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A N N A L E S H I S T O R I C O - N A T U R A L E S M U S E I N
A T I O N A L I S H U N G A R I C I Tomus 87. Budapest , 1995 p.
11-33
The Sarmatian flora from Erdőbénye-Ligetmajor, NE Hungary
by B. ERDEI , Budapest
ERDEI, B. (1995): The Sarmatian flora from Erdőbénye-Ligetmajor,
NE Hungary. - Annls. hist.-nat. Mus. natu. hung. 87: 11-33.
Abstract - The Sarmatian flora of Ligetmajor was studied. The
floristical analysis was followed by a floristical and
palaeoclimatological reconstruction. Since mainly Mediterranean and
xcrophytic elements form the flora the climate must have been
Mediterranean with dry and hot summer and mild winter. Finally the
flora of Ligetmajor was compared with the classical floras of
Erdőbénye (Barnamáj and Kővágó-oldal). Although the floras of
Ligetmajor and Kővágó-oldal are quite similar, some interesting
differences exist between the two floras. With 22 figures and 4
tables.
I N T R O D U C T I O N
I have been studying the Sarmatian flora of the siliceous earth
mine in Erdőbénye-Ligetmajor since 1992 (Figs 1-2). Geologically
there are formations of the middle and upper Sarmatian and
Pannonian namely hydroquartzite, diatom, silicified argillaceous
silt, rhyolitic tuff and andésite (Fig. 3).
I have collected more than 700 prints of 36 plant species. The
private collection is deposited in Debrecen in the eastern part of
Hungary. I have compared the palaeoflora of Ligetmajor with two
other floras of Erdőbénye, notably Barnamáj and Kővágó-oldal. After
a detailed analysis I attempted to reconstruct the palaeogeography
and palaeociimatology of the area in Ligetmajor.
The Tokaj Mountains form a part of the Tokaj-Eperjes Mountains
(extending from Eperjes to Tokaj) in the northeastern region of
Hungary.
The siliceous earth of Ligetmajor can be found in a tectonic
subsidence bounded by pyroxene andésite and siliceous earth stones.
From the north and northeast the area is bounded by the Sarmatian
rhyolitic and volcanic centre of a hill called Erdőhorváti-Szokolya
(the age of its olivine-pyroxene andésite according to a total K/Ar
dating examination is 10.9+0.5 mil l ion years; PÉCS KAY 1983).
From the south and from the east the region is closed by the
pyroxene andésite stones of a hill called Szár
Erdőhorváti-Szokolya. The tectonic subsidence is filled with
rhyolitic tuff and tufite. Pyroxene andésite laccolith has
penetrated into these stones. Because of the erosion some of them
have come to the surface such as Barnamáj and Mulató H i l l .
Two other localities of Sarmatian plant fossils have been
recorded in Erdőbénye for a long time (KoVATS 1856a, A N D R E Á N
S Z K Y 1959).
In Barnamáj , in the southern part of the village plant fossils
are in a fine, grey clay layer which was lifted up by an andésite
laccolith. Unfortunately it has been worked out totally.
One-and-a-half to two km northwest of Barnamáj a new locality
was opened up in Kővágó-oldal. White rhyolitic tuff contains the
fossils in this region. The floras of Barnamáj and Kővágó-oldal are
so similar that their age is considered to be the same. H A J Ó S
& P Á L F A L V Y (1964) described plant fossils from the
siliceous earth mine in Ligetmajor. The bedding stone is clayey
silt which was soaked secondarily wi th silica. Presumably silicic
acid originated from the upper rhyolitic tuff and diatomaceous
earth by solution and migrated into the clayey silt.
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Figs 1-2. Fig. 1. Map of Hungary (locality =
Erdőbénye-Ligetmajor). - Fig 2 . Geological map of the Erdőbénye
basin
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SYSTEMATIC PART
S P H E N O P S I D A
There is a small (4.5 cm) fragment of a stem available for us.
The verticillate leaves are clearly recognizable. However, a more
precise taxonomical ranging is impossible.
M a t e r i a l : No. 531.
G Y M N O S P E R M A T O P H Y T A
ABIETACEAE
The order is represented by pine needles and cataphylls of
cones. The leaves are five needles and they presumably belong to
the group of Pinus paleostrobus (ETTINGSHAUSEN) HEER (ANDREÁNSZKY
1959).
M a t e r i a 1: No. 548, 576-579.
C U P R E S S A C E A E
Cupressus cf. sempervirens L . (Fig. 4)
1959 Cupressus cfr. sempervirens L.: ANDREÁNSZKY, p. 56, Tafel
10, Fig. 9.
We have found a small fragment of a branch characterized by
decussate cataphylls. These are short, obtuse and
rhombus-shaped.
M a t e r i a l : No. 37c.
Fig. 3. Geological profile of the Erdőbénye basin
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A N G I O S P E R M A T O P H Y T A
ULMACEAE
Ccltisjapeti U N G E R 1 8 5 2
1959 Celtis japeti UNG.: ANDREÁNSZKY, p. 135. 1987 Celtis japeti
UNGER: PALAMAREV & PETKOVA, p. 58, Tab. 15, Fig. 2.
One leaf fragment with petiole. Lamina asymmetrical. Shape
narrow ovate. Lamina length 3.1 cm, width 1.6 cm. Apex is attenuate
and base is asymmetrical rounded but approaching the midvein it
becomes decurrent. Margin cannot be observed. Venation
semicraspedodromous or brochidodromous (it is not determinable).
Midvein weak and straight. A pair of basal veins nearly as thick as
the midvein arises from the base. A vein seems to branch off
basally from the latter veins towards the margin. The angle of
divergence of the secondaries is acute (45°-60°-70°). Toward the
apex they are getting thinner. Their course is curved and they do
not reach the margin. Tertiary veins arise at angles of about 60°
and the neighbouring tertiaries anastomose with each other at
obtuse angles.
M a t e r i a 1: No. 271.
Ulmus plurinervia U N G E R 1843
1856 Ulmus plurinervia UNG.: KOVÁTS, p. 26, Tab. 4, Figs 6-7.
1959 Ulmus plurinervia UNG : ANDREÁNSZKY, p. 129, Tafel 34, Fig. 4,
Tafel 36, Figs 5, 7.
Only one leaf was found. It is badly preserved and the petiole
is hardly observable. Lamina symmetrical, shape narrow ovate.
Lamina length 3.4 cm, width 2.0 cm. Apex acute, base cordate. On
the margin there are compound teeth. Under the main teeth small
secondary teeth are also observable. The apical and basal sides of
the teeth are convex. Al l the teeth are attenuate. Venation
craspedodromous. The midvein is straight and the secondaries arise
at acute angle. Approaching the margin they bifurcate and the lower
ones run into the secondary teeth. Due to the fossil's bad
condition the tertiary venation is not observable.
M a t e r i a 1: No. 69b.
Ulmus sp.
One leaf fragment. Only the lower part of the leaf has remained.
Lamina length 6.0 cm, width 2.4 cm. Shape must have been narrow
elliptic. Base asymmetrical and obtuse. The margin is toothed.
Because of the bad condition of the leaf the teeth cannot be
described precisely. Presumably they are compound teeth. Venation
craspedodromous. The course of the midvein is straight. The angle
of divergence of the secondaries is acute. The veins are straight
and run into the teeth. The tertiary venation is extremely thin and
its pattern is orthogonal reticulate.
