GeolCarp_Vol65_No3_227

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, JUNE 2014, 65, 3, 227—239 doi: 10.2478/geoca-2014-0016

Introduction

The  Lower  Miocene  deposits  of  the  Karpatian  stage  (late
Burdigalian)  in  the  Central  Paratethys  contain  rare  plant
macrofossils,  with  only  a  few  directly  dated  localities  in
Austria and Moravia (Knobloch 1967, 1968; Kvaček 2003).
However,  diverse  Miocene  plant  assemblages  of  Western
Europe (Mai 1995, p. 367) that are not directly dated may be
actually coeval with the late Early—Middle Miocene and may
correspond to the “Early-Middle Miocene Optimum” (Flower
& Kennett 1994; Zachos 2001).

Although highly detailed palynological datasets are avail-

able from the Karpatian in Slovakia (Planderová 1967, 1990)
and Moravia (Doláková & Slamková 2003), macrofloral as-
semblages of the Karpatian in the Central Paratethys remain
largely  unknown.  Our  goal  is  to  describe  a  new,  relatively-
well preserved plant assemblage of Early Miocene age from
the Vienna Basin (western Slovakia) and to reconstruct paleo-
vegetation  and  ecosystem  conditions  during  the  late  Early
Miocene  in  the  eastern  part  of  the  Vienna  Basin  (Central
Paratethys).  The  new  plant  assemblage  implies  that  it  was
probably  derived  from  coastal-marsh  habitats  and  allows
comparisons  with  other  assemblages  in  the  Central  Para-
tethys (e.g. Kovar Eder et al. 1998, 2008; Bruch et al. 2004,
2011; Kern et. al. 2011). Our study also contributes to better

Lower Miocene plant assemblage with coastal-marsh

herbaceous monocots from the Vienna Basin (Slovakia)

ZLATKO KVAČEK

, VASILIS TEODORIDIS

1, 2

, MARIANNA KOVÁČOVÁ

, JÁN SCHLÖGL

and VILIAM SITÁR

Institute of Geology and Paleontology, Faculty of Sciences, Charles University in Prague, Albertov 6, 128 43 Prague 2, Czech Republic;

zlatko.kvacek@gmail.com

Department of Biology and Environmental Studies, Faculty of Education, Charles University in Prague, Magdalény Rettigové 4,

116 39 Prague 1, Czech Republic; vasilis.teodoridis@pedf.cuni.cz

Department of Geology and Paleontology, Faculty of Natural Sciences, Comenius University in Bratislava, Mlynská dolina G,

842 15 Bratislava, Slovak Republic; kovacova@fns.uniba.sk; schlogl@fns.uniba.sk

(Manuscript received December 19, 2013; accepted in revised form March 11, 2014)

Abstract: A new plant assemblage of Cerová-Lieskové from Lower Miocene (Karpatian) deposits in the Vienna Basin
(western Slovakia) is preserved in a relatively deep, upper-slope marine environment. Depositional conditions with high
sedimentation rates allowed exceptional preservation of plant remains. The plant assemblage consists of (1) conifers
represented by foliage of Pinus hepios and Tetraclinis salicornioides, a seed cone of Pinus cf. ornata, and by pollen of the
Cupressaceae, Pinaceae, Pinus sp. and Cathaya sp., and (2) angiosperms represented by Cinnamomum polymorphum,
Platanus neptuni, Potamogeton sp. and lauroid foliage, by pollen of Liquidambar sp., Engelhardia sp. and Craigia sp., and
in particular by infructescences (so far interpreted as belonging to cereal ears). We validate genus and species assignments
of the infructescences: they belong to Palaeotriticum Sitár, including P. mockii Sitár and P. carpaticum Sitár, and probably
represent herbaceous monocots that inhabited coastal marshes, similar to the living grass Spartina. Similar infructescences
occur in the Lower and Middle Miocene deposits of the Carpathian Foredeep (Slup in Moravia), Tunjice Hills (Žale in
Slovenia), and probably also in the Swiss Molasse (Lausanne). This plant assemblage demonstrates that the paleovegetation
was represented by evergreen woodland with pines and grasses in undergrowth, similar to vegetation inhabiting coastal
brackish marshes today. It also indicates subtropical climatic conditions in the Vienna Basin (central Paratethys), similar to
those implied by other coeval plant assemblages from Central Europe.

Key words: Early Miocene, Karpatian, Vienna Basin, Western Slovakia, paleobotany, palynology.

understanding of climate during the Early and Middle Mio-
cene in this region.

Geographical and geological setting

Samples were collected in the western part of the Slovak

Republic at the Cerová-Lieskové locality. The outcrop is sit-
uated in the foothills of the Malé Karpaty Mountains, which
represent the eastern margin of the central part of the Vienna
Basin (Fig. 1A—C). The Vienna Basin was a part of the Cen-
tral Paratethys Sea during the Miocene. Karpatian (late Bur-
digalian)  sediments,  assigned  to  the  Lakšárska  Nová  Ves
Formation (Špička & Zapletalová 1964), are well exposed in
a  former  clay  pit.  These  sediments  are  represented  by  mas-
sive,  locally  laminated,  calcareous  clays  and  clayey  silts
with  thin  tempestite  intercalations  (up  to  10 mm  thick)  and
several thin sandstone/siltstone layers. The section is almost
20  meters  thick.  The  macrofossil  assemblage  consists  of
vertebrates, mainly teleosts, and a wide spectrum of inverte-
brates  –  bivalves,  gastropods,  scaphopods,  cephalopods,
decapods,  isopods,  and  barnacles  (Harzhauser  &  Schlögl
2012),  regular  and  irregular  echinoids,  asteroids,  siliceous
sponges  (Lukowiak  et  al.  2013),  and  solitary  corals.  Plant
remains  are  also  locally  abundant.  Microfossil  assemblages
include  benthic  and  planktonic  foraminifera,  radiolarians,

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sponge spicules, bryozoans, ostracods, crinoid ossicles, co-
leoid statoliths, fish otoliths, shark teeth (Underwood &
Schlögl 2013), and locally extremely abundant diatoms.

