its inflated last whorl (see Sacco 1897: p. 3, figs 41 – 42). Paleoenvironment. Unknown.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFAB6761F2BD0636FDA5FDEB.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Făget Basin: Coşteiu de Sus, Lăpugiu de Sus (Romania) (Boettger 1907; hoc opus).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFAC6760F2BD059EFD52F823.taxon	description	Figs 5 A – C	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFAC6760F2BD059EFD52F823.taxon	materials_examined	Type material. We are not aware of the whereabouts of the type material. Illustrated material. NHMW 2018 / 0248 / 0484, SL: 6.0 mm, MD: 2.9 mm, Coşteiu de Sus (Romania), Figs 5 A 1 – A 2. NHMW 2018 / 0248 / 0484 a, SL: 6.0 mm, MD: 2.9 mm, Coşteiu de Sus (Romania), Figs 5 B 1 – B 2. NHMW 1859 / 0038 / 0121, SL: 12.8 mm, MD: 5.5 mm, Grund (Austria), Figs 5 C 1 – C 2.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFAC6760F2BD059EFD52F823.taxon	description	Revised description. Shell large, moderately slender (SL / MD = 2.3), with moderately high spire and broad apex (MD / ApD = 1.3). Protoconch depressed of one smooth whorl. Teleoconch of up to five whorls separated by narrowly canaliculate, linear suture. Spire whorls suture, nearly straight sided, with periphery at abapical suture, apical angle 59 °. Last whorl 78 % of total height, ovate with weak, shoulder, weakly concave above, weakly convex below, with periphery just above mid-whorl (SL / PH = 1.9). Base moderately constricted. Sculpture of last whorl of prominent, bifid spiral cords on base, separated by narrow, weakly punctate grooves. Spiral cords rapidly weakening adapically; last whorl smooth above. Aperture 62 % of total height, moderately wide (MD / MAW = 2.6); outer lip not preserved; basal lip strongly rounded, slightly flaring. Columella weakly thickened, slightly erect abapically, with a single strong fold. Columellar callus below columellar fold forming broad, detached rim; columellar callus above columellar fold forming thin, sheet. Parietal callus not developed.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFAC6760F2BD059EFD52F823.taxon	discussion	Discussion. For descriptions of Pliocene specimens and for a discussion on the variability of this species see Landau et al. (2023: 365). This species is extraordinarily variable in shape and sculpture and might represent an unresolved species complex. We include here also two specimens with up to three punctate spiral grooves below the adapical suture, delimiting broad, flattened spiral cords (Figs 3 A – B). These shells seem to represent juvenile specimens. Paleoenvironment. Shallow marine, inner neritic (Harzhauser 2002; Landau et al. 2023: 365)	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFAC6760F2BD059EFD52F823.taxon	distribution	Distribution in Central Paratethys. Eggenburgian (Early Miocene): Petroşeni Basin: Sălătruc (Romania) (Răileanu & Negulescu 1964). Karpatian (Early Miocene): Pannonian Basin: Várpalota (Hungary) (Steininger 1973);? Korneuburg Basin: Stetten (Austria) (Harzhauser 2002). Badenian (Middle Miocene): Korytnica Basin: Korytnica (Poland) (Friedberg 1928); Polish-Ukrainian Fore-Carpathian Basin: Zboriv (Ukraine) (Friedberg 1928); North Alpine-Carpathian Foreland Basin: Grund (Austria) (hoc opus); Pannonian Basin: Szob, Várpalota (Hungary) (Strausz 1966); Southern Pannonian Basin: Miljevići (Bosnia and Herzegovina) (Atanacković 1985); Dacian Basin: Dobrusha, Opanets (Bulgaria) (Kojumdgieva in Kojumdgieva & Strachimirov 1960); Făget Basin: Coşteiu de Sus (Romania) (hoc opus). Proto-Mediterranean Sea. Burdigalian (Early Miocene): Colli Torinesi (Italy) (Sacco 1897). Serravallian (Middle Miocene): Karaman Basin: Akpınar, Lale, Seyithasan (Turkey) (Landau et al. 2013); Tortonian (Late Miocene): Po Basin (Italy) (Sacco 1897). Pliocene: Italy, Spain, France, Tunisia (see Landau et al. 2023 for details). Northeastern Atlantic. Aquitanian and Burdigalian (Early Miocene): Aquitaine Basin (France) (Peyrot 1932); Langhian and Serravallian (Middle Miocene): Aquitaine Basin (France) (Peyrot 1932); Langhian (Middle Miocene): Loire Basin (France) (Glibert 1952 a); Tortonian and Messinian (Late Miocene): France (Landau et al. 2020), Algarve Basin (Portugal) (Dollfus et al. 1903); Pliocene: France, Spain (see Landau et al. 2023 for details). North Sea. Burdigalian / Langhian (Middle Miocene): North Sea Basin: Belgium (Glibert 1952 b), Denmark (Sorgenfrei 1958), Germany (Anderson 1964; Wienrich 2007), The Netherlands (A. W. Janssen 1984); Tortonian (Late Miocene): Denmark (Schnetler 2005).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFAE6763F2BD0786FCEEF845.taxon	description	Figs 3 D – F	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFAE6763F2BD0786FCEEF845.taxon	materials_examined	Type material. Syntypes illustrated in Grateloup (1838: pl. 6, figs 24 – 25), St-Paul-lès-Dax (France). We are not aware of the whereabouts of the type material. Illustrated material. Lectotype of Acteon subpunctulatus Boettger, 1902 (designated by Zilch 1934: 276), SMF 363155 - 12.2544 a (glass plate 392), SL: 5.3 mm, MD: 2.8 mm, Coșteiu de Sus (Romania), illustrated in Zilch (1934: pl. 22, fig. 11). The specimen was destroyed during WWII, Fig 3 D. NHMW 2020 / 0004 / 1666 b, SL: 4.3 mm, SL: 2.0 mm, Bad Vöslau (Austria), Figs 3 E 1 – E 2. RGM 776 928 / 1, SL: 2.3 mm, MD: 1.2 mm, Seyithasan (Turkey), Serravallian (Middle Miocene), illustrated in Landau et al. (2013: pl. 52, fig. 14), Fig 3 F. Revised description. Shell small, moderately slender (SL / MD = 2.1), with moderately high spire and broad apex (MD / ApD = 1.2 – 1.4). Protoconch low conical of 1.25 convex, smooth whorls. Teleoconch of up to four whorls separated by narrow, deeply impressed, linear suture. Spire whorls convex, smooth with periphery just above abapical suture, apical angle 48 – 52 °. Last whorl 75 – 78 % of total height, weakly convex, with periphery just above mid-whorl. Sculpture of weak, moderately broad, flattened, spiral cords, separated by faintly punctate grooves on base. Cords fading towards mid-whorl. Aperture 62 – 63 % of total height, moderately wide (MD / MAW = 2.5 – 2.8); outer lip not preserved. Basal lip strongly rounded and slightly flared abapically. Columella thickened, slightly erect abapically, with a single strong fold. Columellar callus below columellar fold forming narrow, detached rim; columellar callus above fold and parietal callus forming thin sheet. Color pattern of three indistinct bands of blotches below adapical suture, at insertion of aperture and at base.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFAE6763F2BD0786FCEEF845.taxon	discussion	Discussion. This species is characterized by its convex whorls and its smooth, glossy surface. We have compared the Paratethyan specimens against specimens of Acteon laevigatus from the lower Burdigalian of St-Paul-lès-Dax (France) and found them to be identical. Therefore, we treat Actaeon subpunctulatus Boettger, 1902 as subjective junior synonym of Tornatella laevigata Grateloup, 1828. Specimens from the Miocene of the North Sea Basin described as this species by Janssen (1984) and Wienrich (2007) are excluded from Acteon laevigatus herein. Their spiral sculpture extends along most if not all of the last whorl quite unlike from French Atlantic specimens. Moths et al. (2010) illustrated these specimens as Acteon sp. Paleoenvironment. Unknown.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFAE6763F2BD0786FCEEF845.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienna Basin: Bad Vöslau (Austria) (hoc opus); Făget Basin: Coşteiu de Sus (Romania) (Boettger 1902); Dacian Basin: Islaz (Romania) (Rado & Muţui 1970). Proto-Mediterranean Sea. Serravallian (Middle Miocene): Karaman Basin: Seyithasan (Turkey) (Landau et al. 2013). Northeastern Atlantic. Aquitanian (Early Miocene): Aquitaine Basin: Léognan, Saucats) (France) (Peyrot 1932); Burdigalian (Early Miocene): Aquitaine Basin: Cabannes, Cestas, Mimbaste, Saint-Médard-en-Jalles, St-Paul-lès-Dax (France) (Peyrot 1932); Langhian (Middle Miocene): Loire Basin: Manthelan, Ferrière-Larçon, Paulmy, Pontlevoy (France) (Glibert 1952 a; own data B. L.).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFAF6762F2BD0786FBD2F830.taxon	description	Figs 6 A – F	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFAF6762F2BD0786FBD2F830.taxon	materials_examined	Type material. Holotype, NHMW 1851 / 0002 / 0071, SL: 15.2 MD: 8.9 mm, Grund (Austria), illustrated in Hörnes (1856: pl. 46, figs 24 a – b), Figs 6 D 1 – D 3. Paratypes: NHMW 2010 / 0004 / 1662, SL: 8.4 mm, MD: 4.4 mm, Bad Vöslau (Austria), Figs 6 A 1 – A 2. NHMW 2010 / 0004 / 1662 a, SL: 10.0 mm, MD: 5.5 mm, Bad Vöslau (Austria), Figs 6 B 1 – B 2. NHMW 2010 / 0004 / 1662 b, SL: 6.9 mm, MD: 3.8 mm, Bad Vöslau (Austria), Figs 6 C 1 – C 2. NHMW 1864 / 0001 / 0619, SL: 15.2 mm, MD: 8.3 mm, Grund (Austria), Figs 6 E 1 – E 2. NHMW 1851 / 0002 / 0071, SL: 7.8 mm, MD: 4.5 mm, Grund (Austria), Figs 6 F 1 – F 2. Additional paratypes. 13 spec., NHMW 1851 / 0002 / 0071 a, Grund (Austria). Type locality. Grund (Austria). Type stratum. Grund Formation. Age. Early Badenian, Langhian, Middle Miocene.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFAF6762F2BD0786FBD2F830.taxon	etymology	Etymology. Latin trifasciatus (= three bands), referring to the color pattern.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFAF6762F2BD0786FBD2F830.taxon	diagnosis	Diagnosis. Large, broadly ovate shell with moderately high spire; smooth except for spiral cords on base; with characteristic color pattern of three dark spiral bands below adapical suture, along mid-whorl and on base.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFAF6762F2BD0786FBD2F830.taxon	description	Description. Shell large, relatively solid, broadly ovate (SL / MD = 1.7 – 1.8), with moderately low spire and broad apex (MD / ApD = 1.4). Protoconch small, slightly tilted, depressed, 1 smooth whorl. Teleoconch of up to six convex whorls separated by narrow, weakly canaliculate suture. Spire whorls weakly convex, smooth with periphery at abapical suture, apical angle 78 – 89 °. Last whorl 81 – 84 % of total height, moderately convex with faint shoulder, with periphery above mid-whorl. Sculpture of moderately broad, flattened, spiral cords, separated by weakly punctate grooves on base. Cords narrowing and fading towards mid-whorl. Aperture 64 – 69 % of total height, moderately wide (MD / MAW = 2.0 – 2.4); outer lip simple, weakly convex, strongly rounded and slightly flared abapically. Columella thickened, slightly erect abapically, with a single strong fold. Columellar callus below columellar fold forming narrow, detached rim; columellar callus not developed above fold; no parietal callus. Color pattern of three intense, narrow dark spiral bands just below adapical suture, at mid-whorl (placed at level of insertion of outer lip) and on base.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFAF6762F2BD0786FBD2F830.taxon	discussion	Discussion. This very distinctive species was confused by Hörnes (1856) with the Pliocene to Recent Acteon tornatilis (Linnaeus, 1758), which differs in its slightly slenderer last whorl, its more prominent spiral sculpture and its color pattern, which consists of two broad, whitish bands below shoulder and below mid-whorl (see Cossignani & Ardovini 2011: 36, 368, unnumbered fig.; Gofas et al. 2011: 352, unnumbered fig.; Landau et al. 2023: pl. 21). Acteon antipinguis sp. nov. is comparable in size but differs in its slenderer, subcylindrical last whorl and much more prominent spiral sculpture over the entire shell. A fragmentary shell from the Eggenburgian (Early Miocene) of Romania, illustrated by Răileanu & Negulescu (1964: pl. 16, fig. 3) as ‘ Actaeon tornatilis ’ displays identical color pattern but is much smaller (SL: 3 mm) and is excluded here. A minute specimen from the Badenian of Várpalota (Hungary) illustrated by Katona et al. (2011: pl. 2, fig. 4) as ‘ Acteon tornatilis ’ is surely another species. If the given size (SL: 0.7 mm) is correct, this species does not correspond to any species described herein. Otherwise, it is reminiscent of Acteon semistriatus (Férussac, 1822). Acteon sp. 2 of Stein et al. (2016: text-fig. 85 j) has identical color pattern and might represent an occurrence of Acteon trifasciatus sp. nov. from the Early Miocene of the North Sea. Paleoenvironment. At Grund (Austria) fossiliferous channel fills, which formed in middle to outer neritic environments, bearing allochthonous assemblages uniting coastal-mudflat faunas with inner neritic ones occur (Zuschin et al. 2005; Roetzel 2009). The preservation of the fossil shells suggests that the shells belong to the allochthonous shallow water fauna.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFAF6762F2BD0786FBD2F830.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): North Alpine Foreland Basin: Grund (Austria) (Hörnes 1856); Vienna Basin: Bad Vöslau, Steinebrunn (Austria) (Hörnes 1856; hoc opus).? North Sea. Hemmoorian (Early Miocene): Germany (Stein et al. 2016).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFB0677CF2BD016AFE28F845.taxon	description	Figs 7 A 1 – A 2	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFB0677CF2BD016AFE28F845.taxon	materials_examined	Type material. Holotype, HNHM INV 2025.97, SL: 17.5 mm, MD: 11.6 mm, Parádfürdő (Hungary), illustrated in Kovács et al. (2023: 42, figs 102 – 103), Figs 7 A 1 – A 2. Type locality. Parádfürdő (Hungary). Type stratum. Sandstone of the Ilonavölgy Mb. of the Pétervására Formation. Age. Ottnangian (Early Miocene, mid-Burdigalian).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFB0677CF2BD016AFE28F845.taxon	etymology	Etymology. In honor of Zoltán Vicián (Budapest, Hungary), in recognition of his contributions to Paratethyan paleontology.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFB0677CF2BD016AFE28F845.taxon	diagnosis	Diagnosis. Large, broadly ovate, globose shell with moderately low spire and sculpture of numerous flattened, bifid spiral cords separated by narrow grooves.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFB0677CF2BD016AFE28F845.taxon	description	Description. Shell large, broadly ovate, globose (SL / MD = 1.5), with moderately low spire and broad apex (MD / ApD = 1.5). Protoconch unknown. Teleoconch of up to five convex whorls separated by narrow, weakly canaliculate suture. Spire whorls weakly convex, sculpture not preserved, apical angle 90 °. Last whorl 78 % of total height, strongly convex with faint shoulder, with periphery above mid-whorl. Sculpture of moderately broad, flattened, bifid spiral cords, separated by narrow, weakly punctate grooves on entire shell. Aperture 70 % of total height, moderately wide (MD / MAW = 2.3); outer lip simple, convex, strongly rounded and slightly flared abapically. Columella thickened, slightly erect abapically, with a single fold. Columellar callus below columellar fold relatively broad, detached rim.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFB0677CF2BD016AFE28F845.taxon	discussion	Discussion. This species is distinguished from all other Paratethyan Acteonidae by its large size, globose outline and peculiar sculpture of close-set bifid spiral cords. Acteon inflatus (Borson, 1821), from the Early Miocene of the Colli Torinesi, has a higher and less convex last whorl (see Sacco 1897; pl. 3, figs 8 – 10) and ‘ Acteon ’ biplicata (Borson, 1821), from the Italian Pliocene), has simple spiral cords and a more acute spire (see Sacco 1897: pl. 3, fig. 54). Paleoecology. The assemblage from Parádfürdő (Hungary) as described by Kovács et al. (2023) points to shallow marine environments.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFB0677CF2BD016AFE28F845.taxon	distribution	Central Paratethys Sea. Ottnangian (Early Miocene): Early Miocene North Hungarian Basin: Parádfürdő (Hungary) (Kovács et al. 2023).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFB2677EF2BD0786FE0FFCC3.taxon	description	Figs 8 A – D	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFB2677EF2BD0786FE0FFCC3.taxon	materials_examined	Type material. Holotype, NHMW 1855 / 0045 / 0523, SL: 2.6 mm, MD: 1.4 mm, Baden (Austria), Figs 8 B 1 – B 2. Paratypes: NHMW 1866 / 0040 / 0219, SL: 8.0 mm, MD: 4.2 mm, Baden Sooss (Austria), Figs 8 A 1 – A 2. NHMW 1855 / 0045 / 0523 a, SL: 3.0 mm, MD: 1.6 mm, Baden (Austria), Figs 8 C 1 – C 2. NHMW 1861 / 0001 / 0469, SL: 2.8 mm, MD: 1.8 mm, Bad Vöslau (Austria), Figs 8 D 1 – D 2. Type locality. Baden (Austria). Type stratum. Baden Formation. Age. Middle Miocene, middle Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFB2677EF2BD0786FE0FFCC3.taxon	etymology	Etymology. Referring to Vienna (Austria).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFB2677EF2BD0786FE0FFCC3.taxon	diagnosis	Diagnosis. Small to medium sized, relatively solid, moderately broad, ovate shell with very prominent, flattened, often bifid spiral cords and broad grooves with large subrectangular pits and large columellar fold.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFB2677EF2BD0786FE0FFCC3.taxon	description	Description. Shell small to medium sized, relatively solid, moderately broad, ovate (SL / MD = 1.8 – 1.9), with moderately high spire and broad apex (MD / ApD = 1.3 – 1.4). Protoconch unknown. Teleoconch of up to five whorls separated by narrowly impressed, linear suture. Spire whorls moderately convex, weakly shouldered, with periphery above abapical suture, apical angle 61 – 65 °. Sculpture of up to five broad, weakly convex spiral cords separated by deep grooves with large subrectangular pits, separated by narrow, opisthocline axial growth lines; adapical cord often bifid. Last whorl 77 – 82 % of total height, moderately convex with weak, rounded shoulder, with periphery just above mid-whorl (SL / PH = 1.8 – 1.9). Base moderately constricted. Sculpture of last whorl of about 20 broad, flattened, spiral cords, separated by broad grooves with large subrectangular pits. Spiral cords often bifid, raising number of spiral cords. Spiral sculpture not weakening mid-whorl. Aperture 63 – 69 % of total height, moderately wide (MD / MAW = 2.6 – 2.7); outer lip simple, weakly convex mid-whorl, strongly rounded and slightly flared abapically. Columella thickened, slightly erect abapically, with a single, very strong fold. Columellar callus below columellar fold forming broad, detached rim; columellar callus above columellar fold and parietal callus forming thin, sharply delimited sheet.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFB2677EF2BD0786FE0FFCC3.taxon	discussion	Discussion. This species is characterized by its relatively stout shape and prominent, coarse sculpture. The extant Acteon monterosatoi Dautzenberg, 1889, from the eastern Atlantic and the Mediterranean Sea, differs in its slightly broader shell and lower spire. The species is superficially reminiscent of Callostracon Repetto & Bianco, 2012 [type species Colostracon tyrrhenicum Smriglio & Mariottini, 1996; present-day, Mediterranean Sea] but Callostracon does not have a columellar fold. Paleoenvironment. The occurrence in the Baden Formation at Bad Vöslau (Austria) suggests middle to outer neritic environments with up to 210 m water depth (Kranner et al. 2021).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFB2677EF2BD0786FE0FFCC3.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienna Basin: Baden, Baden Sooss, Bad Vöslau (Austria) (hoc opus).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFB3677EF2BD0477FF3CFC3B.taxon	type_taxon	Type species. Actaeon aciculatus Cossmann, 1889; original designation by Cossmann (1889: 302). Eocene, France.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFB36779F2BD032FFBD4FA99.taxon	description	Figs 9 A – C	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFB36779F2BD032FFBD4FA99.taxon	materials_examined	Type material. Lectotype (designated by Zilch 1934: 276), SMF 363156 - 12.2545 a (glass plate 393), SL: 5.8 mm, MD: 2.3 mm, Coșteiu de Sus (Romania), illustrated in Zilch (1934: pl. 22, fig. 12). The specimen was destroyed during WWII, Fig. 9 A. Illustrated material. NHMW 2020 / 0004 / 1666, SL: 4.8 mm, SL: 1.9 mm, Bad Vöslau (Austria), Figs 9 B 1 – B 2. NHMW 2020 / 0004 / 1666 a, SL: 4.7 mm, SL: 2.1 mm, Bad Vöslau (Austria), Figs 9 C 1 – C 2.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFB36779F2BD032FFBD4FA99.taxon	description	Revised description. Shell small, slender (SL / MD = 2.4 – 2.5), with high spire and broad apex (MD / ApD = 1.3). Protoconch strongly tilted, of about one smooth, convex whorl. Teleoconch of up to four whorls separated by narrow, deeply impressed, linear suture. Spire whorls weakly convex, glossy with periphery just above abapical suture, apical angle 38 – 42 °. Sculpture of hardly visible spiral cords. Last whorl 71 – 74 % of total height, weakly convex, elongate ovate, glossy, with periphery placed mid-whorl (SL / PH = 1.9 – 2.0). Sculpture of faint spiral grooves strengthening and weakly punctate on base. Aperture 54 – 57 % of total height, moderately narrow (MD / MAW = 2.4 – 2.7); outer lip thin, weakly convex. Basal lip strongly rounded. Columella weakly thickened, with a single moderately strong fold. Columellar callus below columellar fold forming narrow, adherent rim; columellar callus above fold forming thin sheet. No parietal callus.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFB36779F2BD032FFBD4FA99.taxon	discussion	Discussion. This species is unique within the Paratethyan Acteonidae due to its slender, glossy shell. We place it in Crenilabium Cossmann, 1889 based on its general shape. Especially the extant Crenilabium birmani Simone, 2006 from Brazil is exceptional similar to the Paratethyan species in shape, sculpture and protoconch features but differs in its higher spire (see Simone 2006: figs 5 – 9). The extant Mediterranean Crenilabium exile (Jeffreys, 1870) is also highly reminiscent of Crenilabium reussi (Boettger, 1902). The main difference is that Crenilabium exile attains twice the maximum size at the same number of whorls (see Landau et al. 2023: pl. 23). Crenilabium pedemontanum (Sacco, 1897), from the Middle Miocene of the Colli Torinesi (Italy) is even more slender, with a higher spire (see Sacco 1897: pl. 3, fig. 50; Ferrero Mortara et al. 1984: pl. 52, fig. 4). Similarly, Crenilabium dargelasi (de Basterot, 1825), from the Burdigalian of France, is slenderer and has a higher spire (see Peyrot 1932: pl. 11, figs 40 – 41). Paleoenvironment. The occurrence at Várpalota (Hungary) suggests agitated, shallow marine environments (own data, M. H.).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFB36779F2BD032FFBD4FA99.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Pannonian Basin: Várpalota (Hungary) (Katona et al. 2011); Făget Basin: Coşteiu de Sus (Romania) (Boettger 1902).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFB46779F2BD02B1FEB5F9F1.taxon	type_taxon	Type species. Tornatella punctocoelata Carpenter, 1863; original designation by Dall (1871: 136). Present-day, California (USA).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFB4677BF2BD0169FE56FD53.taxon	description	Figs 10 A – C	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFB4677BF2BD0169FE56FD53.taxon	materials_examined	Type material. Holotype, NHMW 2007 z 0086 / 0000 e, SL: 10.4 mm, MD: 4.4 mm, Grund (Austria), Figs 10 A 1 – A 2. Paratypes: NHMW 2007 z 0086 / 0000 f, SL: 11.4 mm, MD: 4.8 mm, Grund (Austria), Figs 10 B 1 – B 2. NHMW 2007 z 0086 / 0000 g, SL: 10.6 mm, MD: 5.0 mm, Grund (Austria), Figs 10 C 1 – C 2. Type locality. Grund (Austria). Type stratum. Grund Formation. Age. Middle Miocene, early Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFB4677BF2BD0169FE56FD53.taxon	etymology	Etymology. Referring to Austria.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFB4677BF2BD0169FE56FD53.taxon	diagnosis	Diagnosis. Large, moderately slender shell with moderately high spire, elongate ovate last whorl; sculpture of moderately broad, flattened, spiral cords, separated by narrow, weakly punctate grooves.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFB4677BF2BD0169FE56FD53.taxon	description	Description. Shell large, relatively solid, moderately slender to rarely moderately broad (SL / MD = 1.7 – 2.3), with moderately high spire and broad apex (MD / ApD = 1.4 – 1.5). Protoconch conical of 1.5 smooth whorls with tilted initial part. Teleoconch of up to four whorls separated by narrow, deeply impressed, linear suture. Spire whorls weakly convex, with periphery above abapical suture, apical angle 65 – 71 °. Sculpture of up to eight broad, flattened spiral cords separated by narrow, relatively shallow, indistinctly punctate grooves; spiral cords occasionally bifid. Last whorl 65 – 68 % of total height, weakly convex, elongate ovate with weak, rounded shoulder, with periphery at or above mid-whorl (SL / PH = 1.7 – 1.9). Base moderately constricted. Sculpture of last whorl of up to 30 moderately broad, flattened, spiral cords, separated by narrow, weakly punctate grooves. Cords rarely bifid. Spiral sculpture not weakening at mid-whorl. Aperture 66 – 68 % of total height, moderately narrow (MD / MAW = 2.2 – 2.8); outer lip simple, weakly convex at mid-whorl, strongly rounded, somewhat flared and weakly crenulated abapically. Columella weakly thickened, slightly erect abapically, with a single strong fold. Columellar callus below columellar fold forming narrow, weakly detached rim; columellar callus above columellar fold and parietal callus forming thin, delicate (rarely preserved) sheet.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFB4677BF2BD0169FE56FD53.taxon	discussion	Discussion. Rictaxis austriacus sp. nov. is reminiscent of the northeastern Atlantic Miocene to Pliocene Rictaxis tornatus (Millet, 1854) in size and general sculpture but differs in its weakly punctate spiral grooves and the relatively mammillate protoconch. It is also slightly slenderer than most specimens of Rictaxis tornatus (see Ceulemans et al. 2018: pl. 7, fig. 2; Landau et al. 2023: pl. 24). Rictaxis siedli sp. nov. is much smaller and differs in its prominent, subcircular to subquadratic pits. Rictaxis achatinus (Sacco, 1897), from the Mediterranean Pliocene, differs in its weaker columellar fold and the more prominent spiral grooves with deeper pits (see Ferrero Mortara et al. 1984: pl. 52, fig. 1). Acteon romanicus sp. nov. is larger and has a broader last whorl with wider base, distinctly punctate spiral grooves, and the abapical part of the outer lip is not as expanded as it is in Rictaxis species. Paleoenvironment. At Grund (Austria) fossiliferous channel fills, which formed in middle to outer neritic environments, bearing allochthonous assemblages uniting coastal-mudflat faunas with inner neritic ones occur (Zuschin et al. 2005; Roetzel 2009). The preservation of the fossil shells suggests that the shells belonged to the allochthonous shallow water fauna.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFB4677BF2BD0169FE56FD53.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): North Alpine Foreland Basin: Grund (Austria) (hoc opus).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFB6677AF2BD05C6FF0BFD53.taxon	description	Figs 10 D – E	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFB6677AF2BD05C6FF0BFD53.taxon	materials_examined	Type material. Holotype, NHMW 1997 z 0178 / 1214, SL: 5.9 mm, MD: 2.7 mm, Bad Vöslau (Austria), Figs 10 D 1 – D 2. Paratype, NHMW 2010 / 0004 / 2663 a, SL: 6.6 mm, MD: 3.1 mm, Bad Vöslau (Austria), Figs 10 E 1 – E 2. Type locality. Bad Vöslau (Austria). Type stratum. Baden Formation. Age. Middle Miocene, middle Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFB6677AF2BD05C6FF0BFD53.taxon	etymology	Etymology. In honor of Wolfgang Siedl (OMV, Vienna), in recognition of his contributions to the stratigraphy of the Vienna Basin.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFB6677AF2BD05C6FF0BFD53.taxon	diagnosis	Diagnosis. Medium sized, moderately slender, with moderately high spire with ovate last whorl and about 30 spiral cords separated by punctate grooves with vaguely cancellate pattern of pits and axials; weak columellar fold.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFB6677AF2BD05C6FF0BFD53.taxon	description	Description. Shell medium sized, moderately slender (SL / MD = 2.1), with moderately high spire and broad apex (MD / ApD = 1.4 – 1.5). Protoconch conical of one smooth whorl with tilted initial part. Teleoconch of up to three whorls separated by narrow, deeply impressed, linear suture. Spire whorls weakly convex, with periphery above abapical suture, apical angle 50 – 53 °. Sculpture of nine moderately broad, weakly convex spiral cords separated by punctate grooves. Last whorl 79 – 83 % of total height, weakly convex, ovate with weak, rounded shoulder, with periphery at or above mid-whorl (SL / PH = 1.7 – 2.0). Base moderately constricted. Sculpture of last whorl of about 30 relatively narrow, weakly convex spiral cords, separated by broad, punctate grooves. Pits close-set, subcircular to subquadratic, separated by narrow axials, resulting in vaguely cancellate pattern. Cords not bifid. Spiral sculpture not weakening mid-whorl. Aperture 65 – 70 % of total height, moderately wide (MD / MAW = 2.1 – 2.4); outer lip simple, weakly convex mid-whorl, crenulated, strongly rounded, somewhat flared and crenulated abapically. Columella weakly thickened, only slightly erect abapically, with a single, relatively weak fold. Columellar callus below columellar fold forming narrow rim; columellar callus above columellar fold forming thin, narrow, delicate sheet. Parietal callus not developed. Color pattern of three dark bands; narrow band at adapical suture, broader band at intersection of upper lip, slightly narrower band on base.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFB6677AF2BD05C6FF0BFD53.taxon	discussion	Discussion. Rictaxis siedli sp. nov. is highly reminiscent of Rictaxis achatinus (Sacco, 1897) from the Pliocene of Italy (see Sacco 1897: 36, pl. 3, fig. 42 only; Ferrero Mortara et al. 1984: 283, pl. 52, figs 1 a – b). Rictaxis achatinus differs in its much larger size (SL: 15 mm), the low conical protoconch and the subcylindrical shape of the last whorl. [The lectotype of Rictaxis achatinus was erroneously listed as holotype by Ferrero Mortara et al. (1984), although several syntypes exist. Therefore, the specimen represents a lectotype (by erroneous definition as holotype) and the other specimens are paralectotypes]. There has been considerably confusion on the identity of Rictaxis achatinus in the literature, which started with Glibert (1952 a) who illustrated a specimen from the Langhian of the Loire Basin. This specimen differs from the Italian species by its much more prominent columellar fold and lower spire. Glibert’s specimen might represent the northeastern Atlantic Rictaxis tornatus (Millet, 1854). Due to this confusion, Ceulemans et al. (2018: 117) suggested to synonymize both species. The Atlantic Rictaxis tornatus (Millet, 1854) is also large like Rictaxis achatinus (SL up to 18 mm) but has a weaker columella and a lower spire. Note that none of the other Mediterranean specimens described as Acteonidea [sic] achatina in the literature represents this species; e. g., Pavia 1976 a: 114, pl. 12, fig. 8; Cavallo & Repetto 1992: 164, fig. 471; Ferrero et al. 1998: 49, pl. 2, fig. 5; Sosso & Dell’Angelo 2010: 53, fig. 68; Chirli 2013: 26, pl. 7, figs 18 – 23. As already suggested by Ceulemans et al. (2018), these shells represent another species [most of the records might represent Acteon levidensis (S. V. Wood, 1848)]. Rictaxis tornatus (Millet, 1854) differs from Rictaxis siedli sp. nov. in its larger size, stronger columella, shorter spire, and the abapical part of the outer lip is not as expanded as it is in Rictaxis species (Ceulemans et al. 2018: pl. 7, fig. 2; Landau et al. 2023: pl. 24). Paleoenvironment. The occurrence in the Baden Formation at Bad Vöslau (Austria) suggests middle to outer neritic environments with up to 210 m water depth (Kranner et al. 2021).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFB6677AF2BD05C6FF0BFD53.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienna Basin: Bad Vöslau (Austria) (hoc opus).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFB7677AF2BD05C6FE56F82C.taxon	description	Figs 6 G, 11 A – E	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFB7677AF2BD05C6FE56F82C.taxon	materials_examined	Type material. Holotype, NHMW 2007 z 0086 / 0000, SL: 7.1 mm, MD: 2.8 mm, Grund (Austria), Figs 11 A 1 – A 2. Paratypes: NHMW 2007 z 0086 / 0000 a, SL: 7.9 mm, MD: 3.1 mm, Grund (Austria), Figs 11 B 1 – B 2. NHMW 2007 z 0086 / 0000 b, SL: 7.7 mm, MD: 2.8 mm, Grund (Austria), Figs 11 C 1 – C 2. NHMW 2007 z 00860 / 000 c, SL: 7.3 mm, MD: 2.7 mm, Grund (Austria), Figs 11 D 1 – D 2. NHMW 2007 z 0086 / 0000 d, SL: 5.6 mm, MD: 2.3 mm, Grund (Austria), Figs 6 G, 11 E 1 – E 2. Type locality. Grund (Austria). Type stratum. Grund Formation. Age. Middle Miocene, early Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFB7677AF2BD05C6FE56F82C.taxon	etymology	Etymology. In honor of Ursula Göhlich, vertebrate paleontologist at the NHMW.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFB7677AF2BD05C6FE56F82C.taxon	diagnosis	Diagnosis. Medium sized, slender, elongate ovate shell with moderately high spire and sculpture of broad, flattened spiral cords separated by shallow, weakly punctate grooves; columella with single, sharp, prominent fold.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFB7677AF2BD05C6FE56F82C.taxon	description	Description. Shell medium sized, slender, elongate ovate (SL / MD = 2.5 – 2.6), with moderately high spire and broad apex (MD / ApD = 1.4). Protoconch conical with tilted nucleus, of 1.5 smooth, moderately convex whorls. Teleoconch of up to four whorls separated by narrowly impressed, linear suture. Spire whorls high, weakly convex, with periphery just above abapical suture, apical angle 44 – 48 °. Sculpture of up to eight broad, flattened spiral cords separated by narrow, weakly punctate grooves; some cords occasionally bifid. Last whorl 80 – 84 % of total height, weakly convex with weak, rounded shoulder, with periphery just above mid-whorl (SL / PH = 1.8 – 1.9). Base weakly constricted. Sculpture of last whorl of about 25 broad, flattened, spiral cords, separated by narrow, weakly punctate grooves. Widest cords in shoulder region. Spiral cords often bifid, raising number of spiral cords. Spiral sculpture not weakening mid-whorl. Aperture 65 – 67 % of total height, relatively narrow (MD / MAW = 2.3 – 3.0); outer lip simple, almost straight mid-whorl, strongly rounded and broadly flared abapically. Columella weakly thickened, with a single strong, sharp fold. Columellar callus below columellar fold forming narrow rim adherent to base; columellar callus above columellar fold and parietal callus forming thin sheet.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFB7677AF2BD05C6FE56F82C.taxon	discussion	Discussion. This species is characterized by its slender outline and sculpture of broad spiral cords separated by shallow spiral grooves. We are not aware of comparable species from the Circum-Mediterranean Neogene. The wide-spread and long-lived Acteon semistriatus (Férussac, 1822) has a less elongate last whorl, a wider base, reduced spiral sculpture mid-whorl on the last whorl, and the abapical part of the outer lip is less expanded (see Landau et al. 2023: pl. 20 and references therein). Paleoenvironment. At Grund (Austria) fossiliferous channel fills, which formed in middle to outer neritic environments, bearing allochthonous assemblages uniting coastal-mudflat faunas with inner neritic ones occur (Zuschin et al. 2005; Roetzel 2009). The preservation of the fossil shells suggests that the shells belong to the allochthonous shallow water fauna.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFB7677AF2BD05C6FE56F82C.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): North Alpine Foreland Basin: Grund (Austria) (hoc opus).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFB86774F2BD006AFD6DFDCF.taxon	type_taxon	Type species. Marginella costata Eichwald, 1830; by monotypy. Middle Miocene, Ukraine.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFB86774F2BD006AFD6DFDCF.taxon	discussion	Discussion. Ringiculocosta Sacco, 1892 is currently listed as subjective junior synonym of Ringicula Deshayes, 1838 by MolluscaBase eds (2025 b). Ringiculocosta costata (Eichwald, 1830), Ringiculocosta plicatula (Mayer, 1871), from the Burdigalian (Early Miocene) of France, and Ringiculocosta cancellarioides (Seguenza, 1881), from the Middle and Late Miocene of the Central Paratethys Sea and the Proto-Mediterranean Sea, differ distinctly from Ringicula in their peculiar, cancellate sculpture. Therefore, we prefer to keep these species separated at genus level. The genus appeared during the Early Miocene in the northeastern Atlantic and persisted to the Pliocene of the Mediterranean Sea (Ringiculocosta costata astensis Sacco, 1892). Ringiculocosta occurred also in the Eastern Paratethys Sea during the Tarkhanian and Chokrakian (Middle Miocene) (Guzhov 2022 a). During the Early Miocene it reached also the North Sea where it was represented by two species (Ringiculocosta costata (Eichwald, 1830) and Ringiculocosta costata sensu Moths et al. 2010).