GEOLOGICA CARPATHICA, FEBRUARY 2009, 60, 1, 43—57 doi: 10.2478/v10096-009-0005-4
The Hungarian record of Eocene floras is limited which may
be due first of all to the rare occurrence of sediments pre-
serving macroscopic remains of Eocene plants. Accordingly,
reports on Eocene plants and floras from Hungary have ex-
clusively been recorded from the Transdanubian region of
The joint bauxite and brown coal exploration was started
in the Csordakút opencast mine (Csordakút Basin, Gerecse
The Middle Eocene flora of Csordakút (N Hungary)
and LÁSZLÓ RÁKOSI
Hungarian Natural History Museum, Botanical Department, P.O.Box 222, H-1476 Budapest, Hungary; firstname.lastname@example.org
Templomkert str. 14, 2094 Nagykovácsi, Hungary
(Manuscript received September 17, 2007; accepted in revised form June 12, 2008)
Abstract: The Middle Eocene fossil plant assemblage from Csordakút (N Hungary) comprises plant remains preserved
exclusively as impressions. Algae are represented by abundant remains of Characeae, including both vegetative frag-
ments and gyrogonites. Remains of angiosperms comprise Lauraceae (Daphnogene sp.), Fagaceae (cf. Eotrigonobalanus
furcinervis), Ulmaceae (Cedrelospermum div. sp.), Myricaceae (Myrica sp., Comptonia div. sp.), Leguminosae (leaves
and fruit), Rhamnaceae (?Zizyphus zizyphoides), Elaeocarpaceae (Sloanea nimrodi, Sloanea sp. fruit), Smilacaceae
(Smilax div. sp.). The absence of gymnosperms is indicative of a floristic similarity to the coeval floras of Tatabánya (N
Hungary) and Girbou in Romania. Sloanea nimrodi (Ettingshausen) Kvaček & Hably, a new element for the Hungarian
fossil record indicates a floristic relation to the Late Eocene flora of Kučlin (Bohemia).
Key words: Eocene, Lutetian, paleobotany, plant macrofossils, impressions.
Mountains, Fig. 1) in 1982 (Végh-Neubrandt et al. 1985).
Connected to the exploration of the opencast mine collecting
activities started at that time and have been carried out by the
second author continuously for more than 15 years. This is
the first time that the fossil plant assemblage of Csordakút
has been described and published.
Though the number of fossils collected through the years
is high, indeterminable specimens are numerous since plant
remains are poorly preserved lacking organic matter suitable
for cuticular studies.
Fig. 1. Location of Csordakút.
ERDEI and RÁKOSI
The total thickness of Middle and Upper Eocene sedi-
ments in Hungary does not exceed 500—700 m. Eocene rocks
always discordantly overlie older (Mesozoic or Paleozoic)
strata (Kecskeméti 1998).
The Csordakút Basin and the westward adjacent Nagyegy-
ház Basin have a similar sequence of strata (Csordakút 1
borehole, Gerecse Mountains, Fig. 2) since the two basins
formed one tectonic unit during the coal formation and their
separation started after the Oligocene due to the Savian oro-
genic cycle (Sólyom 1972).
The basement is formed by Triassic dolomites which are
overlain by bauxite of varying thickness and extent. They are
usually intermingled or even replaced by clays of various co-
lours and sandy clays. Two browncoal seams, an upper and a
lower one, are separated by freshwater sediments, so-called
“interstones”. In the Csordakút area up to even 5 seams are
sometimes developed. At the same time a decrease of coal
quality is experienced with the occurrence of clayey coals or
even coaly clays in the seams. The “interstones” comprise
freshwater limestones and carbonaceous marls. In the upper
section they comprise grey, bluish-grey clay layers and clay
marls separating the lower and upper coal seams. The upper
coal seam is often dissected by freshwater and brackish
clays, sandy clays and freshwater limestones into several
(even 4—6) sections. The lower seam is definitely of limnic
origin whereas, the upper seam is paralic. The upper seam is
overlain by freshwater sandy clays, and brackish molluscan
clays, clay marls with a characteristic macrofauna. The next
layers, which were formed in an environment becoming
gradually open marine, are marls with foraminifers and mol-
luscs (Sólyom 1972). The occurrence of Nummulites sub-
planatus Hantken and N. variolarius (Lamarck) Hantken
indicates the first marine sediments of the Eocene transgres-
sion (Kecskeméti in Hably 1985a) and thus, the fossiliferous
sediments are assignable to the Lutetian (Middle Eocene).
(Furthermore, the sediments overlying the upper seam are re-
garded as the heteropic facies of the “operculine marl” based
on its microfossil content.) The Eocene sediments are uncon-
formably overlain by Oligocene sediments, such as variegat-
ed clays, clays, sandstones, conglomerates, etc. (Sólyom
1972). Plant fossils were collected from the freshwater lime-
stone and clayey limestone layers of the so called “inter-
stone” between the lower and upper coal seams.
Material and method
The collection at our disposal is stored in the Hungarian
Natural History Museum, Budapest (Hungary). It comprises
mainly leaves, and rarely fruits.
The fossils are preserved in limestones or clayey lime-
stones, exclusively as impressions. The leaves are often
poorly preserved, entire specimens with apex, base and leaf
margin are relatively rare. The leaf margin is frequently de-
stroyed which may suggest transport and/or decay of the
plant material before final burial. However, it must be added
that even fossiliferous matrix is relatively coarse-grained not
favouring the preservation of finer details. Digital photos
were processed with the software ‘Adobe Photoshop’.
Abbreviations for collections used in the text: BP – pale-
obotanical collection of the Hungarian Natural History Mu-
M a t e r i a l : BP.2001.781.1; BP.2001.792.1.
D e s c r i p t i o n : 2—5 cm long fragments of algae. Leaf-
like branchlets ( ~ 10) of equal length grow in whorls
around the stem. Branchlets undivided, clustered at regu-
larly spaced (3—4 mm) joints. Thorn-like projections on the
branchlets not observable.
D e t e r m i n a t i o n a n d d i s c u s s i o n : The structure of
the fragment and some gyrogonites embedded in the fossilif-
erous matrix support the occurrence of Characeae. Species of
Nitella (Characeae) display a similar structure but with
forked branchlets (Wood & Imahori 1964, 1965), as well as
Fig. 2. Geological profile of the Csordakút 1 borehole. 1 – Triassic
dolomites; 2 – bauxite; 3 – clay; 4 – freshwater limestone; 5 –
coal, clayey coal; 6 – limestone; 7 – marl; 8 – sand, sandstone.
THE MIDDLE EOCENE FLORA OF CSORDAKÚT (N HUNGARY)
Ceratophyllum (Ceratophyllaceae) which bears also forked
leaves. Species of Chara are widely distributed in habitats
ranging from fresh to brackish water, inland to coastal, shal-
low to deep water environments.
M a t e r i a l : BP.2001.687.1; BP.2001.689.1; BP.2001.693.1.