These clays and silts were deposited under upper bathyal

conditions  in  a  relatively  oxygen-depleted  environment
(Schlögl et al. 2011). Foraminiferal assemblages from the in-
tervals  14—20  (Fig. 1C)  were  analysed  with  the  two-step
depth equations of Hohenegger (2005) and yielded a paleo-
depth  range  between  240  and  330  meters  (Schlögl  et  al.
2011).  The  upper  bathyal  aphotic  zone  is  also  reflected  by
gastropod  and  bivalve  assemblages,  with  the  predominance
of carnivorous gastropods (along with scavengers and para-
sites),  and  bivalves  composed  mainly  of  chemosymbionts,
and detritus and suspension feeders (Harzhauser et al. 2011).
The  shark  fauna  mainly  of  small-sized  bathyal  squaliform
sharks,  with  pelagic  lamniform  sharks  and  sharks  from  the
Centrophoridae and Hexanchidae families completely absent
(Underwood  &  Schlögl  2013).  Decapods  (Hyžný  &  Schlögl
2011) and siliceous sponges (Lukowiak et al. 2013) also im-
ply  bathyal  conditions.  Assemblages  of  benthic  foraminifera
are dominated by infaunal and deep-infaunal, suboxic/dysoxic
taxa,  in  some  samples  only  with  minor  contribution  of  oxi-
phyllic  taxa.  Benthic  Foraminiferal  Oxygen  Index  (BFOI)  of

Kaiho (1994) indicates dysoxic and low-oxic conditions dur-
ing the deposition of the intervals 14—20 (Schlögl et al. 2011).

The exceptional preservation of easily disarticulating ani-

mals such as ghost shrimps, minute pilumnid crabs, or ciro-
lanid isopods implies a relatively high net sedimentation rate
(Hyžný & Schlögl 2011; Hyžný et al. 2013). Thin sandstone/
siltstone layers (up to 5 cm), silt “pavements” or laterally re-
stricted  thin  silt  lenses  generally  very  poor  for  organic  re-
mains  probably  represent  distal  turbiditic  deposits.  Thin
laminae up to 10 mm in thickness, rich in molluscs, echino-
derms  and  wood  debris  most  probably  represent  tempestite
deposits. These are frequently dominated by one or two fos-
sil groups, most commonly irregular echinoids and/or ptero-
pods. Deposits also contain various redeposited microfossils
from  areas  surrounding  the  Vienna  Basin,  including  well
preserved,  most  probably  Lower  Cretaceous  freshwater  os-
tracods (Pipík et al. 2010). These allochthonous remains and
plant material were apparently transported by rivers from ad-
jacent emerged land into the basin.

Co-occurrence of the foraminifera Uvigerina graciliformis

Papp & Turnovsky, 1953 and Globigerinoides bisphericus
Todd in Todd, Cloud, Low & Schmidt, 1954 and the absence
of the genus Praeorbulina Olsson, 1964 suggest Late Karpa-

Fig. 1. A – Position of the Vienna Basin in the Carpathian—Pannonian system: 1 – European platform units, 2 – Carpathian-Alpine ex-
ternides, 3 – Pieniny Klippen Belt, 4 – Alpine-Carpathian-Dinaride and Pannonian internids, 5 – Neogene volcanics, 6 – Neogene basins;
B – Bükk, NCA – Northern Calcareous Alps, TCR – Transdanubian Central Range. B – Location of the Cerová-Lieskové clay pit, in-
dicated by arrow. C – Simplified section through the Cerová-Lieskové clay pit: a – Massive calcareous clay, b – Thin tempestite layers
with plant debris, and lithified clayey silt layers, c – Thin siltstone/sandstone layers, and silt lenses (modified from Łukowiak et al. 2013).

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tian age (Cicha & Rögl 2003; Berggren et al. 1995; see also
Schlögl et al. 2011). The regional Paratethyan Karpatian
stage is the time-equivalent of the latest Burdigalian (Rögl et
al. 2003; Piller et al. 2007).

Material and methods

Several stratigraphic intervals contain dispersed or accu-

mulated  plant  debris  that  shows  two  major  preservation
states  (Fig. 1C).  The  first  one  is  represented  by  abundant
plant  debris  associated  with  various  invertebrate  remains
within  thin,  most  probably  tempestite  layers.  The  debris  is
poorly  preserved,  more  or  less  fragmented  due  to  transport
and  mainly  conposed  of  wood  fragments.  The  second  one
represents  scattered,  but  generally  better  preserved  leaves,
grass-blades,  fir-needles  and  wood  fragments.  These  well-
preserved  remains  are  studied  here  (coming  from  inter-
vals 13—14,  14—15,  15—16,  16—17,  19—20,  30—31,  43,  see
Fig. 1C).  Nine  intervals  were  selected  for  the  palynological
analysis  out  of  47  intervals  sampled  for  benthic  and  plank-
tonic  foraminifers  (samples 12,  18,  20,  23,  30,  33,  46,  49,
and interval 49—50). Most of the samples are either barren or
contain very few and poorly preserved pollen grains. There-
fore, palynological data are inappropriate for statistical eval-
uation and paleoenvironmental interpretation.

Sample preparation followed standard laboratory methods

(Erdtman 1954; Faegri & Iversen 1989; Moore et al. 1991).
The description of the taxa follows the system based on mo-
lecular phylogeny (e.g. Judd et al. 2002; Reveal 2012). Leaf
architecture  is  described  using  current  terminology  pub-
lished by Ash et al. (1999) and Ellis et al. (2009). Below, we
document  the  composition  of  the  flora  and  rectify  the  no-
menclature of the previously published data (Sitár 2001). All
the  investigated  material  is  housed  in  the  Slovak  National
Museum  in  Bratislava  (SNM B).  We  compare  this  assem-
blage with similarly preserved plant remains from the Karpa-
tian Slup holostratotype locality (southern Moravia), housed
in the Moravian Museum, Brno (MZM).

Systematic paleobotany

Pinaceae

Pinus L.

Pinus hepios (Unger) Heer

Fig. 2.1

1850 Pinites hepios Unger, p. 362
1852 Pinites hepios Unger – Unger, p. 97, pl. 35, figs. 6—8, 9
1855 Pinus hepios (Unger) Heer, p. 57, pl. 21, fig. 7

M a t e r i a l : A single incomplete fascicle of needles

(SNM B 1792/a,b), several needle fragments.

D e s c r i p t i o n : A single isolated two-needled fascicle,

needles fragmentary, 56.5 to 37.8 mm long and 0.6 to 0.8 mm
broad, straight or slightly curved, secondarily compressed,
needle apices, bases and the sheath not preserved, margin en-
tire, venation consisting of one medial vein (to 0.3 mm broad),
parallel striation poorly observed.

