HACQUETIA 6/1 • 2007, 61-76 diversity of dry grasslands in the považsky inovec mts (slovakia) - a numerical analysis Daniela MICHÄLKOVÄ* Abstract The paper reveals a numerical and ecological analysis of 128 published relevés of xerothermic vegetation of the Považsky Inovec Mts. Four associations have been recognised: Festuco valesiacae-Stipetum capillatae Sillinger 1930, Festuco pallentis-Caricetum humilis Sillinger 1930 corr. Gutermann et Mucina 1993, Poo badensis-Festucetumpallentis Klika 1931 corr. Zolyomi 1966 nom. invers. propos. and Minuartio setaceae-Seslerietum coeruleae Klika 1931 nom. mut. propos. Through the numerical methods, the study reconsiders the conception of the dry grassland associations, which are traditionally used in the phytocoenological literature. It also defines the indication taxa group of the associations and characterises their environmental conditions. Major environmental gradients influencing the vegetation were interpreted using average Ellenberg indication values. Finally, the classification within the higher vegetation units, the nomenclature of the associations and the environmental threats are discussed. Key words: classification, cluster analysis, dry grasslands, Festucetalia valesiacae, ordination, Považsky Inovec Mts, Slovakia, syntaxonomy Izvleček V članku je prikazana numerična in ekološka analiza 128 objavljenih popisov kserotermne vegetacije hribovja Považsky Inovec. Ugotovljene so štiri asociacije: Festuco valesiacae-Stipetum capillatae Sillinger 1930, Festuco pal-lentis-Caricetum humilis Sillinger 1930 corr. Gutermann et Mucina 1993, Poo badensis-Festucetum pallentis Klika 1931 corr. Zolyomi 1966 nom. invers. propos. in Minuartio setaceae-Seslerietum coeruleae Klika 1931 nom. mut. propos. Z numeričnimi metodami je avtorica ponovno preverila sinsistem asociacij suhih travnikov, ki so omenjene v fitocenološki literaturi. Opredelila je skupine značilnih vrst asociacij in značilnosti rastiščnih razmer. S povprečnimi Ellenbergovimi indikacijskimi vrednostmi so prikazani glavni ekološki gradienti, ki vplivajo na floristično sestavo vegetacije suhih travišč. Članek obravnava uvrstitev asociacij v višje sintaksonomske enote, nomenklaturo asociacij in njihovo ogroženost. Ključne besede: klasifikacija, klastrska analiza, suha travišča, Festucetalia valesiacae, ordination, Považsky Inovec, Slovaška, sintaksonomija 1. INTRODUCTION The Považsky Inovec Mts are located in the western part of Slovakia along the river Vâh in the prae-Car-pathian limestone zone (Fig. 1). The central part of the mountains, the Tematmske kopce Hills, is built up by dolomites. In this area, the numerous dry grasslands are located. However, the potential vegetation of this area are the forests, mostly the oak-hornbeam forests (Carici pilosae-Carpinetum) and locally the thermophilous oak forests ( Quercion pubescentis-petraeae; Michalko et al. 1987). The major part of the forest cover had been destroyed during the Turkish war approximately in the 16th and 17th century. These areas later became grasslands. The dry grassland vegetation is geographically influenced from two directions. From the south, the study area is in near contact with the eu-Pan-nonian lowland region, which influences the vegetation by numerous Pontic-Pannonian and sub-Mediterranean thermophilous species. From the north, the Carpathian mountain species descend * Slovak Academy of Sciences, Institute of Botany, Dubravskâ cesta 14, SK-845 23 Bratislava, Slovakia. E-mail: daniela. michalkova@savba.sk 61 HQ_6-1_2007.indd 61 12.6.2007 8:44:16 Hacquetia 6/1 • 2007, 61-76 and build up some dealpine and praealpine elements (Devan et al. 2006). This vegetation is interesting from the floris-tical, historical as well as the successional point of view. It was studied in detail only in two older works (Sillinger 1930, Maglocky 1979). Although some works have recently been done (Michalkova, Škodova & Mertanova 2006; Janišova 2005, 2006a, b, 2007a, b), no numerical analysis has been performed so far. The presented paper brings a numerical classification of all published relevés. The use of numerical methods endeavours to support or eventually correct the current phytosociological scheme of the dry grasslands of the study area. The aim is also to define the indication taxa group of the associations and some characteristics of their environmental conditions. 