original scientific article UDC 581.5:581.9(234.323.6) received: 2005-10-19 PHYTOGEOGRAPHY AND SYNTAXONOMY OF SNOW-BED VEGETATION ON CALCAREOUS SUBSTRATES IN THE SOUTH-EASTERN ALPS: A NUMERICAL APPROACH Boštjan SURINA University of Primorska, Science and Research Centre Koper, SI-6000 Koper, Garibaldijeva 1 E-mail: Bostjan.surina@zrs-kp.si ABSTRACT Based on cluster analysis and the phytogeographic peculiarities of the area, an assessment of the vegetation of snow-beds on calcareous soils (Arabidetalia caeruleaej in the South-eastern Alps was made. Eight distinct associations belonging to three alliances were recognised: Saxifragetum stellaro-sedoidis, Saxifragetum hohenwartii and Ranunculo traunfellneri-Festucetum nitidae (alliance Arabidion caeruleaej, Salici herbaceae-Arabidetum caeruleae (alliance Salici herbaceae-Arabidion caeruleaej, Salicetum retuso-reticulatae, Homogyno discoloris-Salicetum re-tusae, Salici retusae-Geranietum argentei and Potentillo brauneanae-Homogynetum discoloris (alliance Soldanello alpinae-Salicion retusaej. Saxifragetum stellaro-sedoidis was further subdivided into three geographical variants, such as variants of Achillea oxyloba, Ranunculus traunfellneri and Campanula pulla. The Drepanoclado uncinati-Heliospermetum pusilli from the Liburnian Karst (Dinaric Mts.j showed clear floristic and phytogeographic distinctions and its placement into the Dinaric alliance Salicion retusae was confirmed. Key words: biogeography, syntaxonomy, endemic species, cluster analysis, snow-beds, the Alps, Arabidetalia caeruleae, Thlaspietea rotundifolii FITOGEOGRAFIA E SINTASSONOMIA DELLA VEGETAZIONE DEL MANTO NEVOSO SU SUBSTRATI CALCAREI NELLE ALPI SUD-ORIENTALI: APPROCCIO NUMERICO SINTESI In base alla "cluster analysis" ed alle peculiarity fitogeografiche dell'area, gli autori hanno preparato una valutazi-one della vegetazione del manto nevoso di terreni calcarei (Arabidetalia caeruleaej delle Alpi sud-orientali. Otto as-sociazioni distinte, appartenenti a tre alleanze, sono state riconosciute: Saxifragetum stellaro-sedoidis, Saxifragetum hohenwartii e Ranunculo traunfellneri-Festucetum nitidae (alleanza Arabidion caeruleaej, Salici herbaceae-Arabidetum caeruleae (alleanza Salici herbaceae-Arabidion caeruleaej, Salicetum retuso-reticulatae, Homogyno dis-coloris-Salicetum retusae, Salici retusae-Geranietum argentei e Potentillo brauneanae-Homogynetum discoloris (alleanza Soldanello alpinae-Salicion retusaej. Saxifragetum stellaro-sedoidis e stato ulteriormente suddiviso in tre varianti geografiche, ossia le varianti di Achillea oxyloba, Ranunculus traunfellneri e Campanula pulla. Drepanoclado uncinati-Heliospermetum pusilli proveniente dal Carso liburniano (montagne dinarichej si e chiaramente distinto floristicamente e fitogeograficamente, ed e stata confermata la sua collocazione nell'alleanza dinarica Salicion re-tusae. Parole chiave: biogeografia, sintassonomia, specie endemiche, cluster analysis, snow-beds, Alpi, Arabidetalia caeruleae, Thlaspietea rotundifolii INTRODUCTION The order Arabidetalia caeruleae comprises pioneer or permanent hygrophilous stands occurring on more or less stabilised calcareous screes and moraines with a long duration of snow cover, and from the altimontane to subnival belt in Central and south-eastern Europe (Braun-Blanquet, 1926; Horvat, 1931; Englisch et al., 1993). The floristic composition, ecology and physiognomy of Arabidetalia stands are relatively well characterised and distinctions from other scree stands are usually not questionable. Recently, a comprehensive study of the snow-bed vegetation in the Northern Calcareous Alps (Englisch, 1999) resulted in the recognition of three floristically and ecologically well defined alliances in the Alps, including the Arabidion caeruleae, Salici her-baceae-Arabidion caeruleae and Soldanello alpinae-Salicion retusae. However, although this study also included some of the relevés from the South-eastern Calcareous Alps, no similar treatise of syntaxa from Arabidetalia in the South-eastern Alps has yet been provided. The aim of the paper is thus to study, on the basis of a comprehensive synoptic table and cluster analyses, the ecology and syntaxonomy of the order, with an emphasis on the phytogeographic peculiarities of the area, as well as to summarise current knowledge on the phyto-sociology and ecology of syntaxa of Arabidetalia in the South and South-eastern Alps. The phytogeographic peculiarities of the flora and vegetation of the South-eastern (Calcareous) Alps (Fig. 1) have been recognised for a long time. In comparison to other Alpine regions, they are characterised by a significant number of conservative, progressive, absolute and relative endemics, southern-, south-eastern - Alpine, north - Illyrian and Illyrian (Illyricoid) species (Engler, 1901; Mayer, 1960a, 1960b; Wraber, 1995; Tribsch & Schonswetter, 2003; Surina, 2004a). According to Paw-lowski (1970), the South-eastern Calcareous Alps are flo-ristically the second richest region of the Alps. The particularity of its flora and vegetation are due to their specific origin as a result of the historical, geographical and ecological peculiarities of the area. The first phytosociological studies in the area on frigiphilous and hygrophilous scree vegetation were performed by Aichinger (1933: columns 1, 2, 14 in Table 1) in the Karavanke Mts. Further research in the South-eastern Calcareous Alps (Julian Alps, Kamnik Alps and the Karavanke Mts.) was implemented by Wraber (1972: 7, 8, 15) and Surina (2004c: 4, 12, 13) in the Julian Alps and by Haderlapp (1982: 14) in the Kamnik Alps. Poldini & Martini (1993: 17) studied the calcareous hygrophilous and frigiphilous snow-bed and talus slope vegetation in the Carnic Alps, Wikus (1959: 3, 9, 10) in the Dolomites of Lienz, and Lasen (1983: 18) in the Dolomites of Belluno, while Gerdol & Piccoli (1982: 18) engaged in its research in Monte Baldo. Pignatti & Pig- natti (1983) studied the vegetation of Vette di Feltre (Dolomites of Belluno) and with two unpublished relevés identified the association Salicetum retuso-reticulatae. The same phytosociologists published an analysis and overview of the calcareous scree vegetation (Thlaspietalia rotundifolii) in the South-eastern Alps (Pignatti & Pignatti, 1984), while Wraber (1970) published a paper on vegetation of the SE Alps. In comparison to the SE Alps, the snow-bed vegetation on calcareous soils in the Northern Calcareous Alps has been studied by many phytosociologists. The results were summarised by Englisch et al. (1993) and lately by Englisch (1999). North-easter JJ Central Alps (át) ^South-eastern ^fg^^ O Dolomites A ^ ÏÇ^ Calcareous*^ (HU) ( IT) Alps (SLÇ) (o A©' (HR.) Li burn ¡an Karst VB) t Fig. 1: Research area. 1 - Bernina; 2 - Monte Baldo; 3 -Dolomites of Belluno; 4 - Dolomites of Lienz; 5 - Carnic Alps; 6 - Mt. Dobrač; 7 - Julian Alps; 8 - Karavanke Mts.; 9 - Kamnik Alps; 10 - Gesause; 11 - Hochschwab; 12 - Trnovski gozd plateau; 13 - Mt. Snežnik. Sl. 1: Raziskovano območje. 