OPCE-STR.vp Acta Bot. Croat. 70 (2), 147–155, 2011 CODEN: ABCRA 25 ISSN 0365-0588 Ecological conditions, flora and vegetation of a large doline in the Mecsek Mountains (South Hungary) ZOLTÁN BÁTORI*, RÓBERT GALLÉ, LÁSZLÓ ERDÕS, LÁSZLÓ KÖRMÖCZI Department of Ecology, University of Szeged, 6726 Szeged, Közép fasor 52, Hungary Abstract – Vegetation-environment relationships were investigated in a large doline of the Mecsek Mts (South Hungary). To reveal the vegetation pattern, we collected vegeta- tion data and environmental variables along a 243 m long transect. A total of 144 vascular plant species and 4 vegetation types were identified in the doline. We found that both the species composition and the vegetation pattern are significantly influenced by air tempera- ture, air humidity, soil moisture and altitude. Our results confirm the putative temperature and vegetation inversion in the doline. Keywords: direct gradient analysis, doline, flora, vegetation types, vegetation inversion, redundancy analysis, species composition, Mecsek Mountains, Hungary Introduction Karst surfaces and environments are extremely vulnerable to degradation and pollution (CALÓ and PARISE 2006, BREG 2007), thus they are currently in the focus of research and conservation efforts. Dolines are funnel- and bowl-shaped closed depressions formed by water infiltration, which range from a few meters to a few hundred meters in diameter and depth (VERESS 2004). At night, cold-air lakes develop in the depressions (BÁRÁNY-KEVEI 1999), resulting in low air temperatures and high air humidities, which in turn strongly influences the compo- sition of local flora and vegetation (BECK-MANNAGETTA 1906, MORTON 1936, ÖZKAN et al. 2010). Many karst depressions around the world have developed into excellent refuge ar- eas for relict, mountain (HORVAT 1953) and endemic species (EGLI et al. 1990, BRULLO and GIUSSO DEL GALDO 2001), which play a central role in the knowledge about vegetation his- tory. The study was carried out in the karst area of 30 km2 in the Mecsek Mountains (South Hungary), near the city of Pécs. Sub-Mediterranean type, middle-aged oak-hornbeam (Asperulo taurinae-Carpinetum Soó et Borhidi in Soó 1962) and beech forests (Helleboro odori-Fagetum Soó et Borhidi in Soó 1960) dominate the present vegetation of the plateaus ACTA BOT. CROAT. 70 (2), 2011 147 *Corresponding author, e-mail: zbatory@gmail.com Copyright® 2011 by Acta Botanica Croatica, the Faculty of Science, University of Zagreb. All rights reserved. U:\ACTA BOTANICA\Acta-Botan 2-11\Batori.vp 8. rujan 2011 15:57:15 Color profile: Disabled Composite 150 lpi at 45 degrees and slopes of the study site. The bottom of larger valleys is covered by beech forests or ra- vine forests (Scutellario altissimae-Aceretum (Horvát 1958) Soó et Borhidi in Soó 1962). This latter vegetation type is also found in the deeper and larger dolines of the Mecsek Mountains (cf. BÁTORI et al. 2009). The purpose of this study was to analyse the vegetation pattern of the herb layer in a doline of the Mecsek Mountains. The following questions were addressed: How many vege- tation types and vascular plant species can be identified in the doline? What environmental variables influence the vegetation pattern on the slopes? 148 ACTA BOT. CROAT. 