GeolCarp_Vol62_No3_267

www.geologicacarpathica.sk

GEOLOGICA CARPATHICA

, JUNE 2011, 62, 3, 267—278 doi: 10.2478/v10096-011-0021-z

Introduction

The Pannonian Basin is a large internal basin structure with-
in continental Europe surrounded by the Alps, Dinarides,
and Carpathians. It resulted from back-arc basin extension
during the Miocene and subsequent compression during the
Pliocene to Quaternary (Horváth & Cloetingh 1996; Cloetingh

ing the chronostratigraphic division (Papp et al. 1985; Magyar
1995; Rögl 1999; Magyar et al. 1999; Harzhauser et al. 2002,
2004;  Magyar  et  al.  2006,  2007  and  reference  therein).  De-
spite the well-known facts about the paleogeographical evolu-
tion  of  the  Pannonian  Basin,  there  are  still  many  open
questions.  The  scarcity  and  endemism  of  dinoflagellates  as
well  as  calcareous  nannoplankton  in  Lake  Pannon  sediments

Upper Miocene Pannonian sediments from Belgrade (Serbia):

new evidence and paleoenvironmental considerations

LJUPKO RUNDIĆ

, MERI GANIĆ

, SLOBODAN KNEŽEVIĆ

and ALI SOLIMAN

University of Belgrade, Faculty of Mining and Geology, Department of Geology, Kamenička 6, 11000 Belgrade, Serbia;

rundic@rgf.bg.ac.rs; merig@rgf.bg.ac.rs; knezevic.slobodan@gmail.com

University of Graz, Institute for Earth Science, Geology and Paleontology, Heinrichstrasse 26, A 8010 Graz, Austria; ali.soliman@uni-graz.at

(Manuscript received January 6, 2010; accepted in revised form December 16, 2010)

Abstract: The Late Miocene sublittoral marls of the Pannonian Stage (the long-lived Lake Pannon) were studied. From
neotectonic point of view, the investigated area represents a natural border between two different morphostructural do-
mains: the Pannonian Basin to the north and the Peri-Pannonian Realm to the south. More than 20 mollusc and 34 ostracod
species were identified which indicate the upper part of the Lower Pannonian and the lower part of the Middle Pannonian
(“Serbian”) predominantly. The identified dinoflagellate cyst assemblage (21 taxa) hinders assignment of the studied
samples to a Pannonian substage but supports the high endemism of the Pannonian flora. The lithostratigraphical, paleon-
tological, and paleoecological analyses indicate a mesohaline (8—16 ‰), sublittoral ( < 90 m deep) environment of the early
Lake Pannon. The estimated stratigraphic range for the investigated deposits is 9.8—11.4 Ma.

Key words: Late Miocene, Lake Pannon, Belgrade, sublittoral environment, endemism.

Fig. 1. Geological sketch map of the Belgrade City area and the position of geological
cross-sections and the studied boreholes. Note: Quaternary deposits with small thickness
are ommited (the right bank of the Sava River).

et  al.  2006;  Horváth  et  al.  2006).  At
about  the  Middle—Late  Miocene  bound-
ary  (ca.  11.6 Ma),  following  the  demise
of the Central Paratethys Sea a long-lived
Lake  Pannon  was  formed  (Magyar  et  al.
1999;  Piller  et  al.  2007;  Harzhauser  &
Mandic  2008).  It  shows  high  endemism
in both fauna and flora (e.g. Müller et al.
1999;  Harzhauser  &  Piller  2007).  Lake
Pannon  enabled  a  spectacular  adaptive
radiation of molluscs including over 900
described  species  and  many  endemic
genera  (e.g.  the  families  Cardiidae  with
more  than  220  species  and  Dreissenidae
with more than 130 species, Geary et al.
2000).  Among  gastropods,  the  proso-
branch  families  Hydrobiidae  ( ~ 180  spe-
cies)  and  Melanopsidae  ( ~ 100  species)
are  dominant  (Geary  et  al.  2000).  Many
of  the  ostracod,  nannoplankton,  di-
noflagellate, diatom, and other fossil spe-
cies are also recognized as endemic.

During the last couple of decades, many

new data on Lake Pannon have been pub-
lished. Application of an integrated strati-
graphic approach led to modification of
the previously accepted concept concern-

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GEOLOGICA CARPATHICA, 2011, 62, 3, 267—278

made biostratigraphic correlation to the Late Miocene marine
sequences impossible.

The intention of this study was to explain: (1) the general

distribution and structure of the Pannonian sediments from the
Belgrade City area in order to determine the paleogeography
at the time of deposition; (2) paleontological analysis in order
to  determine  biostratigraphic  position  and  the  depositional
environment. In addition, estimated time span is discussed in
terms  of  the  different  chronostratigraphic  divisions  of  the
Late  Miocene  in  the  Pannonian  realm  (Magyar  et  al.  1999;
Harzhauser & Mandic 2008; Harzhauser et al. 2008).