1 - Bernina; 2 - Monte Baldo; 3 - Bellunški Dolomiti; 4 - Lienški Dolomiti; 5 -Karnijske Alpe; 6 - Dobrač; 7 - Julijske Alpe; 8 - Karavanke; 9 - Kamniške Alpe; 10 - Gesause; 11 -Hochschwab; 12 - Trnovski gozd; 13 - Snežnik. MATERIAL AND METHODS Phytosociological research of calcareous scree vegetation of Arabidetalia was conducted by applying the sigmatistic method (Braun-Blanquet, 1964; Westhoff & Maarel, 1973; Dierschke, 1994). Altogether, 232 relevés from the SE Alps (between Monte Baldo and the Kamnik Alps) and NE Alps were used in the analysis (see the Appendix). With the aid of SYN-TAX 2000 computer programme (Podani, 2001) and a synoptic phytosociological table (Tab. 1), comparisons between stands were made after the transformation of cover-abundance values according to van der Maarel (1979) had been performed. The measure of dissimilarity was the complement of the "Similarity Ratio" coefficient. We used the Furthest Neighbour - Complete Linkage clustering method (CL) and the Principal Coordinates Analysis (PCoA) ordination method. Coverage index (D%, see Surina, 2004b, 2005) was calculated for each taxon separately. Groups of diagnostic species were formed on the basis of our own criteria, but with regard to numerous authors. Three floristically and/or ecologically similar syntaxa were included in the analyses, i.e. Saxifra-getum hohenwartii (Thlaspion rotundifolii) from the Karavanke Mts., Poo supinae-Cerastietum cerastoidis (Salicion herbaceae, Salicetalia herbaceae) from the Dolomites of Lienz, and Drepanoclado uncinati-Heliospermetum pusilli from the Liburnian Karst (sensu Beck, 1901) in the Dinaric Mts. Nomenclature of the syntaxa follows Englisch et al. (1993), Theurillat et al. (1995), Englisch (1999), and Aeschimann et al. (2004b); the list is given in the Appendix. Plant names and chorological groups are in agreement with the Flora alpina (Aeschimann et al., 2004a) and the Register of the Flora of Slovenia (Trpin & Vres, 1995). Successive number 1 2 S 4 5 6 7 8 e 10 11 12 1S 14 15 16 17 18 1e Syntaxa Arabidion caeruleae s. lat. Soldanello-Salicion retusae No. of relevés 6 10 16 11 22 17 8 18 3 12 24 16 13 9 18 7 5 8 9 No. of species 26 SS Se 42 Se 47 41 72 22 54 58 88 86 65 e7 56 42 66 Se Characteristic and differential (d) species Arabidion caeruleae (ACs) Soldanella minima** 50 90 63 9 33 67 31 23 11 33 29 13 67 Saxifraga androsacea 17 60 13 9 55 94 50 22 33 92 15 33 44 20 38 Saxifraga sedoides 100 100 100 100 75 39 67 33 15 17 43 Gnaphalium hoppeanum 6 38 28 100 100 11 33 80 25 100 Hutchinsia alpina ssp. austroalpina* 83 90 75 64 75 33 11 14 11 Hutchinsia alpina ssp. brevicaulis 100 88 100 33 33 20 Potentilla brauneana 38 56 83 11 88 100 Thlaspi alpestre* 20 5 18 6 8 17 Arabis caerulea* 29 88 6 100 92 Saxifraga hohenwartii* 100 100 6 11 Rumex nivalis 18 63 89 Geographical differential species TR2 Achillea oxyloba* 75 100 83 8 33 43 AC2 Ranunculus traunfellneri** 83 40 6 64 63 72 25 62 44 38 TR2 Achillea clusiana* 5 29 AC3 Campanula pulla* 45 76 22 ES Viola calcarata ssp. calcarata** 100 SS Soldanella alpina 44 42 4 56 38 44 44 29 100 50 44 Soldanello alpinae-Salicion retusae (SS) Salix retusa 70 6 5 41 13 56 58 81 100 100 100 100 80 88 100 Salix reticulata 24 33 11 78 100 40 Tab. 1: Synoptic table of syntaxa from the order Arabidetalia caeruleae s. lat. in the South-eastern and Northeastern Alps. 1, 2 - Saxifragetum hohenwartii; 3 - Saxifragetum stellaro-sedoidis var. geogr. Achillea oxyloba; 4 -Saxifragetum stellaro-sedoidis var. geogr. Ranunculus traunfellneri; 5, 6 - Saxifragetum stellaro-sedoidis var. geogr. Campanula pulla; 7, 8 - Ranunculo traunfellneri-Festucetum nitidae; 9 - Poo supinae-Cerastietum cerastoidis; 10 -Salici herbaceae-Arabidetum coeruleae; 11 - Drepanoclado uncinati-Heliospermetum pusilli; 12 - Salici retusae-Geranietum argentei; 13, 14 - Homogyno discoloris-Salicetum retusae; 15-17 - Salicetum retuso-reticulatae; 18, 19 - Potentillo brauneanae-Homogynetum discoloris. Tab. 1: Sinoptična tabela sintaksonov iz reda Arabidetalia coeruleae s. lat. v jugovzhodnih in severovzhodnih Alpah. 1, 2 - Saxifragetum hohenwartii; 3 - Saxifragetum stellaro-sedoidis var. geogr. Achillea oxyloba; 4 - Saxifragetum stellaro-sedoidis var. geogr. Ranunculus traunfellneri; 5, 6 - Saxifragetum stellaro-sedoidis var. geogr. Campanula pulla; 7, 8 - Ranunculo traunfellneri-Festucetum nitidae; 9 - Poo supinae-Cerastietum cerastoidis; 10 - Salici herbaceae-Arabidetum coeruleae; 11 - Drepanoclado uncinati-Heliospermetum pusilli; 12 - Salici retusae-Geranietum argentei; 13, 14 - Homogyno discoloris-Salicetum retusae; 15-17 - Salicetum retuso-reticulatae; 18, 19 - Potentillo brauneanae-Homogynetum discoloris. Arabidetalia caeruleae (AG) Veronica aphylla 50 25 36 9 25 44 17 13 15 33 22 20 50 33 Arabis alpina ssp. alpina 83 80 63 82 95 35 38 22 17 8 11 Carex atrata ssp. atrata 9 6 8 6 23 56 6 29 80 Carex parviflora 9 65 25 11 6 38 61 29 88 Carex ornithopodoides 6 13 6 58 15 11 6 75 Festuca nitida 55 38 100 4 50 38 50 Ranunculus alpestris 9 50 82 8 61 100 60 Anemone baldensis 10 33 8 15 33 11 Galium noricum** 41 38 17 11 39 63 Soldanella austriaca* 50 71 28 Doronicum glaciale ssp. calcareum* 18 Festuca rupicaprina* 9 Thlaspion rotundifolii (TR3) Thlaspi rotundifolium ssp. rotundifolium* 56 9 5 50 33 Leontodon montanus ssp. montanus 19 5 25 11 Papaver aurantiacum 38 13 Papaver ernesti-mayeri 13 17 Alyssum ovirense 13 Minuartia austriaca** 13 Petasition paradoxi (PP) Silene quadrifida** 17 30 9 6 17 4 44 15 13 67 Adenostyles glabra 6 18 5 6 6 8 13 8 Polystichum lonchitis 13 11 8 Rumex scutatus 25 8 Aconitum angustifolium* 13 Athamanta cretensis 6 Cystopteris montana 8 Dryopteris villarii 4 Gypsophila repens 6 Thlaspietalia rotundifolii (TR¡) Achillea atrata** 50 90 19 9 9 65 100 94 6 15 78 22 40 38 67 Taraxacum alpinum agg. 17 40 31 18 88 83 100 75 6 23 11 28 57 25 78 Moehringia ciliata 83 70 88 36 45 82 88 83 33 42 6 15 11 17 Poa minor 75 18 53 11 11 17 43 11 Sedum atratum ssp. atratum 9 12 25 17 6 11 20 Cerastium carinthiacum ssp. cari nthiacum* 9 11 43 Doronicum grandiflorum 17 8 Trisetum argenteum 6 Androsacetalia alpinae (AA) Veronica alpina 50 9 23 24 75 83 100 75 11 17 100 75 44 Gentiana bavarica* 6 50 39 60 Oxyria