Hexaplex (Trunculariopsis) trunculoides (Pusch, 1837)

Harzhauser, Mathias, Landau, Bernard M. & Merle, Didier, 2025, The Muricidae (Gastropoda, Muricoidea) of the Miocene Central Paratethys Sea (Haustrinae, Muricinae, Ocenebrinae, Pagodulinae, Typhinae, Muricidae incertae sedis), Zootaxa 5572 (1), pp. 1-162 : 23-26

Hexaplex (Trunculariopsis) trunculoides (Pusch, 1837)

Figs 1 , 8A–D

* Murex trunculoides m.—Pusch 1837: 136, pl. 11, figs 23a–b.

[ Murex ] trunculus Lin. — Hauer 1837: 418 [non Hexaplex (Trunculariopsis) trunculus ( Linnaeus, 1758) ].

[ Murex ] trunculus Lin. — Hörnes 1848: 18 [non Hexaplex (Trunculariopsis) trunculus ( Linnaeus, 1758) ].

Mur [ex]. pomiformis m.— Eichwald 1853: 191 [non Murex pomiformis Mörch, 1852 , = Phyllonotus pomum ( Gmelin, 1791) ].

Mur [ex]. trunculoides Pusch — Eichwald 1853: 191.

Murex Sedgwicki Micht.—Hörnes 1853: 220, pl. 23, figs 1–5 [non Hexaplex (Trunculariopsis) sedgwicki ( Michelotti, 1847) ].

Murex Austriacus — Tournouër 1875: 158 [pro Murex sedgwicki sensu Hörnes 1853 : pl. 23, figs 1–5].

Murex amberus De Greg.—De Gregorio 1885a: 271 [pro Murex sedgwicki sensu Hörnes 1853: 220 , pl. 23, fig. 1].

Murex ( Phyllonotus ) Hoernesi d’Anc. —Hoernes & Auinger 1885: 211 [non Hexaplex (Trunculariopsis) hoernesi (D’Ancona, 1871) ].

Murex austriacus Tourn. — Friedberg 1912: 169, pl. 10, fig. 10.

Murex austriacus Tourn. — Friedberg 1938: 116.

Murex (Muricantha) sedgwicki vindobonensis sp. nov. — Csepreghy-Meznerics 1950: 46, pl. 2, fig. 17 [non Murex vindobonensis Hörnes, 1853 ].

Murex (Muricantha) hörnesi d’Ancona — Csepreghy-Meznerics 1950: 47, pl. 2, fig. 18 [non Murex hoernesi d’Ancona, 1871 , = Hexaplex (Trunculariopsis) campanii ( de Stefani & Pantanelli, 1878) ].

Murex (Muricantha) hoernesi d’Anc. — Strausz 1954: 25, pl. 5, figs 123.

Murex ( Hexaplex , sect. Phyllonotus ) austriacus Tourn. — Korobkov 1955: plate captions, pl. 70, figs 8a–8b.

Murex (Muricantha) sedgwicki vindobonensis Mezn. — Csepreghy-Meznerics 1956: 433 [non Murex vindobonensis Hörnes, 1853 (= Ocenebra vindobonensis (Hörnes, 1853) ].

H [exaplex]. (M [uricanthus].) hoernesi (Anc.) — Sieber 1958: 144 [non Hexaplex (Trunculariopsis) hoernesi (D’Ancona, 1871) ].

Murex (Muricanthus) pomiformis Eichwald 1853 —Kojumdgieva in Kojumdgieva & Strachimirov 1960: 146, pl. 40, fig. 1.

Murex (Muricanthus) turonensis pontileviensis Tournouër, 1875 — Strausz 1966: 264, pl. 32, figs 1, 4, pl. 33, fig. 2, pl. 56, figs 9–11, pl. 79, fig. 1 [non Murex turoniensis var. pontileviensis Tournouër, 1875 = Hexaplex (Trunculariopsis) turonensis ( Dujardin, 1837) ].

Murex (Muricanthus) rudis sedgwicki Michelotti, 1841 — Strausz 1966: 265, pl. 56, fig. 12 [non Hexaplex (Trunculariopsis) sedgwicki ( Michelotti, 1847) ].

