Melongena cornuta ( Agassiz, 1843 )

Harzhauser, Mathias & Landau, Bernard M., 2024, The Colubrariidae, Eosiphonidae, Melongenidae, Pisaniidae, Prodotiidae and Tudiclidae (Gastropoda, Buccinoidea) of the Miocene Paratethys Sea, Zootaxa 5427 (1), pp. 1-110 : 16-19

Melongena cornuta ( Agassiz, 1843)

Figs 8A–D

P [yrula]. Melongena Lam. — Grateloup 1845: pl. 26, figs 1, 7, pl. 28, figs 12, 15 [non Melongena melongena ( Linnaeus, 1758) ].

* P [yrula]. Cornuta — Agassiz 1843: 89.

[ Pyrula ] melongena Lam — Hörnes 1848: 19 [non Melongena melongena ( Linnaeus, 1758) ].

Pyrula cornuta Ag. —Hörnes 1853: 274, pl. 29, figs 1–3, pl. 30, figs 1–3.

Pyrula cornuta Agassiz — Neugeboren 1854: 155.

Pyrula Melongenoides — Millet 1865: 591.

Pyrula denudata — Millet 1865: 582.

Myristica cornuta (Ag.) — Bellardi 1873: 157.

Pyrula melongena — Quenstedt 1884: 615, pl. 209, figs 65–67.

Melongena cornuta Ag. — Dollfus 1888: 31, pl. 1, figs 1–2.

[ Melongena cornuta ] var. semispinosa G.D. — Dollfus 1888: 31, pl. 1, fig. 3.

[ Melongena cornuta ] var. minor G.D. — Dollfus 1888: 56, pl. 1, fig. 4 [non G.B. Sowerby III, 1879].

[ Melongena cornuta ] var. bispinosa G.D. — Dollfus 1888: 56, pl. 1, fig. 5 [non Melongena bispinosa (Philippi, 1844) ].

[ Melongena cornuta ] var. patuloidea G.D. — Dollfus 1888: 56, pl. 3.

Pyrula (Melongena) cornuta Ag. —Hoernes & Auinger 1890: 247, pl. 28, figs 14–16.

P [yrula]. Cornuta — Cossmann 1901: 86, pl. 5, fig. 11.

Melongena cornuta (Ag.) — Sacco 1904: 32, pl. 9, figs 18–21.

Melongena cornuta var. tudicloides (Myl.) — Sacco 1904: 32, pl. 9, fig. 22.

Pyrula (Melongena) cornuta Ag. var. Gauderndorfensis Schff.— Schaffer 1912: 140, pl. 49, fig. 29.

Melongena cornuta Agassiz — Peyrot 1928: 37, pl. 8, figs 3–6.

Melongena cornuta Agas. Var. semispinosa Dollf.— Peyrot 1928: 39, pl. 8, fig. 1.

Melongena cornuta Agas. Var. bispinosa Dollf. — Peyrot 1928: 39, pl. 8, fig. 2 [non Melongena bispinosa (Philippi, 1844) ].

Melongena cornuta (Agassiz) — Rutsch 1929: 23, pl. 1, fig. 11.

Melongena (Pyrula) cornuta (Agassiz) — Stchepinsky 1939: 38, pl. 10, fig. 30.

Pyrula (Melongena) cornuta Lk. Var. pseudobasilica nov. var. — Strausz 1943: 137, pl. 1, figs 1, 3, 6–8.

Galeodes (Volema) cornuta Agassiz — Csepreghy-Meznerics 1950: 50. Melongena (Volema) cornuta hungarica n. sp. — Csepreghy-Meznerics 1950: 51, pl. 3, fig. 3.

Galeodes (Volema) cornuta Lk. — Csepreghy-Meznerics 1954: 41.

Galeodes (Volema) cornuta Ag. — Strausz 1954: 27, pl. 7, figs 154a–b.

