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Introduction
The present study focuses on the Badenian Ostracoda from
the surroundings of Pokupsko, north Croatia. The investigated
area is part of the Banovina region, central Croatia, bordered
by the rivers Sava, Una, and Kupa. During the Middle Mio-
cene, the investigated area of Banovina was located at the
south-western margin of the North Croatian Basin (NCB fur-
ther on), which represented the south-western margin of the
Pannonian Basin system and the Central Paratethys Sea
(Fig. 1).
The Central Paratethys extended from Bavaria to the Car-
pathian mountain chain (Steininger & Rögl 1979, 1984; Rögl
& Steininger 1983, 1984; Rögl 1998, 1999). It was part of an
intercontinental bioprovince that, began to evolve in the Oli-
gocene, and was formed due to collision of the European
(Tisa-Moesia) and African Plates (Horváth & Royden 1981;
Kováč et al. 1998). During the Miocene, connections of the
Central Paratethys with the Mediterranean and the Indo-Pa-
cific Ocean were repeatedly established and interrupted
(Steininger et al. 1988; Rögl 1996). Such unstable connections
resulted in sea-level oscillations (Haq 1991), different deposi-
tional modes and paleoecological conditions, thus resulting in
the development of different ostracod faunas.
The formation of the NCB was associated with a passive
continental rifting. A syn-rift phase began during the Ottnan-
gian and lasted until the Middle Badenian. The post-rift
phase lasted from the Middle Badenian to the Pliocene
(Pavelić 2001).
In the Central Paratethys, a threefold subdivision of the
Badenian (16.3—12.7 Ma) is generally accepted (Papp et al.
1978; Harzhauser & Piller 2007). The Early Badenian
(Langhian) sea-level rise and trangression are related to the
global sea-level cycle TB 2.3 (Haq et al. 1988). In the NCB,
Badenian Ostracoda from the Pokupsko area
(Banovina, Croatia)
VALENTINA HAJEK-TADESSE
*
and BOŽO PRTOLJAN
Croatian Geological Survey, Department of Geology, Sachsova 2, HR-10000 Zagreb, Croatia; *valentina.tadesse@hgi-cgs.hr
(Manuscript received May 14, 2010; accepted in revised form April 14, 2011)
Abstract: In this paper we present the results of the investigations on the Badenian (Middle Miocene) ostracods from
the Pokupsko area. For the first time the presence of Badenian aged sediments in Croatia can be supported by the
occurrence of ostracod biozonal markers. Four Badenian ostracod zones are established: Lower Badenian Biozone NO7
Acanthocythereis hystrix—Bythocypris lucida, Middle Badenian Biozone NO8 Eocytheropteron inflatum—Olimfalunia
spinulosa, and the two Upper Badenian Biozones NO9 Neomonoceratina laskarevi—Miocyprideis sarmatica elongata
and NO10 Carinocythereis carinata—Phlyctenophora farkasi. On the basis of the generally accepted paleoecology of
selected genera, we identified the following ostracod faunas: shallow-water marine, shallow-water brackish-marine,
shallow-water reef, and deep-water marine. The paleontological and trace element analyses suggest that the Pokupsko
ostracod fauna lived in shallow (50 m deep), warm, and limpid waters, connected to a deeper sea and occasionally
exposed to freshwater inflows.
Key words: Miocene, Badenian, North Croatian Basin, Ostracoda, biostratigraphy, paleoecology, trace elements.
the Early Badenian deepening event resulted in deposition of
marls and gravelly calcarenites in the offshore areas (Pavelić
et al. 1998).
The second Badenian cycle corresponds to the global cycle
TB 2.4. (Haq et al. 1988). It was followed by a regression and
the lowstand Ser2 of Hardenbol et al. (1998). During the Late
Badenian, the transgression flooded even the tips of the ex-
posed blocks that had formed isolated islands during the Early
Badenian. Marine sedimentation was initiated by the deposi-
tion of gravels, which are overlain by coralline algal beds.
Fig. 1. Geographical setting of the Pannonian Basin system with
marked position of the investigated area.
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Finally, the renewed flooding belongs to the Late Bade-
nian Langhian/Serravallian cycle TB 2.5 (sensu Haq et al.
1988). Further deepening in the area of the NCB resulted in
the deposition of marls.
Generally, the Early Badenian transgressions were con-
trolled by both, tectonics (induced mainly by the back-arc
basin rifting) and eustacy, followed by forced regression.
The Late Badenian transgression and regression were domi-
nantly controlled by sea-level changes within the Central
Paratethys realm (Pavelić et al. 1998; Bakrač et al. 2010).
The results presented in this paper are exclusively based
on paleontological and geochemical analyses of the ostracod
fauna. The richness and diversity of the ostracod fauna in the
investigated area is the consequence of important changes in
the Badenian Paratethys, which had a positive effect on ma-
rine organisms. The enrichment of marine fauna was the
consequence of open seaways between the Mediterranean
Sea/Indo-Pacific and the Pannonian basin complex during
the Badenian (Rögl 1998; Popov et al. 2004; Harzhauser &
Piller 2007).
In addition to identification of the ostracode fauna, this study
focuses on (1) paleoenvironmental reconstructions; (2) Bade-
nian ostracod-based biozonation, recognized for the first
time in Croatia after the Miocene Paratethys Biozonation of
Jiříček (1983) and Jiříček & Říha (1990); and (3), compari-
son of the investigated ostracod fauna with coeval faunas
from other parts of the Central Paratethys and Tethys.
In our analysis of the ostracod fauna, we included all the
available samples; some of them were recently recovered
from logged outcrops and cores, others were already pre-
served at the Croatian Geological Survey.
Geological settings
The available literature on the Miocene deposits of the inves-
tigated area of Banovina is relatively sparse. The first geological
map of the Glinsko Pokuplje area, along with a short discus-
sion, was published by Štúr (1863). Tietze (1871) investigated
the stratigraphy of the older deposits, whereas Šuklje (1914)
suggested the existence of Tortonian, Sarmatian, and freshwater
Pliocene Congeria beds in the environs of Pokupsko.
Takšić (unpublished 1951) described geological and mining
relationships in the environs of Abes, and reported on the shaft
in Stipan. Kochansky-Devide & Slišković (1978) concluded
that the deposits of Banovina are of the Middle Miocene,
probably Late Helvetian (Karpatian) age. This conclusion is
based on determinations of the Congeria species and com-
parison with similar deposits on Mt Medvednica, as well as
on the fact that the freshwater deposits are overlain by Leitha
Limestone.
Jurišić Polšak (1979) determined the Lower Sarmatian
mollusc fauna of Glinsko Pokuplje. In an unpublished report
related to the geological map of the Vrginmost—Topusko—
Viduševac area, Šikić K. & Šikić L. (1960) described the
Dugo Selo and Stipan coal-bearing deposits.
