GEOLOGICA CARPATHICA
, AUGUST 2017, 68, 4, 329 – 349
doi: 10.1515/geoca-2017-0023
www.geologicacarpathica.com
The Miocene “Pteropod event” in the SW part of
the Central Paratethys (Medvednica Mt., northern Croatia)
MARIJA BOŠNJAK
1
, JASENKA SREMAC
2
, DAVOR VRSALJKO
1
,
ŠIMUN AŠČIĆ
2
and LUKA BOSAK
2
1
Croatian Natural History Museum, Demetrova 1, 10000 Zagreb, Croatia; marija.bosnjak@hpm.hr, davor.vrsaljko@hpm.hr
2
Department of Geology, Faculty of Science, University of Zagreb, Horvatovac 102 a, 10000 Zagreb, Croatia;
jsremac@geol.pmf.hr, simeascic@geol.pmf.hr, luka.bosak1812@gmail.com
This paper was inspired by the scientific opus of Professor Vanda Kochansky-Devidé,
the first woman elected to the Croatian Academy of Sciences and Arts, whose 100
th
birthday we celebrated in 2015.
(Manuscript received August 9, 2016; accepted in revised form June 6, 2017)
Abstract: Deep marine Miocene deposits exposed sporadically in the Medvednica Mt. (northern Croatia) comprise
pelagic organisms such as coccolithophores, planktic foraminifera and pteropods. The pteropod fauna from yellow marls
at the Vejalnica locality (central part of Medvednica Mt.) encompasses abundant specimens of Vaginella austriaca
Kittl, 1886, accompanied with scarce Clio fallauxi (Kittl, 1886). Calcareous nannoplankton points to the presence of NN5
nannozone at this locality. Highly fossiliferous grey marls at the Marija Bistrica locality (north-eastern area of
Medvednica Mt.) comprise limacinid pteropods: Limacina valvatina (Reuss, 1867), L. gramensis (Rasmussen, 1968) and
Limacina sp. Late Badenian (NN5 to NN6 nannozone) age of these marls is presumed on the basis of coccolithophores.
Most of the determined pteropods on species level, except V. austriaca have been found and described from this region
for the first time. New pteropod records from Croatia point to two pteropod horizons coinciding with the Badenian marine
transgressions in Central Paratethys. These pteropod assemblages confirm the existence of W–E marine connection
(“Transtethyan Trench Corridor”) during the Badenian NN5 nannozone. Limacinids point to the possible immigration of
the “North Sea fauna” through a northern European marine passage during the Late Badenian (end of NN5- beginning of
NN6 zone), as previously presumed by some other authors.
Keywords: Planktic gastropods, palaeoenvironment, Middle Miocene, Badenian, Central Paratethys, northern Croatia.
Introduction
Pteropoda are marine gastropods adapted to a holoplanktic
way of life by transforming their foot into the “wings” (para-
podia) for buoyancy and swimming. They live in oceans
around the world at various depths. Most of them are epi- or
mesopelagic (up to 1000 m), but some species live bathy-
pelagic, at depths of 1000 m or more. During the day, they
float at greater depths, and at sunset they rise up near the
surface (negative fototaxis). Pteropods are divided into two
orders: Thecosomata de Blainville, 1824 (“sea butterflies”)
and Gymnosomata de Blainville, 1824 (“sea angels”). Theco-
somata include mainly forms with aragonitic shell in the adult
stage, but in Gymnosomata a shell is only present in the larval
stage (e.g., Herman 1998; Janssen & Little 2010; Corse et al.
2013; Janssen & Peijnenburg 2014 and references therein).
The most abundant occurrences of pteropods in Paratethys
are recorded from the Middle Miocene deposits (e.g., Bohn-
Havas & Zorn 1993, 1994, 2002; Bohn-Havas et al. 2004),
coinciding with peak marine transgressions. The genera
Limacina Bosc, 1817, Vaginella Daudin, 1800 and Clio
Linnaeus, 1767 are the most diverse and numerous taxa
(Janssen 1984; Zorn 1991, 1995, 1999; Bohn-Havas & Zorn
1993, 1994, 2002; Bohn-Havas et al. 2004). In northern
Croatia, on the south-western margin of the Central Paratethys,
Miocene planktic gastropods were recorded by Gorjanović-
Kramberger (1908, p. 36), Kochansky (1944), Kochansky-
Devidé (1973), Basch (1983 b, p. 29), Magaš (1987, p. 19),
Pikija (1987, p. 20), Korolija & Jamičić (1989, p. 16) and
Avanić et al. (1995). Although pteropods were not studied in
detail by the aforementioned authors, we were able to conclude
from their records that Vaginella austriaca Kittl, 1886 was
the most widely distributed pteropod species (Gorjanović-
Kramberger 1908; Kochansky-Devidé 1973; Basch 1983 b;
Korolija & Jamičić 1989; Avanić et al. 1995), sporadically
associated with Clio pedemontana (Mayer, 1868) (Kochansky
1944; Kochansky-Devidé 1973; Basch 1983 b), “Spiratella
sp.” (Magaš 1987; Pikija 1987) and “Spirialis (= Limacina)
andrussowi Kittl, 1886” (Kochansky-Devidé 1973).
A significant number of pteropods were collected during the
recent field research in central and north-eastern parts of
Medvednica Mt. in northern Croatia. This paper describes the
Middle Miocene pteropod assemblages and palaeoenviron-
ments with implications for the wider palaeogeographical area
(Central Paratethys), and indicates possible faunal “migration
routes”.
Abbreviations used in this article are as follows (alpha-
betical order): A — aperture diameter; A
1
/A
2
-ratio — aperture
330
BOŠNJAK, SREMAC, VRSALJKO, AŠČIĆ and BOSAK
GEOLOGICA CARPATHICA
, 2017, 68, 4, 329 – 349
diameters ratio; α — apical angle; α
1
— aperture angle;
CNHM — Croatian Natural History Museum, Zagreb, Croatia;
H — shell height; H/W-ratio — shell height and width ratio;
Inv. No. — inventory number; H
t
— height of teleoconch;
MB — Marija Bistrica locality; UZFS-DG — University of
Zagreb, Faculty of Science, Department of Geology, Croatia;
V — Vejalnica locality; W — shell width.
Geological setting
The investigated localities are situated in the northern
Croatia: Vejalnica locality in the central part of the Medvednica
Mt. (45°54’17” N, 16°4’19” E) and the Marija Bistrica locality
in the north-eastern part of the Medvednica Mt. (45°58’56” N,
16°6’52” E), near Zagreb (Fig. 1).
During the Miocene, the area of northern Croatia formed the
south-western margin of the Central Paratethys, and geo-
tectonically belonged to the Pannonian Basin System (Pavelić
2001, 2002, 2005; Kováč et al. 2007). The Paratethys sea
flooded different types of basement in northern Croatia, and
significant palaeogeographic changes occurred during the
Middle Miocene (Badenian), when the marine transgressions
affected the Central Paratethys area (e.g., Rögl 1998; Rögl et
al. 2007; Kováč et al. 2007). There are disputes regarding age
estimations of the first Miocene marine transgression in
northern Croatia. It was initially dated as the Early Miocene
(Karpatian) transgression (e.g., Šikić et al. 1977, 1979; Basch
1983 a, b; Pavelić 2001), and recently some authors have con-
sidered it to be of the Middle Miocene (Badenian) age (Ćorić
et al. 2009; Pavelić 2015).
The Middle Miocene deposits crop out along the Medvednica
Mt. slopes (Kochansky 1944; Kochansky-Devidé 1957; Šikić
et al. 1977, 1979; Kochansky-Devidé & Bajraktarević 1981;
Basch 1983a,b; Avanić et al. 1995, 2003; Sremac et al. 2005;
Vrsaljko et al. 2006; Pezelj et al. 2007; Pezelj 2015; Brlek et
al. 2016 and references therein; Sremac et al. 2016 and refe-
rences therein; Pezelj et al. 2016) (Fig. 1A). Kochansky (1944)
described three types of Middle Miocene (Badenian) marine
developments on the Medvednica Mt. (“South-western”,
“Central” and “North-eastern” development), and areas
studied in this paper belong to the “Central” or “Čučerje”
development (Vejalnica outcrop), and to the “North-eastern”
or “Zelina” development (Marija Bistrica outcrop) (Fig. 1B).
The “Central-” part is characterized by the Miocene marl
deposits with open sea organisms (nautiloids, pteropods,
planktic foraminifers, nannoplankton) and rare specialized
benthic bivalve genus Solemya Lamarck, 1818 (Gorjanović-
Kramberger 1908; Kochansky 1944; Kochansky-Devidé 1957;
Basch 1983 b; Avanić et al. 1995). In contrast, the “North-
eastern” development is marked with rich planktic associa-
tions of foraminifera and calcareous nannoplankton, and
benthic molluscs findings (Kochansky 1944; Avanić et al.
2003). Regarding the character of Miocene marine deposits on
Medvednica Mt., pteropods were expected and found in both
the “Central-” and “North-eastern” development.
Material and methods
The study areas presented here were chosen according to the
previously published data (Kochansky 1944; Kochansky-
Devidé 1957; Avanić et al. 1995, 2003). Furthermore,
Miocene marine fauna from investigated localities studied by
V. Kochansky-Devidé (Kochansky 1944; Kochansky-Devidé
1957) is housed at CNHM and has recently been undergoing
revision (M. Bošnjak, doctoral thesis).
Research methods included field and laboratory work. Field
work conducted between 2014 and 2016 included collecting
fossil fauna and recording geological columns. For this
research, the authors chose the outcrops of the Middle Miocene
deep marine marls in the central area of the Medvednica Mt.,
and north-eastern Medvednica Mt. (for more information
see Basch 1983 a, b; Avanić et al. 2003; Sremac et al. 2016)
(Fig. 1).
During field work, marls from Vejalnica and Marija Bistrica
localities were collected for more detailed analyses, and 0.3 kg
of each sample were collected for the wet sieving technique.