D i s c u s s i o n : Foliage of Pinus hepios (Unger) Heer is

usually associated with seed cones of  P. ornata (Sternberg)
Brongniart (e.g. Engelhardt 1876a; Menzel 1901). P. hepios
usually  represents  a  cumulative  taxon  name  for  pines  with
two-needle fascicles. It was based by Unger (1850, 1852) on
foliage  and  seeds  from  the  locality  Parschlug  (e.g.  Kovar-
Eder et al. 2004). Recently, Teodoridis & Sakala (2008) re-
vised  cone  and  foliage  records  of  P.  hepios/P.  ornata  from
the Paleogene and Neogene deposits of the Czech Republic
described  by  Sternberg  (1825),  Engelhard  (1876a),  Menzel
(1901) and Knobloch et al. (1996). They considered morpho-
logical  affinities  to  other  pine  species  assigned  to  P. engel-
hardtii Menzel emend. Mai/P. rigios (Unger) Ettingshausen,
Pinus urani  (Unger)  Schimper  and  Pinus  sp.  div.  described
from North Bohemia. Bůžek et al. (1996) recorded isolated 2
or 3-needled fascicles as P. hepios from several drill-cores of
the  Lower  Miocene  Cypris  Formation  in  the  Cheb  and
Sokolov basins of North Bohemia. This record of P. hepios
is  of  Ottnangian  to  Karpatian  age  and  corresponds  strati-
graphically to the plant assemblage of Cerová-Lieskové. It is
associated with foliage of P. rigios and P. cf. saturni Unger,
and with male and female cones and isolated seeds of Pinus
sp.  div.  (Bůžek  et  al.  1996).  Other  coeval  deposits  of  the
Berzdorf Basin (Saxony, Germany) contain foliage of P. cf.
hepios, P. cf.  palaeostrobus Ettingshausen and P. cf. rigios
associated with seed cone of P. spinosa Herbst (Czaja 2003).
Mai  (1986)  revised  the  type  and  original  material  of  Pinus
species  from  the  Paleogene  and  Neogene  strata  of  Europe,
stressing that mainly seed cones allow reliable identification
to species level. Teodoridis & Sakala (2008) suggested that
the xerophytic P. halepensis Mill. (Mediterranean and West-
ern Asia) and P. merkusii Jungh. & de Vries (Vietnam, Laos,
Cambodia,  China,  Philippines,  Malaysia  and  Indonesia)  are
the extant species that are most comparable to the pine repre-
sented by the fossil species of P. hepios/P. ornata.

Pinus cf. ornata (Sternberg) Brongniart

Fig. 2.2—3

1825 Conites ornatus Sternberg, p. 39, pl. 55, figs.1, 2
1828 Pinus ornata (Sternberg) Brongniart, p. 107
1901 Pinus ornata Sternberg – Menzel, p. 54, pl. 2, figs. 6—7, 9
1994 Pinus ornata (Sternberg) Brongniart – Mai, p. 213, pl. 3, figs. 1—3,

text-fig. 1b

M a t e r i a l : An incomplete seed cone (SNM B 1809).
D e s c r i p t i o n : Incomplete seed cone symmetrical, prob-

ably oblong, 79.0 mm long, 16.8 mm broad, apophysis
rhombic, flat to slightly arched, 5.9—(6.9)—8.1 mm long and
5.8—(7.1)—9.9 mm broad, distinctly keeled and radially striated,
umbo rhombic up to 3.4 mm in diameter flat to slightly
arched, excentro-denticulatomucronate, mucro small and in-
distinctly erected.

R e m a r k s : P. ornata was described on the basis of a seed

cone  from  the  Lower  Oligocene  locality  Valeč  (Sternberg
1825,  p. 39,  pl. 55,  figs. 1—2)  but  the  holotype  disappeared
in  the  late  19

century. Additional seed cones were de-

scribed later from North Bohemia, particularly from the
Valeč type locality (Menzel 1901), but also from other Lower
Eocene to Lower Miocene localities of the České středohoří

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Mts, Staré Sedlo, Český Chloumek and Žitenice (Engelhardt
1876b; Knobloch 1962; Knobloch et al. 1996; Teodoridis et
al. 2012) and the Most Basin, Hradiště near Černovice (En-
gelhardt 1876a, 1877) and Břeš any (Menzel 1901). Němejc
(1968, p. 384, pl. 39, fig. 3) re-figured Menzel’s cone from
Valeč which was suggested as the neotype by Kvaček &
Kvaček (1992, pl. 4, fig. 1).

Mai (1986, p. 575; 1994, p. 213) assigned the seed cones

of P. ornata on the basis of the flat apophyses in the basal
cone part and a relatively long stalk to the “Merkusii” group,
together with other related fossil species and their synonyms,
mainly known from Miocene localities of France.

Seed cones of a xerophytic extant species of P. halepensis

Mill. from the Mediterranean and western Asia and those of
the  extant  species  P.  merkusii  Jung.  &  de  Vries  from  Viet-
nam, Laos, Cambodia, China, Philippines, Malaysia and In-
donesia  are  most  comparable  to  P.  ornata.  Kvaček  et  al.
(2004b)  noted  that  P.  massoniana  Lamb.  from  Taiwan  and
China  could  represent  an  extant  equivalent  but  this  species
differs by having shorter cones (only 40—(25)—70 mm long),
which are also more shortly stalked (Fu et al. 1999). Teodo-
ridis  &  Sakala  (2008)  applied  principles  of  the  holistic  ap-
proach (i.e. the whole plant concept sensu Kvaček 2004) to
the  fossil  coniferous  remains  from  the  Most  Basin  (Czech
Republic)  and  defined  the  Pinus  ornata  plant  which  com-
bines seed cones of P. ornata and foliage of P. hepios.

Cupressaceae

Tetraclinis Masters

Tetraclinis salicornioides (Unger) Kvaček

Fig. 2.4

1847 Thuites salicornioides Unger, p. 11, pl. 2, figs. 1—4, pl. 20, fig. 8
1866 Libocedrus salicornioides (Unger) Heer – Ettingshausen, p. 33,

pl. 10, fig. 5 (non fig. 14)

1982 Libocedrites salicornioides (Unger) Endlicher – Kovar, p. 36,

pl. 6, figs. 1, 2, 10, pl. 16, figs. 2—4, pl. 33, figs. 2, 3

1989 Tetraclinis salicornioides (Unger) Kvaček, p. 48, pl. 1, fig. 11,

pl. 2, figs. 2—14, pl. 3, figs. 1—4, text-fig. 1

Material: One complete foliar segment (SNM B 1793/a,b).
Description: Isolated cladode-like branch segment,

15.3 mm long, 1.8 and 5.5 mm broad in its lower and upper
part, corresponding to pseudo-whorl, composed of pairs of di-
morphic facial and lateral leaves with bluntly mucronate api-

ces and rounded base, simple pseudo-whorl obovate, margin
entire, 5 prominent longitudinal lines on upper surface.

Discussion: Kovar (1982, as Libocedrites), Kvaček (1989),

Mai & Walther (1991) and Kvaček et al. (2000) summarized
morphology and taxonomy of Tetraclinis. The relatively ro-
bust  foliage  segment  completely  fused  in  the  pseudo-whorl
shows  morphological  affinity  to  Tetraclinis  salicornioides
rather  than  to  the  more  xeromorphic  T.  brachyodon
(Brongniart) Mai & Walther. Impressions of sterile foliage
of T. salicornioides are almost indistinguishable from an un-
related extinct cupressoid conifer Ditaxocladus S.X. Guo &
Z.H.  Sun  from  the  Upper  Cretaceous  to  Paleocene  of  the
Northern Hemisphere (Guo et al. 2012). The foliage of both
conifers differs in the general shape of foliage sprays, which
are  slender,  elongate  in  Ditaxocladus  and  widely  spread  in
Tetraclinis  salicornioides.  The  seed  cone  morphology  and
leaf anatomy is also significantly different (Guo et al. 2012).

