GeolCarp_Vol57_No4_295

GEOLOGICA CARPATHICA, AUGUST 2006, 57, 4, 295—310

www.geologicacarpathica.sk

Miocene evolution of landscape and vegetation in the Central

Paratethys

ZLATKO KVAČEK

, MICHAL KOVÁČ

, JOHANNA KOVAR-EDER

, NELA DOLÁKOVÁ

, HENRIETTE

JECHOREK

, VALENTIN PARASHIV

, MARIANNA KOVÁČOVÁ

and UBOMÍR SLIVA

Charles University, Faculty of Science, Albertov 6, CZ-128 43 Praha 2, Czech Republic; kvacek@natur.cuni.cz

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

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

Staatliches Museum für Naturkunde Stuttgart, Rosenstein 1, D-07191 Stuttgart, Germany; eder.smns@naturkundemuseum-bw.de

Department of Geology and Paleontology, Faculty of Sciences, Masaryk University, Kotlářská 2, CZ-611 37 Brno, Czech Republic;

nela@sci.muni.cz

Carl-von-Ossietzky-Straße 5, D-02826 Görlitz, Germany; hjechorekgr@surfeu.de

Faculté de Géologie, Université de Bucharest, Str. N. Balcescu, Bucharest, Romania; paleovaly@yahoo.com

(Manuscript received June 27, 2006; accepted in revised form December 8, 2005)

Abstract: The digital elevation model (DEM) helps to express Neogene landscapes and vegetation on palinspastic maps
with reconstructed orography. To reconstruct ancient vegetation cover, basic zonal vegetation formations and their char-
acteristics have been defined based on diversity and proportions of zonal woody evergreen, deciduous, sclerophyllous
and legume-type elements, besides intrazonal (azonal, e.g. coal-forming, aquatic and riparian) and extrazonal (montane
conifer-rich) vegetation. Three time intervals have been analysed – Karpatian to Early Badenian, Late Badenian to earliest
Sarmatian and Early to Middle Pannonian. After evaluating respective local sites of leaf, fruit/seed and spore/pollen
assemblages, paleogeobotanical maps have been constructed for the area of the Central Paratethys and its periphery.

Key words: Miocene, Central Paratethys, paleogeography, palinspastic maps, digital elevation models, vegetation mapping.

Introduction

It is a common task of geobotany today to express interpret-
ed vegetation over larger areas on maps, because the extent
of various types of plant communities is an important fac-
tor, for example, in tracing human influence or migrations
of terrestrial animals. However, several aspects of such stud-
ies are different considering Neogene vegetation (Kovar-
Eder et al. submitted). The paleogeographic configuration
of land and sea was different from the present situation. The
regional relief changed in connection with orogeny pro-
cesses. The floristic spectra included elements mostly ex-
tinct or no longer living in Europe. The time slices for the
respective maps are many million years distant from the Re-
cent. Global climate, atmospheric circulations and the
world ocean varied depending on the time interval studied.
To overcome these problems a team of specialists is needed.
The paleogeographic background with an approximate de-
marcation of sea, basins and approximate relief is the first
premise to attempt such a paleogeobotanical mapping.

Complex and well determined spectra of plant elements

from the reference sites with both megafossil and spore/pol-
len records are most relevant. The megafossil record usually
reflects the situation near the site and is differentiated ac-
cording to the lithofacies. It also reflects the presence of
plants producing poorly preservable pollen (e.g. Lauraceae)
and indicates more strongly floristic changes than the
spore/pollen spectra. Leaves may convey information on
the vegetation physiognomy. Fruits and seeds are better in-
dicators of systematic affinities. Pollen and spores may un-

dergo long distance transport by wind and their spectra thus
include information on the composition and changes of up-
land vegetation, which usually consists of mountain coni-
fer-rich forests. It is problematic to transfer frequencies of
any kind of plant organs in the fossil spectra into true abun-
dances of plants in a community or landscape, because the
fossil record is biased either by overproduction of fossil or-
gans (e.g. pollen, diaspores) and taphonomic processes (e.g.
deciduous vs. evergreen foliage, more rapid decay of deli-
cate leaves). In our analyses, we relied mostly on qualitative
proportions of elements, that is the floral diversity, within
the given assemblage.

A synthetic view over a large area that includes several

countries should be obtained based both on own experi-
ences including authentic knowledge of plant fossil sites
and the review of the published data. In the latter cases, it
is often a difficult task to critically re-evaluate older taxo-
nomical interpretations, both in the megafossil and pollen
spectra. Particularly the interpretations of various pollen
types within the natural system look very different today
from the traditional morphological or semi-natural sys-
tems used previously (or even at present). Particularly by
efforts of large-scale comparisons with living plants
(Stuchlik 1994) and electron scanning microscopy
(Walther & Zetter 1993; Ferguson et al. 1998; Zetter
1998; Liu et al. 2001), surprising solutions for several
sporomorphs have been suggested and important natural
affinities have been recognized (e.g. Mastixia, i.e. Cor-
naceaepollis satzveynsis, Trigonobalanopsis, i.e. Casta-
neoipollenites pusillus, etc.). Studies of pollen in situ and

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KVAČEK et al.

co-occurrence with megafossils are also important. Thus it
has recently become obvious that the tilioid pollen does not
belong in most cases to Tilia, a deciduous zonal element, but
to Craigia & Dombeyopsis lobata or Banisteriaecarpum &
Byttneriophyllum tiliaefolium (alias “Alangium”) plants both
intrazonal, frequent constituents of the Glyptostrobus swamp
forests (e.g. Kvaček et al. 2002).

This paper introduces a new methodology of the Neo-

gene paleogeobotanical mapping presenting it on three
time slices of the Neogene in the Central Paratethys.

Methodology

Digital Elevation Models

(DEMs) of the Central Parat-

ethys (Fig. 1) in individual time intervals of the Miocene,
which help us to express landscapes with expected orogra-
phy, were constructed on the basis of present knowledge
of geodynamic evolution of the Alpine-Carpathian-Pan-
nonian region (Csontos et al. 1992; Kováč et al. 1993,
1997, 2001, 2002; Meulenkamp et al. 1996; Baráth et al.
1997; Plašienka et al. 1997; Plašienka & Kováč 1999;
Bezák et al. 2002; Konečný et al. 2002; Soták & Kováč
2002; Bielik et al. 2004), existing palinspastic maps
(Kováč et al. 1989, 1998, 2003; Magyar et al. 1999;
Kováč 2000; Popov et al. 2004), as well as burial and up-
lift history of sedimentary or other rock complexes (Hor-
váth et al. 1988; Dunkl 1992; Kováč et al. 1994; Hurai et
al. 1995; Dunkl & Demény 1997; Danišík et al. 2004,
etc.).