2. MATERIAL AND METHODS The numerical analyses were performed using the published relevés of xerothermic vegetation of the Považsky Inovec Mts, created by applying the standard Central-European method (Braun-Blanquet 1964, Westhoff & van der Maarel 1973). The analysed data set comprised 128 relevés from the works Maglocky (1979), Sillinger (1930), Klika (1931) and Michalkova, Škodova & Mertanova (2006). The taxa determined only on the level of genus were excluded from the analyses. The layer E0 was excluded as a whole, because mosses and lichens were not determined in all relevés analysed. Some taxonomically problematic species, which were not distinguished by all authors of the relevés, were classified within higher or more broadly defined taxa: Anthyllis vulneraria agg. (subsp. polyphylla), Artemisia campestris (subsp. lednicensis), Colymbada scabiosa agg. (C. badensis), Dactylis glomerata agg. (D. polygama), Acosta rhenana agg. (Acosta biebersteinii), Helianthemum grandiflomm s. l. (subsp. grandiflorum, subsp. obscurum, H. nummularium), Pulsatilla slavi-ca (P. subbslavica), Primula veris (subsp. canescens), Thalictrum minus (subsp. pseudominus). Initial data analysis to remove outlier relevés (relevés, of which the species composition did not correspond to the rest of the analysed data set) was carried out using the CANOCO 4.5 package (ter Braak & Šmilauer 2002). It was necessary to exclude 4 relevés (Maglocky 1979: Table 23, relevés 1-3; Table 25, relevé 20). There were 124 relevés used in the subsequent classification. We compared the results of more types of cluster analyses carried out by using different algorithms. The paper presents the results of the cluster analysis processed by the program PC-ORD 4 (McCune & Mef-ford 1999) where Ward's method and the relative Euclidean distance as a resemblance measure were applied. These results proved to be floristically and ecologically well interpretable and were supported by our field experience as well. Diagnostically important taxa for the individual clusters were determined by calculating the constancy and fidelity measure of each species to each cluster, using the phi coefficient of association (Sokal & Rohlf 1995, Chytry et al. 2002) in the program Juice (Tichy 2002). The results of the classification were summarised in a synoptic 62 HQ_6-1_2007.indd 70 12.6.2007 8:44:29 Daniela Michälkovä: Diversity of Dry Grasslands in the Povazsky Inovec Mts (Slovakia) - A Numerical Analysis table (Table 1), in which both percentage species frequencies (constancies) and fidelities (9 coefficient multiplied by 100, p<0.001) were shown, and diagnostic species were ranked by decreasing fidelity (Chytry et al. 2002). Fisher's exact test excluded the accidental species with high fidelity measures (Tichy & Holt 2006). The threshold values of the indication taxa group were selected subjectively: fidelity of diagnostic species was higher than 40 %; constant species were those with a relative constancy value higher than 60 %; dominant species were defined as all species that had cover values higher than 30 %. The number values indicate fidelity for the diagnostic species and constancy for the constant species in the chapter "Nomenclature and species characteristics of the associations". There is also a number given to each dominant species, which stands for the percentage of relevés meeting the threshold criteria. The indication taxa group of each cluster was compared to the diagnostic species from Holub et al. (1967), Chytry et al. (2007) and Mucina & Kolbek (1993), which enabled interpretation of the associations in terms of higher phy-tosociological units (orders and alliances, Tab. 1). Figure 2: Dendrogram of the cluster analysis (Ward's method, relative Euclidean distance). Slika 2: Dendrogram klastrske analize (Wardova metoda, relativna Evklidska razdalja). A - Festuco valesiacae-Stipetum capillatae (1 - festucetosum rupicoae, 2 - botryochloetosum ischaemii, 3 - caricetosum caryo-phyllae), B - Festuco pallentis-Caricetum humilis (4 - typicum, Koeleria macrantha-variant), C - Poo badensis-Festucetum pal-lentis (5 - jovibarbetosum hitrae), D - Minuartio setaceae-Seslerietum caeruleae (6 - festucetosum pallentis). Sources of relevés (Viri popisov): Cluster A: 1-2 - d: rel. 1, 2; 3-24 - a: Table 27, rel. 4, 12, 16, 5, 7, 6, 20, 9, 8, 28, 29, 32, 34, 33, 17, 18, 19, 21, 22, 23, 24, 25; 25 - d: rel. 3; 26-34 - a: Table 27, rel. 26, 27, 30, 31, 10, 11, 13, 14, 15; 35-36 - d: rel. 4, 5; 37 - Chytry 1995, ined.; 38-40 - a: Table. 27, rel. 1, 2, 3; 41-43 - b: p. 36, rel. 2, 1, 3; 44-46 - d: rel. 6-8; Cluster B: 47-48 - d: rel. 9, 10; 49-52 - b: p. 36, rel. 4, 5, 6, 7; 53 - c: p. 363, rel. 14; 54 - a: Table 26, rel. 18; 55 - c: p. 363, rel. 15; 56-78 - a: Table 26, rel. 14, 21, 23, 5, 15, 8, 19, 20, 22, 11, 12, 13, 16, 17, 1, 6, 7, 4, 24, 10, 3, 2, 9; 79-88 - d: rel. 11-20; Cluster C: 89 - c: p. 363, rel. 13; 90-108 - a: Table 22, rel. 2, 6, 7, 9, 11, 1, 15, 8, 10, 14, 16, 13, 17, 18, 19, 3, 4, 5, 12; Cluster D: 109 - c: p. 370, rel. 29; 110-124 - a: Table 25, rel. 3, 2, 6, 10, 13, 15, 12, 21, 24, 25, 22, 16, 18, 17, 23. Abreviations (Okrajšave): a - Maglocky (1979), b - Sillinger (1930), c - Klika (1931), d - Michalkova, Škodova & Mertanova (2006). 