Polish Botanical Journal 49(2): 181–202, 2004

SYNTAXONOMICAL REVISION OF COMMUNITIES WITH FIRMA AND (ALLIANCE CARICION FIRMAE) IN THE WESTERN CARPATHIANS

JOZEF ŠIBÍK, ANTON PETRÍK & JÁN KLIMENT

Abstract: A syntaxonomical revision of plant communities with dominants Carex fi rma and Dryas octopetala in the Western Carpathians is presented, including a comparison with related plant communities from the Alps. The syntaxonomical position of the studied communities has been re-evaluated, and a new classifi cation concept is offered. Communities from the subalpine belt of the Krivánska Malá Fatra, Chočské vrchy and Nízke Tatry Mts are classifi ed within the association Dryado octopetalae- Caricetum fi rmae Sillinger 1933, and two new subassociations primuletosum auriculae and saxifragetosum aizoidis are described. The plant communities from the subalpine and alpine belts of the Západné Tatry and Belianske Tatry Mts are different. We classify them within the association Arenario tenellae-Caricetum fi rmae (Br.-Bl. 1930) nom. nov., with two subassociations recognized: typicum and salicetosum reticulatae. Phytocoenological relevés obtained in recent years confi rm the well-pronounced separation of the association Androsaceo lacteae-Festucetum versicoloris Sillinger 1933. According to our revision, phytocoenoses in the montane belt or on inversion sites of the Veľká Fatra, Muránska planina and Slovenský raj Mts, including some communities of the subalpine belt of the Tatras, cannot be classifi ed in the alliance Caricion fi rmae Gams 1936, but belong to the alliances Seslerio-Asterion alpini Hadač ex Hadač et al. 1969, Potentillion caulescentis Br.-Bl. in Br.-Bl. & Jenny 1926 emend. Sutter 1969 and Cystopteridion Richard 1972. Key words: Caricion fi rmae, alpine vegetation, syntaxonomy, numerical classifi cation, Alps, Western Carpathians, Slovakia Jozef Šibík, Institute of Botany, Slovak Academy of Sciences, Dúbravská cesta 14, SK-845 23 Bratislava, Slovak Republic; e-mail: [email protected] Anton Petrík, Botanical Garden of Comenius University, Botanická 3, SK-841 04 Bratislava, Slovak Republic Ján Kliment, Botanical Garden of Comenius University, SK-038 15 Blatnica, Slovak Republic; e-mail: [email protected]

INTRODUCTION

In the Western Carpathians, the distribution center the marked similarity of fl oristic composition be- of communities with dominance of Carex fi rma tween stands of these two dominant in the and Dryas octopetala occurs in the subalpine and Western Carpathians, Sillinger (1933) treated them alpine belts. These communities occur especially as facies of the association Dryadeto-Firmetum. on north-facing steep slopes and in rocky grooves, In comparing the vegetation of the Central Alps but can be found also on inversion sites or on and the Tatra Mts, Braun-Blanquet (1930) used steep south-facing rocky walls. Initial syntaxo- the name Caricetum fi rmae tatricum to designate nomical studies of these communities appeared phytocoenoses from the Tatras. Pawłowski (1935) in the 1920s. Using the criterion of the dominance employed the geographical epithet Caricetum of one of the determinant species, Szafer et al. fi rmae carpaticum to delimit and distinguish West (1923) described associations Caricetum fi rmae Carpathian stands from those of the Alps. Hadač et and Dryadetum from the Polish side of the West al. (1969) distinguished two units in the Belianske Tatras. Pawłowski and Stecki (1927) and Klika Tatry Mts, namely Caricetum fi rmae carpaticum (1932) considered stands with dominance of Dryas and Dryadetum octopetalae tatricum. Two names octopetala to be only a facies or stage of the as- proposed by Hadač (cf. Mucina & Maglocký 1985), sociation Caricetum fi rmae (Firmetum). Based on Saxifrago caesiae-Caricetum fi rmae and Festuco 182 POLISH BOTANICAL JOURNAL 49(2). 2004 versicoloris-Dryadetum octopetalae, reversed the alpestris), Pulsatilla slavica (P. subslavica), Schistidium original physiognomical classifi cation. apocarpum (S. atrofuscum, S. strictum) and Thymus In recent decades, detailed fi eld research on pulcherrimus (subsp. pulcherrimus, subsp. sudeticus). communities with dominance of Carex fi rma Numerical classifi cation employed the NCLAS program from the SYN-TAX 5 package (Podani 1993). The β-fl ex- and Dryas octopetala has been carried out in the ible method (β = -0.25) with Jaccard’s, Ružička’s and central part of the Western Carpathians and their Wishart’s similarity coeffi cients was used. The hypoth- foothills. These studies, followed by numerical eses obtained were evaluated by comparison and through classifi cation, have brought new insights for the analysis of phytocoenological tables. syntaxonomical classifi cation of these communi- To determine the content of the alliance Caricion ties, which we present in this paper. fi rmae and the syntaxonomical position of other com- munities with the dominance of Carex fi rma and Dryas octopetala, our phytocoenological relevés were also MATERIAL AND METHODS compared with closely related plant communities of the alliance Seslerio-Asterion alpini (Elyno-Seslerietea) The syntaxonomical revision presented here included and classes Asplenietea trichomanis, Juncetea trifi di and 413 phytocoenological relevés of plant communities Carici rupestris-Kobresietea from the Western Carpath- with the dominance and higher abundance of Carex ians, and to a lesser extent also from the Eastern and fi rma and Dryas octopetala, from the central part of Southern Carpathians and the Alps. the Western Carpathians and their foothills. Since these The synoptic table (Table 1) shows the combined communities with Carex fi rma have been often merged results of several partial syntheses, which could not with the association Caricetum fi rmae Rübel 1911 from be presented as a single dendrogram due to the high the Alps (cf. Grabherr & Mucina 1993: 406), we also number of relevés. The caption of each column contains included 21 relevés of this association originating from the number of relevés used for the synthesis and the the Alps. In this study, only differences between the average number of species in a given community. The Alpine and West Carpathian communities are high- frequency as a percentage (99 = 100%) and mean value lighted, so the Alpine association and its diagnostic of abundance (upper index) are given for each taxon. taxa are not evaluated in more detail. All relevés used They were calculated with FYTOPACK (Jarolímek in this study were collected in accordance with the prin- & Schlosser 1997). If only four or fewer relevés were ciples of the Zürich-Montpelliér school (Braun-Blan- available (column Ds), only the presence of taxa in a given quet 1964), but applying different scales of abundance community is indicated, in italics. Individual columns and dominance used by various authors: the fi ve- or also contain abbreviated references (for unpublished data seven-degree scale of Braun-Blanquet, the 10-degree only author names are given), the number of relevés, and or a combined 11-degree scale of Hadač and Domin their position on the level of orographic units according (cf. Sillinger 1933; Hadač et al. 1969), and a modifi ed to the map for the Database of Fauna of Slovakia (scale 9-degree scale (Barkman et al. 1964). 1:500,000). Diagnostically important taxa of individual To obtain data comparable for numerical classifi ca- plant communities are marked in bold. tion, all relevés were transformed to a nine-degree ordinal The nomenclature of taxa generally follows Mar- scale (van den Maarel 1979). Taxa determined only at hold and Hindák (1998); names of taxa occurring only the level of genus were excluded, and some taxa were in the Alps follow Ehrendorfer (1973). Subspecies (given classifi ed within higher or more broadly defi ned taxa: without the species epithet) appearing in Table 1 in indi- Anthoxanthum odoratum agg. (A. alpinum, A. odoratum), vidual relevés (holotype) or repeating in the text are aster- Carex sempervirens (subsp. tatrorum, subsp. silicicola), isked (*). Names of syntaxa are according to Mucina and Cardaminopsis arenosa agg. (C. borbasii), Empetrum Maglocký (1985); the names of diagnostic taxa of higher hermaphroditum (E. nigrum), Gentianella lutescens rank follow recent synthetic studies from the Tatra Mts (subsp. tatrae), Helianthemum grandifl orum (subsp. gran- (Hadač et al. 1969; Unar et al. 1985), Slovakia (Valachovič difl orum, subsp. glabrum, subsp. obscurum), Jovibarba et al. 1995; Dúbravcová 1996), the Eastern Alps (Grabherr globifera (subsp. glabrescens), Leucanthemum vulgare & Mucina 1993) and Romanian Carpathians (Coldea et al. agg. (L. margaritae), Lotus corniculatus (var. alpicola 1997). The names of newly distinguished and described Beck), Luzula luzuloides (subsp. luzuloides, subsp. syntaxa include authors’ citations. rubella), Pimpinella major (subsp. rhodochlamys), Soli- In the descriptions of communities, the following dago virgaurea (subsp. minuta), Swertia perennis (subsp. abbreviations are used: Art. = article of International J. ŠIBÍK ET AL.: SYNTAXONOMICAL REVISION OF PLANT COMMUNITIES 183

Code of Phytosociological Nomenclature (Weber et al. NOMENCLATURAL TYPE: Braun-Blanquet 1930: 25. 2000); Ass. = association; const. = constant companion taxon (frequency higher than 60%); dif. = differential Two-layered dwarf shrub-herbaceous plant taxon; dom. = dominating taxon; r. = relevé. community, usually open. Along with tuft-rosette- forming hemicryptophytes, cushion-forming and DESCRIPTION OF PLANT COMMUNITIES tufted chamaephytes, small dwarf shrubs with dense branches are also present. Among them, Elyno-Seslerietea Br.-Bl. 1948 Dryas octopetala together with the dominant Seslerietalia coeruleae Br.-Bl. in Br.-Bl. & Jenny sedge, Carex fi rma, determine the physiognomy 1926 of this community. Several other herb species are represented in the community as well (Si- Caricion fi rmae Gams 1936 lene acaulis, Arenaria tenella, Saxifraga caesia, Arenario tenellae-Caricetum fi rmae (Br.-Bl. Minuartia gerardii, Androsace chamaejasme, 1930) nom. nov. hoc loco Pedicularis oederi, Ranunculus breyninus, R. al- pestris, Minuartia sedoides, etc.). The layer of Table 1, column B mosses and lichens (E0) is enriched by species BASIONYM: Caricetum fi rmae tatricum Br.-Bl. 1930 like Vulpicida tubulosus and Dactylina madrepori- (Art. 34). formis, contributing to the color variation of the SYNONYMS: Firmetum Pawłowski & Stecki 1927 p.p. community. Most reach height of 3–10 cm; maj. (Art. 31), Firmetum (= Caricetum fi rmae) car- only fl owering stems or stems of some grasses paticum Pawłowski 1935 p.p. maj. (Art. 34), Caricetum overreach them to 10–25 cm in height. On sites fi rmae Šmarda 1956 (Art. 31), Dryadetum octopetalae exposed to extremely strong winds, a conspicuous tatricum Hadač et al. 1969 (Art. 34), Caricetum fi rmae Domin 1936 (Art. 2b, 31), Saxifrago caesiae- belt-and-terrace structure of the vegetation can be Caricetum fi rmae (Szafer et al. 1923) Hadač in Mucina observed. Narrow vegetation belts are situated on & Maglocký 1985 p.p. maj. (Art. 2b), Festuco versi- the steep front of a terrace, which retains exposed coloris-Dryadetum (Szafer et al. 1923) Hadač in Mucina gravel material. & Maglocký 1985 p.p. min. (Art. 2b), Carex fi rma-ass. The optimum of chionophobic sub-hygrophi- Domin 1923 (nom. nud.), Dryadeto-Caricetum fi rmae lous plant communities of the association Arenario Domin 1925a (nom. nud.), Caricetum fi rmae Domin tenellae-Caricetum fi rmae occurs in the alpine and 1925b (nom. nud.), Dryadetum Domin 1925b (nom. subalpine belts (alt. 1550–2150 m) in limestone- nud.), Caricetum fi rmae (= Firmetum) Domin 1928 dolomite parts of the Tatra Mts. They inhabit (nom. nud.), Caricetum fi rmae tatrense Domin 1930, 1931 (nom. nud.), Caricetum fi rmae tatricum Klika very shallow to shallow, strongly humic, neutral 1932 (nom. nud.). – NON: Caricetum fi rmae Rübel to slightly basic, skeletal soils, in the winter cov- 1911, Caricetum fi rmae Szafer et al. 1923, Dryadetum ered with snow only briefl y or without any snow. Szafer et al. 1923. Due to the humid and cold climate, processes of DIAGNOSTIC TAXA: Carex fi rma (dom., const.), Dryas plant material decomposition and formation of octopetala (dom., const.), Androsace chamaejasmae fi ne humic earth are very slow. Plant communi- (dif.), Arenaria tenella (dif.), Cerastium eriophorum ties of this association mostly inhabit moderate to (dif.), alpina (dif.), Lloydia serotina steep, NW–NE-facing windy slopes. More rarely (dif.), Minuartia gerardii (dif.), M. sedoides (dif.), Pe- they occur on sites oriented in other directions: dicularis oederi (dif.), Rhodax alpestris (dif.), Silene in these cases they inhabit steep rocky walls with acaulis (dif.), Alectoria ochroleuca (dif.), Dactylina a minute fi ne earth layer deposited in fi ssures madreporifi rmis (dif.), Thamnolia vermicularis (dif.), Vulpicida tubulosus (dif.), Bistorta vivipara (const.), and on exposed terraces. On slopes with a suf- Crepis jacquinii (const.), Festuca versicolor (const.), fi ciently thick layer of detritus exposed to harsh Galium anisophyllon (const.), Ranunculus alpestris high-mountain climate with strong winds, forma- (const.), Saxifraga caesia (const.), Ditrichum fl exicaule tion of festooned soils typical for this community (const.), Tortella tortuosa (const.). is observed (cf. Šmarda 1956: 27). 184 POLISH BOTANICAL JOURNAL 49(2). 2004 ,

