DOI: 10.2478/HACQ-2013-0010 HACQUETIA 12/2 • 2013, 23-37 black alder dominated forest vegetation in the western part of central Slovakia - species composition and ecology Richard HRIVNÂK1*, Jaroslav KOSTÂL2, Michal SLEZÀK1, 3, Anna PETRÂSOVÂ4 & Melania FESZTEROYÀ5 Abstract In some regions of Slovakia, black alder forest vegetation has not been documented appropriately yet. This paper is the first vegetation study presenting the phytosociological data and measured environmental parameters from the western part of central Slovakia. The data set was classified by using a modified TWINSPAN algorithm, which allowed us to discern floristically and ecologically distinctive plant communities. They correspond to the associations Stellario nemorum-Alnetum glutinosae Lohmeyer 1957 (riparian alder vegetation on mesic to humid sites along small brooks) and Carici acutiformis-Alnetum glutinosae Scamoni 1935 (eutrophic black alder carr forests in the colline zone) with the variants of Ligustrum vulgare and Galium palustre. The community Carici elongatae-Alnetum glutinosae Schwickerath 1933 (mesotrophic to eutrophic alder carr vegetation growing on permanently waterlogged soils), documented only with two phytosociological relevés, was distinguished following expert knowledge. A floristic and ecological pattern of these associations is presented. The major compositional gradients were interpreted based on Ellenberg's indicator values and the values of environmental variables recorded during the field sampling in the growing season 2011. The principal component analysis revealed the importance of soil moisture, light availability, portion of open water and soil surface for species composition variability at the association level, whereas the variants of Carici acutiformis-Alnetum glutinosae were sorted along the acidity gradient. Key words: Alnion glutinosae, Alnion incanae, Ellenberg indicator values, forest vegetation, soil variables, syn-taxonomy. Izvleček Gozdna vegetacija črne jelše dosedaj v nekaterih predelih Slovaške še ni bila dovolj raziskana. Članek predstavlja prvo vegetacijsko raziskavo fitocenoloških popisov in merjenih rastiščnih dejavnikov iz zahodnega dela osrednje Slovaške. Podatkovni niz smo klasificirali z modificiranim TWINSPAN algoritmom, s čemer smo lahko floristično in ekološko ločili značilne rastlinske združbe. Ugotovili smo asociaciji: Stellario nemorum--Alnetum glutinosae Lohmeyer 1957 (obvodna črnojelševja na srednje do vlažnih rastiščih ob manjših potokih) in Carici acutiformis-Alnetum glutinosae Scamoni 1935 (evtrofna črnojelševja v kolinskem pasu) z dvema variantama Ligustrum vulgare in Galium palustre. Asociacijo Carici elongatae-Alnetum glutinosae Schwickerath 1933 (mezotrofni do eutrofni črnojelševi grezi, ki uspevajo na tleh stalno prepojenih z vodo), ki je dokumentirana samo z dvema popisoma smo določili na podlagi strokovnega poznavanja. Prikazali smo floristični in ekološki vzorec pojavljanja teh asociacij. Glavni gradienti so pojasnjeni na podlagi Ellenbergovih indikatorskih vrednosti, ekološki dejavniki pa so bili merjeni med terenskim vzorčenjem v vegetacijski sezoni 2011. Analiza glavnih komponent je razkrila pomen vlažnosti tal, svetlobe, deleža odprtih vodnih in talnih površin za variabilnost vrstne sestave na nivoju asociacij, medtem ko se varianti asociacije Carici acutiformis-Alnetum glutinosae ločita na gradientu kislosti. Ključne besede: Alnion glutinosae, Alnion incanae, Ellenbergove indikatorske vrednosti, gozdna vegetacija, talne spremenljivke, sintaksonomija. 1 Institute of Botany, Slovak Academy of Sciences, Dubravska cesta 9, SK-845 23 Bratislava, Slovak Republic; e-mail: richard.hrivnak@savba.sk; 2 State Nature Conservancy of Slovak Republic, Samova 3, SK-949 01 Nitra, Slovak Republic; e-mail: kostaljar@gmail.com 1. INTRODUCTION Forest vegetation with dominance of black alder (Alnus glutinosa (L.) Gaertn.) is common on swampy soils with high groundwater table, or it creates more mesophilous stands in alluvial zones along rivers and brooks (Oberdorfer 1953, Douda 2008, Ellenberg 2009, Sburlino et al. 2011). The understorey of these forests in general consists of species having diverse moisture-requirements. It is composed of either mesophilous forest plants or marshland and wet species. Apparent environmental differentiation, considerable floristical variability and inconsistent physiognomy indicate existence of various alder forests with different syntaxonomical positions. At the level of phy-tosociological alliances, the major types of alder vegetation include hygro- or meso-hygrophilous streamside forests (Alnion incanae, class Querco-Fagetea) and alder carr forests (Alnion glutinosae, class Alnetea glutinosae). The syntaxonomical concept of two separate alliances has also been recognized in the central Slovak synthesis (Slezák et al. 2013) representing a preliminary revision of alder vegetation in Slovakia. It uses a more formalized classification method (cf. Chytry 2000) and provides further evidence for uneven distribution of phytosocio-logical material in Western Carpathians. Except for few vegetation analyses (Somsák 1961, Kár-páti et al. 1963, Miadok 1978, Balázs 1996), most of the relevés collected in the past were published in local studies containing not only data of alder-dominated forests (e.g. Berta 1970, Neuhauslová-Novotná 1970, Kontris 1981). Recently, there has arisen an increased interest in many aspects of their syntaxonomy and synecology (Somsák 2000, Kollár