63 HQ_6-1_2007.indd 63 12.6.2007 8:44:18 Hacquetia 6/1 • 2007, 61-76 Figure 3: Detrended correspondence analysis (DCA) ordination diagram of the species occurring in the dry grassland vegetation of the Považsky Inovec Mts. Slika 3: Ordinacijski diagram korespondenčne analize z odstranjenim trendom (DCA) floristične sestave suhih travnikov na hribovju Považsky Inovec. Species list (Seznam vrst): Allium flavum, A. senescens ssp. montanum, Alyssum montanum, Anthericum ramosum, Anthyllis vulneraria, Asperula cynanchica, Bothriochloa ischaemum, Bromus erectus, Campanula moravica, C. sibirica, Carex caryophyl-lea, C. humilis, Dianthuspraecox ssp. lumnitzerii, Eryngium campestre, Festucapallens, F. rupicola, F. valesiaca, Fumanaprocum-bens, Genista pilosa, Globularia punctata, Helianthemum grandiflorum, Inula ensifolia, Jovibarba hirta ssp. glabrescens, Koeleria macrantha, Leontodon incanus, Linum tenuifolium, Medicago falcata, Melica ciliata, Pilosella bauhinii, P. officinarum, Pimpinella nigra, Poa badensis, Potentilla arenaria, Sanguisorba minor, Scabiosa ochroleuca, Sedum sexangulare, Seseli hippomarathrum, S. osseum, Sesleria albicans, Silene otites, Stipa capillata, S. joannis, S. pulcherrima, Teucrium chamaedrys, T. montanum, Thlaspi perfoliatum, Thymus pannonicus, T. praecox, Tithymalus cyparissias. The ordination techniques, using the detrended correspondent analysis (DCA) from the CANO-CO 4.5 package, defined the major gradients in the spatial arrangement of species of the analysed data set. For ecological interpretation of the ordination axes, average Ellenberg indication values (Ellenberg et al. 2001) for relevés were plotted onto a DcA ordination diagram as supplementary environmental data (Fig. 3 and 4). The nomenclature of vascular plants follows Marhold & Hindâk (1998). The nomenclature of higher syntaxa is in accordance with Chytry et al. (2007) and Mucina & Kolbek (1993). Phytosocio-logical nomenclature of associations is reconsidered according to Weber, Moravec & Theurillat (2000). For the individual associations, these abbreviations were used in Fig. 4 and 5: Fv-Sc - Fes-tuco valesiacae-Stipetum capillatae, Fp-Ch - Festuco pallentis-Caricetum humilis, Pb-Fp - Poo badensis-Fes-tucetum pallentis and Ms-Sc - Minuartio setaceae-Ses-lerietum caeruleae. 64 HQ_6-1_2007.indd 64 12.6.2007 8:44:20 Daniela Michälkovä: Diversity of Dry Grasslands in the Povazsky Inovec Mts (Slovakia) - A Numerical Analysis Associations O Fv-Sc □ Fp-Ch O Pb-Fp Ms-Sc Figure 4: Detrended correspondence analysis (DCA) ordination diagram of the relevés, based on average Ellenberg indicator values. Slika 4: DCA ordinacijski diagram popisov na osnovi povprečnih Ellenbergovih indikacijskih vrednosti. -♦-light -■—temperature -^k— continentalily -s—moisture —s—soil reaction —♦—nutrients Fv-Sc Fp-Ch Pb-Fp Ms-Sc Figure 5: Average Ellenberg indication values of the ecological factors of vascular plants occurring in individual clusters (associations) produced as outputs of the cluster analysis. Legend (Ellenberg et al. 2001): Light: 7 - plants in well lit places, sometimes occurring in partial shade, 8 - light-loving plants. Temperature: 5 - indicator of fairly warm condition, 7 - warmth indicator. Continentality: 4 - sub-oceanic, 5 - intermediate. Moisture: 1 - indicator of extreme dryness, 3 - dry sites indicator. Soil reaction: 7 - indicator of weekly acid to weekly basic conditions, never found on very acid soils, 9 - basic reaction and lime indicator. Nutrients: 1 - indicator of sites extremely poor in available nitrogen, 3 - indicator of sites more or less poor in available nitrogen. Slika 5: Povprečne Ellenbergove indikacijske vrednosti posameznih klastrov (asociacij). ki so rezultat klastrske analize. Legenda (Ellenberg et al. 2001): Svetloba: 7 - rastline svetlobe, včasih uspevajo v delni senci, 8 - rastline svetlobe. Temperatura: 5 - nakazovalke zmerno toplih razmer, 7 - indikatorji relativno toplih razmer. Kontinentalnost: 4 - sub-oceanske razmere, 5 - vmesne razmere. Vlažnost: 1 - indikatorji skrajno sušnih razmer, 3 - indikatorji sušnih razmer. Kemična reakcija tal: 7 - indikatorji slabo kislih oziroma slabo bazičnih tal, tovrstnih rastlin nikoli ne najedmo na zelo kislih tleh, 9 - nakazovalke bazične reakcije ali zelo karbonatnih tal. Hrabnila: 1 - nakazovalke rastišč, najrevnejših z razpoložljivim dušikom, 3 - nakazovalke rastišč, revnih z razpoložljivim dušikom. 65 HQ_6-1_2007.indd 63 12.6.2007 8:44:18 Hacquetia 6/1 • 2007, 61-76 3. RESULTS 3.1 Cluster analysis Interpreting the results of the cluster analysis, four associations have been distinguished in the dendrogram at the third highest level of dissimilarity (Fig. 2). They are: Festuco valesiacae-Stipetum capillatae Sillinger 1930 (Fig. 6), Festuco pallentis-Caricetum hu-milis Sillinger 1930 corr. Gutermann et Mucina 1993 (Fig. 7), Poo badensis-Festucetum pallentis Klika 1931 corr. Zolyomi 1966 nom. invers. propos. and Minu-artio setaceae-Seslerietum calcariae Klika 1931 nom. mut. propos. All of these vegetation units, using different synonyms, have been traditionally diagnosed in the phytocoenological literature (cf. Maglocky 1979). Figure 6: Festuco valesiacae-Stipetum capillatae in the Nature Reserve Beckovské skalice (township of Beckov). Photo: D. Michalkova, 19. 7. 2005 Slika 6: Asociacija Festuco valesiacae-Stipetum capillatae v naravnem rezertvatu Beckovské skalice (občina Beckov). Foto: D. Michalkova, 19. 7. 