M a t e r i a l : No. 82
Zelkova zelkovifolia ( U N G E R 1843) BÔ/TiK et KOTLÁBA in
KOTLÁBA 1963 (Figs 13, 14)
1856 Zelkova ungeri Kov.: KOVÁTS. p. 27, Tab. 5, Figs 1-12. 1959
Zelkova ungeri Kov.: ANDREÁNSZKY, p. 133, Tafel 37, Fig. 5, Tafel
39, Fig. 6, Tafel 41, Fig. 2. 1971 Zelkova zelkovaefolia (UNGER)
BUZEK et KOTLÁBA in KOTLÁBA: BUZEK, p. 58, Pl. 21, Figs 8-9, Pl.
22, Figs 4-14. 1991 Zelkova zelkovaefolia (UNGER) BÜÍEK et KOTLÁBA:
FISCHER & H ABLY. p. 29. Pl. 2 Figs 1-3, 5, Figs 19, 23.
The species is represented with a great number of leaves which
have petiole. Lamina asymmetrical. Shape lanceolate. Lamina length
1-7.4 cm, width 0.6-3.3 cm. Apex attenuate, base obtuse. Margin
regularly and simply toothed.
The teeth are acute. The apical side of them is acuminate, the
basal side is convex. The sinuses between the teeth are angular.
Venation is craspedodromous and the midvein is straight.
Secondaries arise at acute angle (50°-60°) but in the upper pail of
the leaf the angle of divergence is growing. Their course is curved
and they run into the teeth. Smaller veins are observable, arising
from several secondary veins toward the margin and running into the
sinuses under the tooth that belongs to the secondary vein. The
tertiary venation is extremely thin. They arise at right angle and
several of them bifurcate and anastomose with the neighbouring
tertiary vein. The quaternary venation is orthogonal
reticulate.
M a t e r i a l : No. 1-11, 12a, c, 13. 14, 15a, 16, 17, 19a,
20-23a, 25-33a, 34, 36, 37a, b, 38a, 63b, 66, 87, 88, 112b, 128b,
161b, 167b, 170c, 173a, 192a, 206d, e, 229c, 252b, c, 257c, 264c,
273b, 275c, 291, 293b, 299a, 302c,
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d, 303-308a, 309a, 310a, 311-313, 315. 317b, 318b, 320-323,
324b-327a, 328a, 329b-332, 333b, 334a, 335b, 337-341a, 342a, 344,
345a, 346a, 347a, 348, 349a, 351a, 353. 358a, 380b, c, 381c, 390a,
405b, 420b, 429c, 455d, 462b, 465c, 467c, 480b, 488d, 495a,
498-506, 508, 509b, 5 L1 a, 512513a, 514-516, 517b, 526a, 528b,
530, 542, 552, 553.
FABACEAE
Podogonium knorrii (A. B R A U N 1851 ) H E E R 1859 (leaflet)
(Figs 10, 11)
1856 Copaifera longestipitata Kov.: KOVÁTS, p. 51, Tab. 1, Figs
3-4. 1959 Podogonium knorrii (A. BR.) HEER: ANDREÁNSZKY. p. 146,
Tafel 43, Fig. 6, Tafel 44, Fig. 8, Tafel 46, Fig. 3. 1992
Podogonium knorrii (BRAUN) HEER: HERENDEEN, p. 4, Figs 1-5.
Leaflets have turned up in a great number. The leaflets are
symmetrical, only their base is asymmetrical, showing that this
species had compound leaves. The leaflets have no petiole and their
shape is narrow oblong. Lamina length 0.7-3.7 cm, width 0.4-1.3 cm.
Apex and base are obtuse (some specimens have acute base). Margin
is entire. Venation camptodromous, presumably brochidodromous. The
midvein is thin and its course is straight. The secondaries arise
at acute angle. A thick, basal vein, being typical, is observable
on one side of the leaflets. The tertiary venation is so thin that
it cannot be described.
M a t e r i a 1: No. 15b, 33b, 72b, 99b, 100a, c, 111, 112a,
113, 114, 116-119, 122-128a, 129a, 135, 160b, 170b, d. 173d, 190b,
198b, 206b. 211, 214, 217-220, 223a, 229a, 231-235a, c, e, 236-238,
243-246, 249-250a, 251a, 253a, 254a, b, 255-256a, 257a, 261-264b,
265, 266, 273d, 275a, 279b, 285a, 30la, 302b, 308b, 310b, 333c,
342b, 345b, c, d, 351b, 352b, 364a, c, 367b, 390c, 411b, 420c, d,
428b, 453, 454, 455b, c, 456-460b, 461, 463-465a, 466, 467b, 468,
472-474, 476, 477. 479, 488b, 490b, 494b, 513b, c, 529c, 533b.
536b, 551a, 556, 558b.
Podogonium knorrii (A. BRAUN 1851) HEER 1859 (fruit)
(Fig. 12)
1992 Podogonium knorrii (BRAUN) HEER: HERENDEEN, p. 4, Figs
1-5.
Wc have collected 42 legumes of Podogonium knorrii. Their length
is 2-2.5 cm. They have a petiole which is almost as long as the
fruit itself. In some cases the seed is also observable as it is
falling out from the fruit.
M a t e r i a I: No. 12b, 39-51, 53-59, 21 Od, 224-228, 229b,
247b, 254c, 258b, 259-260, 267, 278c, 389b, 465b, 467a, 469-470a,
475, 478a, 529d, 547, 550a.
Lcguminosae gen. et sp.
We have 24 leaves in this group. They are symmetrical with small
petiole. Shape is wide elliptic but some of them is orbiculate.
Lamina length 1.2-3.8 cm, width 0.7-1.9 cm. Apex and base are
obtuse or rounded. Margin is entire. On the base of the No. 241
specimen venation is camptodromous, brochidodromous (they are badly
preserved). The midvein is relatively thick and straight. The angle
of divergence of the secondaries is acute. Their course is curved
and approaching the margin they anastomose with the neighbouring
secondaries at right angle and form loops. Tertiary venation is not
observable.
M a t e r i a 1: No. 19b, 131a, 141. 142, 145, 146, 148, 212,
213, 215. 221, 222b. 239, 241, 242, 274b, 298b, 301b, 335c, d,
346c, 359, 364b, 397b, 431b, 470
"Dalbergia bella" H E E R 1859
1969 Dalbergia bella HEER: K N O B L O C H , p. 30, Taf. 9, Figs
8, 8a, 9, 9a, Taf. I I , Figs 4, 4a.
Only some specimens represent the species. They have small
petiole. The leaves seem to be a bit asymmetrical; however, the
base is symmetrical. Lamina length 1.9-2.5 cm, width 0.8-1.2 cm.
Shape obovate, apex rounded, in some cases emarginate. Base acute
and decurrent. Margin entire. Venation cannot be observed precisely
but presumably it is brochidodromous or semicraspedodromous.
M a t e r i a l : No. 136, 240b, 329a, 521b.
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BETULACEAE
Betulaceae gen. et sp.
One badly preserved leaf. It is symmetrical, petiole is not
noticeable (it has not remained). Shape elliptic. Lamina length 5.7
cm, width 3.4 cm. Apex has not remained but presumably it was
acute. Base is obtuse and seems to be a bit cordate. Margin is
toothed. The teeth are compound. Under the main teeth smaller teeth
are also visible. Venation is craspedodromous with a straight
midvein. Secondary venation arises at an angle of about 45° -50°
and its course is straight. The tertiary venation is not
observable.
M a t e r i a l : No. 258a.
Carpinus grandis U N G E R 1 8 5 0 sensu H E E R 1 8 5 6
1959 Carpinus grandis UNGER: ANDREÁNSZKY, p. 89, Tafel 21, Figs
6-7, Tafel 22, Figs 3, 5-6. 1971 Carpinus grandis UNG. sensu HEER:
B8ZEK, p. 50, Pl. 16, Figs 1-3.