From the Vejalnica locality, Miocene sediments alongside the
forest road crop out over a length of app. 15 m, and the total
thickness of layers is app. 2 m, of which 1.50 m is visible in
the road cutting, and 0.50 m is in the gully. We recognized 12
layers, each around 1 to 3 cm thick, and the majority of ptero-
pods were collected from the middle part of the gully. At the
second locality, Marija Bistrica, we investigated an outcrop of
sediments app. 10 m in length and 5 m in height. They are less
distributed and more eroded than the Vejalnica sediments, and
following the changes in lithology 4 layers could be distin-
guished. From these layers, samples for laboratory work were
collected for the purpose of another research, and during the
analyses of the washed material the limacinids presented here
were recognized.
Laboratory work included the wet sieving technique, analy-
ses and determination of molluscs and calcareous nanno-
fossils. Soft marls were crushed and soaked in water. After
twenty-four hours, samples were washed over meshes of 0.5,
0.2, 0.125 and 0.63 mm. After drying, the residue was studied
using stereo-microscope Olympus−SZX10 and polarizing
microscope Leica Laborlux 11, photographed by a Canon
EOS 1100 camera and saved through the Quick PHOTO
CAMERA 3.0 programme at UZFS-DG. Samples with fossil
molluscs were cleaned and photographed by a Canon EOS 6D
camera at CNHM. Pteropod specimens were photographed
under a TESCAN VEGA TS 5136 MM/Oxford scanning elec-
tron microscope at UZFS-DG. Calcareous nannofossil probes
were prepared by standard method (after Bown & Young
1998) in the Wet Laboratory of UZFS-DG. Thin sections were
studied at UZFS-DG by “Zetoplan Reichert” polarizing micro-
scope, using the 1250× and 1600× magnification, and photo-
graphed by a Canon EOS 400D camera. Nannofossils were
determined according to Perch-Nielsen (1985), Bown (1998),
Bartol (2009) and Young et al. (2014).
Pteropod samples are housed at CNHM, with assigned tem-
porary inventory numbers: CNHM MBa-LV 1 to 21 and
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THE MIOCENE “PTEROPOD EVENT” IN THE SW PART OF THE CENTRAL PARATETHYS
GEOLOGICA CARPATHICA
, 2017, 68, 4, 329 – 349
ZAGREB
0
50
100 km
A
d
r
i
a
t
i
c
S
e
a
HUN
BIH
SLO
I
SCG
SCG
A
C
R
O
A
T
I
A
CENTRAL
(ČUČERJE)
NORTH EASTERN
-
(ZELINA)
DEVELOPMENT
MIOCENE
MIDDLE
UPPER
LOWER
P
ANNONIAN
SARMA
TIAN
BADENIAN
KAR
P
A
TIAN
?
PAL EOZOIC-
A
MESOZOIC
MB
V
MB
V
BOUNDSTONE
CLAYSTONE
MARLSTONE
LITHOTHAMNIUM
LIMESTONE AND
SANDSTONE
SANDSTONE
BIOCLASTIC
LIMESTONE
VOLCANIC ASH
LACUSTRINE
LIMESTONE
COAL
BRECCIA
CONGLOMERATE
TRANSGRESS.
BOUNDARY
HIATUS
PAL EOZOIC-
A
MESOZOIC
BASEMENT
LACUSTRINE - MARINE
MIOCENE
PAL EOZOIC-
A
MESOZOIC
VEJALNICA
(N 45°54´17˝; E 16°4´19˝)
MARIJA BISTRICA
(N 45°58´56˝; E 16°6´52˝)
5 km
M
e
d
v
e
d
n
i c
a
M
t .
MB
V
LEGEND:
Z A G R E B
POST-MIOCENE
A
B
MEDITERRANEAN
ST
AGES
CENTRAL
P
ARA
TETHYS
ST
AGES
EPOCH
T
O
R
T
ONIAN
SERRA
V
ALIAN
LANGHIAN
BURDIGALIAN
CALCAREOUS
NANNOFOSSILS
NN4
NN5
NN6 - NN1
1
Fig. 1. A — Location of the investigated area on the Medvednica Mt., northern Croatia. B — Schematic geological columns of the studied
outcrops (after Vrsaljko et al. 2015). Calcareous nannofossils after Harzhauser & Piller (2007), Kováč et al. (2007), Piller et al. (2007) and
Pezelj et al. (2013).
332
BOŠNJAK, SREMAC, VRSALJKO, AŠČIĆ and BOSAK
GEOLOGICA CARPATHICA
, 2017, 68, 4, 329 – 349
CNHM MBb-LV 1 to 14 (Limacina valvatina, Marija Bistrica
locality), CNHM MB-LG 1 to 6 (Limacina gramensis, Marija
Bistrica locality), CNHM MB-L 1 to 2 (Limacina sp. (? nov.),
Marija Bistrica locality), CNHM V-CF 1 to 5 (Clio fallauxi,
Vejalnica locality) and CNHM V-VA 1 to 17 (Vaginella
austriaca, Vejalnica locality). Due to the importance of new
records of the Middle Miocene (Badenian) pteropods on the
Medvednica Mt., we present planktic gastropods together with
the biostratigraphically important nannoplankton. Other fossil
fauna will be addressed in future studies.
Systematic Palaeontology
Phylum: Mollusca Linnaeus, 1758
Class: Gastropoda Cuvier, 1795
Subclass: Heterobranchia Burmeister, 1837
Superorder: Pteropoda Cuvier, 1804
Order: Thecosomata de Blainville, 1824
Suborder: Euthecosomata Meisenheimer, 1905
Family: Limacinidae Gray, 1847
Genus: Limacina Bosc, 1817
Type species: Limacina helicina (Phipps, 1774)
Three limacinid species were recognized: Limacina
valvatina (Reuss, 1867), Limacina gramensis (Rasmussen,
1968) and Limacina sp. (? nov.). For measurements see
Fig. 2.
Limacina valvatina (Reuss, 1867)
1867 Spirialis valvatina – Reuss, p. 32, 146, pl. 6, fig. 11 a−b.
1886 Spirialis valvatina Reuss – Kittl, p. 69, pl. 2, fig. 38.
1981 Spirialis valvatina (Reuss) 1867 – Krach, p. 125, pl. 3, figs.
2−4, 7−8, pl. 5, figs. 1−2, ?9, 10−11, pl. 6, figs. 1−2.
1984 Spirialis valvatina Reuss, 1867 – Janssen, p. 72.
1991 Limacina valvatina (Reuss, 1867) – Zorn, p. 97, pl. 1, figs.
1‒6, pl. 10, figs. 1, 2, pl. 11, figs. 4, 5.
1993 Limacina valvatina (Reuss, 1867) – Janssen & Zorn, p. 179, pl.
1, figs. 4‒11, pl. 2, figs. 1‒11, pl. 3, figs. 1‒12.
1999 Limacina valvatina (Reuss, 1867) – Zorn, p. 728, pl. 1, figs. 4,
6‒10.
2005 Limacina valvatina (Reuss, 1867) – Suciu et al., p. 455, pl. 1,
figs. 15‒17.
2010 Limacina valvatina (Reuss, 1867) – Cahuzac & Janssen, p. 47,
pl. 12, figs. 6‒12.
Material examined: Thirty-five casts from grey marls,
Marija Bistrica locality (temporary inventory numbers CNHM
MBa-LV 1 to 21 and CNHM MBb-LV 1 to 14).
Description: Shells with low spire, up to 4 whorls (Fig. 3).
Height/width-ratio is below 110, a value considered
a reliable distinguishing characteristic between L. valvatina
and L. gramensis (Janssen & Zorn 1993). The apical angle
varies from ≤ 88° to max ≤158°. Dimensions of shell height,
width and apical angles are given in Table 1 and Fig. 4.
Measured specimens show normal variability in H/W-ratio
(Fig. 4 A, B), but in the histogram of apertural shape variability
is larger (Fig. 4 C). This variability of measurement span could
be partly result from a compression.
Remarks: Specimens are pyritized. The shape of fossil
Limacinidae species, including L. valvatina, is more diverse
than in the few existing recent species, as demonstrated, for
example, by the missing umbilicus in some species, size and
shape of body whorl and the morphology of the apertural
margin, or ornamentation of the shell (Janssen 2003).
Specimens shown on Fig. 3 could be juvenile, and are very
low spired. Similar forms identified as “Spiratella (Spiratella)
krutzschi Tembrock, 1989” were found in northern Germany
(Gorlosen, south of Ludwigslust) (Tembrock 1989), which
Janssen (1999) considered a junior synonym of Limacina
valvatina.
Occurrence: These are the first records of Limacina
valvatina from the Badenian sediments of northern Croatia. Its
occurrence in the Middle Miocene of Paratethys has been
docu mented from the Early to the Late Badenian, and records
from the Lower Sarmatian deposits are mentioned (Bohn-
Havas & Zorn 2002), e.g., in Romania (Bohn-Havas & Zorn
1994, p. 78, Abb. 4) and in Bulgaria (Nikolov 2010). However,
the species was most widely distributed during the Badenian
(Bohn-Havas & Zorn 2002). Paratethyan records are available
from Hungary (Bohn-Havas & Zorn 1993, 1994; Bohn-Havas
et al. 2004) in the NN5 nannozone after Bohn-Havas & Zorn
(1993) and Selmeczi et al. (2012), Austria (Zorn 1991; Bohn-
Havas & Zorn 1993, 1994), Poland (Bohn-Havas & Zorn
1993, 1994; Janssen & Zorn 1993; Janssen 2012), Czech
Republic (Bohn-Havas & Zorn 1994; Zorn 1999), Slovakia
(Zorn 1999), Romania (Bohn-Havas & Zorn 1994; Suciu et al.
2005), Bulgaria (Nikolov 2010) and Ukraine (Janssen & Zorn
1993; Bohn-Havas & Zorn 1994). The largest specimens are
recorded in Poland (Janssen & Zorn 1993). Outside Paratethys,
L. valvatina is known from younger Miocene deposits of the
North Sea Basin (Zorn 1999; Bohn-Havas & Zorn 2002;
W
H
α
A
1
A
2
Fig. 2. Measured elements of Limacina shell (after Zorn 1991).
Abbreviations: H — shell height; W — shell width; α — apical angle;
A
1
, A
2
— aperture diameters.
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THE MIOCENE “PTEROPOD EVENT” IN THE SW PART OF THE CENTRAL PARATETHYS
GEOLOGICA CARPATHICA
, 2017, 68, 4, 329 – 349
Janssen 2012), Early and Middle
Miocene of the Aquitaine Basin,
France (Cahuzac & Janssen 2010;
Janssen 2012), Langhian of north-
ern Italy and Sicily (Jannsen
2012), and Miocene deposits of
the Maltese Archipelago (Janssen
2012) and Germany (Tembrock
1989; Jannsen 1999). Also,
Limacina cf. valvatina is known
from the older Miocene sediments
of Val Ceppi, Italy (Janssen 1995;
Zorn 1999).
