GeolCarp_Vol62_No3_251

www.geologicacarpathica.sk

GEOLOGICA CARPATHICA

, JUNE 2011, 62, 3, 251—266 doi: 10.2478/v10096-011-0020-0

Miocene vegetation pattern and climate change in the

northwestern Central Paratethys domain

(Czech and Slovak Republic)

MARIANNA KOVÁČOVÁ

, NELA DOLÁKOVÁ

and MICHAL KOVÁČ

Department of Geology and Paleontology, Faculty of Sciences, Comenius University, Mlynská dolina, 842 15 Bratislava,

Slovak Republic; kovacova@fns.uniba.sk; kovac@fns.uniba.sk

Institute of Geological Sciences, Masaryk University, Kotlářská 2, 611 37 Brno, Czech Republic; nela@sci.muni.cz

(Manuscript received October 13, 2010; accepted in revised form December 16, 2010)

Abstract: The case study area covers the slopes of the tectonically quiet European platform and foreland of the tectonically
active Carpathian mountain chain (Carpathian Foredeep and Vienna Basin). Therefore the research on pollen spectra
mirrors not only the evolution of landscape in two areas with different geodynamics, but also climatic changes in the
Central Paratethys domain during the studied time interval. According to the pollen data, the Early to Middle Miocene
vegetation reflects subtropical climate with very mild (negligible) cooling events during this period. This is indicated by
common occurrence of thermophilous taxa in the whole sedimentary record. The Middle Miocene landscape evolution,
conditioned by uplift of the Carpathian mountain chain and subsidence of adjacent lowlands, led to commencement of the
altitudinal zonation. The terrestrial and aquatic ecosystems confirm a subtropical climate (Miocene Climatic Optimum,
Mi3 event) with some possible long term changes in humidity. The Late Miocene paleogeographical changes, but also
general climatic oscillations in the northwestern Central Paratethys realm, resulted in decrease of the number of thermo-
philous taxa during this time (change in latitudinal position of the vegetation cover). Variously high mountain relief of
the uplifted mountain chains (altitudinal zonality) created ideal conditions for mixed mesophytic forests (to open wood-
land – open grassland type), still with presence of evergreen taxa. A subtropical climate with gradual transition to
warm temperate climatic conditions is supposed on the basis of the reconstructed vegetation cover.

Key words: Miocene, Paratethys, Carpathian Foredeep, Vienna Basin, paleoclimate, palynology.

Introduction

The  Miocene  vegetation  pattern  and  climatic  changes  were
studied by means of palynology in the northwestern part of the
Central  Paratethys  domain  (Fig. 1).  To  determine  changes  in
vegetation  pattern  (altitudinal  zonation)  and  influence  of  the
global  climatic  changes  (latitudinal  zonation)  two  areas  with
different geodynamics and therefore also with different land-
scape  evolutions  have  been  choosen.  To  the  West  there  was
the  tectonically  quiet  Variscan  Bohemian  Massif,  and  to  the
East the neo-Alpine tectonically active uplifting Western Car-
pathian mountain chain. The samples were taken from marine,
brackish to freshwater sediments of the Carpathian Foredeep
(Czech  Republic)  and  Vienna  Basin  (Czech  and  Slovak  Re-
public)  in  the  time  interval:  Early  to  Late  Miocene—Eggen-
burgian  to  Pannonian  (Burdigalian  to  Tortonian,  sensu
Harzhauser & Piller 2007). The analysed sediments were well
biostratigraphically  dated  (Hladilová  1988;  Nehyba  et  al.
1997; Doláková et al. 1999; Rögl et al. 2003; Hudáčková et al.
2003; Kováč et al. 2004, 2006, 2007, 2008).

Paleogeography

The study area – the contact zone between the North Eu-

ropean Platform and Western Carpathian orogen had a very
complicated Neogene geodynamic history (e.g. Royden

1985,  1988;  Ratschbacher  et  al.  1991a,b;  Kováč  &  Hók
1993; Lankreijer et al. 1995; Meulenkamp et al. 1996; Nehyba
et al. 1997; Kováč et al. 1997, 1998a,b, 2001, 2003; Kováč
2000;  Kvaček  et  al.  2006;  Harzhauser  &  Piller  2007).  All
processes  such  as  subduction,  collision,  back  arc  basin  de-
velopment and its tectonic inversion are recorded in great pa-
leogeographical  changes  (Tomek  &  Hall  1993;  Konečný  et
al. 2002). These changes strongly influenced not only the re-
lation between areas flooded by the sea and continental ar-
eas, but also development of landscape and the evolution of
altitudinal zonation in this region (Fig. 2).

