Theodisca theodisca ( Rolle, 1856 )

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 : 89-94

Theodisca theodisca ( Rolle, 1856)

Figs 14A–L

* Cerithium Theodiscum Rolle — Rolle 1856: 573.

Cerithium Gamlitzense Hilb. — Hilber 1879a: 437, pl. 4, figs 2–3.

Cerithium Rollei Hilb. — Hilber 1879a: 24, pl. 4, figs 4a–c.

Cerithium theodiscum Rolle — Hilber 1879a: 439, pl. 4, figs 5a–c.

Cerithium biquadratum Hilb. — Hilber 1879a: 441, pl. 4, figs 6a–c.

Cerithium Noricum Hilb. — Hilber 1879a: 442, pl. 4, figs 7a–c.

Cer [ithium]. theodiscum — Hilber 1879b: 124.

Cerithium Schaueri Hilber nova forma — Hilber 1882: 7, pl. 1, fig. 14 [non Hilber, 1882].

Potamides (Pirenella) Gamlitzensis Hilber — Hilber 1891: 239.

Cerithium Gamlitzense Hilb. — Bauer 1900: 30.

Cerithium Rollei Hilb. — Friedberg 1905: 27, pl. 1, figs 4a–d.

Pirenella Schaueri Hilb. — Friedberg 1914: 284, pl. 17, fig. 19 [non Hilber, 1882].

Pirenella gamlitzensis gamlitzensis (Hilber) — Papp 1952: 114, pl. 3, figs 1–5.

Pirenella gamlitzensis rollei (Hilber) — Papp 1952: 115, pl. 3, figs 6–9.

Pirenella gamlitzensis theodisca (Rolle) — Papp 1952: 115, pl. 3, fig. 10.

Potamides (Pirenella) gamlitzensis Hilb. — Strausz 1954: 17, pl. 2, fig. 27.

Pirenella nodosoplicata biquadrata Hilber, 1879 — Strausz 1955a: 64, pl. 4, fig. 40.

Pirenella gamlitzensis theodisca Rolle (in Hilber), 1879— Strausz 1955a: 69, 192, pl. 7, fig. 117.

Pirenella gamlitzensis Hilber, 1879 — Strausz 1955a: 67, 90, pl. 7, figs 99–105, 111, pl. 8, figs 124–126, 128–134, text-fig. 4. Pirenella gamlitzensis rollei Hilber, 1879 — Strausz 1955a: 68, 191, pl. 7, figs 107, 118, pl. 8, fig. 127.

Pirenella gamlitzensis pseudotheodisca nov. f.— Strausz 1955a: 69, 192, pl. 7, figs 119–123.

P [otamides]. gamlitzensis gamlitzensis (Hilb.) — Sieber 1958: 136.

P [otamides]. gamlitzensis theodisca (Rolle) — Sieber 1958: 136.

P [otamides]. gamlitzensis rollei (Hilb.) — Sieber 1958: 136.

Pirenella picta biseriata (Friedberg) — Švagrovský 1959: 222, pl. 6, fig. 14 [non Friedberg, 1914].

Potamides (Pirenella) gamlitzensis transdanubicus nov. var. — Strausz 1959: 149, pl. 2, figs 5–6.

Potamides (Pirenella) gamlitzensis Hilber — Strausz 1966: 153, pl. 7, figs 43–46, pl. 8, figs 1–4.

Pirenella gamlitzensis theodisca Rolle (in Hilber), 1879— Strausz 1966: 154, pl. 7, figs 14–15.

Potamides (Pirenella) nodosoplicata biquadratus Hilber, 1879 — Strausz 1966: 153, pl. 7, fig. 22.

Potamides (Pirenella) gamlitzensis rollei Hilber, 1879 — Strausz 1966: 154, pl. 7, figs 38–40.

Pirenella gamlitzensis pseudopicta nov. ssp. — Kókay 1966: 42, pl. 4, fig. 7.

Potamides (Pirenella) gamlitzensis pseudotheodiscus Strausz, 1955 — Strausz 1966: 155, pl. 7, figs 16–18.

