Tiarapirenella tabulata (Hörnes, 1856) Harzhauser & Guzhov & Landau, 2023

Harzhauser, Mathias, Guzhov, Aleksandr & Landau, Bernard, 2023, A revision and nomenclator of the Cainozoic mudwhelks (Mollusca: Caenogastropoda: Batillariidae, Potamididae) of the Paratethys Sea (Europe, Asia), Zootaxa 5272 (1), pp. 1-241 : 141-144

Tiarapirenella tabulata (Hörnes, 1856) nov. comb.

Figs 28A–H

* Melanopsis tabulata Ĥrn. —Ĥrnes 1856: 600, pl. 49, fig. 15. Cerithium nodoso-striatum Hilb. — Hilber 1879a: 437, pl. 4, figs 1a–c. Cerithium Sturi Hilb. — Hilber 1879a: 442, pl. 4, figs 8a–c [non Stoliczka in Stur 1863: 50]. Cerithium dionysii Hilber 1882: 7 [nov nom. pro Cerithium sturi Hilber, 1879a , non Stoliczka in Stur 1863]. Potamides variabilis Friedb. — Friedberg 1914: 282, pl. 17, figs 27–28. C [erithium]. Kontkiewiczi — Friedberg 1914: 283 [nomen nudum]. Pirenella Schaueri Hilber var. Eichwaldi R. Ĥrn. U. Auing.— Sieber 1937: 482, pl. 24, figs B 3, 4 [non Friedberg, 1914]. Potamides variabilis Friedb. — Friedberg 1938a: 102. Potamides moravicus M. Ĥrnes — Csepreghy-Meznerics 1950: 28, pl. 1, fig. 13 [non Granulolabium moravicum ( Ĥrnes,

1855)]. Pirenella moravica variabilis (Friedberg) — Papp 1952: 113, pl. 3, figs 11–17. Pirenella moravica (M. Hoernes) — Papp 1952: 113. Pirenella moravica dionysii (Hilber) — Papp 1952: 113. Pirenella moravica Ĥrnes, 1856 — Strausz 1955a: 47, 168, text-fig. 2, 8 [non Granulolabium moravicum ( Ĥrnes, 1855) ]. Pirenella moravica variabilis Friedberg, 1928 — Strausz 1955a: 52, 175, pl. 4, figs 44–45, 48, 49, 53, pl. 5, figs 54–57, pl. 6,

figs 80–86, 88–94. Pirenella moravica palatinotiara nov. f.— Strausz 1955a: 53, pl. 5, figs 63–66. Pirenella moravica pseudonympha nov. f.— Strausz 1955a: 53, pl. 5, figs 75–79. Pirenella sturi Hilber, 1879 — Strausz 1955a: 56, 179, pl. 6, figs 97–98 [non Stoliczka in Stur, 1863: 50]. P [otamides]. moravica dionysii (Hilb.) — Sieber 1958: 136. Pirenella moravica variabilis (Friedberg) — Pavlovsky 1960: 215, pl. 1, 6a. Pirenella picta var. bicostata ( Eichwald 1853) —Kojumdgieva in Kojumdgieva & Strachimirov 1960: 108, pl. 31, fig. 17 [non

Cerithium bicostatum Eichwald, 1852 = Tiaracerithium pictum (de Basterot, 1825) ]. Pirenella moravica variabilis Friedb. — Florei 1961: 683, pl. 7, fig. 46. Pirenella moravica variabilis (Friedberg) — Mariani & Papp 1966: 143. Potamides (Pirenella) moravicus Ĥrnes, 1856 — Strausz 1966: 147, pl. 7, figs 23–29, pl. 8, figs 5, 10–11, pl. 10, figs 9–10 [non