Models of the Central Paratethys vegetation

(Figs. 2—4)

use a simplified system of vegetation units (formations),
which are usually sufficient to get tentative pictures of pa-
leovegetation in space. Our attention has been paid to dis-
tinguish zonal, intrazonal (azonal) and extrazonal
formations on the basis of autecology and leaf physiogno-
my of elements, whose grouping has been attempted in
this respect (Kovar-Eder & Kvaček 2003; Kovar-Eder et al.
submitted). The characteristics of the elements have been
mostly derived from autecologies of their nearest living
relatives or analogues. Reference fossil localities/plant as-
semblages often included taxa of different vegetation for-
mations, of which zonal elements are relevant for the maps
of the reconstructed fossil vegetation. However, it is ap-
parent from the sedimentary settings that assemblages
dominated by intrazonal elements prevailed in the record
of megafossils, mainly from the basin deposits. Extrazonal
conifer-rich mountain vegetation was represented in pol-
len spectra, exceptionally in megafossil records from the
intra-montane basins. Not only altitude, but also the direc-
tion of exposure of mountain slopes and substrate may
have influenced the composition of the conifer stands.
Volcanic settings are the best environments to bring infor-
mation on zonal vegetation of mesic habitats. Percentages
of zonal herbs as well as Non Arboreal Pollen (NAP) as a
whole in pollen spectra may refer to close canopy forests
versus open woodland to steppe vegetation. Even intra-
zonal elements can bring information on the character of
climate (e.g. the presence of palms), although the intrazon-

al assemblages usually bear “cool” aspects due to higher
proportion of deciduous arboreal elements.

For the purpose of the presented paleovegetation maps

several formations have been distinguished and character-
ized mostly based on the proportion of broad-leaved de-
ciduous, broad-leaved evergreen, sclerophyllous and
legume-type components of zonal woody angiosperms
(Kovar-Eder et al. submitted).

Zonal formations

1. (Warm-) temperate Broad-leaved Deciduous Forest

with very low proportion of evergreen woody elements
(vegetation unit 1) includes more than 80 % of zonal de-
ciduous woody elements of angiosperms, such as Parro-
tia, Zelkova, Ostrya, Acer angustilobum etc.

2. Warm-temperate Mixed-Mesophytic Forest (vegetation

unit 2) includes less than 80 % deciduous woody elements of
zonal angiosperms, less than 30 % evergreen broad-leaved
woody taxa of zonal angiosperms and less than 20 % sclero-
phyllous and legume type elements, regular admixture of
Tetraclinis salicornioides and other thermophilous elements,
less than 30 % of zonal herbs of zonal angiosperms.

3. Subtropical Broad-leaved Evergreen Forests includ-

ing the “Younger Mastixioid Floras” sensu Mai (1964)
(vegetation unit 3) includes equal or more than 30 %
broad-leaved evergreen and thermophilous elements, rep-
resented mainly by Lauraceae, Theaceae, Mastixiaceae,
Symplocaceae, Sapotaceae, Engelhardia, and evergreen
Fagaceae (represented in pollen spectra by morpho-spe-
cies Castaneoideoipollenites pusillus, Quercoidites henri-
ci, Quercoidites microhenrici – types) and less than 25 %
of zonal herbs among zonal angiosperms.

4. Subtropical Sub-humid Sclerophyllous Forest (vegeta-

tion unit 4) includes more than 20 % sclerophyllous taxa
(Quercus mediterranea, Quercus drymeja) and legume-type
microphyllous woody elements of zonal angiosperms.

Intrazonal formations

5. Swamp forest and coal-forming mire (not expressed

by patterns on maps, vegetation unit 7) is dominated by
coal-forming woody and herbaceous elements (e.g. Glyp-
tostrobus and other Taxodiaceae, Byttneriophyllum, Nys-
sa, Myrica, Calamus, Spirematospermum, etc.).

6. Marsh and aquatic vegetation (not expressed by pat-

terns on maps, vegetation unit 8) is dominated by aquatic
herbs and helophytes (Cyperaceae, Typha, Potamogeton,
Stratiotes, etc.).

7. Deciduous riparian forest (not expressed by patterns

on maps, vegetation unit 9) is dominated by woody ele-
ments of moist substrates (Taxodium, Alnus, Salix, Popu-
lus, Fraxinus, Acer tricuspidatum, etc.).

Extrazonal formations

8. Mountain conifer-rich forest is dominated (mostly in

pollen records, vegetation unit 10) by Pinaceae (including
Cedrus, Tsuga, Picea, Cathaya, etc.).

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KVAČEK et al.

vegetation formations between and around the reference
sites. The intrazonal vegetation was omitted because of its
limited extent compared with the scale of the maps. The
approximate position of extrazonal conifer belts has been
placed according to the nearby palynodata and orography.

Early Miocene – model of the Karpatian landscape

and vegetation of the Central Paratethys

During Early Miocene, the Central Paratethys was situ-

ated at least 200—300 km to the south of its present posi-
tion. The sea, located along the southern slopes of the
European platform, covered the southwestward dipping
subduction zone in front of the developing Alpine-Car-
pathian orogene. The accretion wedge, build up by exter-
nal Alpine and Carpathian Flysch Belt units, was marked
in that time only with islands, forming an archipelago
along the platform margin. The uplifted islands, composed
of units of the Outer Carpathian nappe pile, were sur-
rounded by residual flysch troughs with turbidite deposi-
tion. Toward the south, the epicontinental sea stretched
over a large part of the internal zones of the Alpine and
Carpathian mountain chains, included the northern lithos-
phere fragment of ALCAPA and the southern fragment of
Tisza-Dacia (Csontos et al. 1992; Csontos 1995). The
lithospheric fragments (or microplates) moved towards the
subduction zone separately, their amalgamation set at first
since the Middle Miocene. The southern boundary of the
Central Paratethys Sea was represented by the Dinaride
mountain chain, dividing it from the Mediterranean Sea.

