ANNALES HISTORICO-NATURALES MUSEI NATIONALIS HUNGARICI
Tomus 78. Budapest, 1986 p. 31-40-
The macroflora of the borehole Kiscell-1 in Budapest
by L. HABLY, Budapest
Abstract — The macroflora of the borehole Kiscell-1 has been studied. It became clear during the
complex geological investigation that the age of the flora is Late Oligocène and belongs to the
NP-23 nannoplankton zone. The flora is typically palaeotropical. Most of the species are thermophilous
elements and a lot of them show xerophilous characteristics. With 1 figure and 3 photoplates.
The age of the flora
Analysis of several exposures and cores were initiated in Hungary durind the investiga
tion of the Eocene-Oligocene boundary problem (BÁLDI, HORVÁTH, NAGYMAROSY & VARGA
1984). The analysis was based on the results of biostratigraphy (nannoplanktons, dino-
flagellates, pollens, macroflora, foraminiferids, large foraminiferids, ostracods, molluscs),
magnetostratigraphy, sedimentology (SEM-records, mineralogy, gcochcmical O and C
isotopic and sedimentological analyses) and the K-Ar method of radiometrical datind.
Among the numerous exposures and cores only the borehole Kiscell-1 contained some
macroflora. The geological section can be seen in Fig. 1. The last presence of Discoaster
saipanensis and D. barbadiensis and the abundance of Zygrhablitus bijugatus, Lanternitus
minutus at a depth of 28 m indicates the boundary of the NP 20/21 zone. The planctonic
foraminiferids, that is, the disappearance of Subbotina linaperta and the rare appearance
of ,,small globigerinoides" indicate the Eocene-Oligocene boundary at the same depth.
The boundary of the NP 21/22 zone is at 78 m and that of the NP 22/23 zone is at 55.5 m.
The transitional planktonic foraminiferid zone can be estimated as between 98-84 m.
The P 18 zone is marked with the abundance of Globigerina linerovskae, Globorotalia post-
cretacea, Globorotalia mundo and Globorotalia brevispira specimens. There is practically
no foraminiferid fauna above 55 m.
However, macroflora appears just above this level, the first leaf remains were found
at a depth of 48.1 m. It seems probable that this is in connection with the petrological change
that occurs at about 51.3 m where slightly laminated Tard Clay is replaced by strongly la
minated Tard Clay.
Thus we can assume the NP 23 nannoplankton zone to be probable for the age of the
flora. This zone contains the euxin layers of the Upper Tard Clay.
Relatively well-preserved macroflora could be found in borehole Kiscell-1. Most of the remains
are leaf fragments, there is only one fruit and there are some flower imprints. Not only the imprints
of the leaves have been preserved but also their texture, so they can also be used in the analysis of the
ABI ETA CE AH
Pinus sp. (PI. II: Fig. 4)
Two needle fragments — both of them double needles — were found between 34.7-48.4 m
in the core. Their entire length cannot be measured but their maximum length is 5 cm. Such double
needles are not rare in Tard Clay Formation. They were found in boreholes marked H and in brick
factories in Óbuda and in Kiseged at Eger.
Since cone remains were not found in borehole Kiscell-1 the precise identification of the needles
is not possible. As the genus Pinus is ecologically widespread today, its climatological value is small
and rather it indicates altitude above sea level.
Doliostrobus hungaricus (RÁSKY) BŰZEK, HOLY & KVAÖEK
(Pl. II: Fig. 3)
Doliostrobus is abundant in Tard Clay. On the basis of this the Doliostrobus level can be marked
out in the Late Oligocène (Hably 1979). This correlates well with the data from the Czech Mountains
where this genus also marks the Late and the Middle Oligocène. There were several sprout remains
but no cone scales were found in borehole Kiscell-1. RÁSKY (1943) mentioned it earlier under the
names of Araucaria hungarica and Sequoia sternbergii from the "Kiscell Clay". NOVAK 1950 also
published it under the name of Sequoia sternbergii from the flora of Kiseged.
Platanus neptuni (ETT.) BÛZEK, HOLY & KVAÖEK
(Pl. II: Fig. 1)
This species was first described as identified from its epidermis in Tard Clay (HABLY 1979, 1980).
Some apical and basal fragments and better preserved fragmented leaf were found in borehole Kiscell-1.
