VEGETATION OF THE DOLINES IN MECSEK MOUNTAINS (SOUTH HUNGARY) IN RELATION TO THE LOCAL PLANT
COMMUNITIES
VEGETACIJA V KRAŠKIH DEPRESIJAH IN NJIHOVA POVEZAVA Z LOKALNIMI RASTLINSKIMI ZDRUŽBAMI, PRIMER IZ GOROVJA MECSEK (MADŽARSKA)
Zoltán BÁTORI1, János CSIKY2, László ERDÖS2, Tamás MORSCHHAUSER2, Péter TÖRÖK3
& László KÖRMÖCZI1
Abstract	UDC 630*1(493-13):574
Zoltán Bátori, János Csiky, László Erdös, Tamás Morschhaus-er, Péter Török & László Körmöczi: Vegetation of the dolines in Mecsek mountains (South Hungary) in relation to the local plant communities
This paper deals with the forest Vegetation of the lower part of the dolines in Mecsek Mts. (South Hungary). In order to char-acterize this Vegetation type, samples were compared to the 6 plant communities occurring in the neighbourhood of the dolines. Considering the Vegetation texture and species composition, the Vegetation of the dolines resembles mainly the extrazonal beechwoods (Helleboro odori-Fagetum) and local ravine forests (Scutellaria altissimae-Aceretum) that preserve several mountain, subatlantic relict species in this area. Our study revealed that the plant communities characteristic of the karst surface of Western Mecsek are arranged along a moisture and nutrient gradient. In this system, the habitat conditions of the dolines are similar to those of the beech forests and the local ravine forests, fresh and relatively rieh in nutrients. In the karst, dominated by oak-hornbeam and beech forests, effects of the thermal inversion are the most spectacular where beech forests follow turkey oak-sessile oak forests and oak-hornbeam forests on the lower part of the doline slopes. The described vegetation type of these depressions is developed by edafic factors; its identification as a separate association is not supported by the analyses.
Keywords: doline, vegetation, diagnostic species, habitat conditions, ecological indicator values.
Izvleček	UDK 630*1(493-13):574
Zoltán Bátori, János Csiky, László Erdös, Tamás Morschhaus-er, Péter Török & László Körmöczi: Vegetacija v kraških depresijah in njihova povezava z lokalnimi rastlinskimi združbami, primer iz gorovja Mecsek (Madžarska)
V članku obravnavamo gozdno vegetacijo spodnjega dela kraških depresij v gorovju Mescek (južna Madžarska). Da bi ovrednotili tip vegetacije, smo vzorce primerjali s šestimi rastlinskimi združbami v soseščini vrtač. Vegatacijska tekstura in vrstna sestava spominja na ekstraconalne bukove gozdove (Helleboro odori-Fagetum) in lokalne ravninske gozdove (Scutellaria altissimae-Aceretum), v katerih je ohranjenih več gorskih subatlantskih reliktnih vrst tega območja. Naša raziskava je pokazala, da so rastlinske združbe na kraškem površju zahodnega dela gorovja Mecsek nanizane vzdolž gradientov vlage in hranil. V tem sistemu so habitatni pogoji v dolinah podobni pogojem v bukovih gozdovih in lokalnih ravninskim gozdovom, kjer so tla relativno bogata s hranili in vlažna. Na krasu, kjer prevladujejo hrastovo - gabrovi in bukovi gozdovi so učinki toplotne inverzije še posebej vidni. Tam bukovi gozdovi v spodnjih delih pobočij dolin sledijo cerovo - gradnovim (Quercus cerris - Q. petraea) in hrastovo - gabrovim združbam. Na tip vegetacije vplivajo predvsem talni faktorji, pri čemer naše analize kažejo, da ne gre za novo asociacijo. Ključne besede: vrtače, doline, vegetacija, diagnostične vrste, habitatni pogoji, vrednosti ekoloških indikatorjev.
1	University of Szeged, Department of Ecology, e-mail:*îbatory@gmail.com, kormoczi@bio.u-szeged.hu
2	University of Pécs, Department of Plant Taxonomy and Geobotany, e-mail:,Moon@ttk.pte.hu, erdosl@gamma.ttk.pte.hu, morsi@gamma.ttk.pte.hu
'University of Debrecen, Department of Ecology, e-mail:fn olinia@gmail.com Received/Prejeto: 30.04.2009
introduction
Dolines are small to large sized closed depressions, from a few meters to a few hundred meters in diameter and depth, formed by water infiltration (Veress 2004). Previ -ously, they were considered exclusively as collapse forms, but nowadays the process of doline formation is explained primarily by the change of soil activity on the bedrock (Jakucs 1980). Due to the special morphological, climati-cal and vegetational peculiarities, dolines have become a centre of interest. Several researchers have investigated the special microclimate and the thermal inversion of the dolines (Bacsó & Zólyomi 1934; Polli 1961, 1984; Futó 1962; Wagner 1963; Bárány 1967; Lehmann 1970; Jakucs 1971; Boros & Bárány 1975; Bárány-Kevei 1999). Most of these studies also refer to the microclimatical différences between the slopes with a different exposure. The rela-tionship between the thermal inversion and the vegetation was published, among others, by Beck v. Mannaget-ta (1906), Boros (1935), Morton (1936), Geiger (1950), Grom (1959), Jakucs (1961), Lausi (1964), Sauli (1972), Favretto and Poldini (1985), Kranjc (1997), Pericin and Hürlimann (2001), Borhidi (2002), Polli (2004).
Due to the special climatical conditions of the dolines and sinkholes, they often serve as habitats of rare and valuable species (Budai 1913; Yannitsaros et al. 1996; Tan et al. 1997; Vojtkó 1997; Varga et al. 2000; Bátori et al. 2006; Virók & Farkas 2008), among which we can also find endemisms (Egli et al. 1990; Egli 1991; Brullo & Giusso del Galdo 2001; S0ndergaard & Egli 2006). In the case of climate changes, the dolines and sinkholes may serve as shelters and their vegetation is often more ancient than the neighbouring vegetation (Jakucs 1952). Because of their ability to preserve the reliefs (Bartha 1933; Horvat 1953; Atalay 2006), dolines have an important role in vegetation history.
A major part of Western Mecsek Mountains (South Hungary) is made up of red and gray Permic to lower Triassic sandstone and Triassic limestone. On the 30 km2 karstic surface nearby Abaliget, Mánfa, Orfü and Pécs, more than 2.200 dolines can be found. 1.702 of these dolines have a very small size (d < 20 m; Rónaki 1972). The diameter of the largest one is more than 200 m, its depth exceeds 25 m (Lovász 1971). The formation of these depressions started during the Pleistocene era and it is stili intensive due also to the woodland and the abundant precipitation that exceeds 700 mm per year.
