taxonID	type	description	language	source
D446F043FFF2FFDA8C92FEC1FA140700.taxon	type_taxon	Type species. Mitra sturi Hoernes & Auinger, 1880, middle Miocene, Paratethys Sea, Romania.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFF2FFDA8C92FEC1FA140700.taxon	diagnosis	Diagnosis. Small, broad to moderately broad fusiform to nassariform shells, with beaded sculpture differing from Thala and Mitromica in the development of a parietal denticle and less cyrtoconoid spire, and from Thala in the absence of internal denticles as signs of episodic growth.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFF2FFDA8C92FEC1FA140700.taxon	description	Description. Shell small, broad to moderately broad fusiform, with conical to slightly cyrtoconoid, weakly gradate spire with incised suture. Protoconch multispiral, high conical with about three convex to weakly convex whorls. Sculpture of axially and spirally arranged subquadrate nodes, resulting in beaded appearance. Aperture narrow with incised anal sinus bordered by distinct parietal denticle. Outer lip with strong denticles, columella with four folds of which adapical one is most prominent. Siphonal canal moderately long, slightly bent.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFF2FFDA8C92FEC1FA140700.taxon	etymology	Etymology. Combination of anti (ancient Greek: against) and Thala. Referring to Thala H. Adams & A. Adams, 1853 into which the Paratethyan Antithala species have been placed so far. Included species. Antithala claviformis nov. sp., Antithala filipescui nov. sp., Mitra neugeboreni Hoernes & Auinger, 1880, Mitra sturi Hoernes & Auinger, 1880. Stratigraphic and geographic range. Only known so far from middle Miocene (Badenian) deposits of the Central Paratethys Sea, recorded from Romania, Hungary, Bosnia and Herzegovina, and Poland. Palaeoenvironment. The localities represent middle neritic palaeoenvironments with soft bottoms. No occurrences from coastal-neritic settings are known.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFF2FFDA8C92FEC1FA140700.taxon	discussion	Discussion. Rosenberg & Salisbury (2003) separated the genus Mitromica Berry, 1958 from Thala H. & Adams, 1853 based, most importantly, on the evidence of episodic growth in Thala, attested by the presence of former outer lips retained as internal denticles ‘ varices’ in members of that genus, whereas Mitromica shows continuous growth. Other shell features cannot separate the two genera. According to Rosenberg & Salisbury (2003), Mitromica ranges from the Caribbean, Bahamas, Florida and Bermuda, the Eastern Pacific, southern Madagascar, South Africa, and West Africa, whereas Thala is restricted to the Indo-Pacific, with an overlapping range in South Africa. Antithala shares features in common with Vexillum Röding, 1798, Mitromica and Thala. The prominently beaded sculpture and shell shape are reminiscent of some Mitromica and Thala species, but those genera lack a parietal denticle, their suture is distinctly less incised, and the base is less constricted. Vexillum develops a similar aperture with a parietal denticle bordering the anal canal, but differs in its sculpture, which is dominated by axial elements, lacking prominent spirally arranged subquadrate beads. Bellardithala nov. gen. differs in its episodic growth and the absence of a parietal denticle.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFF2FFD58C92F8FDFB7704C5.taxon	description	Figs 3 A 1 – A 2, 3 B 1 – B 2, 3 C 1 – C 2	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFF2FFD58C92F8FDFB7704C5.taxon	materials_examined	Type material. Holotype: NHMW 1869 / 0001 / 0670, SL: 4.4 mm, Coşteiu de Sus (Romania), MD: 1.8 mm, figs 3 A 1 – A 2. Paratypes: NHMW 2020 / 0056 / 0001, SL: 4.5 mm, MD: 1.9 mm, Coşteiu de Sus (Romania), figs 3 B 1 – B 2; NHMW 2020 / 0056 / 0002, SL: 4.1 mm, MD: 1.9 mm. Coşteiu de Sus (Romania), figs 3 C 1 – C 2; 11 specimens, NHMW 2020 / 0056 / 0004, Coşteiu de Sus (Romania). Type locality. Coşteiu de Sus (Romania), Făget Basin. Type stratum. Silt and clay of the Dej Formation. Age. Middle Miocene, Badenian.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFF2FFD58C92F8FDFB7704C5.taxon	etymology	Etymology. “ Clava ” (Latin: club), referring to the club-like shape.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFF2FFD58C92F8FDFB7704C5.taxon	diagnosis	Diagnosis. Antithala species of small size, with prominently beaded sculpture, three beaded cords on spire whorls, four columellar folds.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFF2FFD58C92F8FDFB7704C5.taxon	description	Description. Shell small, stout fusiform. Protoconch unknown. Teleoconch of 4.5 straight-sided whorls with periphery at abapical suture; suture distinctly incised. Spire weakly gradate, conical to slightly cyrtoconoid. First teleoconch whorl with prominent axial ribs intersected by two weaker spiral grooves forming three spiral rows of nodes; spiral rows strengthen abapically, forming prominent subquadrate nodes on later whorls. Fourth spiral cord at abapical suture largely covered by succeeding whorl. Last whorl regularly convex at periphery, passing via distinct concavity into strongly constricted base. Seven to eight spiral rows of nodes on last whorl forming orthocline to weakly sigmoidal axial ribs. Fasciole weak, bearing about five spiral cords with reduced nodes. Aperture elongate, narrow with distinct anal sinus narrowed by broad, low parietal denticle. Columella narrow, sharply delimited, bearing four folds, abapically rapidly decreasing in strength. Outer lip thickened, with six small denticles within. Siphonal canal moderately long, recurved, bent slightly to left, with shallow siphonal notch. Shell measurements and ratios. SL: 3.5 – 4.5 mm, MD: 1.8 – 1.9 mm; AA: 45 – 51 °, SL / MD: 2.2 – 2.6, AL / AW: 4.8 – 5.6, AH / S: 2.2 – 2.5.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFF2FFD58C92F8FDFB7704C5.taxon	discussion	Discussion. The specimens here described as Antithala claviformis nov. sp. were probably identified by Hoernes & Auinger (1880) as Mitra sturi (Hoernes & Auinger, 1880) due to them having almost identical sculpture. Antithala claviformis, however, is constantly smaller, broader and has a strongly constricted base, resulting in a club-like last whorl. Most importantly, A. claviformis has four columellar folds whereas A. sturi has only three. Antithala filipescui nov. sp. and A. neugeboreni both differ in having four rows of nodules on the spire whorls. Palaeoenvironment. The assemblages from Coşteiu de Sus suggests middle neritic palaeoenvironments with soft bottom (own data).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFF2FFD58C92F8FDFB7704C5.taxon	distribution	Distribution in Central Paratethys. Badenian (middle Miocene): Făget Basin: Coşteiu de Sus.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFFDFFD68C92FAB8FA0C00F9.taxon	description	Figs 3 D 1 – D 2, 3 E 1 – E 2, 3 F 1 – F 2, 4 A.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFFDFFD68C92FAB8FA0C00F9.taxon	materials_examined	Type material. Holotype: NHMW 1876 / 0011 / 0012, SL: 4.3 mm, MD: 1.9 mm, Lăpugiu de Sus (Romania), illustrated in Hoernes & Auinger (1880, pl. 10, figs 31 a – c), figs 3 E 1 – E 2, Fig. 4 A. Paratypes: NHMW 2020 / 0057 / 0001, SL: 4.4 mm, MD: 1.9 mm. Lăpugiu de Sus (Romania), illustrated in Hoernes & Auinger (1880, pl. 10, figs 28 a – c), figs 3 F 1 – F 2; NHMW 2020 / 0057 / 0002, SL: 4.9 mm, MD: 2.1 mm. Lăpugiu de Sus (Romania), figs 3 D 1 – D 2. Additional material. NHMW 2020 / 0057 / 0003, illustrated in Hoernes & Auinger (1880, pl. 10, figs 29 a – c); NHMW 2020 / 0057 / 0004, illustrated in Hoernes & Auinger (1880, pl. 10, figs 30 a – c); 18 specimens NHMW 2020 / 0057 / 0005; all Lăpugiu de Sus (Romania). Type locality. Lăpugiu de Sus (Romania), Făget Basin. Type stratum. Silt and clay of the Dej Formation. Age. Middle Miocene, Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFFDFFD68C92FAB8FA0C00F9.taxon	etymology	Etymology. In honour of Sorin Filipescu (Babeş-Bolyai University, Cluj, Romania), in recognition of his contributions to the Neogene stratigraphy of the Transylvanian Basin.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFFDFFD68C92FAB8FA0C00F9.taxon	diagnosis	Diagnosis. Antithala species of small size, with beaded sculpture, four beaded cords on spire whorls, four columellar folds, adapical much stronger than lower three folds.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFFDFFD68C92FAB8FA0C00F9.taxon	description	Description. Shell small, broad biconical with weakly gradate spire. Protoconch (fig. 4 A) high conical of 2.7 weakly convex whorls (height: 500 μm, diameter: 460 μm). Teleoconch of 4 – 4.5 straight sided whorls with periphery at abapical suture; suture distinctly incised. Axial ribs on first teleoconch whorl intersected by weaker spiral grooves forming four spiral rows of subquadrate nodes on later whorls. Nine to ten beaded spiral rows on last whorl accompanied by four spiral cords on fasciole. Spiral groove below subsutural row of nodes slightly more pronounced throughout. Last whorl high; maximum convexity at periphery, weakly convex above, narrowly constricted. Aperture elongate, narrow with narrow anal sinus bordered by broad, low parietal denticle. Columella indistinct, narrow, bearing prominent adapical columellar fold and three weak folds below. Outer lip thickened, with five to six small irregular denticles within, confined to adapical two thirds of aperture. Siphonal canal moderately long, straight with shallow siphonal notch. Shell measurements and ratios. SL: 4.3 – 4.9 mm, MD: 1.9 – 2.1 mm; AA: 44 – 49 °, SL / MD: 2.2 – 2.4, AL / AW: 5.1 – 6.7, AH / S: 2.1 – 2.3.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFFDFFD68C92FAB8FA0C00F9.taxon	discussion	Discussion. Hoernes & Auinger (1880) discussed this species as ‘ variety’ of Mitra sturi (Hoernes & Auinger, 1880), but it differs from that species in its stout biconical outline. Moreover, its sculpture is less regular, less tuberculose and the axial ribs are more prominent. The adapical columellar fold of Antithala filipescui nov. sp. is stronger and the abapical folds weaker compared to A. sturi. The higher and narrower spire allows a separation from Antithala claviformis. The prominent adapical columellar fold distinguishes A. filipescui also from the otherwise similar A. neugeboreni (Hoernes & Auinger, 1880). In addition, the periphery of the last whorl of A. filipescui is in a more abapical position compared to A. neugeboreni. Palaeoenvironment. Unknown; the collections from Lăpugiu de Sus indicate a mixing of coastal-inner neritic and middle neritic elements.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFFDFFD68C92FAB8FA0C00F9.taxon	distribution	Distribution in Central Paratethys. Badenian (middle Miocene): Făget Basin: Lăpugiu de Sus (Romania).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFFEFFD68C92FEE4FECF06CC.taxon	description	Figs 5 A 1 – A 2, B 1 – B 2, C 1 – C 2 [Callithea] Neugeboreni nov. form. — Hoernes 1880: 125 [nomen nudum].	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFFEFFD68C92FEE4FECF06CC.taxon	materials_examined	Type material. Lectotype (designated herein): NHMW 1863 / 0040 / 0005, SL: 6.1 mm, MD: 2.7 mm, Lăpugiu de Sus (Romania), illustrated in Hoernes & Auinger (1880, pl. 10, figs 24 a – c), figs 5 A 1 – A 2. Paralectotypes: NHMW 2020 / 0055 / 0001, SL: 5.8 mm, MD: 2.3 mm, Lăpugiu de Sus (Romania), illustrated in Hoernes & Auinger (1880, pl. 10, figs 23 a – c), figs 5 B 1 – B 2; NHMW 2020 / 0055 / 0002, SL: 6.2 mm, MD: 2.1 mm, Lăpugiu de Sus (Romania), figs 5 C 1 – C 2; 5 specimens, NHMW 2020 / 0055 / 0003, Lăpugiu de Sus (Romania). Additional material. SMF 360348 (= SMF XII 2215 a), lectotype of Mitra (Costellaria) annae Boettger, 1902, illustrated by Zilch (1934, pl. 22, fig. 9), Senckenberg Museum, Frankfurt / Main, Germany, Coşteiu de Sus (Romania). Revised description. Shell small to medium-sized, broad fusiform with slightly gradate spire and weakly undulating suture. Protoconch largely missing, last protoconch whorl high, smooth. Teleoconch of 5.5 straight-sided to weakly convex whorls. Axial sculpture of spire whorls of prominent axial ribs separated by narrower interspaces, intersected by three weak spiral grooves forming four rows of subquadrate rounded nodes at intersections. Subsutural row of nodes more prominent, separated from following spiral rows by slightly deeper spiral groove. A narrow secondary spiral row of elongate nodes is intercalated in this groove in some specimens. Last whorl strongly constricted with distinct basal concavity and long base, bearing 10 – 12 spiral rows of nodes plus four spirals on fasciole; spiral sculpture usually weaker than axial ribs; nodes often reduced. Aperture elongate, narrow with distinct, narrow anal sinus, bordered by weak parietal denticle. Columella narrow, sharply delimited, bearing four folds, weakening abapically. Outer lip thickened with up to ten denticles of variable strength within. Siphonal canal moderately long, straight or bent slightly to left, with shallow siphonal notch. Shell measurements and ratios. SL: 5.8 – 6.2 mm, MD: 2.3 – 2.9 mm, AA: 41 °, SL / MD: 2.4 – 2.6, AL / AW: 7.2 – 7.4, AH / S: 2.1 – 2.4.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFFEFFD68C92FEE4FECF06CC.taxon	discussion	Discussion. Antithala neugeboreni (Hoernes & Auinger, 1880) can be differentiated from A. sturi (Hoernes & Auinger, 1880) by its broader outline, gradate spire and the reduced nodes on the last whorl, resulting in dominant axial sculpture. Mitra annae was based by Boettger (1902) on two specimens from Coşteiu de Sus. Boettger (1902) recognized the similarity with A. neugeboreni, but separated M. annae based on the weak spiral sculpture on the last whorl. The reduction of spiral elements is typical in A. neugeboreni and, after having studied the lectotype of M annae, we consider it to be simply a morphotype of A. neugeboreni. Palaeoenvironment. Unknown; the collections from Lăpugiu de Sus indicate a mixing of coastal-inner neritic and middle neritic elements whereas the assemblages from Coşteiu de Sus suggest middle neritic palaeoenvironments with soft bottom (own data).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFFEFFD68C92FEE4FECF06CC.taxon	distribution	Distribution in Central Paratethys. Badenian (middle Miocene): Făget Basin: Lăpugiu de Sus, Coşteiu de Sus (Romania).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFFEFFD08C92F8B2FBC60411.taxon	description	Figs 4 B, 5 D 1 – D 2, E 1 – E 2, F 1 – F 2 [Callithea] Sturi nov. form. — Hoernes 1880: 125 [nomen nudum].	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFFEFFD08C92F8B2FBC60411.taxon	description	Th [ala]. sturi (R. Hörn, et Au.) — Sieber 1958 a: 153.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFFEFFD08C92F8B2FBC60411.taxon	materials_examined	Type material. Lectotype (designated herein): NHMW 2020 / 0058 / 0001, SL: 5.3 mm, MD: 1.9 mm, Lăpugiu de Sus (Romania), figs 4 B, 5 D 1 – D 2. Paralectotypes: NHMW 1863 / 0015 / 0155, SL: 4.6 mm, MD: 1.9 mm, Lăpugiu de Sus (Romania), illustrated in Hoernes & Auinger (1880, pl. 10, figs 27 a – c), figs 5 E 1 – E 2; NHMW 2020 / 0058 / 0003, SL: 5.9 mm, MD: 2.2 mm, Lăpugiu de Sus (Romania), illustrated in Hoernes & Auinger (1880, pl. 10, figs 26 a – c), figs 5 F 1 – F 2; NHMW 2020 / 0056 / 0003, Coşteiu de Sus (Romania); NHMW 2020 / 0058 / 0002, Lăpugiu de Sus (Romania), illustrated in Hoernes & Auinger (1880, pl. 10, figs 25 a – c). 15 specimens, NHMW 2020 / 0058 / 0004, Lăpugiu de Sus (Romania). Revised description. Shell small, moderately slender, fusiform, with conical to faintly cyrtoconoid spire. Protoconch (fig. 4 B) high conical of three weakly convex, smooth whorls (height: 530 μm, diameter: 450 μm). Teleoconch of 4.5 straight-sided whorls, with periphery at abapical suture, separated by deeply incised suture. First teleoconch whorl with blunt, weakly opisthocline axial ribs separated by deep, narrow interspaces. Axial ribs intersected by two weaker spiral grooves resulting in three spiral rows of prominent, subquadrate nodes on spire whorls. Fourth spiral row of nodes intercalated on third and fourth teleoconch whorls along abapical suture, but largely covered by succeeding whorl. Axial and spiral interspaces deeply cut, resulting in regular, prominent sculpture of square nodes. Last whorl moderately convex, moderately constricted base, with nine spiral rows of nodes axially arranged in weakly sigmoidal ribs. About six prominent spiral cords with strongly reduced nodes on fasciole. Aperture elongate, narrow with distinct anal sinus narrowed by broad, low parietal denticle. Columella narrow, sharply delimited, bearing three prominent folds, weakening abapically, plus weaker fourth fold bordering siphonal canal. Outer lip strongly thickened, with five to six stout denticles within. Siphonal canal moderately long, bent slightly to left, with shallow siphonal notch. Shell measurements and ratios. SL: 4.6 – 5.9 mm, MD: 1.9 – 2.2 mm; AA: 40 °, SL / MD: 2.7 – 2.9, AL / AW: 5.7 – 7.9, AH / S: 2.2 – 2.3.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFFEFFD08C92F8B2FBC60411.taxon	discussion	Discussion. Antihala sturi (Hoernes & Auinger, 1880) is characterised by its regularly beaded sculpture and displays little variability in shape. Some specimens develop a slightly wider spiral groove below the first adapical spiral cord (e. g. Bałuk 1997, pl. 11, figs 4 – 6). We are not aware of any similar species from the European Neogene except for the Paratethyan species discussed herein. Palaeoenvironment. Unknown; the collections from Lăpugiu de Sus indicate a mixing of coastal-inner neritic and middle neritic elements.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFFEFFD08C92F8B2FBC60411.taxon	distribution	Distribution in Central Paratethys. Badenian (middle Miocene): Korytnica Basin: Korytnica (Poland) (Bałuk 1997); Ukrainian Fore-Carpathian Basin: Zborów (Zboriv) (Ukraine); (Friedberg 1911,); Pannonian Basin: Herend (Hungary) (Kókay 1966); southern Pannonian Basin: Hrvaćani (Bosnia and Herzegovina) (Atanacković 1985); Făget Basin: Lăpugiu de Sus (own data), Coşteiu de Sus (Romania) (Boettger 1906).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFF8FFD28C92FBCCFA14034D.taxon	type_taxon	Type species: By typification of replacement name: Voluta obsoleta Brocchi, 1814, Pliocene, Italy [for figures of holotype see Rossi Ronchetti (1955: fig. 129) and Pinna & Spezia (1978, pl. 68, figs 2, 2 a).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFF8FFD28C92FBCCFA14034D.taxon	diagnosis	Diagnosis. Medium-sized Thala - and Mitromica - like shells with moderately slender fusiform shells, cancellate, weakly beaded or strongly reduced sculpture. Differing from Mitromica in their episodic growth, indicated by the formation of denticles in juvenile specimens and from Thala in the predominant axial sculpture on early spire whorls. Occasionally with elongate denticles within outer lip.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFF8FFD28C92FBCCFA14034D.taxon	description	Description. Shell medium-sized, moderately slender fusiform with conical to weakly cyrtoconoid spire. Protoconch high conical, mammillate, consisting of about three smooth whorls; teleoconch of 4.5 to 5.5 weakly convex whorls. Sculpture of early spire whorls dominated by axial ribs crossed by weak or subordinate spiral cords, often confined to axial interspaces. Axial sculpture prominent or largely reduced on late spire whorls and last whorl, crossed by numerous spiral cords forming cancellate or weakly beaded sculpture. Aperture narrow to moderately narrow; anal sinus indistinct. Inner lip indistinct, narrow; columella with four folds. Outer lip weakly to distinctly thickened, dentate only in adults; elongated denticles occur at least in several species (own data B. L). Siphonal canal moderately long, bent weakly to left with shallow siphonal notch. Synonyms. Micromitra Bellardi (1888) [non Meek, 1873].	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFF8FFD28C92FBCCFA14034D.taxon	etymology	Etymology. Combination of the surname of Luigi Bellardi (1818 – 1889), Italian malacologist, who has establish the costellariid genus Micromitra (name preoccupied) for the species placed here in Bellardithala, and the genus Thala. Included species. Voluta obsoleta Brocchi, 1814 (for non-Paratethyan species see Bellardi 1888, Sacco 1904, Rossi Ronchetti 1955; Pinna & Spezia 1978, Giannuzzi Savelli & Reina 1883; 1978; Ferrero-Mortara et al. 1984, and Landau et al. 2019), Bellardithala baluki nov. sp., B. dacica nov. sp., B. fedosovi nov. sp., B. kovaci nov. sp., Micromitra abbreviata Bellardi, 1888, M. granosa Bellardi, 1888, M. intermedia Bellardi, 1888, M. mangeliaeformis Bellardi, 1888, M. propinqua Bellardi, 1888, M. pusilla Bellardi, 1888, M. seminuda Bellardi, 1888, M. taurina Bellardi, 1888, Mitra (Micromitra) boehmi Boettger 1906, Mitra (Thala) kostejana Boettger 1906, M. lapugyensis Hoernes & Auinger 1880, M. laubei Hoernes & Auinger 1880, M. partschi Hörnes 1852, M. pupa Dujardin, 1837, Thala angustolonga Sacco, 1904, T. elatocostata Sacco, 1904, T. inflatoparva Sacco, 1904, T. subnullecostata Sacco, 1904.