Geological framework

More than 70 % of all Miocene sediments in the Belgrade

City  area  (Fig. 1)  correspond  to  the  Pannonian  Stage.  The
Pannonian  deposits  occur  at  the  top  of  the  borehole  sections
under a thin layer of loess sediments and alternating loess de-
posits. In some places, they cover older rocks (Badenian reef
deposits, borehole B-1) or they continue from Sarmatian sand-
stones and sandy limestone. The contact between the Sarma-
tian  and  Pannonian  is  conformable  in  general,  but  there  are
localities  where  the  Pannonian  deposit  lies  transgressively
over  the  Sarmatian  sediments.  In  the  City  centre,  numerous
shallow  boreholes  and  outcrops  revealed  Pannonian  deposits

overlain by a thin cover of loess deposits (Knežević & Šumar
1993, 1994). Along other localities and the street outcrops,
very similar stratigraphic positions of the investigated deposits
were noted (Krstić 1973). The total thickness of the Pannonian
sediments is more than 50 m (Stevanović 1977; Eremija
1989).

Material and methods

The data acquired from the different core samples of the

PdUS set of boreholes have been used for the construction of
two geological cross-sections (Table 1, Fig. 2). Stratigraphic
correlation of the Pannonian deposits from an additional three
boreholes is shown in Fig. 3.

Molluscs were analysed from fifteen core samples of the

boreholes B-1 (Kalemegdan), ZV-3 (Zeleni venac), PdUS-3
and PdUS-7 (Sava River banks).

For analysis of the ostracods, seven dried samples from

borehole  ZV-3  were  washed  using  dilute  hydrogen  peroxide
and  standard  63—500 µm  sieves.  The  specimens  were  picked
out from the residue and stored in the collection of the Depart-
ment  of  Geology  and  Paleontology,  Faculty  of  Mining  and
Geology, University of Belgrade.

Four samples from the borehole ZV-3 were selected for the

contents of dinoflagellate cysts. Approximately 15 grams

Fig. 2. Geological cross-sections through the Miocene sediments near the Sava-Danube confluence: A—A’ – along the left bank of the
Sava River, B—B’ – across the Sava River, K

– Upper Cretaceous, Bd – Badenian limestone and sand, Sm – Sarmatian sand, marl and

limestone, Pn – Pannonian marl and silty marl, Q – Loess and other soft deposits.

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were  treated  with  cold  HCl  (34 %)  to  remove  carbonates.
Then after washing with distilled water, the residue was treat-
ed with HF (48 %) and cold HCl to fully remove silicates and
colloids. The residue was ultrasonicated (ca. 30°) and sieved
at  125 µm  and  20 µm.  No  oxidation  at  all  was  applied.  The
residue was washed and stained with Safranine “O”. Two mi-
croscope  slides  were  made  from  each  sample  using  glycerin
jelly  as  the  mounting  medium.  The  first  300  dinoflagellate
cysts were counted using a Zeiss Axioplan 2 microscope fitted
with a Leica DFC 320 digital camera. Additionally, two SEM
stubs  were  prepared  from  a  sample  of  21.60 m  and  scanned
with  DSM  Gemini  SEM  operating  at  a  working  voltage  of
10 kV.  The  dinoflagellate  cyst  nomenclature  generally  fol-
lowed that of Fensome et al. (2008), namely, the new online
version of DINOFLAG2, http://dinoflaj.smu.ca.

Structural and stratigraphic setting

The first structural stage forms the substrate for the Miocene

and Quaternary cover (Fig. 2) in Belgrade City and its vicini-
ty. It occurs on elevated structures such as small highs repre-
sented  by  Jurassic  serpentinite;  Jurassic  and  Cretaceous
carbonate rocks and rare igneous rocks. The second structural
stage is composed of Miocene deposits, the most widespread
rocks in the investigated area on the right bank of the Sava and
Danube  Rivers.  The  last  structural  stage  corresponds  to  the
Quaternary sediments. These are various alluvial deposits on
the left bank of the Sava River, covering the older formations.
The older rocks have very complex structural settings as a re-
sult of long-range tectonic activities (Marović et al. 2007). For
these reasons, there are big differences in the core sections be-
tween boreholes PdUS-1, PdUS-2, and PdUS-3 (Fig. 2). Core
analysis  of  the  last  borehole  as  well  as  the  borehole  UDP-1
showed complete successions of the Badenian, Sarmatian, and
Pannonian  deposits  (Fig. 3).  On  the  contrary,  a  prior  study
shows that the Miocene sediments are thinner and the Sarma-
tian sediments are missing (Knežević & Ganić 2008).