The oak-hornbeam woods (Asperulo taurinae-Car-pinetum) and extrazonal beechwoods (Helleboro odori-Fagetum) are the dominant plant communities on the plateaus and slopes between the dolines. Near to these forms, fragments of turkey oak-sessile oak forests (Potentino micranthae-Quercetum dalechampii) and rock forests (Tilio tomentosae-Fraxinetum orni) also occur. In the deep valleys sorrounding the karstic surfaces there are ravine forests (Scutellaria altissimae-Aceretum). Aider forests (Carici pendulae-Alnetum) can be found in sections of the valleys where streams meander in the deep alluvial deposit. The coenological surveys of these communities were published by Horvát (1956, 1958, 1959, 1972), Kevey (1997), Kevey and Borhidi (1998) and Kevey and Baranyi (2002). The vegetation map of the smaller part of this area has been completed recently (Morschhauser et al. 2000).
The aim of this study was to analyse the species composition and vegetation texture in the dolines of Mecsek Mts. with the Braun-Blanquet method. More-over, we used species indicator values in order to reveal the habitat conditions in the karst depressions.
material and methods
Phytosociological relevés were made in the lower part of the larger dolines (d > 50 m) in Western Mecsek ap-plying the Central European method (Braun-Blanquet 1928). Relevés included the bottom of the dolines or part of them; their size is 400 m2. Due to their funnel-shape they show simultaneously different kinds of expositions and slopes, thus in the analytical table (Tab. 2) only the main ones are presented. Each vegetation relevé was recorded twice: during the spring and the summer aspects. We arranged the species in the table info syntaxonomical groups according to Kevey and Hirmann (2002).
Our 20 relevés were compared to the 120 relevés made by Kevey (1997), Kevey and Borhidi (1998) and Kevey and Baranyi (2002) from the surrounding vegetation of the dolines (6 plant associations).
The relationships among the species composition of the relevés were analysed with standard multivariate Statistical methods (Pielou 1984; Podani 1994). Presence-absence (binary) dataseis were analysed using De-trended Correspondence Analysis (DCA) ordination (Hill & Gauch 1980), which is a common method for indirect gradient analysis. DCA results in an n-dimen-
sional distribution pattern of objects and the rank of the dimensions, in the ordination scatter plot (Fig. 1; two-dimensional hyperspace with the two most important dimensions or axes from the n-dimension), similar relevés are close to each other and less similar relevés are further from each other, thus the distribution of the objects refers to the gradient phenomenon of the background factors. We used the program CANOCO 4.5 (Ter Braak & Šmilauer 2002) for ordination.
Habitat conditions were characterized by the relative ecological indicator values (TWRN) built on the Ellenberg system and adapted to the Hungarian flora by Borhidi (1993). Distributions of indicator values were calculated for each relevé and végétation unit concerning presence-absence and cover (weighted) data, in the case of cover data only the herb layer was considered.
When determining the diagnostic (differential) species, we used the method based on fidelity measurements (Chytry et al. 2002; Tichy & Chytry 2006). The phi coefficient (O) was defined with the JUICE 7.0.25 program (Tichy 2002). This coefficient ranges from -1 to 1, but for convenience, it is multiplied by 100 in the program. The highest phi value of 1 is achieved if the species occurs in ali relevés of the target végétation unit and is absent else-where. The species having a high fidelity (O > 0,45) and at the same time belonging to only one végétation unit were considered diagnosticai.
Plant community names were used according to Borhidi (2003), and plant species names according to Simon (2000).
results
FLOR1ST1CAL CHARACTERISTICS AND DIAGNOSTIC SPECIES
The DCA shows remar kable similarities among the relevés made in the lower part of the dolines in Western Mecsek. Considering the species composition, their végétation re-sembles mainly the beechwoods and ravine forests, less the oak-hornbeam woods and aider forests. There is little resemblance between the végétation of turkey oak-sessile oak forests, rock forests and dolines (Fig. 1).
The végétation of the dolines is dominated by me-sophilous forest plants (Fagetalia), but deciduous forests elements (Querco-Fagea), lllyrian beechwood elements (Aremonio-Fagion) and indifférent species also play an important role in this végétation unit, in dolines, Helle-boro odori-Fagetum and Scutellaria altissimae-Aceretum stands there are several subatlantic relict and mountain species. Some of them (e.g. Dryopteris affinis, Dryopteris dilatata, Dryopteris expansa, Stachys alpina) occur both in dolines, beechwoods and ravine forests, while others (e.g. Actaea spicata, Aruncus dioicus, Lunaria rediviva, Silene dioica) primarily in ravine forests. Due to the cool and humid microclimate of the dolines, they may play a considerable role in the future in the preservation of these species.
Taking the above-mentioned végétation units as the basis of the comparison, there are 4 diagnostic species for the dolines, 36 for the turkey oak-sessile oak forests, 8 for the rock forests, 3 for the oak-hornbeam forests, 7 for the ravine forests, 34 for the aider forests, but in this case, beechwoods do not have diagnostic species (Tab.
3). Diagnostic species of the dolines are Athyrium filix-femina, Paris quadrifolia, (Fagetalia), Atropa bella-donna (Atropion bella-donnae), Dryopteris carthusiana (Alne-tea glutinosae).
Comparing the végétation of the dolines only to the most similar beechwoods and ravine forests, there are 20 diagnostic species between the dolines and the beechwoods, and 41 between the dolines and the ravine forests. In this comparison, Athyrium filix-femina, Circaea lutetiana, Dryopteris filix-mas, Paris quadrifolia, Stachys sylvatica (Fagetalia), Chrysosplenium alternifolium (Alno-Padion), Polystichum aculeatum (Tilio-Aceri on), Galium aparine, Urtica dioica (indifferent), Atropa belladonna, Dryopteris carthusiana are diagnostic in dolines, while Géranium robertianum (Querco-Fagea), Aegopodi-um podagraria, Hepatica nobilis, Lathyrus vernus, Prunus avium, Tilia platyphyllos (Fagetalia), Lathyrus venetus, Ruscus aculeatus, Tamus communis (Aremonio-Fagion) in beechwoods.
Only 4 species occur in the dolines (Athyrium fi-lix-femina, Atropa bella-donna, Dryopteris carthusiana, Paris quadrifolia) that can be considered differential species compared to the ravine forests, while in these forests there are 37 species that can not be found in the dolines, or just occasionally. Most of these are Querco-Fagea (Campanula rapunculoides, Clematis vitalba, Corylus avellana, Crataegus monogyna, Crataegus laevigata, Eu-onymus verrucosus, Géranium robertianum, Ligustrum vulgare, Staphylea pinnata, Tilia platyphyllos), Fagetalia (Aconitum vulparia, Aegopodium podagraria, Carda-mine enneaphyllos, Cerastium sylvaticum, Corydalis cava,
Fig. 1: DCA ordination diagram of 140 relevés (7 Vegetation units) of the study area. Notations: I: Potentillo micranthae-Quercetum dalechampii, II: Tilio tomentosae-Fraxinetum orni, III: Aspe-rulo taurinae-Carpinetum, IV: Helleboro odori-Fagetum, V: Vegetation ofthe dolines, VI: Scutellaria altissimae-Aceretum, VII: Carici pendulae-Alnetum. Eigenvalues for the lst and2"d axis were 0.309 and 0.103, respectively.