digyna 36 13 Trifolium pallescens 94 13 Epilobium anagallidifolium 8 Geum reptans 6 Gymnocarpium dryopteris 33 Saxifraga seguieri* 6 Drabetalia hoppeanae (DH) Doronicum glaciale ssp. glaciale* 25 18 6 15 22 Sesleria ovata* 18 35 17 Thlaspietea rotundifolii (TR) Campanula cochleariifolia 33 60 6 27 5 25 50 8 25 31 11 22 29 13 Minuartia sedoides 6 6 28 8 44 15 17 14 100 Armeria alpina 5 24 13 22 13 8 33 14 Biscutella laevigata ssp. laevigata 12 17 8 8 22 17 13 Leontodon hispidus 6 25 8 17 20 Linaria alpina ssp. alpina 5 13 28 Saxífraga oppositifolia ssp. oppositi-folia 13 25 6 Rhodiola rosea 6 15 Salicetea herbaceae (SH) Sagina saginoides 5 24 38 11 100 58 11 17 50 56 Alchemilla fissa 50 15 11 17 6 56 Salix herbacea 100 67 14 80 Sibbaldia procumbens 13 14 20 33 Gnaphalium supinum 13 13 22 Cerastium cerastoides 100 8 Soldanella pusilla 11 13 Asplenietea trichomanis (AT) dAC2 Silene pusilla 6 9 45 24 100 11 dAC3 Arabis bellidifolia ssp. stellulata 17 10 38 18 5 59 6 50 44 22 13 dAC3 Valeriana elongata** 50 80 31 9 9 18 11 11 17 dAC3 Cystopteris regia 33 40 27 14 33 13 8 Phyteuma sieberi* 10 9 11 19 23 33 14 25 Saxifraga paniculata 9 12 13 6 6 8 6 Paederota lutea 6 17 13 15 11 Asplenium viride 6 50 19 15 Cystopteris fragilis 46 6 8 29 Saxifraga crustata 9 38 8 14 Cerastium subtriflorum** 6 38 8 Valeriana saxatilis 18 6 11 Petrocallis pyrenaica 11 13 Primula clusiana* 6 6 Elyno-Seslerietea (ES) dAC2 Aster bellidiastrum 33 40 6 18 9 6 25 67 33 50 21 81 69 78 67 43 75 33 dAC2 Silene acaulis 9 14 41 75 39 58 50 15 44 78 86 80 100 dSS Carex fi rma 6 5 47 50 33 33 4 38 31 78 61 14 38 dSS Campanula scheuchzeri 5 12 39 58 8 56 8 11 11 100 100 67 dSS Homogyne discolor 30 6 13 11 25 31 31 89 56 14 100 78 dSS Euphrasia salisburgensis 5 25 17 75 23 22 6 20 50 78 dSS Bartsia alpina 20 6 17 4 13 67 44 29 40 dSS Carex sempervirens 6 8 88 4 11 56 57 80 38 dSS Dryas octopetala 5 6 4 6 8 22 39 14 25 dSS Potentilla crantzii 11 4 63 23 11 17 43 63 dSS Ranunculus carinthiacus 38 38 33 11 14 38 67 dSS Homogyne alpina 19 15 17 29 100 dSS Salix serpillifolia 13 22 40 dSS Geranium argenteum 100 23 Poa alpina 50 100 38 100 73 71 100 100 83 42 69 54 78 72 57 20 100 89 Myosotis alpestris 50 45 5 18 50 78 58 8 56 15 33 61 86 80 38 Carex ferruginea ssp. ferruginea 6 33 17 13 6 22 17 100 25 11 Galium anisophyllum 10 6 56 8 29 77 56 6 38 78 Juncus monanthos 18 6 8 88 23 33 17 14 13 Sesleria caerulea 6 9 6 11 38 78 44 100 63 Cerastium carinthiacum ssp. austroalpinum* 33 50 6 27 50 89 6 4 Gentiana pumila* 12 13 17 8 67 17 100 33 Pedicularis verticillata 12 11 69 8 11 6 20 13 Festuca quadriflora 6 9 5 24 44 40 38 Arabis vochinensis 38 94 15 33 63 11 Gentiana anisodonta 13 75 15 11 38 56 Gentiana verna ssp. verna 6 25 22 28 14 38 Pedicularis rostratocapitata 6 17 6 31 11 17 Ranunculus montanus 5 11 50 44 43 100 Salix alpina 13 17 19 31 17 14 Achillea clavenae 17 63 24 17 14 Agrostis alpina 6 50 8 17 88 Festuca norica* 50 50 13 9 44 Saxifraga caesia 13 8 11 6 29 Thymus praecox ssp. polytrichus 22 38 15 6 33 Crepis aurea 22 6 38 78 Draba aizoides ssp. aizoides 10 25 17 40 Erigeron glabratus 4 63 8 6 Gentiana clusii 25 11 6 38 Ligusticum mutellina 12 75 11 100 Anthyllis vulneraria ssp. alpestris 8 13 22 Astrantia bavarica** 11 31 11 Carex ornithopoda 19 8 13 Cerastium arvense ssp. strictum 6 13 6 Elyna myosuroides 8 11 25 Festuca violacea agg. 67 22 100 Gentiana orbicularis 50 8 17 Gentiana terglouensis* 9 25 17 Helianthemum alpestre 75 8 13 Hieracium bifidum 8 8 14 Hieracium villosum 56 8 6 Phyteuma orbiculare 13 6 6 Ranunculus hybridus 6 8 11 Sesleria sphaerocephala* 19 13 13 Juncetea trifidi (JT) Potentilla aurea 33 14 80 13 78 Euphrasia minima 42 6 6 29 Luzula spicata ssp. spicata 11 11 20 38 Agrostis rupestris 8 11 13 Botrychium lunaria 14 13 Primula minima 8 28 Veronica bellidioides 6 14 Nardetea strictae (N) Coeloglossum viride 8 6 8 11 Luzula multiflora ssp. multiflora 6 8 Nardus stricta 14 22 Loiseleurio-Vaccinietea (LV) Rhodothamnus chamaecistus** 25 8 6 Vaccinium gaultherioides 13 6 38 Juniperus communis ssp. alpina 8 14 Montio-Cardaminetea (MC) dAC2 Saxifraga aizoides 67 40 19 55 9 12 22 17 19 23 44 17 14 13 dAC3 Saxifraga stellaris 31 9 91 88 75 44 100 8 8 11 Scheuchzerio-Caricetea fuscae (SC) Selaginella selaginoides 33 56 8 67 50 80 63 67 Carex capillaris 8 71 19 15 33 11 63 Parnassia palustris 67 88 46 11 14 13 Pinguicula alpina 13 22 17 Tofieldia calyculata 15 11 11 Molinio-Arrhenatheretea (M-Ar) Lotus corniculatus 6 8 31 6 29 Anthoxanthum odoratum 69 8 17 Trifolium pratense ssp. pratense 6 38 6 Trifolium repens ssp. repens 4 8 Trollius europaeus 4 11 Muljedio-Aconitetea (MA) dAC2 Viola biflora 17 60 27 9 44 8 88 13 38 43 38 78 Salix waldsteiniana 8 6 23 22 14 Veratrum album ssp. lobelianum 22 19 15 6 Aconitum lycoctonum ssp. neapolitanum 11 4 6 Adenostyles alliariae 6 29 Geranium sylvaticum 6 4 Salix appendiculata 71 8 Saxifraga rotundifolia L. 2O 4 Erico-Pinetea (EP) Rhododendron hirsutum** 38 19 8 14 Pinus mugo ssp. mugo 8 8 Rubus saxatilis 42 8 Querco-Fagetea (QF) Luzula sylvatica ó 8 ó 14 Daphne mezereum ó ó 8 Solidago virgaurea (incl. minuta) 4 ó Vaccinio-Piceetea (VP) Huperzia selago 21 ó 8 ó Vaccinium vitis-idaea 71 13 15 ó Picea abies ó 58 Other species Polygonum viviparum 9O 31 18 23 29 5O 5ó 83 ó3 88 85 89 94 8ó 1OO 1OO 78 RESULTS AND DISCUSSION An extensive analytical table was provided and analyses were performed applying hierarchical classification methods. Two distinct groups of relevés were recognised in the cluster analysis: stands from Arabidion s. lat. and Salicion s. lat. (Fig. 4). Within the "Arabidion" group, seven distinct clusters were recognised (Fig. 2): A - Saxifragetum hohenwartii (Thlaspion, Thlaspietalia) from the Karavanke Mts. (Tab. 1: columns 1, 2), B -Ranunculo traunfellneri-Festucetum nitidae from the Julian Alps (7, 8), C - Poo-Cerastietum (Salicetea herba-ceae; 9) and Salici herbaceae-Arabidetum caeruleae (Salici-Arabidion; 10) from the Dolomites of Lienz, D -Saxifragetum stellaro-sedoidis var. geogr. Achillea oxy-loba (3) from the Dolomites of Lienz, E -from the Julian Alps (var. geogr. Ranunculus traunfellneri; 4), F - from Hochschwab and Gesauseberge (5), and G from the Hochschwab only (F-G var. geogr. Campanula pulla; 6). The "Salicion" group consisted of nine clusters (Fig. 3): A - Drepanoclado-Heliospermetum from the Libur-nian Karst, B - Salici retusae-Geranietum argentei from the Julian Alps (12), C - Homogyno discoloris-Salicetum retusae from the Krn Mts. (Julian Alps) and Carnic Alps (13), and D - from the Karavanke Mts. and the Kamnik Alps (14), E - Salicetum retuso-reticulatae from the Dolomites of