et al. 2005, Hrivnák et al. 2009, Slezák et al. 2011, 2013). Petrásová et al. (2011) devoted a special attention to a detailed analysis of bryophyte species composition, as the moss layer plays an important role in the community structure of these vegetation types. Local phytosociological studies offering new relevés are thus significant tool for compilation of national data set, as there are still territories without any or with only few available relevés. This was primary reason for investigation of alder forest variability in the Poni-trie region (Figure 1) which is characterized only by negligible and descriptive information about such vegetation (cf. Elias 1980, 2008). The purpose of our vegetation study was (i) to identify the main types of alder forests in the study area by field sampling and numerical analysis, and (ii) to find environmental drivers responsible for variation in their species composition. 2. METHODS We studied black alder dominated forests in the western part of central Slovakia (Ponitrie region; Figure 1) following the principles of the Zürich-Montpellier approach (Westhoff & van der Maarel 1973). Phytosociological sampling of 19 vegetation plots was carried out in valleys consisting of diverse geological substrates in 2011. The cover of vascular plants and bryophytes was recorded using the Braun-Blanquet sampling scale extended by Barkman et al. (1964). Phytosociological relevés were stored in a TURBOVEG database (Hennekens & Schaminée 2001) and exported into Juice program (Tichy 2002). Modified TWINSPAN algorithm (Rolecek et al. 2009) was run for numerical classification with three pseudospecies cut levels (0%, 5%, 25%) and total inertia as a measure of cluster heterogeneity. The TWINSPAN division was accepted at the 3-clus-ter level following the outcomes of crispness curve method (Botta-Dukat et al. 2005). This partition produced clusters which closely matched the established phytosociological units. The same species recorded in various layers (herb - E1S shrub - E2 and tree - E3) were merged prior to the analysis. Lists of diagnostic species for individual clusters were derived on the basis of frequency concept, i.e. the species considered as diagnostic needed to display a frequency above 50% in particular cluster, and the difference in frequencies among the clusters more than 40%. Two relevés 8 Faculty of Education, Catholic University, Hrabovska cesta 1, SK-034 01 Ruzomberok, Slovak Republic; e-mail: slezak. miso@gmail.com 4 Faculty of Natural Sciences, Matej Bel University, Tajovskeho 40, SK-974 01 Banska Bystrica, Slovak Republic; e-mail: anniepetrasov@gmail.com 5 Faculty of Natural Sciences, Constantine the Philosopher University in Nitra, A. Hlinku 1 SK-94901 Nitra; Slovak Republic; e-mail: mfeszterova@ukf.sk * Corresponding author. with partially outlying species composition (co-dominance of Carex elongata in the herb layer), which were merged with the floristically closest cluster in the numerical classification, were arbitrarily arranged in the text in order to simplify the results. The expert knowledge was used for their syntaxonomical interpretation. Subsequently, all the next analyses were performed without them. The species-environmental relationships were quantified and tested by the linear ordination method (Principal component analysis; PCA) in CANOCO for Windows 4.5 package (ter Braak & Smilauer 2002), as the length of gradient in the Detrended correspondence analysis was 2.56 SD units. Non-weighted Ellenberg indicator values (EIV) for vascular plants (Ellenberg et al. 1992) and selected environmental data (see Table 1) for relevés were plotted into the PCA ordination diagrams as supplementary variables and correlated to the relevé scores on the first two ordination axes using the Pearson correlation coefficient. The one-way ANOVA and post-hoc Tukey HSD test was applied for multiple comparisons of environmental parameters in the STATISTICA software (StatSoft 2001). Soil samples were randomly taken in three places in sampling plot from the uppermost mineral horizon (0-10 cm depth, litter removed) and mixed to form a single sample per plot in order to reduce the soil heterogeneity. They were dried at a laboratory temperature, crushed and passed through a 2 mm sieve (Slezâk et al. 2011). Soil pH —"ï* - J Figure 1: Distribution of phytosociological relevés of alder-dominated forests in the western part of central Slovakia. Slika 1: Porazdelitev fitocenoloških popisov gozdov v katerih dominira črna jelša v zahodnem delu osrednje Slovaške. o -1.0 0.8 -1.0 0.8 Figure 2: PCA ordination diagrams. The first two ordination axes explain 22.6% and 13.5% of the total species variability. A) Distribution of diagnostic species for particular units in ordination space (normal bold - Stellario nemorum-Alnetum glutinosae; italic - Carici acutiformis-Alnetum glutinosae, underline normal - variant with Ligustrum vulgare, underline italic - variant with Galium palustre); B, C) Position of sampling sites with environmental factors (EIV, number of all species - B; directly measured or calculated factors - C) displayed as supplementary variables. Empty circles - Stellario nemorum-Alnetum glutinosae, squares - Carici acutiformis-Alnetum glutinosae (empty squares - variant with Ligustrum vulgare, full squares - variant with Galium palustre). Slika 2: PCA ordinacijski diagram. Prvi dve osi pojasnita 22,6% in 13,5% skupne variabilnosti vrst. A) Razširjenost značilnih vrst določenih enot v