The leaf is about 8 cm long and about 2.2 cm wide so it is of a smallish medium size, similarly
to the Platanus neptuni leaves found in the boreholes marked H, in Tard Clay. The leaf is oblonged
ovate, the apex is missing. Its margin is entire at the base and from one third of the lamina upwards
is specifically toothed. As the camptodromous venation is entirely covered by remains of organic
matter, only the main vein can be observed.
Its earliest appearance in the Eocene is known from Haselbach, Germany (MAI & WALTHER
1978). It is often present in the Oligocène of Europe and can also be found in the Carpatian stage
of the Miocene. Several Oligocène localities have been found both in the Late and Early Oligocène in
Trans-Danubia and in the northern part of Hungary. Its latest and most abundant locality in Hungary
is in the Late Miocene of Ipolytarnóc (HABLY 1985a). This species seems to be predominant in the
Oligocène. Its shape in the Egerian is substantially different from that in the Kiscellian. In the Kiscellian
it became smaller in size indicating that it became acclimatized to aridity. Almost all the
species have large leaves in the Egerian, therefore we can conclude that their ecological diversification
took place not in space but in time.
Platanus neptuni is a paleotropical element. KNOBLOCH (1973) thinks that its presence is connect
with warm periods. This plant lived relatively far from water-sides, mainly on hilly or slopy regions.
It was a thermophilous and xerophilous plant in the Kiscellian. Due to the changing conditions it lost
its xerophilous characteristics in the Egerian. Since it is very frequent and is almost always present
in the regions under investigation, we can assume that Platanus neptuni had been a basic element in
the association of Tard Clay.
Dryophyllum furcinerve (Ross.) SCHMALH.
It is predominant in the flora of Tard Clay. It was largely found in boreholes marked H at
Óbuda and they are also frequent in Kiseged. As the shape of the lamina varies enormously, it has a
large number of synonyms. In Hungary KOVÁCS (1958) reported it under the name of Castanopsis
furcinervis from the Eocene locality at Lábatlan. ANDREÁNSZKY also published it under the name of
Castanopsis furcinervis from several Oligocène localities eg. Kiseged at Eger, Csillaghegy in Budapest,
Novaj—Vécsey valley, the sorroundings of Nagyimány and the Wind brickyard in Eger (RÜFFLE et al.
1967). Considering all the European localities of this species we can observe that the lamina of the
3 Term. Tud. Muz. Évk. 1986.
plant in the Late Oligocène of Hungary is smaller than that anywhere else. Both ANDREÁNSZKY and
some German authors (RÜFFLE et al. 1967) assume that this difference is due to the presence of an
arid-atlantic climate. This was supported by the analysis of the materials from boreholes maiked
H and from borehole Kiscell-1.
Two specimens of D. furcinerve were found at a depth of 17.4 and 19.3 in the borehole. The
findings are small leaf-fragments with some well preserved morphological marks on them. The
structure of the epidermis can also be easily analysed.
Palaeocarya orsbergensis (WESS. et WEB.) JÄHNICHEN, WALTHER et TAKAC
(Pl. III: Figs 1-4)
No large leaf fragments or fruit remains were found in the borehole, only a piece of lamina
(1 cm 2
), the taxonomical identity of which could only be defined by the analysis of the epidermis.
The character of its texture is quite different from those already published (KVACEK 1972, KNOBLOCH
& KVACEK 1976, JÄHNICHEN et al. 1977).
Similarly to the specimens found in the localities of the Tertiary of Europe, it has very characteristic
large umbrella-like glands. These are all strongly cutinized, they have round shapes and
slightly waved outlines. There are basic differences in the outlines of the cells and their rate of cutinization.
The cell wall of this specimen is usually slightly cutinized and wavy. The polygonal cells of the
epidermis of P. orsbergensis found in borehole Kiscell-1 have heavily cutinized, straight cell walls.
JÄHNICHEN (1977) mentioned a remain from the locality of Haselbach-Lucka where the cell outlines
are straight and heavily cutinized at the margin of the leaf. Therefore it seems certain that our remain
also belongs to the margin area of the lamina.
Stomata are frequent on the lower part of the epidermis and are positioned asymmetrically.
The structure of the stomata is anomocytic, the pores are usually narrow and the guard cells have a
kidney-shaped figure. Both the upper and lower parts of the epidermis are heavily cutinized.
This species has been described under several names in Hungary. The species of the same age
from Kiseged is referred to Schinus oligocaenicum by ANDREÁNSZKY & NOVAK (1957) and by ANDRE
ÁNSZKY (1963). It was mentioned under name Myrica lignitum (RÁSKY 1959) from the Ottnangian
flora of Ipolytarnóc and Schinus molle (ANDREÁNSZKY 1959) from the sarmatian flora of Erdőbénye.