Gagea lutea, Géranium phaeum, Isopyrum thalictroides, Knautia drymeia, Lathyrus vernus, Oxalis acetosella, Ra-nunculus lanuginosus, Salvia glutinosa), or Tilio-Acerion (Asplenium scolopendrium, Cystopteris fragilis, Lunaria rediviva, Silene dioica) species. Alliaria petiolata (Galio-Alliarion), Asplenium trichomanes (Asplenio-Festucion pallentis), Cardamine amara (Cardamini-Montion), Carex remota (Alnetea glutinosae), Polystichum setiferum (Aremonio-Fagion), Chelidonium majus, Ranunculus repens, Stellaria media (indifferent), Robinia pseudo-acacia (adventiva) and Chrysosplenium alternifolium are also diagnostic in ravine forests.
STRUCTURAL CHARACTERISTICS
The structure of the canopy in the dolines is very similar to that of the ravine forests. Upper canopy cover varies between 55 and 90%, while the canopy height varies between 23 and 30 m. Upper canopy is primarily composed of Acer campestre, Acer platanoides, Acer pseudoplatanus, Carpinus betulus, Fagus sylvatica and Tilia tomentosa. Some other species (Fraxinus excelsior, Populus tremula, Quercus petraea, Tilia cordata, Ulmus glabra) also play an important role, while other trees (e.g. Quercus cerris) occur sporadically. Younger trees form a lower canopy
with 0-20% cover and 7-17 m height.
The shrub layer is primarily composed of young trees of the canopy layer, with a cover of 0-60% and 1-5 m in height. While Sam-bucus nigra is also typical in this level, others (e.g. Prunus avium, Tilia platyphyllos) occur sporadically.
The herb layer is mostly well developed with a cover varying between 60-100%, and an average height of 25-70 cm. Frequent species include: Allium ursinum, Athyrium filix-femina, Carex pilosa, Carex sylvatica, Cir-caea lutetiana, Dryopteris carthusiana, Dryopteris filix-mas, Galeobdolon luteum s.L, Galium odoratum, Hederá helix, Helleborus odorus, Mercurialis perennis, Moeh-ringia trinervia, Paris quadri-folia, Polygonatum multiflo-rum, Polystichum aculeatum, Pulmonaria officinalis, Rubus hirtus agg., Ruscus hypoglossum, Stachys sylvatica, Veronica montana, Viola reichenbachiana. During the springtime Anemone ranunculoides, Arum maculatum s.str., Cardamine bulbifera, Ficaria verna, Galanthus nivalis are also common.
In the herb layer of deeper dolines, the presence of fern species (Athyrium filix-femina, Dryopteris affinis, Dryopteris carthusiana, Dryopteris dilatata, cf. Dryopteris expansa, Dryopteris filix-mas, Polystichum aculeatum, Polystichum setiferum) and wet woodland species (e.g. Chrysosplenium alternifolium, Urtica dioica) indicates a cool and humid microclimate, and high soil humidity. Because of the open canopy, Atropa bella-donna appears as well in the lower part of the dolines. The species of the herb layer also often occur on the decayed, mossy trees fallen into the bottom of the dolines.
HABITAT CONDITIONS BASED ON ECOLOGICAL INDICATOR VALUES
Habitat conditions of the compared vegetation units can be characterized by the ecological indicator values. The 7 vegetation units show différences according to temperature (T), moisture supply (W), soil reaction (R), and
Tab. 1: W and N indicator spectra of the vegetation of the study area (maximum values are set in bold).
Binary data	I.	II.	III.	IV.	V.	VI.	VII.
(%)	PmQd	TtFo	AtC	HoF	Dolines	SaA	CpA
W2	0.32	0.11	0.00	0.00	0.00	0.00	0.12
W3	9.51	4.96	1.28	0.42	0.22	1.72	0.90
W4	19.68	10.66	11.25	4.68	2.63	5.63	4.87
W5	52.04	45.07	45.75	41.37	36.86	31.57	24.59
W6	14.56	31.97	35.31	43.97	43.86	40.44	34.58
W7	3.88	7.24	6.33	9.56	15.88	16.28	18.88
W8	0.00	0.00	0.07	0.00	0.22	2.51	7.40
W9	0.00	0.00	0.00	0.00	0.33	1.79	7.70
W10	0.00	0.00	0.00	0.00	0.00	0.07	0.48
W11	0.00	0.00	0.00	0.00	0.00	0.00	0.48
N1	0.39	0.00	0.00	0.00	0.00	0.00	0.00
N2	5.70	1.25	1.68	1.46	0.55	0.40	0.96
N3	16.65	7.24	4.18	0.94	2.08	1.85	2.83
N4	25.45	20.74	17.45	14.35	9.08	15.68	11.85
N5	25.65	25.75	31.40	31.70	26.48	22.83	21.05
N6	12.50	14.47	16.71	19.75	23.63	16.41	14.25
N7	8.48	20.28	19.14	24.01	29.43	30.05	30.61
N8	3.82	9.23	8.49	7.59	6.89	10.99	14.07
N9	1.36	1.03	0.94	0.21	1.86	1.79	4.39
Cover data	I.	II.	III.	IV.	V.	VI.	VII.
(%)	PmQd	TtFo	AtC	HoF	Dolines	SaA	CpA
W2	0.10	0.06	0.00	0.00	0.00	0.00	0.03
W3	2.28	2.37	0.08	0.02	0.00	0.36	0.22
W4	19.37	11.29	3.37	1.03	0.54	1.28	0.44
W5	71.90	32.65	28.84	26.92	15.60	12.65	8.52
W6	4.89	48.83	63.44	64.21	77.06	68.22	67.68
W7	1.37	3.23	3.89	7.40	6.72	9.83	18.85
W8	0.00	0.00	0.02	0.00	0.03	0.68	2.55
W9	0.00	0.00	0.00	0.00	0.05	0.56	1.24
W10	0.00	0.00	0.00	0.00	0.00	0.02	0.11
W11	0.00	0.00	0.00	0.00	0.00	0.00	0.11
N1	0.12	0.00	0.00	0.00	0.00	0.00	0.00
N2	1.37	0.15	0.12	0.05	0.00	0.02	0.23
N3	7.90	3.26	0.65	0.14	0.37	0.65	0.53
N4	55.29	21.39	14.68	7.93	1.86	7.86	4.17
N5	24.66	18.19	17.78	32.19	29.71	21.23	23.27
N6	4.40	9.93	6.59	7.29	13.35	6.68	3.51
N7	4.11	31.61	10.62	16.38	13.23	21.13	19.30
N8	1.64	13.36	48.96	35.58	40.61	35.56	45.66
N9	0.44	0.53	0.23	0.05	0.86	0.49	3.09
nitrogen supply (N). The biggest différences are shown in the case of W and N values, which can be successfully applied in local scale (Sala-mon-Albert & Morschhauser 2002) as well.