The Early Miocene geodynamic development of the Bohe-

mian  Massif  and  Western  Carpathians  junction  area  was
strongly  affected  by  subduction  of  the  flysch  troughs  base-
ment below the East Alpine—Western Carpathian orogen and
development  of  the  Outer  Western  Carpathian  accretionary
wedge  from  Flysch  Belt  units  (Kováč  et  al.  1997,  1998a;
Kováč 2000; Konečný et al. 2002; Kvaček et al. 2006). Ter-
ritory  in-between  the  platform  and  the  front  of  the  Car-
pathian  orogen  was  covered  by  a  wide  arm  of  the  Central
Paratethys  Sea  during  this  time.  The  Eggenburgian  marine
transgression flooded the Carpathian Foredeep on the slopes
of the Bohemian Massif in the West, continuing eastwards to
the Pouzdřany and Ždánice residual flysch troughs. The east-
ern margin of the Early Miocene sea arm was represented by
the  western  and  northern  parts  of  the  present  Vienna  Basin
(Fig. 2A).  This  sea  arm  represented  a  system  of  particular

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GEOLOGICA CARPATHICA, 2011, 62, 3, 251—266

Fig. 2. Paleogeography of the Central Paratethys (according to Kováč 2000).

with the first Burdigalian global sea-level rise (sensu Vail et
al. 1977; Haq et al. 1988; Haq 1991; Kováč 2000).

At the end of the Ottnangian, the transpressive tectonics

resulted  in  a  partial  uplift  of  the  Alpine-Carpathian  chain,
followed by closing of the sea connections with the Mediter-
ranean in front of the Alps. Short term isolation of the West-
ern  Carpathian  sedimentary  basins  took  place.  This  process
was  accompanied  by  closing  of  the  residual  flysch  troughs
and  folding  and  thrusting  of  their  sedimentary  fill  towards
the platform. Gradual uplift of the Outer Western Carpathian
accretion  wedge  started  (Kováč  et  al.  1998a).  On  the  other
side, initial rifting of the Pannonian back arc basin began in
the Western Carpathians hinterland (Horváth et al. 1988).

The Karpatian transpressive tectonics led to extrusion of

the Western Carpathian orogene (ALCAPA Microplate)
from the Alpine domain (Ratschbacher et al. 1991a,b; Kováč
et al. 1998a). Oblique collision between the orogen and the
Bohemian Massif resulted in left lateral displacement along
this zone followed by opening of the Vienna Basin “pull

apart”  depocentres  (Royden  1985;  Tomek  &  Hall  1993;
Lankreijer et al. 1995; Fodor 1995; Kováč et al. 2004). Ini-
tial rifting in the Pannonian back arc basin opened new ma-
rine connections in this time. The sea transgression from the
Mediterranean  advanced  via  the  Trans-Tethydian  Trench
Corridor  (Rögl  1998;  Pavelič  2001).  The  broad  Karpatian
sea flooding extended beside the back arc basin also into the
Vienna Basin and the Western Carpathian Foredeep domain
(Fig. 2B). The continental areas were represented beside the
European platform by uplifted parts of internal zones of the
Eastern Alps and Western Carpathians; the accretion wedge
of the Outer Carpathian Flysch Belt showed a minimal uplift
in its western part only.

The Middle Miocene geodynamic development influenced

factors such as the end of subduction in front of the Western
Carpathian  orogen,  its  soft  docking  on  the  slopes  of  the
European platform (Konečný et al. 2002) and back arc syn-
rift  subsidence  in  the  Pannonian  Basin  domain  (Horváth  et
al.  1988).  Voluminous  acid  and  calc-alkaline  volcanism  is

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GEOLOGICA CARPATHICA, 2011, 62, 3, 251—266

