SUCCESSIONAL STAGES IN THE DEVELOPMENT OF FOREST VEGETATION IN CIRQUES OF TWO VALLEYS IN THE JULIAN ALPS (NW SLOVENIA) SUKCESIJSKE STOPNJE V RAZVOJU GOZDNE VEGETACIJE V KRNICAH DVEH ALPSKIH DOLIN V JULIJSKIH ALPAH (SEVEROZAHODNA SLOVENIJA) Igor DAKSKOBLER1 ABSTRACT Successional stages in the development of forest vegetation in cirques of two valleys in the Julian Alps (NW Slovenia) We investigated different types of vegetation in cirques of two valleys in the Julian Alps, Zadnjica and Kot, which develop on slope screes and are heavily influenced by avalanches that occur here every year. Our phytosociological analysis identified the following shrub-forest developmental stages: Saliceteum eleagno-purpureae hieracietosum porrifolii var. Achnatherum calamagrostis, Homogyno sylvestris-Sali-cetum glabrae, Polysticho lonchitis-Rhamnetum fallacis, Rho-dodendro hirsuti-Laburnetum alpini, Rhodothamno-Larice-tum deciduae anemonetosum trifoliae and Rhododendro hir-suti-Fagetum sylvaticae var. Sorbus chamaemespilus. On un-consolidated fluvio-glacial deposits in the Kot Valley we determined the following successional stages: Salicetum ele-agno-purpurae hieracietosum porrifolii var. Pinus mugo, Amelanchiero ovalis-Pinetum mugo laricetosum and Rhododendro hirsuti-Fagetum sylvaticae petasitetosum paradoxi. Key words: phytosociology, succession, glacial valley, Julian Alps, Natura 2000, Triglav National Park, Slovenia http://dx.doi.org/10.3986/fbg0052 IZVLEČEK Sukcesijske stopnje v razvoju gozdne vegetacije v krnicah dveh alpskih dolin v Julijskih Alpah (severozahodna Slovenija) Raziskali smo različne oblike vegetacije v krnicah dveh dolin v Julijskih Alpah, Zadnjici in Kotu, ki nastajajo na pobočnem grušču in nanje odločilno vplivajo vsakoletni snežni plazovi. S fitocenološko analizo smo lahko prepoznali naslednje grmiščno-gozdne razvojne stopnje: Saliceteum eleagno-purpureae hieracietosum porrifolii var. Achnatherum calamagrostis, Homogyno sylvestris-Salicetum glabrae, Poly-sticho lonchitis-Rhamnetum fallacis, Rhododendro hirsuti--Laburnetum alpini, Rhodothamno-Laricetum deciduae anemonetosum trifoliae in Rhododendro hirsuti-Fagetum sylvati-cae var. Sorbus chamaemespilus. Na nesprijetih rečno-ledeniško nanosih v dolini Kot pa smo ugotovili naslednje sukcesijske stopnje: Salicetum eleagno-purpurae hieracietosum porrifolii var. Pinus mugo, Amelanchiero ovalis-Pinetum mugo laricetosum in Rhododendro hirsuti-Fagetum sylvaticae petasitetosum paradoxi. Ključne besede: fitocenologija, sukcesija, ledeniška dolina, Julijske Alpe, Natura 2000, Triglavski narodni park, Slovenija 1 Scientific Research Centre of the Slovenian Academy of Sciences and Arts, Institute of Biology, Regional Unit Tolmin, Brunov drevored 13, SI-5220 Tolmin, Igor.Dakskobler@zrc-sazu.si FOLIA BIOLOGICA ET GEOLOGICA 60/1, 101-127, LJUBLJANA 2019 I. DAKSKOBLER: SUCCESSIONAL STAGES IN THE DEVELOPMENT OF FOREST VEGETATION IN CIRQUES OF TWO VALLEYS 1 INTRODUCTION In recent years, successional processes and development of forest vegetation in Alpine valleys in Slovenia were studied in particular by M. Zupančič and collaborators (Zupančič & Žagar 2009, Zupančič, Skuma-vec & Mihorič 2017) as well as several geographers (Lovrenčak 2002, Blatnik & Repe 2013, Geršič et al. 2014). On our sporadic visits to the mountains through the Alpine valleys of Zadnjica in Trenta and Kot in the Upper Sava Valley (both in the Triglav National Park) our attention was drawn to different types of vegetation on slope screes and alluvial fans in their cirques, where annual avalanche activity is obviously one the main ecological factors. The human impact on the natural processes can be seen only in Zadnjica (the vicinity of a marked mountain trail) and is insignificant in the gable end of the Kot Valley (above the Ko-tarica falls). We made a phytosociological inventory of these, spatially relatively well-delimited shrub and forest communities and tried to classify them into a syn-taxonomic system. 2 METHODS The vegetation of gable ends of the Zadnjica and Kot method (Braun-Blanquet 1964). A total of 28 relevés valleys was inventoried using the Central-European (Figure 1) were entered into the FloVegSi database (T. Figure 1: Research area with approximate localities of investigated stands (yellow circles: stands on slope screes, red circles: stands on fluvio-glacial deposits) Slika 1: Raziskovano območje s približnimi nahajališči preučenih sestojev (rumeni krogci: sestoji na pobočnem grušču, rdeči krogci: sestoji na rečno-ledniških nanosih) 102 FOLIA BIOLOGICA ET GEOLOGICA 60/1 - 2019 I. DAKSKOBLER: SUCCESSIONAL STAGES IN THE DEVELOPMENT OF FOREST VEGETATION IN CIRQUES OF TWO VALLEYS Seliskar, Vres & A. Seliskar 2003). The relevés were arranged in two tables. Those on slope screes in the cirques of Zadnjica and Kot (above the Kotarica falls) and the relevés on gravelly plains in the upper part of Kot (Klin) were analysed separately. They were mutually compared by means of hierarchical classification, using the (unweighted) average linkage - UPGMA method with Wishart's similarity ratio. For this purpose, we transformed combined cover-abundance values into ordinal values 1- 9 (van der Maarel 1979). We used SYN-TAX 2000 (Podani 2001) software package. The nomenclatural source for the names of vascular plants is Martincic & al. (2007), except for Sesleria caerulea (Aeschimann et al. 2004b). Martincic (2003, 2011) is the nomenclatural source for the names of mosses and Suppan et al. (2000) for the names of lichens. The nomenclatural source for the names of syntaxa are Silc & Carni (2012), except for the name of the class Querco-Fagetea Braun-Blanquet et Vlieger in Vlieger 1937. 2.1 Ecological description of the study area Both cirques, in Zadnjica and Kot, have a predominantly shady aspect (northeast and partly northwest); parent material is unconsolidated moraine (till) and above all slope screes (Jurkovšek 1987 a, b). The elevation of both cirques is similar, spanning 1000 to 1300 m. There are no significant differences in the precipitation volume either, with the average exceeding 2000 mm (B. Zupančič 1998), a large portion of which falls as snow. The mean annual temperature is similar too, ranging between 4 and 6 °C (Cegnar 1998), with the Zadnjica Valley still showing some sub-Mediterranean influence. In recent decades the area has seen a distinctly downward trend in precipitation and higher average temperatures (Dolinar 2018). Both cirques have a characteristically cold local climate and over the winter they receive snow avalanches from their rock walls or steps (Volk Bahun, Zorn & Pavšek 2018). The soil is therefore very shallow, partly collu-vial and undeveloped, lithosol and moder rendzina. 