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFF8FFD28C92FBCCFA14034D.taxon	discussion	Stratigraphic and geographic range. Early Miocene to Pliocene. The earliest representatives of Bellardithala appeared during the Burdigalian in the Proto-Mediterranean Sea, recorded from the Colli Torinesi in Italy (e. g. B. taurina Bellardi, 1888, B. propinqua Bellardi, 1888, B. seminuda Bellardi, 1888). Middle Miocene occurrences are documented from the Badenian (Langhian, Serravallian) of the Central Paratethys Sea as described herein. There, the genus displays its maximum diversity in the southern basins and during the early / middle Badenian (= Langhian). From the northeastern Atlantic the genus is documented from the Langhian to the Messinian (Landau et al. 2019). The last occurrences are described from the Pliocene of the Mediterranean Sea (i. e. B. obsoleta) (Landau et al. 2019).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFF8FFD28C92FBCCFA14034D.taxon	description	Palaeoenvironment. The occurrences suggest a distribution from shallow marine, inner neritic sand environments down to middle neritic soft bottom habitats.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFF8FFD28C92FBCCFA14034D.taxon	discussion	Discussion. Bellardi (1888: 5) established Micromitra as new genus for “ small subovate species with moderately inflated spire, long aperture with canal, thickened, varix-like outer lip with internal denticles and four columellar folds ” (free translation of parts of the original diagnosis in Latin). Bellardi (1888: 5) designated Mitra obsoleta Brocchi, 1814 from the Pliocene of Italy as type species of Micromitra and placed several Burdigalian and Pliocene species from Italy in his new genus [Coan (1966: 130) designated Mitra taurina Bellardi, 1888 as type species, obviously overlooking that Bellardi (1888) had already designated a type species]. The name Micromitra, however, was already preoccupied by Meek (1873: 479) for a brachiopod and the Neogene gastropod genus would have needed a new name. No replacement name was proposed subsequently because Micromitra Bellardi, 1888 was treated as subjective junior synonym of Thala H. Adams & A. Adams, 1853 by most subsequent authors (Cossmann 1899; Peyrot 1903, 1928; Sacco 1904; Sphon 1969; Cernohorsky 1970; Giannuzzi-Savelli & Reina 1983). Only Boettger (1906) used Micromitra Bellardi, 1888 as subgenus of Mitra for his Mitra (Micromitra) boehmi, separating it from Thala species. A further potential name for the Neogene European species is Mitromica Berry, 1958, which was established for extant species from the Eastern Pacific (type species: Mitra solitaria C. B. Adams, 1852). Conchological differences between Mitromica and Thala are subtle and Rosenberg & Salisbury (2003) stressed the continuous growth of Mitromica versus episodic growth of Thala, reflected by the development of denticles in juvenile stages and growth stops on the shell surface. Furthermore, Rosenberg & Salisbury (2003) assumed a geographic separation of both genera. Thala is restricted to Indo-Pacific species whereas western African, western Atlantic and Eastern Pacific species are placed in Mitromica Berry, with overlap in distribution of the two genera only around South Africa. Separation of a ‘ western’ Mitromica - clade from an Indo-Pacific Thala - clade is also supported by radular analyses by Fedosov et al. (2017; pers. comm. May 2020). Molecular data to support these clades is, however, not yet available. Rosenberg & Salisbury (2003) observed episodic growth and denticles in juveniles in the Paratethyan ‘ Thala ’ partschi kostejana Boettger, 1906 and therefore, treated it as Thala. Herein, we document episodic growth also for Bellardithala kovaci nov. sp., B. pupa, B. boehmi, and B. kostejana. Thus, despite some morphological similarities, the Paratethyan species cannot be placed in Mitromica. Placement in Thala, however, is problematic as well. The middle Miocene Paratethyan species and all early Miocene to Pliocene species, originally placed in Micromitra by Bellardi (1888), are characterised by predominant axial sculpture during early stages of growth, whereas the spiral sculpture is subordinate. None of the species develops pointed nodes and prominent beads like Indo-Pacific Thala. In some species, such as M. kovaci, M. obsoleta and M. taurina, the sculpture is largely reduced on the last whorls, which would be an atypical feature for Indo-Pacific Thala (see Thala species described in Rosenberg & Salisbury, 2003, 2014). Therefore, we consider the Old World species to represent a clade distinct from Thala and Mitromica, and erect the replacement name Bellardithala nom. nov. for Micromitra Bellardi, 1888, non Meek, 1873. Early ontogeny. The protoconchs of Thala and Mitromica seem to have little taxonomic value above the species level. The extant species currently listed as Mitromica and Thala by Rosenberg & Salisbury (2003, 2014) and Fedosov et al. (2017) suggest two different modes of development. Paucispiral bulbous protoconchs with inflated whorls are found in both genera [e. g. M. solitaria (C. B. Adams, 1852), M. foveata (Sowerby, 1874) and Thala todilla (Mighels, 1845), T. merrilli Rosenberg & Salisbury, 2014]. Likewise, multispiral protoconchs occur in both genera [e. g. M. africana (Rolán & Fernandes, 1995) and T. mirifica (Reeve, 1845), T. jaculanda (Gould, 1860)]. The species of Bellardithala display high conical protoconchs. In Thala, this type of protoconch correlates with wide geographic distributions (Rosenberg & Salisbury 2014) and the endemicity of Paratethyan species has to be treated with caution, because it may rather be a collecting artifact. We note that all protoconchs of Bellardithala species preserved are tall, multispiral, and in most species mammilate. Species with paucispiral protoconchs have not been found until now.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFFAFFCC8C92FC31FE77011D.taxon	description	Figs 6 A 1 – A 2, B 1 – B 2, C 1 – C 2	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFFAFFCC8C92FC31FE77011D.taxon	materials_examined	Type material. Lectotype: SMF 360342 (= SMF XII. 2217 a), SL: 6.4 mm, MD: 2.3 mm, illustrated by Zilch (1934, pl. 22, fig. 14), Senckenberg Museum, Frankfurt / Main, Germany, Coşteiu de Sus (Romania), figs 6 A 1 – A 2. Additional material. NHMW 2020 / 0059 / 0001, SL: 7.9 mm, MD: 2.8 mm, Lăpugiu de Sus (Romania), figs 6 B 1 – B 2; NHMW 1854 / 0035 / 0107, SL: 7.9 mm, MD: 2.7 mm, Lăpugiu de Sus (Romania), figs 6 C 1 – C 2; 7 specimens, NHMW 2020 / 0059 / 0002, Lăpugiu de Sus (Romania); BL collection, 4 specimens, Lăpugiu de Sus (Romania). Revised description. Shell medium-sized, moderately slender fusiform with conical spire. Protoconch high mammillate of three smooth, weakly convex whorls; height: 600 μm, diameter: 550 μm. Teleoconch of 5.5 weakly convex whorls with periphery slightly above abapical suture. Suture narrowly incised. First three teleoconch whorls with, convex axial ribs, separated by narrower interspaces. Axial sculpture intersected by about 8 spiral grooves. Sculpture weakening abruptly after third whorl. Penultimate whorl nearly smooth; last whorl with faint spiral cords. Prominent spiral cords on base and fasciole with delicate axial growth lines in interspaces. Aperture moderately narrow, elongate with narrow anal sinus; inner lip narrow. Columella with four prominent folds (adapical two being stronger). Outer lip thickened with 10 – 12 small denticles, becoming nearly obsolete in abapical third of aperture. Last whorl convex, moderately constricted at base. Siphonal canal moderately short, straight with shallow anterior notch. Shell measurements and ratios. SL: 6.4 – 7.9 mm, MD: 2.3 – 2.8 mm, AA = 37 – 42 °, SL / MD: 2.7 – 3.0, AL / AW: 4.4 – 5.0, AH / S: 2.5 – 2.7.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFFAFFCC8C92FC31FE77011D.taxon	discussion	Discussion. This rare species is reminiscent of Thala kovaci nov. sp. (= Thala obsoleta sensu Hörnes, 1852, non Brocchi, 1814) due to its reduced sculpture on the last whorl, but differs in its more slender fusiform outline, the much more prominent axial and spiral sculpture on the early teleoconch whorls, and the lower aperture / spire ratio (0.9 versus 1.1). Bellardithala seminuda (Bellardi, 1888), from the Burdigalian of the Colli Torinesi (Italy), is reminiscent of B. boehmi, but differs in its larger size and in the absence of spiral sculpture on the last whorl (see Bellardi 1888: 7, pl. 6, figs 46 a – b). Palaeoenvironment. Unknown; the collections from Lăpugiu de Sus indicate a mixing of coastal-inner neritic and middle neritic elements.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFFAFFCC8C92FC31FE77011D.taxon	distribution	Distribution in Central Paratethys. Badenian (middle Miocene): Făget Basin: Lăpugiu de Sus (own data), Coşteiu de Sus (Boettger 1906).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFE4FFCF8C92FECDFCC502A9.taxon	description	Figs 4 C, 6 D 1 – D 2, E 1 – E 2, F 1 – F 2, G 1 – G 2, H 1 – H 2, I 1 – I 2	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFE4FFCF8C92FECDFCC502A9.taxon	materials_examined	Type material. Holotype: NHMW 1846 / 0037 / 0114, SL: 8.3 mm, MD: 3.1 mm, Steinebrunn (Austria), illustrated in Hörnes (1852 b, pl. 10, figs 32 a – b), figs 6 D 1 – D 2. Paratypes: NHMW 1858 / 0047 / 0007, SL: 8.9 mm, MD: 3.3 mm, Sedlec (= Porzteich) (Czech Republic), figs 6 E 1 – E 2; NHMW 1866 / 0001 / 1019, SL: 6.8 mm, MD: 2.7 mm, Forchtenau (Austria), figs 6 F 1 – F 2; NHMW 1871 / 0010 / 0331, SL: 6.0 mm, MD: 2.2 mm, Steinebrunn (Austria), figs 4 C, 6 G 1 – G 2; NHMW 1871 / 0011 / 0004, SL: 5.9 mm, MD: 2.2 mm, Steinebrunn (Austria), figs 6 H 1 – H 2. Additional material. NHMW 2020 / 0062 / 0001, SL: 5.7 mm, MD: 2.1 mm, Steinebrunn (Austria), figs 6 I 1 – I 2. 18 specimens NHMW 2020 / 0062 / 0002, all Steinebrunn (Austria). Type locality. Steinebrunn (Austria). Type stratum. Silty sand of the Baden Formation. Age. Middle Miocene, Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFE4FFCF8C92FECDFCC502A9.taxon	etymology	Etymology. In honour of Michal Kováč (Comenius University, Bratislava, Slovakia), in recognition of his contributions on the stratigraphy and paleogeography of the Paratethys Sea.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFE4FFCF8C92FECDFCC502A9.taxon	diagnosis	Diagnosis. Medium-sized, moderately slender fusiform shell differing from other Bellardithala species by its weak spiral sculpture and broad flat axial ribs separated by very narrow interspaces.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFE4FFCF8C92FECDFCC502A9.taxon	description	Description. Shell medium-sized, moderately slender fusiform with slightly cyrtoconoid spire. Protoconch (fig. 4 C) high conical of 3.5 weakly convex whorls; height: 660 μm, diameter: 580 μm. Teleoconch of moderately convex 5.5 whorls with periphery slightly below mid-whorl; suture narrowly incised. Last whorl high, weakly convex with weak basal constriction. Sculpture of first teleoconch whorl of broad, slightly opisthocline axial ribs with flattened tops separated by narrow interspaces. Spiral sculpture reduced to five spiral threads in axial interspaces. Axial ribs becoming broad and low on later spire whorls with irregularly spaced narrow interspaces. Spiral threads restricted to axial interspaces causing tiny pits between axial and spiral elements. Interspaces very narrow, present throughout growth. Spiral grooves somewhat stronger in adapical third of last whorl causing tiny pits at intersections with axial interspaces. Axial sculpture weakening towards fasciole. Aperture elongate, moderately narrow with distinct, narrow anal sinus. Inner lip narrow, indistinct. Columella with four broad folds, adapical two most prominent. Outer lip thickened with 10 denticles on inner side. Siphonal canal moderately short, moderately wide, straight with shallow siphonal notch. Shell measurements and ratios. NHMW specimens: SL: 6.8 – 8.5 mm, MD: 2.7 – 3.3 mm; AA: 44 – 46 °, SL / MD: 2.8, AL / AW: 4.7 – 5.3, AH / S: 2.4 – 2.6.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFE4FFCF8C92FECDFCC502A9.taxon	discussion	Remarks. Bellardithala kovaci is documented also by subadult specimens (figs 6 G 1 – G 2, H 1 – H 2) illustrating the development of denticles in juveniles, which was also observed in B. partschi by Rosenberg & Salisbury (2003). This species was misidentified so far as Bellardithala obsoleta (Brocchi, 1814), which was originally described from the Italian Pliocene. The holotype of Voluta obsoleta Brocchi, 1814, illustrated in Rossi Ronchetti (1955) and Pinna & Spezia (1978), differs clearly from the Paratethyan species in having prominent axial sculpture, on spire whorls and last whorl. Pliocene specimens of B. obsoleta from Altavilla (Italy), described by Giannuzzi-Savelli & Reina (1983), differ from the Paratethyan species also in the more prominent axial and spiral sculpture and have a more slender fusiform outline. Serravallian specimens from the Karaman Basin in Turkey, described by Landau et al. (2013) as Thala obsoleta, differ from the Paratethyan species clearly in the prominent axial ribs and the subcylindrical spire whorls. The same features distinguish Tortonian specimens from Montegibbio in Italy from Bellardithala kovaci (see Davoli, 2000). Bellardithala boehmi (Boettger, 1906) differs in its more slender fusiform outline, much more prominent axial and spiral sculpture on the early teleoconch whorls, and the lower aperture / spire ratio (0.9 versus 1.1). Palaeoenvironment. Coastal, inner neritic. The assemblage from Steinebrunn suggests shallow sublittoral environments with sea grass (Harzhauser & Landau 2019). At the Niederleis section, sediment and fossils from coastal and lagoonal environments not exceeding 30 m water depth were transported by tempestites into offshore settings (Mandic et al. 2002).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFE4FFCF8C92FECDFCC502A9.taxon	distribution	Distribution in Central Paratethys. Badenian (middle Miocene): North Alpine-Carpathian Foreland Basin: Unkovice (Czech Republic) (Rzehak 1894); Korytnica Basin: Korytnica (Poland) (Bałuk 1997); Vienna Basin: Niederleis, Steinebrunn (Austria); Mikulov, Sedlec (= Porzteich) (Czech Republic) (Hoernes & Auinger 1880); Eisenstadt-Sopron Basin: Forchtenau (Austria) (Sieber 1956).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFE7FFCF8C92FD41FA9007F2.taxon	description	Figs 7 A 1 – A 2, B 1 – B 2, C 1 – C 2	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFE7FFCF8C92FD41FA9007F2.taxon	materials_examined	Type material. Holotype: NHMW 1859 / 0047 / 0015, SL: 8.1 mm, MD: 2.7 mm, Vienna / Pötzleinsdorf (Austria), figs 7 A 1 – A 2. Paratypes: NHMW 1859 / 0047 / 0015 a, SL: 8.5 mm, MD: 2.9 mm, Vienna / Pötzleinsdorf (Austria), figs 7 B 1 – B 2; NHMW 2020 / 0063 / 0001, SL: 8.5 mm, MD: 2.9 mm, Vienna / Pötzleinsdorf (Austria), figs 7 C 1 – C 2. Type locality. Vienna / Pötzleinsdorf (Austria), Type stratum. Silty sand of the Baden Group. Age. Middle Miocene, Badenian (Serravallian).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFE7FFCF8C92FD41FA9007F2.taxon	etymology	Etymology. In honour of Wacław Bałuk (University of Warsaw, Warszawa, Poland) in recognition of his contributions on Paratethyan gastropod systematics.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFE7FFCF8C92FD41FA9007F2.taxon	diagnosis	Diagnosis. Bellardithala species of medium size, moderately slender fusiform shell, with cyrtoconoid spire, and beaded sculpture differing from congeneric species by its slender bullet-like spire and delicate sculpture.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFE7FFCF8C92FD41FA9007F2.taxon	description	Description. Shell medium-sized, moderately slender fusiform with cyrtoconoid spire and narrowly incised suture. Protoconch unknown. Teleoconch of 4.5 weakly convex whorls, with periphery at abapical third of whorl, rapidly increasing in height. First teleoconch whorl with blunt axial ribs, crossed by six weak spiral threads, most prominent in axial interspaces. Regularly spaced axial ribs crossed by ten spiral cords on penultimate whorl, forming low beads at intersections. Last whorl weakly convex, with faint subsutural concavity, weakly constricted at base, bearing about 22 beaded spiral cords, adapical subsutural spiral cord most prominent, bearing prosocline nodes. Cords over fasciole prominent, where they develop as extensions of columellar folds. Aperture moderately narrow with shallow anal sinus. Columellar callus indistinct, poorly delimited. Columella with four prominent columellar folds. Outer lip weakly thickened; ten delicate denticles within. Siphonal canal moderately long, straight or bent slightly to left, with distinct siphonal notch. Shell measurements and ratios. SL: 8.1 – 8.5 mm, MD: 2.7 – 2.9 mm, AA: 40 – 45 °, SL / MD: 3.0 – 3.1, AL / AW: 4.6 – 5.2, AH / S: 2.1 – 2.2.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFE7FFCF8C92FD41FA9007F2.taxon	discussion	Discussion. Specimens of Bellardithala baluki nov. sp. were mixed with B. laubei by Hoernes & Auinger (1880). However, B. baluki nov. sp. differs from B. laubei (Hoernes & Auinger, 1880), as redefined herein, in its broader shell, beaded sculpture especially on the last whorl, and it develops only 4.5 teleoconch whorls instead of 5.5 in B. laubei. Bellardithala fedosovi nov. sp. is similar in shell shape, but differs in its smaller size, the more convex last whorl, the wider interspaces between the axial ribs, and the overall coarser sculpture. The type species, B. obsoleta (Brocchi, 1814) from the Mediterranean Pliocene, differs in being slightly broader and the axial ribs are well developed and elevated. In B. baluki axial sculpture consists of axial rows of small beads, but do not form elevated ribs.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFE7FFCF8C92FD41FA9007F2.taxon	distribution	Distribution in Central Paratethys. Badenian (middle Miocene): Vienna / Pötzleinsdorf (Austria).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFE6FFCE8C92FF39FACC06CD.taxon	description	Figs 7 D 1 – D 2, E 1 – E 2	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFE6FFCE8C92FF39FACC06CD.taxon	materials_examined	Type material. Holotype: NHMW 1864 / 0001 / 0563, SL: 7.5 mm, MD: 2.8 mm, Steinebrunn (Austria), illustrated in Hoernes & Auinger (1880, pl. 10, fig. 22 a – c), figs 7 D 1 – D 2. Paratype: NHMW 2020 / 0052 / 0001, SL: 7.1 mm, MD: 2.7 mm, Steinebrunn (Austria), figs 7 E 1 – E 2. Type locality. Steinebrunn (Austria). Type stratum. Silty sand of the Baden Formation. Age. Middle Miocene, Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFE6FFCE8C92FF39FACC06CD.taxon	etymology	Etymology. In honour of Alexander Fedosov (Russian Academy of Sciences, Moscow), in recognition of his contributions on the molecular phylogeny of Costellariidae and Mitridae.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFE6FFCE8C92FF39FACC06CD.taxon	diagnosis	Diagnosis. Bellardithala species of medium size, moderately slender fusiform shell, with weakly cyrtoconoid spire, slowly contracting last whorl, beaded axial ribs, distinguished from congeneric species by the high number of spiral cords on the penultimate and last whorls, and the bifid subsutural spiral cord.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFE6FFCE8C92FF39FACC06CD.taxon	description	Description. Shell medium-sized, moderately slender fusiform with weakly cyrtoconoid spire and narrowly incised suture. Protoconch unknown. Teleoconch of 4.5 weakly convex whorls, with periphery in abapical third. Axial sculpture on first teleoconch whorl consisting of about ten prominent, orthocline to weakly opisthocline axial ribs, separated by narrower interspaces, crossed by five spiral cords. Spiral cords forming weak, rectangular nodes at intersections with ribs and narrow grooved in interspaces. Later teleoconch whorls with densely spaced axial ribs, crossed by seven to eight spiral cords resulting in somewhat blurred beads. Prominent, nodular bifid subsutural spiral cord, separated from following cord by deeper spiral groove, especially on last whorl; secondary nodular spiral cord intercalated in this groove in some specimens. Last whorl weakly convex with weakly constricted base, bearing about 15 beaded spiral cords, arranged in narrowly sigmoidal axial ribs. Fasciole weak, bearing weak spiral cords. Aperture elongate, moderately narrow, with indistinct anal sinus. Columellar callus narrow. Columella with four columellar folds, weakening abapically. Outer lip only weakly thickened, with about eight delicate denticles within. Siphonal canal wide, moderately long, straight, with shallow siphonal notch. Shell measurements and ratios. SL: 7.1 – 7.5 mm, MD: 2.3 – 2.7 mm, AA: 45 – 49 °, SL / MD: 2.7 – 2.9, AL / AW: 3.7 – 4.4, AH / S: 2.3 – 2.4.