Localities of this species are known from the Miocene in Hungary as well as in other parts of Europe.
As regards its specific marks, we can arrive at interesting conclusions concerning its ecology and
climatology. ANDREÁNSZKY (1963) allocated it to the genus Schinus and beleived it to be a xerophilous
element. Though the morphology of the leaf suggests that it is a xerophilous genus, according to
KVACEK the slightly cutinized epidermis and the wavy cell wall are inconsistent with that theory.
The recent Engelhardtia live in subtropical-tropical rain forests mainly in South-East Asia, Malaysia
and Eastern Himalayas. Concerning the localities of this species in Europe, JÄHNICHEN also concluded
that it could have formed the association of the ancient subtropical-tropical rain forests. However,
its surprisingly small leaves in some floras and its simultaneous occurrence with the Myrtaceae and
Leguminosae indicate that it was at least partly meso-xerophilous.
On the basis of both the plant communities and the morphological or anatomical marks, we
classify this Kiscellian species as a xerophilous element. This is supported by the straight, heavily
cutinized cell-walls, the small leaves of the Kiseged flora and the xerophilous associations found in
borehole Kiscell-1. It seems therefore, that P. orsbergensis was euryecious within wide boundaries.
On the basis of this we could even presume that it was eurythermous, but the detailed analysis of its
localities show that it always appeared in warm periods and in thermophilous associations. As regards
water requirements, it was also euryecious since it is present in both the tropical and meso-xerophilous
Dalbergia beila HEER (Pl. I: Fig. 1)
Only a small leaf remain of this species was found at a depth of 31.5 m in the core. It is 2.3 cm
long and 1 cm wide. The shape of the lamina is ovate and slightly asymmetrical, its base is acute and
the margin is entire, the apex is chipped out. The main veins are stout. Other parts of the venation
cannot be seen because of the presence of organic matters.
This plant is common in the localities of Tard Clay around Budapest, for example, in borehole
marked H (HABLY 1979) and in the exposures of Szépvölgy and Csillaghegy in Budapest (RÁSKY
1943). Similarly small remains also occurred in boreholes marked H, where the largest remain was
4.5 cm long ang 2 cm wide, while one of the D. bella remains described by RÁSKY was 7.5 cm long
and 4 cm wide.
Cercis hungarica RÁSKY (Pl. II : Fig. 4)
Two specimens appeared from borehole Kiscell-1, one of them together with its ipmprint. The
remains are very fragmented. On one of the remains apical part of a leaf can be observed and on the
other one the middle part of a leaf can be seen with the venation on it. The proper size of the leaves
cannot be measured but on the basis of the space and the angle between the veins a leaf of about
10 cm can be reconstructed. Two pairs of prominant veins branch out of the midrib from a single point
at the base. The lower veins are almost horizontal, that is, they form an angle of 90° with the midrib
while the upper veins enclose an angle of 45°. After a distance of about 3 cm secondary veins brach
out of the midrib from different points, the distance of which is 0.1 cm. Other parts of the venation
are covered by fine, mosaic-like fragments of organic matter in all remains. The margin of the leaf
is entire. The apex at one of the specimen is obtuse.
RÁSKY (1943) described the species from the Late Oligocène of Csillaghegy in Budapest, also
based on leaf fragments. The size of the holotype is larger but their morphological features are the
same. RÁSKY considered it to be closely related to the recent species Cercis canadensis.
Cercis parvifolia LESQU.
A 6 cm long and 4 cm wide specimen occurred at a depth of 30.9 m in borehole Kiscell-1.
The margin of the leaf is entire, the apex is missing. The basal part is very fragmented but its dissimilarities
from the C. hungarica can clearly be observed. While the base of C. hungarica is cordate, no
such projection can be observed for C. parvifolia. Thus the base is nearly acute. Its venation is similar
to that of C. hungarica but it has no horizontal veins running to the projection at the base of the
lamina. Newertheless we can see the predominant veins forming an angle of 45° with the midrib and
the secondary veins branching out of the midrib after larger species. The secondary veins are camptodromous.
The size of the space between the veins is smaller towards the axis. RÁSKY also mentions
it from the Late Oligocène from Csillaghegy, together with C. hungarica. The remains which were
found at Csillaghegy are a bit larger than those under investigation, but the typical specimens are
considerably smaller than the remains from Csillaghegy. Though the epidermis is preserved in the
specimen, it is so thin and fragmented that further preparation is impossible.