observed during this time. The Central Paratethys Sea can be
characterized  in  this  time  as  an  epicontinental  sea  with  a
number of archipelagoes and a lot of separate basins. The sea
flooding extended in front of the orogen (foredeep basin), as
in  the  Pannonian  back  arc  basin  area.  In  the  study  area
(southern  part  of  the  Carpathian  Foredeep  and  Vienna  Ba-
sin),  the  late  Early  Badenian  transgression  was  completely
controlled by tectonics (Lankreijer et al. 1995; Kováč et al.
2001,  2004)  and  associated  with  basin  subsidence  and  con-
tinuous mountain chain uplift. During the Middle Badenian a
part of the Western Carpathian basins were isolated with the
resulting salinity crisis (Kováč et al. 1998b); in the southern
part  of  the  Carpathian  Foredeep  the  sedimentation  ended.
The Vienna Basin still subsided and was connected towards
the  Pannonian  domain  by  straits  in  the  uplifting  Malé  Kar-
paty and Leitha Mountains. The changes in relief led to de-
velopment  of  a  drainage  system,  the  paleo-Danube  river
delta  entered  the  Vienna  Basin  and  voluminous  deltaic  se-
quences started to be deposited (Kováč et al. 2004; Lambert
et al. 2008). The Late Badenian flooding covered the whole
northen  part  of  the  Pannonian  back  arc  basin  system
(Fig. 2C). In the Vienna Basin the transgression was accelerat-
ed by basin subsidence and sea-level rise (Kováč et al. 2001,
2006).  The  basins  were  filled  by  clastic  material  transported
by rivers, entering the basin from elevated mountain ranges in
the basin surroundings. In front of the orogene, the Carpathian
Foredeep started to disintegrate and depocentres moved from
West towards East (Meulenkamp et al. 1996). During the Sar-
matian, closing of the Central Paratethys Sea connections with
the Mediterranean Sea led to isolation and salinity decrease in
all the Western Carpathian basins. The sea started to be shal-
lower, but its extent did not change significantly.

The Late Miocene geodynamic development represents the

final stage of the Pannonian back arc basin evolution, with re-
lated  thermal  subsidence  (Horváth  1988;  Kováč  et  al.  1993;
Lankreijer et al. 1995; Konečný et al. 2002). The Vienna Ba-
sin represented a partly isolated bay at its northwestern bound-
ary during this time (Kováč et al. 1998a; Kvaček et al. 2006).
Uplift of the Western Carpathian mountain chain was accom-
panied  by  the  next  development  of  the  river  net,  resembling
the  Pliocene  paleogeography  (Fig. 2D).  The  Central  Para-
tethys  brackish  sedimentary  environment  –  represented  by
Lake Pannon in the study area (Magyar et al. 1999) gradually
changed  into  a  freshwater  lake  environment,  particularly  in
the back arc basin domain. During the Pannonian, the Vienna
Basin  was  filled  by  a  huge  amount  of  deltaic  sediments
(Kováč et al. 1998, 2004, 2006; Harzhauser et al. 2004). The
shallow-water  fluvial  to  lacustrine  environment  changed  to
swamps and alluvial plains with ephemeral lakes representing
the greater part of its territory until the end of the Late Mio-
cene. The Pannonian and Pontian mountain ranges gained fea-
tures similar to their present form.

Material and methods

In this study, 44 outcrops and boreholes (Carpathian Fore-

deep – Eggenburgian—Early Badenian (23 localities), Czech
and Slovak parts of Vienna Basin – Karpatian—Pannonian

(21  localities))  were  analysed.  Due  to  the  absence  of  index
fossils,  the  studied  Lower  Miocene  sediments  are  undistin-
guishably  (Upper  Eggenburgian—Ottnangian).  The  analysed
samples come from the localities in the southern part of the
Carpathian Foredeep – boreholes Šafov 12, Šafov 13, Čej-
kovice, Únanov, Miroslav, Trboušany, Nosislav 3, Židenice.
Carpathian Foredeep marine and brackish sediments, Karpa-
tian in age, were evaluated from several stratotype localities
Slup,  Hevlín,  Dolní  Dunajovice,  Medlov,  boreholes  No-
sislav 3,  Ždánice 67,  68  and  from  the  Vienna  Basin  bore-
holes  Zohor 1  and  Gbely 139  were  analysed  (Doláková  &
Slamková 2003). Palynological data, Early Badenian in age,
come from the Carpathian Foredeep marine sediments from
the  localities  Židlochovice,  Lysice,  boreholes  Moravské
Knínice,  Sivice,  Chrlice,  Opatovice  and  Otmarov.  Pollen
data,  Late  Badenian  in  age,  come  from  outcrop  Devínska
Nová Ves and borehole Lozorno 1 in the Vienna Basin. Stud-
ied  sediments,  Late  Miocene  in  age,  with  well-determined
plant macrofossils (Knobloch 1968, 1985) come from the Poš-
torná, Dubňany, Moravská Nová Ves outcrops; clay pit Gbely,
boreholes  Suchohrad 32,  Suchohrad 38,  Jakubov 54  and  six
shallow Pohansko boreholes near Břeclav city.