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFE6FFCE8C92FF39FACC06CD.taxon	discussion	Discussion. The holotype of Bellardithala fedosovi nov. sp. was mixed up with Bellardithala lapugyensis (Hoernes & Auinger, 1880) by those authors, but is readily distinguished from that species by its tubercular sculpture (compared to the cancellate sculpture of B. lapugyensis), and the greater number of spiral cords on the penultimate and last whorls. Bellardithala partschi (Hörnes, 1852) is somewhat similar, but also has fewer spiral cords, and the spire is less cyrtoconoid. Bellardithala baluki nov. sp. differs in having finer sculpture composed of more numerous axial rows of smaller beads rather than tubercles separated by narrower interspaces. As with the previous species, the type species B. obsoleta (Brocchi, 1814) differs in having more strongly developed and elevated axial ribs that are less strongly tubercular, separated by deeper interspaces. Palaeoenvironment. The Niederleis section exposed sediment and fossils from coastal and lagoonal environments not exceeding 30 m water depth, which were transported by tempestites into offshore settings (Mandic et al. 2002).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFE6FFCE8C92FF39FACC06CD.taxon	distribution	Distribution in Central Paratethys. Badenian (middle Miocene): Vienna Basin: Niederleis (Austria).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFE6FFC88C92F8B3FADD03A6.taxon	description	Figs 7 F 1 – F 2 [Callithea] Laubei nov. form. — Hoernes 1880: 125 [nomen nudum].	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFE6FFC88C92F8B3FADD03A6.taxon	materials_examined	Type material. Lectotype (designated herein): NHMW 1999 z 0076 / 0002, SL: 8.2 mm, MD: 2.4 mm, Bad Vöslau (Austria), illustrated in Hoernes & Auinger (1880, pl. 10, figs 19 a – c), figs 7 F 1 – F 2. Revised description. Shell medium-sized, very slender fusiform. Protoconch unknown. Teleoconch of 5.5 weakly convex whorls with periphery slightly above abapical suture. Suture incised. First teleoconch bearing about 11 opisthocline axial ribs, separated by narrow interspaces, crossed by faint spiral grooves. Spiral grooves strengthening slightly on later whorls, forming spiral rows of blurred nodes at intersections with axial ribs. Spiral cord at adapical suture most prominent, followed below by two spiral rows of nodes. Nodes on subsequent cords indistinct, blurred and largely reduced below periphery. Base slowly contracting. Aperture elongate, narrow, anal sinus indistinct. Narrow columellar callus. Columella with four oblique folds. Outer lip weakly thickened with faint denticles within. Siphonal canal long and wide, bent slightly to left, with shallow, wide siphonal notch. Shell measurements and ratios. SL: 8.2 mm, MD: 2.4 mm, AA: 29 °, SL / MD: 3.5, AL / AW: 6.94, AH / S: 2.3 – 2.4.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFE6FFC88C92F8B3FADD03A6.taxon	discussion	Discussion. Hoernes & Auinger (1880) based Mitra laubei on a series of syntypes, which, in our opinion, are not conspecific. As first revisers we designate specimen NHMW 1999 z 0076 / 0007 from Bad Vöslau (Austria), illustrated by Hoernes & Auinger (1880, pl. 10, figs 19 a – 19 c) as lectotype. All other specimens are described herein as new species (Bellardithala baluki, B. dacica). Bellardithala laubei (Hoernes & Auinger, 1880) differs from all of its Paratethyan congeners in having a far more slender fusiform outline and slightly more convex whorls. In sculpture it is similar to B. cf. dacica nov. sp., but that species is broader fusiform, and the spire whorls are slightly scalate in profile. Palaeoenvironment. The clay at Bad Vöslau formed in middle neritic settings with up to 250 m water depth (Hohenegger et al. 2008).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFE6FFC88C92F8B3FADD03A6.taxon	distribution	Distribution in Central Paratethys. Badenian (middle Miocene): Vienna Basin: Bad Vöslau (Austria).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFE0FFCB8C92FC58FBAD0286.taxon	description	Figs 8 A 1 – A 2, B 1 – B 2, C 1 – C 2	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFE0FFCB8C92FC58FBAD0286.taxon	materials_examined	Type material. Holotype: NHMW 1854 / 0035 / 0105, SL: 6.9 mm, MD: 2.4 mm, Lăpugiu de Sus (Romania), figs 8 A 1 – A 2. Paratypes: NHMW 2020 / 0061 / 0001, SL: 7.3 mm, MD: 2.4 mm, Lăpugiu de Sus (Romania), figs 8 B 1 – B 2; NHMW 2020 / 0061 / 0002, SL: 6.9 mm, MD: 2.4 mm, Lăpugiu de Sus (Romania), figs 8 C 1 – C 2. Type locality. Lăpugiu de Sus (Romania), Făget Basin. Type stratum. Silt and clay of the Dej Formation. Age. Middle Miocene, Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFE0FFCB8C92FC58FBAD0286.taxon	etymology	Etymology. Referring to Dacia, a province of the Roman Empire, covering also the territory of the Transylvanian Basin.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFE0FFCB8C92FC58FBAD0286.taxon	diagnosis	Diagnosis. Bellardithala species of medium size, moderately slender fusiform shell, with faintly cyrtoconoid slightly gradate spire, differing from congeneric species by its sculpture, which changes from prominent axial ribs and spiral cords with distinct nodes on early teleoconch whorls to densely spaced, cancellate pattern, with blurred nodes on later spire whorls and last whorl.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFE0FFCB8C92FC58FBAD0286.taxon	description	Description. Shell medium-sized, slender fusiform with faintly cyrtoconoid and slightly gradate spire. Protoconch largely damaged; last two protoconch whorls suggesting high conical shape. Teleoconch of 4.5 weakly convex whorl, separated by impressed suture, with periphery shifting from abapical suture to mid-whorl during growth. First teleoconch whorl with axial ribs separated by interspaces of about same width, crossed by five close-set spiral cords, forming spirally elongated nodes at intersections with ribs, persisting into interspaces. Nodes on adapical spiral row most prominent. Axial ribs broader and less prominent on later teleoconch whorls. Five to six spiral cords on later spire whorls, forming cancellate pattern with axial ribs. Spiral cord at adapical suture broader than others, separated by wider and deeper spiral groove from next spiral cord. Last whorl high, weakly convex, slowly contracting, with up to 25 densely spaced spiral cords. Nodes on spiral cords somewhat blurred above periphery and largely reduced below. Aperture elongate, narrow to moderately narrow, with indistinct anal notch. Columellar callus narrow, indistinct. Columella with four prominent columellar folds, weakening only slightly abapically. Outer lip weakly thickened, with about 11 delicate denticles within. Siphonal canal moderately long, bent slightly to left, with shallow siphonal notch. Shell measurements and ratios. SL: 6.9 – 7.3 mm, MD: 2.4 mm, AA: 35 – 40 °, SL / MD: 2.8 – 3.1, AL / AW: 5.0 – 5.5, AH / S: 2.1 – 2.3.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFE0FFCB8C92FC58FBAD0286.taxon	discussion	Discussion. The holotype of Bellardithala dacica nov. sp. was mixed with Mitra laubei (Hoernes & Auinger, 1880) by those authors. Despite some superficial similarity due to the prominent subsutural spiral cord, the species are distinguished by the slender shape of B. laubei and the different sculpture of early spire whorls (broad axial ribs with subordinate spiral sculpture in B. laubei vs narrower axial ribs and prominent spiral cords in M. dacica). A specimen described by Kókay (1966) as Thala laubei from Herend (Hungary) differs from B. laubei also in its broader shape. It is reminiscent of B. cf. dacica but differs in its higher number of axial ribs on the spire and the distinct subsutural spiral cord. This specimen might indicate a further undescribed species.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFE0FFCB8C92FC58FBAD0286.taxon	distribution	Distribution in Central Paratethys. Badenian (middle Miocene): Ukrainian Fore-Carpathian Basin: Żukowce (Zhukivtsi) (Ukraine) (Friedberg 1938); Făget Basin: Lăpugiu de Sus (Romania).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFE2FFCA8C92FF39FD9904DF.taxon	description	Figs 7 G 1 – G 2, H 1 – H 2, I 1 – I 2	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFE2FFCA8C92FF39FD9904DF.taxon	materials_examined	Material. NHMW 1866 / 0001 / 0806, SL: 7.5 mm, MD: 2.6 mm, Niederleis (Austria), illustrated in Hoernes & Auinger (1880, pl. 10, figs 20 a – c), figs 7 G 1 – G 2; NHMW 2020 / 0054 / 0001, SL: 6.7 mm, MD: 2.6 mm, Niederleis (Austria), figs 7 H 1 – H 2; NHMW 2020 / 0054 / 0002, SL: 7.1 mm, MD: 2.4 mm, Niederleis (Austria), figs 7 I 1 – I 2.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFE2FFCA8C92FF39FD9904DF.taxon	description	Description. Shell medium-sized, moderately slender fusiform, with weakly gradate spire. Protoconch unknown. Teleoconch of 4.5 weakly convex to subcylindrical whorls. Axial sculpture of orthocline, regularly spaced axial ribs on first teleoconch whorl, separated by narrower interspaces. Spiral sculpture consisting of four narrow spiral cords on first whorl, increasing in number to up to nine cords on last spire whorl. Spiral cords most prominent in axial interspaces, forming indistinct nodes at intersections with axial ribs. Axial ribs predominate sculpture on spire whorls and weaken on last whorl. Axial and spiral sculpture resulting in densely spaced pattern of blurred nodes. Last whorl subcylindrical above periphery, slowly contracting below. Spiral cords more prominent on base, weak fasciole. Aperture elongate, moderately narrow with indistinct anal sinus. Columella with four prominent folds. Outer lip thickened, with about 11 distinct denticles within; ten narrow lirae close behind peristome, distinctly separated from denticles. Siphonal canal moderately long, straight with shallow siphonal notch. Shell measurements and ratios. SL: 6.7 – 7.5 mm, MD: 2.4 – 2.6 mm, AA: 35 – 39 °, SL / MD: 2.9 – 3.0, AL / AW: 4.5 – 5.2, AH / S: 2.1 – 2.2.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFE2FFCA8C92FF39FD9904DF.taxon	discussion	Discussion. Some specimens from the Vienna and Pannonian basins differ slightly from Bellardithala dacica nov. sp. in having inner lirae or denticles present in all specimens seen, whereas they are not present in the specimen of B. laubei (Hoernes & Auinger, 1880) at hand, nor in specimen of B. dacica nov. sp. However, in other congeners, these inner teeth or lirae are only seen in a small proportion of specimens, and therefore we hesitate to use their presence / absence as a species-specific character. Sculpture is similar to that of B. dacica, possibly on the stronger end of the range of variability, but again insufficiently consistent to separate the two. Although probably just a local variety, we leave these specimens in open nomenclature, and highlight the similarity and differences with typical B. dacica.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFE2FFCA8C92FF39FD9904DF.taxon	distribution	Distribution in Central Paratethys. Badenian (middle Miocene): Vienna Basin: Niederleis (Austria);? Pannonian Basin: Herend (Hungary).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFE2FFC58C92FAB3FECF02F1.taxon	description	Figs 8 D 1 – D 2, E 1 – E 2, F	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFE2FFC58C92FAB3FECF02F1.taxon	materials_examined	Type material. Lectotype: SMF 360340 (= SMF XII. 2214 a), SL: 5.7 mm, MD: 2.1 mm, designated and illustrated by Zilch (1934, pl. 17, fig. 13), Senckenberg Museum, Frankfurt / Main, Germany, Coşteiu de Sus (Romania), figs 8 D 1 – D 2. Paralectotypes: 28 specimens, SMF XII 2214 b, Senckenberg Museum, Frankfurt / Main, Germany, Coşteiu de Sus (Romania). Additional material. NHMW 2020 / 0064 / 0001, SL: 5.8 mm, MD: 2.3 mm, Lăpugiu de Sus (Romania), figs 8 E 1 – E 2; NHMW 2020 / 0064 / 0002, SL: 5.5 mm, MD: 2.3 mm, Lăpugiu de Sus (Romania), fig. 8 F. Revised description. Shell small, moderately broad fusiform with conical to slightly cyrtoconoid spire. Protoconch poorly preserved, high conical with three regularly convex whorls. Teleoconch of 4.5 weakly convex to nearly straight-sided whorls with periphery at abapical suture, separated by incised suture. Axial sculpture of densely spaced, orthocline axial ribs, crossed by densely spaced spiral cords, forming delicate, spirally elongated pits in the interspaces. Spiral sculpture most prominent in axial interspaces, but strongly reduced on axial ribs. Last whorl moderately constricted at base. Aperture moderately narrow, with indistinct anal sinus. Columellar callus narrow. Columella with four columellar folds, weakening rapidly abapically. Outer lip weakly thickened, bearing about nine prominent denticles within. Siphonal canal moderately short, bent slightly left, with shallow siphonal notch. Shell measurements and ratios. SL: 5.5 – 5.8 mm, MD: 2.1 – 2.3, mm, AA: 26 – 43 °, SL / MD: 2.6 – 2.8, AL / AW: 3.4 – 5.0, AH / S: 2.4 – 2.6.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFE2FFC58C92FAB3FECF02F1.taxon	discussion	Discussion. Bellardithala kostejana (Boettger, 1906) can be separated from all its Paratethyan congeners in having a strongly oblique suture. It is superficially similar to B. partschi (Hörnes, 1852), which led Boettger (1906) to described it as variety of the latter. However, it differs in its broader, regularly conical spire, its less incised suture, its finer sculpture, comprising a higher number of axial ribs and spiral cords, and the absence of a prominent subsutural spiral cord, as well as more oblique suture. In addition, B. kostejana is constantly smaller. The occurrence of B. kostejana in offshore marls indicates also an environmental separation from B. partschi, which is restricted to near shore occurrences. Hölzl (1973: 93) mentioned Thala cf. partschi (M. Hoernes) from the Ottnangian (middle Burdigalian) of Gernergraben in Bavaria (without illustration). This would be the oldest record for this group from the Paratethys Sea. We have not seen this specimen but the generally poor preservation of the material from Gernergraben usally does not allow unambiguous identifications. Palaeoenvironment. The assemblages from Coşteiu de Sus suggests middle neritic palaeoenvironments with soft bottom (own data).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFE2FFC58C92FAB3FECF02F1.taxon	distribution	Distribution in Central Paratethys. Badenian (middle Miocene): Făget Basin: Coşteiu de Sus, Lăpugiu de Sus (Romania).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFEDFFC48C92FCECFDF301D1.taxon	description	Figs 9 A 1 – A 2, B 1 – B, C 1 – C 2 [Callithea] Lapugyensis nov. form. — Hoernes 1880: 125 [nomen nudum]. * Mitra Lapugyensis nov. form. — Hoernes & Auinger 1880 (pars): 89, pl. 10, figs 21 a – c [non pl. 10., figs 22 a – 22 c = Bellardithala fedosovi nov. sp.].	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFEDFFC48C92FCECFDF301D1.taxon	materials_examined	Type material. Lectotype (designated herein): NHMW 1854 / 0035 / 0101, SL: 5.4 mm, MD: 2.0 mm, Lăpugiu de Sus (Romania), illustrated in Hoernes & Auinger (1880, pl. 10, figs 21 a – c), figs 9 A 1 – A 2. Paralectotypes: NHMW 2020 / 0053 / 0001, SL: 5.1 mm, MD: 2.0 mm, Lăpugiu de Sus (Romania), figs 9 B 1 – B 2; NHMW 2020 / 0053 / 0002, SL: 5.4 mm, MD: 2.1 mm, Lăpugiu de Sus (Romania), figs 9 C 1 – C 2; 2 specimens, NHMW 2020 / 0053 / 0003, 2 specimens, NHMW 2020 / 0053 / 0002, all Lăpugiu de Sus (Romania). Revised description. Shell small, moderately broad fusiform with weakly cyrtoconoid spire. Protoconch high conical with 2.7 regularly convex whorls (height: 500 μm, diameter: 510 μm). Transition to teleoconch marked by onset of axial sculpture. Teleoconch of 4.5 weakly convex whorls, with periphery close above abapical suture, separated by deeply incised suture. Axial sculpture of about 15 prominent axial ribs on first teleoconch whorl, separated by slightly narrower interspaces. Spiral sculpture consisting of indistinct spiral grooves separating four spiral rows of blurred nodules. Cancellate sculpture on later whorls, with axial ribs and interspaces of similar width, crossed by four broad, regularly spaced spiral cords on second teleoconch whorl and five on subsequent whorl. Spiral cords form low subquadrate nodules at intersection with axial ribs, and well defined grooves in interspaces. Abapically, spiral interspaces reduced to rectangular pits. Last whorl weakly convex, with weakly constricted base, bearing about 13 spiral cords; 7 spirals on weak fasciole. Aperture narrow, elongate, with indistinct anal sinus. Columellar callus very narrow, indistinct. Columella with four columellar folds, weakening abapically. Outer lip only slightly thickened, bearing eight small denticles within. Siphonal canal moderately long, moderately wide, straight or bent slightly left, with wide, shallow siphonal notch. Shell measurements and ratios. SL: 5.1 – 5.4 mm, MD: 2.0 – 2.1 mm, AA: 40 – 43 °, SL / MD: 2.5 – 2.7, AL / AW: 4.1 – 4.8, AH / S: 2.3 – 2.4.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFEDFFC48C92FCECFDF301D1.taxon	discussion	Discussion. Hoernes & Auinger (1880) based the description of Mitra lapugyensis on specimens from Lăpugiu de Sus (Romania) and from Steinebrunn (Austria). In our opinion they are not conspecific, and as first revisers we designate the specimen illustrated by Hoernes & Auinger (1880, pl. 10, fig. 21) from Lăpugiu de Sus as lectotype. The specimen from Steinebrunn illustrated by Hoernes & Auinger (1880, pl. 10, fig. 22) is described herein as Bellardithala fedosovi nov. sp. Bellardithala lapugyensis differs from congeneric Paratethyan species in its cancellate sculpture and small size. The specimen described by Friedberg (1928) from the Badenian of Ukraine differs from typical M. lapugyensis in its more slender outline and the narrower spiral cords. It is doubtful that it is conspecific with B. lapugyensis. Palaeoenvironment. Unknown; the collections from Lăpugiu de Sus indicate a mixing of coastal-inner neritic and middle neritic elements.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFEDFFC48C92FCECFDF301D1.taxon	distribution	Distribution in Central Paratethys. Badenian (middle Miocene): Korytnica Basin: Korytnica (Poland) (Bałuk 1997),? Ukrainian Fore-Carpathian Basin: Zborów (Zboriv) (Ukraine) (Friedberg 1928); Făget Basin: Lăpugiu de Sus (Romania) (Hoernes & Auinger 1880).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFECFFC78C92FD8CFC170219.taxon	description	Figs 4 D, 9 D 1 – D 2, E 1 – E 2, F 1 – F 2	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFECFFC78C92FD8CFC170219.taxon	materials_examined	Type material. Lectotype (designated herein): NHMW 1846 / 0037 / 0113, SL: 6.9 mm, MD: 2.5 mm, Steinebrunn (Austria), illustrated in Hörnes (1852 b: pl. 10, figs 33 a – c), figs 9 D 1 – D 2. Paralectotypes: NHMW 1855 / 0045 / 0390, SL: 7.5 mm, MD: 2.4 mm, Steinebrunn (Austria), illustrated in Hoernes & Auinger (1880, pl. 19, fig. 15); NHMW 2020 / 0060 / 0004, SL: 6.5 mm, MD: 2.3 mm, Steinebrunn (Austria), illustrated in Hoernes & Auinger (1880, pl. 19, fig. 16), figs 9 E 1 – E 2; NHMW 1860 / 0001 / 0110, SL: 6.9 mm, MD: 2.4 mm, Steinebrunn (Austria), illustrated in Hoernes & Auinger (1880, pl. 19, fig. 18), figs 9 F 1 – F 2; NHMW 2020 / 0060 / 0003, SL: 6.5 mm, MD: 2.3 mm, Steinebrunn (Austria), fig. 4 D; NHMW 1855 / 0011 / 0407, Steinebrunn (Austria), illustrated in Hoernes & Auinger (1880, pl. 19, fig. 17). Additional material. NHMW 2020 / 0060 / 0001, 968 specimens, Steinebrunn (Austria), NHMW 2020 / 0060 / 0002, 382 specimens, Steinebrunn (Austria), NHMW 2020 / 0060 / 0003, 315 specimens Steinebrunn (Austria). Revised description. Shell medium-sized, moderately slender fusiform, with high conical to faintly cyrtoconoid, weakly scalate spire. High conical protoconch (fig. 4 D) comprising 2.7 weakly convex whorls (height: 500 μm, diameter: 550 μm). Teleoconch of five subcylindrical to weakly convex whorls, separated by incised suture. First teleoconch whorl with prominent axial ribs, crossed by three narrow spiral cords, forming indistinct nodules at intersections. Axial sculpture remains dominant on later whorls and especially on last whorl; spiral cords often strongly reduced forming four to five spiral rows of indistinct nodules on spire whorls, and up to ten cords on last whorl. Last whorl weakly convex to subcylindrical, slowly contracting below periphery. Aperture elongate, moderately narrow with indistinct anal sinus. Columellar callus indistinct. Columella with four columellar folds; adapical two columellar folds prominent, third one weak and fourth abapical fold strongly reduced. Outer lip only weakly thickened, with about five small denticles within, sometimes strongly reduced. Siphonal canal moderately long, straight with shallow siphonal notch. Shell measurements and ratios. SL: 6.5 – 7.5 mm, MD: 2.3 – 2.5 mm, AA: 25 – 37 °, SL / MD: 2.8, AL / AW: 4.8 – 5.6, AH / S: 2.2 – 2.4.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFECFFC78C92FD8CFC170219.taxon	discussion	Discussion. Bellardithala partschi (Hörnes, 1852) is characterised by its weakly scalate spire, strongly predominant axial sculpture, and flattened, relatively broad spirals separated by narrow grooves. It is most similar to B. lapugyensis (Hoernes & Auinger, 1880), but that species is smaller, with rounded spire whorls and more evenly cancellate sculpture. Bellardithala partschi is rather variable concerning slenderness and convexity of the whorls. Similarly, the spiral sculpture varies considerably and is reduced in many specimens. Intercalations of secondary spiral grooves may raise the number of spiral rows to six or seven on the penultimate whorl. Palaeoenvironment. This species is very common in shallow water settings, partly indicating sea grass meadows (e. g. Steinebrunn, Gainfarn; Zuschin et al. 2007, own data).