ordinacijske prostoru (običajno krepko - Stellario nemorum-Alnetum glutinosae; poševno - Carici acutiformis-Alnetum glutinosae, običajno podčrtano - varianta z Ligustrum vulgare, podčrtano poševno - varianta z Galium palustre); B, C) Položaj vzorčenih sesto-jav z rastiščnimi dejavniki (EIV, število vseh vrst - B; neposredno merjeni ali izračunani- C) prikazano kot dopolnilne spremenljivke. Prazni krožci - Stellario nemorum-Alnetum glutinosae, kvadrati - Carici acutiformis-Alnetum glutinosae (prazni kvadrati - varianta z Ligustrum vulgare, polni krožci -varianta z Galium palustre). □ □ □ Moisture Light --- B) Soil Reaction Continentality j/n o No. of all species ^ ■ ■ ■ ■ Temperature Cfo Nutrients O Abbreviations of species names and environmental variables (Okrajšava imen vrst in ekoloških dejavnikov): Angesyl - Angelica sylvestris, Athyfil - Athyrium filix-femina, Cardama - Cardamine amara, Careacu - Carex acutiformis, Carerem - Carex remota, Ceraluc - Cerastium lucorum, Coryave - Corylus avellana, Descces - Deschampsia cespitosa, Dryocar - Dryopteris carthusiana, Eupacan - Eupatorium cannabinum, Fraxexc - Fraxinus excelsior, Galespe - Galeopsis speciosa, Galipal - G. palustre, Humulup - Humulus lupulus, Junceff - Juncus effusus, Lapscom - Lapsana communis, Liguvul - Ligustrum vulgare, Lycoeur - Lycopus europaeus, Lysinem - Lysimachia nemorum, Lysvulg - L. vulgaris, Myospal - Myosotis scorpioides agg., Pershyd - Persicaria hydropiper, Plagund - Plagiomnium undulatum, Poa tri - P. trivialis, Ranurep - Ranunculus repens, Scirsyl - Scirpus sylvaticus, Rubucae - Rubus caesius, Rumesan - Rumex sanguineus, Soladul - Solanum dulcamara, Valedio - Valeriana dioica, Verobec - Veronica beccabunga, Vibuop - Viburnum opulus; Ca_max - maximum concentration of calcium in soil solution, Min_width_ lenght - minimum width or length of the studied stands, %_soil - percentage of open soil, %_water - percentage of open water. was measured in a distilled water solution (soil/ water ratio of 1/2.5) and inorganic N content as sum of N-NH4+ and N-N03" (extracted with K2S04 and determined colorimetrically using a Nessler's reagent and acid phenoldihydrosulphate, respectively). Calcium content was extracted in 10% HCl solution and determined in accordance with the recommendation by Simansky (2010). Altitude and geographical coordinates (WGS 84) were measured in the field with GPS equipment. Percentage cover of open soil (without any vegetation) and open water areas was estimated visually in the field. Since several recent studies have shown the pronounced effect of habitat quality and landscape configuration-related factors on species composition pattern in floodplain forests and/or alder-dominated vegetation (e.g. Hérault & Honnay 2005, Douda 2010, Košir et al. 2013), total area, minimal width or length of the studied alder forests in each sampling site and width of the valley were obtained from GIS layers as well. The nomenclature of bryophytes and vascular plants follows the checklist by Marhold & Hindâk (1998). The names of plant communities and their assignment into the higher units are in accordance with Jarolimek et al. (2008). Full scientific name of vegetation units with the author's name and year of description were used in case when they were not presented in the above-mentioned paper. 3. RESULTS Species composition patterns Numerical classification led to delimitation of two main relevé's groups that correspond to the associations of Stellario nemorum-Alnetum gluti-nosae (cluster A) and Carici acutiformis-Alnetum glutinosae (cluster B; Table 2). They are characterized by distinctive floristic composition with clearly defined lists of diagnostic species. While mesophilous or slightly hygrophilous species are related to the stands of the cluster A, hygrophil-ous and marshland plants are typical for the forests of cluster B (Table 2). The second level of TWINSPAN division separated relevés of cluster B into two floristically well-differentiated variants, namely variant with Ligustrum vulgare and variant with Galium palustre. Similar patterns in composition structure of data set were also found in the ordination analysis (Figure 2). Forest stands recorded within the association Stellario nemorum-Alnetum glutinosae are usually three-layered. The canopy closed tree layer is almost exclusively created by Alnus glutinosa, whereas the shrub layer is most often built up by Acer campestre and Corylus avellana (Table 2). The forest understorey contains species typical for various environmental microhabitats. In more detail, forest mesophilous plants (e.g. Brachypodi-um sylvaticum, Circaea lutetiana, Pulmonaria offici-nalis agg.) constantly grow in combination with more or less hygrophilous (e.g. Caltha palustris, Impatiens noli-tangere) and nitrophilous species (Aegopodium podagraria, Galium aparine, Glecho-ma hederacea, Urtica dioica; Table 2, Figure 2a). The physiognomy of herb layer is determined by dominant presence of the Urtica dioica. Other species such as Glechoma hederacea and Petasites hybridus reach higher cover values only occasionally (Figure 3). Higher frequency is charac- Figure 3: Mesophilous streamside forests with dominant species Petasites hybridus (Photo: J. Koštal, 19. 07. 2011, Velke Pole - Tomov štal). Slika 3: Mezofilni gozdovi ob vodotokih s prevladujočo vrsto Petasites hybridus (Foto: J. Koštal, 19. 07. 