T. salicornioides diverged from T. brachyodon during the

Eocene and adapted to forest conditions in humid subtropical
to  warm  temperate  zones  (Kvaček  et  al.  2000).  Both  species
went  extinct  during  the  Pliocene.  Fragments  of  foliage  and
cones of Tetraclinis also occur in the Cypris Formation (Cheb
and  Sokolov  basins,  Czech  Republic)  and  in  deposits  of  the
Zittau  and  Berzdorf  basins  (e.g.  Bůžek  et  al.  1996;  Czaja
2003; Holý et al. 2012). The only living Tetraclinis articulata
(Vahl)  Masters  is  a  sclerophyllous  relict  occurring  in  the
western  Mediterranean  (native  in  the  Atlas  Mountains  of
Morocco, Algeria and Tunisia, in Malta and Cartagena in SE
Spain)  and  cannot  be  considered  as  the  most  similar  living
relative to T. salicornoides. T. salicornioides may be ecologi-
cally more similar to Calocedrus macrolepis Kurz distributed
in  SE  China,  Myanmar  (Burma),  Thailand  and  Vietnam  on
the basis of its foliage physiognomy (Kvaček et al. 2011).

Angiosperms

Lauraceae

Cinnamomum L. sect. Camphora Nees

Cinnamomum polymorphum (A. Braun) Heer

Fig. 2.5—6

1845 Ceanothus polymorphus A. Braun, p. 171
1847 Ceanothus bilinicus Unger, p. 145, pl. 49, fig. 9
1851 Daphnogene polymorpha (A. Braun) Ettingshausen, p. 16, pl. 2,

figs. 23—25

1855 Camphora polymorpha (A. Braun) Heer, p. 112, pl. 1, fig. 11

Fig. 2. 1 – Pinus hepios (Unger) Heer, incomplete isolated double—needled fascicle, SNM B 1792/a, scale bar 10 mm. 2—3 – Pinus cf. or-
nata (Sternberg) Brongniart, seed cone with its latex mold SNM B 1809, scale bar 10 mm. 4 – Tetraclinis salicornioides (Unger) Kvaček,
isolated cladode-like branch segment, SNM B 1793/a, scale bar 5 mm. 5—6 – Cinnamomum polymorphum (A. Braun) Heer: 5 – basal part
of narrow leaf, SNM B 1795/1, scale bar 5 mm; 6 – almost complete narrow leaf with well-preserved venation, SNM B 1796, scale bar
5 mm. 7—13 – Palaeotriticum mockii Sitár: 7 – incomplete cylindrical infructescence, holotype, SNM B 1786, scale bar 5 mm; 8 – basal
fragment of similar infructescence, SNM B 1787, scale bar 5 mm; 9 – poorly preserved fragment of infructescence with adhering ichnofossils
on the basal part, SNM B 1788, scale bar 5 mm; 10 – detached spikelet, SNM B 1789, scale bar 5 mm; 11 – detail of a spikelet isolated from
the holotype, SNM B 1786, scale bar 1 mm; 12—13 – details of spikelets, holotype, SNM B 1786, scale bar 2 mm. 14 – Palaeotriticum car-
paticum Sitár, spiny cylindrical infructescence, holotype, SNM B 1791, scale bar 5 mm. 15—16 – Palaeotriticum mockii Sitár, Slup locality,
MZM Ge 28923: 15 – cylindrical infructescence, scale bar 10 mm; 16 – detail of spikelets, scale bar 5 mm. 17—18 –Monocotyledonae fam.
et gen. indet.: 17 – linear leaf fragment, SNM B 1795/2, scale bar 10 mm; 18 – accumulated leaf debris, SNM B 1798, scale bar 10 mm.
19 – Potamogeton sp., incomplete leaf, SNM B 1797, scale bar 10 mm. 20—21 – Dicotylophyllum sp. 1.: 20 – fragmentary leaf lamina,
SNM B 1799, scale bar 10 mm; 21 – lower part of an incomplete leaf, SNM B 1802, scale bar 10 mm.

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1856 Cinnamomum polymorphum (A. Braun) Heer, p. 88, pl. 91,

fig. 11c—d, pl. 93, figs. 25—28, pl. 94, figs. 1—16

M a t e r i a l : Three incomplete leaves (SNM B 1794,

B 1795/1, B 1796) and fragments of leaf lamina.

D e s c r i p t i o n : Leaves simple, lanceolate, elliptic to obo-

vate,  22—89 mm  long,  5—34 mm  broad,  base  slightly  asym-
metric,  cuneate  to  broadly  cuneate,  petiole  not  preserved,
apex  incomplete,  acuminate  and  blunt,  margin  entire,  vena-
tion  suprabasal  acrodromous,  midrib  strong,  straight  or
slightly curved in the apical part, lateral veins thinner, alter-
nate,  originating  at  an  angle  of  20—35°,  running  along  the
margin, secondary veins thinner, alternate or opposite, at an
angle of 40—55°, curved and looping near margin or straight
to forked between midrib and lateral veins, venation of higher
orders poorly preserved.

D i s c u s s i o n : Holý et al. (2012) followed the original

view of Heer (1856) refreshed by Ferguson (1971) and asso-
ciated  the  fruits  of  the  Cinnamomum  sect.  Camphora  type
with variable leaves usually identified as Daphnogene poly-
morpha  (e.g.  Mai  1960,  1999;  Holý  1977;  Pingen  et  al.
1994).  The  whole  plant  is  closely  related  to  the  extant  spe-
cies of Cinnamomum camphora L. Remains of Daphnogene/
Cinnamomum are dominant elements in many Oligocene and
Miocene  floras  of  Europe,  including  mastixioid  ones  (e.g.
Bůžek et al. 1996; Kvaček & Walther 1974; Mai & Walther
1978, 1991; Kvaček et al. 2004; Holý et al. 2012). Kvaček &
Walther (1974) revised morphologically variable leaf forms
(i.e.  narrow  lanceolate  to  broadly  oval  forms)  into  a  single
species  of  C.  polymorphum  that  produced  the  above  men-
tioned  type  of  fruits.  The  leaf  variability  was  considered  as
an ecotypical variation (Kvaček & Walther 1974). However,
Kvaček & Walther (1974) treated leaves of Daphnogene cin-
namomifolia  (Brongniart) Unger from the Paleogene locali-
ties  separately  from  Cinnamomum  polymorphum,  which  is
preserved in branches with attached fruits and differs in de-
tails  of  epidermal  anatomy.  Knobloch  (1967,  1969)  de-
scribed  abundant  leaves  of  Daphnogene  bilinica  (Unger)
Kvaček & Knobloch and one broader incomplete leaf of D.
cinnamomeum  (Rossmässler)  Knobloch  from  the  Karpatian
flora  of  Dolní  Dunajovice,  which  morphologically  corre-
sponds to those from the Cerová-Liesková site.