Geological investigations associated with mapping of the
Karlovac sheet, Basic Geological Map of former Yugoslavia
1 : 100,000, were carried out from 1974 to 1976, and all the
field and laboratory reports are stored at the Croatian Geo-
logical Survey. The micropaleontological report on the Cre-
taceous and Tertiary samples of the Karlovac sheet (Šikić
1974 unpubl.) concluded that the oldest Neogene deposits
are represented by freshwater sediments immediately under-
lying Tortonian deposits with poorly preserved ostracods of
reputed Helvetian age. According to the same author, the
marine Tortonian deposits start with the Spiroplectammina
carinata (Spirorutilus carinatus) Biozone. Later, Pikija
(1984; unpubl. report) concluded that the Badenian deposits
transgressively and unconformably overlie the older depos-
its, the latter ranging from the Triassic dolomites to basal
breccias, Paleogene clastites, tuffs and tuffites.
During the investigations on the thickness and spatial distri-
bution of subsurface coal-bearing deposits in the village of
Stipan, Vrginmost—Lasinja area (Lukšić 1985, unpubl.), Miknić
(1985, unpubl.) micropaleontologically analysed 16 cores and
confirmed the presence of Badenian strata for two of them.
Bajraktarević (1983) analysed foraminiferal fauna and
nannoplankton from Glinsko Pokupje (localities Čremušnica,
Babin potok, and Ilovačak) and had assigned the Badenian
age to the deposits (Spiroplectammina carinata Zone).
In a paper on the Neogene ostracod assemblages of Yugo-
slavia, Sokač & Krstić (1987) described the Šljivovac locality
as one with a rich and well-preserved freshwater ostracod fauna.
In her Master’s thesis, Hajek-Tadesse (unpublished, 2000)
described the ostracod fauna from Pokupsko.
Study sites and methods
Study sites
The investigated area is situated near the village of Pokupsko,
Banovina region, south of the Kupa River in Croatia (Fig. 2).
In addition to the Pokupsko outcrops, two cores at the Stipan
locality (BS-3 and BS-10) and ten samples acquired during
the 1974 Basic Geological mapping, Karlovac sheet, have
been available for this study. We have also analysed five
previously collected samples from the area; these samples
are available at the Department of Geology, Croatian
Geological Survey.
Pokupsko
The bulk of the samples has been collected at the Pokups-
ko outcrop near the Kupa River, along the unpaved road run-
ning parallel to the Kupa fluvial terrace, one kilometer
southwest from Pokupsko. The outcrop extends 15 m in
width and it is 5 to 6 meters thick. It consists of yellowish,
poorly cemented sandstones, alternating laterally with yel-
lowish-white sandy marls. The outcrop is very friable, and
the bedding cannot be observed.
Stipan
The samples labelled Stipan well (BS), previously recov-
ered near the Stipan village between Vrginmost and Lasinja
were assigned to the Badenian age (Lukšić 1985, unpubl.).
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The BS-3/84 well is situated in the Minići village, in an
area known as Ostrožin. The sediment core is composed of
sandy-limy Badenian deposits, whose thickness reaches up
to 96.0 m; coal intercalations occur near the bottom of the
core. Micropaleontological samples were collected from the
intervals 42.00—42.10 m and 59.90—60.00 m. In the first
42.00—42.10 m interval, the Badenian deposits are fine-
grained, and sandy-limy. In the second interval (59.90—
60.00 m), the deposits are marly-sandy.
The BS-10/84 well is situated in a ditch 200 m south of the
Šljivovac village, and it penetrated through 93 meters of
Badenian deposits. The analysed samples were collected
from sandy fossiliferous marls (interval 66.00—66.10 m).
Based on Lukšić’s (1985, unpubl.) results it is obvious that
the Badenian deposits in the Stipan wells and the surround-
ing area are stratigraphically younger than the coal-bearing
series, and/or unconformably overlie the older bedrock com-
posed of Cretaceous flysh-like facies and the volcanogenic-
sedimentary complex.
The Lower Badenian deposits consist of decimeter-thick
interbedded Leitha Limestone, calcareous sandstones, and
light-yellow massive Nullipore Limestone, composed of
Lithothamnium lumps. The upper parts of the Badenian are
composed of marls with varying amounts of carbonate com-
ponent, so that transitions from marls to calcareous marls to
marly limestones can be observed. Marls are sandy and fos-
siliferous, and sporadically they contain tuffitic intercala-
tions. The most remarkable Badenian lithologic member is
the grey to light-yellow bedded to massive organogenous
Leitha Limestone, which is often accompanied by lime-
stones with abundant macrofossil shells. The peaks of all
major neighbouring hills are composed of the same type of
sediment, because the underlying sandy-marly parts are
more easily eroded.
Samples from mapping of the Karlovac Sheet
During the 1974 geological mapping of the Karlovac
sheet, sections Sjeničak and Čremušnica were sampled for
micropaleontological analyses. Ten samples were collected
and these have contributed to the completion of the sampling
net in the investigated area. According to the field-note data,
the samples have been collected from various sediments.
Samples KA/13816, 13817, 13818 and 13821 were collected
from sandy-clayey sediments; 13822, 16630, and 15533
were collected from marly-sandy and sandy sediments;
15271 was collected from sandy sediments; and 13703 and
13671 from clays.
Samples from the micropaleontological collection
The five samples found in the micropaleontological col-
lection of the Department of Geology were found among the
stored Neogene material labelled Jabukovac, Šljivovac, Pet-
rinja, Trepča and Bučica. Since the above samples do not
have any additional marks, it was impossible to pinpoint pre-
cise locations as well as additional information on accompa-
nying fauna, sediment type, and the environment in which
they were found.
Methods
The collected field samples were disaggregated by soaking
in a hydrogen peroxide solution for 24 hours, followed by
washing through sieves (0.5; 0.25; 0.125; 0.063 mm) and
drying. Some of the samples needed extra cleaning and these
were re-soaked in hydrogen peroxide and treated ultrasoni-
cally for approximately 20 seconds. A hundred grams of
each dried residue were observed under a stereomicroscope.
Ostracods and other fossil remains (foraminifers and gastro-
pods) were hand-picked, counted and determined. Ostracods
were qualitatively, not selectively picked in order to preserve
the relative composition of the thanatocoenosis.
The stored material was subjected to an additional clean-
ing process in an ultrasound dish after additional cleaning
the material was picked again.
For the geochemical analyses of trace elements, the avail-
able amount of shells (about 50 mg) was weighed into 30 ml
glass test tubes for one-time use, and 3 mg of concentrated
HCl p.a. was added. The solutions were then analysed in the
Fig. 2. Location map: study
area with marked position of
Pokupsko outcrops (
l); two
wells Stipan (BS-3 ); and sam-
pling points (KA13671 •).
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Croatian Geological Survey by a simultaneous ICP atomic
emission spectroscope Jobin-Yvon JY-5OP. Concentrations
of Mg, Ca, Mn, and Sr were determined; small sample and di-
lution prevented determination of other elements. The accu-
racy of the analyses was checked by the parallel analysis of
the International Geological Standard Soil Samples GXR-5,
GXR-6, and SJS-1, verified by USGS. The data were statisti-
cally analysed using the statistical package STATISTICA.
