Conilithes brezinae ( Hoernes & Auinger, 1879 )

Harzhauser, Mathias & Landau, Bernard, 2016, A revision of the Neogene Conidae and Conorbidae (Gastropoda) of the Paratethys Sea, Zootaxa 4210 (1), pp. 1-178 : 48-50

Conilithes brezinae ( Hoernes & Auinger, 1879)

Figs 3 F, 5D1–D3, 5E1–E3, 5F1–F3, 5G1–G3, 5H1

Conus Dujardini Desh. —Hörnes 1851: 40 (partim), pl. 5, figs 8a–f [non Conus dujardini Deshayes, 1845 ].

[ Leptoconus ] [ Conus ] Brezinae n. f.— Hoernes 1878a: 195 (nomen nudum).

Conus (Leptoconus) Brezinae nov. form.— Hoernes & Auinger 1879: 36.

Conus Brezinae R. Hoern. i Auing.—Friedberg 1911: 51, pl. 2, figs 13–14.

Conus (Conospira) dujardini Desh. —Strausz 1954: 113, pl. 7, fig. 144.

Conus (Conospira) dujardini brezinae H. et Au.—Strausz 1954: 113, pl. 4, fig. 80.

Conus (Conuspira) brezinae R. Hörnes und Auinger, 1879 —Moisescu 1955: 161, pl. 14, figs 9, 12–14.

Conus (Conolithus) dujardini var. brezinae ( Hoernes und Auinger, 1879)—Kojumdgieva in Kojumdgieva & Strachimirov 1960: 209, pl. 49, fig. 7.

Conus (Conolithus) dujardini brezinae Hoernes & Auinger—Strausz 1962: 151 , pl. 22, fig. 16, pl. 43, figs 3–5, pl. 62, figs 8–9.

Conus (Conolithus) dujardini brezinae Hoernes & Auinger, 1879 — Strausz 1966: 452, pl. 22, fig. 16, pl. 43, figs 3–5, pl. 62, figs 8–9.

Conus (Conolithus) dujardini Deshayes 1845 — Hinculov 1968: 151, pl. 38, figs 6a–7 [non Conus dujardini Deshayes, 1845 ].

Conus (Conospira) dujardini brezinae Hoernes & Auinger 1879 —Bohn-Havas 1973: 1067, pl. 8, fig. 6.

Conus (Conolithus) dujardini Deshayes 1845 — Krach 1981: 75 (partim), pl. 21, figs 10, 14, 20, 23 [non Conus dujardini Deshayes, 1845 ].

Conus (Conolithus) dujardini brezinae R. Hoernes et M. Auinger, 1879 — Švagrovský 1981: 154, pl. 48, fig. 8.

Conus (Conolithus) dujardini Deshayes 1845 — Švagrovský 1982: 404, pl. 5, fig. 5.

Conus (Conolithus) dujardini brezinae Hoernes & Auinger—Atanacković 1985: 179 , pl. 40, figs 1–2.

Conus dujardini brezinae Hoernes et Auinger—Ionesi & Nicorici 1994: 62 , pl. 5, figs. 13–15.

Conus (Conolithus) dujardini Deshayes, 1845 — Bałuk 1997: 55, pl. 19, figs 1–4 [non Conus dujardini Deshayes, 1845 ].

Conus (Conolithus) brezinae Hoernes & Auinger—Schultz 1998: 72 , pl. 29, fig. 10.

Conus (Conolithus) dujardini Deshayes, 1845 — Mikuž 2009: 35, pl. 12, figs 159–161 [non Conus dujardini Deshayes, 1845 ].

Conilithes canaliculatus ( Brocchi, 1814) — Kovács & Vicián 2013: 91, figs 152–156 [non Conus canaliculatus Brocchi, 1814 ].

Conilithes dujardini ( Deshayes, 1845) — Janssen et al. 2014: 87, fig. 18 [non Conus dujardini Deshayes, 1845 ].

non Conus (Conolithus) brezinae Hoernes et Auinger, 1879 — Hinculov 1968: 151, pl. 38, figs 5a–b [= unidentified but not C. brezinae ].

non Conus (Conospira) brezinae Hoernes et Auinger—Csepreghy-Meznerics 1972: 33, pl. 17, figs 15–17 [= Conilithes exaltatus ( Eichwald, 1830) ].

Type material. Syntype NHMW 1999 z0077/0023a, illustrated in Hörnes (1851, pl. 5, fig. 8a) ; syntype NHMW 1999 z0077/0023b, illustrated in Hörnes (1851, pl. 5, fig. 8b) ; syntype NHMW 1999 z0077/0023c, illustrated Hörnes (1851, pl. 5, fig. 8c) ; syntype NHMW 1999 z0077/0023d, illustrated in Hörnes (1851, pl. 5, fig. 8d) ; syntype NHMW 1999 z0077/0023e, illustrated in Hörnes (1851, pl. 5, fig. 8e), all Steinebrunn ( Austria); middle Miocene , Badenian (late Langhian).