We have lots of specimens of this species. The leaves aie
symmetrical and have petiole. Lamina length 1.7-6 cm, width 1 -3.2
cm. Shape narrow ovate, apex acute, base obtuse or rounded, in some
cases cordate. Margin is irregularly toothed and the teeth are
compound. It has main and secondary teeth and in some cases a
tertiary tooth also exists. The teeth are weakly attenuate and the
sinuses are angular. Venation is craspedodromous. The midvein is
straight. The angle of divergence of the secondaries is acute and
they run into the main teeth. Their course is straight. Basally
from two or three pairs of secondaries a pair of vein arises and
runs into the teeth. The first pair of them arises at an obtuse
angle. The tertiary venation is very thin, presumably orthogonal
reticulate.
M a t e r i a l : No. 18, 62, 67b, 70a, 71, 76, 85, 86, 93, 98,
108, 250b, 269, 274a, 275b, 276a, 277, 279a 282, 284, 292, 295,
298a, 301c, 356, 357, 363, 413, 421, 429b, 430b, 436b, 518, 520,
540b, 551b.
Carpinus neilreichii KovÁTS 1 8 5 6 (Fig. 8 )
1856 Carpinus neilreichii Kov.: KOVÁTS, p. 23, Tab. 4, Figs 1-3.
1959 Carpinus neilreichii Kov: ANDREÁNSZKY. p. 90, Tafel 22, Figs
7-8.
The species is represented with 9 winged fruits. The fruits are
leaf-like and asymmetrical. They consist of one main part and the
width of it is the largest in its middle region. Margin is toothed,
bigger and smaller teeth are presented. The teeth themselves aie
attenuate and the sinuses are angular. A thick venation arising
from one point of the base is observable.
M a t e r i a 1: No. 137b-140, 210e, 247b, 438c, 446b, 521a.
Carpinus pyramidalis G A U D I N in G A U D I N et STROZZI 1 8 5
8
(Fig. 9 )
1856 Carpinus producta UNG.: KOVÁTS, p. 24, Tab. 4, Fig. 5. 1959
Carpinus pyramidalis (GOEPP.) HF.ER: ANDREÁNSZKY, p. 89, Tafel 21,
Fig. 5.
We have 4 specimens of the other fruit of the Carpinus genus
being present in this flora. It is leaf-like, asymmetrical and
composed of three parts. The middle lobe is much bigger than the
others. Their margin is entire. A thin venation is also
observable.
M a t e r i a l : No. 121, 240a, 289a 360.
FAGACEAE
Fagus haidingieri KOVÁTS 1 8 5 6
1856 Fagus haidingieri Kov: KOVÁTS, p. 24, Tab. 4, Figs 6-7.
1969 Fagus haidingieri Kov.: KNOBLOCH, p. 79, Taf. 36.
Only some specimens have turned up. The leaves are symmetrical
and have relatively short petiole. Lamina length 6.2-7.7 cm, width
2.9-4 cm. Shape narrow ovate, apex presumably acute, base weakly
cordate. Margin is simply toothed. The teeth are acute and
small-sized. They are hardly visible to the naked eye. Their apical
side is concave,
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(he basal is acuminate. The sinuses are rounded. Venation is
craspedodromous. The midvein is weakly curved, its course is
sinuous. The secondary veins arise at an angle of about 45°, but
the angle approaching the base is getting bigger. The secondaries
are relatively thick, their course is straight and they ran into
the teeth. The tertiary venation is extremely thin and random
reticulate.
M a t c r i a 1: No. 109, 278b, 290, 300, 517a.
Quercus kubinyii (KOVÁTS 1 8 5 1 ) BERGER 1 9 5 2 (Figs 5, 6
)
1856 Castanea kubinyii Kov: KOVÁTS, p. 25, Tab. 3, Figs 1-7.
1959 Quercus kubinyii (Kov.) CZECZOTT: ANDREÁNSZKY. p. 106, Tafel
28, Fig. 4. 1991 Quercus kubinyii (KOVÁTS) BERGER: FISCHER &
HABLY, p. 26, PI. 3 Fig. 2, PI. 4 Figs 1-5, PI. 5 Fig. 7. Figs
1, 5, 9, 11.
We have a great number of specimens in the flora. The leaves are
symmetrical and have petiole. Shape lanceolate, lamina length
2.4-11.7 cm, width 1.2-5.0 cm. This species has the largest leaves
in the flora. Apex acute, base rounded, in some cases weakly
cordate or cuneate. Margin is lobale. The lobes are attenuate and
terminate in a long bristle. Their apical and basal sides are
acuminate. The sinuses are rounded. The lobes are missing on the
base. Venation is craspedodromous. The midvein is relatively thick
and straight. The relatively thick secondaries arise at an angle of
about 50°-60°. Their course is straight and they ran into the
teeth. The angle of divergence of the teitiaries is right and they
arise from both of the upper and lower sides of the secondaries.
Their course is straight and they are parallel with each other. The
quaternary venation is extremely fine and orthogonal
reticulate.
M a t e r i a l : No. 19c, 23b, 24a, 38b, 63a, 64, 65, 72a, 89,
90, 92, 131b, 137a, 151, 152, 154, 156, 158, 159, 200b, 206c, 210c,
223d, 235d, 251b, 258c, 272c, 278d, 280, 281, 283, 293a, 301d, 314,
316, 317a, 319, 328c, 341b, 343, 347c, 350, 374-376a, 377, 379,
380a, 381a, 382a, 383a-389a, c, 390b, 391, 392, 395-397a, 398-405a,
406-409, 415-419, 422-428a, 429a, 430a, 431a, 436a, 437, 438a, 441,
443, 445, 446a, 447a, 448a, 449-451a, 452, 484b, 495b, 538-540a,
546.
Quercus pseudocastanea GÖPPERT 1 8 5 2
1959 Quercus pseudocastanea GOEPPERT: ANDREÁNSZKY, p. 116 1986
Quercus pseudocastanea GOEPPERT: KNOBLOCH & VELITZELOS, p. 30,
Tafel 15, Figs 1-2. 1988 Quercus cf. pseudocastanea GOEPPERT:
KNOBLOCH, p. 8, Tafel 13, Figs 1-2. 1991 Quercus pseudocastanea
GOEPPERT: WALTHER & ZASTAWNIAK, p. 169, PI. 2, Figs 2-6.
The species is represented with some specimens. The leaves are
symmetrical, their petiole has not remained. Lamina length 3.8-11.1
cm, width 1.1-5.3 cm. They belong also to the group of the largest
leaves. Shape narrow elliptic, apex acute, base acute and cuneate.
Margin is lobed. The lobes are acute and both of their apical and
basal sides are convex. The sinuses are rounded. Venation is
craspedodromous. The midvein is relatively thick and basally it is
weakly curved. The secondaries arise at an angle of about 45° and
they run into the teeth. Their course is different: concave in the
lower ones and convex in the upper ones. The tertiary veins arise
at right angle from both of the upper and lower sides of the
secondaries. Their course is convex. The quaternary venation is
random reticulate.
M a t e r i a l : No. 149-150, 155, 157, 278a, 393. 410, 414,
435, 444.
Quercus urophylla U N G E R 1 8 5 0
1856 Quercus urophylla UNG.: KOVÁTS, p. 22, Tab. 2, Fig. 7. 1959
Quercus urophylla UNG.: ANDREÁNSZKY. p. 102, Tafel 27, Fig. 4,
Tafel 68, Fig. 5.
