Conilithes exaltatus ( Eichwald, 1830 )

Harzhauser, Mathias & Landau, Bernard, 2016, A revision of the Neogene Conidae and Conorbidae (Gastropoda) of the Paratethys Sea, Zootaxa 4210 (1), pp. 1-178 : 51-53

Conilithes exaltatus ( Eichwald, 1830)

Figs 3 D, 6B1–B3, 6C1–C3, 6D1–D3, 6E1–E3, 6F1–F2

[ Conus ] exaltatus m.— Eichwald 1830: 222.

Conus antidiluvianus Bruguière—Dubois de Montpéreux 1831: 23 , pl. 1, fig. 1 [non Conus antidiluvianus Bruguière, 1792 ).

Conus Dujardini Desh. — Deshayes 1845: 158.

Conus Dujardini Desh. —Hörnes 1851: 40 (partim), pl. 5, figs 3, 5–7 [non pl. 5, figs 8a–f = Conilithes brezinae ( Hoernes & Auinger, 1879) ].

Conus subacutangulus d’Orb. —d’Orbigny 1852: 58, nr. 1003 (nov nom. pro Conus antidiluvianus in Dubois de Montpéreux 1831: 23 , pl. 1, fig. 1].

Conus exaltatus — Eichwald 1852: plate captions, pl. 9, figs 3a–b.

Con. exaltatus m.— Eichwald 1853: 208.

Conus dujardini Desh. var. taurostriolata Sacco—Moisescu 1955b: 261 .

Conus (Leptoconus) Dujardini Desh. — Hoernes & Auinger 1879: 35.

Conus (Leptoconus) dujardini Desh. — Boettger 1906: 2.

Conus cf. avellana Lam. —Friedberg 1911: 56 (partim), pl. 2, fig. 20.

Conus Dujardini Desh. —Friedberg 1911: 47, pl. 2, fig. 11.

Conus Dujardini Desh. var. exaltatus Eichw. —Friedberg 1911: 51, pl. 2, fig. 12.

Conus (Conuspira) [sic] dujardini Deshayes, 1831 — Moisescu 1955a: 159, pl. 14, figs 15–18.

Conus (Conuspira) antideluvianus Bruguière var. buiturica var. nov . — Moisescu 1955a: 162, pl. 14, figs 7–8.

Conus (Conolithus) dujardini Deshayes 1845 —Kojumdgieva in Kojumdgieva & Strachimirov 1960: 209, pl. 49, fig. 4.

Conus (Conolithus) dujardini Deshayes—Atanacković 1963: 77 , pl. 15, figs 3–3a.

Conus (Conolithus) dujardini Deshayes, 1845 — Strausz 1966: 451, pl. 67, figs 2–5.

Conus (Conolithus) dujardini Deshayes—Bałuk 1970: 119 , pl. 13, figs 15–16.

Conus (Conolithus) dujardini Desh. — Stancu et al. 1971: 126, pl. 8, fig. 9.

Conus dujardini Deshayes 1845 — Eremija 1971: 79, pl. 5, fig. 9.

Conus (Conospira) dujardini astensis Sacco—Csepreghy-Meznerics 1972: 34, pl. 17, figs 12–13.

Conus (Conospira) dujardini brezinae Hoernes & Auinger—Csepreghy-Meznerics 1972: 34, pl. 17, figs 15–17.

Conus (Conolithus) dujardini Deshayes 1845 — Nicorici & Sagatovici 1973: 175, pl. 27, figs 10–12.

Conus (Conospira) dujardini Deshayes 1845 —Bohn-Havas 1973: 1066, pl. 8, figs 1–2.

Conus (Conolithus) dujardini Deshayes, 1845 — Atanacković 1985: 178, pl. 39, figs 18–19.

Conus (Conolithus) exaltatus Eichwald, 1853 — Bałuk 1997: 57, pl. 19, figs 5–8.

Conus (Conolithus) dujardini Deshayes—Schultz 1998: 72 , pl. 29, fig. 11.