Murex (Muricantha) gavardanensis Tourn. — Kókay 1966: 56, pl. 7, figs 16–17 [non Murex gavardanensis Benoist, 1880 , =? Hexaplex (Trunculariopsis) esplus (De Gregorio, 1885a) ].

Hexaplex austriacus (Tournouër) — Kókay 1966: 456, pl. 3, fig. 3.

[ Hexaplex ] austriacus Tournouër, 1875 — Vokes 1971: 20.

[ Hexaplex ] trunculoides Pusch, 1837 — Vokes 1971:111.

Murex (Muricanthus) pomiformis Eichwald — Nicorici 1972: 67.

Murex (Muricanthus) pomiformis Eichwald, 1853 — Nicorici & Sagatovici 1973: 171, pl. 25, figs 18–22.

Hexaplex austriacus ( Tournouer, 1875) — Stojaspal 1978: 336, pl. 4, fig. 4.

Trunculariopsis hörnesi (d’Ancona) — Martinell 1982: 375, pl. 1, figs 1–2 [non Murex hoernesi D’Ancona, 1871 = Hexaplex (Trunculariopsis) campanii ( de Stefani & Pantanelli, 1878) ].

Murex (Muricanthus) sedgwicki ( Michelotti, 1841) — Atanacković 1985: 138, pl. 31, figs 7–8 [non Hexaplex (Trunculariopsis) sedgwicki ( Michelotti, 1847) ].

Hexaplex (Muricanthus) bourgeoisi ( Tournouër, 1875) — Nikolov 1994: 48, pl. 1, figs 1–2 [non Hexaplex (Trunculariopsis) bourgeoisi ( Tournouër, 1875) ].

Hexaplex (Phyllonotus) pomiformis ( Eichwald, 1853) — Bałuk 1995: 214, pl. 21, figs 1–6.

Hexaplex austriacus (Tourn.) — Kókay 1996: 456, pl. 3, fig. 3.

Murex (Bolinus) subtorularius Hoernes & Auinger, 1885 —Harzhauser 2002: 94, pl. 6, figs 4, 5 [non Bolinus subtorularius (Hoernes & Auinger, 1885) ].

Hexaplex (Phyllonotus) pomiformis ( Eichwald 1853) —Harzhauser 2002: 94, pl. 6, figs 1–3.

Hexaplex (Phyllonotus) pomiformis ( Eichwald 1853) — Mikuž 2009: 26, pl. 8, figs 103–106.

Hexaplex (Trunculariopsis) austriacus ( Tournouër, 1875) — Merle et al. 2011: 88, 318, pl. 37, fig. 7.

Hexaplex (Trunculariopsis) austriacus (Tournouër) — Landau et al. 2013: 145, pl. 21, fig. 4.

Hexaplex (Trunculariopsis) austriacus ( Tournouër, 1875) — Kovács et al. 2018: 113, figs 2J–K.

Hexaplex (Trunculariopsis) austriacus ( Tournouër, 1875) — Kovács 2019: 114, fig. 4.

non Hexaplex (Trunculariopsis) cf. austriacus ( Tournouër, 1875) — Goret & Pons 2013: 56, pl. 1, fig. 3.

Type material. Holotype: specimen illustrated in Pusch (1837: pl. 11, fig 23), SL: 47 mm. MD: 25 mm, Pusch (1837) stated that his specimen comes from Volhynia (= Ukraine) without locality information. The specimen is lost.

Illustrated material. NHMW 1846/0037/0215, SL: 42.4 mm, MD: 28.3 mm, Gainfarn ( Austria), illustrated in Hörnes (1853: pl. 23, figs 4a–b), Figs 8A 1 –A 3 .

NHMW 1856/0050/0120, SL: 68.8 mm, MD: 47.4 mm, Gainfarn ( Austria), illustrated in Hörnes (1853: pl. 23, figs 2a–b), Figs 8B 1 –B 3 . NHMW 1855/0045/0428, SL: 92.6 mm, MD: 56.4 mm, Steinebrunn ( Austria), Figs 8C 1 –C 3 . NHMW 1856/0050/0455, SL: 89.1 mm, MD: 58.5 mm, Lăpugiu de Sus ( Romania), Figs 8D 1 –D 3 .