2011, Velke Pole -Tomov štal). teristic for mosses Brachythecium rutabulum and Eurhynchium hians. The species richness of this community, ranging from 29 to 52 species with a mean of 40 taxa per relevé, is significantly lower than the species number in both variants of Carici acutiformis-Alnetum glutinosae (Table 1). The conspicuous feature in vertical structure of the community Carici acutiformis-Alnetum glu-tinosae is the homogenous tree layer composed of Alnus glutinosa and the species-rich shrub layer (Table 2). The floristic spectrum depends on specific environmental conditions which are suitable especially for numerous hygrophilous and marshland species, including Angelica sylvestris, Caltha palustris, Carex remota, Lycopus europaeus, Lysimachia vulgaris, Myosotis scorpioides agg., Poa trivialis and Solanum dulcamara. The tall-sedge Carex acutiformis together with Impatiens noli-tan-gere and Caltha palustris most often alternate as dominant species in the forest understorey (Table 2, Figure 4). Unlike the constant water-toler- ant species, mesophilous plants are presented only with lower frequency. In addition to the regular occurrence of species Plagiomnium undulatum, moss layer is locally formed by generalists of wet habitats (e.g. Atrichum undulatum, Plagiomnium affine s. lat.). In accord with the species composition pattern, we distinguished two alder forests at the level of variants. The Carici acutiformis-Alne-tum glutinosae variant with Ligustrum vulgare consists of species-rich shrub layer including mainly species such as Acer campestre, Fraxinus excelsior, Ligustrum vulgare, Prunus spinosa and Swida sanguinea, whereas the stands of variant with Galium palustre encompass only weakly developed shrub layer. On the other hand, the herb layer of latter one is also enriched with ferns (Athyrium filixfemina, Dryopteris spec. div.) and moisture-demanding species (e.g. Cardamine amara, Juncus effusus, Persicaria hydropiper). These variants are equally species-rich with means of 58 and 63 taxa per relevé, respectively (Table 1). Figure 4: Physiognomy of forest understorey in the community of Carici acutiformis-Alnetum glutinosae created by Carex acutiformis (Photo: J. Koštal, 22. 07. 2011, Tesare). Slika 4: Fizionomija zeliščne plasti asociacije Carici acutiformis-Alnetum glutinosae, kjer prevladuje Carex acutiformis (Foto: J. Koštal, 22. 07. 2011, Tesare). Alder carr forests with dominant Carex elon-gata are documented only with two phytoso-ciological relevés given below. They exclusively settle waterlogged soils. Species composition of these stands reflects typical microrelief heterogeneity (Figure 5), i.e. species adapted to waterlogged hollows and species showing affinity to hummocks are accompanied by plants of wet meadows along with liana elements. This species combination supports their assignment into the association Carici elongatae-Alnetum glutinosae. Relevé I, Jedl'ové Kostol'any, Zitava river at the mouth of the Osná valley (Tribec Mts.), altitude 320 m, area 400 m2, cover E3 85%, E2 2%, Ej 95%, E0 5%, 18° 29' 54.29" E, 48° 27' 05.86" N, date 09. 07. 2011, author J. Kostal. E3: Alnusglutinosa 5. E2: Fraxinus excelsior +, Viburnum opulus +, Acer campestre r, A. pseudoplatanus r, Crataegus monogyna r, Ligustrum vulgare r, Sambucus nigra r. Ej! Calthapalustris 3, Carex elongata 3, Thelypterispalustris 2b, Carex appropinquata 2a, Crepis paludosa 2a, Impatiens parviflora 2a, Carex remota 1, Dryopteris carthusiana 1, Equisetum arvense 1, Impatiens noli-tangere 1, Lycopus europaeus 1, Lysimachia vulgaris 1, Solanum dulcamara 1, Alnus glutinosa +, Athyrium filix-femina +, Bidens frondosus +, Brachypodium syl-vaticum +, Cardamine amara +, Circaea lutetiana +, Dryopteris dilatata +, Epilobium tetragonum subsp. tetragonum +, Festuca gigantea +, Filipendula ulma-ria +, Galium palustre +, Humulus lupulus +, Myosotis scorpioides agg. +, Poa trivialis +, Ranunculus repens +, Rubus idaeus +, Scirpus sylvaticus +, Viburnum opulus +, Acer pseudoplatanus r, Angelica sylvestris r, Deschampsia cespitosa r, Dryopteris filix-mas r, Eu-onymus europaeus r , Melica uniflora r, Poa nemora-lis r, Rumex sanguineus r, Stellaria nemorum r, Swida sanguinea r, Veronica beccabunga r. E0: Plagiothecium denticulatum 1, Atrichum undulatum +, Brachythecium rutabulum +, Herzogiella seligeri r, Lophocolea heterophylla r. Figure 5: Microrelief heterogeneity in alder carr forests of Carici elongatae-Alnetum glutinosae with presence of hummocks and stagnant water on soil surface in hollows (Photo: J. Koštal, 22. 07. 2011, Nemecky). Slika 5: Mikroreliefna heterogenost v črnojelševem grezu Carici elongatae-Alnetum glutinosae z grbinami dvignjenimi nad stoječo vodo (Foto: J. Koštal, 22. 07. 2011, Nemecky). Relevé II, Nemecky, water reservoir (Povazsky Inovec Mts.), altitude 282 m, area 400 m2, cover E3 90%, E2 5%, E1 50%, E0 2%, 18° 06' 36.68" E, 48° 41' 09.43" N, date 22. 07. 2011, author J. Kostal. E3: Alnusglutinosa 5. E2: Alnus glutinosa +, Padus avium r, Salix fragilis r. E1: Carex elongata 3, Carex remota 2a, Iris pseudacorus 1, Lycopus europaeus 1, Lysimachia vulgaris 1, Scirpus syl-vaticus 1, Solanum dulcamara 1, Callitriche sp. +, Cal-thapalustris +, Carex acutiformis +, C. canescens +, De-schampsia cespitosa +, Filipendula ulmaria +, Galium palustre +, Glyceria fluitans +, Impatiens noli-tangere +, I. parviflora +, Juncus effusus +, Lemna minor +, Lysimachia nummularia +, Myosotis scorpioides agg. +, Phalaroides arundinacea +, Ranunculus repens +, Athy-rium filix-femina r, Calystegia sepium r, Cardamine amara r, Circaea lutetiana r, Equisetum arvense r, Festuca gigantea r, Fraxinus excelsior r, Galeopsis speciosa r, Glechoma hederacea r, Poa palustris r, Poa trivialis r, Quercus robur r, Rubus idaeus r, Salix fragilis r, Stachys sylvatica r, Urtica dioica r. E0: Atrichum undulatum +, Brachythecium rutabulum +, Hypnum cupressiforme +, Plagiomnium undulatum +. Syntaxonomical scheme of the documented alder forests is: Class: Alnetea glutinosae Br.