In the chemical treatment 20—30 g of dry sediment was

used. The samples were treated with cold HCl (35%) and HF
(70%), removing carbonates and silica. Separation of the pa-
lynomorphs from the rest of the residue was carried out using
ZnCl

(density = 2 g/cm

). Sieving was done using 10 µm ny-

lon  sieve.  The  palynological  residue,  mixed  with  glycerine,
was  prepared  on  slides.  A  transmitted  light  microscope  with
250, 400, 630, 1000 (oil immersion) magnifications and SEM
microscope  was  used  for  pollen  counting  and  identification.
Original micrographs are housed at the Institute of Geological
Sciences MU in Brno. The pollen diagrams have been created
using POLPAL 4 software (Walanus & Nalepka 1999).

To have a better idea about the vegetation composition the

differentiated vegetation groups (zonal, azonal, extrazonal)
were used sensu Kovar Eder et al. (2008a,b) and Kvaček et
al. (2006). A semiquantitave evaluation of climate evolution
has been done based on the proportion of paleotropical (ther-
mophilous) and arctotertiary elements (sensu Mai 1981,
1991) in terms of mesophytic plants. We devided the floris-
tic elements of zonal vegetation into two groups.

In the thermophilous-mesophytic group we included Engel-

hardia,  Sapotaceae,  Palmae,  evergreen  Fagaceae  (including
morphospecies  Quercoidites  microhenrici  and  Quercoidites
henrici),  Trigonobalanopsis,  Symplocos,  Cornaceaepollis
satzveyensis, Tricolpopollenites liblarensis, Araliaceae, Ruta-
ceae. Mostly broad-leaved deciduous elements of warm-tem-
perate  mixed  mesophytic  forests  such  as  Quercus,  Celtis,
Carya, Tilia,  Zelkova, Ostrya,  Carpinus,  Betula,  Juglans  are
included into the group of arctotertiary—mesophytic elements.
Extrazonal  mountain  vegetation  is  represented  by:  Cedrus,
Tsuga,  Picea,  Cathaya.  Azonal  vegetation  is  influenced  by
edaphic  factors  and  in  our  study  it  is  represented  by  riparian
forests  with  Alnus,  Salix,  Ulmus,  swamps  with  Taxodiaceae,
Myricaceae, Nyssaceae and aquatic plant communities.

All the studied material is housed at the Institute of Geo-

logical Sciences MU in Brno and the Faculty of Sciences of
Comenius University in Bratislava.

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Results and discussion

Early Miocene

Eggenburgian—Ottnangian—Karpatian (Late Aquitanian—

Late Burdigalian)

During the Early Miocene thermophilous taxa Engelhardia,

Platycarya,  Sapotaceae,  Palmae  and  ferns  Lygodium,  Pteri-
daceae,  ?Davalliaceae,  Schizaeaceae—Cyatheaceae  were  fre-
quent.  Evergreen  Fagaceae  were  represented  by  the
Trigonobalanopsis  type,  morphotaxa  Tricolporopollenites
microhenrici and Tricolpopollenites liblarensis, Tricolporopol-
lenites  henrici.  Also  Symplocos,  Reevesia,  Parthenocissus,
Araliaceae,  Rutaceae  and  morphotaxa  Cornaceaepollis
satzveyensis,  Tricolporopollenites  pseudocingulum  were  com-
mon  in  pollen  spectra.  Arctotertiary  elements  Carya,  Juglans,
Quercus, Betula, Liquidambar were less frequent (Fig. 3).

The vegetation of the salt marshes and also insolated places

(Chenopodiaceae up to 37 %, Ilex, Tamarix, Ericaceae, Poaceae
less Ephedra, Asteraceae and Buxaceae) was typical. Due to
the salinity oscillations as well as occasional higher evapora-
tion sensu Hladilová (1988), the coasts of individual sea gulfs
and lagoons could be repeatedly salinized and overgrown by
the halophilous flora (Doláková et al. 1999). Pollen grains of
the formal genus Monocirculipollis assigned to the family
Caryophyllaceae (sensu Doláková 2004), were typical for this
time span being absent in younger ones (Fig. 6). Salt marsh
vegetation was sometimes replaced by swamp plants.