3 RESULTS AND DISCUSSION 3.1 Phytosociological analysis of the shrub-forest vegetation sequence on slope screes in the avalanche area in the gable end of the Zadnjica and Kot valleys Table 1 comprises 22 relevés of stands on slope screes that grouped by floristic similarity as indicated in Figure 2. In Table 1 we arranged the relevés by taking into account the dominant species of the highest stand layer, which means that our classification does not fully follow the results of hierarchical classification. The most unique form, the community of red and grey willow (Salicetum eleagno-purpureae hieracietosumpor-rifolii var. Achnatherum calamagrostis) with Achnathe-rum calamagrostis, Athamanta cretensis and Petasites paradoxus dominating the herb layer, was found only along a small torrential brook running from under the waterfall in the Kanjavec rock face down the slope scree until it sinks (relevé 1in Table 1). Open Rhamnus fallax shrub stands were found only in the gable end of Kot, on rather coarse and not entirely consolidated gravel - relevés 3-5 in Table 1. In terms of species composition they obviously belong to the association Polysticho lonchitis-Rhamnetum fallacis, which was described a few years ago (Dakskobler, Franz & Rozman 2013) and is more frequent in the Slovene Alps than previously thought (see also Blatnik & Repe 2013). In recent years it was spotted, but not yet inventoried, also in the Krma Valley. The dominant shrub layer species in one of the Rhamnus fallax stands in the gable of Kot (relevé 2 in Table 1) is Salix glabra, and this stand could be classified also into the association Homogyno sylvestris-Salicetum glabrae (comp. Dakskobler & Surina 2017). Similarly, larch dominated the highest stand layers only in the gable end of Kot, on one location on fully consolidated gravel. Its initial stand (relevé 6 in Table 1) can be classified into the association Rhodothamno-Laricetum. Although Anemone trifolia was not found in the recorded stand, based on the whole floristic composition it should be classified into the subassociation anemone-tosum trifoliae, which is characterised by abundant Sesleria caerulea in the herb layer (comp. Dakskobler, Seliskar & Rozman 2018: 259) Most relevés on the left side of the dendrogram in Figure 2, i.e. relevés in the central part of Table 1, characterise pioneer shrub communities with Laburnum alpinum dominating the highest stand layer. Laburnum alpinum is a south-European montane species distributed across most of the Alps (except the Northeastern and partly Central Alps), in the south and central Jura, the north and central Apennines, the Carpathians and the Dinaric Alps (Aeschimann et al. 103 FOLIA BIOLOGICA ET GEOLOGICA 60/1 - 2019 I. DAKSKOBLER: SUCCESSIONAL STAGES IN THE DEVELOPMENT OF FOREST VEGETATION IN CIRQUES OF TWO VALLEYS 2004a, Brus 1999, 2005). It occurs on calcareous, mixed and silicate bedrock, in the colline, montane and partly the subalpine belt, mainly on beech forest sites. As a shade-tolerant species it frequently occurs in the lower tree and shrub layer, and establishes itself as a pioneer in windthrow and landslide areas, and on erosion slopes. Figure 3 shows its distribution in Slovenia. Laburnum alpinum is the most frequent in the stands of associations Arunco-Fagetum, Ostryo-Fage-tum, Seslerio autumnalis-Fagetum, Lamio orvalae-Fage-tum, Ranunculo platanifolii-Fagetum, Anemono trifoli-ae-Fagetum, Homogyno sylvestris-Fagetum, Saxifrago cuneifolii-Fagetum, Rhododendro hirsuti-Fagetum, Polysticho lonchitis-Fagetum, Luzulo-Fagetum sylvati-cae, Laburno alpini-Piceetum, Rhodothamno-Larice- tum, Saxifrago petraeae-Tilietum, Seslerio albicantis--Ostryetum carpinifoliae, Fraxino orni-Ostryetum car-pinifoliae, Rhododendro hirsuti-Ostryetum carpinifoliae, Rhodothamno-Pinetum mugo, Amelanchiero ovalis--Pinetum mugo, but it occurs also in several other forest and shrub communities (source: FloVegSi database, T. Seliškar, Vreš & A. Seliškar, 2003). The highest it was recorded in the association Rhodothamno-Pine-tum mugo at 1560 m a.s.l. (pasture Stržiškarska Planina under Mt. Poljanski Vrh and Mt. Hohkovbl), in the association Polysticho lonchitis-Fagetum at 1570 m (Planinica under Loška Stena rock wall), in the subassociation Anemono-Fagetum laricetosum at 1615 m (Pod Pečmi above the Tamar Valley, under Srednja Ponca), in the association Rhodothamno-Laricetum at ¡y •a CS 0,850,80,750,70,650,60,55 0,5 0,450,40,350,30,250,20,150,10,05 0 J J j j j j j j j j -a -a -a -a -a -a -a -a -a -a U rt rt rt rt rt rt rt rt rt -a rt -a rt -a rt -a Pi -a Pi -a Pi rt S rt s rt S Figure 2: Dendrogram of relevés of shrub- and forest communities on slope screes in cirques of Zadnjica and Kot valleys (UPGMA, similarity ratio) Slika 2: Dendrogram popisov grmiščnih in gozdnih združb na pobočnem grušč v krnicah dolin Zadnjice in Kota (UPGMA, similarity ratio) Legend / Legenda ChLa Carici humilis-Laburnetum alpini nom. prov. RhLa Rhododendro hirsuti-Laburnetum alpini RhF Rhododendro hirsuti-Fagetum sylvaticae PlRf Polysticho lonchitis-Rhamnetum fallacis HsSa Homogyno sylvestris-Salicetum glabrae Sep Salicetum eleagno-purpureae hieracietosum porrifolii var. Achnatherum calamagrostis 104 FOLIA BIOLOGICA ET GEOLOGICA 60/1 - 2019 I. DAKSKOBLER: SUCCESSIONAL STAGES IN THE DEVELOPMENT OF FOREST VEGETATION IN CIRQUES OF TWO VALLEYS 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 Figure 3: Distribution of Laburnum alpinum in Slovenia, according to the data in the FloVegSi database (authors of the map are I. Dakskobler, B. Anderle, A. Seliškar and B. Vreš) Slika 3: Razširjenost vrste Laburnum alpinum v Sloveniji po podatkih v bazi FloVegSi (avtorji zemljevida so I. Dakskobler, B. Anderle, A. Seliškar in B. Vreš) 1660 m a.s.l. (Skok above Zadnjica under Mt. Triglav; under the peak Kumlehova Glava above the Suha Pisnica brook). These are the highest elevations of its occurrence known to us in the Julian Alps. Brus (1999: 206, 2005: 241) reports that on southern (the warmest, sunny) slopes this species occurs up to 1900 m a.s.l., which almost certainly does not refer to the Julian Alps, but to the Alps in general, (Brus, in litt.). As a pioneer it establishes itself on windthrow areas, in particular in stands of associations Anemono trifoliae-Fa-getum, Ranunculo platanifolii-Fagetum and Rhododen-dro hirsuti-Fagetum. In the northern part of the Di-naric Alps we described a successional stage in natural gaps of montane beech and fir-beech forest as the association Laburno alpini-Rhamnetum fallacis (Dakskobler, Franz & Rozman 2013). In the Jura, the association Sorbo mougeotii-Laburnetum alpini Gehu & Gehu-Franck ex Foucault 2012 is described and designated as the edge shrub community on the sites of thermo-basophilous beech forests (Richard 1968, Gehu & Gehu-Franck 1987, Biondi, Pinzi & Gubellini 2004, Foucault 2012, Theurillat 2019, in litt.). In the Apennines, a successional stage Geranio nodosi-Laburnetum alpini Castelli, Biondi & Ballelli 2001 and its subassociation sorbetosum ariae Biondi, Pinzi et Gubellini 2004 were described on the sites of beech forests from the association Cardamino kitaibe-lii-Fagetum sylvaticae (Castelli, Biondi & Ballelli 2001, Biondi, Pinzi & Gubellini 2004). Our relevés are distinctly different from the shrub communities in the Jura and the Apennines. The species shared with the stands of the association Sorbo mougeotii-Laburnetum alpini from the Jura are Laburnum alpinum, Sorbus mougeotii, S. aria, S. chamaeme-spilus, Salix appendiculata, Fagus sylvatica, Lonicera xylosteum, Mercurialis perennis, Acer pseudoplatanus, Polygonatum verticillatum, Fragaria