Platanaceae

Platanus L.

Platanus neptuni (Ettingshausen) Bůžek, Holý & Kvaček

Fig. 3.1—3, 3.5—7

1967 Platanus neptuni (Ettingshausen) Bůžek, Holý & Kvaček, p. 205,

pl. 1. figs. 1—4, 6 (non 5 – Sloanea artocarpites), pls. 2—4 (for a
full synonymy see Kvaček & Manchester 2004)

M a t e r i a l : Leaf impressions (SNM B 1801-1802, 1804),

fruitlets impressions partly with counterparts (SNM B 1806-8).

D e s c r i p t i o n : Leaves simple ovate to slightly obovate,

up to 80 mm long and max. 25 mm broad, margin sub-entire
or  fine  widely  dentate  on  the  upper  part,  midrib  strong,  al-
most  straight,  venation  eucamptodromous  to  semicraspedo-
dromous,  secondary  veins  regularly  spaced,  at  an  angle  of

45—60°, looping near the margin, quite thin, interspaced
partly with single intersecondaries, higher-order veins indis-
tinct. Sessile narrow ovoid to obovoid fruitlets, 6 mm long
and up to 3 mm broad, ending into 0.5 mm thick and 3 mm
long blunt slender and bent style, the body showing a few in-
distinct longitudinal ribs.

D i s c u s s i o n : Foliage of this extinct plane tree is pre-

served as impressions of almost entire-margined leaves and
disintegrated remains of infructescences. This deciduous ele-
ment occurred in Europe in the Paleogene and continued up
to the middle Miocene (Kvaček & Manchester 2004).

Monocotyledonae

Palaeotriticum Sitár, gen. nov.

Palaeotriticum Sitár (2001), pp. 115—116, nom. inval.

(typification lacking)

The genus name Palaeotriticum was proposed by Sitár

(2001, p. 115) for: “imprints of grass ears”.....“related to
modern Triticum”. The diagnosis of this taxon needs to be
validated as follows:

D i a g n o s i s : Infructescences spike-like, cylindrical, con-

sisting of serially arranged ovoid, tightly crowded bodies
corresponding to spikelets with glume-like bracts.

T y p e : Palaeotriticum mockii Sitár.
The material consists of several specimens of cylindrical

spike-like infructescences preserved as compressed moulds.
Two fossil species are recognized: Palaeotriticum mockii Sitár
(form A) and Palaeotriticum carpaticum Sitár (form B). After
a more detailed study of the material, the following character-
istics are offered to complement the original descriptions.

Palaeotriticum mockii Sitár

Fig. 2.7—13, 2.15—16

1850 Pinites spiciformis Unger, p. 529, pro parte
1852 Pinites spiciformis Unger, p. 28, pro parte, pl. 14, fig. 14 (non

fig. 15)

1967 Spirematospermum cf. wetzleri (Heer) Chandler – Knobloch,

p. 253, pl. 1E, fig. 4

1969 Spirematospermum cf. wetzleri (Heer) Chandler – Knobloch,

p. 48, pl. 13, fig. 3

2001 Palaeotriticum mockii Sitár, pp. 115—116, fig. 2 (generotypus)

D i a g n o s i s : Sessile cylindrical infructescence with

rounded base containing slender ovoid bi-serially arranged
bodies interpreted as single-seeded spikelets with tri-veined
unarmed glumes recalling bracts stretching over the seed by
a short sharp tip.

H o l o t y p e : The specimen illustrated in Sitár (2001,

fig. 2) and re-illustrated here in Fig. 2.6, housed in the Slo-
vak National Museum in Bratislava under inventory number
SNM B 1786.

A d d i t i o n a l m a t e r i a l : 2 fragments of infructescences

(SNM B 1787, B 1788) and 2 isolated spikelets (SNM B 1789,
B 1790).

D e s c r i p t i o n : The holotype is the most complete speci-

men attaining the maximum width near the rounded base of
12 mm and very slightly narrowing upwards. It is preserved

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in the length of ca. 60 mm, which may represent half of the
complete  infructescence.  It  shows  an  arrangement  of  spike-
lets coming obliquely from the center in three visible rows.
The median row apparently covers the spindle. The spikelets
are  obviously  one-seeded,  devoid  of  surface  tissue.  Incom-
plete  remnants  of  glumes  are  seen  on  some  bodies  and  ap-
pear  ventrally  to  be  tri-veined  (Fig. 2.11).  The  tightly
crowded  spikelets  are  exposed  and  show  smooth  surface  of
the seed, which is not preserved. Spikelets of the lateral rows
are sub-parallel, attached at an angle of ca. 35°. Spikelets of
the  medial  row  are  compressed  longitudinally.  The  spikelet
separated  mechanically  from  one  lateral  row  (Fig. 2.11)  is
obovoid, 8 mm long and max. 3 mm broad, bluntly pointed
at the apex, wedge-shaped and flattened on the basis. On the
ventral side, it shows the remnant of the glume with 3 darker
lines of veins that disappeared during the treatment in diluted
hydrofluoric  acid.  The  specimen  was  damaged  during  the
preparation. The loose spikelet (Fig. 2.10) is spindle-shaped,
12 mm long and 3 mm broad, smooth on the surface, without
apparent details except for slight striation on the lower part
coming from a stout short stalk on the base. Another incom-
plete specimen (Fig. 2.8) represents a basal fragment of the
same spike-like infructescence. It is 10 mm wide, and rounded
at the base, without any stalk, showing less distinctly ovoid
tightly  crowded  bodies  interpreted  as  single-seeded  spike-
lets. The last specimen (Fig. 2.9), showing probably the mid-
dle part of a spike-like infructescence, includes only 2 rows
of wedge-shaped spikelets with smooth surface.

D i s c u s s i o n : Unger (1850, 1852) described two infructes-

cences as  Pinites spiciformis from the Slovenian Tertiary lo-
cality  of  Saalberg  at  Stein,  called  now  Žale  at  Kamnik  (late
Badenian, Tunjice Hills). One of these specimens (Unger 1852,
pl. 14,  fig. 14)  strongly  resembles  our  Palaeotriticum  mockii
and differs only by slightly larger dimensions of its spikelets
(max. 10 mm long and 6 mm wide, if the illustration reflects
the  object  in  natural  size).  Unger  believed  that  both  figured
specimens may represent slender conifer seed cones similar to
Pinus strobus L. We were unable to inspect the type material
from Slovenia, which is missing in the collections of Joanneum
(Graz, Austria), where most of the Unger’s original material is
housed,  and  it  is  also  not  present  in  the  collections  of  the
Geologische  Bundesanstalt,  Vienna.  From  the  same  locality,
Unger (1860, p. 41) listed fruits of Carya costata (Sternberg)
Brongniart. The Žale site in Slovenia seems to be of the late
Badenian  age  on  the  basis  of  the  geological  surroundings
(Bogomir Jelen, personal communication Nov. 2013).