D e s c r i p t i o n : A leafy twig (9.5 cm) with 5 attached and
some isolated leaves. Simple leaves alternate, attached at
distances of 1—1.5 cm. Leaves petiolate, petiole stout, 0.7—
1 cm long. Lamina lanceolate, 5.5—6 cm long, 1.2—1.6 cm
wide. Apex long acute, or attenuate, base acute, often cu-
neate. Margin entire. Venation acrodromous (?suprabasal).
Midvein moderately thick and curved. A pair of secondaries
nearly as thick as the midvein diverges from the midvein
close to the base and run up into the upper third of the leaf.
Higher order venation not preserved.
D e t e r m i n a t i o n a n d d i s c u s s i o n : The morphology of
the leaves is comparable to the lanceolate leaf forms of Daph-
nogene cinnamomifolia (Brongniart) Unger regarded as the sun
leaves of the species (Kvaček & Walther 1974, 1998). In the
flora of Csordakút broader leaves (suspected shade leaves) were
not found, however, the total number of leaves recalling the
morphology of Daphnogene is definitely low. Thus, far reach-
ing conclusions should not be drawn. In the lack of cuticular
structures we refrain from assigning the fossils to a species.
Fossil leaves of Daphnogene have frequently been recorded
in the Paleogene floras of Europe, e.g. Late Eocene of Staré
Sedlo (Knobloch et al. 1996) – Daphnogene cinnamomea
(Rossmässler) Knobloch, Daphnogene pseudopolymorpha
Knobloch et Kvaček – leaves with generally broad and
rounded lamina; the Middle Eocene flora of Messel (Wilde
1989) Daphnogene leaves with usually broader lamina, etc.
The stratigraphic occurrence of Daphnogene leaves in the
Paleogene-Neogene deposits of Hungary ranges up to the
Late Miocene. Fossil leaves of Daphnogene are typical ele-
ments of the Hungarian Early and Late Oligocene, e.g.
Nagybátony-Újlak, Eger-Kiseged, Eger-Wind, Csolnok, etc.
(Kvaček & Hably 1991; Manchester & Hably 1997; Hably &
Manchester 2000; Erdei & Wilde 2004) and Early Miocene
(Ipolytarnóc, Hably 1985b) floras.
Lauraceae gen. et sp. indet.
M a t e r i a l : BP.2001.683.1; BP.2001.722.1; BP.2001.807.1;
BP.2001.828.1; BP.2001.862.1; ?BP.2001.731.1.
D e s c r i p t i o n : Fragmentary leaves, length (fragmentary)
more than 4 cm, width 1—1.5 cm. Apex long acute (when pre-
served), base not preserved. Margin entire. Venation campto-
dromous, with secondaries arising at angles of 40—60°.
Secondaries curved when running toward the margin and
D e t e r m i n a t i o n a n d d i s c u s s i o n : Leaf morphology,
entire margin and type of venation suggest leaves of Lauraceae.
cf. Eotrigonobalanus furcinervis
(Rossmässler) Walther et Kvaček
M a t e r i a l : BP.2007.132.1; BP.2007.137.1; BP.2001.678.1;
BP.2001.788.1;BP.2001.795.1; BP.2001.882.1; ?BP.2001.786.1.
D e s c r i p t i o n : Simple mostly fragmentary leaves. Petiole
not preserved. Leaves lanceolate, length even 13 cm, width
1.4—3.4 cm. Apex attenuate, base cuneate (preserved in one
specimen). Margin toothed, teeth small, acute. Venation
craspedodromous. Midrib straight and moderately thick, sec-
ondaries curved upwards. Tertiary veins not observable due
to poor preservation.
D e t e r m i n a t i o n a n d d i s c u s s i o n : The fragmentary
and poorly preserved leaves recall in morphology Eotrigo-
nobalanus. The leaves of Eotrigonobalanus furcinervis display
a wide morphological range (Kvaček & Walther 1989), namely
lanceolate to ovate forms, entire to toothed margin, camptodro-
mous to semicraspedodromous and craspedodromous venation.
Fossils of the species are widespread in the Paleogene of
Europe (for details see Kvaček & Walther 1989; Knobloch et
al. 1996). A further record of the species may be added from
the Eocene flora of Ovce Polje, Macedonia (Mihajlovič &
Ljubotenski 1994). In Hungary the species was widespread
in the Early Oligocene floras of the Tard Clay Formation
(Hably 1986; Hably & Manchester 2000), as at Eger-Ki-
seged, localities in Budapest – borehole Kiscell-1, Vörös-
vári street, Nagybátony-Újlak brickyard.
The species was an important element of the forest vegeta-
tion (both in intrazonal and zonal types) in the European Pa-
leogene, from the Middle Eocene up to the Early Miocene
(Mai & Walther 1985; Kvaček & Walther 1989). Its wide-
spread occurrence attests to a wide ecological tolerance but
this element was certainly not xerophytic (Mai 1970; Kvaček
& Walther 1989).
Cedrelospermum flichei (Saporta) Hably et Thiébaut
2002 Cedrelospermum flichei (Saporta) Hably et Thiébaut, p. 81, pl. 4,
figs. 6—7; pl. 7, figs. 1—9
M a t e r i a l : BP.2001.676.1; BP.2001.829.1; BP.2007.128.1;
D e s c r i p t i o n : Simple, asymmetric leaves. Leaves peti-
olate, petiole short, 1.5—3 mm long. Leaves lanceolate, or
narrow ovate, length 2.5—5 cm, width 0.5—1.3 cm. Base
asymmetric and cuneate, apex acute, attenuate. Margin ser-
rate, at the basal part of leaves entire. Teeth simple and acute
or obtuse. Venation craspedodromous, midvein stout, sec-
ondaries arising at angles of 50—65° and entering the teeth.
Some secondaries forked towards the margin.
ERDEI and RÁKOSI
Fig. 3. 1 – Chara sp. (BP.2001.792.1). 2, 3 – Daphnogene sp. (BP.2001.693.1 ). 4 – Lauraceae gen. et sp. (BP.2001.722.1). 5 – cf. Eotri-
gonobalanus furcinervis (Rossmässler) Walther et Z. Kvaček (BP.2007.137.1). 6 – cf. Eotrigonobalanus furcinervis (Rossmässler) Walther
et Z. Kvaček (BP.2001.678.1). 7 – cf. Eotrigonobalanus furcinervis (Rossmässler) Walther et Z. Kvaček (BP.2001.882.1). 8 – Ce-
drelospermum flichei (Saporta) Hably et Thiébaut (BP.2007.128.1). 9 – Cedrelospermum flichei (Saporta) Hably et Thiébaut
(BP.2001.676.1). 10, 11 – Cedrelospermum flichei (Saporta) Hably et Thiébaut (BP.2001.829.1). Scale bars are 1 cm.