Granulolabium moravicum ( Ĥrnes, 1855) ]. Potamides (Pirenella) moravicus variabilis Friedberg, 1928 — Strausz 1966: 149, pl. 7, figs 30–35. Potamides (Pirenella) moravicus pseudonympha Strausz, 1955 — Strausz 1966: 150, pl. 8, figs 12–14. Potamides (Pirenella) sturi Hilber, 1879 — Strausz 1966: 150, pl. 7, figs 36–37 [non Stoliczka in Stur, 1863: 50]. Potamides (Pirenella) moravicus palatinotiara Strausz, 1955 — Strausz 1966: 150, pl. 8, figs 6–9. Pirenella moravica palatinotiara Strausz 1955 — Boda 1964: 130. Pirenella moravica pseudonympha Strausz 1955 — Boda 1964: 130. Potamides moravica (Ĥrnes) — Kókay 1966: 41, pl. 4, fig. 6 [non Granulolabium moravicum ( Ĥrnes, 1855) ]. Pirenella picta pseudonympha Strausz — Kókay 1966: 41. Pirenella sturi (Hilber) — Kókay 1966: 42 [non Stoliczka in Stur 1863: 50]. Potamides variabilis Friedberg — Csepreghy-Meznerics 1972: 21, pl. 3, fig. 13. Potamides mitralis Eichw. — Csepreghy-Meznerics 1972: 21, pl. 3, fig. 17 [non Tiarapirenella bicincta ( Brocchi, 1814) ]. Potamides (Pirenella) moravicus variabilis Friedberg, 1923 — Nicorici & Sagatovici 1973: 167, pl. 24, figs 8–9. Pirenella moravica (Hörnes, 1856) — Bałuk 1975: 129, pl. 15, figs 9–15 [non Granulolabium moravicum ( Ĥrnes, 1855) ]. Pirenella tabulata (Hörnes, 1856) — Bałuk 1975: 130, pl. 15, figs 5–6. Granulolabium plicatum (Bruguière, 1792) —Harzhauser 2002: 73, pl. 1, figs 17–20 [non Granulolabium plicatum (Bruguière,

1792)]. Pirenella moravica Hörnes, 1856 — Bałuk 2006: 197, pl. 8, fig. 5. [non Granulolabium moravicum ( Ĥrnes, 1855) ].

non Potamides variabilis Friedbg. — Baumberger 1931: 249, pl. 11, fig. 31 [= Thericium].

Type material. Lectotype designated herein: NHMW 1855/0045/0790, SL: 30.4 mm, MD: 10.0 mm, Natural History Museum Vienna, illustrated in Ĥrnes (1856: pl. 49, fig. 15), Figs 28A 1 –A 3 . Paralectotypes: NHMW 1851/0002/0060, SL: 31.7 mm, MD: 10.9 mm, Grund ( Austria) , Figs 28B 1 –B 3. NHMW 1851/0002/0060, SL: 30.6 mm, MD: 10.9 mm, Grund ( Austria), Figs 28C 1 –C 3. 25 spec., NHMW 1851/0002/0060, Grund ( Austria)

Type locality. Grund ( Austria) .

Stratigraphy. Middle Miocene, Badenian.

Illustrated material. ZNP PAN A-I50/688.2, SL: 23.1 mm, MD: 8.2 mm, Małoszów ( Poland), Badenian (Middle Miocene), lectotype (designated herein) of Potamides variabilis Friedberg, 1914 ; illustrated in Friedberg (1914, pl. 17, fig. 28), Figs 28D 1 –D 2. NHMW1859/0010/0066, SL: 26.3 mm, MD: 9.8 mm, Ŗckersdorf ( Austria), Figs 28E 1 –E 2. NHMW1859/0010/0066, SL: 24.3 mm, MD: 9.4 mm, Ŗckersdorf ( Austria), Figs 28F 1 –F 2. NHMW 1999z0004/0007, SL: 19.8 mm, MD: 8.0 mm, St. Veit an der Triesting ( Austria), illustrated in Sieber (1937: pl. 24, fig. B4) as Pirenella schaueri eichwaldi , Figs 28G 1 –G 2. M.114, SL: 18.5 mm, MD: 6.0 mm, Várpalota ( Hungary), Badenian (Middle Miocene), holotype of Pirenella moravica pseudonympha Strausz, 1955a , Fig 28H.