Additional material. NHMW 1860 /0001/0188, SL: 41.0 mm, MD: 25.9 mm, Steinebrunn ( Austria), illustrated in Hörnes (1853: pl. 23, figs 3a–b) ; NHMW 1846 /0037/0216, SL: 26.4 mm, MD: 17.7 mm, Gainfarn ( Austria), illustrated in Hörnes (1853: pl. 23, figs 5a–b); 1 spec. , NHMW 1859 /0010/0013, Rückersdorf ( Austria); 3 spec. , NHMW 1846 /0037/0216, Enzesfeld and Gainfarn ( Austria); 1 spec. , NHMW 1855 /0045/0428, Gainfarn ( Austria); 1 spec. , NHMW 1855 /0045/0429, Gainfarn ( Austria); 2 spec. , NHMW 1855 /0003/0027, Gainfarn ( Austria); 2 spec. , NHMW 1856 /0050/0120, Gainfarn ( Austria); 3 spec. , NHMW 1869 /0001/0350, Steinebrunn ( Austria); 2 spec. , NHMW 1855 /0045/0079, Steinebrunn ( Austria); 1 spec. , NHMW 2024 /0066/0001 (A239), Steinebrunn ( Austria); 2 spec. , NHMW 1851 /0013/0022, Bad Vöslau ( Austria); 6 spec. , NHMW 1869 /0001/0479, Forchtenau ( Austria); 1 spec. , NHMW 1848 /0003/0019, Ritzing ( Austria); 5 spec. , NHMW 1870 /0037/0016, Ritzing ( Austria); 1spec. , NHMW 1853 /0010/0019, Enzesfeld ( Austria); 3 spec. , NHMW 1847 /0037/0065, Mikulov ( Czech Republic); 3 spec. , NHMW 1847 /0046/0026, Szob ( Hungary); 8 spec. , NHMW 1854 /003570173, Lăpugiu de Sus ( Romania) .

Revised description. Large, solid, broadly fusiform shell with gradate spire; apical angle ~65°. Protoconch conical of three convex, smooth whorls. Teleoconch of up to five whorls. Suture impressed, shallowly undulating. Early teleoconch whorls convex with convex subsutural ramp, rounded shoulder, weakly convex below. Sculpture of 10 to 11 broad, prominent axial ribs overrun on antepenultimate and penultimate whorls by adis, IP and abis and slightly more prominent P1–P3; s1, s2 weak, recessed, adjoined by weakly scabrose tertiary threads. Later whorls with about six foliaceous varices alternating with weak intervarical rib. P1 with short spines on varices at shoulder; shorter spines may occur on P2 and P3. Last whorl attaining ~75% of total height; broad, shallow, weakly convex subsutural ramp, rounded shoulder, convex periphery, strongly constricted below. Axial sculpture of six prominent, raised, foliaceous varices alternating with weak intervarical rib. Subsutural ramp with weak adis, only slightly stronger IP, and prominent abis, occasionally bearing short spines. Interspaces and primary cords covered by slightly scabrose tertiary threads. P1–P6 (P6 on base of convex part of whorl), ADP, MP, ABP prominent, s1–s6 weaker, recessed, occasionally reduced. Primary cords with short spines on varices; spines of P1 most prominent; spines of ADP and MP prominent. Entire surface covered by delicate, narrow tertiary threads. Fasciole strongly swollen, twisted, delimiting moderately wide, moderately deep pseudoumbilicus. Aperture ovate to subcircular; outer lip thickened, crenulate with numerous short lirae starting at peristome, slightly incised at P1; lirae corresponding to the internal denticles ID, D1 to D6 (denticles usually bifid). Anal canal distinctly incised, U-shaped, accentuated by parietal denticle and weak ID. Siphonal canal long, open, moderately wide, bent to the left, curved to the right, dorsally recurved at tip. Columella strongly excavated, smooth, markedly angled at siphonal canal. Columellar callus forming broad, sharply delimited rim, erect at siphonal canal, adherent in parietal area.