1 2 3 2 2 2 3 2 2 2 2 2 2 ...... D-C. 3 8 5 18 90 63 48 13 60 48 65 15 , C1 – 3 3 3 2 4 2 2 2 2 2 2 2 2 3 3 2 2 2 2 2 2 . 7 9 2 rmae 11 11 13 18 33 20 22 71 31 fi Seslerio-Asterion alpini / 3 2 2 2 3 2 2 1 2 2 1 3 2 . 2 . . 2 . . 2 . .36 .31 .24 . . 7 .67 9 9 9 9 9 9 27 91 18 18 27 Dryado octopetalae-Salicetum Rübel 1911 (A) and other plant Rübel 1911 6 3 3 2 3 4 2 2 3 2 3 3 3 2 2 2 2 2 3 3 2 2 3 rmae .73 . .11 5 . .99 fi 8 8 3 5 5 11 99 97 29 95 24 74 89 74 21 99 18 66 74 66 45 34 55 ...... Caricetum 2 4 2 4 3 1 2 2 Dryado octopetalae-Caricetum Potentillion caulescentis , C – 2 2 2 3 3 2 2 3 2 2 2 ...... ), G – 11 11 11 11 33 56 99 78 78 56 56 (Ds – Sillinger 1933: 229–230), E 3 3 2 2 3 2 3 2 3 2 2 2 2 2 2 2 2 5 3 9 9 2 82 77 68 41 47 18 18 (B–Ds) with association 3 3 2 3 2 1 3 2 2 2 2 2 .10 . 6 . 8 . 3 . 2 8 3 8 3 11 11 89 83 61 42 31 33 rmae fi A-C. salicetosum reticulatae Potentillion caulescentis 3 3 2 2 3 2 3 2 3 2 2 2 2 2 2 2 1 ...... ( 6 4 2 4 8 8 2 17 10 13 77 73 73 40 58 25 , B2 – Caricion (E–H) from the West Carpathians (a brief synoptic table). West (E–H) from the 3 2 2 2 3 2 2 3 2 2 2 2 2 3 2 2 2 2 2 2 2 2 3 2 2 5 7 3 11 73 64 60 60 59 54 53 46 42 34 29 25 24 23 13 34 12 19 58 46 41 A-C. typicum 3 2 2 2 3 3 2 3 2 2 2 3 2 3 2 2 2 2 2 2 2 2 2 2 2 8 78 83 58 39 38 66 58 45 43 25 38 34 29 12 24 48 10 15 23 32 21 12 Androsaceo lacteae-Festucetum versicoloris Androsaceo Dryas octopetala , B1 – 4 3 2 2 3 3 2 2 2 2 2 2 2 2 3 3 2 2 2 2 2 2 3 2 3 and , D – rmae 86 70 68 80 41 49 48 40 43 20 14 71 fi rmae fi rma fi 2 3 1 2 1 2 2 1 1 2 .82 .19 . .10 . 1 . 7 . 6 . . 8 ...... 13 .16 .33 .20 . 2 ?46 ?63 Caricion 5 5 38 10 57 90 10 71 38 38 Carex Carex Drabo tomentosae-Artemisietum petrosae ) 0 (E ) 0 ) ), F – 0 ) (E 0 ) (E 0 (E (E D-C. saxifragetosum aizoidis Arenario tenellae-Caricetum Arenario , C2 – , B – Seslerio-Asterion alpini / ora fl rmae fi Festucion versicoloris eldia calyculata fi (? Dactylina madreporiformis Dactylina madreporiformis Hypnum cupressiforme Poa alpina Silene acaulis tenella Arenaria Rhodax alpestris Minuartia sedoides Thamnolia vermicularis Minuartia gerardii Cerastium eriophorum Dianthus nitidus To lactea Androsace Saxifraga moschata bi Viola Cortusa matthioli Cystopteris montana alpestre Trisetum Euphrasia salisburgensis Ranunculus breyninus Alectoria ochroleuca Pedicularis oederi chamaejasme Androsace tubulosus Vulpicida alpina Chamorchis Lloydia serotina . Comparison of the Western Carpathian plant communities of the alliance Western . Comparison of the Caricetum Cystopteridion CommunityNumber of relevés species numberAverage taxa of the associations alliance Differential cf, fv cf, pt cf 25oe, cf 21 B1 A 36 90cf B2 42 92 B 39 182 oe fv, C1 33Sc, Pc 52cy C2 35 36 C 34pc 88fv D 27(cy) 9cy Ds ?aa 4 taxa of the subassociations Differential ESc, pc 61 38Sc Sc F 22 11 G 35 45 37 H 40 cf, fv cf cf cf oe fv, primuletosum auriculae reticulatae communities with higher abundance of species A – H – Table 1 Table J. ŠIBÍK ET AL.: SYNTAXONOMICAL REVISION OF PLANT COMMUNITIES 185

2 2 3 3 3 3 3 2 3 5 2 2 3 3 2 1 1 2 2 6 6 ...... 5 3 8 3 58 18 63 38 88 65 45 85 95 78 40 13 15 28 18 85 20 (cont.) 2 2 3 2 3 2 3 2 2 2 3 2 3 5 2 2 2 2 2 2 5 2 2 2 3 3 3 2 6 5 . 9 27 73 78 49 20 22 78 42 2 3 3 3 2 2 2 2 5 2 . 7 .42 .11 .44 .16 . . 9 . .44 . 4 . 2 . 7 .13 . 7 .24 . .13 . 2 .49 . 2 18 18 18 99 27 2 6 2 3 3 3 5 2 5 1 2 3 3 2 3 2 2 2 3 2 5 2 3 5 5 .27 .45 .55 . .78 .99 . . 9 . . . 7 . . 9 . . 83 . . . 3 8 5 18 37 84 95 87 61 32 71 99 16 16 29 89 45 18 16 39 71 13 26 76 39 ...... 4 2 4 2 3 3 3 1 1 1 2 2

.. . . 4 6 3 3 3 2 2 2 2 3 5 2 2 5 5 3 3 5 ...... 11 11 11 11 11 11 11 33 89 89 44 67 22 22 44 33 67 5 5 2 3 5 3 2 3 3 2 2 2 3 2 2 2 2 2 2 5 3 5 3 2 2 3 2 3 2 5 5 7 6 8 18 73 42 26 73 58 28 57 39 35 36 44 41 31 30 23 23 25 15 13 99 90 . 14 5 3 3 5 3 2 3 2 2 3 3 5 3 2 2 3 2 5 2 5 5 7 7 . 1 . 8 .47 .23 .18 .18 .17 .16 .13 11 19 36 31 36 81 44 19 89 61 28 14 17 78 78 75 67 56 47 47 42 99 99 5 6 2 2 5 2 3 3 3 2 2 2 3 2 2 2 2 2 2 5 2 3 2 2 2 2 2 2 7 5 ...... 2 6 8 6 6 67 40 31 62 56 13 40 79 77 56 48 38 31 31 29 27 23 21 21 17 12 13 99 83

2 2 3 3 3 2 2 3 2 2 3 2 2 2 2 3 3 3 2 2 2 2 3 3 2 2 2 2 2 2 2 2 2 7 5 4 2 4 4 3 3 11 25 18 18 47 34 32 32 25 24 17 27 46 20 55 13 13 10 25 18 15 54 44 31 10 10 99 74 2 2 3 3 2 2 3 2 2 3 2 3 2 2 2 3 3 2 2 2 2 5 3 2 2 2 2 2 2 2 2 2 7 5 .12 9 2 4 5 1 4 8 8 5 4 3 22 80 53 51 49 47 46 33 46 34 22 47 17 18 28 27 22 64 55 34 10 99 90 3 2 3 3 2 2 2 2 1 2 2 2 3 2 3 2 3 2 2 2 2 2 2 2 2 2 2 2 3 1 2 2 6 5 . 1 9 31 42 34 24 12 14 19 63 20 28 10 98 57

. 3 2 1 2 2 2 2 1 2 2 3 2 1 7 3 . 1 . 9 .43 . 1 . 2 .58 . 2 .16 . 2 . . 6 .16 . 3 . 8 . .32 . 1 . 2 . . . .12 .22 . 9 . 5 5 5 99 24 24 81 19 99 99 19 29 29 24 ) 0 ) 0 (E ) ) ) ) 0 0 0 ) ) 0 (E ) 0 0 0 ) (E (E 0 (E (E (E (E ) (E 0 (E ) (E 0 (E rma fi rmae fi Squamarina gypsacea Campylium stellatum Hypnum bambergeri alpina Viola Racomitrium lanuginosum Entodon concinnus Hymenostylium recurvirostrum Didymodon giganteus Saxifraga aizoides Huperzia selago Parnassia palustris Dryas octopetala Leontopodium alpinum Aster alpinus Swertia perennis *hungarica Campanula cochleariifolia Sesleria albicans Asplenium viride Scabiosa lucida Festuca supina Orthothecium rufescens myrtillus Vaccinium Soldanella carpatica Mnium thomsonii Carex Oxytropis carpatica Oxytropis Salix reticulata Selaginella selaginoides Salix retusa Cololejeunea calcarea Salix alpina Biscutella laevigata carpatica Pyrola Dicranum scoparium Pogonatum urnigerum pc Sc, pc cy pc Sc, pc pc Sc, pc cy ES JT Caricion pc st, oe ac ES ac ES, Tr cf, ac 186 POLISH BOTANICAL JOURNAL 49(2). 2004