THE MIDDLE EOCENE FLORA OF CSORDAKÚT (N HUNGARY)
D e t e r m i n a t i o n a n d d i s c u s s i o n : The leaves are
comparable to the leaves of C. flichei from the Hungarian Oli-
gocene and Miocene (Hably & Thiébaut 2002). The species
was described from the Early Oligocene of Hungary from the
Tard Clay Formation, namely Nagybátony-Újlak (Budapest)
and Eger-Kiseged (Bükk, NE Hungary), and from the Middle
Miocene flora of Magyaregregy. Outside Hungary the species
was recorded among others from the Oligocene of Cereste and
Rott, the Miocene of Randecker Maar, etc. (Hably & Thiébaut
2002). Kovar-Eder et al. (2004) discuss an additional species
C. ulmifolium (Unger) Kovar-Eder et Kvaček based on leaves
from the Early-Middle Miocene locality of Parschlug and also
assign C. flichei leaves from Magyaregregy to this species. Al-
though the leaves from Parschlug are morphologically similar,
it is probable that the Eocene and Miocene leaves represent
different species. The characteristic fruits of Cedrelospermum
quite frequently recorded in the Hungarian Early Oligocene
have not been detected.
Based on the abundance of Cedrelospermum remains in
lacustric deposits of volcanic areas together with the abundant
production of fruits Manchester (1989, p. 274) considers its
species to be early successional colonizers of open habitats.
Cedrelospermum ?flichei (Saporta) Hably et Thiébaut
M a t e r i a l : BP.2007.219.1.
D e s c r i p t i o n : Leaf fragmented. Shape asymmetric, lan-
ceolate, length (fragmentary) 8.8 cm, width 1.2 cm. Apex
acute, attenuate. Margin serrate along most of its length,
teeth simple, prominent. Teeth apices obtuse but getting
slightly acute close to the apex. Venation craspedodromous,
midvein stout and slightly curved. Secondaries, hardly ob-
servable due to poor preservation, arise at angles of 50—60°
and end in the teeth.
D e t e r m i n a t i o n a n d d i s c u s s i o n : The leaf shows
different characters from Cedrelospermum flichei (Saporta),
which Hably et Thiébaut described from the Early Oligocene
of Hungary (Hably & Thiébaut 2002), by having a relatively
large, lanceolate leaf form and prominent teeth contrasting
the smaller and broader lamina of C. flichei.
The lanceolate leaf with simple and mostly obtuse teeth is
more comparable to the leaves of Tremophyllum sp. from
Messel, one specimen of which was found attached with the
fruits of Cedrelospermum leptospermum (Ettingshausen)
Manchester (Wilde & Manchester 2003). Since we had only
one fragmentary example we hesitated to identify our speci-
men with the leaves of Tremophyllum sp. sensu Wilde &
Manchester (2003). The latter were also compared to some
North American leaves found associated and attached to
fruiting branches of Cedrelospermum (Wilde & Manchester
2003). Our leaf matches C. lineatum (Lesquereux), which
Manchester described from North America, by having prom-
inent simple teeth along most of its length and a more asym-
metric leaf base. By its larger size it differs from both North
American species. The leaves of Tremophyllum tenerrimum
(Weber) Rüffle from the Oligocene of Rott (Rüffle 1963)
differ in having smaller and broader lamina and less pro-
Cedrelospermum leaves and fruits, C. flichei and C. aque-
nse Saporta, have been documented from the Early Oli-
gocene Tard Clay floras (Hably & Thiébaut 2002). This is
one of the earliest leaf records of the genus.
cf. Myrica longifolia Unger
M a t e r i a l : BP.2001.859.1.
D e s c r i p t i o n : Leaf narrow elliptic, petiolate. Length of
lamina 5.8 cm, width 0.8 cm, base acute and decurrent, apex
acute. Margin seems to be irregularly toothed, both acute and
rounded teeth occur. (Margin and venation hardly observ-
able due to poor preservation.) Midrib stout, secondary vena-
tion dense, veins arise at angles of 70—80°.
D e t e r m i n a t i o n a n d d i s c u s s i o n : Leaves described
as M. longifolia are widespread in the Early Cenozoic of Eu-
rope – for example in the Eocene of Geiseltal (Rüffle
1976), the Early Oligocene of Häring – here even dominant
(Ettingshausen 1853), Haselbach (Weisselster Becken; Mai
& Walther 1978), Monte Promina (Ettingshausen 1855), etc.
Rüffle (1976) gives a quite large synonym list and a thor-
ough discussion on the species.
The leaf is quite elongate (l/w > 7) similar to the forms de-
scribed by Mai & Walther (1978) from Haselbach. Rüffle
(1976) and Mai & Walther (1978) evaluated these leaves as
xerophytic forms of the fossil species. Moreover, Rüffle (1976)
defined M. longifolia as a “trockenatlantische Myrica Art”.
Material: BP.2001.820.1;?BP.2001.848.1; ?BP.2001.843.1.
D e s c r i p t i o n : Leaves all fragmented, length of the spec-
imen BP.2001.820.1 (fragmentary) 11.5 cm, width 1.9 cm
(13.6 and 3.4 cm in ?BP.2001.848.1). Apex not preserved,
base decurrent. Margin irregularly toothed, teeth acute or ob-
tuse. Venation dense, semicraspedodromous, midrib moder-
ately thick, secondary veins dense, arising at angles of
70—80°, running to the teeth or terminating between teeth.
D e t e r m i n a t i o n a n d d i s c u s s i o n : The leaf is much
larger than the leaves of M. longifolia. Its macromorphology
shows similarity to M. lignitum (Unger) Saporta occurring in
younger floras, however, due to the lack of cuticular struc-
tures the comparison remains tentative. Nevertheless, M. lig-
nitum proved by cuticular structure (see Kovar 1982) has so
far been recorded mostly from younger floras, such as the
Late Oligocene flora of Linz (Kovar 1982), the Miocene flo-
ras of Parschlug (Austria; Kovar-Eder et al. 2004), and based
on macromorphology from numerous localities (see also
Knobloch & Kvaček 1976).
Comptonia difformis (Sternberg) Berry
1821 Asplenium diforme Sternberg, p. 33, pl. 24, fig. 1
1828 Comptonia acutiloba Brongniart, pp. 141, 143, 209
1971 Comptonia acutiloba Brongn. – Bůžek, pl. 9, figs. 1—8
ERDEI and RÁKOSI
Fig. 4. 1, 2 – Cedrelospermum sp. (BP.2007.219.1). 3 – cf. Myrica longifolia Unger (BP.2001.859.1). 4, 5 – Myrica sp. (BP.2001.820.1).
6 – Comptonia difformis (Sternberg) Berry (BP. 2007.135.2). 7 – Comptonia schrankii (Sternberg) Berry (BP. 2001.734.1). 8 – Compto-
nia schrankii (Sternberg) Berry (BP.2001.711.1). 9 – Comptonia schrankii (Sternberg) Berry (BP.2001.864.1). 10 – Leguminosae gen. et
sp. (BP.2001.828.1). 11 – Leguminocarpon sp. (BP.2007.134.2), the seed is indicated by an arrow. Scale bars are 1 cm.
THE MIDDLE EOCENE FLORA OF CSORDAKÚT (N HUNGARY)
M a t e r i a l : BP.2001.832.1; BP.2007.135.2.
D e s c r i p t i o n : Leaves (all fragmented) lanceolate, peti-
ole, basal and apical parts not preserved. Entire length more
than 3 cm, width 1.2 cm. Lobes alternate or opposite, sessile
to midrib, shape of lobes varying. Apices of lobes acute.