2005 66 HQ_6-1_2007.indd 70 12.6.2007 8:44:29 Daniela Michälkovä: Diversity of Dry Grasslands in the Povazsky Inovec Mts (Slovakia) - A Numerical Analysis Figure 7: Spring aspect of Festuco pallentis-Caricetum humilis. Photo: M. Janišova, 24. 5. 2005. Slika 7: Spomladnski aspekt asociacije Festuco pallentis-Caricetum humilis. Foto: M. Janišova, 24. 5. 2005. 3.1.1 Nomenclature and species characteristics of the associations (cf. Table 1) Cluster A - Festuco valesiacae-Stipetum capillatae Sil-linger 1930 Orig. (Sillinger 1930: 35): Festuceto (valesiacae)-Sti- petum capillatae Phantom name: Festuco valesiacae-Stipetum capillatae Sillinger 1931 (in Mucina & Maglocky 1985: 189), Stipo capillatae-Festucetum valesiacae Sillinger 1931 (in Maglocky 1979: 92) Nomenclatural type: Sillinger 1930: 36, rel. 3, holo-typus Remark: We considered the possibility of suggesting the inversion of the order of species names in Festuco vale-siacae-Stipetum capillatae Sillinger 1930. Festuca valesiaca seems to dominate over Stipa capillata in the growths of this association and therefore it should stay in the second position of the association's name (Art. 10b). However, Stipa capillata grows at the higher herb level compared to Festuca valesiaca and it dominates in a few stands in Southern Moravia (Chytry in litt.). For these reasons we decided to keep the name in its original form described by Sillinger (1930). Diagnostic taxa: Festuca valesiaca 77.8, Medicago falcata 74.4, Eryngium campestre 69.1, Securigera varia 61.5, Pimp-inella nigra 60.8, Acosta rhenana agg. 57.7, Thymus pan-nonicus 54.4, Festuca rupicola 52.9, Carex caryophyllea 49.8, Koeleria macrantha 48.5, Astragalus onobrychis 47.8, Medicago lupulina 47.8, Plantago lanceolata 46.7, Poa angustifolia 46.4, Salvia pratensis 45.7, Fragaria viridis 44.6, Teucrium chamaedrys 44.2, Adonis vernalis 41.5, Agrimonia eupatoria 41.5, Galium verum 41.5, Achillea collina 41.4, Ranunculus bulbosus 40.4 Constant taxa: Sanguisorba minor 83, Potentilla arenaria agg. 74, Tithymalus cyparissias 72, Asperula cynanchica 61, Pilosella officinarum 61 Dominant taxa: Festuca valesiaca 22, Bothriochloa ischae-mum 11 , Carex caryophyllea 9 Cluster B - Festuco pallentis-Caricetum humilis Sillinger 1930 corr. Gutermann et Mucina 1993 Orig. (Sillinger 1930: 28): Asociace Festuca glauca- Carex humilis ( Glauceto-Caricetum humilis) Syntax. syn.: Scabioso suaveolentis-Caricetum humilis Klika 1931 Nomenclatural type: Michâlkovâ, Skodovâ & Merta-novâ (2006): Table 1, rel. 12, lectotypus hoc loco. Diagnostic taxa: Alyssum montanum 55.0, Stipa joannis 67 HQ_6-1_2007.indd 63 12.6.2007 8:44:18 Hacquetia 6/1 • 2007, 61-76 51.5, Poa badensis 47.2, Colymbada scabiosa agg. 41.0, Carex humilis 40.5 Constant taxa: Helianthemum grandiflorum s. lat. 93, Thymus praecox 93, Sanguisorba minor 93, Globularia punctata 86, Tithymalus cyparissias 86, Potentilla arenaria agg. 83, Teucrium montanum 83, Festuca pallens 76, Leontodon inca-nus 69, Anthyllis vulneraria agg. 64, Linum tenuifolium 64, Asperula cynanchica 62 Dominant taxa: Carex humilis 55, Stipa joannis 12 Cluster C - Poo badensis-Festucetum pallentis Klika 1931 corr. Zolyomi 1966 nom. invers. propos. Orig. (Klika 1931: 360): Festuca glauca-Poa badensis-Assoziation Nomenclatural type: Klika 1931: 363, rel. 16, lecto- typus (assigned in Toman 1975) Diagnostic taxa: Fumana procumbens 54.8, Festuca pallens 50.5, Draba lasiocarpa 44.6 Constant taxa: Teucrium montanum 100, Thymus praecox 100, Helianthemum grandiflorum s.lat. 85, Leontodon inca-nus 85, Globularia punctata 65, Potentilla arenaria agg. 65 Dominant taxa: Festuca pallens 30, Poa badensis 5 Cluster D - Minuartio setaceae-Seslerietum caeruleae Klika 1931 nom. mut. propos. Orig. (Klika 1931: 367): Sesleria calcaria-Alsine seta-cea-Assoziation (Alsine setacea = Minuartia setacea, Sesleria calcaria = S. caerulea, Art. 45) Syn.: Seslerio-Caricetum humilis Sillinger 1930 (Art. 31), Carici humilis-Seslerietum calcariae Sillinger 1930 nom. invers. (Art. 31) Non: Carici humilis-Seslerietum calcariae Zlatnik 1928 Nomenclatural type: Klika 1931: 369, rel. 21, lecto- typus (assigned in Toman 1975) Diagnostic taxa: Sesleria albicans 96.9, Genista pilosa 74.2, Thalictrum minus 70.1, Thesium linophyllon 67.0, Phyteuma orbiculare 60.7, Allium senescens ssp. montanum 56.6, Th-laspi montanum 55.7, Bupleurum falcatum 51.2, Polygala amara ssp. brachyptera 51.0, Campanula moravica 50.0, Bro-mus monocladus 44.7, Carex humilis 40.5 Constant taxa: Teucrium montanum 100, Helianthemum grandiflorum s.lat. 88, Anthericum ramosum 81, Leontodon incanus 75, Sanguisorba minor 69, Globularia punctata 62 Dominant taxa: Sesleria albicans 100, Genista pilosa 25 3.2 Ordination and analysis of Ellenberg indication values The detrended gradient analysis (DCA) has been used to define the major gradients in the species composition of the dry grassland vegetation (Fig. 3 and 4, Table 2). Figure 4 shows the species scatter plot of DCA based on individual relevés. Ellenberg indication values, plotted onto the ordination diagram, show that the major variation in species composition of