Photographs were made with the SEM at the Ina Naftaplin,
Zagreb. All ostracod specimens are stored in the repository
of the Croatian Geological Survey.
In this paper we essentially followed the ostracod classifi-
cation of Morkhoven (1963), Hartmann & Puri (1974) and
Horne et al. (2002).
Paleontological and geochemical analyses
Ostracods
The Badenian ostracod fauna from 17 localities (22 sam-
ples) analysed from the vicinity of Pokupsko is interesting due
to its diversity and richness (Table 1; Figs. 3, 4). We have de-
termined a total of 45 genera and 64 species. Samples were
grouped according to differences or similarities of the ostra-
cod fauna at one locality and by correlation between different
localities.
In addition to the presence of zonal markers for the Ostra-
cod Biozone for the Badenian (Jiříček 1983; Jiříček & Říha
1990), we attempted to define the attribution of samples to a
particular zone, out of the four Badenian ostracod zones rec-
ognized so far. However, the ostracod zonations should be
treated with caution, because some new results in ostracod
research have increased the stratigraphic range of several
species (Zorn 2003, 2004; Gross 2006).
Pokupsko
The Pokupsko locality contains the majority of the species
determined in this paper. Due to poor bedding conditions, the
samples in the field were not taken continuously. Following
the analysis of the ostracod fauna, and based on the differ-
ences observed in quantitative and qualitative representation
of genera and species (differences in colour) and on sedimen-
tological determinations, the eleven samples were classified
into two groups, labelled Pokupsko I and Pokupsko II.
In Pokupsko I group, the representatives of the tribe Aurli-
nae are the most abundant, especially the species Aurila angu-
lata, A. cicatricosa, A. haueri, A. punctata, Senesia trigonella,
S. cinctella, Pokornyella deformis; also abundant are the fol-
lowing species: Tenedocythere sulcatopunctata, Pachycau-
dites ungeri, Heliocythere vejhonensis, Costa edwardsi, C.
reticulata, C. punctatissima, Carinocythereis carinata,
Occultocythereis bituberculata, Grinioneis haindigeri, Henry-
howella asperrima, Pterygocythereis calcarata, Olimfalunia
plicatula, O. spinulosa, Olimfalunia sp. and Cytheridea sp.
Less abundant are the following species: Cytherella com-
pressa, Bairdoppilata sp., Callistocythere aff. canaliculata,
Echinocythereis scabra, Cnestocythere lamellicostata, Loxo-
corniculum hastatum, L. punctatella, Neomonoceratina sp.
and Xestoleberis cf. X. glabrescens.
The Pokupsko II group contains all of the above mentioned
species from Pokupsko I, and in addition it contains an in-
creasing number of deeper water species: Krithe sp., Para-
krithe dactylomorpha and Eucythere sp. The genus
Semicytherura is also abundantly represented, with Semi-
cytherura alata, S. acuticostata, S. cf. S. alifera, Semicytheru-
ra sp. as well as Eucytherura textilis, Hemicytherura sp.,
Cytheretta tenuipunctata dentata, Flexus triebeli and a few re-
mains of Ruggieria cf. R. micheliniana, species which oc-
curred only in the Pokupsko II group.
Also, in contrast to the Pokupsko I group, ostracods in the
Pokupsko II group are preserved mostly in adult stage and
with closed valves. The age of both groups, Pokupsko I and
Pokupsko II, is Late Badenian.
The Stipan wells
Two cores have been studied: BS-3 with two intervals and
BS-10 with one interval.
BS-3 (42.00—42.10 m). The most abundant species within
this interval are Olimfalunia plicatula, Olimfalunia sp.,
Cytheridea sp. and C. acuminata; less abundant are Cytherella
compressa, Aurila angulata, Senesia cinctella, Callistocythere
aff. canaliculata and Cnestocythere lamellicostata, whereas
Costa edwardsi, Loxocorniculum hastatum and Bosquetina
dentata are present with only a few specimens. Of particular
significance is the recovery of Cytherelloidea sissinghi, which
represents its first finding within the Paratethys realm.
BS-3 (59.90—60.00 m). This interval contains a rich and di-
verse ostracod fauna. Of particular interest is the finding of
Ruggieria ex gr. carinata and Cytherelloidea sissinghi, forms
that are not frequent in Paratethys. Their findings in the inves-
tigated area represents an additional proof of transgressive
events in the Badenian and migration of ostracods from the
Tethys to Paratethys. Also, abundant are the representatives of
the family Cyprididae, indicative of deeper water environ-
ment, such as Paracypris polita and Phlyctenophora affinis,
as well as Henryhowella asperrima and Pterygocythereis cal-
carata. Similarly to the preceeding interval of the same well,
the species Olimfalunia plicatula, O. spinulosa, Olimfalunia
sp., Cytheridea sp., Cytherella compressa are also abundant,
whereas the folloving are less numerous: Callistocythere aff.
canaliculata, Costa edwardsi, Loxocorniculum hastatum, L.
punctatella, Xestoleberis cf. X. glabrescens, Bosquetina den-
tata and Bythocythere cf. B. neerlandica.
The sample from this interval differs in its faunal content
from the upper interval of the same well, being more similar
to the fauna from the well BS-10 (60.00—60.10 m).
BS-10 (60.00—60.10 m). The ostracod assemblage from
this sample is one of the most interesting in the whole area
investigated. It is dominated by valves of adult individuals,
80 % of which are very well preserved.
The finding of the key species Acanthocythereis hystrix is
important for the age determination, and it supports an earlier
attribution to the older part of the Badenian (unpublished
Miknić 1985). In addition to that species, the sample con-
tains important representatives of the family Cyprididae, as
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Table 1: Distribution of the ostracod species within the samples of the Pokupsko area.
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well as Ruggieria ex gr. carinata, Cytherella inaequalis,
Cytherelloidea sissinghi, Henryhowella asperrima, Pterygo-
cythereis calcarata, Bythocythere cf. B. neerlandica, Olim-
falunia plicatula, Cytheridea sp., Flexus triebeli, Costa
edwardsi and Xestoleberis cf. X. glabrescens.
According to the results obtained from the three intervals
in two cores, we argue that the rich and well preserved ostra-
cod fauna from the Stipan cores is interesting for several rea-
sons. In the BS-3 well, a continuous transition from the
upper part of the Lower Badenian into the lower part of the
Middle Badenian may be discerned based on foraminifers
(unpublished Miknić 1985), and a comparison of ostracod
fauna from the well BS-3 59.90—60.00 m interval with the
BS-10 60.00—60.10 m interval can be carried out.