Zizyphus zizyphoidcs (TJNG.) WLD. (Pl. I: Fig. 2)
Only one specimen was found at a depth of 35.5 m in the core. It become clear studying the other
localities that it is more common in Tard Clay (HABLY 1979). The entire length of the leaf is 6.5 cm,
its width is 2 cm. The lamina is asymmetrical, the margin is irregularly indented and wavy. The lamina
is oblong and its widest part is somewhat above the medial line. The apex is missing. Two stout
secondary veins appear along the midrib, running somewhat closer to the margin than to the midrib.
Other parts of the venation cannot be observed because of the presence of organic materials.
The species can be found either in the Middle Eocene of Trans-Danubia (HABLY 1985b), or in
several Late Oligocène localities (RÁSKY 1956, HABLY 1979). Accordingly it survived trough the
Eocene-Oligocene boundary, and became extinct only in the Late Oligocène. The plant occurs in the
Late Oligocène of Méra in a similar association than in the Tard Clay.
Abelia quadrialata REID. et CHANDL. (Pl. I: Fig. 3, Pl. II: Fig. 2)
ANDREÁNSZKY (1959) mentions this genus from several localities in Tard Clay, eg. Budaújlak
and Kiseged. REID & CHANDLER (1926) also described it from the Late Oligocène from Bembridge
flora, under the name of Abelia quadrialata. It was found in the boreholes marked H in Óbuda, also
from Late Oligocène (HABLY 1979). Two very well preserved imprints were found at depths of 20.4
and 48.4 m in borehole Kiscell-1. In its main characters the one found at 48.4 m is very similar to the
remains found earlier. It consists of four heavily oblonged, obovate petals, two of them close together,
and the other two are far from each other, thus showing zygomorph symmetry. The maximum width
of the petals is 0.3 cm. On the earlier remains radial symmetry can be observed, moreover, ANDRE-
ÁNSZKY actually considered them to have actinomorph symmetry. In the remains that were found
some 20 m above the others the features of zigomorph symmetry can even better be recognized. This
can be seen not only from the distance between the petioles but also because of their distinctness.
The upper petioles that are most distantly placed are much wider (0.5 cm) than the lower ones (0.3 cm)
and their apex is chipped out. So we can see that there are obvious differences between the two
remains. It is conceivable that these imprints represent different stages of development since there
is a time-gap of about half a million years between the dates of their fossilization, according to the
spead of sedimentation in Tard Clay. Considering this the symmetry of the flower became more and
more zygomorph and their petiols which became wider and had their apex chipped out, subdivided
again, but we have no evidence to support this.
The borehole Kiscell-1 contains a flora typical of Tard Clay. There were not enough
remains in the core so that we could find all the characteristic species, but we found some
genera that were not present in the borehole marked H, and the most common and charac
teristic species were present. Though, unlike the boreholes H the few remains of borehole
Kiscell-1 are not sufficient for giving an overall description of the flora in the Tard Clay
Formation, valuable information about the range and characteristics of the Kiscellian flora
can be extracted.
Ecological and palaeoclimatological evaluation
No quantitative information can be gained about the predominant species from the
few remains. Nevertheless, the species that are considered to be predominant on the basis
of the flora from Tard Clay are also represented here. The occurrence of Platanus neptuni
makes it certain that it was a very common and predominant element in the Late Oligocène
of Hungary. The presence of Palaeocarya orsbergensis that could be identified by its epider
mis, as only some of its larger leaffragments have been preserved, is very interesting. The
presence of the flower imprints of Abelia quadrialata in Tard Clay is not much interest, as
it occurs in every such locality. Flower imprints in the floras of other ages are very rare
either in Hungary or in other parts of the world.
The flora in borehole Kiscell-1 is paleotropical. Only an uncertain fruit of Acer can
be mentioned from among the Arcrotertiary elements. The climate can be characterized
very well on the basis of the flora. Most of the species are thermophilous or meso-thermo-
philous, that is, subtropical elements. Some of the remains have xerophilous features. The
conclusion that the climate changed for less moist is supported by the presence of xerophilous
features even on non-xerophilous species. This can be seen either in their morphological
structure or in their texture, like eg. on Palaeocarya orsbergensis.