Taxodiaceae, Myricaceae, Cyrillaceae, Gleicheniaceae,

Decodon,  Lygodium,  Selaginella.  Even  the  aquatic  flora  ap-
peared  –  Sparganium,  Potamogeton,  Onagraceae,  Nelumbo,
Cyperaceae  (Figs. 3,  5,  7).  The  genus  Platanus  was  the  com-
mon member of the pollen spectra since this time span. A per-
manently  low  amount  of  intrazonal  elements  (Taxodiaceae,
Myricaceae and ferns) without strong oscilation occurred in the
palynospectra. Regularly higher ratios of Fagaceae, Carya and
also  heliophilous  taxa  were  observed.  Pinaceae  (up  to  40 %)
and extrazonal vegetation, including abundant Cathaya and less
frequent  Cedrus, Picea,  Abies,  occurred  frequently  (Fig. 3).  A
high  proportion  of  Ulmaceae,  Myrica,  Alnus  was  observed  in
sediments of the Eggenburgian—Ottnangian. Pollen spectra con-
tain a larger amount of spores of thermophilous ferns as Lygodi-
um  (up  to  5 %),  Pteridaceae,  Gleicheniaceae  together  with
Selaginella and bryophyte Riccia (Fig. 3). These findings are in
a  good  conformity  with  macrofloristic  results  of  Knobloch
(1982) who described a unique oryctocoenosis from the rhyolite
tuffites  at  Znojmo  and  Přímětice.  He  considered  them  as  the
shrubby – arboreal heliophilous vegetation with mostly ever-
green  fine  dentate  or  spiny  leave  (sclerophyllous)  similar  to
Mediterranean “macchias”. Swamp vegetation with Glyptostro-
bus,  Myrica,  aquatic  flora  with  Salvinia,  Potamogeton,  Nym-
phaea and coastal reed with Typha, Decodon, Sparganium were
identified  based  on  macrofloristic  remains  too  (Knobloch
1982). The accumulations of Limnocarpus fruits growing in the
brackish water were described by Knobloch (1984). The pollen
often  found  in  clumps  (Myricaceae,  Chenopodiaceae,  Caryo-
phyllaceae,  Oleaceae,  Onagraceae,  Platanus)  support  the  low
water dynamics and a short transport (Figs. 5, 6, 7).

During the Karpatian the thermophilous elements like Ruta-

ceae,  Symplocos  and  Platanus  occurred  less  regularly.  Tem-
perate  taxa  are  represented  generally  in  low  frequencies,  but
the  amount  and  diversity  of  the  temperate  mesophytic  ele-
ments  slightly  increased  (Fig. 4).  Mountain  vegetation  with
Tsuga and Abies was common (Fig. 4). Subtropical humid cli-
mate  was  also  supported  by  the  macrofloristic  remains  de-
scribed  from  the  stratotype  localities  Slup  and  Dolní
Dunajovice.  Leaves  of  the  family  Lauraceae  predominated
with  small  proportion  of  deciduous  trees  in  this  association
(Knobloch 1967, 1982; Kvaček 2003). The azonal vegetation
is  represented  by  swamp  and  riparian  forests  dominated  by
Glyptostrobus and Myrica.

Marshy-palm forest with Calamus, Poaceae, Lygodium,

Sparganium,  Potamogeton  and  riparian  forest  with  Alnus,
Ulmus,  Myricaceae,  Lythraceae  or  Selaginella  are  frequent
(Figs. 4, 8). The associations with Taxodiaceae, Craigia and
Pteridaceae (up to 10 %) and Polypodiaceae document a well
developed  swamp  environment  (Figs. 4, 8).  During  the
present time the genus Craigia occurs in broad-leaved ever-
green  and  deciduous  mixed  forests  and  seasonally  wet  for-
ests. However, according to Kvaček et al. (2002), ecological
tolerances of its fossil representatives may have been greater
during the Tertiary. This tree surely tolerated swampy condi-
tions and entered even coal-forming forests in wetland habi-
tats  namely  swamp  forests  dominated  by  the  Taxodiaceae
and  many  other  swampy  and  riparian  woody  plants  as  well
aquatic herbs. Konzalová (1976) described a very similar ho-
rizon  with  Intratriporopollenites  insculptus  Mai  (Craigia)
from  the  coal  seam  formation  and  Cypris  claystones  of  the
North Bohemian basins.