Lienz and Carnic Alps (15), F - from the Monte Baldo and Dolomites of Belluno (16) and G -from Bernina (Unter-Engadin, above Val Cluoza; 17), H - Potentillo brauneanae-Homogynetum discoloris from the Julian Alps (18) and I - from the Karavanke Mts. (19). Stands of Drepanoclado-Heliospermetum were grouped within the "Salicion" cluster, but several floristi-cal, phytogeographical as well as ecological distinctions were recognised. Due to the lower altitude, such stands were inhabited by many forest (Querco-Fagetea, Vac-cinio-Piceetea, Erico-Pinetea) and tall-herb (Mulgedio-Aconitetea) species (Tabs. 1-4). The presence of Illyrian species in stands, such as Gentiana liburnica, Polygala croatica and Thymus balcanus (Tab. 1) as well as Arabis scopoliana, Scabiosa silenifolia, Carex kitaibeliana etc. (Seslerion juncifoliae, Seslerietalia juncifoliae, Elyno-Seslerietea) were of phytogeographical importance (Surina & Vres, 2004). In comparison to other syntaxa, Alpine geoelements in the broadest sense occurred more sporadically, while Eurosiberian and Eurosiberian/N-American geoelements prevailed (Tab. 2). Secondly, a synoptic table (Tab. 1) was provided with the aid of cluster analysis (Fig. 4) and on the basis of previous results obtained within both groups of relevés. Distinction between the Arabidion and the Soldan-ello-Salicion were evident and delimitation of Arabidion according to Englisch (1999) into at least two alliances was not questionable (Tabs. 1-4). Scree stability, aspect and talus slope may be the three most important environmental factors that determine the floristic composition and differentiation of the alliances: Arabidion prefers shady and slightly less stabilised scree with no or very little accumulation of soils and perhaps steeper slopes in comparison to the Soldanello-Salicion. The coverage indices and number of species of the Thlaspion and the Androsacetalia alpinae, which prefer less stabilised screes, were also much higher. The Poo-Cerastietum and Salici-Arabidetum were grouped together in the cluster analysis, but within the "Arabidion" cluster (Fig. 2: cluster C; Fig. 4: clusters 9, 10). The later syntaxon was placed into the Salici-Arabidion (Englisch, 1999), which was separated from Salicion herbaceae due to floristical and ecological differences derived from the fact that the newly proposed alliance embraces the snow-bed vegetation on humus rich calcareous screes. The number and coverage indices of acidophilous and hygrophilous species of Salicetea herbaceae, Androsa-cetalia alpinae, -uncetea trifidi, Nardetea, Vaccinio-Piceetea and Scheuchzerio-Caricetea fuscae (Tabs. 1, 3, 4) proved the unique syntaxonomic position of Salici-Arabidetum (between Salicion herbaceae and Arabidion caeruleae) and the aforementioned classification of syn-taxa. Fig. 2: Dendrogram based on species frequency in the syntaxa of Arabidetalia caeruleae s. lat. in the Southeastern and North-eastern Calcareous Alps. A - Saxifra-getum hohenwartii; B - Ranunculo traunfellneri-Festu-cetum nitidae; C - Poo supinae-Cerastietum cerastoidis & Salici herbaceae-Arabidetum caeruleae; D - Saxifra-getum stellaro-sedoidis var. geogr. Achillea oxyloba; E -var. geogr. Ranunculus traunfellneri; F-G - var. geogr. Campanula pulla. Sl. 2: Dendrogram, izdelan na osnovi pogostosti vrst v sintaksonu Arabidetalia caeruleae s. lat. v jugovzhodnih in severovzhodnih Apneniških Alpah. A - Saxifragetum hohenwartii; B - Ranunculo traunfellneri-Festucetum nitidae; C - Poo supinae-Cerastietum cerastoidis & Salici herbaceae-Arabidetum caeruleae; D - Saxifragetum stellaro-sedoidis var. geogr. Achillea oxyloba; E - var. geogr. Ranunculus traunfellneri; F-G - var. geogr. Campanula pulla. Fig. 3: Dendrogram based on species frequency in the syntaxa of Soldanello alpinae-Salicion retusae s. lat. in the South-eastern and North-eastern Calcareous Alps. A - Drepanoclado uncinati-Heliospermetum pusilli; B -Salici retusae-Geranietum argentei; C, D - Homogyno discoloris-Salicetum retusae; E-G - Salicetum retuso-reticulatae; H-l - Potentillo brauneanae-Homogynetum discoloris. Sl. 3: Dendrogram, izdelan na osnovi pogostosti vrst v sintaksonu Soldanello alpinae-Salicion retusae s. lat. v jugovzhodnih in severovzhodnih Apneniških Alpah. A - Drepanoclado uncinati-Heliospermetum pusilli; B -Salici retusae-Geranietum argentei; C, D - Homogyno discoloris-Salicetum retusae; E-G - Salicetum retuso-reticulatae; H-l - Potentillo brauneanae-Homogynetum discoloris. Tab. 2: Chorological groups (geoelements) of syntaxa of Arabidetalia caeruleae in the SE Alps (* endemic, ** suben-demic species). Tab. 2: Horološke skupine (geoelementi) sintaksonov Arabidetalia caeruleae v JV Alpah (* endemične, ** suben-demične vrste). Geoelement / No. of spec. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 S-Eur.-Mont. 3 8 12 11 16 18 14 31 3 26 23 30 31 27 27 18 45 23 18 Arct./Alp. 3 6 6 8 9 9 19 12 5 12 7 13 12 13 14 6 21 10 10 SE-Europ.-Mont. 1 1 4 3 3 5 4 6 2 5 8 10 8 7 6 2 9 3 3 E-Alp.** 4 4 5 4 2 4 3 9 1 2 9 6 6 4 2 6 2 1 Eurosib./N-Am. 1 1 2 1 1 1 2 2 5 13 8 8 4 4 2 6 3 2 E-Alp.* 4 6 6 5 7 10 5 6 1 1 7 6 5 3 2 14 4 Euroasiat. 1 1 1 1 1 1 1 1 2 1 2 1 2 1 2 6 1 E-Alp./Illyr. 1 1 2 1 4 7 1 3 7 7 4 2 2 4 5 Europ. 1 1 2 2 1 2 4 6 8 5 2 2 6 2 2 Alp.* 1 1 2 2 2 1 3 3 1 1 1 3 1 1 E-Alp./Apen. 1 2 1 1 1 1 1 1 1 1 1 1 1 1 Europ-/N-Am. 2 2 1 1 1 3 3 1 1 2 1 Alp. 1 2 2 3 3 1 2 3 1 1 Europ./Mont. 1 1 3 1 1 3 1 3 1 1 Other 1 4 5 2 3 3 3 1 1 N E-Alp./Illyr.** 1 1 1 1 2 1 1 1 1 N-Europ.-Alp. 1 1 1 1 1 1 1 1 1 Eurosib. 4 10 3 4 1 1 3 1 Alp./Apen. 1 1 1 1 1 1 1 1 Cosmop. 2 1 1 1 1 2 E-Alp/Carpat 1 2 1 1 1 1 Al p./Illyr. 1 1 1 1 1 Eurasiat./Afr. 1 1 1 1 1 Europ./W-Asiat. 1 2 2 2 Euroasiat./N-Am. 1 3 1 1 Eurasiat./Am. 1 1 1 1 S-Europ. 1 1 1 1 W-Alp.