3 3 2 2 2 2 2 2 5 2 2 1 3 2 2 ...... 8 3 8 3 53 18 35 13 10 93 55 90 28 18 40 3 2 2 3 2 2 2 3 2 2 2 2 2 8 5 2 3 2 2 5 3 2 3 2 3 2 2 2 38 27 27 64 33 2 2 3 2 2 .33 .58 .67 .31 .24 . 2 . 9 55 36 55 18 2 4 2 1 3 2 2 2 2 2 2 2 . . 9 . . .11 2 . . .11 ...... 29 4 . 2 . . .11 . .31 . .29 . .60 . .31 . . .16 9 . 4 . . .20 . .22 5 3 5 3 5 79 99 68 21 61 95 74 ...... 2 1 4 3 4 2 3 3 2 . 3 3 4 2 3 2 3 3 2 3 ...... 11 11 11 89 33 56 22 56 33 78 3 3 3 3 2 2 2 2 2 3 3 2 2 3 2 2 2 3 1 2 2 1 7 5 5 6 99 68 86 61 53 61 53 19 10 20 3 3 3 3 2 2 3 2 2 3 2 2 2 3 1 2 . 1 . 5 . 2 . 1 97 94 97 39 56 64 56 28 22 39 3 2 2 3 2 2 2 3 2 2 2 3 2 2 . 6 . 3 ...... 17 .11 .11 .14 ...... 2 2 8 99 50 79 77 52 60 52 13 2 2 2 2 2 2 3 2 3 1 2 2 2 2 2 2 2 2 2 2 1 2 2 1 3 8 1 2 1 74 72 37 67 34 62 31 35 10 14 16 2 2 2 3 3 3 2 2 2 2 2 2 2 2 2 2 2 . 3 . 4 . 1 . 1 . 1 . 6 9 3 2 11 93 78 51 65 43 48 25 47 12 14 2 2 2 3 2 2 2 3 1 2 2 2 2 2 2 2 2 2 1 2 ...... 3 2 66 23 69 23 76 21 2 2 3 2 2 3 2 2 2 1 2 3 1 . 6 .18 . 1 . 2 . . 4 . . 2 . . . . .37 .22 .11 .12 . . 1 . . 1 . 8 2 . . . 8 ...... 2 . . 2 ?53 5 5 29 29 29 10 81 95 38 57 29 48 90 orum fl . Sesleria sphaerocephala Dianthus *praecox Arenaria ciliata Arenaria Gentiana terglouensis Phyteuma sieberi Leontodon incanus Allium *montanum Pulsatilla slavica Acinos alpinus Allium ochroleucum Galium anisophyllon Phyteuma orbiculare Helianthemum grandi Ranunculus alpestris Bistorta vivipara Saxifraga caesia Bellidiastrum michelii Sesleria tatrae Thymus pulcherrimus aphylla Veronica Polygala *brachyptera Thesium alpinum Festuca tatrae Bartsia alpina Pedicularis rosea Hieracium villosum Rhodax rupifragus Gentianella fatrae fruticans Veronica uva-ursi Arctostaphyllos discolor Saussurea Gentiana verna Anthyllis *alpestris Potentilla crantzii Linum extraaxillare Continued . Seslerio-Asterion alpini Pc Communityac, cy fv pc B1 A B2 B C1 C2 CSeslerietalia coerulae, Elyno-Seslerietea DSc, cy Dsst ESc FSc, pc GSc H fv Sc Sc, fv st Table 1 Table J. ŠIBÍK ET AL.: SYNTAXONOMICAL REVISION OF PLANT COMMUNITIES 187

1 1 3 5 2 . . . . 5 3 13 23 (cont.) 1 2 2 3 2 7 5 3 . . 62 47 3 5 2 2 . . . 2 ...... 9 9 99 3 2 3 1 2 3 3 2 2 2 5 2 5 2 3 3 3 2 2 2 . . 2 . . 4 . 7 . . 4 . 8 ...... 27 . . . 2 . 5 8 8 5 5 8 32 24 24 87 82 29 26 32 24 97 18 47 66 13 ...... 4 3 6 2 3 . . 11 99 89 6 2 5 2 3 3 94 61 5 2 3 . 1 . 1 92 81 6 2 3 2 . . . .11 ...... 8 ...... 2 2 96 48 2 2 3 2 2 1 2 2 2 2 6 2 5 2 2 2 5 2 2 2 2 2 2 2 2 2 3 1 1 4 1 8 4 3 3 8 8 3 2 1 1 14 10 99 34 14 13 2 2 2 2 2 2 5 2 2 2 5 2 2 2 2 2 2 2 2 2 . 1 . 3 . 1 . 2 . 1 . 5 . 1 1 1 4 2 1 1 2 8 4 1 10 10 99 25 22 12 15 2 2 5 2 1 1 2 2 2 6 2 3 2 2 2 5 2 3 3 2 2 2 2 3 ...... 7 4 7 2 11 2 1 2 2 3 5 2 2 2 2 2 1 2 . . . . . 2 . 7 .19 .43 . 1 . 1 1 . .13 . 4 .11 .99 . 8 . 3 . 1 . 4 .10 . 3 1 . 2 . . 1 . 3 . 1 5 5 10 67 10 24 24 90 38 24 10 48 43 Astragalus australis Astragalus alpinus campestris Oxytropis rupestris Carex Gymnadenia conopsea capillaris Carex simpliciuscula Kobresia Pedicularis verticillata Astragalus frigidus *tatrae Oxytropis Ranunculus thora Astragalus norvegicus Bupleurum ranunculoides Globularia cordifolia ericetorum Carex Festuca pumila mucronata Carex Achillea clavennae Pedicularis rostrato-capitata halleri Oxytropis Comastoma tenellum Festuca versicolor Hedysarum hedysaroides Saxifraga oppositifolia Ligusticum mutellinoides Antennaria carpatica Elyna myosuroides Gentiana nivalis fuliginosa Carex alpina Saussurea Luzula *mutabilis Dianthus glacialis alpina Agrostis Tephroseris capitata Tephroseris glaucinus Carduus globosa Traunsteinera st st Sc Carici rupestris-Kobresietea st sa st, cf st st Sc Sc Sc Sc Sc Sc Sc fv Festucion versicoloris cf oe, an oe oe st 188 POLISH BOTANICAL JOURNAL 49(2). 2004

2 2 2 . . . . . 8 10 3 3 2 3 2 1 2 . 1 .24 . . . .24 . . . .20 . . . 9 ...... 13 . . . 2 . 7 . . 3 2 2 2 2 3 5 3 2 2 3 3 3 2 3 2 2 3 2 2 2 3 3 2 2 2 3 2 2 2 . 9 . . 2 . 5 3 8 5 3 3 3 3 3 3 3 3 11 47 61 61 55 53 21 26 45 71 39 82 89 29 89 68 26 29 ...... 1 5 5 3 5 2 2 2 . . . . 11 11 67 33 22 56 22 3 2 3 3 3 2 2 2 2 3 2 3 6 7 2 61 20 39 22 15 5 2 3 3 3 2 2 2 3 . 1 . 2 6 89 25 36 31 19 2 2 2 3 2 2 2 2 ...... 8 .14 ...... 6 . . 8 2 42 17 40 15 12 2 3 2 2 2 2 2 2 2 2 2 2 2 3 2 2 2 2 2 2 2 2 2 2 2 1 1 2 1 9 3 5 1 6 2 8 3 9 1 1 9 11 55 15 13 10 20 14 10 2 2 2 2 3 2 2 2 2 2 2 2 3 3 2 2 2 2 2 2 2 2 2 . 1 . 3 . 1 1 2 1 9 3 9 2 11 17 72 13 10 14 22 12 38 14 22 16 1 3 3 2 2 2 3 2 2 2 2 3 2 2 2 2 2 2 2 2 2 1 2 7 39 10 1 2 1 2 . 1 7 . . 3 . 4 . 1 1 ...... 1 . 7 ...... 1 . . 1 .16 . 3 . 4 1 ...... 4 . 4 . 3 . 1 . 1 . 1 . . 4 . 7 . 7 5 71 19 48 di ) fi 0 ) 0 ) ) ) 0 0 ) 0 (E 0 (E ) 0 (E (E (E (E (E dus fi . Cetraria islandica Empetrum hermaphroditum Sanionia uncinata Myosotis alpestris Vaccinium vitis-idaea Vaccinium Euphrasia tatrae Juncus tri Hieracium alpinum Senecio *carpathicus Sedum atratum Saxifraga wahlenbergii Pritzelago alpina halleri Cardaminopsis Salix kitaibeliana Polytrichum alpinum atrata Carex Campanula tatrae Campanula alpina Primula minima Cetraria cucullata Cetraria nivalis Cladonia arbuscula Cladonia gracilis Pulsatilla scherfelii Gentiana frigida versicolor Avenula Leontodon pseudotaraxaci Vaccinium gaultherioides Vaccinium Luzula sudetica disticha Oreochloa alpina Veronica Saxifraga androsacea Continued . CommunityLoiseleurio-Vaccinion st Caricetalia curvulae, Juncetea tri B1 A B2 B C1 C2 Cjt D Dsjt jt Arabidion coeruleae Ept Fpt Gst H ns an Table 1 Table J. ŠIBÍK ET AL.: SYNTAXONOMICAL REVISION OF PLANT COMMUNITIES 189 5 3 2 3 2 2 2 5 5 2 2 2 2 2 . . . . 5 3 93 10 45 53 35 20 90 85 38 58 (cont.) 3 2 2 2 2 2 3 2 2 3 3 3 2 2 3 2 . . . 9 4 62 36 67 49 2 2 2 2 2 3 3 2 2 ...... 18 .60 . . . 9 82 55 91 72 3 2 5 2 5 3 3 2 1 3 3 5 3 2 2 2 . . 7 . .99 2 . 4 . .36 .36 . .18 . . .28 . . . 3 . . .44 . .38 . .13 9 . . 9 3 8 3 8 5 3 3 11 11 21 18 34 13 71 66 61 ...... 2 2 1 4 1 2 2 2 2 5 5 2 2 2 . . . . . 11 11 22 22 22 44 3 2 2 1 3 2 2 5 3 2 2 2 74 68 65 40 17 3 2 5 3 2 3 . 9 . 2 . 2 . 6 . 6 3 3 53 72 67 50 3 2 2 1 3 2 2 5 3 2 2 ...... 33 . .22 . . . . 2 4 4 88 15 10 10 65 63 33 27 3 2 2 2 3 2 2 2 2 5 3 2 2 2 2 1 2 2 2 1 9 5 2 7 7 1 1 1 74 32 10 92 72 45 3 2 2 2 2 2 5 3 2 3 2 1 2 2 2 . 2 . 5 . 2 . 2 1 9 1 1 8 8 65 30 96 66 48 3 2 2 2 3 2 2 2 2 3 3 2 2 2 2 6 11 83 34 18 88 78 42 2 2 1 1 2 1 1 1 ...... 11 ...... 10 . . . 2 . . . . . 4 . . . . . 6 2 . 2 . 1 ...... 4 .12 1 . . 4 5 48 10 76 43 14 14 10 ) 0 ) 0 ) (E 0 ) (E 0 (E (E exicaule fl eldia pusilla fi Saxifraga bryoides To Neckera crispa tripteris Valeriana brachystachys Carex Luzula alpinopilosa stiriacum Doronicum Festuca picturata jacquinii Crepis Draba aizoides Gypsophila repens Artemisia eriantha pyrenaica Petrocallis Cystopteris fragilis Gymnocarpium robertianum tortuosa Tortella Ditrichum Saxifraga paniculata Fissidens cristatus Asplenium ruta-muraria Hieracium bupleuroides Cerastium *glandulosum Arabis alpina Saxifraga hieraciifolia Papaver tatricum Carex atrofusca Carex alpina Arctous Minuartia langii Kernera saxatilis Draba tomentosa Delphinium oxysepalum an Salicetalia herbaceae, Salicetea herbaceae fp, an fp Potentillion caulescentis cf cf sa Cystopteridion Gp Asplenietea trichomanis Potentilletalia caulescentis, AT Pc Pc, Sc Pc Pc Pc Thlaspietea rotundifolii pt aa, cy an pt, ac sa pt 190 POLISH BOTANICAL JOURNAL 49(2). 2004 2 2 2 2 2 2 2 2 1 3 2 2 1 1 3 4 ...... 5 3 5 3 8 5 3 75 15 20 25 73 23 20 23 2 2 2 2 1 2 2 5 2 2 2 2 3 2 1 2 3 2 2 2 2 2 2 2 2 2 2 2 1 2 22 1 2 . 7 . 7 .11 . 4 .36 . 4 . . . 4 .11 . . 7 .11 . . . 4 . 2 . 2 9 27 1 2 2 2 3 2 2 2 2 2 2 2 2 3 2 2 2 1 2 1 . . 2 . . .29 3 . 4 . 2 . 7 . . .44 7 . . .13 9 . . .16 . .11 2 . 4 . . . .22 . 3 5 3 3 8 8 5 5 8 5 3 3 5 11 16 29 16 53 37 16 ...... 1 2 1 1 1 2 5 2 7 3 3 2 2 3 1 2 2 2 2 ...... 11 11 11 11 11 11 11 44 22 33 22 22 2 2 2 2 2 2 2 3 1 1 2 2 2 2 2 2 2 2 2 2 2 2 2 1 2 2 2 5 2 5 6 6 3 2 7 5 2 1 1 5 1 3 2 1 1 1 31 35 24 32 15 14 2 3 1 1 2 2 2 3 2 2 2 1 2 2 2 2 2 2 1 2 2 2 5 . 1 . 2 . 1 . 1 . 1 . 2 6 3 3 3 8 6 33 25 28 36 28 2 2 2 2 2 1 1 2 2 2 3 2 1 2 2 2 2 .11 .14 .14 . .11 . . . 3 . 3 . 8 . 6 . 3 . . 6 2 2 4 2 2 2 29 42 21 29 19 1 1 2 2 2 2 2 2 2 1 2 2 2 2 2 1 2 1 2 1 1 2 2 1 3 1 1 3 3 1 1 2 1 7 1 2 5 1 11 10 33 23 13 1 2 2 2 2 2 2 2 2 2 2 1 2 2 2 1 2 1 2 2 . 1 . 1 . 1 1 1 3 9 5 3 1 1 9 2 15 15 22 14 1 1 2 2 2 2 2 1 2 2 2 2 1 2 1 1 2 . 1 . 3 . 1 1 1 1 . . 6 . 6 .44 . 1 . 2 . . 6 . . . 3 . . 6 . 2 . 2 . . 8 . . . .33 . . . . . 3 .32 . 1 . . .10 . . 2 . 2 ...... 2 . .17 . 1 . 2 . 1 1 . 1 . . 1 1 . . . . .11 . . . . .11 . 4 . 7 . . 3 . 5 5 5 agg. ora fl dum fi rmum fi . Pinus mugo arenosa Cardaminopsis Gentianella lutescens Jovibarba globifera Botrychium lunaria Hieracium bi Luzula sylvatica Luzula luzuloides albida Pseudorchis Clematis alpina Hieracium caesium Thymus alpestris Primula elatior Pinus sylvestris Picea abies Rhodiola rosea Larix decidua Salix silesiaca Pimpinella major Aconitum Lotus corniculatus Sorbus aucuparia Solidago virgaurea Homogyne alpina Anemone narcissi sempervirens Carex Ranunculus pseudomontanus Coeloglossum viride Bistorta major Potentilla aurea ornithopoda Carex Ligusticum mutellina Saxifraga ×patens Juncus triglumis Campanula serrata Achillea *alpestris Saxifraga mutata Continued . CommunityOther taxa tf B1 A B2 B C1 C2 C D Ds E F G H Table 1 Table J. ŠIBÍK ET AL.: SYNTAXONOMICAL REVISION OF PLANT COMMUNITIES 191