Margin entire. Midrib moderate, secondaries arise at angles
of 70—80° and curve upwards.
D e t e r m i n a t i o n a n d d i s c u s s i o n : For priority of
“acutiloba”, synonym lists and a review of fossil Comptonia
remains see Bůžek (1971), Rüffle (1976) and J. Kvaček
(2004). The species has been recorded mainly in older Ceno-
zoic floras up into the Miocene, as in the North Bohemian
Basin (Chomutov; Bůžek 1971), the Eocene flora of Gei-
seltal (Rüffle 1976), the floras of the Haselbach Series
(Weisselster Basin), the Hungarian Early and Late Oligocene
(H-borehole, Eger-Wind; Andreánszky 1955), etc. Leaf frag-
ments from Csordakút are slender, for example the speci-
mens from Geiseltal (C. difformis (Sternberg) Berry) are
3—4 cm wide (Rüffle 1976). Comptonia specimens from
Messel have similar dimensions (width max. 1 cm) to those
from Csordakút but the lobes are toothed (Wilde 1989). In
the Early Oligocene flora of Santa Giustina (Principi 1916)
Comptonia leaves are more available. Slender leaves have
toothed lobes. Remains recorded as C. acutiloba (or C.
dryandroides Unger, etc.) comprise various leaf forms, thus
the group certainly needs revision. Below we give some re-
marks on the macromorphology of fossil Comptonia leaves
recorded from the Paleogene deposits of Hungary.
Linear lobate leaves of C. dryandroides Unger partly recall-
ing C. difformis (Myrica acutiloba var. dentata Andreánszky
in Andreánszky 1955) were documented in the Late Oli-
gocene Eger-Wind locality, in the upper level flora (Kvaček &
Hably 1991) but its lobes are often finely toothed. Lower Oli-
gocene deposits of Hungary also yielded leaves of C. acutilo-
ba (C. difformis): from the H-boreholes (Budapest, Vörösvári
út) two types are available, one (BP.78.254.3) is morphologi-
cally well comparable to the specimen from Csordakút, the
other type (BP.78.154.3) has more regular lobes and a broader
(2.3 cm) lamina. Both are entire margined. Two specimens
comparable to those from Csordakút (entire margin, similar
dimension) are also available from Eger-Kiseged (North Hun-
gary, BP.2004.491.1, BP.2004.722.1) A Comptonia specimen
recorded from Nagybátony-Újlak (Budapest) has entire-mar-
gined lobes but differs in its broader lamina (3 cm).
According to Rüffle (1976) leaves of the species are often
accompanied by Myrica longifolia Unger. C. difformis is
mostly compared with the modern C. peregrina (Linnaeus)
Coulter, however, leaves of the modern species are consider-
ably smaller (Bůžek 1971). Today the genus is monotypic
spread in warm temperate regions of North America, often
on dry sands (Krüssmann 1960). Considering the relict char-
acter of modern Comptonia, the extrapolation of its ecologi-
cal requirements to the fossil is problematic.
Comptonia schrankii (Sternberg) Berry
1822 Aspleniopteris schrankii Sternberg, p. 29, pl. 21, fig. 2
1906 Comptonia schrankii (Sternb.) Berry, p. 514
1975 Comptonia schrankii (Sternb.) Berry – Palamarev & Petkova,
p. 212, Pl. 2, fig. 1
M a t e r i a l : BP.2001.683.1; BP.2001.711.1; BP.2001.734.1;
BP.2001.795.1; BP.2001.864.1; ?BP.2001.824.1; 2007.131.1;
D e s c r i p t i o n : Leaves (all fragmented) lanceolate, basal
and apical parts not preserved. Lobes alternate or opposite,
and sessile to midrib. Original length even more than 4.5 cm,
width 0.3—0.7 cm, shape of lobes regular. Apices of lobes
acute. Margin entire. Midrib stout, secondaries arise at an-
gles of 75—90°.
D e t e r m i n a t i o n a n d d i s c u s s i o n : The smaller size of
the leaves and regular lobes distinguish these leaves from C.
difformis. However, some transitional leaf forms indicate the
need for revision. Furthermore, leaves of C. difformis and C.
schrankii fall in the morphological variability of leaves of
modern Comptonia peregrina (Berry 1906: figs. 1—6).
C. schrankii has been recorded among others from the Paleo-
gene of Bulgaria (Palamarev & Petkova 1975), from Häring in
Tyrol (Dryandra brongniartii Ettingshausen, Ettingshausen
1853), the Eocene Somod ( = Drieňovice, Slovakia; Hably pers.
com.; BP.97.217.1, BP.97.219.1, BP.97.220.1). From Hun-
gary specimens with similar morphology are available from
the Early Oligocene Eger-Kiseged locality (BP.2004.301.1,
BP.2004.302.1). C. difformis specimens from the Late Oli-
gocene of Austria (Linzer Raum; Kovar 1982: p. 79, Taf. 11,
figs. 14, 15) display similar traits (width 0.5—0.7 cm) to our
specimens, but apices of lobes seem to be obtuse. Leaves com-
parable to C. schrankii also appear in the Early Oligocene flora
of Santa Giustina (Hably, pers. com.).
Leguminosae gen. et sp.
M a t e r i a l : BP.2001.828.1.
D e s c r i p t i o n : Leaflet small, 1.3 cm long and 0.7 cm
wide, narrow obovate, ?sessile. Base acute, apex rounded.
Margin entire. Venation camptodromous, midvein stout at
the base, tapering toward the apex. Higher order venation not
observable due to poor preservation.
D e t e r m i n a t i o n a n d d i s c u s s i o n : The macromor-
phology of the leaflet recalls Leguminosae.
M a t e r i a l : BP.2007.134.2.
D e s c r i p t i o n : Fruit dehiscent, stipitate, single seeded.
Bent at the junction of valves and stipe, valves 3.5 cm long,
1.6 cm wide, shape of valves elliptic. Apex and base acute
(almost obtuse), stipe fragmented but at least 1 cm long,
0.8 mm wide, valve venation not observable. Seed indistinct,
seems slightly oblong, 9 mm long and 8 mm wide but pres-
ervation could cause distortion of shape.
D e t e r m i n a t i o n a n d d i s c u s s i o n : Among fossil le-
gumes Podocarpium podocarpum (A. Braun) Herendeen
(Herendeen 1992a,b), occurring frequently in Neogene flo-
ERDEI and RÁKOSI
ras, shows similarity to our fossil in having dehiscent, stipi-
tate single-seeded fruits, equipped with long or even longer
stipe. However, different characters are the elliptic, more
elongate shape of Podocarpium, the acute base and apex of
its valves, the mostly straight or less bent junction of valves
and stipe and smaller dimension.