Detailed analysis of the geological cross-section B-B’ in the

NE direction (across the Sava River, borehole PdUS-1 to the
Kalemegdan Fortress) showed that there was a strong vertical
movement of up to 75 m along the fault zone. Similar data
was obtained by core analysis of other PdUS boreholes. How-
ever, it is noticed that the sinking is more evident towards the
center of the basin (borehole PdUS-7). It could be supposed
that there was a cascade fault system that separated different
rock units and shifting along the main fault that followed the

Table 1: Geographical position of the investigated boreholes.

No. Boreholes

Coordinates

(WGS84)

PdUS-1

N 44º 49' 00.8"

E 20º 26' 41.5"

PdUS-2

N 44º 49' 07.0"

E 20º 26' 38.2"

PdUS-3

N 44º 48' 57.4"

E 20º 26' 42.4"

PdUS-5

N 44º 49' 18.3"

E 20º 26' 28.0"

PdUS-6

N 44º 49' 06.1"

E 20º 26' 59.3"

PdUS-7

N 44º 48' 58.2"

E 20º 26' 38.9"

UPD-1

N 44º 48' 59.8"

E 20º 26' 29.9"

B-1

N 44º 48' 57.4"

E 20º 26' 42.4"

ZV-3

N 44º 48' 48.9"

E 20º 27' 24.3"

Fig. 3. Stratigraphic columns of the investigated boreholes and po-
sition of the Pannonian deposits. For stratigraphic abbreviations,
see Fig. 2.

Sava River. In addition, a transversal fault cuts pre-Quaternary
sediments.  It  was  documented  by  longitudinal  cross-section
analysis  (A—A’,  Fig. 2)  between  the  boreholes  PdUS-3  and
PdUS-5.  A  very  marked  subsurface  horst-like  structure  was
identified  by  core  analysis  of  borehole  PdUS-2.  Away  from

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both sides of this location, there is a trend towards increased
thickness of the Miocene deposits.

In the lower part of borehole B-1 (Fig. 3), the Pannonian

marly limestone and marl transgressively lay over the Middle
Miocene  Badenian  reef  sediments.  The  inhomogeneous  marl
has a thickness of up to 10 m. The uppermost part of that marl
section is marked by marly clay. Core samples from the bore-
hole  ZV-3  represented  by  silty  marl  contain  very  scarce  and
damaged molluscs (depth ca. 17—24 m). Above, there is a 2 m
thick  zone  of  deformed  and  altered  silty  marl.  Core  section
analysis  shows  that  the  maximal  thickness  of  the  Pannonian
sediments is up to 30 m as recorded in the borehole UDP-1.

Paleontological analysis

For the appropriate biostratigraphic and paleoenvironmental

interpretations, the Late Miocene Lake Pannon division after
Magyar et al. (1999) was used as well as the modern chronos-
tratigraphic and biostratigraphic correlation scheme for the
Mediterranean and Paratethys (Gradstein et al. 2004; Vasiliev
et al. 2004; Kováč et al. 2006; Piller et al. 2007; Harzhauser &
Mandic 2008 – Fig. 4).

Molluscs

New data obtained by stratigraphic and paleontological

analysis of fifteen core samples from boreholes B-1 (Kale-
megdan), ZV-3 (Zeleni venac), PdUS-3 and PdUS-7 from
both sides of the Sava River are discussed here. Although cer-

tain specimens were put in open nomenclature, 20 mollusc
species were determined.

The Pannonian marl in the borehole B-1 contains a poor but

very characteristic mollusc association with Paradacna cekusi
(Gorjanović-Kramberger),  Gyraulus  praeponticus  (Gorjano-
vić-Kramberger),  Congeria  sp.,  Micromelania  sp.  Shells  are
relatively  small,  broken  and  without  prevailing  orientation.
Biostratigraphically, another more important assemblage was
recorded.  It  corresponds  to  the  uppermost  part  of  the  Lower
Pannonian where Congeria banatica  (R. Hoernes), Paradac-
na syrmiense (R. Hoernes),  Parvidacna laevicostata (Wenz),
Orygoceras fuchsi (Kittl) and Gyraulus cf. praeponticus (Gor-
janović-Kramberger) were recognized.