Paratethyan synonyms. This species has a complex nomenclatorial history. Eichwald (1853) introduced the name Murex pomiformis for this species, overlooking the fact that this name was preoccupied. It had already been used by Mörch (1852) for an extant species. Later, Hörnes (1853) misidentified this species as Murex sedgwicki Michelotti, 1847 and provided the first good illustrations. Based on these, D’Ancona (1871, p. 30) introduced Murex hoernesi for Murex sedgwicki of Hörnes (1853) non Michelotti 1841. However, this is a junior homonym of Murex hörnesi Speyer, 1863 , a species from the Oligocene of Germany. Whilst D’Ancona (1871, p. 30) said his species encompassed the specimen illustrated by Hörnes (1856), he described and illustrated a specimen from the Italian Pliocene ‘colline Senesi’. Soon after, Tournouër (1875: 158) introduced Murex austriacus as new name for the specimens illustrated in Hörnes (1853: pl. 23, figs 1–5). Ten years later, De Gregorio (1885a) introduced Murex amberus for the specimen illustrated in Hörnes (1853: pl. 23, fig. 1). Finally, Csepreghy-Meznerics (1950) described Murex (Muricantha) sedgwicki vindobonensis from Hidas ( Hungary) (syntype illustrated in Csepreghy-Meznerics 1950: pl. 2, fig. 17). This name is preoccupied by Murex vindobonensis Hörnes, 1853 (= Ocenebra vindobonensis (Hörnes, 1853) . Consequently, most authors, such as Merle et al. (2011) and Landau et al. (2013), assumed that Murex austriacus Tournouër, 1875 was the earliest available name for the Paratethyan species. However, the first available name is Murex trunculoides Pusch, 1837 , based on a specimen from the Badenian of Poland. It would seem reasonable to treat Murex austriacus Tournouër, 1875 as a nomen protectum and to abandon Murex trunculoides Pusch, 1837 as a nomen oblitum. However, Article 23.9.1.1. of the ICZN (1999) requires that the senior synonym should not have been used as a valid name after 1899. This requirement is violated by the fact that Vokes (1971:111) listed Murex trunculoides as an available name. For Murex austriacus Tournouër, 1875 to be declared a nomen protectum, Article 23.9.1.2. requires that the junior synonym for a given taxon has been used as the presumably valid name in at least 25 papers published by at least 10 authors during the immediately preceding 50 years and covering a period of not less than 10 years.

As obvious from the chresonymy list provided, above, we were not able to fulfill this requirement. Therefore, we resurrect Murex trunculoides Pusch, 1837 as the name for this Paratethyan species, which is currently placed in Hexaplex (Trunculariopsis) .

Discussion. Hexaplex (Trunculariopsis) trunculoides (Pusch, 1837) shows slight variation in spire height. Specimens with high spire are reminiscent of Hexaplex (Trunculariopsis) bourgeoisi ( Tournouër, 1875) , from the Langhian of the Loire Basin ( France). Nevertheless, H. (T.) bourgeoisi is slenderer, its spire is slightly scalate and it develops prominent spines on P2–P6 on varices and its last whorl is less constricted. (see Merle et al. 2011: pl. 40, figs 5–6). Hexaplex (Trunculariopsis) turonensis ( Dujardin, 1837) , from the Langhian of the Loire Basin ( France), might be closely related but differs in its stronger P1, shorter spire and wider aperture (see Merle et al. 2011: pl. 37, figs 8–12; Landau et al. 2013: pl. 21, fig. 10). Hexaplex (Trunculariopsis) ledoni Ceulemans, Van Dingenen, Merle & Landau, 2016 , from the Tortonian the Loire Basin ( France), differs in its foliaceous varices and the slender conical protoconch (see; Landau et al. 2019: pl. 6, figs 1–3).