We have four specimens of the species. The leaves are
symmetrical and have petiole. Lamina length 2.9-7 cm, width 1.1-3.2
cm. Shape oblanceolate, apex obtuse, base acute and cuneate. There
are some small acute teeth on the margin. Venation is camptodromous
and seems to be cladodromous. The midvein reaches the apex and its
course is weakly curved. The angle of divergence of the relatively
thin secondaries is acute. The angle varies near the apex but it
remains acute. The tertiary venation is orthogonal reticulate.
M a t e r i a 1: No. 120, 216, 489, 534.
Quercus mediterranea U N G E R 1 8 4 7
1847 Quercus mediterranea UNGER: UNGER, p. 114, Tab. 32, Figs
5-9. 1856 Quercus szirmayana Kov: KOVÁTS, p. 21, Tab. 2, Figs 1-5.
1959 Quercus mediterranea UNGER: ANDREÁNSZKY, p. 101, Tafel 27,
Fig. 5. 1986 Quercus cf. mediterranea UNGER: KNOBLOCH &
VELITZELOS, p. 30, Tafel 14, Figs 4, 11, Tafel 15, Fig. 7.
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Only three small and badly preserved specimens have turned up.
The leaves are symmetrical. The petiole has not remained due to the
fragmentation. Lamina lengtb of the first leaf 2.6 cm, width 1.4
cm. The length of the second 6.7 cm, width 2.2 cm. The third one is
a small, hardly recognizable leaf. Its length 1.2 cm, width 0.6 cm.
I have described the first and the second leaves. Shape narrow
obovate, apex rounded, base obtuse. Margin seems to be entire. Only
the midvein is observable. It branches under the apex.
M a t e r i a I: No. 115, 133b, 187.
Quercus neriifolia A . B R A U N 1 8 4 5
1959 Quercus neriifolia A. BR. ex HEER: ANDREÁNSZKY, p. 105,
Tafel 28, Fig. 2. 1964 Quercus neriifolia AL. BR.: KUTUZKINA
(TAKHTAJAN) p. 195, Tab. 6, Figs 1, 2, Tab. 17, Fig. 12. 1987
Quercus neriifolia A. BR.: PALAMAREV & PETKOVA, p. 71, Tab. 20,
Figs 2-3, 7.
One fragmented leaf the apex and base of which have not
remained. Presumably it is symmetrical with petiole. Lamina length
6.3 cm, width 1.9 cm. Shape very narrow elliptic. Margin is entire.
Venation is camptodromous and reticulodromous, the course of the
midvein is curved. The secondaries arise at angles of about
50°-60°, and they are curved. They do not reach the margin. The
tertiary venation cannot be observed.
M a t e r i a l : No. 447b.
Quercus sapperi ( M E N Z E L 1933) M A I 1967 (fruit)
(Fig- 7) 1986 Quercus sapperi (MENZEL) M A I : GREGOR, p. 52,
Tafel 25, Figs 1-4. 1991 Quercus sapperi (MENZEL) M A I : WALTHER
& ZASTAWNIAK, p. 172, PI. 7, Figs lb, la, Fig. 9: 1.
The remain of the cupule of a fruit the diameter of which is 4
cm. An outer, concentrical "fringe" is also observable. Its width
is 0.8 cm.
M a t e r i a 1: No. 543.
AQUIFOLIACEAE
Hex gracilis K O L A K O V S K U 1 9 6 4
1964 Ilex gracilis KOL.: KOLAKOVSKU, p. 46, Tab. 9, Figs
2-8.
Only one leaf represents the species. It is symmetrical and has
petiole. Lamina length 1.9 cm, width 0.9 cm. Shape elliptic, apex
obtuse, base cuneate. Some teeth are observable on the margin. The
teeth are obtuse and look like small emarginations. Venation
camptodromous and cladodromous. The midvein is weakly curved. The
secondary venation arises at acute angle. The course of the
secondaries is sinuous and they branch near the margin. The
tertiary venation is orthogonal reticulate.
M a t e r i a 1: No. 35.
ROSACEAE
Rosa lignitum H E E R 1 8 6 9
1971 Rosa bohemica ENGELH.: BUZEK, p. 61, PI. 24, Figs 1-19.
1990 Rosa lignitum HEER: HABLY, p. 33, PI. 29, Fig. 3.
We have 3 specimens which arc symmetrical with relatively small
petiole. Lamina length 1.6-2.7 cm, width 0.8-1.4 cm. Shape
elliptic, apex acute, base acute or nearly obtuse. Margin is
regularly toothed and the teeth are small. Both of their apical and
basal sides are convex, the sinuses aie angular. Venation is
presumably craspedodromous.
M a t e r i a l : No. 223b, unnumbered.
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THYMELAEACEAE
Daphne oehningensis ( A . B R A U N 1 8 5 1 ) W E Y L A N D 1 9
3 8
1985 Daphne oehningensis (A. BRAUN) WEYLAND: HABLY, p. 114, PI.
32, Figs 1-4.
Two fragmented leaves the base of which has not remained. They
are symmetrical and their shape is obovate. Lamina length 3-3.5 cm,
width 1-1.1 cm. Apex rounded, base presumably decurrent. Margin is
entire. Their venation is hardly observable only the midvein is
recognizable. Its course is straight and does not reach the
apex.
M a t e r i a l : No. 134, unnumbered.
ACERACEAE
Acer integerrimum ( V I V I A N I 1 8 3 3 ) M A S S A L O N G O
1 8 5 8
(Fig. 15)
1959 Acer decipiens A. BR.: ANDREÁNSZKY, p. 164, Tafel 52, Fig.
3 1971 Acer integerrimum (Vív.) MÁSSAL.: BÖZEK, p. 79, PI. 37, Figs
1-8, PI. 38, Figs 1-10 1986 Acer integerrimum (VIVIANI) MASSALONGO:
KNOBLOCH & VELITZELOS, p. 14, Tafel I , Fig. 1.
Symmetrical leaves which are composed of 3 or 5 lobes and have
petiole. Lamina length 2.9-7.2 cm, width 3.8-10.2 cm. Apex
attenuate, base cordate. Margin entire. Venation is basal
actinodromous; namely, 3 or 5 primary veins run into the apex of
the lobes. The angle of divergence of the secondaries is acute. The
tertiary venation is reticulate.
M a t e r i a l : No. 205, 206a, 207-209a, 210a, 273c, 367a,
368, 371, 524, 525, 526b, 528a.
Acer sp. (fruit) (Fig. 16)
Some disamara fruits, unfortunately their parts have not
remained together. Length 4-4.5 cm. Some curved veins running from
the upper margin toward the lower part of the fruit are
observable.
M a t e r i a 1: No. 202-204, 554.
VITACEAE
Vitis teutonica A . B R A U N 1845
1959 Vitis teutonica A. BR.: ANDREÁNSZKY, p. 171, Tafel 53, Fig.
1.
One fragmented, symmetrical leaf. Its petiole has not remained.
Base is cordate, margin is lobed. Venation is basal actinodromous
with 5 primary veins running into the teeth. The secondary veins
arise at acute angle (60°-70°) from the lower sides of the
primaries (except the middle vein) and run into the lobes. The
tertiary venation is thin reticulate.
M a t e r i a l : No. 523.
JUGLANDACEAE
Engelhardtia orsbergensis ( W E S S E L et W E B E R 1 8 5 6 )
JÄHNICHEN, M A I et W A L T H E R 1 9 7 7
1976 Engelhardia détecta SAPORTA: KNOBLOCH & KVACEK, p. 27,
Tafel 11, Figs 3, 11, Tafel 12, Figs 1, 2, 8, Tafel 19, Fig. 6,
Tafel 20, Fig. 2 .