In late Early Miocene, the subduction gradually convert-

ed to a collision from the west to the east. The weight of the
overriding Carpathian orogene front and deep subsurface
load of the submerging plate led to development of a flex-
ure on the platform margin. The foredeep basin developed
along the whole front of the Outer Carpathian accretion
wedge. The evolution of the accretion wedge was associat-
ed with compression, controlling folding and thrusting of
the Flysch Belt nappe piles (Kováč et al. 1998).

The internal units of the Carpathians, belonging to the

ALCAPA and Tisza-Dacia microplates, started to collapse
due to stretching in consequence of the subduction pull
(Royden 1993a,b), as well as due to asthenospheric mantle
upheaval in the western part of the back-arc region. The
extension led to initial rifting of the Pannonian basin sys-
tem (Horváth 1993). By basin opening, besides normal
and low angle faults, strike slip faults also played an im-
portant role (Vass et al. 1988, 1993; Tari et al. 1992;
Fodor 1995; Kováč et al. 1998; Konečný et al. 2002). In
the west a sinistral shear dividing the Alpine and Car-
pathian orogenes opened the Vienna Basin, in the east a
dextral shear along the external and internal Carpathians
boundary (Pieniny Klippen Belt) opened the Transcar-
pathian Basin towards the northern part of the East Slovak
Basin. Extension in the western part of the back-arc region
led to beginning of the formation of the Danube Basin, as-
sociated with structural unroofing of the deepest Alpine-
Carpathian structural units.

During the Karpatian, a new marine connection opened

between the Central Paratethys and the Mediterranean Sea.
This connection is supposed through the trans-Dinaride cor-
ridor situated in the area of Slovenia and northern Croatia
(Rögl 1998). Apart from tectonics, the global sea-level rise
during the late Burdigalian had an important role in the de-
velopment of this seaway (TB 2.2 cycle, sensu Haq et al.
1988; Hardenbol et al. 1998; Kováč et al. 2001). The sea
transgression, with new elements of marine fauna and flora,
flooded the present territory of the Drava and Sava Basins
(Pavelić 2001), from where the sea penetrated into the
Mura, Zala and Styrian Basins. The NE oriented flooding
then followed the way between the northern margin of the
Mecsek Mts and the southern margin of the Transdanubian
Range reaching the North Hungarian—South Slovak sedi-
mentary area. Northwards, the sea spread to the Bánovská
kotlina Depression, the Vienna Basin, the Váh river valley
and the East Slovak Basin (Kováč et al. 1993). The Karpa-
tian sea covered especially the western part of the Car-
pathian Foredeep, in the east the sea extended especially
over the area of the present Outer Carpathian units of accre-
tionary wedge with wide marine connections into the East
Slovak Basin (Rudinec 1989, 1990; Kováč et al. 1995).

The DEM paleogeographical model of the Central

Paratethys during the Karpatian

(Fig. 2) documents the

beginning of the Carpathian orogene uplift. The ratio be-
tween continental and marine environments (ratio of land
and surface covered by the sea) can be very roughly inter-
preted due to the enormous erosion of the Early Miocene
sediments at the begin of the Middle Miocene (Kováč et
al. 2003). The erosion is documented by a total absence of
marginal facies particularly in northern areas of the Cen-
tral Western Carpathians and by very sporadic findings of
the Karpatian sediments in the Outer Carpathians, folded
together with the Flysch Belt deposits (Cieszkowski 1992;
Oszczypko 2003). The results of study of fluid inclusions
also confirm erosion of 2 to 5 km thick pile of deposits
(Hurai et al. 2002), as well as an important angular uncon-
formity between the Karpatian and Badenian strata at
many places in the Pannonian basin system.

The Carpathian paleo-relief was probably low at this

time. In many places, the pre-Tertiary basement units were
covered by Paleogene and Early Miocene sediments,
much larger in extent than those preserved today in the
Paleogene and Neogene basins. In the Western Car-
pathians, for example, a continuous sedimentary area cov-
ering the territory from the Bánovská kotlina Depression
to the Vienna Basin has been recorded (Kováč et al. 1993),
without indication of an uplift of the Považský Inovec
core mountain (Kováč et al. 1994, 1997).

The paleo-river net started to develop in areas with

higher relief. The Eastern Alps belonged to such places,
where rivers fed deltas in the Alpine Foredeep and the
southern part of the Vienna Basin (Aderklaa Formation),
and later in the uplifted parts of both the Central Western
Carpathians and the Outer Carpathians in the western seg-
ment of the Carpathian collision zone (Kováč 2000).

Considerations about prevailing low paleo-relief are

also supported by the paleobotanical study, which docu-

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KVAČEK et al.

ments an altitudinally not much differentiated character of
vegetation in the whole Carpathian-Pannonian region dur-
ing the Karpatian. Similarly, no obvious latitudinal and
longitudinal changes were observed there. Some zonation
marks, indicated by the diferentiation of the paleovegeta-
tion cover, are visible in the lowlands and margins of the
marine sedimentary area.

Karpatian vegetation

(Fig. 2) is documented by refer-

ence sites, of which only some have been dated by marine
fauna. The age of the others is indicated by the mammali-
an MN5 Zone and thus may overlap with the Early Bade-
nian. In other cases the boundary to the Ottnangian may
also be uncertain (MN4 Zone).