The use of W values (Tab. 1) with binary data shows that the proportion of W5 (plants of semi-humid habitats, under intermediate conditions) species is the highest in the W indicator spectra of turkey oak-sessile oak forests, oak-hornbeam forests and rock forests, while in the other units the rate of W6 (plants of fresh soils) species is the most dominant. The proportion of W3 (xero-tolerants, but eventually oc-curring on fresh soils), W4 (plants of semi-dry habitats) and W6 species in turkey oak-sessile oak forests is also important, and this is the same in the case of W5 and W7 (plants of moist soils not drying out and well aerated) species in ravine forests and W5, W7, W8 (plants of moist soils tolerating short floods) and W9 (plants of wet, not well aerated soils) species in alder forests. Vegetation units are also divided by the weighted data, showing the maximum at W5 in turkey oak-sessile oak forests, and in the rest of the units the maximum is found at W6. The proportion of W4 spe-cies is also high in turkey oak-sessile oak forests, while W5 species are frequent in oak-hornbeam, rock, and beech forests, and W7 species in alder forests. The W spec-tra of the dolines resemble mostly the spectra of beech forests and ravine forests at the maximum of W6.
Tab. 2: Analytical table of the doline vegetation of the Western Mecsek.
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 A-D K %
								Phragmitetea											
Eupatorium cannabinum	C											1					1	I	5
Lycopus europaeus	C	-	-	-	-	-	-	------	+	-	-	-	-	-	-	-	+	I	5
Solanum dulcamara	C	-	-	-	-	-	-	------	-	-	-	+	-	-	-	-	+	I	5
								Galio-Alliarion											
Alliaria petiolata	C	-	-	-	-	+	-	------	-	-	-	+	-	-	-	-	+	I	10
								Calystegion sepium											
Lamium maculatum	C													2	+	-	+-2	I	10
								Bidentetea (incl. Bidentetalia)											
Persicaria mitis	C	-	-	+	-	-	-	------	-	-	-	-	-	-	-	-	+	I	5
								Atropion bella-donnae											
Atropa bella-donna	C	-	-	+	-	-	+	- - - + - -	1	+	+	+	-	+	-	1	+-1	III	45
								Querco-Fagea											
Acer campestre	A1	2	-	1	2	-	2	2 -----	2	-	1	-	2	2	2	-	1-2	III	50
	A2	2						--- 1 --									1-2	I	10
	B	-	-	+	+	-	+										+	I	15
	C	-	+	+	+	+	+	+ - - + + -	+	+	+	+	+	-	+	-	+	IV	70
	S																	IV	80
Ajuga reptans	C	+	+	-	+	-	-	+ - - + + -	+	+	-	-	-	+	+	+	+	III	55
Brachypodium sylvaticum	C	-	-	-	-	-	-	------	-	-	-	+	-	-	+	-	+	I	10
Bromus ramosus agg.	C	-	-	+	-	-	-	.....+	-	-	-	+	-	-	-	-	+	I	15
Carex divulsa	C	+	-	+													+	I	10
Clematis vitalba	C	+	-	+	-	-	-	.....+	-	-	-	+	-	-	-	-	+	I	20
Cornus sanguinea	C	-	-	+	-	-	-	------	-	-	-	+	-	-	-	-	+	I	10
Crataegus laevigata	C	-	-	-	-	-	-	- + - - - -	+	-	-	-	-	-	+	-	+	I	15
Dactylis polygama	C	-	-	+	-	-	-	------	-	-	-	+	-	-	-	-	+	I	10
Euonymus europaeus	C	-	-	+	-	-	-	- - - - + -	+	-	-	-	+	-	+	-	+	II	25
Ficaria verna	C	2	+	2	+	+	+	+ - + + - 1	+	+	+	+	+	-	2	+	+-2		85
Fragaria vesca	C	-	-	-	-	-	-	------	+	-	-	-	-	-	-	-	+	I	5
Fraxinus excelsior	A1	-	-	-	-	3	-	------	-	-	-	2	-	-	-	-	2-3	I	10
	B	-	-	+	1	+	-	- + - - - +	-	-	-	1	-	+	-	+	+-1	II	40
	C	-	-	-	+	+	+	+ + + - + -	+	+	-	2	-	+	-	+	+-2		60
	S																	IV	70
Geranium robertianum	C	-	+	+				- - - + + -	-	-	+	+	-	-	+	-	+	II	35
Geum urbanum	C	-	-	+	-	-	-	------	+	-	-	+	-	-	-	-	+	I	15
Heracleum sphondylium	C	-	-	-	-	+	-	------	-	-	-	+	-	-	-	-	+	I	10
Hypericum hirsutum	C	-	-	-	-	-	+	------	-	-	-	+	-	+	-	+	+	I	20
Melica uniflora	C	-	+	+	-	1	+	- - - + + +	-	-	+	-	+	-	+	+	+-1	III	55
Mycelis muralis	C	+	+	+	-	+		.....+	+	+	+	+	-	+	-	+	+	III	55
Polygonatum multiflorum	C	-	+	+	-	-	+	+ + -+ + +	+	-	+	+	+	-	+	+	+	IV	70
Populus tremula	A1	-	-	-	-	-	-	--- 2 --	-	-	-	-	1	-	-	-	1-2	I	10
	C	-	-	-	-	-	-	- - - + - -	-	-	-	-	-	-	-	-	+	I	5