Galeodes (Volema) cornuta palatina nov. var. — Strausz 1954: 27, pl. 8, figs 155a–b.

Galeodes (Volema) cornuta Ag. — Strausz 1954: 207, pl. 12, figs 12–16.

Galeodes (Galeodes) cornutus Agass. — Korobkov 1955: plate captions, pl. 99, fig. 8.

Galeodes cornuta ( Agassiz, 1843) — Hölzl 1958: 232, pl. 20, fig. 10.

Galeodes cornuta semispinosa ( Dollfus, 1888) — Hölzl 1958: 233.

G [aleodes]. (G [aleodes].) cornutus (Ag.) — Sieber 1958: 151.

Galeodes (Galeodes) cornutus ( Agassiz 1843) —Kojumdgieva in Kojumdgieva & Strachimirov 1960: 192, pl. 47, figs 1a–b.

Melongena (s. s.) cornuta Agassiz , sp. 1843— Glibert 1963: 86.

Melongena cornuta Agassiz, 1843 — Strausz 1966a: 298, pl. 59, figs 2–3, pls. 60–62, pl. 63, figs 1–13.

Melongena cornuta pseudobasilica Strausz, 1943 — Strausz 1966a: 299, pl. 57, figs 10–12, pl. 58, pl. 59, figs 1–2.

Melongena cornuta palatina Strausz, 1954 — Strausz 1966a: 300, pl. 57, figs 7–9.

Melongena cornuta Agassiz — Florei & CriŞan 1967: 207, pl. 3, figs 7a–b.

Galeodes (Galeodes) cornutus ( Agassiz, 1843) — Steininger et al. 1971: 400.

Galeodes (G [aleodes].) cornutus (Agassiz) — Stancu et al. 1971: 125, pl. 7, figs 4–5 [non fig. 6 = Strombidae ].

Galeodes (Volema) cornuta Ag. juv.— Csepreghy-Meznerics 1972: 29, pl. 11, figs 23, 27.

Galeodes (Galeodes) cornutus ( Agassiz, 1843) — Steininger 1973: 421.

Galeodes (Volema) cornuta (Agassiz) — Bohn-Havas 1973: 1114, pl. 5, fig. 9.

Melongena cornuta pseudobasilica Strausz, 1943 — Nikolov 1994: 55, pl. 5, figs 3–4.

Melongena cornuta ( Agassiz, 1843) — Švagrovský 1982: 388, pl. 2, fig. 3.

Galeodes cornutus ( Agassiz, 1843) — Bałuk 1995: 249, pl. 36, figs 7–8.

Galeodes cornutus (Agassiz) — Schultz 1998: 68, pl. 27, unnumbered fig.

Melongena cornuta ( Agassiz, 1843) — Lozouet et al. 2001: 62, pl. 27, figs 1–2.

Melongena cornuta ( Agassiz, 1843) —Harzhauser 2002: 102, pl. 7, figs 1–3.

Melongena cornuta ( Agassiz, 1843) — Harzhauser 2003: 200, pl. 1, fig. 8.

Galeodes (Volema) cornuta hungarica Csepreghy-Meznerics, 1950 — Pálfy et al. 2008: 106.

Euthria (Euthria) puschi ( Andrzejowski, 1830) — Mikuž 2009: 24, pl. 7, fig. 89 [non Euthria puschii ( Andrzejowski, 1830) ].

Melongena cornuta ( Agassiz, 1843) — Kovács 2022: 87, figs 72–73.

Melongena cornuta ( Agassiz, 1843) — Kovács & Vicián 2023: 238.

Non Galeodes cornuta (Agassiz) — Erünal-Erentöz 1958: 60, pl. 10, figs 1–2 [= Melongena jaapi Landau, Harzhauser, İslamoğlu & Silva, 2013 ].