1984 Palaeocarya orsbergensis (WESSEL et WEBER) JÄHNICHEN,
FRIEDRICH et TAKAC: JÄHNICHEN, FRIEDRICH & TAKAC, p. 110, Pl.
1, Figs 1-6, Pl. 3, Fig. 1, Pl. 4 , Figs 1-4, PI. 5, Figs 1-5.
-
Some fragmented leaflets have turned up. They are asymmetrical
and have petiole. Only one of them is measurable, its length is 3.7
cm, width is 1.3 cm. Shape is very narrow elliptic, apex is
attenuate, base is presumably asymmetrical cuneate or dccurrent.
Margin is toothed. The teeth are very small and much more sparse
than that of the species of Pterocarya and Carya. The teeth are
acute and charactcristical. Venation is hardly observable but it is
likely to be semicraspedodromous. The midvein is thick and weakly
curved. The angle of divergence of the secondaries arising from the
lower part of the midvein is right and that of those that arise
from the upper part is acute. The neighbouring secondaries
anastomose at obtuse angle and form loops. Before the anastomosis a
vein branches toward the margin and runs into a tooth. The tertiary
venation is orthogonal reticulate.
M a t e r i a l : No. 178, 201, 376c, 394, 483a, 536c.
Pterocarya paradisiaca ( U N G E R 1 8 4 9 ) ILJ INSKAJA
1962
1959 Pterocarya denticulata (O. WEB.) HEER: ANDREÁNSZKY, p. 120,
Tafel 32, Fig. 5, Tafel 33, Figs 2-3.
The species is represented with a great number of leaves. They
are symmetrical. Their petiole cannot be observed (presumably it
has not remained). Lamina length 2.5-10 cm, width 1.2-3.9 cm. Shape
elliptic, in some cases weakly obovate. Apex acute, base obtuse or
weakly acute. Margin is toothed similarly to the species of Carya.
The only difference between their teeth is the density of the teeth
namely the teeth of Pterocarya paradisiaca is more dense than that
of the species of Carya. The teeth are acute. Their apical side is
acuminate, the basal is concave. The sinuses are angular. Venation
is semicraspedodromous. The midvein is curved. The secondaries
arise at angles of about 70°-80° and their course is curved. The
neighbouring secondaries anastomose and form loops. Smaller veins
branch from the secondaries toward the margin and run into the
teeth. The tertiaries arise at right angle from both of the apical
and basal sides of the secondaries and connect them. A vein is
observable, arising from the apical initial part of the
secondaries. Later it bends back to the midvein.
M a t e r i a l : No. 52a, 67a, 79, 91, 97, 103, 104, 106a, b,
160a, 162a, 163, 164b-165, 168, 174, 175, 181, 183b, 184, 189,
192b, 194, 195, 199, 200a, 235b, 253b, 302e, 390d, 436c, 478b,
480a, 491-494a 558a.
Carya minor SAPORTA et M A R I O N 1 8 7 6
1964 Carya minor SAPORTA et MARION: KOLAKOVSKU, p. 96, Tab. 8,
Fig. 2. 1986 Carya minor SAPORTA et MARION: KNOBLOCH, p. 27, Taf.
14, Figs 1-4, Taf. 15, Figs 2, 3.
We have only some leaflets among the remains. They are
symmetrical and have petiole. Shape elliptic or obovate. Lamina
length 0.7-6.2 cm, width 0.5-2.2 cm. Apex acute, base cuneate or
weakly decurrent. Margin is toothed. The apical side of the teeth
is acuminate, the basal is concave. The sinuses are angular.
Venation is semicraspedodromous but the loops are not so expressed
as it is in the case of Pterocarya paradisiaca. The course of the
midvein is straight. The secondaries arise at angles of about
60°-70°. They are curved and do not reach the margin.
M a t e r i a 1: No. 73, 80, 81, 147, 191, 197, 198a, 289b,
455a.
Carya denticulata ( W E B E R 1852) ILJINSKAJA 1 9 6 4
1964 Carya denticulata (WEB.) ILJINSKAJA: KOLAKOVSKU, p. 95,
Tab. 36, Figs 3-6.
Weakly asymmetrical leaflets, presumably they had petiole.
Lamina length of the two leaflets is 2.2 cm and 4.1 cm, width is
1.2 cm and 1.3 cm. Shape lanceolate, apex attenuate, base obtuse.
Margin is toothed like in the previous species of Carya, namely,
the apical side of the teeth is acuminate and the basal is concave.
Venation is semicraspedodromous and the loops are formed
imperfectly.
M a t c r i a 1: No. 186, 361.
Carya sp.
We have a great number of leaflets belonging to the genus of
Carya. They are symmetrical and have petiole. Lamina length 3.2-9
cm, width 0.9-3.4 cm. Shape obovate, apex acute (but more weakly
than that of Pterocarya paradisiaca), base obtuse. Teeth are
similar to that of Pterocarya paradisiaca but they are bigger and
more sparse. Venation is semicraspedodromous and the loops are
formed imperfectly.
M a t e r i a 1: No. 60, 78, 84, 99a 100b, 101, 105, 107, 129b,
161a, 164a, 166, 167a, 169, 171, 177b, 180, 182, 188a, 190a, 196,
200c, 209b, 223c, 256b, 334b, 381b, 482, 488a, 557.
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ANACARDIACEAE
Rhus s p.
Asymmetrical leaves, petiole is not observable. Shape
lanceolate. Lamina length 4.2-4.9 cm, width 1.7-2.3 cm. Apex
attenuate, base presumably obtuse. Margin is toothed. The teeth are
acute and both of their apical and basal sides are convex. The
sinuses are angular. Venation craspedodromous. The course of the
midvein is curved. The secondaries arise at acute angle. Their
course is curved and they run into the teeth. Some of them have a
vein branching from the lower side of the secondary veins and it
runs into a sinus. The tertiary venation cannot be observed.
M a t e r i a 1: No. 183a, 272b.
R U T A C E A E
Pteleaecarpum europaeum ( B R O N N 1838) B U Z E K et K N O B L
O C H in K N O B L O C H 1969
1856 Ptelea macroptera Kov.: KOVÁTS, p. 51, Tab. 1, Fig. 2. 1959
Ptelea macroptera Kov: ANDREÁNSZKY, p. 151. 1971 Pteleaecarpum
europaeum (BRONN) BBZEK et K N O B L O C H in KNOBLOCH: BBZEK, p.
70, PI. 31, Figs 1-21.
A fragment of a fruit. Its length is 2.8 cm, width 0.9 cm. The
seed is in the middle of the samaroid fruit. A thin venation is
recognizable on the outer paît of the fruit.
M a t e r i a l : No. 451c.
LAURACEAE
Lauraceae gen. et sp.
Some symmetrical leaves which have relatively thick petiole. The
leaves are thick and coriaceous. Shape narrow elliptic. Lamina
length 4.6 cm, width 1.5-2.8 cm. Apex acute, base acute. Margin
entire. Venation camptodromous, brochidodromous. Midvein straight.
The secondaries arise at acute angle (60°) and they do not reach
the margin. They anastomose with each other at acute and right
angles and form loops. The tertiary venation is thin and hardly
visible. Presumably its pattern is random reticulate.
M a t e r i a l : No. 270, 335a, 365, 536a.
Laurus sp.
Symmetrical leaf with relatively long petiole. Lamina length 3.4
cm, width 1.2 cm. Shape narrow elliptic. Apex presumably acute,
base cuneate. Venation camptodromous, brochidodromous. Midvein
straight. The angle of divergence of the secondaries is acute
(45°-50°). The neighbouring veins anastomose at right angle and
form loops. The tertiary venation is reticulate.