The recovery of the biostratigraphic marker-species Acan-
thocythereis hystrix in the sample BS-10 (60.00—60.10 m) is
extremely important, as it proves the Early Badenian age of
the sample. The occurrence of typical representatives of a
deeper water environment, such as: Acanthocythereis hystrix,
Henryhowella asperrima, Bythocythere cf. B. neerlandica and
Paracypris polita suggests deepening of the basin; Cytherel-
loidea sissinghi and Ruggieria ex gr. carinata, which have
previously not been recorded from the Badenian deposits of
the Central Paratethys, indicate migration of ostracod faunas
from the Tethys into this region. The data about the reverse
migration of the ostracod fauna, namely from the Paratethys
to Tethys (Szczechura & Abd-Elshafy 1988) also support
these considerations.
Samples collected during mapping of the Karlovac sheet
Ten samples collected during the geological mapping of
the Karlovac sheet are classified into six groups according to
the similarity of the ostracod fauna observed.
Karlovac I (KA 16630, 15533). This group is distinguished
from the other groups by the presence of the index species
Acanthocythereis hystrix, indicating the Early Badenian age.
Other abundant species include Ruggieria ex gr. carinata,
Pterygocythereis calcarata, Henryhowella asperrima, Flexus
triebeli, Cytheridea sp., and less numerous Grinioneis haidin-
geri, Aurila haueri, Costa edwardsi and Bairdoppilata sp.
Karlovac II (KA 13703). The recovery of Eocytheropteron
inflatum in sample KA 13703 has been decisive in separating
this sample into a different group. In addition to Eo-
cytheropteron inflatum, this group is characterized by a mixed
shallow-water/deep-water ostracod fauna. The abundant os-
tracod fauna includes Ruggieria ex gr. carinata, Phlycteno-
phora affinis, Paracypris polita, Olimfalunia spinulosa,
Olimfalunia sp., Cytheridea sp., Pontocythere cf. P. curvata
and, less numerous, Aurila cicatricosa, Bosquetina dentata
and Senesia sp.
Karlovac III (KA 13817, KA 13816). In the third group,
the most numerous species are shallow-water representatives
of the genus Aurila, with A. angulata, A. cicatricosa, A.
haueri, A. punctata, Senesia trigonella, S. cinctella, as well
as the species Costa edwardsi, C. reticulata, Cytheridea sp.,
Pontocythere cf. P. curvata, Callistocythere aff. canalicula-
ta, Cnestocythere lamellicostata, Phlyctenophora affinis,
Grinioneis haidingeri, Pterygocythereis calcarata, Olim-
falunia plicatula, O. spinulosa and Olimfalunia sp. Similar
to the above-mentioned groups, this group also contains
Ruggieria ex gr. carinata, and the age of the sample is deter-
mined as Middle Badenian; this age was also documented by
foraminifers (unpublished Šikić L. 1974).
Karlovac IV (KA 13818, KA 13821). The fourth group has
been based on findings of Miocyprideis sp. in two samples.
The dominant species are Cytheridea sp., Costa edwardsi and
C. reticulata, but Olimfalunia plicatula, Carinocythereis
carinata, Senesia cinctella and A. cicatricosa are also
present, though less numerous. Specimens of the genera
Costa and Olimfalunia show environmentally cued polymor-
phous changes (Peypouquet et al. 1988) on their valves. The
Late Badenian age of the sample has been determined based
on the ostracod fauna. According to the foraminiferal assem-
blage, the “Upper Tortonian” Ammonia beccarii Zone was
determined (unpublished Šikić L. 1974).
Karlovac V (KA 15271). The dominant species in this
group are Cytherella compressa, Cytherella inaequalis,
Paranesidea sp., Phlyctenophora affinis and Aurila angulata.
Contrary to previous groups, this group is impoverished with
respect to both the number of species and the number of
specimens. On the basis of the ostracod fauna and foramini-
fers (unpublished Šikić L. 1974), the Late Badenian age of
that sample has been determined.
Karlovac VI (KA 13822, KA 13671). This group has been
identified based on the shallow-water fauna, which, in con-
trast to the KA III group, is not dominated by the genus Aurila.
This group is characterized by Phlyctenophora affinis, Costa
edwardsi, C. reticulata, Cytheridea sp., C. acuminata and
less abundant deeper water species Paracypris polita,
Bosquetina dentata, Echinocythereis scabra and Pterygo-
cythereis calcarata. The age of the sample is Late Badenian.
The ostracod fauna in the samples collected during mapping
yields some important information on Badenian deposits in
the investigated area. The key species Acanthocythereis hys-
trix in the first group and Eocytheropteron inflatum in the
third group define the age of these deposits as Early and Mid-
dle Badenian. The common characteristic of the first three
groups is the presence of the “new-entry” species Ruggieria
ex gr. carinata, whereas the presence of the genus Miocypri-
deis in the fourth group indicates global paleoecological
changes in the Paratethys realm during the transition from
Middle to Late Badenian.
Šljivovac
Although the Šljivovac locality is situated close to Stipan,
its faunal content is more similar to the ostracod fauna from
the B-3 well (59.90—60.00 m interval), except for the species
Parakrithe dactylomorpha, Pontocythere cf. P. curvata, Para-
cytheridea sp. and Cytherella beyrichi, which were found only
in that sample. In the Šljivovac sample, the most abundant
species are Henryhowella asperrima, Pterygocythereis cal-
carata, Cytheridea sp., Costa edwardsi, Olimfalunia plicatula
and Callistocythere aff. canaliculata. Of particular importance
is findings of the “new-entry” species Cytherelloidea sissinghi,
which prior to this study has not been identified in the
Paratethys realm. The environmentally cued polymorphous
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Fig. 3. SEM microphotographs of selected ostracods. 1 – Cytherella compressa (Münster), sample Pokupsko I, right valve, external view.
2 – Bairdoppilata sp., sample Pokupsko I, right valve, external view. 3 – Cytherelloidea sissinghi Szczechura, sample Stipan (BS-10), right
valve, external view. 4 – Henryhowella asperrima (Reuss), sample Stipan (BS-10), left valve, external view. 5 – Bythocypris sp., sample
Pokupsko II, left valve, external view. 6 – Acanthocythereis hystrix (Reuss), sample Stipan (BS-10), right valve, external view. 7 – Pterygo-
cythereis calcarata (Bosquet), sample Pokupsko I, left valve, external view. 8 – Echinocythereis scabra (Münster), sample Pokupsko I, right
valve, external view. 9 – Ruggieria ex gr. carinata Moyes, sample Stipan (BS-10), right valve, external view. 10 – Krithe sp., sample
Pokupsko II, right valve, external view. 11 – Eocytheropteron inflatum (Schneider), sample KA 13703, left valve, external view. 12 – Ponto-
cythere curvata (Bosquet), sample KA 13816, right valve, external view. 13 – Semicytherura acuticostata (Sars), sample Pokupsko I, left valve,
external view. 14 – Hemicytherura sp., sample Pokupsko II, right valve, external view. 15 – Semicytherura sp., sample Pokupsko II, right valve,
external view. 16 – Cytheretta tenuipunctata dentata Brestenska, sample Pokupsko I, left valve, external view. 17 – Flexus triebeli (Ruggieri),
sample Pokupsko I, left valve, external view. 18 – Semicytherura alata (Lienenklaus), sample Pokupsko II, left valve, external view.