Conus (Conolithus) dujardini dujardini Deshayes, 1845 — Chira & Voia 2001: 153, pl. 1, fig. 5.

Conus (Conolithus) dujardini Deshayes, 1845 —Harzhauser 2002: 112, pl. 9, fig. 12.

Conus (Conolithus) dujardini Deshayes, 1845 — Caze et al. 2010: 32, fig. 5N.

Conilithes dujardini ( Deshayes, 1845) — Kovács & Vicián 2013: 92, figs 157–159.

Type material. Syntypes described by Eichwald (1830) from Bilozirka (= Bialazurka, Bialozurka) and Shushkivtsi (= Shukowze) about 50 km NE of Ternopil ( Ukraine) (see map in Dubois de Montpéreux 1831). The specimens were most probably stored in the Zoological Museum of St. Petersburg ( Russia) but could not be identified on request; middle Miocene, Badenian (Langhian).

Studied material. 33 spec. NHMW 1846 /0037/0037, Baden ( Austria) including specimen illustrated in Hörnes (1851, pl. 5, fig. 3d), 32 spec . NHMW 1846 /0037/0036, Baden ( Austria) including specimens illustrated in Hörnes (1851, pl. 5, figs 3a, 3b, 3c), 12 spec . NHMW 1866/0011/0182, 12 spec. NHMW 2007z0078/007032, 24 spec. NHMW 2013 /0078/052-2013/0078/0523, Baden ( Austria); 22 spec . NHMW 1862/0001/0310,> 100 spec.

NHMW 2010 /0004/ 1343-2010 /0004/1345, Bad Vöslau ( Austria) ; 5 spec. NHMW 1973/1615/0214; 19 spec. NHMW A 1616, Baden-Sooß ( Austria) including specimen illustrated in Schultz (1998, pl. 29, fig. 11); 1 spec . NHMW 2013/0479/1603, 1 spec. NHMW 1970 /1396/1231, Gainfarn ( Austria) ; 26 spec. NHMW 1867 /0019/0013, Coşteiu de Sus ( Romania) ; 59 spec. NHMW 1870 /0033/0012, 16 NHMW A 1617, Lăpugiu de Sus ( Romania) .

Illustrated material. Figs 6 B1–B3: Möllersdorf (Austria): SL: 18.6 mm, MD: 7.2 mm, NHMW 2016 /0002/ 0001; Figs 6 C1–C3: Baden ( Austria): SL: 48.5 mm, MD: 17.9 mm , NHMW 1866 /0011/0182; Figs 6 D1–D3: Baden ( Austria): SL: 44.9 mm, MD: 17.6 mm , NHMW 1846 /0037/0036, illustrated in Hörnes (1851, pl. 5, fig. 3), Figs 6 E1–E3: Baden ( Austria): SL: 48.6 mm, MD: 19.6 mm , NHMW 1846 /0037/0036; Figs 6 F1–F2: SL: 26.2 mm, MD: 11.3 mm , MNHN A31836, Coşteiu de Sus ( Romania) (pic: P. Loubry, MNHN) ; 3D: Bad Vöslau ( Austria), SL: 27.7 mm, MD: 11.6 mm, NHMW 1862 /0001/0310.

Revised description. Moderately small to medium-sized elongate shells with very high, scalariform spire and long last whorl. Protoconch high and conical, consisting of 3 whorls. Spire whorls faintly striate with sharp carina slightly below mid-whorl; sutural ramp flat to weakly concave. Tubercles on carina present only on first 3–4 spire whorls. Several deeply incised, punctate spiral grooves below carina on spire whorls and last whorl. Subsutural flexure deep, strongly curved, moderately asymmetrical. Last whorl strongly angular at shoulder with maximum diameter at carina; constricted at base with long, slightly widening and weakly recurved canal. Punctate spiral grooves on lower half of last whorl, extending over the adapical half in occasional specimens. Last whorl of subadult shells often completely covered by grooves. Colour pattern showing irregular blotches on the spire and shoulder and spirally arranged subquadratic dots on the cords on the last whorl, amalgamating to irregular blotches.