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFECFFC78C92FD8CFC170219.taxon	distribution	Distribution in Central Paratethys. Badenian (middle Miocene): Nowy Sącz Basin: Niskowa (Poland) (Bałuk 1970); Voronyaky Hills: Podhorce (Pidhirtsi) (Ukraine) (Friedberg 1911); Ukrainian Fore-Carpathian Basin: Zborów (Zboriv) (Ukraine) (Friedberg 1928); Vienna Basin: Niederleis, Gainfarn, Pötzleinsdorf / Vienna, Steinebrunn (Austria) (Sieber 1953, 1958 b), Mikulov (Czech Republic); Pannonian Basin: Herend (Hungary) (Kókay 1966); Eisenstadt-Sopron Basin: Forchtenau (Austria) (Sieber 1956).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFEEFFC68C92FF39FA1406E4.taxon	description	Figs 8 G	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFEEFFC68C92FF39FA1406E4.taxon	materials_examined	Material. Inv. Z 23408 - P 2003 / 27, Natural History Museum in Bratislava, SL: 9.4 mm, MD: 3.3 mm.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFEEFFC68C92FF39FA1406E4.taxon	description	Description. Shell medium-sized, moderately slender with slightly gradate spire. Protoconch and first teleoconch whorls unknown. Early teleoconch whorls with broad, strongly opisthocline axial ribs, with interspaces of similar width. Axial ribs becoming sigmoidal and densely beaded on later whorls, with two more spiral rows of slightly more prominent beads along adapical suture. About eight spiral rows of beads on penultimate whorl. Beads replaced by prominent, convex spiral cords on strongly constricted base. Aperture is largely destroyed with three prominent columellar folds and much weaker abapical fourth fold. Siphonal canal long.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFEEFFC68C92FF39FA1406E4.taxon	discussion	Discussion. Tomašových (1998) described a single fragmentary specimen of a Bellardithala from basinal clays at Devínska Nová Ves. Due to its peculiar sculpture of strongly opisthocline to sigmoidal, beaded axial ribs, this specimen differs from all other Paratethyan Bellardithala species and might represent a yet undescribed species. Due to its fragmentary preservation, we refrain from naming it formally. Palaeoenvironment. The mollusc assemblage of Devínska Nová Ves indicates deeper neritic environments (Tomašových 1998).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFEEFFC68C92FF39FA1406E4.taxon	distribution	Distribution in Central Paratethys. Badenian (middle Miocene): Vienna Basin: Devínska Nová Ves (Slovakia). Genus Ebenomitra Monterosato, 1917	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFEEFFC68C92FF39FA1406E4.taxon	type_taxon	Type species. Mitra ebenus Lamarck, 1811, subsequent designation by Coan (1966: 130); Present-day, Mediterranean Sea.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFEEFFC68C92FF39FA1406E4.taxon	description	Description. “ Shell small, turriform or fusiform to widely fusiform. Protoconch bulbous, paucispiral, with about two glossy whorls. Teleoconch of up to 7.5 moderately or strongly convex whorls. Early spire whorls with strong ribs or very strong and wide folds; later whorls with strong wide folds vanishing below periphery of last adult whorl, or smooth; spiral sculpture not pronounced. Siphonal canal short to moderately long, sculptured with between two and four strong oblique cords. Aperture ovate to elongate, sometimes constricted towards siphonal canal; outer aperture lip lirate within. Inner apertural lip with three strong oblique columellar folds, adapicalmost fold strongest ” (Fedosov et al., 2017: 601).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFEEFFC68C92FF39FA1406E4.taxon	discussion	Discussion. Fedosov et al. 2017 considered Ebenomitra as a subgenus of Pusia. The pros and cons of the usage of subgenera was controversially discussed in a plethora of papers (e. g. Laurin 2010; Puillandre et al. 2014; Teta 2018 and references therein). As stated in our previous contributions (Harzhauser & Landau 2016, 2019) we consider supraspecific classification levels as arbitrary and try to avoid the usage of subgenera. Therefore, we prefer to raise Ebenomitra to full genus rank.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFEEFFC08C92F899FC2A0795.taxon	description	Figs 4 E, 10 A 1 – A 2, B 1 – B 2, C 1 – C 2, 11 A 1 – A 2	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFEEFFC08C92F899FC2A0795.taxon	description	Mitra leucozona Andrzeiowski — Pusch 1837: 119, pl. 11, figs 6 a – b.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFEEFFC08C92F899FC2A0795.taxon	description	V [exillum]. (V [exillum].) ebenus leucozona (Andrz.) — Sieber 1958 a: 153.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFEEFFC08C92F899FC2A0795.taxon	materials_examined	Type material. Syntype described and illustrated by Andrzejowski (1830, pl. 4, figs 6 a – b) from Szuszkowce (Shushkivtsi) (Ukraine). The whereabouts of the Andrzejowski collection are unknown (Daszkiewicz & Bauer, 2008). Additional material. NHMW 1846 / 0037 / 0110, SL: 21.0 mm, MD: 8.9 mm, Gainfarn (Austria), illustrated in Hörnes (1852 b, pl. 10, fig. 11), figs 10 B 1 – B 2. NHMW 1868 / 0001 / 0401, SL: 16.7 mm, MD: 8.2 mm, Lăpugiu de Sus (Romania), illustrated in Hoernes & Auinger (1880, pl. 7, figs 11 a – b, figs 10 C 1 – C 2. NHMW 1870 / 0033 / 0034, SL: 13.8 mm, MD: 6.3 mm, Lăpugiu de Sus (Romania), illustrated in Hoernes & Auinger (1880, pl. 7, figs 12 a – b), figs 10 A 1 – A 2. NHMW 1876 / 0011 / 0008, SL: 13.5 mm, MD: 6.4 mm, Lăpugiu de Sus (Romania), figs 4 E; NHMW 1854 / 0035 / 0099, SL: 23.4 mm, MD: 9.8 mm, Lăpugiu de Sus (Romania), illustrated in Hoernes & Auinger (1880, pl. 7, figs 10 a – b); NHMW 1857 / 0014 / 0008, SL: 14.0 mm, MD: 6.9 mm, Steinebrunn (Austria), figs 11 A 1 – A 2. NHMW 2020 / 0065 / 0001, SL: 14.5 mm, MD: 6.6 mm, Ternopil (Ukraine), illustrated in Hoernes & Auinger (1880, pl. 7, figs 14 a – b). NHMW 1857 / 0014 / 0008, 12 specimens; NHMW 2020 / 0066 / 0001, 3 specimens, Steinebrunn (Austria); NHMW 2020 / 0083 / 0001, 4 specimens, Baden (Austria); NHMW 1872 / 0030 / 0022, 1 specimen, NHMW 1866 / 0001 / 0626, 1 specimen, Baden-Soo (Austria); NHMW 1997 z 0178 / 1106, 20 specimens, Bad Vöslau (Austria); NHMW 2020 / 0068 / 0001, 6 specimens, NHMW 1997 z 0178 / 1429, 17 specimens, Gainfarn (Austria); NHMW 2020 / 0067 / 0001, 1 specimen, Grund (Austria); NHMW 2020 / 0070 / 0001, 36 specimens, Vienna / Pötzleinsdorf (Austria); NHMW 1861 / 0001 / 0235, 5 specimens, Pöls (Austria); NHMW 1863 / 0015 / 0293, 2 specimens, Szob (Hungary); NHMW 1858 / 0047 / 0004, 1 specimen, Sedlec (= Porzteich) (Czech Republic); NHMW 1851 / 00010 / 0016, 2 specimens, Mikulov (Czech Republic); NHMW 1876 / 0011 / 0008, 46 specimen, Lăpugiu de Sus (Romania). Revised description. Shell moderately large, solid, broadly biconic, with moderately high, gradate spire. Protoconch (fig. 4 E) paucispiral, with low nucleus and about two moderately convex whorls (height: 550 μm, diameter: 550 μm). Transition to teleoconch not well preserved. Teleoconch of about eight whorls; early spire whorls straight sided, with convex, orthocline axial ribs, separated by slightly narrower interspaces. Ribs and interspaces wider on fourth to sixth teleoconch whorls, ribs broad, poorly delimited on last whorl (eight to ten on last whorl) causing faint convexity of whorls. Suture impressed, undulating. Last whorl short, barrel-shaped to faintly convex, moderately constricted at base. Aperture wide, with shallow, narrow anal canal. Columellar callus narrow, indistinct. Columella with three folds, fourth fold subobsolete; adapical fold most prominent, sharp. Outer lip thin, without denticles; about 10 delicate long lirae inside aperture. No spiral sculpture except for weak spiral threads on base and more prominent spiral cords on fasciole, extending from columellar folds. Siphonal canal short, straight or bent weakly to left, with shallow siphonal notch. Colour pattern frequently preserved, consisting of orange-brown background and white band along abapical suture on spire whorls, continuing above mid-whorl on last whorl; two narrower white bands along base and fasciole. Shell measurements and ratios. SL: 12.0 – 21.2 mm, MD: 5.5 – 9.5 mm; AA: 45 – 52 °, SL / MD: 2.2 – 2.3, AL / AW: 2.1 – 2.8, AH / S: 3.4 – 3.6.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFEEFFC08C92F899FC2A0795.taxon	discussion	Discussion. Eichwald (1830, 1840, 1852, 1853) described this species as Mitra striata although he was aware that it was identical to M. leucozona Andrzejowski, 1830 (e. g. Eichwald 1840: 12). Andrzejowski’s paper, however, appeared earlier than Eichwald’s paper (Nosowska 2020) and M. leucozona takes precedence. Ebenomitra leucozona is a very characteristic species due to its subcylindrical last whorl bearing broad, poorly delimited ribs. Generally, it displays only moderate variability concerning size, slenderness and axial ribbing on the last whorl. Therefore, it is surprising that this species was mixed by Eichwald (1840, 1853), Hörnes (1852 b) and Hoernes & Auinger (1880) with smooth and slender shells of other species such as Ebenomitra pseudopyramidella (Boettger, 1906) and Pusia avellanella (Boettger, 1906). The large, barrel-shaped, smooth specimen from Lăpugiu de Sus (Romania), illustrated in Hoernes & Auinger (1880, pl. 7, figs 10 a – b), seems to be an aberrant specimen of E. leucozona based on its identical colour pattern and columellar folds. The specimens from the Badenian of Węglinek i Łychów (Poland), described by Krach (1981, pl. 20, figs 1 – 5), are not conspecific with E. leucozona. They are relatively slender with high spire whorls, bear well defined axial ribs and have an almost smooth last whorl. They are slightly reminiscent of Pusia pseudorecticosta (Boettger, 1906) but are less elongate. These specimens might represent an undescribed species. The specimens described by Davoli (2000) from the Tortonian of Montegibbio in Italy differ in their broader outline and wider apical angle (e. g. figs 27 – 30) and are most probably not conspecific with E. leucozona. Similarly, the specimen from the middle Miocene of the Loire Basin (France) illustrated by Glibert (1952, pl. 12, figs 2 a – b) might represent a separate species. Iljina (1993) mentioned this species also from the Konkian (middle Miocene) of Agara (Georgia) and the Ustyurt plateau, which would expand the distribution area also into the Eastern Paratethys Sea. The illustrated specimens, however, differ clearly from Ebenomitra leucozona (Andrzejowski, 1830) in their slender outline, convex spire whorls blunt axial ribs. It represents a yet undescribed Ebenomitra species. Palaeoenvironment. This species is most abundant at localities suggesting silty-sandy inner neritic environments partly with sea grass meadows (e. g. Gainfarn, Steinebrunn) (Zuschin et al. 2007).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFEEFFC08C92F899FC2A0795.taxon	distribution	Distribution in Central Paratethys. Korytnica Basin: Korytnica (Poland) (Bałuk 1997); Roztocze Hills: Łychów (Poland) (Bałuk 1997); Nowy Sącz Basin: Niskowa (Poland) (Bałuk 1970); Voronyaky Hills: Podhorce (Pidhirtsi) (Ukraine) (Friedberg 1911); Ukrainian Fore-Carpathian Basin: Dryszczów (Nadrichne), Tarnoruda (Tarnoruda), Ternopol (Ternopil), Żabiak, Zaleśce (Salisze), Zborów (Zboriv), Żukowce (Zhukivtsi) (Ukraine) (Friedberg 1911); North Alpine Carpathian Foredeep: Grund, Windpassing (Sieber 1947, 1953); Vienna Basin: Baden, Gainfarn, Enzesfeld, Vienna / Pötzleinsdorf, Steinebrunn (Austria), Mikulov (Czech Republic), Borský Mikulás (Slovakia) (Hoernes & Auinger 1880; Švagrovský 1982); Eisenstadt-Sopron Basin: Forchtenau (Austria) (Sieber 1956); Pannonian Basin: Sámsonháza, Letkés, Illés-Street Budapest (Hungary); Krka Basin: Stara vas pri Šentjerneju (Slovenia) (Mikuž 2009); Mehadia Basin (Romania) (Hinculov, 1968); Zarand Basin: Minişul de Sus (Romania) (Nicorici & Sagatovici 1973); Făget Basin: Lăpugiu de Sus (Romania); Dacian Basin: Staropatica, Târnene (Bulgaria) (Kojumdgieva in Kojumdgieva & Strachimirov 1960).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFEBFFFD8C92FF39FE1B0449.taxon	description	Figs 4 F, 10 D 1 – D 2, E 1 – E 2, F 1 – F 2, 11 B 1 – B 2 V. [oluta] laevis n. — Eichwald 1829: 297, pl. 5, fig. 14 [non Donovan, 1804].	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFEBFFFD8C92FF39FE1B0449.taxon	description	Mitra laevis Dubois — Pusch 1837: 119. [Mitra incognita] var. β laevis m. — Pusch 1837: 119, pl. 11, fig. 5.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFEBFFFD8C92FF39FE1B0449.taxon	materials_examined	Type material. Holotype: NHMW 2020 / 0071 / 0001, SL: 16.9 mm, MD: 6.3 mm, Ternopil (Ukraine), illustrated in Hoernes & Auinger (1880, pl. 7, fig. 13), figs 10 E 1 – E 2. Additional material. NHMW 2020 / 0071 / 0002, SL: 17.0 mm, MD: 6.5 mm, Ternopil (Ukraine), figs 10 F 1 – F 2; NHMW 2020 / 0071 / 0003, SL: 15.8 mm, MD: 6.1 mm, Ternopil (Ukraine), figs 10 D 1 – D 2; NHMW 2020 / 0071 / 0004, SL: 16.1 mm, MD: 5.6 mm, Ternopil (Ukraine), figs 11 B 1 – B 2; NHMW 1857 / 0014 / 0009, SL: 16.5 mm, MD: 6.1 mm, Steinebrunn (Austria), fig. 4 F; NHMW 2020 / 0071 / 0005, 8 specimens, Ternopil (Ukraine); NHMW 1859 / 0040 / 0006, 1 specimen. Zaleśce (Ukraine); NHMW 1857 / 0014 / 0009, 51 specimens, Steinebrunn (Austria). Revised description. Shell moderately large, solid, glossy, moderately broad fusiform, with high, conical to weakly cyrtoconoid spire. Protoconch (fig. 4 F) paucispiral, with low nucleus and about two weakly convex whorls (height: 540 μm, diameter: 530 μm) (note that transition to teleoconch is abraded in available specimen overemphasizing the weak convexity of the last protoconch whorl). Teleoconch of about eight weakly convex to subcylindrical whorls, with periphery at or slightly above abapical suture. Suture narrowly impressed, slightly undulating. Axial sculpture on early teleoconch whorls consisting of low, convex axial ribs, separated by narrower interspaces. Axial sculpture fading out within fourth to fifth whorl, rarely persisting on penultimate whorl. Last whorl moderately contracting. Aperture wide, ovoid, with indistinct anal sinus. Columellar callus narrow, sharply delimited. Columella with three oblique folds; adapical two folds most prominent. Outer lip with 12 – 15 lirae placed distinctly behind peristome; lirae more prominent at their adapertural termination, fading out inside. Outer lip weakly thickened at lirae, thinning towards peristome. Siphonal canal moderately wide, short, straight or weakly bent to left. Spiral sculpture absent, except for few poorly defined spiral cords on weak fasciole; most prominent spiral cord extending from adapical columellar fold. Colour pattern frequently preserved, consisting of orange-brown background with narrow white band slightly below mid-whorl on spire whorls and above periphery on last whorl. Shell measurements and ratios. SL: 12.6 – 17.0 mm, MD: 4.5 – 6.5 mm; AA: 38 – 42 °, AA: 45 – 52 °, SL / MD: 2.5 – 2.7, AL / AW: 3.2 – 3.5, AH / S: 3.1 – 3.4.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFEBFFFD8C92FF39FE1B0449.taxon	discussion	Discussion. Eichwald (1829, 1853) described Voluta laevis from Biłka, Poczajów Stary, Żukowice, Tarnoruda and Zaleśce in Ukraine and Hilber (1882) added material from Hołubica (Ukraine). Boettger (1906) introduced Mitra pseudopyramidella as new name for a specimen described by Hoernes & Auinger (1880 pl. 7, fig. 13) from Ternopil, situated close to the other Ukrainian localities. The shells from Ternopil are clearly conspecific with ‘ Mitra laevis ’ specimens from Hołubica, Żukowice and Zaleśce in the NHMW and GBA collections. Therefore, we treat both taxa as synonyms. Voluta laevis is preoccupied by Donovan (1804) and therefore, Mitra pseudopyramidella Boettger, 1906 is the next available name. Pusch (1837) discussed this species as Mitra turgidula, referring to Voluta turgidula Brocchi, 1814 from the Pliocene of Italy. Thus, although he added “ m ” (for mihi) when describing the species, he did not intend to establish it as new species. Ebenomitra pseudopyramidella differs from Pusia avellanella (Boettger, 1906) in its larger size, glossy shell, and absence of a fourth columellar fold. Ebenomitra sublaevis Landau, Ceulemans & Van Dingenen, 2019, from the Tortonian of France, is highly reminiscent of E. pseudopyramidella, but differs in its smaller size and stout outline (see Landau et al., 2019, pl. 52, figs 1 – 4).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFEBFFFD8C92FF39FE1B0449.taxon	distribution	Distribution in Central Paratethys. Badenian (middle Miocene): Voronyaky Hills: Hołubica (Holubytsia) (Ukraine) (Friedberg 1911); Ukrainian Fore-Carpathian Basin: Dryszczów (Nadrichne), Tarnoruda (Tarnoruda), Zaleśce (Salisze) (Ukraine), Zborów (Zboriv), Żukowce (Zhukivtsi) (Ukraine) (Friedberg 1911); Vienna Basin: Steinebrunn (Austria).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFD5FFFC8C92FB3CFBF20284.taxon	description	Figs 10 G 1 – G 2, H 1 – H 2, I 1 – I 2, J	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFD5FFFC8C92FB3CFBF20284.taxon	materials_examined	Type material. Lectotype (designated herein): ZNG PAN A-I- 50 / 35, SL: 14.0 mm, MD: 5.1 mm, Zboriv (Zborów) (Ukraine), stored in the Geological Museum of the Institute of Geological Sciences, Polish Academy of Sciences, illustrated in Friedberg (1928, pl. 37, fig. 25), figs 10 G 1 – G 2. Paralectotypes: ZNG PAN A-I- 50 / 36.1 – 36.6, SL: 15.2 mm, MD: 5.1 mm, Zboriv (Zborów) (Ukraine), stored in the Geological Museum of the Institute of Geological Sciences, Polish Academy of Sciences, figs 10 H 1 – H 2; ZNG PAN A-I- 50 / 36.1 – 36.6, SL: 9.1 mm, MD: 3.5 mm, Zboriv (Zborów) (Ukraine), stored in the Geological Museum of the Institute of Geological Sciences, Polish Academy of Sciences, figs 10 I 1 – I 2; ZNG PAN A-I- 50 / 36.1 – 36.6, SL: 12.1 mm, MD: 4.8 mm, Zboriv (Zborów) (Ukraine), stored in the Geological Museum of the Institute of Geological Sciences, Polish Academy of Sciences, fig. 10 J. Revised description. Shell moderately large, moderately slender pupiform, with high, weakly gradate spire. Protoconch paucispiral with broadly convex, low nucleus and about one weakly convex whorl. Transition to teleoconch indistinct in available specimens. Teleoconch of six glossy, weakly convex whorls, rapidly increasing in height, with periphery in abapical third, separated by deeply impressed, narrowly canaliculated suture. No axial sculpture on spire whorls. Last whorl weakly convex to subcylindrical, moderately constricted at short base, bearing subobsolete axial ribs, most prominent along periphery and base. Aperture wide, ovoid, anal canal indistinct with weak parietal denticle. Columella with three to four columellar folds. Adapical two folds prominent, abapical folds weak, fourth fold only present in one specimen. Outer lip thin, with about 12 long, delicate lirae within. Siphonal canal short, straight or slightly bent to left, with wide siphonal notch. Shell measurements and ratios. SL: 9.1 – 15.2 mm, MD: 3.5 – 5.1 mm; AA: 25 °, AA: 45 – 52 °, SL / MD: 2.9 – 3.1, AL / AW: 3.0 – 3.4, AH / S: 3.0.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFD5FFFC8C92FB3CFBF20284.taxon	discussion	Discussion. We place this rare species in Ebenomitra based on its paucispiral protoconch, which excludes a placement in Pusia (see Fedosov et al. 2017). Nevertheless, the high spire and canaliculate suture are atypical for Ebenomitra and the occasional presence of a fourth columellar fold is also problematic. The species is somewhat reminiscent of Ebenomitra pseudopyramidella (Boettger, 1906), but differs in its higher spire, more slender profile, deeper suture, and lacks axial sculpture on early spire whorls. The same features allow a separation from Pusia avellanella (Boettger, 1906), which is similar concerning the axial ribs along the periphery, but that species is immediately separated by its multispiral protoconch, which places it in the genus Pusia Swainson, 1840, rather than Ebenomitra. Palaeoecology. The mollusc assemblage from the type locality suggest shallow marine near-shore environments (own data).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFD5FFFC8C92FB3CFBF20284.taxon	distribution	Distribution in Central Paratethys. Badenian (middle Miocene): Korytnica Basin: Korytnica (Poland) (Bałuk 2006); Ukrainian Fore-Carpathian Basin: Zborów (Zboriv), (Ukraine) (Friedberg 1928).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFD4FFFC8C92FD78FB700770.taxon	type_taxon	Type species. Mitra (Callithea) fuchsi Hoernes & Auinger, 1880, middle Miocene, Paratethys Sea, Romania.