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Fig. 4. SEM microphotographs of selected ostracods. 1 – Aurila haueri (Reuss), sample Pokupsko I, right valve, external view. 2 – Pokor-
nyella deformis (Reuss), sample Pokupsko I, right valve, external view. 3 – Aurila punctata (Münster), sample Pokupsko I, right valve, exter-
nal view. 4 – Elofsonella sp., sample Pokupsko II, left valve, external view. 5 – Aurila angulata (Reuss), sample Bučica, right valve,
external view. 6 – Carinocythereis carinata (Roemer), sample Pokupsko II, left valve, external view. 7 – Senesia sp. sample KA 13703, left
valve, external view. 8 – Tenedocythere sulcatopunctata (Reuss) sample Pokupsko I, left valve, external view. 9 – Callistocythere aff.
canaliculata (Reuss), sample KA 13816, right valve, external view. 10 – Xestoleberis glabrescens (Reuss) sample Pokupsko II, left valve,
external view. 11 – Tenedocythere cf. T. salebrosa Uliczny, sample Pokupsko I, left valve, external view. 12 – Ruggieria cf. R. micheliniana
(Bosquet), sample Pokupsko II, right valve, external view. 13 – Pachicaudites ungeri (Reuss), sample Pokupsko II, left valve, external view.
14 – Tenedocythere sp. sample Pokupsko I, left valve, external view. 15 – Costa edwardsi (Roemer), sample Bučica, left valve, external
view. 16 – Loxoconcha punctatella (Reuss), sample KA 13818, right valve, external view. 17 – Phlyctenophora affinis (Schneider), sample
KA 13671, right valve, external view. 18 – Heliocythere vejhonensis Prochazka, sample Pokupsko I, right valve, external view.
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Mg
ppm
Mn
ppm
Sr
ppm
Mg/Ca Sr/Ca
Aurila
9214
84
1497
0.033
0.0055
Cytherella
5942
46
1650
0.019
0.0052
Bairdoppilata
5327 163 1170 0.012 0.0026
Cytheridea
3868 108 1512 0.012 0.0045
Tenedocythere
2935
85
983
0.014
0.0063
Table 2: Obtained results of trace element analyses for selected
ostracod genera from the Pokupsko area.
changes (Peypouquet et al. 1988) have been observed on
valve surfaces of Costa and Olimfalunia. The Middle Baden-
ian age of this sample has been determined on the basis of the
ostracod fauna and Sokač’s observations (pers. comm.).
Trepča and Bučica
The faunas from the two samples are very similar. They
are both dominated by Callistocythere aff. canaliculata,
Cnestocythere lamellicosta, Aurila angulata, A. haueri, Costa
edwardsi, Olimfalunia plicatula and O. spinulosa. In addi-
tion to the species mentioned in the Trepča sample, less nu-
merous are Senesia sp. and Bosquetina dentata, absent in the
Bučica sample, which contains, instead, Cytheridea sp. Ac-
cording to the analysis of the ostracod fauna and comparison
with fauna from other localities, the Late Badenian age has
been determined.
Jabukovac and Petrinja
The main characteristic of the two localities is the predom-
inance of adult specimens of Pontocythere cf. P. curvata.
Other species common to both localities include Xestoleberis
cf. X. glabrescens and Olimfalunia plicatula. In addition, the
Jabukovac locality contains Cytherella compressa, C.
inaequalis and Paracypris polita.
The Petrinja sample contains a richer fauna than the above
described. In addition to the above species it also includes
Bairdoppilata sp., Phlyctenophora affinis, Callistocythere
aff. canaliculata, Aurila angulata, Pokornyella deformis,
Tenedocythere sulcatopunctata, Costa edwardsi, Grinioneis
haidingeri, Olimfalunia plicatula and Cytheridea acuminata.
The Late Badenian age of this sample has been determined
on the basis of the ostracod fauna.
Trace element analyses of the Upper Badenian ostracod
valves
Ostracods are organisms sensitive to environmental
changes that control their occurrence, distribution, and re-
production (Barbieri et al. 1999; Holmes & Chivas 2002).
Ostracod valves are made of Ca and Mg-carbonates, but other
chemical elements, including Sr, K, Na, Mn, Fe, and Ba,
may also be present.
Chivas et al. (1983) were among the first who showed that
certain geochemical parameters of ostracod valves yielded more
detailed infomation on the environment in which the fauna
lived. The analysis of concentrations and ratios of Ca, Mg, Sr,
and Mn in ostracod valves from the vicinity of Pokupsko (all
samples studied come from the sixth biofacies, thus being
Late Badenian in age) enabled us to do the paleoenviron-
mental reconstruction. Generically determined valves were
separated for geochemical analyses. According to Barbieri et
al. (1999), this is acceptable since species of the same genus,
or even phylogenetically related genera, have similar Sr and
Mg concentrations.
The selected genera inhabited both shallow-water (Aurila,
Cytheridea, Tenedocythere) and deep-water environments
(Cytherella, Bairdoppilata).
The concentrations of elements vary in different deposits,
depending on regional features, differences in mineralogy, and
ratio between inorganic and organic components. Chivas et
al. (1983) argued that the amount of Mg in ostracod valves
depends on the temperature and concentration of Mg
2+
in wa-
ter, whereas the Sr content depends directly on the concentra-
tion of Sr
2+
in water. In sea water, the concentration of Mg and
Sr increases with the increasing salinity and decreases with the
decreasing temperature, whereas the higher Mn concentrations
are due to increased influx of terrigenous material (Prasada
1996). The ratio of Sr/Ca and Mg/Ca concentrations in ostra-
cod valves remains unchanged if the water, temperature and
salinity remain constant. If the value of Mg/Ca ratio varies,
and the Sr/Ca ratio remains constant, the water temperature
varies and salinity is constant. Finally, if both the Sr/Ca and
Mg/Ca ratios vary, changes in salinity and insignificant
changes in water temperature are expected.
Dweyer et al. (1995) were among the first to postulate a
connection between the variations in Mg/Ca ratio in valves
of the deep-water genus Krithe and the water depth in which
the genus lived. The authors concluded that the lower Mg/Ca
values in valves correspond to lower water-temperature, and
higher values correspond to higher temperatures.
In both organic and inorganic aragonite, the Sr/Ca ratio of-
ten increases with the lowered temperature (Prasada 1996),
whereas the rather stable Sr/Ca values are indicative of an
equalized temperature.
Certain conclusions can be drawn from the above men-
tioned data and the results obtained by geochemical analyses
shown in Table 2 and Fig. 5a—c.
The greatest deviations from the group of analysed genera
in the first two diagrams (Fig. 5a,b) is found in the genus
Bairdoppilata. The low Mg/Ca and Sr/Ca ratios, as well as the
increased concentrations of Mn (163 ppm), distinguish this
genus from the group of other genera. This may be explained
through the interpretation of different micro-environments.