The fruit structure indicates a relationship to the Legumino-
sae family, to the Caesalpinioideae or Papilionoideae subfami-
ly. The angle formed at the junction of the stipe and valves,
and the fruit apex shape are not useful characters since they
are variable in many genera, also in single-seeded ones (Her-
endeen 1992a; Herendeen & Crane 1992). Single-seeded stipi-
tate fruits occur in Caesalpinioideae and Papilionoideae, in the
tribes of Caesalpinieae, Cercideae, Detarieae, Mimoseae,
Swartzieae, Sophoreae and in several unrelated genera, such
as Gleditsia, Apuleia, Adenolobus, Brenierea, Griffonia, Co-
paifera, Peltogyne, Zenkerella, Ormosia, etc. Since charac-
ters, such as the position of placentation and pattern of valve
venation which are important to distinguish fruits of these
genera (Herendeen 1992a) are indistinctly observable in our
fossil its comparison with modern legume taxa is limited.
Leguminosae gen. et sp. vel Anacardiaceae gen. et sp.
M a t e r i a l : BP.2001.839.1.
D e s c r i p t i o n : Leaflet slightly asymmetric, narrow obo-
vate, 2 cm long and 1 cm wide, probably sessile. Base asym-
metric, acute, apex rounded. Margin entire. Venation
camptodromous, midvein moderately thick, slightly curved.
Higher order venation not observable due to poor preservation.
D e t e r m i n a t i o n a n d d i s c u s s i o n : Leaflet morpholo-
gy refers to an affinity with Leguminosae or Anacardiaceae.
?Zizyphus zizyphoides (Unger) Weyland
M a t e r i a l : BP.2001.714.1; (counterpart BP.2001.720.1).
D e s c r i p t i o n : Leaf simple petiolate leaf 9.3 cm long and
2.9 cm wide, petiole 1.7 cm long. Leaf lanceolate, apex
acute, base acute and slightly asymmetric. Margin serrate,
teeth acute, in the apical part of the lamina teeth more round-
ed, margin even seems to be crenate. Sinuses angular. Vena-
tion imperfect basal acrodromous. Midrib moderately thick
and straight. Besides the pair of stout secondary veins arising
at the base, secondaries and tertiary veins form a fine net-
work. Tertiary veins arising from the basal secondaries run
toward the margin forming loops. Although faintly observ-
able some small veins originating from where the loops seem
to enter the teeth.
D e t e r m i n a t i o n a n d d i s c u s s i o n : The morphology
of the leaf (acrodromous venation, teeth) corresponds more
or less to Z. zizyphoides. The imperfect acrodromous vena-
tion, however, contrasts with the generally perfect acrodro-
mous venation in the leaves of this species. The margin
seems to be more serrate than crenate and the leaf size is
quite large as compared to the leaves of Z. zizyphoides.
Z. zizyphoides has frequently been recorded, e.g. in the
Early Oligocene of Häring (Ceanothus zizyphoides Unger,
Ettingshausen 1853), Sotzka (Ceanothus zizyphoides Unger,
Unger 1847), Eocene of Macedonia (Ovce Polje; Mihajlovic
& Ljubotenski 1994), Paleogene of Bulgaria (Palamarev &
Petkova 1975), the Eocene flora of Akhaltsikhe (Georgia;
Avakov 1989), etc. In the Hungarian Paleogene its frequent
occurrence is known from the Early Oligocene Tard Clay
floras (Andreánszky 1963; Hably 1979), and it was also de-
scribed in the Eocene flora of Tatabánya (Hably 1985b).
M a t e r i a l : BP.2001.738.1.
D e s c r i p t i o n : Leaf small petiolate, lanceolate. Length
3.4 cm, width 0.7 cm. Apex and base acute. Margin serrate
with acute teeth along the entire margin. Sinuses acute. Ve-
nation perfect basal acrodromous with a pair of distinct sec-
ondary veins almost reaching the leaf apex. Higher order
venation not preserved.
D e t e r m i n a t i o n a n d d i s c u s s i o n : The morphology
of this leaf recalls Zizyphus zizyphoides. A distinct trait con-
trasting leaves of the species is the pair of basal secondaries
that run nearly into the leaf apex.
Sloanea nimrodi (Ettingshausen) Kvaček et Hably
1869 Cissus nimrodi Ettingshausen, p. 3, pl. 40, figs. 3—4, 6—10
2001 Sloanea nimrodi (Ettingshausen) Kvaček et Hably – Kvaček,
Hably & Manchester, p. 117, pl. 4, figs. 6—7
M a t e r i a l : BP.2001.715.1; BP.2001.771.1; BP.2001.781.1;
?BP.2001.713.1; BP.2007.128.1; BP.2007.135.2.
D e s c r i p t i o n : Leaves simple, petiolate, lamina 2.5—5.2 cm
long (or more) and 1.5—3.8 cm wide, petiole 1.3—1.5 cm long.
Leaves narrow or wide obovate to elliptic, base acute to nearly
obtuse, apex rounded to obtuse. Margin entire in the lower half
of the lamina, in the upper part subentire, crenulate or toothed.
Teeth obtuse. Venation craspedodromous/semicraspedodro-
mous. Midrib moderately thick and straight. 5(—6) pairs of sec-
ondaries, the lowermost pair basal or suprabasal, almost
opposite, run nearly straight and opposite, the higher pairs alter-
nate, course curved. The uppermost pair (observable in the
specimen BP.2001.715.1, Fig. 5.5) nearly rounded curving
back towards the midrib. Secondaries craspedodromous, often
fork near the margin and exmedially give off a vein entering a
tooth. Tertiary veins perpendicular to the secondaries, percur-
rent, higher order venation reticulate, areoles well developed.
D e t e r m i n a t i o n a n d d i s c u s s i o n : The gross mor-
phology of the leaves is closely comparable to the leaves of
Sloanea. Sloanea elliptica (Andreánszky) Kvaček et Hably
described from the Early Oligocene of Hungary and Slove-
nia (Kvaček et al. 2001) shows different characters, namely a
definitely larger size of leaves, more elongated lamina, higher
number of secondaries (6—8). S. artocarpites (Ettingshausen)
THE MIDDLE EOCENE FLORA OF CSORDAKÚT (N HUNGARY)
Fig. 5. 1 – Leguminosae gen. et sp. vel Anacardiaceae gen. et sp. (BP.2001.839.1). 2 – ?Zizyphus zizyphoides (Unger) Weyland
(BP.2001.714.1). 3 – ?Zizyphus sp. (BP.2001.738.1). 4 – Sloanea nimrodi (Ettingshausen) Z. Kvaček et Hably (BP.2007.135.2), the fruit of
Sloanea sp. is indicated by an arrow. 5 – Sloanea nimrodi (Ettingshausen) Z. Kvaček et Hably (BP.2001.715.1). 6 – Sloanea nimrodi (Et-
tingshausen) Z. Kvaček et Hably (BP.2001.771.1). 7 – Sloanea nimrodi (Ettingshausen) Z. Kvaček et Hably (BP.2007.128.1). 8 – Berberi-
daceae gen. et sp. (BP.2001.870.1). 9 – Dicotylophyllum sp. 1 (BP.2001.796.1). Scale bars are 1 cm.