Most evidence concerning the Pannonian molluscs was col-

lected from boreholes PdUS-3 and PdUS-7. Ten core samples
from  both  boreholes  were  analysed.  The  basal  part  of  the
Pannonian in the borehole PdUS-3 is represented by grey marl
with abundant small limnocardids such as Limnocardium maor-
ti  Strausz,  L.  winkleri  lukae  Stevanović,  L.  gr.  praeponticum
Gorjanović-Kramberger, Paradacna lenzi (R. Hoernes), P. syr-
miense (R. Hoernes), as well as Mytilopsis zujovici Brusina and
Congeria banatica (R. Hoernes). Above the fossiliferous marl,
there is a thin marly bed with imprints of different small limno-
cardids without prevailing orientation of the shell remains. The
upper  part  contains  grey,  silty  marl  with  Mytilopsis  subdigiti-
fera  (Stevanović),  Paradacna  syrmiense  (R.  Hoernes),  Para-
dacna lenzi (R. Hoernes), Paradacna sp., Monodacna vienensis
Papp,  and  Gyraulus  praeponticus  (Gorjanović-Kramberger).
Additionally,  it  contains  numerous  Mytilopsis  czjzeki  M.
Hoernes and Congeria zsigmondyi Halaváts (Figs. 5, 6).

Fig. 4.

strati-

graphic correlation
scheme for the Mio-
cene

Central

Paratethys and the
Lake Pannon bio-
stratigraphy

(after

Magyar et al. 1999;
Gradstein

al.

2004;  Kováč  et  al.
2006;  Piller  et  al.
2007;  Harzhauser  &
Mandic  2008  –
modified).

Time

span  for  the  Pontian
and  Dacian  accord-
ing to Vasiliev et al.
(2004).  A  light  grey
area marks the strati-
graphical  range  of
the investigated Lake
Pannon deposits.

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Fig. 5. PdUS-3 borehole section and distribution of the mollusc fau-
na. 1 – marls, 2 – mollusc shells.

Fig. 6. Molluscs from the borehole PdUS-3: A – Mytilopsis zujovici,
B – Mytilopsis czjzeki. Scale bar: 1 cm.

Pannonian sediments in the lower part of borehole PdUS—7

(Figs. 7,  8)  start  with  Undulotheca    pancici  Brusina  and
Mytilopsis gr. czjzeki M. Hoernes. From the upper parts of this
borehole,  three  marly  samples  were  taken.  The  lowermost
one, greyish blue marl contains abundant specimens of Para-
dacna  lenzi  (R.  Hoernes),  Congeria  banatica  (R.  Hoernes),
and Mytilopsis czjzeki M. Hoernes. The central part contains a
small  lens  with  Paradacna  cekusi  (Gorjanović-Kramberger),
Parvidacna  laevicostata  Wenz,  and  Gyraulus  praeponticus
(Gorjanović-Kramberger)  and  finally,  the  uppermost  one,
silt  and  sandy  silt  has  numerous  Paradacna  syrmiense  (R.
Hoernes)  and  Paradacna  cekusi  (Gorjanović-Kramberger).
As  opposed  to  borehole  PdUS-3,  it  was  noticed  that  the
marker  zone  of  the  Middle  Pannonian,  M.  czjzeki—C.  zsig-
mondyi is missing here.

Biostratigraphically, the above-mentioned associations of

molluscs correspond to the M. czjzeki Zone (Magyar et al.

1999). Moreover, this zone can be divided into two subzones:
a) the lower, small limnocardids subzone with Limnocardium
gr. praeponticum, L. winkleri lukae, L. maorti, Paradacna
lenzi, etc. and b) the upper, czjzeki—zsigmondyi Subzone. Ac-
cording to the previous biozonation, the investigated Pannon-
ian sediments from the Belgrade City area correspond to the
zones B, C, D and the lowermost part of zone E (Papp 1953;
Papp et al. 1985).

Ostracods

The current study of seven silty marl samples taken between

17.10 to 21.60 m of the upper part of the borehole ZV-3 made
it possible to identify 34 species of ostracods (see position in
Fig. 1).  Core  sediment  shows  minor  evidence  of  taphonomic
processes  (e.g.  bioturbations,  pyrite  occurrences).  The  ostra-
cod  association  consists  of,  mainly,  numerous  candonids,  in-

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cluding Reticulocandona reticulata (Méhes), Zalanyiella ruri-
ca Krstić, Z. venustoidea Krstić, Z. buchii Krstić, Serbiella cf.
bacevicae Krstić, S. maxiunguiculata Krstić, Turkmenella sp.,
Typhlocyprella cf. ankae Krstić, T. cf. lineocypriformis Krstić,
Typhlocyprella  sp.,  Fabaeformiscandona  lineata  Krstić,
Camptocypria subpontica Krstić, C. praebalcanica Krstić, C.
alasi  alasi  Krstić,  Serbiella  sagitosa  Krstić,  S.  unguiculus
(Reuss).  Most  of  them  have  a  thin,  more  or  less  transparent,
and  elongated  carapace  (e.g.  Camptocypria,  Typhlocyprella,
Zalanyiella, and Serbiella). Among cytherids, the genus Cyp-
rideis is represented by three taxa (C. longa Krstić, C. longi-
testa  Krstić,  and  C.  cf.  brevis  Krstić)  that  dominate  the
assemblages. All the other species of ostracods make another
important  part  of  the  Pannonian  assemblages  in  these  core
samples. They are represented by large Hungarocypris hiero-
gliphica  (Méhes)  –  genus  Herpetocyprella  after  Danielopol
et al. (2007) as well as Amplocypris major Krstić, Hemicythe-
ria  croatica  Sokač  and  H.  marginata  Sokač.  Small  forms  of
Loxoconcha  rhombovalis  Pokorny,  L.  granifera  (Reuss),  L.
fistulosa  Krstić,  L.  subrugosa  Zalanyi,  Loxocorniculina
hodonica Pokorny, Amnicythere naca (Méhes), A. lacunoidea
Krstić, A. larga Krstić, A. cf. stanchavae Krstić, Cypria dorso-
concava  Krstić,  C.  cf.  siboviki  Krstić  are  present,  too.  Adult
valves of the two large species H. hierogliphica and A. major