vesca, Rubus idae-us, Lonicera alpigena and Aconitum lycoctonum s. lat. The species shared with the stands of the subassociation Gerenio nodosi-Laburnetum alpini sorbetosoum ariae from the Apennines are Laburnum alpinum, Sor-bus aria, Rhamnus fallax, Mercurialis perennis, Fagus sylvatica, Solidago virgaurea, Hepatica nobilis, Carex digitata, Ostrya carpinifolia, Fragaria vesca, Thalic- 105 FOLIA BIOLOGICA ET GEOLOGICA 60/1 - 2019 I. DAKSKOBLER: SUCCESSIONAL STAGES IN THE DEVELOPMENT OF FOREST VEGETATION IN CIRQUES OF TWO VALLEYS trum aquilegiifolium, Euonymus latifolia, Galium album and Hypericum montanum. Despite the listed species that they have in common, the studied Laburnum alpinum community from the Julian Alps differs from its communities from the Jura and the Apennines mainly with the presence of distinctly frigophi-lous species characteristic for subalpine beech, larch, spruce and dwarf pine communities, such as Rhododendron hirsutum, Polystichum lonchitis, Pinus mugo, Clematis alpina, Homogyne sylvestris, Picea abies and Larix decidua. Most of the stands with dominating Laburnum alpinum in the highest layer are therefore classified into the new association Rhododendro hirsuti--Laburnetum alpini. Its nomenclatural type, holotypus, is relevé 16 in Table 1. The diagnostic species of the new association are Laburnum alpinum, Rhododendron hirsutum, Polystichum lonchitis, Pinus mugo, Gymnocarpium robertianum, Clematis alpina, Homo-gyne sylvestris, Larix decidua, Sorbus chamaemespilus and Salix appendiculata and its phytogeographical differential species are Rhodothamnus chamaecistus, Cyclamen purpurascens and Anemone trifolia. The listed species characterise cold, stony sites where beech can still establish itself in favourable conditions (absence of avalanches), as manifested by the entire species composition. The new association is thus classified into the alliance Aremonio-Fagion, order Fagetalia syl-vaticae and class Querco-Fagetea (Carpino-Fagetea), even though it could also be classified into the classes Erico-Pinetea or Roso pendulinae-Pinetea mugo. There are several differences between the stands in the Zadnjica and Kot valleys. The stands in Kot on moister and coarser gravel are characterised in particular by a considerable admixture of sycamore maple (Acer pseudoplatanus), so they are described as a new subassociation aceretosum pseudoplatani (its nomen-clatural type is the same as the nomenclatural type of the new association, relevé 16 in Table 1). The stands in the gable end of the Zadnjica Valley are mainly situated on finer gravel, on slightly drier and warmer sites, and are differentiated by Ostrya carpinifolia and Carex alba. The nomenclatural type, holotypus, of the new subassociation ostryetosum carpinifoliae is relevé 11 in Table 1. The most unique among the relevés of this subassociation is relevé 14, where the shrub layer is completely dominated by dwarf pine (Pinus mugo). Although this stand could also be classified into the association Amelanchiero-Pinetum mugo, it was included in the association Rhododendro-Laburnetum alpini, because in terms of entire species composition it grouped with other relevés of the latter association. However, the same cannot apply to relevé 7 in Table 1, which characterises an open Laburnum alpinum shrub on the debris cone under Mt. Zadnjiski Ozebnik. It is characterised by character scree and stony grassland species such as Achnatherum calamagrostis, Ade-nostyles glabra, Calamagrostis varia and Carex humilis. This stand is tentatively classified into the provisional association Carici humilis-Laburnetum alpini nom. prov. and is syndynamically related to the stands of the association Anemono trifoliae-Fagetum. The tree layer in the forest stands located on the edge or at the bottom of the studied glacial cirques, in areas that are less exposed to avalanches (relevés 19-22 in Table 1), is dominated by beech. The entire species composition indicates a community on extreme sites, very similar to beech stands on the upper forest line, classified into the association Polysticho lonchitis-Fage-tum. Also possible is classification into the association Rhododendro hirsuti-Fagetum, as these stands comprise almost all of the diagnostic species of this association that otherwise characterises beech sites in the submontane and montane belt up to some 1200 m a.s.l. Given the elevation of our relevés (under 1200 m) the latter association is considered more appropriate, but to demonstrate the similarity with the stands of the first, Polysticho lonchitis-Fagetum, it is described as a special altitudinal variant with Sorbus chamaemespilus (Rhododendro hirsuti-Fagetum var. Sorbus chamaemespilus). In the Julian Alps, this species otherwise characterises communities of the altimontane and subalpine belts, and in the (lower) montane belt it is usually limited to frost hollows. In the stands of the association Rhododendro hirsuti-Fagetum in the Julian Alps it was reported on only a few occasions (see Dakskobler 2003). From the relevés in Table 1 we can infer the likely sequence in the development of shrub and forest vegetation on slope screes in avalanche areas in cirques of glacial valleys in the Julian Alps: Polysticho lonchitis-Rhamnetum fallacis (Homo-gyno sylvestris-Salicetum glabrae) - Rhododendro hir-suti-Laburnetum alpini (Amelanchiero-Pinetum mugo) - Rhododendro hirsuti-Fagetum sylvaticae (Rhodotha-mno-Laricetum). 3.2 Phytosociological analysis of the sequence in shrub-forest vegetation on fluvio-glacial deposits in the Kot Valley (Klin). In Table 2 we arranged six relevés made on the gravelly plain Klin in the upper part of the Kot Valley. They were mutually compared using hierarchical classification, which produced the dendrogram in Figure 4. 106 FOLIA BIOLOGICA ET GEOLOGICA 60/1 - 2019 I. DAKSKOBLER: SUCCESSIONAL STAGES IN THE DEVELOPMENT OF FOREST VEGETATION IN CIRQUES OF TWO VALLEYS Successional development of vegetation in the upper part of the Kot Valley was studied in detail by Zupančič, Skumavec & MiHORič (2017). They determined and presented with a phytosociological table the sequence of the following plant communities: Petasitetum paradoxi - Petasiti-Salicetum purpu-reo-albae - Calamagrostio variae-Pinetum mugo -Dryado-Piceetum - Rhodothamno-Pinetum mugo -Rhododendro-Fagetum - Anemono-Fagetum lycopodie-tosum. In this area, i.e. on the gravelly plain under the Ko-tarica falls, we were interested mainly in beech stands with hairy alpenrose (Rhododendro hirsuti-Fagetum), as the findings by Zupančič et al. (ibid.) were new to the forest vegetation in this part of the Julian Alps. We inventoried several typical stands and contact communities of Pinus mugo and Salix eleagnos. Only six relevés, however, are not sufficient to allow us to infer the right sequence in the development of shrub-forest vegetation. Our goal is therefore not to correct or supplement the findings of the much more detailed recent analysis, but merely to classify our relevés into a syn-taxonomic system. Relevé 1 in Table 1 could be classified also into the association Salicetum eleagno-purpureae. Although a comparison with the relevés of this