70 (2), 2011 BÁTORI Z., GALLÉ R., ERDÕS L., KÖRMÖCZI L. U:\ACTA BOTANICA\Acta-Botan 2-11\Batori.vp 8. rujan 2011 15:57:21 Color profile: Disabled Composite 150 lpi at 45 degrees Material and methods Our surveys were conducted in a large (diameter > 200 m) and deep (depth > 25 m) doline of the Mecsek Mountains. The herb layer was sampled along a 2 m wide and 243 m long transect consisting of 1 m × 2 m contiguous plots (Figs. 1, 2). The transect was estab- lished in a north-south direction traversing the deepest point of the depression. Each plot was surveyed twice: in summer 2008 and spring 2009. In addition, a complete plant species list was compiled for the total area of doline. The total area included the area of the slopes (where the slope angle was over 10°) and the area of the edge (an approximately 15–20 m wide stripe around the doline where the slope angle was less than 10°). ACTA BOT. CROAT. 70 (2), 2011 149 FLORA AND VEGETATION OF A LARGE DOLINE Fig. 1. Species occurrences and relief profile along the doline transect. Numbers on the right indi- cate plant species as follows: 1 – Fraxinus excelsior L., 2 – Cardamine bulbifera (L.) Crantz, 3 – Allium ursinum L., 4 – Tilia tomentosa Moench, 5 – Arum maculatum L., 6 – Asarum europaeum L., 7 – Hedera helix L., 8 – Alliaria petiolata (M. B.) Cavara et Grande, 9 – Galium aparine L., 10 – Veronica hederifolia L., 11 – Helleborus odorus W. et K., 12 – Viola reichenbachiana Jord., 13 – Acer platanoides L., 14 – Acer campestre L., 15 – Fraxinus ornus L., 16 – Stellaria holostea L., 17 – Carex pilosa Scop., 18 – Melica uniflora Retz., 19 – Dactylis polygama Horvátovszky, 20 – Polygonatum multiflorum (L.) All., 21 – Quercus petraea (Mattuschka) Lieblein, 22 – Euphorbia amygdaloides L., 23 – Ligustrum vulgare L., 24 – Rosa arvensis Huds., 25 – Lathyrus vernus (L.) Bernh., 26 – Galium schultesii Vest, 27 – Hepatica nobilis Mill., 28 – Crataegus laevigata (Poir.) DC., 29 – Bromus ramosus Huds. agg., 30 – Fallopia dumetorum (L.) Holub, 31 – Quercus cerris L., 32 – Fagus sylvatica L., 33 – Viola alba Bess., 34 – Carpinus betulus L., 35 – Festuca drymeja M. et K., 36 – Lathyrus venetus (Mill.) Wohlf, 37 – Campanula rapunculoides L., 38 – Symphytum tuberosum L. subsp. nodosum (Schur), 39 – Convallaria majalis L., 40 – Iris graminea L., 41 – Glechoma hirsuta W. et K., 42 – Ajuga reptans L., 43 – Clinopodium vulgare L., 44 – Taraxacum officinale Weber ex Wiggers, 45 – Sorbus torminalis (L.) Cr., 46 – Luzula luzuloides (Lam.) Dandy et Wilm., 47 – Luzula forsteri (Sm.) DC., 48 – Primula vulgaris Huds., 49 – Hordelymus europaeus (L.) C. O. Harz, 50 – Mycelis muralis (L.) Dum., 51 – Tamus communis L., 52 – Milium effusum L., 53 – Geum urbanum L., 54 – Stachys alpina L., 55 – Euonymus europaeus L., 56 – Galium odoratum (L.) Scop., 57 – Arabis turrita L., 58 – Geranium robertianum L., 59 – Moehringia trinervia (L.) Clairv., 60 – Cardamine im- patiens L., 61 – Isopyrum thalictroides L., 62 – Galeobdolon luteum Huds. s.l., 63 – Ruscus hypoglossum L., 64 – Galanthus nivalis L., 65 – Cardamine enneaphyllos (L.) Crantz, 66 – Anemone ranunculoides L., 67 – Mercurialis perennis L., 68 – Pulmonaria officinalis L., 69 – Fragaria vesca L., 70 – Carex divulsa Stokes, 71 – Pyrus pyraster (L.) Burgsdorf, 72 – Hypericum hirsutum L., 73 – Veronica chamaedrys L., 74 – Erigeron annuus (L.) Pers., 75 – Cornus sanguinea L., 76 – Rumex sanguineus L., 77 – Dryopteris affinis (Löwe) Fras.-Jenk., 78 – Rubus hirtus W. et K. agg., 79 – Clematis vitalba L., 80 – Scrophularia nodosa L., 81 – Solanum dulcamara L., 82 – Paris quadrifolia L., 83 – Dryopteris carthusiana 8Vill.) H. P., 84 – Rubus fruticosus L. agg., 85 – Veronica montana L., 86 – Aconitum vulparia Rchb., 87 – Sambucus nigra L., 88 – Brachypodium sylvaticum (Huds.) Roem. et Schult., 89 – Carex sylvatica Huds., 90 – Ranunculus ficaria L., 91 – Polystichum setiferum (Forskål) Woynar, 92 – Corydalis cava (L.) Schw. et Koerte, 93 – Epilobium montanum L., 94 – Gagea lutea (L.) Ker-Gawl., 95 – Polystichum aculeatum (L.) Roth, 96 – Stachys sylvatica L., 97 – Eupatorium cannabinum L., 98 – Dryopteris filix-mas (L.) Schott, 99 – Atropa bella-donna L., 100 – Athyrium filix-femina (L.) Roth, 101 – Acer pseudoplatanus L., 102 – Circaea lutetiana L., 103 – Urtica dioica L. � U:\ACTA BOTANICA\Acta-Botan 2-11\Batori.vp 8. rujan 2011 15:57:21 Color profile: Disabled Composite 150 lpi at 45 degrees Presence-absence data were analysed using hierarchical classification (HC; Complete link, Sorensen index) to identify the vegetation types on the slopes. However, field obser- vations were also performed to specify the result of the classification. Cluster analysis was done with the program package SYN-TAX 2000 (PODANI 2001). Air temperature (°C) and humidity (%) were measured for 24 hours by SN21140CA sensors 25 cm above the ground surface in 50 sampling plots at 5 m intervals. Spectrum – Fieldscout TDR-300 was used to detect soil moisture values (%; in 12 cm depth) at the same 50 plots. All measures were carried out in summer, after a dry period, under clear weather conditions. Slope angles were also measured along the transect, from which alti- tude values were calculated for each plot. We performed constrained ordinations using the Vegan R package (OKSANEN et al. 2009, R development core team 2009) and SYN-TAX 2000 to determine the main environ- mental parameters affecting the distribution pattern of plant species along the transect. We 150 ACTA BOT. CROAT. 70 (2), 2011 BÁTORI Z., GALLÉ R., ERDÕS L., KÖRMÖCZI L. Fig. 2. Cover of the 12 most abundant plant species along the transect. U:\ACTA BOTANICA\Acta-Botan 2-11\Batori.vp 8. rujan 2011 15:57:23 Color profile: Disabled Composite 150 lpi at 45 degrees used a series of the linear ordination method, redundancy analysis (RAO 1964) to identify parameters that explain significant variation of species. We calculated the marginal and conditional effect of each term and assessed their significance using Monte-Carlo permuta- tion tests with 5000 permutations (e.g. MUFF et al. 2009, GALLÉ and TORMA 2009). The fol- lowing data were used in the analysis: presence-absence data of species, mean air tempera- ture, mean air humidity, soil moisture and altitude values (Fig. 3). For interpretability, data of the summer aspect were used only in the redundancy analysis. Figures were prepared with Microsoft EXCEL and Adobe Photoshop CS2. Plant com- munity names were used according to BORHIDI (2003), while the names of plant taxa follow SIMON (2000). Results A total of 144 vascular plant species were detected in the doline (Fig. 1, Tab. 1). We found a strong correlation between species composition and doline morphology. Some spe- cies occur in every part of the doline (e.g. Asarum europaeum, Cardamine bulbifera, Fraxinus excelsior), while others occur only on the south-facing slope (e.g. Clinopodium vulgare, Festuca drymeja, Lathyrus venetus), the north-facing slope (e.g. Arabis turrita, Isopyrum thalictroides, Milium effusum) or in the doline bottom (e.g. Dryopteris affinis, Polystichum aculeatum, Solanum dulcamara). From a floristic point of view, the doline ACTA BOT. CROAT. 