the dry grasslands corresponds to the combination of several gradients. The first axis of the ordination diagram interprets 7.6 % of the variability of the data set (lengths of gradient 4.763, eigenvalue 0.642) and it may be more or less associated with the temperature gradient. The second axis (lengths of gradient 3.587, eigenvalue 0.398) is associated with the soil reaction gradient, which negatively correlates to the moisture (which could be later combined with the soil nutrient availability gradient). Individual clusters strikingly differ in their relationships to major environmental factors (Fig. 3 and 4). Connected to the temperature gradient, vegetation of cluster A (Festuco valesiacae-Stipetum capillatae) occupies very warm sites, while cluster D (Minuartio setaceae-Seslerietum caeruleae) is confined to the coolest sites. The gradients of soil nutrients and moisture are mutually correlated, with cluster A (Festuco valesiacae-Stipetum capillatae) associated with high values and clusters B (Festuco pallentis-Caricetum humilis) and C ( Poo badensis-Festucetum pallentis) with low values. There is a relatively variable association of Fes-tuco valesiacae-Stipetum capillatae dissociated in the left lower part of the diagram. In comparison to the rest of the data set, this unit includes the vegetation of warm and moist stands, which are the richest in soil nutrients and the most continental (Fig. 5). This closed vegetation occurs in the stands with deeper soil, which indicates also its species composition (Table 1). These dry grasslands were used as pastures in the past. They occur in the near vicinity of the rural settlements on the slight slopes at lower altitudes. There are relevés of Minuartio setaceae-Seslerietum caeruleae separated in the right part of the ordination space (Fig. 4). This is the most cool-liking association, in which some mountain plant species are present. Sesleria albicans, Phyteuma orbiculare and Aci-nos alpinus are the examples of the dealpine species, while Leontodon incanus, Thlaspi montanum and Biscutella laevigata are the prealpine taxa. In comparison to the other associations, Minuartio setaceae-Sesleri-etum caeruleae grows in the humid stands and it does not have high light requirements (Fig. 5). It occurs at the highest altitudes in the NE to NW expositions. Bromus monocladus (B. pannonicus ssp. monocladus), a remarkable endemic of the Western Carpathians, prefers less extreme, mesophilous stands. 68 HQ_6-1_2007.indd 70 12.6.2007 8:44:29 Daniela Michälkovä: Diversity of Dry Grasslands in the Povazsky Inovec Mts (Slovakia) - A Numerical Analysis Table 2: The outputs of the detrended correspondence analysis (DCA) of the phytocoenological relevés of the dry grassland vegetation of the Povazsky Inovec Mts. Axes 1 2 3 4 Total inertia Eigenvalues: 0.642 0.398 0.329 0.192 8.461 Lengths of gradient: 4.763 3.587 3.533 2.228 Cumulative percentage variance of spec. data: Sum of all eigenvalues 8.461 7.6 12.3 16.2 18.5 There are two associations (Festuco pallentis-Cari-cetum humilis and Poo badensis-Festucetum pallentis) located in the central part of the diagram (Fig. 4). Their inter-separation is rather poorly presented in this projection of the ordination space. Both associations positively correlate to the increasing of the basic character of the substrate and negatively to the amount of soil nutrients and moisture. The more extreme open vegetation of Poo badensis-Festucetum pallentis naturally occurs on the dolomite and limestone screes with the minimal amount of soil available, which contains nutrients and holds water. 3.3 Exposition and inclination of the slopes The exposition of slopes is an important environmental factor, which influences the distribution of dry grassland associations in the study area (Ma-glocky 1978, Janisovä 2005). While the Minuartio setaceae-Seslerietum caeruleae evidently prefers the northern expositions, all the other associations tend to occur mostly on the south facing slopes (Fig. 8). The Festuco valesiacae-Stipetum capillatae occurs on Festuco valesiacae-Stipetum capillatae Festuco pulle ntis-Caricetum humilis NWW W SWW EEN EES SSW SSE S Poo badensis-Festucetum pallentis NWW SWW EEN EES Minuartio setaceae-Seslerietum caeruleae NWW W SWW EEN EES NWW W SWW EEN EES Figure 8: Circle diagrams indicating the slope and exposition priorities of the associations. Green circle scale measures show 20, 40, 60 and 80 %. The number values are marked starting from 9 %. Slika 8: Krožni diagrami prikazujejo nebesno lego in naklon rastišč posameznih asociacij. Zeleni krogi prikazujejo naklon: 20, 40, 60 and 80 %. Vrednosti so označene od 9 % naprej. 69 HQ_6-1_2007.indd 63 12.6.2007 8:44:18 N N E E S N N E S S Hacquetia 6/1 • 2007, 61-76 the slight slopes (Tab. 1, average inclination 11°) in the warm expositions (S, SW, W). The driest association Poo badensis-Festucetum pallentis prefers south to east facing slopes, which are relatively steep (average inclination 26°). Festuco pallentis-Caricetum humilis grows usually in south to west expositions, mostly at the ridges of the hills (Fig. 9). 