are numerous and strongly calcified in the upper core samples.
Different  Typhlocyprella  species  are  recorded  in  the  upper-
most  sample.  In  general,  preservation  of  valve/carapace  is
good. There are more adult specimens than juvenile ones. The
above-mentioned candonids are the most diversified ostracod
group  and  represent  more  than  50 %  of  the  total  number  of
species  found.  Biostratigraphically,  the  determined  ostracods
suggest the  Amplocypris abscissa and Hemicytheria croatica
Zones  (Krstić,  1985)  or  older  part  of  the  Middle  Pannonian
(Serbian,  sensu  Stevanović  1985).  Some  of  these  Pannonian
ostracods are shown in Fig. 9.

Dinoflagellates

The four investigated samples are very rich in dinoflagellate

cysts, however, the diversity is low and the encountered as-
semblage is quite similar in all samples. Spiniferites, Impagi-
dinium, and Pyxidinopsis were the dominant dinoflagellate
cysts with considerable occurrences of heterotrophic taxa (e.g.

Fig. 8. Molluscs from the borehole PdUS-7: A – Congeria banatica,
B – Undulotheca pancici. Scale bar: 1 cm.

Fig. 7. PdUS-7 borehole section and distribution of mollusc fauna.
1 – marls, 2 – mollusc shells.

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Fig. 10. Pannonian dinoflagellates from the borehole ZV-3. All microphotographs are from the borehole ZV-3 (depth 20.60 m). The scale is
20 µm. 1, 2, 8 – Spiniferites bentorii subsp. pannonicus Sütőné Szentai, 1986; right-lateral view. 2 – Spiniferites bentorii subsp. pannonicus
Sütőné Szentai, 1986; left-lateral view, note the weak parastures. 3 – Impagidinium sphaericum (Wall) Lentin & Williams, 1981; dorsal view,
a specimen with fine surface ornamentation. 4 – Impagidinium sp.; later view, note the coarse reticulate surface. 5 – Pyxidinopsis psilata
(Wall & Dale) Head, 1994; lateral view. 6 – Spiniferites sp.; dorsal view, a specimen with robust processes. 7 – Spiniferites bentorii subsp.
budajenoensis Sütő-Szentai, 1986; ventral view. 8 – Spiniferites bentorii subsp. pannonicus Sütőné Szentai, 1986; dorsal view. 9 – Nema-
tosphaeropsis sp., uncertain orientation. 10 – Achomosphaera argesensis Demetrescu, 1989; ventral view. 11 – Impagidinium sp.; oblique
antapical view. 12 – Impagidinium spongianum Sütő-Szentai, 1985; crumpled specimen in ventral view. 13 – Impagidinium sp.; oblique
antapical view, note the coarse gammae on the surface. 14 – Spiniferites tengelicensis Sütő-Szentai, 1982; dorsal view. 15 – Spiniferites
bentorii subsp. oblongus Sütőné Szentai, 1986; dorsal view. 16 – Spiniferites bentorii subsp. oblongus Sütőné Szentai, 1986; ventral view.
17 – Impagidinium sp.; apical view, note the prominent apical boss. 18 – Impagidinium sphaericum (Wall) Lentin & Williams, 1981; dorsal
view, a specimen with coarse surface ornamentation and apical boss. 19 – Impagidinium sphaericum (Wall) Lentin & Williams, 1981; ob-
lique antapical view, a specimen with a smooth surface. 20 – Botryococcus colony.