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFD4FFFC8C92FD78FB700770.taxon	diagnosis	Diagnosis. Large, moderately broad fusiform shell, with beaded sculpture, differing from Thala in the development of a parietal denticle and the less cyrtoconoid spire. Protoconch paucispiral, of weakly convex, smooth whorl. Teleoconch whorls with dense pattern of axial ribs and spiral cords forming densely spaced ‘ pillow-like’, subquadrate nodes at intersections. Axial ribs of early teleoconch whorls of type species bearing three spiral rows of nodes with tiny spines on middle spiral row [not preserved in F. zibinica]. Aperture moderately narrow to wide, with three columellar folds; adapical two folds very prominent. Columellar callus narrow, abapically contracting, no lirae inside aperture. Siphonal canal moderately long, straight or slightly bent to left. No signs of episodic growth.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFD4FFFC8C92FD78FB700770.taxon	description	Description. As for type species.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFD4FFFC8C92FD78FB700770.taxon	etymology	Etymology. In honour of Alexander Fedosov (Russian Academy of Sciences, Moscow), in gratitude for the discussions on Paratethyan Costellariidae. Included species. Mitra (Callithea) fuchsi Hoernes & Auinger, 1880, middle Miocene, Romania; Mitra zibinica Toldo, 1889, late Miocene, Italy.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFD4FFFC8C92FD78FB700770.taxon	description	Stratigraphic and geographic range. Middle Miocene (Badenian) deposits of the Central Paratethys Sea and late Miocene of the proto-Mediterranean Sea. Palaeoenvironment. Unknown; most probably from middle neritic depositional environments with pelitic bottom (based on sediment from the aperture).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFD4FFFC8C92FD78FB700770.taxon	discussion	Discussion. Fedosovia differs from other costellariid genera by the combination of a peculiar ‘ pillow-like’ sculpture and a paucispiral protoconch (as opposed to Ebenomitra, which has a paucispiral protoconch, but lacks the peculiar sculpture). Protoconch of paucispiral or multispiral type is not a useful distinguishing character in many gastropod groups. However, amongst the Costellariidae, Fedosov et al. (2017) placed importance at supraspecific level to protoconch type in many of their genera, and specified the protoconch type in his generic descriptions. We follow this lead here and consider the protoconch type a genus-specific character in Fedosovia. The paucispiral protoconch excludes a relation with Vexillum, although the extant Vexillum piceum (Pease, 1860), from Réunion and Hawaii, develops a comparable sculpture (but is much smaller, SH <10 mm). A paucispiral protoconch occurs in the extant Atlantilux Huang, 2015 but Atlantilux species are characterised by a wide, slightly concave subsutural ramp and convex periphery close to the abapical suture. Atlantilux is also distinguished from Fedosovia by the sculpture of dense axial riblets in the subsutural area opposed by wide fold-like axial ribs along the periphery (see Fedosov et al., 2017). Fedosovia is superficially reminiscent of Costapex Fedosov, Herrmann & Bouchet, 2017 in shape and sculpture. A closer relationship with this extant genus, however, is unlikely in respect to the paucispiral protoconch and the presence of only three columellar folds instead of four.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFD7FFFE8C92FF39FDB10662.taxon	description	Figs 4 G, 12 A 1 – A 2, B 1 – B 2, C 1 – C 2, D [Callithea] Fuchsi nov. form. — Hoernes 1880: 125 [nomen nudum].	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFD7FFFE8C92FF39FDB10662.taxon	materials_examined	Type material. Lectotype (designated herein): NHMW 1857 / 0024 / 0012, Lăpugiu de Sus (Romania), SL: 13.8 mm, MD: 5.0 mm, illustrated in Hoernes & Auinger (1880, pl. 10, figs 13 a – c), figs 4 G, 12 B 1 – B 2. Paralectotypes: NHMW 1999 z 0076 / 0003, Jaroměřice nad Rokytnou (Czech Republic), SL: 14.7 mm, MD: 5.4 mm, illustrated in Hoernes & Auinger (1880, pl. 10, figs 12 a – c), figs 12 A 1 – A 2; NHMW 2020 / 0086 / 0001, Lăpugiu de Sus (Romania), SL: 15.1 mm, MD: 5.2 mm, illustrated in Hoernes & Auinger (1880, pl. 10, figs 14 a – c); NHMW 2016 / 0177 / 0878, Lăpugiu de Sus (Romania), SL: 20.8 mm, MD: 7.3 mm, figs 12 C 1 – C 2; NHMW 2016 / 0177 / 0878, Lăpugiu de Sus (Romania), SL: 20.0 mm, MD: 7.5 mm, fig. 12 D. Revised description. Shell large, moderately broad to moderately slender fusiform. Protoconch (fig. 4 G) paucispiral, low conical, 1.7 smooth, weakly convex whorls (height: 540 μm, diameter: 560 μm). Teleoconch boundary marked by beginning of axial sculpture. Teleoconch of seven whorls, separated by impressed suture. First teleoconch whorl convex, with about ten blunt, convex, weakly opisthocline axial ribs, separated by narrow interspaces. Axial ribs bearing three spiral rows of nodules; nodules of middle row with tiny spines on first teleoconch whorl. Additional spiral rows intercalated on subsequent whorls. Six spiral rows on penultimate whorl, number may rise to eight by bifurcation. Axial and spiral interspaces very narrow, resulting in densely spaced, pillow-like, subquadrate nodes. Second to third teleoconch whorls weakly angulated slightly above mid-whorl. Later whorls straight-sided or weakly convex, with periphery at or slightly above abapical suture. Last whorl weakly convex to straight-sided, strongly constricted. Aperture ovoid, moderately narrow to wide, with narrow anal sinus. Columellar callus narrow, weakly delimited, bearing three columellar folds; adapical two folds very prominent. Outer lip thin, abapically contracting, no lirae inside aperture. Siphonal canal moderately long, narrow, straight or slightly bent to the left, with shallow siphonal notch. Shell measurements and ratios. SL: 13.8 – 20.8 mm, MD: 5.0 – 7.5 mm; AA: 38 – 43 °, SL / MD: 2.6 – 2.7, AL / AW: 3.8 – 4.3, AH / S: 2.2 – 2.8.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFD7FFFE8C92FF39FDB10662.taxon	discussion	Discussion. This species is unique within the Paratethyan Costellariidae due to its conspicuous sculpture of densely spaced nodules. Fedosovia zibinica (Toldo, 1889), from the late Miocene of Montegibbio (Italy) is a closely related species, but differs in its strongly convex whorls (see holotype in Davoli 2000, pl. 3, figs 18 a – b). Pusia textilosa Bellardi, 1888, from the early Miocene of Italy, is slightly reminiscent of the Paratethyan species concerning sculpture, but differs clearly in its gradate spire and short siphonal canal (see Ferrero-Mortara et al. 1981, pl. 53, figs 10 a – b). Palaeoenvironment. Unknown; most probably from middle neritic depositional environments with pelitic bottom (based on sediment from the aperture).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFD7FFFE8C92FF39FDB10662.taxon	distribution	Distribution in Central Paratethys. Badenian (middle Miocene): Carpathian Foreland Basin: Jaroměřice nad Rokytnou (Czech Republic); Făget Basin: Lăpugiu de Sus (Romania). Genus Pusia Swainson, 1840	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFD7FFFE8C92FF39FDB10662.taxon	type_taxon	Type species. Mitra microzonias Lamarck, 1811, by monotypy.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFD7FFFE8C92FF39FDB10662.taxon	description	Description. “ Shell small, fusiform to broadly fusiform or ovate, last adult whorl 64 – 74 % of shell height. Protoconch multispiral, narrowly conical, with three or more glossy whorls. Suture distinct, impressed. Sculpture of dense, rounded, axial ribs on early spire whorls (sometimes absent), turning into broad and low folds on last teleoconch whorls; spiral sculpture of fine regular grooves, pronounced in interstices between axials. Shell base sculptured with several broad and flattenned, somewhat gemmate, spiral cords. Siphonal canal short or very short, stout, bearing several strong oblique cords. Aperture narrow, elongate, its outer lip lirate within. Inner apertural lip with three or four strong columellar folds, adapicalmost fold strongest ” (Fedosov et al., 2017: 597).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFD7FFFE8C92FF39FDB10662.taxon	discussion	Discussion. The species grouped herein in Pusia are somewhat heterogeneous. While Pusia avellanella and Pusia paraleucozona fit well in the genus as defined by Fedosov et al. (2017), Pusia falsitranssylvanica and Pusia confunda are atypical due to their moderately high orthoconoid spires and distinct, crested (rather than low and rounded) axial ribs. Currently, no genus is available to place these species and provisionally we keep them in Pusia.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFD6FFF88C92F91DFA8803C9.taxon	description	Figs 4 H, 13 A 1 – A 2, B 1 – B 2, C 1 – C 2, D 1 – D 2, E 1 – E 2, F 1 – F 2	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFD6FFF88C92F91DFA8803C9.taxon	materials_examined	Type material. Lectotype: SMF 360343 (= XII 2209 a), Senckenberg Museum, Frankfurt / Main, Germany, SL: 4.75 mm, MD: 2.1 mm, Coşteiu de Sus (Romania), designated and illustrated in Zilch (1934, pl. 17, fig. 19), figs 13 A 1 – A 2. Paralectotypes: SMF XII 2209 b – c, Senckenberg Museum, Frankfurt / Main, Germany, Coşteiu de Sus (Romania). Additional material. SMF 360344 (= SMF XII 2210 a), SL: 6.5 mm, MD: 2.8 mm, Senckenberg Museum, Frankfurt / Main, Germany, Coşteiu de Sus (Romania), lectotype of Mitra (Uromitra) avellana pseudavellana Boettger, 1906, figs 13 D 1 – D 2; NHMW 2020 / 0072 / 0001, SL: 10.1 mm, MD: 3.1 mm, Gainfarn (Austria), figs 13 C 1 – C 2, 4 H; NHMW 2020 / 0072 / 0002, SL: 9.9 mm, MD: 4.0 mm, Gainfarn (Austria), figs 13 B 1 – B 2; NHMW 2020 / 0072 / 0003, SL: 9.2 mm, MD: 3.9 mm, Gainfarn (Austria), NHMW 2020 / 0072 / 0004, SL: 9.2 mm, MD: 3.) mm, Gainfarn (Austria), illustrated in Hoernes & Auinger (1880, pl. 9, figs 23 a – c), syntype of Mitra (Costellaria) miocaenica Dollfus & Dautzenberg 1886, figs 13 E 1 – E 2; NHMW 2020 / 0073 / 0001, SL: 11.2 mm, MD: 3.9 mm, Jerutek at Lysice (Czech Republic), illustrated in Hoernes & Auinger (1880, pl. 9, figs 24 a – c), syntype of Mitra (Costellaria) miocaenica Dollfus & Dautzenberg 1886; NHMW 1847 / 0037 / 0048, SL: 14.1 mm, MD: 5.4 mm, Steinebrunn (Austria), illustrated in Hoernes & Auinger (1880, pl. 9, figs 22 a – c), syntype of Mitra (Costellaria) miocaenica Dollfus & Dautzenberg 1886, figs 13 F 1 – F 2: NHMW 1859 / 0037 / 0006, SL: 11.9 mm, MD: 4.3 mm, Lăpugiu de Sus (Romania), illustrated in Hoernes & Auinger (1880, pl. 9, figs 25 a – c), syntype of Mitra (Costellaria) miocaenica Dollfus & Dautzenberg 1886; NHMW 1865 / 0015 / 0018, 8 specimens, Jerutek at Lysice (Czech Republic), NHMW 2020 / 0074 / 0001, 19 specimens, Steinebrunn (Austria), NHMW 1846 / 0037 / 0115, 24 specimens, Gainfarn (Austria), NHMW 2020 / 0072 / 0005, 50 specimens, Gainfarn (Austria), NHMW 2020 / 0075 / 0001, 3 specimens, Lăpugiu de Sus (Romania). ZNG PAN A-I- 50 / 72, syntype of Turricula intermittens R. Hoern. i Auinger var. dryszczoviensis Friedberg, 1911, illustrated in Friedberg (1911, pl. 1, fig. 19), Dryszczów (Ukraine), stored in the Geological Museum of the Institute of Geological Sciences, Polish Academy of Sciences). ZNG PAN A-I- 50 / 73.1 – 73.12, 12 specimens, syntypes of Turricula intermittens R. Hoern. i Auinger var. dryszczoviensis Friedberg, 1911, Dryszczów (Ukraine), stored in the Geological Museum of the Institute of Geological Sciences, Polish Academy of Sciences). Revised description. Shell moderately large, moderately broad to moderately slender fusiform, with high spire. Protoconch (fig. 4 H) high conical, mammillate, consisting of 3.5 convex whorls (height: 800 μm, diameter: 650 μm). Protoconch axis tilted from shell axis in many specimens. Teleoconch of eight weakly convex to subcylindrical whorls, with periphery at abapical suture, separated by narrowly impressed suture. Axial sculpture of 15 – 18 closeset orthocline, convex ribs, separated by slightly narrower interspaces; axials very variable, typically weakening on early spire whorls or rarely persisting over entire spire. Last whorl subcylindrical or weakly convex, with rounded shoulder placed close below adapical suture, rapidly contracting, bearing reduced axial sculpture, often more distinct along basal convexity fading out quickly towards shoulder and abapically on basal concavity. Aperture moderately narrow, ovoid with indistinct, narrow anal sinus, faint parietal denticle. Columella narrow, sharply delimited. Columella with four folds, weakening rapidly abapically. Outer lip thin, with about ten prominent, long lirae within. No spiral sculpture except for few spiral cords on weak fasciole, partly forming continuations of columellar folds. Siphonal canal short, straight or slightly bent to left, with very shallow siphonal notch. Shell measurements and ratios. SL: 4.8 – 14.1 mm, MD: 2.1 – 5.4 mm; AA: 40 – 45 °, A / S: 0.8 – 0.9.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFD6FFF88C92F91DFA8803C9.taxon	discussion	Discussion. This species was confused by Hörnes (1852 b) and Hoernes & Auinger (1880) with the extant Pusia (Ebenomitra) ebenus (Lamarck, 1811), which differs in its much larger size and much broader axial ribs. Consequently, Dollfus & Dautzenberg (1886) introduced Mitra miocaenica (written miocaenica) as new name for the specimens illustrated by Hoernes & Auinger (1880, pl. 9, figs 22 – 25) in a list of Miocene molluscs from the Touraine (France), without any description or comment. Aside from being a nomen nudum, the name Mitra miocaenica, was already preoccupied for an early Miocene species from Italy by Michelotti (1847: 310). According to ICZN Article 58.1 “ species-group names established for different nominal taxa that differ in spelling only in any of the following respects and that are of the same derivation and meaning are deemed to be homonyms when the nominal taxa they denote are included in the same genus or collective group: use of ae, oe or e. ” Therefore, Mitra miocaenica Dollfus & Dautzenberg, 1886 is not available. Boettger (1902) compared this species with Pusia avellana (Bellardi, 1887 b) from the early Miocene of Italy and separated it as variety avellanella. This ‘ variety’ was based on juvenile specimens (see lectotype), whereas Boettger (1906) described fully grown specimens as variety pseudavellana. As first revisers we chose Pusia avellanella as the valid name for this species. Pusia avellana (Bellardi, 1887) differs from the Paratethyan species by its larger size and stouter outline (see Ferrero-Mortara et al., 1981 pl. 51, figs 10 a – b). Bałuk (1997) identified the Paratethyan species as Vexillum cognatum, which was described by Bellardi (1887 b) from the Tortonian of Stazzano (Italy). The holotype, illustrated in Ferrero-Mortara et al. (1981 pl. 51, figs 7 a – b), differs from Pusia avellanella in its more prominent axial ribs, which persist over the entire last whorl. Turricula intermittens dryszczoviensis Friedberg, 1911 is based on slender morphotypes of Pusia avellanella. A relation with Pusia intermittens (Hoernes & Auinger, 1880) is unlikely due to the less fusiform shell with shorter spire and shorter siphonal canal. Pusia avellanella is quite variable concerning size and intensity of its axial sculpture, but specimens with persisting axial sculpture are exceptional. Palaeoenvironment. This species is most abundant at localities suggesting silty-sandy inner neritic environments partly with sea grass meadows (e. g. Gainfarn, Steinebrunn) (Zuschin et al. 2007) but is also found in pelitic deposits, which might correspond to deeper environments.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFD6FFF88C92F91DFA8803C9.taxon	distribution	Distribution in Central Paratethys. Badenian (middle Miocene): Korytnica Basin: Korytnica (Poland) (Bałuk 1997); Ukrainian Fore-Carpathian Basin: Dryszczów (Nadrichne) (Ukraine) (Friedberg 1911); North-Alpine-Carpathian Foredeep: Jerutek at Lysice (Czech Republic) (Hoernes & Auinger 1880); Vienna Basin: Gainfarn, Steinebrunn (Austria) (Sieber 1958 b); Făget Basin: Lăpugiu de Sus (Romania) (Hoernes & Auinger 1880).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFD0FFFB8C92FBB5FED801AD.taxon	description	Figs 4 I, 13 H 1 – H 2	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFD0FFFB8C92FBB5FED801AD.taxon	materials_examined	Type material. Holotype: SMF 360341 (= XII. 2211 a), Senckenberg Museum, Frankfurt / Main, Germany, SL: 5.75 mm, MD: 2.8 mm, figs 4 I, 13 H 1 – H 2. Type locality. Coşteiu de Sus (Romania), Făget Basin. Type stratum. Silt and clay of the Dej Formation. Age. Middle Miocene, Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFD0FFFB8C92FBB5FED801AD.taxon	etymology	Etymology. From confundere (Latin: to mix), referring to the confusion with Pusia pseudorecticosta from the Vienna Basin.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFD0FFFB8C92FBB5FED801AD.taxon	diagnosis	Diagnosis. Pusia species of medium size, stout fusiform profile, with slightly cyrtoconoid spire, and prominent, wide spaced, opisthocline axial ribs.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFD0FFFB8C92FBB5FED801AD.taxon	description	Description. Shell medium-sized, broad fusiform with slightly cyrtoconoid spire. Protoconch (fig. 4 I) multispiral, tall conical of 3.5 smooth convex (height: 580 μm, diameter: 530 μm). Teleoconch of five whorls; first teleoconch whorl low, convex, with periphery close to abapical suture, later whorls increasing in height, moderately convex, with periphery mid-whorl. Suture distinctly incised, weakly undulating along axials ribs. Axial sculpture of very prominent, opisthocline, convex ribs, separated by slightly wider, concave interspaces (14 axial ribs on penultimate whorl). Last whorl convex along periphery, weakly convex above, rapidly contracting below; axial ribs on last whorl weakly sigmoidal, persisting on base. Aperture wide, ovoid with indistinct columellar callus. Columella with four prominent folds, weakening rapidly abapically. Outer lip edge damaged, with seven prominent lirae within. Fasciole with four strong, rounded spiral cords. Siphonal canal short, narrow, straight, with shallow siphonal notch. Shell measurements and ratios. SL: 5.75 mm, MD: 2.8 mm; AA: 46 °, AA: 45 – 52 °, SL / MD: 2.4, AL / AW: 3.3, AH / S: 3.2.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFD0FFFB8C92FBB5FED801AD.taxon	discussion	Discussion. This species is reminiscent of Pusia falsitranssylvanica nov. nom. in sculpture, but differs clearly in its much broader profile, its slightly cyrtoconoid and lower spire, and the coarse lirae. Boettger (1906) confused this species with Pusia pseudorecticosta (Boettger, 1906) from Steinebrunn (Austria) (which was established by him in the same paragraph). These species, however, have little in common and the small size, stout shape, cyrtoconoid spire, blunt axial ribs of Pusia confunda are in sharp contrast with Pusia pseudorecticosta. Palaeoenvironment. The assemblages from Coşteiu de Sus suggest middle neritic palaeoenvironments with soft bottom (own data).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFD0FFFB8C92FBB5FED801AD.taxon	distribution	Distribution in Central Paratethys. Badenian (middle Miocene): Făget Basin: Coşteiu de Sus (Romania) (Boettger 1906).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFD3FFFB8C92FE50FED80780.taxon	description	Figs 13 G 1 – G 2	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFD3FFFB8C92FE50FED80780.taxon	materials_examined	Type material. Holotype: SMF 360346 (= XII. 2212 a), Senckenberg Museum, Frankfurt / Main, Germany, SL: 7.3 mm, MD: 2.2 mm, Coşteiu de Sus (Romania), illustrated in Zilch (1934, pl. 17, fig. 15), figs 13 G 1 – G 2. Type locality. Coşteiu de Sus (Romania), Făget Basin. Type stratum. Silt and clay of the Dej Formation. Age. Middle Miocene, Badenian.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFD3FFFB8C92FE50FED80780.taxon	etymology	Etymology. A combination of falsus (Latin for wrong) and transsylvanica.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFD3FFFB8C92FE50FED80780.taxon	diagnosis	Diagnosis. Pusia species of medium-size, moderately slender fusiform profile, with high spire, convex spire whorls, convex last whorl, and prominent, widely spaced opisthocline axial ribs.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFD3FFFB8C92FE50FED80780.taxon	description	Description. Medium sized, slender fusiform, glossy with high spire. Protoconch largely broken, last protoconch whorl convex, smooth (diameter: 0.5 mm). Teleoconch of six convex whorls, with periphery slightly below mid-whorl, separated by incised suture. Axial sculpture of prominent, opisthocline, convex ribs, separated by slightly wider, concave interspaces (14 axial ribs on penultimate whorl); ribs weakening towards adapical suture, prominent at abapical suture, undulating suture. Last whorl convex, constricted at base, bearing sigmoidal axial ribs, fading over base. Four prominent spiral cords on fasciole. Aperture wide; columellar callus indistinct; columella with three prominent folds. Outer lip not preserved; no lirae inside aperture. Siphonal canal moderately short, moderately wide, straight, with very shallow siphonal notch. Shell measurements and ratios. SL: 7.3 mm, MD: 2.2 mm; AA: 31 °, AA: 45 – 52 °, SL / MD: 2.9, AL / AW: 3.4, AH / S: 2.8.