4. SYNTAXONOMY Festuco-Brometea Br.-Bl. et Tx. ex Soo 1947 Festucetalia valesiacae Br.-Bl. et Tx. ex Br.-Bl. 1949 Festucion valesiacae Klika 1931 Festuco valesiacae-Stipetum capillatae Sillinger 1930 Bromo pannonici-Festucion pallentis Zolyomi 1936 corr. 1966 Festuco pallentis-Caricetum humilis Sillinger 1930 corr. Gutermann et Mucina 1993 Poo badensis-Festucetum pallentis Klika 1931 corr. Zolyomi 1966 nom. invers. propos. Diantho lumnitzeri-Seslerion abicantis (Soo 1971) Chytry et Mucina in Mucina et Kolbek 1993 Minuartio setaceae-Seslerietum caeruleae Klika 1931 nom. mut. propos. The classification of the associations within the class Festuco-Brometea is rather problematic. The traditional conception of the class division into orders and alliances (cf. Mucina & Maglocky 1985) is not identical to the more recent works (cf. Mucina & Kolbek 1993, Michâlkovâ & Šibik 2006, Chytry et al. 2007). Only the classification of the Festuco valesiacae-Stipetum capillatae appears to be definite, because both in the traditional and modern literature sources, the association is classified within the subcontinental alliance Festucion valesiacae (cf. Maglocky 1979, Mucina & Maglocky 1985, Moravec et al. 1995, Chytry et al. 2007). The author of the description of Festuco pal-lentis-Caricetum humilis did not classify this association within any higher syntaxon. At the time of its publication in Sillinger (1930), no alliances of the class Festuco-Brometea had yet been described. On the other hand, the association Scabioso suaveolentis-Caricetum humilis (syntax. syn. of the Festuco pallen-tis-Caricetum humilis) was included by the author of its description in Festucion valesiacae (Klika 1931). Some authors later followed this conception (Maglocky 1979, Mucina & Maglocky 1985). These dry grasslands are classified in the works of Mucina & Kolbek (1993: 475) and Chytry et al. (2007) within Bromo pannonici-Festucion pallentis. This alliance includes the circum-Pannonian thermophilous grassland biotopes on dolomite bedrocks. The association Poo badensis-Festucetum pallentis is classified in Mucina & Kolbek (1993: 473) and Chytry et al. (2007) within the alliance Bromo pan-nonici-Festucion pallentis. On the other hand, in older works (Klika 1931, Maglocky 1979, Mucina & Maglocky 1985) it was considered as a part of the broadly defined alliance Seslerio-Festucion pallentis Klika 1931 corr. Zolyomi 1966. Considering the nomenclatural remark in Mucina & Kolbek (1993: 460), we regard it as a nomen ambiguum. According to the presence of numerous diagnostic taxa of the alliance Bromo pannonici-Festucion pallentis in the vegetation of both associations (cf. Table 1), we agree with this newer concept. The alliance Seslerio-Festucion pallentis has been divided into two alliances in recent literature. One of them is Bromo pannonici-Festucion pallentis. The other, Diantho lumnitzeri-Seslerion albicantis, includes the closed dealpine xerophilous vegetation on the limestone bedrock, mostly dominated by Sesleria al-bicans. The association Minuartio setaceae-Seslerietum caeruleae belongs to this alliance (cf. Mucina & Kolbek 1993, Chytry et al. 2007). 5. DISCUSSION AND CONCLUSION The results of the presented numerical analyses support the traditional conceptions (Sillinger 1930, Klika 1931, Maglocky 1979), which were created through the deductive classification methods. It is very important that the clusters representing individual associations include the relevés of the original association description (cf. Fig. 2, sources of relevés). The relevés of the original description of the Festuco valesiacae-Stipetum capillatae (cluster A) are in Sillinger (1930: 36, rel. 1, 2 and 3). Cluster B (Festuco pallentis-Caricetum humilis) contains the relevés of the original description also from Sillinger (1930: 36, rel. 4, 5, 6 and 7). Cluster C (Poo badensis-Festucetum pallentis) includes a relevé from Klika (1931: 363, rel. 13). Cluster D (Minuar-tio setaceae-Seslerietum caeruleae) does not involve any relevés from Sillinger (1930), because there is only a synoptic table in this work. However, cluster D contains only one relevé from the Považsky Inovec Mts, which occurs in Klika (1931: 370, rel. 29). On the other hand, the cluster analyses did not support the validity of the numerous subasso- 70 HQ_6-1_2007.indd 70 12.6.2007 8:44:29 Daniela Michälkovä: Diversity of Dry Grasslands in the Považsky Inovec Mts (Slovakia) - A Numerical Analysis Figure 9: Influence of slope exposition on the distribution of the dry grassland associations. In front is the Festuco pallentis-Caricetum humilis. The Poo badensis-Festucetum pallentis is marked by a red rectangle in the background. Locality Hradlova nivka (township of Luka). Photo: M. Janišova, 7. 5. 2005. Slika 9: Vpliv naklona in nebesne lege na razširjenost asociacij suhih travnikov. V ospredju sestoji asociacije Festuco pallentis-Caricetum humilis. Sestoj asociacije Poo badensis-Festucetum pallentis je označen z rdečim pravokotnikom v ozadju. Lokacija Hradlova nivka (občina Luka). Foto: M. Janišova, 7. 