* 1 1 1 Accumulation of soil and formation of tiny horizons, slow decomposition of dead plant material, and subsequent acidification of soils are well suited to species of the Juncetea trifidi, Nardetea, Loiseleurio-Vaccinietea, Scheuchzerio-Caricetea fuscae, and Vaccinio-Piceetea (Tabs. 3, 4), which were significantly more abundant and achieved higher coverage indices in syntaxa of the Soldanello-Salicion. Associations of Soldanello-Salicion, restricted mainly to shallow depressions with long lasting snow, were less frigiphilous (Braun-Blanquet, 1926; Englisch et al., 1993) and were species richer, which was mainly due to the occurrence of the Elyno-Seslerietea species (Fig. 5). They also achieved relatively high coverage indices. The high share of species of subalpine and alpine calcareous grasslands in the Salicetum retuso-reticulatae was already mentioned by Pignatti & Pignatti (1983), while the Diantho alpinae-Salicetum retusae, the Homogyno-Salicetum and the Salici-Geranietum are transitional to the Elyno-Seslerietea (mainly the Caricion firmae; see Englisch, 1999; Surina, 2004c) and play an important role in the subsequent succession of calcareous scree vegetation towards subalpine and alpine calcareous grasslands. Our study revealed phytogeographical differentiation of stands of Saxifragetum stellaro-sedoidis (Englisch et al., 1993; Englisch, 1999) and several geographical variants or races based on the floristical peculiarities of the area could be distinguished (Tab. 1, Figs. 2, 4). This was mainly owing to disjunctions of the distribution area of closely related endemic taxa (Tabs. 1, 2), which vicariate in a SE-NE direction (NE-Calcareous Alps - SE-Calcareous Alps), e.g. Saxifraga sedoides - S. hohenwartii, Soldanella austriaca - S. minima, Ranunculus alpestris - R. traunfellneri, Cerastium carinthiacum subsp. carinthiacum - C. c. subsp. austroalpinum etc. (see Hörandl, 1993). As geographical differential species we chose Achillea clusiana, Soldanella austriaca and Campanula pulla for the NE Calcareous Alps, Ranuncu- 6 5 fr 4 'Ë Fig. 4: Dendrogram based on species frequency in the syntaxa of Arabidetalia caeruleae s. lat. in the South-eastern and North-eastern Calcareous Alps. 1, 2 - Saxifragetum hohenwartii; 3 - Saxifragetum stellaro-sedoidis var. geogr. Achillea oxyloba; 4 - Saxifragetum stellaro-sedoidis var. geogr. Ranunculus traunfellneri; 5, 6 - Saxifragetum stel-laro-sedoidis var. geogr. Campanula pulla; 7, 8 - Ranunculo traunfellneri-Festucetum nitidae; 9 - Poo supinae-Cerastietum cerastoidis; 10 - Salici herbaceae-Arabidetum coeruleae; 11 - Drepanoclado uncinati-Heliospermetum pusilli; 12 - Salici retusae-Geranietum argentei; 13, 14 - Homogyno discoloris-Salicetum retusae; 15-17 - Salicetum retuso-reticulatae; 18, 19 - Potentillo brauneanae-Homogynetum discoloris (numbers correspond with Table 1). Sl. 4: Dendrogram, izdelan na osnovi pogostosti vrst v sintaksonu Arabidetalia caerule s. lat. v jugovzhodnih in severovzhodnih Apneniških Alpah. 1, 2 - Saxifragetum hohenwartii; 3 - Saxifragetum stellaro-sedoidis var. geogr. Achillea oxyloba; 4 - Saxifragetum stellaro-sedoidis var. geogr. Ranunculus traunfellneri; 5, 6 - Saxifragetum stel-laro-sedoidis var. geogr. Campanula pulla; 7, 8 - Ranunculo traunfellneri-Festucetum nitidae; 9 - Poo supinae-Cerastietum cerastoidis; 10 - Salici herbaceae-Arabidetum coeruleae; 11 - Drepanoclado uncinati-Heliospermetum pusilli; 12 - Salici retusae-Geranietum argentei; 13, 14 - Homogyno discoloris-Salicetum retusae; 15-17 - Salicetum retuso-reticulatae; 18, 19 - Potentillo brauneanae-Homogynetum discoloris (številke se ujemajo s tistimi v Tabeli 1). lus traunfellneri for the SE Calcareous Alps and Achillea oxyloba for Mt. Baldo, the Dolomites of Lienz and Belluno, and the Carnic Alps. It has recently been established that Saxifraga sedoides from the Dolomites of Lienz was wrongly identified as S. hohenwartii by Wikus (Wikus, 1959: Tab. 7), since the species is restricted to the Karavanke Mts. and the Kamnik Alps. We thus treated selective stands (Dirnböck et al., 1999: Tab. 12, rel. 8) as the Saxifragetum stellaro-sedoidis var. geogr. Achillea oxyloba. Lectotypes: - Saxifragetum stellaro-sedoidis Englisch 1999 var. geogr. Campanula pulla var. geogr. nova: Dirnböck et al. (1999: Tab. 12, rel. 8) - Hochschwab, lectoty-pus hoc loco. - Saxifragetum stellaro-sedoidis Englisch 1999 var. geogr. Ranunculus traunfellneri var. geogr. nova: Surina (2004c: Tab. 1, rel. 2) - Julian Alps, lectotypus hoc loco. - Saxifragetum stellaro-sedoidis Englisch 1999 var. geogr. Achillea oxyloba var. geogr. nova: Wikus (1959: Tab. 7, rel. 5) - Dolomites of Lienz, lectoty-pus hoc loco. Stands of the Saxifragetum hohenwartii, in line with recent syntaxonomical literature on calcareous scree vegetation (Englisch et al., 1993; Englisch, 1999), were placed into Thlaspion rotundifolii as earlier proposed by Wikus (1959), Zollitsch (1967) and Wraber (1970). Although stands with predominating Saxifraga hohenwartii were transitional between the Thlaspion and the Arabi-dion, our views on the syntaxonomy of the Saxifragetum hohenwartii was in agreement with the author of the unit (Aichinger, 1933), who placed this association into the Arabidion. We considered those associations to be a phytogeographical vicariant (in the Karavanke Mts. and Kamnik Alps) to the Saxifragetum stellaro-sedoidis s. lat. Within the Soldanello-Salicion (Fig. 3), four associations were clearly identified, such as the Salici-Gera-nietum, the Homogyno-Salicetum, the Salicetum retuso-reticulatae, and the Potentillo-Homogynetum. Floristical distinctions between the Homogyno-Salicetum and the Salicetum retuso-reticulatae were not only due to the presence or absence of Salix reticulata, Carex ferruginea and Homogyne discolor (e.g. Englisch, 1999), but also owing to the significantly higher coverage indices and number of acidophilous species of the Androsacetalia al- pinae, the Salicetea herbaceae and the Juncetea trifidi (Tabs. 1, 3, 4). Stands with predominating Salix retusa showed certain phytogeographical and ecological differentiation. Several vicariant syntaxa were therefore described: the Homogyno-Salicetum and the Salici-Geranietum for the SE Calcareous Alps, and the Diantho-Salicetum and the Selaginello selaginoidis-Salicetum re-ticulatae for the NE Calcareous Alps. In contrast to stands of the Homogyno-Salicetum and the Salicetum retuso-reticulatae, stands of the Salici-Geranietum and the Di-antho-Salicetum were transitional to stands of alpine calcareous grasslands (the Caricion firmae), and stands of Potentillo-Homogynetum transitional to stands of Loise-leurio-Vaccinietea. Several succession stages of Salicetum retusae s. lat. were observed: in the SE Calcareous Alps towards stands of Gentiano terglouensis-Caricetum firmae and Dryadetum octopetalae s. lat., and in the NE Calcareous Alps towards stands of Caricetum firmae (Wendelberger, 1971; Englisch et al., 1993; Englisch, 1999; Surina, 2004b). For a more detailed description of the ecology, chorology and syntaxonomy of snow-bed vegetation, see previously cited references. Fig. 5: Coverage index (D%) and number of alpine and subalpine calcareous grasslands species (Elyno-Sesle-rietea) in syntaxa of Arabidetalia caeruleae in SE Alps. Sl. 5: Indeks pokrivnosti (D%) ter število alpinskih in subalpinskih apneniških traviščnih vrst (Elyno-Seslerie-tea) v sintaksonih Arabidetalia caeruleae v ¡V Alpah. Tab. 3: Phytosociological groups of