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Selenopemphix spp. and “small brown round cysts” (Brigante-
dinium  spp.)),  Achomosphaera  spp.,  Protoperidinium  sp.,
Nematosphaeropsis  sp.,  and  Komewuia  sp.  (see  Appendix).
Sporadic  occurrences  of  the  freshwater  green  algae,  Pedias-
trum,  Botryococcus  and  fungal  spores  are  recorded.  On  the
other hand, saccate and non-saccate pollen were common but
no attempt has been made to identify them. No marker taxa for
the Upper Miocene similar to those contemporaneously recov-
ered  from  the  circum-Mediterranean  and  other  regions  have
been  recorded.  This  could  go  back  to  the  Upper  Miocene
(Pannonian),  however,  the  Central  Paratethys  is  highly  en-
demic in both fauna and flora and there was no seaway con-
nection with the adjacent basins (Magyar et al. 1999; Piller et
al.  2007;  Harzhauser  &  Mandic  2008).  The  encountered  as-
semblage  is  quite  similar  to  the  recorded  dinoflagellate  cyst
assemblages  from  many  sections  and  drill-holes  in  Hungary
(e.g. Sütőné-Szentai 2000), and the Vienna Basin (Harzhauser
et al. 2008). Thus, a Early Pannonian age could be suggested
for  the  investigated  samples  (Fig. 10).  Additionally,  the  ab-
sence  of  Spiniferites  paradoxus  could  confirm  the  suggested
age (Magyar et al. 1999).

Discussion

The whole area near the Sava—Danube confluence repre-

sents the marginal zone between two different geographical
and morphostructural entities: 1) The Great Pannonian Plain
that is located on the left bank of the Sava River and, 2) the
Balkan Peninsula terrain on the right bank of the Sava River
(Marović & Knežević 1985). It is generally accepted that the
structural  setting  and  morphological  features  of  the  investi-
gated  area  were  consequences  of  younger  phases  of  Alpine
tectogenesis (the so-called Rodanian and the Vlaška phases,
Marović & Knežević 1985). Radial deformations (faults and
differential  fault  block  movements)  were  more  significant
than the plicative structures. As a result, during the Pliocene
and  Pleistocene,  a  great  area  of  relatively  depressed  blocks
was  established  in  the  Pannonian  Plain,  opposite  relatively
elevated  terrains  with  more  morphostructural  forms  –  the
Šumadija  Hills  (Marović  et  al.  2002).  The  studied  area  be-
longs to the complex horst of the Vardar Zone with a SSE—
NNW  direction  (Marović  et  al.  2007).  On  the  eastern  and
western  sides  of  this  structure,  there  are  two  graben-like
structures,  which  are  recognized  as  subsiding  zones  during
the Miocene: 1) the Velika Morava Graben in the east and 2)
the  Kolubara-Tamnava  Graben  in  the  west  (Marović  &
Knežević  1985;  Marović  et  al.  2007).  This  very  complex
structure  was  separated  by  faults  into  numerous  local  gra-
bens and small horsts during its evolution through the Mio-
cene.  Uplifted  horst  structures  were  affected  by  erosion
(Marović et al. 2002).

During the Pannonian age, the water level of Lake Pannon

increased due to regional isolation and the Belgrade City area
was a zone of maximal flooding (Magyar et al. 1999). Accord-
ing to the new interpretations, the early stage of Lake Pannon
(ca. 11.6 Ma) was still influenced by the latest Middle Mio-
cene dry climate (e.g. Harzhauser et al. 2004). This phase co-
incided with a pronounced radiation of melanopsid gastropods

(Magyar et al. 1999; Harzhauser & Mandic 2008). During the
warm early Late Miocene (ca. 10 Ma) humidity increased and
culminated in a phase with high summer precipitation (Bruch
et al. 2007; Utescher et al. 2007). This caused a reorganization
of the coastal-deltaic faunas, suppressing the radiation of mel-
anopsids    (Harzhauser  &  Mandic  2008).  On  the  other  hand,
high  nutrient  loads  favoured  the  dispersion  of  filter-feeding
dreissenids  (Harzhauser  et  al.  2007;  Harzhauser  &  Mandic
2010). Despite continually declining salinity, Lake Pannon re-
mained an alkaline lake (Harzhauser et al. 2007). Similar sedi-
mentological data from the Kolubara Basin (ph and Eh values)
indicate a slightly alkaline environment (Rundić 2006).