It is known, that the genus Bairdoppilata inhabits both
very shallow and deep marine environments. According to
the geochemical analyses of the Pokupsko sample, we sug-
gest a shallow-water environment with considerable influx
of terrigenous material, and so close to lower salinity fresh-
water sources enriched in Mn as indicated by the lower val-
ues of the Mg/Ca and Sr/Ca ratios.
As shown in Fig. 5b,c, the genus Bairdoppilata is joined
by the genus Tenedocythere, in its deviation from the group
of shallow-water marine forms. The lowest Sr concentrations
(985 ppm) in the valves of this genus, as well as the low Mg
concentration, indicate the change in water salinity. Because
we are dealing with a marine environment, such deviations
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Fig. 5. a – Mg/Ca versus Sr/Ca plot in the valves of analysed gen-
era. b – Sr/Ca versus Mn concentration plot in the valves of analy-
sed genera. c – Sr concentration versus Mn concentration plot in
the valves of analysed genera.
may be explained by decreased magnesium and strontium
values as a consequence of crystal growth (Prasada 1996).
The other three analysed genera do not show considerable
deviations regarding the concentrations of elements, the ex-
ception being genus Cytherella, in which the lowest Mn con-
centration (46 ppm) and the highest Sr concentration
(1650 ppm) have been determined. The deviations in Mn and
Sr concentrations may be explained in two ways, either as an
adaptability to greater depths, or life in a protected marine
shallow-water environment, devoid of freshwater influx. The
decision, on which hypothesis should be accepted can be
reached only after analysis of the whole ostracod assemblage
in which Cytherella has been found.
Taking into consideration all the above-mentioned data, the
environment during life of the selected genera corresponded to
the model of an open/half-closed lagoon with periodical fresh-
water influx, presented in the paleoenvironment reconstruc-
tion section of this paper.
Discussion
Correlation of samples based on analysed fauna
The studied ostracod faunas were collected in a relatively
small investigated area on the basis of common characteris-
tics, namely key species, facies indicative species and com-
mon biocoenoses. Accordingly, we were able to classify 25
samples into 14 groups; groups were classified into six bio-
facies. These biofacies do not necessarily represent biostrati-
graphic units; sometimes, they are identified exclusively
according to the paleoecological diversity of the constituent
fauna (Fig. 6.).
First biofacies. It includes the Stipan BS-10 (60.00—60.10 m)
fauna, and its temporal and faunistic equivalent, KA I (15533,
16630); these two groups share three important characteristics.
Both groups contain the key species Acanthocythereis hys-
trix, which determines the attribution of the samples to the
homonym biozone of the Lower Badenian Biozone (NO7)
Acanthocythereis hystrix—Bythocypris lucida (Jiříček 1983;
Jiříček & Říha 1990).
Also, both groups contain a mixed deeper and shallow-wa-
ter fauna. Among the deeper water forms, the most abundant
are Henryhowella asperrima, Pterygocythereis calcarata
and Bythocythere neerlandica, whereas the shallower envi-
ronment is characterized by four genera, namely Olimfalu-
nia, Cytheridea, Flexus and Costa.
The findings of the so-called “new-entry” species Rug-
gieria ex gr. carinata and Cytherelloidea sissinghi imply,
the transgressive changes occurring during the Early Bade-
nian, and the migration of ostracod species from southern
parts of the Mediterranean, indicating the establishment of
the Tethys—Paratethys connection.
The second biofacies continuously succeeds the first biofa-
cies. It includes samples from the Stipan BS-3 well (59.60—
60.00 m) and the Šljivovac locality. Like the first facies
described above, it contains a mixed fauna of deeper water os-
tracods such as Henryhowella asperrima, Pterygocythereis
calcarata, Bosquetina sp., Paracypris polita, Phlyctenophora
affinis and Bythocythere neerlandica and shallow-water
forms, dominated by genus Olimfalunia with several species
of the Olimfalunia—Cytheridea—Callistocythere—Costa associ-
ation. Due to the abundance of genus Olimfalunia and contin-
uous sedimentation that is clearly observed in the Stipan
well samples, it is possible to assign the samples from this
biofacies to the Middle Badenian ostracod Biozone (NO8),
Eocytheropteron inflatum—Olimfalunia spinulosa (Jiříček
1983; Jiříček & Říha 1990).
Both samples contain the species Ruggieria ex gr. carinata
and Cytherelloidea sissinghi.
The third biofacies continuously overlies the second biofa-
cies. The samples of this biofacies BS-3 (42.00—42.10 m) and
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Fig. 6. Ostracod biozonation of the Badenian samples from the Pokupsko area.
KA (13817, 13816) are distinguished from the previous facies
on the basis of paleoecological differences. The biofacies is
characterized by shallow-water ostracod assemblages, that
dominate over the less numerous deeper water forms. The
most abundant form is genus Olimfalunia (with three species),
associated with genera Aurila, Costa and Cytheridea. On the
basis of the predominance of the Olimfalunia genus, this bio-
facies is assigned to the Middle Badenian ostracod Biozone
(NO8) Eocytheropteron inflatum—Falunia spinulosa (Jiříček
1983; Jiříček & Říha 1990).
The two samples of this biofacies differ in two details. In
sample BS-3 we found the species Cytherelloidea sissinghi,
which is missing in the KA II group, whereas in KA II we
found Ruggieria ex gr. carinata, which is not present in the
BS-3 sample. In this level we notice the last occurrence of
the new-entry species Cytherelloidea sissinghi.
Fourth biofacies. Samples from the KA III group
(13703). The recovery of well preserved and numerous adult
specimens of Eocytheropteron inflatum
(which is found only in this sample) is impor-
tant biostratigraphical information. It is well
known as the key species of the Middle
Badenian Biozone in the Paratethys (Jiříček
1983; Jiříček & Říha 1990) and therefore its
recovery in the fourth biofacies indicates the
Middle Badenian, Biozone (NO8) Eocythero-
pteron inflatum—Olimfalunia spinulosa. This
biofacies is characterized by mixed deep-wa-
ter and shallow-water ostracod assemblages,
with Phlyctenophora affinis, Paracypris
polita, Olimfalunia spinulosa, Cytheridea
sp., Pontocythere cf. P. curvata, and, less
numerous, Aurila cicatricosa, Bosquetina sp.
and Senesia sp. We also noticed the last
occurrence of Ruggieria ex gr. carinata in
this level.
Fifth biofacies. The samples from the
KA IV group (13818, 13821) are classified as
a separate biofacies. The most significant fea-
ture of this facies is the presence of Miocypri-
deis sp. Though it is not the marker species of
the well known Upper Badenian Zone Mio-
cyprideis elongata (Jiříček 1983), the recovery
of this genus nevertheless indicates paleoeco-
logical changes, such as decreasing water sa-
linity and depth, which are characteristic of
the region during the lowest part of the Upper
Badenian, namely the Miocyprideis elongata
(NO9) Biozone (Jiříček 1983; Jiříček & Říha
1990).