One of the type specimens of Pinus lardyana Heer (1855,

p. 38,  pl. 20,  fig. 5e)  from  the  Lower  Miocene  deposits  of
Lausanne  also  resembles  Palaeotriticum  mockii  but  the  in-
spection of the actual specimen is required to achieve an accu-
rate comparison. One of the illustrated specimens differs from
our  material  in  spikelets  parallel  to  the  main  axis.  The  other
type material from localities near Lausanne differs still more
by  crowded  irregularly  disposed  bodies  resembling  Spire-
matospermum (see below). One of the syntypes shows a rha-
chis or a twig fully devoid of bracts or foliage and thus may
partly  belong  to  Pinus.  The  species  requires  a  more  detailed
revision. Mai (1986, p. 577) assigns Pinus lardyana to invalid
or  insufficiently  described  species.  Knobloch  (1967,  1969)

described  an  incomplete  infructescence  from  the  Karpatian
locality Slup SE of Znojmo in southern Moravia (Carpathian
Foredeep), fully corresponding to Palaeotriticum mockii. He
assigned it with doubts to Spirematospermum and described it
as a pod 85 mm long and 13 mm broad with serially arranged
moulds of oval seeds (Knobloch 1967– p. 253, pl. 1E, fig. 6;
1969 – p. 48, pl. 13, fig. 5). The bodies, which we interpret
as  single-seeded  spikelets,  are  arranged  in  one  medial  and
two (or double) lateral rows. The overall shape of the fossil
is cylindrical, slightly narrowing to the apex and rounded at
the  base.  The  surface  is  abraded,  not  showing  glume-like
processes  but  otherwise  matching  Palaeotriticum  mockii
(Fig. 2.15—16).  This  material  is  housed  in  the  Moravské
Zemské Museum Brno (MZM, Ge 28923).

The affinity of Palaeotriticum mockii is uncertain due to

poor preservation. Spiral surface sculpture typical of seeds of
Spirematospermum  is  not  visible.  One  of  the  isolated  bodies
from the holotype does not show any sculpture under scanning
electron  microscope  (SEM).  The  incomplete  infructescenses
widely deviate in form from capsules of  Spirematospermum,
in which the crowded seeds are arranged more chaotically and
the capsules are clearly stalked (Fischer et al. 2009, pl. 3). The
bodies in the holotype look like fillings of empty spaces.

We disagree with Unger’s interpretation because the seed

cones of conifers are woody but the plant remains from both
localities in Slovakia and Moravia are devoid of any woody
tissue or its possible traces. The arrangement of fruitlets also
strongly  differs  from  the  structure  of  pine  cones,  in  which
the  seeds  are  more  loosely  arranged,  in  the  case  of  Pinus
strobus  L.  with  adnate  wings.  The  fossils  assigned  to
Palaeotriticum mockii resemble earls of grasses, namely the
wheat with unarmed fruitlets. However diagnostic characters
of  caryopsides  such  as  scales  (glumes,  lemmas)  enclosing
the fruitlet and seed are absent. The bodies are filled with the
sediment, and thus were apparently hollow when buried, and
their surfaces bear darker thin remnants of plant tissue.

The fruit remains of grasses described from the Miocene

until now (e.g. Lańcucka-Środoniowa 1966, 1979) are much
smaller and show inner structural traits of embryo. Accord-
ing to the morphological similarity of Palaeotriticum and its
relatively  good  preservation  implying  a  rather  short-term
transport  from  the  coast,  the  most  likely  living  analogue  of
Palaeotriticum  mockii  appears  to  be  Spartina  alterniflora
Loisel., an invasive halophytic grass spread on salt marshes
of the Americas from Canada to Argentina, which produces
similar spikes. Another species with similar infructescences
is Spartina cynosuroides (L.) Roth, called Big Cordgrass or
Giant  Cordgrass,  which  forms  dense  stands  in  brackish
marshes on tidal wetlands of the Atlantic USA from the Gulf
Coast to New England (see Hitchcock 1951).

Palaeotriticum carpaticum Sitár

Fig. 2.14

2001 Palaeotriticum carpaticum Sitár, p. 118, fig. 3

D i a g n o s i s : Cylindrical infructescence well flattened

covered by impressions of bract-like processes and showing
long spines arising from the spikelet surface.

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H o l o t y p e : The specimen illustrated in Sitár (2001,

fig. 3) and re-illustrated here in Fig. 2.14, housed in the Slo-
vak National Museum in Bratislava under inventory number
SNM B 1791.

D e s c r i p t i o n : Only one specimen is available. Contrary

to the infructescences assigned to P. mockii, its surface is flat
compressed,  not  showing  any  distinct  spikelets-like  bodies.
It  comes  from  the  same  strata  as  the  previous  taxon.  The
compression  is  almost  complete,  63 mm  long  and  max.
10 mm  broad,  with  a  blunt  apex  without  its  very  terminal
part, and the slightly damaged base; this base is widely coni-
cal, without any sign of a stalk. On the sides of the infructes-
cence,  sharp  ca.  2—3 mm  long  spine-like  processes  stretch
apically recalling awns in some grasses.

D i s c u s s i o n : We are not aware of any similar fossil re-

mains matching Palaeotriticum carpaticum Sitár. Our speci-
men may represent an empty infructescence of the previous
species. It does not show any diagnostic trait that would al-
low a better comparison.

Potamogeton sp.

Fig. 2.19

M a t e r i a l : Incomplete leaf (SNM B 1797).
D e s c r i p t i o n : Leaf simple, elliptic, 34 mm long and

13 mm broad, base almost complete, cuneate, narrowed con-
tinually into 7.8 mm long and 2 mm broad petiole, apex in-
complete, probably acute or obtuse, margin entire, venation
poorly preserved, ?paralellodromous, with 3 thin parallel
veins bent towards the apex on either side of a thin midrib.

D i s c u s s i o n : The entire margin, the narrowed base with

a broad petiole and paralellodromous venation type are mor-
phological  features  that  may  correspond  to  leaves  of  Pota-
mogeton.  Broader  leaf  forms  of  this  genus  were  rarely
described,  e.g.  by  Knobloch  (1969)  from  the  Karpatian    of
Moravia at Nový Jičín as Potamogeton sp. or by Sitár (1969)
from the middle to upper Miocene deposits of the Turiec Ba-
sin as P. martinianus Sitár.