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFD3FFFB8C92FE50FED80780.taxon	discussion	Discussion. This species was treated by Boettger (1902, 1906) as variety transsylvanica of Mitra recticosta sensu Hoernes & Auinger, 1880 [= Pusia pseudorecticosta (Boettger, 1906)]. Mitra (Costellaria) recticosta transsylvanica Boettger, 1902, however, is a primary homonym of Mitra (Cylindra) transsylvanica Hoernes & Auinger 1880, and we therefore introduce falsitranssylvanica as replacement name. Mitra (Costellaria) recticosta transsylvanica Boettger, 1902 (= Pusia falsitranssylvanica), was treated as senior synonym of Mitra pseudorecticosta Boettger, 1906 by Landau et al. (2013). Pusia falsitranssylvanica differs distinctly from P. pseudorecticosta in its convex whorls (instead of subcylindrical) and the more prominent, wider spaced, opisthocline axial ribs. Moreover, the occurrence at Coşteiu de Sus suggests a preference for middle neritic soft bottom environments, whereas P. pseudorecticosta is typically found in silty-sandy deposits of coastal environments. The placement in Pusia is tentative. Palaeoenvironment. The assemblages from Coşteiu de Sus suggest middle neritic palaeoenvironments with soft bottom (own data).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFD3FFFB8C92FE50FED80780.taxon	distribution	Distribution in Central Paratethys. Badenian (middle Miocene): Făget Basin: Coşteiu de Sus (Romania) (Boettger 1906).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFDDFFF58C92FF39FE450430.taxon	description	Figs 13 I 1 – I 2, J	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFDDFFF58C92FF39FE450430.taxon	materials_examined	Type material. Lectotype (designated herein): NHMW 1862 / 0029 / 0006, SL: 9.8 mm, MD: 4.7 mm, Rudice (Czech Republic), figs 13 I 1 – I 2. Paralectotype: NHMW 2020 / 0097 / 0001, SL: 12.6 mm, MD: 6.5 mm, Rudice (Czech Republic), fig. 13 J. Revised description. Shell moderately large, stout fusiform, with slightly gradate spire. Protoconch and early teleoconch unknown. Teleoconch whorls low, weakly convex, with periphery in abapical third; suture impressed. Sculpture of sharp crested, orthocline to weakly prosocline ribs, separated by interspaces of about same width (about 15 axial ribs on penultimate whorl). Adapical weak subsutural spiral groove cuts ribs, forming narrow subsutural spiral cord of weak nodes. Further delicate spirals placed below subsutural cord, abapically rapidly weakening. Last whorl convex with short, moderately constricted base. Axial ribs becoming subobsolete on last whorl. Columella with four prominent, oblique columellar folds. Siphonal canal moderately long, straight. Shell measurements and ratios. SL: 9.8 – 12.6 mm, MD: 4.7 – 6.5 mm (fragments; restored height: 13 – 16 mm); assumed SL of complete specimen: 18 mm, AA: 33 °.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFDDFFF58C92FF39FE450430.taxon	discussion	Discussion. Only two fragments are available, which Hoernes & Auinger (1880) described as variety of “ Mitra plicatula ” (= Vexillum transalpinum nov. sp.). A close relationship between Pusia moravica (Hoernes & Auinger, 1880) and V. transalpinum, however, is unlikely in respect to the short spire whorls and broad, convex last whorl of P. moravica. In addition, the subsutural spiral row of weak nodes and the presence of spiral grooves allows a clear separation from V. transalpinum. Palaeoenvironment. The specimens derive from basinal clay, suggesting a middle neritic depositional environment.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFDDFFF58C92FF39FE450430.taxon	distribution	Distribution in Central Paratethys. Badenian (middle Miocene): Vienna Basin: Rudice (Czech Republic) (Hoernes & Auinger 1880).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFDDFFF48C92FB2CFE46021A.taxon	description	Figs 14 A 1 – A 2, B 1 – B 2	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFDDFFF48C92FB2CFE46021A.taxon	materials_examined	Type material. Lectotype: SMF 360345 (= SMF 12.2208 a), Senckenberg Museum, Frankfurt / Main, Germany, SL: 11.5 mm, MD: 5.0 mm, Coşteiu de Sus (Romania), illustrated in Zilch (1934, pl. 17, fig. 18), figs 14 B 1 – B 2. Paralectotypes: SMF 360.2208 b – c, Senckenberg Museum, Frankfurt / Main, Germany, 17 + 7 specimens, Coşteiu de Sus (Romania). Additional material. NHMW 1865 / 0036 / 0010, SL: 9.0 mm, MD: 3.9 mm, Drnovice u Vyškova (Czech Republic), figs 14 A 1 – A 2. Revised description. Shell moderately large, broad fusiform, with distinctly incised suture, and broad conical to weakly cyrtoconoid spire. Protoconch unknown. Teleoconch of seven whorls. Spire whorls weakly convex, with periphery at abapical suture. Axial sculpture of orthocline, convex ribs, separated by interspaces of about same width (about 20 – 25 axial ribs on last whorl). Adapical tips of axial ribs may be slightly swollen resulting in slightly gradate spire. Last whorl strongly convex, rapidly contracting. Aperture wide, ovoid with indistinct anal sinus. Columellar callus indistinct. Columella with four prominent folds, weakening abapically. Outer lip thin, abapically contracting, about 10 – 12 lirae within. No spiral sculpture, except four to six spiral cords on weak fasciole, partly extending from columellar folds. Siphonal canal straight, narrow, moderately short, with shallow siphonal notch. Shell measurements and ratios. SL: 9.0 – 12.0 mm, MD: 3.9 – 5.5 mm; AA: 45 – 55 °, SL / MD: 2.3 – 2.4, AL / AW: 3.7, AH / S: 2.8 – 3.0.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFDDFFF48C92FB2CFE46021A.taxon	discussion	Discussion. Boettger (1906) described this species as variety of “ Mitra ebenus ” [= Ebenomitra leucozona (Andrzejowski 1830)], which is clearly incorrect. Pusia paraleucozona has four prominent columellar folds and numerous delicate axial ribs, whereas E. leucozona has three folds and few blunt axial swellings. At first glance it is similar to Pusia moravica (Hoernes & Auinger, 1880) in being broadly fusiform with narrow ribs persisting on all whorls, but that species is larger [reconstructed height about 13 – 16 mm vs 9.0 – 12.0 mm in Pusia paraleucozona (Boettger, 1906)], at the same size the axial ribs are wider spaced in P. moravica, and the spiral sculpture in the subsutural area is weak, but present, whereas in P. paraleucozona spiral sculpture is restricted to the siphonal fasciole. Ebenomitra renauleauensis Landau, Ceulemans & Van Dingenen, 2019, from the Tortonian of France, is reminiscent of P. paraleucozona but differs in its smaller size and the coarser axial ribs. Palaeoenvironment. Unknown; probably offshore settings.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFDDFFF48C92FB2CFE46021A.taxon	distribution	Distribution in Central Paratethys. Badenian (middle Miocene): North Alpine-Carpathian Foredeep: Drnovice u Vyškova (Czech Republic); Korytnica Basin: Korytnica (Poland) (Bałuk 1997); Făget Basin: Coşteiu de Sus (Romania) (Boettger 1906).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFDCFFF68C92F886FB1200D4.taxon	description	Figs 4 J, 15 A 1 – A 2, B 1 – B 2, C 1 – C 2, D	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFDCFFF68C92F886FB1200D4.taxon	materials_examined	Type material. Lectotype (designated herein): NHMW 2020 / 0076 / 0002, SL: 10.2 mm, MD: 3.6 mm, Steinebrunn (Austria), illustrated in Hoernes & Auinger (1880: pl. 10, figs 6 a – c), figs 15 A 1 – A 2. Paralectotype: NHMW 2020 / 0076 / 0001, SL: 16.0 mm, MD: 5.5 mm, Steinebrunn (Austria), illustrated in Hoernes & Auinger (1880, pl. 10, figs 5 a – c), figs 15 B 1 – B 2. Additional material. NHMW 2020 / 0076 / 0003, SL: 14.4 mm, MD: 4.9 mm, Steinebrunn (Austria), fig. 15 D; NHMW 2020 / 0076 / 0004, SL: 14.0 mm, MD: 5.0 mm, Steinebrunn (Austria), figs 15 C 1 – C 2; NHMW 1887 / 0018 / 0019, Steinebrunn (Austria), illustrated in Meznerics (1933, pl. 14, figs 7 a – b); NHMW 2020 / 0076 / 0005, SL: 9.8 mm, MD: 3.7 mm, Steinebrunn (Austria), fig. 4 J; NHMW 1855 / 0045 / 0035, 80 specimens, Steinebrunn (Austria); NHMW 1866 / 0001 / 1018, 24 specimens Forchtenau (Austria); NHMW 1871 / 0010 / 0015, 2 specimens, Sedlec (= Porzteich) (Czech Republic). Revised description. Shell moderately large, moderately slender fusiform, with high, slightly gradate spire. Protoconch (Fig. 4 J) multispiral, high conical of 3.5 smooth, nearly straight-sided whorls (height: 940 μm, diameter: 750 μm). Teleoconch of about eight subcylindrical to weakly convex whorls, separated by narrowly impressed suture. Axial sculpture of close-set orthocline ribs separated by slightly narrower interspaces (about on 20 penultimate whorl). Last whorl strongly constricted. Axial sculpture on last whorl often reduced. Aperture wide, ovoid with indistinct anal canal. Columellar callus indistinct, barely delimited from base. Columella with four oblique columellar folds, weakening abapically. Outer lip convex, thin, with 10 – 12 long delicate lirae placed deep within aperture. About five weak to prominent spiral cords on weak fasciole, partly extending from columellar folds. Siphonal canal moderately short, bent slightly to left, with shallow siphonal notch. Shell measurements and ratios. SL: 10.2 – 16.0 mm, MD: 3.6 – 5.0 mm; AA: 32 – 40 °, SL / MD: 2.8 – 2.9, AL / AW: 3.0 – 3.9, AH / S: 2.5 – 2.7.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFDCFFF68C92F886FB1200D4.taxon	discussion	Discussion. Hoernes & Auinger (1880) identified this species as Mitra recticosta, which was described by Bellardi (1850) from the Pliocene of Italy. Bellardi (1887 b) doubted this identification and separated the Viennese shells based on their smaller size, more numerous and narrower axial ribs and the shorter last whorl from his M. recticosta but did not propose a new name. This was done by Boettger (1906) who introduced Mitra pseudorecticosta as new name. It is important to note that although Boettger (1906) described specimens from Coşteiu de Sus (Romania), he clearly referred to the Austrian specimens illustrated by Hoernes & Auinger (1880), when introducing Mitra pseudorecticosta as new name. Curiously, the specimens from Coşteiu de Sus, which Boettger (1906) had at hand when discussing Mitra pseudorecticosta are not conspecific with the Austrian shells and are described here as Pusia confunda nov. sp. Friedberg (1910), who was unaware of Boettger’s paper, introduced Mitra dubia as new name for Mitra recticosta Hoernes & Auinger, 1880. This name, however, was preoccupied by Hutton (1873) and therefore, Friedberg (1911) proposed Turricula vindobonensis as replacement name a year later. Boettger (1906) and Friedberg (1910, 1911) refer to the same lot of illustrated specimens and therefore, Turricula vindobonensis is an objective junior synonym of Mitra pseudorecticosta Boettger, 1906. The placement of this species in Pusia is tentative. Hölzl (1973: 93) mentioned Vexillum (Costellaria) vindobonense (Friedberg) from the Ottnangian (middle Burdigalian) of Kaltenbachgraben in Bavaria. This would be the oldest record of this group from the Paratethys Sea. Palaeoenvironment. Most abundant at localities suggesting silty-sandy inner neritic environments partly with sea grass meadows (e. g. Steinebrunn) (own data).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFDCFFF68C92F886FB1200D4.taxon	distribution	Distribution in Central Paratethys. Badenian (middle Miocene): Korytnica Basin: Korytnica (Poland) (Bałuk 1997); Ukrainian Fore-Carpathian Basin: Dryszczów (Nadrichne), Zborów (Zboriv) (Ukraine) (Friedberg 1911); Vienna Basin: Niederleis, Steinebrunn (Hoernes & Auinger 1880), Sedlec (= Porzteich) (Czech Republic); Eisenstadt-Sopron Basin: Forchtenau (Hoernes & Auinger 1880). Eastern Paratethys. Tarkhanian (early Middle Miocene) of Tomakivka (S. Ukraine) (Mikhailovsky 1903). Proto-Mediterranean Sea. Karaman Basin: Seyithasan (Turkey) (Landau et al. 2013).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFDEFFF08C92FE88FE770165.taxon	description	Figs 4 K, 15 E 1 – E 2, F 1 – F 2, G 1 – G 2, H	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFDEFFF08C92FE88FE770165.taxon	description	[Costellaria] Borsoni Bell. — Hoernes 1880: 125 [non Bellardi, 1850].	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFDEFFF08C92FE88FE770165.taxon	materials_examined	Type material. Holotype: NHMW 1887 / 0018 / 0018, illustrated in Meznerics (1933, pl. 14, figs 6 a – b). The Meznerics (1933) collection is stored in the NHMW but the specimen could not be found. Additional topotypic material. NHMW 2020 / 0078 / 0001, SL: 10.9 mm, MD: 3.8 mm, Steinebrunn (Austria), 15 G 1 – G 2; NHMW 2020 / 0078 / 0002, SL: 9.8 mm, MD: 3.5 mm, Steinebrunn (Austria), 15 E 1 – E 2, 4 K; NHMW 2020 / 0078 / 0003, SL: 11.8 mm, MD: 4.0 mm, Steinebrunn (Austria), 15 F 1 – F 2; NHMW 2020 / 0079 / 0001, SL: 11.0 mm, MD: 3.6 mm, Steinebrunn (Austria), 15 H; NHMW 1884 / 0037 / 2895, 61, specimens, Steinebrunn (Austria); NHMW 1860 / 0001 / 0113, Steinebrunn (Austria), illustrated in Hoernes & Auinger (1880 pl. 10, fig. 9); NHMW 1846 / 0037 / 0111, Steinebrunn (Austria), illustrated in Hörnes (1852 b, pl. 10, figs 31 a – c) Revised description. Shell moderately large, moderately slender fusiform, with high spire. Protoconch (fig. 4 K) high conical of 3.5 smooth, weakly convex whorls (height: 900 μm, diameter: 720 μm). Teleoconch of six to seven weakly convex to subcylindrical whorls, with periphery close above abapical suture; suture deeply incised. Sculpture comprising prominent, weakly opisthocyrt axial ribs, separated by interspaces of roughly equal width. Axial ribs often slightly swollen at adapical tip, resulting in weakly gradate spire. Delicate spiral cords restricted to interspaces between axial ribs. Axial ribs slightly broader on last whorl. Last whorl slightly inflated, evenly convex, strongly constricted at base. Aperture wide, ovoid, with indistinct anal sinus. Columellar callus narrow, sharply delimited. Columella with three prominent folds, weakening abapically; very weak fourth fold in some specimens. Outer lip thin, bearing about ten delicate lirae within. About four weak spiral cords on base. Siphonal canal short, bent slightly to left, with shallow siphonal notch. Shell measurements and ratios. SL: 8.4 – 12.9 mm, MD: 3.3 – 4.9 mm; AA: 33 – 35 °, SL / MD: 2.7 – 3.0, AL / AW: 2.2 – 3.5, AH / S: 3.0 – 4.5.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFDEFFF08C92FE88FE770165.taxon	discussion	Discussion. This species was originally identified by Hörnes (1852 b) as Mitra recticosta Bellardi, 1850, which lacks spiral sculpture. Consequently, Hoernes & Auinger (1880) identified it as Uromitra borsoni Bellardi, 1850, based on the rather schematic drawings in Bellardi (1850). Meznerics (1933) doubted this identification and described the specimens from Steinebrunn as new species. This view is supported by the illustration of a syntype of Vexillum borsoni (Bellardi, 1850) from the Tortonian S. Agata (Italy) (Ferrero-Mortara et al., 1981, pl. 52, figs 14 a – b). It differs from the Austrian species in its sigmoidal axial ribs, the strongly constricted base and the longer siphonal canal. Pusia schafferi was also mentioned from the Ottnangian (Burdigalian) of Bavaria (Kaltenbachgraben) by Steininger (1973) based on a spire fragment. The specimen has never been illustrated and the identification remains unclear. Pusia schafferi (Meznerics, 1933) has a similarly elongated fusiform shape as P. pseudorecticosta (Boettger, 1906), but differs in having fine spiral grooves visible in the axial interspaces, whereas in P. pseudorecticosta only the strong spiral cords over the siphonal fasciole are present. Palaeoenvironment. Steinebrunn represents a silty-sandy inner neritic environment with sea grass meadows (own data).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFDEFFF08C92FE88FE770165.taxon	distribution	Distribution in Central Paratethys. Badenian (middle Miocene): North-Alpine-Carpathian Foredeep: Hrušovany nad Jevišovkou, Jerutek at Lysice, Drnovice u Vyškova (Czech Republic) (Hoernes & Auinger 1880); Vienna Basin: Gainfarn, Niederleis, Steinebrunn, Vienna / Pötzleinsdorf (Austria), Sedlec (= Porzteich), Mikulov (Czech Republic), (Hoernes & Auinger 1880; Sieber 1958 b), Rohožník – Konopiská (Slovakia) (Biskupič 2020); Eisenstadt-Sopron Basin: Forchtenau (Austria) (Sieber 1956); Danube Basin: Hlohovec (= Bischofswart) (Slovakia) (Hoernes & Auinger 1880).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFD8FFF08C92FE19FEBF06A2.taxon	description	Figs 4 L, 15 I 1 – I 2, J 1 – J 2, K 1 – K 2	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFD8FFF08C92FE19FEBF06A2.taxon	materials_examined	Type material. Lectotype: SMF 360339 (= SMF XII 2213 a), Senckenberg Museum, Frankfurt / Main, Germany, SL: 6 mm, MD: 2.4 mm, Coşteiu de Sus (Romania), illustrated in Zilch (1934, pl. 17, fig. 16), figs 15 J 1 – J 2. Paralectotypes: SMF XII 2213 b – c, Senckenberg Museum, Frankfurt / Main, Germany, Coşteiu de Sus (Romania). Additional material. NHMW 2020 / 0081 / 0001, SL: 5.4 mm, MD: 2.1 mm, Coşteiu de Sus (Romania), 15 I 1 – 15 I 2, 4 L; NHMW 2020 / 0081 / 0002, SL: 6.0 mm, MD: 2.5 mm, Coşteiu de Sus (Romania), 15 K 1 – K 2; NHMW 2018 / 0248 / 0327, 47 specimens, Coşteiu de Sus (Romania). Revised description. Shell small, moderately broad fusiform, with high conical spire. Protoconch (fig. 4 L) multispiral, high conical, mammillate, of about 3.5 weakly convex whorls (height: 730 μm, diameter: 550 μm). Teleoconch consisting of five convex whorls, with periphery in abapical third, separated by impressed suture. Last whorl convex, somewhat inflated, constricted at base. Axial sculpture of swollen, convex axial ribs separated by concave interspaces of roughly equal width. Axial ribs becoming slightly more prominent towards abapical suture. Axial ribs and interspaces widening on last whorl, subobsolete towards aperture in some specimens. Aperture moderately narrow, ovoid, posteriorly narrowly angulated, with weak parietal swelling. Columellar callus narrow. Columella with four prominent columellar folds, weakening abapically. Outer lip thin, bearing about five prominent lirae within. Five to seven prominent spiral cords on weak fasciole. Siphonal canal moderately short to moderately long, narrow, straight or bent slightly to left, with very shallow siphonal notch. Shell measurements and ratios. SL: 4.9 – 6.0 mm, MD: 1.9 – 2.2 mm; AA: 38 – 41 °, SL / MD: 2.5 – 2.6, AL / AW: 3.4 – 4.3, AH / S: 2.4 – 2.7.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFD8FFF08C92FE19FEBF06A2.taxon	discussion	Discussion. This species is characterised by its small size, convex whorls and swollen axial ribs. There are no similar species in Paratethyan faunas. Ebenomitra cf. renauleauensis Landau, Ceulemans & Van Dingenen, 2019, from the Tortonian of France, is superficially similar, but is distinguished by its lower spire, wider apical angle and lower number of axial ribs (see Landau et al., 2019, pl. 50, figs 2 – 4). Palaeoenvironment. The assemblages from Coşteiu de Sus suggests middle to outer neritic palaeoenvironments with soft bottom (own data).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFD8FFF08C92FE19FEBF06A2.taxon	distribution	Distribution in Central Paratethys. Badenian (middle Miocene): Korytnica Basin: Korytnica (Poland) (Bałuk 1997); Făget Basin: Coşteiu de Sus (Romania) (Boettger 1902); Hațeg Basin: Răhchitova (Romania) (Popa & Ianoliu 2000).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFD8FFF38C92F95AFB660410.taxon	description	Figs 18 C 1 – C 2	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFD8FFF38C92F95AFB660410.taxon	materials_examined	Type material. Lectotype (designated herein): SL. 12.0 mm, MD: 5.0 mm, Hołubica (Ukraine), stored in the State Museum of Natural History of the National Academy of Sciences of Ukraine in Lviv (formerly Muzeum Dzieduszyckich), illustrated in Friedberg (1911: 24, pl. 1, fig. 17). Unfortunately, it was not possible to obtain pictures of the lectotype and we reproduce the illustration from Friedberg (1911), figs 18 C 1 – C 2. Revised description (based on description and illustration in Friedberg, 1911). Shell moderately large, stout fusiform to biconical. Protoconch multispiral, high conical, mammillate, of three smooth, weakly convex, whorls. Teleoconch boundary marked by beginning of axial sculpture. Teleoconch of five convex whorls, with periphery slightly below mid-whorl, separated by incised suture. Axial sculpture of prominent, close-set axial ribs, separated by narrower interspaces (16 – 20 axial ribs on penultimate whorl), crossed by delicate spiral threads, most prominent in axial interspaces. Slightly deeper spiral groove separates nodulose subsutural spiral cord. Last whorl evenly convex, strongly constricted. Aperture narrow, elongate. Columellar callus narrow, indistinct. Columella with four folds, abapically decreasing in strength. Outer lip thin, prominent lirae inside aperture. Four prominent spiral cords on fasciole, partly from extensions of columellar folds. Siphonal canal moderately long, narrow, straight, with deep siphonal notch. Shell measurements and ratios. SL: 10 – 12 mm, MD: 4.2 – 5.0 mm; AA: 40 °.