community from the foothills of the Julian Alps, in the Nadiza Valley (Suc & Cusin 2000) does indicate some distinct differences, also with the variant with Petasites paradoxus. Oriolo & Poldini (2002) classify slightly similar Figure 4: Dendrogram of relevés of shrub and forest vegetation on fluvio-glacial gravel sediments in Klin, in the upper part of the Kot Valley (UPGMA, similarity ratio) Slika 4: Dendrogram popisov grmiščne in gozdne vegetacije na rečno-ledeniških sedimentih v Klinu v zgornjem delu doline Kot (UPGMA, similarity ratio) Legend/ Legenda: Sep Salicetum eleagno-purpureae hieracietosum porrifolii var. Pinus mugo AoPm Amelanchiero ovalis-Pinetum mugo laricetosum nom. prov. RhFpp Rhododendro hirsuti-Fagetum sylvaticae petasitetosum paradoxi 107 FOLIA BIOLOGICA ET GEOLOGICA 60/1 - 2019 I. DAKSKOBLER: SUCCESSIONAL STAGES IN THE DEVELOPMENT OF FOREST VEGETATION IN CIRQUES OF TWO VALLEYS stands into the montane altitudinal form of the subassociation typicum. There is also a certain similarity with relevés of the subassociation Salicetum eleagno--purpureae hieracietosum porrifolii described by ČušiN & ŠiLC (2006) on gravel bars of the Soča River near Bovec. The species that differentiate our relevé against the relevés published by Šilc & ČušiN (2000) and Ori-OLO & Poldini (2002) are in particular Pinus mugo, Hieracium porrifolium, Rhododendron hirsutum, Rhamnus fallax, Salix waldsteiniana, Dryas octopetala, Primula clusiana and many species of the classes Elyno-Se-slerietea and Thlaspietea rotundifolii. Some of the diagnostic species shared with the stands of the subassociation -hieracietosum porrifolii are Hieracium porrifolium, H. piloselloides, Carduus crassifolius, Biscutella laevigata, Campanula cespitosa, Silene vulgaris subsp. glareosa, Achnatherum calamagrostis, Rumex scutatus, Sesleria caerulea and Dryas octopetala. Our relevé at least slightly resembles also the relevé of the sy ntaxon Petasiti-Sali-cetumpurpureo-albae (Zupančič, Skumavec & Miho-rič 2017, relevé 2 in the phytosociological table), but with a significant difference - the dominant species there is Salix eleagnos and not Salix alba - which is a result of different site conditions (M. Zupančič, in litt.). The above comparisons allow us to classify our relevé into the new variant Salicetum eleagno-purpureae hieracietosum porrifolii var. Pinus mugo var. nov. The differential species of the new variant are Pinus mugo, Rhododendron hirsutum, Salix waldsteiniana and Potentilla clusiana. Relevés 2 and 3 in Table 2 can be classified into the association Amelanchiero ovalis-Pinetum mugo, as it comprises several its diagnostic species, but not Fraxinus ornus and Ostrya carpinifolia. Another option would be classification into the association Rhodotha-mno-Pinetum mugo. Given the elevation of occurrence we prefer (like in the case of similar associations Rho-dodendro hirsuti-Fagetum / Polysticho lonchitis-Fage-tum) the dwarf pine community of lower altitudes (Amelanchiero-Pinetum mugo), indicating its similarity with the stands of the association Rhodothamno-Pi-netum mugo with the provisional name of the subassociation, laricetosum. Stands in relevés 4-6 can be classified into the association Rhododendro hirsuti-Fagetum sylvaticae, as they comprise most of its diagnostic species. However, these stands could also be classified into the association Polysticho lonchitis-Fagetum. The species supporting this classification is Sorbus chamaemepsilus, so these stands can also be treated within the already mentioned altitudinal variant. On the other hand, the studied stands differ from other previously described forms of the association Rhododendro hirsuti-Fagetum with certain ecological characteristics. The stands of this association dominate on steep to precipitous shady slopes, whereas ours were found on levelled terrain and gravel deposits. Based on the above, they can be described and typified as a new subassociation Rhodo-dendro hirsuti-Fagetum sylvaticae petasitetosum para-doxi subass. nov. The nomenclatural type, holotypus, is relevé 6 in Table 2. The differential species of the new subassociation are Petasites paradoxus and Brachypo-dium rupestre. 3.3 An overview of syntax described or discussed in the article: Salicetea purpureae Moor 1958 Salicetalia purpureae Moor 1958 Salicion eleagno-daphnoidis (Moor 1958) Grass 1993 Salicetum eleagno-purpureae Sillinger 1933 -hieracietosum porrifolii Čušin et Šilc 2006 var. Pinus mugo var. nov. var. Achnatherum calamagrostis var. nov. Querco-Fagetea Br.-Bl. & Vlieg. 1937 Rhamnetali fallacis Fukarek 1969 Seslerio calcariae-Rhamnion fallacis Dakskobler, Franz et Rozman 2013 Polysticho lonchitis-Rhamnetum fallacis Dakskobler, Franz et Rozman 2013 Fagetalia sylvaticae Pawl. in Pawl. & al. 1928 Aremonio-Fagion (Ht. 1938) Borhidi in Tôrôk, Podani & Borhidi 1989 Rhododendro hirsuti-Laburnetum alpini ass. nov. hoc loco -ostryetosum carpinifoliae subass. nov. hoc loco -aceretosum pseudoplatani subass. nov. hoc loco Carici humilis-Laburnetum alpini nom. prov. Rhododendro hirsuti-Fagetum sylvaticae Accetto ex Dakskobler 1998 -petasitetosum paradoxi subass. nov. hoc loco Vaccinio-Piceetea Br.-Bl. in Br.-Bl., Sissingh et Vlieger 1939 Piceetalia excelsae Pawlowski in Pawlowski et al. 1928 Erico-Pinion mugo Leibundgut 1948 Rhodothamno-Laricetum deciduae Willner et Zuk-rigl 1999 -anemonetosum trifoliae Dakskobler 2006 Erico-Pinetea Horvat 1959 Erico-Pinetalia Horvat 1959 Fraxino orni-Pinion nigrae-sylvestris Zupančič 2007 Helleboro nigri-Pinenion (Horvat 1959) Zupančič 2007 Amelanchiero-Pinetum mugo Minghetti in Pe-drotti 1994 108 FOLIA BIOLOGICA ET GEOLOGICA 60/1 - 2019 I. DAKSKOBLER: SUCCESSIONAL STAGES IN THE DEVELOPMENT OF FOREST VEGETATION IN CIRQUES OF TWO VALLEYS -laricetosum nom. prov. Rhododendro hirsuti-Ericetea carneae Schubert et al. 2001 Rhododendro hirsuti-Ericetalia carneae Grabherr, Greimler et Mucina 1993 Ericion carneae Rübel ex Grabherr, Greimler et Mucina 1993 Homogyno sylvestris-Salicetum glabrae Dakskobler et Surina 2017 4 CONCLUSIONS On slope screes and gravel deposits in the gable end of two glacial valleys in the Julian Alps, in Zadnjica and Kot (Triglav National Park), we phytosociologically described several shrub and forest communities whose development is affected not only by the size, consolidation and moisture level of the gravel, but most decisively by avalanches that occur there every year. Only two shrub and one forest community were documented with a sufficient number of relevés. Stands of associations Polysticho lonchitis-Rhamnetum fallacis and Amelanchiero ovalis-Pinetum mugo are new to the Kot Valley. We described a new association Rhododendro hirsuti-Laburnetum alpini, which belongs to the group of shrub-forest communities of the montane-subalpine belt (Rhododendro hirsuti-Sorbetum aucupariae, Rho-dodendro hirsuti-Salicetum appendiculatae, Rhododen-dro hirsuti-Betuletum carpaticae, Alno viridis-Sorbe-tum aucupariae, Calamagrostio arundinaceae-Sorbe-tum ariae) - comp. Dakskobler, Kutnar & Rozman (2016) that are a more or less long-term stages on potential beech sites and their occurrence could, although not necessarily, be associated