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFB96776F2BD057BFAA8FE23.taxon	description	Figs 12 C, E, 13 A	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFB96776F2BD057BFAA8FE23.taxon	materials_examined	Type specimen. Holotype, SL: 3 mm, MD: 1.7 mm, Sassuolo (Italy), Tortonian (Late Miocene), illustrated in Seguenza (1881: pl. 2, figs 11, 11 a). We are not aware of the whereabouts of the type material. Illustrated material. NHMW-ZOO-MO- 75000 - E- 3283, SL: 3.7 mm, MD: 1.8 mm, Bad Vöslau (Austria), illustrated in Berger (1954, pl. 12, fig. 69), Figs 12 C 1 – C 2, 13 A. NHMW-ZOO-MO- 75000 - E- 57959, SL: 2.6 mm, MD: 1.5 mm, Grund (Austria), Figs 12 E 1 – E 2. Revised description. Shell small, moderately broad fusiform (SL / MD: 1.8 – 1.9) with moderately high conical spire (AA = 38 – 39 °). Protoconch paucispiral of about 1.5 whorls (dp = 400 μm). Teleoconch of three whorls separated by narrowly impressed, linear suture. Spire whorls high, moderately convex with rounded shoulder. Sculpture of close-set axial ribs separated by slightly wider interspaces, overrun by weaker spiral cords, resulting in cancellate pattern. Last whorl 65 – 66 % of total height, with periphery at or just below mid-whorl (SL / PH = 2.1 – 2.4), faintly concave subsutural ramp, rounded shoulder, moderately convex periphery; axial interspaces widening on last whorl, spiral cords broaden with narrower interspaces, resulting in rectangular pattern; base conical, constricted. Aperture 49 – 50 % of total height, moderately narrow, widening abapically (MD / MAW = 3.4); insertion of outer lip distinctly below shoulder, thickened by convex labial varix; anal sinus broad, shallow, U-shaped; siphonal canal very short, open, broad, notched at tip. Columella with two strongly elevated, robust folds abapically, parietal denticle prominent, trigonal. Columellar and parietal callus thickened, sharply delimited, narrowly expanded over base.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFB96776F2BD057BFAA8FE23.taxon	discussion	Discussion. This species differs from Ringiculocosta costata (Eichwald, 1830) in its slenderer shell, and the much lower point of insertion of the outer lip. In addition, the two species seem to have been ecologically separated with Ringiculocosta costata in shallow water environments and Ringiculocosta cancellarioides in offshore habitats. Most specimens have a prominent, rectangular sculpture (Fig. 12 C) but specimens with weak sculpture also occur (Fig. 12 E). Ringiculocosta plicatula (Mayer, 1871), from the Burdigalian of France, differs only in its more prominent axial sculpture (see Mayer 1871: 347, pl. 9, fig. 8; Morlet 1878: 267, pl. 6, fig. 9; Peyrot 1932: 148, pl. 11, figs 29 – 31). It is possible that R. cancellarioides will become a subjective junior synonym of R. plicatula after revision of French and Italian specimens. Paleoenvironment. The occurrence in the Baden Formation at Bad Vöslau (Austria) suggests middle to outer neritic environments with up to 210 m water depth (Kranner et al. 2021).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFB96776F2BD057BFAA8FE23.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Roztocze Hills: Węglinek, (Poland) (Krach 1981); North Alpine-Carpathian Foreland Basin: Grund (Austria) (hoc opus); Vienna Basin: Bad Vöslau (Austria) (hoc opus); Pannonian Basin: Várpalota (Hungary) (Strausz 1966). Proto-Mediterranean Sea. Tortonian (Late Miocene): Emilia-Romagna: Sassuolo (Italy) (Seguenza 1881).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFBB6771F2BD052EFB60FA17.taxon	description	Figs 12 A – B, D, 13 B	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFBB6771F2BD052EFB60FA17.taxon	materials_examined	Type material. Lectotype (designated by Guzhov 2022 a: 1126), SL: 2.4 mm, MD: 1.6 mm, PSM, no. 3 / 469 a, Zhukivtsi (Ukraine), illustrated in Guzhov 2022 a: pl. 6, fig. 1). Illustrated material. NHMW 1855 / 0045 / 0036, SL: 2.2 mm, MD: 1.5 mm, Steinebrunn (Austria), Figs 12 A 1 – A 2. NHMW 1863 / 0015 / 0616, SL: 2.5 mm, MD: 1.7 mm, Niederleis (Austria), Figs 12 B 1 – B 2, 13 B. NHMW 1863 / 0015 / 0616, SL: 2.0 mm, MD: 1.4 mm, Buituri (Romania), Figs 12 D 1 – D 2. Revised description. Shell small, with globose last whorl, very broad (SL / MD: 1.4 – 1.5) with moderately high conical spire (AA = 58 – 64 °). Protoconch paucispiral of about 1.5 whorls (dp = 400 μm). Junction with teleoconch marked by beginning of cancellate sculpture. Teleoconch of three whorls separated by narrowly impressed, linear suture. Spire whorls strongly convex with rounded shoulder. Sculpture of close-set axial ribs separated by interspaces of about same width, overrun by weaker spiral cords, resulting in cancellate pattern. Last whorl 72 – 75 % of total height, with periphery just above mid-whorl (SL / PH = 1.9), weakly convex to faintly concave subsutural ramp, rounded shoulder, inflated below; sculpture occasionally more prominent and wider spaced along shoulder; base broad conical, constricted. Aperture 58 – 61 % of total height, moderately narrow, widening abapically (MD / MAW = 4.0 – 4.5). Outer lip thickened by convex labial varix, bearing low central palatal swelling; anal sinus broad, shallow, U-shaped; siphonal canal very short, open, broad, deeply notched at tip. Columella with two strongly elevated, robust folds abapically, parietal denticle prominent, trigonal. Columellar and parietal callus strongly thickened, sharply delimited, narrowly expanded over base.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFBB6771F2BD052EFB60FA17.taxon	discussion	Discussion. Ringiculocosta costata is quite variable concerning its sculpture, which led Seguenza (1881) to introduce the varieties tenuis (finely cancellate) and sicula (coarsely cancellate). Paratethyan specimens display a comparable variability within the same locality and taxonomic separation of the various morphs is unjustified. The situation seems to be different in the Eastern Paratethys Sea from where Guzhov (2022 a) described three subspecies of Ringiculocosta costata from the Tarkhanian and Chokrakian (Middle Miocene). These taxa seem to represent a distinct radiation in the largely isolated Eastern Paratethys Sea. Ringiculocosta plicatula (Mayer, 1871), from the Burdigalian (Early Miocene) of Saucats (France), differs in its slenderer shell, higher last whorl, weaker shoulder and predominant axial sculpture (see Mayer 1871: 347, pl. 9, fig. 8; Morlet 1878: 267, pl. 6, fig. 9). Similarly, Ringiculocosta cancellarioides (Seguenza, 1881) differs in its slenderer outline. Paleoenvironment. Shallow marine, inner neritic based on associated fauna (own data, M. H.).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFBB6771F2BD052EFB60FA17.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Polish-Ukrainian Fore-Carpathian Basin: Borki Wielkie (Poland), Hołdy, Hryts’kiv, Faszczówka, Tarnoruda, Ripyntsi, Shushkivtsi, Staryi Pochaiv, Zboriv, Zalistsi, Zhabyak, Zhukivtsi, Varivtsi, Yaseniv (Ukraine) (Friedberg 1938; Guzhov 2022 a); Voronyaky Hills: Golubytsya (Hołubica), Pidgirtsi (Ukraine); North Alpine-Carpathian Foreland Basin: Lysice (Czech Republic) (Berger 1954); Vienna Basin: Bad Vöslau, Niederleis, Steinebrunn, Vienna / Grinzing (Austria) (Berger 1954), Mikulov (Czech Republic) (Hörnes 1852), Borský Mikulás (Slovakia) (Švagrovský 1984); Eisenstadt-Sopron Basin: Forchtenau (Austria) (Berger 1954); Oberpullendorf Basin: Ritzing (Austria) (Berger 1954); Pannonian Basin: Budapest, Hidas (Hungary) (Strausz 1966; Kókay 1996); Southern Pannonian Basin: Hrvaćani (Bosnia and Herzegovina) (Atanacković 1985); Făget Basin: Coşteiu de Sus (Romania) (Boettger 1902); Dacian Basin: Islaz (Romania) (Rado & Muţui 1970); Borod Basin: Cetea Valley (Romania) (Popa & Chira 1999). Proto-Mediterranean Sea. Tortonian (Late Miocene): Po Basin: Montegibbio (Italy) (Sacco 1892); Emilia-Romagna: Modena (Italy), Sicily: Ciminna, Rometta (Italy) (Seguenza 1881). Atlantic. Tortonian (Late Miocene): Loire Basin, NW France (BL unpublished data). North Sea. Hemmoorian (Early Miocene): North Sea Basin: Germany (Stein et al. 2016).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFBC6771F2BD0103FE57F94F.taxon	type_taxon	Type species. Marginella auriculata Ménard de la Groye, 1811; subsequent designation by Anton (1838: 48). Present-day, Europe.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFBC6773F2BD01FBFF0BFBE3.taxon	description	Figs 14 F – J	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFBC6773F2BD01FBFF0BFBE3.taxon	materials_examined	Type material. Holotype, NHMW 2025 / 0245 / 0001, SL: 1.7 mm, MD: 1.15 mm, Nemeşeşti (Romania), Figs 14 F 1 – F 2. Paratypes, NHMW 2025 / 0245 / 0002, SL: 1.35 mm, MD: 0.9 mm, Nemeşeşti (Romania), Figs 14 1 – G 2. NHMW 2025 / 0245 / 0003, SL: 1.4 mm, MD: 0.85 mm, Nemeşeşti (Romania), Figs 14 H 1 – H 2. NHMW 2025 / 0245 / 0004, SL: 1.35 mm, MD: 0.85 mm, Nemeşeşti (Romania), Figs 14 I 1 – I 2. NHMW 2025 / 0245 / 0005, SL: 1.55 mm, MD: 1.1 mm, Nemeşeşti (Romania), Figs 14 J 1 – J 2. Additional paratypes. 10 specimens, NHMW 2025 / 0245 / 0006, Nemeşeşti (Romania). Type locality. Nemeşeşti (Romania), Făget Basin. Type stratum. Silt and clay of the Dej Formation. Age. Middle Miocene, middle Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFBC6773F2BD01FBFF0BFBE3.taxon	etymology	Etymology. In honor of the daughter of B. L.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFBC6773F2BD01FBFF0BFBE3.taxon	diagnosis	Diagnosis. Minute, moderately slender ovate, glossy shell with flattish protoconch, conical spire; sculpture of prominent spiral cords with narrow, punctate grooves; columella with two very prominent, sharp folds abapically and small, prominent parietal denticle.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFBC6773F2BD01FBFF0BFBE3.taxon	description	Description. Shell minute, moderately globose, broad (SL / MD = 1.4 – 1.6), with moderately low conical spire (AA = 48 – 52 °). Protoconch of one smooth, convex with flattish top. Teleoconch of 2.5 whorls separated by narrowly impressed, linear suture. Spire whorls moderately convex with periphery placed some distance above abapical suture. Sculpture of distinct, broad spiral cords separated by narrow, weakly punctate spiral grooves. Last whorl 77 – 79 % of total height, ovate, with periphery placed mid-whorl (SL / PH = 2.1 – 2.2); weakly constricted at base, glossy, with numerous, broad, flattish spiral cords separated by narrow grooves. Aperture ~ 60 % of total height, adapically moderately narrow, widening abapically (MD / MAW = 3.2 – 3.7). Outer lip very strongly thickened by wide labial varix bearing broad palatal swelling. Anal sinus broad, moderately deep, U-shaped; siphonal canal very short, open, broad, notched at tip. Columella with two prominent, sharp folds persisting on fasciole. Parietal denticle narrow but prominent; fused with broad parietal pad. Columellar and parietal callus weakly thickened, sharply delimited.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFBC6773F2BD01FBFF0BFBE3.taxon	discussion	Discussion. This species is characterized by its minute size. The fully developed apertural features exclude that the specimens are immature shells of another species. Ringicula minor (Grateloup, 1838) is comparable in outline and sculpture but differs in its larger size, the pointed protoconch and the thick parietal callus, which expands over the base. Ringicula laevigata (Eichwald, 1830) has similar apertural features but is larger and differs in its smooth surface. We are not aware of any other Ringiculidae from the European Neogene of comparable size. Paleoenvironment. Unknown.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFBC6773F2BD01FBFF0BFBE3.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Făget Basin: Nemeşeşti (Romania) (hoc opus).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFBE674DF2BD0397FF37FF03.taxon	description	Figs 13 E, 15 F 1 – F 2	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFBE674DF2BD0397FF37FF03.taxon	materials_examined	Type material. Holotype, NHMW-ZOO-MO- 75000 - E- 57968, SL. 7.3 mm, MD: 4.8 mm, Bad Vöslau (Austria), illustrated in Berger (1954: pl. 10, fig. 51), Fig 13 E, 15 F 1 – F 2 2.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFBE674DF2BD0397FF37FF03.taxon	description	Revised description. Shell medium sized, globose, broad (SL / MD = 1.5), with high conical spire (AA = 53 °). Protoconch low conical, paucispiral, of about 1.5 whorls (dp = 350 μm). Teleoconch of four whorls separated by narrowly impressed, linear suture. Spire whorls relatively high, weakly convex. Distinct spiral cords on first two teleoconch whorls, rapidly weakening abapically. Last whorl 71 % of total height, globose. Periphery placed mid-whorl (SL / PH = 2.0); not constricted at base, glossy, with several blurred, convex, narrow spiral cords separated by smooth, wide interspaces over base. Several narrow spiral cords over base. Aperture 74 % of total height, adapically moderately narrow, widening abapically (MD / MAW = 3.6). Outer lip very strongly thickened by wide labial varix with broad, poorly delimited palatal swelling bearing stout denticles at inner edge. Anal sinus broad, moderately deep, U-shaped; siphonal canal very short, open, broad, notched at tip. Columella with two strongly elevated, relatively thin folds abapically, parietal denticle prominent, fused with broad, thickened parietal pad, overhanging and narrowing aperture adapically. Columellar and parietal callus thickened, sharply delimited, broadly expanded and thickened over medial half of venter. Color pattern of thin dark stripes, coinciding with spiral cords on last whorl.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFBE674DF2BD0397FF37FF03.taxon	discussion	Discussion. This rare species is reminiscent of Ringicula hoernesii (Seguenza, 1881), with which it co-occurs. Especially apertural features and sculpture are almost identical. However, Ringicula dentata Berger, 1954 differs in its considerably higher conical spire and the denticles within the outer lip. Similarly, Ringicula guzhovi Landau, Harzhauser & Malaquias, 2023 differs mainly by its smooth outer lip and marked palatal swelling but without denticles. Berger (1954) mentioned several specimens, but we could only trace the holotype. We tentatively follow Berger (1954) and separate this shell from its congeners based on the peculiar dentition on the outer lip and color pattern, not seen in its Paratethyan congeners. Paleoenvironment. The occurrence in the Baden Formation at Bad Vöslau (Austria) suggests middle to outer neritic environments with up to 210 m water depth (Kranner et al. 2021).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFBE674DF2BD0397FF37FF03.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienna Basin: Bad Vöslau (Austria) (Berger 1954).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF80674CF2BD0637FE3DFAF7.taxon	description	Figs 13 C – D, 15 A – D	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF80674CF2BD0637FE3DFAF7.taxon	materials_examined	Type material. Lectotype (designated herein): PSM no. 3 / 466 - 1, SL: 5.6 mm, MD: 3.7 mm, Zhukivtsi (Ukraine), illustrated in Guzhov (2022 a: pl. 9, fig. 8). Paralectotypes: PSM no. 3 / 468, SL: 5.2 mm, SL: 3.7 mm, Zhukivtsi (Ukraine), illustrated in Guzhov (2022 a: pl. 9, fig. 9). Illustrated material. NHMW 1870 / 0014 / 0001, SL. 5.7 mm, MD: 3.8 mm, Golubytsya (Hołubica) (Ukraine), Figs 13 C, 15 A 1 – A 2. NHMW 1870 / 0014 / 0001 a, SL. 5.5 mm, MD: 3.9 mm, Golubytsya (Hołubica) (Ukraine), Figs 15 B 1 – B 2. NHMW 1870 / 0014 / 0001 b, SL. 5.3 mm, MD: 4.1 mm, Golubytsya (Hołubica) (Ukraine), Figs 15 C 1 – C 2. NHMW-ZOO-MO- 75000 - E- 57957, SL: 8.8 mm, MD: 5.8 mm, Steinebrunn (Austria), Figs 13 D, 15 D 1 – D 2.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF80674CF2BD0637FE3DFAF7.taxon	description	Original description. Because the illustration Eichwald (1829) is very poor and because of the confusion in the literature concerning the status of this species, we provide the description given by Eichwald (1853: 258, translated from Latin): “ Small ventricose shell, gibbous-ovate, very finely transversely striate, apex very acute, aperture longer up to penultimate whorl, wider in its middle, margin reflexed, thickened, base emarginate, columellar margin triplicate; var. exilis with thin, very acute whorls, last whorl suddenly increasing. ” Revised description. Shell medium sized, moderately to strongly globose, very broad (SL / MD = 1.3 – 1.5), with moderately high conical spire (AA = 60 – 73 °). Protoconch pointed, paucispiral of 1.5 convex whorls (dp = 350 – 400 μm). Teleoconch of four whorls separated by narrowly impressed, linear suture. Spire whorls moderately convex with periphery just above abapical suture. Sculpture of weak spiral cords with narrow spiral grooves. Last whorl 74 – 78 % of total height, strongly globose. Periphery placed just above mid-whorl (SL / PH = 1.9 – 2.0); not constricted at base. Sculpture variable ranging from nearly smooth to numerous, broad, flattish spiral cords separated by narrow grooves. Aperture 66 – 72 % of total height, adapically moderately narrow, widening abapically (MD / MAW = 3.3 – 3.7). Outer lip strongly thickened by wide labial varix bearing broad palatal swelling within. Anal sinus moderately deep, narrowly U-shaped; siphonal canal very short, open, broad, notched at tip. Columella with two prominent, robust folds abapically, parietal denticle weak to subobsolete placed on thickened parietal pad. Columellar and parietal callus thickened, sharply delimited, slightly expanding over base.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF80674CF2BD0637FE3DFAF7.taxon	discussion	Discussion. Ringicula guzhovi Landau, Harzhauser & Malaquias, 2023 displays considerable variability concerning its spiral sculpture and is characterized by its weak parietal denticle, relatively high spire and globose last whorl. Our material from the type region comprises nearly smooth shells, however Guzhov (2022 a) illustrated several specimens with distinct spiral grooves from the same region. Ringicula hoernesii (Seguenza, 1881) has a similarly inflated last whorl but differs in its lower and broader spire and narrower columellar folds. Ringicula minor (Grateloup, 1838) agrees in its spire morphology but is smaller and has a narrower and less convex last whorl. Specimens from the Pliocene of Viale (Italy), identified by Sacco as Ringicula auriculata var. exilis (1904: pl. 24, figs 29 – 30), differ in their elongate outline and prominent parietal denticle and probably represent a morph of the highly variable R. auriculata (Ménard de la Groye, 1811) (see Landau et al. 2023: 371, p. 25). Similarly, a specimen from the Miocene of France, illustrated by Morlet (1878: pl. 7, fig. 5) as R. exilis, most probably represents another species. Paleoenvironment. Shallow marine, inner neritic based on the associated fauna (own data, M. H.).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF80674CF2BD0637FE3DFAF7.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Roztocze Hills: Węglinek, (Poland) (Krach 1981); Korytnica Basin: Korytnica (Poland) (Friedberg 1938); Zhukivtsi, Trostyanets, Yaseniv (Ukraine) (Guzhov 2022 a); Voronyaky Hills: Golubytsya (Hołubica), Oles’ko, Pidgirtsi, Yaseniv (Ukraine) (Friedberg 1938); Polish-Ukrainian Fore-Carpathian Basin: Borki Wielkie, Chomentów (Poland) (Friedberg 1938); Tarnoruda, Shushkivtsi, Zalistsi, Zhabyak, Zhukivtsi, Zboriv (Ukraine) (Friedberg 1938); North Alpine Foreland Basin: Grund, Guntersdorf (Austria) (Berger 1954);? Žabčice (Czech Republic) (Montanaro Gallitelli & Tacoli 1951); Vienna Basin: Enzesfeld, Gainfarn, Steinebrunn, Vienna / Pötzleinsdorf (Austria) (Berger 1954); Borský Mikuláš (Czech Republic) (Švagrovský 1984); Pannonian Basin: Várpalota (Strausz 1966); Southern Pannonian Basin: Hrvaćani, Milijevići (Bosnia and Herzegovina) (Atanacković 1963, 1985); Southern Carpathians: Bahna (Romania) (Tiţă 2007); Făget Basin: Nemesti (Romania) Duşa 1967). Proto-Mediterranean Sea.? Serravallian (Middle Miocene): Karaman Basin: Akpınar, Lale, Seyithasan (Turkey) (Landau et al. 2013). Eastern Paratethys. Konkian (Middle Miocene): Ciscaucasia: Chumnaya, Maikopskaya Kuzhora, Vetrennaya (Republic of Adygea, Russia).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF81674EF2BD0263FE13FDB7.taxon	description	Figs 16 A – D	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF81674EF2BD0263FE13FDB7.taxon	materials_examined	Type material. Syntype, SL. 9.0 mm, MD: 6.4 mm, Baden (Austria), illustrated in Seguenza (1881: pl. 1, figs 4, 4 a – b). We are not aware of the whereabouts of the type material. Illustrated material. NHMW 1846 / 0037 / 0090, SL: 8.4 mm, MD: 6.0 mm, Baden (Austria), Figs 16 A 1 – A 2. NHMW 1846 / 0037 / 0090 a, SL: 8.2 mm, MD: 5.9 mm, Baden (Austria), Figs 16 B 1 – B 2. NHMW 1855 / 0045 / 0866, SL: 8.0 mm, MD: 5.5 mm, Bad Vöslau (Austria), Figs 16 C 1 – C 2. NHMW 1862 / 0001 / 0195, SL: 5.9 mm, MD: 4.3 mm, Baden (Austria), Figs 16 D 1 – D 2.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF81674EF2BD0263FE13FDB7.taxon	description	Revised description. Shell medium sized, very globose, with short conical spire (AA = 73 – 77 °). Protoconch paucispiral, pointed, of about 1.5 whorls with slightly oblique nucleus (dp = 410 μm). Junction with teleoconch marked by beginning of spiral sculpture. Teleoconch of four strongly convex whorls separated by narrowly impressed, linear suture. Weak spiral cords on first two teleoconch whorls, rapidly weakening abapically. Growth lines may form vaguely cancellate pattern on first teleoconch whorl. Last whorl 80 – 85 % of total height, strongly globose, with periphery placed around mid-whorl (SL / PH = 1.4 – 1.8); not constricted at base, glossy, with several blurred, convex, narrow spiral cords separated by wide interspaces with more or less distinct spiral threads. Aperture 80 – 85 % of total height, narrow in upper third, widening abapically (MD / MAW = 3.5 – 3.8). Outer lip very strongly thickened by wide labial varix; anal sinus broad, moderately deep, U-shaped, not distorting adapical end of outer lip; siphonal canal very short, open, broad, deeply notched at tip. Columella with two strongly elevated, relatively thin folds abapically, parietal denticle subobsolete or forming small denticle; fused with broad, robust, parietal pad overhanging and narrowing aperture adapically. Columellar and parietal callus thickened, sharply delimited, widely expanded and thickened over medial half of venter.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF81674EF2BD0263FE13FDB7.taxon	discussion	Discussion. Ringicula hoernesii Seguenza, 1881 is very common in offshore clays of the Vienna Basin and was confused by most authors with the Mediterranean Pliocene Ringicula buccinea (Brocchi, 1814), until Ceulemans et al. (2018) and Landau et al. (2023) excluded the Paratethyan records. Ringicula buccinea differs in its thicker columellar folds and weaker to subobsolete spiral sculpture on the last whorl. It is surprising that until now this taxon has been consistently overlooked, considering that Seguenza (1881) introduced Ringicula hoernesii as name for the Austrian shells, even illustrating a syntype from Baden (Austria). This is undoubted due to the very broad species concept adopted by most authors for R. buccinea, rejected by us. Paleoenvironment. The occurrence in the Baden Formation at Bad Vöslau (Austria) suggests middle to outer neritic environments with up to 210 m water depth (Kranner et al. 2021).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF81674EF2BD0263FE13FDB7.taxon	distribution	Distribution in Central Paratethys. Polish-Ukrainian Fore-Carpathian Basin: Błonie (Poland) (Friedberg 1938; Urbaniak 1974); Vienna Basin: Baden, Bad Vöslau, Baden Sooss, Möllersdorf (Austria) (hoc opus); Bükk Mountains: Borsodbóta (Hungary) (Csepreghy-Meznerics 1972); Southern Pannonian Basin: Miljevići (Bosnia and Herzegovina) (Atanacković 1985); Dacian Basin: Portitovtsi, Urovene (Bulgaria) (Kojumdgieva in Kojumdgieva & Strachimirov 1960). Proto-Mediterranean Sea. Seguenza (1881) mentioned an incomplete specimen from the Tortonian of Sassuolo (Italy) with question mark.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF836748F2BD05A3FF37FA17.taxon	description	Figs 13 F – H, 17 A – H	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF836748F2BD05A3FF37FA17.taxon	materials_examined	Type material. We are not aware of the whereabouts of the type material. Guzhov (2022 a) was not able to trace the syntypes in the Eichwald collection stored in the Borissiak Paleontological Institute (Russian Academy of Sciences) in Moscow (Russia). Illustrated material. NHMW 1949 / 0005 / 0009, SL: 2.9 mm, MD: 1.7 mm, Steinebrunn (Austria), syntype of Ringicula hochstetteri Hoernes & Auinger, 1880, illustrated in Hoernes & Auinger (1880: pl. 8, fig. 20), Figs 13 F, 17 A 1 – A 2. NHMW 1949 / 0005 / 0009, SL: 3.5 mm, MD: 1.9 mm, Steinebrunn (Austria), syntype of Ringicula hochstetteri Hoernes & Auinger, 1880, illustrated in Hoernes & Auinger (1880: pl. 8, fig. 21), Figs 17 B 1 – B 2. NHMW-ZOO-MO- 75000 - E- 57969, SL: 2.8 mm, MD: 1.5 mm, Grund (Austria), Figs 17 C 1 – C 2. NHMW-ZOO-MO- 75000 - E- 57964, SL: 3.8 mm, MD: 2.1 mm, Steinebrunn (Austria), illustrated in Berger (1954: pl. 8, fig. 25), Figs 13 G, 17 D 1 – D 2. NHMW-ZOO-MO- 75000 - E- 57966, SL: 3.3 mm, MD: 1.7 mm, Ritzing (Austria), illustrated in Berger (1954: pl. 8, fig. 30), Figs 13 H, 17 E 1 – E 2. NHMW 2018 / 0248 / 0490, SL: 2.7 mm, MD: 1.5 mm, Coşteiu de Sus (Romania), Figs 17 F 1 – F 2. NHMW 2018 / 0248 / 0490, SL: 2.8 mm, MD: 1.5 mm, Coşteiu de Sus (Romania), Figs 17 G 1 – G 2. NHMW 2018 / 0248 / 0490, SL: 2.6 mm, MD: 1.4 mm, Coşteiu de Sus (Romania), Figs 17 H.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF836748F2BD05A3FF37FA17.taxon	description	Revised description. Shell small, glossy, moderately broad fusiform, with relatively high conical spire (AA = 50 – 57 °). Protoconch paucispiral, conical, of about 1.5 whorls (dp = ~ 300 μm). Teleoconch of four whorls separated by narrowly impressed to narrowly canaliculate, linear suture. Spire whorls weakly convex, with periphery placed just above abapical suture. Weak to subobsolete spiral cords on first two teleoconch whorls, rapidly weakening abapically. Last whorl 71 – 74 % of total height, moderately convex, with periphery placed mid-whorl (SL / PH = 2.2); weakly constricted at base. Last whorl smooth, glossy. Aperture 60 – 64 % of total height, narrowing in upper third, widening abapically (MD / MAW = 3.7 – 4.4). Point of insertion of outer lip far below adapical suture. Outer lip moderately thickened by convex labial varix bearing broad, central palatal swelling; anal sinus narrow, moderately deep, V-shaped, terminating at adapical end of outer lip; siphonal canal very short, open, broad, deeply notched at tip. Columella with two strongly elevated, folds abapically; parietal denticle prominent or weak; fused with broad, robust parietal pad. Columellar and parietal callus thickened, sharply delimited, narrowly expanded over base. One specimen shows reddish flammulae on last spire whorl (Fig 17 A 2). Paratethyan synonyms. Ringicula hochstetteri Hoernes & Auinger, 1880, four illustrated syntypes from Steinebrunn (Austria) and Coşteiu de Sus (Romania). Hoernes & Auinger (1880) did not refer to Ringicula laevigata (Eichwald, 1830) when establishing Ringicula hochstetteri. As already stated by Berger (1954), the shells do not differ from Ringicula laevigata and Ringicula hochstetteri is a subjective junior synonym. Paleoenvironment. Shallow marine, inner neritic (own data M. H).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF836748F2BD05A3FF37FA17.taxon	discussion	Discussion. Ringicula laevigata (Eichwald, 1830) was discussed in detail by Guzhov (2022 a: 1137). Slender specimens with high spire were described by Berger (1954) as Ringicula acutior Morlet, 1878, which was originally described from the Tortonian (Late Miocene) of Saint-Jean-de-Marsacq (France). The French species is larger (SL: 5 mm) and differs in its more convex last whorl (see Morlet 1878: 268, pl. 7, fig. 6). Glibert (1952 a: 390, pl. 14, fig 11 f) illustrated a specimen from the Langhian (Middle Miocene) of the Loire Basin (France) as Ringicula acutior. It represents a rather small Ringicula species with an unusually tall spire and seems to have only the adapical of the two basal columellar folds usually seen in members of the genus. We have not encountered any further specimens from the area discussed by Glibert, and we suspect this to be a teratogenic specimen. In any case, it is not conspecific with Ringicula laevigata.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF836748F2BD05A3FF37FA17.taxon	distribution	Distribution in Central Paratethys. Karpatian (Early Miocene): Korneuburg Basin: Niederkreuzstetten (Austria) (Berger 1954). Badenian (Middle Miocene): Ukrainian Fore-Carpathian Basin: Varivtsi, Zalistsi, Zhabyak, Zhukivtsi (Ukraine) (Friedberg 1938; Guzhov 2022 a); North Alpine-Carpathian Foreland Basin: Grund, Guntersdorf (Austria), Hrušovany nad Jevišovkou (Czech Republic) (Berger 1954); Vienna Basin: Bad Vöslau, Enzesfeld, Gainfarn, Niederleis, Poysdorf, Steinebrunn (Austria), Mikulov (Czech Republic) (Berger 1954); Oberpullendorf Basin: Ritzing (Austria) (Berger 1954); Pannonian Basin: Várpalota (Hungary) (Strausz 1966); Southern Pannonian Basin: Miljevići (Bosnia and Herzegovina) (Atanacković 1969); Hațeg Depression: Răchitova (Romania) (Popa & Ianoliu 2000); Făget Basin: Buituri, Coşteiu de Sus, Lăpugiu de Sus (Romania) (Hoernes & Auinger 1880; Boettger 1902); Dacian Basin: Urovene, Trifonovo, Dobrusha (Bulgaria) (Kojumdgieva in Kojumdgieva & Strachimirov 1960); Islaz (Romania) (Rado & Muţui 1970). Eastern Paratethys Sea. Konkian (Middle Miocene): Agara (Georgia), Chumnaya (Republic of Adygea), Aksengir, Karatyulei, Mynsualmas (Kazakhstan) (Guzhov 2022 a). Proto-Mediterranean Sea. Serravallian (Middle Miocene): Karaman Basin: Lale (Turkey) (Landau et al. 2013).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF85674AF2BD0103FDE7FDB7.taxon	description	Figs 14 A – E	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF85674AF2BD0103FDE7FDB7.taxon	materials_examined	Type material. Syntype illustrated in Grateloup (1838: 286, pl. 6, fig. 8), Aquitaine Basin, no type locality given, Early Miocene. Illustrated material. NHMW-ZOO-MO- 75000 - E- 57962, SL: 5.8 mm, MD: 3.7 mm, Bad Vöslau (Austria), Figs 15 E 1 – E 2. NHMW 1862 / 0001 / 0521, SL: 6.1 mm, MD: 3.8 mm, Buituri (Romania), Figs 14 A 1 – A 2. NHMW 1862 / 0001 / 0521 a, SL: 6.0 mm, MD: 3.6 mm, Buituri (Romania), Figs 14 B 1 – B 2. NHMW 1850 / 0002 / 0012, SL: 5.0 mm, MD: 3.1 mm, Grund (Austria), Figs 14 C 1 – C 2. NHMW 1862 / 0001 / 0521 b, SL: 4.9 mm, MD: 2.9 mm, Buituri (Romania), Figs 14 D 1 – D 2.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF85674AF2BD0103FDE7FDB7.taxon	description	Revised description. Shell medium sized, moderately globose, broad (SL / MD = 1.5 – 1.6), with high conical spire (AA = 58 – 62 °). Protoconch unknown. Teleoconch of four whorls separated by narrowly impressed, linear suture. Spire whorls moderately convex with periphery placed some distance above abapical suture. Sculpture of distinct, broad spiral cords separated by narrow spiral grooves. Last whorl 70 – 73 % of total height, globose, with periphery placed mid-whorl (SL / PH = 2.0 – 2.1); weakly constricted at base, glossy, with numerous, broad, flattish spiral cords separated by narrow grooves. Aperture 64 – 66 % of total height, adapically moderately narrow, widening abapically (MD / MAW = 3.6 – 4.6). Outer lip very strongly thickened by wide labial varix bearing broad palatal swelling. Anal sinus broad, moderately deep, U-shaped; siphonal canal very short, open, broad, notched at tip. Columella with two prominent, robust folds abapically, parietal denticle prominent or weak; fused with broad parietal pad. Columellar and parietal callus thickened, sharply delimited, slightly expanded over base.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF85674AF2BD0103FDE7FDB7.taxon	discussion	Discussion. Ringicula minor (Grateloup, 1838) differs from Ringicula guzhovi Landau, Harzhauser & Malaquias, 2023 in its higher spire, less inflated last whorl and prominent spiral sculpture. See Landau et al. (2013: 326) for further discussion. Paleoenvironment. Shallow marine, inner neritic (own data M. H).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF85674AF2BD0103FDE7FDB7.taxon	distribution	Distribution in Central Paratethys. Eggenburgian (Early Miocene): North Alpine Foreland Basin: Fels am Wagram (Austria) (Steininger 1963). Karpatian (Early Miocene): North Alpine-Carpathian Foreland Basin: Laa (Austria) (Berger 1954), Malé Karpaty Mts.: Cerová (Slovakia) (Harzhauser et al. 2011); Korneuburg Basin: Stetten (Austria) (Harzhauser 2002). Badenian (Middle Miocene): North Alpine-Carpathian Foreland Basin: Kalladorf, Grund, Guntersdorf (Austria) (Berger 1954); Badenian (Middle Miocene): Voronyaky Hills: Oles’ko (Ukraine) (Friedberg 1928); Korytnica Basin: Korytnica (Poland) (Friedberg 1928); Vienna Basin: Borský Mikuláš (Czech Republic) (Švagrovský 1984); Bad Vöslau, Steinebrunn, Vienna / Grinzing, Vienna / Pötzleinsdorf (Austria) (Berger 1954); Boršov (Porstendorf), Mikulov, Jaroměřice (Czech Republic) (Berger, 1954); Oberpullendorf Basin: Ritzing (Austria) (Berger 1954); Pannonian Basin: Herend-Márkó, Letkés, Várpalota, Szob (Hungary) (Strausz 1954; Csepreghy-Meznerics 1956; Kókay 1966); Southern Pannonian Basin: Miljevići (Bosnia and Herzegovina) (Atanacković 1985); Făget Basin: Buituri (Romania) (hoc opus). Proto-Mediterranean Sea. Serravallian (Middle Miocene): Karaman Basin: Akpınar, Lale, Seyithasan (Turkey) (Landau et al. 2013). Tortonian (Late Miocene): Livorno Hills: Quarata (Italy) (Dominici et al. 2020); Calabria: Cessaniti (Italy) (D’Amico et al. 2012). Northeastern Atlantic. Burdigalian (Early Miocene): Aquitaine Basin: Cestas, Léognan, Martillac, Mérignac, Saucats, Pourquey, Peloua, Saint-Paul-lès-Dax (France) (Peyrot 1932; Lesport & Cahuzac 2005). Serravallian (Middle Miocene): Salles, Sallespisse, Salies-de-Béarn (France) (Peyrot 1932). Tortonian (Late Miocene): Loire Basin (France) (Landau et al. 2020).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF87674AF2BD032FFE5FF997.taxon	materials_examined	Type material. Holotype, SL: 2.65 mm, MD: 2.12 mm, specimen illustrated in Bałuk & Jakubowski 1968: 294, pl. 1, figs 4 a – b; Korytnica (Poland), stored in the Faculty of Geology, University of Warsaw, Poland.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF87674AF2BD032FFE5FF997.taxon	description	Description. See Bałuk & Jakubowski (1968: 294).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF87674AF2BD032FFE5FF997.taxon	discussion	Discussion. No new material was found since the first description of this rare or overlooked bivalved gastropod and we refer to Bałuk & Jakubowski (1968) for description and illustrations. For discussion see Le Renard et al. (1996) and Schneider et al. (2008). Paleoenvironment. Extant Juliidae appear to feed exclusively on Caulerpa algae (Schneider et al. 2008).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF87674AF2BD032FFE5FF997.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Korytnica Basin: Korytnica (Poland) (Bałuk & Jakubowski 1968).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF87674AF2BD049BFA50FC3B.taxon	type_taxon	Type species. Candinia pliocaenica Le Renard, Sabelli & Taviani, 1996; by original designation. Pliocene, Italy.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF876745F2BD007BFE69FF03.taxon	type_taxon	Type species. Bulla ampulla Linnaeus, 1758; as ruled by Opinion 196 (ICZN 1954 a). Present-day, Indo-West Pacific.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF876745F2BD007BFE69FF03.taxon	discussion	Discussion. For a detailed discussion on the complex taxonomic and nomenclatorial history of the genus see Malaquias & Reid (2008).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF886745F2BD0608FEB6FABD.taxon	description	Figs 18 B 1 – B 3	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF886745F2BD0608FEB6FABD.taxon	materials_examined	Type material. Holotype, NHMW 1859 / 0045 / 0420, SL: 3.3 mm, MD: 2.1 mm, Oles’ko (Lviv Region, Ukraine), Figs 18 B 1 – B 3. Type locality. Oles’ko (Ukraine). Type stratum. Unknown. Age. Middle Miocene, late Badenian (Serravallian).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF886745F2BD0608FEB6FABD.taxon	etymology	Etymology. In honor of Abder El Albani (Université de Poitiers), who revolutionized our understanding of the evolution of early life.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF886745F2BD0608FEB6FABD.taxon	diagnosis	Diagnosis. Small, biconic-ovate with truncated, narrowly umbilicate apex and conical sides above periphery.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF886745F2BD0608FEB6FABD.taxon	description	Description. Shell small, thin-shelled, biconic-ovate, broad (SL / MD = 1.5). Only last whorl visible; convex periphery above mid-whorl (SL / PH = 1.7), whorl sides nearly straight above periphery forming angle of about 58 °. Moderately narrow, truncated apex (MD / ApD = 2.8); with deep, narrow umbilicus; apical plane weakly oblique (AP = 8 °). Smooth except for weak growth lines. Aperture moderately narrow in apical third (MS / AW = 9.7), widening greatly anteriorly (MD / MAW = 2.5), resulting in moderate expansion rate (ER = 3.9). Outer lip thin, not rising above apex, strongly convex at periphery, rounded abapically. Columella short, nearly straight, without fold, forming narrow columellar callus, delimiting narrow umbilical chink. Columellar callus passing into narrow, thin parietal callus.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF886745F2BD0608FEB6FABD.taxon	discussion	Discussion. We are not aware of comparable species from the circum-Mediterranean region. The placement in Bulla is tentative and is based on the broad, open, involute spire, whereas the profile differs from typical Bulla. The ovate-biconical outline is reminiscent of some extant Relichna species [e. g., Relichna sumatrana (Thiele, 1925), from New Caledonia, and Relichna sp. from Vanatu in Valdés 2008: figs 84 E – I, 88 G – H]. Relichna, however, is unknown from Paratethyan deposits and therefore, we assume that this species rather belongs to Bulla and is a further endemic species from the Polish-Ukrainian foredeep as documented for many other gastropod families (Harzhauser et al. 2024). It is quite likely that this species represents a juvenile shell. However, the form is quite unlike juveniles of any other European Neogene Bulloidea, which has led us to formally describe the species, despite the scant and probably immature nature of the specimen. Paleoenvironment. Inner neritic, shallow marine according to accompanying mollusk assemblage. Central Paratethys. Badenian (Middle Miocene): Polish-Ukrainian Fore-Carpathian Basin: Oles’ko (Lviv Region, Ukraine).