3 2 1 2 2 5 2 2 3 3 2 2 5 3 2 2 2 2 2 2 2 . . . . . 3 8 8 5 3 3 30 20 23 53 20 58 88 18 30 23 10 (cont.) 3 2 5 3 2 3 2 3 3 3 2 2 2 2 2 2 2 2 3 3 2 2 . . 4 2 2 2 9 27 31 3 2 2 2 2 2 2 2 2 . 9 .13 .33 .22 . 7 .13 . 2 . 4 .29 .11 . 7 . 9 9 73 18 45 27 2 6 3 3 3 2 3 3 2 2 2 3 2 2 2 2 2 2 . .11 . . 4 . 7 . . .18 . .15 . .10 . .25 ...... 4 . .45 .55 .36 3 3 5 3 5 3 8 3 89 92 45 50 13 21 53 24 24 47 ...... 5 5 3 5 5 ...... 11 11 11 56 22 3 2 5 3 3 2 3 3 2 3 2 2 2 2 3 3 2 3 2 2 3 2 2 7 7 8 2 3 11 66 31 28 19 52 3 3 5 3 3 2 5 3 5 3 3 3 . 8 . 1 . 6 . 6 . 3 . 2 . 2 . 3 . 1 . 3 3 6 8 3 6 11 11 89 36 22 56 19 2 2 3 2 3 2 3 2 2 3 2 2 2 2 2 2 2 3 2 2 3 2 2 8 4 6 2 6 6 2 4 4 6 2 2 6 50 27 33 27 50 13 10 10 2 2 3 2 2 2 2 2 2 2 2 2 2 1 2 2 3 3 2 2 2 2 8 8 7 5 2 5 7 2 6 11 19 19 16 40 37 43 10 27 32 14 20 4 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 3 2 2 3 2 2 3 2 5 1 9 1 3 4 13 12 33 27 40 50 57 12 18 29 25 14 20 2 2 5 3 2 2 2 2 2 2 2 2 1 2 2 2 3 2 2 1 2 ...... 8 2 4 14 2 2 2 1 2 2 1 1 1 2 1 1 1 ...... 26 . 4 . 6 .40 .23 .30 .11 .18 . 3 . 4 . 2 . 2 .21 . 1 . 8 . . . . .13 .24 .40 5 5 5 10 14 14 10 10 10 10 24 19 38 ) 0 polytrichus * Valeriana saxatilis Valeriana Gentianella campestris Soldanella minima Potentilla nitida Polygala alpestris Soldanella alpina Thymus Daphne arbuscula Cladonia pyxidata Calamagrostis varia Calamagrostis Melampyrum sylvaticum Arabis pumila Campanula scheuchzeri Genista pilosa Galium austriacum humilis Carex Schistidium apocarpum Hylocomium splendens Rhytidiadelphus triquetrus Rhytidium rugosum Distichium capillaceum Ctenidium molluscum Campylium chrysophyllum Bryum elegans Hypnum vaucheri Fulgensia bracteata Plagiochila porelloides Distichium inclinatum Thuidium philibertii schreberi Pleurozium Megaspora verrucosa Ptilidium ciliare Encalypta streptocarpa Acer pseudoplatanus Homalothecium philippeanum Encalypta alpina Ctenidium procerrimum julacea Myurella Bryophytes & Lichens (E 192 POLISH BOTANICAL JOURNAL 49(2). 2004 , , Pc 2 2 2 3 5 3 2 2 2 2 2 . . 3 15 30 58 35 28 Trisetion Trisetion rded in the 1980 (Msc.), , tf – 2 2 2 2 2 2 5 2 . 3 . . 2 Elyno-Seslerietea et al. Oxytropido-Elynion Dryadetum octopetalae , ES – 2 2 2 3 . 2 . . . 2 . . 2 . .15 8 . . .15 . .13 . . . . , oe – Seslerion tatrae , st – 2 2 3 3 2 2 2 2 2 2 2 3 2 5 .18 .82 4 . .18 .27 . 2 . . 4 ); 7 – Dúbravcová 5 Cystopteridion 26 21 21 16 21 26 37 21 50 47 24 47 34 Nardion strictae Nardion , cy – ...... owski 1935, Table 2, r. 3, 7, Belianske Tatry Mts Tatry 3, 7, Belianske 2, r. Table owski 1935, ł , ns – rmae fi rmae tatricum di fi fi Seslerietalia coerulae 2 ...... ); 67 – Petrík ined., Belianske Tatry Mts (65), Vysoké Tatry Tatry Vysoké Mts (65), Tatry ); 67 – Petrík ined., Belianske Caricion , Sc – ); 2 – Paw Caricetum Juncion tri rmae fi , cf – 2 2 2 2 5 2 2 2 2 2 2 3 , jt – di fi 2 . 1 . 9 .16 . 5 . 1 . 2 . 3 . 5 . 6 . 3 Caricetum 3 rmae carpaticum fi 2 2 2 2 5 2 2 2 2 2 2 Juncetea tri Seslerio-Asterion alpini ...... 11 ...... 4 . . 4 2 8 2 4 6 8 6 15 27 10 1969: 63–64, r. 136, 214, 218, 231, 240, Belianske Tatry Mts ( Tatry 136, 214, 218, 231, 240, Belianske 1969: 63–64, r. , sa – , JT – , JT Asplenietea trichomanis Caricetum et al.

č 2 2 2 2 2 3 2 2 2 1 2 3 2 2 2 2 2 2 2 2 . 1993: 407) 4 5 5 1 2 5 6 2 8 6 1 1 1 1 1 18 14 15 , AT – , AT cinalis fi et al 2 2 2 2 2 2 2 2 2 2 3 2 2 2 2 2 2 2 . 1 . 1 1 9 1 2 8 ); 5 – Hada 11 18 16 10 25 Papaverion tatrici rmae fi 2 2 2 2 2 3 2 2 2 1 2 2 , pt – 7 1 Galio-Parietalia of Androsacion alpinae Androsacion : 1 – Braun-Blanquet 1930: 25, Belianske Tatry Mts ( Tatry : 1 – Braun-Blanquet 1930: 25, Belianske 1971, Table 2, r. 1, 3, Belianske Tatry Mts ( Tatry 1, 3, Belianske 2, r. Table 1971, 2 1 .12 . 1 . 1 . 1 . 6 . .15 . .12 1 . 2 . 2 ...... 17 1 . . 1 . 1 3 . 1 . . 2 . . . . . 2 , Gp – 5 80 82 124 35 23 55 3 0 57 4 32 37 , an – 12 6 8 13 2 7 8 8 0 6 0 40 31 19 14 et al. Dryadeto-Caricetum rmae typicum fi , (cy) – taxon with lower relation to the relevant syntaxon and coenological optimum also in other syntaxon; ? taxa reco Potentillion caulescentis 1 0 Festucion pictae ); 2 – Šmarda Arabidion coerulae Rübel 1911 by other authors (cf. Grabherr Rübel 1911 , pc – 1969: 58–59, r. 24, 153, 154, Belianske Tatry Mts ( Tatry 24, 153, 154, Belianske 1969: 58–59, r. , fp – : 21 – Braun-Blanquet & Jenny 1926, Table 7, r. 1–21, Alps; 1–21, 7, r. Table : 21 – Braun-Blanquet & Jenny 1926, ined., Belianske Tatry Mts; Tatry ined., Belianske rmae , ac – č fi et al.

č rmae fi . rmae carpaticum fi Caricetum Thlaspietalia rotundifolii Lecanora epibryon Campylium halleri Barbula crocea Racomitrium canescens Cirriphyllum cirrosum Meesia uliginosa trichophyllum Blepharostoma quinquedentata Tritomaria austriaca Timmia Scapania cuspiduligera Isopterygiopsis pulchella Solorina bispora Psora decipiens Lophozia bantriensis Dicranum spadiceum Dicranum fuscescens Plagiopus oederiana Philonotis tomentella Xanthoria elegans Polytrichum formosum Solorina saccata Pohlia cruda Continued Arenario tenellae-Caricetum Arenario ); 3 – Hada Caricetum . Arabidion alpinae , Tr – , Tr Festucion versicoloris A: B1: Potentilletalia caulescentis XPLANATIONS OURCES Community B1 A B2 B C1 C2 CNumber of the accesoric taxa E D Ds E F G H Number of the accesoric taxa E Caricetum S association fusci – Mts (2); 3 – Valachovi Mts (2); 3 – Table 1 Table Table 1, r. 12–18, Západné Tatry Mts ( Tatry 12–18, Západné 1, r. Table ( fv – aa – E tatricum J. ŠIBÍK ET AL.: SYNTAXONOMICAL REVISION OF PLANT COMMUNITIES 193 ), ); ské č 1923, et al. ined., Nízke 1969: 58–59, č ); 1 – Bernátová et al.