70 (2), 2011 151 FLORA AND VEGETATION OF A LARGE DOLINE Fig. 3. Redundancy analysis ordination diagram with 50 plots (�: plots of the south-facing edge, �: plots of the south-facing slope, �: plots of the north-facing edge, �: plots of the north-fac- ing slope, �: plots of the doline bottom), environmental variables (arrows) and the most fre- quent plant species (italic numbers 1, 8, 15, 17–19, 62, 67, 103, according to Fig. 1) of the plot groups. U:\ACTA BOTANICA\Acta-Botan 2-11\Batori.vp 26. rujan 2011 13:54:59 Color profile: Disabled Composite 150 lpi at 45 degrees bottom is the most important, because it preserves the highest number of mountain species (e.g. Aconitum vulparia, Actaea spicata, Asplenium scolopendrium, Dryopteris affinis and Polystichum aculeatum). The glacial relict plant Stachys alpina was found only on the up- per part of the north-facing slope. Species cover is also strongly correlated with slope posi- tion and exposition along the transect (Fig. 2). For example, Festuca drymeja is abundant only on the south-facing slope, Allium ursinum and Cardamine enneaphyllos on the north-facing slope and in the doline bottom, while Galeobdolon luteum s.l. is abundant only in the doline bottom. Four vegetation types were detected along the transect. A turkey oak-sessile oak forest (Potentillo micranthae-Quercetum dalechampii Horvát A. O. 1981) and an oak-hornbeam forest transition occurs on the south-facing edge (0–43 m), an oak-hornbeam forest on the south-facing slope and on the north-facing edge (44–71 m and 218–243 m), a beech forest on the lower part of the south-facing slope and on the north-facing slope (72–90 m and 146–217 m), and a ravine forest fragment in the bottom of the doline (91–145 m). Summary statistics for redundancy analysis are shown in table 2. The subsequent Monte-Carlo permutation test indicates the significance of both the marginal and the condi- tional effect of the studied parameters (p < 0.001). The redundancy analysis triplot shows that doline vegetation is arranged along a moisture and temperature gradient (Fig. 3). The south-facing edge and south-facing slope are the driest and warmest, while the doline bot- tom is the moistest and coldest. Plots of the north-facing edge and north-facing slope oc- cupy a transitional area in the ordination space. The maximum mean diurnal air tempera- ture was detected in a plot of the south-facing edge (19.87 °C) and the minimum in a plot of the doline bottom (17.56 °C). In contrast, the maximum (57.9%) and minimum (7.3%) soil moisture, and the maximum (93.86%) and minimum (78.58%) mean diurnal air humidity values showed a reverse distribution. 152 ACTA BOT. CROAT. 70 (2), 2011 BÁTORI Z., GALLÉ R., ERDÕS L., KÖRMÖCZI L. Tab. 1. List of plant species outside the doline transect. South-facing edge and slope 104: Allium oleraceum L., 105: Astragalus glycyphyllos L., 106: Buglossoides purpureo-coerulea (L.) I. M. Johnst., 107: Campanula persicifolia L., 108: Chaerophyllum temulum L., 109: Cornus mas L., 110: Crataegus monogyna Jacq., 111: Cruciata glabra (L.) Ehrend., 112: Euonymus verrucosus Scop., 113: Euphorbia cyparissias L., 114: Festuca heterophylla Lam., 115: Galium lucidum All., 116: Lamium maculatum L., 117: Lapsana communis L., 118: Lathyrus niger (L.) Bernh., 119: Lilium martagon L., 120: Lysimachia nummularia L., 121: Lysimachia punctata L., 122: Melittis carpatica Klok., 123: Poa nemoralis L., 124: Potentilla micrantha Ram., 125: Scutellaria altissima L., 126: Silene viridiflora L., 127: Vicia sepium L., 128: Viola odorata L. North-facing edge 129: Asperula taurina L. Doline bottom 130: Aethusa cynapium L., 131: Actaea spicata L., 132: Arctium minus (Hill) Bernh, 133: Asplenium scolopendrium L., 134: Calamagrostis epigeios (L.) Roth, 135: Carex pendula Huds., 136: Cirsium arvense (L.) Scop., 137: Festuca gigantea (L.) Vill., 138: Galeopsis speciosa Mill., 139: Heracleum sphondylium L., 140: Knautia drymeia Heuff., 141: Phytolacca americana L., 142: Staphylea pinnata L., 143: Tilia cordata Mill., 144: Tilia platyphyllos Scop. U:\ACTA BOTANICA\Acta-Botan 2-11\Batori.vp 8. rujan 2011 15:57:26 Color profile: Disabled