Studied material. UMJG &P 200.148, SL: 14.5 mm, MD: 6.3 mm, Gamlitz ( Austria), holotype of Cerithium nodosostriatum Hilber, 1879a , illustrated in Hilber (1879a: pl. 4, figs 1a–c). M.17, SL: 21 mm, MD: 5.9 mm, Várpalota ( Hungary), Badenian, Middle Miocene, holotype of Pirenella moravica palatinotiara Strausz, 1955 . Karpatian (Early Miocene): 91 spec., NHMW 1859/0010/0067, Ŗckersdorf ( Austria); Badenian (Middle Miocene): 13 spec., NHMW A866, Grund ( Austria); 20 spec., NHMW 1865/0001/1100, Grund ( Austria); 53 spec., NHMW 2022/0064/0001, Pernersdorf ( Austria); 8 spec., NHMW 1866/0033/0013, Hrušovany nad Jevišovkou ( Czech Republic).

Revised description. Shell medium sized, slender to moderately stocky conical, with weakly gradate spire; apical angle ~20–35°. Protoconch unknown. Early teleoconch whorls moderately convex with two prominent spiral cords bearing broad, flattened beads. Later teleoconch whorls subcylindrical with two primary spiral cords with subquadratic beads. Third cord intercalated between primaries on about third whorl, then fourth cord appears at abapical suture on 4 th to 6 th whorls. Weaker secondary spiral cords with beads of variable strength appear between primary cords. Secondary spiral cords equal primary cords in strength in some specimens. Last whorl high, subcylindrical to slightly convex, bearing about eight primary spiral cords alternating with weaker secondaries highly variable in strength, beads on secondary cords often reduced; base high, slowly contracting. Aperture ovoid, not wide; outer lip not thickened; without granules within; anal canal narrowly incised, delimited medially by weak parietal swelling; siphonal canal short, incised, moderately wide, weakly deflected to the left. Columella excavated, smooth, weakly twisted at siphonal canal. Columellar callus forming moderately broad rim, sharply delimited from base. Top of beads often with intense reddish color.

Synonyms. Cerithium dionysii Hilber, 1882 [replacement name for Cerithium sturi Hilber, 1879a non Stoliczka in Stur, 1863] and Potamides variabilis Friedberg, 1914 represent subjective junior synonyms of Tiarapirenella tabulata . These taxa are based on specimens of rather similar morphology. Slender to extremely slender specimens have been described as Pirenella moravica palatinotiara Strausz, 1955 . Another slender morph with strongly reduced beads and weak spiral cords was named Pirenella moravica pseudonympha Strausz, 1955a . Cerithium nodosostriatum Hilber, 1879a seems to represent an aberrant specimen with multiple secondary spiral threads intercalated between the beaded primary cords. The name Cerithium nodosostriatum is preoccupied by Peters (1855: 364). Cerithium kontkiewiczi Friedberg, 1914 is a nomen nudum based on a collection name by Mathias Auinger (1810–1890) for specimens from Niederkreuzstetten ( Austria). In his description of Potamides variabilis, Friedberg (1914) referred to these specimens, without making the name available.

Discussion. Tiarapirenella tabulata is characterized by its sculpture of primary and secondary spiral cords and differs from G. plicatum by its broader shell, marked secondary spiral sculpture, and the high, large whorl. Moreover, the beads of T. tabulata show very strong reddish coloring, which is unknown from G. plicatum . This species was treated as synonym of Tiarapirenella bicincta ( Brocchi, 1814) by Lozouet (1986: 196). This hypothesis is rejected herein. Tiarapirenella tabulata is much larger than T. bicincta , has a larger last whorl and has more numerous spiral cords and the beads are more rounded and closer-set.