syntaxa of Arabidetalia caeruleae in the SE Alps. Tab. 3: Fitosociološke skupine sintaksonov Arabidetalia caeruleae v ¡V Alpah. Syntaxa / No. of spec. 1 2 3 4 5 6 7 8 9 10 Me 11 12 13 14 15 16 17 18 19 Me Arabidion caeruleae 5 6 6 7 6 8 7 9 3 7 6.5 3 5 8 7 3 9 3 3 4.0 Soldanello-Salicion 1 1 1 2 1 2 3 1.0 1 2 2 3 2 2 3 3 3 2.5 Arabidetalia caeruleae 1 2 3 6 3 5 5 6 1 4 3.5 2 4 6 5 4 2 7 3 3 4.0 Thlaspion rotundifolii 4 1 3 4 2 1 1.0 1 0.0 Petasition paradoxi 1 1 2 2 1 2 1 3 1.0 5 5 3 1 1 0.5 Thlaspietalia rotundifolii 3 3 6 2 6 6 4 6 3 3 3.5 6 5 5 2 3 6 4 2 4.5 Androsacetalia alpinae 1 2 1 2 2 1 1 3 1.0 1 1 1 2 1 4 2 1.0 Drabetalia hoppeanae 2 1 1 2 1 1 1.0 2 2 1 3 1 1.0 Thlaspietea rotundifolii 2 2 3 2 4 3 5 6 1 2.5 2 4 6 3 2 5 2 1 2.5 Salicetea herbaceae 1 1 3 1 3 4 1.0 1 2 3 4 4 2 3 2.5 Asplenietea trichomanis 2 3 4 7 3 5 2 8 1 1 3.0 6 12 8 5 3 4 3 3.5 Elyno-Seslerietea 3 8 11 11 11 2 14 37 1 21 11.0 18 47 37 32 32 15 53 23 19 32.0 Vaccinio-Piceetea 1 1 1 0.0 1 3 4 3 1 1 1.0 Scheuczerio-Caricetea fuscae 3 0.0 3 5 4 5 3 1 6 1 1 3.5 Juncetea trifidi 2 0.0 1 2 6 1 11 4 2 2.0 Nardetea 1 0.0 2 4 1 1 1 1.0 Loiseleurio-Vaccinietea 0.0 1 3 1 1 3 1 1.0 Molinio-Arrhenatheretea 1 1 1 1 1 1 1 4 1 2 1.0 5 7 3 1 1 2 6 1 2.0 Montio-Cardaminetea 1 1 2 2 2 2 1 2 1 2 2.0 1 1 2 1 1 2 1 1.0 Mulgedio-Aconitetea 1 2 1 1 8 1 1.0 7 4 4 1 1 1 2 1.5 Erico-Pinetea 0.0 6 1 3 2 1 0.5 Querco-Fagetea 3 0.0 20 3 2 2 1 0.5 Artemisietea vulgaris 1 1 1 0.0 1 1 0.0 Festuco-Brometea 1 0.0 1 1 1 0.0 Trifolio-Geranietea 0.0 1 1 0.0 Seslerion juncifoliae 0.0 3 0.0 Epilobietea angustifolii 0.0 2 0.0 Other species 1 4 0.0 5 2 3 3 3 1 1 1.5 CONCLUSIONS 1. Delimitation among the alliances Arabidion, Salici-Arabidion and Soldanello-Salicion, based on flo-ristic composition and the phytogeographical peculiarities of the area as well as the specific ecology of stands, was distinct and several phytogeographical and/or ecological vicariant syntaxa were recognised. Within the Arabidion, three associations were recognised in the study area, such as the Ranunculo-Festucetum (an endemic association in the Julian Alps), the Saxifragetum hohenwartii (restricted to the Karavanke Mts. and the Kamnik Alps), and the Saxifragetum stellaro-sedoidis (distributed throughout the Eastern Alps). The latter was subdivided into three geographical variants, such as variants of Achillea oxyloba (western part of the SE Alps: Monte Baldo, Dolomites, Carnic Alps), of Ranunculus traunfellneri (mainly SE Calcareous Alps), and of Campanula pulla (NE Alps). 2. The Salici-Arabidetum was placed into the Salici-Arabidion and our analysis partly confirmed its unique syntaxonomic position between the Arabidion and the Salicion herbaceae. This association is distributed in the Northern, Central and Southern Alps. 3. Within the Soldanello-Salicion, four associations were clearly recognised, such as the Homogyno-Salicetum, the Salicetum retuso-reticulatae, the Salici-Geranietum, and the Potentillo-Homogynetum. The distribution area of the Salicetum retuso-reticulatae is restricted mainly to the NE, SE and Central Alps, and is replaced by the Homogyno-Salicetum in the SE Calcareous Alps. To date, stands of the Salici-Geranietum have only been documented in the Julian Alps and constitute somewhat transitional unit of the Elyno-Seslerietea (Caricion firmae). 4. Our study confirmed the placement of the Drepa-noclado-Heliospermetum from the Liburnian Karst into the Dinaric alliance Salicion retusae. Since no synoptic treatment of Dinaric syntaxa has yet been provided, syntaxonomical and geographical delimitations of the Soldanello-Salicion and the Salicion retusae have remained unresolved. Nevertheless, stands from the Trnovski gozd plateau (NW Dinaric Mts., Slovenia) resembled those of Mt. Snežnik and are probably situated in the north-westernmost part of the distribution area of the Salicion retusae. Syntaxa / D% 1 2 3 4 5 6 7 8 9 10 Me 11 12 13 14 15 16 17 18 19 Me Arabidion caeruleae 33.78 27.59 27.69 27.56 16.93 34.45 19.28 15.66 2.00 19.17 23.42 1.57 7.94 6.70 8.33 17.76 1.29 1.36 5.59 6.14 Soldanello-Salicion 3.24 0.40 0.27 2.94 0.53 4.59 6.35 0.46 0.18 7.28 16.99 14.67 5.58 7.19 15.45 19.58 13.47 14.07 Arabidetalia caeruleae 9.00 7.29 7.38 13.35 15.49 13.95 8.43 11.54 8.1 8 4.47 8.71 0.67 2.21 7.96 6.64 8.00 1.26 5.96 3.28 4.46 5.21 Thlaspion rotundifolii 9.56 0.85 1.64 4.32 1.30 1.15 1.00 0.79 0 Petasition paradoxi 1.32 1.37 4.58 1.75 0.27 0.45 0.53 0.97 0.75 1.78 2.63 1.37 0.28 2.72 0.14 Thlaspietalia rotundifolii 12.58 12.13 22.80 3.50 8.98 14.28 18.52 11.74 13.64 9.23 12.36 1.76 3.42 4.39 1.76 6.98 4.98 1.36 2.23 2.82 Drabetalia hoppeanae 2.19 2.56 1.92 2.95 0.72 0.43 0.58 1.58 1.54 2.41 1.66 0.60 1.07 Androsacetalia alpinae 2.29 3.50 1.64 1.25 4.52 2.59 1.82 5.19 2.05 0.98 3.30 0.35 2.26 1.88 1.98 5.39 1.93 Thlaspietea rotundifolii 9.93 8.89 5.78 8.87 9.64 3.24 5.68 3.81 0.29 5.73 1.36 1.87 3.43 1.76 0.56 3.65 1.79 0.67 1.77 Salicetea herbaceae 0.42 1.13 3.54 0.36 28.75 1.43 0.39 0.75 0.73 2.68 8.37 2.27 1.19 5.59 1.73 Asplenietea trichomanis 9.93 8.47 3.79 6.68 5.48 4.38 5.30 2.59 6.36 4.32 5.39 7.67 4.62 4.29 3.17 1.13 1.34 2.39 1.86 Elyno-Seslerietea 11.29 2.73 9.37 14.65 6.98 14.49 23.88 3.72 2.82 27.55 10.33 15.88 56.13 38.66 51.54 48.78 42.26 38.19 35.44 54.55 45.52 Vaccinio-Piceetea 0.60 0.22 0.29 0 9.50 1.15 3.13 0.95 1.19 3.37 1.05 Scheuczerio-Caricetea fuscae 5.98 0 5.42 5.74 3.59 5.32 3.53 2.72 2.92 0.60 2.69 3.22 Juncetea trifidi 1.87 0 0.38 0.73 2.82 3.35 4.82 2.99 3.37 2.91 Nardetea 0.29 0 0.29 1.58 0.84 0.32 0.60 0.30 /oiseleurio-Vaccinietea 0 0.24 1.81 0.38 0.99 0.64 0.60 0.49 Molinio-Arrhenatheretea 3.97 5.26 2.59 8.87 6.16 3.29 6.32 4.64 13.51 3.75 4.95 3.73 7.00 3.52 3.34 4.66 5.23 3.78 4.18 0.67 3.98 Montio-Cardaminetea 7.95 2.59 4.38 7.12 8.43 5.90 3.77 1.80 7.27 0.86 5.14 3.47 0.43 1.30 1.58 0.28 0.79 0.60 0.52 Mulgedio-Aconitetea 1.32 3.89 2.85 0.55 3.88 0.72 0.64 9.15 1.86 4.97 1.23 1.41 3.14 1.29 4.18 1.63 Erico-Pinetea 0 5.22 0.52 1.22 1.23 0.60 0.26 Querco-Fagetea 0.54 0 7.92 0.87 0.75 0.39 1.00 0.20 Festuco-Brometea 0.23 0 0.29 0.38 0.42 0 Trifolio-Geran ietea 0 0.43 0.1 6 0 Artemisietea vulgaris 1.53 4.89 0.43 0 0.28 0.32 0 Seslerion juncifoliae 0 0.36 0 Epilobietea angustifolii 0 0.82 0 Other species 0.53 1.44 0 13.37 3.12 1.13 5.96 0.47 1.19 0.67 0.90 Tab. 4: Index of coverage (D%) of phytosociological groups of syntaxa of Arabidetalia caeruleae in the SE Alps. Tab. 4: Indeks pokrivnosti (D%) fitosocioloških skupin sintaksonov Arabidetalia caeruleae v JV Alpah. ACKNOWLEDGEMENTS My sincere thanks go to Dr. Igor Dakskobler (Institute of Biology, Scientific and Research