č Dryadeto-Firmetum ); 5 – Szafer ské vrchy Mts (4), Krivánska č ské vrchy Mts; ); 2 – Hada č ); 1 – Bernátová ined., Krivánska Sesleria varia-Dryas octopetala Firmetum Dryadeto-Firmetum rmae fi & Jarolímek 1994, Table 1, r. 29–31, 1, r. Table & Jarolímek 1994, č ské vrchy Mts; 5 – Valachovi ské vrchy Mts; 5 – č 1971, Table 2, r. 2, 4 – 6, Belianske Tatry Mts Tatry 2, 4 – 6, Belianske 2, r. Table 1971, Dryado-Firmetum et al. ); 3 – Valachovi ); 3 – ), Table 2, r. 1–3 (comm. 2, r. Table ), Dryadeto-Caricetum ); 48 – Petrík ined., Belianske Tatry Mts (29), Vysoké Tatry Tatry Vysoké Mts (29), Tatry ); 48 – Petrík ined., Belianske ká Fatra Mts (15); 16 – Petrík ined., Muránska planina (9), rmae fi ľ ); 4 – Šmarda Dryadetum ské vrchy Mts; 1 – Mucina ined., Cho č ká Fatra Mts (12); 13 – Petrík ined., Cho ľ Caricetum rmae carpaticum fi rmae carpaticum fi ); 5 – Sillinger 1933: 223, r. 1–5, Nízke Tatry Mts ( Tatry 1–5, Nízke ); 5 – Sillinger 1933: 223, r. Caricetum 1990, Table 2, r. 1–10, Belianske Tatry Mts; Šmarda 1971, Table 1, r. 103, Belianske Tatry Tatry 103, Belianske 1, r. Table Mts; Šmarda 1971, Tatry 1–10, Belianske 2, r. Table 1990, lohlávková & Fišerová 1976, Table 1, r. 1, 5, Krivánska Malá Fatra Mts ( 1, r. Table lohlávková & Fišerová 1976, č anská Malá Fatra Mts (1), Ve anská Malá Fatra Mts (1), ě owski & Stecki 1927, Table 2, r. 7–9, 13–17, West Tatras, Table 2., r. 21, 22, Belianske Tatry Tatry 21, 22, Belianske 2., r. Table Tatras, West 7–9, 13–17, 2, r. Table owski & Stecki 1927, č ł Caricetum ká Fatra Mts (2), Západné Tatry Mts (1); 1 – Šibík 2003, Table 3, r. 34, Krivánska Malá Fatra Mts; 3, r. Table Mts (1); 1 – Šibík 2003, Tatry ká Fatra Mts (2), Západné ské vrchy Mts (1); 22 – Petrík ined., Krivánska Malá Fatra Mts (10), Nízke Tatry Mts (9), Cho Tatry ské vrchy Mts (1); 22 – Petrík ined., Krivánska Malá Fatra (10), Nízke ľ č : 2 – B rmae carpaticum 1923, Table 12, r. 1–2, West Tatras ( Tatras West 1–2, 12, r. Table 1923, fi owski & Stecki 1927, Table 2, r. 5, 6, 10, 11, 18–20, 23, West Tatras ( Tatras West 18–20, 23, 5, 6, 10, 11, 2, r. Table owski & Stecki 1927, ł ); 10 – Paw et al. 1980 (Msc.), Table 1, r. 1–11, Západné Tatry Mts ( Tatry Západné 1–11, 1, r. Table 1980 (Msc.), ské vrchy Mts; 1 – Bernátová ined., Cho č lohlávková 1980, Table 8, r. 12–15, Krivánska Malá Fatra Mts; 2 – Milová & Urbanová 1989: 295, 8, r. Table lohlávková 1980, ě et al. : 8 – Paw Caricetum : 4 – B : 4 – Sillinger 1933: 229–230, Nízke Tatry Mts; Tatry : 4 – Sillinger 1933: 229–230, Nízke : 10 – Marhold & Valachovi : 10 – Marhold & : 1 – Bernátová ined., Belianske Tatry Mts; 37 – Petrík ined., Belianske Tatry Mts; Tatry Mts; 37 – Petrík ined., Belianske Tatry : 1 – Bernátová ined., Belianske ined., Belianske Tatry Mts; Tatry ined., Belianske ); 13 – Bernátová ined., Západné Tatry Mts (1), Ve Mts (1), Tatry ); 13 – Bernátová ined., Západné ); 2 – Szafer č rmae carpaticum : 7 – Šimeková & Pitoniak 1976, Table 1, r. 1–4 ( 1, r. Table : 7 – Šimeková & Pitoniak 1976, fi ); lohlávková & Fišerová 1976, Table 1, r. 2–4, 6–10, Krivánska Malá Fatra Mts ( 1, r. Table lohlávková & Fišerová 1976, ě rma fi rmae fi rmae primuletosum auriculae fi ské vrchy Mts ( č : 8 – B : 11 – Dúbravcová : 11 Carex Carex 1984, Table 15, r. 1–10, Západné Tatry Mts ( Tatry 1–10, Západné 15, r. Table 1984, Caricetum et al. Caricetum ká Fatra Mts (1). owski 1935, Table 2, r. 1, 2, 4, 6, 8, West Tatras ( Tatras West 1, 2, 4, 6, 8, 2, r. Table owski 1935, ľ ł ské vrchy Mts (1); č ); 10 – Unar ); 5 – Paw Festuca versicolor-Androsace lactea Festuca versicolor-Androsace rmae fi ské vrchy Mts; 2 – Mucina ined., Nízke Tatry Mts (1), Cho Tatry ské vrchy Mts; 2 – Mucina ined., Nízke č Artemisia petrosa-Draba tomentosa Artemisia petrosa-Draba owski 1935, Table 2, r. 5, Cho 2, r. Table owski 1935, A.-C. salicetosum reticulatae Dryado octopetalae-Caricetum D.-C. saxifragetosum aizoidis ł Potentillion caulescentis/Seslerio-Asterion alpini Cystopteridion/Seslerio-Asterion alpini Androsaceo lacteae-Festucetum versicoloris Androsaceo Drabo tomentosae-Artemisietum petrosae Dryado octopetalae-Salicetum reticulatae Firmetum C1: B2: C2: D: F: Ds: ass. E: G: H: Caricetum Mts (1), Západné Tatry Mts (18); 2 – Valachovi Mts (18); 2 – Tatry Mts (1), Západné 1 – Paw ( ined., Cho r. 149, 150, Belianske Tatry Mts ( Tatry 149, 150, Belianske r. Mts ( vrchy Mts (3); 6 – Šibík 2003, Table 3, r. 28–33, Krivánska Malá Fatra Mts; 8 – Švandová-Ursíniová 1966, Table 1, r. 1–8, Cho 1, r. Table 28–33, Krivánska Malá Fatra Mts; 8 – Švandová-Ursíniová 1966, 3, r. Table vrchy Mts (3); 6 – Šibík 2003, Mts (4), Cho Tatry Malá Fatra Mts; 27 – Šibík 2003, Table 3, r. 1–27, Krivánska Malá Fatra Mts; 3, r. Table Malá Fatra Mts; 27 – Šibík 2003, Fatra Mts; 1 – Švandová-Ursíniová 1966, Table 1, r. 9, Cho 1, r. Table Fatra Mts; 1 – Švandová-Ursíniová 1966, Mts (ass. Muránska planina Mts (comm. with Table 13, r. 1–5, West Tatras ( Tatras West 1–5, 13, r. Table Slovenský raj Mts; 17 – Bernátová ined., Krivánska Malá Fatra Mts (1), Lú Slovenský raj Mts (6), Ve Slovenský raj Mts (6), Malá Fatra Mts (1), Muránska planina Mts (2), Nízke Tatry Mts (3), Ve Mts (3), Tatry Malá Fatra Mts (1), Muránska planina (2), Nízke 194 POLISH BOTANICAL JOURNAL 49(2). 2004