with human impact or past activities. In our case, the stands of the new association are a stage in development towards beech forest on extreme sites in the montane belt classified into the association Rhododendro hirsuti-Fagetum. The stands of this association in the gable ends of Zadnjica and Kot valleys are characterised by their similarity with stands of the association Polysticho lonchitis-Fagetum (and are therefore treated as a special altitudinal variant with Sorbus chamaemespilus). Its form on fluvioglacial gravel deposits that can be treated as a new subassociation petasitetosum paradoxi was found only on levelled terrain in the Kot Valley. Our findings confirm and supplement the findings of Zupančič, Sku-mavec & Mihorič (2017) on the exceptionally diverse and fascinating shrub-forest vegetation in the gable end of the Kot Valley and its substantial biotic value. Most of this vegetation belongs to Natura 2000 habitat types: 91K0 Illyrian Fagus sylvatica forests (Aremonio--Fagion), 4070 Pinus mugo and Rhododendrum hirsu-tum shrubs (Mugo-Rhododendretum hirsuti) and 9420 Alpine Larix decidua forests. It comprises several protected (Cypripedium calceolus - also a Natura 2000 species, Gymnadenia odoratissima, G. conopsea, Epi-pactis helleborine, Dianthus sternbergii, Gentiana clu-sii, Lycopodium annotinum), endemic (Cerastium sub-triflorum, Papaver alpinum subsp. ernesti-mayeri = P. ernesti-mayeri) and rare species (Sorbus mougeotii). The study area is situated in the inner Triglav National Park area, so human impact on the development of vegetation here (other than increased mountaineering in recent years and human-induced climate changes) is insignificant. 5 POVZETEK V zatrepu dveh ledeniških dolin v Julijskih Alpah, v Zadnjici in Kotu (Triglavski narodni park), smo na po-bočnih gruščih in prodnatih nanosih fitocenološko opisali več grmiščnih in gozdnih združb, na katerih razvoj poleg velikosti in ustaljenosti ter vlažnosti grušča najbolj odločilno vplivajo predvsem vsakoletni snežni plazovi. Z zadostnim številom popisov smo v obeh dolinah dokumentirali le dve grmiščni in eno gozdno združbo. Sestoji asociacij Polysticho lonchitis--Rhamnetum fallacis in Amelanchiero ovalis-Pinetum mugo so novost v dolini Kota. V obeh raziskovanih do- linah smo našli sestoje subasociacije Salicetum elea-gno-purpureae hieracietosum porrifolii, ki smo jo do zdaj poznali le v Zgornjem Posočju (ČušiN & Šilc 2006). Kot novo smo opisali asociacijo Rhododendro hirsuti-Laburnetum alpini, ki sodi v skupino grmišč-no-gozdnih združb montansko-subalpinskega pasu (Rhododendro hirsuti-Sorbetum aucupariae, Rhodo-dendro hirsuti-Salicetum appendiculatae, Rhododendro hirsuti-Betuletum carpaticae, Alno viridis-Sorbetum aucupariae, Calamagrostio arundinaceae-Sorbetum ariae) - prim. Dakskobler, Kutnar & Rozman 109 FOLIA BIOLOGICA ET GEOLOGICA 60/1 - 2019 I. DAKSKOBLER: SUCCESSIONAL STAGES IN THE DEVELOPMENT OF FOREST VEGETATION IN CIRQUES OF TWO VALLEYS (2016), ki so bolj ali manj dolgotrajen stadij na potencialno bukovih rastiščih in je njihov nastanek lahko, ne pa nujno, povezan tudi s človekovimi vplivi oz. njegovo preteklo dejavnostjo. V našem primeru so sestoji nove asociacije razvojna stopnja proti bukovemu gozdu na skrajnih rastiščih v montanskem pasu, ki jih uvrščamo v asociacijo Rhododendro hirsuti-Fagetum. Za sestoje te asociacije v zatrepih dolin Zadnjice in Kota je značilna podobnost s sestoji asociacije Polysticho lon-chitis-Fagetum (zato jih obravnavamo kot posebno višinsko varianto z vrsto Sorbus chamaemespilus). Samo v dolini Kot pa smo doslej našli njeno obliko na rečno--ledeniških prodnatih nanosih, na uravnavi, ki jo lahko vrednotimo kot novo subasociacijo petasiteto-sum paradoxi. Naša spoznanja potrjujejo in dopolnjujejo ugotovitve Zupančiča in sodelavcev (2017) o izredno raznolikem in zanimivem grmiščno-gozdnem ra- stju v zatrepu doline Kot in o njegovi veliki biotski vrednosti. Večinsko sodi v evropsko varstveno pomembne habitatne tipe: 91K0 Ilirski bukovi gozdovi (Aremonio-Fagion), 4070 *Ruševje z dlakavim slečem (Mugo-Rhododendretum hirsuti) in 9420 Alpski mace-snovi gozdovi. V njem raste več zavarovanih vrst (Cypripedium calceolus - tudi Natura 2000 vrsta, Gymnadenia odoratissima, G. conopsea, Epipactis hel-leborine, Dianthus sternbergii, Gentiana clusii, Lycopo-dium annotinum), endemitov (Cerastium subtriflorum, Papaver alpinum subsp. ernesti-mayeri = P. ernesti-ma-yeri) in redkih vrst (Sorbus mougeotii). Ker raziskano območje leži v strožje varovanem delu Triglavskega narodnega parka, so človekovi vplivi na tukajšnji razvoj vegetacije (z izjemo povečanega obiska planincev v zadnjih letih in od človeka povzročenih podnebnih sprememb) majhni. ACKNOWLEDGEMENTS Academician Dr. Mitja Zupančič peer-reviewed my paper and offered some useful corrections and improvements. Prof. dr. Andraž Čarni kindly helped with literature references. Sincere thanks to Prof. Dr. Jean-Paul Theurillat for his valuable information regarding the valid name of the Laburnum alpinum community in the Jura. Brane Anderle, Andrej Seliškar and Branko Vreš are co-authors of Figure 3. Iztok Sajko prepared Figure 1 for print. I acknowledge the financial support from the Slovenian Research Agency (research core funding No. P1-0236). English translation by Andreja Šalamon Verbič. REFERENCES - LITERATURA Aeschimann, D., Lauber, K., Moser, D. M. & Theurillat, J.-P. 2004a: Flora alpina. Bd. 1: Lycopodiaceae-Apiaceae. Haupt Verlag, Bern, Stuttgart, Wien. Aeschimann, D., Lauber, K., Moser, D. M. & Theurillat, J.-P. 2004b: Flora alpina. Bd. 2: Gentianaceae-Orchi-daceae. Haupt Verlag, Bern, Stuttgart, Wien. Biondi, E., M. Pinzi & L. 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Folia biologica et geologica (Ljubljana) 50 (1): 189-211. 111 FOLIA BIOLOGICA ET GEOLOGICA 60/1 - 2019 I. DAKSKOBLER: SUCCESSIONAL STAGES IN THE DEVELOPMENT OF FOREST VEGETATION IN CIRQUES OF TWO VALLEYS Zupančič, M. J. Skumavec & J. Mihorič, 2017: Floristične in vegetacijske zanimivosti Kota (Julijske Alpe, SZ Slovenija). Naravoslovno društvo Bled in Slovenska akademija znanosti in umetnosti. Zgornje Laze-Ljubljana. Photos / Fotografije (Photo/Foto: I. Dakskobler) Figure 5: Stand of the syntaxon Salicetum eleagno-purpureae hieracietosum porrifolii var. Achnatherum calamagrostis in the Zadnjica Valley Slika 5: Sestoj sintaksona Salicetum eleagno-purpureae hieracietosum porrifolii var. Achnatherum calamagrostis v dolini Zadnjice 112 FOLIA BIOLOGICA ET GEOLOGICA 60/1 - 2019 I. DAKSKOBLER: SUCCESSIONAL STAGES IN THE DEVELOPMENT OF FOREST VEGETATION IN CIRQUES OF TWO VALLEYS Figure 6: Stand of the syntaxon Rhododendro hirsuti-Laburnetum alpini ostryetosum in the Zadnjica Valley Slika 6: Sestoj sintaksona Rhododendro hirsuti-Laburnetum alpini ostryetosum v dolini Zadnjice Figure 7: Stand of the association Polysticho lonchitis-Rhamnetum fallacis in the Kot Valley Slika 7: Sestoj asociacije Polysticho lonchitis-Rhamnetum fallacis v dolini Kot 113 FOLIA BIOLOGICA ET GEOLOGICA 60/1 - 2019 I. DAKSKOBLER: SUCCESSIONAL STAGES IN THE DEVELOPMENT OF FOREST VEGETATION IN CIRQUES OF TWO VALLEYS Figure 8: Successional stages of vegetation on slope