Potamides (Pirenella) gamlitzensis transdanubicus Strausz, 1959 — Strausz 1966: 156, pl. 7, figs 41–42.

Pirenella gamlitzensis gamlitzensis (Hilber, 1879) —Hinculov in Iliescu et al. 1968: 125, pl. 29, figs 11–12.

Pirenella gamlitzensis theodisca (Rolle in Hilber, 1879)—Hinculov in Iliescu et al. 1968: 126, pl. 30, figs 1–2.

Potmides (Pirenella) biseriatus Friedberg — Bałuk 1970: 117, pl. 10, fig. 5 [non Friedberg, 1914].

Pirenella gamlitzensis (Hilber, 1879) — Bałuk 1975: 132, pl. 15, fig. 7.

? Pirenella gamlitzensis (Hilber, 1879) — Calzada-Badia et al. 1978: 114, pl. 1, fig. 6.

Pirenella gamlitzensis (Hilber, 1879) — Švagrovský 1982: 22, pl. 6, fig. 2.

Pirenella mitralis — Aguilà & García García 1997: 20, pl. 1, fig. 8 [non Tiarapirenella bicincta ( Brocchi, 1814) ].

? Pirenella gamlitzensis gamlitzensis (Hilber, 1879) — Ýslamoðlu & Taner 2003: 38, pl. 1, figs 1, 13.

Potamides theodiscus ( Rolle 1856) — D’Amico et al. 2012: 160, pl. 1, fig. 5.

Potamides nodosoplicatus ( Ĥrnes, 1855) — Landau et al. 2013: 54, pl. 4, figs 13–14 [non Ĥrnes, 1855].

non Potamides petersi Auing. (an biquadrata Hilb. )— Friedberg 1914: plate captions, pl. 17, fig. 21 [= Pustulosia submitralis

( Eichwald, 1851)]. non Pirenella nodosoplicata biquadrata Hilber — Boda 1959: 618, pl. 26, fig. 14 [= Pustulosia submitralis ( Eichwald, 1851) ]. non Pirenella gamlitzensis gamlitzensis (Hilber) — Boda 1959: 616, pl. 25, figs 8–11 [= Theodisca biseriata ( Friedberg,

1914)]. non Pirenella biquadrata (Hilber) — Mariani & Papp 1966: 143, pl. 1, figs 6–7 [= Granulolabium submitrale ( Eichwald,

1851)]. non Potamides (Pirenella) gamlitzensis (Hilber, 1879) — Iljina 1993: 73, pl. 9, figs 10–14 [= Theodisca biseriata ( Friedberg,

1914)]. non Potamides (Pirenella) gamlitzensis (Hilb.) — Goncharova & Iljina 1997: 69, unnumbered plate, fig. 30 [? = Potamides

(Pirenella) cicur ( Zhizhchenko, 1934)].

Type material. Lectotype designated herein: NHMW 1846/0049/0019, SL: 14.8 mm, MD: 4.8 mm, St. Florian ( Austria), Badenian, Middle Miocene, illustrated in Hilber (1879a: pl. 4, figs 5a–c), Figs 14A 1 –A 2 . Paralectotypes: NHMW 1846/0049/0019a, SL: 11.5 mm, MD: 3.9 mm, St. Florian ( Austria), Badenian , Middle Miocene , Figs 14B 1 –B 2; NHMW 1846/0049/0019b, SL: 11.4 mm, MD: 4.2 mm, St. Florian ( Austria), Badenian, Middle Miocene, Figs 14C 1 –C 2 .

Type locality. St. Florian ( Austria).

Stratigraphy. Middle Miocene, Badenian.