5. 2005. ciations and variants assigned in Maglocky (1979). According to Art. 5 of the International Code of Phytosociological Nomenclature (Weber, Moravec & Theurillat 2000), these subassociations were in-validly described, because there was no nomencla-torical typus designated in the paper. In this study, only some relevés of Maglocky's subassociations aggregated at the lower hierarchical level of the dendrogram (Fig. 2, numbers 1-6). In the Festuco vale-siacae-Stipetum capillatae festucetosum rupicoae, 8 from 9 relevés published in Maglocky (1979) grouped together. In the F. v.-S. c. botryochloetosum ischaemii it was 6 from 9 relevés and in the F. v.-S. c. cariceto-sum caryophyllae 6 from 7 relevés (Fig. 2, numbers 1-3). None of the subassociations described in the dry grasslands, which Maglocky named Sca-bioso suaveolentis-Caricetum humilis, were confirmed by the analysis. Only 5 from 8 relevés of S. s.-C. h. typicum, Koeleria macrantha-variant aggregated in the dendrogram (Fig. 2, number 4). All seven relevés of Poo badensis-Festucetum pallentis jovibarbeto-sum hirtae from Maglocky (1979) gathered together. Moreover two other relevés joined this group (Fig. 2, number 5). A small cluster was created by all of the three relevés of Minuartio setaceae-Seslerie-tum caeruleae (syn. Carici humilis-Seslerietum calcariae) festucetosum pallentis (Fig. 2, number 6). This study does not attempt to solve the problems of variability within the studied associations. For this purpose, it is necessary to analyse a representative data set consisting of the relevés from the whole area of the associations' occurrence. Nowadays, the dry grasslands in the Považsky Inovec Mts have to face many threats. First of all, the abandonment of the pastures (mostly Festuco valesi-acae-Stipetum capillatae) causes successional changes in the species composition as well as shrub progression. Mostly deer and mouflons ( Ovis ammon subsp. 71 HQ_6-1_2007.indd 71 12.6.2007 8:44:31 Hacquetia 6/1 • 2007, 61-76 musimon) graze the vegetation of Festuco pallentis-Caricetum humilis, which can be found at the higher altitudes, away from the settlements. However, the non-native mouflons, introduced into this area in the past, can cause the erosion (particularly in the Poo badensis-Festucetum pallentis) and increase the amount of nitrogen in the soil. In the past decades, an intensive planting of non-native Pinus nigra and Fraxinus ornuswas accomplished in large areas. This destroyed or eventually reduced the size of many sites where mostly Festuco pallentis-Caricetum humilis and Minuartio setaceae-Seslerietum caeruleae had occurred. In some areas, the grasslands of Festuco vale-siacae-Stipetum capillatae were completely destroyed by ploughing in order to intensify the agricultural production in the communistic times (Maglocky in verb., Devan et al. 2006). Because of the consistent reduction of the area of the dry grasslands, it is necessary to continue with the management activities. Although this vegetation is secondary and human-influenced, it represents a refuge for many rare thermophilous species and it is a valuable contribution to the diversity of habitats in Slovakia. ACKNOWLEDGEMENTS The author is grateful to Milan Chytry, Monika Janišova, Jin Kolbek, Jozef Šibik, Milan Valachovič and Maria Zaliberova for their valuable comments on the manuscript and willingness to help with the numerical analyses. I also thank Jana Sadlonova for helping to create the charts and Sylvia Mertanova who made the map. Lucia Kwon performed the language correction. For intensive help in solving the nomenclatural problems, I am deeply indebted to Jan Kliment. The research study was funded by projects APVT-51-015804 and VEGA 2/5084/25. REFERENCES Braun-Blanquet,J. 1964: Pflanzensoziologie. Grundzüge der Vegetationskunde. Ed. 3. Springer Verlag, Wien, 865 pp. Chytry, M., Hoffmann, A. & Novak, J. 2007: Suché travniky. Tfîda Festuco-Brometea. In: Chytry M. (ed.): Vegetace Ceské republiky. 1. Travinna a keričkova vegetace. Academia, Praha. 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Sci., Uppsala 11: 739-768. Westhoff, V., van der Maarel, E. 1973: The Braun-Blanquet approach. In: Whittaker, R. H. (ed.): Ordination and Classification of Communities. Dr. W. Junk Publishers, The Hague, pp. 617727. Recieved 21. 11. 2006 Revision recieved 29. 1. 2007 Accepted 19. 3. 2007 73 HQ_6-1_2007.indd 71 12.6.2007 8:44:31 Hacquetia 6/1 • 2007, 61-76 Table 1: A synoptic table produced by cluster analysis. The values are percentage frequencies. The fidelities (9 coefficient multiplied by 100) are in upper index. Tabela 1: Sinoptična tabela narejena na osnovi rezultatov klastrske analize. Vrednosti so frekvence v odstotkih. Nadpisana vrednost je navezanost (9 koeficient pomnožen s 100). Association (cluster) A B C D Number of relevés Average number of species Average inclination_ 46 34 11° 42 32 22° 20 20 26° 16 26 28° Festuco valesiacae-Stipetum capillatae Festuca valesiaca Medicago falcata Eryngium campestre Securigera varia Pimpinella nigra Acosta rhenana agg. Thymus pannonicus Festuca rupicola Carex caryophyllea Koeleria macrantha Astragalus onobrychis Medicago lupulina Plantago lanceolata Poa angustifolia Salvia pratensis Fragaria viridis Teucrium chamaedrys Galium verum Adonis vernalis Agrimonia eupatoria Achillea collina Ranunculus bulbosus Festuco pallentis-Caricetum humilis 87 77,8 21 - -- . 