	S																	I	10
		1	2	3	4	5	6	7	8	9 10	11 12	13	14	15	16	17	18	19	20	A-D	K	%
Quercus petraea	A1	-	-	-	-	1	-	-	-	- -	--	2	-	-	-	-	-	-	-	1-2	I	10
	C	-	+										+	+	+	-	-	-	+	+	II	25
	S																				II	35
Scrophularia nodosa	C	-	+	+	-	-	+	-	-	--	--	-	-	-	-	-	+	-	+	+	II	25
Staphylea pinnata	C	-	-	-	-	-	-	-	-	--	--	-	-	-	+	-	-	-	-	+	I	5
Stellaria holostea	C								+	- +	+ -	+	-	+	+	-	-	-	-	+	II	30
Symphytum tuberosum	C	+	-	+	-	-	-	-	-	--	+ -	-	-	-	-	-	-	-	-	+	I	15
Tilia cordata	A1	-	1	-	-	-	1	-	2	--	-2	-	-	-	-	-	-	-	-	1-2	I	20
	B	-	2	-	-	-	1	-	2	--	-2	-	-	-	+	-	-	+	+	+-2	II	35
	C	-	1						+	- +	+ -	-	-	+	-	-	-	-	-	+-1	II	25
	S																				III	50
Veronica chamaedrys	C	-	+	+	-	+	-	-	-	- +	+ -	+							+	+	II	35
Veronica hederifolia	C	-	-	-	-	-	-	-	-	--	--	-	-	-	-	+	-	+	-	+	I	10
Viola alba	C	-	-	-	-	-	-	-	-	--	--	-	-	-	-	+	-	-	-	+	I	5
									Alnetea glutinosae													
Dryopteris carthusiana	C	+	+	+	+	-	+	+	+	-1	+ +	+	-	+	+	+	+	+	-	+-1	IV	80
Dryopteris dilatata	C	+	-	-	+	+	-	+	-	--	--	-	+	-	-	-	-	-	-	+	II	25
Dryopteris expansa	C	-	-	-	-	-	-	-	-	--	--	+	-	-	-	-	-	-	-	+	I	5
							Carpino-Fagetea (incl. Fagetalia)															
Acer platanoides	A1	-	2	-	-	-	-	2	1	--	--	-	-	-	2	-	2	-	-	1-2	II	25
	B	-	-	-	-	-	-	-	-	--	--	-	+	-	-	-	-	+	-	+	I	10
	C	-	+	-	-	+	-	+	+	+ -	--	+	+	-	+	-	+	+	-	+	III	50
	S																				III	50
Acer pseudoplatanus	A1	-	-	4	2	1	-	2	3	3 -	3 -	-	2	3	-	3	2	3	2	1-4	IV	65
	A2	-	-	1	-	-	-	2	-	--	-1	-	-	-	-	1	-	-	-	1-2	I	20
	B	-	-	-	2	-	-	-	+	1 -	-2	1	1	-	+	-	-	3	2	+-3	III	45
	C	+	-	1	1	-	-	-	1	+ -	2 1	+	+	1	1	+	+	1	1	+-2	IV	75
	S																				V	85
Aconitum vulparia	C	-	-	-	-	-	-	+	-	--	--	-	-	-	+	-	-	-	-	+	I	10
Actaea spicata	C														1					1	I	5
Aegopodium podagraria	C	-	-	-	-	-	-	-	-	--	2 -	-	-	-	-	-	-	-	-	2	I	5
Allium ursinum	C	2	+	2	5	4	5	3	5	5 5	5 1	4	5	5	5	+	5	1	4	+-5	V	100
Anemone ranunculoides	C	1	+	+	1	+	+	+	+	+ +	+ +	+	+	+	+	1	-	+	1	+-1	V	95
Arum maculatum s.str.	C	+	+	+	+	+	+	+	+	+ +	+ +	+	+	+	+	+	+	1	+	+-1	V	100
Asarum europaeum	C	-	-	-	+	+	+	1	-	1 1	-1	+	+	1	1	-	-	-	1	+-1	III	60
Athyrium filix-femina	C	2	+	1	2	+	1	-	-	1 +	1 1	2	1	+	1	+	2	1	+	+-2	V	90
Cardamine bulbifera	C	2	+	1	+	+	+	1	+	+ +	+ 1	+	+	+	+	2	+	+	1	+-2	V	100
Cardamine enneaphyllos	C	-	2	-	2	1	-	-	1	2 -	+ -	1	2	2	1	-	-	1	-	+-2	III	55
Cardamine impatiens	C	-	-	+	-	+	-	-	-	--	--	+	-	-	-	-	+	-	-	+	I	20
Carex digitata	C	-	-	-	-	-	+	-	-	--	--	-	-	-	-	-	-	-	-	+	I	5
Carex pilosa	C	-	1	+	+	+	1	1	1	+ +	+ 1	1	-	+	+	2	+	1	1	+-2	V	90
Carex sylvatica	C	+	-	+	+	+	+	-	-	- +	+ +	+	+	+	+	+	+	+	-	+	IV	75
Carpinus betulus	A1	2	2	-	2	-	3	2	2	3 4	+ 1	2	1	-	2	2	2	2	2	+-4	V	85
	A2	1	1	1	2	2	2	-	2	1 2	--	-	1	-	2	1	2	-	-	1-2	IV	65
	B	-	+	1	-	+								1					+	+-1	II	25
	C	-	-	1	-	+	-	-	+	--	-1	+	-	+	+	+	-	+	+	+-1	III	50
	S																				V	100
		1	2	3	4	5	6	7	8	9	10	11	12	13	14	15	16	17	18	19	20	A-D	K	%
Circaea lutetiana	C	+	+	+	+	2	+	+	+	+	+	+	2	1	1	+	+	-	+	1	1	+-2	V	95
Corydalis cava	C	-	-	-	-	-	-	-	-	-	-	-	-	-	1	-	1	-	-	+	-	+-1	I	15
Dryopteris filix-mas	C	+	1	1	1	+	+	1	1	1	1	1	1	+	+	1	2	+	1	1	+	+-2	V	100
Epilobium montanum	C	-	-	-	-	-	-	-	-	-	-	-	-	-	-	-	+	-	-	-	-	+	I	5
Euphorbia amygdaloides	C	-	-	-	-	-	-	+	-	+	-	-	-	-	-	+	-	-	-	-	+	+	I	20
Fagus sylvatica	Al	3	2	-	-	2	2	-	-	2	-	3	2	2	2	2	-	2	-	3	2	2-3	IV	65
	A2	-	-	-	-	1	1	-	-	1	-	1	1	-	1	-	1	-	-	-	1	1	II	40
	B	-	1	-	+	-	-	1	1	+	-	1	2	-	-	1	-	-	-	1	+	+-2	III	50
	C	+	+	-	-	-	-	+	+	-	-	+	1	+	1	+	-	+	-	+	+	+-1	III	60
	S																						V	85
Festuca drymeja	C													+	-	+	+	-	-	-	-	+	I	15
Gagea lutea	C	+	+	-	+	-	-	-	-	+	-	-	-	-	-	-	+	-	+	-	-	+	II	30
Galanthus nivalis	C	-	+	+	1	1	+	1	1	+	+	+	1	+	+	+	+	+	+	+	1	+-1	V	95
Galeobdolon luteum s. l.	C	2	2	3	3	3	2	3	3	2	3	3	2	2	4	4	4	2	4	3	3	2-4	V	100