Distribution. This species is endemic to the Central Paratethys Sea. It appeared during the Karpatian (late Burdigalian) and attained its widest distribution during the early and middle Badenian (Langhian). Last records derive from the late Badenian (Serravallian).

Central Paratethys. Karpatian (Early Miocene): Korneuburg Basin: Niederkreuzstetten, Ŗckersdorf ( Austria) (hoc opus). Badenian (Middle Miocene): Korytnica Basin: Korytnica ( Poland) ( Bałuk 1975, 2006); Polish-Ukrainian Fore-Carpathian Basin: Małoszów ( Poland) ( Friedberg 1914); North Alpine-Carpathian Foreland Basin: Grund, Pernersdorf ( Austria) (Ĥrnes 1856), Hrušovany nad Jevišovkou ( Czech Republic) (hoc opus); Vienna Basin: Berndorf (St. Veit an der Triesting) ( Austria) ( Mariani & Papp 1966); Oberpullendorf Basin: Ritzing ( Austria) ( Papp 1952); Klagenfurt Basin: Lavam̧nd ( Austria) ( Papp 1952); Styrian Basin: Gamlitz, St. Florian (( Austria) ( Hilber 1879a; Papp 1952); Pannonian Basin: Herend–Márkó, Hidas, Sámsonháza, Várpalota, ( Csepreghy-Meznerics 1954; Strausz 1966), Samobor ( Croatia) ( Pavlovsky 1960); Bükk Mountains: Borsodbóta ( Csepreghy-Meznerics 1972); Banat: Zorlenţul-Mare ( Romania) ( Florei 1961); Zârand Basin: Minişul de Sus ( Romania) ( Nicorici & Sagatovici 1973); Dacian Basin: Tyrnene ( Bulgaria) (Kojumdgieva in Kojumdgieva & Strachimirov 1960).

Genus Tiaracerithium Sacco, 1895

Type species. Cerithium pseudotiarella d’Orbigny, 1852 (= Cerithium thiarella Grateloup, 1832 ), original designation by Sacco (1895: 35). Early Miocene, France.

Original diagnosis. ” Forme generalmente piccolo, turrito-acute, a spiccato cingula suturale più o meno granuloso.” ( Sacco 1895: 35). [Generally small, turreted, acute shell with more or less prominent sutural cord.]

Revised description. Small to medium sized, slender with gradate spire. Protoconch of 1.5 whorls. Early teleoconch whorls with three faintly beaded or smooth spiral cords, often adjoined by weaker secondary cords. Sculpture of later teleoconch whorls variable, but typically consisting of dominant spiral row of rounded beads at adapical suture. Mid-whorl spiral cord and spiral cord at abapical suture variably beaded, smooth, or reduced. Whorl profile subcylindrical. Last whorl subcylindrical with moderately constricted, convex base. Aperture ovate; outer lip not thickened; devoid of granulae within; siphonal canal short, moderately wide, slightly twisted to the left; anal canal distinctly incised. Columella excavated, smooth with broad callus rim.

Illustrated material. Tiaracerithium thiarella ( Grateloup, 1832) , MNHN.F.R06864, SL: 16.5 mm, MD: 5.2 mm, Léognan ( France), Aquitanian (Early Miocene), photo: Jacques Mouchart, 2018 (e-recolnat Project, MNHN), illustrated in Lozouet (1986: pl. 2, fig. 5) as Granulolabium (Tiaracerithium) pseudotiarella (d’Orbigny) , Figs 29A 1 –A 2. MNHN.F.J05910, SL: 14.1 mm, MD: 4.2 mm, Léognan ( France), Aquitanian (Early Miocene), photo: Jacques Mouchart, 2018 (e-recolnat Project, MNHN), syntype of Pirenella pseudotiarella pictoides Cossmann & Peyrot, 1922 , illustrated in Cossmann & Peyrot (1922: pl. 6 fig. 41), Figs 29B 1 –B 2. MNHN.F.J05908, SL: 15.1 mm, MD: 4.9 mm, Léognan ( France), Aquitanian (Early Miocene), photo: Peter Massicard, 2018, (e-recolnat Project, MNHN), illustrated in Cossmann & Peyrot (1922: pl. 6, fig. 56) as Pirenella pseudotiarella ventricosa (Grateloup, 1847) , Figs 29C–C 2.