Shell measurements and ratios. n = 26 adult specimens: largest specimen: SL: 48.6 mm, MD: 19.6 mm, mean SL: 40.8 mm (σ = 4.9), mean MD: 16.4 mm (σ = 0.4), spire angle: µ = 59.7° (σ = 4.9°), last whorl angle: µ = 30.9° (σ = 1.5°), LW: µ = 2.5 (σ = 0.15), RD: µ = 0.56 (σ = 0.02), PMD: µ = 0.9 (σ = 0.03), RSH: µ = 0.29 (σ = 0.04).

Discussion. The status of Conus dujardini Deshayes, 1845 and Conus exaltatus Eichwald, 1830 is controversially discussed in the literature (e.g. Bałuk 1997; Kovács & Vicián 2013). Conus dujardini was introduced by Deshayes (1845) without clearly stating type material or type area because he referred to material from Ukraine, Austria and France. From the context, it is obvious, that he had French specimens at hand. In any case he referred to the illustration of Dubois de Montpéreux (1831, pl. 1, fig. 1), which was later designated the lectotype of Conus dujardini by Glibert (1952b). The specimen of Dubois de Montpéreux originates from one of three villages in the area of Bilozirka (= Bialazurka, Bialozurka), 50 km ENE of Ternopil (=Tarnopol) in Ukraine (see map in Dubois de Montpéreux 1831). The specimens described by Eichwald (1830, 1853) as Conus exaltatus were collected at the same localities. As already discussed by Kovács & Vicián (2013), both specimens seem to be conspecific, which is also supported by the PCA; based on shell measurements, both specimens clearly plot with the specimens described herein as Conilithes exaltatus ( Fig. 7B ). In a PCA plot based on shell-ratios ( Fig. 7A ), both specimens plot close to C. brezinae , together with other stout morphologies of C. exaltatus (see remarks paragraph for C. brezinae for the separating features). Hence, it is highly likely that C. exaltatus and C. dujardini are conspecific, making C. dujardini a subjective junior synonym. The name C. dujardini has been extensively used in the literature, however it cannot be favoured over C. exaltatus as Friedberg (1911) used the name as a variety of C. dujardini , and Bałuk (1997) figured a specimen under this name. D’Orbigny (1852) introduced Conus subacutangulus as new name for the specimen, which was illustrated by Dubois de Montpéreux (1831, pl. 1, fig. 1) erroneously as Conus antidiluvianus . Conus subacutangulus is thus a subjective junior synonym of C. exaltatus Eichwald, 1830 .

Paleoenvironment. The majority of specimens in the NHMW collection derive from offshore clays suggesting inner to outer shelf environments as preferred habitat.

Conilithes brocchii in the Paratethys. Hörnes (1851, pl. 5, figs. 5–7) illustrated three Austrian shells as variation of C. dujardini (= exaltatus ), which are characterized by a low spire and concave sutural ramp, which accentuate the carina as a slightly raised ridge. Later, Friedberg (1911, text-fig. 9) and Csepreghy-Meznerics (1972, pl. 17, fig. 11) documented comparable specimens from Ukraine and Hungary. Hall (1966) placed fig. 7 of Hörnes (1851) in Conus brocchii Bronn, 1828 , which was followed by other authors (e.g. Csepreghy-Meznerics 1972; Landau et al. 2013; Kovács & Vicián 2013). A comparison with Pliocene specimens of Conilithes brocchii from Italian localities (NHMW collection) revealed several differences. Aside from being smaller, the Paratethyan shells have a regularly conical, less ventricose last whorl and the spiral grooves on the base are punctate, deep and well defined, whereas in C. brocchii they are smooth and rather shallow without sharp margins. Also the formation of the characteristic sutural rim is not strictly homologous. It is a raised margin in C. brocchii , as opposed to a carina close to the suture in the Paratethyan specimen. Moreover, the “ brocchii ”-morphs are extremely rare in the Paratethyan material. Consequently, we consider these shells to represent aberrant specimens of Conilithes exaltatus (or C. brezinae ). Conilithes brocchii sensu Kovács & Vicián, 2013 from the Badenian of Letkés ( Hungary) has a slightly ventricose outline and rounded shoulder and is not conspecific with C. brocchii and C. exaltatus .