Limacina gramensis
(Rasmussen, 1968)
1886 Spirialis stenogyra (Philippi)
– Kittl, p. 67, text-figs. 35, 36
(non Philippi).
1968 Spiratella gramensis – Ras mus-
sen, p. 244, pl. 27, figs. 4−7.
1984 Spirialis stenogyra (Philippi,
1844) – Janssen, p. 71, pl. 3,
figs. 1, 2 (non Philippi).
1993 Limacina gramensis (Ras mus-
sen, 1968) – Janssen & Zorn,
p. 175, pl. 3, figs. 13, pl. 4,
figs. 1‒9, pl. 5, figs. 1‒3.
1999 Limacina gramensis (Ras mus-
sen, 1968) – Zorn, p. 726, pl. 1,
figs. 1‒3, 5.
Material examined: Six casts
from grey marls at the Marija
Bistrica locality (temporary
inventory
numbers
CNHM
MB-LG 1 to 6).
Description: Slender small shells
(possibly juvenile) with high spire,
3 to 5 whorls (Fig. 5 A, B). Height/
width-ratio above 110. Apical
angle varying from ≤ 44° to ≤ 80°
may be consequence of compres-
sion. Straight spire tangents.
Dimensions of shell height, width
and apical angle are given in Table 2. Fig. 6 shows shell height
and width relationship of studied specimens.
Remarks: Casts are pyritized and slightly deformed
by pressure. Fig. 6 shows shell height/width-ratio of
L. gramensis, but the number of collected specimens is too
small to perform conclusive statistical analyses.
Occurrence: The sporadic presence of pteropods (deter-
mined as Spiratella and Spirialis) in the Middle Miocene
deposits of northern Croatia were reported by Magaš (1987)
and Pikija (1987), during the preparation of the Basic
Geological Map, sheets Sisak and Osijek. Unfortunately, this
material was not studied later and is today unavailable.
The present research proves the first record of Limacina
gramensis in the Badenian deposits of northern Croatia. Up to
today the occurrence of L. gramensis within the Central
Paratethys is limited to the Upper Badenian deposits and the
species is considered to be an index fossil (e.g., Bohn-Havas
& Zorn 2002). It is known from Poland (Bohn-Havas &
Zorn 1993; Janssen & Zorn 1993; Zorn 1999), Czech Republic
(Zorn 1999), Romania (Janssen & Zorn 1993; Zorn 1999),
Bulgaria (Nikolov 2010) and Ukraine (Janssen & Zorn 1993;
Zorn 1999). Outside Paratethys, this species is known from
Miocene of the North Sea Basin (Zorn 1999; Bohn-Havas &
Zorn 2002).
Fig. 3. Limacina valvatina (Reuss, 1867) from the Marija Bistrica locality. A — CNHM MBa-LV 6;
B — CNHM MBa-LV 8; C — CNHM MBa-LV 10; D — CNHM MBa-LV 14; E — CNHM MBa-LV
18; F — CNHM MBb-LV 3; G — CNHM MBb-LV 6; H — CNHM MBb-LV 12. Scale bars 100 μm.
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BOŠNJAK, SREMAC, VRSALJKO, AŠČIĆ and BOSAK
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Limacina sp. (? nov.)
Material examined: Two casts from grey marls at the Marija
Bistrica locality (temporary inventory numbers CNHM MB-L
1 and 2).
Description: Small shells (possibly juvenile), oval-conical
shaped with high spire and 3 preserved whorls (Fig. 5 C, D).
Apical region blunt; aperture not preserved. Height/width-
ratio above 110. Straight spire tangents. Dimensions of shell
height, width and apical angle are given in Table 2.
Remarks: Casts are pyritized and slightly deformed by
pressure. These two specimens (MB-LG 1 and 2, Fig. 5 C, D)
differ from the other six L. gramensis specimens by shape,
which is more rounded than in L. gramensis (Fig. 5 A, B) and
lacks lateral widening of last whorls. The described two speci-
mens are indicated as Limacina sp. (? nov.), but their number
is not sufficient for a valid description of a new species.
Representatives of the families Cliidae and Cavoliniidae
were also determined and presented in this paper. The mea-
sured elements are shown on Fig. 7.
.
Family: Cliidae Jeffreys, 1869
Genus: Clio Linnaeus, 1767
Type species: Clio pyramidata Linnaeus, 1767
Clio fallauxi (Kittl, 1886)
1886 Balantium Fallauxi Kittl, p. 62, pl. 2, figs. 23‒26.
1984 Balantium fallauxi Kittl, 1886 – Janssen, p. 65.
1993 Clio fallauxi (Kittl, 1886) – Janssen & Zorn, p. 195, pl. 8,
figs. 1‒9, pl. 9, figs. 1‒3.
1999 Clio fallauxi (Kittl, 1886) – Zorn, p. 730, pl. 2, figs. 3‒5.
Material examined: Five casts with preserved ornament
(inner shell ornament) on yellow marls at the Vejalnica loca-
lity (temporary inventory numbers CNHM V-CF 1 to 5). The
measured elements are shown on Fig. 7.
Description: Elongated triangular shell sculptured with
transversal ribs overrun by multiple riblets. The ribs are flat-
tened, but in the lateral parts of the shell they are flexuous.
Measurements of specimens are shown in Table 3.
Inv. No.
H (in mm)
W (in mm)
H/W-RATIO
α
A
1
(in mm)
A
2
(in mm)
A
1
/A
2
- RATIO
MBa-LV 1
0.255
0.288
0.885
≤119°
0.21
0.105
2
MBa-LV 2
0.317
0.399
0.794
≤120°
0.279
0.168
1.661
MBa-LV 3
0.238
0.3
0.793
≤133°
0.196
0.098
2
MBa-LV 4
0.301
0.409
0.736
≤ 93°
0.257
0.112
2.295
MBa-LV 5
0.267
0.337
0.792
≤118°
0.219
0.095
2.305
MBa-LV 6
0.265
0.343
0.773
≤ 98°
0.196
0.098
2
MBa-LV 7
0.284
0.341
0.833
≤116°
0.223
0.123
1.813
MBa-LV 8
0.367
0.437
0.84
≤107°
0.278
0.122
2.279
MBa-LV 9
0.282
0.328
0.86
≤102°
0.179
0.089
2.011
MBa-LV 10
0.211
0.262
0.805
≤120°
0.152
0.076
2
MBa-LV 11
0.24
0.273
0.879
≤113°
0.156
0.095
1.642
MBa-LV 12
0.206
0.265
0.777
≤ 97°
0.145
0.089
1.629
MBa-LV 13
0.272
0.358
0.76
≤106°
0.19
0.078
2.436
MBa-LV 14
0.257
0.282
0.911
≤119°
0.19
0.101
1.881
MBa-LV 15
0.259
0.294
0.881
≤107°
0.179
0.101
1.772
MBa-LV 16
0.352
0.399
0.882
≤105°
0.246
0.156
1.577
MBa-LV 17
0.257
0.29
0.886
≤124°
0.195
0.116
1.681
MBa-LV 18
0.382
0.426
0.897
≤122°
0.267
0.124
2.153
MBa-LV 19
0.331
0.368
0.899
≤102°
0.246
0.101
2.436
MBa-LV 20
0.185
0.257
0.72
≤158°
0.143
0.067
2.134
MBa-LV 21
0.243
0.331
0.734
≤154°
0.21
0.095
2.211
MBb-LV 1
0.262
0.334
0.784
≤126°
0.214
0.129
1.659
MBb-LV 2
0.278
0.349
0.797
≤114°
0.223
0.134
1.664
MBb-LV 3
0.239
0.286
0.836
≤110°
0.193
0.1
1.93
MBb-LV 4
0.27
0.303
0.891
≤111°
0.204
0.107
1.907
MBb-LV 5
0.267
0.326
0.819
≤125°
0.2
0.1
2
MBb-LV 6
0.249
0.313
0.796
≤ 98°
0.184
0.112
1.643
MBb-LV 7
0.23
0.294
0.782
≤126°
0.179
0.101
1.772
MBb-LV 8
0.238
0.279
0.853
≤ 92°
0.171
0.1
1.71
MBb-LV 9
0.269
0.319
0.843
≤132°
0.19
0.112
1.696
MBb-LV 10
0.259
0.302
0.858
≤ 95°
0.134
0.101
1.327
MBb-LV 11
0.282
0.355
0.794
≤119°
0.2
0.114
1.754
MBb-LV 12
0.349
0.375
0.931
≤ 88°
0.229
0.114
2.009
MBb-LV 13
0.401
0.445
0.901
≤101°
0.271
0.171
1.585
MBb-LV 14
0.249
0.271
0.919
≤120°
0.184
0.112
1.643
Table 1: Measurements of Limacina valvatina (Reuss, 1867) from the Marija Bistrica locality. Abbreviations: Inv. No. — inventory number;
H — shell height; W — shell width; H/W-ratio — shell height and width ratio; α — apical angle; A
1
and A
2
— aperture diameters;
A
1
/A
2
-ratio — aperture diameters (A
1
and A
2
) ratio.
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0
0.05
0.1
0.15
0.2
0.3
0.25
0.35
0.4
0.45
0.5
0
0.05
0.1
0.15
0.2
0.25
0.3
0.35
0.4
0.45
Shell height, H (in mm)
Shell width, W (in mm)
A
70-75
75-80
80-85
85-90
90-95
3
11
7
10
4
B
H/W-ratio
0
1
2
3
4
5
6
7
8
130-135
135-140
155-160
160-165
165-170
170-175
175-180
180-185
185-190
190-195
195-200
200-205
205-210
210-215
215-220
220-225
225-230
230-235
235-240240-245
A /A -ratio
1
2
Number of specimens
C
Remarks: The pteropod casts here presented resemble the
species Clio fallauxi (Kittl, 1886). Kochansky (1944) listed
Clio pedemontana (Mayer, 1868) from the Medvednica Mt.