During this time interval the plant assemblages with high-

er portion of arctotertiary elements were described by several
authors from the Silesian part of the Carpathian Foredeep in
the Polish Lowland (Oszast & Stuchlik 1977; Stuchlik 1980;
Sadowska  1989;  Ważyńska  et  al.  1998).  The  decrease  of
thermophilous elements during the Ottnangian—Early Karpa-
tian  has  been  recognized  and  defined  as  the  microfloristic
Zone  MF-3  by  Planderová  (1990)  and  Planderová  et  al.
(1993a,b).  Such  an  event  has  never  been  found  in  the  Car-
pathian  Foredeep.  This  fact  is  probably  related  to  different
paleogeography. The most similar pollenspectra of Karpatian
age  was  published  by  Nagy  (1999)  from  the  Mecsek  Mts.
Environmental  interpretation  of  data  from  the  Carpathian
Foredeep is similar to the conditions in the Korneuburg Basin
(Hofmann  et  al.  2002),  except  for  absence  of  Avicenia  and
the  lower  portion  of  the  Palmae  in  the  studied  area.  Early
Miocene was the warmest period of the Miocene in the Pan-
nonian Basin and the sporomorphs indicate a warm subtropi-
cal climate (Nagy 2005).

Middle Miocene

Badenian—Sarmatian (Langhian—Serravallian)

Swamp elements (Taxodiaceae) have more regular occur-

rence without oscillations in comparison with the Karpatian.
Olea type pollen was less frequent in comparison with Low-
er Miocene pollen spectra. In the Badenian pollen spectra the

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higher differentiation of the Fagaceae in thermophilous ever-
green  (morphotypes  Tricolporopollenites  henrici  and  Tri-
colporopollenites  microhenrici)  and  deciduous  oaks,  some
thermophilous taxa Gothanipollenites gothani (Loranthaceae)
or  Tricolporopollenites  indeterminatus  (Hammamelidaceae)
occurred (Fig. 9). Herbs such as Caryophyllaceae (Minutipol-
lis granulatus Krutzsch) were common. Early Badenian Lau-
raceae  and  Betulaceae  leaves  from  the  Carpathian  Foredeep
have been found at the Smolín locality (Sitár et al. 1978). Dur-
ing the Late Badenian the following were frequently present:
Pinaceae (Pinus, Picea, Abies, Tsuga) and deciduous elements
with Quercus, Alnus, Ulmus, Carya. Subtropical taxa are com-
monly  represented  by  Magnolia,  Platycarya,  Engelhardia,
Myrica,  Trigonobalanopsis  and  Distylium.  Paleoecological
conditions favoured development of swamp forests with Taxo-
diaceae, Nyssa  and  Myrica  and  riparian  forest  elements  with
Alnus,  Ulmus  and  Pterocarya.  Drier  areas  were  overgrown
with mixed mesophytic forest represented by Pinus, Juglans,
Carya, Sciadopitys and the extrazonal vegetation type is docu-
mented by the presence of Picea, Tsuga and deciduous oaks.
Herbs  are  represented  mostly  by  Poaceae.  During  the  Early
Sarmatian time interval the azonal vegetation was well devel-
oped  in  swamps  with  Taxodiaceae,  Myrica,  Nyssa  and  salty
marshes  with  Poaceae  and  halophytes  (Chenopodiaceae).
Riparian forests with Alnus, Salix and Ulmus were also com-
mon.  The  extrazonal  vegetation  portion  increased  mainly  in
the mountain vegetation elements. The decrease to disappear-
ance  of  the  Taxodiaceae,  Nyssaceae,  Myricaceae  and  halo-
phytes  suggests  a  large  reduction  of  the  swamp  biotops.
Riparian  forests  with  Ulmus,  Salix,  Alnus  and  Poaceae  were
still present. Gradual decrease of thermophilous taxa indicates
moderate cooling. Syabryaj & Vodoryan (1975) described simi-
lar,  well  diversified  pollen  spectra  from  the  NE  Carpathian
territory in Čop—Munkacevo. Ivanov (1995) and Ivanov et al.
(2002)  described  presence  of  Symplocos  in  Badenian  pollen
spectra from NW Bulgaria. In our studied material we noticed
Symplocos presence only from Early Miocene localities, prob-
ably due to temperature gradient.

There was a warm subtropical climate. Early Badenian

transgression and uplift of the Alpine and Carpathian Moun-
tains produced favourable local climate for vegetation
change (Nagy 2005).