Three zonal forest formations were spread over the Cen-

tral Paratethys region during the Karpatian. Subtropical
broad-leaved forests with high to medium proportion of
evergreen elements were spread in the western part and
continued into the Boreal Province westwards in the form
of the typical Younger Mastixioid Floras sensu Mai
(1964). The corresponding phytostratigraphic unit of the
Boreal Province has been called the Floral Assemblage
(“Florenkomplex”) Františkovy Lázně – Kleinleipisch
(Mai 1995, 2001; Czaja 2003), but direct dating to the
lower part of the MN5 Zone is available only for Františkovy
Lázně in the Cheb Basin outside the Central Paratethys.
Domination of thermophilous elements (Engelhardia, Platy-
carya,  Sapotaceae, Symplocaceae, evergreen Fagaceae,
such as Trigonobalanopsis,  in pollen records expressed by
Castaneoideaepolis pusillus-, Castaneoideaepolis ovifor-
mis-,  Tricolporopollenites liblarensis-,  Quercoidites  henri-
ci-  types, etc.) is apparent for the Karpatian—Early  Badenian
time span (Planderová 1990, Zone MF5; Doláková &
Slamková 2003). Only in the central and northern parts
near mountains (e.g. Lipnica Mała, Nowy Sącz), the pro-
portion of the broad-leaved deciduous elements increased
resulting in the warm-temperate Mixed Mesophytic and
Broad-leaved Deciduous Forest types. Some sites (Mydlo-
vary – Knobloch & Kvaček 1996; Parschlug – Kovar et
al. 2004; Magyaregregy – Hably 2002) have a sub-hu-
mid character (subtropical forests with high proportion of
sub-humid and sclerophyllous elements). This may indi-
cate some heterochronity, which we are unable to resolve
from the paleobotanical record and short-time fluctuation
of perhumid and seasonal climate, as proposed by exother-
mic vertebrates (Böhme 2003). A distinct East-West gradi-
ent is apparent, when broader parts of Europe are
compared (Kovar-Eder et al. submitted).

Extrazonal mountain zones with conifers were probably

as high as 1500—2000 m a.s.l. and more. The upland for-
ests were dominated by Pinus, Abies, Cathaya and only at
still higher altitudes with the admixture of Cedrus, Tsuga,
and Picea, as demonstrated in the pollen spectra (Nagy
1992; Doláková & Slamková 2003). Various Pinaceae (Pi-
nus, Cathaya) and Sciadopitys entered also intrazonal
lowland formations.

Intrazonal coal-forming forests appeared mainly in the in-

ter-Alpine basins (Leoben-Bruck Basin, Parschlug, Fohns-
dorf, and Mecsek Mts – Hably 2002, p. 92). Besides
Glyptostrobus these thermophilous communities occasion-

ally included palms, e.g. Calamus-type at Teiritzberg, ever-
green oaks, Nyssa, Myrica,  Cyrilla  (Zittau Basin),  and not
yet clarified enigmatic Rhoipites pseudocingulum (= Rhus-
type). Among other mostly intrazonal elements, ferns of
Gleicheniaceae, Schizaeaceae (Lygodium), Osmunda and
Polypodiaceae  sensu lato (including, e.g. Pronephrium)
were well represented. Pollen of Avicennia  (Korneuburg Ba-
sin) suggests the presence of impoverished mangrove
shrubs in the NW part of the Central Paratethys.

Paleobotanical data are lacking for this time interval from

the eastern part of the studied region because of unfavour-
able conditions for the preservation of fossil plants there
(Syabryaj 2003). Assemblages from the upper part of the
Smoliarka Horizon (Rylova et al. 1999) and Rozhok (zone
IV FC sensu Yakubovskaya 1993) in Belarus give evidence
that the subtropical vegetation of the mastixioid type may
have extended north-eastwards from the Central Paratethys.

Middle Miocene – model of the Late Badenian

landscape and vegetation of the Central Paratethys

During the Middle Miocene, the active collision/subduc-

tion in front of the Carpathians shifted eastwards due to
gradual break-down of the submerging slab (Tomek & Hall
1993). In the west, this process led to the termination of col-
lision between the orogene and the European platform, fol-
lowed by gradual uplift of the Outer Carpathian
accretionary wedge and by the migration of foredeep depo-
centres from the Western Carpathian foreland towards the
Eastern Carpathians (Jiříček 1979). The sea flooding in the
front of the Carpathian orogene gradually schifted from
west to east. At the end of the Early Badenian, the sea aban-
doned the western part of the foredeep (Czech Republic),
marine sedimentary areas extended only in the northern
front of the Western Carpathians and in the front of the
Eastern Carpathians. The foredeep at the edge of the West-
ern and Eastern Carpathians reached its maximum extent
during Late Badenian—Early Sarmatian time, when 2500 m
of sediments were deposited (Meulenkamp et al. 1996). The
Sarmatian compression associated with uplift of the Outer
Western Carpathians draw the sea away from the northern
part of the Carpathian Foredeep for ever (Kováč et al. 1998).
A connection between the Carpathian Foredeep and the
Pannonian basin system remained preserved only in the
Eastern Carpathian region (Kováč 2000; Kováč et al. 1998).

The Middle Miocene development of the back-arc basin

region was controlled by two geodynamic factors: in the
western and central parts of the Pannonian basin system it
was upheaval of asthenosphere mantle masses, in the east-
ern part it was stretching of an overriding plate induced by
subduction pull in front of the Eastern Carpathians. In the
western part of the back-arc basin, subsidence of the Vienna
and Danube Basins caused depocentres above the thinned
crust and lithosphere associated with volcanic activity in
the hinterland of the Central Western Carpathians (Wernike
1985; Nemčok & Lexa 1990; Tari et al. 1992; Konečný et
al. 2002). In the east, orogene that was parallel to the back-
arc basin depocentres opened in the area of the Transcar-

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MIOCENE EVOLUTION OF LANDSCAPE AND VEGETATION IN THE CENTRAL PARATETHYS

pathian and Transylvanian Basins. Mighty acid and later
also “island arc” type volcanic activity appeared in a belt
along the eastern border of the back-arc region in the hinter-
land of the Eastern Carpathians (Konečný et al. 2002).

Synrift subsidence of the back-arc basin in the Pannonian

basin system was associated with mighty acid and calc-al-
kaline volcanic activity (Kováč 2000; Konečný et al.
2002). Individual depocentres formed in the extensional
tectonic regime. The main types were grabens and half-gra-
bens associated with normal and low angle faults, although
some pull apart basins opened along active strike-slip faults
(Vass et al. 1988, 1993; Tari et al. 1992; Fodor 1995; Kováč
et al. 1998; Konečný et al. 2002; Kováč 2000).

The Badenian marine conection of the Central Parat-

ethys with the Mediterranean Sea is supposed through a
trans-Dinaride corridor (Rögl 1998), at a similar place as
during the Karpatian. The Early Badenian sea-level rise,
which can be correlated with the early Langhian global
sea-level change (TB 2.3 cycle sensu Haq et al. 1988;
Hardenbol et al. 1998), is documented only in the SW part
of the Pannonian basin system – from the Styrian Basin
(Rögl et al. 2002). In the northern parts of the back-arc re-
gion wide-ranging erosion of Early Miocene deposits is
observed. In depressions (future basin depocentres), fan
deltas or terrestrial red coloured sediments were deposited
during this time.