-Bl. et R. Tx. ex Westhoff et al. 1946 Order: Alnetalia glutinosae R. Tx. 1937 Alliance: Alnion glutinosae Malcuit 1929 Association: Carici acutiformis-Alnetum glutinosae Scamoni 1935 variant with Ligustrum vulgare variant with Galium palustre Association: Carici elongatae-Alnetum glutinosae Schwickerath 1933 Class: Querco-Fagetea Br.-Bl. et Vlieger in Vlieger 1937 Order: Fagetalia Pawlowski in Pawlowski et al. 1928 Alliance: Alnion incanae Pawlowski in Pawlowski et al. 1928 Association: Stellario nemorum-Alnetum glutinosae Lohmeyer 1957 Environmental conditions Statistically significant (P < 0.05) Pearson correlations with the first PCA ordination axis stressed the role of EIV for moisture (r = -0.86), light (r = -0.84), portion of open water in soil surface (r = 0.50) and portion of soil surface without any vegetation (r = -0.49). Based on this evidence, the left side of the ordination diagram with higher moisture- and light-requirements was occupied by relevés of Carici acutiformis-Alnetum glutinosae (cluster B), whereas stands of Stellario nemorum-Alnetum glutinosae (cluster A) having inverse the above-mentioned values were concentrated in the opposite end (Figure 2b, c). The second PCA axis can be interpreted as an acidity gradient (Figure 2c) following the strongest positive correlation with the soil pH (r = 0.69, P <0.01). Along this axis, there were scattered especially relevés of particular variants of the Carici acutiformis-Alnetum glutinosae. Multiple comparisons indicated only slight differences in environmental factors (Table 1), but they mostly coincided with the main gradients governing species composition in alder forest vegetation (Figure 2b, c). Alder carr forests confirmed expected relation on more hy-grophilous sites with higher portion of open soil surface than riparian streamside vegetation of Stellario nemorum-Alnetum glutinosae. 4. DISCUSSION The present study emphasized floristical and environmental differentiation of alder plant communities. The identified forest vegetation types belong to either alder carr forests of Carici acuti-formis-Alnetum glutinosae and Carici elongatae-Alne-tum glutinosae (Alnion glutinosae) or mesophilous floodplain forests of Stellario nemorum-Alnetum glutinosae (Alnion incanae). They differ in the species composition and in several ecological characteristics, e.g. water regime, soil properties and microrelief heterogeneity. Alder carr forests grow on permanently waterlogged soils with stagnant or very slow flowing water on soil surface (Ellenberg 2009). These conditions are favourable for occurrence of numerous moisture-demanding species including the marshland plants, spring specialists and common species of wet habitats (Table 2). The higher light-requirements of the Alnion glutinosae vegetation (Figure 2b) can be similarly addressed to the heavy and swampy soils, as these edaphic properties naturally control mosaic structure of the tree layer permitting more light penetration through the canopy (Douda 2008, Slezák et al. 2011). Analogous sites also showed higher portion of soil surface without any vegeta- tion (Table 1) what can be attributed to partial decline of water table in sampling period. In spite of such water fluctuation, soils remain still sufficiently moist. Although the significant impact of soil reaction on variability of alder forests has already been proven by Douda (2008), acidity gradient was important environmental driver only for delimitation of two variants within Carici acutiformis-Alnetum glutinosae in study region. The unsupervised method of numerical classification (cf. Cerna & Chytry 2005) did not split meso- to eutrophic alder carr vegetation growing on periodically waterlogged soils (Carici elon-gatae-Alnetum glutinosae) from eutrophic black alder carr forests confined to colline areas (Carici acutiformis-Alnetum glutinosae). This mergence most likely results from small number of relevés with dominance of Carex elongata in the analysed data set. Both of these vegetation types belong to commonly recorded alder carr forests in Slovakia (e.g. Balâzs 1996, Somsâk 2000, Kollâr et al. 2005, Slezâk et al. 2011). Plant communities with similar floristic structure and habitat preference are known from several Central European countries (Pott 1992, Neuhauslovâ 2000, Willner & Grabherr 2007, Borhidi et al. 2012). Eutrophic black alder stands of low-lying damp and marshland localities have also been reported from the Pan-nonian Basin (Szmorad 2011, Borhidi et al. 2012) and adjacent regions in the Western Carpathians (Kliment & Watzka 2000) as Angelico sylvestris-Alnetum glutinosae. To achieve reliable resolution of its syntaxonomical position within the alliance Alnion glutinosae in Central European context and to clarify its relationship to Carici acutiformis-Alnetum glutinosae, phytosociological synthesis of large-scale data set using numerical approach is necessary. Floristic spectrum and higher cover values of species such as Caltha palustris, Carex appropin-quata, C. elongata and Thelypterispalustris in relevé no. I is interesting