The samples are dominated by the shallow-
water genera Cytheridea and Costa; less
abundant are Olimfalunia plicatula, Carino-
cythereis carinata, Senesia cinctella and
Aurila cicatricosa. Environmentally cued
polymorphous changes (Peypouquet et al.
1988) on valves of the Olimfalunia and Costa
genera have also been noticed.
Sixth biofacies. This biofacies includes
samples from the Upper Badenian groups of Pokupsko I,
Pokupsko II, Trepča, Bučica, Jabukovac, Petrinja, KA V
(13822, 13671), and KA VI (15271). This is the richest biofa-
cies, according to both the number of analysed samples and
the number of determined species.
The differences in ostracod faunas within the analysed sam-
ples are most clearly visible in the distribution of deep-water
species and different shallow-water assemblages. Thus, the
Pokupsko II group contains more abundant, typically deep-
water species, Henryhowella asperrima, Pterygocythereis
calcarata, Krithe sp., Parakrithe dactylomorpha and Echino-
cythereis scabra, whereas in the Pokupsko I group we found
only three deep-water species, Henryhowella asperrima,
Pterygocythereis calcarata and Echinocythereis scabra.
The recovery of Echinocythereis scabra is particularly in-
teresting, because in addition to Pokupsko, it has been iden-
tified only in the KA V sample, and is generally rarely found
in the Paratethys. That species is most frequently found in
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the contemporaneous deposits of the Aquitanian Basin,
France (Moyes 1965; Ducasse & Cahuzac 1997).
Each of the above-mentioned samples or groups also contains
a recognizable dominant shallow-water ostracod association.
Thus, Pokupsko I is characterized by the Aurila—Tene-
docythere—Costa association, and Pokupsko II by the Semi-
cytherura—Cytheridea—Cytheretta association.
In the Karlovac IV group, the dominant genera are Cyther-
ella, Paranesidea, Aurila, and in Karlovac V, Costa and
Cytheridea. The Jabukovac and Petrinja samples are charac-
terized by the shallow-water Pontocythere—Costa—Cytheridea
assemblage, whereas Trepča and Bučica contain the Aurila—
Costa—Olimfalunia association.
The entire ostracod assemblage – except some species – is
well known from the Badenian deposits of Central Para-
tethys. It is most similar to the Badenian fauna of the Vienna
Basin, Lower Austria (unpublished Huber-Mahdi 1984; Zorn
2003, 2004; Gross 2006; Gebhardt et al. 2009). The shallow-
water part of the fauna correlates well with the shallow-water
Badenian fauna of Podvin, Vienna Basin, southern Moravia
(Zelenka 1985), and the deeper water fauna of Pokupsko
shows similarities with the Middle Miocene deeper water
fauna of the pre-Karpatian depression of Poland (Szczechu-
ra 1994). The described fauna is also comparable with the
Badenian fauna from six wells in southwest Poland (Paruch-
Kulczycka 1992).
A non-isolated realm for the studied samples is further
suggested on the basis of similarities with the Middle Mio-
cene ostracod assemblages of a wider geographical area.
Thus, we may distinguish a dozen species that are common
within the Middle Miocene ostracod fauna of Libya and
Egypt (Szczechura & Abd-Elshafy 1988), the island of Crete
(Sissingh 1972), the Langhian ostracods from Sicily
(Dall’Antonia et al. 2001), Northern Sardinia (Bossio et al.
2006), the Rhodanian Basin (Carbonnel 1969), the Miocene
fauna of the NW Basin of northern Germany (Uffenorde
1981) and the Middle Miocene ostracods from the West
Coast of India (Bhandari et al. 2001).
Ostracod paleoecology of the surroundings of Pokupsko
The regional Middle Miocene geological events are closely
associated with the ostracod fauna distribution. The open-
ings or closings of sea straits, transgressions, regressions, as
well as climatic changes, had a deep impact on composition
of the ostracod association, preservation state of the valves,
frequency of particular genera and species, and the occur-
rence of polymorphous changes on ostracod valves.
Differences in the state of preservation and composition of
biocoenoses are probably the result of different energy con-
ditions and sedimentation rates (Whatley 1983; Boomer et
al. 2003).
Slow sedimentation typically results in well preserved,
separated valves, and complete biocoenoses are found. The
Lower and Middle Badenian samples analysed in this paper
are an example of such a rate of sedimentation.
During high-energy conditions and/or rapid sedimenta-
tion, unfavourable conditions for preservation occurred, re-
sulting in selectively sorted valves. Complete, selectively
sorted carapaces (usually lacking juvenile specimens), with
non-separated valves, are found mostly in Upper Badenian
samples of the Pokupsko surroundings.
The lack of autoecological data on Badenian species im-
pedes the paleoecological reconstruction. However, general
assumptions on living conditions of the representative gen-
era may partly compensate for the absence of data and yield
reliable conclusions. Thus, in this paper, indicative genera
were selected, and their occurrence may be associated with a
particular type of environmental conditions.
In the ostracod fauna of the surroundings of Pokupsko,
heavily calcified genera with strongly ornamented valve sur-
faces are dominant, indicating shallow marine habitats.
These include Aurila, Senesia, Tenedocythere, Callisto-
cythere, Costa, Grinioneis, Hemicytherura, Cytheridea,
Xestoleberis, Semicytherura and Olimfalunia.
Within this shallow-water fauna, two specific groups of
genera may be distinguished. The species within the first
group are limited to tropical and subtropical seas, and most
frequently suggest reefal areas. These include Paranesidea,
Loxoconcha and Cytherelloidea. The second group is distin-
guished by the Miocyprideis genus, indicating a brackish en-
vironment.
The transitional, shallow-water/deep-water group is identi-
fied by the following genera: Pterygocythereis, Echino-
cythereis, Bosquetina, Cytherella and Ruggieria.
A separate group is represented by the deep-water genera
Krithe, Parakrithe, Eocytheropteron, Henryhowella, Argil-
loecia, Acanthocythereis, Paracypris and Bythocythere,
whose recent representatives live at a depth of about 100 m.
On the basis of the different genera identified, the ostracod
fauna from the surroundings of Pokupsko can be defined as a
shallow-water marine fauna, a shallow-water brackish-ma-
rine fauna, a shallow-water reef marine fauna, and a deep-
water marine fauna. Note that the majority of samples
contain predominantly mixed fauna.
The dependence of ostracods on environmental conditions
can be very helpful in the paleoenvironment reconstruction.
The analysis of the entire fauna suggests that the ostracod
fauna of the Pokupsko surroundings lived in a shallow ma-
rine environment, down to 50 m depth, and had a connection
with the open sea. The water was warm, as indicated by find-
ings of several warm-water new-entry species, as well as by
an abundance of the foraminifer Borelis (the Pokupsko sam-
ple), which prefers very warm water and implies tropical to
subtropical water temperatures. The species is also found in
a similar ostracod assemblage from the Middle Miocene de-
posits of North Africa (Szczechura & Abd-Elshafy 1988).