The association Arenario tenellae-Caricetum two variants can be distinguished: the typical one, fi rmae occurs on limestone and dolomite bedrock, and a variant with Kobresia simpliciuscula. The on moderate to steep slopes with a rather thick typical variant comprises communities on stabi- layer of detritus, but also on steep rocky slopes. lized dolomite scree slopes. The communities on The occurrence of species of individual higher rocky sites are positively differentiated by Gen- syntaxa indicates that the association shows tiana clusii, Bellidiastrum michelii and Anthyllis close syngenetic relationships to communities of vulneraria subsp. alpestris. The variant with the alliance Potentillion caulescentis (in the alpine Kobresia simpliciuscula is concentrated on lime- belt) and Seslerio-Asterion alpini (in the subalpine stone of high-mountain ridges. Unlike the typical belt). On sites with a deeper soil profi le which variant, it is differentiated by Oxytropis carpatica diminishes the infl uence of basic bedrock, the and Dactylina madreporiformis. Based on the con- occurrence of species of the alliances Arabidion stant presence of those differential taxa and overall coerulae, Festucion versicoloris and Juncion trifi di fl oristic similarities (except for a higher propor- increases, depending on moisture and slope ori- tion of Lloydia serotina and Carex fuliginosa), entation. This phenomenon evidences the process communities with Carex rupestris codominant to of succession, which eventually may lead to the dominant can also be assigned to this variant. formation of those communities. Arenario tenellae-Caricetum fi rmae salicetosum Arenario tenellae-Caricetum fi rmae typicum reticulatae Petrík subass. nov. hoc loco Petrík subass. nov. hoc loco Table 1, column B2 Table 1, column B1 DIFFERENTIAL TAXA: Salix reticulata, S. retusa, Sela- DIFFERENTIAL SPECIES: Aster alpinus, Euphrasia salis- ginella selaginoides, Soldanella carpatica, Swertia burgensis, Leontopodium alpinum, Oxytropis carpatica, perennis subsp. alpestris, Viola alpina, Campylium stel- Ranunculus breyninus, Trisetum alpestre, Squamarina latum, Hypnum bambergeri, Orthothecium rufescens. gypsacea. NOMENCLATURAL TYPE: Petrík ined., holotypus NOMENCLATURAL TYPE: identical with the type of the LOCALITY: Belianske Tatry Mts, Zadné Jatky Mt., association name. northern slope, alt. 1920 m, aspect: N, slope: 35°, 2 relevé area: 20 m , cover total: 90%, E1: 80%, E0: Plant communities of this subassociation are 30%, 3. 8. 1995 open, show average abundance of about 55–70%, and inhabit various sites on limestone and dolomite E1: Carex fi rma 3, Dryas octopetala 3, Silene in the alpine and sometimes the subalpine belt of acaulis 2b, Saxifraga paniculata 2a, Bistorta vivipara the Tatra Mts. Usually they occur on moderate 1, Festuca versicolor 1, Minuartia gerardii 1, Pedicu- laris oederi 1, Salix reticulata 1, Saxifraga aizoides to steep, mostly north-facing slopes with geests 1, Swertia *alpestris 1, Androsace chamaejasme +, of various depths, but often can be found also Bartsia alpina +, Campanula alpina +, Carex fuligi- on steep to abrupt rock walls situated on peaks nosa +, Cerastium eriophorum +, Crepis jacquinii +, and edges of various orientations. The layer of Dichodon cerastoides +, Draba aizoides +, Galium detritus is 15–30 cm (sometimes even 40–50) cm anisophyllon +, Hedysarum hedysaroides +, Huperzia deep, and contains only a small amount of fi ne selago +, Ligusticum mutellinoides +, Lloydia serotina earth, retaining only a small amount of water. +, Minuartia sedoides +, Myosotis alpestris +, Phyteuma These communities therefore appear to be more orbiculare +, Pinguicula alpina +, Pyrola carpatica +, xerophilous. Ranunculus alpestris +, Rhodax alpestris +, Salix retusa +, Saxifraga wahlenbergii +, Selaginella selaginoides Plant communities of the typical subassocia- +, Soldanella carpatica +, Vaccinium vitis-idaea +, tion occur mainly in the Belianske Tatry Mts. In Chamorchis alpina r. the Západné Tatry Mts, these communities usually E0: Dicranum spadiceum 2a, Ditrichum fl exicaule occur on rather steep and dry rocky sites. 2a, Hylocomium splendens 2a, Tortella tortuosa 2a, Based on differences in ecological conditions, Campylium stellatum +, Cirriphyllum cirrosum +, Cte- J. ŠIBÍK ET AL.: SYNTAXONOMICAL REVISION OF PLANT COMMUNITIES 195 nidium molluscum +, Distichium capillaceum +, Enca- indicate that they are transitional towards the asso- lypta alpina +, Hypnum bambergeri +, Orthothecium ciation Dryado octopetalae-Caricetum fi rmae, and rufescens +, Pleurozium schreberi +, Polytrichum sp. thus are interesting phytogeographically. Kliment +, Rhytidium rugosum +, Schistidium apocarpum +, et al. (2004) observed a similar phenomenon sup- Cetraria islandica +, Cladonia *pocillum +. porting the intermediate position of vegetation on Within the association Arenario tenellae- Sivý vrch Mt. between the Chočské vrchy Mts and Caricetum fi rmae, stands of A.-C. salicetosum limestone parts of the Západné Tatry Mts (cf. Futák reticulatae are slightly more hygrophilous. They 1976: 109–110), in the associations Diantho nitidi- occur on moderate to steep, mostly north-facing, Caricetum tatrorum (Sillinger 1933) Kliment et al. wind-exposed slopes in the subalpine and alpine 2004, and Trifolio kotulae-Caricetum tatrorum Kli- belts. They occupy rather deep soils, usually with ment et al. 2004. well-developed but often discontinuous layers of black fi ne humic earth 5–15 cm thick. The soils Dryado octopetalae-Caricetum fi rmae Sillinger exhibit high retention of water; this determines 1933 the occurrence of hygrophilous and humicolous Table 1, column C species, which are the differential species of the association. In comparison with the typical ORIGINAL FORM OF THE NAME: Dryadeto-Firmetum Sillinger 1933 subassociation, these communities have a higher proportion and abundance of mosses. SYNONYMS: Caricetum fi rmae Klika 1932 p.p. maj. Based on differences in ecological conditions, (Art. 31), Firmetum (= Caricetum fi rmae) carpaticum Pawłowski 1935 p.p. min. (Art. 34), Saxifrago cae- three variants can be distinguished within this su- siae-Caricetum fi rmae (Szafer et al. 1923) Hadač in bassociation: with Cerastium eriophorum, with Mucina & Maglocký 1985 p.p. min. (Art. 2b), Festuco Viola alpina, and with Dianthus nitidus. versicoloris-Dryadetum (Szafer et al. 1923) Hadač in Plant communities of the variant with Ceras- Mucina & Maglocký 1985 p.p. maj. (Art. 2b), Caricetum tium eriophorum occur mostly in the alpine belt fi rmae chočense Klika 1932 (nom. nud.). – NON: Cari- of the Belianske Tatry Mts. They are positively cetum fi rmae Szafer et al. 1923, Dryadetum Szafer et differentiated by the species Kobresia simplicius- al. 1923, Firmetum Klika 1926, Firmetum Pawłowski cula, Comastoma tenella, Primula minima, and by & Stecki 1927, Dryadetum octopetalae tatricum Hadač higher constancy of Arenaria tenella, Minuartia et al. 1969. gerardii, Rhodax alpestris and some other taxa DIAGNOSTIC TAXA: Carex fi rma (dom., const.), Dryas 1 as well. octopetala (dom., const.), Bartsia alpina (dif. ), Dian- The variant with Viola alpina comprises com- thus nitidus (dif.), Pinguicula alpina (dif.), Selaginella selaginoides (dif.1), Soldanella carpatica (dif.1), Tofi eldia munities occurring in the subalpine to alpine belts calyculata (dif.), Vaccinium vitis-idaea (dif.1), Hyloco- of the Červené vrchy Mts in the Západné Tatry mium splendens (dif.1), Hypnum cupressiforme (dif.), Mts. It is positively differentiated by Viola alpina Bistorta vivipara (const.), Crepis jacquinii (const.), Fes- and characterized by higher constancy of Belli- tuca versicolor (const.), Galium anisophyllon (const.), diastrum michelii and Pedicularis verticillata. In Pedicularis verticillata (const.), Ranunculus alpestris contrast to the previous variant, the variant with (const.), Saxifraga caesia (const.), Ditrichum fl exicaule Viola alpina is positively differentiated also by (const.), Tortella tortuosa (const.). Bartsia alpina, Sesleria tatrae and Selaginella 1 Differential taxa against the association Arenario selaginoides. tenellae-Caricetum fi rmae. Plant communities treated within the variant NOMENCLATURAL TYPE: Sillinger 1933: 223, rel. 2, with Dianthus nitidus occur on Sivý vrch Mt. in lectotypus hoc loco. the Západné Tatry Mts. These communities occupy a marginal position in the association Arenario This is a two-layered, moderately species- tenellae-Caricetum fi rmae. The occurrence of Di- rich plant community (average 34 taxa per anthus nitidus and the overall fl oristic composition relevé) with dominance of hemicryptophytes and 196 POLISH BOTANICAL JOURNAL 49(2). 2004 chamaephytes. Generally speaking, the cushion- frequent occurrence of species of the class Asple- forming dwarf shrub Dryas octopetala and tufted nietea trichomanis (Table 1) indicates its close sedge Carex fi rma are prominent, giving Dryado relationship to rock fi ssure communities. When octopetalae-Caricetum fi rmae its distinctive stamp. it inhabits windy slopes subject to erosion, with Differences in the abundance of these two species rather deep skeletal soils developed, succession do not basically change the fl oristic composition processes may lead to the formation of the as- of the phytocoenoses. The height of the stands sociation Seslerio tatrae-Festucetum versicoloris. ranges from 5 to 15 cm (tufts, leaves) and from 15 On lee and often rather concave sites with thick to 30 cm (stems, stalks), with Festuca versicolor soils developed, species of the genus Sesleria being a subdominant species. Numerous conspicu- (S. albicans, S. tatrae) and Carex sempervirens ously fl owering species (Bartsia alpina, Bistorta subsp. tatrorum are common; the community is vivipara, Crepis jacquinii, Dianthus nitidus, gradually replaced by phytocoenoses of the as- Galium anisophyllon, Pedicularis verticillata, sociation Diantho nitidi-Caricetum tatrorum. On Pinquicula alpina, Ranunculus alpestris, Saxi- less exposed sites with typically humid climate fraga caesia, Selaginella selaginoides, Soldanella enforced by a north orientation and a thick layer carpatica, Tofi eldia calyculata) and dwarf shrubs of accumulated litter, the association Dryado-Cari- (Vaccinium vitis-idaea) are constant elements of cetum fi rmae is transformed into shrub communi- these stands. The layer of mosses and lichens is ties of the alliance Loiseleurio-Vaccinion. well-developed; the most common species are Two subassociations are distinguished here, Tortella tortuosa, Ditrichum fl exicaule and Hylo- based on differences in the ecology and fl oristic comium splendens. composition of the stands: The association Dryado-Caricetum fi rmae is a chionophobic and sub-hygrophilous plant Dryado octopetalae-Caricetum fi rmae primule- community inhabiting shallow soils developed tosum auriculae subass. nov. hoc loco on calcareous and dolomite bedrock (lithosol) in the supramontane and subalpine belts (alt. Table 1, column C1 1300–1750 m). It occurs mainly on sites with DIFFERENTIAL TAXA: subsp. hun- extreme climatic conditions, such as ridges and garica, Asplenium viride, Campanula cochleariifolia, edges exposed to strong winds, where the snow Gentiana clusii, Sesleria albicans, Cololejeunea layer is thin and of brief duration. It occupies calcarea, Didymodon giganteus, Entodon concinnus, Hymenostylium recurvirostrum, Mnium thomsonii, Or- moderate to steep (20–80°), mostly north-facing thothecium rufescens, Racomitrium lanuginosum. slopes, rocky walls and rocky crevices with cold and humid climate. Rarely it occurs also on sites NOMENCLATURAL TYPE: identical with the type of the association name. oriented in other directions (SW, W); in these cases it is restricted to rocky walls and stabilized Stands of the subassociations D.-C. primu- screes, and forms a type transitional towards rock letosum auriculae occur mainly on steep, north- fi ssure communities. The soils inhabited by these facing rocky sites, small narrow terraces and rocky communities are black, strongly humic, neutral crevices, but can be found also on windy rocky to slightly basic protorendzinas (= Rendzi-Lithic ledges and slopes. Several taxa of the class Asple- Leptosols), with many bedrock fragments; due nietea trichomanes indicate relationships to rock to the extreme climatic conditions on mountain communities. As already noted by Sillinger (1933: edges, raw litter accumulates among tufts of 224), two ecological variants can be distinguished, Carex fi rma (Sillinger 1933: 226; Bělohlávková with more or less identical fl oristic composition. & Fišerová 1976: 138–139). On mainly north-facing rocky terraces and shady This is an autochthonous community growing rocky crevices, stands with a well-developed moss on limestone and dolomite bedrock, inhabiting layer occur. Compact tufts of Carex fi rma and mats steep, often shady rocky slopes and cliffs. The of Dryas octopetala, together with mosses, play J. ŠIBÍK ET AL.: SYNTAXONOMICAL REVISION OF PLANT COMMUNITIES 197 an important role in retaining moisture on the grophilous and much more closed, due to the pres- site. On very exposed sites, such as windy slopes ence of compact tufts of Carex fi rma and mats of and mountain edges, the layer of mosses is less Dryas octopetala. On sites with extremely exposed developed, and more xerophilous species like slopes, typical garland soils can be observed. Racomitrium lanuginosum also occur at higher The subassociation is developed on north- frequency. This subassociation is documented (rarely west)-facing sites on mountain edges from the Krivánska Malá Fatra, Chočské vrchy with very foggy climate. So far it has been docu- and Nízke Tatry Mts. mented by phytocoenological relevés only from the Krivánska Malá Fatra Mts, where it occurs Dryado octopetalae-Caricetum fi rmae saxifrage- extensively on north and northwest slopes of Veľký tosum aizoidis subass. nov. hoc loco Kriváň Mt., Malý Kriváň Mt. and Chleb Mt. In the past, communities of this subassociation Table 1, column C2 probably covered smaller areas. After destruction DIFFERENTIAL SPECIES: Saxifraga aizoides, Biscutella of the subalpine shrubs during the Walachian laevigata, Festuca supina, Huperzia selago, Parnassia colonization in the 16th and 17th centuries (cf. palustris, Pyrola carpatica, Salix alpina, Scabiosa Janík 1971: 69; Plesník 1955: 30) they spread lucida, Vaccinium myrtillus, Dicranum scoparium, Pogonatum urnigerum. from enclaves among Pinus mugo stands and from steep slopes. NOMENCLATURAL TYPE: Šibík & Dúbravcová ined., On sites with a thicker accumulated layer of holotypus undecomposed humus, the occurrence of acido- LOCALITY: Krivánska Malá Fatra Mts, Veľký Kriváň philous species (e.g., Empetrum hermaphroditum) Mt., near the hiking trail giving from ground elevation and bryophytes (Sphagnum spp.) increases; succes- ‘Hrana Veľkého Kriváňa’ to height, alt. 1679 m, as- sion processes may further lead to the formation pect: NNW, slope: 20°, relevé area: 25 m2, cover total: of Pinus mugo stands. These processes are very 95%, E1: 95%, E0: 30%, 49°11′20,9″; 19°01′47,0″, 1. 8. 2002 slow, however, because stands of the subasso- ciation D.-C. saxifragetosum aizoidis represent E1: Carex fi rma 4, Dryas octopetala 3, Festuca a blocked successional stage with a fi xed and versicolor 2a, Tofi eldia calyculata 2a, Bartsia alpina closed species composition. 1, Biscutella laevigata 1, Bistorta vivipara 1, Pinguicula alpina 1, Ranunculus alpestris 1, Saxifraga aizoides The strongly hygrophilous character of the 1, Selaginella selaginoides 1, Vaccinium vitis-idaea 1, stands indicates some transition to the association Anemone narcissifl ora +, Anthyllis *alpestris +, Belli- Androsaceo lacteae-Festucetum versicoloris. diastrum michelii +, Campanula cochleariifolia +, Di- At present, communities of the association anthus nitidus +, Empetrum hermaphroditum +, Galium D.-C. saxifragetosum aizoidis are endangered anisophyllon +, Gentiana verna +, Gentianella *tatrae by planting of Pinus mugo (cf. Bernátová et al. +, Helianthemum grandifl orum +, Huperzia selago +, 1998: 50, 51), as compact tufts of Carex fi rma are Parnassia palustris +, Pedicularis verticillata +, Phy- often destroyed, and alien species introduced. Near teuma orbiculare +, Salix alpina +, Saxifraga paniculata hiking trails there is also a risk of trampling. +, Scabiosa lucida +, Soldanella carpatica +, Vaccinium myrtillus +, Veronica aphylla +, Viola bifl ora +. Androsaceo lacteae-Festucetum versicoloris E0: Hylocomium splendens 2a, Tortella tortuosa 2a, Bryum sp. 1, Campylium stellatum 1, Ditrichum fl exicaule Sillinger 1933 1, Hypnum cupressiforme 1, Orthothecium rufescens 1, Table 1, columns D, Ds Rhytidium rugosum 1, Sanionia uncinata 1, Ctenidium ORIGINAL FORM OF THE NAME: Ass. Festuca versicolor- molluscum +, Hypnum bambergeri +, H. vaucheri +, Androsace lactea Sillinger 1933 Rhizomnium punctatum +, Rhytidiadelphus triquetrus +, Cetraria islandica +, Icmadophila ericetorum +. DIAGNOSTIC TAXA: Festuca versicolor (dom., const.), Androsace lactea (dif.), Cortusa matthioli (dif.), In comparison with the previous subasso- Cystopteris montana (dif.), Poa alpina (dif.), Saxi- ciation, D.-C. saxifragetosum aizoidis are more hy- fraga moschata (dif.), Viola bifl ora (dif.), Bartsia alpina 198 POLISH BOTANICAL JOURNAL 49(2). 2004