The Early Badenian tectonically-controlled transgres-

sion, followed by rapid subsidence (deep sedimentary en-
vironment), started about 15 Ma. The basins were filled by
clastics transported by rivers running from uplifted areas
of the Eastern Alps and the Western Carpathians. After sea-
level fall (documented by erosion of the Early Badenian
carbonate platforms in the SW part of back-arc basin – Vi-
enna and Styrian Basins), the “Middle Badenian” transgres-
sion, which can be correlated with the global sea-level
change in late Langhian (TB 2.4 cycle sensu Haq et al.
1988; Hardenbol et al. 1998), took place. The Central
Paratethys sea reached the present extent in the intra-Car-
pathian Neogene basins, except the uplifted North Hun-
garian—South Slovak sedimentary area. Since that time, a
gradual filling up of the Pannonian basin system by deltas
has been observed, leading to shallowing of sedimentary
environment and development of isolated depocentres. Ba-
sins situated in the north and east suffered from isolation.
A salinity crisis took place in the Carpathian Foredeep as
well as in the Transcarpathian and Transylvanian Basins.

The Late Badenian transgression, which can be correlated

with the global sea-level change at the Langhian/Serraval-
ian boundary (TB 2.5 cycle, sensu Haq et al. 1988; Harden-
bol et al. 1998; 13.65 Ma.) represents the last full marine
flood of the Central Paratethys. Since the end of the Late
Badenian (12.7 Ma), an isolation of epicontinental sea in
the Intra-Carpathian region can be documented and a direct
connection of the Central Paratehtys with the Mediterranean
is not expected. The Sarmatian flood, from the Eastern
Paratethys region, can be correlated with the late Serraval-
ian global sea-level change (TB 2.6 cycle sensu Haq et al.
1988; Hardenbol et al. 1998). The sedimentary environ-
ment was dominantly shallow marine with decreased salinity.

The DEM model of the Central Paratethys during the

Late Badenian

(Fig. 3) documents uplift of the Western

Carpathians, including accretionary wedge in front of the
orogene and broad marine flood in the back-arc basin re-
gion. The intra-Carpathian region gained the characteris-
tic features of an archipelago sea, with many small islands
surrounded by shallow epicontinental sea. The paleo-
geography was still strongly influenced by tectonic pro-
cesses (Styrian phase), as it is well marked by rapid
changes of subsiding depocentres (sea bays, small basins)
and position of the coastal line.

The paleo-relief of the Carpathians significantly changed

during the Middle Miocene, due to strong tectonic influ-
ence. The belt of the Outer Carpathians, bordered by inter-
nal parts of the orogene, started to be uplifted. The land
surface was also strongly differentiated by volcanic activi-
ty, some stratovolcanoes reached heights of 2000 to
3000 m a.s.l. The river net transporting eroded clastic mate-
rial headed mostly towards the back-arc area.

Fair-sized altitudinal differences between lowlands and

mountains are documented by paleobotanical studies.
Mixed pollen spectra with mountain and lowland vegeta-
tion taxa indicate only seemingly the decrease of thermo-
philous taxa and the increase of more dominant temperate
taxa (Sitár & Kováčová-Slamková 1999; Slamková 2004).
The Badenian vegetation in the Central Paratethys can
generally be characterized as thermophilous without dom-
inance of typical boreal plant elements. Hence, we cannot
document sufficiently a gradual cooling of climate indi-
cated in the literature (Böhme 2003) that influenced the
European flora from the Late Badenian onwards.

Late Badenian vegetation

(Fig. 3) has been document-

ed from sites more often dated by marine fauna (due to an
extensive marine transgression), but also from some others
that are dated by mammals to Zone MN6 or by regional
correlation. The differentiation of the levels within the
Badenian has not always been accomplished and such
sites are exceptionally included into the map. Some sites
overlap with the lowermost Sarmatian, which is floristical-
ly hardly distinguishable (see Syabryaj & Stuchlik 2004;
merged into a single “Florenkomplex” Stare Gliwice-Un-
terwohlbach by Mai 1995).

The southernmost sites in Romania, Serbia and Hungary

differ from the remaining ones by thermophilous, partly
sub-humid aspects under subtropical climatic conditions,
which continued from previous times. A general cooling
trend appears in other sites by an increasing role of decidu-
ous elements, while thermophilous plants withdrew step-
wise southwards and only a part survived, mainly
Tetraclinis, Amentotaxus, Magnolia, Lauraceae, Engelhar-
dia and others. This is in the contrast to the preceding
“Wieliczien” thermophilous humid mastixioid assemblage
even in the Polish part of the Carpathian Foredeep (Łan-
cucka-Środoniowa & Zastawniak 1997). From the differ-
entiation of vegetation in the Late Badenian it is obvious
that the climatic gradient between the southern and north-
ern parts of the Central Paratethys increased at that time,
partly due to altitudinal differenciation, as noted above.
A noteworthy forest-forming tree was Fagus accompa-

303

MIOCENE EVOLUTION OF LANDSCAPE AND VEGETATION IN THE CENTRAL PARATETHYS

nied by other deciduous Fagaceae. Acer was diversified
in several species, among which Acer aegopodifolium
(syn. A. quercifolium –  see Ströbitzer-Hermann 2002;
Walther & Zastawniak 2005)  appears for the first time in
the Central Paratethys (Kovar-Eder et al. 1994), together
with  Ginkgo, Eucommia both arriving from Asia via the
Turgay migration route. Over most of the Central Parat-
ethys, warm-temperate Mixed Mesophytic Forest thrived,
only in the Western Carpathians and the Transcarpathian
Ukraine some sites already acquired the character of
Broad-leaved Deciduous Forest, probably due to influence
of mountains or due to cooling effect of intensive volca-
nic activity (Navrotskaja et al. 1991; Syabryaj 1992). The
proportion of herbs was generally low, not indicating low-
land open vegetation (Syabryaj & Stuchlik 2004).

Extrazonal mountain forests are well discernible in pol-

len assemblages, as at Devinská Nová Ves (Sitár &
Kováčová-Slamková 1999), where the coniferous belt re-
flected by the dominant pollen of Pinus includes addi-
tional high-mountain elements, such as Cedrus and
Tsuga. Its lower boundary may have decreased towards
1200 m a.s.l. However, this guess is only inferred from the
analogous Recent situation in the Colchis area (Stuchlik
& Kvavadze 1987; Klotz 1990).