Illustrated material. NHMW 2022/0127/0001, SL: 13.0 mm, MD: 4.5 mm, Gamlitz ( Austria), Badenian, Middle Miocene, Figs 14D 1 –D 2. NHMW 1860/0005/0050a, SL: 13.3 mm, MD: 4.2 mm, St. Florian ( Austria), Badenian, Middle Miocene, Figs 14E 1 –E 2. M.269, SL: 10.1 mm, MD: 3.0 mm, holotype of Pirenella gamlitzensis pseudopicta Kókay, 1966 , Herend ( Hungary), Badenian, Middle Miocene, Fig. 14F. UMJG &P 200.147, SL: 12.2 mm, MD: 5.1 mm, Gamlitz ( Austria), Universalmuseum Joanneum (Graz, Austria), illustrated in Hilber (1879a: pl. 4, figs 6a–c), holotype of Cerithium biquadratum Hilber, 1879 , Fig. 14G. UMJG &P 200.129, SL: 13.3 mm, MD: 4.6 mm, Gamlitz ( Austria), Universalmuseum Joanneum (Graz, Austria), illustrated in Hilber (1879a: pl. 4, figs 3a–c), paralectotype of Cerithium gamlitzense Hilber, 1879 , Figs 14H 1 –H 2. UMJG &P 200.149, SL: 15.9 mm, MD: 6.2 mm, Gamlitz ( Austria), Universalmuseum Joanneum (Graz, Austria), illustrated in Hilber (1879a: pl. 4, figs 2ac), lectotype (designated by Švagrovský 1982) of Cerithium gamlitzense Hilber, 1879 , Fig. 14I. M. 2 SL: 15.8 mm, MD: 6.0 mm, Várpalota ( Hungary), Badenian, Middle Miocene, holotype of Pirenella gamlitzensis transdanubicus Strausz, 1959 , Fig. 14J. UMJG &P 200.144, SL: 13.6 mm, MD: 5.3 mm, Universalmuseum Joanneum (Graz, Austria), Gamlitz ( Austria), illustrated in Hilber (1879a: pl. 4, figs 7a–c), lectotype of Cerithium noricum Hilber, 1879 , Fig. 14K. NHMW 1851/0021/1961a, SL: 4.2 mm, MD: 2.1 mm, Vienna /Mauer, Badenian (Middle Miocene), Fig. 14L.

Material studied. 46 spec., 1860/0005/0050, St. Florian ( Austria), Badenian, Middle Miocene ; 14 spec., NHMW 1856/0020/0020, St. Florian ( Austria), Badenian, Middle Miocene ; 62 spec., NHMW 1861/0035/0132, Steinebrunn ( Austria), Badenian, Middle Miocene .

Revised description. Small sized, elongate conical to elongate pupoid shell of 10 teleoconch whorls; apical angle ~ 30–35°. Protoconch turreted of three convex whorls. Early teleoconch whorls convex with weakly convex subsutural ramp and two prominent, beaded spiral cords (in mid-whorl position and at some distance from abapical suture); third spiral cord intercalated on subsutural ramp on subsequent whorls. Later teleoconch whorls conical to subcylindrical; typically, with three prominent, beaded spiral cords. Central spiral cord often reduced to broad spiral band between adsutural spiral cords. Suture incised, undulating slightly around large, rounded adapical row of beads. Last whorl subcylindrical with moderately convex base with three to five broad spiral cords. Two peribasal cords present. Narrow, twisted fasciole. Aperture ovoid, moderately wide. Inner lip forming broad rim with narrow pseudumbilical chink. Columella deeply excavated, smooth. Anal canal weakly incised. Outer lip weakly thickened, convex, not flaring. Siphonal canal moderately narrow, short, slightly deflected to the left.