Kvaček et Hably described from the Oligocene of North Bo-
hemia is differentiated from S. elliptica by slightly devel-
oped and sharper dentation of the margin and a less distinct
subbasal pair of secondaries (Kvaček et al. 2001). Leaves of
Sloanea peolai (Principi) Hably, Tamás et Cioppi (Hably et
al. 2007) described from the Oligocene flora of Chiavon (Ita-
ly) differ from our leaves by having a lanceolate shape with
small teeth on the margin. Leaves of S. nimrodi (Etting-
shausen) Kvaček et Hably recorded from the Late Eocene of
North Bohemia are of smaller dimension than S. elliptica (Z.
ERDEI and RÁKOSI
Kvaček et al. 2001), in this respect resembling our fossils.
The type specimen of the species (Pl. 4, figs. 6—7 in Z.
Kvaček et al. 2001) shows a more elongated lamina in con-
trast to our fossils, however, studying other S. nimrodi speci-
mens in the collection of Ettingshausen’s type and original
material from Bilina (Cissus nimrodi, figured on Pl. 40,
figs. 6—10 in Ettingshausen 1869; Pl. 10, figs. 1—6 in Hably
et al. 2001) leaves with obovate lamina, and wide obtuse
apex were also found. The specimens from Csordakút are of
distinctly obovate shape with rounded apex. These features
may represent the natural variability of the leaves since elon-
gate leaves also occur.
The earliest record of Sloanea so far, Sloanea nimrodi has
been recorded in North Bohemia from the Late Eocene/?Ear-
ly Oligocene flora of Kučlín (Z. Kvaček et al. 2001). Thus,
the specimens from Csordakút proves an even earlier occur-
rence of the genus.
Sloanea sp. (fruit)
M a t e r i a l : BP.2007.135.2.
D e s c r i p t i o n : Impression of a fruit fragment, the capsule
(or isolated valve?), elliptic, 1.4 cm long, 1.1 cm wide.
Pedicel not preserved. On external surface (at sides of the im-
pression) hardly observable structure – ?spines. Due to poor
preservation it is not clear whether the complete capsule or an
isolated valve, the external or the inner surface is fossilized.
D e t e r m i n a t i o n a n d d i s c u s s i o n : The fruit remain is
associated with a Sloanea leaf. The morphological traits of
the impression recall that of Sloanea fruits, such as Sloanea
eocenica (Rásky) Kvaček, Hably et Manchester described
from the Early Oligocene of Hungary (Nagybátony-Újlak, Z.
Kvaček et al. 2001). The size of the fossil, however, is small-
er. The morphology of the spines covering the valves, which
may serve as distinguishing character among species is very
indistinct in our fossil. Other Sloanea fruits recorded from
Suletice-Berand or Markvartice (Late Oligocene), namely
Sloanea sp. (Z. Kvaček et al. 2001) are slightly smaller than
S. eocenica. Due to the lack of detailed morphology of the
fossil from Csordakút a comparison remains tentative.
?Berberidaceae gen. et sp.
M a t e r i a l : BP.2001.777.1; BP.2001.870.1.
D e s c r i p t i o n : Fragmentary remains of leaves, 2.2—5.9 cm
(fragmentary) long, 0.8—1 cm wide. Leaves ?coriaceous, very
narrow elliptic, base not preserved, apex acute-attenuate. Mar-
gin toothed with distinct, regularly spaced acute teeth. Teeth
concave at the apical and basal sides. Sinuses rounded. Mid-
vein stout, higher order venation not preserved.
D e t e r m i n a t i o n a n d d i s c u s s i o n : The few observ-
able traits of the leaf fragments recall Berberidaceae. The
teeth appear sharper, more regular and distinct than in leaves
Dicotylophyllum sp. 1
M a t e r i a l : BP.2001.796.1; (counterpart BP.2001.878.1).
D e s c r i p t i o n : Fragment of a petiolate leaf. Petiole stout,
0.9 cm long. Leaf obovate, 3.5 cm (fragmentary) long and
1.7 cm wide. Base rounded slightly asymmetric, apex not
preserved. Margin entire. Venation basal acrodromous. The
basal pair of secondaries run almost straight into the upper
half of the lamina. Tertiary veins arising from the basal sec-
ondaries run towards the margin forming loops.
D e t e r m i n a t i o n a n d d i s c u s s i o n : The leaf fragment
resembles leaves described as Matudaea, (e.g. cf. Matudaea
menzelii Walther, Early Oligocene flora of the Tard Clay
Formation; Kvaček & Hably 1998), however, acrodromous
venation occurs in a great number of angiosperm families,
including Lauraceae, Hamamelidaceae, Rhamnaceae, Melas-
tomataceae, etc. Due to the lack of cuticular structure and di-
agnostically valuable traits the systematic affinity remains
Dicotylophyllum sp. 2 (?Juglandaceae)
M a t e r i a l : BP.2007.135.2.
D e s c r i p t i o n : Simple petiolate leaf of 6 cm length and
1.6 cm width. Petiole stout, 3 mm long. Leaf narrow elliptic,
apex ?obtuse, base asymmetric, acute. Margin toothed, teeth
acute, regularly spaced along the margin. Sinuses rounded.
Venation craspedodromous, midvein moderately thick. Sec-
ondaries, min. 15 pairs, arise at angles of 50—60° from the
primary vein and run mostly straight to the teeth. Close to
the teeth secondaries slightly curved. Tertiary veins percur-
rent, observable only at the basal part of the lamina.
D e t e r m i n a t i o n a n d d i s c u s s i o n : This leaf shows
similarity to Dryophyllum callicomifolium (Andreánszky)
Kvaček et Hably (Z. Kvaček & Hably 1991) in the slender
shape and form of teeth. This species occurs in the lower-
most level of the Late Oligocene Eger-Wind assemblage and
the Early Oligocene Tard Clay floras. However, the slightly
asymmetric base and the more or less straight secondaries
distinguish our leaf from both Dryophyllum callicomifolium
and Eotrigonobalanus (cuneate, often decurrent base, curved
secondaries) and raise an affinity to Juglandaceae.
Dicotylophyllum sp. 3
M a t e r i a l : BP.2001.804.1; BP.2001.727.1.
D e s c r i p t i o n : Compound leaf, imparipinnate, with 7 at-
tached, and one isolated leaflet. Leaflets petiolate, petiolule
0.7—1 cm. Leaflets opposite, asymmetric and lanceolate,
6.3—6.5 cm long and 1—1.4 cm wide. Isolated leaflet of big-
ger dimension, fragmentary, 3 cm wide, length much more
than 6 cm. Base of leaflets distinctly asymmetric, acute or
nearly obtuse. Margin serrate. Teeth simple and acute or
acuminate. Sinuses angular. Venation camptodromous-semi-
craspedodromous. Midvein moderately thick, a great number
of secondaries, more than 15 pairs, arise at angles of 70—80°.
Between secondaries intersecondary veins observable. Adja-
cent secondaries often join close to the margin and a vein
branches off the loop and ends in the teeth.
THE MIDDLE EOCENE FLORA OF CSORDAKÚT (N HUNGARY)
Fig. 6. 1 – Dicotylophyllum sp. 2 (BP.2007.135.2). 2, 3 – Dicotylophyllum sp. 3 (BP.2001.804.1). 4 – Carpolithus sp. (BP.2001.708.1).
