Artemidiconus granularis ( Borson, 1820 )

Harzhauser, Mathias & Landau, Bernard, 2016, A revision of the Neogene Conidae and Conorbidae (Gastropoda) of the Paratethys Sea, Zootaxa 4210 (1), pp. 1-178 : 36-41

Artemidiconus granularis ( Borson, 1820)

Figs 3 A, 4A1–A3,4B1–B3

Conus Granularis — Borson 1820: 196, pl. 1, fig. 3.

Conus (Stephanoconus) Stachei nov. form.— Hoernes & Auinger 1879: 16, pl. 6, figs 14–16.

[ Conus granuliferus Grat. var.] Drnowitzensis de Greg.— De Gregorio 1885: 376.

[ Conus granuliferus Grat. var.] opellus de Greg. — De Gregorio 1885: 376 [nov. nom. pro Conus stachei in Hoernes & Auinger 1879 , pl. 6, fig. 14].

[ Conus ] H.[emiconus] granularis var. Stachei (H. A.)—Sacco 1893: 123, pl. 11, fig. 37.

[ Conus (Hemiconus) granularis var. Stachei ] var. druowitzensis [sic] De Greg.—Sacco 1893: 123 [nov. nom. pro Conus stachei in Hoernes & Auinger 1879 , pl. 6, fig. 15].

[ Conus (Hemiconus) granularis var. Stachei ] var. lissitzensis Sacc. —Sacco 1893: 123 [nov. nom. pro Conus stachei in Hoernes & Auinger 1879 , pl. 6, fig. 16].

Hemiconus granularis Bors. —Friedberg 1911: 45, pl. 2, fig. 10.

Conus (Hemiconus) granularis Bors. var. stachei Hörn. u. Auin.— Meznerics 1932 /1933: 346, pl. 14, fig. 2.

Conus (Stephanoconus) stachei Hoernes et Auinger—Csepreghy-Meznerics 1956: 421, pl. 3, fig. 7.

Conus (Hemiconus) granularis Borson—Kojumdgieva in Kojumdgieva & Strachimirov 1960: 208 , pl. 49, fig. 2.

Conus (Hemiconus) granularis stachei Hoernes & Auinger—Strausz 1966: 450 , pl. 66, fig. 9.

Conus (Stephanoconus) granularis stachei Hoernes et Auinger, 1879 — Hinculov 1968: 152, pl. 38, figs 11a–b.

Conus granularis Bronn, 1820 [sic]— Zelinskaya et al. 1968: 228, pl. 51, fig. 19.

Conus (Hemiconus) granularis stachei Hoernes et Auinger—Csepreghy-Mezneric 1972: 34, pl. 17, figs 23–24.

Conus granularis Borson—Pavia 1976: 157 , pl. 2, fig. 12 (holotype).

Conus (Hemiconus) granularis stachei ( Hoernes & Auinger, 1879) — Atanacković 1985: 181, pl. 40, figs 13–14.

Hemiconus granularis ( Borson, 1820) — Bałuk 1997: 54, pl. 20, figs 1–4.

Conilithes granularis ( Borson, 1820) — Kovács & Vicián 2013: 93, fig. 160.

Type material. Lectotype (following ICZN Article 74.6), illustrated in Borson (1820, pl. 5, fig. 3) and Pavia (1976, pl. 2, fig. 12), stored in the Museo Regionale di Scienze Naturali, Torino. The type locality is given as Valle Andona ( Italy), which is of Pliocene age. Pavia (1876) assumed that this was an error by Borson (1820) and pointed out that the preservation corresponds to material from the Messinian of Borelli (Turin Hills). The type localities of the taxa considered to be junior synonyms of A. granularis are: Conus stachei Hoernes & Auinger, 1879 , Baden ( Austria) ; Conus drnowitzensis De Gregorio, 1885 , Drnovice u Vyškova (Czech Republic); Conus opellus De Gregorio, 1885 , Baden (Austria); Conus lissitzensis Sacco, 1893 , Lysice (Czech Republic); all are of middle Miocene (Badenian/Langhian) age.

