A revision of the Neogene Conidae and Conorbidae (Gastropoda) of the Paratethys Sea
Author
Harzhauser, Mathias
Author
Landau, Bernard
text
Zootaxa
2016
4210
1
1
178
journal article
37280
10.11646/zootaxa.4210.1.1
e782e07d-76b7-4e9b-ba34-ed3286254ec6
1175-5326
252966
D39416B8-CF85-440B-84C2-D4380BECC4E3
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
.
FIGURE 3.
Oblique views of selected specimens.
A3.
Artemidiconus granularis
(Borson, 1820)
, Lysice (Czech Republic), NHMW 1865/0015/0004.
3B.
Conasprella berwerthi
(Hoernes & Auinger, 1879)
, Steinebrunn (Austria), NHMW 1846/0037/ 0 0 38.
Fig. 3C.
Conilithes antidiluvianus
(Bruguière, 1792)
, Möllersdorf (Austria), NHMW 1869/0001/0247.
3D.
Conilithes exaltatus
(Eichwald, 1830)
, Bad Vöslau (Austria), NHMW 1862/0001/0310.
3E.
Conasprella minutissima
nov. sp., Steinebrunn (Austria), NHMW 1869/0001/031c, paratype.
3F.
Conilithes brezinae
(Hoernes & Auinger, 1879)
, Steinebrunn (Austria), NHMW 1999z0077/0023e, syntype.
3G1–G2.
Kalloconus berghausi
(Michelotti, 1847)
, Lăpugiu de Sus (Romania), NHMW 1856/0050/0413.
Fig. 3H.
Kalloconus hendricksi
nov. sp., Pöls (Austria), NHMW 1861/0001/0226.
Fig. 3I.
Kalloconus hendricksi
nov. sp., Lăpugiu de Sus (Romania), NHMW 1870/003/0005c.
3J.
Kalloconus cacellensis
(Pereira da Costa, 1866), Lăpugiu de Sus (Romania), NHMW 1855/0043/0002.
3K.
Kalloconus gallicus
(Mayer-Eymar, 1890)
, Nemeşeşti (Romania), NHMW 2016/0036/0001.
3L.
Kalloconus hungaricus
(Hoernes & Auinger, 1879)
, Coşteiu de Sus (Romania), NHMW 1867/0029/0004.
3M.
Kalloconus letkesensis
nov. sp., Letkés (Hungary), private collection Anton Breitenberger.
3N.
Kalloconus moravicus
(Hoernes & Auinger, 1879)
, Mikulov-Kienberk (Czech Republic), NHMW 2016/003/0001.
3P.
Kalloconus ponderoaustriacus
(Sacco, 1893)
, Lăpugiu de Sus (Romania), NHMW 1854/0035/0041, holotype.
3Q.
Kalloconus ponderovagus
(Sacco, 1893)
, Niederkreuzstetten (Austria), NHMW 1849/0004/0016, holotype.
3R.
Lăpugiu de Sus (Romania),
Kalloconus pseudohungaricus
nov. sp., NHMW 1868/0001/0380c, paratype.
3S.
Kalloconus tietzei
(Hoernes & Auinger, 1879)
, NHMW 1847/0046/0004, Szob (Hungary), holotype.
3T.
Kalloconus tschermaki
(Hoernes & Auinger, 1879)
, Niederkreuzstetten (Austria), NHMW 1864/0001/0499, syntype.
3U.
Kalloconus voeslauensis
(Hoernes & Auinger, 1879)
, Bad Vöslau (Austria), NHMW 1849/0023/0005b, syntype.
3V.
Lautoconus eschewegi
(Pereira da Costa, 1866), Lăpugiu de Sus (Romania), NHMW 1858/0043/0007.
3W.
Lautoconus kovacsi
nov. sp., Letkés, (Hungary), NHMW 2016/0003/0004.
FIGURE 4A1–A3.
Artemidiconus granularis
(Borson, 1820)
, Lysice (Czech Republic), NHMW 1865/0015/0004.
4B1–B3.
Artemidiconus granularis
(Borson, 1820)
, Baden (Austria), NHMW 1999z0077/0025.
4C1–C2.
Conasprella berwerthi
(Hoernes & Auinger, 1879)
, Steinebrunn (Austria), NHMW 1860/0001/0074b.
4D1–D3.
Conasprella berwerthi
(Hoernes & Auinger, 1879)
, Steinebrunn or Gainfarn (Austria), NHMW 2015/0416/0002.
4E1–E3.
Conasprella berwerthi
(Hoernes & Auinger, 1879)
, Steinebrunn or Gainfarn (Austria), NHMW 2015/0416/0001.
4F1–F4.
Conasprella berwerthi
(Hoernes & Auinger, 1879)
, Steinebrunn (Austria), NHMW 1846/0037/0038.
4G1–G3.
Conasprella minutissima
nov. sp., Steinebrunn (Austria), NHMW 1869/0001/0331c, paratype.
4H1–H3.
Conasprella minutissima
nov. sp., Steinebrunn (Austria), NHMW 1869/0001/0331b, paratype.
4I1–I5.
Conasprella minutissima
nov. sp., Steinebrunn (Austria), NHMW 1869/0001/0331a, holotype.
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
).