5 – Smilax sp. 1 (BP.2001.774.1). 6 – ?Smilax sp. (BP.2001.739.1). Scale bars are 1 cm.
Determination and discussion: Compound leaves with peti-
olate leaflets occur in numerous families (e.g. Simaroubaceae,
Oleaceae, Meliaceae etc.). Due to the lack of epidermal struc-
ture the systematic affinity of these leaves remains obscure.
M a t e r i a l : BP.2001.708.1 (counterpart BP.2001.804).
D e s c r i p t i o n : Fruit 4.4 cm long and 1.7 cm wide, ?bilat-
eral symmetric, the central part (seed?) rounded 1 cm long
and 1.3 cm wide. No venation of the central or lateral parts
D e t e r m i n a t i o n a n d d i s c u s s i o n : The detailed mor-
phology is not observable. At first glance it resembles a
winged fruit, however, the homogeneous texture of the cen-
tral and lateral parts and the distinct margin of the lateral
parts contrast this. Its structure may recall Leguminosae.
ERDEI and RÁKOSI
Smilax sp. 1
M a t e r i a l : BP.2001.716.1; BP.2001.774.1.
D e s c r i p t i o n : Simple petiolate leaves. Petiole (pre-
served in BP.2001.716.1) thick, 0.6 cm long. Leaves ovate,
3.5—4.5 cm long and 2.7—3 cm wide. Base rounded, apex
rounded and slightly emarginate (preserved only in
BP.2001.716.1). Margin entire. Venation perfect, basal acro-
dromous with 5 primary veins originating from the base at
the attachment of the petiole. Three central veins almost
equally thick, the two veins close to the margin slightly thin-
ner. Veins run forming strongly developed, curved arches
and join at the apex. Finer veins forked, percurrent and arise
at angles of 80—90° from the primary veins.
D e t e r m i n a t i o n a n d d i s c u s s i o n : Fossils of this ge-
nus have been frequently described from the Paleogene. Our
specimens differ from those of Smilax sp. 1 and sp. 2 de-
scribed from the Eocene of Staré Sedlo (Knobloch et al.
1996) in having primary veins diverging from one point at
the base, at the attachment of the petiole to the lamina and in
having a rounded base, slightly emarginate apex and ovate
leaf form. From the Middle Eocene of Messel (Wilde 1989)
three species of Smilax (with cf.) were recorded based on
macro- and cuticular structures. Our leaves displaying ovate
shape, rounded base and apex thus differing from the narrow
ovate leaves from Messel.
Other Paleogene records of Smilax mostly differ by a char-
acteristic cordate leaf base (e.g. Kovar 1982), however, the co-
occurrence of oval and hastate/cordate forms may reflect the
variability of leaves of the same species, for example, Smilax
weberi Wessel in Wessel et Weber in the Late Oligocene
Eger-Wind flora (Andreánszky 1966; Kvaček & Hably 1991).
M a t e r i a l : BP.2001.739.1 (BP.2001.730.1).
D e s c r i p t i o n : Leaf lanceolate, lamina 9.1 cm long and
2.9 cm wide and slightly asymmetric, base obtuse. Apex
probably acute. No teeth observed, however, margin heavily
damaged. Venation perfect basal acrodromous with 5 prima-
ries originating from one point at the base. Higher order ve-
nation not preserved.
D e t e r m i n a t i o n a n d d i s c u s s i o n : The specimen
with elongate lamina and asymmetric leaf base may also rep-
The flora is preserved in lacustric facies in freshwater
(? brackish), sometimes clayey limestones overlying the first
coal seam. The main factors presumed to play an essential
role in the transport of plant material into the place of burial
are water and wind. However, winged fruits and seeds as po-
tential evidence of wind transport are totally missing from
the assemblage. This may be attributable to various facts, for
example, wind played a minor role in transport – a relative-
ly closed vegetation around the lake impeding wind trans-
port, a low proportion of taxa producing winged fruits.
Leaves are always randomly oriented, spaced at nearly regu-
lar distances from each other without overlapping suggesting
calm depositional settings.
Flora and vegetation
The fossil plant assemblage from Csordakút comprises
mostly remains of angiosperms. In addition, algae are repre-
sented by numerous “stem”-like fragments of fresh-(brackish)
water Characeae. Angiosperms documented macromorpho-
logically belong to Lauraceae (Daphnogene sp.), Ulmaceae
(Cedrelospermum div. sp.), Fagaceae (cf. Eotrigonobalanus
furcinervis), Myricaceae (Myrica sp., Comptonia div. sp.), Le-
guminosae (leaves and fruit), Rhamnaceae (?Zizyphus zizy-
phoides), Elaeocarpaceae (Sloanea nimrodi, Sloanea sp. fruit)
and Smilacaceae (Smilax div. sp.).
The absence of gymnosperms in the fossil assemblage
may reflect their subordinate role in the vegetation, or it is
attributable to the depositional setting, perhaps gymno-
sperms flourished far from the depositional basin.
Although the evaluation of taxon frequencies or the role of
individual taxa in forming the vegetation is greatly hindered
due to the great number of undeterminable specimens, taxa
of both zonal and intrazonal vegetation types are recogniz-
able. A lacustric environment is unequivocally proved by the
frequent occurrence of Characeae. Myrica and Comptonia
were members of the intrazonal vegetation (swamp) sur-
rounding the lake. Eotrigonobalanus characterized by a wide
ecological tolerance was probably the member of both zonal
and intrazonal vegetation types. The zonal associations com-
prised Lauraceae, Sloanea, Eotrigonobalanus and Zizyphus.
Open habitats are suggested by the occurrence of Ce-
drelospermum and Leguminosae. A “subxerophytic” charac-
ter of the zonal vegetation is supported by some evidence
namely the occurrence of small-leaved Leguminosae and co-
riaceous leaves with teeth on the margin. Other taxa suggest-
ing a “subxerophytic” vegetation, like Zizyphus zizyphoides
are documented by a single, uncertain specimen or are even
absent as conifers with scale-like foliage.
Comparisons with Early Paleogene assemblages of Hungary
The Eocene flora of Tatabánya (Hably 1985a; Erdei &
Wilde in prog.) was fossilized in marine clays, clay marls
contrasting the lacustric environment of Csordakút. A simi-
lar feature of the Tatabánya assemblage is the absence of
gymnosperms (a single fragmentary specimen in Tatabánya).