screes in the gable end of the Kot Valley (from the left to the right): Rhododen-dro-Laburnetum - Rhodothamno-Laricetum - Polysticho-Rhamnetum - Rhododendro-Laburnetum - Rhododendro-Fagetum Slika 8: Sukcesijske stopnje rastja na pobočnem grušču v zatrepu doline kot (od leve proti desni): Rhododendro-Laburnetum - Rhodothamno-Laricetum - Polysticho-Rhamnetum - Rhododendro-Laburnetum - Rhododendro-Fagetum Figure 9: Stand of the syntaxon Salicetum eleagno-purpureae hieracietosum porrifolii var. Pinus mugo in the Kot Valley Slika 9: Sestoj sintaksona Salicetum eleagno-purpureae hieracietosum porrifolii var. Pinus mugo v dolini Kot 114 FOLIA BIOLOGICA ET GEOLOGICA 60/1 - 2019 I. DAKSKOBLER: SUCCESSIONAL STAGES IN THE DEVELOPMENT OF FOREST VEGETATION IN CIRQUES OF TWO VALLEYS Figure 10: Stand of the syntaxon Rhododendro hirsuti-Fagetum petasitetosum paradoxi in the Kot Valley Slika 10: Sestoj sintaksona Rhododendro hirsuti-Fagetum petasitetosum paradoxi v dolini Kot 115 FOLIA BIOLOGICA ET GEOLOGICA 60/1 - 2019 •n O t-; > w O t-1 o o o > M o M o t-1 o 2 o > Table 1: Successional stages of shrub-forest vegetation on slope screes in glacial cirques of Zadnjica and Kot valleys in the Julian Alps Preglednica 1: Sukcesijski stadiji rastja na pobočnem grušču v krnicah dolin Zadnjice in Kota v Julijskih Alpah Number of releve (Zaporedna številka popisa) Database number of releve (Delovna številka popisa) Elevation in m (Nadmorska višina v m) Aspect (Lega) Slope in degrees (Nagib v stopinjah) Parent material (Matična podlaga) 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 co co o CO co co 258936 S \o Ln S co S 71 O w f tu ?? ^ ei O O W o z; > f v: > O M § X tn a M < M f O TJ S M z; ^ o •n •n O M < M O M ^ > ¡j O z § o 53 o a tn O •n t O g t-1 t-1 M ►C o t-; > M O f o o > to ^ o tu O t-1 O O o > Number of relevé (Zaporedna številka popisa) Coordinate GK X (D-48) 9 10 11 12 13 14 15 16 17 18 19 20 21 22 Pr. Fr. iTt \o o ^ t— iN Lil rr, rr, r-J rr, \C CT\ Ln c-, rr, rr, rej co m rr, rr, r. i r i r | r , r r \r r . r r , r r ¡r ,r ¡r ,r , r Ir Ir, r Diagnostic species of the associations and subassociations (Diagnostične vrste asociacij in subasociacij) SP Salixpurpurea SP Salix eleagnos BA Salix glabra AF Rhamnus fallax AF Rhamnus fallax AF Rhamnus fallax EP Rhododendron hirsutum VP Polystichum lonchitis MuAAconitum lycoctonum subsp. ranunculifolium TR Gymnocarpium robertianum FS Laburnum alpinum FS Laburnum alpinum FS Laburnum alpinum FS Laburnum alpinum EP Pinus mugo EP Pinus mugo VP Clematis alpina VP Homogyne sylvestris BA Salix appendiculata BA Sorbus chamaemespilus VP Larix decidua VP Larix decidua VP Larix decidua EP Carex alba QP Ostrya carpinifolia QP Ostrya carpinifolia QP Ostrya carpinifolia QP Ostrya carpinifolia TA Acerpseudoplatanus TA Acer pseudoplatanus TA Acer pseudoplatanus TA Acer pseudoplatanus FS Fagus sylvatica FS Fagus sylvatica FS Fagus sylvatica FS Fagus sylvatica FS Fagus sylvatica EP Rhodothamnus chamaecistus ES Carex ferruginea AT Carex brachystachys E2b E2b E2 + E2b . E2a . El . E2a + El . El + El + E3 . E2b + E2a + El E2b E2a El El E2 E2a E3a E2b E2a El E3 E2b E2a El E3a E2b E2a El E3b E3a E2b E2a El El El El 1 1 4 2 2 + 1 + 1 2 1 + + 1 + 1 i + + + + + + + 1 1 1 + + + + + 4 4 5 4 1 1 + + + + + i t- + 1 1 Pr. 1 1 11 11 5 2 17 15 15 20 14 13 16 2 14 7 9 7 7 6 7 5 5 4 2 3 8 9 4 4 2 6 10 1 7 4 2 Fr. 5 5 50 50 23 9 77 91 64 59 73 9 64 32 41 36 41 36 32 32 28 32 23 23 18 9 14 36 41 18 18 9 27 45 5 32 18 9 a > « r, o w — tfl ?? CI O o M o z; > r* s H > O M 22 ^ X M a tu < to t-1 O ►o g M z; ^ o •n TI G 5« M < M O M -3 O Z S O >3 /O cl to H S o g t-1 i-1 to ►C ^ Number of relevé (Zaporedna številka popisa) O AF Aremonio-Fagion ^ Cyclamen purpurascens El w Anemone trifolio El Knautia drymeia El O Heüeborus niger El ^ Cardamine enneaphyüos El TA Tilio-Acerion i-j Thalictrum aquilegiifolium El ^ Polystichum aculeatum El O Geranium robertianum El t-1 O Euonymus latifolia E2b 2 Aruncus dioicus El > FS Fagetalia sylvaticae o Mélica nutans El Mercurialis perennis El kj Galium laevigatum El to Daphne mezereum E2a Lonicera alpígena E2a Galeobdolon flavidum El Dryopteris filix-mas El Euphorbia amygdaloides El Mycelis muralis El Salvia glutinosa El Actaea spicata El Brachypodium sylvaticum El Luzula nivea El Paris quadrifolia El Epilobium montan um El Epipactis helleborine El Poa nemoralis El QP Quercetalia pubescenti-petraeae Sorbas aria (Aria edulis) E3a Sorbus aria E2b Sorbas aria E2a Sorbus aria El Primula veris El Hypericum montanum El Convaüaria majalis El QF Querco-Fagetea Hepático nobilis El Carex digit at a El QR Potentilla erecta El Lonicera xylosteum E2a Piatanthera bifolia El AI Dryopteris carthusiana El 9 10 11 12 13 14 15 16 17 18 19 20 21 22 Pr. Fr. + + r 1 + 1 1 1 + 1 + 1 1 1 1 + 1 1 1 . + + + 1 + + 1 + 1 . 1 + 1 1 1 1 19 86 1 1 1 + 13 59 . + 3 14 2 9 1 5 1 1 1 1 1 1 + .... 1 1 . +...........+ + +. + +............. + . . +......... .....+.......... ............r 6 27 4 18 3 14 1 5 1 5 + + + 1 + 2 1 1 1 1 2 1 + 1 + + 1 1 1 + 20 91 1 1 1 2 1 1 1 1 1 + + 1 + 1 + + 1 + 20 91 + 1 + 1 1 + + + + + + 1 1 + + 15 68 + + 1 1 1 + + + + + + + + 13 59 + + 1 + + + + + + 1 + + 12 55 + + 1 + + 1 7 32 + + + + + + 6 27 + + + + 1 5 23 + + + + + 5 23 23 9 9 9 9 5 5 5 36 41 41 9 5 5 5 32 23 9 9 5 5 a > O W t-1 M ?? C o o M o z; > t-1 v: > O M ^ k M a M < M t-1 O T3 S M z; ^ o •n •n O M < M O M ^ > ¡j O z 2 o £ /O a M O •n O g t-1 t-1 M ►C Number of relevé (Zaporedna številka popisa) VP Vaccinio-Piceetea Picea abies E2b Picea abies E2a Vaccinium myrtillus El Maianthemum bifolium El Valeriana tripteris El Lonicera nigra E2a Solidago virgaurea El Vaccinium vitis-idaea El Veronica urticifolia El Abies alba E2b Abies alba E2a Gymnocarpium dryopteris El Oxalis acetosella El Calatnagrostis villosa El Lycopodium annotinum El Gentiana asclepiadea El Saxífraga cuneifolia El Huperzia selago El Rosa pendulino E2a Phegopteris connectilis El Aposeris foetida El Hieracium murorum El Luzula pilosa El Dryopteris expansa El Luzula sylvatica El EP Erico-Pinetea Calatnagrostis varia El O Molinia arundinacea El Rubus saxatilis El Erica carnea El O Peucedanum rablense El O Amelanchier ovalis E2b 2 Amelanchier ovalis E2a > Polygala chamaebuxus El ^ Cotoneaster tomentosus E2a Ci Euphrasia cuspidata El O Asperula aristata El q Rhamnus saxatilis El Q Aquilegia nigricans El P BA Betulo-Alnetea g Sorbus mougeotii E3a Sorbus mougeotii E2b Sorbus austriaca subsp. austriaca E2b 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 + 1 + + + + + r 1 + + + + + + 2 + + + 1 + + + + + 1 + + + 1 + 1 + + + + 5 11 8 6 6 5 4 4 4 3 3 3 3 2 2 2 2 2 1 1 1 1 1 1 1 23 50 36 27 27 23 18 18 18 14 14 14 14 9 9 9 9 9 5 5 5 5 5 5 5 1 1 1 1 2 3 1 1 1 2 1 1 + 1 2 1 2 1 2 + 1 22 100 1 + + 1 + + + + 3 3 + + 3 + 1 1 17 77 + + 1 + + 1 1 1 1 + 1 + + 1 + + 16 73 2 . + 2 2 3 2 3 1 2 + + 3 12 55 1 + 1 1 . + + r 1 + 9 41 + + + 3 14 r + 1 + + + 6 27 + + 1 + + 5 23 + + 1 + 4 18 18 9 5 5 5 27 5 a > ^ O w t-1 w ?? cl O o w o z; > t-1 v: > O M § k M a M < M t-1 O T3 S M z; ^ o •n •n O M < M O M ^ O z 2 o /O a tn o •n O g t-1 t-1 M ►C to o •n O t-; > w O t-1 o 2 o > tfl ^ o tfl O t-1 o 2 o > Number of relevé (Zaporedna številka