Monocotyledonae fam. et gen. indet.

Fig. 2.17—18

M a t e r i a l : Numerous leaf fragments (SNM B 1795/2,

B 1798).

D e s c r i p t i o n : Leaf fragments simple linear, various in

size up to 170 mm long and 29 mm broad, base and apex not
preserved, margin entire, venation paralellodromous and
poorly preserved.

D i s c u s s i o n : Sitár (2001, Fig. 1; Fig. 2.17) described

several  incomplete  leaf  impressions  (5—7 mm  broad)  with
parallel venation, which co-occurred with Paleotriticum and
were  preliminary  assigned  to  the  Poaceae  family.  Similar
leaf  fragments  are  common  in  the  Paleogene  and  Neogene
plant  assemblages  of  Europe  and  are  usually  interpreted  as
azonal  herbaceous  elements  because  they  are  accompanied
by fruits of Cyperaceae, typical of moist habitats. Our speci-
men  (Fig. 2.17)  is  poorly  preserved  and  more  slender.  It
could have some affinity to Pinus, but no morphological fea-
ture resembling leaf midrib is visible. Bands of leaf remains
accompanying  the  infructescences  of  Palaeotriticum  at
Cerová-Lieskové as well as at Slup are not referable to any
particular monocot.

Angiospermae incertae sedis

Dicotylophyllum Saporta

Dicotylophyllum sp. 1

Fig. 2.20—21

M aterial : Incomplete leaves and fragments (SNM B 1799,

B 1802).

Description: Leaves simple, elliptic to obovate, 28—72 mm

long, 11—24 mm broad, base cuneate, petiole not preserved,
apex incomplete, probably shortly acuminate and blunt, mar-
gin entire, venation brochidodromous, midrib strong,
straight or slightly curved in the apical part, secondary veins
thinner, alternate, originating at an angle of 40—60°, curved
and looping near margin, intersecondaries parallel to major
secondaries, venation of higher orders poorly preserved.

Discussion: These remains show some affinity to Lower

Miocene  leaf  morphotypes  of  the  Lauraceae,  Magnoliaceae
and Fabaceae families. The leaves may not represent a single
fossil  species.  The  impression  character  and  poor  preserva-
tion  do  not  allow  discrimination  of  the  leaf  texture.  The
specimen  shown  in  Fig. 2.20  is  similar  to  the  leaf  material
from several Miocene localities in the Rhineland and Lower
Lusatia described as  Papilionaceophyllum liblarense Kräusel
&  Weyland  ( = Magnolia  liblarensis  (Kräusel  &  Weyland)

Fig. 3. 1—3 Platanus neptuni (Ettingshausen) Bůžek, Holý & Kvaček: 1 – negative impression of a complete leaf with fine serrate apex, SNM
B 1800, scale bar 10 mm; 2 – positive incomplete leaf impression with distinctly serrate leaf margin and semicraspedodromous venation,
SNM B 1801, scale bar 10 mm; 3 – incomplete leaf with poorly preserved venation, SNM B 1804, scale bar 5 mm. 4 – Dicotylophyllum
sp. 2, very narrow fragmentary leaf lamina, SNM B 1803/a, scale bar 10 mm. 5—7 – Platanus neptuni (Ettingshausen) Bůžek, Holý & Kvaček:
5 – immature fruitlet with long slightly curved style, SNM B 1806/b, scale bar 1 mm; 6 – immature fruitlet with almost straight style, SNM
B 1807, scale bar 1 mm; 7 – impression of an immature fruitlet with long and curved style showing a few indistinct longitudinal ribs, SNM
B 1808, scale bar 1 mm. 8 – Concavisporites sp. (Gleicheniaceae), spore, interval 49—50. 9—10 – Cupressaceae, pollen grains, sample 12.
11—13 – Pinaceae, pollen grains, interval 49—50. 14—15 – Pinus sp., pollen grain, interval 49—50 and sample 18. 16 – cf. Potamogeton, pol-
len grain, interval 49—50. 17 – cf. Liquidambar, pollen grain, sample 46. 18 – Fagaceae, pollen grain, sample 23. 19 – Engelhardia sp.,
pollen grain, interval 49—50. 20 – Craigia sp. (Intratriporopolenites cf. insculptus type), pollen grain, interval 49—50. 21 – Tri-
colporopollenites sp., pollen grain, interval 49—50. 22 – Zygnemataceae, Algae, sample 46. 23 – Algae, sample 46. 24 – Mecsekia sp.,
Algae, interval 49—50. 25 – Dinoflagellata, Algae, sample 18. 26—27 Dinoflagellata, Algae, interval 49—50. 28 – Dinoflagellata, Algae,
sample 46. 29 – Dinoflagellata, Algae, sample 23. 8—29 – scale bar 20 µm.

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Kvaček)  and  related  to  the  genus  Magnolia  L.  (Kvaček
1979; Schneider 2007). In the absence of the epidermal anat-
omy, the affinity to this species or to other numerous species
of  Laurophyllum/Lauracaeae  and  Fabaceae  is  inconclusive
(e.g.  Bůžek  et  al.  1996;  Kvaček  1971;  Holý  et  al.  2012).  A
stratigraphically  and  geographically  close  plant  assemblage
from  Dolní  Dunajovice  contains  similar  leaves  with  entire
margins  assigned  to Laurophyllum  sp.  and Dicotylophyllum
sp. 8 by Knobloch (1969).

Dicotylophyllum sp. 2

Fig. 3.4

M a t e r i a l : Fragment of leaf/leaflet lamina (SNM B

1803/a,b).

D e s c r i p t i o n : Medial part of leaf/leaflet lamina, 53 mm

long, 4 mm broad, base and apex not preserved, margin
probably entire, venation probably brochidodromous, midrib
strong, slightly curved, lateral veins thinner, alternate, origi-
nating at an angle of 40—55°, venation of higher orders poorly
preserved.

D i s c u s s i o n : The incomplete character of the lamina

does not allow any assignment to the genus and family levels.
Similar leaf impressions from the Karpatian of Moravia
(Dolní Dunajovice) were assigned to Echitonium sophiae
O. Weber by Knobloch (1969).