(const.), Dryas octopetala (const.), Galium anisophyllon DISCUSSION AND CONCLUSION (const.), Pedicularis verticillata (const.), Ranunculus alpestris (const.), Salix alpina (const.), Saxifraga pa- In phytocoenological studies, Carex fi rma has niculata (const.), Sesleria tatrae (const.), Soldanella been treated mostly as a diagnostic taxon of the carpatica (const.), Swertia perennis subsp. alpestris alliances Caricion fi rmae (char.) and Potentillion (const.), Vaccinium vitis-idaea (const.) caulescentis (dif.). West Carpathian populations of NOMENCLATURAL TYPE: Milová & Urbanová 1989: 295, this species occupy steep calcareous and dolomite rel. 1, neotypus hoc loco slopes and rocky walls in the montane and alpine This is a two-layer community, not very rich belts; more rarely they grow also on inversion in species (at average 27 taxa per relevé), deter- sites in the Muránska planina, Slovenský raj, mined by compact tufts of the dominant species Malá Fatra, Veľká Fatra, Chočské vrchy, Nízke and mosses. The occurrence of several hygrophi- Tatry, Západné Tatry and Belianske Tatry Mts, lous species (Androsace lactea, Cortusa matthioli, including calcareous foothills of the Polish Tatra Ranunculus alpestris, Saxifraga moschata, Swertia Mts. In most of those mountain ranges (except *alpestris, Viola bifl ora) indicates the high moisture for the Veľká Fatra Mts), Carex fi rma often co- of these stands. The communities are developed occurs with Dryas octopetala, which also has its in the highest parts of mountains on steep rocky coenological optimum in the alliance Caricion slopes, and on lee terraces in relatively protected fi rmae. The abundance ratio of these two species areas in the supramontane to subalpine belt (alt. in plant communities varies, ranging from almost 1450–1650 m). They inhabit shallow black skel- equal abundance to the strong dominance of one etal soils on limestone and dolomite bedrock (pro- of them. torendzinas) with high content of humus. Very wet Signifi cant differences in the abundance of and shady, mostly north-facing sites are occupied. Carex fi rma and Dryas octopetala, altering the In the winter, snow lies longer, especially on less physiognomy of the respective stands, led Szafer steep terraces or on lee sites free of strong winds. et al. (1923: 43–46, Tabs 12, 13) to distinguish It is an autochthonous and stable community two separate associations: Caricetum fi rmae and growing on shady rocks, developed from some Dryadetum. Similarly, Hadač et al. (1969: 59–64) rock fi ssure communities with Festuca versicolor reported two associations from the Dolina Sied- as the dominant species. Its fl oristic composition mich prameňov valley in the Belianske Tatry Mts: resembles very wet stands of the association Caricetum fi rmae carpaticum (Br.-Bl.) Pawł. 1956 Dryado octopetalae-Caricetum fi rmae (Sillinger and Dryadetum octopetalae tatricum. Hadač (in 1933: 229). Higher chionophility together with Mucina & Maglocký 1985: 187) followed the high constancy of Ranunculus alpestris and concept of Szafer et al. (1923), and published the Saxifraga wahlenbergii point to its intermediate names Saxifrago caesiae-Caricetum fi rmae and position towards communities of the alliance Festuco versicoloris-Dryadetum, replacing ille- Arabidion coerulae. gitimate names (the names Caricetum fi rmae and The community is documented by phytocoeno- Dryadetum octopetalae were already published logical relevés or a synoptic table from the highest by Rübel 1911: 94, 95, 176). Plant communi- parts of the Krivánska Malá Fatra, Chočské vrchy ties with Dryas octopetala have been included and Nízke Tatry Mts; Sillinger (1933: 232) re- in the association Caricetum fi rmae (Firmetum) ported it also from the Lúčanská Malá Fatra Mts by other authors also, either as facies (Pawłowski (Kľak Mt.), and from the Veľká Fatra Mts. Judging & Stecki 1927: 91) or as a stage (Klika 1932: by its fl oristic composition, a relevé of the associa- 150). Sillinger (1933: 221) was the fi rst to show tion Festucetum versicoloris Domin 1925 from the that dominance change does not bring alterations supramountane belt of the Krivánska Malá Fatra in the overall composition of the community, and (Bělohlávková & Fišerová 1978: 108) can be also stressed this by describing a new association, assigned to this subassociation. Dryadeto-Firmetum. J. ŠIBÍK ET AL.: SYNTAXONOMICAL REVISION OF PLANT COMMUNITIES 199

The plant communities of the alpine and sub- surrounding classifi cation of plant communities alpine belts in the central Western Carpathians are with dominating Carex fi rma and Dryas octopetala treated variously by different authors. Pawłowski stimulated us to make the syntaxonomical revision and Stecki (1927: 90) noted the variability of presented here. Along with already published phy- fl oristic composition, and distinguished two alti- tocoenological relevés, numerous unpublished data tudinal elements (variants?): an alpine one with from the area of Slovakia were used. The results of Viola alpina, Minuartia sedoides, Salix retusa and this revision can be summarized as follows: Ligusticum mutellinoides: and a subalpine one with 1. Our results do not support division of the Gentiana clusii, Leontopodium alpinum, Tofi eldia plant communities with Carex fi rma and Dryas calyculata, and a higher proportion of Campanula octopetala growing on limestone-dolomite bed- cochleariifolia. Klika (1932: 152) used two names rock into two associations on the basis of their to differentiate the subalpine communities of the physiognomy (as reported by Szafer et al. 1923; Krivánska Malá Fatra Mts from the alpine com- Hadač et al. 1969; Hadač in Mucina & Maglocký munities of the Tatra Mts, but without presenting 1985). Our revision supports the concept proposed relevés: Firmetum chočense and Caricetum fi rmae by Sillinger (1933: 221), that is, it shows that dif- tatricum, respectively. Sillinger (1933: 224) also ferences in the abundance of the two species do emphasized the higher species diversity of phyto- not signifi cantly alter the fl oristic composition of coenoses in the Tatras. As there were only nega- the respective phytocoenoses. Nevertheless, the tive differences between the communities from the dwarf phytocoenoses with prevalence of Dryas Nízke Tatry Mts and those from the Tatra Mts, he octopetala and Salix reticulata occurring in the considered the former to be only a type of the as- alpine belt of the Belianske Tatry Mts (Table 1, sociation Caricetum fi rmae (Dryadeto-Firmetum) column E), fi rst analyzed by Domin (1929: 9–10) tatricum from the Tatras, somewhat poorer in fl o- under the name Dryadeto-Salicetum reticulatae, ristic composition. In line with this, several other were found to represent a separate, well-defi ned authors included both alpine and subalpine com- plant community, different from phytocoenoses of munities under the single name Caricetum fi rmae the alliance Caricion fi rmae despite their sharing (Firmetum) carpaticum (Pawłowski 1935: 132; of several species. These communities will be ana- Hadač et al. 1969: 55; Unar et al. 1985: 32). lyzed in detail elsewhere (Petrík et al. in prep.). Grabherr et al. (1993: 406) merged West 2. In accord with Klika (1932: 152), we found Carpathian subalpine and alpine phytocoenoses signifi cant differences in fl oristic composition be- with Carex fi rma with those of the Alpine asso- tween the subalpine communities of the Krivánska ciation Caricetum fi rmae Rübel 1911. However, Malá Fatra, Chočské vrchy and Nízke Tatry Mts, Vierhapper (1930: 146) and Pawłowski (1935: and those of the alpine to subalpine belts of the 132) had already pointed to differences in their Západné and Belianske Tatry Mts (Table 1, col- fl oristic composition, and this was also noted, umns B, C). These two groups of phytocoenoses although indirectly, by Braun-Blanquet (1930: are treated here as two different associations. 25) when he described the association Caricetum Stands from the subalpine belts can be attributed fi rmae tatricum. to the association Dryado octopetalae-Caricetum Somewhat deviating plant communities from fi rmae Sillinger 1933. Phytocoenoses from the the montane belt of the Veľká Fatra Mts, from cold Tatra Mts were already clearly defi ned and de- inversion sites of the Krivánska Malá Fatra and limited by Braun-Blanquet (1930: 25). For their Slovenský raj Mts with dominating Carex fi rma, designation, we therefore use the name Arenario have often been assigned to the association Cari- tenellae-Caricetum fi rmae (Br.-Bl. 1930) nom. cetum fi rmae or the alliance Caricion fi rmae (Klika nov. (basionym: Caricetum fi rmae tatricum Br.- 1932: 155–156; Šimeková & Pitoniak 1976: Table Bl. 1930). 2; Bernátová et al. 1982: 142–143). 3. The treatment of West Carpathian phyto- These confl icting treatments and the confusion coenoses with Carex fi rma as identical with the 200 POLISH BOTANICAL JOURNAL 49(2). 2004

Alpine association Caricetum fi rmae Rübel 1911 Eduard Králik (Euphrasia, Myosotis), Martin A. Lysák (Grabherr et al. 1993: 406) was found to be erro- (Sesleria), Pavol Mártonfi (Thymus) and Patrick Mráz neous. Besides the numerous phytogeographically (Hieracium). For determination of moss specimens we important species occurring in only one of these are grateful to Katarina Mišíková, Katarina Kresáňová, Barbora Lučeničová and Zdeněk Pilous; lichens were two mountain ranges (Alps: Achillea clavenae, kindly identifi ed by Anna Guttová and Eva Lisická. Arabis pumila, Campanula scheuchzeri, Carex We thank Zuzana Dúbravcová and Ivana Krajčiová- mucronata, Festuca pumila, Gentiana terglouensis, Šibíková, the co-authors of several phytocoenological Pedicularis rosea, P. rostrato-capitata, Sesleria relevés included in this revision, as well as Rad- sphaerocephala, Soldanella alpina, S. minima, mila Bělohlávková, Dana Bernátová, Ladislav Mucina, etc.; Western Carpathians: Arenaria tenella, Jana Uhlířová and Milan Valachovič for providing Campanula tatrae, Cerastium arvense subsp. their unpublished phytocoenological relevés, and glandulosum, Festuca versicolor subsp. versicolor, the anonymous reviewer for valuable suggestions on Leontodon pseudotaraxaci, Oxytropis carpatica, the manuscript. We are indebted to Viera Banásová, Ivan Jarolímek and especially to Judita Lihová for re- Primula auricula subsp. hungarica, Pyrola car- viewing the English. This work was supported by the patica, Saxifraga wahlenbergii, Sesleria tatrae, VEGA grant agency (grant nos 1/7452/20, 1/9149/02 Soldanella carpatica, Thymus pulcherrimus, and 1/0045/03). etc.), there were also differences in the fl oristic composition of the respective stands; in particular, several common species are lacking in the poorer REFERENCES Alpine stands (Table 1, columns A–C). 4. According to our results, plant communities BARKMAN J. J., DOING H. & SEGAL S. 1964. Kritische Be- with dominance of Carex fi rma occurring on lime- merkungen und Vorschläge zur quantitativen Vegetation- sanalyse. Acta Bot. Neerl. 13: 394–419. stone-dolomite peaks of the Veľká Fatra, Lúčanská Malá Fatra and Muránska planina Mts, and from BĚLOHLÁVKOVÁ R. 1980. Rostlinná společenstva alpínského stupně Kriváňské Malé Fatry. Ph.D. Thesis, Faculty of inversion sites of the Krivánska Malá Fatra and Natural Sciences, Charles University, Prague. Slovenský raj Mts (Table 1, columns G, H), usually BĚLOHLÁVKOVÁ R. & FIŠEROVÁ D. 1976. Pyrola carpatica classifi ed in the alliance Caricion fi rmae, in fact – eine neue Art in Gebirge Krivánska Malá Fatra. Preslia belong to rock fi ssure communities of the alliances 48: 137–142 (in Czech with German summary).