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFD8FFF38C92F95AFB660410.taxon	discussion	Discussion. This species is characterised by its dense axial / spiral sculpture. At first sight, the illustration in Friedberg (1911) is reminiscent of an Antithala species. The thin outer lip, the wider aperture with lirae inside, and the presence of delicate spiral cords in the axial interspaces exclude a closer relationship with that genus. Pusia brevior was established by Friedberg (1911) as variety of Mitra recticosta Bellardi, 1850, but differs from this Pliocene Italian species distinctly by its squatter shape, the presence of spiral sculpture, the smaller size, and wider apical angle. The absence of spiral sculpture distinguishes also Pusia confunda nov. sp., from P. brevior, which develops a similar shape. Pusia schafferi Meznerics, 1933 has comparable sculpture, but is much more slender and has a higher spire. Palaeoenvironment. Unknown.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFD8FFF38C92F95AFB660410.taxon	distribution	Distribution in Central Paratethys. Badenian (middle Miocene): Polish Fore-Carpathian Basin: Borki Wielkie (Poland) (Friedberg 1911); Voronyaky Hills: Hołubica (Holubytsia) (Ukraine) (Friedberg 1911).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFDBFFF28C92FBCCFA150216.taxon	type_taxon	Type species. Vexillum plicatum Röding, 1798 (= Voluta plicarium Linnaeus, 1758), subsequent designation by Woodring (1928: 244), Present-day, Indo-West Pacific.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFDBFFF28C92FBCCFA150216.taxon	description	Description. “ Shell small to medium-sized, fusiform or broadly fusiform to turriform, ovate or biconical; spire very tall to low. Protoconch tall, conical, with three or more evenly convex glossy whorls. Teleoconch whorls evenly convex to subcylindrical or with flattened outline, typically distinctly shouldered. Sculpture dominated by axial elements, represented by ribs of varying strength, sharp or rounded, usually slightly undulating. Spiral sculpture variable, absent in some species, in others represented by fine, evenly spaced grooves, pronounced between axial ribs, or throughout shell surface, overriding axial ribs. Siphonal canal short to very long, straight and tapering or recurved at its tip, notched at its end. Aperture elongate, narrow; outer aperture lip lirate within, sometimes with strong denticles formed by lirae at outer lip’s edge. Inner aperture lip with three or four distinct, sometimes very strong columellar folds, uppermost fold strongest ” (Fedosov et al., 2017: 617). Synonyms of relevance for Paratethyan faunas (see Fedosov et al., 2017 for full list):	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFDBFFF28C92FBCCFA150216.taxon	discussion	Tiara Swainson, 1831, type species: Mitra corrugata Lamarck, 1811, subsequent designation by Gray (1847: 142); present-day, Indo-West Pacific. Turricula H. Adams & A. Adams, 1853 [non Schumacher, 1817], type species: Voluta vulpecula Linnaeus, 1758, subsequent designation by Cossmann (1899: 162); present-day, Indo-West Pacific. Uromitra Bellardi, 1887, type species: Uromitra antegressa Bellardi, 1887 (Bellardi, 1887 b: 23), subsequent designation by Harris (1897: 125); Oligocene, Italy.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFDAFFEC8C92FDC7FE9501AD.taxon	description	Figs 16 A 1 – A 2, B 1 – B 2, C 1 – C 2, D 1 – D 2	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFDAFFEC8C92FDC7FE9501AD.taxon	materials_examined	Type material. Lectotype (designated herein): NHMW 1846 / 0037 / 0102, Baden (Austria), SL: 26.7 mm, MD: 8.0 mm, illustrated in Hörnes (1852 b, pl. 10 figs 28 a – b), figs 16 A 1 – A 2. Paralectotypes: NHMW 2020 / 0084 / 0001, Vienna / Grinzing (Austria), SL: 26.5 mm, MD: 7.6 mm, figs 16 B 1 – B 2; NHMW 2020 / 0085 / 0001, Baden (Austria), SL: 24.7 mm, MD: 7.5 mm, figs 16 C 1 – C 2; NHMW 2020 / 0085 / 0002, 5 specimens, Baden (Austria), SL: 21.6 mm, MD: 7.4 mm, figs 16 D 1 – D 2. Revised description. Shell large, slender fusiform with weakly coeloconoid early spire, changing to slightly cyrtoconoid spire during ontogeny. Protoconch unknown. Teleoconch of eight whorls, with narrowly impressed suture. Early spire whorls straight-sided, later whorls weakly convex, with periphery in abapical third. Last whorl high, weakly constricted at base. Sculpture on early spire whorls consisting of narrow, sharp-crested, orthocline to weakly opisthocline axial ribs, separated by slightly wider interspaces. Abapically, ribs becoming low, wide-spaced (about 10 on penultimate whorl), most prominent along abapical suture; last whorl smooth. Spiral sculpture consisting of faint spiral grooves (rarely preserved), visible only under strong magnification. Few weak or subobsolete spiral cords on fasciole. Aperture moderately narrow, elongate, posteriorly narrowly angulated, with indistinct anal sinus. Columellar callus narrow, indistinct. Columella with four oblique folds, weakening abapically. Outer lip thin with short, delicate lirae within. Siphonal canal moderately long, straight or weakly bent to left, with shallow anterior notch. Shell measurements and ratios. SL: 19.9 – 26.7 mm, MD: 6.3 – 8.0 mm; AA: 23 – 30 °, AA: 45 – 52 °, SL / MD: 3.4 – 3.5, AL / AW: 4.6 – 4.8, AH / S: 2.6 – 2.9.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFDAFFEC8C92FDC7FE9501AD.taxon	discussion	Discussion. The species is characterised by its peculiar axial sculpture of widely spaced ribs, which fade towards the adapical suture. Its axial ribs and basal spiral cords may become subobsolete in some specimens, resulting in an almost smooth shell. Vexillum intermittens (Hoernes & Auinger, 1880) is distinguished by its opisthocline axial ribs on early spire whorls, the deeper suture, and the narrower spire angle. Palaeoenvironment. The species is found in clay of the Baden Formation suggesting middle neritic settings with up to 250 m water depth (Hohenegger et al. 2008). It is unclear, however, if all other localities represent the same depositional environment.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFDAFFEC8C92FDC7FE9501AD.taxon	distribution	Distribution in Central Paratethys. Badenian (middle Miocene): Ukrainian Fore-Carpathian Basin: Dryszczów (Nadrichne), Żukowce (Zhukivtsi) (Ukraine) (Friedberg 1911); Vienna Basin: Baden, Vienna / Grinzing, Vienna / Pötzleinsdorf, Steinebrunn (Hoernes & Auinger 1880; Sieber 1953, 1958 b); Pannonian Basin: Szokolya (Hungary) (Báldi 1960); Bükk Mountains (Hungary) (Csepreghy-Meznerics 1972); Southern Carpathians: Bahna (Romania) (Tiţă 2007).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFC4FFEF8C92FE50FECA011E.taxon	description	Figs 17 A 1 – A 2, B 1 – B 2, C 1 – C 2, D 1 – D 2, E 1 – E 2	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFC4FFEF8C92FE50FECA011E.taxon	materials_examined	Type material. Holotype: M. 61.4361, Hungarian Natural History Museum, SL: 24.0 mm, MD: 6.2 mm, Sámsonháza (Hungary) (Pálfy et al. 2008), figs 17 A 1 – A 2. Paratypes: M. 61.4362, 3 specimens, Hungarian Natural History Museum, SL: 15.0 mm, MD: 3.9 mm, Sámsonháza (Hungary) (Pálfy et al. 2008), Figs 17 B 1 – B 2, C 1 – C 2, D 1 – D 2, E 1 – E 2. Revised description. Shell moderately large, slender fusiform, with very high spire. Protoconch unknown. Teleoconch of nine high, subcylindrical to weakly convex whorls, with periphery mid-whorl, separated by incised suture. Sculpture of widely spaced, sharp-crested, orthocline to weakly opisthocline axial ribs, most prominent along periphery, weakening towards adapical and abapical suture. Axial sculpture fading out on seventh spire whorl; penultimate and last whorls smooth. Last whorl high, subcylindrical, weakly shouldered, moderately constricted at base. No spiral sculpture except for faint spiral cords on weak fasciole. Aperture narrow elongate. Columellar callus indistinct, columella with four oblique folds. Outer lip not preserved. Siphonal canal moderately long, wide, slightly bent to left, with wide, shallow siphonal notch. Shell measurements and ratios. SL: 15.0 – 24.0 mm, MD: 3.9 – 6.2 mm; AA: 24 – 30 °, A / S: 0.6 – 0.7.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFC4FFEF8C92FE50FECA011E.taxon	discussion	Discussion. This species is reminiscent of Vexillum badense (Hoernes & Auinger, 1880) and Vexillum intermittens (Hoernes & Auinger, 1880), but is distinguished from both species by its distinctly more slender shell and higher spire whorls. Its axial ribs are most prominent mid-whorl, whereas in the other two species they strengthen towards the abapical suture. Finally, the occurrence in silty-sandy nearshore deposits of Sámsonháza indicates an ecological separation from V. badense and V. intermittens, which are confined to basinal settings. Palaeoenvironment. The Sámsonháza Formation represents inner neritic shallow water environments (Kovács & Vicián, 2017).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFC4FFEF8C92FE50FECA011E.taxon	distribution	Distribution in Central Paratethys. Badenian (middle Miocene): Pannonian Basin: Sámsonháza (Hungary) (Strausz 1966).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFC7FFEF8C92FEC0FD0F07EA.taxon	description	Figs 4 M, 16 E 1 – E 2, F 1 – F 2, G 1 – G 2, H 1 – H 2 Costellaria intermittens n. form. — Hoernes 1880: 125 [nomen nudum].	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFC7FFEF8C92FEC0FD0F07EA.taxon	description	Halaváts, 1884]. non Vexillum (Costellaria) intermittens (R. Hörnes et Auinger) — Stancu et al. 1971: 125, pl. 7, fig. 9 [=?].	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFC7FFEF8C92FEC0FD0F07EA.taxon	materials_examined	Type material. Lectotype (designated herein): NHMW 1846 / 0037 / 0105: Bad Vöslau (Austria), SL: 14.0 mm, MD: 4.6 mm, illustrated in Hoernes & Auinger (1880, pl. 10, figs 2 a – c), figs 16 G 1 – G 2. Paralectotypes: NHMW 1863 / 0015 / 0624: Bad Vöslau (Austria), SL: 17.7 mm, MD: 4.8 mm, illustrated in Hoernes & Auinger (1880, pl. 10, figs 3 a – c), figs 16 E 1 – E 2; NHMW 2020 / 0087 / 0001: Baden or Bad Vöslau (Austria), SL: 14.4 mm, MD: 4.2 mm, illustrated in Hoernes & Auinger (1880, pl. 10, figs 1 a – c), figs 16 F 1 – F 2; NHMW 2020 / 0087 / 0002: Baden or Bad Vöslau (Austria), SL: 12.2 mm, MD: 3.4 mm, illustrated in Hoernes & Auinger (1880, pl. 10, figs 4 a – c), figs 16 H 1 – H 2. NHMW 2020 / 0087 / 0003: Baden or Bad Vöslau (Austria), SL: 15.3 mm, MD: 4.0 mm, fig 4 M; NHMW 2020 / 0087 / 0004, 26 specimens, Baden or Bad Vöslau (Austria). Revised description. Shell moderately large, slender fusiform, with high conical spire. Protoconch (fig. 4 M) high conical, of three smooth, weakly convex whorls (height: 990 μm, diameter: 740 μm). Teleoconch of seven straight-sided to faintly convex whorls, with periphery close above abapical suture; suture deeply incised. Last whorl slightly more convex or subcylindrical, strongly constricted at base. Sculpture of first three to four teleoconch whorls consisting of seven to ten broad, convex, weakly opisthocline axial ribs, separated by slightly wider interspaces. Axial ribs usually weakening on middle spire whorls, reappearing on penultimate and last whorls, typically fading out towards adapical suture and slightly swollen towards abapical suture. Axial sculpture fading out on last whorl below periphery. Rarely, axial sculpture persisting throughout growth, or completely reduced on last whorl. Aperture moderately narrow, elongate; columellar callus indistinct. Columella with four folds, weakening abapically; fourth fold very weak and short. Outer lip thin, 10 to 16 delicate, subparallel lirae within. Spiral sculpture represented by few faint spiral threads visible only under strong magnification; five prominent spiral cords on weak fasciole. Siphonal canal moderately long, moderately narrow, straight, with wide and shallow anterior notch. Shell measurements and ratios. SL: 12.2 – 17.7 mm, MD: 3.4 – 4.8 mm; AA: 25 – 28 °, SL / MD: 3.2 – 3.5, AL / AW: 4.4 – 5.1, AH / S: 2.1 – 2.4.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFC7FFEF8C92FEC0FD0F07EA.taxon	discussion	Discussion. This species displays little variability in shape and size, but varies considerably in sculpture. In most specimens, spiral sculpture is absent or strongly reduced along the last two or three spire whorls. Vexillum intermittens (Hoernes & Auinger, 1880) is documented so far only from the type area of Baden and Bad Vöslau in Austria. Other Paratethyan occurrences are misidentifications; e. g. Kojumdgieva in Kojumdgieva & Strachimirov (1960) described a specimen from the Badenian of Bulgaria, which differs from V. intermittens in its extraordinarily slender and smooth shell. The specimen from Letkés (Hungary), described by Strausz (1966) develops spiral cords in the axial interspaces and is most probably conspecific with Vexillum szobbiensis (Halaváts, 1884). Palaeoenvironment. The species is only found so far in clay of the Baden Formation suggesting middle neritic settings with up to 250 m water depth (Hohenegger et al. 2008).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFC7FFEF8C92FEC0FD0F07EA.taxon	distribution	Distribution in Central Paratethys. Badenian (middle Miocene): Vienna Basin: Baden-Sooss, Bad Vöslau, Baden (Austria) (Hoernes & Auinger 1880).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFC6FFEE8C92FF39FB7406C2.taxon	description	Figs 4 N, 16 I 1 – I 2, J 1 – J 2, K 1 – K 2, L 1 – L 2, M	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFC6FFEE8C92FF39FB7406C2.taxon	description	cf. Mitra (Callithea) Michelottii M. Hoernes — Hoernes & Auinger 1880: 87, figs 11 a – c.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFC6FFEE8C92FF39FB7406C2.taxon	materials_examined	Type material. Lectotype (designated herein): NHMW 1851 / 0013 / 0001, SL: 20.3 mm, MD: 5.4 mm, Baden (Austria), illustrated in Hörnes (1852 b, pl. 10, figs 30 a – b), figs 4 N, 16 I 1 – I 2. Paralectotype: NHMW 2020 / 0088 / 0001, SL: 19.8 mm, MD: 4.7 mm, Baden (Austria), figs 16 J 1 – J 2. Additional material. NHMW 1869 / 0001 / 0249, SL: 26.5 mm, MD: 7.3 mm, Möllersdorf (Austria), figs 16 L 1 – L 2; NHMW 2020 / 0089 / 0001, SL: 25.1 mm, MD: 7.3 mm, Möllersdorf (Austria), figs 16 K 1 – K 2; NHMW 1875 / 0009 / 0007, SL: 15.3 mm, MD: 4.3 mm, Lăpugiu de Sus (Romania), illustrated in Hoernes & Auinger (1880, pl. 10, figs 11 a – c), fig. 16 M; NHMW 1874 / 0024 / 0003, 2 specimens, Baden (Austria); NHMW 1869 / 0040 / 0035, Hrušovany nad Jevišovkou (Czech Republic), NHMW 1862 / 0039 / 0009, 2 specimens, Rudice (Czech Republic), NHMW 1854 / 0035 / 0098, 5 specimens, Lăpugiu de Sus (Romania). Revised description. Shell large, slender fusiform, with high spire. Protoconch multispiral, high conical, of three smooth, weakly convex whorls (height: 730 μm, diameter: 550 μm). Teleoconch of nine convex whorls, with periphery at mid-whorl or slightly below, separated by deeply incised suture. Axial sculpture on early teleoconch whorls of about ten prominent, convex, weakly opisthocline, widely spaced axial ribs, crossed by deep spiral groove close below adapical suture, forming subsutural band. Adapical tips of axials form nodes on subsutural band. Subsutural band may split into two or three narrow-spaced spiral cords by intercalation of secondary spiral grooves. Spiral cords appear between third to fifth teleoconch whorl below subsutural band, forming nodes or beads at intersections with axial ribs and grooves in axial interspaces. Number of spiral cords variable due to intercalations of secondary cords. Last whorl convex, moderately constricted at base. Axial sculpture weakening on base and fasciole, replaced by about 10 to 12 spiral cords, some extending from columellar folds. Aperture elongate, moderately narrow; columellar callus indistinct. Columella with three folds, weakening abapically. Outer lip thin, with about 11 to 15 long, delicate lirae within. Siphonal canal long, weakly deflected, with very shallow siphonal notch. Shell measurements and ratios. SL: 15.3 – 26.5 mm, MD: 4.3 – 7.3 mm; AA: 23 – 30 °, SL / MD: 3.4 – 4.1, AL / AW: 4.7 – 5.6, AH / S: 2.1.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFC6FFEE8C92FF39FB7406C2.taxon	discussion	Discussion. Vexillum michelottii is a rather polymorphic species concerning the spire angle and especially the sculpture on fully grown specimens. The sculpture ranges from rectangular nodulose to densely beaded depending on a variable number of intercalations of secondary spiral cords. Nevertheless, all available specimens display identical sculpture on early teleoconch whorls and have only three columellar folds. Therefore, we consider all specimens to represent a single species. The specimen from Lăpugiu de Sus, discussed by Hoernes & Auinger (1880) as “ variety ” of V. michelottii, has numerous healed fractures from repeated attacks by crabs or other predators (Fig. 16 M). The aberrant outline and sculpture is probably pathological. Palaeoenvironment. The localities Baden and Möllersdorf belong to the Baden Formation suggesting middle neritic settings with up to 250 m water depth (Hohenegger et al. 2008).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFC6FFEE8C92FF39FB7406C2.taxon	distribution	Distribution in Central Paratethys. Badenian (middle Miocene): North Alpine-Carpathian Foredeep: Hrušovany nad Jevišovkou (Czech Republic) (Hoernes & Auinger 1880); Vienna Basin: Baden, Möllersdorf (Austria), Rudice (Czech Republic) (Hoernes & Auinger 1880); Făget Basin: Lăpugiu de Sus (Romania).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFC6FFE98C92F8BDFB5C05FC.taxon	description	Fig. 18 B	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFC6FFE98C92F8BDFB5C05FC.taxon	materials_examined	Type material. The syntypes have been deposited in the collection of Natural History Museum Vienna (Schaffer 1898) but cannot be located and no additional material is known. Therefore, we reproduce the illustration of Schaffer (1898), fig. 18 B. Shell measurements and ratios. SL: 20.1 mm, MD: 8.0 mm; AA: 36 °.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFC6FFE98C92F8BDFB5C05FC.taxon	discussion	Discussion. The description by Schaffer (1898) is not clear and does not fully agree with the illustration. The illustration shows a large broad fusiform shell. The teleoconch whorls bear a broad coronate subsutural band delimited by a deep spiral groove, coinciding with a slightly angulated periphery. The sculpture consists of a dense pattern of opisthocline axial ribs and delicate spiral cords, being most prominent in the axial interspaces. The last whorl is convex, strongly constricted, with orthocline axial ribs and a moderately long siphonal canal. According to Schaffer (1898), the columella bears five columellar folds. Without additional material, the status of this species remains dubious. A further problem arises from the fact that Vexillum nitidum is secondary homonym of Vexillum nitidum Bellardi, 1887 (originally introduced as Uromitra nitida by Bellardi, 1887 b: 45, pl. 5, fig. 49). Therefore, the Paratethyan species would need a new name. As there is no material available of this species, we refrain from introducing a new name. Palaeoenvironment. Only known from the upper Badenian Studienka Formation, which represents middle to outer neritic depositional environments (Biskupič, 2020).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFC6FFE98C92F8BDFB5C05FC.taxon	distribution	Distribution in Central Paratethys. Badenian (middle Miocene): Vienna Basin: Devínska Nová Ves (Slovakia) (Schaffer 1898). Vexillum nitidum (Schaffer, 1898) [non Bellardi, 1887], reproduction of illustration of syntype by Schaffer (1898, fig. 4). C 1 – C 2. Pusia brevior (Friedberg, 1911), reproduction of illustration of lectotype by Friedberg (1911, pl. 1, fig. 17).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFC1FFE88C92F9E3FE280219.taxon	description	Figs 14 E 1 – E 2, F 1 – F 2, G	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFC1FFE88C92F9E3FE280219.taxon	materials_examined	Type material. Holotype: Z 40033, SL: 11.7 mm, MD: 4.4 mm, Rohožník – Konopiská (Slovakia), illustrated in Biskupič (2020, figs 6 A – B), 14 E 1 – E 2. Paratypes: Z 40034, SL: 6.8 mm, MD: 2.7 mm, Rohožník – Konopiská (Slovakia), figs 14 F 1 – F 2; Z 40035, SL: 6.3 mm, MD: 2.6 mm, Rohožník – Konopiská (Slovakia), fig 14 G. All types stored in the Natural History Museum of Slovak National Museum, Bratislava, Slovakia. Revised description. Shell medium sized with slightly scalate spire. Protoconch high conical of three smooth whorls. Teleoconch of seven subcylindrical whorls. Sculpture of prominent, convex, orthocline to weakly opisthocline axial ribs (up to 20 on penultimate whorl), separated by interspaces of about equal width. Interspaces with spiral cords, separated by narrow grooves, not crossing axial ribs. Weakly nodulose subsutural cord. Last whorl rapidly contracting with deep basal concavity. Rounded spiral cords on base and fasciole. Aperture moderately wide. Columellar callus indistinct. Columella with four prominent folds. Outer lip lirate. Siphonal canal moderately short, straight, with shallow siphonal notch.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFC1FFE88C92F9E3FE280219.taxon	discussion	Discussion. This species is reminiscent of the extant Indo-West Pacific Vexillum micra Pilsbry, 1921, which differs mainly in its subhorizontal columellar folds and more prominent spiral cords. Vexillum pseudoschafferi differs from Pusia schafferi (Meznerics, 1933) in its sculpture, but differs in its stout shape, scalate spire, and the presence of a subsutural cord. Palaeoenvironment. Shallow marine environments in corallinacean marls (Biskupič 2020).