Type material. Not defined. Agassiz (1843: 89) mentioned occurrences from the region around Bordeaux (Burdigalian) without referring to specific specimens. From the text alone, it is not even clear if he had specimens at hand. Specimens described by de Basterot (1825: 68) from the Burdigalian of Saint-Paul-lès-Dax ( France) are listed as syntypes in the collection of the Muséum national d’Histoire naturelle (Paris) ( MNHN.F.A70864), but this is incorrect, because Agassiz (1843) did not refer to de Basterot (1825).

Illustrated material. NHMW 1858/0015/0124, SL: 106.1 mm, MD: 84.0 mm, Mikulov ( Czech Republic), Figs 8A 1 –A 2 . NHMW 1851/0026/0065, SL: 79.2 mm, MD: 58.7 mm, Niederkreuzstetten ( Austria), illustrated in Hoernes & Auinger 1890: pl. 28, fig. 14), Figs Figs 8B 1 –B 2 . NHMW 1864/0001/0507, SL: 175 mm, MD: 139 mm, Niederkreuzstetten ( Austria), Figs 8C 1 –C 2 . NHMW 1848/0003/0071, Niederkreuzstetten ( Austria), SL: 58.1 mm, MD: 44.8 mm, illustrated in Hörnes (1853: pl. 30, fig. 2), Figs 8D 1 –D 2 .

Additional material. Karpatian (Early Miocene): NHMW 1851/0026/0065, Niederkreuzstetten ( Austria), SL: 197 mm, MD: 145 mm, illustrated in Hörnes (1853: pl. 30, fig. 1); 1 spec., NHMW 1861 /0050/0106, Niederkreuzstetten ( Austria) ; 1 spec., NHMW 1849 /0004/0018, Niederkreuzstetten ( Austria) ; 1 spec., NHMW 1864 /0001/0506, Niederkreuzstetten ( Austria) ; 4 spec., NHMW 1849 /0004/0019, Niederkreuzstetten ( Austria) ; 1 spec., NHMW 1859 /0010/0015, Rückersdorf ( Austria) ; 1 spec., NHMW 1851 /0026/0651, Weinsteig ( Austria) ; 5 spec., NHMW 1846 /0037/0238, Weinsteig ( Austria) ; 2 spec., NHMW 1864 /0001/0537, Grossrussbach ( Austria) . Badenian (Middle Miocene): 1 spec., NHMW 1862 /0001/0321, Baden ( Austria) ; 1 spec., NHMW 1866 /0001/0679, Baden ( Austria) ; 8 spec., NHMW 1851 /0026/0032, Grund ( Austria) ; 1 spec., NHMW 1855 /0045/0712, Grund ( Austria) ; 1 spec., NHMW 1868 /0001/0365, Gamlitz ( Austria) ; 2 spec., NHMW 1868 /0001/0366, Gamlitz ( Austria) ; 1 spec., NHMW 1870 /0038/0019, Ritzing ( Austria) ; 2 spec., NHMW 1864 /0001/0538, Hrušovany nad Jevišovkou ( Czech Republic) ; 1 spec., NHMW 1867 /0019/0109, CoŞteiu de Sus ( Romania) ; 2 spec., NHMW 1868 /0001/0457, Lăpugiu de Sus ( Romania) .

Revised description. Large to very large, very robust, broad shell of up to six whorls. Apical angle ~70–80°, rapidly increasing during ontogeny. Protoconch unknown. First two or three teleoconch whorls strongly coronate at shoulder placed below mid-whorl. Surface smooth except for delicate growth lines or delicate spiral threads on very broad, weakly concave subsutural ramp. Subsequent whorls forming prominent subsutural collar covering shoulder of preceding whorl, often resulting in suture strongly undulating around shoulder tubercles. Last whorl high, attaining up to 90% of total height, with very broad, weakly concave subsutural ramp, usually delimited by strong shoulder, rarely with convex periphery, bearing large, widely spaced, often pointed tubercles of very variable morphology, conical mid-whorl, base weakly concave; sculpture variable, ranging from smooth, except for growth lines, to distinct spiral sculpture of broad, often slightly undulating cords over base; oblique spiral row of smaller pointed tubercles delimiting base; fasciole prominent, broad, bearing strongly developed lamellar growth lines. Aperture wide, ovate. Columella slightly twisted, straight or weakly excavated in adapical half. Columellar callus very broad, relatively thin, well delimited from base and fasciole. Fasciole and abapical part of columella forming characteristic broad, flattened area. Anal canal distinctly incised. Outer lip rarely preserved, moderately thickened, thinning towards edge, smooth within. Siphonal canal moderately short, very wide.