from syntaxonomical point of view. There is well-established oligotrophic alder carr vegetation with marsh fern (Thelypteris palus-tris) in Europe (Prieditis 1993, Willner & Grabherr 2007). Although analogous forests have been recorded under different names, this community was firstly described as an association Thelyp-terido palustris-Alnetum glutinosae by Klika (1940) from the Czech Republic. Species-poor herb layer, acidophilous and mire plants along with Sphagnum species, which are completely lacking in our relevé, represent diagnostic features of this association (cf. Klika l. c., Douda 2008). Their absence in combination with constant presence of typical marshland species indicates its relation to the Carici elongatae-Alnetum glutinosae. Habitat qualities of the riparian streamside forests are strongly influenced by periodic floods during the spring and by markedly declining water table in vegetation period (Neuhauslovâ 2000, Ellenberg 2009). Seasonal fluctuation of groundwater table, affecting large variability in soil moisture, results in mixed appearance of forest mesophilous and slightly hygrophilous species (Table 2) typical for the association Stellario nemorum-Alnetum glutinosae (Neuhauslovâ & Kol-bek 1993). It belongs to the most frequently documented Alnion incanae community in the colline and submontane zones of Central Europe (Pott 1992, Willner & Grabherr 2007, Douda 2008, Ma-tuszkiewicz 2012), including also Slovakia (e.g. Miadok 1978, Kontris 1981, Hrivnâk et al. 2009, Slezâk et al. 2011). In Hungarian phytosociologi-cal literature, analogous forest vegetation type has usually been mentioned as Aegopodio-Alne-tum glutinosae V. Kârpâti, I. Kârpâti et Jurko ex Somsâk 1961 (e.g. Borhidi et al. 2012). Although this association's name was several times used for the riparian alder vegetation on mesic to humid sites along small brooks in Slovakia as well (Somsâk 1961, Kârpâti et al. 1963, Balâzs 1996), preliminary analysis of our own data with those published in the above-mentioned botanical studies indicates certain overlapping with Stellar-io nemorum-Alnetum glutinosae in terms of species composition, physiognomy and ecology. However, to create consisting and unambiguous classification of Slovak alder woodlands, the existing relevé material from various authors should be used in accordance with the nomenclatural rules in comparative analysis (cf. Slezâk et al. 2013). 5. ACKNOWLEDGEMENTS The authors thank H. Vojtekovâ and R. Pozgaj for assistance during the field sampling, V. Gru-lich for determination and/or revision of herbarium specimens of Carex genera, D. Kudelovâ for language improvement and anonymous reviewers for their comments to previous version of the manuscript. 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Microcomputer Power, Ithaca, NY, 500 pp. Tichy, L. 2002: JUICE, software for vegetation classification. Journal of vegetation Science 13:451-453. Westhoff, V. & van der Maarel, E. 1973: The Braun-Blanquet approach. In: Whittaker, R. H. (ed.): Ordination and classification of communities. Junk, The Hague, pp. 617-727. Willner, W. & Grabherr, G. 2007: Die Wälder und Gebüsche Österreichs. Ein Bestimmungswerk mit Tabellen. Spektrum Akademischer Verlag, Heidelberg, 302 pp. Received 21. 12. 2012 Revision received 14. 5. 2013 Accepted 29. 5. 2013 Table 1: Descriptive statistics and multiple comparisons of the environmental variables. Only statistically significant differences (P <0.05) in post-hoc Tukey HSD test indicating by different letters are presented. Tabela 1: Opisna statistika in primerjave rastiščnih spremenljivk. Z različnimi črkami so prikazane samo statistično značilne razlike (P <0.05) v post-hoc Tukey HSD testu. Environmental variable Mean SD Max Min ClusterA ClusterB1 ClusterB2 Number of species per relevé 55 12 74 29 40b 58b 63a Ellenberg indicator values Light 5.59 0.30 6.15 4.88 5.33 5.69 5.71 Temperature 5.27 0.45 5.53 3.58 5.35 5.44 5.02 Continentality 3.79 0.90 7.26 3.35 3.63 3.57 4.15 Moisture 6.61 0.38 7.14 5.85 6.22b 6.70a 6.85a Soil Reaction 6.61 0.22 6.93 6.23 6.64 6.72 6.48 Nutrients 6.26 0.25 6.73 5.82 6.40 6.19 6.23 Measured environmental values Altitude (m) 302 75 500 215 348 286 281 Area (m2) 29345.06 56222.25 236128.00 4250.00 13061.60 47830.83 24428.83 Valley width (m) 102.35 53.91 230.00 20.00 94.00 126.67 85.00 Minimum width or length (m) 79.94 57.73 260.00 30.00 60.80 105.00 70.83 Open soils (%) 10.71 9.03 35.00 0.00 5.40a 7.50ab 18.30b Open water (%) 10.12 5.79 19.00 1.00 12.20 11.67 6.83 Maximum content of Ca (%) 2.85 3.48 10.00 0.50 3.30 4.58 0.75 N content (mg/l) 21.30 10.30 49.29 11.13 29.26 18.91 17.04 pH H2O 6.48 0.85 7.68 4.97 6.03 7.13 6.20 Table 2: Phytosociological table of alder-dominated plant communities with dendrogram of the TWINSPAN analysis. Tabela 2: Fitocenološka tabela rastlinskih združb s prevladujočo črno jelšo z dendrogramom TWINSPAN analize. Taxon/Releve's number 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 Cluster Diagnostic species of cluster A; Alnion incanae, Stellario nemorum-Alnetum glutinosae Aegopodium podagraria b+1a a + + 1 1 1 + a Acer campestre (E2) +a1a 1 + 1 1 + r + + Galium aparine 1 a + + a + + + + 1 1 + Pulmonaria officinalis agg. r 1 r . + r r + Corylus avellana (E2) . r r + + + 1 Diagnostic species of cluster B; Alnion glutinosae, Carici acutiformis-Alnetum glutinosae Caltha palustris +.