The majority of ostracods are robustly built, indicating
high-water energy and well-oxygenated water conditions,
except for the Šljivovac sample that contains pyritized
valves. The pyritized valves in a sample of the Middle Bade-
nian age agree well with the conclusions of Šútovská (1990),
who, in the Middle Badenian of the Carpathians, assumes the
occurrence of an anoxic facies although; stratification is not
fully developed due to shallow-water depths.
Salinity of the water was similar to the salinity of the
present-day seas. In closed lagoons it was higher, and in
near-shore areas lower due to freshwater influxes.
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Lithologies of the analysed samples (clay, sandy clay,
marly sand, and sand) reinforce the conclusions obtained by
the microfauna analyses and suggest a tripartite division of
the marine environment into a near-shore zone, transitional
zone and shelf.
The near-shore zone limits the estuaries and semi-open la-
goonal environments shorewards. The near-shore environ-
ment is wave-influenced, especially during storms. Locally,
it is also influenced by tides, wind-driven currents, and
freshwater influx. This type of environment is moderately
stable, characterized by moderately high-energy, with sand,
silt and clay as typical sediment types.
The transitional zone is made up of commonly alternating
fine-grained sand, silt, and clay, and is characterized by
maximum development of both the species and individuals.
The environments more distal from the shore are beyond the
river-induced freshwater influence and wave motion, except
during storms. This is a stable, low energy environment.
The ostracod fauna of the shelf environment varies from
poor to rich; the sediment is clayey, partly sandy or silty.
Each of the above zones, striking parallel to the shore, has
its own “inhabitants” that may be grouped into ecozones that
coincide with the above mentioned depth distribution of os-
tracods.
Given all the available data, the paleoenvironment of the
Pokupsko ostracods was probably shallow marine and often
turbulent and periodically connected to the open sea and
freshwater influx.
Conclusion
A very rich and diverse fauna of ostracods has been identi-
fied within the Badenian layers at seventeen locations
around Pokupsko.
The ostracods belong to the suborders of Platycopa and
Podocopa, and their families of Cytherellidae, Bairdiidae,
Cyprididae, and Cytheridae. Forty-four genera and sixty-six
ostracod species have been identified altogether. The entire
ostracod fauna has locally been classified into six biofacies
based on similarities/differences between different faunas.
The Lower Badenian Zone NO7 Acanthocythereis hystrix—
Bythocypris lucida, encompasses the samples of the first bio-
facies and was identified based on findings of the index
species Acanthocythereis hystrix, “new-entry” species Rug-
gieria ex gr. carinata, and Cytherelloidea sissinghi, and nu-
merous deep-water ostracods. According to the faunal
composition and comparison with the same biozone recog-
nized in other areas of the Paratethys, it is presumed that
samples from Pokupsko belong to the youngest part of the
NO7 Zone.
The second Biozone NO8 (Middle Badenian) Eocytheropte-
ron inflatum—Olimfalunia spinulosa, continuously overlies the
previous zone. The samples of the second, third, and fourth
biofacies were included in this biozone. This is based on identi-
fication of the index species Eocytheropteron inflatum in the
fourth biofacies, the numerous species of Olimfalunia in the
second and third biofacies, as well as the different composition
of ostracod communities in the three biofacies.
The third Biozone NO9 Paijenborchella (E.) laskarevi—
Miocyprideis sarmatica elongate marks the beginning of the
Late Badenian in the Central Paratethys. The samples of the
fifth biofacies belong to this zone; they are also character-
ized by the occurrences of Miocyprideis sp.
The fourth ostracod Biozone NO10 Carinocythereis cari-
nata—Phlyctenophora farkasi, marks the latest Late Bade-
nian in the Central Paratethys. The samples of the sixth
biofacies are assigned to this zone and are the most abundant
in this area. There are certain indications that two subzones
can be distinguished within this zone in the wider area in
Croatia.
As in some other areas, changes in the ostracod distribu-
tion closely follow regional geological events. Transgressive
phases resulted in very diversified faunas, while regressive
phases resulted in a great number of specimens and species.
Diversified ostracod fauna is the main feature of the Lower
Badenian and Middle Badenian samples. The discovery of
“new-entry” species from the Tethys, Cytherelloidea sissinghi
and Ruggieria ex gr. carinata, both of which were not fre-
quent in the Paratethys, shows free connections between the
Tethys and Paratethys areas.
Sea-level drop and the disconnection between the Tethys
and Paratethys caused paleobiogeographic ‘disintegration’.
According to the isotopic studies of Badenian foraminifers oc-
curring below evaporates in the Carpathian Foredeep basin
Peryt D. & Peryt T. (2010) suggest that the interrupted com-
munication of the Paratethys with the ocean was a conse-
quence of eustatic sea-level fall, possibly related to climatic
cooling, and it was coupled with a tectonic closure of the con-
nection with the Tethys. The deep-water ostracods became
less abundant and shallow brackish-marine fauna became
dominant in all paleogeographic domains. Sea-level drop and
the probable isolation of the area in the south-western margin
of the North Croatian Basin resulted in the Badenian freshwa-
ter sediments and coal-bearing horizons (Bakrač et al. 2010).
A regressive trend, which continued in the later Badenian,
resulted in the recovery of numerous shallow-water species.
However, the presence of deep-water species in the Upper
Badenien samples implies oscillations of water level, as well
as occasional penetration of deep-water ostracod fauna in re-
sponse to renewed flooding in the Late Badenian (sensu Haq
et al. 1988).
The analyses of Ca, Mg, Sr and Mn were carried on cara-
paces of the selected Late Badenian genera, including Aurila,
Cytherella, Cytheridea, Bairdoppilata and Tenedocythere.
The results obtained contributed to the reconstruction of the
paleo-environment.
Acknowledgments: The authors wish to thank professors Ana
Sokač, Ivan Gušić and Zlatan Bajraktarević for their helpful
and fruitful suggestions. Particular thanks to Slobodan Miko,
for performing the analysis of trace elements. The authors are
very grateful to Ivo Suša from the Croatian Geological Survey
for technical support, and to Renata Slavković and Zdenka
Barbić from INA, Zagreb, for preparing SEM microphoto-
graphs. The present investigation is part of the research that
was supported by the Department of Geology, Croatian Geo-
logical Survey, and the Ministry of Science, Education and
460
HAJEK-TADESSE and PRTOLJAN
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA, 2011, 62, 5, 447—461
Sports Project Grant No. 181-1811096-1093: “Basic Geo-
logical Map of the Republic of Croatia 1 : 50,000”. For im-
provement of the English text we warmly thank to Ivan
Gušić and Antun Husinec. We are grateful to Elsa Gliozzi
and Irene Zorn for their valuable and helpful reviews.
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