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFC1FFE88C92F9E3FE280219.taxon	distribution	Distribution in Central Paratethys. Badenian (middle Miocene): Vienna Basin: Rohožník – Konopiská (Slovakia) (Biskupič 2020).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFC0FFE88C92FDC4FF69065B.taxon	description	Figs 14 C 1 – C 2, D	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFC0FFE88C92FDC4FF69065B.taxon	materials_examined	Type material. Lectotype (designated herein): Illustrated in Halaváts (1884, pl. 4. fig. 5), Inv. nr. M 107, Mining and Geological Survey of Hungary, SL: 16.9 mm, MD: 5.0, Szob (Hungary), figs 14 C 1 – C 2. Paralectotype: Inv. nr. M 107, Mining and Geological Survey of Hungary, SL: 14.0 mm, MD: 4.2 mm, Szob (Hungary), fig. 14 D. Revised description. Shell, moderately large, slender fusiform, with high spire. Protoconch unknown. Teleoconch of nine whorls. Early spire whorls weakly convex, with periphery slightly below mid-whorl, passing to subcylindrical on late spire whorls, separated by weakly incised suture. Early teleoconch whorls nearly smooth, with indistinct opisthocline axial swellings, abapically strengthening into wide-spaced, low, convex axial ribs on sixth to eighth teleoconch whorls. Onset of spiral sculpture coinciding with axial sculpture, consisting of numerous weak spiral cords, most prominent in axial interspaces, but also overriding axial ribs. Last whorl short, subcylindrical, moderately constricted. Aperture wide. Columellar callus indistinct. Columella with four oblique columellar folds. Siphonal canal, moderately long, wide, straight, with shallow siphonal notch. Shell measurements and ratios. SL: 14.0 – 16.9 mm, MD: 4.2 – 5.0 mm; AA: 25 – 28 °, SL / MD: 3.5, AL / AW: 3.9, AH / S: 2.2.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFC0FFE88C92FDC4FF69065B.taxon	discussion	Discussion. This species is unique amongst the Paratethyan costellariids in being characterised by a change of sculpture from nearly smooth on early spire whorls to distinct axial and spiral sculpture on later whorls, with the transition occurring on about the seventh teleoconch whorl. Palaeoenvironment. The mollusc fauna and lithofacies of Szob suggest a sandy coastal environment with sea grass (Dulai 1996).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFC0FFE88C92FDC4FF69065B.taxon	distribution	Distribution in Central Paratethys. Badenian (middle Miocene): Pannonian Basin: Szob (Hungary) (Strausz 1966).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFC0FFEA8C92F905FE6400D4.taxon	description	Figs 19 I 1 – I 2, J 1 – J 2, K 1 – K 2, L 1 – L 2	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFC0FFEA8C92F905FE6400D4.taxon	materials_examined	Type material. Holotype: NHMW 1846 / 0037 / 0106, Steinebrunn (Austria), SL: 19.1 mm, MD: 6.3 mm, illustrated in Hörnes (1852 b, pl. 10 figs 29 a – b), figs 19 I 1 – I 2. Paratypes: NHMW 2020 / 0094 / 0001, SL: 16.1 mm, MD: 5.7 mm, Lăpugiu de Sus (Romania); figs 19 J 1 – J 2; NHMW 2020 / 0095 / 0001, SL: 19.3 mm, MD: 6.3 mm, Steinebrunn (Austria); figs 19 L 1 – L 2; NHMW 2020 / 0096 / 0001, SL: 19.1 mm, MD: 5.9 mm, Niederleis (Austria); figs 19 K 1 – K 2. Additional material. NHMW 1860 / 0001 / 0109, Steinebrunn (Austria); NHMW 1860 / 0001 / 0109, 36 specimens, Steinebrunn (Austria); NHMW 1865 / 0001 / 0693, Baden (Austria); NHMW 1869 / 0001 / 0440, Forchtenau (Austria) NHMW 1868 / 0001 / 0080, Möllersdorf (Austria); NHMW 1847 / 0037 / 0046, Rohrbach bei Mattersburg (Austria); NHMW 2020 / 0096 / 0002, 6 specimens, Niederleis (Austria); NHMW 1867 / 0019 / 0042, 4 specimens, Coşteiu de Sus (Romania); NHMW 1868 / 0001 / 0406, 38 specimens, Lăpugiu de Sus (Romania); NHMW 1863 / 0015 / 0070, Bujtur (Romania). Type locality. Steinebrunn (Austria). Type stratum. Silty sand of the Baden Formation. Age. Middle Miocene, Badenian (Langhian).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFC0FFEA8C92F905FE6400D4.taxon	etymology	Etymology. Latin transalpinum, meaning beyond the Alps seen from Rome, referring to the paleogeographic occurrence of the new species.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFC0FFEA8C92F905FE6400D4.taxon	diagnosis	Diagnosis. Vexillum species of large size, slender to moderately slender profile, with conical, weakly gradate spire, multispiral protoconch, dense axial sculpture on early teleoconch whorls, wider spaced abapically, sharp crested or convex orthocline to opisthocline axial ribs on penultimate and last whorls.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFC0FFEA8C92F905FE6400D4.taxon	description	Description. Shell large, solid, slender to moderately slender fusiform, with moderately high conical spire; early spire weakly gradate. Protoconch multispiral, high conical, of 3.5 convex whorls (height: 990 μm, diameter: 750 μm). Teleoconch of seven weakly convex to subcylindrical whorls, separated by impressed suture. Axial sculpture on early teleoconch whorls of broad, convex, orthocline to opisthocyrt axial ribs separated by narrow interspaces (about 11 axial ribs on first teleoconch whorl). Axial ribs and interspaces usually distinctly widening during ontogeny, ribs becoming sharp-crested. No spiral sculpture, except for about five indistinct spiral cords on weak fasciole, subobsolete in most specimens. Last whorl moderately convex, constricted at base. Aperture elongate, moderately narrow, with faint anal canal. Parietal region slightly thickened in many specimens. Columellar callus indistinct. Columella with four oblique folds, weakening abapically. Outer lip thin, with about ten lirae within. Siphonal canal moderately short, slightly bent to left, with wide, shallow siphonal notch. Shell measurements and ratios. SL: 14.8 – 22.0 mm, MD: 4.2 – 7.0 mm; AA: 30 – 38 °, SL / MD: 2.9 – 3.2, AL / AW: 3.7 – 4.5, AH / S: 2.1 – 2.9.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFC0FFEA8C92F905FE6400D4.taxon	discussion	Discussion. This species was confused with the Pliocene proto-Mediterranean Vexillum pyramidellum (Brocchi, 1814) by all authors describing middle Miocene Paratethyan faunas [note that Vexillum plicatulum (Brocchi, 1814) is a subjective synonym of V. pyramidellum (Brocchi, 1814) (Della Bella & Tabanelli 1995; Landau et al. 2013)]. Like Vexillum pyramidellum, V. transalpinum is variable in having a more-or-less slender outline and sharply crested axial ribs. The number of axial ribs on the penultimate and last whorls is variable ranging from prominent, densely set axial ribs (rare) to wide-spaced ones. Nevertheless, the axial sculpture is never completely reduced on penultimate and last whorls, as is typical for the Pliocene specimens of Vexillum pyramidellum (Brocchi, 1814), as described and illustrated by Bellardi (1887 b), Cerulli-Irelli, S. (1911), Rossi-Ronchetti (1955), Malatesta (1974), Pinna & Spezia (1978) and Chirli (2002). Vexillum intermittens (Hoernes & Auinger, 1880) is reminiscent of V. transalpinum, but is distinguished by its more slender outline, higher spire and opisthocline axial ribs on early teleoconch whorls. Palaeoenvironment. The species is abundant in inner neritic and middle neritic depositional environments and is found in sandy-silty nearshore deposits and in basinal clay.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFC0FFEA8C92F905FE6400D4.taxon	distribution	Distribution in Central Paratethys. Badenian (middle Miocene): Vienna Basin: Niederleis, Baden, Möllersdorf, Steinebrunn (Hoernes & Auinger 1880; Sieber 1958 b); Eisenstadt-Sopron Basin: Mattersburg (Austria) (Hörnes 1852 b);? Pannonian Basin: Sámsonháza (Hungary) (Csepreghy-Meznerics 1954); Făget Basin: Lăpugiu de Sus, Coşteiu de Sus, Bujtur (Romania); Southern Carpathians: Bahna (Romania) (Tiţă 2007). Proto-Mediterranean Sea. Serravallian (middle Miocene): Karman Basin: Lale, Akboðaz, Akpýnar, Tavþanlý (Turkey) (Landau et al. 2013).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFCDFFE58C92FF39FA1404FE.taxon	type_taxon	Type species. Mitropifex isaoi Kuroda & Sakurai in Kuroda, 1959, by monotypy, Present-day, Indo-West Pacific.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFCDFFE58C92FF39FA1404FE.taxon	description	Description. “ Shell elongate fusiform, sometimes widely fusiform or turriform, typically with high spire and long to very long siphonal canal. Suture typically canaliculated or impressed, often resulting in telescopic appearance of spire. Whorls outline varying from subcylindric to flattened or evenly convex. Axial sculpture of straight ribs strong and widely set to fine and closely set. Spiral sculpture varying from fine grooves, restricted to siphonal canal or pronounced in interspaces between axials only, to strong cords overriding axial ribs to form beads or strong gemmae at intersections. Aperture elongate, its outer lip smooth or bearing fine lirae on its inner surface. Inner lip with between three and five columellar folds, with the upmost fold being the strongest ” (Fedosov et al., 2017: 564).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFCDFFE58C92FF39FA1404FE.taxon	discussion	Discussion. The large and slender fusiform Mitra neudorfensis Schaffer, 1898 and Vexillum pseudocupressinum Bałuk, 1997 have been placed so far in Vexillum by Biskupič (2020) and Bałuk (1997). Alternatively, we propose a placement in Tosapusia Habe, 1964. The genus Tosapusia differs from Vexillum especially in its longer siphonal canal, which lacks a deep notch. In addition, axial ribs in Tosapusia are typically straight, but tend to be arcuate and undulating in Vexillum (Fedosov et al. 2017). Tosapusia pseudocupressina is strikingly similar to the extant Indo-West Pacific Tosapusia evelynae (Guillot de Suduiraut, 2007) in outline and both species display a comparable variability in sculpture. Tosapusia isaoi Kuroda & Sakurai in Kuroda, 1959 is a modern analogue of T. neudorfensis, but lacks a subsutural groove. Moreover, the preference for offshore settings of the two discussed Miocene species is consistent with the deep water habitat of extant Tosapusia species. A further species from the European Neogene, which should be placed in Tosapusia, is T. cupressina (Brocchi, 1814) from the middle Miocene to Pliocene of the proto-Mediterranean Sea (Landau et al. 2013). Extant Tosapusia species have an Indo-West Pacific distribution (Fedosov et al. 2007) but reached to the protoMediterranean Sea and the Central Paratethys during the Miocene. The genus persisted in the proto-Mediterranean Sea until the Pliocene, clearly postdating the Tethys closure during the early Miocene (Harzhauser et al. 2007). Therefore, our data document a restriction of the former distribution area of Tosapusia and should not be misinterpreted as immigration from the IWP region.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFCDFFE48C92FAFCFCFE0434.taxon	description	Figs 19 A 1 – A 2, B 1 – B 2, C 1 – C 2, D 1 – D 2, 18 A	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFCDFFE48C92FAFCFCFE0434.taxon	materials_examined	Type material. The syntypes have been deposited in the collection of Natural History Museum Vienna (Schaffer, 1898) but cannot be located. Additional material. SNM-PM Z 40040, SL: 29.2 mm, MD: 9.6 mm, Konopiská clay pit at Rohožník, Slovakia, middle Miocene (Badenian), Natural History Museum of Slovak National Museum, Bratislava, Slovakia, illustrated in Biskupič (2020, figs 2 A – B), holotype of Vexillum svagrovskyi Biskupič, 2020, figs 19 A 1 – A 2; SNM-PM Z 40045, SL: 40.2 mm, MD: 12.5 mm, paratype of Vexillum svagrovskyi Biskupič, 2020, illustrated in Biskupič (2020, figs 3 C – D), figs 19 B 1 – B 2; SNM-PM Z 40046, SL: 30.3 mm, MD: 10.4 mm, paratype of Vexillum svagrovskyi Biskupič, 2020, illustrated in Biskupič (2020, figs 3 E – F), figs 19 C 1 – C 2; SNM-PM Z 40053, illustrated in Biskupič (2020, figs 2 G – H), figs 19 D 1 – D 2. 7 specimens, SNM-PM Z 40041 – Z 40047, all from Konopiská clay pit at Rohožník, Slovakia, middle Miocene (Badenian), Natural History Museum of Slovak National Museum, Bratislava, Slovakia, paratypes of Vexillum svagrovskyi Biskupič, 2020. Revised description. Shell large, moderately slender fusiform, solid, with weakly gradate spire. Protoconch unknown. Teleoconch of 11 whorls; spire whorls subcylindrical, with impressed suture. Axial sculpture of early teleoconch whorls of densely set opisthocline axial ribs, separated by narrower interspaces. Axial ribs weakening on seventh to ninth teleoconch whorl, becoming slightly more widely spaced and blurred on penultimate and last whorls. Spiral cords most prominent on early teleoconch whorls in axial interspaces, separated by narrow grooves. Broad subsutural band formed by deep spiral groove cutting axial ribs. Spiral band solid, or split into three to five densely spaced spiral cords. All spiral sculpture becoming subobsolete on penultimate and last whorls, rarely subsutural band persisting as weak spiral cord. Last whorl moderately convex, slowly contracting at base, covered by weakly sigmoidal growth lines. Several broad, flat spiral cords with delicate axial threads on weak fasciole. Aperture narrow with indistinct anal sinus; columellar callus indistinct, poorly delimited. Columella with four to five folds, weakening rapidly abapically. Outer lip thin without lirae or denticles. Siphonal canal moderately long and wide, straight, with shallow siphonal notch. [Note that this description is based on the holotype and paratypes of Vexillum svagrovskyi Biskupič, 2020, except for the fifth columellar fold, which is only known from the syntypes of V. neudorfense]. Shell measurements and ratios. SL: 28.5 – 40.2 mm, MD: 9.6 – 12.5 mm; AA: 30 – 34 °, SL / MD: 3.0 – 3.2, AL / AW: 4.3 – 5.1, AH / S: 1.6 – 2.2.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFCDFFE48C92FAFCFCFE0434.taxon	discussion	Discussion. This species is characterised by the presence of a distinct spiral groove a short distance below the suture that bisects the ribs, resulting in a broad subsutural band. The syntypes of Tosapusia neudorfensis seem to be lost, but the illustration in Schaffer (1889) is rather precise. Based on that illustration, it is most likely that Vexillum svagrovskyi Biskupič, 2020 is a junior synonym of T. neudorfensis. Both taxa are only known from the upper Badenian Studienka Formation in Slovakia. Both species are identical in shape and sculpture and the only difference is the weak fifth columellar fold of Tosapusia neudorfensis. We assume that the presence of a fifth columellar fold is rather a matter of intraspecific variability. Palaeoenvironment. Only known from the upper Badenian Studienka Formation, which represents middle to outer neritic depositional environments (Biskupič 2020).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFCDFFE48C92FAFCFCFE0434.taxon	distribution	Distribution in Central Paratethys. Badenian (middle Miocene): Vienna Basin: Devínska Nová Ves, Rohožník – Konopiská (Slovakia) (Schaffer 1898; Biskupič 2020).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFCCFFE78C92FB28FA000771.taxon	description	Figs 4 O, 19 E 1 – E 2, F 1 – F 2, G 1 – G 2, H 1 – H 2	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFCCFFE78C92FB28FA000771.taxon	materials_examined	Type material. Holotype. NHMW 1846 / 0037 / 0103, SL: 42.7 mm, MD: 13.5 mm, Baden (Austria), illustrated in Hörnes (1852 b, pl. 10, fig. 25), figs 19 G 1 – G 2. Additional material. NHMW 2020 / 0090 / 0001, SL: 38.1 mm, MD: 11.5 mm, Baden (Austria), illustrated in Hörnes (1852 b, pl. 10, fig. 26), figs 19 F 1 – F 2; NHMW 2020 / 0091 / 0001, SL: 34.9 mm, MD: 9.3 mm, Möllersdorf (Austria), figs 19 E 1 – E 2; NHMW 1870 / 0033 / 0037, SL: 27.7 mm, MD: 7.8 mm, Lăpugiu de Sus (Romania); figs 19 H 1 – H 2; NHMW 2020 / 0091 / 0002, SL: 23.8 mm, MD: 6.5 mm, Möllersdorf (Austria), illustrated in Hörnes (1852 b, pl. 10, fig. 27); NHMW 2020 / 0091 / 0003, 8 specimens, Möllersdorf (Austria); NHMW 2020 / 0091 / 0004, SL: 31.8 mm, MD: 10.0 mm, Möllersdorf (Austria), illustrated in Hoernes & Auinger (1880, pl. 10, figs 10 a – b); NHMW 1862 / 0001 / 0250, 17 spec., Möllersdorf (Austria); NHMW 1846 / 0037 / 0103, Vienna / Grinzing (Austria); NNHM 2020 / 0092 / 0001 (= A 1562), 5 specimens, Baden-Sooss (Austria); NHMW 1969 / 0001 / 0023, 25 specimens, Bad Vöslau (Austria); NHMW 1846 / 0037 / 0104, 24 specimens, Baden (Austria); NNHM 1866 / 0001 / 0631, 13 specimens, Baden-Sooss (Austria); NHMW 1853 / 0003 / 0089, Forchtenau (Austria); NHMW 1862 / 0029 / 0008, Rudice (Czech Republic); NHMW 1854 / 0035 / 0095, 45 specimens, Lăpugiu de Sus (Romania), fig. 4 O; NHMW 1870 / 0033 / 0037, 17 specimens, Lăpugiu de Sus (Romania); NHMW 1867 / 0019 / 0035, 4 specimens, Coşteiu de Sus (Romania). Revised description. Shell large, slender fusiform, with deeply incised suture. Protoconch (fig. 4 O) high conical, of three smooth, convex whorls (height: 820 μm, diameter: 620 μm). Teleoconch of ten weakly convex to subcylindric whorls. Last whorl moderately convex, slowly contracting into high base. Some specimens with weak shoulder and faint angulation close below adapical suture, especially on penultimate and last whorls. Sculpture on spire whorls consisting of close-set, prominent, weakly opisthocline rounded axial ribs, separated by slightly narrower interspaces, crossed by close-set spiral cords, separated by narrow grooves, forming indistinct nodules at intersections with ribs, and distinct, flat cords in axial interspaces (about 20 axial ribs on penultimate whorl). Axial ribs crossed by subsutural spiral groove, delimiting weak subsutural band, most prominent on early teleoconch whorls. Subsutural band often disintegrating into narrow spiral cords during growth. Axial ribs slightly more widely spaced on last three whorls. Axial and spiral sculpture often reduced on last whorl. Spiral cords more wide-spaced on base and fasciole. Aperture narrow, elongate; columellar callus narrow, weakly delimited, bearing four prominent columellar folds, weakening abapically. Outer lip thin, without lirae in adult specimens; delicate lirae deep inside aperture only in subadult specimens. Siphonal canal long, moderately narrow, straight, with shallow siphonal notch. Shell measurements and ratios. SL: 20.1 – 42.0 mm, MD: 5.8 – 12.8 mm; AA: 25 – 35 °, SL / MD: 3.3 – 3.6, AL / AW: 5.0 – 5.8, AH / S: 1.9 – 2.1.	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFCCFFE78C92FB28FA000771.taxon	discussion	Discussion. This species was identified as Vexillum cupressinum (Brocchi, 1814) by all authors dealing with Paratethyan mollusc faunas until Bałuk (1997) separated it as Vexillum pseudocupressinum. Bałuk (1997) stressed the more densely spaced axial ribs and higher number of spiral cords being restricted to the axial interspaces. Landau et al. (2013) doubted if these features justify a separation, referring to the high variability in sculpture in late Miocene and Pliocene populations of Tosapusia cupressina, as documented by Pelosio (1967), Malatesta (1974), Davoli (2000), Chirli (2002) and Chirli & Richard (2008). After having studied a large number of Paratethyan specimens, we agree with Bałuk (1997). Despite the variability of T. cupressina, it differs constantly by its lower number of axial ribs (especially on early spire whorls). Its spiral cords are less numerous and form a cancellate or nodulous pattern with the axials ribs, lacking the deeply cut grooves seen in the axial interspaces in T. pseudocupressina. Moreover, the Paratethyan species is nearly twice as large as T. cupressina and it lacks the twisted siphonal canal of the Pliocene species. Note that the specimen illustrated by Hoernes & Auinger (1880, pl. 10, fig. 10) with strongly deflected siphonal canal is a pathological specimen with healed fractures. Palaeoenvironment. The species is most abundant in clay of the Baden Formation suggesting middle neritic settings with up to 250 m water depth (Hohenegger et al. 2008).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
D446F043FFCCFFE78C92FB28FA000771.taxon	distribution	Distribution in Central Paratethys. Badenian (middle Miocene): Korytnica Basin: Korytnica (Poland) (Bałuk 1997); Ukrainian Fore-Carpathian Basin: Dryszczów (Nadrichne) (Ukraine), Żukowce (Zhukivtsi) (Friedberg 1911); Vienna Basin: Baden, Bad Vöslau, Möllersdorf, Steinebrunn, Vienna / Grinzing (Austria), Rudice (Czech Republic) (Hoernes & Auinger 1880; Sieber 1958 b); Eisenstadt-Sopron Basin: Forchtenau (Sieber 1956); Făget Basin: Coşteiu de Sus (Boettger 1906); Pannonian Basin: Szob (Hungary) (Strausz 1966); Bükk Mountains (Hungary) (Csepreghy-Meznerics 1972); Făget Basin: Lăpugiu de Sus, Coşteiu de Sus (Romania) (Hoernes & Auinger 1880); Transylvanian Basin: Zorlentul-Mare (Romania) (Florei 1961); Southern Carpathians: Bahna (Romania) (Tiţă 2007); Dacian Basin: Opanec, Orehovica (Bulgaria) (Kojumdgieva in Kojumdgieva & Strachimirov 1960).	en	Harzhauser, Mathias, Landau, Bernard (2021): An overlooked diversity-the Costellariidae (Gastropoda: Neogastropoda) of the Miocene Paratethys Sea. Zootaxa 4982 (1): 1-70, DOI: https://doi.org/10.11646/zootaxa.4982.1.1