Stratigraphy and paleogeography. The first records of this genus occur in the Aquitanian of the northeastern Atlantic. It is documented during the Burdigalian in the Proto-Mediterranean Sea, the northeastern Atlantic and the Central Paratethys and became ubiquitous during the Middle Miocene, when it spread also into the Eastern Paratethys Sea. It survived the Badenian–Sarmatian Extinction Event ( Harzhauser & Piller 2007) and was the dominant gastropod on Sarmatian mudflats in the Paratethys. Tiaracerithium disappeared from the Paratethys by the end of the Late Miocene Bessarabian and is not known from the latest Miocene Maeotian. In the Proto-Mediterranean Sea it persisted throughout the Tortonian and Messinian ( Sacco 1895; D’Amico et al. 2012; Dominici et al. 2020). Tiaracerithium eceae Landau, Harzhauser, Ýslamoðlu & Marques da Silva, 2013 , is a second Middle Miocene Tiaracerithium species in the Proto-Mediterranean Sea, but is known so far only from the Serravallian of Turkey. Pliocene records from the Mediterranean (usually misidentified as “ Granulolabium bicinctum ” are rare (“ alquanto rara ” Sacco 1895: 60) or even based on misidentifications (e.g., Chirli 2006). Similarly, records from the North Sea are based on misidentifications (e.g., Wesselingh et al. 2012). Therefore, we assume that the genus disappeared at the end of the Messinian and the scattered Pliocene records may represent reworked specimens. The genus did not manage to spread into the Indo-West Pacific region, probably because its main radiation postdated the closure of the Tethyan Seaway and because its ecological niches were settled there by Pirenella .

Ecology. Species of Tiaracerithium occurred in enormous numbers in the littoral zone. Harzhauser & Kowalke (2002) documented clear differences in morphology between specimens of Tiaracerithium pictum (misidentified as Granulolabium bicinctum ) from oligohaline mudflats and specimens from sandy littoral to shallow sublittoral marine environments. Specimens from mudflat assemblages display reduced sculpture and conical shells, whereas specimens from more marine settings display more prominent sculpture and more gradate spires. Pure mangrove environments, however, seem to have been avoided by Tiaracerithium .

Discussion. Early teleoconch whorls of Granulolabium plicatum bear three prominent spiral cords of beads on conical to subcylindrical whorls, whereas Tiaracerithium lacks distinct beads on early teleoconch whorls ( Table 2). The adapical spiral row of beads is always very prominent in Tiaracerithium , whereas the mid-whorl cord and the cord at the abapical suture may be reduced. The formation of the beads is not strictly homologous in both genera. In Granulolabium the beads develop along the spiral cords and are limited by the deep spiral grooves, which often results in axially elongated morphologies. Thus, the beads of Granulolabium are threaded like pearls on the spiral cords whereas the beads in Tiaracerithium are spirally arranged without being placed on top of the underlying spiral cords. Consequently, the beads of Tiaracerithium are broadly rounded or subquadratic without marked horizontal limitations. Moreover, Tiaracerithium lacks any granulae inside the aperture, which are eponymous for Granulolabium [we note, however, that the presence of granulae is a variable feature even within G. plicatum and granulae may be entirely absent in some species (see Kadolsky 1995: 13 for G. omnimo Kadolsky, 1989 )].