Galeopsis speciosa	C	+	-	+	-	-	-	-	-	-	-	+	-	-	-	-	-	-	-	-	-	+	I	15
Galium odoratum	C	+	2	1	+	1	-	1	1	+	+	1	1	+	+	1	+	1	-	1	1	+-2	V	90
Geranium phaeum	C	-	-	-	-	-	-	-	-	-	-	+	-	-	-	-	-	-	-	-	-	+	I	5
Hedera helix	Al	-	+	-	-	-	-	-	-	-	-	+	-	-	-	-	-	-	-	1	-	+-1	I	15
	C	+	2	+	+	-	1	+	1	1	+	1	1	1	+	1	+	+	+	1	1	+-2	V	95
	S																						V	95
Hepatica nobilis	C	-	+								+	+	-	-	-	+	-	-	-	1	+	+-1	II	30
Hordelymus europaeus	C	-	+	-	-	-	+	-	-	-	-	-	-	-	+	+						+	I	20
Isopyrum thalictroides	C	-	+	-	+	-	-	+	-	+	-	+	-	-	-	+	-	-	-	-	+	+	II	35
Lathraea squamaria	C	-	+	-	+	-	+	+	+	+	+	-	-	-	+	-	-	+	+	-	-	+	III	50
Lathyrus vernus	C	-	-	-	+	-	-	-	-	-	-	+	-	-	-	-	-	-	-	-	-	+	I	10
Lilium martagon	C	-	-	-	-	-	-	-	-	-	-	+	-	-	-	-	-	-	-	-	-	+	I	5
Mercurialis perennis	C	-	1	-	1	-	1	2	1	1	+	1	2	+	+	1	+	-	+	1	1	+-2	IV	80
Milium effusum	C	-	-	-	-	-	-	-	-	-	+	-	+	-	-	-	+	-	-	-	+	+	I	20
Moehringia trinervia	C	+	-	+	+	+	+	+	-	-	+	+	+	+	-	+	+	-	+	-	+	+	IV	70
Oxalis acetosella	C	+	+	1	+	+	+	1	1	+	1	1	-	-	-	-	-	-	-	-	-	+-1	III	55
Paris quadrifolia	C	+	+	1	-	+	-	+	+	+	-	+	+	-	+	+	+	-	-	+	+	+-1	IV	70
Prunus avium	B																			+	-	+	I	5
	C	-	+	-	-	-	-	-	-	-	-	-	-	-	-	-	-	+	-	+	-	+	I	15
	S																						I	15
Pulmonaria officinalis	C	+	+	+	+	+	-	+	+	+	+	+	1	-	-	+	+	+	-	+	+	+-1	IV	80
Ranunculus lanuginosus	C	-	-	-	-	-	-	-	-	-	+	-	-	-	-	-	-	-	-	-	-	+	I	5
Rubus hirtus agg.	C	-	1	2	+	2	2	+	1	2	1	2	+	1	+	2	+	1	1	2	2	+-2	V	95
Stachys alpina	C	-	+																			+	I	5
Stachys sylvatica	C	+	-	+	+	+	+	-	+	+	-	1	+	+	+	+	+	-	-	-	+	+-1	IV	70
Tilia platyphyllos	B	-	-	-	-	-	-	-	-	-	-	-	-	2	-	-	-	-	-	-	-	2	I	5
	C	-	-	-	-	-	-	-	-	-	-	-	-	+	-	-	-	-	-	-	-	+	I	5
	S																						I	5
Ulmus glabra	Al	-	-	-	-	-	-	-	-	-	-	-	-	-	1	-	-	-	-	2	-	1-2	I	10
	B	+	+	-	-	-	-	-	+	-	-	+	-	-	-	+	-	-	-	+	-	+	II	30
	C	+	-	+	-	-	+	+	-	-	-	+	-	-	-	+	-	-	-	1	+	+-1	II	40
	S																						III	55
Veronica montana	C	+	+	+	+	+	+	+	+	+	+	+	+	+	+	+	+	+	+	1	+	+-1	V	100
Viola reichenbachiana	C	+	+	+	+	+	+	+	+	+	+	+	+	+	+	+	+	+	+	+	+	+	V	100
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20	A-D
Alno-Padion
C.............+ -	+ -- --	+
C - - - 1 ----+ + + + 1 1 -	- - + - +	+-1
C............+ --	+ ----	+
C-- +............+ -- --	+
Eu-Fagion
C...........+ -- -	+ + +- -	+
Tilio-Acerion
C + + -+ + - - + + + - + 1+ +	1 + - - -+-1
Aremonio-Fagion
C...........+ ---	+ ----	+
C + + + + - + -+ + + + + - + +	+ + + 1 1+-1
C +...........+ --	-- + + -	+
C...........+.......+	+
C + 1......+ --+ + - +	--- + -	+-1
C + + - - + + + + + + - - + - -	- + + + +	+
C.....+..........+ -- -	+
Quercetea pubescentis-petraeae
C -------------- +	--- + -+
A1 - 2 - - - - - - - - - - - - -	- - - - -	2
C - - + + -+ + - - + - - - - +	- + - - -	+ S
Quercion farnetto
A1 - - - - 1 - - 2 - - - 3 - 2 2	3 - 2 - 2	1-3
A2 - - - - - - - - - - - - 1 - -	- - - - -	1
B - 1 - 1 2 1 - + - - + 2 2 3 2	2 - + - 1	+-3
C-1+ + 1- + + + --1+ + 11-- + +	+-1 S
Indifferent
C - - + - + - - - - + + - - + +	+- - - -	+
C 1.............+.....+-1
A2 - - - - - - - - - - - - - - -	1 - - - -	1
B.....1 - + - - + 1 - 2 -	+ - + + -	+-2
C - - + + + + - - - + - + - + +	+ + + - +	+ S
C-- + +......+ - + 1 +	2- + + -+-2
Adventiva (incl. Culta, Subspontanea & Indigena)
C ------------------------------+ -+ --	+
Other
C ------------------------------+ --------+
K %
Carex pendula Chrysosplenium alternifolium Festuca gigantea Rumex sanguineus
Dryopteris affinis
Polystichum aculeatum
Aremonia agrimonoides Helleborus odorus Polystichum setiferum Rosa arvensis Ruscus aculeatus Ruscus hypoglossum Tamus communis
Fraxinus ornus Quercus cerris
Tilia tomentosa
I 10 III 45
Galium aparine Rubus fruticosus agg. Sambucus nigra
Urtica dioica Erigeron annuus Arctium sp.