+. r a + b 1 + + 1 a b b 4 + Lysimachia vulgaris . . . + r + r 1 a + + r + 1 + 1 + Poa trivialis r + + r r r 1 b a + b Solanum dulcamara r 1 + 1 + + + 1 + a a r Lycopus europaeus 1 + 1 1 a 1 1 1 + 1 + Angelica sylvestris r . . . r + r r 1 r + + r + r Festuca gigantea r r . . + r r r + 1 + + + + + Carex acutiformis + + 4 3 a 3 1 r + + a Scirpus sylvaticus + + + a + 1 a + 1 b Rumex sanguineus r . . . + + + r + + + r r 34 Taxon/Relevé's number 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 + + a + ++ ++ + . ++ Ranunculus repens ....+ .a + + +. a.al Athyrium filix-femina ..1.. ...+ + + .1r + Rubus caesius ....1 a 1 1 a r . ...1 Plagiomnium undulatum (E0)...........1 + 1.. +..1 Valeriana dioica . . . . . . r 1 . + . r r . r Viburnum opulus (E2) ..........rar..r .rr + Eupatorium cannabinum . . . . . r 1 a . + . r 1 . . Plagiomnium affine s. lat. (E0)........... + 1... ++.. Diagnostic species of cluster B1; Carici acutiformis-Alnetum glutinosae, variant with Ligustrum vulgare Fraxinus excelsior (E2) .+ r. . + r 1 r r Humulus lupulus +. + 1 r . r + . + . . .+ Ligustrum vulgare (E2) r 1 r + r r + . r . 4 1 1 1 + r + r r 1 + + a 1 3 a 1 + + r 1 + a + r 4 3 1 + a r r r + 4 a + 1 1 + + r 1 + r 1 + + 1 a r + + r r r r r r + r 1 + 1 r + r + r + a + + 1 a a 1 1 a 3 + 1 a + + + r + + 1 1 + 1 r + 1 + 1 + + r r + + + 1 1 + + 1 + + + + + 1 r a + + + + 1 + 3 + r 5 r + + Diagnostic species of cluster B2; Carici acutiformis-Alnetum glutinosae, variant with Galium palustre Lapsana communis . + Myosotis scorpioides agg. . + Juncus effusus Persicaria hydropiper . . + . . . r . 1 . 1 Carex remota r . . . . ... a + a Galium palustre . . . . . . r . + Veronica beccabunga . . . . . . r . + Cardamine amara Cerastium lucorum Galeopsis speciosa Tree layer (E^ Alnus glutinosa 5 5 5 5 5 5 5 5 5 5 Salix fragilis . + . + r r . . . + Shrub layer (E2) Sambucus nigra Crataegus monogyna Euonymus europaeus Prunus spinosa Swida sanguinea Acer pseudoplatanus Carpinus betulus Malus sylvestris Ribes rubrum Herb layer (EJ Urtica dioica Glechoma hederacea Impatiens noli-tangere Geum urbanum Circaea lutetiana Brachypodium sylvaticum Lysimachia nummularia Geranium robertianum Equisetum arvense Stachys sylvatica Ajuga reptans Cirsium oleraceum + a . . a Filipendula ulmaria +..++ +.+ + + a Impatiens parviflora a + . 1 a . 1 . + . 1 Alliaria petiolata 1 r a . . 3 . . + r a 5 + b r r r r + r + r 1 r 3 a b + 1 a + r r + 1 + 3 b a + 1 + + + + r + r r r 1 b 1 1 1 + r r + r + 1 3 1 b + 1 1 b 1 a + a 1 + + r 1 + + a + r r r a b 1 r + r + r + r 1 r 1 1 + 5 5 5 5 5 5 r + + r + + r r + 1 r r r + r + r 1 + r 1 1 1 + + 1 + a + 1 1 + b 3 1 1 3 a + r r + + r 1 1 r + 1 + + + + + + + b 1 1 + + + r + + r 1 + + r + + + + + + + r r + + r + a a 1 r + r Taxon/Relevé's number 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 Lamium maculatum Rubus idaeus Deschampsia cespitosa Dryopteris carthusiana Chrysosplenium alternifolium Acer campestre Petasites hybridus Viburnum opulus Carex sylvatica Oxalis acetosella Poa palustris Myosoton aquaticum Euonymus europaeus Lythrum salicaria Scrophularia umbrosa Scutellaria galericulata Epilobium roseum Dryopteris dilatata Moss layer (E0) Brachythecium rutabulum Eurhynchium hians Brachythecium rivulare Amblystegium serpens Atrichum undulatum 1 + a + + + + + + + + + . . r . + a + r 1 . + rrr .r 1 + .. r 1 r r +. +r r+ .r . . + r r . . + . r . . . + . + . . + 1 . . 1 + + . + + . . . . . + . . + . . ++ 1+ r+ .r .+ 1r .r +. r + + r + + + .r +r + . + + + . . r . + + + +a + rr . + b r 1+ ++ a + 1 + 1 r r 3 b + + + + r r r r + + + r + 1 + + + r + r 1 Species occurring in less than four (< 3) phytosociological relevés: E3: Acer campestre (10, r), Fraxinus excelsior (2, 2a), Populus tremula (5, +), Salix caprea (2, +), Ulmus minor (10, +). E2: Acerplatanoides (9, r), Alnus glutinosa (8, +; 13, +; 14, r), Cerasus avium (4, r; 8, r; 15, +), Crataegus laevigata (12, r), Frangula alnus (7, +; 17, r), Lonicera xylosteum (8, +), Padus avium (4, r; 7, 2a; 9, r), Rhamnus catharticus (6, r; 13, r), Rosa canina agg. (9, r; 13, r; 15, r), Salix cinerea (8, +; 12, r), S. fragilis (8, r; 10, r), Tilia cordata (9, r; 12, r), Ulmus glabra (8, r), U. minor (10, r; 11, r). Et: Acerpseudoplatanus (7, r), Aethusa cynapium (2, 1; 13, +), Alisma plantago-aquatica (14, r; 17, r), Asarum europae-um (2, 1), Astrantia major (16, r), Bidens frondosa (11, r; 14, 2b; 17, +), B. tripartita (9, +; 12, r; 14, r), Calamagrostis epigejos (15, r), Callitriche sp. (17, +), Calystegia sepium (7, +; 10, r), Cardamine flexuosa (13, r; 16, r), Cardamine impatiens (5, r; 13, 2a; 14, +), Cardamine pratensis agg. (9, +; 11, +), Carduus crispus (2, 1; 13, r), Carex acuta (3, +), C. flava agg. (15, r), C. nigra (4, +), C. paniculata (8, 2a; 13, 2b), C. pendula (12, 1), Chaerophyllum aromaticum (2, 2b; 12, +), Chelidonium majus (2, r), Chenopodium album agg. (17, r), C. polyspermum (14, r), Corylus avellana (4, r), Crataegus monogyna (6, +), Crepis paludosa (8, 1; 16, +), Cucubalus baccifer (7, r; 10, r), Dactylis polygama (4, r; 8, r), Dryopteris filix-mas (4, r; 7, r), Epilobium ciliatum (11, r), E. parviflorum (14, r), E. tetragonum (14, +), Equi-setum fluviatile (13, r), E. palustre (8, r), Fagus sylvatica (2, r), Fallopia dumetorum (11, r; 13, +), Fragaria vesca (8, r), Fraxinus excelsior (6, r; 11, +), Galeobdolon luteum (3, 1), Galeobdolon montanum (2, +; 15, r; 16, 3), Galeopsis bifida (3, r; 11, r), Galinsoga urticifolia (12, r), Galium odoratum (7, +; 9, r), G. rivale (13, +), Geranium palustre (15, r), G. phaeum (2, 2a), Glyceria fluitans (9, 1; 15, r; 17, r), Hol-cus lanatus (8, r; 13, r; 