M a t e r i a l : No. 268.
cf. Daphnogene sp.
One badly preserved, small leaf. The apex and the base cannot be
observed. The base must have been acute. Lamina length 1.1 cm,
width 0.7 cm. Margin is entire. Venation seems to be acrodromous.
Midvein is weakly curved. A strong, basal pair of veins arises from
the base. The angle of divergence of the secondary veins is acute.
The tertiary venation is not observable.
M a t e r i a l : No. 133a.
ARALIACEAE
Araliaceae gen. et sp. I.
One fragmented, badly preserved leaf. Only its base has remained
(without the petiole). Presumably it was
-
symmetrical. It must have been a quite big leaf but
unfortunately its shape cannot be concluded from the remains. The
base is cordate, its margin seems to be entire. Venation is
actinodromous with 7 thick primary veins arising from one point of
the base. The angle of divergence of the secondaries is acute,
their course is curved. The tertiary and quaternary venation is
hardly visible, their pattern seems to be random reticulate.
M a t e r i a l : No. 369.
Araliaceae gen. et sp. II .
A fragmented remain of the lower part of a leaf. Its shape
cannot be described but it must have been quite a big leaf. The
petiole and the apex have not remained. The base is wide, rounded
and weakly cordate. There are two lobes on the margin: a smaller
and a bigger one. They aie acute, their apical and basal sides are
convex, the sinuses are angular. Venation palinactinodromous. One
of the primaries runs into the lower lobe and from the basal side
of the other primary vein other veins arise. One of them runs into
the second lobe. The secondary venation arises at right angle.
Intersecondary veins are also observable. The tertiary venation is
orthogonal reticulate.
M a t e r i a 1: No. 370.
EBENACEAE
Diospyros aff. pannonica E T T I N G S H A U S E N 1 8 5 1
1986 Diospyros aff. pannonica ETTINGSHAUSEN: KNOBLOCH, p. 33,
Taf. 2, Fig. 11, Taf. 3, Fig. 9, Taf. 4, Fig. 4.
Two specimens represent the species. They are symmetrical and
have petiole. Lamina length is 3.8 cm and 3.0 cm and width is 1.7
cm and 1.5 cm. Shape is narrow ovate. Apex is acute or weakly
obtuse and the base is rounded. Margin is entire. Venation is
camptodromous, reticulodromous. The midvein is straight and
relatively thin. The secondary veins arise at acute angle but
toward the apex the angle decreases (near the base the angle is
60°-70° whereas close to the apex it is only 45°). The secondaires
do not reach the margin and they disappear without branching. Their
course is curved. The tertiary venation is so thin that I could not
describe it.
M a t e r i a I : No. 285b, 487.
Diospyros sp.
We have some symmetrical leaves in the collection. They have
petiole. Lamina length 2.8-5.3 cm, width 1.7-2.7 cm. Shape narrow
ovate, apex acute, base obtuse. The margin is toothed. The teeth
are very small and acute. Their apical side is acuminate, the lower
is concave. The sinuses are angular. Venation is camptodromous,
reticulodromous, the midvein is straight. The secondaiy veins arise
at acute angle (50°-60°). The third and fourth pairs of veins do
not reach the margin and they are strongly curved. The tertiary
venation is extremely thin and its pattern is orthogonal
reticulate.
M a t e r i a 1: No. 74, 167c, 230, 288.
SAPINDACEAE
Sapindus falcifolius ( A L . B R A U N 1 8 3 6 ) A L . B R A U N
1851
1959 Sapindus falcifolius A. BRAUN: ANDREÁNSZKY, p. 156, Tafel
47, Figs 3-4. 1971 Sapindus falcifolius ( A L . BRAUN) A L . BRAUN:
BÔZEK, p. 82, Pl. 35, Figs 1-6.
The species is represented with a small number of specimens.
Asymmetrical leaves with petiole. Lamina length 5.1-6.1 cm, width
1.7-2.5 cm. Shape lanceolate, apex acute, base asymmetrical obtuse
and weakly rounded. Margin entire. Venation is hardly observable.
The midvein is curved. The secondary and tertiary venation is not
visible.
M a t e r i a 1: No. 83, 96, 162b, 177a, 188b, 193. 324a.
H A M A M E L I D A C E A E
Parrotia pristina (ETTINGSHAUSEN 1 8 5 1 ) S T U R 1 8 6 7
1959 Parrotia fagifolia (GOEPP.) HEER: ANDREÁNSZKY, p. 73, Tafel
15, Figs 2-4, 6, Tafel 16, Fig. 1.
-
1971 Parrotia pristina (ETTINGSHAUSEN) STUR: B8ZEK, p. 52, Pl.
17, Figs 1-11. 1988 Parrotia pristina (ETTINGSHAUSEN) STUR:
KNOBLOCH, p. 4, Tafel 13, Fig. 6.
We have only few leaves among the remains. They are symmetrica],
the petiole is not visible. Lamina length 5.9-6 cm, width 3.2-4.3
cm. Shape wide obovate, apex attenuate, base presumably obtuse.
There are some obtuse teeth on the upper part of the margin. The
apical and basal sides of the teeth are convex and the sinuses are
rounded. Venation is craspedodromous, the midvein is straight. The
secondary veins arise at acute angle (50° -60°) but on the basal
part of the leaves there is a pair of relatively strong veins the
angle of divergence of which is 45°. These pairs of veins are
straight whereas the other secondaries are curved and run into the
teeth. The tertiary venation is not observable.
M a t e r i a 1: No. 272a, 327b, 355.
M O N O C O T Y L E D O N O P H Y T A
Monocotyledonae gen. et sp.
Thick pieces of stem the width of which is 1.5 cm. Parallel
venation is undoubtedly recognizable but unfortunately a more
precise taxonomical description is impossible.
M a t e r i a 1: No. 70b, 376d, 429c.
POTAMOGETONACEAE
Potamogeton sp. (Fig. 1 7 )
It is alliaceous, presumably the remain of a rhizome. Length 1.3
cm, width 1.2 cm. Its upper part is attenuate. Parallel venation is
recognizable, the veins arise from one point of its lower part and
converge in the "apex".
M a t e r i a 1: No. 317c.
LILIACEAE
Smilax weberi W E S S E L in W E S S E L et W E B E R 1 8 5
5
(Fig. 18)
1971 Smilax weberi WESSEL in WESSEL et WEBER: BGZEK, p. 89, PI.
44, Figs 1-5, Pl. 45, Figs 1-4. 1992 Smilax weberi WESSEL et WEBER:
HABLY, p. 204. PI. 2, Figs 5- 6.
One fragmented leaf which was presumably symmetrical. The
petiole has not remained. Its shape cannot be defined due to its
fragmentation. Its apex is rounded, the base is saggitate or
hastate, its margin is entire. Venation is parallelodromous
presumably campylodromous with 7 primary veins. The veins run
toward the apex.
M a t e r i a 1: No. 535.
P A L M A E
Sabal major ( U N G E R 1 8 4 7 ) H E E R 1855 (Figs 19, 2 0
)
1985 Sabal major (UNGER) HEER: HABLY, p. 120, PI. 36 Fig. 4; PI.
37 Figs 2-3; PI. 38 Figs 1-3.
Incomplete leaf which is palmate. The petiole, apex, basis and
margin of the flabellate leaf have not remained. The pinnae are
observable, their width is 0.4-0.6 cm. The species has turned up
from Ipolytamóc in a great quantity. However, in general it is a
rare and accessory element.
M a t e r i a l : No. 600.