I
10 10
I	20
IV	65
I	10
V	85 I	20
I	10
II	35 IV	65 I	10
I	10
I	5
II	35 II	40
II	40
I	5 IV	65
IV	70
V	85
II	35 I	10
I	5
II	40
III	60
IV	75 III	45
I	10
I	5
	1	2	3	4	5	6	7	8	9	10
Year, first relevé	2006	2006	2006	2006	2006	2006	2006	2006	2006	2006
Date of first relevé	10.05	08.13	10.05	08.10	08.10	08.15	08.15	08.20	08.20	08.18
Year, second relevé	2007	2007	2007	2007	2007	2007	2007	2007	2007	2007
Date of second relevé	03.17	03.25	03.17	03.05	03.06	03.06	03.06	03.05	03.05	03.07
Altitude m a.s.l.	325	340	305	315	350	325	330	330	325	365
Exposition	NE	E	NE	E	W	SE	N	N, NW	W	S
Declination	30	28	20	15	35	28	15	20	25	28
Upper canopy, cover (%)	70	75	65	60	65	60	75	80	80	70
Lower canopy, cover (%)	10	5	5	10	5	20	-	10	5	5
Shrub layer, cover (%)	0.1	15	4	15	20	5	3	5	5	-
Herb layer, cover (%)	90	70	95	100	100	100	95	100	100	100
Upper canopy, height (m)	27	27	25	27	26	25	26	25	23	26
Lower canopy, height (m)	12	17	13	12	8	12	-	14	13	10
Shrub layer, height (cm)	140	170	150	200	450	200	150	150	150	-
Herb layer, height (cm)	70	40	45	60	50	40	40	40	40	60
	11	12	13	14	15	16	17	18	19	20
Year, first relevé	2007	2006	2006	2007	2007	2008	2006	2006	2006	2006
Date of first relevé	03.15	08.06	08.10	03.22	03.25	06.17	09.05	08.17	09.25	08.20
Year, second relevé	2007	2007	2007	2007	2007	2009	2007	2007	2007	2007
Date of second relevé	10.03	03.11	03.05	10.03	10.03	04.03	03.16	03.06	03.11	03.05
Altitude m a.s.l.	295	350	325	355	360	475	350	335	345	315
Exposition	W, -	N	W	N, -	N, W, -	E	Var.	Var.	N, -	N
Declination	20	15	28	35	40, 30	30	30	20	30	30
Upper canopy, cover (%)	75	80	60	55	70	55	90	55	85	55
Lower canopy, cover (%)	-	5	3	10	-	5	5	5	-	4
Shrub layer, cover (%)	15	60	40	10	15	15	-	0.5	50	15
Herb layer, cover (%)	90	80	100	90	100	90	60	100	75	100
Upper canopy, height (m)	23	30	25	27	30	25	25	30	28	25
Lower canopy, height (m)	-	10	10	15	-	10	15	10	-	7
Shrub layer, height (cm)	200	200	500	400	200	200	-	150	250	500
Herb layer, height (cm)	65	45	60	50	70	50	25	50	45	50
Plot size (m2)	400	400	400	400	400	400	400	400	400	400
Locality: 1, 3, 17: Orfü "Vásáros út"; 2, 10-12, 19: Orfü "Cigány-fold"; 4, 13, 18: Orfü "Szuadó"; 5-9, 20: Orfü "Száraz kút-piheno"; 14: Pécs "Zsidó-
völgy"; 15: Pécs "Lóri"; 16: Pécs "Lyukas-hárs"
Relevés 1-5, 10-20 made by Bátori, Z. (ined.); 6-9 made by Bátori, Z., Erdös, L. (ined.)
Tab. 3: Synoptic table ofthe Vegetation ofthe study area with fidelity values (0 x 100) and diagnostic species.
I.
PmQd
TtFo
AtC
V.
Dolines
VI. SaA
VII. CpA
Festuca heterophylla
Lathyrus niger
Clinopodium vulgare
Tanacetum corymbosum
Sedum telephium subsp. maximum
Prunus spinosa
Trifolium alpestre
Silene viridiflora
Potentina micrantha
Euphorbia cyparissias
Luzula forsteri
Veronica officinal is
Lactuca quercina
Hieracium sabaudum
Poa nemoralis
Astragalus glycyphyllos
Brachypodium rupestre
Lychnis coronaria
Poa pratensis
Trifolium rubens
Vincetoxicum officinale
Pyrus pyraster
Chamaecytisus supinus
Securigera varia
Galium schultesii
Lysimachia punctata
Serratula tinctoria
Fallopia dumetorum
Car ex fiacca
Euphorbia epithymoides Genista ovata subsp. nervata Rosa gallica Vicia tetrasperma Convallaria majalis Carexdivulsa Galium mollugo Arabis turrita
Calamintha sylvatica subsp. sylvatica
Buglossoides purpureo-coerulea
Cystop ter i s frag il i s
Tilia platyphyllos
Campanula trachelium
Viola odorata
Campanula rapunculoides
Asperula taurina
Scutellaria altissima
Heracleum sphondylium
Dryopteris carthusiana
Athyrium fitix-femina
Atropa bella-donna
Paris quadrifolia
91.4
91.4
78.1
76.5
71.2 70.7 68.0 67.7
64.2 63.9 61.1 60.4 60.0 57.4 56.9
56.3 56.3 56.3 56.3
56.3
53.4 52.7 51.9 51.9 51.9 51.9 51.9
51.5 47.2 47.2 47.2 47.2 47.2 47.1 45.5 45.1
37.2
32.7
31.3
37.5
32.9
89.6
60.7
53.4
53.1 46.7
46.2 46.2
45.5
50.1 47.8 46.8
64.2 57.4 50.1 46.8
I.	II.	III.	V.	VI.	VII. _PmQd_TtFo_AtC_Dolines_SaA_CpA
Asplenium scolopendrium	—	—	—	—	77.9
Lunaria rediviva	—	—	—	—	68.0
Cardamine amara	—	—	—	—	64.2
Polystichum aculeatum	—	—	—	37.5	56.2
Silene dioica	—	33.6	—	—	51.7
Robinia pseudo-acacia	—	—	—	—	51.5
Oxalis acetosella	—	—	—	—	46.1
Alnus glutinosa	—	—	—	—	—	94.4
Persicaria mitis	—	—	—	—	—	91.4
Calystegia sepium	—	—	—	—	—	84.9
Salix fragilis	—	—	—	—	—	79.9
Equisetum arvense	—	—	—	—	—	78.2
Rubuscaesius	—	—	—	—	—	75.7
Myosoton aquaticum	—	—	—	—	—	74.8
Poa trivial is	—	—	—	—	—	71.6
Ranunculus repens	—	—	—	—	37.0	71.3
Eupatorium cannabinum	—	—	—	—	—	70.7
Lycopus europaeus	—	—	—	—	—	70.7
Solanum dulcamara	—	—	—	—	—	69.6
Angelica sylvestris	—	—	—	—	—	68.0
Festuca gigantea	—	—	—	—	—	66.3
Petasites hybridus	—	—	—	—	—	64.3
Symphytum officinale	—	—	—	—	—	64.3
Equisetum telmateia	—	—	—	—	—	64.2
Knautiadrymeia	—	—	—	—	—	64.2
Arctium minus	—	—	—	—	—	62.2
Berula erecta	—	—	—	—	—	60.4
Humuluslupulus	—	—	—	—	—	60.4
Aethusa cynapium	—	—	—	—	—	59.0
Ranunculus lanuginosus	—	—	—	—	39.1	57.4
Lythrum salicaria	—	—	—	—	—	56.3
Galeopsis speciosa	—	—	—	—	—	55.1
Carexremota	—	—	—	—	38.0	53.7
Adoxa moschatellina	—	—	—	—	—	53.1
Epilobium lanceolatum	—	—	—	—	—	51.9
Carexpendula	—	—	—	—	—	51.6
Cornus sanguinea	—	—	—	—	—	48.4
Galium aparine	—	—	—	—	—	48.2
Urtica dioica	—	—	—	—	31.0	47.7
Veronica beccabunga	—	—	—	—	—	46.8
Corylus avellana___2^_™_™_46.2
Tota! number of species (in the 20 relevés)	200	174	154	114	174	215
Number of diagnostic species	36	8	3	4	7	34
Proportion of diagnostic species (%)	18	5	2	4	4	16
Abbreviations:
PmQd: Potentillo micranthae-Quercetum dalechampii; TtFo: Tilio tomentosae-Fraxinetum orni; AtC: Asperulo taurinae-Carpinetum; HoF: Helle-boro odori-Fagetum; SaA: Scutellario altissimae-Aceretum; CpA: Carici pendulae-Alnetum; Var.: variable, Al: upper canopy layer, A2: lower canopy layer, B: shrub layer, C: herb layer, S: sum.