Taxa shared by the Csordakút flora are Lauraceae, ?Zizyphus
zizyphoides, Leguminosae, (Anacardiaceae). Remains of
palms recorded in Tatabánya are missing in Csordakút,
whereas Daphnogene, Eotrigonobalanus, Sloanea and Myri-
caceae were not found in Tatabánya. Furthermore, the
THE MIDDLE EOCENE FLORA OF CSORDAKÚT (N HUNGARY)
Tatabánya assemblage is dominated by a lanceolate leaf-type
(?Lauraceae) characterized by an entire, slightly wavy leaf
margin and presumably coriaceous texture which did not ap-
pear in Csordakút. Flora and vegetation disparities may re-
flect the different facies of the two localities. In contrast to
the Eocene assemblages with the scarcity of gymnosperms,
the Early Oligocene floras of the Tard Clay Formation are
well-characterized by conifers (Z. Kvaček & Hably 1998),
like Doliostrobus taxiformis (Sternberg) Kvaček var. hun-
garicus (Rásky) Kvaček et Hably, Tetraclinis salicornioides
(Unger) Kvaček as well as T. brachyodon (Brongniart) Mai
et Walther, Calocedrus suleticensis (Brabenec) Kvaček,
moreover, cycads were recorded (Ceratozamia floersheimen-
sis (Engelhardt) Kvaček; Z. Kvaček 2002). Winged fruits ap-
pearing with high diversity in the Tard Clay floras have not
been found in Csordakút.
Shared taxa are Eotrigonobalanus furcinervis, ?Zizyphus
zizyphoides, Daphnogene, Myrica, Comptonia, Cedrelosper-
mum, Leguminosae and Sloanea.
Cedrelospermum frequently recorded by C. flichei in the
Early Oligocene Tard Clay floras, seems to occur with at
least two species during the Eocene. These are C. flichei and
Cedrelospermum sp. indicating morphological similarity to
leaves of Tremophyllum sp. from the Eocene flora of Messel
(Wilde & Manchester 2003).
Similarly, Sloanea is quite widespread in the Hungarian
Early Oligocene with one species, S. elliptica (valid name
according to priority S. olmediaefolia, Hably & Kvaček
2007, in prog.). It appears with an additional species in Csor-
dakút, S. nimrodi, new to the Hungarian fossil record. S.
nimrodi suggests a floristic relation to the Late Eocene of
North Bohemia (Kučlin).
The Eocene flora of Lábatlan (Kovács 1959, 1961) needs a
revision to provide a precise comparison. Zlatko Kvaček stud-
ied the assemblage during a study trip to Budapest in 1989
and gave some remarks on the flora (Knobloch et al. 1996).
He recognized two leaf types dominating the assemblage, one
with an affinity to Juglandaceae and another resembling the
leaf form of Apocynophyllum helveticum Heer. Remains of
Eotrigonobalanus were not proved. Except for some taxodia-
ceous twigs Z. Kvaček did not find common elements with the
Eocene assemblage of the Staré Sedlo layers in Bohemia. To
give floristic similarities between the Lábatlan and Csordakút
floras the fossil collection from Lábatlan should be revisited.
However, it may be noted that the subordinate role of conifers
parallels other Eocene assemblages from Hungary.
Comparisons to the Paleogene floras of Europe
Since leaves of the Csordakút assemblage are fossilized as
impressions without organic matter resulting in a great num-
ber of indeterminable specimens a precise systematic com-
parison with other coeval floras of Europe is rather limited.
However, some remarks on floristic relations should be
Numerous coeval floras of Central Europe, the Early Paleo-
gene floras of eastern Central Europe, southeastern Europe,
such as the Early Oligocene flora of Häring in Tyrol (Etting-
shausen 1853), Tard Clay floras (Hably 2006) and the
Eocene—Early Oligocene floras of Serbia and Macedonia (Mi-
hajlovič & Ljubotenski 1994), often indicate a “subxerophyt-
ic” character of the vegetation which is poorly manifested in
the Csordakút assemblage. The Eocene-Early Oligocene floras
of Serbia and Macedonia display the “subxerophytic” charac-
ter with Zizyphus zizyphoides and leaflets of Leguminosae, as
in the Late Eocene flora of Ovče Polje (Macedonia; Mihajlov-
ič & Ljubotenski 1994). Additional shared elements are
Comptonia, Daphnogene and Eotrigonobalanus. A shared
feature of our flora and the coeval Girbou assemblage in Ro-
mania (Petrescu et al. 1976) is the absence of gymnosperms.
The coeval floras from Central/Western Europe are indica-
tive of a lower degree of floristic similarity which is also
manifested in their generally non-xerophytic character. The
Late Eocene flora of Staré Sedlo in Bohemia (Knobloch et
al. 1996) or the Eocene floras of the Weisselster-Becken,
Messel and Geiseltal in Germany (Rüffle 1976; Rüffle et al.
1976; Mai & Walther 1978; Wilde 1989) share Daphnogene,
Comptonia, Eotrigonobalanus (missing in Messel) and Smi-
lax with the Csordakút flora, though these were not proved
by epidermal structures in the Hungarian Eocene.
In this context, Sloanea nimrodi present in Csordakút also
occurs in the Eocene flora of Kučlin (Bohemia).
The Middle Eocene (Lutetian) fossil plant assemblage
from Csordakút (N Hungary, Gerecse Mountains) has been
investigated. Fossil remains, mostly leaves and rarely fruits
are preserved as impressions in lacustric facies, in freshwater
limestones. Owing to the often fragmented state of leaves
and absence of organic matter the systematic determination
of a great number of specimens was hindered. Algae are rep-
resented by abundant remains of Characeae, including both
vegetative fragments and gyrogonites. Remains of an-
giosperms include Lauraceae (Daphnogene sp.), Fagaceae
(cf. Eotrigonobalanus furcinervis), Ulmaceae (Cedrelosper-
mum div. sp.), Myricaceae (Myrica sp., Comptonia div. sp.),
Leguminosae (leaves and fruit), Rhamnaceae (?Zizyphus
zizyphoides), Elaeocarpaceae (Sloanea nimrodi, Sloanea sp.
fruit), Smilacaceae (Smilax div. sp.). Taxa of both zonal
(Lauraceae, Sloanea, Eotrigonobalanus and Zizyphus) and
intrazonal (Characeae, Myrica, Comptonia and Eotrigo-
nobalanus?) associations are recognizable. Open habitats are
referred to by the occurrence of Cedrelospermum and Legu-
minosae. The “subxerophytic” character of the flora un-
equivocal in the Early Oligocene Tard Clay assemblages and
Early Paleogene floras of Serbia/Macedonia is indefinitely
supported by a few pieces of evidence, namely the occur-
rence of small-leaved Leguminosae, ?Zizyphus zizyphoides
and coriaceous leaves with distinct teeth on the margin (e.g.
Berberidaceae). Nevertheless, winged fruits are definitely
subordinate in the assemblage. The absence of gymnosperms
indicates a floristic relationship with the coeval floras of
Tatabánya (extremely few remains of gymnosperms, N Hun-
gary) and Girbou in Romania. Sloanea nimrodi, a new ele-
ment for the Hungarian fossil record may reflect floristic
relation to the Early Paleogene flora of Kučlin (Bohemia).
ERDEI and RÁKOSI
Acknowledgments: The authors are grateful to Lilla Hably
(Hungarian Natural History Museum, Budapest) for her use-
ful suggestions, scientific discussions and advice related to
the manusript and to Johanna Eder and Zlatko Kvaček for re-
viewing the manuscript. The study was supported by the
Hungarian Scientific Research Fund (OTKA, T037200) and
Bolyai Research Fellowship for the first author.
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