(“Central” or “Čučerje” area), but unfortunately her original
material is missing. The species C. fallauxi resembles
C. pede montana, but it differs from the latter in its larger api-
cal angle and presence of riblets (e.g., Janssen & Zorn 1993;
Zorn 1999). In Tables 3 and 4, and Figs. 8 and 9, we present
specimens of C. fallauxi from selected Paratethyan localities
(see Table 4). Based on the morphology, measurements and
comparison of the investigated Clio specimens from the
Medvednica Mt. with other records from the Central
Paratethys, we identify the sculpted casts as C. fallauxi. As
shown on Fig. 9, shell height and width fits the trend line of
other Central Paratethys specimens. The apical angle is
variable in comparison to other specimens, but that might be
a preservation artifact.
One specimen (V-CF 5, Fig. 8 D) could only be determined
as Clio cf. fallauxi due to its deviating apical angle value,
although its aperture angle is similar
to the other collected specimens,
and the ribs and riblets are present.
Occurrence: In the Central Para-
tethys the occurrence of Clio fallauxi
was documented only in the Lower
Badenian deposits (e.g., Zorn 1999;
Bohn-Havas & Zorn 2002; Selmeczi
et al. 2012). It was known from
Hungary (Bohn-Havas & Zorn 1993,
1994; Zorn 1999; Bohn-Havas et al.
2004) in NN5 nannozone after
Bohn-Havas & Zorn (1993) and
Selmeczi et al. (2012), Poland
(Bohn-Havas & Zorn 1993, 1994;
Janssen & Zorn 1993; Zorn 1999),
Czech Republic (Janssen & Zorn
1993; Zorn 1999), Romania (Bohn-
Havas & Zorn 1994; Zorn 1999),
Bulgaria (Zorn 1999; Nikolov 2010)
and possibly Slovenia (NN5 nanno-
zone after Mikuž et al. 2012).
C. fallauxi locally occurs together
with C. pedemontana (see Bohn-
Havas & Zorn 1993, 1994; Zorn
1999; Bohn-Havas et al. 2004) in
NN5 nannozone after Bohn-Havas
et al. (2004).
Fig. 4. A — Shell height and width diagram of Limacina valvatina
(Reuss, 1867) from the Marija Bistrica locality. B — Shell height/
width ratio of collected specimens with number of L. valvatina
belonging to each H/W range (70–75, 75–80, 80–85, 85–90, 90–95).
C — Aperture diameters (A
1
/A
2
) ratio of presented L. valvatina with
number of specimens from each range. Abbreviations: A
1
/A
2
-ratio —
aperture diameters A
1
and A
2
ratio; H/W-ratio — shell height and
width ratio.
Fig. 5. A–B: Limacina gramensis (Rasmussen, 1968) from the Marija Bistrica locality. A — CNHM
MB-LG1; B — CNHM MB-LG 8. C-D: Limacina sp. (? nov.) from the Marija Bistrica locality.
C — CNHM MB-L 1; D — CNHM MB-L 2. Scale bars 100 μm.
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0
0.05
0.1
0.15
0.2
0.25
0.3
0.35
0.4
Shell height, H (in mm)
0.225
0.23
0.235
0.24
0.245
0.25
0.255
0.26
0.265
0.27
0.275
Shell width, W (in mm)
A
0.105-0.1
10
0.1
10-0.1
15
0.1
15-0.120
0.120-0.125
0.125-0.130
0.130-0.135
0.135-0.140
0.140-0.145
0.145-0.150
0.150-0.155
0.155-0.160
1
1
1
2
1
0
0
0
0
0
0
H/W-ratio
B
Family: Cavoliniidae Gray, 1850 (1815)
(= Hyalaeidae Rafinesque, 1815)
Genus: Vaginella Daudin, 1800
Type species: Vaginella depressa Daudin, 1800
Vaginella austriaca Kittl, 1886
1886 Vaginella austriaca Kittl, p. 54, pl. 2, figs. 8‒12.
1981 Vaginella austriaca Kittl, 1886 – Krach, p. 124, pl. 1, figs. 15−18,
20; pl. 2, figs. 1−3, 21−24; pl. 4, fig. 2.
1984 Vaginella austriaca Kittl, 1886 – Janssen, p. 73, pl. 4, figs. 1‒8.
1991 Vaginella austriaca Kittl, 1886 – Zorn, p. 120, pl. 6, figs. 1‒6,
pl. 7, figs. 1‒9, pl. 12, figs. 4, 5, pl. 14, figs. 1‒8, pl. 16, figs. 1‒4.
1993 Vaginella austriaca Kittl, 1886 – Janssen & Zorn, p. 203, pl. 6,
figs. 8‒15, pl. 10, figs. 1‒5, pl. 11, figs. 1‒6.
1999 Vaginella austriaca Kittl, 1886 – Zorn, p. 732, pl. 4, figs. 1‒3,
pl. 5, figs. 1, 2.
Material examined: Casts and imprints found in yellow
marls at the Vejalnica locality (temporary inventory numbers
CNHM V-VA 1 to 17) (Fig. 10, Table 5).
Description: Lance-shaped specimens with smooth shell
surface, widest in the aperture area. Absence or presence of
lateral carinae could not be determined since the material is
preserved as casts and imprints. Pre-apertural constriction is
not visible. The middle part of two shell casts is widened and
these specimens are identified as Vaginella cf. austriaca, due
to the lack of diagnostic elements for precise determination,
and are not included in statistical analyses presented in this
paper. Dimensions of the studied specimens (Vejalnica loca-
lity) are shown in Table 5. Measurements of the specimens
presented in this paper are shown in Fig. 11, together with
other Vaginella austriaca specimens from the selected
Paratethyan localities (Table 6).
Remarks: Different species of the genus Vaginella Daudin,
1800 such as V. depressa Daudin, 1800, V. austriaca Kittl,
1886, V. acutissima Audenino, 1897 and V. lapugyensis Kittl,
1886 are similarly shaped, developing more elongated shells
during their evolution. Therefore it can be difficult to deter-
mine them to species level, particularly in poorly preserved
specimens with the apical part with protoconch missing, and
damaged apertural part. Beside general shape differences, the
apical angle is used as a distinguishing criterion, which can
also be misleading due to the curvature of shell sidelines.
V. depressa has the widest apical angle of ≥40°, V. austriaca
c. ≥20°, V. acutissima c. 15–18° and V. lapugyensis c. 8–13°
(Cahuzac & Janssen 2010, p. 100). Apical angle comparison
of V. austriaca from the Vejalnica locality and selected
avai lable Paratethyan specimens is shown in Fig. 12.
Occurrence: Vaginella austriaca from the Medvednica Mt.
was recorded by Gorjanović-Kramberger (1908), Kochansky
(1944), Basch (1983b) and Avanić et al. (1995). In the Central
Paratethys V. austriaca is known from the Karpatian and
Badenian deposits in Austria (Janssen 1984; Bohn-Havas &
Zorn 1993, 1994; Janssen & Zorn 1993; Zorn 1991, 1999),
Poland (Lower and Middle Badenian) (Zorn 1991, 1999;
Bohn-Havas & Zorn 1993, 1994; Janssen & Zorn 1993),
Hungary (Lower Badenian) (Zorn 1991; Bohn-Havas & Zorn
1993, 1994; Bohn-Havas et al. 2004) in NN5 nannozone after
Bohn-Havas & Zorn (1993), Bohn-Havas et al. (2004) and
Selmeczi et al. (2012), Czech Republic (Karpatian and Lower
Badenian) (Zorn 1991, 1999), Slovenia (Mikuž et al. 2012 and
references therein), Serbia (Stevanović 1974), Romania
(Lower Badenian) (Zorn 1991, 1999; Janssen & Zorn 1993;
Bohn-Havas & Zorn 1994) and Bulgaria (Badenian) (Zorn
1999; Nikolov 2010). Outside the Paratethys, Vaginella
austriaca is known from Middle Miocene deposits in the
Mediterranean area (Zorn 1991, 1999 and references therein;
Bohn-Havas & Zorn 2002; Janssen & Little 2010; Janssen
2012), Miocene deposits in the North Sea Basin (Zorn 1991,
Table 2: Measurements of Limacina gramensis (Rasmussen, 1968)
from the Marija Bistrica locality. * Measurements of Limacina sp.
(? nov.) from the same locality. Abbreviations: Inv. No. — inventory
number; H — shell height; W — shell width; H/W-ratio — shell
height and width ratio; α — apical angle.
Fig. 6. Limacina gramensis (Rasmussen, 1968) from the Marija
Bistrica locality. A — Shell height and width relationship. B — Shell
height/width-ratio with marked number of collected specimens
belonging to each range (after Table 2). Abbreviation: H/W-ratio —
shell height and width ratio.
Inv. No.
H (in mm)
W (in mm)
H/W-RATIO
α
MB-LG 1
0.394
0.246
1.602
≤ 44°
MB-LG 2
0.259
0.228
1.136
≤ 80°
MB-LG 3
0.281
0.23
1.222
≤ 70°
MB-LG 4
0.31
0.254
1.22
≤ 61°
MB-LG 5
0.288
0.243
1.185
≤ 53°
MB-LG 6
0.294
0.271
1.085
≤ 46°
MB-L 1 *
0.344
0.25
1.376
≤ 62°
MB-L 2 *
0.292
0.232
1.259
≤ 65°
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1999 and references therein; Bohn-Havas & Zorn 2002;
Cahuzac & Janssen 2010; Janssen & Little 2010; Janssen
2012), and the Aquitaine Basin (Cahuzac & Janssen 2010;
Janssen & Little 2010; Janssen 2012).
Regardless of the poor preservation of Vaginella specimens
from the Medvednica Mt. the authors consider casts to belong
to Vaginella austriaca. The apical angle can be used as
a reliable determination criterion characteristic for Vaginella
species only in very well-preserved specimens (Janssen &
Zorn 1993; Cahuzac & Janssen 2010). Since the specimens
presented here are preserved as casts, measurements of the
apical angle could be uncertain and unreliable for the specific
determination (e.g., Fig. 12). Therefore the authors based their
identification on the morphology, previously known data and
comparison with available data from the neighbouring area.
As seen in Fig. 11, studied specimens fit the trend line of the
shell width and height diagram of the compared Paratethyan
material (see Table 6), which confirmed our determination.