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF896744F2BD0786FE86FBBE.taxon	description	Fig. 18 A	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF896744F2BD0786FE86FBBE.taxon	materials_examined	Type material. Lectotype (designated by Zilch 1934: 277), SMF 363157 - 12.2550 a (glass plate 395), SL: 7.5 mm, MD: 2.3 mm, Coșteiu de Sus (Romania), illustrated in Zilch (1934: pl. 22, fig. 14). The specimen was destroyed during WW II, Fig. 18 A.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF896744F2BD0786FE86FBBE.taxon	description	Revised description. Shell medium sized, moderately broad, ovate (SL / MD = 1.8). Only last whorl visible; periphery just above mid-whorl (SL / PH = 1.9). Low, truncated, moderately narrow apex (MD / ApD = 2.4), with weakly oblique apical plane (AP = 8 °); deep and funnel-shaped umbilicus, delimited by round-edged, elevated crest. Sculpture of weak, flattened spiral cords over base separated by shallow grooves, weakening towards mid-whorl, mid-portion and apex smooth. Aperture moderately wide in apical third (MD / AW = 8.4), widening moderately abapically (MD / MAW = 2.2), resulting in moderate expansion rate (ER = 3.8). Outer lip thin, slightly rising above apex, moderately convex at periphery, rounded and slightly flaring abapically. Columella short, moderately excavated. Columellar callus forming broad, detached rim, delimiting narrow, umbilical chink. Parietal region covered by sediment. Color pattern of axially arranged dark blotches, separated by slightly wider light stripes.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF896744F2BD0786FE86FBBE.taxon	discussion	Discussion. Bulla hungarica Boettger, 1907 is reminiscent of the Pliocene to Recent Bulla striata Bruguière, 1792, which is larger, and its truncated apex is narrower resulting in a more convex adapical half of the last whorl. The Pliocene Mediterranean Bulla subampulla d’Orbigny, 1852, is also larger and differs in its much narrower apex and more globose shape (see Landau et al. 2023: pl. 31). Paleoenvironment. Unknown.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF896744F2BD0786FE86FBBE.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Făget Basin: Coşteiu de Sus (Romania) (Boettger 1907).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF896744F2BD02C8FE8DF952.taxon	type_taxon	Type species. Bulla pyriformis A. Adams, 1850; typification of replacement name. Present-day, Chinese Sea [nom. nov. pro Sao H. & A. Adams, 1854, non Billberg, 1820 (Crustacea), nec Barrande, 1846 (Trilobita), nec Kölliker, 1853 (Cnidaria)].	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF896744F2BD03A8FEA2FA5E.taxon	discussion	Comment. The columella in the family Retusidae may or may not bear a structure called by most authors a ‘ fold’. Rather than being columellar fold / s along the columellar inner edge, as seen in the Volutoidea, this fold seems to be a terminal columellar ridge delimiting the medial border of the siphonal canal. For consistency, we continue to call it a columellar fold.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF896741F2BD01A0FED3FCEF.taxon	description	Figs 19 A – E	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF896741F2BD01A0FED3FCEF.taxon	materials_examined	Type material. Lectotype (designated herein): specimen illustrated in Eichwald (1851: pl. 11, figs 15 a – c). probably from Żukowce (Zhukivtsi) (or Tarnoruda, Zaleśce) (Ukraine). Probably stored in the St. Petersburg State University, Paleontological Museum (Russia). Illustrated material. NHMW 2010 / 0004 / 1656, SL: 5.7 mm, MD: 2.3 mm, Bad Vöslau (Austria), Figs 19 A 1 – A 3. NHMW 2010 / 0004 / 1656 a, SL: 6.0 mm, MD: 2.2 mm, Bad Vöslau (Austria), Figs 19 B 1 – B 3. NHMW 1869 / 0001 / 0775, SL: 4.8 mm, MD: 2.0 mm, Coşteiu de Sus (Romania), Figs 19 C 1 – C 2. NHMW 2010 / 0004 / 1656 b, SL: 5.1 mm, MD: 2.2 mm, Bad Vöslau (Austria), Figs 19 D 1 – D 2. NHMW-ZOO-MO- 75000 - E- 58763, SL: 6.6 mm, MD: 2.5 mm, Gainfarn (Austria), Figs 19 E 1 – E 3.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF896741F2BD01A0FED3FCEF.taxon	description	Revised description. Shell small, medium-thickness, slender (SL / MD = 2.5 – 2.6), elongate pyriform, only last whorl visible. Last whorl with almost straight sides, forming angle of about 19 – 21 °. Periphery placed close to one-third whorl height (SL / PH = 3.0 – 3.6), rounded at moderately contracting base. Apex moderately narrow (MD / ApD = 2.5 – 2.6), truncated, apical plane slightly oblique (AP = 12 – 15 °), deeply and narrowly umbilicate, delimited by round-edged, faintly swollen, elevated crest. Sculpture of a few, faint spiral grooves over base. Aperture longer than shell, very narrow down to periphery (MD / AW = 13.8 – 15.7), abruptly widening anteriorly below periphery (MD / MAW = 1.9 – 2.2), resulting in high expansion rate (ER = 6.6 – 8.0). Outer lip thin, slightly alate apically, extending some distance above apex, weakly concave just above periphery. Columella short, with narrow but prominent columellar callus with low, broad columellar fold. Columellar callus short, narrow, delimiting deep umbilical chink. Parietal callus not developed.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF896741F2BD01A0FED3FCEF.taxon	discussion	Discussion. This is a very distinctive species due to its slender conical outline and weak columellar fold. Nevertheless, the name was erroneously used for many Miocene shells, which in our opinion are not conspecific with Pyrunculus elongatus (Eichwald, 1830). Middle Miocene specimens from the northeastern Atlantic and North Sea illustrated by Peyrot (1932), Glibert (1952 b), A. W. Janssen (1984) and many others listed above are comparatively stout with more convex whorls. We have not reviewed this group but note that Pyrunculus subconulus (d’Orbigny, 1852) might be an available name for at least some of these specimens. A comment on Pyrunculus hoernesi (Weinkauff, 1866). Weinkauff (1866: 238) established Bulla (Cylichna) hoernesi when describing extant specimens from Algeria but referred to the illustration of a Middle Miocene specimen from the Vienna Basin illustrated in Hörnes (1856: pl. 50, figs 4 a – b) as Bulla conulus Deshayes, 1824 [= Pyrunculus elongatus (Eichwald, 1830)]. Thus, as pointed out by Tringali & Oliverio (2001: 157), this Miocene specimen belongs to the type material of Bulla (Cylichna) Hoernesi following ICZN 1999 Art., 72.4. No lectotype was designated by Tringali & Oliverio (2001) from the Algerian shells and therefore the status of Pyrunculus hoernesi (Weinkauff, 1866) is still problematic. Choosing the Paratethyan specimen as lectotype would make Bulla hoernesi Weinkauff, 1866 a subjective junior synonym of Bulla elongata Eichwald, 1830. Therefore, we select the syntype stored in the Zoological Museum Rome (Italy), Monterosato coll. box 16118, as lectotype. The specimen was illustrated in Tringali & Oliverio (2001: fig. 16), SL: 2.8 mm, MD: 1.3 mm, Alger (Algeria). Paleoenvironment. Occurs in inner neritic to outer neritic assemblages.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF896741F2BD01A0FED3FCEF.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Voronyaky Hills: Golubytsya (Hołubica), Yaseniv (Friedberg 1938); Korytnica Basin: Korytnica (Poland) (Friedberg 1938); Polish-Ukrainian Fore-Carpathian Basin: Wieliczka (Poland) (Reuss 1867); Fashchivka, Staryi Pochaiv, Tarnoruda, Zaleśce (Żabiak), Zhukivtsi (Ukraine) (Friedberg 1938); Vienna Basin: Baden, Gainfarn, Steinebrunn, Vienna / Pötzleinsdorf (Austria) (Bergen 1953); Pannonian Basin: Letkés, Szob, Szokolya (Hungary) (Báldi 1960; Strausz 1966); Făget Basin: Coşteiu de Sus, Lăpugiu de Sus (Romania) (Boettger 1907). Proto-Mediterranean Sea. Serravallian (Middle Miocene): Karaman Basin: Seyithasan (Turkey) (Landau et al. 2023).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF8C6740F2BD046CFBD8FE5F.taxon	description	Figs 19 F, 20 A – C	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF8C6740F2BD046CFBD8FE5F.taxon	materials_examined	Type material. Lectotype (designated by Zilch 1934: 277), SMF 363160 - 12.2553 a (glass plate 398), SL: 6.0 mm, MD: 1.8 mm, Coşteiu de Sus (Romania), illustrated in Zilch (1934: pl. 22, fig. 17). The specimen was destroyed during WW II, Fig 20 C. Illustrated material. NHMW 1865 / 0015 / 0155, SL: 6.2 mm, MD: 3.3 mm, Lysice (Czech Republic), Figs 19 F, 20 A 1 – A 2. NHMW-ZOO-MO- 75000 - E- 47049, SL: 8.0 mm, MD: 4.0 mm, Lysice (Czech Republic), Figs 20 B 1 – B 2.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF8C6740F2BD046CFBD8FE5F.taxon	description	Revised description. Shell medium sized, solid, moderately broad (SL / MD = 1.9), only last whorl visible. Periphery placed below mid-whorl (SL / PH = 2.2), almost straight to faintly concave sides above periphery, forming angle of about 23 °. Rounded at moderately contracting base. Apex moderately broad (MD / ApD = 2.3 – 2.5), truncated, apical plane slightly oblique (AP = 6 °), deeply and narrowly umbilicate, delimited by sharply edged, trigonal crest. Surface smooth, slightly glossy. Growth lines delicate. Aperture only slightly longer than shell, very narrow down to mid-whorl (MD / AW = 11.2 – 11.3), distinctly widening anteriorly (MD / MAW = 2.0 – 2.4), resulting in high expansion rate (ER = 4.6 – 5.5). Outer lip thin, extending slightly above apex. Basal lip abapically slightly widening. Columella short, with narrow but prominent columellar callus bearing prominent central columellar fold. Columellar callus delimiting narrow umbilical chink. Parietal callus not developed.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF8C6740F2BD046CFBD8FE5F.taxon	discussion	Discussion. This rare species is characterized by its relatively large size, solid shell for the genus, broad-conical outline, trigonal apical crest, and prominent columellar fold. Pyrunculus frequens (Boettger, 1902) might be related to the Pliocene to recent Mediterranean Pyrunculus ovatus (Jeffreys, 1871), which may form very similar shapes and has a trigonal crest (see Tringali & Oliverio 2001: figs 1 – 14). Pyrunculus ovatus, however, is much thinner-shelled, has a narrower, less truncated apex and lacks a columellar fold (see Landau et al. 2023: pl. 32). Pyrunculus frequens is superficially reminiscent of the extant Mediterranean Pyrunculus hoernesi (Weinkauff, 1866), which differs in its prominent apical sculpture and the lack of a columellar fold. The Late Miocene to Early Pleistocene Mediterranean Cylichna brocchii (Michelotti, 1847) lacks the truncated apex of Pyrunculus frequens and is weakly convex above the periphery (see Landau et al. 2023: pl. 40, figs 1 – 2). Paleoenvironment. Probably middle to outer neritic based on the occurrence at Lysice (own data, M. H.).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF8C6740F2BD046CFBD8FE5F.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Carpathian Foredeep: Lysice (Czech Republic) (hoc opus); Făget Basin: Coşteiu de Sus (Romania) (Boettger 1907).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF8D6742F2BD0345FD60FE97.taxon	description	Figs 21 A – F	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF8D6742F2BD0345FD60FE97.taxon	materials_examined	Type material. Lectotype (designated by Zilch 1934: 277), SMF 363159 - 12.2552 a (glass plate 397), SL: 4.1 mm, MD: 1.8 mm, Coşteiu de Sus (Romania), illustrated in Zilch (1934: pl. 22, fig. 16), Fig. 21 C. The specimen was destroyed during WWII. Illustrated material. NHMW 1855 / 0045 / 0258, SL: 4.2 mm, MD: 1.9 mm, Steinebrunn (Austria), Figs 21 A 1 – A 3. NHMW 1855 / 0045 / 0258 a, SL: 4.6 mm, MD: 2.1 mm, Steinebrunn (Austria), (illustrated in Berger 1953: pl. 16, fig. 32), Figs 21 B 1 – B 3. NHMW-ZOO-MO- 75000 - E- 47032, SL: 2.4 mm, MD: 1.1 mm, Lăpugiu de Sus (Romania), Figs 21 D 1 – D 3. NHMW 2010 / 0004 / 1655, SL: 5.2 mm, MD: 2.2 mm, Bad Vöslau (Austria), Figs 21 E 1 – E 2. NHMW 1855 / 0045 / 0258 a, SL: 3.9 mm, MD: 1.8 mm, Steinebrunn (Austria), Figs 21 F 1 – F 2.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF8D6742F2BD0345FD60FE97.taxon	description	Revised description. Shell small to medium sized, medium-thickness, moderately slender (SL / MD = 2.2 – 2.3), elongate conical, only last whorl visible. Last whorl with almost straight to faintly concave sides, forming angle of about 16 °. Periphery below mid-whorl, placed just above lower third (SL / PH = 2.7), rounded at moderately contracting base. Apex relatively narrow (MD / ApD = 2.1 – 2.2), truncated, apical plane slightly oblique (AP = 11 – 12 °), deeply and narrowly umbilicate, delimited by round edged, faintly swollen, elevated crest. Sculpture of a few, faint, widely spaced spiral grooves, prominent over base, fading towards mid-whorl, subobsolete on adapical third. Growth lines prominent on apical crest. Aperture slightly longer than shell, very narrow down to periphery (MD / AW = 11.8 – 14.0), abruptly widening in anterior third below periphery (MD / MAW = 2.0 – 2.1), resulting in high expansion rate (ER = 5.8 – 6.7). Outer lip thin, extending some distance above apex, narrowly trigonally alate, weakly concave just above periphery. Columella short, with narrow but prominent columellar callus with subobsolete columellar fold. Columellar callus delimiting narrow umbilical chink. Parietal callus not developed.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF8D6742F2BD0345FD60FE97.taxon	discussion	Discussion. Until now, this species was poorly known because it was based on a fragmentary specimen without the outer lip. Later, Berger (1953) gave a drawing of a specimen from Steinebrunn (Austria), which strongly overemphasized spiral sculpture and the strength of the apical crest (see same specimen illustrated herein as Figs 21 B 1 – B 3). Pyrunculus latesulcatus (Boettger, 1902) is most like the extant Mediterranean Pyrunculus hoernesi (Weinkauff, 1866). Both have an identical shape and agree in their apertural features. The extant species has a similar variability in spiral sculpture but differs in the much more prominent axial folds or ribs on the apical crest. Paleoenvironment. The occurrence at Steinebrunn (Austria) suggests shallow marine environments (own data M. H.).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF8D6742F2BD0345FD60FE97.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienna Basin: Bad Vöslau, Steinebrunn (Austria) (hoc opus); Pannonian Basin: Pécsvárad, Tekeres (Hungary) (Bohn-Havas 1973); Făget Basin: Coşteiu de Sus, Lăpugiu de Sus (Romania) (Zilch 1934).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF8F6742F2BD0683FD2DF9DB.taxon	description	Figs 22 A – E	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF8F6742F2BD0683FD2DF9DB.taxon	materials_examined	Type material. Holotype, NHMW-ZOO-MO- 75000 - E- 11815 A, SL: 3.0 mm, MD: 1.5 mm, Bad Vöslau (Austria), Figs 22 A 1 – A 3. Paratypes, NHMW-ZOO-MO- 75000 - E- 11815, SL: 2.9 mm, MD: 1.4 mm, Bad Vöslau (Austria), Figs 22 B 1 – B 3. NHMW-ZOO-MO- 75000 - E- 11815, SL: 3.5 mm, MD: 1.7 mm, Bad Vöslau (Austria), Figs 22 C 1 – C 3. Illustrated material. NHMW-ZOO-MO- 75000 - E- 3308, SL: 4.1 mm, MD: 2.0 mm, Bad Vöslau (Austria), Figs 22 D 1 – D 3. NHMW-ZOO-MO- 75000 - E- 3308, SL: 4.3 mm, MD: 1.9 mm, Bad Vöslau (Austria), Figs 22 E 1 – E 3.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF8F6742F2BD0683FD2DF9DB.taxon	description	Revised description. Shell small, glossy, moderately slender (SL / MD = ~ 2.2), moderately pyriform, only last whorl visible. Last whorl with short conical apical region (47 – 50 °) passing into almost straight to weakly convex sides, forming angle of about 18 – 20 °, with periphery below mid-whorl (SL / PH = 2.7 – 2.8), rounded at moderately contracting base. Apex narrow (MD / ApD = 2.7 – 3.0), truncated, apical plane slightly oblique (AP = 10 – 12 °), deeply and narrowly umbilicate, delimited by round-edged, elevated crest. Sculpture of a few, shallow spiral grooves at both extremities, rarely persisting to mid-whorl. Aperture longer than shell, very narrow just below mid-whorl (MD / AW = 8.0 – 10.0), widening anteriorly (MD / MAW = 1.8 – 2.0), resulting in high expansion rate (ER = 4.0 – 5.4). Outer lip thin, apically extending some distance above apex, faintly pinched inwards above periphery. Columella short, with narrow columellar callus. Columellar fold, low, weak or subobsolete. Columellar callus adherent or erect, delimiting narrow umbilical chink. Parietal callus not developed.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF8F6742F2BD0683FD2DF9DB.taxon	discussion	Discussion. Pyrunculus leuchsi (Berger, 1949) is strongly reminiscent of Pyrunculus szobiensis sp. nov. concerning the changing angle between apical region and sides, but it is slenderer, the apex is narrower, and the mid-portion is less cylindrical. Pyrunculus parangistoma (Boettger, 1902) is even more conical and has a narrower apex and Pyrunculus elongatus (Eichwald, 1830) differs in its much higher and slenderer shell. Paleoenvironment. The occurrence in the Baden Formation at Bad Vöslau (Austria) suggests middle to outer neritic environments with up to 210 m water depth (Kranner et al. 2021).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF8F6742F2BD0683FD2DF9DB.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienna Basin: Bad Vöslau (Austria) (Berger 1953), Borský Mikuláš (Slovakia) (Švagrovský 1984).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF8F675CF2BD014FFE28FDCF.taxon	description	Figs 23 A – C, 24 G – H	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF8F675CF2BD014FFE28FDCF.taxon	materials_examined	Type material. Holotype, NHMW 2018 / 0248 / 0475, SL: 2.8 mm, MD: 1.2 mm, Coşteiu de Sus (Romania), Figs 23 A 1 – A 2, 24 G. Paratypes, NHMW 2018 / 0248 / 0475 a, SL: 2.9 mm, MD: 1.3 mm, Coşteiu de Sus (Romania), Figs 23 B 1 – B 2, 24 H. NHMW-ZOO-MO- 75000 - E- 147038, SL: 2.8 mm, MD: 1.2 mm, Lăpugiu de Sus (Romania), Figs 23 C 1 – C 2. Type locality. Lăpugiu de Sus (Romania). Type stratum. Dej Formation. Age. Middle Miocene, middle Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF8F675CF2BD014FFE28FDCF.taxon	etymology	Etymology. Referring to the small size and the occurrence in Romania.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF8F675CF2BD014FFE28FDCF.taxon	diagnosis	Diagnosis. Small, slender, elongate pyriform shell with narrowly truncated, umbilicate apex and nearly straight sides and moderately constricted, convex base; surface smooth; indistinct columellar fold.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF8F675CF2BD014FFE28FDCF.taxon	description	Description. Shell small, slender (SL / MD = 2.3 – 2.5), elongate pyriform, only last whorl visible. Last whorl with almost straight to weakly convex sides, forming angle of about 15 – 17 °. Periphery below mid-whorl at about one-third whorl height (SL / PH = 2.6 – 2.9), rounded at moderately contracting base. Apex narrow (MD / ApD = 1.8 – 2.0), truncated, apical plane slightly oblique (AP = 12 – 13 °), deeply and narrowly umbilicate, delimited by round-edged, elevated crest. Sculpture restricted to faint growth lines at apex. Aperture longer than shell, very narrow down to just above mid-whorl (MD / AW = 9.4 – 12.1), widening anteriorly below periphery (MD / MAW = 1.8 – 2.1), resulting in high expansion rate (ER = 4.5 – 5.2). Outer lip thin, slightly alate apically, extending some distance above apex. Columella short, with narrow columellar callus with indistinct columellar fold, almost obsolete. Columellar callus delimiting narrow umbilical chink. Parietal callus not developed.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF8F675CF2BD014FFE28FDCF.taxon	discussion	Discussion. Pyrunculus microromanicus sp. nov. is reminiscent of Pyrunculus leuchsi (Berger, 1949) in general shape. It differs in its slightly smaller size and slenderer outline; SL / MD: 2.3 – 2.5 and AP: 15 – 16 ° in P. microromanicus versus SL / MD: 2.2 and AP: 18 – 20 ° in P. leuchsi). In addition, the slightly conical apical region of P. leuchsi is separated from the sides by a faint angulation, which is missing in P. microromanicus sp. nov. Paleoenvironment. Unknown.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF8F675CF2BD014FFE28FDCF.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Făget Basin: Coşteiu de Sus, Lăpugiu de Sus (Romania) (Boettger 1907).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF91675EF2BD0246FED3FCEF.taxon	description	Figs 24 A – F	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF91675EF2BD0246FED3FCEF.taxon	materials_examined	Type material. Lectotype (designated by Zilch 1934: 277), SMF 363161 - 12.2551 a (glass plate 399), SL: 4.3 mm, MD: 2.0 mm, Coşteiu de Sus (Romania), illustrated in Zilch (1934: pl. 22, fig. 18). The type was destroyed during WWII, Figs 24 F. Illustrated material. NHMW 1855 / 0045 / 0258, SL: 4.9 mm, MD: 1.9 mm, Steinebrunn (Austria), Figs 24 A 1 – A 2. NHMW 1855 / 0045 / 0258 a, SL: 4.8 mm, MD: 1.8 mm, Steinebrunn (Austria), Figs 24 B 1 – B 3. NHMW 1855 / 0045 / 0258 b, SL: 4.5 mm, MD: 2.1 mm, Steinebrunn (Austria), Figs 24 C 1 – C 2. NHMW-ZOO-MO- 75000 - E- 42592, SL: 3.2 mm, MD: 1.4 mm, Bad Vöslau (Austria), Figs 24 D 1 – D 3. RGM 776934, SL: 3.5 m, MD: mm, Seyithasan (Turkey), Serravallian), illustrated in Landau et al. (2013: pl. 53, fig. 10), Figs 24 E 1 – E 2.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF91675EF2BD0246FED3FCEF.taxon	description	Revised description. Shell small, medium-thickness, moderately slender (SL / MD = 2.7 – 2.9), moderately elongate pyriform, only last whorl visible. Last whorl with only weakly convex sides, forming angle of about 21 – 23 °. Periphery below mid-whorl at about one-third whorl height (SL / PH = 2.7 – 2.9), rounded at rapidly contracting base. Apex narrow (MD / ApD = 2.5 – 3.0), truncated, apical plane strongly oblique (AP = 21 – 23 °), narrowly umbilicate, delimited by round-edged elevated crest. Sculpture of a few, faint spiral grooves on base. Aperture longer than shell, very narrow down to periphery (MD / AW = 9.6 – 10.8), abruptly widening anteriorly below periphery (MD / MAW = 1.9 – 2.1), resulting in high expansion rate (ER = 5.0 – 5.4). Outer lip thin, apically, extending only slightly above apex. Columella short, with narrow but prominent columellar callus with faint, low, broad columellar fold. Columellar callus delimiting elongated umbilical chink. Parietal callus not developed.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF91675EF2BD0246FED3FCEF.taxon	discussion	Discussion. Pyrunculus parangistoma (Boettger, 1902) is characterized by its relatively stout conical outline. Pyrunculus elongatus (Eichwald, 1830) differs in its slenderer shape, less contracted base and higher position of the periphery. Pyrunculus szobiensis sp. nov. differs in its less conical, more subcylindrical last whorl and the conical apical part of the shell. Paleoenvironment. Coastal marine, inner neritic; the frequent occurrence at Steinebrunn suggest a preference for seagrass environments (own data).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF91675EF2BD0246FED3FCEF.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienna Basin: Bad Vöslau, Steinebrunn (Austria) (hoc opus); Făget Basin: Coşteiu de Sus, Lăpugiu de Sus (Romania) (Zilch 1934); Dacian Basin: Opanets (Bulgaria) (Kojumdgieva & Strachimirov 1960). Proto-Mediterranean Sea. Serravallian (Middle Miocene): Karaman Basin: Seyithasan (Turkey) (Landau et al. 2013).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF936759F2BD049BFCCCFEB3.taxon	description	Figs 25 A – D	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF936759F2BD049BFCCCFEB3.taxon	materials_examined	Type material. Holotype, NHMW 186100010507, SL: 4.4 mm, MD: 2.1 mm, Szob (Hungary), Figs 25 A 1 – A 3. Paratypes, NHMW 186100010507 a, SL: 3.9 mm, MD: 1.8 mm, Szob (Hungary), Figs 25 B 1 – B 2. NHMW 186100010507 b, SL: 4.4 mm, MD: 1.9 mm, Szob (Hungary), Figs 25 C 1 – C 2. NHMW-ZOO-MO- 75000 - E- 42592, SL: 3.3 mm, MD: 1.5 mm, Bad Vöslau (Austria), Figs 25 D 1 – D 3. Type locality. Szob at the SW slopes of the Börzsöny Mountains. Type stratum. Yellowish, grey sand (Hungary) (Dulai 1996). Age. Middle Miocene, eMiddle Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF936759F2BD049BFCCCFEB3.taxon	diagnosis	Diagnosis. Small, moderately slender, bullet-shaped shell with conical apical region passing via indistinct angulation into almost straight, subcylindrical sides. Umbilicate apex. Weak sculpture of indistinct spiral grooves at both extremities. Narrow aperture in adapical half moderately widens in abapical part. No columellar fold.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF936759F2BD049BFCCCFEB3.taxon	description	Description. Shell small, moderately slender (SL / MD = 2.3 – 2.4), bullet-shaped, only last whorl visible. Last whorl with short conical apical region (43 – 49 °) passing into almost straight sides, forming angle of about 8 – 9 °. Periphery placed below mid-whorl between half and one-third height (SL / PH = 3.0 – 3.6), rounded at moderately contracting base. Apex narrow to moderately narrow (MD / ApD = 2.3 – 2.7), truncated, apical plane slightly oblique (AP = 8 – 9 °), deeply and narrowly umbilicate, delimited by round-edged, faintly swollen, elevated crest. Sculpture of a few, shallow spiral grooves at both extremities. Aperture longer than shell, very narrow down to periphery (MD / AW = 8.0 – 11.9), widening anteriorly below periphery (MD / MAW = 1.8 – 2.0), resulting in high expansion rate (ER = 4.4 – 6.0). Outer lip thin, apically extending some distance above apex, weakly concave just above periphery. Columella short, with narrow columellar callus without columellar fold. Columellar callus adherent or delimiting narrow umbilical chink. Parietal callus not developed.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF936759F2BD049BFCCCFEB3.taxon	discussion	Discussion. This species was confused with Pyrunculus parangistoma (Boettger, 1902) by Berger (1953) and Strausz (1962, 1966) but differs in its bullet-shaped outline and lacks a columellar fold. Pyrunculus leuchsi (Berger, 1949) is smaller, broader and has more conical sides of the last whorl. At a smaller size P. leuchsi has a weakly developed columellar fold whereas Pyrunculus szobensis sp. nov. has no columellar fold developed Paleoenvironment. Inner neritic, shallow marine with seagrass meadows (Dulai 1996).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF936759F2BD049BFCCCFEB3.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienna Basin: Bad Vöslau (Austria) (hoc opus); Pannonian Basin: Szob (Hungary) (Strausz 1966).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF94675BF2BD0196FCB1FF03.taxon	description	Figs 25 E – F, 26 A – B	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF94675BF2BD0196FCB1FF03.taxon	materials_examined	Type material. Holotype, NHMW-ZOO-MO- 75000 - E- 42898 A, SL: 3.7 mm, MD: 1.6 mm, Vienna / Ottakring (Austria), Figs 25 E, 26 A 1 – A 2. Paratype, NHMW 1859 / 0045 / 0604, SL: 2.8 mm, MD: 1.4 mm, Lăpugiu de Sus (Romania), Figs 25 F, 26 B 1 – B 2. Type locality. Vienna / Ottakring (Austria). Type stratum. Baden Formation. Age. Middle Miocene, middle Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF94675BF2BD0196FCB1FF03.taxon	etymology	Etymology. In honor of Martin Zuschin (Institute for Paleontology, University Vienna), in recognition of his contributions to Conservation Paleobiology.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF94675BF2BD0196FCB1FF03.taxon	diagnosis	Diagnosis. Small, moderately slender, moderately elongate pyriform shell without distinct concavity in adapical half, narrow crest at umbilicate apex characterized by prominent growth lines crossed by weak spiral grooves, resulting in vaguely reticulated pattern; weak columellar fold.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF94675BF2BD0196FCB1FF03.taxon	description	Description. Shell small, moderately slender (SL / MD = 2.1 – 2.3), moderately elongate pyriform, only last whorl visible. Last whorl with nearly straight sides, forming angle of about 24 °. Periphery below mid-whorl at about one-third whorl height (SL / PH = 2.5 – 2.6), rounded at moderately constricted base. Apex narrow (MD / ApD = 1.9 – 2.0), truncated, apical plane moderately oblique (AP = 14 °), deeply and narrowly umbilicate, delimited by round-edged elevated crest. Sculpture of prominent, widely spaced growth lines and weakly rugose spiral cords separated by shallow, widely spaced spiral grooves resulting in vaguely reticulate pattern. Spiral grooves most prominent on base. Aperture longer than shell, very narrow down just above periphery (MD / AW = 12.1 – 12.2), abruptly widening anteriorly below periphery (MD / MAW = 2.0 – 12.2), resulting in high expansion rate (ER = 6.0 – 6.4). Outer lip thin, apically, extending only slightly above apex. Columella short, with narrow columellar callus with faint, low, broad columellar fold. Columellar callus delimiting narrow umbilical chink. No parietal callus.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF94675BF2BD0196FCB1FF03.taxon	discussion	Discussion. The Paratethyan species is reminiscent of the extant Pyrunculus hoernesi (Weinkauff, 1866) from the Mediterranean Sea but lacks the characteristic axial folds or ribs at the apical crest and has a much narrower umbilicus. Pyrunculus zuschini sp. nov. is intermediate in shape between the even more elongate Pyrunculus elongatus (Eichwald, 1830) and the stouter Pyrunculus parangistoma (Boettger, 1902) and differs from both species in its prominent sculpture. Indeed, it is the only axially and spirally sculptured Pyrunculus species in the Paratethyan assemblages. Its sculpture is somewhat reminiscent of the Upper Miocene to Pliocene Mediterranean Retusa decussata Sacco, 1897, which usually has even stronger reticulated surface sculpture (see Landau et al. 2023: pl. 33), but that species has a broadly and flatly truncated apex and a more cylindrical shell, characters of the genus Retusa Brown, 1827. Paleoenvironment. The occurrence in the Baden Formation at Bad Vöslau (Austria) suggests middle to outer neritic environments with up to 210 m water depth (Kranner et al. 2021).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF94675BF2BD0196FCB1FF03.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienna Basin: Vienna / Ottakring (Austria) (hoc opus); Făget Basin: Lăpugiu de Sus (Romania) (hoc opus).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF96675BF2BD0637FE20FE7B.taxon	type_taxon	Type species. Bulla obtusa Montagu, 1803; Opinion 568 (IZCN 1959), subsequent designation by Iredale (1915: 300). Present-day, British Isles.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF96675BF2BD06E3FB53F821.taxon	description	Figs 27 A – E	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF96675BF2BD06E3FB53F821.taxon	materials_examined	Type material. Holotype, NHMW 1855 / 0043 / 0086, SL: 3.0 mm, MD: 1.7 mm, Lăpugiu de Sus (Romania), Figs 27 D 1 – D 3. Paratypes, NHMW 1855 / 0045 / 0260 c, SL: 2.4 mm, MD: 1.3 mm, Steinebrunn (Austria), Figs 27 A 1 – A 3. NHMW 1973 / 1615 / 0124, SL: 2.5 mm, MD: 1.3 mm, Lăpugiu de Sus (Romania), Figs 27 B 1 – B 3. NHMW 1855 / 0045 / 0260 d, SL: 1.9 mm, MD: 1.1 mm, Steinebrunn (Austria), Figs 27 C 1 – C 3. NHMW 1997 z 0178 / 1212, SL: 2.8 mm, MD: 1.4 mm, Bad Vöslau (Austria), Figs 27 E 1 – E 2. Type locality. Lăpugiu de Sus (Romania), Făget Basin. Type stratum. Silt and clay of the Dej Formation. Age. Middle Miocene, middle Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF96675BF2BD06E3FB53F821.taxon	etymology	Etymology. Latin mysticus (= mysterious), referring to the mysterious status of this species in the literature.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF96675BF2BD06E3FB53F821.taxon	diagnosis	Diagnosis. Small, solid, moderately broad, subcylindrical shell with rounded shoulder angulation, truncated apex with wide umbilicus showing early spire whorls; weak, arcuate growth lines, weak, adherent columellar callus without fold.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF96675BF2BD06E3FB53F821.taxon	description	Description. Shell small, solid, moderately broad, subcylindrical (SL / MD = 1.8 – 1.9), of about three teleoconch whorls. Protoconch unknown. Periphery at one-third whorl height (SL / PH = 2.0 – 2.3), almost straight to faintly concave sides above periphery, forming angle of about 14 – 33 °. Rounded at rapidly contracting base. Apex truncated, with rounded shoulder angulation, moderately sunken, broad (MD / ApD = 1.7 – 2.0), apical plane faintly oblique (AP = 0 – 3 °), deeply and widely umbilicate, showing convex spire whorls. Spire whorls and apical crest convex. Surface with weak to subobsolete arcuate growth lines; usually more prominent on apical crest and spire whorls. Aperture entire length of shell, narrow down to mid-whorl (MD / AW = 8.5 – 10.2), distinctly widening anteriorly (MD / MAW = 2.2 – 2.4), resulting in moderate expansion rate (ER = 3.6 – 4.2). Outer lip thin, very slightly extending above apex. Columella short, nearly straight without columellar fold. Columellar callus forming narrow, adherent rim. No umbilicus or umbilical chink. Parietal callus weak.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF96675BF2BD06E3FB53F821.taxon	discussion	Discussion. Retusa mystica sp. nov. was confused so far with the Miocene to Recent Retusa truncatula (Bruguière, 1792). Although Retusa truncatula is highly variable, it differs from Retusa mystica in its generally slenderer shell, often with a pinched-in section above the periphery, and more prominent axial ribs. Its apical crest typically bears relatively prominent axial folds or ribs. Retusa mystica sp. nov. could easily be mistaken for a weakly sculptured Retusa sarmatica Berger, 1953 but differs in its much wider apical umbilicus (compare Figs 27 A 3 – D 3 versus 29 A 3 – C 3). Paleoenvironment. The occurrence at Steinebrunn (Austria) suggests shallow marine environments with seagrass (own data M. H.). Nearshore environments are also reflected by Várpalota (Hungary) and Kuzmice (Slovakia) (own data M. H.).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF96675BF2BD06E3FB53F821.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienna Basin: Bernhardsthal (Austria) (Harzhauser et al. 2018 a); Eastern Slovakian Basin: Kuzmice (Slovakia) (Švagrovský 1964); Pannonian Basin: Várpalota (Hungary) (Strausz 1954); Făget Basin: Lăpugiu de Sus (Romania) (hoc opus).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF976754F2BD0185FF37FC73.taxon	description	Figs 28 A – F	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF976754F2BD0185FF37FC73.taxon	materials_examined	Type material. Lectotype, SMF 363158 - 12.246 a (glass plate 396), SL: 5.6 mm, MD: 1.6 mm, Coșteiu de Sus (Romania), illustrated in Zilch (1934: pl. 22, fig. 15), Fig 28 C. The specimen was destroyed during WW II. Illustrated material. NHMW-ZOO-MO- 75000 - E- 47025, SL: 4.1 mm, MD: 2.1 mm, Guntersdorf (Austria), Figs 28 A 1 – A 3; NHMW-ZOO-MO- 75000 - E- 47026, SL: 3.5 mm, MD: 1.9 mm, Grund (Austria), Figs 28 B 1 – B 3. NHMW 2018 / 0248 / 0477, SL: 2.3 mm, MD: 1.2 mm, Coşteiu de Sus (Romania), Figs 28 D 1 – D 2. NHMW 1864 / 0001 / 0224, SL: 2.3 mm, MD: 1.2 mm, Gainfarn (Austria), Figs 28 E 1 – E 3. NHMW 1864 / 0001 / 0224 a, SL: 3.5 mm, MD: 1.8 mm, Gainfarn (Austria), Figs 28 F 1 – F 3. Revised description. Shell small, solid, moderately broad, subcylindrical (SL / MD = 1.8 – 2.0), with almost straight to faintly concave sides, forming angle of 9 – 10 °. Periphery just below mid-whorl (SL / PH = 2.3). Protoconch mammillate, heterostrophic with only last whorl visible. Teleoconch of about three whorls separated by deeply incised suture. Spire whorls occasionally partly covered by subsequent whorls, greatly depressed, resulting in almost flat, broad spire (MS / ApD = 1.6 – 1.7; apical angle 150 – 170 °), angled at shoulder. Last whorl ~ 97 % of total height, with narrow, flat subsutural platform, cylindrical below, widening slightly abapically, rounded at base. Surface smooth except for shallowly arcuate growth lines. Aperture narrow in upper half (MD / AW = 7.8 – 9.8), widening distinctly anteriorly (MD / MAW = 1.9 – 2.2), resulting in moderate to high expansion rate (ER = 3.6 – 4.7), outer lip simple, moderately thin. Columella moderately excavated, narrowly expanded over base, without fold. Columellar callus narrow, usually adherent, occasionally slightly erect, bordering small umbilical chink (Fig. 28 E 1). No parietal callus.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF976754F2BD0185FF37FC73.taxon	discussion	Discussion. The Pliocene to Recent Mediterranean Retusa mammillata (Philippi, 1836) differs in its strongly protruding protoconch, which exposes also the first whorl, the slenderer outline and the more prominent axial sculpture. Retusa promamillata Boettger, 1907 is also reminiscent of the Miocene to Recent Mediterranean Retusa truncatula (Bruguière, 1792), which differs in its more prominent axial sculpture and the sunken spire. Retusa voeslauensis (Berger, 1953) shows close similarities with R. promamillata in apical view and has an identical protoconch but differs clearly in its elevated spire and more rounded shoulder. Paleoenvironment. The occurrence at Gainfarn (Austria) suggests shallow marine conditions with seagrass (Zuschin et al. 2007).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF976754F2BD0185FF37FC73.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): North Alpine-Carpathian Foreland Basin: Grund, Guntersdorf (Austria) (Berger 1953); Vienna Basin: Bad Vöslau, Gainfarn (Austria) (Berger 1953); Danube Basin: Modra-Kráľová (Slovakia) (Hladilová & Fordinál 2013); Pannonian Basin: Budapest (Hungary) (Kókay 1996); Făget Basin: Coşteiu de Sus (Romania) (Boettger 1907); Dacian Basin: Islaz (Romania) (Rado & Muţui 1970).