Studied material. 6 spec. NHMW 1864 /0001/0441, Drnovice u Vyškova ( Czech Republic), including holotype of Conus granuliferus drnowitzensis De Gregorio, 1885 ; 3 spec . NHMW 1865 /0015/0004, Lysice ( Czech Republic), including holotype of Conus granularis lissitzensis Sacco, 1893 ; 2 spec . NHMW 1865 /0036/0103, Lysice ( Czech Republic) ; 4 spec. NHMW 1871 /0011/0001, Sedlec ( Czech Republic) ; 1 spec. NHMW 1861 /0040/ 0 0 28, Hrušovany ( Czech Republic) ; 1 spec. NHMW 1999 z0077/0025, Baden ( Austria), designated herein as lectotype of Conus stachei Hoernes & Auinger, 1879 , holotype of Conus granuliferus opellus De Gregorio, 1885 ; 20 spec. NHMW 1863 /0015/0604, Niederleis ( Austria) ; 1 spec. NHMW 1887 /0018/0014, Steinebrunn ( Austria), illustrated in Meznerics (1932 /1933, pl. 14, fig. 2); 5 spec . NHMW 1846/0037/0038, 17 spec. NHMW 1868 /0001/ 0 0 93, Steinebrunn ( Austria) .

Illustrated material. Figs 3 A, 4A1–A3: Lysice ( Czech Republic): SL: 15.1 mm, MD: 8.0 mm, NHMW 1865/ 0015/0004, specimen illustrated in Hoernes & Auinger (1879, pl. 6, fig. 16) (= holotype of Conus granularis lissitzensis Sacco, 1893 ); Figs 4 B1–B3: Baden ( Austria): SL: 15.2 mm, MD: 7.9 mm, NHMW 1999z0077/0025, specimen illustrated in Hoernes & Auinger (1879, pl. 6, fig. 14) (= holotype of Conus granuliferus opellus De Gregorio, 1885 ).

Revised description. Small shells of about 13–15 mm in length, biconical, with moderately high conical spire and stout, ventricose last whorl; slightly constricted at base. Protoconch high conical, comprising at least three whorls. Early spire whorls forming a narrow, beaded upper sutural band and a tuberculate lower part. Subsequent spire whorls consisting of two adsutural spiral bands separated by a narrow groove; a secondary spiral cord appears between the spiral bands on late spire whorls; upper band usually bearing small beads. Subsutural flexure very shallow, moderately curved, nearly symmetrical. Last whorl weakly angulated in subadult shells, rounded in adults; broad and weak nodes appear along the shoulder in some specimens resulting in a wavy appearance. Widely spaced pustulose spiral ridges on lower half of last whorl grading into sharp spiral cords on ventral side. Upper half of last whorl with sharp spiral cords, which rarely bear beads. Base faintly constricted. Aperture straight, slightly broadening in abapical half. Fasciole short, moderately swollen, covered with spiral threads. No colour pattern observed.

Shell measurements and ratios. n = 13 adult specimens; largest specimen: SL: 15.2 mm, MD: 7.9 mm, mean SL: 13.6 mm (σ = 0.8), mean MD: 7.2 (σ = 0.4), spire angle: µ = 68° (σ = 2.9°), last whorl angle: µ = 41° (σ = 4.6°), LW: µ = 1.9 (σ = 0.07), RD: µ = 0.74 (σ = 0.04), PMD: µ = 0.9 (σ = 0.05), RSH: µ = 0.29 (σ = 0.03).

Discussion. This species is synonymized with the Italian early Miocene to Pliocene Conus granularis ( Borson, 1820) (e.g. Bałuk 1997). Conus granuliferus Grateloup, 1835 , from the early Miocene of the Aquitaine Basin, was also considered a synonym of C. granularis by most authors (e.g. Hall 1966, Lozouet et al. 2001). The specimens from the Badenian of the Paratethys seemingly differ from the Italian and French specimens in their mode of spire formation. Hall (1966) and Davoli (1972) described and figured a spire sculpture consisting of two cords separated by a deep groove. The same sculpture is seen in the somewhat abraded holotype of Conus granularis as illustrated in Pavia (1976) and in the French specimen illustrated in Lozouet et al. (2001). The Paratethyan shells, however, develop a secondary cord on the groove separating the two main cords. Some specimens of A. granularis from the Tortonian of Italy in the NHMW-collection, however, show a clear secondary cord as well. Therefore, this feature seems to reflect intraspecific variability and consequently we do not separate the Paratethyan specimens as a distinct species.

Sacco (1893b) placed the Italian Conus granularis Borson, 1820 in Hemiconus Cossmann, 1889 , which in our opinion is an exclusively Eocene genus. Consequently, Friedberg (1911) and Bałuk (1997) treated the Paratethyan species also as Hemiconus . Its type species, Hemiconus stromboides ( Lamarck, 1803) , from the Eocene of the Paris Basin, is high spired fusiform species with blunt nodes on the spire whorls. Other species placed in Hemiconus by Cossmann & Pissarro (1911) show a similar spire sculpture of nodes and a granulose cord along the upper suture. These morphologies have nothing in common with the Miocene species and therefore the placement in Hemiconus is herein rejected. Tucker & Tenorio (2009) listed Conus granularis Borson, 1820 within Conilithes Swainson, 1840 , but the mode of spire formation in C. granularis is different from the scalariform spire of Conilithes . Moreover, the spirally elongate beads on the last whorl of C. granularis do not occur in any Conilithes species and we follow the recommendation by John K. Tucker (pers. comm.) to treat Conus stachei (= granularis ) tentatively as Artemidiconus , despite the large geographic gap between the European Miocene record and the extant Western Atlantic occurrence.