Paratethyan synonyms. Several varieties and subspecies names have been established for French and Italian specimens based on variations in size and sculpture (e.g., Melongena cornuta tudicloides (Mylius, 1891) ; Melongena cornuta semispinosa Dollfus, 1888 ; Melongena cornuta minor Dollfus, 1888 [non G.B. Sowerby III, 1879]; Melongena cornuta bispinosa Dollfus, 1888 [non R.A. Philippi, 1844)]; Melongena cornuta patuloidea Dollfus 1888 ( Dollfus 1888; Sacco 1904). Four names have been established for Paratethyan occurrences: Melongena cornuta gauderndorfensis ( Schaffer 1912) , from the Eggenburgian (Burdigalian) of Gauderndorf ( Austria), represents specimens with completely reduced nodes at the convex periphery. Melongena cornuta pseudobasilica Strausz, 1943 , from the Badenian (Middle Miocene) of Pécsvárad ( Hungary), is based on dwarfed specimens. Melongena cornuta hungarica Csepreghy-Meznerics, 1950 , from the Badenian (Middle Miocene) of Hidas ( Hungary), is a subadult specimen with relatively prominent spiral sculpture on the subsutural ramp. Galeodes cornuta palatina Strausz, 1954 , from the Badenian (Middle Miocene) of Várpalota ( Hungary), represents juvenile shells with coronate spire whorls and prominent spiral cords on the youngest preserved whorls.

Discussion. This species is enormously variable concerning size, shape and sculpture. (note the considerable size difference between Figs 8C and 8D !). Comparable variability is observed in populations of the extant western Atlantic Melongena melongena ( Linnaeus, 1758) and Melongena corona Gmelin, 1791 ( Bruggeman-Nannenga & Hummelinck 1986; Karl & Hayes 2023). Genetic data of Melongena corona did not support the various morphosubspecies described in the literature but documented a single , albeit clearly geographically genetically structured, species (Karl & Hayes 2023). Therefore, we assume that the various ‘subspecies’ and ‘varieties’ of Melongena cornuta result from a comparable intraspecific plasticity.

The largest Paratethyan specimens, attaining more than 20 cm in height, are found in the Karpatian of the Korneuburg Basin and Badenbian of Hungary. Comparable gigantic specimens are found in the Langhian of the northeastern Atlantic of the Touraine ( France). Late Badenian (Serravallian) specimens of the Paratethys Sea are generally small and have reduced sculpture. Therefore, the huge size might correlate with the duration of the Miocene Climatic Optimum. Nevertheless, at Niederkreuzstetten ( Austria), from where several gigantic specimens derive, small and dwarfed specimens co-occur in large numbers.

In the Paratethys, Melongena cornuta was most common during the Karpatian (late Burdigalian), when Avicennia mangroves flourished ( Harzhauser et al. 2002). The successive decline of mangroves during the late Langhian and their loss during the Serravallian, followed the cooling at the Miocene Climate Transition ( Shevenell et al. 2004; Westerhold et al. 2020) and was also reflected by a decline of potamidid diversity ( Harzhauser et al. 2023a, b). Interestingly, the probably mangrove-associated potamidid Ptychopotamides cinctus (Bruguière, 1792) displays the same pattern of abundance and occurrence as Melongena cornuta , which is an additional hint that both species preferred the same habitat.

Paleoenvironment. Coastal marine, intertidal, probably in Avicennia mangroves (Harzhauser 2002).