A specimen from the Burdigalian of Montpeyroux ( France), described by Goret & Pons (2013) as Hexaplex (Trunculariopsis) cf. austriacus ( Tournouër, 1875) differs in its higher spire and comparatively blunt sculpture with subobsolete secondary cords, and represents another species.

Paleoenvironment. Shallow marine, inner neritic (e.g., Gainfarn: Zuschin et al. 2007).

Distribution in Central Paratethys. Karpatian (Early Miocene): Korneuburg Basin: Stetten ( Austria) (Harzhauser 2002); Vienna Basin: Niederkreuzstetten ( Austria) (Harzhauser 2002). Badenian (Middle Miocene): Polish-Ukrainian Fore-Carpathian Basin: Zborów (Zboriv) ( Friedberg 1938); Korytnica Basin: Korytnica ( Bałuk 1995); Vienna Basin: Baden, Bad Vöslau, Enzesfeld, Gainfarn, Steinebrunn ( Austria), Mikulov ( Czech Republic) (Hoernes & Auinger 1885); Eisenstadt Sopron Basin: Forchtenau ( Austria) (Hoernes & Auinger 1885); Oberpullendorf Basin: Ritzing ( Austria) (Hoernes & Auinger 1885); Bakony Mountains: Herend-Márkó ( Hungary) ( Kókay 1966), Bánd ( Hungary) ( Kovács 2020); Mecsek Mountains ( Hungary) ( Strausz 1966); Pannonian Basin: Budapest Illés street, Hidas, Hosszúhetény, Kisbattyán, Letkés, Szob, Várpalota ( Hungary) (Hoernes & Auinger 1885; Csepreghy-Meznerics 1950; Strausz 1966; Kókay 1996; Kovács et al. 2018; Kovács 2020); Southern Pannonian Basin: Hrvaćani ( Bosnia and Hercegovina) ( Atanacković 1985); Făget Basin: Coşteiu de Sus, Lăpugiu de Sus ( Romania) ( Kovács 2019); Şimleu Basin: Tusa ( Romania) ( Nicorici 1972); Zârand Basin: Minişul de Sus ( Romania) ( Nicorici & Sagatovici 1973); Dacian Basin: Opanec, Târnene, Bivolare, Dobrusha (Kojumdgieva in Kojumdgieva & Strachimirov 1960; Nikolov 1994).

Proto-Mediterranean Sea. Serravallian (Middle Miocene): Karaman Basin: Seyithasan ( Turkey) ( Landau et al. 2013).

Figures

FIGURE 1. Cord diagrams and apertural features for Haustrinae [Haustrum? pyrulatum (Hoernes & Auinger, 1882)] and Muricinae [Bolinus gapus (De Gregorio, 1885a), Bolinus partschi (Hörnes, 1853), Chicoreus (Triplex) aquitanicus (Grateloup, 1833), Chicoreus (Triplex) hirmetzli (Kovács, 2018), Chicoreus (Triplex) notatus (Eichwald, 1830). Hexaplex (Trunculariopsis) trunculoides (Pusch, 1837), Hexaplex (Trunculariopsis) esplus (De Gregorio, 1885a), Hexaplex(Trunculariopsis) subasperrimus (d’Orbigny, 1852), Promurex spinicosta (Bronn, 1831), Exquisitor graniferus (Michelotti, 1841)].

FIGURE 3. Bolinus gapus (De Gregorio, 1885a). A1–A3. NHMW 1847/0046/0030, Szob (Hungary). B1–B3. paralectotype, NHMW 1860/0015/1270, Grund (Austria). C1–C3. NHMW1869/0001/0280, Grund (Austria). D1–D3. lectotype, NHMW 1856/0050/0122, Gainfarn (Austria).

FIGURE 8. Hexaplex (Trunculariopsis) trunculoides (Pusch, 1837). A1–A3.NHMW 1846/0037/0215, Gainfarn (Austria). B1–B3. NHMW 1856/0050/0120, Gainfarn (Austria). C1–C3. NHMW 1855/0045/0428, Steinebrunn (Austria). D1–D3. NHMW 1856/0050/0455, Lăpugiu de Sus (Romania).