Composite 150 lpi at 45 degrees Discussion Our study revealed the close relationships between the variation of geomorphological and vegetation pattern in a karst depression at the spatial scale of the study. The most im- portant finding of this study is that the vegetation pattern is strongly correlated with slope exposition and the depth of the doline. A former study by BÁTORI et al. (2009) showed that plant associations of the karst of Mecsek Mountains are arranged along a moisture and nutrient gradient. However, despite being the first detailed description of the doline vegetation and its surroundings of this area, that study only addressed the large-scale vegetation pattern (Braun-Blanquet scale) of the bottom of the dolines, and used exclusively ecological-indicator values to reveal the habitat conditions in the depressions. In contrast, our present study reveals the fine-scale vegetation pattern along a doline transect and shows that there are remarkable differences among the species composition of the south-facing slope, the north-facing slope and the bottom of the doline. This is a conse- quence of the special microclimate that often determines both abiotic and biotic parameters of karst depressions (GEIGER 1950, WHITEMAN et al. 2004, GARGANO et al. 2010, OZIMEC et al. 2010, VRBEK et al. 2010). In Hungary, south-facing slopes receive higher solar radiation (JAKUCS 1971), which affects temperature, air humidity and soil moisture, and ultimately, affects the vegetation. In the case of the investigated doline, mixed-oak forests cover the major part of the south-facing slope. In contrast, the north-facing slope receives lower solar radiation, which results in lower mean temperatures and lower evapotranspiration. Due to the cooler climate, the north-facing slope and the lower part of the south-facing slope are covered by beech forests. Therefore, a vegetation inversion is formed, which is a common phenomenon in karst depressions (BECK-MANNAGETTA 1906, HORVAT 1953, LAUSI 1964, FAVRETTO and POLDINI 1985). The vegetation pattern also changes in the bottom of the doline, where the special ecological conditions have led to the development of a ravine for- est. Our results are in good agreement with those of HOYK (1999), who pointed out that dolines of the Mecsek Mts play an important role in the landscape due to their characteristic shape and valuable flora. Considering the special microclimatic conditions, geomorphological features, vegeta- tion pattern and species composition, dolines are especially important and valuable for sci- entific research and nature conservation. ACTA BOT. CROAT. 70 (2), 2011 153 FLORA AND VEGETATION OF A LARGE DOLINE Tab. 2. Summary statistics for redundancy analysis with the four environmental variables. Axes 1 2 3 4 Eigenvalues 0.110 0.044 0.038 0.029 Species-environment correlations 0.9305 0.8834 0.8298 0.8047 Cumulative % variance of species data 11.0 15.4 19.2 22.1 of species-environment relationship 50 69.7 86.8 100 Sum of all eigenvalues: 1 Sum of all canonical eigenvalues: 0.22 U:\ACTA BOTANICA\Acta-Botan 2-11\Batori.vp 8. rujan 2011 15:57:26 Color profile: Disabled Composite 150 lpi at 45 degrees Acknowledgement We would like to thank András Bíró, Sándor Csete, Miklós Maróti, Tamás Morsch- hauser, Csaba Németh and János Podani for the useful comments and suggestions. This research was supported by the TÁMOP-4.2.2/08/1/2008-0008 program of the Hungarian National Development Agency. References BÁRÁNY-KEVEI, I., 1999: Microclimate of karstic dolines. Acta Climatologica Universitatis Szegediensis 32–33, 19–27. 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