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF996756F2BD04E6FB44FEB3.taxon	description	Figs 29 A – D	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF996756F2BD04E6FB44FEB3.taxon	materials_examined	Type material. Holotype, NHMW-ZOO-MO- 75000 - E- 47021 A, SL: 1.9 mm, MD: 1.1 mm, Vienna / Hernals (Austria), Sarmatian, illustrated in Berger (1953: pl. 16, fig. 20), Figs 29 C 1 – C 2. Illustrated material NHMW 1855 / 0045 / 0260, SL: 1.9 mm, MD: 1.0 mm, Steinebrunn (Austria), Figs 29 A 1 – A 3. NHMW 1870 / 0037 / 0063, SL: 1.8 mm, MD: 0.9 mm, Ritzing (Austria), Figs 29 B 1 – B 3. NHMW-ZOO-MO- 75000 - E- 47018, SL: 2.4 mm, MD: 1.3 mm, Hölles (Austria), identified as Retusa truncatula jekelii by Berger 1949, Figs 29 D. Revised description. Shell small, solid, moderately broad, subcylindrical (SL / MD = 1.9 – 2.2), only last whole visible. Protoconch unknown. Periphery at or just below mid-whorl (SL / PH = 1.8), almost straight to faintly convex sides above periphery, forming angle of about 11 – 14 °. Rounded at rapidly contracting base. Apex truncated, moderately broad (MD / ApD = 1.7 – 2.0), apical plane faintly oblique (AP = 7 – 9 °), deeply and narrowly umbilicate, delimited by convex apical crest. Surface with prominent, arcuate growth lines; most prominent on apical crest, sigmoidal in apical view. Aperture entire length of shell, narrow down to mid-whorl (MD / AW = 7.6 – 8.3), distinctly widening anteriorly (MD / MAW = 2.0 – 2.2), resulting in moderate to high expansion rate (ER = 3.5 – 4.0). Outer lip thin, not extending above apex. Columella short, slightly oblique, without columellar fold. Columellar callus forming very narrow rim, delimiting very narrow umbilical chink. Parietal callus not developed.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF996756F2BD04E6FB44FEB3.taxon	discussion	Discussion. Retusa sarmatica Berger, 1953 is almost identical to Retusa mystica sp. nov. in outline but differs in its narrowly umbilicate apex. Retusa sarmatica is reminiscent of the extant Mediterranean Retusa minutissima (Monterosato, 1878), which has similar sculpture but differs in its subquadratic outline and the open spire. Paleoenvironment. The occurrences at Steinebrunn, Ritzing (Austria) and Borský Mikuláš (Slovakia) suggest shallow marine environments (own data M. H.).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF996756F2BD04E6FB44FEB3.taxon	distribution	Distribution in Central Paratethys. Despite its name, this species is not restricted to the Sarmatian but appears already during the Badenian. Badenian (Middle Miocene): Vienna Basin: Steinebrunn (Austria) (hoc opus); Oberpullendorf Basin: Ritzing (Austria) (hoc opus). Sarmatian (Middle Miocene): Vienna Basin: Hölles, Vienna / Hernals (Austria) (Berger 1953); Eisenstadt-Sopron Basin: Wiesen (Austria) (Berger 1953); Eastern Slovakian Basin: Nižná Myšľa, Nižný Čaj, Skaroš, Slanec, Slančík, Sečovce, Trstené pri Hornáde (Slovakia) (Švagrovský 1971). Poiana Ruscă Mountains: Răcăştia (Romania) (Tămaș et al. 2013); Mehadia Basin: Valea Tudoroviţa (Romania) (Hinculov in Iliescu et al. 1968).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF9B6751F2BD06A6FA5BFF2F.taxon	description	Figs 29 E – F	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF9B6751F2BD06A6FA5BFF2F.taxon	materials_examined	Type material. Lectotype (designated herein), specimen illustrated in Kolesnikov (1935: pl. 33, fig. 20), Tishchenskoe (Stavropol region, Russia), SL: 7.2 mm, MD: 2.0 mm, Bessarabian (Late Miocene), probably stored in TSNIGR Museum Saint-Petersburg. Illustrated material. NHMW-ZOO-MO- 75000 - E- 47058, SL: 7.9 mm, MD: 3.6 mm, Bílovice u Uherského Hradiště (Czech Republic), Figs 29 E 1 – E 3. NHMW-ZOO-MO- 75000 - E- 47058, SL: 7.4 mm, MD: 3.5 mm, Bílovice u Uherského Hradiště (Czech Republic), Figs 29 F 1 – F 2.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF9B6751F2BD06A6FA5BFF2F.taxon	description	Revised description. Shell medium sized, solid, moderately slender, elongate cylindrical with faintly convex to concave sides (SL / MD = 2.1 – 2.3). Protoconch tilted, slightly sunken, largely covered by teleoconch. Teleoconch of about three whorls separated by canaliculated suture. Spire truncated, nearly flat, moderately broad (MD / ApD = 1.7 – 1.8). Early teleoconch whorls low, only weakly convex, with periphery at abapical suture. Last whorl 100 % of total height, with weak, round shoulder and rapidly contracting, wide base. Periphery placed around mid-whorl (SL / PH = 1.8 – 1.9). Surface with delicate, shallowly arcuate growth lines. Aperture 92 – 95 % of total height, moderately narrow in upper half (MD / AW = 6.0), strongly widening anteriorly (MD / MAW = 1.6 – 1.8), resulting in moderate expansion rate (ER = 3.3 – 3.6). Outer lip thin, shallowly arcuate in profile, straight to weakly concave around mid-whorl, wide and rounded at base. Columella short, strongly excavated, strongly thickened, forming broad rim without columellar fold. Columellar callus adherent to base without umbilical chink, persisting continuously into slightly weaker parietal callus.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF9B6751F2BD06A6FA5BFF2F.taxon	discussion	Discussion. We have not studied the type specimens and have no material from the type area at hand. Therefore, the identification of the Central Paratethyan specimens is tentative. Retusa sinzovi (Kolesnikov, 1935) was treated in the literature as subspecies of Acteocina volhynica (Eichwald, 1830) [erroneously listed as Acteocina lajonkaireana (de Basterot, 1825)]. Retusa sinzovi differs from Acteocina volhynica in its much larger size, truncated apex, sunken protoconch and elongate cylindrical shape. The sunken protoconch and truncated apex place this species in Retusa. Paleoenvironment. Inner neritic, shallow marine based on associated fauna (own data, M. H.).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF9B6751F2BD06A6FA5BFF2F.taxon	distribution	Distribution in Central Paratethys. Sarmatian (Middle Miocene): Vienna Basin: Čejč, Bílovice u Uherského Hradiště, Holič, Strážnice, Rohožník (Slovakia) (Švagrovský 1971); Eisenstadt-Sopron Basin: Wiesen (Austria) (Papp 1954); Pannonian Basin: Várpalota (Hungary) (Boda 1959); Eastern Slovakian Basin: Bohdanovce, Nižná Myšľa, Nižný Čaj, Slanec, Skaroš (Švagrovský 1971); Transylvanian Basin: Soceni (Romania) (Jekelius 1944). Eastern Paratethys. Volhynian (Middle Miocene): Ciscaucasia (Russia) (Kolesnikov 1935). Bessarabian (Late Miocene): Moldavian Platform: Chişinău (Moldova) (Simionescu & Barbu 1840); Kerch Peninsula (Russia), Ciscaucasia: Tishchenskoe (Stavropol region, Russia), Azov Sea region: Melitopol (Ukraine) (Kolesnikov 1935).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF9C6753F2BD065AFF26FBC7.taxon	description	Figs 30 A – E	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF9C6753F2BD065AFF26FBC7.taxon	description	Retusa truncatula (Bruguière, 1792) — Ionesi & Ionesi 1972: pl. 4, figs 6 – 10.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF9C6753F2BD065AFF26FBC7.taxon	materials_examined	Type material. We are not aware of the whereabouts of the type material. Illustrated material. NHMW 2018 / 0248 / 0476, SL: 2.9 mm, MD: 1.4 mm, Coșteiu de Sus (Romania), Figs 30 A 1 – A 3. NHMW-ZOO-MO- 75000 - E- 41686, SL: 3.9 mm, MD: 1.8 mm, Steinebrunn (Austria), illustrated as Retusa subtruncatula in Berger (1953: pl. 16, fig. 15), Figs 30 B 1 – B 2. NHMW-ZOO-MO- 75000 - E- 47002, SL: 3.1 mm, MD: 1.5 mm, Grund (Austria), Badenian, holotype of Retusa grundensis Berger, 1953, Figs 30 C 1 – C 3. NHMW 2025 / 0152 / 0001, SL: 1.8 mm, MD: 0.9 mm, Bad Vöslau (Austria), Figs 30 D 1 – D 3. NHMW 2025 / 0152 / 0002, SL: 1.3 mm, MD: 1.1 mm, Bad Vöslau (Austria), Figs 30 E 1 – E 3.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF9C6753F2BD065AFF26FBC7.taxon	description	Revised description. See Landau et al. (2023: 384). Paratethyan synonyms. Retusa truncatula pappi Berger, 1953. Holotype, NHMW-ZOO-MO- 75000 - E- 47018, Hölles (Austria), illustrated in Berger (1953: pl. 16, fig. 19, the specimen is lost. Retusa truncatula pappi was introduced by Berger (1953) for specimens with slight concavity below the shoulder, which does not justify a separation from Retusa truncatula. Specimens treated by Papp (1954) as Retusa truncatula pappi support this synonymization. For a list of synonyms based on extant specimens see MolluscaBase eds. (2025 a).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF9C6753F2BD065AFF26FBC7.taxon	discussion	Discussion. For a discussion on the variability of this species in the Paratethys Sea see Guzhov (2022 b: 1239). Retusa truncatula (Bruguière, 1792) is extremely similar to Retusa mystica sp. nov. but differs in usually being more cylindrical and having a pinched region just above the periphery. Moreover, when seen in apical view, the axial folds or riblets are present at the rounded shoulder that are not, or hardly, present in R. mystica sp. nov. Paleoenvironment. This species lives in sandy and muddy habitats up to 50 m depth (Gofas et al. 2011).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF9C6753F2BD065AFF26FBC7.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Roztocze Hills: Łychów (Poland) (Krach 1981); Korytnica Basin: Korytnica (Poland) (Friedberg 1938); Nowy Sącz Basin: Niskowa (Poland) (Bałuk 1970); Voronyaky Hills: Oles’ko, Pidgirtsi, Yaseniv (Ukraine) (Friedberg 1938); Polish-Ukrainian Fore-Carpathian Basin: Wieliczka (Poland) (Reuss 1867); Borki Wielkie (Poland); Tarnoruda, Staryi Pochaiv, Zaleśce, Zhabyak (Ukraine) (Friedberg 1938); Korytnica Basin: Korytnica (Poland) (Friedberg 1938); Nowy Sącz Basin: Niskowa (Poland) (Bałuk 1970); North Alpine-Carpathian Foreland Basin: Grund, Guntersdorf (Austria) (Berger 1953): Vienna Basin: Baden, Bernhardsthal, Steinebrunn (Austria) (Berger 1953; Harzhauser et al. 2018 a), Borský Mikuláš (Slovakia) (Švagrovský 1984); Pannonian Basin: Várpalota (Hungary) (Strausz 1966); Făget Basin: Coşteiu de Sus (Romania) (Boettger 1907); Eastern Slovakian Basin: Kuzmice (Slovakia) (Švagrovský 1960); Dacian Basin: Islaz (Romania) (Rado & Muţui 1970). Sarmatian (Middle Miocene): Polish-Ukrainian Fore-Carpathian Basin: Krzemieniec Żolobki (Poland); Rydomyl’, Shushkivtsi, Zalistsi (Ukraine) (Friedberg 1938); Vienna Basin: Hölles, Nexing (Austria) (Berger 1953; hoc opus), Vienna / Theresiengasse (Austria) (Harzhauser et al. 2018 b); Holič, Podivín, Rohožník (Slovakia) (Švagrovský 1971); Eisenstadt-Sopron Basin: Wiesen (Austria) (Berger 1953); Pannonian Basin: Sopron, Pécsvárad, Karád (Hungary) (Boda 1959); Eastern Slovakian Basin: Slanec, Trstené pri Hornáde, Nižná Myšľa, Slančík, Bidovce, Sečovce (Slovakia) (Švagrovský 1971); Transylvanian Basin: Soceni (Romania) (Jekelius 1944); Poiana Ruscă Mountains: Răcăştia (Romania) (Tămaș et al. 2013). Eastern Paratethys. Widespread in the Volhynian and Bessarabian of Romania, Moldova and Russia (Guzhov 2022 b). Proto-Mediterranean Sea. Serravallian (Middle Miocene): Karaman Basin: Akpınar, Lale, Seyithasan (Turkey) (Landau et al. 2013). Tortonian (Late Miocene): Livorno Hills: Quarata (Italy) (Dominici et al. 2020). Tortonian and Messinian (late Miocene): Po Basin (Italy) (Sacco 1897; Venzo & Pelosio 1963; d’Amico et al. 2012). Pliocene to present-day. North Sea, northeastern Atlantic and Mediterranean Sea (see Landau et al. 2023 for details).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF9E6752F2BD0372FDF1F972.taxon	description	Figs 31 A – C	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF9E6752F2BD0372FDF1F972.taxon	materials_examined	Type material. Holotype, NHMW-ZOO-MO- 75000 - E- 3311 A, SL: 3.6 mm, MD: 1.7 mm, Bad Vöslau (Austria), illustrated in Berger (1953: pl. 16, fig. 22), Figs 31 A 1 – A 3. Berger (1953) mentioned further occurrences from Lăpugiu de Sus (Romania) and Guntersdorf (Austria). After revision we exclude these specimens because they are not conspecific with Retusa vindobonensis. Illustrated material. NHMW-ZOO-MO- 75000 - E- 7041, SL: 3.5 mm, MD: 3.1 mm, Grund (Austria), Figs 31 B 1 – B 3. NHMW-ZOO-MO- 75000 - E- 7041, SL: 4.5 mm, MD: 2.0 mm, Grund (Austria), Figs 31 C 1 – C 3.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF9E6752F2BD0372FDF1F972.taxon	description	Revised description. Shell small, solid, moderately slender (SL / MD = 2.1 – 2.3). Only last whorl visible. Protoconch unknown. Periphery just below mid-whorl (SL / PH = 2.2 – 2.6) with almost straight sides, weakly widening abapically; adapically forming angle of about 10 – 20 °. Rounded at slowly contracting base. Apex truncated, moderately narrow (MD / ApD = 2.1 – 2.3), apical plane faintly oblique (AP = 7 – 10 °), deeply and narrowly umbilicate, delimited by rounded crest with delicate, fold-like axial ribs on umbilical side. Sculpture of arcuate growth lines; most prominent on apical crest and extremities, subobsolete mid-whorl. Aperture only slightly longer than shell, narrow down to mid-whorl (MD / AW = 9.0 – 9.5), distinctly widening anteriorly (MD / MAW = 1.8 – 2.0), resulting in high expansion rate (ER = 3.3 – 4.7). Outer lip thin, moderately extending above apex. Columella short, nearly straight; columellar fold weak to subobsolete. Columellar callus forming very narrow, adherent rim delimiting faint umbilical chink. Parietal callus not developed.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF9E6752F2BD0372FDF1F972.taxon	discussion	Discussion. This rare species might be related to Retusa sarmatica Berger, 1953 due to its nearly identical apical morphology and prominent axial sculpture. It differs from R. sarmatica in its slenderer shape, more conical last whorl and narrower apex. Retusa subangystoma (d’Orbigny, 1852), from the Early to Late Miocene of France, differs in its more subcylindrical outline and less convex periphery (see Peyrot 1932: pl. 13, figs 34 – 35; Landau et al. 2020: pl. 15). Berger (1953) confused this species with Retusa pliosimplex (Sacco, 1897) from the Pliocene of Italy. The Italian species has a similar shape but is slightly more elongate and differs in its wider, cylindrical apical umbilicus. In addition, it has a wider abapical aperture (see Sacco 1897: 51, pl. 4, figs 15 – 18; Ferrero Mortara et al. 1984: pl. 52, figs 10 a – c; Chirli 2023: pl. 12, figs 1 – 6). Paleoenvironment. The occurrence in the Baden Formation at Bad Vöslau (Austria) suggests middle to outer neritic environments with up to 210 m water depth (Kranner et al. 2021).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF9E6752F2BD0372FDF1F972.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): North Alpine-Carpathian Foreland Basin: Grund, Guntersdorf (Austria) (Berger 1953); Vienna Basin: Bad Vöslau (Austria) (Berger 1953); Pannonian Basin: Budapest (Hungary) (Kókay 1996).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF9F672DF2BD01E7FE54F9B1.taxon	description	Figs 32 A – E	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF9F672DF2BD01E7FE54F9B1.taxon	description	R [etusa]. (R [etusa].) mammillaris regulbensis [sic] (Ad.) — Sieber 1958: 163 [non Bulla regulbiensis Kanmacher, 1798 = Retusa obtusa Montagu, 1803)].	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF9F672DF2BD01E7FE54F9B1.taxon	materials_examined	Type material. Holotype, NHMW-ZOO-MO- 75000 - E- 3318 A, SL: 3.7 mm, MD: 1.7 mm, Bad Vöslau (Austria), Figs 32 A 1 – A 3. Paratype, NHMW-ZOO-MO- 75000 - E- 3318, SL: 3.6 mm, MD: 1.7 mm, Bad Vöslau (Austria), Figs 32 B 1 – B 3. NHMW-ZOO-MO- 75000 - E- 3318, SL: 3.7 mm, MD: 1.7 mm, Bad Vöslau (Austria), Figs 32 C 1 – C 3. Illustrated material. NHMW-ZOO-MO- 75000 - E- 3318, SL: 4.3 mm, MD: 2.1 mm, Steinebrunn (Austria), Figs 32 D 1 – D 2. NHMW-ZOO-MO- 75000 - E- 3318, SL: 3.4 mm, MD: 1.6 mm, Steinebrunn (Austria), Figs 32 E 1 – E 3. Additional material. NHMW 1863 / 0015 / 0107, SL: 2.9 mm, MD: 1.3 mm, Buituri (Romania); NHMW 1863 / 0015 / 0107 a, SL: 2.9 mm, MD: 1.4 mm, Buituri (Romania); NHMW 1863 / 0015 / 0107 b, SL: 2.6 mm, MD: 1.3 mm, Buituri (Romania). Revised description. Shell small, solid, moderately slender, subcylindrical (SL / MD = 2.1 – 2.6), with almost straight to faintly concave sides, forming low angle of 7 – 8 °. Periphery below mid-whorl (SL / PH = 2.3 – 2.6). Protoconch mammillate, heterostrophic with one or two visible whorls. Teleoconch of about three whorls separated by deeply incised suture. Spire whorls occasionally partly covered by subsequent whorls, convex with rounded shoulder, of variable height, resulting in more or less elevated spire (apical angle = 80 – 126 °); spire broad (MS / ApD = 1.6 – 1.7). Last whorl ~ 87 – 93 % of total height, with rounded shoulder angulation, subcylindrical below, widening slightly abapically, rounded at base. Surface smooth except for shallowly arcuate growth lines. Aperture narrow in upper half (MD / AW = 8.2 – 8.7), widening distinctly anteriorly (MD / MAW = 2.1), resulting in moderate to high expansion rate (ER = 3.8 – 4.1); outer lip simple, moderately thin. Columella moderately excavated, narrowly expanded over base, without fold. No parietal callus.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF9F672DF2BD01E7FE54F9B1.taxon	discussion	Discussion. This species was placed by Landau et al. (2013) in Acteocina but lacks the characteristic canaliculate suture of that genus. The spire morphology is identical with that of Retusa promamillata Boettger, 1907 and Retusa mammillata (Philippi, 1836), suggesting a close relation with these species. It differs from both of these species in its more elevated spire. Paleoenvironment. The occurrence at Steinebrunn (Austria) and at Seyithasan (Turkey) suggests shallow water environments (own data M. H, B. L) whereas the occurrence in the Baden Formation at Bad Vöslau (Austria) might also indicate deeper water environments (Kranner et al. 2021).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF9F672DF2BD01E7FE54F9B1.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Roztocze Hills: Łychów (Poland) (Krach 1981); Vienna Basin: Bad Vöslau, Vienna Pötzleinsdorf (Austria) (Berger 1953); Pannonian Basin: Pécsvárad (Hungary) (Kókay 1985); Bükk Mountains: Borsodbóta (Hungary) (Csepreghy-Meznerics 1972). Southern Pannonian Basin: Hrvaćani (Bosnia and Herzegovina) (Atanacković 1985); Făget Basin: Buituri (Romania) (hoc opus). Proto-Mediterranean Sea. Serravallian (Middle Miocene): Karaman Basin: Lale, Seyithasan (Turkey) (Landau et al. 2013).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFE0672DF2BD0010FF10F851.taxon	type_taxon	Type species. Bulla acuminata Bruguière, 1792; by typification of replaced name. Present-day, Mediterranean Sea.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFE1672FF2BD01CEFB89F845.taxon	description	Figs 33 A – C	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFE1672FF2BD01CEFB89F845.taxon	materials_examined	Illustrated material. NHMW 2010 / 0004 / 1651, SL: 4.0 mm, MD: 1.5 mm, Bad Vöslau (Austria), Figs 33 A 1 – A 2. NHMW 1876 / 0011 / 0171, SL: 2.9 mm, MD: 1.1 mm, Lăpugiu de Sus (Romania), Figs 33 B 1 – B 2. NHMW 2010 / 0004 / 1652, SL: 3.3 mm, MD: 1.4 mm, Bad Vöslau (Austria), Figs 33 C 1 – C 2.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFE1672FF2BD01CEFB89F845.taxon	description	Revised description. See Landau et al. (2023: 387).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFE1672FF2BD01CEFB89F845.taxon	discussion	Discussion. As discussed by Landau et al. (2013: 340), Volvulella acuminata (Bruguière, 1792) is either a species complex or an unusually long-lived and widely distributed species. We note that Paratethyan specimens at hand have the periphery above mid-whorl (SL / PH = 1.7) with slightly contracting sides below the periphery, whereas typical shells of V. acuminata tend to be roughly subcylindrical (e. g., Chirli 2013: pl. 12, figs 18 – 23; Landau et al. 2023: pl. 36, figs 1 – 3). The Paratethyan morph is reminiscent of the extant Volvulella fortis (Thiele, 1925) from New Caledonia (See Valdés 2008: fig. 81 C). Specimens described by Csepreghy-Meznerics (1952) from Szob (Hungary) correspond to the subcylindrical morph typical for V. acuminata. Tentatively, we treat slender and stout morphs as conspecific. Paleoenvironment. Volvulella acuminata is found circalittoral muddy sands (Gofas et al. 2011). Paratethyan occurrences derive from inner neritic and middle neritic settings. Many extant Volvulella species are documented from deep water habitats (Valdés 2008).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFE1672FF2BD01CEFB89F845.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienna Basin: Bad Vöslau (Austria) (Berger 1953); Pannonian Basin: Szob (Hungary) (Strausz 1966). Făget Basin: Coşteiu de Sus, Lăpugiu de Sus (Romania) (Boettger 1907; hoc opus). Distribution. The status of the various Early Miocene to Pleistocene records in the Mediterranean region, the north-eastern Atlantic and North Sea might need revision. For details on the distribution see Landau et al. (2013, 2020, 2023). Volvulella acuminata occurs in the Atlantic from Norway (Thompson 1988) to Angola (Nicklès 1950) and in the entire Mediterranean Sea (Cossignani & Ardovini, 2011; Gofas et al. 2011).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFE3672EF2BD07CEFB62FCEF.taxon	type_taxon	Type species. Acteon wetherellii Lea, 1833; original designation by Gray (1847: 160). Miocene, New Jersey, USA.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFE3672EF2BD07CEFB62FCEF.taxon	discussion	Discussion. Acteocina Gray, 1847 is a junior synonym of Alicula Eichwald 1830 [type species Alicula okenii Eichwald, 1830; subsequent designation by Glibert (1952 a: 395), Middle Miocene, Ukraine]. Since Acteocina is a widely used name and Alicula is rarely used, Guzhov (2022 b: 1237) proposed to treat Alicula Eichwald, 1830 as nomen oblitum following ICZN Art. 23.9.2 and Acteocina Gray, 1847 as a nomen protectum. However, Alicula was used as subgenus of Retusa by Glibert (1952: 395). Later, Aristov & Storozhenko (2011: 112) treated Alicula Schlechtendal, 1913 [Grylloblattida] as a junior homonym of Alicula Eichwald, 1830. As a result, Alicula Eichwald cannot be rejected without an application to the ICZN. We have not done this yet. For the sake of stability, however, we prefer to use Acteocina herein but emphasize that the problem must be solved formally by the ICZN. Acteocina was represented in the Karpatian and Badenian of the Central Paratethys by few morphologically ‘ stable’ species. During the early Sarmatian (Volhynian) and middle Sarmatian (Bessarabian), the genus underwent a radiation and displayed a high morphological disparity. This phase peaked during the Late Miocene Bessarabian in the Eastern Paratethys. These taxa are not included in our revision, which focuses on the Volhynian occurrences of the Central Paratethys (see Guzhov 2022 b for references on Eastern Paratethyan literature).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFE36728F2BD049AFED3F89B.taxon	description	Figs 34 A – J	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFE36728F2BD049AFED3F89B.taxon	materials_examined	Type material. Lectotype (designated by Zilch 1934: 277): XII 2546 a, SL: 2.5 mm, MD: 1.1 mm, Coşteiu de Sus (Romania), illustrated in Zilch (1934: pl. 22, fig. 19). The specimen was destroyed during WW II and the photographic glass plate from Zilch (1934) is damaged. Illustrated material. NHMW 1876 / 0011 / 0170, SL: 2.7 mm, MD: 1.2 mm, Lăpugiu de Sus (Romania), Figs 34 A 1 – A 2. NHMW 2020 / 0004 / 1661, SL: 3.8 mm, MD: 1.7 mm, Bad Vöslau (Austria), Figs 34 B 1 – B 3. NHMW 2020 / 0004 / 1661 a, SL: 3.3 mm, MD: 1.6 mm, Bad Vöslau (Austria), Figs 34 C 1 – C 2. NHMW 1867 / 0019 / 0322, SL: 3.2 mm, MD: 1.5 mm, Coșteiu de Sus (Romania), Figs 34 D 1 – D 2. NHMW 1997 z 0178 / 1210, SL: 3.8 mm, MD: 1.6 mm, Bad Vöslau (Austria), Figs 34 E 1 – E 3. NHMW 1997 z 0178 / 1210 a, SL: 3.2 mm, MD: 1.4 mm, Bad Vöslau (Austria), Figs 34 F 1 – F 2. NHMW 1876 / 0011 / 0170 a, SL: 4.1 mm, MD: 1.8 mm, Lăpugiu de Sus (Romania), Figs 34 G 1 – G 3. NHMW 2008 / 0248 / 0479, SL: 2.4 mm, MD: 1.1 mm, Coșteiu de Sus (Romania), Figs 34 H 1 – H 3. NHMW 1859 / 0037 / 0570, SL: 3.3 mm, MD: 1.5 mm, Vienna / Pötzleinsdorf (Austria), Figs 34 I. NHMW-ZOO-MO- 75000 - E- 3317 A, SL: 3.4 mm, MD: 1.5 mm, Bad Vöslau (Austria), holotype of Acteocina heraclitica Berger, 1949, illustrated in Berger (1953: pl. 19, fig. 70), Fig. 34 J.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFE36728F2BD049AFED3F89B.taxon	description	Revised description. Shell small, solid, moderately slender, elongate ovate with slightly convex sides (SL / MD = 2.2 – 2.3). Protoconch of about 1.5 whorls, flattened, strongly tilted (~ 140 °), with nucleus two-thirds covered by first teleoconch whorl. Teleoconch of about three whorls separated by deeply canaliculated suture. Spire conical, weakly cyrtoconoid (apical angle 81 – 95 °), broad (MD / ApD = 1.5 – 1.6). Early teleoconch whorls weakly convex, nearly straight sided with periphery at abapical suture, rapidly increasing in height. Last whorl 90 – 93 % of total height, no shoulder, rounded at slowly contracting base. Periphery placed just above mid-whorl (SL / PH = 1.7 – 1.8). Surface glossy with delicate, shallowly arcuate growth lines. Aperture 80 – 83 % of total height, moderately narrow in upper half (MD / AW = 7.3 – 7.6), widening anteriorly (MD / MAW = 2.1 – 2.5), resulting in moderate expansion rate (ER = 3.0 – 3.5). Outer lip simple, thin, shallowly arcuate in profile, straight to slightly pinched inwards just below mid-whorl, rounded at base. Columella short, moderately to weakly excavated, thickened, forming narrow rim with prominent central columellar fold. Columellar callus delimiting narrow, shallow umbilical chink, persisting continuously into narrow, moderately thickened parietal callus. Paratethyan synonyms. Acteocina heraclitica Berger, 1949 is a subjective junior synonym of Acteocina fusiformis (Boettger, 1907). Berger (1949, 1953) did not discuss Acteocina fusiformis when describing Acteocina heraclitica and most probably overlooked this species, although he mentioned Acteocina heraclitica also from Coşteiu de Sus (Romania).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFE36728F2BD049AFED3F89B.taxon	discussion	Discussion. Acteocina fusiformis (Boettger, 1907) is a widespread species in the Badenian of the Central Paratethys Sea, which is characterized by its olivoid shell, without a shoulder, and by the prominent columellar fold. The northeastern Atlantic occurrences referred to as Acteocina lajonkaireana in the literature might be a species complex. Many specimens from the Early to Middle Miocene of France, described by Peyrot (1932), Glibert 1952 a) and Lozouet et al. (2001) as Acteocina lajonkaireana agree with Acteocina fusiformis in their slender outline and columellar fold, whilst the syntype of Acteocina lajonkaireana is comparatively stout and has a weak fold (see Fig. 37 H). Specimens from the Tarkhanian and Chokrakian of Russia, illustrated as this species by Iljina (1993), differ in their high spire, strongly convex spire whorls and incised but not canaliculate suture and represent another species. Paleoenvironment. Shallow marine, inner neritic; e. g., Bernhardsthal (Austria) in Harzhauser et al. (2018 a).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFE36728F2BD049AFED3F89B.taxon	distribution	Distribution in Central Paratethys. Karpatian (Early Miocene): Korneuburg Basin: Kleinebersdorf, Stetten, Weinsteig (Austria) (Harzhauser 2002). Badenian (Middle Miocene): Polish Fore-Carpathian Basin: Błonie (Poland) (Urbaniak 1974). North Alpine Foreland Basin: Guntersdorf (Austria) (Berger 1953); Vienna Basin: Bernhardsthal, Bad Vöslau, Enzesfeld, Steinebrunn, Vienna / Pötzleinsdorf (Austria) (Berger 1953; Harzhauser et al. 2018 a), Borský Mikuláš (Slovakia) (Švagrovský 1984); Danube Basin: Modra-Kráľová (Slovakia) (Hladilová & Fordinál 2013); Oberpullendorf Basin: Ritzing (Austria) (Berger 1953); Pannonian Basin: Várpalota (Hungary) (Strausz 1966); Eastern Slovakian Basin: Kuzmice (Slovakia) (Švagrovský 1960); Dacian Basin: Islaz (Romania) (Rado & Muţui 1970); Banat: Zorlenţul-Mare (Romania) (Florei 1961); Făget Basin: Coşteiu de Sus, Lăpugiu de Sus (Romania) (Boettger 1907). Proto-Mediterranean Sea. Serravallian (Middle Miocene): Karaman Basin: Seyithasan (Turkey) (Landau et al. 2013).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFE6672AF2BD0786FDEEFB8F.taxon	description	Figs 35 A – D	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFE6672AF2BD0786FDEEFB8F.taxon	materials_examined	Type material. Syntype illustrated in Eichwald (1852: pl. 11, fig. 19), probably Żukowce (Zhukivtsi) (Ukraine), stored in the St. Petersburg State University, Paleontological Museum (Russia). Illustrated material. NHMW-ZOO-MO- 75000 - E- 47062, SL: 3.4 mm, MD: 2.0 mm, Żukowce (Zhukivtsi) (Ukraine), Figs 35 A 1 – A 3. NHMW 1859 / 9945 / 0507, SL: 4.0 mm, MD: 2.4 mm, Golubytsya (Hołubica), (Ukraine), Figs 35 B 1 – B 2. NHMW 1843 / 0009 / 0043, SL: 3.4 mm, MD: 1.9 mm, Żukowce (Zhukivtsi) (Ukraine), Figs 35 C 1 – C 4. NHMW-ZOO-MO- 75000 - E- 47067, SL: 4.3 mm, MD: 2.5 mm, Golubytsya (Hołubica) (Ukraine), Figs 35 D 1 – D 3.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFE6672AF2BD0786FDEEFB8F.taxon	description	Revised description. Shell small, solid, broad, subcylindrical ovate with slightly convex sides (SL / MD = 1.6 – 1.8). Protoconch mammillate of about 1.5 whorls, flattened, strongly tilted (~ 140 °), with nucleus more than half covered by first teleoconch whorl. Teleoconch of about 3.5 whorls separated by deeply canaliculated suture. Spire conical (apical angle 89 – 105 °), broad (MD / ApD = 1.5 – 1.6). Early teleoconch whorls cylindrical with narrow subsutural shelf, rapidly decreasing in height. Periphery placed just above mid-whorl (SL / PH = 1.7 – 1.8). Last whorl inflated, 83 – 90 % of total height, rounded shoulder, weakly convex below, rounded at rapidly contracting base. Surface smooth except for delicate, shallowly arcuate growth lines. Aperture 78 – 83 % of total height, moderately narrow in upper third (MD / AW = 7.2 – 9.3), widening in anterior third (MD / MAW = 2.4 – 2.8), resulting in moderate expansion rate (ER = 3.0 – 3.6). Outer lip simple, thin, shallowly arcuate in profile, straight in upper two-thirds, rounded at base. Columella short, moderately excavated, thickened, forming narrow, adherent rim, without or with only subobsolete columellar fold, persisting continuously into narrow parietal callus. Paratethyan synonyms. Bulla clandestina Dubois de Montpéreux, 1831, from Żukowce (Zhukivtsi) (Ukraine), is a subjective junior synonym of Acteocina lichtensteinii (Eichwald, 1830).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFE6672AF2BD0786FDEEFB8F.taxon	discussion	Discussion. Acteocina lichtensteinii (Eichwald, 1830) differs from all its Paratethyan congeners in its broadly barrel-shaped last whorl. It is known so far only from the late Badenian (Serravallian) of the Polish-Ukrainian Fore-Carpathian Basin, which was a hotspot of endemism at that time (Harzhauser et al. 2024 a). Acteocina lichtensteinii is most probably another of these endemic species, which did not spread to other Paratethyan basins. Paleoenvironment. Inner neritic, shallow marine based on associated fauna (own data M. H.).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFE6672AF2BD0786FDEEFB8F.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Voronyaky Hills: Golubytsya (Hołubica), Yaseniv (Ukraine); Polish-Ukrainian Fore-Carpathian Basin: Biłka, Oles’ko, Tarnoruda, Zalistsi, Zhukivtsi, ravine Zhabyak (Ukraine) (Friedberg 1938).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFE76724F2BD03BAFCEBF91B.taxon	description	Figs 36 A – H	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFE76724F2BD03BAFCEBF91B.taxon	materials_examined	Type material. Lectotype (designated herein), NHMW 1846 / 0037 / 0526 b, SL: 15.2 mm, MD: 7.8 mm, Vienna / Gaudenzdorf (Austria), illustrated in Hörnes (1856: pl. 50, figs 9 a – b), Figs 36 B 1 – B 3. Akera (Bullina) melinda De Gregorio, 1885 was introduced for Bulla lajonkaireana of Hörnes 1856 (note that De Gregorio 1885 a erroneously referred to fig, 7 on pl. 50 in Hörnes (1856), which shows “ Bulla convoluta ” but clearly indicated discussed Hörnes’ Bulla lajonkaireana. Unfortunately, Hörnes (1856) illustrated two different species in his figs 9 a – b and 9 c – d. Herein, we select the specimen illustrated by Hörnes (1856: pl. 50, figs 9 a – b) as lectotype of Akera (Bullina) melinda De Gregorio, 1885, whereas the specimen of figs 9 c – d is Acteocina okenii (Eichwald, 1830). Illustrated material. NHMW 1846 / 0037 / 0527, SL: 15.9 mm, MD: 7.4 mm, Vienna / Gaudenzdorf (Austria), Figs 36 C 1 – C 3. NHMW 1846 / 0037 / 0526 a, SL: 15.4 mm, MD: 7.4 mm, Vienna / Gaudenzdorf (Austria), Figs 36 A 1 – A 2. NHMW-ZOO-MO- 75000 - E- 42947 A, SL: 8.0 mm, MD: 4.2 mm, Vienna / Ottakring (Austria), illustrated in Berger (1953: pl. 20, fig. 97), holotype of Acteocina lajonkaireana ventricosa Berger, 1953, Figs 36 D 1 – D 2. NHMW-ZOO-MO- 75000 - E- 42947, SL: 7.7 mm, MD: 4.0 mm, Vienna / Ottakring (Austria), paratype of Acteocina lajonkaireana ventricosa Berger, 1953, Figs 36 E 1 – E 2. NHMW-ZOO-MO- 75000 - E- 42947, SL: 8.9 mm, MD: 4.5 mm, Vienna / Ottakring (Austria), paratype of Acteocina lajonkaireana ventricosa Berger, 1953, Figs 36 F 1 – F 3. NHMW-ZOO-MO- 75000 - E- 47061, SL: 11.3 mm, MD: 5.4 mm, Vienna / Hundsturm (Austria), paratype of Acteocina lajonkaireana maxima Berger, 1953, Figs 36 G 1 – G 3. NHMW-ZOO-MO- 75000 - E- 47061 A, SL: 12.3 mm, MD: 6.2 mm, Vienna / Hundsturm (Austria), illustrated in Berger (1953: pl. 20, fig. 97), holotype of Acteocina lajonkaireana maxima Berger, 1953, Figs 36 H 1 – H 2. Additional material. 22 spec., NHMW 1862 / 0001 / 0515, Vienna / Gaudenzdorf (Austria). 4 spec., NHMW 1866 / 0040 / 0402, Vienna / Hernals (Austria).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFE76724F2BD03BAFCEBF91B.taxon	description	Revised description. Shell large, extraordinarily thick-shelled, moderately broad, subcylindrical with almost parallel to slightly convex sides (SL / MD = 1.9 – 2.0). Protoconch about 1.5 whorls, flattened, strongly tilted (~ 140 – 160 °); usually partly covered by first teleoconch whorl, rarely completely protruding. Teleoconch of about three whorls separated by deeply canaliculated suture. Spire conical (apical angle 73 – 100 °), broad (MD / ApD = 1.5 – 1.7), spire whorls rapidly increasing in height, with rounded shoulder and moderately convex periphery placed just above mid-whorl, rarely in abapical third (SL / PH = 1.5 – 1.8). Last whorl 88 – 95 % of total height, rounded shoulder, subcylindrical below, rounded at rapidly contracting base. Surface smooth except for delicate to prominent, arcuate growth lines. Aperture 75 – 88 % of total height, moderately narrow in upper half (MD / AW = 6.9 – 7.0), widening in anterior third (MD / MAW = 2.0 – 2.1), resulting in moderate expansion rate (ER = 3.2 – 3.5). Outer lip simple, solid, shallowly arcuate in profile, straight in upper two-thirds, rounded at base. Columella short, excavated, thickened, expanded over base, without columellar fold, persisting continuously into thick, broad parietal callus. Paratethyan synonyms. Berger (1949, 1953) was unaware of the paper by De Gregorio (1885 a) and introduced Acteocina lajonkaireana ventricosa for the specimens from Vienna / Gaudenzdorf, which included also the herein designated lectotype of Acteocina melinda. Therefore, Acteocina ventricosa is a subjective junior synonym of Acteocina melinda. In addition, Berger (1953) introduced Acteocina lajonkaireana maxima as new subspecies for very large specimens. These specimens do not differ from the slightly smaller specimens and therefore, we treat Acteocina maxima also as subjective junior synonym of Acteocina melinda. Friedberg (1928) illustrated two syntypes of Tornatina okenii Eichw. var. buhlovensis Friedberg, 1928 (Friedberg 1928: pl. 35, figs 13, 14). Of these, fig. 13 represents Acteocina volhynica (Eichwald 1830), whereas his fig. 14 corresponds to Acteocina melinda. By selecting this specimen as lectotype of Acteocina buhlovensis (Friedberg, 1928), this species becomes a subjective junior synonym of Acteocina melinda (De Gregorio, 1885).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFE76724F2BD03BAFCEBF91B.taxon	discussion	Discussion. Acteocina melinda (De Gregorio, 1885) is characterized by its unusually large size for the genus, extraordinarily thick shell and very thick, band-like columellar-parietal callus. It is a common species in offshore deposits and is clearly separated from its Sarmatian congeners without intermediate morphologies. Therefore, we treat it at full species rank. Acteocina melinda appears abruptly in the late Volhynian (Ervilia Zone) of the Vienna Basin and might be an offshoot of Acteocina volhynica (Eichwald, 1830). Its appearance coincides with a phase of increased shell thickness and shell size of many Sarmatian mollusk lineages correlating with increased sea water salinity and alkalinity (Piller & Harzhauser 2005; Neubauer et al. 2013). Paleoenvironment. Muddy bottoms in middle to outer neritic environments (own data, M. H.).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFE76724F2BD03BAFCEBF91B.taxon	distribution	Distribution in Central Paratethys. Sarmatian (Middle Miocene): Ukrainian-Polish Fore-Carpathian Basin: Buhłowa (Poland) (Friedberg 1928); Vienna Basin: Vienna / Gaudenzdorf, Vienna / Hundsturm, Vienna / Hernals, Vienna / Nussdorf, Vienna / Ottakring (Austria) (Berger 1953).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFE96727F2BD000FFC69F99A.taxon	materials_examined	Type material. Lectotype (designated herein). PM SPbGU, no. 3 / 524 b, stored in the Paleontological Museum at the Department of Dynamic and Historical Geology of St. Petersburg State University (Russia), SL: 4.1 mm, MD: 2.0 mm, illustrated in Guzhov (2022 b: pl. 36, fig. 1). According to Guzhov (2022 b), two localities are indicated on the labels: Chișinău (Moldova) (= Bessarabian) and Kuncha (Ukraine, Khmelnitskyi Region) (= Volhynian). It is unclear where the specimen was collected.