-
8 9
Figs 4-9. Fossils from Erdőbénye-Ligetmajor: 4 = Cupressus
sempervirens L. No. 373, 5 = Quercus kubinyii (KOVÁTS et Err.)
BURGER No. 377. 6 = Quercus kubinyii (KOVÁTS et Err.) BERGER No.
390b, 7 = Quercus sapperi (MENZEL) Mai No. 544, 8 = Carpinus
neilreichii KOVÁTS NO. 139, 9 = Carpinus pyramidalis GAUDIN in
GAUDIN
et STROZZI NO. 121
-
15 ,1cm, 16 1 cm
Figs 10-16. Fossils from Erdőbénye-Ligetmajor: 10 = Podogonium
knorrii (A. BRAUN) HEER (leaflet) No. 479, 11 = Podogonium knorrii
(A. BRAUN) HEER NO. 590, 12 = Podogonium knorrii (A. BRAUN) HEER
(fruit) No.
225, 13 = Zelkova zelkovifolia (UNGER) BUZEK et KOTLÁBA in
KOTLÁBA NO. 7, 14 = Zelkova zelkovifolia (UNGER) BUZEK et KOTLÁBA
in KOTLÁBA NO. 327, 15 = Acer integerrimum (VIVIANI) MASSALONGO NO.
207
, 16 = Acer sp. (fruit) No. 202
-
Figs 17-20. Fossils from Erdőbénye-Ligetmajor: 17 = Potamogeton
sp. No. 317, 18 = Smilax weberi WESSEL in WESSEL et WEBER No. 535,
19 = Sabal major (UNGER) HEER NO. 600, 20 = Sabal major (UNGER)
HEER NO. 600
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A N A L Y S I S O F T H E FLORA
The species of the flora in Erdőbénye-Ligetmajor form a
taphocoenosis; therefore, they do not represent a uniform
ecological type. We have found mostly fossils of angiosperms and
only some specimens of ferns and angiosperms have turned up.
Comparing the number of specimens o f certain families to the total
number of specimens (Table 1) we can conclude that more than 70% of
the flora is formed by the families of Fabaceae, Fagaceae and
Ulmaceae. The families of Juglandaceae and Betulaceae are also
represented in a relatively great percentage. The other groups of
plants can be regarded as accessory and relict elements o f the
flora. From Table 2 we can come to the same conclusion. Podogonium
knorrii (A. B R . ) H E E R forms 25% of the fossils and more than
20% of the flora is composed of Quercus kubinyii ( K O V Á T S et E
r r . ) B E R G E R (Fagaceae) and Zelkova zelkovifolia ( U N G E R
) B Ö Z E K et K O T L Á B A
in K O T L Á B A (Ulmaceae). The proportion of the species of
Carya sp. , Pterocarya paradisiaca ( U N G E R ) ILJ INSKAJA and
Acer integerrimum ( V I V I A N I ) M A S S A L O N G O is similar
to the proportion of their families. More than 60% of the species
existing in the flora belongs to the group of deciduous, mesophytic
elements. Presumably some of them were thermophyl and
xerophytic.
R E C O N S T R U C T I O N OF T H E FLORA
On the basis of the species, families and elements o f the flora
3 types o f vegetation are distinguished: 1. Mesophyl, subtropical
forest. The dominant species are Quercus kubinyii ( K O V Á T S
et
E r r . ) BERGER, Podogonium knorrii (A. B R . ) H E E R and
Acer integerrimum ( V I V I A N I ) M A S S A L O N G O . Though,
it must be added that a reason for the great quantity of Podogonium
knorrii (A. B R . ) H E E R might be that it had compound leaves
and the leaflets were small-sized. The probability of the
fossilization of a small leaf is much higher than that of a bigger
leaf. On the basis o f the main characteristic features in recent
similar forests the mesophyl, subtropical forest of Ligetmajor was
formed by shrubs and trees which were not higher than 20 m.
Presumably some species of Fagus and Carpinus growing in higher
areas in more humid patches must have risen from the average level
of the foliage crown. Presumably mesophyl elements also appeared
but only in a small number. Besides the latter genera such
accessory elements can be mentioned as species of Laurus,
Diospyros, Araliaceae, Sapindus falcifolius A . B R A U N , Vitis
teutonica A. B R A U N and Smilax weberi W E S S E L in W E S S E L
et W E B E R . Comparing with the classical localities in Erdőbénye
(Barnamáj, Kővágó-oldal) we can conclude that the deciduous
elements have much more representatives in Ligetmajor.
2. Riparian and grove forests. Its presence is proved by the
appearance of the family of Juglandaceae. Mainly species of Carya
and Pterocarya paradisiaca ( U N G . ) I L J I N S K A J A must
have formed the vegetation but mesophyl species from the mesophyl
subtropical forest must have also appeared. The presence of the
rare Engelhardtia orsbergensis ( W E S SEL et W E B E R ) J Ä H N I
C H E N , M A I et W A L T H E R , is worth mentioning.
3. Water or swampy coenosis. The remains of a sphenopsid, the
stems of monocotyledonous plants and the Potamogeton sp. prove the
presence of a water or swampy coenosis. However, it must have
formed only a small ratio o f the flora; moreover, we have to
conclude that it had the smallest extension. Especially i f we take
the fact into consideration that the probability of being
fossilized is much higher in such coenosis than in the other ones.
I t is an interesting difference that Cystoseirites partschii
STERNB. (Phaeophyta) being characteristic of the classical floras
is totally missing here.
-
Figs 21-22. The leaf distribution of the flora in: 21 =
Ligetmajor, 22 = Kővágó-oldal
DIFFERENCES BETWEEN T H E FLORAS O F LIGETMAJOR A N D K Ő V Á G
Ó - O L D A L
Cystoseiritespartschii STERNB. being mass in Kővágó-oldal is
totally missing in Ligetmajor. We can draw the conclusion that
presumably Kővágó-oldal was close to the sea whereas in Ligetmajor
there must have been a fresh-water inlet. The genera o f Ginkgo,
Liquidambar, Glyptostrobus, Sequoia and Libocedrus were found in
Kővágó-oldal; however, they are missing in Ligetmajor. The species
of Glyptostrobus and Liquidambar are riparian elements. It refers
to that the extension of a riparian coenosis must have been bigger
in Kővágó-oldal than in Ligetmajor. A reason for this must have
been the quicker change of the terrain in Ligetmajor; that is to
say that a steeper slope must have existed in this area. There is
another interesting difference in respect of the species of
Quercus. In Kővágó-oldal more than 2 5 % of the flora is formed by
Quercus mediterranea U N G E R and Quercus urophylla U N G E R
(Quercus kubinyii is represented with only 2 . 8 % ) . In
Ligetmajor the ratio of Quercus kubinyii (KOVÁTS et ETT. ) BERGER
is similar and it seems to replace Quercus mediterranea U N G E R
and Quercus urophylla U N G E R in this flora (these species are
represented with only some specimens). Two kinds of explanation are
conceivable: namely, the ecological features must have been
different in Kővágó-oldal and Ligetmajor or their floras are not
uniform in respect of their age (Table 3 ) .
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P A L A E O C L I M A T O L O G I C A L RECONSTRUCTION
I have attempted the palaeoclimatological reconstruction by
means of two methods. 1. O n the basis of the relations of species
and their climatic demands. The xerophytic
elements are dominant in the flora. However, it cannot be
considered as being the same as the stereophyllous forests since
the ratio of deciduous elements is extremely high. It can be
compared principally with stereophyllous forests of the east
Mediterranean region in which deciduous elements are also
represented. The climate of that region and Ligetmajor must have
been similar. The reason for the presence of mesophyl and riparian
elements must have been the closeness of water. On the basis of the
relations of species and the spread o f the relative species we can
conclude that the climate must have been a Mediterranean one with
dry and hot summer and mild winter. Presumably it was drier than
the climate of Kővágó-oldal. However, some tropical species could
withdraw to the forest though only in a small number.