More than 60 mollusc species from the different Pannonian

levels inside the Belgrade City’s centre have been recorded by
our  study  and  previously  published  data.  According  to  the
overall biostratigraphical range of the investigated species as
well  as  comparison  with  the  magnetostratigraphic  data
(Magyar et al. 1999, 2007), the total estimated time span for
all the investigated samples from 9.8 to 11.4 Ma includes the
Early and Middle Pannonian (“Serbian”). Small limnocardids
(Paradacna  cekusi,  Parvidacna  laevicostata,  Limnocardium
praeponticum)  and  snails  (Radix  croatica  and  Gyraulus  cf.
praeponticus) form the basis of the Early Pannonian molluscs
(Magyar et al. 2007). They were found in the marly limestone
and  marl  (borehole  B-1).  The  occurrence  of  limestone  sug-
gests a shallow lake environment without considerable influ-
ence  of  the  land  area  whilst  marl  was  deposited  from
suspension in a sublittoral zone. The appearance of numerous
pulmonate gastropods and small limnocardids, which live on
water  grasses  as  epibionts,  suggest  a  high  degree  of  aeration
and  desalinization  of  the  lake.  This  association  have  a  very
high  endemicity  ( > 86 %,  Harzhauser  &  Mandic  2004).  It  is
correlative with the evolution, species number and the size of
gastropod shells within the Lake Pannon Phase I (Harzhauser
& Mandic 2008). Stevanović (1957) gave the evolution of the
small  limnocardids  (recently  accepted  as  a  separate  genus
Lymnocardium)  and  concluded  that  they  originated  from  the
Sarmatian  cardids  (genus  Cerastoderma).  Later,  Vrsaljko
(1999) showed their phylogenetic lineage based on the num-
ber and type of ribs (however, Mandic et al. (2008) assigned
these  forms  as  endemic  genera,  such  as  Obsoletiforma,
Plicatiforma,  etc.).  According  to  Stevanović  (1977),  occur-
rence of Mytilopsis zujovici, Limnocardium lukae and Radix
croatica indicate a sublittoral lake deposition of water depth
between  15  and  80  meters.  Similarly,  Tarasov  (1997)  esti-
mated the lower limit of the sublittoral zone in the Caspian
Sea at  80—100 m. The presence of Congeria banatica, Para-
dacna  syrmiense,  Paradacna  lenzi  and  Undulotheca  pancici,
as well as Gyraulus praeponticus can suggest a quiet and sta-
ble regime in generally sublittoral condition. Additionally, ab-
sence  of  prevailing  orientation  of  shells  excludes  a  dynamic
water  system.  Similar  salinity  conditions  were  reported  by
Korpás-Hódi (1983) who designated the Late Miocene assem-
blage with Mytilopsis czjzeki—Paradacna abichi as pliohaline
environment (9—16 ‰).

Relatively diversified and abundant ostracods correspond to

the Middle Pannonian – Serbian (Krstić 1972, 1973; Rundić
1998, 2006). This period was assigned as the “bloom time” for
many ostracods, both in terms of diversity and abundance. It

276

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GEOLOGICA CARPATHICA

GEOLOGICA CARPATHICA, 2011, 62, 3, 267—278

corresponds  to  the  first  appearance  of  some  genera  such  as
Zalanyiella,  Camptocypria,  Sinegubiella,  etc.  (Krstić  1985;
Rundić 2006) which radiated during the Late Pannonian. Os-
tracods might settle the environment of high nutrient produc-
tivity  (e.g.  estuaries  along  the  southern  Baltic  Sea  coast,
Frenzel & Boomer 2005). The elongated forms of candonids
with thin carapace can indicate quiet and relatively stable wa-
ter  conditions  (Rundić  2006).  The  brackish  species  (Hemi-
cytheria,  Loxoconcha)  as  well  as  more  freshwater  and
river-marsh genera with a strong massive carapace (Herpeto-
cyprella,  Amplocypris)  also  suggest  a  mesohaline,  infralit-
toral/sublittoral  environment  (Rundić  2002).  According  to
Meisch (2000), the recent Cyprideis torosa (Jones, 1850) sug-
gests a wide range of salinities, from 2—16.5 ‰. It is a typical
euryhaline species (Boomer & Eisenhauer 2002; Rostovtseva
& Tesakova 2009). Similar observations and discussion about
environmental  changes  and  diversification  of  Late  Miocene
Cyprideis  from  the  Lower  Pannonian  of  the  Styrian  Basin
have been  reported  by Gross (2008)  and  Gross  et  al. (2008).
Analogous forms of Amnicythere species are living in the sub-
littoral zones of the Black Sea and the Caspian Sea in brackish
environments  as  well  as  in  more  freshwater  bays  (Cziczer  et
al. 2009). Similar to this, recent Cypria species are generally
freshwater  forms  but  can  tolerate  mesohaline  conditions
(Meisch 2000; Starek et al. 2010). There is other evidence that
confirms this model (see the occurrence of different molluscs).
There are no ostracod species that would indicate an oligoha-
line  environment,  such  as  Ilyocypris,  Cyprinotus,  and  Vesta-
lenulla  (Gross  2008)  which  are  well  known  in  the  Late
Pannonian from other parts of the southern margin of the Pan-
nonian Basin (Sokač 1972; Krstić 1972; Rundić 2006).