In the intrazonal Glyptostrobus peat-forming forests,

Byttneriophyllum and Alnus constituted a basic communi-
ty, which became wide spread later in the Neogene.
Among riparian elements, Platanus leucophylla is occa-
sionally present. Lignite-forming communities are typical-
ly developed in intra-montane depressions (e.g. Handlová
and Nováky) and they are also common in the lowlands
outside the Paratethys area in Poland and Germany (Lusa-
tia seam 1). The corresponding phytostratigraphical level
in the Boreal Province is probably represented by the Flo-
ral Assemblage Schipkau-Konin sensu Mai (2001), but
this correlation is opposed by Krutzsch (2000.) The paleo-
floristic differentiation around the Late Badenian and Ear-
ly Sarmatian boundary has been discussed with little
success to give clear-cut differences based on plant
megafossils (Němejc 1951, 1967; Shvareva 1965; Sitár
1967, 1982). It is still uncertain, which Early Miocene ele-
ments did not enter the Sarmatian flora, where broad-
leaved deciduous trees predominate. In her pollen
assemblages, Planderová (1990) created a transitional
Zone MF7 for this type of flora in the Slovak Neogene. Its
pollen spectra include a very low proportion of or no ther-
mophilous Symplocaceae, Sapotaceae and Cyrillaceae.

Late Miocene – model of the Middle Pannonian

landscape and vegetation of the Central Paratethys

During the Late Miocene, the area of the Central Parat-

ethys gained paleogeographical features similar to the
present situation in the Carpathian-Pannonian region. The
main difference was represented by flooding of the intra-
Carpathian region and the foredeep depocentres, which
were restricted to the southeastern foreland of the Eastern
Carpathians in that time (Jiříček 1979; Meulenkamp et al.

1996). Thus a connection of the brackish Eastern Paratethys
with the Lake Pannon originated (Magyar et al. 1999).

The Late Miocene geodynamic development of the Car-

pathians can be characterized by termination of collision
between the Western Carpathian and the European plat-
form and reinforced collision connected with subduction
in front of the Eastern Carpathians. This process was fol-
lowed not only by the uplift of the accretionary wedge
loop, but also by the uplift of the whole Carpathian oro-
gene mountain chain.

Pull of the active subduction in front of the Eastern Car-

pathians southern edge led to the “second” rifting phase
in the back-arc basin at the begining of the Pannonian,
followed by thermal post-rift subsidence (Lankreijer et al.
1995; Kováč 2000; Konečný et al. 2002). Among the Late
Miocene basins of the intra-Carpathian domain formed in
an extensional regime, flexural basins without important
fault activity prevailed, although in the Early Pannonian
normal and strike slip faults allowed development of small
pull-apart basins (Vass et al. 1988, 1993; Tari et al. 1992;
Fodor 1995; Kováč et al. 1998; Kováč 2000; Konečný et
al. 2002). The Lake Pannon – the Pannonian basin sys-
tem was gradually filled up by deltaic deposits, generally
from the northwest toward the southeast. The sedimentary
environment gradually changed from a brackish deepwa-
ter to a shallow water – lake environment due to isolation
from the Mediterranean and Eastern Paratethys (Magyar et
al. 1999). At the northern margin of the Lake Pannon,
marshes, swamps and deltaic systems spread in an ever-
larger extent. Due to the retreat of the coastal line the
lacustrine environment changed generally into alluvial
also in the Late Pannonian. Later, in the Pliocene a broad
area of lowlads appeared in the hinterland of the Car-
pathian chain. Scattered mountains (e.g. the Transdanubi-
an Range Mts, Bükk Mts, Apuseni Mts) and basalt
volcanoes formed higher morphological elevations.

At the end of the Late Miocene, tectonic inversion of

the basin system (Horváth 1993, 1995; Horváth & Cloet-
ingh 1996) led to the retreat of the aquatic sedimentary
environment in the whole intra-Carpathian area, exept the
central and southeastern regions. Uplift of the Eastern
Alps, the Western and Eastern Carpathians was associated
with angular unconformity between the Middle and Late
Miocene (or younger) strata.

The architecture of the Late Miocene fill of the Pan-

nonian basin system was distinctly influenced by paleo-
geographical changes. From the sequence stratigraphy and
depositional systems point of view, the succession of the
Late Miocene sediments can be characterized at first by a
dominant portion of proximal deltaic deposits (A—C zones,
sensu Papp 1951), which pass upward into distal deltaic
to basinal fine-grained clay—silt—sandy facies (D, E zones).
Fluvio-lacustrine sediments with coal seams (F—H zone)
formed the terminal part of the Pannonian. The overlying
Pliocene strata were deposited mostly in an alluvial sedi-
mentary environment. The Pannonian cyclicity can be cor-
related by global changes Tor-1 cycle (11.6—9.3 Ma, sensu
Hardenbol et al. 1998) and Tor-2 cycle (9.3—7.2 Ma, sen-
su Hardenbol et al. 1998).

304

KVAČEK et al.

The DEM model of the Central Paratethys during the

Middle Pannonian

(Fig. 4) documents the extinction of

the sedimentary area of the Carpathian Foredeep, except
its southeastern part and uplift of the Carpathian mountain
chain. At that time, a maximum extent of brackish seawa-
ters covered the intra-Carpathian domain.

The expected paleo-relief of the Carpathian orogenic

chain in the Late Miocene began to match the present-day
situation, characterized by the presence of both mountains
and lowland areas. A difference can be seen in the more el-
evated belt of the Outer Carpathian accretionary wedge,
which formed a natural barrier between flat territories of
the European platform and the ever-subsiding Pannonian
basin system covered by the Lake Pannon. During the
Pannonian and Pliocene, basalt volcanic activity also par-
ticipated in paleo-orography of the back-arc region
(Kováč et al. 1998; Kováč 2000).