76 74,4 7 - -- 6 63 69,1 7 - -- . 48 61,5 2- -- . 63 60,8 19 - -- . 50 57,7 2- -- 6 39 54,4 2- -- . 50 52,9 2- -- 12 39 49,8 7- -- . 72 48,5 43 - -- 15 - -- . 28 47,8 . - -- . 28 47,8 . - -- . 35 46,7 .- -- 6 37 46,4 2- -- 6 26 45,7 .- -- . 33 44,6 .- -- 6 80 44,2 52 - -- 38 22 41,5 .- -- . 22 41,5 .- -- . 22 41,5 .- -- . 35 41,4 7- -- 5 - -- . 24 40,4 2- -- . Alyssum montanum B-F 37 - - 81 55 5 --- 19 --- Stipa joannis 4- -- 38 51,5 --- --- Poa badensis B-F 7- -- 40 47,2 5 --- --- Colymbada scabiosa agg. 11 - -- 33 41 --- --- Poo badensis-Festucetum pallentis Festuca pallens B-F 13 - -- 76 22,7 100 50,5 38 --- Fumana procumbens B-F 7- -- 57 21,8 85 54,8 6 --- Draba lasiocarpa -- 21 --- 50 44,6 6 --- Minuartio setaceae-Seslerietum caeruleae Sesleria albicans D-S -- 5 --- --- 100 96,9 Genista pilosa 17 - -- 29 --- 5 --- 100 74,2 Thalictrum minus -- . --- --- 56 70,1 Thesium linophyllon 2- -- 7 --- --- 62 67 Phyteuma orbiculare D-S -- . --- --- 44 60,7 Allium *montanum D-S 4- -- 31 --- 10 --- 75 56,6 Thlaspi montanum D-S -- . --- --- 38 55,7 Bupleurum falcatum 7- -- 29 --- --- 62 51,2 Polygala *brachyptera D-S 4- -- . --- --- 38 51 74 HQ_6-1_2007.indd 76 12.6.2007 8:44:36 Daniela Michälkovä: Diversity of Dry Grasslands in the Považsky Inovec Mts (Slovakia) - A Numerical Analysis Association (cluster) A B C D Campanula moravica Bromus monocladus Festucion valesiacae Carex humilis Silene otites Asperula cynanchica Erysimum odoratum Pilosella bauhinii Alliumflavum Scabiosa ochroleuca Jurinea mollis Onosma visianii Stachys recta Veronica prostrata Pseudolysimachion spicatum Artemisia campestris Seseli hippomarathrum Vincetoxicum hirundinaria Bothriochloa ischaemum Dianthus carthusianorum Stipa capillata Arenaria leptoclados Elytrigia intermedia Medicago minima Bromo pannonici-Festucion pallentis Thymus praecox Teucrium montanum Melica ciliata Potentilla arenaria Campanula sibirica Scorzonera austriaca Sedum album Jovibarba *glabrescens Scabiosa canescens Diantho lumnitzeri-Seslerion albicantis Leontodon incanus Biscutella laevigata Seslerio-Festucion pallenis Anthericum ramosum Linum tenuifolium Seseli osseum Dianthus *lumnitzerii Minuartia rubra Dorycnium herbaceum agg. Cyanus triumfetti Galium glaucum Saxifraga tridactylites Hornungia petraea Festucetalia valesiacae Inula ensifolia Linaria genistifolia 28 15 61 11 33 24 52 4 2 15 22 4 17 15 39 28 33 7 17 17 37 28 20 74 11 4 2 7 2 17 4 7 33 22 7 13 17 2 9 2 9 15 37,1 39,5 36,9 36,9 24 100 55 62 17 38 40 40 26 36 26 7 21 10 12 31 24 7 36 10 93 83 57 83 52 48 24 38 24 69 19 60 64 48 38 17 12 5 7 2 19 33 24 40,5 34,2 33,3 33,5 23,8 38,8 60 40 30 55 5 60 45 20 25 20 10 5 100 100 45 65 50 5 45 45 5 85 40 45 30 30 45 20 15 10 10 39,3 30,8 88 25 100 12 38 19 44 19 44 12 25 6 6 12 44 100 38 19 25 25 19 19 75 50 81 25 31 38 6 6 19 25 6 50 44,7 40,5 75 HQ_6-1_2007.indd 71 12.6.2007 8:44:31 7 5 Hacquetia 6/1 • 2007, 61-76 Association (cluster) A B C D Viola collina Veronica austriaca Lactuca viminea Festuco-Brometea Helianthemum grandiflorum Sanguisorba minor Tithymalus cyparissias Carlina vulgaris Acinos arvensis Arenaria serpyllifololia Allium sphaerocephalon Arabis hirsuta Pimpinella saxifraga Plantago media Potentilla heptaphylla Trinia glauca Seseli annuum Leontodon hispidus Trifolium campestre Crinitina linosyri Avenula pubescens Phleum phleoides Other taxa Pilosella officinarum Globularia punctata Anthyllis vulneraria Sedum acre Asperula tinctoria Sedum sexangulare Thlaspi perfoliatum Arabis auriculata Cerastium brachypetalum Cerastium pumilum Pulsatilla grandis Acinos alpinus Galium austriacum Hypericum perforatum Euphrasia stricta agg. Hippocrepis comosa Pulsatilla slavica Inula hirta Silene vulgaris Linum catharticum Sideritis montana Chondrilla juncea 17 48 83 72 17 30 13 2 20 11 28 17 17 20 20 17 11 9 20 61 15 37 11 2 46 37 24 7 7 37 13 9 2 2 2 30 17 15 35,1 35.7 30.8 93 93 86 10 10 26 7 31 10 10 12 7 2 10 10 38 86 64 12 10 33 29 5 7 7 7 2 5 24 5 5 7 5 2 33,4 29 10 85 50 60 5 5 25 5 25 65 20 20 5 25 20 15 25 5 10 15 20 31,9 36,9 34,4 31 19 88 69 50 6 12 19 44 44 6 62 38 6 25 12 19 19 6 19 12 6 6 6 19 5 Abreviations: A - Festuco valesiacae-Stipetum capillatae Sillinger 1930, B - Festuco pallentis-Caricetum humilis Sillinger 1930 corr. Gutermann et Mucina 1993, C - Poo badensis-Festucetum pallentis Klika 1931 corr. Zolyomi 1966 nom. invers. propos., D - Minuartio setaceae-Seslerietum caeruleae Klika, B-F - Bromo pannonici-Festucion pallentis Zolyomi 1936 corr. 1966, D-S - Diantho lumnitzeri-Seslerion albicantis (Soo 1971) Chytry et Mucina in Mucina et Kolbek 1993. 76 HQ_6-1_2007.indd 76 12.6.2007 8:44:36