The development of the flora within the lake was controlled

by  its  (bio)geographic  restriction  as  well  as  by  the  gradual
freshening of the water body (Harzhauser et al. 2007). These
environmental conditions led to the evolution of endemic spe-
cies of gonyaulacoid dinoflagellate cysts, such as Spiniferites
and Impagidinium. Both are well recorded in the Upper Bade-
nian  (Jiménez-Moreno  et  al.  2006),  however,  they  exhibit  a
much higher morphological variability in the Pannonian. This
variability  has  been  used  to  introduce  several  endemic  spe-
cies/subspecies  (e.g.  Sütő-Szentai  1985).  The  dominance  of
Spiniferites  and  Impagidinium  taxa  with  their  characteristic/
endemic  morphotypes,  characterized  by  the  presence  of  a
well-developed  apical  boss  and  variable  surface  ornamenta-
tion,  could  indicate  a  change  in  the  water  chemistry  of  the
Pannon Lake (Harzhauser et al. 2007). In addition, the abun-
dance of heterotrophic taxa, especially of Selenopemphix and
“small round brown cysts” (cf. Brigantedinium), indicates nu-
trients-rich  surface  waters.  Besides,  the  presence  of  Pyxidi-
nopsis psilata, the green algae, Pediastrum, and Botryococcus
indicates a freshwater input to the basin as well as a brackish
water environment (Marret et al. 2007).

Conclusion

The investigated Pannonian sediments from the Belgrade

City area represent a sublittoral environment of the early Lake
Pannon (estimated time interval: 9.8—11.4 Ma). They are blue-

grey,  sandy  and  silty  marls  belonging  to  the  Limnocardium
praeponticum and  Mytilopsis czjzeki mollusc Zones (Magyar
et  al.  1999)  that  correspond  to  the  Lower  Pannonian  and  the
Middle  Pannonian  (Serbian,  sensu  Stevanović  1985).  On  the
basis  of  the  ostracod  biozonation,  the  investigated  sediments
belong to the Amplocypris abscissa and Hemicytheria croatica
Zones (Middle Pannonian). The endemic dinoflagellate asso-
ciations suggest the Lower Pannonian.

All the analysed fossil assemblages indicate a mesohaline

(8—16 ‰)  sublittoral  environment  within  Lake  Pannon
( < 90 m) that was very similar to the recent sublittoral zones
of the Black Sea and the Caspian Sea. Most of the encountered
species  from  studied  boreholes  and  outcrops  are  considered
endemic.  Brackish  cardids  and  dreissenids  (species  found
here) are typical representatives of the sublittoral assemblages.
Common findings of the basinal form Congeria banatica sug-
gest a calm, stable water regime rather than a deeper, basinal
influence. Among the ostracods, the dominant forms are can-
donids. Their diversity and the numerous presences in the up-
permost parts of some boreholes (ZV-3) indicate a more stable
brackish environment during the early phase of the maximum
extent of Lake Pannon.

All the collected data suggest that the investigated Pannon-

ian sediments of the Belgrade City area represent the older
phase in the deposition of the long-lived Lake Pannon (parts
of phases I and II, Harzhauser & Mandic 2008).

Acknowledgments: The Ministry of Science and Technologi-
cal  Development,  Republic  of  Serbia  (Project  No. 176015)
supported the study. AS wishes to thank the Austrian Acade-
my  of  Sciences  and  FWF-Project  No. P  21414-B16  for  fi-
nancial  support.  We  are  deeply  grateful  to  Tadeusz  Peryt
(Polish  Geological  Institute,  Warszawa),  Martin  Gross
(Landesmuseum Joanneum, Graz) as well as three unknown
reviewers  who  gave  helpful  remarks  and  critical  comments
on the manuscript.

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Achomosphaera ramulifera (Deflandre) Evitt, 1963
Achomosphaera argesensis Demetrescu, 1989
Achomosphaera cf. A. fenestra Kirsch, 1991
Batiacasphaera hirsuta Stover, 1977
Impagidinium sphaericum (Wall) Lentin & Williams, 1981
Impagidinium spongianum Sütő-Szentai, 1985
Impagidinium spp.
Komewuia spp.
Lejeunecysta communis Biffi & Grignani, 1983
Nematosphaeropsis sp.
Pyxidinopsis psilata (Wall & Dale) Head, 1994
Selenopemphix brevispinosa Head, Norris & Mudie, 1989

Appendix

List of the identified dinoflagellate cysts (for taxonomic references see, Fensome et al. 2008)

Selenopemphix sp.
Selenopemphix nephroides Benedek emend. Bujak in Bujak

et al., 1980

Spiniferites bentorii (Rossignol) Wall & Dale, 1970
Spiniferites bentorii subsp. budajenoensis Sütő-Szentai,

1986

Spiniferites bentorii subsp. oblongus Sütőné Szentai, 1986
Spiniferites bentorii subsp. pannonicus Sütőné Szentai,

1986

Spiniferites galeaformis Sütő-Szentai, 1994
Spiniferites tengelicensis Sütő-Szentai, 1982
Spiniferites virgulaeformis Sütő-Szentai, 1994