popisa) Mu A Mulgedio-Aconitetea Aconitum degenii subsp. paniculatum Chaerophyllum hirsutum Athyrium filix-femina Polygonatum verticillatum Agropyron caninum Veratrum album Viola biflora Senecio ovatus Phyteuma ovatum Crepis paludosa SSC Sambuco-Salicion capreae, Rhamno-Prunetea Sorbus aucuparia Sorbus aucuparia Sorbus aucuparia Sorbus aucuparia RP Juniperus communis RP Juniperus communis Betula péndula FB El El El El El El El El El El E3a E2b E2a El E2b E2a E3a Epilobietea angustifolii Fragaria vesca El 1 Rubus idaeus E2a Trifolio-Geranietea Vincetoxicum hirundinaria El + + + Viola hirta El + Anthericum ramosum El + Clinopodium vulgare El Hypericum perforatum El Laserpitium siler El + Polygonatum odoratum El Laserpitium latifolium El Valeriana wallrothii (V. collina) El Festuco-Brometea Buphthalmum salicifolium El 1 Stachys recta agg, El 1 + Galium lucidum El 1 Cirsium erisithales El + 1 Brachypodium rupestre El Carex humilis El Prunella grandiflora El + + Pimpinella saxífraga El + Carlina acaulis El + Euphorbia cyparissias El + Linum catharticum El + + Gentianella ciliata El + 9 10 11 12 13 14 15 16 17 18 19 20 21 22 Pr. Fr. + + + 1 + + + + + + + 1 1 1 + + + 1 . + + + + + . H • 1 . + 1 + + + + + + 1 1 4 4 4 3 2 2 2 1 1 1 6 5 4 8 2 2 1 11 2 11 7 3 2 2 1 1 1 1 17 7 7 5 4 4 4 3 2 2 2 1 18 18 18 14 9 9 9 5 5 5 27 23 18 36 9 9 5 50 9 50 32 14 9 9 5 5 5 5 77 32 32 23 18 18 18 14 9 9 9 5 a > ^ O w t-1 w ?? cl o o w o z; > t-1 v: > O M ^ k M a M < M t-1 O TJ S M z; ^ o •n •n O M < M O M ^ > ¡j O z 2 o £ /O a M O •n g t-1 t-1 M ►C ^ Number of relevé (Zaporedna številka popisa) Arabis hirsuta El Hippocrepis comosa El Teucrium montanum El Gymnadenia conopsea El Briza media El MA Molinio-Arrhenatheretea Galium album El PaT Ranunculus nemorosus agg. El Achillea millefolium El Lathyrus pratensis El Trifolium pratense El + CD Caricetalia davallianae Parnassia palustris El Tofieldia calyculata El ES Elyno-Seslerietea Betónica alopecuros El Laserpitium peucedanoides El + Sesleria caerulea El Scabiosa lucida subsp. stricta El + Carduus crassifolius El 1 Thymus praecox subsp. polytrichus El Linum julicum El Festuca calva El Rhinanthus glacialis El Lotus corniculatus s. lat. (L. alpinus ?) El + Carex mucronata El Helianthemum nummularium subsp. grandiflorum El Phyteuma orbiculare El + O Aster bellidiastrum El " Thesium alpinum El + > r M Dryas octopetala El + O Achillea clavenae El + t-1 O Acinos alpinus El + 2 Alchemilla alpígena El + > Carex sempervirens El ® Euphrasia picta El O ¡uncus monanthos El tu O Astrantia bavarica El O Arabis vochinensis El 2 Globularia nudicaulis El P Selaginella selaginoides El g TR Thlaspietea rotundifolii ^ Astrantia carniolica El + t-o Valeriana montana El Adenostyles glabra El + to 2 3 4 5 + + . . . 10 11 12 13 14 15 16 17 18 19 20 21 22 Pr. + + + + + 1 + + 2 1 1 1 2 2 + + 1 1 1 + + 2 + 1 + + 1 1 + + 1 1 1 + + + + + + + + + + + + + + + + 1 1 1 + 1 1 + Fr. 5 5 5 5 5 5 23 4 18 2 9 2 9 1 5 2 1 21 15 13 10 9 5 95 68 59 45 36 36 27 23 18 14 14 14 14 9 5 5 5 5 5 5 5 5 5 5 5 5 + 11 50 + 7 32 6 27 a > ^ O w t-1 w ?? cl O o w o z; > t-1 v: > O M z; ^ X M a M < M t-1 O TJ S M z; ^ o •n •n O M < M O M ^ O z z o /O a M O •n t O g t-1 t-1 M ►C to to ^ Number of relevé (Zaporedna številka popisa) Achnatherum calamagrostis El > Biscutella laevigata El S Campanula cespitosa El i-1 Festuca nitida El Q Aquilegia einseleana El O Gypsophila repens El ^ Hieracium bifidum El ^ Rumex scutatus El tu Trisetum argenteum El Scrophularia juratensis El q Asplenium fissum El O Arabis alpina El ^ Hieracium porrifolium El S Silene vulgaris subsp. glareosa El Viola pyrenaica El o Athamanta cretensis El Petasites paradoxus El Hieracium piloselloides El Leontodon hyoseroides El Heliosperma alpestre El Dianthus sternbergii El Cerastium subtriflorum El Cystopteris montana El Soldanella alpina El AT Asplenietea trichomanis Asplenium viride El Paederota lutea El Asplenium trichomanes El Kernera saxatilis El Moehringia muscosa El Asplenium ruta-muraria El Cystopteris regia El Cystopteris fragilis El ML Mosses and lichens (Mahovi in lišaji) Ctenidium molluscum EO Tortella tortuosa EO Hylocomium splendens EO Rhytidiadelphus triquetrus EO Fissidens dubius EO Dicranum scoparium EO Peltigera canina EO Homalothecium lutescens EO Conocephalum conicum EO Schistidium apocarpum EO Mnium thomsonii EO 4 5 6 7 8 1 + + . . + . + . . . . + . . . . + . . + . + . . . + 9 10 11 12 13 14 15 16 17 18 19 20 21 22 Pr. ..........+ + +.......+ . ...........+ + 1 . 1 + + + + 1 1 1 1 1 1 Pr. Fr. 5 23 5 23 5 23 5 23 4 18 4 18 4 18 4 18 4 18 3 14 3 14 2 9 2 9 2 9 2 9 1 5 1 5 1 5 1 5 1 5 1 5 1 5 1 5 1 5 + 7 32 5 23 4 18 2 9 2 9 2 9 2 9 2 9 1 14 64 12 54,5 1 11 50 + 11 50 1 5 23 3 14 3 13,6 2 9 2 9 + 2 9 1 5 a > ^ O w t-1 w ?? cl o o w o z; > t-1 v: > O M § X M a M < M t-1 O TJ S M z; ^ o •n •n O M < M O M ^ > ¡j O z § o £ /O a tn o •n t O g t-1 t-1 M ►C TI O £ > ro O r* O 2 o > ta H O m C r- O O n > Number of releve (Zaporedna številka popisa) Plagiochila porelloides Plagiomnium undulatum Mnium sp. Polytrichum formosum Neckera crispa 10 11 12 13 14 15 16 17 18 19 20 21 22 Pr. E0 E0 E0 E0 E0 Fr. 5 5 5 5 5 Legend - Legenda Pr. Presence (number of releves in which the species is presented) - število popisov, v katerih se pojavlja vrsta Fr. Frequency in % - frekvenca v % Gr Gravel - prod Sc Scree - grušč Mo Moraine (Til) - morena (til) Ta Talus - vršaj Rs Rockslide - podorno skalovje D Dolomite - dolomit Co Colluvial soil - koluvialna tla Li Lithosols - kamnišče Re Rendzina - rendzina QR Quercetalia roboris Al Alnion incanae PaT Poo alpinae-Trisetetalia to I. DAKSKOBLER: SUCCESSIONAL STAGES IN THE DEVELOPMENT OF FOREST VEGETATION IN CIRQUES OF TWO VALLEYS Table 2: Succession of vegetation on fluvial-glacial gravel sediments of the Kotarica in Klin (the Kot Valley), the Julian Alps Preglednica 2: Sukcesijski razvoj rastja na rečno-ledeniških prodnatih nanosih Kotarice v Klinu v dolini Kot, Julijske Alpe Number of relevé (Zaporedna številka popisa) 1 2 3 4 5 6 Database number of relevé (Delovna številka popisa) 273197 273196 273194 273191 273190 273192 Elevation in m (Nadmorska višina v m) 990 990 990 980 990 980 Aspect (Lega) NE NE NE NE NE NE Slope in degrees (Nagib v stopinjah) 1 1 1 1 1 1 Parent material (Matična podlaga) Gr Gr Gr Gr Gr Gr Soil (Tla) Li Li Re Re Re Re Stoniness in % (Kamnitost v %) 60 20 5 5 0 0 Cover in % (Zastiranje v %): Upper tree layer (Zgornja drevesna plast) E3b 60 60 60 Lower tree layer (Spodnja drevesna plast) E3a 5 10 20 10 Shrub layer (Grmovna plast) E2 20+60 70+20 80 30+30 20+20 40+60 Herb layer (Zeliščna plast) El 30 70 90 90 95 90 Moss layer (Mahovna plast) E0 20 5 10 5 5 Maximum diameter of trees cm 20 35 35 35 (Največji prsni premer dreves) Maximum height of trees (Največja drevesna višina) m 15 22 20 19 Number of species (Število vrst) 50 73 69 54 60 47 Relevé area (Velikost popisne ploskve) m2 100 200 200 400 400 400 Date of taking relevé (Datum popisa) 9/19/2018 9/19/2018 9/19/2018 9/19/2018 9/19/2018 9/19/2018 Locality (Nahajališče) Kot-Klin Kot-Klin Kot-Klin Kot-Klin Kot-Klin Kot-Klin Quadrant (Kvadrant) 9549/3 9549/3 9549/3 9549/3 9549/3 9549/3 Coordinate GK Y (D-48) m 415054 415051 