The Middle Pannonian vegetation

(Fig. 4) corresponds

to the warm temperate climatic zone with evidence of spo-
radically present termophilous and evergreen taxa. A higher
percentual proportion of non-arboreal pollen (10 to 14 %)
indicates local vegetation of partly open woodland (i.e.
woods with open canopy). An increase of halophytic taxa
documents the presence of coastal lagoons and marshlands
during the lowstand of the brackish sea. Swamp vegetation,
which grew directly on swamp substrates, is characterized
mainly by noteworthy Taxodiaceae trees. They are often
present in the association with Myricaceae and subordinary
Nyssaceae. The riparian forest elements subdominantly oc-
curred with Alnus and Ulmus. The extrazonal vegetation of
the mountain areas with Picea, Tsuga, Abies, Cedrus is well
represented in the pollen spectra.

During the Late Pannonian, the Western Carpathian paleo-

geography started to change. The Lake Pannon withdrew
southwards, the nothern margin of the back-arc basin was
slightly uplifted with the progradation of deltaic and alluvial
facies, especially in lowlands. These areas were often covered
by hygrophilous plants: Myrica, Salix, Ulmus, Alnus. Herbs
were represented by  Chenopodiaceae, Asteraceae, Ericaceae,
Poaceae  and  Artemisia.  Unevenly  high moutain relief of the
uplifted mountain chains created ideal conditions for the
mixed mesophytic forests with Carya,  Quercus,  Craigia,
Carpinus, Fagus,  Picea, Abies, Tsuga and Pinus.

The reference sites considered on the map have been as-

signed to the Pannonian zones C—E sensu Papp 1951,
namely the time slice before the major spreading of the
lignite facies over the Pannonian Basin (zone F). The dat-
ing is based mostly on molluscs or regional correlation,
rarely on mammals (MN9 Zone). Additional sites of Early
Pannonian age (zones A—B) are shown in brackets on the
map. They indicate the previous vegetation type at the
Sarmatian/Pannonian boundary. Most leaf assemblages of
Pannonian age are at least partly intrazonal and it is diffi-
cult to obtain a true picture of zonal vegetation from them.
Most spore/pollen assemblages, mainly from the Hungari-
an Pannonian, have not been revised and the stratigraphic
position of pollen samples remains partly uncertain.

At the very beginning of the Pannonian, subtropical condi-

tions returned to some parts of the Central Paratethys. The

thermophilous vegetation from southern Austria (Mataschen,
Styrian Basin, Pannonian B – Kovar-Eder 2004) can also be
found to the north-western periphery outside the Paratethys
(Gozdnica) and may correspond to the Floral Assemblage
(“Florenkomplex”) Düren sensu Mai (1995) in western Eu-
rope with the latest occurrences of mastixioid plants. Howev-
er, the dating of Gozdnica is still under dispute (see Dyjor et
al. 1992, 1998; Mai, personal communication).

Later in the Pannonian, thermophilous evergreen, partly

sub-humid vegetation remained in the south and south-east-
ern parts (Serbia, southern Hungary, the Borod Basin in Ro-
mania). In the Middle Pannonian, broad-leaved deciduous
and Mixed Mesophytic warm-temperate to temperate for-
ests with a low proportion of evergreen elements generally
became widespread over the Central Paratethys (Styrian Ba-
sin, Molasse Zone, Vienna Basin). Characteristic elements
in these communities included Fagus haidingeri—plioceni-
ca  complex, Quercus (?Castanea) kubinyii, Quercus
pseudocastanea—pseudorobur  complex, Carpinus sp. div.,
Betula, Acer integrilobum, Acer vindobonense and Acer
subcampestre  (syn.  Acer jurenakyi)  (Kovar-Eder 1988;
Ströbitzer-Hermann 2002; Ströbitzer-Hermann & Kovar-
Eder 2003). Cooling trends can be traced in spore/pollen
spectra from the cores in the Pannonian Basin (Nagy &
Planderová 1985; Nagy 1992). At some sites (Alsóvadász,
Suchohrad) somewhat higher frequencies of Chenopodi-
aceae and Artemisia  (max. 15 %) may indicate patches of
herbaceous vegetation on marshes within broad-leaved
deciduous forests. However, high mean annual precipita-
tion (Bernor et al. 2003) prevented the development in
this aera of open woodland and grassland vegetation like
that suggested for south-eastern to southern Ukraine out-
side the Central Paratethys (Syabryaj 1999, 2003). Tem-
perate purely deciduous broad-leaved forests were also
spread during the Middle Pannonian in the Western Car-
pathians (e.g. sites at Nové Ustie in the Orava Basin and
Martin in the Turiec Basin). In most cases these assem-
blages include a considerable proportion of intrazonal ele-
ments, both woody and aquatic (Trapa). Characteristic
riparian woody elements were Salicaceae, Platanus leuco-
phylla, Alnus ducalis, Alnus cecropiifolia,  and partly in-
trazonal  Quercus gigas and Pterocarya paradisiaca. The
dominant peat-forming community during the whole Pan-
nonian consisted mostly of the Glyptostrobus-Byttnerio-
phyllum-Alnus  swamp forest. This swampy coal-forming
vegetation reached its widest distribution over the Pan-
nonian Basin during the Late Pannonian, as the large ex-
tension of the lignite facies in Hungary, southern Moravia,
Slovakia and Serbia corroborates (Knobloch 1969; Gi-
vulescu 1992; Pantić & Dulić 1993; Hably 2003).

Extrazonal montane conifers of all kinds (including

Tsuga, Cedrus, Picea, Abies, Keteleeria) were found in
pollen spectra throughout the Pannonian Basin as regular
accessories. In the intra-montane basins macrofossils of
these conifers were also formed (Nove Ustie, Martin).
Therefore we may expect high mountain conifer belts
reaching over 1500 m a.s.l. on the Carpathian and Alpine
ridges and descending to medium altitudes, mixed with
deciduous broad-leaved elements.

306

KVAČEK et al.

Paleoclimatic trends

Palaeoclimatic proxies are available for many sites con-

sidered in the presented maps (see Mai 1995). Most of
them have been derived from “intuitive” comparisons
with extant vegetation, while the objective co-existence
and leaf physiognomical (CLAMP) methodologies have
not been largely applied so far.