Late Miocene

Pannonian—Pontian (Tortonian—Messinian)

In the pollen spectra from the Early Pannonian sensu

Harzhauser et al. (2004) or Early Tortonian sensu Harzhauser &
Piller  (2007),  mostly  broad-leaved  deciduous  elements  domi-
nate,  with  some  thermophilous  elements  admixture  of  Engel-
hardia,  Ilex,  Castanopsis  and  Castanea,  suggesting  a  warm
temperate mixed mesophytic forest with low representation of
evergreen  elements.  The  proportion  of  NAP  –  non  arboreal
pollen  –  Ericaceae  and  Chenopodiaceae  is  higher  (10  and
14 %  respectively)  suggesting  local  marshes  and  open  herba-
ceous plant communities within the forests. Mountain conifers,
such as Picea,  Tsuga, Abies, Cedrus are common accessories.
Lowland vegetation was comprised of the azonal Alnus, Pinus,

Ulmus  mixed  and  broad-leaved  riparian  forest  with  common
deciduous oaks, and swamp taxa Taxodiaceae, Nyssa, Myrica.
Sporadic  occurrences  of  dinoflagellates  and  green  algae  Tas-
manaceae indicate a slightly higher salinity, Botryococcus can
thrive  in  both  brackish  or  freshwater  environments,  whereas
green  algae  Pediastrum,  Mougeotia,  aquatic  ferns  Azolla,  and
aquatic  and  coastal  plants  (Nelumbo,  Nymphaea,  Myriophyl-
lum, Sparganium, Potamogeton etc.) represent a freshwater en-
vironment (Doláková & Kováčová 2008). In the pollen spectra,
from the middle Pannonian (sensu Harzhauser et al. 2004), co-
niferous  woody  plants  of  mountain  vegetation  (Picea,  Abies,
Tsuga, Cedrus, Pinus) and deciduous oaks were abundant. An-
giosperm  trees  and  shrubs  with  Alnus,  Betula,  Liquidambar,
Myrica, Nyssa and Salix indicate a well developed riparian for-
est. The subdominance of herb species is good evidence of the
local open woodland environment.

The facies mutually changing in time and space in individ-

ual pollen spectra are created by azonal types of vegetation
(marshes,  riparian,  coastal  and  aquatic)  or  by  high  amounts
of  herbaceous  plants  Artemisia,  Plantago,  Polygonum,
Asteraceae, Lamiaceae, Daucaceae, Caryophyllaceae,  which
indicate existence of local open areas.

In the Slovak part of the Danube Basin Planderová (1972,

1990) described reduced marshes, isolated lakes surrounded
by  steppe  meadows  (dominance  of  Artemisia)  with  rare
woody  plants.  In  comparison  with  Hungary  she  considered
the climate cooler and drier (Nagy 1985; Nagy & Planderová
1985;  Planderová  1990).  Hoffmann  &  Zetter  (2005)  deter-
mined a pollen assemblage rich in herbs from the Late Pan-
nonian in the Styrian Basin.

The extensive Pannonian and Pontian sea and the protec-

tive mountain range provided a very equable, warm temper-
ate climate, where even the summer season was not too dry
(Nagy 2005).

Conclusions

Development of Miocene vegetation patterns in the area of

the  northwestern  Central  Paratethys  was  derived  (above  all)
from  palynological  analysis.  The  case  study  area  covers  the
slopes  of  the  tectonically  quiet  European  platform  and  the
foreland of the tectonically active Carpathian mountain chain
(Carpathian  Foredeep  and  Vienna  Basin).  Interpretation  of
pollen  spectra  reflects  both,  the  landscape  evolution  in  two
areas with different geodynamics, and the climatic changes in
the  Central  Paratethys  domain  during  the  studied  time  inter-
vals. Based on pollen data, the Early to Middle Miocene vege-
tation  document  a  subtropical  climate  with  very  mild
(negligible)  cooling  during  this  period.  This  is  indicated  by
common  occurrence  of  thermophilous  taxa:  Sapotaceae,
Palmae,  Engelhardia,  Platycarya  and  Tricolporopollenites
henrici, Lygodium and Pteridaceae. Reevesia, Cornus-Mastixia,
Symplocos,  Parthenocissus,  Tricolporopollenites  pseudocin-
gulum,  Rutaceae  and  Araliaceae.  The  proportion  of  the  tem-
perate  elements  such  us  Carya,  Pterocarya,  Juglans,  Celtis,
Fagus  is  noticeably  lower.  The  lower  portion  of  extrazonal
(mountain)  vegetation  and  well  developed  riparian  forests
with Alnus, Craigia, Pteridaceae, Polypodiaceae, Lythraceae,

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GEOLOGICA CARPATHICA, 2011, 62, 3, 251—266

Cyperaceae, Sparganium, Potamogeton, Nelumbo and swamps
with  Taxodiaceae,  Myricaceae  alternated  with  salt  marshes
represented by Chenopodiaceae up to 37 %, Poaceae, Caryo-
phyllaceae,  Asteraceae,  Ericaeae,  Lythraceae,  which  docu-
ment  a  moderate  relief  of  landscape  during  the  whole  Early
Miocene. The frequent pollen clumps support a theory of the
low water dynamics and a short transport distance. The alter-
ation in palynomorphs caused by the crystallization of pyrite
in anoxic conditions was observed.