415090 415145 415070 415105 Coordinate GK X (D-48) m 5140955 5140996 5141082 5141160 5141009 5141081 Diagnostic species of the associations (Diagnostične vrste asociacij) Pr. Fr. SP Salix eleagnos E3a 1 + 2 33 SP Salix eleagnos E2b 1 + 2 33 SP Salix eleagnos E2a 4 + 2 33 SP Salix eleagnos El 2 1 17 SP Salix purpurea E2b + 1 17 TR Petasites paradoxus El 3 + + 1 + + 6 100 EP Pinus mugo E2b + 4 4 1 1 5 83 EP Pinus mugo E2a + 1 1 + + 5 83 EP Pinus mugo El + + 2 33 EP Amelanchier ovalis E2b + 1 17 EP Amelanchier ovalis E2a + r r 3 50 TR Hieracium porrifolium El 1 1 + 3 50 FS Fagus sylvatica E3b 3 3 4 3 50 FS Fagus sylvatica E3a 1 1 2 33 FS Fagus sylvatica E2b + + 1 1 4 67 FS Fagus sylvatica E2a + + 1 1 4 67 FS Fagus sylvatica El + + 1 + + 5 83 FS Laburnum alpinum E3a + + 1 r 4 67 FS Laburnum alpinum E2b 1 + + 1 + 5 83 FS Laburnum alpinum E2a + + 1 + 4 67 EP Rhododendron hirsutum E2a + 1 + 3 2 1 6 100 VP Clematis alpina El 1 1 1 3 50 FB Brachypodium rupestre El 1 1 + 3 50 VP Homogyne sylvestris El + + 2 33 EP Erico-Pinetea Calamagrostis varia El 1 1 + 2 3 3 6 100 Erica carnea El + 4 4 4 4 4 6 100 Molinia arundinacea El 1 2 4 + 1 1 6 100 Carex alba El + + 1 2 1 5 83 Euphrasia cuspidata El + 1 1 3 50 Rubus saxatilis El 1 1 1 3 50 Allium ericetorum El + + 2 33 Aquilegia nigricans El + + 2 33 124 FOLIA BIOLOGICA ET GEOLOGICA 60/1 - 2019 I. DAKSKOBLER: SUCCESSIONAL STAGES IN THE DEVELOPMENT OF FOREST VEGETATION IN CIRQUES OF TWO VALLEYS TA FS QP QF AI VP Number of relevé (Zaporedna številka popisa) 1 2 3 4 5 6 Pr. Fr. Asperula aristata E1 + + 2 33 Carex ornithopoda E1 1 + 2 33 Polygala chamaebuxus E1 1 1 2 33 Rhodothamnus chamaecistus E1 + 1 2 33 Crepis slovenica E1 + 1 17 Gymnadenia odoratissima E1 r 1 17 Cotoneaster tomentosus E2a r 1 17 Aremonio-Fagion Helleborus niger E1 1 1 2 1 1 5 83 Cyclamen purpurascens E1 + + + 1 1 5 83 Anemone trifolia E1 + + + + 1 5 83 Rhamnus fallax E2a + 1 17 Rhamnus fallax E2b r 1 17 Tilio-Acerion Acer pseudoplatanus E3b 1 1 + 3 50 Acer pseudoplatanus E3a + 1 + 3 50 Acer pseudoplatanus E2b + 1 17 Acer pseudoplatanus E2a + 1 17 Acer pseudoplatanus E1 + + + + 1 1 6 100 Fagetalia sylvaticae Melica nutans E1 + 1 1 + 4 67 Daphne mezereum E2a 1 1 1 3 50 Lonicera alpigena E2a 2 1 + 3 50 Galium laevigatum E1 + + + 3 50 Euphorbia amygdaloides E1 + + 2 33 Cypripedium calceolus E1 + 2 33 Mercurialis perennis E1 + + 2 33 Prenanthes purpurea E1 + + 2 33 Galeobdolon flavidum E1 + 1 17 Dryopteris filix-mas E1 + 1 17 Quercetalia pubescenti-petraeae Sorbus aria (Aria edulis) E3a + 1 17 Sorbus aria (Aria edulis) E2b 1 1 2 33 Sorbus aria (Aria edulis) E2a + + + 1 1 + 6 100 Sorbus aria (Aria edulis) E1 + + 2 33 Querco-Fagetea Potentilla erecta E1 1 1 + 1 4 67 Hepatica nobilis E1 + + 1 3 50 Viola riviniana E1 + r 2 33 Frangula alnus E2a + 1 17 Vaccinio-Piceetea Picea abies E3b + 1 1 3 50 Picea abies E3a + + r + 4 67 Picea abies E2b 1 + + 1 + 5 83 Picea abies E2a + + + + 4 67 Picea abies E1 + 1 17 Pyrola rotundifolia E1 + + 2 1 1 5 83 Larix decidua E3b r + + 3 50 Larix decidua E3a 1 + 2 33 Larix decidua E2b 1 + + + 4 67 Larix decidua E2a + + + + 4 67 Larix decidua E1 + + 2 33 Vaccinium myrtillus E1 + 3 3 3 4 67 Vaccinium vitis-idaea E1 + 2 2 2 4 67 Lonicera nigra E2a 1 1 + 3 50 Orthilia secunda E1 1 + + 3 50 Lycopodium annotinum E1 + + + 3 50 Gentiana asclepiadea E1 + + 2 33 Veronica urticifolia E1 + 1 17 Hieracium murorum E1 + 1 17 Lonicera caerulea E2a r 1 17 Maianthemum bifolium E1 + 1 17 125 FOLIA BIOLOGICA ET GEOLOGICA 60/1 - 2019 I. DAKSKOBLER: SUCCESSIONAL STAGES IN THE DEVELOPMENT OF FOREST VEGETATION IN CIRQUES OF TWO VALLEYS Number of relevé (Zaporedna številka popisa) l 2 3 4 Abies alba E2a Aposeris foetida El Solidago virgaurea El Luzula pilosa El BA Betulo-Alnetea Salix appendiculata E2b t t Salix appendiculata E2a 2 t Salix appendiculata El l Juniperus sibirica E2a Sorbus chamaemespilus E2a t Salix waldsteiniana E2a t Salix glabra E2a t MuA Mulgedio-Aconitetea Polygonatum verticillatum El Athyrium filix-femina El EA Epilobietea angustifolii Fragaria vesca El t SSC Sambuci-Salicion capreae, Rhamno-Prunetea Sorbus aucuparia E3a t t Sorbus aucuparia E2b t Sorbus aucuparia El t t RP Berberis vulgaris E2a FB Festuco-Brometea Buphthalmum salicifolium El t t l t Galium lucidum El t t t Carlina acaulis El t t Cirsium erisithales El t t Hippocrepis comosa El l t Prunella grandiflora El t l Gentianella ciliata El l Festuca rupicola El t Coronilla vaginalis El t Stachys recta s. lat. El t Gymnadenia conopsea El t TG Viola hirta El t PaT Poo alpinae-Trisetetalia, Molinio-Arrhenatheretea MA Angelica sylvestris El t r Campanula scheuchzeri El t Ranunculus nemorosus El r ES Elyno-Seslerietea Laserpitium peucedanoides El t l t Dryas octopetala El l 3 2 t Lotus corniculatus s. lat. (L. alpinus) El t l t Betonica alopecuros El t l Aster bellidiastrum El t l 2 Carex mucronata El t l t Globularia cordifolia El t l t Carex firma El t t l Senecio abrotanifolius El t t t Scabiosa lucida subsp. stricta El t l Carduus crassifolius El t t Sesleria caerulea El t t NS Antennaria dioica El t t CD Tofieldia calyculata El t t Achillea clavenae El t Phyteuma orbiculare El t Globularia nudicaulis El t Thymus praecox subsp. polytrichus El t Arctostaphylos alpinus El t Gentiana clusii El t Rhinanthus glacialis El t Thesium alpinum El t Pr. Fr. l ly l ly l ly l ly 2 33 3 so l ly 2 33 3 so l ly l ly l ly l ly 3 so 3 so l ly 4 6y l ly s 83 4 6y 3 so 2 33 2 33 2 33 2 33 l ly l ly l ly l ly l ly 2 33 l ly l ly s 83 4 6y 4 6y 4 6y 3 so 3 so 3 so 3 so 3 so 3 so 2 33 2 33 2 33 2 33 l ly l ly l ly l ly l ly l ly l ly l ly s 6 t t t t t t t t t t t 26 FOLIA BIOLOGICA ET GEOLOGICA 60/1 - 2019 I. DAKSKOBLER: SUCCESSIONAL STAGES IN THE DEVELOPMENT OF FOREST VEGETATION IN CIRQUES OF TWO VALLEYS Number of relevé (Zaporedna številka popisa) 1 2 S 4 s 6 Pr. Fr. Ranunculus carinthiacus E1 + 1 17 TR Thlaspietea rotundifolii Campanula cespitosa E1 1 1 + + 4 67 Astrantia carniolica E1 + + + + 4 67 Heliosperma alpestre E1 2 + + S so Dianthus sternbergii E1 + + + S so Hieracium bifidum E1 + + + S so Adenostyles glabra E1 + + + S so Gymnocarpium robertianum E1 1 + S so Achnatherum calamagrostis E1 + + 2 SS Biscutella laevigata E1 + + 2 SS Hieracium piloselloides E1 + + 2 SS Rumex scutatus E1 + + 2 SS Trisetum argenteum E1 + + 2 SS Valeriana montana E1 + 2 SS Silene vulgaris subsp. glareosa E1 1 1 17 Minuartia austriaca E1 + 1 17 Papaver alpinum subsp. ernesti-mayeri E1 + 1 17 Scrophularia juratensis E1 + 1 17 Linaria alpina E1 + 1 17 AT Asplenietea trichomanis Potentilla clusiana E1 + 1 17 Campanula thyrsoides E1 + 1 17 ML Mosses and lichens (Mahovi in lišaji) Tortella tortuosa Eo 2 + 1 + 4 67 Hylocomium splendens Eo + 1 + S so Rhytidiadelphus triquetrus Eo + 1 2 SS Dicranum scoparium Eo + + 2 SS Cladonia rangiferina Eo + 1 17 Legend - Legenda Pr. Presence (number of relevés in which the species is presented) - število popisov, v katerih se pojavlja vrsta Fr. Frequency in % - frekvenca v % Gr Gravel - prod Li Lithosols - kamnišče Re Rendzina - rendzina QR Quercetalia roboris AI Alnion incanae TG Trifolio-Geranietea NS Nardetea strictae CD Caricetalia davallianae 127 FOLIA BIOLOGICA ET GEOLOGICA 60/1 - 2019