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.

Figs 1-2. Fig. 1. Map of Hungary (locality = Erdőbénye-Ligetmajor). - Fig 2 . Geological map of the Erdőbénye basin

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

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,

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.

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,

(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.

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.

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.

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

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 ) .

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

REFERENCES

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U. U. G. 36: 5-118. CAIN, S. A. & CASTRO, D. M. O. (1959): Manual of Vegetation Analysis. - Harper, New York, 255 pp. DILCHER, D. L. (1974): Approaches to the identification of Angiosperm leaf remains. - Bat. Rev. 1: 1-157. FISCHER, O. & HABLY, L. (1991): Pliocene flora from the alginite at Gerce. - Annls hist. -nat. Mus. natn. hung. 83:

25-47. GREGOR. H. J. (1986): Die Früchte und Samen aus der Oberen Süsswassermolasse von Achldorf (Vilsbiburg, Nie

derbayern). - Documenta Nat. 30 : 49-59. HABLY, L. (1985): Ipolytarnóc alsó-miocén korú flórája. (Early Miocene plant fossils from Ipolytarnoc, N Hungary.)

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Geophytology 22 : 199-205. HAJÓS, M. & PÁI.FALVY, I . (1964): A tokaji-hegység szarmata növénytársulásai. (Sarmatische Pflanzengemeinschaft

des Tokaj-Gebirges.) - Földt. Int. Évi Jel. 1962-ről: 427-435. HERENDEEN, P. S. (1992): A réévaluation of the fossil genus Podogonium. HEER. - In: HERENDEEN, P. S. & Dilcher,

D. L. (eds): Advances in Legume Systematics part 4. The Fossil Record. The Royal Botanic Gardens, Kew, pp. 3-18.

JÄHNICHEN, H , FRIEDRICH, W. L. & TAKAC, M. (1984): Engelhardioid leaves and fruits from the European Tertiary. Paît I I . - Tertiary Res. 6: 109-134.

KNOBLOCH, E . (1969): Tertiäre Floren von Mähren. - Moravské Museum Brno in Zusammenarbeit mit dem Musejni Spolek Brno, Brno, 201 pp.

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KNOBLOCH, E . (1988): Neue Ergebnisse zur Flora aus der Oberen Süsswassermolasse von Aubenham bei Ampfing (Krs. Mühldorf am Inn). - Documenta Nat. 4 2 : 1-27.

KNOBLOCH, E . & VELITZELOS, E. (1986): Die Obermiozäne Flora von Likud bei Elassona (Thessalien, Griechenland). - Documenta Nat. 29: 5-20.

KNOBLOCH, E. & VELITZELOS, E. (1986): Die Obermiozäne Flora von Prosilion bei Kozáni (Süd-Mazedonien, Griechenland). - Documenta Nat. 29 : 29-33.

KOLAKOVSKU, A. A. (1964): Plioccnovaja Flora Kodora. - Izdatelstvo Akademii nauk Graz. CCP, Szuhumi, 209 pp. KOVAR-EDER, J. (1988): Obermiozäne (Pannoné) Floren aus der Molassezone Österreichs. - Beitr. Palüont. Österr.

14: 19-121. KOVÁTS, GY. (1856a): Fossile Flora von Erdőbénye. - Arb. Geol. Ges. Ungarn 1: 1-37. KOVÁTS, GY. (1856b): Fossile Flora von Tällya« - Arb. Geol. Ges. Ungarn 1 : 39-52.

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PÉCS KAY, Z . (1983) : A Tokaji-hegység és a Tiszántúl vulkáni kőzeteinek geomorfológiai vizsgálata. [Geomorpholo-gical study of the volcanic rocks of Tiszántúl and the Tokaj Mountains.] - Egyetemi doktori értekezés. KLTE Ásvány- és Földtani Tanszék, Debrecen, 147 pp.

SZILÁGYINÉ CZIII'ERY, G. (1955) : Beiträge zur Kenntnis der Sarmatischen Flora von Erdőbénye. - Jarb. Ung. Geol. Anst. 44 (1) : 23-32 .

UNGER, F. (1850) : Genera et Species Plantarum Fossilum. - Braumüller, Vindobonae, 627 pp. UNGER, F. (1847) : Chloris protogea. - Engelmann, Leipzig. 110 pp. VÁGNER, I . & MIKLÓS, T. (1990) : Az Erdőbénye-ligetmajori kovaföldbánya földtani felépítése, szarmata flórájának

összehasonlító paleoökológiai elemzése. [The geological structure of the siliceous earth mine in Erdőbénye-Ligetmajor and the palaeoecological study of its Sarmatian flora.j - Szakdolgozat. KLTE Ásvány- és Földtani Tanszék, Debrecen, 35 pp.

WALTHER, H . & ZASTAWNIAK, E. (1991) : Fagaceae from Sosnica and Malczyce (near Wroclaw, Poland). A revision of original materials by Goeppert 1852 and 1855 and a study of new collections. - Ada Palaeobot. 3 1 (1 -2 ) : 153-199.

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.

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