17, r), Iris pseudacorus (1, r; 11, +; 12, +), Lysimachia nemorum (2, +; 3, +), Malus sylvestris (12, r), Melica uniflora (11, r), Mentha aquatica (7, 1), M. arvensis (14, +), M. longifolia (8, r), Mercurialis perennis (9, r), Milium effusum (4, r; 16, +), Moehringia trinervia (13, r; 17, r), Mycelis muralis (9, r; 15, r), Persicaria dubia (13, r; 14, +), P maculosa (10, r; 11, r), Phalaroides arun-dinacea (13, r), Phragmites australis (7, +), Plantago major (14, r), Poa nemoralis (4, r; 11, r), Prunella vulgaris (15, r), Quercus cerris (4, r; 9, r), Q. dalechampii (4, r), Q. robur (4, r; 6, r), Ranunculus auricomus agg. (4, r; 17, r), R. lanugi-nosus (12, 1; 13, r), R. sceleratus (14, +), Ribes rubrum (6, r), Roegneria canina (2, r; 13, +), Rorippa palustris (12, r; 14, +), Rosa canina agg. (6, r; 14, r), Rumex obtusifolius (12, r), Sambucus nigra (14, +), Senecio germanicus (4, +), S. ovatus (2, r), Sonchus asper (14, r), Sorbus aucuparia (4, r), Sparganium erectum (14, r), Stellaria alsine (15, r; 17, +), S. holostea (9, r), S. media (9, r; 11, +), Symphytum officinale (10, r), Swida sanguinea (7, +; 14, r), Torilis japonica (2, r; 13, r), Ulmus minor (11, r), Valeriana officinalis (8, +), Veronica chamaedrys (2, r; 17, r), V. montana (13, r; 17, r), Vicia sepium (16, r), Viola odorata (15, r), V reichenbachiana (9, r). E0: Amblystegium varium (13, r), Brachythecium salebrosum (4, +; 7, +), Bryum subelegans (4, +), Calliergon cordifoli-um (17, +), Calliergonella cuspidata (8, r), Climacium den-droides (8, r), Cratoneuron filicinum (12, +), Herzogiella seligeri (4, r), Homalia trichomanoides (8, +), Lophocolea heterophylla (3, +; 4, +), Plagiomnium cuspidatum (10, +), Plagiothecium denticulatum (7, +; 16, +), Rhizomnium punctatum (9, +). Localities of phytosociological relevés Relevé number, locality-village (full headers with higher accuracy location of individual relevés are presented in the Slovak Phytosociological Database; http://ibot.sav.sk/cdf/ index.html), altitude (m), aspect (in letters), slope (degrees), relevé area, cover of tree layer (E3), cover of shrub layer (E2), cover of herb layer (E1), cover of mosses and lichens (E0), longitude, latitude, date (day/month/year), relevé author. 1. Bojna, Bojnianka walley, 2 km NW from Ranč pod Babicou (Považsky Inovec Mts.), 240 m, -, 0°, 400 m2, E3 90%, E2 1%, E1 98%, E0 5%, 18° 015' 09.18" E, 48° 36' 56.24" N, 24. 07. 2011, J. Koštal. 2. Velké Pole, Tomov štal, 1.5 km NE from village Velké Pole (Vtačnik Mts.), 500 m, -, 0°, 400 m2, E3 90%, E2 20%, E1 95%, E0 2%, 18° 34' 35.20" E, 48° 32' 52.28" N, 19. 07. 2011, J. Koštal. 3. Kostolany pod Tribečom, Babova walley, 800 m NE from the Jedliny (Tribeč Mts.), 300 m, -, 0°, 400 m2, E3 95%, E2 8%, E1 90%, E0 5%, 18° 14' 11.19" E, 48° 26' 25.31" N, 27. 07. 2011, J. Koštal. 4. Jedlové Kostolany, Žitava walley, on the confluence with Žitavica (Tribeč Mts.), 400 m, -, 0°, 400 m2, E3 90%, E2 2%, E1 100%, E0 5%, 18° 31' 56.16" E, 48° 28' 23.79" N, 19. 07. 2011, J. Koštal, H.Vojtekova & R. Požgaj. 5. Horné Hamre, Brod, Pilansky stream, 1.5 km SWW from Brod (Vtačnik Mts.), 300 m, -, 0°, 400 m2, E3 95%, E2 5%, E1 90%, E0 2%, 18° 38' 41.30" E, 48° 30' 03.87" N, 19. 07. 2011, J. Koštal, H.Vojtekova & R. Požgaj. 6. Kolačno, water reservoir (Tribeč Mts.), 250 m, -, 0°, 400 m2, E3 90%, E215%, E1 95%, E0 2%, 18° 24' 37.88" E, 48° 35' 11.55" N, 22. 07. 2011, J. Koštal, H. Vojtekova& R. Požgaj. 7. Brodzany, Brodziansky stream, 2 km SE from Brodzany (Tribeč Mts.), 240 m, -, 0°, 400 m2, E3 90%, E2 30%, E1 90%, E0 10%, 18° 21' 27.65" E, 48° 35'40.20" N, 07. 07. 2011, J. Koštal. 8. Pila, Pilansky stream, 1 km NW from Pila (Vtačnik Mts.), 400 m, -, 0°, 450 m2, E3 80%, E2 15%, E1 95%, E0 5%, 18° 35' 01.67" E, 48° 31' 32.86" N, 09. 07. 2011, J. Koštal. 9. Kostolany pod Tribečom, by Drevenica stream, 1.2 km N from Jedliny (Tribeč Mts.), 330 m, -, 0°, 400 m2, E3 90%, E2 20%, E1 85%, E0 5%, 18° 13' 23.46" E, 48° 26' 35.58" N, 27. 07. 2011, J. Koštal. 10. Tesare, Zlavsky stream, 0.5 km NW from water reservoir Tesare (Považsky Inovec Mts.), 215 m, -, 0°, 400 m2, E3 80%, E2 5%, E1 95%, E0 7%, 18° 04' 28.93" E, 48° 36' 52.95" N, 22. 07. 2011, J. Koštal, H. Vojtekova & R. Požgaj. 11. Kostolany pod Tribečom, Jedliny, by Drevenica stream (Tribeč Mts.), 280 m, -, 0°, 450 m2, E3 95%, E2 5%, E1 95%, E05%, 18° 13' 19.55" E, 48° 26' 11.37" N, 27. 07. 2011, J. K0oštal. 12. Bojna, Bojnianka walley, 2.5 km NW from Ranč pod Babicou (Považsky Inovec Mts.), 245 m, -, 0°, 400 m2, E3 90%, E25%, E1 65%, E0 2%, 18° 01' 45.45" E, 48° 37' 14.52" N, 24. 07. 2011, J. Koštal. 13. Hostie, Hlboka walley, Hostiansky stream, 1 km N from Skycovsky mlyn (Tribeč Mts.), 300 m, -, 0°, 400 m2, E3 85%, E2 1%, E1 85%, E0 2%, 18° 28' 27.91" E, 48° 29' 16.51" N,08. 07. 2011, J. Koštal. 14. Kolačno, water reservoir (Tribeč Mts.), 250 m, -, 0°, 400 m2, E3 90%, E2 1%, E1 70%, E0 5%, 18° 24' 39.53" E, 48° 35' 13.05" N, 22. 07. 2011, J. Koštal. 15. Skycov, Uhrovska walley, Vyčoma stream (Tribeč Mts.), 310 m, -, 0°, 400 m2, E3 90%, E2 1%, E1 75%, E0 7%, 18° 24' 20.85" E, 48° 31' 47.61" N, 12. 07. 2011, J. Koštal. 16. Velke Uherce, Drahožicka walley, Drahožicky stream, 4 km S from water reservoir Velke Uherce (Tribeč Mts.), 350 m, -, 0°, 450 m2, E3 85%, E2 5%, E1 85%, E0 5%, 18° 27' 20.96" E, 48° 33' 10.17" N, 07. 07. 2011, J. Koštal. 17. Solčany, Lazkovy stream (Tribeč Mts.), 230 m, -, 0°, 400 m2, E3 80%, E2 1%, E1 90%, E0 5%, 18° 14' 36.24" E, 48° 31' 03.51" N, 21. 06. 2011, J. Koštal.