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFE96727F2BD000FFC69F99A.taxon	description	Revised description. Shell small, solid, moderately slender, subcylindrical to abapically slightly tapering with weakly convex sides (SL / MD = 2.0). Protoconch, flattened, strongly tilted; nucleus covered more than half covered by first teleoconch whorl. Teleoconch of about three whorls separated by deeply canaliculated suture. Spire broad conical, weakly gradate (apical angle 97 °), moderately broad (MD / ApD = 1.6). Early teleoconch whorls nearly straight sided with periphery placed at abapical suture, moderately increasing in height. Last whorl 92 % of total height, angled shoulder, rapidly contracting base. Periphery slightly above mid-whorl (SL / PH = 1.8). Surface smooth with delicate, shallowly arcuate growth lines. Aperture 85 % of total height, moderately narrow in upper half (MD / AW = 6.4), widening in anterior third (MD / MAW = 2.1), resulting in low rate of expansion 2.9). Outer lip thin. Columella short, distinctly excavated, thickened, forming broad rim with sharp, low, central columellar fold. Columellar callus adherent, without umbilical chink. Columellar callus into broad, thin parietal callus.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFE96727F2BD000FFC69F99A.taxon	discussion	Discussion. Two names are available for Sarmatian Acteocina species from Eichwald (1830). Alicula volhynica Eichwald, 1830 and Alicula okenii Eichwald, 1830. The illustration of Alicula okenii corresponds well to typical Sarmatian shells discussed herein as Acteocina volhynica. It lacks a columellar fold and Eichwald (1853) described the columella as smooth. However, Guzhov (2022 b) illustrated a syntype of Acteocina okenii, which shows a low but distinct columellar fold. This specimen is designated herein as lectotype of Alicula okenii Eichwald, 1830. None of the specimens of Acteocina volhynica studied by us shows a columellar fold and therefore, we refrain from treating them as conspecific with Acteocina okenii. Therefore, we restrict Acteocina okenii (Eichwald, 1830) to specimens with columellar fold. Paleoenvironment. Shallow marine, inner neritic based on co-occurring assemblage (own data, M. H.).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFE96727F2BD000FFC69F99A.taxon	distribution	Distribution in Central Paratethys. Not known from the Central Paratethys. Eastern Paratethys. Volhynian (early Sarmatian, Middle Miocene): Kuncha (Ukraine, Khmelnitskyi Region) or Bessarabian (middle Sarmatian, Late Miocene): Chișinău (Moldova).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFEA6723F2BD018FFF39FB13.taxon	description	Figs 37 A – G	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFEA6723F2BD018FFF39FB13.taxon	materials_examined	Type material. Syntype PM SPbGU, no. 3 / 528 a, stored in the Paleontological Museum at the Department of Dynamic and Historical Geology of St. Petersburg State University (Russia), SL: 3.2 mm, Zalistsi (Ukraine), Sarmatian, illustrated in Guzhov (2022 b: pl. 35, fig. 9). Illustrated material. NHMW 1870 / 0013 / 0012, SL: 3.5 mm, MD: 1.7 mm, Vienna / Heiligenstadt (Austria), Figs 37 A 1 – A 3. NHMW 1870 / 0013 / 0012 a, SL: 3.8 mm, MD: 1.8 mm, Vienna / Heiligenstadt (Austria), Figs 37 B 1 – B 2. NHMW 1860 / 0001 / 0346 a, SL: 3.7 mm, MD: 1.9 mm, Lednice (= Eisgrub) (Czech Republic), Figs 37 C 1 – C 3. NHMW 1861 / 0001 / 0355, SL: 4.3 mm, MD: 2.1 mm, Podivín (= Kostel) (Czech Republic), Figs 37 D 1 – D 2. NHMW 1861 / 0001 / 0355 a, SL: 4.7 mm, MD: 2.3 mm, Podivín (= Kostel) (Czech Republic), Figs 37 E 1 – E 2. NHMW 1861 / 0001 / 0355 b, SL: 4.3 mm, MD: 2.1 mm, Podivín (= Kostel) (Czech Republic), Figs 37 F 1 – F 2. NHMW 1872 / 0003 / 00 o 6, SL: 4.3 mm, MD: 2.3 mm, Vienna / Hernals (Austria), Figs 37 G 1 – G 2.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFEA6723F2BD018FFF39FB13.taxon	description	Revised description. Shell small, solid, moderately slender, subcylindrical to abapically slightly tapering with weakly convex to rarely slightly concave sides (SL / MD = 2.0 – 2.1). Protoconch of about 1.5 whorls, flattened, strongly tilted (~ 140 °); nucleus covered, usually more than half covered by first teleoconch whorl, rarely largely protruding. Teleoconch of about three whorls separated by deeply canaliculated suture. Spire broad conical, often slightly dome-shaped (apical angle 90 – 115 °), moderately broad (MD / ApD = 1.7 – 1.8). Early teleoconch whorls weakly convex, nearly straight sided with periphery placed at abapical suture, usually moderately increasing in height. Last whorl 88 – 92 % of total height, weak, rounded shoulder, rapidly contracting base, with periphery slightly above mid-whorl (SL / PH = 1.6 – 1.7). Surface glossy with delicate, shallowly arcuate growth lines. Aperture 83 – 89 % of total height, moderately narrow in upper half (MD / AW = 5.5 – 6.6), widening in anterior third (MD / MAW = 2.0 – 2.2), resulting in low rate of expansion 2.7 – 3.0). Outer lip thin, shallowly arcuate in profile, straight to faintly concave mid-whorl, rounded at base. Columella short, distinctly excavated, thickened, forming broad rim without columellar fold. Columellar callus adherent, without umbilical chink. Columellar callus persisting continuously into broad, prominent parietal callus. Paratethyan synonyms. Bulla terebellata Dubois de Montpéreux, 1831, from Żukowce (Zhukivtsi) (Ukraine), is a subjective junior synonym of Acteocina volhynica (Eichwald, 1830). Bulla polymorpha Andrzejowski 1832. Andrzejowski (1832: 551) introduced Bulla polymorpha as name for the specimens illustrated by Dubois de Montpéreux 1831: pl. 1, fig. 8 [Bulla terebellata Dubois de Montpéreux 1831], fig. 11 [‘ Bulla spirata ’] and fig 13 [‘ Bulla ovulata ’]. These three specimens represent two different species: figs 8, 11 = Acteocina volhynica (Eichwald, 1830) and fig. 13 = Pyrunculus elongatus (Eichwald, 1830). By selecting the specimen illustrated in Dubois de Montpéreux (1831: pl. 1, fig. 11) as type specimen for Bulla polymorpha Andrzejowski, 1832, this species becomes a subjective junior synonym of Acteocina volhynica (Eichwald, 1830). Weinkauffia pumila Švagrovský, 1971. Holotype No, 2251, Comenius University, Bočiar (Slovakia) (now stored in the Slovak Natural History Museum, Bratislava), SL: 3.0 mm, MD: 1.6 mm). This species was based on a single specimen of Acteocina volhynica (Eichwald, 1830) in which the last whorl covers nearly the entire spire.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFEA6723F2BD018FFF39FB13.taxon	discussion	Discussion. Acteocina volhynica (Eichwald, 1830) has been confused with Acteocina lajonkaireana (de Basterot, 1825) since Hauer (1837) and Hörnes (1856). Acteocina lajonkaireana was originally described by de Basterot (1825: 22, pl. 1, fig. 25) from the Burdigalian of Saucats (France) (see syntype MNHN. F. A 71213, SL: 3.9 mm, MD: 1.9 mm, Figs 37 H 1 – H 3). This small species has a moderately broad, elongate ovate outline with low spire and broad columellar / parietal callus with weak columellar fold. Since then, this species became a dumping ground for Miocene Tornatinidae. The Sarmatian A. volhynica differs from A. lajonkaireana in its broader shell, the more convex sides, the more exposed spire, which is largely covered by the last whorl in the French species resulting in a bullet-shaped outline. Paleoenvironment. Ubiquitous in intertidal to shallow sublittoral environments (Mandic et al. 2008; Lukeneder et al. 2011).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFEA6723F2BD018FFF39FB13.taxon	distribution	Distribution in Central Paratethys. Sarmatian (Middle Miocene): Lublin upland: Modliborzyce (Poland) (Bielecka 1967); Polish-Ukrainian Fore-Carpathian Basin: Chlebów, Chmielnik, Dwikozy, Słupcza, Żolobki (Poland); Bialozurka, Krzemieniec, Mizocz, Rydomyl’, Słupcza, Zaleśce (Ukraine) (Friedberg 1938); North Alpine Foreland Basin: Hollabrunn / Aspersdorf (Austria) (Mandic et al. 2008); Vienna Basin: Hauskirchen, Höflein, Hölles, Kollnbrunn, Wiesen, Siebenhirten, Vienna / Döbling, Vienna / Penzing, Vienna / Ottakring, Vienna / Sechshaus (Austria) (Papp 1954); Lednice (Czech Republic) (hoc opus); Čejč, Bílovice u Uherského Hradiště, Holič, Podivín, Šaštinské Strážne, Rohožník, Trkmanec (Slovakia) (Švagrovský 1971); Eisenstadt-Sopron Basin: Wiesen (Austria) (Papp 1954); Danube Basin: Modra-Kráľová (Slovakia) (Hladilová & Fordinál 2013); Styrian Basin: Osek- 1 in Slovenske gorice (Slovenia) (Mikuž & Križnar 2015); Pannonian Basin: Budapest, Pécsvárad, Karád, Sopron, Úny, Várpalota, Vérkút (Hungary) (Boda 1959); Eastern Slovakian Basin: Bidovce, Bohdanovce, Bočiar, Herľany, Hnojné, Izra, Kalša, Nižná Myšľa, Nižný Čaj, Nové Mesto, Sečovce, Skaroš, Slanec, Slančík, Stankovce, Slanec, Studený potok, Trebišov, Trstené pri Hornáde (Slovakia) (Švagrovský 1954, 1971); Pădurea Craiulu Mountains: Vida Valley (Romania) (Filipescu et al. 1999); Poiana Ruscă Mountains: Răcăştia (Romania) (Tămaș et al. 2013); Făget Basin: Buituri (Romania) (Moisescu 1955). Eastern Paratethys. Numerous records from the Volhynian (Middle Miocene) and Bessarabian (Late Miocene) of Romania, Moldova and Russia (Kolesnikov 1935; Iljina 1993; Ionesi 2006; (Frolov et al. 2020; Guzhov 2022 b).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFEF6722F2BD04C7FBCDFA8F.taxon	type_taxon	Type species. Bulla coronata Lamarck, 1804; original designation by Cossmann (1889: 314). Eocene, France.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFEF6722F2BD04C7FBCDFA8F.taxon	discussion	Discussion. This genus is characterized by its strongly elongate pyriform shape with a prominent, raised, convex apical crest delimiting the deep and narrow umbilicus. It was originally known by several species from the Eocene of the Paris Basin (e. g. Cossmann 1889) but persisted at least to the Langhian in the Central Paratethys Sea (Berger 1953) and to the Serravallian in the Proto-Mediterranean Sea (Landau et al. 2013). In the northeastern Atlantic it is documented from the Aquitanian and Burdigalian (Peyrot 1932). At present the genus Acrostemma Cossmann, 1889 is considered a subjective synonym of Cylichna Lovén, 1846 by WoRMS (MolluscaBase eds. 2025 c). However, this synonymy is not supported by any published research and the elevated coronate edge of the apical umbilicus is quite different to that of the type species of Cylichna, C. cylindracea (Pennant, 1777).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFEF673CF2BD02BBFF37FE07.taxon	description	Figs 39 B – C	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFEF673CF2BD02BBFF37FE07.taxon	materials_examined	Type material. Holotype, NHMW-ZOO-MO- 75000 - E- 3310 A, SL: 5.8 mm, MD: 2.2 mm, Bad Vöslau (Austria), Figs 39 C 1 – C 3. Paratype, NHMW-ZOO-MO- 75000 - E- 3310, Bad Vöslau (Austria), SL: 4.8 mm, MD: 1.8 mm, Bad Vöslau (Austria), Figs 39 B 1 – B 3. Revised description. Medium sized, slender, elongate pyriform shell (SL / MD = 2.6 – 2.8), only last whorl visible. Protoconch unknown. Last whorl greatly elongated, weakly convex, periphery placed mid-whorl (SL / PH = 1.9 – 2.0), almost straight above periphery, forming angle of about 23 – 28 °. Rounded at slowly contracting base. Apex sharply truncated, moderately narrow (MD / ApD = 1.9 – 2.0), apical plane moderately oblique (AP = 14 – 15 °), deeply and narrowly umbilicate, delimited by prominent, elevated, narrowly rounded apical crest, laterally protruding over adjacent whorl side. Surface of apical crest with close-set axial growth lines. Shell surface smooth except for moderately deep, apically fading spiral grooves over base. Aperture longer than shell, very narrow down to mid-whorl (MD / AW = 8.4 – 8.6), weakly widening anteriorly (MD / MAW = 2.5), resulting in moderate expansion rate (ER = 3.3 – 3.5). Outer lip thin, relatively high, narrowly alate apical extension. Columella short, distinctly excavated. Columellar callus forming delicate, narrow rim delimiting deep, narrow umbilical chink. Parietal callus not developed.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFEF673CF2BD02BBFF37FE07.taxon	discussion	Discussion. Landau et al. (2013) treated Acrostemma edlaueri (Berger, 1953) and Acrostemma kuehnelti (Berger, 1953) as subjective junior synonyms of Coelophysis latesulcata Boettger 1902 [= Pyrunculus latesulcatus (Boettger, 1902)]. This decision was based on the poor drawings in Berger (1953) and the poorly preserved holotype of Pyrunculus latesulcatus. Herein, we could study all type specimens and agree with Berger (1953) to treat all three taxa as distinct species. The type species Acrostemma coronata (Lamarck, 1804), from the Middle Eocene Lutetian of the Paris Basin, differs from Acrostemma edlaueri in its even slenderer shape and the absence of spiral sculpture (see Cossmann & Pissarro 1913: pl. 44, fig. 242 - 8). Acrostemma boutillieri (Cossmann, 1889), from the Upper Eocene Bartonian of the Paris Basin, has a broader apex and its outer lip rises only slightly above the apex (see Cossmann 1889: pl. 11, fig. 25). Acrostemma bezanconi (Morlet, 1885), also from the Upper Eocene of the Paris Basin, is more ovate with convex periphery, has a nearly straight columella and a more prominent apical spiral rim (see Cossmann 1889: pl. 11, fig. 12). Acrostemma tournoueri (Peyrot, 1932), from the Lower Miocene Aquitanian and Burdigalian of the Aquitaine Basin is slightly larger (SL: 7.5 mm), has a more convex periphery and a straight columella (see Peyrot 1932: pl. 12, figs 3 – 5). Paleoenvironment. The occurrence in the Baden Formation at Bad Vöslau (Austria) suggests middle to outer neritic environments with up to 210 m water depth (Kranner et al. 2021).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFEF673CF2BD02BBFF37FE07.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienn Basin: Bad Vöslau (Austria) (Berger 1953).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFF1673CF2BD0532FEEBFA5B.taxon	description	Figs 39 A 1 – A 3	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFF1673CF2BD0532FEEBFA5B.taxon	materials_examined	Type material. Holotype, NHMW-ZOO-MO- 75000 - E- 42897, SL: 5.2 mm, MD: 1.6 mm, Vienna / Ottakring (Austria), illustrated in Berger (1953: pl. 16, fig. 31), Figs 39 A 1 – A 3.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFF1673CF2BD0532FEEBFA5B.taxon	description	Revised description. Medium sized, strongly elongate, very slender pyriform shell (SL / MD = 3.2), only last whorl visible. Protoconch unknown. Last whorl subcylindrical, weakly convex; periphery placed just above mid-whorl (SL / PH = 1.9), almost straight in adapical third, forming angle of about 23 °. Rounded at slowly contracting base. Apex sharply truncated, moderately broad (MD / ApD = 1.9), apical plane strongly oblique (AP = 19 °), deeply and narrowly umbilicate, delimited by prominent, round-edged apical crest. Surface of apical crest with broad, low, short growth lines. Sculpture of numerous moderately deep, partly punctate spiral grooves over entire last whorl. Broad, flat spiral cords occasionally bifid. Aperture longer than shell, very narrow down to mid-whorl (MD / AW = 8.3), weakly widening anteriorly (MD / MAW = 2.7), resulting in moderate expansion rate (ER = 3.1). Outer lip thin, relatively high, very narrowly alate apical extension. Columella obscured by sediment. Columellar callus forming delicate, narrow rim delimiting umbilical chink.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFF1673CF2BD0532FEEBFA5B.taxon	discussion	Discussion. This species differs from all congeners in its very slender outline and prominent spiral sculpture. Acrostemma anatolica sp. nov. is broader, the apical crest is more prominent, and the spiral grooves become increasingly wider spaced adapically on the last whorl. Paleoenvironment. Probably middle neritic based on co-occurring assemblage (own data, M. H.).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFF1673CF2BD0532FEEBFA5B.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienna Basin: Vienna / Ottakring (Austria) (Berger 1953).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFF1673FF2BD02CEFED3FDB7.taxon	description	Figs 39 D 1 – D 2	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFF1673FF2BD02CEFED3FDB7.taxon	materials_examined	Type material. Holotype, RGM 776925, SL: 3.5 mm, MD: 1.2 mm, Seyithasan (Turkey), Figs 39 D 1 – D 2 Type locality. Seyithasan (Turkey). Type stratum. Silt and clay of the Týrtar Formation. Age. Middle Miocene, Serravallian.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFF1673FF2BD02CEFED3FDB7.taxon	etymology	Etymology. Referring to Anatolia in Turkey.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFF1673FF2BD02CEFED3FDB7.taxon	diagnosis	Diagnosis. Small, slender ovate shell with truncated apex and prominent, convex apical crest, sculpture of somewhat irregular spiral grooves, close-set over base, adapically becoming distinctly wider spaced.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFF1673FF2BD02CEFED3FDB7.taxon	description	Description. Shell small, slender ovate (SL / MD = 2.6), only last whorl visible. Protoconch unknown. Last whorl weakly convex; periphery placed just below mid-whorl (SL / PH = 2.3). Rounded at slowly contracting base. Apex sharply truncated, moderately narrow (MD / ApD = 2.2), apical plane moderately oblique (AP = 13 °), deeply and narrowly umbilicate, delimited by prominent, elevated, round-edged apical crest. Surface of apical crest with delicate, close-set axial riblets. Sculpture of numerous moderately deep, somewhat uneven spiral grooves, close-set on base, gradually wider spaced adapically. Aperture longer than shell, very narrow down to mid-whorl (MD / AW = 10.2), weakly widening anteriorly (MD / MAW = 2.7), resulting in moderate expansion rate (ER = 3.8). Outer lip thin, relatively high, alate apical extension. Columella short, weakly excavated with weak columellar fold. Columellar callus forming delicate, narrow rim delimiting narrow umbilical chink.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFF1673FF2BD02CEFED3FDB7.taxon	discussion	Discussion. Acrostemma anatolica sp. nov. is characterized by its peculiar spiral sculpture, which distinguishes it from the morphologically similar Eocene Acrostemma boutillieri (Cossmann, 1889) and Acrostemma bezanconi (Morlet, 1885) (see Cossmann 1889: pl. 11, figs 12, 25). Acrostemma kuehnelti (Berger, 1953) is much slenderer and has more numerous and relatively evenly spaced spiral grooves that cover the entire last whorl. Paleoenvironment. Inner neritic, shallow marine (own data B. L.).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFF1673FF2BD02CEFED3FDB7.taxon	description	Proto-Mediterranean Sea. Serravallian (Middle Miocene): Karaman Basin: Seyithasan (Turkey) (Landau et al. 2013).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFF2673FF2BD05A3FEF2FCEF.taxon	type_taxon	Type species. Bulla cylindracea Pennant, 1777; subsequent designation by Herrmannsen (1852: 42). Present-day, British Isles.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFF26739F2BD049BFDB5FB77.taxon	description	Figs 40 A – E	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFF26739F2BD049BFDB5FB77.taxon	materials_examined	Type material. Holotype, NHMW-ZOO-MO- 75000 - E- 18735, SL: 12.2 mm, MD: 4.8 mm, Guntersdorf (Austria), Figs 40 A 1 – A 3. Paratypes: NHMW-ZOO-MO- 75000 - E- 18735, SL: 8.9 mm, MD: 3.5 mm, Guntersdorf (Austria), Figs 40 B 1 – B 3. NHMW-ZOO-MO- 75000 - E- 18735, SL: 9.5 mm, MD: 3.9 mm, Guntersdorf (Austria), Figs 40 C 1 – C 3. NHMW 1859 / 0038 / 0135, SL: 13.7 mm, MD: 5.8 mm, Grund (Austria), Figs 40 D 1 – D 3. NHMW 2025 / 0153 / 0001, SL: 8.7 mm, MD: 3.6 mm, Stetten (Austria), Figs 40 E 1 – E 2. Additional paratypes. NHMW 1846 / 0037 / 0524, SL: 13.2 mm, MD: 4.8 mm, Niederkreuzstetten (Austria), illustrated in Hörnes (1856: pl. 50, fig. 6). NHMW 1846 / 0037 / 0524, SL: 17.7 mm, MD: 7.2 mm, Niederkreuzstetten (Austria). Type locality. Guntersdorf (Austria). Type stratum. Grund Formation. Age. Early Badenian, Langhian, Middle Miocene.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFF26739F2BD049BFDB5FB77.taxon	etymology	Etymology. Latin caudex, = tree trunk or stem; noun in apposition.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFF26739F2BD049BFDB5FB77.taxon	diagnosis	Diagnosis. Large, solid, slender cylindrical with elongate last whorl, nearly parallel sides, truncated, narrowly umbilicate, no columellar fold; delicate, wide-set spiral grooves and color stripes.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFF26739F2BD049BFDB5FB77.taxon	description	Description. Large, solid, slender cylindrical (SL / MD = 2.4 – 2.5), only last whorl visible. Last whorl elongate, with almost parallel, straight to very weakly convex sides, weakly rounded at base; apex truncated, moderately narrow (MD / ApD = 2.3 – 2.6) with moderately oblique apical plane (AP = 13 – 15 °), narrowly and shallowly umbilicate, delimited by low, rounded crest. Surface with delicate growth lines and delicate, wide-set spiral grooves. Growth lines and spiral grooves coincide with brownish color stripes. Aperture as long as the shell, narrow adapically (MD / AW = 5.4 – 7.0) and mid-section, widening anteriorly (MD / MAW = 1.8 – 2.0), with moderate expansion rate (ER = 3.0 – 4.0). Outer lip slightly extending above apex, straight to very weakly convex below. Columella short, forming narrow rim, sharply delimited from base; no columellar fold. Parietal callus subobsolete.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFF26739F2BD049BFDB5FB77.taxon	discussion	Discussion. Cylichna caudex sp. nov. was confused with Cylichna subcylindrica (d’Orbigny, 1852) by Csepreghy-Meznerics (1954) and Harzhauser (2002). Cylichna subcylindrica occurred during the Early to Middle Miocene in the Circum-Mediterranean Region, but the shells described by Peyrot (1932: 13, figs 21 – 23) from France, by A. W. Janssen (1984: pl. 19, fig. 3) from the Netherlands, and by Landau et al. (2013) from Turkey, are much smaller than the Paratethyan specimens (SL 3 – 6 mm versus 8.9 – 17.7 mm). The Paratethyan specimens were identified by Hörnes (1856) and Berger (1953) as the Pliocene Mediterranean Cylichna brocchii (Michelotti, 1847), which attains a comparable size but lacks the color pattern, differs in its more convex last whorl and the presence of a distinct columellar fold (see Landau et al. 2023: pl. 40). Cylichna caudex sp. nov. is subcylindrical whereas C. brocchii is more conical. Retusa pliocrassa (Sacco, 1897), from the Mediterranean Pliocene, is also large but differs in its wider and deeper apical umbilicus and stronger spiral grooves at both extremities (see Landau et al. 2013: pl. 42). Cylichna caudex sp. nov. differs from the morphologically similar Cylichna cylindracea (Pennant, 1777) in its much larger size and the spiral sculpture / color pattern (although this may not always be preserved). Cylichna clathrata (Defrance, 1825) also has a color pattern, is similar in size and does not have a columellar fold but differs in its much broader, interrupted spiral bands of color looking like rows of square spots. Paleoenvironment. The occurrence at Niederkreuzstetten and Stetten (Austria) indicate shallow marine, inner neritic environments (Harzhauser et al. 2002).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFF26739F2BD049BFDB5FB77.taxon	distribution	Distribution in Central Paratethys. Karpatian (Early Miocene): Korneuburg Basin: Niederkreuzstetten, Stetten (Austria) (Harzhauser 2002); Badenian (Middle Miocene: Nowy Sącz Basin: Niskowa (Poland) (Bałuk 1970); North Alpine-Carpathian Foreland Basin: Grund, Guntersdorf (Austria) (Hörnes 1856; hoc opus); Cserhát Hills: Mátraverebély, Márkháza, Piliny (Hungary) (Csepreghy-Meznerics 1954); Bükk Mountains: Borsodbóta (Hungary) (Csepreghy-Meznerics 1972).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFF4673BF2BD03E3FECEFBE3.taxon	description	Figs 41 A – E	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFF4673BF2BD03E3FECEFBE3.taxon	materials_examined	Type material. We are not aware of the whereabouts of the type material. Illustrated material. NHMW 1846 / 0037 / 0525, SL: 4.8 mm, MD: 1.8 mm, Steinebrunn (Austria), illustrated in Hörnes (1856: pl. 50, figs 7 a – c), Figs 41 A 1 – A 3. NHMW 2013 / 0078 / 06554, SL: 5.8 mm, MD: 2.2 mm, Baden (Austria), Figs 41 B 1 – B 3. NHMW 1997 z 0178 / 1209, SL: 5.4 mm, MD: 2.1 mm, Bad Vöslau (Austria), Figs 41 C 1 – C 3. NHMW 2013 / 0078 / 06554 a, SL: 5.8 mm, MD: 2.4 mm, Baden (Austria), Figs 41 D 1 – D 2. NHMW 2013 / 0078 / 06554 d, SL: 4.9 mm, MD: 2.0 mm, Baden (Austria), Figs 41 E 1 – E 3.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFF4673BF2BD03E3FECEFBE3.taxon	description	Revised description. Shell medium sized, solid, slender cylindrical (SL / MD = 2.5 – 2.6), only last whorl visible. Last whorl elongate, with almost parallel, straight to very weakly convex sides, weakly rounded at base; apex moderately broad (MD / ApD = 1.7 – 1.8), truncated, weakly oblique (AP = 11 – 13 °) widely and shallowly umbilicate, delimited by low, sharp ridge. Surface glossy with few, extremely fine spirals visible on tangential light in rare specimens. Aperture as long as the shell, very narrow adapically and in mid-section (MD / AW = 8.6 – 10.3), widening anteriorly (MD / MAW = 2.5 – 3.0), with moderate expansion rate (ER = 2.7 – 3.5). Outer lip only slightly extending above apex, straight to very weakly convex below, rounded and somewhat truncated abapically. Columella short, not excavated; columellar callus forming narrow rim with weak, low columellar fold. Narrow umbilical chink.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFF4673BF2BD03E3FECEFBE3.taxon	discussion	Discussion. Hörnes (1856) and several subsequent authors compared the Paratethyan shells with the Pliocene Mediterranean Cylichna convoluta (Brocchi, 1814), which is a subjective junior synonym of Cylichna cylindracea (Pennant, 1777) (see Landau et al. 2023 for discussion). Paleoenvironment. Cylichna cylindracea lives in inner neritic environments on muddy sands between 20 - 60 m depth (Gofas et al. 2011).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFF4673BF2BD03E3FECEFBE3.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene: Polish-Ukrainian Foreland Basin: Borki Wielkie, Dryszczów (Poland), Shushkivtsi, Tarnoruda, Yaseniv, Zboriv (Ukraine) (Friedberg 1938); Błonie (Poland) (Urbaniak 1974); Vienna Basin: Baden, Bad Vöslau, Steinebrunn (Austria) (Hörnes 1856); Pannonian Basin: Mátraverebély (Hungary) (Strausz 1966); Bükk Mountains: Borsodbóta (Hungary) (Csepreghy-Meznerics 1972). Northeastern Atlantic. Tortonian (Late Miocene): southern Portugal (Landau et al. 2023). Pliocene to present-day: North Sea, northeastern Atlantic and Mediterranean Sea (see Landau et al. 2023: 393 for details).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFF66735F2BD0397FD74FE5F.taxon	description	Figs 42 A 1 – A 3	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFF66735F2BD0397FD74FE5F.taxon	materials_examined	Type material. Holotype, NHMW 1847 / 0037 / 0130, SL: 7.1 mm, MD: 3.5 mm, Mikulov (Czech Republic), illustrated in Hörnes (1856: pl. 50, figs 8 a – b), Figs 42 A 1 – A 3. Type locality. Mikulov (Czech Republic). Type stratum. Coastal part of Baden Formation. Age. Middle Miocene, middle Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFF66735F2BD0397FD74FE5F.taxon	etymology	Etymology. Latin dolium (= barrel), referring to the elongate barrel-shaped outline; noun in apposition.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFF66735F2BD0397FD74FE5F.taxon	diagnosis	Diagnosis. Medium sized, solid, moderately slender subcylindrical shell with truncated, deeply, narrowly umbilicate apex, delicate spiral sculpture at extremities, excavated columella with weak columellar fold, faint color pattern of interrupted dark spiral bands.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFF66735F2BD0397FD74FE5F.taxon	description	Description. Shell medium sized, solid, moderately slender subcylindrical (SL / MD = 2.0), only last whorl visible. Last whorl elongate, with weakly convex, subparallel sides, rapidly contracting at base; periphery above mid-whorl (SL / PH = 1.6); apex truncated, moderately narrow (MD / ApD = 2.4), moderately oblique (AP = 12 °), deeply and narrowly umbilicate, delimited by rounded ridge. Surface smooth except for faint spiral grooves at apex and narrow spiral cords over base. Aperture as long as the shell, moderately narrow adapically and in mid-section (MD / AW = 7.3), widening anteriorly (MD / MAW = 2.4), with moderate expansion rate (ER = 3.0). Outer lip not extending above apex, straight to very weakly convex below, rounded abapically. Columella short, slightly excavated; columellar callus forming relatively broad rim, adherent to base. Weak columellar fold. No parietal callus. No umbilical chink. Faint color pattern of numerous dark spiral bands.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFF66735F2BD0397FD74FE5F.taxon	discussion	Discussion. This species was confused by Hörnes (1856) and Berger (1953) with the Early Miocene Cylichna clathrata (Defrance, 1825), from the northeastern Atlantic. This was probably due to the similar color pattern of dotted / interrupted bands preserved in both species. However, the French species, as illustrated by Peyrot (1932: pl. 13, figs 11, 15), has its periphery distinctly below mid-whorl, lacks a columellar fold and its outer lip extends distinctly above the apex. Cylichna clathrata as illustrated by de Basterot (1825: pl. 1, fig. 10) has a broad, convex apex and lacks a columellar fold. Cylichna dolium sp. nov. differs from the similar Cylichna intermedia (Sacco, 1897) in its truncated apex, less elongate shell and rapidly contracting base. Cylichna miosimplex sp. nov. is much smaller, its outer lip extends slightly above the apex, and it lacks a columellar fold. Steininger (1963: 57) mentioned Cylichna clathrata from the Eggenburgian (Early Miocene) of Fels am Wagram (Austria). We have not seen this material and therefore, its identity remains unclear. Paleoenvironment. Shallow marine, inner neritic based on the associated fauna (own data, M. H.).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFF66735F2BD0397FD74FE5F.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienna Basin: Mikulov (Czech Republic), Bad Vöslau, Gainfarn (Austria) (Berger 1953).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFF86735F2BD06DCFED3F821.taxon	description	Figs 43 A – E	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFF86735F2BD06DCFED3F821.taxon	materials_examined	Type material. Syntype, BS. 090.02.019, SL: 8.8 mm, MD: 3.8 mm, Tetti Borelli (Italy), Late Miocene, illustrated in Ferrero Mortara et al. (1984: pl. 53, fig. 1). Illustrated material. NHMW-ZOO-MO- 75000 - E- 7083, SL: 8.3 mm, MD: 3.6 mm, Gainfarn (Austria), Figs 43 A 1 – A 3. NHMW-ZOO-MO- 75000 - E- 47087, SL: 7.9 mm, MD: 3.7 mm, Steinebrunn (Austria), Figs 43 B 1 – B 3. NHMW-ZOO-MO- 75000 - E- 33312, SL: 8.5 mm, MD: 3.9 mm, Guntersdorf (Austria), Figs 43 C 1 – C 2. NHMW 1854 / 0035 / 0386, SL: 6.7 mm, MD: 3.1 mm, Lăpugiu de Sus (Romania), Figs 43 D 1 – D 3. NHMW 1997 z 0178 / 1884, SL: 9.2 mm, MD: 4.2 mm, Gainfarn (Austria), Figs 43 E 1 – E 3.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFF86735F2BD06DCFED3F821.taxon	description	Revised description. Shell medium sized, solid, moderately slender subcylindrical (SL / MD = 2.2 – 2.3), only last whorl visible. Last whorl elongate, with weakly convex sides, slowly contracting at base; periphery just above mid-whorl (SL / PH = 1.8 – 1.9); apex low, convex, narrow (MD / ApD = 2.8 – 2.9), weakly oblique (AP = 9 – 11 °), deeply and narrowly umbilicate, delimited by convex ridge. Surface smooth except for fine spirals over base. Aperture as long as the shell, moderately narrow adapically and in mid-section (MD / AW = 6.7 – 7.0), widening anteriorly (MD / MAW = 2.2 – 2.3), with low to moderate expansion rate (ER = 2.9 – 3.3). Outer lip slightly extending above apex, straight to very weakly convex below, rounded abapically. Columella short, slightly excavated; columellar callus forming relatively broad rim, slightly extending over base, with distinct, narrow columellar fold. Columellar callus short, slightly expanded over medial side of base. No parietal callus. Narrow umbilical chink.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFF86735F2BD06DCFED3F821.taxon	discussion	Discussion. Cylichna clathrata (Defrance, 1825), from the Early Miocene of France as illustrated by Peyrot (1932: pl. 13, figs 11, 15), differs from Cylichna intermedia (Sacco, 1897) in its nearly straight sided last whorl, has its periphery distinctly below mid-whorl (SL / PH = 2.5), lacks a columellar fold and has a much wider maximum aperture width (MD / MAW = 1.8) (versus 2.2 – 2.3 in Cylichna intermedia). Landau et al. (2013: 331, pl. 52, fig. 18) mistakenly attributed specimens from the Serravallian Karaman Basin of Turkey to C. clathrata. The illustrated specimen clearly shows a terminal columellar ridge and they are herein reassigned to C. intermedia. Bulla clathrata sensu de Basterot (1825: pl. 1, fig. 10) has a comparable shape to C. intermedia but lacks a columellar fold. Paleoenvironment. Shallow marine, inner neritic based on the associated fauna (own data, M. H.).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFF86735F2BD06DCFED3F821.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): North Alpine-Carpathian Foreland Basin: Grund, Guntersdorf (Austria) (Berger 1953); Vienna Basin: Mikulov (Czech Republic), Gainfarn, Steinebrunn, Vienna / Pötzleinsdorf (Austria) (Berger 1953); Pannonian Basin: Budapest (Hungary) (Kókay 1996); Southern Pannonian Basin: Hrvaćani, Miljevići (Bosnia and Herzegovina) (Atanacković 1985); Făget Basin: Buituri, Lăpugiu de Sus (Romania) (Berger 1953). Proto-Mediterranean Sea. Serravallian (Middle Miocene): Karaman Basin: Akpınar, Seyithasan (Turkey) (Landau et al. 2013). Tortonian / Messinian (Late Miocene): Colli Torinesi: Tetti Borelli (Italy) (Ferrero Mortara et al. 1984).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFF96737F2BD0040FB64FB15.taxon	description	Figs 42 B 1 – B 3	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFF96737F2BD0040FB64FB15.taxon	materials_examined	Type material. Holotype, NHMW-ZOO-MO- 75000 - E- 19695, SL: 3.6 mm, MD: 1.7 mm, Poysdorf (Austria), Figs 42 B 1 – B 3. Type locality. Poysdorf (Austria). Type stratum. Baden Formation. Age. Middle Miocene, middle Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFF96737F2BD0040FB64FB15.taxon	etymology	Etymology. Referring to the Pliocene Retusa pliosimplex, with which this species was confused by Berger (1953).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFF96737F2BD0040FB64FB15.taxon	diagnosis	Diagnosis. Small, thin-shelled, glossy, smooth, moderately slender subcylindrical shell with low, weakly truncated apex, oblique, columella not excavated forming narrow rim, without fold.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFF96737F2BD0040FB64FB15.taxon	description	Description. Shell small, thin-shelled, moderately slender subcylindrical (SL / MD = 2.2), only last whorl visible. Last whorl elongate, with weakly convex, subparallel sides, rapidly contracting at base; periphery placed mid-whorl (SL / PH = 2.1); apex low, weakly truncated moderately narrow (MD / ApD = 2.4), moderately oblique (AP = 10 °), deeply and narrowly umbilicate, delimited by convex ridge. Surface smooth, glossy, no spiral grooves. Aperture as long as the shell, moderately narrow adapically and in mid-section (MD / AW = 6.9), widening anteriorly (MD / MAW = 2.2), with moderate expansion rate (ER = 3.2). Outer lip only slightly extending above apex, straight below, with rather abrupt transition into weakly rounded base. Columella short, oblique, not excavated; columellar callus forming narrow rim, delimiting narrow umbilical chink. No columellar fold. No parietal callus.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFF96737F2BD0040FB64FB15.taxon	discussion	Discussion. The illustrated specimen (Figs 42 B 1 – B 3) was identified as Retusa pliosimplex (Sacco, 1897) by Berger (1953), which was described by Sacco (1897) from the Pliocene of Italy. The syntype of Retusa pliosimplex was re-illustrated by Ferrero Mortara et al. (1984: pl. 52, figs 10 a – c) and differs from the Paratethyan species distinctly in its elongate, slightly pyriform outline with its periphery far below mid-whorl, the flattened apical crest and the much wider umbilicus. Cylichna intermedia (Sacco, 1897) is larger, has more convex sides and a columellar fold. Similarly, Cylichna pliocrassa (Sacco, 1897), from the Pliocene of Italy and Spain, differs in its larger size, broader, funnel-shaped apical umbilicus and in its prominent spiral sculpture at both extremities (see Ferrero Mortara et al. 1984: pl. 53, figs 4 – 5; Landau et al. 2023: pl. 42). Paleoenvironment. Shallow marine, inner neritic based on associated fauna (own data M. H.)	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFF96737F2BD0040FB64FB15.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Carpathian Foreland Basin: Žabčice (Czechia) (Montanaro Gallitelli & Tacoli 1951); Vienna Basin: Poysdorf (Austria) (hoc opus).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFFA6736F2BD0205FDB0FCEF.taxon	description	Figs 42 C 1 – C 3	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFFA6736F2BD0205FDB0FCEF.taxon	materials_examined	Type material. Holotype, NHMW-ZOO-MO- 75000 - E- 20707, SL: 8.6 mm, MD: 4.1 mm, Žabčice (Czech Republic), illustrated in Berger (1953: pl. 17, fig. 47), Figs 42 C 1 – C 3. Type locality. Žabčice (Czech Republic) - Type stratum. Badenian silt and sand. Age. Middle Miocene, middle Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFFA6736F2BD0205FDB0FCEF.taxon	etymology	Etymology. In honor of Anita Eschner, zoologist at the NNMW, in recognition for her kind help.