Calcareous nannoplankton
Calcareous nannoplankton is found at both of the investi-
gated localities on the Medvednica Mt., as shown in Table 7
and Figs. 13 and 14. Table 7 shows comparison with available
data covering pteropod and nannoplankton assemblages (after
Chira et al. 2000; Suciu et al. 2005; Vulc & Silye 2005; Ćorić
et al. 2009; Bartol 2009; Mikuž et al. 2012 and this paper).
Different nannoplankton assemblages of high stratigraphic
range are here recorded, and the time span of the deteremined
taxa is from NN4 to NN6. The age of deposits cropping out at
Vejal nica has been determined by the occurrence of
Sphenolithus hetero morphus as the NN5 Zone. Nanno plankton
W
H
t
W
α
α
1
α
H
t
A
B
Fig. 7. Measured elements of A — Clio fallauxi (Kittl, 1886) and
B — Vaginella austriaca Kittl, 1886 (after Zorn, 1991). Abbreviations:
H
t
— height of teleoconch; W — shell width; α — apical angle;
α
1
— aperture angle.
Table 3: Measurements of Clio fallauxi (Kittl, 1886) from the Vejalnica locality (Medvednica Mt., northern Croatia). * Clio cf. fallauxi.
Abbreviations: Inv. No. — inventory number;H
t
— height of teleoconch; W — shell width; α — apical angle; α
1
— aperture angle.
Inv.No.
H
t
(in mm)
W (in mm)
α
α
1
1st ORDER RIBS
RIBS WIDTH (in mm)
CONVEX / CONCAV
RATIO
MACRORIBS
MICRORIBS
convex
concav
V-CF 1
10
7.3
≤ 74°
≤ 28°
≥11
0.54
0.27
2
V-CF 2
5.78
5.22
not saved
≤ 33°
≥9
0.33
0.22
1.5
V-CF 3
6.81
5.42
≤ 68°
≤ 32°
≥9
0.42
0.28
1.5
V-CF 4
6.5
5.63
≤ 72°
not saved
≥5
0.25
0.13
1.92
V-CF 5*
10.15
7.12
≤ 58°
≤ 37°
≥9
0.45
0.3
1.5
Table 4: Measurements of Clio fallauxi (Kittl, 1886) from the Badenian deposits of the Central Paratethys (after literature data). * Note that the
measures are obtained from published photos, not the original specimens. ** Measurements from cited papers. *** Material mentioned by
Krach (1981) derived from boreholes. Abbreviations: H
t
— height of teleoconch; W — shell width; α — apical angle.
REFERENCES
SPECIMEN
LOCALITY
H
t
(in mm) *
W (in mm) *
α *
1
st
ORDER RIBS *
Kittl (1868)
Figs. 23–25
Peterswald, Czech Republic
13.7 **
10 **
Janssen & Zorn (1993)
Pl. 8, Fig. 1
Dębowiec, Poland***
13.14
9.71
39° – 49° **
≥ 12
Pl. 8, Fig. 2
10.29
8.29
≥ 12
Pl. 8, Fig. 3
Simoradz, Poland***
6.6
7.6
≥ 7
Pl. 8, Fig. 4
Roczyny, Poland***
7.82
6.55
≥ 6
Pl. 8, Fig. 5
8.6
7.2
?
Pl. 8, Figs. 6
Dębowiec, Poland***
13.71
8.57
≥ 12
Pl. 8, Figs. 8
Roczyny, Poland***
13.14
8.57
≥ 12
Pl. 9, Fig. 1
Dombrau, Czech Republic
7.27
6.91
45° **
≥ 7
Bohn-Havas & Zorn (1994)
Pl. 3, Fig. 13
Sopron well, Hungary
6.38
4.75
≤ 38°
≥ 7
Zorn (1999)
Pl. 2, Fig. 3
Dražovice, Czech Republic
4.5
2.63
≤48°
≥ 13
4.75
2.25
≤37°
≥ 15
4.88
2.38
≤36°
≥ 7
4
2.38
≤36°
?
5.63
3.38
≤40°
≥ 15
Pl. 2, Fig. 4
11.54
6
≤41°
≥ 13
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BOŠNJAK, SREMAC, VRSALJKO, AŠČIĆ and BOSAK
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assemblages from the Marija
Bistrica locality indicate younger
horizons, on the transition from
the NN5 to NN6 Zone. Coccoliths
are abundant and comprise some
reworked specimens.
Discussion
Pteropod ecology
Pteropods show horizontal dis-
tribution along climate zones,
depending on temperature, sali-
nity, food, oxygen, water depth,
ocean currents etc., with tempera-
ture as the main factor of their
distribution. They are limited to
narrow salinity ranges and can
tole rate slight salinity deviations,
e.g., according to Ivanova (1983)
up to 28–30 ‰, 35 to 36 ‰ (Herman
1998) and 39 ‰ (Velcescu 1997).
Pteropods also have a limited tole-
rance to temperature change, with
exceptions, for example, of recent
representatives of the genus
Creseis Rang, 1828 and Limacina
trochiformis (d’Orbigny, 1834)
(see Herman 1998). Recent ptero-
pods are distributed in worldwide
seas (e.g., Hunt et al. 2010). Most species live in tropical and
subtropical regions, however, they are also represented in
polar regions (e.g., Velcescu 1997; Herman 1998; Janssen &
Peijnenburg 2014).
Pteropod species diversity in tropical seas is high and they
are concentrated at greater depths than temperate and high-
latitude species (Bednaršek et al. 2012b and references
therein), and in colder and polar waters pteropods are particu-
larly abundant and represented by two species, Limacina
helicina (Phipps, 1774) and Limacina retroversa (Fleming,
1823) (e.g., Hunt et al. 2010; Lischka & Riebesell 2012).
Limacina helicina is a polar pteropod species, a main compo-
nent of the polar zooplankton (Hunt et al. 2010; Lischka &
Riebesell 2012). Limacina retroversa is a boreal-temperate
species of subpolar and transitional waters of the North
Atlantic (Lischka & Riebesell 2012).
Modern pteropods nowadays are widely used as first indica-
tors of ocean acidification effects on marine communities, due
to their aragonitic shell (e.g., Hunt et al. 2010; Bednaršek et al.
2012a; Burridge et al. 2015). The process of ocean acidifica-
tion (reducing pH value) and reduction with a lowering of the
CaCO
3
saturation as a result affects marine organisms building
their skeletons from aragonite, a more soluble form of CaCO
3
than calcite. The consequences of the ocean acidification
process are expected to be visible first in the polar region, due
to the higher solubility of CO
2
in colder waters, which impacts
organisms with calcified shells, particularly pteropods (e.g.,
Lischka & Riebesell 2012). Due to the higher aragonite under-
saturation, pteropods can migrate in the aragonite saturated
waters changing their depth strata (Lischka & Riebesell 2012),
Fig. 9. Comparison of shell height and width of Clio fallauxi
(Kittl, 1886) specimens described in previously published papers
(references in Table 4 — white) and from the Medvednica Mt. (this
study — black).
0
2
4
6
8
10
12
14
16
0
2
4
6
8
10
12
Height of teleoconch, H (in mm)
t
Shell width, W (in mm)
Fig. 8. Clio fallauxi (Kittl, 1886) from the Vejalnica locality. A — CNHM V-CF 1; B — CNHM
V-CF 2; C — CNHM V-CF 3; D — Clio cf. fallauxi, CNHM V-CF 5. Scale bars 5 mm.
339
THE MIOCENE “PTEROPOD EVENT” IN THE SW PART OF THE CENTRAL PARATETHYS
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which means that their migration is not influenced only by
surface water temperature (e.g, research presented in Oliveira-
Koblitz & Larrazábal 2014).
“Pteropod events” in the Para
tethys
Pteropods
inhabited
the
Paratethys Sea from its early days
(Rögl 1999), but their abundance
varied in different horizons, with
two peaks during the Middle
Miocene (e.g., Bohn-Havas et al.
2003). Vaginella austriaca Kittl,
1886 is the most abundant ptero-
pod species in the Middle Miocene
of this area (Janssen & Zorn 1993;
Bohn-Havas & Zorn 1993, 1994,
2002; Bohn-Havas et al. 2003,
2004). It first appeared in the
Karpatian NN4 nannozone and
only a small number can be found
in the Early Badenian, about
14.9 Ma ago (Bohn-Havas et al.
2003). According to the magneto-
stratigraphic data from Hungary,
mixed pteropod fauna, compri sing
Clio pedemontana (Mayer, 1868),
Clio fallauxi (Kittl, 1886) and
Limacina valvatina (Reuss, 1867)
appeared in that area 14.5 Ma ago,
in the Badenian NN5 nannozone,
marking the general bloom of
plan ktic molluscs, which lasted till
about 14.2 Ma ago (Bohn-Havas et
al. 2003; Selmeczi et al. 2012).
Mass occurrences of limacinids (Limacina or, previously,
“Spiratella” horizon), were recor ded in the Upper Badenian
deposits throughout the Central Paratethys, except in Hungary
(e.g., Rögl 1998, 1999; Bohn-Havas et al. 2004; Kováč et al.
2007; Śliwiński et al. 2011). This horizon comprises the
species L. valvatina and the Upper Badenian index fossil,
L. gramensis. In Poland this horizon is positioned within
the “Pecten Beds”, in the uppermost Badenian NN6 zone,
aged 13.06 Ma (Śliwiński et al. 2011). Limacinid bloom coin-
cides with the Middle Miocene climatic transition and cooling
event, when CO
2
pressure was in dec line (e.g., Zhang et al.
2013).
Recent and fossil limacinids are usually abundant in cold,
even polar waters and represented by a small number of spe-
cies. Mass mortality of Lima cina from the older, Oligo cene
deposits was attributed to a short-term decrease in surface
water salinity, or rise of H
2
S and oxygen depleted environ-
ments (Rögl 1998; Báldi 2006 and refe rences therein;
Studencka et al. 2016 and references therein). Similar environ-
mental conditions could have triggered the Late Badenian
limacinid crisis (e.g., Báldi 2006 and refe rences therein), or
we can explain this event with the influx of colder ocean
waters into the Paratethys during the Middle Miocene (e.g.,
Kováč et al. 2007).
Fig. 10. Vaginella austriaca Kittl, 1886 from the Vejalnica locality. A — CNHM V-VA 2;
B — CNHM V-VA 7; C — CNHM V-VA 11; D — CNHM V-VA 19. Scale bars 5 mm.