Potentillion caulescentis and Cystopteridion (Le- BĚLOHLÁVKOVÁ D. & FIŠEROVÁ D. 1978. New localities of ontopodio alpini-Campanuletum cochleariifoliae, some vascular plants in Slovakia. Zprávy Českoslov. Bot. Bellidiastro michelii-Campanuletum cochleariifo- Společn. Českoslov. Akad. Ved 13: 107–112 (in Czech with liae) or to phytocoenoses of the association Bel- English summary). lidiastro michelii-Seslerietum calcariae (alliance BERNÁTOVÁ D., ŠKOVIROVÁ K. & KLIMENT J. 1982. Flora and Seslerio-Asterion alpini) advanced in succession. extrasilvatic phytocoenoses of the State nature reserva- tion Čierny Kameň in the mountains Veľká Fatra. Ochr. Also, stands from the montane to subalpine belts Prír. (Bratislava) 3: 137–152 (in Slovak with English of the Polish West Tatras (Szafer et al. 1923; summary). Pawłowski & Stecki 1927) are much more similar BERNÁTOVÁ D., UHLÍŘOVÁ J. & TOPERCER J. 1998. Some new to the communities of these alliances. The question information on subalpine vegetation of the Krivánska Fatra of their fi nal classifi cation will be addressed in Mts, and management recommendations. In: M. KORŇAN (ed.), Výskum a ochrana Krivánskej Fatry, pp. 49–51. another paper (Šibík & Kliment in prep.). Správa Národného parku Malá Fatra, Varín (in Slovak with English summary). ACKNOWLEDGEMENTS. The authors are grateful to Viera Banásová, Zuzana Dúbravcová, Ivan Jarolímek, BRAUN-BLANQUET J. 1930. Zentralalpen und Tatra, eine František. Krahulec, Ján Ripka and Milan Valachovič for pfl anzensoziologische Paralelle. Veröff. Geobot. Inst. Rübel Zürich 6: 81–123. their valuable comments and suggestions on this paper, as well as for providing publications diffi cult to access. BRAUN-BLANQUET J. 1964. Pflanzensoziologie. Grundzüge der Specimens of some taxonomically diffi cult species were Vegetationskunde. 3. Aufl . Springer Verlag, Wien. determined or revised by Jan Kirschner (Gentianella), BRAUN-BLANQUET J. & JENNY J. 1926. Vegetationsentwicklung J. ŠIBÍK ET AL.: SYNTAXONOMICAL REVISION OF PLANT COMMUNITIES 201

und Bodenbildung in der alpinen Stufe der Zentralalpen. JANÍK M. 1971. Pastva na holiach krivánskej Malej Fatry a jej Denkschr. Schweiz. Naturforsch. Ges. 63: 183–349. negatívny vplyv na krajinu. životné prostredie 5: 69–75.

COLDEA G., SANDA V., POPESCU A. & ŞTEFAN N. 1997. Les JAROLÍMEK I. & SCHLOSSER G. 1997. FYTOPACK – a system associations végétales de Roumanie. 1. Les associations of programs to process phytocoenological tables. Biologia herbacées naturelles. Presses Universitaires de Cluj, (Bratislava) 52: 53–59. Cluj-Napoca. KLIKA J. 1926. Quelques remarques géobotaniques sur le Velká DOMIN K. 1925a. Které rostliny stoupají až na nejvyšší vrc- Fatra en Slovaquie. Sborník Přírodovědské Společnosti holek Bielských Tater? Věda Přír. 6: 258–259. v Moravské Ostravě 3(1924–1925): 38–85 (in Czech with French summary). DOMIN K. 1925b. The flora of the kettle-like mountain-valley between Žďárská Vidla and Havran in the Tatras of Biela. KLIKA J. 1932. Der Seslerion coerulae-Verband in den West- Spisy Přír. Fak. Karlovy Univ. 45: 1–30 (in Czech with karpathen. Beih. Bot. Centralbl. 49B: 133–175. English summary). KLIMENT J., BERNÁTOVÁ D., JAROLÍMEK I. & UHLÍŘOVÁ DOMIN K. 1928. The relations of the Tatra mountain vegetation J. 2004. Floristic composition and syntaxonomy of the to the edaphic factors of the habitat. Acta Bot. Bohem. communities with Carex sempervirens subsp. tatrorum 6–7: 133–163. in the West Carpathians. Biologia (Bratislava) 59 (in DOMIN K. 1929. Contribution to the knowledge of the veg- press). etation and fl ora of the Mt. Malý Havran in the Tatras of MAAREL VAN DEN E. 1979. Transformation of cover-abundance Biela. Spisy Přír. Fak. Karlovy Univ. 101: 3–18 (in Czech values in phytosociology and its effect on community simi- with English summary). larity. Vegetatio 39: 97–114.

DOMIN K. 1930. Zur Soziologie der chionophytischen Pflan- MARHOLD K. & HINDÁK F. (eds) 1998. Checklist of non-vas- zenassoziationen des Tatragebirges. Veröff. Geobot. Inst. cular and vascular plants of Slovakia. Veda, Bratislava. Rübel Zürich 6: 167–190. MARHOLD K. & VALACHOVIČ M. 1990. Distribution and DOMIN K. 1931. Květena našich Tater, její společenstva ecology of Petrocallis pyrenaica (L.) R. Br. in Belanské a vztahy k podnebí a půdě. Českoslovanská akciová Tatry Mts. Biológia (Bratislava) 45: 433–439 (in Slovak tiskárna, Praha. with English summary).

DOMIN K. 1936. Schedae ad Floram Čechoslovenicam exsic- MILOVÁ M. & URBANOVÁ V. 1989. Non-forest plant associa- catam. Centuria IV. Acta Bot. Bohem. 11: 3–50. tions of the State nature reserve Prípor. Ochr. Prír. (Bratis- DÚBRAVCOVÁ Z. 1996. Spoločenstvá triedy Juncetea trifi di lava) 10: 289–309 (in Slovak with English summary). v Západných Karpatoch. Ph.D. Thesis, Faculty of Natural MUCINA L. & MAGLOCKÝ Š. (eds) 1985. A list of vegetation Sciences, Comenius University, Bratislava. units of Slovakia. Docum. Phytosociol. N. S. (Camerino) DÚBRAVCOVÁ Z. & HAJDÚK J. 1986. Ein Beitrag zur erfor- 9: 175–220. schung der Vegetation der subalpinen Stufe des Berges PAWŁOWSKI B. 1935. Über die Klimaxassoziation in der al- Sivý vrch im westlichen Tatragebirge. Zborn. Slov. Nár. pinen Stufe der Tatra. Bull. Int. Acad. Polon. Sci. Lett., Cl. Múz., Prír. Vedy 32: 33–54 (in Slovak with German sum- Sci. Math. Nat., Ser. B, Sci. Nat. (1)(1935): 115–146. mary). PAWŁOWSKI B. & STECKI K. 1927. Die Pflanzenassoziationen EHRENDORFER E. (ed.) 1973. Liste der Gefässpflanzen Mit- des Tatra-Gebirges. IV. Teil: Die Pfl anzenassoziationen teleuropas. Gustav Fischer Verlag, Stuttgart. des Miętusia-Tales und des Hauptmassivs der Czerwone FUTÁK J. 1976. Phytogeographical division of the Tatra national Wierchy. Bull. Int. Acad. Polon. Sci. Lett., Cl. Sci. Math. park and its relation to the neighbouring regions. Zborník Nat., Ser. B, Sci. Nat., Suppl. 2: 79–121. prác o Tatranskom národnom parku 17(1975): 109–131 PODANI J. 1993. SYN-TAX-pc. Computer Programs for Mul- (in Slovak with English summary). tivariate Data Analysis in Ecology and Systematics. Ver. GRABHERR G. & MUCINA L. (eds) 1993. Die Pflanzengesells- 5,0. Scientia Publ., Budapest. chaften Österreichs. 2. Natürliche waldfreie Vegetation. PLESNÍK P. 1955. Vplyv pasenia na lesy Krivánskej Malej Gustav Fischer Verlag, Jena. Fatry. Les 2(1–2): 29–37. GRABHERR G., GREIMLER J. & MUCINA L. 1993. Seslerietea RÜBEL E. 1911. Pflanzengeographische Monographie des albicantis. In: G. GRABHERR & L. MUCINA (eds), Die Berninagebietes. Bot. Jahrb. Syst. 47: 1–646. Pfl anzengesellschaften Österreichs. 2: 402–446. Gustav Fischer Verlag, Jena. SILLINGER P. 1933. Monografická studie o vegetaci Nízkých Tater. Orbis, Praha. HADAČ E., BŘEZINA P., JEŽEK V., KUBIČKA J., HADAČOVÁ V., VONDRÁČEK M. et al. 1969. Die Pfl anzengesellschaften des SZAFER W., PAWŁOWSKI B. & KULCZYŃSKI S. 1923. Die Tales “Dolina Siedmich prameňov” in der Belaer Tatra. Pfl anzenassoziationen des Tatra-Gebirges. I. Teil: Die Vegetácia ČSSR, Ser. B 2: 1–343. Pfl anzenassoziationen des Chochołowska-Tales. Bull. 202 POLISH BOTANICAL JOURNAL 49(2). 2004

Int. Acad. Polon. Sci. Lett., Cl. Sci. Math. Nat., Ser. B, Sci. Tomanova dolina valley and žľab spod Diery glen in the Nath. Suppl. (1)(1923): 1–66. Západné Tatry Mts. 1. Phytocoenological tables. Folia Fac. Sci. Nat. Univ. Purkynianae Brun., Ser. Biol. 25(10): 5–101 ŠIBÍK J. 2003. Nelesné spoločenstvá subalpínskeho stupňa (in Czech with English summary). Krivánskej Malej Fatry. M.Sc. Thesis, Faculty of Natural Sciences, Comenius University, Bratislava. UNAR J., UNAROVÁ M. & ŠMARDA J. 1985. Vegetation of the Tomanova dolina valley and žľab spod Diery glen in ŠIMEKOVÁ J. & PITONIAK P. 1976. Relic locality of the eight- petaled dryad (Dryas octopetala L.) near the Vernár in the the Západné Tatry Mts. 2. Description of natural condi- natural area of the Slovenský raj Mts. Biológia (Bratislava) tions and plant communities. Folia Fac. Sci. Nat. Univ. 31: 783–793 (in Slovak with English summary). Purkynianae Brun., Biol. 26(14): 5–78 (in Czech with English summary). ŠMARDA J. 1956. The vegetable cover of erosively denuded and tundra-like soils in the Tatra Mts. Biol. Práce Slov. Akad. VALACHOVIČ M. & JAROLÍMEK I. 1994. Plant communities Vied 2(8): 5–50 (in Czech with English summary). with Daphne arbuscula Čelak. in Muránska planina Plain. Bull. Slov. Bot. Spoločn. 16: 75–82 (in Slovak with English ŠMARDA J., LAZEBNÍČEK J., MATOUŠOVÁ L., NETOPIL R., summary). PACLOVÁ L. & PIKULA J. 1971. To the ecology of plant communities of the Valley Dolina Sedmi pramenů in VALACHOVIČ M., OŤAHEĽOVÁ H., STANOVÁ V. & M AGLOCKÝ Belanské Tatry. Práce a štúdie Československej ochrany Š. 1995. Rastlinné spoločenstvá Slovenska. 1. Pionierska prírody pri SÚPSOP v Bratislave), Ser. 3 4: 5–204 (in vegetácia. Veda, Bratislava. Czech with English summary). VIERHAPPER F. 1930. Vergleichende Studien über die Pflan- ŠVANDOVÁ-URSÍNIOVÁ Ľ. 1966. Skalné spoločenstvá zenassoziationen der Nordkarpathen und Ostalpen. Veröff. a spoločenstvá plytkých vápencových pôd Chočského Geobot. Inst. Rübel Zürich 6: 134–166. pohoria. M.Sc. Thesis, Faculty of Natural Sciences, WEBER H. E., MORAVEC J. & THEURILLAT J. P. 2000. Inter- Comenius University, Bratislava. national Code of Phytosociological Nomenclature. Ed. 3. UNAR J., UNAROVÁ M. & ŠMARDA J. 1984. Vegetation of the J. Veget. Sci. 11: 739–768.

Received 2 February 2004