Synonyms. Hilber (1879a) described four species, which all derived from the Badenian of Gamlitz ( Austria) in the Styrian Basin. These species are treated herein as subjective junior synonyms of Theodisca theodisca : Cerithium gamlitzensis was based on two syntypes of which the specimen illustrated in Hilber (1879a: pl. 4) as fig. 2 was designated by Švagrovský 1982 as lectotype. This specimen has a bicinctum -like morphology and at first sight one might place the specimen in Granulolabium . The central spiral cord, which is typical for Theodisca theodisca is completely reduced on most of the teleoconch. The early teleoconch whorls, however, develop three spiral rows of beads, atypical for Granulolabium . The paralectotype (fig. 3) has also a tripartite early teleoconch sculpture and does not differ from Theodisca theodisca . Cerithium noricum Hilber, 1879 is a comparatively stout morph of T. theodisca with strongly reduced central spiral cord. The early teleoconch, however, shows the typical tripartite sculpture. Cerithium rollei Hilber, 1879 , does not differ from other specimens of T. theodisca . The fourth species from Gamlitz is Cerithium biquadratum and one might hesitate to synonymize it with T. theodisca . Cerithium biquadratum is known from a single specimen, co-occurring with numerous shells of T. theodisca . We consider it to be an extreme, unusually stout morph, which lacks any central spiral cord and develops blunt beads. Potamides (Pirenella) gamlitzensis transdanubicus Strausz, 1959 has a reduced central spiral cord and prominent beads on the adsutural spiral cord. It was described by Strausz (1959) from Várpalota ( Hungary), where it co-occurs with various other morphologies of T. theodisca . Pirenella gamlitzensis pseudotheodisca Strausz, 1955a represents a stout morph of T. theodisca with strongly reduced central spiral cord and blunt nodes on the abapical spiral cord. Pirenella gamlitzensis pseudopicta Kókay, 1966 is a slender morph with prominent spiral cords..

Discussion. For data on ecology and separation from Theodisca biseriata ( Friedberg, 1914) see T. biseriata .

Distribution. Theodisca theodisca appeared in the Central Paratethys during the Late Burdigalian (Karpatian) and might have been distributed at that time also in the Proto-Mediterranean Sea. It was widespread during the Badenian (Langhian, Serravallian) but vanished with the onset of the Sarmatian. In the Proto-Mediterranean Sea, it persisted from the Serravallian to the Tortonian.

Central Paratethys. Karpatian (Early Miocene): North Alpine-Carpathian Foredeep: Laa an der Thaya ( Austria) ( Hilber 1879a); Badenian (Middle Miocene): Korytnica Basin: Korytnica ( Poland) ( Bałuk 1975); Polish-Ukrainian Fore-Carpathian Basin: Sobów, Miechocin ( Poland) ( Friedberg 1905); Vienna Basin: Vienna /Mauer ( Austria) (hoc opus), Borský Mikuláš ( Slovakia) ( Švagrovský 1982); Danube Basin: Kuzmice ( Slovakia) ( Švagrovský 1960); Styrian Basin: Gamlitz, Jamm, St. Florian, Windisch-Graz ( Austria) ( Hilber 1879a; Bauer 1900); Pannonian Basin: Várpalota, Herend–Márkó ( Hungary) ( Strausz 1966); Mehadia Basin: Valea Bela Reca, Valea Calvei ( Romania) (Hinculov in Iliescu et al. 1968).

Proto-Mediterranean Sea.? Burdigalian (Early Miocene): Spain ( Calzada-Badia et al. 1978), Turkey ( Ýslamoðlu & Taner 2003); Langhian (Middle Miocene): Vallès-Penedès ( Spain) ( Aguilà & García García, 1997); Serravallian: Karaman Basin ( Turkey) ( Landau et al. 2013); Tortonian (Late Miocene): southern Italy ( D’Amico et al. 2012).

of Pirenella gamlitzensis pseudopicta Kókay, 1966 . G. UMJG&P 200.147, Gamlitz ( Austria), Middle Miocene, Badenian,

holotype of Cerithium biquadratum Hilber, 1879 . H 1 –H 2. Theodisca theodisca ( Rolle, 1856) , UMJG&P 200.129, Gamlitz ( Austria), Middle Miocene, Badenian, paralectotype of Cerithium gamlitzense Hilber, 1879 . I. Theodisca theodisca ( Rolle,

1856), UMJG & P 200.149 , Gamlitz ( Austria), Middle Miocene, Badenian, lectotype of Cerithium gamlitzense Hilber, 1879 . J .

M.2, Várpalota ( Hungary), Badenian, Middle Miocene, holotype of Pirenella gamlitzensis transdanubicus Strausz, 1959 . K .