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFFA6736F2BD0205FDB0FCEF.taxon	diagnosis	Diagnosis. Medium sized, solid, moderately slender shell with weakly convex periphery placed mid-whorl, moderately broad, truncated apex and deep, narrow apical umbilicus; sculpture of prominent spiral cords at extremities and within apical umbilicus; no columellar fold.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFFA6736F2BD0205FDB0FCEF.taxon	description	Description. Shell medium sized, solid, moderately slender SL / MD = 2.1). Only last whorl visible. Protoconch unknown. Periphery placed mid-whorl (SL / PH = 1.9) with weakly convex sides. Rounded at rapidly contracting base. Apex truncated, moderately broad (MD / ApD = 1.9), apical plane moderately oblique (AP = 11 °), deeply and narrowly umbilicate, delimited by rounded apical crest. Sculpture of prominent spiral cords within apical umbilicus and on apical crest. Prominent spiral grooves at extremities, subobsolete mid-whorl (note that spiral grooves are poorly preserved on available specimen). Aperture as long as shell, moderately narrow down to below mid-whorl (MD / AW = 6.9), slowly widening anteriorly (MD / MAW = 2.5), resulting in low expansion rate (ER = 2.7). Outer lip thin, rising moderate distance above apex. Columella short, straight, oblique, without fold. Columellar callus forming very narrow rim delimiting narrow deep umbilical chink. Parietal callus not developed.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFFA6736F2BD0205FDB0FCEF.taxon	discussion	Discussion. Cylichna elongata (Grateloup, 1837), from the Burdigalian of Saint-Paul (France), might be a closely related species. The French species is comparable in size (SL: 7 mm) and sculpture; prominent spiral grooves at the extremities but differs in its narrower apex and wider apical umbilicus (see Peyrot 1932: pl. 12, figs 6 – 7, 12, pl. 13, figs 1, 3). Cylichna eschnerae sp. nov. was identified by Berger (1949, 1953) as Retusa (Cylichnina) testiculina (Bonelli). Cylichna testiculina (Sacco, 1897) is based on at least six syntypes from the Middle Miocene of the Colli Torinesi (Italy), representing two different species. To clarify the situation, we select specimen BS. 090.02.014, illustrated in Ferrero Mortara et al. (1984: pl. 53, fig. 3) as lectotype of Bullinella (Cylichnina) testiculina Sacco, 1897. Thus, Cylichna testiculina differs from the Paratethyan species in its cylindrical shape, rounded apex, delicate spiral sculpture and adherent columellar callus. Boettger (1907: 206) mentioned Cylichna testiculina also from the Badenian of Coşteiu de Sus (Romania), but it remains unclear which species he had at hand, because the material was destroyed during WW II. Paleoenvironment. Unknown.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFFA6736F2BD0205FDB0FCEF.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): North Alpine-Carpathian Foreland Basin: Žabčice (Czech Republic) (Berger 1953).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFFB6731F2BD049BFDE2FE97.taxon	description	Figs 42 D 1 – D 3	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFFB6731F2BD049BFDE2FE97.taxon	materials_examined	Type material. SL: 8.1 mm, MD: 3.6 mm, Institute for Paleontology, University Vienna, no. nr., illustrated in Steininger (1963: pl. 12, fig. 18), Figs 42 D 1 – D 3. Type locality. Fels am Wagram (Austria). Type stratum. Fels Formation. Age. Early Miocene, early Eggenburgian (Aquitanian).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFFB6731F2BD049BFDE2FE97.taxon	etymology	Etymology. In honor of Fritz F. Steininger (Eggenburg, Austria), mentor of the first author, in recognition of his impact on paleontological research in the Eggenburg region.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFFB6731F2BD049BFDE2FE97.taxon	diagnosis	Diagnosis. Medium sized, solid, moderately slender, subcylindrical shell with periphery below mid-whorl and prominent, moderately broad, flattish spiral cords covering entire shell being most prominent at extremities; broad, semicircular columellar callus slightly extending over base, no columellar fold.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFFB6731F2BD049BFDE2FE97.taxon	description	Description. Shell medium sized, solid, moderately slender, subcylindrical (SL / MD = 2.3), only last whorl visible. Last whorl elongate, with almost parallel sides forming angle of 5 °; straight to very weakly convex, weakly rounded at rapidly contracting base; periphery below mid-whorl (SL / PH = 2.7); apex truncated, moderately narrow (MD / ApD = 1.8) with moderately oblique apical plane (AP = 13 °), moderately narrowly and shallowly umbilicate, delimited by low, rounded crest. Surface with moderately wide-set spiral grooves, delimiting broad, flattish spiral cords, most prominent at extremities but also present along mid-whorl. Aperture as long as the shell, narrow adapically (MD / AW = 5.8) and mid-section, widening anteriorly (MD / MAW = 2.5), with moderate expansion rate (ER = 2.4). Outer lip slightly extending above apex, straight to very weakly concave below. Columella short, strongly excavated, forming broad, semicircular rim, sharply delimited from base; no columellar fold. Parietal callus subobsolete.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFFB6731F2BD049BFDE2FE97.taxon	discussion	Discussion. This species was confused by Steininger (1963) with Cylichna elongata (Grateloup, 1837) from the Burdigalian of St-Paul-lès-Dax (France). The French species has a comparable size (SL: 7 mm) and prominent spiral grooves at the extremities but differs in its convex mid-whorl periphery and the less excavated columella (see Peyrot 1932: pl. 12, figs 6 – 7, 12, pl. 13, figs 1, 3). The extant IWP species Cylichna consobrinoides Kuroda & Habe, 1952, is reminiscent of Cylichna steiningeri sp. nov. in general shape and sculpture but its alate outer lip extends distinctly above the apex (see Valdés 2008: fig. 35 A). The Atlantic Early Miocene Cylichna clathrata (Defrance, 1825) differs in its color pattern of broader interrupted spiral bands looking like rows of square spots, but if not present further differs in the outer lip being more alate apically and the columella far less excavated. Paleoenvironment. Sandy coastal environments (Steiniger 1963).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFFB6731F2BD049BFDE2FE97.taxon	distribution	Distribution in Central Paratethys. Eggenburgian (Early Miocene): North Alpine Foreland Basin: Fels am Wagram (Austria) (Steininger 1963).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFFC6730F2BD0683FA81FDCF.taxon	description	Figs 42 E 1 – E 2, 44 C	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFFC6730F2BD0683FA81FDCF.taxon	materials_examined	Type material. Holotype, NHMW-ZOO-MO- 75000 - E- 2201, SL: 2.2 mm, MD: 1.0 mm, Bad Vöslau (Austria), Figs 42 E 1 – E 2, 44 C.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFFC6730F2BD0683FA81FDCF.taxon	description	Revised description. Shell small, moderately slender cylindrical (SL / MD = 2.2), only last whorl visible. Last whorl elongate, with almost parallel, straight to very weakly convex sides, rounded at base; apex moderately narrow (MD / ApD = 2.6), convex, weakly oblique (AP = 10 °) very shallowly umbilicate. Glossy, entire surface with delicate, regular spiral grooves. Aperture slightly longer than shell, very narrow adapically down below mid-whorl (MD / AW = 8.0), abruptly widening anteriorly (MD / MAW = 1.7), with high expansion rate (ER = 4.6). Outer lip slightly extending above apex, straight to very weakly concave below, rounded abapically. Columella short, strongly excavated; columellar callus forming delicate rim. No columellar fold, no umbilical chink, no parietal callus.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFFC6730F2BD0683FA81FDCF.taxon	discussion	Discussion. When establishing Cylichna subbacillus, Berger (1953) referred to the extant Bulla bacillus Ehrenberg from the Red Sea, described by Pilsbry (1895: 311). This species has never been illustrated, and its status is unknown to us. Cylichna subbacillus differs from its Paratethyan congeners in its small size and comparatively stout outline. Cylichna? subovularis (Sacco, 1897), from the Late Miocene of Italy, differs in its weakly convex sides and weak columellar callus (see Ferrero Mortara et al. 1984: pl. 52, figs 9 a – b). Paleoenvironment. The occurrence in the Baden Formation at Bad Vöslau (Austria) suggests middle to outer neritic environments with up to 210 m water depth (Kranner et al. 2021).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFFC6730F2BD0683FA81FDCF.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienna Basin: Bad Vöslau (Austria).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFFD6730F2BD057BFDE2F8F5.taxon	description	Figs 44 A 1 – A 3	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFFD6730F2BD057BFDE2F8F5.taxon	materials_examined	Type material. SL: 4.4 mm, MD: 2.1 mm, Institute for Paleontology, University Vienna, no. nr., illustrated in Steininger (1963: pl. 12, fig. 12), Figs 44 A 1 – A 3. Type locality. Fels am Wagram (Austria). Type stratum. Fels Formation. Age. Early Miocene, early Eggenburgian (Aquitanian).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFFD6730F2BD057BFDE2F8F5.taxon	etymology	Etymology. In honor of Philipp Strauss (OMV, Vienna) in recognition of his contributions to the geology of the Northern Calcareous Alps.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFFD6730F2BD057BFDE2F8F5.taxon	diagnosis	Diagnosis. Small, moderately slender, subcylindrical shell with faintly angled periphery mid-whorl; surface smooth; columella nearly straight, with delicate rim; no fold.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFFD6730F2BD057BFDE2F8F5.taxon	description	Description. Shell small, moderately slender, subcylindrical (SL / MD = 2.2), only last whorl visible. Last whorl elongate, faintly angled at periphery; sides nearly straight-sided above, forming angle of 19 °; straight to very weakly convex below, rounded at rapidly contracting base; periphery placed just above mid-whorl (SL / PH = 1.8); apex truncated, moderately narrow (MD / ApD = 1.8) with moderately oblique apical plane (AP = 11 °), not umbilicate but with narrow apical depression, delimited by low, relatively narrow, rounded crest. Surface smooth. Aperture as long as the shell, narrow adapically (MD / AW = 6.0) and mid-section, slowly widening anteriorly (MD / MAW = 2.0), with moderate expansion rate (ER = 3.0). Outer lip slightly extending above apex, straight to very weakly concave below. Columella short, nearly straight, weakly twisted, forming delicate, narrow rim, delimiting narrow umbilical chink. No columellar fold. Parietal callus subobsolete.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFFD6730F2BD057BFDE2F8F5.taxon	discussion	Discussion. Steininger (1963) confused this species with Cylichna cylindracea (Pennant, 1777), which also has smooth shell and a shallow apical depression but differs in its larger size and slenderer cylindrical shell. Cylichna caudex sp. nov. is distinctly larger and has prominent spiral sculpture and color pattern. Cylichna sublaevis (d’Orbigny, 1852), from the Burdigalian of France, is comparable in outline and size (SL: 5.5 mm) but differs in its narrow, deep apical umbilicus and its outer lip extends slightly above the apex (see Peyrot 1932: pl. 13, figs 31 – 33; Lozouet et al. 2001: pl. 37, fig. 2). Cylichna ligeriensis (Glibert, 1952), from the Langhian and Tortonian of the Loire Basin (France), is also comparable in its slightly angled periphery (e. g., Landau et al. 2020: pl. 19, figs 2 – 3) but is relatively stouter and differs in its prominent columellar fold. Paleoenvironment. Sandy coastal environments (Steiniger 1963).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFFD6730F2BD057BFDE2F8F5.taxon	distribution	Distribution in Central Paratethys. Eggenburgian (Early Miocene): North Alpine Foreland Basin: Fels am Wagram (Austria) (Steininger 1963).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFFE6733F2BD0786FF0BFCEF.taxon	description	Figs 44 B 1 – B 3	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFFE6733F2BD0786FF0BFCEF.taxon	materials_examined	Illustrated material. NHMW-ZOO-MO- 75000 - E- 3324, SL: 4.3 mm, MD: 2.3 mm, Bad Vöslau (Austria), Figs 44 B 1 – B 3.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFFE6733F2BD0786FF0BFCEF.taxon	description	Description. Small, thin shelled, subcylindrical, moderately broad shell with weakly convex sides and periphery placed below mid-whorl. Apex, moderately broad, slightly truncated, umbilicus reduced to shallow pit with a narrow notch at its base, apical plane shallowly concave. Shell surface glossy with delicate growth lines and faint spiral grooves in apical region. Outer lip extending above apex, thin, straight below. Aperture moderately narrow in adapical part, strongly widening anteriorly. Base not preserved. Columella short, weakly excavated with delicate, narrow rim; no columellar fold.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFFE6733F2BD0786FF0BFCEF.taxon	discussion	Discussion. Only a single fragmentary specimen is available, which lacks the base. It differs from Cylichna dolium sp. nov. in its much wider aperture. Cylichna castexi (Peyrot, 1932), from the Aquitanian of France might be a related species and is comparable in size and shape but seems to differ in its more truncated apex and the fully sealed apical umbilicus (see Peyrot 1932: pl. 13, figs 18 – 20). Paleoenvironment. The occurrence in the Baden Formation at Bad Vöslau (Austria) suggests middle to outer neritic environments with up to 210 m water depth (Kranner et al. 2021).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFFE6733F2BD0786FF0BFCEF.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienna Basin: Bad Vöslau (Austria) (hoc opus).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFFE6733F2BD049AFB47FC3B.taxon	type_taxon	Type species. Bulla bidentata d’Orbigny, 1841, by monotypy. Present-day, Caribbean Sea.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFFE670DF2BD032EFCF9FB8F.taxon	description	Figs 45 A – D	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFFE670DF2BD032EFCF9FB8F.taxon	materials_examined	Type material. Lectotype (designated by Zilch 1934: 276: XII 2549 a, SL: 2.6 mm, MD: 1.7 mm, Coşteiu de Sus (Romania), illustrated in Zilch (1934: pl. 22, fig. 13). The specimen was destroyed during WW II and the photographic glass plate of Zilch (1934) is damaged. Illustrated material. NHMW 1997 z 9178 / 1903, Bad Vöslau (Austria), SL: 2.7 mm, MD: 1.6 mm, Figs 45 A 1 – A 3. NHMW 1997 z 9178 / 1903 a, Bad Vöslau (Austria), SL: 2.8 mm, MD: 1.7 mm, Figs 45 B 1 – B 3. NHMW 187600110169, SL: 3.1 mm, MD: 1.6 mm, Lăpugiu de Sus (Romania), Figs 45 C 1 – C 3. NHMW 185900190131, SL: 2.0 mm, MD: 1.4 mm, Porzteich (Czech Republic), Figs 45 D 1 – D 3.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFFE670DF2BD032EFCF9FB8F.taxon	description	Revised description. Shell small, relatively solid, glossy, ovate, inflated, broad (SL / MD = 1.6 – 1.8), only last whorl visible. Periphery placed just above mid-whorl (SL / PH = 1.7 – 1.8). Apex with shallow umbilical pit, truncated, narrow (MD / ApD = 3.4 – 3.6), weakly oblique (AP = 10 – 14 °), surface smooth. Aperture narrow in adapical half (MD / AW = 10.1 – 10.3), widening distinctly anteriorly (MD / MAW = 2.7 – 3.1), with moderate expansion rate (ER = 3.3 – 3.8). Outer lip extending short distance above apex adapically, convex below, rounded abapically. Columella short, thickened, moderately expanding over base, moderately excavated with one prominent columellar fold. Parietal callus not present. Color pattern of two bands of subquadratic dots (see Kókay 1966: pl. 10, figs 13 – 14). Paratethyan synonyms. Cylichna (Cylichnella) maculata Kókay, 1966 from the Badenian of Hungary is based on two specimens of Cylichnella bitaeniata with preserved color pattern of two bands of spirally aligned rows of dark dots.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFFE670DF2BD032EFCF9FB8F.taxon	discussion	Discussion. The strength of the columellar fold is slightly variable and the shell outline ranges from ovate (typical) to subcylindrical (rare). This species has been confused by many authors with the Mediterranean Pliocene Atys miliaris (Brocchi, 1814), which differs in it its more ovate profile, the outer lip rising above the apex and the lack of a columellar fold (see Pinna & Spezia 1978: pl. 50, fig. 3). The generic placement is tentative, because the Paratethyan species has just one columellar fold, whereas the type species of the genus has two. Only few Cylichnella species are described so far from the Oligocene and Miocene of the Circum-Mediterranean Region and the early Indo-West Pacific Region. Cylichnella vasatensis (Cossmann, 1895), from the Aquitanian and Burdigalian (Early Miocene) of France, is exceptionally similar in outline and size but bears two columellar folds (see Cossmann 1895: pl. 4, figs 14 – 16; Peyrot 1932: pl. 13, figs 24 – 26; Lozouet et al. 2001: pl. 37, fig 4). Cylichnella neuvillei (Peyrot, 1932), from the Burdigalian of France, is much larger (SL: 7 mm), and has a very prominent, sharp adapical columellar fold and a deeper apical umbilical pit (see Peyrot 1932: pl. 13, figs 4 – 6). Similarly, Cylichnella duqmensis Harzhauser, 2007, from the Chattian of the Sultanate of Oman is much larger (SL: 8 mm). Cylichnella durgae (Beets, 1941), from the Late Miocene of Borneo has a slightly truncated apex, prominent spiral grooves and its columella fold persists onto the base (see Beets 1941: pl. 7, figs 291 – 293). Paleoenvironment. The occurrence in the Baden Formation at Bad Vöslau (Austria) suggests middle to outer neritic environments with up to 210 m water depth (Kranner et al. 2021) whereas occurrences at Borský Mikuláš (Slovakia) represent inner neritic environments.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFFE670DF2BD032EFCF9FB8F.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Polish-Ukrainian Fore-Carpathian Basin: Wieliczka (Poland) (Reuss 1867); Vienna Basin: Bad Vöslau, Baden-Sooss, Steinebrunn (Austria) (Boettger 1902; Berger 1953; hoc opus), Porzteich at Břeclav (Czech Republic) (hoc opus); Borský Mikuláš, Devínska Nová Ves (Slovakia) (Švagrovský 1981, 1984); Pannonian Basin: Budapest, Herend-Márkó, Letkés (Hungary) (Kókay 1966; Strausz 1966; pers comm. Zoltán Kovács); Southern Pannonian Basin: Hrvaćani (Bosnia and Herzegovina) (Atanacković 1985); Mecsek Mountains: Hidas (Hungary) (Csepreghy-Meznerics 1950); Făget Basin: Coşteiu de Sus, Lăpugiu de Sus (Romania) (Boettger 1907; hoc opus).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFC0670DF2BD022BFBEAF86F.taxon	type_taxon	Type species. Atys (Alicula) lapugyensis Berger, 1949; Middle Miocene, Central Paratethys Sea.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFC0670DF2BD022BFBEAF86F.taxon	etymology	Etymology. In honor of Thomas A. Neubauer (Bavarian Natural History Collections, Munich, Germany), in recognition of his contributions to Neogene mollusks.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFC0670DF2BD022BFBEAF86F.taxon	diagnosis	Diagnosis. Large, solid, moderately slender club-shaped; only last whorl visible; periphery placed at two-thirds whorl height; apex convex, deeply and narrowly umbilicate; sculpture of numerous flattened spiral cords separated by narrow grooves; outer lip alate; weak columellar fold.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFC0670DF2BD022BFBEAF86F.taxon	description	Description. As for type species.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFC0670DF2BD022BFBEAF86F.taxon	discussion	Discussion. The type species was originally established in Alicula Ehrenberg, 1831 (as subgenus of Atys) by Berger (1940) [Alicula Ehrenberg, 1831 is a junior homonym of Alicula Eichwald, 1830 and consequently Pilsbry (1896; 237) introduced Aliculastrum as replacement name]. Neubaueriella nov. gen. is indeed reminiscent of Aliculastrum Pilsbry, 1896 [type species Bulla cylindrica Helbling, 1779; present-day, Tropical Indo-West Pacific] but differs in the very high placed periphery and in its smaller size (Aliculastrum can become more than twice as large; Too et al. 2014). In addition, the adapical and abapical sides of Aliculastrum are straighter, resulting in a somewhat biconical outline in contrast to the club-shaped outline of Neubaueriella.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFC1670FF2BD0786FDA7FF2F.taxon	description	Figs 46 A – B	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFC1670FF2BD0786FDA7FF2F.taxon	materials_examined	Type material. Holotype, NHMW-ZOO-MO- 75000 - E- 24786 A, SL: 7.0 mm, MD: 3.9 mm, Lăpugiu (Romania), illustrated in Berger (1853: pl. 19, fig. 88), Figs 46 A 1 – A 3. Illustrated material. NHMW 1862 / 0029 / 0920, SL: 11.2 mm, MD: 5.2 mm, Rudice (Czech Republic), Figs 46 B 1 – B 3.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFC1670FF2BD0786FDA7FF2F.taxon	description	Revised description. Large, solid, moderately slender shell with slowly contracting base (SL / MD = 2.2). Only last whorl visible; periphery weakly convex, placed at two-thirds whorl height (SL / PH = 1.5), conical with weakly convex sides below. Apex convex, moderately narrow (MD / ApD = 2.4), deeply and narrowly umbilicate, delimited by indistinctly elevated, round-edged crest. Apical plane around 12 °. Sculpture of numerous prominent, low spiral cords separated by shallow grooves; spiral cords often bifid over base. Grooves weakly punctate over base. Spiral sculpture most prominent on base, weakening slightly along periphery. Aperture longer than shell, narrow below apex (MD / AW = 10.3), slowly widening anteriorly (MD / MAW = 3.6), resulting in low expansion rate (ER = 2.9). Outer lip distinctly alate, extending above apex. Columella moderately short, with thickened, adherent columellar callus expanding slightly over base, bearing weak central columellar fold. Parietal callus not developed.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFC1670FF2BD0786FDA7FF2F.taxon	discussion	Discussion. We are not aware of comparable species from the Circum-Mediterranean Neogene. Neubaueriella lapugyensis (Berger, 1949) is superficially reminiscent of the extant Aliculastrum cylindricum (Helbing, 1779). Aliculastrum cylindricum is larger (SL: 30 mm) and its periphery is lower, in the mid-whorl region (see Too et al. 2014: fig 1 A). In addition, Aliculastrum cylindricum differs in its conical apex, whereas Aliculastrum lapugyensis has a more dome-shaped apex. Paleoenvironment. Shallow marine, inner neritic based on the occurrence at Várpalota and Steinebrunn (own data, M. H.).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFC1670FF2BD0786FDA7FF2F.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienna Basin: Steinebrunn (Austria) (Berger 1953); Carpathian Foreland Basin: Rudice (Czech Republic) (Berger 1953);? Pannonian Basin: Várpalota (Hungary) (Strausz 1966); Bükk Mountains: Borsodbóta (Hungary) (Csepreghy-Meznerics 1972); Făget Basin: Lăpugiu de Sus (Romania) (Berger 1953).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFC3670EF2BD067EFED3F823.taxon	description	Figs 48 A – C	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFC3670EF2BD067EFED3F823.taxon	materials_examined	Type material. Holotype, NHMW-ZOO-MO- 75000 - E- 3290, SL: 3.4 mm, MD: 2.4 mm, Bad Vöslau (Austria), illustrated in Berger (1953: pl. 16, fig. 7), Figs 48 A 1 – A 3. Paratype, NHMW-ZOO-MO- 75000 - E- 36704. SL: 3.4 mm, MD: 2.3 mm, Guntersdorf (Austria), Figs 48 B 1 – B 3. Illustrated material. NHMW 1997 z 01781213, SL: 6.9 mm, MD: 4.6 mm, Bad Vöslau (Austria), Figs 48 C 1 – C 3.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFC3670EF2BD067EFED3F823.taxon	description	Revised description. Shell medium sized, delicate, glossy, very broad, ovate-subcylindrical (SL / MD = 1.5). Only last whorl visible; periphery placed just above mid-whorl (SL / PH = 1.1). Broad, rounded apex (MD / ApD = 1.4), with moderately oblique apical plane (AP = 12 °), moderately wide, shallow umbilicus. Sculpture of weak growth lines. Aperture relatively wide in anterior third (MD / AW = 4.1), greatly widening anteriorly (MD / MAW = 2.0), resulting in low expansion rate (ER = 2.1). Outer lip thin, only slightly rising above apex, weakly convex at periphery, rounded abapically. Columella short, strongly excavated in lower half. Columellar lip forming narrow rim, delimiting narrow umbilical chink.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFC3670EF2BD067EFED3F823.taxon	discussion	Discussion. We follow Berger (1949, 1953) and Landau et al. (2013) and separate the Middle Miocene specimens from Haminoea hydatis (Linnaeus, 1758), which is larger, has a broader, fully adherent columellar callus and an outer lip usually raised above the apex. However, more Miocene specimens will be needed to decide if these features really justify separation from the Pliocene to extant species. Haminoea hydatis var. hydatina (Dod.), from the Tortonian of Montegibbio (Italy) as illustrated by Sacco (1897: pl. 4, fig 6), was listed by Landau et al. (2013) as synonym of Haminoea miocaenica (Berger, 1949). This name would have priority over Haminoea miocaenica, but the fragmentary Italian specimen does not even allow a clear generic identification. Friedberg (1928) listed Bulla inflata Eichwald, 1830, from the Middle Miocene of Ukraine, as synonym of this species (as Bulla hydatis). We have not seen the type specimens of ‘ Bulla ’ inflata and its status remains unclear. The illustration in Eichwald (1852: pl. 11, fig 14) shows an ovate-conical shell with its periphery distinctly below mid-whorl, with narrow adapical aperture and deeply excavated columella, which is quite unlike Haminoea miocaenica (Berger, 1949). Therefore, we tentatively remove it from the chresonymy but note that this name would also have priority over Haminoea miocaenica. Paleoenvironment. The occurrence in the Baden Formation at Bad Vöslau (Austria) suggests middle to outer neritic environments with up to 210 m water depth (Kranner et al. 2021), whereas the occurrences at Steinebrunn (Austria) and Seyithasan (Turkey) point to shallow water environments (own data M. H., B. L.).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFC3670EF2BD067EFED3F823.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Polish-Ukrainian Fore-Carpathian Basin Wieliczka (Poland), Faszczówka, Staryi Pochaiv, Shushkivtsi, Zhukivtsi (Ukraine) (Friedberg 1938); Voronyaky Hills: Oles’ko (Ukraine) (Friedberg 1938); North Alpine-Carpathian Foreland Basin: Hrušovany nad Jevišovkou (Czech Republic), Guntersdorf (Austria) (Berger 1953); Vienna Basin: Bad Vöslau, Steinebrunn, Gaaden (Austria) (Berger 1953). Dacian Basin: Islaz (Romania) (Rado & Muţui 1970). Proto-Mediterranean Sea. Serravallian (Middle Miocene): Karaman Basin: Seyithasan (Turkey) (Landau et al. 2013).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFC3670EF2BD0786FC34FECB.taxon	type_taxon	Type species. Bulla hydatis Linnaeus, 1758; by monotypy. Placed on the Official List and spelling ruled to be Haminoea by Opinion 1942 (ICZN 2000). Present-day, Mediterranean Sea.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFC46709F2BD0236FB0BF8C3.taxon	type_taxon	Type species. Atys muscarius Gould, 1859; original designation. Present-day, Indo-West Pacific-	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFC46709F2BD0236FB0BF8C3.taxon	discussion	Discussion. Vellicolla was established by Oskars et al. (2019) for a group of extant Indo-West Pacific species, characterized by ovate shells with inflated central part, produced apex with round-edged elevated crest, rapidly constricting base, alate outer lip that extends at both extremities past the last whorl, and spiral grooves at both ends. The fossil Paratethyan species herein included in Vellicolla differ from extant species in that the periphery is lower placed. However, the features of the elevated apical crest and the alate portion of the outer lip fit well within the generic concept. The Middle Miocene occurrence in the Central Paratethys Sea indicates a western Tethyan origin of the genus, which is now restricted to the Indo-West Pacific. As the Paratethyan occurrences postdate the closure of the Tethyan Seaway, the roots of the genus must reach back to the Oligocene or Early Miocene, when the Indian Ocean and the Proto-Mediterraean Sea were still connected via the Tethyan Seaway (Harzhauser et al. 2007; Piller et al. 2024). However, we are not aware of Early Miocene Vellicolla species from the Proto-Mediterraean Sea.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFC4670BF2BD0076FCC9FC57.taxon	description	Figs 49 D – E	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFC4670BF2BD0076FCC9FC57.taxon	materials_examined	Type material. Holotype, M. 61.4301, SL: 3.0 mm, MD: 1.7 mm, Szob (Hungary), illustrated in Csepreghy-Meznerics (1952: pl. 14, figs 15 – 17). Picture made by Péter Kóbor, HUN-REN Centre for Agricultural Research, Plant Protection Institute in Martonvásár; Figs 49 D 1 – D 2. Illustrated material. NHMW-ZOO-MO- 75000 - E- 3330, SL: 3.5 mm, MD: 2.1 mm, Bad Vöslau (Austria), holotype of Retusa (Pyrunculus) austriaca Berger 1953, illustrated in Berger (1953: pl. 17, fig. 48), Figs 49 E 1 – E 3.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFC4670BF2BD0076FCC9FC57.taxon	description	Revised description. Shell small, medium-thickness, ovate to ovate-pyriform, broad (SL / MD = 1.7), only last whorl visible. Last whorl strongly inflated; periphery strongly convex, just below mid-whorl (SL / PH = 2.2), rounded at rapidly contracting base. Apex produced, narrowly truncated, moderately narrow (MD / ApD = 2.3), deeply and narrowly umbilicate, strongly oblique (AP = 27 °), delimited by round-edged elevated crest, distinctly concave below. Sculpture of a few spiral grooves just below apex weakening abapically, and over base, strengthening abapically. Aperture longer than shell, slightly widening at apex, moderately narrow below, narrowing towards mid-section (MD / AW = 4.6), distinctly widening anteriorly (MD / MAW = 2.5), resulting in low expansion rate (ER = 1.8). Outer lip narrowly alate apically, extending far above apex, weakly concave below. Columella moderately short, nearly straight with narrow, erect, columellar callus, delimiting broad, deep umbilicus. No columellar fold. Parietal callus not developed. Paratethyan synonyms. Retusa (Pyrunculus) austriaca Berger, 1953, from the Badenian of Bad Vöslau (Austria) is junior synonym of Vellicolla pannonica.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFC4670BF2BD0076FCC9FC57.taxon	discussion	Discussion. Vellicolla pannonica (Csepreghy-Meznerics, 1952) is reminiscent of Vellicolla postneglecta (Berger, 1953) with which it co-occurs, but is smaller, more inflated, with wider aperture and deeper concavity below the apical crest. Its apex is wider and differences in apertural features are well expressed in the smaller expansion rate of Vellicolla pannonica (1.8 versus 2.9 – 3.2). The extant Indo-West Pacific Vellicolla muscaria (Gould, 1859) differs mainly in the higher position of the periphery and the rapidly contracting base. Paleoenvironment. The occurrence at Szob (Hungary) suggests shallow marine environments (Dulai 1996). Occurrences in the Baden Formation at Bad Vöslau (Austria) suggests middle to outer neritic environments with up to 210 m water depth (Kranner et al. 2021).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFC4670BF2BD0076FCC9FC57.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienna Basin: Bad Vöslau (Austria) (Berger 1953); Pannonian Basin: Szob (Hungary) (Strausz 1966).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFC6670BF2BD04C3FC80F827.taxon	description	Figs 49 A – C	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFC6670BF2BD04C3FC80F827.taxon	materials_examined	Type material. Holotype, NHMW-ZOO-MO- 75000 - E- 47055 A, SL: 4.3 mm, MD: 2.1 mm, Lăpugiu (Romania), Figs 49 A 1 – A 3. Paratypes, NHMW-ZOO-MO- 75000 - E- 11797, SL: 4.2 mm, MD: 2.4 mm, Bad Vöslau (Austria), Figs 49 B 1 – B 2; NHMW-ZOO-MO- 75000 - E- 11797, SL: 4.2 mm, MD: 2.3 mm, Bad Vöslau (Austria), Figs 49 C 1 – C 3.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFC6670BF2BD04C3FC80F827.taxon	description	Revised description. Shell small, medium-thickness, ovate-pyriform, broad (SL / MD = 1.8 – 2.0), only last whorl visible. Last whorl strongly inflated; periphery placed just below mid-whorl (SL / PH = 2.2 – 2.5), rounded at rapidly contracting base. Apex produced, truncated, moderately narrow (MD / ApD = 2.5 – 2.8), deeply and narrowly umbilicate, apical plane strongly oblique (AP = 27 – 33 °), delimited by round-edged elevated crest. Sculpture of a few spiral grooves just below apex weakening abapically, and over base, strengthening abapically. Aperture longer than shell, slightly widening at apex, moderately narrow below, narrowing towards mid-section (MD / AW = 7.3 – 7.9), distinctly widening anteriorly (MD / MAW = 2.3 – 2.7), resulting in moderate low rate of 2.9 – 3.2. Outer lip narrowly alate apically, extending some distance above apex, weakly concave below. Columella short, with narrow columellar callus, delimiting narrow, deep umbilicus. No columellar fold. Parietal callus not developed.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFC6670BF2BD04C3FC80F827.taxon	discussion	Discussion. The sculpture of spiral grooves was overemphasized in the illustration of Berger (1953: pl. 17, fig. 49). For comparison with Vellicolla pannonica (Csepreghy-Meznerics, 1952), see above. Vellicolla postneglecta is reminiscent of the extant Pyrunculus pyriformis (Adams, 1850), from the Philippines, but differs in its less inflated last whorl and the wider umbilicus. Vellicolla okomotoi (Habe, 1952) is very similar but the latter has a higher placed periphery and the apical crest is more strongly constricted below. Pyrunculus cf. postneglectus of Stein et al. (2016: pl. 74, figs 4 – 5), from the early Miocene of the North Sea differs in its much broader apex and its outer lip is less raised above the apex. Paleoenvironment. The occurrence in the Baden Formation at Bad Vöslau (Austria) suggests middle to outer neritic environments with up to 210 m water depth (Kranner et al. 2021).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFC6670BF2BD04C3FC80F827.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienna Basin: Bad Vöslau (Austria) (Berger 1953); Făget Basin: Lăpugiu de Sus (Romania) (Berger 1953).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFC7670AF2BD0636FB10FE5F.taxon	type_taxon	Type species. Bulla hirundinina Quoy & Gaimard, 1833; by monotypy. Present-day, Mauritius.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFC76705F2BD06CAFF37FECB.taxon	description	Figs 50 B 1 – B 2	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFC76705F2BD06CAFF37FECB.taxon	materials_examined	Type material. Holotype, MWG UW ZI / 93 / 1251, SL: 1.6 mm, MD: 1.2 mm, Badenian, Korytnica (Poland), paratype of Chelidonura radwanskii Bałuk, 2018, illustrated in Bałuk (2018: text-figs 2 A 1 – 2 A 2), Figs 50 B 1 – B 2. Type locality. Korytnica (Poland). Type stratum. Pińczów Formation. Age. Middle Miocene, middle Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFC76705F2BD06CAFF37FECB.taxon	etymology	Etymology. In honor of Andrzej Pisera (Polish Academy of Sciences, Warsaw, Poland), in recognition of his contributions to paleontology.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFC76705F2BD06CAFF37FECB.taxon	diagnosis	Diagnosis. Small, delicate shell with tiny, stalked protoconch and rapidly expanding wing without adapical crest; weak, wide-spaced growth lines.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFC76705F2BD06CAFF37FECB.taxon	description	Description. Shell small, delicate. Protoconch tiny, oval knob-shaped, stalked. Teleoconch of one strongly expanding, weakly convex wing-like shield, with delicate, wide-spaced growth lines, becoming almost flat and thin. Adapical margin of shield strengthened but without crest. Inner surface glossy, smooth, slightly uneven.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFC76705F2BD06CAFF37FECB.taxon	discussion	Discussion. This species was lumped by Bałuk (2018) with his Chelidonura radwanskii Bałuk, 2018 [= Melanochlamys radwanskii (Bałuk, 2018)] although both species are very different. Melanochlamys radwanskii (Bałuk, 2018) is larger and differs from Chelidonura piserai sp. nov. in its very prominent and solid crest and its protoconch is not stalked. The extant Mediterranean Chelidonura mediterranea (Swennen, 1961) [listed as Biuve fulvipunctata Baba, 1938 in MolluscaBase eds. (2025 d)] differs in its subquadratic shape and less stalked protoconch. Chelidonura jolkii (Anistratenko in Anistratenko & Prisyazhnyuk, 1996) from the Sarmatian of the Eastern Paratethys Sea (Ukraine) differs in its sharp crest along the adapical margin of the more trigonal wing (see Guzhov 2023: pl. 4). Paleoenvironment. Chelidonura species live on soft, sandy bottoms or on rocky surfaces (Ornales-Gatdula et al. 2011).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFC76705F2BD06CAFF37FECB.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Korytnica Basin: Korytnica (Poland) (Bałuk 2018).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFC86705F2BD067FFAE1FDCF.taxon	type_taxon	Type species. Melanochlamys cylindrica Cheeseman, 1881, by monotypy. Present-day, New Zealand.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFC86705F2BD067FFAE1FDCF.taxon	discussion	Discussion. Melanochlamys has a wide geographic distribution and occurs from temperate to tropical waters where it lives on soft bottoms prying on polychaetes and nemerteans cryptic diversity (Cooke et al. 2014)	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFC86705F2BD057BFF37F9BC.taxon	description	Figs 50 A 1 – A 2	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFC86705F2BD057BFF37F9BC.taxon	materials_examined	Type material. Holotype, MWG UW ZI / 93 / 1252, SL: 2.8 mm, MD: 2.2 mm, Badenian, Korytnica (Poland), illustrated in Bałuk (2018: text-fig. 2 B), Figs 50 A 1 – A 2.