Palynology

Terrestrial plants sporomorphs (Fig. 3.8—21) of Pinaceae,

Cupressaceae,  Pinus  sp.,  Engelhardia  sp.,  Tricolporopolle-
nites sp., cf. Potamogeton, cf. Craigia sp. (Intratriporopoleni-
tes  cf.  insculptus)  and  aff.  Concavisporites  (Gleicheniaceae)
occur  in  interval  49—50,  in  addition  to  palynomorphs  of
Mecsekia  sp.  (Fig. 3.24)  and  dinoflagellates  (Fig. 3.25—29)
typical  of  marine  environment.  Similarly,  the  samples  46
and 23 contain various dinoflagellates and marine algae rem-
nants (46) as well as Mecsekia sp. (23) and freshwater algae
(Zygnemataceae) (Fig. 3.22), which are associated with very
rare  and  poorly  preserved  cf. Liquidambar  and  unidentified
pollen  Tricolporopollenites  sp.  (46)  and  single  grains  of
Fagaceae,  Cupressaceae  and  Pinaceae  (23).  Sporomophs
occur rarely in sample 12, being represented by single grains
of  Cupressaceae  and  cf.  Potamogeton  sp.  The  sample 18
shows a palynological composition almost identical to that of
sample 23  and  is  characterized  by  Mecsekia  sp.,  dinoflagel-
lates, Pinaceae, and by re-worked Lygodium spore. The sam-
ples  49  and  30  are  totally  barren  of  palynomorphs.  Single
and  poorly  preserved  grains  of  Pinaceae  and  Cathaya  sp.
were found in samples 20 and 33.

Composition and habitat of source plant assemblage

The macrofossil plant assemblage of Cerová-Lieskové con-

sists of two conifers and four morphotypes of angiosperm
foliage as well as of several molds of spike-like infructes-
cences of azonal monocots. The palynological assemblage is

also not diverse and taxonomically matches the macrofossil
plant assemblage. The low taxonomic richness and common
co-occurrence of these taxa during the Miocene do not allow
precise  phytostratigraphical  correlation  with  other  plant  as-
semblages  in  Central  Europe.  However,  the  occurrence  of
the infructescence of Palaeotriticum mockii at the locality of
Slup  near  Znojmo  implies  floristic  similarity  between  the
Vienna Basin and the Carpathian Foredeep. The single speci-
men at this locality co-occurs with Tetraclinis salicornioides
(cited  by  Knobloch  1969  as  Libocedrites  salicornioides),
Cinnamomum  polymorphum  (as  Daphnogene  bilinica)  and
with  several  lauroid  and  betuloid  leaf  fragments.  Knobloch
(1967,  1969)  described  Karpatian  plant  assemblages  from
Dolní Dunajovice, Velké Dyjákovice and from the drill-core
of  Kl 147  at  Žilina  near  Nový  Jičín  from  the  Czech  part  of
the Carpathian Foredeep. Two other conifers – Glyptostro-
bus  europaeus  (Brongniart)  Unger  and      Pinus  sp.  (cited  in
Knobloch  1969,  p. 45,  as  P.  aff.  goethanus  Unger,  P.  aff.
saturni  Unger)  and  several  angiosperms  (Potamogeton  sp.,
Arundo  sp.  vel  Phragmites  sp.,  Liquidambar  europaea  A.
Braun, Carpinus sp., Betulaceae gen. et sp. indet., Echitonium
sophiae  Weber,  and  Dicotylophyllum  sp.  div.)  also  occur  in
the Carpathian Foredeep. These plant assemblages consist of
taxa that occur frequently during the   Miocene at other sites in
the Czech Republic (the Cypris Formation, the Kristina Mine
in the Zittau Basin in North Bohemia, the Mydlovary Forma-
tion  in  South  Bohemia  –  Bůžek  et  al.  1996;  Holý  et  al.
2012),  Germany  (Berzdorf,  Wackersdorf  –  Knobloch  &
Kvaček  1976;  Czaja  2003),  Poland  (Turów  –  Czeczott  &
Skirgiełło 1959, 1961, 1967, 1975, 1980) and Austria (Kovar-
Eder 1998; Meller 1998; Hofmann et al. 2002).

We suggest that the source vegetation of Cerová-Lieskové

represents  a  mixture  of  azonal  and  zonal  elements.  Arboreal
elements  of  Tetraclinis  salicornoides/Cupressaceae,  Ca-
thaya,  Pinus  sp.,  Cinnamomum  polymorphum,  Engelhar-
dia,  Platanus  neptuni,  and  probably  lauroid  elements  of
Dicotylophyllum sp. 1—2 belong to mesophytic vegetation of
the  Broad-leaved  Evergreen  or  Mixed  Mesophytic  forest
type  (e.g.  Kovar-Eder  et  al.  2008).  These  vegetation  types
are  also  captured  by  other  Karpatian  plant  assemblages  of
Central  Europe  (Kovar-Eder  et  al.  2001;  Holý  et  al.  2012).
Azonal  elements  are  represented  by  herbaceous  monocots,
including Paleotriticum, Potamogeton sp., and arboreal ele-
ments  such  as  Liquidambar  and  partly  mesophytic  Craigia
(Kvaček & Walther 2004) corresponding to riparian vegeta-
tion.  The  analogy  with  the  Spartina  type  grasses  allows  re-
construction of the herbaceous vegetation type as coastal salt
marshes and brackish marshes as known on tidal wetlands of
North America.

Conclusions

Fully marine, upper slope bathyal clays and silts at

Cerová-Lieskové locality contain relatively well-preserved
plant remains that reflect terrestrial vegetation of the sur-
rounding land. The plant assemblage corresponds to alloch-
thonous remains transported by currents into the upper-slope
environments. The macrofossil and pollen assemblages corre-

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spond to thermophile subtropical forests typical of the Karpa-
tian (late Burdigalian) time interval in Europe (Brzobohatý et
al.  2003).  Enigmatic  monocots  resembling  wheat  infructes-
cence  probably  belong  to  herbaceous  salt  marsh  vegetation
growing  close  to  the  shoreline,  similar  to  the  living  grass
Spartina  Schreb.  These  monocots  are  similar  to  specimens
described  from  Moravia  (Early  Miocene—Karpatian)  and
Slovenia  (Middle  Miocene—Badenian).  These  specimens
were  previously  wrongly  assigned  to  Spirematospermum
fruits or Pinaceae seed cones, respectively. We suggest that
these monocots represent herbaceous vegetation bounded to
intra-tidal flats.

Acknowledgments:  We  thank  for  information  on  geology,
paleobotany,  access  to  collections  and  technical  assistance
Edoardo Martinetto, Barbara Meller, Bogomir Jelen, Mathias
Harzhauser, Růžena Gregorová, Nela Doláková, and Libuše
Plchová. We greatly appreciate the comments by Steven R.
Manchester,  Johanna  Kovar-Eder,  and  Adam  Tomašových.
The  study  was  supported  by  the  grant  Projects  of  GA ČR
(Grant Agency of the Czech Republic) 14-23108S, the Min-
istry  of  Education,  Youth  and  Sports  (scheme  MSM
002162085  and  PRVOUK  P  44)  and  research  Projects  of
SRDA  APVV-0644-10  and  APVV-0099-11.  This  is  also  a
contribution to the NECLIME Project (www.neclime.de).

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