Centre of the Slovenian Academy of Sciences and Arts), and Prof. Tone Wraber (University of Ljubljana, Biotechnical Faculty, Department of Biology) for consultations and valuable suggestions. The work was carried out at the Institute of Biology, Scientific and Research Centre of the Slovenian Academy of Sciences and Arts. FITOGEOGRAFSKA IN SINTAKSONOMSKA ANALIZA VEGETACIJE SNEŽNIH DOLINIC NA KARBONATNI PODLAGI V JUGOVZHODNIH ALPAH: NUMERIČNI PRISTOP Boštjan SURINA Univerza na Primorskem, Znanstveno-raziskovalno središče Koper, SI-6000 Koper, Garibaldijeva 1 E-mail: Bostjan.surina@zrs-kp.si POVZETEK Avtor v prispevku s pomočjo klastrske analize in opirajoč se na fitogeografske posebnosti območja podaja pregled vegetacije snežnih dolinic na karbonatni podlagi v Jugovzhodnih Alpah (Arabidetalia caeruleae, Thlaspietea rotundifolii). Obravnava osem asociacij, ki se dobro razlikujejo ekološko in fitogeografsko, te pa združuje v tri zveze. Asociacije Saxifragetum stellaro-sedoidis, Saxifragetum hohenwartii in Ranunculo traunfellneri-Festucetum nitidae pripadajo zvezi Arabidion caeruleae, asociacija Salici herbaceae-Arabidetum caeruleae, ki je v sintaksonomskem oziroma ekološkem oziru sorodna s sintaksoni snežnih dolinic na silikatni podlagi iz razreda Salicetea herbaceae, pa zvezi Salici herbaceae-Arabidion caeruleae. Asociacija Saxifragetum stellaro-sedoidis se fitogeografsko dobro diferencira v tri nove geografske variante, in sicer Achillea oxyloba v Lienških Dolomitih, Ranunculus traunfellneri v Jugovzhodnih Apneniških Alpah ter Campanula pulla v Severovzhodnih Apneniških Alpah. Analiza je nadalje potrdila smiselnost obravnavanja sestojev, v katerih prevladujeta vrbi Salix retusa in S. reticu-lata v okviru samostojne zveze Soldanello alpinae-Salicion retusae. Na območju Jugovzhodnih Alp pripadajo tej zvezi štiri asociacije: Salicetum retuso-reticulatae, Homogyno discoloris-Salicetum retusae, Salici retusae-Geranietum argentei in Potentillo brauneanae-Homogynetum discoloris. Asociacija Drepanoclado uncinati-Heliospermetum pusilli, ki jo poznamo z Liburnijskega krasa (severozahodni Dinaridi), se floristično, fitogeografsko in ekološko zelo dobro loči od sintaksonov vzhodno-alpske zveze Soldanello-Salicion retusae in jih uvrščamo v dinarsko zvezo Salicion retusae. Ključne besede: biogeografija, sintaksonomija, endemiti, klastrska analiza, vegetacija snežnih dolinic, Arabidetalia caeruleae, Thlaspietea rotundifolii REFERENCES Aeschimann, D., K. Lauber, D. M. Mosser & J.-P. Theu-rillat (2004a): Flora alpina. Band 1-3. Haupt Verlag, Bern, Stuttgart, Wien. Aeschimann, D., K. Lauber, D. M. Mosser & J.-P. Theu-rillat (2004b): Flora alpina. Band 3. Register. Haupt Verlag, Berlin, Stuttgart, Wien, 323 pp. Aichinger, E. 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Slovenian National Commission for Unesco & National Institute of Chemistry, Ljubljana, p. 53-58. Zollitsch, B. (1967): Soziologische und ökologische Untersuchungen auf Kalkschiefern in hochalpinen Gebieten. Teil I. Die Steinschuttgesellschaften der Alpen unter besonderer Berücksichtigung der Gesellschaften auf Kalkschiefern in den mittleren und öslichen Zentralalpen. Ber.Bayer.Bot.Ges., 40, 67-100. APPENDIX List of relevés considered in the analysis (Tab. 1 ) 1 - Saxifragetum hohenwartii, Aichinger (1933): Tab. 9, relevés 1, 4, 11, 14-16; 2 - Saxifragetum hohenwartii, Aichinger (1933): Tab. 9, relevé 2, 3, 5-10, 12, 13; 3 -Saxifragetum stellaro-sedoidis var. geogr. Achillea oxy-loba, Wikus (1959): Tab. 7, relevé 1-12, 14 (sub nom. Saxifragetum hohenwartii), Englisch (1999): Tab. G, relevé 45; Hörandl (1993) in Englisch (1999): Tab. G.: 35, 36, 4 - Saxifragetum stellaro-sedoidis var. geogr. Ranunculus traunfellneri, Wraber (1972): Tab. 9, relevés 1 (sub nom. Saxifrago sedoidi-Arabidetum caeruleae), Surina (2004b): Tab. 9, Wikus (1959): Tab. 7, relevé 13 (sub nom. Saxifragetum hohenwartii), Englisch (1999): Tab. G, relevés 22, 38; 5 - Saxifragetum stellaro-sedo-idis var. geogr. Campanula pulla, Dirnböck et al. (1999): Tab. 12, relevés 12, 19, 20, Englisch (1999): Tab. G, relevés 20, 21, 25-34, 39-42; 6 - Saxifragetum stellaro-sedoidis var. geogr. Campanula pulla, Dirnböck et al. (1999): Tab. 12, relevés 8-11, 13-17, Englisch (1999): Tab. G, relevé 13, 14, 15, 16, 17, 18, 19, 23; 7 -Ranunculo traunfellneri-Festucetum nitidae, Wraber (1972): Tab. 9, relevés 3, 4, 6-9, 11, 12 (sub nom. Saxifrago sedoidi-Arabidetum caeruleae); 8 - Ranunculo traunfellneri-Festucetum nitidae, Wraber (1972): Tab. 9, relevés 5, 10, 13 sub nom. Saxifrago-Arabidetum, Tab. 10 (sub nom. Festuco violaceae-Rumicetum nivalis); 9 -Poo supinae-Cerastietum cerastoidis, Wikus (1959): Tab. 10, relevés 1-3 (sub nom. Arabidetum caeruleae poeto-sum supinae); 10 - Salici herbaceae-Arabidetum caeruleae, Wikus (1959): Tab. 10, relevés 4-15 (sub nom. Arabidetum caeruleae); 11 - Drepanoclado uncinati-Heliospermetum pusilli, Surina & Vres (2004): Tab. 1; 12 - Salici retusae-Geranietum argentei, Surina (2004c): Tab. 11; 13 - Homogyno discoloris-Salicetum retusae, Surina (2004c): Tab. 12, Poldini & Martini (1993): Tab. 1, relevés 1, 2, 7 (sub nom. Salicetum retuso-reticulatae), Aichinger (1933): Tab. 10, relevé 6; 14 -Homogyno discoloris-Salicetum retusae, Aichinger (1933): Tab. 10, relevés 1-5, Haderlapp (1982): Tab. 1, relevés 62-64; 15 - Salicetum retuso-reticulatae, Gerdol & Piccoli (1982): Tab. 4, relevé 4, Poldini & Martini (1993): Tab. 1, relevés 3-6, 8-13, Dirnböck et al. (1999): Tab. 12, relevés 1-7; 16 - Salicetum retuso-reticulatae, Gerdol & Piccoli (1982): Tab. 4, relevés 1-3, 5, Lasen (1983): relevé 19-21; 17 - Salicetum retuso-reticulatae, Braun-Blanquet (1926): Tab. 4. 