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFC86705F2BD057BFF37F9BC.taxon	description	Description. “ Protoconch very small, several microns in cross-section, with oval knob shape. Remaining part of shell composed of 1.5 strongly bent shield, straightening gradually towards front, becoming almost flat and much thinner. Also present wing increasing in size towards the posterior. Distinct thickening, in form of ridge running along posterior margin till end of wing, visible both on inner and outer side. Growth lines visible on external surface; internal surface smooth. ” (Bałuk 2018: 500).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFC86705F2BD057BFF37F9BC.taxon	discussion	Discussion. Only two Aglajidae species have been described from the Paratethys prior to our revision: Chelidonura radwanskii Bałuk, 2018, from the Badenian of the Central Paratethys Sea, and Chelidonura jolkii (Anistratenko in Anistratenko & Prisyazhnyuk, 1996) from the Sarmatian of the Eastern Paratethys Sea (Ukraine). These species were placed by Guzhov (2023) in Chelidonura A. Adams, 1850, whereas Chaban & Martynov (2000) placed the Eastern Paratethyan species in Melanochlamys Cheeseman, 1881. For the Central Paratethys species Chelidonura radwanskii, we recognize that Bałuk (2018) illustrated two specimens representing two different genera. The holotype of Chelidonura radwanskii is a Melanochlamys, characterized by a very prominent crest along the wing, whereas the paratype of Chelidonura radwanskii is indeed that genus. Melanochlamys radwanskii is reminiscent of Melanochlamys cylindrica as discussed by Ortea et al. (2024) but differs in its even more prominent crest and weak concavity adjoining the ridge abapically. Chelidonura jolkii is exceptionally similar to Genus 3 of Ortea et al. (2024: fig. 6), which nests within the Chelidonura clade. Therefore, we follow Guzhov (2023) for the Eastern Paratethyan species. Paleoenvironment. Melanochlamys species live on sandy-muddy bottoms on the littoral zone (Montety et al. 2024).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFC86705F2BD057BFF37F9BC.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Korytnica Basin: Korytnica (Poland) (Bałuk 2018).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFC86705F2BD0015FC8FF820.taxon	type_taxon	Type species. Bulla cranchii Fleming, 1828; by monotypy. Present-day, British Isles.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFC86705F2BD0015FC8FF820.taxon	discussion	Discussion. Some Paratethyan Roxania species have been placed in the Scaphandridae genus Sabatia Bellardi, 1877 in the literature (e. g., Sabatia callifera Boettger, 1907, Sabatia callifera helvetica Berger, 1949). Sabatia, however, has a thick and tuberculated parietal callus, not smooth as in the Paratethyan species. Sabatia species have a more angulated periphery and are not rounded as in Roxania.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFC96707F2BD0786FD5BFC1F.taxon	description	Figs 51 D 1 – D 3	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFC96707F2BD0786FD5BFC1F.taxon	materials_examined	Type material. Holotype, NHMW-ZOO-MO- 75000 - E- 18181 A, SL: 4.9 mm, MD: 3.7 mm, Guntersdorf (Austria), Figs 51 D 1 – D 3.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFC96707F2BD0786FD5BFC1F.taxon	etymology	Etymology. Latin bomba (= bomb), referring to the shape; noun in apposition. Revised description. Shell small, solid, very broadly ovate (SL / MD = 1.4), only last whorl visible. Last whorl strongly globose, apex rounded, narrow (MD / ApD = 3.0), deeply and narrowly umbilicate; low angled apical plane (AP = 5 °). Sides of adapical part slightly conical; periphery convex, placed just above mid-whorl (SL / PH = 1.9), rounded at base. Sculpture of punctate spiral grooves at extremities extending some distance towards mid-whorl, mid-portion smooth. Aperture slightly longer than shell, moderately narrow adapically (MD / AW = 7.1), strongly widening anteriorly (MD / MAW = 2.3), resulting in moderate expansion rate (ER = 3.1). Outer lip extending distinctly above apex adapically, rounded abapically. Columella short, thickened, forming broad, sharply delimited columellar rim bordering narrow umbilical chink. Parietal callus bearing small parietal pad.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFC96707F2BD0786FD5BFC1F.taxon	discussion	Discussion. This species is difficult to characterize. Based on a single specimen, we have no information as to its intraspecific variability. Originally placed in Sabatia, it differs from members of that genus in its sculpture of punctate grooves at the extremities and its evenly and strongly rounded last whorl. Although it has a poorly defined parietal pad, it is not as strong or sharply delimited as in Sabatia species. Therefore, we follow Berger (1953) and place it in Roxania. This makes it a secondary homonym of Roxania helvetica Peyrot, 1932 from the Middle Miocene of France (Peyrot 1932: 197, pl. 13, figs 36, 40 – 42, 55). Therefore, we propose Roxania bomba nov. nom. as new name for this species. Roxania bomba nov. nom. is most similar to Roxania callifera (Boettger, 1907) but differs in its weaker parietal pad and more strongly alate apex of the outer lip. It is possible that it is an extreme form of R. callifera, but until more material becomes available, we prefer to keep them separate. Sabatia isseli Bellardi, 1877, from the Mediterranean Pliocene, is strongly expanded abapically and bears a distinct ridge on the parietal callus rather than a poorly delimited pad (see Landau et al. 2023: pl. 57). The extant Sabatia pustulosa Dall, 1895, from New Caledonia, is also strongly inflated but has an even wider aperture and a very prominent, pustulose parietal callus (see Valdés 2008: fig. 48). Paleoenvironment. Unknown.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFC96707F2BD0786FD5BFC1F.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): North Alpine-Carpathian Foreland Basin: Grund, Guntersdorf (Austria) (Berger 1953).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFCA6706F2BD030AFAA1F80D.taxon	description	Figs 52 A – B	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFCA6706F2BD030AFAA1F80D.taxon	materials_examined	Type material. Lectotype (designated by Zilch 1934: 278), SMF 363162 - 12.2548 a (glass plate 401), SL: 4.8 mm, MD: 2.9 mm, Coşteiu de Sus (Romania), illustrated in Zilch (1934: pl. 22, fig. 20). The specimen was destroyed during WW II. Illustrated material. NHMW 1847 / 0037 / 0131, SL: 4.4 mm, MD: 2.9 mm, Baden (Austria), illustrated in Hörnes (1856: pl. 50, fig 2 a – c), Figs 52 A 1 – A 2. NHMW-ZOO-MO- 75000 - E- 47090, SL: 4.2 mm, MD: 2.6 mm, Bad Vöslau (Austria), Figs 52 B 1 – B 2.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFCA6706F2BD030AFAA1F80D.taxon	description	Revised description. Shell small to medium sized, solid, broad, ovate, only last whorl visible (SL / MD = 1.5 – 1.6). Last whorl globose, periphery placed just above mid-whorl (SL / PH = 1.8). Apex rounded, deeply and narrowly umbilicate, moderately narrow (MD / ApD = 2.7 – 2.8), low angled apical plane (AP = 8 – 9 °). Sculpture of punctate spiral cords at extremities extending a variable distance towards mid-whorl, surface glossy. Aperture entire length of shell, narrow adapically (MD / AW = 9.9 – 11.4), widening anteriorly (MD / MAW = 2.1 – 2.2), with high expansion rate (ER = 4.7 – 5.1). Outer lip extending adapically only slightly above apex, convex below, rounded abapically. Columella short, strongly excavated. Columellar callus forming prominent, thick, sharply delimited rim, adherent to base. Columellar callus passing into prominent expanded parietal callus with characteristically (for genus) swollen parietal pad in adapical third. No umbilicus or chink.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFCA6706F2BD030AFAA1F80D.taxon	discussion	Discussion. Roxania callifera (Boettger, 1907) is based on a rather slender specimen with prominent spiral sculpture, whereas the herein described specimens are comparatively stout and have reduced sculpture mid-whorl. Both bear a low parietal callus pad, which does not extend far over the base and a strongly excavated columella and adherent columellar rim. Roxania callifera differs from Roxania bomba nov. nom. In being slightly less rounded and having the apical tip of the outer lip less alate. As discussed under that species, it is represented by a single specimen, and it is possible that it represents an extreme form of R. callifera. These two species differ from all other Paratethyan congeners in having a parietal pad developed. Roxania callifera differs from Sabatia isseli Bellardi, 1877, from the Mediterranean Pliocene, in its slenderer outline, less expanding aperture and weaker parietal callus (see Landau et al. 2023: pl. 57). In its relatively weak parietal callus, the Paratethyan species is reminiscent of the extant Sabatia nivea (Watson, 1883) from Vanuatu, which is much larger and more globose (see Valdés 2008: fig. 46). Paleoenvironment. The species is documented from deep water and shallow water assemblages. In the Vienna Basin it is most abundant in the clay of the Baden Formation, suggesting outer neritic water depth (Kranner et al. 2021). The occurrence from Cerová (Slovakia) suggests even upper bathyal environments (Harzhauser et al. 2011)	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFCA6706F2BD030AFAA1F80D.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienna Basin: Baden Vöslau, Baden, Steinebrunn (Austria) (Berger 1953); Făget Basin: Lăpugiu de Sus, Coşteiu de Sus (Romania) (Boettger 1907).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFCC6701F2BD0786FAE6FB13.taxon	description	Figs 51 C 1 – C 3	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFCC6701F2BD0786FAE6FB13.taxon	materials_examined	Type material. Holotype, NHMW 2025 / 0152 / 0003, SL: 3.4 mm, MD: 2.3 mm, Bad Vöslau (Austria), Figs 51 C 1 – C 3. Type locality. Bad Vöslau (Austria). Type stratum. Baden Formation. Age. Middle Miocene, middle Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFCC6701F2BD0786FAE6FB13.taxon	etymology	Etymology. Referring to the ovate outline.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFCC6701F2BD0786FAE6FB13.taxon	diagnosis	Diagnosis. Small, thin-shelled, very broadly ovate, narrowly umbilicate shell with sculpture of punctate grooves and relatively wide aperture; straight, narrow, erect columella, delimiting deep, narrow umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFCC6701F2BD0786FAE6FB13.taxon	description	Description. Shell small, thin-shelled, very broadly ovate (SL / MD = 1.4). Only last whorl visible; periphery placed just above mid-whorl (SL / PH = 1.7). Narrow, rounded to weakly conical apex (MD / ApD = 4.2), with weakly oblique apical plane (AP = 7 °); deeply and narrowly umbilicate, delimited by round-edged, elevated crest. Sculpture of weak, punctate spiral grooves at extremities delimiting broad, flat spiral cords. Spiral grooves obsolete mid-whorl, leaving mid-portion smooth. Aperture relatively wide in apical third (MD / AW = 4.7), moderately widening anteriorly (MD / MAW = 2.6), resulting in low expansion rate (ER = 1.8). Outer lip thin, distinctly rising above apex, strongly convex at periphery, rounded abapically. Columella moderately short, nearly straight. Columellar callus forming narrow, detached rim, delimiting narrow, deep umbilicus.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFCC6701F2BD0786FAE6FB13.taxon	discussion	Discussion. Roxania oviformis sp. nov. is reminiscent of Roxania utriculus (Brocchi, 1814) in general shape and sculpture but differs in its more convex periphery and wider adapical part of the aperture resulting in a much lower expansion rate (1.8 versus ~ 4.0 in Roxania utriculus). Roxania pila sp. nov. differs in its more spherical outline, narrower adapical part of the aperture, the adherent and broad columellar callus and the prominent parietal callus. The extant Mediterranean Roxania monterosatoi Dautzenberg & H. Fischer, 1896 is superficially similar but has a broad and adherent columellar callus and its outer lip is only weakly rising above the apex (see Dautzenberg & Fischer 1896: pl. 15, figs 3 – 4). Paleoenvironment. The occurrence in the Baden Formation at Bad Vöslau (Austria) suggests middle to outer neritic environments with up to 210 m water depth (Kranner et al. 2021).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFCC6701F2BD0786FAE6FB13.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Bad Vöslau (Austria) (hoc opus).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFCC6700F2BD0206FEC2FE5F.taxon	description	Figs 51 A – B	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFCC6700F2BD0206FEC2FE5F.taxon	materials_examined	Type material. Holotype, NHMW-ZOO-MO- 75000 - E- 24799 A, SL: 5.0 mm, MD: 3.7 mm, Lăpugiu de Sus (Romania), Figs 51 A 1 – A 3. Paratype, NHMW-ZOO-MO- 75000 - E- 24799, SL: 3.9 mm, MD: 3.0 mm, Lăpugiu de Sus (Romania), Figs 51 B 1 – B 3. Type locality. Lăpugiu de Sus (Romania). Type stratum. Dej Formation. Age. Middle Miocene, middle Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFCC6700F2BD0206FEC2FE5F.taxon	etymology	Etymology. Latin pila, ball. Referring to the ball-like shape; noun in apposition.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFCC6700F2BD0206FEC2FE5F.taxon	diagnosis	Diagnosis. Medium sized, solid, very broad ovate shell, narrow apical umbilicus, with deep, punctate spiral grooves at extremities; broad, thin columellar and parietal callus.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFCC6700F2BD0206FEC2FE5F.taxon	description	Description. Shell medium sized, solid, very broadly ovate (SL / MD = 1.3). Only last whorl visible; periphery placed above mid-whorl (SL / PH = 1.6). Broad, rounded apex, with very narrow, shallow umbilicus (MD / ApD = 2.8 – 2.9); apical plane moderately oblique (AP = 12 – 13 °). Sculpture of prominent, punctate spiral grooves at extremities delimiting broad, flat spiral cords. Spiral grooves extending some distance towards mid-whorl, mid-portion smooth. Aperture moderately narrow in apical third (MS / AW = 7.9 – 9.2), greatly widening anteriorly (MD / MAW = 2.5), resulting in moderate expansion rate (ER = 3.1 – 3.6). Outer lip moderately thin, slightly rising above apex, strongly convex at periphery, rounded abapically. Columella short, strongly excavated in lower half, forming broad, adherent columellar callus, passing into broad, thin parietal callus.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFCC6700F2BD0206FEC2FE5F.taxon	discussion	Discussion. The Paratethyan species is highly reminiscent of the Pliocene to extant Mediterranean-Eastern Atlantic Roxania pinguicula (Seguenza, 1880), which differs in its convex base, which extends down to the basal lip, whereas the base terminates along the columellar callus in Roxania pila sp. nov. (see Seguenza 1880: pl. 16, fig. 7; https: // www. marinespecies. org / aphia. php? p = taxdetails & id = 139485 # images). Paleoenvironment. Unknown.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFCC6700F2BD0206FEC2FE5F.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Făget Basin: Lăpugiu de Sus (Romania) (hoc opus).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFCD6702F2BD06CAFDC9FADB.taxon	description	Figs 53 A – F	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFCD6702F2BD06CAFDC9FADB.taxon	description	R [oxania]. (R [oxania].) utriculus totornata Sacco — Sieber 1958: 164.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFCD6702F2BD06CAFDC9FADB.taxon	materials_examined	Type material. Lectotype, Nr. 4624, Museo Civico di Storia Naturale di Milano, Piacentino (Italy), Pliocene, illustrated in Brocchi (1814: pl. 1, fig. 6). Rossi-Ronchetti (1955) and Pinna & Spezia (1976) listed this specimen as holotype, but Pinna & Spezia (1978: 134) referred also to three paratypes (Nr. 4625, 4626, 4627). Therefore, the specimen illustrated by Brocchi (1814) represents a lectotype (by erroneous definition as holotype by Rossi-Ronchetti 1955) and the other specimens are paralectotypes. Illustrated material. NHMW 2013 / 0078 / 0555, SL: 2.4 mm, MD: 1.35 mm, Bad Vöslau (Austria), Figs 53 A 1 – A 3. NHMW 2013 / 0078 / 0555 a, SL: 2.8 mm, MD: 1.6 mm, Bad Vöslau (Austria), Figs 53 B 1 – B 3. NHMW 1867 / 0019 / 0324, SL: 5.5 mm, MD: 3.3 mm, Coşteiu de Sus (Romania), Figs 53 C 1 – C 2. NHMW 2013 / 0078 / 0555 b, SL: 3.1 mm, MD: 1.8 mm, Bad Vöslau (Austria), Figs 53 D 1 – D 3. NHMW 1867 / 0019 / 0321, SL: 5.5 mm, MD: 2.9 mm, Coşteiu de Sus (Romania), Figs 53 E 1 – E 2. NHMW 1997 z 01781211, SL: 3.4 mm, MD: 2.2 mm, Bad Vöslau (Austria), Figs 53 F. Revised description. Shell small to medium sized, solid, moderately broad to broad, elongate ovate (SL / MD = 1.7 – 1.8), with moderately convex sides and convex base. Protoconch unknown. Only last whorl visible. Periphery placed just above mid-whorl (SL / PH = 1.7 – 1.9). Apex narrow (MD / ApD = 2.8 – 2.9), roundly truncated, slightly oblique (AP = 10 – 11 °), deeply and narrowly umbilicate, delimited by round-edged, elevated crest. Apical umbilical wall with sculpture of spiral cords. Sculpture of prominent narrow, spiral cords separated by prominent, punctate grooves at both extremities, occasionally weakening around mid-whorl. Spiral cords occasionally bifid with weaker, shallow, punctate spiral groove. Aperture as long as shell, moderately narrow in adapical half (MD / AW = 8.4 – 9.3), widening anteriorly (MD / MAW = 2.2 – 2.4), resulting in moderate to high expansion rate (ER = 3.9 – 4.0). Outer lip thin, faintly crenulated along base. Columella short, weakly excavated. Columellar callus forming narrow, relatively thin but distinct rim, delimiting narrow, crescent-shaped umbilicus. No columellar fold.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFCD6702F2BD06CAFDC9FADB.taxon	discussion	Discussion. The Paratethyan specimens differ from typical Pliocene to present-day specimens in their slenderer outline (e. g., Pinna & Spezia 1978; Chirli 2013, Landau et al. 2023). Alongside these slender shells, occasional broader ovate specimens corresponding to typical Roxania utriculus co-occur. Moreover, as discussed by Landau et al. (2023: 407), typical Pliocene to extant specimens have spiral sculpture on the last whorl restricted to the extremities, with the mid-portion of the whorl smooth. However, it is not uncommon for Miocene specimens to have spiral sculpture extending across the entire last whorl. This is also seen in the Miocene Paratethyan series illustrated herein. The completely striate form also occurs, albeit rarely, in the Pliocene, and is usually referred to as Roxania utriculus totornata (Sacco, 1897). Provisionally we keep all types in R. utriculus but would not exclude that the common slender morph represents a distinct species. For further discussions on Roxania utriculus see Landau et al. (2023: 407). Paleoenvironment. Roxania utriculus lives in sandy bottoms in inner neritic to outer neritic environments from 30 – 200 m depth (Gofas et al. 2011; Trigo et al. 2018). Paratethyan occurrences occur in similar settings.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFCD6702F2BD06CAFDC9FADB.taxon	distribution	Distribution in Central Paratethys. Karpatian (Early Miocene): Malé Karpaty Mts.: Cerová (Slovakia) (Harzhauser et al. 2011). Badenian (Middle Miocene): North Alpine-Carpathian Foreland Basin: Braunsdorf, Grund, Guntersdorf (Austria); Vienna Basin: Bad Vöslau, Baden, Baden-Sooss, Steinebrunn (Austria) (Berger 1953); Pannonian Basin: Szob (Hungary) (Bohn-Havas 1973); Southern Pannonian Basin: Jakovac (Bosnia and Herzegovina) (Atanacković 1985); Zapresic-Brijeg (Croatia) (Pavlovsky 1957); Bükk Mountains: Borsodbóta (Hungary) (Csepreghy-Meznerics 1972); Mecsek Mountains: Hidas, Pécsszabolcs, Tekeres, (Hungary) (Csepreghy-Meznerics 1950); Făget Basin: Lăpugiu de Sus, Coşteiu de Sus (Romania) (Boettger 1907); Dacian Basin: Opanets (Bulgaria) (Kojumdgieva in Kojumdgieva & Strachimirov 1960). (Proto-) Mediterranean Sea. Serravallian (Middle Miocene): Karaman Basin: Seyithasan (Turkey) (Landau et al. 2013). Tortonian and Messinian (Late Miocene): Po Basin (Italy) (Sacco 1897; Venzo & Pelosio 1963). Pliocene: entire Mediterranean (Landau et al. 2023). Pleistocene: entire Mediterranean (Cerulli-Irelli 1910; Chirli & Linse 2011). Present-day: entire Mediterranean (Ardovini & Cossignani 2004; Repetto et al. 2005; Cossignani & Ardovini 2011; Gofas et al. 2011). North Sea. Late Burdigalian-Langhian (Early-Middle Miocene): Belgium (Glibert 1952 b), Denmark (Moths & Tu ̈ xen 2008), Germany (Wienrich 2007; Moths et al. 2010), The Netherlands (A. W. Janssen, 1984). Tortonian (Late Miocene): Denmark (Schnetler 2005). Pliocene: Belgium (Marquet 1997, 1998). Present-day: Scandinavia, Shetlands, British Isles (T. E. Thompson 1988).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFCD6702F2BD06CAFDC9FADB.taxon	description	Northeastern Atlantic. Tortonian (Late Miocene): Cacela Basin (Portugal) (Dollfus et al. 1904). Present-day: Bay of Biscay (Trigo et al. 2018), Portugal, Canaries (Hernández et al. 2011), Madeira (Nordsieck & García-Talavera 1979; Segers et al. 2009).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFCF6702F2BD02B7FBC2F9DF.taxon	type_taxon	Type species. Laona zonata A. Adams, 1865; by monotypy. Present-day, Japan.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFCF671DF2BD014AFAAEFF2F.taxon	discussion	Discussion. Berger (1953) illustrated a specimen from Bad Vöslau (Austria) (SL: 3.6, MD: 2.7) which is lost. The drawing corresponds well to shells of the Pliocene to Recent Laona quadrata as discussed by Landau et al. (2023). For description and discussion see Landau et al. (2023: 403). Paleoenvironment. The occurrence in the Baden Formation suggests middle to outer neritic environments with up to 210 m water depth (Kranner et al. 2021).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFCF671DF2BD014AFAAEFF2F.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienna Basin: Bad Vöslau (Austria) (Berger 1953). Pliocene to present-day. North Sea, Atlantic, Mediterranean Sea (see Landau et al. (2023: 404 for details).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD0671DF2BD0683FD03FC73.taxon	type_taxon	Type species. Cosmophiline marinellii Berger, 1949; by monotypy. Middle Miocene, Austria.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD0671DF2BD0683FD03FC73.taxon	diagnosis	Diagnosis. “ Shell very thin, small apex, involute, last whorl strongly expanding (as in Philine Ascanius). Surface with strong spiral cords, which diverge arcuately on the last whorl diverge towards the upper and lower margin and terminate in strong teeth. ” (Berger 1953: 115; translated from German).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD0671DF2BD0683FD03FC73.taxon	discussion	Discussion. Oskars et al. (2015) showed the family Philinidae to be polyphyletic with four lineages warranting familial status (Philinidae, Philinorbidae, Laonidae, and an unnamed clade informally designated by the authors as Philinidae Clade 4). Since then, a new family has been added, namely Antarctophilinidae for Southern Ocean species (Moles et al. 2019). Although the separation between these families relies mostly on anatomical characters such as presence / absence of gizzard plates as well as the nature of these plates, shells are also fairly informative for family and generic assignments. The genus of Cosmophiline is characterized by a very distinct shell which to our best knowledge is not present among extant philinid snails. Based on the quadrangular and flattish shape of these shells we assign the genus to the family Philinidae.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD0671DF2BD04E7FF37F97B.taxon	description	Figs 54 E 1 – E 3	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD0671DF2BD04E7FF37F97B.taxon	materials_examined	Type material. Holotype, NHMW-ZOO-MO- 75000 - E- 4172, SL: 2.6 mm, MD: 1.4 mm, Bad Vöslau (Austria), Figs 54 E 1 – E 3. Revised description. Shell small, fragile, translucent, rectangular (SL / MD = 1.9). Spire covered by last whorl. Last whorl greatly expanded, wing like, with weak rounded angulation corresponding to shoulder and base. Sculpture above and below angulations of narrow, relatively strong spiral cords terminating as long pointed spines at periphery. Between angulations much weaker cords becoming subobsolete in broad mid-portion on last whorl (outer lip edge of mid-portion missing). Columella forming narrow rim in adapical part, thinning below. Aperture extremely wide, auriculate.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD0671DF2BD04E7FF37F97B.taxon	discussion	Discussion. This species is unique and is documented so far only by a single specimen. Paleoenvironment. The occurrence in the Baden Formation at Bad Vöslau (Austria) suggests middle to outer neritic environments with up to 210 m water depth (Kranner et al. 2021).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD0671DF2BD04E7FF37F97B.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienna Basin: Bad Vöslau (Austria) (Berger 1953).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD0671DF2BD01EFFBEFF8B7.taxon	type_taxon	Type species. Bulla scabra O. F. Müller, 1784; by monotypy. Present-day, Norway.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD2671FF2BD0786FECEF976.taxon	description	Figs 54 A – C	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD2671FF2BD0786FECEF976.taxon	materials_examined	Type material. We are not aware of the whereabouts of the type material. Illustrated material. NHMW 2010 / 0004 / 1648, SL: 3.7 mm, MD: 2.3 mm, Bad Vöslau (Austria), Figs 54 A 1 – A 3. NHMW 2010 / 0004 / 1648, SL: 2.1 mm, MD: 1.5 mm, Bad Vöslau (Austria), Figs 54 B 1 – B 2. NHMW-ZOO-MO- 75000 - E- 47001, SL: 3.5 mm, MD: 2.3 mm, Lăpugiu de Sus (Romania), Figs 54 C 1 – C 3.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD2671FF2BD0786FECEF976.taxon	description	Revised description. Shell small, relatively fragile, elongate ovate, with flattened spire (SL / MD = 1.6). Last whorl inflated, ovate, apex roundly truncated (MD / ApD = 2.7), whorl profile convex below, rounded at base. Periphery placed above mid-whorl (SL / PH = 1.5). Sculpture of regular, chain-like spiral grooves; the links small, elongated and connected. Aperture total height of shell, moderately wide in upper quarter (MD / AW = 5.5), greatly widening anteriorly (MD / MAW = 1.2), resulting in high expansion rate (ER = 4.7); outer lip rising slightly above apex to slightly below, convex below, rounded abapically. Columella much reduced, narrow. Parietal callus very narrow.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD2671FF2BD0786FECEF976.taxon	discussion	Discussion. See Landau et al. (2023: 399). Paleoenvironment. The species lives from the littoral zone in its more northern distribution to the bathyal zone in its southern distribution (Gofas et al. 2011). In the Paratethys it is known from shallow and deeper water environments.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD2671FF2BD0786FECEF976.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Vienna Basin: Bad Vöslau (Austria) (Berger 1953); Pannonian Basin: Szob (Hungary) (Strausz 1966); Făget Basin: Lăpugiu de Sus (Romania) (hoc opus). Proto-Mediterranean Sea. Serravallian (Middle Miocene): Karaman Basin: Akpınar, Lale, Seyithasan (Turkey) (Landau et al. 2013). Northeastern Atlantic. Tortonian (Late Miocene): Cacela Basin (Portugal) (Landau et al. 2023). North Sea. (Burdigalian / Langhian (Middle Miocene): the Netherlands (A. W. Janssen 1984). Pliocene to present-day. From North Sea to West Africa and Mediterranean Sea (see Landau et al. 2023: 399 for details).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD2671FF2BD01E0FACBF882.taxon	type_taxon	Type species. Type species. Philine quadripartita Ascanius, 1772; by monotypy. Present-day, Norway.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD3671EF2BD0786FF08F9BE.taxon	description	Figs 54 D 1 – D 3	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD3671EF2BD0786FF08F9BE.taxon	materials_examined	Type material. Syntype, illustrated in Montagu (1803: pl. 7, fig. 7), Bigberry bay at Devon (UK); not in Montagu collection at the Royal Albert Memorial Museum & Art Gallery (RAMM) in Exeter (UK) nor in NHMUK in London (Oliver et al. 2017). Illustrated material. NHMW-ZOO-MO- 75000 - E- 9340, SL: 2.7 mm, MD: 2.0 mm, Enzesfeld (Austria), illustrated in Berger (1953: pl. 18, fig. 82), Figs 54 D 1 – D 3.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD3671EF2BD0786FF08F9BE.taxon	description	Revised description. Shell small, relatively fragile, relatively broad subrectangular, with flattened spire (SL / MD = 1.4). Last whorl strongly inflated, apex roundly truncated (MD / ApD = 2.9), whorl profile convex below, only weakly rounded to nearly straight at base. Periphery above mid-whorl (SL / PH = 1.7). Sculpture of regular, chain-like spiral grooves (poorly preserved in Paratethyan material). Aperture total height of shell, wide in upper quarter (MD / AW = 4.0, greatly widening abapically (MD / MAW = 1.1), resulting in large expansion rate (ER = 3.6); outer lip thin. Columella much reduced, narrow.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD3671EF2BD0786FF08F9BE.taxon	discussion	Discussion. See Landau et al. (2023: 399). Paleoenvironment. Extant Philine catena has a wide bathymetric range from low tide down to 2000 m water depth (Ohnheiser & Malaquias 2013: 280).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD3671EF2BD0786FF08F9BE.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Polish - Ukrainian Fore-Carpathian Basin: Wieliczka (Poland) (Reuss 1867); Vienna Basin: Bad Vöslau, Enzesfeld (Austria) (Berger 1853). Proto-Mediterranean Sea. Serravallian (Middle Miocene): Karaman Basin: Akpınar, Lale, Seyithasan (Turkey) (Landau et al. 2013). Northeastern Atlantic. Tortonian (Late Miocene): Loire Basin (France) (Landau et al. 2020). For Pliocene and Pleistocene occurrences in the North Sea, the northeastern Atlantic and the Mediterranean Sea see Landau et al. (2023). Present-day. From Arctic waters and North Sea down to Madeira and Canary Islands (Nordsieck & García-Talavera 1979; Hernández et al. 2011; Landau et al. 2023) and Mediterranean Sea (Repetto et al. 2005; Gofas et al. 2011).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD3671EF2BD0010FC19F8B2.taxon	type_taxon	Type species. Sabatia isseli Bellardi, 1877; by monotypy. Pliocene, Italy.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD46719F2BD0786FDE0FAAF.taxon	description	Figs 52 C – D	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD46719F2BD0786FDE0FAAF.taxon	materials_examined	Type material. Holotype, NHMW 1863 / 0015 / 1154, SL: 4.2 mm, MD: 3.1 mm, Rudice (Czech Republic), Figs 52 C 1 – C 2. Paratype, NHMW 1863 / 0015 / 1154 a, SL: 5.2 mm, MD: 3.4 mm, Rudice (Czech Republic), Figs 52 D 1 – D 2. Type locality. Rudice (Czech Republic). Type stratum. Middle Miocene clay and silt. Age. Middle Miocene, middle Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD46719F2BD0786FDE0FAAF.taxon	etymology	Etymology. After the protagonist of the French novel The Hunchback of Notre-Dame (1831) by Victor Hugo, referring to the hunched back-like parietal callus.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD46719F2BD0786FDE0FAAF.taxon	diagnosis	Diagnosis. Small to medium sized, ovate shell with broadly conical adapical part bearing deep, punctate spiral grooves, deeply excavated columella and prominent, pustulose, hump-like parietal pad.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD46719F2BD0786FDE0FAAF.taxon	description	Description. Shell small to medium sized, solid, broadly ovate (SL / MD = 1.7 – 1.9), only last whorl visible. Last whorl strongly globose, apex rounded, moderately narrow (MD / ApD = 2.2 – 2.6), deeply and narrowly umbilicate; low angled apical plane (AP = 6 – 7 °). Sides of adapical part conical forming angle of 74 – 78 °; periphery convex, placed just above mid-whorl (SL / PH = 1.7), rounded at base. Sculpture of spiral cords separated by deep, prominent, punctate spiral grooves along conical adapical part of shell. Weaker punctate spiral grooves over base; mid-portion smooth. Aperture as long as shell, moderately narrow adapically (MD / AW = 7.8 – 8.9), strongly widening anteriorly (MD / MAW = 1.7 – 2.0), resulting in high expansion rate (ER = 4.4 – 4.5). Outer lip extending slightly above apex adapically, rounded abapically. Columella short, strongly excavated, strongly thickened, forming broad, sharply delimited adherent columellar rim, continuous with broad, sharply delimited parietal callus bearing prominent, gibbose, finely pustulose parietal pad. No umbilicus or umbilical chink.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD46719F2BD0786FDE0FAAF.taxon	discussion	Discussion. Sabatia quasimodoi sp. nov. differs from the Pliocene Mediterranean Sabatia isseli Bellardi, 1877 in its stronger parietal pad and the spiral grooves at the extremities are stronger with mid-whorl smooth, whereas in Sabatia isseli the spirals are finer and cover the whole last whorl. Sabatia quasimodoi sp. nov. differs from other Paratethyan Roxania species in its deeper spiral grooves on the adapical part of the shell and the very prominent, gibbose and finely pustulose parietal pad, which extends far over the base. In this feature, it is reminiscent of the extant Sabatia pustulosa Dall, 1895, from the Indo-West Pacific, which differs in its auriculate aperture (see Valdés 2008: fig. 48). The extant Sabatia supracancellata (Schepman, 1913) and Sabatia pyriformis Valdés, 2008, both from the Indo-West Pacific, have also a pustulose parietal pads and agree with Sabatia quasimodoi sp. nov. in the prominent adapical sculpture but have wing-like expanding adapical tips of the outer lip. Paleoenvironment. Probably middle to outer neritic based on co-occurring fauna (own data, M. H.).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD46719F2BD0786FDE0FAAF.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): North Alpine-Carpathian Foreland Basin: Rudice (Czech Republic) (hoc opus).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD46719F2BD02DAFDEFF984.taxon	type_taxon	Type species. Bulla lignaria Linnaeus, 1758; original designation, placed on the Official List by Opinion 287 (ICZN 1954 b)]. Present-day, Europe.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD4671BF2BD01B2FD28FDB7.taxon	description	Figs 55 A – D	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD4671BF2BD01B2FD28FDB7.taxon	materials_examined	Type material. Holotype, NHMW 2007 z 0086 / 0000, SL: 19.5 mm, MD: 8.6 mm, Grund (Austria), Figs 55 A 1 – A 3. Paratypes: NHMW 2007 z 0086 / 0000 a, SL: 17.1 mm, MD: 8.0 mm, Grund (Austria), Figs 55 B 1 – B 3. NHMW 2007 z 0086 / 0000 b, SL: 19.0 mm, MD: 9.0 mm, Grund (Austria), Figs 55 C 1 – C 3. NHMW 2007 z 0086 / 0000 c, SL: 15.8 mm, MD: 6.9 mm, Grund (Austria), Figs 55 D 1 – D 3. Type locality. Grund (Austria). Type stratum. Grund Formation. Age. Middle Miocene, early Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD4671BF2BD01B2FD28FDB7.taxon	etymology	Etymology. Superlative of Latin = alatus (= winged), referring to the wing-like extension of the outer lip.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD4671BF2BD01B2FD28FDB7.taxon	diagnosis	Diagnosis. Large, relatively fragile, moderately narrow shell with alate outer lip, forming narrow, sharp, trigonal extension at apex; sculpture of broad spiral cords separated by indistinctly punctate grooves.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD4671BF2BD01B2FD28FDB7.taxon	description	Description. Shell large, relatively fragile, moderately narrow (SL / MD = 2.1 – 2.2), narrow posteriorly, expanding abapically, slightly antero-dorsally compressed. Only last whorl visible. Apex narrow (MD / ApD = 3.6 – 3.9), truncated, apical plane oblique (AP = 11 – 16 °), apical umbilicus sealed with semicircular depression, delimited by sharp-edged, trigonal crest. Periphery placed mid-whorl or just below (SL / PH = 2.1 – 2.3), faintly concave sides above periphery, forming angle of about 40 – 44 °. Rounded at slightly flaring base. Sculpture of widely spaced, relatively regular, indistinctly punctate spiral grooves, delimiting broad, flat spiral cords. Punctuations delicate, coalescent within groove. Spiral cords occasionally bifid, closer set at apex, broader mid-whorl. Aperture slightly longer than shell, moderately wide in adapical third (MD / AW = 4.8 – 5.2), widening greatly anteriorly (MD / MAW = 1.2 – 1.3), resulting in high expansion rate (ER = 4.0 – 4.3). Outer lip distinctly alate apically, extending above apex as narrow, sharp, trigonal extension. Outer lip expanding and almost straight below, broadly rounded abapically. External spiral sculpture shines through. Columella narrow, broadly excavated. Columellar callus forming narrow, thin, adherent rim; parietal callus thin, narrow.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD4671BF2BD01B2FD28FDB7.taxon	discussion	Discussion. Scaphander alatissimus sp. nov. differs from Scaphander grundincola (De Gregorio, 1885) in its smaller size, elongate outline and narrow wing-like adapical expansion of the outer lip. Berger (1953) confused this species with the Early Miocene Scaphander tarbelliana (Grateloup, 1837) [= Scaphander grateloupi (Michelotti, 1847)] which is broader and lacks an alate outer lip (see Lozouet et al. 2001: 81, pl. 37, fig. 6 and references therein). Scaphander alatissimus sp. nov. is reminiscent of the extant Scaphander darius Ev. Marcus & Er. Marcus, 1967 from the Western Atlantic, which differs in its weaker spiral sculpture and smaller adapical extension (see Eilertsen & Malaquias 2012: fig 4 / 6, 4 / 8 – 9). Paleoenvironment. At Grund (Austria) fossiliferous channel fills, which formed in middle to outer neritic environments, bearing allochthonous assemblages uniting coastal-mudflat faunas with inner neritic ones occur (Zuschin et al. 2005; Roetzel 2009). Preservation of the fossil shells suggests that the shells might have belonged to the deeper water fauna.