The Middle Miocene landscape evolution, conditioned by

the uplift of the Carpathian mountain chain and subsidence of
adjacent lowlands, led to commencement of the altitudinal zo-
nation. This process is documented by changes in paleovege-
tation cover. In spite of this presence of zonal vegetation with
evergreen  broadleaved  forests  supplemented  by  azonal  vege-
tation (riparian forests, swamps) is typical of the Early Bade-
nian. Only several thermophilous plants, such as Engelhardia
and Platycarya, which were frequent in all of the Early Mio-
cene  associations,  decreased  in  the  Early  Badenian  pol-
lenspectra.  From  the  Late  Badenian  a  higher  proportion  of
extrazonal (mountain) vegetation were present in pollen spec-
tra (Picea, Abies, Tsuga, Cedrus). The terrestrial and aquatic
ecosystems confirm a subtropical climate with visible changes
at the boundary between the Early and Late Badenian. An in-
creased  proportion  of  the  arctotertiary  taxa  during  the  Late
Badenian is documented in pollenspectra by Quercus, Ulmus
and Carya, whereas Platycarya, Engelhardia, Myrica, Distylium
and thermophilous Fagaceae are less frequent. Herbs are rep-
resented mainly by the halophytes (Chenopodiaceae).

Vegetation during the Early Sarmatian time interval was

formed  by  swamp  elements  with  Taxodiaceae,  Myricaceae,
Nyssaceae.  High  elevation  species  of  woody  plants  Tsuga,
Picea, Cedrus, Abies are indicative of mountainous relief re-
sulting from volcanic activity. During the Late Sarmatian the
proportion  of  swamp  elements  decreased  and  was  replaced
mostly by riparian forests.

The Late Miocene paleogeographical changes and general

climatic  oscillations  in  the  northwestern  Central  Paratethys
realm  reflected  the  decrease  especially  of  the  thermophilous
taxa Engelhardia, Castanea, evergreen Fagaceae Quercoidites
microhenrici,  and  to  a  lesser  extent  of  Quercoidites  henrici,
Trigonobalanopsis, Symplocos, Cornaceaepollis satzveyensis,
Tricolpopollenites  liblarensis.  An  apart  from  the  mountain
vegetation the amount of herbaceous plants in the pollen spec-
tra increased during this time span. The varying height’s of the
moutain  relief    of  the  uplifted  mountain  chains  (altitudinal
zonality)  created  ideal  conditions  for  extrazonal  vegetation
(Cedrus, Tsuga, Picea) and dominance of mixed mesophytic
forests  with  Quercus,  Celtis,  Carya,  Tilia,  Zelkova,  Ostrya,
Liquidambar,  Carpinus,  Betula,  Juglans  and  with  regular
presence of evergreen taxa.

The swamp, riparian, often hydrophilous (Azolla, Nym-

phaea,  Potamogeton)  and  halophytic  (Chenopodiaceae)
plants  represent  coastal  swamps,  local  lagoons,  and  marsh-
lands. The higher percentage of the herbs (Artemisia, Aster-
aceae, Lamiaceae, Polygonum, Daucaceae, Caryophyllaceae,
Plantago) and shrubs in the comparison with older time in-
tervals, shows that local open woodland  – open grassland
started to develop during the Pannonian.

The gradual retreat of areas flooded by the sea, as well as

following  retreat  of  the  lake  and  swamp  environment  was
confirmed by the decrease of azonal vegetation towards the
end  of  this  period.  The  reconstructed  vegetation  cover  sug-
gest  a  subtropical  climate  with  gradual  transition  to  warm
temperate climatic conditions.

Acknowledgments: The authors wish to express their grati-
tude  to  reviewers  L.  Hably,  Z.  Kvaček  and  D.  Ivanov  for
useful and constructive comments. This work was supported
by the Slovak Research and Development Agency under the
contracts  APVV-LPP  0120-06,  APVV-0280-07,  ESF-EC-
0006-07,  ESF-EC-0009-07  and  the  Slovak  Grant  Agency
VEGA (Projects No. 2/0060/09, 1/0483/10) and Czech Grant
Agency under the contract No. 205/09/0103.

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