Distribution in Central Paratethys. Eggenburgian (Early Miocene): North Alpine Foreland Basin: St. Gallen ( Switzerland) ( Rutsch 1929); Gernergraben, Kaltenbachgraben ( Germany) ( Hölzl 1958); Gauderndorf ( Austria) ( Schaffer 1912); Pannonian Basin: Budafok-Budapest ( Hungary)( Steininger et al. 1971); Dubova Region ( Romania) ( Stancu et al. 1971). Karpatian (Early Miocene): Korneuburg Basin: Grossrussbach, Weinsteig, Niederkreuzstetten, Stetten, Teiritzberg ( Austria) (Harzhauser 2002, hoc opus). Badenian (Middle Miocene): Korytnica Basin: Korytnica ( Poland) ( Bałuk 1995); Vienna Basin: Borský Mikuláš ( Czech Republic) ( Švagrovský 1982); Oberpullendorf Basin: Ritzing ( Austria) (hoc opus); Pannonian Basin: Borsod, Hidas, Letkés, Pécsvárad, Sámsonháza, Várpalota ( Csepreghy-Meznerics 1954; Kovács & Vicián 2023); Krško Basin: Dolenja Brezovica, Gorenje Vrhpolje ( Slovenia) ( Mikuž 2009); Banat: Zorlenţul Mare ( Romania) ( Florei & CriŞan 1967); Făget Basin: CoŞteiu de Sus, Lăpugiu de Sus ( Romania) ( Kovács 2022; hoc opus); Dacian Basin: Opanec ( Bulgaria), Dobrusha ( Bulgaria) (Kojumdgieva in Kojumdgieva & Strachimirov 1960; Nikolov 1994).

Proto-Mediterranean Sea. Burdigalian (Early Miocene): Colli Torinesi: Rio della Batteria, Termô-Fôurà, Baldissero ( Italy) ( Bellardi 1873); Mesohellenic Basin: Argos Orestikon, Damaskinea, Odria ( Greece) (own data, M.H.); Sivas Basin: İŞhan ( Turkey) ( Stchepinsky 1939).

Northeastern Atlantic. Aquitanian and Burdigalian (Early Miocene): Aquitaine Basin: Cestas, Léognan, Mérignac, Saint-Paul-lès-Dax, Saucats ( France) ( Peyrot 1928); Langhian and Serravallian (Middle Miocene): Aquitaine Basin: Salles, Salies-de-Béarn, Saubrigues ( France) ( Peyrot 1928); Touraine ( France) ( Dollfus 1888).

Figures

FIGURE 1. Measurements for shell morphology; AA: apical angle, SL: shell length, MD: maximum diameter, LWH: last whorl height, AH: aperture height, AL: aperture length, AW: aperture width.

FIGURE 2. Colubraria subobscura (Hoernes & Auinger, 1884). A1–A3. NHMW 1854/0035/0170, paralectotype, Lăpugiu de Sus (Romania). B1–B2. NHMW 1859/0037/0018a, paralectotype, Lăpugiu de Sus (Romania). C1–C3. NHMW 1865/0001/0183, paralectotype, Lăpugiu de Sus (Romania). D1–D2. NHMW 1859/0037/0018b, paralectotype, Lăpugiu de Sus (Romania). E1–E3. NHMW 1874/0025/0021, lectotype, Lăpugiu de Sus (Romania). F1–F2. NHMW 2023/0338/0001, paralectotype, Lăpugiu de Sus (Romania).

FIGURE 8. A. Melongena cornuta (Agassiz, 1843). A1–A2. NHMW 1858/0015/0124, Mikulov (Czech Republic). B1–B2. NHMW 1851/0026/0065, Gauderndorf (Austria). C1–C2. NHMW 1864/0001/0507, Niederkreuzstetten (Austria). D1–D2. NHMW 1848/0003/0071, Niederkreuzstetten (Austria).