Inv. No.
H
t
(in mm)
W (in mm)
α
V-VA 1
4.38
1.69
≤ 27°
V-VA 2
6.88
2.38
≤ 19°
V-VA 3
3.62
1.16
≤ 28°
V-VA 4
3.5
1.5
≤ 21°
V-VA 5
3.5
1.5
≤ 24°
V-VA 6
7.43
2.71
≤ 20°
V-VA 7
4.56
1.62
≤ 20°
V-VA 8
2.5
1.05
≤ 26°
V-VA 9
4.36
1.54
≤ 20°
V-VA 10
4.29
1.43
≤ 21°
V-VA 11
4.19
1.63
≤15°
V-VA 12
4.07
1.05
≤14°
V-VA 13
4.65
2.21
≤ 22°
V-VA 14
4.25
1.5
≤18°
V-VA 15
4.23
1.41
≤13°
V-VA 16
2.56
1.28
≤ 24°
V-VA 17
5
2.1
≤ 28°
Table 5: Measurements of Vaginella austriaca Kittl, 1886 from the
Vejalnica locality (Medvednica Mt., northern Croatia). Abbreviations:
Inv. No. — inventory number; H
t
— height of teleoconch; W — shell
width; α — apical angle.
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Table 6: Measurements of Vaginella austriaca Kittl, 1886 from the Badenian deposits of the Central Paratethys (after literature data). * Note
that the measures are obtained from published photos, not the original specimens. ** Lectotype. Abbreviations: H
t
— height of teleoconch;
W — shell width; α — apical angle.
Fig. 11. Comparison of shell height and width of Vaginella austriaca
Kittl, 1886 from the Medvednica Mt. (black triangels; after Table 5)
and the comparative material from the Central Paratethys (white
triangels; after Table 6).
Fig. 12. Apical angle values of Vaginella austriaca Kittl, 1886 from
the Vejalnica locality, and comparative material from the Central
Paratethys (after Table 6). Numbers above the columns mark number
of specimens within three ranges of measured apical angles (8°–14°,
15°–19°, 20°– 40°).
REFERENCES
SPECIMEN
LOCALITY
H
t
(in mm) *
W (in mm) *
α *
Janssen (1984)
Pl. 4, Fig. 1 **
Baden, Austria
7.67
1.83
≤ 20°
Pl. 4, Fig. 3
6.33
1.58
≤ 23°
Pl. 4, Fig. 4
6.92
1.83
≤ 23°
Pl. 4, Fig. 7
Lapugy, Rumania
4.08
1
≤ 22°
Pl. 4, Fig. 8
4.83
1.17
≤ 22°
Zorn (1991)
Pl. 6, Fig. 3
Bad Vöslau, Austria
6.37
1.95
≤ 27°
Pl. 6, Fig. 4
7
2.09
≤ 25°
Pl. 6, Fig. 5
6.91
2.09
≤ 29°
Pl. 6, Fig. 6
7.09
2.18
≤ 30°
Pl. 7, Fig. 2
7.74
2.43
≤ 23°
Pl. 7, Fig. 3
7.17
1.83
≤ 18°
Pl. 14, Fig. 1
Baden, Austria
7.13
1.75
≤ 21°
Pl. 14, Fig. 2
6.25
1.75
≤ 24°
Pl. 14, Fig. 3
6
1.5
≤ 24°
Pl. 14, Fig. 4
6.5
1.88
≤ 21°
Pl. 14, Fig. 5
Bad Vöslau, Austria
6.19
1.5
≤ 18°
Pl. 14, Fig. 6
Sooss, Austria
4.44
1.38
≤ 18°
Janssen & Zorn (1993)
Pl. 6, Fig. 8
Korytnica, Poland
5.67
1.58
≤ 24°
Pl. 6, Fig. 9
5.83
1.67
≤ 15°
Pl. 6, Fig. 12
4.83
1.5
≤ 29°
Pl. 10, Fig. 1
Brzeszcze, Poland
7.63
3
≤ 28°
Pl. 10, Fig. 2
Miçdzyrzecze, Poland
6.63
2.75
≤ 33°
Pl. 10, Fig. 3
Łapczyca, Poland
7.38
2.38
≤ 34°
Pl. 10, Fig. 4
Brzeszcze, Poland
6.38
2.38
≤ 25°
Pl. 10, Fig. 5
6.63
2
≤ 30°
Pl. 11, Fig. 1
Poremba, Czech Republic
7.94
3.25
≤ 24°
Pl. 11, Fig. 2
Grudna Dolna, Poland
8.31
3.5
≤ 31°
Pl. 11, Fig. 3
7.88
3.63
≤ 37°
Pl. 11, Fig. 4
5
2.75
≤ 38°
Pl. 11, Fig. 5
10.94
3.88
≤ 34°
Pl. 11, Fig. 6
11.81
3.75
≤ 27°
Bohn-Havas & Zorn (1994)
Pl. 2, Fig. 5
Vöslau, Austria
6.2
1.67
≤ 24°
Pl. 3, Fig. 3
6.55
2.55
≤ 15°
Pl. 3, Figs. 4
7.54
2.45
≤ 21°
Zorn (1999)
Pl. 4, Fig. 1
Tučapy, Czech Republic
9.6
3.6
≤ 26°
Pl. 4, Fig. 2
7.9
3
≤ 38°
Pl. 4, Fig. 3a
8.33
3.17
≤ 24°
Pl. 4, Fig. 3b
8.33
3.33
≤ 27°
0
0.5
1
1.5
2
2.5
3
3.5
4
4.5
0
2
4
6
8
10
12
14
Shell height, H (in mm)
Shell width, W (in mm)
2
4
3
14
12
44
8-14
15-19
20-40
Medvednica Mt.
Central Paratethys
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Pteropods from Mt. Medvednica
During this research, pteropods were collected from two
different areas and associated with different biota, indicating
two different stratigraphic horizons (Table 8).
Yellow marls from the Vejalnica locality belong to the
“Central” Miocene belt (sensu Kochansky 1944). Deposits
from this area yielded the first record of pteropods (Vaginella
austriaca Kittl, 1886) in the Medvednica Mt. (Gorjanović-
Kramberger 1908). This finding was later supported by
a record of Clio pedemontana (Mayer, 1868) mentioned by
Kochansky (1944) and Basch (1983b, p. 29). Unfortunately,
the original specimens from these studies are missing. Species
C. fallauxi, collected during this study, occurs in the same area
as the above mentioned C. pedemontana. These records sup-
port the existence of marine environments on the Medvednica
Mt. prior to the Late Badenian, and are in accordance with the
pteropod bloom positioned by Bohn-Havas et al. (2003)
between the 14.5 and 14.2 Ma (Table 9).
The age of the yellow marls from the “Central” Miocene
belt, previously compared to the Austrian “Schlier”, was addi-
tionally estimated on the basis of calcareous nannoplankton,
pointing to the probable Badenian NN5 nannozone, although
the time span of the most determined taxa ranges from NN4
to NN6.
Pteropods in the northeastern part of the Medvednica Mt.
(“Zelina” development) were found in grey marls exposed
near Marija Bistrica, and comprise only limacinids (L. valva-
tina, L. gramensis, Limacina sp. ?). Limacina gramensis was
present in the Central Paratethys only during the Late Badenian
(Table 10), which points to the younger transgressive cycle
(probably NN6 Zone) than the one visible at Vejalnica
locality.
Due to the small size and poor preservation potential of
pteropods, it is possible that during the previous investigations
pteropods were not recorded, or remained unpublished. In
addition, some older fossil collections are not available, so
part of the information may be missing.
“Migration routes”
Despite the fact that pteropods are not active swimmers and
their distribution depends on ocean currents and available sea-
ways, we discuss here about their possible “migration routes”
based on their recorded geographical distribution in the
Paratethys during the Badenian. Open seaways are connected
with maximum flooding, tectonic events and climatic condi-
tions and it is not easy to position them in time and space. New
pteropod findings from this study contribute to the palaeobio-
geographic reconstructions, indicating the possible seaway
connections during the different Miocene transgressive cycles,
but they do not provide clear evidence for the “migration
routes”. We hope that our further research will give us more
insight on this subject.
Available data on the geographical distribution of C. fallauxi,
C. pedemontana, V. austriaca, L. valvatina and L. gramensis
in the Paratethys during the Miocene are shown on Figs. 15
and 16. There are additional pteropod records (Vaginella
SPECIES
REGION/LOCALITY
CENTRAL PARATETHYS
EASTERN PARATETHYS
NE SLOVENIA
NW CROATIA - Medvednica Mt.
ROMANIA
Central
Eastern
Čučerje
Vejalnica*
M. Bistrica*
Cluj-Napoca
Cepari
Calcidiscus permacintyrei Theodoridis
C. tropicus (Kamptner)
Coccolithus miopelagicus Bukry
C. pelagicus (Wallich) Schiller
Cycliargolithus abisectus Muller (Wise)
Discoaster formosus Martini & Worsley
Helicosphaera carteri (Wallich) Kamptner
H. intermedia Martini
H. perch-nielseniae (Haq) Jafar & Martini
Pontosphaera multipora (Kamptner) Roth
P. plana (Bramlette & Sullivan) Haq
Reticulofenestra bisecta (Hay, Mohler & Wade) Roth
R. dictycoda (Deflandre) Stradner
R. minuta Roth
R. perplexa (Burns) Wise
R. pseudoumbilicus (Gartner) Gartner
Sphenolithus heteromorphus Deflandre
S. moriformis (Bronniman & Stradner) Bramlette & Wilcoxon
Umbilicosphaera rotula (Kamptner) Varol
Table 7: Comparison of the Medvednica Mt. Badenian nannoplankton findings with available data on pteropod and calcareous nannoplankton
assemblages from Slovenia (Bartol 2009; Mikuž et al. 2012), Croatia (Čučerje locality after Ćorić et al. 2009, * this research) and Romania
(Cepari after Vulc & Silye 2005; Cluj-Napoca after Chira et al. 2000, Suciu et al. 2005).