Artemidiconus granularis is a moderately variable species. Adult specimens are more ventricose than the comparatively more slender subadult ones, which have a more pronounced shoulder. Similarly, the sculpture of beads on the last whorl displays some variability. This intraspecific and ontogenetic variability led de Gregorio (1885) and Sacco (1893) to propose new variation names for each of the Conus stachei specimens illustrated by Hoernes & Auinger (1879). Consequently, one of the three names of de Gregorio (1885) and Sacco (1893) is an objective synonym of C. stachei and the other two are subjective synonyms. To settle this, a lectotype of C. stachei has to be selected. Thus, we designate the specimen illustrated as fig. 14 by Hoernes & Auinger (1879) as lectotype of Conus stachei .

Paleoenvironment. In the Vienna Basin, this species is found in shallow sublittoral settings partly with sea grass meadows (e.g. Zuschin et al. 2007).

Distribution in Paratethys. Badenian (middle Miocene): Vienna Basin: Baden, Steinebrunn, Niederleis ( Austria), Hrušovany nad Jevišovkou, Sedlec ( Czech Republic) ( Hoernes & Auinger 1879; Sieber 1956); Bükk Mountains: Borsodbóta ( Hungary); Pannonian Basin: Szob ( Hungary) (Csepreghy-Mecnerics 1956; Strausz 1966; Kovács & Vicián 2013); Carpathian Foredeep: Drnovice u Vyškova, Lysice ( Czech Republic), Korytnica, Małoszów, Dryszczów ( Poland), Sboriw ( Ukraine) ( Hoernes & Auinger 1879; Zelinskaya et al. 1968; Bałuk 1997); Transylvanian Basin: Coşteiu de Sus, Lăpugiu de Sus ( Romania) ( Hoernes & Auinger 1879; Boettger 1902); Caransebeş-Mehadia Basin: Valea Bela Reca ( Romania) ( Hinculov 1968); Dacian Basin: Portitovci ( Bulgaria) ( Kojumdgieva & Strachimirov 1960); southern Pannonian Basin: Hrvaćani ( Bosnia and Herzegovina) ( Atanacković 1985).

Proto-Mediterranean Sea and north eastern Atlantic. Aquitanian ( France: Aquitaine Basin) ( Lozouet et al. 2001), Burdigalian ( Italy: Colli Torinesi; France Aquitaine Basin) ( Peyrot 1931; Hall 1966), Tortonian ( Italy: Sant'Agata Fossili, Stazzano Montegibbio) ( Davoli 1972).

Figures

FIGURE 4 A 1 – A 3. Artemidiconus granularis (Borson, 1820), Lysice (Czech Republic), NHMW 1865 / 0015 / 0004. 4 B 1 – B 3. Artemidiconus granularis (Borson, 1820), Baden (Austria), NHMW 1999 z 0077 / 0025. 4 C 1 – C 2. Conasprella berwerthi (Hoernes & Auinger, 1879), Steinebrunn (Austria), NHMW 1860 / 0001 / 0074 b. 4 D 1 – D 3. Conasprella berwerthi (Hoernes & Auinger, 1879), Steinebrunn or Gainfarn (Austria), NHMW 2015 / 0416 / 0002. 4 E 1 – E 3. Conasprella berwerthi (Hoernes & Auinger, 1879), Steinebrunn or Gainfarn (Austria), NHMW 2015 / 0416 / 0001. 4 F 1 – F 4. Conasprella berwerthi (Hoernes & Auinger, 1879), Steinebrunn (Austria), NHMW 1846 / 0037 / 0038. 4 G 1 – G 3. Conasprella minutissima nov. sp., Steinebrunn (Austria), NHMW 1869 / 0001 / 0331 c, paratype. 4 H 1 – H 3. Conasprella minutissima nov. sp., Steinebrunn (Austria), NHMW 1869 / 0001 / 0331 b, paratype. 4 I 1 – I 5. Conasprella minutissima nov. sp., Steinebrunn (Austria), NHMW 1869 / 0001 / 0331 a, holotype.