Distribution in Paratethys. Karpatian (early Miocene): Korneuburg Basin: Kleinebersdorf (Harzhauser 2002); Badenian (middle Miocene): Vienna Basin: Gainfarn, Enzesfeld, Steinebrunn, Bad Vöslau, Baden-Sooß, Möllersdorf, Pfaffstätten ( Austria), Sedlec ( Czech Republic); Alpine-Carpathian Foredeep: Grund, Windpassing ( Austria) ( Hoernes & Auinger 1879; Schultz 1998); Carpathian Foredeep: Węglinek, Łychów, Korytnica ( Poland); Hołubica, Biłka, Jasinów, Bilozirka, Shushkivtsi ( Ukraine) (Dubois de Montpéreux 1831; Bałuk 1997); Southern Carpathians: Dubova ( Romania); Bükk Mountains: Borsodbóta ( Hungary); Pannonian Basin: Bánd, Diósd, Herend, Hetvehely, Hidas, Kovácsvágás, Letkés, Márkháza, Mátraverebély, Nógrádszakál, Sámsonháza, Szob, Szokolya, Várpalota, Zebegény, Budapest: Rákos, Illés street ( Hungary) ( Strausz 1966; Csepreghy- Meznerics 1972; Kovács & Vicián 2013); Banja Luka Basin: Jazovac ( Bosnia and Herzegovina) (Atanacković 1963); Transylvanian Basin: Lăpugiu de Sus, Coşteiu de Sus, Bujtur ( Romania); Zârand Basin: Minişul de Sus ( Romania); Caransebeş-Mehadia Basin: Valea Bela Reca ( Romania) ( Moisescu 1955a; Stancu et al. 1971; Chira & Voia 2001; Caze et al. 2010); Dacian Basin: Opanec ( Bulgaria) ( Kojumdgieva & Strachimirov 1960).

Proto-Mediterranean Sea and northeastern Atlantic. This species is clearly also present in the Miocene of the Mediterranean area but the confusion with C. brezinae makes a critical evaluation of the literature data difficult.

Figures

FIGURE 6 A 1 – A 3. Conilithes antidiluvianus (Bruguière, 1792), Bad Vöslau (Austria), NHMW 2010 / 0004 / 1338. 6 B 1 – B 3. Conilithes exaltatus (Eichwald, 1830), Möllersdorf (Austria), NHMW 2016 / 0002 / 0001. 6 C 1 – C 3. Conilithes exaltatus (Eichwald, 1830), Baden (Austria), NHMW 1866 / 0011 / 0182. 6 D 1 – D 3. Conilithes exaltatus (Eichwald, 1830), Baden (Austria), NHMW 1846 / 0037 / 0036. 6 E 1 – E 3. Conilithes exaltatus (Eichwald, 1830), Baden (Austria), NHMW 1846 / 0037 / 0036. 6 F 1 – F 2. Conilithes exaltatus (Eichwald, 1830), Coşteiu de Sus (Romania), MNHN A 31836.

FIGURE 7 A – B. PCA based on shell measurements of specimens of 3 Conilithes species (NHMW collection, Table 1). A: shell ratios, B: shell measurements. A separation of C. antidiluvianus, C. exaltatus and C. brezinae is evident in both analyses. The measurements for the lectotype of C. dujardini and the syntype of C. exaltatus were taken from the illustrations in Eichwald (1952) and Dubois de Montpéreux (1831). SL = shell length, AH = apertural height, HMD: height of maximum diameter, SA = spire angle, LWA = angle of the last whorl, LW = length width ratio, RD = relative diameter, RSH = relative height of spire.