2. Another comparative method is leaf statistics. The area of
the leaves were calculated from a formula ( C A I N & CASTRO
1959):
A=2/3(LxW) A = area of the leaf or leaflet in sq cm L = leaf
length in cm W = leaf width in cm
The leaves of Ligetmajor (916 specimens) and Kővágó-oldal (607
specimens) were ranged among the following leaf-size
categories:
leptophyll (up to 0.25 sq cm in area) nanophyll (0.25 - 2.25 sq
cm in area) microphyll (2.25 - 20.25 sq cm in area) notophyll
(20.25 - 45.0 sq cm in area) mesophyl 1 (45.0 - 182.25 sq cm in
area) macrophyll (182.25 - 1640.2 sq cm in area) megaphyll ( 1640.2
sq cm in area)
I have represented the results in Table 4. The leaf distribution
of the two localités (Figs 21- 22) does not differ essentially;
only the
proportion of notophyll leaves is higher in the flora of
Ligetmajor. However, we have to take into consideration that the
probability of a bigger leaf's fossilization is much lower than
that of a smaller one and the difference may also arise from the
collection itself. On the basis of all these we can draw the
conclusion that mainly small-sized leaves (nano- and microphyll)
formed the floras. It is in accordance with the results of the
previous observation since xerophytic elements have generally
small-sized leaves.
DISCUSSION
Besides the classical localities in Erdőbénye (Barnamáj,
Kővágó-oldal), another palaeobo-tanical site was found in a
siliceous earth mine in the middle of the 50's. During exploitation
deeper layers were opened and remains of plants were found in large
quantities under a well stratified rhyolitic tuff layer which had
fallen into water. On the basis of the findings we can draw some
conclusions:
The predominant species of the flora are Podogonium knorrii (A.
B R A U N ) HEER, Quercus kubinyii ( K O V Á T S et ETT.) BERGER
and Zelkova zelkovifolia ( U N G E R ) B Ö Z E K et K O T L Á B A
in
K O T L Á B A . Since the majority of the flora is formed by
Mediterranean and xerophytic elements
-
the climate must have been Mediterranean with hot and dry summer
and mild winter. Mostly it can be compared with the climate of the
east Mediterranean since those stereophyllous forests in which
deciduous elements also exist can be found there. Floristically the
two localities (Ligetmajor and Kővágó-olda l ) are quite similar.
In respect o f the species o f Quercus arose a question, that is to
say why Quercus kubinyii ( K O V Á T S et E T T . ) B E R G E R
replaces the stereophyllous species (Quercus mediterranea U N G E R
and Quercus urophylla U N G E R ) in Ligetmajor.
Two kinds of explanation are conceivable: namely, the floras are
not uniform in respect of their age and the other conception is
that the ecological conditions were different.
Another difference existing between the two floras is that
Cystoseirites partschii STERNB. is totally missing in Ligetmajor.
From this the conclusion can be drawn that in this region a
freshwater inlet must have existed. On the other hand, the riparian
elements form a less significant group in Ligetmajor than in
Kővágó-oldal. This can presumably be attributed to differences in
field conditions: namely, in Ligetmajor there must have existed a
steeper slope than in Kővágó-oldal.
Table 1. The percentage of the plant families in Ligetmajor
Families Percentage of the families (number of specimens) in
proportion to the total number
of specimens
Fabaceae 29.6
Fagaceae 21.9
Ulmaceae 21.3
Juglandaceae 10.9
Betulaceae 6.7
Aceraceae 2.8 Lauraceae Aquifoliaceae Anacardiaceae Rutaccae
Araliaceae Sapindaceae 6.1 Rosaceae Uamamelidaceae Ebenaceae
Rhamnaceae Thymelaeaceae
Monocotyledonophyta Gymnospermatophyta 0.7 Sphenopsida
-
Table 2. The percentage of the species (genera) in
Ligetmajor
Species or genera Percentage of the species/genera (number of
specimens) in
proportion to the total number of specimens
Podogonium knorrii (A. BR.) HEER 25.5
Quercus kubinyii (KOVÁTS et Ett.) BERGER 21.46
Zelkova zelkovifolia (UNGER) BUZEK et KOTLÁBA in KOTLÁBA
21.3
Carpinus sp. 6.6
Carya sp. 6.4
Pterocarya paradisiaca (UNGER) ILJINSKAJA 3.9
Acer integerrimum (VIVIANI) MASSALONGO 2.7
Table 3. Differences between the floras of Ligetmajor and
Kővágó-oldal
Species or genera Kővágó-oldal Ligetmajor
Cystoseirites partschii STERNB. X
Ginkgo sp. x
Glyptostrobus sp. x
Sequoia sp. x
Libocedrus sp. x
Liquidambar sp. x
Carya sp. x
Engelhardia orsbergensis (WESSEL el WEBER) JÄHNICHEN, M A I et
WALTHER
x
Pistacia sp. X
Vitis teutonica A. BRAUN - x
Quercus kubinyii (KOVÁTS et Err.) BERGER X < X
Quercus mediterranea UNGER X > X
Quercus urophylla UNGER X > X
-
Table 4. The leaf-size distribution in Ligetmajor and
Kővágó-oldal
leaf-size categories
Ligetmajor number of the leaves
Ligetmajor percentage of
the leaves
Kővágó-oldal number of the leaves
Kővágó-oldal percentage of
the leaves
leptophyll 1 0.1% 1 0.2%
nanophyll 308 33.6% 204 33.6%
microphyll 556 60.7% 391 64.4%
notophyll 50 5.5% 10 1.6%
mesophyll 1 0.1% 1 0
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WALTHER, H . & ZASTAWNIAK, E. (1991) : Fagaceae from Sosnica
and Malczyce (near Wroclaw, Poland). A revision of original
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Author's address: BOGLÁRKA ERDEI
Department of Botany Hungarian Natural History Museum H-1087
Budapest, Könyves Kálmán kit. 40 Hungary
-
Magyarország zuzmóflórájának kézikönyve
[The lichen flora of Hungary] b y K . V E R S E G H Y
The fifth item of the series Studia Naturalia (Scientific
studies of the Hungarian Natural History Museum) represents a
long-needed work, which fills a wide gap in the Hungarian
lichenology. It is the result of 30 years of research work giving a
critical survey of earlier and current approaches of the taxonomy
of the Hungarian lichen flora.
The book comprises two parts. The general part discusses the
definition, thallus morphology, reproduction, ecology, distribution
and environmental conditions of lichens.
The systematical part gives an account on the history of
systematics, taxonomic ranks, preparation practice, chemicals for
identification, systematical outline of the Hungarian lichen genera
and the identification keys of genera and species.
The book covers 717 species. The nomenclature follows literature
sources before 1989. The species name is usually followed by
synonyms used most frequently. Each species has a detailed
discussion on its ecological requirements, floral element category,
distribution pattern in Hungary and additionally some remarks on
the taxonomic status of pheno-logical stages, ecological and
teratological modifications.
Distribution data of the appr. 40,000 specimens revised by the
author based mainly on the botanical collection of the Hungarian
Natural History Museum (BP) and partly on the lichenological
collection of the Eszterházy Károly Teachers' Training College
(EGR).
The book written in Hungarian with an English summary.
ISBN 963 7093 22-2
415 pages, paperback, with 233 figures of black-ink
line-drawing. Price: USD 40, excl. p. & p.
O r d e r s should be sent to:
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