Studied material. Syntypes and 37 spec. NHMW 1846/0037/0040, 25 spec. NHMW 1846 /0037/0039, Gainfarn or Enzesfeld ( Austria) ; 12 spec. NHMW 1853/0010/0008, 9 spec. NHMW A 1614, Enzesfeld ( Austria), including specimen illustrated in Schultz (1998 pl. 29, fig. 10); 2 spec . NHMW 1997z0178/1848, 3 spec. NHMW 1985/0083/0159a,>100 spec. NHM 2013 /0479/ 1583-1602, Gainfarn ( Austria) ; 10 spec. NHMW A451, 29 spec. NHMW 1860 /0001/0076, Steinebrunn ( Austria) ; 2 spec. NHMW 2012 /0213/0104, Pfaffstätten ( Austria) ; 11 spec. NHMW A994, Guntersdorf ( Austria) ; 16 spec. NHMW A 995, Grund ( Austria) ; 24 spec. NHMW 1854 /0035/ 0 0 54, Lăpugiu de Sus ( Romania) .

Illustrated material. Figs 5 D1–D3: Steinebrunn (Austria), SL: 37.9 mm, MD: 15.9 mm, syntype NHMW 1999 z0077/0023a, illustrated in Hörnes (1851, pl. 5, fig. 8a); Figs 5 E1–E3: Steinebrunn ( Austria), SL: 36.9 mm, MD: 14.5 mm , syntype NHMW 1999 z0077/0023b, illustrated in Hörnes (1851, pl. 5, fig. 8b); Figs 5 F1–F3: Steinebrunn ( Austria), SL: 35.1 mm, MD: 14.1 mm , syntype NHMW 1999 z0077/0023c, illustrated in Hörnes (1851, pl. 5, fig. 8c); Figs 5 G1–G3: Gainfarn or Enzesfeld ( Austria): SL: 37.3 mm, MD: 17.8 mm , NHMW 1846 / 0037/0039; Fig. 5 H1: Gainfarn or Enzesfeld ( Austria) ; SL: 36.2 mm, MD: 16.1 mm, NHMW 1846 /0037/0039; Fig. 3 F: Steinebrunn ( Austria), SL: 26.6 mm, MD: 12.2 mm , syntype NHMW 1999 z0077/0023e, illustrated in Hörnes (1851, pl. 5, fig. 8e).

Revised description. Moderately small to medium sized, biconical shells, consisting of 10–12 teleoconch whorls. Protoconch high, conical and multispiral. Very high conical spire; early spire whorls with weakly tuberculate angulation just above suture; nodes become obsolete on 3rd to 4th spire whorl grading into a prominent carina, which successively migrates towards the lower suture abapically. Sutural ramp flat to slightly concave with prominent, raised growth lines. Subsutural flexure deep, strongly curved, moderately asymmetrical. Faint spiral threads appear on the sutural ramp of the early spire whorls in a few specimens. Last whorl slightly allometric with wider and less steep sutural ramp; weakly ventricose with slight constriction at base; siphonal canal very short and moderately wide. Aperture broadening slightly abapically, with thin, prosocyrt outer lip. Surface smooth, except for deep, broad spiral grooves on base, confined to the lower quarter of last whorl. Colour pattern in UV light consisting of irregularly spaced thin spiral lines on last whorl intercalated by 2–3 spirals of delicate dots. These stripes appear on the sutural ramp parallel to the subsutural flexure. Beads on spire coincide with short prosocyrt dots.

Shell measurements and ratios. n = 25: largest specimen: SL: 37.9 mm, MD: 17.8 mm, mean SL: 32.8 mm (σ = 2.9), mean MD: 14.3 (σ = 1.5), spire angle: µ = 58.4° (σ = 5.2°), last whorl angle: µ = 32.7° (σ = 2.4°), LW: µ = 2.3 (σ = 0.13), RD: µ = 0.63 (σ = 0.03), PMD: µ = 0.9 (σ = 0.03), RSH: µ = 0.31 (σ = 0.03).

Discussion. There is considerable confusion about the taxonomic status of Conus exaltatus Eichwald, 1830 , Conus dujardini Deshayes, 1845 and Conus brezinae Hoernes & Auinger, 1879 and in the literature all combinations of synonymizations can be found. The validity of C. brezinae as distinct species was especially doubted by many authors (e.g. Bałuk 1997; Landau et al. 2013; Kovács & Vicián 2013). After a re-examination of the type material we reject this decision. A principal components analysis (PCA) of the shell ratios ( Fig. 7A ) and measurements ( Fig. 7B ) reveals a very clear separation of Conilithes brezinae from C. exaltatus . Despite some variability in slenderness and spire height, C. brezinae is characterized by its marked shoulder and somewhat broader sutural ramp of the last whorl, the absence of striae or nodes on late spire whorls and the spiral grooves confined to the base. Conilithes brezinae further differs from C. exaltatus in the lower position of the carina, which is also less sharp, the lower height of the spire whorls, the shorter last whorl, which lacks a pronounced basal constriction, and the lack of the punctate spiral grooves.