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Fig. 13. Vejalnica locality. A — Helicosphaera carteri (Wallich, 1877) Kamptner, 1954; B — H. carteri (Wallich, 1877) Kamptner, 1954;
C — Pontosphaera multipora (Kamptner, 1948 ex Deflandre in Deflandre and Fert, 1954) Roth, 1970; D — Coccolithus miopelagicus Bukry,
1971; E — C. miopelagicus Bukry, 1971; F — Sphenolithus heteromorphus Deflandre, 1953; G — Discoaster formosus Martini and Worsley,
1971; H — Cyclicargolithus abisectus (Muller, 1970) Wise, 1973; I — Coccolithus pelagicus (Wallich, 1877) Schiller, 1930;
J — Reticulofenestra perplexa (Burns, 1975) Wise, 1983; K — Reticulofenestra minuta Roth, 1970; L — Reticulofenestra pseudoumbilicus
(Gartner, 1967) Gartner, 1969. Figs. B, E, G PPL, others XPL. Scale bars 5 μm.
343
THE MIOCENE “PTEROPOD EVENT” IN THE SW PART OF THE CENTRAL PARATETHYS
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Fig. 14. Marija Bistrica locality. A — Pontosphaera plana (Bramlette and Sullivan, 1961) Haq, 1971; B — Sphenolithus moriformis
(Bronnimann and Stradner, 1960) Bramlette and Wilcoxon, 1967; C — Reticulofenestra dictyoda (Deflandre in Deflandre and Fert, 1954)
Stradner in Stradner and Edwards, 1968; D — Reticulofenestra bisecta (Hay, Mohler and Wade, 1966) Roth, 1970; E — Helicosphaera inter-
media Martini, 1965; F — Reticulofenestra perplexa (Burns, 1975) Wise, 1983; G — Helicosphaera perch-nielseniae (Haq, 1971) Jafar and
Martini, 1975; H — Helicosphaera carteri (Wallich, 1877) Kamptner, 1954; I — Umbilicosphaera rotula (Kamptner, 1956) Varol, 1982;
J — Reticulofenestra pseudoumbilicus (Gartner, 1967) Gartner, 1969; K — Calcidiscus premacintyrei Theodoridis, 1984; L — Calcidiscus
tropicus (Kamptner, 1956) Varol 1989 sensu Gartner, 1992. Figs. G, H, I and K PPL, others XPL. Scale bars 5 μm.
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austriaca and “Spiratella sp.”) from the
eastern part of northern Croatia (Magaš
1987; Pikija 1987) and vicinity of Belgrade,
Serbia (Stevanović 1974), but we did not
include them in Figures 15 and 16, since
we do not know the exact stratigraphic
position of those findings.
In the Early-Middle Badenian (e.g., Rögl
1998; Harzhauser & Piller 2007) fauna
may have migrated in a northwestern–
southeastern direction. Mikuž et al. (2012)
described C. pedemontana and possibly
C. fallauxi in the Badenian NN5 nannozone
deposits of Slovenia, and indicated the
existence of a marine connection between
the Central Paratethys and the Medi-
terranean during the Early Badenian trans-
gression (“Transtethyan Trench Corridor”,
e.g., Rögl 1998). Absence of C. fallauxi in
the Eastern Paratethys may indicate the
existence of a palaeogeographical barrier
between the Central and Eastern Paratethys
during that period, as described in
Studencka et al. (1998). Rögl (1998) dis-
cussed an open marine connection between
the Central and Eastern Paratethys, and the
existence of a southern marine connection
to the Mediterranean and Central Paratethys
through the area between the Black Sea and
the Pontids.
Recent studies show different interpre-
tations of the Badenian subdivision, the age
of the NN5 zone, and number and timing of
Badenian marine transgressions in the
Paratethys (Harzhauser & Piller 2007;
Piller et al. 2007; Kováč et al. 2007; Rögl et
al. 2007; Kováč et al. 2008; Pezelj et al.
2013; Bartol et al. 2014; Hohenegger et
al. 2014). Therefore, the stratigraphic range
of Clio fallauxi should be extended to
the lower part of the Middle Badenian
(Badenian subdivision sensu Papp & Cicha
1968 and Piller et al. 2007). Considering
the appearance of C. fallauxi in the NN5
nannozone e.g., Mikuž et al. (2012) and
Selmeczi et al. (2012) (Early Badenian
sensu Kováč et al. 2007, i.e. Early-Middle
Badenian sensu Piller et al. 2007), our new
records, together with previous research
from Croatia (Kochansky 1944) contribute
to the possible suggested northwestern-
southeastern fauna migration direction in
the Paratethys.
The existence of the seaway between the
Mediterranean and the Central Paratethys
(“Transtethyan Trench Corridor”) during
Table 8: List of the collected fossil fauna presented in this paper. Abbreviations:
V — Vejalnica locality; MB — Marija Bistrica locality.
Species
Locality
V
MB
Pteropoda
Limacina valvatina (Reuss)
Limacina gramensis (Rasmussen)
Clio fallauxi (Kittl)
Vaginella austriaca Kittl
?
Coccolithales
Calcidiscus premacintyrei Theodoridis
C. tropicus (Kamptner)
Coccolithus miopelagicus Bukry
C. pelagicus (Wallich) Schiller
Cyclicargolithus abisectus Muller (Wise)
Discoaster formosus Martini & Worsley
Helicosphaera carteri (Wallich) Kamptner
H. intermedia Martini
H. perch-nielseniae (Haq) Jafar & Martini
Pontosphaera multipora (Kamptner) Roth
P. plana (Bramlette & Sullivan) Haq
Reticulofenestra bisecta (Hay, Mohler & Wade) Roth
R. dictyoda (Deflandre) Stradner
R. minuta Roth
R. perplexa (Burns) Wise
R. pseudoumbilicus (Gartner) Gartner
Sphenolithus heteromorphus Deflandre
S. moriformis (Bronniman & Stradner) Bramlette & Wilcoxon
Umbilicosphaera rotula (Kamptner) Varol
Other
Foraminifera
Ostracods
Bivalves
Benthic gastropods
Scaphopods
Bryozoans
Fish remains
Table 9: Regional distribution and stratigraphic range of Clio fallauxi (Kittl, 1886),
Clio pedemontana (Mayer, 1868) and Vaginella austriaca Kittl, 1886 in the Paratethys
during the Badenian (Middle Miocene). * Badenian threefold subdivision sensu Piller
et al. (2007). Palaeogeographic map of distribution is shown on Fig. 15.
References:
1
Gorjanović-Kramberger 1908;
2
Kochansky 1944:
3
Basch 1983;
4
Janssen
1984;
5
Zorn 1991;
6
Bohn-Havas & Zorn 1993;
7
Janssen & Zorn 1993;
8
Bohn-Havas &
Zorn 1994;
9
Avanić et al. 1995;
10
Zorn 1999;
11
Bohn-Havas et al. 2011;
12
Janssen &
Little 2010;
13
Nikolov 2010;
14
Mikuž et al. 2012;
15
Selmeczi et al. 2012;
16
this paper.
References
Species
Sedimentological
basin
Country
MIOCENE
MIDDLE MIOCENE
Badenian
Lower-Middle * Upper
6; 8; 10; 11; 15
Clio fallauxi
Central Paratethys
Hungary
6; 7; 8; 10
Poland
7; 10
Czech Republic
14
Slovenia
?
16
Croatia
8; 10
Eastern Paratethys
Romania
10; 13
Bulgaria
6; 8; 12; 15
Clio pedemontana
Central Paratethys
Hungary
6; 7; 12
Poland
14
Slovenia
2; 3
Croatia
6; 12
Eastern Paratethys
Romania
13
Bulgaria
4; 6; 8; 5; 10
Vaginella austriaca
Central Paratethys
Austria
5; 6; 8; 11; 15
Hungary
5; 6; 7; 8; 10
Poland
5; 10
Czech Republic
14
Slovenia
1; 2; 9
Croatia
6; 5; 10
Eastern Paratethys
Romania
10; 13
Bulgaria
345
THE MIOCENE “PTEROPOD EVENT” IN THE SW PART OF THE CENTRAL PARATETHYS
GEOLOGICA CARPATHICA
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the Late Badenian is debated in several
papers. According to some authors (e.g.,
Rögl 1998; Harzhauser & Piller 2007;
Piller et al. 2007; Kováč et al. 2007),
the “Transtethyan Trench Corridor” was
closed during the Late Badenian, so the
migration of fauna from west-northwestern
direction would not be possible. Rögl
(1998) assumes the re-opening of the
southern Early Badenian passage. Based
on a bivalve fauna, Studencka et al. (1998)
conclude on a connection of the Central
and Eastern Paratethys and existence of
a corridor between the Paratethys and the
Eastern Mediterranean. Bartol et al. (2012,
2014) considered the connection between
the Mediterranean and the Central Para-
tethys through the “Transtethyan Trench
Corridor” during the Late Badenian, in
contrast to some previous studies (Bistričić
& Jenko 1985; Rijavec 1985; Rögl 1998;
References
Species
Sedimentological
basin
Country
MIOCENE
MIDDLE MIOCENE
Badenian
Lower-Middle * Upper
1; 2; 4
Limacina valvatina
Central Paratethys
Austria
2; 4; 6; 9
Hungary
2; 3; 4;
Poland
4; 5
Czech Republic
5
Slovakia
10
Croatia
4; 7
Eastern Paratethys
Romania
8
Bulgaria
3; 4
Ukraine
2; 3; 5
Limacina gramensis
Central Paratethys
Poland
5
Czech Republic
10
Croatia
5
Eastern Paratethys
Romania
8
Bulgaria
3; 5
Ukraine
Table 10: Regional distribution and stratigraphic range of Limacina valvatina (Reuss,
1867) and Limacina gramensis (Rasmussen, 1968) in the Paratethys area during the
Badenian (Middle Miocene). * Badenian threefold subdivision sensu Piller et al. (2007).
Palaeo geographic map of distribution is shown on Fig. 16. References:
1
Zorn 1991;
2
Bohn-Havas & Zorn 1993;
3
Janssen & Zorn 1993;
4
Bohn-Havas & Zorn 1994;
5
Zorn
1999;
6
Bohn-Havas et al. 2004;
7
Suciu et al. 2005;
8
Nikolov 2010;
9
Selmeczi et al. 2012;
10
this paper.