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD4671BF2BD01B2FD28FDB7.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): North Alpine-Carpathian Foreland Basin: Grund, Guntersdorf (Austria) (Berger 1953; hoc opus).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD66715F2BD05A3FA87FC57.taxon	description	Figs 56 A – C	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD66715F2BD05A3FA87FC57.taxon	materials_examined	Type material. Holotype, NHMW 1851 / 0002 / 0080, SL: 54.5 mm, MD: 28.9 mm, Grund (Austria), illustrated in Hörnes (1856: pl. 50, figs 1 a – b), Figs 56 A 1 – A 2. Illustrated material. NHMW 1855 / 0045 / 0795, SL: 52.8 mm, MD: 28.8 mm, Grund (Austria), Figs 56 B 1 – B 3. NHMW 1997 z 0178 / 1216, SL: 45.8 mm, MD: 25.3 mm, Bad Vöslau (Austria), Figs 56 C 1 – C 3. Additional material. NHMW 1870 / 0037 / 0061, Ritzing (Austria); 4 spec., NHMW 1874 / 0024 / 0068, Ritzing (Austria); 5 spec., NHMW 1848 / 0003 / 0036, Ritzing (Austria); 5 spec. NHMW 1878 / 0041 / 0024, Vienna / Pötzleinsdorf (Austria); 1 spec., NHMW 1851 / 0026 / 0077, Vienna / Pötzleinsdorf (Austria); 2 spec., NHMW 1861 / 0001 / 0285, Pöls (Austria).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD66715F2BD05A3FA87FC57.taxon	description	Revised description. Shell very large, solid, moderately broad (SL / MD = 1.8 – 1.9), narrow posteriorly, expanding abapically, slightly antero-dorsally compressed. Only last whorl visible. Apex narrow (MD / ApD = 3.8 – 4.1), slightly truncated, apical plane strongly oblique (AP = 20 – 23 °), apical umbilicus almost sealed, reduced to semicircular slit, delimited by sharp-edged, trigonal crest. Periphery placed just below mid-whorl (SL / PH = 2.2), weakly concave sides above periphery, forming angle of about 42 – 45 °. Rounded at slightly flaring base. Sculpture of widely spaced, relatively regular, very finely punctate, sharply delimited spiral grooves, delimiting broad, flat spiral cords. Spiral cords not bifid. Spiral grooves occasionally somewhat irregular at intersections with growth lines. Punctuations not discernible as individual pits. Spiral cords closer set adapically, becoming wider spaced towards periphery. Aperture slightly longer than shell, moderately wide in anterior third (MD / AW = 5.3 – 5.4), widening greatly below (MD / MAW = 1.3 – 1.4), resulting in high expansion rate (ER = 3.9 – 4.1). Outer lip slightly alate apically, expanding and almost straight below, broadly rounded abapically. External spiral sculpture not shining through. Basal lip slightly thickened. Columella broadly excavated. Columellar callus forming thick, broad, sharply delimited, adherent rim continuous with parietal callus of similar width and strength.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD66715F2BD05A3FA87FC57.taxon	discussion	Discussion. This is the largest and most widespread Scaphander species in the Paratethys Sea, which is characterized by its very thick shell and broad and thick columellar callus and parietal callus. Already De Gregorio (1885 b) separated it from Scaphander lignarius as new variety grundicola based on the specimen illustrated by Hörnes (1856) as “ Bulla lignaria ”. This name was overlooked by all subsequent authors. Soon later, Sacco (1897) introduced Scaphander lignarius dertonensis for specimens from the Tortonian of San Agata (Italy) [lectotype designated herein: BS. 088.01.009, SL: 31 mm, MD: 16 mm, illustrated in Sacco (1897: pl. 3, figs 102 – 103]. We treat Scaphander dertonensis Sacco, 1897 as junior synonym of Scaphander grundicola (De Gregorio, 1885). Literature records are difficult to evaluate as most Paratethyan records have been erroneously lumped in ‘ Scaphander lignarius’. Some of these records may in fact represent small sized Scaphander species. See Landau et al. (2013: 335) for additional discussion. Paleoenvironment. Known from a broad range of shallow marine and deeper marine settings.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD66715F2BD05A3FA87FC57.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Polish-Ukrainian Fore-Carpathian Basin: Golubytsya (Hołubica) (Ukraine) (Friedberg 1928); Nowy Sącz Basin: Niskowa (Poland) (Bałuk 1970); North Alpine-Carpathian Foreland Basin: Hrušovany nad Jevišovkou (Czech Republic), Grund (Austria) (Berger 1953); Vienna Basin: Bad Vöslau, Gainfarn, Vienna / Kalksburg, Vienna / Pötzleinsdorf (Austria) (Berger 1953); Oberpullendorf Basin: Ritzing (Austria) (Berger 1953); Styrian Basin: Pöls (Austria) (hoc opus); Pannonian Basin: Nógrádszakál, Várpalota (Hungary) (Bogsch 1936; Strausz 1966); Cserhát Hills: Mátraverebély (Hungary) (Strausz 1966); Făget Basin: Buituri (Romania) (Moisescu 1955); Şimleu Basin: Tusa (Romania) (Nicorici 1972); Caransebeş Basin: Deline ş ti, Rugi (Romania) (Stancu & Andreescu 1968); Dubova Region (Romania) (Stancu et al. 1971).? Dacian Basin: Urovene (Bulgaria) (Kojumdgieva in Kojumdgieva & Strachimirov 1960). Proto-Mediterranean Sea. Serravallian (Middle Miocene): Karaman Basin: Seyithasan (Turkey) (Landau et al. 2013). Tortonian (Late Miocene): Po Basin: S. Agata (Italy) (Sacco 1897). Northeastern Atlantic. Langhian (Middle Miocene): Loire Basin: Manthelan (France) (Glibert 1952 a).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD86714F2BD04C3FEEEFE97.taxon	description	Figs 56 D 1 – D 2	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD86714F2BD04C3FEEEFE97.taxon	materials_examined	Type material. Lectotype (designated herein), specimen illustrated in Peyrot (1932: pl. 12, figs 25 – 26), Serravallian (Middle Miocene), Salles (France). Illustrated material. NHMW 2000 z 0001 / 0053, SL: 46.0 mm, MD: 28.0 mm, Niederkreuzstetten (Austria) (illustrated in Harzhauser 2002: pl. 11, fig. 17), Figs 56 D 1 – D 2.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD86714F2BD04C3FEEEFE97.taxon	description	Revised description. Shell very large, solid, broad (SL / MD = 1.6), narrow posteriorly, strongly expanding abapically, slightly antero-dorsally compressed. Only last whorl visible. Apex narrow (MD / ApD = 4.6), slightly truncated, apical plane strongly oblique (AP = 18 °), apical umbilicus fully sealed, delimited by cord-like, narrowly rounded crest. Periphery placed mid-whorl (SL / PH = 2.2), weakly concave sides below apex, forming angle of about 55 °. Rounded at strongly flaring base. Sculpture of close-set, relatively regular, indistinctly punctate, spiral grooves, delimiting broad, flat spiral cords. Spiral grooves occasionally somewhat irregular at intersections with growth lines. Punctuations not discernible as individual pits. Spiral cords even closer-set at apex. Aperture slightly longer than shell, moderately wide in adapical third (MD / AW = 5.8), widening greatly anteriorly (MD / MAW = 1.4), resulting in high expansion rate (ER = 4.1). Outer lip slightly alate and pointed apically, expanding and almost straight below, broadly rounded abapically. Basal lip solid, Columella deeply and broadly excavated. Columellar callus forming thick, broad, sharply delimited, adherent rim continuing into parietal callus of similar width and strength.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD86714F2BD04C3FEEEFE97.taxon	discussion	Discussion. Scaphander helveticus Peyrot, 1932 differs from Scaphander grundincola (De Gregorio, 1885) in its very broad, drop-shaped outline with faintly concave sides below the apex, more crowded spiral cords, and the thickened apical margin bordering the apical umbilicus. The type specimen from Salles (France) differs from the Paratethyan shell only in its slightly broader spiral cords. Specimens from the Atlantic Upper Miocene of Cacela illustrated by Dollfus et al. (1903: 22, pl. 36, figs 13 – 14) as Scaphander lignarius were discussed by Landau et al. (2013: 335). They are not that species, neither are they conspecific with any of the species discussed herein. They probably represent an undescribed species that will be discussed in a monograph describing the gastropod assemblage of the locality (Landau et al., in prep). Paleoenvironment. Shallow marine, lagoonal (Harzhauser et al. 2002).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD86714F2BD04C3FEEEFE97.taxon	distribution	Distribution in Central Paratethys. Karpatian (Early Miocene): Korneuburg Basin: Niederkreuzstetten (Austria) (hoc opus). Northeastern Atlantic. Serravallian (Middle Miocene): Aquitaine Basin: Salles, Sallespisse, Manciet (France) (Peyrot 1932).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD96714F2BD0683FB2CF91D.taxon	description	Figs 57 D 1 – D 3	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD96714F2BD0683FB2CF91D.taxon	materials_examined	Type material. Holotype, NHMW 1997 z 0178 / 1217, SL: 12.7 mm, MD: 3.3 mm, Lăpugiu de Sus (Romania), Figs 57 D 1 – D 3. Type locality. Lăpugiu de Sus (Romania), Făget Basin. Type stratum. Silt and clay of the Dej Formation. Age. Middle Miocene, middle Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD96714F2BD0683FB2CF91D.taxon	etymology	Etymology. Latin catena (= chain), referring to the sculpture in the spiral grooves; noun in apposition.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD96714F2BD0683FB2CF91D.taxon	diagnosis	Diagnosis. Small, solid, moderately broad shell with prominent, deep, relatively close-spaced spiral grooves with chain-like punctuation.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD96714F2BD0683FB2CF91D.taxon	description	Description. Shell small, medium thickness, moderately broad (SL / MD = 1.8), narrow posteriorly, expanding abapically, slightly antero-dorsally compressed. Only last whorl visible. Apex narrow (MD / ApD = 3.4), slightly truncated, apical plane slightly oblique (AP = 10 °), apical umbilicus nearly sealed, reduced to semicircular slit, delimited by sharp-edged, trigonal crest. Periphery placed mid-whorl (SL / PH = 1.8), weakly concave sides above periphery, forming angle of about 48 °. Rounded at slightly flaring base. Sculpture of relatively close-spaced, irregular, flat spiral cords, separated by relatively broad, punctate spiral grooves; punctations within grooves subcircular with weak crests, in apposition, forming chain-like sculpture within grooves. Spiral cords close-set at apex, becoming slightly wider below periphery, occasionally weakly bifid. Aperture as long as shell, relatively wide in adapical third (MD / AW = 3.9), widening greatly anteriorly (MD / MAW = 1.4), resulting in relatively low expansion rate (ER = 2.7). Outer lip only slightly rising above apex, expanding and almost straight below, broadly rounded abapically. Columella broadly excavated. Columellar callus forming narrow rim, delimiting narrow umbilical slit. Parietal region covered by sediment.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD96714F2BD0683FB2CF91D.taxon	discussion	Discussion. The specimen herein described as Scaphander catena sp. nov. was mixed with shells of Scaphander grundincola (De Gregorio, 1885) in the NHMW collection, but differs from its congeners in its smaller size, much weaker columellar callus and chain-like spiral sculpture in the deeper and broader spiral grooves. It is quite possible that at 12.7 mm height it is a juvenile specimen and the columellar callus has still to mature. However, its sculpture of irregular cords separated by grooves with chain like punctuations differs from all its congeners; at similar size the punctuations are much coarser. We therefore consider formal description warranted despite the likely subadult nature of the holotype. We are not aware of similar species in the Circum-Mediterranean Neogene. Paleoenvironment. Unknown.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD96714F2BD0683FB2CF91D.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene):? Southern Pannonian Basin: Miljevići (Bosnia and Herzegovina) (Atanacković 1985); Făget Basin: Lăpugiu de Sus (Romania) (hoc opus).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD96716F2BD000CFEC2FDCF.taxon	description	Figs 57 C 1 – C 3	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD96716F2BD000CFEC2FDCF.taxon	materials_examined	Type material. Holotype, NHMW 1860 / 0004 / 0375, SL: 9.6 mm, MD: 2.6 mm, Lăpugiu de Sus (Romania), Figs 57 C 1 – C 3. Type locality. Lăpugiu de Sus (Romania), Făget Basin. Type stratum. Silt and clay of the Dej Formation. Age. Middle Miocene, middle Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD96716F2BD000CFEC2FDCF.taxon	etymology	Etymology. In honor of Wout Krijgsman (University Utrecht, the Netherlands), in recognitions in its contributions to the stratigraphy of the Circum-Mediterraean Neogene.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD96716F2BD000CFEC2FDCF.taxon	diagnosis	Diagnosis. Small, fragile, moderately broad shell with narrowly umbilicate apex and sculpture of moderatelyclose-set, relatively regular, punctate spiral grooves, crossed by arcuate growth lines resulting in delicate, subquadratic pattern in apical part of shell.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD96716F2BD000CFEC2FDCF.taxon	description	Description. Shell small, fragile, moderately broad (SL / MD = 1.9), ovate, moderately narrow posteriorly, expanding abapically, slightly antero-dorsally compressed. Only last whorl visible. Apex narrow (MD / ApD = 3.3), apical plane slightly oblique (AP = 5 °), very narrowly umbilicate, delimited by round-edged crest. Periphery placed just above mid-whorl (SL / PH = 1.9), weakly concave sides above periphery, forming angle of about 53 °. Rounded at slightly flaring base. Sculpture of moderately close-set, relatively regular, spiral cords, separated by narrow grooves made up of very fine, close-set, punctuations in apposition in apical region, becoming less prominent below. Spiral cords occasionally bifid in apical region. Spiral sculpture crossed by arcuate growth lines resulting in delicate, cancellate pattern in apical region. Aperture slightly longer than shell, moderately wide in adapical third (MD / AW = 5.3), widening greatly anteriorly (MD / MAW = 1.4), resulting in moderate expansion rate (ER = 3.9). Outer lip slightly extending above apex, forming small convex extension. Outer lip expanding and almost straight below, broadly rounded abapically. External spiral sculpture shines through. Columella narrow, broadly excavated. Columellar callus forming moderately narrow, sharply delimited, adherent rim, continuous with thin, narrow, parietal callus.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD96716F2BD000CFEC2FDCF.taxon	discussion	Discussion. Like the rest of the species figured on Fig. 57 (hoc opus), it is small for a Scaphander species. It is possible that it is also a juvenile, although the columellar and parietal callus region is relatively well developed. The sculpture of relatively regular spiral cords separated by extremely fine pitted grooves interrupted by relatively strong growth lines resulting in vaguely cancellate sculpture in the apical region is peculiar and distinguishes Scaphander krijgsmani sp. nov. from the slightly similar Scaphander alatissimus sp. nov. Scaphander grundincola (De Gregorio, 1885) differs in its much larger size and does not develop axial sculpture even on subadult shells. Scaphander catena sp. nov. is reminiscent of the extant Scaphander japonicus A. Adams, 1862 in general shape and prominent spiral grooves (see Valdés 2008: fig. 38). Paleoenvironment. Unknown.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFD96716F2BD000CFEC2FDCF.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Făget Basin: Lăpugiu de Sus (Romania) (hoc opus).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFDB6716F2BD057BFE8BF820.taxon	description	Figs 58 A 1 – A 2	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFDB6716F2BD057BFE8BF820.taxon	materials_examined	Type material. Holotype, NHMW 1836 / 0012 / 6576 a, SL: 14.3 mm, MD: 7.3 mm, Hrušovany nad Jevišovkou (Czech Republic), Figs 58 A 1 – A 2. Type locality. Hrušovany nad Jevišovkou (Czech Republic). Type stratum. Middle Miocene silt and clay. Age. Middle Miocene, middle Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFDB6716F2BD057BFE8BF820.taxon	etymology	Etymology. In honor of Wolfgang E. Schollnberger (Potomac, MD, USA), in recognition of his contributions to Alpine geology.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFDB6716F2BD057BFE8BF820.taxon	diagnosis	Diagnosis. Small, moderately solid, ovate shell with sealed apical umbilicus, weak apical depression; sculpture of numerous, close-set spiral grooves separating spiral cords of equal width.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFDB6716F2BD057BFE8BF820.taxon	description	Description. Shell small, moderately solid, moderately broad, ovate (SL / MD = 1.9), moderately narrow posteriorly, slowly expanding abapically, not antero-dorsally compressed. Only last whorl visible. Apex narrow (MD / ApD = 2.9), slightly truncated, apical plane slightly oblique (AP = 10 °), apical umbilicus fully sealed with shallowly concave depression. Periphery placed mid-whorl (SL / PH = 2.0), weakly concave sides above periphery, forming angle of about 31 °. Rounded at slightly flaring base. Sculpture on base of numerous, crowded spiral cords separated by punctate spiral grooves of similar width to cords. Punctations within grooves fused. Aperture as long as shell, moderately wide in adapical third (MD / AW = 6.0), widening greatly anteriorly (MD / MAW = 1.6), resulting in moderate expansion rate (ER = 3.7). Outer lip not rising above apex, expanding and almost straight below, broadly rounded abapically. Columella broadly excavated. Columellar callus forming moderate width, sharply delimited rim, narrowing adapically. No parietal callus.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFDB6716F2BD057BFE8BF820.taxon	discussion	Discussion. Unlike the small Scaphander species figured on Fig. 57 (hoc opus), this specimen is relatively thick shelled and the columellar callus is well developed, which suggests it represents an almost / fully adult shell. Scaphander schollnbergeri sp. nov. is characterized by its ovate outline, fully sealed umbilicus and dense spiral sculpture, which strongly weakens adapically. It is also unusual in that it is not dorso-ventrally compressed. Scaphander parvulinus Sacco, 1897, from the Tortonian and Pliocene of the Po Basin (Italy), might be a related species of comparable size (SL: 12 mm), which differs in its wider adapical aperture, wider maximum aperture width and larger expansion rate (see Sacco 1897: pl. 3, figs 100 – 101). Two specimens from Borsodbóta (Hungary) illustrated by Csepreghy-Meznerics (1972: pl. 18, figs 16 – 17) might be conspecific with Scaphander schollnbergeri sp. nov. but differ in their slightly more elongate outline and more prominent spiral sculpture. Their ovate outline and only weakly expanding base exclude a relation with Scaphander grateloupi (Michelotti, 1847) [= Scaphander tarbelliana (Grateloup, 1837)] as suggested by Csepreghy-Meznerics (1972). Paleoenvironment. Unknown.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFDB6716F2BD057BFE8BF820.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): North Alpine-Carpathian Foreland Basin: Hrušovany nad Jevišovkou (Czech Republic) (hoc opus);? Bükk Mountains: Borsodbóta (Hungary) (Csepreghy-Meznerics 1972).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFDC6710F2BD011AFAEAFC73.taxon	description	Figs 57 A – B	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFDC6710F2BD011AFAEAFC73.taxon	materials_examined	Type material. Holotype, NHMW 1866 / 0050 / 0227, SL. 10.4 mm, MD: 5.4 mm, Baden Sooss (Austria), Figs 57 A 1 – A 3. Paratype, NHMW 1836 / 0012 / 6576, SL: 10.5 mm, MD: 6.0 mm, Hrušovany nad Jevišovkou (Czech Republic), Figs 57 B 1 – B 3. Type locality. Baden Sooss (Austria). Type stratum. Baden Formation. Age. Middle Miocene, middle Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFDC6710F2BD011AFAEAFC73.taxon	etymology	Etymology. Latin perforatus (perforated), referring to the peculiar sculpture.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFDC6710F2BD011AFAEAFC73.taxon	diagnosis	Diagnosis. Small, relatively fragile, elongate ovate shell with narrowly umbilicate apex; sculpture of spiral cords separated by rows of prominent punctuations that mid-whorl are not touching each other, on apex and periphery, grooves become deeper and punctuations in apposition.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFDC6710F2BD011AFAEAFC73.taxon	description	Description. Shell small, relatively fragile, broad to moderately narrow (SL / MD = 1.8 – 2.0), elongate ovate, moderately narrow posteriorly, weakly expanding abapically, slightly antero-dorsally compressed. Only last whorl visible. Apex narrow (MD / ApD = 3.4 – 3.5), apical plane slightly oblique (AP = 8 – 9 °), narrowly umbilicate, delimited by narrow, round-edged crest. Periphery placed just above mid-whorl (SL / PH = 1.8 – 1.9), weakly concave sides above periphery, forming angle of about 61 – 66 °. Rounded at slightly flaring base. Sculpture of widely spaced, irregular, spiral cords separated by rows of prominent punctuations that mid-whorl are not touching each other, on apex and periphery, grooves become deeper and punctuations in apposition. Pits subcircular in apical region, becoming subrectangular along periphery, apex and mid-portion perforations not placed in grooves, on base within grooves that widen and deepen abapically. Spiral cords bifid only on anterior part of base. Aperture as long as shell, moderately wide in adapical third (MD / AW = 3.4 – 3.5), widening greatly anteriorly (MD / MAW = 1.5 – 1.6), resulting in moderate to high expansion rate (ER = 3.7 – 4.2). Outer lip not extending above apex, expanding and almost straight below, broadly rounded abapically. External spiral sculpture shines through. Columella narrow, broadly excavated. Columellar callus forming thin, narrow, adherent rim; parietal callus thin, narrow.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFDC6710F2BD011AFAEAFC73.taxon	discussion	Discussion. Like the new species described above, at 10.4 mm height it is quite possible that the specimen represents a juvenile with the shell and columellar / parietal callus not fully thickened. However, with its “ Roxania - like ” sculpture in which the perforations are widely spaced and not set in grooves, apart from over the base, it differs from all other Circum-Mediterranean Neogene Scaphander species. The extant Scaphander mundus Watson, 1883, from New Caledonia, develops vaguely similar sculpture of closer-set punctations but differs distinctly in its flaring adapical tip of the outer lip (see Valdés 2008: fig. 42). Paleoenvironment. The occurrence in the Baden Formation at Bad Vöslau (Austria) suggests middle to outer neritic environments with up to 210 m water depth (Kranner et al. 2021).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFDC6710F2BD011AFAEAFC73.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): North Alpine-Carpathian Foreland Basin: Hrušovany nad Jevišovkou (Czech Republic) (hoc opus); Vienna Basin: Baden-Sooss (Austria) (hoc opus).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFDD6710F2BD032FFB2EF94F.taxon	type_taxon	Type species. Tornatina (Pseudavena) tauroglandula Sacco, 1897; original designation by Sacco (1897: 39). Early or Middle Miocene, Italy.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFDD6710F2BD032FFB2EF94F.taxon	discussion	Discussion. Pseudavena was established by Sacco (1897) as subgenus of Tornatina A. Adams, 1850 [= Acteocina J. E. Gray, 1847]. His diagnosis was very brief: “ It includes forms similar to Tornatina, but with the spire hidden by the last whorl of the spire, with a transversely striated surface and longitudinally plicate in the upper part. ” (Sacco 1897: 39, translated from Italian). Pseudavena is characterized by slender to moderately broad subcylindrical shells with dome-shaped apex in which the first teleoconch whorl is covered partly by the subsequent whorl and by a peculiar spiral sculpture, which is most prominent along the shoulder and on the base. Utriculus spatha Watson, 1883, from the Caribbean Sea, might be a living representative of this genus (see Pilsbry 1895: 218, pl. 25, figs 53 – 55). Some extant species placed in Cylichnium Dall, 1908, type species Utriculus domitus Dall, 1889; present-day, Caribbean Sea, are reminiscent of Pseudavena. Cylichnium ancillarioides (Schepman, 1913), from Fiji, and Cylichnium nanum Valdés, 2008, from Vanatu, are comparable in outline and especially Cylichnium nanum is reminiscent of Pseudavena due to its spiral sculpture along the shoulder and the prominent columellar fold. In Cylichnium species, the last whorl covers the spire completely (see Valdés 2008: figs 55 A – E, G – H).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFDD6713F2BD01FBFAC3FAE7.taxon	description	Figs 58 B – D	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFDD6713F2BD01FBFAC3FAE7.taxon	materials_examined	Type material. Holotype, NHMW 1863 / 0015 / 1155, SL: 8.0 mm, MD: 3.3 mm, Rudice (Czech Republic), Figs 58 B 1 – B 3. Paratypes, NHMW 1863 / 0015 / 1155 a, SL: 6.2 mm, MD: 2.8 mm, Rudice (Czech Republic), Figs 58 C 1 – C 3; NHMW 2013 / 0300 / 0556, SL: 8.5 mm, MD: 3.1 mm, Baden-Sooss (Austria), Figs 58 D 1 – D 2. Additional paratypes. 8 spec., NHMW 2013 / 0300 / 0556 a, Baden-Sooss (Austria), 3 spec., NHMW 1863 / 0015 / 1155 b, Rudice (Czech Republic). Type locality. Rudice (Czech Republic). Type stratum. Middle Miocene clay and silt. Age. Middle Miocene, middle Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFDD6713F2BD01FBFAC3FAE7.taxon	etymology	Etymology. Latin bellus meaning beautiful; referring to the beautiful appearance.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFDD6713F2BD01FBFAC3FAE7.taxon	diagnosis	Diagnosis. Medium sized, solid, subcylindrical shell with prominent spiral cords on subsutural ramp and along shoulder; prominent, flattened columella with sharp, low adapical columellar fold and weaker second fold below.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFDD6713F2BD01FBFAC3FAE7.taxon	description	Description. Shell medium sized, solid, subcylindrical, with slowly contracting base, moderately slender (SL / MD = 2.2 – 2.4). Protoconch unknown. Teleoconch of about three whorls separated by narrowly canaliculated suture. Apex low dome-shaped, broad (MD / ApD = 1.3 – 1.5). First whorl sunken and largely covered by subsequent whorl without forming umbilicus. Spire whorls low with subsutural collar and weakly concave subsutural ramp, apical angle 92 – 95 °. Last whorl ~ 95 % of total height with slightly concave subsutural ramp, rounded shoulder, faintly convex periphery in adapical third, slowly contracting below. Periphery placed above mid-whorl (SL / PH = 1.4 – 1.7). Sculpture of prominent narrow, spiral cords separated by narrow grooves on subsutural ramp and below shoulder; weak to subobsolete at periphery; broader spiral cords and grooves over base; delicate growth lines. Aperture ~ 87 % of total height, moderately narrow in uppermost part (MD / AW = 6.0 – 6.4), evenly widening anteriorly (MD / MAW = 2.4 – 2.7) with low expansion rate (ER = 2.4 – 2.6); outer lip simple, straight in upper two-thirds, rounded at base. Columella short, weakly excavated, strongly thickened, markedly flattened, expanded over base, sharply delimited from base, with sharp, low, adapical columellar fold and weaker second fold below. Parietal callus not present.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFDD6713F2BD01FBFAC3FAE7.taxon	discussion	Discussion. This species is characterized by its spire morphology and peculiar sculpture along the shoulder. These features place it in Pseudavena Sacco, 1897, which was known so far only from the Mediterranean region. There it is documented by two species and several varieties, which will need revision. Pseudavena tauroglandula (Sacco, 1897), from the Burdigalian or Langhian of the Colli Torinesi (Italy), differs in its convex spire and the weaker and narrower columella (see Sacco 1897: pl. 3, figs 66 – 69; Ferrero Mortara et al. 1984: pl. 52, fig. 7). Pseudavena coclis (Ponzi, 1876), from the Mediterranean Pliocene, has a similar spire with weakly concave whorls and prominent sculpture along the shoulder, but differs in its much broader shell and weaker columella (see Sacco 1897: pl. 3, figs 70 – 76; Cavallo & Repetto 1992: fig. 472; Chirli 2013: pl. 12, figs 13 – 17). Paleoenvironment. The occurrence in the Baden Formation at Bad Vöslau (Austria) suggests middle to outer neritic environments with up to 210 m water depth (Kranner et al. 2021). Pseudavena tauroglandula (Sacco, 1897) and Pseudavena coclis (Ponzi, 1876) were also associated with muddy bottoms according to Sacco (1897: 39)	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFDD6713F2BD01FBFAC3FAE7.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): North Alpine-Carpathian Foreland Basin: Rudice (Czech Republic) (hoc opus). Vienna Basin: Bad Vöslau, Baden-Sooss (Austria) (Berger 1953).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFDE6713F2BD02DBFDDBF897.taxon	materials_examined	Material NHMW-ZOO-MO- 75000 - E- 18179, SL: 11.1 mm, MD: 6.1 mm, Guntersdorf (Austria).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFDE6713F2BD02DBFDDBF897.taxon	discussion	Discussion. Berger (1953) identified a single specimen as Acera [= Akera] [type species Akera bullata Müller, 1776; present-day, Denmark]. The specimen is still stored in the NHMW collection. The solid shell, the angled periphery and the absence of the characteristic channelled suture exclude this placement. Instead, the specimen represents a juvenile Cypraeidae.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFDE6712F2BD0082FE94FE73.taxon	discussion	Discussion. The specimens illustrated by Strausz (1954, 1966) are reminiscent of the extant Amathinidae species Leucotina casta (A. Adams, 1853) and Leucotina natalensis E. A. Smith, 1910 and are excluded herein from Acteonidae.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFDE6712F2BD0082FE94FE73.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Pannonian Basin: Várpalota (Hungary) (Strausz 1966).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFDF6712F2BD06E6FE7AFC87.taxon	discussion	Discussion. Seneš (1955) based this species on poorly preserved material from the Badenian of Slovakia. The name is preoccupied by Grateloup (1838) but as the shells represent juvenile Ringiculidae, no replacement name is needed.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFDF6712F2BD06E6FE7AFC87.taxon	distribution	Distribution in Central Paratethys. Late Badenian (Middle Miocene): Eastern Slovakian Basin: Sol’ (Slovakia) (Seneš 1955).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFDF6712F2BD04B3FE7AFB62.taxon	discussion	Discussion. The shells represent juvenile Ringiculidae, which are very abundant at the locality.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFDF6712F2BD04B3FE7AFB62.taxon	distribution	Distribution in Central Paratethys. Late Badenian (Middle Miocene): Eastern Slovakian Basin: Sol’ (Slovakia) (Seneš 1955).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFDF6712F2BD0214FF37F84E.taxon	discussion	Discussion. Papp (1954) illustrated a small Sarmatian shell (SL: 3.8, MD. 2.0 mm), assuming that it represented a species described by Handmann (1882: 564) as Conovulus (?) vindobonensis. The holotype of Conovulus (?) vindobonensis from the Sarmatian of Bad Vöslau (Austria) was stored in the Kollegium Kalksburg in Vienna, a Catholic private school, but is lost. Handmann (1883) described a small (SL: 2 mm, MD: 1 mm), smooth, inflated shell of five whorls with a thick outer lip and small columellar denticle. It was established in the ellobiid genus Conovulus Lamarck, 1816 [= Melampus Montfort, 1810] and might represent a Pyramidellidae. In any case the specimen illustrated by Papp (1954) differs considerably in its prominent spiral sculpture of deep, punctate grooves. This morphology is reminiscent of the Amathinidae genus Leucotina A. Adams, 1860 [type species Leucotina niphonensis A. Adams, 1860; present-day, Japan] and therefore we exclude it from Acteonidae.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FFDF6712F2BD0214FF37F84E.taxon	distribution	Distribution in Central Paratethys. Sarmatian (Middle Miocene): Vienna Basin: Hölles (Austria) (Papp 1954).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF2067EDF2BD0786FD3BFCF2.taxon	materials_examined	Type material. The holotype was stored in the Kollegium Kalksburg in Vienna, a Catholic private school, but is lost; SL: 5 mm, MD: 3 mm; Hölles (Austria).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF2067EDF2BD0786FD3BFCF2.taxon	discussion	Discussion. Handmann (1883) described a thin shell of five whorls with globose last whorl, attaining more than half of the total height, sculpture of broader and narrower, punctate spiral grooves. Columella with small, oblique fold; columellar callus forming narrow, adherent rim delimiting narrow umbilical chink. Fordinál & Zlinská (1998) illustrated a small shell, which corresponds largely to the description of Handmann (1883) (SL: 2.4 mm). It lacks a clear columellar fold, which may be due to its subadult growth stage. We tentatively place this dubious species in the Amathinidae genus Adelactaeon Cossmann, 1895 [type species Myonia japonica A. Adams, 1860; present-day, Japan]	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF2067EDF2BD0786FD3BFCF2.taxon	distribution	Distribution in Central Paratethys. Sarmatian (Middle Miocene): Vienna Basin: Hölles (Austria) (Handmann 1883); Skalica (Slovakia) (Fordinál & Zlinská 1998).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF2067EDF2BD0464FDC1FAD0.taxon	discussion	Discussion. The illustration shows a globose shell with prominent spiral grooves and deeply excavated columella. Eichwald (1853) compared it with Roxania utriculus (Brocchi, 1814). But the apertural features are rather reminiscent of the Pliocene Mediterranean Bulla subampulla d’Orbigny, 1852. We consider this a species inquirenda.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF2067EDF2BD0464FDC1FAD0.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Polish-Ukrainian Fore-Carpathian Basin: Shushkivtsi, Zhukivtsi, (Ukraine).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF2067EDF2BD0249FAE2FA2B.taxon	discussion	Discussion. Mentioned by Handmann (1883) in a faunal list from Gainfarn (Austria) without description.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF2067EDF2BD015EFDE5F8B6.taxon	discussion	Discussion. The illustration shows a minute Retusidae with prominent spiral grooves. The specimen is lost, and we are unsure if it corresponds to one of the species described herein.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF2067EDF2BD015EFDE5F8B6.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Polish-Ukrainian Fore-Carpathian Basin: Krzemieniec, Shushkivtsi (Ukraine).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF2167ECF2BD0786FF08FE3C.taxon	discussion	Discussion. Berger (1953) described a single specimen from Kaisersteinbruch in the Vienna Basin (Austria) (SL: 20 mm, MD: 13 mm). This occurrence is highly dubious because this limestone locality does not yield aragonitic gastropod shells. The specimen was probably misplaced in the collection. The shell is lost and therefore, it is impossible to clarify its status. We exclude this Pliocene to Recent species from the Miocene fauna of the Vienna Basin.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF2167ECF2BD052CFDD5FCF9.taxon	discussion	Discussion. Kókay (1985: 67, pl. 9, figs 17 – 20) referred to this Eastern Paratethyan Sarmatian species when describing internal casts from the upper Badenian of Budapest and Pécsvárad (Hungary). The illustrated specimens are internal casts of Pyrunculus sp. (or Retusa truncatula) (figs 17, 19, 20) and probably Cylichna sp. (fig. 18) but surely not Retusa melitopolitana (Sokolov, 1899) as described by Kolesnikov (1935: 292, pl. 33, figs 21 – 24) and Frolov et al. (2020: 11, fig. 4 / 3).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF2167ECF2BD0490FC60FB98.taxon	discussion	Discussion. This Pliocene to present-day species was recorded by Berger (1953: 99, pl. 17, figs 40 – 44) from Bad Vöslau and Vienna / Pötzleinsdorf (Austria). We have not been able to trace his illustrated specimens to clarify their status, but based on the drawings, Berger (1953) lumped different species. If some of these correspond to Retusa umbilicata as reviewed by Landau et al. (2023: pl. 35) remains unclear.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF2167ECF2BD03B1FE30FA06.taxon	discussion	Discussion. Răileanu & Negulescu (1964) illustrated a specimen with very high spire and close-set spiral grooves from the Eggenburgian (Early Miocene) of Sălătruc in the Petroşeni Basin (Romania). It differs from Ringicula longominor Sacco, 1892 in its higher spire and more globose last whorl. The specimen lacks part of the columella and its status remains unclear.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF2167ECF2BD0130FDBEF86B.taxon	discussion	Discussion. Friedberg (1928) described a small, globose shell with delicate spiral sculpture and sharp columellar fold, which he identified as Tornatellaea biplicata Borson, 1821 [introduced by Borson (1821: 347, pl. 2, fig. 16)]. Some years later, however, Fridberg (1936: 113) recognized that this specimen was based on a strongly demaged Ringicula.	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
A67987E6FF2167ECF2BD0130FDBEF86B.taxon	distribution	Distribution in Central Paratethys. Badenian (Middle Miocene): Polish-Ukrainian Fore-Carpathian Basin: Borki Wielkie (Poland) (Friedberg 1928).	en	Harzhauser, Mathias, Landau, Bernard M., Malaquias, Manuel António E. (2026): Marine shelled Heterobranchia in part (Gastropoda) of the Miocene Central Paratethys Sea. Zootaxa 5742 (1): 1-150, DOI: 10.11646/zootaxa.5742.1.1, URL: https://doi.org/10.11646/zootaxa.5742.1.1