Species-level taxa placed in Tiaracerithium reported from the Paratethys ascalarata. Potamides . Friedberg, 1914 → Tiaracerithium pictum (de Basterot, 1825) athanasiui . Cerithium . Simionescu & Barbu, 1940 → Tiaracerithium pictum (de Basterot, 1825) bessarabica. Pirinella. Simion. & Barbu, 1940 → Tiaracerithium pictum (de Basterot, 1825) bicinctum . Cerithium . Eichwald, 1852 → Tiaracerithium pictum (de Basterot, 1825) bijugum . Cerithium . Eichwald, 1852 → Tiaracerithium pictum (de Basterot, 1825) bizonata. Cerithium . Eichwald, 1851 → Tiaracerithium pictum (de Basterot, 1825) bulgarica. Potamides . Friedberg, 1938a → Tiaracerithium pictum (de Basterot, 1825) conitesta . Seila . Švagrovský, 1954 → Tiaracerithium pictum (de Basterot, 1825) costatonodosa . Seila . Švagrovský, 1954 → Tiaracerithium pictum (de Basterot, 1825) elongatum . Cerithium . Handmann, 1883b → Tiaracerithium pictum (de Basterot, 1825) feldbachensis. Cerithium . Stiny, 1925 → Tiaracerithium pictum (de Basterot, 1825) florianum . Cerithium . Hilber, 1879a → Tiaracerithium pictum (de Basterot, 1825) heptastichus . Cerithium . Quenstedt, 1884 → Tiaracerithium pictum (de Basterot, 1825) hexastichus . Cerithium . Quenstedt, 1884 → Tiaracerithium pictum (de Basterot, 1825) interpolis. Cerithium . Handmann, 1883a → Tiaracerithium pictum (de Basterot, 1825) laevigatum . Cerithium . Eichwald, 1830 → Tiaracerithium pictum (de Basterot, 1825) lineata. Cerithium . Handmann 1883b → Tiaracerithium pictum (de Basterot, 1825) lineolatum . Cerithium . Sowerby, 1832 → Tiaracerithium pictum (de Basterot, 1825) mitrale . Cerithium . Eichwald, 1830 → Tiaracerithium pictum (de Basterot, 1825) mitreolum . Cerithium . Eichwald, 1851 → Tiaracerithium pictum (de Basterot, 1825) nodosa . Cerithium Simionescu & Barbu, 1940 → Tiaracerithium pictum (de Basterot, 1825) nympha . Cerithium . Eichwald, 1851 → Tiaracerithium pictum (de Basterot, 1825) peneckei . Potamides . Hilber, 1891 → Tiaracerithium pictum (de Basterot, 1825) pentastichus . Cerithium . Quenstedt, 1884 → Tiaracerithium pictum (de Basterot, 1825) pictum . Cerithium . de Basterot, 1825 → Tiaracerithium pictum (de Basterot, 1825) pseudogamlitzensis. Pirenella . Strausz, 1955a → Tiaracerithium pictum (de Basterot, 1825) pulchellum . Cerithium . Sowerby, 1832 → Tiaracerithium pictum (de Basterot, 1825) rumanum . Cerithium . Pilide, 1878 → Tiaracerithium pictum (de Basterot, 1825) soceni . Pirenella . Jekelius, 1944 → Tiaracerithium pictum (de Basterot, 1825) stefanescui. Cerithium . Fontannes, 1887 → Tiaracerithium pictum (de Basterot, 1825) striata. Cerithium . Friedberg, 1905 → Tiaracerithium pictum (de Basterot, 1825) tetrastichus . Cerithium . Quenstedt, 1884 → Tiaracerithium pictum (de Basterot, 1825) thiara . Cerithium . auctores → Tiaracerithium pictum (de Basterot, 1825) trijugum . Cerithium . Eichwald, 1851 → Tiaracerithium pictum (de Basterot, 1825) trizonata. Cerithium . Eichwald, 1851 → Tiaracerithium pictum (de Basterot, 1825) turritella . Cerithium . [ Deshayes, 1835] → Tiaracerithium pictum (de Basterot, 1825) unicostata. Potamides . Friedberg, 1914 → Tiaracerithium pictum (de Basterot, 1825) varicosum . Cerithium . Handmann, 1883a → Tiaracerithium pictum (de Basterot, 1825)