Broad-leaved evergreen forests, including the “Late Mas-

tixioid Floras”, were usually compared with similar forests
in monsoon East Asia. In general, the climate corresponding
to this type of vegetation was humid to per-humid (annual
sum of precipitation about 1000 to 3000 mm), with heavy
rains prevailing in the summer, but without any month with
a humidity deficit. The temperature of the coldest month
varied between 4—10 ºC and absolute minima rarely
reached under zero. The Mean Annual Temperature (MAT),
according to various estimates (e.g. Mai 1995) varied in
larger spans, according to the percentage of evergreen taxa,
from 13 to almost 20 ºC. The climate must have been quite
equable, with a range of temperature less than 25 ºC. This is
a prevailing condition in the Karpatian to Lower Badenian
stages over the Central Paratethys. Only in some parts, high
summer mean temperatures over 20 ºC and also dry substrate
caused a relative humidity deficit and local expansions of
sub-humid sclerophyllous (microphyllous) evergreen forests.
This type of forest has nothing to do with the etesian (Medi-
terranean-type) climate and was probably more similar to
the microphyllous montane forests in drier parts of the Hi-
malayas today.

The cooling trends expressed by the decline of MAT in

the Late Badenian over northern and eastern parts of the
Central Paratethys caused changes in the forests. Ever-
green elements mostly withdrew southwards and only less
frost-sensitive trees remained forming the Mixed Meso-
phytic Forest with high representation of deciduous taxa.
MAT, under which this type of vegetation optimally
thrives, is variously estimated, depending on what type of
forests is brought for comparison – East Asiatic or North
American (Wolfe 1979). While in the former, the Coldest
Month Mean Temperature (CMMT) is usually quite low
(up to —2 ºC), the latter thrives in warmer, subtropical con-
ditions (northern Florida – mean annual temperature ca.
20 °C, January mean up to 10 ºC). In general, the follow-
ing estimation of the decline in temperature can be ex-
pected for the Central Paratethys from the published data
(e.g. Mai 1995) – MAT 16(—?10) ºC, that is lowering by
about 3 ºC in comparison with the Karpatian, and ade-
quate lowering of the January mean, which may have been
even stronger, because of decrease of climatic equability.
The sum of Mean Annual Precipitation (MAP) remained
high enough for humid conditions in any case, also thanks
to the colder climate. Such warm temperate conditions
predominated during the Late Badenian and the Middle
Pannonian. In the Pannonian in northern and easterly parts
of the area studied, particularly near the mountains, the
Deciduous Broad-leaved Forests indicate still more severe
deterioration of climatic conditions. These temperate for-
ests dominated by deciduous oaks and beech and inter-

mixed with various Pinaceae withstood decrease of
CMMT up to —10 °C, particularly on higher altitudinal
habitats. The impact of such severe winters was certainly
milder due to heavy snows, as is the case today in East
Asia, particularly in Japan. Equally high precipitation
throughout the year also prevented expansion of herba-
ceous steppe vegetation, although patches of it can be no-
ticed in the pollen record within deciduous broad-leaved
forests in the Middle Pannonian. Due to high humidity of
climate, extensive lignite deposits originated over most of
the Paratethys area and its north and west periphery.

Conclusions

After having compiled the presented Miocene geobotani-

cal maps according to the methodology applied above, the
following conclusions can be drawn for the future research
in other areas and time slices of the Cenozoic.

Contrary to various models of ancient Cenozoic vegeta-

tion that rely on the physiognomy and composition (di-
versity) as well as abundance of elements (e.g. Wolfe
1979; Mai 1995), the presented system of vegetation units
is much more simplified. It surely suffers from various de-
ficiencies. It neglects abundance. But this parameter is, in
our opinion, not objectively derivable from frequencies of
fossils in a given site or core level. Another weak point of
the system employed above is that individual elements
with a broader ecological span can enter more units or
they are transitional and their foliar physiognomy (ever-
green vs. deciduous) cannot be precisely identified (e.g.
Symplocos, Engelhardia). The defined vegetation forma-
tions were certainly not profoundly clear-cut in ancient
landscapes and transitions between them existed. Still the
diversity percentages are most objective characteristics for
a given assemblage and can be verified any time, the as-
signment into the system of the defined vegetation forma-
tions as characterized above (see also Kovar-Eder et al.
submitted) is easy and mostly unequivocal.

According to our experience, it is advisable to use pa-

leogeographical and geobotanical maps of narrow time
slices, because they reveal better consistent patterns of
vegetation and its dynamics in spite of fewer reference lo-
calities. When a longer time interval has been considered,
local differences between the sites sometimes expressed
trends in time rather than climatic gradients in space (see
Fig. 3 for the Early-Middle Pannonian).

Megafossil and spore/pollen plant records were com-

bined, whenever feasible, to gain better understanding of
taxonomy and ecology of elements composing assem-
blages. Of course, great problems exist, concerning how to
transfer spore/pollen diagrams with various enigmatic
plant elements into a formation with known physiogno-
my. In the fututre, it would be desirable to unify views on
the taxonomy of those elements of uncertain affinities
both in megafossil and spore/pollen records.

Older data of paleobotanical and palynological research

have not been neglected but revised and transferred to a
common nomenclature, when the documentation (illustra-

307

MIOCENE EVOLUTION OF LANDSCAPE AND VEGETATION IN THE CENTRAL PARATETHYS

tions, preparations) was available and re-studied. In many
cases, such assemblages and their elements were wrongly
interpreted, or wrongly assigned to the natural system. Yet
the current progress in knowledge of whole fossil plants is
improving these inconsistencies. Actuopalynological stud-
ies of Recent vegetation (e.g. Stuchlik & Kvavadze 1987;
Kvavadze & Stuchlik 1990, 1993) offered important clues
for converting spore/pollen spectra into various types of
real vegetation. It would be worth attempting to apply for
the same sets of fossil data the new methodology pre-
sented in this account, which is based on proportions of
components and diversity, along with that currently em-
ployed by palynologists, which uses abundance percent-
ages of elements.

Acknowledgments:

We appreciate discussion, data and

exchange of views with our colleagues L. Stuchlik,
Kraków, V. Mosbrugger and A. Bruch, Frankfurt am
Main and I. Magyar, Budapest. L. Stuchlik, L. Hably and
D. Ivanov as the reviewers and P. Bosák as the editor who
suggested useful improvements to the first version of the
text. This research has been sponsored by the European
Science Foundation (Project EEDEN), by the Czech
Grant Agency GAČR (Project No. 205/04/0099), Slovak
Grant Agency VEGA (Projects No. 2/5016/05, 1/2035/05,
1/0080/03), project of the Slovak Research and Devel-
opment Support Agency APVV-51-011305 and the Slo-
vak Ministry of Education (Project No. AV/808/2002).

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