GeolCarp_Vol60_No1_43

GEOLOGICA CARPATHICA, FEBRUARY 2009, 60, 1, 43—57 doi: 10.2478/v10096-009-0005-4

www.geologicacarpathica.sk

Introduction

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
Hungary.

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)

BOGLÁRKA ERDEI

and LÁSZLÓ RÁKOSI

Hungarian Natural History Museum, Botanical Department, P.O.Box 222, H-1476 Budapest, Hungary; erdei@bot.nhmus.hu

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

Geology

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-
seum, Budapest.

Systematics

Charophyta

Characeae

Chara sp.

Fig. 3.1

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.

Angiospermae

Dicotyledoneae

Lauraceae

Daphnogene sp.

Fig. 3.2,3

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.

Fig. 3.4

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
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 : Leaf morphology,

entire margin and type of venation suggest leaves of Lauraceae.

Fagaceae

cf. Eotrigonobalanus furcinervis

(Rossmässler) Walther et Kvaček

Figs. 3.5,6,7

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).

Ulmaceae

Cedrelospermum flichei (Saporta) Hably et Thiébaut

Fig. 3.8,9,10,11

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;

?BP.2001.803.1.

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.

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

Fig. 4.1,2

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-
nounced teeth.

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.

Myricaceae

cf. Myrica longifolia Unger

Fig. 4.3

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”.

Myrica sp.

Fig. 4.4,5

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

Fig. 4.6

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

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

Fig. 4.7,8,9

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;
?BP.2001.808.1.

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

Leguminosae gen. et sp.

Fig. 4.10

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.

Leguminocarpon sp.

Fig. 4.11

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.

Fig. 5.1

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.

Rhamnaceae

?Zizyphus zizyphoides (Unger) Weyland

Fig. 5.2

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).

?Zizyphus sp.

Fig. 5.3

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.

Elaeocarpaceae

Sloanea nimrodi (Ettingshausen) Kvaček et Hably

Fig. 5.4,5,6,7

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)

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)

Fig. 5.4

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.

Fig. 5.8

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
of Myricaceae.

Dicotylophyllum sp. 1

Fig. 5.9

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
obscure.

Dicotylophyllum sp. 2 (?Juglandaceae)

Fig. 6.1

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

Fig. 6.2,3

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.

ERDEI and RÁKOSI

Monocotyledoneae

Smilacaceae

Smilax sp. 1

Fig. 6.5

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).

?Smilax sp.

Fig. 6.6

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-
resent Smilax.

Discussion

Taphonomy

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
made.

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).

Conclusions

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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