Theodisca theodisca ( Rolle, 1856) , UMJG&P 200.144, Gamlitz ( Austria), Middle Miocene, Badenian, lectotype of Cerithium noricum Hilber, 1879 . L. Theodisca theodisca ( Rolle, 1856) , NHMW 1851/0021/1961a, Vienna /Mauer, Badenian, Middle

Miocene.

Genus Terebralia Swainson, 1840

Type species. Strombus palustris Linnaeus, 1767 ; subsequent designation by Sacco (1895: 51). Recent, Indo-West Pacific.

Original diagnosis. “ Shell moderately large, solid brown, with numerous flat-sided whorls, and sculptured with spiral cords and axial ribs; aperture ovate, outer lip sweeping in arc to join moderately-developed anterior canal frequently forming complete peristome. Columella with paired internal plaits; internal palatal teeth opposite varices.” ( Houbrick 1991: 304).

Stratigraphy and paleogeography. The oldest records of Terebralia go back to the middle Eocene of the northeastern Atlantic ( Thivaiou et al. 2023). During the Oligocene and Miocene, Terebralia was well represented in the entire Circum-Mediterranean region. It even reached the North Sea during the Early Miocene (Hemmoorian) ( Moths et al. 2010) and can be traced during the Burdigalian as far east as the Qom Basin in Iran (Harzhauser et al. 2002). Terebralia vanished in Europe during the Late Miocene and seems to have been absent from Messinian environments ( Thivaiou et al. 2023). The genus expanded its range into the Indo-West Pacific at least during the Burdigalian, when it is documented by several species in Pakistan and southern Iran ( Vredenburg 1925; Harzhauser et al. 2017). The closure of the Tethys Seaway during the middle Burdigalian seems to have been a vicariance event for Terebralia , resulting in morphologically very similar species at both sides of the closed gateway (Harzhauser et al. 2017). In the Indonesian region, however, Terebralia was still absent or rare during the Miocene, because all species listed by Leloux & Wesselingh (2009) as Terebralia seem to represent other genera. Herein we do not discuss the subsequent late Neogene rise of Terebralia in the IWP region.

Ecology. Modern Terebralia species are usually bound to mangroves, where species may attain significant densities of more than 100 specimens per m 2 ( Ellison et al. 1999; Wells & Lalli 2003). Population densities decline outside the mangrove pneumatophores on mudflats, which might be explained by a preference to leaf litter between the mangroves ( Crowe & McMahon 1997) and the increasing stress by predators on open mudflats ( Wells & Lalli 2003). Mangroves and mudflats seem to have been the preferred habitat of Terebralia also during the Paleogene and Neogene ( Reid et al. 2008; Thivaiou et al. 2023). Terebralia prolignitarum is found in assemblages with Mesohalina margaritacea and Ellobium , suggesting the presence of mangroves (see Mesohalina ). Similarly, Terebralia lignitarum from the Early Miocene (Karpatian) of the Korneuburg Basin co-occurred with Avicenna mangroves (Harzhauser et al. 2002; Kern et al. 2010) and stable isotope data of Latal et al. (2006) confirmed a marginal, oligohaline habitat. In the Paratethys, mangroves disappeared around 13.8 Ma with the onset of the Serravallian (late Badenian) ( Jiménez-Moreno et al. 2008) due to the global cooling of the Middle Miocene climate transition ( Holbourn et al. 2013). Nevertheless, Terebralia persisted in the Central Paratethys until the Middle/Late Miocene boundary (when Lake Pannon formed at 11.6 Ma) and survived in the Eastern Paratethys even into the Bessarabian (Late Miocene). This clearly documents that Sarmatian populations of Terebralia lignitarum and T. menestrieri were not strictly bound to mangroves.

Extant Terebralia species may co-occur within the same habitat, although Wells & Lalli (2003) discussed microhabitat partitioning between species. The Paleogene and Neogene European records suggest a similar diversity of up to two co-occurring species.