The use of N values (Tab. 1) with binary data shows that the maximum is at N4 (plants of submesotrophic habitats) and N5 (plants of mesotophic habitats) in turkey oak-sessile oak forests, at N5 in rock forests, oak-hornbeam forests and beech forests, at N7 (plants of soils rich in minerai nitrogen) in dolines, ravine forests and aider forests. The proportion of N5 species is considerable in ali of the units, while N6 (plants of moderately nutrient rich habitats) and N7 species show similarly important values except for the turkey oak-sessile oak forests. When using the weighted data, the maximum is at N4 in the case of turkey oak-sessile oak forests, while concerning ali the other units the maximum is found at higher values than the binary data.
in the case of T indicator values, the proportion of T5 (montane mesophilous broad-leaved forest belt) or T6 (submontane broad-leaved forest belt) species is the highest in the végétation units, while the R indicator values show the maximum at R6 (mostly on neutral soils but also in acidic and basic ones, generally widely tolerant, more or less indifferent plants) or R7 (basifrequent plants, mostly on basic soils). According to the weighted data, the rate of R8 (basiphilous plants) in rock forests and R5 (plants of slightly acid soils) in turkey oak-sessile oak forests is also considerable.
IMPORTANCE OF THE VEGETATION IN NATURE CONSERVATION
The bottom of the dolines in Western Mecsek is not influenced much by human activity. Besides their végétation, their near-natural state is shown by the presence of the fallen, mossy and decayed trees. Several protected species were detected in the lower part of the dolines. These are: Aconitum vulparia, Aremonia agrimonoides, Dryopteris affinis, Dryopteris carthusiana, Dryopteris dilatata, cf. Dryopteris expansa, Helleborus odorus, Hepatica nobilis, Galanthus nivalis, Lïlium martagón, Polystichum aculea-tum, Polystichum setiferum, Ruscus aculeatus, Ruscus hy-poglossum, Stachys alpina, Tamus communis. Dryopteris affinis, cf. Dryopteris expansa and the relict Stachys alpina are rarities not only in the dolines but also in the whole country. Dryopteris affinis - a new species of the Mecsek Mts. - was recently discovered in the dolines and valleys of Western Mecsek (Bátori et al. 2006). Some other protected plants (Aruncus dioicus, Asperula taurina, As-plenium scolopendrium, Carex strigosa, Chaerophyllum aureum, Epipactis helleborine, Iris gramínea, Lathyrus venetus, Lunaria rediviva, Neottia nidus-avis, Primula vulgaris, Silene dioica) occur also in the slopes and edges of these depressions.
discussion and conclusion
The deep valleys and the karst depressions contribute to the geomorphological and végétation diversity of the landscape of the Western Mecsek. In this study, the végétation and habitat conditions were investigated in the lower part of the dolines by the Braun-Blanquet method.
The multivariate analysis shows remarkable simi-larities among the relevés made in the dolines in Western Mecsek. Considering the species composition, the végétation of the dolines resembles mainly the beechwoods and ravine forests.
The canopy layer of the dolines resembles that of the local ravine forests, in which Acer pseudoplatanus, Car-pinus betulus, Fagus sylvatica, Fraxinus excelsior, or Tilia tomentosa are dominant (Kevey 1993,1997). in contrary, the species composition of the herb layer of the dolines is markedly different. Many Fagetalia, Querco-Fagea and Tilio-Acerion elements are missing from the dolines that appear in the ravine forests. The végétation of the dolines differs also from the beech forests of the Mecsek Mts. Several fern species and wet woodland species are missing from the beech forests that are typical in dolines.
Athyrium filix-femina, Atropa bella-donna, Dryopteris carthusiana and Paris quadrifolia are the most important differential species of the dolines.
Hoyk ( 1999a, b, 2002) and Hoyk and Keveiné Bárány (2000) studied the habitat conditions and the végétation of the karst of Western Mecsek with soil analyses and ecological indicator values. They found that the majority of the species indicate a deciduous forest climate, togeth-er with a sub-mediterranean deciduous forest climate, and a mediterranean, atlantic evergreen forest climate. According to moisture supply, fresh, moderately fresh, and moderately wet conditions are indicated by most of the plants, while the distribution of soil reaction values shows near neutral and moderately calciphilous conditions. Our study revealed that the plant communities characteristic of the karst surface of Western Mecsek are arranged along a moisture and nutrient gradient. Turkey oak-sessile oak forests grow in the driest habitats with the highest rate of W3, W4 and W5 species in the indicator spectra. Rock forests and oak-hornbeam forests prefer moister habitats and, along the gradient, they are followed by beech forests, dolines and ravine forests. Al-
der forests occur in the moistest habitats with the highest proportion of W7, W8 and W9 species in the indicator spectra. We can see a very similar arrangement between the habitats with the highest (aider forests) and the lowest (turkey oak-sessile oak forests) nitrogen supply. In this system, the habitat conditions of the dolines are similar to those of the beech forests and the local ravine forests: fresh and relatively rieh in nutrients.
Due to the special funnel and bowl shape of the dolines, thermal inversion is evident, which results in sudden changes in the vegetation. This phenomenon causes the inversion of the zones of pine forests, dwarf pine thickets and alpine grasslands in the dolines of South-Eastern Alps (Borhidi 2002), or the séparation of the fresh Festuco ovinae-Nardetum from the dry Festuce-tum sulcatae in the dolines in Bükk Mountain (North Hungary; Bacsó & Zólyomi 1934). Some plant associations of the Slovenian Karst (Asaro-Carpinetum betuli, Ornithogalo pyrenaici-Carpinetum) also occur in the
bottom of the dolines that are cooler and more humid than their environment (Kranjc 1997). Because of the climate-balancing effect of the forests, this phenomenon is not so striking in Mecsek Mts. (Lehmann 1970). In the karst, which is dominated by oak-hornbeam and beech forests, effects of the thermal inversion are the most spectacular where beech forests follow turkey oak-sessile oak forests and oak-hornbeam forests on the lower part of the doline slopes. The described vegetation type in the lower part of these depressions is developed by edafic factors, mainly by increased soil humidity.
Considering the vegetation texture and species composition, the vegetation of the dolines in Western Mecsek resembles mainly the local beechwoods and ravine forests. This vegetation type occurs only in small patches, its stands are disjunct and isolated fragments in the matrix of Illyrian beech and oak-hornbeam woods. Distinction of the vegetation of the dolines as a separate association is not supported by the analyses.
acknowledgement
The Authors are indebted to the Department of Plant the Department of Ecology of the University of Szeged Taxonomy and Geobotany of the University of Pécs, and for their indispensable help.
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