18 - Poten-tillo brauneanae-Homogynetum discoloris, Wraber (1972): Tab. 10; 19 - Potentillo brauneanae-Homo-gynetum discoloris, Aichinger (1933): Tab. 12. Species with lower frequencies in the Table 1 (*/** endemic and subendemic species in the SE Alps) Artemisietea - Cirsium spinosissimum 25 (7), 89 (8), 8 (10), 13 (15), 11 (17); AT - Campanula cespitosa** 6 (8), C. justiniana 25 (11), Moehringia muscosa 8 (11), Festuca stenantha 13 (12), Saxifraga hostii / hostii** 19 (12), S. squarrosa** 19 (12), Potentilla nitida 13 (15); ES - Anemone narcissiflora 31 (12), 5 (15), Arenaria ciliata 8 (13), Carex mucronata 6 (2), 4 (11), Chamorchis alpina 8 (13), 6 (17), Erigeron uniflorus 8 (13), 60 (19), Festuca brachystachys ssp. brachystachys** 12 (6), 6 (17), Festuca melanopsis* 6 (17), 43 (18), Galium baldense* 6 (17)m 57 (18), Gentiana nivalis 31 (12), 31 (13), He-lianthemum nummularium ssp. grandiflorum 6 (12), 6 (17), Koeleria eriostachya 33 (11), 50 (12), Leontopodium alpinum 19 (12), 29 (18), Luzula glabrata* 6 (6), 6 (17), Minuartia verna ssp. verna 17 (8), 38 (15), Oxytro-pis jacquinii 13 (12), 13 (15), Pedicularis rosea ssp. rosea** 6 (8), 6 (17), Pimpinella alpina 38 (12), 8 (13), Polygala alpestris 8 (10), 15 (13), Stachys alopecuros / alopecuros 6 (8), 13 (12), Trifolium thalii 22 (16), 29 (18), Androsace chamaejasme 11 (17), Aster alpinus 13 (12), Callianthemum coriandrifolium 100 (19), Carduus defloratus agg. 6 (6), Dianthus alpinus* 6 (17), Festuca calva* 6 (13), Gentiana froelichii ssp. froelichii* 11 (14), G. liburnica* 4 (11), Globularia nudicaulis 4 (11), He-dysarum hedysaroides ssp. hedysaroides 6 (17), Helicto-trichon parlatorei* 13 (12), Heracleum austriacum ssp. siifolium* 6 (8), Horminum pyrenaicum 17 (17), Laser-pitium peucedanoides 6 (8), Leucanthemum atratum ssp. atratum* 6 (17), L. a. ssp. lithopolitanicum* 11 (14), Li-num alpinum 8 (13), Lloydia serotina 25 (15), Plantago atrata ssp. atrata 8 (10), Polygala croatica 1 (11), Pulsatilla alpina ssp. alpina 6 (17), Rhinanthus glacialis 19 (12), Saussurea pygmea 6 (8), Saxifraga exarata ssp. atropurpurea* 19 (12), S. e. ssp. moschata 5 (5), Scabiosa lucida ssp. lucida 6 (17), Thymus balcanus* 8 (11), Viola calcarata ssp. calcarata** 100 (19);EP -Calamagrostis varia 33 (11), Cirsium erisithales 4 (11), Erica carnea 11 (14), Galium austriacum 22 (14), Rosa pendulina 4 (11); Epilobietea angustifolii - Fragaria vesca 21 (11), Senecio ovatus ssp. ovatus 4 (11); FB -Asperula aristata 13 (12), Carex caryophyllea 13 (15), Gentiana germanica 6 (6), Polygala amarella 8 (13); JT -Carex curvula ssp. curvula 13 (15), Carex fuliginosa 13 (15), Festuca nigrescens 6 (17), Juncus jacquinii 6 (17), J. trifidus 6 (17), Leontodon helveticus 22 (17), Oreochloa disticha 11 (17), Veronica fruticans 6 (17); LV - Arc-tostaphylos alpina 13 (12), Loiseleuria procumbens 11 (17); M-Ar - Astrantia major ssp. major 6 (8), Des-champsia cespitosa 46 (11), Euphrasia picta ssp. picta 13 (12), Helictotrichon versicolor 6 (17), Poa annua 11 (16), P. supina 100 (9), Ranunculus tuberosus 6 (12), Senecio ovirensis ssp. gaudinii 8 (11), Vicia sepium 6 (12); MC -Chrysosplenium alternifolium 63 (11); MA - Aconitum napellus ssp. tauricum 14 (18), Chaerophyllum hirsutum 6 (8), Cortusa matthioli 14 (18), Hypericum richeri ssp. grisebachii 8 (11), Pedicularis recutita* 6 (8), Rumex al-pestris 6 (8), Salix glabra 14 (18), S. hastata 33 (14); N -Antennaria dioica 8 (13), Carex leporina 8 (13); QF -Adoxa moschatellina 8 (11), Anemone nemorosa 8 (11), Aquilegia nigricans 4 (11), Athyrium filix-femina 4 (11), Cardamine enneaphyllos 4 (11), Cardamine trifolia 4 (11), Carex digitata 29 (11), Fagus sylvaticae 4 (11), Festuca altissima 13 (11), Hieracium murorum 38 (11), Homogyne sylvestris 42 (11), Lamium orvala 4 (11), Lu-zula luzuloides ssp. luzuloides 25 (12), Maianthemum bifolium 17 (11), Paris quadrifolia 4 (11), Phegopteris connectilis 4 (11), Poa nemoralis 29 (11), Primula elatior 6 (8), Salix alba ssp. alba 6 (17), Stellaria nemorum ssp. montana 4 (11); SH - Luzula alpinopilosa ssp. alpinopi-losa 22 (17); SC - Carex nigra 92 (10), Gentiana utricu-losa 6 (12), Luzula sudetica 31 (12), Phleum alpinum 6 (17), Pinguicula vulgaris 6 (17); TG - Seseli libanotis 19 (12), Veronica chamaedrys agg. 6 (17); VP - Aposeris foetida 6 (8), Clematis alpina 25 (11), Drypteris dilatata 4 (11), Lonicera caerulea 46 (11), Pyrola minor 4 (11), P. rotundifolia 8 (10), Rhododendron ferrugineum 15 (13), Vaccinium myrtillus 50 (11), Valeriana tripteris 8 (11), Veronica urticifolia 4 (11); Other species - Alchemilla sp. 17 (8), 8 (11), 38 (15), 20 (19), A. flabellata 56 (12), 8 (13), Festuca sp. 44 (12), 8 (13), Thymus sp. 22 (8), 75 (15), Alchemilla anisiaca 6 (17), A. sericoneura 8 (13), A. subcrenata 6 (17), A. vallesiaca 29 (18), Campanula sp. 6 (8), Erigeron sp. 6 (17), Hieracium sp. 6 (8). Less frequent geoelements to Table 2 E-Alp./Apenn. - 7, 8; Europ./SW-Asiat. - 3, 13; SW-Europ.-Mont. - 17; lllyr. - 11; N-Europ./Am. - 11; E-Europ. - 14; Europ./Mont. - 12; SE-Europ-Mont./NE-Europ. - 18; Medit.-Mont. - 1; E-Europ. - 11: 1; W-Alp.** - 19: 1; W-Europ. - 11. List of syntaxa mentioned referred to in the text and Table 1 Thlaspietea rotundifolii Br.-Bl. 1948 Androsacetalia alpinae Br.-Bl. in Br.-Bl. & Jenny 1926 Drabetalia hoppeanae Zollitsch 1968 Thlaspietalia rotundifolii Br.-Bl. in Br.-Bl. & Jenny 1926 Thlaspion rotundifolii Jenny-Lips 1930 Petasition paradoxi Zollitsch ex Lippert 1966 Arabidetalia caeruleae Rübel ex Br.-Bl. 1949 Arabidion caeruleae Br.-Bl. in Br.-Bl. & Jenny 1926 em. Englisch 1999 Ranunculo traunfellneri-Festucetum nitidae (Wra-ber 1972) Englisch 1999 Saxifragetum stellaro-sedoidis Englisch 1999 var. geogr. Achillea oxyloba Surina 2005 var. geogr. Ranunculus traunfellneri Surina 2005 var. geogr. Campanula pulla Surina 2005 Saxifragetum hohenwartii Aichinger 1933 Salici herbaceae-Arabidion caeruleae Englisch 1999 Salici herbaceae-Arabidetum caeruleae Englisch 1999 Soldanello alpinae-Salicion retusae Englisch 1999 Salicetum retuso-reticulatae Br.-Bl. in Br.-Bl. & Jenny 1926 Homogyno discoloris-Salicetum retusae Aichinger 1933 Diantho alpinae-Salicetum retusae Englisch 1999 Selaginello selaginoidis-Salicetum reticulatae Englisch 1999 Salici retusae-Geranietum argentei Surina 2005 Potentillo brauneanae-Homogynetum discoloris Aichinger 1933 Salicion retusae Horvat 1949 Drepanoclado uncinati-Heliospermetum pusilli Surina & Vreš 2004 Elyno-Seslerietea Br.-Bl. 1948, Seslerietalia juncifo-liae Horvat 1930, Seslerion juncifoliae 1930, Caricion firmae Gams 1936, Gentiano terglouensis-Caricetum firmae T. Wraber 1970, Caricetum firmae Rübel 1911, Dryadetum octopetalae Rübel 1911, Salicetea herbaceae Br.-Bl. 1948, Salicion herbaceae Br.-Bl. in Br.-Bl. & Jenny 1926, Poo supinae-Cerastietum cerastoidis Söyrinki ex Oberd. 1957, Asplenietea trichomanis (Br.-Bl. in Meier & Br.-Bl. 1926) Oberd. 1957, Juncetea trifidi Hadač in Hadač & Klika 1944, Nardetea strictae Rivas Goday & Borja Carbonell 1961, Loiseleurio-Vaccinietea Eggler 1952, Montio-Cardaminetea Br.-Bl. & Tüxen ex Klika & Hadač 1944 em. Zechmeister 1993, Mulgedio-Aconite-tea Hadač & Klika in Klika & Hadač 1944, Epilobietea angustifolii Tüxen & Preising ex von Rochow 1951, Scheuchzerio-Caricetea fuscae Tüxen 1937, Molinio-Arrhenatheretea Tüxen 1937, Festuco-Brometea Br.-Bl. & Tüxen 1943, Trifolio-Geranietea Müller 1962, Artem-isietea vulgaris Lohmeyer, Preising & Tüxen ex von Rochow 1951, Erico-Pinetea Horvat 1959, Querco-Fagetea Br.-Bl. & Vlieg. 1937, Vaccinio-Piceetea Br.-Bl. 1930 em. Zupančič 1976.