POLAND
UKRAINE
ROMANIA
BOSNIA AND
HERZEGOVINA
CROATIA
AUSTRIA
HUNGARY
CZECH
REPUBLIC
SLOVAKIA
SERBIA
SLOVENIA
W E S T E R N
C A R P A T H I A N S
Krakow
Lviv
Budapest
Bucuresti
Brno
E A S T E R N
A L P S
E
A
S
T
E
R
N
C
A
R
P
A
T
H
I A
N
S
S O U T H E R N
C A R P A T H I A N S
D
I N
A
R
I D
E
S
S O U T H E R N
A L P S
S t y r i a n
B a s i n
Z a l a
B a s i n
T r a n s y l v a n i a n
B a s i n
Zagreb
?
Fig. 15. Palaeogeographic distribution of Limacina valvatina (Reuss, 1867) (white circles), Clio fallauxi (Kittl, 1886) (white rectangulars),
Clio pedemontana (Mayer, 1868) (black rectangulars) and Vaginella austriaca Kittl, 1886 (black triangels) during the Early−Middle
Badenian sensu Piller et al. (2007) in the Paratethys. For references and localities see Tables 9 and 10. Palaeogeographic map after
Kováč et al. 2007.
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BOŠNJAK, SREMAC, VRSALJKO, AŠČIĆ and BOSAK
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Kováč et al. 2007). Spreading of pteropod fauna (Limacina
horizon) during the Late Badenian in the eastern part of the
Paratethys is shown on Fig. 16. L. gramensis and L. valvatina
are also recorded from northern palaeogeographical areas
(Tembrock 1989; Janssen & Zorn 1993; Janssen 1999).
Bearing that in mind, their distribution could possible point to
a connection or brief existence of a northern European passage
between the Central Paratethys and the North Sea Basin during
the Miocene.
Conclusions
Middle Miocene marls from Mt. Medvednica (northern
Croatia) comprise two different pteropod assemblages.
The pteropod fauna at the locality Vejalnica (central part of
the Medvednica Mt.) is characterized by low pteropod diver-
sity and predominance of the species Vaginella austriaca Kittl,
1886, accompanied by Clio fallauxi (Kittl, 1886). Yellow
marls additionally comprise other pelagic and deep marine
biota.
Grey Miocene marls at the locality Marija Bistrica (north-
eastern area) are highly fossiliferous and, among other fossils,
comprise limacinid pteropods: Limacina valvatina (Reuss,
1867), L. gramensis (Rasmussen, 1968) and Limacina sp.
Determined pteropod taxa, except V. austriaca and Limacina
sp. have been found in this region for the first time. Pteropods
Clio pedemontana (Mayer, 1868) and Limacina andrussowi
(Kittl, 1886) (in original: “Spirialis” or “Spiratella”) were
recorded in the same region by previous authors.
Pteropod marls at the Vejalnica locality were deposited
during the older Badenian transgressive cycle, and grey marls
with limacinids from the north-eastern part of the Medvednica
Mt. are dated to the Late Badenian.
Pteropod records from Croatia, compared with other
published data, indicate pteropod immigration into the
Paratethys from the west during the Badenian NN5 nanno-
zone. Therefore the Mediterranean connection (“Transtethyan
Trench Corridor”), proposed by previous authors, could be
a possible immigration seaway. A northern European marine
passage may have been active during the Late Badenian,
enabling the immigration of the “North Sea fauna” (including
limacinid pteropods) into the Paratethys. Such theory was pre-
sumed by several authors during the 1990s (e.g., Janssen &
Zorn 1993), but has not been confirmed, due to the lack of
fossil evidence.
POLAND
UKRAINE
ROMANIA
BOSNIA AND
HERZEGOVINA
CROATIA
AUSTRIA
HUNGARY
CZECH
REPUBLIC
SLOVAKIA
SERBIA
SLOVENIA
W E S T E R N
C A R P A T H I A N S
Krakow
Lviv
Budapest
Bucuresti
Brno
E A S T E R N
A L P S
E
A
S
T
E
R
N
C
A
R
P
A
T
H
I A
N
S
S O U T H E R N
C A R P A T H I A N S
D
I N
A
R
I D
E
S
S O U T H E R N
A L P S
S t y r i a n
B a s i n
Z a l a
B a s i n
T r a n s y l v a n i a n
B a s i n
Zagreb
Fig. 16. Palaeogeographic distribution of Limacina valvatina (Reuss, 1867) (white circles) and Limacina gramensis (Rasmussen, 1968) (black
circles) during the Late Badenian in the Paratethys. For references and localities see Table 10. Palaeogeographic map after Kováč et al. 2007.
347
THE MIOCENE “PTEROPOD EVENT” IN THE SW PART OF THE CENTRAL PARATETHYS
GEOLOGICA CARPATHICA
, 2017, 68, 4, 329 – 349
Acknowledgements: We are very grateful to both the revie-
wers and Handling editor who helped us to improve the paper.
The authors thank Marin Šoufek (CNHM) and Professor
Vladimir Bermanec (UZFS-DG) for the access to the scanning
electron microscope, Robert Košćal (UZFS-DG) and Nives
Borčić (CNHM) for technical support, Professor Tihomir
Marjanac (UZFS-DG), Marina Čalogović, Bojan Karaica and
Marko Repac for help during the field work, and to Davorka
Radovčić (CNHM) for linguistic help.
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GEOLOGICA CARPATHICA
, OCTOBER 2017, 68, 5, i–ii
doi: 10.1515/geoca-2017-00xx
www.geologicacarpathica.com
Erratum
ʻThe Miocene “Pteropod Event” in the SW part of the Central Paratethys (Medvednica Mt., Northern Croatia)’ by MARIJA
BOŠNJAK, JASENKA SREMAC, DAVOR VRSALJKO, ŠIMUN AŠČIĆ and LUKA BOSAK published in GEOLOGICA
CARPATHICA Vol. 68, No. 4, 2017, pages 329‒349, doi: 10.1515/geoca-2017-0023, error in the Figure 5 on page 335, Table 8
on page 344, Table 10 on page 345 and Figure 16 on page 346:
In Fig. 5, determinations of gastropods are erroneous and the figure caption is incorrect.
Fig. 5. A–B: Limacina gramensis (Rasmussen, 1968) from the Marija Bistrica locality. A — CNHM MB-LG1; B — CNHM MB-LG 8.
C-D: Limacina sp. (? nov.) from the Marija Bistrica locality. C — CNHM MB-L 1; D — CNHM MB-L 2. Scale bars 100 μm.
The corrected Figure caption is shown below:
Fig. 5. Juvenile benthic gastropods from the Marija Bistrica locality. A — CNHM MB sp. A-1; B — CNHM MB sp. A-2; C — CNHM MB sp.
B-1; D — CNHM MB sp. B-2. Scale bars 100 µm.
Specimens illustrated in Fig. 5 as the pteropods Limacina gramensis and Limacina sp. (nov. ?) do not represent pteropod
species, but juvenile benthic gastropods. Description of Limacina gramensis and Limacina sp. (nov.?) should be excluded from
the chapter Systematic Palaeontology, p. 333‒334, as well as Table 2 and Fig. 6, page 336.
This change affects the following tables/figures:
In Table 8 Limacina gramensis and Limacina sp. should be excluded from pteropod list. Benthic gastropoda from the Marija
Bistrica locality are added among the category: Other. The corrected Table 8 is shown below:
Species
Locality
V
MB
Pteropoda
Limacina valvatina (Reuss)
Clio fallauxi (Kittl)
Vaginella austriaca Kittl
?
Coccolithales
Calcidiscus premacintyrei Theodoridis
C. tropicus (Kamptner)
Coccolithus miopelagicus Bukry
C. pelagicus (Wallich) Schiller
Cyclicargolithus abisectus Muller (Wise)
Discoaster formosus Martini & Worsley
Helicosphaera carteri (Wallich) Kamptner
H. intermedia Martini
H. perch-nielseniae (Haq) Jafar & Martini
Pontosphaera multipora (Kamptner) Roth
P. plana (Bramlette & Sullivan) Haq
Reticulofenestra bisecta (Hay, Mohler & Wade) Roth
R. dictyoda (Deflandre) Stradner
R. minuta Roth
R. perplexa (Burns) Wise
R. pseudoumbilicus (Gartner) Gartner
Sphenolithus heteromorphus Deflandre
S. moriformis (Bronniman & Stradner) Bramlette & Wilcoxon
Umbilicosphaera rotula (Kamptner) Varol
Other
Foraminifera
Ostracods
Bivalves
Benthic gastropods
Scaphopods
Bryozoans
Fish remains
ii
GEOLOGICA CARPATHICA
, 2017, 68, 5, i–ii
In Table 10, distribution of Limacina gramensis in Central Paratethys is incorrect. The corrected Table 10 is shown below:
In Fig. 16. palaeogeographic distribution of Limacina gramensis in Croatia is incorrect. The corrected Figure 16 is
shown below:
Although L. gramensis was not found in the research area, the species is considered to be a Late Badenian index fossil,
therefore it is important for the discussion regarding the Badenian palaeogeography (parts of Discussion and Conclusion,
Fig. 16, Table 10).
The erroneous determination of gastropods on Fig. 5 does not significantly affect presented discussion and conclusions, since
the age of the investigated sites was based on the nannoplankton assemblages as well as on pteropods.
References
Species
Sedimentological
basin
Country
MIOCENE
MIDDLE MIOCENE
Badenian
Lower-Middle * Upper
1; 2; 4
Limacina valvatina
Central Paratethys
Austria
2; 4; 6; 9
Hungary
2; 3; 4;
Poland
4; 5
Czech Republic
5
Slovakia
10
Croatia
4; 7
Eastern Paratethys
Romania
8
Bulgaria
3; 4
Ukraine
2; 3; 5
Limacina gramensis
Central Paratethys
Poland
5
Czech Republic
5
Eastern Paratethys
Romania
8
Bulgaria
3; 5
Ukraine
POLAND
UKRAINE
ROMANIA
BOSNIA AND
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CROATIA
AUSTRIA
HUNGARY
CZECH
REPUBLIC
SLOVAKIA
SERBIA
SLOVENIA
W E S T E R N
C A R P A T H I A N S
Krakow
Lviv
Budapest
Bucuresti
Brno
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A
S
T
E
R
N
C
A
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P
A
T
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N
S
S O U T H E R N
C A R P A T H I A N S
D
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A
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I D
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B a s i n
Z a l a
B a s i n
T r a n s y l v a n i a n
B a s i n
Zagreb