LIST OF THE SPECIES
(see also Fig. 1)
Platanus neptuni 35.5 m
ANDREÁNSZKY, G. (1954): Ősnövénytan. — Akadémiai Kiadó, Budapest: 320 p.
ANDREÁNSZKY, G. & NÓVÁK, E. (1957) : Neue und interessante tertiäre Pflanzenarten aus Ungarn III.
— Annls hist.-nat. Mus. natn. hung. (ser. n.) 8: 43-55.
ANDREÁNSZKY, G. (1959): Contributions à la connaissance de la flore de l'oligocène inférieur de la
Hongrie et un essai sur la reconstitution de la flore contemporaine. — Acta bot. hung. 5(1-2): 1-37
ANDREÁNSZKY, G. (1959): Die flora der sarmatischen Stufe in Ungarn. — Akad. Kiadó, Budapest:
ANDREÁNSZKY, G. (1963): Das Trockenelement in der Alttertiären Flora Mitteleuropas auf Grund
paläobotanischer Forschungen in Ungarn. — Vegetáció Acta Geobot. 11 (3): 95-111.
BÁLDi, T., HORVÁTH, M., NAGYMAROSY, A. & VARGA, P. (1984): The Eocene-Oligocene boundary in
Hungary. The Kiscellian stage. — Acta geol. hung. 27 (1-2): 41-65.
HABLY, L. (1979) : Some Data to the Oligocène Flora of the Kiscellian Tard Clay, Hungary. — Annls
hist.-nat. Mus. natn. hung. 71: 33-53.
HABLY, L. (1980): Platanus neptuni (Ett.)BûZEK, HOLY & KVACEK in the Hungarian Oligocène.—
Acta bot. hung. 26 (3-4): 299-316.
HABLY, L. (1985a) : Early Miocene plant fossils from Ipoly tarnóc, N Hungary. — Geol Hung. ser. Pal.
HABLY, L. (1985b): New data to the Eocene flora of Hungary. — Annls hist.-nat. Mus. natn. hung.
JÄHNICHEN et al. (1977): Blätter und Früchte von Engelhardtia Lesch. ex Bl. (Juglandaceae) aus dem
europäischen Tertiär. — Feddles Reperotorium 88 (5-6): 323-363.
KNOBLOCH, E. (1973): Eine bemerkenswerte Platane des europäischen Tertiärs. — Der Aufschluss
KNOBLOCH, E. & KVACEK, Z. (1976) : Miozäne Blätterfloren vom Westrand der Böhmischen Masse. —
Rozpravy ÚŰG 42: 1-131.
KOVÁCS, É. (1958) : Középső-eocén flóra Lábatlanról. — M. Áll. Földt. Int. évi jel. 1957-58. p. 473-495
KVAÖEK, Z. (1972): Engelhardtia-leaves in the European Tertiary. — Casopis miner, geol. roc. 17
MAI, D. & WALTHER, H. (1978): Die Floren der Haselbacher Serie in Weisselster-Becken (Bezirk
Leipzig, DDR). — Abh. Staatl. Mus. Mineral. Geol. Dresden 28: 1-101.
NÓVÁK, E. (1950): A kisegedi oligocén flóra fenyőféléi. — Bud. Tud. egy. Biol. Int. évk. 1(1): 48-61.
RÁSKY, K. (1943) : Die oligozäne Flora des Kisceller Tons in der Umgebung von Budapest. — Földt.
közi. 73: 503-610.
RÁSKY, K. (1956): Fosszilis növények a Budapest környéki „budai" márga összlctből. — Földt. közi.
RÁSKY, K. (1959) : The fossil flora of Ipolytarnóc. — Joiirn. Paleontology 33 (3) : 453-461.
REID, E. M. & CHANDLER, M. E. J. (1926): The Bembridge Flora. — Catalogue of Cainoz. Plants. I.
British Mus. London : 206 p.
RÜFFLE et al. (1967): Eozäne Floren des Geiseltales. — Abh. Zentr. Geol. Inst. 26: 200-282.
Author's address: DR. LILLA HABLY
Hungarian Natural History Museum
Budapest, pf. 222
Annls hist.-nat. Mus. natn. hung., 78, 1986
rialata (81.3.2.) X4
hungaricus (81.5.3.) Xl.3 — 4 = Cercis hungarica (81.14.2.) Xl.3; Pinus sp. (81.14.2.) Xl.3
Palaeocaryas orbergensis (81.8.2.): 1, 2, 3 — upper epidermis, 4=1 ower epidermis