Discussion. Terebralia is a genus of low diversity, which is represented by only three species in the modern IWP-region: Terebralia palustris ( Linnaeus, 1767) , Terebralia sulcata ( Born, 1778) and Terebralia semistriata (M̂rch, 1852). In their review of the Cenozoic European occurrence of Terebralia, Thivaiou et al. (2023) observed a maximum of about four coeval species in the Western Tethys during the Oligocene and Early Miocene. In contrast, about 24 species level taxa of Terebralia have been described or mentioned from Paratethyan deposits. Two of these names ( abbreviata Schaffer, 1912 = schafferi Finlay, 1927 ) have been used for specimens of Granulolabium plicatum (Bruguière, 1792) , whereas all others represent Terebralia . This enormous diversity would be in strong contrast to the poor modern diversity of the genus. After revision of the taxa, we conclude that only five species occurred in Paratethyan environments: Terebralia prolignitarum ( Sacco, 1887) (Egerian) , T. lignitarum ( Eichwald, 1830) (Eggenburgian to Badenian), T. duboisi ( Ĥrnes, 1855) (Eggenburgian to Bessarabian), T. subcorrugatum (d’Orbigny, 1852) (early/middle Badenian) and T. menestrieri (d’Orbigny, 1844) (Volhynian to Bessarabian).

Species-level taxa placed in Terebralia reported from the Paratethys abbreviata . Clava . Schaffer, 1912 → Granulolabium plicatum (Bruguière, 1792) ampullosum . Cerithium . auctores → Terebralia lignitarum ( Eichwald, 1830)

angustata. Terebralia . Friedberg, 1936 → Terebralia lignitarum ( Eichwald, 1830) andrzejowskii . Cerithium . Friedberg, 1914 → Terebralia duboisi ( Ĥrnes, 1855)

bidentatum . Cerithium . Defrance, 1832 → Terebralia lignitarum ( Eichwald, 1830) bidentatum . Cerithium , auctores [pars] → Terebralia prolignitarum ( Sacco, 1887) cingulatior . Potamides . Sacco, 1888 → Terebralia lignitarum ( Eichwald, 1830) corrugatum . Cerithium . auctores → Terebralia sp.

dollfusi . Clava . Hoernes, 1901 → Terebralia subcorrugata (d’Orbigny, 1852) duboisi . Cerithium . Hörnes, 1855 → Terebralia duboisi ( Ĥrnes, 1855)

fusiformis. Cerithium (Clava) . Schaffer, 1912 → Terebralia lignitarum ( Eichwald, 1830)

holleri . Clava . Hoernes, 1901 → Terebralia lignitarum ( Eichwald, 1830) lignitarum . Cerithium . Eichwald, 1830 → Terebralia lignitarum ( Eichwald, 1830)

macoveii . Cerithium Simionescu & Barbu, 1940 → Terebralia menestrieri (d’Orbigny, 1844) margaritifera. Potamides (Terebralia) . Sacco, 1895 → Terebralia lignitarum ( Eichwald, 1830) mehadiensis. Terebralia . Hinculov et al., 1968 → Terebralia lignitarum ( Eichwald, 1830) menestrieri . Cerithium . d’Orbigny, 1844 → Terebralia menestrieri (d’Orbigny, 1844)

pauli . Cerithium . Hoernes, 1874 → Terebralia menestrieri (d’Orbigny, 1844) penetrabile . Cerithium . Kolesnikov, 1935 → Terebralia menestrieri (d’Orbigny, 1844) perrugata . Cerithium . Hilber, 1879a → Terebralia subcorrugata (d’Orbigny, 1852) praebidentata . Terebralia . Seneš in Steininger et al., 1971 → Terebralia lignitarum ( Eichwald, 1830) prolignitarum . Potamides . Sacco 1887 → Terebralia prolignitarum ( Sacco, 1887) quinquecinctum . Cerithium . Schaffer, 1912 → Terebralia duboisi ( Ĥrnes, 1855)

rahtii. Cerithium . auctores → Terebralia prolignitarum ( Sacco, 1887)

schafferi . Clava . Finlay, 1927 → Granulolabium plicatum (Bruguière, 1792) subcorrugatum . Cerithium . d’Orbigny, 1852 → Terebralia subcorrugata (d’Orbigny, 1852) vignali . Terebralia . Cossmann 1906 → Terebralia lignitarum ( Eichwald, 1830)