With respect to this re-definition, the Serravallian specimens from the Turkish Karaman Basin, described by us ( Landau et al. 2013) as Conilithes dujardini [which we consider to be a subjective junior synonym of Conilithes exaltatus ( Eichwald, 1830) , see below], should be treated as C. brezinae . Kovács & Vicián (2013) proposed to synonymize Conilithes brezinae with the Pliocene C. canaliculatus ( Brocchi, 1814) . We do not accept this conclusion because C. canaliculatus has a more ventricose last whorl, its maximum diameter is below the shoulder, the sutural ramp of the last whorl is narrower and the spiral grooves cover a larger part of the base (see Malatesta 1974; Pinna & Spezia 1978; Chirli 1997).

Paleoenvironment. The occurrences in the Vienna Basin represent shallow water settings with sea grass meadows (e.g. Gainfarn; Zuschin et al. 2007). This ecological preference is an additional argument for a separation from C. exaltatus , which is typically found in offshore clays.

Distribution in Paratethys. Badenian (middle Miocene): Vienna Basin: Gainfarn, Enzesfeld, Steinebrunn, Grinzing, Pötzleinsdorf, Bad Vöslau, Niederleis ( Austria), Mikulov, Kienberk, Hrušovany, Sedlec ( Czech Republic), Devínska Nová Ves ( Slovakia) ( Hoernes & Auinger 1879; Sieber 1958b; Švagrovský 1981); Alpine- Carpathian Foredeep: Grund, Windpassing ( Austria), Lysice, ( Czech Republic) ( Hoernes & Auinger 1879; Sieber 1947, 1949); Korytnica, Babice, Błonie, Hołubica, Podhorce, Zukowce, Biłka, Tarnopol (Friedberg 1911; Bałuk 1997); Oberpullendorf Basin: Ritzing ( Austria) ( Hoernes & Auinger 1879: Sieber 1956); Bükk Mountains: Borsodbóta ( Hungary); Pannonian Basin: Várpalota, Balaton, Diósd, Hont, Letkés, Mátraverebély, Pécsszabolcs, Szob, Budapest: Illés street Szob, Hidas, ( Hungary) (Csepreghy-Meznerics 1954; Strausz 1966; Kovács & Vicián 2013); Styrian Basin: Pöls ( Austria) ( Hoernes & Auinger 1879); Transylvanian Basin: Lăpugiu de Sus, Coşteiu de Sus ( Romania); Caransebeş-Mehadia Basin: Valea Bela Reca ( Romania); Buzău Basin: Crivineni, Valea Muscel ( Romania) ( Hoernes & Auinger 1879; Hinkulo 1968; Ionesi & Nicorici 1994); Banja Luka Basin: Jazovac ( Bosnia and Herzegovina) ( Atanacković 1985); Krka Basin: Dolenja Brezovica ( Slovenia) ( Mikuž 2009).

Proto-Mediterranean Sea and north eastern Atlantic. This species is clearly also present in the Mediterranean area (e.g. Serravallian, Karman Basin, Turkey, Landau et al. 2013) but the confusion with C. exaltatus makes a critical evaluation of the literature data difficult.

Figures

FIGURE 5 A 1 – A 3. Conilithes allioni (Michelotti, 1847), Kleinebersdorf (Austria), NHMW 1976 / 1785 / 0044. 5 B 1 – B 2. Conilithes eichwaldi nov. nom., Zalisce (Ukraine), ZISP 62085. 5 C 1 – C 2. Conilithes eichwaldi nov. nom., Zalisce (Ukraine), ZISP 62085. 5 D 1 – D 3. Conilithes brezinae (Hoernes & Auinger, 1879), Steinebrunn (Austria), NHMW 1999 z 0077 / 0023 a, syntype. 5 E 1 – E 3. Conilithes brezinae (Hoernes & Auinger, 1879), Steinebrunn (Austria), NHMW 1999 z 0077 / 0023 b, syntype. 5 F 1 – F 3. Conilithes brezinae (Hoernes & Auinger, 1879), Steinebrunn (Austria), NHMW 1999 z 0077 / 0023 c, syntype. 5 G 1 – G 3. Conilithes brezinae (Hoernes & Auinger, 1879), Gainfarn or Enzesfeld (Austria), NHMW 1846 / 0037 / 0039. 5 H 1. Conilithes brezinae (Hoernes & Auinger, 1879), Gainfarn or Enzesfeld (Austria), NHMW 1846 / 0037 / 0039. 5 I 1 – I 3. Conilithes sceptophorus (Boettger, 1887), Lăpugiu de Sus (Romania), SMF XII. 2245 a, syntype. 5 J 1 – J 3. Conilithes antidiluvianus (Bruguière, 1792), Baden (Austria), NHMW 1869 / 0001 / 0161. 5 K 1. Conilithes antidiluvianus (Bruguière, 1792), Möllersdorf (Austria), NHMW 1869 / 0001 / 0247.

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.